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brain. h e complex web of interacting genes and regulatory pathways
ultimately modulates neurobiochemical systems that modulate
sensory-response systems. However, I will leave the challenge of map-
ping the neurobiology of social behavior to others.
It is clear that it is not sui cient to look at any one level if one wants
to understand the mechanisms and evolution of social behavior. How-
ever, looking at multiple levels at once requires dif erent models than
currently exist. h e question is too complex for stone soup, but we don't
want to make Mulligan stew. We want to be able to understand the nec-
essary and sui cient components, even though they are complex. Many
today are still looking for “a gene for trait X” that is dif erent from “a
gene for trait Y.” Ernst Mayr, one of the architects of the modern evolu-
tionary synthesis, referred to this kind of thinking as “beanbag gene-
tics.” Beanbag thinking is not dif erent from the analogy of trying to
understand what the beach looks like by examining it one grain of sand
at a time. With the growing set of available genomic tools, it is tempting
to treat the honey bee genome as a very large collection of bean bags
and continue to look for the gene or the collection of genes responsible
for some trait. Instead, we need to go up to 10,000 meters and study the
beach. We need to understand the ecology of genes, where the genome
and the physiology of the individual constitute the ecological back-
ground. We should also extend this approach to include the ecology of
the environment, an equal partner during development, and recognize
that phenotypic traits are inl uenced by networks of processes and are
linked together mechanistically. h e balloon metaphor used in Section
5.12 comes to mind.
h is has been the story of more than 30 years of research. I have tried
to link together topics seemingly as disparate as sex determination,
mating behavior, variation in individual behavior, phenotypic and ge-
netic architectures, development, reproductive anatomy and physiology,
and social foraging. Finally, I return to the second part of Maeterlinck's
puzzle. Where does the spirit of the hive reside? At least to some extent
it is in the ovaries of “a crowd of bees working in a dark hive.”
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