Mechanisms ofWoody Plant Defenses Against Insects William J. Mattson, Jean Levieux, C. Bernard-Dagan Editors Mechanisms of Woody Plant Defenses Against Insects Search for Pattern With 106 Illustrations Springer-Verlag New York Berlin Heidelberg London Paris Tokyo William J. Mattson U.S. Department of Agriculture Forest Service North Central Forest Experiment Station East Lansing, Michigan 48823, USA Jean Levieux Institut National de la Recherche Agronomique Station de Zoologie Forestiere Ardon, 45160 Olivet. France C. Bernard-Dagan Universite de Bordeaux I Laboratoire de Physiologie Cellulaire Vegetale Avenue des Facultes 33405 Talence Cedex, France Library ofCongress Cataloging-in-Publication Data Mechanisms ofwoody plant defenses against insects. Ine!udes index. I. Woody plants-Disease and pest resistance. 2. Insect pests. I. Mattson, William J. II. Levieux. Jean. III Bernard-Dagan. C. SB76I.M46 1988 634.9'67 87-32102 © 1988 by Springer-Verlag New York Inc. Softcoverreprintofthehardcover1stedition1988 All rights reserved. This work may not be translated or copied in whole or in part without the written permissionofthe publisher(Springer-Verlag, I7S FifthAvenue. New York. NY 10010. USA). except for briefexcerptsinconnectionwith reviewsorscholarlyanalysis. Use inconnectionwithany form ofinformation storage and retrieval. electronic adaptation. computer software. or by similar or dissimilar methodology now known or hereafterdeveloped is forbidden. The use ofgeneraldescriptive names. trade names, trademarks. etc. in this publication. even ifthe former are notespeciallyidentified. is not tobe takenasasignthatsuch names. as understood bytheTradeMarks and Merchandise Marks Act, may accordingly be used freely by anyone. Camera-readycopy provided by the editors. 9 8 7 6 S 4 3 2 I ISBN-I3:978-I-4612-8368·3 e-ISBN-I3:978-1-4612-3828-7 001:10.1007/978-1-4612-3828-7 Preface In 1984, a new research working party on mechanisms of woody plant resistance against insects and pathogens (lUFRO S2.05-06) was formed in the International Union of Forestry Research Organizations. The purpose ofthis working party is to stimulate and facilitate research progress in this area through enhanced communication among the many research scientists scattered among the various countries of the world. This volume is the result ofthe first international symposium ofthis working party, which took place in Orleans. France on August 26-29, 1986. Thanks are due to Institute National de la Recherche Agronomique, Station de Zoologie, Forestiere, for their in strumental part in organizing this symposium, and to the University and the City of Orleans for generous financial support. The intent of this proceedings is to (a) bring together a diverse array of research results on the mechanisms ofwoody plant resistance against different kinds ofinsects. and (b) to search for threads of commonality among these different plant/insect as sociations to further our fundamental understanding ofhow plants defend themselves against phytophagous organisms. The research papers are organized into three main groups. The first group examines plant defenses from various basic physiological and ecological considerations. The remaining papers, which are mainly case studies of plant resistance against insects. are categorized on the basis of insect intimacy with host tissues: (a) highly mobile. free feeders such as moth larvae and sawflies, and (b) poorly mobile, "attached" or imbedded feeders such as aphids, scales, miners, and bark beetles. Each ofthese sections is further organized on the basis ofkind oftissue attacked by the insects, either leaves or stern/shoot cortex and vascular tissues. Contents Preface v Contributors IX I. Basic Physiological and Ecological Considerations I. Defensive Strategies of Woody Plants Against Different Insect-Feeding Guilds in Relation to Plant Ecological Strategies and Intimacy of Association with Insects W.J. Mattson. R.K. Lawrence, R.A. Haack, D.A. Herms, and p.J. Charles 3 2. Towards a Unified Theory of Plant Defense A.A. Berryman........................................................... 3lJ 3. Defensive ResponsesofTrees in Relation toTheirCarbon/Nutrient Balance J. Tuomi, P. Niemela, F.S. Chapin, 1II, J.P. Bryant, and S. Siren 57 4. Growth and Differentiation·Balance Relationships in Pines AffectTheir Re· sistance to Bark Beetles (Coleoptera: Scolytidae) P.L. Lorio, Jr. 73 5. Seasonal Variations in Energy Sources and Biosynthesis ofTerpenes in Maritime Pine C. Bernard-Dagan 93 6. Terpene Biosynthesis Under Pathological Conditions C. Cheniclet, C. Bernard-Dagan, and G. Pauly 117 7. Resistance of Plants at the Population Level to Attack by Phytophagous Insects J. Lunderstadt 13I II. Defenses Against Free-Feeding Insects 8. Induced Defenses in Ponderosa Pine Against Defoliating Insects M.R. Wagner............................................................. 141 VIII 9. Changes in Levels of Foliar Minerals and Phenolics in Trembling Aspen, Populus tremuloides. in Response'to Artificial Defoliation W.J. Mattson and S.R. Palmer........................................... 157 10. The Role of Resin Acids in the Relationship Between Scots Pine and the Sawfly, Diprion pini (Hymenoptera: Diprionidae). I.-Resin Acids in the Needles L. Buratti, J.P. Allais, and M. Barbier................................... 171 II. The Role of Resin Acids in the Relationship Between Scots Pine and the Sawfly, Diprion pini (Hymenoptera: Diprionidae). B.-Correlations with the Biology of Diprion pini C. Geri, L. Buratti, and J.P. Allais 189 12. Variations in Nutrient Levels as a Defense: Identifying Key Nutritional Traits of Host Plants of the Western Spruce Budworm K.M. Clancy, M.R. Wagner, and R.W. Tinus 2m 13. Phenological Resistance ofOaks to the Green Oak Leafroller, Tortrix viridana (Lepidoptera: Tortricidae) P. Du Merle 215 14. Wound Response of Living Bark of Scots Pine Seedlings and Its Influence on Feeding by the Weevil, Hylobius abietis A. Ericsson, R. Gref, C. Hellqvist. and B. Uingstr6m 227 Ill. Defenses Against Attached, Imbedded Insects 15. Interactions Between the Leaf Miner, Phyllocnistis suffusella, and Poplars L. Nef 239 16. A Resistance Response of Picea excelsa to the Aphid, Adelges abietis (Homoptera: Aphidoidea) O. Rohfritsch 253 17. Variations in Susceptibility of Pinus pinaster to Matsucoccusfeytaudi (Homoptera: Margarodidae) D. Schvester .......... 267 18. Beech Resistance to the Beech Scale: A Variety of Defenses D. Wainhouse, I.M. Gate, and D. Lonsdale 277 19. Resistance Mechanisms of Loblolly and Shortleaf Pines to Southern Pine Beetle Attack S.P. Cook and F.P. Hain 295 20. Wound-Induced Oleoresins ofAbies concolor: Is It Part of Host Resistance to the Fir Engraver, Scolytus ventralis? G.T. Ferrell __.................................... 305 ix 21. Elicitation of Defensive Reactions in Conifers F. Lieutier and A.A. Berryman 313 22. Do Fungi Influence the Establishment of Bark Beetles in Scots Pine? F. Lieutier, A. Yart, J. Garcia, B. Poupinel, and J. Levieux 321 23. The Toxicity of Norway Spruce Monoterpenes to Two Bark Beetle Species and Their Associates C. Everaerts, J-c. Gregoire, and J. Merlin 335 24. Traumatized Hosts: Their Influence on the Population Dynamics of the Southern Pine Bark Beetle Guild R.O. Flamm and R.N. Coulson 345 25. The Role of Spruce Monoterpene Derivatives as Oviposition Stimuli for Rhizophagus grandis. a Predator of the Bark Beetle, Dendroctonus micans M. Baisier, J.-c. Gregoire, K. Dehnte, and O. Bonnard 359 26. Host Orientation Behavior of Dendroctonus ponderosae: Integration of Token Stimuli Host and Defenses K.F. Raffa........................................................... ..... 369 27. Investigating the Hypothesis of Primary Attraction in Conifer-Specific Scolytidae Using a Chemometrical Approach D.N. Rutledge, C. Chararas, and C.J. Ducauze 31)\ Index.......................................................................... 4\1 Contributors Jean-Pierre Allais. Centre National de la Recherche Scientifique (CNRS), Institut de Chimie des Substances Naturelles (ICSN) 91190 GIF/YVETTE, France Marianne Baisier. Laboratoire de Biologie Animale etCellulaire. Universite Libre de Bruxelles. 50. Av. Franklin Roosevelt, 1050 Bruxelles, Belgium Michel Barbier. Centre National de la Recherche Scientifique (CNRS), Institut de Chimie des Substances Naturelles (ICSN), 91190 GIFIYVETTE, France Colette Bernard-Dagan. Laboratoire de Physiologie Cellulaire Vegetale. Universitc de Bordeaux I, Avenue des Facultes, 33405 Talence Cedex, France Alan A. Berrvman. Department of Entomology, Washington State University, Pullman. WA 99164, U.S.A. Odile Bonnard. Laboratoire de Zoologie, Universite de Dijon, Boulevard Gabriel, F 21 100 Dijon, France John P. Bryant. Institute of Arctic Biology, University ofAlaska, Fairbanks, Alaska 99701, U.S.A. Lionel Buratti. Centre National de la Recherche Scientifique (CNRS), Institut de Chimie des Substances Naturelles (ICSN), 91190 GIF/YVETTE, France F. Stuart Chapin. Ill. Institute of Arctic Biology, University of Alaska, Fairbanks, Alaska 99701, U.S.A. Constantin Chararas. Ministere de ragriculture, Institut National Agronomique, Paris Grignon, Laboratoire de Chimie Analytique Pierre-Jean Charles. Deceased, I.N.R.A., Stationde Zoologie Forestiere, Ardon. 45160 Olivet, France Catherine Cheniclet. Laboratoire de Physiologie Cellulaire Vegetale, Universite de Bordeaux I, Avenue des Facultes, 33405 Talence Cedex, France Karen M. Clancy. U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station, Forestry Sciences Laboratory, 700 S. Knoles Dr., Flagstaff, AZ 86001, U.S.A. xii Stephen P. Cook, Department of Entomology, Box 7626, North Carolina State Uni versity, Raleigh, NC 27695, U.S.A. Robert N. Coulson, Department of Entomology, Texas A & M University, College Station, TX 77843, U.S.A. Karin Delinte, Laboratoire de Zoologie, Universite de Dijon, Boulevard Gabriel, F 21100 Dijon, France Pau. Du Merle, I.N.R.A., Station de Zoologie Forestiere, Avenue A. Vivaldi, F 84000 Avignon, France Christian J. Ducauze, Ministere de l'agriculture, Institut National Agronomique, Paris Grignon, Laboratoire de Chimie Analytique Anders Ericsson, Department of Forest Genetics and Plant Physiology, Swedish Uni versity of Agricultural Sciences, S-90183 Umea, Sweden Claude Everaerts. Laboratoire de Zoologie, Universite de Bourgogne, 6 Boulevard Gabriel, F-21 100 Dijon, France George T. Ferrell. Research Entomologist, U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range ExperimentStation, Berkeley. California 94701, U.S.A. Richard O. Flamm. Department of Entomology, Texas A & M University, College Station, TX 77843, U.S.A. JacquesGarcia, I.N.R.A., Stationde Zoologie Forestiere, Ardon, 45160Olivet, France Imogen M. Gate. Forestry Commission Research Station, Alice Holt Lodge, Wrec c1esham, Farnham, Surrey, GU10 4LH, UK Claude Geri, I.N.R.A. Station de Zoologie Forestiere, Ardon, 45160 Olivet, France Jean-Claude Gregoire, Laboratoire de Biologie Animaleet Cellulaire, CP 160, Univ ersite Libre de Bruxelles, 50 Av. F.D. Roosevelt, B-1050 Bruxelles, Belgium RolfGre}: Department of Forest Genetics and Plant Physiology, Swedish University of Agricultural Sciences, S-90183 Umea, Sweden Robert A. Haack, U.S. Department ofAgriculture, Forest Service, North Central Forest Experiment Station, 1407 S. Harrison, East Lansing, MI 48823, U.S.A. Fred P. Hain. Department of Entomology, Box 7626, North Carolina State University, Raleigh, NC 27695, U.S.A. Claes Hellqvist. Department of Plant and Forest Protection, Swedish University of Agricultural Sciences, S-77073 Garpenberg, Sweden DanielA. Herms, Department ofEntomology, Michigan State University, East Lansing, MI, 48824, U.S.A. XIII Robert K. Lawrence, Department of Entomology, Michigan State University, East Lansing, MI, 48824, U.S.A. BoUingstrom, Department ofPlant and Forest Protection, Swedish University ofAg ricultural Sciences, S-77073 Garpenberg, Sweden Jean Levieux, I.N.R.A., Station de Zoologie Forestiere, Ardon, 45160 Olivet, France Franr,;ois Lieutier, I.N.R.A, Station de Zoologie Forestiere, Ardon, 45160 Olivet, France DavidLonsdale, Forestry Commission Research Station, Alice Holt Lodge, Wreccle sham, Farnham, Surrey, GUto 4LH, UK Peter L. Lorio, Jr., Research Forester, Forest Insect Research, U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station, 2500 Shreveport Highway, Pineville, Louisiana 71360, U.S.A. Ji)rf?, Lunderstiidt, Institut fUr Forstzoologie der Universitat Gottingen, Busgenweg 3, 0-3400 Gottigen William l. Mattson, U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station, 1407 S. Harrison, East Lansing, MI 48823, U.S.A. loe! Merlin, Laboratoire de Biologie Animale et Cellulaire, CP 160, Universite Librc de Bruxelles, 50 Av. F.D. Roosevelt, B-1050 Bruxelles, Belgium Ludovic Nef Centre pour la Lutte Integree contre les Insectes Forestiers, AGRO/EFOR. Universite Catholique de Louvain, Place Croix du Sud, 2 1348 Louvain-Ia-Neuvc. Belgium Pekka Niemelii, DepartmentofBiology, University ofTurku, SF-20500 Turku, Finland Sean R. Palmer, Department of Entomology, Hodson Hall, University of Minnesota, St. Paul, MN, 55108, U.S.A. Ginette Pauly, Laboratoirede PhysiologieCellulaire Vegetale, Universitede Bordeaux I, Avenue des Facultes, 33405 Talence Cedex, France Blandine Poupinel, I.N.R.A., Station de Zoologie Forestiere, Ardon, 45160 Olivet, France Kenneth F. Raffa, Department ofEntomology, University ofWisconsin, Madison, WI 53706, U.S.A. Odette Rohfritsch, Laboratoire de Cecidologie, U.A. C.N.R.S. 04 1182, Institut de Botanique, 28 rue Goethe, 67083 Strasbourg, France DouRlas N. Rutledge, Ministere de ('agriculture, Institut National Agronomique, Paris Grignon, Laboratoire de Chimie Analytique Daniel Schvestcr, I.N.R.A., Station de Zoologie Forestiere, Avenue Vivaldi, 84000 Avignon, France