Oxidative Stress and Hormesis in Evolutionary Ecology and Physiology: A Marriage Between Mechanistic and Evolutionary Approaches PDF
Preview Oxidative Stress and Hormesis in Evolutionary Ecology and Physiology: A Marriage Between Mechanistic and Evolutionary Approaches
David Costantini Oxidative Stress and Hormesis in Evolutionary Ecology and Physiology A Marriage Between Mechanistic and Evolutionary Approaches Oxidative Stress and Hormesis in Evolutionary Ecology and Physiology David Costantini Oxidative Stress and Hormesis in Evolutionary Ecology and Physiology A Marriage Between Mechanistic and Evolutionary Approaches 123 David Costantini Department of Biology Universityof Antwerp Wilrijk Belgium and InstituteforBiodiversity, AnimalHealth andComparative Medicine Universityof Glasgow Glasgow UK ISBN 978-3-642-54662-4 ISBN 978-3-642-54663-1 (eBook) DOI 10.1007/978-3-642-54663-1 Springer Heidelberg NewYork Dordrecht London LibraryofCongressControlNumber:2014934114 (cid:2)Springer-VerlagBerlinHeidelberg2014 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionor informationstorageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilar methodology now known or hereafter developed. Exempted from this legal reservation are brief excerpts in connection with reviews or scholarly analysis or material supplied specifically for the purposeofbeingenteredandexecutedonacomputersystem,forexclusiveusebythepurchaserofthe work. Duplication of this publication or parts thereof is permitted only under the provisions of theCopyright Law of the Publisher’s location, in its current version, and permission for use must always be obtained from Springer. Permissions for use may be obtained through RightsLink at the CopyrightClearanceCenter.ViolationsareliabletoprosecutionundertherespectiveCopyrightLaw. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publicationdoesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexempt fromtherelevantprotectivelawsandregulationsandthereforefreeforgeneraluse. While the advice and information in this book are believed to be true and accurate at the date of publication,neithertheauthorsnortheeditorsnorthepublishercanacceptanylegalresponsibilityfor anyerrorsoromissionsthatmaybemade.Thepublishermakesnowarranty,expressorimplied,with respecttothematerialcontainedherein. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) To my mother Preface Research programmes on oxidative stress and hormesis are wide ranging. Bio- medicalandtoxicologicalscienceshavetraditionallycentralisedsuchresearch,but in the last years we have seen the merger of laboratory-based biochemical and physiologicalapproachestothestudyofoxidativestressandhormeticmechanisms withethological,ecologicalandevolutionaryprinciples.Althoughresearchinthis area is still in its infancy, many new fields of research are emerging from the marriagebetweenthesemechanisticandfunctionalperspectives.Whatiscritically lackingyetisasynthesisgearedtowardsevolutionaryecologists,ecophysiologists and behavioural ecologists that seeks to consolidate a vast body of literature whilst,atthesametime,remainssensitivetoreaderswhomaynothavethedepth of knowledge often necessary to navigate new fields of scientific enquiry. This book seeks to fill in this gap. Its goal is to synthesise and integrate research on oxidativestressandhormesis,butasseenfromtheperspectiveofanevolutionary ecologistorphysiologist.Workinthisareacanopennewfrontiersforresearchto further our understanding on how and why biological diversity has evolved. This book does not have, however, the presumption to claim that oxidative stress or hormesis are vital, nor does it wish to fully describe biochemical mechanisms. Ratheritwishestoemphasisethatoxidativestressandhormesisaremissingpieces ofacomplexbiologicalpuzzle.Focussingonthesemissingpiecesandintegrating them with other biological features can provide fascinating insights into how organisms work, how they evolve to sustain physiological function under a vast array of environmental conditions and why they work the way they do. The book is organised into ten chapters. Chapter 1 provides general historical information, definitions and background of research on oxidative stress physiol- ogy, hormesis and life history; Chaps. 2–10 analyse how oxidative stress and hormesis have influenced many aspects of life, from the incipient stages of development to the strategies of reproduction and the ways organisms age. While everychapterdealswithaspecifictopic,itshouldnotbeforgottenthattheyareall related to each other. For example, when interpreting the effect of reproductive effortonoxidativedamage,itisalsoimportanttoconsiderthatsucheffectwillbe possibly dependent on other aspects, like early life experiences, predation risk, or food quality and quantity. Moreover, the physiological machinery in turn may influence the way a specific factor impacts on the individual. For example, over evolutionary time, diet has been likely adjusted to the physiological needs of an vii viii Preface individualorspecies,whileatthesametimethephysiologicalstrategieshavebeen adjusted to make use of what is present in the diet itself. We clearly need to integrateallthisinformationinordertofurtherourunderstandingofhowvariation in physiological systems and life history strategies comes out. In this regard, the term adaptation has been used in a general way throughout the book, including both real adaptations and exaptations. For a character to be regarded as an adaptation, it must be a derived character that evolved in response to a specific selective agent. The emergence of an antioxidant mechanism could have been maintained, for example, by natural selection because it confers current selective advantage, but this is not evidence of why it has been evolved. This means that protection against oxidative stress may not be the primary reason for why all antioxidantmechanismshavebeenevolved,butevolutionofacertainantioxidant mechanism could have provided secondarily a selective advantage. Opportunities for studying truly natural processes are diminishing because pristine or near-pristine environments are rapidly shrinking and becoming more difficulttoreach.Thisshouldremindusthatweneedtoknowmoreabouthowthe changing and emerging environments are influencing natural animal populations. The many examples reported throughout the book will actually show how oxi- dative stress and hormesis can represent two important links between environ- mental changes and evolutionary fitness, hence they can represent two relevant tools of investigation for conservation biologists, as well. The book is not exhaustive, hence many relevant studies have not been included. The book has also never taken into consideration discussion of meth- odological approaches to quantify reactive species production, antioxidants or damage levels. Obviously, methods for the quantification of components of the redox machinery have to satisfy specific technical criteria. We have to recognise, however, that each method has limits; hence using a combination of more oxi- dativestatusparametersisessential.Itisalsocrucialthatthebiochemistryofwhat is being measured is taken into consideration when interpreting experimental results.Wehavenottolosesightofthefactthattherearealsospecificrestrictions inherenttomanyecologicalresearchprogrammes,suchastheavailabilityofonly small amounts of blood and the requirement of non-terminal sampling. The biological meaning of many mechanisms or molecules that are currently measured to quantify damage or antioxidant defences is often hard to gauge, because most in-depth information about the physiological functions of many of them come from studies on a limited number of taxa. We also do not know in many cases what differences in oxidative stress mean; this is because we still do not know whether the impact on evolutionary fitness differs among oxidative damage parameters. Consequently, any generalisations and extrapolations about function should be made with considerable caution. Focusing too much on the limited array of model organisms generally used, and on highly conserved path- ways, narrows our understanding of what works under what circumstances. We needtoappreciatethatevolutionhaslikelymodifiedorchangedthebasicdefence toolkitandthatunderstandinghowdifferentorganismshavesolvedthechallenges posed by their environments and lifestyles is interesting and important. Preface ix The complexity and diversity of the natural world is a powerful experimental resourcethatisoftenforgottenbymanyscientificdisciplines.Appreciatingsucha complexity will be instructive and will offer us new opportunities for exploring new frontiers of biological research. David Costantini Acknowledgments Above all I want to thank very much indeed Nadia Macciocchi, Fernando and SerenaCostantini,LiviaandFernandoCarello,whosupportedandencouragedme. I would like to thank very much Gianfranco Brambilla, Claudio Carere, Mauro Carratelli,GiacomoDell’Omo,AlbertoFanfani,NeilMetcalfeandPatMonaghan for valuable advice and support over my career. I also thank very much the following colleagues for providing material or comments that helped me to improve the quality of the book: Doris Abele, Carlos Alonso-Alvarez, John Armstrong, Michaël Beaulieu, Emiliano Bruner, David Bruno, Claudio Carere, Caterina Ferrari, Christopher Guglielmo, Fabrice Helfenstein, Mia Hoogenboom, Valeria Marasco, Scott McWilliams, Anders Pape Møller, Etsuo Niki, Michalis Nikolaidis, Kristin M. O’Brien, Rolf Peterson, Maria Savini, Shona Smith, Gabriele Sorci, Carl Soulsbury and John Speakman. Finally I thank for financial supportofmyresearchtheNaturalEnvironmentResearchCouncil(NERC)ofthe UnitedKingdom,theResearchFoundationofFlanders(FWO),theSwissNational Science Foundation and the Italian association Ornis italica. xi Contents 1 Historical and Contemporary Issues of Oxidative Stress, Hormesis and Life-History Evolution . . . . . . . . . . . . . . . . . . . . . 1 1.1 The Great Oxidation Event: From a Reducing to an Oxidising World. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1.2 Reactive Species, Antioxidants and Oxidative Stress. . . . . . . . 2 1.2.1 On the Nature of Free Radicals and Other Reactive Species . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1.2.2 Antioxidant Mechanisms. . . . . . . . . . . . . . . . . . . . . 5 1.2.3 Oxidative Stress. . . . . . . . . . . . . . . . . . . . . . . . . . . 13 1.2.4 Biochemical Integration and Modularity of Redox Systems. . . . . . . . . . . . . . . . . . . . . . . . . . 16 1.3 Hormesis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 1.3.1 Historical Scenario: Birth, Death and Resurgence of Hormesis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 1.3.2 Types of Hormesis . . . . . . . . . . . . . . . . . . . . . . . . . 21 1.3.3 Quantitative Features of Hormesis and Problems with Its Detection. . . . . . . . . . . . . . . . . . . . . . . . . . 24 1.3.4 Hormesis and Evolutionary Fitness. . . . . . . . . . . . . . 26 1.4 Life-History Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 2 Early-Life Hormesis and Oxidative Experiences Fine-Tune the Adult Phenotype. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 2.1 Early Environment and Phenotypic Development. . . . . . . . . . 39 2.2 Pre-natal Maternal Effects: How Mothers Use Hormones to Shape Their Offspring. . . . . . . . . . . . . . . . . . . . . . . . . . . 41 2.2.1 Effects of Maternal Androgens: Examples from Birds . . . . . . . . . . . . . . . . . . . . . . . 42 2.2.2 Stress Hormones and the Developmental Programming Hypothesis. . . . . . . . . . . . . . . . . . . . . 44 2.3 Epigenetic and Transgenerational Hormetic Effects . . . . . . . . 49 xiii
The list of books you might like
