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The Insects
Structure and Function
FIFTH EDITION
TheInsectshasbeenthestandardtextbookinthefieldsincethefirsteditionwas
publishedover40yearsago.BuildingonthestrengthsofChapman’soriginaltext,
thislong-awaitedneweditionhasbeenrevisedandexpandedbyateamofeminent
insectphysiologists,bringingitfullyuptodateforthemolecularera.
Thechaptersretainthesuccessfulstructureoftheearliereditions,focusingon
particularfunctionalsystemsratherthanontaxonomicgroupsandmakingiteasyfor
studentstodelveintotopicswithoutextensiveknowledgeoftaxonomy.Thefocusis
onformandfunction,bringingtogetherbasicanatomyandphysiologyand
examininghowtheserelatetobehavior.This,combinedwithnearly600clear
illustrations,providesacomprehensiveunderstandingofhowinsectswork.
NowalsofeaturingarichlyillustratedprologuebyGeorgeMcGavin,thisisan
essentialtextforstudents,researchersandappliedentomologistsalike.
R.F.Chapman(1930–2003)wasaneminentinsectphysiologistandProfessorinthe
DivisionofNeurobiologyattheUniversityofArizona.HisfirstfoureditionsofThe
Insectshaveformedthestandardtextinthefieldformorethan40years.
StephenJ.SimpsonisARCLaureateFellowintheSchoolofBiologicalSciencesand
AcademicDirectorofthePerkinsCentreforthestudyofobesity,diabetesand
cardiovasculardiseaseattheUniversityofSydney.Hiscoreresearchaimsareto
understandswarminginlocustsandtodevelopandimplementanintegrative
frameworkforstudyingnutrition.In2012hewasawardedtheWigglesworthMedal
fromtheRoyalEntomologicalSocietyofLondon.
AngelaE.DouglasisDaljitS.andElaineSarkariaProfessorofInsectPhysiologyand
ToxicologyatCornellUniversity,NewYork.Herresearchandteachingismotivated
bythemechanismsunderlyinginsectfunction,andhercoreresearchinterestsarethe
overlappingtopicsofinsectnutritionandinteractionsbetweeninsectsandbeneficial
microorganisms.SheisaFellowofTheRoyalEntomologicalSocietyandThe
EntomologicalSocietyofAmerica.
Comp.by:AbdulMalik Stage:Revises1 ChapterNo.:FrontMatter TitleName:CHAPMANSIMPSONANDDOUGLAS
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The Insects
Structure and Function
FIFTH EDITION
R. F. CHAPMAN
FormerlyoftheUniversityofArizona,USA
Edited by
STEPHEN J. SIMPSON
TheUniversityofSydney,Australia
ANGELA E. DOUGLAS
CornellUniversity,NewYork,USA
Comp.by:AbdulMalik Stage:Revises1 ChapterNo.:FrontMatter TitleName:CHAPMANSIMPSONANDDOUGLAS
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CAMBRIDGE UNIVERSITY PRESS
Cambridge,NewYork,Melbourne,Madrid,CapeTown,
Singapore,Sa˜oPaulo,Delhi,MexicoCity
CambridgeUniversityPress
TheEdinburghBuilding,CambridgeCB28RU,UK
PublishedintheUnitedStatesofAmericabyCambridgeUniversityPress,NewYork
www.cambridge.org
Informationonthistitle:www.cambridge.org/9780521113892
©CambridgeUniversityPress1998,2013
Thispublicationisincopyright.Subjecttostatutoryexception
andtotheprovisionsofrelevantcollectivelicensingagreements,
noreproductionofanypartmaytakeplacewithout
thewrittenpermissionofCambridgeUniversityPress.
FirstpublishedbyEdwardArnold1969
Secondedition1971,6thprinting1980
Thirdedition1982,5thprinting1991
FourtheditionpublishedbyCambridgeUniversityPress1998,7thprinting2011
Fifthedition2013
PrintedintheUnitedKingdombytheMPGBooksGroup
AcataloguerecordforthispublicationisavailablefromtheBritishLibrary
LibraryofCongressCataloguinginPublicationdata
Chapman,R.F.(ReginaldFrederick)
Theinsects:structureandfunction/R.F.Chapman.–5thedition/editedby
StephenJ.Simpson,AngelaE.Douglas.
pages cm
Includesbibliographicalreferencesandindexes.
ISBN978-0-521-11389-2
1. Insects. I. Simpson,StephenJ. II. Douglas,A.E.(AngelaElizabeth),1956– III. Title.
QL463.C482013
595.7–dc23 2012018826
ISBN978-0-521-11389-2Hardback
CambridgeUniversityPresshasnoresponsibilityforthepersistenceor
accuracyofURLsforexternalorthird-partyinternetwebsitesreferredto
inthispublication,anddoesnotguaranteethatanycontentonsuch
websitesis,orwillremain,accurateorappropriate.
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CONTENTS
Listofcontributors ix 4 Nutrition 81
Preface xi Introduction 81
Acknowledgments xii 4.1 Requirednutrients 82
Prologue xiii 4.2 Balanceofnutrients 87
4.3 Nutritionaleffectsongrowth,development,
Part I The head, ingestion, utilization reproductionandlifespan 95
and distribution of food 4.4 Contributionofsymbioticmicroorganisms
toinsectnutrition 98
1 Head 3
Summary 104
Introduction 3
1.1 Head 4
5 Circulatory system, blood and the
1.2 Neck 9
immune system 107
1.3 Antennae 10
Introduction 107
Summary 13
5.1 Thecirculatorysystem 108
5.2 Circulation 113
2 Mouthparts and feeding 15
5.3 Hemolymph 117
Introduction 15
5.4 Hemocytes 124
2.1 Ectognathousmouthparts 16
Summary 129
2.2 Mechanicsandcontroloffeeding 22
2.3 Regulationoffeeding 34
6 Fat body 132
2.4 Otherconsequencesof
Introduction 132
feeding 37
6.1 Fatbodystructureanddevelopment 133
2.5 Headglands 37
6.2 Storageandutilizationofenergy
Summary 43
andnutrients 137
6.3 Functionasanendocrineorganand
3 Alimentary canal, digestionand
nutritionalsensor 142
absorption 46
Summary 144
Introduction 46
3.1 Thealimentarycanal 47
Part II The thorax and locomotion
3.2 Digestion 59
3.3 Absorption 72 7 Thorax 149
3.4 Thealimentarytractasanimmunological Introduction 149
organ 77 7.1 Segmentationofthethorax 150
Summary 78 7.2 Morphologyofthethorax 151
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vi Contents
7.3 Musclesofthethorax 155 12 Reproductive system: male 282
Summary 155 Introduction 282
12.1 Anatomyoftheinternalreproductive
8 Legsandlocomotion 157 organs 283
Introduction 157 12.2 Spermatozoa 286
8.1 Structureofthelegs 158 12.3 Transferofspermto
8.2 Walkingandrunning 166 thefemale 292
8.3 Othermechanismsofterrestrial 12.4 Othereffectsofmating 306
locomotion 173 Summary 310
8.4 Aquaticlocomotion 180
8.5 Otherusesoflegs 186 13 Reproductive system: female 313
Summary 189 Introduction 313
13.1 Anatomyoftheinternalreproductive
9 Wingsand flight 193 organs 314
Introduction 193 13.2 Oogenesis 317
9.1 Structureofthewings 194 13.3 Ovulation 333
9.2 Formofthewings 204 13.4 Fertilizationoftheegg 333
9.3 Movementofthewings 207 13.5 Oviposition 335
9.4 Wingkinematics 214 Summary 343
9.5 Aerodynamicmechanisms 221
9.6 Powerforflight 223 14 The eggand embryology 347
9.7 Sensorysystemsforflightcontrol 225 Introduction 347
Summary 230 14.1 Theegg 348
14.2 Embryogenesis 357
10 Muscles 233 14.3 Alternativestrategiesofacquiringnutrients
Introduction 233 byembryos 379
10.1 Structure 234 14.4 Sexdetermination 388
10.2 Musclecontraction 242 14.5 Parthenogenesis 390
10.3 Regulationofmusclecontraction 244 14.6 Pedogenesis 392
10.4 Energeticsofmusclecontraction 252 Summary 393
10.5 Muscularcontrolintheintactinsect 254
10.6 Changesduringdevelopment 257 15 Postembryonic development 398
Summary 263 Introduction 398
15.1 Hatching 399
Part III The abdomen, reproduction 15.2 Larvaldevelopment 403
and development 15.3 Metamorphosis 417
15.4 Controlofpostembryonic
11 Abdomen 269
development 436
Introduction 269
15.5 Polyphenism 443
11.1 Segmentation 270
15.6 Diapause 448
11.2 Abdominalappendagesandoutgrowths 273
Summary 454
Summary 280
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Contents vii
Part IV The integument, gas exchange 19 Thermal relations 588
and homeostasis Introduction 588
19.1 Bodytemperature 589
16 Integument 463
19.2 Thermoregulation 595
Introduction 463
19.3 Performancecurves 598
16.1 Epidermis 464
19.4 Behaviorandsurvivalatlow
16.2 Thecuticle 469
temperatures 600
16.3 Chemicalcompositionofthecuticle 473
19.5 Activityandsurvivalathigh
16.4 Typesofcuticles 483
temperatures 607
16.5 Molting 488
19.6 Acclimation 610
16.6 Cuticleformation 493
19.7 Cryptobiosis 611
16.7 Functionsoftheintegument 497
19.8 Temperatureandhumidityreceptors 611
Summary 498
19.9 Temperature-relatedchangesinthe
nervoussystem 614
17 Gaseous exchange 501
19.10 Large-scalepatternsininsect
Introduction 501
thermalbiology 616
17.1 Trachealsystem 502
Summary 617
17.2 Spiracles 511
17.3 Cutaneousgasexchange 515
Part V Communication
17.4 Respiratorypigments 515
17.5 Gaseousexchangeinterrestrialinsects 516 A Physiological coordination within the
17.6 Gaseousexchangeinaquaticinsects 528 insect
17.7 Insectssubjecttooccasionalsubmersion 537
17.8 Gasexchangeinendoparasiticinsects 540 20 Nervous system 625
17.9 Otherfunctionsofthetrachealsystem 541 Introduction 625
17.10 Gasexchangeininsecteggs 542 20.1 Basiccomponents 626
Summary 542 20.2 Basicfunctioning 630
20.3 Anatomyofthenervoussystem 642
18 Excretionandsaltandwaterregulation 546 20.4 Brain 647
Introduction 546 20.5 Controllingbehavior 659
18.1 Excretorysystem 547 Summary 669
18.2 Urineproduction 552
18.3 Modificationoftheprimaryurine 555 21 Endocrine system 674
18.4 Controlofdiuresis 559 Introduction 674
18.5 Nitrogenousexcretion 562 21.1 Chemicalstructureofhormones 675
18.6 Detoxification 567 21.2 Endocrineorgans 684
18.7 Non-excretoryfunctionsoftheMalpighian 21.3 Transportofhormones 691
tubules 569 21.4 Regulationofhormonetiter 691
18.8 Nephrocytes 571 21.5 Modeofactionofhormones 696
18.9 Waterregulation 573 Summary 703
Summary 584
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viii Contents
B Perception ofthe environment 25.4 Colorpatterns 807
25.5 Colorchange 807
22 Vision 708 25.6 Significanceofcolor 813
Introduction 708 25.7 Lightproduction 817
22.1 Compoundeyes 709 Summary 821
22.2 Formandmotionvision 715
22.3 Receptorphysiology,colorand 26 Mechanical communication:producing
polarizationvision 721 sound and substrate vibrations 824
22.4 Dorsalocelli 731 Introduction 824
22.5 Stemmata 732 26.1 Natureandtransmissionofacousticand
22.6 Othervisualreceptors 734 vibrationalsignals 825
22.7 Magneticsensitivityandphotoreception 735 26.2 Significanceofacousticandvibrational
Summary 735 signals 826
26.3 Mechanismsproducingsoundsand
23 Mechanoreception 738 vibrations 832
Introduction 738 26.4 Patternsofacousticandvibrational
23.1 Cuticularmechanoreceptors 739 signals 845
23.2 Chordotonalorgans 748 26.5 Neuralregulationofsoundproduction 847
23.3 Stretchandtensionreceptors 764 Summary 853
Summary 768
27 Chemicalcommunication:pheromones
24 Chemoreception 771 and allelochemicals 857
Introduction 771 Introduction 857
24.1 Externalstructureofchemosensorysense 27.1 Definingchemicalsignals 858
organs 772 27.2 Pheromonesusedinintraspecific
24.2 Cellularcomponents 774 communication 858
24.3 Distributionandnumbersofsensory 27.3 Informationcontentofpheromonal
sensilla 776 signals 874
24.4 Howthechemosensorysensillum 27.4 Biosynthesisofpheromones 876
functions 776 27.5 Regulationofpheromoneproduction 882
24.5 Integratingfunctionandbehavior 788 27.6 Perceptionofpheromonesandother
24.6 Projectionstothecentralnervoussystem 789 infochemicals 883
Summary 791 27.7 Informationtransferbetweenspecies:
allelochemicals 885
C Communication with other organisms 27.8 Producing,storingandreleasing
allomones 887
25 Visual signals:color and light 27.9 Allelochemicalsusedindefense 890
production 793 27.10 Mimicry 895
Introduction 793 Summary 898
25.1 Thenatureofcolor 795
25.2 Structuralcolors 795
25.3 Pigmentarycolors 802 Index 901
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CONTRIBUTORS
LarsChittka MichaelF.Land
SchoolofBiologicalandChemicalSciences SchoolofLifeSciences
QueenMary,UniversityofLondon UniversityofSussex,BrightonUK
UK
TomMatheson
BronwenW.Cribb DepartmentofBiology
CentreforMicroscopy&Microanalysisand
UniversityofLeicester
SchoolofBiologicalSciences UK
TheUniversityofQueensland,Brisbane
Australia GeorgeC.McGavin
OxfordUniversityMuseumofNaturalHistory
AngelaE.T.Douglas
Oxford
DepartmentofEntomology
UK
CornellUniversity
Ithaca,NY JeremyMcNeil
USA DepartmentofBiology
UniversityofWesternOntario,London
JulianA.T.Dow
Canada
InstituteofMolecularCellandSystemsBiology
CollegeofMedical,Veterinary&LifeSciences
DavidJ.Merritt
UniversityofGlasgow
SchoolofBiologicalSciences
UK
TheUniversityofQueensland,Brisbane
JonF.Harrison Australia
SchoolofLifeSciences
HansMerzendorfer
ArizonaStateUniversity,AZ
FachbereichBiologie/Chemie,Osnabru¨ck
USA
Germany
RalfHeinrich
JocelynG.Millar
Abtl.Zellula¨reNeurobiologie
DepartmentofEntomology
Schwann-Schleiden-Forschungszentrum,Go¨ttingen
UniversityofCalifornia,Riverside
Germany
USA
DeborahK.Hoshizaki
DivisionKidney,Urologic&HematologicDiseases StuartReynolds
NIDDK,NationalInstitutesofHealth DepartmentofBiology&Biochemistry
Bethesda,MD UniversityofBath
USA UK
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x Contributors
StephenRogers GrahamK.Taylor
DepartmentofZoology DepartmentofZoology
UniversityofCambridge OxfordUniversity
UK UK
LeighW.Simmons JohnS.Terblanche
CentreforEvolutionaryBiology DepartmentofConservationEcology&Entomology
SchoolofAnimalBiology FacultyofAgriSciences
TheUniversityofWesternAustralia,Crawley StellenboschUniversity
Australia SouthAfrica
StephenJ.Simpson PeterVukusic
SchoolofBiologicalSciences SchoolofPhysics
TheUniversityofSydney UniversityofExeter
Australia UK
MichaelT.Siva-Jothy LutzT.Wasserthal
DepartmentofAnimalandPlantSciences Institutfu¨rZoologieI
UniversityofSheffield Universita¨tErlangen-Nu¨rnberg
UK Germany
JohnC.Sparrow
DepartmentofBiology
UniversityofYork
UK
MichaelR.Strand
DepartmentofEntomology
CenterforTropicalandEmergingGlobalDiseases
UniversityofGeorgia,GA
USA
Description:The Insects has been the standard textbook in the field since the first edition published over forty years ago. Building on the strengths of Chapman's original text, this long-awaited 5th edition has been revised and expanded by a team of eminent insect physiologists, bringing it fully up-to-date fo