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Sturkie’s Avian Physiology Seventh Edition Edited by Colin G. Scanes Department of Biological Science, University of Wisconsin, Milwaukee, WI, United States; Center of Excellence for Poultry Science, University of Arkansas, Fayetteville, AR, United States Sami Dridi Center of Excellence for Poultry Science, University of Arkansas, Fayetteville, AR, United States Academic PressisanimprintofElsevier 125London Wall,LondonEC2Y5AS,UnitedKingdom 525BStreet,Suite1650,SanDiego,CA92101,UnitedStates 50HampshireStreet,5thFloor,Cambridge,MA02139,UnitedStates TheBoulevard,Langford Lane,Kidlington,OxfordOX5 1GB,UnitedKingdom Copyright©2022ElsevierInc.Allrightsreserved. Nopart ofthispublicationmay bereproduced ortransmitted inanyform orbyanymeans, electronicor mechanical,including photocopying, recording,oranyinformation storageandretrieval system,withoutpermission inwritingfromthepublisher. Details onhowtoseek permission, furtherinformation aboutthePublisher’spermissions policiesandourarrangements with organizations suchastheCopyrightClearanceCenterandtheCopyrightLicensing Agency,canbefoundatourwebsite:www. elsevier.com/permissions. Thisbookandtheindividual contributionscontainedinitareprotected undercopyrightbythePublisher (otherthanasmay benotedherein). Notices Knowledgeandbestpracticeinthisfieldareconstantlychanging. As newresearchandexperiencebroadenourunderstanding, changesinresearch methods,professional practices,ormedical treatmentmay becomenecessary. Practitionersandresearchers mustalwaysrelyontheir ownexperience andknowledgeinevaluatingandusingany information,methods,compounds,orexperiments describedherein. Inusingsuchinformation ormethodsthey shouldbe mindfuloftheirown safetyandthesafetyofothers,including partiesforwhom theyhaveaprofessional responsibility. Tothefullestextentofthelaw,neither thePublishernortheauthors,contributors, oreditors, assumeany liabilityforany injuryand/ordamagetopersonsorpropertyasamatterofproductsliability,negligence orotherwise,or fromanyuseor operation ofanymethods,products, instructions,or ideascontainedinthematerialherein. LibraryofCongressCataloging-in-Publication Data Acatalogrecord forthisbook isavailablefromtheLibrary ofCongress BritishLibraryCataloguing-in-Publication Data Acataloguerecord forthisbook isavailablefromtheBritishLibrary ISBN:978-0-12-819770-7 Forinformation onallAcademic Presspublications visitourwebsite athttps://www.elsevier.com/books-and-journals Publisher:Charlotte Cockle Acquisitions Editor: AnnaValutkevich EditorialProjectManager:Emerald Li ProductionProjectManager:Selvaraj Raviraj CoverDesigner: Mark Rogers TypesetbyTNQTechnologies To our mentors who have guided and inspired us and our families who have given us so much support. Contributors N.J. Beausoleil, Animal Welfare Science and Bioethics Environment, The Hebrew University, Rehovot, Israel; Centre, School of Veterinary Science, Massey Univer- Miloubar Feedmill, Ashrat Industrial Area, Israel sity, Palmerston North, Manawat(cid:1)u, New Zealand Hans H. Cheng, USDA, ARS, Avian Disease and Charles M. Bishop, School of Natural Sciences, Bangor Oncology Laboratory, East Lansing, MI, United States University, Bangor, Gwynedd, United Kingdom Helen E. Chmura, Institute of Arctic Biology, University Julio Blas, Department of Conservation Biology, Estación of Fairbanks, Fairbanks, AK, United States Biológica de Doñana, Consejo Superior de Inves- LarryClark,METIS,Ltd.,FortCollins,CO,UnitedStates tigaciones Científicas (CSIC), Seville, Spain Mark A. Cline, Department of Animal and Poultry Sci- WalterGayBottje,DepartmentofPoultryScience,Center ences, Virginia Tech, Blacksburg, VA, United States of Excellence for Poultry Science, Division of Agri- Jamie M. Cornelius, Department of Integrative Biology, culture, University of Arkansas, Fayetteville, AR, Oregon State University, Corvallis, OR, United States United States Dane A. Crossley, Department of Biological Sciences, Kathleen R. Brazeal, School of Biological Sciences, University of North Texas, Denton, TX, United States University of Nebraska-Lincoln, Lincoln, NE, United States Veerle M. Darras, Department of Biology, KU Leuven, Leuven, Belgium Lindsay P. Brown, Department of Chemistry, University of Tennessee, Knoxville, TN, United States Karen D.M. Dean, University of Lethbridge, Lethbridge, AB, Canada ShaneC.Burgess,CollegeofAgriculture&LifeSciences, The University of Arizona, Tucson, AZ, United States Eddy Decuypere, Laboratory of Livestock Physiology, Department of Biosystems, Faculty of Bioscience Warren W. Burggren, Developmental and Integrative Engineering, KU Leuven, Leuven, Belgium Biology, Department of Biological Science, University of North Texas, Denton, TX, United States Mike Denbow, Department of Animal and Poultry Sci- ences, Virginia Tech, Blacksburg, VA, United States Johan Buyse, Laboratory of Livestock Physiology, Department of Biosystems, Faculty of Bioscience En- Pierre Deviche, School of Life Sciences, Arizona State gineering, KU Leuven, Leuven, Belgium University, Tempe, AZ, United States Shawn R. Campagna, Department of Chemistry, Univer- Sami Dridi, Center of Excellence for Poultry Science, sity of Tennessee, Knoxville, TN, United States; UniversityofArkansas,Fayetteville,AR,UnitedStates Biological and Small Molecule Mass Spectrometry Joëlle Dupont, PRC (UMR 6175), INRA, Nouzilly, Core, University of Tennessee, Knoxville, TN, United France;Unité dePhysiologie de laReproduction etdes States Comportements, Institut National de la Recherche Rocco V. Carsia, Department of Cell Biology and Neu- Agronomique, Nouzilly, France roscience, Rowan University School of Osteopathic Vijay Durairaj, Huvepharma Inc., Lincoln, NE, United Medicine, Stratford, NJ, United States States Vincent M. Cassone, Department of Biology, University Edward M. Dzialowski, Department of Biological of Kentucky, Lexington, KY, United States Sciences, University of North Texas, Denton, TX, Natalia Cerón-Romero, Food and Animal Sciences, Ala- United States bama A&M University, Huntsville, AL, United States Nima K. Emami, Center of Excellence for Poultry Shira L. Cheled Shoval, Department of Animal Science, Science, University of Arkansas, Fayetteville, AR, The Robert H. Smith Faculty of Agriculture, Food and United States xxvii xxviii Contributors Nadia Everaert, Precision Livestock and Nutrition Unit, Michael H. Kogut, Southern Plains Agricultural Research TERRA Teaching and Research Centre, Gembloux Center,USDA-ARS,CollegeStation,TX,UnitedStates Agro-Bio Tech, University of Liège, Gembloux, DanielT.Ksepka,BruceMuseum,Greenwich,CT,United Belgium States Graham D. Fairhurst, School of Environment and Sus- Christine Köppl, Cluster of Excellence “Hearing4all”, tainability,UniversityofSaskatchewan,Saskatoon,SK, Carl von Ossietzky University, Oldenburg, Germany; Canada Research Center Neurosensory Science, Carl von Alison Ferver, Center of Excellence for Poultry Science, Ossietzky University, Oldenburg, Germany; Department UniversityofArkansas,Fayetteville,AR,UnitedStates ofNeuroscience,SchoolofMedicineandHealthScience, Carl von Ossietzky University, Oldenburg, Germany AlexanderR.Fisch,DepartmentofChemistry,University of Tennessee, Knoxville, TN, United States Wayne J. Kuenzel, Poultry Science Center, University of Arkansas, Fayetteville, AR, United States Joel Gautron, BOA, INRAE, Université de Tours, Fonction et Régulation des protéines de l’œuf, Vinod Kumar, Department of Zoology, University of Développement de l’œuf, Valorisation, Évolution, Delhi, Delhi, India France H. Lehmann, Animal Welfare Science and Bioethics Elizabeth Gilbert, Department of Animal and Poultry Centre, School of Veterinary Science, Massey Univer- Sciences,VirginiaTech,Blacksburg,VA,UnitedStates sity, Palmerston North, Manawat(cid:1)u, New Zealand David L. Goldstein, Department of Biological Sciences, Scott A. MacDougall-Shackleton, Departments of Psy- Wright State University, Dayton, OH, United States chology and Biology, University of Western, London, ON, Canada Elizabeth S. Greene, Center of Excellence for Poultry Science, University of Arkansas, Fayetteville, AR, Graham R. Martin, School of Biosciences, University of United States Birmingham, Birmingham, United Kingdom Christopher G. Guglielmo, Department of Biology, J.E. Martin, Royal (Dick) School of Veterinary Studies Advanced Facility for Avian Research, Western Uni- and the Roslin Institute, University of Edinburgh, versity, London, ON, Canada Edinburgh, Scotland Thomas P. Hahn, Department of Neurobiology, Physiol- Amanda L. May, Center for Environmental Bio- ogyandBehavior,UniversityofCalifornia,Davis,CA, technology, University of Tennessee, Knoxville, TN, United States United States Orna Halevy, The Hebrew University of Jerusalem, Andrew E. McKechnie, South African Research Chair in Rehovot, Israel; The Ohio State University, Wooster, Conservation Physiology, South African National Bio- OH, United States diversity Institute, Pretoria, Gauteng, South Africa; DSI-NRF Centre of Excellence at the FitzPatrick Maxwell Hincke, Department of Innovation in Medical Institute, Department of Zoology and Entomology, Education; Department of Cellular and Molecular University of Pretoria, Hatfield, Pretoria, Gauteng, Medicine, Faculty of Medicine, University of Ottawa, South Africa Ottawa, ON, Canada D.E.F. McKeegan, Institute of Biodiversity, Animal S.E. Holdsworth, Animal Welfare Science and Bioethics Health and Comparative Medicine, University of Centre, School of Veterinary Science, Massey Univer- sity, Palmerston North, Manawat(cid:1)u, New Zealand Glasgow, Glasgow, Scotland Scott R. McWilliams, Department of Natural Resources Christa F. Honaker, Department of Animal and Poultry Science, University of Rhode Island, Kingston, RI, Sciences,VirginiaTech,Blacksburg,VA,UnitedStates United States Anna Hrabia, Department of Animal Physiology and Henrik Mouritsen, Institut für Biologie und Umweltwis- Endocrinology, University of Agriculture in Krakow, senschaften, Universität Oldenburg, Oldenburg, Ger- Krakow, Poland many; Research Centre for Neurosensory Sciences, Alexander Jurkevich, Advanced Light Microscopy Core, University of Oldenburg, Oldenburg, Germany University of Missouri, Columbia, MO, United States Casey A. Mueller, Department of Biological Sciences, John Kirby, College of the Environment and Life Sci- California State University San Marcos, San Marcos, ences, The University of Rhode Island, Kingston, RI, CA, United States United States Contributors xxix YvesNys,BOA,INRAE,UniversitédeTours,Fonctionet Cynthia A. Smeraski, Science Education and Literacy Régulation des protéines de l’œuf, Développement de Foundational Program, Fort Collins, CO, United States l’œuf, Valorisation, Évolution, France Nurudeen Taofeek, Food and Animal Sciences, Alabama Mary Ann Ottinger, Department of Biology and Bio- A&M University, Huntsville, AL, United States chemistry,UniversityofHouston,Houston,TX,United Hiroshi Tazawa, Developmental and Integrative Biology, States Department of Biological Science, University of North Barbara J. Pierce, Department of Biology, Sacred Heart Texas, Denton, TX, United States University, Fairfield, CT, United States Zehava Uni, Department of Animal Science, The Robert Tom E. Porter, Department of Animal and Avian Sci- H. Smith Faculty of Agriculture, Food and Environ- ences, University of Maryland, College Park, MD, ment, The Hebrew University, Rehovot, Israel United States Sandra G. Velleman, The Hebrew University of Jerusa- Frank L. Powell, Department of Medicine Division of lem, Rehovot, Israel; The Ohio State University, Pulmonary,CriticalCare,SleepMedicine,Universityof Wooster, OH, United States California, San Diego La Jolla, CA, United States Jorge A. Vizcarra, Food and Animal Sciences, Alabama Monika Proszkowiec-Weglarz,United StatesDepartment A&M University, Huntsville, AL, United States of Agriculture, Agricultural Research Service, Animal BrynnH.Voy,DepartmentofAnimalScience,University Biosciences and Biotechnology Laboratory, Beltsville, of Tennessee, Knoxville, TN, United States MD, United States Yajun Wang, Key Laboratory of Bio-resources and Eco- Marilyn Ramenofsky, Department of Neurobiology, environment of Ministry of Education, College of Life Physiology and Behavior, University of California, Sciences, Sichuan University, Chengdu, Sichuan, PR Davis, CA, United States China Narayan C. Rath, USDA/Agricultural Research Service WesleyC.Warren,DepartmentofAnimalSciences,Bond and Department of Poultry Science, University of Life Sciences Center, University of Missouri, Colum- Arkansas, Fayetteville, AR, United States bia, MO, United States Nicole Rideau, Recherches Avicoles, (UR 83), INRA, Heather E. Watts, School of Biological Sciences, Nouzilly, France; Unité de Recherches Avicoles, Washington State University, Pullman, WA, United InstitutNationaldelaRechercheAgronomique,Nouzilly, States France J. Martin Wild, Department of Anatomy and Medical Alejandro B. Rodriguez-Navarro, Departmento de Min- Imaging, Faculty of Medical and Health Sciences, eralogia y Petrologia, Universidad de Granada, Spain University of Auckland, Auckland, New Zealand ColinG.Scanes,CenterofExcellenceforPoultryScience, John C. Wingfield, Department of Neurobiology, Physi- UniversityofArkansas,Fayetteville,AR,UnitedStates; ology and Behavior, University of California, Davis, Department of Biological Sciences, University of CA, United States Wisconsin, Milwaukee, WI, United States Takashi Yoshimura, Laboratory of Animal Integrative ElizabethM.Schultz,DepartmentofBiology,Wittenberg Physiology, Graduate School of Bioagricultural Sci- University, Springfield, OH, United States ences, Institute of Transformative Bio-Molecules, Paul B. Siegel, Department of Animal and Poultry Nagoya University, Nagoya, Aichi Prefecture, Japan Sciences,VirginiaTech,Blacksburg,VA,UnitedStates HuaijunZhou,DepartmentofAnimalScience,University Jean Simon, Recherches Avicoles, (UR 83), INRA, of California, Davis, CA, United States Nouzilly, France Chapter 1 The importance of avian physiology John C. Wingfield DepartmentofNeurobiology,PhysiologyandBehavior,UniversityofCalifornia,Davis,CA,UnitedStates The 20th century saw many revolutionary advances in the well as the complex interplay of physiology, behavior, biological sciences. In 100years, biology progressed from cells, and molecules by which individuals function and cataloginganddescribingspeciestosequencingthehuman interact.Managementandcontroloftheseissuesinrelation genome. Along the way, astounding advances were made to natural resources will also be difficult unless there are in cell and molecular biology, biomechanics, physiology, incentives to foster interdisciplinary research, integrate theoreticalecology,genetics,behavioral neurobiology, etc. education,andmaximizeourpotentialtoaddressproblems Theseandotherareasofthebiologicalsciencescontinueto by bringing to bear all of our knowledge in an effective develop with Aves an important model group for links way. Citing a report from the National Academy of Sci- between environment and gene expression (e.g., Konishi ences,Jasanoffetal.(1997)assertthatinthenextdecades, etal.,1989). Environmental change including degradation, young scientists will be impeded in their advancement population, public health, food and energy production, unless they are trained from an early age to diversify their education, and especially the public’s understanding of expertise and career objectives. scienceandtechnologyaresomeofthemostcriticalissues As an example of the need to integrate ecology, facing science and society. All of these pertain to biology. behavior, and evolutionary biology with mechanisms at Technology and basic research in biological sciences have physiological, genetic, cell, and molecular levels, we can thepotentialtoaddresstheaboveissues,buttheycannotbe consider the functions of a differentiated cell in which understood along traditional disciplinary lines. One enor- complex interactions of many proteins occur. These func- mous hurdle awaits 21st century biological research: how tions can be modified by hormones that coordinate gene to integrate our knowledge of biology at all levels so we activity among various cells and tissues of the organism canpavethewayforadeeperandbroaderunderstandingof leading to the ultimate responses to internal and external how life on this planet works? This will also entail how to environmental signals. A critical question is then how do feed a still burgeoning population and educate future gen- hormones orchestrate transitions in morphology, physi- erationsofundergraduateswhileconservingasmuchofthe ology,andbehaviorofindividualorganismsinrelationtoa naturalworldaspossible?Ifthiswerenotenough,wemust changing, and sometimes capricious environment? We can also deal with potentially catastrophic environmental imagine observing an ecosystem and focusing on certain problems resulting from climate change. populations of individuals within it that have problems Amajorrecurringproblemwiththespectacularsuccess dealing with change in the environment triggered by, for biology has experienced is that individual investigators example, human disturbance. It quickly becomes apparent havebecomesospecializedandfocusedthatinmanycases that some populations, and individuals within them, deal they have, understandably, lost track of other disciplines. with the environmental challenges better than others. It is There is a growing consensus that biologists within disci- reasonable to then ask what aspects of their physiology, plines have difficulty communicating with colleagues in behavior, and morphology are the causes of this failure, or otherbranchesofbiology.Clearly,theproblemswefacein success, and what the cell, molecular, and genetic bases of the next decades will be solved only by taking broad these differences might be. The investigation of interindi- integrative approaches involving expansive, multidisci- vidual variation will be another foundation for under- plinarycollaborations.Inotherwords,molecularbiologyor standing how populations will evolve in response to theoretical ecology in isolation will not resolve problems environmental change. It is relatively simple to construct that involve populations, their component individuals, as hypotheticalscenarioswherebywetraversethespectrumof Sturkie’sAvianPhysiology.https://doi.org/10.1016/B978-0-12-819770-7.00004-9 3 Copyright©2022ElsevierInc.Allrightsreserved. 4 PART | I Undergirdingthemes biological science from ecosystem to molecule in either physiologywillbeacentralfocusforsuchintegration.The direction,butitisnotsointuitivelyobvioushowwedothis examples given below are by no means all inclusive and in practical terms. If such an interdisciplinary and integra- new concepts and research directions are inevitable. tive approach can be achieved, it will be possible to determine how we will deal with global changes already 1.1.1 Physiology and poultry production underwayandultimatelypreparefuturegenerationstocope with ongoing changes. Physiologicalfunctionsofdomesticfowlfromreproduction Avian physiology has a long history of providing and growth to responses to diseases and stress such as modelsforintegrationofdisciplines,anditwillcontinuein weather factors (especially heat) also triggered thorough this role in the future (Konishi et al., 1989). In general, investigations of environmental biology and hormonal there are exceptionally broad data bases for birds on ecol- control of these processes (see Sturkie’s Avian Physiology ogyandevolutionacrosstheglobe.Matchthiswithrapidly revised editions from 1954 to 2020). The domestic fowl developingtoolsatgenomic,transcriptomic,andepigenetic and some other domesticated species (e.g., quail. duck, levels, then avian physiology is uniquely poised midway turkey) provide phenomenally broad data bases of physi- alongthespectrumfromgenestoenvironmentaltoexplore ology that rival those of mammals and fish. These in turn the molecular bases of adaptation and the integration of provide endless opportunities for other studies on wild ecological and evolutionary aspects at the interfaces of species and gene-environment interactionsdthe founda- morphology, physiology, and behavior. In addition, many tions of understanding adaptation to a changing world. wild avian species are abundant and easy to study making them ideal subjects for field observations and experiments 1.1.2 Physiological ecology and birds, marine, in their natural environment, as well as in laboratory freshwater, and terrestrial experimentation. Specific examples how avian physiology hasplayedamajorroleinthedevelopmentandintegration Birds have played a central role in the development of of biological processes at many functional levels are dis- physiologicalecology,especiallybuildingonthevastarray cussed next. of data, techniques and tools made available from agricul- tural research (Phillips et al., 1985; Konishi et al., 1989). 1.1 Specific examples of the Avianphysiologicalecologyisathrivingbranchofbiology importance of avian physiology that has provided a framework for other emerging topics such as environmental endocrinology, the regulation of Avianphysiologyhasitsrootsinover100yearsofresearch seasonal changes in physiology, and individual differences on domesticated species, particularly the domestic fowl and fitness. (e.g., Hau et al., 2010; Taff and Vitousek, (Gallus gallus) as presented in the original version of 2016). Avian migration is a prime example of the integra- Sturkie’s Avian Physiology (Sturkie, 1954). Over the de- tion of physiological systems that has benefitted enor- cades since there have been huge advances in avian phys- mously from avian physiology in general (Newton, 2010; iology focused primarily on the poultry industry and food Dingle, 2014;Ramenofsky and Wingfield, 2007). This has production. Then in the early 21st century, the chicken fueled investigations of one of the most integrative bio- genome was sequenced and annotated (e.g., Burt, 2005; logical processesdthe evolution, ecology, morphology, BurtandPourquie,2003).Thisopenedupanewgeneration physiology, and behavior of movements across the envi- of biological studies at many of the interfaces outlined ronment in general. above, and was followed by sequencing the genome of a Fundamentalandappliedresearchondomesticfowlhas songbird, the zebra finch, Taeniopygia guttata (Warren enricheddevelopmentofavianphysiologyinrelationtothe et al., 2010). As technologies developed to sequence comparative biology of environmental stress (Romero and genomes more quickly and cheaply, the genomes of many Wingfield,2016).Allorganismsmustbeabletocopewith otherspecieshavebeensequencedaspartofambitiousand perturbations of the environment that are largely unpre- excitingprojectstoeventuallysequencethegenomesofall dictable, but require individuals to express facultative the extant species of birds and some extinct ones as well physiologicalandbehavioralresponsestocope.The“stress (e.g., Zhang et al., 2014). If successful, such an unprece- response” is common to most, if not all, organisms from dented data base will allow analyses at so many levels of bacteria and plants to invertebrate as well as vertebrate biological function. However, it should be borne in mind animals. Although the mechanisms involved vary funda- that the computational technologies to analyze such enor- mentally from plants to animals, they are well conserved mousamountsofdataandintegratethemwithotherdiverse within the vertebrates making avian systems excellent and unique data bases on morphology, physiology, models. One aspect of climate change is the increase in behavior, and environment are huge challenges. Avian frequency, duration, and intensity of extreme weather Importanceofavianphysiology Chapter | 1 5 events such as unseasonal storms, hurricanes, and ty- fundamental biology are vast. In the 21st century, with phoons. Understanding how some populations cope with technologiesuntilrecentlyunimagined,thesecontributions these unpredictable events and others do not will have will continue. What could not have been envisaged all profound implications for understanding adaptation and those decades ago is how avian physiology has helped consequences for conservation and agriculture. pioneer,initiateanddevelopnew environmental aspects of New emerging avian models such as the zebra finch, biologicalsciencessuchasmorphologyandbehavioratthe greattit(Parusmajor,Santureetal.,2011),white-throated interfaces with gene-environment interactions and coping sparrow (Zonotrichia albicollis, Tuttle, 2003), and dark- withglobalchangefromurbanization,endocrinedisrupting eyed junco (Junco hyemalis, Ketterson and Atwell, 2016) pollutants,andinvasivespeciestotheburgeoningeffectsof and others, provide field and laboratory contexts for climate change. Thereisnoquestionthatavianphysiology understanding physiological ecology as well as advancing isveryimportantandwillcontinuetobeatthecuttingedge basic research in neurobiology such as the physiology ofsomanybranchesofbiologicalsciences.Itwillcontinue underlying behavioral patterns and avian song control to influence not only basic research but also policy in systems (see Konishi et al., 1989; Pfaff and Joëls, 2017). managing planet Earth for future generations. Theseapproachesusingexperimentsbothinthelaboratory and the field have allowed the beginning of investigations References of coping with climate change, shifts in habitat range, urbanization etc. (e.g., Partecke et al., 2006; Fokidis et al., Bonier,F.,2012.Hormonesinthecity:endocrineecologyofurbanbirds. 2009; Martin et al., 2010; Bonier, 2012; Caro et al., 2013; Horm.Behav.61,763e772. Visser and Gienapp, 2019). Birds have become important Burt,D.,2005.Chickengenome:currentstatusandfutureopportunities. examplesofmodelsforconservationphysiology(Wikelski GenomeRes.15,1692e1698. and Cooke, 2006; Madlinger et al., 2016). For example, Burt,D.,Pourquie,O.,2003.Chickengenome-sciencenuggetstocome avian physiology is providing critical insights into one of soon.Science300,1669. the most important aspects of conservationdthe impact of Carere,C.,Costantini,D.,Sorace,A.,Santucci,D.,Alleva,E.,2010.Bird populationsassentinelsofendocrinedisruptingchemicals.Ann.1st. invasive species resulting from human activity as well as SuperSanita46,81e88.https://doi.org/10.4415/ANN_10_01_10. range expansions and contractions that have important and Caro,S.P.,Schaper,S.V.,Hut,R.A.,Ball,G.F.,Visser,M.E.,2013.The often devastating impacts on the survival of indigenous case of the missing mechanisms: how does temperature influence species from plants to mammals. seasonaltiminginEndotherms?PLoSBiol.11,e1001517. Another spin-off from avian physiology is the use of Dawson, A., 2000. Mechanisms of endocrine disruption with particular birds as sentinels of endocrine disruptiondphysiology, referencetooccurrenceinavianwildlife:areview.Ecotoxicology9, morphology, and behavior (National Research Council, 9e69. 1999; Dawson, 2000; Norris and Carr, 2006; Carere et al., Dingle,H.,2014.Migration:theBiologyofLifeontheMove,seconded. 2010). Although the responses of plants, invertebrates, as OxfordUnivOxford. well as vertebrates, in general, are all important foci to Fokidis, H.B., Orchinik, M., Deviche, P., 2009. Corticosterone and corticosteronebindingglobulininbirds:relationtourbanizationina document toxicology and endocrine disruption, the use of desertcity.Gen.Comp.Endocrinol.160,259e270. birds as models, and their well-known physiology both in Hau,M.,Ricklefs,R.E.,Wikelski,M.,2010.Corticosterone,testosterone the laboratory and field once again provide a useful model andlifehistorystrategiesofbirds.Proc.R.Soc.B277,3203e3212. forunderstandinghoworganismsrespondtodisasterssuch Jasanoff, S., Colwell, R., Dresselhaus, M.S., Goldman, R.D., asleakageoftoxicchemicals,oilspills,etc.Anexampleof Greenwood,M.R.C.,Huang,A.S.,Lester,W.,Levin,S.I.,Linn,M.C., a remaining critical issue of endocrine disruption is how Lubchenco, J., Novacek, M.J., Roosevelt, A.C., Taylor, J.E., organisms will cope with the now ubiquitous and global Wexler,N.,1997.Conversationswiththecommunity:AAASatthe distributionofchemicalmixturesinvaryingconcentrations millennium.Science278,2066e2067. and how we may be able to ameliorate some of the effects Ketterson, E.D., Atwell, J.W., 2016. Snowbird: Integrative Biology and of endocrine disruption by broad ranging “clean up” pro- EvolutionaryDiversityintheJunco.UniverChicagoPress,Chicago. grams to levels of chemicals that organisms can at least Konishi,M.,Emlen,S.,Ricklefs,R.,Wingfield,J.C.,1989.Contributions ofbirdstudiestobiology.Science246,465e472. tolerate? Madlinger, C.L., Cooke, S.J., Crespi, E.J., Funk, J.L., Hultine, K.R., Hunt, K.E., Rohr, J.R., Sinclair, B.J., Suski, C.D., Willis, C.K.R., 1.2 Conclusions Love,O.P.,2016.Successstoriesandemergingthemesinconserva- tion physiology. Cons. Physiol. 4, 206. https://doi.org/10.1093/con- Reading Sturkie’s original avian physiology text (Sturkie, phys/cov/057. 1954), it is astonishing to learn how avian physiology has Martin,L.B.,Hopkins,W.A.,Mydlarz,L.D.,Rohr,J.R.,2010.Theeffects developed, expanded, and progressed in almost 70years. of anthropogenic global changes on immune functions and disease The contributions of avian physiology to agriculture, resistance. Annals NY Acad. Sci. https://doi.org/10.1111/j.1749- biomedicine, veterinary science, and practice, as well as 6632.2010.05454.x.

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