Operant conditioning of rat H-reflex affects motoneuron axonal conduction velocity

This study assessed the effects of operant conditioning of the H-reflex on motoneuron axonal conduction velocity in the rat. After measurement of the control H-reflex size, rats were either exposed for at least 40 days to the HRup or HRdown conditioning mode, in which reward occurred only if the soleus H-reflex was greater than (HRup mode) or less than (HRdown mode) a criterion or continued under the control condition (HRcon mode) in which the H-reflex was simply measured. We then measured axonal conduction velocity of triceps surae motor units of HRup, HRdown, and HRcon rats by stimulating the axon in the ventral root and recording from the tibial nerve. Conduction velocity was 8% less in successful HRdown rats than in HRcon rats (P=0.02). Conduction velocity in HRup rats and unsuccessful HRdown rats was not significantly different from that in HRcon rats. Since recording bypassed the intra-spinal portion of the motoneuron, the change was clearly in the axon. This decrease was similar to the 6% decrease previously found in successful HRdown monkeys. Unsuccessful HRdown rats and monkeys did not show this decrease. This result suggests that the mechanism of HRdown conditioning is similar in rats and monkeys and provides further support for the hypothesis that HRdown conditioning decreases motoneuron excitability by producing a positive shift in firing threshold. While traditional theories of learning emphasize synaptic plasticity, neuronal plasticity may also contribute to operantly conditioned behavioral changes.     

Reversal of H-reflex operant conditioning in the rat

In response to an operant conditioning task, rats can gradually increase or decrease soleus H-reflex amplitude without change in background electromyographic activity or M response amplitude. Both increase (under the HRup mode) and decrease (under the HRdown mode) develop over weeks. The present study investigated reversal of conditioned H-reflex change. Following collection of control data, rats were exposed to one mode (HRup or HRdown) for 50 days, and then exposed to the opposite mode for up to 72 days. Rats responded to each mode exposure with gradual, mode-appropriate change in H-reflex amplitude. This finding is consistent with other evidence that H-reflex conditioning depends on spinal cord plasticity. The effects of exposure to the HRup (or HRdown) mode were not affected by whether exposure followed previous exposure to the HRdown (or HRup) mode. In accord with recent studies suggesting that HRup and HRdown conditioning have different spinal mechanisms, these results suggest that reversal of H-reflex change is due primarily to the superimposition of additional plasticity rather than to decay of the plasticity responsible for the initial change.     

Operant conditioning of the primate H-reflex: factors affecting the magnitude of change

Primates can gradually increase or decrease H-reflex amplitude in one leg when reward depends on that amplitude. The magnitude of change varies greatly from animal to animal. This study sought to define the factors that control this magnitude. It evaluated the influence of animal age, muscle size (absolute and relative), background electromyographic activity (EMG) level, M response amplitude, initial H-reflex amplitude, performance intensity, and behavior of the contralateral leg.Fifty-four animals (Macaca nemestrina) underwent operant conditioning of the triceps surae H-reflex in one leg (the trained leg). Twenty-eight were rewarded for larger H-reflexes (HRup animals), and 26 were rewarded for smaller H-reflexes (HRdown animals). In the HRup animals, H-reflex amplitude in the trained leg rose to an average final value of 177% of its initial amplitude. Magnitude of increase varied widely across animals. Nine animals rose to 120–140%, 11 to 160–240%, three to 300% or more, and five remained within 20% of initial amplitude. In the HRdown animals, H-reflex amplitude in the trained leg decreased to an average of 69% of initial amplitude. Magnitude of decrease varied widely. Five animals decreased to 20–40%, seven to 40–60%, six to 60–80%, and eight remained within 20% of initial amplitude.Animal age, as assessed by weight, markedly affected HRdown conditioning, but not HRup conditioning. Heavy HRdown animals (6 kg) were more successful than light HRdown animals (< 6kg). Thirteen of 14 heavy animals and only five of 12 light animals decreased to less than 80% of initial amplitude. One heavy animal and seven light animals remained within 20% of initial amplitude. Established correlations between weight and age indicate that heavy animals were young adults, while many light animals were adolescents. This striking difference in HRdown performance was not attributable to weight-related differences in other factors.Initial H-reflex amplitude varied considerably across animals and affected the magnitude of change. In HRup animals, H-reflex amplitude in the trained leg tended to increase more if initial H-reflex amplitude was small, while in HRdown animals it decreased more if initial amplitude was large. The inter-animal variation in initial H-reflex amplitude was probably largely attributable to variation in Ia afferent excitation by the stimulating electrode pairs and to variation in motoneuron recruitment.Performance intensity, measured as trials per day, had no significant effect on the magnitude of change in either HRup or HRdown animals. Together with available human data, this finding suggests either that the chronic descending influence responsible for the gradual H-reflex change need only be present for a relatively brief period each day, or that it persists between periods of task performance.Final H-reflex amplitude in the control leg varied greatly across animals. It averaged 131% of its initial amplitude in HRup animals and 108% in HRdown animals. Within each group, final control leg amplitude did not correlate with the magnitude of change in the trained leg. Its wide variation and lack of correlation with final amplitude in the trained leg is consistent with evidence that operant conditioning of the H-reflex produces plasticity at multiple spinal and supraspinal sites both ipsilateral and contralateral to the trained leg.     

Sensorimotor cortex ablation prevents H-reflex up-conditioning and causes a paradoxical response to down-conditioning in rats

Operant conditioning of the H-reflex, a simple model for skill acquisition, requires the corticospinal tract (CST) and does not require other major descending pathways. To further explore its mechanisms, we assessed the effects of ablating contralateral sensorimotor cortex (cSMC). In 22 Sprague-Dawley rats, the hindlimb area of left cSMC was ablated. EMG electrodes were implanted in the right soleus muscle and a stimulating cuff was placed around the right posterior tibial nerve. When EMG remained in a specified range, nerve stimulation just above the M response threshold elicited the H-reflex. In control mode, no reward occurred. In conditioning mode, reward occurred if H-reflex size was above (HRup mode) or below (HRdown mode) a criterion value. After exposure to the control mode for > or = 10 days, each rat was exposed for another 50 days to the control mode, the HRup mode, or the HRdown mode. In control and HRup rats, final H-reflex size was not significantly different from initial H-reflex size. In contrast, in HRdown rats, final H-reflex size was significantly increased to an average of 136% of initial size. Thus like recent CST transection, cSMC ablation greatly impaired up-conditioning. However, unlike recent CST transection, cSMC produced a paradoxical response to down-conditioning: the H-reflex actually increased. These results confirm the critical role of cSMC in H-reflex conditioning and suggest that this role extends beyond producing essential CST activity. Its interactions with ipsilateral SMC or other areas contribute to the complex pattern of spinal and supraspinal plasticity that underlies H-reflex conditioning.     

Operant conditioning of primate H-reflex: phases of development

This study sought to determine whether operantly conditioned change in the primate triceps surae (TS) H-reflex develops in distinct phases. Data from 20 animals in which the TS H-reflex in one leg was trained up (i.e., HRup mode) and 18 in which it was trained down (i.e., HRdown mode) were averaged to define H-reflex behavior in trained and control legs. In HRup animals, the trained-leg H-reflex showed a large phase I increase in the first two days followed by gradual phase II increase that continued for weeks. The control-leg H-reflex appeared to show much smaller phase I and phase II increases. In HRdown animals, the trained-leg H-reflex decreased gradually over weeks, while the control-leg H-reflex appeared to increase within 2 days and did not change from then on. The initial rapid increase in the HRdown control leg suggested that two early events occurred in the HRdown trained leg: a nonspecific increase like that in the control leg and an operantly conditioned mode-specific decrease. These two effects may have obscured each other, so that H-reflex size in the HRdown trained leg did not drop rapidly in the first few days. These results improve understanding of adaptive H-reflex change as an operantly conditioned phenomenon, and provide encouragement and direction for efforts to reproduce and study the phenomenon in reduced or anesthetized preparations.     

Synaptic and mechanical coupling between type-identified motor units and individual spindle afferents of medial gastrocnemius muscle of the cat

Experiments were performed to test the possibility that motor unit-muscle spindle pairs that are coupled especially strongly mechanically will also be coupled especially strongly synaptically ("weighted ensemble input": Ref. 4). Synaptic and mechanical coupling between one or two individual muscle spindle afferents and individual motor units of the medial gastrocnemius (MG) muscle were measured in barbiturate-anesthetized cats. Synaptic coupling was assessed by measuring the amplitude of single-fiber monosynaptic excitatory postsynaptic potentials (EPSPs) generated in motoneurons by individual spindle afferents. Mechanical coupling was assessed by measuring the alteration in discharge rate of these spindle afferents caused by tetanic activation of the same motor units. Afferents were classified as primary or secondary on the basis of conduction velocity and response to muscle stretch and contraction. Motor units were classified as slow twitch (S); fast twitch, fatigue resistant (FR); fast twitch, intermediate fatigue resistance (FI); and fast twitch, fatigue sensitive (FF) on the basis of twitch contraction time and resistance to fatigue. In 85% of 138 motor unit-primary afferent interactions tested, tetanic activation of the single motor unit unloaded (i.e., decreased the discharge rate of) the primary afferent. A very weak though significant correlation was found between tetanic contraction strength and primary afferent unloading. In 66% of 155 motor unit-secondary afferent interactions tested, tetanic activation of the single motor unit unloaded the secondary afferent. Again, afferent unloading was but weakly related to tetanic contraction strength. Single-fiber EPSPs generated by primary or secondary muscle spindle afferents were recorded in type-identified motor units. EPSPs generated by primary afferents were significantly larger in oxidative (S + FR) than in glycolytic (FF) motor units. No such differences were seen for EPSPs generated by secondary afferents. The magnitude of the EPSP generated in a motoneuron by a spindle afferent was compared to the magnitude of the unloading of that afferent by tetanic activation of the corresponding motor unit. Overall, no relationship was found between these measures.(ABSTRACT TRUNCATED AT 400 WORDS)     

Whole-muscle and motor-unit contractile properties of the styloglossus muscle in rat.

Investigations of whole muscle and motor-unit contractile properties have provided valuable information for our understanding of the spinal cord and extraocular motor systems. However, no previous investigation has examined these properties in an isolated tongue muscle. The purpose of this study was to determine the contractile properties and muscle fiber types of the rat styloglossus muscle. The styloglossus is one of three extrinsic tongue muscles and serves to retract the tongue within the oral cavity. Adult male Sprague-Dawley rats (n = 19) were used in these experiments. The contractile characteristics of the whole styloglossus muscle (n = 9) were measured in response to stimulation of the hypoglossal nerve branch to the muscle. The average twitch tension produced was 3.30 g with a mean twitch contraction time of 13.81 ms. The mean maximum tetanic tension was 19.66 g and occurred at or near the fusion frequency, which averaged 109 Hz. The styloglossus muscle was resistant to fatigue [fatigue index (F. I.) = 0.76]. In separate experiments (n = 7), the contractile characteristics of 37 single motor units were measured in response to extracellular stimulation of hypoglossal motoneurons. The twitch tension generated by styloglossus motor units averaged 35.7 mg, and the mean twitch contraction time was 12.46 ms. The mean fusion frequency was 92 Hz. Maximum tetanic tension averaged 177.8 mg. Styloglossus single motor units were resistant to fatigue (F. I. = 0.74). The sites of stimulation that yielded a contractile response in the styloglossus muscle were consistent with the location of the styloglossus motoneuron pool reported in earlier anatomy studies. Muscle fiber typing was determined in three animals based on the myofibrillar ATPase reaction at pH 9.8, 4.6, and 4.3. The styloglossus muscle was composed of approximately 99% type IIA fibers with a few scattered type I fibers present in the study sample. On the basis of the combined findings of the physiology and histochemistry experiments, the styloglossus muscle appeared to be a homogeneous muscle composed almost exclusively of fast, fatigue-resistant motor units. These properties of the styloglossus muscle and its motor units were compared with findings in other rat skeletal muscles.     

Progress of age-related changes in properties of motor units in the gastrocnemius muscle of rats

The mechanical properties of individual motor units in the medial gastrocnemius muscle, as well as the whole muscle properties and innervating motor nucleus, were investigated in dietary-restricted, male Fischer 344/DuCrj rats at ages of 4, 7, 12, 21/22, 27, 31, and 36 mo. The tetanic tension of the type S units continuously increased until the age of 36 mo. Those of type FF and FR units declined from 21/22 to 27 mo of age but did not change further while the whole muscle tension decreased greatly. The atrophy of muscle fibers, the decline in motoneuron number and axonal conduction velocity, and the decrease in the posttetanic potentiation of twitch contraction of motor units seemed to start after 21/22 mo of age and were accelerated with advancing age. Prolongation of twitch contraction time was evident for only type S and FR units in 36-mo-old rats. The fatigue index was greatly increased for type FF units in 36-mo-old rats. These findings indicated that the progress of changes in various properties occurring in the senescent muscle was different in terms of their time course and degree and also dependent on the types of motor unit. The atrophy and decrease in specific tension of muscle fibers affected the decline in tension output of motor units. This was effectively compensated for by the capture of denervated muscle fibers over time.     

Properties of self-reinnervated motor units of medial gastrocnemius of cat. II. Axotomized motoneurons and time course of recovery

This study tested the hypothesis that functional connection to muscle is necessary for expression of normal motoneuron electrical properties. Also examined was the time course of self-reinnervation. Properties of individual medial gastrocnemius (MG) motor units were examined following section and reanastomosis of the MG nerve. Stages examined were 3-5 wk (prior to reinnervation, no-re), 5-6 wk (low-re), 9-10 wk (med-re), and 9 mo (long-re, preceding paper) after nerve section. Motor units were classified on the basis of their mechanical response as type fast twitch, fast fatiguing (FF), fast twitch with intermediate fatigue resistance (FI), fast twitch, fatigue resistant (FR), or slow twitch, fatigue resistant (S) (11, 24). Motoneuron electrical properties were measured. Muscle fibers were classified using histochemical methods as type fast glycolytic (FG), fast oxidative glycolytic (FOG), or slow oxidative (SO) (60). Prior to functional reinnervation, MG motoneurons exhibited increased input resistance, decreased rheobase, decreased rheobase/input resistance, and decreased axonal conduction velocity. There was no change in mean afterhyperpolarization (AHP) half-decay time. Normal relationships between motoneuron electrical properties were lost. These data are consistent with dedifferentiation of motoneuron properties following axotomy (35, 47). At 5-6 wk after reanastomosis, motor-unit tensions were small, and motoneuron membrane electrical properties were unchanged from the no-re stage. There were no differences in motoneuron electrical properties between cells that elicited muscle contraction and those that did not. Motor-unit types were first recognizable at the med-re stage. The proportions of fast and slow motor units were similar to normal MG. Within the fast units, there were fewer type-FF units and more type-FI and type-FR units than normal, reflecting a general increase in fatigue resistance at this stage. Neither motoneuron membrane electrical properties nor muscle contractile properties had reached normal values, although both were changed in that direction from the low-re stage. Normal relationships between muscle properties, between motoneuron properties, and between motoneuron and muscle properties were re-established. The correspondence between motor-unit type and motoneuron type was similar to normal or 9 mo reinnervated MG. Muscle-unit tetanic tensions became larger with time after reinnervation. Most of the increase in muscle tension beyond the med-re stage could be accounted for by increase in muscle fiber area. There was an increased proportion of SO muscle fibers observed in the med-re muscles, as at the long-re stage.(ABSTRACT TRUNCATED AT 400 WORDS)     

Changes in contractile properties of motor units of the rat medial gastrocnemius muscle after spinal cord transection

The effects of complete transection of the spinal cord at the level of Th9/10 on contractile properties of the motor units (MUs) in the rat medial gastrocnemius (MG) muscle were investigated. Our results indicate that 1 month after injury the contraction time (time-to-peak) and half-relaxation time were prolonged and the maximal tetanic force in most of the MUs in the MG muscle of spinal rats was reduced. The resistance to fatigue also decreased in most of the MUs in the MG of spinal animals. Moreover, the post-tetanic potentiation of twitches in MUs diminished after spinal cord transection. Criteria for the division of MUs into three types, namely slow (S), fast fatigue resistant (FR) and fast fatigable (FF), applied in intact animals, could not be directly used in spinal animals owing to changes in contractile properties of MUs. The 'sag' phenomenon observed in unfused tetani of fast units in intact animals essentially disappeared in spinal rats and it was only detected in few units, at low frequencies of stimulation only. Therefore, the MUs in spinal rats were classified as fast or slow on the basis of an adjusted borderline of 20 ms, instead of 18 ms as in intact animals, owing to a slightly longer contraction time of those fast motor units with the 'sag'. We conclude that all basic contractile properties of rat motor units in the medial gastrocnemius muscle are significantly changed 1 month after complete spinal cord transection, with the majority of motor units being more fatigable and slower than those of intact rats.     

Properties of self-reinnervated motor units of medial gastrocnemius of cat. I. Long-term reinnervation

This work tested whether the membrane electrical properties of cat motoneurons, the contractile properties of their muscle units, and the normal relationships among them would be restored 9 mo after section and resuture of their muscle nerve. Properties of medial gastrocnemius (MG) motor units were examined 9 mo following section and resuture of the MG nerve in adult cats. Motoneuron electrical properties and muscle-unit contractile properties were measured. Motor units were classified on the basis of their contractile properties as type fast twitch, fast fatiguing (FF), fast twitch with intermediate fatigue resistance (FI), fast twitch, fatigue resistant (FR), or slow twitch, fatigue resistant (S) (8, 20). Muscle fibers were classified as type fast glycolytic (FG), fast oxidative glycolytic (FOG), or slow oxidative (SO) on the basis of histochemical staining for myosin adenosine triphosphatase, nicotinamide adenine dinucleotide diaphorase, and alpha-glycerophosphate dehydrogenase (48). Following 9 mo self-reinnervation, the proportions of each motor-unit type were the same as in normal control animals. Motoneuron membrane electrical properties [axonal conduction velocity, afterhyperpolarization (AHP) half-decay time, rheobase, and input resistance] also returned to control levels in those motoneurons that made functional reconnection with the muscle (as determined by ability to elicit measurable tension). The relationships among motoneuron electrical properties were normal in motoneurons making functional reconnection. Approximately 10% of MG motoneurons sampled did not elicit muscle contraction. These cells' membrane electrical properties were different from those that did elicit muscle contraction. Contractile speed and fatigue resistance of reinnervated muscle units had recovered to control levels at 9 mo postoperation. Force generation did not recover fully in type-FF units. The reduced tensions were apparently due to failure of recovery of FG muscle fiber area. Following reinnervation, relationships between motoneuron electrical and muscle-unit contractile properties were similar to controls. This was reflected in a degree of correspondence between motor-unit type and motoneuron type similar to normal units (84 vs. 86%, as defined by Ref. 61). There was a significantly increased proportion of type-SO muscle fibers and a decrease in the fast muscle fibers (especially type FOG) in 9 mo reinnervated MG. Together with the unchanged proportions of motor-unit types, this led to an estimate of average innervation ratios being increased in type-S motor units and decreased in type-FR units.(ABSTRACT TRUNCATED AT 400 WORDS)     

Sag during unfused tetanic contractions in rat triceps surae motor units.

Contractile properties and conduction velocity were studied in 202 single motor units of intact rat triceps surae muscles activated by intra-axonal (or intra-myelin) current injection in L5 or L6 ventral root to assess the factors that determine the expression of sag (i.e., decline in force after initial increase during unfused tetanic stimulation). Sag was consistently detected in motor units with unpotentiated twitch contraction times <20 ms. However, the range of frequencies at which sag was expressed varied among motor units such that there was no single interstimulus interval (ISI), with or without adjusting for twitch contraction time, at which sag could be detected reliably. Further analysis indicated that using the absence of sag as a criterion for identifying slow-twitch motor units requires testing with tetani at several different ISIs. In motor units with sag, the shape of the force profile varied with tetanic frequency and contractile properties. Simple sag force profiles (single maximum reached late in the tetanus followed by monotonic decay) tended to occur at shorter ISIs and were observed more frequently in fatigue-resistant motor units with long half-relaxation times and small twitch amplitudes. Complex sag profiles reached an initial maximum early in the tetanus, tended to occur at longer ISIs, and were more common in fatigue-sensitive motor units with long half-relaxation times and large twitch amplitudes. The differences in frequency dependence and force maximum location suggested that these phenomena represented discrete entities. Successive stimuli elicited near-linear increments in force during tetani in motor units that never exhibited sag. In motor units with at least one tetanus displaying sag, tetanic stimulation elicited large initial force increments followed by rapidly decreasing force increments. That the latter force envelope pattern occurred in these units even in tetani without sag suggested that the factors responsible for sag were expressed in the absence of overt sag. The time-to-peak force (TTP) of the individual contractions during a tetanus decreased in tetani with sag. Differences in the pattern of TTP change during a tetanus were consistent with the differences in force maximum location between tetani exhibiting simple and complex sag. Tetani from motor units that never exhibited sag did not display a net decrease in TTP during successive contractions. These data were consistent with the initial force decrement of sag resulting from a transient reduction in the duration of the contractile state.     

Modulation of motor unit discharge rate and H-reflex amplitude during submaximal fatigue of the human soleus muscle

Declining motor unit discharge rates and H-reflex amplitude have been observed in separate experiments during fatiguing submaximal contractions in humans. The purpose of this experiment was to investigate motor unit discharge rate, H-reflex amplitude, and twitch contractile properties concurrently during a fatiguing submaximal isometric contraction of the ankle plantarflexors. Eleven healthy subjects performed fatiguing contractions of low force (25% maximal voluntary contraction (MVC)) or high force (42–66% MVC). Hoffmann (H)-reflexes, muscle compound action potentials (M-waves), twitch contractile properties, and motor unit discharges were recorded from the soleus muscle. In the low-force fatigue task, motor unit firing rate increased gradually over time, whereas the resting H-reflex was significantly depressed at 15% of endurance time and remained quasiconstant for the rest of the task. This suggests that the processes mediating the resting H-reflex depression are relatively independent of those modulating the motor unit firing rate during a low-force fatigue task. In the high-force fatigue task, a decline in the average motor unit discharge rate was accompanied by a decrease in the resting H-reflex amplitude and a prolongation of the twitch half-relaxation time (HRT) at the completion of the fatigue task. Overall, motor unit firing rate was modulated in parallel with changes in the twitch HRT, consistent with the muscle wisdom hypothesis.     

Comparison of the histochemical and contractile properties of human triceps surae

Summary Contractile and histochemical properties of the triceps surae were compared in 16 males and 4 females aged 20 to 49 years. Surface electrical stimulation was used to determine twitch, tetanic and fatigue parameters. From these tests, twitch tension (Pt), time to peak tension (TPT), half relaxation time 1/2 RT), tetanic tensions at 10, 20 and 50 Hz and an index of fatigue (FI) were calculated. A maximal voluntary contraction (MVC) was also performed. Muscle samples from the belly of the lateral gastrocnemius were obtained using the needle biopsy technique. The samples were treated histochemically for myosin ATPase and NADH-tetrazolium reductase in order to classify the fibres as either Type I, slow twitch (ST) or Type II, fast twitch (FT) and to determine fibre areas. Correlations were performed between the grouped male and female contractile and histochemical variables. The results demonstrated significant positive relationships between percentage of ST fibres (%ST) and TPT (r=0.49), and %ST and the ratio of tetanic forces at 10 Hz to 50 Hz (Po10/Po50) (r=0.55). No significant relationships were obtained for Pt, 1/2 RT, MVC or FI with any histochemical parameter. The results suggest that fibre type distribution determined using myosin ATPase is related to electrically stimulated isometric contractile speeds and not to voluntary force generation (MVC) or electrically induced fatigue.     

Presynaptic inhibition, EPSP amplitude, and motor-unit type in triceps surae motoneurons in the cat

1. Composite group Ia excitatory postsynaptic potentials (EPSPs) produced by heteronymous nerve stimulation were recorded from triceps surae motoneurons of barbiturate-anesthetized cats. Motoneuron rheobase, input resistance, and axonal conduction velocity were measured, and motor units were classified on the basis of the mechanical responses of their muscle units. 2. The amplitude of EPSPs recorded from 33 medial gastrocnemius (MG) motoneurons ranged from 0.6 to 4.3 mV. The mean EPSP amplitude differed among the major MG motor-unit types, increasing in the order fast twitch, fast fatiguing (FF); fast twitch, fatigue resistant (FR); slow twitch, fatigue resistant (S) (FF less than FR less than S). The amplitude of EPSPs recorded from 15 soleus motoneurons ranged from 0.3 to 3.4 mV, with a mean of 1.4 mV. 3. Presynaptic inhibition of EPSPs was produced by trains of conditioning volleys in the posterior biceps-semitendinosus (PBST) nerve. In 33 MG cells PBST conditioning stimulation reduced the amplitude of EPSPs by 11-50%, with a mean inhibition of 27%. The amplitude of EPSPs in 15 soleus motoneurons was decreased by 5-84%, with a mean inhibition of 37%. 4. When the magnitude of presynaptic inhibition was expressed as percent inhibition, there was no relation between presynaptic inhibition and either motor-unit type or the amplitude of the EPSP. However, when presynaptic inhibition was expressed as the absolute amount of inhibition in millivolts, the magnitude of inhibition was highly correlated with EPSP amplitude both across the entire triceps surae population (MG, lateral gastrocnemius, soleus) as well as within each muscle population. This correlation was also significant within the MG FF and FR motor-unit populations. 5. We conclude that EPSP amplitude and not motor-unit type is the major determinant of the magnitude of presynaptic inhibition. However, because of the effect of motor-unit type on EPSP amplitude, the net effect is that presynaptic inhibition increases in the order FF less than FR less than S.     

Twitch and tetanic properties of human thenar motor units paralyzed by chronic spinal cord injury

Little is known about how human motor units respond to chronic paralysis. Our aim was to record surface electromyographic (EMG) signals, twitch forces, and tetanic forces from paralyzed motor units in the thenar muscles of individuals (n = 12) with chronic (1.5-19 yr) cervical spinal cord injury (SCI). Each motor unit was activated by intraneural stimulation of its motor axon using single pulses and trains of pulses at frequencies between 5 and 100 Hz. Paralyzed motor units (n = 48) had small EMGs and weak tetanic forces (n = 32 units) but strong twitch forces, resulting in half-maximal force being achieved at a median of only 8 Hz. The distributions for cumulative twitch and tetanic forces also separated less for paralyzed units than for control units, indicating that increases in stimulation frequency made a smaller relative contribution to the total force output in paralyzed muscles. Paralysis also induced slowing of conduction velocities, twitch contraction times and EMG durations. However, the elevated ratios between the twitch and the tetanic forces, but not contractile speed, correlated significantly with the extent to which unit force summated in response to different frequencies of stimulation. Despite changes in the absolute values of many electrical and mechanical properties of paralyzed motor units, most of the distributions shifted uniformly relative to those of thenar units obtained from control subjects. Thus human thenar muscles paralyzed by SCI retain a population of motor units with heterogeneous contractile properties because chronic paralysis influenced all of the motor units similarly.     

Effect of force level and training status on contractile properties following fatigue.

This study examined the effects of fatigue on the contractile properties of the twitch contraction evoked by the H-reflex and the maximal M-wave. Untrained subjects demonstrated more pronounced slowing of half relaxation time (HRT) in the H-reflex twitch than in the M-wave twitch and also more slowing of HRT in high force vs. low force contractions. Endurance-trained subjects, however, demonstrated a shortening of the H-reflex twitch. The twitch contractile properties following fatigue are dependent on force level and training status.     

Memory traces in spinal cord produced by H-reflex conditioning: effects of post-tetanic potentiation

Operant conditioning of the wholly spinal, largely monosynaptic triceps surae H-reflex in monkeys causes changes in lumbosacral spinal cord that persist after removal of supraspinal influence. We evaluated the interaction between post-tetanic potentiation and these memory traces. Animals in which the triceps surae H-reflex in one leg had been increased or decreased by conditioning were deeply anesthetized, and monosynaptic reflexes to L6-S1 dorsal root stimulation were recorded before and after tetanization from both legs for 3 days after thoracic cord transection. Animals remained anesthetized throughout and were sacrificed by overdose. Reflex asymmetries consistent with the effect of H-reflex conditioning were present after transection and persisted through the 3 days of study. Tetanization affected conditioned leg and control leg reflexes similarly. This finding suggests that, while post-tetanic potentiation and probably H-reflex conditioning alter Ia synaptic transmission, the two phenomena have different mechanisms.     

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

1. Post-synaptic potentials (PSPs) evoked by electrical stimulation of a variety of input systems have been compared in triceps surae motoneurones innervating slow and fast muscle units, the speed of contraction of which was also determined.2. Stimulation of high threshold afferents in both flexor and extensor muscle nerves, and of joint afferents, evoked polysynaptic PSPs which were predominantly hyperpolarizing in both fast and slow twitch motor units.3. Volleys in cutaneous afferents in the sural and saphenous nerves evoked polysynaptic PSPs composed of mixtures of inhibitory and excitatory components. The inhibitory components were predominant in slow twitch motor units, while in fast twitch units there was a trend towards excitatory predominance.4. Repetitive stimulation of the red nucleus caused predominantly inhibitory PSPs in slow twitch units and mixed or predominantly excitatory PSPs in fast twitch units. There was a correlation in the excitatory/inhibitory balance between PSPs of cutaneous and rubrospinal origin in those motoneurones in which both types of PSPs were studied.5. The amplitudes of group Ia disynaptic inhibitory PSPs were found to be correlated with motor unit twitch type: IPSPs in slow twitch units were larger than those in fast twitch units. Rubrospinal conditioning volleys were found to facilitate group Ia IPSPs in both fast and slow twitch motor units.6. The results suggest that there may be several basic patterns of synaptic input organization to motoneurones within a given motor unit pool. In addition to quantitative variation in synaptic distribution, there is evidence that qualitative differences in excitatory to inhibitory balance also exist in the pathways conveying input from cutaneous afferents and rubrospinal systems to triceps surae motoneurones. These qualitative differences are correlated with the motor unit twitch type.     

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)