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.

     

Development and evaluation of an enzyme-linked immunosorbent assay to detect Pieris rapae remains in guts of arthropod predators

An enzyme-linked immunosorbent assay (ELISA) was developed to detect remains of Pieris rapae L. (Lepidoptera: Pieridae) immature stages in the guts of field collected arthropod predators. The assay can be used to help ascertain the relative importance of arthropod predator species in suppressing P. rapae in cabbage, Brassica oleracea var. capitata L. The ELISA is sensitive to all immature stages of P. rapae, although first and fifth instars can be detected more readily than eggs or pupae and third instars showed intermediate detectability. Assays on whole body homogenates of predators readily detected predation on P. rapae first instars by all seven of the predator species tested, although response generally declined with increasing predator size. Together the results show that the P. rapae ELISA possesses a sufficiently high level of sensitivity and specificity to be a useful tool in helping to elucidate the roles of arthropod predator species in reducing populations of P. rapae in cabbage.     

Identificaton of UV,green and red receptors,and their projection to lamina in the cabbage butterfly,Pieris rapae

Summary The photoreceptors in the compound eye of a cabbage butterfly, Pieris rapae, were examined by conventional and intracellular-labeling electron microscopy by the use of the cobalt(III)-lysine complex as an ionized marker. Five types of spectral sensitivity were recorded intracellularly in electrophysiological experiments. They peaked at about 340, 380, 480, 560 and 620 nm, respectively. One of the distal retinula cells (R2) was a UV receptor, whereas the R4 distal retinula cell was a green receptor. The basal retinula cell, R9, was found to be a red receptor; it was localized near the basement membrane, having a bilobed cell body with an individual nucleus in each lobe. A small number of rhabdomere microvilli were present in a narrow cytoplasmic bridge connecting the two lobes. The axons of six retinula cells (R3–R8) in each ommatidium terminated at the cartridge in the lamina (short visual fiber), whereas those of the other three retinula cells, R1, R2 and R9, extended to the medulla (long visual fiber). The information from the UV and red receptors is therefore probably delivered directly to the medulla neurons, independent of that from the other spectral receptor types.     

Visual learning in assays of sinigrin solution as an oviposition releaser for the cabbage butterfly, Pieris rapae

Pieris rapae (L.) oviposited readily on paper wetted with sinigrin solution, if it was yellow, green, light blue or white, but not if violet, red, or black. On violet discs, feeding responses distracted the butterflies from oviposition and they often laid on neighbouring green discs which lacked sinigrin. When some discs with water and others with sinigrin solution were offered together, butterflies oviposited more on water discs of the same colour as the sinigrin discs, as opposed to those of a different colour. Within a range of sinigrin solutions tested (10–1000 ppm) the higher concentrations elicited most oviposition. Preference was slight for different concentrations applied to physically identical discs, but marked for the higher of two concentrations (100 vs 75 ppm) on discs of different but equally acceptable appearance. It was concluded that by trial-and-error, ovipositing butterflies learnt to associate the appearance of the oviposition site with its contact chemical stimuli, and that learning either increased or decreased the specificity of responses to sinigrin-treated discs, according to the design of the assay. Learning influenced preference differently in butterflies offered two as opposed to three kinds of acceptable site.

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Résumé La ponte de Pieris rapae L., phytophage spécialiste de crucifères, a été étudiée avec des rondelles de papier buvard coloré, imbibées avec une solution de sinigrine, substance spécifique, non volatile, des plantes hôtes. Les papillons ont bien pondu lors de répétitions sur papiers jaune, vert, bleu clair ou blanc, et peu sur papiers rouge, violet ou noir. Avec des rondelles violettes, la prise d'aliments a distrait de la ponte, et des oeufs ont alors été souvent pondus sur des rondelles vertes proches sans sinigrine. Quand des rondelles avec de l'eau et d'autres avec des solutions de sinigrine ont été offertes simultanément, les papillons ont pondu plus sur les rondelles imbibées d'eau de la même couleur que celles avec de la sinigrine, que sur celles d'une autre couleur. Dans la gamme des concentrations de solution de sinigrine examinées (10 à 1000 ppm les plus élevées ont induit les pontes les plus importantes. Il y a eu peu de préférence entre des rondelles de même couleur ayant des concentrations différentes, mais le choix a été net quand les rondelles étaient de couleurs différentes et pourtant acceptables, dans ces conditions la concentration la plus élevée (100 versus 75 ppm) était préférée. On peut en conclure que les papillons, par un apprentissage avec essai et erreur, lors de la ponte ont associé l'aspect du lieu de ponte avec les stimulations chimiques de contact; l'apprentissage a, soit augmenté, soit réduit la spécificité des réponses aux rondelles traitées à la sinigrine, suivant les conditions de l'expérience. La discussion examine la signification du tambourinage par les pattes antérieures comme séquence préliminaires de la ponte.

     

Ultrastructure and migration of screening pigments in the retina of Pieris rapae L. (Lepidoptera,Pieridae)

Summary The retinal morphology of the butterfly, Pieris rapae L., was investigated using light and electron microscopy with special emphasis on the morphology and distribution of its screening pigments. Pigment migration in pigment and retinula cells was analysed after light-dark adaptation and after different selective chromatic adaptations. The primary pigment cells with white to yellow-green pigments symmetrically surround the cone process and the distal half of the crystalline cone, whilst the six secondary pigment cells, around each ommatidium, contain dark brown pigment granules. The nine retinula cells in one ommatidium can be categorised into four types. Receptor cells 1–4, which have microvilli in the distal half of the ommatidium only, contain numerous dark brown pigment granules. On the basis of the pigment content and morphology of their pigment granules, two distal groups of cells, cells 1, 2 and cells 3, 4 can be distinguished. The four diagonally arranged cells (5–8), with rhabdomeric structures and pigments in the proximal half of the cells, contain small red pigment granules of irregular shape. The ninth cell, which has only a small number of microvilli, lacks pigment. Chromatic adaptation experiments in which the location of retinula cell pigment granules was used as a criterium reveal two UV-receptors (cells 1 and 2), two green receptors (cells 3 and 4) and four cells (5–8) containing the red screening pigment, with a yellow-green sensitivity.     

Constituents of host- and non-host plants deterring oviposition by the cabbage butterfly, Pieris rapae

Oviposition by Pieris rapae L. on cabbage was deterred by homogenized cabbage tissue sprayed onto intact plants. Ether extracts of cabbage also were deterrent, but water extracts were not. Hexane extracts of other host plants deterred oviposition and water extracts had little or no effect. Polar as well as non-polar extracts of non-host plants inhibited oviposition. Polar deterrents in the non-host crucifers, Erysimum cheiranthoides and Capsella bursa-pastoris may explain the avoidance of these plants by P. rapae. Chemical deterrents, as well as stimulants, apparently play a major role in the acceptance or rejection of plants as hosts by ovipositing female butterflies.

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Zusammenfassung Die Eiablage von Pieris rapae auf Kohl wurde gehemmt von einem Homogenisat aus Kohlblättern, das auf intakte Pflanzen gesprüht wurde. Ether-Extrakte der Kohlblätter wirkten im Gegensatz zu Wasserextrakten abschreckend. Ebenso zeigten Hexanextrakte anderer Wirtspflanzen eine hemmende Wirkung auf die Eiablage, während Wasserextrakte anderer Wirtspflanzen keine Wirkung aufwiesen. Dagegen aber verhinderten sowohl polare als auch unpolare Extrakte von Nicht-Wirtspflanzen die Eiablagen. Polare, abschreckende Substanzen in den Nicht-Wirtspflanzen Erysimum cheiranthoides und Capsella bursa-pastoris (Cruciferae) könnten Ursache dafür sein, dass P. rapae diese Pflanzen meidet. Chemische Substanzen mit sowohl abschreckender als auch stimulierender Wirkung spielen offensichtlich eine bedeutende Rolle für legebereite Schmetterlingsweibchen bei der Auswahl von Pflanzen als geeigneten Eiablageplatz.

     

A simple action treshold for timing applications of a granulosis virus to controlPieris rapae [Lep.: Pieridae]

Pieris rapae (L.) an important pest of cole crops in the northeastern United States, is susceptible to a granulosis virus,Pieris rapae GV (PrGV), that has been shown to be an effective control measure by researchers in several countries. As an alternative to weekly applications of virus to protect cabbage, we tested the use of an action threshold of one small (first-third instar) larva per plant. Results were compared with those obtained using the same threshold with permethrin, and with weekly applications of virus. Plots treated weekly with virus received 5 applications but the action threshold was exceeded only once. In all virus-treated plots, numbers of large (fourth-fifth instar) larvae remained below 0.35 per plant, and were lower at the end of the season (0.07 in plots treated weekly and 0.1 in plots treated once) than in either the untreated or permethrin-treated plots (0.5). In late August, numbers of large larvae in the check plots reached almost 3 per plant. At harvest the number of feeding holes over 0.3 cm in diameter in the 4 innermost frame and the 4 wrapper leaves were counted. Check plots differed from treated plots by an average of 124.2±6.5 holes per plant in the frame and wrapper leave; virus-treated plots had 51.1±6.9 holes more than the permethrin plots. The difference in overall damage between plots treated 5 times with virus during the season and those treated once was not significant. Plots treated once with virus had significantly more damage (7.6±2.7) to wrapper leaves than those treated five times and marketability ratings were somewhat lower, based on fresh market standards. There were no significant differences in head weight among the treatments. At harvest, a high proportion of larvae collected from the check plots were diseased (77% versus an average of 46% in the treated plots). Because of the high numbers of large larvae in the check plots in late August and the extensive damage to plants, we assumed that virus did not affect a significant number of larvae in these plots until late in the growing season. These results indicate the usefulness of PrGV in a cabbage IPM program and that the use of action thresholds can be highly effective, particularly when insect numbers only occasionally reach damaging levels. While cabbage treated with permethrin had the least amount of injury, that treated weekly with virus was not significantly different by fresh market standards, and all cabbages treated with virus met processing standards. For the fresh market, in which cosmetic standards are more important, PrGV may have to be used weekly or with an action threshold lower than one small larva per plant.      

Response of the wasp (<Emphasis Type="Italic">Cotesia glomerata</Emphasis>) to larvae of the large white butterfly (<Emphasis Type="Italic">Pieris brassicae</Emphasis>)

The large white butterfly (Pieris brassicae L) first invaded northernmost Japan from Siberia around 1994, and after a few years, began to expand its range. The wasp, Cotesia glomerata (L) parasitizes larvae of the small white butterfly (Pieris rapae crucivora Boisduval), a usual host in the same geographic area. Some Pieris brassicae larvae in Hokkaido have been parasitized by Cotesia glomerata, but the parasitism rate of Pieris brassicae larvae tends to be lower than that of Pieris rapae. To examine the process of parasitizing Pieris brassicae larvae, we observed how the parasitoid wasp responded to the host larvae on damaged leaves. Cotesia glomerata females tended to avoid Pieris brassicae larvae, and even when female wasps inserted their ovipositors into Pieris brassicae larvae, none laid eggs. The parasitoids obtained from Pieris rapae larvae failed to parasitize Pieris brassicae during the host-acceptance step.     

Axonal pathfinding in developing labial palps of the butterflies,Pieris rapae and Pieris brassicae

Summary Electron microscopy was used to study the developmental changes in the pupal labial palp of Pieris rapae L. and P. brassicae L. prior to axogenesis of the peripheral receptor organs. Two cells emigrate from the anlage of the labial palp-pit organ (LPPO). They develop growth cones and filopodia, which are directed distally. The filopodia insert into stem cells of the LPPO. The perikarya of the cells proceed into the hemolymph space of the palp such that, eventually, these cells bridge the gap between the LPPO and the apical scolopidial organ (ASO) of the labial palp. This bridge is established at about 11% of total pupal development. Only 6 h later, at 15% of pupal development, the cells are no longer visible. We suggest that these cells are luminal neurons and name them pit-organ luminal (POL) cells. These POL cells may act as primary guiding structures for the axons of the sensory cells of the LPPO, which are assumed to orientate along this structure once they reach the ASO.Supported by the Deutsche Forschungsgemeinschaft (H.A.: SFB4/G1).     

Oviposition and larval performance of Rabdophaga terminalis on Salix spp. with special consideration to bud size of host plants

Females of the gall midge Rabdophaga terminalis (H. Loew) (Diptera: Cecidomyiidae) are very selective in their choice of host plants at oviposition. In preference trials, eggs were laid almost exclusively on S. alba and S. fragilis. One single oviposition occurred on S. viminalis but none on S. triandra, S. daphnoides or S. smithiana. Significantly more eggs were laid on S. alba than on S. fragilis. No differences were detected between clones of the same species regarding oviposition in laboratory trials. Within the species of S. alba, larger buds were preferred. However, this did not explain the preference of S. alba to S. fragilis. There was agreement between plant species selected by the females and survival of the larvae. No galls developed when eggs were transferred to species that were not oviposited on. There was no difference between S. alba and S. fragilis in larval growth, but after two weeks of development there was a significantly higher proportion of larvae in cocoons and pupae on S. alba. There was a negative linear relationship between weight of larvae and larval number per gall, indicating larval competition. The negative slope was steeper for S. alba than for S. fragilis when larvae were weighed after two weeks of development. There were some discrepancies between laboratory and field data on infestation of S. alba and S. fragilis. Higher infestation levels were related to denser stands in the field.

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Résumé Etude de la ponte et des performances larvaires de Rabdophaga terminalis (Diptera; Cecidomyiidae) sur plusieurs espèces de saules avec examen particulier de l'influence de la taille des bourgeons.La ponte et les performances larvaires de la cécidomyie galliforme, Rabdophaga terminalis (H. Loew) ont été étudiées sur plusieurs espèces et clones de saules cultivés en Suède. Les femelles sélectionnent avec soin les plantes hôtes lors de la ponte. Dans des essais préférentiels, les oeufs ont été pondus presque exclusivement sur S. alba et S. fragilis; une ponte a été observée sur S. viminalis et aucune sur S. triandra, S. daphnoïdes ou S. smithiana. Significativement plus d'oeufs ont été pondus sur S. alba que sur S. fragilis. Les essais au laboratoire n'ont pas permis de déceler des différences entre clones de la même espèce. Chez S. alba, les plus gros bourgeons étaient préférés, mais ceci ne permet pas d'expliquer la préférence pour S. alba par rapport à S. fragilis. Il y a coïncidence entre les plantes choisies par les femelles et al survie des larves; il n'y avait pas développement de galles quand des oeufs étaient transférés sur des espèces qui n'avaient pas reçu de pontes. La croissance larvaire sur S. alba et S. fragilis ne présentait pas de différences, mais après 2 semaines de développement il y avait significativement plus de larves en cocons et pupes sur S. alba. La corrélation entre le poids des larves et le nombre de larves par cellule était linéaire et négative, ce qui indique une compétition larvaire; la pente était plus raide pour S. alba que pour S. fragilis quand les larves étaient pesées au bout de 2 semaines de développement. Il y a eu quelques discordances entre les résultats au laboratoire et les données de terrain sur la contamination de S. alba et S. fragilis. Les plus fortes contaminations étaient observées à l'extérieur sur pépinières denses.

     

Sensory cell degeneration in the ontogeny of the chemosensitive sensilla in the labial palp-pit organ of the butterfly,Pieris rapae L. (Insecta,Lepidoptera)

Summary The ontogeny of the chemoreceptive sensilla in the labial palp-pit organ was studied in Pieris rapae by examining twelve successive stages between pupation and emergence of the imago, which takes a period of 160 h under the experimental conditions. Mitoses occur until 20 h after pupation. They lead to anlagen of sensilla, 91% of which are comprised of three sensory cells. However, two sensory cells degenerate in each sensillum during a period of 28 h. The same process occurs in anlagen with four sensory cells resulting in bicellular sensilla. Axons grow out only after the number of sensory cells has been reduced. Further consecutive steps in sensory cell differentiation are: (a) outgrowth of dendritic outer segment and dendrite sheath; (b) outgrowth of trichogen process and change in structure of elongating dendrite sheath; (c) deposition of cuticle and pore tubules in the pegs; (d) retraction of trichogen process; (e) increase in diameter of dendritic outer segment accompanied by increase of microtubule number and appearance of regularly spaced electron-dense bodies at tubular doublets; (f) branching of dendritic outer segment; and (g) transformation of the dendritic branches into curled lamellae and partial destruction of the dendrite sheath. The unique process of sensory cell degeneration is interpreted as an event that revokes a step towards a possible functional improvement of the labial palp-pit organ during further evolutionSupported by the Deutsche Forschungsgemeinschaft (SFB 4/G1)     

Differential selection of host plants by twoPieris species: the role of oviposition stimulants and deterrents

Oviposition responses ofPieris rapae L. andP. napi oleracea Harris to nine crucifers, one Capparidaceae and one Tropaeolaceae were directly compared under controlled conditions. Chemical fractions from these plants were also tested on both insects for the presence of oviposition stimulants or deterrents. The results showed that plant chemistry is a key factor in differential selection of potential hosts by thesePieris species. Some plant species were equally acceptable to bothPieris species. However,P. rapae preferred cabbage over most test plants whereasP. napi oleracea strongly preferred plant species that were avoided byP. rapae. The observed preferences were explained in most cases by the presence of stimulants and deterrents in extracts of the plants. The twoPieris species have apparently evolved differential sensitivities to the chemical stimuli that trigger or deter oviposition. The balance of positively and negatively interpreted sensory signals evoked by plant chemicals obviously plays an important role in acceptance or rejection of a plant by both species. The role of specific glucosinolates and differing structure-activity relationships is suggested.     

Colonization and hyperparasitism ofCotesia rubecula (Hym.: Braconidae), a newly introduced parasite ofPieris rapae,in Virginia

A Yugoslavian strain ofCotesia rubecula (Marshall) (Hymenoptera: Braconidae) was released in spring broccoli for control of the imported cabbageworm,Pieris rapae (L.), in Montgomery Co., Virginia, in 1987.C. rubecula reproduced and parasitized imported cabbageworm larvae in fall broccoli in 1987. It was found in moderate numbers in the summer and fall crops in 1988, and by fall had dispersed 0.8 km from the original release site. However, noC. rubecula were detected in our broccoli plots in Montgomery Country in 1989 or 1990. The hymenopteran hyperparasitesIsdromas lycaenae (Walker)(Ichneumonidae), Spilochalcis torvina (Cresson) (Chalcididae), andTetrastichus galactopus (Ratzeburg) (Eulophidae) were found attackingC. rubecula. Hyperparasites emerged from 31.8% and 41.1% of theC. rubecula cocoons in 1987 and 1988, respectively.T. galactopus was the most numerous hyperparasite in 1987, butS. torvina was dominant in 1988. Hyperparasite activity was low during early spring, but increased in July in both years. By August 1988, only hyperparasites emerged from theC. rubecula cocoons collected in the field. In spite ofC. rubecula's success in overwintering, hyperparasites, especiallyS. torvina, may be a limiting factor in the establishment ofC. rubecula in southwestern Virginia.      

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.     

Field ecology of freshwater macroalgae in pools and ditches,with special attention to eutrophication

A review is presented on the occurrence of macroalgae in dune pools, ditches and moorland pools in The Netherlands, which were studied in the period 1980–1987. There are clear effects of eutrophication on biomass as well as on species composition. A general trend is that under mesotrophic or moderately eutrophic conditions, the filamentous macroalgae form a diversified component of the primary producers in shallow water bodies. Especially spore forming genera asSpirogyra andOedogonium can be represented with up to about 20 species per site. In hardwater ditches alsoVaucheria is an important component, especially the sediment dwellingVaucheria dichotoma. Under distinctly eutrophic conditions dense floating algal masses contain only one or a few species asCladophora glomerata, Enteromorpha intestinalis, Hydrodictyon reticulatum. Under hypertrophic conditions the water surface is covered by dense blanketing mats of duckweeds in summer and autumn. Besides trophic factors, other factors as alkalinity and chloride concentration play a role in species composition. The results are discussed in the framework of water quality assessment of polder waters and possible applications for water purification or use as manuring substance. The need for experimental ecophysiological studies is underlined.     

Nutritional suitability and ecological relevance of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> and <Emphasis Type="Italic">Brassica oleracea</Emphasis> as foodplants for the cabbage butterfly, <Emphasis Type="Italic">Pieris rapae</Emphasis>

The thale cress, Arabidopsis thaliana, is considered to be an important model species in studying a suite of evolutionary processes. However, the species has been criticized on the basis of its comparatively small size at maturity (and consequent limitations in the amount of available biomass for herbivores) and on the duration and timing of its life cycle in nature. In the laboratory, we studied interactions between A. thaliana and the cabbage butterfly, Pieris rapae, in order to determine if plants are able to support the complete development of the herbivore. Plants were grown in pots from seedlings in densities of one, two, or four per pot. In each treatment, one, two, or five newly hatched larvae of P. rapae were placed on fully developed rosettes of A. thaliana. In a separate experiment, the same densities of P. rapae larvae were reared from hatching on single mature cabbage (Brassica oleracea) plants. Pupal fresh mass and survival of P. rapae declined with larval density when reared on A. thaliana but not on B. oleracea. However, irrespective of larval density and plant number, some P. rapae were always able to complete development on A. thaliana plants. A comparison of the dry mass of plants in different treatments with controls (= no larvae) revealed that A. thaliana partially compensated for plant damage when larval densities of P. rapae were low. By contrast, single cress plants with 5 larvae generally suffered extensive damage, whereas damage to B. oleracea plants was negligible. Rosettes of plants that were monitored in spring, when A. thaliana naturally grows, were not attacked by any insect herbivores, but there was often extensive damage from pulmonates (slugs and snails). Heavily damaged plants flowered less successfully than lightly damaged plants. Small numbers of generalist plant-parasitic nematodes were also recovered in roots and root soil. By contrast, plants monitored in a sewn summer plot were heavily attacked by insect herbivores, primarily flea beetles (Phyllotreta spp.). These results reveal that, in natural populations of A. thaliana, there is a strong phenological mismatch between the plant and most of its potential specialist insect herbivores (and their natural enemies). However, as the plant is clearly susceptible to attack from non-insect generalist invertebrate herbivores early in the season, these may be much more suitable for studies on direct defense strategies in A. thaliana.     

Host discrimination and evidence for a host marking pheromone in the almond seed wasp, Eurytoma amygdali

Laboratory experiments and observations on the oviposition behaviour of the almond seed wasp Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) revealed that the females of this species deposit a host-marking pheromone, immediately after an oviposition, by dragging the tip of their abdomen on the fruit surface. This pheromone enables them to discriminate between the infested and uninfested fruit and to select for oviposition the latter. Its primary function is apparently the prevention of repeated ovipositions in already infested fruit, thus contributing to the optimal utilisation of the available resources for larval development. The responses of individual females to different treatments of almonds, in a series of two-choice tests, revealed that the pheromone can be perceived by the females on direct contact and, when at high concentrations, also olfactorily from a short distance. The pheromone was present inside the abdomen and thorax of females but not of males, and, although water soluble, could not be entirely removed from heavily infested almonds when rinsed with water. Direct observations revealed that after an average of 3.7 successive visits to pheromone-bearing almonds, females were induced to walk or, most often, fly away from the experimental set-up. This suggests that the pheromone may also contribute to the dispersion of the wasps.

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Résumé Des expériences de laboratoire et des observations concernant le comportement de ponte de Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) ont montré que, immédiatement après la ponte, les femelles déposent une phéromone de marquage de l'hôte en trainant le bout de leur abdomen sur la surface de l'amande. Cette phéromone les rends capable de distinguer les fruits infectés des non-infectés dt de sélectionner pour la ponte les derniers. Apparemment, la fonction principale de cette phéromone est la prévention de la répétition des ovipositions dans les fruits déjà infectés et la répartition uniforme des oeufs dans les amandes, contribuant ainsi à la meilleure utilisation des ressources disponsibles pour le développement des larves. Des expériences de deux choix entre des fruits de différents traitements ont montré que la phéromone pouvait être perçue par les femelles par le direct contact et, quand elle était à hautes concentrations, par olfaction d'une courte distance. La phéromone était présente dans l'abdomen et dans le thorax des femelles, et bien qu'elle soit soluble à l'eau, elle ne pouvait pas s'éloigner entièrement par lavage des amandes sérieusement infectées à l'eau. Les observations ont démontré qu'après un numéro de 3.7 visites successives sur des amandes portant de la phéromone les femelles s'éloignaient du lieu de ponte en marchant ou, le plus souvant, en s'envolant, ce qui suggère que la phéromone contribue à la dispersion des femelles.

     

Insect-crop interactions in a diversified cropping system: parasitism by Aleochara bilineata and Trybliographa rapae of the cabbage root fly, Delia radicum, on cabbage in the presence of white clover

Parasitism of the cabbage root fly, Delia radicum (L.) by the staphylinid Aleochara bilineata Gyllenhal and the cynipid Trybliographa rapae Westwood was examined in a cabbage monoculture and a mixed stand of cabbage undersown with white clover. Number of overwintering cabbage root fly pupae per plant was consistently reduced in the mixed stand, and the incidence of plants attacked by cabbage root fly was either reduced or not different in the mixed stand compared to cabbage monoculture. For both parasitoids, the probability of D. radicum attacked plants having at least one parasitized pupa increased with density of cabbage root fly pupae around the plant. For A. bilineata, this positive relation between presence of parasitism and host density was consistently stronger in cabbage monoculture than in cabbage undersown with clover. Location of a host plant by T. rapae was not consistently affected by the presence of clover. D. radicum attacked plants situated in the cabbage and clover mixture were found by T. rapae as easily as in cabbage monoculture. Overall, the total risk of parasitism for a cabbage root fly pupa by A. bilineata was reduced in the mixed stand compared to the cabbage monoculture, whereas the risk of parasitism by T. rapae was not consistently affected by clover. For both parasitoids, intensity of parasitism showed a variable relationship with host density on individual plants attacked by the cabbage root fly. Overall, in spite of consistently lower total density of pupae in the mixed cabbage—clover than in cabbage monoculture, the density of unparasitized pupae was reduced by the presence of non-host plants only in two of the four experiments. The results emphasize the need to include not only herbivore and crop, but also other plant species as well as natural enemies when evaluating management methods.     

The responses of some Lepidoptera to labiate herb and white clover extracts

Herbs have been traditionally used as intercrops with crop plants on the assumption that their odour repels pest species. Alcohol extracts and essential oils of labiate herbs were tested in the laboratory for deterrent/repellent responses to ovipositing Plutella xylostella (L.) and feeding larvae of P. xylostella and Pieris brassicae L.Alcohol extracts of hyssop, rosemary, sage, thyme and white clover reduce oviposition by P. xylostella on pieces of brassica leaf in dual-choice tests. Essential oils of sage and thyme reduce oviposition on pieces of brassica leaf. Feeding, in dual-choice tests, by final instar P. xylostella and P. brassicae larvae is reduced by application of alcohol extracts of herbs.

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Résumé Des plantes médicinales ont traditionnellement été utilisées en association avec des plantes cultivées dans l'espoir que leur odeur repousserait les insectes nuisibles. Des extraits à l'alcool et des essences volatiles de labiées médicinales ont été essayés au laboratoire pour examiner leur effets répulsif et dissuadant pour la ponte de Plutella xylostella et l'alimentation des chenilles de P. xylostella et Pieris brassicae. Les extraits alcooliques d'hysope, de romarin, de sauge, de thym et de trèfle blanc réduisent la ponte de P. xylostella sur des morceaux de feuilles de chou traitées. Les essences volatiles de sauge et de thym réduisent la ponte de P. xylostella sur des morceaux de feuilles de chou dans des expériences de choix. L'alimentation des chenilles de dernier stade de P. xylostella et P. brassicae dans des expériences de choix est réduite par l'application d'extraits alcooliques des labiées médicinales.