Glandular trichomes ofSolanum berthaultii alter host preference of the Colorado potato Beetle,Leptinotarsa decemlineata

Choice and no-choice studies were conducted to determine how the glandular trichomes of the wild potato,Solanum berthaultii Hawkes, affect host preference of the Colorado potato beetle,Leptinotarsa decemlineata (Say). Given a feeding choice betweenS. tuberosum andS. berthaultii, larvae and adults preferred the foliage ofS. tuberosum, but adults were more discriminating. When foliage ofS. berthaultii was appressed toS. tuberosum leaflets, fewer adults fed on the appressed leaflets. When given a choice between ‘trichome-intact’ and ‘trichome-removed’S. berthaultii foliage, adults preferred to feed on the latter. The preference for ‘trichome-removed’ foliage and the percent of adults initiating feeding, increased with the degree of trichome removal. These studies provide evidence that the resistance ofS. berthaultii is associated with feeding deterrents localized in the glandular trichomes, thatS. berthaultii possesses more than one mechanism of resistance to the Colorado potato beetle, and that the expression of resistance is dependent on the developmental stage of the insect.     

Glandular trichomes of Solanum berthaultii and resistance to the Colorado potato beetle

The defensive mechanisms of the wild potato, solanum berthaultii Hawkes, to larvae of the Colorado potato beetle, Leptinotarsa decemlineata (Say), were studied by selective removal of glandular trichomes and trichome exudates from leaflets, and by comparing performance on S. berthaultii and on the cultivated potato, S. tuberosum L., which lacks defensively active type A and B glandular trichomes. Removal of type A trichomes increased the proportion of larvae that fed on S. berthaultii. Removal of the exudate from type B trichomes increased the proportion of larvae that fed and led to a decrease in mortality. The predominant active compounds in type B exudate, i.e. fatty acid esters of sucrose, were only effective in the presence of type A trichomes. Sucrose esters did not affect larval feeding on S. tuberosum leaflets or on S. berthaultii leaf discs from which the type A trichomes had been removed. Growth of surviving larvae was not significantly affected by removing type A trichomes or type B exudate. Growth of larvae was significantly increased when S. berthaultii leaflets were presented in artificial diet which eliminated the physical barrier of the type B stalks. Growth was no different on artificial diet containing either S. berthaultii or S. tuberosum leaf material (fresh or lyophilized powder) but was poorer on these diets than on S. tuberosum leaflets. The presence of type A trichomes is a fundamental requirement for expression of S. berthaultii resistance to L1 L. decemlineata. Type B droplets containing sucrose esters increase the expression of resistance in the presence of defensively-active type A trichomes.

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Résumé Les mécanismes de défense de la pomme de terre sauvage, S. berthaultii Hawkes, aux larves de Leptinotarsa decemlineata Say, ont été étudiés par ablation sélective des trichomes glandulaires et par l'élimination de leur exsudat des folioles, et par comparaison avec S. tuberosum L. qui a perdu les trichomes glandulaires défensifs A et B. L'ablation des trichomes A a augmenté la proportion de larves ayant consommé S. berthaultii. L'élimination de l'exsudat des trichomes B a augmenté la proportion de consommatrices et réduit la mortalité. Les principaux composés actifs de l'exsudat B, c'est-à-dire des esters d'acides gras de sucrose, n'étaient actifs qu'en présence de trichomes A. Les esters de sucrose n'ont pas modifié la consommation larvaire sur folioles de S. tuberosum, ou sur disques de feuilles de S. berthaultii dont les trichomes A avaient été enlevés. La croissance des larves survivantes n'a pas été modifiée significativement par l'ablation des trichomes A ou l'élimination de l'exsudat de B. La croissance des larves a été significativement augmentée quand les folioles de S. berthaultii ont été incorporés dans l'aliment artificiel après élimination de la barrière physique due aux pédoncules B. La croissance a été de même importance sur aliments artificiels contenant des feuilles (fraiches ou en poudre lyophylisée) de S. berthaultii ou de S. tuberosum, mais plus faible que sur folioles de S. tuberosum. La présence de trichomes A est indispensable à la résistance de S. berthaultii aux L, de L. decemlineata. Les gouttelettes de type B contenant des esters de sucrose augmentent l'expression de la résistance en présence d'une défense active par trichomes A.

     

Variable responses of tuber moth to the leaf trichomes of wild potatoes

This study examines the response of tuber moth, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae), during the initial stages of attack, to variability in trichome density and composition on foliage of Solanum berthaultii (Hawkes) and Solanum tarijense (Hawkes) (Solanaceae). Solanum berthaultii bears two types of glandular trichome (type A and type B) that together reduced oviposition by the moth. Females were often completely deterred from ovipositing on foliage with >300 trichomes per cm². In contrast, neonate establishment on S. berthaultii was generally positively related to trichome densities, indicating that trichomes may be a poor defense against P. operculella when the moth oviposits in soil and neonate larvae select the host plant. Solanum tarijense has only one type of glandular trichome (type A) and eglandular hairs. Most eggs were deposited on the adaxial leaf surfaces that had lower trichome densities. Although the density of type A trichomes was negatively related to oviposition, high densities of hairs on the abaxial and adaxial leaf surfaces appeared to stimulate oviposition, leading to stronger positive relations between hair densities and oviposition. Larvae generally established on the abaxial surface where hair densities were greatest. Relationships between the abaxial densities of leaf hairs and neonate establishment on S. tarijense were positive. The results indicate that the responses by P. operculella to the types and density of trichomes are complex. Whereas type A and type B trichomes may act synergistically to reduce oviposition by the moth, leaf hairs do not defend against oviposition and neither leaf hairs nor type A and B trichomes reduce neonate establishment by this herbivore species.     

Molecular markers locate genes for resistance to the Colorado potato beetle,Leptinotarsa decemlineata,in hybrid Solanum tuberosum x S. berthaultii potato progenies

The wild Bolivian potato, Solanum berthaultii Hawkes, has been used as a source of resistance to the Colorado potato beetle (CPB), Leptinotarsa decemlineata Say, one of the most significant pests of potato. In this study, two reciprocal backcross S. tuberosum x S. berthaultii potato progenies, BCB and BCT, were mapped with RFLP markers and screened for resistance to CPB consumption, oviposition and defoliation. The genotypic and phenotypic data were combined and analysed to locate quantitative trait loci (QTLs) for resistance to CPB. Three QTLs on three chromosomes in BCB, and two QTLs on two chromosomes in BCT influenced resistance. The QTLs were generally additive but one instance of epistasis was noted. Each QTL accounted for 4–12% of the phenotypic variation observed in resistance. In the more resistant BCB population, a three QTL model explained ca. 20% of the variation in CPB oviposition. When alleles at the three QTLs were homozygous S. berthaultii, oviposition was reduced ca. 60% compared to the heterozygotes. The QTLs for resistance to CPB were compared to those previously identified for the type A and B glandular trichomes, which have been implicated in resistance in the same progenies. Generally, the QTLs for resistance to CPB coincided with loci associated with the glandular trichomes confirming the importance of the glandular trichomes in mediating resistance. However, a relatively strong and consistent QTL for insect resistance in both BCB and BCT on chromosome 1 was observed that was not associated with any trichome traits, suggesting the trichomes may not account for all of the resistance observed in these progenies.     

Host acceptance behaviour of Colorado potato beetle larvae influenced by potato glandular trichomes

ABSTRACT. In no-choice tests, larvae of the Colorado potato beetle, Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), avoided contact with, and were less frequently observed feeding on, excised foliage of the resistant wild potato Solanum berthaultii Hawkes than on leaves of the cultivated potato, S.tuberosum L. For older larvae, reduced feeding was a consequence of less frequent contact with S.berthaultii leaves. However, first instars appeared unable to penetrate the adhesive barrier presented by glandular trichomes of S. berthaultii , and those that remained on the leaflets fed less often than did first instars on S. tuberosum. Removal of the trichome barrier by wiping leaflets with tissue paper did not attenuate the apparent repellent effect of S. berthaulti foliage, but led to increased incidence of feeding by first instars. Inhibition of larval feeding may therefore rely on a mechanical barrier provided by the glandular trichomes of S.berthaultii.     

Recombinant and classically selected factors of potato plant resistance to the Colorado potato beetle, Leptinotarsa decemlineata, variously affect the potato aphid parasitoid Aphidius nigripes

Different forms of crop resistance developed against majorpotato pests such as the Colorado potato beetle (CPB), Leptinotarsadecemlineata, may be variously compatible with biological controlof secondary pests such as aphids. We compared the performance of theparasitoid Aphidius nigripes developing in the aphid hostMacrosiphum euphorbiae, on five potato lines, including atransgenic `Superior-BT' line expressing the CryIIIA toxin ofBacillus thuringiensis, which is specific to Coleoptera; andthe `NYL 235-4' line derived from Solanum berthaultii,characterized by relatively unspecific resistance to herbivores based onmoderately-high density of glandular trichomes. The other lines testedwere a `Kennebec-OCI' transgenic line expressing the protease inhibitorrice cystatin I (OCI), a potential resistance factor against coleopteranpests; and the commercial cultivars `Superior' and `Kennebec' used ascontrols. Parasitoid immature survival and adult size were reducedcompared to controls when the wasps developed on aphids fed the`Superior-BT' potato. In contrast, parasitoid size and fecundityincreased above control when the wasps developed on aphids fed the`Kennebec-OCI' potato. Parasitoids reached the adult stage faster andwere more fecund on `NYL 235-4' than control lines. The results indicatethat the different forms of potato resistance currently developed mainlyagainst the CPB had various unexpected effects on aphid parasitoidfitness. These effects on the parasitoid were complex but were generallyinterpretable in terms of host aphid quality variation among potatolines used as food by the aphids during parasitoiddevelopment.     

Effects of altitude of origin on trichome-mediated anti-herbivore resistance in wild Andean potatoes

The diversity and abundance of herbivores often decreases with increasing altitude. Plants are expected to respond to reduced herbivore pressure by allocating fewer resources to anti-herbivore resistance at higher altitudes. This study uses a greenhouse experiment and parallel bioassay to examine population variability in trichome-mediated defenses across a range of accessions of Solanum berthaultii Hawkes and S. tarijense Hawkes that originated from different altitudes. S. berthaultii bears two types of glandular trichomes, type A and type B, whereas S. tarijense has type A trichomes only, and hairs resembling type B trichomes that are eglandular. Both type A and type B glandular trichomes on S. berthaultii deterred ovipositing female Phthorimaea operculella (Zell.) (Lepidoptera: Gelechiidae). The density of type A, but not type B trichomes decreased with increasing altitude of origin in S. berthaultii populations. The ratio of type A to type B trichomes on the abaxial surface of S. berthaultii negatively affected oviposition and was inversely related to altitude of origin. In S. tarijense, type A trichomes deterred, but eglandular leaf hairs stimulated oviposition. Consequently, the ratio of type A trichomes to eglandular leaf hairs was negatively related to the number of eggs laid. The total numbers of trichomes per leaf generally increased with increasing altitude of origin in S. tarijense whereas the ratio of type A trichomes to eglandular hairs declined. In the oviposition bioassay, we found no direct relationship between resistance and altitude of origin, which may have been due to differences in leaf area at the time of the bioassay. Nevertheless, the results suggest that populations of both plant species that originated from higher altitudes were generally more susceptible to ovipositing P. operculella.     

Resistance of three wild tuber-bearing potatoes to the Colorado potato beetle

The resistance of Solanum okadae Hawkes & Hjert. (PI 458367), Solanum oplocense Hawkes (PI 473368), and Solanum tarijense Hawkes (PI 414150) to the Colorado potato beetle, Leptinotarsa decemlineata (Say) (Chrysomelidae: Chrysomelini), was studied. In replicated field trials all three accessions showed a high level of resistance to the beetle. No significant genetic variability between genotypes of the same species was found. Results from host acceptance behavior experiments, suitability for larval development tests, foliage consumption tests, and adult survival and oviposition tests supported the hypothesis that the mode of resistance differs between the three wild Solanum species. Solanum okadae and S. oplocense affected host acceptance and consumption. Because the beetle reacted differently to these two species it was hypothesized that the antifeedant chemical(s) differed in nature or quantity. S. tarijense contrasted with the other two species by affecting mostly adult colonization and oviposition.     

QTL analysis of potato tuber dormancy

The potential loss of chemical sprout inhibitors because of public concern over the use of pesticides underscores the desirability of breeding for long dormancy of potato (Solanum tuberosum L.) tubers. Quantitative trait locus (QTL) analyses were performed in reciprocal backcrosses between S. tuberosum and S. berthaultii toward defining the complexity of dormancy. S. berthaultii is a wild Bolivian species characterized by a short-day requirement for tuberization, long tuber dormancy, and resistance to several insect pests. RFLP alleles segregating from the recurrent parents as well as from the interspecific hybrid were monitored in two segregating progenies. We detected QTLs on nine chromosomes that affected tuber dormancy, either alone or through epistatic interactions. Alleles from the wild parent promoted dormancy, with the largest effect at a QTL on chromosome 2. Long dormancy appeared to be recessive in the backcross to S. berthaultii (BCB). In BCB the additive effects of dormancy QTLs accounted for 48% of the measured phenotypic variance, and adding epistatic effects to the model explained only 4% more. In contrast, additive effects explained only 16% of the variance in the backcross to S. tuberosum (BCT), and an additional 24% was explained by the inclusion of epistatic effects. In BCB variation at all QTLs detected was associated with RFLP alleles segregating from the hybrid parent; in BCT all QTLs except for two found through epistasis were detected through RFLP alleles segregating from the recurrent parent. At least three dormancy QTLs mapped to markers previously found to be associated with tuberization in these crosses.Paper number 55 of the Department of Fruit and Vegetable Science, Cornell University     

QTL analysis of potato tuberization

Quantitative trait loci (QTLs) affecting tuberization were detected in reciprocal backcrosses between Solanum tuberosum and S. berthaultii. Linkage analyses were performed between traits and RFLP alleles segregating from both the hybrid and the recurrent parent using a set of framework markers from the potato map. Eleven distinct loci on seven chromosomes were associated with variation in tuberization. Most of the loci had small effects, but a QTL explaining 27% of the variance was found on chromosome 5. More QTLs were detected while following alleles segregating from the recurrent S. tuberosum parent used to make the backcross than were detected by following alleles segregating from the hybrid parent. More than half of the alleles favoring tuberization were at least partly dominant. Tuberization was favored by an allele from S. berthaultii at 3 of the 5 QTLs detected by segregation from the hybrid parent. The additive effects of the QTLs for tuberization explained up to 53% of the phenotypic variance, and inclusion of epistatic effects increased this figure to 60%. The most common form of epistasis was that in which presence of an allele at each of 2 loci favoring tuberization was no more effective than the presence of a favorable allele at 1 of the 2 loci. The QTLs detected for tuberization traits are discussed in relationship to those previously detected for trichome-mediated insect resistance derived from the unadapted wild species.Paper number 54 of the Department of Fruit and Vegetable Science, Cornell University     

Tuber production,dormancy and resistance against Phthorimaea operculella (Zeller) in wild potato species

The diversification of resistant potato varieties at a landscape level could slow adaptation by Phthorimaea operculella to potato resistance and promote sustainable crop protection. In this study, we assessed wild potato species as novel sources of foliage and tuber resistance against P. operculella. Tuber resistance was quantified for 136 and foliage resistance for 54 potato accessions representing 14 and nine potato species, respectively. Several accessions were highly resistant to moth damage in tubers and/or foliage. In particular, Solanum chiquidenum and Solanum sandemanii were highly resistant to damage in tubers. Several accessions of Solanum multiinterruptum and a small number of accessions of Solanum bukasovii, Solanum berthaultii, Solanum sparsipilum and Solanum wittmackii also had highly resistant tubers. Larval survival on foliage of S. bukasovii and S. chiquidenum was generally low. New resistance sources are listed, and insect performance on the plants is described with possible resistance mechanisms. The study also examined potential trade‐offs associated with resistance. Tuber resistance was negatively correlated with the number and weight of tubers produced per plant, but positively correlated with the length of dormancy across accessions, indicating that, although long dormancy is not a prerequisite for resistance, species and accessions with extended dormancy will have more resistant tubers. Tuber and foliage resistance were generally positively correlated across all accessions; however, among accessions from within a potato species, there were negative (S. berthaultii), positive (S. chiquidenum) and non‐significant (S. bukasovii) relations. These results indicate that, besides identifying novel resistance sources, an improved understanding of the mechanisms and inherent trade‐offs associated with tuber and foliage resistance will improve the efficiency of potato breeding programmes aimed at enhancing resistance against P. operculella.     

Comparison of polyphenol oxidase expression in glandular trichomes of solanum and lycopersicon species

Tetralobulate glandular trichomes are present on the foliage of many solanaceous species. Resistance of many of these species to insects is conditioned by the ability of trichomes to rupture upon contact and to rapidly polymerize their contents, resulting in entrapment of insects in hardened trichome exudate. In the wild potato, Solanum berthaultii, polymerization of trichome exudate is initiated by a soluble M_r 59,000 polyphenol oxidase (PPO), which is a dominant protein constituent of the organ. PPOs, although ubiquitous in angiosperms, typically display great heterogeneity in molecular weight and are found at low levels in plant cells. Because of the unusually high accumulation and tissue-specific expression of the M_r 59,000 PPO in S. berthaultii glandular trichomes, we analyzed trichome proteins of a number of Lycopersicon and Solanum species to assess the extent to which possession of the M_r 59,000 PPO is conserved. Trichomes were collected manually and examined for PPO activity, immuno-cross-reactivity with S. berthaultiiM_r 59,000 PPO, and protein content. In addition, N-terminal amino acid sequences were obtained for five trichome PPOs. All species analyzed possessed trichome PPOs similar in structure and level of expression to that of S. berthaultii. The relationship between sequences and structures of these conserved PPOs and the variable PPOs of leaf is discussed.     

Behavioural responses of the Colorado potato beetle to trichomes and leaf surface chemicals of Solanum tarijense

Wild Solanum species constitute a source of resistance to several pests and diseases of potato. Several species of wild tuber‐bearing potato have been identified as resistant to the Colorado potato beetle, Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), including Solanum tarijense Hawkes (Solanaceae). Our objective was to determine the mechanism of resistance of S. tarijense to the Colorado potato beetle and, because the resistance is limited to the adult stage of the insect, to study the host selection behaviour on resistant plants. In the field, Colorado potato beetles demonstrated a unique behaviour when in contact with S. tarijense, abandoning the plant by falling to the ground after a few minutes. The abundant trichomes on the leaves of S. tarijense induced the falling behaviour. However, on S. tarijense feeding remained low even after the trichomes were mechanically removed. Observations demonstrated that the normal sequence of behaviour leading to feeding was interrupted before adult beetles fed on S. tarijense leaves. Feeding experiments using volatile and non‐volatile fractions of leaf surface extracts identified a phagodeterrent effect of the volatile fraction. Our results contrast with a similar evaluation of the mode of resistance of Solanum berthaultii Hawkes, a close relative of S. tarijense, on which some feeding occurred and adults did not show falling behaviour. This study presents information on S. tarijense as a new source of resistance to the Colorado potato beetle that can be used for potato breeding.     

Aphid repellent sesquiterpenes in glandular trichomes of Solanum berthaultii and S. tuberosum

Exudate from glandular trichomes of the wild potato species, Solanum berthaultii Hawkes, and the cultivated potato, S. tuberosum L., contained volatile substances including sesquiterpenes when examined with gas chromatography/mass spectrometry procedures. Sesquiterpenes were the major volatile constituents of Type A trichome exudate but were a minor constituent of Type B exudate. Sesquiterpene mixtures of the two potato species differed qualitatively and quantitatively. -Caryophyllene and E--Farnesene were major components identified in Type A trichome exudate of the cultivated potato. Sesquiterpene mixtures of both potato species were repellent to the green peach aphid, Myzus persicae (Sulzer). The role of trichome sesquiterpenes as a determinant of aphid behavior on the two potato species and in the expression of host resistance is discussed.

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Résumé L'examen en chromatographie gazeuse et en spectométrie de masse a révélé la présence de substances volatiles contenant des sesquiterpènes dans l'exsudat des trichômes glandulaires des pommes de terre sauvage (Solanum berthaultii) et cultivée (S. tuberosum). Les sesquiterpènes étaient le principal composé volatile de l'exsudat de type A, tandis qu'ils n'étaient que faiblement représentés dans l'exsudat de type B. Les mélanges de sesquiterpènes des deux espèces différaient qualitativement et quantitativement. Le -Caryophyllène et le E--Farnesène constituaient les principaux composés identifiés dans l'exsudat de trichôme de type A de S. tuberosum. Les mélanges sesquiterpéniques des deux espèces ont repousée Myzus persicae Sulzer. La discussion a porté sur l'influence des sesquiterpènes des trichômes dans la définition du comportement des pucerons sur ces deux plantes et sur la manifestation de la résistance de l'hôte.

     

Population dynamics of the potato tuber moth on processing tomatoes in Israel

The potato tuber moth (PTM),Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae), is a major pest of processing tomatoes,Lycopersicon esculentum Mill. (Solanaceae), in Israel. The larvae penetrate the tomato fruit through the stem end and present a serious threat to crop quality. Foliage and fruit samples were taken in nine commercial tomato fields located in Israel's three main tomato growing areas, two of which are potato growing areas as well. PTM was not found where potatoes were absent. Potato harvest in nearby fields was found to be the most significant factor affecting seasonal trends in PTM population density in tomatoes. All four larval instars were found in foliage on all sampling dates. Significantly higher proportions of first instars were found during the population density increase which followed potato harvest. Damaged fruits did not contain first instar larvae, indicating that PTM never undergoes complete development within tomato fruit. Fruit damage levels at harvest were positively correlated to the peak mean population densities on foliage and the date they were observed. In tomato fields not adjacent to potatoes, infestation was first observed at the edge of the field. Both before and after the potato harvest in nearby fields, population density at the edge of the field was significantly higher than at the center. In tomato fields adjacent to potatoes, no significant differences were found between population densities at the edge and center before the potatoes were harvested. After the potato harvest, population density at the center of tomato fields was higher than at the edge. Deceased, October 1988     

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.     

Glandular Trichomes on Alfalfa Impede Searching Behavior of the Potato Leafhopper Parasitoid

New cultivars of alfalfa, Medicago sativa L., have been released with glandular trichomes for resistance to potato leafhopper, Empoasca fabae (Harris). Yet, the impact of the glandular trichomes on the primary natural enemy of the leafhopper, Anagrus nigriventris Girault, is unknown. We compared the host searching behavior of the egg parasitoid on four alfalfa clones varying in trichome characters. Female wasps were videotaped on Ranger, a susceptible clone with relatively sparse trichomes, B14, a resistant clone with dense but nonglandular trichomes, and FG12 and FG18, two resistant clones with glandular trichomes. Although the number of leafhopper eggs per stem exposed to wasps did not significantly differ among the four clones, the frequency of foraging and total foraging time were less on the two clones with glandular trichomes than on the two clones with nonglandular trichomes. In addition, an analysis of covariance demonstrated that, although the number of ovipositional probes increased with egg density on a stem, the number of probes on stems with glandular trichomes was significantly less than that on stems without glandular trichomes. The allocation of time by wasps among drumming, probing, and grooming behaviors was similar among the clones. Wasps tended to fly off of clones with glandular trichomes more often than off of clones with nonglandular trichomes. This study suggests that cultivars with glandular trichomes may interfere with host searching by A. nigriventris.     

Location of resistance factors in the leaves of potato and wild tuber-bearing Solanum species to the aphid Myzus persicae

Analysis of electrically recorded feeding behaviour of aphids was combined with colony‐development tests to search for sources of resistance to Myzus persicae (Sulzer) (Homoptera: Aphididae) in tuber‐bearing Solanum species (Solanaceae), aiming at a reduction of potato leaf roll virus (PLRV) transmission. Twenty genotypes, originating from 14 gene bank accessions, representing 13 wild tuber‐bearing Solanum spp., three Solanum tuberosum L. (potato) cultivars, and one S. tuberosum breeding line, were selected. Colony‐development tests were carried out in no‐choice experiments by placing adult aphids on plants of each genotype and counting numbers of nymphs and adults on young plants after 8 and 15 days, and on flowering plants after 14 and 30 days. Large differences were observed among genotypes: some developed small colonies and others developed large ones. Also, in a few genotypes, resistance in mature plants was different for leaves of different ages; young leaves were resistant to aphids whereas old senescent leaves were susceptible. The electrical penetration graph (DC‐EPG system) technique was used to study aphid feeding behaviour on each Solanum genotype for 6 h. Electrical penetration graph (EPG) results also showed large differences among the genotypes, indicating resistance at the leaf surface and at three different levels of plant tissue (epidermis, mesophyll, and phloem). Therefore, it was concluded that different mechanisms of resistance to M. persicae exist among the genotypes analysed. EPGs recorded from aphids on Solanum berthaultii Hawkes and Solanum tarijense Hawkes with and without glandular trichomes showed that strong surface resistance can bias EPG parameters associated with resistance located in deeper tissues. Experimental evidence is presented that the resistance to aphids in the genotypes with glandular trichomes strongly depends on these morphological structures.     

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.     

Nonglandular leaf trichomes as short-term inducible defense of the grey alder,Alnus incana (L.), against the chrysomelid beetle,Agelastica alni L.

Summary The grey alder compensates leaf area losses due to insect grazing by continuously producing new leaves throughout the vegetative period. Different degrees of defoliation were attained experimentally by a controlled release of the oligophagous beetle Agelastica alni on arbitrarily selected trees from a homogenous population of young alders. The reduction in leaf area per tree significantly influenced the density of leaf trichomes, assessed 10–30 days later, on newly sprouting leaves only. Cross-correlations between leaf area reduction and trichome density were strongest for leaves which completed unfolding 14–21 days after damage. Dualchoice assays suggested a negative influence of trichomes on oviposition rate of A. alni. Removal of trichomes by shaving demonstrated the highly significant effect of trichomes on feeding behavior of adults and larvae in dual-choice assays. The role of the induced increase in trichome density as a possible short-term defense reaction against herbivorous insects is discussed.