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Reflex Control of Abdominal Flexor Muscles in the Crayfish : II. The Tonic System
1 Department of Biological Sciences, Stanford University, Stanford, California
1. Fibres from the tonic, superficial abdominal flexor muscles in the crayfish receive a complex, highly polyneuronal innervation from among five motor axons and one inhibitor. All efferent nerve fibres show some degree of spontaneous activity.
2. The muscle fibres therefore exhibit a constant flux of membrane potential, and hence of tension, in intact preparations. Depolarization is the result of facilitation and/or summation of junctional potentials of various amplitudes, and in some fibres of superimposed electrogenic responses. Neighbouring fibres tend to show similar innervation patterns, more distant ones dissimilar ones.
3. No useful distinction may be made between fast and slow motor axons. A given axon may produce junctional potentials of very different amplitudes (and some what different rise-times) in neighbouring muscle fibres while another exhibits a precisely reciprocal relationship. The largest axon produces facilitating junctional potentials in all the muscle fibres it innervates, but others may exhibit facilitation in one muscle fibre and antifacilitation in another.
4. Most muscle fibres are innervated by two or three excitatory axons; fibres with single, quadruple or quintuple motor innervation are relatively rare. There is a pronounced tendency for fibres with a rich excitatory innervation to receive the inhibitor as well. The innervation is not shared equally among motor axons: one serves over 90% of the muscle fibres, and two others 20% or less. Statistical analysis of the combinations of motor axons serving muscle fibres reveals that these are apparently random, with all variations from randomness accountable on the grounds of broad regional differences in distribution.
5. The motor axons are selectively activated by specific reflex inputs. Since muscle fibres receive, on the average, only a restricted sample of the available motor supply, it follows that they participate differentially in different reflex actions. Evidence is presented that the firing pattern of motor nerves is appropriate for the temporal properties of their neuromuscular junctions.
6. Reflex inhibition is accomplished by central inhibition of all excitatory motor outflow, accompanied by reciprocal firing in the inhibitor axon. This and the fact that less than half the muscle fibres receive inhibitory innervation demonstrate that, in contrast to the one other crustacean system analysed, reflex inhibition is primarily a central event. Peripheral inhibition in the slow flexor system must serve mainly as a device to achieve repolarization and thus terminate contractions. Such action necessarily depends upon post-synaptic rather than presynaptic mechanisms.
Note:
On leave from the Department of Physiology, University of Tokyo Medical College.
Submitted on January 13, 1965
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