Chilli Leaf Curl Virus an Emerging Threat to Chilli in India

Available online at www.ijpab.com Zehra et al Int. J. Pure App. Biosci. 5 (5): 404-414 (2017) ISSN: 2320 – 7051 DOI: http://dx.doi.org/10.18782/2320-7051.5471 ISSN: 2320 – 7051 Int. J. Pure App. Biosci. 5 (5): 404-414 (2017) Review Article

Syed Berjes Zehra1, Asif Ahmad2, Abhishek Sharma5, Shakeela Sofi3, Azra Lateef1, Zaffar Bashir3*, Mohit Husain4 and Jagdeeh Prasad Rathore5 1Department of Vegetable sciences, 2Department of Plant Pathology, 3Department of Microbiology, 5Division of Fruit Science 4Faculty of Forestry, Benhama-Ganderbal, 5Department of vegetable sciences (PAU-Ludhiana) Sher-e-Kashmir University of Agricultural Sciences and Technology of Kashmir, Shalimar-190025 *Corresponding Author E-mail: [email protected] Received: 19.08.2017 | Revised: 22.09.2017 | Accepted: 26.09.2017

ABSTRACT Chilli is known to be affected by forty five viruses. Twenty four of them have been reported to occur naturally and rest can infect on artificial inoculation. Among the twenty four viruses reported to occur naturally on chilli, eleven viruses have been reported from India. Among all, the chilli leaf curl virus is the most destructive virus in terms of incidence and yield loss. In severe cases 100 percent losses of marketable fruit have been reported. Chilli leaf curl locally known as murda is a most destructive disease of chilli in India. However, the viral nature of the disease has not been proved experimentally until Ashrafi Jha from IARI established this by grafting, later it was proved that leaf curl of chilli is caused by tobacco leaf curl virus (Ruga tabaci) which is transmitted by vector B. tabaci. The diseases caused by begomoviruses are easily recognized by their distinctive symptoms in plants infected with these viruses. The symptoms are broadly of three types: a) vein yellowing, b) yellow mosaic and c) leaf curl. Control of ChiL CV viruses by chemicals or cultural management practices has not been very effective. The planting of resistant cultivars offers the best way to control these viruses. A number of sources of virus resistance have been reported in wild and cultivated lines and several virus- resistant lines have already been released. Many multiple virus-resistant varieties have been developed at Punjab Agricultural University, Ludhiana. Important multiple resistant lines are Perennial, BG-1, Lorai, and Punjab Lal. Weeds act as potential sources of begomoviruses, several weed families particularly Malvaceae have been found to harbor viruses.

Key words: Chilli leaf curl, begomo viruses and B. tabaci and management.

INTRODUCTION Capsicum, family Solanaceae. It has originated Chilli (Capsicum annuum L.) is an important in Mexico, Southern Peru and Bolivia71. There spice grown for its fruits, which are used in are mainly five cultivated Capsicum spp. viz. green as well as ripe dried form for its C. annuum, C. baccatum, C. chinense, C. pungency. Chilli belongs to the genus frutescens, and C. pubescens.

Cite this article: Zehra, S.B., Ahmad, A., Sharma, A., Sofi, S., Lateef, A., Bashir, Z., Husain, M. and Rathore, J.P., Chilli Leaf Curl Virus an Emerging Threat to Chilli in India, Int. J. Pure App. Biosci. 5(5): 404-414 (2017). doi: http://dx.doi.org/10.18782/2320-7051.5471

Zehra et al Int. J. Pure App. Biosci. 5 (5): 404-414 (2017) ISSN: 2320 – 7051 Among these, C. annuum L. is the most widely India, viz., cucumber mosaic virus, tobacco cultivated species all over the world for its leaf curl virus25, Indian chilli mosaic virus, pungent (chilli syn. Hot pepper) and non- potato virus Y, potato virus X, tobacco ring pungent (sweet pepper) fruits. Chilli crop is spot virus, pepper veinal mottle virus and raised over 1832 thousand hectares in the pepper vein bending virus, chilli leaf curl world, with a production of 2959 thousand virus, tomato leaf curl New Delhi virus30, tonnes4. Highest production per unit area of capsicum chlorosis virus a tospo virus. 36,585 kg per ha was recorded in Japan4. It Recently cucumber mosaic virus subgroup II was first introduced in India by Portuguese has been reported to infect chilli6 from western towards the end of 15th century. The top ten Himalayan region of India. In Mexico, two chilli producing countries India, China, Gemini viruses are considered to be the main Ethiopia, Myanmar, Mexico, Vietnam, Peru, pathogens in pepper i.e pepper huastico Virus Pakistan, Ghana and Bangladesh accounted for (PHV) and pepper golden Mosaic virus more than 85% of the world production in (PepGMV) formerly called texas pepper virus. 2009; lion‟s share is taken by India with 36% Among all, the chilli leaf curl virus is the most share in global production, followed by China destructive virus in terms of incidence and (11%), Bangladesh (8%), Peru (8%) and yield loss. In severe cases 100 percent losses 4 33 Pakistan (6%) .In India Chilli is cultivated in of marketable fruit have been reported . all the states including Andhra Pradesh, CHILLI LEAF CURL VIRUS Karnataka, Maharashtra, Orissa, Rajasthan, The etiology of chili leaf curl disease was Tamil Nadu, West Bengal and Punjab over an reported during the 1960s13. Since it is a area of 804790 hectares with total production begomovirus and it is transmitted in a of 1276300 metric tonnes of dry chilli with persistent manner by white flies of the species productivity of 1.5 metric tonnes per hectare. complex Bemisia tabaci7. The typical Andhra Pradesh alone contributes 46 per cent symptoms consisting of leaf curling, rolling, of total chilli production of India. India puckering, blistering of interveinous areas and contributes one-fourth of the total quantity of thickening and swelling of the veins, chilli exported in the world4.Tremendous shortening of internodes and petioles, foreign exchange is being earned through crowding of leaves and stunting of whole export of chilli powder, oleoresin of low, plants, older leaves may become leathery and medium or high pungency. At present 169500 brittle60. quintals of dry chilli of the worth of Rs Chilli leaf curl locally known as murda is a 160408 lakh is exported from the country4. most destructive disease of chilli in India. The Indian chillies are exported to over 90 etiology of chili leaf curl disease was reported countries in the world4. If some varieties are during the 1960s13. Mites famous for red colour because of the pigment (Polyphagotarsonemus latus)) and thrips capsanthin, others are known for biting (Scirtothrips dorsalis) were considered to be pungency attributed by capsaicin. It is a rich responsible for causing chilli leaf curl source of vitamins A and C. India is the only symptoms42. Johnpulle28 considered it to be country rich in many varieties with different caused by both mites and thrips. Amin3 quality factors. World‟s hottest chilli “Naga concluded that leaf curl in chilli was due to Jolokia” is cultivated in hilly terrains of Assam feeding of thrips and mites and not by an 4 in a small town Tezpur, India . infectious agent. However, the viral nature of Chilli is known to be affected by forty five the disease has not been proved experimentally viruses20. Twenty four of them have been until Ashrafi Jha from IARI in 1953 reported to occur naturally and rest can infect established this by grafting, later it was proved on artificial inoculation. Among the twenty that leaf curl of chilli is caused by tobacco leaf four viruses reported to occur naturally on curl virus (Ruga tabaci) which is transmitted chilli, eleven viruses have been reported from by vector B. tabaci56. Copyright © Sept.-Oct., 2017; IJPAB 405

Zehra et al Int. J. Pure App. Biosci. 5 (5): 404-414 (2017) ISSN: 2320 – 7051 Based on partial DNA-A sequences, a types: a) vein yellowing, b) yellow mosaic and monopartite begomovirus was reported to be c) leaf curl. Leaf crinkle or leaf curl complex associated with the disease which was a strain was observed on chillies from India and of the Chili leaf curl virus (ChiLCV) from abroad25. Curling of leaf margin, reduction in Pakistan 30. leaf size, vein clearing were observed in India, VARIABILITY IN BEGOMOVIRUSES Sri Lanka and USA46. Abaxial curling of the INFECTING CHILLI leaves accompanied by puckering, thickening There has been a tremendous increase in the and swelling of the veins were observed by number of begomoviruses that have been many study. Appearance of most prominent isolated in recent years as these are assumed to symptoms such as vein clearing followed by have been co-evolving with their veinal distortion, swelling of veins and dicotyledonous plant hosts for a long time54. veinlets on dorsal side. The typical symptoms Currently, 35-70 geminiviruses are being consisting of leaf curling, rolling, puckering, discovered and characterized every year. Out blistering of interveinous areas and thickening of the total geminiviruses, 200 defined species and swelling of the veins, shortening of belong to begomoviruses, thus making it the internodes and petioles, crowding of leaves largest genus in the family. and stunting of whole plants, older leaves may So far, 34 recognized and 18 tentative become leathery and brittle60. species of begomoviruses have been found to OCCURRENCE naturally infect tomato. However, the Leaf curl of chilli has been reported by several begomoviruses with generic name „tomato leaf investigators from India and abroad25. curl viruses are the most devastating viruses. However, the virus nature of the disease has In India, eight different viruses cause Tomato not been proved experimentally by them. Park leaf curl disease (ToLCD). Three of these and Fernando42 questioned the virus nature of viruses, tomato leaf curl Gujarat virus disease. Pruthi and Samuel44 observed typical (ToLCGV), tomato leaf curl India virus leaf curl symptoms on 15-25% of chilli plants (ToLCIV) and tomato leaf curl New Delhi under field conditions and they considered it to virus (ToLCNDV) occur in northern India be caused by the Leaf curl virus of tobacco while the other two, tomato leaf curl reported by Pal and Tandon39.Latter on chilli Bangalore virus (ToLCBV) and tomato leaf leaf curl disease caused by whitefly (Bemisia curl Karnataka virus (ToLCKV) occur in tabaci) transmitted (WFT) geminivirus, southern India38. Khan30 found the association namely, pepper leaf curl virus (PepLCV) was of ToLCNDV and chilli leaf curl virus reported from India35, United States63, Nigeria2 (ChiLCV) in the infected samples of chilli and several other countries such as Pakistan, showing leaf curl disease based on the partial and Indonesia15. sequencing of a DNA-A like component. LOSSES However, the full length sequence of a The emerging threat of the viruses belonging begomovirus isolated from plants with chilli to the genus Geminivirus has been extensively leaf curl disease in India belongs to the species addressed earlier. The extent of yield loss ChiLCV-IN and has been found to be similar caused by some geminiviruses has been to tomato leaf curl Joydebpur virus58. Chilli estimated as high as 100% and in some areas leaf curl disease is caused by a complex infection with viruses has rendered the consisting of the monopartite chilli leaf curl growing of peppers uneconomical, causing virus and a DNA-β satellite component. whole fields to be abandoned prior to SYMPTOMOTOLOGY harvest21. Up to 96% loss in yield has been The diseases caused by begomoviruses are reported by Bhendi yellow vein mosaic virus45. easily recognized by their distinctive In legumes, the yield losses have been symptoms in plants infected with these estimated to be approximately $300 million viruses. The symptoms are broadly of three per year taking black gram, mungbean and Copyright © Sept.-Oct., 2017; IJPAB 406

Zehra et al Int. J. Pure App. Biosci. 5 (5): 404-414 (2017) ISSN: 2320 – 7051 soybean together. Economic losses due to white flies of the species complex Bemisia geminivirus infections in Cassava are tabaci7. As the begomoviruses are phloem estimated to be US $ 1300-2300 million in inhabited, so they are neither sap transmissible Africa67, US $ 5 billion for cotton in Pakistan nor seed transmissible. They are successfully between 1992-97, US $ 300 million for grain transmitted with the help of whitefly vector legumes in India70, and US $ 140 million in belonging to family Aleyrodidae which are the Florida for tomato alone. Tomato leaf curl and pests of herbaceous and woody plants. The tomato yellow leaf curl diseases are plant viruses are successfully transmitted by widespread in more than 20 countries; it whitefly belonging to genus Bemisia and caused heavy yield losses and wiped out Trialeurodes. In the genus Bemisia, only B. tomato crops in some areas20. Saikia and tabaci has been shown to be vector whereas in Muniyappa50 reported more than 90% fruit the genus Trialeurodes, T. vaporariorum, T. yield loss due to tomato leaf curl disease when abutilonea, and T. ricini transmit viruses. The infection occurred within four weeks of largest and most economically significant transplanting. groups of plant viruses transmitted by Bemisia ETIOLOGY tabaci are the Gemini viruses. Leaf curl of chilli has been reported by several WHITEFLY AS A VECTOR investigators from India and abroad. Mites, Whitefly belongs to family Aleyrodidae (order Polyphagotarsonemus latus and thrips, Hemiptera). It is a tiny insect, only 1–3 mm Scirtothrips dorsalis were considered to be long. The wings and bodies of the adults are responsible for causing chilli leaf curl covered with a powdery or flour-like white symptoms42. Johnpulle28 considered that chilli wax thus called whiteflies. On most host leaf curl disease is caused by mites and thrips. crops, eggs are laid and immature stages Pruthi and Samuel44 observed typical leaf curl develope on the abaxial surface of leaves. symptoms on 15-25% of chilli plants under Although they are mainly tropical insects, field conditions and they considered it to be whiteflies are found in all warmer parts of the caused by the Leaf curl virus of tobacco world, and several are also serious pests in reported by Pal and Tandon39. Latter on chilli greenhouses in temperate areas. Direct crop leaf curl disease caused by whitefly (Bemisia damage occurs when whiteflies feed in plant tabaci) transmitted (WFT) geminivirus, phloem, remove plant sap, and reduce plant namely, pepper leaf curl virus (PepLCV) was vigor. Whiteflies also excrete honeydew, reported from India47, United States55, Nigeria2 which promotes sooty mold that interferes and several other countries such as Pakistan, with photosynthesis and may lower harvest and Indonesia15. quality. In some hosts, damage can result from TRANSMISSION whitefly feeding toxins that cause plant Transmission of Leaf curl virus through M. disorders such as silver leaf of squash persicae, A. nasturtii and A. craccivora was (Cucurbita pepo)8 and irregular ripening of reported by Szalay64. Leaf curl or leaf crinkle tomato53. From a phytopathological occurring on chillies transmitted by Bemisia perspective, however, the main concern about tabaci, Bemisia gossypiperda has been whiteflies is their ability to transmit viruses. Of reported by Husain25. However, the viral the approximately 1,500 species of nature of the disease had not been proved whiteflies36, only a few have been shown to experimentally until Ashrafi Jha from IARI in vector plant viruses, and these vectors include 1953 established this by grafting, later it was the cotton/tobacco/sweet potato whitefly [B. proved that leaf curl of chilli is caused by tabaci (Gennadius)] and the greenhouse tobacco leaf curl virus (Ruga tabaci) which is whitefly (T. vaporariorum Westwood). B. transmitted by vector B. tabaci39. tabaci, by far the most important whitefly in Begomoviruses infect dicotyledonous plants terms of virus transmission, is typically and are transmitted in a persistent manner by polyphagous, is generally found in tropical and Copyright © Sept.-Oct., 2017; IJPAB 407

Zehra et al Int. J. Pure App. Biosci. 5 (5): 404-414 (2017) ISSN: 2320 – 7051 semitropical regions, and vectors circulative is controversial, and the only data begomoviruses, criniviruses, carlaviruses, available have been obtained with TYLCV. ipomoviruses, and torradoviruses. T. Thus, although no direct evidence of vaporariorum is also polyphagous, is found replication has been obtained so far, TYLCV nearly worldwide in both greenhouses and transcripts accumulate in B. tabaci and the fields, and transmits criniviruses and quantity of viral DNA seems to increase with torradoviruses. The banded winged whitefly time in the insect following an acquisition [T. abutiloneus (Haldeman)] is also a vector of period on infected tomato plants11. The a number of criniviruses. The castor bean minimum acquisition and inoculation feeding whitefly [Trialeurodes ricini (Misra)] has been period required by B. tabaci to transmit tomato reported to be a vector of the begomovirus leaf curl virus was found to be 30 minutes Tomato yellow leaf curl virus (TYLCV) in each. After acquisition the vector required 6 Egypt26, but this finding has not yet been hours to become viruliferous. Single whitefly confirmed. Also, Bemisia afer sensu lato has was able to transmit the virus. The whiteflies recently been described as a vector of the remained infective throughout their life span; crinivirus Sweet potato chlorotic stunt virus. the virus was not transmitted to the progeny of B. tabaci corresponds to a complex of genetic the whitefly9. The minimum acquisition and variants or haplotypes, usually referred to as inoculation – feeding periods for the vector, biotypes. Based on mitochondrial cytochrome Bemisia tabaci to transmit yellow leaf curl oxidase subunit (COI) gene consensus virus was 20 and 30 minutes, respectively. The sequences, a recent report has suggested that latent period was 21-24 hours. B. tabaci is a complex of at least 24 distinct MANAGEMENT cryptic species14. Although no morphological The foliar application of cypermethrin (0.01, characteristics can be used to distinguish 0.015%), deltamethrin (0.0028, 0.0042%) and between B. tabaci populations, genetic and dimethoate (0.03, 0.045%) was effective in behavioral differences have been used for reducing the whitefly incidence in green gram haplotype/biotype characterization. These field. In a study carried out with seven include isoenzyme profiling, bar-coding based insecticides, triazophos 0.04 per cent was most on conserved genes like the mitochondrial COI effective for managing populations of gene, life history traits, host range and/or host Scirtothrips dorsalis and preventing incidence preference, virus transmission competency, of leaf curl in chilli41. Neem seed kernal composition of endosymbionts, dispersal extract (5%) was found most effective than behavior, insecticide resistance, and Karanj and Tumba seed extract. Insecticide discontinuous gene flow8. Biological Imadalcoprid 17.8 SL (0.003%) was found differences between B. tabaci biotypes can effective than Malathion 50 EC (0.05%) and radically affect the emergence of a virus Methyl-demeton 25 EC (0.025%) in the disease by causing differences in transmission management of Whiteflies40. Two fold efficiency, host range, or mating behavior, as increase in nodulation and grain yield with is discussed below. Whiteflies can transmit 50% reduction in mung bean yellow mosaic plant viruses in a semipersistent manner (e.g., incidence in mung bean when dry powder of criniviruses) or a persistent manner (e.g., Clerodendron aculeatum was applied as soil begomoviruses). Interestingly, the persistent amendment in addition to six times foliar transmission of some begomoviruses involves spray at weekly interval70. Biological control a third partner. Thus, persistent transmission of includes use of parastoids of genera TYLCV by B. tabaci depends on chaperonin Eretmocerus and Encarsia and predators GroEL homologs produced by endosymbiotic belonging to numerous families5. bacteria36.Whether the persistent transmission Control of ChiLCV viruses by chemicals or of begomoviruses is propagative (i.e., the virus cultural management practices has not been replicates in the insect vector) or only very effective. The planting of resistant Copyright © Sept.-Oct., 2017; IJPAB 408

Zehra et al Int. J. Pure App. Biosci. 5 (5): 404-414 (2017) ISSN: 2320 – 7051 cultivars offers the best way to control these 2 moderately resistance, 19 susceptible and 13 viruses. A number of sources of virus highly susceptible31. The F1 progenies and resistance have been reported in wild and BC2 generations of certain crosses were found cultivated lines and several virus-resistant to be resistant for leaf curl complex. However, lines have already been released. Among these the BC1 generation of the cross, (Punjab lal x are lines with resistance to potato virus Y Pusa sadabahar) x Punjab lal, was found to be (PVY), tobacco etch virus (TEV), pepper highly resistant with a much reduced mottle virus (PeMV) and certain coefficient of infection. Plants collected in tobamoviruses. However, other viruses, thirteen wild populations of Capsicum annuum particularly cucumber mosaic virus (CMV) from Northwest Mexico were tested for and the leaf curl virus complex, continue to resistance to the pepper huasteco begomovirus pose a challenge to breeding programs. (PHV) that is transmitted by the whitefly SCREENING OF PEPPER GENOTYPES Bemisia tabaci Genadius. Plants of only two of AGAINST LEAF CURL DISEASE the populations either did not develop disease In India, breeding for resistance in chilli was symptoms after inoculation55.Two varieties of started in the late sixties but most of the chilli (Puri Red and Puri Orange) were found screening has been under field conditions, resistant to chilli leaf curl virus upon assessing disease incidence and disease subjecting to viruliferous white flies under severity. For disease rating, a coefficient of artificial screening. The results were further infection has been used, for which the confirmed by grafting the scion from typically percentage disease incidence was multiplied leaf curl affected chilli plants on the stocks of with a response value assigned to each Puri Red and Puri Orange varieties35. Pant observed disease severity grade. A good C1and Pant C2 showed significantly higher correspondence was usually obtained between yields compared to the better parent viz.NP- field and greenhouse assessments34. Since 46A by way of plot average as well as plant then, a number of lines with varying degrees yield. Leaf curl incidence was significantly of resistance have been identified; some of lower in both these varieties compared to NP- these lines also exhibit resistance or tolerance 46A, especially in case of Pant C1. Jawala, a to other viruses66.Many multiple virus-resistant selection developed from the cross N.P. 46-A× varieties have been developed at Punjab Puri Red was found to be promising against Agricultural University, Ludhiana. Important leaf curl disease. Sangar51 reported that the multiple resistant lines are Perennial, BG-1, varieties JCA-248, JCA-218, Pant C-1, NP-46, Lorai, and Punjab Lal59. These sources have Pusa Jawala and JCA-196 were resistant been used for development of high yielding against chilli leaf curl virus. GKC-29, BS-35 hybrids of chilli like CH-1 and CH-3. Pusa and EC-497636 genotypes showed no Jwala, a chilli variety resistant to virus was symptoms of leaf curl and they were further developed by Tewari and Ramanujam65 subjected to grafting with scions of susceptible followed by Pant C-1 and Pant C-2. Cultivars Pusa Jwala and later on to PCR amplification, Puri Red, Puri Orange, G2 and Kondiverum the three lines didn‟t show any amplification were known to be resistant to mosaic by confirming absence of viral genome were artificial inoculation. Out of one hundred and declared as symptomless genotypes32. Pusa fifty nine varieties/crosses/selections tested, Jawala, Surajmukhi, Japani lounge and JCA- Pure red, Cfr-10, LCA-135, LCA-412 and 196 were found resistant against Chilli leaf Pant C-1 were found to be resistant /tolerant to curl virus31. These sources have been used for mosaic complex, whereas others showed the development of high yielding hybrids of susceptible reaction27. 37 chilli (Capsicum chilli like CH-1 and CH-3. Kumar32 screened annuum) genotypes were screened for the three hundred and seven genotypes belonging incidence of leaf curl virus, 3 (Pusa Jwala, to four cultivated and one wild species of Surya Mukhi and Loungi) were rated resistant, Capsicum against pepper leaf curl virus Copyright © Sept.-Oct., 2017; IJPAB 409

Zehra et al Int. J. Pure App. Biosci. 5 (5): 404-414 (2017) ISSN: 2320 – 7051 (PepLCV) out of which only three genotypes hosts. A. conyzoides L., Oxalis acetosella L., viz., GKC-29, BS-35 and EC-497636 were Oxalis corniculata L., Euphorbia geniculata found resistant. EC 4020, EC 6589, EC 7299, L., Synedrealla hodiflora L., Galinsoga ED 7338, EC 9293, Puri Red and Puri Orange parviflora Cav., Solanum nigrum L., Cassia -field resistant to leaf curl virus, Surjamani and uniflora L., Parthenium hysterophorus L. and Perennial resistant to leaf curl virus also Walteria americana L. in all the seasons with resistant to chilli mosaic. S 118, S 114 high disease incidence during summer (derived from Perennial x Long Red) resistant compared to Kharif and rabi. The highest to resistant to ChLCV62. Jwala (derived from ToLCV incidence (37.68 %) was recorded on Puri Red and N.P. 46A) resistant to ChLCV Euphorbia geniculata L., reported to be one of and mosaic. C.annuumn var. angulosumn the reservoir hosts for ToLCV Rhynchosia tolerant to leaf curl virus also tolerant to CMV, mosaic geminivirus, whose natural host is Pusa Jwala also resistant to TMV, susceptible Rhynchosisa minima,was transmitted to beans to CMV. Delhi Local tolerant to leaf curl virus in which it caused typical mosaic, dwarfing also tolerant to TMV, immune to CMV and and leaf distortion19. Ramappa48 reported more PVX and C.annuum S 38-3-19, S42-2-4 - than 30 weed species to harbour tolerant to leaf curl virus -also tolerant to PVX begomoviruses under natural conditions. and CMV66.Despite efforts by various research Kang29 reported the presence of cotton leaf groups, it was not possible to establish genetic curl virus in symptom less plants of chilli and control of the resistance gene(s). This could be China rose using polymerase chain reaction. attributed to shortcomings in screening Reddy49 has reported Parthenium techniques, virus detection methods, hysterophorus, a wide spread weed in India, as variability of virus causing leaf curl disease a reservoir host of begomovirus. etc. To sustain production and enhance Sivalingam61 tested seven weed and profitability of chilli cultivation, it is important three cultivated crop plants for the detection of to identify confirmed sources of resistance and begomoviruses by polymerase chain reaction utilize them in resistance breeding programme using universal DNA-β primers of by employing conventional and molecular begomoviruses and primers specific to the coat breeding approaches. protein gene of begomoviruses infecting ALTERTNATE HOSTS cotton (CLCBV-CP) and out of these, only Weeds are widely distributed in the world and two weed species Tribulus terrestris and have high environmental adaptability. The first Cucumis sp. were found to be positive for both studies of whitefly transmitted viruses in Latin DNA-B and CLCBV-CP gene amplification America date from 1930s and concern several which were not showing any visible symptoms weed species in the families Malvaceae and in the field. This indicated that these weed Euphorbiaceae in Puerto Rico and Brazil10. species are the probable alternate hosts for Many reports have demonstrated that weeds CLCBV. serve as reservoir or alternative hosts for Gupta23 tested the efficiency of begomovirus survival and spread. Sastry52 Bemisia tabaci (Gennadius) populations reported that weed hosts such as collected from different plant-hosts viz., Acanthosperomum hispidum, Ageratum cotton, potato, tomato, soybean, brinjal and conyzoides, Parthenium hysterophorus, weed (Sida sp.) and reported that whitefly Datura stramonium, E. geniculata and from different hosts can efficiently transmit Gynandropisis pentaphylla as sources of the virus to cotton if it has been successfully inoculum for tomato leaf curl virus disease. acquired by the whitefly. Valand and Muniyappa37 studied the host Sivalingam61 identified carrot (Daucus range of tobacco leaf curl virus and found carota) as a new host for begomoviruses virus was transmissible to nine crop plants, causing yellow mosaic disease of carrot in the eight ornamental plants and eighteen weed states of Uttar Pradesh, Haryana and Punjab. Copyright © Sept.-Oct., 2017; IJPAB 410

Zehra et al Int. J. Pure App. Biosci. 5 (5): 404-414 (2017) ISSN: 2320 – 7051 The nucleotide similarity index of the engaged Pumpkin (Cucurbita maxima) and snap melon viruses showed the association of Tomato leaf (Cucumis melo momordica) act as potential curl New Delhi virus. These viruses have also reservoirs of begomoviruses as they remain in been reported to infect Duranta species in the field for long. The Euphorbian Mosaic India57 as well as in Pakistan. Virus (EuMV) was found naturally infecting Perveen43 detected the presence of pepper in Mexico22. In Cuba, Begomoviruses begomoviruses in aksan, hulhul, lehli, itsit, also frequently infect and induce conspicuous bhakra, tandla, shoe flower (Hibiscus golden mosaics in weeds Rhyncosia minima rosasinensis), okra, brinjal and chilli using (family leguminosae) and Malvastrum triple antibody sandwich enzyme linked coromandelianum (family malvaceae). immunosorbent assay (TAS-ELISA). Begomovirus diversity is probably highest in Recently, Tiwari68 identified four Latin America and the the Caribbean, where different species of begomoviruses on the these viruses have many wild hosts belonging basis of cloning and sequencing of 800 bp to family malvaceae36. fragment in different cucurbit crops in UP. Based on highest (97–99%) sequence REFERENCES identities and closest phylogenetic 1. Ahmad J., Singh B R., Khedhairy A. A., relationships, four representative Begomovirus Alarifi S., Khan J.A and Musarrat. species were identified as Ageratum enation Characterization of Sun hemp begomo virus (from Trichosanthes dioica), Squash leaf virus and its geographical origin based on curl China virus (from Cucurbita maxima), in silico structural and functional analysis Tomato leaf curl New Delhi virus (from of recombinant coat protein. African Memordica charantia, Luffa cylindrica and Journal of Biotechnology 10: 260-270 Luffa acutangula) and Tomato leaf curl (2011). Palampur virus (from Cucurbita pepo). Their 2. Alegbejo M. D. Screening of pepper findings suggested the existence of a high cultivars for resistance to leaf curl virus. genetic diversity among begomoviruses Capsicum Eggplant Newsletter 18: 69-72 infecting cucurbitaceous crops. (1999). In China, Clerodendron cyrtophylum 3. Amin P. W. Leaf curl disease of chilli has been reported to be a new host for peppers in Maharashtra. PANS 25: 131-34 begomovirus. The sequence characterization (1979). studies revealed the association of a distinct 4. Anonymous. Post-harvest profile of chilli. begomovirus with C. cyrtophylum showing Government of India, Ministry of yellow mosaic type of symptoms. The new Agriculture (Department of Agriculture & virus has been given a name as Clerodendrum Cooperation) (2009). golden mosaic Jiangsu virus. 5. Arono, J., Gabarra, R., Liu, T., Alvin, M. Ahmad and Saeed1 investigated the and Dan G. Natural Enemies of Bemisia role of tomato crop as a potential host for tabaci Predators and Parasitoids. In: cotton leaf curl virus. Ahmad1 identified Stansly PA and Naranjo S E (Ed) Bemisia: sunhemp (Crotolaria juncea) as a new host for Bionomics and Management of a Global begomoviruses in SouthEast Asia. Haider24 Pest. pp: 385-42. Springer, the reported the association of begomovirus with Netherlands (2010). Vinca minor (Periwinkle) showing yellowing 6. Biswas, K., Hallan, V., Zaidi, A. A. and and leaf curl type of symptoms in Pakistan. Pandey, P. K. Molecular evidence of Gaikwad 16,17,18 reported Brinjal, Petunia cucumber mosaic virus subgroup II hybrida, Calendula and Marigold as a natural infecting Capsicum annuum L. in the host for the tomato leaf curl virus in India. The Western region of India. Current extensive host range has consequences for Discovery 2: 97-105 (2013). virus disease epidemiology and is worth 7. Brown, J K., Frohlich, D.R. and Rosell, considering while developing a system-wide R.C. The sweet potato/silverleaf management strategy for these viruses. whiteflies: biotypes of Bemisia tabaci or a

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