Neural basis of a simple behavior: abdominal positioning in crayfish

Crustaceans have been used extensively as models for studying the nervous system. Members of the Order Decapoda, particularly the larger species such as lobsters and crayfish, have large segmented abdomens that are positioned by tonic flexor and extensor muscles. Importantly, the innervation of these tonic muscles is known in some detail. Each abdominal segment in crayfish is innervated bilaterally by three sets of nerves. The anterior pair of nerves in each ganglion controls the swimmeret appendages and sensory supply. The middle pair of nerves innervates the tonic extensor muscles and the regional sensory supply. The superficial branch of the most posterior pair of nerves in each ganglion is exclusively motor and supplies the tonic flexor muscles of that segment. The extension and flexion motor nerves contain six motor neurons, each of which is different in axonal diameter and thus produces impulses of different amplitude. Motor programs controlling each muscle can be characterized by the identifiable motor neurons that are activated. Early work in this field discovered that specific central interneurons control the abdominal positioning motor neurons. These interneurons were first referred to as "command neurons" and later as "command elements." Stimulation of an appropriate command element causes a complex, widespread output involving dozens of motor neurons. The output can be patterned even though the stimulus to the command element is of constant interval. The command elements are identifiable cells. When a stimulus is repeated in a command element, from either the same individual or from different individuals, the output is substantially the same. This outcome depends upon several factors. First, the command elements are not only identifiable, but they make many synapses with other neurons, and the synapses are substantially invariant. There are separate flexion-producing and extension-producing command elements. Abdominal flexion-producing command elements excite other flexion elements and inhibit extensor command elements. The extension producing elements do the opposite. These interactions insure that interneurons of a particular class (flexion- or extension-producing) synaptically recruit perhaps twenty others of similar output, and that command elements promoting the opposing movements are inhibited. This strong reciprocity and the recruitment of similar command elements give a powerful motor program that appears to mimic behavior.