Search

Crayfish escape from threats by either giant neuron-mediated "reflex" tail flexions that occur with very little delay but do not allow for much sensory guidance of trajectory or by "nongiant" tail flexion responses that allow for sensory guidance but occur much less promptly. Thus, when a stimulus occurs, the nervous system must make a rapid assessment of whether to use the faster reflex system or the slower nongiant one. It does this on the basis of the abruptness of stimulus onset; only stimuli of very abrupt onset trigger giant-mediated responses. We report here that stimuli which excite the lateral giant (LG) command neurons for one form of reflex escape also produce a slightly delayed postexcitatory inhibition (PEI) of the command neurons. As a result, only stimuli that become strong enough to excite the command neurons to firing threshold before the onset of PEI, within a few milliseconds of stimulus onset, can cause giant-mediated responses. This inhibition is directed to distal dendrites of the LG neurons, which allows for some location specificity of PEI within the sensory field of a single hemisegment.

Most neurons have inhibitory synapses both "proximally" near the spike-initiating zone and "distally" on dendrites. Although distal inhibition is thought to be an adaptation for selective inhibition of particular dendritic branches, another important distinction exists between proximal and distal inhibition. Proximal inhibition can attenuate excitatory input absolutely so that no amount of excitation causes firing. Distal inhibition, however, inhibits relatively; any amount of it can be overcome by sufficient excitation. These properties are used as predicted in the circuit-mediating crayfish escape behavior. Many neuronal computations require relative inhibition. This could partly account for the ubiquity of distal inhibition.