Interspecific egg load assessment of host plants by Pieris rapae butterflies

Egglaying responses of Pieris rapae L. butterflies to the oviposition deterring pheromone (ODP) of Pieris brassicae L. were studied in the laboratory. Choice experiments with ODP treated leaves and control leaves revealed that females perform a strong preference to lay their eggs on the control leaves. This preference is maintained even when during the experiment the control leaf becomes covered with a large number of conspecific eggs. Choice experiments with cabbage leaves with and without P. rapae eggs seem to indicate the absence of intraspecific egg load assessment of host plants in P. rapae. The deterrent effect of the ODP of P. brassicae to P. rapae females persists for at least 8 days. Behavioural observations suggest olfactory hairs as well as gustatory hairs to be involved in the perception of the ODP but electrophysiological recordings of the various chemoreceptors are necessary to confirm this. Finally the prospects of application of this pheromone/kairomone in cabbage pest control are discussed.

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Appréciation de la charge interspécifique en oeufs sur la plante hôte par Pieris rapae

Résumé La réponse au laboratoire de P. rapae à la phéromone dissuadant la ponte (ODP) de P. brassicae a été étudiée par l'oviposition. Des expériences de choix entre des feuilles traitées à l'ODP et des témoins ont montré que les femelles préfèrent nettement les feuilles témoins. Cette préférence s'est maintenue même quand les feuilles témoins ont été recouvertes d'un grand nombre d'oeufs de P. rapae. Ceci peut indiquer l'absence chez P. rapae d'une évaluation de la charge de ses propres oeufs. L'effet dissuadant du ODP de P. brassicae sur les femelles de P. rapae persiste au moins 8 jours. Les observations comportementales suggèrent que des poils olfactifs aussi bien que des poils gustatifs sont impliqués dans la perception d'ODP mais une confirmation de cette hypothèse par enregistrements électrophysiologiques est nécessaire. Les perspectives d'utilisation de cette phéromone/kairomone dans la lutte contre les insectes du chou sont examinées.

     

The role of sense organs in the feeding behaviour of Chinese perch

Experiments were conducted to identify the roles of the individual sense organs in the feeding behaviour of Chinese perch Siniperca chuatsi by determining the consumption of natural food after selective removal or blocking of eyes, lateral lines and olfactory organs, and also by observing the behavioural response to visual, mechanical and chemical stimulation by artificial prey. Chinese perch were able to feed properly on live prey fish when either eyes or lateral lines were intact or functional, but could scarcely feed without these two senses. Chinese perch recognized its prey by vision through the perception of motion and shape, and showed a greater dependence on vision in predation when both visual and mechanical cues were available. Chemical stimulation by natural food could not elicit any feeding response in Chinese perch, and gustation was only important to the fish for the last stage of food discrimination in the oropharyngeal cavity. The sensory basis of Chinese perch in feeding is well adapted to its nocturnal stalking hunting strategy, and also explains its peculiar food habit of accepting live prey fish only and refusing dead prey fish or artificial diets.     

Variation in feeding behaviour of Nilaparvata lugens on resistant and susceptible rice varieties

The feeding behaviour of Nilaparavata lugens was monitored on three rice varieties showing different levels of resistance in the Philippines, using a video-assisted observation method. N. lugens made more frequent, shorter probes on the moderately resistant IR46 and resistant IR62 rice varieties than on the susceptible IR22. Honeydew production was significantly lower on the resistant varieties though insect weight gains in 24 h were similar on IR46 and IR22, both being significantly greater than on the highly resistant variety.Population development, growth index and damage ratings were low on IR62 indicating antibiosis and/or non preference. When IR46 plants were infested as seedlings population increase, growth index and damage ratings were similar to those on the susceptible IR22. When infested at a later stage of plant growth the damage rating showed a moderate level of resistance though some population development was maintained, indicating antibiosis and tolerance. N. lugens started probing less frequently after surface exploration on both resistant varieties than on IR22 suggesting the presence of a resistance factor associated with the surface waxes of these varieties.

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Résumé Le comportement alimentaire de Nilaparvata lugens sur variétés de riz, sensible (IR22), partiellement résistante (IR46) et fortement résistante (IR62), a été contrôlé avec une méthode associant la vidéo à l'observation. N. lugens faisait des piqûres plus fréquentes et plus brèves sur IR46 et IR62, que sur la variété sensible. La production de miellat était significativement plus faible sur les variétés résistantes, bien que les gains de poids des insectes aient été les mêmes en 24 h sur IR46 et IR22, les deux étant significativement supérieurs à celui sur IR62.La croissance de la population, l'indice de croissance et le taux de dégâts étaient tous plus faibles sur IR62, ce qui révèle une antibiose et/ou une absence de préférence. Quand la contamination des IR46 a au lieu au stade semis, la croissance de population, l'indice de croissance et le taux de dégâts étaient semblables à ceux de la variété sensible IR22. Quand la contamination avait lieu à un stade ultérieur, le laux de dégâts révélait un niveau modéré de résistance bien qu'une certaine croissance de population se soit maintenue, ce qui révèle antibiose et tolérance.Après exploration de la surface des feuilles des deux variétés résistantes, N. lugens sondait moins fréquemment que sur IR22, ce qui laisse présumer un facteur de résistance associé aux cires superficielles de ces variétés.

     

Deterrents extracted from the leaves of Ginkgo biloba: effects on feeding and contact chemoreceptors

Powdered dried ginkgo tree leaves were subjected to various chemical extractions and successive extracts were monitored for antifeedant activity against larvae of Pieris brassicae. Many fractions moderately inhibited food intake, and some were deterrent at levels as low as 25–50 ppm. Some behaviourally highly active fractions were tested electrophysiologically for neural responses in the maxillary taste sensilla. These extracts appeared to stimulate deterrent receptors. There were distinct differences in responses between Pieris brassicae and P. rapae. Ginkgolide A, B, and C each strongly stimulated deterrent receptors in P. rapae, which corresponds with the observation (Matsumoto & Sei, 1987) that these compounds are effective antifeedants for this species. No toxic effects were observed in insects after feeding for 24 h on diets containing ginkgo extracts.

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Résumé Des feuilles de Gingko séchées et broyées ont été soumises à différentes extractions. On a testé l'activité déterrante (antiapprétant) des extraits successifs sur les larves de Pieris brassicae. Un grand nombre de fractions inhibent modérément la consommation, et certaines sont déterrantes à des concentrations aussi faibles que 25–50 ppm. Par électrophysiologie, on a testé la réponse nerveuse des sensilles gustatives maxillaires à certaines fractions qui montraient une forte action sur le comportement. Il s'avère que ces extraits stimulent les récepteurs déterrants. Les réponses diffèrent pour P. brassicae et P. rapae. Les gingkolides A, B et C stimulent chacun fortement les récepteurs déterrants chez P. rapae, ce qui correspond aux observations (Matsumoto & Sei, 1987) selon lesquelles ces composés sont des antiappétants efficaces pour cette espèce. On n'a pas observé d'effet toxique sur les insectes nourris durant 24 heures par des diètes contenant des extraits de Gingko.

     

Cross habituation to feeding deterrents and acceptance of a marginal host plant byPieris rapae larvae

Sensitivity of the cabbage butterfly,Pieris rapae L. to feeding deterrents was compared for larvae reared on different food sources under laboratory conditions. Since cabbage-reared larvae normally reject nasturtium,Tropaeolum majus L., the effects of previous exposure to allelochemicals on larval acceptance or rejection of this plant were also examined. When compared with cabbage-reared larvae, nasturtium-reared larvae were less sensitive to feeding deterrents including cymarin, erysimoside and 2-O-β-d-glucosyl cucurbitacin E. Nasturtium-reared larvae were insensitive to chlorogenic acid, which was deterrent to cabbage-reared larvae. Feeding by larvae reared on a wheat germ diet was not deterred by these compounds. The results indicate that dietary experience can extensively affect larval sensitivity to feeding deterrents and that cross habituation of larvae to deterrents occurs in response to certain chemical constituents of nasturtium and wheat germ diet. Digitoxin, however, proved to be an exception. Larvae reared on either nasturtium or wheat germ diet were as sensitive to digitoxin as those reared on cabbage. Previous results have shown that rejectionof nasturtium by cabbage-reared larvae is due to the presence of strong feeding deterrents in this plant. However, more than 50% of 2nd instar larvae reared from neonate on cabbage leaves treated with strophanthidin, cymarin, erysimoside, digitoxigenin and digitoxin accepted nasturtium as a food source. 2-O-β-d-glucosyl cucurbitacin E, 2-O-β-d-glucosyl cucurbitacin I and rutin were also active in causing larvae to feed on nasturtium. Thus dietary exposure to unrelated plant chemicals can profoundly affect insect acceptance of a plant that contains feeding deterrents.     

Perception of the oviposition deterring pheromone by tarsal and abdominal contact chemoreceptors in Pieris brassicae

Perception of the oviposition deterring pheromone by contact chemoreceptors in female Pieris brassicae was studied employing a tip recording technique. Electrophysiological responses of tarsal taste hairs to eggwash solutions show a marked increase in frequency of spikes originating from only one sensory cell. This suggests that in foretarsal taste hairs females, apart from the glucosinolate cells also possess sense cells specifically sensitive to the oviposition deterring pheromone.Morphological studies by means of the scanning electron microscope revealed that the ovipositor of P. brassicae is provided with two groups of contact chemoreceptors. Electrophysiological recordings from these sensilla indicate the presence of at least three sensory cells, one of them being a mechanoreceptor. Stimulation with eggwash evokes a slight increase in spike frequency which cannot be ascribed to one particular sense cell. This indicates that abdominal taste hairs in some way may participate in the perception of the oviposition deterring pheromone. Responses to glucosinolates do not differ significantly from control stimulations.

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Résumé Une technique d'enregistrement apical a été utilisée pour examiner la perception d'une phéromone dissuadant la ponte par les poils récepteurs chimiques de contact des femelles de Pieris brassicae. Les réponses électrophysiologiques des poils gustatifs des tarses en présence de solutions de rinçage d'oeufs présentent une fréquence marquée des potentiels d'action provenant principalement d'une cellule sensorielle. Ceci suggère que les poils gustatifs des tarses des pattes antérieures des femelles possèdent, en plus de cellules répondant aux glucosinolates, des cellules sensorielles sensibles spécialement à la phéromone dissuadant la ponte.Des études morphologiques au microscope à balayage révèlent que l'oviposition de P. brassicae est pourvu de deux groupes de chimiorécepteurs de contact. Des enregistrements électrophysiologiques de ces sensilles révèlent la présence d'au moins trois cellules sensorielles, l'une d'entre elles étant un mécanorécepteur. La stimulation avec la solution de rinçage des oeufs évoque un léger accroissement de la fréquence des potentiels d'action qui ne peut être attribué à une cellule sensorielle particulière. Ceci indique que les poils gustatifs abdominaux peuvent participer d'une certaine façon à la perception de la phéromone dissuadant la ponte. Les réponses aux glucosinolates ne diffèrent pas significativement des stimulations témoin.

     

Daily changes in the responsiveness of taste receptors correlate with feeding behaviour in larvae of Spodoptera littoralis

Larvae of Spodoptera littoralis were monitored through the final larval stadium (6 days) for weight gain, amount eaten in bioassays and responsiveness of gustatory receptors to stimulation with sodium chloride and four concentrations of sucrose. The age of larvae within the stadium, time of day (am or pm) and interactions between these factors influenced the electrophysiological response to 0.05 M sucrose. Overall, the responsiveness of taste receptors increased in the first half of the stadium, then declined. Neural input correlated with weight gain on days 1 to 4, and with amount eaten in bioassays on days 1 to 5. This correlation between neural input and feeding behaviour indicates that the observed modulation in neural sensitivity has a causal relationship with feeding behaviour. The ways in which the modulation might occur are discussed.     

Oviposition behaviour of Dasineura brassicae on a high- versus a low-quality Brassica host

The oviposition behaviour of the brassica pod midge, Dasineura brassicae Winn. (Diptera: Cecidomyiidae), on a preferred host, Brassica napus L. (summer oilseed rape) was compared to that on a nonpreferred, less suitable host for larval growth, Brassica juncea (L.) Coss & Czern (brown mustard). The experiments were done under field conditions with wild females.The number of landing females was significantly higher on B. napus than on B. juncea, indicating host differences in olfactory and/or visual stimuli. After landing, the behaviour differed in that females stayed longer and laid more egg batches on B. napus than on B. juncea plants. The probability that oviposition would occur after landing and the potentially adjustable egg batch size were similar on the high- and the low-quality host.A larger egg load on B. napus than on B. juncea can thus be attributed mainly to a higher landing rate and more repeated ovipositions occurring on B. napus.

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Comportement de ponte de Dasineura brassicae sur des Brassica de qualites élevée et basse

Résumé La comparaison porte sur le comportement de ponte de D. brassicae Winn. (Dept. Cecidomyiidae) sur Brassica napus L., hôte préféré, et sur B. juncea L., hôte non-préféré, qui convient moins au développement larvaire. Les expériences réalisées en champ ont porté sur des femelles sauvages.Le nombre de femelles qui atterrissent sur B. napus est significativement plus élevé que sur B. juncea, ce qui indique des différences entre les stimuli olfactif et visuel des plantes-hôtes. Aprés atterrisage, le comportement diffère en ce que les femelles restent plus longtemps et déposent plus de pontes sur B. napus que sur B. juncea. La probabilité pour que la ponte suive l'atterrissage et les tailles des ooplaques potentiellement ajustables sont semblables sur les hôtes de qualité élevée ou basse.Une charge supérieure en oeufs sur B. napus que sur B. juncea peut ainsi être principalement attribuée à un taux d'atterrissage plus élevé et à une plus grande fréquence de ponte sur B. napus.

     

Effect of methoxyphenols on grain aphid feeding behaviour

Methoxyphenols might be important in the resistance of cereals to aphids. Electrical penetration graph (EPG) recordings were used to determine the effect of caffeic, ferulic and sinapic acids, and scopoletin on the feeding behaviour of the grain aphidSitobion avenae (F.). Aphids on wheat seedlings treated systemically with these phenols showed reduced ingestion of phloem sap and salivation into sieve elements in most cases. The earlier pathway phases of probing were prolonged. Moreover increase in number of probes as well as reduction of total time of probing was observed. In addition, no O-demethylase activity was found in homogenates of aphids fed on moderately-resistant (phenolic rich) or susceptible (phenolic poor) wheat varieties. The significance of these results for understanding the resistance of cereals to aphids is discussed.     

Behavioural variations among strains of Trichogramma spp.: Host-age selection

Host-age selection by several Trichogramma spp. strains was examined in laboratory choice-tests, in order to use this as a criterion to select a candidate strain for inundative releases against lepidopterous pests of cabbage in the Netherlands. The behaviour of individual females toward combinations of host eggs of two different ages (zero vs 2 and 1 vs 3-day-old) of the host species Mamestra brassicae, Pieris brassicae and P. rapae was observed directly. Host age generally did not appear to affect contact or acceptance of eggs of either host species, except that oviposition was generally lower in the oldest M. brassicae eggs. The duration of the oviposition behaviour was sometimes influenced by host-age. All strains were able to discriminate between parasitized and unparasitized hosts by external or internal examation. The lack of variation among the strains suggests that the host-age selection pattern of a given host-Trichogramma combination is predominantly dependent on the host species. The results are discussed in relation to optimal foraging theory.

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Résumé Afin de sélectionner une souche candidate de Trichogramma sp. pour des lâchés inondatifs contre les Lépidoptères, pestes du chou aux Pays-Bas, la sélection hôte-âge de plusieurs souches de Trichogramma spp., issues d'un éventail de 60 souches, a été examinée en laboratoire par des tests de choix. Le comportement de femelles individuelles, mises en présence de combinaisons d'oeufs-hôtes d'âges différents (zero ou 2 jours et 1 ou 3 jours) des espèces-hôtes Mamestra brassicae, Pieris brassicae et P. rapae, a été observé directement. En général, l'âge de l'hôte ne semble pas affecter le contact et l'acceptation des oeufs quelque soit l'espèce-hôte. La ponte diminue généralement dans les oeufs les plus âgés de M. brassicae, mais pas chez les autres espèces. La durée du comportement de ponte est parfois influencée par l'âge de l'hôte. Toutes les souches ont montré la capacité de discriminer entre les hôtes parasités et nonparasités, a l'examen externe ou interne et ceci, indépendamment de l'âge de l'hôte. Le manque de variation parmi les souces suggère que le mode de sélection hôte-âge est surtout caractéristique de l'espèce-hôte. Les resultats sont discutés en relation avec la théorie de fourrage optimal.

     

Behavioural variation among strains ofTrichogramma spp.: host-species selection

The host-selection behaviour of nine strains ofTrichogramma spp., towards eggs ofMamestra brassicae, Pieris brassicae andP. rapae, was investigated in laboratory experiments in order to select candidate strains for inundative releases against these species. Experiments were carried out by continuously observing the behaviour of individual females, which were offered equal numbers of eggs of two host species arranged in a grid.M. brassicae was a highly acceptable host species for all strains, whereas the acceptability of the twoPieris species was similar within strains, but varied between strains. Considering the variation in acceptance ofPieris eggs, strains either showed: (1) no preference betweenMamestra andPieris (High PierisAcceptance = HPA strains), (2) a preference forMamestra (Variable PierisAcceptance = VPA strains), or (3) an aversion forPieris (Low PierisAcceptance = LPA strains). Females of VPA strains showed a high acceptance ofPieris eggs if the preferredMamestra eggs were absent. They contacted comparatively fewerPieris eggs in presence ofMamestra eggs, which indicates selection of hosts at a distance. HPA strains probably are the best candidates for inundative releases.

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Sélection hôte-espèce par différences souches de l'æuf-parasiteTrichogramma spp.

Résumé Le comportement de sélection de l'hôte par neuf souches deTrichogrammes, vis-à-vis d'ufs deMamestra brassicae, Pieris brassicae etP. rapae a été étudié au laboratoire afin de sélectioner des souches candidates pour des lâchers inondatifs contre ses espèces. Des expériences ont été menées par observations en continu du comportement de femelles auxquelles était offert un nombre égal d'ufs de deux espèces d'hôtes, disposés selon un grille.M. brassicae est un espèce-hôte fortement acceptée par toutes les souches. Au contraire, l'acceptabilité des deux espèces dePieris, semblable pour chaque souche, varie entre les souches. En tenant compte de la variation d'acceptance des ufs dePieris les souches (1) ne montrent aucune préférence entreMamestra etPieris, (2) montrent une préférence pourMamestra, ou(3) une aversion pourPieris. Des femelles des souches du second groupe acceptent fortement les ufs dePieris si les ufs deMamestra, préférés, sont absents. Comparativement, elles rentrent en contact avec moins d'ufs dePieris en présence d'ufs deMamestra. Ceci indique une sélection des hôtes à distance. Les souches du premier groupe sont probablement les meilleures candidates pour des lâchers inondatifs.

     

Behavioural and sensory responses to drimane antifeedants in Pieris brassicae larvae

Fifteen drimane compounds were tested for their feeding inhibiting activity in larvae of Pieris brassicae L. (Lepidoptera: Pieridae) when applied to leaf material of the host plant Brassica oleracea L. The antifeedant efficacy of the drimanes was related to their molecular structure in order to identify important functional groups. Of the drimanes tested, those with a lactone group on the B-ring were the most effective feeding inhibitors. Additionally, the sensory responses to 13 of the drimanes were measured. Neural activity was evoked in the deterrent cell in the medial sensillum styloconicum. Also, inhibition of sensory responses to feeding stimulants was found. Results of behavioural and electrophysiological tests were correlated in an attempt to elucidate the sensory code underlying feeding inhibition by drimanes in Pieris brassicae. It was concluded that the response of the deterrent cell in the medial sensillum styloconicum contributes significantly to inhibition of feeding behaviour in larvae of Pieris brassicae.     

Morphological aspects of feeding and improvement in feeding ability in early stage larvae of the milkfish,Chanos chanos

The osteological development of elements comprising the oral cavity and fins was examined in early stage larvae of laboratory-reared milkfish,Chanos chanos, from hatching to 200 hours after hatching. Fundamental elements of the oral cavity had developed by the time of initial mouth opening, 54 hours after hatching. The oral cavity was long and cylindrical, with a short, robust Meckel's cartilage, and robust quadrate and symplectic-hyomandibular cartilages. The initial ossification of existing elements and addition of new elements occurred between 120–146 hours after initial mouth opening (HAMO), whereas the cartilaginous basihyal and caudal fin-supports appeared at 37.5 and 61.5 HAMO, respectively. Based on the morphology and developmental patterns of characters examined in this study, the feeding mode of early stage larval milkfish was considered to be “straining,” with an improvement in feeding ability occurring between 120–146 HAMO.     

Activity persistence of granulosis viruses [Baculoviridae] in soils in United Kingdom and Denmark

The persistence of granulosis virus infectivity in soil has been investigated. Virus applied to soil was extracted at different times up to 1 year after the original application and the amount of virus was determined using bioassays. In this way the content of infective capsules remained was determined. It was shown that about 99% of the original virus infectivity was lost during 1 year. Together with other investigations this result indicates that leaching, passive lateral movements and degradation accounts for the main part of the loss.      

A test of a model for planktivorous filter feeding by bream Abramis brama

Synopsis The planktivorous feeding of bream, Abramis brama on Daphnia hyalina and Bosmina coregoni was analyzed in a stepwise regression analysis with the average size (and standard deviation) of consumed organisms as dependent variable and the size of the fish, the average size (and standard deviation) of the organisms and their density in the environment as independent variables. Three basic predictions on filter feeding were formulated and tested. It was predicted that the (average) prey size should increase with fish size, but that the standard deviation should decline. Secondly the prey size should be strongly correlated with the prey size available and thirdly the prey density should have little effect on the size selection. These hypotheses could not be rejected for bream>10 cm feeding on B. coregoni and for bream>20 cm feeding on D. hyalina. The hypotheses were not valid for smaller bream as these acted as particulate or combined filter- and particulate feeders.     

On the feeding habits ofRhynchopsilopa [Dip.: Ephydridae]

Further evidence for the myrmecophagous habits of adultRhynchopsilopa flies is supplied. The genus probably evolved in association with ants of the genusCrematogaster. The feeding behaviour ofR. nitidissima Hendel on workers ofCrematogaster is described. Choice experiments show an absolute preference forCrematogaster over 3 other and genera. The mouth parts ofR. nitidissima are briefly described.

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Résumé Un témoignage supplémentaire des habitudes myrmécophages des adultes de DiptèresRhynchopsilopa est fourni. Le genre a probablement évolué en association avec les fourmis du genreCrematogaster. Le comportement alimentaire deR. nitidissima sur les ouvrières deCrematogaster est décrit. Les choix expérimentaux montrent la préférence absolue pour le genreCrematogaster à l'exclusion des 3 autres genres de fourmis. Les pièces buccales deR. nitidissima sont brièvement décrites.

     

Tidal activity pattern and feeding behaviour of the ophiuroid Ophiocoma scolopendrina on a Kenyan reef flat

Ophiocoma scolopendrina exhibits a distinctive pattern of feeding activity on intertidal reef platforms off Kenya. With the first wave of the flooding tide, dense aggregations of these ophiuroids (up to 320 m⁻²) engage in a 1–2 min burst of surface-film feeding, vigorously sweeping the air-water interface and associated sea foam with the ventral surface of 2–4 arms. Suspension feeding (with arms extended in the water column) is the primary feeding mode throughout the rest of the tidal cycle (involving 25–65% of the population at a time), while bottom feeding (with arms extended along the substratum) is infrequent (<10%). Field experiments showed that surface-film feeding is regulated by water depth and can be triggered by suspended particles. This feeding mode appears to be an adaptation to the intertidal habitat, enabling the ophiuroids to exploit a nutrient-rich surface film during a temporal refuge (low tide) from fish predation. Dense populations of O. scolopendrina may represent an important trophic link between producers of particulate organic material and higher-level consumers in coral reef environments.Tara Oak and Robert E. Scheibling contributed equally to this paper. The order of authorship is alphabetical     

Screening of crude plant extracts as feeding deterrents of the wireworm, Melanotus communis

Extracts from 78 plant species (24 families) were screened for antifeedant activity against the corn wireworm, Melanotus communis Gyll. (Coleoptera: Elateridae). Five extracts from four families significantly reduced wireworm feeding damage in a series of choice feeding tests. Two extracts, Asclepias tuberosa and Hedera helix, exhibited exceptional levels of feeding deterrency. An x-ray technique indicated that wireworms burrowed indiscriminately between soil containing either of these extracts and surrounding untreated soil, but were found more frequently in the untreated areas.

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Résumé L'aptitude à empêcher la prise de nourriture de Melanotus communis Gyll. a été recherchée dans les extraits de 78 plantes appartenant à 24 familles. Cinq extraits de 4 familles ont réduit significativement les dégâts alimentaires lorsqu'il y avait choix alimentaire. Deux extraits — Asclepias tuberosa et Hedera helix — présentaient des niveaux exception-nellement élevés de dissuasion de l'alimentation. Une technique aux rayons X a montré que les taupins fouillaient indistinctement le sol contenant de ces extraits ou le sol avoisinant non traité, mais on les recontrait plus fréquemment dans le sol non traité.