Table Of ContentGenetics and the Behavior of
Domestic Animals
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Genetics and the
Behavior of Domestic
Animals
Third Edition
Edited by
Temple Grandin
Department ofAnimalSciences, Colorado State University,
Fort Collins, CO, UnitedStates
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Contents
Listofcontributors xv
Preface xvii
1. Behavioral genetics and animal science 1
TempleGrandinandMarkJ.Deesing
Introduction 1
Geneticsshapesbehavior 3
Geneticeffectsofdomestication 3
Basicgeneticmechanisms 6
Single-nucleotidepolymorphisms 6
Repeats 6
Copynumbervariations 6
Jumpinggenes 7
CodingDNA 7
NoncodingDNA,alsocalledregulatoryDNA 7
Exome 8
RNAtranscriptome 8
Denovomutations 8
Quantitativetraitloci 8
Haplotypes 9
Epigenetics 9
Lamarckism 9
Braingeneticsmorecomplexthanothertraits 10
Abriefhistoricalreviewofanimalbehaviorstudy 10
Behaviorism 11
Instinctsversuslearning 12
Ethology 13
Ethologyandbehaviorismprovidetoolstostudyemotionsand
behaviors 15
Neuroscienceandbehavior 15
Emotionalsystemsmotivatebehavior 16
Confusionofemotionalsystemsmayconfoundstudies 18
Geneticsandemotionalsystems 19
Interactionsbetweengeneticsandexperience 20
Interactionsbetweeninstinctualhard-wiredbehaviorand
experience 21
Theparadoxofnovelty 22
Reactiontonovelty 22
v
vi Contents
Geneticfactorsandtheneedfornovelty 23
Temperamentisnotjustaboutfear 25
Speciesdifferencesinemotionalreactionstosimilartests 26
Biologicalbasisoffear 27
Fearfulnessandthemotheringinstinctconflict 28
Nervoussystemreactivitychangedbytheenvironment 28
Tamingdoesnotchangenervoussystemreactivity 29
Domesticversuswildandfearresponses 30
Neoteny 30
Overselectionforspecifictraits 31
Overselectioninlivestock 32
Linksbetweendifferenttraits 33
Transgenicmice 34
Randomfactorsandnoninheritedeffectsonvariation 34
Concludingthoughts 34
References 35
Furtherreading 46
2. Behavioral genetics and animal domestication 49
PerJensenandDominicWright
Definitionofdomestication 49
Thedomesticatedphenotype 51
Evolutionarymechanismsindomestication 53
Geneticmechanismsindomestication 55
Selection 56
Inbreeding 57
Drift 59
Theroleofhybridizationindomestication 59
Geneticarchitectureofdomestication 60
Whattypesofmutationscausethedomesticphenotype? 63
Copynumbervariation 65
Mappinggenesforbehavior—top-downapproaches 66
Pedigreestudiesandheritabilityanalysis 67
Quantitativetraitlociandassociationmapping 67
Selectivesweepmapping 68
Mappinggenesforbehavior—bottom-upapproaches 69
Effectsofspecificmutations 70
SocialaggregationinC.elegans 70
ForaginginDrosophilalarvaeandApismellifera 71
Pigmentationandbehavior 72
Neurotransmittersandsocialbehavior 74
Differentmeanstosameends 76
Epigenetics 77
Epigeneticchangesandlong-termbehavioralmodifications 78
Epigeneticchangesindomestication 79
Closingremarks 81
References 82
Contents vii
3. How studying interactions between animal emotions,
cognition, and personality can contribute to improve
farm animal welfare 95
AlainBoissyandHansW.Erhard
Prefaceto“Howstudyinginteractionsbetweenanimalemotions,
cognition,andpersonalitycancontributetoimprovefarm
animalwelfare”byDr.TempleGrandin 95
References 96
Introduction 97
Doanimalsfeelemotions? 98
Whatisthenatureofanimalsentience?Whatisanemotion?
Whatisstress? 98
Fearandanxiety 100
Diversityoffear-elicitingevents 100
Diversityinfear-relatedresponses 101
Variouswaysofassessingfearandanxiety 102
Aneedforarefinedandstructuredmethodologyinfearstudies 103
Contributionofcognitivepsychologytoaccessanimalemotions 104
Influenceofcognitiveprocessesonemotions 105
Alterationofthejudgmentanddecision-makingbyemotions 106
Animalindividualityinemotions:theconceptofpersonality 108
Individualvariabilityinemotionalbehaviors 109
Consistencyoffear-relatedresponsesacrossdifferentfrightening
situations 109
Consistencyoffear-relatedresponsesovertime 110
Personalityasacomplexintermediatevariable 110
Pre-existingcharacteristicsoftemperament 111
Geneticbackgroundandgeneticmodelsofemotionalreactivity 111
Developmentalaspects 112
Personalityandhealth 112
Aconceptofpositivewelfarebasedonpositiveexperiences 113
Geneticsandselectivebreedingforalleviatingnegativeemotions 113
Elicitingpositiveemotionsbyenrichingtheenvironmentand
management 114
Environmentalenrichment 115
Positivehumancontactandtrainingprograms 116
Cognitiveenrichment:anewapproachbasedontheanimals’
appraisalabilities 116
Promotingpositiveexperiencestomitigatenegativeexperiences
andtoimproveanimalwelfareandhealth 118
Conclusion 119
References 120
4. Genetics and behavior during handling, restraint, and
herding 131
TempleGrandinandMarkJ.Deesing
Introduction 131
viii Contents
Principlesofherdbehavior 133
Visualandauditorysenses 134
Differencesinthestrengthofherdingbehaviorofcattleandsheep 135
Socialbehaviorandhandling 136
Flightzone 137
Workingtheflightzone 139
Measuringtemperamentinlivestock 139
Testsoriginallydesignedtomeasurefearfulnessincattle 140
Restrainttests 140
Exitspeedtests 141
Startletestsforcattle,sheep,andpigs 142
Teststhatmeasureseparationdistress,fear,andseeking 142
Isolationtests 142
Open-fieldtests 143
Approachanovelobjectornovelpersontest 143
Pentesting 144
Otherbehavioralindicatorswhichmayindicategenetic
differencesinfear 144
Eyewhite 144
Physiologicalmeasures 145
Productiontraits 145
Relationshipbetweenfearandattractiontonovelty 146
Animalswithafearfultemperamenthavemoreviolentreactions
toasuddennovelty 146
Animalreactiontonoveltyisveryspecific 147
Temperamentandstrengthofemotionalresponsesarehighly
heritable 147
Fieldobservationsonbreeddifferencesincattleandsheepbehavior 148
Problemswithcrossbredcattle 149
Sireandbreedeffectsonruminantbehavior 149
Individualdifferenceswithinabreed 150
Effectofpiggeneticsonbehaviorduringhandling 151
Lean,rapid-growingpigsaremoreexcitable 152
Behaviorproblemsincrossbredbeefcattleduringthe1990s 152
Animalsize,bodyshape,coatcolor,andtemperament 153
Bodytypeandtemperamentindomesticanimals 154
Typeandtemperamentinwildherdanimals 155
Coatcolorandtemperament 155
Hairwhorls 157
Hairwhorlheightandvigilance 160
Theeffectsofearlyexperienceonhandling 161
Effectsofenvironmentandexperienceonpigs 162
Excellentmemoryofaversiveexperiences 163
Traininghighlyreactiveanimalsbyintroducingnoveltygradually 164
Innatenervousfearfulnessorreactiontoseparationstress 165
Facilitatinganimalmovementinhandlingfacilities 166
Geneticeffectsonhandlingfacilitydesign 167
Conclusion 167
Contents ix
References 168
FurtherReading 180
5. Reproductive and maternal behavior of livestock 183
P.J.Chenoweth,F.J.McPhersonandA.J.Landaeta-Hernandez
Generalintroduction 183
Evolutionanddomestication 184
Livestockbehavior 185
Reproductivebehavior 186
Geneticconsiderations 192
Inbreeding 194
Unbalancedselectionandoverselection 195
Assistedreproductivetechnologies 196
Balancingresources 197
Maternalbehavior 197
Temperament 199
Earlyweaningandseparationofdamfromoffspring 201
Geneticaspectsofmaternalbehaviorinselectedlivestockspecies 203
Environmentandstress 207
Selectedareasofconcern 210
Reducedmalereproductivepotential 210
Injuriesandpathologies 211
Conclusion 213
References 214
6. Factors influencing the expression of behavior in the
domestic dog 229
DeborahL.Wells
Introduction 229
Nature 230
Breed 231
Sex 235
Cerebrallateralization 236
Nurture 239
Sourceofacquisitionandearlyexperiences 239
Training 241
Ownertraits 243
Conclusions 245
References 245
7. From mind to genome: the effect of domestication
on dog cognition and genetics 253
MorganFerrans,HannahSalomonsandBrianHare
Cognitivelyremarkable 255
Thedomesticationhypothesis 256
