Host-plant susceptibility to the carrot fly, Psila rosae. 1. Acceptability of various host species to ovipositing females

A preference hierarchy of the oligophagous carrot fly for 30 umbelliferous host-plant species or varieties and six non-hosts was assessed. Foliage of the test plants was presented to a laboratory population of the fly in oviposition choice experiments together with leaves of a standard plant, the susceptible carrot cultivar “Danvers”. Only two species (Anthriscus cerefolium, Carum carvi) were more acceptable than the standard plant, while about half the species received significantly fewer eggs. Some umbellifers (Anethum graveolens, Pimpinella major) had a low acceptability similar to that of non-hosts (non-umbelliferous plants). The results obtained with dual and multiple choice assays were in agreement. A multiple choice assay with potted plants yielded a similar ranking of the species as assays using detached leaves, indicating that foliage is representative for whole plants. Variability in the exploratory runs performed by the females on the leaves prior to egg-laying is described and quantified for hosts and non-hosts. Differences in post-alighting pre-ovipositional behaviour of the female flies on the leaves accounted for a major part of the variation recorded in egg-laying.     

Role of chemical cues from cotton in mediating host selection and oviposition behaviour in Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

We investigated the role of chemoreception in the host selection and oviposition behaviour of Helicoverpa armigera in the laboratory using five cotton genotypes and synthetic volatile terpenes. Female moths oviposited on substrates treated with methanol, ethanol, acetone and pentane extracts of leaves, squares and flowers of the cotton genotypes. Phytochemicals soluble in pentane were the most efficient in eliciting oviposition behaviour. In a two-way bioassay, pentane extracts of leaves or squares of a Multiple Host-plant Resistance genotype (MHR11), Deltapine commercial (DP90), and Smith Red Leaf (SRL) received significantly more eggs than solvent-treated controls. Extracts of squares of the native genotype Gossypium nelsonii did not receive more eggs. Females preferred DP90 and MHR11 to SRL and G. nelsonii. Female moths also laid more eggs on pentane extracts of MHR11 flowers than MHR11 leaves from preflowering, early flowering and peak-flowering plants. In a flight chamber, female moths used olfactory cues at short range to mediate oviposition and discrimination between host plants. Egg-laying, mated females were attracted at a distance (1.5 m) to volatile compounds released by whole plants and odours emanating from filter papers treated with synthetic volatile terpenes. Individually, the terpenes did not stimulate any significant oviposition response. However, there was a significant oviposition response to a mixture of equal volumes of the terpenes ( trans -β-caryophyllene, α-pinene, β-pinene, myrcene, β-bisabolol, and α-humulene). Conversely, antennectomised (moths with transected antennae), egg-laying, mated females did not stimulate any significant oviposition response. The significance of these findings in relation to H. armigera host-plant selection are discussed.     

Olfactory response of Trigonotylus caelestialium (Het.: Miridae) to rice plant and gramineous weeds

Abstract:  The olfactory response of Trigonotylus caelestialium , to rice, Oryza sativa L., and two species of gramineous weeds, Poa annua and Digitaria ciliaris , was investigated with an olfactometer to clarify the role of host-plant odours as olfactory cues in their host-finding behaviour. Third-instar nymphs and adult males were significantly attracted to whole plants (above ground parts) of P. annua . However, adult females were not significantly attracted to whole plants of P. annua . Attractancy of rice to T. caelestialium differed with the growth stage and part of the plant. Adult females were significantly attracted to stems and leaves in the panicle-formation stage, and panicles in the flowering stage. They were not attracted to stems and leaves in the fourth-leaf stage and flowering stage, or to panicles in the milk-and full-ripe stages. Although adult males were significantly attracted to stems and leaves in the panicle-formation stage, they were not attracted to the other rice structures tested. Both females and males were significantly attracted to stems and leaves, as well as panicles of D. ciliaris in the flowering stage. The findings suggest that T. caelestialium use host-plant volatiles as olfactory cues to find their host plants and their invasion to paddy fields were caused by olfactory responses to rice plant.     

The chemical world of crucivores: lures, treats and traps

Papilio polytes utilizes only a few plant species of Rutaceae as hosts in the field. We examined in detail the acceptability of Toddalia asiatica (a major host plant) and three other potential rutaceous hosts, Murraya paniculata, Melicope triphylla, and Phellodendron amurense, for ovipositing females of the butterfly. Female responses to the foliage, methanol extracts, and partitioned fractions from these plants were assayed for the presence of oviposition stimulants and/or deterrents. Larval survivorship on these plant species was also compared as an estimate of fitness. The foliage and a methanol extract of T. asiatica readily induced egg-laying, while females responded moderately to the foliage and a methanol extract of P. amurense. By contrast, ovipositing females only marginally accepted Me. triphylla and completely rejected Mu. paniculata. Further experiments to test the biological activity of fractions derived from the respective plant species revealed that T. asiatica contains potent oviposition stimulant(s) and that weak stimulant(s) are present in P. amurense. Poor or negative oviposition responses to both Me. triphylla and Mu. paniculata proved to be attributable to strong deterrent(s) present in these plants. Larvae performed very well on T. asiatica and P. amurense, whereas larval mortality was much higher on Mu. paniculata and Me. triphylla, suggesting the involvement of antifeedant(s) or toxic substance(s) in these plants.     

Oviposition by Mamestra brassicae (L.) (Lep., Noctuidae) in relation to age, time of day and host plant

The age at which females begin to lay, the daily pattern of oviposition, the influence of host-plant material on egg-laying as well as the host preference of Mamestra brassicae L. (Lep., Noctuidae) were investigated under laboratory conditions. The females began to oviposit during the third, fourth and fifth scotophase after emergence. A great variability in daily female fecundity was observed. The highest level of oviposition occurred during the fifth day. Females oviposited during the whole scotophase, but maximal oviposition occurred during the second hour. The females without host-plant material laid fewer eggs than females with host-plant material (cabbage leaves Brassica oleracea var. capitata ). In two-choice tests, the females preferred to oviposit on cabbage rather than chrysanthemum, but there was no difference in the mean number of eggs laid on cabbage and tomato. There were significantly greater mean numbers of total eggs deposited on tomato than on chrysanthemum.     

Foliar form, colour and surface characteristics influence oviposition behaviour of the carrot fly

Various leaf models made of paper were presented to carrot flies, Psila rosae (F.) (Diptera: Psilidae) in choice assays to investigate the effect of non-chemical plant traits on oviposition behaviour. The surrogate leaves differed in colour, shape, surface coating, size and stem length. In the presence of host-plant extracts, physical factors strongly influenced oviposition. Green, yellow and orange three-dimensional models similar in shape to host-plant leaves (pinnately or ternately compound or dissected) and with a thin cover of paraffin wax were most acceptable to the females. Egg-laying was not affected by leaf size, but was negatively correlated with stem length. The results obtained by testing models with simple leaf silhouettes were confirmed in an experiment using more lifelike imitations of real host and non-host leaves. The findings are discussed by an extensive review of similar studies in three other phytophagous fly species (cabbage root fly, onion fly, Hessian fly).     

Shaking Youngsters and Shaken Adults: Female Beetles Eavesdrop on Larval Seed Vibrations to Make Egg-Laying Decisions

Egg-laying decisions are critical for insects, and particularly those competing for limited resources. Sensory information used by females to mediate egg-laying decisions has been reported to be primarily chemical, but the role of vibration has received little attention. We tested the hypothesis that vibrational cues produced by feeding larvae occupying a seed influences egg-laying decisions amongst female cowpea beetles. This hypothesis is supported by three lines of evidence using two strains of the cowpea beetle (Callosobruchus maculatus), an Indian strain with choosy females and aggressively competing larvae and a Brazilian strain with less choosy females and larvae exhibiting an “accommodating” type of competition. First, in free-choice bioassays of seed selection, choosy Indian females selected control seeds (free of eggs, larvae, or egg-laying marker) over seeds with live larvae (free of eggs and egg-laying marker), but did not discriminate between control seeds and those with dead larvae. In contrast, less choosy Brazilian females showed no preference for seeds containing live or dead larvae over controls. Second, laser-doppler vibrometer recordings confirmed that larvae feeding inside seeds generate vibrations that are available to the female during egg-laying decisions. Third, during dichotomous choice experiments where artificial vibrations approximating those produced by feeding larvae were played back during seed selection, Indian females preferred immobile control seeds over vibrating seeds, but Brazilian females showed no preference. These results support the hypothesis that females use larval vibrations in their egg-laying decisions; whether these vibrations are passive cues exploited by the female, or active signals that ‘steer’ the behaviour of the female is unknown. We propose that vibration cues and signals could be important for host selection in insects, particularly those laying on substrates where visual or chemical cues may be unreliable. This seems to be the case with females of the cowpea beetle since visual cues are not important and chemical egg-marking does not last more than two weeks, allowing vibration cues to improve discrimination of egg-laying substrate particularly by choosy females.     

Plant-mediated indirect effects of climate change on an insect herbivore

Anthropogenic climate change is a substantial challenge to biodiversity conservation, exerting direct effects on plants and animals alike. Herbivores may be additionally affected by indirect effects, mediated through, for instance, climate change-induced alterations in host-plant quality. Thus, climate change may pronouncedly impact long-evolved plant-animal interactions, but our knowledge is still in its infancy, particularly with regard to the combined effects of temperature and water availability. We here investigate the effects of simulated climate change, considering variation in both temperature and water availability, on (1) host-plant chemistry, (2) herbivore oviposition and larval feeding preference, and (3) larval and adult performance. As study system, we used the butterfly Pieris napi (Lepidoptera: Pieridae) and its host plant Sinapis alba (Brassicacae). Host-plant chemistry was affected by simulated climate change, with higher temperatures increasing the carbon-nitrogen ratio and concentrations of glucosinolates, while drought stress led to reduced glucosinolate concentrations. Both egg-laying females and larvae preferred plants with the highest concentrations of the glucosinolate glucosinalbin, potentially acting as oviposition and feeding stimulus. Herbivore performance was positively affected by plants grown at control temperatures or under drought stress and thus reduced glucosinolate concentrations. Hence, P. napi was not able to select the most profitable host. Our study indicates that (1) climate-induced changes in plant chemistry may exert indirect effects on herbivores, (2) effects of climate change will depend on the magnitude of change in specific abiotic parameters and their interactions, whereby positive (e.g. drought) and negative (e.g. temperature) effects may even cancel out each other, and (3) changes in critical chemical cues may diminish host-plant detectability and thereby result in reduced realised fecundity. We thus highlight the important role of temperature and water availability on plant chemistry, which may change interactions between insects and plants.     

Leaf volatiles as attractants for neonate Helicoverpa armigera Hbn. (Lep., Noctuidae) larvae

The 1st instar Helicoverpa armigera larvae were bioassayed in the laboratory to study their orientational responses towards leaf volatiles of four leguminous crops: chickpea, Cicer arietinum L.; pigeonpea, Cajanus cajan Millsp.; blackgram, Vigna mungo L.; and cowpea, Vigna unguiculata L. (Walp.). The gram podborer larvae showed positive orientational responses towards leaves of all four test plants. Whole leaves of chickpea, pigeonpea and blackgram were more attractive for gram podborer larvae than cowpea whole leaves. Larval attraction for crushed (damaged) leaves of chickpea, blackgram and cowpea was significantly higher than the attraction for pigeonpea crushed leaves. The orientational responses of gram podborer larvae for crushed leaves of cowpea were significantly higher compared to whole leaves. However, the whole pigeonpea leaves elicited higher orientational responses than the crushed leaves. Maceration was not observed to affect the attractancy of chickpea and blackgram leaves. Further, the leaves were extracted in n-hexane and methanol. It was observed that the crude extracts of all the test leaves elicited positive orientational responses of larvae. In no-choice tests, the orientational preference of the larvae for the hexane extracts of all the test leaves was statistically equal. Similarly, the methanol extracts of leaves of all the test plants also attracted a greater percentage of larvae in no-choice tests. However, under two-choice bioassays, hexane foliage extract of all the test plants elicited higher orientational responses of larvae compared to the methanolic extracts of same leaves. The results of these bioassays clearly indicate that all the test leaves emit kairomones for gram podborer larvae. Moreover, kairomonal components of these leaves are, at least in part, extractable in hexane and methanol, which are higher in hexane than methanol.     

Host plant odor extracts with strong effects on oviposition behavior in Papilio polyxenes

Ovipositing females of the black swallowtail butterfly, Papilio polyxenes Fabricius (Lepidoptera: Papilionidae: Papilioninae), have an innate response to contact chemical cues, but it was not known how naïve females would respond to volatile host plant chemicals. Naïve and experienced females were tested in a free-flight experiment using model plants and volatile extracts from two hosts, carrot [ Daucus carota L. (Apiaceae)] and poison hemlock [ Conium maculatum L. (Apiaceae)], and one non-host, fava bean [ Vicia faba L. (Fabaceae)]. Naïve females responded to both host plant extracts with more approaches and landings than to non-host extracts, with poison hemlock receiving more responses than carrot. Females experienced with one of the two host plants also approached and landed frequently on extracts of host plants, but poison hemlock again received more approaches and landings, regardless of a female's host plant experience. Female P. polyxenes thus respond specifically to volatile extracts of their host plants, even in the absence of both relevant contact chemical cues and adult host plant experience.     

Putting more eggs in the best basket: clutch-size regulation in the comma butterfly

Abstract.  1. Many studies have identified different factors influencing clutch-size regulation, primarily within various groups of insects. One prediction is that ovipositing females should increase clutch size with host quality. However, in many studies it is not clear whether ovipositing females are responding to host quality or quantity.
2. Females of the polyphagous comma butterfly, Polygonia c-album (L.), were allowed to oviposit on two hosts differing greatly in quality: the preferred host, stinging nettle ( Urtica dioica L.), and the low-ranked host, birch ( Betula pubescens Ehrh). Ovipositing females were observed visually and clutch sizes were recorded. The experiment was repeated in three different years; in total, 938 observations of oviposition events were made.
3. In all three years, females ovipositing on U. dioica laid larger clutches (median 1.6–1.85) compared with females ovipositing on B. pubescens (median 1.0–1.3) . The difference was significant in two out of three years and when all three years were pooled.
4. Thus, P. c-album females exhibit clutch-size regulation, with larger clutches on better hosts. It is suggested that the proximate mechanism is likely to be a response to the same stimuli used for female ranking of host plants in the preference hierarchy.     

Particle film deters oviposition by Diaprepes abbreviatus (Coleoptera: Curculionidae)

A hydrophilic formulation of the inert silicate kaolin was tested in a screenhouse for its effect on the behavior of the root weevil Diaprepes abbreviatus (L.), a pest of citrus and ornamental plants in Florida and the Caribbean. Feeding by adults on treated foliage was reduced by 68-84% compared with adults fed untreated foliage. No insecticidal activity was detected after 14 d of exposure to kaolin-treated leaves. Oviposition was completely suppressed on treated foliage. Although females oviposited >19,000 eggs during two trials on untreated foliage, no egg masses were found on foliage treated with the kaolin formulation. These data indicate potential for kaolin as a barrier to oviposition in citrus groves and may prove to be an economically viable and environmentally sound component of an integrated approach for control of D. abbreviatus and related root weevils.     

Long-term changes in the oviposition behaviour of the cabbage butterfly, Pieris rapae, induced by contact with plants

ABSTRACT. Contact between the tarsi and cabbage foliage was found to have important influences on the oviposition behaviour of Pieris rapae , whereas contact between the ovipositor tip and cabbage foliage showed no influence. Oviposition on a leaf disc was not influenced by the presence of previously laid eggs. Gravid female P.rapae showed an increased tendency to approach and land on host or non-host foliage, or on yellow or green cards, after tarsal contact with cabbage foliage. This effect persisted for at least 72h. It was accompanied by an increased tendency to oviposit on non-host plants, most obviously within seconds of contact with cabbage, but with some persistence after 72h. Contact with a non-host leaf (lettuce) reduced the tendency of females to land on plants, but this effect was less pronounced than the one induced by contact with cabbage. These behavioural changes were immediate as well as long-term and were not associated with changes in the ovaries, so they probably result from the direct effects of sensory input on the CNS. Such behaviour probably increased the likelihood that gravid females would locate host-plants for oviposition in the field. For tests of host-plant specificity, the implications are that the behaviour of gravid females towards an array of plants might differ according to the presence or absence of hosts, not only during the test, but also during the preceding several days.     

Manipulation of oviposition patterns of the parasitoidCyzenis albicans (Tachinidae) in the field using plant extracts

We examine the oviposition behavior of the parasitic fly Cyzenis albicans(Fall.) to determine if long-distance or contact chemical cues given off by damaged oak and apple foliage influences the attack of their host the winter moth Operophtera brumata(L.). Wind-tunnel experiments indicate the presence of an attractive odor in oak leaves and the absence of an attractant in apple leaves. The application of oak foliage extracts to apple trees increased the number of parasitoid eggs laid and the level of parasitism of winter moths in the field. The impact of altered host-seeking behavior by the parastoid is discussed in the context of the population ecology of its host.     

Relationship between oviposition preference and offspring performance in Australian Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Abstract We investigated the oviposition preference and larval performance of Helicoverpa armigera under laboratory conditions to determine if the oviposition preference of individual females on maize, cowpea and cotton correlates with offspring performance on the leaves of the same host plants. The host-plant preference hierarchy of females did not correlate with their offspring performance. Female moths chose host plants that contributed less to their offspring fitness. Plant effects accounted for the largest amount of variation in offspring performance, while the effects of female (family) was low. The offspring of most females (80%, n  = 10) were broadly similar, but 20% (two out of 10), showed marked difference in their offspring performance across the host-plant species. Similarly, there was no relation between larval feeding preference and performance. However, like most laboratory experiments, our experi-mental design does not allow the evaluation of ecological factors (for example, natural enemies, host abundance, etc.) that can play an important role in larval performance in the field. Overall, the results highlight the importance of carrying out preference performance analysis on the individual or family level, rather than pooling individuals to obtain average population data.     

Preference for plants damaged by conspecific larvae in ovipositing cabbage root flies: influence of stimuli from leaf surface and roots

In laboratory dual-choice assays females of the cabbage root fly, Delia radicum, prefer for oviposition plants with roots damaged by conspecific larvae to undamaged controls. Cauliflower and kale plants were inoculated with root fly eggs (25 per plant) and the hatching larvae were allowed to feed on the roots for various periods of time (1–17 days). After 4 (cauliflower) or 5 (kale) days of larval feeding the oviposition preference was most pronounced and flies laid between 64% and 68% of their eggs near plants with damaged roots. Later, with increasing damage but fewer surviving, and thus actively feeding, larvae, the magnitude of the preference declined. The preference for plants already damaged by conspecific larvae may contribute to the previously observed aggregated distribution of D. radicum eggs in Brassica crop fields.Further experiments revealed that the sensory cues inducing this oviposition preference originate from the complex consisting of the damaged roots, the surrounding substrate (soil) and associated microbes, rather than from the aerial plant parts. In choice assays using the root-substrate complex of damaged and control plants (aerial parts removed), the observed preference for damaged roots was similar to that found for the entire plant but was more pronounced. The damaged roots alone, compared to control roots, received up to 72% (cauliflower) and 75% (kale) of the eggs. By contrast, surrogate leaves sprayed with methanolic leaf surface extracts from the most preferred plants which had been damaged were not discriminated from surrogate leaved sprayed with extracts of the respective control plants. Analysis of glucosinolate levels in methanolic leaf surface extracts revealed that root damage resulted in enhanced concentrations of indole-glucosinolates on the leaf surface in kale but not in cauliflower. Although indole-glucosinolates are oviposition stimulants for the cabbage root fly, the induced changes were apparently too small to influence oviposition behaviour.     

Genetics of host-plant preference in the comma butterfly Polygonia c-album (Nymphalidae), and evolutionary implications

In the Lepidoptera, sex-linked genes have been found to be of importance for species differences in, for example, host-plant preference, and have been implicated in ecological speciation. Variation within species is typically not sex-linked. However, in the comma butterfly Polygonia c-album (Nymphalidae) an X-linked gene has been found to play a major role in determining differences in host-plant use between two well separated populations. For this reason, we studied the role of sex-linked genes for host-plant preference within a single Swedish population of this species. Three generations of females with known pedigrees were studied in the laboratory, and they were given a choice between Urtica dioica and Salix caprea in flight cages. We found strong variation among females and significant genetic variance for host-plant preference, but no evidence for major importance of sex linkage of host-plant preference on this local scale. To what extent the observed genetic variation was due to additive genes and/or effects of major genes was not clear from the maximum likelihood analysis. In a follow-up study we sampled females over a larger area. We found strong variation among females, but not among localities, suggesting an open population structure with strong gene flow. From the combined stock, a selection experiment was performed over 2 years and six generations. The selection lines diverged after the first generation of selection and remained separate, but did not diverge further, suggesting a low degree of narrow-sense heritability and that the genetic differences may be effects of major genes. We discuss these results in relation to the possible role of genetics in the radiation of the Lepidoptera and other phytophagous insects.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 755–765.     

Dependence on host constituents controlling food acceptance by Manduca sexta larvae

The host range of Manduca sexta L. (Lepidoptera: Sphingidae) is restricted in nature to plants in the family Solanaceae. However, naive hatchling larvae often accept and continue their development on foliage from a wide spectrum of unrelated plants. In contrast, solanaceous-experienced larvae refuse to feed on other plants. Experiments were designed to explore the role of constituents of various plants in this behavioral phenomenon. Fourth instar larvae reared on solanaceous hosts: tomato (Lycopersicon esculentum Mill.), potato (Solanum tuberosum L.), or tobacco (Nicotiana tabacum L.), and the leguminous non-host cowpea (Vigna sinensis Savi), or on artificial diet based on wheat germ were tested for their acceptance or preference when offered a new diet in both choice and no-choice situations. Under no-choice conditions, acceptability of cowpea foliage was strongly affected by the larval dietary experience. Most larvae reared on solanaceous foliage did not feed, whereas those larvae reared on non-solanaceous food readily accepted cowpea. Moreover, solanaceous foliage, leaf discs and extracts were readily accepted by larvae regardless of their dietary experience. Larvae reared on any of the solanaceous plants strongly preferred cowpea discs that were treated with solanaceous extract, while larvae reared on non-solanaceous diets did not discriminate between treated and control discs. Assays of cowpea extracts as well as acceptance of cowpea discs treated with solanaceous extract indicated a lack of detectable deterrent in cowpea. Our results suggest that the mechanism for induced host specificity in M. sexta larvae involves development of a dependence on solanaceous chemical constituents. We argue that this dependence on host plant chemistry has adaptive significance.     

The effects of foods consumed after adult eclosion on the mate-searching behavior and feeding preferences of the white-spotted longicorn beetle Anoplophora malasiaca (Coleoptera: Cerambycidae)

The effects of changes in host plants on the mate-searching behavior and feeding preferences of the white-spotted longicorn beetle Anoplophora malasiaca (Thomson) (Coleoptera: Cerambycidae) were examined. All individuals were raised on the same artificial diet until they became pupae. Analysis of the mate-searching behavior of the males showed that many more newly emerged males were attracted to the odor of the artificial diet than to an unbaited control. We prepared three groups of beetles and fed each group on different host plants for one week. The host plants used included the following: an artificial diet (containing Morus alba Linné), Citrus unshiu Marc. branches, and Vaccinium spp. branches. The mate-searching behavior of the males changed in relation to the plant supplied for feeding. Simultaneously, the preference among the three host plants was tested. The newly emerged males preferred the artificial diet. After a week of feeding on one of the three plants, however, the adult males selected and consumed significantly more of the plant that they had just experienced than the other plants. These results suggest that the male mate-location cue can be acquired after adult eclosion. In addition, the male beetles are capable of changing their host-plant preference. If they do so, they use different odor cues for mate location. Newly emerged A. malasiaca females showed no preference for their first choice of food among the three host plants presented, whereas the consumption was significantly larger on C. unshiu branches. After one week of feeding on different host plants, females chose their host plant after the adult stage as well as C. unshiu, but consumed mostly C. unshiu. These results suggest that the food preferences of females are different from those of males. The behavior of females may not be affected by chemical signals from their original host-plant species (as pupae) or from the host-plant species acquired when they emerge as adults.