Muscle fatigue encompasses a class of acute effects that impair motor performance. The mechanisms that can produce fatigue involve all elements of the motor system, from a failure of the formulation of the descending drive provided by suprasegmental centers to a reduction in the activity of the contractile proteins. We propose four themes that provide a basis for the systematic evaluation of the neural and neuromuscular fatigue mechanisms: 1) task dependency to identify the conditions that activate the various mechanisms; 2) force-fatigability relationship to explore the interaction between the mechanisms that results in a hyperbolic relationship between force and endurance time; 3) muscle wisdom to examine the association among a concurrent decline in force, relaxation rate, and motor neuron discharge that results in an optimization of force; and 4) sense of effort to determine the role of effort in the impairment of performance. On the basis of this perspective with an emphasis on neural mechanisms, we suggest a number of experiments to advance our understanding of the neurobiology of muscle fatigue.

A cinematographic analysis of the unrestrained walking, trotting, galloping, jumping and landing movements of 11 adult cats was undertaken to provide previously unavailable information concerning the demands imposed on the nervous system for the control of low and high speed movements and the demands imposed by such natural movements on muscle performance and proprioceptive response.With due regard for the swing (F and El) and stance (E2 and E3) phases of the step cycle of an individual limb, single frame analysis of the film permitted measurement of instantaneous angles of the lower spine, hip, knee, ankle and metatarsophalangeal joints. Appropriate lever arm measurements were also made on 50 freshly dispatched cats and 25 cadavers such that the Law of Cosines could be used to calculate instantaneous lengths of select hind limb muscles that would apply to the natural movements of adult cats of small (1.5-2.5 Kg), intermediate (2.6-3.5 Kg) and large (3.6-4.5 Kg) size. Muscle displacements were analyzed relative to maximum and minimum i n situ lengths and the lengths associated with quiet standing. Use was also made of a previous electromyographic analysis of hind limb muscles during unrestrained locomotion (Engberg and Lundberg, '69).The sequential relations between the four phases of the step cycle are maintained as forward speed increases from walking (< 2 mph) to high speed galloping (> 16 mph). There are significant differences in the time consumed by each phase, however, with a greater reduction in the ES phase, little reduction in the E2 and E' phases and virtually no reduction in the F phase. When each phase is expressed as a relative percentage of the duration of the total step cycle, the greatest reduction is again in E3 with little change in the E2 phase. In contrast F and E' phases increase in the percent of time they occur in each cycle, with the greatest increase in the F phase. For all speeds, analysis of the phase relations between movements of various sections of the hind limb revealed a remarkable unity of knee and ankle joint movement. The hip joint is largely out of phase with the knee and ankle during E1 and E2, all three joints being in phase in F and E3. The digits are essentially out of phase with the other joints except in the stance phase of the gallop. Rates and extents of muscle displacement during natural movements are greater than might be anticipated when expressed in absolute mm's and mm/sec but not when considered in relation to maximum and minimum in situ length and the length associated with quiet standing (Ls). During stepping a progressive increase in forward speed results in: ( a ) a greater usage of muscles at lengths between Ls and maximum in situ length; ( b ) for knee and ankle extensors, pronounced increase in the lengthening contraction associated with the E2 (yield) phase of the step; and, ( c ) for both flexor and extensor muscles, an increased active phase of lengthening or near isometric contraction immediately prior to periods of active shortening. In contra...

Studies have shown that in the mammalian neuromuscular system stretch reflexes are localized within individual muscles. Neuromuscular compartmentalization, the partitioning of sensory output from muscles, and the partitioning of segmental pathways to motor nuclei have also been demonstrated. This evidence indicates that individual motor nuclei and the muscles they innervate are not homogeneous functional units. An analysis of the functional significance of reflex localization and partitioning suggests that segmental control mechanisms are based on subdivisions of motor nuclei–muscle complexes. A partitioned organization of segmental control mechanisms could utilize (1) the potential functional diversity of muscle fiber types, (2) the variety of mechanical actions of individual muscles arising from their distributed origins and insertions, and (3) diverse architectural features such as intramuscular variations in pinnation and complex in-series and in-parallel arrangements of muscle fibers. The differentiated activity observed in some muscles during natural movements also calls for localized segmental control mechanisms. Partitioning may also play a role in mechanical interactions between contracting motor units and in increasing the stability of neuromuscular systems. The functional advantages of reflex localization and partitioning suggest they are probably common features of segmental systems, whose organization reflects the structure and function of their associated neuromuscular systems.

1. The synaptic connections of 44 single identified muscle spindle Ia afferents and of 21 Golgi tendon organ (Ib) afferents from medial gastrocnemius (MG) were studied in 46 cats by the spike-triggered averaging of synaptic noise in 803 motoneurons of various types. 2. The well-known monosynaptic Ia excitatory connections were confirmed and their characteristics examined in 113 cells. The method was used at greater sensitivity than before and revealed that, in addition to the larger EPSPs of the order of 300 muV, there were many below the previously reported lower 17-muV limit. 3. By studying the Ia disynaptic inhibitory pathway with quick stretch-evoked Ia volleys and by spike-triggered averaging (STA), it was shown that the latter method can reveal disynaptic and possibly trisynaptic excitatory connections. This is believed to depend on having continuous activity in the relevant interneurons. 4. Latencies of individual connections showed broad distributions and arguments are advanced for setting working limits to mono- and disynaptic paths for Ia excitation and inhibition. Monosynaptic EPSP latency from cord entry was 0.4-1.1 ms and disynaptic inhibition was 1.2-2.4 ms. It was recognized that the boundaries are not rigid and monosynaptic Ia EPSPs may have latencies up to 1.5 ms. 5. Rise times of disynaptic PSPs were, on average, significantly longer than monosynaptic, but individual disynaptic responses could have values within the monosynaptic range. 6. A small diphasic wave shortly preceding the monosynaptic EPSPs was interpreted as a presynaptic spike. Its timing was consistent with this and, as such, permitted estimates to be made of central conduction time. 7. An early negative wave (latency less than or equal 1.1 ms) of small emplitude was sometimes detected in antagonist motoneurons when triggering from Ia afferents. It was found tha extracellular fields could be detected due to single Ia afferent excitations and efforts were made to see if the early negative wave could be explained by this. In a few cases there was evidence that a very short-latency IPSP might be occuring. This evidence and its implications are discussed with attention to the new factors which have to be considered in using the spike-triggered averaging method at very high sensitivity. 8. Ib effect were di- or trisynaptic. They were excitatory to 18% of synergists and to 28% of antagonists. They were inhibitory to 41% of synergists and to 19% of antagonists. The Ib IPSPs were larger than the EPSPs.

SUMMARY1. The main purpose of this study was to examine the effects of two subtly different stimulus patterns on the force developed by fast-twitch, fatiguable motor units in a cat hindlimb muscle during control (pre-fatigue) and fatiguing contractions.2. The peak force and the force-time integral responses of nineteen high fatigue (FF) and three intermediate fatigue (FI) motor units of the tibialis posterior muscle in five deeply anaesthetized adult cats were measured at selected times during the course of a 360-s fatigue test.3. The fatigue test involved a pseudo-random alternation of two patterns of stimulation. One pattern (regular) was composed of a train of stimuli with constant interpulse intervals, set at 1-8 x the twitch contraction time of each unit (interval range, 27-51 ms), and delivered for 500 (or 400) ms. For the total (FF + FI) motorunit sample, the mean (±S.D.) stimulation frequency was 26 + 4 Hz (range, 19-37 Hz). The other stimulus pattern (optimized) consisted of three initial stimuli with short (10 ms) interpulse intervals, followed by a constant interpulse-interval train that was adjusted (interval range, 29-62 ms; frequency, 23 + 5 Hz; frequency range, 16-36 Hz) such that the total train had the same number of pulses, and the same average frequency and duration as the regular train.4. The stimulus trains were delivered at 1 s-5 for 360 s, using three-train sequences of each pattern, randomly alternating with one another. The response of the third train in each sequence was selected for the force measurements. The force profile obtained from the fatigue test was subsequently decomposed into two profiles: one attributable to regular and one to optimized stimulation.5. During the initial responses to the fatigue test, the optimized stimulus pattern produced significantly more force than the regular stimulus pattern. For FF units, the mean increase in peak force (141 %) was significantly greater than the increase in the force-time integral (59 %).6. All motor units exhibited an initial potentiation of peak force with the regular stimulation pattern, whereas peak force declined monotonically with the optimized pattern. In contrast, the force-time integral potentiated in the first 30 s for both regular and optimized stimulus patterns. L. BEVAN AND OTHERS 7. Each motor unit maintained an increased force response to optimized stimulation during the fatigue test, with the greatest relative increase occurring about 120 s into the test, well after the potentiation effect had subsided. At 360 s of stimulation, the force enhancement with optimization was still substantial (83 % for peak force and 37 % for the force-time integral), yet force was virtually non-existent in response to regular stimulation.8. These results suggest that subtle changes in the activation pattern are a potential mechanism by which the central nervous system might increase force during fatigue. Furthermore, force optimization and potentiation seem to be motorunit properties that are as fundamental as the more conventionally stu...

SUMMARY1. The main purpose of this study was to quantify the adaptation of spinal motoneurons to sustained and intermittent activation, using an extracellular route of stimulating current application to single test cells, in contrast to an intracellular route, as has been used previously. In addition, associations were tested between firing rate properties of the tested cells and other type (size)-related properties of these cells and their motor units.2. Motoneurons supplying the medial gastrocnemius muscle of the deeply anaesthetized cat were stimulated for 240 s with microelectrodes which passed sustained extracellular current at 1-25 times the threshold for repetitive firing. Many cells were also tested following a rest period with intermittent 1 s current pulses (duration 600 ms) at the same relative stimulus strength. Cell discharge was assessed from the EMG of the motor unit innervated by the test neuron. The motoneurons and their motor units were assigned to four categories (i.e. types FF, FR, S and F; where F = FF + FR) based on conventional criteria. In all, twenty F (16 FF, 4 FR) and fourteen S cells were studied with sustained stimulation. Thirty of these cells (17 F, 13 S) 4. All cells exhibited a delay from the onset of current to the first spike, followed by a brief accelerating discharge that was followed by a slower drop in firing rate. Some cells (21 of 34 with sustained activation; 20 of 32 with intermittent) exhibited doublet discharges (interspike intervals < 10 ms) that were intermingled with the more predominant singlet discharges. Doublets were more common in the S cell type.5. With sustained activation, the mean delay from the onset of current to the first spike was 2-6 + 1 1 s for F cells, and 3-2 + 1-9 s for S cells. The time required to reach peak frequency of singlet discharge following repetitive firing onset was significantly shorter for F than S cells (7-0 + 5 0 vs. 14-3 + 13-6 s) and the peak singlet frequencies also differed significantly (F, 28-0 + 7-7 Hz vs. S, 15-6 + 2-5 Hz). Subsequently, the mean magnitude of firing rate reduction from the peak to 24 s later was significantly greater for F cells than that for S cells (16-2 + 6 Hz vs. 5X8 + 3 Hz). These gradual reductions in firing frequency for both F and S cells during the course of their sustained stimulation were qualitatively similar to the late adaptation observed in previous studies that had employed intracellular stimulation.6. The time course of firing frequency for each unit with sustained activation was fitted with a double-exponential equation: the first time constant (T1) for the initial increase in frequency was relatively short (F, 2X5 + 2-1 s vs. S, 3-7 + 4-1 s). The second time constant (r2) was significantly shorter for F than S cells (130-7 + 98-4 s vs. 750 0 + 402-4 s). It is argued that the r2 values provided a quantitative description of the type of adaptation termed 'late' in previous studies.7. The responses to intermittent stimulation were qualitatively similar to those seen with sustained activat...

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