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Journal of Neuroscience, Vol 9, 2285-2296, Copyright © 1989 by Society for Neuroscience
Spinal cord segments containing key elements of the central pattern generators for three forms of scratch reflex in the turtle
LI Mortin and PS Stein
Biology Department, Washington University, St. Louis, Missouri 63130.
The immobilized, low-spinal turtle produces 3 forms of the fictive scratch
reflex in response to tactile stimulation of specific sites on its body
surface (Robertson et al., 1985). We used complete transections of the
spinal cord at different rostrocaudal levels to reveal the minimum length
of spinal cord sufficient to produce each scratch form. Additional
transections revealed the progressive loss of elements of the motor pattern
and the eventual loss of rhythmogenesis. We have identified, therefore,
spinal cord segments containing key elements of each scratch form's central
pattern generator (CPG). The turtle spinal cord consists of 8 cervical
segments (C1-C8), 10 dorsal segments (D1-D10), 2 sacral segments (S1, S2)
and about 16 caudal segments (Ca1-Ca16; Kusuma et al., 1979). The cell
bodies of motor neurons innervating the hindlimb muscles are located in the
hindlimb enlargement, segments D8-S2 (Ruigrok and Crowe, 1984). The
receptive field for the rostral scratch is innervated by segments D3-D6;
the pocket scratch receptive field is innervated by segments D6-D8; the
caudal scratch receptive field is innervated by segments S2, Ca1, and more
caudal segments (Mortin and Stein, 1985). A rostral scratch motor pattern
could be produced with as few as 5 or 6 segments, i.e., segments D5-D9 or
D3-D8. The anterior 3 segments of the hindlimb enlargement, D8-D10, could
produce a pocket scratch motor pattern. A single segment, either D7 or D8,
is capable of rhythmogenesis in response to stimulation of sites in its
part of the pocket receptive field. A caudal scratch motor pattern could be
produced by D8-End (the hindlimb enlargement and more caudal segments). The
posterior 40-80% of the hindlimb enlargement is not necessary for the
production of a rostral or pocket motor pattern. The anterior segment of
the enlargement is necessary for the production of a normal caudal scratch
motor pattern. Key elements of the CPG for each of the 3 scratch forms
reside in segments D7-D10. The pattern-generating capacity of the anterior
half of the hindlimb enlargement is greater than the posterior half; such
an asymmetric distribution of pattern-generating elements in the
enlargement of the spinal cord has been described for cat scratching
(Deliagina et al., 1983). These results are consistent with the hypothesis
that the CPGs producing different motor patterns for the hindlimb share
neuronal elements (Grillner, 1981; Robertson et al., 1985; Currie and
Stein, 1988, 1989).
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