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The Journal of Neurophysiology Vol. 83 No. 5 May 2000, pp. 2825-2834
Copyright ©2000 by the American Physiological Society
Department of Neurophysiology, Section of Neurophysiology, The Panum Institute, 2200 Copenhagen N, Denmark
Raastad, Morten and
Ole Kiehn.
Spike Coding During Locomotor Network Activity in Ventrally
Located Neurons in the Isolated Spinal Cord From Neonatal Rat. J. Neurophysiol. 83: 2825-2834, 2000. To
characterize spike coding in spinal neurons during rhythmic locomotor
activity, we recorded from individual cells in the lumbar spinal cord
of neonatal rats by using the on-cell patch-clamp technique. Locomotor
activity was induced by N-methyl-D aspartate (NMDA) and 5-hydroxytryptamine (5-HT) and monitored by ventral root
recording. We made an estimator based on the assumption that the number
of spikes arriving during two halves of the locomotor cycle could be a
code used by the neuronal network to distinguish between the halves.
This estimator, termed the spike contrast, was
calculated as the difference between the number of spikes in the most
and least active half of an average cycle. The root activity defined
the individual cycles and the positions of the spikes were calculated
relative to these cycles. By comparing the average spike contrast to
the spike contrast in noncyclic, randomized spike trains we found that
approximately one half the cells (19 of 42) contained a significant
spike contrast, averaging 1.25 ± 0.23 (SE) spikes/cycle. The
distribution of spike contrasts in the total population of cells was
exponential, showing that weak modulation was more typical than strong
modulation. To investigate if this low spike contrast was misleading
because a higher spike contrast averaged out by occurring at different
positions in the individual cycles we compared the spike contrast
obtained from the average cycle to its maximal value in the individual
cycles. The value was larger (3.13 ± 0.25 spikes) than the spike
contrast in the average cycle but not larger than the spike contrast in the individual cycles of a random, noncyclic spike trains (3.21 ± 0.21 spikes). This result suggested that the important distinction between cyclic and noncyclic cells was only the repeated cycle position
of the spike contrast and not its magnitude. Low spike frequencies
(5.2 ± 0.82 spikes/cycle, that were on average 3.5 s long)
and a minimal spike interval of 100-200 ms limited the spike contrast.
The standard deviation (SD) of the spike contrast in the individual
neurons was similar to the average spike contrasts and was probably
stochastic because the SDs of the simulated, noncyclic spike trains
were also similar. In conclusion we find a highly distributed and
variable locomotor related cyclic signal that is represented in the
individual neurons by very few spikes and that becomes significant only
because the spike contrast is repeated at a preferred phase of the
locomotor cycle.
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