Diet-induced food preference by Manduca sexta larvae: acceptable non-host plants elicit a stronger induction than host plants

Diet-induced changes in food preference by fifth instar larvae of the tobacco hornworm, Manduca sexta (Johan.) (Lepidoptera, Sphingidae), were examined. Two groups of larvae with different diet experience were used: larvae reared on a host or on an acceptable non-host plant species. Each group of larvae was offered a choice between leaf discs from each rearing plant species (2-plant choice test) and food preference was measured as the consumption of one plant species relative to that of the other plant species. Diet-induced changes in preferences were larger with the host versus acceptable non-host plant pairs Solanum pseudocapsicum (L.) versus Raphanus sativus (L.), Lycopersicon esculentum (Mill.) versus Vigna sinensis (Savi), and Datura innoxia (L.) versus V. sinensis than with the host versus host plant pairs L. esculentum versus Capsicum annuum (L.) and L. esculentum versus D. innoxia. To examine how much the food preference had been altered for each test plant species alone, two other groups of larvae were offered a choice between leaf discs from a single plant species and filter paper discs laced with distilled water (1-plant choice test). Larvae preferred the dietary plant species more strongly than the non-dietary plant species in tests using the following plant species: for C. annuum with C. annuum and L. esculentum as diets, for V. sinensis with V. sinensis and L. esculentum or D. innoxia as diets, and for R. sativus with R. sativus and S. pseudocapsicum as diets. The preference for the hosts L. esculentum and D. innoxia did not change significantly after rearing larvae on different hosts or on an acceptable non-host. Thus, diet-induction by M. sexta larvae results in an enhancement of preference for the dietary plant species which is much stronger with acceptable non-hosts than with hosts.     

Role of bilateral chemosensory input in food discrimination by Manduca sexta larvae

The function of bilateral chemosensory input in food discrimination was examined using larvae of the tobacco hornworm, Manduca sexta (Johan.) (Lepidoptera: Sphingidae), in 2-choice preference tests. Larvae having three different complements of chemosensory organs remaining after microsurgery were tested: unoperated, unilaterally-ablated, and bilaterally-ablated. Discrimination between Solanum pseudocapsicum (L.) and Pelargonium hortorum (Bailey) was reduced after unilateral removal of chemosensory organs. A similar effect was found in tests offering larvae a choice between one plant species or an ethanolic extract of S. pseudocapsicum and wetted filter paper, indicating an impaired food detection. These results demonstrate that both sets of chemosensory organs are required for normal feeding decisions about these plants.Six additional plant species were each tested against wetted filter paper. The function of bilateral chemosensory input in food discrimination varies with the plant species tested. Bilateral chemosensory input is required for discrimination of Vigna sinensis (Savi) and Raphanus sativus (L.), but not for that of Lycopersicon esculentum (Mill.), Datura innoxia (L.), Brassica napus (L.), and Canna generalis (Bailey). Discrimination of V. sinensis, S. pseudocapsicum, and P. hortorum by unoperated larvae is twice as strong as that by unilaterally-ablated larvae, suggesting algebraic addition of unilateral sensory inputs. For R. sativus unilateral input does not mediate discrimination, but bilateral input does, indicating sensory processing which may be more than additive. These results show that information mediated by bilateral chemosensory organs of M. sexta is not redundant in mediating feeding decisions, but provide the necessary sensory input for normal discrimination. This suggests that, for some plant species, feeding decisions are based on both qualitative and quantitative aspects of chemosensory input.No difference in preferences between left-sided and right-sided operated larvae were observed which suggests functional duplication of each set of unilateral chemosensory organs. Possible neural mechanisms underlying feeding decisions mediated by bilateral versus unilateral chemosensory inputs are discussed.     

Morphological and functional heterogeneity in olfactory perception between antennae and maxillary palps in the pumpkin fruit fly,Bactrocera depressa

The morphology and ultrastructure of the olfactory sensilla on the antennae and maxillary palps were investigated through scanning electron microscopy (SEM) and transmission electron microscopy (TEM), and their responses to five volatile compounds were measured using electroantenogram (EAG) and electropalpogram (EPG) techniques in the pumpkin fruit fly, Bactrocera depressa (Shiraki; Diptera: Tephritidae). Male and female B. depressa displayed distinct morphological types of olfactory sensilla in the antennae and maxillary palps, with predominant populations of trichoid, basiconic, and coeloconic sensilla. Basiconic sensilla, the most abundant type of olfactory sensilla in the antennae, could be further classified into two different types. In contrast, the maxillary palps exhibited predominant populations of a single type of curved basiconic sensilla. High‐resolution SEM observation revealed the presence of multiple nanoscale wall‐pores on the cuticular surface of trichoid and basiconic sensilla, indicating that their primary function is olfactory. In contrast, coeloconic sensilla displayed several longitudinal grooves around the sensillum peg. The TEM observation of individual antennal olfactory sensilla indicates that the basiconic sensilla are thin‐walled, while the trichoid sensilla are thick‐walled. The profile of EAG responses of male B. depressa was different from their EPG response profile, indicating that the olfactory function of maxillary palps is different from that of antennae in this species. The structural and functional variation in the olfactory sensilla between antennae and maxillary palps suggests that each plays an independent role in the perception of olfactory signals in B. depressa.     

Temporal and developmental aspects of diet-induced food preferences in larvae of the tobacco hornworm, Manduca sexta

Diet‐induced changes in food preferences in an oligophagous caterpillar were studied in order to characterize the conditions under which this induction occurs. The time course of acquisition and extinction of induced food preferences by larvae of the tobacco hornworm, Manduca sexta (Johan.) (Lepidoptera, Sphingidae), was examined by varying the food and duration of larval food experience. Larvae were given feeding experience with the host plant tomato, Lycopersicon esculentum (Mill.) (Solanaceae), or the acceptable non‐host plant cowpea, Vigna unguiculata (Walp.) (Fabaceae), or switched from one to the other during different instars. Food preferences for V. unguiculata by the fourth or fifth instar larvae were measured individually in two‐choice tests with discs of V. unguiculata and moist filter paper. The acquisition or extinction of an induced food preference for V. unguiculata was indicated by larvae preferring V. unguiculata to filter paper or the reverse, respectively. Results showed that: (1) the period required for the acquisition of induced preference for V. unguiculata can be short (36 h), (2) food experience in either the third or fourth instar period is sufficient, and (3) the most recent feeding experience appears to be important. In contrast, the period required for extinction of induced preference for V. unguiculata appeared to be longer (1–3 instars), and both the most recent feeding experience and total duration of larval experience with the inducing food seem to play a role. Experience is not restricted to a particular instar period to acquire or extinguish such an induced food preference. The induced food preference for V. unguiculata was not very rigid and could be reversed by having one instar of feeding experience on L. esculentum. The findings indicate that diet‐induced food preferences in M. sexta contains elements of habituation and associative learning, but do not support food imprinting and induction of oligophagy.     

Sensilla of the antennae and the labial and maxillary palps of Blattella germanica (L.) (Dictyoptera: Blattellidae): Their classification and distribution

The various sensilla on the antennae and on the labial and maxillary palps of Blattella germanica (L.) were studied. Thick-walled chemoreceptors with fluted shafts and articulated bases are located on the antennae and on the labial and maxillary palps. Thin-walled chemoreceptors, without fluted shafts or articulated bases, are restricted to the flagellar segments of the antennae and to the distal segments of the palps. Antennae of adult males have more thin-walled chemoreceptors than do those of females. Hair-plate sensilla are found at the scape-head and scape-pedicel joints, and at the joints of segments on the palps. Campaniform sensilla are concentrated as a ring around the distal margin of the pedicel, and are also scattered singly on the scape, pedicel, and flagellar segments of the antennae, and on the first segment of the maxillary palps. Occasionally, a few sensilla coeloconica and cold receptor sensilla are found on the antennal flagellum.     

Maxillary palps can mediate taste rejection of plant allelochemicals by caterpillars

All caterpillars possess a pair of maxillary palps that “drum” the surface of foods during feeding. These chemosensory organs contain over 65% of a caterpillar's taste receptor cells, but their functional significance remains largely unknown. We examined their role in rejection of plant allelochemicals, using the tobacco hornworm (Manduca sexta) as a model insect and an extract from a plant species (Grindeliaglutinosa) as a model stimulus. We selected this system because hornworms reject foods containing Grindelia extract, and because preliminary studies indicated that their maxillary palps respond to this extract. We hypothesized that Grindelia extract elicits rejection through stimulating: (1) olfactory receptor cells, (2) taste receptor cells, (3) oral mechanoreceptors, and/or (4) a postingestive response mechanism. Our results were consistent only with hypothesis 2: caterpillars approached Grindelia-treated diets without apparent hesitation, but rejected it within 6 s of initiating biting; Grindelia-treated solutions stimulated taste receptor cells in the maxillary palp, but not the other gustatory chemosensilla; and ablating the maxillary palps eliminated rejection of Grindelia-treated diets. Our results demonstrate that taste receptor cells in the maxillary palps mediate rejection of Grindelia extract, and provide the first direct evidence for a role of maxillary palps in rejection of plant allelochemicals. Accepted: 25 January 1998     

Comparative morphology of sensilla on antenna,maxillary palp and labial palp of larvae of Eucryptorrhynchus scrobiculatus (Olivier) and E. brandti (Harold) (Coleoptera: Curculionidae)

Eucryptorrhynchus scrobiculatus (Olivier) and E. brandti (Harold) are two wood boring pests of Ailanthus altissima (Mill.) Swingle (tree of heaven) and the variety Ailanthus altissima var. Qiantouchun. These beetles attack healthy trees and bore into roots and trunks during the larval stage. We studied the typology, distribution and morphostructure of the sensilla on the antennae, maxillary palps and labial palps of E. scrobiculatus and E. brandti larvae using scanning and transmission electron microscopy. The results showed the following: (i) the antennae of the two weevil larvae had two types of sensilla, sensilla basiconica (S.b.1 and S.b.2) and sensilla twig basiconica (S.tb.1‐S.tb.3), with S.tb.4 observed only on the antennae of E. brandti larvae; (ii) the maxillary palps had three types of sensilla, S.b.2, S.tb. (S.tb.2, S.tb.3 and S.tb.5) and digitiform sensilla; (iii) the labial palps had two types of sensilla, S.b.2 and S.tb. (S.tb.2, S.tb.3 and S.tb.5); (iv) the quantity and distribution of sensilla on the antennae, maxillary palps and labial palps remained constant between E. scrobiculatus and E. brandti larvae; and (v) sensilla basiconica had distinct sidewall pores, an apical pore was observed on sensilla twig basiconica, and digitiform sensilla were oval in shape, with a distinct apical pore. Based on the microstructure of the cuticle wall and dendrite, we hypothesized that these sensilla functioned as olfactory, gustatory and hygro‐/thermo‐receptors, respectively. We discuss the relationships among types of sensilla and the types of damage caused by the larvae inside the host tree to understand olfactory and gustatory receptor mechanisms. The results of this study will provide a firm basis for future electrophysiological studies.     

Companion planting – do aromatic plants disrupt host‐plant finding by the cabbage root fly and the onion fly more effectively than non‐aromatic plants?

Brassica and Allium host‐plants were each surrounded by four non‐host plants to determine how background plants affected host‐plant finding by the cabbage root fly (Delia radicum L.) and the onion fly [Delia antiqua (Meig.)] (Diptera: Anthomyiidae), respectively. The 24 non‐host plants tested in field‐cage experiments included garden ‘bedding’ plants, weeds, aromatic plants, companion plants, and one vegetable plant. Of the 20 non‐host plants that disrupted host‐plant finding by the cabbage root fly, fewest eggs (18% of check total) were laid on host plants surrounded by the weed Chenopodium album L., and most (64% of check total) on those surrounded by the weed Fumaria officinalis L. Of the 15 plants that disrupted host‐plant finding in the preliminary tests involving the onion fly, the most disruptive (8% of check total) was a green‐leaved variant of the bedding plant Pelargonium × hortorum L.H. Bail and the least disruptive (57% of check total) was the aromatic plant Mentha piperita × citrata (Ehrh.) Briq. Plant cultivars of Dahlia variabilis (Willd.) Desf. and Pelargonium×hortorum, selected for their reddish foliage, were less disruptive than comparable cultivars with green foliage. The only surrounding plants that did not disrupt oviposition by the cabbage root fly were the low‐growing scrambling plant Sallopia convolvulus L., the grey‐foliage plant Cineraria maritima L., and two plants, Lobularia maritima (L.) Desv. and Lobelia erinus L. which, from their profuse covering of small flowers, appeared to be white and blue, respectively. The leaf on which the fly landed had a considerable effect on subsequent behaviour. Flies that landed on a host plant searched the leaf surface in an excited manner, whereas those that landed on a non‐host plant remained more or less motionless. Before taking off again, the flies stayed 2–5 times as long on the leaf of a non‐host plant as on the leaf of a host plant. Host‐plant finding was affected by the size (weight, leaf area, height) of the surrounding non‐host plants. ‘Companion plants’ and aromatic plants were no more disruptive to either species of fly than the other plants tested. Disruption by all plants resulted from their green leaves, and not from their odours and/or tastes.     

Innate and acquired components of oligophagy in the herbivorous lepidopteran,Manduca sexta

Innate and acquired aspects of oligophagy were investigated in the tobacco hornworm, Manduca sexta L. (Lepidoptera: Sphingidae), a species normally restricted to members of the family Solanaceae. Larvae were reared in the laboratory on solanaceous species tobacco (Nicotiana tabacum L.), strawberry groundcherry (Physalis pruinosa L.), and potato (Solanum tuberosum L.), as well as the non‐solanaceous foods cowpea [Vigna sinensis (L.) (Fabaceae)], rapeseed [Brassica napus L. (Brassicaceae)], and artificial diet. Feeding assays were conducted using the above plants plus corkscrew vine [Vigna caracalla (L.) Verdc. (Fabaceae)], lettuce [Lactuca sativa L. (Asteraceae)], and moist glass‐fiber filter paper. Larval feeding was characterized using two camera‐monitored assays: (1) a quantitative no‐choice disc test that determines consumption over time, delays in initiation of feeding, and % of larvae rejecting the food, and (2) a novel no‐choice emigration test that measures how soon a larva abandons a whole plant. Experimental results from both assays revealed that larvae displayed high levels of acceptance of Solanaceae regardless of whether they had been reared on solanaceous or non‐solanaceous food. We conclude that solanaceous oligophagy in M. sexta is primarily innate and does not require (but may be strengthened by) previous feeding experience on Solanaceae. In contrast, larvae tested on non‐solanaceous plants or moist filter paper showed large variation in both acceptance of foods and emigration times that were strongly dependent on the food on which they had been reared (analogous to the ‘induction of preference’ of earlier literature). Two types of induction were identified: an increase in acceptability of a plant of (1) the same species as the one on which the larva was reared, or (2) a related species. These discoveries both challenge and expand our current understanding of oligophagy in the tobacco hornworm.     

Rôle of gustation and olfaction in food plant discrimination in the tobacco hornworm, Manduca sexta

The rôle of the various chemoreceptors in behavioural discrimination among closely related food plants was studied in Manduca sexta larvae. Amputation of the three types of receptors in various combinations showed that: (1) removal of the maxillary styloconica (gustatory) causes a drastic loss of discrimination; (2) ablation of either the maxillary palpi or antennae (olfactory) also reduces discrimination, but to a lesser degree; and (3) amputation of both palpi and antennae simultaneously causes a severe discriminatory loss comparable to that of gustatory loss. We conclude that both gustation and olfaction are important for host plant discrimination. These chemosensory organs were also shown to play a strong role in the induction of preference. Unilateral extirpations of all three sense organs caused no detectable loss, demonstrating their redundancy in normal animals.     

Ultrastructure and distribution of sensilla on the maxillary and labial palps of Chlorophorus caragana (Coleoptera: Cerambycidae)

Chlorophorus caragana is a species of long‐horned beetle that damages Caragana davazamcii Sancz. (Fabales: Papilionaceae) bushes in desert areas in China. The beetles cause substantial damage to local forestry plantations and the environment. Sensilla on the maxillary and labial palps of coleopterans a allow the insects to recognize their host plants. We used scanning and transmission electron microscopy to study the ultrastructure, distribution, and abundance of various sensilla on the maxillary and labial palps of C. caragana. We found four types of sensilla including ten subtypes: one of Böhm's bristles, three of sensilla chaetica, one of digitiform sensilla, and five of sensilla twig basiconica. The types and distribution of the sensilla on the maxillary and labial palps were highly similar between males and females. Finally, this article discusses the functions of the sensilla of related species in recognizing hosts and the significance of gustation studies in the context of the control of C. caragana.     

Host plant preference and performance of the vine weevil Otiorhynchus sulcatus

Abstract 1 The relationship between reproductive performance and preference for potential host plants of the vine weevil is investigated, as shown in tests on contact (or feeding) preference, presented herein, and tests on olfactory preference, published elsewhere. 2 Assessment of reproductive performance shows that the host‐plant range of the adult vine weevil Otiorhynchus sulcatus in Europe is limited to one gymnosperm genus (Taxus sp.) and a broad range of angiosperm plants in two subclasses of the Dicotyledonae, namely Dilleniidae and Rosidae. The successful reproduction on very distantly related plant taxa suggests that the original weevil‐ and plant‐habitat has mediated the current host‐plant range of the vine weevil. 3 Contact‐preference tests with equally suitable hosts, such as Aronia, Fragaria, Euonymus and Taxus, and one less suitable host, Humulus, indicate a mismatch between contact preference and performance and, as far as olfactory preferences are known, these match neither the contact preferences nor the performance. This mismatch may arise because (i) host plant species offered do not occur in weevil habitat in Europe (e.g. Aronia and the cultivated Fragaria come from North America) and (ii) predation (or disease) risks differ among host plants, thereby altering effective reproductive performance. 4 With respect to performance on novel hosts (Thuja, Prunus) and bad hosts (Rhododendron), some between‐individual variation is found within a single population, suggesting that local populations harbour (possibly genetic) variation for adaptation to new hosts. How this variation is maintained in the face of strong selection pressures on local populations of flightless and thelytokous weevils, is an important question for understanding the broad host plant range in the vine weevil.     

Morphology of antennal,maxillary palp and labial palp sensilla in different larval instars of the Asian long‐horned beetle,Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae)

The Asian long‐horned beetle (ALB) is one of the most important international trunk‐boring insect pests of forest trees, which has caused enormous losses in the forests of China, the USA and some countries in Europe. To investigate the morphology of the antennal, maxillary palp and labial palp sensilla in the five larval instars of the ALB, scanning electron microscopy was used to determine the typology, morphology, number and distribution of the sensilla. The antennae had two types of sensilla: the sensilla twig basiconica (s.t.b.) and sensilla basiconica. Three different types of sensilla were observed on the maxillary palps, which were the sensilla digitiformia, the sensilla chaetica and the s.t.b. The labial palps had two types of sensilla: the sensilla chaetica and the s.t.b. The s.t.b. had seven total subtypes, and the sensilla basiconica had two subtypes. The typology, quantity and distribution of sensilla on the antennae, maxillary palps and labial palps remained constant in the five larval instars, but the dimensions of all sensilla types increased throughout the development. We discussed the mechanoreception and the chemoreception of ALB larvae inside the host tree to provide a theoretical understanding and information for further research on the behaviour and the electrophysiology of this devastating forest pest.     

Do grazing caddisfly larvae of Melampophylax mucoreus (Limnephilidae) use their antennae for olfactory food detection?

Abstract. The behaviour of larvae of Melampophylax mucoreus (Limnephilidae; Trichoptera) is studied to test their capacity for food detection over long distances by olfaction. In aquaria experiments with stagnant water, the larvae rapidly accumulate on a biofilm-covered stone (offered food patch) but not on control stones without biofilm-cover. Larvae in an artificial flow channel (flume), with uni-directional flow between an arrangement of stones covered with biofilm and stones without biofilm, can find food patches upstream from their location more rapidly than downstream. Stimuli transported towards the larvae by the water current might assist food location. However, no significant differences could be found in the movement of larvae with intact antennae compared with larvae without antennae. The results lead to the conclusion that the larvae of M. mucoreus find food patches mainly by random movement, which is assisted by their high mobility. Long-distance olfactory orientation towards food patches, mediated by antennae, can be precluded. Gustatory stimuli appear to play an important role in contact-chemoreception to identify biofilm patches on stone surfaces. Scanning electron microscopy studies of the larval head reveal that the antennae of M. mucoreus are small pegs (50 µm in length) with an apical plate (15 µm in diameter). The mouthparts of the larvae, especially the maxillary palps, show a variety of different sensilla types that are possibly used for contact-chemoreception. These sensilla also could be responsible for the orientation towards a food resource upstream of the larvae.     

Premating Behavior of Dealates of the Formosan Subterranean Termite and Evidence for the Presence of a Contact Sex Pheromone

Swarming by alates of the Formosan subterranean termite, Coptotermes formosanus Shiraki, occurs around dusk during April–June in Louisiana and adjoining states. Dispersal is the primary purpose for swarming. We studied the premating behavior, following swarming. Loss of wings, especially in females, was essential for successful formation of tandem pairs between the two sexes. No calling was observed and apparently a long-range volatile sex pheromone, reported in some other species of termites, may not be involved in mate attraction in C. formosanus. Males moving randomly use their antennae to detect a female, and, together with maxillary and labial palps, contact is maintained with her while in tandem. Previously mated males paired with virgin females showed tandem behavior. The incidence was very low when both males and females were previously mated and then paired. Males with complete ablation of antennae did not form tandem pairs. Ablation of 10 terminal antennal segments or the labial palps caused a significant increase in the time to initiate tandem behavior. Electrophysiological recordings from gustatory sensilla on both antennae and maxillary palps revealed increased neural activity in response to female abdominal tip extracts compared to activity elicited by a solvent control. Analysis of extracts of male and female abdominal tips using high-performance liquid chromatography showed a peak unique to the female extract. We discuss the presence of a nonvolatile chemical in female C. formosanus and its involvement in the specific premating behavior.     

The effect of different host plants on the reproduction and longevity of Nysius natalensis

Nysius natalensis Evans (Hemiptera: Orsillidae) is a pest of sunflower in South Africa. Adults invade sunflower fields from their weedy hosts which occur inside crop fields and on surrounding headlands. The host plant suitability for survival and reproduction as well as the effect of within‐generation host switching was studied on different wild host plants and sunflower. Life history parameters used to assess host plant suitability were F₁ adult survival, pre‐oviposition period, fecundity, and longevity. Nymphs and adults were provided with stems and seeds of five host plants, viz., Amaranthus hybridus L. (Amaranthaceae), Portulaca oleracea L. (Portulacaceae), Chenopodium album L. (Chenopodiaceae), Conyza albida Spreng. (Asteraceae), and sunflower, Helianthus annuus L. (Asteraceae). Nymphs were reared on crushed seed of the five plant species. After completion of the nymphal stage, emerging adults of each host plant group were provided with seed of a different host plant species for food. Adults did not survive long on stems only and very few eggs were laid. Seeds of the host plant species were shown to be an essential source of nutrients for N. natalensis reproduction, whereas the vegetative plant parts were unsuitable. Nymphal food and host‐plant switching between the nymphal and adult stages significantly affected the pre‐oviposition period. Nymphal and adult food source also affected female longevity. The number of eggs laid was not influenced by nymphal food, but was influenced by adult food and the switch between nymphal and adult food. The comparative attractiveness of sunflower and wild host plants for oviposition was also investigated and showed that females preferred to lay eggs on wild host plants, compared with sunflower. These results may explain why N. natalensis will lay their eggs on sunflower after weeds in the vicinity are controlled, or senesce toward the end of the growing season.     

Fitness consequences of host plant choice: a field experiment

Determining the relative contributions of different ecological factors for herbivore fitness is one key to understanding the ecology and evolution of host plant choice by herbivores. Natural enemies are increasingly being recognized as an important factor: host plants of inferior quality for development may still be used by herbivores if they provide enemy‐free space (EFS). Here we used the tobacco hornworm, Manduca sexta, to experimentally disentangle the effects of natural enemies from the potentially confounding factors of host plant quality, competition and microhabitat. We explored the consequences for both individual components of fitness and total fitness of M. sexta feeding on a typical high quality host plant, tobacco Nicotiana tabacum and a novel, low quality host plant, devil's claw Proboscidea louisianica in an experimental field environment in the presence of a parasitoid natural enemy, Cotesia congregata. Although early larval survival, development and growth rates, final body size and fecundity were all reduced for M. sexta feeding on devil's claw, a high rate of parasitism on tobacco and an absence of parasitism on devil's claw contributed to similar total fitness (net reproductive rate, R₀) across the two host plant species. Our results suggest M. sexta has adopted a novel host plant (devil's claw) outside its typical host range because this host plant provides enemy free space. In addition, oviposition behavior of adult female M. sexta appears to be well suited to exploiting the enemy‐free space on devil's claw; oviposition by M. sexta on devil's claw appears to correspond with seasonal variation in parasitoid abundance.     

Foodplant selection and induction of feeding preference among host and non-host plants in larvae of the tobacco hornworm Manduca sexta

Ten host plant (Solanaceae) and twelve non-host plant species were tested as foodplants for first instar larvae of the tobacco hornworm, Manduca sexta. Only nine host and three non-host plant species elicited feeding and supported growth up to fifth instar. The range of acceptability suggested that plants be divided into hosts, acceptable non-hosts, and unacceptable non-hosts. Using the two-choice feeding preference test we found that the initial preference for hosts was preserved when larvae were reared on hosts, but was less strong or absent for larvae reared on acceptable non-hosts. Thus oligophagy in the tobacco hornworm is not induced, but must be inherited.Newly-hatched first instar larvae and fifth instar larvae showed a preference hierarchy among both hosts and non-hosts. Fifth instar larvae reared separately on two different host species showed slightly different preference hierarchies among hosts. The preference for the rearing plant was increased and also two other host species changed positions in hierarchies.Feeding preferences of larvae reared on hosts or acceptable non-hosts were determined using plant combinations of host vs. host, host vs. acceptable non-host, and acceptable non-host vs. acceptable non-host. Induction of feeding preference was found in all three of these categories. This shows that induction of feeding preference in the tobacco hornworm is not restricted to host plant species.The degree to which feeding preferences were induced ranged from very strong to undetectable and dependend on the plant species paired. The strength of induction in the tobacco hornworm was found to correlate inversely with taxonomic relatedness of the plant species paired. Analysis of induction data from the literature revealed a similar correlation for other lepidopteran species.

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Résumé Vingt-deux espèces de plantes, dont 10 planteshôtes (Solanées), ont été testés comme plantes alimentaires pour des chenilles de ler stade de Manduca sexta. Sur cet ensemble, seulement 12 plantes (dont 9 plantes hôtes) induisaient la prise de nourriture et permettaient la croissance jusqu'au 5ème stade. La diversité des résultats suggère que les plantes pouvaient être classées en hôtes, non-hôtes acceptables et non-hôtes refusés. En utilisant le test du choix alimentaire préférentiel entre deux rondelles de feuilles, les chenilles néonates de ler stade ont préféré nettement les plantes-hôtes aux autres. Cette préférence initiale pour les plantes-hôtes était préservée quand les cheniles étaient élevées sur plantes-hôtes, mais devenait moins nette ou disparaissait pour des chenilles élevées sur d'autres plantes acceptées. Ainsi l'oligophagie ches M. sexta n'est pas induite, mais doit être héritée.Les chenilles néonates, aussi bien que celles de 5ème stade, présentent des préférences hiérarchisées parmi les plantes hôtes ou non. La seule frontiere nette observée était entre espèces de plantes acceptables ou non. Les hiérarchies préférentielles des chenilles du 5ème stade différaient légèrement lors-qu'elles avaient été élevées sur deux plantes-hôtes différentes. La différence essentielle était l'observation d'une préférence accrue pour l'espèce ayant servi à l'élevage, mais deux autres plantes-hôtes changaient aussi de position hiérarchique.La cause de ces changements de préférence a été approfondie, les chenilles étant élevées sur des feuilles de chaque espèce acceptable (hôte ou non). Leurs préférences alimentaires ont été définies en utilisant des combinaisons diverses (hôte x hôte, hôte x non-hôte acceptable, non-hôte acceptable x non-hôte acceptable). L'induction de la préférence alimentaires a été obtenue dans ces trois associations. Ceci montre que l'induction des choix alimentaires chez M. sexta n'est pas limitée aux plantes-hôtes. Le degré d'induction de la préférence alimentaire variait de très fort à indécelable; il dépendait de l'association examinée. La source de la variabilité de cette induction a été examinée en fonction de la relation entre la force de l'induction et les rapports taxonomiques des plantes associées. La relation obervée était inversée pour M. sexta. L'examen des données de la littérature ont révélé une relation du même type pour les autres espèces de Lépidoptères.