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Journal ArticleDOI

A comparison of peripheral and rubrospinal synaptic input to slow and fast twitch motor units of triceps surae

01 May 1970-The Journal of Physiology (John Wiley & Sons, Ltd)-Vol. 207, Iss: 3, pp 709-732
TL;DR: Post‐synaptic potentials evoked by electrical stimulation of a variety of input systems have been compared in triceps surae motoneurones innervating slow and fast muscle units, the speed of contraction of which was determined.
Abstract: 1. Post-synaptic potentials (PSPs) evoked by electrical stimulation of a variety of input systems have been compared in triceps surae motoneurones innervating slow and fast muscle units, the speed of contraction of which was also determined. 2. Stimulation of high threshold afferents in both flexor and extensor muscle nerves, and of joint afferents, evoked polysynaptic PSPs which were predominantly hyperpolarizing in both fast and slow twitch motor units. 3. Volleys in cutaneous afferents in the sural and saphenous nerves evoked polysynaptic PSPs composed of mixtures of inhibitory and excitatory components. The inhibitory components were predominant in slow twitch motor units, while in fast twitch units there was a trend towards excitatory predominance. 4. Repetitive stimulation of the red nucleus caused predominantly inhibitory PSPs in slow twitch units and mixed or predominantly excitatory PSPs in fast twitch units. There was a correlation in the excitatory/inhibitory balance between PSPs of cutaneous and rubrospinal origin in those motoneurones in which both types of PSPs were studied. 5. The amplitudes of group Ia disynaptic inhibitory PSPs were found to be correlated with motor unit twitch type: IPSPs in slow twitch units were larger than those in fast twitch units. Rubrospinal conditioning volleys were found to facilitate group Ia IPSPs in both fast and slow twitch motor units. 6. The results suggest that there may be several basic patterns of synaptic input organization to motoneurones within a given motor unit pool. In addition to quantitative variation in synaptic distribution, there is evidence that qualitative differences in excitatory to inhibitory balance also exist in the pathways conveying input from cutaneous afferents and rubrospinal systems to triceps surae motoneurones. These qualitative differences are correlated with the motor unit twitch type.
Citations
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Book
05 Jun 1975
TL;DR: Introduction to synaptic circuits, Gordon M.Shepherd and Christof Koch membrane properties and neurotransmitter actions, David A.Brown and Anthony M.Brown.
Abstract: Introduction to synaptic circuits, Gordon M.Shepherd and Christof Koch membrane properties and neurotransmitter actions, David A.McCormick peripheral ganglia, Paul R.Adams and Christof Koch spinal cord - ventral horn, Robert E.Burke olfactory bulb, Gordon M.Shepherd, and Charles A.Greer retina, Peter Sterling cerebellum, Rodolfo R.Llinas and Kerry D.Walton thalamus, S.Murray Sherman and Christof Koch basal ganglia, Charles J.Wilson olfactory cortex, Lewis B.Haberly hippocampus, Thomas H.Brown and Anthony M.Zador neocortex, Rodney J.Douglas and Kevan A.C.Martin Gordon M.Shepherd. Appendix: Dendretic electrotonus and synaptic integration.

3,241 citations

Journal ArticleDOI
TL;DR: Intracellular stimulation of individual motoneurones ensured functional isolation of the muscle units innervated by them in pentobarbitone‐anaesthetized cats.
Abstract: 1. A variety of physiological properties of single motor units have been studied in the gastrocnemius muscle (primarily in the medial head) of pentobarbitone-anaesthetized cats. Intracellular stimulation of individual motoneurones ensured functional isolation of the muscle units innervated by them. 2. A system for muscle unit classification was developed using a combination of two physiological properties. Almost all of the units studied could be classified into one of three major types, including two groups with relatively short twitch contraction times (types FF and FR, which were differentiable from one another on the basis of sensitivity to fatigue) and one group with relatively long contraction times (type S, which were extremely resistant to fatigue and were differentiable from FF and FR units on the basis of the shape of unfused tetani). Post-tetanic potentiation of twitch responses was observed in all three muscle unit types. The distributions of axonal conduction velocities for motoneurones innervating FF and FR muscle units were essentially the same, while conduction velocities for motoneurones innervating type S units were, in general, slower. 3. Histochemical profiles of muscle units representative of each of the physiological classes present in the gastrocnemius pool were determined using a method of glycogen depletion for muscle unit identification. Each of the physiological categories of muscle units exhibited a corresponding unique set of muscle fibre staining reactions, or histochemical profile. Within each physiological type, all of the units examined had the same histochemical profile. The results generally support the hypothesis that the histochemical characteristics of muscle fibres are meaningfully related to the physiological properties of the same fibres. However, certain limitations in the detailed application of the hypothesis were also apparent. 4. Systematic assessment of the histochemical profiles of relatively large numbers of fibres belonging to single muscle units provided strong support for the hypothesis that all of the muscle fibres innervated by a single α-motoneurone are histochemically identical.

1,514 citations

Journal ArticleDOI
TL;DR: Together these mechanisms contribute to the continuous dynamic adjustment of sensorimotor interactions, ensuring that the central program and feedback mechanisms are congruous during locomotion.
Abstract: Locomotion results from intricate dynamic interactions between a central program and feedback mechanisms. The central program relies fundamentally on a genetically determined spinal circuitry (central pattern generator) capable of generating the basic locomotor pattern and on various descending pathways that can trigger, stop, and steer locomotion. The feedback originates from muscles and skin afferents as well as from special senses (vision, audition, vestibular) and dynamically adapts the locomotor pattern to the requirements of the environment. The dynamic interactions are ensured by modulating transmission in locomotor pathways in a state- and phase-dependent manner. For instance, proprioceptive inputs from extensors can, during stance, adjust the timing and amplitude of muscle activities of the limbs to the speed of locomotion but be silenced during the opposite phase of the cycle. Similarly, skin afferents participate predominantly in the correction of limb and foot placement during stance on uneven terrain, but skin stimuli can evoke different types of responses depending on when they occur within the step cycle. Similarly, stimulation of descending pathways may affect the locomotor pattern in only certain phases of the step cycle. Section ii reviews dynamic sensorimotor interactions mainly through spinal pathways. Section iii describes how similar sensory inputs from the spinal or supraspinal levels can modify locomotion through descending pathways. The sensorimotor interactions occur obviously at several levels of the nervous system. Section iv summarizes presynaptic, interneuronal, and motoneuronal mechanisms that are common at these various levels. Together these mechanisms contribute to the continuous dynamic adjustment of sensorimotor interactions, ensuring that the central program and feedback mechanisms are congruous during locomotion.

1,003 citations

Journal ArticleDOI
12 Nov 1971-Science
TL;DR: The correlation among a variety of physiological properties and the histochemical characteristics of muscle fibers belonging to single motor units in a mixed mammalian muscle is directly demonstrated.
Abstract: The correlation among a variety of physiological properties and the histochemical characteristics of muscle fibers belonging to single motor units in a mixed mammalian muscle is directly demonstrated. The population of motor units making up the cat gastrocnemius was classified into three nonoverlapping groups on the basis of a combination of physiological parameters. The muscle fibers belonging to motor units of each physiological type exhibited a distinctive histochemical profile, such that the three basic histochemical "fiber types" exactly matched the three physiologically defined groups. Within each individual motor unit, the muscle fibers were histochemically uniform.

761 citations

OtherDOI
TL;DR: The sections in this article are: Motor Unit Types: Histochemical Profiles and Ultrastructural Correlations, Anatomical Considerations, and Control of Muscular Action: Recruitment and Rate Modulation.
Abstract: The sections in this article are: 1 Motor Unit Types 1.1 Muscle Fiber Types: Histochemical Profiles and Ultrastructural Correlations 1.2 Motor Unit Types: Physiological Profiles in Experimental Animals 1.3 Motor Units in Human Muscle 1.4 Stability of Motor Unit Types 1.5 Developmental Considerations 1.6 Skeletofusimotor Units 2 Anatomical Considerations 2.1 Anatomy of Muscle Units 2.2 Anatomy of Motor Nuclei 2.3 Motoneuron Anatomy in Relation to Unit Type 2.4 Electrophysiological Properties Intrinsic to Motoneurons 2.5 Organization of Synaptic Input 2.6 Control of Motoneuron Excitability: Interactive Factors 3 Control of Muscular Action: Recruitment and Rate Modulation 3.1 Motor Unit Recruitment 3.2 Precision and Stereotypy in Recruitment Process 3.3 Output Modulation by Rate and Pattern of Motoneuron Firing 3.4 Recruitment or Rate and Pattern Modulation? 4 Summary and Concluding Comments

711 citations

References
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Journal ArticleDOI
TL;DR: The present paper is concerned with the central part of the motoneuron and the significance of its size in synaptic transmission and asks whether the cell bodies (and dendrites) connected with large and small motor fibers have different functional properties which can be recognized by their discharge characteristics.
Abstract: SINCE THE BEGINNINGS OF NEUROHISTOLOGY it has been recognized that neurons within the central nervous system vary widely in size, but the functional significance of this basic observation has never emerged from the realm of speculation. The largest cells have surface areas which are at least 100, perhaps 1,000, times greater than those of the smallest cells. Correspondingly, the diameters of axons in the central and peripheral portions of the nervous system range from less than .25 p. to more than 20 c-c. This broad spectrum of physical dimensions invites a search for functional correlates. This is one of a series of studies on the problem of size as it relates to spinal motoneurons. The preceding papers (21, 25) were concerned chiefly with the peripheral part of the motoneuron and the muscle fibers it innervates. They provided experimental evidence that the diameter of a motor nerve fiber is related to the number of muscle fibers it supplies. This finding seemed to make good sense: if a motor fiber innervates many muscle fibers and forms a large motor unit, it must have sufficient axonal substance to give off a large number of terminals. The present paper is concerned with the central part of the motoneuron and the significance of its size in synaptic transmission. It asks whether the cell bodies (and dendrites) connected with large and small motor fibers have different functional properties which can be recognized by their discharge characteristics. In order to investigate this problem one must be able to distinguish the signals of a large motoneuron from those of a small one. This may be done by recording their action potentials from thin filaments of lumbar ventral roots. As Gasser (8) demonstrated, the amplitudes of nerve impulses recorded externally from peripheral nerves are directly related to the diameters of their fibers. If it may be assumed that the diameters of axons are also related to the sizes of their cell bodies, as scattered histo-

2,232 citations

Journal ArticleDOI
TL;DR: The results of experiments designed to resolve the segmental reflex into its functional components are described, finding that group I and group II fibers are the lowest threshold fibers in muscle and cutaneous nerves respectively, and may be excited in isolation by the simple expedient of selecting the appropriate nerves.
Abstract: THE SEGMENTAL REFLEX discharge (7,31,21) must be considered of anatomical rather than functional significance in that it contains, in unnatural combination , those elements which constitute the several distinct ipsilateral reflexes. In the present paper are the results of experiments designed to resolve the segmental reflex into its functional components. The observation that a major division of the segmental reflex into its direct (two-neuron-arc) and indirect (multineuron-arc) components followed segregation of muscle afferent and cutaneous afferent fibers for afferent stimulation (21) provides the point of departure for the experiments to be described. Some of the present observations have been mentioned briefly in a preliminary note (23). A general discussion of these and other results will be found in another paper (25). The afferent fibers of the A group (14) exhibit a range of diameters extending from 20~ to 1.5~ (36). In a dorsal root the whole range of fibers is present, but in the peripheral nerves significant segregation .s are found (36, 8, 29, 14) which permit a degree of selective stimulation of the various components (21). For the purposes of the present discussion the afferent fibers will be classified into groups, each group being marked by a peak in the fiber distribution plots of one or another of the several peripheral nerves. Group I consists of the largest afferent fibers, which are to be found only among the afferent fibers arising from muscle. Approximately these fibers range from 20~ to 12~ in diameter (8, 29), with a distribution peak at 15 to 16~. Group II contains fibers of approximately 12~ to 6~ in diameter, with a mode at 8 to 9p. These fibers form a prominent peak in the fiber distribution plots of cutaneous nerves (8, 30, 14), but they are poorly represented among the muscles afferent fibers (8, 29). Group III consists of fibers gathered about a peak at 3 to 4~ (the delta pile). These last are to be found in both muscle and cutaneous nerves. Another category, to consist of the C fibers, the afferent and reflex function of which is proven (3, 2), should be included as group IV. These fibers have not been studied during the course of the present experiments. Since group I and group II fibers are the lowest threshold fibers in muscle and cutaneous nerves respectively, they may be excited in isolation by the simple expedient of selecting the appropriate nerves …

375 citations

Journal ArticleDOI
TL;DR: Motor units, defined as including a motoneurone (cell body, dendrites and axon) plus the muscle unit innervated, have been examined in the triceps surae motor pool of pentobarbital anaesthetized cats.
Abstract: 1. Motor units, defined as including a motoneurone (cell body, dendrites and axon) plus the muscle unit innervated, have been examined in the triceps surae motor pool of pentobarbital anaesthetized cats.2. The technique of intracellular stimulation and recording which was used permitted measurement of the axonal conduction velocity, post-spike hyperpolarization duration and input resistance of individual motoneurones, and the correlation of these properties with the characteristics of the twitch and tetanus responses of the muscle unit innervated by the cell elicited by direct intracellular stimulation.3. On the basis of muscle unit speed of contraction, motor units were divided into two groups: (a) fast twitch, or F, type with twitch time to peak (TwTp) less than or equal to 30 msec, and (b) slow twitch, or S, type with TwTp of 40 msec or greater. The twitch tensions (TwTen) produced by type F units were significantly larger (median value = 18 g) than the tensions generated by type S units (TwTen median value = 1.6 g). Type F muscle units had much higher tetanus fusion frequencies (median = 85 pulses/sec) than the S type (median 25 pulses/sec), and tended to have smaller tetanus to twitch tension ratios (Tet/Tw) (median = 2.6) than type S units (median = 5.4).4. The gastrocnemius heads contained a mixture of F and S types of muscle units, the proportions found being about 3 to 1 respectively. Units encountered in the soleus muscle were uniformly of type S. The characteristics of gastrocnemius and soleus type S motor units were not identical but appeared to represent quantitative differences in units of the same qualitative type.5. Motoneurones innervating type F muscle units had faster axonal conduction velocities, shorter post-spike hyperpolarizations and lower input resistances than those supplying type S units. However, no combination of motoneurone properties alone was sufficient to separate unambiguously types F and S motor units.

351 citations