Insect-Plant Biology
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1 Insect-Plant Biology Second Edition Louis M. Schoonhoven Joop J.A. van Loon Marcel Dicke Laboratory of Entomology, Wageningen University, The Netherlands Univers!".2is- i^.c! LancNa- I bibiiothok Darmyladt i Bibliothek Biologie 1 OXPORD UNIVERSITY PRESS
2 Contents Foreword to the second edition by Professor Elizabeth Bernays vii Preface to the second edition " ix Preface to the first edition ' x 1 Introduction Increased attention: why? Relationships between insects and plants Relevance for agriculture Insect-plant research involves many biological subdisciplines References 3 2 Herbivorous insects: something for everyone Host-plant specialization Food-plant range and host-plant range Specialization on plant parts Above-ground herbivory Below-ground herbivory Number of insect species per plant species Herbivorous insects: are they plant taxonomists? Host plant is more than food plant Microclimates around plants Extent of insect damage in natural and agricultural ecosystems Compensation for herbivore damage Conclusions References 24 3 Plant structure: the solidity of anti-herbivore protection Insect feeding systems Leaf surface * Epicuticular waxes Trichomes Leaf toughness Mandible wear C 3 and C 4 plants Structures involved in mutualistic relationships Plant galls 41
3 xii CONTENTS 3.6 Plant architecture Conclusions References 44 4 Plant chemistry: endless variety Plant biochemistry Primary plant metabolism Secondary plant substances Alkaloids Terpenoids and steroids Phenolics Glucosinolates Cyanogenics Leaf surface chemistry Plant volatiles v Concentrations of secondary plant substances Production costs Compartmentation Temporal variability _ Seasonal effects Day/night effects Interyear variation Effects of location and fertilizers Sun and shade Soil factors Induced resistance Induced direct resistance Induced indirect" resistance Variation in herbivore-induced changes Genomic and metabolomic changes induced by herbivory Systemic effects Long-term responses Signal transduction Interaction between herbivore-induced and pathogen-induced changes Plant-plant interactions Genotypic variation Inter-individual variation in plant chemistry Intra-individual variation in plant chemistry Plant sex affects insect susceptibility Conclusions Literature References 86 5 Plants as insect food: not the ideal Plants are suboptimal food Nitrogen Water Artificial diets 105
4 CONTENTS xiii 5.3 Consumption and utilization Food quantities eaten Utilization Suboptimal food and compensatory feeding behaviour Allelochemicals and food utilization Detoxification of plant allelochemicals Symbionts Food utilization and supplementation Detoxification of plant allelochemicals Host-plant quality affected by microorganisms Plant pathogens Endophytic fungi Host-plant effects on herbivore susceptibility to pathogens and insecticides Food-plant quality in relation to environmental factors Drought, Air pollution Conclusions References Host-plant selection: how to find a host plant Terminology Host-plant selection: a catenary process Searching mechanisms Orientation to host plants Optical versus chemical cues Visual responses to host-plant characteristics Olfactory responses to host plants Flying moths and walking beetles: two cases of olfactory orientation Chemosensory basis of host-plant odour detection Morphology of olfactory sensilla Olfactory transduction Olfactory electrophysiology and sensitivity Olfactory specificity and coding ~ Host-plant searching in nature Conclusions References Host-plant selection: when to accept a plant The contact phase of host-plant selection: elaborate evaluation of plant traits Physical plant features acting during contact Trichomes Surface texture Plant chemistry: contact-chemosensory evaluation The importance of plant chemistry for host-plant selection: a historical intermezzo 173
5 xiv CONTENTS 7.5 Stimulation of feeding and oviposition., Primary plant metabolites Plant secondary metabolites'promoting acceptance: token stimuli Generally occurring secondary plant metabolites acting as stimulants Inhibition of feeding and oviposition Deterrency as a general principle in host-range determination Host-marking as a mechanism to avoid herbivore competition Plant acceptability: a balance between stimulation and deterrency Contact-chemosensory basis of host-plant selection behaviour Contact chemoreceptors ' Gustatory coding Caterpillars as models for coding principles, Token stimulus receptors: unsurpassed specialists Sugar and amino acid receptors: detectors of nutrients Deterrent receptors: generalist taste neurons Peripheral interactions Host-plant selection by piercing-sucking insects Oviposition preference Host-plant selection: a three-tier system Evolution of the chemosensory system and host-plant preferences Conclusions References Host-plant selection: variation is the rule Geographical variation Differences between populations in the same region Differences between individuals Environmental factors causing changes in host-plant preference Seasonality Temperature Predation risks Internal factors causing changes in host-plant preference Developmental stage Insect sex affects food choice Experience-induced changes in host-plant preference Non-associative changes Associative changes Pre- and early-adult experience Adaptive significance of experience-induced changes in host preference Conclusions References The endocrine system of herbivores listens to host-plant signals Development Morphism Diapause 236
6 CONTENTS xv 9.2 Reproduction Maturation Mating behaviour Conclusions References Ecology: living apart together Effects of plants on insects Plant phenology Plant chemistry " Plant morphology Alternative food " Effects of herbivores on plants Above-ground and below-ground insect-plant interactions Microorganisms and insect-plant interactions Vertebrates and insect-plant interactions Indirect species interactions in communities Exploitative competition Apparent competition Trophic cascades Species interactions and phenotypic plasticity Top-down versus bottom-up forces Food webs and infochemical webs Food webs Infochemical webs Communities Why are so many herbivorous insect species 'rare'? Colonization Community development Molecular ecology Conclusions References Evolution: insects and plants forever in combat Fossilized records of insect-plant interactions Speciation Reproductive isolation Rates of speciation Reciprocal speciation Genetic variation in host-plant preference of insects Interspecific differences Intraspecific differences Preference-performance correlation Genetic variation and local host-plant adaptation Genetic variation in plant resistance against insects Selection and adaptation Evolution of insect diversity 292
7 xvi CONTENTS 11.7 Evolution of host-plant specialization Coping with plant secondary metabolites Competition Reduced mortality from natural enemies Phylogenetic relationships Reciprocal evolution of herbivorous insects and their host plants Criticism of the theory of co-evolution Support for the theory of co-evolution Conclusions References J Insects and flowers: mutualism par excellence Mutualism Flower constancy Flower recognition Flower handling Pollination energetics Distance Accessibility Temperature Food-source evaluation Reward strategy Signalling nectar status Pollinator movement within multiple-flower inflorescences Competition Evolution - Nature conservation Economy Conclusions References Insects and plants: how to apply our knowledge Which herbivorous insect species become pests and why? Characteristics of herbivorous pest species Consequences of crop-plant introductions Agricultural practices promote the occurrence of pest problems Host-plant resistance Host-plant resistance mechanisms Partial resistance Plant characteristics associated with resistance Methodology of resistance breeding Polycultures: why fewer pests? The disruptive-crop hypothesis The enemies hypothesis Trap-cropping and crop-weed systems Diversity as a guiding principle 350
8 CONTENTS xvii 13.4 Plant-derived insecticides and antifeedants Antifeedants Neem tree, azadirachtin Outlook for antifeedants as crop protectants 13.5 Weed control by herbivorous insects Opuntia and Salvinia Success rate of biological weed-control programmes 13.6 Conclusion: diversification holds the clue to the control of pestiferous insects 13.7 References Appendices A: Further reading Books that focus wholly or to a large extent on insect-plant interactions Proceedings of international symposia on insect-plant relationships B: Structural formulae of selected secondary plant compounds C: Methodology C.I Choice of plants and insects C.I.I Plants C.I.2 Insects C.2 Behaviour C.2.1 Olfactory orientation C.2.2 Feeding C.2.3 Oviposition C.3 Sensory physiology C.3.1 Ablation C.3.2 Electrophysiology C.4 Plant chemistry C.4.1 Headspace C.4.2 Leaf surface C.4.3 Plant interior C.4.4 Gene expression patterns C.5 References Taxonomic index Author index Subject index
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