Biology Reference
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for the present, and till we attempt to locate it, we will term the 'spirit of
the hive.' ” (pp. 38-39; English translation 1903). Here he resorted to a
mystical vitalism to explain how colonies full of individuals working in
the dark organize into a cooperative whole, and he let it for someone
else to identify the “spirit of the hive” and where it resides.
How do insect societies evolve complex social organization? h ere is
a hierarchy of organizational levels from genes to the society, but there
is no centralized control of behavior, and there is no social genome
controlling the society on which natural selection can act. Yet it hap-
pens. Insects display the most complex and fascinating social organiza-
tion known, enough to capture the minds and fantasies of both Darwin
and Maeterlinck. In the following chapters, I will dei ne the genetic,
physiological, and behavioral mechanisms behind the mystical “spirit
of the hive” of Maeterlinck. In Chapter 2, I will show how division of
labor among worker honey bees arises from the simple result that bees,
like all other animals, respond to stimuli in their environments and, as
a consequence, change the stimulus environment. In Chapter 3, I show
how genetic variation within colonies resulting from the polyandrous
mating behavior of queens—queens mate with many males—contributes
to variation in the stimulus-response relationships of workers within a
nest and contributes to their social organization. h us there are conse-
quences of polyandry. In Chapter 4, I propose that polyandry may have
evolved in response to its ef ects on within-colony genetic variation
and social organization and dynamics, although there are many com-
peting hypotheses.
Chapters 5 and 6 focus on results of a selective breeding program that
has continued for more than 20 years, designed to study the ef ects of
selection on a single colony-level trait, the amount of surplus stored pol-
len. h is trait is a consequence of the complex interactions of thousands of
individuals who share the nest. I present the phenotypic architecture—
the correlated changes in many dif erent behavioral, physiological, and
anatomical traits of worker honey bees—of pollen hoarding and then
genetically map the traits to reveal the underlying genetic architecture,
which is an extensive network of interacting genes having ef ects on
many correlated traits.
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