Quantification of T- and H-responses before and after a period of endurance training

Summary Tendon (T-) and Hoffmann (H-) responses in the soleus muscle were quantified either separately or in association to compare the mononeurons activated and to study their changes after a period of endurance training. In a first experiment T- and H-responses of the same amplitude were compared: the electrical stimulus (inducing the H-response) and the Achilles tendon tap (inducing the T-response) were associated so that the T-response firstly was concomitant with the H-response, and secondly shifted 10 ms forward or back compared to the H-response. From the study of these combined reflexes we would suggest that the same motoneurons are involved in T- or H-responses of the same amplitude. In a second experiment the maximal H-responses, the T-responses and maximal aerobic power (W _aer,max) were measured on 20 subjects before and after a period of endurance training. For 75% of the subjects the W _aer,max and the reflex parameters (T or H) varied in the same direction: most of them exhibited higher values of both W _aer,max and reflex amplitudes while the others had W _aer,max and reflex values hardly modified or decreased. The different effects of the training period could reflect the heterogeneity of the subject's status and involvement in sport. In most cases the T: Hmax ratios were also influenced, reflecting the fact that T- and H-responses were not identically affected by training. Thus it is suggested that an endurance training programme can influence not only the excitability of the motoneurons but also the response of the muscle receptors to stretch. An interpretation in terms of a change of spindle receptivity and/or a change in their recruitment due to a greater stiffness of the trained muscles is suggested.     

Stretch reflex excitability of the anti‐gravity ankle extensor muscle in elderly humans

Aim: To examine whether the stretch reflex excitability of the soleus muscle changes with age, stretch reflexes at rest (REST) and during weak voluntary contractions (ACT) were elicited in 18 older and 14 younger subjects. Method: The amplitude of the stretch reflex responses and gain, defined as the gradient of the regression line for the relation between stretch reflex responses against the angular velocity of the applied perturbation, were evaluated in each short‐latency (M1) and two long‐latency components (M2 and M3). Results: It was found that in the older group, both the amplitude and gain of the M1 component did not change from the REST to the ACT conditions, whereas in the younger group both variables significantly increased from the REST to ACT conditions. The latency of the M1 component was significantly shorter under the REST condition (older vs. younger: 51.8 ± 7.37 vs. 55.1 ± 8.69 ms), while no group differences were found in those variables under the ACT condition, suggesting that the muscle‐tendon complexes of SOL muscles of the older subjects were less elastic and had less slack, probably due to age‐related histochemical alterations. Further, the Hoffman reflex (H‐reflex), elicited during the REST condition in 10 older and 11 younger subjects showed no significant differences, suggesting that the soleus motoneuron response to the Ia input was comparable between the two subject groups. Conclusion: The histochemical alterations occurring with the ageing process might augment the short‐latency stretch reflex in the SOL muscle without enhancement of motoneuronal excitability, and this effect might be masked when the muscle is voluntarily activated.     

Motor preparation and the achilles tendon reflex: The role of background muscle tension

The effects of instructed tension or relaxation of the soleus muscles on the amplitude of the Achilles tendon reflex were investigated. Reflexes were evoked during the foreperiod of a warned reaction time experiment with a plantar flexion of the right foot serving as the response, and during isometric contractions of the right soleus muscle at different low levels of tension. The tension levels were related to the subject's maximal voluntary contraction force. The results were comparable in the two tasks. When background muscle tension was present in the right leg the reflexes in that leg were depressed, whereas in the contralateral leg reflexes were increased. This difference in amplitude was roughly proportional to the amount of tension and independent of whether the antagonist muscle was also activated during the contraction. It was argued that the differential reflex effect found in the two soleus muscles during motor preparation need not be a reflection of a selective neural inhibition. It could also be explained by a reduced effectiveness of the tendon tap in stretching the muscle, due to stiffening of the muscle during contraction. It was concluded that during motor preparation, only reflex changes recorded in the absence of muscle tension can be considered reflections of subliminal changes in spinal reactivity.     

Monosynaptic and oligosynaptic contributions to human ankle jerk and H-reflex

Studies were undertaken in normal subjects to determine whether it is possible for oligosynaptic reflex pathways to affect motoneuron discharge in the ankle jerk and H-reflex of the soleus. It is argued that if the rising phase of the increase in excitability of the soleus motoneuron pool produced by tendon percussion or by electrical stimulation of the peripheral nerve lasts more than a few milliseconds and if the increase in excitability takes several milliseconds to reach the threshold for motoneuron discharge, these reflexes are unlikely to be exclusively monosynaptic. In relaxed subjects, changes in excitability of the soleus motoneuron pool produced by tendon percussion and by electrical stimulation of the tibial nerve were examined using conditioning stimuli just below threshold and a test H-reflex just above threshold for a reflex response. The increase in excitability due to tendon percussion had an average rise time of 10.8 ms and a total duration of approximately 25 ms. With electrical stimulation the rising phase appeared shorter, but it could not be measured accurately due to afferent refractoriness. In single motor units, the rise times of the composite excitatory postsynaptic potentials (EPSPs) set up by subthreshold tendon percussion and by subthreshold electrical stimulation of the tibial nerve were estimated from changes in the probability of discharge of voluntarily activated single motor units. Rise times were significantly longer with tendon percussion (mean +/- SD, 7.1 +/- 2.3 ms; n = 34) than with electrical stimulation (2.4 +/- 1.4 ms; n = 32). In four experiments in which a number of motor units were studied using identical mechanical and identical electrical stimuli, the poststimulus time histograms (PSTHs) for each stimulus were pooled to provide an estimate of the rise time of the excitability change in the motoneuron pool. The mean rise times of these four samples were 10.5 ms with mechanical stimulation and 4.5 ms with electrical stimulation. The spontaneous variability in latency of reflexly activated single motor units was 0.8-3.1 ms (average SD, 0.34 ms) in the tendon jerk, and 0.6-1.4 ms (average SD, 0.19 ms) in the H-reflex. Comparison of these figures with the measurements of rise time given above suggests that the composite EPSPs are larger than the background synaptic noise. With six motor units, the timing of reflex discharge in the tendon jerk when the subject was relaxed was compared with the timing of the change in probability of discharge due to apparently identical percussion when the units were activated voluntarily.(ABSTRACT TRUNCATED AT 400 WORDS)     

Effects of stretch-shortening cycle training on mechanical properties and fibre type transition in the rat soleus muscle

The effects of exercise training on mechanical properties and fibre type transitions have been investigated in rat soleus muscle. The exercise was a repetition of stretch-shortening cycles. A method of dual controlled releases was applied to obtain tension/extension curves, which characterize the elastic behaviour of the series elastic component (SEC), and the force/velocity relationship characterizing the contractile elements. Other contractile measurements included: contraction time (t _c), half-relaxation time (t _1/2) and twitch/tetanus ratio (P _t/P _o). Additionally, the muscle fibre type composition was determined by a classical histochemical method. A 12-week period of training induced a significantly higher percentage of fast-twitch fibres and a lower percentage of slow-twitch fibres (P<0.01). This fibre adaptation led to a significant (P<0.01) decrease in t _c and an increase in maximum shortening velocity (V _max). An increase in compliance of the SEC was also observed. This elastic adaptation is interpreted in terms of modification of the active components of the SEC. All the histochemical and mechanical data presented in this study show that rat soleus muscles trained by stretch-shortening cycles acquired faster characteristics. Thus the results confirm that a slow-twitch to a fast-twitch fibre transition is obtainable in mature rats.     

Effect of electromyostimulation training on soleus and gastrocnemii H- and T-reflex properties

When muscle is artificially activated, as with electromyostimulation (EMS), action potentials are evoked in both intramuscular nerve branches and cutaneous receptors, therefore activating spinal motoneurons reflexively. Maximal soleus and gastrocnemii H- and T-reflex and the respective mechanical output were thus quantified to examine possible neural adaptations induced at the spinal level by EMS resistance training. Eight subjects completed 16 sessions of isometric EMS (75 Hz) over a 4-week period. Maximal soleus and gastrocnemii M wave (M_max), H reflex (H_max) and T reflex (T_max) were compared between before and after training, together with the corresponding plantar flexor peak twitch torque. No significant changes were observed for electromechanical properties of H_max reflex following EMS. On the other hand, peak twitch torque produced by T_max, but not by equal-amplitude H reflex, significantly increased as a result of training (+21%, P<0.05). These changes were associated with a trend towards a significant increase for normalized gastrocnemii (+21%, P=0.07) but not soleus T_max reflex. It is concluded that, contrary to results previously obtained after voluntary physical training, EMS training of the plantar flexor muscles did not affect alpha motoneuron excitability and/or presynaptic inhibition, as indicated by H-reflex results. On the other hand, in the absence of change in a control group, T_max electromechanical findings indicated that: (1) equal-amplitude H- and T-reflex adapted differently to EMS resistance training; and (2) EMS had an effect on gastrocnemii but not on soleus muscle, perhaps because of the differences in respective motor unit characteristics (e.g., axon diameter).     

Excitability of the soleus H reflex during graded walking in humans

The excitability of the soleus Hoffmann (H) reflex was measured in five healthy male subjects during graded treadmill walking. Uphill and downhill walking at an 8% grade as well as level walking were used to vary the demands for lengthening and shortening contractions of the soleus muscle. These changes were assumed to cause differences in control of the afferent input in the spinal cord and the voluntary output to the soleus muscle. The H reflex was strongly modulated in all three walking conditions, high during the stance phase and low or absent during the swing phase. The shape of the modulations was, however, different. At uphill walking the reflex increased gradually during the whole stance phase and seemed to follow the soleus electromyogram (EMG) pattern closely. In the downhill condition the reflex excitability increased rapidly at heel strike like the soleus EMG and co-contraction of the anterior tibial muscle was observed. At level walking a fast rise in reflex excitability was seen just after heel strike with low or absent soleus EMG. Mean soleus EMG was lower during downhill than during uphill or level walking, but the mean H reflex amplitude was similar in all three conditions. However, when the H reflex was related directly to the EMG activity by linear regression the reflex gain was lower during uphill walking than in the two other conditions. Furthermore, the ratio between H reflex and EMG amplitude was high during the first half of the stance phase at level walking indicating an elevated reflex excitability independent of the voluntary motor output. It is therefore concluded that the modulation of reflexes during walking cannot be interpreted in terms of the idea of automatic gain compensation. The reflexes must be controlled specifically and independently during the different phases of the motor output to meet the mechanical requirements of the movement task. Most explicitly this was seen during downhill walking, where an elevated reflex excitability together with co-contraction at the ankle joint seem to provide increased joint stiffness and security, when the kinetic energy of the body has to be brought under control at heel strike.     

The effects of tenotomy and overload on the postnatal development of muscle fibre histochemistry in the cat triceps surae

Five to seven day-old kittens were subjected to partial tenotomy of the Achilles (triceps surae) tendon. The effects of tenotomy and overload on the development of muscle fibre histochemistry and fibre sizes were investigated when the cats had reached the adult stage. The examined muscles were the uniform soleus and the mixed medial gastrocnemius. Tenotomized muscles of both types had lower weights than their controls. Tenotomized soleus showed a redistribution of succinic dehydrogenase (SDH) activity, together with signs of muscle fibre death. Tenotomized medial gastrocnemius muscles displayed more severe degenerative signs than the soleus, together with signs of fibre death. The fibre death seemed to affect mainly fibres of type IIb. Moreover, a less distinct differentiation in histochemical staining pattern between muscle fibre types was found in these muscles. Overloaded soleus muscles had greater weights than their controls, while no difference could be shown for the overloaded medial gastrocnemius. The overloaded medial gastrocnemius showed a uniform hypertrophy of all fibre types. Also, overloaded soleus showed a uniform hypertrophy. Both types of muscle showed a normal histology as well as normal staining characteristics (SDH and AcATPase). It is concluded that both soleus and medial gastrocnemius are sensitive to loss of muscle tension during development. The basic features of muscle morphology and histochemistry were normal, though, and it is suggested that other factors account for most of the normally occurring development and differentiation.     

Stretch reflex excitability of the anti-gravity ankle extensor muscle in elderly humans

AIM: To examine whether the stretch reflex excitability of the soleus muscle changes with age, stretch reflexes at rest (REST) and during weak voluntary contractions (ACT) were elicited in 18 older and 14 younger subjects. METHOD: The amplitude of the stretch reflex responses and gain, defined as the gradient of the regression line for the relation between stretch reflex responses against the angular velocity of the applied perturbation, were evaluated in each short-latency (M1) and two long-latency components (M2 and M3). RESULTS: It was found that in the older group, both the amplitude and gain of the M1 component did not change from the REST to the ACT conditions, whereas in the younger group both variables significantly increased from the REST to ACT conditions. The latency of the M1 component was significantly shorter under the REST condition (older vs. younger: 51.8 +/- 7.37 vs. 55.1 +/- 8.69 ms), while no group differences were found in those variables under the ACT condition, suggesting that the muscle-tendon complexes of SOL muscles of the older subjects were less elastic and had less slack, probably due to age-related histochemical alterations. Further, the Hoffman reflex (H-reflex), elicited during the REST condition in 10 older and 11 younger subjects showed no significant differences, suggesting that the soleus motoneuron response to the Ia input was comparable between the two subject groups. CONCLUSION: The histochemical alterations occurring with the ageing process might augment the short-latency stretch reflex in the SOL muscle without enhancement of motoneuronal excitability, and this effect might be masked when the muscle is voluntarily activated.     

Different metabolic responses to exercise training programmes in single rat muscle fibres

Summary The aim of this report is to elucidate the effects of exercise training on metabolic properties of different muscle fibre types of the rat hindlimb. Single muscle fibres were dissected from soleus (SOL) or extensor digitorum longus (EDL) muscles of Wistar strain male rats trained on a treadmill for 16 weeks. Each fibre was typed histochemically (SO, slow-twitch oxidative; FOG, fast-twitch oxidative glycolytic; FG, fast-twitch glycolytic). Then glycolytic and oxidative enzymes (CK, LDH, PFK, PK, SDH, and MDH) activities were measured biochemically. Slow,-type fibres (SO) were hypertrophied following endurance training and fast-twitch fibres (FOG and FG) were hypertrophied following sprint training. In EDL muscles the distribution of the slow-type fibres was reduced following the sprint training. The activity of glycolytic enzymes increased significantly in the fast-type fibres (FOG and FG) following sprint training, while oxidative enzymes activities increased in both fast (FOG and FG) and slow (SO) muscle fibres following the endurance training. Neither glycolytic nor oxidative enzymes' activities always increased equally in all types of fibre following exercise training. Consequently, the metabolic profiles in each type of single muscle fibre were affected differently by different intensities of exercise training. These results suggest that the functional (enzymes activity) and structural (muscle fibre hypertrophy) changes of skeletal muscle fibre following exercise training appeared gradually, and would be controlled by different factors.     

Long-loop reflex from arm afferents to remote muscles in normal man

The present study was designed to examine the effects of median nerve stimulation on motoneurones of remote muscles in healthy subjects using H-reflex, averaged EMG and PSTH methods. Stimulation of the median nerve induced facilitation of soleus H-reflex from about 50 ms and it reached a peak at about 100 ms of conditioning-test interval. Afferents that induced the facilitation consisted of at least two types of fibres, the high-threshold cutaneous fibres and the low-threshold fibres. When the effects were examined by the averaged surface EMG and PSTH, no facilitation but rather inhibition or inhibition-facilitation was induced in all tested muscles except for the upper limb muscles on the stimulated side. The inhibition latency was shortest in masseter muscle and longest in leg muscles, while values for the contralateral upper limb muscles were in the middle, indicating that the onset of inhibition was delayed from rostral to caudal muscles. Inputs from the median nerve converged to inhibitory interneurones, which mediate the masseter inhibitory reflex. Our findings suggested that inputs from the median nerve initially ascend to the brain, at least to the brainstem, and then descend to the spinal cord. Therefore, inhibition induced by median nerve stimulation was not considered as an interlimb reflex mediated by a propriospinal pathway, but long-loop reflex, at least via the pons. The discrepancy between the results of reflex and motor units suggests that facilitation of soleus H-reflex following median nerve stimulation was mainly due to reduced presynaptic inhibition.     

Motor unit numbers,motor unit sizes and innervation of single muscle fibres in hyperinnervated adult mouse soleus muscle

The sural and the lateral plantar nerves were implanted simultaneously into the denervated soleus muscle of adult mice. Each of these nerves contained approximately the normal number of soleus motor axons. This procedure therefore allowed a study of how an initial excessive number of motor axons provided by two different, foreign nerves and terminating into the soleus muscle affected the final pattern of muscle innervation. In muscles examined two months or more after the implantation of the foreign nerves all muscle fibres were innervated. The fraction of the muscle innervated by either nerve varied widely from one preparation to another. However, all the motor axons which were implanted into the muscle appeared to make permanent synapses. Moreover, the distribution of motor unit sizes of each foreign nerve relative to the total number of muscle fibres innervated by that nerve was similar to the distribution of motor unit sizes in muscles cross-innervated by that nerve alone, although the absolute motor unit sizes were reduced. Estimated by intracellular recording, 20–30% of the muscle fibres were polyneuronally innervated. A similar fraction of teased muscle fibres stained for acetylcholinesterase had more than one endplate.     

Adaptations in rat skeletal muscle following long-term resistance exercise training

The purpose of this investigation was to determine whether long-term, heavy resistance training would cause adaptations in rat skeletal muscle structure and function. Ten male Wistar rats (3 weeks old) were trained to climb a 40-cm vertical ladder (4 days/week) while carrying progressively heavier loads secured to their tails. After 26 weeks of training the rats were capable of lifting up to 800?g or 140% of their individual body mass for four sets of 12–15 repetitions per session. No difference in body mass was observed between the trained rats and age-matched sedentary control rats. Absolute and relative heart mass were greater in trained rats than control rats. When expressed relative to body mass, the mass of the extensor digitorum longus (EDL) and soleus muscles was greater in trained rats than control rats. No difference in absolute muscle mass or maximum force-producing capacity was evident in either the EDL or soleus muscles after training, although both muscles exhibited an increased resistance to fatigue. Individual fibre hypertrophy was evident in all four skeletal muscles investigated, i.e. EDL, soleus, plantaris and rectus femoris muscles of trained rats, but muscle fibre type proportions within each of the muscles tested remained unchanged. Despite an increased ability of the rats to lift progressively heavier loads, this heavy resistance training model did not induce gross muscle hypertrophy nor did it increase the force-producing capacity of the EDL or soleus muscles.     

Motor Unit Properties in the Soleus Muscle after Its Distal Tendon Transfer to the Plantaris Muscle Tendon in the Rat

The aim of this study was to evaluate how a modification in the mechanical conditions under which a muscle is used could induce changes in the characteristics and the spinal drive of its motor units (MU). The distal tendon of the soleus muscle of Wistar rats was transferred to the distal stump of the plantaris muscle tendon. The EMG activity of the soleus was chronically recorded for 8 weeks, every other day, during a 1-min treadmill walk. After spinal ventral root splitting, individual MU contractile properties were measured in control soleus (102 MUs) or in transposed soleus muscles after 4 weeks (41 MUs) or 8 weeks (28 MUs). Muscle/body weight ratio did not vary after transposition, nor did MU tetanic forces. A decrease in MU twitch contraction times and in their half relaxation times was observed at weeks 4 and 8. MU tension-frequency curves varied significantly after tendon transfer, becoming closer to the curves of the fast MUs of the control group. During locomotion, we observed no change in the amplitude of rectified-filtered electromyographic activity, but a significant decrease in mean burst duration and an increase in the median frequency of the power density spectrum. Tendon transposition of the soleus muscle brought about adaptations in MU contractile properties and soleus spinal control.     

Cross-innervation of fast and slow-twitch muscles by motor axons of the sural nerve in the mouse

The extensor digitorum longus (EDL) or soleus muscles of adult mice were cross-innervated by the sural nerve (SN) and deprived of their original innervation. The number and sizes of motor units and the location of endplates in these muscles were studied 1.5 to 16 months later. In the EDL muscle, the SN cross-innervated the original endplates. Very few ectopic endplates were seen, even when the nerve was implanted well outside of the original endplate area. Only 3% of the fibres were polyneuronally innervated. In the soleus muscle, however, the SN formed large numbers of ectopic endplates whether the nerve was implanted in the original endplate zone or outside of it. In addition, 20% of the muscle fibres were polyneuronally innervated. The SN cross-innervated both EDL and soleus muscles completely. There was no preference for a particular group of the SN motoneurones since all the cross-innervated muscles were innervated by all SN motor axons and the motor unit sizes of the SN were similar in the cross-innervated EDL and soleus muscles. It is concluded that intrinsic properties of a muscle determine the ability to form ectopic synapses. The distribution of the motor unit sizes is determined by the particular pool of motoneurones which innervates the muscle.     

Different modulation pattern of spinal stretch reflex excitability in highly trained endurance runners

This study was undertaken to elucidate the impact of long-term physical training on the modulation of stretch reflex excitability. To this end, electromyographic activities of the soleus muscle in response to quick toe-up rotation were compared between highly trained endurance runners (n?=?8) and non-trained control subjects (n?=?9). We specifically focused on the stretch reflex modulation under different voluntary activation levels, from rest to pre-activated conditions (5, 10, 20, and 30% of the maximal). While the two groups showed similar modulation patterns of the stretch reflex responses, the extent of reflex modulation in accordance with the muscle pre-activation level was larger in the trained group. The present results therefore suggest a different modulation pattern of the stretch reflex responses with changing activation level between individuals with different physical background, and the enhancement of the responses in the trained individuals may particularly be advantageous in exerting high level muscle contraction.     

Changes in the excitability of soleus muscle short latency stretch reflexes during human hopping after 4 weeks of hopping training

Changes in the excitability of the human triceps surae muscle short latency stretch reflexes were investigated in six male subjects before and after 4 weeks of progressive two-legged hopping training. During the measurements the subjects performed 2-Hz hopping with: preferred contact time (PCT) and short contact time. The following reflex parameters were examined before and after the training period: the soleus muscle (SOL) Hoffmann-reflex (H-reflex) at rest and during hopping, the short latency electromyogram (EMG) components of the movement induced stretch reflex (MSR) in SOL and medial gastrocnemius muscle (MG), and the EMG amplitude of the SOL and MG tendon reflexes (T-reflexes) elicited at rest. The main results can be summarized as follows: the SOL T-reflex had increased by about 28% (P < 0.05) after training while the MG T-reflex was unchanged; the SOL MSR (always evident) and the MG MSR (when observable) did not change in amplitude with training, and before training the SOL H-reflex in both hopping situations was significantly depressed to about 40% of the reference value at standing rest (P < 0.05). After training the H-reflex during PCT hopping was no longer depressed. As the value of the measured mechanical parameters (the total work rate, joint angular velocity and the ankle joint work rate) was unchanged after training in both hopping situations, the reflex changes observed could not be ascribed to changes in the movement pattern. To explain the observed changes, hypotheses of changes in the excitability of the stretch reflex caused by the training were taken into consideration and discussed.     

Human proprioceptive adaptations during states of height-induced fear and anxiety

Clinical and experimental research has demonstrated that the emotional experience of fear and anxiety impairs postural stability in humans. The current study investigated whether changes in fear and anxiety can also modulate spinal stretch reflexes and the gain of afferent inputs to the primary somatosensory cortex. To do so, two separate experiments were performed on two separate groups of participants while they stood under conditions of low and high postural threat. In experiment 1, the proprioceptive system was probed using phasic mechanical stimulation of the Achilles tendon while simultaneously recording the ensuing tendon reflexes in the soleus muscle and cortical-evoked potentials over the somatosensory cortex during low and high threat conditions. In experiment 2, phasic electrical stimulation of the tibial nerve was used to examine the effect of postural threat on somatosensory evoked potentials. Results from experiment 1 demonstrated that soleus tendon reflex excitability was facilitated during states of height-induced fear and anxiety while the magnitude of the tendon-tap-evoked cortical potential was not significantly different between threat conditions. Results from experiment 2 demonstrated that the amplitudes of somatosensory-evoked potentials were also unchanged between threat conditions. The results support the hypothesis that muscle spindle sensitivity in the triceps surae muscles may be facilitated when humans stand under conditions of elevated postural threat, although the presumed increase in spindle sensitivity does not result in higher afferent feedback gain at the level of the somatosensory cortex.     

Cross-innervated mammalian skeletal muscle: histochemical, physiological and biochemical observations

1. Cross-innervation of the slow soleus and fast flexor hallucis longus or flexor digitorum longus muscles has been performed in new-born kittens and rabbits and in adult cats.

2. The effects on the histochemical and structural properties of the muscle have been studied and compared with the changes in the contractile properties.

3. Cross-innervation has produced a dramatic change in histochemical pattern in the fast muscles, with the development of areas of muscle fibres indistinguishable from normal soleus muscle. The converse change from the histochemical pattern of slow soleus to that of fast muscle has also occurred, but has been less consistent.

4. It is concluded that the neural influence determining the contractile properties of fast and slow muscle also has a profound controlling influence on the structure and metabolic activity of the muscle fibres.

5. No significant changes could be demonstrated biochemically in the ATPase activities of the fast and slow muscles following cross-innervation.

     

Reinnervation of fast and slow mammalian muscles by a superfluous number of motor axons

In this study peripheral nerves from flexor digitorum longus, (alien nerve) as well as the deep branch of the muscle's own lateral popliteal nerve were cut and connected to the distal stump of the lateral popliteal nerve. Extensor digitorum longus and tibialis anterior muscles then became reinnervated to a similar extent by either nerve, showing no preference for its own nerve. A significant proportion of the endplates in these muscles remained permanently supplied by more than one axon, and a proportion of the muscle fibres was supplied by both nerves. No ectopic endplates were formed on fast muscle fibres. The same two nerves were also connected to the slow soleus muscle and this muscle became preferentially reinnervated by the nerve to flexor digitorum longus. In contrast to fast muscles, endplates of soleus muscle fibres were only rarely contacted by more than one axon, and ectopic endplates were often found in this muscle. In both types of muscles that had an excess of motor nerves, extensive sprouting persisted for many months. Thus, identical motor nerves induce different patterns of innervation in slow and fast muscles, and muscle fibres do not show a preference for their own nerve.