Trichomes of Lycopersicon spp. and their effect on Myzus persicae (Sulzer) (Hemiptera: Aphididae)

Abstract The green peach aphid ( Myzus persicae ) is becoming resistant to many of the pesticides used in the tomato ( Lycopersicon esculentum ) industry. A potential alternative method of control is host-plant resistance, which may be mediated by trichomes. Resistance to the green peach aphid was assessed for six accessions of L. hirsutum, three of L. pennellii and a L. esculentum control. Mortality, entrapment and number of aphids unaffected by trichomes were recorded 24, 48 and 72 h after placement. Using multiple regression against counts of glandular and non-glandular trichomes, the role of foliar pubescence in resistance to the aphid was established. Lycopersicon pennellii possessed the highest level of resistance after 72 h and type IV trichome density accounted for 26.5% of variance in mortality ( P  < 0.001). Low densities of type IV and high densities of type VII trichomes increased the number of aphids 'unaffected' by trichomes, a relationship accounting for 29% of variance ( P  < 0.001). Information on which types of trichomes most strongly affect resistance to insect pests will aid future breeding and L. pennellii shows potential for use in breeding programs to increase levels of host-plant resistance in L. esculentum.     

Trichome‐based host plant resistance of Lycopersicon species and the biocontrol agent Mallada signata: are they compatible?

Trichome‐based host plant resistance of Lycopersicon (Solanaceae) species offers the potential to reduce pesticide use in tomato production, but its compatibility with biocontrol agents is largely unknown. The effect of trichome‐based host plant resistance on the lacewing biocontrol agent, Mallada signata, was assessed for five accessions of L. cheesmanii, four accessions of Lycopersicon hirsutum, two accessions of Lycopersicon pennellii, and one Lycopersicon esculentum cultivar. An intact leaf was isolated from the whole plant using Tangletrap to coat the petiole and 20 green peach aphids [Myzus persicae (Sulzer) (Homoptera: Aphididae)] were placed on the leaf surface. After 24 h, 10 lacewings were placed on the leaf. The numbers of dead, trapped by exudates, untrapped and predated lacewings and aphids, and the numbers that had left the leaf were recorded a further 24 h later. Differences in insect designations between accessions were analysed using ANOVA. A General Linear Model (GLM), consisting of the densities of each trichome type and leaf area, was fitted to the data to determine the role of trichomes on the observed effects on lacewings and aphids. Lacewing mortality was greater on one accession of L. pennellii and one accession of L. hirsutum than on L. esculentum. The GLM indicated that type IV trichomes decreased the numbers of aphids predated, and increased cannibalism and, along with type III trichomes, increased entrapment‐related predator mortality. Although there were no differences in the numbers of predated aphids, with the majority predated for all accessions, the compatibility of trichome‐based host plant resistance of Lycopersicon species and the biocontrol of aphids by lacewings is questionable.     

Influence of Canopy Height and Fertilization Levels on the Resistance of Lycopersicon Hirsutum to Aculops Lycopersici (Acari: Eriophyidae)

The objective of this work was to study the effect of NK fertilization levels and canopy height on the resistance of Lycopersicon hirsutum and Lycopersicon esculentum to Aculops lycopersici (Acari: Eriophydae). The effects of NK fertilization levels and canopy height in the leaf size and density of trichomes and their effects on tridecan-2-one (2-TD) and undecan-2-one (2-UD) limiting the attack of A. lycopersici on tomato plants were assessed. Different NK fertilization levels had no effect on the resistance of L. hirsutum to A. lycopersici. No significant differences were found in attack rates of this mite on leaves of the top and median parts of L. hirsutum canopy. The type and density of trichomes were the main determining factor of A. lycopersici attack on tomato plants. High trichome densities and type VI glandular trichomes which produce tridecan-2-one are important resistance factors on tomato plants. L. hirsutum showed a high resistance level to A. lycopersici due to high densities of type VI glandular trichomes and consequently higher levels of tridecan-2-one in its leaves.     

Effect of fertilization levels, age and canopy height of Lycopersicon hirsutum on the resistance to Myzus persicae

The effects of different levels of fertilization with nitrogen and potassium (NK), height within the canopy and plant age of Lycopersicon hirsutum f. glabratum (PI 134417) C. H. Mull and L. esculentum on the density of foliar cristalliferous idioblasts and trichomes, and on the levels of 2- tridecanone (2-TD) and 2-undecanone (2-UD) and their influence on the resistance to Myzus persicae (Sulzer) were investigated. NK levels were inversely related to densities of trichomes and cristalliferous idioblasts in L. esculentum, but only to densities of cristalliferous idioblasts in L. hirsutum. Trichome density increased with increasing height within the canopy and with plant age in L. esculentum and L. hirsutum. Density of cristalliferous idioblasts in L. hirsutum increased with canopy height reaching a maximum at mid-height within the canopy in opposition to L. esculentum in which a small decrease in density with height within the canopy was found. There was also a small decrease of idioblast density with increasing plant age for L. esculentum, while a greater and opposite variation took place in L. hirsutum. L. esculentum has mainly non- glandular leaf trichomes (90%), while L. hirsutum has mainly glandular trichomes (97%). The main glandular trichomes of L. hirsutum are producers of 2-TD and 2-UD, whose leaf content increases with increase of trichome density. A negative effect of 2-TD concentration and cristalliferous idioblast density on the survival and longevity of M. persicae was observed. Since L. hirsutum presented much higher levels of 2-TD and cristalliferous idioblast density than L. esculentum, L. hirsutum seems a promising source of resistance to M. persicae for tomato breeding programs.     

Foliar pubescence and resistance to potato moth, Phthorimaea operculella, in Lycopersicon hirsutum

Pubescence characteristics for six accessions of Lycopersicon hirsutum Dunal and five accessions of L. hirsutum f. glabratum CH Mull. were determined and compared with those of an accession of cultivated tomato (L. esculentum Mill.). Removal of trichome exudates from excised leaflets using ethanol solution resulted in a reduced mortality and increased survival of potato moth (Phthorimaea operculella (Zeller)) neonates for the accessions that were most lethal when not treated with ethanol solution. No such treatment effect was evident for L. esculentum or for the L. hirsutum accession with least effect on neonates when its trichomes were intact. In a glasshouse experiment with caged intact plants, mortality of neonate P. operculella placed on the abaxial surface was greater on seven accessions than for L. esculentum.Neonates were less severely affected on the adaxial surface. Eleven days after inoculation, no live larvae were found on LA 1927, PI 127827, PI 134418, and PI 134428, and numbers on other accessions were lower than for L. esculentum. Eventual emergence of adults followed a similar trend. Multiple regression of insect data against pubescence indicated a significant correlation between density of type IV and VI trichomes and neonate mortality, decreased larval development and decreased adult emergence. Non-glandular type V trichomes were positively correlated with high survival of insects to 11 days and to adult. Though factors other than glandular trichomes are likely to be important, increased density of type IV and VI, along with reduced type V, are shown to be important to select in breeding for P. operculella resistance.     

Toxicity of methyl ketones from tomato trichomes to Tetranychus urticae Koch

The two methyl ketones, 2-tridecanone and 2-undecanone, are constituents of type VI glandular trichomes of the wild tomato, Lycopersicon hirsutum f. glabratum (PI 134417). They are known to cause mortality in several herbivorous insect species. In this study we investigated the effects of these chemicals on two strains of the two-spotted spider mite, Tetranychus urticae Koch, collected from tomato and cucumber crops in Dutch greenhouses. The two ketones were tested separately, in combination in the ratio found in L. hirsutum f. glabratum and in several other ratios to detect any synergistic interaction between them. We measured both the direct contact and residual toxicity, as well as the viability of the eggs produced by ketone-treated females. The methyl ketones can be considered as quite toxic to spider mites because their LC50 values were comparable to that found for the formulated acaricide amitraz. 2-Tridecanone was slightly more toxic than 2-undecanone but only in the tomato strain. Synergistic effects were not detected. In the bio assays for the residual effects of the two methyl ketones no significant mortality occurred but the mites avoided feeding on the treated surface and the eggs were laid almost exclusively on the untreated area. Furthermore, there was no significant egg viability for most of the treatments and when some egg viability occurred, it was not correlated to the dose of the chemical. Based on published data on the levels of methyl ketones in the trichome tips of wild and cultivated tomato (L. hirsutum f. glabratum and Lycopersicon esculentum Mill, respectively) it was calculated how many contacts are required to reach the LD50 (ng a.i. per mite). It appeared that on wild tomato just a few contacts suffice, but many are needed on cultivated tomato. Nevertheless, it seems justified to conclude that even in cultivated tomato, 2-tridecanone represents a naturally occurring acaricide. The cucumber and tomato strains of the two-spotted spider mite did not differ in their response to the methyl ketones. Because this is surprising, given the selection pressures on their respective host plants, we formulated two hypotheses, one explaining the absence of a difference due to cross-resistance and another one proposing that bioaccumulation obscures the underlying differences between the strains. This revised version was published online in November 2006 with corrections to the Cover Date.     

Leaf volatiles from some wild tomato species

The investigated species are Lycopersicon cheesmanii s.str., L. cheesmanii var. minor, L. chilense, L. hirsutum s. str., L. hirsuturn f. glabratum , L. "parviflorum", L. chmielewskii, L. peruvianum s. str., L. peruvianum var. humifusum, L. pimpinellifolium and Solanum penellii . Wound-emitted leaf volatiles isolated and concentrated by adsorption on Tenax GC were separated by capillary gas chromatography. The different species produce different and rather broad patterns of volatiles especially compared to a modem tomato cultivar. There is also a considerable difference between varieties of the same species. When more than one accession was investigated, the results indicated a chemotype differentiation. The chemotypes are in some cases specialized with one or a few quantitatively dominating main components. L. hirsutum emits the greatest number of components. Three accessions had α-zingeberene, α-santalene, and limonene as main component, respectively. The investigated accession of L. hirsutum f. glabratum has 2-undecanone as main component, instead of 2-tride-canone previously identified as main component in another accession. Both compounds are natural insecticides. The results illustrate the need for basic knowledge of the chemical characters in the ancestors of cultivated plants.     

QTL analysis of quantitative resistance to Phytophthora infestans (late blight) in tomato and comparisons with potato.

Quantitative trait loci (QTLs) for resistance to Phytophthora infestans (late blight) were mapped in tomato. Reciprocal backcross populations derived from cultivated Lycopersicon esculentum x wild Lycopersicon hirsutum (BC-E, backcross to L. esculentum; BC-H, backcross to L. hirsutum) were phenotyped in three types of replicated disease assays (detached-leaflet, whole-plant, and field). Linkage maps were constructed for each BC population with RFLPs. Resistance QTLs were identified on all 12 tomato chromosomes using composite interval mapping. Six QTLs in BC-E (lb1a, lb2a, lb3, lb4, lb5b, and lb11b) and two QTLs in BC-H (lb5ab and lb6ab) were most consistently detected in replicated experiments or across assay methods. Lycopersicon hirsutum alleles conferred resistance at all QTLs except lb2a. Resistance QTLs coincided with QTLs for inoculum droplet dispersal on leaves, a trait in L. hirsutum that may contribute to resistance, and dispersal was mainly associated with leaf resistance. Some P. infestans resistance QTLs detected in tomato coincided with chromosomal locations of previously mapped R genes and QTLs for resistance to P. infestans in potato, suggesting functional conservation of resistance within the Solanaceae.     

Susceptibility of Tuta absoluta (Meyrick) (Lep., Gelechiidae) reared on two species of Lycopersicon to Bacillus thuringiensis var. kurstaki

The mortality of Tuta absoluta (Meyrick) larvae that were fed on leaves of Lycopersicon hirsutum f. glabratum (PI 134417, insect resistant) and Lycopersicon esculentum (cultivar Santa Clara, susceptible) treated with Bacillus thuringiensis var. kurstaki (Btk) was evaluated. Feeding on untreated PI 134417 was detrimental to the survival of T. absoluta larvae. When Btk was applied to the two Lycopersicon plants, mortality occurred in all T. absoluta instars. Application of Btk on tomato leaves had synergistic or additive effects with the resistant genotype on larval survival. This effect was dependent on the instar at which the larvae were fed Btk-treated leaves. Delayed Btk application may cause higher insect mortality if the insects become more susceptible to the pathogen after a longer period of feeding on the resistant crop.     

Host plant resistance among tomato accessions to the spider mite <Emphasis Type="Italic">Tetranychus evansi</Emphasis> in Kenya

The spider mite Tetranychus evansi has a broad range of host plants. Control of T. evansi has been a big challenge to tomato farmers due to its fast rate of reproduction, development of resistance to chemical pesticides and its ability to use weeds as alternative hosts when the tomato plants are not available. The aim of the current study was to determine the host plant acceptance and the relative contributions of trichomes in the control of the red spider mite by comparing the survival, development and oviposition rates of the red spider mite on eight tomato accessions. Leaflets from eight tomato varieties were assayed with the spider mites to determine the egg laying capacity and developmental time of the spider mites on the tomato accessions as well as the trichome densities. Densities of trichome types I, IV, V and VI varied among the tomato accessions. Variation in types I, IV and VI accounted for most of the variation in mite responses. The varieties with high densities of types IV and VI had the highest fecundity and mite development did not go beyond the larval stage. The developmental time varied significantly among the tomato accessions. The results indicated that the higher the density of trichome type I the lower the adult survival. The findings indicated possible resistance of some of the tested tomato accessions against T. evansi which is partially associated with trichomes types and density.     

Role of canopy height in the resistance of Lycopersicon hirsutum f. glabratum to Tuta absoluta (Lep., Gelechiidae)

The objective of this work was to study the effect of plant canopy height on the resistance of the tomato species Lycopersicon hirsutum to the tomato leafminer Tuta absoluta . Determination of the levels of tridecan-2-one (2-TD) and undecan-2-one (2-UD) in L. hirsutum , as well as the leaf area, density and types of trichomes present in L. hirsutum and L. esculentum were made on apical, medium and basal parts of plant canopy. Correlation of these data were made with the following biological characteristics of T. absoluta : rates of oviposition and egg hatching; length of egg, larval and pupal stages; mortality of the larval and pupal stages; pupal weight; and sex ratio. Levels of 2-TD on leaves of L. hirsutum increased from bottom to top of canopy. Tuta absoluta had higher oviposition and egg hatching, as well as lower mortalities and smaller larval and pupal periods in L. esculentum than in L. hirsutum . Longer pupal period and greater number of small and large mines of T. absoluta were verified in the apical and medium parts of L. hirsutum plants. On the other hand, higher pupal mortality and longer larval periods were obtained in the apical and basal parts of the L. hirsutum plants. Larval mortality was higher in the basal part of those plants. In L. esculentum , leaves from the apical part of the plant promoted lower mortality, longer larval period and greater number of large mines of T. absoluta . However, a shorter pupal period was verified on leaves of the basal part. It appears that an increase in glandular trichome density in L. hirsutum leads to an increase in the levels of 2-TD, which, in turn slows down larval development.     

Inhibition of Telenomus sphingis an egg parasitoid of Manduca spp. by trichome/2-tridecanone-based host plant resistance in tomato

The resistance of accession PI 134417 of the wild tomato Lycopersicon hirsutum f. glabratum C. H. Mull to Manduca sexta (L.) (Lepidoptera: Spingidae) and Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) is conditioned by the high densities of 2-tridecanone-containing, glandular trichomes associated with the foliage. In laboratory experiments, rates of parasitism of M. sexta eggs by Telenomus sphingis (Ashmead) (Hymenoptera: Scelionidae) were lower among eggs on PI 134417 foliage than among eggs on foliage of the cultivated tomato L. esculentum Mill. (cv. Better Boy). The latter is characterized by a significantly lower density of type VI glandular trichomes than PI 134417 and an absence of 2-tridecanone. Parasitism by T. sphingis was also reduced among eggs on foliage of the F₁ hybrid between PI 134417 and L. esculentum. The hybrid foliage lacks 2-tridecanone but has a density of type VI glandular trichomes that is intermediate between those of PI 134417 and L. esculentum, indicating that elevated densities of type VI glandular trichomes adversely affect T. sphingis. This conclusion was further substantiated by the finding that there were no differences among plant lines in the levels of parasitism of M. sexta eggs when the eggs were on foliage that had been divested of glandular trichomes.In bioassays in which T. sphingis adults or immatures in host eggs were exposed to filter paper treated with 2-tridecanone at rates comparable to those associated with PI 134417 foliage, 2-tridecanone was acutely toxic and caused high levels of mortality. In addition, at high concentrations, 2-tridecanone vapors were repellent to T. sphingis adults. However, when exposed to PI 134417 foliage, few T. sphingis adults were killed.Parasitism of M. sexta eggs was unaffected when the eggs were deposited by moths reared as fifth instar larvae on diet containing 2-tridecanone and/or 2-undecanone at levels comparable to those associated with PI 134417 foliage.     

Sesquiterpene carboxylic acids from a wild tomato species affect larval feeding behavior and survival of Helicoverpa zea and Spodoptera exigua (Lepidoptera: Noctuidae).

The sesquiterpene carboxylic acids (SCA), (+)-(E)-alpha-santalen-12-oic, (-)-(E)-endo-alpha-bergamoten-12-oic, and (+)-(E)-endo-beta-bergamoten-12-oic acid, are produced in glandular trichomes of Lycopersicon hirsutum f. typicum Humb. & Bonpl. accession (LA) 1777, which is highly resistant to a range of pests of cultivated tomatoes. L. esculentum Mill. Exposure of the larvae of two key tomato pests, tomato fruitworm [Helicoverpa zea (Boddie)] and beet armyworm [Spodoptera exigua (Hübner)], to these compounds in diets and on leaf surfaces resulted in reduced development rates and survival and deterred feeding. These effects were observed when levels of SCA, in artificial insect diet, applied to leaflets of susceptible cultivars, or synthesized in trichomes of leaves of plants, exceeded 2 mg SCA/g of diet or fresh leaf weight. This study suggests that cultivated tomatoes capable of synthesizing SCA, at 2 mg SCA/g of leaf tissue or greater, on their leaves and fruit would display enhanced host plant resistance to H. zea and S. exigua and other insect pests.     

Variation in Response of Wild Lycopersicon and Solanum spp. against Tomato Powdery Mildew (Oidium lycopersici)

A set of 154 accessions of nine wild Lycopersicon spp. and five accessions of three closely related Solanum spp. were tested for resistance to tomato powdery mildew ( Oidium lycopersici ). Screening revealed valuable sources of resistance, mainly among L. hirsutum, L. pennellii, L. cheesmanii, L. chilense, L. peruvianum and L. parviflorum. L. esculentum (all ssp.) and L. pimpinellifolium expressed high susceptibility to O. lycopersici inoculation. Results of variance and cluster analysis of responses to O. lycopersici coincide with recent taxonomic classification and genetic relationships within genus Lycopersicon .     

Effect of cropping season and possible compounds involved in the resistance of Lycopersicon hirsutum f. typicum to Tuta absoluta (Meyrick) (Lep., Gelechiidae)

The effect of cropping season on the resistance of the accession LA 1777 of Lycopersicon hirsutum f. typicum to the tomato leafminer Tuta absoluta (Meyrick) and the preliminary identification of the possible compounds involved in this trait were the objectives of this study. Two experiments were carried out with the accession LA1777 and the commercial cultivar Santa Clara of Lycopersicon esculentum . The first experiment was carried out during the summer season (January to April 1997) and the second was carried out during the autumn–winter season (May to September 1997). The accession LA 1777 was significantly more resistant to T. absoluta than the commercial cultivar, especially during the autumn–winter cropping season. Either antibiosis or antixenosis was probably responsible for the higher larvae mortality (54.2 and 91.2% for the summer and autumn–winter seasons, respectively) and longer larval development of T. absoluta (14.2 and 25.9 days for summer and autumn–winter seasons, respectively) on L. hirsutum f. typicum as compared with L. esculentum (11.0 and 19.6 days for summer and autumn–winter seasons, respectively). Insect fecundity parameters such as oviposition period, number of eggs/female, and rate of larvae eclosion were also negatively affected by L. hirsutum f. typicum . Fourteen main peaks were observed in total ion chromatograms of both tomato species. Eleven of these were present only in chromatograms of the accession, one was only present in chromatograms of the commercial cultivar, and two peaks were present in chromatograms from both when hexane extracts of leaves were subjected to a gas chromatographic–mass spectrometric analysis (GC–MS). However, the matches obtained between the peak spectra and the MS database were too low to justify even preliminary identification of the compounds.     

Local and systemic production of nitric oxide in tomato responses to powdery mildew infection

Various genetic and physiological aspects of resistance of Lycopersicon spp. to Oidium neolycopersici have been reported, but limited information is available on the molecular background of the plant–pathogen interaction. This article reports the changes in nitric oxide (NO) production in three Lycopersicon spp. genotypes which show different levels of resistance to tomato powdery mildew. NO production was determined in plant leaf extracts of L. esculentum cv. Amateur (susceptible), L. chmielewskii (moderately resistant) and L. hirsutum f. glabratum (highly resistant) by the oxyhaemoglobin method during 216 h post-inoculation. A specific, two-phase increase in NO production was observed in the extracts of infected leaves of moderately and highly resistant genotypes. Moreover, transmission of a systemic response throughout the plant was observed as an increase in NO production within tissues of uninoculated leaves. The results suggest that arginine-dependent enzyme activity was probably the main source of NO in tomato tissues, which was inhibited by competitive reversible and irreversible inhibitors of animal NO synthase, but not by a plant nitrate reductase inhibitor. In resistant tomato genotypes, increased NO production was localized in infected tissues by confocal laser scanning microscopy using the fluorescent probe 4-amino-5-methylamino-2',7'-difluorofluorescein diacetate. NO production observed in the extracts from pathogen conidia, together with elevated NO production localized in developing pathogen hyphae, demonstrates a complex role of NO in plant–pathogen interactions. Our results are discussed with regard to a possible role of increased NO production in pathogens during pathogenesis, as well as local and systemic plant defence mechanisms.     

Aspects of potato leafhopper (Homoptera: Cicadellidae) biology on Solanum berthaultii and other potato genotypes

Survival of nymphs and adult males and females of potato leafhopper, Empoasca fabae (Harris), on six potato (Solanum spp.) genotypes with varied levels of resistance was evaluated and related to type, density, and droplet/head size of foliar glandular trichomes. Survival was greater on Solanum tuberosum cultivars ('Allegany' and 'Elba') compared with accessions of the wild S. berthaultii (PI 473331 and PI 473334). The barrier provided by glandular trichomes may affect survival of E.fabae by restricting nymph and adult feeding. PI 473331 was the most pubescent host, bearing type A and type B trichomes and the most unsuitable host for E. fabae. Allegany possesses leaflets with low trichome densities and small gland diameter and was the most suitable host for adult E. fabae survival. Females generally had higher survival rates than males. Nymphal and adult leafhoppers differed greatly in their survival rates, particularly on S. tuberosum x S. berthaultii hybrids (Q174-2 and NY123). Interestingly, nymphs had longer (190 d) mean lethal time (LT50) than adults (31 d) when held on Q174-2, and a lower LT50 (19 d) when held on NY123 compared with adults (104 d). Starvation, dehydration, or both were likely associated with mortality of nymphs confined on PI 473331 and PI 473334. Nymphs on these accessions died at a comparable rate to those confined on a starvation treatment and more rapidly than those supplied with water only, sucrose solution or other potato genotypes. Darkened trichome exudates were observed on insects held on PI 473331 and PI 473334, consistent with previous reports that glandular trichomes of S. berthaultii contain phenolic oxidation chemistry. This study aids in the identification of the probable mechanisms of resistance to E. fabae used by S. berthaulti and its hybrids with S. tuberosum.     

Genetic transfer of a twospotted spider mite (Acari: Tetranychidae) repellent in tomato hybrids

Lycopersicon hirsutum Dunal is very resistant to arthropod herbivory, and research on causes of resistance has often implicated trichomes and their secretions. To better understand relationships among resistance, repellency, and 2,3-dihydrofarnesoic acid, a trichome-borne sesquiterpenoid spider mite repellent, two tomato, Lycopersicon esculentum Miller, varieties were interbred with a highly resistant, spider mite repellent accession (LA1363) of L. hirsutum. Backcross and F2 generations were produced with each tomato variety. Whole leaves of 99 hybrids were bioassayed with twospotted spider mites, Tetranychus urticae Koch, allowing selection of six hybrids (two susceptible and four resistant) for each generation of each family. When these 24 hybrids were characterized for spider mite repellency with thumbtack bioassays, two hybrids had repellent leaflets, demonstrating that repellency was genetically transferred to interspecific tomato hybrids. Leaflet washes containing trichome secretions from each of three hybrids, including the two having repellent leaflets, were repellent in bridge bioassays. For the two hybrids having repellent leaflets and leaflet washes, removal of trichome secretions by dipping leaflets in methanol eliminated leaflet repellency. 2,3-Dihydrofarnesoic acid was present in trichome secretions of the hybrids having leaflet repellency, and it also was present in secretions of other hybrids, indicating that its presence is essential, but not sufficient for leaflet repellency. With regard to resistance, 16 of the hybrids tested had been identified as resistant in a whole leaf bioassay, but only two had repellent leaflets, indicating that other mechanisms of resistance are present in the resistant L. hirsutum parent.     

Role of the glandular trichomes in resistance of perennial alfalfa to the potato leafhopper (Homoptera: Cicadellidae)

Experiments were performed to elucidate resistance of glandular-haired alfalfa, Medicago sativa L., to the potato leafhopper, Empoasca fabae (Harris). The primary objective was to examine the potential role of the glandular trichomes and stem lignification for imparting resistance to this pest. During free-choice foliar discoloration experiments, the resistant alfalfa clone FGplh13 expressed lower levels of injury than the susceptible P5373 after 13 d. No-choice tests performed using nymphal potato leafhoppers showed higher levels of mortality associated with the resistant glandular-haired clone FGplh13, and no nymphs survived after 48 h. Additional no-choice experiments using FGplh13 and P5373 alfalfa with the glandular and nonglandular trichomes intact or removed show that mortality of nymphs and adults decreased after removal of the glandular trichomes from FGplh13. Nymphal and adult mortality and nymphal development time did not differ on FGplh13 with the trichomes removed and P5373 with the trichomes intact or removed. Firstinstar, and possibly second-instar, potato leafhoppers were entrapped by an exudate produced by the glandular trichomes on the FGplh13 alfalfa. Stem lignification and the number of vascular bundles did not differ between FGplh13 and P5373 alfalfas. The glandular trichomes on FGplh13 alfalfa appeared to provide the major host resistance factor, with resistance to adults being chemically based and resistance to nymphs being chemically and mechanically based. To maintain levels of potato leafhopper resistance, breeders would appear to benefit by continuing to select for the expression of the glandular trichome phenotype.