Further observations on the effect of de-afferentation on the locomotory activity of amphibian limbs.

The extent to which co-ordinated rhythmical patterns of locomotory movement are to be regarded as expressions of intrinsic activity within the central nervous system, and how far they are dependent upon the patterns of impulses received from peripheral sense organs, remains a fundamental problem in general physiology. In previous communications we have discussed this problem with reference to the ambulatory activity of Amphibia (Gray & Lissmann, 1940). We found that when the limbs and body of a toad {Bufo bufo) are desensitized by severance of their sensory nerves, the effect on the locomotory pattern depends to a very marked degree upon the extent to which the process of de-afFerentation has been applied to the whole of the somatic musculature. In conformity with the results obtained by previous investigators (Hering, 1893; Weiss, 1936) we found that de-afferentation of either one or two limbs did not substantially affect the co-ordinated, diagonal pattern of limb movements. When three limbs were de-afferentated the ambulatory activity of the animal was substantially reduced; nevertheless, the diagonal coordination of the limbs was maintained whenever movement occurred. When four limbs were de-afferentated, ambulation was laboured and restricted to a few steps at a time. On the other hand, when de-afferentation was applied to all the spinal nerves II-XI (inclusive) no trace of ambulatory rhythm or of diagonal limb movements was observed, although non-ambulatory responses were obtained from the limbs by labyrinthine or optical stimulation. From these observations we were unable to derive any positive support for the theory, advocated by Weiss (1936), that the power to maintain rhythmical ambulatory limb movements is an intrinsic property of the central nervous system and fundamentally independent of the peripheral sense organs. We see no reason to amend our views, but further consideration is desirable in view of the subsequent publications by Weiss (1941 a, b) which not only stress the conception of an inherent central nervous control of ambulatory rhythms but quote our experiments in its support.