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Proceedings of the 11th International Symposium on Insect-Plant Relationships SERIES ENTOMOLOGICA VOLUME 57 The titles published in this series are listed at the end of this volume. Proceedings of the 11 th International Symposium on Insect-Plant Relationships Edited by Jens Kvist Nielsen, Christian Kj<er and Louis M. Schoonhoven Reprinted from Entomologia Experimentalis et Applicata 104 (1) 2002 with additional indices SPRINGER-SClENCE+BUSINESS MEDIA, B.Y. Library of Congress Cataloging-in-Publication Data International Symposium on Insect-Plant Relationships (llth : 2001 : Helsing0r, Denmark) Proeeedings ofthe Eleventh International Symposium on Insect-Plant Relationships / edited by Jens Kvist Nielsen, Christian Kjaer, and Louis M. Sehoonhoven. p. em. -- (Series entomologiea ; v. 57) Held Aug. 4-10, 2001 in Helsing0f, Denmark. ISBN 978-90-481-6129-4 ISBN 978-94-017-2776-1 (eBook) DOI 10.1007/978-94-017-2776-1 1. Insect-plant relationships--Congresses. 1. Nielsen, Jens Kvist. II. Kjaer, Christian. III. Sehoonhoven, L. M. IV. Title. V. Series. 2002032154 ISBN 978-90-481-6129-4 Printed on acidlree paper AII Rights Reserved © 2002 Springer Science+Business Media Dordrecht Originally published by Kluwer Academic Publishers in 2002 Softcover reprint of the hardcover 1s t edition 2002 No part of material protected by this copyright notice may be reproduced or utilized in any form or by any means, electronic or mechanical, inc1uding photocopying, recording or by any information storage and retrieval system, without written permission from the copyright owner. Contents Preface vii Acknowledgements viii Neuronal basis for perception of chemical stimuli H. Mustaparta, Encoding of plant odour information in insects: peripheral and central mechanisms Chemosensory hases of behaviour 1.1 Glendinning, How do herbivorous insects cope with noxious secondary plant compounds in their diet? 15 J.J.A. van Loon, C.Z. Wang, J.K. Nielsen, R Gols & Y.T. Qiu, Flavonoids from cabbage are feeding stimulants for diamondback moth larvae additional to glucosinolates: Chemoreception and behaviour 27 Chemical basis for host plant selection J.A.A. Renwick, The chemical world of crucivores: lures, treats and traps 35 S.M. Cook, E. Bartlet, D.A. Murray & LH. Williams, The role of pollen odour in the attraction of pollen beetles to oilseed rape flowers 43 J. Harmatha & J. Nawrot, Insect feeding deterrent activity oflignans and related phenylpropanoids with a methylenedioxyphenyl (piperonyl) structure moiety 51 A. Barker, U. Schaffner & 1.-L. Boeve, Host specificity and host recognition in a chemically-defended herbivore, the tenthredinid sawfly Rhadinoceraea nodicomis 61 RR Smyth, D.W. Tallamy, J.A.A. Renwick & M.P. Hoffmann, Effects of age, sex, and dietary history on response to cucurbitacin in Acalymma vittatum 69 N. Lombarkia & S. Derridj, Incidence of apple fruit and leaf surface metabolites on Cydia pomonella oviposition 79 Behaviour and performance on plants B. Gabrys & W.E Tjallingii, The role of sinigrin in host plant recognition by aphids during initial plant penetration 89 S. Hannunen & B. Ekbom, Within species variation in host plant quality and movement behavior of Lygus rugulipennis nymphs 95 G. Powell & J. Hardie, Xylem ingestion by winged aphids 103 J. Jansson & B. Ekbom, The effect of different plant nutrient regimes on the aphid Macrosiphum euphorbiae growing on petunia 109 A. Smits, Performance of pine looper Bupalus piniarius larvae under population build-up conditions 117 Effects of plant defences on insects (individuals and populations) E. Haukioja, V. Ossipov & K. Lempa, Interactive effects of leaf maturation and phenolics on consump- tion and growth of a geometrid moth 125 J.K. Holopainen, Aphid response to elevated ozone and C02 137 P.M. J!Ilndrup, J.D. Barnes & G.R. Port, The effect of ozone fumigation and different Brassica rapa lines on the feeding behaviour of Pieris brassicae larvae 143 C. Muller, J.-L. Boeve & P.M. Brakefield, Host plant derived feeding deterrence towards ants in the turnip sawfly Athalia rosae 153 G. Sz6venyi, Qualification of grassland habitats based on their Orthoptera assemblages in the K6szeg Mountains (W-Hungary) 159 Effects of insects on plants (individuals and populations) P.B. McEvoy, Insect-plant interactions on a planet of weeds 165 Interactions between several trophic levels M. Hilker & T. Meiners, Induction of plant responses to oviposition and feeding by herbivorous arthropods: a comparison 181 L.S. Rayor & S. Munson, Larval feeding experience influences adult predator acceptance of chemically defended prey 193 F. Nogueira-de-Sa & J.R. Trigo, Do fecal shields provide physical protection to larvae of the tortoise beetles Plagiometrionafiavescens and Stolas chalybea against natural enemies? 203 Genetics of adaptations between plants and insects P.W. de Jong & J.K. Nielsen, Host plant use of Phyllotreta nemorum: do coadapted gene complexes playa role? 207 Evolution of insect-plant relationships J.M. Scriber, Evolution of insect-plant relationships: chemical constraints, coadaptation, and concor- dance of insect/plant traits 217 Conclusion L.M. Schoonhoven, Concluding remarks 237 Index of authors 241 General index 242 Listing of oral and poster presentations from SIPll 247 List of registered participants 256 Preface Scientific investigations of insect-plant relationships have expanded rapidly in recent years. The first International Symposium on Insect-Plant Relationships (SIP1) was held in Wageningen in 1958, and since 1986 (SIP6) these meetings have been held with regular three-year intervals. These meetings have hosted a number of important debates: on the relative role of nutrients and secondary plant substances, of qualitative and quantitative plant de fences, of predictable and unpredictable resources, of co-evolution and sequential evolution, and on the relationship between adult behaviour and larval performance, to mention only a few. These debates have been important and contributed greatly to the development of the field. In spite of this, we are still far from a situation where we can define all important differences between a good and a bad host plant for any insect. It is becoming clear, however, that knowledge at any level from small molecules to large ecosystems is required. It is also important to realize that plants are not passive food items for insects, but living organisms that respond in different ways to differential environmental conditions and to the presence of insects. Furthermore, the host plants provide environments where insects live exposed to the physical environment and to other organisms which may be competitors or predators. We should also be aware of the fact that interactions between insects and plants evolve, and that the present relationships are formed by past evolutionary interactions. This knowledge calls for an integrated approach to the studies of insect-plant relationships. The 11th International Symposium on Insect-Plant Relationships (SIPll) followed the tradition of previous SIP-meetings and covered topics of different levels from chemistry, physiology and ethology to ecology, genetics and evolution of insect-plant relationships. Thanks to the efforts of 127 participants from 27 countries from all over the world, the meeting lived up to the high reputation that these meetings have always had. The meeting was held from August 4-10,2001 at LO-skolen in Helsinglllr, Denmark and was organised by Jens Kvist Nielsen, Christian Kjrer and Niels Agerbirk. The symposium included 10 keynote lectures, 48 oral presentations and 60 posters. The organisers want to thank all the participants for their clarity during the presentations, their enthusiasm and positive interactions throughout the symposium. It was a pleasure to see you all in Helsinglllr. All speakers and poster presenters were invited to submit manuscripts to be published in the proceedings of SIPll and in a special issue of Entomologia Experimentalis et Applicata. All papers were then subjected to the full rigour of the normal peer-review procedure of the journal. This is a continuation of a trend adopted by previous organisers, and we agree with them that a book of fully refereed papers would provide more service to the field (authors and readers alike) than an unrefereed volume. The proceedings of SIPl1 include 24 full papers and one short communication. Contributions from all sessions but one (Molecular Ecology of Insect Plant Interactions) are included in the proceedings volume. For various reasons many valuable contributions to the symposium are not found in the proceedings, but these will probably be published soon elsewhere. We accept this situation as the best solution to the discrepancy created by the tight schedule for production of the proceedings volume, the full peer reviewing procedure, and the wish to create at the symposium a forum for aerating new and as yet only partially substantiated ideas. - We want to thank all authors for their collaboration and all reviewers for their constructive and insightful comments which have increased the quality of the proceedings tremendously. We want to thank the staff at LO-skolen in Helsinglllr for providing a friendly and stimulating environment for the Symposium, and Helle and Dorthe Klestrup from DIS Congress Service Copenhagen for keeping track of all the participants. Furthermore, we want to thank Peter W. de Jong for drawing the logo, and Bodil Thestrup, Kathe MlIlgelvang and Juana Jacobsen for help with the preparation of announcements, abstract booklet, and the symposium website. Finally we owe many thanks to the staff at Kluwer Academic Publishers, to Franc;oise Kaminker from the editorial office, and to Joop J.A. van Loon and Steph Menken from the editorial board of Entomologia for their help and advice during the preparation of this volume. This volume is dedicated to Liz Bernays and Reg Chapman for their outstanding contributions to the field of insect-plant relationships. JENS KVIST NIELSEN CHRISTIAN KJlER LOUIS M. SCHOONHOVEN Acknowledgements The organisers wish to thank the following institutions and organisations for their support and sponsorship: • Cheminova AlS • Torben & Alice Frimodts Foundation • The Danish Ministry of Foreign Affairs • Danida • Danish Environmental Research Institute, Department of Terrestrial Ecology • Chemistry Department, Royal Veterinary and Agricultural University • Danish Agricultural and Veterinary Research Council Entamalagia Experimentalis et Applicata 104: 1-13, 2002. © 2002 Kluwer Academic Publishers. Encoding of plant odour information in insects: peripheral and central mechanisms Hanna Mustaparta Department of Zoology, Neuroscience Unit, Norwegian University of Science and Technology, MTFS, N-7489 Trondheim, Norway Accepted: April 16, 2002 Key words: olfactory coding, olfactory receptor neurones, antennal lobe neurons, ordinary glomeruli, olfaction, chemical communication, insects, heliothine moths Abstract Insects are suitable model organisms for studying mechanisms underlying olfactory coding and olfactory learning, by their unique adaptation to host plants in which the chemical senses are essential. Recent molecular biological studies have shown that a large number of genes in insects and other organisms are coding for olfactory receptor proteins. In general, one receptor type seems to be expressed in each neurone. The functional characterisations of olfactory receptor neurones have been extensive in certain insect species, demonstrating a fine-tuning of single neurones to biologically relevant odourants; both insect and plant produced volatiles. Stained neurones of the same functional type have been shown to project in one and the same glomerular unit in the primary olfactory centre, the antennal lobe. This corresponds to molecular biological studies, showing projections in one glomerulus by neurones expressing the same receptor type. Comparison of these findings with physiological and morpholog ical characterisations of antennal lobe neurones has indicated correspondence between input and output of the glomerular units. Examples are presented from studies of heliothine moths. From the antennal lobe, the olfactory information is further conveyed to the mushroom bodies, particularly important for learning, and the lateral pro tocerebrum, a premotoric area. The three brain areas are regions of synaptic plasticity important in learning of odours, which is well studied in the honeybee but also in species of moths. Introduction and elicit responses in the neurones. Functional char acterisation of olfactory receptor neurones (ORNs) In evolutionary context, the sense of smell is consid requires electrophysiological recordings which, fortu ered as the oldest of our senses, by being present in nately, have been extensive in certain insect species. organisms of all phyla. Recent molecular biological By knowing the odourants to which the neurones re studies have shown that a large number of genes in spond, allows us to select compounds that might be various organisms codes for olfactory receptor pro biologically relevant and influence the behaviour of teins e.g., in humans 347 functional olfactory genes the insects in nature. Of course, not all electrophysi have been identified (Zozulya et aI., 2001). In contrast, ologically active compounds are relevant or detected the visual system operates with only three types of at physiological concentrations; this can only be de photoreceptors as the basis for colour vision. Thus the termined by behavioural studies. Thus, combinations ability to detect and discriminate different odours de of methods, involving molecular biology, electrophys pends on a large number of olfactory receptor proteins, iology, chemistry, and behaviour are all important for as shown in all organisms studied so far, including resolving the mechanisms involved in chemical com insects. However, simply knowing which receptor pro munication in insects and in insect-plant interactions. teins are present, as identified by molecular biological Insects are interesting model organisms for study methods, does not reveal which odourants they bind ing olfactory coding mechanisms, by their diverse 2 and often unique adaptation to their host plants which The most obvious level of functional organisa they locate and select by the use of chemical, vi tion is the separation of the macroglomerular com sual, and mechanical cues (Schoonhoven et aI., 1998). plex (MGC), which processes pheromone information, Unsuitable plants are avoided by detection of other from the ordinary glomeruli which process plant odour chemical cues. Pheromones are known in more than information. The number and position of ordinary 1000 insect species, and the identification of relevant glomeruli as well as MGC units in the antennallobes plant odourants is in progress in many species, as are constant within a species (Rospars, 1983; Rospars exemplified in this volume. & Hildebrand, 1992), and 3-D anatomical atlases of the glomeruli have been made in some species (Flana gan & Mercer, 1989; Galizia et aI., 1999a; Laissue Neuronal pathways et aI., 1999). In the two heliothine moths, Heliothis virescens and Helicoverpa assulta, the antennallobes An advantage in studying olfactory coding mecha have been reconstructed on the basis of synapsin anti nisms in insects is their relative simple and com body staining and laser scanning confocal microscopy, partmentalised nervous system where neurones can which have resulted in 3-D atlases of the glomeruli be identified, from the receptor neurones detecting (Berg et aI., 2002). It shows four MGC units in the chemical signals to interneurones integrating and H. virescens males, confirming previous results (Berg conveying information to motor neurones that elicit et aI., 1998; Vickers et aI., 1998; Christensen et aI., the behavioural responses. The ORNs are housed in 1991, 1995), and 62 ordinary glomeruli. The male numerous sensilla, mainly located on the antennae. antennal lobe of H. assulta contains only three MGC Among the various sensillum types identified in dif units and 62 ordinary glomeruli. The number of ordi ferent insect species, s. trichodea contain the ORNs nary glomeruli in the heliothine moths is comparable responding to pheromones and s. basiconica house to that in other moths species, 63 in Manduca sexta the ORNs for plant odour detection, as shown in the (Rospars & Hildebrand, 2000) and 67 in male and earliest studies of the silk moth (Schneider, 1992). In 68 in female Mamestra brassicae (Rospars, 1983). our research on olfactory coding mechanisms we are In contrast, a much larger number is found in hy studying moth species of the subfamily Heliothinae. menopteran species, 160 in the honeybee (Galizia Here the filiform female and male antennae differ by et ai., 1999a) and about 180-200 in parasitoid wasps the presence of s. trichodea type 1 exclusively in the (H.M. Smid, pers. commun.). males, whereas s. trichodea type 2 and s. basiconica The neuronal pathway of olfactory information in are present on the antennae of both sexes (Almaas & the brain has been studied in many insect species Mustaparta, 1990, 1991; Koh et aI., 1995). The ax (reviewed by Homberg et aI., 1989; Masson & ons of the receptor neurones form the antennal nerve Mustaparta, 1990; Anton & Homberg, 1999; Hansson which enters the antennal lobe, the primary olfactory & Christensen, 1999). As in other lepidopterans, the centre of the brain. The nerve divides into fascicles neurones of the antennal lobes in heliothine species and forms the glomerular structures together with the consist of two major types, the local interneurones dendrites of the antennal lobe neurones (reviewed by and the projection neurones (Christensen et aI., 1991, Boeckh & Tolbert, 1993; Hildebrand, 1996). The for 1995; Vickers et aI., 1998; Berg et aI., 1998). Both mation of glomeruli during development is dependent excitation and inhibition pathways have been demon on the presence of glia cells, and in the absence of strated: There is excitatory synaptic input from the glia the primary axons grow throughout the anten primary ORN axons to the local interneurones and to nallobe directly into protocerebrum (Baumann et aI., the projection neurones; there is inhibitory input from 1996; Oland et aI., 1988; Rossler et aI., 1999). It is the local interneurones to the projection neurones as in the glomeruli, which contain the synapses between well as excitatory input from projection neurones to receptor neurones and antennallobe neurones, that the local interneurones (Distler & Boeckh, 1996, 1997a,b; processing of odour information takes place in this pri Leitch & Laurent, 1996; Sun et aI., 1997). This cir mary centre. A central question in olfactory research cuitry may create excitation of local and projection is whether the glomeruli are functional units and, if so, neurones as well as inhibition and disinhibition of how they are functionally organised; for example, does projection neurones (Christensen et aI., 1993; Mac one glomerulus primarily process information about Leod & Laurent, 1996; Stopfer & Laurent, 1999; one odorant. Stopfer et aI., 1999). The local interneurones, inner-

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