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Crop Protection 90 (2016) 132e141

Contents lists available at ScienceDirect

Crop Protection
journal homepage: www.elsevier.com/locate/cropro

Post-harvest development of anthracnose in pepper (Capsicum spp):


Etiology and management strategies
Asgar Ali a, *, Paa K. Bordoh a, Ajit Singh b, Yasmeen Siddiqui c, **, Samir Droby d
a
Centre of Excellence for Post-harvest Biotechnology (CEPB), School of Biosciences, The University of Nottingham Malaysia Campus, Jalan Broga, 43500,
Semenyih, Selangor D.E., Malaysia
b
School of Biosciences, The University of Nottingham Malaysia Campus, Jalan Broga, 43500, Semenyih, Selangor D.E, Malaysia
c
Laboratory of Food Crops and Floriculture, Institute of Tropical Agriculture, Universiti Putra Malaysia, 43400, Serdang, Selangor, Malaysia
d
Department of Post-harvest Science ARO, The Volcani Center, Bet Dagan, Israel

a r t i c l e i n f o a b s t r a c t

Article history: Chilli anthracnose, caused by Colletotrichum spp, is one of the main causes for post-harvest decay of chilli.
Received 27 January 2016 It can develop on the eld, during long distant transport, cold storage and shelf-life. In conventional
Received in revised form agriculture, the whole plant including the fruits, are sprayed with fungicides as a prerequisite for post-
15 July 2016
harvest control of chilli anthracnose. Due to consumer concerns regarding the use of synthetic fungicides
Accepted 21 July 2016
Available online 7 September 2016
and the demand for safer storage methods, the use of synthetic fungicides is no longer allowed for the
post-harvest control of chilli anthracnose. As a result, studies on alternative methods to control post-
harvest decay have been developed over the years along with the demand for safer storage methods.
Keywords:
Anthracnose
In this review, results published within the last decade have been summarized and alternative ap-
Chilli proaches to synthetic fungicides for post-harvest control of chilli anthracnose were discussed in detail.
Post-harvest management Overall, the use of natural antimicrobials, biocontrol agents, resistant cultivars and ozone shows promise
Natural products as treatments that can be adopted on a commercial scale to control post-harvest chilli anthracnose
Colletotrichum spp caused by Colletotrichum species.
2016 Published by Elsevier Ltd.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
1.1. Infection process of anthracnose and symptoms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
2. Control management practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
2.1. Conventional practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
2.1.1. Cultural practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
2.1.2. Chemical control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
2.2. Alternative management practice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
2.2.1. Resistant cultivars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
2.2.2. Natural products . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
2.2.3. Biological control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
2.2.4. Physical control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
2.2.5. Salts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
3. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Acknowledgement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

* Corresponding author.
** Corresponding author.
E-mail addresses: Asgar.Ali@nottingham.edu.my (A. Ali), khyx3pkh@nottingham.edu.my (P.K. Bordoh), Ajit.Singh@nottingham.edu.my (A. Singh), yasmeen@upm.edu.my
(Y. Siddiqui), samirdroby@volcani.agri.gov.il, samird@volcani.agri.gov.il (S. Droby).

http://dx.doi.org/10.1016/j.cropro.2016.07.026
0261-2194/ 2016 Published by Elsevier Ltd.
A. Ali et al. / Crop Protection 90 (2016) 132e141 133

1. Introduction C. gloeosporioides. C. capsici generally infects mature fruits whiles


C. gloeosporioides infects both green and mature fruits
Pepper belongs to the nightshade family (Solanaceae) and the (Pakdeevaraporn et al., 2005) (Table 1). However, C. acutatum has
genus Capsicum. It originated from the Americas, and was culti- specically been reported and identied as the cause of early
vated in New Zealand, South Africa, Malaysia and other Asian anthracnose disease in immature green bell peppers in Ohio,
countries. There are approximately 20e27 species of Capsicum; out Florida and Trinidad (Harp et al., 2008, 2013; Lewis-Ivey et al.,
of which ve are domesticated and used as fresh vegetables and 2004; Ramdial and Rampersad, 2014).
spices: Capsicum annum L., Capsicum frutescens L., Capsicum chinese As a result of the serious post-harvest losses caused by due to
Jacq., Capsicum pubescens Ruiz et Pavon. and Capsicum baccatum L chilli anthracnose, several attempts have been made to manage this
and grown worldwide (Than et al., 2008b). Within these ve problem. Traditionally, cultural techniques such as good sanitation
domesticated Capsicum species there are several cultivars. These practices (farm cleanliness); proper disposal of f rotten fruits, using
cultivars are of immense economic importance displaying different clean equipment and proper harvesting techniques have been
shapes, sizes, colors and avors. The most cultivated Capsicum va- employed to control post-harvest anthracnose. Additionally, syn-
rieties are Capsicum annuum (Tong and Bosland, 1999) followed by thetic fungicides such as propiconazole, difenoconazole, carben-
C. frutescens (Bosland and Votava, 2003). These cultivars of dazim, benomyl, maneb and captan (Gopinath et al., 2006;
Capsicum can be broadly categorized under Chilli (hot) or sweet Boonyapipat, 2013) have been used in the pre-harvest control of
peppers. The fruit of Capsicum has a variety of names, such as chilli, chilli anthracnose as a pre-requisite for the post-harvest control of
chilli pepper or pepper depending on differences between the the disease. It is imperative to note that benomyl (not registered or
English-speaking countries and type of fruits. Chilli pepper is hot to approved in Australia) and its associated fungicides carbendezim
mild in taste due to the presence of capsaicin whereas, sweet and thiophanate methyl (both of which registered) has raised major
pepper taste sweet as the name suggest and do not produce health concerns such as causing eye defects, and other birth related
capsaicin. Sweet peppers are native to Mexico, Central America, and effects by disrupting the process of cell division making their use
have been propagated throughout the world including tropics, unacceptable and dangerous (APVMA, 2009). Additionally, the
subtropics and temperate regions. emergence of resistant strains of C. capsici isolates in chilli fruit
Despite the rich nutritional and economic value of Capsicum spp, against benomyl, which were cross-resistant to thiophanate-
which has promoted their cultivation as a major cash crop, their methyl and carbendazim was reported in Malaysia (Sariah, 1989).
production has been greatly hindered by a variety of pests and Recently, resistance of C. truncatum to benomyl has also been re-
diseases. Amongst these, Colletotrichum is a large genus comprising ported in Trinidad (Ramdial and Rampersad, 2014). In addition,
a number of important species that are the most prevalent fungal long term usage of synthetic chemicals is known to have a negative
pathogens causing diseases in diverse tropical and subtropical impact on the environment especially soil and water resources
fruits and vegetables. Almost every crop including chilli grown (Wilson et al., 1999; Northover and Zhou, 2002). The increasing
throughout the world is susceptible to one or more species of health concerns expressed by consumers and the intention of
Colletotrichum. Recently, the genus Colletotrichum was listed as the governments to regulate pesticide use and their residues in fresh
eighth most important group of phyto-pathogenic fungi in the produce have necessitated the development of non-toxic alterna-
world, based on perceived scientic and economic signicance tive management techniques for anthracnose control. Therefore,
(Dean et al., 2012). this review highlights alternative approaches to chilli anthracnose
The prominence of Colletotrichum spp. and the losses caused by management during the post-harvest phase.
anthracnose post-harvest disease, presents a mounting threat to
commercially production of chilli. For instance, it anthracnose is
1.1. Infection process of anthracnose and symptoms
widely known to cause severe losses in yield as it affects immature
fruits in the eld and mature fruits after harvest, during storage and
Understanding the etiology of the disease is relevant for
transit when conditions are favorable (Thind and Jhooty, 1985;
developing strategies to control anthracnose. In most cases, post-
Hadden and Black, 1989; Jayalakshmi et al., 1999; Bosland and
harvest fungal infections occur when conidia from acervuli or
Votava, 2003). Moreover, in Thailand, yield losses of up to 50%
micro-sclerotia from the eld are splashed by irrigation or during
have been reported (Pakdeevaraporn et al., 2005) while in India,
rainfall on healthy fruit and leaves especially. Additionally, humid,
disease incidence levels ranged between 66% and 84% resulting in
warm and wet conditions aggravate the rate of infection since the
yield losses of 12e50% (Thind and Jhooty, 1985; Bagri et al., 2004;
fungus thrives best under such conditions. The splashed conidia
Sharma et al., 2005). In Korea, yield losses of 15% were reported
attach themselves to the fruits surface and begin to germinate
(Kim and Park, 1988) and 50% in Malaysia (Sariah, 1994). Apart from
producing appressoria. The appressoria develops a specialized
pre-harvest losses, fruit quality deterioration of chilli ranging be-
structure called infection pegs which then penetrate into the fruit
tween 21 and 47% due to chilli anthracnose have also been reported
epidermis. At this stage, the fungal infection may remain quiescent
in Sri Lanka (Anon, 1993).
as a biotroph. Activation of quiescent infections occurs during
Cross infection and pathogenicity studies have revealed that
ripening and senescence of the fruit after harvest (Prusky and
chilli anthracnose is caused by several species of Colletotrichum
Lichter, 2007); followed by colonization of fruit tissue, appear-
namely Colletotrichum acutatum (Simmonds), Colletotrichum capsici
ance of symptoms such as circular sunken spots (watersoaked) and
(Syd.) Butler and Bisby, Colletotrichum gloeosporioides (Penz) Penz.
production of acervuli and sporulation (Bailey et al., 1992; Prusky
and Sacc. and C. coccodes (Wallr.) S. Hughes (Simmonds, 1965;
et al., 2000). The known developmental stages where the fungus
Johnston and Jones, 1997; Voorrips et al., 2004; Pakdeevaraporn
might become inactive are during penetration, initiation of
et al., 2005; Sharma et al., 2005; Than et al., 2008a; Ramdial and
germination, germ tube elongation, formation of appressorium or
Rampersad, 2014). More recently Colletotrichum fruticola Prihas-
subsequent infection. Based on the denition of Swinburne and
tuti, L. Cai & K.D. Hyde and C. siamense Prihastuti, L. Cai & K.D. Hyde
Brown (1983), it was widely accepted that failure of spores to
have also been reported as causative agents for chilli anthracnose in
germinate or develop beyond any subsequent stage is due to
India (Sharma and Shenoy, 2014). However, the most economically
temporary adverse physiological conditions imposed by the host
important pathogens of chilli anthracnose are C. capsici and
directly on the fungus or indirectly by exhibiting temporary
134 A. Ali et al. / Crop Protection 90 (2016) 132e141

Table 1
Common Colletotrichum spp that causes post-harvest anthracnose in Chilli.

Causal agent Cultivar Symptoms References

C. capsici Bird pepper (C. frutescens), chilli (C. annuum), sweetbell  Diseased chilli peppers are withered with brown to black (Roberts et al., 2001;
pepper (C. annuum) necrotic patches Chanchaichaovivat et al.,
 Fruits develop water-soaked and small black circular 2007; Warin et al., 2009)
spots on the surface.
C. gloeosporioides Green and red chilli pepper (C. annum)  Lesion usually develops on ripe fruits. Lesions are small, (Kim et al., 1999; Kim et al.,
indenite, slightly sunken, water-soaked spots that may 2004)
enlarge rapidly and coalesce.
 Fruiting bodies form in concentric circles to cover the
surface of the lesions
 Lesions appear tan or brown and are covered with salmon
to orange gelatinous spores.

resistance against the fungus. Temperatures around 27  C and high infected plant debris from the eld (Nelson, 2008). Early planting of
humidity (a mean of 80%) are optimal for anthracnose disease chilli or planting cultivars that bear fruit within a short ripening
development (Roberts et al., 2001). Other reports also state that period to allow the fruit to escape fungal infection is also recom-
temperatures of 20  C and 25  C favor colony growth and sporu- mended. Other alternative sanitation practices such as weeding,
lation (Mello et al., 2004). Kim and Park (1988) determined that the removal of infected or wounded fruits should be carried out regu-
most important climatic parameter related to anthracnose devel- larly to prevent the pathogens from using such wounds as sites of
opment on chilli is relative humidity. Symptoms of chilli anthrac- infection.
nose are similar irrespective of the host species. Typically disease
symptoms on fruits are characterized by small, depressed water-
2.1.2. Chemical control
soaked lesions, which are sub-circular or angular with translucent
Synthetic chemicals have long been used as one of the most
light brown margins (Figs. 1 and 2). At advance stages of infection,
common practices for controlling chilli anthracnose. However, they
the circular or angular sunken lesions are covered with concentric
cannot be used in post-harvest management of anthracnose
rings of wet, gelatinous spores from salmon-colored fungal fruiting
because of health concerns to humans and potential buildup of
bodies (acervuli) due to the colour of the spores with numerous
pathogen-resistance to the fungicides. By ensuring proper cultural
black spines (setae) (Roberts et al., 2001).
practices without using synthetic fungicides, post-harvest
anthracnose can be reduced signicantly. Fungicide tolerance,
2. Control management practices however, often arises quickly, if a single compound is relied upon
too heavily (Staub, 1991). Planting seeds which are free of pathogen
This review reports on alternative post-harvest management infections or pre-treatment of seeds with synthetic chemicals
practices as opposed to the use of synthetic fungicides, which were (Thiram, Captan and Bavistin) are commonly used to control
clustered in ve categories: (i) resistant cultivars (ii) Natural anthracnose (Choudhary et al., 2013). Commonly used fungicides
products (antimicrobials) (iii) biological controls (iv) physical con- which have been traditionally recommended in pre-harvest chilli
trol (v) salts. It is fair to state that, most of these alternatives are still anthracnose management include Manganese ethyl-
far from practical application. enebisdithiocarbamate (maneb), propiconazole, difenoconazole

2.1. Conventional practices

2.1.1. Cultural practices


Several cultural practices have been used as pre-requisites to
control post-harvest chilli anthracnose due to the special etiology
of the pathogen. These precautionary measures are implemented to
minimize the rate of infection and reducing the infection pressure
even before fruits are mature and harvested. Than et al. (2008a,b) in
their review reported that, chilli seeds free of pathogen should be
planted and weeds eliminated. They also reported that rotating
crops that are not alternative hosts to Colletotrichum spp every 2e3
years is very effective for controlling post-harvest chilli anthrac-
nose. Other sanitation practices include the provision of good
drainage systems on the eld to channel out waste water or sewage
during on-farm fruit disinfection such as fruit washing at packing
houses or during irrigation regimes, and nally removal of plant
debris which may serve as source of inoculum (Than et al., 2008b).
If there was history of disease in a particular eld, then other crops
should be rotated in isolation from other solanaceous plant for at
least 2 years (Roberts et al., 2001). By doing so, the life cycle of the
pathogen on the eld to begin an infection process is disrupted and
the chance of infection is reduced since debris of most solanaceous
crop (after harvest) may become inoculum and a haven for the
fungus. At the end of the growing season, it is also recommended to Fig. 1. Anthracnose on green and red chilli pepper (C. annum cv. Nokwang') caused by
deep plough to completely cover diseases plants or removing C. gloeosporioides (source: Kim et al., 1999).
A. Ali et al. / Crop Protection 90 (2016) 132e141 135

They observed that fruits of PBC 595 had the largest lesions, while
fruits of PBC 518 had the smallest lesions (Manandhar et al., 1995).
Recently, Sun et al. (2015) reported on the inheritance of resis-
tance to C. acutatum from a C. chinense accession (PBC932) in a BC1
population derived from hybrid with C. annuum line 77013 (sus-
ceptible) using QTL analyses. Resistance tests were performed on
detached mature green or mature red fruit under laboratory con-
ditions (storage at 28  C, 95% RH for 48 h and additional 5 days
under plastic wrap) and then evaluated for disease incidence
(anthracnose) and overall lesion diameter. The mean disease scores
of the resistant parent PBC932 were 12.2% for GD (Disease inci-
dence in Green fruit stage), 0.8 mm for GO (Overall lesion diameter
in Green fruit stage), 2.10 mm for GT (True lesion diameter in Green
fruit stage), 52.5% for RD (Disease incidence in Red fruit stage),
4.0 mm for RO (Overall lesion diameter in Red fruit stage) and
5.8 mm for RT (True lesion diameter in Red fruit stage). These
values were all signicantly lower than those of the susceptible
parent 77013 (GD: 99.9%, GO: 21.19 mm, GT: 21.2 mm, RD: 97.8%,
RO: 19.6 mm and RT: 19.9 mm). The mean disease scores of the F1
individuals fell in between PBC932 and 77013, but skewed toward
PBC932 in all six phenotyping methods so that score values were
not signicantly different from those of the resistant parent. They
proposed that the resistance from C. chinense PBC932 is completely
Fig. 2. Anthracnose on green bell pepper fruit caused by C. capsici.
dominant over the susceptibility, at both the mature green and red
stages. Based on a map of the 14 linkage groups, that included 385
markers (SSR, InDel and CAPS), covering a length of 1310.2 cM,
and cabendazim (Smith, 2000; Gopinath et al., 2006). The efcacy
inclusive Composite Interval Mapping (ICIM) revealed main effect
of strobilurin fungicides, azoxystrobin (Quadris and Amistar 25
QTLs were located on the P5 chromosome for all fruit stages and
SC), trioxystrobin (Flint), pyraclostrobin (Cabrio) have also been
resistance criteria, and four minor-effect QTLs were detected only at
labeled to control chilli anthracnose but only preliminary reports
the green mature stage. Identication of recombinant individuals
exist on the efcacy of these fungicides against the disease in its
suggested that resistance in green versus red fruit may be
severe form in eld trials (Alexander and Waldenmaier, 2002;
controlled by distinct genes within the QTL interval on P5, Sun et al.
Lewis and Miller, 2003). Despite the positive results obtained in
(2015). Further studies need to be undertaken to investigate the
pre-harvest studies in chilli anthracnose managements using these
importance of these distinct genes in the post-harvest manage-
synthetic chemicals, some Colletotrichum spp like Colletotrichum
ment of chilli anthracnose.
capsici, C. gloeosporioides and C. siamese have developed resistance
to thiophanate-methyl, Strobilurin-fungicides (azoxystrobin and
2.2.2. Natural products
kresoxim-methyl) and carbendazim (Sariah, 1989; Inada et al.,
2.2.2.1. Plant extracts. Certain antimicrobial or metabolic com-
2008; Hu et al., 2015).
pounds synthesized by plants might be very good alternative for
controlling diseases of tropical fruits and vegetables. In chilli,
2.2. Alternative management practice several in vitro studies have shown the efcacy of certain medicinal
herbs or plant extracts against Colletotrichum spp. (Charigkapakorn,
2.2.1. Resistant cultivars 2000; Begum et al., 2007; Nduagu et al., 2008; Johnny et al., 2011;
Research has shown that, the usage of resistant varieties not Saravanakumar et al., 2011; Ajith et al., 2012) (Table 2). A lot of
only reduces damages from diseases, but also minimizes chemical studies have been conducted on the pre-harvest control of chilli
and mechanical expenditures of disease control (Grichar et al., anthracnose (Charigkapakorn, 2000; Ajith et al., 2012; Rashid et al.,
1998; Besler et al., 2001; Monfort et al., 2004). However, the 2015) but very few studies have reported as post-harvest control.
reduction in some cases maybe complete or partial. Resistant cul- Plant parts such as rhizomes, bark, roots and leaves have shown
tivars are mostly used as a control measures to manage pre-harvest signicant antimicrobial properties against Colletotrichum sp that
anthracnose in chilli fruit or plants in the eld. Some genetic re- causes chilli anthracnose (Table 2).
sources resistant to anthracnose in chilli have been independently Research has shown that an aqueous leaf extract of Plumbago
reported from different countries and regions (Kim et al., 1987; Park indica L. had a signicant suppressive effect on C. gloeosporioides in
et al., 1987; Hong and Hwang, 1998; Pae et al., 1998; AVRDC, 1999; terms of conidial germination (98.9%) and radial mycelial growth
Yoon and Park, 2001; Yoon, 2003; Mongkolporn et al., 2004; Kim (98.8%). In vitro studies have shown that botanicals or plant ex-
et al., 2004, 2007; Voorrips et al., 2004; Lee et al., 2010; tracts from Cathranthus roseus L. G.Don., Coleus aromaticus Benth.,
Shivashankar et al., 2010; Kim et al., 2011; Rahman et al., 2011; Manilkara zapota L. P.Royen., and Azidirachta indica A. Juss confers
Syukur et al., 2013; Sun et al., 2015). antifungal effects on the radial mycelial growth of C. capsici
Few studies about the use of resistant cultivars to control post- responsible for anthracnose disease in bell pepper (Capsicum fru-
harvest rot in pepper have been reported. Earlier research in the tescence) (Ajith et al., 2012). Singh and Khirbat (2014), reported on
Philippines reported that C. annuum of different lines show resis- the efcacy of aqueous extract of three wild plants viz., siras
tance to anthracnose caused by C. gloeosporioides isolated from (Albizza lebbeck L. Benth.), babul (Acacia Arabica Lam.) and Cler-
C. annuum. Over 50.0% of the fruits in lines PBC 452, PBC 454 and odendron (Clerodendron infortunatum L.) to control fruit rot in
PBC 595 had lesions in less than 5 days after inoculation, whereas it C. annuum caused by C. capsici. The leaf extracts of siras, babul
took 6 days for fruit in three lines (PBC 365, PBC 371 and PBC 518), 8 and clerodendron caused signicant inhibition of spore germina-
days for fruit in line PBC 370, and 11 days for fruit in line PBC 495. tion in C. capsici with percentages of inhibition of 100%, 86.6% and
136 A. Ali et al. / Crop Protection 90 (2016) 132e141

73.2% respectively. peel colour and soluble solids concentrations. More recently, Ali
Recently, Alves et al., 2015 reported the efcacy of 1% aqueous or et al. (2014b) assessed the efcacy of only ethanol extracted
20% ethanol plant extracts to control bell pepper (C. annuum) propolis (EEP) on C. capsici isolated from matured green bell pepper
anthracnose caused by C. acutatum after fruits were treated with (C. annuum L.). It was observed that mycelial growth inhibition was
plant extracts before inoculation with C. acutatum conidial sus- positively related to the EEP concentration with inhibition values of
pension of C. acutatum (2  105 conidia/ml). Fruits were kept at 66.1%, 84.3% and 89% at dilutions of 0.25%, 0.50% and 0.75%
high RH (90%) for 24 h in moist chambers made from plastic trays respectively. Furthermore, when 1%, 5% or 10% concentrations of
containing four layers of paper towels moistened with sterile EEP were used on fruits, they signicantly delayed anthracnose
distilled water and placed in polyethylene bags. In this study, 6% development for 28 days on articially inoculated bell peppers
aqueous garlic, mallow and ginger extracts reduced disease severity stored at 10  C and 90% relative humidity (Ali et al., 2014b).
by more than 97%. Garlic extract required the highest concentration Recently, 0.5% of lemongrass essential oil incorporated with a 1.0%
to reduce severity by 75% when compared to ginger and mallow chitosan when applied as edible coating, signicantly maintained
extracts, but exhibited the best efcacy in relation to different fruit quality on green bell pepper (C. annuum), with the anthrac-
inoculum concentrations and isolates of C. acutatum. They nose disease incidence results in consideration after 21 days at
concluded that the garlic aqueous extract effectively controlled room temperature. However chitosan alone was more effective in
anthracnose caused by C. acutatum in bell pepper under controlled extending the fruit shelf life (Ali et al., 2015).
conditions. It has been proposed that, because anthracnose is a latent
Even though recent research suggests the use of these plant infection and starts from the germination of conidia, which pro-
extracts as bio-fungicides (Nduagu et al., 2008; Johnny et al., 2011; duce appressoria, cinnamon oil which contain cinnamaldehyde
Ajith et al., 2012; Pranshanth et al., 2014), more studies on their penetrates into the appressoria of fungus to inhibit its growth
efcacy in the control of post-harvest chilli anthracnose need to be thereby slowing down the anthracnose (Maqbool et al., 2010). Citral
conducted. aldehyde or citral present in lemongrass oil is attributed to cause
irreversible cell membrane disruption through cross-linkage reac-
tion leading to leakage of electrolytes and subsequent depletion of
2.2.2.2. Essential oils and propolis. Bautista-Ban ~ os et al., 2013, in a
amino acids and sugars (Inouye et al. (2000), whiles others may
review stated that essentials oils are basically concentrated hy-
selectively be inserted into the lipid rich portion of the cell mem-
drophobic liquid containing a mixture of volatile aroma com-
brane, thereby disturbing membrane function (Regnier et al., 2008).
pounds resulting from plant secondary metabolism and have long
Also the presence of monoterperne hydrocarbons such as limonene
been known to provide effective control over phytopathogens. They
and camphone are reported to exert fungitoxicity by diffusing into
are usually obtained by steam or hydro-distillation and are known
cell membrane and causing deformation (Meincken et al., 2005;
for their antiseptic (bactericidal, viricidal and Fungicidal) properties
Tyagi and Malik, 2010).
and the rich fragrance they produce (Bakkali et al., 2008). Although
several in vitro studies have suggested that the use of essential oil
will control Colletotrichum spp., in different fruits and vegetables, 2.2.2.3. Chitosan and edible coating materials. Chitosan a natural
very few studies have been conducted to control post-harvest chilli compound (linear polysaccharide) composed of randomly distrib-
anthracnose. Hurdle technology which involves the combination of uted b-(1-4)-linked D-glucosamine obtained from chitin in the
two or more natural products as edible coating on fruits has been outer shell of crustaceans had widely been used as an antifungal
used in postharvest control of anthracnose. A study by Ali et al. agent to control anthracnose on several tropical and subtropical
(2014a) revealed that cinnamon oil (0.1%) in combination with fruits and vegetables (Bautista-Ban ~ os et al., 2006, 2013; Ediringhe
Brazilian green propolis (5%) was fungicidal (100% inhibition) on et al., 2012). In chilli, chitosan (0.8%) was as effective as a seed
the mycelial growth and conidial germination of C. capsici. When treatment in reducing Colletotrichum spp infection and improving
these combinations were used on chilli fruits (C. annuum), they seedling growth of chilli cv. Jinda (Photchanachai et al., 2006),
lowered the post-harvest disease incidence by 14% after 33 days of since the fungus is seed-borne and infected seeds had reduce
storage, and signicantly delayed changes in weight loss, rmness, germination and their seedlings had poor vigor (Maude, 1996).

Table 2
Efcacy of different plant extracts to control Colletotrichum capsici (in vitro) and post-harvest chilli anthracnose (in vivo).

Causal agent Bio-fungicide (dose) Level of control Disease Reference

C. capsici Aqueous extract of stem bark of Azadirachta indica A. Juss Reduction of colony by 40.8 In vitro studies (Nduagu et al., 2008)
(5% w/v) and Vemonia amygdalina Del. (5% w/v) and 22.5% respectively
Aqueous extract of root bark of Azadirachta indica A. Juss (5% 39.4% and 54.2% respectively
w/v) and Vemonia amygdalina Del. (5% w/v)
C. capsici Aqueous leaf extracts of Abrus precatorius L. (Gundumuthu) 66.5% disease reduction Fruit rot (Anand and Bhaskaran, 2009)
(10%).
Aqueous leaf extracts of Aegle marmelos L. (vilvum) (10%). 58.3% disease reduction
Pseudomonas uorescens (2%)-as a biocontrol agent. 53.4% disease reduction
C. capsici Methanol crude extract of Piper betle L. (10 mg/ml and 85.3% and 100% inhibition of radial In vitro Studies (Johnny et al., 2011)
12.5 mg/ml) mycelia growth respectively.
Chloroform leaf extract of Piper betle L.(10 mg/ml and 78.5% and 100% inhibition of radial
17.50 mg/ml) mycelia growth respectively
Acetone leaf extract of Piper betle L. (10 mg/ml and 15.00 mg/ 73.6% and 100% inhibition of radial
ml). mycelia growth respectively
C. capsici Methanol leaf extract of Maesa indica Roxb. A.DC (1 mg/ml). 74.2% inhibition of mycelia growth In vitro studies (Kambar et al., 2014)
Methanol leaf extract of Pimenta dioica L. Merill (1 mg/ml). 71.0% inhibition of mycelia growth
Methanol extract of Persea macrantha Nees Kosterm. (1 mg/
ml).
C. capsici  Methanol leaf extract of Citrus reticula Blanco. (1 mg/ml).  Over 50% inhibition of fungal In vitro studies (Pranshanth et al., 2014)
growth
A. Ali et al. / Crop Protection 90 (2016) 132e141 137

Recently, chitosan degraded by gamma irradiation to obtain irra- the consideration of these antagonists as bio-control agents of chilli
diated oligochitosan at 0.01e1% signicantly inhibited the mycelial anthracnose.
growth of C. gloeosporioides isolated from chilli and the effect was In Thailand, it was reported that Pichia guilliermondii Wick
shown to be dose dependent (Kewsuwan et al., 2014). signicantly controlled chilli anthracnose caused by C. capsici dur-
In post-harvest studies, chitosan (1.0% w/v) treated bell pepper ing storage at 28  C and 95% relative humidity in the dark
(C.annuum) fruits stored at 13  C or 20  C (RH 85%) markedly (Nantawanit et al., 2010). In this study, fruits were inoculated with
reduced the weight loss in bell peppers at both temperatures. yeast suspension (30 ml at 5  108 cells/ml) and incubated for
However, increasing the concentration of chitosan from 1.0 to 1.5% various time intervals (until 72 h). For control only sterilized water
(w/v) provided signicantly greater weight retention, while was used. A second wound was made approximately 1 cm away
reducing respiration rate, colour loss, wilting and reduction in from the initial wound and inoculated with 10 ml of a spore sus-
C. gloeosporioides symptoms (EL Ghaouth et al., 1991). Another pension of C. capsici (5  104 spores/ml). Fruits were air-dried and
hurdle technology which include chitosan coating enriched with put in plastic trays and wrapped with polyethylene sleeves under
cinnamon oil (0.25%) on sweet pepper stored at 8  C for 35 days 95% relative humidity for 5 days at 28  C in the dark. The incidence
provided the best control effect on anthracnose (below 5%) (Xing of disease (the percentage of infected wounds that exhibited
et al., 2011). On fresh-cut green peppers (Capsicum annuum), chi- anthracnose symptoms) and the lesion diameter (diameter of
tosan edible coatings at different concentrations (0.5%, 1.0% and rotted tissue) were determined after 5 days of incubation. Lesion
2.0%) resulted in a dose dependent effect with respect to reduction diameter was measured as the average length of the lesion along
in anthracnose symptoms in comparison to the control after stor- two perpendicular axes. To study the development of resistance at
age at 5  C for 15 days. Xing et al. (2011) did not specify the causal different distances from the yeast-treated wounds, similar experi-
agent or the type of decay as the disease incidence was reported as ments at 24 h after yeast treatment were performed. Afterward, the
decay (Raymond et al., 2012). Edirisinghe et al. (2012) reported that fungal-treated wounds were made at 1, 2, and 4 cm away from the
chitosan (1.5%) decreased anthracnose severity in green bell pepper initial yeast-treated sites and all treated fruits were incubated for 5
(Capsicum annuum cv. Meno) caused by C. capsici by approxi- days at 28  C and 95% Relative Humidity (RH) in the dark. The
mately 76% after 28 days of storage. It has been proposed that the incidence of disease and the lesion diameter were also determined
chitosan coating act as a barrier, limiting the penetration of the after incubation for 5 days. In that bio-control system, the antag-
fungus and stimulates structural defense reactions such as the onist P. guilliermondii induced defense response in C. annuum.
formation of hemispherical protuberances along host cell walls, cell Chanchaichaovivat et al. (2007), reported a signicant effect of
wall thickening and occlusion of many intercellular spaces with a Pichia guilliermondii Wick strain R13 (yeast isolated from Thai
brillar material in bell pepper (EL Ghaouth et al., 1997), or acts as a rambutan (Nephelium Lappaceum L.) in reducing the disease inci-
stimulus to induce host defense responses such as activation of dence on C. capsici infected chilli fruit of bird pepper (C. frutescens),
defense related enzymes (Bautista-Baos et al., 2006). chilli (C. annuum) and sweet bell pepper (C. annuum) to as low as
Other possible mechanism of antifungal action of chitosan in- 6.5%. In this experiment, wounded surface-sterilized chilli fruits
cludes interaction between positively charged chitosan molecules were inoculated with 20 ml of cell suspension of yeast (5  109 cells/
due to ammonium groups and negatively charged residues on the ml) and allowed to dry for 1e2 h. About 20 ml of spore suspension of
fungal cell wall surface due to phospholipids head groups, which C. capsici (5  104 spores/ml) was added to the wound. Fruits were
lead to increase in plasma membrane permeability (Leuba and put on plastic trays wrapped with polyethylene sleeves and stored
Stossel, 1986; Kaus et al., 1989). It is also reported that chitosan in a ventilated cabinet in the dark at 28  C and 95%RH. The authors
may enter the fungal cell, interact with DNA, and alter its confor- suggested the use of Pichia guilliermondii as biological antagonist as
mation, thus inhibiting the synthesis of mRNA and proteins preservative to control anthracnose as oppose to the use of chlo-
(Hadwiger and Loschke, 1981). rinated water. It has been proposed that Pichia guilliermondii strain
R13 and other yeasts suppresses Colletotrichum spp through mul-
2.2.3. Biological control tiple modes of action including nutrient competition, their tight
In post-harvest control of chilli anthracnose, it has been re- attachment to host plant resulting in competition for space be-
ported that, antifungal metabolites secreted from Trichoderma tween biological antagonist and the pathogen fungus, production
harzianum Rifai with strain number T-156co5 signicantly of lethal toxin (termed killer toxin), possibly induction of plant
controlled C. capsici isolated from C. annuum L. The puried anti- resistance and production of hydrolytic enzyme (Janisiewicz et al.,
fungal metabolite determined by 1H and 13C Nuclear Magnetic 2000; Chanchaichaovivat et al., 2008; Nantawanit et al., 2010).
Resources spectroscopy was closely related to isoharziadione and When chilli fruits were pretreated with the yeast antagonist, they
was tested on C. annuum at concentrations of 50 and 100 mg/L had signicantly reduced disease incidences and lesion diameters
(Warin et al., 2009). When this treatment was tested on detached caused by C. capsici. Additionally, yeast-treated fruits signicantly
fruits kept in plastic boxes and incubated for 4 days in a plant enhanced the activities of phenylalanine ammonia-lysae, chitinase
growth cabinet 25 2  C, 12 h light and 80% RH), the antifungal and b-1, 3-glucanase and the accumulation of capsidiol spores
metabolite secreted from Trichoderma harzianum T-156co5 at phytoalexins in chilli tissue (Nantawanit et al., 2010). Some studies
100 mg/L was also comparable to 100 mg/L benomyl in controlling have also shown that these antagonists secretes an antifungal
anthracnose and produced the highest disease control efcacy metabolite (isoharziandione) which inhibits spore germination and
score of 89.9%. germ tube growth of certain plant pathogens (Warin et al., 2009).
In Ethiopia, a laboratory control test revealed the efcacy of
Trichoderma species against C. gloeosporioides isolated from 2.2.4. Physical control
C. annuum (Jebessa and Ranamukhaarachchi, 2006). In that study, a The use of physical control methods such as blanching, hot air,
test antagonist Trichoderma harzianum Rifai (mutant, Th38), Tri- vapor or hot water rinsing and brushing alone or in combination
choderma pseudokoningii Rifai and Trichoderma koningii Oudem with other treatments has been reported to control post-harvest
separately inhibited the growth of four isolates of C. gloeosporioides decay in chilli and bell peppers. Treatments of bell pepper
when a mycelia disk of the test antagonist was placed at the edge of (C. annuum) with hot water at 53  C for 4 min or at 45  C for 15 min
petri plate containing potato dextrose amended with 15-day-old was effective in alleviating chilling injury and reducing decay after
C. gloeosporioides spore suspension (106/ml). The authors proposed 14 and 28 days of storage at 8  C (Gonzalez-Aguilar et al., 1998;
138 A. Ali et al. / Crop Protection 90 (2016) 132e141

2000). The researchers conrmed that hot water treatments (4 min Earlier studies revealed that sodium benzoate which, under
dip at 53  C) followed by packaging the bell peppers with a acidic conditions, converts to undissociated benzoic acid and
0.065 mm low density polyethylene lm signicantly reduced the become responsible for its antimicrobial activity (Concise
respiration rate, reduced decay, retained turgidity and green International Chemical Assessment, 2004). Studies have showed
coloration maintained excellent overall quality after 28 days of that sodium metabisulphite are being used as food additive having
storage at 8  C. A rapid method for simultaneously rinsing and antimicrobial function. Several studies about the inhibitory effect of
disinfecting sweet pepper (C. annuum) using hot water and brushes sulphites on yeast, molds and bacteria has also been reported
at 55 1  C for 12 2s reduced decay incidence while maintaining (Lindsay, 1985; Russell and Gould, 1991). The proposed mecha-
fruit rmness and general appearance of the fruit after storage at nisms of sulphites include reactions with protein carbonyl groups,
7  C for 15 days and an additional 4 days at 16e18  C was (Fallik FAD, RNA and DNA (Russell and Gould, 1991). In the case of po-
et al., 1999). In the disinfecting process, physical treatments such tassium metabisulphite against microorganisms, it converts itself
as hot water rinsing and brushing helps to remove dirt, dust and into sulphur dioxide in aqueous solution which acts as an active
even fungal spores from the fruit calyx and skin and also help to antimicrobial compound. The mechanisms include the reduction of
seal the small cracks in the epidermis of fruit, thus maintaining disulphide linkages, formation of carbonyl compounds, reaction
better keeping quality (Fallik et al., 1999). However, short hot water with ketone groups, and inhibition of respiratory mechanisms
treatments might not control latent infections of Colletotrichum (Frazier and Westhoff, 1988; Rengasamy and Poole, 1995).
spp. (Fallik et al., 1999), and its application alone cannot provide or
maintained the post-harvest qualities of pepper (Gonzalez-Aguilar 3. Conclusion
et al., 1998).
Possible disease resistance mechanisms induced by heat treat- As a tropical commodity, Capsicum spp, including both bell
ment during postharvest storage of fruits have been elucidated pepper and chilli can be preserved and the shelf life extended for up
(Yun et al., 2013). It is reported that resistance mechanism is as a to 71 and 38 days at 10  C and 7.5  C respectively, without spoilage
result of associated up-regulated proteins such as beta-1, 3- or any off-avors when proper treatments are applied. According to
glucanase, Class III chitinase, 17.7 kDa heat shock protein and low the scientic literature, high losses are due to various microor-
molecular weight heat-shock protein in fruits tissues which are ganisms, with anthracnose being one of the major problems asso-
expressed and act as defensive mechanism for host tissue to ght ciated with the marketing of high-quality fruits in many Capsicum
against the pathogen when heat treatment is applied (Yun et al., production areas. Traditionally and in most countries, application of
2013). fungicides is the most common method for controlling chilli
Despite the lack of research on the use of ultraviolet-C (UV-C) to anthracnose on the fruit and other part of the plants. Commonly
control chilli anthracnose, it has been reported to signicantly used synthetic chemicals include broad spectrum contact fungi-
reduce post-harvest fruit rot or decay. Exposure of fresh-cut mature cides such as captan and some cases carbendazim. Benomyl which
green pepper at 20 KJ/m2 and stored at 5  C for 5 or 12 days showed was used before to control anthracnose has been discouraged due
reduction in counts of mesophyll bacteria and molds, while main- to fungal resistance and their negative impact on environment and
taining the keeping quality of the fruit (Rodoni et al., 2012). Fresh- human health. However, in some countries its registration has not
cut bell pepper (C. annuum), irradiated with UV-C at 10 kJ/m2 on the been abrogated and it is still produced. It is suggested that its use or
inner and outer surface of the fruit has also been reported to production in such countries should be discouraged due to health
effectively reduced spoilage (Rodoni et al., 2015) after storage at concerns, for instance the treatment of peppers with benomyl
4  C for 12 days. As a simple and environmental-friendly strategy to might have a negative effect on a producer's ability to export the
extend the post-harvest life of fresh vegetables (Civello et al., 2006), peppers.
UV-C radiation induces benecial physiological effects in plant At the post-harvest, biological control, natural products (such as
tissues, delay ripening and senescence-associated changes (Bu chitosan, essential oils and plant extracts), resistant cultivars and
et al., 2013), activates defense-related genes (Pombo et al., 2011) some salts have been used to control anthracnose. However, more
and induces the accumulation of phytoalexins (D'hallewin et al., research needs to be conducted to nd better alternatives to control
1999). For bell pepper and chilli, there are no of any on physical chilli anthracnose, despite already existing practices where syn-
methods to control post-harvest chilli anthracnose. thetic fungicides are used in the eld as a pre-requisite to post-
In a recent post-harvest study, the fumigation of bell pepper harvest disease management of chilli anthracnose. One non-
(C. annuum cv. Zamboni) with 3 ppm ozone gas at 12  C and 95% traditional alternative that is still under experimentation is
relative humidity for 3 days, followed by 24 days of cold storage at Ozone. Despite successful results achieved in minimally processed
12  C demonstrated great potential for increasing ascorbic acid fruits, more research should be carried out to better understand the
content of the fruit, reducing anthracnose disease incidence and mode of action of ozone on Colletotrichum to control chilli
maintaining fruit physic-chemical quality in comparison to fruits anthracnose since it leaves no residual effects. On the other hand,
exposed to 0, 1, 5, 7 and 9 ppm ozone (Alwi and Ali, 2014). the high start-up costs associated with in Ozone technology makes
it likely that only export producers will consider this technology.
2.2.5. Salts The use of non-toxic, natural or biodegradable products to
Salts such as sodium and potassium caused signicant re- control post-harvest diseases is a credible choice, and hence, lab-
ductions in conidial germination, and mycelial growth when so- oratory research has been aimed at identifying safe products in the
dium benzoate, sodium metabisulphite and potassium laboratory. This review identied various compounds such as chi-
metabisulphite at a concentration 1000 mg per ml concentration tosan, plant extracts and essential oils that t this description. Salts
were separately applied to control anthracnose in bell pepper provide another alternative which might either surpass the ef-
(Capsicum frutescence) caused by C capsici. When salt concentration ciency of synthetic chemicals or work synergistically with these
were applied to wounded Capsicum fruit prior to inoculation with natural compounds. There is evidence showing that chitosan
C. capsici, 39%e82% smaller lesions were observed compared to formulation alone or in combination with other natural products
control (Ajith and Lakshmidevi, 2011). Even though this study are excellent candidates for consideration as alternatives for the
suggests the use of these salts in post-harvest disease management control of post-harvest rots of Capsicum spp. Studies on the use of
practices, more research remains to be done. plant extracts in the post-harvest control of chilli anthracnose are
A. Ali et al. / Crop Protection 90 (2016) 132e141 139

still lacking, therefore further studies should be performed in vivo, Boonyapipat, P., 2013. Study of three fungicides to control anthracnose (Colleto-
trichum capsici) in chilli (Capsicum frutescens): case study from Songkhla
since scientic evidence has already demonstrated their efcacy
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