Leaf mechanics and herbivory defence: How tough tissue along the leaf body deters growing insect herbivores

Research on herbivory defence often focuses on leaf chemistry but less on how plant mechanical properties like leaf veins deter herbivores. Herbivores often eat tough, complex plant tissue, yet how mechanical properties affect feeding performance as the consumer grows is unclear. We measured the toughness and strength of five types of leaf tissue – the midrib, the secondary and marginal veins and the lamina inside (inner) and outside (outer) the marginal vein – in mature Eucalyptus viminalis and Eucalyptus ovata leaves with punch tests. Leaf veins were, on average, 6.2 times tougher than lamina. Marginal veins were uniformly strong and tough along the leaf body, while midribs were less strong and secondary veins less tough toward leaf tips. We correlated the force required to puncture leaf tissue with the feeding performance of a chewing insect herbivore (the spiny leaf insect, Extatosoma tiaratum (Phasmida)) across four instar stages to explore the role of tough leaf veins as potential feeding barriers. Larvae more often ate less tough leaf tips and tougher tissue as they grew. However, younger larvae were capable of penetrating the tough marginal vein when starved. We suggest tough leaf veins and consumer position along the leaf body influence insect herbivore feeding performance over their lifetime.     

Leaf injury and food consumption by larvae of Phaedon cochleariae (Coleoptera; Chrysomelidae) and Plutella maculipennis (Lepidoptera Plutellidae) feeding on turnip and radish

Larvae of Plutella maculipennis (Lepidoptera; Plutellidae) and Phaedon cochleariae (Coleoptera; Chrysomelidae) were fed on the leaves of both turnip Brassica rapa and radish Raphanus sativus. Both the weight of the food eaten and the area of leaf injured were measured. The weight eaten depended on the nutritive value of the food, whereas the area of leaf injured depended on the leaf thickness and the method of feeding of the insect. Both species developed fastest on young plants, which contained the most protein. They ate a greater weight of old than of young leaves, and injured a greater area of radish than of turnip leaf, which had more dry matter/unit area. The area of leaf injured by P. maculipennis was equivalent to the weight eaten, but P. cochleariae which had a different method of feeding destroyed much more leaf area than was expected from the weight of food eaten.

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Zusammenfassung Larven des Meerrettichkäfers, Phaedon cochleariae Fab., und der Kohlschabe, Plutella maculipennis Curtis, wurden an intakten Pflanzen von Rettich (Rhaphanus sativus L., var. French Breakfast) und Rübsen (Brassica rapa L., var. Early Milan White) gehalten. Größe und Trockengewicht der verzehrten Nahrung wurden während der gesamten Larvalentwicklung ermittelt. Die Versuche verliefen bei einer konstanten Raumtemperatur von 20°.Beide Insekten frßen eine größere Gewichtsmenge von Rübsen- als von Rettichblättern und von alten mehr als von jungen Blättern. Die mittlere Umsatzrate von P. maculipennis betrug für Rübsen und Rettich 0,41 und der Ausnutzungskoeffizient war an jüngeren Pflanzen sowohl bei Rettich wie Rübsen größer als an älteren. Die Umsatzrate von P. cochleariae variierte zwischen 0,11 und 0,27 in Abhängigkeit von Art und Alter der Nährpflanze, aber es war nicht möglich, den Ausnutzungskoeffizienten zu messen, weil der Kot nicht gesammelt werden konnte. Das Trockengewicht der von P. maculipennis gefressenen Nahrung auf den jüngsten und ältesten Pflanzen variierte bei Rübsen zwischen 10,52 und 23,77 mg und bei Rettich zwischen 9,45 und 15,28 mg; und bei P. cochleariae zwischen 12,24 und 15,70 für Rübsen und 9,79 und 12,29 für Rettich.Beide Insekten fraßen von Rettich eine größere Blattfläche als von Rübsen, wahrscheinlich weil Rübsen einen größeren Gehalt an Trockensubstanz pro Flächeneinheit enthält. Die von P. maculipennis beschädigte Blattfläche entsprach dem Gewicht der gefressenen Nahrung, da sie sauber durch das Blatt hindurch frißt und die Blattadern meidet. P. cochleariae beschädigte eine größere Blattfläche als erwartet wurde, da er die untere Blattoberfläche beschabt, oft nicht eindringt, aber die Blattadern durchnagt, so daß das Gewebe infolge Austrocknung und Ernährungsstörungen abstirbt.

     

Leaf age effects of elevated CO2-grown white oak leaves on spring-feeding lepidopterans

Folivorous insect responses to elevated CO₂-grown tree species may be complicated by phytochemical changes as leaves age. For example, young expanding leaves in tree species may be less affected by enriched CO₂-alterations in leaf phytochemistry than older mature leaves due to shorter exposure times to elevated CO₂ atmospheres. This, in turn, could result in different effects on early vs. late instar larvae of herbivorous insects. To address this, seedlings of white oak (Quercus alba L.), grown in open-top chambers under ambient and elevated CO₂, were fed to two important early spring feeding herbivores; gypsy moth (Lymantria dispar L.), and forest tent caterpillar (Malacosoma disstria Hübner). Young, expanding leaves were presented to early instar larvae, and older fully expanded or mature leaves to late instar larvae. Young leaves had significantly lower leaf nitrogen content and significantly higher total nonstructural carbohydrate:nitrogen ratio as plant CO₂ concentration rose, while nonstructural carbohydrates and total carbon-based phenolics were unaffected by plant CO₂ treatment. These phytochemical changes contributed to a significant reduction in the growth rate of early instar gypsy moth larvae, while growth rates of forest tent caterpillar were unaffected. The differences in insect responses were attributed to an increase in the nitrogen utilization efficiency (NUE) of early instar forest tent caterpillar larvae feeding on elevated CO₂-grown leaves, while early instar gypsy moth larval NUE remained unchanged among the treatments. Later instar larvae of both insect species experienced larger reductions in foliage quality on elevated CO₂-grown leaves than earlier instars, as the carbohydrate:nitrogen ratio of leaves substantially increased. Despite this, neither insect species exhibited changes in growth or consumption rates between CO₂ treatments in the later instar. An increase in NUE was apparently responsible for offsetting reduced foliar nitrogen for the late instar larvae of both species.     

Combined Effects of Host Plant Quality and Predation on a Tropical Lepidopteran: A Comparison between Treefall Gaps and the Understory in Panama

In tropical forests, light‐gaps created from treefalls are a frequent source of habitat heterogeneity. The increase in productivity, through gap formation, can alter food quality, predation and their impact on insect herbivores. We hypothesized that in gaps, herbivores would be less resource‐limited and more predator limited, whereas in the understory, we predicted the reverse. In this study, we investigate the combined effects of food quality and predation on the lepidopteran larva Zunacetha annulata feeding on its host plant Hybanthus prunifolius in two habitats; sunny treefall gaps and the shaded understory in Panama. In bioassays, Z. annulata feeding on sun leaves ate 22 percent less leaf area, grew 25 percent faster, and had higher pupal weights than larvae feeding on shade leaves. However, shade leaves had higher nitrogen content and specific leaf area. In gaps, predation was 26.4 percent compared to 13.8 percent in the understory. Larvae on understory plants traveled greater distances and spent more time searching and traveling than larvae on gap plants. These differences in behavior are consistent with lower predation risk and lower quality food in the understory. Using data from bioassays and field experiments we calculated 0.22 percent and 1.02 percent survival to adulthood for larvae in gaps and the understory, respectively. In conclusion, although these habitats were in close proximity, we found that larvae in the understory are more resource‐limited and larvae in gaps are more predator limited.     

Resistant cabbage cultivars change the susceptibility of Plutella xylostella to Bacillus thuringiensis

1 Laboratory studies demonstrated that the susceptibility of larvae of the lepidopteran crucifer pest Plutella xylostella to the insect pathogen Bacillus thuringiensis (Bt) was influenced by the host plant. 2 Larvae reared on the resistant cabbage cultivars Minicole F₁ and Red Drumhead were significantly more susceptible to Bt (the LC₅₀ fell to one half) than larvae fed leaves of susceptible cultivars. 3 However, a third resistant cultivar, Aquarius F₁, had no synergistic effect on Bt‐related mortality. 4 Actual uptake of Bt was monitored in the bioassays, as a preliminary experiment showed that the plant resistance reduced consumption of Bt‐treated leaf discs. However, differences in feeding rate did not explain the observed differences in mortality.     

Feeding of fall armyworm,Spodoptera frugiperda,on Bt transgenic cotton and its isoline

Studies on insect food intake and utilization are important for determining the degree of insect/plant association and host species’ resistance, and also for helping design pest management programs by providing estimates of potential economic losses, techniques for mass breeding of insects, and identifying physiological differences between species. We studied the feeding and development of fall armyworm, Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae), on transgenic (Bt) and non‐transgenic (non‐Bt) cotton. The larvae of S. frugiperda fed on Bt cotton had a longer development period (23.0 days) than those fed on non‐Bt cotton (20.2 days). Survivorship of S. frugiperda larvae fed on Bt cotton (74.1%) was lower than that of larvae fed on non‐Bt cotton (96.7%). Pupal weight of larvae fed on Bt cotton (0.042 g) was lower than that of larvae fed on non‐Bt cotton (0.061 g). The cotton cultivar significantly affected food intake, feces production, metabolization, and food assimilation by S. frugiperda larvae. However, it did not affect their weight gain. Intake of Bt‐cotton leaf (0.53 g dry weight) per S. frugiperda larva was lower than the intake of non‐Bt‐cotton leaf (0.61 g dry weight). Larvae fed on Bt‐cotton leaves produced less feces (0.25 g dry weight) than those fed on non‐Bt‐cotton leaves (0.37 g dry weight). Weight gain per S. frugiperda larva fed on Bt‐cotton leaves (0.058 g dry weight) was similar to the weight gain for larvae fed on non‐Bt‐cotton leaves (0.056 g dry weight). The cotton cultivar significantly affected the relative growth, consumption, and metabolic rates, as well as other nutritional indices: the figures were lower for larvae fed on Bt‐cotton leaves than for larvae fed on non‐transgenic cotton leaves.     

Suitability of Eucalyptus and Corymbia for Mnesampela privata (Guenée) (Lepidoptera: Geometridae) larvae

Abstract 1 Mnesampela privata (Guenée) has a host list of 40 Eucalyptus and at least one Corymbia species. Larval survival and performance was studied on 19 species to investigate how certain leaf traits influence the suitability of different species. 2 After 7 days, survival on Eucalyptus aggregata and Eucalyptus camphora is greater than 70% even though the toughness of leaves is 0.15–0.19 mg/mm². However, after the same time, survival on genotypes of Eucalyptus melliodora and Eucalyptus sideroxylon was less than 60%, or even 0%, even though the toughness of some leaves was as low as 0.11 mg/mm². An unmeasured allelochemical, rather than toughness, may reduce survival on these species. 3 Dry weights of first‐instar larvae were negatively correlated with leaf toughness for 13 of the species studied. Species that produced the heaviest first‐instar larvae were not the same hosts that produced the heaviest second‐instar larvae. 4 Dry weights of female pupae were negatively correlated with total oil content for five of the species studied. 5 Larvae exhibit age‐related changes in feeding behaviour. Neonates skeletonize leaves (avoid leaf veins and oil glands) and post‐third‐instar larvae ingest whole leaf fragments (consume small leaf veins and oil glands). These findings suggest that neonates are sensitive to high leaf toughness and non‐oil plant secondary metabolites whereas older larvae are less sensitive to high leaf toughness and are likely to become larger adults on hosts with lower oil contents.     

Feeding responses of adapted and non-adapted insects to the defensive properties of Baccharis halimifolia L. (Compositae)

Summary Trirhabda bacharidis (Weber) (Chrysomelidae), a univoltine, monophagous beetle is the dominant herbivore on Baccharis halimifolia (Compositae), a woody, perennial shrub that leafs out in early spring and retains its leaves into November. Available plant biomass increases during the season but T. bacharidis feeds only during spring and early summer. During the remainder of the growing season, there are no major herbivores feeding on B. halimifolia. B. halimifolia leaves increase in toughness and thickness and decrease in moisture and nutrients as the season progresses. In feeding preference tests, T. bacharidis larvae preferred young leaves over leaves of older age classes. When reared on leaves of different ages, larvae fed on young leaves, weighed more, pupated earlier and had greater survivorship. T. bacharidis larvae showed no significant feeding preference for similarly tough B. halimifolia leaves painted with differing concentrations of late season acetone leaf extract. Elm leaves painted with the same leaf extracts were avoided by larvae of the non-adapted specialist Pyrrhalta luteola (Mueller) (Chrysomelidae).These results suggest that the adapted specialist, T. bacharidis, is not deterred by the B. halimifolia acetone soluble secondary chemical which increases in amount over the season. Decreasing leaf nitrogen (perhaps in concert with increasing leaf toughness) seems to be the primary factor that dissuades its feeding. However, acetone soluble secondary chemicals in the leaves of B. halimifolia may be effective in preventing herbivory from non-adapted insects.     

Self-selection and utilization efficiency of Heliothis zea larvae feeding on seeds and the walls of snap bean pods

First instar Heliothis zea larvae tended to feed on the leaves of snap beans, but later instars preferred to burrow in the pods. Fifth instars offered only leaves grew poorly because they ate them in small quantities, presumably because of deterrency. The utilization efficiency and growth of fifth instars fed pulp was slightly but significantly inferior to that of seed-fed larvae but was greatly superior to that of leaf-fed larvae. Since the dry matter content of pulp is less than one-third that of seeds, larvae allowed to eat only pulp had abnormally large fresh weight intakes and devoted about three times as much time to feeding as did seed-fed larvae. Larvae offered a mix of seeds and pulp ate both and had a total fresh weight food intake somewhat greater than that of seed-fed larvae but much less than that of pulp-fed larvae. Eating both seeds and pulp offers no obvious nutritional advantage but presumably offers a significant ecological advantage. It is argued that the number of foraging trip a larva must make to discover pods is minimized by feeding on a mix, thus reducing exposure to mortality factors such as predation.     

Movement and feeding patterns of Epilachna cucurbitae Richards (Coleoptera:Coccinellidae) on pumpkin and zucchini plants2

The oviposition patterns of adults and the movement and feeding patterns of larvae of Epilachna cucurbitae on two species of cucurbits, Cucurbita maxima cv Queensland Blue and C. pepo cv Blackjack, were studied in the field and laboratory. The physical and nutritional characteristics of host plant leaves of different ages were described. Younger leaves had higher nitrogen contents but were less abundant, smaller and had higher trichome densities than older leaves. The development of first instar larvae was delayed by the leaf hairs on young and mature pumpkin leaves which prevented larvae from reaching the leaf surface to feed First instal larvae developed more quickly on leaves rich in nitrogen. Neither the total developmental time of larvae nor the size of pupae was affected by leafage because larvae on poor quality leaves compensated by eating more. Female beetles oviposited on all but the youngest and oldest leaves of the host plant. The trichomes on young leaves prevented females from attaching eggs to the leaf surface. First instar larvae remained where they hatched, but older larvae were more mobile, Changing feeding sites frequently and moving progressively to younger, more nutritious leaves. Final instar larvae moved onto adjacent vegetation to pupate. The adaptive significance of these patterns is discussed in relation to the nutritional value, hairiness and abundance of host plant leaves of different ages and the physical limitations of different larval instars.     

Diamondback moth oviposition: effects of host plant and herbivory

The oviposition behaviour of Plutella xylostella L. (Lepidoptera: Plutellidae) on Chinese cabbage (Brassica rapa L. Pekinensis, cv. Wombok), canola (Brassica napus L. cv. Thunder TT), and cabbage (Brassica oleracea L. Capitata, cv. sugarloaf) (Brassicaceae) was studied in the laboratory. In no‐choice experiments moths laid most eggs on the stems and lower three leaves of cabbage plants, the lower three leaves of canola plants, but on the upper three leaves of Chinese cabbage plants. The effects of conspecific herbivore damage to foliage could be replicated by mechanical damage. When foliage was damaged, injured cabbage and canola plants were preferred for oviposition over intact conspecifics, whereas injured Chinese cabbage plants were less preferred than intact conspecifics. However, when root tissue was damaged, intact cabbage and canola plants were preferred over injured conspecifics, whereas moths did not discriminate between root‐damaged and intact Chinese cabbage plants. Injury to upper leaves significantly affected the intra‐plant distribution of eggs. In cabbage and canola plants, injury to leaf 6 significantly increased the number of eggs laid on this leaf, resulting in a significant decrease in the number of eggs laid on the lower foliage/stem of plants, whereas in Chinese cabbage plants it significantly decreased the number of eggs laid on leaf 6. Following oviposition on intact plants, neonate larvae established the vast majority of feeding sites on leaves 5–8 in all three host plants, indicating that larvae moved a considerable distance from preferred oviposition sites in cabbage and canola plants. The growth rate of neonates fed on leaf‐6 tissue was significantly greater than that of those fed on leaf‐1 tissue; >90% of larvae completed development when fed exclusively on leaf‐6 tissue but no larvae completed development when fed exclusively on leaf‐1 tissue. The study demonstrates the complex and unpredictable interactions between P. xylostella and its host plants and provides a basis from which we can begin to understand observed distributions of the pest in Brassica crops.     

Do birch-feeding caterpillars make the right feeding choices?

Summary The effects of leaf quality on caterpillar performance are frequently investigated, as are the feeding preferences of caterpillars, but rarely are the two considered together to see how preferences match performance. I looked at the preferences and performance of the specialist feeder Achlya flavicornis and the generalist Spilosoma luteum on Betula pendula leaves which were artificially damaged, mined or left undamaged, and on leaves of different ages. Often feeding preferences were not for the leaf types that gave faster development, larger pupae or greater survival. A. flavicornis preferred medium aged to older leaves though medium aged leaves produced smaller pupae. They did not discriminate between young and medium aged leaves, though larvae fed young leaves developed more slowly, and had lower survival and lower adult emergence. S. luteum larvae did not discriminate between damaged and undamaged leaves, though feeding on damaged leaves gave smaller pupae and lower adult emergence. I suggest that generalists like S. luteum might be unable to make feeding choices between leaves that are on the same tree and vary only slightly because populations are exposed to much greater variation across the wide range of potential food plants individuals may encounter. Possible differences in within-plant feeding selectivity between generalists and specialists should be investigated. A. flavicornis is a leaftier for which selection of a suitable leaf for tying could be more important in the first instance than feeding considerations. Larvae preferred younger to older leaves for tying, and larvae tended to feed on the tied leaves, so tying preferences explain the initial feeding preferences.     

Patterns of growth,reproduction, defense,and herbivory in the dioecious shrub Baccharis halimifolia (Compositae)

Summary Patterns of growth, reproduction, defense (leaf resin) and herbivory were compared between the sexes of the dioecious shrub Baccharis halimifolia (Compositae). Male plants possessed longer shoots and more tender leaves, grew faster, and flowered and senesced earlier than female plants. Levels of leaf nitrogen, water content, and acetone-soluble resin (shown to deter feeding by polyphagous insect herbivores) did not differ between male and female plants. When offered a choice between leaves from male and female plants, adults of two leaf beetles (Chrysomelidae), the monophagous Trirhabda bacharidis and the polyphagous Paria thoracica, both preferred to feed on male leaves. Similarly, the daily fecundity of older females of T. bacharidis was higher when they were fed leaves from male compared to female plants. However, adult survivorship and total fecundity of T. baccharidis did not differ between male and female leaf treatments. We attribute the feeding preference for and slight increase in fecundity on male plants to the tenderness of male leaves. Larvae of the fly Tephritis subpura (Tephritidae) fed exclusively in the sterile receptacle of male flower heads (85% infested), but the phenology was such that pollen production was not adversely affected. Larvae of two other flies Dasineura sp. and Contarinia sp. (Cecidomyiidae) occupied >95% of only female flower heads where they fed among and on the developing seeds. We conclude that foliage-feeding herbivores are unlikely candidates to explain the female-biased sex ratio (59% female) of B. halimifolia plants in the field, and that their preference for male plants is a result of plant characteristics (e.g. rapid growth) that have been selected by some other factor. However, our data on selective floral herbivory in B. halimifolia are in accord with the argument that dioecy reduces the inadvertent loss of flower parts of one sex when herbivores feed on flower parts of the opposite sex.     

Onshore–offshore variations of copepod community in northern South China Sea

Previous studies which have tested the feeding preferences of shredders for fungal species and the food quality of fungi used detritus uniformly colonized by a fungus, which is not the case for decaying leaves in streams. It is not known whether shredders in different development stages exhibit variations in feeding preference and larval performance. This study examined the feeding preferences and the growth of the third and the fifth instars of Pycnopsyche gentilis larvae using fungal-colonized patches and whole leaves, respectively, having different fungal species compositions (Alatospora acuminata, Anguillospora filiformis, Articulospora tetracladia, Tetrachaetum elegans, and all species combined). The aquatic hyphomycetes used were co-dominant on leaves in the stream inhabited by the caddisfly. During 14 d of feeding, the larvae of both instars did not show significant differences in feeding preferences for the patches growing on oak leaves, although the third instar larvae were slightly more selective than the fifth instar larvae. When fed with maple leaves for 18 d, larval growth rates, gross growth efficiencies, and survivorship were not significantly different among the fungal treatments. However, the larval growth of both instars fed with fungal-colonized leaves was always significantly greater than the growth of larvae fed with diets of uncolonized leaves. The third instar larvae grew faster than the fifth instar larvae, but the growth efficiencies of the two instars were similar. These results suggest that P. gentilis larvae exhibit less selectivity in their feeding than other caddisfly shredders that have been examined and that the dominant fungi colonizing leaves in their habitat are similar in palatability and food quality for this shredder. Handling editor: B. Oertli     

Host Plant Quality Factors That Influence the Growth and Development of Oxyops vitiosa, a Biological Control Agent of Melaleuca quinquenervia

The Australian weevil Oxyops vitiosa was released in 1997 in Florida as a biological control agent of Melaleuca quinquenervia. The larvae of this agent are flush-feeders, found only on the growing tips of their host. Knowledge of this restriction to feeding on the growing tips and other nutritional requirements may assist in the establishment and dispersal of this species. Therefore, O. vitiosa survival was assessed when neonates were fed M. quinquenervia leaves from branches that had dormant buds or emerging bud leaves. Additionally, the influence of leaf quality from different sites and within sites was determined by the feeding of neonates emerging bud leaves collected at three sites and from three leaf qualities (poor, intermediate, and high). Within-site leaf qualities were described in the field by leaf color and in the laboratory by percentage dry mass and nitrogen. Larval survival was lowest when fed leaves from branches that had dormant buds. Associated with this low survival were high leaf toughness and percentage dry mass. When larvae were fed emerging bud leaves, most of the variation in larval survival and performance was attributed to differences in within-site plant quality. Generally, the highest-quality leaves had relatively low percentage dry mass and high percentage nitrogen. Larval survival generally decreased when fed the poor-quality leaves, and in one site, the intermediate-quality leaves. Larvae required less time to develop to adults when fed the high-quality leaves. Development time increased in females but not in males when the larvae were fed the poor-quality leaves. Adult biomass of both females and males generally increased when the larvae were fed the high-quality leaves from two of the three sites. The results indicate that the larvae of O. vitiosa are restricted to feeding on flush foliage with low toughness. Additionally, variations in foliar percentage dry mass and nitrogen influence larval survival and performance. This knowledge benefited the development of mass-production nursery sites and the selection of suitable release sites, which facilitated the establishment of this biological control agent.     

Differences in parasitic ability of Beauveria bassiana (Moniliales: Moniliaceae) in relation to Pantana phyllostachysae (Lepidoptera: Lymantridae) when feeding on different Phyllostachys pubescens (Poales: Poaceae) leaves

Differences in composition of on‐year, off‐year and new leaves in Phyllostachys pubescens (moso bamboo) forests will affect the resistance of Pantana phyllostachysae larvae and the parasitic ability of Beauveria bassiana. Pantana phyllostachysae fed on mixed leaves of on‐year, off‐year and new bamboo leaves. The activities of acetylcholine esterase (AChE) and glutathione S‐transferase (GSTs) in P. phyllostachysae larvae fed on mixed leaves were significantly higher, greatly reducing the pathogenicity of B. bassiana to P. phyllostachysae larvae compared with the treatments of off‐year and new bamboo leaves. The phenoloxidase (PO) activity in P. phyllostachysae larvae feeding on mixed leaves varied within three days after parasitization by B. bassiana, which indicated higher and less changeable PO activity compared with treatments of off‐year and new bamboo leaves. Inclusion of new leaves differed greatly due to its growth, in which, both AChE and GSTs activities were higher than the off‐year leaf treatment, leading to a lower pathogenicity of B. bassiana to P. phyllostachysae larvae feeding on new leaves compared with off‐year leaves. On the contrary, the pathogenicity of B. bassiana to P. phyllostachysae larvae feeding on on‐year leaves was lower than other treatments. AChE and GSTs activities in P. phyllostachysae larvae were lower and PO activity was less changeable, which might be related to the disadvantage of the larval surface structure to B. bassiana infection in the on‐year leaf treatment.     

Effects of elicitation treatment and genotypic variation on induced resistance in Populus: impacts on gypsy moth (Lepidoptera: Lymantriidae) development and feeding behavior

We examined the effects of various wounding treatments and genotypic variation on induced resistance in Populus (Salicales: Salicaceae) against herbivory by the gypsy moth, Lymantria dispar L. (Lepidoptera: Lymantriidae). Second-instar larvae grew and consumed less on leaves from induced than non-induced trees. Likewise, larvae preferred leaf disks from non-induced trees. Among induction treatments, gypsy moth feeding had the strongest and most consistent effect in behavioral choice tests. Mechanical wounding of leaves and mechanical wounding plus application of gypsy moth regurgitant had intermediate effects, while application of jasmonic acid had the weakest overall effect. Under no-choice conditions, there were no consistent trends across clones in the ability of various treatments to elicit plant responses affecting the herbivore. Levels of constitutive and inducible resistance to herbivory varied significantly among 12 Populus clones. Larvae grew up to 30-fold more, and consumed up to 250-fold more on the most suitable than the least suitable clone. Prior feeding by gypsy moths reduced larval feeding up to 71.4% on the most highly inducible clone, but it had little or no effect for the least inducible clones. There was no evidence for a relationship between levels of inducible and constitutive resistance, or between inducible resistance and phylogenetic relatedness among clones. We discuss implications for the ecology and evolution of plant-insect interactions and the management of insect pests. Received: 12 October 1998 / Accepted: 22 March 1999     

Temporal dynamics of herbivore‐induced responses in Brassica juncea and their effect on generalist and specialist herbivores

Herbivore feeding may induce an array of responses in plants, and each response may have its own temporal dynamics. Precise timing of these plant responses is vital for them to have optimal effect on the herbivores feeding on the plant. This study measured the temporal dynamics of various systemically induced responses occurring in Brassica juncea (L.) Czern. (Brassicaceae) leaves after insect herbivory in India and The Netherlands. Morphological (trichomes, leaf size) and chemical (glucosinolates, amino acids, sugars) responses were analysed. The effects of systemic responses were assessed using a specialist [Plutella xylostella L. (Lepidoptera: Plutellidae)] and a generalist [Spodoptera litura Fabricius (Lepidoptera: Noctuidae)] herbivore. We tested the hypotheses that morphological responses were slower than chemical responses and that generalist herbivores would be more affected by induced responses than specialists. Glucosinolates and trichomes were found to increase systemically as quickly as 4 and 7 days after herbivore damage, respectively. Amino acids, sugars, and leaf size remained unaffected during this period. The generalist S. litura showed a significant feeding preference for undamaged leaves, whereas the specialist herbivore P. xylostella preferred leaves that were damaged 9 days before. Performance bioassays on generalist S. litura revealed that larvae gained half the weight on leaves from damaged plants as compared to larvae feeding on leaves from undamaged plants. These studies show that although morphological responses are somewhat slower than chemical responses, they also contribute to induced plant resistance in a relatively short time span. We argue that before considering induced responses as resistance factors, their effect should be assessed at various points in time with both generalist and specialist herbivores.     

Codling moth granulosis virus: effects of its use on some other orchard arthropods

In four field trials from 1978 to 1980, sprays of codling moth granulosis virus (CpGV) plus 1·0% skimmed milk powder did not significantly affect damage to fruit by leaf rollers (tortrix moths). In laboratory tests, survival of larvae of the leaf roller Archips podana fed on leaves sprayed with CpGV plus milk was unaffected and they grew faster than on unsprayed leaves, because of the milk deposits. This might increase damage by A. podana if CpGV plus milk were applied during the feeding period of this species. In one field trial an unusual infestation of fruit by larvae of pith moth Blastodacna atra was not affected by CpGV. Azinphos-methyl significantly reduced damage by B. atru and, in one field trial where sprays were correctly timed, that by leaf rollers. CpGV had no consistently significant effects on numbers of fruit tree red spider mite Panonychus ulmi or its predators, whereas azinphos-methyl induced outbreaks of P. ulmi by killing its predators.