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Identification of Resistance to Bemisia Tabaci Genn. in Closely Related Wild Genet Resour Crop Evol (2017) 64:247–260 DOI 10.1007/s10722-015-0347-y RESEARCH ARTICLE Identification of resistance to Bemisia tabaci Genn. in closely related wild relatives of cultivated tomato based on trichome type analysis and choice and no-choice assays Mohamed Rakha . Peter Hanson . Srinivasan Ramasamy Received: 1 September 2015 / Accepted: 16 November 2015 / Published online: 7 December 2015 Ó The Author(s) 2015. This article is published with open access at Springerlink.com Abstract The sweetpotato whitefly, Bemisia tabaci in S. pimpinellifolium accession VI030462 based on Genn., is a major pest of tomato (Solanum lycoper- the no-choice bioassay. Whitefly resistance in sicum) and other crops throughout the tropics and VI037240 and VI030462 is noteworthy because these subtropics. The objectives of this study were to species are closely related to cultivated tomato and characterize 255 accessions of S. galapagense, S. introgression of whitefly resistance should be rela- cheesmaniae and S. pimpinellifolium for trichome tively straightforward. High densities of type IV types, and to evaluate selected accessions with high trichomes and low densities of type V trichomes were densities of glandular trichomes for resistance to associated with reduced numbers of whitefly adults, whitefly. Twenty-two accessions classified as either nymphs, puparium, and eggs in the choice bioassay sparse or abundant for type IV trichomes were selected and with high adult whitefly mortality in the no-choice and evaluated for numbers of adults, eggs, nymphs, bioassay. Preliminary trichome analysis followed by and puparium of whitefly in choice bioassays, for adult choice and no-choice assays facilitated rapid identifi- mortality and egg numbers in no-choice bioassays, and cation of whitefly-resistant accessions from a large for densities of type I, IV, V, and VI trichomes. The pool of candidates of different species. highest whitefly resistance was detected in S. galapa- gense accessions VI063177 and VI037239 based on Keywords Bemisia tabaci Á Solanum cheesmaniae Á choice and no-choice bioassays. In addition, we found S. galapagense Á S. pimpinellifolium Á Sweetpotato high levels of whitefly resistance in S. cheesmaniae whitefly Á Trichomes accession VI037240 based on the choice bioassay and Introduction Electronic supplementary material The online version of this article (doi:10.1007/s10722-015-0347-y) contains supple- mentary material, which is available to authorized users. Tomato (Solanum lycopersicum L.) is one of the most widely consumed vegetables worldwide, with a global M. Rakha (&) Á P. Hanson Á S. Ramasamy AVRDC, The World Vegetable Center, production of 162 million tons with a net value of more PO Box 42, Shanhua, Tainan 74199, Taiwan than $59 billion in 2012 (United Nations Food and e-mail: [email protected] Agriculture Organization (FAO 2012) statistics; http:// faostat.fao.org). The sweetpotato whitefly (Bemisia M. Rakha Horticulture Department, Faculty of Agriculture, tabaci Genn.) (family Aleyrodidae), also known as the University of Kafrelsheikh, Kafr El-Sheikh 33516, Egypt silverleaf whitefly (B. argentifolii Bellows et Perring), 123 248 Genet Resour Crop Evol (2017) 64:247–260 is a major pest of cultivated plants in tropical, sub- antibiosis (reduced insect fitness resulting from tropical and warm temperate regions. The most pre- reduced survival, oviposition, and development) (An- dominant and damaging biotypes are biotype B (also tonious et al. 2005; Bleeker et al. 2009, 2011, 2012; known as Middle East-Minor Asia 1 genetic group Freitas et al. 2002; Muigai et al. 2003). Glandular (MEAM1), hereafter referred to as B) and biotype Q trichomes also may act as a physical barrier, and (also known as the Mediterranean genetic group interfere with whitefly landing, feeding and oviposi- (MED), hereafter referred to as Q). These biotypes tion (Dimock and Kennedy 1983; Snyder and Carter have spread from their native ranges to as many as 60 1984; Channaryappa et al. 1992). countries (Pan et al. 2013) causing annual losses of $1 Recent studies of whitefly resistance in the wild to 2 billion (Jiang et al. 2012). Whitefly feed on foliar tomato species S. galapagense S.C. Darwin et Peralta phloem and produce a sticky secretion called honey- (formerly Lycopersicon cheesmaniae f. minor (Hook. dew, a substrate for fungi that results in black sooty f.) C.H. Mull.) reported diverse resistance levels mold on the leaf surface and reduces leaf photosyn- between and within accessions (Firdaus et al. 2012; thetic efficiency; whitefly saliva may also cause Lucatti et al. 2013). Spider mite and whitefly resis- irregular fruit ripening, which reduces fruit quality and tance has been reported in S. pimpinellifolium L. increases the number of unmarketable fruit (Schuster (formerly L. pimpinellifolium (Juslen.) Mill.) acces- et al. 1996). Most importantly, 114 virus species sion TO937 (Alba et al. 2009; Rodrı´guez-Lo´pez et al. belonging to five different genera (Begomovirus, 2011). Solanum galapagense, along with S. cheesma- Crinivirus, Closterovirus, Ipomovirus and Carlavirus) niae (L. Riley) Fosberg (syn L. cheesmaniae L. Riley) are transmitted by whitefly (Jones 2003). Bego- and S. pimpinellifolium are closely related to culti- moviruses that cause tomato yellow leaf curl disease vated tomato and introgression of resistance from are the most numerous of the B. tabaci-transmitted these species may be easier and faster compared to the viruses and early tomato yellow leaf curl infection can green-fruited species S. chilense (Dunal) Reiche cause 100 % yield loss (Saikia and Muniyappa 1989). (formerly L. chilense Dun.), S. peruvianum L. Current whitefly control strategies rely primarily on (formerly L. peruvianum (L.) Mill.), S. pennellii insecticides to which whitefly are notorious for Correll (formerly L. pennellii (Corr.) D’Arcy), and S. developing resistance (Denholm et al. 1996; Palumbo habrochaites S. Knapp and D.M. Spooner (formerly L. et al. 2001). Pesticide control of whitefly is often hirsutum Humb. and Bonpl.) (Peralta et al. 2008; Rick costly and can be hazardous to farmers and the 1971). To date only a limited number of accessions/ environment, including natural enemies. Therefore, populations of S. galapagense, S. cheesmaniae and S. availability of whitefly-resistant cultivars could con- pimpinellifolium have been characterized for trichome tribute to cost-effective and environmentally sound types and evaluated for insect resistance. Screening for pest management. insect resistance can be laborious, especially for large Seven trichomes types have been identified in numbers of accessions and plant populations (Bas cultivated tomato and its wild relatives. Trichomes are et al. 1992). Availability of high throughput methods categorized into non-glandular types without droplets to assess efficiently and accurately large numbers of (Types II, III and V) and glandular types with droplets tomato accessions/plants for insect resistance would (Types I, IV, VI and VII). Trichomes in cultivated facilitate identification of resistance sources and tomato are mostly non-glandular type V, sometimes improve breeding. Information on insect resistance with sparse densities of glandular types I and VI, levels among a large number of accessions and their whereas some wild relatives of tomato have an underlying resistance mechanisms would be very abundance of glandular trichomes, especially types I, useful for tomato breeders worldwide. Based on the IV and VI (Gurr and McGrath 2001; McDowell et al. strong association between insect resistance and 2011; Simmons and Gurr 2005). Glandular trichomes presence of glandular trichomes, the objectives of this play an important role in whitefly resistance through study were to characterize all available S. galapa- release of defense compounds such as acyl sugars, gense, S. cheesmaniae and S. pimpinellifolium acces- methylketones, and sesquiterpenes; these compounds sions in AVRDC – The World Vegetable Center’s may cause antixenosis (insect behavior changes genebank for trichome types, and to evaluate the resulting in reduced or no host colonization) and/or resistance to whitefly of selected accessions with high 123 Genet Resour Crop Evol (2017) 64:247–260 249 densities of glandular trichomes in choice and no- sparse for type IV trichomes and selected for whitefly choice bioassays. choice and no-choice bioassays to determine resis- tance. The 22 accessions included four S. cheesma- niae,11S. galapagense and seven S. pimpinellifolium. Materials and methods Five weeks after sowing, 10 seedlings per accession and 20 seedlings of the susceptible check were Plant materials and growth conditions transplanted into 12 cm2 pots with potting soil and moved from the plastic house to growth rooms. The Seeds of 260 accessions of S. galapagense, S. chees- growth room temperature was increased slowly (one maniae and S. pimpinellifolium were obtained from degree per day) from 23 to 27 °C to allow plants to the AVRDC genebank (Supplementary Table 1). adjust to the higher optimal temperatures AVRDC tomato line CL5915-93D4-1-0-3 was (27 ± 2 °C) and conditions (70 % RH, 16/8 h day/ included as a whitefly-susceptible check. Five acces- night) for whitefly. Plants were fertilized weekly with sions were discarded due to low germination. The NPK 15-15-15 and were watered daily. Whitefly (B. accessions with enough viable seeds were divided into tabaci, biotype B) used in choice and no-choice assays two groups of 130 accessions and groups 1 and 2 were were collected from colonies originating from an sown on 11 and 28 November 2014, respectively. AVRDC field and reared and maintained on cabbage Seeds were sown in 72-plug seedling trays with 40 ml plants (Brassica oleracea L., a non-tomato yellow leaf peat moss per cell. The accessions and check were curl-susceptible host) in muslin-covered cages in a grown in an AVRDC greenhouse (26 ± 4 °C, growth chamber at 23–30 °C. Six weeks after sowing, 6/18 h day/night) and fertilized weekly. each plant was infested with 10 and 5 whitefly pairs for choice and no-choice bioassays, respectively. The Analysis of trichome morphology and density experiments were conducted from November 2014 to February 2015 in an AVRDC greenhouses.
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