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Physiological Stress This thesis has been submitted in fulfilment of the requirements for a postgraduate degree (e.g. PhD, MPhil, DClinPsychol) at the University of Edinburgh. Please note the following terms and conditions of use: • This work is protected by copyright and other intellectual property rights, which are retained by the thesis author, unless otherwise stated. • A copy can be downloaded for personal non-commercial research or study, without prior permission or charge. • This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the author. • The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the author. • When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given. Coping with stress: personality, life history and social dominance in swordtail fishes, Xiphophorus sp. Kay Boulton “Everyone knows what stress is and no-one knows what it is.” (Selye, 1973) Submitted for the degree of Doctor of Philosophy The University of Edinburgh 2014 LAY SUMMARY Competition for resources plays an important role in natural selection, creating winners and losers. Winners become socially dominant, obtain resources and so increase their fitness at the expense of losers. Provided they are heritable, phenotypic traits promoting competitive success will be inherited by subsequent generations. Thus, while resource dependent traits (e.g. growth) that rely on competitive outcomes are widely recognised as being under strong selection, this is also likely to be the case for those traits that determine competitive ability and social dominance. In addition, competition is expected to be an important source of stress, for example, harassment of subordinates by dominant individuals. Consequently individual fitness may depend not only on the ability to win resources, but also on the ability to cope with stress. This thesis proposes that social dominance is not just a simple consequence of body size or weaponry, but rather that the interplay between growth, repeatable behavioural characteristics (i.e., personality), and the ability to cope with social and environmental stressors are equally important factors. Thus the dynamic of dominance arises, a model that highlights the expectation of complex relationships between traits causal and consequent to social dominance. Here, empirical studies of Xiphophorus sp. are used to test each element in the model. First the concept of individual personality is explored, asking to what extent it is really stable over long periods of time (equivalent to life-spans). Next, the links between behaviour, physiological stress and contest outcome are considered and, using a repeated measures approach, the hypothesis that individuals differ in “stress coping style” is evaluated. Finally, using a quantitative genetic approach the genetic relationship is estimated between behavioural and life history traits under experimentally manipulated levels of competition. In this way the contribution of genetic and environmental effects to the patterns of trait (co)variation that make up the dynamic of dominance is assessed. I DECLARATION The work described in this thesis has been carried out by myself or as otherwise acknowledged below. It is entirely of my own composition, the main text and bibliography do not exceed 70,000 words and it has not, in whole or part, been submitted for any other degree. Chapters 2, 4 and 5 are based on experiments designed by K Boulton, AJ Wilson and CA Walling. All data were collected by K Boulton from 2010-2013 with assistance where necessary from AJ Grimmer, at the University of Edinburgh and the Tremough campus of Exeter University. AJ Wilson, CA Walling and JD Hadfield assisted with statistical models. K Boulton analysed the data and wrote the manuscripts. Chapter 3 includes data provided by RL Earley, collated and scored by two students, RM Pearce and B Sinderman, University of Alabama, USA. AJ Wilson assisted with statistical models. K Boulton analysed the data and wrote the manuscript. Chapter 4 includes data from samples collected and extracted by K Boulton with RIA performed by Elsa Couto in the laboratory of AVM Canario, CCMar, Faro, Portugal during 2013. K Watt, AVM Canario and RL Earley provided assistance with hormone collection design. Kay Boulton March 2014 II DEDICATION Patrem meum Bowden Dennis (1935-1964) III ABSTRACT Competition for resources plays an important role in natural selection, creating winners and losers. Winners become socially dominant, obtain resources and so increase their fitness at the expense of losers. Provided they are heritable, phenotypic traits promoting competitive success will be inherited by subsequent generations. Thus, while resource dependent traits (e.g. growth) that rely on competitive outcomes are widely recognised as being under strong selection, this is also likely to be the case for those traits that determine competitive ability and social dominance. In addition, competition is expected to be an important source of stress, for example, harassment of subordinates by dominant individuals. Consequently individual fitness may depend not only on the ability to win resources, but also on the ability to cope with stress. This thesis proposes that social dominance is not just a simple consequence of body size or weaponry, but rather that the interplay between growth, repeatable behavioural characteristics (i.e. personality), and the ability to cope with social and environmental stressors are equally important factors. Thus the dynamic of dominance arises, a model that highlights the expectation of complex relationships between traits causal and consequent to social dominance. Here, empirical studies of Xiphophorus sp. are used to test each element in the model. First the concept of individual personality is explored, asking to what extent it is really stable over long periods of time (equivalent to life- spans). Next, the links between behaviour, physiological stress and contest outcome are considered and, using a repeated measures approach, the hypothesis that individuals differ in stress coping style is evaluated. Finally, using a quantitative genetic approach the additive genetic variance-covariance matrix (G) is estimated between behavioural and life history traits under experimentally manipulated levels of competition. In this way the contribution of genetic and environmental effects to the patterns of trait (co)variation that make up the dynamic of dominance is assessed. IV ACKNOWLEDGEMENTS I am indebted to my main supervisors Alastair J Wilson and Craig A Walling for three and a half years of enthusiasm, help, humour and endless, infinite patience. Without them this thesis would not have been possible, and there are not large enough bottles of whisky that will thank them adequately. Additionally I must also thank Per T Smiseth and Sarah Reece for enthusiasm and encouragement, especially during the first year of this work. Special thanks also to the following: Andrew J “Grim” Grimmer for fish expertise, help in the lab and again, endless patience; Jarod D Hadfield, Ian MS White and Neils J Dingemanse for (extra) stats advice and wit along the way; Gil G Rosenthal for sending the little fishy fishes from Mexico; Kathryn Watt, Ryan L Earley, Adelino VM Canario and Elsa Couto who helped, advised and offered yet more patience in the endless quest for cortisol. I am also indebted to the Institute of Evolutionary Biology PhD committee for trusting me with the opportunity to complete this work, to the Engineering and Physical Research Council for PhD funding, the Edinburgh Alliance for Complex Traits Genetics for statistics course funding and the Genetics Society for a Hereditary Fieldwork Grant that funded the visit to Adelino’s lab in Portugal. To my friends, family and loved ones: thank you all for your support and encouragement. Special thanks to the exceptionally talented Class of 2014 (Sarah Mattey (Smattey), Lucy Carter, Elisa Schaum, Emma Hodcroft (honorary member), Hannah Froy, Reuben Nowell, Kevin Donnelly, Emily Moore and Cheryl Gibbons), Classes of 2013 (especially Steph Smith, Jess Hedge and Laura Appleby) and 2015 (especially Elisa Anastasia, Christine Tansey and Jess Flood), parents June and George, daughters Emma and Jennie, sisters JoJo and Denise, friends Anna Murray from Currie, Tango Bob, Jean Budge, Sarah from JFK, the three Jonathans (Moss, Cooke and Strassberg), Maggie and Geoff and last but not least Marmalade, for helping to keep body and soul together. V TABLE OF CONTENTS Lay Summary ............................................................................................................................. I Declaration ............................................................................................................................... II Dedication ............................................................................................................................... III Abstract ................................................................................................................................... IV Acknowledgements .................................................................................................................. V Table of Contents .................................................................................................................... VI Chapter 1 Introduction ...........................................................................................................
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