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.     

2-tridecanone, tomatoes and Heliothis zea: potential incompatibility of plant antibiosis with insecticidal control

The allelochemic, 2-tridecanone, which conditions resistance in the wild tomato Lycopersicon hirsutum f. glabratum to Manduca sexta and plays an important role in the resistance to Leptinotarsa decemlineate is shown to induce an enhanced level of tolerance to the carbamate insecticide carbaryl in a third important insect pest of tomato, Heliothis zea. This phenomenon has important implications regarding the compatibility of using tomato cultivars having 2-tridecanone-mediated insect resistance in conjunction with insecticides to manage the full spectrum of tomato insect pests.

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Résumé Le 2-Tridécanone,est présent dans les extrémités des poils glandulaires des feuilles de Lycopersicon hirsutum et glabratum (PI 134417). On a précédemmment mis en évidence son rôle capital dans la résistance du PI 134417 à Manduca sexta L. et à Leptinotarsa decemlineata Say, mais sa faible importance dans la résistance à Heliothis zea Boddie. Au cours de cette étude, la paralysie temporaire des larves néonates de H. zea par le 2-Tridécanone associé aux feuilles a été observée, mais cette substance induit chez ces larves un système de détoxification qui entraîne, non seulement une récupération apparemment complète, mais provoque une résistance au carbaryl, insecticide au carbamate, 3–4 fois plus élevée.Dans les programmes d'amélioration de la tomate, toute sélection pour une résistance élevée à M. sexta et L. decemlineata attribuable à une forte teneur en 2-Tridécanone, sans sélection parallèle pour la résistance à H. zea, caractère génétique séparé, produirait des lignées de tomates chez lesquelles la protection contre H. zea serait plus difficile par suite de la résistance au carbaryl induite par le 2-Tridécanone.

     

2-Undecanone, a constituent of the glandular trichomes of Lycopersicon hirsutum f. glabratum: Effects on Heliothis zea and Manduca sexta growth and survival

The effects of 2-undecanone, a compound found in the glandular trichomes of the wild tomato, Lycopersicon hirsutum f. glabratum C. H. Mull, PI 134417, on the larvae of Heliothis zea (Boddie) and Manduca sexta (L.) were studied by rearing the larvae on artificial diets containing this chemical. 2-Undecanone increased larval mortality of H. zea in the first 48 h when combined with 2-tridecanone, but not alone, and caused deformity and mortality of H. zea pupae both alone and in combination with 2-tridecanone. 2-Undecanone had no effect on M. sexta larvae or pupae. Potential implications of these findings for insect pest management are discussed.

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Résumé Le 2-undécanone est une methyle cétone qui se trouve avec le 2-tridécanone dans les extrémités des poils glandulaires des feuilles de la tomate sauvage: Lycopersicon hirsutum f. glabratum C. H. Mull, PI 134417. Notre étude entreprendait à rechercher le rôle du 2-undécanone dans la resistance de cette plante à Heliothis zea et Manduca sexta.Les produits chimiques synthétiques s'étaient incorporés en diète artificièlle sur laquelle les insectes ont élevés. Le 2-undécanone a causé peu de mortalité des larves de H. zea sauf quand il soit combiné avec le 2-tridécanone, dans quel cas on trouve, dans les premiers 48 heures, une augmentation de mortalité, en comparaison au 2-tridécanone seul. Aussi le 2-undécanone a causé les difformités des nymphes de H. zea et la mortalité dans l'étage de nymphe, toutes les deux seul et en combinaison avec le 2-tridécanone. En contrast, le 2-undécanone n'a pas produit un effet sur les larves ou nymphes de M. sexta.L'élevage sélective des lignes de tomates de niveau plus élevé de 2-undécanone pourrait utile pour la lutte contre le H. zea. En combinaison avec des niveaux élevés du 2-tridécanone, c'est possible de reduire les populations nuisibles des larves. Il peut aussi réduire le nombre des lépidoptères adultes produit dans la récolte de tomate. Toutefois, il ne serait pas probablement de valeur dans la lutte contre le M. sexta.

     

2-Undecanone, a pupal mortality factor in Heliothis zea: sensitive larval stage and in planta activity in Lycopersicon hirsutum f. glabratum

2-Undecanone, a compound found in the tips of the Type VI glandular trichomes of the wild tomato species Lycopersicon hirsutum f. glabratum C. H. Mull (accession PI 134417), when incorporated in artificial diet, was previously shown to cause pupal deformity and mortality among Heliothis zea (Boddie). In the present study, the developmental stage of H. zea sensitive to the effects of dietary 2-undecanone was determined by rearing larvae through the fourth instar on diet with or without 2-undecanone, then transferring them to new diet either with or without 2-undecanone. Dietary levels of 2-undecanone (0.055% wet wt) were representative of concentrations found in PI 134417 foliage. Ingestion of 2-undecanone during the fifth instar was both necessary and sufficient to cause pupal mortality. Rearing fifth instar H. zea on PI 134417 foliage with glandular trichomes intact (2-undecanone present at 0.084±0.026% wt wt) or with the trichomes, and hence 2-undecanone, removed indicated that these effects occur in planta. Potential implications of these findings for using 2-undecanone-mediated resistance for insect pest management are discussed.

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Résumé Il a été démontré précédemment que l'undécanone-2, présente dans les extrémités des trichomes glandulaires de type VI de la tomate sauvage Lycopersicon hirsutum glabratum C. H. Mull (accession PI 134417), provoque déformations et mortalité chez les chrysalides de Heliothis zea (Boddie) lorsqu'elle est incorporée dans une nourriture artificielle. Dans cette étude, le stade de développement de H. zea sensible aux effets de l'undécanone-2 a été déterminé en élevant les larves jusqu'au 4e stade sur une diète contenant ou non de l'undécanone-2, puis en les transférant sur une nouvelle diète avec ou sans undécanone-2. Les concentrations en undécanone-2 utilisées (0.055% du poids frais) étaient comparables aux concentrations présentes dans les feuilles du PI 134417. L'ingestion de l'undécanone-2 durant le 5e stade s'est avérée nécessaire et suffisante pour provoquer une mortalité au stade chrysalide. L'élevage du 5e stade de H. zea sur des feuilles de PI 134417 dont les trichomes sont intacts (l'undécanone-2 est présente à la concentration de 0.084±0.025% du poids frais) ou sur des feuilles dont les trichomes, et par conséquent l'undécanone-2, ont été éliminés, a démontré que les effets toxiques se produisent in planta.L'undécanone-2 pourrait être un facteur de résistance efficace des tomates, mais probablement uniquement dans les régions où H. zea est polyvoltime sur la tomate et où les premiéres générations se développent sur le feuillage pendant toute la période larvaire avant la maturation des fruits. Normalement, les premiers stades larvaires de H. zea se nourrissent de feuilles et les derniers stades des fruits. Les fruits ne contenant pas d'undécanone-2, les chenilles qui se développeraient sur les plants en fructification pourraient échapper aux effets toxiques.

     

Relationship of leaf lamellar-based resistance toLeptinotarsa decemlineata andHeliothis zea in a wild tomato,Lycopersicon hirsutum f.glabratum, PI 134417

Segregating populations of hybrids of the insect-resistant wild tomato,Lycopersicon hirsutum f.glabratum, C.H. Mull, PI 134417, and the susceptible tomato cultivar ofL. esculentum Mill, Walter, were screened by bioassays with the Colorado potato beetle,Leptinotarsa decemlineata (Say) and tomato fruitwormHeliothis zea (Boddie). Plant lines with a range of levels of resistance toH. zea were selected from one group of hybrids; plants with a range of resistance levels toL. decemlineata, from another group. The response of both insect species to both groups of plants was evaluated. Resistance to each of these species is under separate genetic control and apparently involves distinct mechanistic components, although it remains possible that at least some factors are important in conditioning resistance to both species.

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Résumé La tomate sauvage,L. hirsutum f.glabratum C.H. Mull, n⁰ PI134417, présente des lamelles foliaires qui lui confèrent une résistance au doryphore,L. decemlineata Say et àH. zea Bodie. Cette étude a voulu préciser si ces résistances étaient héritées ensemble dans des populations d'hybrides entre PI 134417 et la tomate cultivée,L. esculentum Mill. Des lignées de plantes présentant une gamme de résistance àH. zea ont été sélectionnées à partir d'une population hyrbide. De même, des lignées de plantes ayant une gamme de résistance àL. decemlineata ont été sélectionnées à partir d'un second groupe d'hybrides. Les réactions des 2 insectes aux 2 groupes de plantes ont été estimées.Les résistances àH. zea et àL. decemlineata n'étaient pas corrélées nettement. Certaines lignées sont résistantes aux 2 espèces, mais d'autres ne sont résistantes qu'à une espèce. Ainsi, les résistances à chacune de ces espèces ont probablement des déterminismes génétiques différents. Différentes composantes mécaniques sont vraisemblablement impliquées dans ces résistances, bien qu'il soit possible qu'au moins quelques facteurs aient un rôle important dans le conditionnement de la résistance. La création de cultivars de tomates avec une résistance liée aux lamelles foliaires sera compliquée par le besoin d'une sélection en fonction des réponses aux 2 espèces..

     

Induced tolerance of neonate Heliothis zea to host plant allelochemicals and carbaryl following incubation of eggs on foliage of Lycopersicon hirsutum f. glabratum

Summary Incubation of Heliothis zea (Boddie) eggs on foliage of Lycopersicon hirsutum f. glabratum C.H. Mull (accession PI 134417) results in neonates with elevated levels of tolerance to the toxic effects of PI 134417 foliage attributable to 2-tridecanone found in the glandular trichomes which abound on that foliage. The neonates from such eggs are also shown to have elevated levels of tolerance to the carbamate insecticide carbaryl. Incubation of eggs in an atmosphere containing 2-tridecanone similarly produced elevated levels of tolerance to 2-tridecanone among resulting neonates, indicating that 2-tridecanone is the likely inducing agent and that exposure to 2-tridecanone vapor, which is known to emanate from PI 134417 foliage, is sufficient for induction. Analysis of the cytochrome P-450 content in gut microsomes of fifth instar larvae indicated that exposure of larvae to 2-tridecanone in artificial diet or to PI 134417 foliage resulted in significantly elevated levels of cytochrome P-450 relative to larvae fed diet without 2-tridecanone or foliage of L. esculentum which contains no 2-tridecanone. In addition, removal of the glandular trichomes from PI 134417 foliage eliminated the ability of that foliage to induce elevated levels of cytochrome P-450. These results provide circumstantial evidence that cytochrome P-450 may be involved in the induced tolerance to xenobiotics among neonates from eggs exposed to 2-tridecanone or PI 134417 foliage.Support for this research was provided by the USDA Competitive Research Grants Program in Biological Stress under Grant No. 83-CRCR-1-1241 and Grant No. 85-CRCR-1-1615, and the North Carolina Agricultural Research Service. Paper No. 10856 of Journal Series of the North Carolina Agricultural Research Service, Raleigh, NC, USA 27650. Use of trade names does not imply endorsement of products named nor criticisms of similar ones not mentioned     

Growth, food consumption and mortality of Heliothis zea larvae on foliage of the wild tomato Lycopersicon hirsutum f. glabratum and the cultivated tomato, L. esculentum

Larvae of Heliothis zea (Boddie) were fed foliage of the resistant wild tomato Lycopersicon hirsutum f. glabratum C. H. Mull (PI 134417) and susceptible L. esculentum (cultivar Walter), either with the glandular trichomes removed or with the trichomes intact. Response variables measured included larval mortality, larval weight, consumption rate (CR), and efficiency of conversion of ingested material (ECI). Resistant foliage was found to contain factors in both the trichomes and leaf lamellae which increased larval mortality, reduced larval weight, reduced CR, and, except in second instars, reduced ECI. Susceptible foliage also contained trichome-based factors which increased larval mortality and decreased weight, as well as CR of second instars and ECI of fifth instars. Implications of these findings for breeding resistant tomato cultivars are discussed.

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Résumé La tomate sauvage, Lycopersicon hirsutum f. glabratum C. H. Mull (PI 134417) est fortement résistant à Heliothis zea. Les études préalables ont indiqués que la résistance est associée avec la présence de 2-tridécanone dans les extrémités des poils glandulaires de la tomate sauvage, et que les facteurs dans la lamelle des feuilles sont peut-être impliqués. À déterminer la présence des facteurs lamellaires et à caractériser de plus les facteurs de résistance dans les extrémités des poils glandulaires aussi, nous nourrissions les larves du feuillage résistant de PI 134417 et du feuillage susceptible L. esculentum (lignée Walter) les deux avec les extrémités des poils glandulaires enlèvés ou avec les extrémités des poils glandulaires intacts. Nous mesurions la mortalité des larves, les poids des larves, la taux de consommation relative et l'efficacité de conversion de matière mangé.On se trouvait le feuillage résistant à contenir les facteurs, tous deux dans le feuillage lamellaire et dans les extrémités des poils glandulaires, qui causaient l'augmentation de mortalité, les poids plus basses, la taux de consommation relative plus basse, et, hormis dans le cas des larves en phase deuxième, l'efficacité de conversion réduite. Le feuillage susceptible contenait aussi les facteurs dans les extrémités des poils glandulaires qui augmentaient la mortalité, diminuaient les poids et la taux de consommation relative des larves en phase deuxième, et diminuaient l'efficacité de conversion des larves en phase cinquième. Cettes conclusions indiquent la présence des facteurs de résistance en outre de 2-tridécanone et peuvent fournir les indices à l'identité de ces facteurs.

     

Accumulation and turnover of 2-tridecanone in Tetranychus urticae and its consequences for resistance of wild and cultivated tomatoes

In this study we assessed the dynamic changes of 2-tridecanone in a herbivorous mite (Tetranychus urticae) on tomato (Lycopersicon esculentum, cv. Moneymaker), a plant with methyl ketones in the tetracellular tips of the glandular trichomes (Type VI). We showed that spider mites accumulate 2-tridecanone when foraging on cultivated tomato. Thus, the rate of mite–trichome contact multiplied by the amount of toxin per trichome tip exceeded the relative rate of toxin turnover multiplied by the amount of toxin per mite. The relative rate of toxin turnover was estimated to be 1.1 per day on cucumber, a plant without this toxin. The amount per trichome tip varied from 0.33 ng for middle-leaf trichomes to 1.26 ng for main-stem trichomes. Hence, to achieve a static level of 2-tridecanone equal to 8–17 ng per mite – representing the level we found in mites on middle leaves – the rate of mite–trichome contact should be 26–57 per day. Because methyl ketone apparently accumulates in the spider mites on tomato, the rate of mite–trichome contact is probably higher than that. We expect the accumulation of ketones to occur especially on the stems of cultivated tomato, since this is the area most densely occupied with glandular hairs and because here the hairs have higher levels of the methyl ketones.Using dose–response relationships assessed earlier (Chatzivasileiadis and Sabelis, 1997, 1998), we estimated that the number of mite–trichome contacts causing 50% mortality per day is equal to 88 on a tomato stem, whereas it equals 70 for another strain of spider mites collected from cucumber. On wild tomato, L. hirsutum f. glabratum (PI 134417), just one to two contacts would suffice to cause 50% mortality per day. We suggest that methyl ketones from glandular hairs on tomato are an important mortality factor for spider mites on wild tomato and probably also on cultivated tomato.     

Field evaluation of insect resistance in a wild tomato and its effects on insect parasitoids

Populations ofHelicoverpa (=Heliothis) zea (Boddie),Heliothis virescens (F.),Manduca sexta (L.) andM. quinquemaculata (Haw.) and their egg and larval parasitoids were sampled in field plots of the: insect-resistant wild tomato,Lycopersicon hirsutum f. glabratum C. H. Mull, accession PI 134417; susceptible commercial tomato cultivar ‘Better Boy’; F₁ hybrid; and selected, moderately resistant backcross genotype. Densities ofH. zea andH. virescens eggs and small larvae were higher on resistant genotypes than on susceptible genotypes, but densities of large larvae were similar on all genotypes. Densities ofManduca spp. larvae were too low to permit similar analyses of the effects of plant genotype. Rates of egg parasitism byTrichogramma spp. andTelenomus sphingis (Ashmead) were reduced on insect-resistant genotypes. Rates of parasitism by the larval parasitoidsCampoletis sonorensis (Cameron) andCotesia congregata (Say) were reduced on resistant genotypes. No consistent effects on parasitism rates byCotesia marginiventris (Cresson) were observed and parasitism rates byCardiochiles nigriceps Viereck were unaffected.     

Response of insecticide-resistant and susceptible Colorado potato beetles, Leptinotarsa decemlineata to 2-tridecanone and resistant tomato foliage: the absence of cross resistance

Insecticide-resistant Colorado potato beetles (CPB), Leptinotarsa decemlineata, from Long Island, New York and susceptible beetles from North Carolina were tested for response to 2-tridecanone and foliage of the CPB resistant wild tomato, Lycopersicon hirsutum f. glabratum C. H. Mull, PI 134417. Populations did not differ in their response to 2-tridecanone or PI 134417 foliage. Exposure of beetles to sublethal doses of 2-tridecanone did not increase tolerance of either population to subsequent exposures to 2-tridecanone. It was also found that the New York beetles suffered higher mortality than North Carolina beetles on CPB susceptible L. esculentum foliage and PI 134417 foliage from which the glandular trichomes, which contain 2-tridecanone, were removed. Survival of North Carolina beetles was lower on PI 134417 foliage without trichomes than on L. esculentum foliage.

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Résumé La lutte contre Leptinotarsa decemlineata, sur la tomate et des autres récoltes, compte ordinairement sur les insecticides. Cette confiance a causé plusieurs populations de cette espèce à développer une résistance aux insecticides. Accession PI 134417 de la tomate sauvage, Lycopersicon hirsutum f. glabratum C. H. Mull, est résistant à L. decemlineata, en grande partie à cause de la presence de 2-tridécanone dans les éxtremités des poils glandulaires du feuillage. Notre étude entreprendait à rechercher la possibilité d'une résistance croisée aux insecticides et au 2-tridécanone.Les coléoptères de Long Island, New York, ceux qui sont résistants aux insecticides et ceux susceptibles de North Carolina, éprouvaient pour une réponse au 2-tridécanone et au feuillage résistant.Les deux populations de coléoptères ne diffèrent pas dans sa réponse au 2-tridécanone ou au feuillage résistant. Nous ne trouvons pas d'évidence d'une résistance croisée.Aussi, nous trouvons que les coléoptères de New York ne survivaient pas aussi bien que ceux de North Carolina au feuillage résistant sans les éxtremités des poils glandulaires et au feuillage susceptible. La survivance était plus basse sur le feuillage résistant sans les éxtremités des poils glandulaires que le feuillage susceptible seulement avec les coléoptères de North Carolina.

     

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.     

Populations of Predaceous Natural Enemies Developing on Insect-Resistant and Susceptible Tomato in North Carolina

Naturally occurring populations of immature and adultGeocoris punctipes,adultColeomegilla maculataand immature coccinellids were monitored on field-grown tomato lines susceptible and resistant toManduca sextaandHelicoverpa zea. Helicoverpa zeaandHeliothis virescenseggs and small larvae that serve as prey for these predators also were monitored. MoreH. zeaandH. virescenseggs and small larvae were found on resistant than on susceptible plant lines. However, similar populations of largeH. zeaandH. virescenslarvae were found on resistant and susceptible plants. The number of adultGeocoris punctipes,adultColeomegilla maculataand immature coccinellids on resistant plants was always as high or higher than the number on susceptible plants. The data demonstrate no incompatibility of host-plant resistance with biological control provided by these predaceous insects, but indicate that the number ofG. punctipesand coccinellids required to provide effective biological control may develop too late in the season to be of practical value. Large populations of stilt bugs (Jalysus wickhami,Hemiptera: Berytidae) and spiders were observed to occur earlier in the growing season than eitherG. punctipesor coccinellids and may be a significant source of mortality forH. zeaeggs and small larvae.     

Interaction ofManduca sexta resistance in tomato with insect predators ofHelicoverpa zea

Resistance in accession PI 134417 ofLycopersicon hirsutum f.glabratum toManduca sexta L. results from the presence of the methyl ketones 2-tridecanone and 2-undecanone in the type VI trichome glands on foliage of these plants. Short (24 h) and long term (neonate to adult) laboratory experiments usingM. sexta-resistant and susceptible foliage and methyl ketone treated filter paper disks were conducted to evaluate the direct (plant-mediated), and indirect (prey-mediated) effects of this resistance on two predators ofHelicoverpa zea eggs,Coleomegilla maculata (DeGeer) andGeocoris punctipes (Say). Direct effects of resistance were manifest as reduced egg consumption and increased mortality for both predators. Indirect effects were manifest as reduced egg consumption byG. punctipes, but were not observed forC. maculata. Results of experiments using methyl ketone-treated filter paper disks instead of foliage were similar.     

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.

     

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.     

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.     

Wild species of Nicotiana as a new source of tobacco resistance to the tobacco hornworm Manduca sexta

Sixteen species of Nicotiana were examined for resistance to second instar larvae of the tobacco hornworm, Manduca sexta (L.). The tests were designed so as to discriminate between antibiosis and nonpreference. High levels of antibiosis resistance were observed in several wild species such as N. gossei and species in the Repandae section. Several other species, such as N. excelsior, showed antibiosis that appeared to be due to a different mechanism than the alkaloid-trichome exudate based resistance of the above species. The data indicate that these species of Nicotiana may be used as a new source of resistance to larvae of M. sexta.

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D'espèces sauvages de Nicotiania comme nouvelle source de résistance du tabac à Manduca sexta

Résumé La résistance aux chenilles de second stade de Manduca sexta L. a été examinée chez seize espèces de Nicotiana. Les essais ont été conçus pour distinguer absence d'appétance et antibiose. Plusieurs espèces sauvages comme N. gossei et les espèces de la section Repandae ont présenté un niveau élevé d'antibiose. D'autres espèces, comme N. excelsior ont présenté une antibiose vraisemblablement due à un mécanisme différent de la résistance des espèces précédentes provoquée par un exsudat alcaloïde des trichomes. Les résultats indiquent que ces espèces de Nicotiana peuvent être utilisées comme nouvelle source de résistance aux chenilles de M. sexta.

     

Resource availability and the trichome defenses of tomato plants

We conducted two experiments to determine how resource availability influenced allocation by tomato (Lycopersicon esculentum) to trichomes, and how different patterns of trichome allocation by plants grown in different resource environments might then influence the behavior of tobacco hornworm (Manduca sexta) caterpillars. In the first experiment we used high and low levels of light and water, and then, using scanning electron microscopy, determined trichome densities on the leaves and stems. We sampled leaves and stems at several places throughout the plant to determine whether there were within-plant differences in allocation to trichomes. The results of the first experiment showed that resource availability influenced allocation to trichome growth. Patterns in high and low-light supported both the growth-differentiation balance hypothesis (GDBH) and the carbon-nutrient balance hypothesis (CNBH). However, the GDBH was not supported by differences among water treatments. Contrary, to predictions of the GDBH, plants with intermediate growth did not have the highest trichome densities, and plants with similar growth differed in trichome density. Possible biological and artifactual explanations are discussed. The first experiment also showed that there was within-plant variation in allocation to trichomes, and that plant resource availability may influence within-plant variation in allocation to trichomes. In the second experiment, we grew plants in high and low-light, and then monitored the behavior of tobacco hornworms on the stems of these plants in the laboratory. This experiment demonstrated that the light environment that tomato plants were grown in influenced the resting behavior of caterpillars. Furthermore, it demonstrated that both glandular and non-glandular trichomes impeded caterpillars from searching for food. Overall, this study indicated that plant resource availability can influence allocation to trichome defenses, and that these differences may affect insect herbivores.     

Trichome morphology provides phylogenetically informative characters for Tremandra, Platytheca and Tetratheca (former Tremandraceae)

Summary Trichomes of Tremandra R.Br. ex DC., Platytheca Steetz and Tetratheca Sm. (Elaeocarpaceae, former Tremandraceae), together with two outgroup species of Elaeocarpus L., are illustrated using scanning electron microscopy, and their distribution on various plant organs is documented. Various trichomes types were identified that relate taxa: simple hairs, stellate hairs, short glandular trichomes, long glandular trichomes, and three forms of tubercules. Both outgroup and ingroup taxa have simple hairs. Stellate hairs are confirmed as unique to Tremandra. Prominent and sculptured multicelled tubercules, some bearing a stout hair, are characteristic of Platytheca. Smaller multicelled tubercules occur in both Platytheca and Tetratheca, except for the Western Australian taxon Te. filiformis Benth. (possibly plesiomorphic). Unicellular tubercules (papilla) characterise two species of Tetratheca. Short glandular trichomes, usually found on the ovary, also occur in both of these genera but not in all species (possibly secondary losses), while long glandular trichomes, usually on stems and leaves, occur only in some groups of Tetratheca. Within Tetratheca, Western Australian taxa that have five-merous flowers fall into three ‘groups’: seven species (together with one from South Australia) that have short glandular trichomes but no long glandular trichomes; six species that have long glandular trichomes but no short glandular trichomes; and four species that have both trichome types. All other species of Tetratheca have four-merous flowers and form two ‘groups’: 12 eastern species (including one from South Australia) that have both short glandular trichomes and long glandular trichomes; 4 western species and six eastern species that lack short glandular trichomes. On the basis of these characters, a phylogenetic hypothesis for the three genera is presented.