Table Of Content

Genetics and the Behavior of Domestic Animals This pageintentionallyleftblank Genetics and the Behavior of Domestic Animals Third Edition Edited by Temple Grandin Department ofAnimalSciences, Colorado State University, Fort Collins, CO, UnitedStates AcademicPressisanimprintofElsevier 125LondonWall,LondonEC2Y5AS,UnitedKingdom 525BStreet,Suite1650,SanDiego,CA92101,UnitedStates 50HampshireStreet,5thFloor,Cambridge,MA02139,UnitedStates TheBoulevard,LangfordLane,Kidlington,OxfordOX51GB,UnitedKingdom Copyright©2022ElsevierInc.Allrightsreserved. Nopartofthispublicationmaybereproducedortransmittedinanyformorbyanymeans, electronicormechanical,includingphotocopying,recording,oranyinformationstorageand retrievalsystem,withoutpermissioninwritingfromthepublisher.Detailsonhowtoseek permission,furtherinformationaboutthePublisher’spermissionspoliciesandour arrangementswithorganizationssuchastheCopyrightClearanceCenterandtheCopyright LicensingAgency,canbefoundatourwebsite:www.elsevier.com/permissions. Thisbookandtheindividualcontributionscontainedinitareprotectedundercopyrightbythe Publisher(otherthanasmaybenotedherein). Notices Knowledgeandbestpracticeinthisfieldareconstantlychanging.Asnewresearchand experiencebroadenourunderstanding,changesinresearchmethods,professionalpractices, ormedicaltreatmentmaybecomenecessary. Practitionersandresearchersmustalwaysrelyontheirownexperienceandknowledgein evaluatingandusinganyinformation,methods,compounds,orexperimentsdescribedherein. Inusingsuchinformationormethodstheyshouldbemindfuloftheirownsafetyandthesafety ofothers,includingpartiesforwhomtheyhaveaprofessionalresponsibility. Tothefullestextentofthelaw,neitherthePublishernortheauthors,contributors,oreditors, assumeanyliabilityforanyinjuryand/ordamagetopersonsorpropertyasamatterof productsliability,negligenceorotherwise,orfromanyuseoroperationofanymethods, products,instructions,orideascontainedinthematerialherein. ISBN:978-0-323-85752-9 ForInformationonallAcademicPresspublications visitourwebsiteathttps://www.elsevier.com/books-and-journals Publisher:NikkiPLevy AcquisitionsEditor:AnnaValutkevich EditorialProjectManager:LindsayLawrence ProductionProjectManager:NiranjanBhaskaran CoverDesigner:ChristianJ.Bilbow TypesetbyMPSLimited,Chennai,India 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