The Spirit of the Hive: The Mechanisms of Social Evolution

241 Pages·2013·5.39 MB·English

by  Robert E. Page Jr.

Th e Spirit
of the Hive

Th e
Spirit
of the
Hive
Th e Mechanisms of Social Evolution
Robert E. Page, Jr.
Harvard University Press
Cambridge, Massachusetts
London, En gland
2013
Copyright © 2013 by the President and Fellows of Harvard College
All rights reserved
Printed in the United States of America
Library of Congress Cataloging- in- Publication Data
Page, Robert E.
Th e spirit of the hive : the mechanisms of social evolution / Robert E. Page, Jr.
pages cm
Includes bibliographical references and index.
ISBN 978-0-674-07302-9 (alkaline paper)
1. Honeybee—Evolution. 2. Honeybee—Behavior 3. Beehives.
4. Pollen. I. Title.
QL568.A6P24 2013
595.79'9—dc23 2012044923
To Harry Laidlaw (1907– 2003)
My good friend, teacher, and mentor

Contents
Foreword by Bert Hölldobler xi
Preface xiii
1 Darwin’s Dilemma and the Spirit of the Hive 1
1.1 Natural History of the Honey Bee 4
1.2 Summary Comments 8
2 What Is the Spirit of the Hive? 10
2.1 Stimulus- Response Basis of Behavior 10
2.2 Th e Logic of Division of Labor 13
2.3 Case Studies 19
2.4 Adaptive Fine Tuning of Division of Labor 24
2.5 From Stone Soup to Mulligan Stew 26
2.6 Summary Comments 45
3 Individual Variation in Behavior 50
3.1 Ge ne tic Variation and Behavior 51
3.2 Polyandry in the Honey Bee 52
3.3 Ge ne tic Recombination in Honey Bees 57
3.4 Ge ne tic Variation Is Necessary for Evolution 60
3.5 Ge ne tic Variation for Worker Behavior 60
3.6 Behavioral Plasticity and Constraints 63
3.7 Ge ne tic and Behavioral Dominance 64
3.8 Behavioral Plasticity and Colony Resilience 68
3.9 Laying-W orker Behavior 72
3.10 Summary Comments 74
viii Contents
4 Th e Evolution of Polyandry 77
4.1 Why Do Queens Mate with So Many Males? 77
4.2 Sex Determination and Polyandry 79
4.3 Pathogens and Parasites 90
4.4 Genotypic Diversity and Division of Labor 95
4.5 A Pluralistic View of the Evolution of Polyandry 106
5 Th e Phenotypic Architecture of Pollen Hoarding 110
5.1 Levels of Biological Or ga ni za tion 110
5.2 Selective Breeding for Pollen Hoarding 112
5.3 Individual Behavior 122
5.4 Sensory- Response Systems 125
5.5 Associative Learning 131
5.6 Nonassociative Learning 134
5.7 Motor Activity 137
5.8 Neurobiochemistry 137
5.9 Anatomy of Worker Ovaries and Vitellogenin 140
5.10 Phenotypic Architecture of Males 141
5.11 Phenotypic Architecture of Africanized Honey Bees 141
5.12 A Pollen- Hoarding Syndrome 142
6 Th e Ge ne tic Architecture of Pollen Hoarding 148
6.1 Background 148
6.2 Mapping Pollen Hoarding 151
6.3 Verifi cation of Quantitative Trait Loci 156
6.4 Identifi cation of Pln3 158
6.5 Pln4 and Mapping the Interactions of
Pollen- Hoarding QTLs 159
6.6 Mapping the Ovary and Juvenile Hormone Regulation
by Vitellogenin 161
6.7 Candidate QTLs 162
6.8 Caveat 166
7 Reproductive Regulation of Division of Labor 169
7.1 Background 169
7.2 Th e Double- Repressor Model 170
7.3 Th e Reproductive- Ground- Plan Hypothesis and
Early Experiments 172
Contents ix
7.4 How Vitellogenin Aff ects Onset of Foraging and
Foraging Behavior 176
7.5 Evidence for the Reproductive- Ground- Plan Hypothesis 179
7.6 Diffi culties with the Vitellogenin Foraging Model 186
7.7 Summary Comments 188
8 Developmental Regulation of Reproduction 191
8.1 Queen and Worker Phenotypes 192
8.2 Nurses and Larvae Share Developmental Programs 196
8.3 Developmental Signatures of Colony- Level Artifi cial
Selection 201
8.4 Summary Comments 203
9 Th e Regulatory Architecture of Pollen Hoarding 208
9.1 Loading Algorithms 208
9.2 Heritability of the Pollen- Hoarding Syndrome 210
9.3 Social Regulation of Pollen Hoarding 213
10 A Crowd of Bees 217
Ac know ledg ments 219
Index 221