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Conclusion
How does social organization evolve? At least in part, selection on the superorganismal
trait of stored pollen changes frequencies of alternative alleles (forms) of genes
or gene regulatory networks with broad pleiotropic eff ects including aff ects on reproductive
signaling networks. Changes in signaling networks aff ect changes in development
that aff ect the ovaries of workers, that in turn aff ect sensory physiology and
response systems, and thus the behavior and interactions of thousands of individuals.
And where does the “spirit of the hive” reside? At least in part, it resides in the ovaries
of “a crowd of bees working in a dark hive”.
Acknowledgements
I thank the Wissenscha� skolleg zu Berlin for the opportunity to spend this year as
a Fellow pursuing the development of this model for the evolution of a superorganism.
I also thank Rüdiger Wehner for his support and encouragement, and for his
comments on this manuscript. Research presented here was funded primarily with
grants from the National Institutes of Health, the National Science Foundation, and
the United States Department of Agriculture. � e Alexander von Humboldt-Sti� ung
enabled the long and productive collaboration I have had with Joachim Erber.
532 | Ernst Mayr Lecture 2009