University of Groningen. Maternal phenotypic engineering Müller, Wendt

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University of Groningen Maternal phenotypic engineering Müller, Wendt IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from it. Please check the document version below. Document Version Publisher's PDF, also known as Version of record Publication date: 2004 Link to publication in University of Groningen/UMCG research database Citation for published version (APA): Müller, W. (2004). Maternal phenotypic engineering: Adaptation and constraint in prenatal maternal effects Groningen: s.n. Copyright Other than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons). Take-down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons the number of authors shown on this cover page is limited to 10 maximum. Download date: 18-12-2018

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Maternal Phenotypic Engineering Adaptation and Constraint in Prenatal Maternal Effects Introduction 1

The research reported in this thesis was carried out at the Zoological Laboratory, Researchgroup Animal Behavior of the University of Groningen, P.O. Box 14, 9750 AA Haren, The Netherlands. The production of the thesis was funded by the University of Groningen and financially supported by the school for Behavioral and Cognitive Neurosciences (BCN) Cover lay-out: Photos: Printed by: Dick Visser & Wendt Müller Wendt Müller Van Denderen b.v., Groningen 2

RIJKSUNIVERSITEIT GRONINGEN Maternal Phenotypic Engineering Adaptation and Constraint in Prenatal Maternal Effects PROEFSCHRIFT ter verkrijging van het doctoraat in de Wiskunde en Natuurwetenschappen aan de Rijksuniversiteit Groningen op gezag van de Rector Magnificus, Dr. F. Zwarts, in het openbaar te verdedigen op vrijdag 1 oktober 2004 om 14.45 uur door Wendt Müller geboren op 5 februari 1974 te Mönchengladbach Introduction 3

Promotor: Copromotor: Beoordelingscommissie: Prof. Dr. S. Daan Dr. C. Dijkstra Dr. T.G.G. Groothuis Prof. Dr. R.V. Alatalo Prof. Dr. E. Gwinner Prof. Dr. J. Komdeur 4

I was spurred on by an uncertain longing for what is distant and unknown, for whatever excited my fantasy Alexander von Humboldt, 1801 Introduction 5

Addresses of co-authors Rauno V. Alatalo, Heli Siitari Department of Biological and Environmental Sciences, University of Jyväskylä, P.O.Box 35, 40351 Jyväskylä, Finland Viktor Apanius Department of Biology, Wake Forest University, PO Box 7325 Winston Salem NC 27101-7325, USA Jonathan Blount Division of Environmental and Evolutionary Biology, Institute of Biomedical and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK Serge Daan, Cor Dijkstra, Corine Eising, Ton G.G. Groothuis, Alice Kasprzik, Bernd Riedstra Zoological Laboratory, Researchgroup Animal Behavior University of Groningen, P.O. Box 14, 9750 AA Haren, The Netherlands Peter Surai Avian Science Research Centre, Scottish Agricultural College, Auchincruive, Ayr K46 5HW, UK 6

Contents Chapter 1: General introduction 9 Chapter 2: Chapter 3: Chapter 4: Chapter 5: Chapter 6: Chapter 7: Chapter 8: Chapter 9: Within clutch co-variation of egg mass and sex in the black-headed gull (Larus ridibundus) 21 Sex differences in yolk hormones depend on maternal social status in Leghorn chickens (Gallus gallus domesticus) 35 Within-clutch patterns of yolk testosterone vary with the onset of incubation in black-headed gulls 47 Maternal antibody transmission and breeding densities in black-headed gulls 59 Multiple pathways of maternal effects in black-headed gull eggs: Constraint and mutual adjustment 71 Inter-sexual differences in T-cell mediated immunity of black-headed gull chicks (Larus ridibundus) depend on hatching order 87 Consequences of experimentally manipulated brood sex ratios in a sexually dimorphic species 101 An experimental study on the causes of sex-biased mortality in the black-headed gull - the possible role of testosterone 115 Chapter 10: Short- and long-term consequences of prenatal androgen exposure on the immune system of black-headed gulls 129 Chapter 11: Synthesis 147 Dutch summary 163 Postscript 167 References 171 Publications 191 Introduction 7

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