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1 Introduction: Botany and Importance

S.K. Mukherjee1 and R.E. Litz2


1
Calcutta University, Calcutta, India
2
University of Florida, Florida, USA

1.1 Introduction 1
1.2 Description of Mango 2
The tree 2
Flowers 2
The fruit 3
The seeds and polyembryony 4
1.3 History of Cultivation 5
Origin of Mangifera indica 5
Domestication of mango 9
Distribution 10
1.4 Germplasm Conservation 11
Genetic erosion 11
Collection and documentation of Mangifera germplasm 12
Relevance of germplasm resources to mango improvement 12
1.5 Importance of Mango 12
Cultivars 12
1.6 Production and Uses 14

1.1 Introduction
Mango has become a major fruit crop of the tropics and subtropics, particu-larly in
Asia, where the mango has always been the most important fruit crop and where it
has been considered the ‘king of fruits’ (Purseglove, 1972). A generation ago, the
Green Revolution culminated, creating surpluses of sta-ple and horticultural crops
in many developing countries. The Green Revo-lution was the result of nearly a
century of effort of applying Mendelian genetics to crop improvement (i.e.
conventional breeding) together with the optimization of agronomic and
horticultural practices and the successful management of insect pests and diseases.
However, improvement of tree
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 1
2 S.K. Mukherjee and R.E. Litz

crops has lagged far behind field crops for several reasons: their heterogene-ity,
polyploidy, lengthy juvenile period, time required for evaluation of trees in the
field, and the relatively high cost of maintaining tree plantings. For the most part,
fruit cultivars continue to be ancient selections, many of which have serious
problems, including alternate bearing, lack of disease resistance, low yields, etc.
The rapid growth of mango production in recent years has been due to its
expansion into new growing regions of the New World, China and parts of Africa;
the planting of regular bearing selections; and the adop-tion of modern field
practices, which include irrigation management, control of flowering, etc.
Agricultural practices are currently undergoing another revolution, as integrated
pest and disease management replaces the earlier reliance on agrichemicals, and
emerging fields within biotechnology begin to impact cultivar development.

1.2 Description of Mango

The tree

The mango tree is believed to have evolved as a canopy layer or emergent species
of the tropical rainforest of South and South-east Asia (Kaur et al., 1980; Bompard,
Chapter 2, this volume). Mature trees can attain a height of 40 m or more, and can
survive for several hundred years. Mango trees that have been domesticated by
selection from openly pollinated seedling popu-lations show variation in tree
architecture (i.e. shape and size). The tree is an arborescent evergreen. Leaves are
simple and alternate, with petioles that range in length from 1 to 12.5 cm. Leaf
morphology is highly variable, de-pending on the cultivar: leaves can be
lanceolate, oblong, ovate and interme-diate types involving these forms. Leaf
length ranges from 12 to 38 cm and width can be between 2–13 cm. Young leaves
are copper-coloured, changing gradually to light and then dark green with age. The
leaves are spirally arranged in whorls and are produced in flushes. The canopy is
normally oval, elongated or dome shaped. The juvenile period of seedling trees can
range from 3 to 7 years. The root system consists of a long, vigorous taproot and
abundant surface feeder roots.

Flowers

Mango flowers are borne on terminal pyramidal panicles, and are glabrous or
pubescent; the inflorescence is rigid and erect, up to 30 cm long, and is widely
branched, usually tertiary, although the final branch is always cymose. The
inflorescence is usually densely flowered with hundreds of small flow-ers, which
are 5–10 mm in diameter. The flowers are either monoecious or polygamous, and
both monoecious and polygamous flowers are borne within a single inflorescence
(Plate 1). The pistil aborts in male flowers. The ratio of monoecious to polygamous
flowers is strongly influenced by
Introduction: Botany and Importance 3

environmental and cultural factors. The flowers have four or five sepals and petals
that are ovate to ovoid to lanceolate and also thinly pubescent. The floral disc also
is four- or five-lobed, fleshy and large and located above the base of the petals.
There are five large, fleshy stamens, only one or two of them being fertile; the
remaining stamens are sterile staminodes that are sur-mounted by a small gland. In
addition, two or three smaller filaments arise from the lobes of the nectaries. The
stamens are central. The ovule is anatro-pous and pendulous. It is believed that the
flowers are cross-pollinated by flies (see Davenport, Chapter 5, this volume).

Mukherjee (1951a, 1953) investigated the pollen morphology of mango and 12


other Mangifera species. Their pollen grains were tricolpate of almost the same
size. Mondal et al. (1982, cited in Kostermans and Bom-pard, 1993) attempted to
correlate pollen morphology with taxonomic relationships of 17 Mangifera species
based upon different characteristics of the exine and sporoderm. They
demonstrated that all of the species of section II (subgenus Limus) possess coarse
exine; whereas there was no clear correlation with pollen type in species within
section I (subgenus Mangifera).

The fruit

Description
The mango fruit is a large, fleshy drupe, containing an edible mesocarp of varying
thickness. The mesocarp is resinous and highly variable with respect to shape, size,
colour, presence of fibre and flavour. The flavour ranges from turpentine to sweet.
The exocarp is thick and glandular. There is a character-istic beak that develops
laterally on the proximal end of the fruit. A sinus is always present above the beak.
Fruit shape varies, including elongate, oblong and ovate or intermediate forms
involving two of these shapes. Fruit length can range from 2.5 to > 30 cm,
depending on the cultivar. The endo-carp is woody, thick and fibrous; the fibres in
the mesocarp arise from the endocarp.

The mango fruit is climacteric (see Brecht and Yahia, Chapter 14, this
volume), and increased ethylene production occurs during ripening. Chloro-phyll,
carotenes, anthocyanins and xanthophylls are all present in the fruit. The skin is
generally a mixture of green, red and yellow pigments, although fruit colour at
maturity is genotype dependent. During ripening the chloro-plasts in the peel
become chromoplasts, which contain yellow and red pig-ments (Krishnamurthy
and Subramanyam, 1970; Akamine and Goo, 1973; Salunkhe and Desai, 1984;
Mitra and Baldwin, 1997). Peel colour obviously is cultivar dependent (see Knight
et al., Chapter 3, this volume). Fruit of ‘Bom-bay Green’ is green; ‘Carabao’,
‘Manila’, ‘Mulgoa’ and ‘Arumanis’ are greenish-yellow; ‘Dashehari’ and
‘Alphonso’ are yellow; and ‘Haden’, ‘Keitt’ and ‘Tommy Atkins’ have a red blush.
The red blush is due to the presence of anthocyanins (Lizada, 1991). The pulp
carotenoids in ripe fruit also vary with respect to cultivar (Mitra and Baldwin,
1997).
4 S.K. Mukherjee and R.E. Litz

Flavour
Flavour of the mango mesocarp is a function of carbohydrates, organic acids,
lactones, monoterpene hydrocarbons and fatty acids (Mitra and Baldwin, 1997).
During fruit maturation, starch that accumulates in the chloroplasts is hydrolysed to
sucrose, glucose and fructose (Medlicott et al., 1986; Selvaraj et al., 1989; S.
Kumar et al., 1994); sucrose is present in slightly higher concen-trations than either
fructose or glucose. Organic acid content decreases dur-ing ripening
(Krishnamurthy and Subramanyam, 1970). The dominant organic acid is citric
acid, but glycolic acid, malic acid, tartaric acid and oxalic acids are also present
(Sarker and Muhsi, 1981; Medlicott and Thompson, 1985). The peach-like flavour
of mangoes is attributed to the presence of lac-tones (Lakshminarayana, 1980;
Wilson et al., 1990).

Nutrition
Mango fruit contain amino acids, carbohydrates, fatty acids, minerals, organic
acids, proteins and vitamins. During the ripening process, the fruit are ini-tially
acidic, astringent and rich in ascorbic acid (vitamin C). Ripe mangoes contain
moderate levels of vitamin C, but are fairly rich in provitamin A and vitamins B 1
and B2. Perry and Zilva (1932) determined the vitamin A, C and D content of the
fruit of three Indian mango cultivars, and found that the pulp of mangoes is a
concentrated source of vitamin C. The pulp of mango fruit contains as much
vitamin A as butter, although vitamin D is not present in a significant quantity.
Fruit acidity is primarily due to the presence of malic and citric acids. In addition,
oxalic, malonic, succinic, pyruvic, adipic, galac-turonic, glucuronic, tartaric,
glycolic and mucic acids are also present (Jain et al., 1959; Fang, 1965). Acidity is
cultivar related; for example, immature Florida cultivars have low acidity (0.5–
1.0%) in comparison with ‘Alphonso’ (3%). During ripening, acidity decreases to
0.1–0.2%. Following fruit set, starch accumulates in the mesocarp. Free sugars,
including glucose, fructose and sucrose, generally increase during ripening;
however, the sucrose content increases three- to fourfold due to the hydrolysis of
starch. Sucrose is the principal sugar of ripe mangoes. The sucrose content of ripe
fruit of three Indian cultivars, ‘Alphonso’, ‘Pairie’ and ‘Totapuri’, ranges from 11
to 20% representing 15 to 20% of the total soluble solids (Popenoe, 1932).

The seeds and polyembryony

Mango seeds are solitary, large and flat, ovoid oblong and surrounded by the
fibrous endocarp at maturity. The testa and tegumen are thin and papery. Embryos
are dicotyledonous. Seeds of monoembryonic mango types contain a single zygotic
embryo, whose cotyledons can be unequal in size or lobed in shape. The seeds of
polyembryonic mango types contain one or more embryos (Plate 2); usually one
embryo is zygotic, whereas the remaining embryos are derived directly from the
nucellus, a maternal tissue. Nucellar embryos apparently lack a suspensor.
Polyembryony has also been reported in Mangifera casturi, M. laurina and M.
odorata (Bompard, 1993). Certain
Introduction: Botany and Importance 5

polyembryonic cultivars reportedly can produce seeds with adventitious nucellar


embryos only, for example ‘Strawberry’ (Juliano, 1934), ‘Carabao’ and ‘Pico’
(Juliano and Cuevas, 1932) and ‘Olour’ and ‘Cambodiana’ (Maheshwari et al.,
1955). Early studies suggested that polyembryony appeared to be a polygenic trait
(Juliano, 1934; Sturrock, 1968), segregating as a recessive character in the progeny
of controlled crosses. Recent studies, however, have demonstrated that the
polyembryony trait is inherited as a dominant character (Aron et al., 1998). Several
studies have shown that nucel-lar seedlings can be distinguished from the single
zygotic seedling of poly-embryonic seeds by isozymes (Schnell and Knight, 1992;
Degani et al., 1993) and DNA markers, for example single sequence repeats
(SSRs) (Eiadthong et al., 1999a), amplified fragment length polymorphisms
(AFLPs) (Kashkush et al., 2001) and inter-simple-sequence-repeats (ISSRs)
(Gonzalez et al., 2002). Mango seeds are considered to be recalcitrant, and cannot
survive for more than a few days or weeks at ambient temperatures (Parisot, 1988).
This important characteristic of mango seeds would have inhibited the long
distance dis-persal of mango by seed until recent times.

1.3 History of Cultivation


Origin of Mangifera indica

The largest number of Mangifera species occurs in the Malay Peninsula, the
Indonesian archipelago, Thailand, Indochina and the Philippines (Mukher-jee,
1985; Bompard, 1989; see Bompard, Chapter 2, this volume). The most recent
classification of Mangifera species was based upon floral morphology (Kostermans
and Bompard, 1993) and included 69 species, most of which are included in two
subgenera Mangifera and Limus with another 11 species occupying an uncertain
position (Table 1.1). Eiadthong et al. (1999b) described the phylogenetic
relationships among Mangifera species using genomic restriction fragment length
polymorphisms (RFLPs) and amplification of chloroplast DNA (cpDNA), and
suggested that the Mangifera species should be classified using molecular data. In
the next few years, it is likely that molecular biology will have a major impact on
phylogenetic studies involving mango and its relatives.

Mangifera species with a single fertile stamen are distributed in north-eastern


India, Myanmar, Thailand and the Malay Peninsula. Many of the mango relatives
have small fruits with thin, acidic flesh, large seeds, abun-dant fibre and astringent
resinous substances that are localized near the skin. In addition to M. indica, edible
fruit is produced by at least 26 other species in the genus, primarily species found
in South-east Asia (Gruezo, 1992). Mangifera caesia, known as ‘binjai’ or
‘kemang’ in South-east Asia, is culti-vated in Java, where it bears fruit in the
mango off-season (Bompard, 1992a). Mangifera foetida is less commonly
cultivated due to its highly astringent fruit; however, the fruit is widely used for
pickling and as a substitute for tamarind (Bompard, 1992b). Mangifera kemang and
M. altissima are consumed
6
Table 1.1. Classification of Mangifera species according to Kostermans and Bompard (1993).

Genus Subgenus Section Species

Mangifera Mangifera Marchandora Pierre M. gebede Miq


Euantherae Pierre M. caloneura Kurz
M. cochinchinensis Engler
M. pentandra Hooker f.
Rawa Kosterm. M. andamanica King M. minutifolia Evard.

S.K. Mukherjee and R.E. Litz


M. gracilepes M. nicobarica Kosterm.
M. griffithii Hooker f. M. paludosa Kosterm.
M. merrillii Mukherji M. parvifolia Boerl. & Koorders
M. microphylla Griff. ex Hooker f.
Mangifera Ding Hou M. altissima Blanco. M. mucronulata Bl.
M. applanata Kosterm. M. oblongifolia Hooker f.
M. austro-indica Kosterm. M. orophila Kosterm.
M. austro-yunnanensis Hu M. pedicellata Kosterm.
M. casturi Kosterm. M. pseudo-indica Kosterm.
M. collina Kosterm. M. quadrifida Jack
M. dewildei Kosterm. M. rigida Bl.
M. dongnaiensis Pierre M. rubropetala Kosterm.
M. flava Evard. M. rufocostata Kosterm.
M. indica L. M. similis Bl.
M. lalijiwa Kosterm. M. sulauesiana Kosterm.
M. laurina Bl. M. sumbawaensis Kosterm.
M. linearifolia (Mukherji) Kosterm. M. sylvatica Roxb.
M. longipetiolata King M. swintonioides Kosterm.
M. magnifica Kochummen M. timorensis Bl.
M. minor Bl. M. torquenda Kosterm.
M. monandra Merr. M. zeylanica (Bl.) Hooker f.
Limus (Marchand) M. blommesteinii Kosterm. M. leschenaultii Marchand
Kosterm. M. caesia Jack M. macrocarpa Bl.
M. decandra Ding Hou M. odorata Griff.
M. foetida Lour. M. pajang Kosterm.
M. kemanga Bl. M. superba Hooker f.
M. lagenifera Griff.
Species of uncertain M. acutigemma Kosterm. M. persiciformis Wu & Ming
position M. bompardii Kosterm. M. subsessifolia Kosterm.
M. bullata Kosterm. M. taipa Buch.-Hamilton
M. campospermoides M. transversalis Kosterm.

Introduction: Botany and Importance


Kosterm.
M. hiemalis Liang Jian Ying M. utana Utana
M. maingayii Hooker f.

7
8 S.K. Mukherjee and R.E. Litz

as fresh fruit or used green as a salad (Angeles, 1992; Bompard, 1992a). Mangifera
pajang has the largest fruit in the genus, and is an attractive fruit. Mangifera
odorata is grown in the Philippines and Indonesia, and has occa-sionally been used
as a rootstock for mango (Ochse, 1931; Bompard, 1992c). Mangifera odorata is
widely grown in the humid lowlands of South-east Asia in areas that are unsuitable
for mango as a mango substitute. Mangifera lau-rina and M. pentandra are
appreciated as salad ingredients (Bompard, 1992d). In addition, M. griffithii, M.
minor, M. monandra, M. quadrifida and M. similis have palatable fruit that are
considered to have great potential (Gruezo, 1992). All mango cultivars belong to
the species M. indica.
According to De Candolle (1884), ‘It is impossible to doubt that it (the mango)
is a native of south Asia or of the Malay Archipelago, when we see the multitude
of varieties cultivated in those countries, the number of ancient names, in particular
a Sanskrit name, its abundance in the gardens of Bengal, of Deccan peninsula, and
of Ceylon even in Rheede’s time (i.e. 1683).’ Although the centre of origin and
diversity of the genus Mangifera is now firmly established as being in South-east
Asia, the origin of M. indica has been a matter of speculation for many years. The
fossil record provides few clues, as only a single fossil bearing the imprint of a leaf
of M. pentandra has ever been found (Seward, 1912). Mangifera indica is believed
to have first appeared during the Quatenary period (Mukherjee, 1951b). Blume
(1850) considered that mango might have originated from several related species,
primarily located in the Malay archipelago.

On the basis of ancient accounts of travellers and the written historical record,
it was believed for many years that mango must have originated in India and spread
outwards from there to South-east Asia and thence to the New World and Africa.
Because north-eastern India is at the northernmost edge of the distribution of the
Mangifera species, Hooker (1876) suggested that mango might have been
naturalized in India. The historical record pro-vides a sometimes conflicting
account of the distribution of mango. Miquel (1859) did not record it as being wild
in the Indonesian archipelago. Accord-ing to Rumphius (1741), the mango was
introduced into certain islands of the Indonesian archipelago within recent times;
however, the mango was in cul-tivation in Java at least as early as AD 900–1100,
when the temple at Borobo-dur was built and faced with carvings of the Buddha in
contemplation under a mango tree (Plate 3). Based upon taxonomic and recent
molecular evidence, it is now apparent that the mango probably evolved within a
large area including north-western Myanmar, Bangladesh and north-eastern India
(see Bompard, Chapter 2, this volume).

Polyembryonic and monoembryonic M. indica


Within M. indica, there are two distinct types that can be distinguished on the basis
of their mode of reproduction and their respective centres of diversity: a subtropical
group with monoembryonic seed (Indian type) and a tropical group with
polyembryonic seed (South-east Asian). A few polyembryonic cultivars occur
along the west coast of India; however, they may have been introduced into Goa
from South-east Asia, perhaps by the Portuguese from
Introduction: Botany and Importance 9

their colonies of Malacca in the Malay Peninsula or Timor in the Indonesian


archipelago. Kumar et al. (2001) estimated the genetic relatedness among ten
polyembryonic and monoembryonic cultivars from the west coast of south-ern
India using genomic and chloroplast DNA RFLP analysis. The cultivars could be
grouped on the basis of embryo type (i.e. monoembryonic and poly-embryonic)
and had distinctly different genetic backgrounds. They con-cluded that
polyembryonic mangoes could not have originated in India, and must have been
introduced, probably from South-east Asia.

Domestication of mango

Historical record
It is probable that mango cultivation originated in India, where De Candolle (1884)
estimated that mango cultivation appeared to have begun at least 4000 years ago. In
the early period of domestication, mango trees probably yielded small fruit with
thin flesh. Such fruit can be found today in north-eastern India and in the Andaman
Islands (Anonymous, 1992). Folk selections of superior seedlings over many
hundreds of years would have resulted in larger fruit with thicker flesh. Mukherjee
(1950a, b) described many of these primitive selections from Orissa in north-
eastern India; they demonstrated great variation in fruit shape and size.

The mango is a very important cultural and religious symbol of India.


Buddhist pilgrims Fa-Hien and Sung-Yun mentioned in their travel notes that the
Gautama Buddha was presented with a mango grove by Amradarika (c.500 BC) as
a place for meditation (Popenoe, 1932). According to Burns and Prayag (1921), a
mango tree is depicted in friezes on the stupa of Bharut, which was constructed
c.100 BC. Other travellers to India, including the Chi-nese Hwen T’sung (AD 632–
645), the Arabs Ibn Hankal ( AD 902–968) and Ibn Batuta (AD 1325–1349) and the
Portuguese Lurdovei de Varthema (AD 1503– 1508), all described the mango. The
Indian subcontinent was the birthplace of some of the earliest highly developed
civilizations, and over the centuries, India exerted strong cultural, religious and
commercial influence over South and South-east Asia. In successive waves,
Hinduism, Buddhism and Islam were introduced into South-east Asia from India.
To this day, many com-monly used words in Indonesia are derived from both
Sanskrit and Tamil. One of the most widely used words for mango in Malaysia and
Java (Indone-sia) is ‘mangga’, which is derived from the Tamil ‘manga’. Traders
and monks from India possibly introduced superior selections of mango into South-
east Asia; however, vegetative propagation was unknown in India until after the
arrival of the Portuguese in Goa in the 15th century. Moreover, the most im-portant
mango selections of Thailand, Cambodia, Vietnam, Malaysia, Indo-nesia and the
Philippines historically have all been of the polyembryonic type, and have
traditionally been seed propagated. Until the establishment of Portuguese enclaves
on the coast of India beginning in the late 15th century, mango cultivars did not
exist in India, as there was no known method for vegetatively propagating superior
selections (see Iyer and Schnell,
10 S.K. Mukherjee and R.E. Litz

Chapter 4, this volume). However, under the Moghul emperor Akbar (1556–
1605), the best selections of seedling mangoes were propagated by approach
grafting and were planted in large orchards. The ‘Lakh Bagh’, a mango orchard of
100,000 trees, was planted near Darbhanga in Bihar. Perhaps noth-ing more
eloquently attests to the importance of this fruit and the esteem in which it was held
than this vast mango orchard. The Ain-i-Akbari, an ency-clopedic work that was
written during the reign of Akbar, contains a lengthy account of the mango, and
includes information about the quality of the fruit and varietal characteristics.
There was evidently a strong body of informa-tion about mango cultivation that
had accumulated up to that time. Most of the mango cultivars of India had their
origin in those years, and have been maintained under cultivation for over 400
years by vegetative propagation. ‘Alphonso’, ‘Dashehari’, ‘Langra’, ‘Rani Pasand’,
‘Safdar Pasand’ and other mango cultivars were selected during that time. Relics of
orchards from the time of Akbar are found in different parts of India, and it has
been suggested that they could still provide valuable material for selection of
superior mango cultivars.

Distribution

Spreading from the centres of domestication


The global spread of mangoes and their cultivation outside their original centres of
domestication probably did not occur until the beginning of the European voyages
of discovery and colonialization in the 15th and 16th centuries. Because mango
seeds are recalcitrant, and cannot survive for more than a few days or weeks,
mango germplasm in the early days must have been transported as ripe fruit,
seedlings or, later on, as grafted plants. It is believed that the Portuguese
transported the mango from their colonies in India to their African colonies,
although Purseglove (1972) suggested that it might also have been introduced to
Africa via Persia and Arabia in the 10th century by Arab traders. The Portuguese
later introduced the mango into Brazil from their African colonies of Mozambique
and Angola. Spaniards, who encountered a mango-growing civilization in the
Philippines after Magellan’s passage across the Pacific Ocean, introduced
polyembryonic mango types to their New World colonies through the Pacific
trading ports of Mexico and Panama. The most important, traditional mango
cultivar in Mexico remains the ‘Manila’, reflecting its Philippine origin. ‘Carabao’
and ‘Manila’ are probably identical. The mango was introduced to the West Indies
in the mid- to late 18th century, probably from Brazil. The first introductions of
mango into Florida (USA) occurred in 1861, and involved the ‘No. 11’
polyembryonic seedling from Cuba. Seven years later, another polyembry-onic
selection, ‘Peach’ was introduced into the state (Knight and Schnell, 1993). Many
of the early introductions into Florida proved to be unproduc-tive, although
‘Mulgoba’ was planted on a small commercial scale (this culti-var is referred to as
‘Mulgoa’ in India, ‘Mulgoba’ in the USA and ‘Malgoa’ in Malaysia).
Introduction: Botany and Importance 11

Secondary ‘centres of diversity’


In 1910, a seedling of ‘Mulgoba’ came into production in Florida. Its fruit had a
highly attractive red blush, and appeared to bear more heavily than its parent(s)
(Wolfe, 1962). This selection was named ‘Haden’. Although ‘Haden’ was not
superior with respect to fruit quality in comparison to the imported germplasm
from India, its genetic base was much wider. During the 20th century, more
introductions of mango germplasm into Florida occurred from South-east Asia (the
Philippines, Cambodia), India and elsewhere. It was at one time believed that these
introductions of mango germplasm created a secondary centre of diversity of the
species (Knight and Schnell, 1993). ‘Eldon’, ‘Glenn’, ‘Lippens’, ‘Osteen’,
‘Parvin’, ‘Smith’, ‘Springfels’, ‘Tommy Atkins’ and ‘Zill’ are progeny of ‘Haden’.
‘Saigon’ seedlings were selections made from ‘Cambodiana’, a polyembryonic
introduction from Indochina. From ‘Saigon’ seedlings, ‘Alice’, ‘Herman’ and
‘Florigon’ were selected. Based upon more recent genetic analysis involving
microsatellite markers, it is now estimated that the majority of Florida cultivars are
descended from only four monoembryonic Indian mango cultivar accessions, i.e.,
‘Mulgoba’, ‘Sandersha’, ‘Amini’ and ‘Bombay’, together with the polyembryonic
‘Tur-pentine’ from the West Indies (Schnell et al., 2006). The Florida mango culti-
vars have been found to be highly adaptable to many agroecological areas and bear
regularly, whereas many of the outstanding Indian cultivars have been
unproductive outside their centre of domestication, and are alternate bearing. These
selections also have a highly attractive red blush at maturity, firm flesh, a high
flesh to seed ratio and a regular bearing habit. Some of the Florida cultivars, for
example ‘Tommy Atkins’, ‘Keitt’, etc. are also moder-ately resistant to
anthracnose, the most important production and posthar-vest problem of mango in
many areas. In the latter half of the 20th century, plantings of Florida cultivars have
been established in many countries and now form the basis of international trade of
mangoes.

Current distribution
The mango is cultivated commercially throughout the tropics and in many
subtropical areas. It is grown at the equator and at a latitude of 35–37q in southern
Spain. According to Knight and Schnell (1993), ‘The process that began in Florida
– introduction of superior germplasm from abroad followed by selection of
improved cultivars adapted to local conditions – is now underway in many areas.’

1.4 Germplasm Conservation


Genetic erosion

The Mangifera species have their centre of diversity and origin in South-east Asia,
a region that has experienced great economic development in recent years. Vast
wooded areas have been completely or partially deforested either for expanding
agriculture or for removal of tropical hardwoods for export.
12 S.K. Mukherjee and R.E. Litz

This has caused great genetic erosion within many species and genera. The
Mangifera species, like many other tropical fruit trees, are canopy and emer-gent
trees of the tropical rainforest (Kaur et al., 1980). These trees are widely scattered
in the tropical rainforest, flower erratically and reproduce from large seeds that
deteriorate rapidly. As such, they are particularly vulnerable and in danger of
extinction.

Collection and documentation of Mangifera germplasm

The International Plant Genetic Resources Institute (IPGRI), formerly known as


the International Board for Plant Genetic Resources (IBPGR), commis-sioned an
ecogeographical study of known Mangifera genetic resources (Muk-herjee, 1985).
Based upon this documentation, a joint IBPGR-International Union for the
Conservation of Nature (IUCN)-World Wildlife Fund (WWF) project was initiated
to collect wild mangoes on the island of Borneo and in the Malay Peninsula
(Bompard, 1989), the regions that held the highest con-centrations of Mangifera
species. Kostermans and Bompard (1993), in the lat-est revision of the taxonomy
of Mangifera, recognized 69 species, many of which were collected during the
course of this project (Table 1.1). Because of the loss of natural habitat, the
establishment of in situ and ex situ germplasm collections of Mangifera species
was considered to be imperative.

Relevance of germplasm resources to mango improvement

The genetic improvement of mango hitherto has depended on the utilization of the
genetic variability found within a single species, M. indica. According to
Mukherjee (1985), ‘A concerted sampling strategy should be devised for ex situ
samples to meet urgent needs for use in research for improvement of the crop
through breeding or as rootstocks. Sources of resistance to mango mal-formation,
anthracnose, powdery mildew, gall midge are urgently needed.’

1.5 Importance of Mango

Cultivars

A partial list of the principal mango cultivars has been provided in Table 1.2. This
list includes many cultivars that were identified in a survey of world mango
production compiled by Watson and Winston (1984). The distribution of mango
cultivars outside their centres of domestication can be attributed primarily to three
historical events: (i) the movement of Indian varieties (monoembryonic) along the
trade routes of the Portuguese to Africa and South America; (ii) the spread of
South-east Asian varieties (polyembryonic) across the Pacific Ocean to Central and
South America by the Spaniards; and (iii) the identification of improved mango
cultivars initially in Florida and
Introduction: Botany and Importance 13

Table 1.2. Most important mango cultivars in major producing countries.

Continent Country Cultivars

Africa Cote d’Ivoire ‘Amelie’, ‘Kent’


Egypt ‘Alphonso’, ‘Bullock’s Heart’, ‘Hindi be Sennara’,
‘Langra’, ‘Mabrouka’, ‘Pairie’, ‘Taimour’, ‘Zebda’
Kenya ‘Boubo’, ‘Ngowe’, ‘Batawi’
Mali ‘Amelie’, ‘Kent’
South Africa ‘Fascell’, ‘Haden’, ‘Keitt’, ‘Kent’, ‘Sensation’, ‘Tommy
Atkins’, ‘Zill’
Asia Bangladesh ‘Aswina’, ‘Fazli’, ‘Gopal Bhog’, ‘Himsagar’, ‘Khirsapati’,
‘Langra’
China ‘Gui Fei’, ‘Tainong No. 1’, ‘Keitt’, ‘Sensation,’ ‘Zill’, ‘Zihua’,
‘Jin Huang’
India ‘Alphonso’, ‘Banganapalli’, ‘Bombay’, ‘Bombay Green’,
‘Chausa’, ‘Dashehari’, ‘Fazli’, ‘Fernandian’,
‘Himsagar’, ‘Kesar’, ‘Kishen Bhog’, ‘Langra’, ‘Mallika’,
‘Mankurad’, ‘Mulgoa’, ‘Neelum’, ‘Pairi’, ‘Samar
Behisht’, ‘Suvarnarekha’, ‘Totapuri’, ‘Vanraj’, ‘Zardalu’
' '
Indonesia ‘Arumanis’, ‘Dodol’, ‘Gedong’, ‘Golek’, Madu’, Manalagi’
Israel ‘Haden’, ‘Tommy Atkins’, ‘Keitt’, ‘Maya’, ‘Nimrod’, ‘Kent’,
‘Palmer’
Malaysia ‘Apple Rumani’, ‘Arumanis’, ‘Golek’, ‘Kuala Selangor 2’,
‘Malgoa’
'
Myanmar ‘Aug Din’, ‘Ma Chit Su’, ‘Sein Ta Lone’, Shwe Hin Tha’
Pakistan ‘Anwar Ratol’, ‘Began Pali’, ‘Chausa’, ‘Dashehari’,
‘Gulab Khas’, ‘Langra’, ‘Siroli’, ‘Sindhri’,
‘Suvarnarekha’, ‘Zafran’
The Philippines ‘Carabao’, ‘Manila Super’, ‘Pico’
Taiwan ‘Irwin’, ‘Jin-hwung’, ‘Keitt’, ‘Tommy Atkins’, ‘Tainong
No. 1’, ‘Tsar-swain’
Thailand ‘Nam Doc Mai’, ‘Ngar Charn’, ‘Ok Rong’, ‘Keow Savoey’,
‘Pimsen Mum’
Australia ‘Calypso’, ‘Kensington Pride’
North and Costa Rica ‘Haden’, ‘Irwin’, ‘Keitt’, ‘Mora’, ‘Tommy Atkins’
Central Dominican ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
America Republic
Guatemala ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
Haiti ‘Francine’, ‘Madame Francis’
Mexico ‘Ataulfo’, ‘Haden’, ‘Keitt’, ‘Kent’, ‘Manila’, ‘Palmer’,
‘Sensation’, ‘Tommy Atkins’, ‘Van Dyke’
USA ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
South Brazil ‘Bourbon’, ‘Coite’, ‘Coquinho’, ‘Coracao’, ‘Espada’,
America ‘Haden’, ‘Itamaraca’, ‘Keitt’, ‘Mamao’, ‘Palmer’, ‘Rosa’,
‘Tommy Atkins’, ‘Uba’, ‘Van Dyke’
Colombia ‘Vallenato’
Ecuador ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
Peru ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
Venezuela ‘Haden’, ‘Keitt’, ‘Kent’, ‘Tommy Atkins’
14 S.K. Mukherjee and R.E. Litz

later in other new mango-producing areas, as a result of open and controlled


pollination among local and introduced mango germplasm from India and South-
east Asia.
Further information about many of the mango cultivars, including their fruit
characters, is available in Knight et al. (Chapter 3, this volume), and in
publications by Burns and Prayag (1921) for mangoes of Maharashtra, Naik and
Gangolly (1950) for south Indian mangoes, Singh and Singh (1956) for Uttar
Pradesh mangoes, Mukherjee (1948) for Bengal mangoes and Camp-bell (1992)
for Florida mangoes.
Because many clonally propagated mango cultivars have unique local and/or
regional names, there is considerable confusion in nomenclature. The Indian
Agricultural Research Institute (IARI), New Delhi, has been recog-nized by the
International Society for Horticultural Science (ISHS) as the International
Registration Authority for Mango, whose mission is to consoli-date superfluous
names of mango cultivars. The potential for molecular, for example randomly
amplified polymorphic DNA (RAPD), markers, to resolve much of this confusion
has been demonstrated by Schnell and Knight (1992), Degani et al. (1993), Schnell
et al. (1995), Eiadthong et al. (1999a), Kashkush et al. (2001) and Gonzalez et al.
(2002) (see Bompard, Chapter 2 and Iyer and Schnell, Chapter 4, this volume).

There is little variation among seedlings derived from polyembryonic


mangoes. None the less, a certain amount of variability does occur, probably as a
result of somatic mutation. Thus, in Indonesia there are several ‘Aru-manis’
selections that are denoted numerically, for example ‘Arumanis 1’, ‘Arumanis 2’,
etc. In addition, although Philippine mango cultivars are dis-tinguished by different
names, for example ‘Carabao’, ‘Manila’, ‘Philippine’, etc., the differences among
them are quite subtle.

1.6 Production and Uses


The mango is the most important fruit of Asia, and currently ranks fifth in total
production (in metric tonnes) among major fruit crops worldwide, after Musa
(bananas and plantains) (105,815,354 t), Citrus (all types) (105,440,168 t), grapes
(65,584,233 t) and apples (59,444,377 t) (FAOSTAT, 2006). According to the
Food and Agriculture Organization of the United Nations (FAO) database
(FAOSTAT, 2006), world mango production has increased from 16,903,407 t in
1990 to 28,221,510 t in 2005. Much of this new production has occurred outside
the traditional centres of mango culture of South and South-east Asia. In 1990,
India produced approximately 51% of the world’s mangoes, but by 2005, India’s
share had declined to approximately 38%, despite the substantial increase in mango
production since 1990 (from 8,645,405 to 10,800,000 t between 1990 and 2005).
The current leading producing nations after India include (in metric tonnes) China
(3,450,000), Thailand (1,800,000), Pakistan (1,673,900), Mexico (1,600,000),
Indonesia (1,478,204), Brazil (1,000,000) and the Philippines (950,000). Although
world production has increased by 67% between 1990 and 2005, mango exports
have increased almost sixfold
Introduction: Botany and Importance 15

from 158,030 to 907,782 t, with total export value estimated to be US$583,763,000


(FAOSTAT, 2006). The major exporting countries are (in met-ric tonnes) Mexico
(212,505), India (156,222) and Brazil (111,181). As a result, mangoes are widely
available as fresh fruit and as processed products (i.e. dried fruit, dairy products,
juice, pickles, etc.).
Mangoes are an important component of the diet in many less developed
countries in the subtropics and tropics. In regions of the world that have
experienced low living standards and serious nutritional deficiencies, their
attractiveness and flavour have also enhanced the quality of life. Surplus
production has increasingly been processed and fruit of certain cultivars is destined
for export as fresh fruit. Approximately 1% of mango production is utilized for
processing for juice, nectars, preserves (including chutney), fruit leather, dried fruit
slices, frozen pulp and as a flavouring for baked goods, ice cream, yoghurt, etc.
(see Raymundo et al., Chapter 17, this volume). No part of the fruit is wasted. In
India and the subcontinent, the seed is used for extraction of starch ‘amchur’, and
the peels (skin) have been used as a source of anacardic acid. Mango wood is a low
quality timber, and the bark of the tree is an important source of tannins for curing
leather.

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18 S.K. Mukherjee and R.E. Litz

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2 Taxonomy and Systematics

J.M. Bompard
Les Mazes, Montaud, France

2.1 Introduction 19
2.2 The Genus Mangifera L. 20
Distribution 20
Ecology and habitat 20
2.3 Taxonomy and Systematics 22
Taxonomic history 22
2.4 Phytogeography 28
Species distribution 28
Subgenera and section distribution 29
2.5 Interspecific Molecular Characterization 30
2.6 Region of Origin of the Genus 31
2.7 Origin of the Common Mango 32
The common mango in South-east Asia 32
2.8 Conclusion 35
Potential contribution of wild species to mango cultivation 35
Source of rootstock 35
Hybridization 36
Potential of wild species 36

2.1 Introduction
The genus Mangifera is one of the 73 genera (c.850 species) belonging to the
family of Anacardiaceae, in the order of Sapindales. Anacardiaceae is a fam-ily of
mainly tropical species, with a few representatives in temperate regions. Malesia,
which is the phytogeographic region extending from the Malay Peninsula south of
the Kangar-Pattani line to the Bismarck Archipelago east of New Guinea
(Whitmore, 1975) contains more species in the Anacardiaceae than any other area.
Within Malesia occurrence is mainly in Western Malesia (Ding Hou, 1978b).

 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 19
20 J.M. Bompard

Apart from mango, several other cultivated fruit trees belong to the fam-ily,
for example the ambarella or Otaheite apple (Spondias dulcis Forst.) prob-ably
from Melanesia, and the yellow and purple mombins (Spondias mombin L. and S.
purpurea L., respectively) from tropical America, the Bouea species from
IndoMalesia, dragon plums (Dracontomelum spp.) from IndoMalesia and the
Pacific region, kaffir plum (Harpephyllum caffrum Bernh. ex K. Krause) and the
marula plum (Sclerocarya caffra Sond.) of southern Africa. The cashew
(Anacardium occidentale L.) is from tropical America and the pistachio (Pistacia
vera L.) from Iran and Central Asia. Anacardiaceous species also yield other
valuable products: wood (several genera), gums and resins (Pistacia spp.),
varnishes (Rhus spp. and Melanorrhoea spp., ‘lacquer trees’) and tanning materials
(Rhus spp. and Schinopsis spp.). It is also a family well known for the dermal
irritation produced by some of its members, such as the poison ivies and oaks
(Rhus spp.) in North America, rengas (Gluta spp.) in South-east Asia and other
species including some Mangifera species whose resinous sap may induce a mild
to strong allergic reaction.

2.2 The Genus Mangifera L.

Distribution

The range of natural distribution of the 69 Mangifera species is mainly restricted to


tropical Asia, and extends as far north as 27° latitude and as far east as the
Carolines Islands. Wild mangoes occur in India, Sri Lanka, Ban-gladesh,
Myanmar, Sikkim, Thailand, Kampuchea, Vietnam, Laos, southern China,
Malaysia, Singapore, Indonesia, Brunei, the Philippines, Papua New Guinea and
the Solomon and Carolines Islands. The highest species diver-sity, c.29 species,
occurs in western Malesia, especially in peninsular Malay-sia and in Borneo and
Sumatra, which represent the heart of the distribution range of the genus (Fig. 2.1).

Ecology and habitat

The majority of Mangifera species occur as a rule as scattered individuals in


tropical lowland rainforests on well-drained soils. The species are distrib-uted
mostly below 300 m, but can occur up to c.1000 m above sea level, on well-drained
soils (44 species), in periodically inundated areas (ten species) and in certain types
of swamp forest (i.e. M. gedebe, M. griffithii and M. parvi-folia). Three species are
mainly found in sub-montane forests above 1000 m and occasionally up to 1700 m
above sea level (M. bompardii, M. dongnaiensis and M. orophila). There are also
species that are adapted to seasonally dry climates in deciduous or semi-deciduous
forests (e.g. M. caloneura, M. collina, M. timorensis and M. zeylanica). A few
species occur north of the Tropic of Cancer, for example M. austro-yunnanensis
and M. persiciformis in China, M.
Taxonomy and Systematics 21

80° 100 120 140 160


30° N 30° N

4 3
20° N 20° N
4
13
6
10° N
5 5 10° N
2
27

0° N 28 0° N
27 7 4
9 3 2
5
10° N 10° N

80° 100 120 160

Fig. 2.1. Distribution of Mangifera species in the range of the genus. Numbers shown
indicate the number of wild species in each area: Sri Lanka, India and Sikkim, Andaman and
Nicobar Islands, Myanmar, Thailand, Indochina, China, peninsular Malaysia, Sumatra,
Borneo, Java, Lesser Sunda Islands, Sulawesi, Moluccas, the Philippines, New Guinea, and
Solomon Islands (the Caroline Islands not represented).

sylvatica Roxb. in Sikkim and southern China, at altitudes of 600–1900 m above


sea level; apparently wild M. indica can also be found outside the tropics.
Wild mangoes are large trees, 30–40 m (occasionally 54 m) in height, with tall
columnar boles. Several species are exploited for their timber. The major-ity of
wild mangoes occur as scattered individuals at very low densities in lowland
forests on well-drained soils. Some of these are very rare; there are normally one to
three trees above 40 cm in diameter/10 ha. Only a few spe-cies (M. gedebe, M.
griffithii and M. parvifolia) are gregarious in certain types of swamp forest. Most
species are evergreen although a few are deciduous in the rainforests following a
dry period, and stand bare for a short time before flushes of new leaves appear. A
deciduous habit that is not linked to a sea-sonal climate also occurs in other genera
of Anacardiaceae (Ding Hou, 1978b).
In the rainforest of western Malesia, Mangifera species flower and fruit very
irregularly. As with many other genera in the region, mast or general fruiting at
intervals of 3–8 years is the dominant pattern. In mast years, the ground beneath
the trees can be covered with mangoes, whose strong smell attracts many animals.
Isolated flowering may occur after a dry period and is generally followed by a poor
fruit crop. The occurrence of flowering of a few species, for example the ‘lanjut’
(M. lagenifera) is only once every 5–10 years. There seem to be clear reproductive
barriers between species in the wild, although limited hybridization among
cultivated species has been reported (see section 2.8, Conclusion, this chapter).
22 J.M. Bompard

These widely scattered towering tree species, often with an inaccessible


crown, are undercollected and poorly represented in herbarium collections
(Bompard, 1995). Because of their irregular flowering, the flowers and fruits of a
few species are still unknown. Collecting plant material is consequently very
difficult, and plant explorations are still yielding new records or new species. Many
species have been recently recorded for the first time, even from peninsular
Malaysia, a country that has already been rather well combed by botanists, having
one of the highest collecting indices in the Malesian region. Other species still
await to be discovered. Sadly some species of very limited range may already have
been lost to posterity by deforestation.
Our very meagre knowledge of the wild mangoes is due to the fact that
identification at the species level from leaves only is often difficult because of
intraspecific variation in vegetative characters. Moreover, many of the origi-nal
species were based on very poor specimens. Consequently, frequent mis-
identification of herbarium material has resulted in much confusion, requiring a
critical revision of all the specimens in these collections. It is not uncommon that
the same species has been described from different places under differ-ent names.
For instance, M. inocarpoides described by Merrill and Perry from New Guinea in
1941, M. camptosperma and M. reba (recorded by Pierre in South Vietnam in
1897) are now recognized to be a single species M. gedebe Miquel, a species
initially named in 1861 from a specimen collected in Suma-tra. Mangifera longipes
Griffith is now treated as M. laurina Blume, because this name takes precedence as
it was validly published 4 years earlier.
After thorough study of herbarium collections and field collections, a number
of species have been newly described. Sixty-nine species are now recorded,
including 13 species of uncertain affinities, in contrast with the 49 species
recognized by Mukherjee (1949). As more collections are made, there will
doubtless be further taxonomic adjustments made to the genus Mangifera.

2.3 Taxonomy and Systematics


Taxonomic history

Subdivision of the genus


An historical review of the subdivisions of the genus Mangifera shows that two
major groups have been rather consistently recognized in taxonomic treatments.
Hooker (1862) was the first to recognize two sections based on the characters of
the flower disc: section I with a disc broader than the ovary, and section II with a
disc stalk-like or wanting. These sections were later named by Marchand (1869)
Amba, an Indian name for the common mango, and Limus, a Sundanese name for
M. foetida in West Java, respectively. He also added a section Manga for M.
leschenaultii, which in fact belongs to the section Limus.

In his monograph of the Anacardiaceae, Engler (1883) maintained Hook-er’s


sections, and subdivided group A (Hooker’s section I) into two groups,
Taxonomy and Systematics 23

one group with four or five petals and the other group with four petals. He
considered the following sequence of morphological characters to be impor-tant for
classification: (i) texture of the leaves; (ii) number of fertile stamens;
(iii) prominence of veins; (iv) pilosity of inflorescences; and (v) leaf shape. Pierre
(1897) further divided the genus Mangifera into five sections based
on flower characters, i.e. number of stamens, the attachment of stamens to the disk,
and the style. Two of these five sections – namely section I Euan-therae, with a
short thick flower disc and 4–12 fertile stamens, and section V Marchandora then
consisting of M. camptosperma (currently considered a syn-onym of M. gedebe)
are still maintained as they form clear-cut sections.
In his monograph, Mukherjee (1949) recognized two unnamed sections,
conserving Hooker’s subdivision. Ding Hou (1978a) adopted the same method in
his revision of the Malesian Anacardiaceae recognizing only Hooker’s two original
sections and providing them with proper names and synonyms: section Mangifera
(section I Hooker, section Amba Marchand, group A Engler, sections Euantherae
and Marchandora Pierre) and section Limus (section II Hooker, sections Limus and
Manga Marchand, group B Engler, and sections Eudiscus and Microdiscus Pierre).

Most recent classification of the genus


The taxonomic classification referred to herein follows that proposed by Kos-
termans and Bompard (1993). This treatise includes the results of collections and
surveys carried out between 1986 to 1998 in Borneo and peninsular Malaysia,
which were initiated and sponsored by the International Institute for Genetic
Resources (now Biodiversity International) and the World Wide Fund for Nature. 1
It was published under the auspices of the International Board for Plant Genetic
Resources (now International Plant Genetics Resources Institute) and the Linnean
Society of London.
The most recent treatment of Mangifera reflects the current status of what is
still fragmentary knowledge. It can provide a basis for further studies involving all
aspects of the wild relatives of mango, but particularly their potential in mango
breeding. Determining phylogenetic affinities based upon molecular markers could
change our thinking about relationships among Mangifera species and among the
cultivated forms of M. indica (see Interspe-cific Molecular Characterization
section, this chapter).
The morphological characters used for identification have been placed in the
following sequence of importance:
1. Shape of the floral disc (see section Subdivision of the genus).
2. Number of fertile stamens.
3. Seed labyrinthine or not.
4. Shape of secondary branches of the inflorescences: open or lax panicle,
flowers glomerulate or sub-glomerulate, the ramifications racemoid or spike like.

5. Pubescence of the inflorescence.


6. Shape, number and attachment of the nerves (ridges or fingers) at the inner
surface of the petals.
24 J.M. Bompard

7. Shape and size of the petals.


8. Flowers tetra- or pentamerous (not a very constant character and often
overlapping).
9. Reticulation of the leaves, especially of the lower surface.
10. Shape of the leaf (only fully grown leaves of sterile branches can be used).
11. Texture of the leaves.
12. Deciduous or non-deciduous trees.
13. Colour of the flowers.
14. Shape, colour and smoothness of the fruit.
15. Number and size of the stone fibres.
Kostermans (Kostermans and Bompard, 1993) raised the sections to the rank
of subgenus, i.e. subgenus Limus (Marchand) Kosterm., having a disc narrower
than the base of the ovary, stalk-like or even lacking and subgenus Mangifera
(Ding Hou) Kosterm., having a disc broader than the base of the ovary, cushion-
like, often divided in four or five lobes.

SUBGENUS LIMUS (MARCHAND) KOSTERM. Mangifera species of the subgenus Limus


are quite distinctive and show only remote affinity with the common mango. This
taxon is more primitive than the subgenus Mangifera and may be ances-tral to it,
although the two subgenera may have originated from two different ancestors. The
subgenus Limus consists of 11 species, which are native to the rainforests of
western Malesia (peninsular Thailand, Malay Peninsula, Suma-tra, West Java and
Borneo), with the exception of M. foetida, which extends to the east, possibly as far
as New Guinea, and M. odorata which is only known in cultivation.

Kostermans divided the subgenus Limus into two sections: (i) section
Deciduae for deciduous trees (i.e. M. caesia, M. kemanga, M. pajang, M. superba
and possibly M. blommesteinii, M. decandra and M. lagenifera); and (ii) section
Perennes for non-deciduous species (i.e. M. foetida, M. leschenaultii, M. macro-
carpa and M. odorata) (Kostermans and Bompard, 1993). In deciduous trees, the
bracts enclosing the buds leave a characteristic collar of dense, narrow scars, which
persist on old twigs and are especially prominent in M. caesia and M. kemanga.

Mangifera lagenifera and M. decandra have ten stamens, five of which are
fertile. The other nine species have only one (and rarely two) fertile stamen(s) and
two to four staminodes. The two species with five fertile stamens (M. decandra and
M. lagenifera) and M. superba, M. caesia, M. kemanga and M. blom-mesteinii,
whose leaves are apically aggregated into rosettes at the end of mas-sive twigs are
particularly distinctive. The fruits of these species are broadly ellipsoid or pear
shaped, not compressed, and have dirty whitish or pinkish mesocarp and
lanceolate, and fibrous, non-ligneous leathery endocarp.
Mangifera subsessilifolia shows some affinity with M. lagenifera and M.
blommesteinii; however, it has been placed among the species of uncertain
taxonomic position due to a lack of complete study material. This is not a very rare
species, but flowering and fruiting seem to occur at intervals of, or > 5 years,
similar in this respect to M. lagenifera, which can be found growing
Taxonomy and Systematics 25

in old orchards in peninsular Malaysia. The flowers and fruits of M. sub-sessilifolia


are still unknown.
Mangifera foetida, M. odorata, M. caesia and M. kemanga are widely
cultivated in the humid lowlands of the Malay Peninsula, Sumatra, Borneo, Java
and Bali. They have also been introduced elsewhere in South-east Asia; M. caesia,
M. foe-tida and M. odorata are grown in the southern part of the Philippines, M.
foetida is grown in Myanmar, and M. odorata is found in Indochina. They have
been described in general reviews of tropical fruit (Ochse and Bakhuizen, 1931;
Ochse et al., 1961; Molesworth, 1967; Verheij and Coronel, 1991).
Mangifera pajang, an endemic and commonly cultivated species in Bor-neo, is
hardly known outside its native island. This deciduous tree has very stout twigs,
with leaves more or less aggregate at the apices. The globose fruits, up to 20 cm in
diameter, are the largest known fruits in the genus. The rough, potato-brown rind
(0.5–1 cm thick) can be peeled off like that of a banana. Its bright, deep yellow,
thick and fibrous flesh is sweet with a dis-tinctive taste (Kostermans, 1965;
Bompard, 1991a). In orchards in Borneo where M. foetida and M. pajang are both
cultivated, forms with leaves and fruits having intermediate characters are
occasionally found.
Mangifera caesia, M. foetida, M. pajang and especially M. odorata are impor-
tant in tropical humid regions where the common mango cannot be grown
satisfactorily. Mangifera pajang has potential as an ornamental tree, having
brilliant rose-red blossoms (Philipps et al., 1982).

SUBGENUS MANGIFERA. The subgenus Mangifera contains most of the species (47), and
is divided into four sections: (i) section Marchandora Pierre; (ii) section Euan-therae
Pierre; (iii) section Rawa Kosterm.; and (iv) section Mangifera Ding Hou.
Section Marchandora Pierre. This section has only one species, M. gedebe Miquel
(syn. M. camptosperma Pierre, M. inocarpoides Merr. and Perry, M. reba Pierre). The
labyrinthine seed is unique to this species, wherein the inner integu-ments penetrate the
cotyledons and form numerous irregular folds. The flat, discus-like fruit has only a very
thin mesocarp. Mangifera gedebe grows in inundated places along rivers or lakes. The
seed floats in water and is dis-persed during periods of high water, and this may explain
its wide distribu-tion, from Myanmar through Malesia to New Guinea and the
Bougainville Island.

Section Euantherae Pierre. The three species in this section (M. caloneura Kurz (syn.
M. duperreana Pierre), M. cochinchinensis Engler and M. pentandra Hook. f.) appear
to be the most primitive among the species of the subgenus Mangifera. The flowers are
characterized by the presence of five fertile sta-mens. The three species are mainly
confined to Myanmar, Thailand, Indo-china and the north of the Malay Peninsula. The
region is in the transition zone from the humid tropical rainforest to monsoon forest,
and these species show an adaptation to low rainfall. Mangifera cochinchinensis, which
occurs in south-eastern Thailand and in Vietnam, has small oblong fruits with a thin
seed; the fruits are much relished by local people in southern Vietnam, although they
are very acidic. Mangifera caloneura and M. pentandra are closely
26 J.M. Bompard

related, and can be mistaken for M. indica. However, their leaves are more
leathery, have a more conspicuously dense reticulation, and the panicles are much
more hirsute than the common mango. Mangifera caloneura occurs from Myanmar
through Thailand to Indochina, in lowland evergreen forests, as well as in semi-
deciduous forests. It is cultivated for its acidic-sweet fruit, and has been planted
along the streets of Vientiane and Ho Chi Minh City (Saigon). Mangifera
pentandra, apparently native to the northern Malay Pen-insula close to the Kra
isthmus transition zone, is found in old orchards, in scattered locations, especially
in Kedah and possibly also in peninsular Thai-land. It is also grown in the
Anambas Islands and in Sabah, where it might have been introduced in early times.
It is a prolific bearer, with small man-goes, c.8 cm length, and ripening green or
yellow. The pale orange, watery pulp has a sweet taste and few fibres.

Section Rawa Kosterm. This group, consisting of nine species, is not well delimitated.
Most species have thick twigs and rather coriaceous leaves seated on protruding
pedestals. The small, hardly flattened ovoid or ellip-soid fruits that are black or partly
red at maturity in several species are also characteristic. ‘Rawa’ is the Malay word for
marsh, indicating that these spe-cies usually are found in periodically or permanently
inundated areas. The five species that occur in west Malesia (M. gracilipes, M.
griffithii, M. micro-phylla, M. paludosa and M. parvifolia) grow primarily in the
swamps of south peninsular Malaysia, in central coastal areas of east Sumatra and
western Borneo, and occasionally in peripheral uplands. It has also been reported from
the Andaman Islands and from Thailand (Sreekumar et al., 1996; Eiad-thong et al.,
2000a).

Mangifera andamanica and M. nicobarica are endemics from the Andaman


and Nicobar Islands, respectively. Mangifera merrillii is a rare species endemic to
the Philippines and M. minutifolia is known solely from a single collection from
southern Vietnam. Mangifera griffithii and M. microphylla are the only cultivated
species within section Rawa. The former species is considered to be representative
of the section, and is cultivated along the eastern coast of peninsular Malaysia and
in western Borneo, and rarely in Sumatra. The fruits are small (3–5 cm long) and
oblong or ovoid; the skin is rose-red, turning purplish black at maturity. The rind is
thin and easily removed from the orange-yellow pulp, which is juicy and pleasantly
sweet. Different forms are recognized by local people, according to the size and
taste of fruits. Mangifera microphylla is a related, but less well-known species,
having thinner leaves and a rather similar fruit.

Section Mangifera Ding Hou. With more than 30 species, section Mangifera is by far
the largest. The common mango and the related M. laurina belong here. Species within
the section have the same distribution range as the genus. The section may be divided
into three groups based on floral structure and organ number variation: (i) those having
pentamerous flowers; (ii) those having tetramerous flowers; and (iii) an intermediate
group of species hav-ing both pentamerous and tetramerous flowers. Within these three
groups, it is possible to distinguish species with either puberulous or glabrous panicles.
Taxonomy and Systematics 27

Only characteristics of representative species within each group, especially those


found in cultivation, are described below.
Pentamerous flowers (14 species): Three species, M. laurina, M. minor and M.
sylvatica, show affinity with the common mango. Mangifera laurina is a species of
the lowland forests of Malesia, where it is also under cultivation in old orchards. It
can be distinguished from the common mango by having lax and widely pyramidal,
glabrous or sparingly puberulous panicles. The flow-ers are smaller and are not
glomerulate; the petals have a different shape, texture and colour. The fruit
resemble those of a small common mango, with orange-yellow pulp, which is
almost liquid at maturity. It is generally con-sumed when unripe. Several forms are
in cultivation; however these are now becoming rare. Mangifera laurina is well
suited to the humid tropical lowlands, fruiting well in areas where the common
mango cannot be grown satisfacto-rily; moreover, it appears to be highly resistant
to anthracnose (Bompard, 1991b).

Mangifera minor occurs east of Wallace’s line, from Sulawesi to New Guinea
(east Malesia) and to the Carolines Islands in the east. It is adapted to a wide range
of ecological conditions, growing equally well in dry savannahs and in tropical
rainforests up to 1300 m. The fruit is obliquely oblong, 5–10 cm long, much
narrowed, the tip obtuse, with a distinct beak and sinus. It is found in cultivation,
although the yellowish fruit pulp is acidic and scant. Mangifera sylvatica is found
from Sikkim (up to 1200 m) to northern Myan-mar and Thailand, and apparently
also in Yunnan up to 1900 m. The fruit is obliquely ovate, 8–10 cm long, much
compressed distally forming a hook, has scanty whitish-yellow pulp which is
almost fibreless. Other species are occasionally found in cultivation, for example
M. rufocostata, which is esteemed by the Banjarese people of South Kalimantan
for its very sour fruits that are used to prepare a spicy condiment with chilli.

Tetramerous flowers (15 species): Mangifera altissima is apparently endemic


to the Philippines, where it occurs mainly at low elevations in the forests from
northern Luzon to Mindoro (Brown, 1950). The fruit is mango shaped, ovoid or
ellipsoid, slightly compressed, up to 8 cm length, green or some-what yellow when
ripe, with whitish, sweetish-acidic flesh. It is commonly found in dooryards, and
thrives in regions with distinct wet and dry seasons (Angeles, 1991).

Mangifera torquenda occurs wild in west Malesia, and is cultivated in south


Sumatra and in Borneo, where it is common in the forests and orchards of eastern
Kalimantan. The sub-globose fruit, c.7.5 cm long and 6.5 cm in diameter, is
yellow-green with darker spots at maturity, and has a thin rind. The pale yellow
pulp has a rather pleasant sweet-acid, slightly resinous taste and a light turpentine
smell. Short fibres are attached to the seed. It is closely related to M. longipetiolata.

Mangifera magnifica is a common species in the rainforests of western


Malesia, occasionally cultivated in central Sumatra and in West Kalimantan, where
it has a special importance in the myths of Land Dayak peoples. The fruit is ovoid
oblong, up to 12 cm long, 10 cm in diameter, only slightly compressed, greyish
green with brown spots. The pulp is whitish, soft at
28 J.M. Bompard

maturity, sweetish acid. Sweeter forms are reported in central Kalimantan (J.J.
Afriastini, personal communication). The stone is unique in the genus in that it
lacks fibres adhering to it.
Mangifera quadrifida is found from peninsular Malaysia to the Moluccas. The
fruit is ellipsoid-globose, 6–8 cm long, green covered with black dots turn-ing
completely black at maturity, and has a pale yellow, sweet-acid pulp. Another form
is recognized by its more coriaceous leaves, smaller fruits, c.4 cm long, having
dark yellow pulp, purplish around the stone, and a sweet, palat-able taste,
somewhat like prunes. Both forms are cultivated in old orchards.
Tetra- and pentamerous flowers (four species, and also M. indica): Mangifera
casturi is related to M. quadrifida, from which it can be distinguished by leaf and
fruit characters. It has never been collected in the wild, and is a favourite among
the Banjarese people in south Kalimantan. The fruits are small, a little compressed
and up to 6 cm in length, becoming completely black at matu-rity. The orange pulp
is very sweet and palatable, and resembles ‘honey mango’ or ‘mangga madu’
grown in East Java. Although M. casturi bears heavily, it has a strong- to alternate-
bearing habit. It is an excellent fruit for the humid tropical lowlands, and appears to
be resistant to anthracnose. Sev-eral differently named forms exist; these have
polyembryonic seeds. Mangifera rubropetala is also only known in cultivation, and
may be a primitive race of M. indica.

SPECIES OF UNCERTAIN TAXONOMIC POSITION. There is a group of 11 disparate spe-cies of


uncertain taxonomic position that cannot be placed with certainty due to the absence of
adequate material. There are three species only known in China.

2.4 Phytogeography
Species distribution

An examination of the present distribution of the genus shows that the larg-est
number of Mangifera species in either subgenera is found in western Malesia on
the Sunda shelf. A decreasing number of species occurs towards the genus
boundary east of Wallace’s line in east Malesia, and in its northern and western
range of distribution. While peninsular Malaysia and the islands of Sumatra and
Borneo have the highest diversity of species, the number of species becomes
gradually lower in east Malesia, especially in the Lesser Sunda Islands, Moluccas
and New Guinea. This is explained by the geologic and paleogeographic features
of the Malesian region which spans two large partly submerged continental
shelves, the Asiatic shelf (Sunda Shelf linking the Malay Peninsula with the islands
of Sumatra, Java, Borneo and Palawan) and the Australasian shelf (Sahul Shelf
linking the Aru islands and New Guinea with Australia). During the last glaciation
period (c.22,500–11,000 BP) the shelves were regions of land uncovered by the
lowering of sea level, and present day peninsular Malaysia, Sumatra and Borneo
were connected by
Taxonomy and Systematics 29

land bridges during the late period of maximal sea lowering. During the cool
periods of glacial maxima, the Malesian forest was reduced in extent, but there is
no evidence that it was reduced to isolated island forests. The Sunda-land and
Papuasian rainforest blocks are therefore comparable to refugia in terms of species
richness and the high degree of endemism (Whitmore, 1981). Mangifera has
undergone major species development in west Malesia, which has remained
relatively stable over a long period of time. The current vegeta-tion of west Malesia
probably differs very little from that at the end of the Tertiary (van Steenis, 1950).
A lower number of Mangifera species is found in Java and the Philippines, regions
less often connected with Asia during the Pleistocene.

Only three species occur in New Guinea, which is largely covered with
rainforest. These include M. minor, M. mucronulata and the widely distributed M.
gedebe. Mangifera foetida also occurs, but may have been introduced. Mangifera
minor occurs from Celebes and the Philippines to the Solomon Islands; M.
mucronulata is found in the Moluccas, New Guinea and the Solomon Islands. The
distribution of these species suggests a late immigration of a Laurasian genus from
Sundaland via the Philippines, Sulawesi and the Moluccas into New Guinea, which
is supported by the geological history of the region. No Mangifera species have
ever been recorded from northern Australia.
Very few species are found in peninsular India and Sikkim. From present-day
distribution, there is little evidence of migration of species into the sub-continent of
India after its collision with Eurasia in the middle Eocene (Audley-Charles et al.,
1981). According to Mehrotra et al. (1998), fossil leaves described as
Eomangiferophyllum damalgiriensis Mehr. from the Upper Palaeo-cene in north-
eastern India are an analogue of the modern genus Mangifera.
Mangifera sylvatica occurs along the northern limit of the range of Mangifera,
with more or less discontinuity, from Sikkim to northern Thailand and to the
southern part of Yunnan, where it is reported in mountains up to 1900 m above sea
level (Anonymous, 1980). The few species that grow in southern China are very
poorly known: M. austro-yunnanensis from western Yunnan, M. persiciformis
from south-eastern Yunnan and southern Guizhou at latitudes up to 26°N and M.
hiemalis, the ‘winter mango’ from Guangxi near the northern border Vietnam. In
the revised Flora of China (Min and Bar-fod, 2008), M. austro-yunnanensis is
considered to be conspecific with M. indica, M. hiemalis is treated as a synonym of
M. persiciformis, and M. laurina is recorded from the lowland forests of south
Yunnan.

Subgenera and section distribution

The species distribution is especially meaningful when the ranges of the spe-cies of
each subgenus and section are considered separately.

Subgenus Limus
All species of the subgenus Limus are restricted to the Malesian area (M. foetida
and M. macrocarpa occurring in peninsular Thailand), whereas all the species
30 J.M. Bompard

with five fertile stamens, considered the most primitive condition, are con-fined to
west Malesia (M. decandra in Sumatra and Borneo; M. lagenifera in the two latter
areas and in peninsular Malaysia). Only M. caesia, M. foetida and closely related
M. leschenaultii occur in east Malesia.

Subgenus Mangifera
In the subgenus Mangifera, M. gedebe is the only species belonging to the sec-tion
Marchandora, and has the widest range within the genus, extending from
Myanmar through Malesia to New Guinea and Bougainville Island. Section
Euantherae is centred in the region from Myanmar to Vietnam. Mangifera
pentandra is only known from peninsular Malaysia, the Anambas Islands and
Borneo. Section Rawa is mainly in western Malaysia and shows notable
diversification in the swamps and peripheral uplands in the south of penin-sular
Malaysia, east central Sumatra (notably the Riau province) and west Borneo.
During the glacial period this area, termed the ‘Riouw pocket’ (Cor-ner, 1978),
formed a vast plain connecting the Malay Peninsula, Sumatra and Borneo, and is
believed to have been filled with swamps. Mangifera merrillii is an endemic of the
Philippines, M. minutifolia is an endemic of Vietnam, M. andamanica and M.
nicobarica are endemics of the Andamans and Nicobar Islands. None of the
species of section Mangifera occurring in mainland South-east Asia, north of the
isthmus of Kra, are found in eastern Malesia; however, it would be interesting to
assess the genetic relatedness of M. syl-vatica and M. minor, and also M. laurina,
which may prove to be phylogeneti-cally very closely related.

2.5 Interspecific Molecular Characterization


Molecular biology techniques now make it possible to assess genetic related-ness
in a more precise way. Published data support some of the groupings based on
anatomical characters (Kostermans and Bompard, 1993) but not entirely.
RAPD (random amplification of polymorphic DNA) markers were first used
in mango by Schnell and Knight (1993) and Schnell et al. (1995). Nine Mangifera
species were analysed and compared to the traditional taxonomic groupings. The
unweighted pair group method of arithmetic averages (UPGMA) cluster analysis
for the subgenus Limus was not supportive of the separation between sections
Perennes and Deciduae, which, admittedly, has a weak taxonomic basis. It
confirmed the relatedness between M. foetida and M. pajang. The UPGMA cluster
analysis of the subgenus Mangifera supported the current taxonomy based on
flower morphology. It showed the related-ness between M. quadrifida and M.
torquenda (both placed in the group of species with tetramerous flowers), but also
with M. casturi, although the lat-ter species has tetra- and pentamerous flowers.
One of the most significant results was the evidence for the existence of
interspecific hybridization within the studied species of the section Mangifera (see
also Yonemori et al., 2002).
Phylogenetic relationships among 14 Mangifera species of Thailand were
analysed by comparing amplified fragment length polymorphism (AFLP)
Taxonomy and Systematics 31

markers (Eiadthong et al., 2000b), and by comparing sequences of the inter-nal


transcribed spacer (ITS) region of nuclear ribosomal DNA (nrDNA) (Yonemori et
al., 2002). They demonstrated that the common mango was closely related to M.
laurina, M. sylvatica and M. oblongifolia of subgenus Mangifera to which M.
indica belongs. A close relationship between M. indica and M. sylvatica has been
corroborated by Nishiyama et al. (2006), who com-pared signal intensity of
genomic in situ hybridization (GISH) on somatic metaphase chromosomes of M.
indica, using labelled DNA of eight wild Mangifera species.

Furthermore, Eiadthong et al. (2000b) and Yonemori et al. (2002) have


demonstrated that M. odorata, M. foetida and M. macrocarpa (of subgenus Limus)
were related to M. indica. It is not surprising in the case of M. odorata whose
hybrid origin (M. foetida × M. indica) has now been established, but this calls into
question the position of the section Perennes.
Results of molecular studies do not permit a comprehensive view of the
phylogenetic relationships among the genera. So far, they are rather support-ive of
the groupings based on phenotype within the subgenus Mangifera (notably sections
Rawa and Euantherae), but not for the subgenus Limus which will need to be
redescribed, and likely restricted to the group of species related to M. caesia (M.
kemanga, M. lagenifera, M. superba and possibly M. decandra). More studies will
be needed to infer phylogenetic relationships within the section Mangifera.
Keeping in mind the frequent misidentifications in collections and botanic gardens,
herbarium specimens of studied material must be deposited in the national herbaria
so that its taxonomic position can be ascertained in case of doubt.

2.6 Region of Origin of the Genus


Based on morphological, phytogeographical and fossil evidence, Mukherjee (1953)
argued that:
although the highest number of species of both sections is concentrated in the Malay
Peninsula [19 were then recorded], the centre of origin of the genus cannot be
restricted to that area alone, as both the phylogenetically older species, i.e. with
pentacyclic flowers (M. duperreana, now reduced to M. caloneura, and M.
lagenifera), occur in Siam and Indochina, and the former is absent from Malay
Peninsula.
He concluded that the genus had its origin somewhere in the Myanmar– Thailand–
Indochina area or in the Malayan area. Careful identification of the greatest part of
herbarium materials available has allowed a more accurate delimitation of the
distribution ranges of the Mangifera species, notably of the subgenus Limus, and
has revealed, among other things, that M. lagenifera does not occur north of Kra
isthmus contrary to Mukherjee’s assertions. Fur-thermore, the ten-stamen species,
M. decandra, which was described by Ding Hou in 1972 and hence was unknown
to Mukherjee, is confined to Borneo and Sumatra, and to date has not been
recorded from peninsular Malaysia.
32 J.M. Bompard

Without overemphasizing the present great species diversity of subgen-era in


the Malay Peninsula, Borneo and Sumatra, the available evidence points to a
Sundaic origin for the genus. This, however, must not minimize the particular
importance of the region stretching from Myanmar to Indo-china as another centre
of diversification, as attested by the range of species belonging to the section
Euantherae. Unfortunately, many of the species of this region remain poorly
known, and it can be expected that plant collecting in the region will yield
interesting new findings. The speciation that occurred in this region with a likely
radiation centre today traced by the range of the section Euantherae, is of special
significance as it has given rise to the com-mon mango.

2.7 Origin of the Common Mango


The common mango apparently originated in regions on the western border of the
secondary centre of diversification mentioned above. Truly wild mango trees have
been recorded in Bangladesh (Chittagong Hill tract, c.23°N), north-eastern India
(‘undoubtedly indigenous in the evergreen tracts of valley of Assam’ according to
Kanjilal et al., 1937), and in Myanmar where it was reported as ‘not unfrequent in
the tropical and lower mixed forests all over Burma from Arracan and Pegu down
to Tenasserim’ (Kurz, 1877). It would be desirable to assess its affinity with the
species of the section Euantherae, as well as with species of other sections of the
subgenus Mangifera that occur in the same area and region. It is also believed to be
wild ‘in the sub-Himalayan tract, in deep gorges of the Baraitch and Gonda hills in
Oudh, and the outer hills in Kamaon and Garhwal’ (Brandis, 1874). The common
mango has been grown and disseminated for such a long time in India that semi-
wild trees can be found in the forests throughout the subcontinent. The fruits of
wild trees are said to be small and of poor quality. Watt (1891) mentioned two so-
called ‘almost unaltered wild varieties’ existed under cultivation in Tirhoot, ‘one
originating from Kangra, a very variable one, and the other from Sikkim which was
evidently the progenitor of the varieties cultivated in Malda’.

The common mango in South-east Asia

The Linnean binomial (Mangifera indica) indicates in this instance the place where
the common mango was selected and improved, and not necessarily its place of its
origin. It has been traditionally accepted that mango was domesti-cated several
millennia ago in India (see Mukherjee and Litz, Chapter 1, this volume); however,
it cannot be excluded that domestication occurred inde-pendently in several areas,
possibly in the south-western and south-eastern regions of its centre of origin, or
later differentiated in those two regions. This hypothesis would account for the
differences that exist between the local polyembryonic cultivars of Myanmar,
Thailand, Indochina and Indonesia, and the monoembryonic Indian cultivars. Note
that polyembryony occurs
Taxonomy and Systematics 33

also in the cultivated M. casturi, M. laurina and M. odorata. Aron et al. (1998)
have demonstrated that polyembryony in mango is under the control of a single
dominant gene.
According to Juliano (1937), Bijhouwer suggested that there were two main
centres of domestication of mango, ‘one in India with monoembryonic mangoes,
the other in the Saigon area, Indonesia and the Philippines with polyembryonic
mangoes’. The ‘Saigon’ area must in fact be extended to southern Vietnam, other
parts of Indochina, Thailand and Myanmar, which were recognized by Valmayor
(1962) as homes of polyembryonic mangoes. Notwithstanding, the origin of
polyembryonic mangoes is probably better placed in Myanmar, and possibly the
eastern part of Assam. According to Brandis (1874), ‘in Burma, the mango is not
generally grafted, and seeds of a good kind, as a rule, produce fruit of a similar
description’. There are only a few polyembryonic mango cultivars in India. They
are restricted to the south-western coastal region, and geographically isolated from
the polyembryonic mangoes of Myanmar and South-east Asia. Analysis of genetic
relatedness using RAPD markers among polyembryonic and monoembryonic
cultivars grown in the west coast of southern India suggest that the polyembryonic
types are unlikely to have originated from India and might have been intro-duced
from South-east Asia (Ravishankar et al., 2004).

Indian Buddhist monks might have introduced the common polyembry-onic


mango to South-east Asia, first along land trade routes through Myan-mar, where
they might have found better races, and from there into insular South-east Asia. It
is well established that some local names of the common mango currently used in
parts of Indonesia are of Sanskrit origin (‘ampelam’ and its cognates), and are
sometimes used to designate M. laurina, which is a truly native species. Vernacular
names do not always travel with a plant, and even if they did so in the case of the
common mango, it is very unlikely that these introductions were the first ones and
that they came obligatorily from India. In the absence of a comprehensive
classification of the innumerable South-east Asian cultivated forms of the common
and wild mangoes, includ-ing the countless primitive races, we have to rely on
linguistics and the rich history and prehistory of this region.

Vernacular names
The different local names of the common mango in Indonesia (‘pauh’, ‘ampe-lam’
and its variants, and ‘mangga’) bear evidence of a long history of con-tacts with
mainland Asia and India, and point to possible introduction at different times from
different places. In some parts of Indonesia, the vernacu-lar names ‘paoh’ or ‘pauh’
refer either to primitive races of the common mango, or to native species, as a rule
the ones most closely resembling the common mango, for example: ‘pauh asal’ (=
native mango) for M. pentandra in peninsular Malaysia; ‘pahohutan‘ or ‘pahutan’
(= forest mango) for M. altissima in the Philippines; and ‘pao pong’ (= forest
mango) for M. minor in Flores, Lesser Sunda Islands. ‘Pau’ is a word belonging to
Austronesian lan-guages, nowadays spoken over a very wide area from
Madagascar to the Easter Islands by people who originate from mainland Asia.
These languages
34 J.M. Bompard

are still spoken by certain minority populations in Vietnam, Cambodia and the
Mergui Archipelago off the coast of Myanmar (Bellwood et al., 1995). In
Cambodia, which was occupied by the Chams from about the 3rd to the 15th
century AD, ‘pa:uh’ is a Chamic word. ‘Sva:y’, used by the Khmers (as in ‘sva:y
srok’ meaning mango of the village (M. indica), and ‘sva:y prey’, wild mango (i.e.
M. caloneura) as attested in pre-Angkorian Khmer inscriptions dating from the 6th
to the 8th century AD (Pou and Martin, 1981)) is of Austro-Asiatic origin. ‘Sva:y’
has cognates in south Vietnam (‘xoay’) and in Asian languages spoken by
aboriginal people in peninsular Malaysia. ‘Wai’, another cognate, is a vernacular
name of M. minor in several parts of New Guinea. Pawley and Ross (1995)
proposed ‘wai’ and ‘pau(q)’ as the reconstructed Proto-Oceanic terms referring,
respectively, to a generic name for mango, and a species that is probably M. indica.

Nowadays, these two words are generic terms for mango fruits that rather
closely resemble M. indica. In the same way, ‘thayet’ which is the com-mon
vernacular name referring to M. indica in Myanmar (‘sinnin thayet’ and ‘taw-
thayet’ for M. caloneura and M. sylvatica, respectively), or ‘mamuang’ in Thai
languages are probably generic names.
Obviously, linguistic evidence alone provided by these vernacular names is not
sufficient to prove the time and place of an introduction. None the less, it is
significant that in mainland South-east Asia none of the vernacular names of the
common mango exhibits signs of an Indian influence, moreover, cog-nates of these
names are also applied to primitive races in some parts of insular South-east Asia.

Evidence of early trade in South-east Asia


The history of plant domestication in mainland South-east Asia has undoubt-edly
involved introduction of plants by people migrating from the mainland into insular
South-east Asia. In more recent times, there is evidence of con-tacts and sea trade
since at least the first centuries AD between mainland and insular South-east Asia
to indicate that there have been numerous opportuni-ties for introduction of the
common mango from different places at different times prior to the 4th century
(before the Indianization of early South-east Asian states) into present-day
Malaysia and Indonesia.
Recent studies based on archaeological evidence stress the long unrecog-nized
importance of South-east Asian trade (emanating from South-east Asia) between
ports established along the Java Sea, those of mainland Asia, and India, back to the
1st century AD, and possibly earlier (Walker and San-toso, 1984). Trade routes
connected the developing population centres of the mainland, such as the earliest
known South-east Asian political entity, Funan, an advanced agrarian society
located on the southern Vietnam coast, which became influenced by the Indians
and reached the zenith of its commercial prosperity in the middle of the 3rd century
(Hall, 1985). Increasingly, king-doms organized according to the Indian concept of
royalty were established in the Indonesian archipelago, for example Kutai in East
Kalimantan (4th century) and Central Java (8th to 9th century), the latter being
famous for the Buddhist temple at Borobudur, where sculptures depict the mango
tree.
Taxonomy and Systematics 35

It is highly probable that the eventual introductions of superior cultivars of


polyembryonic mangoes from the south-west coast of India, ‘between the 6th and
14th century, the height of classical South-east Asian civilization and also the
golden age of early south Indian civilization’ (Hall, 1985), were not the first ones.

During the 16th and 17th centuries, the Portuguese and Spaniards con-tributed
to the widest distribution of superior varieties in the archipelago, espe-cially to the
east. The name mango itself derives from the Tamil ‘man-kay’ or ‘man-ga’ (see
Mukherjee and Litz, Chapter 1, this volume), which the Portu-guese adapted as
‘manga’ and ‘mangueira’ when they colonized west India.
Superior Philippine cultivars originated through introduction of culti-vars from
Indonesia, for example ‘Dodol’ into Mindanao, and from Indo-china, for example
‘Carabao’ and ‘Pico’ in Luzon, the Visayas and northern Mindanao (Wester, 1920;
Bondad et al., 1984). However, these introductions dating from the first half of the
17th century were also preceded by the intro-duction of primitive races of the
common mango as well as other species into the Sulu Archipelago and Mindanao
through contacts with north Borneo, as attested by their local names quoted by
Wester (1920), that is mampalam (M. indica, and possibly also M. laurina), baonoh
(M. caesia) and wannih (M. odorata).

The South-east Asian M. indica germplasm includes many races that defy
classification. Natural cross-pollination has undoubtedly occurred with native
species, such as M. laurina, which was also brought into cultivation in several
areas before the introduction of M. indica.

2.8 Conclusion

Potential contribution of wild species to mango cultivation

To date, the improvement and breeding of common mango has depended on the
use of genetic variability within a single species, M. indica. Mukherjee (1957)
observed that ‘similarity in chromosome number and pollen morphol-ogy in
different species suggests close compatibility during hybridization and stock-scion
relationship if other species are used as stock for the com-mon mango’.
Biotechnology opens new perspectives for mango improve-ment (Litz, 2004). As
noted by Litz et al. (Chapter 18, this volume), the transformation of mango with
genes from other species could address a number of plant breeding objectives.

Source of rootstock

Grafting experiments between M. indica and other species are reported in the
literature, for example budding of M. indica on M. foetida and M. odorata in Java
(Ochse and Bakhuizen, 1931), M. odorata on M. indica in the Philippines (Wester,
1920), and M. indica on M. zeylanica in Sri Lanka (Gunaratman, 1946).
36 J.M. Bompard

Mangifera indica ‘Madu’ in Java, and M. laurina in Sabah have been used as
rootstocks for M. casturi. Trials of grafted M. caesia on M. indica (Wester, 1920)
and M. indica on M. kemanga or M. caesia (Ochse and Bakhuizen, 1931) were
unsuccessful, as these two species have distinct bark features and only remote
affinity with the common mango. Better compatibility can be expected using
species more closely related to the common mango within the subgenus Mangifera.
In West Kalimantan, M. laurina is occasionally used as a rootstock for the common
mango on periodically inundated riverbanks. It has been tried as a rootstock by the
Department of Agriculture in Sabah (Lamb, 1987). Campbell (2004) reported that
M. casturi, M. griffithii, M. laurina, M. odorata, M. pentandra and M. zeylanica
grafted on M. indica had a high percentage of success.

Several species that can grow in permanently inundated areas (i.e. M. gedebe,
M. quadrifida, M. griffithii and other species of the section Rawa) repre-sent a
potential source of rootstock for the development of mango cultivation on poorly
drained soils or in areas liable to prolonged flood. Other species may be a source of
dwarfing rootstocks.

Hybridization

From our observations in Borneo, natural interspecific hybridization involv-ing


various cultivated Mangifera species can occasionally occur. Suspected hybrids
were observed between wild M. gedebe and cultivated M. laurina in the lakes area
along the Mahakam River in East Kalimantan, where impor-tant populations of M.
gedebe occur; between cultivated M. foetida and M. pajang, two species showing
close affinity, in different areas of Kalimantan where both species are grown
together; and between closely related M. caesia and M. kemanga in cultivation. A
hybrid origin has been suggested for M. odorata (M. indica × M. foetida), which is
unknown in the wild (Ding Hou, 1978a). Based on AFLP analysis, Teo et al.
(2002) and Kiew et al. (2003) have con-firmed that M. odorata is a hybrid between
M. foetida and M. indica. The index of similarity showed that M. odorata is closer
to M. foetida (76% similarity) than it is to M. indica (66%). Yamanaka et al. (2006)
showed a high genetic similarity among 11 landraces of M. odorata from the
Malaysian Agricultural Research and Development Institute (MARDI) gene bank.
Higher variability can be expected from Sumatra and Java samples.

Existing information about experimental interspecific hybridization is scarce.


According to Mukherjee et al. (1968), successful crosses between M. odorata and
M. zeylanica were made in India.

Potential of wild species

There is little doubt that wild mangoes are potentially valuable in breeding
programmes. Some species have important horticultural implications as they
demonstrate many desirable characteristics (Bompard, 1993). Fairchild (1948)
Taxonomy and Systematics 37

noted that crosses between the common mango and related five-stamen spe-cies of
the section Euantherae might produce hybrids with better pollinating quality.
Mangifera pentandra, which is grown in peninsular Malaysia and Sabah, is a
prolific bearer, due to its high proportion of hermaphrodite to male flowers.

Stress resistance
In the Malesian rainforests, wild mangoes thrive well under an ever-humid climate,
without a prolonged dry season, i.e. is in areas with an annual rain-fall > 4000 mm
and no monthly mean < 100 mm and where the common mango cannot be grown
satisfactorily. Species, occurring in subtropical areas, including primitive races of
the common mango, or in high altitude tropical forests, should be evaluated for
cold tolerance, opening up the possibilities for mango production in subtropical and
Mediterranean areas. Mangifera lau-rina and other species related to the common
mango that grow in the rainfor-est (e.g. M. minor in New Guinea) are apparently
immune to anthracnose. Sharma and Choudhury (1976) also observed that trees of
an unknown wild race found in the Tripura State (north-eastern India) were free
from mango malformation.

Potential new fruits


Extensive, yet largely unrecorded variability also exists among the non-indica
species under cultivation. Sadly, this gene pool is barely represented in exist-ing
collections, and is rapidly vanishing. An increasing number of horticul-turists are
demonstrating a keen interest in the wild relatives of the mango. It is hoped that
local peoples who have contributed to the recognition and maintenance of these
species can benefit from future innovative mango breeding.

Since early times, local peoples have planted seeds collected from trees that
were observed to produce better quality fruits in the forests around their
settlements. In areas now completely devoid of lowland primary forest, espe-cially
in Sumatra and Borneo, the only wild relatives still found are those which have
been integrated into indigenous agroforests which represent gene banks for an
amazing diversity of fruit trees. A tenuous but constant selection pressure over
many centuries has resulted in improved selections of several species. Today, some
of these selections hold economic importance for their intrinsic characteristics. In
Malesia, forms of M. odorata and M. foetida with sweeter and less fibrous flesh
have been identified. The ‘wani’, a form of M. caesia from Bali and Borneo, has
green-skinned fruit with milky white soft flesh and a sweet taste quite different
from the fruit of common forms of M. caesia. In addition, there are many
interesting selections of M. casturi, M. grif-fithii and M. torquenda.

Further improvement of these wild mangoes is especially desirable owing to


their local economic importance in the wet tropical regions. Use of vegetative
propagation methods must be encouraged. With proper selection, there is every
reason to believe that other Mangifera species can become valu-able commercial
fruits.
38 J.M. Bompard

Acknowledgement
Thanks are due to Dr Dawn Frame who assisted in correcting the text.

Note
1
Surveys were carried out in Kalimantan in cooperation with the Indonesian Institute
of Science (LIPI) and the Indonesian Commission on Germplasm, and in Malaysia
with the Forest Research Institute of Malaysia (FRIM).

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Mangifera odorata (Anacardiaceae) verified by amplified fragment
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Taxonomy and Systematics 41

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3 Important Mango Cultivars
and their Descriptors

R.J. Knight, Jr,¹ R.J. Campbell² and I. Maguire¹


1
University of Florida, Florida, USA
²Fairchild Tropical Botanic Garden, Florida, USA

3.1 Introduction 43
3.2 Criteria for Cultivar Description 44
3.3 Mango Cultivars 45
‘Alfa’ (Brazil) 45
‘Alphonso’ (India) 45
‘Amelie’ (West Africa) 46
‘Arumanis’ (Indonesia) 46
‘Ataulfo’ (Mexico) 46
‘B74’ (‘Calypso’™) (Australia) 47
‘Banganpalli’ (India) 47
‘Beta’ (Brazil) 47
‘Bombay Green’ (India) 47
‘Cambodiana’ (Vietnam) 48
‘Carabao’ (Philippines) 48
‘Chausa’ (India) 48
‘Cogshall’ (Florida, USA) 49
‘Coração de Boi’ (Brazil) 49
‘Dasheheri’ (India) 49
‘Espada’ (Brazil) 49
‘Ewais’ (Egypt) 50
‘Excellent Succari’ (Egypt) 50
‘Extrema’ (Brazil) 50
‘Fajri’ (India) 50
‘Fernandin’ (India) 51
‘Genovea’ (Egypt) 51
‘Glenn’ (Florida, USA) 51
‘Golek’ (Indonesia) 51
‘Haden’ (Florida, USA) 52
‘Himsagar’ (India) 52
‘Hindi Besennara’ (Egypt) 52
‘Hindi Khassa’ (Egypt) 52
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
42 (ed. R.E. Litz)
Mango Cultivars and Descriptors 43

‘Irwin’ (Florida, USA) 53


‘Julie’ (West Indies) 53
‘Keitt’ (Florida, USA) 53
‘Kensington’ (Australia) 54
‘Kent’ (Florida, USA) 54
‘Khanefy’ (Egypt) 54
‘Kyo Savoy’ (Thailand) 55
‘Langra’ (India) 55
‘Mabrouka’ (Egypt) 55
‘Madame Francis’ (Haiti) 55
‘Mallika’ (India) 56
‘Manila’ (Mexico) 56
‘Manzanillo’ (Mexico) 56
‘Mesk’ (Egypt) 56
‘Mulgoa’ (India to Florida, USA) 57
‘Nabeel’ (Egypt) 57
‘Nam Doc Mai’ (Thailand) 57
‘Neelum’ (India) 58
‘Nuwun Chan’ (Thailand) 58
‘Okrung’ (Thailand) 58
‘Osteen’ (Florida, USA) 59
‘Pairi’ (India) 59
‘Palmer’ (Florida, USA) 59
‘Rosa’ (Brazil) 59
‘Sensation’ (Florida, USA) 60
‘Suvarnarekha’ (India) 60
‘Tahar’ (Israel) 60
‘Taimour’ (Egypt) 61
‘Tommy Atkins’ (Florida, USA) 61
‘Totapuri’ (India) 61
‘Turpentine’ (West Indies) 62
‘Vallenato’ (Colombia) 62
‘Van Dyke’ (Florida, USA) 62
‘White Succari’ (Egypt) 62
‘Zebda’ (Egypt) 63
3.4 Conclusion 63

3.1 Introduction
The mango (Mangifera indica L.) has traditionally been grown in an area that
extends southwards and eastwards from India through Myanmar and Vietnam to
Indonesia. It probably is not indigenous to the Philippines, where it has long been
cultivated (Valmayor, 1962), but Mangifera species are endemic there (Bon-dad,
1982). This crop is best adapted to a warm tropical monsoon climate, with a
pronounced dry season followed by rains. Fruit of the best quality is usually
produced in such areas, but specific races are known to fruit in humid regions. For
example, some Mangifera species bear dependably on the island of Borneo, where
most standard cultivars do not mature normal crops of fruit. Numerous Mangifera
species closely related to the common mango are indigenous to
44 R.J. Knight et al.

Borneo and nearby parts of Malaysia and Indonesia, and this region is the prob-
able centre of diversity for this genus (see Bompard, Chapter 2 this volume).
Most crops long cultivated over an extended time and area show consider-able
diversity, reflecting different selection criteria in different regions of cul-ture as
well as genetic responses to varied environmental influences. Certainly this is true
of the mango. Indian cultivars differ markedly from those grown in South-east
Asian countries and in Egypt. An additional factor that has promoted genetic
diversity in the group recently has been the widespread introduction of this crop
into new areas of cultivation, many in the western hemisphere, over the last 500
years. In this manner, genetically diverse germ-plasm has been brought from
widely dispersed areas of the original range of the species and grown in mixed
plantings where, through the cross-pollination natural to the species, new genetic
combinations have been made and selected under many varying conditions of
microclimate. In Florida, since the late 18th century, enough such importations and
genetic recombinations have occurred to qualify the southern part of this state as a
secondary centre of diversity for the crop. A new group of mango clones
designated the Florida cultivars has been exported to Brazil, Israel, Australia and
other places where the process of increasing diversity under new and varying
cultural and environmental con-ditions continues (Knight and Schnell, 1994;
Schnell et al., 2006).
Some Florida cultivars, most notably ‘Haden’, have been important in aid-ing
the establishment of a modern mango industry in other parts of the world (Knight
and Schnell, 1994), and the phenomenon first observed in Florida is now occurring
elsewhere; we are presented with the prospect of the importation of cultivars of
outstanding merit from their countries of origin to be grown, first experimentally
and then commercially, in new regions. For this reason it is important to become
familiar with the characteristics of a group of cultivars that currently are known in
the commerce and/or horticulture of one or more coun-tries, and that may have
potential for expanded culture or use in breeding.

3.2 Criteria for Cultivar Description


In the recent past, efforts to assemble lists of mango descriptors produced two
publications that cover the subject (Mukherjee, 1985; IBPGR (now Inter-national
Plant Genetic Resources Institute, IPGRI), 1989) and provide people who manage
collections with morphological criteria to identify cultivars. The Descriptor List
used by IPGRI documents passport data (identifying the acces-sion and
information recorded by collectors), characterization (recording char-acters
considered to be highly heritable which can easily be seen in the field and are
expressed in all environments) and preliminary evaluation, which records a limited
number of additional traits thought desirable by a consensus of users of the crop.
Plant data are important in preliminary evaluation, and include for the tree, habit
and height of the mature tree; for the leaf, shape, length and width, and colour of
the young leaf; and for the inflorescence, position, shape, density of flowers,
length, colour, hairiness, presence or absence of leafy bracts, and percentage of
flowers in an average inflorescence. (Some research indicates
Mango Cultivars and Descriptors 45

that both leafy bracts and number of perfect flowers are influenced by local
conditions and vary in their expression with differing environments.)
Additional plant data used in initial evaluation include those for the flower,
diameter in millimetres, type (pentamerous, tetramerous or both), nature of disc
(swollen, broader or narrower than ovary, reduced or absent) and number of
stamens; fruit, length, width and thickness, weight, shape, skin colour (which may
be compared with reference cultivars), skin thickness, skin texture, ratio of pulp to
skin and stone, texture of pulp, adherence of skin to pulp, fibre in pulp and its
quantity and length, and stem insertion; and stone, length, weight, veins and pattern
of venation, presence or absence of fibres and their length.
Additional plant data for leaves, inflorescence and fruit have been col-lected
and some of these, notably season (maturity period), productivity, eat-ing quality
and attractiveness are quite important. Unfortunately, from the viewpoint of those
who expect to apply these criteria outside the Indian sub-continent, reference
cultivars are for the most part Indian and many are not readily available outside
India. Other important characters that have been evaluated or proposed for
evaluation include susceptibility to stress (drought, wind, flooding), susceptibility
to specific diseases and pests, molecular markers, cytological characters and
identified genes. Because of the extreme comprehensiveness of this list and the
limited availability of many of the proposed descriptor evaluations at this time, we
have tried to utilize such information as is available to make the comparison,
identification and evalu-ation of specific well-known cultivars a practical
possibility.

3.3 Mango Cultivars


A list of mango cultivars that are of interest in areas other than their places of
origin, with descriptions intended to help differentiate them, follows (see Plates 4–
40). Spelling and name variants in some cases represent efforts to transliterate from
other orthographies to the Roman alphabet, and in others reflect regional
differences in usage.

‘Alfa’ (Brazil)

A monoembryonic cultivar developed by EMBRAPA Cerrado, Brazil, from


crossing ‘Mallika’ × ‘Van Dyke’. The tree is semi-dwarf in habit and high-
yielding, resistant to Oidium mangiferae and malformation, and moderately
resistant to anthracnose (Colletotrichum gloeosporioides); the fruit is large (435 g),
pink-red, firm, medium fibrous and of good quality (16% total soluble solids
(TSS), 0.23% acidity) (Pinto et al., 2004).

‘Alphonso’ (India)

Also known as ‘Appus’, ‘Badami’, ‘Gundu’, Haphus’, ‘Kagdi’, ‘Khader’ and


‘Khader Pasand’. The tree is moderately large, with broadly rounded, dense
46 R.J. Knight et al.

canopy; the fruit (Plate 4) is yellow, ovate-oblique, averaging 6 cm long by 5 cm


broad, weighing 225–325 g (mean 226 g); the skin is thin; the flesh is firm to soft,
low in fibre, yellow, sweet with characteristic aroma and with a very pleasant taste
preferred by many who know this cultivar, bringing pre-mium prices on Indian and
international markets. Seed is monoembryonic in a large, woody stone; the quality
is excellent; ripening fruit in late to midsea-son. Bearing is irregular, medium to
heavy in India, but light and irregular in Florida (Prasad, 1977; R.J. Knight, Jr,
personal communication, 1995).

‘Amelie’ (West Africa)

Also known as ‘Gouverneur’ in the Caribbean. The tree is tall with a rounded,
dense canopy; the fruit is green to orange-yellow with the advance of the season,
rounded, 10–15 cm long by approximately 10 cm broad by approximately 7.8 cm
thick and weighing 300–600 g (average 360 g). The skin is thick and separated
with difficulty; the flesh is soft, juicy, melting, without fibre, a deep orange colour,
sweet and perfumed, free from turpentine, and provides the best of mango tastes.
Seed is monoembryonic in a medium-sized, elongate, narrow stone that adheres to
the flesh, having a few short, pliable fibres that are not objectionable; the quality is
excellent; the season early. The fruit closely resem-bles that of ‘Julie’. ‘Amelie’ is
exported to France, along with ‘Kent’, from Burkina Faso, Ivory Coast and Mali.
‘Amelie’ is increasing in popularity on the French market, chiefly in Paris and the
surrounding area. It brings lower prices than cultivars with blushed fruit because
the consumer is not always aware when it is ripe (Naville, 1985, 1986; R. LePrette,
personal communication, 1996).

‘Arumanis’ (Indonesia)

Also referred to as ‘Harumanis’. The tree is vigorous and tall with a slightly open
canopy. The fruit (Plate 5) is greenish yellow with large, light-yellow dots,
elongate oblong with rounded base, 11–14 cm long by 6.6–7.5 cm broad by 4.75–
6.5 cm thick, weighing 200–350 g. The skin is thick, tough and easily separated,
the flesh firm and juicy with little fibre, lemon yellow, sweet, slightly insipid with
a strong aroma, of poor to fair quality. Seed is polyem-bryonic in a thick, woody
stone; this cultivar ripens midseason and bears regularly. Relatively easy to
propagate by graftage, scionwood survives well; widely planted in humid parts of
the world where many better-quality culti-vars fail to fruit (R.J. Knight, Jr, personal
communication, 1995).

‘Ataulfo’ (Mexico)

A polyembryonic cultivar sold in North American markets under the name


‘Ataulfo’ and as ‘Champagne’™. Originated in Tapachula, Chiapas, Mexico
reportedly from seed brought from Costa Rica in about 1930. The tree is
Mango Cultivars and Descriptors 47

vigorous and upright, a mid-range producer with production averages of 10–20 t/ha
possible. The tree is not highly adaptive to different climatic/ edaphic conditions. It
is moderately resistant to anthracnose disease. The fruit (Plate 6) is small (200–300
g), elongate, of good quality, sweet with slight acidity, yellow, firm, standing
shipping stress well, and ripens from early to midseason (Campbell et al., 2002;
Magallanes-Cedeño, 2004).

‘B74’ (‘Calypso’™) (Australia)

A monoembryonic cultivar that originated from the controlled cross of ‘Sensa-tion’


× ‘Kensington’. The tree is upright, with low to moderate vigour and is highly
productive, with good tolerance of flower and fruit diseases; the fruit (Plate 7) is
moderately large (457.4 ± 38.1 g), ovate (10.12 ± 0.27 cm long by 9.13 ± 0.28 cm
wide), fibre-free and firm, bright yellow overlaid with red blush, with extended
shelf life and potential for shipment to overseas markets; ripens late in the season;
patented (Whiley, 2001; Whiley and Hofman, 2006).

‘Banganpalli’ (India)

Also called ‘Beneshan’ and ‘Chappatai’. The tree is medium sized with a rounded
canopy; the fruit is primrose-yellow, ovate-oblique, large and the skin smooth, thin
and shiny, flesh firm to meaty with juice moderately abun-dant, without fibre,
maize-yellow, with pleasant aroma and sweet taste. Seed is monoembryonic, in an
oblong stone covered with sparse fibres; quality good; ripens midseason and bears
heavily (Singh, 1960).

‘Beta’ (Brazil)

A cultivar developed by EMBRAPA Cerrado, Brazil, from crossing ‘Amra-pali’ ×


‘Winters’ (M20222 United States Department of Agriculture (USDA)). The tree is
moderately vigorous and free of malformation, high-yielding but irregular,
moderately resistant to anthracnose and Oidium; the fruit is small (310 g), yellow,
firm with low fibre, of excellent quality (24.8% TSS, 0.16% acidity) (Pinto et al.,
2004).

‘Bombay Green’ (India)

Also called ‘Bhojpuri’, ‘Bombai’, ‘Hiralal Bombai’, ‘Kali Bombai’, ‘Laile Alipur’,
‘Malda’, ‘Sarauli’ and ‘Sheeri-Dhan’. The tree is tall and erect; the fruit (Plate
8) is apple green with ochre blush at the base and on some exposed parts, dots
abundant, with brown specks in the middle, ovate with beak almost missing,
medium sized, with tough, thick, non-adhering smooth skin; the flesh is cadmium-
orange, firm and juicy with scanty fibre just under the skin,
48 R.J. Knight et al.

very sweet with pleasant aroma, of very good quality; seed is monoembryonic in a
full, thick, medium-sized stone. This cultivar ripens early in the season and is a
medium bearer. ‘Bombay Yellow’ is said to be practically identical to this cultivar
but for a slight difference in fruit colour. The present ‘Bombay Green’ is said to be
a degenerate form of the original one (Singh, 1960). In Jamaica it is sometimes
called ‘Peter’, which suggests a confusion with ‘Pairi’, but the Jamaican ‘Peter’ is
without the bright red blush normal to ‘Pairi’.

‘Cambodiana’ (Vietnam)

Also known as ‘Xoai Voi’. The tree is moderately vigorous, with a dense, rounded
canopy; the fruit (Plate 9) is greenish yellow, unblushed with a few small white
dots, oblong to ovate, 9–11.5 cm long by 6.5–7.5 cm broad by 5–6 cm thick,
weighing 220–340 g; the skin is thin, tender and adherent; the flesh contains little
fibre, is tender and melting, lemon yellow, sweet and mildly subacid with a
pleasant aroma; the seed is polyembryonic in a thick, woody stone. Ripens early in
the season. Brought to Florida in 1902, where it gave rise to the ‘Saigon’ landrace
(Campbell, 1992).

‘Carabao’ (Philippines)

The tree is vigorous, forming a large and dense canopy; the fruit (Plate 10) is
greenish to bright yellow, brushed with a few small green dots, long and slender,
with base rounded to slightly flattened, 11–13 cm long by 6.5–7 cm broad by 6–6.5
cm thick, weighing 270–440 g; the skin is thick, medium tough and easily
separated; the flesh is without fibre, tender and melting, lemon yellow, spicy and
sweet with a mild aroma, of good to excellent quality; seed is polyembryonic in a
thin, papery stone. Ripens early in the season (Camp-bell, 1992). This is a heavy
bearer that may alternate; however, it can be induced to fruit by potassium nitrate
treatment in the tropics (Bondad and Linsangan, 1979). It is highly resistant to
bacterial black spot (Xanthomonas campestris pv. mangiferaeindicae) in
Queensland (Mayers et al., 1988). It was introduced to Florida in 1909. ‘Carabao’
is important in commerce between the Philippines and Japan and is increasingly
imported into the USA.

‘Chausa’ (India)

Also called ‘Samar Bahisht Chausa’ and ’Khajari’. The tree is tall and spread-ing;
the fruit is canary yellow to raw sienna when fully ripe, with numerous obscure
medium-sized dots with minute specks inside them, oblong with prominent beak,
obtuse to rounded, medium sized; the skin is thin and some-what adhering, pulp
raw sienna, soft and juicy with scanty fine, long fibres near the skin; the fruit is
very sweet with a luscious, delightful aroma, of excellent quality; seed is
monoembryonic in a thick, medium-sized oblong
Mango Cultivars and Descriptors 49

stone with fine, short fibres all over the surface and a tuft of long fibres on the
ventral edge. Ripens late in the season and is a light bearer (Singh, 1960).

‘Cogshall’ (Florida, USA)

A monoembryonic cultivar that originated on Pine Island in Lee County. The tree
is relatively small, forming a rounded canopy, moderately susceptible to
anthracnose and consistently productive; the fruit is medium to large, aver-aging
about 350 g, yellow with a bright crimson blush, oblong (11–14 cm long by 7.5–
8.5 cm broad by 6.2–8 cm thick) of excellent quality, rich and sweet in taste, with
tender skin and soft flesh. Ripens early to midseason over about 4 weeks, a season
longer than some cultivars. It is recommended for the home garden, not
commercial planting, in Florida but is now grown commercially on Mauritius and
marketed in France. Seedling of ‘Haden’ (Campbell and Campbell, 1995; Schnell
et al., 2006).

‘Coração de Boi’ (Brazil)

The tree is vigorous, precocious and productive; the fruit is greenish yellow and
intense red on the side exposed to the sun, cordiform, medium sized, pulp yel-low
and fibrous. The seed is polyembryonic. There are two seasons in São Paulo,
January–February and September–December. This is one of the best-known
commercial cultivars in São Paulo state (Sampaio, 1980; A.C. Pinto, personal
communication, 1996; L.C. Donadio, personal communication, 1996).

‘Dasheheri’ (India)

Also known as ‘Dasheri’ and ‘Aman Dusehri’. The tree is of medium height and
moderate vigour, spreading, with a rounded, medium-dense canopy; the fruit is
primrose to canary yellow with abundant light-yellow dots, oblong to oblong-
oblique with base rounded to obliquely rounded, medium sized, skin smooth,
medium thick, tough and non-adhering; the flesh is yellow, firm, with almost no
fibre, scanty juice and a delightful aroma, very sweet taste, of excellent quality;
seed is monoembryonic in a thick, medium-sized stone. Rip-ens midseason and is
heavy bearing; fruit keeps well (Singh, 1960).

‘Espada’ (Brazil)

The tree is tall and develops rapidly, with a dense canopy, very productive; the fruit
is intense green or greenish yellow, oblong-elongate with a concave base, medium
sized, with smooth, thick skin; the flesh has much fibre, is egg-yellow, with a
strong aroma of turpentine. The quality is considered good for fresh consumption.
The polyembryonic seed is in an oblong stone, covered
50 R.J. Knight et al.

with soft fibres and many nerves. There are two seasons per year in São Paulo,
January–February and November–December (Sampaio, 1980; A.C. Pinto, personal
communication, 1996).

‘Ewais’ (Egypt)

A polyembryonic cultivar of major commercial importance. The tree is vigor-ous,


the fruit small (275 g), yellow with no blush, with small, light-brown slightly corky
dots, ovate-oblong in shape (11.7 cm long by 7.2 cm wide by 6.3 cm thick), with
adherent skin of intermediate thickness, relatively free of disease; flesh orange,
juicy but susceptible to jelly seed, with no objectionable fibre, sweet and agreeable
in taste, of very good quality. The stone is large (38.5 g). Fruit ripens midseason
(Knight and Sanford, 1998). In warm subtrop-ics this cultivar has shown a
tendency for flowering in the warm season, with fruit ripening during the cool
winter. It has good anthracnose tolerance.

‘Excellent Succari’ (Egypt)

A polyembryonic mango of minor commercial importance. The tree is vigor-ous,


ripening fruit in late midseason. The fruit is small (280 g), green with a yellow
overlay and small, yellow smooth dots, ovate-oblong in shape (11 cm long by 7 cm
wide by 6.4 cm thick), with non-adherent skin of intermediate thickness quite free
of surface disease; the flesh is orange, melting (without jelly seed) and juicy with
no objectionable fibres, a delightfully sweet taste and excellent quality; stone large
(36.6 g) (Knight and Sanford, 1998). It has moderate to good anthracnose tolerance
in the warm subtropics.

‘Extrema’ (Brazil)

The tree is upright, vigorous and productive. The fruit is yellow with green-ish
areas, ovate-reniform, weighing 350–400 g, with smooth and thin skin, and yellow,
watery flesh with almost no fibres with a moderately resinous, agreeable taste. The
quality is considered good for fresh consumption and processing. The
polyembryonic seed is in a large, fibrous stone. Ripens early in the season
(Sampaio, 1980; A.C. Pinto, personal communication, 1996).

‘Fajri’ (India)

Also spelled ‘Fazli’. The tree is of medium size and moderately vigorous, with
rounded, open canopy. The fruit is light chrome yellow with small, dark-coloured
fairly sparse dots, obliquely oval with base slightly rounded and beak distinct to
slightly prominent, large (averaging 14.3 cm long by 9.8 cm broad, weighing 500 g
on average) with a medium-thick skin that is
Mango Cultivars and Descriptors 51

smooth with some inclination to be warty, and firm to soft, fibreless flesh of a light
cadmium yellow with a pleasant aroma and a sweet taste, having juice that may be
scanty to moderately abundant, of good to very good quality. The seed is
monoembryonic in a large, oblong stone that is covered with a sparse, short and
soft fibre. Ripens midseason to late (Gangolly et al., 1957; N. Balasundaram, India,
personal communication, 1990).

‘Fernandin’ (India)

The tree is moderately vigorous with a dense, rounded canopy; the fruit is bright
yellow with an attractive bright-red blush, ovate-oblique, averaging 12.2 cm long
by 8.5 cm broad, weighing 450 g; the skin is rough and warty, thick and adherent,
flesh bright yellow, moderately to abundantly juicy, thick, with no objectionable
fibre, with delightful to piquant aroma and sweet to very sweet, delicious taste, of
superior quality; seed is monoembryonic; season late (Gangolly et al., 1957; Singh,
1960).

‘Genovea’ (Egypt)

A polyembryonic cultivar of minor commercial importance. The fruit is small


(234.5 g), green with a yellow overlay and medium-sized smooth yellow dots,
ovate-oblong in shape (11 cm long by 6 cm wide by 5.6 cm thick), a thin adherent
skin relatively free of surface disease; flesh orange, firm (no jelly seed) and juicy
with no objectionable fibres, a sweet agreeable taste of ac-ceptable quality; stone
large (53 g) (Knight and Sanford, 1998).

‘Glenn’ (Florida, USA)

The tree is moderately vigorous, small to medium with dense, rounded can-opy of
compact growth; the fruit (Plate 11) is bright yellow with orange-red blush, with
numerous small yellow and white dots, oval to oblong with a rounded base, 9.5–
12.5 cm long by 7.5–8.5 cm broad by 7–8 cm thick, weigh-ing 400–620 g; the skin
is thin, tough and easily separated, flesh soft and juicy, with little fibre, deep
yellow, rich and spicy with a strong, pleasant aroma, of excellent quality; seed is
monoembryonic in a thick, woody stone. Ripens early in the season and is a regular
bearer. This is a seedling of ‘Haden’ (Campbell, 1992; Schnell et al., 2006).

‘Golek’ (Indonesia)

The tree is moderately vigorous with an upright, open canopy; the fruit (Plate 12) is
greenish yellow with an orange overlay and prominent white dots, oblong with
rounded base, 9.5–12.5 cm long by 6–8 cm broad by 5.5– 6.5 cm thick, weighing
200–365 g; the skin is thin, tough and easily separated;
52 R.J. Knight et al.

the flesh is soft and juicy with abundant fibre (not objectionable), deep yel-low,
sweet, insipid with a mild aroma, of poor to fair quality; the seed is polyembryonic
in large, woody stone with abundant fine fibre. Ripens mid-season (R.J. Knight, Jr,
personal communication, 1995).

‘Haden’ (Florida, USA)

The tree is vigorous, with a large, spreading canopy; the fruit (Plate 13) is bright
yellow with a deep crimson or red blush and numerous large yellow dots, oval with
a rounded base, 10.5–14 cm long by 9–10.5 cm broad by 8.5– 9.5 cm thick,
weighing 510–680 g; the skin is thick, tough and adherent; the flesh is firm and
juicy with abundant fibre, deep yellow, rich and sweet with a pleasant aroma, of
good to excellent quality; the seed is monoembyonic in a medium-thick woody
stone. Ripens early to midseason and bearing is sometimes irregular. This is a
seedling of ‘Mulgoba’ × ‘Turpentine’ and is the first of the Florida mango
cultivars, introduced in 1910 and since grown in many other countries. It is the
seed parent of numerous other cultivars (Campbell, 1992; Knight and Schnell,
1994; Schnell et al., 2006).

‘Himsagar’ (India)

The tree is vigorous, tall, with a dense, spreading canopy; the fruit (Plate 14) is
greenish yellow to bright yellow with no blush, with light-yellow dots, ovate with a
flattened base, 12–15 cm long by 8.5–9.5 cm broad by 7.5–8.5 cm thick, weighing
465–585 g; the skin is thin, tough and easily separated; the flesh is firm and juicy
with no fibre, orange, rich and sweet with a mild aroma, of good to excellent
quality; the seed is monoembryonic in a thick, woody stone. This is a late
midseason cultivar that bears well (R.J. Knight, Jr, per-sonal communication,
1995).

‘Hindi Besennara’ (Egypt)

A polyembronic cultivar of major commercial importance. The tree is of me-dium


vigour, ripening fruit early to midseason. The fruit (Plate 15) is of small to medium
size (319 g), green with orange overlay, with small white corky dots, oblong-
cylindrical in shape (15.4 cm long by 6.7 cm wide by 6.5 cm thick) with thick, non-
adherent skin relatively free of surface disease; the flesh is orange, yielding and
juicy with no objectionable fibres, pleasantly sweet in taste, of very good quality;
the stone is large (47.2 g) (Knight and Sanford, 1998).

‘Hindi Khassa’ (Egypt)

A polyembryonic cultivar of major commercial importance. The tree is vig-orous,


ripening fruit in late midseason. The fruit is of medium size (461 g),
Mango Cultivars and Descriptors 53

yellow with no blush, with intermediate-sized smooth, light-yellow dots, oblong-


cylindrical in shape (16 cm long by 6.6 cm wide by 6.9 cm thick), with thick,
adherent skin relatively free of surface disease; flesh is orange, firm and juicy with
no objectionable fibres, of mediocre taste and a quality not suitable for export;
stone is large (55 g) (Knight and Sanford, 1998).

‘Irwin’ (Florida, USA)

The tree is small to medium, moderately vigorous, with open canopy. The fruit
(Plate 16) is bright yellow with a crimson or bright red blush, numerous large white
dots, ovate with rounded base, 11.5–13 cm long by 8–9 cm broad by 6.5– 7.5 cm
thick, weighing 340–450 g; the skin is medium-thick, tender and adherent; the flesh
is soft, tender, melting and juicy without fibre, lemon yellow, sweet and mild with
a pleasant aroma, of good quality; the seed is monoembryonic in a thin, papery
stone. The stone may be seedless following cool weather at flower-ing time. This is
an early, regular and heavy bearer. The fruit is usually soft with a short postharvest
life, but it is often exported from tropical America to Europe. It is a seedling of
‘Lippens’ × ‘Haden’ (Campbell, 1992; Schnell et al., 2006).

‘Julie’ (West Indies)

Also called ‘St Julienne’. The tree is compact (dwarf), with a dense canopy; the
fruit (Plate 17) is greenish yellow with a light pink to maroon blush and numerous
small white dots, rounded with flattened apex, pronouncedly compressed laterally,
7–9.5 cm long by 4–7.5 cm broad by 2–5.5 cm thick, weighing 200–325 g with a
thin, tender skin and soft, melting, juicy, orange flesh with scanty fibre, of a rich,
spicy flavour with a strong, pleasant aroma, of good quality; seed is
monoembryonic in a thin, papery stone. This cultivar ripens midseason and is a
regular producer of small crops. The fruit is often severely infected with
anthracnose disease, but its unique taste is preferred by many West Indians, and it
is exported to the London market (C.W. Campbell, personal communication,
1996).

‘Keitt’ (Florida, USA)

The tree is medium sized, moderately vigorous, upright with open canopy; the fruit
(Plate 18) is greenish yellow, with a pink or red blush, numerous small white or
yellow dots, oval, with rounded base, 13–15 cm long by 9–11 cm broad by 8.5–10
cm thick, weighing 510–2000 g; the skin is thick, tough and adherent; the flesh is
firm and juicy, with little fibre, lemon yellow, sweet and mild with a pleasant
aroma, of good to excellent quality; the seed is monoembryonic in a thick and
woody stone. This cultivar ripens late in the season. It is a seedling of ‘Brooks’.
After ‘Tommy Atkins’ it is the most com-mercially important cultivar in the export
mango industry of the western
54 R.J. Knight et al.

hemisphere. It is resistant to anthracnose disease, packing and shipping stress and


is heavily productive (Campbell, 1992; Schnell et al., 2006). It is highly sus-
ceptible to bacterial black spot in Queensland (Mayers et al., 1988).

‘Kensington’ (Australia)

Also known as ‘Kensington Pride’ and ‘Bowen’. ‘Kensington’ has a large, vig-
orous tree with spreading canopy; the fruit (Plate 19) is yellow with an orange-red
blush on the shoulder, round ovate with a flattened base and a slight beak, 10.5–13
cm long by 8.5–9.6 cm broad by 7.5–8.5 cm thick, weighing 350–750 g; the skin is
thick, tender and adherent; the flesh is soft and juicy, with moderate to little fibre,
sweet with a characteristic flavour that makes it the most popu-lar cultivar in
Australian markets, of excellent quality; seed is polyembryonic in a moderately
thick, woody stone. This cultivar ripens midseason and it bears well. It is unusually
susceptible locally, in Florida, to damage by red-banded thrips (Selenothrips
rubricinctus (Giard.)), and may be killed by this pest without adequate
countermeasures (R.J. Campbell, personal communication, 1994; R.J. Knight, Jr,
personal communication, 1995). It is moderately suscep-tible to anthracnose and
bacterial spot (Mayers et al., 1988).

‘Kent’ (Florida, USA)

The tree is large and vigorous with a dense, upright canopy; the fruit (Plate
20) is greenish yellow with a red or crimson blush, numerous small yellow dots,
oval, with rounded base, 11–13 cm long by 9.5–11 cm broad by 9.9.5 cm thick,
weighing 600–750 g; the skin is thick, tough and adherent; the flesh is firm, tender,
melting and juicy with little fibre, deep yellow to orange-yellow, sweet with a rich
flavour and pleasant aroma, of excellent quality; the seed is monoembryonic in a
thick, woody stone. Fruit ripens late midseason to late and bearing may be
alternate. It is a seedling of ‘Haden’ × ‘Brooks’, which is a seedling of ‘Totapuri’
(‘Sandersha’) (Schnell et al., 2006). ‘Kent’ is not commonly commercial in Florida
because it is prone to storage disease, but is a successful commercial cultivar in
drier parts of Mexico, Central and South America and West Africa (Campbell,
1992). It is highly susceptible to bacterial black spot in Queensland (Mayers et al.,
1988).

‘Khanefy’ (Egypt)

A cultivar of minor commercial importance. The fruit is large (475 g), green with a
yellow overlay and large, brown, smooth dots, ovate in shape (10.7 cm long by 8.3
cm wide by 8.6 cm thick), with an adherent skin quite free of sur-face disease; the
flesh is yellow, often with jelly seed, juicy, with no objection-able fibres and a
bland flavour unacceptable to many Western palates. The stone is moderately large
(53 g) (Knight and Sanford, 1998).
Mango Cultivars and Descriptors 55

‘Kyo Savoy’ (Thailand)

The tree is large, vigorous, with an open canopy made up of long branches; the
fruit is green when harvested (before the ripening process begins) turn-ing to
greenish yellow, oblong, 11.5–12.5 cm long by 5.5–6.5 cm broad by 5–6 cm thick,
weighing 230–340 g; the skin is thin, tender and adherent; the flesh is medium
firm, tender and not very juicy with no fibre, pale yellow, very sweet with an
insipid taste and a mild, pleasant aroma, of fair to good quality; the seed is highly
polyembryonic in a medium-thin stone. This is a regular producer (C.W. Campbell,
personal communication, 1995). The fruit is often consumed green.

‘Langra’ (India)

Also called ‘Darbhanga’, ‘David Ford’, ‘Hadialaziz’, ‘Hajipur Langra’, ‘Har-doi


Langra’, ‘Lan Garhi’, ‘Langra Faquirwala’, ‘Sylhet’ and ‘Tikari’. The tree is
moderately vigorous, forming a dense canopy; the fruit is greenish yellow with
medium to big dark-green dots, ovalish to oblong, 8–10.5 cm long by 6.5–7.5 cm
broad by 6–7 cm thick, weighing 235–375 g; the skin is medium smooth, thick; the
flesh is firm to soft, fibreless, lemon yellow, very sweet with a strong, pleasant
aroma, juice moderately abundant; seed is monoem-bryonic in a medium-sized,
flattened stone covered with dense, short and soft fibre; quality is very good. Fruit
ripens early to midseason (Gangolly et al., 1957; R.J. Knight, Jr, personal
communication, 1995).

‘Mabrouka’ (Egypt)

A major commercial cultivar considered to have been originally introduced from


India. The tree is moderately vigorous; the fruit (Plate 21) is large (481 g), yellow
with an orange to red blush and small, light-yellow, smooth dots; ovate-oblong in
shape (13.7 cm long by 8.9 cm wide by 8.2 cm thick) with a thick, non-adherent
skin relatively free of surface disease; the flesh is yellow, firm and juicy with no
objectionable fibre, moderately agreeable in taste, of acceptable quality. The
monoembryonic seed is in a moderately large (51 g) stone. Fruit ripens late
midseason, ships well and has been marketed in Poland (Singh, 1960; Knight and
Sanford, 1998).

‘Madame Francis’ (Haiti)

The tree is moderately vigorous, medium sized, forming an open canopy; the fruit
(Plate 22) is greenish to bright yellow, with no blush and a few large rus-set dots,
oblong, sigmoid with rounded base, 15–17 cm long by 8.5–11 cm broad by 5.5–7.5
cm thick, weighing 370–520 g; the skin is thin, tender and adherent; the flesh is
soft and juicy with medium fibre, orange, rich spicy and sweet with a pleasant
aroma, of fair to good quality; seed is polyembryonic
56 R.J. Knight et al.

in a thin, papery stone. This cultivar ripens early to midseason and bears well.
Shipped to North American markets from Haiti nearly 10 months of the year (R.J.
Campbell, personal communication, 2007).

‘Mallika’ (India)

The tree is a moderately vigorous dwarf with a dense canopy; the fruit (Plate
23) is bright yellow with no blush and numerous small, light-yellow dots, oblong
with rounded base, 10–12 cm long by 6.5–7.5 cm broad by 5–5.5 cm thick,
weighing 280–450 g; the skin is thick, tough and easily separating; the flesh is soft,
tender and juicy with little fibre, deep yellow to orange, rich, strongly aromatic and
sweet, of excellent quality; seed is monoembryonic in a medium-thick and woody
stone. This cultivar ripens midseason and is an irregular producer. This cultivar
came from crossing ‘Neelum’ and ‘Dashe-hari’ (Singh et al., 1972; Campbell,
1992).

‘Manila’ (Mexico)

The tree is large, vigorous, with an upright, open canopy; the fruit (Plate 24) is
bright yellow, sometimes with a light-pink blush, a few small reddish dots, long
and slender with rounded base and bluntly pointed apex sometimes with a small
beak, 12.5–14 cm long by 5.5–6 cm broad by 5–5.5 cm thick, weighing 180–260 g;
the skin is thin, medium tough and easily separating; the flesh is medium firm and
juicy, with little to abundant fibre, deep yellow, sweet, rich and spicy in taste with
a pleasant aroma, of good to very good quality; seed is polyembryonic in a
medium-thick and woody stone. This cultivar ripens early midseason and crops
fairly dependably. For a long time ‘Manila’ has been the most popular mango in
Mexico.

‘Manzanillo’ (Mexico)

The tree is large, of medium vigour with an upright canopy; the fruit is yel-lowish
orange with 75% of the surface blushed an intense dark red with numerous dots,
oval with moderately flattened base, averaging 12 cm long by 10 cm broad by 7.5
cm thick, and 660 g in weight; the flesh is low in fibre, slightly subacid and very
palatable, quality high; seed is monoembryonic in a relatively small stone. This
cultivar ripens early in the season but spread over a 60-day harvest period. It bears
heavily without pronounced alterna-tion and the fruit stores and ships well (Núñez-
Elisea, 1984).

‘Mesk’ (Egypt)

A major commercial cultivar. The tree is vigorous, the fruit small to medium sized
(312.5 g), yellow with a red blush, with small, corky yellow dots;
Mango Cultivars and Descriptors 57

ovate-oblong (11.3 cm long by 7.4 cm wide by 6.5 cm thick) with adherent skin
intermediate in thickness and fairly free of surface disease; the flesh is orange,
frequently jelly-seeded, with no objectionable fibres and a sweet, agreeable taste of
very good quality. The polyembryonic seed is in a moder-ately large (52.5 g) stone.
Fruit ripens late in the season (Knight and Sanford, 1998).

‘Mulgoa’ (India to Florida, USA)

Also spelled ‘Mugoba’ and ‘Mulgova’. The tree is large, vigorous with open,
spreading canopy; the fruit (Plate 25) is bright yellow with a pink blush and
numerous large white dots, oval to ovate with flattened base, 8.5–10.5 cm long by
6.5–7.5 cm broad by 5–6 cm thick, weighing 340–450 g; the skin is thick, medium
tough and adherent; the flesh is soft, tender, melting and juicy, with little fibre,
lemon yellow, rich spicy and sweet with strong, pleas-ant aroma, of good to
excellent quality; seed is monoembryonic in a thick, woody stone. This cultivar
ripens midseason to late and is a shy, irregular bearer. Introduced to Florida in 1889
and called ‘Mulgoba’, this is the seed parent of ‘Haden’, first of a series of cultivars
known as the Florida group. A question exists whether the cultivar known in
Florida is identical with the Indian cultivar or is a seedling rootstock that survived
after the scion was killed by cold. In either case its superior quality ensured its
retention and propagation (Campbell, 1992). Literature serves to compound the
nomencla-tural confusion, as illustrated by Gangolly et al. (1957) whose ‘Mulgoa’
fruit, yellow overall and roundish oblique with a deeply depressed stem insertion,
does not resemble the cultivar introduced to Florida. Singh (1960), on the other
hand, portrays a rounded, lightly blushed greenish yellow fruit that closely
resembles the Florida mango. Furthermore, vegetative propagation of selected
chance seedlings has resulted in a variety of clonal types carried under this name in
India (Ratnam and Chellapa, no date, post-1954).

‘Nabeel’ (Egypt)

A minor commercial cultivar. The fruit is large (495 g), green with small yel-low
dots that are smooth; ovate-oblong (14 cm long by 9 cm wide by 7 cm thick), with
adherent skin relatively free of surface disease; the flesh is orange, firm and juicy,
without objectionable fibre, with a passable but not out-standingly pleasing taste
and acceptable quality. The seed is polyembryonic in a large (56.6 g) stone (Knight
and Sanford, 1998).

‘Nam Doc Mai’ (Thailand)

The tree is vigorous, medium sized with upright, dense canopy; the fruit (Plate 26)
is greenish to bright yellow with a slight pink blush and numerous
58 R.J. Knight et al.

small green dots, long and slender, sigmoid in shape with a rounded base, 17–19
cm long by 7.5–8.5 cm broad by 6.5–7.5 cm thick, weighing 340–580 g; the skin is
medium thick, tender and easily separated from the flesh which is soft, tender and
juicy with no fibre, lemon yellow, rich, spicy and very sweet with pleasant aroma,
of excellent quality; seed is polyembryonic in a thin, papery stone. This cultivar
ripens early midseason, fruits regularly and may have multiple crops in one season
(Campbell, 1992). It is highly resistant to foliar infection, and resistant to fruit
infection by bacterial black spot in Queensland (Mayers et al., 1988).

‘Neelum’ (India)

The tree is moderately vigorous with a small, compact canopy; the fruit is bright
yellow with no blush and numerous small white dots, oval with flat-tened or
slightly rounded base, 9.5–11 cm long by 7.5–8.5 cm broad by 6–6.5 cm thick,
weighing 230–300 g; the skin is thick, tender and easily sepa-rating; the flesh is
soft, melting and juicy with no fibre, deep yellow, mild and sweet with a
delightfully pleasant aroma, of good to excellent quality; seed is monoembryonic in
a medium-thick, woody stone. This cultivar is a late, heavy bearer (Campbell,
1992).

‘Nuwun Chan’ (Thailand)

The tree is moderately vigorous, small, upright with a dense canopy; the fruit (Plate
27) is greenish yellow with a pink to red blush, numerous small green dots, long
and slender with a flattened base, 16–18 cm long by 7–8 cm broad by 6–6.5 cm
thick, weighing 340–500 g; the skin is thick, tough and easily separating; the flesh
is soft, melting, juicy with little fibre, pale yellow, mild and sweet with a faint,
pleasant aroma, of good eating quality; seed is poly-embryonic in a thick, woody
stone. This cultivar is an early, regular bearer. Fruit is often eaten green (Campbell,
1992).

‘Okrung’ (Thailand)

The tree is moderately vigorous, medium sized and upright, forming a dense
canopy; the fruit (Plate 28) is green to greenish yellow with no blush and numerous
small white dots, oblong and sigmoid with a rounded base, 11–13 cm long by 5–6
cm broad by 4.5–5.5 cm thick, weighing 160–240 g; the skin is thick, tough and
medium adherent; the flesh is soft and juicy with abundant fibre, yellow or
greenish, mild, somewhat insipid and very sweet with a pleasant aroma, of good
quality; seed is polyembryonic in a thick, woody stone. This cultivar ripens
midseason, is a heavy producer and some-times bears more than one crop/year
(Campbell, 1992).
Mango Cultivars and Descriptors 59

‘Osteen’ (Florida, USA)

The tree is vigorous, medium sized, forming a dense canopy; the fruit (Plate
29) is yellow-orange with a purple or lavender blush and numerous small white
dots, oblong with rounded base, 12–15.5 cm long by 9–10.5 cm broad by 8.6–9.5
cm thick, weighing 500–760 g; the skin is thick, tough and easily sepa-rating; the
flesh is firm and juicy, with little fibre, lemon yellow, mild and sweet with a
pleasant aroma, of good quality; seed is monoembryonic in a thick and woody
stone. This cultivar ripens late midseason to late and is a regular producer. It is a
‘Haden’ seedling (Campbell, 1992; Schnell et al., 2006).

‘Pairi’ (India)

Also written ‘Pairie’, ‘Paheri’ and ‘Pirie’; synonyms are said to be ‘Peter’, ‘Peter
Pasand’, ‘Grape’, ‘Gohabunder’, ‘Nadusalai’, ‘Rasjuri’ and ‘Yerra Goa’. The tree
is moderately vigorous, forming a dense, rounded canopy; the fruit (Plate 30) is
medium sized, green to yellow with a bright red blush, roundish, skin smooth,
thick, flesh golden yellow, slightly juicy, fibreless, with a deli-cious subacid taste,
of excellent quality; the thick stone covered with short, bristly fibre encloses
monoembryonic seed (Popenoe, 1927; Singh, 1960). This cultivar has long been
popular as a dooryard fruit tree in Hawaii.

‘Palmer’ (Florida, USA)

The tree is moderately vigorous, forming a large, upright, tight canopy; the fruit
(Plate 31) is yellow-orange with a dark-red to crimson blush and a few small white
dots, oblong with rounded base, 12–15 cm long by 8.5–10 cm broad by 6.5–7.5 cm
thick, weighing 510–850 g; the skin is medium thick, tough and adherent; the flesh
is firm and melting with little fibre, orange-yellow, mild and aromatic, of good
quality; seed is monoembryonic in a medium-thick woody stone. This is a late
midseason cultivar and is a regular bearer. It is a seedling of ‘Haden’ (Schnell et
al., 2006). In Florida it is of minor commercial importance (Campbell, 1992). It is
grown in Israel and is the seed parent of ‘Naomi’. It is attracting increased attention
in the western hemi-sphere export market as a result of its superior eating quality.

‘Rosa’ (Brazil)

The tree is medium sized, of slow growth with a rounded canopy; the fruit (Plate
32) is yellow to rose-red on the side exposed to sun, oblong-cordiform and medium
sized; the skin is thick and smooth; the flesh is firm and mod-erately juicy, fibrous,
golden yellow, moderately sweet with a turpentine aroma, of ordinary quality,
susceptible to anthracnose disease; the seed is polyembryonic in a small, oblong
stone. This cultivar ripens midseason to
60 R.J. Knight et al.

late. It is one of the most important commercial cultivars in the Federal Dis-trict of
Brazil, used for juice as well as fresh consumption, and is one of the most well-
known cultivars in Brazil (Sampaio, 1980; L.C. Donadio, personal communication,
1996; A. Pinto, personal communication, 1996).

‘Sensation’ (Florida, USA)

The tree is vigorous, with a moderately open, symmetrical canopy; the fruit (Plate
33) is dark yellow with a prominent dark-red to purple blush that cov-ers most of
its surface, oval with rounded base and rounded apex, 9–11.5 cm long by 7–8 cm
broad by 6.5–7 cm thick, weighing 280–340 g; the skin is medium thick, tough and
easily separating; the flesh is firm and medium juicy, fibreless, deep yellow, mild
and sweet with a weak, pleasant aroma, of fair to good quality; seed is
monoembryonic in a thick, woody stone. This cultivar ripens midseason to late
(Campbell, 1992). It is a seedling of ‘Haden’ × ‘Brooks’ (Schnell et al., 2006), and
the seed parent of ‘B74’. It alter-nates severely, and in ‘on’ years the fruit may be
clustered so heavily that it becomes diseased before maturity, thus ‘Sensation’ is
not of commercial importance. It is highly resistant to bacterial black spot in
Queensland (May-ers et al., 1988), but often has severe internal breakdown
(browning, water soaking) (A.W. Whiley, personal communication, 1996).

‘Suvarnarekha’ (India)

Also called ‘Swarnarekha’ and ‘Sundri’. The tree is moderately vigorous and tall,
with a rounded, dense, spreading canopy; the fruit is light cadmium yel-low with a
blush of jasper red and abundant small, light-coloured dots, ovate oblong with a
base slightly flattened, of medium size, 11 cm long by 8.2 cm broad, weighing 400
g; the skin is medium thick, easily separated, flesh soft, fibreless, primrose yellow
with a pleasant aroma, sweet taste and abundant juice, of medium to good quality;
seed is monoembryonic in an oblong-oval stone covered with soft, short fibre.
Ripens early in the season early and is heavy bearing (Gangolly et al., 1957).

‘Tahar’ (Israel)

The tree is vigorous, medium sized, with an upright, dense canopy; the fruit is
bright yellow with a dark-red blush and numerous small white dots, ovate with
flattened base, 11.5–13 cm long by 8.9–9.5 cm broad by 7.5–8 cm thick, weighing
360–520 g; the skin is thick, tough and easily separating; the flesh is soft and juicy
with little fibre, deep yellow, mild, aromatic and slightly insipid with a strong
odour not appreciated by many, of fair to good quality; seed is monoembryonic in a
medium-thick woody stone. This cultivar ripens in late midseason and bears well in
Israel (Campbell, 1992).
Mango Cultivars and Descriptors 61

‘Taimour’ (Egypt)

A major commercial cultivar. The tree is vigorous; the fruit (Plate 34) is large (500
g), ripening late in the season, dark green with large light-brown dots, smooth in
texture, ovate-oblong (12.8 cm long by 8.4 cm wide by 8 cm thick), with non-
adherent skin of intermediate thickness, quite free from surface disease; flesh is
orange, firm (free of jelly seed) and juicy with no objection-able fibre, of a
delightfully rich, sweet taste and excellent quality. The seed is polyembryonic in a
medium-sized (50.8 g) stone (Knight and Sanford, 1998).

‘Tommy Atkins’ (Florida, USA)

The tree is vigorous, with a dense, rounded canopy; the fruit (Plate 35) is orange-
yellow, with a crimson or dark-red blush and numerous small, white dots, oval to
oblong, with broadly rounded base, 12–14.5 cm long by 10–13 cm broad by 8.5–10
cm thick, weighing 450–700 g; the skin is thick, tough and adherent; the flesh is
firm and medium juicy; with a medium amount of fibre, lemon to deep yellow,
mild and sweet with a strong pleasant aroma, of fair to good quality; seed is
monoembryonic in a thick, woody stone. This cultivar ripens early to midseason. It
is a ‘Haden’ seedling (Schnell et al., 2006). ‘Tommy Atkins’ is the most important
commercial cultivar in the western hemisphere export mango market; it is highly
resistant to anthracnose dis-ease and handling and shipping stress, and a consistent,
heavy producer (Campbell, 1992). ‘Jelly seed’ (internal breakdown) is a serious
problem in the moist subtropics and tropics outside Florida, where the mango is
grown on calcareous, well-drained soil (A.W. Whiley, personal communication,
1996).

‘Totapuri’ (India)

Also called ‘Bangalora’, ‘Collector’, ‘Kallamai’, ‘Killi’ (‘Gilli’), ‘Mukku’, ‘Sand-


ersha’ and ‘Thevadiyamuthi’. The tree is of medium size, vigorous, spread-ing with
an open canopy; the fruit (Plate 36) is greenish yellow with a pink blush and a few
small, white dots, oblong, base rounded, apex rounded to bluntly pointed with a
large beak, 17.5–20 cm long by 9–11.5 cm broad by 8.5–10.5 cm thick, weighing
800–1100 g; the skin is thick, tough and adherent; the flesh is firm and medium
juicy with a weak, somewhat repugnant aroma, of poor to fair quality; seed is
monoembryonic in a thin, papery stone. This cultivar ripens late midseason, is
productive and regular bearing. Fruit cracks when exposed to heavy rains at
ripening time. ‘Totapuri’ was imported to Florida twice, as ‘Sandersha’ in 1901
and as ‘Totapuri’ in the early 1960s. It is the seed parent of ‘Anderson’ and
‘Brooks’, which is itself the parent of ‘Kent’. It is called ‘Totapuri’ in Bangalore,
and ‘Bangalora’ in much of the rest of India (Gangolly et al., 1957; Campbell,
1992).
62 R.J. Knight et al.

‘Turpentine’ (West Indies)

The tree is vigorous, with a large, spreading, rounded canopy; the fruit (Plate
37) is bright yellow with a few large white dots, occasionally with a pink blush,
oval with a flattened base, 7.5–8.5 cm long by 6.5–7.5 cm broad by 6–6.5 cm thick,
weighing 140–200 g; the skin is thick, tough and easily sepa-rating; the flesh is
firm and juicy, with abundant coarse fibre, lemon yellow, rich, aromatic, spicy,
resinous and sweet with a strong, pleasant aroma, of poor to fair quality; seed is
polyembryonic in a thick, woody stone. This cul-tivar ripens early midseason to
late midseason and is a heavy producer but may alternate. It is commonly used as a
grafting stock (Campbell, 1992).

‘Vallenato’ (Colombia)

The tree is vigorous, with an upright, dense canopy; the fruit (Plate 38) is bright
yellow, with a crimson blush, oblong with flattened base, 8–9 cm long by 7–8 cm
broad by 6–7 cm thick, weighing 195–340 g; the skin is thin, tough and adherent;
the flesh is firm, juicy with abundant fine fibre (not objection-able), pale yellow,
mild and sweet with a strong, pleasant aroma, of good to excellent quality; seed is
monoembryonic. This cultivar ripens in early mid-season (R.J. Campbell, personal
communication, 1995).

‘Van Dyke’ (Florida, USA)

The tree is moderately vigorous, with a large, open canopy; the fruit (Plate
39) is bright yellow with a bright red or crimson blush, oval with rounded base, 9–
11.5 cm long by 7.5–9.5 cm broad by 7–8 cm thick, weighing 250– 520 g; the skin
is thick, tough and easily separating; the flesh is quite firm, melting and juicy with
little fibre, orange-yellow, rich, spicy and sweet with a strong, pleasant aroma, of
good to excellent quality, but susceptible to inter-nal breakdown; seed is
monoembryonic in a medium-thick, woody stone. This cultivar ripens in late
midseason and is a regular, heavy producer. It is a seedling of ‘Haden’ (Campbell,
1992; Schnell et al., 2006).

‘White Succari’ (Egypt)

A cultivar of major importance. The tree is vigorous; the fruit is medium large (410
g), greenish yellow with yellow overlay and small, brown dots of smooth texture,
ovate-oblong in shape (11.25 cm long by 8.3 cm wide by 8.0 cm thick), with thin
adherent skin reasonably free of surface disease; flesh is orange, yielding and juicy
with no objectionable fibres, of an agree-able sweet taste and very good quality.
The seed is polyembryonic in a moderate to large-sized stone (49 g), the season
early (Knight and Sanford, 1998).
Mango Cultivars and Descriptors 63

‘Zebda’ (Egypt)

A cultivar of major importance. The tree is vigorous and regularly produc-tive; the
fruit (Plate 40) is large (660 g), green with no overlay and small, brown dots of
smooth texture, oblong-cylindrical in shape (14.6 cm long by 9.7 cm wide by 8.3
cm thick), with non-adherent skin quite free of surface disease; flesh is deep
orange, firm and juicy, with no objectionable fibre and a mild, sweet taste, of
acceptable quality. The seed is polyembryonic in a moderately small (52 g) stone.
This cultivar is of late-midseason maturity (Knight and Sanford, 1998). It is highly
tolerant of anthracnose and resistant to malformation (R.C. Ploetz, personal
communication, 2007).

3.4 Conclusion
The mango fruit’s nutritional value, aesthetic and gustatory appeal have assured its
growing importance in non-traditional markets since the late 1950s, as it has been
introduced to consumers previously unacquainted with it. Furthermore, the
migration of large populations from South-east Asia and other regions where this
fruit is a traditional crop to metropolitan centres where it has not been well known
has created a permanent demand for it in these new markets. An additional factor
permitting market expansion has been the growing mango production in areas
previously unimportant in world commerce such as Mexico, Brazil, Australia,
West Africa, Israel, Flor-ida and the Canary Islands. The fact that most new
markets are remote from areas of production has necessitated selection of cultivars
for fresh market sale that are dependably productive and resistant to harvest,
handling and shipping stress, with relatively long shelf life, for example ‘Tommy
Atkins’, ‘Keitt’ and ‘Madame Francis’. The fruit quality of mango cultivars well
suited to packing and shipping has been a secondary consideration, and is gener-
ally not so high as that of cultivars acknowledged to be superior for eating.
Economic factors obviously must dictate what is grown for the fresh market.

The commercial market for processed mango products permits other cultivars
to be utilized, and these may vary with the product that is mar-keted. Cultivars
chosen for purée or juice preparation are likely to be quite different from those
used for manufacture of chutney or other products requiring pulp that maintains its
integrity after it is cooked. ‘Totapuri’ (‘Sand-ersha’) or ‘Turpentine’, for example,
considered mediocre for fresh consump-tion, can be used to prepare excellent
chutneys, as can many ‘criollo’ types in the West Indies. ‘Tommy Atkins’ makes
outstandingly good dried fruit sec-tions, sweet and aromatic, even though its fresh-
fruit quality is generally conceded not to be high. Mango butter and mango leather
are other products that are appreciated by many who know them (see Raymundo et
al., Chapter 17, this volume; Campbell and Campbell, 1983; Campbell and Smith,
1987). As more fruit that is wholesome, but not of export quality, becomes
available in areas of increasing production, it is likely that processed mango
products will become more common.
64 R.J. Knight et al.

Table 3.1. Ratings of selected mango cultivars grown in Florida (Source: Knight, 1993).

a b c d e f g h i
Cultivar Shape Size Firmness Colour Anthracnose Fibre Taste Yield Score

‘Alphonso’ 3 5 7 2 3 7 9 1h x
‘Boribo’ 3 8 8 4 7 9 5 6 x
h
‘Carabao’ 5 6 7 3 5 9 8 6 x
h
‘Haden’ 3 9 8 8 5 7 7 3 x
‘Keitt’ 4 10 9 6 8 9 8 8 ///
‘Kensington’ 3 8 7 7 7 8 7 6 /
‘Langra’ 2 6 8 3 5 8 8 3h x
‘Pope’ 3 9 5 7 2 8 8 1 x
‘Tommy 3 9 9 9 9 6 6 7 ///
Atkins’
‘Van Dyke’ 3 7 10 9 7 8 7 6 ///
a
Ratings of 1 (round) to 5 (long) indicate fruit shape, not its desirability.
b
Ratings below 6 justify discard; those of 7 and above show size only, not merit.
cRatings of 1–10 where 1 = least and 10 = most.
d
Ratings of 1–10 where 1 = least and 10 = most.
e
Ratings of 1–10 where 1 = most and 10 = least susceptible.
fRatings of 1–10 where 1 = most and 10 = least.
g
Ratings of 1–10 where 1 = worst and 10 = best.
hTrends markedly towards alternate bearing.
i
One or more checks (/) show overall value; (x) indicates no commercial acceptability.

Despite the recognized high quality of many well-known mango cultivars,


considerable cultivar improvement is still needed in most regions of culture before
anything approaching perfection is likely to be achieved (Table 3.1). For any given
area, cultivars that combine adequate resistance to disease and packing and
shipping stress, regular heavy production, high quality, and attractive appearance
throughout a long bearing season are all requisites. Production of seedlings from
controlled crossing of different parents having desired characters, followed by
vigorous selection and evaluation of the resultant selections, can produce such
improved cultivars. Pursuit of com-mon goals, including the cooperative exchange
and testing of elite germ-plasm in different regions of production, can accelerate
progress towards this objective (Lavi et al., 1989, 1993; Knight, 1993). Such
interregional and inter-national activities are to be encouraged because of their
potential for advanc-ing mango production and utilization in the world.

Acknowledgements
For their help in reviewing portions of this chapter and/or contributing vast
quantities of information on mango cultivar descriptors and attributes, the authors
are profoundly grateful to the following: Dr N. Balasundaram, Head, Sugarcane
Breeding Institute, Regional Centre, Karnal, India 132001; the late Dr Carl W.
Campbell, Tropical Research and Education Center, 18905 SW 280
Mango Cultivars and Descriptors 65

Street, Homestead, Florida 33031 USA; Luis C. Donadio, Universidade Estad-ual


Paulista, Rodavia Carlos Tonanni KM5, Jaboticabal, 14870, São Paulo, Brazil; Dr
Shmuel Gazit, Department of Horticulture, Hebrew University of Jerusalem,
Rehovot 76100, Israel; Mr Remy LePrette, Directeur, Interfel, 155 Rue F.G.
Poissonierre, Paris 75009, France; Dr Alberto C. Pinto, Lider de Pro-jeto/CPAC,
EMBRAPA, Caixa Postal 08 223, CEP 73301-970, Brasilia, DF, Brazil; Dr Eli
Tomer, Department of Fruit Trees, the Volcani Centre, Institute of Horticulture, PO
Box 6, Bet Dagan 50-250, Israel; Dr Anthony W. Whiley, Maroochy Horticultural
Research Centre, PO Box 5093 SCMC, Nambour, Queensland 4560, Australia.

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nitrate. HortScience 14, 527–528.
Campbell, B.A. and Campbell, C.W. (1983) Preservation of tropical fruits by drying.
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Campbell, B.A. and Smith, J. (1987) An overview of tropical fruit uses in Florida.
Pro-ceedings of the Florida State Horticultural Society 100, 408–411.
Campbell, C.W. and Campbell, R.J. (1995) ‘Cogshall’, a mango for the home garden.
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Campbell, R.J. (ed.) (1992) A Guide to Mangos in Florida. Fairchild Tropical
Garden, Miami, Florida, USA.
Campbell, R.J., Ledesma, N. and Campbell, C.W. (2002) Tropical Mangos: How to Grow
the World’s Most Delicious Fruit. Fairchild Tropical Garden, Miami, Florida, USA.
Gangolly, S.R., Singh, R., Katyal, S.L. and Singh, D. (1957) The Mango. Indian
Council of Agricultural Research, New Delhi, India.
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International Board for Plant Genetic Resources, Rome.
Knight, R.J., Jr (1993) Evaluating important fruit characters in mango germplasm.
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‘Haden’ cultivar’s significance to the modern industry. Economic Botany
48, 139–145.
Lavi, U., Tomer, E. and Gait, S. (1989) Inheritance of agriculturally important traits in
mango. Euphytica 54, 5–10.
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breeding of mango cultivars and rootstocks. Acta Horticlturae 341, 146–151.
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in the Soconusco region of Chiapas, Mexico. Acta Horticulturae 645, 361–363.
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control of bacterial black spot (Xanthomonas campestris pv. mangiferaeindicae)
of mango. 1. Evaluation of 23 cultivars of mango for foliar and fruit resistance to
bacterial black spot under orchard conditions at Childers, south east
Queensland. Maroochy Hor-ticultural Research Report 5, 100–101.
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Mukherjee, S.K. (1985) Systematic and Ecogeographic Studies of Crop Gene Pools. 1.
Mangifera L. International Board for Plant Genetic Resources, Rome.
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Intervari-etal hybridization in mango (Mangifica indica L.): techniques, main
results and their limitations. Acta Horticulturae 645, 327–330.
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Prasad, A. (1977) Bearing behaviour and fruit quality of south Indian varieties of mango
in northern India. Indian Journal of Horticulture 34, 372–376.
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Horticulture Ltd, Nambour, Australia.
4 Breeding and Genetics

C.P.A. Iyer1 and R.J. Schnell2


1
Indian Institute of Horticultural Research, Bangalore, India
2
USDA ARS, National Germplasm Repository, Miami, Florida, USA

4.1 Introduction 68
4.2 Origin of Cultivars and Distribution 68
Mangifera species and mango 68
History of cultivation 69
Impact of Florida mangoes 70
4.3 Reproductive Mechanisms 70
Polyembryony 70
Floral biology and pollination 71
Incompatibility 72
Cytology 72
4.4 Inheritance of Characters 73
Dwarfness, regular bearing and precocity 73
Flesh colour 74
Skin colour 74
Flowering time 74
Beak 74
Disease resistance 74
Other horticultural traits 75
4.5 Breeding Objectives 75
General objectives 75
Specific objectives 76
4.6 Methods of Breeding 78
Selection from open-pollinated seedlings 78
Controlled pollination 79
4.7 Handling of Hybrid Populations and Selection 80
Criteria for initial selection 80
Pre-selection 81
Potential for marker assisted selection (MAS) 81
Molecular markers 81
4.8 Minimizing Problems in Breeding 83
Heavy fruit drop 83
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 67
68 C.P.A. Iyer and R.J. Schnell

Long juvenile phase 83


Polyembryony 84
4.9 Achievements of Conventional Breeding 84
India 85
Other countries 86
4.10 Mutations 87
Somatic mutations 87
Induced mutations 88
4.11 Breeding Potential of Wild Species 88
4.12 Conclusions 89

4.1 Introduction
Mango has been considered to be a difficult plant species to improve in breeding
programmes because of certain inherent characteristics including:
(i) a long juvenile phase; (ii) a high level of heterozygosity resulting in unpre-
dictable outcomes in hybridization; (iii) only one seed per fruit; (iv) heavy fruit
drop leading to low retention of crossed fruits; (v) polyembryony in many
cultivars; and (vi) the large area required for a meaningful assessment of hybrids.
Despite these drawbacks, mango breeding can be successful because of its wide
range of genetic variation and the ease with which a selected hybrid can be
vegetatively propagated. Barring a few hybrid variet-ies resulting from planned
hybridization programmes, which are now gain-ing increased attention, almost all
known cultivars have resulted from the selection of chance seedlings from natural
cross-pollinations. However, in Florida, a number of cultivars have resulted from
the screening of seedlings from known mother plants. Most of the present-day
mango cultivars were selected on the Indian subcontinent; these selections were
made based mainly on fruit quality, with very little emphasis on modern
horticultural and industrial requirements. These requirements include precocity,
dwarf-ness, heavy and regular bearing, absence of physiological disorders, resis-
tance to disease and pests and good shipping qualities. With decreasing land
availability and the rising cost of labour, tree architecture requirements have also
changed. The need for new cultivars to meet these requisites pinpoints the
importance of planned hybridization rather than merely depending on chance
seedlings. Current knowledge of hybridization techniques, inheri-tance patterns,
management of hybrid populations and the development of genetic markers have
greatly reduced the uncertainty in mango breeding.

4.2 Origin of Cultivars and Distribution


Mangifera species and mango

Almost all the commercial cultivars belong to Mangifera indica. However, a few
commercial varieties of South-east Asia belong to other species, i.e. M. altissima,
M. caesia, M. foetida, M. griffithi, M. odorata, M. pentadra, M. sylvatica,
Breeding and Genetics 69

M. zeylanica, M. laurina, M. lagenifera, M. cochinchinensis, etc. The monoem-


bryonic mango (M. indica) originated in north-eastern India (Assam), the Indo-
Myanmar border region and Bangladesh (Chittagong Hill tract), where it is still
found as a wild tree, with very small fruits. It may also occur in the lower
Himalayan tract, near Nepal, Bhutan and Sikkim. Polyembryonic mangoes are
considered to have originated in South-east Asia. Wild man-goes, representing
different Mangifera species, can be found in tropical Asia, particularly north-
eastern India, Sri Lanka, Myanmar, Thailand, Indo-China, southern China,
Malaysia, Indonesia, Papua New Guinea, the Philippines and as far as the Solomon
and Caroline Islands in the east. There are more than 60 species worldwide. The
highest specific diversity is found in the heart of the distribution area of the genus
Mangifera; the Malay Peninsula, Borneo and Sumatra (Bompard, 1993).

History of cultivation

Mango has been cultivated in India for at least 4000 years and over 1000 vari-eties
are recognized there (Mukherjee, 1953). Almost all of them are selec-tions made
from naturally occurring, open-pollinated seedlings. However, based on random
amplification of polymorphic DNA (RAPD) analysis, Rav-ishankar et al. (2004)
felt that the mono- and polyembryonic types of Indian mango cultivars have a
different genetic base, and that the polyembryonic types might have been
introduced from South-east Asia and are unlikely to have originated in India.
Mango culture gradually spread to tropical and subtropical countries throughout the
world, where selections were made that were adapted to particular growing
conditions. Thus, selection by man has played the most significant role in the
development of new mango culti-vars. The explorers who tasted the mango in the
regions of its origin were enchanted with its aromatic qualities, ambrosial flavour
and creamy, smooth and silky texture, and introduced the fruit to other tropical
regions. The spread of Hinduism and Buddhism assisted in the distribution of
mangoes in South-east Asia. The Chinese traveller Hwen T’sang who visited India
in the first half of the 7th century AD returned to China with the mango. The mango
was known in Baghdad in the 7th century.

The Persians or Omanis may have taken mangoes to East Africa around the
10th century AD. The fruit was introduced to East and West Africa in the early 16th
century by the Portuguese and thence into Brazil. After being established in Brazil,
the mango was carried to the West Indies, being first planted in Barbados by about
1742 and later in the Dominican Republic and Jamaica (about 1782). The mango
was introduced into Mexico from the Phil-ippines by the Spanish and also from the
West Indies (Morton, 1987). Duval et al. (2006) developed microsatellite markers
for studying the genetic vari-ability of Caribbean mangoes and concluded that there
were two routes of mango to the French West Indies, namely, cultivars grown in
Central Amer-ica (Mexico) and South America (Colombia) introduced from South-
east Asia and also from former French colonies in the Indian Ocean. As the mango
70 C.P.A. Iyer and R.J. Schnell

adapted to new locales, new cultivars were selected based on local adapta-tion and
fruit preferences.

Impact of Florida mangoes

The first recorded successful introduction of mango into Florida (USA) was made
in 1861 (Knight, 1980). The earliest introductions were from the West Indies and
India, followed by the introduction of several hundred accessions in the 20th
century from South-east Asia, India and from other mango-growing areas of the
world (Florida Mango Forum, 1951). The introduction of mangoes into Florida and
subsequent development of a Florida group of mangoes has been reviewed by
Knight and Schnell (1994). The Florida mango cultivars are unique in that they are
hybrids between Indian cultivars (primarily mono-embryonic) and the South-east
Asian cultivars (primarily polyembryonic) selected under south Florida conditions.
The mango breeding system favours out-crossing. Therefore, the proximity of
numerous genotypes of disparate geographical origins led to the production of
many new seedlings by inter-pollination in Florida (Knight and Schnell, 1993).
Florida selections are therefore not the result of a formal breeding programme.
Early Florida selec-tions were made by growers and enthusiasts and historical
information is often anecdotal. The Florida Mango Forum, established in 1938 for
the advancement of mango production, documented historical information on the
parentage of Florida cultivars in their proceedings. In addition to the United States
Department of Agriculture (USDA) Germplasm Resources In-formation Network
(GRIN) database, several sources compile information on Florida mango selections
and introduction of accessions to Florida (Ruehle and Ledin, 1956; Singh, 1960;
Campbell and Campbell, 1993; Schnell et al., 2006). With the exception of South-
east Asia, Australia and some African countries, which cultivate mostly locally
selected varieties, the majority of countries produce cultivars developed in Florida,
i.e. ‘Haden’, ‘Tommy Atkins’ and ‘Keitt’ (Galan Sauco, 1997). These Florida
selections are now widely grown commercial cultivars affording production
stability across many environments (see Mukherjee and Litz, Chapter 1, this
volume).

4.3 Reproductive Mechanisms

Polyembryony

Nucellar embryos
Mangoes can be classified into two groups, monoembryonic and polyembry-onic,
based on their mode of reproduction from seeds. In general, monoem-bryonic seeds
are found in the sub-tropical group (Indian type) and the polyembryonic seeds in
the tropical group (South-east Asian). Monoembry-onic mango seeds each contain
a single zygotic embryo, and hence only one seedling per seed that is of probable
hybrid origin. Polyembryonic mango
Breeding and Genetics 71

seeds can contain one or more embryos, one of which is usually, but not always
zygotic. Adventitious embryos develop from the nucellus, a maternal tissue
surrounding the embryo sac, and consequently the seedlings of poly-embryonic
mangoes are genetically identical to the maternal parent. Adven-titious embryos
can also originate by direct budding from the cotyledons and hypocotyls of other
nucellar embryos (Juliano, 1934). According to Mahesh-wari and Rangaswamy
(1958), the nucellar cells destined to form adventi-tious embryos are recognizable
by their dense cytoplasm and starchy contents. They gradually push into the
embryo sac cavity where they divide and differentiate into embryos.

Inheritance of polyembryony
Polyembryony is genetically determined. Leroy (1947) considered that adven-tive
embryony probably reflects the effect of one or more recessive genes. This view
was supported by Sturrock (1968), whose study of the progenies of
monoembryonic mango hybridized with polyembryonic cultivars indicated that
monoembryony was possibly a dominant trait. In contrast, Aron et al. (1998) and
Brettell et al. (2004) observed that polyembryony in mango is con-trolled by a
single dominant gene. Schnell et al. (2006) reported that 58 of the Florida cultivars
had been classified with 50 being monoembryonic and eight polyembryonic.
Information from the Florida cultivars parentage analysis using 25 microsatellite
markers supported the findings of Aron et al. (1998) where polyembryony was
found to be dominant. ‘Haden’ is a cross of the monoembryonic ‘Mulgoba’ and the
polyembryonic ‘Turpentine’. If we assume that a single dominant gene controls
this trait, all of the Indian cultivars in Florida must be homozygous recessive and
the ‘Turpentine’ parent of ‘Haden’ must have been heterozygous. The evidence
suggests that ‘Haden’ inherited the recessive allele from ‘Turpentine’, as all
identified progeny of ‘Haden’ are monoembryonic with the exception of ‘Winters’.
The most probable pollen parent of ‘Winters’ is ‘Ono’, a polyembryonic cultivar
from Hawaii. The fre-quency of this dominant allele is low in the Florida
population and absent from the Indian cultivars in Florida. In view of these
interesting findings, and since a thorough knowledge of inheritance of
polyembryony is essential for speculating the origin of M. indica, more work on
these lines is warranted.

Floral biology and pollination

The mango inflorescence is primarily terminal, although axillary and multi-ple


panicles may also arise from axillary buds. Both perfect (hermaphrodite) and
staminate (male) flowers occur in the same inflorescence. The total num-ber of
flowers in a panicle may vary from 1000 to 6000, depending on the cultivar
(Mukherjee, 1953). Initial fruit set in mango is directly related to the proportion of
perfect flowers, although the final fruit set does not necessarily depend on this ratio
(Iyer et al., 1989). It appears that the proportion of per-fect flowers in a cultivar
becomes critical for optimum fruit set only when the proportion drops to 1%.
72 C.P.A. Iyer and R.J. Schnell

Flowers begin to open early in the morning and anthesis has generally been
completed by noon. The greatest number of flowers opens between 9 and 10 a.m.
Although the receptivity of the stigma continues for 72 h after anthesis, it is most
receptive during the first 6 h; however, there are reports that the stigma can become
receptive even before anthesis has occurred (Singh, 1960). The minimum time
required for pollen grains to germinate is 1.5 h (Sen et al., 1946; Singh, 1954;
Spencer and Kennard, 1955). Singh and Singh (1952) observed 98% pollen
viability after 11 months in storage at 7C and 25% relative humidity (RH), and
65.7% viability after 24 months of stor-age at 0C and 25% RH.

Mango is cross-pollinated, which is carried out by insects such as the common


housefly, honeybees and thrips, and possibly by other insects al-though to a lesser
extent. Pollination by wind and gravity has been sug-gested to occur in mango
(Popenoe, 1917; Maheshwari, 1934; Malik, 1951). In nature, > 50% of flowers do
not receive any pollen. Some workers had sug-gested that self-pollination in certain
cultivars can also occur quite frequently (Dijkman and Soule, 1951). Studies by
Issarakraisila and Considine (1994) have shown that for polyembryonic
‘Kensington’, a night temperature of
< 10C results in pollen grains with low viability (< 50%). The optimum
temperature for normal meiosis is between 15 and 33C with 70–85% viability.

Incompatibility

Although the existence of self-sterility in mango was suspected several years ago
(Ruehle and Lynch, 1948, cited in Sharma and Singh, 1970; Dijkman and Soule,
1951), the prevalence of self-incompatibility was clearly established in
monoembryonic ‘Dashehari’ by Singh et al. (1962). Subsequently, detailed studies
indicated that the four popular monoembryonic cultivars of northern India (i.e.
‘Dashehari’, ‘Langra’, ‘Chausa’ and ‘Bombay Green’) were self-incompatible
(Mukherjee et al., 1968; Sharma and Singh, 1970). Embryo-logical studies have
shown that although fertilization takes place after self-pollination, degeneration of
endosperm occurs 15 days after pollination involving self-incompatible parents
(Mukherjee et al., 1968). The self-incompatibility system operating in mango
appears to be of the sporophytic type. Instances of cross-incompatibility among
certain mango cultivars have also been reported (Ram et al., 1976), necessitating
the identification of suitable pollinizers for mango.

Using an approach involving isozyme analysis, Dag et al. (2006) have initiated
studies in many commercial mango cultivars in Israel and con-cluded that self-
pollination is not a yield-limiting factor in monoembryonic ‘Maya’ and the practice
of planting ‘Maya’ in solid blocks is sound. They had obtained similar results
earlier with monoembryonic ‘Tommy Atkins’ (Dag et al., 1997).
Breeding and Genetics 73

Cytology

Chromosome number
Information on the cytology of mango is quite limited. Only a few Mangifera
species (i.e. M. indica, M. caloneura, M. sylvatica, M. foetida, M. caesia, M.
odorata and M. zeylancia) have been studied, and were found to have chromosome
numbers of 2n = 2x = 40 and n = x = 20 (Mukherjee, 1950, 1957; Roy and
Visweswariya, 1951). Chromosome numbers and ploidy status of other species are
yet to be studied. The only exception to this chromosome number that has been
reported to date (Roy and Visweswariya, 1951) involves ‘Vallikolamban’, which
was reported to be tetraploid (2n = 4x = 80), although subsequent stud-ies have
indicated that it is only a diploid (Majumder and Sharma, 1990).

Polyploidy
Mango has been referred to as an allopolyploid. Due to the presence of secondary
associations at metaphase of meiosis, Mukherjee (1950) suggested that the basic
chromosome number of Mangifera is n = 8. In addition, the high number of
somatic chromosomes and the correspondingly high number of nucleolar
chromosomes led him to conclude that mango is an allopolyploid. However, the
evidence used to arrive at this conclusion is not unequivocal. In fact, the molecular
marker evidence is antithetical to this conclusion. Results from Duval et al. (2005),
Viruel et al. (2005) and Schnell et al. (2005, 2006) using microsatellite markers all
indicate that M. indica is diploid.
Although many wild Mangifera species are potentially valuable for crop
improvement, they are yet to be exploited. Mukherjee (1963) felt that the different
Mangifera species could intercross easily, based on the success ob-tained with
interspecific crosses between M. zeylanica and M. odorata.

4.4 Inheritance of Characters


High heterozygosity in the cultivars that are used in hybridization and the
inadequate number of hybrid progenies realized has made accurate genetic analysis
in mango very difficult. However, based on limited data, some indi-cations are
available which would be useful in selecting parents in breeding programmes
designed with specific objectives. In studies of the distribution of different traits in
seedlings derived from open-pollination (where the pol-len parent is unknown),
Lavi et al. (1989) observed: (i) there is no maternal effect on juvenile period and
fertility; (ii) there is a slight effect of the female parent on fruit taste and size; (iii)
there is a maternal parent effect on harvest season and fruit colour.

Dwarfness, regular bearing and precocity

An analysis based on observations of more than 1000 hybrids, involving sev-eral


combinations, has revealed that dwarfness, regular bearing and precocity
74 C.P.A. Iyer and R.J. Schnell

are controlled by recessive genes (Sharma and Majumder, 1988a). Regularity of


bearing appears to be linked with precocity. Characters contributing to biennial
bearing are dominant over those governing regular bearing habit.

Flesh colour

Sharma (1987) considered that additive gene action may be involved in the
inheritance of flesh colour; however, studies involving monoembryonic ‘Alphonso’
and ‘Neelum’ have indicated that light yellow is dominant over orange-yellow
(Iyer, 1991).

Skin colour

With regard to skin colour of fruit, Sharma (1987) observed that when red cultivars
were crossed with green cultivars, the F1 seedlings exhibited vari-ous gradations of
red. Iyer and Subramanyam (1987) also found a wide array of colours in the
hybrids when the coloured monoembryonic ‘Janardhan Pasand’ was crossed with
green-fruited cultivars, indicating that colour is mediated by a number of loci.

Flowering time

The flowering response of mango cultivars in subtropical and tropical envi-


ronments differs greatly (see Davenport, Chapter 5 this volume). Trees can be
stimulated to flower under certain conditions in tropical environments using
ethephon; however, this is ineffective in subtropical environments. Schnell and
Knight (1998) investigated the repeatability of flowering using eight cultivars over
six harvest cycles (years), collecting data weekly. Three characters were evaluated:
days to bloom (DTB) (from 1 November in each year), days in bloom (DIB), and
days in bloom and fruit (DIBF). Significant differences were detected for all three
characters for both years and cultivars. Significant differences were not detected
for replicate trees within cultivars. Repeatability (R) of the flower phenology
characters was high (R = 0.73, 0.88 and 0.77 for DTB, DIB and DIBF,
respectively). This indicates that much of the variation is heritable and useful for
extending flowering times in subtropical environments.

Beak

The presence of a beak on mango fruit appears to be a dominant character since


most of the hybrid plants had this feature when monoembryonic ‘Ban-galora’
(‘Totapuri’) was used as one of the parents in controlled crosses (Iyer and
Subramanyam, 1979). Bunch bearing was found to be a dominant char-acter, as
indicated in many crosses (Sharma et al., 1972) involving bunch-bearing types
with single-fruited cultivars.
Breeding and Genetics 75

Disease resistance

Bacterial canker (Xanthomonas campestris pv. mangiferaeindicae) resistance


appears to have cytoplasmic inheritance. Whenever ‘Neelum’, a susceptible
cultivar, is used as the female parent, susceptibility is transmitted to all the hybrids,
irrespective of the male parent (Sharma and Majumder, 1988a). It has been
suggested that internal breakdown (spongy tissue) is mediated by recessive genes
(Iyer, 1991). Susceptibility to ‘mango malformation’ appears to be dominant, since
crosses involving resistant ‘Bhadauran’ did not yield any resistant hybrids (Sharma
and Majumder, 1988a).

Other horticultural traits

The genetics of inheritance of various horticultural traits appears to be unclear.


More knowledge will be forthcoming only when large-scale con-trolled
hybridization experiments are undertaken at different mango research centres.
However, the information now available for some of the characters is very useful in
deciding which parental genotypes ought to be used in hybridization programmes.

Brettell et al. (2004) subjected the large number of mango hybrids obtained
from the Australian National Mango Breeding Programme to a biometrical
analysis. Their data indicated that many of the important fruit quality aspects,
including fruit weight, fruit shape, ground skin colour, fruit width and pulp depth
have high heritabilities and can therefore be readily selected in a breed-ing
programme. Of particular interest is the observation that a high frequency of
hybrids with a red or burgundy blush can be recovered from crosses where one
parent has an intense red blush. Similarly, while the unique flavour com-pounds
associated with ‘Kensington Pride’ are also found in nearly 50% of the hybrids
involving ‘Kensington Pride’, leaf fragrance was not found to be a reliable
predictor of fruit flavour.

4.5 Breeding Objectives

General objectives

Breeding objectives vary from region to region, depending on the specific trait(s)
for which improvement is sought. However, they can be broadly gen-eralized to
consist of the development of cultivars with: (i) regular bearing;
(ii) dwarf tree habit; (iii) precocity; (iv) attractive, good sized (300–500 g), good
quality fruits (appealing flavour and firm flesh without fibres); (v) resistance to
major diseases and pests; (vi) freedom from physiological dis-orders; and (vii)
good shipping qualities and shelf life. While it would be hard to combine all these
characteristics within a relatively short time, espe-cially resistance to all major
diseases and pests, all of these characteristics are basic for commercial success.
76 C.P.A. Iyer and R.J. Schnell

With regard to the improvement of rootstocks by breeding, the main desirable


features are: (i) polyembryony; (ii) dwarfing; (iii) tolerance of adverse soil
conditions (high pH, calcareous soil, etc.); and (iv) good scion-compatibility.

Specific objectives

In addition to improving general characters such as yield and quality, breed-ing has
also been undertaken for certain specific purposes.

Dwarfness
Because of the obvious benefits of comparatively dwarf trees for orchard man-
agement and fruit quality, attempts have been focused on obtaining hybrids with a
dwarf tree framework. Breeding for dwarfness is important in mango, since a
consistent dwarfing effect of any rootstock has not been established to date. The
Indian cultivars that could be useful as a source for dwarfness include ‘Kerla
Dwarf’, ‘Janardan Pasand’, ‘Manjeera’, ‘Amrapali’, ‘Creeping’ (Iyer and
Subramanyam, 1986) and ‘Nileswar Dwarf’ (Singh, 1990).

Regular bearing
The causes of irregular bearing vary from region to region. In general, the main
reason for alternate bearing, particularly in subtropical India, is the lack of
initiation of vegetative growth soon after fruiting. However, two cul-tivars,
‘Neelum’ and ‘Bangalora’ (‘Totapuri’), which are regular bearers, have been
extensively used as either of the parents in a hybridization programme to transfer
the regular bearing habit to hybrids. ‘Neelum’ has been observed to be a good
combiner and has contributed to the evolution of many regular-bearing Indian
hybrid cultivars. However, ‘Bangalora’ is not a suitable par-ent since the hybrids
possess very prominent beaks and their fruit quality is invariably poor. The regular
bearing Florida cultivars (i.e. ‘Tommy Atkins’, ‘Keitt’, etc.) also have potential as
parents.

Fruit colour
Most of the commercial cultivars in South-east Asia possess green skin. Efforts are
underway to produce new hybrid cultivars that retain the good qualities of these
fruits together with attractive skin colour, so that they will occupy a better position
in international trade. Since good skin colour has been shown to be transmissible to
hybrids from suitably coloured parental cultivars, a number of cultivars with
coloured skin are being used for hybrid-ization. In general, the attractively
coloured Florida cultivars have been found to be suitable parents. In addition, there
are several Indian cultivars (e.g. ‘Janardan Pasand’, ‘Suvarnarekha’, etc.) that
would be suitable for use as parents for this purpose.

In Florida, the skin colour of the mango is an important factor and red skin is
considered essential for mangoes shipped to northern markets. In the past, the
evaluation of mango colour has been subjective and based on visual
Breeding and Genetics 77

ratings. Large errors are associated with these types of ratings, which makes
evaluation based on fruit colour difficult. Ayala-Silva et al. (2005) used a colo-
rimeter to quantify fruit colour, quality and differentiation among cultivars. Mango
colour was measured with a Minolta Chroma Meter CR-400 (Osaka, Japan)
portable tristimulus colorimeter and fruit chromaticity was recorded in
Commission Internationale de l’Eclairage (CIE) L *, a* and b* colour space
coordinates. In this system of colour representation the values L *, a* and b *
describe a uniform three-dimensional CIE colour space. If the L *, a* and b*
coordinates are known, then the colour is not only described, but also located in
quantifiable space. Maternal half-sib families (MHS) of the mango culti-vars,
‘Keitt’, ‘Tommy Atkins’, ‘Tyler Premier’, ‘Mamita’, ‘White Alfonso’ and
‘Sandersha’ were evaluated along with two parental check clones, ‘Tommy Atkins’
and ‘Keitt’. Significant differences were found for each of the L *, a* and b* colour
space coordinates. Further work is underway to estimate the herita-bility of these
traits to estimate their usefulness for breeding and selection.

Disease resistance
MANGO MALFORMATION. Although no breeding work has been reported that spe-cifically
addresses disease or pest resistance/tolerance, cultivars are known to show varying
degrees of susceptibility to biotic stress (see Ploetz and Freeman, Chapter 8, this
volume ). Mango malformation, caused by Fusarium subglutin-ans, is a very serious
disease that has threatened the very survival of the mango industry in many subtropical
mango-growing regions. As there are no reliable cultural and chemical control
measures available, breeding for resistance/tolerance has been attempted using
‘Bhadauran’ as the resistant parent; however, all of the F1 hybrids were susceptible to
the disease (Sharma and Majumder, 1988a). In this respect, the observations of Ram et
al. (1987) are very encouraging. Out of 102 cultivars screened, three of them, namely,
‘Bhydayam Dula’, ‘Samar Bahist Rampur’ and ‘Mian Sahib’, were free of
malformation and could be tried as one of the parents in hybridization.

BACTERIAL CANKER. Bacterial canker is a serious problem with many cultivars. The only
cultivar possessing true resistance to canker is ‘Bombay Green’ (Prakash and
Srivastava, 1987) and hence could be a potential gene donor.

ANTHRACNOSE. Anthracnose, caused by Colletotrichum gloeosporioides Penz., is the


most widespread disease in all mango-growing countries, manifesting itself in blossom
blight, peduncle blight, leaf spot, twig blight, wither tip, fruit russetting and fruit rot.
‘Tommy Atkins’ is moderately tolerant of anthracnose and coupled with its other
desirable qualities (i.e., regular bearing, fruit colour, etc.) should be a good parent in
breeding programmes. In addition, ‘Parish’ and ‘Fairchild’ have been reported to be
relatively resistant (Yee, 1958).

POWDERY MILDEW . Powdery mildew caused by Oidium mangiferae Berthet, has been
reported to cause heavy loss of crops in years when RH is very high and accompanied
by cool nights during flowering. Cultivar differences with respect to susceptibility are
recognized, and ‘Pairi’ (‘Raspuri’) is highly susceptible.
78 C.P.A. Iyer and R.J. Schnell

Gupta (1976) has listed those cultivars that are most tolerant of this disease –
‘Neelum’, ‘Zardalu’, ‘Bangalora’, ‘Totapuri-Khurd’ and ‘Janardan Pasand’ – and
hence could be valuable in breeding programmes.

PEST RESISTANCE. Considerable variation is also known to occur among mango cultivars
with respect to their susceptibility to attack and injury by insect pests. Although no
resistant genotypes have been reported for the mango hopper (Idiocerus spp.), the
insect has been observed to avoid colonizing open plant types where free movement of
wind is possible, an observation that could be useful in selection. Although complete
resistance is not known to either fruit fly (Bactrocera spp.) or seed weevil (Stenochetus
mangiferae), variation in the degree of susceptibility has been reported (Iyer, 1991).
Rossetto et al. (2006) observed that resistance to fruit fly is compatible with fruit
quality and pro-ductivity and advocated that resistance to fruit fly should be one of the
objec-tives of all mango breeding programmes. Their results also indicated that the
main factors for resistance of mangoes to fruit flies lie in the fruit peel and not in the
fruit pulp.

4.6 Methods of Breeding

Selection from open-pollinated seedlings

In India, almost all cultivars are selections that were made from naturally occur-
ring open-pollinated seedlings. All of the Florida cultivars were selected from
open-pollinated seedling progenies; none has come from a controlled breeding
programme. Among the 64 Florida cultivars evaluated in the parentage analysis by
Schnell et al. (2006), the genetic background was found to be based on as few as
four Indian cultivars and the polyembryonic cultivar ‘Turpentine’. Two Indian
cultivars, ‘Mulgoba’ and ‘Sandersha’, are in the background of most Florida types
with ‘Amini’, ‘Bombay’, ‘Cambodiana’, ‘Long’, ‘Julie’ and ‘Nam Doc Mai’
making lesser contributions. In the parentage analysis ‘Turpentine 10’ was iden-
tified as a most probable paternal parent for ‘Haden’. The polyembryonic seed-ling
races of Cuba and Florida were considered the same by Popenoe (1920) who called
them the West Indian race (commonly known as ‘Turpentine’ in Florida). ‘Haden’
was reported as the maternal parent for ten cultivars included in the analysis, but
based on the parentage analysis, 31 cultivars were found to have ‘Haden’ as one of
the most likely parents. Likewise, the other important early Florida selection
‘Brooks’ is the parent of seven cultivars. ‘Haden, ‘Brooks’ and seedlings of
‘Haden’ and ‘Brooks’ have contributed disproportionately to the Florida group. In
Florida, modern selection and breeding programmes for mango have focused on
cultivars with exceptional production, red skin, disease resistance and extended
shelf life. Methodology for crop improvement consists of collecting seeds from
selected maternal parents with desired characteristics and growing them in close
proximity to desirable male parents. Seedlings are screened by leaf aroma and
horticultural traits, leading to a field population of thousands of candidate seedlings
(Campbell and Zill, 2006).
Breeding and Genetics 79

In the Canary Islands, Spain, breeding mainly involves selection of open-


pollinated seedlings of ‘Lippens’. Lavi et al. (2004) indicated that mother trees
should not be chosen entirely on the basis of their phenotypes and trees with
inferior performance could also be included since progeny performance is quite
unpredictable. They observed that most of the variance components of the
agriculturally interesting traits are non-additive (Lavi et al., 1998) and most of
these traits result from dominant and epistatic interactions.
The Israel mango-breeding programme therefore relies on open-pollination
involving many mango cultivars from various parts of the world and screen-ing
approximately 100 seedlings from each mother tree. The seedlings are grown on
their own roots in the nursery for about a year and then planted in the field at
spacings of 2 × 6 m. Fruiting occurs after 3–6 years and first selec-tion is carried
out based on field and laboratory data. Fruit characteristics at this stage are good
skin colour, fruit weight of 400–600 g and high fruit qual-ity (good taste, absence
of fibres and small seed). Where a long harvest sea-son is desired both early and
late harvest seedlings are selected and a general idea about shelf life of these
seedlings is obtained. The second selection is carried out under commercial
conditions by several experienced farmers using grafted plants. Plants that
successfully pass this stage are planted in semi-commercial plots for a final
assessment before recommendation to farm-ers. The two selection stages are aimed
at shortening the breeding programme and minimizing both the false negatives
(loss of interesting seedlings which were not identified) and the false positives
(wrong identification of interest-ing seedlings which should actually be rejected).
New mango cultivars have also been selections made from open-pollinated
seedlings. ‘Maya’ and ‘Nim-rod’ are seedlings of the same mother tree
(Oppenheimer, 1967); ‘Tahar’ is a seedling of ‘Irwin’ (Slor and Gazit, 1982) and
‘Naomi’ is a seedling of ‘Palmer’ (Tomer et al., 1993). The other promising
selections include ‘Shelly’ (late sea-son), ‘Tango’ (early season), ‘Selection 20/1’
(large fruit with aborted seeds) and ‘Selection 1/5’ (shiny red colour).

The South African mango-breeding programme also places a major emphasis


on open-pollinated seedling selection. The screening of seedlings, besides
undergoing tests for various plant and fruit characters, also includes shipping
quality tests in which fruits are packed in 4 kg crates and stored at 11°C for 28 days
to simulate shipping conditions. After storage the fruits are allowed to ripen at
room temperature (25 ± 2°C) and screened thoroughly. Promising seedlings are
again field-tested with ‘Sabre’ as rootstock with a row of 40 trees along with ten
trees of commercial varieties. Two cultivars, ‘Joa’ (‘Palmer’ seedling) and ‘Chene’
(‘Kent’ seedling), were released from the breeding programme in 1996. Another
high yielding selection ‘A2-CD28’ (‘Fascell’ seedling) is a midseason clone with
an attractive pink blush (Human et al., 2006) and has been recommended for Plant
Breeders’ Rights in 2005. The other promising selections are ‘Osteen’ (‘Haden’
seedling) and ‘Neldica’ (‘Palmer’ seedling).

The 'R2E2' cultivar developed in Australia is a seedling progeny of the Florida


cultivar 'Kent', and is now the second most popular mango grown in Australia.
80 C.P.A. Iyer and R.J. Schnell

Controlled pollination

Hand pollination
The traditional, cumbersome method involving the continued pollination of flowers
on a panicle over several days when the flowers are open has now been replaced in
India with more efficient methods. The current method involves the pollination of a
limited number of flowers per panicle (maxi-mum of ten), utilizing a larger number
of panicles since it is very rare that a panicle bears more than one fruit to maturity.
Using this method, fruit set as high as 3.85% can be achieved compared to the
0.23–1.57% efficiency involv-ing other methods (Mukherjee et al., 1968; Singh et
al., 1980).

Caging
The enclosure of two desirable parents of synchronous flowering in a screen house
with pollinating insects provides a more practical method of hybridiza-tion. This
method has been used in Israel (Degani et al., 1993), Brazil (Pinto and Byrne,
1993) and South Africa (Cilliers et al., 1996). A standardized caging technique for
mango breeding was previously used in India following the discovery of self-
incompatibility in some of the most popular commercial culti-vars (Sharma and
Singh, 1970). This procedure involves the planting of self-in-compatible (female)
and male parents in specially prepared breeding plots, and are enclosed in an
insect-proof cage into which freshly reared houseflies, or any other suitable
pollinator, are introduced to effect cross-pollination (Sharma et al., 1972).

Polycross mating
A polycross is simply the use of a number of advanced selections or current
commercial clones planted in a design that maximizes the chance for cross-
pollination. The polycross design has been extensively used in sugarcane breeding
where small flower size and low numbers of seedlings per cross make controlled
pollinations difficult. At USDA Subtropical Horti-culture Research Station (SHRS)
in Miami the clones ‘Haden’, ‘Tommy Atkins’, ‘Kent’, ‘Keitt’ and ‘Nam Doc Mai’
have been used to produce new seedlings for selection. Five clones of each
genotype were planted, with at least one plant of each genotype next to all other
genotypes. Over 1000 seed-lings from known mother trees are planted as maternal
half-sib families. The pollen parent of superior selections is determined using
microsatellite markers.

4.7 Handling of Hybrid Populations and Selection


Criteria for initial selection

Primary selection from the hybrid progeny is based on: (i) precocity; (ii) fruit size
and shape; (iii) skin colour; (iv) fruit characteristics (high pulp to stone ratio and
freedom from fibre and physiological disorders); and (v) fruit qual-ity. Following
this preliminary evaluation, selected hybrids are retained for
Breeding and Genetics 81

further screening. It is important to graft the hybrids onto proper rootstocks as early
as possible, as grafted plants are precocious. At least ten grafted plants of each
selected hybrid are used in the final selection, which is based on yield, regularity in
bearing and response to diseases and pests, in addition to other desirable fruit
characters. At least 3 consecutive years’ performance data should be collected
before deciding on their suitability for release as new cultivars.

Pre-selection

Trees have a long juvenile phase, and the development of pre-selection meth-ods is
important for discarding inferior seedlings at a very early stage, obvi-ating the need
for maintaining a large number of seedlings for long periods. This can save time,
land and labour. Leaf flavour has been reported to be directly correlated with fruit
flavour (Majumder et al., 1972; Whiley et al., 1993). Emergence of new growth
flushes, simultaneously with fruiting or immediately after harvest, is indicative of
regular bearing (Sharma et al., 1972). A higher phloem to xylem ratio, associated
with dwarfing, has been used effectively as a pre-selection criterion. Genotypes in
which the ratio exceeds 1.0 are least vigorous, those with a ratio between 0.6 and
1.0 are of medium vigour and those with a ratio of less than 0.6 are most vigorous
(Kurian and Iyer, 1992). In addition, higher levels of phenolics in the apical bud is
associated with reduced vigour and dwarfing (Iyer, 1991). Although Majumder et
al. (1981) indicated that low stomatal density is an indicator of dwarfness this has
not been confirmed by other workers (Iyer, 1991). Regular bearing mango cultivars
have low polyphenol oxidase (PPO) activity (cate-cholase and cresolase) compared
to alternate bearers (Sharma, 2003). Sharma et al. (2000) observed that a strong
positive correlation existed between the incidence of floral malformation and both
enzyme activity (catecholase and cresolase) and phenolic content and speculated
that PPO activity can be used as a biochemical index for screening mango
germplasm against malforma-tion disease.

Potential for marker assisted selection (MAS)

More than 65 microsatellite markers have been developed for mango and these are
easily used to verify parentage using a software package such as CERVUS (Marshall
et al., 1998). When caging trees or using the polycross mat-ing design it is possible to
identify the male parent from a set of potential male parents. This has been useful in
cacao breeding where mistakes in pol-lination have lead to the estimation of unreliable
breeding values for parental clones. The development of linkage maps and
identification of quantitative trait loci (QTL) for productivity and quality traits has led
to a very successful MAS in cacao (Schnell et al., 2007). This could serve as a model
for future mango breeding and selection efforts.
82 C.P.A. Iyer and R.J. Schnell

Molecular markers

Molecular markers can be used for estimating genetic relationships among clones,
for parentage analysis and for the development of a saturated linkage map.
Isozymes were the first markers to be used for fingerprinting mango cultivars, to
determine self- versus cross-pollination and to estimate genetic relationships
(Degani et al., 1990; Knight and Schnell, 1994). RAPD markers were also used to
fingerprint cultivars and estimate genetic relationships in mango (Schnell et al.,
1995). A group of ‘Haden’ seedlings and a random group of seedlings were
evaluated using 11 RAPD primers. This study sup-ported the ‘Haden’ parentage of
‘Eldon’, ‘Lippens’, ‘Tommy Atkins’ and ‘Zill’; however, the parentage of ‘Glenn’
and ‘Osteen’ was questioned. Adato et al. (1995) used DNA fingerprinting (DFP)
to evaluate genetic relationships between 26 mango cultivars and 14 rootstocks.
They provided a pedigree that further confirmed the relationship between many of
the ‘Haden’ seed-lings. Lopez-Valenzuela et al. (1997) used RAPD markers to
estimate genetic diversity among 15 rootstock cultivars using 13 markers, and
identified a specific RAPD band associated only with the polyembryonic types.
Eiad-thong et al. (1999) utilized anchored simple sequence repeat markers to anal-
yse 22 mango cultivars; they were able to distinguish genotypes, but were unable to
find markers unique to either monoembryonic or polyembryonic types, or for the
Thai cultivars selected for green harvest (crispy mango) from the cultivars selected
for ripe fruit production. Kashkush et al. (2001) utilized amplified fragment length
polymorphisms (AFLP) to estimate genetic rela-tionships between 16 cultivars and
seven rootstock cultivars. They also anal-ysed 29 progeny from a cross of
‘Tommy-Atkins’ and ‘Keitt’ and produced a crude linkage map that identified 13
of the 20 linkage groups.

Viruel et al. (2005) developed the first reported set of 16 microsatellite


markers for mango, of which 14 produced the expected one or two amplifi-cation
products per genotype. These 14 microsatellites were used to evaluate 28 mango
genotypes that included 14 Florida cultivars. Discrimination of all 28 genotypes
was possible and the average number of alleles per locus was 5.3. Previously
known pedigree information for the ‘Haden’ family of man-goes was confirmed
and was in agreement with previously published RAPD and DFP analyses (Adato
et al., 1995; Schnell et al., 1995) with one exception. Viruel’s clone of ‘Zill’ was
not resolved as a seedling of ‘Haden’. Schnell et al. (2005) developed a second set
of 15 microsatellite markers and analysed 59 Florida cultivars and four related
species. Two of the microsatellites were monomorphic among the Florida cultivars;
the other 13 had an average num-ber of alleles per locus of 4.2 with polymorphism
information content (PIC) values varying from 0.21 to 0.63.

Schnell et al. (2006) used 25 microsatellite loci to estimate genetic diversity


among 203 unique mangoes (M. indica L.), two M. griffithii Hook. f. and three M.
odorata Griff. accessions maintained at the National Germplasm Repository
(NGR) and by Fairchild Tropical Botanic Garden (FTBG) in Miami, Florida. The
25 microsatellite loci had an average of 6.96 alleles per locus and an average PIC
value of 0.552 for the M. indica population. The total
Breeding and Genetics 83

propagation error in the collection (i.e. plants that had been incorrectly labelled or
grafted) was estimated to be 6.13%. When compared by origin, the Florida
cultivars were more closely related to Indian than to South-east Asian cultivars.
Unbiased gene diversity (Hnb) of 0.600 and 0.582 was found for Indian and South-
east Asian cultivars, respectively, and both were higher than H nb among Florida
cultivars (0.538). When compared by horticultural type, H nb was higher among the
polyembryonic types (0.596) than in the monoembryonic types (0.571).
To date 63 microsatellite markers have been developed for mango (Duval et
al., 2005; Honsho et al., 2005; Schnell et al., 2005; Viruel et al., 2005). This
number is more than adequate for genetic diversity studies and for par-entage
analysis as has been demonstrated by Schnell et al. (2006); however, it is not
enough to develop a saturated linkage map for the 20 linkage groups of mango.
Developing an additional 200 microsatellite or single nucleotide polymorphic
markers is a major objective of the USDA Agriculture Research Service (ARS)
programme in Miami over the next 2 years. Three experimen-tal populations have
been developed and planted in the field as mapping populations. The first
population is an F2 population derived from self-polli-nation of ‘Tommy Atkins’
consisting of 168 seedlings that was planted in the field in 1995. The second
population is an F2 population derived from self-pollination of ‘Haden’. A total of
224 seedlings from a single isolated ‘Haden’ tree have been in the field for 3 years.
Phenotypic data collection is in prog-ress for both of these populations. The
development of a saturated linkage map and the identification of QTL for
important traits are objectives for the USDA-ARS programme in Miami for the
next 5 years.

4.8 Minimizing Problems in Breeding

Heavy fruit drop

Heavy fruit drop ultimately results in few hybrid fruits, despite the large number of
flowers used for cross-pollination. While many recommendations are available to
minimize mango fruit drop with growth regulators, these have not been very useful
in breeding programmes where the number of flowers remaining in a panicle is
very low. Iyer and Subramanyam (1972) suggested that embryo culture could be
used to rescue hybrid embryos, and Sahijram et al. (2005) developed in-vitro
techniques to rescue immature mango embryos from controlled crosses and
recovered hybrid plants.

Long juvenile phase

Normally, mango seedlings require 3–10 years to flower, thereby prolonging the
breeding programme. Grafting individual hybrids on the proper rootstocks at the
earliest possible stage and growing them in a location where climatic stress
(particularly cold weather) prevails, induces precocious
84 C.P.A. Iyer and R.J. Schnell

flowering. Iyer (1991) has reported significant differences between seedlings on


their own roots and grafted plants of the same genotype with respect to fruit size,
quality and even colour in the early years. However, different results have been
reported in a recent study of the effect of rootstocks on the performance of seedling
scions with respect to ten horticultural traits (Lahav et al., 1995). No difference of
practical importance was found between the original seedlings and their grafted
duplicates.
Singh (1969) has suggested that young mango seedlings can be induced to
flower and fruit if they are grafted onto comparable shoots of a bearing tree (a few
days before flowering). The scions are defoliated and girdled. Using this technique,
it has been reported that the fruit characteristics of F 1 hybrids can be determined
within 2 years and F2 hybrids within 4 years, thus eliminating at least 10 years
from the period required to raise and evaluate F2 populations (Singh, 1963).
Ethephon (Chacko et al., 1974) and paclobutrazol (Anonymous, 1984) have
flower-inducing properties in mango and could also be utilized for shortening the
juvenile phase. However, they must be used with caution since chemical induction
of flowering can alter fruit size and this could lead to errors in judgement when
making selections within the hybrid progeny.

Polyembryony

Seeds of polyembryonic mango cultivars characteristically contain several nucellar


embryos, and may also contain a zygotic embryo. While nucellar seedlings are
preferred as rootstocks for mango because of their uniformity, the breeder, on the
other hand, is generally interested in sexual seedlings for the selection of improved
rootstocks. Until recently, crosses involving poly-embryonic cultivars as the
maternal parents were generally not performed, since reliable methods for
identifying zygotic seedlings were not available. The use of polymorphic enzyme
systems (isozymes) (Degani et al., 1990, 1992) to identify zygotic seedlings
(Schnell and Knight, 1992; Truscott, 1992; Degani et al., 1993) is based on the fact
that nucellar seedlings should have the same isozyme alleles as the maternal parent.
A variation at a locus coding for an enzyme indicates that the plant has originated
by sexual reproduction. Zygotic seedlings arising from self-pollination are
distinguished from nucel-lar seedlings by being homozygous at one or more loci at
which the female parent is heterozygous. Statistically, when three or four
heterozygous loci are examined, up to 88 or 94%, respectively, of the selfed
zygotic seedlings are identifiable (Moore and Castle, 1988). Cross-pollination by
another cultivar of the same genotype is equivalent to self-pollination. Zygotic
seedlings from cross-pollination are distinguishable from those resulting from self-
pollination if they express an allele not carried by the female parent.

The frequency of occurrence of zygotic seedlings varies among the poly-


embryonic mango cultivars, i.e. 22% with ‘13-1’ (Degani et al., 1993), 20 and 24%
with ‘Turpentine’ (Degani et al., 1993 and Schnell and Knight, 1992, respectively),
2 and 4% with ‘Sabre’ (Truscott, 1992 and Schnell and Knight,
Breeding and Genetics 85

1992, respectively) and 36 and 64% with ‘Madu’ and ‘Golek’, respectively
(Schnell and Knight, 1992).

4.9 Achievements of Conventional Breeding


Despite the many problems associated with mango breeding for cultivar devel-
opment, many useful hybrids have been released. The earliest attempts were
probably made in the West Indies to combine the good qualities of the Indian
mango with the indigenous types by controlled pollination (Brooks, 1912).

India

Intervarietal hybridization in India has resulted in the release of many culti-vars.


The work at Sabour initially yielded two promising hybrids: ‘Mahmood Bahar’ and
‘Probashanker’, both combinations of ‘Bombay’ and ‘Kalapady’ (Roy et al., 1956).
Subsequently, four more hybrids have been developed. These are: ‘Sundar Langra’
(‘Sardar Pasand’ × ‘Langra’) having ‘Langra’ quality and regular bearing habit;
‘Alfazli’ (‘Alphonso’ × ‘Fazli’) with ‘Fazli’ quality and early ripening; ‘Sabri’
(‘Gulabkhas’ × ‘Bombai’) having ‘Bombai’ fruit shape and colour of ‘Gulabkhas’
with regular bearing habit; and ‘Jawa-har’ (‘Gulabkhas’ × ‘Mahmood Bahar’)
having high pulp and early bearing habit (Hoda and Ramkumar, 1993). Developed
in Kodur, the hybrid ‘Swarna-jehangir’, combining the high quality of ‘Jehangir’
and the attractive colour of ‘Chinnaswarnarekha’, is a prolific bearer and is the best
of all hybrids devel-oped at this centre. The other hybrids released from Kodur are
‘Neeludin’ (‘Neelum’ × ‘Himayuddin’), ‘Neelgoa’ (‘Neelum’ × ‘Yerra Mulgoa’)
and ‘Neeleshan’ (‘Neelum’ × ‘Baneshan’).

Two excellent, regular-bearing hybrids, ‘Mallika’ and ‘Amrapali’, were


developed and released by the Indian Agricultural Research Institute (IARI), New
Delhi (Singh et al., 1972). ‘Mallika’ is a hybrid between ‘Neelum’ and ‘Dashehari’
with a high total soluble solids (TSS) content, a higher percentage of pulp, fibreless
flesh and a fruit size of about 300 g. ‘Amrapali’ (‘Dashehari’ × ‘Nee-lum’) is
precocious, distinctly dwarf and hence amenable to high-density planting, a regular
bearer with excellent quality and is also very rich in vitamin A. Recently, two more
cultivars, ‘Arunima’ (‘Amrapali’ × ‘Sensation’) and ‘Pusa Surya’ (a selection from
‘Eldon’) have been released from the IARI. A promising mango hybrid ‘Ambika’,
a cross between ‘Amrapali’ and ‘Janardhan Pasand’, having a yellow colour with
red blush, firm flesh and scanty fibre was released from the Central Institute of
Sub-Tropical Horticulture, Lucknow.
Four hybrid cultivars were released from the Indian Institute of Horticul-tural
Research in Bangalore: ‘Arka Aruna’ (‘Banganapalli’ × ‘Alphonso’), ‘Arka
Puneet’ (‘Alphonso’ × ‘Banganapalli’), ‘Arka Anmol’ (‘Alphonso’ × ‘Janardhan
Pasand’) and ‘Arka Neelkiran’ (‘Alphonso’ × ‘Neelum’). ‘Arka Aruna’ is dwarf,
and large fruited with a high percentage of pulp and high TSS content. It is ideal
for homesteads. ‘Arka Puneet’ is very similar to
86 C.P.A. Iyer and R.J. Schnell

‘Alphonso’ but free of ‘spongy tissue’, has a good shelf life and is not suscep-tible
to fruit fly attack. ‘Arka Anmol’ is a heavy bearer with good keeping quality (Iyer
and Subramanyam, 1993). ‘Arka Neelkiran’ is free of spongy tissue and has
excellent skin colour.
‘Ratna’ is a cross between ‘Alphonso’ and ‘Neelum’ that was carried out at the
Fruit Research Station, Vengurla, Maharashtra; it has a larger fruit size, fruit
quality similar to ‘Alphonso’ and is free of ‘spongy tissue’ (Salvi and Gunjate,
1988). A parthenocarpic mango cultivar, ‘Sindhu’, has been devel-oped at this
station as a result of back-crossing ‘Ratna’ with ‘Alphonso’ (Gun-jate and
Burondkar, 1993).
Two hybrid cultivars were released from the Fruit Research Station in
Sangareddy, Andhra Pradesh. ‘Au-Rumani’ (‘Rumani’ × ‘Mulgoa’) is a regular
and prolific bearer with fibreless flesh. ‘Manjira’ (‘Rumani’ × ‘Neelum’) is a
dwarf, regular and prolific bearer with good quality fruits.
The Paria Research Station in Gujarat developed three mango hybrids,
‘Neelphonso’ (‘Neelum’ × ‘Alphonso’), ‘Neeleshan Gujarat’ (‘Neelum’ × ‘Bane-
shan’) and ‘Neeleshwar’ (‘Neelum’ × ‘Dashehari’). These hybrids are supe-rior in
TSS, total sugars and vitamin C, in addition to their dwarfing habit, with respect to
their parents (Sachan et al., 1988).

Other countries

USA
Mango hybridization was reported from Hawaii in the 1920s, but no out-standing
problem appears to have been addressed or solved (Pope, 1929). A number of
crosses have been reported in Florida (Young and Ledin, 1954; Sturrock, 1969), but
all of the Florida cultivars are chance seedlings and none came from controlled
pollinations.

Israel
There is an extensive breeding programme in Israel aimed at producing higher
yielding cultivars with good quality, attractive fruit and with longer harvest
periods. Several hundred seedlings from open and controlled polli-nations have
been evaluated, and 14 of them have been identified as being of interest (Lavi et
al., 1993). The rootstock breeding programme is aimed at developing rootstocks
resistant to or tolerant of soil stresses, i.e. calcareous soils, saline irrigation water
and heavy non-aerated soils that predominate in the mango-growing regions of
Israel. Several interesting monoembryonic and polyembryonic rootstocks have
been selected (Lavi et al., 1993), but none has performed better than ‘13-1’, the
currently preferred rootstock in Israel (Gazit and Kadman, 1980).

Australia
A breeding programme to develop a new cultivar which retains the charac-teristic
flavour of ‘Kensington’, but with improved productivity, greater dis-ease
resistance, enhanced skin colour and better postharvest performance,
Breeding and Genetics 87

was initiated in Queensland, Australia. These features are found in many Florida
cultivars (i.e. ‘Irwin’, ‘Sensation’ and ‘Tommy Atkins’) which are being used as
maternal parents in crosses with ‘Kensington’ (Whiley et al., 1993). Promising
hybrids have been identified in crosses involving ‘Sensa-tion’, for example
‘Calypso’™ (see Knight et al., Chapter 3, this volume). ‘Calypso’™ has increased
shelf life, firmer fruit, extra blush for cosmetic appeal, a higher flesh-to-seed ratio
and consistent yields of high-quality fruit. The Australian mango breeding
programme was strengthened since 1994 by launching a major effort involving
various organizations located in different agro-climatic zones in hybrid production,
as well as regional testing.

Brazil
Breeding has been initiated in the tropical savannah of Brazil to develop cul-tivars
that are dwarf and with good quality fruit. Hybridizations have involved local,
Indian and Florida cultivars. ‘Amrapali’ and ‘Imperial’ were good male parents to
confer dwarfing in the progeny (Pinto and Byrne, 1993). Out of 2088 seedlings in
the field, 209 seedlings were selected in the first year and 42 of these were later
identified as promising, from which four have been released as new cultivars (Pinto
et al., 2004). These four are: ‘Alfa’ (‘Mal-lika’ × ‘Van Dyke’), which is semi-
dwarf, high yielding and regular bearing; ‘Beta’ (‘Amrapali’ × ‘Winter’), high
yielding and moderately resistant to anthracnose and Oidium; ‘Roxa’ (‘Amrapali’ ×
‘Tommy Atkins’), with excel-lent fruit quality; and ‘Lita’ (‘Amrapali’ × ‘Tommy
Atkins’), high yielding with excellent fruit quality.

South Africa
The South African breeding programme at the Citrus and Subtropical Fruit
Research Institute (CSFRI) is based on introductions, open-pollination and mass
selection. Four new cultivars have been released: ‘Heidi’, ‘Neldawn’, ‘Neldica’
and ‘Ceriese’. In addition, 12 promising selections have been iden-tified for further
evaluation (Marais, 1992).

4.10 Mutations

Somatic mutations

Asexual propagation enables the preservation of accumulated mutations (macro


and micro), which would normally be eliminated during sexual propogation. In
many fruit crops, bud mutations and chimeras occur rather frequently and can
provide an additional source of variability for selection. However, such reported
instances are relatively few in mango. Roy and Visweswariya (1951) observed
mutants of ‘Puthi’ in which the number of palisade cell layers differed from the
original cultivar. Naik (1948) observed significant variation among trees of the
same clone with respect to fruit shape, size, colour and quality, which was ascribed
to bud mutations. ‘Davis Haden’, a sport of ‘Haden’, is larger than ‘Haden’ and its
season of maturity is about
88 C.P.A. Iyer and R.J. Schnell

a month earlier (Young and Ledin, 1954). ‘Rosica’ from Peru, is a bud mutant of
‘Rosado de lca’. Unlike its parent, ‘Rosica’ is high yielding and regular bearing,
and does not produce seedless fruits (Medina, 1977).
Oppenheimer (1956), after a survey of many orchards in India, reported wide
variability in the performance of trees of the same clone within a single orchard.
Mukherjee et al. (1983) conducted a survey of mangoes in eastern India and
identified some superior clones. Singh and Chadha (1981), in a study of orchards
of ‘Dashehari’, located four clones which were superior in performance. Singh et
al. (1985) isolated two high-yielding clones from orchards of ‘Langra’. Within
‘Kensington’, strains have also been identified that show improved resistance to
bacterial black spot (Whiley et al., 1993).
Roy (1950) observed a mutant of ‘Alphonso’ with respect to fruit shape, and
suspected it to be a mericlinal chimera. Pandey (1998) has described seven clones
of ‘Alphonso’: ‘Alphonso Behat’ and ‘Alphonso Bihar’ from Bihar, ‘Alphonso
Batli’, ‘Alphonso Black’ and ‘Alphonso Bombay’ from Maharashtra, ‘Alphonso
Punjab’ from Punjab and ‘Alphonso White’ or ‘Bili Ishada’ from the North Canara
district of Karnataka. Rajput et al. (1996) assembled several ‘Dashehari’ variants
and after 14 years of observation, reported that the clone ‘Dashehari 51’ was
superior with respect to yield and regular bearing. Other somatic mutants include:
‘Cardozo Mankurad’ with large fruits of attractive colour and high yields from
‘Mankurad’ of Goa; dwarf selections from the ‘Rumani’ and ‘Bangalora’
(Ramaswamy, 1989); development of ‘Paiyur’, a dwarf selection from ‘Neelum’
(Vijaya Kumar et al., 1991); ‘Rati Bangana-palli’ and ‘Nuzuvid’ from
‘Banganapalli’ (Anonymous, 1999); and ‘MA-1’, regular bearing and high yielding
with resistance to ‘spongy tissue’ from ‘Alphonso’ (Mukunda, 2003).

In Thailand, Chaikiattiyos et al. (2000) selected clone ‘SKoo7’ (now known as


‘Kaew Sisaket’) from 320 ‘Kaew’ plants; ‘SKoo7’ has higher yield and superior
quality. Jintanawongse et al. (1999) also made superior selections for yield and
fruit quality from ‘Nam Dok Mai’, ‘Khiew Sawoey’, ‘Rad’ and ‘Nang Klang Wau’
and DNA fingerprints of all these clones were made for comparison with the
parental clone.
For these studies, it is important to conduct a replicated cultivar evalua-tion
trial against standard commercial cultivars to establish that these varia-tions are
stable and not due to environmental responses. The use of genetic markers should
be explored to confirm that the new clones are genetically distinct from the original
cultivar.

Induced mutations

Mutation induction using ionizing radiation was attempted by Siddiqui et al.


(1966). Siddiqui (1985) irradiated dormant buds of ‘Langra’ with high doses of J
rays, and grafted them onto 1-year-old seedlings. A bud graft exposed to 3.0 kR
bore fruits which were heavier, larger and had a more cream-yellow pulp than the
control. This variability was stable over three seasons. Sharma and Majumder
(1988b) irradiated bud sticks, topworked them onto 10-year-old
Breeding and Genetics 89

seedlings, and found that dosages above 5 kR are lethal for mango and that the
lethal dose required for 50% mortality (LD50) lies between 2 and 4 kR. Effective
dosages of the chemical mutagens, ethane methyl sulfonate (EMS) and N-nitroso
methyl urea (NMU), were 1.5 and 0.05%, respectively. The spectrum of mutations
induced by physical and chemical mutagens was observed to be more or less the
same, indicating the high sensitivity of certain loci. The mutants included
dwarfness, changes in shape and serration of leaves and in TSS content in
‘Dashehari’. As in other perennial crops, muta-genesis techniques that can allow
useful traits to be targeted, as well as isolating mutated sectors from a chimera, are
essential.

4.11 Breeding Potential of Wild Species


Bompard (1993; see Bompard, Chapter 2, this volume) has made a compre-hensive
study of the wild Mangifera species and enumerated their potential use in breeding.
Mangifera laurina, which has subglabrous and laxly flowered panicles and is well
adapted to areas with perpetual wet climates, is resistant to anthracnose. Mangifera
orophila from Malaysia and M. dongnaiensis from Vietnam are both restricted to
mountain forests 1000–1700 m above sea level and their hybrids with mango could
extend cultivation into temperate zones.
Mangifera magnifica is completely free of fibres; M. rufocostata and M.
swintonioides have an off-season bearing habit; M. pajang (endemic to Borneo)
and some strains of M. foetida have good quality fruits. Similarly ‘Wani’ from Bali
and Borneo, the best variety of M. caesia, has a distinctive taste. Mangifera casturi
from South Kalimantan is a prolific bearer with small, black, sweet fruits having
good potential. Mangifera altissima is reportedly resistant to mango pests, such as
hoppers, tip borers and seed borers (Angeles, 1991). Sharma and Choudhury
(1976) observed that wild Mangifera trees identified in Tripura State (north-eastern
India) were free of mango malformation. The wild species could also contribute to
higher productivity. Fairchild (1948) observed that crosses between five-stamened
mango and the Indian mango (only one fertile stamen) could produce hybrids
having better pollinating quality. The interspecific compatibility of these species
with M. indica must be verified before they can be utilized in hybridization
programmes (as sug-gested by Bompard, 1993; Kostermans and Bompard, 1993;
see Bompard, Chapter 2, this volume).

4.12 Conclusions
Until recently, all mango cultivars arose as chance seedlings or as seedling
selections from known mother trees. Enthusiasm for controlled hybridiza-tion by
means of hand pollination waned because of the tedious nature of the task and
heavy fruit drop, resulting in only very few hybrids. This low hybrid population
was inadequate for selection and hence not many outstanding hybrids were
obtained. However, improvements in pollinating techniques
90 C.P.A. Iyer and R.J. Schnell

and more rapid screening of hybrid populations have enabled the release of many
hybrid mango cultivars of commercial value. Because of the world market’s
demand for mangoes with specific qualities, the synthesis of new cultivars has
become imperative. Rapid strides in molecular biology and in other aspects of
biotechnology have opened up new approaches in plant breeding. The development
of polymerase chain reaction (PCR)-based genetic markers, specifically
microsatellites, and their application to classical breeding offer tremendous
potential for mango improvement. The develop-ment of a saturated linkage map
and the identification of QTL for important traits will allow the implementation of
a MAS programme. The introduction of specific genes for disease resistance from
cultivars and wild species into popular cultivars should soon be a reality. Without
resorting to these new technologies, mango breeding will continue to be a slow
process.

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5 Reproductive Physiology

T.L. Davenport
University of Florida, Florida, USA

5.1 Introduction 98
5.2 Phenology 99
5.3 Shoot Development 100
Vegetative shoots 102
Reproductive shoots 104
5.4 Flowering Mechanisms 105
Shoot initiation 105
Induction 106
Florigenic promoter (FP) or stimulus 108
Vegetative promoter (VP) 110
5.5 Environmental Influence on Vegetative and Reproductive Development 111
Temperature 111
Water relations 113
Effect of N on flowering 114
Photoperiod 116
5.6 Hormonal Influence on Flowering 116
Ethylene 116
Auxin 117
Cytokinins 118
Gibberellins 119
Plant growth retardants 121
5.7 Photoassimilate Influence on Flowering 123
5.8 Horticultural Manipulation of Flowering 123
5.9 Conceptual Flowering Models 124
Carbohydrate-regulated flowering models 124
Hormone-regulated flowering models 127
5.10 Floral Management 133
5.11 Floral Biology 134
Sex ratio 134
Environmental determinants of sex ratio 134
Physiological determinants of sex ratio 135
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 97
98 T.L. Davenport

Anthesis and dehiscence 136


Pollen 136
Pollination 137
5.12 Fruit Development 139
5.13 Stenospermocarpy 139
5.14 Fruit Set and Retention 140
Sex ratio 140
Mineral nutrients 141
Hormonal control 141
5.15 Alternate Bearing 145
5.16 Conclusions 145

5.1 Introduction
Flowering and fruit set are the most critical of all events occurring after estab-
lishment of a tree crop. Given favourable growth conditions, the timing and
intensity of flowering greatly determine when and how much fruit are pro-duced.
Many important details about flowering are becoming clearer, espe-cially in
herbaceous plants, at the physiological, biochemical and molecular levels (see
reviews by Searle, 1965; Zeevaart, 1976, 2006; Bernier et al., 1981, 1993; Halevy,
1985–1986; Bernier, 1988; Kinet, 1993; Boss et al., 2004; Komeda, 2004; Putterill
et al., 2004; Corbesier and Coupland, 2005).
Cool temperatures in the subtropics stimulate mango flowering and age of the
last vegetative flush has an important bearing on its ability to flower in marginally
cool or warm temperatures of the tropics (van der Meulen et al., 1971; Davenport,
2000, 2003). Consequently, mango flowering can be en-hanced during its normal
season or manipulated to occur at other times of the year in the tropics. For
example, potassium nitrate (KNO3) can stimulate out-of-season flowering in
mangoes in tropical latitudes (Barba, 1974; Núñez-Elisea, 1985; Davenport, 1993;
Protacio, 2000), although this treatment has not always been dependable. Various
aspects of mango flowering and/or fruit set have been reviewed (Singh, 1958a,
1979; L.B. Singh, 1960, 1977; Chacko, 1986, 1991; Chadha and Pal, 1986;
Davenport, 1993, 2000, 2003; Dav-enport and Núñez-Elisea, 1997; Singh et al.,
2005), and M.J. Soule (1950) pub-lished an extensive annotated bibliography of the
older literature related to mango reproduction.

Understanding mango flowering is essential to efficiently utilize man-agement


systems that extend the flowering and crop production seasons. Recent studies of
mango flowering have resulted in conceptual models that help explain the
physiological basis of flowering (Chacko, 1991; Cull, 1991; Kulkarni, 1991, 2004;
Whiley et al., 1991; Davenport and Núñez-Elisea, 1997; Davenport, 2000, 2003).
Control of flowering allows growers to harvest their crops at the most profitable
times. Increasing the season of availability improves competitiveness in the
international marketplace, and promotes the most efficient use of resources as costs
of inputs continue to rise.
This chapter addresses the physiology of mango flowering, early fruit set and
retention. Cultivar names and type of embryony (i.e. monoembryonic or
Reproductive Physiology 99

polyembryonic) are purposefully left out to focus on the physiological aspects of


reproduction regardless of whether they are tropically or subtrop-ically adapted or
from Indian or South-east Asian origin. Cultivars are selected for their productivity
in specific environments. Transfer of a cultivar to a different environment often
results in some alteration in performance. It is reasonable to assume that the
underlying mechanisms by which all culti-vars respond to their environment within
the framework of their genetic limi-tations are similar. The concepts described
herein, therefore, apply to all cultivars, regardless of origin.

5.2 Phenology
Growth of mango and other tropical trees is not continuous (Nakasone et al., 1955;
Halle et al., 1978; Verheij, 1986; Davenport, 1993, 2000, 2003). Apical buds spend
most of the time in rest. Growth occurs as intermittent, ephem-eral flushes of
shoots from apical or lateral buds (Naik and Mohan Rao, 1942; Singh, 1958a, b).
Stems are quiescent or resting terminal vegetative structures on branches from
which shoot growth occurs. Shoots are elongating vegeta-tive or reproductive
structures that emerge from apical or lateral buds of stems. Vegetative shoots
develop a prescribed number of nodes during growth before entering a resting state
as a stem. Depending on environment, periods of stem rest are generally short in
young plants but usually last sev-eral months between episodes of growth in
mature trees. Vegetative growth generally occurs up to three or four times a year
on individual branches, depending upon cultivar and growth conditions.

Development of the vegetative shoot from initiation of growth to full


elongation requires 3–6 weeks, depending on the cultivar and climatic condi-tions
(Whiley et al., 1991). During this period, 10–20 new leaves are generally produced
before returning to a resting state. These rhythmic episodes of extension growth are
recorded on each branch as segments consisting of compressed internodes
interspersed with long internodes, that is articulate growth (Tomlinson and Gill,
1973). Davenport (1992, 2003, 2006) referred to regions of compressed internodes
as intercalations and the entire segment of long internodes terminating in an
intercalation as an intercalary unit. The number of intercalations between each
branching point indicates the number of vegetative growth episodes or flushes that
have occurred between each flowering flush.

Flushes of vegetative growth occur on groups of stems borne on scaffold-ing


branches in isolated sections of tree canopy. Flushing stems are usually connected
at some common branch point within the tree limbs. Asynchro-nous flushes of
growth at various times in random portions of a tree canopy may appear to be
continuous growth but are simply flushes occurring in various parts of the total
canopy over time. Flowering flushes generally occur after extended periods of stem
rest in the low-latitude tropics or during cool winter months in the high-latitude
tropics and subtropics. Like vegetative flushes, reproductive flushes are usually
asynchronous in tropical climates
100 T.L. Davenport

(Verheij, 1986). In the subtropics, however, trees exposed to cold tempera-tures


(3–10°C) display synchronized flowering flushes throughout the tree canopy
approximately 1 month later. Subsequent vegetative flushes also tend to be
synchronous for one or two growth cycles depending upon the number of retained
fruit. Less intense, cool weather (10–18°C), however, results in asynchronous
reproductive flushes in responsive stems as is typical of trees growing in the
tropics. The timing of flowering flushes of cultivars in various locations has been
reviewed by L.B. Singh (1960), Chadha and Pal (1986) and Pandey (1989).
Variations in flowering patterns occur in all cultivars depend-ing on their age and
whether they are growing in dry or humid tropics or subtropics (L.B. Singh, 1960).

5.3 Shoot Development


Flushes of vegetative extension growth of mango stems terminate with for-mation
of determinate panicles. Several weeks to a few months after separa-tion of the last
flower or fruit from these panicles are required for the central axis of the panicle or
rachis to dry and mechanically separate from the sup-porting stem, depending on
the longevity of attached fruit. Five to ten lateral vegetative shoots typically
develop from axillary buds located at the termi-nal intercalation positioned in a
compact whorl surrounding the panicle scar of each stem (see Fig. 1 of Reece et
al., 1949). These lateral shoots become the branch points of stems. These
branching shoots form 10–15 leaves before the apical buds return to a resting state
to establish them as individual stems. Ini-tiation of these lateral vegetative shoots
may occur 2–3 months after desicca-tion of panicles which fail to set fruit. Fruit-
bearing stems do not initiate new lateral shoots until several months after
separation of fruit and rachis from the stem (Kulkarni and Rameshwar, 1989). Such
delayed vegetative growth can reduce the potential for new shoots to flower during
the next flowering sea-son (Singh and Khan, 1939; L.B. Singh, 1960, 1972;
Monselise and Gold-schmidt, 1982). The apical bud of stems is at rest for most of
the year in mature trees. Stems on centennial trees typically produce only one
vegetative flush during the year (N. Golez, personal communication, the
Philippines, 1989).
The apical resting bud of each newly established lateral stem (intercalary unit)
is surrounded by a compact whorl of 10–12 leaves with short inter-nodes
(intercalation) (Fig. 5.1). Protective bud scales are green but may be brown at the
tips due to desiccation (Sen and Mallik, 1941; Mustard and Lynch, 1946; Singh,
1958b; Ravishankar et al., 1979). Resting buds possess a number of pre-formed
nodes, each of which contains a leaf bract or leaf pri-mordium and a lateral
meristem (Fig. 5.2; see Figs 7–12 in Chaikiattiyos et al., 1994). The outermost,
proximally located dried leaf bracts (bud scales) pro-tect the more distal interior
leaf bracts, leaf primordia and lateral meristems from mechanical damage and
desiccation. Leaf bracts are vestigial non-developed leaves. Scales abort upon
evocation of new shoots. Proximally located bracts in apical buds fail to further
develop beyond some enlarge-ment and also abort with elongation of shoots.
Reproductive Physiology 101

Fig. 5.1. Apical bud of resting mango stem.

Lateral meristematic
primordia Apical dome
(includes meristem)

Leaf primordia (bracts)

Fig. 5.2. Stylized cross-section of apical bud showing positions of apical


meristem, lateral meristems and leaf primordia.

If apical buds are initiated during vegetatively inductive conditions, bracts


develop as small leaves and the leaf primordia develop as the full-sized leaves of
vegetative shoots. Additional leaves result from nodes formed by renewed activity
of the apical meristem. The number of leaves (nodes) is dependent upon the mean
temperature during initiation, and increases as temperatures rise (Whiley et al.,
1989). The lateral meristems of the apical bud develop as axillary buds at the base
of petioles in the elongating vegeta-tive shoot, each bearing protective bracts, leaf
primordia and lateral mer-istems (Fig. 5.3; see Fig. 1 of Reece et al., 1949).

In contrast, if shoot growth is initiated under floral inductive conditions, the


leaf bracts and primordia fail to fully develop, but the lateral meristems
102 T.L. Davenport

Stem

Fig. 5.3. Axillary bud of resting mango stem. Leaf petioles (arrows).

begin to elongate and branch at each node forming secondary, tertiary and
quaternary lateral meristems. Each branch point in the lateral inflorescence from
the panicle axis to the floral pedicels bears a floral bract (i.e. partially developed
vestigial leaf primordium) (Fig. 5.4). The distal half of the panicle structure is
derived from newly formed nodes laid down by cell divisions in the apical
meristem prior to returning to a resting state. Mixed shoots, bear-ing both leaves
and inflorescences at each node, result from development of both the primary leaf
primordia and the lateral meristems, which form the inflorescences in the same
nodes as leaves.
Vegetative shoot induction, thus, involves stimulating development of leaf
primordia from resting buds while repressing development of lateral meristems.
Leaf primordia then follow a predetermined cascade of genetic signals resulting in
leaf development at each node. Because all shoots emerge from resting buds, a
vegetatively induced event does not involve simply inhibition of flowering. The
putative inductive signal directing differentia-tion of leaf primordia onto leaves
upon initiation is termed a vegetative pro-moter (VP) rather than a floral inhibitor.

Shoots bearing only inflorescences (generative shoots) result from induc-tive


development of lateral meristems and suppression of leaf primordial development.
A predetermined cascade of flowering gene signals is activated in lateral meristems
resulting in lateral cymose inflorescences terminating with flowers. A distinct
florigenic promoter (FP) may be responsible for spe-cific activation of the lateral
meristems of mango. Mixed shoot induction results in combined development of
leaf primordia and lateral meristems.

Vegetative shoots

Vegetative shoots bear only leaves (Fig. 5.5). The anatomy of mango vege-tative
shoot development has been described (Singh, 1958b; Chaikiattiyos
Reproductive Physiology 103



º
4° 1°

1° Bract

Axis


Pedicel

1° Bract
1° 3° Bract
Pedicel

2° Bract

Pedicel 1°
1° Bract 1° 2° Bract Pedicel
Axis Pedicel

Fig. 5.4. Diagram and photos of mango inflorescence depicting the panicle axis
and primary (1°), secondary (2°) and succeeding levels of pedicel and cymose
floral archi-tecture. Vestigial leaf promorida (floral bracts) are depicted at the base
of each level of pedicel architecture.

et al., 1994). Vegetative shoots may arise either from axillary buds, if no apical bud
exists due to flowering in the previous flush, or from the apical bud when present.
The latter is considered extension growth or addition of an intercalary unit on the
existing stem, but the developmental events during shoot formation from either
apical or lateral buds are basically the same. Cells in the leaf primordia of initiating
buds begin to form individual leaves in the proximal portion of the vegetative
shoot. Soon thereafter, the apical meristem activates to form more nodes bearing
leaf primordia and lateral meristems. These newly formed leaf primordia develop
as the distal portion of the vegetative shoot if environmental conditions remain
vegetatively inductive (Núñez-Elisea et al., 1996). Newly elongating vegetative
shoots are green in most cultivars but may be bronze or red in others. Fully
expanded
104 T.L. Davenport

VEGETATIVE GENERATIVE MIXED CHIMERIC V/F TRANSITION F/V TRANSITION

GENERATIVE CHIMERIC V/F TRANSITION VEGETATIVE MIXED F/V TRANSITION

Fig. 5.5. Stylized diagrams and photomontage of shoot types found in mango. Transition
shoots shift from vegetative to floral (V/F) or floral to vegetative (F/V). Arrow ( )
represents individual leaves; floral diagram ( ) represents lateral inflorescences.

leaves are a shade of red, depending upon cultivar and cultural conditions and are
thin and limp from lack of lignification. The apical buds of vegetative shoots
generally become quiescent before completion of the limp, red-leaf stage (Núñez-
Elisea and Davenport, 1995). Internodes are compressed at the apex, and leaf
development is arrested thereby forming a bud with protec-tive outer scales, inner
leaf primordia, lateral meristems and the apical mer-istem. Fully expanded leaves
become light green and stiff as they become lignified and suberized. Vegetative
shoots are mature when leaves become dark green, which occurs when they are c.2
or 3 months old.

Reproductive shoots

Two types of reproductive shoots typically occur in mango. Generative shoots


display only flowers and have floral bracts or non-developed leaves at the base of
each lateral inflorescence (Fig. 5.5). Terminal inflorescences, i.e. panicles or
thyrsoids (Weberling, 1989), develop from dormant apical buds. The anatomy of
panicle development has been described (Juliano and Cuevas, 1932; Musahib-ud-
din, 1946; Mustard and Lynch, 1946; Singh,
Reproductive Physiology 105

1958b; L.B. Singh, 1960; Sturrock, 1966; Ravishankar et al., 1979; Scholefield,
1982; Scholefield et al., 1986). The complexes of primary to quaternary branch-ing
lateral structures of the inflorescence each terminate with three cymose flowers.
The terminal flower opens first, followed by two subtending lateral flowers. These
complexes form the lateral inflorescence structures emerging from the central axis
of the panicle. The central axis extension also terminates in a similar fashion.
Morphological stages of floral buds and panicle develop-ment were described by
Shu (1981) and Oosthuyse (1991a). Reece et al. (1949) described the development
of inflorescences initiated in lateral buds when the terminal bud is missing. There
are more nodes in dormant apical buds and their bracts are more developed than in
axillary buds; however, floral evocation is indistinguishable.

Generative shoot development in apical buds initially involves swelling of the


lateral meristems and their bud scales. Each axillary meristem devel-ops as an
inflorescence on a primary peduncle. The apical meristem then forms new lateral
meristems and leaf primordia for the distal portion of pan-icle development if floral
inductive conditions persist (Núñez-Elisea et al., 1996). Panicles may be open or
compact, depending upon internode elonga-tion, which is cultivar dependent (L.B.
Singh, 1960), but the architecture gen-erally conforms to that in Fig. 5.5. Mixed
shoots develop under weak floral inductive conditions (i.e. in the low-latitude
tropics). Both leaves and pri-mary pedunculate inflorescences develop from the
same nodes (Fig. 5.5). Leaf primordia and lateral meristems develop as leaf and
floral structures, respectively.

5.4 Flowering Mechanisms


Mango stems undergo varying periods of rest between episodes of growth,
depending on tree age and environmental influences. Resting mango buds must,
therefore, respond to two distinctly different signals for shoots to occur. The first
signal initiates growth of the shoot and the second determines if it will be
vegetative or reproductive. The signals that regulate initiation of shoot growth in
resting buds differ from the inductive signals that regulate shoot type.

Shoot initiation

Initiation is the onset of shoot development, regardless of the type of shoot evoked.
It involves cell division and elongation of cells in leaf primordia (vegetative
shoots), lateral meristems (generative shoots) or both (mixed shoots) in the nodes
of the resting buds, and is followed by cell divisions in the apical meristem to form
more nodes. Shoot initiation is stimulated by pruning, defoliation and irrigation
during dry conditions, or transition from the dry to rainy season in the tropics.
Application of nitrogen (N)-containing fertilizers, exposure to ethylene, or a shift
from cool to warm temperatures
106 T.L. Davenport

also stimulates shoot initiation. Reece et al. (1946, 1949), Mustard and Lynch
(1946), Núñez-Elisea and Davenport (1992b), Núñez-Elisea et al. (1996) and
Davenport et al. (2006a) observed that the vegetative or reproductive fate of mango
buds remains undetermined until after shoot growth is initiated. Reece et al. (1949)
proposed that a putative signal that triggers initiation of shoot development is
separate and different from the inductive signal, which determines the fate of the
shoot. Removal of apical buds by pruning stimu-lates initiation of axillary shoots
(Singh and Singh, 1956; Núñez-Elisea and Davenport, 1992b; Núñez-Elisea et al.,
1996; Davenport et al., 2006a). Defolia-tion of the apical whorl of five to ten
leaves also stimulates shoot initiation in dormant apical buds (Núñez-Elisea et al.,
1991; Núñez-Elisea and Daven-port, 1995). The fate of shoots that emerge in
response to these initiation stim-uli, however, is determined by other factors that
are prevalent at the time of initiation. Tip pruning, for example, during warm
summer months results in initiation of vegetative shoots from axillary buds,
whereas pruning during cool winter months usually results in initiation of axillary
inflorescences.

Induction

Induction in mango is the temporary commitment of buds to evoke a par-ticular


developmental pathway (i.e. vegetative shoot, generative shoot or mixed shoot)
when growth is initiated. Initiation of herbaceous plant flower-ing refers to the
onset of floral bud growth in actively growing vegetative shoots after the floral
inductive event (Bernier et al., 1981, 1993; Halevy, 1985– 1986; Bernier, 1988;
Huala and Sussex, 1993; Kinet, 1993). The inductive sig-nal is formed in leaves,
but the responsive buds are in continuous vegetative growth at the time of floral
induction in herbaceous plants and floral initia-tion follows; whereas mango buds
are in rest. Although the mango bud must be initiated to grow, that growth is
induced according to forces already present.

Whereas the floral inductive signal in mango may be present prior to bud
initiation, it must be present at the time of initiation for flowering to occur
(Kulkarni, 1988a; Núñez-Elisea and Davenport, 1995; Núñez-Elisea et al., 1996;
Davenport and Núñez-Elisea, 1997; Davenport et al., 2006a). The induc-tive signal
can be shifted from floral (F) to vegetative (V) or vegetative to floral, forming F/V
or V/F transition shoots, by altering temperatures dur-ing early shoot development
(Batten and McConchie, 1995; Núñez-Elisea et al., 1996) (Fig. 5.5). This shift in
morphogenic responses during shoot devel-opment demonstrates the plasticity and
temporal nature of induction, indi-cating that cells of the apical meristem do not
become irreversibly determined under inductive conditions. These results
demonstrate that, rather than being irreversibly committed to a vegetative or
reproductive fate at the onset of shoot initiation, the mango apical meristem
provides progenitor cells, some of which differentiate into specific target cells at
each node in the apex. The apical meristem, therefore, may not be directly involved
in the flowering process.
Reproductive Physiology 107

Target cells within leaf primordia and lateral meristems are competent to
respond to inductive signals; for example when initiated to grow under veg-
etatively inductive conditions, individual leaf primordia develop as leaves and
subtending lateral meristems associated with each developing leaf develop as
dormant axillary buds with protective bracts. These axillary buds may develop in
subsequent flushes as vegetative shoots when initiated in vegetatively inductive
conditions or as axillary inflorescences under floral inductive conditions. Under
strongly floral-inductive conditions, leaf pri-mordia fail to develop beyond the
bract stage, become dormant, and lateral meristems develop. Each lateral meristem
forms nodes consisting of leaf pri-mordia and meristems that are influenced by the
putative floral-inductive stimulus, which suppresses development of newly formed
leaf primordia. Subsequently formed meristems form pedunculate structures that
terminate in cymose inflorescences borne on each tertiary peduncle (Fig. 5.4).
Forma-tion of the primary, secondary, tertiary and quaternary peduncles, as well as
pedicels of inflorescences are always accompanied by a subtending, aborted bract
or vestigial leaf at each node (Fig. 5.4). Such development is attributed to a
sequence of gene expression (Coen et al., 1990; Coen and Meyerowitz, 1991;
Weigel et al., 1992; Coen and Carpenter, 1993; Lumsden, 1993; Yanofsky, 1995).
Shoot initiation during weakly floral-inductive conditions activates growth of leaf
primordia to develop leaves and the lateral meristems to pro-duce peduncles
bearing lateral inflorescences in each node of mixed shoots. The bases of each
pedicel branch within each lateral inflorescence also bear a vestigial leaf.

Upon termination of cell divisions in the apical meristem at the end of a


flushing period, no more nodes are formed. The apical bud of vegetative shoots
becomes quiescent, and the resting leaf primordia, bracts and lateral meristems are
poised to resume growth at a later date. When reproductive or mixed shoots
become quiescent, the lateral meristems ultimately develop determinant cymose
inflorescences. The most distally located meristem is possibly the determinant
extension of the central axis forming the terminal cymose floral group.

Chimeric shoots (Fig. 5.5) can occur in mango trees when shoot initiation
occurs during floral inductive conditions. They display inflorescences on one side
of the longitudinally bisected shoot and leaves on the other. The shoot axis is red
on the floral side of red fruiting cultivars (typical of panicles) and green on the
vegetative side (typical of vegetative shoots). This difference in the two sides
extends to the apical bud, which bears an undeveloped inflorescence on the floral
side and leaf bracts on the vegetative side. The explanation for this spatial
differentiation is that target nodes on each side of the apical bud respond to the
different inductive signals at the same time. The apical meristem is not implicated
except to form more nodes for the lat-eral inductive responses on each side in the
second portion of growth. Differ-ences in inductive signals on each side of an
existing shoot probably cause the differential response. This phenomenon indicates
that the fate of nodes on each side of the shoot cannot be attributed to a single
mother cell in the apical meristem. The inductive response must involve cells
formed in later
108 T.L. Davenport

cell divisions and would be determined by their location within nodes of the bud.

Florigenic promoter (FP) or stimulus

Early flowering work provided evidence for the presence of a graft transmis-sible
floral stimulus (i.e. florigen) that was induced in leaves and was trans-located to
buds to stimulate floral development (Chailakhyan, 1936; Zeevaart and Boyer,
1987). Florigen was functionally conserved across plant species (Lang, 1965, 1984;
Zeevaart, 1976; Lang et al., 1977). Floral induction in most plants involves sensing
of some environmental cue (i.e. daylength, water stress or vernalizing temperature)
in some organ (e.g. leaves). A putative flo-ral stimulus or alteration in the ratio of
florigenic to anti-florigenic compo-nents may be translocated to target cells in
meristems (Bernier et al., 1981). Photoassimilate movement from leaves in phloem
facilitates its transport to buds where it can interact to initiate flowering (King and
Zeevaart, 1973). Until recently, a floral stimulus could not be identified.
Alternative hypoth-eses were proposed that nutrient diversion to the meristems
could be involved (Sachs and Hackett, 1983) or that floral induction might be con-
trolled by multiple factors, including the putative floral stimulus, photoas-similates
and phytohormones (Bernier et al., 1993).

Molecular biology of flowering in the facultative, long-day, model plant,


Arabidopsis thaliana (reviewed in Zeevaart, 2006 and Aksenova et al., 2006), has
provided insight into the nature of the floral stimulus (FP). A network of four
interacting genetic signalling pathways may result in flowering in response to
photoperiodic, vernalization, gibberellin and autonomous envi-ronmental cues
(Perilleux et al., 1994; Mouradov et al., 2002; Perilleux and Bernier, 2002; Boss et
al., 2004; Komeda, 2004; Putterill et al., 2004; Corbesier and Coupland, 2005).
The photoperiodic pathway involves activation of the CONSTANS (CO) gene that
encodes a zinc-finger protein, which in turn induces expression of the
FLOWERING LOCUS T (FT) gene in the phloem tissue of leaves. FT is the
terminal, integrating gene of the four path-ways regulating flowering in
Arabidopsis. Its transcribed mRNA was initially thought to be the FP that is
transported in phloem to buds (Huang et al., 2005); however, evidence indicates
that the translated protein product of FT is translocated to Arabidopsis buds
(Corbesier et al., 2007). Analogous proteins encoded by Hd3a, an ortholog of FT
in rice (Tamaki et al., 2007), and the aspen ortholog, PtFT1, which along with CO
regulates the timing of flowering and growth cessation of Populus trichocarpa
(Bohlenius et al., 2006), appear to be the FP. In the buds, the protein product of FT
is thought to combine with the bZIP transcription factor (FD) protein to activate
transcription of floral iden-tity genes (i.e. APETALA1) to begin floral expression
(Abe et al., 2005; Wigge et al., 2005). Similar mechanisms are likely to exist in
mango.
Zhang et al. (2005) and Davenport et al. (2006b) isolated a CONSTANS-like
gene (MiCOL) from mango leaf DNA. CO is a circadian expression gene
interacting with the photoperiodic pathway in Arabidopsis (Putterill et al.,
Reproductive Physiology 109

2004), and is central to activation of the FT gene in Arabidopsis during long days.
Its role in mango flowering is unclear. The mango ortholog has 79%, 76% and 62%
homology with two apple CO genes, MdCOL2 and MdCOL1, and the Arabidopsis
CO gene (AtCO), respectively. Isolation of the FT or homologous gene responsible
for synthesis of the FP has been unsuccessful.
Studies with mango indicate that a FP is synthesized in leaves during exposure
to cool, floral-inductive temperatures and moves to buds to induce flowering
(Reece et al., 1946, 1949; Singh and Singh, 1956; L.B. Singh, 1959, 1962, 1977;
R.N. Singh, 1961; Sen et al., 1972; Núñez-Elisea and Davenport, 1989, 1992b;
Davenport and Núñez-Elisea, 1990; Davenport et al., 1995, 2006a). Unlike
receptor sites in buds of Thlaspi arvense (Metzger, 1988) and other plants requiring
vernalization for floral induction (Zeevaart, 1976; Bernier et al., 1981), mango
leaves appear to be where the putative floral stim-ulus is produced. Complete
defoliation of girdled branches during inductive conditions results in vegetative
shoots instead of generative shoots (Reece et al., 1949; Sen et al., 1972; Núñez-
Elisea and Davenport, 1989, 1992b; Núñez-Elisea et al., 1996; Davenport et al.,
2006a). It appears to be transported over long distances from leafy branches to
defoliated branches (Sen et al., 1972; Núñez-Elisea et al., 1996).

The putative, temperature-regulated FP is short-lived in situ (Núñez-Elisea and


Davenport, 1989, 1992b; Davenport et al., 1995; Núñez-Elisea et al., 1996).
Leafless cuttings from trees during cool, floral inductive conditions produce
inflorescences when stimulated to grow within 7 days of transfer to warm, non-
inductive conditions; the influence of the removed leaves lasts for 13 days when
cuttings are stored at cool temperatures (Davenport et al., 2001a). The same
cuttings produce only vegetative shoots in both storage conditions after the initial
loss of reproductive shoot production. There are more leaves on mango stems than
are necessary for floral induction in cool temperatures. Stems bearing as little as
one-quarter of a cross-sectioned leaf induce 95% generative shoots (Davenport et
al., 2006a); the remaining shoots are vegetative. Half of a leaf or more resulted in
100% generative shoots. Thus, the limiting amount of leaf necessary for floral
induction is less than a quarter of a leaf per stem. Davenport et al. (2006a)
demonstrated the quanti-tative movement of mango FP from half to five leaves on
a donor stem to five leafless receiver stems located as far as 100 cm from the donor
stem in isolated branches during exposure to cool, floral inductive temperatures.
The FP moves with photoassimilates in phloem from donor leaves to buds in the
receiver stems.

The mango floral stimulus is graft transmissible (L.B. Singh, 1959, 1962;
Kulkarni, 1986, 1988b, 1991). Flowering of seedling stems is stimulated by
grafting onto mature trees or by grafting mature stems onto juvenile plants (L.B.
Singh, 1959, 1962). Some mango cultivars selected in the tropics can flower at
higher temperatures than others and are not restricted to winter flowering
(Kulkarni, 1991). Transfer of the FP from tropical to subtropical selec-tions was
accomplished using reciprocal grafts between the two cultivar types (Kulkarni,
1986, 1988b, 1991). Subtropical cultivars that seldom flower in warm temperatures
flower in the ‘off’ season using these techniques. Three conditions
110 T.L. Davenport

were essential for summer flowering to occur in the low-temperature-requiring


cultivars (receptors) when grafted to the summer flowering type (donors): (i) the
summer-flowering donor cultivar stocks or scions were in a flowering cycle; (ii)
buds on the receptor scions or stocks of grafted plants had initi-ated shoot growth
during this cycle; and (iii) receptor stocks or scions had been completely defoliated
for transfer and/or expression of the floral stimu-lus. The presence of any leaves on
the receptor plants resulted in vegetative shoots.

Girdling experiments to isolate treated mango branches from the rest of the
tree suggest that the FP is translocated via phloem to apical buds (King and
Zeevaart, 1973; Bernier et al., 1981; Núñez-Elisea and Davenport, 1989, 1992b;
Núñez-Elisea et al., 1996; Davenport et al., 2006a). Shading experiments to reduce
photosynthate loading into the phloem also support this (Kulkarni, 1991). Reduced
flowering responses were observed in isolated leafy branches that were provided
with 90% and complete shading, which stopped photosyn-thate production entirely,
mimicked defoliation during cool, floral inductive conditions, resulting in a
vegetative growth response (R. Núñez-Elisea, T.L. Davenport and B. Schaffer,
Florida, 1991, unpublished results).

Vegetative promoter (VP)

An independently regulated VP probably contributes to induction of vegeta-tive


shoots as opposed to a floral inhibitor or expression of a default vegeta-tive status
in the absence of sufficient FP at the time of shoot initiation. Grafting studies (L.B.
Singh, 1959, 1962; Kulkarni, 1986, 1988b, 1991, 2004) demonstrated that complete
removal of leaves from receptor stems is required to express flowering of those
receptors when they are grafted to flowering donor stems. Kulkarni (1986, 1988b,
1991, 2004) considered that a putative floral inhibitor in leaves of the non-induced
receptor stems might antagonize the influence of the floral stimulus from donor
leaves. Others have noted a relationship between leaf age and the ability of shoots
to be reproductive (Singh et al., 1962a; Scholefield et al., 1986). KNO3-stimulated
early flowering in the tropics is successful only on stems that are at least 4
(Davenport, 2003) to 7 months old (Astudillo and Bondad, 1978; Bondad and
Apostol, 1979; Núñez-Elisea, 1985). Young stems often produce vegetative shoots
when ini-tiated under conditions that are floral inductive for more mature stems
(Núñez-Elisea and Davenport, 1995; Davenport, 2003). The putative VP appears to
be most active in leaves of young stems and slowly dissipates over time to allow
expression of the FP when shoots are initiated to grow in warm conditions.

The VP may be a gibberellin or closely associated with the gibberellin


synthesis pathway as indicated by enhanced flowering responses of trees to plant
growth retardants. Mangoes growing in wet and humid, low-latitude tropics tend to
produce frequent vegetative flushes and flower sporadically, perhaps due to higher
levels of the VP in the young stems combined with low levels of the putative FP
when shoot initiation occurs. Paclobutrazol
Reproductive Physiology 111

(PBZ) reduces the time in rest necessary to allow floral induction during warm
temperature conditions by c.1 month (Davenport, 2003), thus increas-ing the
potential to produce reproductive shoots in younger stems when ini-tiated to grow.
PBZ and uniconazole, triazole compounds that inhibit kaurene oxidase in the
gibberellin-synthesis pathway (Dalziel and Lawrence, 1984; Rademacher, 1991),
stimulate production of flowering shoots during weakly inductive conditions
(Burondkar and Gunjate, 1991, 1993; Tongumpai et al., 1991a; Voon et al., 1991;
Nartvaranant et al., 2000; Yeshitela et al., 2004a). Application of PBZ to mango
trees bearing 1-month-old stems produced inflorescences when bud break was
initiated 3 months later by foliar applica-tion of KNO3 (Davenport, 2003).

Vegetative or reproductive induction at the time of shoot initiation is governed


by the ratio of the putative floral promotive to inhibitory compo-nents (Lang et al.,
1977; Lang, 1984; Kulkarni, 1988a; see Bernier et al., 1981 for additional
references). The mango floral inhibitor should be viewed as an age-dependent VP.
The presence of an age-regulated VP in mango leaves, which moves with the
temperature-regulated FP and photoassimilates in phloem, may explain the
induction of specific receptors by this promoter in targeted leaf primordia to cause
development of leaves in vegetative or mixed shoots. A gradual decrease in the
level or influence of the VP may cause vegetative shoots to develop when initiation
occurs on 2-month-old stems, and generative or mixed shoots when initiation
occurs in stems from 4- to 7-month-old stems, given the constantly warm daily
temperatures maintaining a low level of FP in both situations.

5.5 Environmental Influence on Vegetative and


Reproductive Development
The effects of temperature and water relations on determinating vegetative and
reproductive growth of mango have been addressed (Davenport and Núñez-Elisea,
1997; Davenport, 2000; Kulkarni, 2004; Bangerth, 2006). This section focuses on
the impacts of temperature, plant water relations, mineral nutrition and photoperiod
on shoot initiation and induction.

Temperature

The developmental fate of mango buds is strongly influenced by tempera-ture


(Davenport and Núñez-Elisea, 1997). Cool night temperatures < 15°C in
combination with day temperatures < 20°C typically induce flowering if shoot
initiation occurs when plants are exposed to these conditions (Ou, 1980, 1982;
Wolstenholme and Mullins, 1982a, b; Shu and Sheen, 1987; Whi-ley et al., 1988,
1989, 1991; Núñez-Elisea et al., 1993; Núñez-Elisea, 1994; Núñez-Elisea and
Davenport, 1994a, b). The physiological and molecular basis for temperature
perception in leaves with respect to floral induction is not understood (Samach and
Wigge, 2005). Whiley et al. (1988, 1989, 1991)
112 T.L. Davenport

described the vegetative growth and flowering responses of several mo-


noembryonic and polyembryonic cultivars to four temperature regimes rang-ing
from vegetatively inductive (30C day/25C night) to floral inductive (15C
day/10C night). The effect of temperature on marcotted, container-grown plants
that were tip pruned or defoliated in order to stimulate shoot initia-tion was also
studied (Davenport, 1987; Núñez-Elisea et al., 1991, 1993, 1996; Núñez-Elisea and
Davenport, 1994b). Mango trees develop vegetative shoots when shoot initiation
occurs in warm temperatures (30C day/25C night), whereas inflorescences
develop when shoots initiate growth in cool tempera-ture conditions (18C
day/10C night; or 15C day/10C night) (Whiley et al. 1989; Núñez-Elisea and
Davenport, 1991b, 1995; Núñez-Elisea et al., 1993, 1996; Batten and McConchie,
1995). Bangerth et al. (2004) reported changes in the major phytohormones in
stems of containerized mango trees during exposure to cool, floral inductive
temperatures. The minimum leaf age and time of exposure to a low temperature
regime (18C day/10C night) required by stems for floral induction was examined
(Núñez-Elisea and Davenport, 1995). Leaves are competent to respond to cool
temperatures at 7 weeks, forming a small percentage of generative shoots. As they
age, higher propor-tions of generative shoots are induced and warmer temperatures
can stimu-late floral induction. The response to temperature is moderated by age of
the previous flush. Stems that are 4–5 months beyond the limp, red-leaf stage of
development will be induced to form generative shoots if initiated to grow at 25–
30C (Davenport, 2003).

Whiley et al. (1988, 1989, 1991) observed that at least 17 weeks are required
for initiation of reproductive shoots on non-clipped stems of trees maintained at
15C day/10C night. In similar experiments with different cultivars with-out
previous clipping of distal leaves to stimulate initiation, inflorescences were
observed after 5 weeks at 15C day/10C night (Chaikiattiyos et al., 1994).
Although inductive conditions were present in each of these studies, shoot
initiation was delayed by the presence of distal leaves. The earlier ini-tiation of
inflorescence development in tip-pruned or tip-defoliated stems compared to intact
ones demonstrates that the floral stimulus may be pres-ent, but the buds are not
induced until initiation occurs. It demonstrates the importance of stimulating
initiation of stems by tip defoliation or pruning at the onset of incubation in
controlled environment conditions so that the inductive response can be observed
within a reasonable length of time. The variable delays in shoot initiation in these
studies occurred because the experimental protocols depended on the plants’
internal initiation cycle to initiate shoots. This cycle slows down when plants are
exposed to lower tem-peratures (Whiley et al., 1988, 1989, 1991).

Floral or vegetative induction occurs when shoots are initiated. Resting buds
of plants that are exposed to cool temperatures (18C day/10C night) for > 3
weeks and then transferred to a warm temperature (30C day/25C night) before
initiation, produce only vegetative shoots (Núñez-Elisea et al., 1996). Thus, the
stems do not ‘remember’ that they had been exposed to floral inductive conditions
while still in rest. They responded to warm conditions present when shoot initiation
occurred.
Reproductive Physiology 113

This response to temperature conditions at the time of shoot initiation extends


to the formation of transition shoots if conditions change during early shoot
development. First reported by Naik and Mohan Rao (1943), transition shoots are
an unusual transition in expression of shoot type during a single growth flush
(Kulkarni, 1988b; Núñez-Elisea and Davenport, 1989, 1992b; Batten and
McConchie, 1995). The transition typically occurs near the middle of the extending
shoot. Resting buds possess preformed nodes, each of which contains a primordial
leaf or bract and a lateral meristem. The api-cal meristem initiates cell division at
the same time or soon after the nodal target tissues begin development (Mustard
and Lynch, 1946; L.B. Singh, 1960; Núñez-Elisea et al., 1996). Vegetative or
inflorescence development in the pre-formed primordia is underway before the
apical meristem begins to pro-duce differentiating cells. Transfer from a warm,
vegetatively inductive con-dition to a cool, floral inductive environment at early
bud break results in formation of V/F transition shoots (Fig. 5.5). Transfer from
cool to warm con-ditions at the same stage of bud break results in formation of F/V
transition shoots (Batten and McConchie, 1995; Núñez-Elisea et al., 1996).

The flowering response to temperature occurs in mangoes growing in


subtropical latitudes where cool temperature is the dominant induction fac-tor.
Many cultivars flower erratically in the low-latitude tropics, providing
continuously warm temperatures with high soil and atmospheric moisture. Under
such conditions, the age of stems is the dominant inductive factor (Buell, 1954;
Nakasone et al., 1955; Ravishankar et al., 1979; Ou and Yen, 1985; Issarakraisila
et al., 1992), and occasional cool night temperatures in the upper latitude tropics
have a positive moderating effect (Davenport, 2003).

Water relations

In the absence of cool temperatures, mango trees in the tropics may flower in
response to irrigation or rain following periods of water stress lasting 6–12 weeks
or more (Pongsomboon, 1991). Plant water stress has been presumed to provide the
stimulus for flowering (reviewed in Whiley, 1993; Chaikiatti-yos et al., 1994;
Schaffer et al., 1994; Davenport and Núñez-Elisea, 1997); how-ever, most of these
studies have failed to substantiate prolonged tree water deficit as a successful agent
for floral induction.
Experiments with container-grown trees fail to produce inflorescences after 8
weeks of water deficit (Wolstenholme and Hofmeyr, 1985). Under glasshouse
conditions (27°C day/22°C night; relative humidity (RH) ≥ 90%), container-grown,
monoembryonic cultivars were water stressed through deficit irrigation for 14 days,
resulting in an average leaf xylem water poten-tial of −3.9 MPa (Davenport, 1992;
Núñez-Elisea and Davenport, 1992a, 1994b). Following resumption of irrigation,
all trees grew vegetatively. Sim-ilarly, only vegetative growth was obtained when
container-grown trees were deprived of irrigation for 36 days during summer,
although leaf xylem water potentials of −3.78 MPa were attained (Núñez-Elisea
and Davenport, 1994b). Water stress imposed on plants during the cool autumn
months
114 T.L. Davenport

(night temperatures < 15°C) do not increase the proportion of apical buds forming
inflorescences, but expedited shoot initiation after rewatering (Núñez-Elisea and
Davenport, 1994b). These results demonstrated that cool temperatures provide
inductive conditions, whereas relief of water stress accelerated shoot initiation
under cool, inductive temperatures. Flowering was delayed when container-grown
monoembryonic mangoes were water-stressed at 18°C day/15°C night
(Chaikiattiyos et al., 1994). Water-stressed trees held at 29°C day/25°C night did
not flower.
Mango trees growing in the low-latitude tropics may flower after an extended
period of mild water stress (Harris, 1901; Collins, 1903; Kinman, 1918; Gangolly
et al., 1957; Gangolly, 1960; L.B. Singh, 1960). Pongsomboon et al. (1991)
observed flowering in field-grown trees in the tropics following 6 weeks of
withholding water. The primary impact of water stress appears to be prevention of
shoot initiation during stress. The accumulating age of stems is greater in water-
stressed trees than in trees maintained under well-watered conditions that promote
frequent vegetative flushes (Davenport, 1992, 1993; Schaffer et al., 1994). This
delay in flushing may provide more time for accu-mulation of a putative FP
(Schaffer et al., 1994) or reduction in the level of a putative VP (Davenport and
Núñez-Elisea, 1997; Davenport, 2000). Some cultivars appear to be better adapted
to such delays in growth and perform better in dry environments in the tropics.

Effect of N on flowering

Subsequent to the discovery of ethephon to stimulate mango flowering (Gon-zalez,


1923; Alcala and San Pedro, 1935), Barba (1974), Bueno and Valmayor (1974),
Astudillo and Bondad (1978), Bondad et al. (1978), Bondad and Apos-tol (1979),
Pantastico and Manuel (1978) and Bondad and Linsangan (1979) reported that
KNO3 could be used for the same purpose. This has been exploited in the low- and
mid-latitude tropics (Mosqueda-Vázquez and de los Santos de la Rosa, 1981;
Mosqueda-Vázquez and Avila-Resendiz, 1985; Núñez-Elisea, 1985, 1986; Ou and
Yen, 1985; Winston and Wright, 1986; Tongumpai et al., 1989; Goguey, 1993;
Ravishankar et al., 1993; Sergent et al., 1996; Yeshitela et al., 2004b, 2005). The
nitrate (NO3–) anion is the active component of KNO3 (Bueno and Valmayor,
1974), and ammonium nitrate (NH4NO3) is twice as effective as KNO3 (Núñez-
Elisea, 1988; Núñez-Elisea and Caldeira, 1988). In the low- and mid-latitude
tropics, receptive trees respond by developing visible reproductive buds within 2
weeks after application. The effective spray concentration is 1–10% KNO3,
depending on the age of the trees and climate. Two to four per cent KNO 3 or
calcium nitrate (Ca(NO3)2) and 1–2% NH4NO3 are effective for stimulating
flowering in most conditions. The physiological and temporal timing of application
is important. Old trees, non-vigorous trees, and trees in which vegetative flushes
have been discouraged by low water potentials produce the best response to NO3–
induction (N. Golez, personal communication, the Philip-pines, 1989).
Reproductive Physiology 115

Chemical bud forcing is most effective in the tropics where distinct wet and
dry seasons prevail. The response to chemical bud forcing by NO3− and ethephon
diminishes at latitudes > 22° N or S (Mosqueda-Vázquez and de los Santos de la
Rosa, 1981). Their effect may involve the decline of night temperatures from 
20°C around the equator to  10°C between 22° and 27° N or S latitude during
winter months or by late summer vegetative flushes. Trees in the wet or dry
subtropics at 25° N or S have not responded to treat-ments (Davenport, 1993).

Stems must be sufficiently mature, dark green with a minimum age of 4


months since the previous limp, red-leaf stage in easily induced cultivars and 5
months for more recalcitrant cultivars to obtain a reproductive shoot response in
the low-latitude tropics (Davenport, 2003). Bueno and Valmayor (1974) indicated
that leaves must be brittle when hand-crushed. Núñez-Elisea (1986, 1988) reported
that stems must be at least 6 months old. Trees that experience autumn dry periods
become responsive to treatments as early as October (northern hemisphere).
Groups of stems within tree cano-pies are produced through asynchronous flushes
of growth, and vary in age; only a few are responsive to the first inductive spray.
Subsequent biweekly applications cause flowering in canopy sectors as they reach
the age-depen-dent requirement for initiation. Early and out-of-season flowering
and fruit-ing can thereby be achieved.

KNO3 may be floral inductive in mango (Barba, 1974); however, trees in the
upper latitude tropics typically flush vegetatively rather than produce bloom when
either KNO3 or NH4NO3 is sprayed between June and Septem-ber (N. Golez,
personal communication, the Philippines, 1989). The warm, rainy season
producing frequent flushes of growth during this period is con-ducive to a
vegetative response to the sprays. These results indicate that KNO3 and NH4NO3
stimulate shoot initiation but do not determine bud morphogenesis. In buds
released after KNO3 or NH4NO3 treatments, the ratio of leaf-generated FP to VP
and not NO3– causes initiating buds to become reproductive. Kulkarni (1988b,
2004) suggested that the floral stimulus is present in stems when buds are forced in
response to KNO3 and suggested that KNO3 may also sensitize buds to the floral
stimulus. Davenport (2003), T.L. Davenport and J. Oleo (2006, unpublished data)
and F. Ramirez and T.L. Davenport (submitted for publication) observed 100%
vegetative shoots when 4% KNO3 was foliar applied to 2-month-old stems;
whereas, applica-tion of the same spray treatment to 4.5-month-old stems on trees
in the same orchards resulted in 100% reproductive shoots.
Trees with high leaf N levels rarely flower in the tropics. Lack of flower-ing is
always due to frequent vegetative flushes of growth, especially during the rainy
season. Mango trees must have leaf N levels of 1.4% or less in order to suppress
frequent flushes of vegetative growth (Davenport, 2003). Leaf N levels of < 1.1%
suppress frequent flushes but also provide insufficient nutri-tion to support good
cropping. Thus, 1.1–1.4% N levels in leaves appear to be optimum for good
commercial production and control of flowering time in a managed orchard. The
application of KNO3 to the foliage of the resting stems 4–5 months after the limp,
red-leaf stage will cause a flowering response.
116 T.L. Davenport

Photoperiod

Flowering in most trees does not appear to be under photoperiodic control


(Kozlowski et al., 1991). Mango cultivation is concentrated between 27° N and 27°
S where the shortest annual photoperiod is c.10.5 h and the longest photoperiod is
c.13.5 h. Cultivars in the upper-latitude tropics and subtropics flower during the
winter when photoperiods are short; however, trees in the low-latitude tropics,
where a 12-h photoperiod is nearly constant, can flower at any time of the year.
Furthermore, flowering on spring-initiated shoots in the subtropics occurs during
summer (Schaffer et al., 1994). Studies have failed to demonstrate a correlation
between 8-h photoperiods and flowering (Maiti, 1971; Maiti and Sen, 1978; Maiti
et al., 1978). Núñez-Elisea and Daven-port (1995) studied the effects of 11-, 12-,
13- and 24-h photoperiods at 18°C day/10°C night, or 11- and 13-h photoperiods at
30°C day/25°C night on flowering of container-grown trees. Photoperiod had no
effect on the fate of buds, and the promotive effect of cool temperatures on
flowering was inde-pendent of photoperiod. Photoperiods of 11-, 12- or 13-h with
18°C day/10°C night, caused flowering in trees within 40 days. The 24-h
photoperiod with 12-h thermoperiods of 18°C and 10°C caused flowering of trees
within 35 days. Photoperiods of 11- or 13-h at 30°C day/25°C night resulted in
vegeta-tive growth only. With warm temperatures, vegetative shoots were
produced in 17 days. These results confirm that floral induction is caused by cool
tem-peratures and not by short photoperiods and that warm temperature, not a long
photoperiod, caused vegetative induction.

5.6 Hormonal Influence on Flowering


FP is a protein product of the FT gene in Arabidopsis (Corbesier et al., 2007) and
the Hd3a gene in rice (Tamaki et al., 2007) and moves in phloem from leaves to
buds; there is little evidence that phytohormones are directly involved as the FP.
Phytohormones appear to be responsible for shoot initia-tion in conditions that are
floral inductive.

Ethylene

Smudging has been utilized to stimulate mango flowering in the Philippines. Only
branches that attain sufficient age respond to smudging by forming reproductive
shoots (Acala and San Pedro, 1935; Bueno and Valmayor, 1974). Rodriguez
(1932), investigating smoke-induced flowering of pineapple, pro-posed that
ethylene, generated by burning material, may stimulate flower-ing. Dutcher (1972)
confirmed that smoke from smudge fires contained ethylene. Smudging and the use
of ethephon in 1968 by F. Manuel (Barba, 1974) and others (Bondad, 1972, 1976)
to promote mango flowering sug-gested that endogenous ethylene is integral for
floral induction (Barba, 1974; Bondad, 1976; Chadha and Pal, 1986). Ethephon
effectively promotes flowering
Reproductive Physiology 117

of mangoes under specific conditions in the low-latitude tropics (Davenport and


Núñez-Elisea, 1997).
The involvement of endogenous ethylene in flowering is supported by
observations that indirectly link it to symptoms of ethylene production. Extrusion
of latex from terminal buds occurs at the time of inflorescence ini-tiation, and
epinasty of mature leaves near the apex during expansion of the panicle has been
observed (Davenport and Núñez-Elisea, 1990, 1991). Both are symptoms of plants
exposed to high ethylene levels (Abeles, 1973). Indi-rect support also comes from
reports that KNO3-stimulated flowering of mango is mediated by increased levels
of endogenous ethylene (Thuck-Thye, 1978; Lopez et al., 1984). Mosqueda-
Vázquez and Avila-Resendiz (1985) reported that the efficacy of KNO 3 was
negated by cobalt chloride (CoCl2) and silver nitrate (AgNO3), which inhibit the
synthesis and action of ethyl-ene, respectively, when sprayed 1–4 h after KNO3.
Saidha et al. (1983) reported a gradual increase in endogenous leaf ethylene
production as the season of floral initiation approached. Ethylene production by
stems producing repro-ductive shoots was up to fivefold that of resting stems.
Inconsistent (Pandey et al., 1973; Sen et al., 1973; Winston and Wright, 1986)
or non-responsive results with ethephon (Pandey and Narwadkar, 1984; Ou and
Yen, 1985; Pandey, 1989) or smudging (Sen and Roy, 1935), especially during
warm, non-inductive conditions, have been reported. Dav-enport and Núñez-Elisea
(1990, 1991) reported elevated ethylene production in mango stems in response to
ethephon sprays without an accompanying floral response. Experiments were
conducted during floral-inductive and non-inductive periods. Unlike Saidha et al.
(1983), they observed no increase in ethylene production rates prior to or during
panicle development.
The effect of ethylene on flowering is unresolved. It is likely that ethylene
stimulates shoot initiation by inhibiting auxin transport from leaves to buds and
stems (Morgan and Gausman, 1966; Beyer and Morgan, 1971; Riov and Goren,
1979, 1980; Ramina et al., 1986). This may increase the ratio of cytoki-nin to auxin
in buds and stimulate shoot initiation (Davenport, 2000). Other factors (i.e. cool
temperatures or aged leaves) may be responsible for floral induction (Ona and de
Guzman, 1982; Davenport, 1993).

Auxin

Although auxin may have a critical role in floral induction of mango (Chadha and
Pal, 1986; Hegele et al., 2006), there is little supporting evidence. The application
(L.B. Singh, 1961; Singh and Singh, 1963; Bakr et al., 1981; Pandey and
Narwadkar, 1984) and analysis of auxin in leaves (Paulas and Shanmu-gavelu,
1989; Sivagami et al., 1989), stems (Chen, 1987) and shoots (Chacko et al., 1972b)
have been reported in relation to mango flowering. These studies are inconclusive
due to inconsistencies in purification and analytical methodolo-gies (Davenport
and Núñez-Elisea, 1997).
Auxin may indirectly stimulate root-produced cytokinins through initia-tion of
new root growth. Auxin is transported basipetally from growing
118 T.L. Davenport

shoots and leaves to roots (Goldsmith, 1968; Cane and Wilkins, 1970; Wilkins and
Cane, 1970; Goldsmith and Ray, 1973; Lomax et al., 1995) and stimulates root
initiation (Hassig, 1974; Wightman et al., 1980). The efficacy of various auxins for
stimulating adventitious rooting of mango marcots and cuttings was reviewed by
Davenport and Núñez-Elisea (1997).
Auxin inhibits shoot initiation (Davies, 1995) and confers apical domi-nance
by preventing axillary bud break. Leaf-produced auxin and petiolar auxin transport
capacity declines as leaves age (Veen, 1969; Veen and Jacobs, 1969; Davenport et
al., 1980). The interaction of decreasing auxin and accu-mulating cytokinins in
resting buds may explain the cyclic nature of shoot initiation. The ratio of cytokinin
to auxin levels in buds regulates shoot initiation (Skoog and Miller, 1957;
Bangerth, 1994; Cline et al., 1997; Beveridge et al., 2003).

Cytokinins

Relationships between mango flowering and the endogenous levels of cyto-kinins


in leaves (Paulas and Shanmugavelu, 1989; Kurian et al., 1992), stem tips
(Agrawal et al., 1980) and xylem sap (Chen, 1987) and the effect of cyto-kinin
applications on bud break and shoot development have been reported. Chen (1985)
described precocious flowering of mango shoots in response to early October
application of 6-benzylaminopurine (BA). Flowering was observed 1 month
following application and 3 months later on non-treated trees. Núñez-Elisea et al.
(1990) reported numerous reproductive shoots per stem in response to the synthetic
cytokinin, thidiazuron, during cool, floral inductive conditions; however, numerous
vegetative shoots per stem were initiated when thidiazuron was applied during
warm, vegetatively induc-tive conditions. Early bud break was not achieved
following foliar applica-tion of Promalin (commercial formulation of BA and
gibberellins A4+A7) (Oosthuyse, 1991b), BA (A.K. Singh and Rajput, 1990) or
kinetin (Singh and Singh, 1974).

Chen (1987) reported the lowest levels of putative trans zeatin and its riboside
were translocated from roots during the vegetative shoot growth and resting stages,
whereas the highest levels occurred during early flower-ing and full bloom. Paulas
and Shanmugavelu (1989) observed no significant difference in cytokinin levels of
the fourth and fifth leaves during resting bud and flowering. Cytokinin levels in
mango stem buds increased during expo-sure to cool, floral inductive temperatures
(Bangerth et al., 2004). Agrawal et al. (1980) described 11 cytokinin-like
substances isolated from stem tips of an alternate-bearing cultivar in ‘on’ and ‘off’
years. Kurian et al. (1992) reported a link between PBZ applications and reduction
in cytokinins in mango leaves with treatments, perhaps caused by reduction in
feeder root development and formation of thick, blunt roots (Bausher and
Yelenosky, 1987; Peng et al., 1991; Burrows et al., 1992; Yelenosky et al., 1993).
Concurrent with this response was suppression of bud initiation and reduced
internode lengths for c.2 years.
Reproductive Physiology 119

The role of cytokinins in flowering is unresolved due to sampling of dif-ferent


organs at non-comparable times or conditions. The elevated cytokinin levels found
prior to and during flowering and the flowering response to applied BA led to the
conclusion that cytokinins are involved in flowering of mango (Chen, 1985, 1987;
Bangerth, 2006); however, such responses can be explained if cytokinins are
involved in stimulation of bud break (i.e. shoot initiation) during floral inductive
conditions.
A well-documented role for cytokinins in higher plants, especially evi-dent in
vitro, is bud organogenesis (Skoog and Miller, 1957; Miller, 1963; Takahashi,
1986; Salisbury and Ross, 1992; Davies, 1995; Haberer and Kieber, 2002). The
primary cytokinins in higher plants are trans zeatin, dihydrozeatin, isopentenyl
adenine and their ribosides. They are translocated from roots and accumulate in
resting buds (Hendry et al., 1982a, b) or can possibly be synthe-sized in nearby
tissues as regulated by auxin (Nordstrom et al., 2004; Tanaka et al., 2006). Their
rate of accumulation may relate to periodic root flushes that alternate with shoot
flushes (Krishnamurthi et al., 1960; Bevington and Castle, 1986; Cull, 1987, 1991;
Parisot, 1988; Williamson and Coston, 1989).

Gibberellins

Gibberellins are tetracyclic diterpenoid compounds that vary in biological activity


according to the type and location of substituted side groups on a basic ent-
gibberellane skeleton. The number of known gibberellins is > 100 (Pearce et al.,
1994). Reproductive shoot initiation is suppressed in many woody angiosperms by
gibberellic acid (GA3) (Pharis and King, 1985). GA3 inhibits mango flowering
(older literature reviewed in Davenport and Núñez-Elisea, 1997; Núñez-Elisea and
Davenport, 1998).
GA3 inhibition of mango flowering is correlated with the applied con-
centration (Kachru et al., 1971, 1972) and may cause buds to develop vegeta-tively
under floral-inductive conditions. Núñez-Elisea and Davenport (1991a, 1998)
reported a delay in initiation of axillary shoots when GA3 was foliar applied to
deblossomed stems during cool, floral inductive temperatures. Higher
concentrations caused longer delays in shoot initiation. GA 3 did not inhibit floral
induction, so long as cool, inductive temperatures were present during axillary
shoot initiation. Late initiating buds, which grew during warm, spring
temperatures, however, formed vegetative shoots. Similar delays in reproductive
shoot initiation in response to GA3 application was reported by Shawky et al.
(1978) and Turnbull et al. (1996). Multiple applica-tions, even at lower rates, are
more effective than a single application (Tomer, 1984; Turnbull et al., 1996;
Davenport and Smith, 1997). GA3 treatment has been recommended in the Canary
Islands to delay flowering until the danger of frost has passed (Galán-Saúco, 1990).
In the subtropics of Australia, it is used to prevent flowering in newly planted trees
during the spring so that the full growing period can be utilized for vegetative
growth, thereby hastening orchard establishment (A.W. Whiley, personal
communication, Queensland, 1996).
120 T.L. Davenport

Response to GA3 varies among cultivars, growing conditions and timing of


application (Tomer, 1984; Oosthuyse, 1995a; Turnbull et al., 1996; Sánchez-
Sánchez et al., 2004). GA3 can delay shoot initiation beyond the floral induc-tive
window, resulting in a vegetative flush when shoots develop in warm weather
(Kachru et al., 1971, 1972; Núñez-Elisea and Davenport, 1991a, 1998; S. Gazit,
personal communication, Israel, 1993). The variable response to GA3 may be
related to levels of active gibberellin in buds at the time of applica-tion,
inconsistent uptake or differential sensitivity of buds, depending on their position
(apical versus axillary) or age (Núñez-Elisea and Davenport, 1991a, 1998).
Efficacy is related to the timing of application; immediately prior to normal shoot
initiation appears to be most effective (Davenport and Smith, 1997).
Reports of endogenous gibberellins in mango tissues, especially in buds, are
difficult to interpret with respect to a regulatory role in bud break or flowering.
Problems include sampling of tissues other than apical buds, i.e. whole stems
(Tongumpai et al., 1991b), leaves (Paulus and Shanmugavelu, 1989; Sivagami et
al., 1989) and xylem sap (Chen, 1987), or at times when developing shoots may
contribute to the overall result (Chen, 1987). Pal and Ram (1978) tentatively
identified the presence of gibberellins A1, A3, A4, A5,
A6, A7 and A9. Chen (1987) identified gibberellins A1/3, A4/7, A5, A17, A20 and
A29. The estimated levels of gibberellins in apical buds for 6 months prior to
the flowering season were reported to be higher in the ‘off’ year than in the ‘on’
year of an alternate-bearing cultivar (Pal and Ram, 1978). Chen (1987) reported the
highest levels of gibberellins in xylem sap during leaf differen-tiation and lower
concentrations during rest, panicle emergence and full flowering. Tongumpai et al.
(1991b) observed increasing levels of gibberellins in whole stems over the 16
weeks prior to vegetative shoot emergence and decreasing levels over the same
period prior to panicle development. Gib-berellins A1, epi-A1, A3, A19, A20 and
an unidentified gibberellin in buds and leaves from shoot and stem tips of different
ages have been quantified (Dav-enport et al., 2001b). The detected gibberellins are
members of the early 13-hydroxylation pathway of gibberellin synthesis
(Takahashi, 1986; Pearce et al., 1994). Gibberellins A3 and A19 were the most
abundant gibberellins in apical stem buds. The concentration of GA3 increased
within buds with increasing age of stems, although concentrations of other GAs
were variable. The concentration of GA3 did not change significantly with age in
leaves, whereas that of most of the other GAs declined. Davenport et al. (2001b)
concluded that elevated GA3 levels in buds may enhance or maintain the synthesis
or activity of endogenous auxin to maintain low cytokinin/auxin ratios and enhance
inhibition of shoot initiation (Jacobs and Case 1965; Scott et al., 1967; Pharis et al.,
1972; Ross et al., 1983; Law 1987; Law and Hamilton, 1989).

The roles of gibberellins and other phytohormones in shoot initiation and


induction is unclear. Endogenous levels in buds and leaves must be cor-related
with physiological events in individual stems. Experimental approaches should
include examination of resting buds up to both vegetative and repro-ductive shoot
initiation to avoid misinterpretation of results. Experiments
Reproductive Physiology 121

should utilize plants grown under defined conditions with specific environ-mental
controls for evaluation of cause and effect. Finally, extraction and purification
protocols should include quantifiable internal standards and use of sensitive
unambiguous analytical techniques.

Plant growth retardants

Plant growth retardants have been evaluated to stimulate early or more intense
flowering, especially in the ‘off’ year of alternate-bearing cultivars (Davenport and
Núñez-Elisea, 1997). They are in three main classes: (i) the gibberellin transport
inhibitor, daminozide (N-dimethylamino-succinamic acid), known as alar or B-
Nine; (ii) the onium type, chloremquat chloride (2-chloroethyl trimethylammonium
chloride), known as cycocel and CCC; and (iii) the steroid-synthesis-inhibiting
triazoles, for example PBZ (PP-333), known as Cultar®, and uniconazole, known
as XE-1019 or Sumagic (Rademacher, 1991, 2000a). The latter two classes of
compounds inhibit ent-kaurene syn-thetase, an enzyme in the gibberellin synthesis
pathway (Nickell, 1983; Dalziel and Lawrence, 1984; Rademacher, 1991, 2000a).
Applying daminoz-ide results in increased gibberellin levels, perhaps due to the
inability to dis-tribute it properly (Rademacher, 1991). Plant responses may depend
upon whether target tissues are near the site of gibberellin synthesis or sufficiently
removed from it to be affected by the inhibited translocation.

Daminozide and cycocel


The efficacy of daminozide and cycocel for increasing flowering in the ‘off’ sea-
son of alternate-bearing cultivars has been studied (Maiti et al., 1972; Mukhopad-
hyaya, 1978; Rath and Das, 1979; Suryanarayana, 1980; Rath et al., 1982; Ou and
Yen, 1985), together with their ability to stimulate early flowering (Suryanarayana
and Rao, 1977; Chen, 1985; Kurian and Iyer, 1993a, b). Enhanced, inconsistent
flowering occurs in response to these compounds, especially cycocel.

Triazoles
PBZ is being used (except in the USA where it has not been cleared for use) to
stimulate enhanced or early flowering. It is best applied to the soil due to its low
solubility, long residual activity and lack of efficient foliar uptake (Rademacher,
2000b). PBZ applied as a soil drench (1–20 g active ingredient (ai)/tree) reduces
internode lengths and causes earlier and enhanced flower-ing in mango trees
(Hasdiseve and Tongumpai, 1986; Haw, 1986; Hongsb-hanich, 1986). Depending
on climate, residual activity lasts for c.2 years (Kulkarni, 1988a). These results
have been confirmed in different locations in the tropics (Davenport and Núñez-
Elisea, 1997; Yeshitela et al., 2004a, b). Nartvaranant et al. (2000) recommended
soil application of PBZ at 1–1.5 g ai/m of canopy diameter to achieve flowering in
90–120 days if the trees are stimulated to flush. Davenport (2003) observed that
such treatments allowed a reduction of c.1 month in the time required for stem rest
before stimulating them to initiate reproductive shoots using KNO 3. PBZ also
reduces alternate
122 T.L. Davenport

bearing of some cultivars (Hillier and Rudge, 1991; Burondkar and Gunjate, 1993;
Rao, 1997; Rao et al., 1997; Rao and Srihari, 1998; Vijayalakshmi and Srinivasan,
1999). Cultivars that tend to flower with minimal inductive impe-tus are more
responsive and can be induced to flower out-of-season using PBZ (Tongumpai et
al., 1989). Núñez-Elisea et al. (1993) demonstrated that application of PBZ and
uniconazole advanced bud break of containerized trees in controlled environment
chambers, but cool temperatures were neces-sary to induce flowering. Initiated
shoots were induced to be vegetative in warm temperatures. The greater proportion
of purely reproductive panicles in treated plants (compared with controls) suggests
that triazoles impact the level of a putative VP, probably a gibberellin. Whiley
(1993) suggested a sec-ondary mechanism for the floral promotive action of PBZ
on mangoes, not-ing inconsistent responses in the literature between cultivars,
environments and application times.

Application of PBZ reduces the number of panicles, despite increased fruit set
(Goguey, 1990). Davenport (1987, 1994) observed neither growth inhibition nor
enhanced or early flowering in response to root drenches or bark banding with
uniconazole (1–5 g ai/tree) in trees growing in alkaline, calcareous soil. He
reported that new shoot growth was stunted with extremely short internodes when
trees were severely pruned soon after or as long as 3 years after treatment. Yield
was severely reduced due to the lack of normal growth flushes. The growth
stunting effect continued for 7 years after pruning. Davenport (1994) warned that
use of triazole plant growth retar-dants for control of tree growth, flowering or
yield must be done with con-siderable caution, especially if severe pruning of the
trees is anticipated. Residual uniconazole or PBZ applied as a soil drench or bark
band is appar-ently retained in high concentrations in main scaffolding branches. In
Cen-tral and South America, growers utilize PBZ annually to stimulate early
flowering. A test tree should be severely pruned to determine if the trees are
affected by PBZ to anticipate the orchard response to later severe pruning.

Certain gibberellins (i.e. GA1) are necessary for shoot elongation. Inhibi-tion
of bud break and shoot elongation in response to application of the growth
retardants cycocel (Maiti et al., 1972) and triazoles (Kulkarni, 1988a; Burondkar
and Gunjate, 1991, 1993; Tongumpai et al., 1991a; Kurian et al., 1992; Winston,
1992; Kurian and Iyer, 1993a, b; Núñez-Elisea et al., 1993; Wer-ner, 1993) have
been reported. Elongation of panicles is inhibited, especially by high levels of
triazoles (Kulkarni, 1988b; Winston, 1992; Davenport, 1994; Salomon and
Reuveni, 1994). Inflorescences in treated trees may become compact, improving
opportunities for disease and insect attack (Winston, 1992). Kurian et al. (1992)
associated reduced cytokinin levels in leaves with inhibition of shoot initiation in
plants treated with soil drenches of PBZ. Ele-vated, concentration-dependent levels
of phenolic compounds were also found in resting apical buds of PBZ-treated trees
(Kurian et al., 1994). They suggested that low cytokinin activity and high phenolic
levels in buds con-tributed to inhibition of shoot initiation.

The combined impact of the gibberellin synthesis-inhibiting triazoles on shoot


initiation, induction, and elongation implies that several different
Reproductive Physiology 123

gibberellins regulate specific activities in mango plants. This is supported by the


inhibitory effect of GA3 on shoot initiation in contrast with early initia-tion of
flowering in triazole-treated trees. Compression of reproductive and vegetative
shoot internodes may involve inhibition of GA1 synthesis. Stimu-lation of
flowering instead of vegetative growth during early initiation in triazole-treated
plants in marginal or non-floral inductive conditions, sug-gests that the putative
VP, a gibberellin other than GA3 or GA1, is reduced when gibberellin synthesis is
inhibited.

5.7 Photoassimilate Influence on Flowering

Flowering may be regulated by C:N ratios with high levels being conducive to
flowering (Kraus and Kraybill, 1918). Photoassimilates reaching the apical bud
from leaves was central to several theories of floral induction (Sachs, 1977; Bernier
and Sachs, 1979; Bernier et al., 1981, 1993; Bernier, 1988) includ-ing mango
(Mallik, 1951; L.B. Singh, 1960; Chacko and Ananthanarayanan, 1982;
Rameshwar, 1989) and other species (Allsopp, 1965; Sachs, 1977; Mishra and
Dhillon, 1978; Ramina et al., 1979; Bernier et al., 1981; Sachs and Hackett, 1983).
The theory of photoassimilate diversion to the apical bud (Sachs et al.,
1979) is the basis for the carbohydrate-regulated flowering models (see below).
Sugars are utilized during panicle development (Ravishankar and Mohan Rao,
1982). Starch reserves and C:N ratios have been correlated with flowering (Mishra
and Dhillon, 1978; Suryanarayana, 1978a, b, c; Chacko and Ananthanarayanan,
1982; Whiley et al., 1988, 1989, 1991; Robert and Wolsten-holme, 1992;
Shivashankara and Mathai, 1995), and the subject has been reviewed (L.B. Singh,
1960, 1972; Singh, 1979; Chacko, 1986, 1991; Chadha and Pal, 1986; Pandey,
1989). Starch accumulation during extended periods of canopy rest prior to
flowering provides supportive evidence, but there is little consensus regarding the
role of carbohydrates and N in flowering.
Photoassimilates may be necessary for floral induction. If a florigenic
promoting gene product is synthesized in leaves in small amounts, it must be able
to move to those buds via phloem. Due to the requirement for high sol-ute
concentrations to motivate phloem flow, the low concentration of the FP could not
cause fluid movement through sieve tubes of the phloem on its own. The much
higher concentrations of photoassimilated sugars carried by water loading into the
phloem in leaves passively transports the FP towards the various sinks, including
respiring buds, where they are utilized for floral induction.

5.8 Horticultural Manipulation of Flowering


Mango flowering can be stimulated by trunk or branch girdling, defoliation and
deblossoming (Pandey, 1989). Responses vary with cultivar and envi-ronment.
Trunk girdling of mango trees to promote flowering is inconsis-tently effective
(Kinman, 1918; Gaskins, 1963; Winston and Wright, 1986) and
124 T.L. Davenport

can be detrimental to trees, especially if done in subsequent years. It has been


shown to increase flowering in the ‘off’ year of alternate-bearing cultivars;
however, it either has no effect or is only marginally beneficial in the ‘on’ year
(Mallik, 1951; Rath and Das, 1977, 1979; Rath et al., 1982; Rameshwar, 1989).
Girdling in late summer or early autumn usually results in less vegetative flushing
prior to flowering, which is enhanced in trees exposed to marginally inductive
conditions. Tree response is dependent on the width of the girdle. Narrow cuts
result in either a short-term or no response; whereas, girdles that are too wide can
kill trees if they do not close within a reasonable time. Gir-dling cuts phloem
transport, starves roots of photoassimilates and interrupts auxin transport to roots
(Morris and Thomas, 1978; Hegele et al., 2004). These are detrimental to root
development and can alter the bud cytokinin:auxin ratio due to reduced cytokinin
translocation from roots. This results in delayed shoot initiation, which can impact
the level of the age-dependent, putative VP when shoot initiation occurs. The delay
in flushing, therefore, enhances flowering. Defoliation of trees stimulates flushing,
possibly by altering the cytokinin:auxin ratio in buds because leaves are the
primary source of auxin.
Bloom delay is useful where recurring temperatures < 15°C stimulate
flowering, but continued low temperatures hamper pollination, fertilization and
early fruit development (Young and Sauls, 1979; Wolstenholme and Mul-lins,
1982a, b; Whiley et al., 1988; Galán-Saúco et al., 1991). Low temperatures cause
production of seedless, underdeveloped fruit. Deblossoming stimu-lates growth of
dormant axillary buds, which produce inflorescences if initia-tion occurs under
conditions conducive to floral morphogenesis (Singh et al., 1974). Late blooms can
also be promoted with ethephon (Chadha and Pal, 1986; Galán-Saúco et al., 1993)
and cycloheximide (Pal and Chadha, 1982; Shu, 1993). Sprays of these compounds
cause abscission of apical panicles, thereby releasing dormant axillary buds that
will produce inflorescences under cool, floral-inductive conditions of the
subtropics.

5.9 Conceptual Flowering Models


Several conceptual models have been proposed that attempt to explain the
physiological basis of mango flowering. Each model should be viewed as a
collection of integrated ideas, which require rigorous testing for validity within the
context of the models. A useful model should explain how flower-ing and
vegetative growth is regulated in all cultivars and races in both humid and dry
climates in the tropics and subtropics. It should also be sup-ported by the
preponderance of research evidence. The flowering models are either carbohydrate-
regulated or hormone-regulated.

Carbohydrate-regulated flowering models

Cull (1987, 1991) presented a holistic approach for tree crop research and
management to maximize sustainable fruit production. This concept is based
Reproductive Physiology 125

on the axiom of genotype/environment adaptability expressed through the annual


phenological cycle and is an alternative to the traditional reductive-based approach
to crop research and development. He proposed that pro-ductive cultivars follow
‘normal’ phenological patterns from year to year due to gene expression in specific
environments. A significant departure from this pattern results in reduced or total
crop failure. Annual variations in climatic conditions that alter tree phenology can
be countered by strategic applications of nutrients, water, plant growth regulators
and canopy manip-ulation. The model does not attempt to explain the intricacies of
shoot ini-tiation or induction, but takes a broader approach in detailing temporal
relationships between reproductive and vegetative growth that lead to reli-able
cropping.

The fundamental principle underlying this model is that yield is the product of
photoassimilate (carbohydrate) accumulation and subsequent redistribution during
the annual growth cycle. Accumulated photoassimi-lates would drive critical
growth events that require higher levels of resources than are available from
current photoassimilate supplies. Cultivars that pro-ceed with balanced
reproductive, vegetative and rest phases are more likely to have sufficient carbon
resources to meet periods of critical demand and therefore will sustain high yields.
The model illustrates floral initiation as occurring after a 2- to 3-month rest period
during autumn/winter when a critical threshold level of carbohydrate is reached in
buds together with a putative floral stimulus. Bud break during cool weather results
in a high per-centage of flowering stems (> 90%; Searle et al., 1995) with fruit set
and reten-tion suppressing vegetative flushing on individual fruiting stems until
after they have matured and harvested. Shortly after harvest, vegetative buds are
released and a flush of growth occurs during the summer, which is followed by a
period of strong root growth. The regenerated canopy becomes a source for
rebuilding photoassimilate reserves that are stored in the roots, bark and resting
stems. In the tropics, growth events are less orderly, and cultivar and management
skills are of greater importance. The pre-flowering rest period is usually achieved
by drought as temperatures remain above the critical threshold for shoot growth
(15C) (Whiley et al., 1989). Other practices used with some success to enforce
canopy quiescence are girdling and the applica-tion of growth retardants.

The principles of phenological modelling have been advanced into work-ing


pheno/physiological models for avocado (Whiley, 1994) and mango (Searle et al.,
1995). The advantage of this approach is that the annual pro-gression of growth
cycles with associated physiological changes is studied concurrently, adding a
further dimension to our understanding of tree growth and development.
Information gathered in this way provides opportunities to identify and assess
critical yield-limiting events, which in the case of man-goes largely relates to the
success or failure of flowering.
Chacko (1991) proposed a flowering model driven by assimilate supply and
diversion to apical meristems (Fig. 5.6). Environmental conditions, such as water
stress, cool temperatures, high evaporative demand, flooding, girdling and other
events that inhibit vegetative growth result in a shift in
126
FLORAL FLORAL

INDUCTION INHIBITION

· Water stress KNO3


· Low temperature (cultivar and location
· High VPD specific)
· Flooding
Exogenous
? gibberellin

· Stem girdling High


· Root pruning starch MISSING LINK
GROWTH
INCREASED ASSIMILATE STIMULATION · High temperature
GROWTH ASSIMILATE supply DIVERSION
CHECK and high · High humidity
to SHOOT APEX from SHOOT APEX gibberellin · High soil moisture

T.L. Davenport
· Mild nitrogen
stress Sugar

High
nitrogen
· High reserves
· Growth · Low reserves
· Efficient
retardants · More wood
assimilate
· Inhibitors formation
partitioning
High
Dwarf/precocious Over vigorous Frequent
gibberellin
cultivars cultivars flushing of
levels
e.g. ‘Irwin’ e.g. ‘Kensington’ roots and shoots

HEREDITY JUVENILITY

Fig. 5.6. Chacko’s Assimilate Supply and Diversion Flowering Model, a carbohydrate-regulated flowering model (Source: Chacko, 1991).
Reproductive Physiology 127

carbohydrate partitioning and a diversion of soluble assimilates to stem api-ces.


The elevated carbohydrate status in buds, together with a floral stimulus, results in
floral induction. Vigorously growing cultivars and juvenile plants have low starch
reserves (Whiley et al., 1988, 1989, 1991) and a diversion of soluble assimilates
from stem apices results in floral inhibition. Conditions that promote vegetative
growth, i.e. high temperature and moisture, high gibberellins and N, also lead to
floral inhibition. Experiments involving chemical girdling of trees are based on this
model (Blaikie et al., 1999).

Hormone-regulated flowering models

Tri-factor Hypothesis of Flowering


Extensive work on movement of the putative floral stimulus across grafts from
donor to receptor stems (Kulkarni, 1986, 1988b) and the inhibitory influ-ence of
fruit on subsequent flowering (Kulkarni and Rameshwar, 1989) form the basis of a
flowering model proposed by Kulkarni (1991): the Tri-factor Hypothesis of
Flowering in mango (Kulkarni, 2004). This theory (Fig. 5.7) proposes an
interactive role for a putative, cyclically produced floral stimu-lus in leaves, a floral
inhibitor in leaves and fruits, and bud activity during the floral cycle. During
dormancy following a vegetative cycle, genetic and

Genetic and Environmental Factors

Flowering promoter Flowering inhibitor Bud activity


synthesized in the and vegetative in synchrony with
leaves in the floral promoter the floral cycle
cycle synthesized in the
leaves and possibly
other organs

Pure panicles Mixed leafy panicles Vegetative flush

Fig. 5.7. Kulkarni’s Tri-factor Hypothesis of Flowering in mango, a hormone-


regulated flowering model (Source: Kulkarni, 2004).
128 T.L. Davenport

Mango flowering model

AUXIN
PHOTOASSIMILATES FRUIT
GIBBERELLINS A3

Ax

AUXIN VEGETATIVE SHOOT MIXED SHOOT GENERATIVE SHOOT


GIBBERELLINS
GA3 GA1
GAx

FREQU ENT INDUCTION


PROMOTER
IN LEAVES
CHILLING TEMP.
OTHER FACT ORS?

VEGETATIVE
GROWTH WATER STRESS
SHOOT INITIATION
PRUNING
DEFOLIATION CHILLING TEMP.
NITRATE SPRAY
ETHYLENE
ROOT INITIATION GIRDLING

STORAGE CARBOHYDRATES ROOTS CYTOKININS

Fig. 5.8. Davenport’s Comprehensive Conceptual Hormone-regulated Flowering Model


(Source: Davenport and Núñez-Elisea, 1997; Davenport, 2000). Single lines indicate
promotive impact. Double lines indicate inhibitory impact.

environmental factors determine the level of synthesis of the putative floral


stimulus. Flowering occurs only if certain correlative factors are present, for
example if the receptor bud becomes active. If fruits are or have been recently
present on the stem, vegetative growth will result. Presence of the putative floral
inhibitor in leaves interferes with expression of the floral stimulus resulting in
vegetative growth. The level of the floral stimulus determines the response: high
levels give rise to normal panicles, intermediate levels give rise to mixed panicles
and low levels result in vegetative growth.

Comprehensive Conceptual Flowering Model


This is a model of flowering involving the various classes of phytohormones
(Davenport, 1992, 1993, 2000, 2003; Davenport and Núñez-Elisea, 1997) (Fig. 5.8)
based on many lines of experimental evidence as well as on research of other
tropical and subtropical fruit crops with similar phenological cycles (Menzel, 1983;
Davenport, 1990, 1992; Menzel et al., 1990; Menzel and Simp-son, 1994;
Davenport and Stern, 2005). Focusing on events occurring in indi-vidual buds, it is
applicable to monoembryonic and polyembryonic cultivars in the tropics and
subtropics and attempts to explain the physiological basis for the annual
progression of the phenological cycle.

SHOOT FORMATION. Two distinct events must occur for vegetative or repro-ductive
growth to occur in resting apical or lateral buds of mango: (i) the
Reproductive Physiology 129

bud(s) must be initiated to grow (shoot initiation); and (ii) at the time of ini-tiation,
shoot development (i.e. vegetative, mixed, or generative) is deter-mined
(induction). Although conditions for floral induction may be present prior to shoot
initiation, determination of that inductive condition in buds is not made until
initiation occurs. Initiation and induction events are regulated by different signals
and each may be manipulated by different stimuli. Removing the apical whorl of
leaves or tip pruning physiologically mature stems stimulates shoot initiation in
apical or lateral buds, respectively. If containerized plants are maintained in warm
temperatures (30°C day/25°C night) following initiation, vegetative shoot growth
is induced. If they are kept under cool conditions (18°C day/10°C night), initiating
shoots are induced to be generative. In either of the two temperature regimes
without pruning, they do not initiate shoots until the natural flushing event occurs
much later. They become vegetative or reproductive according to the tem-perature
at the time of shoot initiation. If transferred from cool to warm tem-peratures
before shoot initiation, new shoot growth is induced to be vegetative. Induction is
therefore determined at the time of shoot initiation, and plants rapidly lose their
floral inductive potential when removed from the cool envi-ronment.
Determination of shoot type can be reversed during morphogenesis by transferring
containerized trees from warm-to-cool or cool-to-warm con-ditions (Batten and
McConchie, 1995; Núñez-Elisea et al., 1996).

INITIATION CYCLE. The cyclic initiation of vegetative or reproductive shoots is common


to all mango cultivars. Developing vegetative shoots are rich sources of auxins and
gibberellins, which may be inhibitors in an internal cycle that regulates shoot initiation.
Auxins are actively transported basipe-tally to roots from production sites in stems
(Goldsmith, 1968; Cane and Wilkins, 1970; Wilkins and Cane, 1970; Goldsmith and
Ray, 1973), and they stimulate adventitious root growth in mango and other crops
(Hassig, 1974; Wightman et al., 1980; Sadhu and Bose, 1988; Rajan and Ram, 1989;
Núñez-Elisea et al., 1992). Elevated auxin synthesis in periodically flushing shoots is
likely to form a concentrated pulse of auxin, which inhibits recurring bud break and
moves basipetally to the roots. This pulse of elevated auxin may stimulate initiation of
new root flushes following each vegetative flush. Alteration of root and shoot growth
occurs in mango (Krishnamurthi et al., 1960; Cull, 1987, 1991; Parisot, 1988) and other
tropical and subtropical trees (Bevington and Castle, 1986; Williamson and Coston,
1989; Ploetz et al., 1991, 1993). Timing of the root flush may depend on the distance
from stems to roots, the physiological condition of the transport path, and
environmental conditions (i.e. temperature or water relations).

New roots that develop following growth stimulation are a primary source of
cytokinins (Davies, 1995). Cytokinins are transported passively to stems via the
xylem sap in all plants and are active in bud break (Went, 1943; Kende and Sitton,
1967; Sitton et al., 1967; Itai et al., 1973; Haberer and Kieber, 2002). Cytokinins
stimulate shoot initiation in mango (Chen, 1985; Núñez-Elisea et al., 1990) and
other plants (Oslund and Davenport, 1987; Belding and Young, 1989; Williamson
and Coston, 1989; Davenport, 1990; Davies, 1995;
130 T.L. Davenport

Henny, 1995). Auxin inhibits shoot initiation (Davies, 1995) and confers api-cal
dominance by preventing axillary bud break. Leaf-produced auxin and petiolar
auxin transport capacity declines as leaves age (Davenport et al., 1980). Auxin and
cytokinins may therefore be involved in the periodic cycle of bud break.

A critical balance of these two phytohormones, possibly modulated by GA 3,


may regulate shoot initiation. During a rest period, the inhibitory action of auxin
transported to buds decreases with time; whereas, cytokinin lev-els in buds increase
(Chen, 1987). When a critical cytokinin/auxin ratio is achieved, the buds are
stimulated to grow, thereby resetting the cycle with initiation of new shoots. The
interaction of auxin and cytokinin to control bud break in plants is a concept that is
supported by molecular studies (see review by Nordstrom et al., 2004). Moreover,
initiation of shoot growth occurs following pruning, defoliation or the application
of thidiazuron (Núñez-Elisea et al., 1990). Vigorous cultivars (Whiley et al., 1989)
and young, small trees under vegetatively promotive conditions flush frequently
with only short periods of rest; however, this cycle slows with age. Old centennial
trees flush infrequently (N. Golez, personal communication, the Philippines, 1989).

Foliar or soil-applied NO3− stimulates initiation of reproductive shoots only if


applied after resting stems have attained an age to overcome any veg-etatively
inductive influence. In contrast, high N in soils leads to high N lev-els in leaves
resulting in frequent vegetative flushes. The mechanism whereby NO 3− stimulates
shoot initiation is unknown.
Seeds are rich sources of auxin and gibberellins, which contribute to the strong
inhibition of bud break commonly observed on fruit-bearing mango stems. The
longer that fruit are attached to stems, the longer the postharvest inhibition may last
in the stem (Kulkarni and Rameshwar, 1989; Kulkarni, 1991).

Water stress inhibits shoot initiation by its direct impact on cell division and
elongation possibly by interfering with translocation of cytokinins from roots.
There is little evidence that water stress is directly involved in induc-tive processes.
During water stress, roots continue to grow and produce cyto-kinins (Itai and
Vaadia, 1965; Itai et al., 1968; Wu et al., 1994). Reduced xylem flux due to limited
soil hydration, and transpiration due to increased sto-matal resistance during water
stress may reduce the amount of cytokinins reaching stems. After rewatering, the
increased levels of cytokinins in roots may translocate to and accumulate in buds.
Auxin synthesis and transport from leaves are reduced during water stress
(Davenport et al., 1980) and may require several days for correction after
rewatering. This rapid shift in the cytokinin/auxin ratio of buds may explain the
shooting response that occurs soon after relief of water stress. GA 3 may act with
auxin to inhibit shoot ini-tiation (Davenport et al., 2001b). Early flowering in
plants treated with PBZ may be a response to lowered gibberellin levels, thus
lowering the level of initiation inhibitor.

This model could explain why sectors of tree canopies flush in the trop-ics.
Mango trees flush often and synchronously throughout the canopy when they are
young. With advancing age, the frequency of flushing is reduced
Reproductive Physiology 131

and synchrony is lost, resulting in sporadic flushes of vegetative or reproduc-tive


growth in sections of the canopy. As the distance between stems and roots
increases, the time required for transport of the putative pulses of ele-vated auxin
levels to roots, formed during a vegetative flush, is increased. Groups of stems
exhibiting simultaneous flushing ultimately connect to a common branch. Dye
trace studies indicate that water transport remains in strict phylotaxic alignment
from secondary roots to the canopy, even in large trees (T.L. Davenport,
unpublished results, Florida, 1991). Unless disturbed by girdling or by pruning of
branches or roots, specific branches in the can-opy communicate only with those
roots in phylotaxic alignment with them. The hormone transport time may vary
among sections of the canopy as the tree grows. This generates individual initiation
cycles in sections of the can-opy that are separately maintained unless
resynchronized with the rest of the tree following a canopy-wide environmental
trigger.
Synchronization of growth throughout trees occurs following exposure to low
temperature, water stress, light pruning of the entire tree and any condition that
would increase the postulated cytokinin/auxin ratio in buds throughout the canopy.
An increased ratio may occur by inhibiting auxin transport from leaves to buds, or
increasing cytokinin translocation from roots to stems. Winter in the subtropics
would reduce auxin transport; whereas, water stress in the tropics may impact the
availability of cytokinins from roots and auxin from leaves. The intensity of the
initiation response (i.e. synchronization of flushes in the canopy) may be regulated
by decreased auxin transport at low temperatures, the base level of which may be
deter-mined by the age of individual stems. Passage of a strong, extended cold
front during subtropical winters produces synchronized flowering. Milder winters
with weak cold fronts result in asynchronous flowering in sections of trees. The
oldest sectors of canopies flower first, followed by sectors bearing sequentially
younger flushes in subsequent cold fronts. Vegetative flushes occur when night
temperatures are > 18°C for significant periods between cold fronts.

INDUCTION SWITCH. Floral or vegetative induction is possibly governed by the interactive


ratio of a FP that is up-regulated in low temperatures to an age-regulated VP in leaves
at the time of shoot initiation. High FP:VP ratios would be conducive to induction of
generative shoots, low ratios conducive to veg-etative shoots and an intermediate ratio
of the two would be conducive to mixed shoots. Regardless of the endogenous levels of
the two components perceived in buds at the time of initiation, flowering and vegetative
growth responses can best be explained by the ratio of the two.

Although the putative FP seems to be up-regulated during leaf exposure to


cool temperatures (< 18°C), there appears to be a basal level present at all times in
leaves exposed to higher temperatures. Flowering of mango occurs in low-latitude
tropics lacking cool night temperatures when stems become sufficiently aged so
that the ratio of the basal level of resident FP to decreas-ing VP increases to a
critical threshold to provide floral induction when shoots are initiated. This could
explain how flowering on non-synchronized
132 T.L. Davenport

branches may occur at any time of the year in trees growing in low-latitude tropics.
High proportions of mixed shoots are commonly found in these con-ditions,
indicating the marginally floral-inductive ratios present under these conditions. In
contrast, flowering in younger stems having higher levels of VP is observed only
when initiation occurs in cool, floral-inductive tempera-tures. More flowering
occurs throughout the canopy when stems are exposed to cool temperatures,
attributable to the higher ratio of up-regulated FP to resident VP.

Genetic differences in base levels of the putative FP and/or VP or the receptors


of these components could explain the range in flowering responses in tropical and
subtropical cultivars and why a cultivar grown in an environ-ment different from
that in which it was selected is less productive. Cultivars selected in the subtropics
usually flower as well in the low-latitude tropics as those selected in the tropics.
Cool temperatures in the subtropics sometimes cause earlier flowering in tropical
cultivars than those selected in the sub-tropics. Kulkarni (1991) demonstrated that
several multi-flowering cultivars can induce flowering in receptor graft plants and
cause a range of the flower-ing response of the receivers to donors. Some cultivars
may produce higher base levels of putative FP than others. These are the same
cultivars that read-ily flower under warm temperatures and flower early during cool
winter months. The Comprehensive Conceptual Flowering Model suggests that
flowering can occur at any time in any cultivar regardless of origin so long as stems
are sufficiently old to reduce the VP level to below the critical FP/VP ratio when
initiation occurs.

Although the putative FP, perhaps a product of an ortholog of the Arabi-dopsis


FT gene, has not been identified, the VP may be a gibberellin. Triazoles and other
plant growth retardants that inhibit gibberellin synthesis, promote strong and out-
of-season flowering under conditions that would normally be marginally or non-
floral inductive.

PHOTOASSIMILATES. Photoassimilates produced by leaves provide carbohy-drates


essential for development of roots and other plant organs, including fruit. They are
either used immediately by the nearest sinks (Finazzo et al., 1994) or are stored in
locations throughout the tree to be used when demand for carbon resources exceeds the
existing photosynthetic supply (Whiley et al., 1988, 1989, 1991). A direct role for
carbohydrates in shoot initiation or induction is not part of this model, although they
facilitate mass flow in phloem from leaves to passively carry the FP to buds.

ALTERNATE BEARING. High levels of auxin and gibberellins produced in seeds possibly
inhibit shoot initiation on fruit-bearing stems for weeks or months following fruit
removal. Rapid production of new shoots following light pruning of fruit-bearing stems
after harvest indicates that residual levels of auxin and gibberellins linger only in the
rachis and last intercalary unit. If fruit are not set on the lingering rachis, there is less
inhibition. Heavy fruit set in 1 year impacts the timing of subsequent shoot initiation on
the large num-ber of fruit-bearing branches. Substantial delays in subsequent vegetative
Reproductive Physiology 133

flushes until close to the normal flowering period impact the flowering abil-ity of
young shoots. This may explain the occurrence of chronic alternate bearing in
some cultivars.

5.10 Floral Management


The Comprehensive Conceptual Flowering Model is consistent with growth and
development patterns of mango trees in the tropics and subtropics. It provides a
reasonable explanation for the various events in the phenologi-cal model of Cull,
predicts what will happen under a defined set of cir-cumstances and is being used
to develop strategies for consistent mango flowering. Flowering can be obtained at
any time of the year in a flowering management programme (Davenport, 2003).

The flowering management programme begins each season with tip pruning of
the entire canopy of orchard trees. Tip pruning can be done imme-diately after
harvest to move production forward in the following year or c.1–2 months after
harvest, depending upon cultivar, in order to achieve har-vest at the same time as
the previous year. If sufficient soil water is available at the time of pruning, a
vegetative flush will occur on all pruned stems c.1 month later. The number of new
shoots that will mature to become stems will be five- to eightfold greater than the
original number of pre-pruned stems due to initiation of many lateral vegetative
shoots on each stem. This increase in terminal stem number in the canopy will be
reflected in a concomitant increase in yield. The frequent flushes that can cause an
early second flush of vegetative growth tend to be suppressed.

The new stems must not flush a second time until at least 5 months after
pruning (Davenport, 2006). If they flush within 3–4 months after pruning, they will
be induced to be vegetative. Pre-prune leaf N levels in the stems must be 1.1–1.4%
in order to suppress a second flush of vegetative growth during the rainy season.
Mild water stress after the post-prune flush during the dry season will suppress a
second, undesired vegetative flush when leaf N levels are above the optimum
range. Pruning near the end of the dry sea-son in non-irrigated or furrow-irrigated
trees should be avoided. Transition from dry to wet season 2–3 months after
pruning causes a rain-stimulated vegetative flush prior to achieving sufficient age
of stems from the last flush. Test sprays of 4% KNO 3 on two to three
representative trees should be applied 5 months (for easily induced cultivars) and 6
months (for more dif-ficult to flower cultivars) after pruning. If no developing
shoots occur within 2 weeks, the spray is repeated. A flowering response is usually
evident after the second application. The other trees that were pruned on or near the
same date can then receive the foliar spray and will respond by synchronized flow-
ering. Although Davenport (2003) described the appropriate timing of PBZ in a
flowering management programme, it is not recommended because flowering can
be achieved without it. For orchard trees to be amenable to tip pruning, efficient
spray application of KNO3 and easy harvesting, they should be no taller than 4 m.
Pruning to rejuvenate large mango trees and
134 T.L. Davenport

properly shape trees for the annual flowering management programme is


recommended (Davenport, 2006).

5.11 Floral Biology


Detailed descriptions of generative shoots were reviewed in Davenport and Núñez-
Elisea (1997). Juliano and Cuevas (1932) described ovary devel-opment.

Sex ratio

Sex ratio (i.e. the proportion of perfect to staminate flowers) is a variable


component within panicles, trees and among cultivars. This ratio varies with
cultivar, but is usually < 50% (Davenport and Núñez-Elisea, 1997). Most per-fect
and staminate flowers are borne in the proximal portion of panicles due to their
architecture (Musahib-ud-din and Dinsa, 1946; Cobin, 1950). The variability in the
perfect/staminate flower ratio may be governed by physi-ological and
environmental conditions. Most studies indicate that although the total number of
flowers is substantially less in the distal half of panicles, there is a greater
proportion of perfect flowers in this region (Davenport and Núñez-Elisea, 1997);
however, this condition may be reversed in some culti-vars (Hussein et al., 1989).

Perfect flowers tend to form in the terminals of individual inflorescences while


staminate flowers are displayed in the earlier forming flowers located closer to the
panicle axis. When panicles begin to elongate in the lower inflo-rescences, only
staminate flowers form and the perfect flowers form at the terminus of each lateral
inflorescence. As more distally located lateral inflo-rescences begin elongation and
anthesis, they too first display staminate flowers before perfect flowers. These
inflorescences are progressively shorter than previously formed proximal
inflorescences, and there are fewer stami-nate flowers. The final vertical spike of
the panicle is composed almost exclu-sively of perfect flowers. Flowers abscise
soon after anthesis, thereby shifting the sex ratio. The sex ratio should include sex
type of all flowers in each panicle; however, sex ratios are normally determined at
some arbitrary moment during panicle development. Thus, the sex ratio is naturally
vari-able, increasing from extremely low to extremely high values so that timing of
observations during panicle development is critical.

Environmental determinants of sex ratio

Tropical cultivars yield poorly in the subtropics due to a small proportion of perfect
flowers on inflorescences (Singh and Singh, 1959; Singh et al., 1965; Singh, 1971).
Cool weather during inflorescence development contributes to fewer perfect
flowers (Naik and Mohan Rao, 1943; Singh et al., 1965, 1966).
Reproductive Physiology 135

Inflorescences that emerge during the middle and end of the flowering sea-son
produce two and seven times more perfect flowers, respectively, than the early
breaking inflorescences (Majumder and Mukherjee, 1961; Singh et al., 1966). This
response correlates with higher temperatures later in the flower-ing season. In
controlled-environment studies, low temperatures (15°C day/10°C night) reduced
the proportion of perfect flowers, particularly in tropical, polyembryonic cultivars
relative to subtropical, monoembryonic cultivars (Sukhvibul et al., 1999).

Physiological determinants of sex ratio

Endogenous factors affect the ratio of perfect to staminate flowers. Bajwa et al.
(1956), Majumder and Mukherjee (1961) and Joubert et al. (1993) reported that
lateral inflorescences on mixed shoots carried higher proportions of per-fect
flowers. Inflorescences on older trees produce higher proportions of per-fect
flowers than those on young trees (Naik and Mohan Rao, 1943; Majumder and
Mukherjee, 1961; Chacko and Randhawa, 1971; Pandey, 1989). This also occurs in
inflorescences borne on grafted compared with seedling trees (Musahib-ud-din and
Dinsa, 1946). The effect of tree maturity or rootstocks on sex ratio of flowers is not
understood. Panicles carried within the canopy of some cultivars (Majumder and
Mukherjee, 1961; Singh et al., 1966) or on particular sides of the canopy
(Mukherjee, 1953; Majumder and Mukherjee, 1961) have been reported to have
higher proportions of perfect flowers.
Application of some hormones and growth regulators alters the sex ratio of
inflorescences. GA3, applied at concentrations of 50–100 mg/l just prior to
inflorescence shoot initiation, substantially reduces the proportion of perfect
flowers (Maiti, 1973), as do combination sprays of urea (0, 3 and 6%) and GA3 (0,
15 and 30 mg/l) (Rajput and Singh, 1989). Soil-applied PBZ (10 g ai/tree)
significantly increases the ratio of perfect/staminate flowers (Kurian and Iyer,
1993a). Increases in floral ratio also occur with daminozide, whereas maleic
hydrazide either had no effect or lowered the ratio (Singh et al., 1965; Subhad-
rabandu, 1986). Foliar application of 50 mg/l BA with 2% calcium ion (Ca 2+)
increased the proportion of perfect flowers (Singh and Rajput, 1990). Naphtha-lene
acetic acid (NAA) at concentrations of 50, 100 and 200 mg/l increased the
perfect/staminate flower ratio (Mallik et al., 1959; Singh et al., 1965).
Other factors influencing sex ratios of inflorescences include stem age and
mineral nutrients. Gunjate et al. (1983), Desai et al. (1986) and Hussein et al.
(1989) reported that inflorescences from stems that grew at different times during
the previous summer/autumn period had significantly differ-ent perfect/staminate
flower ratios. Singh and Dhillon (1987) found that boron (B) levels affect sex ratio.

Sex ratios can be manipulated with growth regulators, but has no com-mercial
advantage (A.W. Whiley, personal communication, Queensland, 1996). Increases
in fruit yield resulting from chemically increased perfect/ staminate flower ratios
have not been observed, suggesting that perfect flower numbers are not the primary
limitation to crop performance (Schaffer
136 T.L. Davenport

et al., 1994). If only one or two fruits were set on each terminal, the tree would
carry an unusually heavy crop. It is unlikely that reduced perfect flower numbers
due to cool temperatures during inflorescence development is directly responsible
for poor fruit set and yields. Pollen viability, growth and ovule fertilization are
probably the main factors contributing to low fruit set under these conditions.

Anthesis and dehiscence

Floral anthesis generally occurs at night in polyembryonic cultivars (Wagle, 1929;


Torres, 1931; Galang and Lazo, 1937; Pimentel et al., 1984) and at night or early
morning in monoembryonic types (Popenoe, 1917; Musahib-ud-din and Dinsa,
1946; Singh, 1954a; Randhawa and Damodaran, 1961a, b). Subsequent dehiscence
of the four-lobed anthers occurs during the daylight morning hours revealing pale
blue pollen grains (Torres, 1931; Galang and Lazo, 1937; Mallik, 1957; L.B.
Singh, 1960). Anthesis and anther dehiscence are delayed by low temperatures or
overcast days (Singh, 1954a; De Wet and Robbertse, 1986a). Dehiscence is also
delayed by high RH, and pollination occurs primarily around midday (Mallik,
1957; Randhawa and Damodaran, 1961a, b).
Stigmas are receptive from c.18 h prior to anthesis to at least 72 h after
anthesis with optimum receptivity within 3 h from anthesis (Popenoe, 1917;
Wagle, 1929; Sen et al., 1946; Singh, 1954a; Spencer and Kennard, 1956; Rand-
hawa and Damodaran, 1961a, b; Gunjate et al., 1983; Pimentel et al., 1984;
Robbertse et al., 1994). Receptive stigmas are shiny and white-green, whereas non-
receptive stigmas are desiccated and yellow-brown. Pollen germination generally
occurs within 90 min of deposition, although the percentage ger-mination of pollen
deposited on stigmas is relatively poor (Singh, 1954a; S.N. Singh, 1961).
Pollination is initiated by the formation of two unusual protu-berances that meet to
form a bridge or ponticulus connecting the dorsal side of the ovule with the ovary
wall in line with the base of the style (Joel and Eisenstein, 1980). The ponticulus
may guide the elongating pollen tube to the ovule. Ovule fertilization occurs 48–72
h after pollination (S.N. Singh, 1961; Ram et al., 1976). Both zygote cell and
endosperm nuclear division appear to rest for about 2 weeks following pollination
despite cell division and growth of the ovary (Sharma and Singh, 1972; Ram et al.,
1976). A description of embryo development is presented by U.R. Singh (1961).
Application of B may improve stigma receptivity, pollen tube germination and
growth and ovule fertilization (De Wet and Robbertse, 1986b; Robbertse et al.,
1988; De Wet et al., 1989) as well as fruit development (Chen, 1979; Robbertse et
al., 1990); however, Jutamanee et al. (2002) could not verify the effect of B.

Pollen

Pollen grains are 20–45 Pm long and are oblong when dry and more spherical
when hydrated (Popenoe, 1917; Jivanna Rao, 1923; Bijhouwer, 1937;
Reproductive Physiology 137

Mukherjee, 1950; Singh, 1954a; Randhawa and Damodaran, 1961b; S.N. Singh,
1961). There are generally three equilateral, tapering furrows along the longitudinal
sides of dry pollen that give hydrated grains a roughly tri-angular shape when
viewed on end (Popenoe, 1917; Singh, 1954a; S.N. Singh, 1961; U.R. Singh and
A.P. Singh, 1973). Each furrow has a germpore in its centre (Mukherjee, 1950;
S.N. Singh, 1961). Anthers produce c.250–650 pollen grains with a mean of 410
grains per anther (Popenoe, 1917, 1920; Spencer and Kennard, 1955).

In vitro germination of mango pollen has been reported (Popenoe, 1917;


Spencer and Kennard, 1955; Young, 1958; Randhawa and Damodaran, 1961b;
S.N. Singh, 1961). Germination on stigmas was c.10% less than that on artificial
media, that is 78% across cultivars (Spencer and Kennard, 1955), although lower
rates of germination have been reported (Mukherjee, 1950). Mango pol-len is most
viable soon after anther dehiscence and rapidly degrades (Sen et al., 1946; Spencer
and Kennard, 1955; Mallik, 1957; S.N. Singh, 1963). Although the initial
percentage of viable pollen is generally  90% during warm weather (Popenoe,
1917; Mukherjee, 1949a, b; Singh, 1954a; S.N. Singh, 1961), cool tem-peratures
early in the flowering season result in abnormal, non-viable pollen grains
(Popenoe, 1917; U.R. Singh and A.P. Singh, 1973; Shu et al., 1989; Gazit et al.,
1992; Issarakraisila et al., 1992). The pre-vacuolate stage of meiosis during
microsporogenisis is the most sensitive period to temperatures < 10°C (Issara-
kraisila and Considine, 1994). Germination and pollen tube growth are reduced by
cool temperatures (S.N. Singh, 1961; Mullins, 1986; Robbertse et al., 1988;
Whiley et al., 1988; De Wet et al., 1989) and completely inhibited at tempera-tures
< 15C (Popenoe, 1917; Young, 1955; Sukhvibul et al., 2000).

Pollination

Pollination is a major yield-limiting constraint, due to the large number of flowers


on trees and low fruit set. Unlike polyembryonic cultivars, which produce nucellar
embryos, pollination is necessary for fruit set with mo-noembryonic cultivars
(Popenoe, 1917; Young, 1942; Spencer and Kennard, 1955; Gunjate et al., 1983;
Pimentel et al., 1984; Robbertse et al., 1994). Pollen compatibility within and
between cultivars has been widely investigated. Complete or partial self-
incompatibility has been reported (Mukherjee et al., 1961; Singh et al., 1962b;
Sharma and Singh, 1970, 1972; Ram et al., 1976; De Wet et al., 1989; Robbertse et
al., 1993). Incompatibility is evident by degen-eration of embryonic and nucellar
tissues and excessive loss of fruitlets. Cross incompatibility between some cultivars
has also been noted (Saha and Chhonkar, 1972; Ram et al., 1976; Robbertse et al.,
1993).

Wind
Early investigators concluded that the species is wind pollinated (Hartless, 1914).
Initially wet pollen dries to a powdery consistency on anthers soon after anthesis in
dry conditions (Pimentel et al., 1984), whence it is likely to be liberated in moving
air or via gravity to adjacent stigmas on the same and
138 T.L. Davenport

nearby flowers (Naik and Mohan Rao, 1943; Mallik, 1957). Singh (1954a) and
S.N. Singh (1961) suggested, however, that the amount of pollen moving in air
streams was too low for wind to be a pollination vector. They did not report the
location of pollen-collecting slides or take into account the close proximity of
flowers within inflorescences or numbers of open flowers in the canopy. Panicles
bagged to exclude pollinating insects were reported to set fruit (Free and Williams,
1976), which were retained to maturity, thereby con-firming that mango pollen can
be transferred by air movement or gravity (Bijhouwer, 1937; Mallik 1957). The
tacit assumption that open-pollinated flowers are exclusively crossed is likely to be
incorrect, although mango may favour cross-pollination.

Insect
Popenoe (1917) reasoned that pollen transfer occurs primarily within flowers by
insects. Panicles bagged to exclude insect visitation generally result in less fruit set
than on panicles in the open (Popenoe, 1917; Musahib-ud-din and Dinsa, 1946;
Mallik, 1957; Free and Williams, 1976; Jiron and Hedstrom, 1985). Insects
working mango flowers include Diptera, Hymenoptera, Lepidoptera and
Coleoptera (Popenoe, 1917; Simao and Maranhao, 1959; Randhawa and
Damodaran, 1961b; McGregor, 1974; Anderson et al., 1982; Jiron and Hed-strom,
1985). Flies of various genera are common on mango flowers (Pope-noe, 1917;
Burns and Prayag, 1921; Bijhouwer, 1937; Singh, 1954a; Spencer and Kennard,
1955; Eardley and Mansell, 1993). Polistes wasps are observed on mango flowers
but are considered to be ineffectual for pollen transfer (Spencer and Kennard,
1955; Free and Williams, 1976; Wolfenbarger, 1977). Honeybees (Hymenoptera)
are occasional visitors (Young, 1942; Simao and Maranhao, 1959; Smith, 1960;
Morton, 1964; Jiron and Hedstrom, 1985; Mac-Millan, 1991; Du Toit and Swart,
1993, 1994; Eardley and Mansell, 1993, 1994), but only if other more inviting
flowers are not present (Spencer and Kennard, 1955; Free and Williams, 1976;
McGregor, 1976). They are assumed to be the most effective pollinators of mango
and may be more effective if hives are placed in orchards during flowering (Du
Toit and Swart, 1993, 1994). Ander-son et al. (1982) recorded actual pollen
transfer on mango flowers by insects and found, in order of importance, the most
efficient pollinators to be wasps, bees, large ants and large flies.

With few exceptions (Mallik, 1957), pollen deposition rates are generally low
(Naik and Mohan Rao, 1943; Mukherjee, 1951). Differences in pollination rates
can be attributed to environmental conditions during flowering, differ-ing attraction
of insects to specific cultivars, proximity of more attractive flowering species or a
combination of the above. Young (1942) observed that insects visit only 10–12%
of available flowers. Depending on weather condi-tions, insect activity on mango
flowers is usually continuous from early morning to late afternoon, but nocturnal
activity of some species has also been reported (Jiron and Hedstrom, 1985).

The role of insects in cross-pollination is not understood. Anderson et al.


(1982) observed various insects carrying pollen to and from flowers and noted
pollination subsequent to those visits; however, they made no distinction
Reproductive Physiology 139

between actual pollen depositions by visiting insects and pollen transferred by


other means. Wester (1920) considered that pollination is facilitated by wind and to
a lesser extent by insects, and this conclusion is probably correct in most
environments. Self-pollination within flowers by insects while the pollen is still
damp is likely to occur. Use of isozyme (Degani et al., 1990; Rob-bertse et al.,
1993) and microsatellite DNA markers (Adato et al., 1995; Schnell et al., 2005) to
discern ratios of cross- versus self-pollinated fruitlets and off-spring is the most
accurate procedure to confirm self- and cross-pollination. Initial fruit set of
pollinated flowers is inconsequential since most of these fruitlets abscise before
reaching maturity (Lynch and Mustard, 1950; Singh, 1954a; Randhawa and
Demodaran, 1961a).

5.12 Fruit Development


Fruit growth is correlated with several growth regulating substances. Enlarge-ment
is sigmoidal reaching a constant size c.2–3 weeks before maturity (Singh, 1954a;
Randhawa and Damodaran, 1961a, b; Ram, 1983; Prakash and Ram, 1984). The
highest rates of fruit growth have been correlated with peak levels of putative
endogenous auxins found in seeds (Chacko et al., 1970a, b; Singh and Singh, 1974;
Chen, 1981; Ram, 1983; Prakash and Ram, 1984). Baghel et al. (1987a, b) reported
increased fruit mass with a combination spray of NAA and urea to pre-anthesis
panicles. Free and bound gibberellins, especially in seeds (Ogawa, 1963; Ram,
1983), peak similarly to putative aux-ins during fruit development (Chacko et al.,
1970c, 1972a; Ram and Pal, 1979; Chen, 1981). Cytokinins tentatively identified
as zeatin and zeatin riboside and other active fractions appear to vary in
concentration in seeds and peri-carp with two distinct peaks of activity. No
particular relationship to growth was evident (Ram, 1983; Ram et al., 1983). Seeds
produced more cytokinin activity than did the pericarp. In contrast, Chen (1983)
found only one peak of cytokinin activity in seeds and pulp occurring at about half
full size of both seed and pericarp. Seed tissues contain the highest cytokinin
activity. Levels of endogenous auxin, gibberellins and cytokinins in leaves during
fruit set were compared to production in leaves during other periods with-out
conclusive results (Paulas and Shanmugavelu, 1989).

5.13 Stenospermocarpy
Abscission of non-fertilized and fertilized flowers is normal. Fruitlet abscis-sion
from pea size on is often associated with embryo abortion (Chandler, 1958; U.R.
Singh, 1961; Singh, 1964; Lakshminarayana and Aguilar, 1975; Ram et al., 1976)
and is referred to as stenospermocarpy (Soule, 1985). Steno-spermocarpy in mango
is unusual (Chacko and Singh, 1969a) but occurs regularly in some cultivars
(Núñez-Elisea and Davenport, 1983; Whiley et al., 1988). Stenospermocarpic
fruitlets have slower growth rates than seeded fruit, generally become misshapen
and fail to reach full size.
140 T.L. Davenport

Stenospermocarpy in some cultivars has been correlated with low tem-


peratures during flowering and early fruit set in the subtropics (Lakshmina-rayana
and Aguilar, 1975; Young and Sauls, 1979; Whiley et al., 1988; Schaffer et al.,
1994). Núñez-Elisea and Davenport (1983) reported that stenospermo-carpic fruit
often occur distal to seeded fruitlets within panicles and sug-gested that embryo
abortion is associated with high temperatures when these latter fruit set. Secondary
spring flowering of some monoembryonic culti-vars under high temperatures has
resulted in high proportions of steno-spermocarpic fruit (E.K. Chacko, personal
communication, Australia, 1995). Application of auxins, gibberellins and
cytokinins produce seedless fruit in some cultivars, suggesting that the abscission
zone is protected by these hor-mones despite the loss of the endogenous supply
from the aborted seed (Ven-kataratnam, 1949; Chacko and Singh, 1969a, b;
Kulkarni and Rameshwar, 1978).

5.14 Fruit Set and Retention


Fruit set and retention of mango was recently reviewed by Singh et al. (2005).
Abscission of flowers and fruitlets is accomplished by rapid formation of a
separation layer in the abscission zone in the pedicel-peduncle junction (Bar-nell,
1939). U.R. Singh (1961) described formation of the abscission zone dur-ing floral
ontogeny and of the separation layer during abscission of flowers and fruitlets. The
majority of panicles lose all fruitlets (Núñez-Elisea and Davenport, 1983). The
pattern of fruitlet abscission is asymptotic with the greatest losses occurring during
the first weeks after anthesis (Núñez-Elisea and Davenport, 1983; Prakash and
Ram, 1984; Searle et al., 1995). Except for the tendency to retain fruit in the distal
portion of panicles, abscission of flowers and fruitlets is random. It can involve
fruitlets regardless of size or location.

Of the 8–13% of perfect flowers setting fruit, < 1% reach maturity (Bijhou-
wer, 1937; Sen, 1939; Naik and Mohan Rao, 1943; Mukherjee, 1949b; U.R. Singh,
1960; Randhawa and Damodaran, 1961a; Singh, 1978; Gunjate et al., 1983;
Prakash and Ram, 1984). Generally, most fruit are set on the most distal spike
portion of panicles (Chadha and Singh, 1963; Núñez-Elisea and Daven-port, 1983).
Fruit loss has been associated with embryo abortion, resulting in blackened or
shrivelled embryos (Singh, 1954a, 1964; Chandler, 1958; U.R. Singh, 1961;
Sharma and Singh, 1972; Ram et al., 1976) after the fruit is sepa-rated from the
tree (Núñez-Elisea and Davenport, 1983).

Sex ratio

The perfect/staminate floral ratio in panicles may influence fruit set and pro-
ductivity (Naik and Mohan Rao, 1943; Singh, 1954b; Singh and Singh, 1959; U.R.
Singh, 1960). Mallik (1957) noted that more perfect flowers are formed in ‘on’
than ‘off’ years of alternate-bearing cultivars. Other studies, however,
Reproductive Physiology 141

have demonstrated that the number of perfect flowers does not correlate with
subsequent yield (Randhawa and Damodaran, 1961a) so long as the proportion of
perfect flowers is not < 4% (Singh, 1964, 1971). Most fruit are borne in the distal
portion of panicles (Shawky et al., 1977), which may be correlated with the high
ratio of perfect to staminate flowers there. Schole-field and Oag (1984) estimated
that one mature fruit is harvested for each 169 perfect flowers in the distal half of
the panicle; whereas 592 perfect flowers are required to produce one fruit in the
proximal half. Therefore, intrinsic factors other than sex ratio regulate fruit set.

Mineral nutrients

Boron is one of seven micronutrients required for normal plant growth. The
physiological function of B is unknown (Hu and Brown, 1994), although it is
essential for floral development, pollen germination, pollen tube growth, embryo
development and growth of organs (i.e. fruit) (Vasil, 1963; Agarwala et al., 1981;
Dell and Huang, 1997; Shorrocks, 1997). Deficient soils are com-monly found in
mango-producing areas of Australia, Thailand, Central and South America and
Africa where symptoms are common (Aitken et al., 1987; Singh et al., 2005).
Boron applications to deficient mango trees increase normal fruit set (Robbertse et
al., 1990; Raja et al., 2005). Fruitlet abscission in mangoes has also been attributed
to zinc (Zn) deficiency (Jiron and Hedstrom, 1985).

Hormonal control

Auxin
Research demonstrating improved fruit set and retention following applica-tion of
several auxin analogues to pre-anthesis panicles or to panicles bearing fruitlets of
various sizes has been reviewed (Davenport and Núñez-Elisea, 1997; Singh et al.,
2005). NAA is the most effective auxin analogue for improv-ing fruit retention
(Prakash and Ram, 1986; Khan et al., 1993). Initial fruit set was substantially
increased when sprays of 200 mg/l indole acetic acid (IAA) were applied to
developing panicles (Singh et al., 1965). A 300–400% increase in fruit set resulted
when NAA (40 or 50 mg/l) was sprayed at the pre-anthesis stage (Ram, 1983;
Singh and Ram, 1983; Prakash and Ram, 1986). Chen (1981) reported no effect on
fruit retention when 5 mg/l of either naphthaleneacet-amide or β-naphthoxoyacetic
acid were applied three times at 2-week inter-vals to panicles in which fruit had
reached 4 mm in diameter.
Despite increased fruit retention of mango using exogenous applications of
auxins, few studies have examined endogenous auxins in fruit as related to
retention (Chacko et al., 1970a, b; Ram et al., 1983; Prakash and Ram, 1984).
Singh and Singh (1974) were unable to detect significant differences in endog-
enous auxins or inhibitors when comparing alternate and regular bearing cultivars.
Chen (1981) observed lower levels of auxin-like substances in
142 T.L. Davenport

mesocarp and calyx tissues of abscised fruits than those of intact fruits. Simi-lar
decreases in auxin and gibberellins with an increase in abscisic acid as fruitlets
abscised were reported by Bains et al. (1999). The interaction of auxin in fruit and
abscission zones to maintain mango fruit retention is not clear.
Continuous auxin synthesis and basipetal transport to the abscission zone is
critical for maintenance of plant organs, including fruit (Crane, 1964; Nitsch, 1965;
Morgan et al., 1977; Davenport et al., 1980; Roberts and Osborne, 1981). Increased
mango fruit set and retention in response to exogenously applied auxins confirms
this requirement; however, other hormonal factors also appear to be involved.
Developing seeds are rich sources of all the known classes of phytohormones,
including auxins (Crane, 1964; Nitsch, 1965; Chacko et al., 1970a, b, c; Chen,
1981). Hence, exogenous enrichment of auxin in the presence of other seed-
produced phytohormones facilitates increased fruit retention. In contrast, NAA (10
and 20 mg/l) spray-applied to bagged, self-pollinated flowers, does not result in
development of stenospermocarpic fruits beyond the marble size (Venkataratnam,
1949; Chacko and Singh, 1969a, b). Similarly, applications of 250 or 500 mg/l GA 3
or 250 mg/l BA alone to panicles does not promote production of
stenospermocarpic fruits (Chacko and Singh, 1969a, b). Supplying exogenous β-
naphthoxyacetic acid (10 mg/l), BA (250 mg/l) and GA3 (250 and 500 mg/l)
together in multiple sprays until half grown, however, resulted in retention of
several seedless fruit to maturity. Chen (1983) and Oosthuyse (1995b) observed
that gibberel-lin, cytokinin and auxin reduce fruit drop of open-pollinated fruitlets
of some cultivars. Thus, although auxin is important for maintaining the abscission
zone, the presence of other phytohormones appears to be important for fruit-let
development (Chacko et al., 1970a, b; Ram, 1983; Ram et al., 1983).

Cytokinins
Although cytokinins are not generally thought to be associated directly with
abscission, Ram (1983) and Ram et al. (1983) concluded that low cytokinin levels
during fruit development might contribute to fruit loss. Chen (1983) observed a
correlation of low cytokinin levels in stenospermocarpic fruits with abscission at
the marble stage of growth. Application of 250 mg/l BA to bagged panicles does
not promote production of seedless fruits (Chacko and Singh, 1969a, b). The
synthetic cytokinin, N-(2-chloro-4-pyridyl)-N’-phenylurea (CPPU) also does not
improve fruit set when applied alone at a rate of 10 mg/l to post-anthesis panicles
(Oosthuyse, 1995b). The role of cytokinins in separation events remains
inconclusive.

Gibberellins
Gibberellins do not appear to be directly linked to the onset of abscission (Chacko
et al., 1970c, 1972a; Ram and Pal, 1979; Chen, 1981; Ram, 1983). Spray
applications of GA3 to pre- and post-anthesis panicles to increase fruit set and
retention have been inconsistent. Increased yield (Teaotia et al., 1967; Singh and
Ram, 1983; Rajput and Singh, 1989) and production of seedless fruit (Kulkarni and
Rameshwar, 1978) have been reported from these treatments, but Chacko and
Singh (1969a, b) observed no such effects. Chen (1983) and
Reproductive Physiology 143

Oosthuyse (1995b) investigated the effects of several foliar applications of GA3


starting at the 4 mm diameter stage, but were unable to improve fruit set.
Several classes of gibberellin-synthesis inhibitors have been tested for
reducing fruit drop. The growth retardants, daminozide and cycocel, increased fruit
set when applied to post-anthesis panicles (Singh and Ram, 1983). The authors
suggested that increased fruit retention might have been mediated through
increased cytokinin-like activity of the growth retardants. Although initial fruit set
was promoted by PBZ, yield was not improved (Kurian and Iyer, 1993a). It is not
clear whether the contrasting results of increased yield (Kurian and Iyer, 1993b)
were due to reduced fruit loss or more intense flow-ering in response to treatment.
Goguey (1990) reported increased fruit set and retention using soil-applied PBZ at
5 g ai/tree. Spray application of uni-conazole, a more biologically active triazole
(500–2000 mg/l), reportedly increased fruit set and yield (Galila and El-Masry,
1991). It is difficult to resolve the contradictory results demonstrating enhancement
of fruit reten-tion by GA3 and inhibitors of its synthesis.

Inhibitors
Abscisic acid (ABA) is possibly involved in fruitlet abscission. Although cor-
relations exist between certain inhibitors and abscission of mango fruitlets, no clear
cause and effect relationships have been established. Fruit drop was correlated with
levels of an acidic inhibitor, possibly ABA (Chacko et al., 1970b, 1972a; Singh
and Singh, 1974; Ram, 1983; Prakash and Ram, 1984). Chen (1981) reported
similar changes in putative ABA with maximum levels occurring during early fruit
drop and with advancing age of fruits. Putative ABA levels in abscised and
retained fruits were compared and were highest in the calyx and mesocarp of
abscised fruitlets.

Ethylene
Ethylene has the greatest immediate impact on flower and fruitlet abscission. Van
Lelyveld and Nel (1982) reported higher levels of ethylene in abscised fruitlets
compared with those retained on trees. Núñez-Elisea and Davenport (1983, 1984,
1986) examined the dynamics of ethylene production in intact and excised fruitlets
from onset to separation. Increased production began in explants about 26 h
postharvest and increased logarithmically until fruit sep-aration. Abscission of the
fruitlets began 48 h after the onset of enhanced ethylene production. Similar results
with avocado fruitlet abscission experi-ments (Davenport and Manners, 1982)
indicate that the onset of ethylene production in intact fruitlets is spontaneous in
individual fruitlets followed by abscission 48 h later. The pericarp provided the
bulk of ethylene for induc-tion of abscission processes; the pedicel produced no
ethylene. There was reduced fruit drop in response to inhibitors of ethylene
production and action (Singh and Ram, 1983; Naqvi et al., 1990, 1992). Whereas
increased peroxi-dase (Van Lelyveld, 1978) and polyphenol oxidase activities have
been reported in abscissed mango fruitlets (Van Lelyveld and Nel, 1982), Núñez-
Elisea and Davenport (1984) observed no changes in peroxidase activity or protein
levels prior to separation of fruitlets.
144 T.L. Davenport

Abscission of stenospermocarpic fruits has been associated with small


increments in ethylene production (Núñez-Elisea and Davenport, 1983). Sen-
sitivity to low levels of endogenous ethylene may reflect the absence of seed-
produced auxins. Protection of the abscission zone depends on a constant supply of
auxin, and ethylene production levels in tissues correlate with endogenous auxin
levels (Roberts and Osborne, 1981).
Despite their roles in cell division, cell enlargement and maintenance of the
abscission zone in developing fruit, a specific recommendation for exog-enous
application of plant growth regulators, either alone or in combination, to improve
yield of mango has not been adopted. Phytohormones have little residual effect on
fruit development, and multiple applications of products to counteract the short-
term responses are prohibitively expensive.

Photoassimilates
Wolstenholme and Whiley (1995) discussed the ecophysiology of the mango as a
basis for preharvest management. They proposed that the adaptive sur-vival
strategies of the mango explain its notoriously poor cropping perfor-mance.
Mechanisms that impart tolerance to heat, drought and flood stresses, which the
tree has developed for survival in harsh environments, have come at considerable
carbon cost with the resultant diversion of photoassimilate resources away from
fruiting.
There is abundant evidence that heavy cropping in tree crops exhausts stored
reserves (Jones et al., 1975; Kaiser and Wolstenholme, 1994; Whiley et al., 1996)
and that current photosynthate is often unable to satisfy the demands of fruit set
and fruit growth after heavy and prolonged flowering (Chacko et al., 1982). There
are significant genotypic differences in photoassimilation rates between low- and
high-yielding cultivars growing in both the tropics and the subtropics of Australia
(Chacko et al., 1995; Searle et al., 1995). At each location, photoassimilation rates
were considerably greater on the higher-yielding cultivar, and this difference was
maintained from flowering through to fruit maturation. 14C studies during the fruit
set and abscission period also demonstrated strong discrimination in the movement
of assimilates, which was dominated by randomly located fruit on panicles of the
low-yielding cultivar (Chacko et al., 1995). In contrast, assimilate discrimination to
fruitlets was less severe in the high-yielding cultivar with a more even distribution
of photoassimilates. It was concluded that the availability and distribution of
photoassimilates during the fruit set and establishment stages was largely
responsible for the yield differences between the cultivars.

Supporting evidence for the role of photoassimilates in fruit set and retention
also comes from enrichment studies (Schaffer et al., 1999). Container-grown plants
that flowered in the open were transferred to controlled-environment glasshouse
rooms immediately after the completion of anthesis. Temperatures were 28°C
day/20°C night while the atmospheric carbon diox-ide (CO2) concentrations were
350 or 600 Pmol/mol. Photoassimilation of trees in the CO2-enriched rooms was
approximately 60% greater than those held at partial pressures of 350 Pmol/mol
CO2. Fruit retention and final yield were significantly higher on those trees grown
at the partial pressure of
Reproductive Physiology 145

600 μmol/mol CO2. Higher levels of available assimilates during the fruiting cycle
appear to benefit fruit retention and yield.

5.15 Alternate Bearing


Alternate bearing of certain mango cultivars has plagued growers (Hartless, 1914;
Rao, 1997). Lack of production in the ‘off’ year is usually a result of lack of floral
initiation (R.N. Singh, 1959), and has been reviewed (Mukherjee, 1953; Gangolly
et al., 1957; L.B. Singh, 1960, 1972; Chadha and Pal, 1986; Pan-dey, 1989). Shoot
initiation and induction described in this chapter perhaps offers a clearer
understanding of this phenomenon.

5.16 Conclusions
This chapter has provided a comprehensive review of investigations of vari-ous
factors potentially involved in mango flowering, fruit set and retention. Many of
the reports cited are contradictory. Such variable results reflect: (i) the different
experimental approaches utilized, especially in field experi-ments; (ii) the range of
environments in which experiments have been con-ducted; and (iii) differences in
the responses of cultivars to treatments. As a consequence, it is difficult to draw
unambiguous conclusions with respect to the role of specific factors on flowering,
fruit set and retention. More research is clearly needed in these areas, particularly
in controlled environments. For example, although KNO 3 is utilized with great
success to stimulate flowering in tropical conditions, confusion remains as to
whether it effects initiation or floral induction. Future studies on flowering research
should include results of the proportion of stems that remain in rest and those that
produce vegeta-tive shoots as well as the proportion of reproductive shoots.
Providing this information allows an analysis of the impact of treatments on
initiation and inductive events.

This chapter also presents several hypothetical models for shoot devel-opment
and flowering. Each of the proposed models consists of a series of hypotheses that
invite further study to test their validity. Future research should challenge these
models so that flowering and crop yield can be better understood in both the tropics
and the subtropics.

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6 Ecophysiology

B. Schaffer,1 L. Urban,2 P. Lu3 and A.W. Whiley4


1
University of Florida, Florida, USA
2
Centre INRA de Corse, San Giuliano, France
3
EWL Sciences, PO Box 39443, Winnellie, Northern Territory, Australia
4
Sunshine Coast Horticultural Services Pty Ltd, Nambour, Queensland, Australia

6.1 Introduction 170


6.2 Photosynthesis 172
Introduction 172
Light 177
Leaf temperature 179
Elevated atmospheric CO2 concentration 181
Humidity 182
Flooding 182
Internal factors 184
Photosynthetic contributions by fruit 187
6.3 Plant Water Relations 188
6.4 Tree Growth and Development 190
Light 190
Temperature 192
Drought 193
Flooding 194
Wind 195
Salinity 196
Elevated atmospheric CO2 concentration 197
6.5 Crop Production 198
Temperature limitations to crop production 198
Light interception and orchard design 199
6.6 Conclusions 200

6.1 Introduction
The genetic composition of mango cultivars is the primary determinant of yield
potential. However, actual yield, as well as tree growth and develop-ment, are
mediated by several endogenous factors including previous fruit load, postharvest
vegetative growth, preflowering maturity of terminal shoots,
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
170 (ed. R.E. Litz)
Ecophysiology 171

production and mobilization of carbohydrates, nutritional status, plant growth


substances and carbon to nitrogen ratios (Schaffer et al., 1994). These factors are
either directly or indirectly affected by environmental variables such as light,
temperature and water availability. Environmental conditions outside the range for
optimum growth may also impose stress which results in physiological changes
that reduce growth or cause permanent damage to mango trees. For example, major
climatic events (i.e. extended drought, floods, wind storms, heat waves and freezes)
can cause severe damage to crops due to development of excessive stress.
However, mediated stress and the release from stress imposed by normal seasonal
changes provide condi-tions that result in the progression of cropping cycles due to
phenological changes in the plants. An example of beneficial stress in mango is the
improved synchrony and reliability of flowering in subtropical climates due to cool
winter temperatures. Thus, understanding the impact of the environ-ment on tree
physiology and growth, and the particular adaptive strategies developed through
the processes of evolution, can provide a framework to manage the crop to
maximize genetic yield potential (Schaffer et al., 1994).

Physiological responses of mango to environmental variables can be related to


the evolutionary centre of origin of a specific cultivar. Mango culti-vars are
classified into two ecotypes based on embryony (see Mukherjee and Litz, Chapter
1, this volume). A race with a single zygotic seed, monoembry-onic types, evolved
in the dry subtropical, monsoonal regions of the Indian subcontinent with very hot
summers but cooler winters. The polyembryonic types, produced through nucellar
embryony, largely evolved in the consis-tently hot, humid tropics of South-east
Asia where the monsoonal pattern still predominates but the dry season is shorter
than that of the Indian sub-continent (Mukherjee, 1972). Hybridization occurs
freely within and between the two ecotypes and has led to a proliferation of
cultivars of widely varying genetic composition. Differences in growth and
flowering responses to tem-perature have been observed between the two
embryonic ecotypes (Whiley et al., 1989) and selection and breeding offer
potential for increasing the crop-ping performance of this notoriously low-yielding
and recalcitrant tree across a wider range of environmental conditions (Schaffer et
al., 1994).
Inevitably, the many unique features of the mango tree represent its evo-
lutionary response to an indigenous environment that is particularly hostile, with
sustained extreme heat and high evaporative demand for much of the year. This
chapter provides an overview of the impact of environmental fac-tors on
physiology, growth and productivity of mango. Plant responses will be considered
in the context of the evolutionary origin and adaptability of the mango tree.
Responses to light, temperature and water are emphasized, while the effects of
atmospheric carbon dioxide (CO2) concentration, wind and salinity are also
discussed.
Photosynthesis and plant water relations are closely associated with
environmental conditions and directly affect plant growth and productivity. The
principles of leaf gas exchange and plant water relations are discussed to provide a
theoretical basis for interpreting physiological responses to the environment. The
impact of photosynthesis and tree water relations on
172 B. Schaffer et al.

growth and development under different environmental conditions is dis-cussed.


Pollination, fertilization, flowering and fruit set, which are strongly influenced by
environmental factors, have been addressed elsewhere (see Davenport, Chapter 5,
this volume; Schaffer et al., 1994). In the final sec-tion of this chapter, mango crop
production is integrated with aspects of ecophysiology.

6.2 Photosynthesis

Introduction

The net CO2 assimilation rate (Anet) in C3 plants is a function of the carboxyla-tion
rate (Vc), the oxygenation rate (Vo) and the rate of CO2 evolution in light that
results from processes other than photorespiration, sometimes called ‘day
respiration’ (Rd):
Anet = Vc – 0.5Vo – Rd (6.1)
Rd is usually inferred from measurements of leaf CO 2 exchanges after 5 min in the
dark (i.e. ‘night respiration’ Rn). However, it has been repeatedly shown that Rd is
lower than Rn (see Atkin et al. (2000) for review), so that light is known to inhibit
respiration, with a Rd/Rn value ranging from 30 to 100% (see Peisker and Apel
(2001) for review). Urban et al. (2008) established the following linear regression
for Rd of mango leaves: Rd = 0.35Rn – 0.21, which may be used to infer Rd from
Rn for photosynthetic photon flux (Q) values above 170 Pmol photons/m2/s.
Currently, modelling of Anet often uses the Harley et al. (1992) version of the
Farquhar et al. (1980) model. According to this model, Anet can be expressed as:

Anet = (1 – 0.5O/(WCi))min(Wc, Wj, Wp) – Rd (6.2)


where O represents the partial pressure of oxygen (O2) in the intercellular air
spaces (Pa), W the specificity factor of ribulose-1,5-bisphosphate carboxylase/
oxygenase (Rubisco). Ci is the partial pressure of CO2 in the intercellular air
spaces (Pa), Wc the carboxylation rate limited by the amount, activation state or
kinetic properties of Rubisco (Pmol CO2/m2/s), Wj the carboxylation rate limited
by the rate of ribulose bisphosphate regeneration (Pmol CO2/m2/s), and Wp the
carboxylation rate limited by triose phosphate utilization in sucrose and starch
synthesis (Pmol CO2/m2/s).
Usually O is set as 21 kPa (21%). The variable W, which characterizes the
ratio of the affinities of CO2 and O2 for ribulose-1,5-bisphosphate in the active site
of Rubisco, can be calculated from the CO2 compensation point ** (the CO2
concentration at which photosynthesis equilibrates with respiration):
W = 0.5O/* * (6.3)
where W = 2220 mol CO2/mol O2 at 25C for ‘Cogshall’ mango leaves (Urban et
al., 2008), which is lower than those given by Epron et al. (1995): 2100–2900
Ecophysiology 173

mol CO2/mol O2. Rubisco’s large subunit is encoded by a single gene in the
chloroplast genome, and no post-transcriptional modifications have been
discovered so far. It is thus very unlikely that W can change in the short term
(Spreitzer and Salvucci, 2002).
The internal partial pressure of CO2 (Ci) is one of the two major variables of
photosynthesis (with the photosynthetically active photon flux density). It may be
calculated from the supply function:
Ci = Ca – Anet/gb – Anet/gs (6.4)
where Ca is the partial pressure of CO2 (Pa) in ambient air, gb represents the leaf
boundary layer conductance (mol H2O/m2/s), and gs is the stomatal conductance of
water (H2O) (mol H2O/m2/s).
Stomatal conductance is the major factor controlling Anet. It ranges from
c.0.02 to c.0.4 mol H2O/m2/s in ‘Cogshall’ mango leaves and may be linearly
related to Anet (Urban et al., 2002, 2003, 2006). The slope of the relationship
between gs and Anet however is affected by drought (Fig. 6.1). Variations in the
slope of this relationship reflect changes in photosynthetic water use effi-ciency
and are not well understood.
It must be stressed that using Ci as the driving variable of photosynthesis is
much debated. It has been advocated that Cc, the partial pressure of CO2 at the site
of carboxylation, should be utilized instead. Using Ci implies that the following
assumptions have been made: Cc = Ci and gm = 0, where gm repre-sents mesophyll
conductance, also called liquid phase resistance, which

0.50

0.45

0.40

0.35
Wet
gs (mol H2O/m2/s)

0.30 y = 0.025x + 0.028


R2 = 0.86
0.25

0.20

0.15
Dry
0.10 y = 0.009x + 0.024
0.05 R2 = 0.695

0
0 5 10 15 20
Anet (μmol CO2/m2/s)

Fig. 6.1. The relationship between stomatal conductance (gs) and net photosynthesis
(Anet) in mango leaves from well-irrigated (■) and drought-stressed (▲) 12-year-old
‘Cogshall’ trees (Source: redrawn from Urban et al., 2006).
174 B. Schaffer et al.

encompasses diffusion from the intercellular leaf spaces to the carboxylation sites
in the chloroplasts. There is a growing body of evidence that gm is not negligible in
most species. The average value of gm in unstressed mango leaves (0.21 Pmol
CO2/m2/s) (Urban et al., 2008), calculated using the method of Epron et al. (1995),
is within the range of values for broadleaf species sur-veyed by Ethier and
Livingston (2004) and Manter and Kerrigan (2004). The carboxylation rate (in Eqn
6.2) limited by the amount, activation state or kinetic properties of Rubisco (Wc)
can be calculated as:
Wc = VcmaxCi/(Ci + Kc(1 + O/Ko)) (6.5)
where Vcmax represents the maximum rate of carboxylation (Pmol CO2/ m2/s),
and Kc (Pa CO2) and Ko (Pa O2) are the Michaelis constants of Rubisco
carboxylation and oxygenation, respectively. The Vcmax values of well-exposed
mango leaves at a leaf temperature of 30C are typically in the range of 80–100
Pmol CO2/m2/s (Urban et al., 2006). Specific values of Kc and Ko for mango
leaves have not been estimated and are approximated using data from other species
(i.e. cotton or tobacco).
The carboxylation rate limited by the rate of ribulose bisphosphate regen-
eration (Wj) is controlled by the rate of electron flow J (Pmol electrons/m2/s):
Wj = JCi/(4(Ci + O/W)) (6.6)
with
J = DT Q/(1 + D2T 2Q2/J )
2 0.5 (6.7)
max
where Q is the photosynthetically active photon flux density (Pmol quanta/ m2/s),
T represents leaf absorbance (no units), D is the apparent efficiency of light energy
conversion (mol electrons/mol photons) and Jmax is the light-saturated rate of
electron transport (Pmol electrons/m2/s). Leaf absorbance of mango leaves,
measured from 390–760 nm using an integrating sphere, was found to be close to
0.81 (Urban et al., 2008) and is in the normal range of T values of the literature
(Bauerle et al., 2004). Leaf absorbance, which is pos-itively correlated with leaf
chlorophyll content, may increase as a conse-quence of paclobutrazol treatments
(Gonzalez and Blaikie, 2003). The apparent efficiency of light energy conversion
in mango reaches 0.32 Pmol electrons/ Pmol photons (Urban et al., 2004b), in the
absence of photoinhibition or pho-todamage. This value corresponds to the mean
value of operational D (Sin-gaas et al., 2001). The Jmax values of well-exposed
mango leaves at a leaf temperature of 30C are typically in the 120–150 Pmol
CO2/m2/s range. The
Jmax as well as the Vcmax values are rather low when compared to values from
other species and partly explain why maximal rates of leaf photosynthesis
(Amax) are rather low, typically 12–15 Pmol CO2/m2/s.
The carboxylation rate limited by triose phosphate utilization during sucrose
and starch synthesis (Wp in Equation 2), can be calculated by:
Wp = 3TPU + Vo/2 = 3TPU + Vc0.5Ci /(CiW) (6.8)
where TPU is the rate of phosphate release in triose phosphate utilization during
starch and sucrose production. The TPU is usually not included in
Ecophysiology 175

most studies on photosynthetic capacity because of methodological difficul-ties.


However, Urban et al. (2003) found that TPU = 8–12 Pmol CO2/m2/s at a leaf
temperature of 30C in well-exposed ‘Cogshall’ mango leaves.
The variables Vcmax and Jmax are temperature dependent and their depen-
dency is described by:
Parameter (Vcmax, Jmax, TPU) = exp(c – 'Ha/(RTl))
/(1+exp(('STl – 'Hd)/(RTl))) (6.9)
where c is a scaling factor, 'Ha (J/mol) the activation energy of the given
parameter, R the gas constant (8.3143 J/K/mol), Tl (K) the leaf temperature, 'S
(J/mol) an entropy term and 'Hd (J/mol) the deactivation energy of the given
parameter.
Similarly, the temperature dependency of Rd, W, Kc and Ko is described
by:
Parameter (Rd, W, Kc, Ko) = exp(c – 'Ha/(RTl)) (6.10)
Proteins of the Calvin cycle and thylakoids represent the majority of leaf nitrogen
(N). Therefore, photosynthetic capacity is strongly related to leaf N content
expressed on an area basis (Na) (Field and Mooney, 1986; Evans, 1989; Kellomäki
and Wang, 1997; Walcroft et al., 1997). To account for the relation-ship commonly
observed between the parameters defining photosynthetic
capacity (Vcmax, Jmax, TPU and Rd mainly) and Na (Field and Mooney, 1983;
Harley et al., 1992) (Fig. 6.2), scaling factors c of Vcmax, Jmax, TPU and Rd may
be related to Na, either linearly or slightly non-linearly.
In summary, leaf net photosynthesis depends on five major classes of factors,
either variables (external or internal factors) or parameters (more or less constant
factors), provided that plants are not exposed to too extreme conditions; we may
consider the internal factors as genetic factors. The five classes of factors are:

1. The photosynthetically active photon flux density (Q), which is the major
driving variable of photosynthesis. Gross photosynthesis is determined by Q while
Ci determines the proportion of photorespiration, and thus net photosynthesis. One
of the major environmental factors affecting Ci is water availability in the root zone
through its effect on gs.
2. Leaf nitrogen concentration (Na), which is not a rate-determining factor of
photosynthesis, unlike Q, but may be considered as a rate-limiting factor. In other
words, Na sets the photosynthetic potential of a leaf (i.e. photosyn-thetic capacity).
We shall see below which factors influence Na in mango leaves.
3. Leaf temperature influences leaf photosynthesis. Net photosynthesis is
positively correlated with leaf temperature in a normal range. Leaf tempera-ture
(Tl) is not a driving variable of photosynthesis but it is the single most important
rate-determining factor after Q. In addition, extreme temperatures may influence
photosynthesis through their damaging effects. Kinetics of enzymes involved with
photosynthetic reactions collectively comprise an additional set of factors that
influence leaf net photosynthesis.
176 B. Schaffer et al.

140

Vcmax (μmol CO 22/m /s) 120

100

80

60 y = 41.52x – 15.52
R2 = 0.87
40 y = –201.64x–1 + 173.41
R2 = 0.88
20

0
1.0 1.5 2.0 2.5 3.0 3.5 4.0 Na (g N/m2)
(a)

250

200
/s) μ 2 mol/m

150
Jmax (

100
y = 66.94x – 15.40
R2 = 0.83
50 y = –330.44x–1 + 291.55
R2 = 0.86
0
1.0 1.5 2.0 2.5 3.0 3.5 4.0
(b) Na (g N/m2)

Fig. 6.2. Relationship between (a) the maximum rate of carboxylation (Vcmax) and (b)
the light-saturated rate of electron transport (Jmax), and nitrogen concentration per unit
leaf area (Na). Measurements were performed on mango leaves of 3-year-old
‘Cogshall’ trees (●), standard leaves ({) and leaves close to developing fruits (…) of
_
11-year-old ‘Cogshall’ trees. Best fit lines for pooled data correspond to the linear ( )
–1 …
and the ax + b ( ) models (Source: redrawn from Urban et al., 2003).

4. Several parameters related to enzymes include the specificity factor of Rubisco


(W), the Michaelis constants of Rubisco carboxylation and oxygen-ation, Kc and
Ko, the activation and deactivation energies of the different pa-rameters 'Ha and
'Hd, the entropy terms 'S, c factors and leaf absorbance (T). With the exception of
Kc and Ko, the specific values of all these parame-ters have been estimated for
‘Cogshall’ mango (Urban et al., 2003).
5. The apparent efficiency of light energy conversion (D). This factor be-longs to
a category of its own since it should theoretically not differ from one
Ecophysiology 177

species to another and may be considered as a constant in the absence of


photoinhibition and photodamage.

Light

Light exposure
Plants allocate nitrogen resources within the canopy to enhance photosyn-thetic
capacity at locations exposed to high incident light levels, thus maxi-mizing whole
plant carbon gain (Field and Mooney, 1983; Hollinger, 1996; Carswell et al.,
2000). For leaves of a given age and for a given nitrogen sup-ply, leaf N per unit
leaf area appears to be strongly related with light expo-sure (DeJong and Doyle,
1985; Le Roux et al., 1999, 2001; Rosati et al., 1999, 2000). Photosynthetic light
acclimation of leaves may result from changes in either leaf nitrogen concentration
(Nm) or mass-to-area ratio (Ma) because Na = MaNm. Lynch and González (1993)
observed a negative correlation between Nm and light exposure in the tropical fruit
tree Borojoa patinoi, but such a behaviour is rare; positive correlations between Nm
and light exposure are more commonly observed. In addition, photosynthetic light
acclimation of leaves may result from changes in partitioning of total leaf N among
the different pools of the photosynthetic machinery (Evans, 1989). In mango, light
acclimation of photosynthesis results mainly from changes in Ma, and to a lesser
extent from changes in allocation of total leaf N at low irradiance; whereas changes
in Nm play only a minor role (Fig. 6.3). Light acclimation of mango leaves thus
follows a pattern similar to peach leaves (Le Roux et al., 1999; Walcroft et al.,
2002).

Light intensity
Photosynthesis of ‘Cogshall’ mango trees increases with increasing levels of light
intensity to reach a maximum at Q = 1200 Pmol photons/m2/s (L. Urban,
unpublished data). Whiley et al. (1999) measured Q at 1284 Pmol pho-tons/m2/s
for field-grown ‘Kensington Pride’ trees growing in subtropical Queensland,
Australia, which is well below full sunlight (full sunlight  2000 Pmol
photons/m2/s). Such a high threshold is a typical feature of sun plants. Individual
leaves are rarely able to utilize full sunlight; whole trees consist of many leaves
that shade each other, so that only a small fraction of a tree’s leaves are exposed to
full sun at any given time of the day, while the rest of the leaves receive
subsaturating photon fluxes in the form of small patches of light that penetrate
through gaps of the leaf canopy. Because the photosyn-thetic response of whole
trees is the sum of the photosynthetic activity of all the leaves, only rarely is
photosynthesis saturated with light at the whole-tree level.
While most leaves experience subsaturating light intensities, well-exposed
leaves of the upper-crown may receive excessive quantities of light. Those leaves
must dissipate the absorbed light energy in excess to prevent damage to the
photosynthetic apparatus. Moderate decreases in maximal quantum efficiency (i.e.
quantum efficiency of dark-adapted leaves Fv/FmPredawn) are
178 B. Schaffer et al.

3.0

(gN/gdrymatter)

2.0

1.0
N m

0.0
0.0 0.2 0.4 0.6 0.8 1.0
(a) Gap fraction

3.0
Tree # 1
y = 1.42x + 1.43
2.5
R2 = 0.90
2.0
Tree # 2
a2(g/m)

y = 1.33x + 1.44
1.5
R2 = 0.79
N

1.0

0.5

0.0
0.0 0.2 0.4 0.6 0.8 1.0
(b) Gap fraction

Fig. 6.3. Relationship between (a) leaf nitrogen concentration per unit mass (Nm) and
(b) leaf nitrogen concentration per unit leaf area (Na) and the gap fraction for
mango leaves measured in the crown of two 3-year-old ‘Cogshall’ trees. Gap
fractions were measured as an indicator of light exposure. Measurements were
performed on leaves < 2 months old (●), 8 months old (■), 12–14 months old (▲)
and 17–20 months old (♦) (Source: Urban et al., 2003).

typical features of moderate photoinhibition and should be interpreted in terms of


non-photochemical quenching, an adaptative mechanism involving the xanthophyll
cycle and allowing excess energy to be dissipated in the form
of heat (Adams et al., 2005). Such small decreases in Fv/FmPredawn are com-
monly observed in mango leaves even from well-irrigated trees (Urban and
Alphonsout, 2007).
When temperature (30C) and water vapour pressure deficit (VPD < 1 kPa) are
non-limiting, and in the absence of photoinhibition, maximal rates of net leaf
photosynthesis (Amax) may reach 12–15 Pmol CO2/m2/s at saturating
Ecophysiology 179

Q, on ‘Cogshall’ trees. Whiley et al. (1999) measured Amax of 15.2 Pmol CO2/
m2/s for ‘Kensington Pride’ trees growing in a subtropical climate in Queen-sland,
Australia. However, values > 16 Pmol CO2/m2/s were observed on field-grown
trees of ‘Tommy Atkins’, ‘Haden’ and ‘Irwin’ on sunny days during the wet season
in tropical regions of Australia (P. Lu, unpublished data). This is much higher than
citrus (< 10 Pmol CO2/m2/s), but substan-tially lower than plum (approx. 26 Pmol
CO2/m2/s). Whiley et al. (1999) esti-mated the light compensation point to be 29
Pmol photons/m2/s for leaves of non-stressed, field-grown mango trees, which is
much higher than that attributed to shade-tolerant species (< 10 Pmol
photons/m2/s) (Harvey, 1979). The data show that mango trees are basically sun-
adapted plants.

Leaf temperature

Effect of temperatures in a normal range on leaf photosynthetic capacity


Medlyn et al. (2002) calculated optimal temperatures for Vcmax and Jmax to be 35–
41C and 30–38C, respectively. Tree species native to cold climates had the
lowest temperature optima for both Vcmax and Jmax. For ‘Cogshall’ mango
trees, calculated temperature optima for Vcmax and Jmax are 44 and 45.5C,
respectively. They demonstrated that mango photosynthesis increases with
temperature well above 40C (Fig. 6.4). Although there is a clear lack of refer-
ences for other tropical trees, it is tempting to attribute the temperature response of
mango photosynthesis to its tropical origin.
Estimates of 'Ha, 'Hd and 'S are 7.0695, 17.0799 and 536 J/mol for Vcmax, and
3.8782, 10.2211 and 317 J/mol for Jmax, respectively. Both 'Hd and 'S are
within the range of published values for Vcmax and Jmax (Dreyer et al., 2001). In
addition, 'Ha for Vcmax is within the 60–80 kJ/mol range for many species,
including crop species as well as deciduous and evergreen trees (Medlyn et al.,
2002). For mango, 'Ha for Jmax is consistent with data published for evergreen
species (Medlyn et al., 2002), which is consistent with the fact that mango leaves
commonly are 2–4 years old before senescence and abscision. However, the
Jmax/Vcmax at 25C for mango is about 1.86, approximately 11% higher than the
mean value calculated over the whole range of species stud-
ied by Medlyn et al. (2002). Lower activation energies for Jmax than Vcmax result
in a temperature-induced decrease in Jmax/Vcmax, confirming previous
observations by Walcroft et al. (1997) and Dreyer et al. (2001). The estimate of 'Ha
for Rd is 4.5710 J/mol. 'Ha is higher than the range of published values for Rd
(Dreyer et al., 2001).

Chilling temperatures
Fv/FmPredawn decreases in mango leaves with decreasing temperature, while
chilling reduces quantum efficiency (Whiley et al., 1999; Sukhvibul et al.,
2000; Weng et al., 2006a, b). The decrease in Fv/FmPredawn may be interpreted to
reflect sustained engagement of zeaxanthin in photoprotective energy dis-
sipation. Decreases in quantum efficiency correspond to a decrease in the rate of
electron flow. It may be argued that sustained zeaxanthin-dependent
180 B. Schaffer et al.

4.0

3.5

3.0
Vcmax / Vcmax at 25°C

2.5

2.0

1.5

1.0

0.5

0.0
15 20 25 30 35 40 45 50
(a) Tl (°C)

2.0
1.8
1.6
1.4
Jmax /Jmax at 25°C

1.2
1.0
0.8
0.6
0.4
0.2
0.0
15 20 25 30 35 40 45 50
(b) Tl (°C)

Fig. 6.4. Temperature response functions adjusted to the (a) maximal rate of car-
boxylation (Vcmax) and (b) the light-saturated rate of photosynthetic electron flux
(Jmax), normalized to the mean value at 25°C in leaves from ‘Cogshall’ mango
seedlings. The data scatter represents the real scatter at each temperature. Reference
values at 25°C were computed for each of eight leaves, taken from young trees from
two origins (● and ), and a unique temperature response was adjusted over the range
of normalized data. Tl, the leaf temperature; the dotted lines correspond to Equation 9;
the solid lines correspond to Equation 10 (no deactivation energy component).

energy dissipation reduces the risk of formation of singlet oxygen 1O2 in the
antennae, while decreases in J lower the risk of electrons reducing O2 to anion
superoxide O2− in the photosynthetic electron transport chain (Adams et al.,
2005). In other words, decreases in Fv/FmPredawn and quantum efficiency cor-
respond to adaptative mechanisms against the effect of cold, when photosyn-
thesis is low and there is an imbalance between the quantity of light energy
absorbed and the quantity of energy used in the photochemical reactions of
Ecophysiology 181

photosynthesis (Adams et al., 2005). Interestingly, Weng et al. (2006b) found that
mango leaves transferred from warm and dark to chilling conditions showed only
slight down-regulation of PSII efficiency when compared to leaves moved from
dim light to chilling conditions. Of course, long-term exposure to cold and very
low temperatures ( 10C) may eventually result in true photodamage, not just
photoinhibition. Very low values of Fv/
FmPredawn, decreases in chlorophyll content and slow recovery kinetics are all
indicators of photodamage. Sukhvibul et al. (2000) observed that susceptibil-
ity to cold-induced photodamage was more pronounced in polyembryonic cultivars
than in monoembryonic cultivars, possibly reflecting their different eco-
evolutionary development.

Elevated atmospheric CO2 concentration

The CO2 concentration in the earth’s atmosphere has been increasing rapidly since
the early 20th century and is continuing to rise, primarily due to burn-ing of fossil
fuels (Houghton, 2005). Earth’s atmospheric CO2 concentration is currently about
370 Pmol CO2/mol (Houghton, 2005) and is projected to reach 600 Pmol CO2/mol
by 2050. Elevated ambient CO2 levels will undoubt-edly affect cropping systems
since atmospheric CO2 concentrations can sig-nificantly affect plant growth and
productivity (Idso and Kimball, 1991; Houghton, 2005). There is little published
information concerning the effects of elevated ambient CO 2 levels on physiology,
growth and production of tropical fruit trees, including mango.
Schaffer et al. (1997) exposed leaves of field- and container-grown ‘Kens-
ington’ (syn. ‘Kensington Pride’) trees to short durations (several minutes) of
varying ambient CO2 concentrations. They found that under saturating light levels
for photosynthesis, net photosynthesis increased as ambient CO 2 con-centration
increased up to 1200 Pmol CO2/mol. At ambient CO2 concentra-tions > 1200
Pmol CO2/mol, net photosynthesis stabilized, probably due to leaves reaching their
maximum biochemical capacity to fix carbon. Studies with ‘Cogshall’ mango trees
indicated that when Ca increases, stomata close swiftly and Ci may become very
unpredictable (L. Urban, unpublished data). Therefore, using Ci may be preferable
to Ca for quantifying short-term effects of elevated CO2 concentations on Anet of
mango. Saturating CO2 levels may often be reached at Ci = 800 Pmol CO2/mol air.
Long-term (6–12 months) exposure of ‘Kensington’ mango trees to an
atmospheric CO2 concentration of 700 Pmol/mol resulted in higher net CO 2
assimilation rates than in leaves of plants grown at atmospheric CO 2 concen-
trations of 350 Pmol/mol when net CO2 assimilation was measured at the same
CO2 concentration as the growth environment. However, carboxylation efficiency
(the amount of CO2 fixed per mole of ambient CO2) was lower for plants in the
CO2-enriched environment compared to plants in the ambient (350 Pmol
CO2/mol) environment (Schaffer et al., 1997). Although further studies are needed
to determine the effects of long-term exposure to elevated CO2 concentrations on
mango growth and productivity, it appears that
182 B. Schaffer et al.

mango will benefit from increases in atmospheric CO2 concentrations. How-ever,


the effects of increased atmospheric CO2 concentrations associated with global
warming on mango production may be offset by higher respiratory losses and
increased assimilate partitioning to shoot growth in highly vegeta-tive cultivars
(Schaffer et al., 1999). Therefore, responses of mango cultivars to elevated
atmospheric CO2 concentrations need to be evaluated over a range of temperatures
to ascertain their likely performance under changing atmospheric conditions.

Humidity

Although mango production occurs in the tropics and subtropics in areas of high
and low relative humidity (RH) (Campbell, 1984), there are very few published
reports on the effects of RH or VPD on physiology and tree growth.

In a study with container-grown ‘Kensington’ plants, Pongsomboon et al.


(1992) reported that stomatal conductance was inversely correlated with VPD.
Differences between cultivar responses to VPD have been observed in field-grown
‘Irwin’ (monoembryonic) and ‘Kensington’ (polyembryonic) mango trees during
the wet and dry season in tropical Australia. During the wet season and for well-
irrigated trees during the dry season, both ‘Irwin’ and ‘Kensington’ showed
decreasing stomatal conductance with increasing leaf-to-air vapour pressure deficit
(LAVPD) but ‘Kensington’ showed a more rapid decrease than ‘Irwin’ (Fig. 6.5)
(P. Lu, unpublished data). It was also observed that daytime leaf xylem water
potential was lower in ‘Irwin’ than in ‘Kensington’ while predawn water potentials
were similar for both culti-vars (P. Lu, unpublished data). These results indicate
that under similar soil water conditions, ‘Kensington’ tends to close stomata much
more rapidly than ‘Irwin’ to conserve water under dry atmospheric conditions. This
water conservation strategy is probably a reflection of ‘Kensington’s’ adaptive
responses to the hot and dry seasonal tropical environment under which it evolved
(Wolstenholme and Whiley, 1995). Other studies in tropical Austra-lia revealed
that polyembryonic ‘Nam Doc Mai’ behaved like ‘Irwin’ (P. Lu, unpublished data).
However, ‘Nam Doc Mai’ in Thailand has comparatively low vigour compared to
‘Kensington’ when grown in the tropics.

Further research is required to determine if differences in photosynthetic or


stomatal responses of mango to VPD are indeed based on embryonal char-
acteristics. Clarification of the reasons for variation would undoubtedly facil-itate
breeding and selection of cultivars for dry and humid areas.

Flooding

The primary effect of flooding on plants is due to a reduction in soil oxygen


concentration. Oxygen levels in the soil can decrease from 20% to < 5% within 1–2
days of flooding (Crane and Davies, 1988) and soils eventually become
Ecophysiology 183

0.6

0.5

Stomatal conductance (mol/m2/s)


0.4
‘Irwin’: R2 = 0.594

0.3

0.2

0.1 ‘Kensington’: R2 = 0.635

0.0
1 2 3 4 5 6 7 8
LAVPD (kPa)

Fig. 6.5. Correlation between leaf stomatal conductance and leaf-to-air vapour
pressure deficit (LAVPD) during the dry and wet season for ‘Irwin’ (closed circles) and
‘Kensington’ mango trees (open circles). Trees were well irrigated during the dry
season and all measurements were taken when the Q > 300 Pmol photons/m2/s
(n = 12) (Source: P. Lu, unpublished data).

anoxic (no oxygen). Mango is considered to be a moderately flood-tolerant species


(Schaffer et al., 1994, 2006) and waterlogging or flooding of trees peri-odically
occurs in many of the regions where the crop is grown (Plate 41). Mango trees
have evolved a mechanism to cope with temporary flooding (see ‘Flooding’ section
under Tree Growth and Development, this chapter).
Typically, the first easily measurable responses of fruit trees to flooding are
reductions in Amax, gs and transpiration, which occur within 2–3 days fol-lowing
flooding (Larson et al., 1991c; Schaffer et al., 1992, 2006). Short-term anoxia
results in a decrease in net photosynthesis which cannot be related to a gs-
associated decrease in Ci (Zude-Sasse et al., 2001). Removing trees from flooded
conditions after 28 days reversed the flooding-induced decrease in leaf gas
exchange, resulting in a gradual increase in photosynthesis and tran-spiration to
preflooded rates.
Although flooding adversely affects mango trees, short-term flooding of trees
in limestone soils can result in increased micronutrient availability with improved
plant nutritional status. In calcareous soils of south Florida, in which iron (Fe) was
withheld from the fertilizer programme, short-term flooding (10–20 days) of
polyembryonic ‘Peach’ mango trees resulted in an increase in net photosynthetic
rates to above preflooding levels following the release of trees from flooding
(Larson et al., 1992). This increase in photosyn-thesis has been correlated with
improved Fe and manganese (Mn) uptake as
184 B. Schaffer et al.

a result of these elements becoming more soluble when calcareous soils are flooded
(Larson et al., 1991b, 1992).

Internal factors

Leaf age
Leaf characteristics (i.e. photosynthetic capacity and the amount of N per unit area)
are generally strongly influenced by leaf age, with maximum val-ues being
observed when leaves have just completed full expansion (Con-stable and Rawson,
1980; Marshall and Biscoe, 1980; Dwyer and Stewart, 1986; Field, 1987; Wilson et
al., 2000; Frak et al., 2001). Chlorophyll content is three to four times lower in
young than in mature mango leaves (Zude and Ludders, 1997). Similarly, the
concentration of Rubisco is lower in young than in mature, green leaves (Nii et al.,
1995). In contrast to many other plant species, once mango leaves are mature the
relationship between Na and irra-diance does not seem to be affected by leaf age
(Urban et al., 2003). The Na values may remain high in old leaves experiencing
high irradiance. This indicates that changes in Na in mango leaves are influenced
by irradiance and not age, at least during the first year.

Carbohydrate accumulation and source-sink balance


Source-sink imbalances can exert feedback down-regulation or repression of leaf
photosynthesis through carbohydrate accumulation in leaves (Azcon-Bieto, 1983;
Foyer, 1988; Koch, 1996; Schaffer et al., 1997; Whiley et al., 1999; Paul and
Foyer, 2001; Paul and Pellny, 2003). Transient accumulations of car-bohydrates in
leaves, as they have been observed during the diurnal period, may impair the rate
of electron transport (Pammenter et al., 1993). Changes in photosynthetic capacity,
not just assimilation rates, are more likely to be observed in association with lasting
source-sink imbalances. One hypotheti-cal mechanism is that high levels of
carbohydrates repress the expression of genes coding for several photosynthetic
enzymes (Krapp and Stitt, 1995; Koch, 1996; Drake et al., 1997). Alternatively,
carbohydrates may interact with hormonal signals to control gene expression
(Thomas and Rodriguez, 1994). There is also some evidence that photosynthetic
capacity is related to leaf carbohydrate status through the effect of the latter on
phosphate availability (Riesmeier et al., 1993; Sun et al., 1999). In the long term,
carbohydrate accu-mulation may eventually lead to cell death. High sugar
concentration has been associated with senescence in leaves of several species
(Noodén et al., 1997; Wingler et al., 1998; Quirino et al., 2001). Reduced energy
utilization by CO2 assimilation, like the one resulting from carbohydrate
accumulation, in combination with high energy capture is potentially dangerous
and can result in over-reduction of the electron transport chain, photoinhibition and
oxidative stress caused by photoreduction of oxygen to superoxide O 2− in the
Mehler-ascorbate peroxidase reaction (Badger, 1985). Moreover, reactive sin-glet
oxygen 1O2 can be formed through reaction of oxygen with triplet chlo-rophyll
released by the breakdown of the chlorophyll-protein complexes in
Ecophysiology 185

thylakoids (Merzlyak and Hendry, 1994). Formation of reactive oxygen spe-cies


can lead to membrane damage and eventually cell death.
Whiley et al. (1999) observed that Amax, which is closely related to photo-
synthetic capacity, the quantum yield and Fv/FmPredawn are substantially lower in
mango trees grown in containers (root-restricted) when compared
to field-grown trees. These observations were confirmed by Urban and Alphonsout
(2007) who studied the effect of the removal of a 1 cm-wide band of bark on leaf
photosynthesis and leaf N content of 3-year-old and 11-year-old ‘Cogshall’ mango
trees. Girdling is a common horticultural practice used to manipulate tree growth
and development in many fruit species. Its most immediate effect is to stop the
basipetal movement of assimilates through the phloem, which results in an
accumulation of carbohydrates above the girdle (Roper and Williams, 1989;
Schaper and Chacko, 1993; Di Vaio et al., 2001). Girdling can promote floral
induction in mango (Chacko, 1991), but it has also been shown to reduce net
photosynthesis. The major effect of girdling is a dramatic increase in leaf
carbohydrate concentration and a concomitant decrease in photosynthetic electron
transport and net photosynthesis (Fig. 6.6) (Gonzalez and Blaikie, 2003; Urban and
Alphonsout, 2007). Urban and Alphonsout (2007) observed that Anet was reduced
by 77% within 28 days from girdling and remained at about 2 Pmol CO2/m2/s
until the beginning of flowering. The decrease in photosynthetic electron transport
rate (J) and
sustained photoprotection (reflected by the decrease in Fv/FmPredawn) pro-tected
leaves of girdled branches effectively from photodamage, as shown by
the vigorous recovery of Anet and J observed immediately after the appear-ance of
inflorescences. This increase in Anet and J was associated with no

250
A Q = 400 μmol photons/m2/s
y = 84.5e–0.0313x
200 R2 = 0.69
J (μmol electrons/m2/s)

Q = 1200 μmol photons/m2/s


150 y = 120.3e–0.0405x
R2 = 0.74
Q = 2000 μmol photons/m2/s
100 y = 155.4e–0.0401x
R2 = 0.74
50

0
0 10 20 30 40
2
Starch (g/m )

Fig. 6.6. The relationship between the total photosynthetic electron flux (J) measured
at photosynthetic photon flux (Q) = 400 ('), 1200 (●) and 2000 () Pmol photons/
m2/s, and the amount of starch per unit leaf area. Best fit lines at each Q were assessed
from measurements performed on both girdled and non-girdled mango leaves before
flowering. Data were used to establish the following relationship: J = (0.0434Q +
–0.0412[starch]a
72.8)*e (Source: Urban and Alphonsout, 2007).
186 B. Schaffer et al.

decrease in leaf carbohydrate content during the first month following the onset of
flowering, suggesting that the effect of carbohydrate accumulation on
photosynthesis is mediated by sink activity. Apart from its negative effect on the
carbon budget of mango trees, girdling appeared to be rather harm-less. However,
leaf N concentration decreased, which indicates that there may indeed exist long-
term negative effects of girdling on photosynthetic capacity. The width of bark
(phloem) removed may be critical with respect to the intensity of the effect of
girdling on the tree. Whiley et al. (2006) girdled the trunks of ‘B74’ (‘Calypso’™)
mango trees in the Northern Territory of Australia in autumn (as soon as they had
come out of the wet season). The girdles were no more than the thickness of a
pruning saw (about 1 mm) and healed within 6 weeks. In the first and third years
after girdling, the trees had significantly higher yields than non-girdled trees on
which, coincidentally, fruit matured early, thus giving market advantage. There
was no significant difference in yield in the second year of treatment between
girdled and non-girdled trees. The first and third years had strong natural induction
while the second year gave poor flowering across all varieties in the district. Thus,
dur-ing years of strong induction, this type of girdling most likely provided extra
carbohydrate reserves to drive flowering and support fruit set and retention while in
the off-flowering year there was sufficient carbohydrate reserves to support
reproductive activity. In contrast to observations with ‘Cogshall’ mango trees
(Urban and Alphonsout, 2007), there was no evidence of long-term effects of
narrow girdles on leaf N of ‘B74’ mango trees (Whiley et al., 2006). However,
when wider girdles are made, tree recovery may take much longer leading to
sustained physiological disruption.

Proximity of inflorescences
While the effects of water stress and high light, temperature and atmospheric CO 2
concentration on photosynthesis are increasingly well described, very little is
known about the effect of phenology, and especially of flowering on
photosynthesis of mango. There is some evidence that flowering may have an
effect on photosynthesis. Flowering-associated decreases in Anet and gs were
observed in sweet cherry (Roper et al., 1988) and mango (Shivashankara and
Mahai, 2000; Urban et al., 2004a). Lack of precise knowledge about the effect of
flowering on photosynthesis may impair our ability to adequately simulate
photosynthesis, especially for tropical fruit trees for which flower-ing often extends
over a long period of time. Mango flowering can last for > 2 months. Therefore, its
effect on photosynthesis should not be overlooked. Urban et al. (2004a) showed
that the decrease in Anet in mango leaves close to inflorescences is not attributable
to a gs-associated decrease in Ci or to an increase in Rd. Rd was lower in leaves
close to inflorescences than in standard leaves. If any, the effect of Rd on Anet was
a positive one. This study suggested
strongly that the decrease in Anet was due to a decrease in the electron flow in
photosystem II, but failed to provide direct evidence for it as well as the ele-
ments for interpretation. Using a modelling approach, Urban et al. (2008)
confirmed that there is a decrease in the total light-driven photosynthetic electron
flux in leaves close to inflorescences and showed that the decrease
Ecophysiology 187

in Anet is also attributable to an increase in photorespiration. The latter appears to


be the consequence of a gm-associated decrease in Cc, while the former results
from an increase in electron flow towards alternative sinks, a decrease in the
amount of leaf N per unit leaf area, and, hypothetically, either a decrease in leaf N
allocation to the bioenergetic pool of the photosynthetic machinery, inorganic
phosphorus depletion in leaves, or feedback inhibition of photosynthesis. The latter
hypothesis is least probable in the absence of carbohydrate accumulation in leaves
close to inflorescences. Both of these hypotheses need to be tested to further our
understanding of the inhibiting effect of inflorescences on photosynthesis of nearby
leaves. Interestingly, net photosynthesis measured on leaves close to panicles
bearing set fruits are intermediary between those measured on standard leaves and
those mea-sured on leaves close to inflorescences, suggesting that changes in
photosyn-thesis associated with flowering are reversible. Urban et al. (2008) also
showed that processes other than temperature or light acclimation, and acclimation
to reduced sink activity, may cause leaf N concentration and photosynthetic
capacity to vary in mango. Whiley (unpublished data) observed that the mango
leaves immediately adjacent to inflorescences lose colour intensity as the
inflorescence grows out. Although leaf N over this period was not mea-sured in
mango, it has been measured in avocado (Whiley, 1994) which has a similar
intense burst of flowering in which a large biomass is produced in a short time.
Leaf N declines rapidly in avocado leaves as the inflorescences break from buds
and grow and then N stabilizes (at a lower concentration) by mid-bloom. This can
be reversed by N applications during flowering and the additional application of a
growth retardant (paclobutrazol (PBZ)) giving leaf N and A a significant boost.
Similarly to avocado, it is likely that the reduction in A close to mango
infloresences is related to reduced leaf N.

Photosynthetic contributions by fruit

Fruit of many species have chlorophyll and photosynthetic activity, particu-larly


during the early stages of growth (Jones, 1981; Whiley et al., 1991). How-ever, for
most crops, respiratory losses from fruit exceed photosynthetic gains throughout
ontogeny (Kriedemann, 1968; Whiley et al., 1991). An exception to this is
blueberry (Vaccinium spp.) fruit in which there is a net photosyn-thetic gain from
petal fall through to colour break, with an estimated 15% of the total fruit carbon
requirement contributed from fruit photosynthesis (Birkhold et al., 1992). Studies
with monoembryonic ‘Dashehari’ mangoes showed that when fruit were
approximately 10 mm in diameter, the photo-synthetic rate of fruit was 2.7% that
of leaves and declined to 1.2% of leaf photosynthesis at fruit maturity (Chauhan
and Pandey, 1984). However, even this comparatively small carbon contribution
may be important during the critical fruit set period when trees rely on stored
carbohydrates and a rela-tively inefficient canopy to supply current photosynthates.
Further studies with mangoes to establish optimum light regimes for fruit
photosynthesis at different stages of ontogeny are warranted.
188 B. Schaffer et al.

6.3 Plant Water Relations


In this section, theoretical concepts of plant water relations are briefly out-lined to
help interpret the effects of environmental factors on mango water relations (see
also Nobel (1983) and Baker (1984)).
An important concept in plant water relations is water potential (Ψ), which is a
measure of the free energy of water. For pure water, Ψ = 0. As sol-utes are added
to water, its free energy decreases and < becomes more nega-tive. Water moves
along a gradient from higher to lower (more negative) <.
< can be expressed as:
< = <S + <p + <m (6.11)
where <S is osmotic (or solute) potential which refers to the effect of solutes on the
change in free energy of water; <p is the hydrostatic or pressure poten-tial also
referred to as the turgor pressure; and <m is the matric potential, which is generally
negligible in plant cells.
In plant cells, <p is generally positive or equal to 0. However, in xylem tissue
of transpiring plants, <p is negative (under tension). The driving force for
transpiration is the vapour pressure difference between the leaf (consid-ered to be
water saturated) and the surrounding air. Thus, water moves from a greater to a
lower (or more negative) <p and hence along a decreasing < gradient. The cohesive
forces of the H2O molecules allows the xylem water to remain in a continuous
column even though there is a negative <p.
Plant water stress can be determined from Eqn 6.11 and from changes in <.
The components of <, such as <S and <p, can often be used to define the sources
of water stress. The drought tolerance of mango highlights some unique aspects of
physiology of this tree with respect to its water manage-ment. Typical mango
environments in the tropics impose extreme water stress and high evaporative
demand for prolonged periods. Adaptive strate-gies of mango trees include a deep
root system (Sukonthasing et al., 1991), desiccation-tolerant surface feeder roots
and drought avoidance mechanisms thought to be mediated by a comprehensive
system of resin canals distrib-uted throughout the tree (Venning, 1948; Joel, 1980;
Joel and Fahn, 1980a, b; Pongsomboon, 1991) and rapid stomatal closure. Plants
with laticfers or resin ducts/canals have been reported to be drought tolerant due to
extended maintenance of turgor following the withdrawal of water (Downton,
1981; Kramer, 1983; Kallarackal et al., 1990). While the mechanism of turgor
main-tenance remains unresolved, it is believed that the latex or resin is probably
involved in the modulation of plant water status (Kallarackal et al., 1990). The
differentiation, structure and distribution of resin canals in mango has been
described by Venning (1948), Joel (1980) and Joel and Fahn (1980a, b, c). Resin
canals are present in trunks, shoots, leaves and fruit (exocarp) of mango in close
association with the vascular tissues. The resin contains mainly ter-penes, but
phenols and protein-carbohydrate mucilage are also present (Joel and Fahn, 1980c).
In well-watered trees, the resin is under positive pressure and freely exudes from
damaged or cut surfaces (Pongsomboon, 1991). In studies of the development of
water deficit in container-grown ‘Kensington’
Ecophysiology 189

mango trees, loss of turgor occurred in expanding leaves when leaf water potential
(<l) reached –1.2 Mpa. In mature leaves turgor was not lost until <l reached –1.75
MPa (Pongsomboon, 1991). Necrotic leaf areas appeared when <l reached
approximately –3.2 MPa with permanent wilting developing at −3.45 MPa. This is
high (less negative) compared with a <l of –6.6, –5.0 MPa for orange (Citrus
sinensis) and macadamia (Macadamia integrifolia), respec-tively (Fereres et al.,
1979; Stephenson et al., 1989). Thus, mango leaves toler-ate less internal water
stress than woody perennial fruit trees from more mesic environments. With
mango, however, the permanent wilting point was reached 36 days after
withholding water compared to 10 days for simi-larly sized macadamia trees
(Stephenson et al., 1989). The higher critical threshold of <l and the longer period
of survival for ‘Kensington’ mango indicates that drought tolerance is based on
more effective water regulation to prevent desiccation and on maintenance of leaf
turgor rather than resis-tance by tissues to damage.
Reich and Borchert (1988) observed that stomatal regulation in mango
significantly reduced the rate of development of internal water deficit when
compared with four other tropical tree species. In water-withholding studies, the
radial expansion of mango trunks continued when most of the other species were
shrinking, indicating that mango trees could better tolerate drought conditions and
maintain photoassimilation rates. This is consistent with the decrease in gs/Anet
observed by Urban et al. (2006) as a consequence of drought (Fig. 6.1). A decrease
in gs/Anet indicates that there is an increase in photosynthetic water use efficiency.

In containerized ‘Kensington’ mango trees, there was a linear correlation


between stomatal conductance and <l during the development of water stress
(Pongsomboon, 1991). In contrast, with avocado and macadamia, stomatal
response was much more rapid with a curvilinear relationship between <l and
stomatal conductance, and stomatal closure reached at –1.2 and 3.0 MPa,
respectively. The slower response of stomatal conductance to <l in mango trees
appears to be related to a more effective mechanism for the mediation of water
deficit development compared with avocado and macadamia.
Pongsomboon (1991) monitored leaf water potential, osmotic potential of resin
(<r) and osmotic potential of the whole leaf tissue (<S) in container-grown
‘Kensington’ mango trees when water was withheld for a 45-day period. When
tissues were fully hydrated, <l and <r were higher than <S. For 40 days of the
drying cycle, <l and <r declined at a similar rate; however, <S declined to about –
1.2 MPa within 4 days where it remained stable until 18 days into the drying cycle.
There was a subsequent decline in <S for 28 days after withholding water when it
stabilized at –2.0 MPa, remaining constant for another 12 days. Pongsomboon
(1991) suggested that osmotic adjustment occurred, probably mediated through the
resin as water deficit developed. It appears, therefore, that the energy investment
by the tree in a resin canal sys-tem is justified by the vital drought-avoidance
benefits conferred by main-taining turgor and preventing wilting under prolonged
periods of water stress. Further investigations are required to substantiate these
conclusions.
190 B. Schaffer et al.

In a recent experiment of 2-year-old potted ‘Cogshall’ mango trees grafted on


‘Maison Rouge’ rootstock, midday minimum leaf water potential remained
relatively constant, about –1.0 MPa, for the range of predawn leaf water potential
(a surrogate of soil water potential) from 0 to –0.5 MPa, but when the predawn leaf
water potential dropped below –0.5 MPa, leaf water potential fell rapidly to about –
1.8 MPa (Gaelle Damour, Centre de coopéra-tion Internationale en Recherche
Agronomique pour le Développement (CIRAD), personal communication). This
‘isohydric’ behaviour seemed to be associated with rapid stomatal closure in
response to decrease in water avail-ability (Urban and Jannoyer, 2004).

Development of novel techniques to study xylem integrity have pro-vided


some unique insights into stomatal function (Tyree and Sperry, 1988; Cochard et
al., 2002). Xylem sap is transported under negative pressures in plants and
therefore is susceptible to cavitation events that render xylem conduits non-
conductive. Cavitation occurs when the negative sap pressure exceeds a threshold
value defined by anatomical characteristics (Tyree and Sperry, 1988). Many
species function very close to the point of embolism. Therefore, stomata control
both plant water losses and sap pressure and thus may actively control the risk of
xylem embolism (Jones and Sutherland, 1991). Xylem vulnerability to cavitation
was studied in twigs of 13-year-old ‘Cogshall’ mango trees. Xylem vessels started
to cavitate at a xylem water potential of about of –1.5 MPa and underwent a rapid
and substantial loss (90%) of hydraulic conductivity when xylem water potential
dropped from –2.0 to –3.0 MPa (Fig. 6.7) (H. Cochard, unpublished data). This
result is con-sistent with the observation that leaves lose turgor when <l reaches –
1.75 MPa and permanent wilting appears when <l is above –3.0 MPa (Pongsom-
boon, 1991) when Fig. 6.7 predicts 90% of xylem hydraulic conductivity is lost. It
also confirms that mango, like many other trees, operates close to the point of
embolism via effective stomatal control.

Direct measurement of xylem water potential using a pressure chamber is


problematic due to the presence of latex (Castro Neto et al., 2004). It is more
problematic in some cultivars, for example ‘Kensington’, than in others, such as
‘Irwin’. Alternative indicators of plant water status such as measurements of
whole-tree water use (sap flow) and stem/branch shrinkage were found to be
sensitive and integrated indicators of whole-tree water status in mango and were
successfully used to schedule irrigation in a mango orchard in the seasonal dry-wet
tropical region of northern Australia (Lu, 2002, 2006).

6.4 Tree Growth and Development


Light

In an orchard, light distribution within and between tree canopies can have a
profound effect on growth and development of the fruit. We have previ-ously
discussed the effect of light on photosynthesis and defined the opti-mum light
levels required for mango leaves. When light levels fall below the
Ecophysiology 191

100
2006 light
2006 shade
80 2005

60
PLC

40

20

0
–4 –3 –2 –1 0
Xylem pressure (MPa)

Fig. 6.7. Vulnerability of xylem of 13-year-old ‘Cogshall’ mango trees to cavitation in


twig segments from sun-exposed (light) or shaded sides of the tree. Cavitation is
expressed as percentage loss of conductivity (PLC) with decreasing xylem water
potential. Symbols represent means and error bars represent one standard error.
Vulnerability curves were obtained with the centrifuge technique (Source: Cochard et
al, 2005; and from H. Cochard, unpublished data, with permission).

threshold required for light saturation of photosynthesis, the subsequent reduction


in available photoassimilates will affect growth of the tree. In many tree fruit crops,
flower-bud induction, fruit size and fruit colour are reduced when low light levels
occur due to crowding within and between tree cano-pies (Jackson, 1980; Flore,
1994; Whiley and Schaffer, 1994). There is no pub-lished information on the effect
of light levels on mango fruit size, although fruit are photosynthetically active and
a reduction in size under low Q could be expected.

Fruit skin colour is an important feature of mango with fruit of many cultivars
developing attractive pink to red coloration. Fruit colour is geneti-cally determined
and the reddish blush is generally more developed in monoembryonic cultivars,
while fruit from most polyembryonic cultivars remain green/yellow at maturity.
Skin coloration of mature fruit is partly due to anthocyanins which develop when
tissues are exposed to light. While this subject is well researched in other fruit
crops (Proctor and Creasey, 1971), light levels required for skin coloration of
mango fruit have not been quanti-fied. Studies in Australia with the polyembryonic
cultivar ‘Kensington’, which develops a blush only on the exposed side of the fruit,
indicated that the position of fruit on trees had a significant effect on the
development of colour due to differences in the penetration of light into the canopy
during fruit ontogeny (Schaffer et al., 1994). The intensity of redness was greatest
on fruit from the eastern side of the tree followed by fruit from the south-western
192 B. Schaffer et al.

and northern sides of the tree. This information establishes an important con-cept
with respect to the light regime but does not quantify the absolute light levels
required for anthocyanin development. Further research is necessary to establish
physiological parameters from which pruning and orchard man-agement strategies
can be developed.

Temperature

Mango is a predominantly tropical species although the tree will usually grow and
produce more successfully in frost-free subtropical latitudes with a marked dry
season and high heat accumulation. Under optimum tempera-tures with non-
limiting nutrients and water, the tree remains vegetative with growth flushes
occurring at regular intervals. The large size and poor crop-ping of trees in the
humid lowland tropics are well known, and there is a direct relationship between
temperature and the frequency of vegetative flushes. Trees grown at 20C
days/15C nights (20/15C) required 20 weeks (mean of ten cultivars) to complete
a growth/dormancy cycle while at 30/25C the same cycle was completed in 6
weeks (Whiley et al., 1989). There are marked differences between cultivars with
respect to their tendency towards vegetative growth. For instance, in controlled
temperature studies over a 20-week period, at 30/25C, ‘Irwin’ produced 2.0
growth flushes with approximately 45 days of dormancy between active growth
periods while ‘Kensington’ produced 4.7 growth flushes with only 5 days of
quiescence between flushes (Whiley et al., 1989). Dry matter accumulation over
the 20 weeks was similar for the two cultivars; however, starch accumulation in the
woody trunk tissues of ‘Irwin’ and ‘Kensington’ was 13 and 3.6% of dry mat-ter,
respectively. The response differences between these two cultivars may be the
contributing factor to their performance at tropical latitudes where temperature is
non-limiting for growth. Under these conditions ‘Irwin’ has more reliable cropping
than ‘Kensington’, suggesting that the genetically determined low-vigour trait is
more sensitive to environmentally precipi-tated stresses that induce flowering.

The number and size of leaves which develop on each growth flush are also
influenced by temperature. Whiley et al. (1989) reported that on trees growing at
20/15C an average of 7.1 leaves per flush were produced while at 30/25C there
were 13.6 leaves on each growth flush (data are mean values from ten cultivars). At
30/25C the mean leaf size was 300% greater than those on trees growing at
20/15C. Soil temperatures have also been reported to have a strong effect on the
growth of mangoes. In studies with ‘Irwin’ grafted on ‘Turpentine’ rootstocks,
episodic shoot growth occurred when soil temperatures were held at 27C or 32C
for 120 days but an extended dormant period developed when soil temperatures
were held at 21C (Yusof et al., 1969). These results indicate that environmental
control over shoot growth of mangoes may in part be related to soil temperatures.

From controlled temperature studies it has been calculated that the median
daily temperature (mean of the maximum and minimum daily temperatures)
Ecophysiology 193

at which shoot growth ceases is approximately 15C (mean value for ten cul-tivars)
(Whiley et al., 1989). Subsequent studies (Issarakraisila et al., 1991) have
confirmed that 15C is the critical minimum growth temperature for shoots of
‘Kensington’.
Stress-inducing temperatures which prevent shoot growth have been shown to
promote floral induction in mangoes, but this is outside the scope of this
discussion. For further information of the effects of temperature on pollination,
floral initiation and fruit development, see Davenport, Chapter 5, this volume and
Schaffer et al. (1994). We again emphasize that although mango is a ‘heat-loving’
crop well adapted to the hot, semi-arid subtropics and monsoonal tropics, in these
environments it experiences extremes of heat, drought and evaporative demand that
may cumulatively reduce potential production capacity.

Drought

Although mango is considered to be drought tolerant and may survive with-out rain
or irrigation for > 8 months (Gandhi, 1955), water deficits during the reproductive
cycle can have severe effects on the retention and early growth of mango fruit. In
studies with bearing, container-grown ‘Irwin’ trees, pre-dawn <l levels were
maintained at either less than –0.3 MPa (non-stressed) or –1.2 MPa (water stressed)
for the first 2 months after fruit set. For the first 5 days following fruit set, all trees
lost a similar percentage of fruit, but there-after fruit abscission was greater on
water-stressed trees. After 1 month, drought-stressed trees had retained
approximately 4% of their initial fruit set compared with approximately 8% on
non-stressed trees. During the first 30 days following fruit set, the rate of fruit
growth for non-stressed trees was twice that of drought-stressed trees, and final
fruit size (measured 60 days after fruit set) of non-stressed trees was 20% greater
than on water-stressed trees. In a separate study, another group of ‘Irwin’ trees was
maintained stress-free (pre-dawn water potential above –0.3 MPa) during the first
month following fruit set with water-stress (–1.2 MPa) imposed on some trees dur-
ing the second month of fruit development (Pongsomboon, 1991). There was no
effect of drought on fruit retention but final fruit size was 34% smaller for stressed
compared with non-stressed trees.

In field studies, Singh and Arora (1965) compared fruit drop of monoem-
bryonic ‘Dashehari’ trees irrigated at 1-week intervals with trees irrigated at 3-
week intervals. During the first 6 weeks of fruit growth, weekly irrigation reduced
fruit drop compared with the irrigation at 3-week intervals. During the latter stages
of fruit development, these gains were lost as more fruit dropped from the weekly
irrigated trees. In another study, field-grown monoembryonic ‘Tommy Atkins’
trees were managed under different irriga-tion regimes from early fruit set until the
start of the rainy season (approxi-mately 43 days) (Larson et al., 1989). Trees were
irrigated on a 7- or 14-day schedule or received no irrigation. Pre-dawn <l was –0.3
MPa for trees irri-gated on the 7-day schedule and decreased to –0.5 MPa for the
non-irrigated
194 B. Schaffer et al.

trees. Irrigation at 7-day intervals resulted in the greatest yield with the larg-est
fruit, especially during the early harvest period (Larson et al., 1989).
Irrigation, therefore, particularly during the first 4–6 weeks following fruit set,
increases individual fruit size and yield. This is a critical period of fruit
development since it is when cell division is most rapid and cell walls are
developed. Even slight reductions in plant water status during this period can have
adverse effects on fruit growth and retention (Pongsomboon, 1991). Although
drought tolerance of the mango tree is well known, this comes at considerable cost
to tree performance, particularly in areas with prolonged dry seasons that extend
through flowering and fruiting. Irrigation is there-fore one of the most powerful
tools to alleviate non-lethal yet potentially yield-reducing drought stress.

Flooding

Studies with container-grown mango trees have reported variable responses with
respect to tree survival. Larson et al. (1991c) observed that as many as 45% of
trees died after their roots were submerged in water for 4–10 days, but in the
surviving population no further mortality occurred when flooding was extended for
up to 110 days. In other experiments, there was no tree mortality after container-
grown mango trees were flooded from 1 to several months although tree growth
was significantly reduced (Larson et al., 1991c; B. Schaffer unpublished data,
Homestead, Florida, 1993).
The ability of mango trees to survive prolonged flooding appears to be
dependent on the development of hypertrophic (swollen) stem lenticels
immediately above the water line (Plate 42). The initial stages of lenticel
hypertrophy are characterized by the development of intercellular spaces in the
phellem tissue and production of additional phellem tissue by increased phellogen
activity. Later stages of hypertrophy are characterized by the development of
intercellular spaces in the phellem tissue and cortex (Larson et al., 1991a).
Observations vary among studies whether or not trees devel-oped hypertrophic
lenticels or how quickly after flooding they formed. These anomalies have been
attributed to environmental differences at the time of root submersion (Larson et
al., 1991c). In trees that died as a result of flooding stress there was no lenticel
hypertrophy; however, stem lenticels hypertro-phied within 4–10 days on mango
trees that survived flooding (Larson et al., 1991a, c, 1993). Sealing hypertrophic
lenticels of mango trees with silicone grease or petroleum jelly resulted in tree
death within 3 days of flooding, thereby demonstrating their necessity for tree
survival. The role of hypertro-phic lenticels in flood-tolerant species is not clear,
although they are thought to eliminate potentially toxic metabolites such as ethanol,
acetaldehyde and ethylene which result from anaerobic respiration in the roots
(Chirkova and Gutman, 1972; Larson et al., 1993). They may also confer flood
tolerance by enhancing O2 diffusion to the roots (Kozlowski, 1984).

In some instances, adventitious roots have developed above the water line
when container-grown mango trees have been flooded for long periods
Ecophysiology 195

(Schaffer et al., 1994). It is likely that these roots facilitate the absorption and
translocation of O2 to submerged roots and their development is a common
morphological response of many woody plants to root anoxia. The develop-ment of
adventitious roots has not been reported for flooded, field-grown trees and they
may only form on young trunks after extended flooding peri-ods, which usually do
not occur under normal production conditions.
Vegetative growth of mango trees generally declines when trees become
flooded for > 2–3 days. When trees in a limestone soil in containers were flooded
for > 110 days, there was a 94% reduction in shoot extension growth, while
flooding for approximately 10 days resulted in a 57% reduction in shoot extension
growth (Larson et al., 1991c). In a subsequent study, the stem radial growth (a
more sensitive indicator of tree growth than shoot extension growth) of mango
trees decreased 2 weeks after roots were submerged. Flooding for > 14 days also
significantly reduced root dry weight, resulting in an increased shoot to root ratio
(Larson et al., 1991c). These adverse effects of flooding on the growth of mango
trees are expected as reduced net photo-synthesis and presumably higher root
respiration limit the availability of carbon-based assimilates required for growth.

Wind

Most fruit trees benefit from wind protection, particularly during the estab-lishment
years when the disruption of physiological processes results in a significant
depression of growth in young trees. In addition, wind also causes abrasions to the
skin of fruits, particularly when they are small, which develop into unsightly
blemishes by the time they are fully grown thereby reducing quality and market
value. However, the cost of windbreaks may not be offset by higher returns. In
some mango-producing regions, winds are not sufficiently strong to justify the cost
of wind protection. Until recently, wind protection in South Africa was not
recommended for mangoes due to the loss of potential cropping space by ‘living’
windbreaks, their potential to create frost pockets, and the likelihood of promoting
the incidence of flower and fruit diseases through increased humidity (Van der
Meulen et al., 1971); however, the value of windbreaks is well appreciated today in
South Africa (B.N Wolstenholme, personal communication, Pietermaritzburg,
South Africa, 1995).

In studies with ‘Kensington’ mangoes in Australia, where artificial wind-


breaks were constructed to shelter trees from the prevailing summer south-easterly
winds, a 600% increase in yield was recorded in the first year following wind
protection (Mayers et al., 1984). This significant improvement in tree performance
was a result of better growth of trees which set and held more fruit per panicle,
suffered less damage to leaves (cuticle fracturing) and had reduced fruit loss from
bacterial black spot (caused by Xanthomonas camp-estris pv. mangiferaeindicae)
compared to the wind-exposed trees. These results indicate that wind can have a
significant effect on mango productivity from the reduction of both growth and
yield through undisclosed physiological
196 B. Schaffer et al.

mechanisms, and a decreased level of bacterial black spot infection. The pro-vision
of windbreaks in orchards is expensive with decisions to be made on the use of
either ‘living’ or artificial shelters. Requirements for wind protec-tion will vary
depending on site circumstances, and all factors pertaining to crop performance
will require careful consideration.

Salinity

Salt stress in mango trees produces symptoms similar to those described for other
species (Harding et al., 1958; Ehlig, 1960; Kadman, 1964; Bingham et al., 1968).
Mild symptoms of chloride toxicity are scorched leaf tips and margins and leaf
curling, while in more severe cases growth ceases, leaves abscise and trees die.
Necrotic areas develop on leaves of trees exposed to high sodium levels (Jindal et
al., 1976; Kadman et al., 1976; Gazit and Kadman, 1980). Irri-gation water
concentrations of 20–60 mM sodium chloride (NaCl) or sodium sulfate (Na2SO4)
reduced leaf area and changed the branching structure of container-grown mango
trees, suggesting that salinity resulted in reduced leaf cell elongation, and affected
the activity of the terminal meristem (Schmutz and Lüdders, 1993). As the duration
of the exposure to saline con-ditions increased, transpiration decreased
exponentially (Schmutz and Lüd-ders, 1993). In a later study, Schmutz (2000)
found that following a gradual increase in salinity of the nutrient solution applied
to potted polyembryonic ‘13-1’ rootstocks (from 0 to 120 mM NaCl over 15 days),
Amax significantly declined despite there being no visible leaf symptoms of salt
toxicity. The decline in Amax occurred within 6 days of beginning the salinity
treatment, which was 15 mM NaCl for 3 days followed by 30 mM NaCl for 3 days.
This indicates that photoassimilation in mango is quite sensitive to exposure of
trees to salinity.

There is considerable variation in salinity stress of mango, both within and


between populations of mono- or polyembryonic mango ecotypes. Based on the
results of limited studies, there appears to be greater salt tolerance in
polyembryonic than in monoembryonic populations (Jindal et al., 1975; Kad-man
et al., 1976). In seedling populations from mono- and polyembryonic cultivars
irrigated for 2 years with water containing approximately 10 mM chloride, most
plants developed leaf scorching after 6 months which gradu-ally became more
severe, culminating in degeneration and death. However, some seedlings which
had no damage or only slight toxicity symptoms were mostly of the polyembryonic
‘13-1’ rootstock cutivar or related types (Kad-man et al., 1976). Leaf analyses
revealed that the chloride concentration in tolerant seedlings (0.68–0.77%) was
greater than in susceptible seedlings (0.43–0.55%). In addition, tolerant plants had
lower leaf concentrations of potassium, calcium and magnesium than saline-
sensitive seedlings, possibly a result of comparative nutrient dilution since
vegetative growth was greater for saline-tolerant than for saline-sensitive seedlings.
Kadman et al. (1976) also suggested that the mechanism of chloride tolerance in
‘13-1’ was based on greater physiological tolerance of chloride concentrations in
leaf tissues,
Ecophysiology 197

rather than ion exclusion or a selective uptake mechanism common in other species
(Collander, 1941; Walker, 1986). However, the relative sodium toler-ance of ‘13-1’
was due to exclusion of sodium from shoots and its accumula-tion in root cell
vacuoles (Schmutz and Lüdder, 1993). More recently Hoult et al. (1996) reported
significant cultivar differences within a population of 21 polyembryonic mango
cultivars exposed to saline (480 mg/l NaCl) irrigation water for 10 months.
Differences were measured in leaf Na (0.37–1.34%) and Cl (0.39–1.07%)
concentrations but these were poorly correlated with toxicity symptoms on leaves.

Salinization of agricultural land is increasing and in many areas salinity


management is critical. There appears to be sufficient genetic diversity within
Mangifera indica to enable the selection and development of saline-tolerant
rootstocks (Kadman et al., 1976; Gazit and Kadman, 1980; Hoult et al., 1996).
However, quantitative data on the critical limits of soil and water salinity which
mango trees will tolerate without reductions in yield and fruit quality are needed.

Elevated atmospheric CO2 concentration

Growing ‘Kensington’ mango trees for 6 months in a controlled atmosphere


glasshouse with an ambient CO2 concentration of 600 Pmol/mol resulted in more
dry matter partitioned to the roots compared to plants grown in an ambient CO 2
environment of 350 Pmol/mol (Schaffer et al., 1999) (Fig. 6.8). Fruit dry weight
was greater for mango trees grown in an atmospheric CO 2 concentration of 600
Pmol/mol compared to trees grown at 350 Pmol/mol (Schaffer et al., 1999) (Fig.
6.9). Most of the increased total fruit dry weight at

500
350 μmol CO2/mol
600 μmol CO2/mol
400
Dry weight (g)

300

200

100

0
Old Old New New Trunk Roots
leaves branches leaves branches
Plant part

Fig. 6.8. Partitioning of dry matter in ‘Kensington’ mango trees grown for 6 months
in atmospheric CO2 concentrations of 350 or 600 Pmol/mol. Bars represent means
(n = 6 trees) ± standard error (Source: redrawn from Schaffer et al., 1999).
198 B. Schaffer et al.

50
350 μmol CO2/mol
40 600 μmol CO2/mol
Dry weight (g)

30

20

10

0
Skin Flesh Testa Seed Total fruit
Fruit tissues

Fig. 6.9. Partitioning of dry matter in ‘Kensington’ fruit of trees grown for 6 months in
atmospheric CO2 concentrations of 350 or 500 Pmol/mol. Bars represent means (n =
33–49 fruit)  standard error (Source: redrawn from Schaffer et al., 1999).

the higher atmospheric CO2 concentration was a result of increased amount of


pulp; whereas, there were no significant effects of increased atmospheric CO2
concentration on dry matter accumulation in the skin, testa or seed (Schaffer et al.,
1999) (Fig. 6.9). Thus, increasing the atmospheric CO 2 assimi-lation rate increased
growth and partitioning to the mesocarp. Therefore, increased atmospheric CO2
concentration (at least to 600 Pmol/mol) as a result of global climate change may
increase the economic yield of mango (Schaffer et al., 1997). However, under
actual environmental conditions result-ing from global climate change, water and
nutrient availability may be limit-ing and is likely to offset any increases in
biomass or economic crop yield resulting from elevated ambient CO 2
concentrations (Gitay et al., 2001).

6.5 Crop Production

Temperature limitations to crop production

Temperature is probably the most important environmental variable to con-sider


when selecting mango cultivars for particular sites. The mean tempera-ture range
for optimum growth of mango is about 24–30°C (Mukherjee, 1953; Whiley et al.,
1989). However, mango trees can tolerate temperatures up to 48°C for short
periods (Mukherjee, 1953). Mango trees have limited tolerance to cold and trees
are usually severely damaged or killed after a few hours at temperatures < 0°C
(Carmichael, 1958; Campbell et al., 1977). Although mature trees have withstood
temperatures of −4°C for a few hours with lim-ited damage, juvenile trees were
killed after 13 h at −4°C to −6°C (Campbell et al., 1977).
Ecophysiology 199

Monoembryonic mango cultivars tend to be more cold tolerant than


polyembryonic cultivars, probably due their probable subtropical origin. There are
also differences in fruit setting capacity between mono- and poly-embryonic
cultivars at subtropical latitudes, where monoembryonic culti-vars crop more
successfully when minimum temperatures fall below 12°C during flowering
(Searle et al., 1995). Differences in low temperature toler-ance have important
implications for the selection of suitable cultivars for production under specific
climatic conditions.

Light interception and orchard design

Light interception and utilization within tree canopies is a primary consider-ation


in orchard design. Thus, tree spacing as well as pruning practices in orchards are
primarily based on maximizing light for photosynthesis (see previous section on
‘Light’ under Photosynthesis, this chapter). The follow-ing equation describes the
relationship between light interception, photosyn-thesis and yield in fruit tree
orchards:
Biological Yield = (Light Available) (% Light Intercepted) (Photosynthe-
sis) − Respiration (6.12)
where biological yield refers to dry matter production, including that of fruit
(Lakso, 1994). The amount of light available is a function of climate and can-not
be manipulated, and potential net photosynthetic efficiency of a crop is inherent
and cannot be altered without genetic manipulation. Thus, optimiz-ing biological
yield is based on maximizing the percentage of light inter-cepted by the orchard
canopy and minimizing stress so that photosynthetic potential is not compromised.
With respect to light, the mango tree presents special problems due to long-lived
leaves, dense canopies, and the potential for vigorous growth in some cultivars.

Maximizing light interception by the photosynthetic surface of an orchard is a


function of tree spacing, canopy density and height. For some fruit tree trees, for
example apple (Malus domestica) and citrus (Citrus spp.), the use of low vigour
rootstocks and pruning provides opportunities for an array of management options.
However, the lack of dwarfing rootstocks and the com-plications in pruning due to
the floral morphology of mango (inflorescences are predominantly borne
terminally on the most recently produced shoots) limit the efficient harvest of light
with respect to maintenance of productiv-ity. Improved productivity can be
obtained by grafting, which either short-ens the juvenility phase and/or exerts some
control over vigour. Compared to temperate fruit orchards, canopies of commercial
mango orchards have a higher proportion of ‘shade’ to ‘sun’ leaves (Schaffer et al.,
1994). Mango trees are rarely selectively pruned (Young and Sauls, 1981),
although novel ideas of ‘heading back cuts’ after harvest have been investigated
(Oosthuyse, 1992). In some areas, trees are mechanically topped and hedged to
control tree size. Hedging encourages increased complexity (ramification),
resulting in a dense outer canopy. Where costs are not prohibitive, strategically
timed, selective
200 B. Schaffer et al.

pruning to increase the percentage of leaves exposed to > 60% of full sunlight will
increase the photosynthetic efficiency of the canopy with a potential improvement
in yield. Schaffer and Gaye (1989) increased light interception of mango by
removing 25% of the canopy. This resulted in higher chlorophyll concentrations in
leaves of pruned canopies later in the year. Although net photosynthesis was not
measured in that study, it is likely that the higher leaf chlorophyll concentrations in
the pruned canopies resulted in higher photo-synthetic rates.

Light utilization of mango can be enhanced by pruning, but the timing of such
treatments is critical. For example, in the subtropics, shoots produced following
harvest generally flower 3–5 months after being exposed to induc-tive (cool)
temperatures. Therefore, trees should be pruned immediately after harvest to
improve light penetration and contain tree size. However, in the tropics there is a
shorter period between the cessation of summer growth and flowering and
summer-grown shoots of many cultivars fail to induct that year (Scholefield et al.,
1986; see Davenport, Chapter 5, this volume). There-fore, due to the removal of
potential flowering sites, summer pruning of mango trees in the tropics generally
reduces yield in the following season.
Growth control of mango trees through the development of dwarfing
rootstocks, low-vigour cultivars and pruning strategies to increase light pen-
etration within tree canopies and by entire orchards, may improve yield per-
formance of this crop. Improved information on light interception, critical leaf to
fruit ratios and relationships between shoot maturity and floral induc-tion with
respect to genotype/environmental interactions will enhance the development of
improved cultural practices for mango production. It is per-tinent to emphasize that
few evergreen fruit trees are as precocious as mango, or as large at maturity.
Orchardists should take advantage of the precocity and of light interception
principles by initial high-density planting, with hedgerows perhaps the best option.
As trees become crowded, however, effi-ciency of light interception is
compromised and remedial action before this occurs is required to maintain
productivity.

6.6 Conclusions
Although much literature in the past stressed problems associated with low
temperature on mango growth and production, sustained high temperatures with
associated soil moisture stress during fruit ontogeny and high evapora-tive demand
are perhaps the major reasons for relatively low yields in mango orchards
worldwide, especially in the tropics. Although mango trees have a number of
survival mechanisms that allow them to cope with stressful envi-ronments, these
come at a considerable energy cost thereby potentially reduc-ing the availability of
carbon-based inputs for fruiting. It is likely that annual assimilation gains and
resource availability during critical developmental periods are inadequate for
sustained high yields of quality fruit. These prob-lems can be alleviated by
development of improved germplasm with adapta-tion to specific environments
(Whiley et al., 2006). In the future, greater effort
Ecophysiology 201

is required for rootstock development and understanding the manipulative effect on


whole tree physiology. Expansion of human populations and cli-mate change
scenarios predict lesser and poorer quality water available for cropping systems
across tropical and subtropical latitudes. Hence, greater salinity tolerance within
the species should be identified. Knowledge of mango physiology, particularly in
relation to the tree’s responses to varying environmental conditions remains basic
and must be expanded. A greater understanding of these principles together with
their application will assist in the development of more productive cropping
systems.

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7 Fruit Diseases

D. Prusky, I. Kobiler, I. Miyara and N. Alkan


Agricultural Research Organization, Bet Dagan, Israel

7.1 Introduction 210


7.2 Anthracnose 211
The pathogenesis strategy 212
Epidemiology – the disease cycle 213
Mechanisms of fungal pathogenicity and fruit resistance 214
Management 216
7.3 Alternaria Rot (Black Spot) 219
Pathogenesis 219
Mechanisms of fungal pathogenicity and fruit resistance 220
Management 220
7.4 Stem-end Rots 221
Pathogenesis 221
Epidemiology 222
Management 223
7.5 Other Minor Diseases 224
7.6 Conclusions 224

7.1 Introduction
Postharvest diseases can reduce fruit quality and cause severe economic losses, due
to decay resulting in completely unmarketable and blemished fruits that are often
sold in less demanding local markets, where the prices are considerably lower than
export prices. It is clear to the producer that quality at the time of harvest cannot be
improved but merely maintained for a limited period of time. Harvesting fruits at
the optimal stage, with respect to size and maturity, can, therefore, ensure peak
quality and maximum shelf-life potential. Thus, managing total tree health can
contribute to reducing postharvest losses. It is known that older and neglected
orchards may become a profuse inoculum source for postharvest diseases.
Furthermore, preharvest
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
210 (ed. R.E. Litz)
Fruit Diseases 211

stress factors such as excess or shortage of water, fluctuating or extreme envi-


ronmental conditions and high nitrogen levels (Hawthorne, 1989) can render fruit
more susceptible to postharvest diseases. After harvest, improper treat-ment of
fruits through storage at non-optimal temperatures accelerates fruit deterioration as
a result of enhancement of normal physiological processes such as respiration and
ethylene production (Thompson et al., 2002). More-over, excessive temperatures in
the field during harvesting, in transit and in the packing house can render tropical
fruit more susceptible to chilling injury and can contribute to the development of
disease. Combinations of high tem-peratures and high relative humidity (RH)
favour the growth of postharvest pathogens, and can contribute to the development
of disease at the retail end (Banik et al., 1998). Proper, uninterrupted cooling,
therefore, protects quality and extends the shelf life of produce. Precooling of
products (Thompson et al., 2002), and application of top or liquid icing, vacuum,
hydrocooling and forced air cooling (Thompson et al., 2002) are examples of
effective alterna-tive methods that can be used to remove field heat and to restrict
pathogen growth. Effective cold-chain management is crucial to ensuring product
integrity and preventing postharvest pathogens from spoiling produce in transit or
shortly after arrival (Lizada, 1993). Furthermore, low-temperature storage
conditions are generally not conducive to disease development, a fact that can be
exploited to ensure quality and extend shelf life (Banik et al., 1998). However,
improper cooling during shipping, or interrupted cooling will also promote
microbial growth, resulting ultimately in product spoilage. In practice, several
breaks in the cold chain might have significant impacts on decay development.

Among the postharvest diseases of mango, anthracnose is the most prev-alent


in humid growing regions. The incidence of this disease can reach almost 100% in
fruit produced under wet or very humid conditions. Other common postharvest
diseases of mango in humid areas are stem-end rot, caused by Lasiodiplodia
theobromae (Pat.) Griffon & Maubl. (Sangchote, 1998) and Dothiorella
dominicana Petrak. et Cif. (Johnson and Sangchote, 1994), and black mould rot,
caused by Aspergillus niger van Tieghem. Mango black spot, caused by Alternaria
alternata (Fr.:Fr.) Keissl. (Prusky et al., 1983), is prevalent in dry countries.

7.2 Anthracnose
Mango anthracnose is caused by Glomerella cingulata (Stoneman) Spauld. & H.
Schrenk (anamorph: Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. In
Penz.), C. gloeosporioides Penz. var. minor J.H. Simmonds (Fitzell and Peak,
1984) and Colletotrichum acutatum J.H. Simmonds (Freeman et al., 1998). The
pathogen also causes blossom blight, leaf blight and, in severe cases, tree dieback
(Ploetz, 1994; Ploetz and Prakash, 1997). The form-genus Colletotri-chum Corda
(form-order Melanconiales; form-class Coelomycetes; subdivi-sion
Deuteromycotina) comprises imperfect fungal species which exist as Glomerella
(subdivision Ascomycotina) in their sexual, teleomorphic or perfect
212 D. Prusky et al.

state. These fungal pathogens occur worldwide, and the genus is synony-mous with
anthracnose. Leaf anthracnose appears as irregular black necrotic spots on both
sides of the mango leaves. Lesions often coalesce and form large necrotic areas,
frequently along the leaf margins. Lesions develop pri-marily on young tissue, and
conidia are formed in lesions of all ages. Under favourable conditions, the fungus
can invade the twigs and cause dieback (Ploetz et al., 1996). Panicle anthracnose or
blossom blight can affect both the inflorescence stalk and the individual flowers; in
the stalk, elongated dark grey to black lesions appear; the blighted flowers are dry,
and their colour ranges from brown to black. Fruits smaller than pea-size can be
infected and abort; whereas, larger fruits that are aborted because of normal self-
thinning or other physiological causes are usually mummified. The resulting
mummi-fied fruit are invaded saprophytically by C. gloeosporioides, and the
fungus sporulates abundantly on them.

Although field anthracnose causes considerable damage, the vast losses


inflicted by postharvest anthracnose are of far greater economic importance.
Postharvest anthracnose appears on the fruit surface, as rounded brown to black
lesions with an indefinite border. Lesions > 2 cm in diameter are fairly common.
Lesions of various sizes can coalesce and cover extensive areas of the fruit,
typically in a tear-drop pattern that develops from the basal towards the distal end
of the fruit. Lesions are usually restricted to the peel, but in severe cases the fungus
can invade the pulp. In advanced stages of the dis-ease, the fungus produces
acervuli, and abundant orange to salmon-pink masses of conidia appear on the
lesions.

The pathogenesis strategy

Colletotrichum gloeosporioides is regarded as a hemibiotrophic species that


commences its invasion with a transient post-penetrative asymptomatic bio-trophy,
characterized by a temporary confinement with a localized mode of intracellular
hemibiotrophy. This is succeeded by a phase of destructive necrotrophy that
culminates in the appearance of disease symptoms and pro-duction of conidiomata
by the pathogens (Prusky and Plumbley, 1992; Latunde-Dada et al., 1999).
Quiescent species pass through a prolonged phase of pre-penetrative growth that is
arrested in synchrony with the physiological state of the infected organ.

Following its landing on the fruit or host tissue, the ungerminated aseptate
conidium differentiates to a melanized appressorium. In the case of Colletotrichum
lindemuthianum, appressorial adhesion is mediated by the man-nose- and
galactose-rich glycoproteins of the extracellular matrices, which are secreted
during appressorial differentiation (Pain et al., 1996). Both the surface wax of the
specific fruit and the hard surface of the tissue are recog-nized as host signals that
selectively trigger germination and appresso-rium formation solely by the conidia
of C. gloeosporioides (Podila et al., 1993; Liu and Kolattukudy, 1998). The
melanin of the melanized appressoria pro-tects the fungus and behaves as a
permeability barrier, and the appressoria
Fruit Diseases 213

contain osmolytes (i.e. glycerol) that are needed for generating the osmoti-cum
from which high turgor pressure will drive the invasive forces of the penetrating
hyphae through the small appressorial pores (0.5 Pm in Col-letotrichum
sublineolum). The osmolyte is generated by the metabolism of stored glycogen,
trehalose and lipids and, at an in vivo concentration of at least 3.3 M in mature-
melanized appressoria (de Jong et al., 1997), this osmo-lyte is capable of
generating turgor pressures as high as 8 MPa (Howard et al., 1991; Money and
Howard, 1996). Melanized appressoria appear to be quite capable of a forcible,
non-enzymatic penetration of an intact host surface. However, Dickman et al.
(1982) suggested that attack on papaya by C. gloeo-sporioides depended on
cutinase production by the pathogen. The germi-nated appressoria in
Colletotrichum develop single infection hyphae that grow and extend into the waxy
cuticle, reach the first layers of pericarp cells, and then become quiescent for long
periods of time (Coates et al., 1993; Prusky, 1996).

In recent years, the taxonomy of Colletotrichum has been clarified by the


adoption of molecular biological techniques that involve the use of poly-merase
chain reaction amplification, alignment of nucleotide sequences, and the
construction of dendograms, phylogenetic trees and similarity matrices from the
data generated. For example, Sherriff et al. (1994) and Sreenivasap-rasad et al.
(1996) used homologies between the nucleotide sequences from amplified non-
transcribed regions (internally transcribed spacers 1 and 2 (ITS1, ITS2)) and the
large subunits (domains 1 and 2) of ribosomal DNA extracted from a wide range of
isolates to both justify and resolve the taxo-nomic status of Colletotrichum species.
Bailey (1997) proposed the adoption of species aggregates based on these and other
data.

Epidemiology – the disease cycle

Conidia are formed abundantly in the mango canopy (Fitzell and Peak, 1984)
which, therefore, is considered to be the primary source of inoculum. In the field,
C. gloeosporioides produces conidia on lesions on leaves, twigs, panicles and
mummified fruits (Arauz, 2000). Conidia can be rain-splashed onto other leaves or
flowers, where they can cause secondary infections. Developing fruit can be
infected, and some isolates can cause preharvest fruit loss (Gan-totti and Davis,
1993). In the case of postharvest anthracnose, developing fruits are infected in the
field, but the infections remain quiescent until the onset of ripening, which occurs
after harvest. Once the climacteric period of the fruit starts, lesions begin to
develop but there is no fruit-to-fruit infection. In this context Prusky and co-
workers (Guetsky et al., 2005) suggested that the capability of C. gloeosporioides
to cause early disease symptoms in unripe fruits depends on the activation of
laccases by specific strains of the fungus. Details of these systems are discussed in
the following section.
Colletotrichum gloeosporioides requires free water or RH > 95% to enable
conidial germination and appressorium formation (Fitzell et al., 1984; Dodd et al.,
1991). However, conidia can survive for 1–2 weeks under low RH and
214 D. Prusky et al.

then germinate if exposed to 100% RH (Estrada et al., 1993). In general, infec-tion


is favoured at temperatures ranging from 20 to 30C.

Mechanisms of fungal pathogenicity and fruit resistance

Unripe mango fruits are reservoirs of extremely high concentrations of pre-formed


antimicrobial compounds. This arsenal of constitutive resistance weapons may
accumulate in the immature pericarp, but not in the mesocarp, at concentrations in
fruit fresh weight of up to 220 Pg/g. They include mix-tures of 5-substituted
resorcinols such as resorcinol-5-(12-heptadecadienyl) and resorcinol-5-
(pentadecyl) (Droby et al., 1986, 1987; Prusky and Plumbley, 1992; Prusky, 1996)
(Fig. 7.1). Faced by such a chemically adverse environ-ment, the fungus usually
postpones its development until the concentrations of the compounds in the host
decline. The capability to penetrate the host without encountering the arsenal of
passive chemical resistance or triggering the weapons of active resistance is
inherent in a number of fruit-infecting Colletotrichum species such as C.
gloeosporioides, C. acutatum and A. alternata (Prusky and Keen, 1993). The
conidia, melanized appressoria and penetra-tion pegs of these quiescent species
have evolved mechanisms of physiologi-cal inactivity that enable them to pause
until the host-ripening process and the decline of antifungal compounds starts, thus
enabling the avoidance of host defences that exist within unripe, pre-climacteric
fruits. Colletotrichum gloeosporioides is a field-to-store pathogen whose conidia
infect during the preharvest growth of fruits. In fruits such as mango and avocado,
termina-tion of fungal quiescence is associated jointly with the climacteric
production of ethylene during fruit ripening and the onset of dramatic reductions in
the levels of preformed fungitoxic resorcinols (Droby et al., 1986, 1987).
Flaishman

Resorcinol-5-(12-heptadecenyl) Resorcinol-5-(pentadecyl)
HO HO HO HO

(CH2)11 (CH2)14

CH CH2

CH CH3

(CH2)3

CH3

Fig. 7.1. Chemical structure of the 5-substituted resorcinols isolated from


mango fruits.
Fruit Diseases 215

and Kolattukudy (1994) hypothesized that ethylene is involved in terminat-ing


quiescence, by inducing appressoria formation and hyphal growth, which strongly
suggests that Colletotrichum spp. must have coevolved with their hosts, to develop
a mechanism that uses the host’s ripening hormone as a signal to reactivate the
infection process. However, when Prusky and co-workers (Prusky et al., 1996)
applied ethylene to unripe fruits they could not enhance the termination of
quiescence as had been suggested, possibly indi-cating that ethylene is a signal that
induces appressoria formation in vitro, but that does not enhance fungal growth in
the fruit. Furthermore, since fun-gal infection and appressoria formation occur in
the orchard, it is difficult to conceive how ethylene produced much later, during
ripening and storage, could enhance appressoria formation on the fruit. Regardless
of the mecha-nisms that may be involved in the onset and termination of
quiescence, qui-escent infection appears to be a case of coevolution; it is
advantageous to both pathogen and host in a natural ecosystem to allocate chemical
defences to the immature fruit but not to the ripe fruit (Prusky and Keen, 1993).

Quiescence also may result from a localized host response that is often
associated with an oxidative burst, i.e. production of reactive oxygen species
(ROS). Localized generation of ROS during quiescence was found to be one of the
earliest (within 2–3 h) detectable cytological defence responses to attempted
penetration of unripe, resistant avocado fruits by C. gloeosporioides. Beno-
Moualem and Prusky (2000) proposed that fungal infection of unripe fruits leads to
production of ROS during quiescence, at the infection loci surrounding the
germinated appressoria and their penetration hyphae. This localized ROS
production would activate the synthesis of antifungal com-pounds or compounds
that inhibit fungal metabolism (Ardi et al., 1998) at the infection loci, thereby
enhancing and/or preserving the levels of antifungal compounds and, in turn,
inhibiting fungal development and imposing quiescence.

Guetsky et al. (2005) suggested that initiation of quiescence could result from
the fungal capability to induce laccase activity that modulates the metab-olism of
preformed antifungal compounds and the activation of the quies-cent C.
gloeosporioides that occurs during fruit ripening. Early activity of aggressive
isolates in unripe fruits included increased laccase activities that resulted in early
metabolism of preformed antifungal compounds leading to early appearance of
symptoms.
When the levels of toxic compounds in the fruit peel decline, C. gloeospo-
rioides can also enhance its colonization of ripening fruits dynamically by locally
altering the pH of the fruit at the infection site to suit the increased expression of
pathogenicity factors and the enzymatic arsenal (Yakoby et al., 2000; Prusky et al.,
2001; Eshel et al., 2002; Prusky and Yakoby, 2003). In the pathogen C.
gloeosporioides, the gene pelB encoding for the enzyme pectate lyase, a key factor
for virulence, is expressed when the pH is > 6.0, a value at which decay is initiated
in the tissue (Yakoby et al., 2000, 2001). Also in the case of A. alternata, another
pathogen of mango fruits, the expression of the endoglucanase gene AaK1 is
maximal at pH levels > 6.0 (i.e. values that are characteristic of decayed tissue).
AaK1 is not expressed at the lower pH
216 D. Prusky et al.

values at which the pathogen is quiescent (Eshel et al., 2002). Ambient alkal-
ization by Colletotrichum is achieved by active secretion of ammonia, which is
produced as a result of protease activity and deamination of amino acids. The
pathogenicity of C. gloeosporioides and expression of the virulence factor PL-B
both depend on raising the ambient pH (Drori et al., 2003). This modu-lation of
environmental pH has been used as the basis for a new approach to disease control
in mango fruits, and is discussed below.

Management

Control of postharvest anthracnose can be achieved by field management,


postharvest treatments or a combination of both. Management of mango
anthracnose in the field involves cultural and chemical practices, as well as cultivar
selection.

Cultural control
Since the development of mango anthracnose is dependent on moisture or high RH,
orchards ideally should be established in areas with a well-defined dry season, to
allow for fruit development under conditions unfavourable for disease
development. In the tropics, mango flowering usually occurs dur-ing dry seasons,
but anthracnose incidence of > 90% is common in fruits that develop during the
rainy season (Arauz, 1999). In contrast, the incidence and severity of mango
anthracnose can be close to zero in fruits that develop completely in the dry season,
without the application of any other control measure (Arauz, 2000).

Considerable effort has been invested in understanding and managing mango


flowering. Flowering can be advanced by several weeks by applying potassium
nitrate sprays to mature foliage (Núñez-Elisea, 1985). The growth retardant
paclobutrazol, alone or followed by potassium nitrate sprays, can also be used to
advance flowering (Núñez-Elisea et al., 1993). Both treatments could contribute to
the manipulation of the flowering season to a less sensi-tive period. Field sanitation
of the tree itself is difficult to practise. Elimina-tion of dry panicles and mummified
fruits is time consuming. Bagging results in reduced anthracnose severity, but it
also reduces the red colour of the fruit, which could reduce consumer appeal
(Hofman et al., 1997).

Resistant cultivars
Although all commercial mango cultivars are susceptible to anthracnose, some are
less susceptible than others; ‘Tommy Atkins’ and ‘Keitt’ are less susceptible than
‘Irwin’, ‘Kent’, ‘Haden’ and ‘Edward’ (Campbell, 1992).

Chemical control
In extreme situations, in which fruit develops entirely under disease-favouring
conditions, seasonal applications of up to 25 sprays of protective and sys-temic
fungicides have been reported (Dodd et al., 1997). However, few fungi-cides are
approved in importing countries for use on mango. Therefore, the
Fruit Diseases 217

choice of fungicides depends on the intended destination of the fruit. Dithio-


carbamate fungicides are highly effective for anthracnose control but can be used
for only a few specific countries, whereas copper fungicides are recom-mended,
but their efficacy is lower (Arauz, 2000). Fungicides with erradicant activity for
mango anthracnose include benzimidazoles and the imidazole prochloraz. Benomyl
has been used under calendar-based schedules, usually in a mix with protectant
fungicides, to delay the build-up of resistance in the pathogen population.
Prochloraz has been used as a protectant or as an erad-icant spray (Estrada et al.,
1996), but is mainly used only as a postharvest treatment.

Biological control
A number of microorganisms with in vitro or in vivo activity against C. gloeo-
sporioides have been isolated (Jeffries and Koomen, 1992), but few examples had
been used commercially in the field until Korsten (2004) isolated Bacillus spp.
from leaf and fruit surfaces, and effectively controlled anthracnose of mango.
Postharvest control was achieved with semi-commercial preharvest sprays or
postharvest packing house dips, sprays or ultra-low-volume appli-cations.
Integrated treatments involving antagonists combined with quarter-strength or
recommended dosages of fungicides such as prochloraz or sodium hypochlorite
also effectively suppressed postharvest anthracnose of mango. Commercializing
the antagonists in South Africa (Korsten, 2004) and in the Philippines proved to be
difficult because of the limitations set by the local registration guidelines, and the
effect of product formulation on antagonist performance in commercial
applications.

Postharvest control
Traditionally, postharvest control of mango anthracnose has aimed at eradi-cation
of quiescent infections on the fruit. Such eradication is achieved com-mercially by
thermal and chemical treatments, or a combination of both (McMillan, 1987).
Dipping fruit in hot water alone is moderately efficient; temperatures of 50–55C
for 3–15 min have been recommended, with the higher temperatures corresponding
to the shorter exposures. Fruit from Latin America entering the USA market must
undergo a quarantine hot water treatment to eliminate fruit fly (Ceratitis capitata
and Anastrepha spp.) larvae; the fruit is immersed in water at 46C for 90–120
min, depending on variety and fruit size. The efficacy of the fruit fly quarantine
treatment varies from 60 to 85% for elimination of anthracnose infections
(McGuire, 1991). Hot water treatments leave no chemical residue on the fruit and
could be a good anthra-cnose control option for organically-produced mangoes or
for mangoes tar-geted for markets in the USA, where no fungicides are currently
approved for postharvest use. Temperature and time controls are critical, because
fruits can easily be damaged by overexposure to heat. Several fungicides have been
applied after harvest to control mango anthracnose. Benomyl, at rates vary-ing
from 500 to 1000 Pg/ml, was used as a postharvest dip in the past, but its use is no
longer permitted. Thiabendazole at 1000–2000 Pg/ml is also effec-tive, and there is
interest in its registration for postharvest use with mango,
218 D. Prusky et al.

as it is currently used on other fruits such as citrus. Prochloraz can be used with
fruit shipped to the European Union (EU), at rates up to 1000 Pg/ml; its efficacy
ranges from 65% under very high disease pressure to 94% under moderate disease
pressure (Arauz, 2000). One advantage of benzimidazole fungicides (i.e. benomyl
or thiabendazole) is that they are also effective in controlling stem-end rot caused
by L. theobromae, which is the second most important postharvest disease of
mango in tropical areas. Imidazoles such as prochloraz and imazalil are not
effective against L. theobromae on mango (Estrada et al., 1996). The combination
of hot water and fungicides is the most effective commercial postharvest treatment
for the control of mango anthra-cnose; the rate of fungicide application and the
duration of exposure to hot water are both lower, and efficacy is higher, than either
treatment used sepa-rately. The hot water and the fungicides can be applied
sequentially or together. Irradiation of fruit to control anthracnose has been
attempted: gamma irra-diation was not successful (Spalding and Reeder, 1986), but
a short-wave infrared radiation treatment developed in South Africa resulted in
anthra-cnose levels similar to those resulting from the commercial hot-water treat-
ment, and was much faster and less expensive (Saaiman, 1996a).

Prusky and Keen (1993) suggested that it might be possible to prolong the
period of fruit resistance and to delay the onset of anthracnose develop-ment until
the fruit ripens. The climacteric rise occurs simultaneously with the production of
ethylene and with changes in external and internal colour, flavour, aroma and
firmness, and with a reduction in fruit resistance to fun-gal attack (Prusky and
Keen, 1993). Postharvest practices such as cold stor-age and controlled atmosphere
storage maintain resistance to decay by delaying the ripening process, but there are
two limitations to the potential benefit of this approach in mango. First, mangoes,
similarly to many other tropical and subtropical fruits, are sensitive to chilling and
are injured at tem-peratures < 10–13C, depending on the cultivar and the duration
of expo-sure. Second, once the fruits are allowed to ripen under ambient
conditions, disease develops normally. Nevertheless, some progress has been made
towards anthracnose control through maintaining fruit resistance beyond the onset
of ripening. In the last decade, a better understanding of the physi-ological basis of
quiescent infections on tropical and subtropical climacteric fruits has been
achieved, mainly through the work of D. Prusky and his co-workers (1993). The
decline in antifungal compounds, which is brought about by oxidative processes,
can be delayed so that it occurs closer to full ripeness. In avocadoes and mangoes,
the concentrations of antifungal com-pounds were enhanced and, consequently, the
decline in concentration was delayed, by exposure of fruit to an atmosphere
containing 30% carbon diox-ide (CO2) for 24 h, or by treatment of fruit with the
antioxidant compound butylated hydroxy anisole (Prusky, 1988; Kobiler et al.,
1998). The delay resulted in less disease in ripe fruit.

Postharvest biological control of anthracnose in mangoes has been attempted.


In an investigation with a strain of a Bacillus sp. that exhibited in vitro activity
against C. gloeosporioides, it was found that disease control in vivo was obtained
when fruits were inoculated with the bacterium 24 h prior
Fruit Diseases 219

to inoculation with the fungus, but not when fruit were inoculated with the
pathogen first (Korsten et al., 1992), which indicated that the quiescent phase of
the fungus was not affected by the antagonist. Other approaches to disease control
using biological methods included the use of a non-pathogenic strain of
Colletotrichum magna that colonizes the fruit endophytically and prevents
infection by C. gloeosporioides (Prusky et al., 1993); and the expression of an
antifungal peptide in the yeast Saccharomyces, which controlled postharvest
diseases caused by C. coccodes (Jones and Prusky, 2001).

7.3 Alternaria Rot (Black Spot)


Alternaria alternata (Fr.:Fr.) Keissl. causes black spot of mango. Conidiophores,
arising singly or in small groups, are simple or branched, straight or bent, and
sometimes geniculate, and pale- to mid-olivaceous or gold in colour, and smooth in
texture. The conidia are oboclavata; they are borne in long chains in culture, and
most have three to five septa.

Pathogenesis

Germinated conidia penetrate mainly through wounds and specifically through


lenticels of the fruits, and then become quiescent.

Symptoms
Alternaria rot of mango has been increasingly reported as an important pathogen
that causes blossom disease and postharvest fruit rot in ripening fruits in Australia,
Egypt, India, Israel and South Africa. The symptoms are small, black circular spots
that develop around the lenticels. Initially, the spots are concentrated around the
stem end of the fruits, where there are large numbers of lenticels. The spots can
grow and coalesce to become a sin-gle spot that covers a significant part of the fruit
surface. At first, the decay is firm and does not penetrate the pulp more than 1–2
mm, but later the disease progresses into the flesh, which darkens and becomes soft
(Prusky et al., 1983). Symptoms of alternaria rot are more limited, darker and
firmer than those of anthracnose. The former pathogen also attacks mango leaves,
and symptoms can be observed throughout the year. The pathogen may also attack
mango inflorescences, resulting in a significant decrease in fruit set.

Epidemiology
The main sources of inoculum are conidia released from infected leaves, twigs and
inflorescences; however, Alternaria spores easily can be found in all the dry tissues
of mango trees in the orchard. Conidia are transferred to the fruit by air currents
and in dew runoff (Ploetz et al., 1994). Germination of conidia depends on the RH
in the orchard during fruit growth. The area of quiescent Alternaria infection on
mango fruit at harvest increased as the num-ber of hours of exposure to RH  80%
increased over 320 h (Prusky et al.,
220 D. Prusky et al.

1983). Regions with the highest potential for disease incidence are located close to
the 85–90% RH isolines during the fruit growth period. The interme-diate regions
lie between 75 and 85% RH, and the lowest potential risk is in the dry regions,
where the prevailing RH is < 75% (Prusky et al., 1992).

Mechanisms of fungal pathogenicity and fruit resistance

As in the case of anthracnose, the mechanism of resistance against Alternaria is


also related to the presence of high concentrations of preformed antimicro-bial
compounds (Droby et al., 1986, 1987). Susceptibility was found to depend on the
decline of the compound to subfungitoxic concentrations, which occurred faster in
‘Tommy Atkins’ than in ‘Keitt’.
Alternaria alternata can also alter the local fruit pH at the infection site
dynamically, to match the increased expression of pathogenicity factors and the
enzymatic arsenal (Eshel et al., 2002; Niem et al., 2007). In the case of A.
alternata, the expression of the endoglucanase gene AaK1 was found to be
maximal at pH levels > 6.0, which are characteristic of decayed tissue; it was not
expressed at the lower pH values at which the pathogen was quiescent (Eshel et al.,
2002). Alkalization of the ambient environment by Alternaria is achieved by active
secretion of ammonia (Eshel et al., 2002), probably as a result of protease activity
and deamination of amino acids. This modulation of environmental pH has been
used as the basis of a new approach to disease control in mango fruits, and is
discussed below.

Management

The losses caused by alternaria rot can be minimized by a regular field-spraying


programme and by postharvest fungicide treatments.

Field and postharvest chemical control


Preharvest treatments with dithiocarbamate fungicides inhibit the develop-ment of
latent infection. Three sprays with the protectant fungicide maneb, starting 2 weeks
after fruit set, are most effective for disease control (Prusky et al., 1983). However,
since quiescent infections do not develop until after harvest and ripening, the
application of a postharvest treatment by spraying the fruits on the packing line
with 900 Pg/ml prochloraz is simpler and more efficient than the preharvest
fungicide treatment.
Control of alternaria rot is significantly improved by a combination of physical
and chemical treatments. The physical treatment includes a 15–20 s hot water
spraying and brushing (HWB) treatment at temperatures between 50 and 55C
(Prusky et al., 1999). This new approach improved fruit quality at the same time as
it reduced disease incidence. If this physical treatment is followed by a 250 Pg/ml
prochloraz spray it can further improve disease control. Prusky et al. (1999)
concluded that the type and strength of the post-harvest treatment should be varied
according to the level of quiescent infection
Fruit Diseases 221

of A. alternata at the time of harvest. Although the fungicide prochloraz is very


effective for the control of postharvest disease, a milder postharvest treat-ment,
such as chlorine at 500 Pg/ml, can be applied to fruit in which a low incidence of
quiescent infections is found at harvest (Prusky et al., 2002).
This postharvest physical-chemical treatment has been further improved in
light of the recent finding that A. alternata pathogenicity may modulate the pH of
the host environment to promote colonization (Eshel et al., 2002; Prusky and
Yakoby, 2003; Prusky and Lichter, 2007). Application of a combination of HWB
for 15–20 s, followed by spraying with 50 mM hydrochloric acid (HCl), effectively
controlled alternaria rot in stored mango fruit. Similar HWB treat-ments followed
by spraying with reduced concentrations of prochloraz at 45 Pg/ml in 50 mM HCl
inhibited alternaria rot development better than treatment with HCl alone (Prusky
et al., 2006). The enhancement of prochlo-raz activity in acidified solutions was
attributed to its enhanced solubility under acidic conditions, which resulted in an
increase in the fungicidally active ingredient in the solution. This technology
represents the latest devel-opment in the control of postharvest diseases, and
provides a simple treat-ment for the control of diseases that alkalinize the host
environment, including both alternaria rot and anthracnose.

7.4 Stem-end Rots


Stem-end rots of mango fruit present one of the most serious postharvest problems
that affect this industry worldwide. They become more prominent as orchards
become older. Losses increase when fruits are stored for pro-longed periods at low
temperatures or when fruits ripen at temperatures
> 28C. The stem-end rot diseases are caused by a variety of fungal patho-gens
including: D. dominicana (anamorph of Botryosphaeria dothidea), Dothi-orella
mangiferae, L. theobromae (Botryodiplodia theobromae), Phomopsis mangiferae
and Pestalotiopsis mangiferae, among which Botryosphaeria is the dominant
pathogen (Darvas 1991; Johnson et al., 1992; Sangchote, 1998). Stem-end rot
diseases cause heavy losses in mangoes during storage (Prusky et al., 1992; Mitra,
1997; Kobiler et al., 2001).

Pathogenesis

Johnson and co-workers (1993) have suggested that spores of the pathogen may
germinate and penetrate into the host tissue through wounds, and remain as
endophytes in the branches of mango trees.

Symptoms
Depending on the fungus involved, a variety of symptoms may develop at the stem
and as the fruit ripens. Infections by L. theobromae (B. theobromae), D.
dominicana synonym Fusicoccum aesculi (anamorph of B. dothidea) and D.
mangiferae, cause the formation of diffuse, water-soaked tissue that spreads
222 D. Prusky et al.

from the stem end as fingerlike projections, which darken and coalesce into
circumpedicular lesions or lobed margins (Johnson et al., 1992; Slippers et al.,
2004, 2005). Necrosis remains beneath the cuticle and may penetrate through-out
the fruit flesh. Superficial mycelia may appear around the pedicel and ruptures of
the skin or, in some cases, may penetrate through the epidermis. The ascomata of
D. mangiferae are initially embedded, either separately or grouped in complex
stromata, and they finally erupt through the epidermis and open. The spore wall is
dark and thick, and becomes thinner towards the end. Conidiophores are hyaline,
cylindrical, smooth and branched at the base. A watery fluid may drain from the
stem or from surface ruptures (Kor-sten, 2004). Diseased fruit could infect a whole
box of fruits by direct contact, and thereby spread the pathogen in the box. In the
case of injured fruit, lesions could appear at some distance from the stem. Stem-
end rot diseases also have a major effect on the flavour of the fruit.

The disease may also cause tip and branch dieback and cankers on mango trees
(Ramos et al., 1991). Anamorph morphology is commonly used to iden-tify
Botryosphaeria spp. (Jacobs and Rehner, 1998; Slippers et al., 2004), but the
morphological distinctions among the anamorphs of some of the closely related
species are not clear. Recent studies based on DNA sequence data have highlighted
taxonomic groups and relationships in Botryosphaeria (Jacobs and Rehner, 1998;
Slippers et al., 2004). These data, combined with morphological characteristics,
could clarify the current taxonomic confusion.
Phomopsis mangiferae is a weak parasite of less economic importance than the
species above; it produces a dark, circumpeduncular lesion with defined edges that
spreads relatively slowly but penetrates deeply into the flesh. Fruiting bodies may
develop on the affected surface. Phomopsis mangiferae can also be distinguished
by a dark pinhead-size pycnidial fruiting body (Johnson et al., 1992). The lesion
caused by Phomopsis may be similar to the stem-end symptoms cause by C.
gloeosporioides and A. alternata. However, the lesions of the latter two diseases
penetrate only to a depth of 10–20 mm.
Lesions of L. theobromae can be distinguished by their wrinkled black edges
which have a velvety appearance. In the dark zone, pycnidial masses are formed
(Johnson et al., 1992). In affected plants, twigs die back from the tips and into old
wood, giving a scorched appearance to the limb with abun-dant gum secretions
from branches, stems and the main trunk. Pestalotiopsis mangiferae appears as
silvery grey spots that vary in size, and are usually sharply outlined by a dark
border. The fungus may appear as a member of the complex of stem-rot pathogens.

Epidemiology

The pathogens causing stem-end rots initiate inoculation in the orchards as the trees
age. They are enabled to do so by mismanagement or neglect of orchards, and are
affected by periods of rain and high RH at the beginning and end of the dry season
(Johnson et al., 1991; Lonsdale, 1993; Johnson and Sangchote, 1994; González et
al., 1999). Hyphae of the fungi colonize the
Fruit Diseases 223

floral parts of mango trees, develop endophytically in healthy tissue of all parts of
mango plants, especially in the fruit pedicel, and remain quiescent until the fruit
matures, at which stage the parasite is ready to infect through the stem end by
developing in the xylem vessels of the maturing fruit (John-son et al., 1992).
Pathogens can colonize flower parts, remain inactive pend-ing button abscission
and then penetrate the stem end, as in the case of Diplodia natalensis in citrus
(Nadel, 1944), of which no sexual stage has been found in nature. Fruits can be
also infected at the stem by the soilborne patho-gen L. theobromae (anamorph of B.
theobromae), if the fruits are placed on the ground (Johnson et al., 1992) after
harvest. It is also possible that infection can result from transfer of spores by
insects; decayed fruits produce volatiles, which are hypothesized to be attractants
of an insect that could be a vector of the fungal spores (Nago and Matsumoto,
1994).
The range of pathogens that cause stem-end rot is influenced by tempera-ture,
moisture stress and the nutrition status of the host. The initial systemic infection
plays a crucial role in establishing blossom-blight infection. How-ever, secondary
infection is apparently an even more important factor in devel-opment and
incidence of soft stem-end rots. Secondary infections occur when rain washes
spores away from various inoculum sources such as leaves and stems (Saaiman,
1996b). Endophytic Botryosphaeria spp. were found to be espe-cially prominent in
trees continually exposed to water shortage and salt stress (Grobler et al., 2002),
and Botryosphaeria spp. were found to move into develop-ing fruit, resulting in
postharvest stem-end rot development (Lonsdale, 1993).

Management

Field and postharvest control


A variety of postharvest practices can be used to delay disease develop-ment.
These include low-temperature storage and modified- or controlled-atmosphere
storage; however, the management of stem-end rots is far from being perfected.

Cultural control
Johnson et al. (1992) demonstrated that infection of mango fruit before har-vest
occurred through endophytic colonization of pedicel tissue by Botry-osphaeria spp.
present from previous growth flushes, and the possibility of pruning to promote
growth flushing was tested as a means to reduce inocu-lum in the stem tissue from
which new-season inflorescences emerged. Cooke et al. (1998) reported that the
levels of endophytic organisms such as Botry-osphaeria spp. were reduced
significantly when a pruning programme was implemented in mango orchards as a
preharvest control measure. Korsten (2006) found that prevention of water stress
during fruit development and maturation, and avoidance of placing fruits on the
ground suppressed dis-ease development. He also suggested that harvesting of
immature fruit should be avoided; fruit should be cooled to 13C immediately after
harvest and chemical treatment, and stored in a well-ventilated place.
224 D. Prusky et al.

Chemical control
Postharvest control of Botryosphaeria spp. was achieved by postharvest dip-ping,
spraying or ultra-low-volume application of benomyl (where possible). Prochloraz
or sodium hypochlorite also effectively suppressed postharvest stem-end rot of
mango (Plan et al., 2002; Korsten, 2006). A combined treat-ment of wax and hot
water (55qC) provide very effective control of most postharvest pathogens
(Sangchote, 1998), but in some cases partial-vacuum infiltration improved disease
control, which suggests that control efficiency may have been reduced because the
fungicide did not reach the pathogen (Plan et al., 2002).

A treatment consisting of HWB and prochloraz followed by 2,4-dichloro-


phenoxyacetic acid (2,4-D) diluted in wax, reduced side and stem-end decay by
50–70%, and improved fruit quality during prolonged storage (Kobiler et al.,
2001). The best control was obtained by concentrations of 2,4-D ranging from 75
to 175 Pg/ml; this efficiently controlled side rots caused by A. alternata and the
stem-end rots caused by A. alternata, Phomopsis spp. and Lasiodiplodia spp. The
combination of HWB, prochloraz application and 2,4-D treatment reduced the
incidence of stem-end rot after 4 weeks of storage at 14C and 7 days of shelf life
at 20C from 86 to 10% in ‘Tommy Atkins’ and from 63 to 12% in ‘Keith’
(Kobiler et al., 2001).

Biological control
Bacillus licheniformis, on its own or alternated with copper oxychloride, has been
evaluated as a preharvest spray treatment to control mango fruit dis-eases.
Preharvest applications of B. licheniformis at 3-week intervals from flowering until
harvest controlled moderate levels of anthracnose, and of soft rot caused by
Botryosphaeria, which suggests a potential treatment for com-mercial preharvest
applications (Silimela and Korsten, 2006).

7.5 Other Minor Diseases


Other pathogens may attack mango fruits after harvest through occasional wounds,
and cause severe diseases, such as black mould caused by Aspergillus spp. and
transit rot caused by Rhizopus spp. Disease control begins in the field, and is
followed by postharvest sanitation, and avoidance of latex burn (stain) and
mechanical injury. A hot water treatment consisting of 46C for 60–120 min and
fungicides can be used, depending on the cultivar (Spalding and Reeder, 1986).
HWB at 55C for 20 s shows good control (Prusky et al., 1999).

7.6 Conclusions
Long periods in transit, new marketing approaches for mangoes (e.g. ‘Ready to
Eat’) and stringent international standards and requirements have raised the need
for improved approaches to disease control, in order to preserve fruit quality.
Integrated postharvest treatment has provided a more durable,
Fruit Diseases 225

consistent, sustainable and practical solution than previously available to enable


producers to control postharvest diseases. Ensuring product quality and safety
starts in the orchard, with proper handling conditions and is fol-lowed by optimal
postharvest treatments; however, the combination of post-harvest treatments with
the proper use of the cold-chain concept may provide the ultimate in favourable
conditions for preserving fruit quality.

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8 Foliar, Floral and Soilborne Diseases

R.C. Ploetz1 and S. Freeman2


1
University of Florida, Florida, USA
2
Agricultural Research Organization, Bet Dagan, Israel

8.1 Introduction 232


8.2 Foliar and Floral Diseases 232
Algal leaf spot 232
Alternaria leafspot 233
Anthracnose 235
Apical necrosis 240
Bacterial black spot (black canker) 241
Black-banded disease 245
Black mildew, sooty blotch and sooty mould 246
Blossom blight 248
Decline disorders 249
Galls and scaly bark 256
Grey leafspot 259
Leaf blight 260
Malformation 261
Parasitic plants 270
Phoma blight 270
Phoma leafspot 270
Pink disease 271
Powdery mildew 271
Scab 274
Seca and sudden decline 274
Stigmina leafspot 279
8.3 Soilborne Diseases 281
Black root rot 281
Nematode damage 281
Phytophthora diseases 282
Root rot and damping off 284
Sclerotium rot 285
Verticillium wilt 286
White root disease 286
8.4 Conclusions 289
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 231
232 R.C. Ploetz and S. Freeman

8.1 Introduction
Diseases affect all tissues and developmental phases of mango. Mango dis-eases
have been reviewed in the first edition of this book (Litz, 1997) and by Singh
(1968), Cook (1975), Snowdon (1990), Ploetz et al. (1994) and Ploetz (2003). Lim
and Khoo (1985), Prakash and Srivastava (1987) and Ridgeway
(1989) have also reviewed this subject. This chapter focuses on foliar, floral and
soilborne diseases of mango. Each is discussed with respect to signifi-cance,
geographical distribution, history, symptoms, causal agent(s), epide-miology and
management. These diseases are caused mainly by eukaryotes (Domain Eukaryota)
among which the true fungi, Eumycota (Ascomycota and Basidiomycota),
predominate. Other less important eukaryotes include the fungus-like oomycetes
(Oomycota), nematodes (Metazoa), and parasitic plants and green algae (Plantae).
With one exception, relatively minor pathogens of mango are prokaryotes in the
Domain Eubacteria; all are Gram-negative J-proteobacteria. None of these diseases
is caused by other plant-pathogenic eukaryotes (protozoa), prokaryotes (D- and E-
proteobacteria, Mollicutes and Firmicutes), or the nucleic acid-based pathogens,
the viruses and viroids.

8.2 Foliar and Floral Diseases


Diseases above ground are the most important and conspicuous problems on
mango. Since many of the pathogens that incite foliar diseases also affect panicles,
diseases on each are combined in this section, as are a few diseases that affect the
branches and trunks of mature trees.

Algal leaf spot

Algal leaf spot, also known as red rust, is caused by a parasitic green alga,
Cephaleuros virescens Kunze (synonyms: Cephaleuros parasiticus Karst, Ce-
phaleuros mycoidea Karst) (family Trentepohliaceae, division Chlorophyta) (Lim
and Khoo, 1985). It is a common problem on mango and many other tropical and
subtropical plants (Joubert and Rijkenberg, 1971). Although algal leaf spot can
cause serious problems on tea Camellia sinensis (L.) O. Kuntz, black pepper, Piper
nigrum L., and other important crops, it is usually debilitating on mango only in
poorly managed orchards (Lim and Khoo, 1985). In the latter cases, abiotic or
biotic stresses, such as mites, insects and other foliar diseases, can increase the
severity of this disease.
Conspicuous symptoms of algal leafspot are orange to rust-coloured,
velutinous patches on both leaf surfaces (Plate 43) (Lim and Khoo, 1985). They are
initially 5–8 mm in diameter, but can merge to involve large, irregu-lar sections of
the leaf. They later assume a dull, greyish green colour, and eventually become
bleached patches. The alga can also affect twigs and branches, causing the bark to
crack as the pathogen’s filaments extend into the host cortical tissues. The orange
tufts produced by C. virescens are the algal thallus located beneath the host cuticle.
They produce erect cells, some
Foliar, Floral and Soilborne Diseases 233

Fig. 8.1. Branch of the thallus of Cephaleuros virescens that has terminated in
several oval sporangia (Photograph courtesy of T.-K. Lim).

of which enlarge to produce stalked, terminal or ovoid, 30 × 24 Pm, sporangia


(Fig. 8.1). Sporangia produce biflagellate zoospores that are dispersed by
rainsplash and wind and are the primary infective propagules. Flask-shaped
gametangia in the thallus are responsible for sexual reproduction. In free water,
they release 8–32 biflagellate gametes, pairs of which fuse and give rise to dwarf
sporophytes. The sporophytes produce microsporangia that bear quadriflagellate
zoospores; their role is not known.
Cephaleuros virescens requires a humid environment to establish and spread
(Lim and Khoo, 1985). Pruning the canopy, mowing beneath trees and wider row
spacings to increase air circulation and sunlight penetration all help combat the
disease. Proper fertilization and irrigation, and control of insect pests, mites and
other foliar diseases increase the tree’s ability to cope with algal leafspot.
Algacides or fungicides (i.e. fentin acetate or those con-taining copper (Cu)) can be
used to control the disease in severe cases.

Alternaria leafspot

Alternaria alternata (Fr.) Kreissler (synonyms: Alternaria fasciculata (Cooke and


Ellis) L. Jones and Grout, Alternaria tenuis Nees, and Macosporium
234 R.C. Ploetz and S. Freeman

fasciculatum Cooke and Ellis; no teleomorph is known) causes black spot on


mango, alternaria leaf spot and lesions on inflorescences (Prusky et al., 1983;
Cronje et al., 1990). Although the fungus is cosmopolitan and has a large number
of host plants (Neergaard, 1945; Domsch et al., 1980), the diseases it causes on
mango are most prevalent in arid environments. In Israel, it is a more important
disease on fruit than leaves.
Symptoms on leaves are round, dark spots, 1–3 mm in diameter; they are most
prevalent on the underside of leaves (Prusky, 1994). Similar spots that occur on the
rachis of inflorescences can reduce fruit set (Cronje et al., 1990). Conidiophores of
the causal fungus originate alone or in small groups, and are smooth and pale to
mid-olivaceous or golden brown. They are sometimes geniculate, and vary between
simple and branched, and straight to flexuous. Conidia are obclavate, 20–36 × 9–
9.5 Pm, three- to five-septate and borne in long chains (Fig. 8.2).

Fig. 8.2. Conidia and conidiophores of Alternaria alternata (Source: Ellis, 1971).
Foliar, Floral and Soilborne Diseases 235

Certain antifungal compounds are associated with the latency of A. alter-nata


infections on fruit (Droby et al., 1986, 1987). Whether the same com-pounds are
formed in leaves and inflorescences has not been reported. Infected leaves, twigs,
inflorescences and leaf litter are significant sources of inocu-lum for fruit infection.
Conidia of A. alternata are dispersed by air currents (Prusky, 1994). Several
different fungicides and combined treatments are effective against the diseases that
are caused by this pathogen. Postharvest development of alternaria fruit rot during
storage is usually prevented by a combination of hot water brushing in combination
with prochloraz at 225 Pg/ml (Prusky et al., 2002).

The finding that A. alternata alkalinizes the host pH (Eshel et al., 2002)
prompted investigations into modulating environmental pH with acid treat-ments to
reduce fungal colonization. Spore germination and germ-tube elon-gation of A.
alternata in vitro were inhibited by, respectively, 95 and 65% by exposure to 1.25
mM hydrochloric acid (HCl). Germination was completely inhibited by 2.5 mM
HCl (Prusky et al., 2006). Acidified solutions containing 45 Pg/ml prochloraz
inhibited alternaria rot development better than treat-ment with HCl alone (Eshel et
al., 2002). These results have not been extended to control foliar and floral diseases
that are caused by this pathogen.

Anthracnose

Anthracnose is the most important disease of mango (Cook, 1975; Lim and Khoo,
1985; Ploetz, 2003), particularly on fruit in humid, high rainfall envi-ronments. It is
usually replaced by, and is less important than, other diseases in dry production
areas. Anthracnose can also be a serious problem on foli-age and flowers. Crowded
and moist conditions in nurseries can result in significant damage to young leaves
and, in extreme cases, new orchards have been devastated (Bose et al., 1973).
Blossom blight, which has been attributed to one of the anthracnose agents but is
also caused by other fungi, is covered separately below.

Symptoms
On panicles, necrotic flowers abscise leaving persistent peduncles. Small, cir-cular
dark spots also develop on pedicles and peduncles. Lesions may enlarge and
coalesce to form large patches of necrotic, dark brown tissue. With suf-ficient
rainfall, salmon- to orange-coloured fructifications of the pathogen develop on the
affected tissues. On leaves, lesions are dark brown and sur-rounded by chlorotic
haloes, have irregular, rounded margins, and are not delimited by veins (Fig. 8.3).
Lesions are 0.5–1.0 cm in diameter on mature leaves, but can expand on young
leaves. Eventually, large, necrotic patches can develop that deteriorate and fall
from the leaf giving it a tattered appear-ance. Although different mango cultivars
are known to vary considerably in their resistance to anthracnose on fruit, reports
on the foliar and floral resis-tance of different cultivars and whether resistances of
the various organs are related have not been published.
236 R.C. Ploetz and S. Freeman

Fig. 8.3. Foliar symptoms of anthracnose (Photograph courtesy of S. Freeman).

Aetiology
In most production regions, anthracnose is caused by the ascomycete fungus,
Colletotrichum gloeosporioides Penz. (teleomorph: Glomerella cingulata
(Stonem.) Spauld. and Schrenk.) (Cook, 1975; Snowdon, 1990; Ploetz, 2003). In
Austra-lia, Florida USA, India, Japan and Taiwan, Colletotrichum acutatum
Simmonds (teleomorph: Glomerella acutata) plays a minor role (Fitzell, 1979;
Prakash, 1990; Weng and Chuang, 1995; Taba et al., 2004; Tarnowski,
unpublished). In Colombia, Colletotrichum boninense J. Moriwaki, Toy. Sato and
T. Tsukiboshi has been reported (Afanador-Kafuri et al., 2003).
Colletotrichum spp. cause diseases on several subtropical and tropical hosts
(Jeffries et al., 1990; Freeman et al., 1998). Cultures of the fungus on potato
dextrose agar (PDA) are greyish white to dark grey and usually pro-duce an aerial
mycelium ranging from a thick mat to sparse tufts (Holliday, 1980). Conidia are
hyaline, unicellular and either cylindrical with obtuse ends or ellipsoidal with a
rounded apex and a narrow, truncate base. They are 7–20 × 2.5–5 Pm, and are
formed on hyaline to faintly brown conidiophores in
Foliar, Floral and Soilborne Diseases 237

(d)
50μ

(a)

(e)

(b)

10μ

(c)
(g) (f )

Fig. 8.4. (a) Acervulus and emergent setae, (b) conidiophores, (c) conidia, (d) perithecium,
(e) asci, (f) ascospores and (g) appressoria at hyphal termini of Glomerella cingulata
(anamorph: Colletotrichum gloeosporioides) (Source: from Commonwealth Mycological
Institute (CMI) description no. 315).

acervuli that are irregular in shape and about 500 Pm in diameter. Setae are 4–8 ×
200 Pm, one- to four-septate, brown and slightly swollen at the base and tapered at
the apex. Hyphopodia have been used to distinguish isolates of C. gloeosporioides
and C. acutatum (Du et al., 2005), but provided ambiguous results in Florida
(Palmateer et al., 2007).
Characterization and taxonomic identification of Colletotrichum spp. has
relied on morphology and host range (Freeman et al., 1998; Du et al., 2005). In
general, C. gloeosporioides (Fig. 8.4) produces longer and narrower conidia than
C. acutatum (Fig. 8.5), as well as circular vs lobed hyphopodia. However,
variability in culture and host range has made these criteria unreliable for species
identification (Adaskaveg and Hartin, 1997; Freeman et al., 1998). Col-letotrichum
gloeosporioides and C. acutatum are species complexes with a large number of
hosts that are so broadly defined that the names are of ‘limited use to the
taxonomist or plant health practitioner’ (Du et al., 2005). Several molec-ular tools
have been implemented to differentiate within and among Col-letotrichum spp.,
including: species-specific polymerase chain reaction (PCR) primers, random
amplified polymorphic DNA (RAPD) and arbitrarily primed PCR (Freeman and
Rodriguez, 1995; Afanador-Kafuri et al., 2003); A+T-rich DNA analyses (Freeman
et al., 1993); sequence analyses of the inter-nal transcribed spacer (ITS) regions of
ribosomal DNA (rDNA) (Sreenivasap-rasad et al., 1996; Freeman et al., 2001;
Afanador-Kafuri et al., 2003) and MAT1-2 mating type sequences (Du et al.,
2005).
238 R.C. Ploetz and S. Freeman

(c)
10 μ

(d)

(a) (b)
25 μ 10 μ

Fig. 8.5. (a) Acervulus, (b) conidiogenous cells and setae, (c) conidia and (d)
appres-soria of Colletotrichum acutatum, anamorph of Glomerella acutata (Source:
from CMI description no. 630).

Relatively few studies have examined isolates of C. gloeosporioides from


mango. Gantotti and Davis (1993) reported that isolates from different organs
produced different pectin-degrading enzymes, whereas Alahakoon et al. (1994b)
indicated that > 80% of isolates from mango leaves, inflorescences and fruit in Sri
Lanka were identical, based on restriction fragment length polymorphisms
(RFLPs). Rivera-Vargas et al. (2006) reported that isolates of C. gloeosporioides
from leaves, fruit, flowers and panicles caused similar dis-ease on detached leaves.
Work is needed to clarify whether, and the extent to which, isolates from different
mango organs differ, and whether there is a relationship among disease reactions on
various organs.
Analyses of RFLPs and RAPDs indicated that isolates from mango are
genetically uniform and differ from those recovered from avocado, caram-bola and
papaya (Hodson et al., 1993; Alahakoon et al., 1994a; Hayden et al., 1994).
Isolates genetically similar to those from the latter crops occur infre-quently on
mango, and mango isolates were not found on the other plants and were usually
virulent only on mango. Alahakoon et al. (1994b) suggested
Foliar, Floral and Soilborne Diseases 239

that the mango isolates comprise a C. gloeosporioides population that was dis-
seminated worldwide from a single source, perhaps as an endophyte.
A recent study identified Colletotrichum spp. that infect mango, passion-fruit
(Passiflora spp.) and tamarillo (Solanum betacea cav. Sendt.) in Colombia and
assessed whether cross-infection between the host species occurred (Afanador-
Kafuri et al., 2003). With species-specific PCR primers, most of the mango isolates
were identified as C. gloeosporioides. However, DNA of the passionfruit isolates
and single isolates from tamarillo and mango were not amplified by either C.
acutatum- or C. gloeosporioides-specific primers; they were identified later as C.
boninense (Freeman, unpublished data). Further molecular analyses determined
that the isolates of C. gloeosporioides from mango were heterogeneous, but that the
population of C. boninense from pas-sionfruit, tamarillo and mango was uniform; it
may not be host specific.
The origins and diversity of C. gloeosporioides on mango require more study.
Furthermore, whether distinct populations of this diverse species occur on different
mango cultivars and organs, and whether disease reac-tions on one mango organ
could be used to predict those on another should be determined. These results
would be relevant to host resistance and improvement of the crop, as well as the
chemical and cultural control of this important disease.

Epidemiology and management


Fitzell and Peak (1984) determined that conidia were the most important inoculum
in Australia. They are produced on branch terminals, mummified inflorescences,
flower bracts and leaves. New leaf flushes are the most signifi-cant source of
inoculum, and this was corroborated by Dodd et al. (1991) in the Philippines.
Optimum production of conidia occurred between 25 and 30C when free moisture
was available, but also formed at 95–97% relative humid-ity (RH). The pathogen’s
teleomorph played no apparent role in the spread of the disease (Fitzell and Peak,
1984). The threat posed by C. gloeosporioides to fruit production is greatest in
areas with heavy rainfall. In general, this begins with the onset of flowering (Lim
and Khoo, 1985; Jeffries et al., 1990). During commercial production these
diseases are managed with diverse chemicals. Registration of the various pesticides
varies in different production areas (Jeffries et al., 1990).

Many of the most effective fungicides are systemic. They have selective
modes of action, but are prone to lost or reduced efficacy due to the develop-ment
of resistance in the targeted pathogens. Resistance in C. gloeosporioides to
benomyl is now common (Maymon et al., 2006), and there is inherent toler-ance in
C. acutatum to this fungicide (Freeman et al., 1998; Peres et al., 2002). Newer
strobilurin fungicides that are also susceptible to resistance problems must be used
properly (no more than three applications per season and alter-nated with
fungicides with different modes of action) to ensure that their efficacy is not lost.

Although fungicide applications usually focus on reducing damage to fruit,


disease control on foliage is indicated in some, and on inflorescences in most
situations. Trees in nurseries usually require protection if they are
240 R.C. Ploetz and S. Freeman

crowded or receive overhead irrigation. Since infected foliage and branch terminals
represent important reservoirs of inoculum for blossom and fruit infection, fruit set
and anthracnose control on fruit are enhanced if disease control is exercised prior
to flowering (Jeffries et al., 1990). Off-season control measures are beneficial in
production environments that receive significant rainfall.

Apical necrosis

Apical necrosis was first reported in Spain in 1991, and now occurs in Israel, Italy,
Portugal and possibly Egypt (Cazorla et al., 1998, 2006; F.M. Cazorla, Málaga,
2007, personal communication). The disease can be quite damaging, and limits
production when panicles are affected. Apical buds, leaves and panicles are
susceptible (Fig. 8.6a–c), but fruit are not (Cazorla et al., 1998). Vegetative and
floral apices are affected by a dark-brown to blackish necrosis (Fig. 8.6a, c).
Necrosis on leaves begins as blackened, water-soaked lesions, 1–3 mm in diameter,
that can coalesce and expand to cover large areas (Fig. 8.6c). Necrosis also extends
from affected buds to petioles, through the leaf midrib, and can cover large areas
(Fig. 8.6a). A milky bacterial exudate often develops on affected apical buds, but
infrequently on petioles (Fig. 8.6b). Large portions of the canopy and high numbers
of flowers can be killed.

(a) (b) (c)

Fig. 8.6. Symptoms caused by the apical necrosis pathogen, Pseudomonas syringae
pv. syringae. (a) Extensive necrosis on young stem, apical bud, petioles and leaves;
(b) bacterial exudate on necrotic stem; and (c) death of a developing floral panicle
and associated leaf necrosis (Photographs courtesy of F.M. Cazorla).
Foliar, Floral and Soilborne Diseases 241

The causal bacterium, Pseudomonas syringae pv. syringae van Hall, affects
many perennial fruit crops (Hirano and Upper, 1983; Kennelly et al., 2007). It is an
epiphyte and is generally not an aggressive pathogen. Disease usually develops
after high populations of the bacterium develop in host tissues as a result of host
predisposition. Strains from mango produce an antimetabolite toxin, mangotoxin,
which plays a role in pathogen virulence and symptom development (Arrebola et
al., 2007).
Cold, wet weather and host genotype are primary factors in the develop-ment
of apical necrosis (Cazorla et al., 1998, 2006). ‘Tommy Atkins’, ‘Lippens’ and
‘Manzanillo’ are very susceptible whereas ‘Keitt’ and ‘Sensation’ are less so.
Apical necrosis is managed in commercial orchards with Cu-containing pesticides,
although there has been a recent increase in control failures and Cu resistance in
Spain and Portugal (Cazorla et al., 2002, 2006). These out-breaks have been
associated with several different Cu-resistance plasmids in the causal bacterium.
Carzorla et al. (2006) determined that the plant resis-tance activator acibenzolar-S-
methyl and the phosphonate derivative fosetyl-Al provided control comparable to
Bordeaux mixture, and that the latter treat-ment might protect plants due to the
protective film it provides against wound entry for the pathogen.

Bacterial black spot (black canker)

Bacterial black spot is a destructive leaf, stem and fruit disease in many pro-
duction areas (Gagnevin and Pruvost, 2001). In India, the disease is called bacterial
canker or black canker due to the cankers it causes on the stems of some cultivars
(Prakash et al., 1994). It can be the most important disease where fungal-induced
diseases are well controlled (Gagnevin and Pruvost, 2001). Bacterial black spot has
been identified in Australia, Myanmar, the Comoros, India, Japan, Kenya,
Malaysia, Mauritius, New Caledonia, Paki-stan, the Philippines, Réunion,
Rodrigues, South Africa, Taiwan, Thailand and the United Arab Emirates (UAE)
(Fukuda et al., 1990; Pruvost et al., 1992; Prakash et al., 1994; Gagnevin and
Pruvost, 1995, 2001; Kishun, 1995; Ah-You et al., 2007b). Given the ease with
which the pathogen is disseminated in propagation materials and its wide,
confirmed distribution, bacterial black spot may be more widely spread than is
currently recognized.

Symptoms
Mango leaves, stems and fruit are affected (Manicom and Pruvost, 1994; Gagnevin
and Pruvost, 2001). On leaves, water-soaked spots are initially 1–3
mm in diameter. As they enlarge, they become raised, black and angular, are
limited by veins and surrounded by chlorotic haloes (Fig. 8.7). These lesions are
larger and more conspicuously raised than those caused by other xan-thomonads
that have been recovered from other species in the Anacardiaceae (Ah-You et al.,
2007a). Lesions can merge to form large necrotic patches, and bacteria may ooze
from lesions during wet conditions. Old lesions dry out, turn white or grey, and
crack. Defoliation occurs in severe cases. Anthracnose
242 R.C. Ploetz and S. Freeman

(a)

(b)

Fig. 8.7. (a) Symptoms of bacterial black spot on the undersurface of a mango leaf,
caused by Xanthomonas axonopodis pv. mangiferaeindicae (Photograph courtesy of
O. Pruvost). (b) Symptoms of bacterial spot on the undersurface of a mango leaf,
caused by a yellow-pigmented xanthomonad (Photograph courtesy of R.C. Ploetz).
Bacterial black spot lesions are larger and more raised than those of bacterial spot.

lesions are not raised or as black and angular as those caused by bacterial black
spot. On branches, bacterial black spot lesions are dark and cracked along the long
axis (Fig. 8.8). They develop only on highly susceptible culti-vars and are often
associated with wounds. Conspicuous, star-shaped lesions are produced on fruit.

Aetiology
Diverse xanthomonads have been recovered from mango and other hosts in the
Anacardiaceae (Gagnevin and Pruvost, 2001; Ah-You et al., 2007a). Only some of
these cause symptoms of bacterial black spot. Early reports that this disease is
caused by Pseudomonas mangiferae-indicae (Patel et al., 1948) and
Foliar, Floral and Soilborne Diseases 243

Fig. 8.8. A twig canker caused by X. axonopodis pv. mangiferaeindicae that


is associated with a wound (Photograph courtesy of O. Pruvost).

Erwinia mangiferae (Steyn et al., 1974) are erroneous. The pathogen’s place-ment
in Pseudomonas was probably due to its production of non-pigmented colonies in
culture (P. syringae pv. syringae causes a different disease of mango, apical
necrosis – see above). Cook (1975) indicated that E. mangiferae is a saprophyte
that reached high populations in old lesions.
Pathological, cultural, biochemical, physiological, serological and genetic data
indicate that strains of the pathogen from different production areas are diverse
(Sanders et al., 1994; Gagnevin and Pruvost, 1995, 2001; Kishun, 1995; Pruvost et
al., 2005). Genetic diversity is greatest among strains from South-east Asia,
suggesting that this region of host diversity is also a centre of pathogen
diversification (Gagnevin and Pruvost, 1995).
The pathogen has a single flagellum, is Gram-negative, rod shaped and 0.4–0.5
× 1.0–1.5 Pm (Manicom and Wallis, 1984). It is oxidase negative and does not
reduce nitrates to nitrites. It cannot use asparagine as a sole carbon and nitrogen
source, but is able to hydrolyse starch, esculin, gelatin and casein. On artificial
media, colonies are cream coloured. (The latter trait is atypical for xanthomonads,
which are usually yellow in culture.) Yellow-pigmented xanthomonads have been
recovered from mango in Brazil, Réunion, South Africa and Florida USA. These
strains cause non-raised leaf lesions (Fig. 8.7b), do not cause fruit or stem lesions,
and should not be
244 R.C. Ploetz and S. Freeman

classified as pv. mangiferaeindicae (Ah-You et al., 2007a). Three genetically and


pathologically distinct groups were identified from different geographic regions
and hosts in the Anacardiaceae by Ah-You et al. (2007a). Group I strains were
from the Old World, multiplied in mango and cashew (Anacar-dium occidentale
L.), fell in amplified fragment length polymorphism (AFLP) group 9.5 of
Xanthomonas axonopodis, and contained strains that produced typical bacterial
black spot symptoms on mango. These strains of Xanthomo-nas campestris pv.
mangiferaeindicae sensu lato were redescribed as X. axonopo-dis pv.
mangiferaeindicae sensu novo (s.n.) (Ah-You et al., 2007a). In contrast, Group II
strains were from Brazil, multiplied in cashew, but not mango, and fell in AFLP
group 9.6 of X. axonopodis. They are associated with symptoms on mango that
differ from those of bacterial black spot, including brown, flat leaf lesions, and
black, depressed lesions on fruit of only a few cultivars that is sometimes
associated with pulp rot. These strains were responsible for previous reports of
bacterial black spot in Brazil (Gagnevin and Pruvost, 2001; Ah-You et al., 2007a).
Group III strains are responsible for a unique syn-drome on Spondias dulcis and
Spondias mombin in the French West Indies; they fell in AFLP group 9.4. Groups
II and III were described, respectively, as X. axonopodis pv. anacardii and X.
axonopodis pv. spondiae (Ah-You et al., 2007a).

Epidemiology and management


The pathogen is disseminated by wind-driven rain (Gagnevin and Pruvost, 2001).
Long-distance dissemination occurs via infected propagation material and, less
frequently, in infected seed. True seed are not infected, but surface contamination
is known. Insects may play a role in dissemination, but these interactions are little
studied. The pathogen is an epiphytic colonist of leaves (Manicom, 1986; Pruvost
et al., 1990), buds (Pruvost et al., 1993) and fruit (Pruvost and Luisetti, 1991b).
Infection occurs through wounds and, less often, stomata on old leaves (young
leaves are thought to be resistant due to their non-functional stomata) (Gagnevin
and Pruvost, 2001). High RH (> 90%) and moderate temperatures (25–30C)
favour disease development (Kishun and Sohi, 1983; Pruvost and Luisetti, 1991b).
There is a direct rela-tionship between the level of disease that develops on leaves
and fruit (Man-icom, 1986; Pruvost et al., 1990). Pruvost and Luisetti (1991b)
considered that leaf susceptibility was an important criterion when cultivars were
selected for lower fruit susceptibility.

Resistance to bacterial black spot varies among mango cultivars, and resistant
cultivars should be used where disease pressure is high (Manicom and Pruvost,
1994). When new orchards are established, pathogen-free plant-ing material should
be utilized. Since the pathogen moves short distances in wind-blown aerosols
(usually within orchards), the long-distance spread of the pathogen depends almost
entirely upon the movement of infected plants (Manicom and Pruvost, 1994).
Windbreaks can be used to reduce wounding and infected twigs (Fig. 8.8) should
be pruned to reduce inoculum in the canopy.

Bacterial black spot can be quite difficult to control, particularly on sus-


ceptible cultivars. Available chemicals may be only marginally effective
Foliar, Floral and Soilborne Diseases 245

under high disease pressure (Pruvost et al., 1989). During rainy weather,
applications of Cu-based bactericides are recommended. The application schedules
for these compounds focus on protecting fruit, and vary according to the length of
time fruit are exposed to wet conditions (Manicom and Pru-vost, 1994). Although
agricultural antibiotics (e.g. various formulations of streptomycin sulfate or nitrate)
have been reported to be effective (Misra and Prakash, 1992; Viljoen and Kotzé,
1972), resistance that develops to these products after continuous use limits their
long-term effectiveness against this disease. Biological control measures have not
been widely researched. Pruvost and Luisetti (1991a) reported little success. In
India, Kishun (1994) indicated that a strain of Bacillus coagulans from the
phylloplane of mango was effective against strains of the pathogen, although
control of bacterial black spot in the field was not reported.

Black-banded disease

This is a relatively unimportant disease that affects mango leaves and branches
(Reddy et al., 1961). The causal fungus, Rhinocladium corticola Mas-see
(described as ‘corticolum’) (teleomorph: Peziotrichum corticolum (Massee)
Subrumanian), was described on the bark of trees in Poona, India (Hughes, 1980).
It produces repent, intricately branched, septate, olivaceous hyphae 5–7 Pm in
diameter. Erect hyphae bear globose, olivaceous, densely and minutely tuberculate,
15–18 Pm conidia. Hughes (1980) questioned whether this was a species of
Rhinocladium since it was quite different from other spe-cies in the genus. It forms
a black, velvety mass of hyphae on affected sur-faces in conspicuous blotches or
bands. The fungus is restricted to the outer portions of bark. Bordeaux mixture
controls the disease, but is not required in most situations.

Black mildew, sooty blotch and sooty mould

Several ascomycetes produce dark-coloured, usually superficial growths on the


surfaces of stems, leaves and fruit (Lim and Khoo, 1985). These range from thin,
diffuse webs of dark hyphae to opaque, felty layers; in extreme cases, a thick crust
of hyphae forms. The variations in appearance result, presumably, from the
different species of fungi that are involved. These are usually not important
problems in well-maintained orchards. However, layers of hyphae that the black
mildew and sooty mould fungi form may be thick enough to block sunlight and
inhibit photosynthesis. These blemishes also detract from the appearance and
marketability of fruit.

Sooty moulds develop in the presence of aphids, mealybugs, scales and other
sucking insects that produce honeydew (excreta) when feeding. Hon-eydew is a
required food source for these fungi, and the problems they cause dissipate if the
associated insects are controlled. In contrast, black mildews
246 R.C. Ploetz and S. Freeman

and sooty blotch are not dependent on honeydew and grow directly on host
surfaces.

Aetiology
Black mildew and sooty mould are similar in appearance, but their respec-tive
causal agents are distinct (Lim and Khoo, 1985). The black or dark mil-dews are a
group of mostly tropical obligate plant pathogens that produce two types of
hyphopodia (Alexopoulos et al., 1996). Capitate hyphopodia are lobed appressoria
from which infection haustoria are formed, whereas mucronate hyphopodia
function as conidiogenous cells. Black mildew of mango is caused by Meliola
mangiferae Earle (Sordariomycetes, Ascomycota) (Fig. 8.9).

In contrast, the fungi that cause sooty moulds are diverse saprophytes that
require honeydew to colonize plant surfaces. In Malaysia, Lim and Khoo (1985)
listed coelomycetes (Polychaeton), hyphomycetes (Tripospermum) and
loculoascomycetes (Antennulariella, Chaetothyrium, Limacinula and Scorias),
whereas the reported agents in India were hyphomycetes (Leptoxyphium,

(b)
10 mm
(a)

1 mm

10 μm

(c) 250 μm

(d)

Fig. 8.9. (a–c) Signs of black mildew, and (d) microscopic features of the
causal agent, Meliola mangiferae (Source: from CMI description no. 1355).
Foliar, Floral and Soilborne Diseases 247

Microxyphium and Tripospermum) and loculoascomycetes (Capnodium) (Butler


and Bisby, 1931; Prakash, 1988). In Pakistan, 18 species in eight genera were
associated with sooty mould, including the foliar pathogens Aspergillus,
Alternaria, Botryodiplodia, Capnodium, Cladosporium, Curvularia, Fusarium and
Helminthosporium spp. (Hamid and Jalaluddin, 2006). Since some of the sooty
mould fungi may not sporulate on plants and because they are often found in
combination with one another, it is usually difficult to identify the specific species
that are involved.
Although ‘sooty blotch’ has been used as a synonym for ‘sooty mould’ (Singh,
1968), sooty blotch refers specifically to disease complexes on tem-perate and
tropical plants that are not associated with honeydew and are caused by a diverse
group of Dothidiomycetes (Ascomycota) (Johnson et al., 1997; Ploetz et al., 2000;
Batzer et al., 2005). The specific agents that are associ-ated with sooty blotch on
mango are not known, but resemble those from apple, carambola and pear (Plate
44).

Epidemiology and management


Sooty moulds develop on honeydew that is produced on plant surfaces by aphids,
mealybugs, scales and other sucking insects. They are managed by controlling the
associated insects with oils and insecticides (Lim and Khoo, 1985). In Pakistan,
spraying of fungicides (sulfur (S) and mancozeb) and insecticides (malathion,
diazinon and coal tar at 1 kg/tree) separately reduced the incidence of sooty mould
on foliage, whereas a mixture of fungicides and insecticides further decreased
sooty mould incidence (Hamid and Jalalud-din, 2006). In India, sooty mould,
caused by species of Microxyphium, Lep-toxyphium and Tripospermum, was best
controlled with a spray of S and parathion-methyl (Prakash, 1991).

Sooty blotch management has not been investigated on mango; however, signs
of these fungi have been removed from apple with various postharvest washes
(Batzer et al., 2002), and managed in apple orchards with diverse contact and
systemic fungicides (Williamson and Sutton, 2000).

Blossom blight

Blossom blight can reduce fruit set and production considerably since flow-ers and
large areas of the panicle can be killed. When this disease was con-trolled with
fungicides in the Philippines, a 55–80% increase in fruit set occurred (Pordesimo,
1982).

Symptoms
Blossom blight starts as a wilt of the affected part of the inflorescence that is often
curved, the ‘shepherd’s crook symptom’ (Fig. 8.10). The peduncle blackens and
dies back from the tip. Internally, discoloration advances beyond the surface lesion.
Large black lesions can develop lower on the peduncle, and once it is girdled the
apex dies.
248 R.C. Ploetz and S. Freeman

Fig. 8.10. Symptoms of blossom blight on panicles of mango. Note the


blighted, curved terminals and almost complete lack of fruit set (Photograph
courtesy of D. Benscher).

Aetiology
The cause of blossom blight is confused. Colletotrichum gloeosporioides has been
reported most frequently as the responsible fungus (Fitzell et al., 1984; Jeffries et
al., 1990), and A. alternata has also been reported to attack panicles and reduce
fruit set (Cronje et al., 1990). Powdery mildew (see section below) also damages
panicles, but its symptoms are distinct from those of blossom blight. In South
Africa, symptoms caused by A. alternata and C. gloeosporioides are small, mainly
superficial black spots, 1–2 × 2–5 mm, on the peduncle (Darvas, 1993; Lonsdale
and Kotzé, 1993). Rather than blossom blight, Lonsdale and Kotzé (1993)
indicated that these pathogens caused blossom spot. In con-trast, Lonsdale and
Kotzé (1993) reported that Dothiorella mangiferae caused extensive, systemic
damage, and Darvas (1993) indicated that Dothiorella domini-cana is the only
fungus that caused typical symptoms of blossom blight. Crous et al. (2006) placed
these fungi in a new genus, Neofusicoccum (see Decline disorders section below).
Work is needed to determine the distribu-tion of Neofusicoccum-incited panicle
disease, and the identity of the most important blossom blight pathogens
worldwide.

Epidemiology and management


Little is known about the epidemiology of Neofusicoccum-incited panicle dis-ease.
Studies on the stem-end rot diseases have shown that the causal fungi are
endophytes. The roles of internal and external sources of inoculum in the
development of panicle disease are unknown. For optimal fruit set and
Foliar, Floral and Soilborne Diseases 249

development, blossom blight must be controlled. Once flowering begins, early and
frequent fungicide applications are necessary in many areas, depending on rainfall.
Previously published infection models can be used to time applications
appropriately (Fitzell et al., 1984; Dodd et al., 1991). Fitzell et al. (1984)
investigated environmental conditions that were conducive to infection by C.
gloeosporioides, and indicated that temperature and free mois-ture were important
determinants of infection. They developed a model for scheduling fungicide
application, which reduced the number of applications that were needed to control
blossom blight. Presumably, systemic fungicides would be needed to control
disease caused by endophytic agents.

Decline disorders

Several diseases of mango have been variously termed blight, canker, decline,
gummosis, twig blight, tip dieback and stem bleeding. They have similar symptoms
and aetiologies.

Symptoms
These widespread problems are not well understood. Symptoms include: (i)
marginal scorching of leaf lamina; (ii) foliar symptoms of nutritional defi-ciencies,
particularly of iron (Fe) and manganese (Mn); (iii) vascular discolor-ation (Fig.
8.11a); (iv) dieback of small branches basipetally from the terminal that may or
may not progress to defoliation (Fig. 8.11b and c); (v) gummosis, an oozing of a
clear or cloudy exudate either from terminal buds or from branches, scaffold limbs
or trunks (Plate 45); and (vi) root degeneration (Lim and Khoo, 1985; Ploetz et al.,
1996a; Ploetz and Prakash, 1997).

Aetiology
Diverse biotic and abiotic factors may be primary causes of decline symp-toms or
predisposing agents (McSorley et al., 1980; Kadman and Gazit, 1984; Schaffer et
al., 1988; Ploetz and Prakash, 1997). Fungi are the most common agents. They are
endophytes that also cause stem-end rots on mango fruit, and are usually secondary
pathogens that cause disease on weakened, pre-disposed hosts (Johnson et al.,
1992; Ploetz and Prakash, 1997; Slippers et al., 2005; Slippers and Wingfield,
2007). Several species cause all or some of the above symptoms when used to
individually inoculate plants (Ploetz et al., 1996a). Their frequent association with
one another in affected tissues may indicate that these symptoms usually develop,
or develop more severely, after multiple infections.

Several of these pathogens are in the Botryosphaeriaceae (Dothidiomy-cetes,


Ascomycota). The taxonomy and nomenclature of these fungi has been confused,
and ‘phylogenetic understanding of major groups within Botry-osphaeria remains
poor’ (Crous et al., 2006). With 28S rDNA sequence data, Crous et al. (2006)
examined natural relationships among available members of the family. Ten
lineages were distinguished, most of which contained ana-morphs with distinct
morphological features. New relationships were revealed
250 R.C. Ploetz and S. Freeman

) ) )
(a) (b) (c)

Fig. 8.11. Among the symptoms that are associated with mango decline are:
(a) internal/vascular discoloration and branch terminal death (tip dieback) that may not
(b), or may be associated with defoliation (c) (Photographs courtesy of D. Benscher).

in some of the lineages that necessitated the renaming of several taxa (Crous et al.,
2006). These new names and holomorphs are used below when discuss-ing the taxa
that occur on mango.
Lasiodiplodia theobromae (Pat.) Griffon and Maubl.) (synonyms: Botryodi-
plodia theobromae Pat., Diplodia natalensis Pole-Evans, and Diplodia theobromae
(Pat.) W. Nowell) is the most common and widespread cause of decline dis-eases
of mango (Ploetz and Prakash, 1997), and affects many other host plants in the
tropics (Punithalingam, 1976). Crous and Palm (1999) declared B. theo-bromae, a
nomen dubium. Denman et al. (2000) reduced D. natalensis and L. theobromae to
synonymy with D. theobromae. However, adopting this change was questioned by
Burgess et al. (2006), who noted that five species of Lasio-diplodia fell in a
phylogenetic clade that had 100% bootstrap support; it was distinct from a clade
that included species of Diplodia and Dothiorella. The teleomorph of L.
theobromae, formerly Botryosphaeria rhodina (Cooke) Arx (synonym:
Physalospora rhodina Cooke), is usually not found in nature. In the study of Crous
et al. (2006), the genus Botryosphaeria was reserved for the type species
Botryosphaeria dothidea (Moug.:Fr.) (anamorph: Fusicoccum aesculi Corda),
which was in a different clade than L. theobromae. However, Crous et al. (2006)
refrained from renaming B. rhodina until the poorly resolved clade in which it
resided could be clarified with work with additional isolates and analyses.

Lasiodiplodia theobromae attacks weakened trees that are predisposed by:


high and low temperatures; drought; high RH; hardpan soils; sun scorch; and tar
and tanglefoot (Muller, 1940; Das Gupta and Zacchariah, 1945; Alvarez-García
and López-Gracía, 1971; Acuña and Waite, 1977; Ploetz et al., 1996a). It is often
an endophyte, infects wounded plants, and is found in soil, on dead twigs,
mummified fruit and on organic debris beneath trees (Johnson et al., 1992).
Foliar, Floral and Soilborne Diseases 251

Dieback caused by L. theobromae has been recognized as a significant dis-


ease in India since the 1940s. It was the most serious disease of mango in the
Jaipur district (Verma and Singh, 1970), and affected 30–40% of the planta-tions in
the Moradabad region of Uttar Pradesh (Prakash and Srivastava, 1987). Das Gupta
and Zacchariah (1945) indicated that only L. theobromae caused dieback; Phoma
sp. and two Fusarium spp. were not pathogenic. Lasio-diplodia theobromae caused
a canker on mango in Indonesia (Muller, 1940) and Malaysia (Lim and Khoo,
1985), dieback in Egypt and the Sonsonate area of El Salvador (Acuña and Waite,
1977), and gummosis and dieback in Puerto Rico (Alvarez-García and López-
Gracía, 1971).
Lasiodiplodia theobromae produces fluffy, grey-black mycelium on oatmeal
agar (OA) and PDA (Johnson, 1994b). Conidiomata may be simple or develop into
aggregated stromatic bodies (Burgess et al., 2006). Cirri of conidia may ooze from
ostioles. Conidia are initially hyaline, aseptate, granular, ovoid to ellipsoid and
thick-walled (Fig. 8.12). Mature conidia are two celled, 17–33 × 10–15 Pm, and
brown walled with numerous longitudinal striations. Paraphyses are usually
present. The teleomorph was produced when indi-vidual isolates were cultured on
caimito fruits (Chrysophyllum cainito L.) or papaya (Carica papaya L.) stems,
suggesting that it was homothallic (Alvarez-García and López-Gracía, 1971).

One of the most important mango pathogens causes stem-end rot on fruit,
dieback and blossom blight. Crous et al. (2006) refer to it as Neofusicoccum
parvum (Pennycook and Samuels) Crous, Slippers and A.J.L. Phillips (formerly
Fusicoccum parvum Pennycook and Samuels) (teleomorph: Botryosphaeria-like;
formerly Botryosphaeria parva Pennycook and Samuels). Slippers et al. (2005)
argued that D. dominicana Petro and Cif. may be synonymous with this fun-gus.
Johnson (1992), an author of the Slippers et al. (2005) paper, had placed D.
dominicana in synonymy with F. aesculi Corda (teleomorph B. dothidea
(Moug.:Fr.) Ces. and De Not.). They indicated that this fungus had been mis-
identified as Botryosphaeria ribis Gross. and Duggar (anamorph: Fusicoccum sp.)
and B. dothidea (anamorph: F. aesculi), due to overlapping host ranges and spore
dimensions. They felt that the tip dieback fungus reported by Ramos et al. (1991)
as B. ribis was probably N. parvum (= ‘B. parva’).
Neofusicoccum parvum produces cottony grey mycelium and discrete pyc-
nidia or stromatic multilocular fruiting bodies on, respectively, PDA and OA
(Johnson et al., 1991). Discrete, immersed pycnidia in a subcuticular pseudostroma
are produced on mango. Conidia are fusiform to navicular, 14.7–25.5 (19) × 4.5–7
(5.2) Pm, hyaline and unicellular (Slippers et al., 2005). Sometimes, brown,
biseptate conidia are observed. The teleomorph develops infrequently on OA, and
has been found on mango twigs in tree litter in Australia (Johnson et al., 1991). On
twigs, pseudothecia are subglobose to pyriform, 210 × 120 Pm, and form beneath
the epidermis. Ascostromata are hemi-lenticular and up to 10 mm wide on OA.
Asci are eight spored, bituni-cate and irregularly biseriate. Ascospores are hyaline,
single celled, fusiform and 16–25 × 4.5–9.5 Pm.

Neofusicoccum mangiferum (Syd. and P. Syd.) Crous, Slippers and A.J.L.


Phillips (basionym: Dothiorella mangiferae Syd. and P. Syd.; synonyms:
252 R.C. Ploetz and S. Freeman

(b)

(d)

(c)
(a)

500 μ
10 μ

Fig. 8.12. Pycnidia (a and b), conidiogenous cells and immature conidia (c)
and mature and immature conidia (d) of Lasiodiplodia theobromae (Source:
from CMI description no. 519).

Nattrassia mangiferae (Syd. and P. Syd.) B. Sutton and Dyko; Fusicoccum


mangiferum (Syd. and P. Syd.) Johnson, Slippers and M.J. Wingf.) causes blos-
som blight and postharvest fruit rot in South Africa (Lonsdale and Kotzé, 1993;
Saaiman, 1996). On PDA, N. mangiferum produces grey, felted myce-lium with
gregarious, partly immersed, discrete conidiomata, ‘pepper-spot’ patterns of
pycnidial initials, and dark grey mycelium that lacks the white tufts found in
similar species (i.e. N. parvum) (Johnson, 1994b; Slippers et al., 2005). On mango,
the fungus produces unilocular conidiomata in subcuticu-lar pseudostroma. The
conidia of N. mangiferum differ from those produced by other Neofusicoccum spp.
by their shorter average length (13–14 Pm) and smaller length/width ratio (2–2.5)
(Slippers et al., 2005). They are usually unicellular, ellipsoid to ovoid, 13.6 × 5.4
Pm and hyaline, although conidia often become one or two septate, light brown
with distinctly darker middle cells. The teleomorph (not identified) resembles ‘B.
parva’, and develops occasionally on OA.
Foliar, Floral and Soilborne Diseases 253

(d)

(e)

(c)

10 μ

(b)
50 μ

(a)

Fig. 8.13. (a) Vertical section of stroma, (b) part of pycnidial wall and conidiophores,
(c) conidiophores, (d) conidia, and (e) the Scytalidium-like synanamorph of
Neoscyta-lidium dimidiatum (Source: from CMI description no. 274).

A dieback disease of mango has been recognized in Niger (Reckhaus and


Adamou, 1987). Neoscytalidium dimidiatum (Penz.) Crous and Slippers
(basionym: Torula dimidiata Penz.; synonyms: Scytalidium dimidiatum (Penz.) B.
Sutton and Dyko (Fig. 8.13); Fusicoccum dimidiatum (Penz.) D.F. Farr;
Hendersonula toruloidea Natrass) causes sudden wilting of shoots to large
branches, firing of leaves and trunk cankers from which a clear exudate orig-inates.
Reckhaus and Adamou (1987) believed that water stress was a pri-mary,
predisposing factor in the development of this disease.
Botryosphaeria dothidea (anamorph: F. aesculi) is an uncommon mango
pathogen (Ploetz and Prakash, 1997; Slippers et al., 2005). It produces a fluffy grey
mycelium with discrete pycnidia on PDA or stromatic multilocular fruiting bodies
on OA. Discrete, immersed pycnidia are produced on mango.
254 R.C. Ploetz and S. Freeman

Conidia are 18.8–30.4 × 4.5–7 Pm, hyaline, and single celled (Fig. 8.14). The
teleomorph is occasionally produced on OA and has been found in litter beneath
avocado and mango trees (Johnson, 1994b; Michailides et al., 1999). On twigs,
pseudothecia are subglobose to pyriform, 210 × 120 Pm, and im-mersed beneath
the epidermis. On OA, ascostromata are hemi-lenticular and up to 10 mm wide.
Asci are eight spored, bitunicate and irregularly biseriate. Ascospores are hyaline,
single celled, fusiform and 16–25 × 4.5–9.5 Pm.
Mango decline is an important disorder in Florida USA, Israel and other areas
that have calcareous soils (Schaffer, 1994; Ploetz et al., 1996a). Symptoms include
interveinal chlorosis and marginal necrosis of leaves,

(b)

(a)
10 μ

(c)
500 μ

Fig. 8.14. (a) Conidia, (b) conidiophores and (c) a vertical section of a conidioma of
Fusicoccum aesculi, anamorph of Botryosphaeria dothidea (Source: Sutton, 1980).
Foliar, Floral and Soilborne Diseases 255

dieback of young twigs that progresses to larger branches, reduced growth of


secondary roots, gummosis and vascular discoloration. Several different factors
have been associated with mango decline in Florida. Schaffer et al. (1988) used the
Diagnosis and Recommendation Integrated System (DRIS) to assess the nutritional
status of declining and healthy ‘Tommy Atkins’ trees. The nutrient imbalance
index, an indication of the overall nutrient balance in trees, was greatest for
declining trees. DRIS identified Mn, Fe or both ele-ments as the most deficient in
declining trees, and in two of the three declin-ing orchards that they investigated,
concentrations of these elements were below the critical range. Mineral
deficiencies may be predisposing factors in the development of mango decline,
since pathogenic fungi are recovered from symptomatic trees (see below).

McSorley et al. (1980) detected the nematode Hemicriconemoides mangiferae


Siddiqi at low, but consistent levels on declining mango trees. They sug-gested that
it may have been responsible for the reduced root growth in affected trees, and
could play a role in the development of the disorder.
Smith and Scudder (1951) reported that Diplodia sp. caused a dieback of
mango. Additional species of fungi were examined by Ploetz et al. (1996a).
Alternaria alternata, C. gloeosporioides, N. parvum (D. dominicana), L.
theobromae and two Phomopsis spp. were recovered from trees with diverse
decline symptoms, and caused one or more of these symptoms on ‘Keitt’ and
‘Tommy Atkins’. Colletotrichum gloeosporioides, N. parvum and L. theobromae
were most damaging, and caused significant bud necrosis, tip dieback, gummosis
and vascular discoloration (Fig. 8.15); these symptoms were distinguishable only
when C. gloeosporioides sporulated on inoculated branches.

In summary, several different fungi cause, or are associated with, decline


symptoms worldwide; most are endophytes that have Botryosphaeria or
Botryosphaeria-like teleomorphs (Botryosphaeriaceae). Stress and wound pre-
disposition are usually associated with symptom development.

Management
Controlling decline disorders of mango is difficult. Techniques that could detect
these pathogens in plants would be useful to identify pathogen-free propagation
materials. The internal location and the diversity of fungi that are involved
decrease the opportunities for controlling these disorders with fungicides (Peterson
et al., 1991). Because significant movement of some of these pathogens may occur
via wind and rainsplash, strategic applications of broad-spectrum protectant
fungicides may be effective at certain times of the year (Lonsdale and Kotzé,
1993), but have not been tested. In India, dieback was managed by pruning affected
portions of the canopy and treating the wounded areas with a 5:5:50 Bordeaux mix
(Prakash and Raoof, 1989). Man-agement of the controllable predisposing factors,
such as drought stress, may also be beneficial. A better understanding of the
epidemiology of these dis-eases would assist these efforts. Pruning to force
synchronous flushes of foliar growth might enable the avoidance of windows of
infection for certain pathogens (Johnson, 1994b).
256 R.C. Ploetz and S. Freeman

(a) (b)) (c)

Fig. 8.15. Decline symptoms induced on ‘Tommy Atkins’ plants artificially inoculated
with isolates of: (a) C. gloeosporioides; (b) Neofusicoccum parvum; and (c) L.
theobro-mae (Photographs courtesy of D. Benscher).

Galls and scaly bark

Gall and scaly bark disorders of mango are known in several producing regions.
These diseases are usually minor problems.

Symptoms
In India, bark scaling develops as deep cracks along the entire rootstock por-tion of
the plant, and cracks may penetrate the phloem and become necrotic (Prakash and
Srivastava, 1987). These symptoms resemble those of a scaly bark disorder,
‘cuarteado’, in Colombia (Cook, 1975). In Hawaii, similar symptoms developed on
mango seedlings (Cook et al., 1971). The bark from the soil line to the first
branches was rough and scaly, and xylem pegs, 5 mm long, were evident when the
bark was removed around leaf scars and secondary branches.

In Mexico, a disorder known as ‘nanahuate’, ‘bolas’ or ‘buba of mango’,


causes galls, 5–10 cm in diameter, which resemble a cauliflower, are initially light
green, but become dark brown as they die (Fig. 8.16) (Angulo and Villapudua,
1982). The galls remain attached to trees for many years, and severely affected
branches die. Similar symptoms are found in Florida USA, and are associated with
pruning injuries. Larger galls have also been noted in Puerto Rico, as well as the
US Department of Agriculture (USDA) Agriculture Research Service (ARS) in
Miami and University of Florida in Homestead (Fig. 8.17) (Ploetz et al., 1996b; R.
Rodriguez, personal communication). Some of the latter galls are large, have
rough, scaly exteri-ors, and are usually found on the main trunk or scaffold limbs
of affected trees. Cracks may penetrate the phloem and become necrotic, but the
branch death that is associated with galls in Mexico and India has not been
observed.
Foliar, Floral and Soilborne Diseases 257

Fig. 8.16. Galls of the ‘buba’ type in Haiti (Photograph courtesy of Carolyn
Cohen, USDA, Animal and Plant Health Inspection Service (APHIS)).

Fig. 8.17. Large, 30-year-old gall on ‘Langra’ in the USDA-ARS mango


germplasm repository in Miami (Photograph courtesy of R.C. Ploetz).

Aetiology
Fusarium decemcellulare C. Brick (synonym: Fusarium rigidiuscula (Brick) Snyd.
and Hans.) causes these diseases in Florida USA, Mexico and Venezuela (Malaguti
and de Reyes, 1964; Angulo and Villapudua, 1982; Ploetz et al., 1996b). Colonies
on PDA are dark carmine-red on the underside. The fungus produces microconidia
in false heads or chains on branched and non-branched monophialides (Fig. 8.18).
Large macroconidia, 92–55 × 7–5.5 Pm, are produced in slimy yellow
sporodochia, c.1 mm in diameter. The fungus’s
258 R.C. Ploetz and S. Freeman

(a)

(b)

(c)
20 μ

Fig. 8.18. (a) Ascus and ascospores of Albonectria rigidiuscula, and (b) micro-
conidia and conidiophores, and (c) macroconidia and conidiophores of its
anamorph, Fusarium decemcellulare (Source: from CMI description no. 21).

teleomorph, Albonectria rigidiuscula (synonyms: Nectria rigidiuscula Berk. and


Broome, and Calonectria rigidiuscula) (Rossman et al., 1998), has not been
observed on mango.
Fusarium decemcellulare causes corky bark, gall, canker and dieback dis-
eases on diverse woody hosts in the subtropics and tropics (Holliday, 1980; Farr et
al., 1989; Alfieri et al., 1994). It causes an important disease of cacao (Theobroma
cacao L.), cushion gall, as well as a stem gall on loquat (Eriobotrya japonica
(Thunb.) Lindl.) and scaly bark of pongam (Pongamia pinnata (L.) Pierre). Host
specialization in the fungus has not been reported. Fusarium decemcellulare has
not been reported to cause gall and scaly bark disorders of mango in other areas.
The possible role of Agrobacterium tumefaciens was examined in Miami; however,
the bacterium could not be recovered from affected tissues (R. McGuire, Miami,
1993, personal communication). In Hawaii, microorganisms were not recovered
from affected plants (Cook, 1975).

Epidemiology
Isolates of F. decemcellulare from mango are only mildly aggressive (Ploetz et al.,
1996b), and require wounding in order to infect. A recent outbreak of scaly bark in
a commercial mango orchard in Florida USA was associated with pruning wounds.
In the cushion gall disease on cacao, F. decemcellulare interacts with several
different insect pests and pathogenic agents (Holliday,
Foliar, Floral and Soilborne Diseases 259

1980; Ploetz, 2007). These insects may facilitate infection and disseminate the
pathogen. Insect-F. decemcellulare interactions have not been investigated on
mango.

Management
No pesticides have been identified to control this problem. Measures that should be
helpful include the removal and destruction of affected branches and trees in the
orchard, disinfestation of pruning equipment to ensure that the pathogen is not
spread during pruning operations, and the use of healthy planting material in new
orchards.

Grey leafspot

Pestalotiopsis mangiferae (Henn.) Steyaert (synonym: Pestalolia mangiferae


Henn.; no teleomorph of the fungus is known) causes grey leafspot and stem-end
rot of mango fruit (Lim and Khoo, 1985; Johnson, 1994b). It is a weak pathogen
that usually requires wounding in order to infect mango. Grey leafspot is usually
unimportant and occurs mainly on unhealthy or poorly maintained trees.

Pestalotiopsis mangiferae produces abundant conidia in acervuli that develop


in grey leafspot lesions and necrotic areas on fruit (Lim and Khoo, 1985). As
lesions age, black columns of spores emanate through ruptures in the host
epidermis. Conidia are produced that have three thick-walled, brownish,
concolorous median cells and thin-walled, hyaline apical and basal cells; the apical
cells bear three characteristic appendages (Fig. 8.19). Conidia are 20 × 5 Pm,
fusiform and straight to slightly curved. Two other species of Pestalotiopsis that
occur on mango produce larger conidia, Pestalo-tiopsis mangifolia Guba and
Pestalotiopsis versicolor Speg. (synonyms: Pestaloti-opsis cliftoniae Tracy and
Earle and Pestalotiopsis coccolobae Ellis and Everh.).
On leaves, symptoms are light grey spots, usually 2–20 mm in diameter (Lim
and Khoo, 1985). These may coalesce to form large patches of necrotic tissue on
leaves. Lesions are surrounded by dark, raised margins, and as they mature, raised
black dots (which are columns of the pathogen’s conidia) are evident in lesion
centres. Although most cultivars are susceptible, specific control measures are
usually not required. Dithiocarbamate fungicides con-trol this disease.

Leaf blight

This disease has been reported in India and Nigeria (Hingorani et al., 1960; Cook,
1975; Okigbo, 2001; Okigbo and Osuinde, 2003), and the causal fun-gus,
Macrophoma mangiferae Hingorani and Sharma (Ascomycota), has also been
intercepted in shipments to the USA from Mexico (Systematic Mycol-ogy and
Microbiology Laboratory, USDA-ARS, Beltsville). This is not a serious problem.
260 R.C. Ploetz and S. Freeman

50 μm (a)

(b)
(c)
10 μm

Fig. 8.19. (a) Vertical section of an acervulus, (b) mature conidia, and (c)
conidiog-enous cells and developing conidia of Pestalotiopsis mangiferae
(Source: from CMI description no. 676).

Macrophoma mangiferae produces subepidermal, globose pycnidia, 77–231


Pm in diameter. Hyaline conidiophores, 1.5–2 × 8–11 Pm, produce unicellular
conidia, 5.3–7 × 10.5–24.5 Pm. No teleomorph is known. Since the genus Mac-
rophoma has been placed in synonymy with Sphaeropsis, this species should be
redescribed.
Leaves, twigs and fruit are affected, especially when the latter are stored.
Small, yellow spots gradually enlarge to become brown with wide purple margins.
The lesions are initially circular, but develop an irregular appear-ance and may
encompass large portions of the leaf surface. Pycnidia form most frequently on the
underside of leaves. Elliptical stem lesions are infre-quent but can girdle stems. In
India, the disease is most serious during the rainy season (Verma and Singh,
1996b). Macrophoma mangiferae survives in pycnidia that develop on bark of
twigs of young mango plants, blighted leaves and as dormant mycelium in wood
(Verma and Singh, 1996a). Okigbo (2001) reported that the fungus survived
adverse conditions best in stems and branches.

Four applications of captan, Bordeaux mixture, captafol, carbendazim and


thiophanate-methyl were effective on young plants (Verma and Singh, 1996a). The
bacterium Bacillus subtilis NCIB 3610, isolated from soil under a mango tree,
inhibited M. mangiferae on agar plates, and symptoms were reduced in the field by
the application of the antagonist (Okigbo and Osuinde, 2003).
Foliar, Floral and Soilborne Diseases 261

Malformation

Malformation is one of the most destructive mango diseases (Ploetz, 2001).


Although trees are not killed, the vegetative phase of the disease impedes canopy
development and the floral phase dramatically reduces fruit yield. Based on the
incidence and severity of malformation in Egypt, an estimated US$15 million of
fruit were lost in 1998 (Ploetz et al., 2002). Losses in more important producing
countries (i.e. India) are undoubtedly much greater.
Malformation was first reported in India in 1891 (Kumar and Beniwal, 1991),
and has subsequently been observed in Brazil, Myanmar, Egypt, El Salvador, India,
Israel, Malaysia, Mexico, Nicaragua, Oman, Pakistan, South Africa, Sudan, Spain,
Swaziland, Uganda and the USA (Flechtmann et al., 1973; Crookes and
Rijkenberg, 1985; Lim and Khoo, 1985; Kumar and Beni-wal, 1991; Ploetz, 2001;
Kvas et al., 2007; S. Freeman, Bet Dagan, 2007, unpub-lished data; G.I. Johnson,
Jamison, Australia, 2007, personal communication; J.F. Leslie, Manhattan, Kansas,
2007, personal communication). Since the pathogen is easily disseminated in
infected germplasm and there are con-spicuous gaps in the disjunct distribution
(note African and American records), malformation is probably more widely
spread.

Symptoms
Malformation affects vegetative and floral meristematic tissues (Fig. 8.20) (Ploetz,
2001). Vegetative malformation is most serious on seedlings and small plants in
nurseries, especially where seedlings are grown beneath affected trees, a common
practice in the Middle East (Ploetz et al., 2002; Youssef et al., 2007). Vegetative
malformation also occurs on mature trees. Apical and axillary buds produce
misshapen shoots with shortened inter-nodes and dwarfed leaves that are brittle and
recurve towards the sup-porting stem (Fig. 8.20). Shoots may not expand fully,
resulting in a bunched appearance (i.e. the ‘bunchy-top’ symptom of the disease).

Floral malformation is most important. Affected inflorescences usually do not


set fruit or fruit are aborted. Primary or secondary axes on affected panicles are
shortened, thickened and highly branched (Fig. 8.20). Malformed panicles produce
up to three times the normal number of flowers, range from one-half to two times
the normal size, and have an increased proportion of male to perfect flowers.
Malformed panicles may also produce dwarfed and distorted leaves (exhibit
phyllody).

Aetiology
The aetiology of malformation has been controversial for almost as long as the
disease has been recognized (Ploetz, 2001). Suggested causes include mites
(Narasimhan, 1954), nutritional problems (Prasad et al., 1965), physio-logical or
hormonal imbalances (Dang and Daulta, 1982; Singh and Dhillon, 1989), viruses
(Kauser, 1959) and unknown causes (Kumar and Beniwal,
1991). Summanwar et al. (1966) demonstrated that a fungus, identified then as
Fusarium moniliforme Sheld., was responsible for the floral phase of this disease.
Varma et al. (1974) later showed that F. moniliforme also caused
262 R.C. Ploetz and S. Freeman

(a)

(b)

Fig. 8.20. Among the symptoms caused by malformation are: (a) in panicles, an in-
crease in the size and number of flowers and interspersed floral and vegetative organs
(phylody); and (b) in vegetative shoots, compact or retarded growth of buds and brittle,
dwarfed and recurved leaves. Symptoms in (a) are on ‘Haden’ in Michoacan, Mexico and
are associated with an undescribed species in the Gibberella fujikuroi species complex,
whereas (b) is on a ‘Van Dyke’ plant artificially inoculated with an isolate of Fusarium
mangiferae (Photographs courtesy of R.C. Ploetz).
Foliar, Floral and Soilborne Diseases 263

vegetative malformation. This pathogen has had several synonyms in the literature,
including: Fusarium subglutinans (Wollenweb. and Reinking) Nelson, Toussoun
and Marasas; F. moniliforme Sheldon var. subglutinans Wollenweb. and Reinking;
and F. moniliforme Sheldon emend. Snyd and Hans. ‘Subglutinans’ sensu Snyd.,
Hans. and Oswald.
In 2002, 29 strains of the pathogen from Egypt, Florida USA, Israel, Malaysia
and South Africa were redescribed as a new species in the Gibberella fujikuroi
species complex (GFSC), Fusarium mangiferae Britz, Wingfield and Marasas
(Steenkamp et al., 2000; Britz et al., 2002). Fusarium mangiferae resem-bles
morphologically other members of the GFSC. It was established based on E-
tubulin and histone H3 DNA sequences, subtle morphological differ-ences, and
because most of the examined strains had been shown in previous studies to cause
malformation on artificially inoculated mango. The presence of F. mangiferae has
been verified in India (O’Donnell et al., 1998; Zheng and Ploetz, 2002), Oman
(Kvas et al., 2007) and Spain (S. Freeman, Bet Dagan, 2007, unpublished results).
Although a recent report from Pakistan mentions F. mangiferae, the identity of the
pathogen there is not clear since the authors only discussed morphological
characteristics of the pathogen (Iqbal et al., 2006).

Based on DNA sequence data (O’Donnell et al., 1998, 2000; Steenkamp et al.,
1999, 2000), F. mangiferae is related to a lineage that includes Fusarium fujikuroi
Nirenberg, Fusarium proliferatum (Matsushima) Nirenberg, and Fusarium
sacchari (E. J. Butler) W. Gams (Marasas et al., 2006), and corresponds to the
‘Asian Clade’ of O’Donnell et al. (1998). Based on combined sequence data for
five genes, the closest known relative of F. mangiferae is an isolate from tropical
rainforest soil in Papua New Guinea (Marasas et al., 2006).
Fusarium mangiferae produces white, floccose mycelium on PDA with light-
to dark-purple pigments in the agar (Leslie and Summerell, 2006). Cream-coloured
sporodochia on carnation leaf agar (CLA) produce abun-dant thin-walled, long,
slender and straight to slightly curved, three- to five-septate macroconidia with
curved apical cells and foot-shaped basal cells (Fig. 8.21). Single celled, rarely
single septate, obovoid microconidia are produced in false heads on polyphialides
with two to five conidiogenous openings and on monophialides. Microconidial
chains and chlamydospores are absent.

A second species, Fusarium sterilihyphosum Britz, Wingfield and Marasas,


was described originally for isolates from a small area in South Africa (Britz et al.,
2002). In subsequent work, it was detected in Brazil (Ploetz, 2003; K. O’Donnell,
unpublished results), and was recently shown to cause malfor-mation in Brazil after
artificial inoculation (Lima et al., 2006b). On PDA, colo-nies of F. sterilihyphosum
produce white, floccose mycelium with rose to light purple pigmentation in the
agar (Leslie and Summerell, 2006). Uncommon, cream- to orange-coloured
sporodochia are produced on CLA that produce rare, long, slender, three to five-
septate macroconidia (Fig. 8.22). On mono-and polyphialides, obovoid, oval to
allantoid microconidia that are usually single celled are produced on false heads.
Distinctive sterile coiled hyphae are produced by some isolates of this species.
264 R.C. Ploetz and S. Freeman

(a) (c) (e)

(b) (d) (f )

Fig. 8.21. Microscopic features of Fusarium mangiferae: (a) and (b), macroconidia;
(c) and (d), microconidia, and (e) and (f), microconidia in situ on carnation leaf
agar. Scale bars for (a)–(d) = 25 Pm, and (e)–(f) = 50 Pm (Photographs courtesy
of Suzanne Bullock).

Fusarium sterilihyphosum is relatively uncommon in South Africa and Brazil


where, respectively, F. mangiferae and a third, unnamed species that resembles F.
sterilihyphosum morphologically, predominate. The latter taxon is phylogenetically
distinct from F. mangiferae and F. sterilihyphosum, produces a unique teleomorph
in the GFSC, and has been shown to cause malforma-tion (Lima et al., 2006a, b).

Fusarium mangiferae has not been found in either Brazil (Lima et al., 2006a,
b) or Mexico (Rodríguez-Alvarado et al., 2006, 2008). In the latter studies, isolates
that were recovered from malformed trees resembled F. sterilihypho-sum in that
they induced malformation symptoms, formed sterile coiled hyphae and produced a
PCR fragment that is also produced by isolates of F. sterilihyphosum (see below).
Translation elongation factor-1D DNA sequences for isolates from several areas in
Mexico are identical, but differ signifi-cantly from other taxa in the GFSC; they
probably represent a new species (Rodríguez-Alvarado et al., 2006, 2008; K.
O’Donnell, Peoria, 2007, personal communication). Additional work is needed to
clarify relationships among the strains in Brazil and Mexico, and whether they are
found elsewhere in the Amer-icas. Likewise, whether F. mangiferae is found
outside Florida USA in the western hemisphere should be determined; it
predominates in the eastern hemisphere.
PCR primer pairs have been used to diagnose some of the above taxa. Zheng
and Ploetz (2002) developed a pair, 1-3F/R, that amplifies a 608 bp fragment for F.
mangiferae. It has been used extensively for diagnostic pur-poses (Youssef et al.,
2007). Another pair, 61-2F/R, developed to diagnose Fusarium verticilloides
(published as F. moniliforme in Müller et al., 1999), did not amplify F. mangiferae
DNA, but when amplification protocols were mod-ified, amplified a 445 bp-
fragment for strains of F. sterilihyphosum and the new species from Mexico
(Zheng and Ploetz, 2002; Rodríguez-Alvarado et al., 2008). It has not been tested
with the unnamed pathogen from Brazil.
Foliar, Floral and Soilborne Diseases 265

(a) (c) (e) (g)

(b) (d) (f ) (h)

Fig. 8.22. Microscopic features of Fusarium sterilihyphosum: (a) and (b), macroconidia; (c)
and (d), microconidia; (e) and (f), coiled hyphae; and (g) and (h), microconidia in situ on
carnation leaf agar. Scale bars for (a)–(d) = 25 Pm and (e)–(f) = 50 Pm (Photographs
courtesy of Suzanne Bullock).

Three other taxa have been associated with mango malformation. Fusar-ium
oxysporum Schlecht emend. Snyder and Hansen (Fig. 8.23) was reported in Egypt
and Mexico (Bhatnagar and Beniwal, 1977; Kumar and Beniwal, 1991), but these
reports appear to be erroneous since bona fide, vouchered specimens have neither
been described nor shown to cause the disease (Plo-etz, 2001; Rodríguez-Alvarado
et al., 2008). Fusarium sp. nov. (Britz et al., 2002) and F. proliferatum (Gibberella
intermedia (Kuhlman) Samuels, Nirenberg and Seifert) were recovered from
malformed mango trees in Malaysia (Leslie, 1995), but their pathogenicity has not
been determined.

Epidemiology
Although malformation has been reproduced with F. sterilihyphosum and the
unnamed taxa from Brazil and Mexico, no work has been conducted on the
epidemiology of disease that is caused by these pathogens. Thus, results below are
from work on F. mangiferae or what is presumed to be this species. Fusarium
mangiferae is spread by grafting and in infected nursery stock (Prakash and
Srivastava, 1987). Since seed do not appear to harbour the fungus (Saeed and
Schlosser, 1972; Youssef et al., 2007), seedlings should be disease free.
Microconidia of F. mangiferae are probable infective propagules since they are the
primary spores that are produced by the fungus and form profusely on dead
malformed tissues. The disease spreads slowly in orchards, perhaps because
conidia of the pathogen die quickly when exposed to sun-light (Manicom, 1989).
Experimentally, populations of F. mangiferae in infected panicles in Egypt and
Israel declined rapidly during the summer (Youssef et al., 2007). Wounding
enhances infection and subsequent disease develop-ment (Ploetz, 2001).
266 R.C. Ploetz and S. Freeman

(a) (b)

(c)

(e) (d)

Fig. 8.23. Microscopic features of Fusarium oxysporum: (a) sporodochia, (b)


macro-conidia, (c) microconidia in false head on monophialide, (d) terminal and
intercalary chlamydospores, and (e) macroconidia and microconidia (Photographs
courtesy of K. O’Donnell).

The mango bud mite, Aceria (Eriophyes) mangiferae Sayed, is often observed
in high numbers on malformed trees and has been indicted as the cause, or a factor
in the development, of this disease (Narasimhan, 1954, 1959; Nariani and Seth,
1962). Circumstantial evidence indicates that the mite does not cause malformation
(Ploetz and Prakash, 1997); for example, A. mangiferae is present in Australia, but
the disease is not (Ridgeway, 1989). However, A. mangiferae probably vectors the
pathogen. It has been recovered from the mite’s body on culture media (Crookes
and Rijkenberg, 1985; Sattar, Ismailia, 2006, personal communication), and was
recently shown to adhere to its body (Gamliel-Atinsky, Freeman and Palevsky,
unpublished data) (Fig. 8.24). The mite cannot ingest the pathogen, due to its small
mouthparts. However, it was able to move spores of F. mangiferae to infection
courts in mango buds via external contamination of its body, and increased
infection
Foliar, Floral and Soilborne Diseases 267

GFP-labelled microconida of
Fusarium mangiferae

Aceria mangiferae

20.0 μm

Fig. 8.24. Body of the mango bud mite, Aceria mangiferae, to which green
fluorescent protein (gfp)-labelled microconidia of Fusarium mangiferae have
adhered (Photograph courtesy of E. Gamiel-Atinsky).

of buds by the pathogen (Gamliel-Atinsky et al., 2007). Presumably, the mite’s


feeding on buds facilitated infection. Aceria mangiferae does not appear to play a
significant role in disseminating the pathogen among trees. In Israel, mites were
not found in traps that were designed to monitor their movement in a malformed
mango orchard, although high numbers of F. mangiferae conidia were recorded in
the air in this orchard (Gamliel-Atinsky et al., 2007).

The distribution of F. mangiferae in affected trees suggests that vegetative and


floral buds are the primary sites of infection and that systemic coloniza-tion of
older, subtending tissues does not occur. Freeman et al. (1999) trans-formed
isolates of F. mangiferae from mango with the uidA reporter gene (E-
glucuronidase), and used them to artificially inoculate mango. Their results verified
that bud and flower tissues of the host are primary infection sites, and that wounds
provide points of entry for the pathogen. In Florida USA, F. mangiferae was
restricted almost entirely to malformed floral and vegetative tissues (Ploetz, 1994).
Levels of infection were highest in malformed flowers and vegetative shoots (c.65–
85%), were much lower or non-existent in asymp-tomatic tissues (0–11%), and
were rare in branches (0–4%) even when they supported malformed flowers or
shoots. Remnant infections of F. mangiferae in scaffold branches and trunks were
restricted almost exclusively to branch scars or dormant apices (Freeman,
unpublished data). Reports of root infection by F. oxysporum (Kumar and Beniwal,
1991) or F. mangiferae (Abdel-Sattar, 1973; Kumar and Beniwal, 1991) causing
malformation in seedling plants have not been corroborated. Although roots can be
infected by
268 R.C. Ploetz and S. Freeman

F. mangiferae, these infections are not systemic and do not appear to result in
symptom development (Youssef et al., 2007).
The localized and variable levels of infection by F. mangiferae that have been
noted in diseased and non-symptomatic tissue (Ploetz, 1994; Youssef et al., 2007),
suggest that there are thresholds of infection, whereby malfor-mation develops
only after a sufficient proportion of a host meristem is colo-nized by the pathogen.
This hypothesis is supported by the long latent period that exists before symptoms
develop in artificially inoculated plants and the hormonal perturbations that
probably occur when meristematic tissues are infected with this pathogen (van
Staden et al., 1989; van Staden and Nichol-son, 1989; Ploetz, 2001).

Management
Management of malformation can be difficult. New plantings should be established
with pathogen-free nursery stock. Scion material should never be taken from an
affected orchard, and any affected plants that are observed in the nursery should be
removed and burned immediately. Nurseries should not be established in orchards
that are affected by malformation. Once the disease is found in an orchard, control
is possible, but time consuming. In these cases, cultural management has been most
effective (Narisimhan, 1959; Singh et al., 1974; Manicom, 1989). All affected
terminals and the subtending three nodes are cut from trees, removed from the field
and burned. Unfortu-nately, producers may be unwilling to devote the effort that is
required to ensure that this approach succeeds. In addition, it may be difficult to
impose this treatment on large trees.

A diverse array of pesticides, hormones and growth regulators has been tested
against malformation, but these measures have been marginally effec-tive. Singh et
al. (1994) obtained moderate control with sulfates of cobalt (Co), cadmium (Cd)
and nickel (Ni) in India, but it is unlikely that these toxic com-pounds could be
used safely. Darvas (1987) reduced the percentage of mal-formed inflorescences
from 96% to 48% by injecting ‘Keitt’ trees with the fungicide fosetyl-Al. This
reduction was significant (P < 0.05), but the increase in fruit yield, 46–95 kg of
fruit/tree, was not. Other fungicidal compounds have been generally less effective
(Diekman et al., 1982; Chakrabarti and Ghosal, 1989). In general, the protected,
internal location of the pathogen in affected trees makes it difficult to control this
disease. When applied as foliar sprays or via continuous drip irrigation, prochloraz
reduced the severity of malformation significantly in Israel, but this was dependent
on the timing of application (Freeman et al., unpublished data). Although disease
was not completely controlled, this and other systemic fungicides might be useful
in future integrated management programmes that would incorporate other
measures such as removal of symptomatic terminals and use of tolerant cultivars.

Prakash and Srivistava (1987) indicated that ‘There is great variation in the
susceptibility of existing varieties.’ Unfortunately, controlled inoculations have not
been used to determine cultivar resistance, and these reports have come from non-
replicated tests; cultivars listed as ‘resistant’ may have come
Foliar, Floral and Soilborne Diseases 269

from healthy nursery stock or may have escaped infection once planted in the field
(Ploetz, 2001). For example, Bastawros (1996) reported that two newly introduced
cultivars in Egypt, ‘Kent’ and ‘Keitt’, were immune (0% disease); however, these
cultivars are susceptible in Florida USA (Ploetz, unpublished data). Controlled
inoculations with grafted plants of different genotypes and quantified levels of
virulent isolates of F. mangiferae have not been reported.

Recently, Ploetz (unpublished data) utilized previously described proto-cols


(Freeman et al., 1999) to assess malformation development on grafted plants of
diverse genotypes. Disease development was affected by: the length of time after
inoculation and inoculated apical buds remained dormant after inoculation; the
isolate of F. mangiferae that was used for inoculation; and mango genotype.
Virulent isolates, patience (latent period ranged from 40 to 210 days), and
sufficient replication were needed to successfully conduct screenings for response
to malformation. Future work is warranted to investi-gate attributes that are related,
and might predict resistance, to this disease.
The symptoms of malformation suggest that a hormone imbalance occurs in
affected tissues. Singh and Dhillon (1989) assayed levels of indole acetic acid
(IAA), gibberellic acid (GA3) and zeatin in malformed and healthy mango
seedlings. Whereas IAA and GA3 levels were, respectively, about ten and five
times lower in malformed plants, levels of zeatin were about five times higher. Van
Staden and colleagues (Nicholson and van Staden, 1988; van Staden and
Nicholson, 1989; van Staden et al., 1989) examined specific cytokinins produced
by mango and ‘F. moniliforme’ (presumably F. mangiferae). They determined that
the cytokinin complements in healthy and malformed panicles differed
qualitatively and quantitatively, and that the pathogen pro-duced some of the
hormones and metabolites that were implicated in dis-ease development. However,
it was impossible to assign unequivocal roles for production of hormones by the
host and pathogen and the subsequent development of symptoms. For example,
whether production of hormones by the pathogen directly caused the noted changes
or whether hormone pro-duction by the host was somehow altered in the presence
of the pathogen was not clear. Additional work would be needed to clarify these
interactions, and whether F. sterilihyphosum and the unnamed pathogens in Brazil
and Mexico also produce cytokinins or other hormones in affected plants.

Parasitic plants

The family Loranthaceae contains several parasitic plant species that affect mango.
In Malaysia, Dendrophthoe (fomerly Loranthus) pentandra Linn. is the most
important species (Lim and Khoo, 1985). Other Dendrophthoe spp., Elytranthe
spp. and Viscum spp. are also known in Malaysia, but are less important. In India,
Dendrophthoe falcata (formerly Loranthus longiflorus) is common, and other, less
frequently encountered, species include Macrosolen cochinchinensis, Helicanthes
elasticus and Elytranthe capitellata (Majumder and Sharma, 1990). These parasites
are usually only important in neglected
270 R.C. Ploetz and S. Freeman

orchards. Although they are photosynthetically self-sufficient, the plants obtain


water and minerals from the host plant via haustoria that penetrate and colonize the
host vascular system. In severe cases, the removal of water and minerals from
parasitized branches is sufficient to reduce the vitality and yields of trees.

Since the appearance of these plants is distinct from the mango host, they are
easily distinguished in infected trees (Lim and Khoo, 1985). The points at which
the mango host is penetrated are usually characterized by swollen growths called
burrs. Lim and Khoo (1985) and Majumder and Sharma (1990) indicated that
affected portions of trees should be removed far enough below burrs to remove
haustoria of the parasite. After affected tissues are removed, cut sur-faces can be
treated with creosote or other wound dressings. These plants can also be treated
with herbicides, such as 2,4-dichlorophenoxyacetic acid (2,4-D), but these are
dosage sensitive treatments and pose a risk to the host plant.

Phoma blight

Phoma blight is widespread in India (Prakash and Singh, 1977). It occurs only on
old leaves. Initially, lesions are minute and yellow-brown (Prakash and Singh,
1977). As they expand they darken to brown or cinnamon, become irregular, and
may ultimately develop dark margins and dull-grey centres. In severe cases,
necrotic patches as large as 13 cm in diameter may form that cause defoliation. The
disease is caused by Phoma glomerata (Corda) Wollenw. and Hochapf (Prakash
and Singh, 1977). It forms globose to obpyriform, light-coloured to car-bonaceous
pycnidia that average 30–400 Pm in diameter. Pycnidia have one to three ostioles,
form singly or in clusters, and have short necks. On PDA, pyc-nidia and conidia
are abundant. Conidia are hyaline to dark coloured, ovoid to ellipsoid, unicellular
or occasionally bicellular, and average 8.3 × 3.2 Pm.

Phoma leafspot

Another Phoma sp. causes a leafspot in India (Prakash and Singh, 1976b), and is
referred to as phoma leafspot. On young leaves, Phoma sorghina (Sacc.) Boerema.
Doren. and Vankest causes irregular to roughly circular, water-soaked spots, up to
2.5 mm in diameter. Lesions are brown with a yellow to brown margin. Lesions on
midribs are elongated and more conspicuous, and may coalesce to up to 14 cm in
diameter. They can be confused with those caused by anthracnose.

Pink disease

A basidiomycete, Erythricium salmonicolor (Berk. and Broome) Burdsall (syn-


onyms: Corticium salmonicolor Berk. and Broome, and Phanerocbaete salmoni-
color (Berk. and Broome) Jülich; anamorph: Necator decretus Massee) causes
Foliar, Floral and Soilborne Diseases 271

pink disease. Pink disease affects many economically important woody plants in
the humid tropics, where it is one of the most destructive diseases of mango
(Holliday, 1980). The disease is also known as cobweb, rubellosis and thread blight
(Prakash and Srivistava, 1987). It has been most thor-oughly studied on rubber,
Hevea brasiliensis, an important host in South-east Asia (Rao, 1975). On mango,
pink disease can significantly reduce tree vigour and yield, especially in 6- to 15-
year-old trees (Lim and Khoo, 1985).
Symptoms first appear as white, felty mycelial threads on twigs and branch
crotches (Lim and Khoo, 1985). If favourable conditions persist, the mycelial
threads coalesce to form a rough, pink crust on the bark surface. This stage usually
takes 1 to several months to develop and coincides with the penetration of bark and
internal colonization of the tree. Affected bark often cracks. As the fungus kills the
vascular and cambial areas beneath the bark, branches above the colonized areas
die, resulting in a sparse, patchy canopy.
Two types of sporulation occur (Holliday, 1980; Lim and Khoo, 1985).
Erythricium salmonicolor produces a smooth, clammy, pinkish white hyme-nium
over the pink crust it forms on bark. Basidiospores form on the hyme-nium and are
borne on sterigmata on narrowly clavate to cylindrical basidia (Fig. 8.25).
Basidiospores are hyaline, broadly ellipsoidal and 8–10 × 5–7 Pm. Conidia of N.
decretus, which are produced in reddish-orange sporodochia, are hyaline,
ellipsoidal, unicellular and 10–18 × 6–12 Pm. Although the infec-tion process has
apparently not been studied in mango, basidiospores can infect rubber trees
(Holliday, 1980). The anamorph and teleomorph are

(a)
100 μ
(b) (f )

(e)
(c)

(d) 20 μ

Fig. 8.25. (a) Conidium-bearing pustule, and (f) conidiogenous cells and conidia
of Necator decretus, and (b) hymenium, (c) basal hyphae, (d) immature and
mature basidia, and (e) basidiospores of its teleomorph, Erythricium
salmonicolor (Source: from CMI description no. 511).
272 R.C. Ploetz and S. Freeman

formed during wet conditions, and conidia and basidiospores are dispersed by
rainsplash and wind.
Pink disease management relies on early detection and removal of affected
tissues from orchards. When removal of syptomatic tissues is imprac-tical, control
depends upon treatment with fungicides. Several protectant and systemic
fungicides are effective (Lim, 1994). They should be used as soon as symptoms are
evident, and as long as the disease is present. All cul-tivars of mango that have
been tested in Malaysia are susceptible (Lim and Khoo, 1985).

Powdery mildew

Powdery mildew is a widespread and important disease of leaves, panicles and


fruit. The disease can result in yield reductions as high as 90%, due mainly to its
effect on fruit set and development (Schoeman et al., 1995).

Symptoms
Mango cultivars vary in their response to powdery mildew (Palti et al., 1974). On
the most susceptible cultivars, virtually all foliar, floral and fruit parts of the plant
are affected (Plate 46). Powdery growth can cover all tissues on panicles, resulting
in a brown, shrivelled necrosis. Since fruit set and reten-tion can be affected, the
disease can have a profound impact on yield. Foliage can also be damaged
significantly, and young leaves are most susceptible. White, powdery coatings of
conidia develop on either side or both sides of leaves, depending on the cultivar.
When damage occurs on the undersides of leaves it is often restricted to the mid-
rib. In all cases, leaves become dis-torted, and affected areas turn purple and
ultimately necrotic.

Aetiology
Powdery mildew is caused by Oidium mangiferae Berthet, a host-specific fun-gus
(Prakash and Srivistava, 1987; Ploetz and Prakash, 1997). It was first described in
Brazil (Berthet, 1914), and is now recognized in most mango-producing regions
(Palti et al., 1974). Conidium and haustorium traits indi-cate that O. mangiferae
belongs to the Erysiphe polygoni group (Johnson, 1994a). Although the pathogen
was originally classified as Erysiphe cichoracearum by Wagle (1928), Uppal et al.
(1941) noted that it produced saccate and lobed appressoria, which are not
characteristic of E. cichoracearum. The pathogen has no known teleomorph, a
common trait for powdery mildew fungi in the tropics (Holliday, 1980). Conidia of
O. mangiferae are unicellular, hyaline, elliptical to barrel-shaped and measure 33–
43 × 18–28 Pm (Uppal et al., 1941; Palti et al., 1974). They are produced in large
numbers on host surfaces, and impart a powdery appearance to affected tissues
(Plate 46). The lengths of germ tubes vary depending upon RH, and they terminate
in appressoria. Glob-ular haustoria form in host epidermal cells. Conidiophores are
of the pseudoid-ium type, with two to four septa and a straight basal cell
(Boesewinkel, 1980).
Foliar, Floral and Soilborne Diseases 273

Epidemiology
Powdery mildew is most severe during cool, dry weather. Conidia are dis-
seminated by wind and are released on a diurnal basis (Schoeman et al., 1995).
Peak spore release, between 11:00 to 16:00 h, was positively correlated with hourly
temperature and negatively correlated with RH, vapour pres-sure deficit and leaf
wetness (all P < 0.01). Conidia germinate at temperatures ranging from 9 to 32C
(23C is optimal), and at RH as low as 20% (Palti et al., 1974). Since germination
occurs in such a wide range of RH, disease develop-ment is usually independent of
this parameter. Infection can occur within 5–7 h, and conidia are produced within 5
days of infection. Disease develop-ment occurs within a very broad range of
temperatures, 10–31C. Gupta (1989) reported that dry weather encouraged disease
development.

Management
Mango cultivars vary in their resistance to powdery mildew (Palti et al., 1974).
‘Zill’, ‘Kent’, ‘Alphonso’, ‘Seddek’ and ‘Nam Doc Mai’ are very sus-ceptible;
‘Haden’, ‘Glenn’, ‘Carrie’, ‘Zebda’, ‘Hindi be Sennara’, ‘Ewaise’ and ‘Keitt’ are
moderately susceptible; and ‘Sensation’, ‘Tommy Atkins’ and ‘Kensington’ are
slightly susceptible (Ploetz et al., 1994; Nofal and Haggag, 2006). In India, Tiwari
et al. (2006) reported that ‘Baigan Phalli’, ‘Barbalia’, ‘Dabari’, ‘Dilpasand’,
‘Khirama’, ‘Nagarideeh’, ‘Oloor’ and ‘Totapari’ were highly resistant and
‘Amrapali’ was most susceptible.
Schoeman et al. (1995) recommended that the first fungicide application to
control this disease should occur when panicles begin to change colour. Assuming
an effective period of 3 weeks for a given application, they con-cluded that
applications should continue every third week until panicle sus-ceptibility
decreased at the end of fruit set. Powdery mildew is easily controlled with S
(Johnson, 1994a). Other fungicides are effective, but many have negative
environmental impacts (Ray, 2003; Tavares et al., 2004). Foliar sprays of di-
potassium hydrogen orthophosphate (K2HPO4) and potassium di-hydrogen
orthophosphate (KH2PO4), systemic fungicides, and an alterna-tion of fertilizer
and systemic fungicides inhibited powdery mildew on pan-icles (Reuveni et al.,
1998; Nofal and Haggag, 2006). Treatments of the fertilizers with half or a quarter
of the recommended rate of sterol-inhibitor fungicides and Kresoxym-methyl
provided protection comparable with or superior to that of standard fungicides
alone (Oosthuyse, 1998; Reuveni et al., 1998). Sulfur can burn flowers and young
fruit during warm, sunny condi-tions (Johnson, 1994a), and three fungicides used
during bloom, dinocap, fenbuconazole and hexaconazole, can reduce pollen
germination (Dag et al., 2001).

Scab

Elsinoë mangiferae Bitancourt and Jenkins (anamorph: Sphaceloma mangiferae


Bitancourt and Jenkins) causes scab on mango (Bitancourt and Jenkins, 1943;
Cook, 1975). The disease was first recognized in Cuba and Florida USA in the
274 R.C. Ploetz and S. Freeman

1940s and is now widespread in the western hemisphere. Scab is important in


nurseries since young host tissues are most susceptible, and because moist
environments aid infection (Ruehle and Ledin, 1955). Lesions, usually first
observed on the underside of leaves, are dark brown to black, and 1–2 mm in
diameter. They may enlarge to 5 mm in diameter and become light grey with
narrow, dark borders. Affected foliage develops a distorted appearance, and
greyish blotches are produced on stems.
Elsinoë mangiferae produces brownish ascocarps, 30–48 × 80–160 Pm, in the
host epidermis. Globular asci, 10–15 Pm in diameter, are dispersed in ascocarps,
and contain one to eight hyaline ascospores. Ascospores are 4–6 × 10–13 Pm, three
septate and constricted at the median septum; the sub-apical cell is longitudinally
septate. Conidiophores of S. mangiferae are erect, sinuous, 2.5–3.5 × 12–35 Pm,
and wider at the base. Conidia are brown, one or two celled, and 2–4 × 6–29 Pm.

Young host tissues are most susceptible. Rainy weather promotes sporu-lation
of the fungus, but specific information is not available on the epidemi-ology of
scab. Whether conidia and ascospores are infectious is not known.

Seca and sudden decline

This is a disease that is known by several different names in Brazil and the Middle
East and is the only one that routinely kills mango trees. ‘Seca’ (dry-ing), ‘murcha’
(withering), branch blight and Recife sickness in Brazil, was first recognized in
Pernambuco in 1938, and is now also found in Bahia, Goias, the Federal District,
Rio de Janiero and São Paulo (Ribeiro, 1997; Colo-simo et al., 2000; Silveira et al.,
2006). It threatens neighbouring states due to its efficient movement in infected
propagation materials, on pruning equip-ment, and via a mobile beetle vector.

In 1998, a disease termed ‘sudden decline’ began to kill trees in Oman (Al
Adawi et al., 2003, 2006), about the same time a similar problem (i.e. quick decline
or sudden death) was observed in Pakistan (Malik et al., 2005). In many ways these
diseases resembled seca. Circumstantial evidence sug-gested that the disease was
introduced from Brazil by a producer with orchards in Oman and Pakistan (M.
Deadman, Muscat, 2005, personal com-munication). By 2007, many mango-
producing areas in Oman and Pakistan were affected and uncontrolled
dissemination of infected germplasm may have spread the disease elsewhere in the
region. Its spread into India should be investigated (A.W. Cooke, Indooropilly,
2007, personal communication).

Symptoms
Symptoms include: discoloration of the vascular cambium; exudation of an amber-
coloured gum from the trunk and branches, particularly from galler-ies of the
putative beetle vector of the pathogen; wilting; rapid death of branches and entire
trees without defoliation; and a scorched appearance of dead trees (Plate 47)
(Junqueira et al., 2002; Al Adawi et al., 2006). On grafted trees, scions, rootstocks
or both may be susceptible and exhibit vascular
Foliar, Floral and Soilborne Diseases 275

symptoms. In Oman, where susceptible Omani seedlings are used as root-stocks,


the disease is frequently a problem of rootstocks (Al Adawi et al., 2006), whereas
in Brazil, the disease is associated with the scion (P < 0.01) (Colosimo et al.,
2000); rootstock cultivar had an insignificant impact on dis-ease development in
the latter work (P > 0.05). When disease development begins in the canopy,
symptoms may initiate in a branch or portion of a tree, but death of the entire plant
usually follows. Where root/rootstock infection is involved, sudden death of the
entire tree usually occurs.

Aetiology
Ceratocystis fimbriata Ellis and Halst. sensu lato (s.l.) (anamorph: Thielaviopsis
sp.) was reported in Brazil in the 1930s (Viegas, 1960; Ribiero, 1980; Silveira et
al., 2006), and is recognized as the primary cause of seca. Diplodia recifiensis
Batista (= Lasiodiplodia theobromae?) was indicted as the cause of Recife sick-
ness in Brazil (Batista, 1947), but it probably plays no role or a secondary role in
the development of this disease (see below). In Oman, C. fimbriata s.l. causes
sudden decline, but L. theobromae and Ceratocystis omanensis Al Subhi, M.J.
Wingf., M. van Wyk and Deadman are also associated with the disease (Al Adawi
et al., 2006; van Wyk et al., 2007). The ease with which L. theobromae and the
difficulty with which C. fimbriata s.l. are recovered from affected trees may have
been responsible for previous assumptions that ‘D. recifiensis’ caused Recife
sickness in Brazil and L. theobromae caused sudden decline in Oman (Batista,
1947; Ploetz and Prakash, 1997; Al Adawi et al., 2003, 2006).
Ceratocystis contains many pathogens, particularly of trees (Kile, 1993). The
wide host range of C. fimbriata s.l. led Webster and Butler (1967) to hypothesize
that it was a species complex, and DNA sequences have begun to delineate some of
the host-specific, often morphologically indistinct, taxa in the species (van Wyk et
al., 2007). A contemporary view is that C. fimbriata sensu stricto (s.s.) specifically
refers to the cause of black rot of sweet potato (Ipomoea batatas L.) on which it
was first described (Halsted and Fairchild, 1891). Other cryptic, monophyletic
lineages of C. fimbriata s.l. have been described as distinct species (Engelbrecht
and Harrington, 2005; Johnson et al., 2005; van Wyk et al., 2005, 2007), and more
will likely follow.
Two new Ceratocystis spp. have been described on mango in the Oman Gulf
region. Ceratocystis omanensis belongs to the Ceratocystis moniliformis Hedgc.
s.l. species complex (Al Subhi et al., 2006), which are typically not primary
pathogens. Ceratocystis omanensis is a minor pathogen on mango. The primary
sudden decline agent in Oman and Pakistan, C. fimbriata s.l., represents a
monophyletic lineage based on ITS, E-tubulin and translation elongation factor
(TEF) 1-D DNA sequence comparisons, and it has unique morphological
characteristics; it was renamed Ceratocystis manginecans M. van Wyk, A Al
Adawi and M.J. Wingf. sp. nov. (van Wyk et al., 2007).
On 2% malt extract agar (MEA), colonies of C. manginecans are greyish olive
and have a banana odour (van Wyk et al., 2007). Hyphae are smooth and
segmented (Fig. 8.26). Ascomatal bases are globose, black and (153–)192– 254(–
281) Pm in diameter; ascomatal necks are dark brown, lighter towards the apices
(514–)557–635(–673) Pm long, (25–)32–42(–48) Pm, wide at the
276 R.C. Ploetz and S. Freeman

(b) (e) (f)

(c)

(a) (d) (g)

Fig. 8.26. Microscopic features of Ceratocystis manginecans: (a) globose ascoma,


(b) divergent ostiolar hyphae, (c) hat-shaped ascospore, (d) segmented hypha,
(e) primary phialidic conidiophore with emerging cylindrical conidia, (f) secondary
conidiophore with emerging chain of barrel-shaped conidia, and (g) dematiaceous
chlamydospores and cylindrical- and barrel-shaped conidia. Scale bars: (a) = 100 Pm,
(b) = 20 Pm, (c) = 5 Pm, (d) = 20 Pm, (e) = 20 Pm, (f) = 20 Pm, (g) = 20
Pm. (Source: van Wyk et al., 2007).

base, and (14–)16–22(–26) Pm wide at the tip; and ostiolar hyphae are hya-line,
divergent and (42–)45–59(–69) Pm long. Asci are evanescent, and ascospores are
hyaline, hat-shaped, 3–4 Pm long, and 4–5 Pm wide without, and 7–8 Pm wide
within the sheath. Primary conidiophores are phialidic, lageniform, hyaline, (72–
)81–109(–144) Pm long, 5–7(–9) Pm wide at the base, 6–8(–9) Pm wide at the
broadest point, and 3–6 Pm wide at the tip. Secondary conidiophores are tube like,
flared at the mouth, short, hyaline, (59–)65– 77(–84) Pm long, 5–8 Pm wide at the
base and (5–)6–8 Pm wide at the tip. Primary conidia are hyaline, cylindrical, (15–
)23–29(–33) Pm long, and 3–6 Pm wide; and secondary conidia are hyaline, barrel
shaped, (8–)9–11(–12) Pm long, and 5–7(–8) Pm wide. Chlamydospores are
brown, thick-walled, glo-bose to subglobose, (11–)12–14 Pm long and 9–11(–12)
Pm wide.
Two isolates of C. fimbriata s.l. from mango in Brazil (CBS 114721 and CBS
600.70) have been compared to isolates of C. manginecans (van Wyk et al., 2005,
2007). They are similar to, but differ significantly from, C. manginecans. They
reside with C. manginecans in a clade that contains other New World
Foliar, Floral and Soilborne Diseases 277

species, Ceratocystis cacaofunesta and Ceratocystis platani. Research is needed to:


(i) examine additional isolates of C. fimbriata s.l. from mango in Brazil; (ii)
describe the putative, new species; (iii) determine whether C. manginecans is
present in Brazil; and (iv) clarify pathogenic variation in the agent(s) in Oman and
Brazil. At least two pathotypes of C. fimbriata s.l. are evident in Brazil (Rossetto et
al., 1996; Junqueira et al., 2002; Silveira et al., 2006).
Rossetto et al. (1996) evaluated 15 cultivars against two isolates of the
pathogen in Brazil. Eight-year-old trees were inoculated c.40 cm beneath branch
apices with IAC FITO 4905, which is pathogenic to ‘Jasmim’, and IAC FITO 334-
1, which is not. ‘São Quirino’, ‘Irwin’, ‘Edwards’ and ‘Van Dyke’ were resistant,
and ‘IAC 100 Bourbon’ was moderately resistant. ‘Glenn’, ‘Joe Welch’, ‘Zill’ and
‘Haden’ were susceptible, and ‘Kent’ responded as ‘Jas-mim’, resisting IAC FITO
334-1 and succumbing to IAC FITO 4905.

Epidemiology
Genotype has a profound impact on disease development, and severe epi-demics
occur wherever susceptible rootstocks and/or scions are used. Greater disease
develops when trees are stressed, although it is not clear whether this results from
an increased attraction of the vector to stressed trees or reduced disease resistance
in the host. The associated pathogens are moved easily in infected germplasm, the
route by which the diseases have spread in Brazil and probably to Oman. Pruning
implements also move the pathogen, and soil, once infested with chlamydospores
of the pathogen, can be a long-term reser-voir of inoculum. Insect dissemination
plays a particularly insidious role.
Beetles (Coleoptera: Scolytidae) are closely associated with seca in Brazil
(Batista, 1947; Viegas, 1960; Piza, 1966; Ribiero, 1980). Batista (1947) indicated
that Xyleborus affinis was the sole vector of D. recifiensis. In contrast, Ribiero
(1980) reported that Hypocryphalus mangiferae Stebbing was the primary vec-tor
of C. fimbriata s.l. (Fig. 8.27). It produced galleries in the cambium of affected
trees (Plate 47a), and was the only scolytid found on healthy, as well as diseased,
trees. Hypocryphalus mangiferae is also associated with the dis-eases in Oman and
Pakistan, where C. manginecans is recovered from the insect and trees are
commonly found with insect probing damage before dis-ease develops (Al Adawi
et al., 2006; van Wyk et al., 2007).
The interactions between H. mangiferae and the Ceratocystis pathogens of
mango are incompletely understood. In olfactometer tests in Brazil, H. mangiferae
was attracted to cultures of C. fimbriata s.l., and larvae of the insect were raised to
adulthood on the fungus (Ribiero, 1980). Several other species, many of which are
in the genus Xyleborus, were also associated with seca, but because they were
found only in diseased trees they appeared to be opportunistic feeders on C.
fimbriata s.l. rather than primary vectors. Although the sequence of events in
Brazil and the Oman Gulf has not been studied, it is probable that H. mangiferae
contaminates its body with these pathogens while feeding in diseased trees and
subsequently disseminates the pathogen to healthy trees.

Hypocryphalus mangiferae is thought to be native to some of the same areas in


southern Asia where mango evolved (Wood, 1982; Butani, 1993;
278 R.C. Ploetz and S. Freeman

Fig. 8.27. Hypocryphalus mangiferae, vector of the seca and sudden


decline pathogens (Photograph courtesy of R.C. Ploetz).

Atkinson and Peck, 1994; Mukherjee, 1997). Thus, the insect would have been
introduced into Brazil and would have been a new encounter, rather than
coevolved, relationship with C. fimbriata s.l. (van Wyk et al., 2007). In contrast, if
C. manginecans were introduced into Oman and Pakistan from Brazil, it may have
then established a relationship with a native insect. Although the available
information suggests that the H. mangiferae– Ceratocystis interactions on mango
were recent, opportunistic encounters in the New World, additional work is needed.

Management
Given the ease with which these pathogens are moved and their destructive impact,
preventing their dissemination to new areas must be a high priority. Pathogen-free
propagation material should be used whenever new plantings are established and
germplasm is moved. Clean pruning implements should be used in affected areas,
and should be frequently disinfested with bleach, formalin or other disinfestants
(Junqueira et al., 2002). Trees that have been killed by the disease must be
removed and destroyed as they are significant reservoirs of inoculum and infested
vectors. Where partially resistant culti-vars are grown, the removal and burning of
affected branches and treatment of the exposed branch stubs with Cu fungicides is
recommended (Ribeiro et al., 1995; Ribeiro, 1997).

Managing these diseases with fungicides on susceptible cultivars would be a


challenge. External applications of protectant or systemic fungicides would
probably be ineffective, given the internal, protected location of the pathogen.
Injecting fungicides, as is done to control Dutch elm disease, might be effective.
However, this would probably not be allowed where concerns
Foliar, Floral and Soilborne Diseases 279

with pesticide contamination of fruit exist. Treatment of germplasm collec-tions


and young, non-bearing trees might be the only situations in which fungicide
injection would be possible.
Genetic resistance offers the best hope for managing these diseases. Var-ious
levels of tolerance have been observed in Brazil and resistant clones have been
developed. However, pathogenic variation in the causal fungus in Brazil has
hindered progress (Rossetto et al., 1996; Junqueira et al., 2002; Sil-veira et al.,
2006). Although disease responses of some genotypes vary in dif-ferent production
areas in the country, ‘Manga D’agua’, ‘Pico’, ‘IAC 101’, ‘IAC 102’, ‘Edwards’,
‘Van Dyke’ and ‘Carabao’ resist two races of the patho-gen, and ‘Rosa’, ‘Sabina’,
‘Sao Quirino’, ‘Oliveira Neto’, ‘Jasmim’, ‘Sensation’, ‘Irwin’ and ‘Tommy Atkins’
are generally tolerant (Ribiero, 1997; Junqueira et al., 2002). ‘Kent’ and ‘Jasmim’
respond differentially (see above), and ‘Coquinho’, ‘Glenn’, ‘Joe Welch’, ‘Zill’
and ‘Haden’ are susceptible. Although ‘Espada’ is also reported to be tolerant, old
trees are frequently attacked. In commercial orchards, the disease on ‘Espada’ is
managed by grafting onto resistant rootstocks and pruning diseased branches.
Colosimo et al. (2000) worked with other scion cutivars, although in a single
location (and against a single pathotype?). They reported that ‘Oliveira’ was most
resistant, ‘Car-lota’, ‘Imperial’, ‘Extrema’ and ‘Pahiri’ had intermediate resistance,
and ‘Bourbon’ was most susceptible.

One must also recognize the impact of other diseases on different cultivars.
Carvalho et al. (2004) described two new cultivars, ‘IAC 103 Espada Vermelha’
and ‘IAC 109 Votupa’, which had moderate resistance to seca. ‘IAC 103 Espada
Vermelha’ also had moderate resistance to powdery mil-dew but was susceptible to
anthracnose. Both cultivars were susceptible to malformation.

Stigmina leafspot

Stigmina leafspot is caused by Stigmina mangiferae (Koorders) Ellis (synonym:


Cercospora mangiferae Koorders; a teleomorph for the fungus is not known). Lim
and Khoo (1985) indicated that the disease occurs on a range of cultivars, and is
most severe during rainy weather. Both young and old leaves are affected. Dark-
brown spots, 1–2 mm in diameter, are formed initially by the fungus. These can
enlarge and coalesce to 1 cm or larger, and are surrounded by conspicuous
chlorotic haloes that aid diagnosis of this disease. The fungus produces large, olive-
grey conidia, 30–60 × 3.5–5.0 Pm, usually on the lower leaf surface (Fig. 8.28).
Conidia are wider at the base than the apex, are straight to curved, have three to
seven septa, and are borne in subglobular, dark stromata that are 20–60 Pm in
diameter.
Although the fungus sporulates sparsely on artificial media, it produces
copious conidia on necrotic host tissue, especially in leaf litter. Thus, Lim and
Khoo (1985) suggested that removing such debris from orchards and burning it
would assist control efforts. Several different fungicides are effective (Lim and
Khoo, 1985).
280 R.C. Ploetz and S. Freeman

Fig. 8.28. Conidia of Stigmina mangiferae, cause of stigmina leaf spot (Source:
from CMI description no. 097).

8.3 Soilborne Diseases


Although soilborne diseases of mango are relatively less important than foliar and
floral diseases, they can cause significant damage to seedlings, nursery stock and
mature trees. In general, the pathogens that are involved are different from those
that cause problems above ground. Chemical
Foliar, Floral and Soilborne Diseases 281

management is indicated rarely for these diseases; sanitation and other cultural
measures are used most often.

Black root rot

Black root rot is reported to be an uncommon problem on young mango trees (Lim
and Khoo, 1985). Canopies of affected plants wilt suddenly and subse-quently
defoliate. Roots exhibit a water-soaked, blackened decay, and have an unpleasant,
putrid odour. Although black root rot is associated with pro-longed flooding, its
precise aetiology is not known. Several species of fungi have been recovered from
affected plants, including Fusarium solani, F. oxyspo-rum and L. theobromae, but
these were thought to be secondary colonizers of roots (Lim and Khoo, 1985).
Although mango is generally considered to be flood intolerant, its flood tolerance
is actually variable (Larson, 1991). Varia-tion in the responses of individual trees
in orchards is evident after flooding, and when potted plants are flooded
experimentally, they usually adapt by forming hypertrophied lenticels
(intumescence) (Larson et al., 1993). Plants that do not adapt in this manner
succumb fairly rapidly. Roots of the latter plants have symptoms of black root rot
(R.C. Ploetz, Homestead, Florida, 1988, personal observations). Although flood
tolerance is environmentally and biochemically complex (Larson et al., 1993),
some of the fungi reported by Lim and Khoo (1985) may interact with flood-
induced stress in this host to cause black root rot.

Nematode damage

Decline of mango trees due to nematodes has been reported in various regions
(Khan et al., 1971, 2005; McSorley et al., 1980; Anita and Chaubey, 2003).
Infestations occur in areas where warm temperatures and sandy, moist, well-
drained soils predominate (Ploetz et al., 1994). Many nematode species have been
recovered from mango roots, including: Helicotylenchus dihystera (Cobb) Sher,
Quinisulcius acutus (Allen) Siddiqi, Rotylenchulus reniformis Linford and
Oliveira, Criconemella sp., Pratylenchus brachyurus (Godfrey) Fil-ipjev and
Schuurmans Stekhoven, Xiphinema sp., Meloidogyne sp., Praty-lenchus sp. and
Hoplolaimus sp. However, only Hemicriconemoides mangiferae Siddiqi is
pathogenic (Powers and McSorley, 1994). Although high popula-tions of R.
reniformis are often found on mango trees, no correlation has been shown between
their density and tree health.
Populations of H. mangiferae vary according to soil moisture and tem-perature
(Khan et al., 1971). Soil moisture < 10% and > 30%, as well as tem-peratures <
15C and > 35C are detrimental to nematode survival and are likely to reduce
populations. In addition, tree age appears to be a significant factor, since H.
mangiferae is found more frequently on old (> 10 years) than young trees (< 3
years). Management is difficult and may depend on pre-plant chemical applications
plus cultural control measures (McSorley et al.,
282 R.C. Ploetz and S. Freeman

1981). Phenamiphos and 1,2-dibromo-3-chloropropane reduce levels of H.


mangiferae after planting; however, neither are registered for use in the USA.
Anita and Chaubey (2004) reported that high organic matter content in the
rhizosphere had a detrimental effect on nematode populations in the field.

During orchard establishment, nematode-free nursery stock should be used.


Since H. mangiferae is partially endoparasitic, it is moved easily to clean field
sites. The use of clean planting material in infested fields should also be avoided. If
such land must be used, soil fumigation prior to planting should be conducted.
Fruit yields may still be maintained if infected trees are well irrigated and
fertilized.

Phytophthora diseases

Phytophthora palmivora (E.E. Butler) (Oomycota) causes diseases of mango in


several areas. It caused wilt, crown rot, root rot and the death of nursery trees in
Arizona USA, the Philippines and Thailand (Kueprakone et al., 1986; Matheron
and Matejka, 1988; Tsao et al., 1994). Gumming and conspicuous bark lesions
develop above ground on these plants, whereas root and crown rots are evident at
or below the ground level. Crowded conditions and exces-sive irrigation and
rainfall exacerbate these diseases. Sanitation, the use of less-crowded conditions
and reduced irrigation would be beneficial.
Damage has also been recorded on the trunks of field-grown, mature trees in
the Ivory Coast (Lourd and Keuli, 1975), and on fruit in Australia, Malaysia and
West Africa (Turner, 1960 cited in Chee, 1969; Cooke, 2007). Mortality of trees is
not observed, but substantial stem cracking and bleed-ing does occur. The
symptoms that occur on fruit have not been recorded in Malaysia and West Africa,
but on ‘Calypso’ fruit in Australia, a firm chocolate-brown decay is produced that
has a sweet odour. Fruit isolates in Australia caused leaf blight and crown canker
on mango seedlings (Fig. 8.29).
Phytophthora palmivora has coenocytic hyphae, up to 7 Pm in diameter,
papil-late sporangia, 31–56.4 × 20.7–36.7 Pm, which germinate either directly with
germ tubes or indirectly with motile zoospores (Fig. 8.30) (Waterhouse, 1970;
Erwin and Ribiero, 1996). Zoospores are the primary infective propagule, and
require free water for movement. Phytophthora palmivora is heterothallic.
Antheridia are amphigynous and oogonia are spherical. Chlamydospores, c.35 Pm
in diameter, are also formed.
Recently, a Phytophthora sp. was isolated in Spain from mango trees that were
wilted, chlorotic and had sparse canopies and cracked bark (Zea-Bonilla et al.,
2007). On V8 agar, sporangia were semi-papillate, obovoid and 51 (28–52) × 36
(22–37) Pm. Paragynous antheridia, spherical oogonia and oospores of 28 (19–32)
Pm in diameter were produced homothalli-cally. Ribosomal DNA sequences
(ITS1, 5.8S rDNA and ITS2) (GenBank Accession No. AM235209) were
compared with those in GenBank; the closest match, 99% identity, was with
Phytophthora citricola, corroborating the above morphological identification (Fig.
8.31). An isolate deposited in
Foliar, Floral and Soilborne Diseases 283

(a) (b)

Fig. 8.29. Symptoms induced by Phytophthora palmivora after artificial inoculation of


(a) stems and (b) foliage of mango seedlings (Photographs courtesy of A.W. Cooke).

Fig. 8.30. (a) Sporangia, (b) oogonia with amphygynous antheridia and
oospores, and (c) chlamydospore of Phytophthora palmivora (Source: from CMI
description no. 831).
284 R.C. Ploetz and S. Freeman

Fig. 8.31. (a) Semipapillate sporangia and (b) oogonia of Phytophthora


citricola (Source: from CMI description no. 114).

the Spanish Type Culture Collection, CECT 20567, caused root rot on ‘Florida’
and lesions on leaves and stems of seedlings of ‘Gomera 3’.

Root rot and damping off

The oomycete, Pythium vexans de Bary, can cause root rot and wilt of seed-lings
(Lim and Khoo, 1985). In Malaysia, this pathogen caused seedling losses of up to
30% in nurseries. Symptoms included wilting of foliage, which initially becomes
pale green, but later develops necrotic patches. Roots develop a wet, blackened
necrosis that begins in fine roots and progresses to larger roots and the root collar.
Death of seedlings often occurs. Lim and Khoo (1985) indicated that
overcrowding, excessive moisture and the use of polybags favoured this disease.

Prakash and Singh (1980) reported that the basidiomycete Rhizoctonia solani
Kuhn (teleomorph: Thanatephorus cucumeris (Frank) Donk) caused root and
damping off of seedlings in India (Fig. 8.32). Affected tissues become soft, dark
brown or black, and seedlings may ultimately become completely girdled and
collapse. Mycelia and sclerotia of the pathogen form conspicu-ously on affected
tissues.

Sclerotium rot

This disease has been reported in Brazil (Almeida et al., 1979), India (Prakash and
Singh, 1976a) and the Philippines (Palo, 1933). The causal fungus, Sclero-tium
rolfsii Sacc. (teleomorph: Athelia rolfsii (Curzi) Tu and Kimbrough;
Foliar, Floral and Soilborne Diseases 285

(d)

(a)

(b)

20 μ

(c)

Fig. 8.32. (a) Sclerotial cells, (b) mycelium and (c) monilioid cells of Rhizoctonia
solani, and (d) basidia and basidiospores of its teleomorph, Thanatephorus
cucumeris (Source: from CMI description no. 406).

synonyms: Corticium rolfsii Curzi and Pellicularia rolftii E. West), produces


globular, brown sclerotia, 1.0–2.6 mm in diameter. Sclerotia are resilient struc-
tures that enable the pathogen to survive adverse environmental conditions.
Symptoms begin with the formation of felty white tufts of mycelium of the
pathogen around the base of seedlings. The fungus can girdle the entire stem to a
height of 5 cm or more above the soil line. It eventually forms conspicu-ous
sclerotia in high numbers. Ultimately, seedlings wilt and die. Seed may also rot
prior to germination. The disease is controlled via sanitation and the disinfestation
of seedbeds.

Verticillium wilt

Verticillium wilt of mango was first reported in Florida USA (Marlatt et al., 1970).
The disease was originally attributed to Verticillium albo-atrum Reinke and Berth.,
but this was before Verticillium dahliae Kleb. was recognized as a distinct species.
Since the causal fungus described by Marlatt et al. (1970) formed microsclerotia, it
is clear that V. dahliae was involved (Fig. 8.33).
Symptoms of the disease include a ‘firing’ and necrosis of leaves, usually in a
portion of the canopy. Sectoral development of the disease often does not progress
to other portions of the trees, which may recover. Killed leaves usu-ally remain
attached to the tree, and the xylem of affected branches is dis-coloured brown (Fig.
8.34). Verticillium wilt is relatively uncommon, and is
286 R.C. Ploetz and S. Freeman

(b)

10 μ

(a) (c) (d)

Fig. 8.33. (a) Verticilliate conidiophore, (b) phialospores and (c) immature and (d)
mature microsclerotia of Verticillium dahliae (Source: from CMI description no. 256).

found on land where susceptible vegetable crops (i.e. potato, tomato and aubergine)
were recently grown (Pohronezny and Marlatt, 1982). New mango orchards should
not be planted on such sites.

White root disease

Rigidoporus lignosus (Klotzsch) Imazeki, the basidiomycete that causes white root
disease, is a common soil inhabitant in the humid tropics of Africa and Asia
(Holliday, 1980). It has also been reported in the western hemisphere, but the
identity of the fungus there is unclear. Rigidoporus lignosus has a wide host range
on woody perennials, including rubber, the host on which the pathogen was first
reported (1904 in Malaysia). The significant losses in rub-ber plantations in the
eastern hemisphere have resulted in considerable research on this pathogen
(Nandris et al., 1987).
Rigidoporus lignosus produces white rhizomorphs on the surfaces of roots and
root crowns that later darken to a yellowish and then reddish colour (Lim and
Khoo, 1985; Nandris et al., 1987). The leading edge of the rhizo-morph is well
defined and seldom appears above ground. It undergoes a morphogenic change to
produce infectious hyphae that penetrate the host epidermis and subsequently
degrade wood. Rigidoporus lignosus produces a non-differentiated white rot that
affects lignin in host cell walls.
Foliar, Floral and Soilborne Diseases 287

Fig. 8.34. Vascular discoloration caused by V. dahliae (Photograph courtesy


of R.C. Ploetz).

The fungus is most damaging on mango if orchards are established in old


rubber plantations or newly cleared jungle sites (Lim and Khoo, 1985). Previously
colonized stumps and debris from rubber and other hosts are pri-mary sources of
inoculum. Orange-yellow, bracket-like sporophores are pro-duced during the rainy
season on the root collar, trunk or exposed roots (Fig. 8.35). Basidiospores are
viable, but may play a secondary role in dis-seminating the disease. Rhizomorphs
are more significant epidemiologically, since they grow rapidly and can advance
great distances in soil in the absence of woody substrates.
288 R.C. Ploetz and S. Freeman

(a) (b)

10 μ (c)
20
μ (d)

Fig. 8.35. (a) Upper and (b) lower surface of sporophore, (c) basidiospores, and (d) hyphae
of Rigidoporus lignosus (Source: from CMI description no. 198).

White root disease is managed by eliminating or avoiding colonized woody


debris when new orchards are established (Lim and Khoo, 1985). Unfortunately,
this is difficult and often impractical. Alternative measures include: (i) treating
planting holes with S to promote the growth of antago-nistic microorganisms; (ii)
treating stumps after clearing operations with chemicals that discourage their
colonization by basidiospores; and (iii) estab-lishment of leguminous cover crops
that promote growth of the pathogen and the eventual exhaustion of its energy
reserves.

8.4 Conclusions
As mango production continues to increase in different regions, so will the scope
and types of disease problems. Although new fungicides and bacteri-cides will be
developed in the future, it is probable that reliance on these tools will diminish. In
recent years, the regulation of pesticides has increased while the number of new
compounds that have been registered for use has decreased. Since it appears certain
that this trend will continue, the problems posed by diseases must be solved
increasingly with alternative disease control strategies.
Foliar, Floral and Soilborne Diseases 289

A willingness among producers to utilize new technologies will play a role in


this process. New methods to detect important pathogens have been developed
(Zheng and Ploetz, 2002), but more work is needed. Effective detection protocols,
especially for those pathogens that can colonize host tis-sues without causing
symptoms, could be used to interdict important, exotic pathogens and identify
pathogen-free propagation materials. Detection pro-tocols could also indirectly
assist disease control efforts through their use in epidemiological studies, research
on disease resistance, or to clarify portions of disease cycles.

Acknowledgements
The authors thank Francisco Cazorla for comments on apical necrosis; Ber-nard
Slippers and Mike Wingfield for comments on the decline section; Syl-via
Fernandez, John Leslie, Christiano Lima, Kerry O’Donnell and Gerardo Rodriquez
for information on malformation; Ali Obaid Al-Adawi and Mike Wingfield for
comments on seca and sudden decline; and Robert Knight for translating
Portuguese articles on seca. The following individuals are thanked for pictures:
Francisco Cazorla, T.-K. Lim, Oliver Pruvost, Carolyn Cohen, Suzanne Bullock,
Efrat Gamliel-Atinsky, Eric Palevsky and Tony Cooke. Kevin Hyde, editor of
Fungal Diversity, is thanked for permission to use micrographs of Ceratocystis
manginecans.

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9 Physiological Disorders

V. Galán Saúco
Instituto Canario de Investigaciones Agrarias, Tenerife (Canary Islands), Spain

9.1 Introduction 303


9.2 Internal Fruit Breakdown 304
Symptoms 305
Histology 305
Altered physiology 306
Causes 306
Cultivar susceptibility 307
Control 308
9.3 Lumpy Tissue 309
9.4 Ricey Tissue 310
9.5 Fruit Cracking 310
9.6 Black Tip Disorder 310
9.7 Lenticel Spot 311
9.8 Other Fruit Disorders 311
9.9 Stem Disorders 312

9.1 Introduction
Physiological disorders can be defined as ailments that have not been caused by
infecting organisms. Unlike mango malformation, for example, which has been
considered as a physiological disorder in the past (IBPGR, 1989) but has a
phytopathological origin, true physiological disorders cannot be transmit-ted from
plant to plant, mechanically or by insect bites. For the most part, they are a result of
some form of physical damage or of an altered physiology of the tree or fruit. In
the particular case of fruit disorders – the most preva-lent and important
physiological disorders of mango – according to Subra-manyan et al. (1971), they
are essentially the result of imbalances in metabolism induced by some factor or
factors in the pre- or postharvest environment that leads to cell collapse and,
typically, the appearance of waterlogged or
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 303
304 V. Galán Saúco

brown areas on some part of the fruit. The complexity of events leading to the
occurrence of physiological disorders makes it more difficult to pinpoint the causal
factors than is the case of symptoms caused by pathogens or pests.
Physiological disorders occur relatively frequently in many fruit and vegetable
species. Some well-known examples include bitter pit, water core and internal
breakdown in apple (Atkinson et al., 1980) and in watermelon (Singh et al., 1975;
Cirulli and Ciccarese, 1981), blossom end rot in tomato and pepper (Winsor and
Adams, 1987), mesocarp discoloration and pulp spot in avocado (Van Lelyveld et
al., 1984; Bower and Cutting, 1987) and yellow pulp in banana (Melin and Aubert,
1973).
Although physiological disorders of annual crops and temperate fruits have
been studied extensively, tropical fruits have only recently been stud-ied in order to
understand and control physiological disorders. This is largely due to the rapid
expansion of these crops during the past few decades.
Preharvest factors that predispose mango fruit to physiological disorders
include growing location, orchard condition, tree nutrition and condition at the time
of harvest. Postharvest storage conditions, such as temperature, oxygen and carbon
dioxide levels, packaging and surface coating treatments, are also potential
contributing factors to the occurrence of these disorders (Subramanyan et al.,
1971).
Evidence of specific metabolic changes that occur in mango fruits that express
this type of disorder is scarce. Probably due to this lack of knowl-edge, disorders
are usually named for the associated environmental factor, for example chilling
injury, or by the altered appearance that the physiologi-cal disorder confers to the
fruit. Clear examples of the latter are internal fruit breakdown, which is by far the
most prevalent physiological disorder in mango, lenticel spot, fruit cracking and
black tip disorder.

9.2 Internal Fruit Breakdown


Most opinion (Winston, 1984; Schaffer, 1994) is that several physiological dis-
orders currently listed as separate disorders are in fact different degrees or aspects
of the same problem, internal fruit breakdown (IFB). These include tip pulp, soft
nose (i.e. waterlogging of the flesh near the distal end), jelly seed (i.e. overripe
flesh around the seed, surrounded by firm flesh), stem end breakdown (i.e. an open
cavity in the pulp at the stem end), spongy tissue (i.e. areas of the flesh that appear
spongy and have a greyish black discolor-ation), soft centre and soft flesh. This
aggregation is still open to some debate. Raymond et al. (1998) asserted that
temporal and spatial differences in symp-tom development within the fruit have
been found between soft nose, jelly seed and stem end cavity, validating their
return to separate disorder status, but they recognized that no major microscopic
differences could be found. None the less, irrespective of the disparity of
symptoms, all of these disor-ders have a common denominator: calcium imbalance.
With this defining criterion, IFB would also include the physiopathy described in
Malaysia as yeasty or insidious fruit rot, which shows a marked interdependence
with
Physiological Disorders 305

calcium (Lim and Khoo, 1985) and also what Velasco Cárdenas (1974) calls
‘ablandamiento del pico’ (literally peak softening).
Cheema and Dani (1934) first reported an IFB disorder, but perhaps the most
comprehensive work published on physiological disorders of the mango fruit was
that of Malo and Campbell (1978), who described the different degrees of tissue
decomposition in fruit of ‘Tommy Atkins’. The comprehen-sive work of
Wainwright and Burbage (1989) reviews the symptomatol-ogy, chemical changes
and causes of IFB, illustrating its occurrence in many cultivars in virtually all the
producing areas of the world. Losses from IFB vary geographically and also among
cultivars, and can affect 100% of the harvested fruit (Subramanyan et al., 1971;
Malo and Campbell, 1978; Bro-drick and Thord-Gray, 1982; Galán Saúco et al.,
1984; Santos Filho et al., 2002; Cracknell Torres et al., 2003a).

Symptoms

Symptoms usually begin to appear early during fruit development and advance
rapidly, eventually making the fruit inedible (see Plates 48–52). The process
commences while the fruit is still hanging on the panicle, with an interruption
occurring in the vascular tissues of the peduncle and endocarp, and usually
followed by the formation of a cavity close to the funiculum (stem end
breakdown). In advanced stages, the affected tissues around the cavity become
grey or blackish. As the disorder progresses, the proximal end of the fruit becomes
mushy to the touch (soft nose). In some cases, a yellow-ing of the skin between the
apex and the stigmatic end of the fruit can also occur. The fibre surrounding the
endocarp may fully disintegrate and in severe cases an accumulation of a
transparent, gelatinous substance devel-ops around the seed (jelly seed). The
affected mesocarp matures more rap-idly than the healthy flesh and acquires a
characteristic deeper yellow colour; the affected tissue may be so extensive that
greyish, watery tissues appear over the whole mesocarp (spongy tissue) (Malo and
Campbell, 1978; Winston, 1984; Joshi and Roy, 1985; Wainwright and Burbage,
1989; Katrodia and Chuva, 1993). The absence of latex and the lack of firmness in
the proxi-mal end of the fruit at harvest can be clear signs of the disorder (Chaplin,
1986), although the lack of latex alone has also been observed in black tip disorder
(see below).

Histology

Histological differences have been found between fruits of different cultivars


affected by IFB. For example, in the very susceptible ‘Tommy Atkins’ the xylem
elements in the mesocarp and the mesocarp cells are all highly deteriorated. In the
case of a less sensitive cultivar (i.e. ‘Lippens’) the affected mesocarp cells have
intact cell walls, even though the tissue is obviously affected, showing a translucent
mesocarp (Cracknell Torres and Galán Saúco, 2003).
306 V. Galán Saúco

Altered physiology

A reduction in firmness, total soluble solids, total pectin and pectinase activ-ity was
observed by these researchers in the affected mesocarp tissue, con-firming the
results of Van Lelyveld and Smith (1979) and Roe and Bruemmer (1981), both of
whom also observed greater activity of both pectinase and malic enzyme in affected
pulp of ‘Alphonso’. On the other hand, the higher level of nitrogen, the lower levels
of calcium, and the higher nitrogen to cal-cium ratio observed by Cracknell Torres
and Galán Saúco (2003) in the affected tissue of ‘Tommy Atkins’ and ‘Lippens’ is
in agreement with the studies of Young (1960) and Young and Miner (1961) but in
conflict with results of Krishnamurthy (1981), who found no differences among
calcium levels between affected and unaffected mesocarp tissues. Other differences
in chemical composition have been detected in affected and unaffected tis-sues
(Subramanyan et al., 1971; Patkar et al., 1983; Nuevo et al., 1984; Gupta et al.,
1985) and further research should be devoted to ascertain the true nature of this
disorder.

Causes

Despite many attempts to isolate a pathogen from affected tissues, no patho-gen has
yet been linked with IFB. In Malaysia, spongy tissue has been found in
‘Harumanis’ fruit which has been affected by the fruit-piercing moth Othreis spp.
and in ‘Fan Siamese’ fruit, which has been associated with dam-age caused by
phytotoxic sprays of some fungicides (Lim and Khoo, 1985). Postharvest vapour
heat treatment at 46C for 10 min has also been reported to induce IFB in
‘Carabao’ fruit (Esquerra et al., 1990).
Several environmental factors have been linked to IFB. For example, in India
the disorder appears to be more prevalent in coastal areas (Subraman-yan et al.,
1971). Gunjate et al. (1982) observed that it is associated with fruit that remains
exposed to the sun following harvest. IFB has also been attributed to heat
convection from the soil at air temperatures around 55C (Katrodia and Rane,
1989). In several experiments with the cultivar ‘Alphonso’ in India, a positive
correlation was observed between IFB inci-dence and relative density. Mangoes
that were harvested with a relative den-sity between 1.00 and 1.20 showed no IFB,
while fruits greater than 1.2 had a 30% incidence of IFB (Krishnamurthy, 1980).
Fruit weight also seems to be an important factor at least in the case of some
cultivars like ‘Alphonso’ in India (Subramanyan et al., 1971) and in Spain with
‘Sensation’ (Hermoso et al., 1996), with a positive correlation in both cases
between fruit weight and incidence of IFB.

Cultural practices such as excessive irrigation have also been considered to be


possible causes of IFB. Katrodia (1988) observed a higher incidence of IFB
following periods of rainfall just prior to harvesting of ‘Alphonso’ in India;
however, trials done elsewhere with ‘Tommy Atkins’ (Malo and Camp-bell, 1978)
or ‘Sensation’ (Farré and Hermoso, 1993) showed no correlation
Physiological Disorders 307

between excessive irrigation and IFB. Furthermore, in Malaysia the incidence of


IFB (insidious or yeasty fruit rot) has been reported to be higher during the drier
months of the year (Lim and Khoo, 1985).
Most observations and field studies associate IFB with calcium deficiency,
which is also the main cause of similar disorders in other fruits (Bangerth, 1979;
Winsor and Adams, 1987). In general, low levels of leaf calcium at har-vest
coincide with high IFB incidence. In Florida, IFB appears to occur more in acid
and sandy soils with low calcium content than in basic, calcareous soils (Young,
1957; Schaffer, 1994), but in ‘Harumanis’ yeasty or insidious fruit rot occurs
irrespective of various types of soils, ranging from acidic to alkaline. It has been
suggested that the degree of IFB is aggravated in the presence of excess nitrogen
fertilization (Young, 1960; Young and Miner, 1961; Young et al., 1962, 1965).

The direct relationship between IFB and calcium and nitrogen content has
been confirmed by analysis of affected and non-affected fruits of orchard-grown
‘Harumanis’ (Ahmad Tarmizi et al., 1993), and in soilless cul-tivation trials
involving ‘Tommy Atkins’ (Cracknell Torres et al., 2003b). In the latter study,
positive correlations were observed between mesocarpic nitrogen levels and IFB
and between the nitrogen:calcium ratio and IFB, but there was a negative
relationship between calcium and IFB.

Cultivar susceptibility

Although IFB has been observed in at least 65 cultivars in 23 countries (Galán


Saúco, 1999), it is clear that varietal differences play a role in the intensity and
expression of the symptoms, as these have been described in different degrees of
intensity according to countries and cultivars.
According to Young (1957), cultivars of Indian origin and their offspring show
a higher incidence of IFB, for example ‘Alphonso’ whose susceptibility has been
well documented (Subramanyan et al., 1971; Gunjate et al., 1979, 1982;
Krishnamurthy, 1980, 1981; Patkar et al., 1983; Gupta et al., 1985; Lim and Khoo,
1985; Katrodia and Rane, 1989; Lad et al., 1992). There is, however, conflicting
evidence; some Indian cultivars, for example ‘Rajapuri’ (Katrodia, 1988),
‘Banganapalli’, ‘Kalapardy’, ‘Janradhan Pasand’ (Iyer and Subraman-yan, 1992),
‘Pairi’, ‘Kesar’ and ‘ Neelum’ (Lad et al., 1992), rarely demonstrate these
disorders. All Florida cultivars are affected due to their Indian parent-age
(Schaffer, 1994); ‘Haden’, ‘Kent’, ‘Keitt’, ‘Sensation’, ‘Tommy Atkins’ and ‘Van
Dyke’ are repeatedly mentioned as particularly susceptible (De Larous-silhe, 1980;
Galán Saúco and Fernández Galván, 1987; Campbell, 1992; Oost-huyse, 1993;
Knight, 1997; Galán Saúco, 1999; Cracknell Torres et al., 2003a). IFB problems
are also cited for the Malaysian cultivar ‘Harumanis’ or ‘Aru-manis’ (Lim and
Khoo, 1985; Tengku Ab.Malik, 1992a, b; Ahmad Tarmizi et al., 1993).

IFB incidence in some cultivars is so low as to be virtually non-existent,


particularly the polyembryonic and fibrous-fleshed types like ‘Turpentine’ in
Florida (Malo and Campbell, 1978), ‘Turpentine’, ‘Coquinho’ and ‘Espada’
308 V. Galán Saúco

in Brazil (Ferreira, 1989), and ‘Gomera 1’ in the Canary Islands (Cracknell Tor-res
et al., 2003a). There is at least one documented case of IFB in Australia involving
fruit of the polyembryonic ‘Kamerunga White’ (Winston, 1984).
Cultivar-related differences with respect to IFB incidence have been described
by Cracknell Torres et al. (2003a). They studied 28 cultivars and observed that
‘Edward’, ‘Gomera 1’, ‘Irwin’, ‘Valencia Pride’, ‘Mabroka’, ‘Ah Ping’ and ‘Heidi’
were almost free of this disorder. Modern Israeli cultivars, such as ‘Shelly’ and
‘Tango’ have also exhibited low frequency of IFB (Lavi et al., 1997a, b).
Observations of the extent of spongy tissue in various hybrids and selfed progenies
of ‘Alphonso’ clearly indicate that this disorder is genetically determined, and
‘Alphonso’ is apparently homozygous recessive for this character (Iyer and
Subramanyan, 1992).

Control

Despite the close relationship that exists between nutrition and IFB, very few
practical recommendations have been proposed for controlling this problem. There
is general agreement with the observations of Young (1960) and Young and Miner
(1961) that maintaining a low leaf content of nitrogen (<1.2%) and a calcium level
≥ 2.5% minimizes the amount of affected fruits. High nitrogen levels enhance
vegetative growth and rapid fruit development. Calcium moves only in the xylem.
Calcium concentration in organs such as fruits, which have a low rate of
transpiration and which are preferably supplied via the phloem, can result in
calcium levels falling below the critical level required for membrane integrity and
cell wall stability (Marschner, 1995). The antagonistic effect of nitrogen
fertilization, especially when ammonium salts are applied, and calcium uptake and
accumulation on leaves and fruits, is well documented in many fruit species (Shear,
1975; Lewis et al., 1977; Lud-ders, 1979) and has been clearly demonstrated for
mangoes (Young et al., 1962, 1965).

Lime application has also been recommended to increase the cation exchange
percentage to values 7.0 (Ferreira, 1989). According to Schaffer (1994), IFB has
been corrected in ‘Keitt’ by calcium application either to the soil as calcium
carbonate (CaCO3) or as a foliar calcium nitrate (Ca (NO3)2) spray.
Cultural practices such as mulching have been cited as being beneficial for
reducing IFB. Mulching lowers the soil temperature and thereby miti-gates the
reflected or rising heat that eventually affects the fruit (Katrodia, 1988; Lad et al.,
1992). The accompanying reduction in the transpiration stream of the tree and the
consequent minor mobilization of calcium to the fruit (Bangerth, 1979) may also be
important.
Certain rootstocks can improve calcium uptake and accumulation and provides
an important new procedure for controlling IFB. Fieldwork was initiated in
Malaysia at the end of the last century (Tengku Ab.Malik, 1996), but further
studies remain unreported.
Early harvesting at the green-ripe stage has been recommended as a measure to
reduce IFB (Young, 1957; Young and Miner, 1961; Galán Saúco
Physiological Disorders 309

et al., 1984; Winston, 1984), but for some cultivars this practice results in lower
quality fruits. The elimination at harvesting of fruits that do not exude latex (Mead
and Winston, 1991) and avoiding the exposure of fruit to direct sunlight during
harvest (Gunjate et al., 1982) have been recommended as prac-tical methods to
reduce the incidence of IFB in mangoes in the market. Accord-ing to Santos Filho
et al. (2002), postharvest hydrothermal treatments should be avoided in fruits
coming from orchards with a history of IFB incidence.
Preharvest dips of 0.5–2.0% of calcium chloride (CaCl2), applied from the
second month after fruit set until harvest, can increase calcium content in the fruit,
and has been reported to be an effective control measure for spongy tissue in fruit
of ‘Alphonso’ (Gunjate et al., 1979). In contrast, foliar sprays with calcium have
been reported to be ineffective for increasing leaf cal-cium content (McKenzie,
1995), and IFB has even been increased by applica-tion of calcium, which indicates
that a nutrition imbalance within the fruit was induced (Oosthuyse, 1997).

An integrated approach to control insidious fruit rot incidence in ‘Haru-manis’


has been developed in Malaysia. This includes maintaining soil pH close to 6.0,
late pruning, application of calcium sprays (2% CaCl2) to the fruits 4–6 weeks after
flowering, supplemental irrigation, monitoring nitro-gen and potassium fertilization
and anticipated harvesting (Tengku Ab. Malik, 1992a, b). Galán Saúco (1999)
suggested the following general inte-grated management measures: avoid planting
sensitive cultivars (i.e. ‘Sensa-tion’, ‘Tommy Atkins’, etc.); use appropriate
rootstocks (i.e. ‘Tangkai Panjang’ or others with high calcium absorption capacity);
harvest at the green-ripe stage; use an appropriate fertilization programme that is
rich in calcium and poor in nitrogen; avoid excessive irrigation close to fruit
maturity; and mulch soil in regions with hot summers.

There are notable differences in IFB resistance among cultivars. Because of


the widespread incidence of this problem, breeding for rootstocks and sci-ons for
IFB resistance is critical for resolving this problem (Iyer and Degani, 1997). Iyer
and Subramanyan (1992) have described the progress of breeding studies to
overcome IFB in India, obtaining hybrids of ‘Alphonso’ by other Indian cultivars
free from spongy tissue.
Biotechnological approaches also merit attention. Litz and Lavi (1997), based
on work done in tomatoes to control jelly seed (Smith et al., 1990; Oeller et al.,
1991), have suggested that control of IFB could be achieved by manipulating
ripening using the antisense strategy, either by blocking ethyl-ene biosynthesis or
with polygalacturonase, but no work has been reported in mango so far to test this
suggestion.

9.3 Lumpy Tissue


This disorder, of unknown etiology, occurs frequently in Thailand, where it occurs
mainly in the cultivars ‘Namta-an’, ‘Parg-grabong’ and ‘Pinsen Dang’, and in the
Philippines in ‘Pico’. The disorder becomes apparent as fruit ripening begins, with
indentations or scoring of the peel, which become
310 V. Galán Saúco

increasingly marked as ripening progresses. Opened fruits display a meso-carp full


of white lumps of intact, starch-filled cells (Lizada et al., 1984).

9.4 Ricey Tissue


This disorder is of unknown origin, but is common in ‘Carabao’ in the Philip-pines.
No external symptoms are visible, but internal symptoms are found in ripening fruit
and even at physiological maturity, and consist of small lesions. The lesions
resemble grains of rice in size and aspect (hence the name) on the mesocarp,
surrounded by cotton wool-like tissue. Except for the small changes in the texture
of the affected area, no other organoleptic changes have been detected (Lizada et
al., 1984).

9.5 Fruit Cracking


The only symptom of this disorder is that the fruit crack suddenly while still on the
tree, the cracks forming clean, knifelike cuts breaking the skin and into the flesh.
Secondary infections by Colletotrichum gloeosporioides (the causal agent of
anthracnose) or other fungal pathogens can occur. The cause of fruit cracking
appears to be related to water tension differences in the skin during periods of high
relative humidity and it occurs when trees are heavily irrigated after a prolonged
dry period, and if heavy rains are intermixed with dry spells. Frequently almost all
of the fruits on the tree are affected, although those near maturity are the most
susceptible. The incidence of fruit cracking seems to be higher in cultivars with
little or no fibre. Control measures may include regular watering and mulching, but
the ultimate solution may lie in breeding as in IFB (Lim and Khoo, 1985).

Another cause for fruit splitting is infection by bacterial black spot


(Xanthomonas campestris), which is considered to be a major cause of fruit
splitting. Consequently, prevention is the only solution to the problem by ensuring
adequate wind protection, which is also beneficial if the damage is caused by
differences in water tension of the skin cells, and/or appropriate chemical sprays to
prevent infection (Meurant et al., 1997).

9.6 Black Tip Disorder


This disorder is widespread in India (Prakash and Srivastava, 1987). Ram (1988)
indicated that it was first described in 1909. Three disorders frequently cited prior
to 1958, namely taper tip, tip pulp and girdle necrosis, are in fact variants of black
tip. The first symptom of this disorder is the etiolation and yellowing of the distal
end of fruit, which then turns black and hardens, causing the fruit to ripen
prematurely and making it unmarketable (Plate 53). The vascular bundles in the
pedicel may turn brown and decay. Affected fruits do not secrete latex at harvest.
Physiological Disorders 311

Outside India, the only known occurrence of black tip is from the Guang-dong
province of China (Zhang et al., 1995). In both India and China, the disorder
occurs in areas close to brick-making kilns, particularly where trees are more
exposed to fume-laden winds coming from the brick works. Although not fully
assessed, it seems that fluorine is the causal agent of this disorder (Zhang et al.,
1995), although a dry hot climate may enhance the effect of gases. Differences in
susceptibility among Indian cultivars have been reported by Ram (1988), who also
suggested the possibility of varietal selection for orchards in the vicinity of brick
kilns. Majumder and Sharma (1985) recom-mend a minimum distance between
kiln and trees of 1.6 km in the path of prevailing winds and 0.8 km on the other
sides, as well as recommending prevention by spraying three times with borax
(0.6%) and caustic soda (0.8%), for example prior to flowering, during flowering
and at fruit set.

9.7 Lenticel Spot


This disorder is characterized by the development of small spots of corky tissue in
the skin lenticels that darken as the fruit changes colour during ripening, which
makes the fruit unmarketable. The causes are not entirely uncertain, but it is most
often associated with incorrect postharvest practices, including low storage
temperatures, high humidity of the storage atmos-phere, excessive immersion time
in postharvest dips and excessive detergent in wash water (Oosthuyse, 1993;
Meurant et al., 1997). Some growers link it to other causes, such as delayed
harvesting or wet conditions during picking. Differences among cultivars with
respect to the severity of this problem have been observed. It is very common in
the late harvest of ‘Keitt’ in the subtrop-ics at the beginning of winter under cooler
and wetter conditions.

9.8 Other Fruit Disorders


Spots and marks on green fruit can have several different causes: cold dam-age or
chilling injury, usually seen as small raised brown or discoloured spots on the skin,
or pitting surrounded by sunken areas affecting the epidermis and the endocarp
(Lizada et al., 1984). Storage temperatures below 13C have been identified as
major causes of chilling injury, and must be avoided (Meurant et al., 1997). High
temperatures or sudden exposure to sunlight can cause sunburn, which is
characterized by bleached or yellow patches on the skin, which may become
leathery, yellow-brown to black, and lightly sunken. Trees should be well irrigated
during fruit filling and harvested fruit must always be kept (even briefly) in full
shade.
Wind damage, resulting from prolonged rubbing of the fruit against leaves or
dead twigs, requires adequate windbreaks and timely pruning of dead wood. If hail
damage is likely to be a frequent problem, the whole orchard will need to be
protected by nets. Damage caused during harvest, storage and transport, can only
be reduced by careful handling. Sapburn,
312 V. Galán Saúco

caused by prolonged contact with latex leaking onto the skin from a cut stem, can
be minimized by suitable harvesting and packing operations. High levels of carbon
dioxide (CO2) during storage provoke the development of off-flavours and small
internal lesions, as well as exudating brown tissues (Chaplin, 1986) and inhibition
of ripening (Thompson, 1971).
A disorder named pulpa negra, literally ‘black flesh’, which speaks quite
clearly as to the symptoms, has been reported in Mexico, especially in ‘Haden’
(Mora et al., 1998). Although the problem was detected in fruits that had been
stored at 13C for more than 20 days, it is not entirely clear that low tempera-ture is
the only cause as it also occurred in fruits reportedly stored only at room
temperature.

9.9 Stem Disorders


While some slight cracking of the bark around the trunk may be normal, some
cultivars, like ‘Manzanillo’ or ‘Gomera 4’, are prone to fissuring, with an unusually
high degree of suberification. This disorder is common in the Canary Islands,
Spain, and it is more pronounced on the sides of the trunk that are wetted by
sprinklers than on the less exposed sides (Fernández Galván and Galán Saúco,
unpublished). This phenomenon of stem cracking has been also associated with bad
orchard management, particularly with bad irrigation practices (Mora et al., 1998).
Some rootstocks, including ‘Gomera 1’ in the Canary Islands, frequently exhibit
gall-like growths along the branches, thought to be linked to climatic factors which
facilitate swelling of lateral buds without being enough to provoke flushing and the
corresponding shoot elongation.

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calci-um fertilization on ‘Kent’ mangoes in deep, acid, and sandy soil.
Proceedings of the Florida State Horticultural Society 75, 364–371.
Young, T.W., Koo, R.C.J. and Miner, J.T. (1965) Fertilizer trials with ‘Kent’
mangoes. Pro-ceedings of the Florida State Horticultural Society 78, 369–375.
Zhang, Ch., Huang, H. and Kuang, Y. (1995) A study of the cause of the mango
black tip disorder. Scientia Horticulturae 64, 49–54.
10 Pests
J.E. Peña,1 M. Aluja2 and M. Wysoki3
1
University of Florida, Florida, USA
2
Instituto de Ecología, AC, Xalapa, Veracruz, Mexico
3
Institute of Plant Protection, Bet Dagan, Israel

10.1 Introduction 317


10.2 Mango Fruit Pests 318
Fruit flies 318
Fruit fly control: brief overview 322
Mango seed weevils 328
Mango seed borer (Lepidoptera: Pyralidae) 331
Fruitspotting bugs (Hemiptera; Coreidae) 332
Thrips 333
Blossom pests 334
Pests of buds and leaves 338
Pests of trunks, twigs and roots 344
10.3 Discussion 345

10.1 Introduction
Mango, like most fruit trees, is usually attacked by two or three key pests, several
secondary pests and by a large number of occasional pests in local-ized areas
where it is grown. Worldwide lists of pests of mango have been published by
Laroussilhe (1980), Tandon and Verghese (1985) and Veeresh (1989). The pests of
mango in India (Srivastava, 1998; Anonymous, 2006), Australia (Anonymous,
1989), Pakistan (Mohyuddin, 1981), Israel (Wysoki et al., 1993; Swirski et al.,
2002), the USA (Peña, 1993), West Africa (Vannière et al., 2004), Brazil (Assis
and Rabelo, 2005), Central America (Coto et al., 1995) and Puerto Rico (Martorell,
1975) have also been described. Some publications contain check lists of mango
pests and most contain details of life histories and control of mango pests (Morin,
1967; Golez, 1991; Murray, 1991).
Of c.322 species of insects and mites that have been recorded as minor and
major pests of mango, 127 (39%) are foliage feeders, 87 (27%) are fruit feeders, 36
(12%) feed on the inflorescence, 33 (10%) inhabit buds and 39
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 317
318 J.E. Peña et al.

(12%) feed on branches, the trunk and roots. The four key pests (fruit flies, seed
weevils, tree borers and mango hoppers) require annual control mea-sures.
Secondary pests generally occur at sub-economic levels, but can become serious
pests as a result of changes in cultural practices and cultivar or because of
indiscriminate use of pesticides against a key pest. For exam-ple, Mohyuddin and
Mahmood (1993) reported that scale insects became serious pests following non-
judicious use of insecticides against fruit flies. Similarly, mites, Oligonychus spp.,
are secondary pests of mango, which can become serious because of human
intervention. Occasional or incidental pests also can cause economic damage only
in localized areas at certain times. The majority of pests reported here are of this
category.

10.2 Mango Fruit Pests


With current world emphasis on quality fruit for local consumption and export,
insects that blemish fruit by feeding, scratching or ovipositing in the pulp or seed
can cause high losses. Only fruit flies, seed weevils and lepi-dopterous larvae
actually penetrate the fruit pulp and seed. The feeding of other pests (e.g. Othreis
materna (L.), Gonodonta pyrgo (Cram.), Gonodonta clo-tilda (Stoll) and
Leptoglossus stigmai (Herbest)) often extends only into the pulp of ripening
mangoes (Angeles and Requena, 1966).

Fruit flies

Most mango-producing countries are in fruit-fly infested areas, and produc-ers


suffer significant direct economic losses (larval feeding renders fruit unmarketable)
and indirect economic losses (quarantine restrictions hinder-ing exports), resulting
from the presence of fruit flies (Hill, 1975; Umeya and Hirao, 1975; Anonymous,
1987; Yee, 1987; Singh, 1991; Aluja, 1993; Aluja et al., 1996; Vannière et al.,
2004; Aluja and Mangan, 2008). Few insects have a greater impact on international
marketing and world trade in agricultural produce than tephritid fruit flies
(Hendrichs, 1996; Aluja and Mangan, 2008). Approximately 60 species of fruit
flies are reported to attack mango and a related species, Mangifera foetida (White
and Elson-Harris, 1992; Aluja et al., 1996; Malavasi and Zucchi, 2000; Clarke et
al., 2001, 2005; Norrbom, 2004; Vayssières et al., 2005). Fruit flies attacking
mango belong to the genera Anastrepha (c.12 species), Bactrocera (c.33 species),
Ceratitis (eight species) and Dirioxa (two species) (White and Elson-Harris, 1992;
Vayssières and Kala-bane, 2000; Lux et al., 2003; Norrbom, 2004; Vayssières et
al., 2004, 2005; Sec-retariat of the Pacific Community, 2005). Some of these
species are referred to as the ‘mango fruit fly’: Anastrepha obliqua Macquart,
Bactrocera frauenfeldi (Shiner), Ceratitis cosyra (Walker) (Aluja, 1993; Leblanc
and Allwood, 1997; Lux et al., 2003; Steck, 2003). All Dacus species that attack
mango have been recently placed in the genus Bactrocera (Drew) (Thomson,
2005).
Pests 319

The following discussion is mainly restricted to the biology, ecology and


management of mango-infesting fruit flies. Regulatory issues, such as risk analysis
and postharvest treatments are discussed by Johnson and Hofman in Chapter 15,
this volume. Postharvest treatments specifically related to mangoes and fruit flies
have been discussed by Sharp et al. (1988, 1989a, b), Hallman and Sharp (1990),
Nascimento et al. (1992), Mangan and Hallman (1998), Shellie and Mangan
(2002a, b) and Bustos et al. (2004). Broad regula-tory issues were recently
reviewed by Follet and Neven (2006).
General reviews on biology, ecology and behaviour of economically important
and non-pestiferous fruit flies, many of them infesting mangoes, were written or
edited by Christenson and Foote (1960), Bateman (1972), Fletcher (1987),
Robinson and Hooper (1989), Aluja (1994), Aluja and Norrbom (2000) and
Malavasi and Zucchi (2000).

Anastrepha
Anastrepha spp. are endemic to the western hemisphere and their range extends
from the southern USA to northern Argentina and includes the Caribbean islands
(Aluja, 1994) (Plate 54). Twelve Anastrepha species have been purportedly
associated with mango (Norrbom, 2004). Of these, A. obli-qua, A. ludens (Loew)
and A. suspensa (Loew) stand out as economically important pests of mangoes
(Aluja et al., 1996; Norrbom, 2004). Anastrepha obliqua is reportedly the most
common fruit fly pest in the Americas (Jirón and Hedström, 1988; Nascimento et
al., 1992). This species is the most com-mon fruit fly pest of mangoes in Mexico,
Costa Rica, Honduras and Guate-mala (Soto-Manitiú et al., 1987; Jirón and
Hedström, 1991; Aluja et al., 1996; Camargo et al., 1996; Sponagel et al., 1996),
but it also attacks mangoes in Cuba, Puerto Rico, Jamaica, El Salvador and
Venezuela (Norrbom, 2004). In Mexico, A. ludens commonly attacks mangoes at
higher elevations, while A. obliqua dominates at lower altitudes (Aluja et al.,
1996). In Brazil and Ecuador, mangoes are mainly attacked by A. fraterculus
(Wiedemann), but A. turpiniae Stone, A. serpentina Wiedemann, A.
pseudoparallela (Loew) and A. zuelaniae Stone have been reported (Zucchi, 1988;
Rebouças et al., 1996; Arias and Jines, 2004; Norrbom, 2004; Barbosa et al., 2005;
A. Malavasi, January 2008, São Paulo, personal communication).

BIOLOGY OF ANASTREPHA FRUIT FLIES. Anastrepha fruit fly biology today is based on
basic studies carried out between 1900 and 1944 (Aluja, 1994, 1999). The basic life
cycle is very similar among most pestiferous Anastrepha species. For example, egg
incubation of A. ludens in mango requires 3.8 days, larval devel-opment requires 14.2
days and pupal development has been known to need 14.2 days at 27  2C (Leyva et
al., 1991). Larvae pass through three instars before emerging from the fruit and
burrowing into the ground to pupate (Aluja, 1994). Clutch size varies between one
egg/clutch in A. obliqua and >100 eggs in A. grandis Mcquart (Aluja, 1994).
Importantly, in species laying sev-eral eggs per clutch (e.g. A. ludens), clutch size is
determined by females on a case-by-case basis and is greatly influenced by degree of
ripeness and the concomitant degree of epicarp hardness (Díaz-Fleischer and Aluja,
2003;
320 J.E. Peña et al.

Birke et al., 2006). Life expectancy varies greatly depending on the host on which
the larvae developed and environmental conditions (for example see Toledo and
Lara 1996, and Malavasi and Zucchi, 2000), but some adults can live for >150 days
(Aluja et al., 2000).
Abundance of A. obliqua populations has been positively correlated with
temperature and negatively correlated with relative humidity (RH) (Herrera and
Viñas, 1977). However, studies by Celedonio-Hurtado et al. (1995) and Aluja et al.
(1996) demonstrated the lack of a clear relationship between rain-fall and
Anastrepha fly captures in mango orchards in Mexico. They indicated that overall
population fluctuation patterns can vary greatly among orchards within a fairly
small geographic region.

Bactrocera
Bactrocera spp. are pests of mango in Africa and Australasia (Drew, 1989; Leblanc
and Allwood, 1997; Leblanc et al., 1997; Tenakanai, 1997; Hancock et al., 2000;
Hollingsworth et al., 2003; Clarke et al., 2005) (Plate 55). The common species
reported on mango include B. tryoni (Frogatt), B. zonata (Saunders), B. dorsalis
(Hendel), B. neohumeralis (Hardy), B. jarvisi (Tryon), B. papayae Drew and B.
frauenfeldi (Schiner) (Umeya and Hirao, 1975; Drew and Hancock, 1994;
Hollingsworth et al., 2003). Two species, B. phillippiensis Drew
& Hancock and B. occipitalis Bezzi, have been recorded for Palau, Pacific Islands
(Secretariat of the Pacific Community, 2005), and recently, a new spe-cies, B.
invadens Drew, Tsuruta and White, was reported for West Africa (Kenya, Benin)
(Lux et al., 2003; Vayssières et al., 2005). Bactrocera correcta (Bezzi), B. caryeae
(Kapoor), B. curcubitae (Coquillett), B. diversa (Coquillett) and B. tau (Walker)
have been reported in India (Australian Government, 2004).

BIOLOGY OF BACTROCERA FRUIT FLIES. The biology of dacine fruit flies was most recently
reviewed by Fletcher (1987); additional details can be found in Christenson and Foote
(1960), Bateman (1972), Robinson and Hooper (1989), White and Elson-Harris (1992)
and Aluja and Norrbom (2000). As with most pestiferous flies, females within
Bactrocera insert their eggs beneath the fruit skin, especially in ripening fruit; white
banana-shaped eggs are usually deposited in clusters, hatching after 1.5–20 days (White
and Elson-Harris, 1992; Messing, 1999). A single female can lay >1000 eggs over her
lifetime (White and Elson-Harris, 1992). One generation requires c.37 days with a
period of 19 days for egg to adult transformation; eggs hatch 38 h after ovi-position;
larvae develop in 7–8 days and adults emerge in 10–11 days (Mess-ing, 1999). There
are usually three larval instars. The larvae tunnel into the fruit, contaminating it with
frass and providing entry for fungi and bacteria. Depending on factors such as
temperature conditions and type of host, larval development can be completed in 7–8
days (Messing, 1999) but can take up to 2 weeks (White and Elson-Harris, 1992).
When the infested fruit is imma-ture, the fruit ripens prematurely and is unfit for
marketing. Fully-grown larvae c.7 mm long drop to the ground and enter the soil where
they pupate.
Pests 321

After emergence, the females require a protein source for egg maturation (White
and Elson-Harris, 1992). Studies with B. dorsalis in India (Singh, 1991) indicated
that pupal period was longest (18 days) at 15C and shortest (6 days) at 35C.
Warm, humid weather is favourable for Bactrocera fruit flies and pest populations
build up as mango ripening occurs. Bactrocera popula-tions decrease during dry
periods.
Syed et al. (1970) reported that up to 30% of mango fruit were attacked by B.
dorsalis in July and August. Mohyuddin and Mahmood (1993) reported that mango
fruit are heavily attacked in Central Punjab during July and August, with up to 35%
of the fruit being damaged by B. dorsalis and B. zonata. Vayssières et al. (2005)
reported the presence of B. invadens after the first sig-nificant rains in mid-April,
reaching >900 males captured/trap/week. Trap captures peaked at 1800
flies/trap/week in mid-June when the presence of B. invadens was related to
ripening of different mango cultivars. Ekesi et al. (2006) observed that B. invadens
shared mango fruit with Ceratitis cosyra in Africa and suggested that B. invadens
is predominantly a lowland pest.

Ceratitis
Eight Ceratitis spp. have been reported to attack mango fruit. The Mediter-ranean
fruit fly C. capitata (Wiedemann) is a common polyphagous pest in mango-
growing areas of Hawaii USA, Israel, Australia, Spain, Mexico, Réunion and
Brazil and elsewhere in South America (Etienne, 1966; Morin, 1967; Galán-Saúco,
1990; Harris et al., 1993; Barbosa et al., 2005; Woods et al., 2005) (Plate 56). In
Africa the most common species are C. cosyra (Walker), C. fasciventris (Bezzi), C.
rosa (Karrsch), C. anonae (Graham) and less frequently C. capitata (Wiedemann)
(Lux et al., 2003); whereas, C. catoirii Guer. occurs in Réunion (Etienne, 1968).
Ceratitis quinaria and C. silvestrii are considered of economic importance in Benin
(Vayssières et al., 2005). Ceratitis cosyra is broadly distributed across Africa and
causes enormous damage, which can result in total loss of the crop. On average
about 20–30% of mango produc-tion is lost due to this fly species in various
African countries (Lux et al., 2003).

BIOLOGY OF CERATITIS FRUIT FLIES. Flies within Ceratitis, particularly C. capitata, are
quite cosmopolitan, and their basic life cycle varies greatly according to site
(Papadopoulos et al., 1996). In Israel females seek suitable sites for ovipo-sition and
puncture mango fruit early in the season, before the fruit has rip-ened. According to
Wysoki et al. (1993) these ‘barren’ punctures damage the fruit, due to the leakage of
resins from the fruit. The female can oviposit all over the fruit, with no preference for
any part. Later, when fruit development is suitable for maggot development, the
oviposition sites become light in colour and the tissue softens. The fully-grown
maggots leave the fruit and pupate in the soil. The developmental period is c.3–4 weeks
and 8–10 genera-tions/year can occur depending on temperature and other factors
intrinsic to the fly population (Hill, 1975).
322 J.E. Peña et al.

Fruit fly control: brief overview

The current trends in fruit fly control call for coordinated, area-wide approaches
(Hendrichs, 1996, 2001; Tan, 2000; Huang et al., 2006; Enkerlin, 2007; Orankanok
et al., 2007) whose major objective is to overcome the often ineffective and
environmentally unsustainable control schemes resulting from uncoordinated
actions by individual producers. Aluja et al. (1996) pro-pose that since fruit flies
that attack mango also attack other fruit crops in the same area, their management
must be based on mango, wild hosts and other commercially grown host plants.
Thus, to improve the efficiency of fruit fly management, host plant blooming and
fruiting factors need to be elucidated. Hendrichs (1996) stated that when fruit
growers pursue a concerted fruit fly population management strategy over
significantly large areas, the number of fruit flies moving into orchards from
neighbouring orchards is largely reduced. Aluja (1993, 1996) and Aluja et al.
(1996) suggest a fruit fly manage-ment scheme based on border trapping,
enhancement of host-plant resistance through use of plant growth regulators, use of
the sterile insect technique and bait stations and augmentative parasitoid releases
(Sivinski, 1996; Sivin-ski et al., 1996; Malavasi and Zucchi, 2000; Montoya et al.,
2000; Tan, 2000; Dyck et al., 2005; Mangan and Moreno, 2007). Aluja (1993) and
Aluja and Liedo (1986) state that accomplishment of these goals depends on
grower status (rich versus poor), access to technology, cost, scale (single orchard,
regional level, national level), globalization of markets and local regulations
restricting impact on the environment.

Monitoring and sampling


Monitoring fruit flies attacking mango serves different purposes: (i) to apply a
control or management tactic after the presence of the fruit fly is noticed; and (ii) to
verify if fruit fly species will attack mango under natural condi-tions. In general,
thresholds for adult fruit flies are quarantine-mediated (Beers et al., 1993). These
thresholds vary from location to location, but de-pending on the fruit fly species
they are typically based on the capture of a single fruit fly. In other fruit crops, a
threshold of five flies/trap is recom-mended resulting in a reduction from four
chemical sprays to 1.5 sprays/ season (Beers et al., 1993). Sampling for fruit flies
in mango is mostly per-formed using adult traps, because eggs and young larvae
are often difficult to see in the fruit and because the primary aim of management
programmes is to prevent fruit damage.

In the case of pestiferous species within Anastrepha and some Bactrocera


species, the most widely used traps since the early 1970s for monitoring and
controlling populations are glass and plastic versions of the McPhail trap, which is
baited with a mixture of protein (occasionally hydrolysed cotton seed together with
borax, molasses or fermented juices) and water (Balock and Lopez, 1969; Jirón,
1995). More recently, human urine has been success-fully tested as bait for
McPhail and McPhail-type traps for resource-poor farmers in tropical countries
(Piñero et al., 2003; Aluja and Piñero, 2004). The McPhail trap has provided
different results in mango orchards. Balock and
Pests 323

Lopez (1969) reported that high concentrations of McPhail traps reduced the build-
up of fly populations and protected mangoes from severe injury dur-ing certain
periods of the year. However, the McPhail trap has several draw-backs. It is
expensive, breaks easily, is cumbersome to service and, most importantly, is quite
inefficient. Aluja et al. (1989), working in a mixed mango orchard in Chiapas,
Mexico, found that only 31.1% of Anastrepha spp. flies landing on the McPhail
trap were caught with many flies entering the trap but then escaping. Due to the
low efficiency of the McPhail trap it is being replaced with Multi-Lure® traps,
which provide new trap designs. Dry synthetic-food-based lures have also been
developed, i.e. BioLure® (Suterra LLC, Inc., Bend, Oregon) (Heath et al., 1995,
1997; Epsky et al., 1999) and Nu-Lure® (Advanced Pheromone Technologies)
(Robacker and Warfield, 1993; Robacker et al., 1997; Robacker, 2001).

Fruit fly presence has also been monitored in Australia using Dakpot ® fruit fly
traps hung beneath the tree canopy (Anonymous, 1989). Methyl eugenol is
considered the most powerful male lure for oriental fruit flies. Methyl eugenol was
used for successful monitoring, control and erradication of B. dorsalis in Oahu
Hawaii USA (Steiner and Lee, 1955), Rota Island (Steiner et al., 1965) and
Okinawa, Kume, Miyako and Uaekama Islands (Japan) (Iwa-hashi, 1984). It has
been used for monitoring B. umbrosa (F.) in the Philippines (Umeya and Hirao,
1975), and is used to lure B. invadens in Africa, which is unlike other African
Dacini species that are attracted to Cue-lures (Lux et al., 2003; Anonymous, 2005).
In Palau, Pacific Islands, two lures are used to attract mango flies: Bactrocera
fraeunfeldi (Schiner) is attracted to Cue-lure, and B. occipitalis and B.
philippinensis Drew and Hancock to methyl eugenol (Secretariat of the Pacific
Community, 2005). Bactrocera dorsalis and B. umbrosa were monitored and
controlled by mass trapping of males with methyl eugenol and infestations were
brought to sub-economic levels in Pakistan (Mohyuddin and Mahmood, 1993).
However, concern over the carcinogenic-ity of methyl eugenol (Waddell et al.,
2004) calls for the development of other para-pheromones to attract Bactrocera
fruit flies. Trimedlure is still consid-ered an important para-pheromone for the
Mediterranean fruit fly, with the exception of C. cosyra adults, which are attracted
to terpinyl acetate and not to trimedlure (Steck, 2003). The attractiveness of mango
compounds is currently being investigated. For example some of the volatiles
emitted by ‘Tommy Atkins’ mangoes, i.e. terpenes (p-cymene and limonene), are
attractive to C. capitata adults (Hernández-Sánchez et al., 2001).

Many questions linger with respect to the optimal trap number and time for
trap placement in mango groves. In Naru, to produce mango free of B. frauenfeldi,
300–400 traps baited with methyl eugenol plus a toxicant were placed every square
kilometre and trapping density was increased around mango plantings
(Anonymous, 2002). Even though large numbers of traps can be utilized to increase
detection sensitivity, the cumulative costs and logistical considerations do not
make this option practical. Traps with spe-cific and effective lures that can detect
the F1 generation at low trap densities (5–10 traps/km2) would fit the description of
a good detection and monitor-ing device (Tween, 1993).
324 J.E. Peña et al.

Sampling for earlier fruit fly stages can be used to demonstrate that the fruit is
not susceptible to fruit fly attack. For instance, the Caribbean fruit fly, A. suspensa,
may not attack green mango fruit (Peña et al., 2006b). Peña et al. (2006b) initiated
research to determine if the Caribbean fruit fly will attack green ‘Tommy Atkins’
mangoes and infest it under field and forced labora-tory conditions. Through a
sequential collection of fruit from fruit-fly infested mango orchards, fruit were
dissected for eggs and larvae. At the same time, fruit were stored and held for
puparia emergence. In addition fruit were exposed in cages to wild fruit flies and
traps were placed to verify the pres-ence of fruit flies. Estimating the time that
‘Tommy Atkins’ fruit remain immature and therefore non-hosts for fruit flies, may
provide a window for mango exports in some fruit fly-infested areas. Lux et al.
(2003) also mention that small growers tend to harvest fruit before maturation as a
strategy to evade fruit infestation.

Chemical control
Mango plantations account for major insecticide use in the tropics (Cunning-ham,
1984). From the late 1960s to date, the conventional control of fruit flies was
through toxic bait sprays that combine proteinaceous bait (e.g. hydroly-sed protein)
with an insecticide (López et al., 1969; Soto-Manatiú et al., 1987; Mangan et al.,
2006; Mangan and Moreno, 2007). For many years the insecti-cide of choice has
been malathion (Peck and McQuate, 2000; Burns et al., 2001). Fruit flies are
highly susceptible to any insecticide, and other com-pounds have also been widely
used. For example, Singh (1991) reported that 5% aldrin dust, when mixed in soil,
provided the highest residual toxicity to falling mature larvae (23.4% after 15
days), compared to BHC endosulfan and quinolphos. Vergherse et al. (2004),
working on the control of B. dorsalis in India, used a rotation of fenthion (0.05%),
deltamethrin (0.0028%), carbaryl (0.2%) and dimethoate (0.06%) to reduce the risk
of resistance development. Yee (1987) concluded that weekly applications of
malathion for 3 months can also provide effective control.

Since the late 1990s, there has been a concerted effort to find environmen-tally
friendly alternatives to malathion (Peck and McQuate, 2000). Cyromaz-ine,
imidacloprid (organochlorinated compound), spinosad (bacteria-derived
insecticide) and phototoxic dyes (Phloxine B) have been successfully tested against
various fruit fly species (Díaz-Fleischer et al., 1996; King and Hen-nessey, 1996;
Peck and McQuate, 2000; Vargas et al., 2002; Liburd et al., 2004; McQuate et al.,
2005). Despite their success, and as is typical with insecticides intensively applied
on a large scale, resistance has already been documented in the case of spinosad
(Wang et al., 2005; Hsu and Feng, 2006) or collateral damage (i.e. negative impact
on natural enemies) (Stark et al., 2004). In Pakistan, application of pesticides
caused reduction of fruit fly infestations, but their use has created scale insect
problems by eliminating their natural enemies (Mohyuddin and Mahmood, 1993).
Such an effect had been reported by Ehler and Endicott (1984) with pests of olive,
citrus and walnut. Another recent development with respect to chemical control of
fruit flies has been the refinement of the bait-station concept (Mangan and Moreno,
2007).
Pests 325

Without chemical control, the damage from C. capitata as a result of ‘bar-ren’


and fertile punctures can be as high as 60%. Control is achieved by aerial bait
spraying, ground cover spraying and spot spraying of trees. In Spain, chemical
control has been achieved by applying organophosphates and hydrolysed albumen
(Khanzada and Naqvi, 1985), but this approach is under pressure because of
environmental concerns. Decisions on when to apply insecticides are based on the
appearance of the first trapped males (trimedlure-baited traps are used). When
applying insecticides directly, dimethoate (0.1%) and fenthion (0.15%) are used
(Khanzada and Naqvi, 1985). Bait sprays are based on naziman (1:1 protein
hydrolysate: malathion/4 l water; Wysoki et al., 1993). Removal of fallen fruit can
also prevent build-up of Mediterranean fruit fly populations.

Pestiferous Anastrepha spp. are susceptible to most insecticides (Shaw and


Spisshakoff, 1958; Shaw, 1961). Bait sprays applied from the ground and from the
air are successful, but they can cause environmental damage, surges of secondary
pest populations and reductions in parasitoid populations (López et al., 1969; Soto-
Manitiu et al., 1987). In Peru, control measures against Anastrepha in mango begin
when McPhail trap catches average two adults/ trap/week (Herrera and Viñas,
1977). In Mexico, control starts when the fruit is 85-days-old and is suspended 2
weeks before harvest (Cabrera et al., 1993). In Costa Rica, dipterex and malathion
are sprayed weekly and reduce mango damage up to 40% (Soto-Manitiu et al.,
1987). In Brazil, malathion with protein and sugarcane bagasse is used (Carvalho
and De Queiroz, 2002). In Ecuador, Arias and Jines (2004) recommend a spray of
malathion (1%) with protein (4%) once the fruit fly population reaches 0.14 fruit
flies/trap/day (FTD).

Biological control
PARASITOIDS. Classical biological control and repeated augmentative releases of mass-
reared parasitoids have been used to suppress Anastrepha, Ceratitis and Bactrocera
populations (Wharton, 1978; Sivinski, 1996; Sivinski et al., 1996, 1997, 2000; Montoya
et al., 2000). In Florida USA, Mexico, Costa Rica, Brazil, Colombia and Peru,
parasitoid species (i.e. Diachasmimorpha longicau-data (Ashmead), Fopius
vandenboschi (Fullaway) and Aceratoneuromyia indica (Silvestri)) have been imported
and released for the control of A. suspensa, A. ludens and A. fraterculus (Ovruski et al.,
2000). Despite the widespread use of exotic parasitoids over the past 80–100 years, the
current trend is to resort to native species as they pose less of an environmental threat to
local fauna (García-Medel et al., 2007; Aluja et al., 2009).

Use of parasitoids with mango is hindered by the fact that fruit are very large
and therefore provide larvae a refuge from parasitism (López et al., 1999). As a
consequence, Aluja (1993) and Montoya et al. (2000) recommended that
parasitoids should be released outside the mango orchards to attack fly larvae in
their much smaller native hosts and thereby significantly reduce the size of
populations entering mango orchards.
Several parasitoids, for example Opius fullawayi (= Diachasmimorpha
fullawayi (Silvestri)), Diachasmimorpha kraussi, D. Fullaway, D. tryoni (Cam-
eron), Opius bellus Gahan, Biosteres longicaudatus Ashmead (= D. longicaudata),
326 J.E. Peña et al.

B. tryoni (Couron) (= D. tryoni (Cameron)) and Biosteres oophilus Fullaway (=


Fopius arisanus (Sonan)), parasitize C. capitata (Beardsley, 1961; Wharton and
Marsh, 1978). Bess et al. (1961) reported that the most important parasi-toids
collected from C. capitata in Hawaii USA were F. vandenboschi, B. oophilus (=
Opius oophilus) (= F. arisanus) and B. longicaudatus. In Brazil, mainly Doryc-
tobracon areolatus (Szépligeti) (97%) and D. longicaudata (3%) parasitize fruit fly
larvae attacking mango (Carvalho and De Queiroz, 2002). In Kenya, Ghana,
Tanzania, Uganda and Cote d’Ivoire, the most important parasitoids obtained from
Ceratitis spp. emerging from mangoes were D. fullawayi, Fopius cauda-tus
(Szépligeti), Psyttalia cosyrae (Wilkinson) and Tetrastichus giffardianus Sil-vestri
(Lux et al., 2003). In Mexico and other parts of Latin America, the most common
parasitoids attacking fruit flies that infest mangoes (A. obliqua, A. ludens, A.
pseudoparallela, A. turpiniae) are D. areolatus, Doryctobracon brasilien-sis
(Szépligeti), Doryctobracon crawfordi (Viereck), Doryctobracon fluminensis
(Lima) and Utetes anastrephae (Viereck) (López et al., 1999; Ovruski et al., 2000;
Zucchi, 2000). In Pakistan, the parasitoids attacking B. zonata include Opius
longicaudatus (= D. longicaudata), Dirhinus giffardii Silvestri, and Bracon sp.; O.
longicaudatus (= D. longicaudata), D. giffardii and Spalangia grotiusi Girault
were reported to attack B. dorsalis, albeit in small numbers (Syed et al., 1970).

Microbial control
Use of pathogens/disease agents (fungi, bacteria, nematodes) has been attempted
with varying degrees of success. For example, Metarhizium anisopliae has been
evaluated in small-scale mango orchards in Kenya using bait stations laced with the
pathogen. Results do not show differences between use of pathogens and use of
insecticides (malathion) (Lux et al., 2003). Lezama-Gutierrez et al. (2000) also
evaluated isolates of M. anisopliae against larvae of A. ludens. They suggested that
M. anisopliae can cause a 22–43% reduction in adult emergence, depending on the
soil where the lar-vae pupariates. De la Rosa et al. (2002) evaluated the fungus
Beauveria bassi-ana (Bals.) under laboratory conditions and concluded that highest
mortality was achieved at the adult stage, while Dimbi et al. (2003) reported on the
pathogenicity of M. anisopliae and B. bassiana on different species of Ceratitis.
Poinar and Hislop (1981), Lindegren and Vail (1986) and Toledo et al. (2006) have
investigated the use of various nematodes, Heterorhabditis bacteriophora,
Heterorhabditis heliothidis (Khan, Brooks and Hirschmann) and Steinernema
feltiae Filipjev, against Anastrepha, Bactrocera and Ceratitis. Finally, Robacker et
al. (1996) and Toledo et al. (1999) have tested various strains/isolates of Bacillus
thuringiensis (Berliner) against larvae of A. ludens, A. obliqua and A. serpentina.
For additional details on microbial control of pestiferous fruit flies, we recommend
the recent review by Dolinski and Lacey (2007).

Predators
In addition to parasitoids, pathogens and nematodes, ants have been used to control
fruit flies in mango orchards. Peng and Christian (2006) used the weaver ant,
Oecophylla smaragdina (Fabricius) for control of the Jarvis fruit fly, B. jarvisi, in
mango orchards in Australia. Van Mele et al. (2007 and references
Pests 327

therein) in Benin used an African weaver ant (Oecophylla longinoda). Aluja and
colleagues (Aluja et al., 2005) investigated the potential of ants as possi-ble
biological control agents in various tropical orchards and backyard gar-dens in
which mango trees were growing with other fruit trees.

Cultural fruit fly control


Fruit bagging is one of the best solutions to prevent fruit fly attack of mango and
other tropical fruits (Aluja, 1996; Peña et al., 1999; Paderes and Orden, 2004).
Success with mangoes can be quite high, but Bondad (1985) demon-strated that
bagging materials do not always resist the effect of rain/wind. Therefore, while
bagged mangoes tend to produce a greater amount of mar-ketable fruit than those
not bagged, more research is needed to determine the type of bags to use for
different mango varieties and the best time to bag fruit (Love et al., 2003).

Jirón (1995) reported that A. obliqua populations could be reduced by


increasing planting distances in order to reduce RH and increase solar radia-tion
within orchards. In India, cultural control practices include removal of fallen fruit
and inter-tree ploughing and raking (followed by insecticide cover sprays). Such
practices can reduce fruit fly infestation between 77% and 100% (Verghese et al.,
2004).
A cultural practice that can impinge on the success of management pro-
grammes is the widespread use of potassium nitrate (KNO 3) sprays to accel-erate
and synchronize flowering of mango trees. As a result, fruit harvests can be
advanced and synchronized. Such a procedure can, under certain cir-cumstances,
help control fruit flies, but can also exacerbate the problem. For example, in the
case of the Mexican fruit fly (A. ludens), a notorius pest of citrus that also attacks
mangoes, advancing the mango harvest offers ideal conditions for adult pests to
move from citrus groves to mango orchards. This would not occur if mango trees
flowered naturally since fruit ripen sev-eral months after the citrus harvest (Martin
Aluja, personal observation).

Host resistance
Yee (1987) reported that B. dorsalis does not attack all mango cultivars to the same
extent. The most susceptible cultivars in Hawaii USA are ‘Hawaiian’, ‘Pirie’ and
‘Sandersha’. Singh (1991) indicated that the frequency of Bactro-cera injury to
physiologically mature fruit of ‘Dashehari’ ranged from 3.6 to 10%, while in fully
ripe fruit the frequency of injured fruit ranged from 10 to 25.9%. Highest damage
was reported in fully ripe fruit of ‘Mallika’ followed by ‘Totapari’.

Susceptibility of different mango cultivars to attack by A. obliqua was


measured by Carvalho et al. (1996) who observed that ‘Espada’ showed no
infestation by A. obliqua, whereas ‘Carlota’ was highly infested. In this study, the
survival of adults of A. obliqua was lower when the larvae were fed on ‘Espada’
compared to ‘Carlota’. Furthermore, ‘Espada’ had an adverse effect on the
longevity of A. obliqua females, possibly due to the presence of toxic substances
(Carvalho and De Queiroz, 2002) or absence of essential nutri-ents. Jirón and Soto-
Manitiu (1987) also observed that susceptibility of
328 J.E. Peña et al.

mangoes to A. obliqua differed among cultivars. ‘Rosinha’, ‘Coquinho’ and


‘Espada’ were resistant to A. obliqua attack, whereas ‘Smith’ and ‘Pope’ were
highly susceptible. According to Joel (1980), mangoes contain resin ducts in the
exocarp that confer protection against the vertical movement of the ovi-positor and
larval movement. Other studies have shown that resistance is related to degree of
maturity (Díaz-Fleischer and Aluja, 2003; Aluja and Mangan, 2008); immature
mango fruit are less susceptible to A. suspensa than mature mangoes when infested
artificially (Hennesey and Schnell, 2001). In Brazil, use of gibberellic acid (GA 3)
reduces the susceptibility of ‘Tommy Atkins’ mangoes to attack by C. capitata
based on artificial delay of fruit maturity (de Macedo, 1988). Differences on attack
by A. ludens to mango might be influenced by volatiles from green or yellow fruits
(Garcia-Ramirez et al., 2004).

Quarantine treatments
Quarantine treatments have been reviewed by Johnston and Hofman (Chap-ter 15,
this volume). Several quarantine treatments have been developed for harvested
mangoes: irradiation, hot water or hot water followed by immer-sion cooling are
widely used (Sharp et al., 1988, 1989a, b, c; Hallman and Sharp, 1990; Nascimento
et al., 1992; Mangan and Sharp, 1994; Mangan and Hallman, 1998; Shellie and
Mangan, 2002a, b; Bustos et al., 2004; additional references in reviews by Mangan
and Hallman, 1998 and Follet and Neven, 2006).

Mango seed weevils

The mango seed weevil, Sternochetus mangifereae (Fabricius), and the mango pulp
weevil, Sternochetus frigidus (Fabricius) (Coleoptera: Curculionidae) are important
pests of mango (Plates 57–59). Quarantine restrictions prevent the export of fresh
weevil-infested mangoes into uninfested areas. The flesh of ripe fruit is damaged
when mango seed weevil adults emerge from the seed, and weevil-damaged seed
may limit plant propagation in nurseries and orchards (Johnson, 1989). Early fruit
drop may be caused by severe weevil infestations (Subramanian, 1925). The
mango seed weevil occurs from India through South-east Asia to Australia, on
tropical Pacific Islands, in parts of Africa, in the Caribbean region and in northern
South America (Balock and Kozuma, 1964; Shukla and Tandon, 1985; Johnson,
1989; Schotman, 1989).

Biology
Srivastava (1998) reports that S. mangiferae is a greyish brown weevil, 8 mm long
and about 4 mm wide; its habits are nocturnal, usually feeding and ovi-positing at
dusk. After emergence, adults enter diapause and the duration varies with the
geographic range (Schotman, 1989). According to Shukla and Tandon (1985), all
adults emerging in southern India during June enter dia-pause from July until late
February in the following year. The beginning and the end of diapause seem to be
associated with long-day and short-day
Pests 329

photoperiods, respectively (Balock and Kozuma, 1964). The mango seed weevil
produces only a single generation each year. In Tamil Nadu, India, adults feed on
leaves and tender mango shoots in March and April (Subra-manian, 1925). Shukla
and Tandon (1985) report that females began oviposit-ing 3–4 days after mating
when fruit reaches a marble-size. Oviposition varied from 3 to 5 weeks
(Subramanian, 1925; Shukla and Tandon, 1985; Hansen et al., 1989). The female
uses its snout to make a cavity in the fruit, lays a single egg and then covers it with
a secretion (Pradhan, 1969).
According to Srivastava (1998), about six larvae can be found within a mango
seed. Generally, only a single larva completes development within each fruit.
Larval development and pupation occurs within the seed. Adults are formed 1
week later; however, adults generally emerge from the seed c.1–2 months after
fruit drop (Balock and Kozuma, 1964). The weevils over-season under bark and
stone walls, where they remain dormant until the next flowering season (Van Dine,
1906; Balock and Kozuma, 1964; Shukla and Tandon, 1985).

Sternochetus frigidus attacks Mangifera indica, M. foetida and M. sylvatica.


The weevil occurs in Bangladesh, Brunei, India, Indonesia, Myanmar, Paki-stan,
Papua New Guinea, the Philippines, Singapore and Thailand (CAB International,
2003). Sternochetus frigidus lays eggs on mango fruit with a minimum diameter of
6 cm (De and Pande, 1988). Newly hatched larvae tun-nel directly through the fruit
pulp. Pupation takes place in a brown cocoon within a chamber adjacent to the
kernel. The weevils leave the ripe fruit through a hole in the peel. This pest is likely
to survive storage and transpor-tation (Australian Government, 2004).
Reproductively immature adult wee-vils overwinter inside seed or other protective
places from May until February in India (De and Pande, 1988).

Sampling
Shukla et al. (1988) reported the intra-tree distribution of eggs of S. mangiferae on
‘Baganpalli’ mango; the highest number of eggs per fruit occurred on fruit in the
lower region of the tree. With increasing tree height, egg deposition on fruit
decreases. No statistical differences on fruit infestation were observed on north,
south, east or west directional quadrats of the tree. Eggs were deposited in the
lower region of the fruit rather than the pedicel. Weevils enter diapause in crevices
in the tree trunk. Most of the weevils (87%) are at a height of 0–2 m in the trunk
compared to 7% at 2–4 m and only 4% above 4 m. Emery (2002) considered that
since both the mango pulp weevil (S. frigi-dus) and the mango seed weevil (S.
mangiferae) infest fruit at an early stage, any fruit is a viable sample; however, as
infested fruit all ripen precociously, the sensitivity of surveys is enhanced by
seeking out nearly ripe and fallen fruit prior to harvest. If the survey coincides with
the mango harvest, rejected or fallen fruit should be inspected. Fruit should be
sampled by longitudinal dissection of fruit through the seed to expose the kernel. If
the fruit is ripe, it should be struck along the longitudinal axis with a hammer, and
the seed should be opened with pliers. The random sampling of 600 fruit from ran-
domly selected properties in each area provides a 9% chance of detecting a
330 J.E. Peña et al.

0.5% infestation of fruit. Sample size can be determined from the following
formula:
Probability of > 1 infested fruit = 1 – probability of no infected fruit in total
sample = 1 – (1 – 0.5%)600 = 1 – (1 – 0.005)600 = 1 – (0.995)600 = 95%

Economic damage
Follett and Gabbard (2000) report that germination rates for infested seed of
polyembryonic ‘Common’ are equal to those of uninfested seed. Germina-tion is
significantly reduced for infested seed of monoembryonic ‘Haden’ compared with
uninfested control seed, although germination of infested seed was >70%. Direct
feeding damage to the pulp was found in only 0.11% of 3602 mango fruit, which
suggests that S. mangiferae is a less serious pest of mangoes than previously
considered.

Cultural control
Field sanitation, i.e. the removal of all fallen fruit and seed, is very labour
intensive, and demands complete removal and disposal of fallen fruit. This
procedure has been inconsistent in demonstrating pest control. In India, field
sanitation reduced infestation of the mango nut weevil, Sternochetus gravis
(Fabricius), by only 22% (De and Pande, 1987). In Hawaii USA, field sanita-tion
failed to reduce infestation rates (Hansen and Amstrong, 1990).

Chemical control
Various insecticides have been evaluated for controlling adult weevils, par-ticularly
during oviposition (Balock and Kozuma, 1964; Shukla and Tandon, 1985). The
most effective control was provided by the organophosphate fen-thion, which
reduced infestation to <17%. In another field test, the pyrethroid deltamethrin and
the carbamate carbaryl were most effective, both resulting in <15% infestation
rates. Spot application of diazinon on tree trunks was recommended based on cost,
efficiency and least environmental damage. Verghese et al. (2004) reported that
commercially available azadirachtin was not effective for management of S.
mangiferae in India.

Resistant cultivars
Mango cultivars resistant to the mango weevil would be beneficial. Potential
mechanisms of resistance are seedless cultivars, those that form seed early or those
that fruit off-season. Most cultivars grown in Hawaii and India are equally
susceptible (Bagle and Prasad, 1984; Hansen et al., 1989), although ‘Itamaraca’
has shown some resistance (Balock and Kozuma, 1964).

Biological control
The mango weevil has few natural enemies. Parasitoids are unknown, prob-ably
because of the concealed nature of most life stages. Adults may be sus-ceptible to
predation by ants, rodents, lizards and birds (Hansen, 1993). A baculovirus has
been reported that affects the larvae of S. mangiferae (Shukla et al., 1984).
Pests 331

Mango fruit infested with seed weevil do not show any visible external
symptoms and cause considerable quality control problems and economic loss to
the mango-processing industry as well as restriction in export of fresh fruit. A non-
destructive X-ray inspection method has been developed to detect weevil-infested
fruit. X-ray radiographs of infested mangoes show dark areas in the seed
corresponding to disintegrated kernel tissue as a con-sequence of feeding by
developing grubs. Non-infested mangoes show a uni-formly light-grey area
representing healthy kernel. There is a close agreement between fruit showing
weevil infestation based on their X-ray images and physical examination of cut
fruit, indicating the reliability of the technique. X-ray imaging has good potential
for application in the processing industry and the export trade as a quality control
measure (Thomas et al., 1995).

Mango seed borer (Lepidoptera: Pyralidae)

Distribution and biology


The red banded mango caterpillar or mango seed borer, Deanolis sublimbalis
Snellen, also referred to as Noorda albizonalis Hampson (Waterhouse, 1998), is an
important pest of mangoes in the Philippines (Anonymous, 1984), India
(Zaheruddeen and Sujatha, 1993), Vietnam (Nguyen et al., 1998; van Mele et al.,
2001), China (Li et al., 1997), Thailand, Indonesia and Papua New Guinea
(Cunningham, 1984). The oval white eggs are laid in groups at the fruit apex and
take 3–4 days to hatch. The larvae develop through five instars in 14–20 days and
they pupate in cocoons in the soil. The period from egg to adult requires 28–40
days. The insect apparently prefers mango, but M. odorata and M. minor have also
been recorded as hosts. Adults are generally nocturnal and spend most of the day
under leaves on the tree. A shorter developmental period has been observed when
larvae develop on pulp rather than in seed of ‘Carabao’ fruit, although those reared
on the seed were larger and lived longer (Waterhouse, 1998).

Damage
Mango fruit in all stages of development are susceptible to attack (Water-house,
1998). The first larval instars feed on tissues beneath the skin, and bore through the
mango pulp to the seed, which is consumed. Up to 11 larvae have been recorded in
a single fruit, but usually there is only one. Infested fruit split and rot, and fall to
the ground (Anonymous, 1984). In the Guimaras Islands, the Philippines, Golez
(1991) recorded 12.5% fruit infestation and in serious outbreak years, 40–50%
yield reductions are possible. Waterhouse (1998) considered that since D.
sublimbalis is capable of causing such levels of damage, it might be a more
important pest of mangoes than has generally been realized. It may have been
overlooked as a pest or has recently spread to new areas and has become evident as
a pest there. Van Mele et al. (2001) suggested that damage caused by D.
sublimbalis in the Mekong Delta has been wrongly attributed to fruit flies;
however, Waterhouse (1998) states that soon after boring by D. sublimbalis,
secondary infestations with fruit flies
332 J.E. Peña et al.

(Bactrocera ferrugineous, B. frauenfeldi) occur, together with infections by bac-


teria and fungi.

Biological control
According to Waterhouse (1998) no parasitoids were detected in Java, Indo-nesia.
However, in the Guimaras Islands of the Philippines, the vespid wasp, Rychium
attrisimum, preys on the larvae as they exit the fruit to pupate. Lar-vae are used to
stock the wasps’ nests as food for their young. The egg para-sitoids Trichogramma
chilonis Ishii and Trichogramma chilotreae attack the pest in Luzon (Golez, 1991).
Leefmans and van der Vecht (1930) reported that an entomopathogenic fungus
infected the larvae in Indonesia.

Monitoring and control


Infested fruit can be detected by the presence of a dark-brown ring and cat-erpillar
frass at the point of entry (CAB International, 2003). Mango fruit become
susceptible to the seed borer c.60 days post-induction, and insecti-cide applications
should commence then. Further sprays at 75, 90 and 105 days post-induction are
required to fully protect the fruit. The most effective chemicals are deltamethrin
and cyfluthrin (Golez, 1991). Infested fruit should be removed from trees before
the larvae can leave them to attack neighbour-ing fruit; fruit should be wrapped in
protective bags at 55–65 days after pol-lination, and fallen fruit should be
destroyed (Anonymous, 1984).
Other Lepidoptera that can attack fruit have been reported in India and the
Philippines (Australian Government, 2004). The pomegranate fruit borer,
Deudores isocrates (Fabricius) (Lepidoptera: Lycaenidae), which is also a pest of
loquat, lychee, guava and pear, could attack mango by laying single eggs on
shoots; the emerging larva bore into the fruit (Srivastava, 1998). In the Philippines,
the larvae of the cocoa tussock moth, Orgyia postica (Walker) (Lepidoptera:
Lymantriidae), regularly attack leaves of cocoa, but its larva also attack mango
fruit and panicles (Fasih et al., 1989).

Fruitspotting bugs (Hemiptera; Coreidae)

The yellowish green coreid bugs, Amblypelta lutescens lutescens (Distant) and
Amblypelta nitida Stål occur along the coast of Queensland, Australia, and attack
most of the tropical and subtropical fruit crops there (Waite and Huwer, 1998).
They prefer to feed on young, green fruit, but A. l. lutescens also dam-ages the
terminals of a number of hosts. In tropical north Queensland, A. l. lutescens is the
dominant species and feeds on young fruit causing black lesions to develop and the
fruit to fall. It also feeds on the terminals and leaf petioles, causing wilting and
dieback (Cunningham, 1989). In the subtropical south, both species attack mango,
but A. nitida confines its attention to the fruit, while A. l. lutescens also attacks fruit
and terminal growth (G.K. Waite, 1995, unpublished results). The bugs breed in
natural rainforest areas, and fly into the orchards to feed on the fruit and terminals.
Female bugs lay individual, opalescent green eggs under the leaves. There are five
nymphal instars and a generation takes c.40 days.
Pests 333

The main predators of fruitspotting bugs are spiders, particularly mem-bers of


the family Thomisidae. Several species of egg parasitoids have been recorded. In
north Queensland, Ooencyrtus sp. (Encyrtidae), Anastatus sp. (Eupelmidae) and
Gryon sp. (Scelionidae) parasitized 37.5–91.6% of eggs of A. l. lutescens (Fay and
Huwer, 1994). In south Queensland, Anastatus sp. and Gryon meridianum (Dodd)
parasitize eggs of A. nitida and A. l. lutescens to a similar extent (Waite and Petzl,
1994).
Because fruitspotting bugs continuously migrate into orchards, more than one
insecticide spray may be required to protect the young fruit. How-ever, the fruit are
safe from attack once they are c.50 mm long, and two or three sprays of endosulfan
at intervals of 2 weeks are generally sufficient to protect them.

The coconut bug, Pseudotheraptus wayi Brown, was first recorded on man-
goes in South Africa in 1977, and now also attacks guavas, pecans, macada-mias,
avocados and loquats. It causes damage similar to that of Amblypelta spp. (De
Villiers, 1990).
Helopeltis spp. (Miridae) are minor pests of mango, cashew and cacao in the
Philippines and in northern Australia, where they feed on fruit and cause
superficial corky blemishes. Insecticides are used to control them, but in the
Philippines, bagging is also effective (Anonymous, 1984).
Plant bugs within the Lygaeidae and Pyrrhocoridae, i.e. the Indian milk-weed
bug, Spilostethus pandurus (Scopoli) and the red cotton bug, Dysdercus koenigri
(Fabricius), can injure fruit, inflorescences and leaves of mangoes in India
(Australian Government, 2004). However, D. koenigri is an important pest of
cotton rather than mango (Schaefer and Ahmad, 2000), while S. pan-durus feeds
preferentially on members of the Asclepediaceae (i.e. Calotropis) (Sweet, 2000).

A hymenopterous parasitoid complex attacking the eggs of the banana-spotting


bug, A. l. lutescens, was first reported in north Queensland. It includes an Anastatus
sp. (Eupelmidae), Ooencyrtus sp. nov. (Encyrtidae) and a Gryon sp. (Scelionidae),
and is similar to other complexes known to attack eggs of related coreids in Africa,
Indonesia and Papua New Guinea. Parasitism ranged from 37.5–91.6% in eggs
collected at three sites from orange jessa-mine, Murraya paniculata (L.) Jack.
Anastatus sp. was the dominant parasitoid (Fay and Huwer, 1994).

Thrips

Grove et al. (2001) reported that in South Africa, the citrus thrips, Scirtothrips
aurantii Faure and the red banded thrips Selenothrips rubrocinctus (Giard) are the
only thrips that caused lesions on fruit (Plates 60–63). However, Grove et al.
(2001) considered S. aurantii to have more economic importance than S.
rubrocinctus. Grove and Pringle (2000), using a two-stage sampling system to
determine population levels of S. aurantii, showed that S. aurantii has a clumped
distribution, and recommended that 50 fruit per orchard should be examined in
order to obtain accurate population estimates for pest management.
334 J.E. Peña et al.

Blossom pests

Midges, caterpillars, hoppers, thrips and mites are the most important pests
attacking mango inflorescences.

Midges
The mango gall midge or mango blister midge, Erosomya mangiferae Felt, is a
major pest, destroying flowers and up to 70% of set fruit (Plate 64). It was first
described by Felt (1911) in St Vincent (West Indies). Barnes (1948) recog-nized
nine gall midges from mango; two of these, Asynapta sp. and E. man-giferae, are
from the West Indies. Butani (1979) reported five cecidomyiid species on mango
blossoms, including Erosomya indica (Grover and Prasad). Dasyneura mangiferae
(Felt) was reported in Hawaii USA (Vannière et al., 2004). In recent times, Gagne
and Etienne (2006) reported the species Gephy-raulus mangiferae (Felt), n.comb.
infesting mango flowers on the island of Guadeloupe, French West Indies. Male
adults of E. mangiferae are 1.61 mm and females 1.32 mm long. Eggs are
deposited in folds between sepals and petals of flower buds. The larval stage has
four instars. Young larvae are cream coloured and late instar larvae are yellowish.
Larval feeding prevents flower opening and consequently development of the fruit
does not occur. Infested buds develop as long pointed galls, in which pupation
occurs (Van-nière et al., 2004). Studies of population fluctuation of Erosomya sp.
have been conducted in India by Grover (1986a), who reported that emergence of
adults was higher at 24C and 60–82% RH compared to lower temperatures and
RH. Abbas (1985) described systematic surveys to determine the percentage of
infestation of E. indica, and showed that infestation follows a negative binomial
infestation. The midge infests the newly emerged panicles by ovi-positing at bud
burst stage, and the first instar maggots bore into the grow-ing panicle. The second
generation then infests very young fruit, which drop before the marble stage.
Sampling of mango midges needs to include affected tissue, different trapping
devices, pheromones, etc. On citrus, use of coloured sticky traps placed in the tree
canopy provides a more efficient method of sampling the citrus midge, Prodiplosis
longifila Cagné, than ground emergence traps and collection of larval samples
(Peña and Duncan, 1992).

In a survey of parasitoids of cecidomyiid pests of mango in India, Grover


(1986b) reported that Platygaster sp., Systasis sp. and Eupelmus sp. were asso-
ciated with Dasineura sp., and Tetrastychus sp. was associated with E. indica. An
external parasitoid, the pteromalid, Pirene sp., attacked Procystiphora mangiferae
(Felt). Predators of the cecidomyiids include Formicai sp., Oeco-phila sp. and
Camponotus sp.

Mango hoppers
Approximately 18 species of leaf hoppers have been reported as pests of mango. Of
these, Idioscopus clypealis Leth., Idioscopus niveosparsus Leth., Idiosco-pus
magpurensis Pruthi and Amritodus atkinsoni Leth., are important (Virakta-math
and Viraktamath, 1985; Viraktamath, 1997; Fletcher and Dangerfield, 2002) (Plate
65). The females deposit their eggs in panicles or midribs of tender
Pests 335

leaves. The adults and nymphs preferentially feed on young leaves and flow-ers or
shoots, and excrete honeydew upon which sooty mould develops (Ahmed et al.,
1981). This interferes with photosynthesis, adversely affecting plant growth and
yield (Godase et al., 2004). Affected inflorescences turn brown, become
dehydrated and fruit set does not occur.
There has been no systematic study of the biology of most of the leaf hoppers
that attack mango; however, biology of A. atkinsoni, I. clypealis and I. niveosparus
has been studied by Sohi and Sohi (1990). Both A. atkinsoni and I. niveosparsus
are multivoltine. In A. atkinsoni, the egg, nymphal (five instars) and adult stages
require 7–9, 15–17 and 3–4 days, respectively (Patel et al., 1977). Development
from egg to adult is normally complete in 25–30 days. There can be between one
and six generations of A. atkinsoni in different areas of India. In Pakistan there are
four to five generations in the Central Punjab (Mohyuddin, unpublished data). In
the Philippines, I. clypealis is reported to have one to four generations, whereas it
has five or six generations in India.
Idioscopus nagpurensis is univoltine. In Pakistan, it normally oviposits in
mango inflorescences during March. Nymphs feed on inflorescences during March
and April. From May to February of the following year, only aestivat-ing adults are
found (Mohyuddin, unpublished data). Most of these species are quite fecund.
Amritodus atkinsoni reportedly lays 200 eggs during its life-time (Rahman, 1940)
and I. clypealis lays 100–190 eggs in the Punjab (Husain and Pruthi, 1924).
Amritodus atkinsoni eggs are laid in the midribs of tender leaves, flower buds and
inflorescences (Babu et al., 2002). Idioscopus niveospar-sus lays c.238 eggs in 9
weeks under laboratory conditions (Mohyuddin, unpublished data). Mohyuddin
and Mahmood (1993) reported that A. atkin-soni and I. niveosparsus occur in upper
portions of mango trees during differ-ent times of the year. Amritodus brevistilus
and I. niveosparus populations increase from February to peak in March–April in
Sri Lanka, while peaks of I. clypealis occur in March and September (Kudagamage
et al., 2001). Idiosco-pus clypealis populations peak in south-eastern India during
March and April (Tandon et al., 1983). Idioscopus nivesoparus and I. clypealis
peaks coincided with major and minor flowering periods while population peaks
for A. bre-vistilus coincide with the occurrence of vegetative flushing
(Kudagamage et al., 2001).

Azizur Rahman and Singh (2004) demonstrated that A. atkinsoni popula-tions


on panicles of ‘Langra’ mango were negatively correlated with high RH; whereas
no significant relationships were observed with rainfall, sun-shine and wind
velocity.

SAMPLING. Very few sampling studies have been reported for hoppers on mango. A
sequential sampling plan for mango hoppers was recommended by Verghese et al.
(1985) in India. Mohyuddin and Mahmood (1993) reported sampling by direct visual
examination: A. atkinsoni and I. niveosparsus were found on upper portions of mango
trees during different times of the year. They moved to the lower parts of the stems and
the leaves during summer. Tandon et al. (1989) reported that distribution of I.
niveosparsus was aggre-gated and was best explained by Iwao’s patchiness regression.
To assess
336 J.E. Peña et al.

damage, they recommended a sampling size of 59–98 panicles/tree. Verghese et al.


(1985) developed a sequential sampling plan classifying infestations of adults and
nymphs of I. clypealis as light, moderate and severe.

BIOLOGICAL CONTROL. Several natural enemies have been described from west and
South-east Asia. Mohyuddin and Mahmood (1993) reported the egg par-asitoids,
Gonatocentrus sp., Miurfens sp. nr. mangiferae Viggiani and Hayat, Centrodora sp. nr.
scolypopae Valentine, Aprostocetus sp. and Quadrastichus sp., and the adult
ectoparasitoid Epipyrops fuliginosa Tames in Pakistan. Fasih and Srivastava (1990)
reported that Aprostocetus sp., Gonatocerus sp. and Polynema sp. parasitize eggs. Five
species of predators, including Chrysopa lacciperda (Kimmins), Mallada boninensis
(Okomote), Bochartia sp. and two unindenti-fied species (one each of Mantidae and
Lygaeidae) prey on nymphs (Fasih and Srivastava, 1990). In India, Sadana and Kumari
(1991) studied the effi-cacy of the lyssomanid spider, Lyssomanes sikkimensis on I.
clypealis. Classical biological control of mango hoppers has not been attempted.
Whitwell (1993) described four genera of parasitoids from Dominica, the most
common being Aprostocetus sp., followed by Platygaster sp., Synopeas sp. and
Zatropis sp. Peng and Christian (2005a, b) reported that the weaver ant, Oecophylla
smarag-dina (Hymenoptera: Formicidae) is an efficient biocontrol agent of I. nididulus
in northern Australia. The entomopathogens, Verticillium lecanii (Zimmer-man)
Viegas, Beauveria bassiana Balsamo (Vuillemin) and Isaria tax, infect I. clypealis in
India (Kumar et al., 1993; Srivastava and Tandon, 1986) while the effectiveness of
Metarhizium anisopliae var. anisopliae was tested under laboratory conditions against
A. atkinsoni (Vyas et al., 1993).

CHEMICAL CONTROL. Several pesticides have been tried for controlling mango hoppers
(Tandon and Lai, 1979; Yazdani and Mehto, 1980; Shah et al., 1983; Shukla and
Prassad, 1984; Islam and Elegio, 1997; Kudagamage et al., 2001). Khanzada and Naqvi
(1985) reported that six sprays of fenitrothion/year were effective for controlling mango
hoppers in Pakistan. Nachiappan and Baskaran (1986) tested eight insecticides:
phasalone, endosulfan, carbaryl, penthoate, fenitrothion, monocrotophos, quinalphos
and phosphamidom. Endosulfan provided the best control when spraying was done 1
week after flowering and then 14 days later. Mohyuddin and Mahmood (1993) reported
that monitor applied at 5 m on tree trunks and leaves in May provided control of mango
hoppers. Jhala et al. (1989) considered that sprays of carbaryl during the off-season
maintained the hopper population at low-density levels.

Godase et al. (2004) demonstrated that sprays of 0.05% monocrotophos at the


first panicle emergence and a second spray 15 days later are essential to prevent
yield loss. Kudagamage et al. (2001) found that imidacloprid (Admire SL 200)
controlled mango hoppers if applied just after flowering and again 10 days later.

Lepidoptera
The lepidopteran flower feeders are the second most important inflorescence pests
of mango. Geometrids, for example Chloropteryx glauciptera Hampson and
Pests 337

Oxydia vesulia (Cramer) infestation was reported in Dominica by Whitwell (1993).


Infestations increase during the flowering season, averaging three
larvae/inflorescence (87% infestation) to c.100% infestation later in the flow-ering
season. Eggs of the noctuidae Penicillaria jocosatrix Gueneé are laid pre-
dominantly on or near the inflorescences or new leaves. The microlepidoptera
complex attacking mango in Florida USA consists of Pococera attramentalis
Lederer, Pleuroprucha insulsaria (Gueneé) (Plate 66), Platynota rostrana
(Walker), Tallula spp. and Racheospila gerularia (Hübner) (Plate 67). Most of the
damage to inflorescences (Plate 68) is caused by P. attramentalis and P. rostrana.
Pococera attramentalis and P. attramentalis are also common pests of sorghum
(Kring et al., 1987) and other tropical fruit trees. The larvae of both species feed on
the axis of the inflorescence, petals and ovaries late in the flowering season; dried
fallen flowers are webbed together and fastened to flower clusters to form nests
(Patel et al., 1977). The Lepidoptera complex attacking mango flowers in Australia
consists of several species from the families Geometridae, Lymanthridae,
Noctuidae, Pyralidae and Tortricidae. In Brazil, Barbosa (2005) and Barbosa et al.
(2005) reported Pleuroprucha asth-enaria (Walker) (Lepidoptera: Geometridae)
and Cryptoblabes gnidiella (Milliere) (Lepidoptera: Pyralidae) affecting mango
inflorescences. The Pleuroprucha asthenaria life cycle from egg to adult is 17 days
and the C. gnidiella life cycle is 36 days (Barbosa, 2005). Pleuroprucha asthenaria
can cause premature ripen-ing of injured fruit while C. gnidiella infests
inflorescences that are compacted from paclobutrazol applications (Barbosa, 2005).

According to Schreiner (1987), Dipel® reduced caterpillar damage, but careful


monitoring or constant spraying was necessary to prevent significant damage. In
Brazil, the pesticides Bacillus thuringiensis, trichlorfon and lamb-dacyhalothrin
provided 66–59% mortality (Barbosa, 2005). Control with pes-ticides is mostly
unjustifiable in Florida USA and Australia, but regular monitoring is needed for
early detection of population increases (Cunning-ham, 1984; Peña, 1993).

Classical biological control of lepidopteran insects attacking mango in


Dominica was initiated with the introduction of the wasps, Aleiodes sp. and
Euplectrus sp., and the fly Blepharella lateralis Macquart. Populations of the pest
were reduced to 25% of pre-release levels; parasitization rates were 20–99%, with
Euplectrus sp. being the most abundant parasitoid (Nafus, 1991). The parasitoid
Macrocentrus prob. delicatus attacks P. attramentalis; however, the parasitism rate
is unknown (Peña, 1993). In Brazil, C. gnidiella is parasitized by Brachymeria
pseudoovata Blanch (Hymenoptera: Chalcididae).

Thrips
The western flower thrips, Frankliniella occidentalis (Pergande) damages flow-ers
and fruit in Israel (Wysoki et al., 1993). The developmental time of F. occi-
dentalis from egg to egg at 25C occurs between 14.8 and 16.65 days. The duration
of development of F. occidentalis from egg to adult is closely related to
environmental conditions, especially temperature. Frankliniella (possibly cubensis)
is present in mango flowers during the dry season in Costa Rica, requiring several
applications of systemic insecticides (Jirón, 1993). In Florida,
338 J.E. Peña et al.

the thrips complex consisting of Frankliniella bispinosa (Morgan) and F. kelliae


(Sakimura) is the most frequently observed blossom pest on flowers and causes
damage by ovipositing in the panicle and feeding on the floral necta-ries and
anthers, which may result in premature loss of pollen. The biology of F. bispinosa
has been reviewed by Watson (1917), and Sakimura (1981) studied the taxonomy
of F. kelliae. In South Africa, Grove et al. (2001) reported that Thrips acaciae
Thybom, Thrips tenellus Trybom and S. aurantii were the most abundant species
collected from mango flowers.

SAMPLING. Thrips density is related to flower phenology and the prevalent dry season in
Florida USA. Peña (1993) suggested that aerial trapping is superior to flower
inspection, but because there is no method for determin-ing the true population size, the
aerial trap method cannot be shown to be an unbiased estimator. In India, Verghese et
al. (1985) determined that the lower mango canopy is better for sampling, and
recommended a sample size of 55 panicles/tree for surveying. Verghese et al. (1988)
indicated that distribution of Thrips palmi (Karny) on mango panicles is better
explained by Iwao’s patchiness regression, which indicated an aggregated distribution
and suggested that the lower canopy should be sampled. Sample sizes should be 55
panicles/tree for control and survey studies, with a 20% error of the mean and 92
panicles/tree to obtain a lower (5%) percentage error of the mean. In Florida, Peña et al.
(2006a) showed that a cumulative number of 400–700 thrips/panicle during 4 weeks
causes 33–50% yield reduction of ‘Keitt’ mangoes.

CHEMICAL CONTROL. The efficacy of different pesticides (acetamiprid, fenpro-patrin,


milbemectin and zeta-cypermethrin, novaluron) was tested by Peña et al. (2006a)
against mango flower thrips. All treatments, except milbemec-tin, reduced thrips
densities 5 days after application of the first spray. Danitol and zeta-cypermethrin had
the lowest numbers of thrips 12 and 20 days after the first spray. All treatments had
lower thrips densities compared to the control at 28, 35 and 43 days after the second
application.

BIOLOGICAL CONTROL. Several parasitoids and predators, such as Ceranisus menes


(Walker) in Israel (Rubin and Kuslitizky, 1992) and the predators, Orius sp., Anystis
agilis Banks and Hypoaspis aculifer (Canestrini) (Loomans et al., 1995), are candidates
for biological control of F. occidentalis, while Dasyscapus parvipennis Gahan is
considered to have good potential for the biological con-trol of flower thrips in Puerto
Rico (Bartlet, 1938).

Pests of buds and leaves

Foliage feeders is one of the largest groups of injurious insects of mango. Pests of
mango buds and foliage may cause damage by reducing the photo-synthetic area of
the plant, thereby reducing the quantity of photosynthates translocated to the fruit.
The most destructive mango leaf feeders are thrips,
Pests 339

midges, mites, scales, whiteflies, mealybugs, weevils, ants, locusts and cater-pillars
(Jeppson et al., 1975; Bhole et al., 1987; Jadhav and Dalvi, 1987; Tigvatt-nanont,
1988). Formation of leaf galls in India is caused by the Eurytomidae, Mangoma
spinidorsum (Subba Rao, 1986), but there is little information on their importance
as foliage pests.

Thrips
The Mediterranean mango thrips, Scirtothrips mangiferae Priesner, is a severe pest
of mango in Israel, causing the young leaves to curl along the midrib, distorting
their shape and leading to premature drop (Wysoki et al., 1993). The twigs of
infested shoots are much shorter than those of uninfested ones. The population of
the thrips is low during winter, increases in early spring and reaches its peak during
summer (Wysoki et al., 1993). Yellow sticky traps can be used for monitoring
thrips densities. Ganz et al. (1990) established that an average population of ten
Mediterranean mango thrips per young shoot was the threshold above which
chemical control is required. Efficient control has been achieved by spray
application of fluvalinate or acephate (Ganz et al., 1990).
The red banded thrips, Selenothrips rubrocinctus (Giard), is an important pest
of cacao in the Caribbean islands and attacks mango and avocado in Australia and
Florida and Hawaii USA. The adults feed on the underside of leaves, causing
necrosis and subsequent leaf drop. According to Hill (1975), S. rubrocinctus is
only a pest in mango nurseries, and rarely damages mature trees. Its biology was
reviewed by Moznette (1922). Adult thrips are dark bodied with a red band on the
first abdominal segment. The immature stages are light orange with abdominal
segments one and two and the anal seg-ments bright red. The population of this
species peaks during the dry season and declines during the rainy season.
According to Yee (1987), the thrips are controlled by malathion (25% v/w).

The weaver ant, O. smaragdina (Hymenoptera: Formicidae) is considered an


effective biological control of S. rubrocinctus in the Northern Territory of Australia
(Peng and Christian, 2004).

Midges
Mango leaves are attacked by different Cecidomyiidae species, especially in Asia,
but also in the Caribbean region. Two genera, Procontarinia Kieffer and Cecconi
and Erosomyia Felt, are particularly associated with mango and all known species
have been reared from mango (USDA, 1981; Schreiner, 1991; Harris and
Schreiner, 1992; Uechi et al., 2002; Gagne and Medina, 2004). Prasad (1971)
described the biology of the main species attacking mango in India. A new species
of gall midge, Procontarinia schreineri Harris, which attacks mango foliage in
Guam, lays eggs on young mango leaves and larvae develop rapidly over c.5 days
and induce blister galls. According to Harris and Schreiner (1992), the main factors
affecting populations of this midge are rainfall and location. More galls are present
during rainy periods, possibly because RH improves larval and pupal survival. No
differences were observed in gall densities collected from lower and top portions of
the tree. Askari and Radjabi (2003) observed overlapping generations of
Procontarinia
340 J.E. Peña et al.

matteiana in Iran related to the different leaf flushing patterns and found that the
optimum pest temperatures were 10–26C. Differences on susceptibility of mango
cultivars to P. matteiana might indicate that susceptible cultivars should not be
grown in areas infested by this gall midge (Jhala et al., 1987; Githure et al., 1998).
Daneel et al. (2000) suggested that products to control P. matteiana should be
applied after harvest, coinciding with the first major flush and a second spray 6
weeks later. Austin (1984) and Sankaran and Mjeni (1989) have reported several
platygastrid species parasitizing Procontarinia spp. and their prospects for
biological control of the pest.

Mites
The mango bud mite, Aceria mangiferae (Sayed) (Acari: Eriophyidae), attacks
buds and inflorescences (Keifer et al., 1982; Ochoa et al., 1994) (Plates 69–71).
According to Jeppson et al. (1975) this mite stunts and brooms twigs, causing bud
proliferation and appears to be responsible for necrosis of bud tissue cells (Varma
et al., 1974). In Hawaii, Tegonotus mangiferae (Acari: Eriophyidae) feeds on the
underside of leaves (Jeppson et al., 1975), while another species, Metaculus
mangiferae (Attiah) (Acari: Eriophyidae) causes russeting of termi-nal leaves, buds
and inflorescences. The latter is an important pest in Egypt, India, Palestine and
Angola (Jeppson et al., 1975). The puncture wounds of several acarines (Acari:
Tetranychidae) cause serious damage to leaves, which may dry and fall. The main
pest in Mauritius, India, Egypt, Israel and Peru is Oligonychus mangiferus
(Rahman and Sapra); in Israel, the spider mite Tet-ranychus cinnabarinus
(Boisduval), which lives on the underside of the leaves, causes bronzing around the
puncture wounds. The adult life span of O. mangiferus is 10.11 days for females
and 4.21 days for males (Rai et al., 1988). The avocado red mites, Oligonychus
yothersi McGregor and Oligonychus puni-cae (Hirst), feed on the upper surface of
leaves and cause considerable sti-pling around the midrib at high population
densities (Andrews and Poe, 1980). If the tetranychid mites are sufficiently
abundant, infested leaves may drop. There is little or no information on sampling
techniques or for their economic thresholds.

Aceria mangiferae occurs wherever mango is grown (Denmark, 1983; Doreste,


1984). There has been controversy regarding a possible association between this
mite and floral and foliar galls, i.e. mango malformation (Sayed, 1946;
Narasimhan, 1959; Summanwar and Raychoudhury, 1968; Denmark, 1983; Ochoa
et al., 1994). However, A. mangiferae does not cause mango mal-formation, but
may be a carrier of Fusarium mangiferae, which is recognized as the causal agent
of mango malformation (Varma et al., 1974; Freeman et al., 2004). Aceria
mangiferae life history has been described by Abou-Awad (1981); it is completed
in 15 days at 25–27C.

DAMAGE. According to Keifer (cited in Jeppson et al., 1975), A. mangiferae infestation


of buds results in arrested growth, with the stunted, short, young stems close together at
the terminal branch. When the leaves fall, the overall effect is scanty growth of twiggy
branches, with a few stubby, short branch-lets and discoloured buds. The mite appears
to be responsible for necrosis of
Pests 341

bud tissue cells, which is initiated externally at the edge of the bud and pro-gresses
toward the centre and internal areas of the bud (Peña et al., 2005).

SAMPLING. Peña et al. (2005) reported preliminary results of the distribution and
sampling techniques for A. mangiferae. Taylor’s power law and Iwao’s patchiness
regression were used to analyse spatial distributions of the mango bud mite in mango
orchards. Taylor’s power law generally provided a better description of variance-mean
relationships for the species than did Iwao’s patchiness regression. The species
exhibited aggregated patterns of spatial distribution. More mites were found on apical
buds than on lateral-latent buds. Sample size requirements for fixed levels of precision
were determined by using variance-mean relationships (Peña et al., 2005). For a given
mean, and desired precision, different numbers of samples are required. For instance,
for a 10% precision and at 0.5 mites/bud, c.220 samples are required, whereas for a
30% precision at 0.5 mites/bud, only 25 samples are required. Sampling small
arthropods (i.e. mites) is operationally difficult and often time consuming. To ease this
burden, presence-absence, or binomial, sam-pling was tested in place of complete
counts for estimating or classifying densities of these organisms. Binomial sampling is
based on defining the pro-portion of one or more individuals – the percent of incidence
(the incidence P(I)) – and the density of animals (m) per sampling unit. At densities of
three and ten mites/bud, 71% and 42% buds, respectively, are considered unin-fested.
According to the equation, P(I) = 0.28 + 0.01 (mites/bud) an infesta-tion level of more
than ten mites/bud or a P(I) > 38% could be used as the nominal threshold (Peña et al.,
2005).

BIOLOGICAL CONTROL. The phytoseiid Amblyseius swirski Athias Henriot is associated


with A. mangiferae (Abou-Awad, 1981). In Florida USA, several unidentified
phytoseiids occur on buds infested with A. mangiferae. Tenuipal-pid (Brevipalpus
phoenicis), Tydeid and Tarsonemid (Tarsonemus confusus (Ewing)) mites also inhabit
mango buds. Therefore, it is difficult to deter-mine the mite species that is the host prey
(Peña, personal observation).

CHEMICAL CONTROL. Osman (1979) reported that applications of four full cov-erage
sprays of dichlorvos were effective for controlling A. mangiferae in Egypt. Rai et al.
(1966) cautioned that chemical control should be directed to apparently healthy and not
malformed tissues. In Florida USA, agrimek plus citrus oil, fenproximate and
fenpropathrin resulted in the lowest mite densi-ties 12 days after application. Agrimek
plus citrus oil, and acequinocyl resulted in the lowest mite densities 26 days after
treatment (Peña et al., 2005). In Brazil, Nascimento et al. (2002) recommended sulfur
applications.

PLANT RESISTANCE. In Florida USA, the densities of A. mangiferae were mea-sured on


22 mango cultivars from December 1997 to June 1998. ‘Keenan’, an unknown cultivar,
‘cv. 9819’, ‘Brander’ and ‘Bombay Green’ had significantly more mites than ‘Joellen’,
‘Duncan’, ‘Red Itamaraca’, ‘Smith’, ‘Wally’ and ‘Hindi’ (Peña et al., 2005).
342 J.E. Peña et al.

Scales
ARMORED SCALES. At least 26 species of diaspidids attack mangoes worldwide (Chua
and Wood, 1990). In India, Aspidiotus destructor Signoret causes serious damage,
while Parlatoria pergandii Comstock and Lepidosaphes gloverii (Pack-ard) damage 3-
year-old plants (cited in Chua and Wood, 1990). Radionaspis indica (Marlatt) (=
Leucaspis indica) encourages growth of black mould, which covers young branches
(Dekle, 1976). Several diaspidids, e.g. Aulacaspis mangiferae (tubercularis) Newstead,
attack shoots and leaves; the oleander scale (Plate 72) in Florida USA (Miller and
Davidson, 2005) and the mango scale in Ghana (van Halteren, 1970) cause similar
damage. They are damag-ing not only because they feed on sap, but also because of the
toxicity of their saliva (Singh, 1991). Scales inhabit both leaf surfaces and also are on
fruit. Van Halteren (1970) concluded that A. mangiferae development is completed in
35–40 days for females and 23–28 days for males.

SOFT SCALES. Other species of Coccidae, Coccus viridis (Green), Coccus longu-lus,
Ceroplastes actiniformis, Philephedra tuberculosa Nakahara and Gill and the mango
shield scales Milviscutulus mangiferae (Green) and Viusonia stellifera (Westwood) in
Asia, Africa, Australia, Israel and the Americas cause similar damage. These coccids
are generally polyphagous, attacking different genera and species. They are mobile and
injure mango because of the production of honeydew and the subsequent accumulation
of sooty mould on the honey-dew (Escalante, 1974; Silva and Cavalcante, 1977). Most
of these scales can be suppressed at sub-economic levels, either by application of
selective pesti-cides (i.e. oils) or by biological control agents.

SAMPLING. Because of their small size, it is laborious to sample all stages of scales.
Pheromones in tent-style traps have been used with other fruit crops, as well as
monitoring crawler movement using sticky bands close to infested leaves. Either
double-sided sticky tape, or tape coated with Vaseline ® is effec-tive for trapping
crawlers. Bands are removed and examined under a micro-scope to determine crawler
numbers. Dark-coloured tape provides a better contrast to detect crawlers (Beers et al.,
1993).

BIOLOGICAL CONTROL. In a survey of mango-producing areas in South Africa,


Labuschagne (1993) determined that the predatory thrips Auleurodothrips fasciapennis
Franklin and the parasitoid Aspidiotiphagus citrinus (Crawford) are the most important
biocontrol agents of A. tubercularis. In South Africa, Joubert et al. (2000) obtained
46% parasitism of A. tubercularis using an unidentified species of the parasitoid,
Aphytis sp. Arias et al. (2004a) observed Coccidophilus spp. (Coleoptera:
Coccinellidae) and Chrysopa spp. preying on A. tubercularis in Ecuador; the exotic
predator Cybocephalus nipponicus (Coleoptera: Nitidulidae) was introduced to
supplement predation of the former scale (Arias et al., 2004b). Several parasites have
been recorded in Israel parasitizing the mango shield scale: Coccophagus lycimnia
(Walker), C. eritraensis Compere, C. scutellaris (Dalman), C. bivittatus (Compere),
Pests 343

Microterys flavus (Howard) and Metaphicus flavus Howard. Usually no chem-ical


control is required for this scale in Israel due to the activity of natural enemies
(Kfir and Rosen, 1980). Natural enemies for the control of the pink wax scale
Ceroplastes rubens Maskell in Australia include the parasitic wasps Anicetus
beneficus Ishii and Yasumatsu, Aenasioidea varia Girault and Rhopal-encyrtoidea
dubia Girault (Cunningham, 1984).
Whiteflies and blackflies
The two fly species of economic importance are the whitefly, Aleurodicus dis-
persus Russel and the blackfly, Aleurocanthus woglumi Ashby. The whiteflies suck
cell sap from leaves, which wilt when whitefly populations are high. High
infestations can almost blacken entire trees, reducing photosynthetic efficiency and
causing defoliation (Angeles et al., 1971; Peña, 1993). A number of parasitoids,
e.g. Encarsia opulenta (Silvestri) and Amitus hesperidus (Silves-tri), attack the
immature stages and provide good control.
Mealybugs
Mealybugs injure mango by sucking sap through their stylets, and excreting large
amounts of honeydew onto fruit and leaves. Sooty mould fungus growth on the
honeydew can render the fruit unmarketable, and reduce the photosynthetic
efficiency of leaves and cause leaf drop (CAB International, 2003). The
margarodid mango mealybug Drossicha stebbingi (Green) is a seri-ous pest of
mango in India and Pakistan (Prasad and Singh, 1976; Mohyud-din, 1981;
Mohyuddin and Mahmood, 1993). It is univoltine. After mating, females enter the
soil in June and die after laying eggs at a depth of up to 15 cm. These begin to
hatch at the end of December or early January. The nymphs emerge from the soil
and move to tender shoots where they settle. Prasad and Singh (1976) reported that
the intensity of attack varied with respect to year and locality in India, probably
because of soil and environ-mental conditions. Moderate rainfall (55–60 mm)
during oviposition and dry conditions during hatching appear to favour
development. Adults develop in April. They mate, and males die soon afterwards.
The females enter the soil in May for oviposition, and the diapausing eggs remain
in the soil until the end of December.

The pseudococcid fruit tree mealybug, Rastrococcus invadens Williams, is a


serious pest of several crops, including mango in West Africa (Agounké et al.,
1988). Mealybugs feed on leaves and fruit. Females have three moults and males
have four. The entire life cycle can be completed in 31–84 days. The mealybugs
weaken plants by puncturing the tissues and consuming sap, but the major damage
is caused by the production of large amounts of honeydew upon which saprophitic
fungi develop. The resultant thick black layer of sooty mould causes a drastic
reduction in photosynthetic efficiency, resulting in premature leaf drop.
Rastrococcus invadens severely reduces fruit produc-tion in some areas of Africa
(Moore and Cross, 1992).
SAMPLING. Boavida et al. (1992) devised sampling plans for R. invadens, but advised
that the sampling strategy was only practical for estimating medium to high mealybug
populations in the field. Narasimham and Chacko (1991)
344 J.E. Peña et al.

determined that densities of R. invadens Williams, Rastrococcus iceryodes (Green)


and Rastrococcus mangiferae (Green) were significantly higher on the abaxial than
the adaxial surface. Mealybug density was also higher from ground level to 2 m
compared with >2 m; they also observed that spatial distribution of R. iceryodes
did not differ among internal and external cano-pies, whereas densities of R.
invadens and R. mangiferae were higher in the external canopy. They also
determined that there were statistical differences causing some inter-tree variation,
but did not determine if the mealybugs followed a random or contagious
distribution.

CONTROL. Various control methods, including banding tree trunks with vari-ous
materials to prevent D. stebbingi nymphs from climbing (Lakra et al., 1980; Srivastava,
1981) and dusting chlorinated hydrocarbons on the soil (Srivastava, 1981), have been
tried with little success. In Pakistan the mango mealybug was controlled by hoeing or
ploughing the soil and conservation of the predator, Sumnius renardi Weise, by
wrapping burlap around the trunks of the trees (Mohyuddin and Mahmood, 1993).

Pests of trunks, twigs and roots

Coleopterans and scales


Stem-boring Coleoptera and scales as a group of injurious mango insects have not
been studied in great detail (van Whervin, 1968; Woodruff, 1985). The wide host
range of borers and overlapping borer species has compli-cated their study.
Tunnelling of borer larvae in branches and trunks of mango and the slow feeding of
some scale species in certain seasons and regions may cause serious reductions in
yields and might also contribute to mango decline. However, borer occurrence and
injury tends to be sporadic and below levels requiring direct action. Few natural
enemies have been reported for suppression of borer populations in mango.
Detailed coverage of stem-boring species is beyond the scope of this chapter;
however, Hypocryphalus mangiferae (Stebbing), Apate monachus Fabricius and
Batocera rubus L. are pests within this group.

Infestations of the mango scale, Radionaspis indica (Marlatt), and plumose


scale, Morganella longispina, commonly occur on the trunk, branches and buds.
Severe infestations include cracking of bark, exudation of sap and decline of the
upper branches. Peña (1993) demonstrated that branches with both species of scale
showed more decline symptoms than branches with low-scale density. Research on
the bionomics and control of these scales is necessary to confirm their role in
mango decline symptomatology.
The scolytids, Hypocryphalus mangiferae (Stebbing) and Xylosandrus com-
pactus (Eichoff) directly attack the main stem and branches (Silveira, 1960;
Wysoki et al., 1993). Growth of fungal mycelia can extend terminally and basally
from the beetle gallery in the mango tree and can kill the affected branches. The
insects prefer trees that have been weakened by pathogens, wind, etc., but after a
population has been established, the infestation spreads
Pests 345

to healthy trees. Hypocryphalus mangiferae has been associated with mango wilt
disease in Brazil and Oman (van Wyk et al., 2007). Following initial trap-ping
results from Berti Filho and Fletchman (1986), Rocha da Silva (2006) established
that H. mangiferae is attracted to trees where the fungus is pres-ent. Scolytid
beetles are attracted to mango trees in response to visual stimuli, to host-specific
chemicals and to species-specific aggregation pheromones (Lindgreen et al., 1982).
The evaluation of traps as tools for managing ambro-sia beetles on mangoes in
Florida USA is necessary in order to reduce their damage in newly established
groves.

Termites
Mango orchards are becoming more common in dry and semi-arid areas with vast
termite populations. Mango growing in infested areas often results in plant growth
suppression as a result of reduced root establishment, inva-sion and pruning of
roots (Rogers et al., 1999). For example, six termite spe-cies (Odontotermes
indicus Thakur, O. lokanandi Chatterjee and Thakur, O. obesus (Rambur), O.
giriensis (Roonwal and Chhotani), O. bhagwatii Chatter-jee and Thakur and
Microtermes obesi Holm) were recorded from mango orchards in India (Srivastava
and Singh, 2004). More species of termites were observed in winter than during the
summer and rainy season. Veeresh et al. (1989) observed that O. wallonensis, O.
horni and O. obesus constructed earthen sheeting on the stem of small mango trees.
Singh (1960), cited by Srivastava (1998), reports Eutermes (Nasutitermes) costali,
Calutermes (Cryptotermes) bre-bis, Heterotermes tenuis, Coptotermes gestroi,
Neotermes (Kelatermes) bosei (Gard-neri) Synder, Microcereoutermes peroffinis,
Calotermes (Neotermes) greeni and Coptotermes curvignathus also affecting
mango in India. According to Srivas-tava (1998), the most important termite
species affecting mango are O. obesus and O. wallonensis; O. wallonensis nests in
the root zone in Uttar Pradesh, India. The workers feed on roots, stems and
branches.
Colonies of the subterranean termite, Coptotermes curvignathus Holmgren,
were monitored in Malaysia using bait matrices containing 0.5% hexaflu-muron
(Said Sajap et al., 2000). In Florida, urban dwellings infested with the Formosan
termite, C. formosanus Shiraki, were treated with baits containing 0.5%
weight/weight noviflumuron (Cabrera and Thoms, 2006). These baits might be
useful when mango orchards are planned for areas infested with termites. In India,
termite infestations are controlled with a combination of monthly irrigation, hoeing
and application of neem cake (Singh and Singh, 2003).

10.3 Discussion
In general, most mango pests also occur on other fruit crop species. Fruit flies,
scales, mites, thrips, lepidopteran flower feeders, mirids, weevils and beetles are
mostly generalists, and some of their management schemes need to be
implemented with this in mind. In the case of fruit flies, Aluja (1996) suggests
surveying vegetation adjacent to infested mango orchards as
346 J.E. Peña et al.

populations are sustained and multiplied in these locations and from them adult
flies move into commercial orchards to attack ripening fruit (Aluja et al., 1996).
Management of key pests (i.e. fruit flies, seed weevils, etc.) must be mandatory, in
order to have an effect on a large region. The use of some mea-sures (i.e.
quarantine, etc.) must involve neighbouring producing countries in order to have a
positive effect on sanitation. The most progressive exam-ples in management of
mango pests are in Australia, Mexico and Israel, while other producing countries,
such as South Africa, Brazil and Ecuador, are tak-ing serious steps to reconcile the
opposing forces of globalization of markets and sustainibility. For other areas
where maximizing yields and blemish-free fruit is not a priority, the emphasis
should be biological control. Management tactics that can be improved include the
following:

1. Selective pesticides. Pesticides that are used in integrated pest man-agement


programmes must have selective toxicity. The current trend is the development of
chemicals that are highly effective for a limited group of in-sects. Díaz-Fleischer et
al. (1996) suggested the use of cyromazine to reduce fertility of A. obliqua.
Cunningham (1989) suggested that oils could be uti-lized for control of scales in
mango; however, most of the recommendations are based on highly toxic or illegal,
non-registered persistent chemicals (Singh, 1991; van Mele et al., 2001; de Bie,
2004). In South Africa, Joubert et al. (2004) tested kaolin, which is the active
ingredient of Surround®, a non-toxic natural clay mineral, against the mango seed
weevil, mango scale and citrus thrips. Surround® was effective against citrus
thrips, mango weevil and co-conut bug, P. wayi, but caused outbreaks of mango
scale and long-tailed mealybug, Pseudococcus longispinus. However, producers of
export fruit rely on calendar-based chemical control when trees are heavily infested
with mango scale (Joubert et al., 2004).

Biological control. Biological control has great potential as a tactic for regulating
pest populations in integrated pest management programmes in mango orchards.
However, it will be difficult for biological control alone to reduce a pest from an
economic to a completely non-economic status for pests attacking fruit. A
combination of augmentative releases of parasi-toids and the use of sterile insects,
at least from a theoretical perspective, has been considered to be more effective for
fruit flies than either method applied alone (Barclay, 1987). Biological control
should be highly effective for indirect pests. Indeed, numerous studies have been
conducted in many mango-producing countries to promote the use of parasites and
predators for this type of pest (Cunningham, 1989; Mohyuddin and Mahmood,
1993; Moore and Cross, 1993; Whitwell, 1993; Wysoki et al., 1993; Labuschagne
et al., 1995).

2. Host plant resistance. Tolerance of mango to pests is mentioned for Noorda sp.
and Idioscopus sp. (Bagle and Prasad, 1984; Cunningham, 1989), while man-go
resistance to Sternochetum mangiferae is mentioned by Hansen (1993). Carvalho et
al. (1996) have also demonstrated the different degrees of suscepti-bility of mango
cultivars to A. obliqua. Most of this research, however, needs
Pests 347

to be assessed further. Angeles (1991) reported that Mangifera altissima does not
seem to be affected by leaf hoppers, tip borers and seed borers in the Philippines.
There is little doubt that wild mangoes have potential in breed-ing. Determining the
tolerance or insect resistance of mango cultivars and related species should be done
in natural stands or in established germplasm collections. After the initial selection
has been made, evidence for the pattern of resistance must be established and
changes in the environment, whether geographic or temporal, should not disrupt or
decrease the resistance to any great extent. Therefore, tests for resistance in mango
to insects should in-clude provision for exposure to insects under varying
conditions whenever possible.

3. Pheromones and trapping devices. Developments in the identification and


synthesis of sex pheromones have resulted in their possible use for pest
management in mango orchards (Chu et al., 1994; Khan et al., 2002, 2005; Sheikh
et al., 2008). Food attractants, however, remain the most common monitoring
tools. Trapping techniques can be utilized to reduce pesticide use by improving
timing of sprays as a result of better monitoring of pest popu-lations. It remains
uncertain if trapping techniques can be used to predict infestations by fruit-feeding
pests and if they can be used for direct control (by mass trapping) over several
years.
4. Cultural and physical control. Use of cultural and physical techniques (i.e.
pruning, bagging, etc.) depends on costs of control, availability of techni-cal
assistance and market purposes. Paderes and Orden (2004) observed that bagging
and pruning of ‘Manila’ in the Philippines is mostly influenced by the availability
of technical assistance for growers.
Greatly increased regulation of pesticides, heightened public aware-ness of
environmental contamination, pesticide resistance problems in pests and the high
cost of chemical pest control has resulted in increasing reliance on integrated pest
control as an important strategy in sustainable agriculture.

Acknowledgements
We are particularly indebted to Walther Enkerlin (Joint IAEA/FAO Division,
Vienna), Andrea Birke-Biewendt, Alberto Anzures-Dadda and Larissa Guillén-
Conde (Instituto de Ecología, AC) and Aldo Malavasi (private consultant) for their
assistance and for providing many valuable references. M. Aluja acknowledges the
financial support of the Mexican Campaña Nacional Contra Moscas de la Fruta
(Convenio SAGARPA-IICA-INECOL) and the Instituto de Ecología, AC. He also
acknowledges support from CONACyT through a Sabbatical Year Fellowship
(Ref. 79449) and thanks Benno Graf and Jörg Samietz (Forschungsanstalt
Agroscope Changins-Wädenswil ACW), for pro-viding ideal working conditions
during the final phase of the publication process of this chapter.
348 J.E. Peña et al.

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11 Crop Production: Propagation
S. Ram1 and R.E. Litz2
1
GB Pant University of Agriculture and Technology, Pantnagar, India
2
University of Florida, Florida, USA

11.1 Introduction 367


11.2 Seed Propagation 369
Monoembryonic seed 369
Polyembryonic seed 369
11.3 Vegetative Propagation 374
Preparation of rootstock 376
Attached methods 378
Detached methods of grafting 380
Effect of rootstock 385
Top and double working 386
Rooting 387
Micropropagation 391
11.4 Comparative Performance of Trees Propagated by Different Methods 391
11.5 Conclusions 392

11.1 Introduction
Mango reproduces naturally by seed, although this is rarely a horticultural practice,
particularly for monoembryonic cultivars. Instead, vegetative prop-agation is
utilized to preserve the unique phenotypes of superior selections, and has been
based upon grafting and rooting methods, growing plants from nucellar seedlings
of polyembryonic mangoes and micropropagation (Fig. 11.1). Grafting of mango
can be either attached or detached. In the former, the scion is not severed from the
mother plant until its union with the rootstock is complete, i.e. approach grafting,
tongue, saddle and root grafting. In the latter, the scion is removed from the mother
tree and then joined with the rootstock, and both are allowed to grow prior to
cutting of the rootstock above the graft union. Detached methods include rind or
crown grafting, cleft or wedge grafting, whip or splice grafting, side grafting,
veneer grafting,
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 367
368
Propagation

Sexual Asexual

Monoembryonic Polyembryonic
Grafting Rooting Micropropagation
seed seed

Attached Detached Layering Cutting Organogenesis

Somatic
Approach Cleft or wedge Ground
embryogenesis

S. Ram and R.E. Litz


Tongue Splice or whip Pot Shoot tip culture

Saddle Side Air

Root Notch or inlaying

Veneer

Rind or crown

Budding

Fig. 11.1. Methods of mango propagation.


Crop Production: Propagation 369

notch or inlaying, ‘T’ or shield budding, forkert budding, modified forkert budding,
patch budding, modified patch budding, chip budding, etc. Root-ing methods
include layering and cutting techniques. Although in vitro meth-ods for
regenerating mango have been reported via somatic embryogenesis, organogenesis
and shoot tip culture (see Litz et al., Chapter 18, this volume), their practical
application for propagation has not yet been demonstrated. The various methods of
mango propagation are shown in Figs 11.2–11.8. Mango propagation has been
discussed recently by Singh and Singh (1998), Galán-Saúco (1999) and Neto et al.
(2002).

11.2 Seed Propagation

Monoembryonic seed

Seed propagation does not ensure true-to-type plant reproduction of


monoembryonic mango selections; however, it was extensively used before
vegetative methods for mango propagation were known (Singh, 1960). Large
seedling orchards were planted during the medieval period of Indian history until
vegetative propagation was introduced into India by the Portuguese in the late 15th
century. Monoembryonic seeds contain only a single sexual embryo, and a single
plant grows from a seed of a monoembryonic cultivar.

Polyembryonic seed

Polyembryony is common in mango cultivars that originated and are grown in the
moist tropics. In Knight et al. (Chapter 3, this volume), there is a survey of the
major polyembryonic and monoembryonic mango cultivars. Trees from nucellar
seedlings are identical to the mother plant (see Mukherjee and Litz, Chapter 1, this
volume). In polyembryonic seed, only one embryo is zygotic in origin; it either
degenerates or produces weak and stunted seed-lings (Maheshwari et al., 1955;
Sachar and Chopra, 1957; Singh, 1960; Stur-rock, 1968). Approximately three to
eight seedlings normally originate from a single polyembryonic seed (Garner and
Chaudhri, 1976), although 30 or more embryos have been recorded in a single
polyembryonic mango seed (Juliano, 1934, 1937). Nucellar seedlings can be
distinguished from the zygotic seedling on the basis of their greater vigour c.1
month after germination.
Polyembryonic mango cultivars have traditionally been seed propagated in
South-east Asia and in some tropical Latin American countries. In many mango-
growing areas, polyembryonic and monoembryonic mango selec-tions are grafted
onto uniform, nucellar seedling rootstock. Zygotic and nucellar seedlings can be
morphologically similar. Therefore, detection and separation of nucellar seedlings
is important in breeding programmes involv-ing controlled pollinations. Different
protein and DNA markers have been used. Degani et al. (1993) and Schnell and
Knight (1993) demonstrated that isozymes and RAPD markers can be used to
distinguish zygotic embryos from
370 S. Ram and R.E. Litz

(a) Approach grafting

(b) Tongue grafting

(c) Saddle grafting

(d) Root grafting

Fig. 11.2. Grafting methods of mango propagation: (a) approach grafting; (b)
tongue grafting; (c) saddle grafting; (d) root grafting.
Crop Production: Propagation 371

(a) Cleft grafting

(b) Soft wood grafting

(c) Epicotyl grafting

(d) Splice (whip) grafting

Fig. 11.3. Grafting methods of mango propagation: (a) cleft grafting; (b) soft
wood grafting; (c) epicotyl grafting; (d) splice or whip grafting.
372 S. Ram and R.E. Litz

(a) Side grafting

(b) Notch grafting

(c) Veneer grafting

(d) Rind or crown grafting

Fig. 11.4. Grafting methods of mango propagation: (a) side grafting; (b) notch
grafting or inlaying; (c) veneer grafting; (d) rind or crown grafting.

nucellar seedlings. Eiadthong et al. (1999), using single sequence repeats (SSRs)
could distinguish among mango cultivars, and separate them according to their
geographic origin; however, monoembryonic and polyembryonic geno-types could
not be differentiated. Amplified fragment length polymorphism (AFLP) markers
have been used for cultivar identification (Kashkush et al.,
Crop Production: Propagation 373

(a) Shield budding

(b) Patch budding

(c) Modified patch budding

(d) Forkert budding

Fig. 11.5. Budding methods of mango propagation: (a) shield budding; (b)
patch budding; (c) modified patch budding; (d) forkert budding.
374 S. Ram and R.E. Litz

(a) Modified forkert budding

(b) H-budding

(c) Chip budding

Fig. 11.6. Budding methods of mango propagation: (a) modified forkert


budding; (b) H-budding; (c) chip budding.

2001) and Gonzalez et al. (2002) identified a range of inter-simple-sequence-


repeats (ISSR) primer sequences with potential for cultivar identification.

11.3 Vegetative Propagation


Vegetative or asexual propagation preserves the genotype and phenotype of elite
selections by means of grafting, rooting and micropropagation. Trees
Crop Production: Propagation 375

(a) Modified chip budding

(b) Flap budding

(c) Window budding

Fig. 11.7. Budding methods of mango propagation: (a) modified chip


budding; (b) flap budding; (c) window budding.

propagated by grafting onto seedling rootstocks usually flower after 3 or 4 years in


comparison to the 5–10 years for seedling trees and they are gener-ally smaller
than seedling trees because they begin to bear fruit earlier. The oldest method for
vegetatively propagating mango is probably the attached method, also known as
inarching or approach grafting.
376 S. Ram and R.E. Litz

(a) Top working

(b) Double working

Fig. 11.8. Top working (a) and double working (b) of mango.

Preparation of rootstock

Rootstocks are either zygotic or nucellar seedlings. Nucellar seedling root-stocks


are clonal, and have many advantages over heterogeneous monoem-bryonic
seedlings as rootstocks (see Crane et al., Chapter 13, this volume). Clonal
rootstocks have generally been selected for specific soil types and
Crop Production: Propagation 377

stress tolerance, and behaviour of the scion cultivar on clonal rootstock is highly
predictable. Monoembryonic seedlings are generally used as rootstocks in India.
Polyembryonic ‘Turpentine’ and ‘Kensington Pride’ seedlings are used as
rootstocks in Florida and Australia, respectively, whereas polyembryonic ‘Sabre’,
‘13-1’ and ‘4-9’ seedlings are used for rootstocks in Israel. Throughout South-east
Asia, polyembryonic seedlings are used for rootstocks, e.g. poly-embryonic ‘Saing’
and ‘Thalapt’ in Myanmar (Grant and William, 1949). In Senegal, polyembryonic
‘Amelioree’, ‘De Cameron’ and ‘Madame Francis’ are used as rootstocks (Furon,
1966). In Brazil, seedlings of polyembryonic ‘Carabao’, ‘Mangua d’Agua’ and
‘Pico’ are used for rootstock; they are consid-ered to have resistance to
Ceratocystis fimbriata (Neto et al., 2002); the IAC series (IAC100-104) are used to
control ‘seca de mangueira’ (Neto et al., 2002).
Freshly extracted mango seeds from ripe fruits germinate with higher
frequency (76–91%) than those from overripe, firm or green fruits (Shant and
Saproo, 1974). Seeds with large cotyledons from seedling trees germinate earlier,
store better and seedling growth is more vigorous than seeds with small cotyledons
(Naik, 1949; Simao, 1960). Naik (1941) noted differences in germination and
vigour in monoembryonic seedling progenies of different parentages. Seedlings of
‘Chausa’ reportedly produced better height and girth than ‘Langra’ and seedling
vigour was closely related to the weight of the seed stone (Giri and Chaudhery,
1966). Mango seeds are recalcitrant, and lose viability (Ledin and Ruehle, 1954;
Ruehle and Ledin, 1955; Singh, 1960) within 30 days. Therefore, seed should be
collected and sown within 1 week after collection. About 80% of seed germination
occurs if they are sown within 1 month of extraction (Stephens, 1960; Sturrock,
1968; Chacko and Singh, 1971). Parisot (1988) recorded that seeds that are free of
pulp could be stored for up to 84 days at 15C on sterile cotton with deionized
water. Husk-ing or decortication of the hard endocarp of the seed improves
germination (Paul and Guneratnam, 1938; Simao, 1960; Chauran et al., 1979;
Abdel-Galil, 1992). At the time of sowing, treatment with fungicides also improves
germi-nation, particularly of dehusked seeds (Chauran et al., 1979). Seeds should
be sown without injuring the cotyledons and preferably early in the season.

Seedbed preparation
The standard practice for seed germination varies. In India, seedbeds are used,
whereas in most other countries, seeds are planted individually in pots or plastic
bags. Nurseries are usually under partial shade to prevent exces-sive evaporation
from the plant and soil and to protect seedlings from inclem-ent weather; shade is
particularly useful in areas with dry, hot climates. Seedbeds can also be used in the
open provided the soil is adequately moist. Highly sandy or loamy soils are
generally unsuitable for mango seed germi-nation. Soil should be well drained with
plenty of organic matter, and seeds should be sown c.15 cm apart in a special
seedbed in which 25 cm top soil of the bed is excavated and the bed bottom is
hardened with concrete or lined with an iron or polyethylene sheet to prevent
elongation of the taproot and to encourage fibrous root development (Stephens,
1948). Germination can be expedited if seeds are soaked in 20–100 mg/l
gibberellic acid (GA3) for 24–48 h
378 S. Ram and R.E. Litz

or sprayed with 50–300 mg/l GA3; however, these seedlings are usually
unsatisfactory for grafting. Seeds should be planted with convex edge up at a depth
of 5–7 cm (Singh, 1960; Bakhshi, 1963) or with a small portion exposed above the
ground level (Pope, 1929; Ruehle and Ledin, 1955; Bakhshi, 1963; Anonymous,
1975). Soil moisture should be maintained. Seeds germinate 2–3 weeks after
planting (Ruehle and Ledin, 1955; Ahmed and Rashid, 1961; Singh and Jawanda,
1962). Seedlings that are 1-month-old with coppery leaves and with their stones
attached survive transplanting better than older plants without much damage to
their root system (Ruehle and Ledin, 1955). The taproot should be slightly pruned
at the time of planting.

Nursery
Seedlings should be transplanted without disturbing the roots to nursery beds or
pots at least 1 month before grafting. Either the crown of the seed-lings should be
pruned or the distal half of the leaves should be removed to reduce transpiration.
Seedbed-grown plants generally have better germina-tion rates and lower
production costs than those grown directly in nursery beds. Seedlings should be
actively growing at the time of grafting. Success in grafting depends on several
factors, such as age and vigour of rootstock and scion, diseases, insect pests,
environment and the skill of the grafter.

Attached methods

Approach grafting is expensive, cumbersome, requires a long time to pro-duce


grafts and the success rate is often low. Other methods are now used because they
are cheaper and easier.

Approach grafting
A scion shoot, while still attached to the parent plant, can be grafted onto the
seedling rootstock by making a long cut (5–7 cm) of similar length and depth on
one side of both rootstock and scion through the cambium and slightly into the
wood (Fig. 11.2a). The cut surfaces of rootstock and scion are pressed firmly
together and secured with waxed string, raffia or grafting tape. After union, the
scion is severed below the graft union and the rootstock above the union. Several
years ago, approach grafting was standard practice for propa-gating mango in many
countries, but has been largely replaced by detached grafting methods, except for
parts of India. For approach grafting, seedlings 45–60 cm long with a diameter of
1.25 cm are utilized (Asadullah and Khan, 1960; Singh, 1960). They are planted in
pots or plastic bags containing soil mixture, and are kept beneath the mother tree.
Alternatively, the ball of earth around the root system of the seedling grown for
rootstock is tied in ‘jutties’ made of dried grass, paddy straw or plastic bags and
planted under the mother tree or raised to the shoot height of the mother tree on a
raised plat-form or directly suspended from branches of the mother tree. The
seedling rootstock can be a few months- to 2-years-old (Naik, 1949; Garg, 1954;
Ahmed, 1960; Singh, 1960). Juvenile seedlings whose leaves are copper coloured
and
Crop Production: Propagation 379

with the seed still attached to the seedling are often used. The root system is
covered with moist sphagnum moss or similar material and wrapped in
polyethylene to prevent drying. Production costs are less with young seed-lings
than with older material.
In the subtropics, approach grafting in spring can achieve 88–100% suc-cess
with some cultivars (Asadullah and Khan, 1960; Majhail and Singh, 1962a, b;
Talukdar and Ahmed, 1965). Majhail and Singh (1962b) observed that seedling
size and age do not affect grafting success in spring, but that larger seedlings
yielded better success from July to September. Approach grafting should be
avoided during winter months when plants are dormant. In the tropics, mangoes
can be approach grafted year-round (Ahmed, 1960; Singh, 1960). Approach
grafting should be done when the trees are in active growth (Singh, 1960; Rao,
1967). In low rainfall areas, approach grafting should be done at the time of the
onset of rains, while in regions of heavy rainfall, it should be done soon after the
rainy season is over, provided tem-peratures are not <15C, the critical temperature
for the growth of mango trees (Whiley, 1993). Healthy trees should be used for
scion wood. The suc-cess of approach grafting is cultivar dependent, for example
‘Langra’ scions establishes better than either ‘Dashehari’ or ‘Chausa’ (Asadullah
and Khan, 1960). Better success has also been reported with seedlings of 1.3–1.6
cm diameter than with smaller shoots (Giri, 1966). Variations of approach graft-ing
include multistock, inflorescence, top working and adjuvant grafting. At one time,
south Indian nurserymen would approach graft a single large scion with two to five
rootstocks (Patwardhan and Deshmukh, 1931; Singh, 1960). Grafts with large and
old scions having several branches become top heavy, and fail to thrive. New
growth from inflorescences that are approach grafted is veg-etative. Approach
grafting has also been used to top work old seedling trees with cultivars (Singh,
1960). Adjuvant grafting refers to approach grafting of diseased or damaged trees
in order to rejuvenate them on new rootstocks.

Tongue grafting
Tongue grafting is practised with thicker rootstock than scion. The rootstock is first
cut as in approach grafting; a second cut is made into the wood of the stem
approximately halfway down the first cut in a sloping direction point-ing tongue
upwards. A similar cut is made in the scion shoot. Both cuts are made in such a
manner that one fits into the other tightly (Fig. 11.2b). Root-stock and scion are
tied together with grafting tape and a graft union is com-plete in c.1–2 months. The
scion is then severed from the mother tree and the rootstock is decapitated above
the graft union. Tongue grafting is slightly more complicated than simple approach
grafting; however, the success rate is better because three surfaces of the cambium
layer are involved in the graft union (Burns and Prayag, 1921).

Saddle grafting
With saddle grafting, the rootstock is decapitated c.25 cm above ground level, and
two upward sloping cuts are made on the sides of the rootstock to form a 5–7 cm
long wedge on its cut end. On the stem of the scion shoot, a tongue
380 S. Ram and R.E. Litz

is made and the outer surface of the tongue is not pared. The wedge of the
rootstock is then fitted into the groove of the tongue (Fig. 11.2c). Subsequently, the
joint is secured with grafting tape and the scion shoot is separated from the mother
tree after the union is complete (Singh, 1960). Saddle grafting is easier to perform
than tongue grafting but is more difficult than approach grafting.

Root grafting
Root grafting is similar to bench grafting. A seedling plant (c.l year old) is potted
in a U-shaped pot with a notched edge (7–8 cm deep and 5 cm wide) (Singh, 1960).
The 7–10 cm taproot is projected through the notch and the pot itself contains the
root. The seedling above the collar is staked, and placed in the shade for 1–1.5
months for establishment. Seedlings are approach grafted with the scion (Fig.
11.2d), and the graft is separated from the mother tree after the union is complete.
Grafts are maintained in the shade and watered regularly. Naik (1941, 1949)
reported 100% success with root grafting.

Detached methods of grafting

Cleft or wedge grafting


Singh (1960) suggested that cleft grafting can be used with rootstocks of greater
diameter than the scion and also used for top working (Fig. 11.3a). Rootstock of 5–
7 cm or more in diameter should be used and cleft grafted after decapitating the
stock c.45 cm above the ground. The decapitated root-stock is split to a depth of
about 5 cm through the centre of the stem with a knife and a mallet. After the knife
is removed, a hard wooden wedge is inserted to keep it open for the subsequent
insertion of the scion. Scions (15–20 cm long) from a terminal shoot >3 months old
is wedged securely (to a depth of 6–7 cm). The cleft of the scion is slipped into the
split of the stock that has been forced open with the wooden wedge. The thicker
edge of the cleft is placed towards the outer edge of the rootstock in such a way
that the back of the rootstock and scion meet firmly. Another scion is also inserted
diametrically opposite the first one. The wooden wedge is removed and cut
surfaces are sealed with grafting wax. The graft union requires c.2 months. Cleft or
wedge grafting is appropriate for replacing the crown of young trees; however,
with young seedling rootstock, this has also been used for large-scale propagation.
In Brazil, Pinheiro et al. (1970) reported that cleft grafting was more successful
(97%) than four other grafting methods tried. Cleft grafting is easier to use than
veneer grafting and has a similar rate of success (Ram, 1993). Success can be
improved when leaves are retained below the point where the rootstock is severed,
when the grafting portion of the rootstock is a new flush and when the stem is
pinkish green; however, the scion should be mature.

Soft wood grafting


Insertion of the scion into the bronze-coloured stem of the rootstock has been
referred to as soft wood grafting (Fig. 11.3b). Amin (1974, 1978) reported 100%
success if leaves are not removed below the graft union. Seedling rootstocks
Crop Production: Propagation 381

of various ages can be used; however, success decreases with age, i.e. 80% with 1-
year-old rootstocks and less with 6-month-old seedlings (Gaur, 1984; Reddy and
Melanta, 1988; Kulwal and Tayde, 1989; Panickar and Desai, 1989; Gandhoke,
1993). This technique is easy and is effective in dry, hot weather and in areas with
low precipitation.

Epicotyl or stone grafting


Epicotyl grafting involves cleft grafting of scions onto 2-week-old seedling
rootstock (Traub and Auchter, 1934). Predefoliation of the scion is not essen-tial,
although it is widely practised (Fig. 11.3c). Moderate temperature and high relative
humidity (RH) are important for success. A 2–3 cm long slant-ing cut is made in
the epicotyl with a matching cut on the proximal portion of the scion in the splice
method for grafting. For wedge grafting, a vertical cut 4–5 cm long is made in the
decapitated epicotyl to receive the wedge-shaped scion. The scion and rootstock
are tied together with grafting tape, and the small grafted plant is planted
immediately and watered. The scions sprout within a month. The success rates of
splice and wedge grafting meth-ods are reported to be >50% (Bhan et al., 1969;
Gupta et al., 1988; Dhunaga et al., 1989; Gunjate, 1989; Jinturkar and Narwadkar,
1989; Kulwal and Tayde, 1989; Madalgeri et al., 1989; Narwadkar and
Anserwadekar, 1989; Radhamony et al., 1989; Singh et al., 1989; Srivastava, 1989;
Patil et al., 1991); under green-house conditions in the subtropics, success can be
>90% (Ram, unpublished). Grafted plants are maintained under partial shade in a
moderate and moist environment. In the subtropics, growth is initially slow and can
require c.2 years to attain the size of a veneer graft; however, in the tropics, growth
is much more rapid.

Splice or whip grafting


This is the easiest method for propagating mango and is widely used in the
subtropics. The rootstock should be c.1–2 years old, the diameter of both rootstock
and scion should be 0.6–1.0 cm. The rootstock is severed c.10–20 cm above the
soil and a diagonal cut (3–4 cm long) is made at the distal end of the rootstock (Fig.
11.3d). A similar slanting cut is made on the proximal end of the scion and the cut
surfaces of both the rootstock and the scion are bound together with grafting tape.
The union (usually 90% successful) takes place in c.2 months (Ahmed, 1974).
Torres (1949, 1960) used this method with 3- to 9-month-old seedling rootstock
with a high success rate. Majumder and Rathore (1970) reported 50% success with
splice grafting with 2-week-old seedlings and up to 60% success with 30-day-old
seedlings; however, sur-vival of the grafts was poor.

Side grafting
Side grafting, also known as the ‘Nakamura method’ (Fig. 11.4a), was for-merly
popular (Burns and Prayag, 1921; Pope, 1929; Pope and Storey, 1933; Walters,
1932; Tanaka, 1939; Lynch and Mustard, 1950; Thrower, 1954; Ruehle and Ledin,
1955; Lynch and Nelson, 1956; Ahmed, 1960; Singh, 1960; Serpa, 1964; Rao,
1967). This method is supposed to be highly effective in mild
382 S. Ram and R.E. Litz

weather in the absence of strong winds, intense heat and heavy rain (Rao, 1967),
and success has also varied (50–100%) with different cultivars (Veera-raghavan,
1945). In India, side grafting is generally used in humid, coastal areas with 1.0–1.5
cm diameter rootstock. Scions should be c.10 cm long and defoliated c.1 week
prior to grafting. The rootstock age has varied from 4 to 36 months with 72–90%
success (Mulat, 1959; Kashyap et al., 1972; Faruque and Fakir, 1973; Kanwar and
Bhajwa, 1974; Ben-ya’acov, 1976). The scion is inserted into the wedge and tied
with grafting tape. The union is complete after 2–3 months. The top of the
rootstock can be removed above the graft union after new scion growth occurs.

Notch grafting or inlaying


This method is used in the humid tropics with >8 cm diameter rootstock (Singh,
1960). The rootstock is decapitated c.45 cm above the ground level. V-shaped
notches (4–5 cm) are made at equal distances, depending upon the thickness of the
rootstock, from the periphery of the cut surface. The proxi-mal end of each scion is
fitted into each notch of the rootstock (Fig. 11.4b) and scions are secured to the
rootstock with grafting wax. The union is normally complete in c.2–3 months
(Singh, 1960).

Veneer grafting
This grafting technique was first described by Lynch (1941), and has been widely
adopted (Cooper and Furr, 1945; Ruehle and Ledin, 1955; Mukherjee and
Majumder, 1961, 1962, 1964; Wolfe, 1963; Ahmed, 1964; Serpa, 1964; Jagirdar et
al., 1968; Bhambota et al., 1971; Prasad et al., 1973; Ramirez Diaz, 1973; Gun-jate
and Limaye, 1978; Ram and Bist, 1982; Pinto et al., 2004). Veneer grafting is
usually more effective than other methods of grafting (Bhambota et al., 1971;
Prasad et al., 1973), with approximately 90% success (Ram and Bist, 1982). For
veneer grafting, 3- to 6-month-old 1.0–1.5 cm diameter scion shoots with lush
green leaves should be defoliated 3–15 days prior to grafting (depending upon the
season), leaving the petioles attached (Mukherjee and Majumder, 1961; Singh and
Srivastava, 1979a, b; Ram and Bist, 1982; Rajan and Sinha, 1987; Bajpai et al.,
1989). Prior defoliation may not be required under humid conditions (Gunjate et
al., 1976) or when extremes of temperature and RH do not occur. The scion should
be 10–15 cm long, although smaller scions (5–10 cm) have also been used (Ram
and Bist, 1982). Older shoots can also be used (Mukherjee and Majumder, 1961;
Jagirdar et al., 1968; Ram and Bist, 1982). Scions stored for 8 days at ambient
temperature (25–35C) in moist sphagnum moss covered with polyethylene can
still be grafted with a 50% success rate during March through to July (Ram and
Bist, 1982). Cultivar differences may be important.

The use of seedling rootstock at different ages (3 months to 2 years) has


resulted in 40–100% success, depending upon the season, scion maturity,
predefoliation period, storage condition of scion, etc. (Zill, 1951; Subra, 1954;
Mukherjee and Majumder, 1961, 1962, 1964; Ahmed, 1964; Serpa, 1964; Jagirdar
and Bhatti, 1968; Bhambota et al., 1971; Prasad et al., 1973; Ramirez Diaz, 1973;
Gunjate and Limaye, 1978; Ram and Bist, 1982). Dipping or smearing
Crop Production: Propagation 383

growth regulator onto the cut surfaces of the scion and rootstock has been
ineffective (Kulkarni et al., 1989). The rootstock is prepared for veneer grafting by
making a slanting cut (5 cm long); an oblique cut is then made at the base of the
first cut, so that a piece of wood along with bark is removed (Fig. 11.4c). The base
of the scion wood is then fitted into the rootstock so that the cut surfaces with the
cambium layers of scion and rootstock are in contact with each other. The
rootstock and scion are secured together with grafting tape. The union takes place
after 1.5–2.0 months. After scion growth begins, the rootstock is removed above
the graft union. Some workers have recom-mended first ringing and then removing
the rootstock shoot 1–2 weeks later, while others have partially cut through the
rootstock shoot before severing the shoot completely a few days later. After new
leaves of the scion turn green, the rootstock may be removed completely together
with grafting tape (Mukherjee and Majumder, 1961; Ram and Bist, 1982).

Rind or crown grafting


Rind or crown grafting has been used in the Americas and India. Its success
depends upon several factors, such as season, rootstock, scion maturity, mod-erate
temperature and high RH (Gangolly et al., 1957; Popenoe, 1959; Singh, 1960). The
rootstock is decapitated 30 cm above the soil. A longitudinal cut (8 cm) is made in
the bark from the top of the decapitated rootstock, down-wards, without injuring
the wood below the bark. The bark is loosened from the wood with a wooden
wedge or spatula and a freshly prepared 3-month-old scion is inserted after creating
a wedge c.6 cm long at the proximal end of the scion. The flap of bark is then
secured over the inserted scion with grafting tape. Several scions may be inserted
into a single thick rootstock (Fig. 11.4d). High RH around the graft is important for
success. The graft union occurs in c.1 month, but the grafting tape should be
removed from the grafts only after the leaves of the scion turn green. Rind or
crown grafting is a cumbersome method and is not often used in commercial
nurseries; however, it can be useful for top working.

Budding
Budding involves the grafting of a single bud, with or without wood attached, with
the rootstock. Budding methods are referred to by the shape of the bud, flap of the
rootstock covering the bud before insertion, etc. The various meth-ods for budding
are illustrated in Figs 11.5–11.7. The inserted bud eventually develops as the
crown. Rootstock above the bud is severed after bud estab-lishment. Budding
provides a stronger union and the graft union occurs ear-lier than with other
grafting methods. Both rootstock and scion should be actively growing, and
excision of the bud is easy. Stimulation of the bud prior to its excision by
predefoliation, application of growth regulators and gir-dling can improve the
efficiency of the procedure. A bud is normally selected from a 3- to 4-month-old
scion shoot, and is budded onto rootstock stems of 1.0–1.25 cm diameter c.25–30
cm above ground level where the rootstock is greyish green, except in cases of very
juvenile rootstock. Generally 1- and 2-year-old rootstocks are better for grafting
than younger ones. The bud is
384 S. Ram and R.E. Litz

completely covered by grafting tape for a few weeks. Bud growth can be
stimulated by removing grafting tape c.2 weeks after grafting.

Shield or ‘T’ budding


Shield or ‘T’ budding of mango (Fig. 11.5a) occurs with varying success, ranging
from 32–94% (Singh and Khan, 1943, 1946; Singh and Ram, 1946; Singh, 1946;
Singh, 1960; Singh and Srivastava, 1961). In northern India and Pakistan, shield or
‘T’ budding is generally most effective from March to July (India) and March–
April (Pakistan). Defoliating the scion shoot for 10–15 days before budding (Pope,
1929; Parsons, 1931; Khan, 1960; Rao, 1967; Teaotia and Maurya, 1970) and
girdling of the scion shoot several weeks prior to budding can increase bud take
(Anonymous, 1950). Scion cultivar differences also appear to be important, for
example success is better with ‘Langra’ than with ‘Aman Dashehari’ and ‘Chausa’
(Ahmed and Chatha, 1960); however, success is higher with ‘Sindhri’ than with
‘Langra’ and ‘Banganpalli’ (Jagirdar and Ali, 1968). Marked differences in success
have also been reported for dif-ferent agroecological zones in the same season:
45% in the dry hot plains of the Punjab and 90% in relatively cool areas (Bajwa
and Ram, 1946).
Temperature, RH and storage media influence bud storage (Walters, 1932;
Sundara Rao, 1956; Srivastava, 1963a, b; Rao, 1967). Storage of bud-wood in
moist sphagnum moss for 10 days is generally satisfactory. Sealing the cut ends of
budwood with melted wax and storage in a thermos flask with ice for 48 h does not
adversely affect budding (Singh and Khan, 1943). Buds can be inserted in inverted
‘T’ cuts using 2- to 5-year-old rootstock (Higgins, 1910; Ruehle and Ledin, 1955).
‘T’ budding in situ is also successful (Singh and Khan, 1942, 1946; Bajwa and
Ram, 1946; Khan, 1960) in March– April and August–October (Ullah and Ali,
1955). The bud should be secured with grafting tape. Soule (1971) described four
stages in the formation of the bud union: (i) pre-callus with wound periderm up to
4 days after budding;
(ii) callus proliferation 8 days after budding; (iii) cambial bridge between rootstock
and scion and vascular tissue differentiation 12 days after budding; and (iv) the
healed union after 2–4 weeks. If the buds are green up to 15 days after budding, the
grafting tape should be removed and the rootstock should be girdled 10 cm above
the union to stimulate the bud to sprout. The root-stock is finally removed above
the budding point after a growth flush of the scion (Luthra and Sharma, 1946;
Ruehle and Ledin, 1955; Hayes, 1957). Budded plants are transplanted only after at
least one season’s growth.

Patch budding
Patch budding (Figs 11.5b and c) (Verma, 1942; Ullah and Ali, 1955; Singh, 1960;
Moreuil, 1963; Teaotia, 1963; Jauhari and Singh, 1970; Teaotia and Maurya, 1970;
Prasad and Singh, 1972; Prasad et al., 1973) is used as an alterna-tive to approach
grafting (Garner and Chaudhri, 1976). Under subtropical north Indian conditions,
March and May through to July is the optimum period for patch budding. In
tropical Malaysia, patch budding is successful from December through to February.
The scion is defoliated 2 weeks prior to budding. Rootstocks are severed above the
budding point 1–2 weeks after budding.
Crop Production: Propagation 385

Patch budding is apparently superior to forkert budding with ‘Langra’ and


‘Dashehari’ (Teaotia, 1963).

Forkert budding
Over 90% grafting success has been reported with forkert budding (Fig. 11.5d) in
tropical South-east Asia and Sri Lanka (Paul and Guneratnam, 1937a, b). In
Maharashtra, India during July and August, success with this technique using 1-
year-old rootstock was 60–70% (Gandhi, 1942), and 100% with ‘Langra’ and
‘Dashehari’ (Singh and Srivastava, 1962; Teaotia, 1963). A modified forkert
budding method (Fig. 11.6a) can be more effective than shield budding (Garner and
Chaudhri, 1976). H-budding is another modifi-cation of the forkert method (Singh,
1960), and derives its name from an ‘H’-shaped cut made in the rootstock (Fig.
11.6b).

Chip budding
For chip budding, 2- to 3-month-old seedlings are used (Fig. 11.6c). The bud is
excised with a piece of wood attached to it. Graft union can occur 3–4 weeks after
budding because rootstock tissues are partially undifferentiated and possess a
broadly exposed cambium. The method is very efficient, and is widely used (Lynch
and Nelson, 1949, 1956; Soule, 1954; Subra, 1954; Ruehle and Ledin, 1955;
Ahmed, 1960; Bhan et al., 1969; Rajput and Haribabu, 1971), often with
modifications (Lynch and Mustard, 1950; Singh, 1960) (Fig. 11.7a).

Flap budding
The excised bud shape is boat-shaped instead of rectangular (Fig. 11.7b). This
procedure has been used in South-east Asia (Naik, 1949; Garner and Cha-udhri,
1976).

Window budding
This technique is similar to flap budding except that a window is created in the flap
for the bud so that flap does not press the bud while being tied (Fig. 11.7c). This
method has been used in Queensland, Australia. The bud union occurs in c.4 weeks
(Stephens, 1948).

Effect of rootstock

Relative advantages of polyembryonic rootstocks


Swamy et al. (1972) described the results of a rootstock trial involving six
polyembryonic rootstocks for ‘Baneshan’ and four for ‘Neelum’. Based on growth
and fruit yield from 1942 to 1965, ‘Pahutan’ and ‘Goa’ rootstocks were
recommended for ‘Neelum’ and ‘Olour’ rootstock was recommended for
‘Baneshan’. Oppenheimer (1960) reported the highest yield for ‘Haden’ mango
occurred on polyembryonic ‘Sabre’ rootstock among three different rootstocks
tried. George and Nair (1969) conducted a rootstock trial using polyembryonic
‘Chandrakaran’ and ‘Bappakai’ and monoembryonic ‘Puliyan’ rootstocks for
386 S. Ram and R.E. Litz

‘Bennett’, ‘Alphonso’ and ‘Baneshan’. ‘Chandrakaran’ and ‘Bappakai’ grew more


vigorously and produced higher yields during the first 6 years of growth compared
to monoembryonic ‘Puliyan’ rootstock. In south India, polyembryonic ‘Olour’ and
‘Bappakai’ and monoembryonic rootstocks were evaluated using ‘Neelum’ as a
scion (Gowder and Irulappan, 1970). ‘Bap-pakai’ was recommended as a rootstock
based on fruit yield and uniform growth of ‘Neelum’.

Performance of ‘Dashehari’ on 24 rootstocks (ten polyembryonic and 14


monoembryonic) was reported by Rajan and Pandey (1991a, b). Rootstocks
strongly affected tree vigour with tree height ranging from c.4.0 to 6.0 m after 14
years of growth. ‘ST-9’ and ‘Latra’ rootstocks stimulated the most vigorous
growth. ‘Ambalvi’, ‘Pahutan’, ‘Olour’, ‘Nakkare’, ‘Mylepelian’, ‘Rumani’,
‘Willard’, ‘Mundappa’, ‘Pahutan’, ‘Ambalvi’ and ‘Moovandan’ caused dwarfing.
Stem swelling was recorded with ‘Mahmuda Vikarabad’ rootstock; however, the
scion girth varied with rootstock. ‘Rumani’ and ‘Ambalvi’ rootstocks caused less
scion girth (0.75–0.8 m) than the vigorous ‘ST-9’ and ‘Latra’ rootstocks (0.79–0.84
m). Crown spread was also greater with vigorous rootstocks than with dwarfing
rootstocks. The canopy of ‘Dashehari’ trees on ‘ST-9’ and ‘Latra’ rootstock was
2.15 and 2.25 times greater than on ‘Rumani’, respectively. Fruit yield varied from
8 kg to 25 kg/tree. Dwarfing rootstock resulted in higher fruit yield per unit canopy
volume without much change in fruit quality. Polyembryonic rootstocks
‘Ambalvi’, ‘Mylepelian’, ‘Olour’ and ‘Villaikolamban’ were compared with
‘Dashehari’ seedling rootstock using ‘Dashehari’ as the scion (Jauhari et al., 1972;
Singh and Singh, 1976). The ‘Dashehari’ seedling rootstock was most vigorous and
produced higher yields, whereas ‘Villaikolamban’ resulted in the least vigour and
yield. ‘Villaikolamban’ caused dwarfing of ‘Alphonso’ and low yields
(Anonymous, 1994), and has been recommended as a rootstock for high density
‘Alphonso’ orchards, based upon higher yield/m3 of canopy.
‘Neelum’ fruit quality on ‘Bappakai’ rootstock was better compared to
‘Neelum’ grown on ‘Olour’ and monoembryonic seedlings (Gowder and Iru-
lappan, 1970). Larger ‘Neelum’ fruits having high total soluble solids devel-oped
on ‘Pahutan’ rootstock in comparison with ‘Goa’, ‘Olour’ and ‘Salem’ rootstocks
(Swamy et al., 1972). ‘Madu’ and ‘Gurih’ rootstocks delayed fruit-ing in Indonesia
compared to ‘Gurung’, ‘Kopjor’, ‘Budadaja’, ‘Nanas’ and ‘Saigon’ (Kusumo et al.,
1971). Polyembryonic rootstock ‘13-1’ has been dem-onstrated to tolerate
calcareous soil containing 20% calcium carbonate and saline irrigation water
containing >600 ppm chloride (Stoler, 1976; Kadman et al., 1978). Gazit and
Kadman (1980) grew ‘Maya’ on ‘13-l’ rootstock on cal-careous soils with up to
20% lime and 250 ppm chloride. ‘Ann’ and ‘Gomera’ polyembryonic rootstocks
also show salt tolerance (Galán-Saúco, 1993).

Top and double working

Top working is used to rejuvenate unproductive trees and for grafting on seedling
trees (Fig. 11.8a). Two methods are used for top working mango. Branches or main
limbs are cut back to 30 cm during winter months or with
Crop Production: Propagation 387

the onset of spring. The new shoots are budded or grafted in the following summer
or autumn, usually by shield budding or veneer grafting. Singh (1990) suggested
that half of a tree should be top worked in the first year and the other half in the
second year, although top working of a complete tree in a single year has been
highly successful (Lynch, 1941; Ruehle and Ledin, 1955; Ahmed, 1960; Singh,
1960). Cutting back of limbs in October or early November and veneer grafting in
March–April has been recommended in Florida (Carmichael, 1957/58; Miner,
1962); in Uttar Pradesh (India), top working on 8-year-old mango trees was more
successful during June (Singh, 1960). After top working, trees come into bearing
within 2 years (Furon and Plaud, 1972).

Grafting on an already grafted or budded tree is referred to as double working


(Singh, 1960), and double-worked trees therefore consist of three genetically
different parts, i.e. the rootstock, interstock and crown (Fig. 11.8b). In Florida, old
plantings of ‘Haden’ have been double worked with ‘Tommy Atkins’ (Mitchell,
1971). Double working can also be used to repair trees affected by disease or cold
injury and to develop a strong framework. ‘Kala-pady’ has reportedly been used as
an interstock in order to dwarf ‘Langra’ (Sen, 1939). In one trial, 12 interstocks
were grafted on two rootstocks for ‘Dashehari’; rootstock-interstock combinations
significantly affected tree height and vigour. Fruit yield was also influenced by
rootstock in all the com-binations. ‘Kurukkan’ interstock on ‘ST-9’ rootstock
yielded more fruit in comparison with ‘Ambalvi’ on the same rootstock. The
maximum yield was obtained with ‘Nakkare’/‘Chandrakaran’ and
‘Ambalvi’/‘Chandrakaran’; whereas the lowest yield was obtained with
‘Ambalvi’/‘ST-9’. Iyer et al. (1990) reported varying success (20–73%) with
double-worked ‘Langra’.

Rooting

Mango air layers have been induced to root either while they are still attached to
the parent tree or by cutting shoots into pieces containing one to several buds.
Several rooting methods have been successful with mango.

Layering
Layering is used chiefly to propagate monoembryonic mango cultivars on their
own roots. Rooting methods involve the initial training of the main stem. Shoots
close to ground level are ringed and covered with soil, while other shoots are
covered with soil mixture or wet sphagnum moss at the ringed portion and wrapped
with polyethylene to maintain moisture (Singh, 1960; Majumder and Mukherjee,
1961; Mukherjee and Majumder, 1963). Auxins such as E-naphthalene acetic acid
(NAA) and indole butyric acid (IBA) are generally essential in order to induce
rooting even after ringing.

Ground layering
Vigorous, 1- or 2-year-old shoots are selected near the base of the parent tree. The
greyish-green bark portion of the shoot (3–5 cm length) is ringed without
388 S. Ram and R.E. Litz

injuring the wood. On the proximal end of the shoot just above the ring, growth
regulators are applied and the ringed portion is buried in the soil. Depending upon
the growth regulators and their concentration, ringed shoots (or layers) generate
roots within 2–3 months. The rooted shoot can then be detached from the mother
tree and planted.

Mound layering or stooling


This technique involves the use of juvenile shoots or shoots with induced juvenility
(through heading back and etiolation followed by use of growth regulators), and is
more efficient than simple layering (Majumder and Mukherjee, 1961). One-year-
old mango seedlings should be decapitated 25–30 cm above ground level in
February–March, and several new shoots will develop from the stem by May.
When leaves of the new shoots turn green, the shoots are ringed at their proximal
end and treated with 5000 ppm IBA in lanolin paste just above the ring. A soil
mound is created around the base of ringed shoots. Rooting occurs after 1–2
months and rooted layers can be detached from the parent plant and established in
individual pots. Each layer can generate between four and seven roots. Stooling has
been success-ful with many cultivars using growth regulators and rooting cofactors
in stool beds (Ram et al., 1981; Sirohi and Ram, unpublished data). Dwarf culti-
vars root poorly, but can be stimulated to form thin and fibrous roots by applying
auxin, for example IBA (15,000–30,000 ppm) and NAA (2500–10,000 ppm)
mixtures in lanolin paste with or without rooting cofactors, such as caf-feic acid
(0.046M) and catechol (0.046M). Caffeic acid appears to be highly synergistic to
IBA.

Pot layering
With pot layering, the ringed portion of the shoot is maintained in a soil mix-ture in
special pots at a convenient height within the tree canopy. This can be done only
during periods of high RH, but the success is low (c.15–20%) (Singh, 1960).

Air layering
Although air layering or marcottage is one of the oldest methods for propagat-ing
mango, good success has been elusive (Grove, 1947; Garg, 1954; Rangacha-rlu and
Rao, 1956; Rao and Rao, 1956; Choudhury, 1959; Jauhari and Nigam, 1962; Rao
et al., 1963; Mukherjee and Majumder, 1963; Srivastava, 1963a, b; Mukherjee and
Bid, 1965; Sen and Bose, 1966; Bid and Mukherjee, 1969; Basu et al., 1972;
Prasad and Singh, 1972; Núñez-Elisea et al., 1992). Juvenility, high RH and
moderate temperature are important factors for air layering mango (Singh, 1954;
Rao et al., 1963; Rao, 1967; Chhonkar and Singh, 1972; Singh et al., 1979; Ram
and Sirohi, 1982a). Cultivar effects on air layering are also pronounced; and
vigorous cultivars root better than dwarf cultivars (Garg, 1954; Rao and Rao, 1956;
Rao, 1967; Basu et al., 1972; Ram and Sirohi, 1982b).

Layering success can be improved with auxins, such as 2–3% indole ace-tic
acid (IAA) and 0.5–2.0% NAA (Thakurta and Dutt, 1941; Singh and Teotia,
Crop Production: Propagation 389

1951; Srivastava, 1963a, b; Núñez-Elisea et al., 1992). Although Núñez-Elisea et


al. (1992) used NAA alone, NAA and IBA mixtures not only improve root-ing but
also accelerate the production of fibrous roots, which increases sur-vival. Etiolated
layers just above the girdle require less auxin than non-etiolated branches. Root
regeneration in air layers requires up to 90 days, but emer-gence of roots is first
noticeable a month after layering. Sen et al. (1961) and Mukjerjee and Bid (1965)
observed that IBA depletes sugar in the bark and wood of the rooting zones. Ram
and Sirohi (1982a) studied the interaction of rooting cofactors involved with air
layering of mango. Application of 0.01– 0.08M IBA in lanolin paste increased root
numbers from 2.2 to 2.8 with ‘Dashehari’, but root number could be increased from
2.8 to 6.0 with 0.01– 0.10M catechol. Survival was correlated with increased
numbers of roots; however, IBA and catechol did not show any additive or
synergistic effect on air layering. July was the best month for rooting of layers
(>90% success) and winter months were the worst.

Rajan and Ram (1989) showed that 0.0573M NAA and 0.05 × 102M chlo-
rogenic acid had a synergistic effect on rooting of layers and the number of roots
formed during March when applied separately with 0.0735M IBA. In October,
NAA and chlorogenic acid showed synergism with respect to num-ber of roots
produced, but their effect on rooting was additive. Pyrogallol (0.06 × 10−2M) and
6-benzylaminopurine (0.22 × 10−3M) also had a synergistic effect on root
regeneration during October and March. IBA increases endogenous root-ing
cofactors and lowers rooting inhibitors, resulting in early rooting.

Cuttings
The potential of mango cuttings to form roots depends on maturity of the cutting,
rooting medium, temperature, RH, etc. Girdling or etiolation of stems for a few
weeks and use of growth regulators have improved rooting frequency (Thakurta
and Dutt, 1941; Gardner and Piper, 1943; Van Overbeek and Gregory, 1945; Said
and Shoushan, 1945; Khalifa et al., 1964; Dijkman, 1950; Ledin and Ruehle, 1954;
Gowda, 1962; Ahmed, 1964; Sen and Bose, 1964; Mukherjee et al., 1966, 1967;
Maurice, 1969; Sen et al., 1969; Ali and Nazir, 1970; Prasad and Pathak, 1970; Bid
and Mukherjee, 1972; Bid et al., 1972). Rooting is improved with cuttings from the
lower bole of the tree rather than from the upper bole. Mukherjee et al. (1966)
reported that cut-tings from 4-year-old trees root more readily than those from 10-
year-old trees, alhough Sen et al. (1968) had better success with older shoots.
Hard-wood cuttings reportedly root better (Hussain, 1953; Benincasi, 1970) than
semi-hardwood cuttings (Singh and Teaotia, 1951). Mango cuttings should be c.15
cm long and 1.2–1.8 cm girth with between three and five buds (Garner and
Chaudhri, 1976). Retention of one to two leaves at the apex of the cut-tings has
helped rooting, and is attributed to the presence of rooting cofactors in the leaves.

In India and Pakistan, cuttings are generally made during the spring months
(Hussain, 1953). Cuttings can be stored for a short time before apply-ing the
rooting treatment. Khalifa et al. (1964) rinsed cuttings for 24 h with running tap
water, which increased their shelf life. Longevity of cuttings can
390 S. Ram and R.E. Litz

also be increased by inserting them in a solution consisting of 100 ppm IBA, 10


mg/litre vitamin B1, 2% ammonium sulphate ((NH4)2SO4) and 2% sucrose.
Cuttings can be stored at 4C for 20 days. Rooting of hardwood cuttings under mist
can be improved with IBA and NAA (Lambourne, 1935; Cooper, 1936; Thakurta
and Dutt, 1941; Cooper and Stoutemyer, 1945; Dijkman, 1950; Jauhari and Nigam,
1958; Sen and Bose, 1964; Mukherjee et al., 1965; Basu et al., 1966; Sen et al.,
1969; Ali and Nazir, 1970; Rajan and Ram, 1983a, b). IBA increases reducing and
non-reducing sugars, and stimulates a favourable carbon to nitrogen (C:N) ratio in
ungirdled cuttings comparable to girdled cuttings without IBA. Application of
NAA to the ring 1 week before cuttings are made and dipping their basal ends in an
IBA solution at the time of detaching, increases rooting efficiency (Van Overbeek
and Gregory, 1945; Sen et al., 1969). Low pH potting mixtures (pH 4.5–7.0)
reportedly are better for rooting than higher pH mixtures (Kains and McQuestan,
1955). Reddy and Majumder (1975) and Mitra and Bose (1986) used bottom heat at
30C  2 and 35C, respectively, to stimulate root formation. Reuveni et al.
(1991) and Reuveni and Castoriano (1993) treated semi-hardwood cuttings of
‘Turpen-tine’, ‘Gomera’, ‘Sabre’ and ‘13-1’ with bottom heat at 20, 25, 30 and
35C under mist, and observed most rooting occurred at 25 and 30C. The percent-
age of rooting in cultivars varied from 60–95% with between four and ten
roots/cutting. Wounding of the proximal ends of cuttings from 7-month-old
seedlings together with bottom heat increases the number of roots per cut-ting
(Reddy, 1970).

The involvement of phytochrome in root regeneration of mango cuttings was


proposed by Reddy et al. (1975), who observed 92% rooting with etio-lated and
red-light-treated cuttings of 7-month-old seedlings, whereas cut-tings in a hot bed
treated with 5000 ppm IBA and planted under normal light resulted in 41% rooting.

Reddy and Majumder (1975) achieved 100% rooting of cuttings from 7-


month-old mango seedlings treated with 5000 ppm IBA combined with 2000 ppm
each of umbelliferone, ruten or quercetin. Phenols and flavonoids acted as rooting
cofactors of IBA with juvenile cuttings. Sadhu et al. (1978) and Reddy and
Majumder (1978) observed a synergistic effect of some phe-nolic compounds (e.g.
p-hydroxybenzoic acid, p-coumaric acid and ferulic acid) used as a preplanting
treatment with auxin for induction of rooting in hardwood mango cuttings.
Synergism was more pronounced with IBA than IAA.

Basu et al. (1970) observed no significant difference in the levels of endog-


enous rooting substances (i.e. p-hydroxybenzoic acid, p-coumaric acid and abscisic
acid) between juvenile and non-juvenile cuttings. Rajan and Ram (1983a, b)
studied the levels of endogenous rooting hormones of mango cut-tings with bottom
heat under mist and measured increased levels of rooting hormones and low levels
of rooting inhibitors. Fayek et al. (1981) reported that shoots of 35-year-old
‘Madu’ contained more rooting inhibitors than shoots of 1-year-old plants. Using
15,000 ppm IBA and 10,000 ppm NAA, Rajan and Ram (1983a, b) obtained c.70%
rooting with mature cuttings; about eight fibrous roots were regenerated as a result
of each treatment, and all
Crop Production: Propagation 391

cuttings survived in the nursery. Sadhu (1979) and Sadhu and Bose (1980a, b)
found that 10,000 ppm cycocel pretreatments of 10-year-old ‘Langra’ resulted in
41% rooting with 2.2 roots/cutting.

Micropropagation

Somatic embryogenesis from cultured nucellar explants of polyembryonic cultivars


of Mangifera indica was first reported by Litz et al. (1982, 1984). Sub-sequently,
somatic embryogenesis was induced from the cultured nucellus of monoembryonic
cultivars (Litz, 1984). DeWald et al. (1989a, b) optimized the protocols for large-
scale production of embryogenic cultures in suspension and for somatic embryo
maturation and germination. Shoot tip culture of ‘Turpentine’, ‘Gomera’ and ‘13-1’
seedlings has been described by Yang and Ludders (1993); the rate of
multiplication was low, and performance of plants in the field has not been tested.
The current status of somatic embryogenesis and organogenesis of mango is
reviewed by Litz et al., Chapter 18, this volume.

11.4 Comparative Performance of Trees Propagated by


Different Methods
Ram and Sirohi (1989) compared the performance of ‘Dashehari’ propagated by
cleft grafting, approach grafting, veneer grafting, epicotyl grafting, stool-ing and
air layering. The trees were of uniform size at the time of planting. Approach-
grafted plants did not establish in the field as well as the other methods. During the
first 12 years, epicotyl grafts grew more rapidly than cleft-grafted, veneer-grafted,
approach-grafted, stooled and air-layered trees, respectively (Ram, 1993). The
development of the crown also followed the same pattern. The trees propagated by
rooting did not develop an erect main stem to an adequate height (1.2 m), which
hindered cultural operations under the tree. Trees propagated by approach grafting,
veneer grafting and epicotyl grafting all produced branches close to the ground.
Maximum yields were obtained in trees propagated by epicotyl and cleft grafting
followed by veneer grafting, stooling, air layering and approach grafting,
respectively. The architecture of cleft-grafted plants was much better than trees
propa-gated by other methods; whereas, rooting methods produced twiggy, spread-
ing and dwarf trees (Ram, 1993). After 18 years, cleft-grafted trees were superior
with respect to architecture and yield, followed by epicotyl and veneer-grafted
trees. Rajan and Pandey (1991a, b) conducted experiments with ‘Dashehari’ and
‘Langra’ that were propagated by different vegetative methods using
monoembryonic seedling rootstock. Approach-grafted ‘Dashehari’ attained the
greatest height followed by epicotyl-grafted, bud-ded and air-layered trees after 7
years. Budded ‘Langra’ grew most vigor-ously, followed by approach-grafted,
veneer-grafted, epicotyl-grafted and air-layered trees. Similar growth of scion girth
and crown spread was recorded with both cultivars. The type of shoot used for
approach grafting
392 S. Ram and R.E. Litz

affected tree growth of ‘Dashehari’. Grafting height should be minimized and close
contact should be maximized to achieve faster growth when vigorous rootstocks
are used.

11.5 Conclusions
Standard methods are widely utilized for propagating mango scion cultivars with
increasing efficacy. In many regions, including India and Mexico, scion cultivars
are still being propagated on heterogeneous monoembryonic seed-ling rootstocks
(see Crane et al., Chapter 13, this volume), despite the demon-strated advantages of
clonal nucellar rootstocks. The potential of clonally propagated monoembryonic
mango rootstock has not been properly investi-gated. The genetic heterogeneity of
monoembryonic mangoes has been explored neither for stress tolerance nor for
their effects on scion growth and development. Somatic embryogenesis could play
an important role in such investigations as an alternative propagation method (see
Litz et al., Chapter 18, this volume). Other Mangifera species also have interesting
attributes, and should be screened for graft compatibility with mango (see
Bompard, Chap-ter 2, this volume). The species that could be tested as rootstock
for mango might extend mango cultivation to areas where abiotic and biotic
stresses currently limit production and could provide a better source for dwarfing
rootstock for high-density orchards. Mango species growing in swamps or
seasonally inundated areas (i.e. Mangifera decandra, Mangifera gedebe, Mangifera
inicarpoides, Mangifera griffithii and Mangifera quadrifida) represent a promis-
ing source of rootstock for the development of mango cultivation on poorly drained
soils and inundated lands. In West Kalimantan, Mangifera laurina is occasionally
used as a rootstock for commercial mango cultivars on periodi-cally inundated
riverbeds, and is now being tried at Sabah (Bompard, 1993). Mangifera zeylanica
has been tried as a rootstock for several mango cultivars in Sri Lanka (Parsons,
1931). Interspecific hybridization of other Mangifera species with the common
mango could also increase the available genetic variability for rootstock
development.

The potential for using species in other genera in the Anacardiaceae as


rootstock has scarcely been explored. Burns and Prayag (1921) investigated the use
of Semecarpus anacardium, Spondias mangifera (Spondias pinnate), Spondias
acuminata and Horigarna grahami as rootstocks for common mango without any
success. Anacardium occidentale (cashew) seedlings have been reported to be graft
compatible with mango scions and mango fruit on cashew rootstock were
reportedly almost double the size of normal fruit with smaller seeds and free from
fibre (Garner and Chaudhri, 1976). Mango rootstock development should receive
as much attention as breeding of scion cultivars.
Classical breeding and grafting studies should be greatly expanded to include
the enormous genetic variability within the genus. Emerging bio-technological
approaches also should not be overlooked as alternative prop-agation methods and
as the means to engineer specific horticultural traits in candidate rootstocks.
Crop Production: Propagation 393

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12 Crop Production: Mineral Nutrition
I.S.E. Bally
Department of Primary Industries and Fisheries, Queensland, Australia

12.1 Introduction 405


12.2 Soils, Mineral Diagnosis and Sampling 405
12.3 Tissue Mineral Diagnosis and Sampling 406
12.4 Interpreting Soil and Leaf Analyses 406
12.5 Nitrogen 407
Sources, uptake and translocation of N 409
Nitrogen deficiency and toxicity 409
Effect of N on crop production and fruit quality 410
Nitrogen management 411
12.6 Phosphorus 411
Sources, uptake and translocation of P 411
Phosphorus deficiency and toxicity 412
Effect of P on crop production and fruit quality 412
12.7 Potassium 412
Sources, uptake and translocation of K 413
Potassium deficiency and toxicity 413
Effects of K on crop production and fruit quality 414
12.8 Magnesium 414
Sources, uptake and translocation of Mg 414
Magnesium deficiency and toxicity 414
12.9 Sulfur 415
Sources, uptake and translocation of S 415
Sulfur deficiency and toxicity 415
12.10 Zinc 416
Sources, uptake and translocation of Zn 416
Zinc deficiency and toxicity 416
Effect of Zn on crop productivity 417
12.11 Manganese 417
Role of Mn 417
Sources, uptake and translocation of Mn 417
Manganese deficiency and toxicity 418
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
404 (ed. R.E. Litz)
Crop Production: Mineral Nutrition 405

Effect of Mn on crop production 418


12.12 Iron 418
Sources, uptake and translocation of Fe 418
Iron deficiency and toxicity 419
12.13 Calcium 419
Sources, uptake and translocation of Ca 419
Calcium deficiency and toxicity 420
Effects of Ca on crop production and fruit quality 420
Calcium in leaves and fruit 421
12.14 Boron 422
Sources, uptake and translocation of B 422
Boron deficiency and toxicity 423
Effect of B on crop production and fruit quality 423
12.15 Conclusion 424

12.1 Introduction
Assessment of the mineral status of mango trees is not without its challenges. Like
many other tropical woody perennial tree crop species, mangoes have complicated
and variable phenological cycles that influence the trees’ uptake and translocation
of minerals. Their extensive root system enables them to exploit unevenly
distributed minerals throughout the soil profile; these min-erals are often not
assessed during routine soil analysis. The leaves, trunk, bark and roots act as
mineral reserves that buffer many short-term mineral shortages (Robinson, 1986b).
Soil and leaf mineral analyses used as short-term indicators of tree mineral status,
tree productivity or fruit quality are therefore difficult and unreliable (Catchpoole
and Bally, 1995).

12.2 Soils, Mineral Diagnosis and Sampling


Statistical relationships between the soil and tree mineral status have been difficult
to establish because of the large mineral reserves in trees and the variable
distribution of minerals in the soil profile (Robinson, 1986b). How-ever, soil
mineral analyses can be useful for determining the availability of the essential
minerals, and if they are within the optimal range for mango. Soil analysis also
provides valuable information on other soil properties that can influence mineral
availability to the tree, i.e. pH, electrical conductivity (Ec) and concentrations of
organic matter and clays. Regular soil analysis in conjunction with tissue analysis
in mature cropping orchards can provide useful information on the changes in soil
minerals over time and on the influences of fertilizer programmes on soil and tree
status.
There are several publications that outline the techniques that can be uti-lized
for obtaining representative soil samples for mineral analysis (Dow et al., 1991;
Peverill et al., 1999; Anonymous, 2007). Before a new orchard is planted, the soil
should be sampled from several depths and analysed to capture a full picture of the
mineral status throughout the soil profile. This
406 I.S.E. Bally

will facilitate the adjustment of pH and deep placement of minerals before the trees
are planted. In established orchards the sites of soil sampling will vary according to
the age of the trees and the soil type. In sandy or duplex soil types where minerals
are easily leached from the upper to the lower pro-file, deep (1–1.5 m) monitoring
will indicate any mineral build-up and help avoid unnecessary fertilizer
applications. In lighter sandy soils, surface sam-pling to depths of 10–15 cm can
underestimate soil mineral concentrations as these layers are often leached and are
low in clay colloids.

12.3 Tissue Mineral Diagnosis and Sampling


Leaf mineral analysis is commonly used to assess mango tree mineral status, and is
useful for developing and monitoring tree fertilizer programmes. Leaves often
display visual symptoms of toxic and deficient concentrations of many minerals.
Sampling mango leaves for mineral analysis should be done when the tree is at its
most phenologically quiescent stage, i.e. when leaf mineral concentrations are most
stable. One of the most stable periods in the mango phenological cycle is the
dormant phase, which occurs after the completion of summer flushing and
approximately 2 weeks before the emer-gence of flower panicles. The common
practice of withholding irrigation water leading up to flowering makes this period
the most inactive of the year and an ideal time for leaf sampling. At other times of
the year, leaf mineral concentrations sharply decrease when the tree is flowering
and fruiting and increase in the months following harvesting (Catchpoole and
Bally, 1995; Oosthuyse, 2000b). Sampling at an inactive growth stage reduces
variability between leaf samples and provides a stable reference point for annual
com-parisons. Guides with respect to the most appropriate leaves for sampling
have been variously reported (Kumar and Nauriyal, 1979; Chadha et al., 1980;
Smith, 1992; Catchpoole and Bally, 1995), and generally concur that the most
appropriate leaves to sample are the third or fourth leaf behind the apical bud, or
the first full-size leaf of the most recently matured dormant flush where leaves are
fully expanded and hardened off.

Leaves recently sprayed with foliar nutrients or fungicides should be avoided


or noted, as analyses of manganese (Mn), zinc (Zn), boron (B) and copper (Cu) are
commonly affected by mineral residues on the outside of leaves or imbedded in the
cuticle, and which are not available to the tree. Some publications recommend that
sampled leaves should be washed with deionized water to reduce residues from
sprays and dust from the operator’s hands (Reuter et al., 1986; Shu et al., 1992).

12.4 Interpreting Soil and Leaf Analyses


Several published soil and leaf mineral standards are available to assist in
interpretating analysis results (Smith and Scudder, 1951; Young and Koo, 1969;
Kumar and Nauriyal, 1977; Robinson, 1986a; Peverill et al., 1999; Stassen
Crop Production: Mineral Nutrition 407

et al., 1999) (Table 12.1). Most of these standards are in reasonable agreement with
respect to the optimal ranges of individual minerals (Samra and Arora, 1997). Soil
and leaf analyses should be interpreted by comparing with these standards and with
previous soil and leaf analyses and past seasons’ crop-ping and fertilizer
application history. The recommended optimal ranges of leaf mineral
concentrations should be considered as general guides only, as high-yielding trees
producing good quality fruit have been found to vary widely in leaf mineral
composition (Catchpoole and Bally, 1995; Oosthuyse, 2000b; Medeiros et al.,
2004).
Positive relationships between leaf minerals and tree productivity have been
reported. Oosthuyse (1997) determined that leaf concentrations of nitro-gen (N),
phosphorus (P), potassium (K), magnesium (Mg) and Zn influenced the number of
fruit retained, and that leaf Zn and Mg also influenced fruit size. Rao and
Mukherjee (1989) observed positive correlations between tree yields and leaf N
and K from non-fruiting terminals in five Indian mango cultivars with generally
low N and K concentrations. Although these rela-tionships have been observed,
there are many factors that can influence pro-ductivity. Therefore, predicting
productivity based on leaf analysis alone is difficult. However, soil and leaf
analyses are useful for identifying major mineral imbalances and long-term trends
in tree nutrition, and can be used to adjust fertilizer programmes.

Leaf age can affect the mineral concentration in assays. Minerals that are
mobile within the plant, for example N, P, K and Mg, generally decrease during
leaf aging and the less mobile minerals, such as calcium (Ca), sulfur (S), B and
Mn, accumulate in leaves with age (Chadha et al., 1980; Medeiros et al., 2004).
Mango leaves that have been sampled from fruiting terminals generally
display increasing concentrations of N, K, Ca, Mn, iron (Fe), Cu and Zn dur-ing
early fruit development and decreasing concentrations during late fruit
development and maturation. These minerals, with the exception of Mg and P, also
vary greatly among fruiting terminals (Oosthuyse, 1997, 2000b).
The Diagnosis and Recommendation Integrated System (DRIS) for inter-
preting leaf mineral analysis, uses ratios of mineral concentrations rather than the
absolute mineral concentration to identify limiting minerals in order of their effect
on the tree (Beaufils, 1973). With mango, DRIS has been used with varying
success. Raghupathi et al. (2004) observed that DRIS was unable to diagnose the
nutrient imbalance of a particular nutrient in isolation; how-ever, others have used
the technique more successfully. Schaffer et al. (1988) used DRIS to identify Mn
and Fe as the most deficient elements in orchards with tree decline, a disorder of
unknown aetiology, and Hundal et al. (2005), Raj and Rao (2006) and Wadt et al.
(2007) utilized DRIS to identify yield-limiting mineral combinations in mangoes
grown in India and Brazil.

12.5 Nitrogen
Nitrogen is one of the most important elements for crop production, and has a
significant role in mango growth, yield and fruit quality. Nitrogen is an
408
Table 12.1. Suggested optimal mango leaf mineral concentration ranges according to different sources.

References

Young and
Smith and Koo (1969, Crane Catchpoole Kumar Pimplasker Stassen Bhargava
Robinson Scudder 1971), Young and et al. and Bally and Nauriyal and Bhargava et al. and Chadha
a
Mineral Unit et al. (1997) (1951) Sauls (1981) (1997) (1995) (1977) (2003) (1999) (1988)
N 1.0–1.5 1.54 1.0–1.5 1.2–1.6 0.8–1.9 1.0 0.89–1.93 1.25 1.23
P % 0.08–0.18 0.05 0.09–0.18 0.09–0.12 0.12–1.3 0.10 0.06–0.11 1.45 0.06

I.S.E. Bally
K % 0.3–1.2 0.97 0.5–1.0 0.4–0.8 0.4–2.5 0.50 1.02–2.01 0.1 0.54
Ca % 2.0–3.5 0.91 3.0–5.0 2.0–3.5 1.5–2.8 1.50 – 0.8–1.05 1.71
Mg % 0.15–0.4 0.26 0.15–0.47 0.25–0.35 0.2–0.4 0.15 – 2.8 0.91
S % 0.5–0.6 – – – 0.1–0.23 0.50 0.11–0.17 0.3 0.12
B % 50–80 – 24–84 – 20–140 – – 50 –
Fe mg/kg 7–200 – 38–120 70–100 30–120 – – 80 1.71
Mn mg/kg 60–500 – 92–182 – 160–980 – – 80 66
Zn mg/kg 20–150 – 10–119 20–40 20–63 – 11–26 40 25
Cu mg/kg 10–20 – 28–35 – 10–150 – – 20 12
Mo mg/kg – – – – 0.2–0.4 – – 50 –
a
N, nitrogen; P, phosphorus; K, potassium; Ca, calcium; Mg, magnesium; S, sulfur; B, boron; Fe, iron; Mn, manganese; Zn, zinc; Cu,
copper; Mo, molybdenum.
Crop Production: Mineral Nutrition 409

essential component of many plant tissues, and occurs in chlorophyll, amino acids,
proteins, enzymes and growth hormones and is a major driver of plant growth,
having a direct effect on tree vigour (Marschner, 1995). Nitrogen at 100 g/tree/year
has been shown to be sufficient for mango tree growth (Kanwar et al., 1987), and
N concentrations influence concentrations of other elements when N is either low
or in excess (Sen et al., 1947).

Sources, uptake and translocation of N

Nitrogen in the soil occurs in many forms, but for the most part as large and
complex organic molecules that comprise the organic matter. These mole-cules are
too large for roots to absorb, and are broken down to nitrate (NO 3–) and
ammonium (NH4+). The concentration of N in soil is dependent on the
concentrations of soil organic matter and mineral N. Nitrogen that is avail-able to
the plant is determined by the processes of mineralization, immobili-zation, de-
nitrification, volatilization and leaching, which are influenced by temperature,
moisture, pH and aeration. Nitrogen is easily leached from the soil by rain and
irrigation, and consequently, soil N often limits tree growth, especially in sandy
soils (Strong and Mason, 1999).
Common N fertilizers include: urea (CO(NH2)2, 46% N), potassium nitrate
(KNO3, 13% N, 38% K), calcium nitrate (CaNO3, 15.5% N, 19% Ca), ammonium
nitrate (NH4NO3, 35% N), sulfate of ammonia ((NH4)2SO4, 21% N, 23.6% S).
Nitrogen is taken up by mango trees primarily through the roots as NO 3– and
NH4+; NO3– is the preferred source. Nitrogen can also be adsorbed through leaves
as ammonia (NH3), urea and amino acids. Nitrate, after being adsorbed by the
roots, is either reduced to NH4+ in the roots or translocated to the leaves, stems or
other tissues, where it is reduced to NH4+ in a two-step process in which NO3– is
reduced to nitrite (NO2–) which is in turn reduced to NH4+ (Marschner, 1995).
Ammonium metabolism is complex, and several pathways are necessary to
produce amino acids, proteins, enzymes and hor-mones. Within the tree, N
concentrations vary among tissues and are depen-dent on N availability in the soil
and demand within the tree. If N is limiting, the plant can translocate N from older
tissues to new growing tissues where it is required (Marschner, 1995).

Nitrogen deficiency and toxicity

Nitrogen deficiency symptoms appear in leaves as yellowing or chlorosis initially


of the older leaves, slow growth and lack of vigour. Severe N defi-ciency can cause
complete leaf yellowing, leaf and fruit abscission and death of twigs and branches
(Sen et al., 1947; Smith and Scudder, 1951). Nitrogen toxicity is not often seen in
mango; however, trees with high N concentra-tions have dark green leaves and
excessive vegetative vigour, often at the expense of flowering and cropping (Tiwari
and Rajput, 1976). High N con-centrations are also associated with internal fruit
disorders, e.g. jelly seed
410 I.S.E. Bally

and internal breakdown in many cultivars (Tarmizi et al., 1993; Cracknell Torres et
al., 2004; see Galán Saúco, Chapter 9, this volume).

Effect of N on crop production and fruit quality

Nitrogen has a major effect on mango tree vigour, stimulating both vegeta-tive and
floral growth. Increases in shoot length, leaves per shoot and leaf area have been
demonstrated by Singh et al. (1973), Tiwari and Rajput (1975), Syamal and Mishra
(1989), Reddy et al. (2000) and Sergent et al. (2000).
Yeshitela et al. (2005) reported that N in combination with K, as KNO 3 and
urea, improved the percentage of terminal shoots that flower; however, excessive N
can stimulate vegetative growth at the expense of flowering, fruit set and fruit
quality (Scholefield et al., 1986; Monselise and Goren, 1987; Silva et al., 2002).
Nitrogen can cause increased fruit set and retention (Singh et al., 1973; Oosthuyse,
1997) and fruit weight and yield (Alvian, 1974; Tiwari and Rajput, 1975; Young
and Koo, 1975; Alvian and Figueroa, 1977; Syamal and Mishra, 1989; Reddy et
al., 2003). Kanwar et al. (1987) demonstrated that N at 100 g/tree/year-of-age was
sufficient for tree growth and Young and Koo (1975) reported that maximum
yields were increased by N applications of 0.8–1.1 kg/tree/year. Many reports of
the effect of N on increased mango fruit size and yields are based on data of N in
combination with other ele-ments (i.e. P and K) and may be partly attributed to
these combinations; however, N has a major influence on productivity in mango.

High N application rates that stimulate yield increases can also have negative
effects on fruit quality. Nitrogen has been negatively associated with fruit colour in
mango (Oosthuyse, 1993; McKenzie, 1994, 1995). Nguyen et al. (2004)
demonstrated that high N applications during fruit growth inhibited the de-greening
of ripening fruit, causing green skin at ripeness. Bally (2007) also reported a
negative relationship between N and fruit colour, demonstrating that high leaf N
concentrations reduce the percentage of yellow skin in ripe fruit, reduced the
lightness and chroma (vividness) of the yellow colour, the percentage of skin
covered with blush and the intensity of the blush colour (Plate 73). He also
identified a significant exponential rela-tionship between the severity of sunburn
and skin N concentrations; the effects of sunburn were reduced as N concentrations
increased in the fruit skin.

Early indications of the negative effect of N on postharvest rots in mango were


demonstrated by Weng and Chuang (1997), who observed that N posi-tively
affected the germination rate, hyphal growth and appressoria forma-tion of
Colletotrichum gloeosporioides. When Nguyen et al. (2004) investigated the effect
of N on fruit quality, they found that 300 g/tree applied as a foliar spray
significantly increased the severity of anthracnose caused by C. gloeosporioides.
These observations were confirmed by Bally (2007), who found that the severity of
fruit anthracnose during ripening increased with applied N (Plate 74). He also
demonstrated that the N effect was because N enhanced the decline of antifungal
resorcinol compounds in the fruit exocarp during
Crop Production: Mineral Nutrition 411

ripening. Nitrogen has been a recognized factor in determining internal fruit quality
of mangoes, with imbalances of N and other minerals, principally Ca, being
implicated as a major factor causing the various forms of internal breakdown
(Young and Miner, 1961; Subramanyam et al., 1971; Malo and Campbell, 1978;
Burdon et al., 1992; Tarmizi et al., 1993; Cracknell Torres et al., 2004; Torres et
al., 2004; see Galán Saúco, Chapter 9, this volume).

Nitrogen management

Leaf N concentrations between 1 to 1.5% dry weight (DW) are generally con-
sidered to be in the optimal range (Robinson et al., 1997). Optimum concen-
trations for fruit skin N have not been published, although Catchpoole and Bally
(1995) reported skin N concentrations of 0.069% DW in mature ‘Kens-ington
Pride’ fruit. Leaf N concentrations vary throughout the year, and are influenced by
the growth events during the phenological cycle. Nitrogen concentrations are
generally greatest at the end of the summer vegetative flushing period and decrease
during panicle growth, flowering and fruiting (Catchpoole and Bally, 1995;
Medeiros et al., 2004).
Because N is highly mobile within the tree and a primary driver of growth and
fruit quality, the general practice of monitoring leaf and soil N annually is
inadequate for assessing tree N status at all stages of tree phenol-ogy. A cheap,
rapid test for N is needed to provide closer monitoring of changes in leaf N status
and allow the rates and timing of supplementary N applications to be matched to
tree and fruit demands. Bally and Still (per-sonal communication) have calibrated
the Konica-Minolta Soil Plant Analy-sis Diagnostic (SPAD-502) meter to measure
the chlorophyll content of leaves, which is directly related to their N status
(Gonzalez et al., 2005).

12.6 Phosphorus
Phosphorus is important for cell division and growth and is a component of many
essential plant molecules such as sugar-phosphates that are involved in respiration,
photosynthesis and other metabolic pathways, nucleotides such as DNA and RNA,
phospholipids in membranes and as pryophosphate (PPi) in ATP and other cellular
energy metabolism molecules (Salisbury and Ross, 1992). Phosphorus is also
involved in root tip elongation, fruit ripening and leaf expansion.

Sources, uptake and translocation of P

Phosphorus that is available for plants occurs in two forms in soil, primarily as the
monovalent phosphate anion (H2OP4–) and secondly as the divalent anion
(HPO42–) in the soil water solution. The balance of these anions is dependent on
soil pH, with HPO42– favoured in soils >pH 7 and H2OP4–
412 I.S.E. Bally

favoured in soils <pH 7 and most readily available in soils with pH between 6 and
7. In higher pH soils, calcium phosphate compounds tie up P from plant
availability, and in lower pH soils, P oxidizes with Fe, aluminium (Al) and Mn to
become unavailable to plants. Other forms of soil P include liable P that is bound
on clays and organic matter, which can become available to trees when dissolved in
water.
Phosphorus is most rapidly taken up by the tree as HPO 42– and slower as
H2OP4– and often enhanced by chemical association with N. After entry into the
roots, the anions are either converted to organic phosphates immediately or after
transport to other tissues. Within the tree, P is easily translocated between tissues,
with redistribution usually occurring from older leaves to younger actively growing
tissues. Phosphorus concentrations within the mango tree are highest in the roots,
wood and bark and lowest in the leaves (Vuuren and Stassen, 1997). Leaf P
concentrations are generally at their low-est during fruit development and at their
highest in the vegetative growth period (Medeiros et al., 2004).

Phosphorus deficiency and toxicity

As P is a mobile element, deficiencies are initially seen in the older leaves.


Phosphorous deficiency and toxicity are rare in mango trees, with toxicity
symptoms occasionally seen in nursery stock where trees are grown in pot-ting
media based on sugarcane mill/press mud. Symptoms appear as stunted seedling
growth with bronzing of the leaves. Phosphorus deficiency appears first in older
leaves as marginal necrosis with brown taints, tip necrosis, pre-mature abscission
and stem dieback (Smith and Scudder, 1951; Singh and Saxena, 1994). Sen et al.
(1947) describe deficiency symptoms in leaves as developing a reddish-purple
colour on the undersides of leaves that spreads over the entire leaf, eventually
encompassing the veins, with leaves becom-ing thick and stiff. Phosphorus-
deficient trees have restricted root development that reduces tree and fruit size
(Stassen et al., 1999).

Effect of P on crop production and fruit quality

There are several reports of P applied in combination with N and/or K and resulting
in increased yields (Samra and Arora, 1997); however, these reports do not
separate the P from the other elements. Reddy and Majmudar (1985) reported an
association between higher concentrations of P in terminals and floral induction
(Narwadkar and Pandey, 1989).

12.7 Potassium
Potassium in the cytoplasm is an activator of enzymes involved in photosyn-thesis,
respiration and starch and protein synthesis (Bhandal and Malik, 1988;
Crop Production: Mineral Nutrition 413

Marschner, 1995). Potassium is important for cell growth due to its role in cell
expansion and development of thick epidermal cell walls that increase the
resistance of trees to pathogens and insect pests. Potassium is involved in tree
water status by regulating water uptake by the roots and water loss through the leaf
stomata (Salisbury and Ross, 1992).

Sources, uptake and translocation of K

Most of the K in soils (90–98%) is in the form of insoluble crystalline minerals that
are unavailable to plants. Available K occurs in the soil solution as K + ions and
attached to cation exchange sites on clays (Gourley, 1999). Potas-sium moves
readily between the exchange sites and the soil solution and is influenced by
moisture and temperature. Heavy applications of Ca and Mg fertilizers compete
with K for exchange sites, thereby reducing the availability and uptake of K (Lim
and Khoo, 1985). Potassium concentrations are lower in soils with low cation
exchange capacities, e.g. granites, sands, highly leached and acidic soils, and are
higher in clay soils (Lim and Khoo, 1985; Gourley, 1999).

The main K fertilizers used in mango production are potassium chloride (KCl)
also known as muriate of potash, potassium sulfate (K 2SO4) and potas-sium nitrate
(KNO3). Potassium sulfate is generally preferred to KCl because it has a neutral
effect on pH and because mangoes are very sensitive to chlo-rides. Mango trees
take up K as the K+ ion from the soil solution. Potassium is readily redistributed,
typically from old leaves to young growing tissues. The highest concentrations of
K occur in leaves, fruit, roots and bark (Vuuren and Stassen, 1997). Leaf K
concentrations are generally at their highest at the end of the postharvest summer
vegetative flushing period, decreasing with flowering and fruiting (Catchpoole and
Bally, 1995; Medeiros et al., 2004). Mango trees can absorb more K than is
required for maximum tree growth; excessive uptake of K can cause imbalances
with other minerals in the tree (Gourley, 1999).

Potassium deficiency and toxicity

Potassium deficiency first appears in older mango leaves as the tree redistrib-utes
K to young growing tissues. Sen et al. (1947) induced K deficiency by growing
mangoes in sand culture and observed brown necrosis on the leaf margins
extending from the leaf tip towards the base. Smith and Scudder (1951) also
generated K deficiency symptoms in sand culture and described them as irregularly
distributed yellow spots and necrotic areas along the margins of small, thin
attenuated leaves which are very persistent. Lim and Khoo (1985) described the
non-necrotic areas of the leaf as dull, yellowish green to light green; symptoms
usually develop in the dry season or when irrigation has been stopped.
414 I.S.E. Bally

Effects of K on crop production and fruit quality

Potassium nitrate applications just prior to and at the flowering stage promote
flowering, increase fruit set and fruit retention (Sergent and Leal, 1989; Lyan-naz,
1994; Ferrari and Sergent, 1996; Oosthuyse, 1997; Rojas and Leal, 1997; Sergent
et al., 1997; Saleh and El-Monem, 2003; Shinde et al., 2006). In the low and mid-
latitude tropics, KNO3 is used to stimulate out-of-season flowering; however, this
effect is lost in higher latitudes (Davenport and Núñez-Elisea, 1997; see
Davenport, Chapter 5, this volume). Shongwe and Roberts-Nkrumah (1997)
suggested the KNO3 effect on flowering is a result of lowering the tran-spiration
rate and increasing water use efficiency. Protacio (2000) suggests the KNO 3 effect
on flowering is primarily due to N stimulation rather than K and he postulated that
KNO3 overcomes the inhibitory effects of gibberellic acid (GA3) on starch
accumulation by elevating the N concentrations over the N threshold to
synchronize bud break from apices with an existing floral initial.
Potassium influences fruit quality of many species (Marschner, 1995);
however, in mango there are only a few studies that link K nutrition with increased
fruit quality. Shinde et al. (2006) observed that increased K fertiliza-tion increased
fruit weight (5.15%), ascorbic acid (26.99%), organoleptic score for texture,
flavour, colour and shelf life and reduced physiological weight loss (22.79%) and
spongy tissue (68.08%). Potassium in the form of monopotassium phosphate
(KH2PO4) suppresses powdery mildew on mango (Reuveni et al., 1998;
Oosthuyse, 2000a), but it is not clear if the effect is due to P or K.

12.8 Magnesium
Magnesium is a component of enzymes that transport P, develop green col-oration
in chlorophyll, activate other enzymes and is involved in carbohydrate metabolism
and nucleic synthesis (Marschner, 1995; Stassen et al., 1999).

Sources, uptake and translocation of Mg

Magnesium is present in soil as the minerals biotite, serpentine, olivine and


hornblende, in organic matter, and as exchangeable Mg2+ in the soil solution.
Mg2+ is the second most dominant cation after Ca2+ and successfully com-petes
with K+ and Na+, replacing them on exchange sites (Aitken and Scott, 1999).
Magnesium is a mobile element that is taken up by the plant as Mg 2+. Magnesium
concentrations in the tree can be depressed by high concentra-tions of other
cations, such as Mn2+, Ca2+, K+, NH4+ and Al3+, in the root environment (Aitken
and Scott, 1999).

Magnesium deficiency and toxicity

Magnesium deficiency first appears in older leaves, due to its high mobility within
the tree, as pale green or yellow leaves with the inner vein areas of the
Crop Production: Mineral Nutrition 415

leaf lamina becoming mottled and necrotic while the leaf veins remain green.
Young trees are stunted by Mg deficiency. Smith and Scudder (1951) gener-ated
symptoms of Mg deficiency in sand culture and described them as a green wedge
pattern formed by the lateral intrusion of a bronzed chlorosis along the leaf margin
(Plate 75). Deficiency symptoms are more likely to occur in textured, sandy or
highly leached soils due to their low cation exchange capacity or when heavy
applications of liming products, K fertil-izers or green manuring have been applied
(Stassen et al., 1999). Magne-sium deficiency has been associated with a browning
skin discoloration in ‘Kensington Pride’ fruit.

12.9 Sulfur
Sulfur is a macro element required in large amounts by the tree. Sulfur is an
essential component of the amino acids cystine and methionine that make up
photosynthetic proteins, the vitamins thiamine and biotin and coenzyme-A used in
the synthesis and breakdown of fatty acids (Salisbury and Ross, 1992).

Sources, uptake and translocation of S

Up to 70–90% of S in soil is present in organic matter (Marschner, 1995), and


sulfates that are available for plants are mainly found in the surface layers and the
soil solution with only small amounts adsorbed to soil colloids. As a result, S is
very prone to leaching. High rainfall, light-textured and dry soils are adverse to S
accumulation and uptake (Anonymous, 1988).
Supplementary sources of S include sulfates as minor components of fer-
tilizers, i.e. superphosphate, sulfate of ammonia, potassium sulfate and gyp-sum as
well as some fungicides and miticides. Organic-based sulfate compounds are not
available to plants until they have been converted to the sulfate ion (SO 42–) by
bacteria in the mineralization process. The divalent sul-fate anions (SO42–) are
actively taken up by the roots and translocated to shoots or growing tissues. Sulfur
can also be adsorbed through the leaves as sulfur dioxide (SO 2), usually from
pollution from burning fossil fuels. Sulfur is not readily re-translocated from one
tissue to another. The competitive effects of other anions such as phosphates and
nitrates can reduce the uptake of S (Marschner, 1995).

Sulfur deficiency and toxicity

Sulfur deficiency in mango is uncommon as it is generally present in ade-quate


quantities in most soils and is a component of many fertilizers, fungi-cides and
miticides. Smith and Scudder (1951) describe S deficiency in mango as lateral
necrotic spots that occur on the vascular bundles and lamina on a very deep-green
leaf and premature defoliation. In other species symptoms
416 I.S.E. Bally

often appear to be similar to N deficiency, but are distributed more evenly between
young and old leaves, with the younger tissues developing symp-toms first as S is
not easily redistributed from older to younger tissues (Salisbury and Ross, 1992;
Marschner, 1995).
Sulfur toxicity occurs when pollutants high in SO2 content are converted to
bisulfate (HSO3–) when combined with water, and is further oxidized to sulfuric
acid (H2SO4), often known as acid rain. Sulfur dioxide adsorbed by leaves reacts
with water to form HSO3– that is toxic to the leaf, inhibiting photosynthesis and
degrading chlorophyll (Marschner, 1995).
Klumpp et al. (2003) reported soil S concentrations of between 53.2 and 86.0
mg/kg DW in contaminated soils in the vicinity of a copper smelter in Brazil
compared with 33–48 mg/kg DW in uncontaminated soils. Leaves from mango
trees growing in the contaminated soils had S concentrations of 3.8 mg/g DW,
double that of trees growing in uncontaminated soils (1.9 mg/g DW). At these
concentrations no obvious leaf symptoms were visible, indicating that mango has a
high tolerance of S in the soil and atmosphere. Information on mango leaf S
concentrations is scarce; however, a wide range of concentrations are consid-ered
to be optimal for mango growth (Table 12.1): 0.6–6.4 mg/g DW S (Marchal et al.,
1991), 1.07–1.69 mg/g DW S (Pimplasker and Bhargava, 2003). Chaudhary and
Nauriyal (1988) induced leaf deficiency symptoms in 1-year-old seedlings in sand
culture when leaf S concentrations were between 0.32% and 0.74% DW.

12.10 Zinc
Zinc is chemically bound to Fe and Mn to form components of chlorophyll, and is
essential for photosynthesis (Weir and Cresswell, 1995). Zinc is essential for the
synthesis of proteins and hormones, including auxins, and is required for the
maintenance of biomembranes (Salisbury and Ross, 1992; Marschner, 1995).

Sources, uptake and translocation of Zn

Zinc is present in soil in a range of inorganic minerals and organic matter with the
solubility of the inorganic forms decreasing as soil pH increases. Common Zn
fertilizers include zinc oxide (ZnO, 80% Zn), zinc chelates (ZnNa 2 EDTA and
Zn(NH4)2 EDTA, 6.5–14% Zn), zinc sulfates (ZnSO4, 23% Zn). Zinc is taken up
by the tree in the form of the zinc ion (Zn2+), and trans-located to regions of
growth such as shoot tips. Zinc is not readily translo-cated to other tissues. Zinc is
readily taken up through the leaves, and foliar Zn application is a common method
of managing Zn in mango orchards.

Zinc deficiency and toxicity

Zinc deficiency is often referred to as ‘little leaf’ because leaves fail to reach full
size. Symptoms first appear in immature leaves in the coloured stage, as
Crop Production: Mineral Nutrition 417

a thickening of the leaf and failure to fully expand. Leaf expansion is often stunted
on one side of the blade, causing it to form a sickle shape (Dilly et al., 1997) (Plate
76). Symptoms in fully mature leaves are prominent light-yellow or olive-green
veins on the upper surface that are thick and brittle. Internode length is reduced,
thereby causing a rosetting effect (Lim and Khoo, 1985; Agarwala et al., 1988;
Marschner, 1995). Zinc and Fe deficiency often occur together because of their
association with calcareous high pH soils.

Effect of Zn on crop productivity

Singh and Rajput (1977) reported that foliar sprays of 2.0–8.0% ZnSO4 prior to
flowering increased the number of perfect flowers in panicles and later increased
fruit yield, fruit sugar, ascorbic acid and total soluble solids (TSS). Daulta et al.
(1981) also observed that foliar Zn sprays increased fruit set and improved fruit
quality, but had no effect on sex ratio. Kumar and Kumar (1989) demonstrated that
1% ZnSO4 sprays applied to ‘Dashehari’ trees improved postharvest fruit life by
reducing weight loss, spoilage, increasing sugars, reducing acidity and slightly
increasing vitamin A. Littlemore et al. (1991) observed that 1% ZnSO4 foliar
sprays applied quarterly to ‘Kensing-ton Pride’ was sufficient to maintain leaf Zn
concentrations above critical concentrations and avoid symptoms of little leaf.
Although leaf Zn concen-trations were improved and symptoms cured, no effect on
tree yields was observed. Soil applications are often ineffective due to adverse soil
conditions making it unavailable to trees.

12.11 Manganese

Role of Mn

Manganese is a trace element that is required in small amounts by the tree; it is


important in the redox processes and is a cofactor, activating many enzymes that
catalyse oxidation, reduction, decarboxylation and hydrolytic reactions. Manganese
is an essential mineral for photosynthesis and the biosynthesis of proteins,
carbohydrates and lipids (Marschner, 1995).

Sources, uptake and translocation of Mn

Manganese is found in soil in the insoluble form of manganese oxide (MnO 2)


which is reduced to the water soluble exchangeable ion (Mn 2+) that is taken up by
tree roots. Mn2+ is mainly found in the soil solution but can also be loosely bound
to organic matter and clay colloids. Reduction of MnO 2 to Mn2+ is carried out by
organic reducing agents in aerobic soils and is favoured by reduced pH. As the soil
pH increases, microbial oxidation of Mn2+ increases. Both reduction and oxidation
processes occur in the soil simultaneously, but
418 I.S.E. Bally

the balance of available Mn is governed by soil pH, with low pH favouring


reduction and high pH favouring oxidation (Peverill et al., 1999).

Manganese deficiency and toxicity

Manganese deficiency in mango is initially expressed by light necrosis between the


main veins of middle and younger leaves that later become necrotic. Buff-coloured
spots appear on the primary leaf veins at the margins of the leaves and coalesce to
form necrotic patches before leaf abscission. In severe Mn deficiency, necrosis
begins at the leaf tip, spreading to the base of the lamina (Smith and Scudder,
1951; Agarwala et al., 1988).

Effect of Mn on crop production

There are few reports on the effect of Mn on crop production or fruit quality.
Schaffer (1994) suggested that mango decline was associated with trees that are
deficient in Mn and Fe.

12.12 Iron
Iron is a trace element that is required in small quantities, and is a component of
several enzymes and molecules. It is a component of chlorophyll and leaf Fe
concentrations directly impact photosynthesis. Iron-containing enzymes participate
in oxidation processes that release energy from sugars and starches, the conversion
of nitrates to ammonia and the biosynthesis of eth-ylene (Marschner, 1995). In
proteins, Fe acts as an electron carrier by alternate oxidation and reduction between
Fe2+ to Fe3+ (Salisbury and Ross, 1992).

Sources, uptake and translocation of Fe

Available Fe occurs in the soil solution in very low concentrations (~10 –15 M) as
the ionic forms Fe2+ and Fe3+. Fe2+ is far more soluble and more easily taken up
than Fe3+; however, in well-aerated soil, Fe2+ is readily oxidized to Fe3+ and
precipitated, becoming unavailable. Fe2+ can also be displaced by other cations or
by alkaline soils (Salisbury and Ross, 1992). Plants partly overcome Fe
unavailability by producing ligands that bind to Fe3+ to form soluble chelates that
maintain Fe in solution. On the root surface the Fe3+ is reduced to Fe2+ which is
immediately adsorbed by the roots. This process is often inhibited in high pH soils
and in calcareous soils. Iron deficiency is com-mon in mango (Schmidt, 1999).
Iron is readily absorbed through leaves, which is a useful method for management
of Fe when soil conditions are not conducive to uptake.
Crop Production: Mineral Nutrition 419

Iron deficiency and toxicity

Iron deficiency initially appears in young leaves as reduced concentrations of


chlorophyll. Iron-deficient leaves are pasty yellow/green with developing chlorosis
of the whole leaf as severity increases (Plate 77). Leaves fail to develop to full size
and necrosis of the leaf tips occurs in severe cases. Iron deficiency is often seen on
calcareous soils with >20% calcium carbonate (CaCO3), with poor aeration and
drainage in the coolest part of the year (Marschner, 1995). Iron deficiency reduces
the ability of mango trees to photosynthesize, thereby stunting tree growth and
reducing yields.

12.13 Calcium

A major role of Ca is membrane stability, which is achieved when Ca2+ binds to


phosphate and carboxylate groups of phospholipids and proteins on the surface of
the plasmalemma, and thereby preventing leaking of solutes to the cytoplasm
(Kirkby and Pilbeam, 1984). Calcium protects cells from toxins, slowing the aging
of plant tissues and promoting longer shelf life of many fruits. Calcium is
important for pectin polymers that strengthen cell walls and provide defence from
pathogens (Ferguson, 1984). Most Ca in trees is fixed in cell walls and is not easily
translocated. Calcium is essential for new root hair and leaf development.

Sources, uptake and translocation of Ca

Soil Ca occurs as exchangeable, non-exchangeable and soluble forms.


Exchangeable Ca makes up 65–85% of the total exchange capacity of normal soils,
and is weakly bonded, allowing rapid exchange with the soil solution, and is thus
prone to leaching (Kirkby, 1979; Mengel and Kirkby, 1987). Non-exchangeable Ca
is bound to minerals, for example feldspars, amphiboles, phosphates and
carbonates. Soil Ca concentrations are often up to ten times higher than other
cations such as K+ and Mg2+ and usually similar or slightly higher than sodium
ions (Na+) (Mengel and Kirkby, 1987).
Calcium is taken up passively through limited unsuberized tips of actively
growing roots and moves from the root cortex to the stele mainly through the
apoplast or free space of the root tips. It then enters the xylem, moving upwards
with the mass transpiration flow (Bangerth, 1979; Mengel and Kirkby, 1987; Ho
and Adams, 1989; Marschner, 1995). Root Ca concentra-tions are lower than other
cations such as K+ and Mg2+ because it is passively taken up and easily replaced
from the exterior surface of the plasma mem-brane by other cations (Marschner,
1995). Root Ca concentrations can there-fore be influenced by the relative
concentrations of other cations (K+, Mg2+, NH4+) in the soil solution. The
suppression of Ca uptake by high soil concen-trations of NH4+ may account for the
link between high N nutrition and many Ca-related disorders. On the other hand,
high NO3– can stimulate the uptake
420 I.S.E. Bally

of cations by stimulating organic anion synthesis, which attracts Ca 2+ (Kirkby,


1979; Mengel and Kirkby, 1987).
Calcium has low mobility in the phloem (Bangerth, 1979; Kirkby and Pilbeam,
1984), limiting translocation within the tree. Ca required by devel-oping organs is
supplied via the xylem. Calcium deficiencies are often caused by low availability,
poor uptake and/or poor translocation within the plant. Root pressure can also
contribute to the mass flow, but the extent to which root pressure contributes to Ca
flow in mango is unclear.
The intensity of evaporation from any organ will directly influence the amount
of Ca the organ receives (Marschner, 1995). Low transpiring plant organs, such as
fruit, are more prone to Ca deficiencies than actively transpir-ing leaves. This is
especially the case if the organ is fast growing and requires high amounts of Ca
(Mengel and Kirkby, 1987; Marschner, 1995). Limited Ca translocation occurs in
the absence of mass flow through exchange adsorp-tion in the xylem vessels,
similar to an exchange column (Mengel and Kirkby, 1987). As tissues grow, the
cation exchange sites of the new cell walls act as a sink for the Ca in the xylem
exchange column, drawing the Ca up the column (Clarkson, 1984). This exchange
movement of Ca may be important for lower transpiring organs such as the shoot
apex and fruit (Mengel and Kirkby, 1987; Marcelle et al., 1990). The translocation
of Ca is induced by the polar move-ment of indole-3-acetic acid (IAA) that is
produced in the shoot apex and rapidly growing organs (Banuelos et al., 1987; Ho
and Adams, 1989).

Calcium deficiency and toxicity

Calcium deficiency symptoms reflect the low mobility of Ca within the tree, first
appearing in young, actively growing tissues. Ca-deficient plants generally display
symptoms of membrane degeneration, which have been likened to senescence and
fruit ripening (Fallahi et al., 1977; Bangerth, 1979). Many symp-toms of internal
disorders in mango appear to be associated with premature ripening or cell
degeneration (Burdon et al., 1991, 1992). Raymond et al. (1998a) observed that
cell disruption and rupture of the cell walls were the first micro-scopic indicators of
soft nose, stem-end cavity and jelly seed. High Ca concen-trations reduce the
binding sites for enzymatic degradation (Rhodes, 1980). No direct symptoms of Ca
toxicity have been recorded in mango, but high concen-trations of Ca in soils can
displace other minerals such as Mg, Zn, B, Cu and P.

Effects of Ca on crop production and fruit quality

In mango, fruit Ca has been implicated as a factor in fruit quality, for example with
regards to internal physiological disorders, shelf life and disease resis-tance. Many
symptoms of internal physiological disorders appear to be asso-ciated with low Ca-
related membrane degeneration, i.e. premature ripening (Burdon et al., 1992;
Raymond et al., 1998a). Links between internal break-down and Ca nutrition in
mango were reported by Young (1957), who found
Crop Production: Mineral Nutrition 421

that increasing Ca fertilization generally resulted in decreased incidence of soft


nose. Young and Miner (1961) and Shear (1975) suggested a link existed between
Ca and N with respect to physiological disorders by demonstrating the negative
effect of N on internal breakdown could be partly overcome by high Ca
concentrations in the fruit. When Malo and Campbell (1978) were unable to alter
internal breakdown in ‘Tommy Atkins’ by N or K fertilization over a 4-year
period, the high Ca in the calcareous soils of Florida was suggested as a possible
reason.
Recent investigations have monitored Ca concentrations in fruit as well as in
soil. Subramanyam et al. (1971) observed that fruit with spongy-tissue disorder had
Ca concentrations of 74 mg/100 g DW, compared with 85 mg/100 g DW for
healthy fruit. Rane et al. (1976), Kadiyala (1995) and Tarmizi et al. (1993) also
reported significantly lower Ca concentrations in fruit with physiological disorders.
Burdon et al. (1991, 1992) measured lower Ca concentrations (5.85 mg/100 g fresh
weight (FW)) in the distal portions of ‘Kent’ fruit with soft-nose symptoms than in
healthy fruit (8.02 mg/100 g FW), and concluded that physiological disorders are
associated with Ca defi-ciency, but that Ca may be only one of many causal factors
and not necessar-ily the primary cause. There have been other reports that
increased Ca concentration is associated with physiological disorders; however,
these have been attributed to mineral redistribution after tissue breakdown (Burdon
et al., 1991; Raymond et al., 1998b).

Comparing fruit Ca data from different publications is difficult, as some quote


Ca concentrations on a FW basis and others on a DW basis, and many authors do
not specify whether they measured whole fruit or fruit pulp. None the less,
evidence allows the conclusion that low Ca concentrations are a causal factor of
physiological disorders in mango fruit.
There are mixed reports of the effect of fruit Ca on postharvest fruit
breakdown in mango. Singh et al. (1987) reported that spray-to-runoff appli-
cations of calcium chloride (CaCl2, 36% Ca) and calcium nitrate (Ca(NO3)2, 18%
Ca) at rates between 5 and 20 ml/l a week before harvest, increased shelf life of
‘Alphonso’ mangoes by 10 days. However, the authors did not discuss whether the
extended shelf life was due to delayed ripening or to reduced postharvest disease.
When Duttaray et al. (1993) attempted to repeat this experiment to control stem-
end rot caused by Diplodiia natalensis Pole Evans using 2500–5000 μl/l Ca as
CaCl2 and 450–900 μl/l Ca as Ca(NO3)2, they sig-nificantly increased the
incidence of stem-end rot. Simmons et al. (1997) used preharvest Ca sprays and
manipulation of leaf:fruit ratios to increase fruit Ca, but were unable to
significantly reduce postharvest disease.

Calcium in leaves and fruit

Leaf concentrations of Ca increase with age, and are considered to be ade-quate if


they are between 2–3.5% DW in acid soils and 3.0–5.0% in alkaline soils (Reuter
and Robinson, 1997). Leaf Ca concentrations generally do not correlate well with
fruit Ca concentrations (Bally, 2007).
422 I.S.E. Bally

Calcium concentrations in mango fruit can vary between and within fruit.
Gunjate et al. (1979) measured a gradient in Ca concentration from the stem end
(122 mg/100 g DW) to the apex (110 mg/100 g DW) of ‘Alphonso’ fruit, and from
the skin (142 mg/100 g DW) to the centre (130.4 mg/100 g DW) and to the seed
(128.4 mg/100 g DW). Gradients of Ca concentrations from the inner-mesocarp to
the outer-mesocarp, and from the stem end to the fruit apex have been reported
(Burdon et al., 1991; Tarmizi et al., 1993; Shorter and Joyce, 1998). Variation in
Ca concentrations within fruit tissues may be related to the position of the vascular
tissue and the transpirational properties of the tissue that influence the diffusion
path of Ca (Shorter and Joyce, 1998).
Concentrations of Ca on a DW basis increase rapidly during early fruit
development (cell division stage) followed by a gradual reduction later (cell
expansion stage); however, concentrations vary between reports. Gunjate et al.
(1979) measured Ca concentrations in ‘Alphonso’ of 241 mg/100 g DW at fruit set
decreasing to 68 mg/100 g DW at harvest. Chattopadhyay and Sarkar (1990) also
observed a similar decline, but measured higher concen-trations of 600–700
mg/100 g DW Ca at fruit set and 350–450 mg/100 g DW Ca at harvest in four
cultivars. More recently, Bally (2007) measured fruit mesocarp Ca between 0.22%
and 0.33% 30 days after fruit set and between 0.08% and 0.14% at harvest (155
days after fruit set) in ‘Keitt’. Both Gunjate et al. (1979) and Bally (2007) also
noted large fluctuations in Ca concentra-tions in the first half of fruit development
that reflected imbalances in supply and demand of developing fruit.

12.14 Boron
Boron binds as cis-diol borate complexes to mannose and certain other sug-ars in
cell wall polysaccharides and also has a role in sugar movement in the tree. Boron
also is important in nucleic acid synthesis (Salisbury and Ross, 1992), and is
essential for pollen and flower development, pollen germina-tion and pollen tube
growth (Stanley and Lichtenberg, 1963; Gupta et al., 1985) and is thus essential for
mango fruit set. Boron also is important in the synthesis of proteins that translocate
sugars (Gupta et al., 1985).

Sources, uptake and translocation of B

Most B in soil occurs as axenite, tourmaline, ulexite, colemenite and kermite; they
are relatively insoluble and are released very slowly so that small amounts are
available to plants (Gupta et al., 1985). Plant-available B is mainly associated with
soil organic matter and soil solution and is dependent on soil physical and chemical
properties, with the highest concentrations found in soils derived from marine
sediments, and the lowest concentrations in light sandy soils derived from granites
(Weir and Cresswell, 1995; Peverill et al., 1999). Low B concentrations are
associated with low organic matter, high rainfall or irrigation, high pH, high Ca and
dry soils (Gupta et al., 1985).
Crop Production: Mineral Nutrition 423

Supplementary forms of B include sodium borate as borax (Na 2B4O7, 10% B) and
as solubor (Na2B8O13·4H2O, 20% B), boric acid (HBO3-, 18% B), calcium borate
(Ca3(BO3)2, 12% B) and calcium-sodium borate (CaNa3B5O10, 18% B).
Boron is primarily taken up by the roots as un-dissociated boric acid (B(OH)3)
and transported in the xylem vessels by mass flow, accumulating in organs with the
highest transpiration rates, that is leaves and growing shoots. There is only limited
redistribution of B via the phloem (Raven, 1980); how-ever, this limited
redistribution and xylem supply is usually sufficient for normal tree health (Shelp
et al., 1995).

Boron deficiency and toxicity

Boron deficiency is expressed in tissues that are rapidly expanding and have low
transpirational rates, for example roots, fruits and shoots (Shelp et al., 1995). In
mango, B deficiency can result in poor flowering, pollination and reduced fruit set.
It is expressed in growing shoots by uneven cell division, causing leaves to grow
lop-sided with a curved sickle shape and deformed lamina and margins (Plate 78).
Leaves often have shot-holes that are sur-rounded by a light-green halo and ragged
margins. Apical dominance can be lost with swelling of the internodes. The main
raceme of panicles can develop a slight bend or kink towards the tip and in some
cultivars the bark splits and oozes black gummy sap from the cracks, known as
gummosis (Plate 78) (Nartvaranant et al., 2002). Agarwala et al. (1988) generated
B deficiency in 1-year-old seedlings using sand culture, and described mild
deficiency symp-toms, as mild chlorosis with a marked reduction in length and
width of the leaves. In more severe cases, older leaves become chocolate-brown at
the base, spreading to the tip before becoming completely chlorotic. Stems turned
black, lost apical dominance and eventually stopped growing. Boron deficiency can
be aggravated by high N status of trees (Ram et al., 1989; Raja et al., 2005).

In mango fruit, B deficiency causes cracks that split open; there is brown
discoloration of the mesocarp. Lumpy, deformed fruit may also be a symp-tom of
B deficiency. Meneses et al. (1994) examined normal and deformed fruit using
neutron capture radiography, and suggested that the deformities were due to B
toxicity. Boron toxicity is common in mango with typical symp-toms appearing in
leaves as dark spots on leaf margins that coalesce, eventu-ally leading to marginal
leaf necrosis in more severe cases (Plate 79). Boron toxicity often occurs after
excessive application of B fertilizers. Symptoms can be amended through leaching
of fertilizer from the root zone, raising soil pH with applications of lime or
stimulating growth through application of N; however, these measures may have
other implications for crop production.

Effect of B on crop production and fruit quality

Boron deficiency symptoms of trunk gummosis in ‘Kyo Savoy’ and ‘Nam Doc
Mai’ in Thailand were successfully remedied by soil applications of 20 or
424 I.S.E. Bally

25 g/m borax (11% B) during the summer wet season; however, the response time
and effect on gummosis of the two cultivars differed (Nartvaranant et al., 2002).
Foliar application of B solutions at the pre-flowering stage increased yield and fruit
quality in several studies. Dutta (2004) observed that 3000 ppm boric acid is
optimal for maximizing yield and quality of ‘Him-sagar’ in West Bengal, India.
Coetzer et al. (1991) reported that foliar applica-tion of B at flowering raised leaf B
concentrations to 60 mg/kg and increased yields from 14 to 22 kg/tree as well as
improving fruit quality. When Loría-Meneses et al. (1992) applied boric acid to the
skin of developing fruit they found it was not significantly translocated into the
mesocarp and remained in the surface layers of the skin around secretary glands.
The response of mango trees to soil applications of B varies between cultivars.
Rossetto et al. (2000) observed that ‘Winter’ was least sensitive, whereas, ‘Tommy
Atkins’, ‘Haden’ and ‘Van Dyke’ in declining order were more sensitive.

12.15 Conclusion
Our understanding of mineral nutrition in mango is incomplete and lacks detailed
knowledge of the effects of many minerals, as demonstrated above. A better
understanding has generally come from observations of the effects of minerals on
fruit quality rather than yield. Studying the effects of minerals on mango
production and fruit quality is difficult because of their delayed response to applied
minerals and large tree reserves. The effects of minerals on productivity and fruit
quality can be greatly enhanced by also determin-ing their effects on other
phenological events that contribute to productivity and fruit quality. More regular
and targeted assessment of mango tree min-eral status will facilitate improved
management of mango trees. This may require the determination of fruit mineral
concentration standards for differ-ent stages of fruit development. Development
and employment of low-cost, rapid and non-destructive techniques to measure
minerals will enable mineral fluxes within trees to be monitored closely.

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13 Crop Production: Management
J.H. Crane,1 S. Salazar-García,2 T.-S. Lin,3
A.C. de Queiroz Pinto4 and Z.-H. Shü5
1
University of Florida, Florida, USA
2
Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias, Santiago
Ixcuintla, Mexico
3
National Taiwan University, Taipei, Taiwan
4
Private Consultant on Tropical Fruits, Brasilia, Brazil
5
Meiho Institute of Technology, Pingtung, Taiwan

13.1 Introduction 432


13.2 Production Areas and Yields 434
13.3 Climate of Production Areas 436
13.4 Soils and Soil Preparation 440
13.5 Plant Propagation and Rootstocks 443
13.6 Major Cultivars 445
13.7 Plant Spacing 449
13.8 Fertilizer Practices 452
13.9 Irrigation Practices 460
13.10 Vegetative Growth and Reproduction Manipulation 463
13.11 Environmental Stress Management 468
13.12 Harvesting Practices 470
13.13 Conclusions 472

13.1 Introduction
Mangoes have been in cultivation for several thousand years (Mukherjee, 1953,
1972; Kostermans and Bompard, 1993), and crop production practices have
continually improved (Singh, 1960, 1978; Crane et al., 1997). A detailed
understanding of mango plant physiology and behaviour in relation to cli-matic and
edaphic conditions, genetics and cultural practices dates back to the late 1950s
(Mukherjee, 1953; Chadha and Pal, 1986; Chacko, 1989; Whiley, 1993; Schaffer et
al., 1994; Kulkarni, 2004; Davenport, 2006). Recent reviews (Cull, 1991; Whiley,
1993; Schaffer et al., 1994; Davenport and Núñez-Elisea, 1997; Kulkarni, 2004;
Davenport, 2006) on mango crop management and physiology illustrate that the
best prospect for improving mango production must involve a holistic approach to
cultural practices that considers the spe-cific climatic and edaphic environment,
cultivar and tree phenology for a
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
432 (ed. R.E. Litz)
Crop Production: Management 433

given production area. Most importantly, the development of a phenological


approach to understanding and managing mango orchards is very promis-ing from
a practical standpoint (Cull, 1991). The phenological approach to crop management
is based on monitoring environmental conditions and tree-growth stages and
conditions, and manipulating the tree through cul-tural practices for optimum
production (Fig. 13.1). Published research can be used to develop a holistic crop
management approach.
Mango production in India has been reviewed extensively (Singh 1960, 1978;
Majumder and Sharma, 1990; Kostermans and Bompard, 1993; Negi, 2000). The
purpose of this chapter is to discuss current concepts, strategies and innovations for
mango culture and to describe current production prac-tices in four commercial
mango-producing areas: Brazil, Mexico, Taiwan and the USA. Crane et al. (1997)
has described the cultural practices in Australia, Israel, Mexico and the USA.

The Brazilian and Mexican mango industries now account for about 62% and
8% of exports to the USA, respectively (Perez and Pollack, 2007). Improve-ments
in cultural practices in both countries have increased production effi-ciency and
exports. Taiwan’s mango industry, although relatively small, is innovative and
provides top-quality mangoes for the local and Asian mar-kets. Mango production
in the USA is small, and production is geared towards national speciality markets.

Fruit
development
Flowering and
fruit set
Flower bud Postharvest
development Unwanted vegetative flush
vegetative
flush
Harvest
Amount of development

Premature Root flush


fruit drop

Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct
n

s May Jun July Aug Sept Oct Nov Dec Jan Feb Mar Apr

Fig.13.1. Theoretical mango phenological cycle (Source: after Cull, 1991). s,


southern hemisphere; n, northern hemisphere.
434 J.H. Crane et al.

13.2 Production Areas and Yields


The mango industry of Brazil is extensive due to favourable soil and climatic
conditions. Bahia and São Paulo are the most important production areas of Brazil
with 34,600 and 28,800 ha, respectively, of the estimated 70,000 ha of mango
production in 2001 (Table 13.1). The north and south of Brazil account for only
6.0% of the total mango-production area (Souza et al., 2002). The most recent
mango census (2005 data) estimated mango production in Brazil to be c.970,700 t
(Gazeta, 2006; Agrianual, 2007), which is a slight increase since 2001. Bahia and
São Paulo also have the largest production, respec-tively (Table 13.1). Brazil
exports 133,300 t of mangoes, 13.7% of the total mango production. Although pest
and disease problems and the highly com-petitive internal and external markets
have limited the expansion of the mango industry in some Brazilian regions,
important technologies have been

Table 13.1. Regions of mango production in Brazil, Mexico, Taiwan and the USA
(Source: see table footnotes).

Country State Region Hectares Production (t)


a
Brazil Bahia North-east 19,000 306,000
Pernambuco 8,000 146,000
São Paulo South-east 18,000 245,000
Minas Gerais 7,000 130,000
Miscellaneous Other regions 18,000 143,000
b
Mexico Sinaloa Nayarit Northern and Central 106,740 1,138,361
Jalisco Colima Pacific region
Michoacán
Guerrero
Oaxaca Southern Pacific region 42,180 367,257
Chiapas
Tamaulipas Gulf of Mexico region 28,778 191,996
Veracruz
Compeche
Miscellaneous Other states 7,827 37,151
c
Taiwan Tainan Central-South 18,200 191,332
Pingtung South
Miscellaneous Other prefectures
d
USA Florida Southern region 324 5,986
Puerto Rico South, South-West Coast 1,079 14,206
California Coachella Valley 105 1,905
Hawaii Islands of Hawaii, Oahu, 142 317
Maui, Kauai
a
Source of data on Brazil: Agrianual (2006, 2007); Anonymous
(2006). b Source of data on Mexico: SIIAP (2007).
c
Source of data on Taiwan: Agriculture and Forestry Department (1996).
d
Source of data on USA: Linden (2006); Anonymous (2007); J.H. Crane, personal communication.
Crop Production: Management 435

developed to improve and support the mango industry (Santos Filho et al., 2002).
This has resulted in an increase of 58% in mango exports from 1998 to 2005
(Souza et al., 2002; Agrianual, 2007).
There are c.181,525 ha of mangoes in 23 of Mexico’s 32 states (SIIAP, 2007);
there are four large mango-producing regions (Table 13.1), ranging from 1433N
to 27N latitude. They are distinguished by differences in cli-mate, soils and
cultivars. There are 106,740 ha of mango in the Northern and Central Pacific
region; the major cultivars are ‘Tommy Atkins’, ‘Ataulfo’, ‘Kent’, ‘Haden’ and
‘Keitt’. The Southern Pacific region has 42,180 ha and produces ‘Manila’
(‘Carabao’), ‘Ataulfo’, ‘Tommy Atkins’, ‘Haden’ and ‘Kent’. The leading cultivars
of the Gulf of Mexico region (28,778 ha) are ‘Manila’ followed by ‘Tommy
Atkins’. Mango production in Mexico was about 1.7 million t in 2006 and the
states of Sinaloa, Guerrero, Nayarit, Oax-aca, Chiapas, Veracruz and Michoacán
accounted for 92% of the crop (SIIAP, 2007). All these states except Veracruz
(which abuts the Gulf of Mexico) are situated along the Pacific coastal and inland
areas.
In 2006 Mexico exported 196,120 t to the USA valued at >US$132 million
(EMEX, 2007). Mexico supplies 87.5% of the mangoes consumed in the USA and
mango exports in 2006 were 17.9% higher than in the 2005. The major export
cultivars are ‘Tommy Atkins’ (33%), ‘Kent’ (23%), ‘Ataulfo’ (19%), ‘Keitt’ (14%)
and ‘Haden’ (11%). None the less, most mango production (c.85%) in Mexico is
for the domestic market. The most important mango warehouses and distributors
are in Mexico City, Guadalajara and Monterrey.
The 7-month mango season in Mexico is due to differences in latitude,
phenological cycles, rainfall and soil moisture patterns and the use of growth
regulators. Chiapas, along the South Pacific coast, is usually the first to har-vest
mangoes (‘Ataulfo’). Full bloom of ‘Ataulfo’ occurs from mid-November to mid-
February and fruit is harvested from the end of January to the end of May. Mango
harvesting continues northwards from Chiapas to the states of Oaxaca, Guerrero,
Michoacán, Colima, Jalisco, Nayarit, Sinaloa and Sonora. Sinaloa is a major
mango producer with the latest harvest season for ‘Manila’ (July), ‘Haden’ (July),
‘Tommy Atkins’ (July), ‘Kent’ (July–August) and ‘Keitt’ (August–September).
The ‘Manila’ mango is produced mainly in Veracruz and Guerrero. Full bloom
occurs in January–February and fruit is harvested in May–June.

The production area in Taiwan increased from 17,000 ha in 1987 to 21,000 ha


in 1992 and stabilized at c.20,000 ha for c.10 years when the yield was 17 kg/tree;
however, as production has increased to c.27 kg/tree, the plant-ing area has
decreased to c.18,200 ha in 2006 (Table 13.1). There are six important mango-
growing prefectures in Taiwan: Nantou, Taichung, Chiayi, Tainan, Kaohsiung and
Pingtung from north to south in western and eastern Taiwan. Tainan and Pingtung
are the most important production areas, com-prising about 90% of the total
harvested area.
Mangoes are grown commercially in four areas of the USA: Florida (25–
28N), Puerto Rico (18–185´N), Hawaii (22–30N) and California (33–335´N)
(Table 13.1). Puerto Rico has the largest area (1079 ha) (Alvarado-Ortiz and Acin,
2004), followed by Florida (324 ha) (FASS, 2003; Anonymous,
436 J.H. Crane et al.

2007; J.H. Crane, personal communication), Hawaii (142 ha) (NASS-Hawaii,


2006; Anonymous, 2007) and California (105 ha) (Linden, 2006; Anonymous,
2007). Mango production in the USA is estimated to be only 22,414 t with Puerto
Rico accounting for >63% of the production (Alvarado-Ortiz and Acin, 2004;
Linden, 2006; Anonymous, 2007). The four areas have different climates which
present different production opportunities and challenges.
The main production areas of Florida are along the south-east coast in Miami-
Dade County (25–27N), the east (Palm Beach, Broward and Indian River
Counties) and west (Lee County) coasts. The season in Florida is from May
through to September. Puerto Rico is a Caribbean island (18N, 67W)
(Espenshade, 1992; Bahr and Johnston, 1993c). Most of the commercial pro-
duction is near Ponce on the south coast and Mayaguez in the south-west (23.9 ha).
The main mango season in Puerto Rico is May through to Septem-ber, although
some mangoes are harvested as late as mid-November. In Hawaii (19–22N),
production occurs on the islands of Hawaii, Oahu, Maui and Kauai. The season in
Hawaii is from May to August, although some pro-duction occurs year-round. The
California industry is on the western side of the Salton Sea in the Coachella Valley
of southern California (33N) (Scott, 1990). Mango production is from mid-August
to October (Linden, 2006).

13.3 Climate of Production Areas


Mangoes are mainly cultivated in the tropics (25N, 25S) and subtropics (35N,
35S), although limited production also occurs in warm temperate/ subtropical, i.e.
Mediterranean-type areas, for example Israel, southern Spain and the Canary
Islands and California. The ideal areas for mango production have a cool and/or
dry period prior to flowering followed by moderate soil moisture and moderately
hot temperatures (30–33C) (Chacko, 1986). The diversity of climates and soils in
mango-production areas reflects the adapt-ability of the species and improvements
in cultural practices.
Temperature and availability of water are the most significant environ-mental
factors that influence commercial mango production by affecting the frequency,
intensity, duration and time of vegetative growth and flowering (Chacko, 1986,
1989; Whiley, 1993; Schaffer et al., 1994; Núñez-Elisea and Davenport, 1995;
Davenport, 2006) and disease incidence (Johnson et al., 1989; Ploetz, 1994, 2003;
Ploetz et al., 1994). Temperatures <15C and >30C inhibit pollen tube
germination, thereby impeding fertilization and resulting in embryo abortion and
fruit abscission.
Mango production in Brazil extends from 3N, close to the Amazon region,
where a humid, hot tropical climate predominates almost year-round to
approximately 25S in Paraná, which has a subtropical climate. Between these two
extremes, there are various soil and climatic conditions where mango trees are
grown intensively. The semi-arid tropical climate of the north-east region of
Petrolina in Pernambuco (2113´N and 4750´W) is more appropriate for high-
quality mango production than Votuporanga in São Paulo (2113´S and 4750´W),
which has a subtropical climate (Table 13.2).
Table 13.2. Climatic parameters for Brazil, Mexico, Taiwan and the USA.

Temperature (C)a Annual mean rainfall (mm)a


Annual Summer Winter Annual Season

Yearly Yearly
Country State Region Climatic category mean Mean Mean mean Wet Dry

Brazilb Pará North region Tropical hot humid 26.0 31.5 22.0 2893 436.2 111.8
Goiás Central region Savannah dry tropics 29.8 31.9 17.7 1576 270.3 6.2
Bahia North-east region Tropical semi-arid 27.9 33.8 22.0 545 139.6 1.7

Crop Production: Management


Pernambuco Tropical semi-arid 26.2 32.0 20.5 570 136.2 4.8
Piauí Tropical hot subhumid 27.5 32.9 22.1 1449 286.3 11.6
São Paulo South-east region Subtropical cool 21.3 25.1 17.6 1623 127.1 40.0
Minas Gerais Tropical mesothermic 26.5 35.0 18.0 840 105.2 6.0
Paraná South region Subtropical very cool 17.6 22.7 12.4 1408 165.0 74.5
Mexicoc Sinaloa Northern and Warm subhumid 24.7 27.5 (22–38) 22.0 (13–29) 922 799 123
Central Pacific tropics
region
Nayarit Warm subhumid tropics 24.7 26.3 (24–38) 23.1 (13–30) 1396 1346 50
Colima Warm semi-arid tropics 26.5 27.6 (25–38) 25.4 (14–30) 660 605 55
Michoacán Very warm semi-arid 28.3 29.6 (28–40) 27.0 (18–32) 709 601 108
tropics
Guerrero Warm subhumid tropics 28.0 29.0 (28–40) 26.9 (18–32) 1011 910 101
Oaxaca Southern Pacific Isothermal warm 25.9 26.9 (26–35) 25.0 (18–28) 1779 1731 48
Chiapas region humid tropics 24.7 24.9 (26–35) 24.4 (18–28) 3732 3661 71
Central Gulf of Mexico Isothermal warm 24.8 26.7 (26–38) 22.9 (14–30) 1026 898 128
Veracruz region subhumid tropics
Southern Isothermal warm 25.5 27.6 (26–38) 23.4 (14–30) 2093 1807 286
Veracruz humid tropics
Taiwand Tainan Central-South Subtropical 23.6 26.8 (23.9–28.3) 19.1 (17.1–20.4) 1733 264 (71–415) 25 (15–41)
Pingtung South Subtropical 24.7 26.8 (24.9–27.9) 21.8 (20.4–22.7) 2158 329 (170–511) 30 (19–47)
Average Whole island Subtropical 25.0 25.8 (23.1–27.5) 19.1 (17.3–21.6) 1885 273 (92–392) 42 (24–60)

437
(Continued)
438
Table 13.2. Continued

Temperature (C)a Annual mean rainfall (mm)a


Annual Summer Winter Annual Season

Yearly Yearly
Country State Region Climatic category mean Mean Mean mean Wet Dry

USAe Florida Southern coasts Marine subtropics – 23.2 26.8 (26–27) 18.9 (18–19) 1643 1313 333
east coast
Marine subtropics – 23.3 27.1 (24–28) 18.0 (17–18) 1354 1064 290

J.H. Crane et al.


west coast
Puerto Rico South Coastal area Marine tropical 26.3 27.7 (23–32) 24.8 (19–30) 904 632 272
California Coachella Valley Dry warm subtropical 22.5 31.8 (30–33) 13.3 (11–15) 79.7 55.1 24.6
Hawaii Island of Hawaii Marine tropical 23.3 24.2 (16–33) 22.5 (13–33) 3279 1926 1353
Island of Oahu Marine tropical 25.1 30.1 (18–34) 24.0 (13–32) 559 414 145
Island of Maui Marine tropical 24.2 25.6 (15–35) 22.9 (11–32) 531 442 89
Island of Kauai Marine tropical 24.2 25.6 (17–32) 23.1 (12–31) 1091 698 393
a
Numbers in parentheses are ranges.
b
Brazil: rainfall figures for the wet and dry season are per
month. c Source of data on Mexico: García (1973).
d Taiwan: temperature ranges are averages for lowest and highest summer (April–October) and winter (November–March) months and the yearly mean; precipitation
is averages for wettest (May–October in Pintung; April–September in Tainan and island average) and driest (October–March) months and yearly
annual. Available at http://www.cwb.gov.tw/
e
Source of data on USA: Butson and Prine (1968); Getz (1979); E.E. Toro, personal communication; C.L. Chia, personal communication; Quayle et al.
(1995); Garczynski (1995); SERCC (2007).
Crop Production: Management 439

Some cultivars, for example ‘Haden’ in north-eastern Brazil, have poor fruit set
when temperatures are >35C.
Solar radiation is very important for fruit development, since its dura-tion and
intensity are directly related to photosynthesis and carbohydrate production
(Mukherjee, 1953). Light incidence depends on the season (Allen et al., 1998).
According to Lima Filho (2000), mango production in the semi-arid region of
Brazil is where maximum solar radiation occurs in summer (October southern
hemisphere; 528 cal/cm2/day) and the minimum in win-ter (June southern
hemisphere; 363 cal/cm2/day), which corresponds to the flowering and fruit
development stages, respectively.
Mango production in Mexico is in the tropics. Most rainfall occurs dur-ing the
summer and may be accompanied by hurricanes (Table 13.2). Climate in the
Northern and Central Pacific region (17–27N) ranges from warm sub-humid to
warm/very warm semi-arid tropics with a 7-month dry season (García, 1973). In
Colima and Michoacán, mangoes are irrigated due to low annual precipitation
(semi-arid condition with a 6- to 8-month dry season; the mean annual temperature
is 26.5–28.3C). The Southern Pacific and Gulf of Mexico coastal regions have
isothermal (< 5C monthly temperature oscil-lation) warm, subhumid to humid
climates with a 6–8-month dry season (García, 1973).

The Gulf of Mexico coastal region is affected by winds from the north (11–28
m/s) from October to April. These winds cause direct damage, such as limbs
breaking and flower and fruit drop, and also increase plant respiration and
transpiration rates that stress the trees. In the Pacific coastal regions hur-ricanes
usually occur from August to October. Recent, unusually warm win-ters have
resulted in undesirable late autumn or winter vegetative flushes and poor flowering.
Normal bloom occurs in late January and February; how-ever, late autumn or
winter shoots delay anthesis during hot spring tempera-tures (April to early May)
which result either in low fruit set or parthenocarpic fruit.

The primary production areas in Taiwan are from 22N (Fungshan, Ping-tung)
to 24N (Nanto and Taichung prefectures). The average temperature of these areas
is c.23C, with a mean of 20C during flowering (December– March) and c.18.6C
during development (April–August) (Table 13.2). Flower induction in mango is not
a problem in Taiwan because of its subtropical climate; however, fruit set can be
affected by low temperature, rainfall, etc. A monsoon-type climate prevails in
southern Taiwan with precipitation occurring mostly from May through to
September. There is little rainfall (usually <50 mm) during flower bud formation
and flowering (autumn-winter) (Table 13.2).

The production areas of the USA have different climates. South Florida has a
marine subtropical climate (Table 13.2). South-east Florida has a mean annual
temperature of 23C, a mean annual rainfall of 1643 mm and 62% relative
humidity (RH) (Butson and Prine, 1968; Getz, 1979; Barrick and Black, 1980).
Constant ocean-spawned winds of 4–9 km/h buffet the region from February
through to October. The wet season (two-thirds annual rain-fall) occurs during the
late spring into autumn (May–October, northern
440 J.H. Crane et al.

hemisphere) and the dry season occurs during winter and early spring (November–
April). Lowest temperatures (-4 to -6C) occur from December through to February
with a 70% probability of 0C at least once each year (Bradley, 1975).

The Puerto Rican industry is mostly along the south and south-west coast at
elevations between sea level and 50 m (Table 13.2) (Aponte-Morán et al., 1977).
The La Cordillera central mountain range divides the island from north to south
and has a major impact on the island’s climate, with annual rainfall of 1550 mm in
the north and 904 mm in the south (Espenshade, 1992; SERCC, 2007). In the main
production areas, the dry season is from Decem-ber to May, although July and
August are also dry (Aponte-Morán et al., 1977). Mean maximum and minimum
temperatures vary with altitude; how-ever, along the coastal production areas, 24–
29C is normal (Espenshade, 1992; SERCC, 2007). The lowest temperatures (18–
20C) occur along the coast during January and February.

The Hawaii mango industry is located mostly on the leeward coasts of Oahu,
Hawaii and Maui islands; however, limited production occurs on the windward
side of Kauai Island (Table 13.2). Each commercial planting has a distinct
climatological niche that varies with altitude and location with respect to
mountains and predominant north-east trade winds. Annual rain-fall is 531 mm on
Maui, 559 mm on Oahu and 3279 mm near Hilo (east coast of Hawaii) (Table 13.2;
C.L. Chia, personal communication). Temperatures, like rainfall, vary with
elevation and location but generally range between 21 and 27C (Bahr and
Johnston, 1993b). Lower (13C) and higher temperatures (32C) occur
occasionally.
The Coachela Valley of California is in the Salton Trough Desert area (Bahr
and Johnston, 1993a) (Table 13.2). The valley ranges from 80 m below sea level to
488 m above sea level (Espenshade, 1992; Aslan et al., 1993) and the climate is
arid. Temperatures in the valley vary considerably depending upon season,
elevation and exposure. Some data reported herein represent the average for the
area and may not accurately reflect the particular micro-climate of the mango
orchards in the valley (Garczynski, 1995). The mean annual temperature is 22.5C;
the average summer temperature is 31.8C, and the average winter temperature is
13.3C (Aslan et al., 1991; Garczynski, 1995). Winter temperatures range from
22.1C to 4.5C (Garczynski, 1995); however, Schacht (1992) reported a low of -
4.4C for 2 h. Summer tempera-ture extremes range from a low of 11C to 50C
(Garczynski, 1995). The aver-age annual rainfall is 79.8 mm with two-thirds of this
occurring during the winter (December–March) (Aslan et al., 1991; Garczynski,
1995). The valley is buffeted by wind and sandstorms during late spring, and these
can cause severe damage (Schacht, 1992; Aslan et al., 1993).

13.4 Soils and Soil Preparation


Mangoes tolerate many soil types (Majumder and Sharma, 1990; Crane and
Campbell, 1991; Kostermans and Bompard, 1993). Trees grow most vigorously
Crop Production: Management 441

in deep, fertile, moderately acid to neutral pH, loam-type soils. They tolerate
infertile sands, volcanic ash and limestone-based soils, excessively drained and or
periodically flooded soils and soils with acid (pH 4.5–7) to alkaline pH (pH 7–8.5).
Mangoes are sensitive to saline and sodic soil conditions and proper irrigation
practices and the use of salt-tolerant rootstocks is imperative for successful crop
production in some areas (Schaffer et al., 1994).
Land preparation includes clearing virgin or existing orchard trees, disk-ing,
destruction of subsurface hardpans (slip ploughing), mixing of upper and lower soil
profiles, crushing superficial bedrock and mixing rock and soil layers, formation of
land contours to facilitate drainage and/or flood irriga-tion, bedding and amending
soils with organic matter and inorganic chemi-cals, for example hydrated lime
(Ca(OH)2), dolomite and preplant fertilizers.
Mango is grown on a wide range of soil types, from latossols with a high
percentage of sand in the north-east of Brazil to loamy oxysols in the south-east.
Some areas in the north-east have shallow soils that need improved drainage. Soils
of the central region are chemically poor and acid (pH 3.7–4.7) and require lime
and/or gypsum application prior to orchard establishment. Soil analysis is used for
determining the suitability of soil for production and potential fruit quality,
especially in areas cultivated for export fruit. Typically soil analysis is conducted
prior to land clearing and ploughing to determine nutrient levels and the need for
lime and/or gypsum application. Soil sam-ples are taken from 0–20 cm and 20–40
cm depth of the soil profile. Soil sam-ples are usually taken at a 0–30 cm and 30–
60 cm soil depth for mature and established orchards. Ploughing, harrowing and
lime and/or gypsum incor-poration are recommended at 30 cm depth and at least
30 days prior to rainy weather (Pinto and Ramos, 1998). Liming is very important
for acid soils (<pH 4.0) in the central region of Brazil in order to increase soil pH
to 6.0–6.5. It also improves the soil base saturation to 60–70% and results in better
soil conditions for growth and production (Pinto, 2000). The amount of lime to be
applied is based on a basis saturation equation (where hydrogen (H), aluminium
(Al), calcium (Ca), magnesium (Mg) and potassium (K)):

Lime rate (t/ha): (T × 0.5) – S where T = (H + Al) + S and S = Ca + Mg + K


Gypsum applications are recommended for acid subsoils with >20% Al
saturation and <0.5 cmol/dm3 Ca at soil depth of 60 cm (Andrade, 2004). This has
been shown to significantly reduce fruit physiological disorders, such as soft nose.

Production areas of Mexico vary in soil characteristics, and include flat coastal
areas and steep mountain slopes from sea level to >600 m above sea level.
Orchards on sloped land commonly utilize individual tree terraces. Soil depth
ranges from 30 cm to >3 m (alluvial soils). Shallow and eroded soils are common
on hilly terrain. The presence of medium to large (10–50 kg) boulders can impede
the use of machinery but they reduce erosion and increase soil-water storage
capacity. In general, poor soil drainage is not a problem; however, areas with clay
soils have drainage problems during peri-ods of heavy rain, especially if there is a
shallow water table. A range of soil types is used for mango production in Mexico.
In the Northern Pacific region,
442 J.H. Crane et al.

mangoes are planted in cambisols (pH 6.0–7.0), luvisols (pH 6.5–7.5) and feozem
(pH 5.0–6.0) soils with loamy textures (Anonymous, 1970, 1982). These soils are
well drained, with 2–3% organic matter, moderate to high water holding capacity
and cation exchange capacity (CEC) of 10–20+ meq/100 g soil. In the Central
Pacific region mangoes are planted on cambi-sols, feozem and regosols with light
textures (Anonymous, 1970, 1982). The regosols (pH 6.5–7.5) are well drained,
with <2% organic matter content, low water holding capacity and CEC <10
meq/100 g soil. Soils planted to man-goes in the Southern Pacific region include
nitosols, luvisols and feozem (Anonymous, 1970, 1982). The nitosols (pH 5.0–6.0)
are well drained, with >3% organic matter content, low to moderate water holding
capacity and low CEC (<10–20 meq/100 g soil). Soils in the Gulf of Mexico region
include fluvisols, cambisols and vertisols of loamy and clayey textures
(Anonymous, 1970, 1982). The fluvisols and vertisols are moderately and slowly
drained soils, respectively. Soil pH for fluvisols ranges from 5.5 to 7.5 with <2%
organic matter content. Soil pH for the vertisols is alkaline (7.5–8.0) and the
organic matter content is moderate (2.1–3%). The water holding capacity and CEC
is low to moderate for the fluvisols (<10–20 meq/100 g soil) and high for the
vertisols (10–20+ meq/100 g soil).

Soil tillage is used in flat lands before planting. Dimensions of planting holes
range from 40–60 cm depth and 30–50 cm diameter in light-textured soils. Planting
holes can be larger in heavy-textured or stony soils. On steep hills only planting
holes are made and individual terraces are built up to hold soil, water, organic
matter, fertilizers or soil amendments. Preplant soil analyses are rarely taken;
however, chemical or organic fertilizers are com-monly applied at planting time.
With rapid expansion of the mango industry, orchards have been planted in shallow
soils (0.75–1.2 m depth) underlain with a hardpan. These soils are poorly drained
and root growth and exten-sion are limited. Trees growing in such soils are prone
to drought stress dur-ing dry periods, flooding stress after rains and nutritional
deficiencies. The weakened trees appear to be more susceptible to pathogens, for
example Botryodiplodia theobromae, which causes cankers or stem dieback and
eventu-ally tree death (Ponce-González and Salazar-García, 1992). Cultural
practices are not available to ameliorate these problems and planting in such soils
is not recommended.

Mangoes are grown in Taiwan on sandy loam, loamy sands, clay and coarse
sandy soils. Soil pH ranges from 5.0 to >7.8. Most trees are grown on sloping land.
Silt clay loam with pH 5.0–6.0 is mostly found in the Pingtung area, while the
Tainan area has clay soils with pH 7.3–7.8. Acid soils are amended with Ca(OH)2
or dolomite and alkaline soils are amended with sulfur (S) or acid-based fertilizers.

The topography in the mango-production areas of Florida is flat and ranges


from sea level to c.6.1 m above sea level. Soils in Florida include vari-ous sands,
muck and oolitic limestone. In the main production area (Miami-Dade County),
mangoes are planted in extensively scarified (crushed) and trenched oolitic
limestone rock (Krome and Chekika very gravelly loam) (Colburn and Goldweber,
1961; Noble et al., 1996). The limestone soil is very
Crop Production: Management 443

permeable (1.5–5.1 cm/h), with high pH (7.4–8.4), low organic matter con-tent (3–
10%), low water holding capacity (0.2–0.3 cm/cm of soil) and low CEC (16.0–37.0
meq/100 g soil) (Calhoun et al., 1974; Anonymous, 1989). In areas subject to
flooding, crushed rock is formed into beds (0.6–1.0 m high and 1.0–1.5 m wide)
before planting. The sandy soils in other Florida produc-tion areas are poorly to
well drained with or without an undulation hard-pan 0.5–3.0 m down in the soil
profile (Henderson et al., 1984). The highly organic muck soils in Palm Beach
County are poorly drained and underlain by dense limestone bedrock. Beds of
varying dimensions are made in the sandy and muck areas to increase the
proportion of the root system above flood levels. The sand and muck soils are
characterized by an acid to alka-line pH (3.6–8.4) and low cation exchange
capacities (Carlisle et al., 1978; Henderson et al., 1984).

In Puerto Rico, mangoes are grown on flat and gently sloping land con-sisting
of alluvial fans and terraces level with or slightly above the flood plain (Bierbolini
et al., 1979). Some orchards in western Puerto Rico are located on moderately
steep to very steep slopes (12–60% slope) and rounded hill tops that are somewhat
eroded and superficial (Bierbolini et al., 1975). Soils in southern Puerto Rico are
very deep, moderately well to well drained and consist of fine-textured sediment of
sandy and clayey loam over gravelly fine-textured sediment. The pH ranges from
moderately acid (pH 6) to alka-line (pH 8 or above). Soils have good to very good
native fertility and good water holding capacity. Soils in the west are well drained
and moderately acid (pH 5.0–6.0).

Mango production in Hawaii occurs on volcanic ash soils varying from recent
to highly weathered (R. Yost, personal communication). They tend to be well
drained; some soils must be slip ploughed to break up the hardpans in preparation
for planting. The pH ranges from 5 to 8 and the fertility varies substantially.
Mangoes in California are on lacustrine deposits consisting of fine-textured
sediments that are highly stratified sandy loam soils with clay lenses (S. Aslan,
personal communication). The soil is alkaline (pH 7–8.4) and calcareous (Aslan et
al., 1991). Land is slip ploughed to 1.5–1.8 m depth to break up and mix the
stratified soil layers.

13.5 Plant Propagation and Rootstocks


Mango production areas rely on traditional propagation, i.e. seedage, graft-ing and
budding. Marcottage (air layering) methods have been tried for many years (Rajan
and Ram, 1988; Majumder and Sharma, 1990) and although recently improved
(Núñez-Elisea et al., 1989, 1991, 1992; Lambe et al., 1991; Núñez-Elisea and
Davenport, 1995), it is still not used commercially. In vitro techniques hold great
promise for rootstock propagation and cultivar improvement, and are currently
being perfected and investigated (Litz, 1984, 1986, 2005; Litz et al., 1984; see Litz
et al., Chapter 18, this volume).
The monoembryonic Florida cultivars in Brazil are vegetatively propa-gated
by grafting and budding on polyembryonic rootstocks. Rootstock and
444 J.H. Crane et al.

grafted mangoes are established in nurseries by sowing seeds in a soil sub-strate


consisting of a mixture of organic matter and semi-sterilized soil plus 3 kg of
superphosphate (3Ca(H2PO4)2) and 0.5 kg of potassium nitrate (KNO 3) for each
m3 of mixture (Pinto et al., 1981; Castro Neto et al., 2002). In the south-east, the
indirect seeding technique is used: seed is sown in the soil to germi-nate and then
transplanted to small softwood containers or ‘jacás’. Subsequently, seedlings are
patch budded. In the central and north-east regions, seedlings are grown in
substrate in plastic bags and subsequently whip or veneer grafted. The endocarp is
removed prior to sowing to accelerate germination and improve uniformity of
rootstock growth (Pinto and Genú, 1981).
The most important characteristics for mango rootstocks are vigour, high yield
potential, compatibility with the scion, environmental adaptability and resistance to
pests and diseases (Rosetto et al., 1996; Nascimento et al., 2002). Polyembryonic
cultivars, that is ‘Espada’ and ‘Fiapo’ in the north-east and ‘Rosinha’ and
‘Coquinho’ in the south-east, are the most important root-stocks. The Agencia
Paulista de Tecnologia Agropecuária (APTA) has developed root-rot-resistant
rootstocks, including IAC-101, IAC-102 and IAC-106; however, they are not
widely used.
There are no specific mango rootstocks in Mexico. Before the mid-20th
century, seedling plants were used; however, uniform plantings did exist because
polyembryonic ‘Manila’ was grown in the Gulf of Mexico region. Elsewhere, there
were monoembryonic trees, resulting in a heterogeneous mix of trees. Since the
late 1950s and early 1960s, polyembryonic criollo root-stocks have been used for
rootstocks; however, due to their vigour, mono-embryonic rootstocks are still in
use in some areas. Commercial production is done in partially shaded nurseries. A
seedbed is prepared using sterilized substrate (loamy soil, composted organic
matter or a mixture). The fibrous seedcoat is removed to expose the smooth, bean-
shaped seed, which is soaked for about 10–30 min in 50 mg/l gibberellic acid
(GA3) to reduce germination time. Seeds are placed on the seedbed (usually in
May–July) with their hump up and watered every other day. After 3–5 weeks,
vigorous and healthy seed-lings are transferred to black plastic bags 20 cm wide ×
40 cm long filled with a sterilized mixture of sandy loam soil and organic matter
(3:1). Rootstocks are veneer grafted when they are 4–6 months old. Grafted plants
are ready for planting by May–July of the next year, during the rainy summer
season.
Some experienced mango growers initially plant seedling trees in the orchard
to ensure a vigorous tree. Grafting is then performed 3–6 months after planting. In
some cases a low vigour interstock like ‘Esmeralda’ is top-worked onto the
seedling rootstock and then grafted to the desired scion. The use of ‘Esmeralda’
interstocks has reduced tree canopy size by 28% and 33% for ‘Manila’ and
‘Ataulfo’, respectively (Mosqueda-Vázquez et al., 1996; Vázquez-Valdivia et al.,
2000).
Mangoes in Taiwan are generally splice or veneer grafted; however, a
combination of the two methods is used when the rootstock and scion are of
different diameters. Polyembryonic ‘Tsar-swain’ seedlings are pre-ferred as clonal
rootstocks although seedlings of monoembryonic ‘Irwin’ are sometimes used.
Crop Production: Management 445

The main commercial cultivars in the USA, ‘Keitt’, ‘Tommy Atkins’, ‘Van
Dyke’, ‘Haden’ and ‘Brooks’, are monoembryonic. Orchards are planted with
grafted or budded trees (Aponte-Morán et al., 1977; Chia et al., 1988; Crane and
Campbell, 1991; Hamilton et al., 1992). The most common rootstocks in Florida
are seedlings of polyembryonic ‘Turpentine’, ‘Number 11’ and ‘Criollo’. In
Hawaii, any mango seedlings are used. In California, ‘Turpen-tine’ or ‘Criollo’ are
used. In Puerto Rico, polyembryonic ‘Mangotino’ and ‘Pasote’ seedlings are used
in the Ponce region and ‘Mayaguezano’, ‘Turpen-tine’ and ‘Colombo Kidney’ in
the Mayaguez region (Aponte-Morán et al., 1977; Toro, 1988; E.E. Toro, personal
communication). Regardless of the seed source, roguing the zygotic seedlings is
important for obtaining uniform tree size, growth characteristics and production
(Schnell and Knight, 1991, 1992; Degani et al., 1993; Schnell et al., 1994). Seed is
removed from mature fruit along with the husk, and the seed is planted in well-
drained container media (Sauls and Campbell, 1980; Young and Sauls, 1989).
Seedlings are large enough to graft or bud after 2–6 months. The most common
vegetative prop-agation methods in Florida are veneer and cleft grafting (Sauls and
Camp-bell, 1980; Crane and Campbell, 1991) and chip and ‘T’ or inverted ‘T’
budding. The trees are usually ready for field planting in 6–12 months. Graft-ing
may be done at any time of year when suitable rootstocks are available, but it is
more successful during warm weather. In Puerto Rico the most common
propagation method is cleft grafting (Aponte-Morán et al., 1977; Toro, 1988).
Grafting is most successful during the spring and from September through to
November.

Topworking of established trees is common. Trees are cut back to several


major limbs or the trunk. In Florida, several sprouts emerging from the pruned
limbs are allowed to develop to 1.3–7.6 cm diameter, and are veneer grafted with
the new cultivar (Sauls and Campbell, 1980). In Puerto Rico, several sprouts are
allowed to develop, and larger sprouts (2.5–7.6 cm diam-eter) are cleft grafted and
smaller diameter sprouts (1.0–1.5 cm) are bark grafted (Toro, 1988).

13.6 Major Cultivars


For detailed information about cultivars, refer to Knight et al., Chapter 3, this
volume.
Mango seeds were introduced by Portuguese and Spanish colonizers to Brazil
during the 16th century. Subsequently, mango seeds were dispersed into the
interior of the country. There are >120 local mango cultivars, many of which are
the result of open pollinations among Indian and Philippine mango germplasm.
There is a large amount of variability among the cultivars with respect to colour,
shape and size of the fruit. Most of these cultivars are for the fresh fruit market
(e.g. ‘Espada’, ‘Bourbon’, ‘Ubá’, ‘Rosa’, ‘Coité’, ‘Coquinho’ and ‘Lira’), or for
juice, jellies and other value-added products (e.g. ‘Ubá’, ‘Coité’, ‘Mamão’) (Table
13.3). Some polyembryonic criollo culti-vars (e.g. ‘Espada’, ‘Coquinho’ ‘Comum’
and ‘Fiapo’) are used as rootstocks.
446
Table 13.3. Major commercial mango cultivars in Brazil, Mexico, Taiwan and the USA.

a b
Country Cultivar Origin Seed type Production season Tree growth habit

Brazil ‘Espada’ Brazil Polyembryonic Oct.–Dec.


‘Bourbon’ Brazil Monoembryonic Oct.–Dec.
‘Ubá’ Brazil Monoembryonic Oct.–Dec.
‘Rosa’ Brazil Polyembryonic Oct.–Dec.
‘Coité’ Brazil Monoembryonic Oct.–Dec.
‘Mamão’ Brazil Monoembryonic Oct.–Dec.
‘Coquinho’ Brazil Polyembryonic Oct.–Dec.
‘Van Dyke’ USA Monoembryonic Nov.–Jan. Large/spreading/open
‘Tommy Atkins’ USA Monoembryonic Nov.–Jan. Large/dense
‘Haden’ USA Monoembryonic Nov.–Jan. Large/spreading
‘Keitt’ USA Monoembryonic Feb.–Mar. Small/spreading/open

J.H. Crane et al.


‘Palmer’ USA Monoembryonic Jan.–Feb. Small/spreading
Mexico ‘Manila’ Mexico Polyembryonic Jan.–Aug. Large/dense
‘Tommy Atkins’ USA Monoembryonic Feb.–Aug. Large/dense
‘Haden’ USA Monoembryonic May–July Large/spreading
‘Kent’ USA Monoembryonic Feb.–Aug. Large/dense
‘Keitt’ USA Monoembryonic May–Sep. Medium/spreading/open
‘Ataulfo’ Mexico Polyembryonic Jan.–June Large/spreading/upright
Taiwan ‘Tsar-swain’ Taiwan Polyembryonic Mar.–July
‘Jin-hwung’ Taiwan Monoembryonic June–Sept.
‘Tainong No. 1’ Taiwan Monoembryonic May–June
‘Irwin’ USA Monoembryonic April–Aug.
‘Keitt’ USA Monoembryonic Aug.–Oct. Small/spreading
USA ‘Tommy Atkins’ USA Monoembryonic June–Aug. Large/dense
‘Keitt’ USA Monoembryonic Aug.–Oct. Large/spreading
‘Kent’ USA Monoembryonic July–Aug. Large/upright/dense
‘Van Dyke’ USA Monoembryonic July–Aug. Large/spreading/open
‘Parvin’ USA Monoembryonic July–Aug. Large/spreading/dense
‘Haden’ USA Monoembryonic June–mid-July Large/spreading
‘Brooks’ USA Monoembryonic Aug.–Oct. Small/open
a
Natural fruit production season for country in which cultivar is being
grown. b Growth habit for country in which cultivar is being grown.
Crop Production: Management 447

Florida selections (i.e. ‘Haden’, ‘Keitt’, ‘Tommy Atkins’, ‘Van Dyke’ and
‘Palmer’) are grown for the export trade. Most Florida cultivars produce from
Decem-ber to January except ‘Palmer’, which is harvested between January and
Feb-ruary and ‘Keitt’, which is harvested between February and March. ‘Tommy
Atkins’ represents 80% of the commercial export volume in Brazil (Pinto, 2004);
‘Palmer’ is increasing in demand in the national market.
Mango production in Mexico relies on the following cultivars (from early to
late beginning of harvest season): ‘Manila’, ‘Ataulfo’, ‘Kent’, ‘Haden’, ‘Tommy
Atkins’ and ‘Keitt’ (Table 13.3). ‘Zill’, ‘Irwin’, ‘Sensation’, ‘Diplomático’,
‘Manililla’, ‘Oro’, ‘Ovo’ and criollos are also grown. Mango harvest seasons are
no longer inflexible as they may be modified by pruning, water manage-ment and
growth regulators. The most widely planted cultivar in Mexico is ‘Manila’ (45,396
ha); no export figures have been reported. Although ‘Manila’ is grown all over the
country it dominates in Veracruz (24,908 ha) and Guerrero (9198 ha). It is an
alternate bearer and the harvest season is from May to June in Veracruz and July in
Sinaloa (De los Santos and Mosqueda-Vázquez, 1988–89; C. Guzmán, personal
communication). There are 22,890 ha of ‘Tommy Atkins’ and it is the major export
cultivar to the USA (33% of total exports). Fruit is harvested from February
(Michoacán) to August (Colima, Jalisco and Sinaloa) (V. Medina and C. Guzmán,
personal commu-nications). ‘Kent’ is cultivated on 13,366 ha and accounts for
23% of exports to the USA. Sinaloa is the major producer of ‘Kent’ (9710 ha) and
the begin-ning of harvest season is similar to ‘Tommy Atkins’ (Campbell, 1992);
however, it can be harvested as late as September in Colima, Jalisco and Nayarit.

‘Ataulfo’ is from Chiapas and is now cultivated nationwide on >34,000 ha.


The major producing states are Chiapas (south) and Nayarit (north) with 18,334 ha
and 7156 ha, respectively. The harvest season is February–May in Chiapas and
June–July in Sinaloa (V. Palacio and C. Guzmán, personal com-munications). In
some years this cultivar is the third most important export to the USA, and
accounted for 19% of exports in 2006 (EMEX, 2007). ‘Keitt’ (6828 ha) is the
fourth most important export cultivar (14% exports). Most ‘Keitt’ orchards are
located in Sinaloa (5198 ha). ‘Keitt’ is harvested from May (Nayarit) to September
(Colima, Jalisco and Sinaloa). Large fruit size and fungal lesions sometimes cause
marketing problems because harvest occurs during the rainy season. ‘Haden’
(27,768 ha) is still an important mango although exports to the USA have
decreased to 11% (EMEX, 2007); the major producing states are Michoacán
(15,573 ha), Guerrero (5053 ha) and Sinaloa (3518 ha). Fruit are harvested from
February/March (Michoacán and Guerrero) to August/September in Colima,
Jalisco and Michoacán. Alternate bearing and mango malformation are problems
and ‘Haden’ orchards are being replaced by or topworked to cultivars like
‘Ataulfo’.
The mango was introduced into Taiwan from South-east Asia by the Dutch in
the 16th century (Chen, 1991). The introductions were all polyem-bryonic and
included ‘Tsar-swain’, ‘Pung-swain’, ‘Va-swain’ and ‘Kee-gway-swain’. ‘Tsar-
swain’ is still very important (Table 13.3). From 1954 to 1973, 35 monoembryonic
cultivars were introduced from the USA, and ‘Irwin’ and ‘Keitt’ have been major
cultivars in Taiwan since 1960. ‘Jin-hwung’, which is
448 J.H. Crane et al.

derived from a chance seedling of ‘White’ and ‘Keitt’, was selected in 1980.
‘Tainong No. 1’ and ‘Tainong No. 2’ are derived from controlled pollinations and
were released in 1985 by the Fengshan Tropical Horticultural Experiment Station,
Taiwan Agricultural Research Institute. Only ‘Tsar-swain’, ‘Jin-hwung’, ‘Irwin’
and ‘Keitt’ and ‘Tainong No. 1’ are commercially important today.
Fruit in the southern prefectures or in lower elevation orchards are har-vested
earlier than the northern prefectures or orchards at higher elevations. ‘Tsar-swain’
comprises about 40% of total production; its season is from March to August,
depending on the prefecture or location of the orchard. The fruit of ‘Tsar-swain’ is
154 g, 100 mm long and 64 mm wide, total soluble solids (TSS) value is 17 Brix,
total titratable acidity 2.2% and seed/pulp weight ratio 0.84 g. ‘Irwin’ comprises
40% of the production and is harvested from Pingtung in April and in Tainan in
August. ‘Jin-hwung’ comprises about 10% of production and is harvested from
June to September. The fruit of ‘Jin-hwung’ is 965 g, 144 mm long and 99 mm
wide, TSS value is 15 Brix, total titrateable acidity 2.3% and seed/pulp weight
ratio 0.94 g. ‘Keitt’ makes up c.5% of production and is harvested from August to
October. The fruit of ‘Tainong No. 1’ is 237 g, 99 mm long and 69 mm wide, TSS
value is 20 Brix, total titrateable acidity 4.0% and seed/pulp weight ratio 0.85 g.

The major cultivars in Florida are ‘Keitt’ and ‘Tommy Atkins’, which account
for c.70% and 20% of the hectarage, respectively (Table 13.3). Small commercial
hectarages of ‘Van Dyke’, ‘Palmer’, ‘Irwin’, ‘Raposa’ and ‘Kent’ are also grown.
In Puerto Rico, the major export cultivars are ‘Keitt’, which makes up c.60% of the
hectarage, ‘Parvin’ (20%), ‘Irwin’ (10%), ‘Tommy Atkins’ (5%) and ‘Haden’ (<
5%). Other cultivars (i.e. ‘Davis-Haden’, ‘Palmer’, ‘Kent’, ‘Mayaguezano’, ‘Poste’
and ‘Cubano’) are grown on a small scale. The local cultivars are grown for the
domestic market (Toro, 1988). The commer-cial hectarage of California is mostly
‘Keitt’, although other cultivars have been evaluated (Scott, 1990; Linden, 2006).

Immature ‘Keitt’ is the major early season cultivar (picked green) and mature
‘Tommy Atkins’ is the major early season cultivar in Florida (Table 13.3) (J.H.
Crane, personal communication). Mature ‘Keitt’ and ‘Kent’ mangoes are the major
late season cultivars. Six- to 8-year-old ‘Tommy Atkins’ trees produce 75–150
kg/tree and older trees may produce up to 300 kg/tree. Internal breakdown varies
from year to year and may be aggravated by over-fertilization with nitrogen (N).
Fruit are considered moderately resistant to anthracnose. The harvest season is June
through to August. ‘Keitt’ trees are precocious and produce large crops regularly
during July through to September.

‘Palmer’, ‘Irwin’ and ‘Van Dyke’ are grown commercially on a limited scale
in Florida and Puerto Rico (except ‘Van Dyke’). In Florida, ‘Palmer’ is harvested
from July to early September, ‘Irwin’ from June to early July, and ‘Van Dyke’
from July to August. In the recent past, ‘Bailey’s Marvel’, ‘Brooks’ and ‘Haden’
were grown commercially; however, their importance has declined due to natural
disasters and susceptibility to anthracnose. ‘Haden’ is no longer grown
commercially in Florida (Crane and Campbell, 1991; Campbell, 1992) but,
‘Glenn’, a seedling of ‘Haden’, has been recommended
Crop Production: Management 449

(Campbell and Campbell, 1996). ‘Harders’ is grown in Manoa, Oahu Island,


Hawaii (Hamilton et al., 1992; Hamilton, 1993). Trees bear regularly and often
produce off-season good-quality fruit in the late autumn and winter. ‘Rapoza’, a
seedling of ‘Irwin’, was selected in Hawaii in 1984 (Hamilton et al., 1992); its
harvest season is from mid-July to October.

13.7 Plant Spacing


Plant spacing and density are influenced by climate, soil type and depth, rootstock
and scion vigour, growth habit and the targeted tree size. Cultural practices
including tree-size control, fertilizer and irrigation availability, methods, rates,
timing and frequencies; current technology and the necessity for orchard access by
farm machinery also influence plant spacing and con-figuration. The cost of
borrowed capital, land, water, irrigation, orchard maintenance and net returns will
dictate what cultivars are grown and how they are managed (see Evans and
Mendoza, Chapter 16, this volume).
Initial plant spacing and density should be planned to maximize early yield
and returns from young orchards and to maintain high yields from mature orchards.
Trees in overcrowded orchards compete for water, nutri-ents and light and
eventually lose production in the lower canopy. The effi-cacy of foliar sprays
(nutrients, pesticides and growth regulators) is reduced and harvest is more difficult
in crowded orchards.
Plant spacing among trees and rows has decreased in recent years to optimize
returns on investments in land, equipment and orchard infrastruc-ture. This has
been possible because of more insight into the physiology of mango trees,
improvements in irrigation system design and efficiency, and better fertilizer
delivery systems and pruning (i.e. intense hand and/or mechanical pruning) and the
use of plant growth regulators. Closer plant spacings require more tree-size control
and expertise on the part of the pro-ducer. Various training systems are advocated
for new trees to force complex branching and increase bearing surface volume and
fruit production poten-tial (Fig. 13.2). Topping and hedging and/or hand pruning
(selective prun-ing and/or heading back) is used to maintain mature tree size and
fruit production and improve fruit colour (Fig. 13.3).

In Brazil, the traditional spacing of 10 m × 10 m in a rectangular or qua-dratic


format, with a density of 100 plants/ha, has been replaced by plant-ings of 8 m
(between rows) × 5 m (within rows) to 5 m × 5 m with higher plant densities,
which vary from 250 to 400 plants/ha (Cunha and Castro Neto, 2000; Mouco et al.,
2002). In general, growers use two types of pruning: for-mation and production
pruning (Albuquerque et al., 2002). Formation prun-ing is used to establish the
intial tree architecture and trees are pruned five times for several years leaving
c.243 branches prior to starting mango pro-duction (Fig. 13.2). Pruning systems
include cleaning, skirting of the lower canopy, lateral, central and top-canopy
pruning, and pruning to correct poor canopy development and maintain adequate
canopy after production commences (Albuquerque et al., 2002).
450 J.H. Crane et al.

Planted Headed First Tip prune After each flush


single to ~80 cm branching shoots 10 mm repeat tip pruning
stem diameter of shoots 10 mm in
diameter once they
have matured

Fig. 13.2. Tree training system for young trees to increase branching and productive
canopy volume (Source: after Oosthuyse, 1995; Campbell and Wasielewski, 2000).

Central leader Central leader topped Closed vase

Rectangle Rectangle with roof Open vase

Fig. 13.3. Various mechanical topping and hedging schemes that may include
selective hand pruning or shoot tipping to maintain tree size and regular bearing.

In most of Brazil, holes for plantings are 60 × 60 × 60 cm and simple or triple


superphosphate, lime and manure are usually mixed with the exca-vated soil c.20
days prior to planting (Pinto and Ramos, 1998; Albuquerque et al., 1999). In the
north-east, planting is during the rainy season and new plantings are mulched to
reduce soil evaporation where solar irradiation and temperature are very intense,
which can lead to death of newly planted mango trees. Intercropping with crops
such as beans, maize, papaya and pineapple improves incomes during the first 3
years of orchard establishment
Crop Production: Management 451

(Mouco et al., 2002). Windbreaks, consisting of pine trees, elephant grass and three
rows of banana plants, are often used during the first 2 years of orchard
establishment.
In Mexico, orchards were originally established at 10 m × 10 m to 16 m × 16
m, and trees were not pruned, which resulted in enormous and very pro-ductive
trees; however, care and harvest from very large trees is problematic. Plant spacing
is based on cultivar vigour (‘Ataulfo’ and ‘Manila’ are most vigorous), land slope
(wider spaces as slope increases), farm machinery, cli-matic conditions and soil
fertility (wider distances for warmer climates and more fertile soils) and water
availability (Cruzaley-Sarabia et al., 2006). Irri-gated orchards may handle closer
spacings if pruning is practised. Under rainfed conditions, soil moisture availability
may have an important impact on tree size. For example, 10-year-old orchards may
have a few big trees (c.69–100 trees/ha) or if trees are spaced more closely (300–
600 trees/ha), tree size is reduced.

Currently, both square and rectangular planting patterns are common in


Mexico although hexagonal planting systems are also used. For square designs,
tree spacing ranges from 5 m (400 trees/ha) to 10 m (100 trees/ha). Rectangular
planting systems have more options and the most common arrangements are 6 m ×
5 m (333 trees/ha), 8 m × 5 m (250 trees/ha), 8 m × 6 m (205 trees/ha) and 10 m ×
5 m (200 trees/ha) (Chávez-Contreras et al., 2001).
In Taiwan, plant spacing for monoembryonic cultivars (i.e. ‘Tsar-swain’)
ranges from 6 m × 6 m to 10 m × 10 m (100–256 trees/ha). In contrast, plant
spacing for polyembryonic cultivars ranges from 4 m × 5 m to 5 m × 6 m,
depending on the topography and soil fertility. Orchards on sloped lands and
infertile soils are less vigorous and are planted at higher densities than orchards on
lowlands and fertile soils.
Recommended plant spacings in Hawaii and Florida are similar. The cooler
climate of south-eastern California allows close spacing, i.e. 3 m in-row by 5.4 m
between-rows (617 trees/ha). Traditional plant spacings in Florida were as high as
11 m in-rows and 11–14 m between-rows (64–82 trees/ha) (Young and Sauls,
1989). Plant spacings in Florida currently include 3.5 m × 6.1 m (468 trees/ha), 4.5
m × 6.1 m (364 trees/ha), 4.5 m × 7.6 m (292 trees/ha), 6.1 m × 6.1 m (268
trees/ha), 6.1 m × 7.6 m (215 trees/ha) and 7.6 m × 7.6 m (173 trees/ha) (Young
and Sauls, 1989; Crane and Campbell, 1991). In Puerto Rico, plant spacings are 3.7
m × 5.5 m (491 trees/ha), 4.6 m × 9.1 m (238 trees/ha), 6.1 m × 9.1 m (180
trees/ha), 7.6 m × 7.6 m (173 trees/ha) and 9.1 m × 9.1 m (120 trees/ha) (Toro,
1988). Interplanting mango trees at moder-ate to wide plant spacings (i.e. 5.4–7.5
m in-row) with banana or plantain (Musa sp.), and papaya (Carica papaya L.) in
Florida and plantains in Puerto Rico is widely practised. Mango trees have also
been planted at close spacing (e.g. 3.0–4.5 m) and every other tree is removed if
crowding becomes a prob-lem. Controlling tree size and maintaining crop
productivity is important, otherwise, competition among trees will reduce yields
and fruit quality (Toro, 1988; Crane and Campbell, 1991; Oosthuyse, 1995).
Annual or biennial hand pruning and/or mechanical hedging and topping is
necessary and should begin several years before trees begin to crowd and should
continue after
452 J.H. Crane et al.

trees grow to their desired size based on plant spacing. Mature trees are topped at
3.5–5 m, and hedged trees are cut on a slight angle (5–10) to leave a 2.5–3.5 m
row middle (J.H. Crane, personal communication). Trees can be maintained at in-
row spacings of 2–3 m; however, this involves intense tree training and hand
pruning, which most producers have been unwilling to adopt (Fig. 13.3) (Oothuyse,
1995; Oosthuyse and Jacobs, 1995; Stassen et al., 1999; Campbell and
Wasielewski, 2000). Severe hedging is utilized to increase light penetration and re-
establish inner productive canopy but this can result in little to no production in the
following year. Some producers utilize a com-bination of mechanical pruning
followed by selective pruning to open the inner canopy to light and air movement,
improving fruit colour and reducing disease pressure.

13.8 Fertilizer Practices


Fertilizer practices are influenced by availability and cost of organic and inorganic
materials, application costs, soil type and depth, irrigation prac-tices and rainfall,
cultivar and production objectives. The response to fertil-izer is influenced by
fertilizer source, rate, timing and method of application, tree-growth stage, climate,
edaphic conditions, soil moisture status and cul-tivar vigour. The objective of
fertilizer practices for young trees during years 1 and 2 is to establish the tree and
maintain constant canopy growth and health while building a structurally strong
tree. Objectives for bearing trees include maintenance of tree health, avoidance of
alternate bearing and allow-ance for vegetative dormancy which results in flower
induction and initiation.
Over-fertilization of bearing trees leads to continuous vegetative growth under
some climatic conditions, reduced flowering and fruit yields and increased
occurrence of physiological disorders of the fruit (Schaffer et al., 1994; Coelho et
al., 2002; Nguyen et al., 2004). Likewise, improper fertilization may lead to
nutrient deficiencies and/or toxicities which result in reduce tree growth, yields and
fruit quality. Fertilizer practices will continue to be refined and developed as our
understanding of the physiological needs and responses to essential plant nutrients
increases and as concerns increase about the effect of excess and/or leached
nutrients on the natural environment.
Fertilizer management is based on soil and leaf analysis in Brazil (Tables
13.4–13.6; Santos et al., 2002). The quantity and type of fertilizer application
varies from the orchard establishment period (mostly vegetative plant growth) to
the orchard production period. At the time of planting, 20–30 l/ hole of cattle
manure is usually mixed with the native soil. Subsequently, N, phosphorus (P) and
potassium (K) are applied 2–3 times annually. During young tree establishment, P
and K are recommended only if these elements are deficient (Tables 13.5 and
13.6). These fertilizers are broadcast mechanically in the planting rows, and then
incorporated into the top soil layer (Andrade, 2004; Sousa et al., 2004). The
quantities of macronutrients applied at the time of planting and at various growth
stages in south-eastern Brazil (mainly São Paulo) and the central regions are based
on plant age and soil P and K content
Crop Production: Management 453

Table 13.4. Standard leaf nutrient content ranges for mature mango trees in Brazil,
Mexico, Taiwan and Florida USA.

a
Range for mature trees
b c d
Mineral Unit Brazil Mexico Taiwan USA
N % 1.2–1.4 1.2–1.5 1.4–1.7 1.0–1.5
P % 0.08–0.16 0.07–0.13 0.1–0.15 0.09–0.18
K % 0.5–1.6 0.6–0.7 0.9–1.2 0.5–1.0
Ca % 2.0–3.5 2.3–3.4 1.0–1.8 3.0–5.0
Mg % 0.25–0.5 0.14–0.19 0.2–0.35 0.15–0.47
e
B ppm 50–100 NR NR 24–54
Fe ppm 50–200 97–114 60–120 38–120
Mn ppm 50–100 191–802 30–200 92–182
Zn ppm 20–40 14–20 20–100 101–119
Cu ppm 10–50 5–12 5–20 28–35
a
Recommended leaf nutrient levels based on research and the literature.
b
Source of data for Mexico: Guzmán-Estrada (2001, 2004, 2006). Range shown for all
cultivars tested (see Table 13.8 for detail by cultivar).
c
Values are for ‘Irwin’ leaves. Source of data for Taiwan: Job (1989).
d
Source of data for USA: Young and Koo (1969, 1971); Young and Sauls
(1989). e NR, not reported.

Table 13.5. Soil phosphate and potash corrective fertilization rates based on soil
analysis in Brazil.
(a) Rate of P2O5 (kg/ha) application.

Level of P availability in the soil

Clay content (%) Low Medium Adequate

15 60 30 0
16–35 100 50 0
36–60 200 100 0
>60 280 140 0

(b) Rate of K2O (kg/ha) application.

Soil K content Level of K availability in the soil


3
(mg/dm ) Low Medium Adequate
3
Cation exchange capacity = <4.0 cmol/dm at pH 7 and <20% clay
<15 50 – –
16–40 – 25 –
>40 – – 0
3
Cation exchange capacity = >4.0 cmol/dm at pH 7 and >20% clay
<25 100 – –
25–80 – 50 –
>80 – – 0
454 J.H. Crane et al.

Table 13.6. Quantity of P2O5 and K2O applications based on young tree plant age, and
P and K soil content of oxisoils for São Paulo and the Brazilian central regions.

3 3
P soil content (mg/dm ) Exchangeable K soil content (mmol/dm )
Trace <6 6–12 13–30 >30 <0.8 0.8–1.5 1.6–3.0 >3.0
Tree age
(years) Rate of P2O5 application (g/plant) Rate of K2O application (g/plant)

0–1 30 0 0 0 0 40 0 0 0
1–2 60 160 120 80 0 80 40 0 0
2–3 120 240 160 100 0 160 120 80 40
3–4 160 320 240 120 0 240 180 120 80

(Table 13.5); however, the quantities of N, P and K for mango production are
based mainly on soil and leaf analysis. Leaves used for analyses are 6–8 months
old, from the mid-canopy and from branches with fruits and from all four sides of
the canopy to reduce variation in analysis results. The proce-dure for sampling
leaves is: (i) divide the orchard into separate areas of no more than 10 ha with trees
of the same age and productivity and growing on similar soil; (ii) collect healthy
leaves from the middle of the tree canopy, from the four cardinal points on normal
branches with recently matured leaves from the previous flush of growth, with
leaves not less than 4 months old. Remove four leaves per plant, from a total of 20
plants selected ran-domly, and take the leaves prior to the application of nitrates or
other foliar fertilizers that are applied to break the dormancy of the floral buds.

There are two distinct periods of mango fertilization in Brazil: pre- and
postharvest fertilizations (Alves et al., 2002). In the preharvest fertilization of non-
irrigated orchards, P should be applied in a single dose, before flower-ing, and
incorporated with a medium-weight plough. At the beginning of the rainy period
40% of the N and K should be applied and the remainder after flowering, during
fruit development. In irrigated orchards c.40% of the P should be applied before
flowering and 60% postharvest. For N, 50% is applied preharvest (i.e. after the start
of fruit set) and 50% postharvest. Potas-sium applications should be distributed
throughout the production cycle but more during fruit development and c.25%
postharvest. In São Paulo and cen-tral Brazil c.40% and 20% of the N and K should
be applied after harvest and at the end of the rainy season (i.e. beginning of March),
respectively.
When the productivity of orchards in north-east Brazil is <10 t/ha and leaf N
concentrations exceed 16 g/kg and the P and K concentrations in the soil profile are
>40 mg/dm3 and 0.45 cmol/dm3, respectively, application of N, P and K is
unnecessary (Table 13.7). On the other hand, if the expected productivity is >50
t/ha and leaf N concentrations are <12 g/kg and the P and K concentrations in the
soil profile are <10 mg/dm3 and <0.16 cmol/ dm3, respectively, 120 kg/ha of N,
150 kg/ha of phosphate (P2O5) and 250 kg/ ha of K2O should be applied.
Table 13.7. Quantity of N, P2O5 and K2O applications based on fruit productivity, leaf N content, and P and K soil content for the semi-arid

Crop Production: Management


regions of Brazil.
3 -3
N leaf content (g/kg) Soil P content (mg/dm ) K soil content (mmol c dm )
Fruit <12 12–14 14–16 >16 <10 10–20 21–40 >40 <1.6 1.6–3.0 3.0–4.5 >4.5
production
(t/ha) N application rate (kg/ha) P2O5 application rate (kg/ha) K2O application rate (kg/ha)

15–20 60 40 20 0 45 30 15 0 80 40 20 0
20–30 75 50 25 0 65 45 20 0 120 60 30 0
30–40 90 60 30 0 85 60 30 0 160 80 45 0

455
456 J.H. Crane et al.

Table 13.8. Standard leaf nutrient content values for selected mango cultivars in Mexico
(Source: Guzmán-Estrada, 2001, 2004, 2006).

Cultivar

Mineral Unit Ataulfo Haden Keitt Kent Manila Tommy Atkins

N % 1.31 1.20 1.28 1.16 1.46 1.28


P % 0.07 0.09 0.15 0.08 0.09 0.13
K % 0.60 0.59 0.56 0.57 0.66 0.62
Ca % 2.53 3.40 2.87 3.23 2.31 2.77
Mg % 0.14 0.16 0.19 0.19 0.18 0.15
S % 0.23 0.17 0.20 0.15 0.15 0.17
a
B ppm NR NR NR NR NR NR
Fe ppm 113.6 101.3 99.4 96.6 112.7 96.2
Mn ppm 801.9 328.1 191.1 243.8 236.3 219.3
Zn ppm 20.2 16.6 21.4 13.6 17.7 16.4
Cu ppm 5.40 5.43 11.70 6.70 6.83 6.96
a
NR, not reported.

Deficiencies of zinc (Zn) and boron (B) are common in orchards in Brazil.
Zinc sulfate and borax are normally used to correct these deficiencies; the rates
depend upon leaf analyses (Silva et al., 2002). Lime is applied when base
saturation is <60%. Gypsum at a rate of 2 t/ha for sand and 3 t/ha for clay soils is
recommended to reduce the incidence of internal breakdown (Silva et al., 2002).
Gypsum (280 g/m2) incorporated at a 30 cm soil depth before orchard
establishment under Cerrados conditions (pH 3.7 and very poor Ca levels) reduces
internal breakdown from 60 to 3% in ‘Tommy Atkins’ (Pinto et al., 1994).

Fertilization is not a common practice in most mango-producing regions of


Mexico. In Colima (Central Pacific region) <30% of orchards are fertilized.
However, in the Soconusco area of Chiapas (Southern Pacific region), 86% of
‘Ataulfo’ orchards are fertilized; 56.7% orchards are fertilized once a year, 39.5%
twice a year, and 2.7% receive three applications per year. In fertilized orchards,
there is a high variation in amounts, materials, timing and forms of fertilizer
applications. Recommendations are empirical since there are few published
guidelines. Standard leaf nutrient levels have been proposed for several cultivars
grown in Sinaloa (Table 13.4, Table 13.8). Only a few growers use leaf or soil
analyses. Fertigation is rarely employed. In the Soconusco area of Chiapas,
‘Ataulfo’ mango trees have K, Ca, Mg, Zn and B deficiencies. In southern Sinaloa,
most mango cultivars (‘Ataulfo’, ‘Haden’, ‘Tommy Atkins’, ‘Kent’, ‘Keitt’) are
deficient in N, P, K, Mg, S, copper (Cu), Zn and B, normal in Ca and iron (Fe) and
high in manganese (Mn) (Guzmán-Estrada, 2001). In Colima, where mango is
cultivated on sandy soils with pH 7–8.4, Fe and Zn are the most common
deficiencies. In Nayarit, irrigated and non-irrigated ‘Haden’ and ‘Tommy Atkins’
trees are deficient in K, P and Ca (in this order) and have excess Mg (Salazar-
García et al., 1993). No Al toxicity has been reported.
Crop Production: Management 457

Except for N, visual symptoms of nutrient deficiencies are uncommon in


commercial mango orchards; however, fertilized trees have significantly increased
yield and fruit size. Nitrogen is most commonly applied, followed by P and K.
Current fertilization practices vary with the mango-producing region, soil type,
cultivar and tree age. The NPK recommendations for ‘Manila’ trees at 1–4 years,
5–10 years, 10–15 years, 16–20 years and >20 years old in the Gulf of Mexico
region are: 0.2-0.1-0.1 kg/tree/year, 0.4-0.2-0.4 kg/ tree/year, 0.6-0.3-0.6
kg/tree/year, 0.8-0.4-0.8 kg/tree/year and 1.0-0.5-1.0 kg/tree/year, respectively.

Young trees in the Southern Pacific region receive NPK at 0.4-0.2-0.2 kg/
tree/year, respectively, from year 1 to year 5, and at 0.7-0.7-0.7 kg/tree/year
thereafter. In Michoacán (Central Pacific region), mature ‘Haden’ and ‘Tommy
Atkins’ trees receive NPK at 1.1-0.4-0.9 kg/tree/year (Chávez-Contreras et al.,
2001). The NPK recommendation in the Northern Pacific region at 1–4 years, 5–10
years, and 10–15 years of age is 0.4-0.2-0.2 kg/tree/year, 1.3-0.55-0.85 kg/tree/year
and 2.8-0.9-1.8 kg/tree/year, respectively. Although soil amendments are needed in
regions with low or high pH soils (i.e. Chiapas and Colima) they are not used.

Micronutrients are not usually included in fertilization programmes; however,


circumstantial evidence suggests there is an interaction between flower B
deficiency and extreme high temperature as a major factor causing stenocarpic
fruit. Foliar B applications at either pre- or during full bloom have become a
routine practice for ‘Ataulfo’ orchards. Zinc, Mn and Cu, are included with
fungicide sprays during bloom and the rainy season.
General fertilizer recommendations for mangoes of different ages vary in
Taiwan (Table 13.9). Fertilizers are applied in two equal rates: (i) after har-vest;
and (ii) when fruit are at the marble stage of development. Fertilizer is applied by
broadcasting, banding, side dressing and hole-application. Leaf analysis is used for
nutrient diagnosis and fertilizer recommendations (Table 13.4). The rates of N, P
and K increase with tree age (Table 13.9). Dolomite is commonly used to adjust
soil acidity and as a Mg supplement. The applica-tion rates are 1 t/ha/year for
sandy soils, 1.5 t/ha/year for loamy or silty soils and 2 t/ha/year for clay soils.
Sulfur is applied as ammonium sulfate ((NH 4)2SO4), magnesium sulfate (MgSO4)
or calcium sulfate (CaSO4). Boron is applied at a rate of 50 g/tree or as a 0.3 %
spray 2–3 times a year. No addi-tional micronutrient supplements are
recommended.
Nutritional needs and fertilizer requirements under Florida conditions have
been well studied, although similar studies have not been done in Puerto Rico
(Aponte-Morán et al., 1977; Toro, 1988), Hawaii and California. The sandy and
calcareous soils in Florida are very low in native nutrient content and cation
exchange capacity. These soils require inorganic and organic fertilizers for
optimum growth and production. The leaf mineral con-tent and deficiency
symptoms of young containerized ‘Haden’ and ‘Zill’ trees in sand culture have
been described for N, P, K, Mg, Mn and S (Smith and Scudder, 1951). Obvious
leaf symptoms of Ca, Cu, Zn and B were not observed during a 3-year
investigation, although classical Zn deficiency symptoms were observed in the
field.
458 J.H. Crane et al.

Table 13.9. General nitrogen, phosphate and potash fertilizer recommendations


(g of nutrient/tree/year) for mango trees in Taiwan.

Tree age (years)

Nutrient 1–2 3–4 5–7 8–10 >11

Nitrogen (N) 150 225 240 300 360


Phosphorus (P2O5) 20 75 160 200 240
Potassium (K2O) 120 225 360 450 540

Leaf nutrient levels in mature mango trees on calcareous and sandy soils have
been investigated (Young and Koo, 1969, 1971; Koo and Young, 1972).
Significant variation in mineral content was due to cultivar, leaf age, position of
sample leaf on the twig, presence or absence of fruit and soil type. Ranges of
mineral nutrient levels for maintaining productive trees under Florida conditions
were developed (Table 13.4). Fertilizer frequency, rates and timing in Florida are
based on observation, leaf and soil nutrient content and experi-ence. Leaves are
sampled at least once a year between December and Febru-ary; mature leaves are
collected from at least 30 trees at 0.9–2.4 m and from all sides of the trees, and
before the trees have been sprayed with nutrients (Young and Koo, 1971; Koo and
Young, 1972; Young and Sauls, 1989). Sam-ples should be taken for trees showing
nutrient deficiencies, different culti-vars and from groves under different cultural
programmes and growing in different soil types.

Nitrogen has the greatest effect on tree growth and yield and is used as the
basis for determining the amount of fertilizer to apply; however, some caution must
be used in recommendations from the past because of the change from highly to
less soluble fertilizer ingredients that has occurred over the last 30 years (J.H.
Crane, personal communication). The aim of the programme for the first 2 or 3
years is to produce a strong, healthy canopy so that when production begins, trees
will bear regularly and heavily. Young non-bearing trees (1–3-years-old) are
fertilized with 100–225 g/tree of a
2–10% N and K (K2O) source (e.g. 4-2-8, 4-8-12, 8-2-8-2, 8-3-9; N-P2O5-K2O-
Mg) or similar material also containing P (P2O5) and Mg, at 6–8 week
intervals during the first year (Young, 1974; Young and Sauls, 1989). About 25–
50% of N should be in an organic or slow-release form. The amount of fertilizer is
gradually increased (up to 1.4 kg application/tree) and the fre-quency decreased (2–
4 times/year) during years 2 to 4. Sources of N recom-mended for mangoes in
Florida soils include ammonium nitrate (NH4NO3) and potassium nitrate (KNO3),
some in a slow-release and/or organic form. Urea is not recommended for
calcareous soils because it volatilizes as ammo-nia gas, but S-coated urea is used.
Currently, low N analysis fertilizers are recommended because excessive
vegetative growth and increased fruit physiological disorders (e.g. internal
breakdown) are caused by moderate to high N applications (Young and Miner,
1960, 1961; Nguyen et al., 2004). Cal-cium applications may be necessary to raise
the pH of acid sandy soils in
Crop Production: Management 459

other areas of Florida. Recommended Ca sources include dolomite, gypsum,


calcium carbonate (CaCO3) and calcium nitrate (Ca(NO3)2).
Microelement application methods depend upon soil type, leaf analysis and
other cultural practices (Young, 1974; Young and Sauls, 1989; Crane and
Campbell, 1991). Foliar applications of Mn, Zn and Cu are made to trees growing
in calcareous soils, because soil applications are ineffective due to high Ca content
and pH. Soil applications of chelated Mn and Zn materials formulated for trees
growing in calcareous soils are recommended. Foliar or soil applications of non-
chelated Mn and Zn are appropriate for trees grow-ing in acid sands, although
foliar applications may be more efficacious and less expensive. Usually Cu is
applied as a fungicide and additional applica-tions are unnecessary. Magnesium is
frequently applied foliarly together with Mn and Zn. Typical sources of Mn include
manganese sulfate (MnSO4), manganese nitrate (Mn(NO3)2) and manganese oxide
(MnO), and of Zn include zinc sulfate (ZnSO4), zinc oxide (ZnO) and zinc nitrate
(Zn(NO3)2). Recommended rates for foliar applications include 3.46–5.7 kg each
of a Zn and an Mn sulfate per 937/l of water/ha. However, commercial microele-
ment formulations containing Mn, Zn, Cu, molybdenum (Mo), S and B are used
because of their convenience. Microelements are applied 2–4 times/ year.

Iron is applied to soil as a drench with chelated materials formulated for


calcareous soils; however, these are expensive (Young and Sauls, 1989). Typically,
chelated iron as EDDHA (sodium ferric ethylenediamine di-(o-
hydroxyphenylacetate)) and EDDTA (sodium ferric diethylnetriamine
pentaacetate) can prevent and correct Fe deficiency in calcareous and acid sandy
soils, respectively. For young trees, 7.1–14.2 g/tree applied as a soil drench 2–4
times/year is recommended. Trees are irrigated for 1–2 days before applying a
mixture of chelated iron and water to the soil around the base of the tree and then
irrigated briefly afterwards to ensure the material has moved into the upper soil
profile where most fibrous roots are located.
Foliar applications of ferrous sulphate (FeSO4) and other Fe compounds
(including chelated materials) with or without adjuvants have been ineffec-tive for
maintaining, preventing or correcting Fe deficiency (Leonard and Stewart, 1953;
Leonard and Calvert, 1971; Green et al., 1999). Studies with mild acids plus
organosilicone adjuvant and FeSO4 show promise as an inexpensive foliar iron
application method (Crane et al., 2007, 2008).
The fertilizer programme for fruit-producing trees involves maintenance of
tree health and fruit production and quality. The early recommendations for mature
mango trees growing in calcareous and acid sandy soils were 111–168 kg of N and
K/ha/year (Young and Miner, 1960, 1961; Young et al., 1962, 1965; Young, 1974;
Young and Koo, 1974); however, the recommended N rate has been reduced to 40–
60% of earlier levels (J.H. Crane, personal communication) due to the slow-release
component of modern mixed fertil-izer formulations. Nitrogen applications >225
kg/ha/year reduce fruit qual-ity, cause excessive vegetative growth and increase
internal breakdown (Young and Miner, 1960, 1961; Raymond et al., 1998); leaf
tissue analysis for N should dictate application (Davenport, 2006). Leaf N levels
>1.5% can
460 J.H. Crane et al.

result in little or no flowering. Fertilizer mixtures containing 2–10% N and 4–16%


K are used. Fertilizer rates may be reduced if leaf N and K levels are within
acceptable ranges and tree performance is acceptable. Phosphorus applications may
be reduced if leaf nutrient levels are within the acceptable range; fertilizer mixtures
with 3–5% P are normal. Additional Ca applications, e.g. gypsum, dolomite,
gypsum and Ca(NO3)2, on acid sandy soils increase Ca leaf levels and reduce the
incidence of internal fruit breakdown (Young et al., 1962).

Microelements are applied to mature trees regardless of leaf analysis as a


preventative measure. This is especially true for Fe and Zn, because correct-ing
deficiency of these elements is sometimes difficult and expensive. Rates for foliar
sprays of Mg, Zn and Mn are similar to those for young trees with 2–4
applications/year. The rates for chelated iron for mature trees range from 14 to 113
g/tree 1–2 times/year.
Fertilizer may be applied by fertigation. Mixed fertilizers containing P are not
recommended because phosphates can precipitate and clog the irri-gation system.
Phosphate can be applied alone as dilute H3PO4. Iron can also be applied through
microsprinklers or a drip system at increased frequency and reduced rates.

In Puerto Rico, fertilizers are also applied in dry form and through low-volume
irrigation systems. Young non-bearing trees (1–3 years old) are fertil-ized with
454–1591 g/tree of a 12–15% N, 5–6% P (P2O5) and 8–10% K (K2O) source (e.g.
12-6-8, 12-6-10, 15-5-10; N-P2O5-K2O), split into two applications
(December/February and April/May) (Toro, 1988). When trees begin to bear (after
3–4 years), 10-5-15, 12-6-16, or 10-5-20 is applied in split applications which
increase in amount with tree age (3.4–6.8 kg/tree maximum). Some orchards are
fertilized through the irrigation system at 7–15 day intervals (E.E. Toro, personal
communication). In these plantings, (NH4)2SO4, urea, phosphoric acid (H3PO4)
and KNO3 or potassium sulfate (K2SO4) are used. Micronutrients are applied
either to the soil or to foliage in Puerto Rico; rec-ommendations for young trees are
foliar applications at 21–30 day intervals during the growing season (Toro, 1988).
Micronutrients are applied as needed in mature orchards and may be applied once
during the spring and autumn. Sources of micronutrients are sulfate forms of Zn,
Cu and Mn.

13.9 Irrigation Practices


Water requirements for mango production are not precisely known and cur-rent
irrigation practices are based on experience and climatic and limited edaphic
measurements. Irrigation practices are influenced by available tech-nology and
cost, soil type and depth, rainfall amount and distribution, fertil-izer practices and
production objectives. The response to irrigation is influenced by the rate, timing
and method of application, tree-growth stage, climate, edaphic conditions and
cultivar.
Irrigation of young trees assists tree establishment, prevents drought stress,
and maintains constant canopy and root growth. Objectives for bearing
Crop Production: Management 461

trees are maintenance of tree health, avoidance of severe drought stress and
enhancement of vegetative dormancy. Mangoes are drought tolerant (Schaffer et
al., 1994); however, in some production areas under non-irrigated condi-tions, fruit
production and quality may be reduced. Excessive irrigation can cause reduced tree
growth and tree decline (Larson et al., 1989a, 1991d; J.H. Crane, personal
communication). Mango production has been displaced onto marginal lands that
possess low water and/or nutrient holding capacity and high pH. Saline water for
irrigation is a concern. Irrigation methods include flood and furrow, high-pressure
volume guns, high-volume under- and over-tree sprinkler and low-pressure
microsprinkler and drip systems.
In north-eastern Brazil, under semi-arid tropical conditions, irrigation is
essential throughout the hot and dry season (Albuquerque et al., 1999). Sev-eral
irrigation systems are used, with c.41% of orchards using microsprinkler systems.
About 21% use other types of irrigation (e.g. furrows, drip, basin, etc.) and 33% of
orchards use no irrigation (Gomes et al., 2002). Mean produc-tivity of irrigated
orchards may be as high as 25 t/ha compared with only 12 t/ha for non-irrigated
orchards (Coelho et al., 2002). Irrigation is used on 14,500 ha (74% of the
cultivated area) of commercial mango in the north-east. Microsprinkler irrigation is
the most common method (30.3% of the irrigated area) (Gomes et al., 2002).
Commercial mango orchards in the south-east, mainly in São Paulo, are not
irrigated.
Fertigated orchards require well-trained people. Some nutrients can cause
corrosion of irrigation pipes, and proper management is essential to prevent
environmental damage. Proper selection of nutrients is critical due to problems of
solubility and compatibility. Urea, (NH4)2SO4 and KNO3 are the primary N
fertilizers; they are highly soluble and are compatible with most nutrients.
However, the SO42– is incompatible with Ca and their mixture causes precipitation
and clogging of emitters.
The oldest commercial orchards in Mexico were rainfed and established in
areas with deep soils and annual rainfall >900 mm (subhumid and humid tropics).
However, growth of the mango industry has forced the planting of semi-arid
tropical areas (annual rainfall 600–700 mm) (Colima and Micho-acán) (Table
13.2). Currently, 67% of orchards are rainfed (SIIAP, 2007) with annual rainfalls
that fluctuate from 900 to 3700 mm; 80% of the rainfall occurs in June–October.
Irrespective of the amount of annual rainfall, since the late 1980s a significant
proportion of the new and old mango orchards (60,000 ha) have installed irrigation
systems. This is to prevent water deficits and to increase yield and fruit size and
quality. Sources of irrigation water include rivers or deep wells and irrigation may
be applied either by gravity or by electrical or diesel engines.

Irrigation management involves either furrow or pressurized systems. The


furrow soil surface system (FSSS) can be simple, crossed, furrow-basin and fish
spine. Low-pressure drip and microsprinkler systems are most com-mon,
particularly the latter. Two microsprinklers (90–120 l/h) or 10–15 drip emitters (8–
12 l/h) are used per mature tree in a low-pressure system. In semi-arid regions,
mango potential evapotranspiration from October to June each year is 16,000
m3/ha. The FSSS system applies >25,000 m3/ha of water;
462 J.H. Crane et al.

however, low-pressure systems can reduce water use to 5000 m3/ha with no
negative effect on yield or fruit quality (L.M. Tapia, personal communica-tion).
Water requirements in the semi-arid region of Michoacán are calculated by using
evapotranspiration measured with a Class A pan and multiplied by the crop
coefficient Kc, which for practical purposes is considered as 0.4 for vegetative
growth and 0.8 for fruit growth and development (Chávez-Contreras et al., 2001).
The FSSS water schedule for obtaining maximum yield and fruit quality in
Michoacán is: September (pre-bloom), one 20 cm irrigation; October–December
(bloom), two 10 cm irrigations spaced 20 days apart; January–April (fruit growth),
eight 10 cm irrigations at 15–17 day inter-vals; May–July (vegetative growth),
three 50 cm irrigations at 21–25 day inter-vals. The amount of water used for drip
irrigation in mature mango trees during fruit growth and development is 1350
l/tree/week and for microsprin-klers it is 1560 l/tree/week. After the first irrigation,
growers water every 8 days in low-pressure irrigation systems and every 15–20
days in FSSS.
Soil-water content influences the frequency of flowering, the number and
length of panicles, yield and quality of mango fruits in Taiwan (Chang and Lu,
1995). Two types of irrigation system are used: (i) furrow flooding in the low
lands; and (ii) microsprinklers, overhead sprinklers and drip (trickle) irrigation on
sloped lands. Microsprinklers have become the most important irrigation system in
recent years and fertigation is utilized by many produc-ers. Irrigation management
is based on producer experience and observation of the soil, current weather and
tree phenology.
In Florida, rainfall is not evenly distributed through the year, with the dry
season occurring during the autumn–winter (October–April). During the wet
spring–summer season (May–September), dry periods of 3 days or more can occur.
Most orchards in Florida are irrigated only during prolonged dry periods. Several
systems are common. High-volume overhead or under-tree sprinklers which run at
high pressures (28,121–70,303 kg/m2) and distribute large amounts of water (0.51–
0.89 cm/ha) and microsprinkler systems which run at low pressure (7030–28,121
kg/m2) and distribute lower volumes of water (37.9–1113.6 l/tree/ha). Some
growers use both systems: the microsprin-kler system for irrigation and fertigation
and the high-volume system for cold protection during freezing weather.

Well-established trees require little to no irrigation. During prolonged drought


conditions (>30 days with no significant rainfall), irrigation may be applied.
Irrigation is not recommended during the cool autumn and winter months, to
enhance the vegetative dormancy period induced by cool tem-peratures, to enhance
synchronization of apical stems and to intensify the flowering response after
growth commences.
Currently, irrigation recommendations and practices are based on obser-vation
and experience with c.6.35 cm water/application/ha. Preliminary research suggests
that irrigation at 7-day intervals during the period of fruit development increases
fruit size, earliness and yield (Larson et al., 1989b); however, this has not been
implemented by growers. Most Florida producers rarely irrigate their mango
orchards during the spring and summer as rain-fall during this period coincides
with fruit development and is sufficient for
Crop Production: Management 463

fruit production. In Puerto Rico, Hawaii and California, microirrigation is used. In


Hawaii, where temperatures are consistently warm in the mango-growing areas,
mature trees are more productive if irrigation is withheld for at least 2 months
before flowering (Chia et al., 1988). Drip and microsprinkler irrigation is used in
California, but irrigation regimes have not been reported. In Puerto Rico, drip
irrigation is more common than microsprinklers (E.E. Toro, personal
communication). The drippers apply 3.79–7.75 l/tree/ha and trees are irrigated 2–3
times/week. When microsprinklers are used, one microsprinkler is placed between
two trees in-row. Trees may be irrigated from 9–11 months of the year; water is
withheld for 1–3 months prior to floral induction or flowering.

13.10 Vegetative Growth and Reproduction Manipulation


Mango trees require new vegetative growth in order to produce fruit each year. The
optimum temperature for vegetative growth is 24–30C (Krishnamurthi et al.,
1961; Shü and Sheen, 1987; Whiley et al., 1989) and when levels of essen-tial
plant nutrients and water are not limiting, vigorous growth results. Mature leaves
and a period of cessation of vegetative growth (i.e. mature api-cal and subtending
meristems) are required for the transformation from veg-etative to reproductive
growth (Núñez-Elisea and Davenport, 1992; Kulkarni, 2004; Davenport, 2006).

Canopy management and reproductive manipulation vary according to


climatic conditions, cultivar and available technology (Davenport, 1993). Ultimate
mango-tree size depends upon the climactic and edaphic condi-tions and cultural
practices. Under optimum conditions, trees can reach heights and canopy diameters
of 30 m or more (Kostermans and Bompard, 1993); however, it is difficult to
protect large trees from insects, diseases and strong winds, and harvesting is
difficult and costly. With the increase in costs for orchard establishment and
maintenance, the number of trees per unit area has increased and tree size has
decreased, whereas production of high quality fruit per unit area has increased.
Growers in some production areas manipulate the period of flowering and fruit
production. Tree size can be controlled to maximize the number of trees per unit
area and maintain productive tree canopy and yields.

The natural period of mango production in Brazil was originally from October
to January. Production has increased from September through to April through the
use of precocious and late-bearing cultivars along with floral induction techniques.
There is potential that whole-year harvesting and mango supply can be achieved,
although there are some months with low mango yields. There are three types of
floral induction and their use depends upon phenology and season. In general, the
steps are: (i) pruning of apical internodes after harvesting; (ii) application of
paclobutrazol (PBZ) to stimulate flowering by inhibiting gibberellin biosynthesis;
(iii) spraying with K2SO4 (2–2.5%) to increase carbohydrate levels in the tree; (iv)
drought stress to facilitate growth cessation; (v) applications of ethylene (optional);
and
464 J.H. Crane et al.

(vi) three applications of 3–4% KNO3 and Ca(NO3)2 to trigger flowering


(Albuquerque et al., 2002). This flower initiation protocol results in off-season
flowering and fruit production when mangoes are scarce and prices are high
(between April and August) in the domestic market.
Tree-size control and pruning strategies appropriate for the various cul-tivars
and diverse climatic regions in Mexico are under development. Researchers in
conjuction with producers are investigating appropriate tech-nologies that take into
account the current tree size, tree spacing and configu-ration in the orchards, the
available technology, and the goal of the pruning programme. In young trees the
central leader is pruned (5 mm above previ-ous growth scar) a year after planting to
promote lateral branching and pre-cocious flowering. Once new branches have
matured the best-positioned 2–5 lateral branches around the main stem are selected.
Trees are pruned for 2–3 years after every 2–3 vegetative flushes by heading back
the wood of the last two growths (Guzmán-Estrada and Vázquez-Valdivia, 2006).
Young tree training is important for ‘Ataulfo’ and ‘Keitt’ and results in strong
canopies that will resist heavy fruit loads and strong winds. Tree training is
unnecessary for ‘Manila’, ‘Haden’, ‘Tommy Atkins’ and ‘Kent’.

Usually mature trees are pruned every year after harvest to promote light
penetration into the canopy and to remove weak, diseased, broken and poorly
positioned branches. Mechanical topping and hedging is not widely practised.
However, this pruning system was recently introduced in orchards in Nayarit to
control the pink hibiscus mealybug (Maconellicoccus hirsutus Green). In Chiapas,
8-year-old trees and older are pruned by hand (machete) in alternate years to delay
or avoid canopy crowding; one side of the canopy is pruned in one year and the
other side is pruned the next year.
To rejuvenate old, very large mango trees and/or overcrowed orchards, trees
are cut back to main limbs at 1–2 m above the soil level. This technique is common
in the subhumid and semi-arid production areas of Chiapas and Veracruz.
However, in the warm humid tropical areas of Chiapas and Veracruz this technique
is not widely used because of the subsequent loss of fruit pro-duction and excessive
regrowth (3–4 vegetative flushes/year) that occur. Selected trees can be removed to
avoid overcrowding.
Climatic conditions are conducive to early bloom and harvest from the
Chiapas to Michoacán production regions, where forced blooming and early fruit
production is used by >80% of producers to obtain higher prices. Even when a
significant amount of advanced bloom is induced, blooming during the normal
flowering period may occur. The intensity of the normal bloom is influenced by the
intensity of the fruit set by the advanced bloom. The most common method to
advance bloom involves canopy sprays of KNO3 or NH4NO3 (Mosqueda-Vázquez
and De los Santos, 1982; Núñez-Elisea, 1986, 1988; Guzmán-Estrada, 1991;
Sandoval-Esquivez et al., 1993). The effect of these compounds is influenced by
cultivar and environmental conditions (probably temperature). In the Gulf of
Mexico region, one to two sprays of 2% KNO3 or 1% NH4NO3 solution are
applied to ‘Manila’ trees any time from 15 October to 30 November to stimulate
early flowering. ‘Manila’ and ‘Ataulfo’ are sprayed with 2% KNO3 during the
same period in the Southern
Crop Production: Management 465

Pacific region. In the Central Pacific region, ‘Haden’, ‘Manila’ and ‘Ataulfo’ trees
are treated with one or two sprays of 2–4% KNO3 or 1–2% NH4NO3 at any time
during the first half of November. Similarly, ‘Haden’ and ‘Manila’ are treated
during November with 8% KNO3 or 4% NH4NO3 in the Northern Pacific region.
‘Tommy Atkins’ does not respond to foliar nitrate treatments for promoting early
flowering. This is due to delayed floral initiation in this cultivar so that when
nitrate treatments are applied the buds are not irrevers-ibly committed to flowering
(Pérez-Barraza et al., 2000). Consequently, veg-etative growth is produced in
response to treatments that stimulate bud break (Pérez-Barraza et al., 2006a).
However, application of PBZ promotes early bloom in ‘Tommy Atkins’ (Salazar-
García and Vázquez-Valdivia, 1997).
Currently, soil applications of Cultar® (25% a.i.) close to the tree trunk is used
for most cultivars in dosages that range from 1 ml/m canopy diameter for ‘Manila’
in Veracruz to 2–4 ml for ‘Tommy Atkins’, ‘Haden’ and ‘Ataulfo’ in Michoacán,
applied at 1–2 year intervals. The response to PBZ treatment is enhanced by 30–45
days water stress and canopy sprays with nitrates (Chávez-Contreras et al., 2001).
In Nayarit and Sinaloa late bloom and harvest are profitable because they are the
last two production areas to be harvested in Mexico. Two canopy sprays of 50 mg/l
GA3 (15 and 30 November) cause delayed bloom and shift 86% of the harvest to 1
month later (Pérez-Barraza et al., 2006b).

In Taiwan, cultivar, latitude, elevation and cultural practices are used for off-
season production (Shü et al., 2000). Several strategies are utilized to stimulate
flowering and fruit set:

1. After harvest the last one or two vegetative flushes on each shoot are cut back
to control tree size. Subsequently, two flushes of healthy shoots are allowed to
grow to serve as fruiting shoots for the next year. Weak or crowd-ed shoots are
removed to facilitate ventilation and light penetration.
2. In general, flowering is not a problem in subtropical Taiwan; however, poor
fruit set due to bad weather and lack of pollinators occurs occasionally. A recent
programme to increase the population of pollinators, mainly the greenbottle flies
(Chrysomyia megacephala Fabricius) in mango orchards has been very successful.
Increased yield has been noted and the practice has been exploited commercially
throughout the island.
3. Most of the mango fruit harvest goes to the domestic market within a short
time period, causing the price to decline rapidly. Off-season fruit pro-duction is
thus very important to avoid this sharp drop in price.
4. Physical trunk damage by girdling and ringing and application of ethrel is used
to promote early flowering of the early season cultivar ‘Tsar-swain’ (Liu, 1996).
Foliar applications of KNO3 can promote early flowering but only if applied to
flower-bud-induced shoots; PBZ is not recommended for use on edible crops in
Taiwan.
5. Panicle removal has been used to postpone flowering and fruiting (Shü and
Sheen, 1987; Shü, 1993). Emerging terminal panicles are removed by hand.
Chemical removal of terminal panicles with hydrogen cyanamide (CH 2N2) or
calcium cyanamide (CaCN2) causes leaf damage and is not as
466 J.H. Crane et al.

effective as pruning (Hwang et al., 2004). Axillary panicle induction has some
advantages, because flowering can be timed to avoid frost or cold tempera-tures or
a period of excessive rainfall during the normal flowering period (Singh et al.,
1974; Shen and Huang, 1980). Axillary panicle removal also reduces mango
malformation and reduces alternate bearing (Majumder et al., 1976; Pal and
Chadha, 1982).

Control of flowering and tree size in the USA varies with respect to differ-ent
climatic, edaphic and soil conditions as well as cultural practices (Daven-port,
1993). In Florida, cool temperatures during the winter months (December through
to February) are usually adequate to arrest vegetative growth and induce flower
bud differentiation. In some years, the duration of cool tempera-tures prohibits
floral expression until late winter/early spring (February/ March) and when
continuous warm temperatures begin (March), profuse, syn-chronized flowering
occurs. In some years, warm and cool periods may occur for a few days to weeks
during the winter and partial flowering may occur 2–4 times during the winter.
This prolongs the flowering and harvest season.
Flowering and fruiting of trees are not actively manipulated in Florida. For
young trees, vegetative growth is encouraged during the first 2–3 years and
panicles may be removed by hand or natural pathogens, i.e. powdery mildew or
anthracnose are allowed to kill panicles and flowers. Tipping and selective pruning
of young trees is recommended to improve tree structure, control tree size and
enhance early fruit production (Oosthuyse and Jacobs, 1995; Campbell and
Wasielewski, 2000); however, selective pruning and mechanical topping and
hedging are used to control mature-tree size. Selec-tive pruning usually involves
removal of selected scaffold limbs to open up the tree canopy to light and to
remove dead wood. Mature trees may or may not be allowed to grow together in
the tree row to form hedgerows. Periodic mechanical topping at 3.5–5 m and
hedging to leave a 2.5–3.5 m row middle is common (Crane and Campbell, 1991;
J.H. Crane, personal communication). Limiting the between-row spread of the trees
to 2.5–3 m improves light pen-etration into the tree canopy. In some orchards,
hedging the inner sides of the canopy of adjacent rows every 2–4 years and/or
topping every third or fourth row every 2–4 years is recommended. Trees are
mechanically pruned immedi-ately after harvest. Timing the pruning to selected
rows each year ensures that most of the planting will always be productive if
continuous flushing of pruned parts of the canopy prohibits reproductive growth
the following spring. Pruning trees shoots of 2–10 cm diameter immediately after
harvest and then tip pruning 3–4 times to force multiple lateral growths has been
advocated (Davenport, 2006). This strategy: (i) reduces or controls tree size;

(ii) shapes trees to facilitate subsequent tip pruning; (iii) synchronizes the veg-
etative flushing; and (iv) inhibits continuous vegetative flushing and prolongs the
period of vegetative dormancy. After vegetative growth has ceased for 5 or more
months, trees will synchronously flower when regrowth occurs.
In Puerto Rico, continuous vegetative growth of 1–2-year-old trees is
promoted. During this time panicles may be removed by hand to promote
vegetative growth, and this encourages rapid development of large trees.
Crop Production: Management 467

Temperatures are insufficient to inhibit vegetative growth and induce reli-able and
consistent bud differentiation, causing erratic or poor flowering and poorly
synchronized fruit yields. The most common method for synchroniz-ing and
enhancing the time of flowering involves a combination of drought stress and
timed application of KNO3. This method depends on cultivar and the desired time
of fruit harvest. To slow (or stop) vegetative growth and to stress the trees,
irrigation is withheld for 1–3 months prior to flowering. The drought stress is
prolonged until leaves become dark green and show slight signs of wilting. A 1–
5% solution of KNO3 is applied to the foliage, which induces flowering 3–4 weeks
later. Regular irrigation is resumed when c.75% of the panicles have set fruit. The
timing of the drought stress and KNO3 spray varies with cultivar and when fruit
harvest is desired.
PBZ has been used to control vegetative growth after trees flush in response to
pruning following harvest. A soil drench of PBZ (7–10 g/tree) is applied and trees
are irrigated for 8–12 h. Irrigation is then withheld for 30–90 days until trees show
signs of drought stress. A foliar application of 1–2% KNO3 is applied, and
flowering occurs 30–45 days later. The amount of tree training depends upon the
cultivar and is practised on young trees when they are c.1–1.5 m high. ‘Keitt’ and
‘Palmer’ tend to have long branches of various lengths and benefit from training to
create a stronger limb structure and more compact tree. Trees are generally trained
to a modified central leader system, and some selective pruning of older trees is
practised to open the canopy to more light and air movement. Mechanical topping
and hedging are used to shape mature trees. Usually trees are topped to 3–4.5 m
immedi-ately after harvest.

In Hawaii, seasons with heavy crops are commonly followed by light or no


crops for 1–2 years (Nagao and Nishina, 1993). The small variation in warm
temperatures and evenly distributed rainfall encourage vegetative growth, which
reduces the potential for fruit production. This is less prob-lematic on the drier
leeward sides of the islands. Diseases (i.e. anthracnose and powdery mildew)
reduce production by infecting panicles and flowers. Preliminary trials with 2 and
4% KNO3 applications during the winter (Feb-ruary) resulted in 66–84% flowering
5 weeks after treatment (Nagao and Nishina, 1993); however, this procedure is not
utilized commercially.
Young ‘Keitt’ mango trees are extensively trained by hand (usually twice a
year) during the first 4–5 years to improve the structural strength of scaf-fold
limbs, increase the number of terminals and to control tree size in California.
Panicles are removed by hand during the first 4 years. Training is necessary
because non-trained ‘Keitt’ trees tend to have a few very long scaf-fold limbs, little
branching and are structurally weak. Bearing trees are pruned annually to maintain
trees at 3.1–4.6 m. In California, cool tempera-tures during the winter months
(November through to February) are ade-quate to arrest vegetative growth and
induce bud differentiation. The duration of cool temperatures may inhibit floral
expression until early spring (March and April), when warm temperatures allow
profuse, synchronized flowering. Warm periods during winter may stimulate early
flowering, which may be damaged by subsequent cold temperatures (Schacht,
1992).
468 J.H. Crane et al.

Early season flowering when cold temperatures and dry windy conditions prevail
(December–February) results in poor fruit set and abnormal fruit. Therefore,
pruning of mature trees just before April (early spring) delays flowering and
induces synchronous axillary flowering after the danger of cold has passed.

13.11 Environmental Stress Management


Mango is an adaptable species that withstands a range of subtropical and tropical
climates and soils. Physiological responses are related to the evolu-tionary history
of mangoes, with monoembryonic cultivars better adapted to the subtropics and
polyembryonic cultivars better adapted to the tropics.
In Brazil, the important commercial mango areas are in the tropical semi-arid
climate of the north-east. Flooding and freezing rarely occur there, but wind and
drought stress are very common and negatively affect growth of young mango trees
and increase fruit drop. Windbreaks reduce wind stress; compact rows of elephant
grass and/or rows of banana trees are utilized (Mouco et al., 2002). In general, 3–4
rows of banana trees are planted around or in perpendicular rows in the orchard
against the main wind flow.
Drought stress, particularly in north-eastern Brazil, can suppress mango
growth and production, and irrigation is used to ameliorate this problem. Drought
stress has been used to increase endogenous ethylene concentra-tions and trigger
floral induction 70–90 days after PBZ application. Drought stress should be
avoided during fruit development. Coelho et al. (2002) reported the crop
coefficient (Kc) for mango increased from 0.39 at flowering to 0.85 during fruit
development.
In Mexico, the Gulf of Mexico coastal region may experience strong, dry
northerly winds (11–28 m/s) from October to April and cause limb breakage and
increase flower and fruit drop by desiccation. To ameliorate this prob-lem, growers
have planted east-west oriented bamboo (Bambusa vulgaris) and Australian pine
(Casuarina equisetifolia) windbreaks. Bamboo is planted outside the Australian
pine trees and two rows of pines are planted in a stag-gered arrangement. All
coastal mango-producing regions in Mexico are potential targets of hurricanes
during the summer rainy season, even in late October. Broken limbs are pruned,
damaged trees are pruned back to sound wood, and some trees are replaced or top-
worked.
Flooding for 2–3 weeks may occur during the summer rains. This is mainly a
problem in low-lying areas of Chiapas and Nayarit that have loamy soils and a high
water table. Drainage canals have been constructed in such areas to minimize the
problem. Drought stress is common as 67% of Mexico’s mango orchards are not
irrigated (SIIAP, 2007). Annual rainfall ranges from 900 to 3700 mm in Mexico
and is concentrated in June to October. The dry season begins in October (close to
the period of floral initiation) and contin-ues through the harvest period for early
and some mid-season cultivars. Drought stress in areas of low rainfall causes
intense fruit drop, which reduces yield and fruit size especially in heavily bearing
trees. More that 60,000 ha of
Crop Production: Management 469

mangoes are irrigated in low rainfall areas. In Veracruz and Chiapas most orchards
are adjacent to riverbanks and the deep soils provide root access to the water table.
No data are available supporting the benefit of irrigation in medium-high
precipitation areas.
Most mango-producing regions in Mexico are frost free. The only report of
frost damage to 2–4-year-old mango orchards was in Sonora where freez-ing
events may occur every 6–8 years (E. Sánchez, personal communication). Low
temperatures (≤10°C) during bloom reduce fruit set, especially in ‘Ataulfo’.
Treatment with GA3 to delay the bloom has been suggested as a method to avoid
low-temperature damage during flowering but flowering under high-temperature
conditions is also detrimental to fruit set and crop yield (Pérez-Barraza et al.,
2006b).
In Taiwan, damage from typhoons occurs periodically and trees are reset and
either pruned to recover or replanted. In general, freezing temperatures are not a
problem in the mango-production areas but cool temperatures can reduce fruit set.
To avoid cool or cold temperatures during the normal flow-ering period, shoot tips
may be pruned to delay and force flowering from lateral buds. Flooding in
production areas is uncommon, and drought stress is not an issue because most
producers irrigate during dry periods.
The production areas of Hawaii and Puerto Rico are frost free; however,
Florida and California may experience temperatures at and below freezing during
the winter months (December through to February). In Florida, freez-ing
temperatures (0 to −6°C) may occur for a few hours for 1–4 nights/year, although
freezing temperatures for 13–15 h within a 24 h period have been reported
(Johnson, 1970; Campbell et al., 1977). In California, temperatures as low as
−6.6°C are common and frosts may occur 10–15 times/year during January and
February (Aslan et al., 1993). Mango trees do not acclimate to cold temperatures
(McKellar et al., 1983), although differences in cold toler-ance and recovery from
cold damage have been observed (Carmichael, 1958). In Florida, high-volume
overhead and under-tree irrigation is used to protect trees during freezing weather.
At least 0.6 cm of water/ha is distributed. Overhead systems are designed for
complete coverage (overlapping spray patterns) of the trees and under-tree systems
are designed to spray 0.9–2.4 m into the tree canopy. Irrigation commences before
freezing temperatures are reached (usually c.2–3°C) and continues until ice has
melted. These systems are powered by diesel or gas engines as electrical power is
unreliable during freezing weather conditions. In California, microsprinklers, wind
machines and helicopters are used to raise the air temperature of plantings
(Schacht, 1992).

Mango trees are relatively tolerant of wind stress (Schaffer et al., 1994; Crane
and Balerdi, 2005); however, newly planted trees are commonly staked at planting
in the calcareous soils of Florida to prevent damage to the bark and cambium
caused by constant movement and rubbing against the rocky soil. Staking also
stabilizes the tree against toppling during hurricanes. In contrast, tolerance to
mature trees depends on tree size, with larger trees being more vulnerable to wind
damage than pruned trees (Crane et al., 1993, 1994, 2001; Crane and Balerdi,
1996; NASS, 2006).
470 J.H. Crane et al.

In the mid-20th century, windbreaks of C. equisetifolia were planted around


many mango orchards in Florida. However, intrusion into the orchard, shading and
damage to mango trees when trees toppled into orchards during hurricanes has
stopped this practice. Some producers have topped remain-ing pine windbreaks at
4.9–6.7 m to reduce orchard shading and their potential to topple (Crane et al.,
1993). In Hawaii, natural windbreaks are recommended for some areas where
constant winds are a problem for establishing young trees (C.L. Chia, personal
communication). In California, constant winds during spring may damage panicles
and young fruit; man-made windscreens are used to protect trees (Scott, 1990;
Schacht, 1992). The windscreens can be raised to prevent trapping of cold air
within the plantings. Puerto Rico is affected by hurricanes but no specific
ameliorating recommendations have been reported (Toro, 1988).

Flooding is not common in production areas of Puerto Rico, California and


Hawaii; however, periodic flooding (1–21 days) is typical during the summer in
Florida. Trees have been planted on beds of crushed limestone rock, 0.6–1.0 m
high and 1.0–1.5 m wide, which allows part of the root system to be above water.
Planting of orchards at sites at or below 2 m above sea level is not recommended.
Mango trees are moderately flood tolerant (Larson et al., 1991c; Schaffer et al.,
1994, 2006), although this is affected by floodwater tem-perature, oxygen content
and anatomical adaptations of the rootstock (Lar-son et al., 1991a, 1993a, b).
Periodic flooding of the limestone-based soils in Florida increases the availability
of soil Mn and Fe (Larson et al., 1991b, 1991d, 1992), alleviating plant
deficiencies of these elements. Furthermore, rhizo-sphere anoxia increases
reduction of Fe3+ to Fe2+ by nicotinamide adenine dinucleotide (NADH) in mango
root tissue and Fe uptake by mango roots in oxygen-depleted media (Zude-Sasse
and Schaffer, 2000).

13.12 Harvesting Practices


Harvesting is done by hand in Brazil, Mexico, Taiwan and the USA (Evans, 2007)
and is one of the most expensive operations in mango production because fruit do
not mature synchronously, and trees require multiple pickings. The technology to
mechanize mango harvest is difficult because of differences in fruit colour, size
and weight among cultivars, difficulty in determining fruit maturity, requirement
for multiple pickings, lack of tree-size-controlling rootstocks and tree training and
moderate to large tree canopies.
Prior to harvesting in Brazil, excessive set fruit and injured and diseased fruit
are removed, part of the rachis is removed to prevent scarring and bruis-ing of the
fruit, and some leaves that shade fruit may be removed to allow for better peel
colour development (Alves et al., 2002). Fruit in the lower part of the canopy are
harvested from the ground; however, ladders are required for picking fruit high in
the tree canopy. Peel damage due to latex exudation at the stem end of the fruit
during harvest is a very common problem, and >50% of harvested fruit may be
affected. This problem occurs mainly when fruit are picked high in the canopy with
a picking pole by severing the fruit
Crop Production: Management 471

near the stem end of the pedicel. An improved picking pole that cuts the petiole c.2
cm above the pedicel reduces latex burn to <10%. Before trans-porting to the
packing house, fruit containers are placed in the shade to avoid increasing the pulp
temperature.
In Mexico, harvesting is done by hand when fruit have reached physio-logical
maturity. Picking poles with bamboo baskets (Gulf of Mexico and Southern Pacific
regions) or nylon net bags or cotton bags (Central and Northern Pacific regions)
and a cutting blade at the distal end are used to reach fruit high in the canopy.
Ladders may be used, although climbing trees is more common. Fruit is usually
placed in 20–30 kg wooden or plastic boxes. The inner walls of the picking crates
are covered by newspaper to absorb latex and decrease sap peel damage.

Several fruit maturity indexes exist for mango: pulp TSS content and/or
acidity, pulp firmness, skin or pulp colour, calendar days or heat units from bloom
to harvest. No index alone is successful because of differences among cultivars and
significant variability in fruit maturity within trees and orchards and among
orchards. Pickers are trained to distinguish the following charac-teristics: (i) size,
form and fruit colour; (ii) shoulder development (higher than the base of peduncle);
(iii) cavity formation at the base of the peduncle; and (iv) increased lenticel size
(Chávez-Contreras et al., 2001). The Associa-tion of Mango Packers and Exporters
(EMEX; Empacadoras de Mango de Exportación) have established some minimal
physical and chemical stan-dards to define maturity for ‘Haden’, ‘Tommy Atkins’,
‘Kent’, ‘Keitt’ and ‘Ataulfo’. A photographic maturity index chart is used by
growers, pickers, companies and packing houses. Mango fruit-peel damage by
latex is common when tree sap pressure is high. To minimize this damage,
irrigation is stopped at least 2 weeks prior to harvest; fruit are also picked with a
long peduncle and are washed immediately after the harvest bin is full.

Mature-green mangoes are the first fruit to be harvested and all the cri-ollo,
Oro and Florida cultivars fit this category. These mangoes are for the domestic
market and usually sell for high prices. ‘Manila’ mangoes are picked when their
colour changes from pale green to greyish green. ‘Ataulfo’ is harvested when the
green peel shows a yellow colour break. The Florida mangoes are picked ‘mature-
green’ and at colour break.
The harvest season in Taiwan begins in March with ‘Tsar-swain’ and ends in
September or October with ‘Keitt’. Depending on the cultivar and market, fruits are
harvested at 70–80% maturity. ‘Tsar-swain’ is harvested either at the green stage
for pickles or at a mature ripe stage for domestic markets. ‘Irwin’ fruit are
harvested at either 80% maturity for export or at >90% maturity for local markets.
Green mature fruits may be triggered to ripen with calcium carbide, ethephon or
ethrel at 30–40C, depending on the cultivar. Mature fruits are stored at at 8–12C
for several days to several weeks. Fruit destined for export are disinfested using the
vapour heat method; the fruit core temperature must reach and be held at 46.5C
for 30 min.
There are two markets for mango producers in the USA: the ‘green’ mar-ket
for non-ripe fruit and the ‘tree-ripened’ market. The green and tree-ripened markets
are speciality, niche markets where the green fruit are used as a
472 J.H. Crane et al.

component of processed foods (e.g. chutneys, preserved pieces), whereas the tree-
ripened fruit target the demand for ready-to-eat mangoes. Green mangoes are
picked before the fruit is mature, whereas the tree-ripened mangoes are allowed to
develop almost full colour and ripeness before being picked. Florida mangoes are
more expensive than imports and the volume of fruit is very limited and cannot
supply the demand of the national market. Usually multiple pickings are required
to harvest the crop but this is influenced by market demand and prices, the number
of blooms producing the crop and weather conditions.

Harvesting in the USA is by hand. A long picking pole with a canvas or nylon
bag attached to a metal ring with a cutting blade at the distal end is commonly used
in Florida (Aponte Morán et al., 1977; Crane and Campbell, 1991). Other picking
aids such as ladders and mobile hydraulic lifts are also used. Time of harvesting
depends upon cultivar, the intended market and market demand. In Florida, green
mangoes may be picked after March, while fruit-picking time for the fresh market
depends upon the cultivar reaching the mature stage desired (Crane and Campbell,
1991). In Puerto Rico, man-goes are harvested from March to November,
depending upon cultivar, mar-ket price and the date when flowering was induced.
The mango season in California is restricted to September/November, and in
Hawaii the main sea-son is May to August. Approximately 50% of mangoes
produced in California are certified organic (Linden, 2006).

13.13 Conclusions
Differences in mango culture are due to the climatic and edaphic conditions,
available information and technology, and tradition in each production area. The
interaction of climate and cultural practices, for example irrigation, fer-tilizer,
pruning, etc., that are essential for optimizing crop yields and quality is not
completely understood. None the less, horticultural systems can be developed and
tested that can impact fruit production and quality. It is important that area- and, in
many cases, site- and cultivar-specific cultural information and practices need to be
developed to optimize production and fruit quality. This chapter has addressed the
current state of mango culture in four different production areas, and has
emphasized improvements that are essential for a prosperous industry.

Ackowledgements
The Mexican co-author acknowledges the following INIFAP researchers, based at
several Research Stations (CE) and states: Ernesto Sánchez-Sánchez, CE-Valle del
Yaqui (Sonora); Camerino Guzmán-Estrada, CE-Sur de Sinaloa (Sinaloa); R.
Mosqueda-Vázquez (deceased) and Enrique N. Becerra-Leor, CE-Cotaxtla
(Veracruz); Fulgencio M. Tucuch-Cauich, CE-EDZNA (Campeche);
Crop Production: Management 473

Rubén Cruzaley-Sarabia, CE-Iguala (Guerrero); Víctor Medina-Urrutia, CE-


Tecomán (Colima); Víctor Palacio-Martínez, CE-Rosario Izapa (Chiapas).

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14 Postharvest Physiology
J.K. Brecht1 and E.M. Yahia2
1
University of Florida, Florida, USA
2
Universidad Autónoma de Querétaro, Querétaro, Mexico

14.1 Introduction 484


14.2 Contribution of Mango Fruit to Human Nutrition and Health 485
14.3 Mango Ripening Physiology 491
Climacteric behaviour 491
Ethylene production and responses 493
14.4 Compositional Changes during Fruit Maturation and Ripening 494
Organic acids 494
Soluble sugars 495
Structural polysaccharides 496
Pigments and colour 498
Phenolic compounds 502
Flavour (taste, aroma) 504
14.5 Transpiration and Water Loss 507
14.6 Physical Damage and Physiological Disorders 507
Chilling injury (CI) 507
Heat injury 508
14.7 Modified Atmospheres (MA) and Controlled Atmospheres (CA) 508
Injuries associated with MA and CA 510
Modified atmosphere packaging (MAP) 511
Semipermeable coatings 513
Insecticidal CA 514
14.8 Manipulation of Mango Postharvest Physiology by Molecular Biology 515
14.9 Conclusions 516

14.1 Introduction
Successful postharvest handling of mangoes requires knowledge of the post-
harvest physiology of the fruit and how the fruit physiology determines the best
handling practices to maintain and develop high fruit quality. For example, mango,
like banana, tomato and avocado, is a climacteric fruit, which means
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
484 (ed. R.E. Litz)
Postharvest Physiology 485

that it may be picked when mature but before ripening has commenced, and
subsequently ripened postharvest. As mango fruit mature on the tree and begin to
ripen, eating quality improves, but potential marketable life de-creases due to the
difficulty of controlling the ripening changes once they have been initiated,
increased bruising susceptibility and increased decay. Susceptible mango cultivars
tend to develop more internal breakdown (jelly seed, soft nose and stem-end
cavity) the longer that harvesting is delayed (Raymond et al., 1998; see Galán
Saúco, Chapter 9, this volume). As a tropical species, mangoes are subject to
chilling injury (CI), which limits the use of refrigeration to maintain postharvest
quality. Mangoes are also subject to other physiological disorders, physical damage
and decay, the symptoms of which may make the fruit unmarketable (Yahia et al.,
2006a).
Mangoes harvested at a mature but unripe stage of development (‘mature-
green’) can be stored in the unripe state as long as the initiation of ethylene
production and hence ripening is avoided. The initiation of ripening can be avoided
by prompt cooling and storage at a low temperature at which ripen-ing does not
occur or, more effectively, by changing the composition of the storage atmosphere
so that the oxygen (O2) level is reduced and carbon diox-ide (CO2) level is raised.
This latter approach is called either modified atmo-sphere (MA) or controlled
atmosphere (CA) storage, depending on the degree of control. These technologies
slow fruit metabolism and specifically inhibit the initiation of ethylene production.
With MA or CA transport or storage, mangoes can typically be maintained in a
firm, green condition for several days longer than can be achieved with normal
refrigerated air storage. How-ever, there are limits to the levels of O2 and CO2 that
can be tolerated by mangoes and these limits are affected by several factors,
including cultivar, maturity or ripeness stage, storage temperature and storage time
(Yahia, 1998).

Mango postharvest physiology and technology have been described in


previous reports, book chapters and reviews (Subramanyam et al., 1975; Lak-
shminarayana, 1980; Ledger, 1986; Peacock, 1986; Lizada, 1991; Coates and
Johnson, 1993; Johnson and Coates, 1993; Lizada, 1993; Heather, 1994; Jacobi et
al., 1994; Johnson et al., 1997; Mitra and Baldwin, 1997; Tharanathan et al.,
2006).

14.2 Contribution of Mango Fruit to Human Nutrition and Health


Consumers are becoming aware of the nutritional and health benefits of fresh fruits
and vegetables. Mango fruit are a rich source of vitamin C (Table 14.1), although
the content decreases during ripening (Thomas, 1975; Vinci et al., 1995). ‘Raspuri’
mango is rich in vitamin C (300 mg/100 g fresh fruit) during the early stages of
development, but the concentration is less (39.1–69.5 mg/100 g) at maturity
(Siddappa and Bhatia, 1954). The content of vitamin C was between 13 and 178
mg/100 g in the ripe fruit of 50 cultivars surveyed by Singh (1960). The vitamin C
content in fully grown mango fruit of culti-vars in Puerto Rico ranged between 6
and 63 mg/100 g (Iguina de George
486 J.K. Brecht and E.M. Yahia

Table 14.1. Composition of the edible portion of mango fruit (Source: USDA/ARS,
2007).

Nutrient Unit Value per 100 g edible portion

Water g 81.71
Energy kcal 65
Energy kJ 272
Protein g 0.51
Total lipid (fat) g 0.27
Ash g 0.50
Carbohydrate, by difference g 17.00
Fibre, total dietary g 1.8
Sugars, total g 14.80
Minerals
Calcium mg 10
Iron mg 0.13
Magnesium mg 9
Phosphorus mg 11
Potassium mg 156
Sodium mg 2
Zinc mg 0.04
Copper mg 0.110
Manganese mg 0.027
Selenium Pg 0.6
Vitamins
Vitamin C (total ascorbic acid) mg 27.7
Thiamine mg 0.058
Riboflavin mg 0.057
Niacin mg 0.584
Pantothenic acid mg 0.160
Vitamin B6 mg 0.134
Folate, total Pg 14
Folic acid Pg 0
Folate, food Pg 14
Vitamin B12 Pg 0.00
Vitamin A IU 765
Retinol Pg 0
Vitamin E (D-tocopherol) mg 1.12
Vitamin K (phylloquinone) Pg 4.2
Lipids
Fatty acids, total saturated g 0.066
4:0 g 0.000
6:0 g 0.000
8:0 g 0.000
10:0 g 0.000
12:0 g 0.001
14:0 g 0.009
16:0 g 0.052
18:0 g 0.003
Fatty acids, total monounsaturated g 0.101

(Continued)
Postharvest Physiology 487

Table 14.1. Continued

Nutrient Unit Value per 100 g edible portion

16:1 undifferentiated g 0.048


18:1 undifferentiated g 0.054
20:1 g 0.000
22:1 undifferentiated g 0.000
Fatty acids, total polyunsaturated g 0.051
18:2 undifferentiated g 0.014
18:3 undifferentiated g 0.037
18:4 g 0.000
20:4 undifferentiated g 0.000
20:5 n-3 g 0.000
22:5 n-3 g 0.000
22:6 n-3 g 0.000
Cholesterol g 0
Amino acids
Tryptophan g 0.008
Threonine g 0.019
Isoleucine g 0.018
Leucine g 0.031
Lysine g 0.041
Methionine g 0.005
Phenylalanine g 0.017
Tyrosine g 0.010
Valine g 0.026
Arginine g 0.019
Histidine g 0.012
Alanine g 0.051
Aspartic acid g 0.042
Glutamic acid g 0.060
Glycine g 0.021
Proline g 0.018
Serine g 0.022
Other
Ethanol g 0.0
Caffeine mg 0
Theobromine mg 0
E-Carotene Pg 445
D-Carotene Pg 17
E-Cryptoxanthin Pg 11
Lycopene Pg 0
Lutein + zeaxanthin Pg 0

et al., 1969). Vitamin C content was 105.2, 65.7 and 17.3 mg/100 g in ‘Langra’,
‘Ashwini’ and ‘Fazli’ mangoes, respectively (Gofur et al., 1994), and decreased
rapidly 5–7 weeks after fruit set, and when ripe fruit were stored at room
temperature. Vitamin B1 (thiamine) in two mango cultivars was 35–60 Pg/100 g,
488 J.K. Brecht and E.M. Yahia

and vitamin B2 (riboflavin) in three cultivars was 45–55 Pg/100 g (Stahl, 1935).
Thiamine content of four Philippine cultivars was 57–600 Pg/100 g, and riboflavin
content of three cultivars was 37–730 Pg/100 g (Quinones et al., 1944). Folic acid
in green mangoes was 36 mg/100 g (Gosh, 1960).
The mango fruit is a rich source of carotenoids, some of which function as
provitamin A: E-carotene (all-trans), E-cryptoxthanin (all-trans and cis),
zeaxanthin (all-trans), luteoxanthin isomers, violaxanthin (all-trans and cis) and
neoxanthin (all-trans and cis) (Mercadante et al., 1997; Yahia et al., 2006b;
Ornelas-Paz et al., 2007, 2008). Total carotenoid content rose from 12.3 to 38.0
Pg/g in ‘Keitt’ and from 17.0 to 51.2 Pg/g in ‘Tommy Atkins’ from the mature-
green to the ripe stage (Mercadante and Rodriguez-Amaya, 1998), and ripen-ing
alterations occurred principally in the major carotenoids, violaxantin and E-
carotene. With ‘Keitt’, all-trans-E-carotene, all-trans-violaxanthin and 9-cis-
violaxanthin increased from 1.7, 5.4 and 1.7 Pg/g, respectively, in the mature-
green fruit to 6.7, 18.0 and 7.2 Pg/g in the ripe fruit (Mercadante and Rodriguez-
Amaya, 1998). In ‘Tommy Atkins’ these carotenoids increased from 2.0, 6.9 and
3.3 Pg/g to 5.8, 22.4 and 14.5 Pg/g, respectively, during ripening. Geographic
effects were reported to be substantial (Mercadante and Rodriguez-Amaya, 1998).
Some of the cis and trans isomers of provitamin A reported in ‘Haden’ and
‘Tommy Atkins’ mangoes include 13-cis-E-carotene (trace amounts), trans-E-
carotene (12.5–15.5 Pg/g) and trans-D-cryptoxanthin (0.3–0.4 Pg/g) (Godoy and
Rodriguez-Amaya, 1994). In processed mango juice, violaxanthin was not
detected, auroxanthin appeared at an appre-ciable level, and E-carotene was the
principal carotenoid (Mercadante and Rodriguez-Amaya, 1998). The major
carotenoid in ‘Bourbon’, ‘Haden’, ‘Extreme’, ‘Golden’ and ‘Tommy Atkins’
mangoes is E-carotene (48–84% of the total), while epoxycarotenoids
(violaxanthin, luteoxanthin and mutatox-anthin) constitute 13–49% of the total
(Godoy and Rodriguez-Amaya, 1989). Mean vitamin A in these mangoes (retinol
equivalents/100 g) ranges from 115.3 (‘Haden’) to 430.5 (‘Extreme’).

Children in Senegal with normal cytology had higher serum retinol and E-
carotene levels than those with abnormal cytology after massive oral doses of
vitamin A and consumption of mangoes (Carlier et al., 1992). Mango retinol is
highly bioavailable (82% efficiency) by estimating vitamin A and carotene reserves
in the liver and plasma of rats (Yuyama et al., 1991). During mango fruit ripening,
vitamin A increases – ripe mangoes are tenfold richer in caro-tene than partially
ripe fruit, while unripe green mangoes contain only trace amounts (Modi and
Reddy, 1967). Mevalonic acid, a precursor of carotenoids, increases progressively
during mango ripening (Modi and Reddy, 1967). Vitamin A equivalents in 100 g of
mango fruit are 1000 to 6000 IU (Singh, 1960). The E-carotene content of the fruit
of 30 mango cultivars in Puerto Rico ranged from 400 to 800 IU/100 g fresh fruit
(Iguina de George et al., 1969). The development of E-carotene in mangoes held at
16–21C was lower than that at 20–28C (Vazquez-Salinas and Lakshminarayana,
1985). Jungalwala and Cama (1963) identified 16 different carotenoids in
‘Alphonso’ mangoes, and E-carotene accounted for 60% of the total. Of the
oxycarotenoids, luteoxan-thin, violaxanthin and cis-violaxanthin were present in
significant amounts.
Postharvest Physiology 489

All the oxycarotenoids were present as E-carotene derivatives, mostly as epoxides


of zeaxanthin. Variation in carotenoid content, as in many other constituents, is due
to several factors, including cultivar, geography, climate, storage/processing
conditions and analytical procedures employed.
Several carotenoids occur in fruit of different mango cultivars (Cano and de
Ancos, 1994; Ben-Amotz and Fishler, 1998; Chen et al., 2004), but only a few of
them occur in significant concentrations (Ornelas-Paz et al., 2007). Mercadante et
al. (1997) quantified several carotenoids in ‘Keitt’ mangoes; the most predominant
ones were all-trans-E-carotene, all-trans-violaxanthin and 9-cis-violaxanthin,
accounting for 27, 38 and 18% of the total carotenoid content, respectively. Similar
findings have been reported for crude extracts from other mango cultivars
(Mercadante and Rodríguez-Amaya, 1998; Pott et al., 2003a, b). Carotenoids are
responsible for the yellow-orange colour of mango mesocarp (Vázquez-Caicedo et
al., 2004). All-trans-E-carotene and the dibutyrates of all-trans-violaxanthin and 9-
cis-violaxanthin are the main car-otenoids in ‘Ataulfo’ and ‘Manila’ mangoes
(Yahia et al., 2006b; Ornelas-Paz et al., 2008; Fig.14.1). The content of these
carotenoids during fruit ripening increased exponentially in ‘Ataulfo’ and
exponentially or in a second order polynomial manner in ‘Manila’, and the highest
correlation coefficients were obtained for the relationships between the internal and
external a* and h colour values and the content of the evaluated carotenoids in
both mango

3.5

All-trans-violaxanthin
3.0 9-cis-Violaxanthin
All-trans-β-carotene
Pulp carotenoid content (mg/100 g)

2.5

2.0

1.5

1.0

0.5

0.0
‘Haden’ ‘Ataulfo’ ‘Tommy ‘Manila’ ‘Criollo’ ‘Kent’ ‘Paraíso’ Atkins’

Cultivar

Fig. 14.1. Content of selected carotenoids in pulp of several mango cultivars.


Data represent the mean of eight individual observations for each cultivar 
standard error (Source: Ornelas-Paz et al., 2007).
490 J.K. Brecht and E.M. Yahia

cultivars (R = 0.81–0.94). Equations to predict the content of the most impor-tant


carotenoids in ‘Manila’ and ‘Ataulfo’ mangoes on the basis of their inter-nal and
external colour values were obtained by Ornelas-Paz et al. (2008).
The content of D-tocopherol is approx. 0.5 mg/100 g in an unidentified
cultivar from Costa Rica (Burns et al., 2003), while the United States Depart-ment
of Agriculture (USDA) Nutrient Database (USDA/ARS, 2007) indicates an D-
tocopherol content of 1.12 mg/100 g. Ornelas-Paz et al. (2007) found that D-
tocopherol is the only detectable tocopherol in seven mango cultivars (Fig. 14.2);
‘Haden’ and ‘Tommy Atkins’ mangoes had the highest amounts (380 and 470
Pg/100 g, respectively), with c.200–250 Pg/100 g in the other cultivars.
Mango fruit are rich in several types of antioxidant phytochemicals, that is
carotenoids and phenolics (Ornelas-Paz et al., 2007; Rocha-Ribeiro et al., 2007).
Botting et al. (1999), showed that mango fruit have antimutagens and the
heterocyclic amine 2-amino-3-methylimidazo[4,5-f]quinoline. Percival et al.
(2006) observed that whole mango juice inhibited cell proliferation in the
leukaemic cell line HL-60 and also inhibited the neoplastic transforma-tion of
BALB/3T3 cells. García-Solís et al. (2008) studied the effect of ‘Ataulfo’ mango
consumption on chemically induced mammary carcinogenesis and plasma
antioxidant capacity in rats treated with N-methyl-N-nitrosourea (MNU). Mango
was administered in the drinking water (0.02–0.06 g/ml) during both short-term
and long-term (LT) periods to rats treated or not with

600

500
α-Tocopherol (μg/100 g)

400

300

200

100

0
‘Haden’ ‘Ataulfo’ ‘Tommy ‘Manila’ ‘Criollo’ ‘Kent’ ‘Paraíso’ Atkins’

Cultivar

Fig. 14.2. The content of D-tocopherol in the pulp of several mango cultivars.
Data represent the mean of eight individual observations for each cultivar 
standard error (Source: Ornelas-Paz et al., 2008).
Postharvest Physiology 491

MNU. Rats treated with MNU showed no differences in mammary carcino-genesis


or in plasma antioxidant capacity measured by both ferric reducing/ antioxidant
power (FRAP) and total oxyradical scavenging capacity assays. However, in
animals not treated with MNU, but with an LT intake of mango, the plasma
antioxidant capacity as measured by the FRAP assay tended to increase in a dose-
dependent manner. This suggests that mango consump-tion by healthy subjects
may increase antioxidants in plasma.

14.3 Mango Ripening Physiology


Ripening is part of the natural senescence of mango fruit. It is an irreversible
process that contributes to organelle disruption and changes in chemical con-
stituents, flavour and texture. While ripening improves the eating quality of mango
fruit, the postharvest life of the fruit is reduced. Natural senescence, and thus
ripening, is aggravated and promoted by ethylene, mechanical injury and high
temperature. This process can be delayed by lower tempera-ture, elimination of
mechanical damage and reducing ethylene production (Yahia et al., 2006a).
Ripening of mango is inhibited while fruit are attached to the tree, and respiration
and ripening are stimulated upon detachment (Lakshminarayana, 1973). Burg and
Burg (1962) reported that ethylene levels in the tissues of mature-green, attached
mango fruit were relatively high (1.87 Pl/l) and suggested that ethylene was
ineffective for promoting ripen-ing due to a ripening inhibitor supplied by the tree.

Changes associated with mango fruit ripening include: (i) flesh colour from
greenish yellow to yellow to orange in all cultivars (Plate 80a); (ii) skin colour
from green to yellow in some cultivars (Plate 80b); (iii) chlorophyll decreases and
carotenoid content increases; (iv) flesh firmness decreases and juiciness increases;
(v) starch is converted into sugars; (vi) total soluble solids (TSS) content increases;
(vii) titratable acidity decreases; (viii) characteristic aroma volatiles increase; (ix)
CO2 production rate increases from 40–50 to 160–200 mg/kg/h at 20C; and (x)
ethylene production rate increases from 0.1–0.2 to 1–3 Pl/kg/h at 20C. Gowda and
Huddar (2000) found the changes in eight mango selections during ripening
included reductions in fruit weight, volume, length, thickness, firmness, pulp
content, pulp:peel ratio, starch and vitamin C, and increases in TSS, pH, total
sugars, sugar:acid ratio, pulp carotenoid content and peel colour.

Climacteric behaviour

Mango is a climacteric fruit, exhibiting a climacteric pattern of respiration and an


increase in ethylene production during ripening (Cua and Lizada, 1990; Reddy and
Srivastava, 1999; Lalel et al., 2003; Fig. 14.3). The initiation of ethylene
production within the fruit triggers and coordinates the changes that occur during
ripening. These changes include colour changes in the peel and flesh, softening of
the flesh, and development of sweet flavour and
492 J.K. Brecht and E.M. Yahia

6 50
Hard green
5 Sprung green
40

Ethylene (mmol/kg/h)
Half ripe
CO2 (mmol/kg/h)

4 Ripe
30
3
20
2

1 Hard green Sprung green 10


Half ripe Ripe
0 0
0 1 2 3 4 5 6 7 8 9 10 Ripening 0 1 2 3 4 5 6 7 8 9 10
period (days) Ripening period (days)

Fig. 14.3. The climacteric pattern of respiration and ethylene production during mango
fruit ripening (Source: Lalel et al., 2003).

aroma. Mangoes can be ripened after harvest when picked at physiological


maturity (mature-green), when they are fully sized, but before ripening has been
initiated. Maturity indices are chosen to predict fruit quality potential and
postharvest behaviour (Peacock et al., 1986; Medlicott et al., 1988). After harvest,
the fruit is then cooled and isolated from possible sources of ethyl-ene (ripening
fruit, engine exhaust, smoke, etc.) during storage or shipping. This is the primary
strategy used to control ripening and thus extend shelf life. Respiration patterns and
ripening behaviour vary among cultivars, with different climatic conditions and
growing locations (Krishnamurthy and Subramanyam, 1970). Respiration is very
high after fruit set and then declines and is maintained at a low rate until fruit
ripening begins.
The rise in respiration and ethylene production during the climacteric is related
to fruit ripening. The respiratory peak in ‘Alphonso’ mangoes har-vested mature-
green occurs 5 days after harvest, and the fruit ripens within 7 or 8 days
(Karmarkar and Joshi, 1941), while in ‘Kent’ and ‘Haden’ mangoes the peak
occurs on days 9 and 11, respectively (Burg and Burg, 1962), and in ‘Pairi’
mangoes on day 9 (Krishnamurthy and Subramanyam, 1970). These dif-ferences
are normal due to differences in location, climatic conditions, orchard and tree
conditions, and postharvest temperature. The rise in the climacteric respiration in
‘Dashehari’, ‘Amrapali’ and ‘Rataul’ mangoes coincides with the highest level of
sucrose and polygalacturonase (PG; EC 3.2.1.15) activity in ripening fruit (Kalra
and Tandon, 1983). Respiration and ethylene production are excellent maturity
indices, but require considerable expense to measure.
The expression of alternative oxidase (Aox) and uncoupling proteins (Ucp) has
been investigated during mango ripening and compared with the expression of
peroxisomal thiolase (EC 2.3.1.16), a ripening marker in mango (Considine et al.,
2001). The multigene family for Aox in mango is expressed differentially during
mango fruit ripening. Abundance of Aox message and protein peaks at the ripe
stage, while expression of the single gene for the Ucp peaks at the turning or half-
ripe stage, and the protein abundance peaks at the ripe stage. Proteins of the
cytochrome chain peak at the mature-green
Postharvest Physiology 493

stage, suggesting that increases in cytochrome chain components are impor-tant for
facilitating the climacteric burst of respiration and that Aox and Ucp are important
in postclimacteric senescence processes (Considine et al., 2001). Because both
message and protein for the Aox and Ucp increase in a similar pattern, their
expression is not controlled in a reciprocal manner but may be active
simultaneously.
Fruit slicing affects respiration rate (Allong et al., 2001). Slicing of mature-
green ‘Julia’ and ‘Graham’ mangoes increased respiration rate immediately after
cutting, but it decreased significantly within the first 12 h of storage at 5 or 10C,
yet still remained at levels above that of the intact fruit throughout the storage
period. The effect of slicing on half-ripe and firm-ripe fruit is an initial increase in
respiration followed by a decline to levels of the intact fruit.

Ethylene production and responses

Mangoes have a moderate ethylene production peak of 1–3 Pl/kg/h during ripening
at 20C. Ethylene, applied directly or as ethrel, induces faster and more uniform
fruit softening (Lakshminarayana, 1973; Barmore, 1974; Lak-shminarayana et al.,
1974; Sornsrivichai and Waru-Aswapti, 1989). Ethylene treatment can be prior to
shipping (Barmore and Mitchell, 1975). There is disagreement regarding the effect
of ethylene treatment on quality (Chaplin, 1988), and this may be related to
maturity when treated. Treatment of imma-ture fruit leads to softening, but the fruit
have poor flavour.
Mango fruit ripening is accompanied by increased ethylene production, which
coordinates the ripening process. Mango expresses an autocatalytic increase in
ethylene production during ripening (Mattoo and Modi, 1969b). Ethylene
production starts before full ripeness is reached (Burg and Burg, 1962; Cua and
Lizada, 1990). Ethylene production in unripe mango fruit is very low (<0.1
Pl/kg/h) (Burdon et al., 1996). Ethylene production decreases as the fruit matures,
is then undetectable for a time and reappears upon ini-tiation of ripening (Akamine
and Goo, 1973). ‘Kent’ and ‘Haden’ mango fruit have internal ethylene
concentrations of c.0.01 Pl/l during the preclimacteric phase, increasing to c.0.08
Pl/l at the initiation of the climacteric, and up to 3.0 Pl/l at the climacteric peak.
Burg and Burg (1962) reported that ethylene production rises when or before CO 2
production rises in ripening mangoes, while Biale and Young (1981) included
mangoes among fruits in which eth-ylene rises after CO2 production rises.

Only a small concentration of exogenous ethylene (0.005 Pl/l) is needed to


initiate mango ripening (Wills et al., 2001). The small amount of ethylene in the
fruit at harvest is sufficient to initiate ripening (Burg and Burg, 1962). While fruit
of ‘Amrapali’ and ‘Deshehari’ mangoes produce a measurable amount of ethylene
during ripening (Reddy and Srivastava, 1999), ethylene production does not follow
a climacteric pattern and two ethylene peaks (at the mature-green and full-ripe
stages) were recorded. This is probably due to the way that ethylene was measured
in the different fruit, and the lack of control exerted on maturity stages of fruit. In
‘Carabao’ mangoes, the peak of
494 J.K. Brecht and E.M. Yahia

ethylene production occurs 110 days after flower initiation, and declines as fruit
approached full maturity (Cua and Lizada, 1990). The content of 1-
aminocyclopropane-1-carboxylic acid (ACC), the immediate precursor of ethylene,
increases in different tissues (peel, outer and inner mesocarp) dur-ing ripening in
both cultivars, ‘Amrapali’ and ‘Deshehari’, while ACC oxi-dase (ACO; EC
1.14.17.4), which catalyse the conversion of ACC to ethylene and ethylene
production, decline (Reddy and Srivastava, 1999). Fruit peel has the highest levels
of ethylene and ACO and less ACC accumulation than the outer and inner
mesocarp at the mature-green stage. The inner mesocarp has less ACO activity and
high ACC accumulation during the ripening pro-cess compared to peel; levels in
the outer mesocarp are intermediate between those in the peel and inner mesocarp.
Changes in the ability to convert ACC to ethylene in the peel are not related to
changes in ripening parameters in the fruit pulp (Lederman et al., 1997). Mango
seed also produces ethylene (Reddy and Srivastava, 1999). Fruit slicing has no
measurable effect on ethyl-ene production in ‘Julia’ and ‘Graham’ mangoes
(Allong et al., 2001).
Treatment of mango fruit with acetaldehyde or ethanol (0.1, 0.5 or 1% ethanol
or acetaldehyde vapour) has concentration-dependent inhibitory effects on ethylene
production (Burdon et al., 1996). Application of ACC to acetaldehyde- or ethanol-
treated fruit discs indicates that acetaldehyde can completely eliminate increased
ACO activity, whereas ethanol cannot (Bur-don et al., 1996). Accordingly, Burdon
et al. (1996) suggested that acetalde-hyde can either inhibit ACO activity directly
or prevent the increase in the enzyme, thereby providing a possible mechanism for
retarding fruit ripening.

14.4 Compositional Changes during Fruit Maturation and Ripening

Several important metabolic changes occur during the maturation and ripen-ing of
mangoes, and some of those are useful as maturity indices (Ketsa et al., 1991). The
ripening changes are irreversible senescence processes that are related to
degradation of organelles or changes in chemical constituents, and thus relate to the
quality and postharvest life of the fruit. Natural senescence is aggravated and
promoted by ethylene, mechanical injury and high tem-perature, and can be
delayed by low temperature, elimination of mechanical damage and reduction of
ethylene production.

Organic acids

Organic acids are important for respiratory activity and as flavour constitu-ents.
During maturation and ripening, mango fruit experience a substantial loss of
organic acids. The predominant acids in mature mango fruit are citric, succinic,
malic and tartaric acids; citric acid has the highest concentration and tartaric acid
the lowest (Shashirekha and Patwardhan, 1976; Sarker and Muhsi, 1981; Medlicott
and Thompson, 1985). Citric acid content increases steadily during fruit
development in ‘Irwin’, reaching a maximum at the beginning of the endocarp-
hardening period, and then decreases steadily
Postharvest Physiology 495

(Ito et al., 1997). In ‘Keitt’ the predominant organic acids are citric and malic
acids, but tartaric, oxalic, ascorbic, and D-ketoglutaric acids also are present, and
the initial loss in acidity is due to a substantial loss in citric acid and a small loss in
malic acid (Medlicott and Thompson, 1985). In ‘Badami’ man-goes, citric acid is
the major organic acid, but malic and succinic acids are also present (Shashirekha
and Patwardhan, 1976). In ‘Fazli’ mangoes, oxalic, citric, malic, pyruvic and
succinic acids have been detected and tartaric acid has been detected in ‘Zardalu’
mangoes (Kumar et al., 1993). In general, citric and succinic acids decrease during
ripening while malic acid shows different changes with different cultivars (Lizada,
1993).
Mango fruit contain organic acids involved in tricarboxylic acid cycle
reactions (i.e. oxalic, succinic, pyruvic, oxaloacetic and D-ketoglutaric acids). In
‘Pairi’ mangoes, maximum concentration of D-oxoglutaric and pyruvic acids occur
before the climacteric peak. Aspartic and glutamic acid concentrations increase for
c.3 days after harvest and then decrease as the climacteric maxi-mum is reached
(Krishnamurthy et al., 1971). Malic enzyme (EC 1.1.1.40), which catalyses the
oxidative decarboxylation of L-malic to pyruvic acid, occurs in the three-quarter-
ripe and ripe stages and the activity pattern during ripening is similar in
‘Alphonso’, ‘Banganpalli’, ‘Dasheri’, ‘Fazli’, ‘Langra’ and ‘Suvar-narekha’
(Selvaraj and Kumar, 1994). In ‘Alfonso’ (sic), the levels of malic dehy-drogenase
(EC 1.1.1.37) and succinic dehydrogenase (EC 1.3.5.1) increase with the onset of
ripening; whereas, the level of citrate synthase (EC 2.3.3.1) increases several-fold
during maturation but decreases markedly during ripening (Baqui et al., 1974). The
activity of malic enzyme increases during ripening, reaching a maximum
immediately after the climacteric peak, and then declines (Dubery et al., 1984).
The activity patterns of phosphoenol pyruvate carboxylase (PEPC; EC 4.1.1.49)
and pyruvate decarboxylase (EC 4.1.1.1) during ripening vary among different
cultivars, while malic enzyme activity increases during ripen-ing. PEPC activity is
relatively high in ‘Alphonso’ and ‘Langara’, but low in ‘Dashehari’ and ‘Totapuri’
during ripening (Selvaraj and Kumar, 1994).

Soluble sugars

The increase in soluble sugars is a major change during mango fruit ripening, and
sweetness is the most important compositional change related to mango flavour.
While starch content increases in chloroplasts during mango fruit development, it is
almost completely hydrolysed to simple sugars during ripening (Medlicott et al.,
1986; Selvaraj et al., 1989; Kumar et al., 1994; Ito et al., 1997). In ‘Alphonso’,
starch content is 14% (by weight) at the immature stage and c.0.3% at the ripe
stage. Similarly, starch is almost undetectable in ‘Irwin’ mangoes after ripening,
whereas sucrose increases significantly and fructose increases slightly (Ito et al.,
1997). Starch content decreases slightly during ripening of ‘Haden’, but is
insufficient to account for the observed increase in the level of sucrose (Castrillo et
al., 1992).
Ripe mango contains up to 10–20% total sugars on a fresh weight (FW) basis,
depending on the cultivar and the stage of ripeness. At the beginning
496 J.K. Brecht and E.M. Yahia

of ripening, reducing sugars make up most of the sugar content, while there are
more non-reducing (c.17%) than reducing (3%) sugars in completely ripe fruit.
Sucrose contributes 57% of the total sugar in ripe ‘Keitt’ mangoes, with fructose
and glucose making up 28% and 15%, respectively (Medlicott and Thompson,
1985). Although Krishnamurthy et al. (1971), Lakshminarayana (1973, 1975) and
Shashirekha and Patwardhan (1976) reported a simultane-ous increase of glucose,
fructose and sucrose during ripening, Vazquez-Salinas and Lakshminarayana
(1985) observed a gradual reduction in glucose and fructose and a continuous
increase of sucrose during ripening in ‘Haden’, ‘Irwin’, ‘Kent’ and ‘Keitt’.
Medlicott and Thompson (1985) and Vazquez-Salinas and Lackshminarayana
(1985) identified the main reducing sugar as fruc-tose, while Selvaraj et al. (1989)
reported that glucose is predominant. Con-flicting reports on the relative
concentrations of individual sugars in mango fruit during ripening is cultivar-
dependent and due to different storage and handling conditions (Medlicott and
Thompson, 1985).
Sucrose content increases during ripening as a result of starch hydrolysis from
increased amylase (EC 3.2.1.1) activity (Mattoo and Modi, 1969a; Fuchs et al.,
1980; Tandon and Kalra, 1983). The high activities of sucrose synthase (EC
2.4.1.13) and invertase (EC 3.2.1.26) in the mesocarp during ripening indicate
active sucrose metabolism (Kumar et al., 1994). Hexoses and hexose phosphates
can be formed from pyruvate by gluconeogenesis (Selvaraj and Kumar, 1994). The
activity of glucose-6-phosphatase (EC 3.1.3.9) reportedly increases up to the three-
quarter-ripe stage; whereas, fructose-1,6-diphosphatase (EC 3.1.3.11) activity
increases as the fruit ripens from the three-quarter-ripe to full-ripe stage (Kumar
and Selvaraj, 1990). The glycolytic enzyme hexokinase (6-phosphofructokinase;
EC 2.7.1.11) has maximum activity at the ripe stage, while pyruvate kinase (EC
2.7.1.40) activity increases until the three-quarter-ripe stage and declines at
ripening (Selvaraj and Kumar, 1994). The pattern of activity changes in
hexokinase/phosphofructokinase and pyruvate kinase demonstrates that glycolysis
is activated during mango fruit ripening.
Reducing sugars, mainly fructose, increase slightly during ripening, and
sucrose synthase (EC 2.4.1.13) activity increases approximately ten times during
the phase of rapid sucrose accumulation (Castrillo et al., 1992). This activity
accounts for the maximum rate of sucrose synthesis. The proportion of sucrose
phosphate synthase (EC 2.4.1.14) activity that is sensitive to inhibi-tion by
inorganic phosphate changes during ripening (Castrillo et al., 1992). Maximum
catalytic activity of sucrose synthase is constant throughout the ripening period and
contributes significantly to sucrose metabolism. The activities of neutral and acid
invertases (EC 3.2.1.26) are very low in com-parison with the other enzymes of
sucrose synthesis. Acid invertase activity increases and later decreases during
ripening.

Structural polysaccharides

Pulp firmness is important for the evaluation of fruit maturity potential for
transport and storage, and as a quality characteristic. Fruit softening and cell
Postharvest Physiology 497

wall changes are principal changes associated with fruit ripening. Fruit tex-ture
changes are due to changes in cell walls and pectic substances in the middle
lamella, and these are cultivar-related (Selvaraj and Kumar, 1989). Softening of
mango fruit is characterized by increased solubility of cell wall pectins (Roe and
Bruemmer, 1981; Tandon and Kalra, 1984; Lazan et al., 1986; Nasrijal, 1993). In
general, water-soluble polysaccharides increase during ripening (Lazan et al.,
1986; Brinson et al., 1988), but water- and alkali-soluble pectins decline in ‘Keitt’
mangoes, and ammonium oxalate-soluble pectins increase as the fruit become soft
(Roe and Bruemmer, 1981). There is an over-all loss of galactosyl and
deoxyhexosyl residues during mango fruit ripen-ing, the latter indicating
degradation of the pectin component of the wall (Muda et al., 1995). The loss of
galactose appears to be restricted to the chela-tor soluble fraction of the wall pectin,
while loss of deoxyhexose seems to be more evenly distributed among the pectin.

Pectinesterase (PE; EC 3.1.1.11), which catalyses the deesterification of


methyl groups from acidic pectins, has been detected in ripening mangoes (Tahir
and Malik, 1977; Roe and Bruemmer, 1981; Ali et al., 1990, 1995; Abu-Sarra and
Abu-Goukh, 1992). Physiological maturity in ripened mangoes is associated with
lower PE activity (van Lelyveld and Smith, 1979) and peel has higher PE activity
than pulp (Ashraf et al., 1981). Endo-polygalacturonase (PG; EC 3.2.1.15), which
is responsible for degrading the 1-4-linked galactur-onic acid residues, occurs in
ripening fruit (Lazan et al., 1986, 1993; Abu-Sarra and Abu-Goukh, 1992).
Enzymatic and/or non-enzymatic processes, in addi-tion to PG activity, are
involved in the extensive softening of fruit (Mitcham and McDonald, 1992). Other
cell wall hydrolases can be detected in ripening fruit, including cellulases (EC
3.2.1.4; Lazan et al., 1986; Abu-Sarra and Abu-Goukh, 1992), E-galactosidase (EC
3.2.1.23; Ali et al., 1990, 1995; Lazan et al., 1993), galactanase (EC 3.2.1.145; Ali
et al., 1990) and xylanase (EC 3.2.1.8; Ali et al., 1990).

Ripening in mangoes, as characterized by decreased tissue firmness, is


initiated in inner mesocarp tissue close to the seed, and progresses outwards (Lazan
et al., 1993). Pectin solubilization in inner and outer mesocarp tissues is
comparable, but pectin solubilization begins earlier in the inner than in the outer
mesocarp (Lazan et al., 1993). The outer mesocarp of ‘Keitt’ remains firm longer
than ‘Tommy Atkins’, and the inner is softer than the outer meso-carp at each stage
of ripening in both cultivars (Mitcham and McDonald, 1992). Cell wall neutral
sugars, particularly arabinosyl, rhamnosyl and galac-tosyl residues, decrease with
ripening in both cultivars. ‘Keitt’ has more loosely associated, chelator-soluble
pectin, accumulates more soluble poly-uronides and retains more total pectin at the
ripe stage than ‘Tommy Atkins’. Both cultivars have similar PG activity, which
increases with ripening. The amount and molecular weight (MW) of cell wall
hemicellulose decreases with ripening in both cultivars. Galactose is the only cell
wall neutral sugar to show a significant decrease during ripening of ‘Sensation’
mangoes (Sey-mour et al., 1990). Losses of neutral sugars can be due to hydrolysis
of galac-tans and arabinogalactans by E-galactosidase having galactanase activity.
E-Galactosidase activity shows a parallel increase to tissue softening during
498 J.K. Brecht and E.M. Yahia

ripening. The close correlations between changes in E-galactosidase during rip-


ening, and between changes in E-galactosidase activity with tissue softening, and
increased pectin solubility and degradation suggests that E-galactosidase has an
important role in cell-wall pectin modification and mango fruit soft-ening during
ripening (Ali et al., 1995).
Postharvest treatments, such as refrigeration, packaging, application of fruit
coatings, etc., can retard mango fruit softening and activity of pectinases (Lazan et
al., 1990; Nasrijal, 1993). Calcium (Ca) joins free carboxyl groups resulting from
PE-catalysed hydrolysis of methyl ester bonds to form Ca-bridges between
adjacent pectin molecules. Calcium application to ‘Haden’ mangoes by infiltration
or dipping extends their storage life by 1 week (Zam-brano and Manzano, 1995).
Postharvest vacuum application of Ca to mango has also been tried (Tirmazi and
Wills, 1981; Wills et al., 1988; van Eeden, 1992; Yuen et al., 1993). Vacuum (300
mm Hg) infiltration of 1–4% calcium chloride (CaCl2) into ‘Amrapali’ and
‘Deashehari’ mangoes ripened at 25C inhibits PG activity, while ethylene
treatment (1 Pl/l) markedly increases its activity (Reddy and Srivastava, 1999).
Pressure (115 kPa for 2 min) or vacuum infiltration (32 kPa) with 1–8% CaCl2
delays ripening of ‘Kensington Pride’ mangoes by 12 or 8 days, respectively (Yuen
et al., 1993). Yuen et al. (1993) reported that vacuum infiltration of CaCl 2 causes
some peel injury, which can be reduced by: (i) increasing the temperature of the
fruit flesh or the CaCl2 solution during pressure infiltration; (ii) packaging the fruit
in sealed polyethylene during pressure infiltration; and (iii) packaging the fruit in
sealed polyethylene bags or cling or shrink wraps after CaCl 2 treatment. Cal-cium
chloride infiltration of ‘Keitt’ mangoes reduces ethylene production, respiration
rate and the incidence of storage decay (van Eeden, 1992).

Pigments and colour

Mango skin colour is important for its role in the perception of overall qual-ity
(González-Aguilar et al., 2001) and can be important for determining the
appropriate maturity for harvesting (Cocozza et al., 2004; Jha et al., 2007), pro-
cessing (Mahayothee et al., 2004) and consumption (Cocozza et al., 2004; Jha et
al., 2007). The loss of green colour is an obvious sign of fruit ripening in many
mango cultivars. The development of the optimum skin colour usually defines
mango quality. Some mango cultivars retain green colour in ripe fruit. Depending
on the cultivar, skin colour can change from dark to olive-green; sometimes
reddish, orange-yellow or yellowish hues appear from the base colour. Some
cultivars develop a reddish blush, which has been attrib-uted to anthocyanins.
Colour changes in mango fruit are due to the disap-pearance of chlorophyll and the
appearance of other pigments (Fig. 14.4). Chloroplasts are transformed to
chromoplasts containing yellow or red pig-ments (John et al., 1970;
Lakshminarayana, 1980; Parikh et al., 1990; Lizada, 1993). Well-arranged grana
and osmiophilic globules occur in chloroplasts of cells in the peel of unripe
mangoes (Parikh et al., 1990), and lose integrity during ripening. Osmiophilic
globules appear, indicating the transformation
Postharvest Physiology 499

6 Green Green/yellow Yellow/green Yellow


10

15
5
8
LSD
(P = 0.05)
Anthocyanin
4
μAnthocyanins( g/cm 2)

cm 2)
Carotenoids (μg/cm2)

μ(g/
10 6

Chlorophylls
3
LSD
Carotenoid (P = 0.05)
4
2
5

2
1 LSD
Chlorophyll (P = 0.05)

0 3 6 9 12 15
Storage time (days)

Fig. 14.4. Carotenoid, anthocyanin and chlorophyll concentrations in the peel of


‘Tommy Atkins’ mango during ripening at 22C (Source: Medlicott et al., 1986).
LSD, least significant difference.

of the chloroplast to a chromoplast. In yellow cultivars, carotenoids and


xanthophylls are the predominant pigments. The anthocyanin paenoidin-3-
galactoside occurs in the skin of some cultivars (Proctor and Creasy, 1969). During
fruit ripening, chlorophyll concentration decreases substantially in ‘Keitt’, while
carotenoid concentration increases and anthocyanin decreases gradually in
‘Tommy Atkins’ (Medlicott et al., 1986). In ‘Keitt’, a substantial loss of
chlorophyll in the peel occurs after the fruit begin to soften. Peel colour is not an
adequate maturity index, since the fruit is already soft when the colour change
occurs. ‘Tommy Atkins’ mangoes develop more red and yellow pigmen-tation in
the peel and mesocarp than ‘Keitt’ (Mitcham and McDonald, 1992).
Mango fruit pulp contains high concentrations of carotenoids (up to 9 mg/100
g), causing the development of an intense yellow to orange colour. Mango is a
good source of vitamin A. The pulp carotenoid level is cultivar-dependent. In
‘Alphonso’, 16 fractions of carotenoids have been reported: 50% of those are E-
carotene (Jungalwala and Cama, 1963; John et al., 1970). No qualitative changes in
carotenoid composition have been reported for ‘Keitt’ and ‘Tommy Atkins’
mangoes from mature-green to the ripe stage, although quan-titative changes occur
during ripening (Mercadante and Rodriguez-Amaya,
500 J.K. Brecht and E.M. Yahia

1998). However, John et al. (1970) detected 15, 14 and 17 carotenoids in ‘Bad-
ami’ mangoes at mature-green, partially ripe and fully ripe stages of fruit,
respectively. Variation with respect to pigment types and quantities is due to
cultivar differences, geography and climate, different maturity stages and
treatments after harvest; discrepancies in results are probably due to differ-ent
analytical procedures.
Mango skin colour can be used to estimate the content of all-trans-E-carotene
(Vázquez-Caicedo et al., 2004), the most important provitamin A carotenoid (Wolf,
1984). Ornelas-Paz et al. (2007) demonstrated that the val-ues of external and
internal colour are similar in ‘Manila’ and ‘Ataulfo’ man-goes (non-blushed) in
contrast to blushed cultivars (‘Criollo’, ‘Paraíso’ and ‘Kent’). The carotenoids in
fruit skin of some mango cultivars can be corre-lated with some non-destructive
colour measurements (Table 14.2; Figs. 14.5–14.8 (Ornelas-Paz et al., 2008).

The most abundant carotene of mango is all-trans-E-carotene, while the most


important xanthophylls are violaxanthin and its isomers (Wilberg and Rodriguez-
Amaya 1995; Chen et al., 2004). Mercadante et al. (1997) quantified many
cartenoids of ‘Keitt’ mangoes and concluded that the most predomi-nant
xanthophylls were all-trans-violaxanthin and 9-cis-violaxanthin, account-ing for
38% and 18% of total carotenoid content, respectively, although other xanthophylls
are important in other cultivars (Ben-Amotz and Fishler, 1998; Setiawan et al.,
2001).
Modi and Reddy (1967) reported an increase during mango ripening of the
carotene precursors, mevalonic acid (MVA) and geraniol, with a concomitant
increase in carotene content. The geraniol concentration of unripe ‘Alphonso’

Table 14.2. Correlation coefficients (R) for the relationships between the content of the main
carotenoids in mesocarp and the internal/external colour values in ‘Ataulfo’ and ‘Manila’
mango fruit. The correlation analysis was performed using D = 0.5 (Source: Ornelas-Paz
et al., 2008).

a
Cultivar Colour value All-trans-violaxanthin 9-cis-Violaxanthin All-trans-E-carotene
‘Ataulfo’ a* 0.84/0.90 0.83/0.87 0.90/0.90
b* –0.05/0.41 0.00/0.41 –0.05/0.45
L* –0.75/0.19 –0.75/0.21 –0.80/0.27
C* 0.31/0.71 0.34/0.70 0.33/0.72
h –0.88/–0.89 –0.86/–0.87 –0.94/–0.90
‘Manila’ a* 0.92/0.87 0.93/0.89 0.86/0.81
b* 0.76/0.69 0.75/0.67 0.67/0.54
L* –0.86/0.35 –0.86/0.32 –0.74/0.18
C* 0.81/0.74 0.81/0.73 0.73/0.61
h –0.90/–0.89 –0.92/–0.91 –0.82/–0.82
a Colour was recorded using the Commission Internationale de l’Eclairage (CIE) L*a*b* uniform colour space,
where L* indicates lightness on a scale of 0 (black) to 100 (white), a* indicates chromaticity on a
green (–) to red (+) axis, and b* indicates chromaticity on a blue (–) to yellow (+) axis. Numerical
values of a* and b* were converted into chroma (C) and hue angle (h), which represent colour purity
and the shade of colour, respectively.
content

40 Mesocarp Mesocarp Mesocarp


g/kg

30
)
Pigme

10
(1
0
nt

20
–3

Postharvest Physiology
0 5 10 15 20 25 30 40 45 50 55 60 60 65 70 75 80 85
content

40 Peel Peel Peel


g/kg

30
0 )
Pigme
(1

10
nt

20
–3

0
–5 0 5 10 15 20 25 30 35 40 45 60 62 64 66 68
a* value b* value L* value

Fig. 14.5. Relationships between the content of all-trans-E-carotene (▲), all-trans-violaxanthin (as dibutyrate, ), 9-cis-violaxanthin (as
dibu-tyrate, ) in mesocarp and the a*, b* and L* values, measured in mesocarp or peel of ‘Ataulfo’ mango fruit during ripening. Each
point repre-sents the mean of two independent measurements  the standard error (vertical bars). The continuous line represents an
exponential regression (Source: Ornelas-Paz et al., 2008).

501
502 J.K. Brecht and E.M. Yahia

40
Mesocarp Mesocarp
content
g/kg)

30
1
0
(
m
P

e
n
t
i

20

3

10

0
40 45 50 55 60 65 55 60 65 70 75 80 85

40
Peel Peel
content
g/kg)

30
1
0
(
m
P

e
n
t
i

20

3

10

0
30 35 40 45 55 60 65 70 75 80 85 90 95
C* value h° value

Fig. 14.6. Relationships between the content of all-trans-E-carotene (▲), all-trans-violaxanthin (as
dibutyrate, ), 9-cis-violaxanthin (as dibutyrate, ) in mesocarp and the C* and h values,
measured in mesocarp or peel of ‘Ataulfo’ mango fruit during ripening. Each point represents the
mean of two independent measurements  the standard error (vertical bars). The continuous line
represents an exponential regression (Source: Ornelas-Paz et al., 2008).

mangoes varies from 0.5 to 3.0 Pmol with 0.0 to 0.5 Pmol MVA; in ripe mangoes
the corresponding levels are 5–10 and 1–5 Pmol, respectively. The increase in free
geraniol and MVA indicates that these compounds are dephosphorylated during
ripening. Acid phosphatase (EC 3.1.3.2) may regulate carotenogenesis in ripe
mangoes (Mattoo et al., 1968). Mangoes stored at low temperatures and then
ripened at room temperature fail to synthesize as much carotenoids as fruit held at
room temperature (Krishnamurthy and Subramanyam, 1973; Thomas, 1975). Hot
water treatments increase the colour intensity of the pulp (Medlicott et al., 1986)
and the peel (Esguerra and Lizada, 1990).
‘Tongdum’ mangoes, which ripen without changing colour, have three-fold
more chlorophyll and slightly more E-carotene in the peel and have higher rates of
ethylene production compared with ‘Nam Dok Mai’ mangoes, which change from
green to yellow upon ripening (Ketsa et al., 1999). Activities of chlorophyllase (EC
3.1.1.14) and peroxidase (EC 1.11.1.7) in the peel of ripe ‘Tongdum’ fruit are
about half of that in ‘Nam Dok Mai’ fruit. Changes in the peel of ripe green
mangoes are due to either or both a lower activity of chloro-phyllase or peroxidase
activity and are not a result of low ethylene production.

Phenolic compounds

The phenolic content of mangoes is high early during development, then decreases
and remains fairly steady during ripening (Lakshminarayana et al.,
40
Pigment content
g/kg)

Mesocarp Mesocarp Mesocarp


30
(10

20

3

10

Postharvest Physiology
-5 0 5 10 15 20 25 20 30 40 50 60 50 55 60 65 70 75 80 85
40
Pigmen conten

g/kg)

Peel Peel Peel


30
t
3 (10

20
t

10

0
-10 -5 0 5 10 15 20 20 25 30 35 40 55 60 65 70 75
a* value b* value L* value

Fig. 14.7. Relationships between the content of all-trans-E-carotene (▲), all-trans-violaxanthin (as dibutyrate, ), 9-cis-violaxanthin
(as dibutyrate, ) in mesocarp and the a*, b* and L* values, measured in mesocarp or peel of ‘Manila’ mango fruit during ripening. Each
point represents the mean of two independent measurements  the standard error (vertical bars). The continuous line represents an
exponential or second order polynomial regression (Source: Ornelas-Paz et al., 2008).

503
504 J.K. Brecht and E.M. Yahia

40
Mesocarp Mesocarp
content

30
g/kg)1
0
(
m
P

e
n
t
i

20

3

10

0
20 25 30 35 40 45 50 55 60 65 60 65 70 75 80 85 90 95

40
Peel Peel
content

30
g/kg)1
0
(
m
P

e
n
t
i

20

3

10

0
20 25 30 35 40 45 60 65 70 75 80 85 90 95 100 105 110
C* value h° value

Fig. 14.8. Relationships between the content of all-trans-E-carotene (▲), all-trans-


violaxanthin (as dibutyrate, ), 9-cis-violaxanthin (as dibutyrate, ) in mesocarp and the C*
and h values, measured in mesocarp or peel of ‘Manila’ mango fruit during ripening. Each
point represents the mean of two independent measurements  the standard error (vertical
bars). The continu-ous line represents an exponential or second order polynomial regression
(Source: Ornelas-Paz et al., 2008).

1970). This is associated with loss of astringency (Selvaraj and Kumar, 1989). The
peel of mango fruit has a higher phenolic content than the pulp at all stages of fruit
development (Jain, 1961; Lakshminarayana et al., 1970).
Polyphenol oxidase (PPO; EC 1.14.18.1) catalyses the oxidation of mono-and
diphenols to o-quinones, which polymerize to produce brown pigments. PPO
activity increases slightly from harvest maturity to the half-ripe stage and then
declines in ‘Banganapalli’, ‘Dashehari’, ‘Fazli’ and ‘Langra’ mangoes, and
decreases in ‘Alphonso’, ‘Suvarnarekha’ and ‘Totapuri’ mangoes (Selvaraj and
Kumar, 1989). The PPO isolated from ‘Haden’ mango is active towards the o-
diphenolic compounds, showing higher activity in the presence of catechol,
followed by chlorogenic acid, but not with monophenols (Park et al., 1980).

Flavour (taste, aroma)

Sugar changes are very important for organoleptic attributes in the mango fruit.
Fruit flavour is mostly a balance between the content of sugars and organic acids
(Medlicott and Thompson, 1985) as well as aromatic volatiles. Kapse et al. (1989)
determined that increasing TSS and decreasing acidity
Postharvest Physiology 505

increases flavour ratings of mango fruit. Sucrose is the predominant sugar in ripe
mango fruit (Tandon and Kalra, 1983; Medlicott and Thomson, 1985; Vazquez-
Salinas and Lakshminarayana, 1985). The predominant acid in mango fruit is citric
(Medlicott and Thompson, 1985; Lizada, 1993). Several factors affect sugar and
acid contents in mango, including cultivar (Kapse et al., 1989; Kundu and Ghosh,
1992; Gowda et al., 1994), stage of maturity at harvest (Shashirekha and
Patwardhan, 1976; Morga et al., 1979; Tandon and Kalra, 1983), postharvest
treatments (Kumar et al., 1993) and storage conditions (Vazquez-Salinas and
Lakshminarayana, 1985).
Ripe mangoes contain >300 volatiles (Pino et al., 2005), but not all of them are
odour-active and thus do not contribute significantly to aroma. Studies have
identified the volatiles of mango, but not their aromatic activity. The predominant
volatiles in some cultivars are monoterpenes and sesquiter-penes (MacLeod and De
Troconis, 1982; Engel and Tressl, 1983; Pino et al., 2005), as well as lactones and
some fatty acids (MacLeod and Pieris, 1984; MacLeod and Snyder, 1985; Wilson
et al., 1990). However, there is no indica-tion of the presence of a single flavour
impact component (Engel and Tressl, 1983). Some mango cultivars have a peach-
like flavour that may be related to the presence of lactones, which contribute to the
flavour of peaches (Prunus persica) (Lakshminarayana, 1980; MacLeod et al.,
1988, Wilson et al., 1990). MacLeod et al. (1988) detected four lactones in
‘Kensington Pride’ that are also the major volatiles of peach. Monoterpene
hydrocarbons represent about 49% (w/w) of the total volatiles in ‘Kensington
Pride’, with D-terpinolene being the most abundant (26%) and 16 esters
representing 33% (MacLeod et al., 1988). The esters, together with some of the
lactones, contribute to the flavour of ‘Kensington Pride’ mangoes.

Indian mangoes have a unique flavour, which has been attributed to (Z)-
ocimine (Engel and Tressl, 1983; Lizada, 1993). Pino et al. (1989) detected 83
volatiles in ‘Corazon’, ‘Bizcochuelo’ and ‘Super Haden’ mangoes, and total
volatiles ranged between 39 mg/kg in ‘Bizcochuelo’ to 70 mg/kg in ‘Corazon’. The
identified volatiles include D-cubebene, E-maaliene, ethyl(Z)-9-hexade-canoate,
ethyl(Z)-9,12-octadecanoate, ethyl(Z)(Z)(Z)-6,9,12-octadecanoate, cucarvone, 2-
methylpropane-2-ol, 3-methylepentan-ol, thymol and carvacrol (Pino et al., 1989).
MacLeod and Snyder (1985) listed the volatile components of several mango
cultivars, including ‘Willard’ and ‘Parrot’ from Sri Lanka; levels of D-terpinolene
were similar to ‘Kensington Pride’.
Kostermans and Bompard (1993) considered that lack of fibre was linked to an
absence of aroma and flat taste and smell, but some cultivars such as ‘Kensington
Pride’ are low in fibre and have a distinctive flavour and aroma profile, and a high
level of D-terpinolene (Bartley and Schwede, 1987; MacLeod et al., 1988). Lipid
content of the pulp is correlated with the flavour character-istics of some mango
cultivars (Bandyopadhyay and Gholap, 1973a; Gholap and Bandyopadhyay,
1975b, 1976). The ripening of ‘Alphonso’ mangoes at ambient temperature is
accompanied by a sharp increase in triglyceride con-tent, together with the
development of a strong aroma and flavour (Gholap and Bandyopadhyay, 1975a,
1976), but ripening at 10C results in a bland aroma and flavour (Bandyopadhyay
and Gholap, 1973b). ‘Totapuri’ mangoes,
506 J.K. Brecht and E.M. Yahia

a bland cultivar, showed no change in the development of aroma or in the pulp


lipid content (Gholap and Bandyopadhyay, 1975b). During ripening at ambient
temperature, palmitoleic acid content is higher than that of palmitic acid in
‘Alphonso’, whereas ripening at low temperature does not affect the proportions of
these two fatty acids (Bandyopadhyay and Gholap, 1973b). The relative
proportions of palmitoleic and palmitic acids in ‘Totapuri’ mango pulp are constant
irrespective of the ripening conditions (Gholap and Ban-dyopadhyay, 1975b).
Gholap and Bandyopadhyay (1976, 1980) suggested that the relative contents of
palmitic and palmitoleic acids determine the fla-vour quality of mango fruit.

The absence of lactones having coconut-like odour notes in ‘Totapuri’


mangoes may be significant for differentiating its aroma characteristics from
‘Alphonso’, together with the presence of certain similar and dissimilar com-
ponents (Bandyopadhyay, 1983). The aroma of green mangoes has been attributed
to cis-ocimine in ‘Alphonso’ and E-myrcene in ‘Batali’ mangoes (Gholap and
Bandyopadhyay, 1976; Bandyopadhyay, 1983). Table 14.3 lists characteristic
aromas of ‘Alphonso’ and ‘Totapuri’ mangoes and their possi-ble chemical
identities.
In almost all fruits, aromatic volatiles are produced at later stages of rip-ening
(Yahia, 1994). Tree-ripe ‘Tommy Atkins’ mangoes produce higher lev-els of all
aroma volatiles except hexanal than do mature-green fruit (Bender et al., 2000a).
Both mature-green and tree-ripe mangoes stored in 25 kPa CO2 tend to have lower
terpene (especially p-cymene) and hexanal concentra-tions than those stored in 10
kPa CO2 and air-stored fruit. Acetaldehyde and ethanol levels tend to be higher in
tree-ripe mangoes held in 25 kPa CO2 than in those from 10 kPa CO2 or air
storage, especially at 8C. Inhibition of volatile production by 25 kPa CO 2 is
greater in mature-green than in tree-ripe

Table 14.3. Characteristic aromas in ‘Alphonso’ and ‘Totapuri’ mangoes and their possible
chemical causes (Source: Bandyopadhyay, 1983).

Aroma ‘Alphonso’ ‘Totapuri’

Fruit, estery Acetaldehyde, methyl acetate, Propionaldehyde


ethyl acetate, n-butyl acetate Methyl acetate
Green-mango-like cis-Ocimine E-Myrcene
Camphoraceous Not detected Detected, but not identified
Earthy Caryophyllene-pinene Not detected
Almond-like Benzaldehyde Not detected
Burnt-sugar-like Benzonitrile Not detected
Spicy Not detected D-Terpinene
Sweet, sugar-like Detected, but not identified Not detected
Coconut oil-like D-Caprolactone, D-octalactone, Not detected
D-undecalactone
Postharvest Physiology 507

mangoes, and at 8C compared to 12C for tree-ripe fruit. However, aroma volatile
levels in tree-ripe mangoes from 25 kPa CO2 are equal to or greater than those in
mature-green fruit treatments. Atmospheres that prolong mango shelf life by
slowing ripening processes can allow tree-ripe mangoes to be stored or shipped
without sacrificing their aroma quality.
Quality enhancement has been used to determine properties critical to fla-vour
acceptability of mangoes, and focus group interviews have been conducted to
determine sensory attributes important to the purchase and consumption of
mangoes (Malundo, 1996). Sugars and acids enhance perception of specific flavour
notes in mango, including aromatics (Malundo et al., 2001).

14.5 Transpiration and Water Loss


Water loss lowers fruit weight, resulting in shrivelling, and may further reduce
quality by causing poor colour development and uneven ripening. Water is lost
from mango fruit through stomata, lenticels and other openings. Relative humidity
(RH) inside the fruit is 100% and water is lost when RH in the envi-ronment
surrounding the fruit is <100%. Water loss is also greatly influenced by
temperature. With constant RH and air movement, water loss increases signifi-
cantly with any increase in temperature. Transpiration rate is influenced by cul-
tivar and ripeness stage. It is correlated with skin thickness, morphological
structure, epidermal cells and surface wax coating. For example, waxes usually
develop on the epidermis of fruit in the later stages of development and thus it is
common for fruit harvested early to shrivel faster compared with those har-vested
at a more advanced stage of development (Yahia et al., 2006a).

14.6 Physical Damage and Physiological Disorders


Mangoes are susceptible to physical damage at every step of the postharvest
handling chain (see Johnson and Hofman, Chapter 15, this volume) and
reduction/elimination of mechanical injury is essential to reduce losses in quality
and postharvest life. Mango fruit are susceptible to various physio-logical disorders
that influence fruit quality (see Galán Saúco, Chapter 9, this volume). These
disorders are either induced or inherent, and several of them become apparent
during fruit ripening. Disorders, i.e. chilling injury (CI) and heat injury, may be
induced after harvest. Inherent physiological disorders include ‘spongy stem end’
in ‘Kensington Pride’ (Brown et al., 1981), ‘soft nose’ in Florida mangoes (Young,
1957) and ‘internal breakdown’, ‘spongy tissue’ or ‘soft nose’ in Indian ‘Alphonso’
mangoes (Subramanyam et al., 1971).

Chilling injury (CI)

Low storage temperatures can injure mature-green mangoes if storage exceeds a


day or so at or near 0C to a few weeks at just below 12C. This problem limits the
use of low storage temperature to manage postharvest ripening
508 J.K. Brecht and E.M. Yahia

and seriously affects the ability of handlers to store mangoes or transport them over
long distances, because temperatures that are low enough to delay ripening, decay
and senescence may damage the fruit. The symptoms of CI include greyish, scald-
like discoloration on the skin, followed by pitting, uneven ripening, and poor
flavour, aroma and colour development (Hatton et al., 1965; Medlicott et al.,
1990). The symptoms often are not apparent at the low temperature, but develop
later, when the fruit are brought to warmer temperatures for ripening or are
displayed for sale. Other symptoms in mango fruit held at room temperature for 1–
2 days after low temperature storage were described as discoloured and with pitted
areas on the surface (Srivastava, 1967; Kane, 1977) followed by increased
susceptibility to micro-bial spoilage (Sadasivam et al., 1971; Subramanyam et al.,
1975).
Chilling susceptibility varies with cultivar (Farooqui et al., 1985); ‘Haden’ and
‘Keitt’ are particularly susceptible. ‘Sensation’ developed more skin symptoms
than ‘Sammar Bahisht’ mangoes (Farooqui et al., 1985). While CI has generally
been reported to occur in mango fruit at temperatures below c.10–13C
(Mukherjee, 1958; Akamine, 1963; Hatton et al., 1965; Musa, 1974; Couey, 1986),
some cultivars (i.e. ‘Dashehari’ and ‘Langara’) were reported to be safely stored at
7–8C for up to 25 days (Mann and Singh, 1976). While most cultivars show injury
at <10C if fruit have just reached maturity, toler-ance of CI increases as fruit ripen
(Medlicott et al., 1990; Mohammed and Brecht, 2002). Tolerance of ‘Keitt’ mango
fruit of CI was induced by pre-storage heat treatments (McCollum et al., 1993).

Heat injury

Mango is highly tolerant of heat (Yahia et al., 2000; Jacobi et al., 2001b). Man-
goes that are not stored in refrigerated conditions after harvest may be exposed to
extremely high ambient temperatures in many production areas. This may lead to
heat injury, especially if the fruit are exposed to >30C for >10 days, but injury can
also occur more rapidly at higher temperatures. The heat disinfestation treatments
of mangoes that are required for insect quar-antine security may injure fruit that are
not fully mature (Jacobi and Giles, 1997; Jacobi et al., 2001a).

External symptoms of heat injury include lenticel spotting and skin browning
(‘scald’) with secondary disease development, while internal symptoms include
mesocarp browning, tissue cavitation and ‘starch spots’ (Jacobi and Wong, 1992;
Jacobi and Giles, 1997; Mitcham and McDonald, 1997; Jacobi et al., 2001a, b).
Ripening of heat-injured mangoes may also be inhibited (Jacobi et al., 2001a, b).

14.7 Modified Atmospheres (MA) and Controlled Atmospheres (CA)

Long-term marine shipping in MA and CA has been used for transit from several
countries (Yahia, 1993). Research results are very contradictory due to
Postharvest Physiology 509

the different cultivars and maturity stages of mangoes used, different atmo-spheres
implemented and lack of experimental controls. Optimum condition for prolonged
shipping or storage is reported to be 3–5 kPa O2 plus 5–10 kPa CO2, which can
delay ripening, but the benefits are not very significant. Use of CA and MA would
most likely be beneficial in delaying fruit ripening during marine transport for 2
weeks or more.
Bender et al. (2000b) determined the tolerance of preclimacteric ‘Haden’ and
‘Tommy Atkins’ to reduced O2 levels for storage times in typical marine
shipments. They reported that mangoes can tolerate 3 kPa O2 for 2–3 weeks at 12–
15C and that tolerance of low O2 decreases as mangoes ripen. All low O2
treatments reduced mature-green mango respiration; however, elevated ethanol
production occurred in 2 and 3 kPa O2 storage, with the levels two to threefold
higher in ‘Tommy Atkins’ than in ‘Haden’. ‘Haden’ fruit at the onset of the
climacteric accumulated ethanol in 4 kPa O2 and produced 10–20 times more
ethanol in 2 and 3 kPa O2 than preclimacteric fruit. There were no visible injury
symptoms, but off-flavour developed in mature-green fruit at 2 kPa O2 and in
ripening-initiated fruit at 2 and 3 kPa O2. Ethanol production was not affected by
storage in 25 kPa CO2. Ethylene production was reduced slightly by low O2;
however, ‘Haden’ fruit also showed a residual inhibitory effect on ethylene
production at 2 or 3 kPa O2 storage, while ‘Tommy Atkins’ fruit stored in 2 kPa
O2 produced a burst of ethylene upon transfer to air at 20C. Fruit firmness, total
sugars and starch levels did not differ among treat-ments, but 2, 3 or 4 kPa O2 and
25 kPa CO2 maintained significantly higher acidity than 5 kPa O2 or air. The
epidermal ground colour responded differ-ently to low O2 and high CO2 in the two
cultivars. Only 2 kPa O2 maintained ‘Haden’ colour better than air, while all low
O2 levels maintained ‘Tommy Atkins’ colour better than air. High CO 2 was more
effective than low O2 in maintaining ‘Haden’ colour, but had about the same effect
as low O2 on ‘Tommy Atkins’.
Properly selected atmospheres, which prolong mango shelf life by slow-ing
ripening processes, can allow tree-ripe mangoes to be stored or shipped without
sacrificing their superior aroma. Mature-green and tree-ripe ‘Tommy Atkins’
mangoes were stored for 21 days in air or in a CA (5 kPa O 2 + 10 kPa or 25 kPa
CO2) at 12C (mature-green) and at either 8 or 12C (tree-ripe) (Bender et al.,
2000a). Tree-ripe mangoes produced much higher levels of all aroma volatiles
except hexanal than mature-green fruit after ripening for 2 days. Both mature-green
and tree-ripe mangoes stored in 25 kPa CO2 had lower terpene (especially p-
cymene) and hexanal levels than those stored in 10 kPa CO 2 and air-stored fruit.
Acetaldehyde and ethanol levels were higher in tree-ripe mangoes from 25 kPa
CO2 than in those from 10 kPa CO2 or air storage, especially at 8C. Inhibition of
volatile production by 25 kPa CO2 was greater in mature-green than in tree-ripe
mangoes, and at 8C com-pared to 12C for tree-ripe fruit. Aroma volatile levels in
tree-ripe mangoes from the 25 kPa CO2 treatment equalled or exceeded those in
mature-green fruit treatments.
Mangoes have high tolerance of short-term elevated CO2 atmospheres (Yahia,
1998). Mangoes can tolerate CO2 atmospheres of up to 25 kPa for
510 J.K. Brecht and E.M. Yahia

2 weeks at 12C (Bender et al., 2000b). High (25 kPa) CO2 inhibits ethylene
production, but increases ethanol production. Aroma volatiles are reduced
following 25 kPa CO2 treatment, while 10 kPa CO2, low O2 atmospheres and
storage temperature did not significantly influence production of terpene
hydrocarbons, which are characteristic of Florida-type mangoes. Mature-green
‘Tommy Atkins’ mangoes can be stored for 21 days in CA (5 kPa O 2 + 10 kPa or
25 kPa CO2) at 12C, while tree-ripe fruit can be stored for 21 days in the same
atmospheres at either 8 or 12C (Bender et al., 2000a).
The quality of ‘Keitt’ mangoes was evaluated during storage for 6 days at
20C in an extremely low O2 (LO) CA (approximately 0.3 kPa) before stor-age in
modified atmosphere packaging (MAP) made from three, low-density polyethylene
(LDPE) films with different gas permeability characteristics (González-Aguilar et
al., 1997). Both LO and MA treatments delayed the losses of colour, weight and
firmness. Fruit maintained good appearance with a significant delay of ripening.
Mangoes are very tolerant of LO treat-ment; however, some MAP fruit developed
a fermented taste after 10 and 20 days at 20C. Short duration (6-day) storage of
mangoes in LO did not other-wise have any deleterious effect on fruit quality
during subsequent storage under MA or normal atmosphere. Properly selected
atmospheres, which pro-long mango shelf life by slowing ripening, permit fruit to
be shipped without sacrificing superior aroma.

Beaulieu and Lea (2003) studied ‘Keitt’ and ‘Palmer’ mangoes without heat
treatment to assess volatile and quality changes in stored fresh-cut man-goes
prepared from firm-ripe (FR) and soft-ripe (SR) fruit, and to assess what effect
MAP may have on cut fruit physiology, overall quality and volatile retention or
loss. Subjective appraisals of fresh-cut mangoes based on aroma and cut edge or
tissue damage indicated that most SR cubes are unmarket-able by day 7 at 4C.
Both cultivars stored in MAP at 4C had almost identical O2 consumption, which
is independent of ripeness. The CO2 and O2 concen-trations measured for cubes
stored in passive MAP indicated that the system is inadequate to prevent potential
anaerobic respiration after 7 days storage.

Injuries associated with MA and CA

A 10 kPa CO2 atmosphere alleviates chilling symptoms in ‘Kensington Pride’


fruit, but higher concentrations are injurious; low O2 (5 kPa) has no signifi-cant
effect (O’Hare and Prasad, 1993). Higher concentrations of CO2 (>10 kPa) are
ineffective for alleviating CI at 7C, and tend to cause tissue injury and high levels
of ethanol in the pulp. Injury in ‘Kensington Pride’ caused by higher levels of CO 2
appears to be more severe at lower temperatures (O’Hare and Prasad 1993; Bender
et al., 1994, 1995), which could be a result of either compounding injury (chilling
+ CO2) or reduced sensitivity of ripe mango
to CO2.
‘Rad’ mangoes develop internal browning and off-flavour in atmo-spheres
containing 6 and 8 kPa CO2 (Noomhorm and Tiasuwan, 1995). The presence of
starchy mesocarp in ‘Carabao’ mangoes, which is characteristic
Postharvest Physiology 511

of internal breakdown, increases during storage in MA (Gautam and Lizada, 1984).


Fruit stored for 4–5 days have severe symptoms, including air pockets in the
mesocarp resulting in spongy tissue (Nuevo et al., 1984a, b). Paren-chyma cells of
affected tissues have c.18 starch granules per cell, compared to c.2 starch granules
in healthy adjacent cells. However, no difference in starch granule shape was
detected between the spongy and healthy tissues. The spongy tissue, which usually
occurs in the inner mesocarp near the seed and becomes evident during ripening,
has almost ten times the starch content of healthy tissue in the same fruit. External
symptoms of internal browning due to MA include failure of the peel to develop
colour beyond the half-yellow stage.

‘Carabao’ mangoes stored in polyethylene bags (0.04 mm thickness) for 1 day


at 25–31C had a faint fermented odour that disappeared during sub-sequent
ripening outside the bags (Gautam and Lizada, 1984). The fermented odour
increases with time, and persists throughout ripening when the fruit are stored for
2–5 days. The respiratory quotient of this cultivar ranged from 0.59 at 21 kPa O 2 to
6.03 at 2.4 kPa O2, which indicates a progressively anaer-obic metabolism (Sy and
Mendoza, 1984). CO2 production decreases as O2 decreases from 21 to 3 kPa, but
increases at <3 kPa O2. Fermentative decar-boxylation could explain the odour
(Lakshminarayana and Subramanyam, 1970).
Pronounced decay appears after storage of ‘Rad’ mangoes for 20 days in
atmospheres containing 4–6 kPa O2 with 4–8 kPa CO2 at 13C and 94% RH, and
severe incidence of decay appears after 25 days (Noomhorm and Tiasu-wan, 1995).
Greater incidence of decay (stem-end rot and anthracnose) occurs in ‘Carabao’
mango stored in MA for 2–5 days at 25–31C (Gautam and Lizada, 1984).

Modified atmosphere packaging (MAP)

Modified atmosphere packaging is used to create a beneficial MA around a


packaged product using semipermeable film to restrict the movement of respiratory
gases into and out of the package; at equilibrium, the respiration rate of fruit in
MAP is equal to the diffusion of the respiratory gases through the film. Fruit
wrapped in 0.08 mm thick polyethylene bags, with and with-out perlite-potassium
permanganate (KMnO4) and stored for 3 weeks at 10C before treatment with
ethylene developed normal colour, texture and flavour (Esguerra et al., 1978).
Individually sealed ‘Keitt’ in low-density (LDPE) and high-density (HDPE)
polyethylene films for 4 weeks at 20C exhibited delayed ripening, reduced weight
loss and did not develop any off-flavours (Gonzalez et al., 1990). The LDPE had a
thickness of 0.010 mm and permea-bilities of 700 cm3 O2/m2/h/atm and 0.257 g
H2O/m2/h/atm. The HDPE film had a thickness of 0.020 mm and permeabilities of
800 cm3 O2/m2/h/ atm and 0.166 g H2O/m2/h/atm.
In a study to model MAP for mango, ‘Keitt’ fruit were individually vacuum
packaged in LDPE film (0.0245 mm thick, 25.0 g/m2) and stored at 7C/80–90%
512 J.K. Brecht and E.M. Yahia

RH, 12C/75–85% RH, 17C/70–80% RH, 22C/65–75% RH or 25C/65–75% RH


(Yamashita et al., 1997). After mass transfer had reached steady state, respiration
rates, moisture loss, permeability of peel and film to water vapour, and
composition of atmosphere around the fruit were determined for 33 days. Daily
rates of weight loss increased from 4.1 g/kg of fruit at 7C to 10.9 g/kg at 25C.
Respiration rates also increased with storage temperature for both packaged and
unpackaged mangoes, and were 21, 38 and 43% less in packaged fruit at 12, 17 and
22C, respectively. Permeability of peel was 600-fold greater than that of the
plastic film. The in-package CO2 levels increased and O2 decreased with time;
concentration changes were greatest during the first 10–15 days of storage and
were more marked at the higher tempera-tures. Experimental and calculated values
for CO2 levels differed by 29%, depending on temperature.

‘Tommy Atkins’ mangoes individually sealed in heat-shrinkable films and


stored for 2 weeks at 12.8C and then ripened at 21C had less weight loss, but did
not show differences in firmness, skin colour development, decay development or
time to fruit ripening, and had more off-flavours than unwrapped fruit (Miller et
al., 1983). Polyethylene films used were: Clysar EH-60 film of 0.01 mm nominal
thickness, Clysar EHC-50 copolymer film of 0.013 mm nominal thickness, and
Clysar EHC-100 copolymer film of 0.025
mm nominal thickness. Individual mature fruit of the same cultivar were later
sealed in Clysar EHC-50 copolymer film with 0.013 mm thickness, and Cryovac
D955 with 0.015 mm thickness, and stored at 21C and 85–90% RH (Miller et al.,
1986). The O2 permeabilities of the films were 620 cm3/24 h/ m2/atm and 9833
cm3/24 h/m2/atm, respectively. Water permeability was 1.5 g/24 h/m2 and 2.0 g/24
h/m2 at 23C, respectively. Fruit in MAP had less weight loss, but higher incidence
of decay and off-flavour at soft-ripeness than unsealed fruit. The authors concluded
that there were no practical ben-efits from wrapping this fruit in these films and
storage at 21C or even at lower temperatures. ‘Film wrapping mangoes at various
stages of ripeness after harvest…will not improve the maintenance of mango
quality during storage or ripening.’
‘Keitt’ mangoes were individually sealed in LDPE films and in a heat-
shrinkable copolymer (Cryovac D-955) film with non-sealed mangoes as the
control and stored for up to 5 weeks at 12C, 17C or 22C (Yamashita et al.,
1999). MAP reduced the rate constant of vitamin C degradation at all tem-peratures
and vitamin C content of individually packaged mangoes was less affected by
storage temperature than the control. Values for Q10 (the ratio of CO 2 production
to O2 consumption in respiration) were 1.3 and 1.0 for man-goes wrapped with the
heat-shrinkable copolymer and the LDPE films, respectively, and 2.8 for the non-
sealed control.
The combined effect of hot benomyl (1000 ppm) at 55C for 5 min and film
packaging in 0.01 mm PVC extended the storage life of mature-green ‘Nam Dok
Mai’ mangoes stored at 13C (Sornsrivichai et al., 1992). Fruit qual-ity was not
affected by film packaging after 4 weeks, but fruit showed infe-rior quality after 6
weeks. The inhibition of carotene pigmentation in the peel of this cultivar may be
related to O2 concentration inside the package and not
Postharvest Physiology 513

to CO2 concentration (Yantarasri et al., 1994). At least 16 kPa O2 is essential for


development of peel colour to the marketable stage (greenish).
‘Kensington Pride’ mangoes treated with heated benomyl (0.5 g/l at 51.5C for
5 min) and sealed in polyethylene bags (0.04 mm thickness) for various durations
at 20C, had off-flavour and lacked normal skin colour when ripened, but ripened
satisfactorily in perforated bags (Chaplin et al., 1982). The postharvest life of these
fruit was not consistently longer than the control. The CO 2 concentration in the
bags was >20 kPa while the O2 concen-tration was <5 kPa. The incidence of off-
flavours was reduced by including ethylene-absorbent blocks in the bags. The
authors concluded that ‘mangoes cannot be stored satisfactorily at ambient
temperature by such technique’; however, Stead and Chithambo (1980) reported
that fruit ripening at 20–30C was delayed 5 days by sealing in polyethylene bags
(0.02 mm thickness) with ethylene-absorbent blocks without any abnormal flavour.
‘Tommy Atkins’ and ‘Keitt’ mangoes were individually sealed in shrink-able
Cryovac polyolefin films (0.015 or 0.019 mm thickness), either non-perforated
(MD film) or perforated with eight holes of 1.7 mm diameter/ inch2 (MPY) or
eight holes of 0.4 mm diameter/inch2 (SM60M) (Rodov et al., 1994). After 2–3
weeks at 14C and an additional week at 17C, mangoes packaged in perforated
polyolefin films ripened normally. Optimum results were achieved when the film
with 0.4 mm perforations was combined with increased free volume inside the
package by sealing the fruit within polysty-rene trays. After 3 weeks of storage and
1 week of shelf life, sealed ‘Keitt’ mangoes were inferior to the control; they were
less ripe, but beyond 4 weeks (up to 6 weeks) sealed fruit had better quality scores
because they were less overripe. Sealing did not reduce decay of fruit stored for
long periods.
Non-perforated PVC film packaging of ‘Nam Doc Mai’ mangoes was not
sufficiently permeable for O2 exchange to allow proper ripening (Yantarasri et al.,
1995). Therefore, a ‘perforated MA’ was used in which fruit were wrapped in
polystyrene trays (three fruit/pack) at 20C with perforation area of 0.004 cm2.
Fruit ripened normally with no off-flavours. Colour devel-opment in the peel
required a higher concentration of O2 than the flesh. A film of pore area 0.008
cm2 allowed fruit colour to develop after 3 weeks while a pore area of 0.39 cm2
allowed the fruit to colour within 2 weeks.

Semipermeable coatings

Some fruit coatings can create an internal MA within the fruit due to semi-
permeable restriction of O2 and CO2 movement in and out of the fruit. Bald-win et
al. (1999) tested two types of fruit coatings – polysaccharide-based and carnauba
wax-based – for their effect on external and internal mango fruit atmospheres and
quality factors during simulated commercial storage at 10 or 15C with 90–99%
RH, followed by simulated marketing conditions at 20C and 56% RH. The
coatings exhibited markedly different O2 permeabil-ity characteristics under
laboratory conditions. Polysaccharide coatings were less permeable to respiratory
gases (i.e. O2 and CO2) and more permeable to
514 J.K. Brecht and E.M. Yahia

water vapour compared to carnauba wax. When applied to fruit under simu-lated
commercial conditions, however, the differences between the coatings with regards
to their permeability to respiratory gases were much reduced, most likely due to
high RH during cold storage. Both coatings created a MA within the fruit, reduced
decay and improved appearance by imparting a sub-tle shine; but only the
polysaccharide coating delayed ripening and increased concentrations of flavour
volatiles. The carnauba wax coating significantly reduced water loss compared to
uncoated and polysaccharide-coating treat-ments.

‘Julie’ mangoes treated with 0.75% w/v aqueous solution of Pro-long


semipermeable fruit coating (a mixture of sucrose esters of fatty acids and sodium
salt of carboxy methyl cellulose) and stored at 25C and 85–95% RH exhibited
reduced weight loss, retarded ripening and increased storage life (6 days longer)
without evidence of adverse effects on quality (Dhalla and Hanson, 1988).
Treatment with 1.0% Pro-long could increase ethanol concen-tration in the pulp.
Treatment with Pro-long (0.8–2.4%) also delayed ripening of ‘Haden’ (Carrillo-
López et al., 1996).

Insecticidal CA

Mangoes are very tolerant of insecticidal atmospheres, and thus a potential


commercial application is feasible, especially in combination with other treat-
ments (i.e. heat). ‘Keitt’ mango tolerated as low as 0.2 kPa O 2 and as high as 80
kPa CO2 for up to 5 days without any injury upon ripening, although fer-mentative
odours could be noted while the fruit were under atmosphere stress (Yahia, 1993,
1994, 1995, 1997; Ortega and Yahia, 2000). Other mango cultivars were also
evaluated and were very tolerant of extreme atmospheres (Yahia, 1998).
Storage of ‘Keitt’ mangoes in an insecticidal MA (0.03–0.26 kPa O2, 72–79 kPa
CO2, balance nitrogen (N2)) and CA (0.2 kPa O2, balance N2; or 2 kPa O2
+ 50 kPa CO2, balance N2) for up to 5 days at 20C delayed fruit ripening as
indicated by respiration, flesh firmness and colour development (Yahia et al., 1989;
Yahia, 1993; Yahia and Tiznado, 1993; Yahia and Vazquez, 1993). The activities
of phosphofructokinase, alcohol dehydrogenase (EC 1.1.1.1) and pyruvate
decarboxylase were enhanced but activity of pyruvate kinase, suc-cinate
dehydrogenase and D-ketoglutarate dehydrogenase (EC 1.2.4.2) was unaffected.
Although these atmospheres caused changes in glycolysis and tricarboxylic acid
cycle, there was no indication of injury and the fruit rip-ened normally in air.
Sensory evaluation conducted after fruit ripening showed no off-flavours, and there
were no differences between fruit main-tained in the MA or CA and those
maintained continuously in air. ‘Keitt’ mango is therefore very tolerant of
insecticidal atmospheres, and 5 days exposure is sufficient to control many insects
(Rojas-Villegas et al., 1996).
Storage of ‘Keitt’ and ‘Tommy Atkins’ mangoes for 21 days at 12C in
atmospheres containing 25, 45, 50 or 70 kPa CO2 plus either 3 kPa O2 or air,
induced ethanol production of 0.18–3.84 ml/kg/h after transfer to air at 20C
Postharvest Physiology 515

for 5 days (Bender et al., 1994). Atmospheres containing 50 or 70 kPa CO 2 caused


fruit injury, and resulted in the highest ethanol production rates. Enclosure of
‘Haden’ and ‘Tommy Atkins’ mangoes in sealed 20 l jars with an initial
atmosphere of 90 kPa CO2 in air or 97 kPa N2 + 3 kPa O2 for 24 h prior to storage
delayed their ripening, and no injury was reported (Pesis et al., 1994).

14.8 Manipulation of Mango Postharvest Physiology by


Molecular Biology
Mango fruit quality is compromised when harvest occurs before the fruit are fully
mature since they are unable to achieve the full complement of flavour and aroma
during the postharvest handling period compared with fruit harvested at a fully
mature or ripening-initiated stage of development. As a climacteric fruit, maturity
in mango corresponds to attainment of ripening competence. The presence of
ethylene is required for the progression and com-pletion of mango ripening. Thus,
strategies for prolonging the postharvest life and maintaining postharvest quality of
mango other than disease control are focused on reducing the effects of ethylene.
This situation provides an excellent opportunity to utilize genetic transformation to
improve mango postharvest quality by manipulating the role of ethylene (see Litz
et al., Chap-ter 18, this volume). Cruz-Hernández et al. (1997) transformed ‘Hindi’
mango with mango ACO and ACC synthase (EC 4.4.1.14) in the antisense orienta-
tion. A cDNA that codes for mango ACO was also isolated and characterized by
Zainal et al. (1999). Suppression of mango ethylene biosynthesis should allow
harvesting of advanced maturity fruit that contain high levels of sug-ars and
possess enhanced capacity to produce ripe aroma volatiles after exposure to
exogenous ethylene. Progression of ripening in such fruit can be easily halted at the
most desirable and convenient time by simply removing exogenous ethylene.

A cDNA homologue of the ethylene receptor gene ETR-1, referred to as


METR1, which codes for a polypeptide of 802 amino acids with a predicted 89 kDa
MW has been isolated (Gutierrez-Martínez et al., 2001). Two or more ETR
homologues exist in mango. The level of METR1 mRNA in the mesocarp increases
transiently during wounding. Repression of genes involved in eth-ylene action in
mango fruit should result in ethylene-insensitive fruit that are minimally affected
by exposure to ethylene in the postharvest environment, resulting in better control
of ripening and senescence to maintain mango postharvest quality.

Internal breakdown in mango fruit is essentially a problem of premature


ripening; the longer harvest of susceptible varieties is delayed, the greater the
incidence of internal breakdown. Using molecular approaches to reduce ethylene
production and action in mature fruit could reduce internal break-down or
premature ripening and lead to greatly improved quality. Another approach to
mitigating internal breakdown would be to target genes involved in the
maintenance of cell wall integrity. Vasanthaiah et al. (2006) demonstrated
516 J.K. Brecht and E.M. Yahia

differential expression of several genes in tissue showing internal breakdown


symptoms compared with healthy tissue. They suggested that oxidative stress may
be one of the causes of the disorder.
Sane et al. (2005) have isolated and characterized an ethylene-dependent a-
expansin gene, MiExpA1, which is correlated with softening in mango. Expression
of MiExpA1 increases with the progression of ripening and treat-ment with 1-
methyl cyclopropene inhibits both ripening/softening as well as MiExpA1 transcript
and protein accumulation. Recently, a pectate lyase (EC 4.2.2.2) homologue from
ripening mango (MiPel1) has been cloned (Chour-asia et al., 2006). A progressive
increase in transcript accumulation was observed during ripening but expression
was delayed significantly in fruit in air without exogenous ethylene. The
expression was specific to fruit and was triggered only during ripening. Increased
transcript accumulation during ripening was associated with pectin solubilization.
Pectate lyase may be closely associated with pectin degradation and have an
important role in mango softening.

14.9 Conclusions
Mango fruit have the potential to develop extremely desirable texture, taste and
aroma that make this fruit highly appreciated and desirable. Strategies used to
extend mango shelf life are based on control of ripening, ethylene action and
ethylene production. Therefore, fruit are usually harvested at the mature-green
stage, prior to ripening initiation, and stored and transported at low temperatures at
or near the threshold for induction of chilling injury. These practices result in poor
quality, immature and chill-injured mangoes on the market. Successful handling of
ripening-initiated mangoes is prob-lematic due to the fruit’s short shelf life and the
increased incidence of inter-nal breakdown that accompanies delayed harvests
makes international transport of ripening mangoes almost impossible.
Consequently, the export market for fresh mangoes, which expanded rapidly in the
1990s, has not con-tinued its rapid expansion in recent years.

Future expansion of mango consumption will require an understanding of


mango postharvest physiology in order to overcome the problems of CI, internal
breakdown, and premature and uneven ripening. This may involve increased
transport of tree-ripe mangoes in CA-equipped marine containers or in MAP. It
may involve development of improved procedures for storage and ripening to offer
preconditioned, ripening-initiated, ready-to-eat man-goes to consumers. It may also
involve genetic transformation of mango to manipulate the progression and
uniformity of ripening and softening.

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15 Postharvest Technology
and Quarantine Treatments

G.I. Johnson1 and P.J. Hofman2


1
Horticulture 4 Development, Jamison, Australian Capital Territory, Australia
2
Department of Primary Industries and Fisheries, Nambour, Queensland, Australia

15.1 Introduction 530


15.2 Considerations Influencing Postharvest Requirements 531
Market and consumer research 531
Quality assurance (QA) and Good Agricultural Practice (GAP) 531
15.3 Preharvest Management 535
Maturity 535
Adjusting maturation time 538
Skin colour and lenticel damage 538
Storage life and physiological disorders 540
Pests and diseases 542
Weather conditions 544
15.4 Flavour and Aroma 544
15.5 Harvesting and Transport to the Packhouse 544
Timing 544
Sapburn 545
Harvesting and desapping 546
Transport to the packhouse 548
15.6 Packhouse Measures 548
Delivery inspection and traceability 550
Desapping and washing 550
Disease control 550
Brushing 554
Grading and sizing 554
Grade standards 555
Disinfestation 557
15.7 Preparing Fruit for Market 566
Surface coatings 566
Packaging 567
Inspection 568
Palletizing 568
Precooling 569
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 529
530 G.I. Johnson and P.J. Hofman

Ethylene and ripening 569


15.8 Pre- and Post-shipping Storage 571
Cool storage 571
Controlled and modified atmosphere storage 573
15.9 Transport 575
15.10 Marketing 578
Networks and cooperatives 578
Promotion and consumer education 578
15.11 Conclusions 579

15.1 Introduction
Postharvest handling of mangoes is the last phase (from the tree to mouth) of an
agribusiness venture. To optimize productivity, profitable uses for all grades of
fruit should be sought, and stable employment provided for key skilled staff.
Sustainable land and water management, and compliance with health, safety and
financial obligations to employees are also necessary. Increasingly, Good
Agricultural Practice (GAP) protocols need to be observed (GAP, 2003; FFTC-
GAP, 2007).
This chapter reviews the technology and quarantine treatments that have been
developed for the postharvest handling of mangoes. Peacock (1986), Led-ger
(1986, 1991b) and Opara and Nguyen (1999) have previously reviewed postharvest
handling of mangoes, while several others have reviewed spe-cific aspects of the
topic (Johnson and Coates, 1993; Heather, 1994; Jacobi et al., 2001b; Singh et al.,
2004; Yahia, 2006). Less than 10% of total world mango production is exported.
Export markets for mangoes have expanded because of social changes and rising
demand, increased international air cargo space for some sectors and promotion of
export fruit production in developing countries (Procter and Cropley, 1994;
FAOSTAT, 2008). Expan-sion of production to meet the supply requirements of
export and distant domestic markets has been possible because of successful
integrated man-agement strategies and disinfestation technologies to control
diseases and insects (Johnson and Coates, 1993; Johnson and Heather, 1995;
Ploetz, 2007), and increased land availability due to deforestation and
diversification away from rice. Market development has also been facilitated
through harmoniza-tion of the rules of trade between nations and regions at global
and near global levels (WTO, 2008), agreement on pest and disease risk
management under the International Plant Protection Convention (IPPC), various
bilateral and regional Free Trade Agreements (FTA, 2007), and the global
expansion of supermarkets. Simultaneously, knowledge of and concerns about
exotic pest risks to domestic fruit production, socio-political concerns about
chemical residues on food, environmental management and labour conditions, and
rising production and marketing costs, have impinged upon market access and
stimulated international dialogue and research initiatives which address these
concerns (Buchanan, 1994; Gullino and Kuijpers, 1994; Ploetz, 2003, 2007).
Postharvest Technology 531

15.2 Considerations Influencing Postharvest Requirements


Market and consumer research

Strategies and procedures for horticultural market research have been out-lined by
Minnis (1993, 1994), Hall et al. (2001) and Kitinoja and Kader (2003).
Increasingly, a supply chain approach is being taken (Johnson and Hofman, 2004).
Hofman and Ledger (2006) proposed that the supply chain approach should be
used to guide research and development and that there needs to be a champion in
the supply chain with significant influence and a desire for improving chain status
and performance. Key features are identification of market demand, through-put,
price and profit flows, seasonal fluctuations in availability and demands, supply
competitors (and their commodity statis-tics), importer-buyer requirements and
relationships, options for value-adding, and consumer expectations and sales
promotions. Point-of-sale transaction data from supermarkets and other sales
outlets in target markets can be an invaluable source of intelligence and can be
purchased from marketing infor-mation specialists. Detailed supply chain
assessments can guide options for innovation and improvement (Johnson and
Hofman, 2004).
The volume, grade and quality of product available for export requires analysis
in relation to buyer specifications, retail customer and provedore preferences,
technological and regulatory requirements for supplying the market, and the
production, packaging, cooling and transportation proto-cols/options that are
needed/available or specified under agreed codes of practice (NRI, 2008a). Procter
and Cropley (1994), Mahendra et al. (2002) and the Natural Resources Institute
(NRI) (2008b) provide some perspectives on these issues.

Quality assurance (QA) and Good Agricultural Practice (GAP)

Postharvest handling assures timely delivery of a product that closely matches


buyer specifications and complies with mandatory regulatory require-ments.
Satisfying customers underpins quality assurance (QA) and obser-vance of GAP
(Box 15.1), which aims to produce a product of the desired standard, compliant
with regional or internationally agreed standards in production and handling, and
encourage regular, larger and more frequent purchases, and brand loyalty. As
export markets become increasingly com-petitive, responsive observance of QA
and GAP certification can be vital for maintaining and expanding market niche
(Johnson and Coates, 1991; Askar and Treptow, 1993; Ledger and Premier, 2006).
Piñeiro et al. (2004) provide generic guidelines for the development of quality and
safety guidelines for fresh produce, and Ledger and Premier (2006) provide
guidelines for a Mango Quality Plan.

Components of a postharvest handling system may be developed in


commercial confidence to enhance brand-name reputation and increase mar-ket
share. Increasingly they are also agreed under contractual agreements
532 G.I. Johnson and P.J. Hofman

Box 15.1. What is Good Agricultural Practice (GAP)?


GAP enhances trader-buyer relationships and the reputation of producers and
traders for the suitability of produce and environmental care. A key feature of
GAP is the role of independent certifying authorities (usually private sector) in
accreditation and compliance monitoring, to provide additional assurance to
buyers of certification creditability. GAP can be applied to any farming system
or scale. Key elements include: sustainable practices such as integrated pest
management (IPM), responsible fertilizer use and care for the environment. It
relies on four key principles (Wikipedia, 2007; NRI, 2008b):
• Economic and efficient production of adequate supplies of safe and nutri-
tious foods or other agricultural commodities.
• Compliance with sustainable practices which improve natural resource
availability.
• Assuring the economic and environmental sustainability of the farming
enterprise.
• Meeting social and cultural norms and expectations in production and
marketing.
GAP has gained global prominence because of the trend to making it a ‘com-
pulsory standard’ for exporters to Europe. A private sector initiative, EurepGAP
sets ‘voluntary standards’ for agricultural product certification anywhere. It involves
partnership between farmers and retailers wanting to establish and implement
procedures and standards for certifying application and compliance in GAP. It is
applied before the farm-gate (from planting to market dispatch), and is not nec-
essarily a standard that is displayed to customers. EurepGAP involves an array of
documentation including general regulations, control points, compliance cri-teria
and checklists (EurepGAP, 2007).
In 2007, EurepGAP transformed into GLOBALGAP (GLOBALGAP, 2007)
that could be tailored and adopted anywhere in the world. In parallel with the
Eurep-GAP process, the South-east Asian countries have been working
towards Association of South-east Asian Nations (ASEAN) GAP standards
suited to fruit marketing within ASEAN (ASEAN-GAP, 2007; FFTC-GAP,
2007; Johnson et al., 2008).

with supermarkets or exporters. No regulations govern the pursuit of supe-rior but


legal agribusiness practices that may enhance grower and seller rep-utations or
provide additional advantages. Regulations govern chemical residues, pests,
diseases, product and packaging specifications. The sections that follow need to be
considered in developing the postharvest components of GAP protocols and
dynamic QA systems.

Regulatory restrictions and quarantine treatments


Quarantine treatments disinfest produce of target pests. They are a critical
component of protocols designed to satisfy market-stipulated prohibitions against
pest entry to countries, or regions within countries. Protocols stipulate procedures
for monitoring, detecting, eliminating and handling pest-affected
Postharvest Technology 533

produce. Frequently, a detailed pest risk analysis (PRA) is required, protocol


efficacy against pests of quarantine concern must be demonstrated, appli-cator
integrity scrutinized and market-access requirements and rights ap-proved by
regulatory and agricultural authorities in the importing and exporting country or
region. Increasingly, systems approaches are being applied. These minimize the
need for reliance on quarantine treatments by including PRA, production-based
pest management systems, consideration of establishing pest-free areas or
identifying commodities which are non-hosts of quarantine pests.

Restrictions or limitations on pesticide residues and other contaminants in


marketed produce are also considerations in the choice of quarantine treat-ment
and market access. Pesticide residue monitoring protocols are often required
(Johnson and Heather, 1995; Sharp and Heather, 2002; McMaugh, 2005). By
restricting or preventing market access, quarantine and pesticide residue
restrictions can unintentionally operate as quasi-trade barriers, effec-tively
reducing competition with domestic production of the same or alter-native
products. However, this is decreasing because of better adherence to science-based
decision making as the foundation for quarantine restrictions (WTO, 2008).
Government authority web sites and export agencies can pro-vide information on
quarantine and tariff barriers, pesticide use restrictions, inspection, packaging and
labelling regulations (PPQ, 2007; IPPC, 2008) (see Disinfestation section under
15.6 Packhouse Measures, this chapter). Phyto-sanitary requirements for fresh
mango exports to some markets are shown in Table 15.1. Marketing through
reputable exporters, known to the importing country as suppliers of produce that
comply with regulatory restrictions, can encourage producer and importer
confidence.

Limitations of the product


Mangoes ripen rapidly and have low tolerance of temperatures <10C. Post-harvest
handling procedures for mangoes aim to optimize quality and mini-mize premature
ripening and fruit damage. Precise maintenance of fruit quality and the storage
environment demands inputs at every stage from picking to the consumer (Ledger,
1991b; Milne, 1994; Ledger et al., 2002b).

Supply, logistics and transport


Market development cannot succeed without a reliable supply of suitable or
market-compliant produce. Production seasons are usually short (2–8 weeks),
cultivar appearance and flavour diverse, maturation time variable, and orchards
sometimes small and scattered. Practical solutions to these limita-tions (i.e.
sourcing from a range of ecoclimates with differing maturation dates, or flower-
induction technologies) are critical for industry develop-ment. Competitive air-
freight rates and rapid road transport, combined with cool-chain handling and
atmosphere management can make nearby export markets almost as accessible as
distant domestic markets. Special perishable produce rates may be negotiated or
subsidized by government, especially during industry establishment. Sea freight is
necessary for moving large volumes when air freight is uneconomical. Out-turn
problems often arise,
534 G.I. Johnson and P.J. Hofman

Table 15.1. Typical phytosanitary requirements for mangoes for some countries.
a
Phytosanitary requirements (usually for mangoes from individual
Country countries or regions on a case-by-case basis)

Australia Approved treatment for fruit fly, area free of pulp weevil
(Sternochetus gravis (F.))
Canada No phytosanitary certificate required
China Phytosanitary certificate required. Field management measures for
specific pests of quarantine concern to China plus approved
disinfestation treatment for fruit flies
EU Phytosanitary certificate required
Indonesia Phytosanitary certificate required plus grown in area free of Queensland
and Mediterranean fruit fly
Japan Phytosanitary certificate required plus disinfestation schedule approved
for nominated mango cultivars and fruit fly species and inspection of
approved quantity of fruit (2–5%)
Korea Phytosanitary certificate required, combined with field surveys
Malaysia Must be free of seed weevil on inspection
New Zealand Phytosanitary certificate required plus approved disinfestation schedule
for nominated fruit fly species
Saudi Arabia Phytosanitary certificate required. Require destructive test of 2% of
consignment for seed weevil, or field survey verification of block freedom
Singapore No restrictions
United Arab Phytosanitary certificate required. Require destructive test of 2% of
emirates consignment for seed weevil, or field survey verification of block freedom
USA Phytosanitary certificate required plus disinfestation approved for
nominated mango cultivars and fruit fly species
a
General requirements: Prior approval to import is required to access the market of many countries.
Packhouse and disinfestation treatment/facility inspections may be required by exporting and
importing regulatory authorities. Import permits may cover multiple importations but usually require
renewal every 3–12 months. Phytosanitary certificates must be issued by a government authority. Fruit
must be free of soil and debris and packed in clean, new containers. Timber packaging and pallets will
be subject to additional requirements. Consignments found to contain quarantinable pests will be
rejected, and either re-exported or destroyed.

especially during market development (Snowdon, 1994). Insurance should be


arranged to cover losses. Specialist inputs may be required to identify causes of the
out-turn problems and in the apportionment of liability amongst producer, shipper
and marketer (Snowdon, 1990, 1994).

Personnel
QA systems and GAP protocols must encompass the human component of an
organization or business (Bunt and Piccone, 1994; Rolle, 2006; Sonneveld, 2006;
FFTC-GAP, 2007). Market agents, exporters, farm/packhouse suppli-ers, finance
providers and transport personnel and companies need to be selected and worked
with from the quality perspective. Human resource development, training and
education have been of major significance in the success of many industries.
Postharvest Technology 535

Ledger and Bagshaw (1994) refer to three general styles of management:


(i) defect detection; (ii) defect prevention; and (iii) continuous improvement. They
consider that quality management of horticulture has traditionally been the defect
detection style, but it is now moving to the continuous improve-ment style via the
implementation of QA, total quality management and supply-chain improvement
systems. Key ingredients of successful imple-mentation of quality systems and
supply chains are: (i) unqualified commit-ment by the owner and senior
management of the human, material and other resources needed to introduce and
maintain the system; and (ii) employee understanding and active participation in
the process (Ledger and Bagshaw, 1994).

Profitability and sustainability


Higher freight rates, tariffs and taxes, pesticide-use monitoring and quaran-tine and
security clearance times can affect sales negotiations and profit mar-gins, as can
socio-cultural differences among retail buyer, importer, exporter and producer
during marketing negotiations and exporter and buyer per-ceptions and consumer
expectations of quality. Intended markets, retailers and regional trade fairs should
be visited to make personal contacts and to assess suitability of the market and
retailing facilities, conditions and pros-pects. Ongoing market monitoring is vital,
with regular out-turn inspections by trained personnel, and contingency
arrangements for product regrading at destination if required. Prompt, personal
attention to client concerns or product problems can be essential for continuing
success (Johnson, 1997; Vinning and Young, 2006).

15.3 Preharvest Management


The effects of production practices on fruit quality have been reviewed by Arpaia
(1994), Hofman and Smith (1994), Hofman (1998) and Hewett (2006). The fruit
characteristics influenced by preharvest factors include internal and external
colour, shape, size, sweetness, vitality (the inherent capacity to maintain quality
after harvest) (Hofman et al., 1997b), cleanliness and residue levels, and the
occurrence of pest or disease infestations or biotic/abiotic damage. The main
production factors influencing at-harvest quality apart from genotype include
chemical treatment regimes and orchard hygiene, weather conditions before and at-
harvest, irrigation, pruning systems and tree nutrition.

Maturity

Peacock (1986) considered that fruit maturity referred to its stage of ontog-eny,
with fruit of different maturities being at different stages of ontogeny. Fully mature
mango fruit are strictly those which have produced a fully developed seed and
which have reached their full physiological potential in
536 G.I. Johnson and P.J. Hofman

relation to size increase and dry matter accumulation within the constraints of the
growth environment. When fruit size and dry matter concentration reach a plateau,
climacteric fruit such as mango can undergo ripening, where colour, texture,
flavour and aroma may change (Watada et al., 1984). In these fruit, a sharp rise,
followed by a decline in respiration also accompanies the transformation from not-
ready-to eat (unripe), to edible (ripe), to senescent (overripe). Ripening signals the
completion of seed ontogeny, and encour-ages dispersal of the seed by attracting
vertebrate fructivores (Cipollini and Stiles, 1992). If fruit are not harvested,
maturation and ripening occur on the tree. Ripe fruit fall to the ground, or are
consumed by bats, primates, pha-langers, birds or humans, either on the tree or
after detachment.
Softening and sweetening of fruit flesh and colour changes can occur at any
stage of ontogeny, even in pea-sized fruit (Oosthuyse, 1995). Fruit drop at any
stage of development is preceded by these events, and the likelihood of their
occurrence increases as fruit size and dry matter levels approach their maxima
(Singh et al., 2004). Although the changes constitute some of the components of
ripening, they can only be regarded as such if the fruit have attained physiological
maturity, i.e. ‘the stage of development when a plant or plant part will continue
ontogeny even if detached’ (Watada et al., 1984; Yashoda et al., 2006).

When fruit are removed from the tree several days before the onset of
ripening, they are initially hard and green. The fruit progressively soften, change
colour and develop aroma at a rate determined by cultivar, storage environment
and at-harvest maturity. Ideally, fruit are picked, treated, packed and transported
while hard-green, and arrive at retail markets at some pre-determined stage of
colour development (usually more yellow or red, than green, and ‘sprung’, but still
firm). The rate at which ripening will occur under particular storage conditions
depends upon the stage of ontogeny at harvest. More mature fruit will ripen more
rapidly than less mature fruit.
Accurately estimating when the fruit are ready for harvest is critical to
consistently meet customer expectations. This is called horticultural matu-rity, and
several criteria of horticultural maturity are possible. One is a legal minimum or
buyer-specified standard of maturity, which confirms that the fruit would be
acceptable for consumption or processing when ripe or ready to eat for green-
eating types. Maturity estimation often relies on visual or calendar-based (days
from flowering) assessment or in some cases the appli-cation of a simple test (e.g.
dry matter or flesh colour assessment). In more exacting cases, more accurate
estimates of horticultural maturity may be required to assess product suitability for
more stringent or narrower quality specifications, as may be required for contract
sales or sea-export consign-ments.

In both cases, easy to assess harvest indices relying on visual, chemo-sensory


or fruit-age attributes are needed, and they must correlate with the commercially
relevant fruit characteristics measured in prescribed tests. Pea-cock (1986), Harvey
(1987) and Askar and Treptow (1993) reviewed methods for assessing fruit
maturity. In mango, dry matter, flesh colour, skin colour, fruit shape, Brix, specific
gravity or days from flowering or heat accumulation
Postharvest Technology 537

7
7 Early harvest
Mid-harvest
6
Late harvest
6

Flavour (1–9)
Flavour (1–9)

5
5
4
4
3
r ² = 0.86** 3 r ² = 0.83

2 4 6 8 10 12 1200 1400 1600 1800 2000


Flesh colour at harvest (chart) Heat units

Fig. 15.1. Relation between flesh colour (1 = white; 15 = very yellow) and accumulated
heat units (>10C) with flavour of ripe ‘B74’ mango grown under Australian conditions. **=
Significant to P = 0.01. (Source: Hofman and Marques, unpublished data).

units (e.g. degree-days) have been used (Baker, 1986; Hofman and Ledger, 2006).
In South Africa flesh colour is favoured, while in Australia, skin colour, dry matter
and accumulated heat units are considered as well (Fig. 15.1). Using several
maturity indicators will usually increase the accuracy of pre-dicting the first
acceptable harvest date. Information would be cultivar-specific. In some cultivars
there may be few reliable visible indicators of maturity that allow picking of the
most mature fruit on the tree, particularly when flowering has occurred over many
weeks. In these instances it is very difficult for pickers to spot pick the more
mature fruit in a cost-effective way. Variation in maturity between fruit can also be
influenced by where fruit develop on the tree. In the cooler subtropical areas of the
southern hemi-sphere, fruit on the northern or western side mature more quickly
than fruit on the southern side (Hofman et al., 1995; Oosthuyse, 1995; Hofman et
al., 1998). In these situations, it may be more feasible for growers to map the
maturity of their blocks or orchards to identify groups of trees that have more
mature fruit, and/or identify parts of the tree (e.g. the northern/western side in the
southern hemisphere) that generally hold more mature fruit. Pickers can then be
instructed to pick all the fruit on a specified canopy position and from specified
areas of the orchard in order to harvest more mature fruit. This can be more cost
effective than selectively picking from individual trees. New technologies such as
portable near infrared spectroscopy (NIRS) units may assist in non-destructively
mapping maturity profiles across orchards (Subedi et al., 2007).

The maturity of fruit at harvest is important for determining fruit quality at


out-turn in overseas markets (see 15.8 Pre- and Post-shipping Storage sec-tion, this
chapter). If fruit in a carton or pallet are of uneven maturity, it may be impossible
to find an effective storage regime(s) which will ensure good quality of all fruit on
arrival. One fruit of more advanced maturity in a box or pallet can accelerate the
ripening of other fruit, which can then arrive with
538 G.I. Johnson and P.J. Hofman

disease symptoms and with little or no shelf life. Variable maturity within
treatment lots can also adversely affect product quality after heat disinfesta-tion
(Jacobi et al., 1995).
On-farm record keeping and analysis of date of flowering, seasonal prod-uct
maturity, orchard management schedules, environmental data, transport regimes,
market destinations and out-turn problems may enable some pre-diction of at-
market quality and better selection of the appropriate destina-tion market. Recent
research is focusing on non-destructive methods that could be used for checking
fruit maturity in automated grading systems (Joyce et al., 1993; Subedi et al.,
2007). Improvements in product quality and performance resulting from the
effective use of such systems over several seasons, can provide considerable
competitive advantages when developing buyer relationships or consumer
‘brand’/country-of-origin loyalty.

Adjusting maturation time

Flowering may be hastened or delayed by pruning, flowering inducers or growth


regulators (Davenport and Núñez-Elisea, 1997), to move the fruit-set period into a
different time period, and bring forward or delay the harvest date to coincide with
higher demand and market prices. Disadvantages could include higher at-harvest
temperatures or greater risk of rainfall prior to harvest adversely affecting fruit
quality.
Some cultivars and growing conditions are more favourable for manipu-lating
flowering date than others. In the Philippines, manipulation of ‘Cara-bao’
flowering with potassium nitrate (KNO3) helps spread production and market
supplies year-round (Bondad and Linsagan, 1979). The treatment also increases
‘Kent’ flowering but does not alter timing (Goguey, 1993). By contrast, chemical
manipulation of flowering has not been effective on ‘Kens-ington Pride’ (Barba,
1974).
Evenness in flowering within and between tree/block/farm lots can contribute
to product uniformity and increased customer confidence. Treat-ments that increase
evenness of flowering can have commercial benefit in this regard also.

Skin colour and lenticel damage

Skin colour can affect sales, with markets preferring colour (green, yellow, orange,
red blush) familiar to past purchase experience, known use and cul-tivar
knowledge, or ethnic-group preferences. Fruit position on the tree af-fects red
colour development, since sun exposure is important for anthocyanin development.
Likewise, bagging of fruit can decrease red and green skin colour on ripe fruit
(Hofman et al., 1997a). Nitrogen (N) can increase the pro-portion of the ripe fruit
with green skin (Fig. 15.2) by retaining skin chloro-phyll (McKenzie, 1994;
Nguyen, 2003; Nguyen et al., 2004; Bally, 2007). In cultivars susceptible to green
skin at ripeness, N fertilization rates should be
Postharvest Technology 539

40

30

20
Green colour (%)

10

2
1

0
0 75 150 300 Fol. 0 150 300 450 Fol. 0 150 300 450 Fol.
HG orchard LG1 orchard LG2 orchard
Total N application (g/tree)

Fig. 15.2. Percentage of green skin colour on ripe ‘Kensington Pride’ mango fruit from
trees in three different orchards (coded HG, LG1 and LG2) following application of N (0–
450 g/tree). (Source: H. Nguyen et al., 2004). Fol. = foliar N sprays to a total of 50 g/tree.
Bars represent least significant difference (LSD) at 5%.

balanced between improving yield and reducing quality. Applying N to trees soon
after harvest can minimize these negative effects in the subsequent crop. Additional
N can be applied just before flowering as long as leaf N concentrations are below a
certain level (Whiley and Hofman, 2007). This level may vary with cultivar.
Excessive fruit calcium (Ca) concentrations in mango will also retard green colour
loss during ripening (Wills et al., 1988). Some cultivars are marketed green (e.g.
‘Keow Savoey’), which are consumed mature-green before softening and colour
development occur.
Enhanced prominence or damage to lenticels on the skin can affect visual
appearance (Plate 81). Various terms have been used, including lenticel dam-age,
discoloration and spotting. Symptoms can be caused by darkening of the cells
immediately around the lenticel producing a brown or black spot, or by a red or
green halo around the lenticel with or without the black or brown spot in the centre
(Bezuidenhout et al., 2005; Self et al., 2006). Recent studies suggest that the
discoloration is not primarily caused by loss of cellular func-tion, but rather by the
deposition of phenolic pigments in the cell wall (du Plooy et al., 2006). It is
possible that leakage of precursors (elicitors) from adjacent resin canals into the
cell wall next to the lenticels contributes to pig-ment formation. Lenticel
discoloration may be a stress-related self-defence
540 G.I. Johnson and P.J. Hofman

Table 15.2. Effects of leaf:fruit ratios on quality of ‘Kensington Pride’ mango fruit after ripen-
a
ing at 22C (Source: Simmons et al., 1998).

Disease
Leaf:fruit Fruit Dry Lenticel spotting
ratio mass (g) matter (%) (1–5) Severity (1–5) Incidence (%)
c b c c b
Control 441.4 13.0 3.5 1.2 13.3
d c c b ab
30 363.2 12.0 3.5 1.8 40.0
b b b b ab
60 532.5 13.7 3.9 2.0 43.3
a a a a a
120 696.6 15.1 4.2 2.7 63.3
a
Treatments were applied by girdling individual branches. Control fruit were from non-girdled
branches. Values are means of 30 fruit per treatment. Values with different letters within columns are
significantly different at P < 0.05.

mechanism against foreign particles and infection entering through the len-ticels
(Bezuidenhout et al., 2005; du Plooy et al., 2006).
The severity of lenticel damage is often difficult to control, but strategies for
minimizing damage exist. There are cultivar differences in susceptibility
(Oosthuyse, 1999), possibly related to differences in lenticel, wax and/or cuticle
structure or composition (du Plooy et al., 2004). Strong negative cor-relations have
been found with maximum and minimum temperature and Class A pan
evaporation, and strong positive correlations with maximum relative humidity
(RH) and rain at harvest (Oosthuyse, 1998). These results suggest that cool, humid
and wet conditions around harvest increase the risk of lenticel damage. Increased
damage following excess irrigation during the latter stages of fruit growth
(Simmons, 1998) support the above conclusions. Lenticel damage can also be more
severe in larger fruit obtained from branches with higher leaf:fruit ratios (Table
15.2), possibly because of greater damage to the lenticels during fruit growth
(Simmons et al., 1998).

Storage life and physiological disorders

Physiological disorders include a range of symptoms that affect shelf life and
marketability (Johnson et al., 1996). These generally result in either prema-ture
ripening of parts of the fruit (e.g. soft nose and jelly seed) or tissue break-down
(i.e. stem-end cavity) (Winston, 1986; Mead and Winston, 1991; Whiley, 1999)
and tissue breakdown in ‘Keitt’ (Bally, 2007). These disorders are more severe in
more mature fruit (Young, 1957; Katrodia, 1989; Mead and Winston, 1991) and
are often evident on the tree or after ripening without storage.
Mango disorders are affected by growing conditions (Young, 1957; Young and
Miner, 1961). Production away from the coast, and higher altitude and/ or lower
temperature are associated with lower incidence of spongy tissue (Subramanayam
et al., 1980; Katrodia, 1989), and susceptibility to the disor-der is also affected by
rootstock (Joshi and Roy, 1985). Stem-end cavity ap-pears to be more severe in
wet conditions near harvest (Wainwright and
Postharvest Technology 541

Burbage, 1989; Mead and Winston, 1991). Incidence of spongy tissue has been
reduced by mulches that decrease radiated and reflected field heat (Katrodia and
Sheth, 1989). Severity of watery pulp breakdown in ‘Keitt’ is lower with higher
crop loads from similar size trees (Bally, 2007), possibly because of smaller fruit
size at high crop loads.
Several reports suggest links between low fruit Ca and mango disorders. High
leaf Ca has been related to reduced soft nose (Young and Miner, 1961) and reduced
stem-end cavity (Mead and Winston, 1991). Soil Ca applications can reduce stem-
end cavity incidence (Whiley, 1999), but these responses are not always consistent.
Applications of Ca to ‘Keitt’ from just before flowering onwards did not increase
leaf or fruit Ca during fruit growth or at commer-cial harvest (Bally, 2007).
However, soil characteristics may also affect responses to soil Ca applications. In
the sandy soils typical of Australian orchards, and even in the heavier clay
subtropical soils with low cation exchange capacity, Ca can be rapidly removed
from the top soil profile, resulting in little long-term increase in soil solution Ca
(Hofman and Mullen, 2005; Bally, 2007). Regular (two/week), small applications
are required to consis-tently increase solution Ca (Hofman and Mullen, 2005).
Other factors (e.g. vegetative vigour and water status) can influence fruit Ca uptake
(Hofman and Smith, 1994).

Foliar Ca treatments have produced inconsistent effects, in some cases having


little or no effect on fruit Ca concentrations or quality (Singh et al., 1987; Simmons
et al., 1995), and in other instances reducing internal disor-ders (Chitarra et al.,
2001). Postharvest dips have also had mixed results, with both positive (Wills et
al., 1988; Singh et al., 2000) and nil or very small effects (Joyce et al., 2001)
reported. As a result there is little commercial use of foliar or postharvest Ca
treatments in mango. Future development of more labile Ca formulations may
provide more consistent results.
Other nutrients have also been associated with fruit quality (Hofman and
Smith, 1994). There are relatively few reports of potassium (K) and mag-nesium
(Mg) effects on mango fruit quality. Higher Ca and Mg, and a ten-dency towards
lower K in mango fruit, have been noted in fruit from orchards with no incidence
of soft nose, compared with fruit from orchards with high incidence of the disorder
(Burdon and Moore, 1991). Conversely, K levels are positively correlated with
disease resistance; Karunanayake (2007) reported reductions in disease on mango
fruit from trees receiving additional K. Application of triple the recommended level
of K significantly reduced stem end rot caused by Lasiodiplodia theobromae, while
the severity of anthracnose was most reduced by application of the recommended
level of K compared to nil and triple rate treatments.

Excess N can reduce storage life and quality, and excessive levels cause
deterioration of avocado fruit quality (Wolstenholme, 2004) where its effect may
be mediated through vegetative:reproductive balance and crop load (Hofman and
Mullen, 2005). High N has been associated with increased dis-orders in mango
(Young and Miner, 1961; Mead and Winston, 1991), possibly through the dilution
effect of increased fruit size on Ca concentrations; how-ever, soil N applications
later during fruit growth do not affect watery pulp
542 G.I. Johnson and P.J. Hofman

breakdown in fruit ‘Keitt’ fruit (Bally, 2007). Heavy rain late during fruit
development can release soil N previously unavailable to trees due to low soil
moisture, potentially resulting in high fruit levels. Boron (B) deficiency has been
related to abnormal fruit development (Lahav and Whiley, 2002) and increased
fruit storage disorders (Yogaratnam and Johnson, 1982; Smith et al., 1997). It may
also be important in mango (Coetzer et al., 1991). The effect of larger fruit size and
maturity on shelf and storage life (Seymour et al., 1990) can also be mediated
through production factors influencing fruit set and leaf:fruit ratio. Fruit position in
the canopy may also play a role here (see above).

Pests and diseases

Postharvest diseases and pests are reduced by various preharvest control measures
including orchard hygiene, manipulation of flowering, integrated management and
the use of chemical and biological controls (Johnson et al., 1989a; Johnson, 1997;
Fonseca et al., 2004b; Ploetz, 2004; Akem, 2006; Astridge and Baron, 2007a, b, c;
Chin et al., 2007; Diedhiou et al., 2007). Prusky et al. (Chapter 7, this volume) and
Ploetz and Freeman (see Chapter 8, this vol-ume) reviewed preharvest
management of several postharvest diseases, while Dann et al. (2005, 2007)
reviewed novel treatment options. Under the range of subtropical to tropical and
dry to wet ecoclimates, combinations of treatments have been recommended to
protect vegetative growth flushes, flower panicles and developing fruit from
infections that lead to anthracnose, bacterial spot, powdery mildew, scab and stem-
end-rot symptoms on fruit during development or after harvest (Poffley et al.,
1999; Ledger, 2004; Sto-volt and Dirou, 2004; Akem, 2006).

Lonsdale (1993) found that monthly applications of copper oxychloride


(CuCl2·3Cu(OH)2) in combination with mancozeb controlled most mango
postharvest diseases. Copper (Cu) alone was less effective in controlling
anthracnose. Copper sprays also provide protection against bacterial spot, while
mancozeb can provide protection against scab (Poffley et al., 1999; Led-ger, 2004).
Timmer and Zitko (1996) and Hardy et al. (2004) discussed the application of
copper treatments to citrus for disease control. Formulations differ in their weather
hardiness and indicated that retention of copper oxide (CuO) is superior to
retention of copper chloride (CuCl2) or oxychloride, and application of Cu with the
pH <7–6 can damage fruit and leaves. Lonsdale (1993) considered there was no
disease control benefit on mango by alternat-ing Cu with prochloraz sprays, but
Ledger (2004) recommended their strate-gic application every 3–4 weeks in
rotation with mancozeb and Cu when rainy conditions favoured anthracnose on
developing fruit.
Azoxystrobin (Amistar®) and other strobilurin fungicides effectively control
anthracnose, alternaria and powdery mildew on mango (Reuveni et al., 1998;
Willingham et al., 1999; Reuveni, 2000; Sundravadana et al., 2006, 2007),
Botryosphaera parva (Syn., Dothiorella dominicana) and Phomopsis sp. causing
stem end rot (Everett et al., 2005), and Cercospora leaf spot (Ane-siadis et al.,
2003) on other hosts. In Australia, no more than three strategic
Postharvest Technology 543

applications of azoxystrobin are recommended for field control of anthra-cnose on


mango in alternation with Cu, prochloraz and mancozeb schedules. Azoxystrobin
should be appled as one or two applications at flowering and/ or early fruit set at no
less than 14-day intervals, and again at 21 and 7 days before harvest (Stovolt and
Dirou, 2004). Application of azoxystrobin to control anthracnose on mango leaves
significantly reduces subsequent fruit disease and boosts yield (Sunarharum, 2007).

Recent research has also focused on the potential of defence-boosting


treatments, applied before or after harvest to reduce disease impacts on yield and
shelf life (Terry and Joyce, 2004; Dann et al., 2007; Karunanayake, 2007).
Anthracnose on mango fruit can be reduced by salicylic acid, its functional
analogue benzothiadiazole (BTH) (= acibenzolar-S-methyl = Bion®), and
ultraviolet (UV-C) irradiation with variable results, including phytotoxic effects
(Zainuri et al., 2001; Zeng and Waibo, 2005; Zainuri, 2006; Zeng et al., 2006;
Karunanayake, 2007).
Orchard hygiene, including reduction of inoculum by removal of old fruit and
branches, and removing prunings from the orchard, can reduce anthracnose and
stem end rots on fruit after harvest (Johnson, 1994; Ledger, 2004; Ploetz, 2004).
Some field diseases can disfigure fruit. Following heavy rain close to harvest,
bacterial spot can appear as discrete raised lesions, fol-lowed by fruit cracking or
rupture and fruit drop. Black spot orchard man-agement practices (i.e. windbreaks,
Cu sprays and use of resistant cultivars) can reduce the risk of damage on fruit
(Johnson et al., 1996; Dodd et al., 1997; Ploetz and Prakash, 1997). In cool
conditions, powdery mildew infection on young fruit can cause a ghosting
symptom on mature fruit similar to that caused by powdery mildew on apples. Scab
and sooty moulds can disfigure fruit. Generally, spray programmes for anthracnose
will control scab (Condé and Pitkethly, 2007), while sooty moulds are managed
through integrated management of scale insects and by postharvest brushing
(Johnson and Coates, 1993).

Tree nutrition can affect fruit disease incidence and severity. High Ca can
reduce fruit diseases in many fruit crops (Hofman and Smith, 1994). Nitro-gen
application before flowering can increase mango fruit disease (H. Nguyen et al.,
2004); applications up to 6 weeks after flowering can increase anthra-cnose
severity (Bally, 2007). Negative correlations occur between exocarp N percentage
and antifungal resorcinol concentrations.
Poffley et al. (1999), Peña (2004) and Peña et al. (see Chapter 10, this vol-
ume) discussed integrated pest management (IPM) for reducing pest dam-age and
quarantine hazards associated with fruit. Preharvest control measures for fruit flies,
seed weevils, scales and other skin defect-causing pests con-tribute significantly to
product quality improvement. IPM strategies can adequately control orchard pests
while reducing reliance on pesticides (Cun-ningham, 1986, 1991a, b, c;
Vijaysegaran, 1994; Peña, 2004; Peña et al., Chap-ter 10, this volume). Bagging of
developing fruit can reduce or eliminate disease infection (Hofman et al., 1997a)
and fruit fly infestation (Kitagawa et al., 1992). However, for export market access,
and regardless of effective field control, postharvest disinfestation measures are
often mandatory.
544 G.I. Johnson and P.J. Hofman

Weather conditions

Rain before harvest, and high RH and temperatures can increase disease lev-els,
fruit susceptibility to heat and brush damage, lenticel damage and reduce storage
life (Dodd et al., 1991a; Estrada et al., 1993; Prusky et al., 1993a, b; Cooke and
Johnson, 1994; Oosthuyse, 1998; Jacobi et al., 2001b). Disease risk predic-tion
based on the monitoring of environmental variables to determine fungi-cide
application frequency, can reduce pesticide residues (Fitzell and Peak, 1984; Fitzell
et al., 1984; Peak et al., 1986; Dodd et al., 1991a, b, 1992; Prusky et al., 1993b).
Irrigation frequency and water availability for tree growth can signifi-cantly impact
postharvest diseases and disorders (Simmons, 1998).

15.4 Flavour and Aroma


Flavour is largely determined by sugars and volatiles in the ripe fruit, both of
which increase in more mature fruit. The aroma produced by ripening and ripe
mango can help attract customers, and provide some indication of fla-vour
development. In mango fruits, more than 280 different aroma volatile compounds
have been reported (Singh et al., 2004). Variation in the constitu-ent aromatic
compounds in mango cultivars results in aroma and flavour diversity (MacLeod
and Snyder, 1985; MacLeod et al., 1988; Torres et al., 2007). The high fruit levels
of D-terpinolene contribute to the characteristic flavour of stronger-flavoured
cultivars such as ‘Kensington Pride’ (Bartley and Schwede, 1987; MacLeod et al.,
1988). ‘Kensington Pride’ harvested at the green-sprung stage have higher
concentrations of total aroma volatiles com-pared with fruit harvested at the hard-
green or coloured stages (Lalel et al., 2003b). Most of the glycosidally bound
aroma compounds increase in the pulp as the fruit matures, which contribute to
improved flavour. During the first 7 days of ripening, D-turpinolene is the major
volatile compound, but in the later stages of ripening ethyl octanoate dominates
(Lalel et al., 2003a).

15.5 Harvesting and Transport to the Packhouse


Harvesting of mango is determined according to attainment of acceptable/ required
maturity: (i) for arrival at market during the time of peak demand/ highest price to
maximize the chance of early sale; or (ii) to minimize loading wait at the shipping
port. Fruit is generally picked into field crates or bins, with or without the use of
mechanical picking platforms.

Timing

Maturity is determined by assessing variables such as days from flowering,


accumulated heat units, flesh dry matter percentage, flesh colour or fruit shape/skin
colour (see Maturity section under 15.3 Preharvest Management,
Postharvest Technology 545

this chapter). Fruit water potential fluctuates diurnally, and can affect fruit quality.
The water potential of fruit at harvest can affect susceptibility to han-dling, heat
damage and product storage potential (Joyce and Patterson, 1994). In hot weather,
fruit should be harvested in the coolest part of the day to reduce fruit overheating
and energy requirements for postharvest cooling, and to minimize worker
discomfort. Harvest during rain can reduce fruit quality (see Weather conditions
section under 15.3 Preharvest Management, this chapter).

Sapburn

Severing the stem from the fruit causes relatively large volumes of latex to spurt or
ooze from the cut stem. The sap is of low pH and high oil content and can burn the
surface of the fruit (Bagshaw and Brown, 1989). The oil frac-tion contains
terpinolene and resorcinols and is the fraction of the latex that causes the damage.
Skin damage is particularly severe with ‘Kensington Pride’ (O’Hare, 1994), and
less serious in Florida cultivars. In Pakistan, ‘Chausa’ is more susceptible than
‘Sindhri’ and ‘Dashehari’ (Maqbool et al., 2007). O’Hare (1994) observed that
latex levels are lower and less phytotoxic in ‘Nam Doc Mai’, ‘Nang Klang Wun’,
‘Tong Dum’ and ‘Keow Savoey’ (0.16– 0.48 ml/fruit), than in ‘Kensington Pride’
(1.67 ml/fruit). The oil compo-nent of the latex of Thai cultivars is much lower
than that of ‘Kensington’ (O’Hare, 1994; Hassan, 2007). The concentrations and
ratio of the two main resorcinols, 5-n-pentadecylresorcinol and 5-n-
heptadecenylresorcinol, differ among cultivars (Hassan, 2007).

Factors affecting the potential of latex to cause sapburn are not well
understood. It appears that both the terpene in the oil fraction of the sap, and
adequate polyphenol oxidase (PPO)/peroxidise concentrations in the skin are
required to develop sapburn; PPO and resorcinols in sap are less signifi-cant
(Loveys et al., 1992; Robinson et al., 1993; John et al., 2002). Rain near harvest
and high N in fruit result in more severe sapburn in ‘Kensington Pride’. However,
negative relationships have been observed between exo-carp N concentration and
alkyl resorcinols (Hassan, 2007). Sap from fruit harvested early in the day causes
less sapburn than sap from fruit harvested later in the day, although early-harvested
fruit exude more sap than late-harvested fruit (Maqbool et al., 2007).

Latex may provide protection against infestation by fruit fly larvae (Joel, 1978,
1980) and may also contribute to disease tolerance. The 5-substituted resorcinols
have antifungal properties (Cojocaru et al., 1985; Droby et al., 1986, 1987; Prusky
and Plumbley, 1992). Karunanayake (2007) extracted a resorci-nol and a resorcinol
derivative from the dichloromethane phase of Sri Lankan mango peel extracts that
had antifungal properties. Differing relationships occur between resorcinol levels
and relative susceptibilities of different mango cultivars to anthracnose
(Karunanayake, 2007; Hassan et al., 2007). Strong positive relationships occur
between resorcinol concentration in the peel and latex, and fruit resistance to
artificially inoculated anthracnose.
546 G.I. Johnson and P.J. Hofman

‘Kensington Pride’ has more non-aqueous latex, higher concentrations of resor-


cinols and greater tolerance of anthracnose than ‘Nam Doc Mai’ (Hassan, 2007).
Less anthracnose occurs on fruit ripened with an intact stem, compared with de-
sapped fruit. Hegnauer (1994) reviewed the phytochemistry of Mangifera.
Cultivars that are prone to sapburn can be harvested with 10–20 mm stems
attached, and re-trimmed at the packhouse. Latex does not usually exude from
longer stems because there is no continuity between the fruit and stem resin ducts
(Joel, 1980), and the fruit lactifers are not severed; however, stems can break in
transit to the packhouse, resulting in latex leakage and sapburn. Long stems left on
the fruit to reduce sapburn has variable effects on disease depending on disease
type and storage time. With shorter storage periods, anthracnose and stem-end-rot
incidence and severity can be lower in fruit with stems attached due to higher
levels of resorcinols (Hassan, 2007), but during longer storage stem-end-rot levels
can increase due to the higher levels of inoculum associated with the retained stalk
(Johnson et al., 1993).
Mango latex can cause skin disorders in humans (Keil et al., 1946; Oka et al.,
2004). Bandyopadhyay et al. (1985) noted that resorcinol derivatives are allergens
in the Anacardiaceae, and suggested that the 5-substituted resorci-nol in mango
latex causes dermatitis. Both heptadec(adi)enyl resorcinols and
pentadecylresorcinol can elicite an allergic reaction in sensitive patients (Oka et al.,
2004). Harvesting and packhouse personnel must avoid contact with the latex of
high-risk cultivars.

Harvesting and desapping

Rough handling at harvest can cause skin damage and internal fracturing or
bruising. Using hooked sticks or shaking the tree to detach fruit causes skin
damage and flesh fracturing (Ledger, 1991a; Abu-Goukh and Mohamed, 2004) and
sapburn. Mechanical damage during harvest also causes soft, darkened areas and
bruises on fruit following hot water treatment. Mangoes should be handled as if
they were eggs. Long-handled secateurs cut and grip the stem, allowing the fruit to
be carefully lowered to the picking bin. Contact with soil and soilborne pathogens
should be avoided (Johnson et al., 1993).
In cultivars where sapburn is a problem, latex should be drained from the fruit
(desapping or bleeding) to minimize the incidence and severity of sapburn. Several
systems have been assessed for reducing damage (Brown et al., 1986; Ledger,
1991b; Holmes et al., 1993; Lim and Kuppelweiser, 1993; O’Hare and Prasad,
1993; O’Hare, 1994; Shorter and Joyce, 1994). In Austra-lia, the main commercial
practices (Plate 82) are:

● Desapping in the field with harvest aids using detergent. The basic de-sign
characteristics include detergent spraying onto a tarpaulin, a trough with the
same detergent and a final spray before fruit are placed in a field bin. The fruit
are either hooked from the tree in the direction of the harvest aid and onto the
catching surface or the fruit are snapped directly off the tree and placed onto
the tarpaulin. The fruit roll from the tarpaulin into
Postharvest Technology 547

the trough containing detergent and then into 300–400 kg field bins. Alkaline
detergents that deactivate damaging sap components are most effective; high
concentrations of surfactant in the detergent are not re-quired. The crucial
factors are that fruit should be exposed to detergent for at least 90 s, the
detergent is either not recycled, or replaced before sap accumulation in the
detergent causes other damage such as skin browning (Bally et al., 1997;
O’Hare et al., 1999).
● Another design includes a motorized hydraulic ladder (cherry picker) with the
fruit desapped for 1–2 s before placing in a basket containing a spray of
alkaline detergent. This system is particularly effective for tall trees, but care
must be taken that fruit are covered by the detergent for at least 90 s.

● Picking fruit with long stems into small 18 kg crates and desapping in the shed.
The fruit are dipped into detergent before desapping and plac-ing on a long
conveyor system that holds the fruit inverted and provides detergent/water
sprays for a few minutes. The fruit are inverted for c.20 min before drying and
packing.

In these systems, the detergent is not strongly alkaline, but the surfactant should be
of sufficient concentration to provide a protective coating around the fruit before
desapping. Stem breakage must be minimized in the crates, as this can cause
sapburn and quality loss (Holmes et al., 1993; Holmes and Bally, 1994). Latex
must not spray or drip onto fruit being desapped. Workers who are sensitive to
mango sap should wear hand protectants, aprons and footwear to minimize skin
contact. The detergent must reduce sapburn and skin browning without causing
other damage (i.e. lenticel spotting) (Fig. 15.3). Desapping in the field by inverting
fruit directly onto racks without detergents has been used for sensitive cultivars and
when particular growing conditions have increased fruit susceptibility to lenticel
spotting or other damage from detergents; however, labour costs are becoming
prohibitive, requiring compromises between cost and quality. Desapping by
inverting and placing on the ground significantly increases the incidence and
earlier appearance of stem end rot caused by soilborne L. theobromae, and is not
recommended (Johnson et al., 1993).

Harvest aids have reduced in-shed desapping. Holmes et al. (1993) found 9–
16% of fruit in field crates were affected by sapburn when harvest aids were not
used. Harvest aids provide the greatest reduction in total sapburn (from 69% to 15–
18%). While harvest aids can significantly reduce sapburn, inappropriate use can
increase some forms of skin browning (Bally et al., 1997). Underhill and Dahler
(1995) described four types of skin browning which produce symptoms distinct
from the sapburn caused by the oil phase of latex. Several forms of skin browning
involve tissue reactions with sap/ detergent mixtures. Symptoms vary if latex enters
fruit through micro-cracks in the cuticle or the lenticels (O’Hare et al., 1999).
Holmes (2003) developed guidelines for the use of harvest aids.

Cost savings associated with the use of harvest aids can be lost if fruit-to-fruit
or fruit-to-ground impact is not minimized during harvest. Rough
548 G.I. Johnson and P.J. Hofman

Block 1

Lenticel spotting (7days after harvest)


Block 2
Block 3
3

c
Control LOC Cl Magi Plus Wash
ear Glo
Mango Mango Mango Mango Man go Superconcentrate

Detergent

Fig. 15.3. Effect of detergents on lenticel spotting on skin of ‘B74’ mango (0 = no


lenticel spotting; 4 = severe spotting). Fruit were obtained from three different
blocks in two orchards, dipped in detergent for 2 min, then held at 20C for 7 days
(Source: Hofman and Ledger, 2006). The bar represents the least significant
difference (LSD) at 5%.

harvesting can increase the incidence of bruising and internal fracturing, and lower
wholesale returns. Thorough training of picking crews and supervi-sion of their
performance is required to maintain good practice.

Transport to the packhouse

Harvested fruit should be transported to the packhouse as soon as possible, with no


prolonged exposure to the sun. Rough handling and transport must be minimized.
Roads/tracks from orchard to packhouse should be smooth, with transport vehicle
tyres correctly inflated, and special suspensions to reduce vibration and damage.

15.6 Packhouse Measures


Harvested fruit are transported to a central packhouse which provides shel-ter from
rain and sun, and facilities for cleaning, treating, packing, cooling and storing fruit
until consignment to market (Schoorl and Holt, 1982, 1985). Mechanized
packhouse systems can offer labour savings and increased re-turns (Murray and
George, 1994). When manual handling was reduced from five to two steps, fruit
appearance improved, disease losses were lowered, sizing accuracy improved,
packing rate increased and space, labour and supervisory requirements were
reduced (Murray and George, 1994).
A typical packhouse sequence is shown in Fig. 15.4. In packhouses that
include a disinfestation facility, the sequence must be modified to allow for
Postharvest Technology 549

1. Deliver 18. Air or sea freight

2. Weigh 17. Cool chain transport

3. Cold wash and brush 16. Palletizing

4. Hot water/fungicide bath 15. Outer boxing

5. Fungicide application 14. Cool/ethylene gas

6. Dry 13. Pack

7. Wax and brush (optional) 12. Disinfestation

8. Dry 11. Crates

9. Grade 10. Size

Processing Second First grade Extra class


grade grade (domestic/export) (export)

Processing Local marketing

Fig. 15.4. Packhouse and marketing activities for mango. Waxes are not applied in some
countries because of abnormal ripening and off-flavour development. When disinfestation is
not required, steps 11, 12 and 15 would be omitted. When fruit are heat disinfested, they
may be packed into an inner box prior to cooling. For some markets accessible by air, the
fruit may be treated with ethylene and stored until near ripe. The boxed fruit may then be
packed into an outer carton prior to palletizing. Pre-ripening allows fruit to be rechecked
prior to despatch, with fruit of unsatisfactory appearance (e.g. skin damage) redirected to
domestic market or processing.
550 G.I. Johnson and P.J. Hofman

sizing before disinfestation, with tray packing after treatment. Packhouse design
and installation consultants can provide substantial savings by elimi-nating
bottlenecks and minimizing product damage points.

Delivery inspection and traceability

Harvested fruit in field crates should be treated, packed and cooled as soon as
possible. Quality and contaminant management systems may require that a record
system tracks the block or trees from which each bin of fruit is har-vested. This
enables records to be kept of tree or block yield, quality perfor-mance and defect
levels as well as labour performance rates and pesticide residues. Individual fruit in
a tray can be traced back to the tree or block from which it was harvested.
Block/tree-to-tray traceability systems and pack-out records allow problems (i.e.
excessive or unapproved pesticide detections) to be traced to the site of the
problem and relevant action taken. They can also be used to motivate producers or
picking teams to deliver high quality produce.

At the packhouse, samples of fruit should be evaluated immediately for


maturity, blemishes and disease and pest incidence and recorded in an appro-
priately designed computerized system. Preharvest orchard inspections can reveal
the defects that can be anticipated in the packhouse. Some degree of in-field sorting
can occur at the point of harvest, and soft or damaged fruit collected separately and
discarded.

Desapping and washing

Unloading should avoid dropping, damaging and wounding of fruit. Fruit are
normally unloaded from field bins into bin dumps if desapping is unnec-essary or
removed manually from the crates for desapping (see Harvesting and desapping
section under 15.5 Harvesting and Transport to Packhouse, this chapter).
Detergents and sanitizers are sometimes added to washing water. Their use requires
careful consideration. Some may cause fruit dam-age, or promote early fruit
disease expression (Korsten et al., 1993). Chlorine is added and carefully regulated
to wash and/or rinse water in some pack-houses, but this is not essential.
Quaternary ammonium disinfectants should not be added to wash water as their
direct application to foodstuffs is gener-ally not permitted.

Disease control

Hot water and fungicide application


Hot water dips, or sprays over brushes, with or without fungicide, and fun-gicide
sprays or dips, can eradicate quiescent fungal infections that have been established
on and beneath the cuticle and within the pedicel prior to
Postharvest Technology 551

harvest (Johnson et al., 1989a, b, 1991, 1992; Kernot et al., 1999; Poffley et al.,
1999; Plan et al., 2002). Suslow (2000) provides generic recommendations for the
postharvest handling of produce. Postharvest disease treatment efficacy varies with
infection level, cultivar, ripening status and storage regime. Hot water treatment
also cleans fruit, but can contribute to increased skin dam-age (Cooke and Johnson,
1994). Anthracnose caused by Colletotrichum gloeo-sporioides Penz. and
Colletotrichum acutatum Simmonds, is controlled more readily than stem end rots
(or soft brown rot) caused by anamorphs of Botry-osphaeria spp. (Fusicossum spp.,
Neofusicoccum spp., L. theobromae (Pat.) (Griff. and Maubl.)) (Johnson, 1994;
Slippers et al., 2005; Crous et al., 2006) and Pho-mopsis mangiferae Ahmad, and
alternaria rot caused by Alternaria alternata (Fr.) Keissler. The latter is generally
only a problem in fruit from dry regions or in fruit from more humid areas during
storage for 3 weeks or more (Prusky et al., 1980, 1993a, b, and Chapter 7, this
volume; Johnson et al., 1990b).
Fruit are moved through a water bath for 5 min at 48–50C for less mature
fruit and hot-water-damage-susceptible cultivars (e.g. ‘Zill’ and ‘Irwin’) and at 50–
55C for mature fruit and less susceptible cultivars (Anonymous, 1994b).
Treatment for 3 min may be adequate for control of anthracnose, while immersion
for up to 7 min may enhance control of stem end rot (Muirhead and Grattidge,
1986; Sepiah, 1986; Johnson et al., 1989b). In large-scale facili-ties, dip tanks may
range from 3000 to 5000 l, with fruit immersed and moved through the tank by a
series of paddles. In tank construction, non-corrodible materials such as stainless
steel and fibreglass are preferred, and the con-veyor system that contains the
paddles should travel along the bottom of the tank to reduce damage to fruit that
sink. Accuracy in temperature control, efficacy of the heating unit and timing of
fruit flow through the bath are critical. Temperature probe placement at pump inlet
and outlet and thorough water circulation to ensure accurate temperature reading
and to minimize hot spots are critical. Impurities (e.g. minerals, sediment and
debris) in dip water can affect fungicide performance and stain or damage the fruit.
In-line filters in the inlet and pump circulation systems should be installed and
cleaned regularly.

Where acceptable, carbendazim can be added to the hot water at the rec-
ommended rate, and topped up and replaced regularly, to provide improved control
of stem end rot and anthracnose at lower temperatures (52C). Beno-myl has been
withdrawn for postharvest use, but much of the benomyl use information (from
earlier research) is relevant for carbendazim (Johnson et al., 1997). Also, hot
thiabendazole (TBZ) is generally as effective as hot benomyl for controlling stem
end rot, but may provide inferior control of anthracnose (Coates et al., 1993). The
active component of benomyl and TBZ in plants, carbendazim (MBC), is identical
(Erwin, 1973; Muirhead, 1976); however, TBZ also contains sulfur (S) which
affects its rate of breakdown and spec-trum of activity (D. Guest, personal
communication, Melbourne, 1995). Beno-myl penetrates plant tissue more
effectively than TBZ, carbendazim or thiophanate methyl (Eckert, 1983).

Dipping fruit in hot, dirty, latex-contaminated water can increase phyto-


toxicity and lenticel damage. Hot fungicide dips lose efficacy due to sap
552 G.I. Johnson and P.J. Hofman

Table 15.3. Registered uses of prochloraz for postharvest treatment of mango


(Source: adapted from Lunn, 2004).

a
Country Form Method Rate (kg ai/100 l)
Australia ec 30 s spray 0.025
Brazil ec 2 min dip 0.05
China ec/wp 1 min dip 0.05–0.1
Colombia ec Not specified 0.025
Peru ec Not specified 0.02–0.045
South Africa ec 20 s dip 0.08
a
ai, active ingredient; ec, emulsifiable concentrate; wp, wettable powder.

build-up in the dip tank and stripping out of fungicide (Wells and Littlemore,
1989). Ledger (2004) optimized dip:fruit ratio and dipping practices. Prochlo-raz
provides good control of anthracnose and alternaria rot, but does not pro-vide
control for stem end rot (Johnson et al., 1990b; Johnson and Coates, 1993). In
South Africa, for local markets prochloraz is added at 405 ppm of active ingredient
(ai) of a 45% emulsifiable concentrate (ec) formulation; for export markets, 810
ppm prochloraz is used. Fruit are immersed for 20 s. In Australia, prochloraz at 250
ppm is applied by overhead spray, and fruit require 15–20 s to pass through the
prochloraz spray race on a roller conveyer system.
A maximum residue level (MRL) of 7.0 mg/kg for prochloraz for assorted
tropical and subtropical fruits with an inedible peel is recommended (CODEX-
MRL, 2008). This group MRL replaces individual fruit commodity MRLs, and
takes into account the lower residues in the flesh compared to the skin (Muller and
Burt, 1989). Some registered use-rates for postharvest applica-tion of prochloraz
are listed in Table 15.3.
Hot water sprays over brushes (55C for 15–20 s) is an effective alterna-tive to
hot water dips containing prochloraz for controlling alternaria rot (Prusky et al.,
1999, 2006). Application of hot water spraying and brushing for 15–20 s (HWB)
followed by a spray of 50 mM hydrochloric acid (HCl), alone or in combination
with prochloraz, also improved control of alternaria rot (Prusky et al., 2006). These
treatments have not been tested for anthracnose. Using 2,4-dichlorophenoxyacetic
acid (2,4-D) diluted in wax after HWB and prochloraz reduces stem end rot
(Kobiler et al., 2001). A hot water and beno-myl combination treatment followed
by a prochloraz spray provides effec-tive control of anthracnose, stem end rot and
alternaria rot during longer storage (Johnson et al., 1989a, 1990b). Similar benefits
are now attributed to hot TBZ dip and cold prochloraz spray (Ledger, 2004).

When fungicides are used in the packhouse, spent dip suspensions and
fungicide containers must be disposed using approved methods, often included
with supplier recommendations. Carbendazim suspensions can be drained into a
trench filled with stones, but runoff must be avoided. Car-bendazim and other
benzimidazole fungicides are toxic to earthworms (Wright and Stringer, 1973).
Postharvest Technology 553

Heat
Pest disinfestation treatments involving heat provide some control of anthra-cnose,
but do not adequately control mango fruit pathogens for export. Tem-perature and
time combinations suitable for non-deleterious fruit disinfestation are sublethal to a
significant percentage of quiescent infections beneath the fruit cuticle and pedicel
tissues (Coates and Johnson, 1993). In many regions, fruit skin temperatures
frequently approach the mid-40C range during pre-harvest development, a natural
selection pressure favouring heat-tolerant fungal infection structures. For
‘Kensington Pride’, hot benomyl in combina-tion with either prochloraz or vapour
heat at 46.5C for 20 min controls stem end rot more effectively than hot benomyl
alone and TBZ, alone or in combi-nation with vapour heat, during storage at 23C
for 15 days (Coates et al., 1993). Disease control in combination with heat
disinfestation has been reviewed by Coates and Johnson (1993) and Jacobi et al.
(1994, 2001a).

Future options
Heat is an ideal disease control treatment, since it is environmentally safe and non-
chemical. Its effectiveness would be enhanced if fruit tolerance could be increased
by genetic manipulation or the development of pre-conditioning treatments. Pre-
treatments to render quiescent structures of pathogens more susceptible to heat
would also improve disease control. Measures to increase efficacy could include
other energy sources, chemicals, adjuvants, fumigants or microorganisms to
damage or soften fungal wall structures.
Treatments to delay fruit ripening also limit or reduce disease losses. Storage
quality would benefit from the development of cultivars or pre-conditioning
treatments to improve tolerance of fruit for cool storage or controlled atmo-sphere
(CA) and modified atmosphere (MA) storage (Brecht and Yahia, Chapter 14, this
volume). With increasing concerns about the use of chemi-cals on food (Gullino
and Kuijpers, 1994), and in view of current limitations on heat treatment and
storage regime disease control efficacy, non-deleteri-ous alternatives to synthetic
fungicides are required. Alternatives to fungi-cides for controlling postharvest
diseases have been reviewed by Johnson and Sangchote (1994) and Korsten
(2006). Options include: (i) biological con-trol, i.e. the use of microorganisms to
control pathogens (Wilson and Pusey, 1985; Jeffries and Koomen, 1992; Korsten et
al., 1993, 1994; Korsten 2006); (ii) enhanced exploitation of naturally occurring
antifungal compounds in fruit (Prusky et al., 1982; Johnson et al., 1998; Joyce et
al., 1999; Zainuri et al., 2003; Hassan et al., 2007); (iii) application of fruit
coatings such as chitosan with both MA and antifungal effects (El Ghaouth et al.,
1992a, b; Wilson et al., 1994; El Ghaouth and Wilson, 1995); (iv) exposure to UV-
C light (wavelength <280 nm) (Chalutz et al., 1992; Wilson et al., 1994). Zainuri
(2006) reported some promise in the use of UV-C radiation for control of
anthracnose, but fruit damage risks and treatment dose accuracy were critical; (v)
containment of fruit in atmospheres containing high levels of carbon dioxide (CO 2)
for 24–48 h after harvest (flushing) (Prusky et al., 1992, 1993c); (vi) regulation of
fruit ripening (Brady, 1994); and (vii) application of naturally occurring plant
products (Fallik and Grinberg, 1992; Wagner and Flores, 1994). Many of these
554 G.I. Johnson and P.J. Hofman

options may delay disease development by eliciting increases in antifungal


compounds in the fruit (Prusky and Keen, 1993; Wilson et al., 1994; Zainuri,
2006).
What are the alternatives to synthetic fungicides for controlling mango dis-
eases? Bacteria active against mango isolates of C. gloeosporioides, stem end and
soft brown rot pathogens have been evaluated (Koomen et al., 1990; Korsten et al.,
1991, 1992, 1993; Jeffries and Koomen, 1992; Coates et al., 1995; Korsten, 2006).
Antifungal resorcinols in the peel of mango fruit interfere with the devel-opment of
anthracnose and alternaria rot (Cojocaru et al., 1985; Droby et al., 1986, 1987;
Prusky and Keen, 1993; Zainuri, 2006; Hassan, 2007), with higher levels present in
some cultivars (Hassan, 2007; Karunanayake, 2007; Hassan et al., 2007). Lonsdale
(1992, 1993) found that enclosure of ‘Keitt’ in high-density polyethyl-ene bags
with 30% CO2 and 15% oxygen (O2) for 24 h at 11C prior to storage, improved
control of anthracnose. However, 24 h exposure to 20% CO 2 signifi-cantly
increased the incidence of soft brown rot (stem end rot) in ‘Keitt’ and ‘Kent’
compared to untreated fruit, especially in the absence of O 2. UV irradia-tion of
fruit for 10–30 s in combination with wax prior to storage is similar to hot water in
reducing the incidence of soft brown rot compared to untreated fruit.

Brushing

Brushing on mango packing lines can occur after, or at the same time as, hot water
and fungicide treatments. Hot water treatment washes sap away, and loosens
superficial debris, scale insect carapaces and sooty mould, which are removed as
the fruit pass over rotating brushes. Brushing also removes superficial deposits of
fungicides that accumulate on fruit from orchard application of Cu fungicides
(Lonsdale, 1993) and incorrect mixing or sedi-mentation of benzimidazole
fungicides resulting from sap accumulation in dip tanks (Wells and Littlemore,
1989). Soft, non-damaging brushes should be used, washed every day and replaced
seasonally.
For ‘Kensington Pride’ mangoes harvested after rain, skin marking, fruit
shrivel and weight loss increase significantly on fruit treated with a hot water and
fungicide dip or a hot water and fungicide dip followed by treatment with
prochloraz, when fruit brushing followed either or both treatments relative to
untreated and untreated/brushed fruit. Prochloraz before brushing resulted in fruit
quality similar to untreated or brushing only (Cooke and Johnson, 1994). Brushing
can increase lenticel spotting (Oosthuyse, 1999). Therefore, the num-ber and type
of brushes must remove foreign matter and polish the fruit, while not increasing
risk of brush and lenticel damage, especially during wet weather and with heat
treatments (see Weather conditions and Skin colour and lenticel damage sections,
both under 15.3 Preharvest Management, this chapter).

Grading and sizing

The purposes of grading are to sort fruit into defined categories of unifor-mity and
to divert out-of-grade fruit from the pack line to either a second
Postharvest Technology 555

grade, processing or reject line. Mangoes with defects outside acceptable lim-its as
defined in a grade schedule or chart are manually removed and trans-ferred (by
conveyer belt) to seconds or processing lines as appropriate. The purpose of sizing
is to categorize fruit into size or weight groups for packing. Fruit must be sized
prior to disinfestation with hot water or vapour heat to ensure consistent treatment
responses. Typical systems include automatic graders that separate fruit by weight
into groupings that correspond to pre-determined categories (Schoorl and Holt,
1982, 1985). Camera vision systems can separate for colour, defects and shape.
Fruit usually accumulate in sepa-rate bins for packing into cartons or into bulk
containers for processing or disinfestation. The fruit are packed manually into
single-layer trays, with plastic or cardboard liners that have depressions designed to
accommodate fruit of a particular size. The depressions provide some support for
individ-ual fruit during packing, while the cardboard liners also provide some
additional buffering against impact damage. The pattern of the depressions
facilitates most efficient utilization of carton space. Mango tray liners com-monly
accommodate 12–25 fruit for 6.5 kg trays. Some tray liners may be inappropriate
for sea export due to interference with vertical airflow.

Organic materials (i.e. paper, leaves or shredded wood) have been used to
cushion individual fruit in cartons. These materials can harbour pathogens, for
example Rhizopus stolonifer (Ehrenb. Fr. Lind.), which causes transit rot of
mangoes and has been detected in shredded wood used in mango packaging.
Shredded wood creates micro-wounds in the fruit skin, providing points of entry
for hyphae growing on the wood. Losses are more severe when fruit have been
removed from cold storage, allowing condensation to develop on the fruit and
shredded wood (Muirhead and Grattidge, 1986).

Grade standards

The International Standardisation Organisation (ISO) is a non-governmental


organization (NGO), and is a network of national standards institutes (157
countries). ISO is the global leader for development and publication of stan-dards.
ISO publishes a range of standards for fruit and vegetables, testing, crop and
postharvest management procedures and food safety, system auditing and nutrient
and water testing that are relevant to mango systems’ benchmarking and
improvement. A range of standards may also be defined within GAP certification
protocols and nationally developed marketing arrangements. Agreed grade
standards provide a reference point for produc-ers and traders in production and
marketing (EurepGAP, 2007; ISO, 2008). The CODEX Alimentarius Commission
also oversees the development of standards for fresh and processed fruit with the
CODEX Committee on Fresh Fruit and Vegetables (CODEX, 2008a). CODEX
standards for fresh fruit spec-ify provisions for quality, sizing, tolerances,
presentation, marking and label-ling, and contaminants (CODEX, 2008a). There
are CODEX standards for fresh mangoes, canned mangoes and mango chutney
(CODEX, 2008b).
556 G.I. Johnson and P.J. Hofman

Minimum requirements for grade standards specify that fruit intended for
international trade should be intact, firm, fresh in appearance, sound, clean, free
from black stains and bruising, free from damage caused by low temperatures and
free from pests and pest damage. Fruit should be carefully picked at the stage of
physiological development which will allow transport and handling and
continuation of the ripening process so that fruit will ripen to consumer
expectations. Class standards can depend on customer specifi-cations, and can be
based on fruit size and appearance. Colour illustrations in Anonymous (1993) and
Amesbury et al. (2002) are indicative of some of the quality standards that can be
specified for appearance, shape and colour, and tolerance levels for superficial skin
defects. Similar charts are often available for individual cultivars and are produced
during the development of QA systems for specific marketing groups and
customers.

Packing-line QA inspections
Packing-line control inspections are used to monitor grading efficacy and packing-
line damage. Packing-line inspection samples are taken soon after fruit pass points
in the line at which defects are most likely to be overlooked and/or induced. For
start and end-of-line pack-out checks, and out-turn inspections, randomly selected
cartons are unpacked, and all fruit are checked for compliance with preset quality
parameters. Most value is gained from quality control checks if records are kept
and evaluated, with feedback/trou-ble shooting as necessary, to constantly improve
the system (Ledger and Bag-shaw, 1994; Ledger and Premier, 2006). Computer
analysis of such information provides a seasonal benchmarking record of QA
improvement, and high-lights areas for attention in packing-line improvement and
personnel train-ing. Record keeping is mandatory under GAP certification systems.

Future options
Greater automation of grading and packing will become necessary as pro-duction
and labour costs increase, and as customers become more demanding (Hilton,
1994). Recent advances in computing have made possible high-speed sorting using
visual systems for colour, shape and externally visible defects. Also, NIRS systems
can now be used in-line to sort for flesh characteristics that influence flavour. In
mango, percentage dry matter and flesh colour are related to ripe fruit flavour, and
can be estimated using NIRS (Saranwong et al., 2004; Subedi et al., 2007).
Estimation is sufficiently accurate to allow acceptable separation into several
categories for final flavour. NIRS may also be useful for predicting ripening
behaviour and weight loss during ripening (Mahayothee et al., 2004). Given the
influence of weight loss in chilling injury (CI) development during cold storage
(Bower et al., 2003), NIRS may also be able to estimate potential for CI during
cold storage.
There is interest in other non-destructive quality assessment for the pack-
house. Joyce et al. (1993) noted that future innovation could lead to proton
magnetic resonance imaging (MRI) technology suitable for packing-line
applications to allow non-destructive detection of internal disorders and pest
infestations. X-ray imaging may have potential for detecting seed weevil
Postharvest Technology 557

damage in mangoes (Thomas et al., 1995; Reyes et al., 2000), but recent inves-
tigations suggest that neither X-ray imaging nor MRI is sufficiently reliable for
quarantine purposes, particularly where larvae are small (R.A. Jordan, personal
communication, 2007). New methods of nuclear magnetic reso-nance (NMR)
(Marigheto et al., 2008) may distinguish internal disorders such as jelly seed.
Acoustic/ultrasonic methods can sort for fruit firmness (Miz-rach, 2008) and may
help identify softening fruit with internal disorders and reduced storage life.
Robotics in sorting and packing will be used increas-ingly where labour costs and
availability are high.

Disinfestation

Disinfestation treatments, backed by integrated field control programmes and/or


area freedom stipulations under market access approvals, provide assurance to
authorities of an importing country that the commodity will be free of target pests
and not pose a quarantine threat (Johnson and Heather, 1995; Follett and Nevin,
2005). Market access application and approval ar-rangements for most countries
operate under national legislation and regula-tions formulated under the framework
of the IPPC and the World Trade Organisation (WTO) Agreement on the
Application of Sanitary and Phyto-sanitary Measures (IPPC, 2008; SPS, 2008).
Key aspects of quarantine regula-tion of crop pests are covered by International
Standards for Phytosanitary Measures (ISPM), developed and agreed in
consultation with signatory cou-ntries to the IPPC under the Food and Agriculture
Organization (FAO) (ISPM, 2007a). ISPM (2007b) cover guidelines on the
phytosanitary measures for regulated pests. Under the IPPC, a technical panel on
phytosanitary mea-sures has oversight of international guidance on phytosanitary
treatments including assessment and recommendations for use as international
stan-dards. In addition to ISPM, a range of regional plant protection measures has
been established. It has been agreed internationally (ISPM, 2007b) that
phytosanitary treatments should fulfil the following requirements:

● Provide effective destruction, inactivation or removal of pests or render them


infertile or devitalized. Normally, stipulation of the destruction effica-cy level,
with quantification or statistical benchmarking, is required. When experimental
data are unavailable or inadequate, other evidence is needed to support a claim
of efficacy, for example historical/practical experience. The technology and
● treatment regime used should be: (i) clearly delin-eated, with evidence to
confirm adequate adherence to scientific meth-ods in generating data
(experimental designs). Data in support of the treat-ment efficacy should be
verifiable, replicable and statistically valid, and preferably published in a peer-
reviewed journal; (ii) appropriate for use in international or regional trade and
research; and (iii) safe to apply, with no undesirable impacts on treated
commodities or the environment.
Approvals for disinfestation treatments and market access are obtained on a
country-by-country basis. A starting point for intending exporters is to
558 G.I. Johnson and P.J. Hofman

Table 15.4. Examples of summary information for exporting Australian mangoes to


the EU and New Zealand (Source: adapted from AQIS, 2008b).

Required for

Documentation EU New Zealand

Import Permit No No
a
Phytosanitary Certificate Yes Yes
b
Additional Declaration No Yes
Post Entry Quarantine No No
EX188 No No
EX46 No No
Radiation Statement No No
a Treatment details, including date of treatment, are to be endorsed on the Phytosanitary
Certificate in the treatment section. The treatment is to be shown as: irradiation at minimum
250 gray.
b
The mangoes in this consignment have been treated in accordance with Appendix 12 of
the Bilateral Quarantine Arrangement between New Zealand Ministry of Farming and AQIS.

determine the disinfestation requirements for mangoes entering target mar-kets, for
example the import health standards for New Zealand (BANZ, 2008) and the Plant
Protection and Quarantine (PPQ) manuals for the USA (PPQ, 2007). Australian
exporters can use the Australian Quarantine and Inspection Service (AQIS) phyto
exports database (AQIS, 2008b). The AQIS web site plant product export database
provides summary information (Table 15.4) and detailed information on
phytosanitary requirements for potential exporters.
Follett and Nevin (2005) noted that increased trade has increased exotic pest
threats and attention to quarantine and regulatory issues. Risk-based alternatives
were replacing the probit 9 standard for quarantine efficacy. Cul-tivar testing was
seen as necessary only for some treatments and commodi-ties, and generic
treatments for broad groups of pests and commodities were seen as a means of
enhancing trade. Area-wide pest management was valued for preharvest pest
control and improvement of quarantine security for export products. However,
some treatments such as γ irradiation were not accepted by all countries and this
slowed their adoption. Follett and Neven (2005) concluded that efforts for
standardization of phytosanitary measures and research would improve information
exchanges and market access nego-tiations.

Target pests
A pest of regulatory concern that could become established in an area where it is
not found is a quarantine pest risk, and requires quarantine action. Mango fruit
pests include internal pulp feeders (i.e. fruit fly immatures), seed and fruit pulp
pests (i.e. mango weevils and fruit caterpillars) and external pests (i.e. scales,
mealybugs, thrips and mites) (see Peña et al., Chapter 10, this volume). External
pests pose detection risks as surface hitchhikers that
Postharvest Technology 559

can be detected visually by inspectors. Such pests need to be controlled in the field
and removed before the fruits are exported. Internal pests, such as wee-vils, fruit
fly immatures or larvae of Lepidoptera, pose additional risks because of difficulties
of detection and their potential to damage fruit flesh and/or mango seed. Immatures
of mango seed weevil (Sternochetus mangiferae (F.)) occur in mango seed (but not
flesh) in most of Africa, Asia, Australia, the Pacific and the Carribbean (Waite,
2002), and are difficult to kill in situ with-out damaging the market quality of the
treated mangoes. Orchard-control measures and surveying are discussed by Hansen
(1991, 1993), Waite (2002) and Wittenberg (2007). The mango pulp weevil
(Sternochetus gravis (F.), syn. Sternochetus frigidus (F.)) occurs in India,
Bangladesh, part of the island of Palawan in the Philippines and a few other
regions in South-east Asia (Waite, 2002; Astridge and Baron, 2007c; Catindig and
Kong, 2007; Walker, 2007a). It causes severe damage to the fruit pulp only (de
Jesus et al., 2007).
Follett and Gabbard (2000) concluded that mango seed weevil does not
seriously affect mango yield or marketability. Nevertheless, the seed weevil is a
major quarantine concern for countries which have not recorded it or claim area
freedom, that is Middle Eastern countries and China (Waite, 2002). Seed and pulp-
attacking Lepidoptera pests are quarantine risks in some countries (Waite, 2002;
Walker, 2007b; Yarrow and Chandler, 2007). Entry of mangoes from countries
having mango weevil and other Sternochetus spe-cies may be restricted or
prohibited into countries free of these pests. Exten-sive surveying, sampling,
implementation of field control measures and/or area-freedom certification and
maintenance may be necessary for approval of market access (Johnson and
Heather, 1995; Waite, 2002). Disinfestation treatments that ensure weevils are not
able to reproduce may be acceptable when dosages for mortality damage fruit
excessively.

Fruit fly disinfestation


Tephritidae are the most important mango pests and occur wherever man-goes are
grown (Waite, 2002). Eggs are oviposited below the peel. The wound provides an
opening for microorganisms and scars the peel. Larvae feed and tunnel throughout
the pulp. Fruit flies infest tropical and temperate fruits. It is the risk to temperate
climate fruits and commodities produced in fly-free areas that has prompted the
development of quarantine restrictions and treatments for fruit fly hosts.

At present, quarantine treatments against fruit flies are not required for fruit
entering the European Union (EU), despite the large production of tem-perate fruit
in fruit-fly-free regions. Fly infestation has not been perceived as a threat because
winter temperatures throughout much of the region effec-tively prevent
establishment of the flies, despite geographical continuity with the distribution
range of the Mediterranean fruit fly (Ceratitis capitata (Wiedemann)). Canada does
not require fruit fly disinfestation of tropical produce for the same reason. Exotic
fruit fly pests could become established in southern Florida, Texas and California
because of their subtropical climate. The USA requires that mangoes be disinfested
by vapour heat, irradiation, hot water or hot air.
560 G.I. Johnson and P.J. Hofman

In Queensland, Australia, fly larvae infestation of mangoes in the mar-keting


chain is rare, despite the widespread occurrence of endemic species of fruit flies
(Bactrocera spp.). Preharvest control measures, and grading out of coloured fruit at
the packhouse, effectively eliminate infestation of most com-mercial
consignments; however, mangoes consigned to the Australian states in temperate
regions free of the flies must be disinfested against fruit flies to help ensure area
freedom of temperate-fruit-production areas (RSPM, 2004; Jessup et al., 2007).
When effective field control and grade-out of ripening fruit is in place, the
mandatory disinfestation of mature-green mangoes entering the exclusion zone is
probably unnecessary.
Potentially acceptable quarantine treatments that disinfest mangoes include
vapour heat, hot air, hot water immersion, irradiation, quick-freezing, combination
treatments and some miscellaneous treatments (Taylor et al., 2002; Ducamp Collin
et al., 2007). The major constraints in the development of treatments have been the
susceptibility of mangoes to heat, cold and irra-diation damage and O2 depletion,
and the extensive research and negotiation required to obtain market access
approvals to high-end markets (i.e. the USA, the EU and Japan). Treatments need
to be verified as non-damaging to a range of cultivars by fruit size by environments
likely to be encountered (Jacobi and Gowanlock, 1995; ISPM, 2007b). Treatments
that cannot be used because they lower fruit quality at dosages that kill pests are
methyl bromide fumigation (Spalding et al., 1977) and cold temperature storage
(Kane and Marcellin, 1978).

Vapour heat
Vapour heat treatment (VHT) involves heating air that is nearly saturated with
moisture, and passing the air stream across the fruit (Jacobi et al., 2001b). When
the temperature of the mango fruit is at or below the dew point of air, condensation
occurs on the fruit surface and rapidly heats the fruit by con-ductive energy
transfer. The core of the fruit next to the seed is heated to c. 45C for the required
time before cooling. Fruit have to be sorted for size before treatment because of
different rates of attaining the required core temperature.

Vapour heat is used worldwide to disinfest mangoes of fruit flies. Jacobi et al.
(2001b) list the VHT protocols approved for importation of mangoes into Japan
from the Philippines, Taiwan, Thailand, Australia and Mexico. Conditions range
from 43–47C pulp core temperature for 10 min to 6 h; however, the most common
treatment conditions are 46–47C for 10–30 min. Melon fruit fly (Bactrocera
cucurbitae Coquillett) immatures in mangoes from Okinawa were killed at 44 
0.3C core temperature for 3 h (Sunagawa et al., 1987). Taiwanese mangoes
infested with melon fly can be disinfested with vapour heat at 47.5C until the
centre pulp is >46.5C for 45 min (Kuo et al., 1987).

A VHT schedule was approved against Queensland fruit fly (Bactrocera


tryoni), in ‘Kensington Pride’, ‘R2E2’, ‘Keitt’, ‘Palmer’ and ‘Kent’ from Aus-tralia
for the Japanese market (AQIS, 2008b), which consists of a core tem-perature of
47C for 15 min. The United States Department of Agriculture,
Postharvest Technology 561

Animal and Plant Health Inspection Service Plant Protection and Quarantine
(USDA-APHIS PPQ) approved VHT as a quarantine treatment for Mexican fruit
fly (Anastrepha ludens (Loew)) and other Anastrepha species in ‘Manila’, and for
mangoes from Taiwan infested with oriental fruit fly (Anonymous, 1994a). Generic
guidelines for use of VHT in treating commodities for the USA market are
provided by the USDA-APHIS PPQ manual on vapour heat. Mangoes from
Taiwan imported into Australia must be treated until the pulp temperature has been
46.5C for 30 min (AQIS, 2008a).

Hot air
Hot or forced hot air systems also heat the air to 40–50C, but at a lower RH.
Relative humidity usually remains >50%, depending on ambient RH, but is never
high enough to produce condensation. Heat is transferred to the fruit by convection,
with no condensation of water on the skin (Gaffney and Arm-strong, 1990; Jacobi
et al., 2001b). Relative humidity should be high enough to prevent fruit desiccation
during treatment. Transfer of heat from the air to the skin is slow compared with
VHT. Mangan and Ingle (1992) reported that a mean centre pulp temperature of
>47C killed all stages of West Indian fruit fly, Anastrepha obliqua (Macquart), in
Mexican mangoes, and Sharp (1992) found a centre pulp temperature of >46C
killed all stages of Caribbean fruit fly, Anastrepha alletis (Loew), in Florida-grown
mangoes.

Hot water
Provided that fruit are not damaged, hot water immersion is environmen-tally safe
and efficient for killing mango pests. Use of hot water to kill fruit fly eggs and
larvae intensified in the USA when the Environmental Pro-tection Agency (EPA)
removed ethylene dibromide from the market as a chemical fumigant because of
health concerns (Anonymous, 1983). Sharp and Spalding (1984) showed that
mangoes could be disinfested of Caribbean fruit fly using hot water. The work led
to more studies in Haiti and a disinfestation method for West Indian fruit fly (Sharp
et al., 1988), as well as Mediterranean fruit fly and other Anastrepha spp. in Texas
and Mexico (Sharp et al., 1989a, b), Puerto Rico (Segarra-Carmona et al., 1990)
and Peru (Sharp and Picho-Martinez, 1990). Nascimento et al. (1992) developed a
hot water treatment for fruit flies in mangoes in Brazil. Hot-water-treated mangoes
may be imported into the USA from Mexico, Central America, South America and
the West Indies (Anonymous, 1994a). Typical treatments include 46.1C for 65
min for smaller fruit to 90 min for larger fruit (Jacobi et al., 2001b). Large
commercial hot-water-treatment facilities have been constructed, certified by the
USDA-APHIS PPQ, and used in Mexico, Central and South America, and the
West Indies. Generic guidelines for the use of hot water are provided by the
USDA-APHIS PPQ manual for hot water treatment.

In Australia, Smith (1992) showed that immersing five Australian mango


cultivars in 48C water for 30 min killed eggs and larvae of Bactrocera aquilo-nis;
however, ‘Kensington Pride’ is more sensitive to hot water than to vapour heat, so
the latter has been adopted for disinfestation of mangoes in Australia (Jacobi et al.,
1994). Grové et al. (1997) found that treatment of several cultivars
562 G.I. Johnson and P.J. Hofman

in hot water at 46.1°C for 90 min followed by refrigeration for 24 h did not damage
fruit, although some cultivars showed severe lenticel damage. Refrigeration of
‘Tommy Atkins’ fruit immediately after treatment resulted in scald development.
Weevils in ‘Alphonso’ mangoes from India were not killed when infested mangoes
were immersed in water at 48–52°C for up to 90 min and 54–70°C for up to 5 min
(Shukla and Tandon, 1985).
Compared with hot air treatments, hot water treatments can damage the skin,
partly because of rapid heat transfer from the water to the skin com-pared with
from the skin to the centre of the fruit. Damage includes skin scald-ing, lenticel
damage, cavities, white starchy areas in the flesh and delayed ripening (Jacobi et
al., 2001b). Several factors influence damage severity after heat treatment, for
example cultivar, temperature and duration (Jacobi et al., 2001b). Immature fruit
have low heat tolerance, and small fruit are damaged by heat more readily than
large fruit. Conditioning treatments (i.e. 37°C core temperature, for at least 12 h in
air) can reduce injury, and preharvest condi-tions, especially rainfall before
harvest, can increase skin damage (Esguerra and Lizada, 1990; Esguerra et al.,
1990; Jacobi and Wong, 1992; Jacobi et al., 1994, 1995; Jacobi and Giles 1997).
Better understanding of these influences could increase the commercial potential
for hot water disinfestation.
Hot water dips could pose human health risks. Sivapalasingam et al. (2003)
reported that an outbreak of Salmonella enterica that infected 72 patients from 13
USA states may have been due to contamination of hot-water-dipped mangoes
from a single farm in Brazil. No outbreaks were reported among consumers in the
EU of mangoes from the same farm, and the EU does not require hot water
disinfestation.

Irradiation
Irradiation involves γ rays (at <1000 Gy), X-rays, electrons and microwaves
(Thomas, 1986; Velasco and Medina, 2004; Follett et al., 2007; Moreno et al.,
2007). A 2005 FAO/International Atomic Energy Agency (IAEA) report indi-cated
that >20 irradiation facilities have been planned, constructed or reno-vated in ten
countries, some of which are mango exporters (Eustice, 2007). Radiation
treatments have been developed for fruit flies in mangoes from Florida, Mexico,
India and Australia. Von Windeguth (1986) treated mangoes with 76 Gy and
disinfested them of Caribbean fruit fly eggs and larvae. Third instar Mediterranean
fruit fly larvae in Mexican mangoes irradiated with 250 Gy did not emerge from
pupae, and 60 Gy applied to third instar Mexican fruit fly, and West Indian fruit fly
in Mexican mangoes prevented adult emer-gence (Bustos et al., 1992). Bustos et
al. (2004) recommended a generic dose of 150 Gy for control of Mexican fruit fly
(A. ludens), the West Indian fruit fly (A. obliqua), the sapote fruit fly (Anastrepha
serpentina) and the Mediterranean fruit fly (C. capitata) in mango. ‘Kensington
Pride’ mangoes infested with eggs and larvae of Queensland fruit fly and
Bactrocera jarvisi (Tryon) are disinfested with 74–101 Gy (Heather et al., 1991).

International guidelines for the use of irradiation as a phytosanitary measure


are available (ISPM, 2003), and recently a fast track process has been proposed as
an Annex to ISPM 28 (ISPM, 2008), which endorses irradiation
Postharvest Technology 563

at 70 Gy as a generic treatment to control Anastrepha spp. in fruit and vegeta-bles


by extrapolating work on mango by Bustos et al. (2004). Heather (2004) provides
generic guidelines for the development of irradiation protocols for disinfestation.
Fruits are never exposed to radioactive materials (Anony-mous, 1986) and most
modern treatment units use an electron beam process rather than a radioactive
source for irradiation.
Irradiation can be used for controlling seed weevil and lepidopterous pests in
fruit. Seo et al. (1974) reported that 206 and 329 Gy killed mango weevil in
Hawaiian mango. Thomas (1975) showed that 500 Gy killed all mango weevil
larvae and pupae and 750 Gy prevented adults from emerging from mangoes in
Africa. A dose of 500 Gy, however, did not disinfest ‘Alphonso’ mangoes of seed
weevil (Shukla and Tandon, 1985). Indian man-goes from approved packhouses
must be irradiated with a minimum of 400 Gy at an approved and certified
irradiation treatment facility using Cobalt-60 (APEDA, 2007). A quarantine
treatment of 300 Gy has been approved to ster-ilize mango seed weevil in mangoes
exported from Hawaii to USA mainland markets (Follett, 2004). Follett and Lower
(2000) demonstrated control of Cryptophlebia illepida (Butler), Cryptophlebia
ombrodelta (Lower) and Cryp-tophlebia illepida (Lepidoptera: Tortricidae), and an
irradiation quarantine dose of 250 Gy has been approved for Hawaiian mangoes.
The treatment also controls fruit flies (Follett, 2004).

USA regulations covering irradation are described in the Code of Federal


Regulations GPO Access (2008), revised annually (Wall, 2008), and this sum-
marizes approved treatments for a range of pests (EPA, 2002) (Table 15.5),

Table 15.5. Minimum absorbed dose of gamma irradiation required by USDA for specific
pests (Source: adapted from EPA, 2002).

Scientific name Common name Minimum absorbed dose (Gy)

Anastrepha ludens Mexican fruit fly 70


Anastrepha obliqua West Indian fruit fly 100
Anastrepha serpentina Sapote fruit fly 100
Anastrepha suspensa Caribbean fruit fly 70
Bactrocera cucurbitae Melon fruit fly 150
Bactrocera dorsalis Oriental fruit fly 150
Bactrocera jarvisi Jarvis fruit fly 100
Bactrocera tryoni Queensland fruit fly 100
Brevipalpus chilensis False red spider mite 300
Ceratitis capitata Mediterranean fruit fly 150
Cryptophlebia illepida Koa seed worm 250
Grapholita molesta Oriental fruit moth 200
Sternochetus mangiferae Mango seed weevil 300
All other fruit flies of the family Tephritidae which are not 150
listed above
Plant pests of the class Insecta not listed above, except 400
pupae and adults of the order Lepidoptera
564 G.I. Johnson and P.J. Hofman

many of which can infest mangoes. The USDA-APHIS PPQ manual on irradia-tion
provides generic guidelines. Irradiation was approved for the USA market as a
phytosanitary treatment for all fresh fruits and vegetables from all coun-tries in
2002. Effects of J-irradiation on mango fruit quality and disease control have been
reported (Mitchell et al., 1992; Moreno et al., 2006; Reyes and Cisneros-Zevallos,
2007; Wall, 2008). Only marginal disease control was obtained with ‘Kensington
Pride’ at the highest non-deleterious doses for mature-green fruit (300 Gy), with
additive effects of disease control treatments and irradiation on disease reduction
(Johnson et al., 1990a). Disease control may be more effective in cultivars with
greater tolerance of irradiation (van der Linde and Thord-Gray, 1986; Johnson et
al., 1990a). Other types of irradiation have been evaluated for mango disinfestation
but none has been adequately suitable.

Quick freezing
Quick freezing of mango, lowering the temperature to –17C and holding at –6C
or below for 48 h is used to disinfest mangoes for processing (Anony-mous, 1994a;
PPQ, 2007). The process is not approved for importing man-goes with seeds from
most of the West Indies, French Guiana, all countries outside of North, Central and
South America, Oceania, Hawaii, South-east Asia, the Philippines and the Republic
of South Africa into continental USA because mango weevil could be present
(Anonymous, 1994a; PPQ, 2007).

Fumigation
Fumigation is an ideal methodology for ensuring effective control when the
fumigant is effective and safe to use. Until 1994, New Zealand required fumi-
gation of mangoes from Australia, the Cook Islands and the Philippines using 33,
29 or 22 g/m3 ethylene dibromide at 10–15, 15.5–19.5, or 20C and above,
respectively, at normal atmosphere pressure (NAP) to disinfest man-goes of fruit
flies before entry. As part of the international phase-out of ozone-depleting
substances, the process was banned in 1994 (Anonymous, 1992; N.W. Heather,
personal communication, Brisbane, 1994) and most applica-tions as a fruit
fumigant have ceased worldwide. Methyl bromide was phased out completely in
the USA in 2005, but some emergency uses for quarantine applications may be
permitted, e.g. to destroy a serious quarantine pest in an imported consignment or
to meet official requirements of an importing coun-try (EPA, 2008). Mangoes
imported into Australia from countries where fruit flies occur must be fumigated
with 16–35 g/m3 ethylene dibromide for 2 h at 21–26C or above (Anonymous,
1985, 1988).
Phosphine is widely used as a fumigant of durable produce (grains and
tobacco). It provides effective control of fruit fly larvae and other pests in
temperate fruits under experimental conditions (Horn and Horn, 2004). However,
phosphine when mixed with water is highly explosive and the vapour is toxic to
humans, so prospects for utilization are not strong.

Miscellaneous treatments
CHEMICAL TREATMENTS. Postharvest chemical treatments using dimethoate are effective
against Queensland fruit fly with ‘Kensington Pride’ (Swaine et al.,
Postharvest Technology 565

1984). The treatment is required for Australian-grown mangoes entering all


Australian states except Queensland and New South Wales, but is under review.
The USA and the EU do not allow the use of chemicals to disinfest mangoes.

NATURAL PRODUCTS. The short shelf life of mango and the high level of insect mortality
required obviates the use of natural products for disinfestation. Suhaila and Halim
(1994) reported the potential of low toxicity, insecticidal compounds from edible plants
that may be effective for topical application to harvested fruit. Extracts of black pepper
(Piper nigrum) were particularly active in laboratory tests against vinegar fly
(Drosophila melanogaster (Meigen)).

ATMOSPHERES. CA and MA regimes could have potential for disinfesting man-goes, but
there has been less interest in the technology because heat treatments and irradiation are
faster (Ke and Kader, 1992; Yahia and Tiznado-Hernandez, 1993; Yahia and Vazquez-
Moreno, 1993; Yahia, 1994; León et al., 2000). Treat-ments are limited to regimes
which do not adversely affect ripe fruit quality. León et al. (2000) found that CA of 1%
O2 and 30 or 50% CO2 disinfested ‘Manila’ mangoes of A. obliqua, but damage (as
spongy tissue) was unaccept-ably high.

Shrink-wrapping has been ineffective as a quarantine treatment to disin-fest


mangoes of fruit fly immatures. Gould and Sharp (1990) reported that the time
needed to disinfest Florida-grown mangoes infested with Caribbean fruit fly eggs
and larvae exceeded the shelf life of wrapped mangoes.

COMBINATION TREATMENTS. Serial applications of two or more treatments, which alone


do not achieve quarantine security, have been used to disinfest mangoes. Seo et al.
(1972) reported that eggs and larvae of Mediterranean fruit fly, orien-tal fruit fly and
melon fly were killed in mangoes immersed in water at 46.3C for 120 min and then
fumigated with ethylene dibromide. Lin et al. (1976) reported that all oriental fruit fly
and melon fly larvae in Taiwan-grown man-goes were killed when fruit were immersed
in 48–50C water for 120 min, hydrocooled, dried and cooled, and then fumigated with
ethylene dibromide.
Controlled Atmosphere/Temperature Treatment System (or CATTS)
technology applies a short heat treatment in a low O 2/high CO2 environ-ment, and
controls quarantine insect pests while maintaining commodity quality (Mitcham,
2007; Neven, 2008). Trials using CATTS with mangoes have been conducted in
Australia with promising results. Varith et al. (2007) evalu-ated a microwave-
vapour heat treatment (MW-VHT) disinfestation technol-ogy for mangoes: the
microwave component for pre-heating and the VHT component for the holding
process. Temperatures of 46–55C and holding times of 2–20 min effectively
disinfested fruit of oriental fruit fly eggs without effects on physico-chemical
parameters, compared to untreated fruit. There was less heat damage compared
with conventional VHT only fruit. MW-VHT shortened the process time by 90%
compared with the conventional VHT.

PACKAGING. Some markets, for example Japan and the USA, require that fruit must be
packed into insect-proof packages following disinfestation to preclude
566 G.I. Johnson and P.J. Hofman

reinfestation during transportation or storage. The disinfestation facility feeds fruit


into an insect-proof area within which waxing (optional), grading and packing
occur.

15.7 Preparing Fruit for Market

Surface coatings

Surface coatings are used to improve fruit appearance and to alter gas per-meability
to reduce moisture loss or retard ripening. Commercial use of sur-face coatings on
mango fruit needs to be considered carefully because of the fine balance between
beneficial and undesirable effects on fruit quality. Neg-ative effects of coatings
include reduction in chlorophyll loss (Fonseca et al., 2004a), anaerobic conditions
and off-flavours (Amarante and Banks, 2001) and skin damage, possibly due to
cytotoxic reactions with other components in the coating formulation (Bower et al.,
2003). Generally, coatings have less effect on delaying ripening during cold
storage, compared with extending the shelf life at typical ripening temperatures
(Amarante and Banks, 2001). Less significant effects are observed in more mature
and in ripening fruit. Coatings often delay skin colour change rather than softening,
which increases the risk of soft, green fruit with less consumer appeal.

Coatings are generally emulsions of synthetic (e.g. polyethylene) or nat-ural


(e.g. polysaccharides, carnauba, beeswax, etc.) origin. Surface coatings containing
waxes, oils (e.g. carnauba, beeswax, etc.) and resins (e.g. shellac) have a greater
effect on limiting water loss then reducing O2 and CO2 perme-ability, compared
with those containing polysaccharides, (e.g. those based on cellulose) (Amarante
and Banks, 2001). Formulations based on shellac result in a shinier appearance
than those based on carnauba wax and polysaccharide-based waxes (Baldwin et al.,
1999; Hoa and Ducamp, 2008).
Factors other than coating formulation can affect fruit gas permeability, i.e.
cultivar, variations in skin permeability between fruit, inconsistency in coating
thickness during application, interference from water during appli-cation causing
coating cracking and coating thickness and evenness-of-spread over the fruit
surface. The effect of coating on fruit quality can vary with holding conditions
because of larger temperature effects on respiration rate than on coating
permeability.
Shorter and Joyce (1994) found commercially formulated Avocado and
Passionfruit Wax, a polyethylene and shellac emulsion, and Technimul 9122 Wax,
a polyethylene-based emulsion, were acceptable with ‘Kensington Pride’ mango,
while Peach Wax, a polyethylene-based emulsion and starch solution was
unacceptable. With Peach Wax, deleterious modified atmosphere effects on colour
development, softening and flavour were obtained (El Ghaouth et al., 1992b;
Shorter and Joyce, 1994). Coating ‘Tommy Atkins’ mango with a carnauba-based
coating and BeeCoat (based on beeswax) reduced water loss, shrinkage,
chlorophyll breakdown, CI and decay after cold storage, and Bee-Coat also
reduced red lenticel discoloration (Feygenberg et al., 2005). With
Postharvest Technology 567

‘Tommy Atkins’, polysaccharide and carnauba-based coatings modified the


atmosphere within the fruit and reduced decay, but only the polysaccharide-based
coating delayed ripening (Baldwin et al., 1999). The carnauba-based coating
significantly reduced water loss compared with the polysaccharide-based coating
treatments; carnauba-based coatings result in lower water per-meability and higher
O2/CO2 permeability.
Coatings may reduce surface defects. Excessive water loss is associated with
increased skin CI in avocado and mango, and carnauba-based coatings reduce CI in
cold-stored mangoes (Bower et al., 2003). In this study, the carnauba-based coating
contained numerous holes, which allowed respiration gas exchange (thereby
preventing anaerobic respiration), while still providing efficient control of water
loss. Surface coatings may also reduce sapburn, skin browning and lenticel damage
(Shorter and Joyce, 1994), but incorporating these potential benefits into
commercial systems may be difficult.
Waxes should be applied by roller brushes in a specifically designed wax
applicator or by very light hand application. Dipping fruit in a wax emulsion is not
recommended. A uniform flow of fruit through the wax applicator must be
maintained to prevent uneven wax application. Fruit should be dry before entering
the wax applicator, otherwise foaming of water-emulsion waxes may occur.
Brushes on the wax applicator need to be completely satu-rated with the wax
mixture before any fruit passes over them. Complete cov-erage of the entire fruit
surface is essential. Patchy application can be caused by insufficient wax, too few
brushes following application (minimum of six brushes required), poor and/or
inadequate drying facilities, and overload-ing of the unit. Brushes should be kept
soft with regular washing with hot water.

Packaging

Packaging provides conveniently sized carriage units for product, protects


individual fruit from contact rub and compression damage, and excludes dirt, pests
and contaminants. McGregor (1987) and Hilton (1994) discussed key aspects of
packaging for tropical produce. Packaging is also a marketing tool. Design and
colours of symbols and text on carton exteriors portray a marketing image.
Manufacturers of consumer products exploit packaging to great advantage (along
with advertising) to increase both first-time and repeat sales. Marketing and design
professionals may be involved in the development and customer evaluation of
product packaging. Cultural pref-erences need to considered, e.g. use or avoidance
of red for some Asian mar-kets. Packaging is the external face of brand loyalty.
Consistent product performance and quality is the core.

Some constraints to packaging may be specified by market regulations,


including carton dimensions and labelling requirements. Country of origin,
cultivar, grower, packing shed, market agent, count (number per carton and weight
range) and class may be required. The word ‘mangoes’ should be clearly visible
(Anonymous, 1993). The information appears on the narrow
568 G.I. Johnson and P.J. Hofman

sides of the cartons. Storage and product use information can also be printed on the
cartons. Many QA systems require adequate labelling linked to appro-priate record
keeping for plate-to-farm traceability. Clear labelling facilitates correct delivery,
allows immediate buyer recognition of product profile and ensures maintenance of
accurate sales records. An exporting country may find it of value to identify
individual packers by barcoding or numbers stamped on cartons, so that sources of
faulty packaging can be traced. Some countries also use date codes which enable
exporters to determine the fresh-ness of the produce at the point of export and
evaluate an importers’ capacity to achieve adequate turnover of the fruit without
prolonged storage. It also provides invaluable feedback on the efficiency of the
total distribution chain.
Cartons used for export should be clean, strong, unbroken and new. The water
absorption capacity of the material should be evaluated as excess absorption will
lead to collapse on the pallet. The cartons’ strength will depend on the starch used
by the manufacturer, the outer liner and the direc-tion and numbers of fluting in the
carton (Anonymous, 1994b). There is increasing pressure in the EU for recyclable
packing material. Cartons that are recyclable should be marked with the
appropriate international symbol. Returnable plastic crates are increasingly being
used for domestic trade, but the return cost would make this less profitable for
international trade.

Inspection

In some countries, independent inspectors check the fruit prior to palletizing to


ensure that the relevant marketing, residue and phytosanitary standards have been
met. Fruit for Japan is disinfested under the supervision of a Japa-nese inspector.
Further inspections are usually made at the port of exit. Some exporting countries
require a declaration by the grower to ensure that fruit will comply with the
standards specified by importing countries.

Palletizing

Handling mangoes on pallets allows convenient movement of large volumes of


fruit. McGregor (1987) described critical features and arrangements for loading.
The disadvantages of pallets for export are the cost, lower numbers of cartons per
sea container and loss. Some domestic markets have pallet share systems. Relevant
markets and transporters should be consulted con-cerning required pallet
dimensions and appropriate access for fork-lift sys-tems. The correctly sized pallet,
for example as designated by the ISO, which is designed to fit snugly into a
standard sea container, should also be used for the local market.

Precision stacking with each box fitted exactly on top of the one below
minimizes risk of damage. Collapsed or lopsided pallet stacks have usually been
due to careless stacking and/or loose placement in the shipping con-tainer. Pallet
slats should not block ventilation holes in the cartons. Cartons
Postharvest Technology 569

should be register-stacked so that ventilation is continuous. Link sheets, which bind


the cartons together at intervals, should also be designed to ensure con-tinuous
ventilation through the pallet. In the cold room, pallets should not be stacked
against a wall or placed directly against each other (Boelema, 1987).

Precooling

Precooling removes field heat from the product and lowers the temperature to that
required for ripening, transportation or storage. Precooling also reduces the cooling
demand on any in-transit cooling system. Precooling concepts and systems are
described by Thompson et al. (2002). Forced-air cooling systems efficiently and
rapidly remove field heat, and are preferred for bringing fruit to storage
temperature. High RH systems are preferred as they reduce fruit water loss.
Hydrocooling can increase the risk of infection by wound pathogens (i.e. Rhizopus
spp.) and are less effective with large fruit. Kitinoja and Kader (2003) describe
low-cost cooling facilities for use in devel-oping countries.

Ethylene and ripening

Induction of ripening is routinely employed with mangoes. There are effec-tive low
technology methods involving calcium carbide (releases acetylene which mimics
ethylene) or the leaves of particular trees (Lizada, 1994). More sophisticated
systems include generation of ethylene from ethanol using catalytic conversion,
pure ethylene gas, or a mixture of ethylene and an inert gas (CO 2 or N2) to reduce
the risk of explosion with 3–30% ethylene in air (Reid, 2002). A number of
automatic ethylene control systems are available (PDS, 2008) to maintain ethylene
concentrations within required limits.
Climacteric fruit have differing sensitivities to ethylene. ‘Kensington Pride’
mango is sensitive to concentrations as low as 0.01 Pl/l (O’Hare et al., 1994).
Ripening is enhanced with concentrations up to 5–10 Pl/l, with very little ben-efit
at >50–100 Pl/l (Nguyen, 2003). There is more yellow colour on the ripe fruit
when ripened at 20C with 10 Pl/l ethylene for 3 days compared with no ethylene,
resulting in a more attractive appearance. Also, diseases are gener-ally less in these
fruit (Table 15.6), presumably because fruit ripen more quickly with less time for
disease development. Good ethylene treatment can improve presentation
appearance and increase saleable life (defined as the days from when the fruit reach
at least 60% yellow skin colour to when the fruit had lost saleability because of
disease) (Ledger et al., 2002a).
Ethylene can also reduce quality if not used appropriately. Ripening
‘Kensington Pride’ fruit at <18C with ethylene can result in soft fruit with less
yellow skin colour, most likely because ethylene stimulated softening to a greater
extent than chlorophyll loss (Nguyen, 2003). Ripe fruit disease can also be greater.
These effects can be aggravated with concentrations above 100 Pl/l (Fig. 15.5).
‘Kensington Pride’ fruit must be cooled to <24C before
570 G.I. Johnson and P.J. Hofman

Table 15.6. Days for fruit to reach the eating soft stage (days to ripe at 20C),
percentage weight loss/day and percentage of fruit surface area affected by stem
rots in ‘Kensington Pride’ mango fruit treated with 25 Pl/l 1-methylcyclopropene (1-
MCP) for 14 h at 20C followed by exposure to 100 Pl/l ethylene for 24 h at 20C.
Fruit were then ripened at 20C. Means followed by the same letter in each column
are not significantly different (P >0.05) (Source: Hofman et al., 2001).

Treatment Days to ripe Weight loss (%)/day Stem rots (%)

Untreated 13.6 b 0.3 a 9.6 b

Ethylene 7.9 a 0.4 b 1.3 a


c
1-MCP 18.7 c 0.3 a 18
d
1-MCP + ethylene 18.2 c 0.3 a 25.8

70
24 h
72 h
60

50

40
Green colour (%)

30

20

10

5
2
1

0
0 10 100 1000 0 10 100 1000 0 10 100 1000
Treatment at 15°C Treatment at 20°C Treatment at 25°C
Ethylene concentration (μl/l)

Fig. 15.5. Effect of ethylene concentration and time in ethylene and ripening
temperature on the percentage skin surface area with green colour of ‘Kensington
Pride’ mangoes at eating soft; least significant difference = 5.16 (P <0.05) (Source:
Nguyen et al., 2002). Note the increased green colour on the skin of ripe fruit with
lower ripening temperature and higher ethylene concentrations and duration.

the start of ethylene treatment; otherwise, skin spotting can develop (Ledger,
2003a). Ripening at 18–22C is recommended for maximum yellow skin colour,
less disease and higher flavour volatiles (Hofman, 1997; Lalel et al., 2004).
The relatively high respiration rate of ripening mangoes can result in CO 2
accumulation in the ripening room, particularly if the room is full and there is poor
ventilation. Carbon dioxide concentrations up to 5.3% have
Postharvest Technology 571

been recorded in ripening rooms (Ledger, 2007), which can cause more green
colour and a dull appearance on the ripe fruit (Nguyen, 2003). Ripening room CO 2
concentrations should be maintained at <1% with adequate ventilation to minimize
fruit quality loss (Kernot et al., 1999; Ledger, 2007).
Accidental exposure of mangoes to ethylene and its analogues from adja-cent
ripening rooms, exhaust fumes from internal compression engines or wound
ethylene produced from damaged/ripening fruit can cause prema-ture ripening.
Various systems can remove unwanted ethylene, for example oxidizing
mechanisms such as potassium permanganate either in sachets or in ethylene
scrubbing units in storage rooms, catalytic oxidizers or ozone-based systems (Reid,
2002). Smartfresh™ (active ingredient 1-methylcyclopropene; 1-MCP) is a
relatively new approach for preventing undesirable ethylene effects. 1-MCP is a
structural analogue of ethylene and irreversibly binds to the ethylene receptors in
the plant, thus preventing ethylene-initiated ripen-ing. Ripening re-commences as
additional ethylene-receptor sites are pro-duced in the fruit (Blankenship and Dole,
2003). Generally, Smartfresh™ treatment is applied in well-sealed cold rooms or
plastic tents as soon as pos-sible after packing. 1-MCP concentrations of 250–
200,000 Pl/l for 12 h are optimum for delaying ripening (Jiang and Joyce, 2000;
Hofman et al., 2001; Adkins et al., 2002; Penchaiya et al., 2006), although most
reports state 250– 1000 Pl/l. 1-MCP treatment completely negated any effect of
subsequent eth-ylene on ripening, and can almost double the days to eating soft
compared with ethylene-treated fruit ripened at 20C (Table 15.6) (Adkins et al.,
2002). However, the 1-MCP effects were less in more mature fruits (Alves et al.,
2004), and ethylene exposure before 1-MCP will negate any 1-MCP benefit
(Adkins et al., 2002). Any beneficial effects of 1-MCP also appear to be less with
longer-term storage (Hofman, unpublished results). 1-MCP treatment can cause
more disease on ripe fruit, because the longer days to ripen allows more disease
development (Hofman et al., 2001). Sourcing fruit from well-managed orchards
can help minimize this effect (Adkins et al., 2005).

For some domestic markets, on-farm treatment of mangoes with ethyl-ene is


used to ensure that fruit have more attractive colour when they are displayed at the
wholesale market 48–72 h after dispatch from the farm. This practice improves
returns as fruit can be delivered to retail outlets ready-to-eat. Ethylene induction of
ripening is undesirable for more distant markets because fruit arrive at the market
too ripe for sale, with greater risk of bruis-ing and disease.

15.8 Pre- and Post-shipping Storage


Cool storage

Cool storage is important when delivery time from harvest to the consumer exceeds
the typical ripening time (5–10 days). The ideal storage temperature is dictated by
the risk of CI, fruit ripening and disease development during stor-age, and storage
time. CI is first noted as greying of the skin, which intensifies
572 G.I. Johnson and P.J. Hofman

with lower temperatures and longer duration (Phakawatmongkol et al., 2004;


Suresh et al., 2004). In more severe cases flesh discoloration and abnormal
ripening can occur. CI development can occur at regimes of 3–12C (Sadasi-vam el
al., 1971; Thomas and Oke, 1983; Chaplin et al., 1986 a, b, 1991a, b; Smillie et al.,
1987; Thomas and Joshi, 1988; Medlicott et al., 1990b). Longer storage times
require greater care with temperature selection, the quality of the fruit being stored
and conditions before and after harvest. Storage should be for the minimum period
necessary. The following factors affect the opti-mum storage temperatures and
durations:
● Genetic differences – cultivars differ in chilling sensitivity (Phakawat-mongkol
et al., 2004).
● Maturity – less mature fruit ripen more slowly at a given temperature, and are
more prone to CI and other storage-related disorders (Medlicott, 1985;
Medlicott et al., 1987, 1990 a, b; Oosthuyse, 1993). Such fruit may not soften
at all when exposed to temperatures that are suitable for stor-age of more
mature fruit. In South Africa, adequately mature fruit can be stored at 8–10C
for 21–28 days (Oosthuyse, 1994). Placement in cold storage without delay
and post-storage exposure to temperatures that promote ripening (e.g. 20C)
are important preconditions for success.
● Duration of storage – ‘Kensington Pride’ mangoes can be stored at 10C for 3
weeks or at 7C for 2 weeks, after which skin colour development can be
affected (McLaughlan and Wells, 1994). Generally, the shorter the storage
time, the greater the tolerance to storage temperatures outside the 10–12C
range.
● Delays between harvest and cold storage, and ripeness stage – the longer the
delay between harvest and cold storage, the greater the risk of ripen-ing during
storage. This applies particularly for more mature fruit. If prolonged, a delay
may render refrigerated storage ineffective in pre-venting fruit from becoming
soft during transit, despite the apparent ab-sence of softening on dispatch
(Oosthuyse, 1994). Fruit should be picked, packed and placed in cold storage
within 24 h. For fruit that have rip-ened, storage temperatures of less than 8C
can be used for up to 21 days without deterioration in quality during storage;
however, the fruit will deteriorate rapidly after removal from storage (Van
Straten and Oost-huyse, 1994). Some cultivars may be more sensitive to ripe
storage, since ‘Kensington Pride’ fruit at the mid-climacteric stage will start to
lose appearance after 3 days at 10C because of increased disease and mild CI
(H. Nguyen et al., 2004).

● Disease load and fruit tolerance of disease – certain mango cultivars are very
tolerant of postharvest pathogens (e.g. see Hassan, 2007). The con-ditions
under which mangoes are grown may be unfavourable for infec-tion. In these
situations, storage temperatures can be higher to reduce the risk of CI.

Development of CI in mango and other fruits is closely associated with


antioxidant activity (Arafat, 2005; Kondo et al., 2005). Mango fruit held at 6C for
10–20 days had lower antioxidant activity in the skin compared with fruit
Postharvest Technology 573

stored at 12C. Application of several jasmonate derivatives before storage reduced


CI at 6–7C (González-Aguilar et al., 2000; Kondo et al., 2005), pos-sibly through
an antioxidant mechanism. Other chemical treatments can also reduce CI.
Polyamines occur naturally in fruit and decrease during storage under chill-
inducing conditions, and application before storage can reduce CI (Nair et al.,
2003). Salicylic acid appears to be involved in cell wall stability. Application of
methyl salicylate, which breaks down to sali-cylic acid, significantly reduced CI in
‘Zill’ mangoes stored at 7C (Han et al., 2006). 2,4-D can also reduce mango CI,
possibly through interaction with natural plant hormones and antioxidant levels in
the fruit (Wang et al., 2008). Some of these treatments could have commercial
application, but may have residue implications.

Decay is a major limitation to storage life. The incidence of postharvest decay


on fruit that ripen after refrigerated storage is positively related to the duration of
storage and the extent of ripening during storage (Oosthuyse, 1991, 1992, 1994).
Disease development after post-storage exposure to ripen-ing temperatures can be
reduced by minimizing the shipping period and by storing fruit at temperatures that
inhibit softening and ground skin colour development. If CI occurs, disease
develops earlier and will be more exten-sive (Oosthuyse, 1990).

Controlled and modified atmosphere storage

Decreasing the O2 and/or increasing the CO2 concentration can have several
advantages with respect to storage (i.e. reduced ethylene production, better flavour
retention, slowing softening and green skin colour loss and reduced CI)
(Thompson, 1998; Yahia, 2006). However, if the O 2 concentration is too low
(dependent on cultivar, storage temperature, fruit maturity and ripeness stage)
anaerobic respiration will commence, with associated production of ethanol and
acetaldehydes, leading to off-flavours and physiological disor-ders (Bender et al.,
2000). Atmosphere modification generally has less benefit for tropical fruit
compared with temperate fruit, but does have commercial potential for sea freight
to distant markets. Atmosphere control can be active or passive, or combinations of
the two. Surface coatings (see Surface coatings section under 15.7 Preparing Fruit
for Market, this chapter) also provide modified atmosphere inside the fruit.

With CA systems, O2 and CO2 concentrations are actively monitored and


controlled by injecting N2 and CO2, or bleeding air into the container as required.
In more passive systems, such as the MaXtrend® system (Maxtend, 2008) fruit
respiration directly lowers O2 concentrations, and its concentra-tion is monitored
and manipulated by venting as required. In some cases, the container is flushed
with N2 at the start of storage to rapidly establish the desired atmospheres. Excess
CO2 is absorbed with hydrated lime. In MA sys-tems, atmospheres are modified by
placing a semi-permeable membrane around the fruit (usually plastic film), and
relying on fruit respiration to modify the atmosphere.
574 G.I. Johnson and P.J. Hofman

McLauchlan and Barker (1994) suggested 4% CO2 and 2–4% O2 for CA


storage of ‘Kensington Pride’ mangoes at 13C, and recommended further research
on atmospheres <2% O2 and >10% CO2. Oxygen had the biggest effect on
retarding skin colour and softening, with significant retardation when decreasing
from 4 to 2%. Subsequent research suggested that concen-trations of 1.5–2% may
be more effective in retarding softening, although these concentrations may
increase the risk of off-flavours. ‘Tommy Atkins’ and ‘Haden’ can tolerate 2–3%
O2 for 2–3 weeks at 12C, but lower concentra-tions were not tested (Bender et al.,
2000). In ‘Kensington Pride’ there was little additional capacity for CO 2
concentrations between 6 and 10% to retard softening or loss of green colour
(McLauchlan and Barker, 1994), although there may be some benefit for storage of
1–2 weeks at concentrations >10% (Bender et al., 2000). For ‘Delta R2E2’
mangoes 3% O2 and 6% CO2 have been recommended (Lalel and Singh, 2006);
however, this is a firm-fleshed culti-var, which can perhaps tolerate higher O2
concentrations to improve vola-tiles, compared with the softer ‘Kensington Pride’.
Longer storage times with CA could cause higher disease levels (Johnson et al.,
1990b), higher acidity in the flesh at eating soft (McLauchlan and Barker, 1994;
Bender et al., 2000) and slower loss of green colour compared with non-stored fruit
(Bender et al., 2000). For cultivars that normally have higher acidity, CA-stored
fruit may need to be ripened for several more days to lower acidity.
Cold storage and CA can reduce volatiles production following ripening at
room temperature. As the skin CI severity increases with decreasing stor-age
temperature, total volatiles production appears to decrease (Singh et al., 2004). In
‘Kensington Pride’ and ‘R2E2’, CA storage significantly reduces total
concentrations of aroma volatile compounds compared with air-stored fruit,
irrespective of storage period between 24–38 days (Singh et al., 2004; Lalel and
Singh, 2006). Decreasing the O2 concentration from 3 to 1% at 6 or 8% CO 2 or
increasing CO2 concentration from 6 to 8% significantly increased most of the
monoterpenes, including terpinolene. Cold-stored fruit are known to have less
aroma than those ripened without storage.
Fruit can tolerate short periods with <1% O2 or >20% CO2 (Yahia, 2006).
This has been utilized for insect disinfestation (see Disinfestation section under
15.6 Packhouse Measures, this chapter). Mango can tolerate low O 2 concentrations
for 5 days at 20C (Yahia, 1994). These short-term CA treat-ments may improve
storage life or reduce CI during subsequent cold storage without atmosphere
modification; this has been noted with avocado (Truter and Eksteen, 1987; Pesis et
al., 1994). Preliminary investigations suggested little benefit of 20–60% CO2 for
1–8 days before cold storage of ‘Kensington Pride’ (Meiburg et al., 1998).
The optimum conditions for storage of each product to provide maximum
storage life without quality loss must be determined taking into account cultivar,
season and growing conditions. Measurement of chlorophyll fluo-rescence has
been used to monitor product performance under CA, with ad-justment of gas
conditions to achieve the optimal storage-life/quality balance. Changes in
chlorophyll characteristics and therefore chlorophyll fluores-cence under CA occur
before CI symptoms are obvious (DeEll and Toivonen,
Postharvest Technology 575

2003a). Thus monitoring changes in chlorophyll fluorescence characteristics can


provide advance warning of the potential for CI, and allow adjustment of storage
conditions to minimize its development (DeEll and Toivonen, 2003b). This concept
has now been marketed as ‘HarvestWatch’ (Harvest-Watch, 2008), and some
preliminary success has been obtained with apples (Stephens and Tanner, 2005;
DeLong et al., 2007).
Modified atmosphere packaging (MAP) generally cannot reliably achieve the
low O2 concentrations required to significantly delay softening without damaging
the fruit, but MAP can still have beneficial effects relative to the costs of CA (Pesis
et al., 2000; Rosa et al., 2001; Singh et al., 2001; Castro et al., 2005; Yahia, 2006).
However, MAP can reduce quality if the cultivar/holding temperature/film
permeability/storage time combination is not optimal (Sornsrivichai et al., 1989),
resulting in anaerobic conditions and off-flavours. Excess moisture retention inside
the bags can increase disease problems (Joyce and Patterson, 1994). Special films
have been developed with higher water vapour transmission rates (Pesis et al.,
2000) or moisture absorption materials can be included. Ethylene absorption
sachets can reduce chloro-phyll loss and red discoloration around the lenticels
(Rosa et al., 2001).
MAP reduces weight loss (Singh and Janes, 2001; Bower et al., 2003), which
maintains saleable weight, but may also reduce CI. There may be a direct relation
between these, since weight loss can contribute to CI in avo-cado, and the
reduction in CI obtained in mango through the use of wax coatings has been
attributed to the same mechanism (Bower et al., 2003). Pesis et al. (2000)
considered that lenticel discoloration is a symptom of mild CI, and noted that MAP
reduced the red coloration around the lenticels in the blushed area and the green
coloration around the lenticels in the green area of ‘Keitt’ mangoes. Less lenticel
spotting occurs in ‘Kensington Pride’ man-goes stored under MAP (Yuen et al.,
1993). It is not clear whether the reduc-tion in lenticel damage was due to CO2/O2
or humidity modification.
Cultivar, film type, number and mass of fruit per package, temperature, RH,
time of storage, maturity of the fruit and production conditions are important for
developing MAP systems (Brecht et al., 2003; Yahia, 2006). Important challenges
are the differential effects of temperature on fruit respi-ration and film
permeability, resulting in differing gas concentrations around the fruit as
temperature fluctuates. Success with MAP depends on a consistent or at least
predictable cold chain removal of the plastic film before significant temperature
fluctuations are likely to occur and using MAP films that are unlikely to cause
anaerobic conditions within the temperature range experi-enced in the cold chain.
Brecht et al. (2003) suggested an approach to designing flexible CA/MA systems
to account for variations in the cold chain.

15.9 Transport
Transportation of tropical fruit and vegetables has been reviewed by McGregor
(1987) and Thompson (2002). For local markets (<3 h access), transportation of
fruit in non-refrigerated carriers is feasible, particularly if the fruit has
576 G.I. Johnson and P.J. Hofman

been precooled and transported at night with few stops. Fruit must be shel-tered
from direct sun and rain. For sea export, fruit must be cooled to the required vessel
carrying temperature (or within 2C thereof) and the cold chain must be maintained
until the fruit is displayed for purchase.
Some retailers prefer fruit that are ready to eat within 1–2 days of receipt. In
these situations, the ideal scenario is to ripen the product as close as possible to the
retail outlet to minimize physical damage to the soft fruit. However, where end-
market location and transport arrangements allow delivery to market within 3 days,
ripening on-farm has advantages by reducing costs for growers, and extending
ownership of the product. Transport time is a major consideration for determining
optimum systems (Ledger, 2003b) (Table 15.7).
When the export dispatch facility is >1 h away from the packhouse, the
following road transport recommendations apply:
● The refrigerated truck should be clean and in good mechanical condi-tion. The
insulation and floor should be in a sound condition, the door seals must be
intact and the doors must close very tightly.

Table 15.7. Ripening and transport recommendations for ‘Kensington Pride’ mango within
Australia, to cater for the ‘ripe-for-tonight’ programme of major retail chains (Source:
Ledger, 2003b).

System Aim Recommended handling conditions

System 1 – ripen To deliver uniformly backward Precool fruit to transport temperature


at market fruit to the market destination within 12–15 h of packing
and then use ethylene to ripen Transport at 12–16C for trips of 1–
fruit ready for retail sale. 2 days and 12C for longer trips
Temperature is managed Ripen at the market using 10 ppm
through the chain to prevent ethylene for 2–3 days at 18–20C
mixed ripening and to avoid Continue to hold fruit at 18–20C
temperatures >22C. This is the until ready for sale
preferred system for Northern Store at 10–12C to slow ripening
Territory and northern Western for a maximum of 3 days
Australia growers send-ing
>2000 km to market
System 2 – ripen To deliver fruit to the market Precool to 18–20C within 12–15 h
on farm destination ready for retail of packing
sale within 1–2 days. Fruit are Ripen using 10 ppm ethylene for
ripened evenly using ethylene 2–3 days at 18–20C
to colour stage 3 (30–50% Hold at 18–20C until colour stage
yellow) before transport and 3 (30–50% yellow)
temperature is managed Transport at 12–16C for trips of
through the chain to avoid high 1–2 days and 12C for 3–4 day
temperatures >22C. Ripening trips
on farm is not recommended Hold at market at 18–20C until
for transport times >4 days ready for sale
Store at 10–12C to slow ripening
for a maximum of 3 days
Postharvest Technology 577

● The refrigeration equipment must be correctly set on air delivery and must be
calibrated for each journey. Equipment needs to function reli-ably and receive
regular servicing. Air should be delivered at the set point and fluctuations
should not exceed 0.5C from set point. Refriger-ated vehicles should be
fitted with temperature loggers monitoring the delivery air, and with a digital
display on the outside of the box. Refrig-erated vehicles are not usually
designed for, or capable of, lowering fruit temperatures so the fruit must be at
the relevant shipping temperature when loading.

● Because of the shorter time involved, air-transported fruit may have less
stringent temperature requirements than sea-export fruit. Airlines carry-ing
cargo may need to be consulted concerning the normal hold tem-peratures in
their aircraft.
● Sea-export fruit should be held under refrigeration until loading. Sea transport
can be in refrigerated vessels, with entire refrigerated decks filled with pallets,
or in sea containers, each of which is linked to a cen-tral ducted refrigeration
system in refrigerated container vessels. Alter-natively, integral containers
with their own individual cooling systems or integral CA containers may be
used.
Close temperature monitoring on the vessels is essential. By monitoring
delivery air temperatures (DAT) and return air temperatures (RAT), it is pos-sible
to assess whether fruit is heating up due to respiration or inadequate precooling,
and to take necessary steps (Anonymous, 1989; Eksteen, 1990). While most
refrigerated container vessels monitor individual container air temperatures,
including DAT and RAT, it is sometimes advisable to include additional
temperature loggers which can measure air and fruit pulp tem-peratures for an
entire journey.
The sea-freight component is generally the most time-consuming part of the
whole field-to-supermarket voyage (for example see Table 15.8). Essen-tial
activities before and after transport can be significant, for a relatively perishable
product like mango. Minimizing time delays in each component of the distribution
chain is important. To reduce product deterioration,

Table 15.8. Typical packing and shipping schedules for mangoes consigned by sea
to the EU from South Africa.

Operation Days required

Picking and packing 1


Precooling and accumulation of load 4
Transport to port 2
Port handling and accumulation of load 3
Voyage time 17
Discharge handling 1
Transport and distribution 2
Total 30
578 G.I. Johnson and P.J. Hofman

producers and marketers should encourage training in perishable product handling


and QA systems for personnel from trucking, sea-freight and air-freight companies
who are responsible for loading, unloading and maintain-ing storage facilities.

Co-shipment or storage with fruit or flowers that produce high levels of


ethylene can cause unanticipated triggering of mango ripening. Co-shipment with
papaya (Carica papaya) increases mango ripening (O’Hare et al., 1994).
Conversely, co-shipment of carambolas (Averrhoa carambola) with mangoes
caused ripening of the carambolas. The development of specialized packaging
materials to eliminate extraneous ethylene may reduce the risk of unwanted
ripening, although mixed transport should be avoided.

15.10 Marketing
Modern supermarket chains require large quantities of uniform produce that can be
purchased on contract for delivery at a particular time to stores across a city or
country. This allows the supermarket chain to promote the product at a special
price. Mangoes are generally priced per fruit rather than by weight, although this is
changing. Barcoding and/or Price Lookup Codes (PLU) on the labels of individual
fruit for electronic checkout processing improves monitoring of purchase habits
and stock control. The International Federation of Produce Standards (IFPS) (2008)
provides a forum for stan-dardization of produce labelling and the PLUs are
applicable internationally. Proctor and Cropley (1994) cautioned the need to ensure
that label adhesives comply with food additive restrictions in the EU.

Networks and cooperatives

Marketing cooperatives or networks can assist individual producers to obtain


critical mass in an industry, and fulfil buyer expectations of large supply and
seasonal spread of production (Glogoski, 1995; Griffin, 1995; Higginbottom, 1995;
M.C. Nguyen et al., 2004).

Promotion and consumer education

Mangoes are increasingly popular among affluent consumers in the EU, North
America and northern Asia. In the tropics, they are reminders of a non-urban
living, which has become less common because of rapid indus-trialization and
migration to the cities. Whether for domestic use or export, mangoes must compete
in the fresh market with other equally attractive, nutritious, aromatic and tasty fruit.
Mangoes must also increasingly compete with the snack food, beverage and
entertainment industries.
Consumer education can encourage consumption and sales. Customers can be
educated how to select and store mangoes and how to use both the
Postharvest Technology 579

fresh and the processed products in a variety of ways, thereby increasing total
demand. Production of mango slices in take-away packs can tap domes-tic and
export markets for ready-to-eat, healthy products and circumvent some
disinfestation requirements (see Raymundo et al., Chapter 17, this vol-ume).
Siriphanich (1994) has reviewed minimal processing of tropical fruit and noted the
advantages of gaining market access and reducing transporta-tion costs.

15.11 Conclusions
Mango production has been based almost entirely on Mangifera indica, albeit a
variable meld of thousands of cultivars which may be derived from inter-specific
hybrids of a few closely related species (Kostermans and Bompard, 1993). Given
its perishable nature, capitalizing more on the diversity of exist-ing germplasm to
develop cultivars with superior storage traits linked to customer appeal could
deliver major benefits.
Future improvements in postharvest technology and quarantine treat-ment will
come from refinement of preharvest management, for example reducing disease
inoculum and increasing fruit resistance to disease, reduc-ing harvest costs and
fruit damage, improving postharvest treatments and systems, and supply chain
approaches to enhance fruit longevity and quality and reduce the risks of product
damage. Improvements will also accrue from the provision of user-friendly
information for supply chain personnel, but only if the information is utilized and
implemented. Increases in throughput via the automation of harvesting and
treatment systems for fruit will increase as production and marketing costs escalate.
Labour saving and work effi-ciency will also become more critical. Innovative
transport arrangements may become necessary as regional development places
greater pressures on transport systems. International, collaborative joint-marketing
ventures will ensure year-round supplies of uniform quality fruit, and per capita
consump-tion of mangoes will increase (Johnson, 1995).

Acknowledgements
The authors acknowledge the contributions of the co-authors of Johnson et al.
(1997), which this book chapter supercedes, and Leanne Taylor and Roberto
Marques for assistance with references. The authors also thank the Depart-ment of
Primary Industries and Fisheries for research programme support.

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16 World Mango Trade and the
Economics of Mango Production

E.A. Evans and O.J. Mendoza


University of Florida, Florida, USA

16.1 Introduction 606


16.2 Recent Trends in World and USA Mango Production, Trade and Consumption 607
World situation 607
USA mango production, imports and consumption 611
16.3 Sample Costs and Returns Associated with the Establishment and Production of
Mango Orchards 613
General approach to estimating cost of production of orchard crops 614
Main assumptions 614
Discussion of establishment phase budget 617
Discussion of production phase budget 621
Profitability analysis 624
16.4 Conclusions 626

16.1 Introduction
Worldwide mango production occurs in over 90 countries. Although only a
relatively small proportion of total mango production enters international trade
(<4%), the volume traded has increased substantially since the late 1990s. Among
the factors responsible for the increased mango production, trade and consumption
are lower prices, year-round availability, fewer horticultural trade barriers, changes
in food consumption preferences, longer shelf life for perishables and consumer
interest in healthier foods. Although not a major mango producer, the USA has
developed most of the popular cultivars traded on the international market, and is
the largest single-country mango importer. The costs and returns and general
practices of establishing and maintaining orchards vary considerably from coun-try
to country and within each country (different regions and production systems).

 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
606 (ed. R.E. Litz)
World Mango Trade and Economics 607

16.2 Recent Trends in World and USA Mango Production,


Trade and Consumption
World situation

Asia accounts for approximately 77% of global mango production, and the
Americas and Africa account for approximately 13% and 9%, respectively
(FAOSTAT, 2007). In 2005, world production of mango was estimated to have
reached 28.51 million t, an increase from the 27.82 million t recorded in the
previous year. Between 1996 and 2005, production grew at an average annual rate
of 2.6%. Table 16.1 shows the world’s top ten mango-producing coun-tries, which
account for about 85% of the world’s production.
India is the largest producer, accounting for 38.58% of global production from
2003 to 2005. During that period, the Indian mango crop averaged 10.79 million t,
followed by China and Thailand at 3.61 million t (12.90%) and 1.73 million t
(6.20%), respectively. Other leading mango-producing coun-tries and their
respective shares of world production during the 2003–2005 period include Mexico
(5.50%), Indonesia (5.29%), Pakistan (4.48%), Brazil (4.30%), the Philippines
(3.53%), Nigeria (2.61%) and Egypt (1.28%).
Although currently only 3.3% of the world production of mango is traded
globally, this represents a noticeable increase over the quantities traded since the
late 1980s. In terms of distribution, Mexico, Brazil, Peru, Ecuador and Haiti supply
the majority of North America’s imports. India and Pakistan are the predominant
suppliers to the West Asian market. South-east Asian coun-tries get most of their
supplies from the Philippines and Thailand. European Union (EU) buyers source
mango mainly from South America and Asia.
In 2005, global exports of mango reached 912,853 t, a slight decrease of
0.73% compared with the previous year, and were valued at US$543,100,000
(FAOSTAT, 2007). Table 16.2 shows the top ten major mango-exporting coun-
tries. India is the largest producer but only recently has overtaken Mexico as the
number one exporter of the fruit. For the 2003–2005 period, Mexico and India
dominated the export trade with shares of 22.64% and 20.25%, respec-tively,
followed by Brazil (13.18%) and Pakistan (6.94%). Other major export-ers include
the Netherlands (major re-exporter), Peru, Ecuador, the Philippines, Thailand and
China.
World imports of mango increased from 397,623 t in 1996 to 826,584 t in
2005. The USA is the number one importer of mango. During the 2003–2005
period, the USA imported 271,848 t, or approximately a third of the total mango
imports (Table 16.3).
The Netherlands imported 88,300 t of mangoes (10.62%), although most of it
is redistributed throughout the EU. Other prominent importing countries that are
also major redistributors are the United Arab Emirates (6.82%) and Saudi Arabia
(5.32%). Most of these imports are redistributed to other countries within the
Middle East. Although China (4.91%) appears as a major importer, the quan-tities
imported have been declining. For example, China imported 57,000 t in 2004 and
only 19,000 t in 2005. This could be due to increases in domestic pro-duction in
response to an increase in domestic demand driven by rising per
608
Table 16.1. World’s ten major mango producers, 1996–2005 (1000 t) (Source: FAOSTAT, 2007).

Countries 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2003–2005 (%)

E.A. Evans and O.J. Mendoza


India 11,000 11,000 10,230 9,780 10,500 10,060 10,640 10,780 10,800 10,800 38.58
China 2,074 2,410 2,562 3,127 3,211 3,273 3,513 3,571 3,582 3,673 12.90
Thailand 1,181 1,198 1,088 1,462 1,633 1,700 1,700 1,700 1,700 1,800 6.20
Mexico 1,189 1,500 1,474 1,508 1,559 1,577 1,523 1,362 1,573 1,679 5.50
Indonesia 783 1,088 600 827 876 923 1,403 1,526 1,438 1,478 5.29
Pakistan 908 914 917 916 938 990 1,037 1,035 1,056 1,674 4.48
Brazil 593 508 469 456 538 782 842 1,254 1,358 1,000 4.30
Philippines 898 1,005 945 866 848 882 956 1,006 968 985 3.53
Nigeria 656 689 731 729 730 730 730 730 730 730 2.61
Egypt 203 231 223 287 299 325 287 319 375 380 1.28
Others 3,248 3,230 3,347 3,656 3,597 3,731 4,001 4,327 4,242 4,308 15.34
World total 22,733 23,773 22,584 23,615 24,730 24,973 26,634 27,609 27,822 28,508 100.00
Table 16.2. World’s ten major mango-exporting countries, 1996–2005 (1000 t) (Source: FAOSTAT, 2007).

World Mango Trade and Economics


Countries 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2003–2005 (%)

Mexico 148 187 209 204 207 195 195 216 213 195 22.64
India 27 45 47 38 39 46 42 179 156 223 20.25
Brazil 24 23 39 54 67 94 104 138 111 114 13.18
Pakistan 18 25 39 41 48 52 48 60 82 49 6.94
Netherlands 21 25 17 37 34 43 33 58 51 69 6.42
Peru 11 6 11 20 21 27 35 40 60 58 5.71
Ecuador 0 2 7 0 26 34 30 38 41 40 4.31
Philippines 40 45 53 35 40 39 36 38 36 25 3.61
Thailand 8 9 10 10 9 11 9 8 33 2 1.55
China 12 7 9 10 5 5 15 22 10 4 1.31
Others 80 104 87 103 132 121 127 126 127 135 14.08
World total 391 478 529 552 628 666 673 923 920 913 100.00

609
610
Table 16.3. World’s top ten major mango-importing countries, 1996–2005 (1000 t) (Source: FAOSTAT, 2007).

Countries 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2003–2005 (%)

E.A. Evans and O.J. Mendoza


USA 171 187 197 219 235 238 263 278 276 261 32.70
Netherlands 25 34 35 63 62 70 71 91 76 98 10.62
United Arab Emirates 28 37 48 48 42 46 52 62 58 51 6.82
Saudi Arabia 10 16 14 9 28 36 35 40 42 51 5.32
China 36 40 47 33 33 34 38 47 57 19 4.91
Bangladesh 5 9 0 11 21 21 14 43 37 36 4.63
UK 16 18 18 23 22 27 24 32 37 47 4.63
Germany 13 17 17 24 23 25 28 32 33 37 4.11
France 18 23 22 31 26 26 27 32 35 35 4.09
Malaysia 14 6 21 1 20 27 31 26 45 19 3.59
Others 61 68 66 84 114 106 101 142 148 173 18.58
World total 398 454 486 545 628 656 684 825 843 827 100.00
World Mango Trade and Economics 611

capita income. Other noticeable importers include Bangladesh and the UK (4.63%
each), Germany (4.11%), France (4.09%) and Malaysia (3.59%).
The most popular export mango cultivars continue to be ‘Kent’, ‘Tommy
Atkins’, ‘Haden’ and ‘Keitt’. These cultivars have fruit with a red blush and are
less fibrous, firmer and more suited for long-distance transportation. The green
cultivars are only now being widely accepted in the international market and
include cultivars such as ‘Ataulfo’ and ‘Amelie’. Other cultivars which are gain-
ing significance in international trade include: ‘Alphonso’, ‘Dudhpeda’, ‘Kesar’,
‘Sindhu’, ‘Pairi’, ‘Desi’, ‘Chausa’, ‘Langra’ and ‘Katchamita’. Most of these
newer cultivars on the international scene are coming from India and Pakistan.
Since the late 1990s, per capita consumption has increased noticeably in the
USA, Japan and China, mainly due to higher income levels, improved advertising
and lower mango prices. On the international scene, prices for most mango
varieties have declined considerably over the decade, dropping about 50%. For
example, the average price for mango in the European mar-ket was US$12/kg in
1996, compared to just over US$6/kg today. The reduc-tion in price is partly due to
the increased availability of tropical fruits. There is consensus, however, that prices
have stabilized but could increase with proper promotional efforts.

Although the quantity of processed mango fruit that is traded internation-ally


is small compared with the fresh fruit trade (<7%) there is evidence to suggest that
it is increasing. Such products include mango juice, pickled man-goes, mango
chutney, mango pulp and paste, mango purée, dried fruit, mango slices in brine and
mango flour. India is the main exporter of processed mango followed by Pakistan,
Brazil and Zimbabwe. Major importers include the United Arab Emirates, Saudi
Arabia, Kuwait, USA, UK and Canada.

USA mango production, imports and consumption

The USA is not a major mango producer even though most of the commer-cially
traded varieties have been developed in Florida. USA production, mainly in
Florida, remains fairly stable at a little under 3000 t/year.
The USA is currently the world’s leading importer of fresh mangoes,
accounting for 32.70% of the total imports during the 2003–2005 period
(FAOSTAT, 2007). Figure 16.1 shows the trend of mango imports into the USA
between 1997 and 2006. Overall, the graph indicates a steady increase in the
volume of mango imports. Between 1997 and 2006, imports increased from
187,193 t to 298,088 t, an average annual growth rate of 5.46%. Mango imports
were valued at about US$233,100,000 in 2006 (USDA, Foreign Agri-cultural
Service, 2007).
The main sources of USA imports of mango are Mexico, Peru, Ecuador and
Brazil. Mexico is the main supplier of mango to the USA (60.78% share in 2006)
(Fig. 16.2). In recent years, Brazil, Peru and Ecuador have become significant
exporters to the USA, competing with Mexico at the beginning and the end of the
season. The USA exports very few of its mango imports, mainly to Canada and the
UK.
612 E.A. Evans and O.J. Mendoza

350,000

300,000

250,000
USA total imports (t)

200,000

150,000

100,000

50,000

0
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year

Fig. 16.1. USA total imports of mango (t), 1997–2006 (Source: USDA
Foreign Agricultural Service, 2007).

200,000
180,000
160,000
140,000
USA total imports (t)

120,000
100,000
80,000
60,000
40,000
20,000
0
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year

Mexico Peru Ecuador Brazil

Fig. 16.2. USA total imports of mango (t) by country, 1997–2006 (Source:
USDA Foreign Agricultural Service, 2007).

USA consumption of mango has increased steadily, from a per capita level of
0.5 kg in 1996 to 1 kg in 2005 (USDA, Economic Research Service, 2007). The
growth in USA consumption of mango is driven by many factors, such as year-
round availability, lower prices, consumer preferences and more
World Mango Trade and Economics 613

Table 16.4. Average cost, insurance and freight (CIF) prices for selected varieties from
main suppliers to the USA, 1998–2006 (US$/kg) (Source: USDA Agricultural Marketing
Service, 2007).

Country of origin

Year Brazil Ecuador Haiti Peru Mexico Year average


a
1998 3.43 3.21 n/a 3.61 2.09 3.09
1999 2.13 1.67 2.24 1.89 1.78 1.94
2000 2.09 1.69 2.05 1.65 1.72 1.84
2001 1.74 1.65 2.24 1.85 1.69 1.83
2002 1.67 1.47 2.13 1.61 1.61 1.70
2003 1.72 1.28 1.96 1.45 1.45 1.57
2004 1.65 1.83 1.98 1.43 1.43 1.66
2005 1.67 1.94 2.11 1.58 1.67 1.79
2006 1.65 1.67 2.11 1.39 1.72 1.71
a
n/a, not available.

disposable income. However, mango consumption levels are still considered


relatively low when compared to other fruits, for example bananas (11 kg) and
oranges (5 kg).
USA prices for mango vary widely by cultivar and season, mainly due to the
fact that the commodity demand is price inelastic (sensitive to variations in
quantities available, a 1% increase in the quantity tends to lead to a >1% fall in the
price). In general, mango prices have been steadily declining over the past decade.
Table 16.4 shows the average cost, insurance and freight (CIF) prices for mango
imports into the USA from main supply sources dur-ing the 1998–2006 period.
Although prices have decreased noticeably from the 1998 level, they do appear to
have stabilized in the last couple of years.

16.3 Sample Costs and Returns Associated with the


Establishment and Production of Mango Orchards
In this section, we have estimated the per hectare costs to establish an 18 ha
commercial mango orchard in Florida USA, as well as the annual estimated costs
and net returns per hectare after establishment. We have not provided too much
detail on production practices (since such information is readily available
elsewhere), but rather focus on the methodology of estimating the costs and returns
of establishment and production. Although somewhat hypo-thetical, the data
presented are based on a combination of information obtained from the growers,
economic engineering using recommended prac-tices, and discussions with
industry experts. We begin by briefly describing the general methodology,
followed by a listing of some of the major assumptions used in the analysis.
Estimates of the establishment costs are then discussed, and the chapter closes with
a discussion of profitability of a mango operation.
614 E.A. Evans and O.J. Mendoza

General approach to estimating cost of production of orchard crops

The general approach for estimating the cost of production of perennial crops
(orchards and vineyards), which usually take more than a year to begin pro-
duction, is to develop two separate budgets: one for the establishment phase and
the other for the production phase. The establishment phase bud-get reflects the
sum total of all expenses (expressed on a yearly and per unit basis) that are
incurred over the years to bring an orchard into meaningful (mature trees)
production. Once this amount is determined, it is treated as if it were an expense
incurred in the purchase of a ‘capital item’, for example machinery to be used in
the production phase of the orchard. As in the case of a capital item, an equal
annual amount (amortized amount) is charged to the production operation as an
expense spread over the estimated future life of the orchard. In other words, the
amortized amount is included as a non-cash overhead expenditure in the production
phase budget when determin-ing net returns from the enterprise. The production
phase budget estimates the costs and returns on an annual per unit basis associated
with the mainte-nance of the orchard after it has been established. Although the
procedure seems daunting, it is made much easier by using spreadsheet software
with built-in formulas for calculating amortization.

Main assumptions

The following assumptions were used to estimate production costs and returns for
18 ha operations.

Land
The hypothetical farm consists of 20 ha. A mango orchard is being estab-lished on
18 ha, with roads, irrigation system and farmstead occupying 2 ha. The 18 ha
orchard is considered large enough to use machinery and equip-ment efficiently.
The orchard is farmed by the owner with the help of hired part-time labour.

Site preparation
It is assumed that the land is relatively clear, with no costs being included for major
land preparation such as timber clearing, rock removal or land level-ling. However,
if these operations are required, they should be included. The main operations
considered here are associated with fencing and road con-struction for travelling
and harvesting. These operations are usually done 1 year prior to planting, but costs
are shown in the first year.

Planting, training and pruning


Trees are planted on 7.5 m × 7.5 m spacing, or 175 trees/ha. The life of the orchard
in this study is projected to be 25 years. The variety considered in this study is
‘Tommy Atkins’. Pruning and training begin in the third year, and labour time
required for pruning increases in the successive years.
World Mango Trade and Economics 615

Table 16.5. Annual fertilizer rates @ 6% N.

Year 1 2 3 4 5 6 7+

Rate (kg/ha) 236 472 707 786 1100 1257 1415

Hedging and topping operations are carried out immediately after fruit har-vesting.

Fertilization
During the first 3 years, an N-P-K fertilizer (6% nitrogen) is spread by hand six
times each year. After year 3, the frequency of application decreases to four times
each year. Table 16.5 shows the annual fertilizer rates assumed. In addition, the
trees are given annual nutritional sprays of copper, zinc, man-ganese and boron.
Iron is applied in chelated form as a soil drench two to three times each year.

Irrigation
Total irrigation costs include the cost of pumping water and irrigation labour.
Water for our irrigation system is supplied from a well; therefore, the cost of the
water is zero. Irrigation costs for individual orchards vary, depending on the
amount of water pumped, pumping system, energy source and irrigation district.

Weed control management


Weeds in the tree rows are controlled with applied pre- and post-emergent
(residual) contact herbicides such as Roundup®. With five applications each year,
spot sprays are applied at the rate of approximately 5 l/ha. However, as the trees
grow larger and shade reduces weed growth, the number of appli-cations is reduced
to approximately three per annum.

Harvest
Mango fruits are best harvested using clippers and hand-carried harvesting bags
(about US$0.11/kg). With large trees, fruits are harvested using picking poles, with
or without attached clippers, which are equipped with bags into which the fruit
falls. After harvesting, the fruits are usually piled under a tree on the ground. The
fruit is then loaded onto trucks in the field and hauled to packing houses for
US$0.11/kg.

Yields and returns


A major assumption is that the orchard requires approximately 7 years to reach
maturity; hence the first 6 years are considered the establishment phase. Even
though trees require about 7 years before reaching maturity, they will start having
saleable fruits from about the third year. Table 16.6 shows the typical yield
assumptions used in the analysis. Annual gross field yields at maturity are assumed
to be 22,000 kg/ha.
616 E.A. Evans and O.J. Mendoza

Table 16.6. Yield assumptions.

Year 1 2 3 4 5 6 7+

Yield (kg/ha) 0 0 1,700 4,500 9,000 15,500 22,000

Labour
Hourly wages for workers are US$10.00 for skilled workers and US$7.00 for field
workers. Adding 34% for the employer’s share of federal and state pay-roll taxes,
insurance and other possible benefits, yields a labour rate of US$13/h for skilled
labour and US$9/h for field labour.

Cash overhead
Cash overhead consists of numerous cash expenses paid during each year that are
assigned to the entire farm, not to a particular operation. These costs include
property taxes, office expenses, capital interest, liability and property insurance,
equipment repairs, sanitation services and crop insurance.

PROPERTY TAXES. In the USA, most counties charge a base property tax rate of
approximately 1% on the assessed value of the property.

INTEREST ON OPERATING CAPITAL. Interest on operating capital is based on cash operating


costs and is calculated at a nominal rate of 5%/year. A nominal interest rate is the going
market cost of borrowed funds.

INSURANCE. Insurance for farm investment differs depending on assets included and
amount of coverage. Property insurance provides coverage for property loss and the
standard practice in the USA according to the American Society of Agricultural
Engineers (ASAE) is to charge about 0.67% of the average value of the assets over
their useful life. Liability insurance covers accidents on the farm (US$580/year).

OFFICE EXPENSES. Office and business operating expenses are estimated at US$355/ha,
and include office supplies, telephone, bookkeeping, account-ing, legal fees, road
maintenance and miscellaneous administrative charges.

SANITATION SERVICES. Sanitation services offer portable toilets for orchards and cost the
farm US$1900/year (US$106/ha). This cost includes delivery and servicing of a single
toilet and washing unit.

CROP INSURANCE. Multi-peril crop insurance is purchased at a cost of US$445/ha.

Non-cash overhead
Non-cash overhead is calculated as the capital recovery cost for equipment and
other farm investments. Capital recovery cost is the annual depreciation
World Mango Trade and Economics 617

and interest costs for a capital investment. It is the amount of money required each
year to be set aside so that a capital item can be fully replaced at the end of its
useful life. It can be viewed as the value of the portion of the capital item that was
utilized in the production process during that year. The for-mula for the calculation
of the annual capital recovery costs is the PMT Excel formula:

Capital Recovery = –PMT (I, N, (PP − SV)) + (I)*(SV)


Where:
● PMT = the formula that calculates the payment for a loan based on con-stant
payments and a constant interest rate.
● I = the interest rate used to represent the cost of borrowed capital (in this case,
5%).
● N = the number of years.
● PP = the purchase price for the capital item.
● SV = an estimate of the remaining value of an investment at the end of its
useful life. In this case, the percentage of remaining value is calculated from
equations developed by the ASAE based on equipment type and years of life.

Farm equipment on a mango orchard in the region is purchased either new or


used. The study shows the current purchase price for new equip-ment. The new
purchase price is adjusted to 60% to indicate a mix of new and used equipment.

The orchard is irrigated using a sprinkler irrigation system. The life of the
irrigation system is estimated at 20 years. The irrigation system is installed before
the orchard is planted and includes costs of a pumping system, instal-lation labour
and design and materials.
Although it is assumed that the grower owns the land, the going rental rate in
South Florida of approximately US$2500/ha is used to reflect the opportunity
(economic) cost of land.

Discussion of establishment phase budget

Table 16.7 shows the layout of the establishment phase budget. The budget reflects
the annual per hectare costs incurred in establishing a mango orchard. Here it is
assumed that a mango orchard reaches maturity in the seventh year. As such, the
budget reflects all associated cash and non-cash expenses incurred over 6 years.

The information required for each year is similar except for the first 2 or 3
years. With reference to Table 16.7, year 1 includes pre-planting costs (site
preparation and orchard layout), which in this case amounts to US$5000/ha. Most
of the pre-planting costs most likely would occur in the year before, but the
practice is to include these costs in year 1. Planting costs include new trees, digging
holes for trees, setting trees, and wrapping and staking trees. Both pre-planting and
planting costs appear only in year 1. Total per hectare
618
Table 16.7. Sample costs to establish a mango orchard in south Florida (source: compiled by the authors).

Year (US$/ha)

Operation Unit 1st 2nd 3rd 4th 5th 6th

Pre-planting costs
Site preparation 4,500
Orchard layout 500
Total pre-planting costs 5,000
Planting costs
Mango trees (175 trees/ha) US$25.00 4,375

E.A. Evans and O.J. Mendoza


Digging, planting, wrapping and staking US$10.00 1,750
Total planting costs 6,125
Replanting costs
Replaced trees – 5% 5% 225
Digging, planting, wrapping and staking 90
replaced trees
Total replanting costs 315
Cultural costs
Orchard pruning 0 0 0 260 260 260
Herbicide 100 100 100 100 100 100
Fungicide 0 0 0 1,360 1,360 1,360
Fertilizer 80 160 242 270 430 600
Insecticide 200 200 200 200 200 200
Mowing 390 390 240 180 180 180
Irrigate and walk lines 90 90 90 90 67 67
Pest control advisor 0 0 150 150 150 150
Pollination 0 0 0 0 450 450
Rodent control (squirrels) 60 60 60 60 60 60
Total cultural costs 920 1,000 1,082 2,670 3,257 3,427
Harvesting and marketing costs
Picking – US$0.11/kg US$0.11 0 0 187 495 990 1,705
Packing and shipping – US$0.11/kg US$0.11 0 0 187 495 990 1,705
a
Sales charge @ 10% of FOB price 10% 0 0 153 405 810 1,395
Total harvesting, marketing and inspection 0 0 527 1,395 2,790 4,805
costs
Interest on operating capital @ 5% 5% 602 66 54 134 163 171
Total operating costs 12,647 1,381 1,663 4,199 6,210 8,403
Cash overhead costs
Liability insurance 30 30 30 30 30 30
Crop insurance 0 0 445 445 445 445
Sanitation fee 0 0 106 106 106 106

World Mango Trade and Economics


Office expenses 355 355 355 355 355 355
Property taxes 150 150 150 150 150 150
Property insurance 250 250 250 250 250 250
Investment repairs 185 190 210 210 210 210
Interest on operating capital (cash overhead) 5% 49 49 77 77 77 77
Total cash overhead costs 1,019 1,024 1,623 1,623 1,623 1,623
Total cash costs 13,666 2,405 3,286 5,822 7,833 10,027
Income from production 0 0 1,530 4,050 8,100 13,950
Net cash costs for the year 13,666 2,405 1,756 1,772 –267 –3,923
Accumulated net cash costs 13,666 16,070 17,827 19,598 19,332 15,408
Non-cash overhead
Land rent 2,500 2,500 2,500 2,500 2,500 2,500
Machinery and equipment 405 405 405 405 405 405
Building 185 185 185 185 185 185
Tools (shovels, picking bags, saws, etc.) 20 20 20 20 20 20
Shop tools 42 42 42 42 42 42
Irrigation system 380 380 380 380 380 380
Drippers 30 100 0 0 0 0
Sprinklers 0 0 30 30 30 30
Total non-cash overhead costs 3,562 3,632 3,562 3,562 3,562 3,562

619
Total net cost for the year 17,228 6,037 5,318 5,334 3,295 –361
a FOB, freight on board.
620 E.A. Evans and O.J. Mendoza

pre-planting and planting costs are estimated at US$11,125 (US$5000 + US$6125).


In year 1, it is also assumed that there are no replanting costs asso-ciated with
replacement trees. However, provisions are made to accommo-date this cost in
subsequent years. Here we assume that 5% of the trees planted in year 1 will be
replaced in year 2. Cultural costs are then estimated, includ-ing standard operations
such as pruning, fertilizing and mowing. In our example, the total cultural costs in
year 1 are estimated at US$920/ha. Next are harvesting and marketing costs. This
subheading includes the costs of such operations as picking, packing and shipping.
Other charges include sales charges (if these operations were commissioned) and
inspection and assessment fees. In both years 1 and 2, it is not expected that trees
will have ‘saleable’ crops and, in fact, it is recommended that efforts should be
made to deflower such trees to prevent fruiting and to foster better tree growth.
Con-sequently, there are no marketing costs in those years. To the sum for pre-
planting, planting, replanting, cultural, and harvesting and marketing costs is added
an interest charge of 5%. This reflects the economic (opportunity cost) cost of
money that was either borrowed or owner-financed, to cover these expenses. Thus
the total operating (variable) costs in year 1 is US$12,647/ha.

Overhead costs are fixed costs (in the sense that they must be paid irre-
spective of the level of production) that relate to the entire farm. For the pur-pose
of budgeting they are allocated on a per hectare basis. There are two subcategories
of overhead costs: cash overhead and non-cash overhead. Cash overhead costs
must be paid with cash during the year. They include liability and crop insurance,
taxes and office expenses. Interest is charged on cash overhead costs. Total cash
overhead costs plus total operating costs equals total cash costs. Total cash costs
represent the total cash expended in any given year for producing and marketing a
crop. In this example total cash costs for year 1 is US$13,666 (US$12,647 +
US$1019)/ha. Non-cash over-head costs reflect capital recovery associated with
asset ownership is added later and discussed below.

The next major category is income from production. Mango trees will start
supporting economic crops from about the third year, even though the trees are not
yet fully mature. The income from the sales of these crops is deducted (credited)
from the total cash costs for that year. In years 1 and 2, there is no income.
However, in year 3 there is income. Based on our analysis, the income (US$1530)
is then deducted from total cash costs for that year (US$3286) to give the net cash
costs for the year (US$1756). The negative net cash costs recorded for year 5
(US$267) implies that, for that year, revenues from the sales of fruit exceeded total
cash expenditures for that year (US$7833 − US$8100).

The accumulated net cash costs category is self-explanatory; it keeps a running


tally of cash costs incurred during the establishment years. Thus, based on our
analysis, the accumulated net cash costs in year 6 amounts to US$15,408/ha,
reflecting the sum of the net cash costs for years 1 through to 6, i.e. US$13,666 +
US$2405 + US$1756 + US$1772 + (−US$267) + (−US$3923). This amount does
not include capital recovery costs for equipment and machinery that a grower owns
and that is used in establishing an orchard.
World Mango Trade and Economics 621

To complete the establishment phase budget, it is necessary to include the non-


cash overhead charges. As previously defined, these are not cash payments. They
include an estimate of the amount of money that should be set aside to help with
replacing an investment item at the end of its useful life. They usually reflect
depreciation and interest charges associated with a capital item (i.e. the
contribution of a capital item to the production process for a given year). If some of
these services were commissioned they would be included in cash charges. The
same is true for land rent. Here it is included among the non-cash overhead charges
because we assumed that the grower already owns the land and therefore is not
paying cash for the land. Since the land could have been sold and the proceeds
placed in a bank to earn interest, the rent charged against production represents, in
a sense, lost interest from not selling the land and placing the proceeds in a money-
bearing bank account. In other words, it is included as payment to the owner for
using his land in this manner. The sum of non-cash overhead and net cash cost
equals the total net cost for that year. Table 16.7 shows that, in year 1, the total net
cost is US$17,228/ha, out of which US$13,666 and US$3562 are the net cash costs
and the non-cash overhead costs, respectively.

The final category is total accumulated net costs, which keeps a tally of the
total cost incurred since the start of the establishment phase. Based on the
information provided in Table 16.7, the data reveal that, at the end of the 6 years,
the investor would have invested US$36,850/ha to establish the orchard. Of this
amount, the cash costs amount to US$15,408 and non-cash costs due to ownership
of fixed assets were estimated at US$21,442. Thus, the total cost to establish the 18
ha mango orchard is US$663,300 (18 × US$36,850) of which the cash amount
(ignoring fixed costs) would be US$277,344 (18 × US$15,408).

The full establishment costs of US$663,300 (US$36,850/ha) is then amortized


at 5% real rate of interest for the estimated life of the orchard (25 years) to give an
annual cost for capital recovery (interest and depreciation) of US$2615/ha. This
amount will be charged as part of the non-cash over-head charges (Table 16.8) in
computing the cost to produce a hectare of mango during the production phase
(after the orchard has been established).

Discussion of production phase budget

Table 16.8 provides the estimates of the annual growing costs (after establish-
ment) for mango in south Florida. The budget is similar to the one used in the
establishment phase except that there are no provisions for pre-planting, planting
and replanting expenses, and the estimates are for a typical year. Also of
importance, and shown in italics in Table 16.8, is the amortized estab-lishment cost
(US$2615/ha) obtained from Table 16.7. This reflects the annual amount that must
be recovered for the investment made in establishing an orchard.

Based on Table 16.8, it can be seen that the total annual cash cost to pro-duce
a hectare of mango, assuming a yield of 22,000 kg/ha (Table 16.6) is
622
Table 16.8. Sample cost per hectare to produce mango in south Florida.

Costs (US$/ha)

Operation Labour Fuel, lube Material Total


Operation Unit time (h/ha) cost and repairs cost Custom/rent cost

Cultural costs (materials, labour, fuel,


lube and repairs)
Orchard pruning (× 1) 15 135 25 0 150 310
Herbicide (× 4) 8 72 37 40 0 149
Fungicide (× 12) 30 390 36 940 0 1,366

E.A. Evans and O.J. Mendoza


Fertilizer (× 4) 12 156 24 792 0 972
Insecticide (× 1) 4 36 25 40 0 101
Mowing (× 6) 12 156 25 0 0 181
Irrigate and walk lines (× 1) 3 27 0 40 0 67
Pest control advisor (× 1) 0 0 0 0 150 150
Pollination (× 1) 0 0 0 0 450 450
Rodent control (squirrels) (× 1) 0 0 0 0 60 60
Machinery repair 5 65 54 20 0 139
Total cultural costs 89 1,037 226 1,872 810 3,945
Harvesting and marketing costs
Picking – US$0.11/kg US$0.11 0 0 0 0 2,420 2,420
Packing and shipping – US$0.11/kg US$0.11 0 0 0 0 2,420 2,420
Sales charge @ 10% of FOB price 10% 0 0 0 0 1,100 1,980
Total harvesting and marketing costs 0 0 0 0 5,940 6,820
Interest on operating capital @ 5% 5% 197
Total operating costs/hectare 1,037 226 1,872 6,750 10,962
Cash overhead costs
Liability insurance 30
Crop insurance 445
Leaf analysis 12
Soil analysis 12
Sanitation fee 106
Office expenses 355
Property taxes 150
Property insurance 250
Investment repairs 207
Interest on operating capital (cash overhead) 5% 78
Total cash overhead costs 1,645
Total cash costs/hectare 12,607

World Mango Trade and Economics


Cost per Annual
producing cost: capital
hectare recovery/ha Total cost

Non-cash overhead
Land rent 2,500 2,500 2,500
Machinery and equipment 3,648 405 405
Building 2,964 185 185
Tools (shovels, picking bags, saws, etc.) 217 20 20
Shop tools 464 42 42
Irrigation system 4,817 380 380
Sprinklers 20 20 20
Amortized establishment cost 2,615 2,615
Total non-cash overhead costs 14,630 6,167 6,167
Total cost/hectare 18,774

623
624 E.A. Evans and O.J. Mendoza

US$12,607 or about US$0.57/kg. Of this total, cultural costs accounts for US$3945
(31%), harvesting and marketing costs amount to US$6820 (54%) and cash
overhead costs US$1645 (13%). When non-cash overhead costs are added in, the
economic cost to produce a hectare of mango in South Florida is estimated at
US$18,774, or about US$0.85/kg.

Profitability analysis

Table 16.9 utilizes the information presented in Tables 16.6 and 16.8 along with
information obtained elsewhere to analyse the profitability of mango production in
South Florida based on 2006 data. Based on an average freight on board (FOB)
price of US$0.90/kg and marketable yield of 22,000 kg/ ha (actual yield would be
slightly more), the gross returns are calculated as US$19,800/ha. Subtracting total
operating costs of US$10,962 (cultural, and harvesting and marketing costs) from
this amount gives a gross margin of US$8838/ha. In other words, based on these
assumptions, the enterprise is profitable in the short run since gross margin is
positive. This implies that the returns from crop sales are more than sufficient to
cover the variable costs incurred in the production and marketing of the crop and
there are still funds left over to go towards paying overhead costs.

To be viable in the long run, the amount remaining should be able to cover
overhead (cash and non-cash) and provide a reasonable return to the operator.
Subtracting cash overhead costs (US$1645) and non-cash overhead costs
(US$6167) from gross margin (US$8838) gives net returns of US$1026/ ha.
Viewed slightly differently, from total revenue of US$19,800 obtained from selling
a hectare of the crop, after paying all costs (variable and fixed) the amount of
US$1026/ha remain as payment to management. This results in a return of
approximately 7% on cash and non-cash investment.1
Given that prices and yield can vary, Table 16.10 shows a sensitivity anal-ysis
for changes in price and yield variables. The sensitivity analysis is cal-culated on
net returns but could be done on gross margin. As seen in Table 16.10, a 10% price
increase combined with a 10% yield increase would result in over a 335% net
return per hectare increase (from US$1026 to US$4460/ ha). A 10% price increase
would have a greater impact on net return than would a 10% yield increase. In the
case of the former, the net returns would increase from US$1026/ha to US$2984/ha
(191%). In the case of the latter, net returns would increase to US$2324 (127%).
This underscores the importance of doing whatever is necessary to improve crop
prices such as improving the quality of the product and engaging in direct
marketing where possible.
Assuming marketable yields of 22,000 kg/ha, the break-even price, or the price
at which net returns (economic profits) would be zero, is computed as US$0.85/kg.
In other words, at this price the grower is just able to cover both variable and fixed
costs. Any FOB price above this would result in pos-itive net return and vice versa.
Likewise, if prices were to remain at US$0.90/ kg, growers would have to ensure
that they obtain yields in excess of about 20,000kg/ha (break-even volume) to
generate positive returns.
World Mango Trade and Economics 625

Table 16.9. Costs and returns to produce mango in south Florida, 2007.

Price or Value or
cost per cost/ha
Unit Quantity/ha unit (US$) (US$)

Gross returns kg 22,000 0.90 19,800


Total gross returns for mangoes 19,800
Operating costs
Pruning Tree 175 1.77 310
Herbicide Application 4 37.25 149
Fungicide Application 12 113.83 1,366
Insecticide Application 1 101.00 101
Fertilizer Application 4 243.00 972
Mow middles Times 6 30.17 181
Pest control advisor ha 1 150.00 150
Irrigation ha 1 67.00 67
Pollination ha 1 450.00 450
Rodent control ha 1 60.00 60
Machinery repair ha 1 139.00 139
Interest on operating capital @ 5% 5% 197
Harvesting and marketing costs
Picking – US$0.11/kg kg 22,000 0.11 2,420
Packing and shipping – US$0.11/kg kg 22,000 0.11 2,420
Sales charge @ 10% of FOB price kg 22,000 0.09 1,980
Total operating costs/hectare 10,962
Gross Margin 8,838
Cash overhead costs
Liability insurance 30
Crop insurance 445
Leaf analysis 12
Soil analysis 12
Sanitation fee 106
Office expenses 355
Property taxes 150
Property insurance 250
Investment repairs 207
Interest on operating capital 5% 78
(cash overhead) @ 5%
Total cash overhead costs 1,645
Total cash costs/hectare 12,607
Non-cash overhead
Land rent 2,500
Machinery and equipment 405
Building 185
Tools (shovels, picking bags, 20
saws, etc.)
Shop tools 42

(Continued)
626 E.A. Evans and O.J. Mendoza

Table 16.9. (Continued)

Price or Value or
cost per cost/ha
Unit Quantity/haunit (US$) (US$)

Irrigation system 380


Sprinklers 20
Amortized establishment cost 2,615
Total non-cash overhead costs 6,167
Total cost/hectare 18,774
Net returns above total costs 1,026

Table 16.10. Sensitivity analysis.

FOB price (US$/kg)

Yield (kg/ha) 0.72 0.81 0.90 0.99 1.08

19,800 –3833 –2052 –272 1508 3288


22,000 –2891 –933 1026 2984 4943
24,200 –1949 187 2324 4460 6597

16.4 Conclusions
The major trends and developments occurring within the world and USA mango
market have been discussed and the methodology used to compute the costs of
establishing and maintaining an orchard has been demon-strated. Several trends
were highlighted, including the increasing levels of production, trade and
consumption of mangoes and the declining or stag-nating prices. Although the case
for establishing an orchard was hypotheti-cal, the statistics are based on
information obtained from growers, economic engineering using recommended
practices, and discussions with industry experts.

Acknowledgements
The authors are indebted to many farmers and agricultural researchers who
provided us with innumerable advice and relevant comments about this study. We
wish to thank those who kindly offered suggestions for improve-ment, including Dr
Jonathan Crane, Dr Carlos Balerdi, Scott Smith, Luis D. Roman, Sikavas
Nalampang, Denys Benjamin and Valderez Gonzalez. A spe-cial thanks to Carol
Fountain for editing the manuscript.
World Mango Trade and Economics 627

Note
1This is obtained by adding the interests charged on operating and cash overhead costs
to the net returns to give an adjusted net income. Next, subtract these interests from the
total costs per hectare to give an adjusted total cost per hectare. Finally, express the
adjusted net income as a percentage of the adjusted total cost per hectare.

References
FAOSTAT (2007) Food and Agriculture Organization of the United Nations database.
Available at: http://faostat.fao.org/ (accessed 30 November 2007).
United States Department of Agriculture (USDA) Agricultural Marketing Service (2007)
USDA, Agricultural Marketing Service: Fruit and Vegetable Market News Website.
Available at: http://www.marketnews.usda.gov/portal/fv (accessed 31 March 2008).
United States Department of Agriculture (USDA) Economic Research Service (2007)
Fruit and Tree Nut Yearbook, 2005. Available at: http://www.ers.usda.gov/Data/
FoodConsumption/FoodAvailSpreadsheets.htm (accessed 31 March 2008).
United States Department of Agriculture (USDA) Foreign Agricultural Service (2007)
USDA Foreign Agricultural Service: Market and Trade Data. Available at: http://
www.fas.usda.gov/markettradedata.asp (accessed 3 November 2007).
17 Fruit Processing
L.C. Raymundo, M.T. Ombico and T.M. de Villa
University of the Philippines Los Baños, Laguna, the Philippines

17.1 Introduction 628


17.2 Dehydrated Mango Products 630
Dried mango 630
Mango fruit bar 631
Mango fruit roll 632
Vacuum-puffed dried mango 632
17.3 Spray-dried Mango Powders 634
Spray-dried mango fruit powder and instant mango juice 634
Spray-dried green mango powder 635
Spray-dried instant green mango shake 636
17.4 Capital Investment Costs 637
17.5 Raw Material Requirements of the Mango Processing Plant 638
17.6 Conclusion 639

17.1 Introduction
Processing is a value-adding step that fills the gap between farm production and
marketing. The objective of processing is to convert highly perishable but prime
quality farm commodities to more stable forms. When the process is accomplished
with the least alteration in the nutritional value and aes-thetic properties of the
food, high consumer acceptance is assured. Com-pletely new product lines can
likewise be created out of fresh raw materials through processing. In other
instances, the raw material may undergo such extensive physical alteration during
processing that the product is used dif-ferently by consumers. Culinary experts
devise new uses for these products to fit the changing lifestyles of present-day
consumers. The availability in the market, for example, of pre-mixed condiments
and various meat powders used for flavouring foods such as instant noodles and
similar convenience
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
628 (ed. R.E. Litz)
Fruit Processing 629

foods, has simplified the life of working women who do not have the time or the
skill to prepare food the way their mothers and grandmothers did at home. The
products are affordable and easy to prepare; hence, consumer acceptance is high. In
addition, these foods are found everywhere, from neighbourhood stores in the
suburbs and countryside to supermarkets in urban centres. The product and process
research and development work of food scientists and food technologists, the
business acumen of entrepreneurs and the marketing expertise of product
distributors ensure the availability of the food products.

The global competitiveness of agricultural produce of a country can be con-


siderably enhanced by utilizing appropriate technologies to produce high-quality
processed foods. Research focusing particularly along the areas of processing
equipment, product design and packaging has made the latest techniques available
for the manufacture of new generations of food prod-ucts. Tropical fruits have had
an excellent record of breaking into the world market because of their exotic
flavours (Plate 83). When processed into purée, juice, nectar or simply dried fruit
slices, they can be shipped long distances with only minimum changes in quality.

The popularity of fruits may be attributed to consumer perception of their


health benefits. In the case of fruit juices, many consumers are now looking for
healthy alternatives to the traditional carbonated beverages. Fruits are rich in
dietary fibre as well as phytonutrients, especially antioxidants, and have no
cholesterol. The availability in the market of natural fruit juices derived from fresh
fruits is a welcome alternative to synthetic juices that are being passed off as fruit
juices but are, in reality, sugar-based, fruit-flavoured beverages prepared from
artificial ingredients.
Mango fruits are usually eaten fresh as dessert or as relish depending on their
stage of maturity when picked. In the Philippines and elsewhere, fresh mangoes are
available throughout the year because the flower-induction technology first
described by Barba (1974) allows growers to produce fruit out-of-season. The off-
season fruits, however, are still expensive. Most mango growers have not put the
technology into practice due to the high cost of additional farm inputs necessary for
its successful implementation during the rainy season.

At the peak of the harvest season, on the other hand, oversupply of fresh
mangoes depresses the market price to the detriment of the growers. Moreover,
high temperatures combined with high relative humidity (RH) and intense sunlight
during the harvest season accelerate the metabolic processes associated with
ripening in fresh mangoes, rendering them sus-ceptible to microbial attack,
particularly by Colletotrichum gloeosporioides Penz., the cause of anthracnose.
The physico-chemical changes that occur during ripening also lead to fruit
deterioration and loss of quality. Thus, fresh mangoes are processed to facilitate
better distribution and to stabilize prices.

Mango processing was previously reviewed by Nanjundaswamy (1997), who


described the status of processing technologies and products in India. This review
discusses current trends in mango processing.
630 L.C. Raymundo et al.

17.2 Dehydrated Mango Products

Dehydration works on the principle that by lowering the water (H2O) con-tent of
foods below a certain threshold level, growth of many spoilage micro-organisms is
prevented, thus preserving the food. As more and more H 2O is removed from the
material, its solids content becomes concentrated, further making the food less
suitable for microbial growth.
The choice of the most appropriate drying system is determined largely by the
cost-effectiveness of the process. Sun drying is the most inexpensive method for
drying foods. The products, however, are rather susceptible to contamination by
dirt, insects, rodents, faecal matter and microorganisms. The process also requires
several days to dry each load or batch of raw mate-rials since it relies on the
availability of sunlight. Temperature control is vir-tually impossible. Other systems
have been designed that are more hygienic and equally cost-effective compared to
sun drying. The use of solar dryers practically eliminates most of the inherent
defects of sun drying. However, the reliance of solar dryers on sunlight as the
source of energy for evaporat-ing H2O from the material being dried makes the
system commercially unre-liable. Solar panel collector-equipped dryers with
provision to store energy from sunlight can operate continuously and efficiently.

A mechanical dryer such as the convection oven provides the processor with
control over the system that is essential for a successful drying operation. It elim-
inates most of the problems mentioned above. Although the initial investment cost
is high due to the acquisition of the dryer, in the long run it is more eco-nomical
than sun drying because the drying time per load is much shorter.
Oil-, steam-, liquid propane gas- or electric-powered air heaters are the
alternative sources of energy for the dryers. Dryers have also utilized a wood-fired
furnace that heats the air entering the drying chamber. Its main advan-tage is that
trimmings from raw materials, packing materials and other trash can be burned in
the furnace, keeping the compound clean.

Dried mango

Dehydrated or dried mango slices are among the first products manufac-tured
commercially from ripe mango fruits that caught the attention of con-sumers in the
international market for processed tropical fruits (Plates 83 and 84). The product
was developed in Cebu, the Philippines, from where it spread to the neighbouring
islands of Panay and Negros. It is now produced in many regions of the Philippine
archipelago where mango is abundant. In addition, it is a popular product in
Thailand and elsewhere in South and South-east Asia.

In the Philippines, the ‘Carabao’ mango is the preferred variety for dehy-
dration or drying. The fruit should be at the firm-ripe stage. When over-ripe fruit is
used as raw material, a dark-coloured product will invariably result. Although the
dried pieces from over-ripe mangoes have a more distinct ripe mango flavour that
attracts customers, the shelf life is considerably shorter.
Fruit Processing 631

The fruit is washed thoroughly. The cheeks are sliced with a sharp stainless-
steel knife. Each slice is then cut into two or three pieces length-wise. The flesh is
scooped from the skin with a stainless-steel scoop or ladle. These operations are
done manually; however, a simple and novel peeling and slicing machine for ripe
mangoes has been developed and patented in Australia (as cited by
Nanjundaswamy, 1997).
Sugar syrup is prepared by adding 175 kg sugar, 1.7 kg citric acid and 0.85 kg
sodium metabisulfite in 175 l H2O to make a 45 Brix sugar concen-tration and
heating to 90°C to dissolve the metabisulfite. The prepared mango slices (1 t) are
added to the syrup. The preparation is heated to 90°C and then allowed to stand for
at least 6 h. Subsequently, the sugar concen-tration of the syrup is adjusted by
dissolving more sugar until the concen-tration reaches 50 Brix using a hand
refractometer. After 6 h, the mango pieces are again removed from the syrup and
the sugar concentration is adjusted to 60 Brix using a hand refractometer for the
‘plumping’ stage. The syrup is reheated to 90°C; the mango slices are added to the
syrup and soaked for a further 6 h.

The final step in the process involves the removal of mango pieces from the
syrup. They are lightly rinsed with H2O to remove the excess sugar that may
crystallize on the surface of the mango during drying. The slices are then loaded in
drying trays and dehydrated at 40–50°C in a convection dryer. Drying time usually
requires 18 h. The dried mango pieces are unloaded from the dryer and
reconditioned at room temperature overnight. Each piece is coated with
confectioner’s sugar. The product is then packed in aluminium-foil-laminated
pouches and sealed. Recent improvements in dryer design can reduce drying time
to 8 h. As a result, up to three batches of prepared mango slices can be processed
daily instead of only two batches every 36 h. The process for the production of
dried mangoes has been described by Raymundo et al. (1999).

Mango fruit bar

The product is also commonly referred to as mango ‘leather’ (Plate 83). The
processing of mango fruit bar has also been described previously by Amor-iggi
(1992) and Raymundo et al. (1999, unpublished work, 2006). Purée pro-cessed
from ripe ‘Carabao’ mango is the preferred raw material for the manufacture of
mango fruit bar in the Philippines, although ‘Pico’ is also suitable because of the
orange colour of the purée.
The total solids content of 1 t of mango purée is adjusted to 25 Brix with the
use of a hand refractometer by adding sugar to the purée. The amount of sugar
required depends on the initial total solids content of the mango purée. Then 2 kg
each of citric acid and sodium metabisulfite are blended into the purée. Juice of
calamansi, also known as the Philippine lemon (× Citrofor-tunella microcarpa Wij.
(Bunge); Citrus mitis Blanco) may be used instead of citric acid at the rate of c.20
kg per batch, although the resulting cost of the product will be slightly higher.
There is no real difference in the appearance
632 L.C. Raymundo et al.

and flavour of the finished product. Citrus juice is generally utilized if the client
prefers an all-natural product.
The prepared purée is heated for 2 min at 80°C with constant stirring to avoid
scorching. The hot mixture is carefully transferred to stainless-steel drying trays.
The trays are loaded into the dryer and dried for 14 h at 55°C. At the end of the
drying operation, the dried sheets of mango should be pli-able and should not stick
to the fingers when touched.
The sheets of mango are removed from the trays. Six layers of the dried mango
leather are stacked on top of each other. The edges are trimmed and bars measuring
2 × 4 cm are cut from the stack. Each bar is coated with con-fectioner’s sugar,
wrapped in cellophane then packed in labelled plastic bags.

Mango fruit roll

The processing of mango fruit roll has been described previously (UPLB, 1996;
Raymundo et al., 1999, unpublished work, 2006). The product and pro-cess are
similar to mango fruit bar, and they only differ in the presentation. The total solids
content of 1 t of mango purée is adjusted to 25 Brix using a hand refractometer by
adding sugar to the purée. The amount of sugar needed depends on the initial total
solids content of the mango purée. Then 2 kg each of citric acid and sodium
metabisulfite are blended into the purée. As with mango fruit bar, citric acid may
be replaced by calamansi juice at the rate of c.20 kg per batch without affecting the
overall sensory quality of the fruit roll, if the client specifies an all-natural product.

The prepared purée is then heated for 2 min at 80°C with constant stir-ring to
avoid scorching. The hot mixture is carefully transferred to stainless-steel drying
trays. The trays are loaded into a convection dryer and dried for 12–16 h at 55°C.

The dried sheets of mango are removed from the trays. Confectioner’s sugar is
sprinkled over a stainless-steel-topped table to avoid sticking of the sheets on
subsequent rolling. Each sheet is rolled manually into 1 cm diameter pieces. The
ends are trimmed; and each roll is sliced into 5 cm long pieces. The pieces are
coated with confectioner’s sugar and wrapped with cellophane. The rolls are then
packed in appropriate containers.

Vacuum-puffed dried mango

The use of vacuum for puffing and drying mango and similar food materials is not
as widespread as explosion puffing (Eskew et al., 1963), high tempera-ture short
time (HTST) pneumatic drying and centrifugal fluidized bed (CFB) drying (Brown
et al., 1972), because the vacuum-puff dryer is still expensive. With the increasing
consumer demand for high-quality processed foods and their willingness to pay a
higher price for such products, vacuum-puff dehy-dration could become an
economically viable investment opportunity for entrepreneurs, particularly in the
mango processing industry.
Fruit Processing 633

Vacuum-puffed mango pieces will readily rehydrate due to the porosity


created during the puffing and drying stages of the process. The significantly
shorter drying time (Candelaria and Raymundo, 1994b) also makes possible the
drying of four loads of prepared mango slices in 24 h.
The ripe mangoes are washed thoroughly in chlorinated H 2O, then sliced
either mechanically or with a stainless-steel knife. The flesh is scooped from the
skin with a sharp stainless-steel ladle (Candelaria, 1993; Candelaria and
Raymundo, 1994a). The fruit pieces are then heated to 90°C in 30 Brix syrup
containing 1% sodium metabisulfite, and steeped in the syrup for 4–6 h. The
mango slices are removed from the syrup, rinsed briefly in H 2O, arranged in
stainless-steel trays and loaded into the vacuum oven.
The mango pieces thus prepared are initially heated at a positive pres-sure of
40–50 kPa until the maximum tissue temperature of 100°C is reached, usually
within 8 min. The pressure is released and the hot mango pieces are dried at −70 to
−80 kPa vacuum at a temperature of 45–50°C. Total dehydration time under
vacuum is 6 h. The above pressure-temperature combinations pro-vide the most
desirable puff and rehydration characteristics (Candelaria and Raymundo, 1994b).

With the present technology, vacuum-puffed dried mango from a 1 t batch of


prepared mango slices is more expensive to produce than convec-tion oven-dried
mango (Table 17.1). The production cost needs to be reduced for the product to be
market competitive. Research that focuses on devising a system to assure a
continuous supply of mango fruits is required. The plant must operate on a year-
round basis in order to optimize the use of its equip-ment and facilities.

The facilities can be used for the production of other vacuum-puffed fruits (i.e.
bananas and muskmelon) as well as vegetables (i.e. white potato and maize
kernels) (Candelaria, 1993; Candelaria and Raymundo, 1994b) among others,
which can be used as raw materials in the manufacture of instant foods.

Table 17.1. A comparison of the profitability of different mango product lines.

Cost of goods Gross profit Net profit before


Product Volume (kg) sold (US$) (US$) tax (US$)
a
Mango powder 124,800 251,460 966,540 439,983
a
Instant mango juice 202,890 467,355 1,560,645 965,088
a
Green mango powder 141,221 245,226 1,166,984 600,216
a
Instant green mango shake 124,800 209,468 1,308,532 473,975
b
Vacuum-puffed dried mango 124,800 1,141,807 730,193 246,588
c
Dried mango 67,392 398,492 140,644 84,386
c
Mango fruit bar 115,200 921,600 461,816 277,090
c
Mango fruit roll 115,200 921,600 461,816 277,090
a US$10/kg powder.
b US$15/kg.
c
US$8/kg.
634 L.C. Raymundo et al.

17.3 Spray-dried Mango Powders


Spray-drying is a process in which a liquid feed is finely dispersed or atomized to
form droplets, which are eventually sprayed into a heated air chamber. The process
facilitates the rapid evaporation of H2O from the feed droplets, thereby forming the
powder particles. The product obtained using the technology is a free-flowing
powder that may or may not be instantly soluble in H 2O, depend-ing on the
formulation of the liquid feed that has been used. By modifying the formulation
and adjusting the process parameters, a plain powder is obtained that can be dry-
mixed with sugar, modified starches or similar components. The powder is used for
flavouring confectioneries and pharmaceutical prepa-rations as well as in the
manufacture of baby foods and tropical fruit drinks fortified with nutrients to
replace those portions lost during processing.
Tropical fruit juice powders that are rapidly soluble in H 2O are produced
directly by spray-drying fruit juices and purées. By dry-mixing spray-dried plain
fruit powder with sweeteners, a ready-to-drink juice is made. The latter method has
the added advantage in that it allows formulation of exclusive blends of fruit
drinks. As a result, consumers have a wider range of products from which to
choose that will suit their individual preferences.
By converting fruit pulps into powder or by instantizing them, their shelf life is
prolonged. Consequently, this value-adding step simplifies exportation since many
of the restrictions normally imposed on fresh produce by import-ing countries are
offset by the process. Transportation cost is also reduced by at least 85%, which
reflects the amount of H2O removed from the fresh juice. Instant juices are more
convenient for consumers since they can be reconsti-tuted easily. In the
pharmaceutical and cosmetics industries, there is a large demand for natural
tropical fruit powders as flavouring and colouring agents to add to their usual line
of fruit-flavoured products. Powders are also more convenient to handle during the
manufacturing process.
Spray-drying is by far the most cost-effective method for transforming fruit
pulps into powder. Fruit pulp is very heat-sensitive and requires special treat-ments
to produce competitively priced powders of superior quality. Further-more,
because of the rapid drying cycle and simplicity of operation, continuous
production is achieved which contributes to the low operational cost. The short
holding time of the powder inside the drying chamber reduces the risks of pow-der
burn. Human contact with the liquid feed and powder is also minimized as a result
of the short holding time. The processes are, therefore, very hygienic and the
product is ready for packaging as it leaves the dryer. Unlike other drying sys-tems,
including convection oven-drying and drum-drying, there is no need to purchase a
hammer mill for grinding the dry flakes into a powder. The main drawback of
spray-drying, however, is the relatively high initial investment cost.

Spray-dried mango fruit powder and instant mango juice

Khalid and Sial (1974), Anonymous (1998) and Diaz (2000) have discussed the
recovery of fruit powder and production of instant mango juice powder
Fruit Processing 635

using the technology. Both products use mango purée as the raw material. They
differ only in the composition of the liquid feed. The liquid feed is mango purée
with the total solids adjusted to the right consistency, thereby allowing the purée to
be discharged through a high-speed nozzle in the form of fine droplets into the
drying chamber that quickly dries to a yellow free-flowing powder. The patent
application for the manufacture of spray-dried mango powders is still pending at
the Philippine Patent Office. The process parameters used, therefore, cannot be
discussed in detail in this chapter. The parameters used are the same as reported
previously, namely, the inlet tem-perature, the feed rate, air speed and the outlet
temperature (Welti and Laflu-ente, 1983; Liang and King, 1991; Anonymous,
1998). The powder of instant mango juice is comparatively dense so that it gets wet
easily, enabling it to sink immediately during reconstitution. The plain mango fruit
powder, on the other hand, does not disperse as readily; it has a tendency to clump
on the H2O surface. However, it will dissolve quickly in hot H 2O or with the use
of an ordinary household blender.

A modest capacity spray-drying plant equipped with a NIRO SD 12.5R model


spray-dryer made in Denmark, valued at approximately US$207,860, with an
evaporative capacity of 25 kg H2O/h will require an initial invest-ment of about
US$313,860 to make it operational. In addition, if fresh man-goes are be used as
raw materials, a purée processing facility will cost around US$98,000. The figure
represents expenses for buying and installation of equipment, and does not include
the costs of the building, land and ancillary expenses such as environmental
pollution control system and spray-dryer accessories.

At the evaporative capacity rate of 25 kg H2O/h, the facility can produce


124.8 t/year of plain mango powder and 202.8 t/year of instant mango juice at a
total cost of US$251,460 and US$467,355, respectively (Table 17.1). The net profit
for plain mango powder is US$439,983 and US$965,088 for instant mango juice
before taxes.

Spray-dried green mango powder

The purée of green mangoes can be converted to a powder (UPLB, 2005) just like
the purée of ripe mangoes by spray-drying (Plate 85a). The spray-dried powder can
be mixed with other condiments and used as a souring agent for exotic or native
dishes, or as the raw material in the manufacture of instant green mango shake (see
below). The powder may be dry-mixed with sugar, powdered honey, caramel
powder or powdered sugar syrup to instantize it. During this process, the mixture
can be fortified with vitamins (e.g. ascorbic acid) and other nutrients.

The estimated cost of goods sold using a commercial spray-dryer with an


evaporative capacity of 25 kg H2O/h is US$245,226/year (Table 17.1). At the rated
capacity of the spray-drying plant of 141,221 kg/year, the net profit before taxes
for green mango fruit powder is US$600,216 at a selling price of US$10/kg of
powder.
636 L.C. Raymundo et al.

Spray-dried instant green mango shake

Green mango shake on cracked ice is a very popular thirst quencher in South-east
Asia and elsewhere (Plate 85b). Its origin can be traced back to the coun-tryside
where finely diced fresh green mango pieces are mixed with H2O, sugar and ice to
make a cheap, wholesome summer drink during the mango season. The practice
has since been modified and upgraded to cater to upscale domestic markets and
abroad.
The beverage is an excellent source of vitamin C. When the green mango
purée is spray-dried, a free-flowing white to greenish-white powder is pro-duced
that will dissolve instantly even in cold H2O. The powdering process offers
unprecedented convenience to the consumer, especially when the pow-der is
packed in sachets. Since no artificial or synthetic colours and flavour-ing agents are
included in the liquid-feed formulation, the natural taste and aroma of green mango
is retained in the powder.
Using a commercial spray-dryer with an evaporative capacity of 25 kg H2O/h,
the estimated cost of goods sold is US$209,468/year. At the rated capacity of
spray-drying plant of 124,800 kg, the net profit before taxes for instant green
mango shake is US$473,975/year at a selling price of US$10/kg of powder (Table
17.1).
The technology for the production of instant green mango shake and green
mango powder was developed at UPLB (2005). If the proper feed for-mulation and
process parameters are applied, spray-drying is an efficient and hygienic method
for producing cheap but high-quality mango fruit powder and instant mango juice.
Table 17.2 demonstrates that the spray-dried mango powders have much lower
microbial load, i.e. 2.8–10.4 × 102 colony forming units (cfu)/g, compared to the
industry standard of 1 × 104 to 5 × 105 cfu/g total plate count. Except for mango
powder, the mould and yeast counts are within the limit of 1 × 102 cfu/g. Locally
refined sugar and imported modified starch, on the other hand, have much higher
mould and yeast counts than the Heinz standard for powdered products (Shapton

Table 17.2. Microbial load of raw materials and spray-dried mango fruit powder and instant
mango juice (cfu/g).

Material Total plate count Mould count Yeast count


2 2
Mango powder 2.8 × 10 2 × 10 <10
2 2
Instant mango juice 3.6 × 10 0.4 × 10 <10
2 2
Green mango powder 10.4 × 10 0.95 × 10 <10
2 2 2
Instant green mango shake 9.8 × 10 0.85 × 10 1 × 10
2 2
Mango purée 0.5 × 10 0.2 × 10 <10
2 2
Green mango purée 0.8 × 10 0.1 × 10 <10
2 2 2
Sugar, refined 11 × 10 43 × 10 20 × 10
2 2 2
Modified starch 63 × 10 7.5 × 10 5 × 10
4 5 2 2
H.J. Heinz standard (Shapton and 1 × 10 to 5 × 10 1 × 10 1 × 10
Shapton, 1991)
Fruit Processing 637

and Shapton, 1991) at 11 × 102 and 63 × 102 cfu/g total plate count, respec-tively.
These materials are essential ingredients of the liquid-feed formula-tion used for
the production of the spray-dried powders.
The formulation and processing of numerous mango products popu-lar in
South Asian countries were reported previously (Nanjundaswamy, 1997).

17.4 Capital Investment Costs


An initial investment of approximately US$100,000 is needed for the estab-
lishment of a mango dehydration plant. The facility can also be used for dry-ing
other farm commodities such as vegetables, spices and other high-value crops. The
cost of acquiring the land, building construction and ancillary costs such as
environmental pollution abatement as well as quality assurance laboratory facilities
are not included in the estimate.
A commercial dryer such as that manufactured by Tsung Hsing Food
Machinery Co. Ltd of Taiwan with a loading capacity of 500–600 kg prepared
mango slices costs c.US$11,000. At the rated capacity of the plant of 1 t per batch,
67,392 kg of dried mango pieces can be produced annually. The cost of goods sold
is US$398,492 with a net profit before taxes of US$84,386/year. If the facility is
used for the manufacture of mango fruit bar and mango fruit roll, the annual
production is 115,200 kg of either fruit bar or fruit roll with a net profit before
taxes of US$277,090/year (Table 17.1).
The total cost of equipment for the vacuum-puffing plant with a loading
capacity of 1 t is approximately US$122,000, broken down as US$100,000 for
fabrication of the dryer and US$22,000 for other plant equipment. The total
capitalization, which includes the cost of raw materials, salaries and equip-ment
per year, is US$648,240 excluding interest on capital.
Spray-dryers come in various capacities ranging from NIRO SD Micro Spray-
dryer ideal for powdering small quantities of raw materials for research and
development to units that produce powders at evaporative rates in excess of 500 kg
H2O/h. The investment cost for equipment increases with the capac-ity of the dryer
(Table 17.3). For a small-scale operation producing powders or flours from
commercial purée, the initial investment is US$182,000 for a 5 kg H 2O evaporative
capacity dryer while a plant equipped with a 12 kg evapora-tive capacity dryer
requires US$262,000 for equipment. On the other hand, an investment of
US$313,860 is needed to equip a medium-scale drying plant with a 25 kg H2O/h
evaporative capacity such as NIRO SD 12.5R.
The estimates do not include land, building and construction costs as well as
the cost of pollution statement facilities. The cost of land is dependent on where the
plant will be located while building and construction costs are determined by the
capacity of the spray-dryer unit to be acquired.
Total capitalization is US$317,200, US$535,947 and US$806,265 for the 5 kg,
12 kg and 25 kg H2O/h evaporative capacity units, respectively. The estimate
includes the cost of raw materials, salaries and equipment per annum. Interest on
capital is not included.
638 L.C. Raymundo et al.

Table 17.3. Basic equipment for the spray-drying plant.

Evaporative rate

Equipment and accessories 5 kg H2O/h 12 kg H2O/h 25 kg H2O/h

NIRO Production Minor 110,000 – –


NIRO SD 6.3R – 156,000 –
NIRO SD 12.5R – – 207,860
Spray-dryer accessories 30,000 25,000 25,000
Homogenizer/emulsifier 20,000 45,000 45,000
Packaging equipment 5,200 5,200 5,200
Refrigerated rooms (modular, 1 t capacity, 5,000 10,000 10,000
@ US$2,500/unit)
Walk-in freezers (modular 1 t capacity 4,000 8,000 8,000
@ US$4,000/unit)
Stainless steel kettles (500 l capacity, 5,000 10,000 10,000
with stirrer @ US$2,500/unit),
Stainless steel work tables 2,800 2,800 2,800
Total 182,000 262,000 313,860

Table 17.4. Estimated volume of raw material needed for the operation of a mango
spray-drying and a mango dehydration plant.

Mango purée/slices Fresh mango

Product line kg/month kg/year kg/month kg/year

Mango powder 21,000 252,000 42,000 504,000


Instant mango juice 19,500 234,000 39,000 468,000
Green mango powder 9,100 109,200 18,200 218,400
Instant green mango shake 7,800 93,600 15,600 187,200
Vacuum-puffed dried mango 104,000 1,248,000 208,000 2,496,000
Dried mango 34,670 416,040 69,340 832,080
Mango fruit roll 24,000 288,000 48,000 576,000
Mango fruit bar 24,000 288,000 48,000 576,000

1.5 Raw Material Requirements of the Mango Processing Plant


The amount of mango purée, mango slices and fresh mango required monthly and
annually to run the plant continuously varies with the prod-uct line (Table 17.4). In
general, more raw materials are needed to process dried mango, mango fruit roll
and mango fruit bar compared to the spray-dried mango powders. As a result, the
net income is lower than that derived from the production and sale of spray-dried
mango powders (Tables 17.1 and 17.4).
Fruit Processing 639

The spray-drying facility requires 187.2–218.4 t/year of green mangoes and


468–504 t/year of ripe mango fruits at the evaporative capacity of 25 kg H 2O/h to
produce the volume of powders in Table 17.1. On the other hand, 576 t/year of ripe
fruits are required to produce mango fruit roll or fruit bar. For dried mango, 832
t/year of fresh fruits are needed to produce 67.4 t/year of the product while to
produce 124.8 t/year of vacuum-puffed dried mango, 2496 t/year of fresh fruits are
necessary.
In terms of area, c.38–48 ha can supply the green mango requirement of the
spray-drying facility. Approximately 100 ha are needed to produce 124.8 t of ripe
mango powder or 202.9 t of instant mango juice from fresh ripe mango fruits. The
fresh fruit requirement for the manufacture of fruit roll and fruit bar can be
supplied by 115 ha of mangoes. The 832 t and 2496 t of fresh fruit are equivalent to
the annual harvest from 166 ha and 500 ha of mangoes to produce 67.4 t and 124.8
t of dried mango and vacuum-puffed dried mango, respectively. The estimates are
based on a yield of 5 t from 100 trees/ha as well as the biennial fruiting habit of
mango.

17.6 Conclusion
Processing of mango is a profitable business venture once economies of scale are
attained, i.e. when the processor has the proper proportions of raw materials, labour
and machinery to meet a given market demand. Fresh mangoes are now available
year-round. But the supply is still insufficient to satisfy the demand by the fresh
fruit market and the mango-processing sector. It is, therefore, essential that a
farming system for mango be designed in order to minimize the cost of production
of off-season fruits and ensure the sustainable operation of mango processing
facilities. Since raw material sourcing is the primary cause of the dif-ficulties
encountered by processors of dried mangoes, mango in syrup and similar products,
growers need more assistance for them to adopt the tech-nology for off-season
mango flower induction. The problem is not as serious for product lines utilizing
mango purée as starting material since the purée can be processed during the peak
of the harvest season and held in cold stor-age for later use. Once the system is in
place, the processed mango industry is expected to develop further and become a
major revenue generator.
In the near future, with the advances in the field of genetic engineering, it may
be possible to eliminate the biennial fruiting habit of many current mango cultivars
or, at the very least, minimize its influence on the perfor-mance of the crop.
Species of Mangifera and some non-cultivated, wild types of mango can be the
source of desirable traits that may be incorporated in the next generation of mango
cultivars, such as their innate ability to bear fruits during the rainy season (see
Bompard, Chapter 2, this volume).
The technologies for processing mangoes are readily available. Others are
being developed in research laboratories to cope with the changing needs of
consumers. The main problem is to ensure a continuous supply of high-quality
fresh mango fruit in order to produce the prime quality commodities that
consumers expect from the industry.
640 L.C. Raymundo et al.

References

Amoriggi, G. (1992) The marvelous mango bar. Ceres 24, 25–28.


Anonymous (1998) Instruction Manual for Spray Drying Plant. NIRO A/S Soeborg, Denmark.
Barba, R.C. (1974) Induction of flowering of the mango by chemical spray. Proceedings
of the Crop Science Society of the Philippines 5, 154–160.
Brown, G.E., Farkas, D.F. and De Marchena, E.S. (1972) Centrifugal fluidized bed.
Blanches, dryers and puff piece-form foods. Food Technology 26, 23–30.
Candelaria, N.M. (1993) Dehydration of vacuum-puffed fruits and vegetables. PhD
dis-sertation, University of the Philippines Los Baños, Laguna, the Philippines.
Candelaria, N.M. and Raymundo, L.C. (1994a) Comparative drying and reconstitution,
characteristics of some fruits and vegetables. Philippine Agriculturist 77, 321–326.
Candelaria, N.M. and Raymundo, L.C. (1994b) Vacuum puffi ng and dehydration of
fruits and vegetables. Philippine Agriculturist 77, 251–260.
Diaz, J.V. (2000) Development of spray-dried instant mango (Mangifera indica L.) juice pow-
der. MSc thesis, University of the Philippines Los Baños, Laguna, the Philippines.
Eskew, R.K., Cording, J., Jr and Sullivan, J.F. (1963) A gun for explosive, puffing of
dehy-drated fruits and vegetables. Food Engineering 35, 91–96.
Khalid, M.A. and Sial, M.B. (1974) Spray-drying of mango juice powder.
Mesopotamia Journal of Agriculture 9, 47–56.
Liang, B. and King, L.J. (1991) Factors influencing flow patterns, temperature and
conse-quence drying rates in spray-drying. Drying Technology 9, 1–27.
Nanjundaswamy, A.M. (1997) Processing. In: Litz, R.E. (ed.) The Mango: Botany,
Pro-duction and Uses. CABI International, Wallingford, UK, pp. 507–542.
Raymundo, L.C., Ombico, M.T. and De Villa, T.M. (1999) Establishment of integrated mango
processing plant. In: Namuco, L.O. and Andam, C.J. (eds) Mango Production Manual.
Philippine Council for Agriculture, Forestry and Natural Resources Research and
Development (PCARRD), Los Baños, Laguna, the Philippines, pp. 97–119.
Raymundo, L.C., Ombico, M.T., De Villa, T.M. and Jaen, R.M. (2006) Processes of
Fruits, Nuts, Vegetables and Other Tropical Foods. Fruit and Vegetable
Laboratory, Food Science Cluster, University of the Philippines Los Baños,
Laguna, the Philippines. (Unpublished)
Shapton, D.A. and Shapton, N.F. (1991) Principles and Practices for the Safe
Processing of Foods. Butterworth-Heinemann Ltd, Oxford, UK.
University of the Philippines Los Baños (UPLB) (1996) Standardization of Processing
Method and Evaluation of Different Packaging Material and Storage Conditions for
Mango Roll. Annual Report, Institute of Food Science and Technology, College of
Agriculture, University of the Philippines Los Baños, Laguna, the Philippines.
University of the Philippines Los Baños (UPLB) (2005) Spray-dried Instant Juice and
Powder from Green Mango. Annual Report, Food Science Cluster, College of
Agri-culture, University of the Philippines Los Baños, Laguna, the Philippines.
Welti, J.S. and Lafluente, B. (1983) Spray-drying of comminuted orange products. I.
In-fluence of air and temperature and feed rate on product quality. Revista de
Agro-nomica y Technologia de Alementos 23, 97–106.
18 Biotechnology
R.E. Litz,1 M.A. Gómez-Lim2 and U. Lavi3
1
University of Florida, Florida, USA
2
CINVESTAV, Irapuato, Mexico
3
Agricultural Research Organization, Bet Dagan, Israel

18.1 Introduction 641


18.2 Cell and tissue culture 643
Organogenesis 643
Somatic embryogenesis 643
Protoplast isolation and culture 650
Potential for other regeneration pathways 651
18.3 Molecular Breeding and Genetics 651
Marker assisted selection (MAS) 651
Gene cloning 651
Genomics 655
18.4 Genetic Engineering 656
In vitro induced mutations 656
Genetic transformation 659
18.5 In vitro conservation 660
Medium-term storage 660
Long-term storage 661
18.6 Conclusions 662

18.1 Introduction
Improvement of monoembryonic mangoes by selection of superior seedling trees
resulting from open pollinations dates from approximately 500 years ago (the late
1500s), when the Mogul emperor Akbar established the Lakh Bagh, a garden of
seedling mango trees near Dabangha in Bihar (see Mukher-jee and Litz, Chapter 1,
this volume). Only a few years before this time, the Portuguese had introduced
grafting techniques into India, and the superior selection mango trees within the
Lakh Bagh were vegetatively propagated and named. Among these early mango
selections were ‘Alphonso’, ‘Dashe-hari’, ‘Langra’, etc. All subsequently named
mango selections have been
 CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 641
642 R.E. Litz et al.

derived by identifying seedling trees within populations either from within


uncontrolled open pollinations or, less commonly, among controlled polli-nated
progenies.
Mango cultivar improvement programmes currently exist in several countries
(see Iyer and Schnell, Chapter 4, this volume), and they address significant
production problems that have a genetic basis. Classical breeding of mango has
obvious limitations, which include, the long juvenile period of mango trees (7 or
more years), the low frequency of fruit set following con-trolled pollination, the
period required for seedling trees to be evaluated for fruit production, tree
architecture, and the cost of maintaining large popu-lations of seedling trees in
order to observe segregation of important hor-ticultural traits. There is no single
ideotype for mango; however, the most important attributes for scion cultivars must
include: compact size, resistance to anthracnose and other limiting diseases, fruit
production (which would include annual bearing and factors that affect fruit
quality, i.e. shape, colour, flavour and size). In many traditional mango-producing
countries, consumer preference for mangoes has been quite conservative, and has
been resistant to the introduction of new cultivars. Although there is a demand for
improved clonal rootstocks, there are few breeding programmes that address this
need. For mango rootstocks, the most important traits include tolerance of abiotic
stress, polyembryony and the ability to limit vegetative growth of the scion (i.e.
dwarfing).

Biotechnology refers to the application of molecular biology and somatic cell


genetics to the improvement of plants. Biotechnology can resolve some of the most
serious production problems of important mango cultivars and improve breeding
methodologies. Genomic studies will ultimately associate genes with specific
functions, and this will impact genetic engineering and molecular breeding of
mango. Marker assisted selection (MAS) would facili-tate the screening of seedling
populations for important horticultural traits. Genetic engineering would permit the
targeting and alteration of specific horticultural traits in existing cultivars, without
altering the integrity of clones. Mango improvement by modern genetics will be
freed from the constraints of the lengthy juvenile period of the species and the
additional years required for tree evaluation. Moreover, the efficient management
of mango plant genetic resources should be greatly facilitated within the next
decade by advances that have been made in molecular biology, cell culture and
cryo-preservation.

Mango biotechnology has been variously reviewed since 2002 (Litz and
Gomez-Lim, 2002, 2005; Litz, 2003, 2004, 2008; Gomez-Lim and Litz, 2007;
Krishna and Singh, 2007). In this chapter, we have addressed the current sta-tus of
mango genomics, gene cloning and cell culture. Within the discussion of cell
culture, the following areas of research are addressed: (i) the utiliza-tion and
potential of cell culture for improving existing mango cultivars by in vitro-induced
mutation followed by selection and genetic transformation; and
(ii) advances in medium- and long-term conservation of genetic resources. The
model species Arabidopsis thaliana has provided a useful tool for studying the
regulation of gene expression of important plant processes, for example
Biotechnology 643

embryo and organ development, including shoots, leaves, roots, flowers and fruit.
Genomic analysis and the identification of horticulturally important genes that
control these and other processes will ultimately enable us to under-stand and
control many aspects of mango fruit production (i.e. disease and pest control, fruit
quality, tree architecture, etc.). The development of molecu-lar approaches will
also facilitate mango breeding through the identification of DNA markers for
important traits.
This chapter is a discussion of the current state of mango cell culture, gene
cloning and the manipulation of cell cultures to address plant breeding objectives
by genetic engineering.

18.2 Cell and Tissue Culture


Organogenesis

The earliest report of morphogenesis from mango cell cultures involved the
differentiation of adventitious roots from callus that was initiated from mango
cotyledons on Murashige and Skoog (1962) semi-solid induction medium (MS)
that was supplemented with kinetin and naphthaleneacetic acid (NAA) (Rao et al.,
1982). Of greater significance was the later report that callus initiated from leaves
from mature mango trees also had morphogenic potential. Raghuvanshi and
Srivastava (1995) induced caulogenic cultures from fully expanded, young leaves
of monoembryonic ‘Amrapali’. Disin-fested leaf pieces were initially explanted
into liquid MS medium supple-mented with the antioxidant 0.05%
polyvinylpyrolidone with 2 h subcultures for up to 24 h on a rotary shaker at 75
rpm. Leaf pieces were then transferred onto MS semi-solid medium supplemented
with 13.0 PM kinetin and 1.1 PM indole acetic acid (IAA), the optimum growth
regulator formulation for mul-tiple shoot development from leaf callus. Subculture
of individual shoots onto semi-solid MS medium supplemented with 9.8 PM indole
butyric acid (IBA) stimulated root induction and development. Cultures were main-
tained at 25C with a 16 h photoperiod provided by cool white fluorescent lights
(50–70 Pmol/m2/s).

The major advantage of the organogenic pathway for regenerating mango is


that cultures can be induced whenever there is a leaf flush; however, genetic
manipulation studies require much higher rates of regeneration than has been
hitherto reported.

Somatic embryogenesis

Genetic engineering of mango has been based upon the efficient recovery of
somatic embryos from embryogenic cultures. Efficient regeneration of woody
plants from cell cultures derived from mature phase materials is often difficult to
achieve, and the optimum procedure generally must be determined empir-ically.
Mangifera indica consists of two ecogeographic races: a monoembryonic
644 R.E. Litz et al.

group that probably originated in north-eastern India and northern Myan-mar and a
polyembryonic group that originated in South-east Asia. The pres-ence of nucellar
embryos in seeds of polyembryonic mangoes demonstrates that cells of the
nucellus of the species have morphogenic potential (see Mukherjee and Litz,
Chapter 1, this volume). Sturrock (1968) reported that polyembryony in mango
appeared to be inherited as a recessive trait; how-ever, Aron et al. (1998) have
demonstrated that polyembryony is under the control of a single dominant gene.
Somatic embryogenesis, which is the equiv-alent morphogenic response in vitro, is
a dominant trait in several other spe-cies, for example lucerne (Reisch and
Bingham, 1980) and orchardgrass (Gavin et al., 1989), although in red clover it
apparently is conferred by a reces-sive gene (Broda, 1984). The presence of
morphogenically competent cells in the nucellus is critical for the induction of
embryogenic mango cultures of both monoembryonic and polyembryonic mango
cultivars.
Induction of embryogenic mango cultures from the excised nucellus of
immature mango seeds of polyembryonic and monoembryonic cultivars was first
described by Litz et al. (1982) and Litz (1984), respectively. The current standard
protocol for induction and maintenance of embryogenic mango cultures and for
development of somatic embryos is derived from DeWald et al. (1989a, b) and Litz
et al. (1993). Table 18.1 provides citations for all in vitro studies that have
involved somatic embryogenesis of mango. The efforts in this field have expanded
considerably since the late 1990s.
It is impossible to predict the embryogenic response of the nucellus of
different mango cultivars, irrespective of the seed type (i.e. polyembryonic or
monoembryonic). Early reports suggested that the nucellus of polyembry-onic
mangoes responded in vitro more readily than the nucellus of monoem-bryonic
mangoes (Litz, 1986); however, this assumption no longer is considered to be
valid. Litz et al. (1997) reported that the induction of embryogenic com-petence in
the explanted nucellus of monoembryonic ‘Tommy Atkins’ can be inhibited by the
ethylene antagonist aminoethoxyvinylglycine (AVG) and by
dicyclohexylammonium sulfate (DCHA), an inhibitor of spermidine synthe-sis. In
contrast, induction of embryogenic competence of the explanted nucel-lus of
polyembryonic ‘Tuehau’ is unaffected by either AVG or DCHA. Litz and Schaffer
(1987) demonstrated that somatic embryogenesis in mango is partially mediated by
spermidine. The biosynthesis of ethylene and/or the sensitivity of nucellar tissue to
ethylene may be important factors for the induction of embryogenic cultures from
this tissue.

Induction
The induction of embryogenic competence is associated with the develop-mental
stage of the nucellus at the time of explanting, and also can be af-fected by the
physiological status of the tree (Litz, 1987). Fruits that are approximately 30–40
days after pollination contain seeds in which the nucel-lus is at the ideal stage for
explanting: relatively thick and intact and easily removed. The embryo (mass) in
fruit and developing seed of this age is usu-ally no more than half the length of the
immature seed. Mango fruit of the appro-priate stage of development are surface-
disinfested with 20–30% (v/v) domestic
Biotechnology 645

Table 18.1. Summary of published reports on somatic embryogenesis of mango


from nucellar explants.

Country Reference

Colombia Flórez-Ramos et al. (2007)


India Ara et al. (1998, 2000a, b)
Chaturvedi et al. (2004)
Deore et al. (2000)
Jana et al. (1994)
Krishna and Singh (2007)
Laxmi et al. (1999)
Singh et al. (2001, 2002)
Sulekha and Rajmohan (2003)
Thomas (1999)
Mexico Manzanilla Ramirez et al. (2000)
Rivera Domínguez et al. (2004)
Philippines Patena et al. (2002)
USA DeWald et al. (1989a, b)
Lad et al. (1997)
Litz (1984)
Litz and Gomez-Lim (2005)
Litz and Lavi (1997)
Litz and Schaffer (1987)
Litz and Vijayakumar (1988)
Litz et al. (1982, 1983, 1984, 1995, 1997, 1998)
Mathews et al. (1992)
Monsalud et al. (1995)
Pliego Alfaro et al. (1996b)

bleach containing Tween 20 for 30 min. The sterilant is rinsed from the fruit with
three changes of sterile deionized or distilled water, and each fruit is bisected along
its longitudinal axis without damaging the immature seed under axenic conditions.
The immature seed is removed from the bisected fruit, which is also bisected
carefully along its longitudinal axis. Manza-nilla Ramirez et al. (2000) obtained
optimum induction with polyembryonic ‘Ataulfo’, monoembryonic ‘Tommy
Atkins’ and monoembryonic ‘Haden’ nucellar explants when the embryo (mass) to
immature seed ratio was 1:3. The zygotic embryo (of a monoembryonic cultivar) or
polyembryonic mass (of a polyembryonic cultivar) is excised and discarded. The
nucellus can be carefully peeled from the interior of the seedcoat using a sterile,
flat spatula. Following the transfer of the nucellus onto induction medium in sterile
Petri dishes, the cultures are incubated in darkness at 25C. Thereafter, it is essen-
tial to subculture the nucellar explants onto fresh induction medium at daily
intervals until the oxidation of the explant ceases; oxidation is associated with
darkening of the medium around the explanted nucellus.

A summary of those mango cultivars that have been successfully estab-lished


as embryogenic cultures is provided in Table 18.2. The efficiency of
646 R.E. Litz et al.

Table 18.2. Induction of somatic embryogenesis from nucellar cultures of mango


(Source: Jana et al., 1994; Litz and Lavi, 1997; Manzanilla Ramirez et al., 2000;
Ara et al., 2000b; Chaturvedi et al., 2004).

Cultivar Seed type Cultivar Seed type Cultivar Seed type

‘Alphonso’ Mono ‘Gedong’ Poly ‘Mulgoa’ Mono


‘Ambalavi’ Poly ‘Golek’ Poly ‘Mundan’ Mono
‘Amrapali’ Mono ‘Heart’ Poly ‘Nam Doc Mai’ Poly
‘Arumanis’ Poly ‘Hindi’ Poly ‘Neelum’ Mono
‘Ataulfo’ Poly ‘Honc Cambodiana’ Poly ‘Ono’ Poly
‘Baneshan’ Mono ‘Irwin’ Mono ‘Parris’ Poly
‘Brander’ Poly ‘James Saigon’ Poly ‘Peach’ Poly
‘Brooks’ Mono ‘Keitt’ Mono ‘Philippine’ Poly
‘Cambodiana’ Poly ‘Kensington Pride’ Poly ‘Sabre’ Poly
‘Carabao’ Poly ‘Kur’ Poly ‘Simmonds’ Poly
‘Chausa’ Mono ‘Langra Benarsi’ Mono ‘Tommy Atkins’ Mono
‘Chino’ Poly ‘Lippens’ Mono ‘Tuehau’ Poly
‘Dashehari’ Mono ‘Madu’ Poly ‘Turpentine’ Poly
‘Everbearing’ Mono ‘Manzano’ Poly ‘White Langra’ Mono
‘Florigon’ Poly ‘Mikongenesis’ Poly

induction is cultivar-dependent. For example, Litz et al. (1998) compared the


induction of four mango cultivars, polyembryonic ‘Hindi’, monoembry-onic
‘Lippens’, polyembryonic ‘Nam Doc Mai’ and monoembryonic ‘Tommy Atkins’,
and observed that ‘Hindi’ has the highest embryogenic response, followed by
‘Lippens’, ‘Tommy Atkins’ and ‘Nam Doc Mai’ in descending order. Manzanilla
Ramirez et al. (2000) compared the induction responses of three mango cultivars
and observed that polyembryonic ‘Ataulfo’ was more embryogenic than either
monoembryonic ‘Tommy Atkins’ or monoembry-onic ‘Haden’ in descending
order. Optimization of conditions for induction of embryogenic mango cultures
using polyembryonic ‘James Saigon’ and polyembryonic ‘Parris’ as models was
described by DeWald et al. (1989a). The current induction procedure has been only
slightly modified since then, and utilizes a basal medium consisting of B5
(Gamborg et al., 1968) major salts without ammonium sulfate ((NH 4)2SO4), MS
minor salts and organic compo-nents, 60 g/l sucrose, 400 mg/l glutamine, 2.4–4.8
PM 2,4-dichlorophenoxy-acetic acid (2,4-D) and 2.0 g/l gellan gum. Patena et al.
(2002) modified the standard induction medium by supplementing it with 100 mg/l
coconut water (CW) to control oxidation of the explants.

The auxin 2,4-D has a temporal effect on induction of embryogenic


competence of explanted nucellus of polyembryonic ‘Carabao’ (Lad et al., 1997).
Induction requires a minimum of 7–14 days exposure to 2,4-D and a maximum
exposure period of 56 days; acquisition of embryogenic compe-tence is optimum
after approximately 28 days exposure to 2,4-D (Plate 86). Nurse cultures, which
consist of highly embryogenic mango cultures (e.g. polyembryonic ‘Hindi’) have
been effective for stimulating induction of
Biotechnology 647

embryogenic competence from the nucellus of cultivars that are normally difficult
to induce, for example polyembryonic ‘Nam Doc Mai’ (Litz et al., 1998). The
nurse culture procedure involves explanting the nucellus onto sterile filter paper
which has been moistened with induction medium and which overlays the highly
embryogenic mango culture (polyembryonic ‘Hindi’) on semi-sterile induction
medium. It is not clear if a nucellar callus is initiated from the explant prior to
acquisition of embryogenic competence; however, somatic embryos can develop
directly from the nucellus without an intermediate callus (Litz, 1987).
Embryogenic nucellar cultures are recog-nizable ap-proximately 30 days after
explanting; they are completely orga-nized, and consist of proembryonal somatic
embryos, embryogenic cells, cell aggregates and proembryonic masses (PEMs)
(Litz et al., 1993, 1995; Litz and Lavi, 1997).

Maintenance
Embryogenic mango cultures are friable and cream to light brown in colour,
although the cultures rapidly darken on semi-solid medium, and must therefore be
subcultured at 3–4 week intervals. PEMs develop from globular somatic embryos
in the presence of the primary induction agent, 2,4-D. The PEMs originate as
globular somatic embryos, but their devel-opment as individual somatic embryos is
arrested in the presence of 2,4-D. The PEMs increase in diameter with cells of the
protoderm dividing rap-idly; secondary globular somatic embryos develop from
these proliferating embryogenic cells. This highly repetitive pattern of somatic
embryogen-esis in the presence of 2,4-D is the basis for maintenance of
embryogenic cultures.

Embryogenic mango cultures can be maintained as proliferating PEMs either


on semi-solid or in liquid induction medium. Maintenance of embryo-genic
cultures of many cultivars is optimal in liquid induction medium sup-plemented
with 4.8 PM 2,4-D (Litz et al., 1984; DeWald et al., 1989a) (Plate 87); however,
rapid proliferation of embryogenic suspension cultures is cultivar-dependent (Litz
et al., 1993). Embryogenic suspension cultures are initiated by inoculating
approximately 400 mg of PEMs into sterile 80 ml maintenance medium in 250 ml
Erlenmeyer flasks (or 200 mg of PEMs into 40 ml mainte-nance medium in 125 ml
Erlenmeyer flasks). The flasks are maintained on a rotary shaker at 100–125 rpm in
semi-darkness at 25C with regular transfers of PEMs into fresh medium at 10–14
day intervals. Regular subculturing is essential to prevent loss of morphogenic
potential and darkening of the tis-sue. The typical embryogenic suspension culture
consists of PEMs, embryo-genic cells and multicellular complexes.

Maturation
Development of somatic embryos from embryogenic cultures maintained on semi-
solid maintenance medium occurs sporadically and without synchroni-zation, due
to the lack of direct contact of parts of a culture with medium containing 2,4-D.
Exposure to 2,4-D is necessary for embryogenic culture proliferation, while at the
same time, somatic embryo development is
648 R.E. Litz et al.

inhibited. In order to stimulate somatic embryo development, embryogenic cultures


must be transferred from maintenance medium to medium without 2,4-D in order
to initiate efficient somatic embryo development. For embryo-genic suspension
cultures, the PEMs are decanted through sterile filtration fabric with a 1000 Pm
opening size. The larger fraction (>1000 Pm diameter) is re-inoculated into liquid
maintenance medium for continued proliferation and the smaller fraction is
transferred either into liquid medium or onto semi-solid medium without 2,4-D in
order to arrest repetitive somatic embryogenesis and to initiate somatic embryo
development (Plate 88).
The plant growth media and conditions for stimulating somatic embryo
development and maturation are based upon the protocol described by DeWald et
al. (1989b) with minor alterations. Initially, a medium consisting of B5 major salts,
MS minor salts and organic components, 60 g/l sucrose and 400 mg/l glutamine
with or without 2.0 g/l gellan gum is utilized. Different mango cultivars require
different periods for cotyledon differentiation following subculture to maturation
medium (Litz et al., 1993). Either 4.65 PM kinetin or 4.44 PM benzyladenine
(BA) can stimulate the development of cotyledons and the apical meristem and
reduce the maturation period. The cultures are incubated in darkness at 25C.

When growth of embryogenic cultures of highly responsive cultivars is


optimized as suspension cultures, the early cotyledonary somatic embryos that
develop in liquid medium are hyperhydric. Mathews et al. (1992) and Monsalud et
al. (1995) demonstrated that hyperhydric somatic embryos can-not develop to
maturity, and become necrotic. Hyperhydricity of mango somatic embryos can be
reversed either by partially desiccating heart stage embryos (2–3 mm length) under
high relative humidity (RH) for 24 h or by plating them on maturation medium
solidified with 6.0 g/l gellan gum (Monsalud et al., 1995). Reversion of
hyperhydricity can result in precocious germination of mango somatic embryos,
although this can be prevented if 500 PM abscisic acid (ABA) is included in the
modified maturation medium.
Embryogenic cultures of different mango cultivars vary in their response to
subculture from liquid maintenance medium onto maturation medium, and this
must be determined empirically for each cultivar. For example, poly-embryonic
‘Hindi’ somatic embryo development (cotyledon differentiation) from
embryogenic suspension cultures follows a step-wise procedure. The <1000 Pm
diameter fraction from liquid maintenance is subcultured into liquid maturation
medium until the early heart stage of somatic embryo development is apparent.
Subsequently, the heart stage somatic embryos are subcultured onto semi-solid
maturation medium. In contrast, the <1000 Pm fraction of monoembryonic ‘Keitt’
and polyembryonic ‘Carabao’ embryogenic suspension cultures can be transferred
directly from liquid maintenance medium onto semi-solid maturation medium.

Mango zygotic embryos require approximately 4–5 months in order to develop


to maturity in vivo, and mature embryos can be >6–8 cm long (Plate 89).
Consequently, the plant growth media that have been formulated to stimulate
growth and development of mango somatic embryos from the heart stage to
maturity reflect the differing requirements of these enlarging
Biotechnology 649

somatic embryos. The medium that has been utilized for development of mango
somatic embryos to maturity consists of B5 major salts, MS minor salts and
organic components, 400 mg/l glutamine, 20% (v/v) filter-sterilized CW, 40 g/l
sucrose and 2.0 g/l gellan gum (DeWald et al., 1989b). As the somatic embryos
enlarge, the sucrose concentration of maturation medium is gradually reduced to 10
g/l.
During somatic embryo development certain developmental anomalies become
apparent, of which the most frequently observed are polycotyly, fas-ciation,
absence of bipolarity, secondary somatic embryogenesis from the hypocotyl and
precocious germination. Polycotyly and fasciation do not affect subsequent plant
development; however, failure to address problems associated with absence of
bipolarity, secondary embryogenesis and preco-cious germination can seriously
impact the recovery of plants. Control of precocious germination of developing
embryos is occasionally necessary, since immature somatic embryos that germinate
before they are physiologi-cally mature cannot survive. Some of these
developmental anomalies can be eliminated by maintaining relatively high sucrose
concentrations and/or by incorporating 100 PM ABA in the maturation medium
(Monsalud et al., 1995; Pliego-Alfaro et al., 1996b). The cultures are maintained in
darkness at 25C during somatic embryo maturation.

Germination
When somatic embryos begin to germinate, they are finally transferred to light
conditions. The radicle elongates, followed by growth of the taproot. The shoot
apical meristem remains quiescent for approximately 2 weeks after germination, at
which time the shoot elongates. Although many mango somatic embryos germinate
under these conditions, their survival or conver-sion ex vitro is low, primarily due
to apical shoot necrosis, a physiological disorder that is associated with calcium ion
(Ca++) deficiency. Different strat-egies have been attempted to improve the
conversion rate (i.e. survival of somatic embryo-derived plants):

1. The period for embryogenic cultures in/on maintenance medium should be


minimal (Litz and Lavi, 1997).
2. Ara et al. (1998) rescued in vitro microshoots obtained from germinated
somatic embryos by pulsing them for 24 h with 24.6 PM IBA in liquid medi-um
followed by transfer to auxin-free medium in darkness for root develop-ment.

3. Somatic embryo shoots have also been rescued by micrografting the shoots on
decapitated in vitro-germinated seedling rootstocks (Plate 90).
4. Enhanced recovery of mango plantlets can occur following the induction of
photoautotropism by transfer of small plantlets onto minimal plant growth medium,
containing <5% sucrose and 1% (w/v) activated charcoal (Litz et al., 1993). A
filter-sterilized air mixture consisting of 20,000 ppm carbon dioxide (CO 2) in a
nitrogen gas carrier is introduced into the growing containers, and the plantlets are
exposed to a 16 h photoperiod at 180 Pmol/s/m2 provided by cool white
fluorescent tubes.
650 R.E. Litz et al.

5. Ara et al. (1999) reported improved conversion following the encapsulation of


early heart stage somatic embryos in calcium alginate-containing modified
standard mango medium with half-strength major salts and supplemented with 2.9
PM gibberellic acid (GA3). The germination of encapsulated somatic embryos was
almost 75% greater than non-encapsulated somatic embryos.

Protoplast isolation and culture

The isolation, culture and regeneration of plantlets from protoplasts isolated from
embryogenic suspension cultures of monoembryonic ‘Amrapali’ has been
described (Ara et al., 2000a). One gram of embryogenic culture was transferred
from a 3–4-week-old suspension into 10 ml filter-sterilized growth medium
consisting of B5 major salts, MS minor salts and organic components, supple-
mented with 0.3 M sucrose, 0.4 M mannitol, 0.1 M sorbitol, 2.74 mM glutamine,
1.0% cellulase, 1.0% hemicellulase and 0.5% pectinase (Sigma) with gentle shak-
ing in darkness at 25C for 24 h. The digestion mixture was then passed through a
sieve (50 Pm) in order to remove debris, and then centrifuged for 5 min at 100 g.
The supernatant was discarded, and cell debris was resuspended and precipitated
by centrifugation three times at 3 min for each centrifugation cycle. The pellet was
finally resuspended in 1 ml medium, and layered on 3 ml sucrose solution (25%
w/v) and centrifuged at 100 g for 7 min. Protoplasts were removed and cultured in
maintenance medium, but modified to contain 0.18 M sucrose, 2.74 mM glutamine
and 4.5 PM 2,4-D. Somatic embryos developed from PEMs following subculture
onto somatic embryo development medium (without 2,4-D), and plantlets were
recovered using standard procedures (see Somatic embryogenesis section in 18.2
Cell and tissue culture, this chapter).
Protoplast technology has not been utilized at this time for mango
improvement, and the likelihood of its exploitation is difficult to predict. The
sexual compatibilities of Mangifera spp. with the common mango are unknown.
Many newly described Mangifera species tolerate stressful environmental
conditions and have pest and disease resistance (see Bompard, Chapter 2, this
volume); however, it is uncertain if they are isolated genetically from mango.
Somatic hybridization might enable the genetic recombination of the common
mango with some of the Mangifera species as a means for develop-ing new
rootstocks. This approach, however, would almost certainly have little utility for
scion development, since the common mango is a tetraploid, and the ploidy level of
many of the Mangifera spp. is unknown. Somatic hybridization would result in
hexaploids or octoploids, and introgression of useful traits into the common mango
would be very difficult.

Potential for other regeneration pathways

The recovery of embryogenic haploid cultures from microspores has been


described for several perennial fruit tree species, including Annona squamosa (Nair
et al., 1983), Citrus aurantifolia (Chaturvedi and Sharma, 1985), Citrus
Biotechnology 651

microcarpa (Chen et al., 1980), Dimocarpus longan (Yang and Wei, 1984) and
Litchi chinensis (Fu and Tang, 1983). Mangifera indica is thought to be an auto-
tetraploid (2n = 4x = 40), and recovery of diploid (n = 2x = 20) plants would reveal
the nature of the ancestral species and simplify genomic analysis. Chromosome
doubling would restore autotetraploidy, and somatic hybrid-ization of diploid
mangoes with wild Mangifera species would allow interest-ing genetic
recombination.

18.3 Molecular Breeding and Genetics


The identification and use of molecular markers for distinguishing mango from
other Mangifera spp. and for identifying different mango cultivars has been
discussed by Bompard, Chapter 2, this volume and by Iyer and Schnell, Chapter 4,
this volume.

Marker assisted selection (MAS)

MAS offers great potential for improvement of quantitative traits in crop plants.
There are clear advantages for the use of molecular markers in plant breeding, such
as a decreased number of breeding generations, the availabil-ity of a uniform
method for scoring, no need to use phenotypic scoring until the end and, finally,
the possibility for obtaining information on the percent-age of genome contributed
by each parent in the offspring. Although molecu-lar markers have been used for
taxonomic purposes with mango (Schnell et al., 1995; López-Valenzuela et al.,
1997; Eiadthong et al., 2000; Ravishankar et al., 2000; etc.) mango has not been
the subject of MAS. It is notable that although mango has 40 chromosomes, it has a
comparatively small haploid genome size (0.91 pg), which is only three times as
large as the recently sequenced genome of A. thaliana (the plant with the smallest
genome size known), about half that of tomato and comparable to that of rice
(Arumuganathan and Earle, 1991). Clearly, the comparatively small mango
genome would facilitate the identifi-cation of molecular markers and the creation
of a genetic map.

Gene cloning

Genetic transformation of mango cultures involves the transfer of genes to


manipulate specific processes. Genetic manipulation of any crop requires that
relevant genes must be available. Mango has been the subject of molecu-lar
research for several years, and several genes have been identified.
Early studies showed that changes in mRNA and protein content occur during
fruit ripening (López-Gómez and Gómez-Lim, 1992; Chaimanee et al., 1999). The
level of a number of mRNAs (as assayed by in vitro translation) changes
throughout the ripening process. This method for detecting changes in mRNAs has
low sensitivity and, therefore, only the most abundant proteins
652 R.E. Litz et al.

can be detected. For that reason, the molecular analysis of fruit ripening requires
construction of a gene library. In mango, cDNA libraries have been constructed,
mainly from ripe fruit, and screened using several approaches. The mRNA for
virtually all the ripening-related genes isolated so far have been shown to be absent
or at a low level in immature fruit, increase during ripening and decline as ripening
progresses (Giovannoni, 2001). Unlike other fruits, none of the identified genes in
mango code for enzymes involved in the ripening process itself (see below).

Mango fruit are highly perishable commodities due to over-ripening, which is


mainly caused by the sharp increase in ethylene production, which occurs
simultaneously with the climacteric peak (Tucker and Grierson, 1987; see Brecht
and Yahia, Chapter 14, this volume). Mangoes have poor storage quality and
storage in controlled or modified atmospheres has been associ-ated with
physiological disorders (Chaplin, 1989). Genetic manipulation of mango fruit
ripening represents an attractive alternative to extend storage life and, therefore, the
isolation of mango genes coding for enzymes involved in ethylene biosynthesis has
been a target for research.
The two key enzymes in the ethylene biosynthetic pathway are those catalysing
the conversion of S-adenosylmethionine (SAM) to 1-aminocyclo-propane-1-
carboxylic acid (ACC) and ACC to ethylene, i.e. ACC synthase and ACC oxidase
or ethylene forming enzyme (EFE), respectively and cDNA clones coding for
mango ACC synthase and ACC oxidase have been identi-fied (Gómez-Lim, 1993).
Their expression during ripening was studied in pulp and peel in northern blot
analysis-type experiments. The ACC synthase message is undetectable in unripe
fruit and starts to appear in turning fruit, reaching a maximum in ripe fruit (Gómez-
Lim, 1993). This pattern of expres-sion is similar in the peel and in the pulp;
however, the message appears in the pulp before the peel. The ACC oxidase
message shows similar kinetics in both types of tissue, but the message is clearly
detectable before any ACC synthase message becomes detectable (Gómez-Lim,
1993). These results sug-gest that ACC oxidase is expressed before ACC synthase
and that ripening starts on the inside of mango fruit and proceeds outwards.
Ethylene-treated mango fruits show a different pattern of expression, with ACC
oxidase and ACC synthase appearing initially in the peel.

If ethylene is being actively produced, the gas must be clearly perceived by


plant cells and therefore ethylene sensitivity is important for the ripening process.
Major advances in understanding ethylene signal transduction have come from a
molecular genetic approach using ethylene responsive mutants of A. thaliana and
tomato. Several genes coding for ethylene receptor homo-logues have been isolated
from various plants (Johnson and Ecker, 1998). Genetic manipulation of the
ethylene receptor represents an interesting alter-native to control ethylene
production and delay ripening, particularly in those fruits where other alternatives,
such as storage in controlled atmo-spheres, have not been effective (Wilkinson et
al., 1997). A cDNA coding for a mango ethylene receptor has been isolated
(Gutiérrez-Martinez et al., 2001). The message seems to be present at low levels in
unripe fruit and to increase
Biotechnology 653

as the fruit ripens. Mechanical wounding appears to up-regulate the expres-sion of


the receptor.
Several studies have convincingly shown that the profile of released volatiles
changes as ripening proceeds (Olle et al., 1998; Sakho et al., 1998; Ansari et al.,
1999; Saby John et al., 1999; Andrade et al., 2000; Bender et al., 2000). It is likely
that many of these compounds, most of which are terpe-noids, are determinants of
flavour and aroma and many of them originated from the metabolism of fatty acids
via the E-oxidation pathway. It is known that several fatty acids, particularly
linoleic and oleic acids, decrease in con-centration during ripening. In this sense, a
cDNA for thiolase, an enzyme from the E-oxidation pathway, was identified as
having a ripening-specific pattern and to be up-regulated during ripening
(Bojórquez and Gómez-Lim, 1995). A cDNA for acyl CoA oxidase, the key
enzyme in the E-oxidation path-way, has isolated from mango fruit and shown to
behave similarly to thiolase (A. Nila-Mendez and M. A. Gómez-Lim, Irapuato,
unpublished data). These enzymes might be involved in metabolism of fatty acids
to produce volatile compounds. The contents of sugars and organic acids can
influence the fla-vour properties of mango (Malundo et al., 2001).

The manipulation of fruit aroma and flavour is a long established research goal
and, accordingly, the isolation of genes coding for enzymes involved in
biosynthesis of these compounds has been targeted. The gene coding for alcohol
acyl transferase, an enzyme presumably involved in the synthesis of compounds
implicated in fruit flavour, has been identified in mango (GB: AX025510; patent
WO 0032789-A 36).
Alternate oxidase is involved in the cyanide-resistant respiratory path-way. It
has been studied mainly in thermogenic species, and its activity is correlated with
heat production, necessary to volatilize foul-smelling com-pounds to attract insect
pollinators. There is a significant participation of this pathway in the climacteric of
many fruit. A cDNA coding for mango alter-nate oxidase has been isolated and the
message was detected by Northern blot analysis in unripe fruit and shown to
increase substantially in ripe fruit (Cruz-Hernandez and Gómez-Lim, 1995). These
results were correlated with similar increases in enzyme activity and protein
accumulation. The tempera-ture in ripe monoembryonic ‘Alfonso’ fruit is up to
10C higher than in unripe fruit and this has been attributed to the activity of
alternate oxidase (Kumar et al., 1990). This extra heat might also serve to volatilize
aroma-giving compounds. The results with alternate oxidase were confirmed by
Considine et al. (2001), who isolated several members of the multigene family of
mango alternate oxidase and showed that they were expressed differen-tially during
ripening. They also identified a gene coding for an uncoupling protein whose
mRNA peaked at the turning stage whereas the protein peaked at the ripe stage
(Considine et al., 2001). They suggested a role for alternate oxidase and the
uncoupling protein in post-climacteric senescence. Because both mRNA and
protein for alternate oxidase and the uncoupling protein increased in a similar
pattern, they hypothesized that their expression is con-trolled simultaneously.
654 R.E. Litz et al.

Ripening of fruit involves a number of metabolic reactions, including synthesis


and turnover of the plant cell wall. In mango, fruit ripening is char-acterized by a
gradual softening, which is caused by progressive depolymer-ization of pectic and
hemicellulosic polysaccharides with significant loss of galactose, arabinose and
mannose residues at the ripe stage (Yashoda et al., 2005). This depolymerization is
caused by the activity of different hydrolytic enzymes that are secreted as ripening
proceeds (i.e. polygalacturonase and E-galactosidase), which are present in
different isoforms in mango pulp (Prasanna et al., 2005, 2006). Cell wall-
metabolizing enzymes from mango, i.e. exopolygalacturonase (Chaimanee et al.,
2000), endo-polygalacturonase (Sun-tornwat et al., 2000), E-galactosidase (S.
Parra-Arenas and M. A. Gómez-Lim, Irapuato, unpublished data) and pectate lyase
(Chourasia et al., 2006), have been cloned and correlate closely with fruit ripening.
Furthermore, an ethyl-ene-responsive, ripening-related expansin has also been
identified as being closely correlated with softening (Sane et al., 2005).

A cDNA clone coding for a homologue of the YPT/Rab class of small


Guanosine-5’-triphosphate (GTP)-binding proteins has been identified from mango
and is induced during ripening (Zainal et al., 1996). These proteins appear to
control the secretion of other proteins as well as the fusion of mem-branes in
animal cells (Fischer von Mollard et al., 1994). Some of these small GTP-binding
proteins have homologues in plants, albeit their role has not been well defined
(Staehelin and Moore, 1995). It is tempting to speculate that, based on the pattern
of expression, these small GTP-binding proteins facilitate secretion of various
hydrolytic enzymes during fruit ripening.
Saiprasad et al. (2004) isolated five ripening-related cDNAs by reverse
transcription-polymerase chain reaction (RT-PCR). They showed similarity to PRl-
1 protein, a transcription initiation factor, a CCR-4 protein and to an 18S ribosomal
RNA gene and a 23S ribosomal RNA gene. Likewise, Lycett et al. (1997)
identified by differential screening six clones up-regulated dur-ing ripening.
Unfortunately, none of these clones code for proteins directly associated with
ripening. Only two of them showed homology to a plastid chromatin and
Ypt/Rab11 class of small GTPase, respectively. I A tomato rab11-like cDNA was
used for tomato transformation in antisense (Lu et al., 2001). The fruits changed
colour as expected but failed to soften normally. This was accompanied by reduced
levels of two cell wall hydrolases, pectin-esterase and polygalacturonase. There
were other phenotypic effects in the plants, including determinate growth, reduced
apical dominance, branched inflorescences, abnormal floral structure and ectopic
shoots on the leaves. In some plants, ethylene production was reduced. These data
suggest an additional role for the gene in other ripening processes.

Vasanthaiah et al. (2006) reported the characterization of 27 differentially


expressed genes during mango internal breakdown and found catalase, superoxide
dismutase, ubiquitin, alcohol dehydrogenase, coproporphyrino-gen oxidase and
keratin-associated protein to be up-regulated; whereas, several ribosomal genes,
cysthathionine gamma synthase and fructose bispho-sphate aldolase were down-
regulated. The increased expression of catalase, coproporhyrinogen III oxidase and
keratin genes during internal breakdown
Biotechnology 655

was interpreted as a sign of oxidative stress, which they hypothesized as one of the
probable causes for this disorder.
In addition to these genes, several mango sequences have been reported in the
GenBank. These include a genomic sequence for the large subunit of ribulose 1,5-
bisphosphate carboxylase (U39269), cDNAs for two unidentified clones
(AF370123 and AF061639), a partial cDNA for LFY (AY189684), a cDNA for
xyloglucan endo-transglycosylase (GenBank accession AY600965) and a putative
Pto-like serine/threonine kinase gene (AY693369). The function of these sequences
in mango development remains to be determined.
There have been several reports on therapeutic properties of extracts from
mango leaves or seeds, including anti-inflammatory effects (Beltrán et al., 2004;
Garrido et al., 2004; Leiro et al., 2004), anthelminthic and antial-lergic properties
(García et al., 2003a), anti-diarrhoeal properties (Sairam et al., 2003) and even an
enhancement of the humoral immune response in mouse (Garcia et al., 2003b). It is
likely that the constituents responsible for these therapeutic effects will be the
subject of intense scrutiny and, if the right genetic elements are identified, the
genetic manipulation of the biosyn-thetic route may become a reality.

Genomics

The human genome project has been the catalyst for the development of sev-eral
high-throughput technologies that have made it possible to map and sequence
complex genomes. Many bacterial genomes and the genomes of Sacchromyces
cerevisiae, Caenorhabditis elegans, Drosophila melanogaster, A. thali-ana, Oriza
sativa and Populus trichocarpa have been fully sequenced. In addition, the
National Center for Biotechnology Information Entrez Genome Projects web site
reports that sequencing of several more plant genomes is in prog-ress.
Nevertheless, the completion of the entire genomic sequence of a par-ticular
organism represents the end of the structural genomics segment of the project. It is
clear, therefore, that the identification of every gene within the genomes of model
organisms is only the initial step to understand what these genes do and how they
interact to make up a living organism. Understand-ing the functions of the 20,000–
50,000 genes comprising plant (and animal) genomes and the variations within a
population and roles in normal devel-opment will represent a possibly greater task
than the mapping and sequenc-ing efforts currently underway.

Understanding the function of genes and other parts of the genome is known as
functional genomics, and research has involved model organisms such as A.
thaliana and rice. Model organisms offer a cost-effective way to follow the
inheritance of genes through many generations in a relatively short time.
Functional genomics is characterized by high throughput or large-scale
experimental methodologies combined with statistical and com-putational analysis
of the results. The fundamental strategy in a functional genomics approach is to
expand the scope of biological investigation from studying single genes or proteins
to studying all genes or proteins at once in
656 R.E. Litz et al.

a systematic fashion. Computational biology will perform a critical and expanding


role in this area. Whereas structural genomics has been character-ized by data
management, functional genomics will be characterized by min-ing the data sets for
particularly valuable information. Functional genomics promises to rapidly narrow
the gap between sequence and function and to yield new insights into the behaviour
of biological systems. The essential requirement for the implementation of
functional genomics is the availability of identified sequences, which can be
subsequently used in studies such as high-density arrays. Currently, there are very
few sequenced genes from mango, and these are mainly fruit-specific, and an effort
to increase this num-ber is needed. Considering the cDNA libraries prepared from
ripe fruit in various laboratories around the world, an effort to determine expressed
sequence tags (ESTs) would be worthwhile. At the same time, more cDNA
libraries from tissues other than fruit are needed. At present, the technology for
successful application of functional genomics to mango is well devel-oped, but the
‘raw material’ (i.e. identified genes) is lacking. Nevertheless, over 1000 ESTs from
ripe mango pulp have been sequenced and are in the process of being analysed by
microarrays using RNA probes from different ripening stages and from different
mango cultivars (G. Ramirez Zavala and M.A. Gómez-Lim, Irapuato, unpublished
results).

On the other hand, proteomics, the large-scale study of protein structure and
function, is still to be applied to mango. Proteomics is often considered the next
step after genomics but in fact it is more challenging, because a pro-teome differs
from cell to cell and constantly changes through its biochemical interactions with
the genome and the environment. The Human Genome Project has revealed that
there are far fewer protein-coding genes in the human genome than proteins in the
human proteome (20,000–25,000 genes versus >500,000 proteins). The protein
diversity is thought to be due to alter-native splicing and post-translational
modification of proteins and protein degradation (Reddy, 2007). Clearly, this
discrepancy implies that protein diversity cannot be fully characterized by gene
expression analysis alone. Thus proteomics may be more useful for characterizing
cells and tissues.

18.4 Genetic Engineering


In vitro induced mutations

There is significant consumer resistance to replacement of local mango cultivars by


newer selections in many traditional mango-producing countries of South and
South-east Asia. Serious production and postharvest problems that have a genetic
cause (e.g. alternate and irregular bearing, susceptibility to anthra-cnose and mango
malformation, tree shape and size, etc.) cannot be addressed in the long term using
applied physiology. For example, paclobutrazol has been applied to alternate-
bearing mango cultivars to promote flowering in the off-years; however, this can
result in severe decline in trees that have been treated for successive years. As a
result, conventional mango breeding in
Biotechnology 657

India has focused upon the development of mango cultivars that are largely
indistinguishable from traditional selections with respect to fruit size, appear-ance,
taste, flavour and overall quality.
Despite its impact on crop breeding (Maluszynski, 2001) and particularly on
banana improvement (Novak et al., 1990), mutation breeding has not been
successfully exploited for mango cultivar improvement by production of use-ful
off-types of existing selections. There is some anecdotal evidence that somatic
mutations can occur naturally in mango on the basis of variation that occurs within
seed-propagated polyembryonic cultivars. There are reported to be marked
phenotypic differences within polyembryonic ‘Kensington Pride’ trees in Australia,
and among polyembryonic cultivars of South-east Asia (e.g. ‘Arumanis’, ‘Golek’,
etc.). Different phenotypes of polyembryonic ‘Aru-manis’, for example, have been
characterized as ‘Arumanis-1’, ‘Arumanis-2’, etc. Gan et al. (1981) and Litz et al.
(1993) described isozyme variation within populations of South-east Asian
polyembryonic mangoes.
The most important production and postharvest problem of mango in the
humid tropics and subtropics is anthracnose, caused by Colletotrichum
gloeosporiodes (Penz.) Penz. and Sacc. In Penz. The current strategies for con-trol
of this disease involve the use of moderately resistant cultivars (i.e.
monoembryonic ‘Calypso’, monoembryonic ‘Keitt’ and monoembryonic ‘Tommy
Atkins’, etc.) and at least weekly applications of fungicides (i.e. benomyl or maneb
or mancozeb) from the time of flowering until harvesting (Dodd et al., 1997). This
can result in as many as 25 spray applications in a season, and is considered an
increasingly unsustainable agricultural practice.

The effect of γ-irradiation on embryogenic cultures of polyembryonic ‘Hindi’,


monoembryonic ‘Keitt’ and monoembryonic ‘Tommy Atkins’ was reported by Litz
(2001) as part of the Food and Agriculture Organization (FAO)/International
Atomic Energy Agency (IAEA) Co-ordinated Research Project entitled
Improvement of Tropical and Subtropical Fruit Trees through Induced Mutations
and Biotechnology. Embryogenic mango cultures on semi-solid maintenance
medium were exposed to 0–200 Gy irradiation provided by 60Co. The lethal dose
for 50% mortality (LD50) for each cultivar was deter-mined: it was approximately
125 Gy for ‘Keitt’ and approximately 100 Gy for ‘Tommy Atkins’. The LD 50 of
embryogenic ‘Hindi’ cultures could not be established within this dosage range,
possibly due to its high rate of prolif-eration relative to ‘Keitt’ and ‘Tommy
Atkins’. This study was confirmed by Manzanilla Ramirez et al. (2000). The main
objective of these studies has been to develop appropriate technologies for utilizing
induced mutations in vitro for recovery of useful off-types of existing cultivars.
Culture filtrates produced by pathogenic fungi and bacteria can be uti-lized not
only to select for resistance to the pathogen in vitro (Hammerschlag, 1992), but
also to induce the host resistance response. In order for in vitro selection to be used
effectively, essential prerequisites include: (i) a highly morphogenic suspension
culture; and (ii) evidence that the culture filtrate or phytotoxin can produce
symptoms of the disease on plant organs as well as with cells in suspension
cultures. Litz et al. (1991) first reported that the C.
658 R.E. Litz et al.

gloeosporiodes culture filtrate was effective as a selective agent in mango sus-


pension cultures growing in maintenance medium formulation. Somatic embryos
developed from embryogenic cells and PEMs that had survived exposure to C.
gloeosporiodes culture filtrate, and regenerants appeared to show resistance to
inoculation with the pathogen. Jayasankar et al. (1999) characterized the in vitro
effects of the purified C. gloeosporiodes phytotoxin, colletotrichin (Gohbara et al.,
1977, 1978), and crude C. gloeosporiodes culture filtrate on the mortality and
growth of polyembryonic ‘Hindi’ and polyem-bryonic ‘Carabao’ embryogenic
cultures. In this study, the LD50 values for the effects of C. gloeosporiodes culture
filtrate and colletotrichin on embryogenic cultures and the growth curves of
challenged cultures were established. Later, embryogenic cultures of the same
cultivars were either exposed con-tinuously for four cycles of
challenge/selection/regrowth or were challenged for one, two, three and four
complete cycles with colletotrichin and the par-tially purified culture filtrate of C.
gloeosporiodes (Jayasankar and Litz, 1998). At the end of each cycle, surviving
PEMs were physically removed from the selection medium, cloned and then either
rechallenged or subcultured onto somatic embryo maturation medium.

In order to induce the expression of anti-fungal genes in vitro, at least three


successive challenges with either crude filtrate or colletotrichin were necessary.
This was determined by co-culturing the challenged material with a virulent strain
of C. gloeosporiodes. Maintenance medium was inoculated with challenged and
selected embryogenic cultures at opposite sides of Petri dishes. After 3 weeks, a
virulent strain of the pathogen was inoculated in the centre of each Petri dish. Co-
culture of the pathogen with resistant cultures resulted in the suppression of
mycelium growth; the anti-fungal nature of the PEMs increased with each cycle of
challenge and selection, and was maxi-mum after three cycles. There was enhanced
production of extracellular anti-fungal proteins chitinase and E-1,3-glucanase in
selected cultures. An additional chitinase isozyme at 45 kDa was observed with
anti-fungal ‘Hindi’ cultures and at 25 kDa with anti-fungal ‘Carabao’ cultures, with
respect to the controls. The anti-fungal nature of selected, resistant lines in
suspension cultures and in somatic embryos was persistent for more than 2 years
follow-ing selection. Various random amplification of polymorphic DNA (RAPD)
markers were associated with selected cultures that were strongly anti-fun-gal
(Jayasankar et al., 1998). RAPD markers of the unchallenged controls and of
leaves from the parent trees were identical, indicating that exposure to either
colletotrichin or culture filtrate is essential for expression of anti-fungal genes. The
RAPD results also demonstrated that embryogenic cultures are quite stable
genetically, and the expression of anti-fungal genes is not the result of somaclonal
variation. Furthermore, it seems highly probable that the phytotoxins themselves
are highly mutagenic.

In vitro-induced mutation followed by selection could be a highly efficient


method for addressing specific breeding problems of mango assum-ing that
effective selection agents are available. Unfortunately, at this time, there are
relatively few such selection agents that can be utilized in this manner.
Biotechnology 659

Genetic transformation

Genetic transformation is currently the only practical solution for improving


existing elite selections of perennial species for specific horticultural traits and for
investigating gene function by interference RNA. Transformation of mango has
been reviewed most recently by Litz and Gomez-Lim (2002, 2005) and Gomez-
Lim and Litz (2007).

General protocols
Mathews et al. (1992, 1993) first reported the genetic transformation of mango
using embryogenic cultures of polyembryonic ‘Hindi’ and of a monoembry-onic
‘Keitt’ zygotic embryo-derived embryogenic line, respectively. These two studies
utilized two different disarmed, engineered strains of Agrobacterium tumefaciens:
(i) strain C58C1 containing the plasmid pGV 3850::1103 with the selectable
marker gene for neophosphate transferase (NPTII) which confers resistance to the
antibiotic kanamycin, both of which were driven by the CaMV constitutive 35S
promoter (Mathews et al., 1993); and (ii) strain A208 containing the plasmid
pTiT37-SE::pMON9749, a co-integrate vector, with genes for NPTII and the
scorable marker E-glucuronidase (gus or uidA) with the 35S promoter (Mathews et
al., 1992). A report by Cruz Hernandez et al. (1997) utilized A. tumefaciens strain
LBA4404 containing NPTII, E-glucuronidase (GUS) and genes that mediate a
horticulturally useful trait in binary plasmid pBI121 with the CaMV 35S promoter.
Mathews and Litz (1990) earlier had demonstrated that 12.5 Pg/ml kanamycin
sulfate is toxic to embryogenic suspension cultures; whereas, much higher levels
(200 Pg/ml kanamycin) are toxic to embryogenic cultures that are grown on semi-
solid medium.
These genetic transformation reports have followed a similar two-step
selection (Mathews et al., 1992; Cruz Hernandez et al., 1997). Embryogenic
suspension cultures in their logarithmic phase of growth are separated by passing
them through sterile filtration fabric (1000 Pm pore size), and the large fraction
(>1000 Pm) is abraded with a sterile brush on sterile filter paper. The abraded
PEMs are then incubated with acetosyringone-activated A. tumefaciens for 3 days
in liquid maintenance medium, with subculture into fresh medium at 24 h intervals.
The PEMs are then transferred onto semi-solid maintenance medium supplemented
with 200 mg/l kanamycin sulfate and 500 mg/l cefotaxime. After 10 months on this
selection medium, the PEMs are transferred to semi-solid maintenance medium
containing 400 mg/l kanamycin sulfate. Proliferating cultures are subcultured in
liq-uid maintenance medium containing 100 mg/l kanamycin sulfate, and somatic
embryo development is initiated by subculture onto semi-solid maturation medium.
Mathews et al. (1993) regenerated transgenic mango plants derived from a ‘Keitt’
zygotic embryo embryogenic culture and which had been transformed with pGV
3850::1103 containing the selectable marker gene nptII. Genetic transformation
was confirmed by: (i) growth in selection medium containing inhibitory levels of
kanamycin sulfate; (ii) positive histochemical reaction for GUS with X-GLUC
(Jefferson, 1987); and (iii) Southern hybridization.
660 R.E. Litz et al.

Transient gene expression in embryogenic polyembryonic ‘Kensington Pride’


and polyembryonic ‘Carabao’ cultures has been described using a biolistic
approach using two vectors: (i) pBI426 with GUS-NPTII under the control of a
double CaMV 35S promoter; and (ii) pBINgfp-Ser, which con-tains NPTII and the
green fluorescent protein gene (gfp) (Cruz Hernandez et al., 2000).

Transformation with genes that mediate horticulturally significant traits


Loss of mango fruit due to spoilage in storage and en route to markets accounts for
a significant proportion of total production in many developing countries that have
poorly developed infrastructure (i.e. cold storage facili-ties, poor roads, unreliable
transportation, etc.) (see Brecht and Yahia, Chap-ter 14, this volume) Mango has
become an important export commodity for several developing countries. Extended
shelf life and absence of physiologi-cal disorders that cause internal breakdown of
fruit (e.g. ‘soft nose’ and ‘jelly seed’) of the most important export cultivars (e.g.
monoembryonic ‘Tommy Atkins’) are potentially very important, therefore, for the
valuable export trade and for domestic markets.

The mango is a climacteric fruit, and ethylene therefore is a critical regu-lator


of the biochemical processes that occur during ripening. Certain rate-limiting genes
that mediate ethylene production in mango have been cloned. Cruz Hernandez et
al. (1997) described the genetic transformation of embryo-genic polyembryonic
‘Hindi’ mango cultures with mango ACC oxidase, ACC synthase and ACC
alternative oxidase cloned in the antisense orientation and under the control of the
CaMV 35S constitutive promoter in the pBI121 binary vector in A. tumefaciens
strain LBA4404. Embryogenic cultures were transformed by the two-step
procedure described above. Although the phe-notype of the transformed lines was
not reported, the genetic transforma-tions were confirmed in each case by the
XGLUC reaction for GUS, growth in the presence of inhibitory levels of
kanamycin sulfate, Southern blot hybrid-ization and NPTII amplification by PCR.
Successful regeneration of plants and inhibition of ethylene production by mature
mango fruit could possibly resolve the production problem of premature ripening
(jelly seed) and post-harvest loss due to spoilage.

18.5 In vitro Conservation

Medium-term storage

Mango embryos cannot tolerate desiccation during maturation, and devel-opment is


not arrested; this is typical of recalcitrant seeds or embryos. With-out a period of
developmental arrest, mango embryos develop to maturity and germinate in a
continuous sequence. Mango seeds (and embryos) cannot survive for more than 3
or 4 weeks under minimal in vitro growth conditions (Parisot, 1988), and would
perish after several days in a conventional seed bank.
Biotechnology 661

Monsalud et al. (1995) demonstrated that 4–5 mm long somatic embryos


representing the late heart stage can be partially desiccated, and stored dry in Petri
dishes for more than 30 days without any loss of viability. On the other hand,
larger somatic embryos cannot survive partial desiccation. This study has
interesting implications for future studies that could focus on the concept of an
‘artificial seed’ (i.e. somatic embryo) genebank for vegetatively propa-gated
tropical fruit trees.
Mango somatic embryos have been manipulated in order to induce
developmental arrest. Abscisic acid (ABA) is associated with the initiation of
developmental arrest and acquisition of desiccation tolerance of orthodox type
seeds and embryos. Abscisic acid causes developmental arrest in vitro at relatively
low concentrations for somatic embryos of the orthodox type (Bew-ley and Black,
1985). Pliego Alfaro et al. (1996a, b) were able to arrest the development of late
heart stage polyembryonic ‘Hindi’ somatic and nucellar embryos with ABA at 100
PM and higher concentrations in the maturation medium. This strategy arrests
growth and development for several months so long as mango embryos are on
ABA-containing maturation medium. ABA also has a persistent residual effect, and
mango somatic embryo develop-ment was inhibited for approximately 1 month
after their transfer onto matu-ration medium without ABA. Increased osmolarity of
the maturation medium also inhibited somatic embryo development; however, there
was no residual effect following transfer of somatic embryos onto maturation
medium with-out osmoticum.

Long-term storage

Embryogenic mango cultures cannot be stored indefinitely, and lose their


regeneration potential over time. The initiation of embryogenic cultures is
dependent on flowering and fruit set, which is strictly related to environmen-tal
stimuli, and normally occurs one time each year or on alternative years with
alternate-bearing selections. Long-term storage of embryogenic mango cul-tures is
essential for genetic manipulation studies and will be increasingly important for the
management of genetic resources. Embryogenic mango cul-tures have been
cryopreserved by different procedures (Wu et al., 2003; Rajani Nadgauda and
Pamela Moon, Homestead, Florida, USA, personal communi-cation). Wu et al.
(2003) compared three cryopreservation protocols for embryo-genic cultures
derived from monoembryonic ‘Zihua’ zygotic embryos: encapsulation-
dehydration, pregrowth-dehydration and vitrification. Encap-sulation-dehydration
was unsuccessful, and only limited survival (8.3%) was obtained following
desiccation of PEMs for 1 h to 58.5% moisture content prior to freezing in liquid
nitrogen. Vitrification, involving treatment of PEMs with plant vitrification
solution 2 (Sakai et al., 1991) for 20 min prior to freezing in liquid nitrogen, was
successful (94.3%). Embryogenic ‘Hindi’ cultures have also been introduced into
cryogenic storage (Rajani Nadgauda and Pamela Moon, Homestead, Florida, USA,
personal communication) and somatic embryos have been recovered from these
cultures. Two procedures were
662 R.E. Litz et al.

successful: (i) stepwise cooling in which cryoprotected (5% DMSO and 5%


glycerol) embryogenic cultures were cooled in ‘Mr Frosty’ containers at the rate of
–1C/min from room temperature (25C) to –75C followed by rapid cooling to –
196C; (ii) rapid cooling (vitrification). After cryovials were removed from liquid
nitrogen and rapidly warmed, cultures were thoroughly washed with maintenance
medium and plated on semi-solid maintenance medium. Somatic embryo
development was initiated by subculturing the PEMs on somatic embryo
maturation medium.

18.6 Conclusions
Biotechnology tools have great potential for mango, including advanced
micropropagation procedures, conservation and cultivar improvement. There are
also several strategies to genetically manipulate the crop for biotechno-logical
purposes. Genes in the antisense or sense orientation or RNAi (Small, 2007) can be
utilized to inhibit specific genes. The increasing availability of identified genes
should facilitate a better understanding and genetic manip-ulation of specific
developmental processes.
Considering the large number of mango microsatellite sequences identi-fied
and reported in the GenBank, parameters such as heterozygosity, aver-age gene
diversity, frequency of outcrossing, cultivar relationships and a mango genetic map
should be determined in the near future. Correlations between morphological and
DNA markers together with a linkage map should eventually enable MAS for
mango improvement.
Most of the research quoted here has been carried out in a few centres;
however, it is now widely acknowledged that biotechnology is mainstream
research. It is hoped that more research centres will be involved.

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constitu-ents of Brazilian varieties of mango fruit. Journal of Food Composition
and Analysis 13, 27–33.
Ansari, S.H., Ali, M., Velasco-Negueruela, A. and Pérez-Alonso, M.J. (1999) Volatile
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