Generation of scratching. I. Activity of spinal interneurons during scratching

Abstract

1. In decerebrate, curarized cats, stimulation of the cervical spinal cord evoked fictitious scratching (9), i.e., periodical activity of the hindlimb motoneurons with a discharge pattern typical of actual scratching (cycle duration about 250 ms, flexor phase about 200 ms, extensor phase about 50 ms). During fictitious scratching, extra-cellular records were obtained from 182 spinal neurons located in different regions of the gray matter cross section (except for the motor nuclei), from segments L4 and L5. 2. The firing rate of 73% of neurons was rhythmically modulated in relation with the scratch cycle. Most of the modulated neurons fired in bursts and were silent between bursts. They were located mainly in Rexed's (22) layer VII. 3. Burst onsets ("switchings on" of the neurons) were distributed rather evenly throughout the scratch cycle except for a small maximum at the very beginning of the cycle (the cycle was assumed to start with the termination of the extensor phase). Burst terminations ("switchings off") in the overwhelming majority of the neurons were distributed over the last-third part of the cycle. As a result, those neurons which began to fire earlier in the cycle usually had longer bursts, compared to the neurons which began to fire later. Besides, since there were very few switchings off in the first half of the cycle, the number of simultaneously active neurons increased during the first half of the cycle, reached the maximum somewhat later than the middle of the cycle, and considerably decreased by the end of the cycle. 4. With more intensive scratching, the firing rate in the bursts considerably increased in all neurons tested, while the duration of the scratch cycle changed only slightly. 5. A correlation between the burst position in the cycle and the behavior during the latent period of scratching (when stimulation of the cervical spinal cord had already been started but rhythmical oscillations had not yet appeared) was found in many neurons. Most of the neurons which began to fire at the beginning of the scratch cycle and had long bursts were tonically activated during the latent period. On the contrary, most of the neurons which fired in short bursts at the end of the cycle were either inhibited or not affected during this period. 6. A correlation betwen the burst position in the cycle and the frequency pattern was found in many neurons. In most of the neurons which began to fire in the first half of the cycle (except for the very beginning), the discharge rate increased in the course of the burst. In the remaining neurons, the discharge rate changed only slightly during the burst. 7. Hypotheses concerning organization of the spinal mechanism of scratching are discussed.