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Encyclopedia of Sustainability Science and Technology Series Editor-in-Chief: Robert A. Meyers Matthew L. Spangler Editor Animal Breeding and Genetics A Volume in the Encyclopedia of Sustainability Science and Technology, Second Edition Encyclopedia of Sustainability Science and Technology Series Editor-in-Chief RobertA.Meyers The Encyclopedia of Sustainability Science and Technology series (ESST) addresses the grand challenge for science and engineering today. It provides unprecedented, peer-reviewed coverage in more than 600 separate articles comprising 20 topical volumes, incorporating many updates from the first edition as well as new articles. ESSTestablishes a foundation for the many sustainability and policy evaluations being performed in institutions worldwide. An indispensable resource for scientistsandengineers indeveloping new technologies and for applying existing technologies to sustainability, the EncyclopediaofSustainabilityScienceandTechnologyseriesispresentedat theuniversityandprofessionallevelneededforscientists,engineers,andtheir studentstosupportrealprogressinsustainabilityscienceandtechnology. Althoughtheemphasisisonscienceandtechnologyratherthanpolicy,the EncyclopediaofSustainabilityScienceandTechnologyseriesisalsoacom- prehensive and authoritative resource for policy makers who want to under- stand the scope of research and development and how these bottom-up innovationsmapontothesustainabilitychallenge. Matthew L. Spangler Editor Animal Breeding and Genetics A Volume in the Encyclopedia of Sustainability Science and Technology, Second Edition With64Figuresand43Tables Editor MatthewL.Spangler DepartmentofAnimalScience UniversityofNebraska Lincoln,NE,USA ISSN2629-2378 ISSN2629-2386(electronic) EncyclopediaofSustainabilityScienceandTechnologySeries ISBN978-1-0716-2459-3 ISBN978-1-0716-2460-9(eBook) https://doi.org/10.1007/978-1-0716-2460-9 ©SpringerScience+BusinessMedia,LLC,partofSpringerNature2023 Thisworkissubjecttocopyright.AllrightsaresolelyandexclusivelylicensedbythePublisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting,reuseofillustrations,recitation,broadcasting,reproductiononmicrofilmsorinany otherphysicalway,andtransmissionorinformationstorageandretrieval,electronicadaptation, computersoftware,orbysimilarordissimilarmethodologynowknownorhereafterdeveloped. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthis publicationdoesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexempt fromtherelevantprotectivelawsandregulationsandthereforefreeforgeneraluse. Thepublisher,theauthors,andtheeditorsaresafetoassumethattheadviceandinformationinthis bookarebelievedtobetrueandaccurateatthedateofpublication.Neitherthepublishernorthe authorsortheeditorsgiveawarranty,expressedorimplied,withrespecttothematerialcontained hereinorforanyerrorsoromissionsthatmayhavebeenmade.Thepublisherremainsneutralwith regardtojurisdictionalclaimsinpublishedmapsandinstitutionalaffiliations. ThisSpringerimprintispublishedbytheregisteredcompanySpringerScience+BusinessMedia, LLC,partofSpringerNature. Theregisteredcompanyaddressis:1NewYorkPlaza,NewYork,NY10004,U.S.A. Series Preface Our nearly 1000-member team recognizes that all elements of sustainability scienceandtechnologycontinuetoadvanceasdoesourunderstandingofthe needs for water, clean air, food, energy, and health and the relation of every singleaspectofthisvastbodyofknowledgetoclimatechange.OurEncyclo- pedia content is at a level for university students, professors, engineers, and other practicing professionals. It is gratifying for our team to note that our online First Edition has been heavily utilized as evidenced by over 500,000 downloads,whichofcourseisinadditiontoscientists’utilizationoftheprint Encyclopediaandprintsinglevolumes. Now we are pleased to have a Living Reference available online which assures the sustainability community that we are providing the latest peer- reviewedcontentcoveringthescienceandtechnologyofthesustainabilityof the earth, and we are also publishing the content as a series of individual topicalbooksforeasyusebythosewithaninterestinparticularsubjects.Our team covers the physical, chemical, and biological processes and measure- ments that underlie the earth system including pollution and remediation, health,andclimatechange,andwecomprehensivelycovereveryenergyand environmenttechnologyaswellasalltypesoffoodproduction,water,trans- portation,andthesustainablebuiltenvironment. Our team members include two Nobel Prize winners (McFaddden and Fischlin), two former Directors of the NSF (Colwell and Killeen), and the ChiefScientistoftheRockyMountainInstitute(Lovins).Andourmorethan 50eminentsectioneditorsandnowbookeditorsassurequalityofourselected authorsandtheirreviewpresentations. Theextentofourcoverageclearlysetsourprojectapartfromotherpubli- cationswhichnowexist,bothinextentanddepth.Infact,currentcompendia ofthescienceandtechnologyofseveralofthesetopicsdonotpresentlyexist, and yetthe content iscrucialtoany evaluation and planning for the sustain- abilityoftheearth.Itisimportanttonotethattheemphasisofourprojectison scienceandtechnologyandnotonpolicyandpositions.Rather,policymakers will use our presentations to evaluate sustainability options. Vital scientific issuesinclude:humanandanimalecologicalsupportsystems;energysupply and effects; the planet’s climate system; systems of agriculture, industry, forestry, and fisheries and the ocean; fresh water and human communities; wastedisposal;transportationandthebuiltenvironment;aswellasinfectious diseasesandthevarioussystemsonwhichtheydependandthebalanceofall v vi SeriesPreface ofthesewithsustainability.Inthiscontext,sustainabilityisacharacteristicofa process or state that can be maintained at a certain level indefinitely even as globalpopulationincreasesfrom7.7billion(2021)toward9.7billionby2050 and11billionaround2100(UNEstimate).Thispopulationgrowthisexpected mostly in Africa and India and southeast Asia, and the hope for decreasing malnutrition coupled with better diet, decreasing poverty, and an increase in wealthimpliessomethinglikea50%increaseinfooddemandbyasearlyas 2030. But, the UN forecasts that if recent trends continue, the number of people affected by hunger will surpass 840 million by 2030. If the world economygrowsatanaverageofanominal3%perannum,therewillbemore thanadoublinginsizeby2050.Thismeansthatglobalproductionofconcrete, steel,andothermetalsaswellasplastics,whichpresentlymakesupnearlya thirdofallgreenhouseemissions,willalsolikelydouble.Thebuildingofnew orexpandedcities,bridges,roads,dams,andmanufacturingplants,aswellas construction of sea walls and pumping capacity for climate adaptation in additiontoairconditioningwillberequired.Thismeansthattherearegoing tobesomerealproblemsinreducingormorelikelyevenslowingthegrowth ofgreenhousegasemissionsaswellasforenergy,agriculture,andwater.Itis increasingly clear that conflicting demands among biofuels, food crops, for- estation, and environmental protection will be difficult to reconcile. The “green revolution” was heavily dependent on fertilizers which are manufacturedusingincreasinglyexpensiveanddiminishingreservesoffossil fuels. In addition, about 70% of available freshwater is used for agriculture. Clearly,manynaturalresourceswilleitherbecomedepletedorscarcerelative to population. All of this highlights the need for the technology and science presentedinthisbookseries aswell astrulymajorincreases inresearchand development. PalmDesert,CA,USA RobertA.Meyers,Ph.D. August2022 Editor-in-Chief Volume Preface Demandforanimalproteinproductscontinuestoincreaseamidanincreasing global population and increased standards of living within numerous coun- tries.Atthesametime,contemporaryconcernsexistrelativetotheimpactof climatic changes on animals (both domestic and wild) and potential impacts that modern animal agriculture may have on the environment. As a conse- quence, the list of phenotypic traits that are of economic importance to livestock producers and consumers have changed over time. The general notionofsustainabilityexplicitlyincludeslivestockenterpriseeconomicvia- bility, and the future of animal breeding and genetics will require directly addressing improvements in firm-level economic outcomes in the context of changing production environments and consumer demands. The field of Animal Breeding and Genetics is well positioned to meet these challenges. Geneticselectioncreatescumulativeadvancestowardsadefinedtargetrequir- ingthatthetargetiswell-definedandcurrent.Newtoolsandmethods,suchas genomicsandgeneediting,builduponaframeworkforthegeneticevaluation and selection of animals that has been successful at creating phenotypic changeandimprovingnumeroustraitsofeconomicimportance. The aims of this volume are to provide primers relative to the theory and applicationofAnimalBreedingandGeneticsacrossnumerousspeciesandto provideinsightintoboththehistoryandfutureofthedisciplinewithrespectto addressingsustainability.Activescientistsinthefield,aspiringstudents,and those working in allied industries will find this work as a valuable resource relative to the current state and future directions of genetic selection and managementeffortsacrossnumerousspecies. I would like to thank all of the authors who contributed their time and expertisetothisvolume.Tohavesuchanaccomplishedlistofinternationally recognizedauthorsisimpressiveandverymuchappreciated.Iwouldalsolike to express my sincere appreciation to Ms. Sunali Mull for her help and guidancethroughouttheprocessofcompletingthiswork. Lincoln,USA MatthewL.Spangler November2022 vii Contents AnimalBreedingandGenetics:Introduction ................. 1 MatthewL.Spangler AnimalBreedingMethodsandSustainability ................. 5 AgustinBlasco QuantitativeMethodsAppliedtoAnimalBreeding ............ 25 GuilhermeJ.M.Rosa FoundationsofMolecularGenetics:FromMajor GenestoGenomics ...................................... 51 RalucaG.Mateescu BuildingGeneticModels .................................. 63 L.DaleVanVleck GenotypebyEnvironmentInteractionsinLivestock Farming ............................................... 77 FrancescoTiezziandChristianMaltecca SociallyAffectedTraits,InheritanceandGenetic Improvement ........................................... 99 PiterBijma PigBreedingforIncreasedSustainability .................... 139 PieterW.Knap PoultryBreeding ........................................ 181 SamuelE.AggreyandRomdhaneRekaya BeefCattleBreeding ..................................... 191 DonaghBerry NovelTraitSelectiontoEnhanceSustainabilityofBeef ProductionSystems ...................................... 223 ElizabethDressler,JasonE.Sawyer,andMeganM.Rolf DairyCattleBreeding .................................... 243 JennieE.Pryce SustainableGeneticImprovementinDairyGoats ............. 261 ErinMassender,LuizF.Brito,andFlavioS.Schenkel ix x Contents HorseBreeding ......................................... 279 JessicaL.Petersen BreedinginDevelopingCountriesandTropics ................ 297 S.W.P.Cloete SustainabilityofWildPopulations:AConservationGenetics Perspective ............................................. 355 JenniferM.Thomson BreedinginanEraofGenomeEditing ...................... 369 EmilyLouiseClark Long-TermChallengesforAnimalBreeding .................. 385 RobertBanks Index .................................................. 411