Integration of directional mechanosensory input by crayfish interneurons.

1. Interneurons activated by mechanosensory hairs on the crayfish telson respond selectively to directional displacements of the medium; the directions of maximum sensitivity lie 180 degrees apart in approximately the rostrocaudal plane, corresponding to the directional sensitivities of the two populations of primary afferent neurons. We have examined the basis for this selectivity by intracellular recording in the interneurons, correlating subthreshold potentials with activity evoked in identified afferents by bending single hairs or by producing nearfield displacements of the medium. 2. Interneurons can usually be caused to discharge by a brief train of impulses in single sensory axons. Unitary EPEPs are associated with arriving affterent spikes in the fourth (sensory) root; each primary interneuron receives convergence from several sensory axons, all sensitive to the same direction of movement. Since each afferent axon is drawn from a pair innervating a single sensory structure, this remarkable specificity of connection is unlikely to depend on an anatomical mode of address. 3. Higher order interneurons receive from directionally sensitive lower order interneurons of the same class, as well as from primary afferents of that class. The responses of such cells may show much more decrement during a train of displacement stimuli than do those of lower order cells. Directionality does not appear to be enhanced. 4. During the "null phase" some interneurons appear to be actively inhibited: bending of single hairs 180 degrees away from the effective direction may produce membrane hyperpolarization and slow spontaneous discharges, and shocks to afferent roots produce mixtures of monosynaptic EPSPs and polysynaptic IPSPs.