Crayfish escape behavior: production of tailflips without giant fiber activity

The giant interneurons of the crayfish nerve cord are well-known mediators of fast tail flexions, "tailflips," that propel animals through the water away from danger. More recent studies have revealed an additional nongiant generator of tailflips. In contrast to giant tailflips, which are stereotyped, nongiant tailflips have variable form. The operating principles and portions of the neural circuitry governing nongiant tailflips were here investigated. Whereas fast flexor motor neurons (FFs) receive excitatory postsynaptic potentials (EPSPs) with large unitary components just prior to giant tailflips, excitation of the FFs during nongiant tailflips is due to summation of many small EPSPs, and these build up for about 60 ms prior to the tailflip; we call the period of excitation prior to FF firing the preflexion phase and the period during which FFs fire, the flexion phase of the tailflip. Even FFs that will not fire during a given tailflip become depolarized during preflexion and flexion periods. Throughout the preflexion and flexion periods there is activity in dorsal nerve cord axons (DCAs) that lie below the giants. Many DCAs are interneurons that excite FFs at short latency. Some DCAs fire uniquely during the preflexion phase, while some fire only during the flexion phase. Which DCAs fire is highly variable, and in some cases firing of particular DCAs can be correlated with particular forms of tailflips. Two identified DCAs, 12 and 13, that fire during the flexion phase were studied. These interneurons originate and receive their synaptic input in the second and third abdominal ganglia, respectively, and project to the last ganglion exciting FFs caudal to their ganglion of origin en route. Their pattern of synaptic input prior to and during nongiant tailflips is indistinguishable from that of FFs. Their input to FFs is weak, but when they fire they tend to promote intersegmental synchrony of FFs in the segments they feed. It appears likely to us that nongiant tailflips are synthesized from a small library of component tailflip movements that can be combined to produce a variety of complete tailflips and that the component movements are produced by a limited group of premotor interneurons, of which 12 and 13 are members.(ABSTRACT TRUNCATED AT 400 WORDS)