Can herbivore‐induced plant volatiles inform predatory insect about the most suitable stage of its prey?

The leaf beetle Plagiodera versicolora (Coleoptera: Chrysomelidae) is a specialist herbivore, all of whose mobile stages feed on the leaves of salicaceous plants. Both the larval and adult stages of the ladybird Aiolocaria hexaspilota (Coleoptera: Coccinellidae) are dominant natural enemies of the larvae of the leaf beetle. To clarify the role of plant volatiles in prey‐finding behaviour of A. hexaspilota, the olfactory responses of the ladybird in a Y‐tube olfactometer are studied. The ladybird adults show no preference for willow plants Salix eriocarpa that are infested by leaf beetle adults (nonprey) over that for intact plants but move more to the willow plants infested by leaf beetle larvae (prey) than to intact plants. Moreover, ladybird larvae show no preference for willow plants infested by leaf beetle larvae or adults over intact plants. Using gas chromatography‐mass spectrometry, six volatile compounds are released in larger amounts in the headspace of willow plants infested by leaf beetle larvae than in the headspace of willow plants infested by leaf beetle adults. In addition, the total amount of volatiles emitted from willow plants that are either intact or infested by leaf beetle adults is much smaller than that from willow plants infested by leaf beetle larvae. These results indicate that volatiles from S. eriocarpa infested by P. versicolora inform A. hexaspilota adults about the presence of the most suitable stage of their prey, whereas A. hexaspilota larvae do not use such information.     

Wheat stem sawfly‐infested plants benefit from parasitism of the herbivorous larvae

1 Parasitoids Bracon cephi (Gahan) and Bracon lissogaster Muesebeck and their herbivorous host the wheat stem sawfly Cephus cinctus Norton, a pest of wheat Triticum aestivum, were investigated for yield in T. aestivum grown in the field. 2 Wheat stem sawfly‐infested stems had a higher yield potential than uninfested stems. However, final reproductive output was not significantly different between ears on infested stems that supported complete larval development compared with ears on uninfested stems. 3 Stems containing parasitized larvae and stems containing larvae that died before completing their development had a higher mean number of seeds and seed weight, when accounting for number of fertile spikelets of each ear, than either infested with live larvae and uninfested stems. 4 The results obtained suggest that larval feeding prevented infested stems from attaining their yield potential, and that the negative impact of the pest on wheat yield was reduced when late instar sawfly larvae were parasitized. Even though some feeding occurs before parasitism, this early damage has a comparatively low impact on yield. 5 This is the first study to show a yield benefit and enhanced plant fitness due to the wheat stem sawfly parasitoids B. cephi and B. lissogaster. This results from the maintenance of increased seed number and seed weight in the higher yielding stems that are preferentially infested by this pest.     

The composite effect of transgenic plant volatiles for acquired immunity to herbivory caused by inter-plant communications

A blend of volatile organic compounds (VOCs) emitted from plants induced by herbivory enables the priming of defensive responses in neighboring plants. These effects may provide insights useful for pest control achieved with transgenic-plant-emitted volatiles. We therefore investigated, under both laboratory and greenhouse conditions, the priming of defense responses in plants (lima bean and corn) by exposing them to transgenic-plant-volatiles (VOCos) including (E)-β-ocimene, emitted from transgenic tobacco plants (NtOS2) that were constitutively overexpressing (E)-β-ocimene synthase. When lima bean plants that had previously been placed downwind of NtOS2 in an open-flow tunnel were infested by spider mites, they were more defensive to spider mites and more attractive to predatory mites, in comparison to the infested plants that had been placed downwind of wild-type tobacco plants. This was similarly observed when the NtOS2-downwind maize plants were infested with Mythimna separata larvae, resulting in reduced larval growth and greater attraction of parasitic wasps (Cotesia kariyai). In a greenhouse experiment, we also found that lima bean plants (VOCos-receiver plants) placed near NtOS2 were more attractive when damaged by spider mites, in comparison to the infested plants that had been placed near the wild-type plants. More intriguingly, VOCs emitted from infested VOCos-receiver plants affected their conspecific neighboring plants to prime indirect defenses in response to herbivory. Altogether, these data suggest that transgenic-plant-emitted volatiles can enhance the ability to prime indirect defenses via both plant-plant and plant-plant-plant communications.     

Females of Cotesia vestalis,a parasitoid of diamondback moth larvae,learn to recognise cues from aphid‐infested plants to exploit honeydew

1. To maximise their reproductive success, the females of most parasitoids must not only forage for hosts but must also find suitable food sources. These may be nectar and pollen from plants, heamolymph from hosts and/or honeydew from homopterous insects such as aphids. 2. Under laboratory conditions, females of Cotesia vestalis, a larval parasitoid of the diamondback moth (Plutella xylostella) which does not feed on host blood, survived significantly longer when held with cruciferous plants infested with non‐host green peach aphids (Myzus persicae) than when held with only uninfested plants. 3. Naïve parasitoids exhibited no preference between aphid‐infested and uninfested plants in a dual‐choice test, but those that had been previously fed aphid honeydew significantly preferred aphid‐infested plants to uninfested ones. 4. These results suggest that parasitoids that do not use aphids as hosts have the potential ability to learn cues from aphid‐infested plants when foraging for food. This flexible foraging behaviour could allow them to increase their lifetime reproductive success.     

Is selection of host plants by Plagiodera versicolora based on plant‐related performance?

Plant-related performance may be one of the most important factors in the selection of host plants by insect herbivores. We investigated the importance of plant-related performance in host selection by the willow leaf beetle, Plagiodera versicolora (Laicharting) (Coleoptera: Chrysomelidae), on four willow species: Salix chaenomeloides Kimura, Salix eriocarpa Fr. et Sav., Salix   integra Thunb., and Salix serissaefolia Kimura (Salicaceae). Bagging experiments in the field revealed that the performance of P. versicolora adults and larvae differed significantly among willow species under enemy-free conditions and at constant densities. Egg clutch and larval abundance were positively related to adult abundance. Plagiodera versicolora adults did not discriminate strongly among willow species for feeding and oviposition. Larval performance did not differ among willow species in the presence of natural enemies, suggesting that interspecific differences in host quality were overridden by mortality from natural enemies. Adult and egg clutch abundance of P. versicolora changed seasonally despite the temporal stability of adult and larval performance under enemy-free field conditions. Thus, plant-related performance of P. versicolora adults and larvae may contribute little to population growth and temporal dynamics of host use in P. versicolora . Potential factors that reduce discrimination of P. versicolora among host willow species are discussed.     

Effect of feeding by larvae of Inopus rubriceps (Diptera: Stratiomyidae) on development and growth of sugarcane.

Pot experiments were used to investigate the effect of root-feeding larvae of the soldier fly Inopus rubriceps (Macquart) on shoot production from sugarcane planting pieces (setts) and on growth and ratooning of sugarcane plants. Shoot elongation was inhibited while setts were exposed to larvae, and it resumed when larvae were removed. Infested setts produced a greater weight of roots than uninfested setts. Similar symptoms were induced by mechanical root pruning, suggesting that the effect of soldier fly larvae on setts may be a redirection of growth from the shoot to roots due to root damage. Larvae had a greater effect on shoot production at lower temperature, particularly in cultivar 'Q151', which had a higher temperature threshold than 'CP44-101'. Temperature and cultivar may influence the harmful effect of soldier fly larvae on sett germination by changing the differential rates of plant growth and larval feeding. When growing plants were exposed to larvae, the infested plants were slightly smaller at harvest and subsequently produced many fewer ratoon shoots from underground buds than uninfested plants. Shoot elongation from buds was also inhibited in setts cut from the above-ground stalks of infested plants. Analysis of nutrient levels in plants did not indicate the mechanism for ratooning inhibition, because levels of time 10 elements analyzed were at least as high or higher in infested plants. Infestation was associated with an increased level of sucrose and a reduced level of fructose in stalks. The inhibitory effect of larval feeding on ratooning was not reversed when larvae were removed from pots 10 wk before harvest. However, new stubble produced from infested plants then ratooned normally after a second harvest, provided the new roots were not attacked. The symptoms of larval feeding in growing plants are unexplained, but may be caused by the prolonged withdrawal of sap from roots or the injection of some inhibitory substance by larvae.     

An Apparent Trade-Off between Direct and Signal-Based Induced Indirect Defence against Herbivores in Willow Trees

Signal-based induced indirect defence refers to herbivore-induced production of plant volatiles that attract carnivorous natural enemies of herbivores. Relationships between direct and indirect defence strategies were studied using tritrophic systems consisting of six sympatric willow species, willow leaf beetles (Plagiodera versicolora), and their natural predators, ladybeetles (Aiolocaria hexaspilota). Relative preferences of ladybeetles for prey-infested willow plant volatiles, indicating levels of signal-based induced indirect defence, were positively correlated with the vulnerability of willow species to leaf beetles, assigned as relative levels of direct defence. This correlation suggested a possible trade-off among the species, in terms of resource limitation between direct defence and signal-based induced indirect defence. However, analyses of volatiles from infested and uninfested plants showed that the specificity of infested volatile blends (an important factor determining the costs of signal-based induced indirect defence) did not affect the attractiveness of infested plant volatiles. Thus, the suggested trade-off in resource limitation was unlikely. Rather, principal coordinates analysis showed that this ‘apparent trade-off’ between direct and signal-based induced indirect defence was partially explained by differential preferences of ladybeetles to infested plant volatiles of the six willow species. We also showed that relative preferences of ladybeetles for prey-infested willow plant volatiles were positively correlated with oviposition preferences of leaf beetles and with the distributions of leaf beetles in the field. These correlations suggest that ladybeetles use the specificity of infested willow plant volatiles to find suitable prey patches.     

Performance of cabbage stem flea beetle larvae (Psylliodes chrysocephala) in brassicaceous plants and the effect of glucosinolate profiles

The cabbage stem flea beetle (CSFB), Psylliodes chrysocephala L. (Coleoptera: Chrysomelidae), is one of the most important pests in European winter oilseed rape production. Adult beetles feed on young leaves whereas larvae mine within the petioles and stems. Larval infestation can cause significant crop damage. In this study, the host quality for CSFB of four oilseed rape (Brassica napus L.) cultivars and seven other brassicaceous species with different glucosinolate (GSL) profiles was assessed under controlled conditions. Larval instar weights and mortality were measured after 14 and 21 days of feeding in the petioles of test plants. To study the impact of GSL on the performance of larvae, the GSL contents in petioles from non-infested and infested plants were analysed before, and 21 days after, the start of larval infestation. Larval performance was not significantly different between the four cultivars of oilseed rape, but differed considerably among the other brassicaceous species tested. In comparison to the weight of larvae in the standard B. napus cv. Robust, the larval weight was higher in turnip rape (Brassica rapa L. var. silvestris) and significantly reduced in white mustard (Sinapis alba L.), oil radish (Raphanus sativa L. var. oleiformis), and cabbage (Brassica oleracea L. convar. capitata var. alba). The duration of larval development increased in white mustard and oilseed radish. The GSL profiles of the petioles showed little difference between non-infested and infested plants of oilseed rape whereas the content of aliphatic GSL increased in the infested turnip rape plants. In contrast, the aliphatic and benzenic GSL decreased in infested Indian rape (B. rapa subsp. dichotoma Roxb.). Larval weight was not correlated with the total GSL content of plants, neither before infestation nor 21 days after. Larval weight was positively correlated with progoitrin and 4-hydroxyglucobrassicin. White mustard, which provides inferior host quality for larval development, has the potential to introduce insect resistance into high-yielding oilseed rape cultivars in breeding programmes.     

Physiological response of Colorado potato beetle and beet armyworm larvae to depletion of wound-inducible proteinase inhibitors in transgenic potato plants

Larvae of Colorado potato beetle (CPB), Leptinotarsa decemlineata, and beet armyworm (BAW), Spodoptera exigua, reared on potato plants in which wound-induced accumulation of proteinase inhibitors (PIs) was largely reduced through antisense-mediated depletion of a specific lipoxygenase (LOX H3) had significantly larger weight gains than those fed on non-transformed plants. The midgut endoproteolytic activities of CPB larvae fed on non-transformed potato were significantly higher than those from larvae fed on LOX-H3-deficient plants. However, none of these proteolytic activities was inhibited by potato leaf extracts, regardless of the plant that they were fed on. Taken together, these data suggest that CPB, a leaf-feeding specialist of solanaceous plants, is largely adapted to the inducible PIs of potato, though the metabolic cost associated with the hyperproduction of digestive proteases may account for the 14-31% lower weight gain of larvae fed on non-transformed plants. The effect of LOX-H3 depletion on insect performance was more evident with larvae of the polyphagous BAW (52-63% higher weight gain and 73% higher fecundity when reared on LOX-H3-deficient plants). The poorer larval performance of BAW on non-transformed plants may be due to the susceptibility to inhibition by potato leaf tissues of most BAW digestive proteases. Indeed, BAW larvae fed on non-transformed potato showed a significant reduction in most endoproteolytic activities compared to larvae fed on LOX-H3-deficient plants, suggesting a that these insects deal poorly with induced plant defences in potato.     

Response of the elm leaf beetle to host plants induced by oviposition and feeding: the infestation rate matters

We investigated by olfactometry and feeding‐ and oviposition‐choice‐tests how the highly specialised elm leaf beetle, Xanthogaleruca luteola Müller (Coleoptera: Chrysomelidae), responds to conspecifically induced defences in the field elm Ulmus minor Miller (Ulmaceae). While egg deposition of the beetle induced elms to release volatiles attractive to the egg parasitoid Oomyzus gallerucae Fonscolombe (Hymenoptera: Eulophidae), feeding alone did not. In the present study, females of the elm leaf beetle showed preferences for the odours of twigs induced by low egg deposition and feeding over odours from uninfested twigs. In contrast, heavy infestation rendered elm odours less attractive to the beetles. Feeding and oviposition bioassays revealed an oviposition preference for leaves from uninfested twigs when compared to locally infested leaves. However, beetles preferred to feed upon systemically induced leaves compared to uninfested ones. The different preferences of the elm leaf beetle during host plant approach might be explained by a strategy that accounts for both gaining access to high quality nutrition and avoiding competition or parasitism.     

Diamondback moth females oviposit more on plants infested by non‐parasitised than by parasitised conspecifics

Abstract 1. When offered a choice, female diamondback moths (Plutella xylostella) oviposited more eggs on plants with non‐parasitised conspecific larvae than on plants with parasitised larvae. 2. The leaf area consumed by parasitised larvae was significantly lower than that by non‐parasitised larvae. However, this quantitative difference in larval damage did not explain the female’s ability to discriminate between plants with parasitised and non‐parasitised larvae, as females showed an equal oviposition preference for plants infested by higher or lower densities of non‐parasitised larvae. 3. Pupal weight and duration of the larval stage of P. xylostella were independent of whether larvae were reared on plants that were previously infested by either non‐parasitised or parasitised larvae. 4. The larval parasitoid Cotesia vestalis did not distinguish between plants infested by non‐parasitised larvae and plants infested by larvae that had already been parasitised by conspecific wasps. 5. Based on these data, it can be concluded that the moth oviposition preference for plants infested by non‐parasitised conspecifics relative to plants infested by parasitised conspecifics was not explained by plant quality or by the attractiveness of plants towards wasps. It is hypothesised that one of the reasons for this preference is avoidance of plants where a relatively high risk of parasitism is expected due to the emergence of parasitoids from the parasitised host larvae.     

Effects of larval densities and the duration since larval infestation on the host-searching behavior of Diadegma semiclausum, a parasitoid of diamondback moth larvae on plants

The host-searching behavior of Diadegma semiclausum, a parasitoid of diamondback moth (DBM) larvae, was studied in a wind tunnel. Wasps flew preferentially to a cabbage plant, freshly infested by 1 DBM larva, or one freshly infested by 10 DBM larvae, than to an intact plant. There was no significant difference between wasps?? responses to plants infested by different DBM larvae densities. We also found that the duration since the last infestation by 10 DBM larvae (1 or 3?days) negatively affected the attractiveness of infested plants. We also studied the time wasps spent searching for larvae on a cabbage plant (residence time). The wasps spent ca. 400?s on a plant freshly infested by 1 larva. Residence time was significantly longer (ca. 1,200?s) on a plant freshly infested by either 5 or 15 DBM larvae. Residence time of D. semiclausum on a plant freshly infested by 5 DBM larvae was significantly longer than on a previously infested plant (1 or 3?days after the last infestation). These results showed that host densities on a plant and the duration since their last infestation affected the host-searching behavior of D. semiclausum.     

Biotypes of the seed beetle Callosobruchus maculatus have differing effects on the germination and growth of their legume hosts

• 1 Populations of the seed beetle Callosobruchus maculatus (F.) exhibit considerable differences in body size and larval behaviour. We examined whether such variation modifies the relationship between beetle infestation and host plant performance.
• 2 Larvae from African and Asian biotypes were reared in seeds of four hosts that represented an almost four‐fold variation in seed mass. We estimated mass lost to larval consumption, and compared germination rates and seedling growth between infested and control seeds.
• 3 In seeds bearing a single larva, the larger‐bodied, contest‐competing larvae of the Asian biotype caused a 38–47% greater reduction in seed mass compared with the smaller‐bodied, scramble‐competing larvae of the African biotype. The amount of seed mass lost per larva remained similar in seeds with one or two scramble‐competing larvae but decreased significantly in seeds bearing two contest‐competing Asian larvae.
• 4 Differences in larval consumption and behaviour produced striking differences in the frequency of germination. Germination of singly‐infested mung bean (i.e. the smallest host) was 71% for African‐infested seeds versus 11% for Asian‐infested seeds. In cowpea (i.e. the largest host), 76% of Asian‐infested seeds germinated, whereas the germination rate of African‐infested cowpeas (92%) was similar to that of uninfested seeds.
• 5 Effects of beetle origin persisted after germination. Seedlings derived from Asian‐infested seeds had greater cotyledon damage 7 days after germination, and displayed lower height and less biomass 15 days after germination. Cotyledon damage was a good predictor of seedling performance (i.e. better than seed mass consumed) 15 days after germination.
• 6 Previous studies have suggested that population differences in larval size and burrowing behaviour (‘centripetal tendency’) reflect adaptation to different‐sized seeds. The present study demonstrates that these differences in turn influence the impact of larval feeding on host viability. Strong biotypic variation makes it difficult to generalize about pest impacts at the level of pest species.

     

Behavior of bollworm (Lepidoptera: Noctuidae) larvae on genetically engineered cotton

Reports of bollworm, Helicoverpa zea (Boddie), larvae feeding in white flowers of Bollgard cotton have been relatively common since the commercialization of this technology in 1996. Field studies were conducted in Louisiana to determine if differences in bollworm larval behavior occuron non-Bollgard (cultivar 'Deltapine 5415') and Bacillus thuringiensis (Bt), Bollgard ('NuCOTN 33B') cottons. Larvae were placed on the terminal foliage of either single cotton plants or on all plants within 1-m row micro-plots. On preflowering cotton plants, significantly more bollworms moved from the site of infestation (terminal) on Bollgard plants compared with that on non-Bollgard plants. On individual flowering plants, the number of nodes larvae moved from the terminal and the number of infested bolls was greater on Bollgard cotton plants. Similar differences between Bollgard and non-Bollgard plants in the percentage of infested terminals and squares were observed at 48-h after infestation when 1-m rows were infested. These data will be used to refine scouting protocols for bollworm larvae on Bollgard cotton.     

Feeding of pea leafminer larvae simultaneously activates jasmonic and salicylic acid pathways in plants to release a terpenoid for indirect defense

The pea leafminer, Liriomyza huidobrensis, is an important pest species affecting ornamental crops worldwide. Plant damage consists of oviposition and feeding punctures created by female adult flies as well as larva-bored mines in leaf mesophyll tissues. How plants indirectly defend themselves from these two types of leafminer damage has not been sufficiently investigated. In this study, we compared the indirect defense responses of bean plants infested by either female adults or larvae. Puncturing of leaves by adults released green leaf volatiles and terpenoids, while larval feeding caused plants to additionally emit methyl salicylate and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene (TMTT). Puncturing of plants by female adults induced increases in jasmonic acid (JA) and JA-related gene expressions but reduced the expressions of salicylic acid (SA)-related genes. In contrast, JA and SA and their-related gene expression levels were increased significantly by larval feeding. The exogenous application of JA+SA significantly triggered TMTT emission, thereby significantly inducing the orientation behavior of parasitoids. Our study has confirmed that larval feeding can trigger TMTT emission through the activation of both JA and SA pathways to attract parasitoids; however, TMTT alone is less attractive than the complete blend of volatiles released by infested plants.     

Nitrogen homeostasis in a willow leaf beetle, Plagiodera versicolora, is independent of host plant quality

While foliar nitrogen (N) content of host plants depends on environmental conditions, N content of herbivorous insects may remain relatively constant due to homeostasis. However, it is unknown to what extent insects can maintain their body elemental composition against natural variation in host plant quality. The present study examined the performance and N content of a willow leaf beetle, Plagiodera versicolora Laicharting (Coleoptera: Chrysomelidae), when fed leaves of host willow, Salix eriocarpa Franchet et Savatier (Salicaceae), with varying nutritional status. Water content, toughness, and N content of willow leaves varied seasonally, and they affected performance of the leaf beetle. The leaf beetle achieved high performance when fed young leaves. On the other hand, the N content of the leaf beetle changed little, and it was independent of that of willow leaves, indicating strong N homeostasis of the leaf beetle. We discussed the function of N homeostasis in herbivorous insects in tritrophic level interactions.     

Efficacy of imidacloprid for control of cereal leaf beetle (Coleoptera: Chrysomelidae) in barley

The toxicity of imidacloprid to the cereal leaf beetle, Oulema melanopus (L.), was measured under laboratory and field conditions. Insect mortality and plant damage were determined from artificial and natural infestations of O. melanopus applied to various growth stages of barley. All rates of imidacloprid formulated and applied as a seed treatment caused >90% mortality to cereal leaf beetle larvae when barley was infested with eggs at the 4-leaf stage, but were ineffective when barley was infested with eggs at the early tillering or flag-leaf stages of barley. This window of susceptibility influenced results obtained in field trials where peak larval emergence did not occur until the early tillering stage of barley. The resulting mortality in plants from treated seeds never exceeded 40% in the field. Foliar imidacloprid, however, caused >90% mortality in the field, and may be another option in the management of the cereal leaf beetle.     

Use of ablated stalks to assess field rates of parasitism of Diatraea saccharalis (Lepidoptera: Crambidae) by Cotesia flavipes (Hymenoptera: Braconidae)

We evaluated a technique that used ablated sections of corn stalks infested with larvae of Diatraea saccharalis F. to assess parasitism by Cotesia flavipes (Cameron) in a corn field. Stalkborer larvae were retrieved successfully from the artificially infested, ablated stalks for at least 96 h after deployment in the field. Levels of parasitism in ablated stalks attached to corn plants were comparable to levels of parasitism measured using whole plants that were artificially infested. Olfactometer comparisons showed preference by female C. flavipes for both whole plants and ablated stalks containing larval D. saccharalis, over uninfested stalks and plants. This technique provides a means to estimate parasitism of stalkborer larvae by C. flavipes without destructive sampling of agronomic plants.     

Leaf beetle grazing does not induce willow trichome defence in the coppicing willow Salix viminalis

Abstract 1 Willows are frequently attacked and defoliated by adult leaf beetles (Phratora vulgatissima L.) early in the season and the plants are then attacked again when new larvae emerge. The native willow Salix cinerea has previously been shown to respond to adult grazing by producing new leaves with an increased trichome density. Subsequent larval feeding was reduced on new leaves. This type of induced plant response may reduce insect damage and could potentially be utilized for plant protection in agricultural systems. 2 Here, we investigated if the willow species most commonly used for biomass production in short rotation coppice, Salix viminalis, also responds to adult beetle grazing by increasing trichome density. Larval performance and feeding behaviour on plants previously exposed to adult beetles was compared with that on undefoliated control plants in a greenhouse. 3 We found an overall decrease in trichome density within all the plants (i.e. trichome density was lower on new leaves compared to that for older basal leaves on S. viminalis). However, leaves of beetle defoliated plants had a higher trichome density compared to control plants. Larval growth and feeding was not affected by this difference between treatments. Larvae appeared to remove trichomes when feeding on S. viminalis, a behaviour that might explain the lack of difference between treatments.     

Effects of leaf pubescence in Salix borealis on host- plant choice and feeding behaviour of the leaf beetle, Melasoma lapponica

Density of leaf trichomes in Salix borealis affected both the choice of individual host plants and feeding behaviour of adults and last instar larvae of the willow feeding leaf beetle, Melasoma lapponica. Beetles clearly preferred shaved disks to unshaved ones taken from the same leaf; this preference was highest in leaves of the most pubescent plants. High leaf pubescence explained the low preference for willow clones from the high density site in among-site preference trials; shaving significantly increased the consumption of these pubescent willow clones. In no-choice experiments, the food consumption by both adults and last instar larvae decreased with an increase in leaf pubescence. The time budget of adults did not depend on leaf pubescence of the host plants, however adults compelled to feed on highly pubescent plants changed their feeding sites twice as often as on less pubescent willow clones. Larvae feeding on highly pubescent plants spend moving three times as much time as larvae feeding on less pubescent plants. Combined with our earlier observations on the increase in leaf pubescence in the year(s) following defoliation, these data suggest that leaf hairiness may have contributed to the delayed induced resistance in S. borealis by disturbing the feeding behaviour of M. lapponica.