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University of Groningen Neglected aspects of hormone mediated maternal effects Kumar, Neeraj

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University of Groningen

Neglected aspects of hormone mediated maternal effects

Kumar, Neeraj

DOI:

10.33612/diss.101325389

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:

2019

Link to publication in University of Groningen/UMCG research database

Citation for published version (APA):

Kumar, N. (2019). Neglected aspects of hormone mediated maternal effects: Studies on early embryonic

modulation of maternal hormonal signals in avian eggs and related methodological aspects. University of

Groningen. https://doi.org/10.33612/diss.101325389

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Neglected aspects of hormone

mediated maternal effects

Studies on early embryonic modulation of maternal hormonal

signals in avian eggs and related methodological aspects

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The research reported in this thesis was carried out in the expertise group ‘Evolutionary Genetics, Development & Behaviour’ at the Groningen Institute for Evolutionary Life Sciences, in collaboration with the Max Planck Institute for Ornithology (Seewiesen, Germany), according to the requirements of the Graduate School of Science, Faculty of Science and Engineering, University of Groningen, The Netherlands.

This work was financially supported by the University of Groningen and the Max Planck Institute for Ornithology.

Cover design: Nele Zickert

Printed by: RidderPrint, The Netherlands ISBN: 978-94-034-2162-9 (printed version) ISBN: 978-94-034-2161-2 (electronic version)

Copyright © N. Kumar 2019, Groningen, The Netherlands.

All rights reserved. No part of this thesis may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, without prior permission of the author.

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Neglected aspects of hormone

mediated maternal effects

Studies on early embryonic modulation of maternal hormonal

signals in avian eggs and related methodological aspects

PhD thesis

to obtain the degree of PhD at the University of Groningen

on the authority of the Rector Magnificus Prof. C. Wijmenga

and in accordance with the decision by the College of Deans. This thesis will be defended in public on Monday 2 December 2019 at 11.00 hours

by

Neeraj Kumar

born on 9 April 1985

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Supervisors

Prof. A.G.G. Groothuis Prof. M. Gahr

Assessment Committee

Prof. R. Bowden Prof. H. Schwabl Prof. G. van Dijk

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CONTENTS

Chapter 1 General introduction and thesis overview 7

Chapter 2

Substantial differences in testosterone measure between LC-MS/MS and radioimmunoassay (RIA) in bird and rat plasma, and in bird eggs, warrant caution for use of RIA kits

21

Chapter 3 Gonadal steroid levels in rock pigeon eggs do not represent

adequately maternal allocation 37 Chapter 4 Avian yolk androgens are metabolized rather than taken up

by the embryo during the first days of incubation 59

Chapter 5

Early embryonic modification of maternal hormones differs systematically among embryos of different laying order: A study in birds

73

Chapter 6

Steroid receptors and their regulation in avian extra-embryonic membranes provide a novel substrate for hormone mediated maternal effects

93

Chapter 7 Summary, synthesis, and future perspectives 107

References 121

Dutch summary 135

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