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Charles Darwin struggled to explain how forty thousand bees working in the dark, seemingly by instinct alone, could organize themselves to construct something as perfect as a honey comb. How do bees accomplish such incredible tasks? Synthesizing the findings of decades of experiments, The Spirit of the Hive presents a comprehensive picture of the genetic and physiological mechanisms underlying the division of labor in honey bee colonies and explains how bees' complex social behavior has evolved over millions of years.
Robert Page, one of the foremost honey bee geneticists in the world, sheds light on how the coordinated activity of hives arises naturally when worker bees respond to stimuli in their environment. The actions they take in turn alter the environment and so change the stimuli for their nestmates. For example, a bee detecting ample stores of pollen in the hive is inhibited from foraging for more, whereas detecting the presence of hungry young larvae will stimulate pollen gathering. Division of labor, Page shows, is an inevitable product of group living, because individual bees vary genetically and physiologically in their sensitivities to stimuli and have different probabilities of encountering and responding to them. A fascinating window into self-organizing regulatory networks of honey bees, The Spirit of the Hive applies genomics, evolution, and behavior to elucidate the details of social structure and advance our understanding of complex adaptive systems in nature.
Foreword [Bert Hölldobler]
Preface
1. Darwin’s Dilemma and the Spirit of the Hive
1.1 Natural History of the Honey Bee
1.2 Summary Comments
2. What Is the Spirit of the Hive?
2.1 Stimulus–Response Basis of Behavior
2.2 The Logic of Division of Labor
2.3 Case Studies
2.4 Adaptive Fine Tuning of Division of Labor
2.5 From Stone Soup to Mulligan Stew
2.6 Summary Comments
3. Individual Variation in Behavior
3.1 Genetic Variation and Behavior
3.2 Polyandry in the Honey Bee
3.3 Genetic Recombination in Honey Bees
3.4 Genetic Variation Is Necessary for Evolution
3.5 Genetic Variation for Worker Behavior
3.6 Behavioral Plasticity and Constraints
3.7 Genetic and Behavioral Dominance
3.8 Behavioral Plasticity and Colony Resilience
3.9 Laying-Worker Behavior
3.10 Summary Comments
4. The Evolution of Polyandry
4.1 Why Do Queens Mate with So Many Males?
4.2 Sex Determination and Polyandry
4.3 Pathogens and Parasites
4.4 Genotypic Diversity and Division of Labor
4.5 A Pluralistic View of the Evolution of Polyandry
5. The Phenotypic Architecture of Pollen Hoarding
5.1 Levels of Biological Organization
5.2 Selective Breeding for Pollen Hoarding
5.3 Individual Behavior
5.4 Sensory-Response Systems
5.5 Associative Learning
5.6 Nonassociative Learning
5.7 Motor Activity
5.8 Neurobiochemistry
5.9 Anatomy of Worker Ovaries and Vitellogenin
5.10 Phenotypic Architecture of Males
5.11 Phenotypic Architecture of Africanized Honey Bees
5.12 A Pollen-Hoarding Syndrome
6. The Genetic Architecture of Pollen Hoarding
6.1 Background
6.2 Mapping Pollen Hoarding
6.3 Verification of Quantitative Trait Loci
6.4 Identification of Pln3
6.5 Pln4 and Mapping the Interactions of Pollen-Hoarding QTLs
6.6 Mapping the Ovary and Juvenile Hormone Regulation by Vitellogenin
6.7 Candidate QTLs
6.8 Caveat
7. Reproductive Regulation of Division of Labor
7.1 Background
7.2 The Double-Repressor Model
7.3 The Reproductive-Ground-Plan Hypothesis and Early Experiments
7.4 How Vitellogenin Affects Onset of Foraging and Foraging Behavior
7.5 Evidence for the Reproductive-Ground-Plan Hypothesis
7.6 Difficulties with the Vitellogenin Foraging Model
7.7 Summary Comments
8. Developmental Regulation of Reproduction
8.1 Queen and Worker Phenotypes
8.2 Nurses and Larvae Share Developmental Programs
8.3 Developmental Signatures of Colony-Level Artificial Selection
8.4 Summary Comments
9. The Regulatory Architecture of Pollen Hoarding
9.1 Loading Algorithms
9.2 Heritability of the Pollen-Hoarding Syndrome
9.3 Social Regulation of Pollen Hoarding
10. A Crowd of Bees
Acknowledgments
Index
Robert E. Page, Jr. is Vice Provost and Dean of the College of Liberal Arts and Sciences and Foundation Professor of Life Sciences at Arizona State University. Bert Holldobler is now Foundation Professor of Biology at Arizona State University; formerly Chair of Behavioral Physiology and Sociology at the Theodor Boveri Institute, University of Wurzburg. He is also the recipient of the U.S. Senior Scientist Prize of the Alexander von Humboldt Foundation and the Gottfried Wilhelm Leibniz Prize of the German government. Until 1990, he was the Alexander Agassiz Professor of Zoology at Harvard University.
