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linked to the stimulus-response relationships best shown by the con-
nections with sucrose sensitivity. h is is not to say that dif erences in
sucrose sensitivity cause the dif erences in foraging behavior—there
could be common, unknown factors causing the variation in both and
generating the correlations—but it does reveal the lack of indepen-
dence of this constellation of phenotypic traits.
h e dominant story that emerges from looking at Figure 5.14 is the
linkage of traits. We selected for one thing, the amount of pollen stored
in the comb, and changed a whole suite of phenotypic traits. h ese
traits are linked causally because we i nd the same associations in wild-
type bees. h e possible phenotypic space, the character associations, is
constrained, not free to vary independently for every trait. Selection is
like squeezing a balloon to change its shape. You can squeeze and
constrict it at one end, but it bulges out at the other, showing that the
entire balloon is connected and interdependent.
What causes these associations? h ere are two plausible explana-
tions. h ese traits could be linked together in a pleiotropic gene net-
work where the ef ects of genes that vary between strains explain dif er-
ences between high- and low-strain bees or between nectar and pollen
specialists, or the suite of traits may be under broad hormonal control,
and the high- and low-strain bees (and nectar and pollen specialists) may
vary for hormonal signals that control the gene network, or both. I will
explore those explanations in Chapters 6, 7, and 8. h e network of asso-
ciated traits reveals the architecture of the “spirit of the hive” through
the networks that af ect sensory-response systems and behavior.
Suggested Reading
Amdam, G. V., Ihle, K. E., and Page, R. E. 2009. Regulation of honeybee worker
(Apis mellifera) life histories by vitellogenin. In Hormones, Brain and
Behavior, 2nd ed., vol. 2, ed. D. W. Pfaf , A. P. Arnold, A. M. Etgen, S. E.
Fahrbach, et al. San Diego: Academic Press, pp. 1003-1025.
Amdam, G. V., and Page, R. E. 2010. h e developmental genetics and physiology
of honeybee societies. Anim. Behav. 79:973-980.
Dreller, C. 1998. Division of labor between scouts and recruits: Genetic inl uence
and mechanisms. Behav. Ecol. Sociobiol. 43:191-196.
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