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.
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).
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).
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
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.
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).
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.
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
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.’
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.
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.’
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
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Singh, L.B. and Singh, R.N. (1956) A Monograph on the Mangoes of UP.
Superintendent of Printing, Uttar Pradesh Government, Lucknow, India.
Sturrock, T.T. (1968) Genetics of mango polyembryony. Proceedings of the Florida
State Horticultural Society 81, 311–314.
Watson, B.J. and Winston, E.C. (1984) Plant genetic improvement. In: Proceedings
of the First Australian Mango Research Workshop. Commonwealth Scientific
and Indus-trial Research Organization (CSIRO), Canberra, pp. 104–138.
Wilson, C.W., Shaw, P.E. and Knight, R.J., Jr (1990) Importance of some lactones
and 2,5-dimethyl-4-hydroxy-3-(2H)-furanone to mango (Mangifera indica L.)
aroma. Journal of Agricultural Food Chemistry 38, 1556–1559.
Wolfe, H.S. (1962) The mango in Florida – 1887 to 1962. Proceedings of the Florida
State Horticultural Society 75, 387–391.
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.
Distribution
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
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).
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).
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
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).
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
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.
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.
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.
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.
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).
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.
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
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
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.
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.
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).
References
Angeles, D.E. (1991) Mangifera altissima Blanco. In: Verheij, E.W.M. and Coronel,
R.E. (eds) Plant Resources of South-east Asia No.2: Edible Fruits and Nuts.
Pudoc-DLO, Wageningen, the Netherlands, pp. 206–207.
Anonymous (1980) Mangifera L. In: Cheng, M. and Ming, T.L. (eds) Flora
Reipublicae Popularis Sinicae. Vol. 45(1). Science Press, Beijing, Peoples
Republic of China, pp. 73–78.
Aron, Y., Czosnek, H., Gazit, S. and Degani, C. (1998) Polyembryony in mango
(Mangifera indica L.) is controlled by a single dominant gene. HortScience 33,
1241–1242.
Audley-Charles, M.G., Hurley, A.M. and Smith, A.G. (1981) Continental movements
in the mesozoic and cenozoic. In: Whitmore, T.C. (ed.) Wallace’s Line and
Plate Tec-tonics. Clarendon Press, Oxford, UK, pp. 9–23.
Bellwood, P., Fox, J.J. and Tryon, D. (eds) (1995) The Austronesians, Historical and
Com-parative Perspectives. The Australian National University, Canberra.
Bompard, J.M. (1991a) Mangifera caesia Jack and Mangifera kemanga Blume,
Mangifera foetida Lour. and Mangifera pajang Kostermans. In: Verheij, E.W.M.
and Coronel, R.E. (eds) Plant Resources of South-east Asia No.2: Edible Fruits
and Nuts. Pudoc-DLO, Wageningen, the Netherlands, pp. 207–211.
Bompard, J.M. (1991b) Mangifera laurina Blume and Mangifera pentandra Hooker
f.; Mangifera odorata. In: Verheij, E.W.M. and Coronel, R.E. (eds) Plant
Resources of South-east Asia No.2: Edible Fruits and Nuts. Pudoc-DLO,
Wageningen, the Neth-erlands, pp. 216–220.
Bompard, J.M. (1993) The genus Mangifera re-discovered: the contribution of wild
spe-cies to mango cultivation. Acta Horticulturae 341, 69–77.
Bompard, J.M. (1995) Surveying Mangifera in the tropical rain forests of South-east Asia.
In: Guarino, L., Ramanatha Rao, V. and Reid, R. (eds) Collecting Plant Genetic
Diver-sity. Technical Guidelines. CAB International, Wallingford, UK, pp. 627–637.
Bondad, N.D., Rivera, F.N., Agcopra, D.B. and Minh Tam Aurin (1984) Philippine
man-goes and their relationship to South-east Asian cultivars. Philippine
Geographical Journal 28, 59–71.
Brandis, D.D. (1874) Forest Flora of North West and Central India. Allen, London,
pp. 125–127.
Brown, W.H. (1950) Useful Plants of the Philippines. Vol. 2. Bureau of Science,
Manila, the Philippines, pp. 336–340.
Campbell, R.J. (2004) Graft compatibility between Mangifera species and Mangifera
indica L. ‘turpentine’ rootstocks and their subsequent horticultural traits. Acta
Horti-culturae 645, 311–313.
Taxonomy and Systematics 39
Whitmore, T.C. (1981) Paleoclimate and vegetation history. In: Whitmore, T.C. (ed.)
Wal-lace’s Line and Plate Tectonics. Clarendon, Oxford, UK, pp. 36–42.
Yamanaka, N., Hasran, M., Xu, D.H., Tsunematsu, H., Idris, S. and Ban, T. (2006)
Ge-netic relationship and diversity of four Mangifera species revealed through
AFLP analysis. Genetic Resources and Crop Evolution 53, 949–954.
Yonemori, K., Honsho, C., Kanzaki, S., Eiadthong, W. and Sugiura, A. (2002)
Phyloge-netic relationships of Mangifera species revealed by ITS sequences of
nuclear ribo-somal DNA and a possibility of their hybrid origin. Plant
Systematics and Evolution 231, 59–75.
3 Important Mango Cultivars
and their Descriptors
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
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.
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.
‘Alfa’ (Brazil)
‘Alphonso’ (India)
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)
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).
‘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)
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).
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).
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)
‘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)
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).
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).
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).
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).
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.
‘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).
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
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)
‘Mabrouka’ (Egypt)
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).
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).
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).
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
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.
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).
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).
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)
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).
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).
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.
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
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Universidade Estadual de Jaboticabal, São Paulo, Brazil, pp. 35–50.
Schnell, R.J., Brown, J.S., Campbell, R.J., Kuhn, D.N., Meerow, A.W. and Olano,
C.T. (2006) Mango genetic diversity analysis and pedigree inferences for
Florida culti-vars using microsatellite markers. Journal of the American Society
for Horticultural Science 131, 214–224.
Singh, L.B. (1960) The Mango: Botany Cultivation and Utilization. Leonard Hill, London.
Singh, R.N., Majumder, P.K., Sharma, D.K. and Mukherkjee, S.K. (1972) Some promis-
ing mango hybrids. Acta Horticulturae 24, 117–119.
Valmayor, R. (1962) The Mango: its Botany and Production. University of the Philip-
pines, Laguna, the Philippines.
Whiley, A.W. (2001) Mango (Mangifera indica) ‘B74’. Plant Varieties Journal 14, 45–46.
Whiley, A.W. and Hofman, P.J. (2006) ‘Calypso’™ Best Practices Manual. (On CD).
Horticulture Ltd, Nambour, Australia.
4 Breeding and Genetics
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
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.
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
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.
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).
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.
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 7C and 25% relative humidity (RH), and
65.7% viability after 24 months of stor-age at 0C and 25% RH.
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.
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
Beak
Disease resistance
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.
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
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.
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.
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
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.
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.
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.
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.
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.
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
Polyembryony
1992, respectively) and 36 and 64% with ‘Madu’ and ‘Golek’, respectively
(Schnell and Knight, 1992).
India
‘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
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).
Induced mutations
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.
4.12 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
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.
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.
Lateral meristematic
primordia Apical dome
(includes meristem)
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.
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
3°
2°
º
4° 1°
5°
1° Bract
Axis
1°
Pedicel
1° Bract
1° 3° Bract
Pedicel
2° Bract
2°
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
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
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.
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
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.
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.
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).
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 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
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).
(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).
Temperature
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
15C day/10C night. In similar experiments with different cultivars with-out
previous clipping of distal leaves to stimulate initiation, inflorescences were
observed after 5 weeks at 15C day/10C 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 (18C day/10C night) for > 3
weeks and then transferred to a warm temperature (30C day/25C 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
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
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).
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
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
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
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
Gibberellins
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 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.
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.
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.
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
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
(15C) (Whiley et al., 1989). Other practices used with some success to enforce
canopy quiescence are girdling and the applica-tion of growth retardants.
INDUCTION INHIBITION
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
AUXIN
PHOTOASSIMILATES FRUIT
GIBBERELLINS A3
Ax
VEGETATIVE
GROWTH WATER STRESS
SHOOT INITIATION
PRUNING
DEFOLIATION CHILLING TEMP.
NITRATE SPRAY
ETHYLENE
ROOT INITIATION GIRDLING
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).
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.
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
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.
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.
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
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).
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.
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).
Pollination
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).
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
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
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
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.
5.16 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
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
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:
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)
0.20
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 30C 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 30C 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
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
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).
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).
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
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 ( 10C) 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.
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.
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.
Flooding
0.6
0.5
0.3
0.2
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).
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.
250
A Q = 400 μmol photons/m2/s
y = 84.5e–0.0313x
200 R2 = 0.69
J (μmol electrons/m2/s)
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
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 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)
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 20C
days/15C nights (20/15C) required 20 weeks (mean of ten cultivars) to complete
a growth/dormancy cycle while at 30/25C 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 30/25C, ‘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
20/15C an average of 7.1 leaves per flush were produced while at 30/25C there
were 13.6 leaves on each growth flush (data are mean values from ten cultivars). At
30/25C the mean leaf size was 300% greater than those on trees growing at
20/15C. 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 27C or 32C
for 120 days but an extended dormant period developed when soil temperatures
were held at 21C (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 15C (mean value for ten cul-tivars)
(Whiley et al., 1989). Subsequent studies (Issarakraisila et al., 1991) have
confirmed that 15C 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).
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.
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.
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).
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
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7 Fruit Diseases
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
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.
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).
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.
Resorcinol-5-(12-heptadecenyl) Resorcinol-5-(pentadecyl)
HO HO HO HO
(CH2)11 (CH2)14
CH CH2
CH CH3
(CH2)3
CH3
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
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).
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
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 50–55C
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 46C 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 < 10–13C, 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.
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).
Pathogenesis
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).
Management
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
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 13C 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).
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).
7.6 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
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Dodd, J.C., Prusky, D. and Jeffries, P. (1997) Fruit diseases. In: Litz, R.E. (ed.) The Mango:
Botany, Production and Uses. CAB International, Wallingford, UK, pp. 257–280.
Droby, S., Prusky, D., Jacoby, B. and Goldman, A. (1986) Presence of an antifungal
com-pound and its relation in the latency of Alternaria alternata in unripe peel of
mango fruits. Physiological and Molecular Plant Pathology 29, 173–183.
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228 D. Prusky et al.
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.
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).
Alternaria leafspot
Fig. 8.2. Conidia and conidiophores of Alternaria alternata (Source: Ellis, 1971).
Foliar, Floral and Soilborne Diseases 235
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
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).
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.
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.
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.
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 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
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
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.
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.
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
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
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.
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.
) ) )
(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.
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.
(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).
(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).
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
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
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).
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.
Fig. 8.16. Galls of the ‘buba’ type in Haiti (Photograph courtesy of Carolyn
Cohen, USDA, Animal and Plant Health Inspection Service (APHIS)).
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).
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
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).
Malformation
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).
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.
(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 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
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)
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).
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.
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
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
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
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 32C
(23C 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, 10–31C. 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
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.
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
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
(c)
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
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.
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).
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
Fig. 8.28. Conidia of Stigmina mangiferae, cause of stigmina leaf spot (Source:
from CMI description no. 097).
management is indicated rarely for these diseases; sanitation and other cultural
measures are used most often.
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 <
15C and > 35C 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
Phytophthora diseases
(a) (b)
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
the Spanish Type Culture Collection, CECT 20567, caused root rot on ‘Florida’
and lesions on leaves and stems of seedlings of ‘Gomera 3’.
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).
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
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.
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
(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).
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
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
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.
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
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 46C 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 55C (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.
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
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).
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.
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 13C 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.
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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.
Fruit flies
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 2C (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 15C and shortest (6 days) at 35C.
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.
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.
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
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.
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.
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’.
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).
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).
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).
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.
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.
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 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).
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 24C 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).
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).
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.
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.
Lepidoptera
The lepidopteran flower feeders are the second most important inflorescence pests
of mango. Geometrids, for example Chloropteryx glauciptera Hampson and
Pests 337
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 25C 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.
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.
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).
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 10–26C. 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.
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).
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.
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).
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).
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:
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.
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
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
Somatic
Approach Cleft or wedge Ground
embryogenesis
Veneer
Rind or crown
Budding
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).
Monoembryonic seed
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
Fig. 11.2. Grafting methods of mango propagation: (a) approach grafting; (b)
tongue grafting; (c) saddle grafting; (d) root grafting.
Crop Production: Propagation 371
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
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
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
(b) H-budding
Fig. 11.8. Top working (a) and double working (b) of mango.
Preparation of rootstock
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 15C 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
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 <15C, 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.
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.
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.
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 (25–35C) 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.
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).
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.
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
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
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).
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.
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
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
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
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.
References
Bhan, K.C., Samaddar, R.N. and Yadav, P.C. (1969) Chip budding and stone-
grafting of mangoes in India. Tropical Agriculture Trinidad 46, 247–253.
Bid, N.N. and Mukherjee, S.K. (1969) Varietal response to etiolation and growth-
regulator treatment in air-layering of mango (Mangifera indica L.). Indian
Journal of Agricul-tural Science 39, 1013–1019.
Bid, N.N. and Mukherjee, S.K. (1972) Studies into the effects of forced shoot
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12 Crop Production: Mineral Nutrition
I.S.E. Bally
Department of Primary Industries and Fisheries, Queensland, Australia
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).
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.
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).
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).
and internal breakdown in many cultivars (Tarmizi et al., 1993; Cracknell Torres et
al., 2004; see Galán Saúco, Chapter 9, this volume).
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.
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.
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).
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).
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 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
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).
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).
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).
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 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.
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
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).
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
12.13 Calcium
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.
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
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).
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 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.
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
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
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
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
Table 13.1. Regions of mango production in Brazil, Mexico, Taiwan and the USA
(Source: see table footnotes).
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 1433N
to 27N 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.
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
437
(Continued)
438
Table 13.2. Continued
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
Some cultivars, for example ‘Haden’ in north-eastern Brazil, have poor fruit set
when temperatures are >35C.
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 (17–27N) 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 26.5–28.3C). The Southern Pacific and Gulf of Mexico coastal regions have
isothermal (< 5C 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 22N (Fungshan, Ping-tung)
to 24N (Nanto and Taichung prefectures). The average temperature of these areas
is c.23C, with a mean of 20C during flowering (December– March) and c.18.6C
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 23C, 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 -6C) occur from December through to February
with a 70% probability of 0C 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–
29C is normal (Espenshade, 1992; SERCC, 2007). The lowest temperatures (18–
20C) 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 27C (Bahr and
Johnston, 1993b). Lower (13C) and higher temperatures (32C) 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 22.5C;
the average summer temperature is 31.8C, and the average winter temperature is
13.3C (Aslan et al., 1991; Garczynski, 1995). Winter temperatures range from
22.1C to 4.5C (Garczynski, 1995); however, Schacht (1992) reported a low of -
4.4C for 2 h. Summer tempera-ture extremes range from a low of 11C to 50C
(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).
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)):
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.
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.
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.
a b
Country Cultivar Origin Seed type Production season Tree growth habit
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.
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
Fig. 13.2. Tree training system for young trees to increase branching and productive
canopy volume (Source: after Oosthuyse, 1995; Campbell and Wasielewski, 2000).
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.
(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.
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.
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.
15 60 30 0
16–35 100 50 0
36–60 200 100 0
>60 280 140 0
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
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
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).
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.
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
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.
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.
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.
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.
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.
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.
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.
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 30–40C, depending on the cultivar. Mature fruits are stored at at 8–12C
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 46.5C
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
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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
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).
Table 14.1. Composition of the edible portion of mango fruit (Source: USDA/ARS,
2007).
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
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
16–21C was lower than that at 20–28C (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
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
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
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 20C; and (x)
ethylene production rate increases from 0.1–0.2 to 1–3 Pl/kg/h at 20C. 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
6 50
Hard green
5 Sprung green
40
Ethylene (mmol/kg/h)
Half ripe
CO2 (mmol/kg/h)
4 Ripe
30
3
20
2
Fig. 14.3. The climacteric pattern of respiration and ethylene production during mango
fruit ripening (Source: Lalel et al., 2003).
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 10C,
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.
Mangoes have a moderate ethylene production peak of 1–3 Pl/kg/h during ripening
at 20C. 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.
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.
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.
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
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)
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).
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
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
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)
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)
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
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).
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 10C results in a bland aroma and flavour (Bandyopadhyay
and Gholap, 1973b). ‘Totapuri’ mangoes,
506 J.K. Brecht and E.M. Yahia
Table 14.3. Characteristic aromas in ‘Alphonso’ and ‘Totapuri’ mangoes and their possible
chemical causes (Source: Bandyopadhyay, 1983).
mangoes, and at 8C compared to 12C 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).
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.10–13C
(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
7–8C for up to 25 days (Mann and Singh, 1976). While most cultivars show injury
at <10C 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 >30C 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).
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–
15C 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 20C. 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 12C (mature-green) and at either 8 or 12C (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 8C. Inhibition of
volatile production by 25 kPa CO2 was greater in mature-green than in tree-ripe
mangoes, and at 8C com-pared to 12C 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 12C (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 12C, while tree-ripe fruit can be stored for 21 days in the same
atmospheres at either 8 or 12C (Bender et al., 2000a).
The quality of ‘Keitt’ mangoes was evaluated during storage for 6 days at
20C 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 20C. 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 4C.
Both cultivars stored in MAP at 4C 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.
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 15C with 90–99%
RH, followed by simulated marketing conditions at 20C 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.
Insecticidal CA
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.
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15 Postharvest Technology
and Quarantine Treatments
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
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.
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.
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
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.
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
Fig. 15.1. Relation between flesh colour (1 = white; 15 = very yellow) and accumulated
heat units (>10C) 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).
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.
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 22C (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).
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).
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).
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).
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).
Timing
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
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
c
Control LOC Cl Magi Plus Wash
ear Glo
Mango Mango Mango Mango Man go Superconcentrate
Detergent
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.
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.
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.
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
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 48–50C for less mature
fruit and hot-water-damage-susceptible cultivars (e.g. ‘Zill’ and ‘Irwin’) and at 50–
55C 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 (52C). 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).
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 (55C 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-40C 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 46.5C 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 23C
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
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).
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
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
Required for
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.
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
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. 45C 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 43–47C pulp core temperature for 10 min to 6 h; however, the most common
treatment conditions are 46–47C for 10–30 min. Melon fruit fly (Bactrocera
cucurbitae Coquillett) immatures in mangoes from Okinawa were killed at 44
0.3C core temperature for 3 h (Sunagawa et al., 1987). Taiwanese mangoes
infested with melon fly can be disinfested with vapour heat at 47.5C until the
centre pulp is >46.5C for 45 min (Kuo et al., 1987).
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
46.5C for 30 min (AQIS, 2008a).
Hot air
Hot or forced hot air systems also heat the air to 40–50C, 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
>47C killed all stages of West Indian fruit fly, Anastrepha obliqua (Macquart), in
Mexican mangoes, and Sharp (1992) found a centre pulp temperature of >46C
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 46.1C 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 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).
Table 15.5. Minimum absorbed dose of gamma irradiation required by USDA for specific
pests (Source: adapted from EPA, 2002).
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 –17C and holding at –6C
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 20C 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 21–26C 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
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.
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
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.
Packaging
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
Palletizing
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
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.
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 20C 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 <18C 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 <24C 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 20C),
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 20C followed by exposure to 100 Pl/l ethylene for 24 h at 20C.
Fruit were then ripened at 20C. Means followed by the same letter in each column
are not significantly different (P >0.05) (Source: Hofman et al., 2001).
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 18–22C 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 20C (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).
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
● 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.
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.
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 2C 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).
● 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 0.5C 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.
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.
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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Postharvest Technology 605
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
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 (%)
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 (%)
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.
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$233,100,000 in 2006 (USDA, Foreign Agri-cultural
Service, 2007).
The main sources of USA imports of mango are Mexico, Peru, Ecuador and
Brazil. Mexico is the main supplier of mango to the USA (60.78% share in 2006)
(Fig. 16.2). In recent years, Brazil, Peru and Ecuador have become significant
exporters to the USA, competing with Mexico at the beginning and the end of the
season. The USA exports very few of its mango imports, mainly to Canada and the
UK.
612 E.A. Evans and O.J. Mendoza
350,000
300,000
250,000
USA total imports (t)
200,000
150,000
100,000
50,000
0
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year
Fig. 16.1. USA total imports of mango (t), 1997–2006 (Source: USDA
Foreign Agricultural Service, 2007).
200,000
180,000
160,000
140,000
USA total imports (t)
120,000
100,000
80,000
60,000
40,000
20,000
0
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year
Fig. 16.2. USA total imports of mango (t) by country, 1997–2006 (Source:
USDA Foreign Agricultural Service, 2007).
USA consumption of mango has increased steadily, from a per capita level of
0.5 kg in 1996 to 1 kg in 2005 (USDA, Economic Research Service, 2007). The
growth in USA consumption of mango is driven by many factors, such as year-
round availability, lower prices, consumer preferences and more
World Mango Trade and Economics 613
Table 16.4. Average cost, insurance and freight (CIF) prices for selected varieties from
main suppliers to the USA, 1998–2006 (US$/kg) (Source: USDA Agricultural Marketing
Service, 2007).
Country of origin
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.
Year 1 2 3 4 5 6 7+
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.
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.
Year 1 2 3 4 5 6 7+
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.
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.
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:
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.
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)
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
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
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 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).
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)
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$)
(Continued)
626 E.A. Evans and O.J. Mendoza
Price or Value or
cost per cost/ha
Unit Quantity/haunit (US$) (US$)
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
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.
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.
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).
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.
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.
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
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.
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.
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.
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).
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).
Evaporative rate
Table 17.4. Estimated volume of raw material needed for the operation of a mango
spray-drying and a mango dehydration plant.
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
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.
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.
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 25C with a 16 h photoperiod provided by cool white fluorescent lights
(50–70 Pmol/m2/s).
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
Country Reference
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 25C. 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.
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.
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.
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 25C 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):
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.
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 25C 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.
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.
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
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).
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
10C 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.
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.
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.
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.
Genetic transformation
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.
Medium-term storage
Long-term storage
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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