Transcript
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 Article history: Received 27 January 2016Received in revised form15 July 2016Accepted 21 July 2016Available online 7 September 2016 Keywords: AnthracnoseChilliPost-harvest managementNatural products Colletotrichum spp a b s t r a c t Chilli anthracnose, caused by Colletotrichum spp, is one of the main causes for post-harvest decayof chilli.It can develop on the eld, during long distant transport, cold storage and shelf-life. In conventionalagriculture, the whole plant including the fruits, are sprayed with fungicides as a prerequisite for post-harvest control of chilli anthracnose. Due to consumer concerns regarding the use of synthetic fungicidesand the demand for safer storage methods, the use of synthetic fungicides is no longer allowed for thepost-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.In this review, results published within the last decade have been summarized and alternative ap-proaches to synthetic fungicides for post-harvest control of chilli anthracnose were discussed in detail.Overall, the use of natural antimicrobials, biocontrol agents, resistant cultivars and ozone shows promiseas treatments that can be adopted on a commercial scale to control post-harvest chilli anthracnosecaused by Colletotrichum species. © 2016 Published by Elsevier Ltd. Contents 1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1331.1. Infection process of anthracnose and symptoms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1332. Control management practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1342.1. Conventional practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1342.1.1. Cultural practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1342.1.2. Chemical control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1342.2. Alternative management practice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1352.2.1. Resistant cultivars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1352.2.2. Natural products . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1352.2.3. Biological control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1372.2.4. Physical control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1372.2.5. Salts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1383. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138Acknowledgement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 * Corresponding author. ** Corresponding author. E-mail addresses:
[email protected] (A. Ali),
[email protected] (P.K. Bordoh),
[email protected] (A. Singh),
[email protected] (Y. Siddiqui),
[email protected],
[email protected] (S. Droby). Contents lists available at ScienceDirect Crop Protection journal homepage: www.elsevier.com/locate/cropro http://dx.doi.org/10.1016/j.cropro.2016.07.0260261-2194/ © 2016 Published by Elsevier Ltd. Crop Protection 90 (2016) 132 e 141 1. Introduction Pepper belongs to the nightshade family (Solanaceae) and thegenus Capsicum . It srcinated from the Americas, and was culti-vated in New Zealand, South Africa, Malaysia and other Asiancountries. There are approximately 20 e 27 species of Capsicum ; outof which ve are domesticated and used as fresh vegetables andspices: Capsicum annum L., Capsicum frutescens L., Capsicum chinese Jacq., Capsicum pubescens Ruiz et Pavon. and Capsicum baccatum L and grown worldwide (Than et al., 2008b). Within these vedomesticated Capsicum species there are several cultivars. Thesecultivars are of immense economic importance displaying differentshapes, sizes, colors and avors. The most cultivated Capsicum va-rieties are Capsicum annuum (Tong and Bosland,1999) followed by C. frutescens (Bosland and Votava, 2003). These cultivars of Capsicum can be broadly categorized under Chilli (hot) or sweetpeppers.Thefruitof Capsicum hasavarietyofnames,suchas ‘ chilli ’ , ‘ chilli pepper ’ or ‘ pepper ’ depending on differences between theEnglish-speaking countriesand type offruits. Chillipepper ishottomild in taste due to the presence of capsaicin whereas, ‘ sweet ’ pepper taste sweet as the name suggest and do not producecapsaicin.SweetpeppersarenativetoMexico,CentralAmerica,andhave been propagated throughout the world including tropics,subtropics and temperate regions.Despitetherichnutritionalandeconomicvalueof Capsicum spp,which has promoted their cultivation as a major cash crop, theirproduction has been greatly hindered by a variety of pests anddiseases. Amongst these, Colletotrichum is a largegenus comprisinga number of important species that are the most prevalent fungalpathogens causing diseases in diverse tropical and subtropicalfruits and vegetables. Almost every crop including chilli grownthroughout the world is susceptible to one or more species of Colletotrichum . Recently, the genus Colletotrichum was listed as theeighth most important group of phyto-pathogenic fungi in theworld, based on perceived scienti c and economic signi cance(Dean et al., 2012).The prominence of Colletotrichum spp. and the losses caused byanthracnose post-harvest disease, presents a mounting threat tocommercially production of chilli. For instance, it anthracnose iswidely known to cause severe losses in yield as it affects immaturefruitsinthe eldandmaturefruitsafterharvest,duringstorageandtransit when conditions are favorable (Thind and Jhooty, 1985;Hadden and Black, 1989; Jayalakshmi et al., 1999; Bosland andVotava, 2003). Moreover, in Thailand, yield losses of up to 50%have been reported (Pakdeevaraporn et al., 2005) while in India,disease incidence levels ranged between 66% and 84% resulting inyield losses of 12 e 50% (Thind and Jhooty, 1985; Bagri et al., 2004;Sharma et al., 2005). In Korea, yield losses of 15% were reported(KimandPark,1988)and50%inMalaysia(Sariah,1994).Apartfrom pre-harvest losses, fruit quality deterioration of chilli ranging be-tween21and47%duetochillianthracnosehavealsobeenreportedin Sri Lanka (Anon,1993).Cross infection and pathogenicity studies have revealed thatchilli anthracnose is caused by several species of Colletotrichum namely Colletotrichum acutatum (Simmonds), Colletotrichum capsici (Syd.) Butler and Bisby, Colletotrichum gloeosporioides (Penz) Penz.and Sacc. and C. coccodes (Wallr.) S. Hughes (Simmonds, 1965; Johnston and Jones, 1997; Voorrips et al., 2004; Pakdeevarapornet al., 2005; Sharma et al., 2005; Than et al., 2008a; Ramdial andRampersad, 2014). More recently Colletotrichum fruticola Prihas-tuti,L.Cai & K.D.Hyde and C. siamense Prihastuti,L. Cai & K.D. HydehavealsobeenreportedascausativeagentsforchillianthracnoseinIndia (Sharma and Shenoy, 2014). However, the most economicallyimportant pathogens of chilli anthracnose are C. capsici and C. gloeosporioides . C. capsici generally infects mature fruits whiles C. gloeosporioides infects both green and mature fruits(Pakdeevaraporn et al., 2005) (Table 1). However, C. acutatum hasspeci cally been reported and identi ed as the cause of ‘ earlyanthracnose ’ disease in immature green bell peppers in Ohio,Florida and Trinidad (Harp et al., 2008, 2013; Lewis-Ivey et al.,2004; Ramdial and Rampersad, 2014).As a result of the serious post-harvest losses caused by due tochillianthracnose,severalattemptshavebeenmadetomanagethisproblem. Traditionally, cultural techniques such as good sanitationpractices (farm cleanliness); proper disposal of f rotten fruits, usingclean equipment and proper harvesting techniques have beenemployed to control post-harvest anthracnose. Additionally, syn-thetic fungicides such as propiconazole, difenoconazole, carben-dazim, benomyl, maneb and captan (Gopinath et al., 2006;Boonyapipat, 2013) have been used in the pre-harvest control of chilli anthracnose as a pre-requisite for the post-harvest control of the disease. It is imperative to note that benomyl (not registered orapproved in Australia) and its associated fungicides carbendezimandthiophanatemethyl(bothofwhichregistered)hasraisedmajorhealth concerns suchas causingeye defects, and other birth relatedeffects by disrupting the process of cell division making their useunacceptable and dangerous ( APVMA, 2009). Additionally, theemergence of resistant strains of C. capsici isolates in chilli fruitagainst benomyl, which were cross-resistant to thiophanate-methyl and carbendazim was reported in Malaysia (Sariah, 1989).Recently, resistance of C. truncatum to benomyl has also been re-ported in Trinidad (Ramdial and Rampersad, 2014). In addition,long term usage of synthetic chemicals is known tohavea negativeimpact on the environment especially soil and water resources(Wilson et al., 1999; Northover and Zhou, 2002). The increasinghealth concerns expressed by consumers and the intention of governments to regulate pesticide use and their residues in freshproduce have necessitated the development of non-toxic alterna-tive management techniques for anthracnose control. Therefore,this review highlights alternative approaches to chilli anthracnosemanagement during the post-harvest phase. 1.1. Infection process of anthracnose and symptoms Understanding the etiology of the disease is relevant fordeveloping strategies to control anthracnose. In most cases, post-harvest fungal infections occur when conidia from acervuli ormicro-sclerotia from the eld are splashed by irrigation or duringrainfall on healthy fruit and leaves especially. Additionally, humid,warm and wet conditions aggravate the rate of infection since thefungus thrives best under such conditions. The splashed conidiaattach themselves to the fruits surface and begin to germinateproducing appressoria. The appressoria develops a specializedstructure called infection pegs which then penetrate into the fruitepidermis. At this stage, the fungal infection may remain quiescentas a biotroph. Activation of quiescent infections occurs duringripening and senescence of the fruit after harvest (Prusky andLichter, 2007); followed by colonization of fruit tissue, appear-ance of symptoms such as circular sunken spots (watersoaked) andproduction of acervuli and sporulation (Bailey et al., 1992; Pruskyet al., 2000). The known developmental stages where the fungusmight become inactive are during penetration, initiation of germination, germ tube elongation, formation of appressorium orsubsequent infection. Based on the de nition of Swinburne andBrown (1983), it was widely accepted that failure of spores togerminate or develop beyond any subsequent stage is due totemporary adverse physiological conditions imposed by the hostdirectly on the fungus or indirectly by exhibiting temporary A. Ali et al. / Crop Protection 90 (2016) 132 e 141 133 resistance against the fungus. Temperatures around 27 C and highhumidity (a mean of 80%) are optimal for anthracnose diseasedevelopment (Roberts et al., 2001). Other reports also state thattemperatures of 20 C and 25 C favor colony growth and sporu-lation(Mello etal., 2004). Kim andPark(1988) determinedthatthe most important climatic parameter related to anthracnose devel-opment on chilli is relative humidity. Symptoms of chilli anthrac-nose are similar irrespective of the host species. Typically diseasesymptoms on fruits are characterized by small, depressed water-soaked lesions, which are sub-circular or angular with translucentlight brown margins (Figs.1 and 2). At advance stages of infection,the circular or angular sunken lesions are covered with concentricrings of wet, gelatinous spores from salmon-colored fungal fruitingbodies (acervuli) due to the colour of the spores with numerousblack spines (setae) (Roberts et al., 2001). 2. Control management practices This review reports on alternative post-harvest managementpractices as opposed to the use of synthetic fungicides, which wereclustered in ve categories: (i) resistant cultivars (ii) Naturalproducts (antimicrobials) (iii) biological controls (iv) physical con-trol(v)salts.Itisfairtostatethat,mostof thesealternativesarestillfar from practical application. 2.1. Conventional practices 2.1.1. Cultural practices Several cultural practices have been used as pre-requisites tocontrol post-harvest chilli anthracnose due to the special etiologyofthepathogen.Theseprecautionarymeasuresareimplementedtominimize the rate of infection and reducing the infection pressureevenbeforefruitsarematureandharvested.Thanetal.(2008a,b)intheir review reported that, chilli seeds free of pathogen should beplanted and weeds eliminated. They also reported that rotatingcrops that are not alternative hosts to Colletotrichum spp every 2 e 3years is very effective for controlling post-harvest chilli anthrac-nose. Other sanitation practices include the provision of gooddrainagesystemsonthe eldtochanneloutwastewaterorsewageduring on-farm fruit disinfection such as fruit washing at packinghouses or during irrigation regimes, and nally removal of plantdebris which may serve as source of inoculum (Than et al., 2008b).If there was history of disease in a particular eld, then other cropsshould be rotated in isolation from other solanaceous plant for atleast 2 years (Roberts et al., 2001). By doing so, the life cycle of thepathogen on the eld tobegin an infectionprocess is disrupted andthe chance of infection is reduced since debris of most solanaceouscrop (after harvest) may become inoculum and a haven for thefungus.Attheendofthegrowingseason,itisalsorecommendedtodeep plough to completely cover diseases plants or removinginfectedplantdebrisfromthe eld(Nelson,2008).Earlyplantingof chilli or planting cultivars that bear fruit within a short ripeningperiod to allow the fruit to escape fungal infection is also recom-mended. Other alternative sanitation practices such as weeding,removal of infected or wounded fruits should be carried out regu-larly to prevent the pathogens from using such wounds as sites of infection. 2.1.2. Chemical control Synthetic chemicals have long been used as one of the mostcommonpracticesfor controlling chilli anthracnose. However, theycannot be used in post-harvest management of anthracnosebecause of health concerns to humans and potential buildup of pathogen-resistance to the fungicides. By ensuring proper culturalpractices without using synthetic fungicides, post-harvestanthracnose can be reduced signi cantly. Fungicide tolerance,however, often arises quickly, if a single compound is relied upontooheavily(Staub,1991). Plantingseeds whichare freeofpathogeninfections or pre-treatment of seeds with synthetic chemicals(Thiram, Captan and Bavistin) are commonly used to controlanthracnose (Choudhary et al., 2013). Commonly used fungicideswhich have been traditionally recommended in pre-harvest chillianthracnose management include Manganese ethyl-enebisdithiocarbamate (maneb), propiconazole, difenoconazole 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 ), sweetbellpepper ( C. annuum ) Diseased chilli peppers are withered with brown to blacknecrotic patches Fruits develop water-soaked and small black circularspots on the surface.(Roberts et al., 2001;Chanchaichaovivat et al.,2007; Warin et al., 2009) C. gloeosporioides Green and red chilli pepper ( C. annum ) Lesion usually develops on ripe fruits. Lesions are small,inde nite, slightly sunken, water-soaked spots that mayenlarge rapidly and coalesce. Fruiting bodies form in concentric circles to cover thesurface of the lesions Lesionsappeartanorbrownandarecoveredwithsalmonto orange gelatinous spores.(Kim et al., 1999; Kim et al.,2004) Fig.1. Anthracnose on green and red chilli pepper ( C. annum cv. Nokwang') caused by C. gloeosporioides (source: Kim et al., 1999). A. Ali et al. / Crop Protection 90 (2016) 132 e 141 134 and cabendazim (Smith, 2000; Gopinath et al., 2006). The ef cacyof strobilurin fungicides, azoxystrobin (Quadris and Amistar 25SC ® ), tri oxystrobin (Flint), pyraclostrobin (Cabrio) have also beenlabeled to control chilli anthracnose but only preliminary reportsexist on the ef cacy of these fungicides against the disease in itssevere form in eld trials (Alexander and Waldenmaier, 2002;Lewis and Miller, 2003). Despite the positive results obtained inpre-harvest studies in chilli anthracnose managements using thesesynthetic chemicals, some Colletotrichum spp like Colletotrichumcapsici, C. gloeosporioides and C. siamese have developed resistanceto thiophanate-methyl, Strobilurin-fungicides (azoxystrobin andkresoxim-methyl) and carbendazim (Sariah, 1989; Inada et al.,2008; Hu et al., 2015). 2.2. Alternative management practice 2.2.1. Resistant cultivars Research has shown that, the usage of resistant varieties notonly reduces damages from diseases, but also minimizes chemicaland mechanical expenditures of disease control (Grichar et al.,1998; Besler et al., 2001; Monfort et al., 2004). However, thereduction in some cases maybe complete or partial. Resistant cul-tivars aremostly used asa control measures tomanage pre-harvestanthracnose in chilli fruit or plants in the eld. Some genetic re-sources resistant to anthracnose in chilli have been independentlyreportedfromdifferentcountriesandregions(Kimetal.,1987;Parket al.,1987; Hong and Hwang,1998; Pae et al.,1998; AVRDC,1999;Yoon and Park, 2001; Yoon, 2003; Mongkolporn et al., 2004; Kimet al., 2004, 2007; Voorrips et al., 2004; Lee et al., 2010;Shivashankar et al., 2010; Kim et al., 2011; Rahman et al., 2011;Syukur et al., 2013; Sun et al., 2015).Few studies about the use of resistant cultivars to control post-harvest rot in pepper have been reported. Earlier research in thePhilippines reported that C. annuum of different lines show resis-tance to anthracnose caused by C. gloeosporioides isolated from C. annuum . Over 50.0% of the fruits in lines PBC 452, PBC 454 andPBC595 had lesionsin lessthan5 daysafter inoculation,whereasittook6daysforfruitinthreelines(PBC365,PBC371andPBC518),8days for fruit in line PBC 370, and 11 days for fruit in line PBC 495.They observed that fruits of PBC 595 had the largest lesions, whilefruits 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 BC 1 population derived from hybrid with C. annuum line 77013 (sus-ceptible) using QTL analyses. Resistance tests were performed ondetached mature green or mature red fruit under laboratory con-ditions (storage at 28 C, 95% RH for 48 h and additional 5 daysunder plastic wrap) and then evaluated for disease incidence(anthracnose) and overall lesion diameter. The mean disease scoresof the resistant parent PBC932 were 12.2% for GD (Disease inci-dence in Green fruit stage), 0.8 mm for GO (Overall lesion diameterin Green fruit stage), 2.10mm for GT (True lesion diameterin Greenfruit stage), 52.5% for RD (Disease incidence in Red fruit stage),4.0 mm for RO (Overall lesion diameter in Red fruit stage) and5.8 mm for RT (True lesion diameter in Red fruit stage). Thesevalues were all signi cantly lower than those of the susceptibleparent 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 F 1 individuals fell in between PBC932 and 77013, but skewed towardPBC932 in all six phenotyping methods so that score values werenot signi cantly different from those of the resistant parent. Theyproposed that the resistance from C. chinense PBC932 is completelydominant over the susceptibility, at both the mature green and redstages. Based on a map of the 14 linkage groups, that included 385markers (SSR, InDel and CAPS), covering a length of 1310.2 cM,inclusive Composite Interval Mapping (ICIM) revealed main effectQTLs were located on the P5 chromosome for all fruit stages andresistancecriteria,andfourminor-effectQTLsweredetectedonlyatthe green mature stage. Identi cation of recombinant individualssuggested that resistance in green versus red fruit may becontrolledbydistinctgeneswithintheQTLintervalonP5,Sunetal.(2015). Further studies need to be undertaken to investigate theimportance of these distinct genes in the post-harvest manage-ment of chilli anthracnose. 2.2.2. Natural products 2.2.2.1. Plant extracts. Certain antimicrobial or metabolic com-pounds synthesized by plants might be very good alternative forcontrolling diseases of tropical fruits and vegetables. In chilli,severalinvitrostudieshaveshowntheef cacyofcertainmedicinalherbsorplantextractsagainst Colletotrichum spp.(Charigkapakorn,2000; Begum et al., 2007; Nduagu et al., 2008; Johnny et al., 2011;Saravanakumar et al., 2011; Ajith et al., 2012) (Table 2). A lot of studies have been conducted on the pre-harvest control of chillianthracnose (Charigkapakorn,2000; Ajith et al., 2012; Rashid et al.,2015) but very few studies have reported as post-harvest control.Plant parts such as rhizomes, bark, roots and leaves have shownsigni cant antimicrobial properties against Colletotrichum sp thatcauses chilli anthracnose (Table 2).Research has shown that an aqueous leaf extract of Plumbagoindica L. had a signi cant suppressive effect on C. gloeosporioides interms of conidial germination (98.9%) and radial mycelial growth(98.8%). In vitro studies have shown that botanicals or plant ex-tracts from Cathranthus roseus L. G.Don., Coleus aromaticus Benth., Manilkara zapota L. P.Royen., and Azidirachta indica A. Juss confersantifungal effects on the radial mycelial growth of C. capsici responsible for anthracnose disease in bell pepper ( Capsicum fru-tescence ) (Ajith et al., 2012). Singh and Khirbat (2014), reported on the ef cacy of aqueous extract of three wild plants viz., siras( Albizza lebbeck L. Benth.), babul ( Acacia Arabica Lam.) and Cler-odendron ( Clerodendron infortunatum L.) to control fruit rot in C. annuum caused by C. capsici . The leaf extracts of ‘ siras ’ , ‘ babul ’ and ‘ clerodendron ’ caused signi cant inhibition of spore germina-tion in C. capsici with percentages of inhibition of 100%, 86.6% and Fig. 2. Anthracnose on green bell pepper fruit caused by C. capsici. A. Ali et al. / Crop Protection 90 (2016) 132 e 141 135
