Glandular trichomes and sesquiterpene lactones of Artemisia nova (Asteraceae)

The leaves of Artemisia nova Nelson are covered with a dense layer of nonglandular trichomes that obstruct the observation of glandular trichomes on the leaf surfaces. An acid-staining technique was developed that eliminates interference from the nonglandular trichomes and stains the glandular ones red, allowing them to be seen under a light microscope. Using this method it was found that these glands develop on both leaf surfaces, except over veins, and cover 21–35% of the leaf surface area at densities of 50–70 per mm². Mature glands are sac-like structure with a thin sheath or cuticle surrounding a clear transparent fluid. This fluid contains all of the sesquiterpene lactones present in the leaves, and some of the monoterpenes. These terpenoids were responsible for the red color of the glands after acid treatment.     

Feeding patterns of a specialist and a generalist grasshopper: electronic monitoring on their host plants

Abstract Hypochlora alba (Dodge) is a specialist grasshopper that lives and feeds almost exclusively on Artemisia ludoviciana Nutt., a sage plant mostly avoided as food by generalist or polyphagous grasshoppers, such as Melanoplus sanguinipes (Fabr.). Adaptations to feeding on the pubescent sage foliage containing high levels of allelochemics may involve behavioural as well as physiological specialization. Using an electronic device to monitor feeding activity, we compared feeding periodicity, bout length, bout frequency, and overall time spent feeding by H.alba on sage foliage and by M.sanguinipes on seedling ryegrass ( Lolium perenne L.), the plants on which they had been reared. Although both species fed for about the same amount of time per day (c. 4–5%), they achieved this by two different feeding patterns. H. alba had feeding bouts averaging c. 10 min each, compared to c. 4–5 min for M.sanguinipes. Bout frequency was reversed, with M.sanguinipes feeding about once every hour and H.alba about half as often. Bout length for both species was unaffected by phase of the photoperiod while bout frequency, and consequently the rate of feeding in min/h, was typically reduced by two-thirds or more during the dark phase. The relationship of grasshopper feeding behaviour to host plant specialization is discussed.     

Attraction of the grasshopper, Melanoplus sanguinipes, to host plant odors and volatile components

The attraction of the polyphagous grasshopper, Melanoplus sanguinipes (F.), to odors from plant foliage or to chemical components of these odors was tested in a glass Y-tube olfactometer. Humidified air was passed at equal flow rates through a sample chamber containing plant material and an empty control chamber and then through the Y-tube to the holding chamber containing the test insect. Solutions of volatile chemicals in water were metered at a constant rate into the sample air stream with a syringe pump. Insects moving upwind were recorded after entering either the sample or control arm of the Y-tube. Both nymphs and adults were strongly attracted to the odor of chopped and intact seedling foliage of perennial ryegrass and wheat. Chopped leaves but not intact leaves of sorghum and alfalfa also were significantly attractive. The major components of ryegrass odor from both chopped and stem-cut leaves were Z-hex-3-en-1-yl acetate, Z-hex-3-en-1-ol, and pent-1-en-3-ol, in that order, with lesser amounts of E-hex-2-enal. Chopped leaves released many more minor components. The major components when tested individually or in binary and ternary mixtures were significantly attractive to grasshoppers compared to humidified air but not to the degree of whole plant odor. However, a quartenary mixture simulating the odor blend had levels of attractancy equal to that of chopped grass odor. Blends of these 5- and 6-carbon unsaturated alcohols, esters, and aldehydes volatilizing from green plants probably play an important role as olfactory cues for orientation of grasshoppers to food plants.

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Résumé L'attraction de M. sanguinipes (F.) par l'odeur du feuillage et des composés chimiques de ces odeurs a été examinée dans un olfactomètre en tube de verre en Y. Des flux identiques d'air humidifié étaient introduits dans une enceinte contenant le matériel végétal et dans une enceinte témoin, et, via le tube en Y, dans l'enceinte contenant l'insecte à étudier. Des solutions aqueuses de substances volatiles étaient introduites à un taux constant dans le courant d'air grâce à une seringue. Les insectes se déplaçant contre le vent étaient enregistrés après pénétration dans une branche déterminée du tube en Y. Larves et adultes étaient fortement attirés par l'odeur de feuillage intact ou coupé de semis de Lolium perenne et de blé (Triticum aestivum). Les feuilles coupées, mais non les feuilles intactes de sorgho (Sorghum bicolor) et de luzerne (Medicago sativa) étaient, elles aussi, significativement attractives. Les principales substances constituant de l'odeur de feuilles de Lolium perenne, tant coupées qu'intactes, étaient dans l'ordre de Z-hex-3-en-1-yl acétate, le Z-hex-3-en-1-ol et le pent-1-en-3-ol, et dans une moindre mesure, E-hex-2-enal. Les feuilles coupées libéraient beaucoup plus de composés secondaires. Les principaux constituants testés, seuls ou en mélanges binaires ou ternaires, étaient significativement attractifs, mais moins que l'odeur totale de la plante. Toutefois, un mélange quaternaire, simulant le mélange odorant, avait le même pouvoir attractif que l'odeur de la plante entière. L'union de ces alcools, esters et aldéhydes à 5- et 6-carbones non saturés, volatilisés à partir des plantes vertes, jouent probablement un grand rôle comme signaux orientateurs lors de l'attraction des criquets par leur plantes alimentaires.

     

Orientation responses of the grasshopper, Melanoplus sanguinipes, to visual, olfactory and wind stimuli and their combinations

Prereproductive adults of the grasshopper, Melanoplus sanguinipes (F.) (Orthoptera, Acrididae), demonstrated orientation and movement towards both visual and olfactory stimulus sources in a still-air chamber. Visual stimuli (wheat and lima bean foliage, vertical black or yellow-green stripes, and a yellow-green broad leaf pattern) were approached more frequently than the control white background surface. Olfactory stimuli (chopped wheat foliage and a four-component, synthetic, grass odor blend of volatiles) elicited an even greater positive response than the visual stimuli. Changing the proportions of the four volatiles in the blend significantly reduced positive orientation responses to the stimulus source. Visual cues of wheat foliage and olfactory cues of either chopped wheat odor or the grass odor blend gave greater responses when combined than when presented separately.In flowing air or wind, nearly all insects demonstrated a rapid positive response to odors of chopped wheat and the grass odor blend, significantly greater than the response to the same stimuli in still air. However, positive responses to visual cues were not significantly greater in wind than in still air. When combined with the olfactory stimuli in flowing air, visual cues did not increase the incidence of response. Grasshoppers responding to grass odors in wind moved more rapidly and directly toward the source, and stopped less often and for shorter durations than insects responding to odor in still air or to visual cues.We conclude from these studies that M. sanguinipes adults show orientation behavior to both visual and olfactory stimuli from food plant sources, although leaf odors elicit a stronger positive response particularly when carried by wind.     

Sagebrush and grasshopper responses to atmospheric carbon dioxide concentration

Summary Seed- and clonally-propagated plants of Big Sagebrush (Artemisia tridentata var.tridentata) were grown under atmospheric carbon dioxide regimes of 270, 350 and 650 μl l⁻¹ and fed toMelanoplus differentialis andM. sanguinipes grasshoppers. Total shrub biomass significantly increased as carbon dioxide levels increased, as did the weight and area of individual leaves. Plants grown from seed collected in a single population exhibited a 3–5 fold variation in the concentration of leaf volatile mono- and sesquiterpenes, guaianolide sesquiterpene lactones, coumarins and flavones within each CO₂ treatment. The concentration of leaf allelochemicals did not differ significantly among CO₂ treatments for these seed-propagated plants. Further, when genotypic variation was controlled by vegetative propagation, allelochemical concentrations also did not differ among carbon dioxide treatments. On the other hand, overall leaf nitrogen concentration declined significantly with elevated CO₂. Carbon accumulation was seen to dilute leaf nitrogen as the balance of leaf carbon versus nitrogen progressively increased as CO₂ growth concentration increased. Grasshopper feeding was highest on sagebrush leaves grown under 270 and 650 μl l⁻¹ CO₂, but varied widely within treatments. Leaf nitrogen concentration was an important positive factor in grasshopper relative growth but had no overall effect on consumption. Potential compensatory consumption by these generalist grasshoppers was apparently limited by the sagebrush allelochemicals. Insects with a greater ability to feed on chemically defended host plants under carbon dioxide enrichment may ultimately consume leaves with a lower nitrogen concentration but the same concentration of allelochemicals. Compensatory feeding may potentially increase the amount of dietary allelochemicals ingested for each unit of nitrogen consumed.     

Parasitism of the grasshopper Melanoplus sanguinipes by a sarcophagid fly, Blaesoxipha atlanis: influence of solitary and gregarious development on host and parasitoid

Blaesoxipha atlanis (Aldrich) (Diptera: Sarcophagidae) is a common parasitoid of the grasshopper Melanoplus sanguinipes (F.) (Orthoptera: Acrididae) in western Canada. We tested the hypothesis that B. atlanis can develop as either a solitary or a gregarious parasitoid, and assessed the influence of parasitism on the growth and survival of infected grasshoppers. Males and females of M. sanguinipes were parasitized manually with one, two, or three first-instar larvae of B. atlanis in the laboratory. Parasitism was more deleterious to males than females of M. sanguinipes; females are larger than males. Host survival and longevity declined with the number of larvae per host in a sex-specific manner. In females, 39%, 24%, and 8% of hosts containing, respectively, one, two, and three sarcophagid larvae survived parasitism. Although 41% of single-parasitized males survived, all males containing more than one larva died. Variations in host quality as measured by dry mass explained much of the response to parasitism in male, but not female, hosts. Parasitoid larvae, apparently, did not cause significant physical damage to host organs and tissues but instead functioned as a metabolic sink. The greater metabolic activity associated with egg production could account for the relatively higher tolerance to parasitism of female, as opposed to male, grasshoppers. Developmental time, adult size, and percentage survival of B. atlanis declined with the intensity of parasitism, especially in parasitoids developing in male hosts. Females developing gregariously contained fewer ovarioles at eclosion than counterparts developing as solitary larvae. The mean body size of field-collected B. atlanis did not differ from that of laboratory-reared parasitoids developing singly in a host. Gregarious development is an alternative strategy to solitary development that may enable B. atlanis to maintain population numbers during periods of grasshopper scarcity.     

Responses of a generalist and a specialist parasitoid (Hymenoptera: Eucoilidae) to Drosophilid larval kairomones

Foraging parasitoids are thought to need more specific information than generalists on the presence, identity, availability, and suitability of their insect host species. In the present paper, we compare responses to host kairomones by two phylogenetically related parasitoid species that attack Drosophilidae and that differ in the width of their host range. As predicted, the behavioral response of the parasitoids to host kairomones reflected their difference in host range. The response of the specialist parasitoid Leptopilina boulardiwas restricted to contact kairomones from its natural hosts and one closely related species. In contrast, the generalist parasitoid Leptopilina heterotomaresponded to contact kairomones of a variety of Drosophilidae species.     

Hemocyte surface changes in the migratory grasshopper,Melanoplus sanguinipes in response to wounding and infection withBeauveria bassiana

Surface changes of hemocytes from the migratory grasshopper,Melanoplus sanguinipes (Fab.), following wounding and/or infection withBeauveria bassiana (Bals.) Vuillemin were assessed using fluorescein labelled wheat germ agglutinin (WGA) and concanavalin A (Con A). Binding was observed on outer wall of hemocytes when either Con A or WGA conjugates were tested. After infection there was an increase in the percentage of hemocytes which bound to WGA and which remained so for 24 h. However, after wounding this increase in binding subsided after only 1 h. Furthermore, it was found that the increase in WGA-binding occurred within the first 15 min of wounding or infection. In addition, ConA binding in males only increased 3-fold in fungus-injected insects within 1 h of treatment and remained high for 24 h. These results indicated the presence of mannose (and/or glucose) residues (Con A specificity) and N-acetyl-D-glucosamine residues (wheat germ agglutinin; WGA specificity) on outer wall surfaces of the hemocytes. The implications of these results in the immune response ofM. sanguinipes to infection withB. bassiana are discussed.     

Pathological effects of the parasite Malameba locustae in males of the migratory grasshopper Melanoplus sanguinipes and its interaction with the insecticide, cypermethrin

Gross pathological changes in Malpighian tubules of adult male Melanoplus sanguinipes (Fab.) infected by the protozoan Malameba locustae (King & Taylor) included hypertrophy, rupture of the wall, loss of pigment and presence of melanotic lesions. Infected grasshoppers also had depleted fat bodies and were hyperactive and more resistant to the insecticide, cypermethrin.

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Résumé Des Melanoplus sanguinipes adultes mâles, infectés par le protozoaire parasite, Malameba locustae mûrissent au même taux et avec le même poids que les sains. Cependant, les acridiens infectés, une semaine et deux semaines après l'aptitude à l'envol, étaient hyperactifs et significativement plus résistants à la prostration provoquée par la cyperméthrine (insecticide pyrèthroïde). Les modifications pathologiques brutes observées sur les tubes de Malpighi des acridiens infectés comprenaient l'hypertrophie, la rupture de la paroi, la perte de la pigmentation et des lésions mélaniques. Les insectes infectés ont aussi un épuisement du corps gras. Ces modifications pathologiques ne sont apparemment pas responsables de l'augmentation de la résistance à la cyperméthrine des individus infectés puisque la plus grande résistance à la prostration a eu lieu chez ceux qui ont été exposés à M. locustae et traités simultanément au Triple-Sulpha-G pour empêcher le développment du parasite.

     

The genetic consequence of differing ecological demands of a generalist and a specialist butterfly species

Many studies aim at testing the impact of recent fragmentation on the genetic diversity and connectivity of populations, while some species do exist naturally in fragmented landscapes because of their habitat requirements. Therefore, it is important to look at the genetic signatures of species occurring in naturally fragmented landscapes in order to disentangle the effect of fragmentation from the effect of habitat requirements. We selected two Nymphalid butterflies for this purpose. While Melanargia galathea is a common butterfly in flower-rich meadows, Melitaea aurelia is closely connected to calcareous grasslands, thus being historically fragmented due to its ecological demands. For the analysis of the genetic response on these opposed patterns, we analysed 18 allozyme loci for 789 individuals (399 individuals of M. galathea and 390 individuals of M. aurelia) in a western German study region with adjacent areas in Luxemburg and northeastern France. Both species showed similarly low genetic differentiations among local populations (M. galathea: F _ST 3.3%; M. aurelia: F _ST 3.6%), both combined with a moderate level of inbreeding. Isolation-by-distance analysis revealed a significant correlation for both species with similar amounts of explained variances (M. galathea: r ² = 27.8%; M. aurelia: r ² = 28.5%). Most parameters of genetic diversity were higher in M. galathea than in M. aurelia, but the latter species had a considerably higher amount of rare or locally restricted genes; the differing ecological demands are thus reflected in these differences. Both species thus seem to be genetically well suited to their respective ecological requirements. In the light of conservation genetics, we deduce that highly fragmented populations are not necessarily prone to extinction. The extinction risk might be linked to the life history of an organism and its population genetic structure.     

Performance of a generalist grasshopper on a C<Subscript>3</Subscript> and a C<Subscript>4</Subscript> grass: compensation for the effects of elevated CO<Subscript>2</Subscript> on plant nutritional quality

The increasing CO₂ concentration in Earths atmosphere is expected to cause a greater decline in the nutritional quality of C₃ than C₄ plants. As a compensatory response, herbivorous insects may increase their feeding disproportionately on C₃ plants. These hypotheses were tested by growing the grasses Lolium multiflorum C₃) and Bouteloua curtipendula C₄) at ambient (370 ppm) and elevated (740 ppm) CO₂ levels in open top chambers in the field, and comparing the growth and digestive efficiencies of the generalist grasshopper Melanoplus sanguinipes on each of the four plant × CO₂ treatment combinations. As expected, the nutritional quality of the C₃ grass declined to a greater extent than did that of the C₄ grass at elevated CO₂; protein levels declined in the C₃ grass, while levels of carbohydrates (sugar, fructan and starch) increased. However, M. sanguinipes did not significantly increase its consumption rate to compensate for the lower nutritional quality of the C₃ grass grown under elevated CO₂. Instead, these grasshoppers appear to use post-ingestive mechanisms to maintain their growth rates on the C₃ grass under elevated CO₂. Consumption rates of the C₃ and C₄ grasses were also similar, demonstrating a lack of compensatory feeding on the C₄ grass. We also examined the relative efficiencies of nutrient utilization from a C₃ and C₄ grass by M. sanguinipes to test the basis for the C₄ plant avoidance hypothesis. Contrary to this hypothesis, neither protein nor sugar was digested with a lower efficiency from the C₄ grass than from the C₃ grass. A novel finding of this study is that fructan, a potentially large carbohydrate source in C₃ grasses, is utilized by grasshoppers. Based on the higher nutrient levels in the C₃ grass and the better growth performance of M. sanguinipes on this grass at both CO₂ levels, we conclude that C₃ grasses are likely to remain better host plants than C₄ grasses in future CO₂ conditions.     

Effects of enhanced UV-B radiation on a weedy forb (Plantago lanceolata) and its interactions with a generalist and specialist herbivore

Assessments of potential impacts of global climate change often focus exclusively on plants; however, as the base of most food webs, plants generally experience abiotic stresses concomitantly with biotic stresses. Longleaf plantain, Plantago lanceolata L., is a cosmopolitan temperate perennial weed that experiences a wide range of environmental conditions throughout its range. We examined the impacts of elevated levels of exposure to shortwave (UV-B) radiation on this plant, on two herbivores associated with this plant, and on the plant-herbivore interaction. Plantains were grown at 6 and 12 kJ m^–2 d^–1 BE₃₀₀ UV-B radiation and concentrations of iridoid glycosides (aucubin and catalpol), verbascosides, and nitrogen were measured. In terms of plant impacts, we found that iridoid glycoside concentrations were unchanged by elevated UV-B radiation, whereas, in one experiment, the concentration of verbascosides in young leaves and levels of nitrogen in old leaves increased under elevated UV-B radiation. Variation in plant chemistry due to leaf age and maternal family was greater than variation due to UV-B exposure. When caterpillars were fed excised leaves from plants grown under elevated UV-B, growth and survivorship of the specialist herbivore, Precis coenia Hbn. (Lepidoptera: Nymphalidae), were unaltered and growth of the generalist herbivore, Trichoplusia ni (Hbn.) (Lepidoptera: Noctuidae), was accelerated. When the caterpillars were reared on potted plants at high and low levels of UV-B radiation, growth and survivorship of P. coenia were unchanged while growth of T. ni was significantly depressed by elevated UV-B. Elevated UV-B altered allocation patterns of above-ground biomass in these plants; masses of crowns and reproductive tissue were reduced. UV-B levels, however, did not affect distribution of damage to foliage inflicted by either species. In two additional experiments with artificial diet, designed to test the direct effect of UV-B radiation on caterpillars, growth and survivorship of P. coenia were unaltered while survivorship of T. ni was significantly depressed when caterpillars were exposed to elevated UV-B radiation. These studies collectively demonstrate that higher trophic level impacts of UV-B-induced changes in plants depend on the identity of the herbivore and its degree of adaptation not only to variation in hostplant quality but also variation in its light environment.     

Constituents of glandular trichomes of Tithonia diversifolia: relationships to herbivory and antifeedant activity

The herbivory activity of the bordered patch larvae (Chlosyne lacinia, Lepidoptera) on leaves of a Brazilian population of Tithonia diversifolia and the antifeedant potential of its leaf rinse extract were investigated. The caterpillars fed only on the adaxial face, where the density of glandular trichomes is very low, and avoided the abaxial face, which contains high levels of trichomes. Deterrent activity against the larvae was observed in leaf discs treated with leaf rinse extract at concentrations of 1-5% of fresh leaf weight. High-performance liquid chromatography (HPLC) analysis indicated that sesquiterpene lactones are the main constituents of the glandular trichomes. Dichloromethane rinse extracts of the leaves and inflorescences were chemically investigated, and 16 compounds were isolated and identified: 14 sesquiterpene lactones, a flavonoid and a diterpenoid. In this study, five sesquiterpene lactones are described for the first time in the genus, including two lactones, one of which has an unusual seco-guaianolide skeleton. Our findings indicate that the caterpillars avoid the sesquiterpene-lactone-rich glandular trichomes, and provide evidence for the antifeedant activity of the dichloromethane leaf rinse extract. In addition, a study of the seasonal variation of the main constituents from the leaf surface throughout a year demonstrated that a very low qualitative but a very high quantitative variation occurs. The highest level of the main metabolite tagitinin C was observed between September and October and the lowest was from March to June, the later corresponding to the period of highest infestation by the larvae.     

Secondary successional patterns in a sagebrush (Artemisia tridentata) community as they relate to soil disturbance and soil biological activity

The relationship between secondary succession, soil disturbance, and soil biological activity were studied on a sagebrush community (Artemisia tridentata) in the Piceance Basin of northwestern Colorado, U.S.A. Four levels of disturbance were imposed. I: the vegetation was mechanically removed and as much topsoil as possible was left; 2: the vegetation was mechanically removed and the topsoil scarified to a depth of 30 cm; 3: topsoil and subsoil were removed to a depth of 1 m, mixed and replaced; 4: topsoil and subsoil were removed to a depth of 2 m and replaced in a reverse order. Plant species composition, dehydrogenase and phosphatase enzymatic activity, mycorrhizae infection potentials, and percent organic matter were the variables measured.Treatment 4 drastically altered the pattern of vegetation succession. Treatments 2, 3, and 4 started with Salsola iberica as the dominant species but six years later, 3 and to lesser extent 2 changed in the direction of the species composition of 1, dominated by perennial grasses and perennial forbs. Treatment 4 developed a shrub dominated community. The rate of succession was not decreased by the increased levels of disturbance. Both dehydrogenase enzyme activity and mycorrhizae infection potential (MIP) increased with the change from Salsola iberica to a vegetation dominated by either perennial grasses and forbs or shrubs. The intensity of disturbance in 2, 3, and 4 reduced drastically dehydrogenase activity and MIP, but in six years they recovered to levels comparable to 1. Phosphatase enzyme activity and organic matter were unrelated to species composition but related to treatment and time elapsed. In both cases a significant decrease was observed throughout the six-year period.Nomenclature followsThis study was funded by the United States Department of Energy under Contract No. DE-AS02-76EV04018.     

The role of host species, age and defensive behaviour on ovipositional decisions in a solitary specialist and gregarious generalist parasitoid (Cotesia species)

The main objective of this study was to determine the extent to which host acceptance behaviour as related to host species, age, and defensive behaviour might explain the differences in host use that exist between two congeneric and sympatric species of parasitic wasps. Cotesia glomerata (L.) (Hymenoptera: Braconidae) is gregarious and generalist on several species of Pieridae, whereas C. rubecula (Marshall) is solitary and specific to Pieris rapae (L.). Cotesia species differed in their responses to host species (P. brassicae (L.), P. napi (L.) and P. rapae) and developmental stage (early and late 1st, 2nd and 3rd instars). In no-choice tests, host acceptance by C. rubecula was higher for p. rapae and females did not distinguish among the 6 host ages. In contrast, when foraging for P. brassicae and P. napi, C. rubecula females more readily attacked early first instar. Cotesia glomerata showed a higher degree of behavioural plasticity towards acceptance of Pieris host species and host age than did C. rubecula. Cotesia glomerata females parasitized the three Pieris species and showed higher acceptance of first and second instars over third instar. Oviposition success was also influenced by host defensive behaviour. The frequency and the effectiveness of defensive behaviour rose with increasing age of the host, P. brassicae being the most aggressive Pieris species. Furthermore, the mean duration of C. glomerata oviposition was significantly reduced by the defensive reactions of P. brassicae, which would likely affect parasitoid fitness as oviposition time is positively correlated to clutch size in C. glomerata. Acceptance frequencies corresponded well to field reports of Pieris-Cotesia associations and to patterns of parasitoid larval performance, suggesting that the acceptance phase might be used as a reliable indicator of Cotesia host-specificity.     

The role of plant genotype,environment and gender in resistance to a specialist chrysomelid herbivore

Summary Patchiness in herbivore attack is a well-documented phenomenon. When neighboring plants suffer vastly different levels of attack, then one suspects genotypic differences among plants to be the underlying mechanism. In this study, I use common garden experiments in two natural, but divergent, habitats at the Cedar Creek Natural History Area in central Minnesota to determine the role of plant genotype, environment and gender in plant resistance to a specialist herbivore. Resistance was measured by larval survivorship and weight. Eight clones ofRhus glabra were selected and 12 equal-aged ramets were dug up and planted in two gardens (each garden received 6 ramets per clone). First instarBlepharida rhois (Coleoptera: Chrysomelidae) larvae of known parentage were transferred to ramets and censused every other day. At the end of the experiment, larvae were collected and weighed. Analysis of variance was used to determine the importance of plant genotype, environment and gender on larval mortality and weight. The experiment was repeated in its entirety one month later. Both plant genotype and environment significantly affected larval survivorship in the first run of the experiment. No interactions were significant. Results from the second run indicated marginally significant genotype and environment main effects, and a genotype by environment interaction in larval survivorship. There was a significant genotype by environment interaction in larval weight on the same run. In neither run did clone gender have significant affects on resistance.     

Antioxidant responses to enhanced generation of superoxide anion radical and hydrogen peroxide in the copper-stressed mulberry plants

The aim of the study was to implicate the generation of reactive oxygen species (ROS) and altered cellular redox environment with the effects of Cu-deficiency or Cu-excess in mulberry (Morus alba L.) cv. Kanva 2 plants. A study of antioxidative responses, indicators of oxidative damage and cellular redox environment in Cu-deficient or Cu-excess mulberry plants was undertaken. While the young leaves of plants supplied with nil Cu showed chlorosis and necrotic scorching of laminae, the older and middle leaves of plants supplied with nil or 0.1 μM Cu showed purplish-brown pigmented interveinal areas that later turned necrotic along the apices and margins of leaves. The Cu-excess plants showed accelerated senescence of the older leaves. The Cu-deficient plants showed accumulation of hydrogen peroxide and superoxide anion radical. The accumulation of hydrogen peroxide was strikingly intense in the middle portion of trichomes on Cu-deficient leaves. Though the concentration of total ascorbate increased with the increasing supply of Cu, the ratio of the redox couple (DHA/ascorbic acid) increased in Cu-deficient or Cu-excess plants. The activities of superoxide dismutase (EC 1.15.1.1), catalase (EC 1.11.1.6), peroxidase (EC 1.11.1.7), ascorbate peroxidase (EC 1.11.1.11) and glutathione reductase (EC 1.6.4.2) increased in both Cu-deficient and Cu-excess plants. The results suggest that deficiency of Cu aggravates oxidative stress through enhanced generation of ROS and disturbed redox couple. Excess of Cu damaged roots, accelerated the rate of senescence in the older leaves, induced antioxidant responses and disturbed the cellular redox environment in the young leaves of mulberry plants.     

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.

     

Seasonal occurrence of specialist and generalist insect predators of spider mites and their response to volatiles from spider-mite-infested plants in Japanese pear orchards

In two adjacent Japanese pear orchards (orchards 1 and 2), we studied the seasonal occurrence of the Kanzawa spider mite, Tetranychus kanzawai, and its predators. Also the response of these predators to the volatiles from kidney bean plants infested with T. kanzawai was investigated using trap boxes in orchard 1. The mite density in orchard 1 was unimodal, with one peak at the end of August. In this orchard, population development of the specialist insect predators, Scolothrips takahashii, Oligota kashmirica benefica and Stethorus japonicus, was almost synchronized with that of the spider mites. These predators disappeared when the density of their prey became very low in mid-September. Both S. takahashii and O. kashmirica benefica abruptly increased in number in orchard 2 when the spider mite population in orchard 1 decreased. These results suggested that some of the predators migrated from orchard 1 to orchard 2. In this period, predator-traps with T. kanzawai-infested bean plants attracted significantly more S. takahashii than traps with uninfested plants. Very few individuals of S. japonicus and O. kashimirica benefica were found in the traps, despite their abundance in orchard 1. The generalist insect predator, Orius sp., was attracted to the traps throughout the experimental period irrespective of the density of spider mites, although this predator was never observed inside the orchards.     

Oviposition and reproductive performance of a generalist parasitoid (Trichogramma pretiosum) exposed to host species that differ in their physical characteristics

The response of generalist egg parasitoids to alternative natural hosts that are present simultaneously is not well known. We investigated the behavior of Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae) in relation to two field hosts Helicoverpa armigera Hübner and Spodoptera litura Fabricius, in choice and no choice tests. We quantified the effects of natal host species and post-emergence adult age on the oviposition preference of the parasitoids. H. armigera eggs were consistently preferred over S. litura eggs, regardless of the natal host and adult age. When only S. litura eggs were available as hosts, they were parasitized at statistically similar rates to H. armigera eggs (average of 17 ± 2.7 vs. 13 ± 3.0, H. armigera to S. litura). The adult lifespan and lifetime fecundity of T. pretiosum were variable but were affected by natal host species and/or host species to which they were exposed. Mean lifespan and fecundity of parasitoids that had developed in H. armigera eggs and were exposed to H. armigera eggs for oviposition were 13.9 ± 1.8 days and 98.7 ± 11.0 adult offspring. By contrast, those that developed in S. litura eggs and were exposed to S. litura eggs for oviposition lived for 7 ± 0.9 days and produced 53.8 ± 8.0 adult offspring. The ovigeny index (OI) was significantly lower in the parasitoids exposed to H. armigera eggs than in those exposed to S. litura eggs, regardless of the natal host, indicating that H. armigera eggs sustain the adult parasitoids better than S. litura eggs. These results are used to predict parasitoid behavior in the field when both hosts are available.