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Worker Policing in the Honeybee, Epigenetics in Locusts, Ageing In The Honeybee as a model to study Worker policing, Epigenetics, and Ageing Ulrich ERNST Supervisor: Prof. Liliane Schoofs Co-Supervisors: Dr. Peter Verleyen Prof. Tom Wenseleers Members of the Examination Committee: Prof. Johan Billen Dissertation presented Dr. Elke Clynen in partial fulfilment of Prof. Arnold De Loof the requirements for Dr. Christoph Grüter the degree of Doctor in Prof. Roger Huybrechts Science January 2016 © 2016 KU Leuven, Science, Engineering & Technology Uitgegeven in eigen beheer, Ulrich Ernst, Naamsestraat 59, B-3000 Leuven, Belgium Alle rechten voorbehouden. Niets uit deze uitgave mag worden vermenigvuldigd en/of openbaar gemaakt worden door middel van druk, fotokopie, microfilm, elektronisch of op welke andere wijze ook zonder voorafgaandelijke schriftelijke toestemming van de uitgever. All rights reserved. No part of the publication may be reproduced in any form by print, photoprint, microfilm, electronic or any other means without written permission from the publisher. Table of contents PREFACE ............................................................................................................................. 1 1 GENERAL INTRODUCTION ....................................................................................... 3 1.1 HONEYBEES ................................................................................................................................. 3 1.1.1 Life history................................................................................................................................... 3 1.1.2 History: of men and bee .............................................................................................................. 4 1.1.3 Economy ..................................................................................................................................... 5 1.1.4 General scientific interest ............................................................................................................ 6 1.1.5 Development and epigenetics ..................................................................................................... 7 1.2 POLICING ..................................................................................................................................... 8 1.2.1 Conflicts within a superorganism ................................................................................................ 8 1.2.2 Conflict resolution and policing in social insects ....................................................................... 11 1.2.3 Discovery of worker policing ..................................................................................................... 15 1.2.4 Explanations for the occurrence of worker policing ................................................................... 16 1.2.5 Origin and evolution of policing in insects ................................................................................. 22 1.2.6 Maintenance of policing behaviour ............................................................................................ 27 1.2.7 Identification of worker-laid eggs (WLE) ................................................................................... 28 1.2.8 The source of the egg marking signal ....................................................................................... 37 1.2.9 Costs and benefits and best strategies in egg policing ............................................................. 38 1.2.10 Policing outside the social insects ............................................................................................. 41 1.3 PROBLEMS AND RESEARCH QUESTIONS ....................................................................................... 44 1.3.1 Discrimination of worker-laid eggs (WLE) ................................................................................. 44 1.3.2 Specialisation on policing .......................................................................................................... 44 1.3.3 Epigenetics and polyphenism ................................................................................................... 45 1.3.4 Theories of and methods to study ageing ................................................................................. 45 2 PEPTIDES MARK THE DIFFERENCE BETWEEN EGGS OF QUEENS AND WORKERS IN HONEY BEES ................................................................................... 47 ABSTRACT .................................................................................................................................. 48 2.1 INTRODUCTION ........................................................................................................................... 49 2.2 MATERIALS AND METHODS .......................................................................................................... 50 2.2.1 Polar compounds involved in egg recognition ........................................................................... 50 2.2.2 Sample collection procedure for eggs ....................................................................................... 51 2.2.3 Identification of peptides on egg surface ................................................................................... 51 2.2.4 Identification of peptides in queen’s spermatheca .................................................................... 52 2.2.5 Identification of potential modifications ..................................................................................... 52 2.2.6 Identification of signal peptides ................................................................................................. 53 2.2.7 Gene Ontology .......................................................................................................................... 53 2.2.8 Label-free quantitation of peptide abundances ......................................................................... 53 2.2.9 Statistics.................................................................................................................................... 53 2.3 RESULTS .................................................................................................................................... 54 I 2.3.1 Evidence for polar compounds as signals ................................................................................. 54 2.3.2 Identification of proteinaceous compounds on the egg surface ................................................ 54 2.3.3 Identification of spermatheca peptides ...................................................................................... 55 2.3.4 Caste and colony differences in peptide abundancies .............................................................. 55 2.3.5 Differential occurrence of peptides ............................................................................................ 57 2.4 DISCUSSION ............................................................................................................................... 59 2.4.1 New candidates for a queen signal ........................................................................................... 59 2.4.2 Chemical egg protection ........................................................................................................... 60 2.4.3 Quantitative differences between QLE and WLE and colonies ................................................. 61 2.4.4 Perception of peptides .............................................................................................................. 62 2.4.5 Formation of peptides ............................................................................................................... 63 2.4.6 Discussion of the identified compounds .................................................................................... 64 2.4.7 Source of the signal .................................................................................................................. 68 2.4.8 Link to fertility ............................................................................................................................ 69 2.4.9 Origin of peptide fragments and potential causes for quantitative differences .......................... 69 2.4.10 Peptides as artefacts of the isolation method?.......................................................................... 70 2.4.11 Future perspectives .................................................................................................................. 70 2.4.12 Conclusion ................................................................................................................................ 71 2.5 ACKNOWLEDGMENTS .................................................................................................................. 71 3 INDIVIDUAL AND PATRILINE SPECIALISATION IN POLICING BEHAVIOUR IN THE EUROPEAN HONEYBEE, APIS MELLIFERA .................................................. 73 ABSTRACT .................................................................................................................................. 74 3.1. INTRODUCTION ........................................................................................................................... 75 3.1.1 Division of labour and specialisation ......................................................................................... 75 3.1.2 Conflict, conflict resolution and policing .................................................................................... 76 3.1.3 Specialisation
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