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The neuronal basis of the excitation received by motoneurones during swimming in curarized Xenopus embryos has been investigated further. Extracellular stimulation of axons in the fibre tracts of the spinal cord has been used to evoke unitary excitatory post-synaptic potentials (p.s.p.s) in motoneurones. The p.s.p.s. had a rise time of 3-5 ms and a long falling phase lasting up to 200 ms. These potentials consist of two components: a 'fast' p.s.p. which is insensitive to 50 microM-(+/-)-2-amino-5-phosphonovaleric acid (APV) but is blocked by 2 mM-cis-2,3-piperidine dicarboxylic acid (PDA) and is therefore probably mediated by kainate/quisqualate receptors, and a 'slow' p.s.p. which is blocked by both APV and PDA and is therefore probably mediated by N-methyl-D-aspartate (NMDA) receptors. Paired intracellular recordings from motoneurones and interneurones have revealed a class of spinal cord interneurone which makes descending excitatory amino-acid-dependent synapses onto motoneurones and commissural interneurones. The p.s.p.s evoked by intracellular stimulation of these excitatory interneurones consist of 'fast' and 'slow' components identical in shape and pharmacological properties to those of the extracellularly evoked potentials. One neurone may, therefore, be able to release a transmitter which activates both NMDA and non-NMDA receptors on the same post-synaptic neurone generating fast and slow post-synaptic potentials. The excitatory interneurones play an important role in the generation of the swimming pattern in the curarized Xenopus embryo. Like motoneurones, they fire once per swimming cycle in phase with the ipsilateral motoneurones and receive a background excitation during swimming that is excitatory amino acid mediated. They are therefore part of the swimming rhythm generator. The temporal summation of the extracellularly evoked p.s.p.s shows that these excitatory interneurones are sufficient to generate the excitatory drive received by motoneurones during swimming.
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