The relative sensitivity of Renshaw cells to orthodromic group Ia volleys caused by static stretch and vibrations of extensor muscles.

1. Activity of Renshaw cells monosynaptically excited by ventral root stimulation and disynaptically excited by electric stimulation of the group Ia afferents in the gastrocnemius-soleus (GS) nerve, was recorded in precollicular decerebrate cats. The response of these units to prolonged vibration applied longitudinally to the deefferented GS muscle was then compared with that elicited by static stretch of the homonymous muscle, for comparable frequencies of discharge of the group Ia afferents. 2. Small-amplitude vibration of the GS muscle at 200/sec for one second produced a sudden increase in the discharge rate of Renshaw cells, which gradually decreased within the first 100 msec of vibration to reach steady albeit lower level than that obtained during the first part of vibration. The response of the Renshaw cells during the first 100 msec of vibration (phasic response) and that elicited during the last 500 msec of vibration (tonic response) were evaluated for different frequencies of sinusoidal stretch. The mean increase in the firing frequency per imp./sec in the Ia afferents was also calculated using the total one-second period. 3. The response of Renshaw cells to muscle vibration increased with the frequency of vibration and, over the value of 10/sec, appeared to be linearly related to the frequency of the input, at least up to the frequency of 150/sec. Since vibration was of sufficient amplitude to produce driving of all the primary endings of muscle spindles, the responses were expressed as mean increases in the discharge rate of Renshaw cells per average impulse/sec in the Ia afferents. The discharge of the Renshaw cell increased on the average by 2.90 and 1.08 imp./sec per each imp./sec in the Ia afferents during the phasic and the tonic component of the response respectively, while the response calculated during the whole period of vibration corresponded on the average to 1.45 imp./sec per each imp./sec in the Ia afferents. 4. The Renshaw cells tested above responded also with increasing frequencies of discharge to increasing levels of static extension of the GS muscle. In particular the discharge frequency of Renshaw cells was on the average linearly related to muscle extension, at least for values ranging from 0 to 8 mm. The mean increase in discharge rate as a function of the static extension corresponded on the average to 0.89 imp./sec/mm. Since the discharge rate of the primary endings of muscle spindles recorded from the deefferented GS muscle increased by 2.62 imp./sec/mm, it appears that the mean increase in the discharge rate of Renshaw cells as a function of static extension corresponded to 0.34 imp./sec per each imp./sec in the Ia afferents.     

Responses of nucleus z neurons to vibration of hindlimb extensor muscles in the decerebrate cat.

1. Small sinusoidal changes of length, applied longitudinally to the ddifferented GS muscle, were uused as a specific stimulus to activate the muscle spindle receptors in precollicular decerebrate cats. In order to determine the relative contribution of the primary and secondary endings of muscle spindles to the response of the nucleus z neurons, the effects of muscle vibration on this unit activity were studied under conditions in which the segmental monosynaptic reflexes produced by stimulation of the primary endings of muscle spindles were continuously monitored. 2. Vibration of the GS muscle (at 180-200/sec and amplitudes up to 250-320 mu) affected the frequency and pattern of discharge in 50 out of 168 units recorded from the lower medulla...     

The sensitivity of Renshaw cells to velocity of sinusoidal stretches of the triceps surae muscle.

1. The electrical activity of Renshaw cells monosynaptically excited by ventral root stimulation and disynaptically excited by electric stimulation of the group I afferents in the GS nerve has been recorded and their response to individual sinusoidal stretches of the deefferented GS muscle tested for different amplitudes and durations of the stimulus. 2. The experimental data indicate that the Rensahw cell responses are not only length dependent but also rate dependent. This finding indicates that the same Renshaw cells receive recurrent collaterals of both tonic and phasic motoneurons. 3. The observation that the discharge of Renshaw cells is particularly sensitive to the velocity of stretch suggests that the recurrent collaterals of large phasic motoneurons, which are recruited during high velocity stretches, exert a stronger excitatory action on Renshaw cells than do axon collaterals of the smaller tonic motoneurons, which are selectively stimulated during low velocity stretches.     

Cholinergic mechanism controlling the response gain of forelimb extensor muscles to sinusoidal stimulation of macular labyrinth and neck receptors

The multiunit EMG activity of the triceps brachii was recorded in precollicular decerebrate cats during roll tilt of the animal or neck rotation at the frequencies of 0.026-0.15 Hz and at the peak amplitude of 10 degrees, leading to selective stimulation of labyrinth or neck receptors. The first harmonic component of the EMG responses to labyrinth stimulation was characterized by an increased activity during side-down tilt of the animal and a decreased activity during side-up tilt; however, just the opposite changes were elicited for the same directions of neck rotation. The peak of the responses was closely related to the extreme animal or neck displacement, thus being attributed to stimulation of position-sensitive macular labyrinth and receptors. Moreover, the modulation as well as the gain of the EMG responses were small in amplitude. Intravenous injections of an anticholinesterase at a dose which in some instances slightly decreased the extensor tonus as well as the background activity of the triceps brachii (eserine sulphate, 0.05-0.075 mg/kg), greatly enhanced the response gain of this extensor muscle to animal tilt or neck rotation at the parameters reported above. This finding was also observed in the absence of any decrease in spontaneous EMG activity of the extensor muscle after injection of the anticholinesterase. In no instance did the phase angle of the response change following these injections. The increased gain of the EMG response of the forelimb extensor muscle to sinusoidal stimulation of labyrinth and neck receptors was first observed 5-10 min after the injection and reached the highest value in about one hour. This effect, was not only time-dependent, but also state-dependent. In fact, the increase in response gain described above either did not occur or was negligible during the sudden recovery of the extensor rigidity which occurred either spontaneously or after somatosensory stimulations. The effects elicited by eserine sulphate were reversed within seconds by a 0.1-0.5 mg/kg dose of atropine sulphate, an anticholinergic drug. It is postulated that for the same labyrinthine or neck signal giving rise to excitatory vestibulospinal volleys acting on extensor motoneurons, the amplitude of the EMG modulation of limb extensor muscles depends on the activity of a cholinergic system.(ABSTRACT TRUNCATED AT 400 WORDS)     

The response of alpha-motoneurons of different size to stretch and vibration of extensor muscles after injection of 5-hydroxytryptophan in spinal cats.

1. Vibration of the GS muscle at 100/sec, and the peak-to-peak amplitude of about 100 mu, produced slight reflex activity of homonymous and heteronymous alpha-motoneurons in the spinal, unanesthetized cat. 2. Intravenous injection of the 5-HT precursor 5-HTP (25 mg/kg, L-form) performed in the spinal preparation facilitated the responses of single extensor alpha-motoneurons to both vibration and stretch of the corresponding muscle. The 5-HT antagonist methysergide (0-5 mg/kg i. v.) blocked almost completely the facilitation of the vibration and stretch induced motoneuronal discharges produced by 5-HTP in the spinal cat, indicating that the facilitation was 5-HT specific. 4. A comparison of the types of alpha-motoneurons participating in the reflex responses elicited by the different stimuli in the spinal cat following injection of 5-HTP showed that both small-tonic and large-phasic alpha-motoneurons take part in both stretch and vibration reflexes. 5. It is concluded that the alpha-motoneuron pool of the GS muscle represents an homogenous population, no matter what kind of synaptic drive is used for its activation, and that the susceptibility of a motoneuron to discharge depends on the amount of the afferent excitatory input rather than on the modalities of sensory stimulation.     

Differential microvascular response to disuse in rat hindlimb skeletal muscles.

The aim of the study was to address discrepant findings in the literature regarding coupling between decreased functional demand during disuse and reduced capillarity. We previously reported [K. Tyml, O. Mathieu-Costello, and E. Noble. Microvasc. Res. 49: 17-32, 1995] that severe disuse of rat extensor digitorum longus (EDL) muscle caused by a 2-wk application of tetrodotoxin (TTX) on the sciatic nerve is not accompanied by capillary loss. Using the same animal model, the present study examined whether this absence of coupling could be explained in terms of 1) too short a duration of disuse and 2) muscle-specific response to disuse. Fischer 344 rats were exposed to either no treatment (control) or to 2- or 8-wk TTX applications. Fiber size, capillary density per fiber cross-sectional area, and capillary-to-fiber (C/F) ratio were determined by morphometry in the EDL muscle (control, 2- and 8-wk groups) and in the superficial portion of medial gastrocnemius (Gas) muscle (control, 2 wk). In both muscles, microvascular blood flow was evaluated by intravital microscopy [red blood cell velocity in capillaries (V(RBC))] and by laser Doppler flowmetry (LDF). Regardless of duration of TTX application or muscle type, TTX-induced disuse resulted in a significant reduction of fiber area (44-71%). However, capillary density increased in EDL muscle (both at 2 and 8 wk) but not in Gas muscle. C/F ratio decreased in EDL muscle at 8 wk (18%) and in Gas muscle (39%). This indicates that the effect on capillarity depended on duration of disuse and on muscle type. V(RBC) and LDF signal were significantly larger in EDL than in Gas muscle. Analysis of change in capillarity vs. V(RBC) suggested that the outcome of disuse may be modulated by blood flow. We conclude that the duration of skeletal muscle disuse per se does not dictate capillary loss, and we hypothesize that discrepant findings of coupling between functional demand and capillarity could be due to the presence/absence of flow-related angiogenesis superimposed on the capillary removal process during disuse.     

Endothelium-dependent vasodilation in different rat hindlimb skeletal muscles.

Few studies have examined potential for endothelium-dependent vasodilation in skeletal muscles of different fiber-type composition. We hypothesized that muscles composed of slow oxidative (SO)- and/or fast oxidative glycolytic (FOG)-type fibers have greater potential for endothelium-dependent vasodilation than muscles composed of fast glycolytic (FG)-type fibers. To test this hypothesis, the isolated perfused rat hindlimb preparation was used with a constant-flow, variable-pressure approach. Perfusion pressure was monitored continuously, and muscle-specific flows were determined by using radiolabeled microspheres at four time points: control, at peak effect of acetylcholine (ACh I; 1-2 x 10(-4) M), at peak effect of ACh after infusion of an endothelial inhibitor (ACh II), and at peak effect of sodium nitroprusside (SNP; 4-5 x 10(-4) M). Conductance was calculated by using pressure and flow data. In the SO-type soleus muscle, conductance increased with ACh and SNP, but the increase in conductance with ACh was partially abolished by the endothelial inhibitor N(G)-nitro-l-arginine methyl ester (control, 0.87 +/- 0.19; ACh I, 2.07 +/- 0.29; ACh II, 1.32 +/- 0.15; SNP, 1.76 +/- 0.19 ml. min(-1). 100 g(-1). mmHg(-1); P < 0.05, ACh I and SNP vs. control). In the FOG-type red gastrocnemius muscle, similar findings were obtained (control, 0.64 +/- 0.11; ACh I, 1.36 +/- 0.21; ACh II, 0.73 +/- 0.16; SNP, 1.30 +/- 0.21 ml. min(-1). 100 g(-1). mmHg; P < 0.05, ACh I and SNP vs. control). In the FG-type white gastrocnemius muscle, neither ACh nor SNP increased conductance. Similar findings were obtained when muscles were combined into high- and low-oxidative muscle groups. Indomethacin had no effect on responses to ACh. These data indicate that endothelium-dependent vasodilation is exhibited by high-oxidative, but not low-oxidative, rat skeletal muscle. Furthermore, endothelium-dependent vasodilation in high-oxidative muscle appears to be primarily mediated by nitric oxide.     

Sarcomere length-joint angle relationships of seven frog hindlimb muscles.

The sarcomere length-joint angle relationship was measured in 7 different muscle-joint complexes (n = 43 muscles) of the frog hindlimb (Rana pipiens). Muscles studied included the cruralis, iliacus internus, gastrocnemius, gluteus magnus, gracilis major, semimembranosus and the semitendinosus. Muscle-joint complexes were mounted in a jig and submerged in chilled Ringer's solution. Joints were rotated throughout their range of motion, while sarcomere length was measured by laser diffraction. Muscles were then formalin fixed and architectural properties determined by microdissection of individual muscle fibers. Sarcomere length change per degree of joint rotation (dLs/d theta) ranged from a low of 3.7 nm/degree for the cruralis muscle acting at the knee to a high of 12.5 nm/degree for the semitendinosus muscle acting at the hip. Values for dLs/d theta were significantly different between all muscles (p < 0.001), and dLs/d theta values for muscles acting at the hip were significantly greater than those for muscles acting at the knee (p < 0.005). dLs/d theta was negatively correlated with fiber length, suggesting a balance between fiber length and moment arm in most muscle-joint systems. However, many exceptions to this generalization were noted. These data suggest that different muscle-joint systems are 'designed' for differential contribution of muscle force production to the joint torque profile. The low variability of these data also suggests that sarcomere number is tightly regulated in these muscle-joint systems but not simply as a result of the total in vivo muscle excursion.     

The effect of stimulation of Golgi tendon organs and spindle receptors from hindlimb extensor muscles on supraspinal descending inhibitory mechanisms

Experiments were performed in precollicular decerebrate cats to investigate whether proprioceptive volleys originating from Golgi tendon organs and muscle spindles may activate supraspinal descending inhibitory mechanisms. Conditioning stimulation of the distal stump of ventral root filaments of L7 or S1 leading to isometric contraction of the gastrocnemius-soleus (GS) muscle inhibited the monosynaptic reflex elicited by stimulation of the ipsilateral plantaris-flexor digitorum and hallucis longus (Pl-FDHL) nerve. The amount and the time course of this Golgi inhibition were greatly increased by direct cross-excitation of the intramuscular branches of the group Ia afferents due to ephaptic stimulation of the sensory fibers, which occurred when a large number of a fibers had been synchronously activated. The postsynaptic and the presynaptic nature of these inhibitory effects, as well as their segmental origin, have been discussed. In no instance, however, did the stimulation of Golgi tendon organs elicit any late inhibition of the test monosynaptic reflex, which could be attributed to a spino-bulbo-spinal (SBS) reflex. Conditioning stimulation of both primary and secondary endings of muscle spindles, induced by dynamic stretch of the lateral gastrocnemius-soleus (LGS) muscle, was unable to elicit any late inhibition of the medial gastrocnemius (MG) monosynaptic reflex. The only changes observed in this experimental condition were a facilitation of the test reflex during the dynamic stretch of the LGS, followed at the end of the stimulus by a prolonged depression. These effects however were due to segmental interactions, since they persisted after postbrachial section of the spinal cord. Intravenous injection of an anticholinesterase, at a dose which greatly potentiated the SBS reflex inhibition produced by conditioning stimulation of the dorsal root L6, did not alter the changes in time course of the test reflex induced either by muscle contraction or by dynamic muscle stretch. Conditioning stimulation of a muscle nerve activated the supraspinal descending mechanism responsible for the inhibitory phase of the SBS reflex only when the high threshold group III muscle afferents (innervating pressure-pain receptors) had been recruited by the electric stimulus. This finding contrasts with the great availability of the system to the low threshold cutaneous afferents. The proprioceptive afferent volleys originating from Golgi tendon organs as well as from both primary and secondary endings of muscle spindles, contrary to the cutaneous and the high threshold muscle afferent volleys, were apparently unable to elicit not only a SBS reflex inhibition, but also any delayed facilitation of monosynaptic extensor reflexes attributable to inhibition of the cerebellar Purkinje cells.     

Force-velocity properties of two avian hindlimb muscles

Recent work has provided measurements of power output in avian skeletal muscles during running and flying, but little is known about the contractile properties of avian skeletal muscle. We used an in situ preparation to characterize the force-velocity properties of two hind limb muscles, the lateral gastrocnemius (LG) and peroneus longus (PL), in Wild Turkeys (Meleagris gallopavo). A servomotor measured shortening velocity for at least six different loads over the plateau region of the length-tension curve. The Hill equation was fit to the data to determine maximum shortening velocity and peak instantaneous power. Maximum unloaded shortening velocity was 13.0+/-1.6 L s(-1) for the LG muscle and 14.8+/-1.0 L s(-1) for the PL muscle (mean+/-S.E.M.). These velocities are within the range of values published for reptilian and mammalian muscles. Values recorded for maximum isometric force per cross-sectional area, 271+/-28 kPa for the LG and 257+/-30.5 kPa for the PL, and peak instantaneous power output, 341.7+/-36.4 W kg(-1) for the LG and 319.4+/-42.5 W kg(-1) for the PL, were also within the range of published values for vertebrate muscle. The force-velocity properties of turkey LG and PL muscle do not reveal any extreme differences in the mechanical potential between avian and other vertebrate muscle.     

Fatigability of rat hindlimb muscles after acute irreversible acetylcholinesterase inhibition.

The purpose of this study was to investigate the functional impact of acute irreversible inhibition of acetylcholinesterase (AChE) on the fatigability of medial gastrocnemius and plantaris muscles of Sprague-Dawley rats. After treatment with methanesulfonyl fluoride (a lipid-soluble anticholinesterase), which reduced their AChE activity by >90%, these muscles were subjected to an in situ indirect stimulation protocol, including a series of isolated twitch and tetanic contractions preceding a 3-min fatigue regimen (100-ms trains at 75 Hz applied every 1.5 s). During the first minute of the fatigue regimen, the effects of AChE inhibition were already near maximal, including marked reductions in peak tension and the force-time integral (area), as well as a decrement of compound muscle action potential amplitudes within a stimulus train. Neuromuscular transmission failure was the major contributor of the force decreases in the AChE-inhibited muscles. However, despite this neuromuscular transmission failure, muscles of which all AChE molecular forms were nearly completely inhibited were still able to function, although abnormally, during 3 min of intermittent high-frequency nerve stimulation.     

Mechanical properties of locust extensor tibiae muscles

• 1.1. The prothoracic and mesothoracic extensor tibiae muscles of the locust respond to activity in the “slow” extensor tibiae motoneuron (SETi) with very slow contractions and a low fusion frequency, while their phasic contractions are more rapid than those of the metathoracic extensor tibiae muscle.
• 2.2. SETi activity can induce a memory or “catch” effect in which a high tension is maintained by a lower frequency than is needed to develop it. “Catch” tension is reduced by phasic contractions of the muscle or by activity in the inhibitory axon.
• 3.3. A bundle of tonic fibres isolated from the metathoracic extensor tibiae muscle exhibits co-ordinated rhythmic contractions similar to those recorded from intact muscles.
• 4.4. Depolarizations of the tonic fibres coincide with the contractions and are sometimes accompanied by bursts of EPSPs and IPSPs.
• 5.5. The tonic fibres are electrically-coupled.

     

The silent period in the stretch response of ia-activated dorsal spino-cerebellar tract neurons to sinusoidal muscle stretch in cats

By means of extracellular recordings of action potentials the stretch responses of single neurons of Clarke's column were analysed. The neurons were monosynaptically activated from Ia afferents of both ipsilateral gastrocnemius muscles. When stretch cycles of more than 0.2 mm amplitude and frequencies above 2 Hz were applied to the gastrocnemius muscles, the discharging was found to cease during the period of stretch release, whereas the average discharge rate was found to increase. In the frequency range between 0.1 and 10 Hz a sinewave of stretch frequency — the response sinewave — fitted to the non-zero bins of cycle histograms described the stretch response at small and large amplitudes equally well. The amount of increase in the average firing rate corresponded quite well to the portion of the response sinewave below the zero discharge rate. This indicates that the occurance of discharge pauses and the relation of the average discharge rate to frequency and amplitude of stretch can be described successfully by a half-wave rectification of the response at zero discharge rate. If one regards the shape of cycle histograms to be a nearly sinusoidal modulation plus a non-linear clipping at zero the application of linear systems analysis is worthwhile in describing the response not only at very small amplitudes but in the whole range of muscle stretch.     

Mechanical characteristics and fiber composition of human leg extensor muscles.

To investigate the influence of skeletal muscle fiber composition on the mechanical performance of human skeletal muscle under dynamic conditions, 34 physical education students with differing muscle fiber composition (M. vastus lateralis) were used as subjects to perform maximal vertical jumps on the force-platform. Two kinds of jumps were performed: one from a static starting position (SJ), the other with a preliminary counter-movement (CMJ). The calculated mechanical parameters included height of rise of center of gravity (h), average force (F), net impulse (NI) and average mechanical power (W). It was observed that the percentage of fast twitch fibers was significantly related (p less than 0.05--0.01) to these variables in SJ condition and also to h and NI of the positive work phase in CMJ. It is concluded that skeletal muscle fiber composition also determines performance in a multijoint movement. The result is explainable through the differences in the mechanical characteristics of the motor units and their respective muscle fibers.     

Free radical activity following contraction-induced injury to the extensor digitorum longus muscles of rats.

The purpose of the study was to investigate the role of free radicals in the injury induced by a protocol of repeated pliometric (lengthening) contractions to the extensor digitorum longus (EDL) muscle in situ in rats. Previous data have indicated that prior treatment with the antioxidant polyethylene glycol-superoxide dismutase reduced the damage that was apparent at 3 days following this type of exercise. Three hours and 3 days following the protocol, the magnitude of the semiquinone-derived free radical signal observed by electron spin resonance spectroscopy (ESR) was not different for exercised and non-exercised skeletal muscles. A reduction in the protein thiol content of muscle was evident at 3 h, and was still apparent at 3 days. Three hours after the protocol, the total muscle glutathione content and the percentage in the oxidized form were unchanged, but by 3 days the percentage of muscle glutathione present in the oxidized form was elevated. The susceptibility of muscle to lipid peroxidation in vitro was reduced 3 days after the pliometric contractions. These data indicate that oxidation of protein thiols and glutathione may be involved in the secondary damage following pliometric contractions, but provide no evidence that the species involved were derived from mitochondrial semiquinone radicals.