Jonathan David Blount Of

Ca r o t e n o id s a n d th e Co st s o f R eproduction : S t u d ie s in th e L e s s e r B l a c k -B a c k e d G u l l Jonathan David Blount UNIVERSITY of GLASGOW This thesis is submitted for the degree of Doctor of Philosophy Division of Environmental & Evolutionary Biology Institute of Biomedical & Life Sciences Faculty of Science University of Glasgow April 2002 © J. D. Blount 2002 ProQuest Number: 13818437 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a com plete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 13818437 Published by ProQuest LLC(2018). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States C ode Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106- 1346 GLASGOW 1 UNIVERSITY .LIBRARY: ^ « ' 1111/ UG50 t o n \ CANDIDATE’S DECLARATION I declare that the work recorded in this thesis is entirely my own unless otherwise stated and that it is of my own composition. No part of this work has been submitted for any other degree. Jonathan Blount April 2002 ii A cknowledgements I am very grateful to my supervisors, David Houston and Anders Pape Moller, for their guidance and logistic support over the last three years. I am also grateful to Peter Surai, who became an unofficial supervisor. Peter’s early enthusiasm and encouragement was vital in helping me to turn plans into reality, and he has taught me a lot of useful technical skills along the way. I could not have hoped for more generous and good- humoured supervisors. I was funded by a postgraduate scholarship from the Institute of Biomedical and Life Sciences, University of Glasgow. Kemin Europa NV and Roche Products Ltd. kindly donated the carotenoids used in the supplemental feeding experiments. Several friends and colleagues at the University of Glasgow and elsewhere have helped me by commenting on experiment designs, laboratory analyses, data analyses and manuscripts. In particular, Ruedi Nager has given much time and expertise. I am also very grateful to Trond Amundsen, Iain Barber, Tim Birkhead, Gary Bortolotti, Martin Fowlie, Geoffrey Hill, Stuart Humphries, Malcolm Kennedy, Martyn Kennedy, Hanna Kokko, Kate Lessells, Neil Metcalfe, Pat Monaghan, Francois Mougeot, Paola Ninni and Graeme Ruxton, and not least my office mates - Stuart Bearhop, Sarah Davis, Stuart Fleming, Steve Votier and Jennifer Ward (room 512), Dave Stevens and Nanette Verboven (room 428). Mike Harris, Malcolm Kennedy, Pat Monaghan and Lucas Keller gave helpful feedback on annual progress reports. I also thank Bob Fumess for supplying some yolk samples, ringing training and help with related paperwork, Pat Monaghan and the Cumbria Wildlife Trust for helping me to obtain other licenses for fieldwork, Roger Downie for help with obtaining funds for conference travel, Tom Sawyer and Bemie Zonfrillo for help with obtaining and printing photographs, Clotilde Biard, Andrea Fidgett, June Freel, Pat McLaughlin, Kate Orr and Aileen Adams for help in the lab, and Loma Kennedy for administrative assistance. All my friends, house mates and drinking pals new and old in Glasgow deserve a big thank you for their support and humour, and for making the last three years so enjoyable: “Wee Andrea” Aitken, “Big Gav” Alexander, Jen Andrew, Kate Arnold, Steve Amott, “Big Stu” Bearhop, Maria Bogdanova, Matt Cottam, James Cotton, Fiona Cubitt, Francis Daunt, Sarah Davis, Darren Evans, Andrea Fidgett, Stuart Fleming, Suki Finney, Martin Fowlie, Mike Fowler, “Big Frank”, “Big Dave” Fraser, Helen Gorman, Kate Greaves, Maggie Hall, Geoff Hilton, Stu Humphries, Kat Jones, Ellen Kalmbach, Martyn Kennedy, “Hong Kong” Kenny Leung, Jan Lindstrom, Amanda Maclean, Fiona McPhie, Ian Morgan, Beth Mouat, Ruedi Nager, Alice Pascall, Jane Reid, Ben Ross, Tom Sawyer, Joanna Smith, Vince Smith, Dave Stevens, “Big Svenni”, Michael Trewby, Nanette Verboven, Tom van Pelt, Steve Votier, Jen Ward and Helen Whitaker. Sorry if I’ve forgotten to mention anyone. I have many great memories of my time on Walney Island. For that I’m grateful to the gulls, and also Maria Bogdanova, Brian, Trish and Emma Chadwick, Darren Evans, Stephen Ewing, Nick Littlewood, Ruedi Nager, Alice Pascall, Nanette Verboven and Helen Whitaker. Michael Trewby (‘bushwacker’) is the best friend and fieldworker anyone could hope for. Michael’s superhuman efforts, enthusiasm and support got me through a particularly difficult period during the 1999 field season. Regular supplies of whisky also helped - thank you David! Thanks also to Steve Amott, Andrea Fidgett, Alice Pascall, Michael Trewby and Steve Votier for helping to clear up the cottage and study plots at the end of the season. Finally, I thank my immediate and extended family for their love and support. I dedicate this thesis to Fiona and Isla. Co n t e n t s Summary 1 Chapter 1: General Introduction 4 Physiological trade-offs and life history evolution 5 Maternal effects 6 Limitation of which resource(s)? 8 Carotenoids: biochemistry and biological activity 10 Carotenoids and immune function 11 Carotenoids and egg quality 12 Carotenoids and sperm quality 14 Carotenoids and oxidative stress 15 Could carotenoid supply be limiting in wild birds? 15 Aims and plan of thesis 20 Study species and site 20 References 24 Chapter 2: Carotenoids and egg quality in the lesser black-backed gull 35 Larus fuscus: a supplemental feeding study of maternal effects Introduction 36 Materials and methods 39 Results 46 Discussion 51 References 56 Chapter 3: Patterns of yolk enrichment with dietary carotenoids in gulls: 60 the roles of pigment acquisition and utilization Introduction 61 Materials and methods 64 Results 69 Discussion 75 References 82 v Chapter 4: Carotenoid supply mediates a trade-off between immune 87 defence and chick-rearing capacity in gullsLarus fuscus Introduction 88 Materials and methods 91 Results 98 Discussion 104 References 109 Chapter 5: The influence of carotenoid supply and immune challenge on 114 egg production capacity: an experimental study in gullsLarus fuscus Introduction 115 Materials and methods 118 Results 126 Discussion 137 References 146 Chapter 6: Antioxidants, showy males and sperm quality: a preliminary 152 investigation in lesser black-backed gullsLarus fuscus Introduction 153 Materials and methods 157 Results 160 Discussion 163 References 167 Chapter 7: General discussion 171 Carotenoids and the costs of reproduction 172 Was carotenoid supply limiting for reproduction per se? 172 Interactions between immune function and reproduction: the role of carotenoid 174 supply Some implications for studies of sexual selection 177 Some implications for studies of nutritional biochemistry 178 Future perspectives 179 References 181 vi List of Ta b l e s Table 1.1 Concentrations of carotenoids in the egg yolk of wild birds. 19 Table 2.1 Effects of supplemental feeding on egg phenotype in control and 50 carotenoid-fed lesser black-backed gulls. Table 3.1 Variation in yolk carotenoid profiles in eggs produced by control and 73 carotenoid-supplemented lesser black-backed gulls, resulting from univariate ANOVAs with feeding treatment as a factor. Table 3.2 Spearman rank correlation coefficients between yolk concentrations and 74 percentages of individual carotenoids in eggs produced by control- supplemented lesser black-backed gulls. Table 4.1 Variation in disappearance and hatching rates of foster eggs, and 103 disappearance and fledging rates of chicks hatching from those eggs, in relation to the challenge and diet treatment of the female parent. Table 5.1 Variation in carotenoid and Ig concentrations in chick plasma resulting 133 from rmANCOVAs with age as a within-subjects factor, maternal challenge, maternal diet and position in the laying sequence as between-subjects factors. Table 6.1 Variation in sperm quality indices resulting from an ANCOVA with 162 treatment group as a factor and male coloration index as a covariate. vii L ist o f F ig u r e s Figure 1.1 Chemical structures of carotenoids. 10 Figure 1.2 Lesser black-backed gulls, Larus fuscus, at South Walney Nature 21 Reserve. Figure 2.1 Effects of supplemental feeding on maternal phenotype in control and 48 carotenoid-fed lesser black-backed gulls Figure 2.2 Effects of supplemental feeding on egg phenotype in the control and 49 carotenoid-fed groups Figure 3.1 Concentrations and relative proportions of carotenoids in the yolk of 71 eggs produced by wild lesser black-backed gulls given a carotenoid-free or a carotenoid supplement in the diet for about one month prior to laying Figure 3.2 Mean ± s.e. susceptibility to lipid peroxidation in the yolk of eggs 72 produced by wild lesser black-backed gulls given a carotenoid-free or a carotenoid supplement in the diet for about one month prior to laying Figure 4.1 Changes in maternal phenotype in relation to challenge and diet 100 treatments. Figure 4.2 Chick-rearing capacity in relation to challenge and diet treatments of 102 female parent. Figure 5.1 Egg production capacity in relation to the challenge and diet treatments 128 of female gulls. viii Figure 5.2 Variation in carotenoid and immunoglobulin concentrations in b2-eggs, 130 in relation to maternal challenge and diet treatments. Figure 5.3 Variation in carotenoid and immunoglobulin concentrations in chick 132 plasma over the nestling period in relation to maternal challenge and diet treatments. Figure 5.4 Immune responses of offspring hatching from eggs of differing quality, 134 as influenced by the diet and challenge treatment of the original female parent.

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