6. Social dominance and serotonin receptor genes in crayfish.

Gene expression affects social behavior only through changes in the excitabilities of neural circuits that govern the release of the relevant motor programs. In turn, social behavior affects gene expression only through patterns of sensory stimulation that produce significant activation of relevant portions of the nervous system. In crayfish, social interactions between pairs of animals lead to changes in behavior that mark the formation of a dominance hierarchy. Those changes in behavior result from changes in the excitability of specific neural circuits. In the new subordinate, circuits for offensive behavior become less excitable and those for defensive behavior become more excitable. Serotonin, which is implicated in mechanisms for social dominance in many animals, modulates circuits for escape and avoidance responses in crayfish. The modulatory effects of serotonin on the escape circuits have been found to change with social dominance, becoming excitatory in dominant crayfish and inhibitory in subordinates. These changes in serotonin's effects on escape affect the synaptic response to sensory input of a single cell, the lateral giant (LG) command neuron for escape. Moreover, these changes occur over a 2-week period and for the subordinate are reversible at any time following a reversal of the animal's status. The results have suggested that a persistent change in social status leads to a gradual change in the expression of serotonin receptors to a pattern that is more appropriate for the new status. To test that hypothesis, the expression patterns of crayfish serotonin receptors must be compared in dominant and subordinate animals. Two of potentially five serotonin receptors in crayfish have been cloned, sequenced, and pharmacologically characterized. Measurements of receptor expression in the whole CNS of dominant and subordinate crayfish have produced inconclusive results, probably because each receptor is widespread in the nervous system and is likely to experience opposite expression changes in different areas of the CNS. Both receptors have recently been found in identified neurons that mediate escape responses, and so the next step will be to measure their expression in these identified cells in dominant and subordinate animals.