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T C Cope

Publications and source records attributed to T C Cope.

At least 37 records · Page 2Linked to original sources

Stability of motor-unit force thresholds in the decerebrate cat.

To further test the hypothesis that some fixed property of motoneurons determines their recruitment order, we quantified the variation in force threshold (FT) for motoneurons recruited in muscle stretch reflexes in the decerebrate cat. Motor axons supplying the medial gastrocnemius (MG) muscle were penetrated with micropipettes and physiological properties of the motoneuron and its muscle fibers, i.e., the motor unit, were measured. FT, defined as the amount of MG force produced when the isolated motor unit was recruited, was measured from 20 to 93 consecutive stretch trials for 29 motor units. Trials were selected for limited variation in base force and rate of rise of force, which have been shown to covary with FT, and in peak stretch force, which gives some index of motor-pool excitability. Under these restricted conditions, large variation in FT would have been inconsistent with the hypothesis. Analysis of the variation in FT employed the coefficient of variation (CV), because of the tendency for FT variance and mean to increase together. We found that CV was distributed with a median value of 10% and with only 2 of 29 units exceeding 36%. Some of this variation was associated with measurement error and with intertrial fluctuations in base, peak, and the rate of change of muscle force. CV was not significantly correlated with motor-unit axonal conduction velocity, contraction time, or force. In three cases FT was measured simultaneously from two motor units in the same stretch trials. Changes in recruitment order were rarely observed (5 of 121 stretch trials), even when FT ranges for units in a pair overlapped. We suggest that the large variation in recruitment threshold observed in some earlier studies resulted not from wide variation in the recruitment ranking of motoneurons within one muscle, but rather from variation in the relative activity of different pools of motoneurons. Our findings are consistent with the hypothesis that recruitment order is determined by some fixed property of alpha-motoneurons and/or by some unvarying combination of presynaptic inputs that fluctuate in parallel.

Animals↗

Recruitment of triceps surae motor units in the decerebrate cat. I. Independence of type S units in soleus and medial gastrocnemius muscles.

1. We tested the hypothesis that reflex inhibition of soleus motor units reflects selective inhibition of slow-twitch (type S) motor units throughout the triceps surae. Physiological properties including type, together with firing behavior, were measured from single motor units in the medial gastrocnemius (MG) muscle of decerebrate cats with the use of intra-axonal recording and stimulation. MG unit firing was contrasted during net inhibition or excitation of the slow-twitch soleus muscle produced by ramp-hold-release stretches of MG. 2. Stretch of the MG muscle increased the firing of type S motor units in the MG regardless of the reflex response of the soleus muscle. When stretch inhibited soleus, each of the 14 type S units sampled from MG either was newly recruited or exhibited increases in the rate of ongoing firing. Increased firing was observed in 320 of 321 stretch trials. For 8 of these 14 units, a total of 155 stretch trials evoked reflex excitation of soleus, and unit firing increased in all trials. 3. For the eight MG type S motor units studied during both reflex inhibition and excitation of soleus, firing rate tended to be higher during inhibition. The higher rates were also associated with the higher MG forces required to elicit soleus inhibition. For one MG type S unit it was possible to compare firing rates during soleus inhibition and excitation for trials of overlapping levels of MG force. For this unit, firing rate was similar, but still appreciably higher, during inhibition. 4. Soleus inhibition was also produced by stretch of the plantaris (PL) or lateral gastrocnemius (LG) muscles. Type S units in PL (n = 2) or in LG (n = 1) were recruited or increased firing rate even when stretch of these muscles produced soleus inhibition. 5. The firing behavior of 12 fast-twitch (type F) units was studied (11 from MG, 1 from PL). All type F units either were recruited or accelerated the rate of firing during soleus inhibition, as well as during soleus excitation. 6. These findings give evidence that reflex inhibition of type S motor units in the soleus muscle does not necessarily reflect an organizational scheme in which there is inactivation of type S units in other active muscles. In the DISCUSSION we point out the absence of direct evidence for selective inactivation of units on the basis of their type classification.

Animals↗

Recruitment of triceps surae motor units in the decerebrate cat. II. Heterogeneity among soleus motor units.

1. On the basis of the orderly activation of motoneurons in a pool, one would predict that motor unit activity and whole muscle force will change at least roughly in parallel: active motor units should continue to fire as net muscle force increases and quiescent motor units should remain inactive as muscle force decreases. We have consistently observed this relationship in our studies of the medial gastrocnemius (MG) muscle, but here we report an uncoupling of the soleus muscle and some of its motor units. 2. Physiological properties and firing behaviors of 20 soleus motor units were characterized in five decerebrate cats with the use of intra-axonal stimulation and recording. Motor unit firing was elicited in reflexes initiated by muscle stretch, nerve stimulation, and mechanical stimulation of the heel. Particular emphasis was placed on the heterogenic reflexes produced in soleus by ramphold-release stretches of the MG muscle. In agreement with previous reports, either net heterogenic excitation or inhibition of the soleus muscle was produced in separate trials of MG stretch. 3. During excitation of soleus in autogenic stretch reflexes and in crossed-extension reflexes, all 20 units were recruited or increased firing, i.e., unit firing was coupled with soleus force. In the other reflexes, however, unit firing and muscle force were uncoupled for 10 of these units. Six tonically active motor units were inhibited during an increase in soleus force produced by MG stretch or by mechanical stimulation of the heel. Four motor units were activated during a decrease in soleus force produced by the same stimuli. 4. Six motor units were studied during both soleus inhibition and excitation evoked by MG stretches. One motor unit was consistently coupled to the soleus muscle response; firing increased during soleus excitation and decreased during inhibition. However, four soleus motor units were inhibited under both conditions, and one unit was excited under both conditions. Thus the firing behavior of five of these six motor units was the same in response to MG stretch, irrespective of the soleus response. 5. The uncoupling was most clearly recognized when tonically active units ceased firing during net excitation of the soleus muscle and when silent units began firing during net inhibition of the soleus muscle. Unit responses were not as striking in all trials of MG stretch (spike number increased or decreased relative to prestretch values by 1-4 spikes), but the responses were consistent across trials; in multiple stretches, spike number commonly either increased or decreased. Intertrial regularity was also observed in units for which firing was coupled with the net reflex response of the soleus muscle. 6. Divergence in the firing of soleus motor units was also observed in three cases in which records were taken simultaneously from two motor units. In one pair, one unit increased and the other decreased firing during MG stretch-evoked inhibition of soleus. In the other two pairs, one unit increased and the other decreased firing when soleus was excited by heel stimulation. In all pairs, the unit that decreased firing under these conditions had the lowest recruitment threshold in response to the soleus stretch. 7. Although all soleus motor units were classified as slow-twitch (type S), variation in their physiological properties bore some relation to firing behavior. Those units recruited during periods of soleus inhibition exhibited among the fastest conduction velocities and contraction times in our sample. In all three unit pairs sampled, the unit expressing decreases in firing had the slower conduction velocity and contraction time. 8. These findings demonstrate that soleus motor units are differentially activated and deactivated by peripheral afferents. (ABSTRACT TRUNCATED)

Action Potentials↗

Reinnervated muscles fail to produce stretch reflexes.

1. We studied the stretch-evoked reflex organization of hind limb muscles in two decerebrate cats 36 mo after unilateral section and immediate surgical repair of the common nerve supplying the lateral gastrocnemius (LG) and soleus (S) muscles. 2. The production of considerable reflex force by reinnervated muscles in response to electrical stimulation of uninjured nerves indicated substantial functional recovery of motor units. However, reduction in the responsiveness of reinnervated muscles to stretch of the untreated medial gastrocnemius (MG) muscle indicated some deficit in recovery of normal synaptic integration. 3. Stretch failed to elicit autogenic excitation of the reinnervated S and LG. This failure was observed whether the reinnervated muscles were quiescent or contracting in other reflexes. 4. The heterogenic reflex organization of reinnervated muscles was abnormal. Stretch of the reinnervated S failed to evoke heterogenic reflexes both in the untreated MG and in the reinnervated LG. Stretch of the reinnervated LG failed to produce excitation of MG. 5. These findings demonstrate deficiencies in proprioceptive feedback from reinnervated muscles and lead us to expect incomplete recovery of motor function after nerve section.

Afferent Pathways↗

Spatial arrangement and metabolic capacity of fiber types in self-reinnervated cat muscle.

The recovery potential of skeletal muscle was explored by examining cat muscle between 10 and 33 mo after complete transection and immediate surgical reunion of its own nerve. Biochemical analysis of single muscle fibers showed that the activities of key enzymes in energy metabolism (malate and lactate dehydrogenase and adenylokinase) were similar to normal for their respective fiber types, suggesting that incomplete recovery of the ability to sustain submaximal contraction in reinnervated muscles (T.C. Cope, C.B. Webb, and B.R. Botterman. J. Neurophysiol. 65: 648-656, 1991) is explained in some other way. Two independent statistical procedures for assessing the randomness of adjacencies of histochemically identified fiber types showed type grouping in some areas, but there were also many regions with randomly distributed fiber types. These findings demonstrate the potential for substantial recovery of both energy metabolism and dispersion of fiber types after self-reinnervation.

Animals↗

Cutaneous stimulation fails to alter motor unit recruitment in the decerebrate cat.

1. An attempt was made to repeat the observation that cutaneous input to the cat medial gastrocnemius (MG) muscle sometimes had the differential effect of inhibiting motoneurons with slow axonal conduction velocity while simultaneously exciting others with fast conduction velocity. Dual microelectrode recording from intact ventral root filaments was used to study the effects of cutaneous inputs on recruitment order and on firing frequency of physiologically characterized MG motor units in decerebrate cats. Motor responses to pinch of the skin over the lateral surface of the ankle as well as electrical stimulation of the caudal cutaneous sural (CCS) nerve were contrasted with the responses to static muscle stretch as well as muscle vibration. 2. In contrast to the prediction, recruitment order in pairwise tests was the same for skin pinch or CCS stimulation as it was for MG stretch or vibration in all 32 tested pairs of motor units. This sample included seven pairs comprising one slow-twitch (S) and one fast-twitch motor unit, where the predicted reversal of recruitment should have been most apparent. Regardless of the source of excitation, recruitment of motor units of the MG was consistent with Henneman's size principle in approximately 90% of trials. 3. Skin pinch increased the firing rate of 30 of 32 individual motor units previously activated by stretch or vibration, including 7 slow-twitch units. In the remaining two units, skin pinch transiently (100-400 ms) slowed the firing of an S unit in 11 of 13 vibration + pinch trials. The other unit (type unknown) showed one or two retarded spikes in each of four vibration + pinch trials. In three S units, including the lone inhibitable unit and two others that were only excited by skin pinch, there was a significant positive rank correlation between change in unit firing frequency and change in soleus integrated electromyographic activity.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Motor-unit recruitment in self-reinnervated muscle.

1. Recruitment order of motor units in self-reinnervated medial gastrocnemius (MG) muscles was studied in decerebrate cats 16 mo after surgical reunion of the cut MG nerve. Pairs of MG motor units were isolated by dual microelectrode penetration of ventral roots to measure their recruitment sequence during cutaneous reflexes in relation to their physiological properties. 2. Physiological properties of reconstituted motor units appeared normal, as expected. Also normal were the relationships among these properties: twitch and tetanic tension tended to increase with axonal conduction velocity and decrease with twitch contraction time. A small fraction of motor units (10/116) in reinnervated muscles produced either no measurable tension or unusually large amounts of tension compared with controls. This was the only distinct feature of the sample of reconstituted units. 3. In muscles reinnervated after nerve section, stretch was notably ineffective in eliciting reflex contraction of MG muscles or their constituent motor units (only 5/116 units). Incomplete recovery from nerve section was probably the cause of this impairment, because stretch reflexes were readily evoked in adjacent untreated muscles and in one reinnervated MG muscle that was studied 16 mo after nerve crush. In contrast with the ineffectiveness of muscle stretch, sural nerve stimulation succeeded in recruiting 49/116 units, a proportion fairly typical of normal MG muscles. 4. The contractions of the first unit recruited in cutaneous reflexes tended to be slower and less forceful than those of the other unit in a pair. By these measures, recruitment obeyed the size principle. This recruitment order with respect to unit contractile properties was not significantly different (P > 0.05) between untreated and reinnervated muscles but was significantly (P < 0.005) different from random order in both groups. The same recruitment pattern was observed for pairs of motor units sampled from the muscle reinnervated after nerve crush, whether units were recruited by muscle stretch or sural nerve stimulation. 5. The usual tendency for motor units with slower conduction velocity (CV) to be recruited in sural nerve reflexes before those with faster CV was not strong in reinnervated muscles. After nerve section the proportion of units exhibiting the usual recruitment pattern was not significantly different (P > 0.05) from a random pattern for CV. 6. The central finding is that the normal recruitment patterns recover from nerve injury in a muscle that is reinnervated by its original nerve. By contrast, stretch reflexes do not recover well from nerve section, and this deficiency may contribute to motor disability.

Animals↗

Influence of synaptic identity on single-Ia-afferent connectivity and EPSP amplitude in the adult cat: homonymous versus heteronymous connections.

1. This study makes use of the pattern of synaptic connections between motoneurons and Ia afferents of triceps surae muscles in the cat to test the relative importance of synaptic identity, neuronal size, and neuronal topography as determinants of Ia-afferent connectivity and excitatory postsynaptic potential (EPSP) amplitude. 2. The synaptic actions of single-Ia medial gastrocnemius (MG) afferents were measured by intracellular recording in MG and lateral gastrocnemius (LG) motoneurons. The spike-triggered averaging technique was used to measure EPSPs generated by homonymous or heteronymous Ia afferents and motoneurons, i.e., neurons supplying the same or different muscles, respectively. In agreement with earlier studies, the pooled sample showed that the number of functional connections and the size of EPSPs were both significantly greater for homonymous than for heteronymous neurons. 3. Afferent conduction velocity, motoneuron conduction velocity, rheobase current, and position of the motoneuron relative to the spinal cord afferent entry were all correlated with EPSP amplitude, but the amplitude difference between homonymous and heteronymous connections remained significant after the statistical removal analysis of covariance (ANCOVA) of the contribution of these variables. Stepwise multiple-regression analysis showed that synaptic identity explained the greatest fraction of the variance in EPSP amplitude (9%), with significant but smaller fractions accounted for by rheobase current or motoneuron conduction velocity. 4. In a separate experiment, the monosynaptic affects from both homonymous and heteronymous single-Ia afferents were examined in each of 88 MG or LG motoneurons. The single-Ia afferents used in this portion of the study were sampled from both MG and LG muscles and selected for similar conduction velocities and spinal cord entry points.(ABSTRACT TRUNCATED AT 250 WORDS)

Afferent Pathways↗

Modulation of la EPSP amplitude: the effects of chronic synaptic inactivity.

In this study, we test the hypothesis that monosynaptic connections between la afferents and spinal motoneurons are strengthened by chronic disuse. Impulse activity along the medial gastrocnemius (MG) nerve was blocked for 2 weeks using TTX delivered by an osmotic minipump to a Silastic cuff placed around the nerve. The duration and specificity of this block were confirmed by chronic EMG recordings from several triceps surae muscles. The effect of TTX-induced inactivity of presynaptic elements on EPSP amplitude was distinguished from the effect of treating the postsynaptic target by comparing the results from heteronymous synaptic connections, where only one or the other element was treated. After 2 weeks of synaptic inactivity, the heteronymous EPSPs generated by MG la afferents in lateral gastrocnemius/soleus (LG-Sol) motoneurons were significantly (p less than 0.005) larger than control values (48%). Sample differences in rheobase current and half-afterhyperpolarization, both of which may covary with EPSP amplitude, did not account for the differences between groups. Segregation of the two samples of motoneurons by rheobase current identified the increase as being confined to those LG-Sol cells whose rheobase fell below 10 nA. In addition, EPSPs generated by untreated LG-Sol la afferents in treated MG motoneurons were significantly enhanced (39%, p less than 0.05). Thus, TTX treatment of either presynaptic or postsynaptic elements increases synaptic strength. This increase in monosynaptic EPSP amplitude following TTX-induced inactivity may reflect an alteration intrinsic to the la afferent to motoneuron synapse, but influences from extrinsic sources cannot be discounted.

Action Potentials↗

Control of motor-unit tension by rate modulation during sustained contractions in reinnervated cat muscle.

1. The aim of this study was to describe the control of tension by rate modulation of single motor units in reinnervated muscle. 2. Single fast-twitch motor units were isolated from medial gastrocnemius (MG) muscles in two groups of anesthetized adult cats: one in which the MG nerve was left untreated and another in which that nerve was sectioned and immediately sutured together 10-33 mo before study. Together with conventional measures of isometric contractile properties, units were tested with the use of computer-controlled feedback regulation of stimulation rate to maintain tension during continuous isometric contraction at a constant submaximal level [25% of maximal tension (Pmax)]. 3. For motor units from both groups, stimulation rate began to decline after target tension was attained and then settled at lower values for variable durations before rapidly increasing, usually within the last 5% of the contraction's duration, until reaching the experimentally selected limit of 100 pulses/s (pps). 4. Measures of the declining phase in stimulation rate occurring at the beginning of sustained contraction were not significantly different in comparison of untreated versus reinnervated muscles. These measures included 1) the magnitude of the decrease in rate, 2) the minimum rate attained, and 3) the time taken to reach minimum stimulation rate expressed as a fraction of endurance time (Et, total duration of the sustained contraction). 5. Most fast-twitch units from reinnervated muscles fell within normal limits for both endurance time and the number of stimuli applied over that period.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Motor-unit recruitment in the decerebrate cat: several unit properties are equally good predictors of order.

1. Recruitment order was studied in pairs of motor units of the medial gastrocnemius (MG) muscle of decerebrate cats with the use of dual microelectrode recording from intact ventral root filaments. Excitation was provided by stretch of MG, stretch of synergists [lateral gastrocnemius (LG), plantaris (PL), and soleus (SOL) muscles] or electrical stimulation of the caudal cutaneous sural (CCS) nerve. Motor units were characterized by axonal conduction velocity (CV), tetanic tension (Pmax), twitch contraction time (CT), and fatigue index (FI). 2. Consistent with the recruitment pattern described by others, most often in relation to either CV or Pmax, the first unit of a pair to be recruited by MG stretch was typically the one with the lower CV and Pmax, and the higher FI and CT. The proportion of pairs that agreed in rank order of each property and recruitment order was as follows: for CT, 94%; for CV, 87%; for Pmax, 84%; and for FI, 75%. With a single marginal exception (CT vs. FI), no motor-unit property proved to be significantly better than the others at predicting recruitment (G test; P greater than 0.05). 3. In all 11 tested pairs containing one slow (type S) and one fast (type F) unit, the S was more easily recruited by stretch. Type F units divided into groups with high (type FR), low (type FF), and intermediate (type FInt) values for FI were recruited in order from FR to FInt to FF in 8/11 pairs. Thus our findings were similar to earlier demonstrations that recruitment proceeds in order by type. 4. Stretch of MG synergists usually recruited units in the same order as MG stretch. In two S-S pairs, recruitment order was switched with synergist stretch. 5. Stimulation of the CCS nerve was generally excitatory to the MG units sampled. Most unit pairs were recruited by CCS stimulation in the same order as by MG stretch, but, for 6 of 39 pairs, CCS stimulation switched the order produced by stretch. Thus, whereas sural afferent input can preferentially excite some units over others as suggested by Kanda et al., that effect is not widespread or selective for unit type under these conditions. 6. Assuming that all MG motor units cooperate as a single functional pool in homonymous stretch reflexes, we support others in concluding that a motoneuron's recruitment threshold is not strictly determined by its size. However, our data do not distinguish other schemes that predict recruitment order more accurately than the size principle.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

Nonuniform fatigue characteristics of slow-twitch motor units activated at a fixed percentage of their maximum tetanic tension.

1. The endurance of slow-twitch motor units from the soleus (SOL) and medial gastrocnemius (MG) muscles of the cat were tested by determining the length of time (endurance time, Et) that a unit could maintain its tension output at 85% of maximum. Motor-unit tension was clamped at the target level by altering the stimulation rate of a unit's motor axon through computer feedback control. Tested in this way, units of both muscles displayed a wide range of Ets, approximately 40- to 50-fold. 2. Electromyographic (EMG) waveforms of motor units subjected to force-clamp contractions were analyzed to access whether any activity-dependent changes in their waveform shape might predict Et. Three measurements of waveform shape were determined: baseline-to-baseline duration, peak-to-peak amplitude, and area. Typically, amplitude decreased and duration increased as a contraction proceeded, whereas area remained fairly constant. Because changes in each measure were very similar for units of widely different Ets, it was concluded that neuromuscular junction failure and changes in the excitability of the sarcolemma (excluding the t-tubule system) play a minor role in determining Et. 3. Et was highly correlated with the mean stimulation rate (Et/number of stimuli) used during the force-clamp contractions. Mean rate was seen to progressively decrease with increasing Et. This correlation could not be explained by measures of isometric contractile speed or relaxation (e.g., twitch contraction time or half-relaxation time) measured before the force-clamp contractions. Both contraction time and half-relaxation time were found to be unrelated to both Et and the rate used to stimulate the unit during the force-clamp contraction. 4. Among type S units of SOL and MG, maximum tetanic tension and Et were not related. A significant relation (r = -0.49) was found between axonal conduction velocity and Et for SOL units (n = 38). In addition, a significant correlation (r = 0.47) was found between conduction velocity and tetanic tension for SOL units. Perhaps because of the small sample of type S units from MG (n = 10), conduction velocity was found not be related to either Et or tetanic tension. 5. Others have shown that a motor unit's maximum tetanic tension and axonal conduction velocity are correlated with its order of recruitment among motoneurons innervating a muscle. Recent work has further shown that among type F units the order in which a motoneuron is recruited is highly correlated with the fatigue resistance of its muscle unit.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Motor-unit stimulation patterns during fatiguing contractions of constant tension.

1. Through computer feedback control, muscle-unit tension was maintained by altering the stimulation rate of a functionally isolated motor axon. The required stimulation patterns and fatigue properties of motor units from the flexor carpi radialis (FCR), flexor digitorum longus (FDL), and medial gastrocnemius (MG) muscles of the cat were studied when tension was maintained or "clamped" at a constant average level (25% of maximum tetanic tension). 2. In each muscle, two distinct stimulation patterns were observed during constant-tension contractions, one associated with slow-twitch units and the other with fast-twitch units. Once target tension was reached, slow-twitch units required fairly constant rates in order to maintain a constant force, whereas fast-twitch units displayed a marked decline in rate during the early phases of the contraction, averaging between 42 and 54% for the three muscles. The decline in rate most likely represented potentiation of the contractile response and slowing of contractile speed. In general, slow-twitch units responded with lower mean rates (approximately 14 pps less), averaged over the course of the contraction, than fast-twitch units. 3. For fast-twitch units of each muscle, resistance to fatigue varied continuously and over a wide range. The duration that tension could be maintained at 25% of maximum, defined as endurance time, ranged between 16 and 2063 s. No categorization of fast-twitch units into groups could be made on the basis of endurance time. Of the 5 slow-twitch units followed beyond 2700 s, only one failed to maintain tension during the observation period. 4. For hindlimb fast-twitch units, endurance was independent of the stimulation rate needed to maintain tension during the contraction. By contrast, there was a significant tendency for an inverse relation between endurance time and mean stimulation rate for FCR fast-twitch units. 5. Recovery of maximum tension was evaluated at 30 s, 1 min, 2 min, and 5 min following a constant-tension contraction. After a 5-min rest, fast-twitch units were able to produce an average of 80-85% of their maximum tetanic tension. By using the median endurance time (approximately 100 s) to divide the fast-twitch population into "low" and "high" endurance groups, recovery of tension was found not to be uniform among the two groups. High endurance units were able to recover a greater percentage of their original maximum tetanic tension. No difference was found between force recovery for low and high endurance units at 30 s.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Maximum tension predicts relative endurance of fast-twitch motor units in the cat.

1. The relationships between maximum tetanic tension (P0), endurance time, and axonal conduction velocity (CV) were investigated in fast-twitch motor units of the cat flexor carpi radialis (FCR) and medial gastrocnemius (MG) muscles, and in one flexor digitorum longus (FDL) muscle. Endurance time was the length of time that a unit could maintain 25% of its maximum tetanic tension during a sustained contraction. Motor-unit tension was "clamped" at 25% of maximum by altering the stimulation rate of a unit's motor axon through computer feedback control. 2. In individual experiments, including the one investigated FDL muscle, an inverse relation was consistently found between maximum tension and endurance time. Pooled data from the FCR and MG muscles also resulted in significant correlations between maximum tetanic tension and endurance time. 3. Following the force-clamp contraction, some motor units were subjected to the standard fatigue test of Burke and colleagues (6). Motor units were classified as type FR (fast twitch, fatigue resistant) or type FF* (fast twitch, fast fatiguing after the force-clamp contraction). For both type FR and FF* units, maximum tetanic tension and endurance time were found to be inversely related. However, no correlation was found between maximum tetanic tension and fatigue index for type FR units. Only when all type F (FR + FF*) units were considered as a population was there a significant correlation between these two properties. 4. Other investigators have shown that maximum tetanic tension and axonal conduction velocity are highly correlated with the recruitment order of motoneurons (e.g., Refs. 2, 26). Endurance time was found to be more tightly coupled with contraction strength than with conduction velocity. In 12 of 14 experiments, significant Spearman rank correlation coefficients were found between endurance time and tension, whereas significant correlations were found in only 3 of 14 experiments for endurance time and conduction velocity. 5. Pairs of motor units isolated from the same muscle were formed to see if the unit with the smaller tension had the slower conduction velocity and the longer endurance time. Across all muscles, the probability that the unit with the smallest tension had the greatest endurance time was 0.91 (441 of 487 pairs). By contrast, the probability that the least forceful unit of the pair had the slowest conduction velocity was 0.61. 6. In four experiments, pairs of type-identified units were examined. Among FR-FR pairs, the least forceful unit had the greatest endurance time in 88% of 43 pairs. For FF*-FF* pairs, the percentage was somewhat lower, 72% of 29 pairs.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Acute effects of spinal transection on EPSPs produced by single homonymous Ia-fibers in soleus alpha-motoneurons in the cat.

1. Excitatory postsynaptic potentials (EPSPs) generated in soleus motoneurons by single homonymous Ia-fibers were measured using intracellular recording and the spike-triggered averaging technique. Two groups of barbiturate-anesthetized adult cats were studied: one with the spinal cord intact and the other with the spinal cord severed at thoracic segment 13 (T13) several hours prior to recording. 2. In cord-transected cats, single homonymous Ia-fibers produced EPSPs in soleus motoneurons that were, on average, larger and faster rising relative to normal, as they are for those produced in medial gastrocnemius (MG) motoneurons (8, 12, 13, 40). Specifically, mean EPSP amplitude and rise time were, respectively, 261 +/- 22 microV and 0.65 +/- 0.05 ms for the transected group vs. 160 +/- 21 microV and 0.96 +/- 0.08 ms for the intact group. The group means for each parameter were significantly different (P less than 0.005). 3. The group difference in EPSP amplitude was largely due to a decrease in number of small EPSPs in the transected group (11% less than 100 microV compared with the normal 41%) and not due to the occurrence of unusually large ones. Ratios of the largest to smallest amplitude EPSPs produced in the same motoneuron were similarly distributed for intact and transected groups, implying that the effect of transection on EPSP size was uniform across different Ia-fiber synapses made with the same motoneuron. Mean EPSP amplitude for each transected cat (n = 5) was larger than normal, but in some cases the increase took greater than 10 h to express itself. 4. The normal tendency for EPSP rise time to decline on average with amplitude was absent in the transected group, wherein rise time was reduced to similar average values in all amplitude categories. This suggests that the decrease in rise time occurred independently of the increase in amplitude. In contrast, EPSP half-width, which tended tow ward lower than normal values [5.63 +/- 0.36 (SE) ms vs. 6.51 +/- 0.44 ms; P greater than 0.10], decreased in proportion with rise time as evidenced by the preservation of the normal relation between those parameters in transected cats. Normalizing EPSPs by motoneuron time constant (tau) reduced the group differences in rise time and half-width, suggesting that a fall in tau contributes to the abbreviation of EPSP time course. 5. The condition of the spinal cord best accounted for differences in synaptic strength between groups.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Cross-correlation assessment of synaptic strength of single Ia fibre connections with triceps surae motoneurones in cats.

1. The relation between excitatory post-synaptic potentials (e.p.s.p.s) by single I a fibres and the resultant cross-correlograms in triceps surae motoneurones was investigated in barbiturate-anaesthetized cats. The e.p.s.p.s. were documented first, using the discharge of single Ia fibres evoked by muscle stretch to compile spike-triggered averages of motoneurone membrane potential. Subsequently, Ia fibre action potentials were cross-correlated with rhythmic discharge of the same motoneurones induced by intracellular injection of current. 2. Primary correlogram peaks were statistically significant for thirty-one of forty-nine single Ia fibre-motoneurone connections. Cumulative sums of correlograms were used to identify the onset and duration of peaks. For twenty cases involving more than 2000 trigger spikes, thirteen showed significant correlogram peaks. For these thirteen, the mean percentage increase (m.p.i.) in motoneurone firing probability, defined as the mean height of the correlogram peak above base line, ranged from 29 to 138%. The k values (maximum height divided by base line) ranged from 2.1 to 5.2. Peak duration varied from 1.8 to 3.2 ms. In the remaining seven cases the Ia e.p.s.p.s produced no significant correlogram peak (i.e. P greater than 0.05). 3. A significant positive relationship (r = 0.76; P less than 0.005) was found between m.p.i. in motoneurone firing probability and e.p.s.p. amplitude (n = 13), with a mean slope of 0.30%/microV. The k values were more weakly related to e.p.s.p. amplitude (r = 0.67; P less than 0.01). The correlogram parameter most strongly related to e.p.s.p. amplitude (r = 0.80) was correlogram peak area (number of spikes above base line per excitatory post-synaptic potential). E.p.s.p. rate of rise was not significantly related (P greater than 0.10) to either m.p.i. in firing probability (r = 0.28) or peak area (r = 0.36). 4. The shapes of the primary correlogram peaks could be accounted for largely by a function proportional to the e.p.s.p. derivative (after temporal alignment). Subtracting a function proportional to the e.p.s.p. derivative from the correlogram peak left either a negligible remainder or a remainder term whose duration was shorter than the e.p.s.p. 5. To investigate properties of single-fibre Ia e.p.s.p.s occurring near motoneurone threshold during repetitive firing, e.p.s.p.s were selectively averaged using Ia spikes occurring near the end of the depolarizing ramp in membrane potential. These 'ramp e.p.s.p.s' tended to be somewhat smaller (by ca.8%) than the 'rest e.p.s.p.s' produced at the same connections with the motoneurone at rest.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

Discharge properties of motoneurons supplying distal forelimb muscles in the cat.

Discharge properties of cat cervical motoneurons innervating distal forelimb muscles were investigated by intracellular current injection. Values for rheobase current, afterhyperpolarization duration and several measures of repetitive discharge characteristics were in most respects similar to those obtained for hindlimb motoneurons.

Action Potentials↗

Soleus motor units in chronic spinal transected cats: physiological and morphological alterations.

Intracellular recording and stimulation were used to study electrical properties of soleus motoneurons and isometric contractile properties of their muscle units. In addition, some muscle units were depleted of glycogen for purposes of studying their morphology. These data were compared for two experimental groups of barbiturate-anesthetized adult cats: those with spinal cords intact (normal) and others with spinal cords transected (ST) 4 mo before study. The greatest change in soleus motoneurons after ST was a decrease in afterhyperpolarization (AHP) duration. Axonal conduction velocity (CV) increased slightly, yet consistently. These data confirm previous findings. The voltage threshold of motoneurons, estimated from the product of rheobase current and input resistance, increased from the normal value of 7.5 mV to 10.1. This resulted from a significant increase in rheobase current after ST, because input resistance was not altered significantly. These data suggest that motoneuron excitability decreased after ST and therefore cannot account for hyperreflexia in chronic ST cats. Measurement of soleus muscle unit isometric contractile properties generally confirmed earlier reports of decreases in twitch time course [both contraction time (CT) and half-relaxation time] and in twitch (Ptw) and maximum tetanic (Pmax) tensions after chronic ST. The ratio Ptw/Pmax increased as a result of a greater decline in tetanic than in twitch tension. No significant change was observed in the fatigue index. Direct measures of muscle unit morphology showed that the specific tension for three ST units fell below the range of four normal units. This was expected given that the innervation ratio was not altered and that the decrease in muscle fiber area after ST was proportionately smaller than the decrease in Pmax. Two lines of evidence showed that motoneuron and muscle unit properties were coordinated before and after ST. First, the normal correlation between AHP duration and CT was nearly as strong after as before ST, and the slope of this relation was not significantly different for normal and ST groups. Second, multivariate statistics (canonical analysis) showed that a combination of all motoneuron electrical properties listed above was strongly correlated with a combination of muscle unit twitch time, Pmax, and Ptw/Pmax for normal and ST groups. In contrast with the conclusions of Gallego et al., we suggest from these data that SOL motor-unit properties are coordinated after ST.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗