Mandibular postural tremor during relaxation and during volitional innervation of the jaw muscles

Jaw tremor was recorded as the changes in the vertical distance between mandible and maxilla. Two states of motor performance of the jaw muscles were chosen: relaxed and actively positioning of the mandible close to toothcontact, referred to as respectively rest- and activity tremor.Rest tremor contained a strong ballistocardiac component (bc power = 50%) and was of small amplitude (9.4 m rms). Activity tremor revealed increased amplitudes (19.2 m rms) with a predominant neuromuscular component (nm power = 75%). Bc tremor patterns differed significantly from nm tremor patterns. A negative linear relationship, which apparently was similar for rest and for activity tremor (P>0.10), existed between the estimated relative bc contribution and tremor amplitude. As tremor amplitude increased neuromuscular activity appeared increasingly time locked to bc activity, which should be attributed to synchronization of spindle activity, either through the stretch reflex or through vascular pulsation. The average bc contribution to tremor amplitude remained of almost constant magnitude (7.0 m rms), with no differences between rest and activity for the group as a whole. During resting conditions, complete relaxation of the jaw muscles was not attained by the majority of the subjects. The latter revealed a heart rate arrhythmia that was strongly correlated with motor (nm) activity (P<0.001). During activity conditions no such correlation was present.The present results demonstrate that for the understanding and the interpretation of the underlying mechanisms of jaw tremor genesis, both the recording technique, i.e., differential jaw displacements, and the signal analysis procedure, i.e., autocorrelograms and power spectra over extended record length, were of crucial importance. A clear distinction was made between cardioballistic tremors and tremors related to muscular activity. Heart rate arrhythmia and reflex activity could be demonstrated as well.     

Control of human jaw elevator muscle activity during simulated chewing with varying bolus size

During chewing, a small part of the observed muscle activity is needed for the basic open-close movements of the mandible, and additional muscle activity (AMA) is needed to overcome the resistance of the food. The AMA consists of two contributions: a large peripherally induced contribution, starting after food contact and a small anticipating contribution, starting before food contact. We investigated whether the latencies of these contributions depend on the expected or actual bolus size.Subjects made rhythmic open-close movements near their natural chewing frequency controlled by a metronome. This frequency was determined while the subjects were chewing gum. Food resistance was simulated by an external force, acting on the jaw in a downward direction during part of the closing movement. Bolus size was simulated by the jaw gape at which the force started. Jaw movement and surface EMG of the masseter and anterior temporal muscles on both sides and the suprahyoid muscles were recorded during experiments in which the jaw gape at which the force started was varied. The peripherally induced contribution to the AMA started about 20 ms after the onset of the force, irrespective of the jaw gape at which the force started. It is concluded thatthe onset of this contribution depends solely on food contact in the actual cycle. The function of the observed mechanism for jaw elevator muscle control may be to enable a highly automatic control of the muscle activity required to overcome the resistance of food of different hardness and different size.The onset of the anticipating contribution to the AMA showed neither a relationship with the actual jaw gape at which force onset occurred nor with the expected jaw gape of force onset. It is suggested that the onset of the anticipating AMA is related to the jaw gape at the onset of closing. The function of this contribution may be the regulation of the mechanical response of the jaw after an expected disturbance of the closing movement by food contact, by tuning the muscle stiffness to the expected hardness of the food.     

Jaw stretch reflexes in children

The substantial morphological transformations that occur during human development present the nervous system with a considerable challenge in terms of motor control. Variability of skilled motor performance is a hallmark of a developing system. In adults, the jaw stretch reflex contributes to the functional stability of the jaw. We have investigated the response properties of the jaw stretch reflex in two groups of young children and a group of young adults. Response latencies increased with development, and all age groups produced stimulus-magnitude-dependent increases in reflex gain and resulting biting force. Reflex gain was largest for the older children (9–10 years), yet net increases in resulting biting force were comparable across age groups. These data and earlier experiments suggest that oral sensorimotor pathways mature throughout childhood in concert with the continued acquisition of complex motor skills.     

Limitation of jaw movement by antagonist muscle stiffness during unloading of human jaw closing muscles

Summary The unloading reflex in the jaw closing muscles in man was investigated with a view to correlating the jaw closing movement with the timing of the electrical activity of the agonist and antagonist muscles. When the resistance to a forceful isometric bite was suddenly and unexpectedly withdrawn, the closing movement of the mandible was always arrested before the teeth came together. The rapid arrest of the jaw closing movement was not adequately accounted for by the timing of the inhibition of the jaw closing muscles and reflex excitation of the jaw opening muscles. It was observed that the jaw opening muscles, as well as the jaw closing muscles, were always active during the phase of isometric biting on an object between the teeth. It is therefore concluded that the resulting stiffness of the antagonist muscles is the mechanism which is principally responsible for limiting the jaw closing movement after unloading of the agonist muscles.Supported by the National Health and Medical Research Council of Australia, and the Australian Dental Education and Research Trust     

Rotation of synergistic activity during isometric jaw closing muscle contraction in man

5 healthy subjects were studied during 10–15 min of isometric jaw elevator contraction above fatigue threshold level. Bite force was measured between upper and lower front teeth and electromyographic (EMG) activity recorded from the right temporal and masseter muscles. Of the two muscles only the masseter was active at the start of the test and usually during the whole test. When the test was repeated, however, great relief from the pain in the fatigued masseter was sometimes experienced and the temporalis took over the load. All subjects experienced this “switch” phenomenon after a varying number of tests, one of them already during his first test. The mechanism seemed to be completely out of voluntary control and showed facilitation at repeated tests.     

Effects of local and remote muscle pain on human jaw reflexes evoked by fast stretches at different clenching levels

Muscle pain imposes significant changes on natural motor tasks, but the consequences for stretch reflexes are still disputed. The present study examined the jaw reflexes to fast (10 ms) stretches of the mandible in an experimental model with local pain in the masseter muscle and remote pain in the tibialis anterior muscle. The stretch reflexes were elicited in healthy volunteers (n=13) before, during, and after periods with constant levels of experimental pain and while the subjects clenched at 0%, 15%, 30%, and 45% of the maximal voluntary contraction (MVC) levels. Surface electromyography (EMG) was used to record the reflex responses. Pain in the masseter muscle (mean ± SEM, 3.8±0.4 on a 10-cm visual analogue scale), but not in the tibialis anterior muscle (3.4±0.3; paired t-test, P=0.318) was associated with significant changes in both prestimulus EMG activity (ANOVA, P=0.002) and in peak-to-peak amplitudes of the stretch reflex (ANOVA, P=0.022). However, when the changes in prestimulus EMG activity were taken into consideration a significant increase in the stretch reflex persisted in the painful muscle at 15% and 30% MVC. Local circuits at the trigeminal level involving the fusimotor system are proposed to mediate a significant part of this modulatory effect. Electronic Publication     

Modulation of H reflexes in the forearm during voluntary teeth clenching in humans

We investigated whether there is any modulation of the H reflex in the forearm during teeth clenching and how any correlation that may be found is modulated. The H reflexes of the flexor carpi radialis (FCR) and the extensor carpi radialis (ECR) muscles were evoked on the right side in five healthy adult volunteers. The H reflexes of the FCR and ECR muscles were facilitated in association with voluntary teeth clenching in a force-dependent manner (r=0.46–0.663, P<0.05). The increase in amplitude of the H reflex of the FCR muscle associated with teeth clenching started before the onset of the EMG activity of the masseter muscle. The results of the present study demonstrate that oral motor activity exerts strong influences on the motor activity of the forearm.A part of this study was presented at the World Congress on Sports Dentistry and Dental Traumatology, 22 June 2001, Boston, Massachusetts, USA     

Wipe and flexion withdrawal reflexes display different EMG patterns prior to movement onset in the spinalized frog

Summary We investigated the hypotheses (1) that the initial flexion part of the wipe reflex elicited in the spinalized frog has the same EMG pattern for wipes to different target locations (Berkinblit et al. 1986), thereby reducing the complexity of the control of this task, and (2) that this initial flexion is the same as occurs in the flexion withdrawal reflex (Easton 1972). The activities of seven muscles of the hindlimb of the spinal frog were recorded via intramuscular electromyograms (EMGs) during the wipe reflex to two target locations and during the flexion withdrawal reflex. The EMGs were analyzed during the interval between stimulus placement and movement onset for mean integrated EMG and duration from EMG onset to movement onset. This analysis revealed significant differences (p<0.0001) in the EMG patterns that preceded the initial flexion posture for all three movements. These findings suggest that the spinal circuitry coordinating the initial flexion part of the wipe reflex to different target locations and the flexion withdrawal reflex may not be uniformly shared.     

Reflex responses of motor units in human masseter muscle to mechanical stimulation of a tooth

The reflex responses evoked by controlled mechanical stimulation of an upper central incisor tooth in single motor units in the human masseter muscle were examined. The stimuli were (brisk) taps and (slow) pushes of about 2 N peak force, applied orthogonally to the labial surface of the ipsilateral upper central incisor tooth. The reflex responses of the motor units were characterised by analysis of the changes in the durations of the first and second interspike intervals (ISIs) immediately following the stimulus. A significant increase in the duration of these ISIs in comparison with pre-stimulus ISIs indicated inhibition, and significant shortening indicated excitation. Twenty masseter motor units were tested with both the pushes and the taps. The brisk taps elicited a significant reflex inhibition in 16 of the 20 motor units at a latency of 13 ms and duration of 37 ms. This inhibition was followed by significant excitation in 11 of the 20 units at latencies of 71 ms, lasting for 29 ms. The short-latency response to slow pushes was significant inhibition in four units: significant excitation in one unit and no response in 15 units. The slow pushes evoked a significant long-latency excitatory reflex response in 12 of the 20 units at latencies of 77 ms and lasting for 40 ms. The shapes and amplitudes of the compound post-synaptic potentials underlying the reflex responses in the motoneurones were estimated. It is concluded that stimulation of periodontal mechanoreceptors usually activates an excitatory reflex pathway to the jaw-closing motoneurones. This probably helps to grip the food bolus between the teeth during chewing. However, when the rate of application of the stimulus is large enough, a short-latency inhibitory response is evoked which, if of sufficient duration, may over-ride the subsequent excitatory response. Inhibition of the jaw-closing muscles will tend to protect the teeth and soft tissues when one bites unexpectedly on a hard object while chewing.     

The effect of electrical stimulation in the hypothalamus on the monosynaptic jaw closing and the disynaptic jaw opening reflexes in the cat

Summary The effects of electrical stimulation in the hypothalamus on the monosynaptic jaw closing and the disynaptic jaw opening reflexes were investigated in cats anaesthetized with chloralose. The hypothalamic electrodes were located by observation of behavioural attack responses in the unanaesthetized animal and by means of Horsley-Clarke coordinates. The locations were verified in histological serial sections.Hypothalamic conditioning with trains of 3–10 pulses, 0.5 ms duration, 0.5 mA, 500 Hz, evoked a strong facilitation of the jaw closing reflex and a facilitation followed by an inhibition of the jaw opening reflex. These effects differed from those elicited from the cerebral cortex. The hypothalamic effects had a longer latency (11–13 ms) and required a longer train of conditioning stimuli than was the case with those evoked from the cortex. Bilateral ablation of the sensorimotor cerebral cortex or lesion of the pyramids at the lower pontine level diminished but did not abolish the hypothalamic effects. They did, however, disappear after lesions including the ventral midbrain tegmentum.The stimulus positions eliciting the largest hypothalamic effects on the jaw reflexes were located in a region extending medio-laterally from the perifornical area to the entrance of the ansa lenticularis in the lateral hypothalamus. Rostro-caudally the location was found at the level of the ventromedial hypothalamic nucleus and the anterior hypothalamus just rostral to this nucleus. The region corresponds to those parts of the hypothalamus from which agonistic and feeding responses have been evoked.It is suggested that the observed hypothalamotrigeminal mechanism may exercise a tonic influence on the trigeminal motoneurones, thereby controlling the set point of the biting force. The implications of this hypothesis on the etiology of bruxism and the myofascial pain dysfunction are discussed.This work was supported by the Swedish Medical Research Council (Proj B80-14X-9945-16A) and by Gunvor and Josef Anérs Stiftelse     

Sensory components facilitating jaw-closing muscle activities in the rabbit

Summary The role of oral and facial sensory receptors in the control of masticatory muscle activities was assessed from the effect of acute deafferentiation on cortically induced rhythmic jaw movements (CRJMs) in anesthetized rabbits. When a thin polyurethane-foam strip (1.5, 2.5 or 3.5 mm thick) was placed between opposing molars during CRJMs, masseteric activities were facilitated in association with an increase in the medial excursion of the mandible during the power phase. The effects varied with the pattern of CRJMs, and the rate of facilitation was greater for small circular movements than for the crescent-shaped movements. Furthermore, the response of the masseter muscle was greater in the anterior half of the muscle, where muscle spindles are most dense, than in its posterior half. It was also demonstrated that the response increased with an increase in the thickness of the test strip. In contrast, the activities of the jaw-opening muscle were not affected significantly. The duration of masseteric bursts increased during application of the test strip and the chewing rhythm tended to slow down. However, the latter effect was not significant. After locally anesthetizing the maxillary and inferior alveolar nerves, the facultative responses of the masseter muscle to the test strip was greatly reduced but not completely abolished. Lesioning of the mesencephalic trigeminal nucleus (Mes V) where the primary ganglion cells of muscle spindle afferents from jaw-closing muscles and some periodontal afferents are located, also reduced the facilitative effects. Similar results were obtained in the animals with the kainic acid injections into the Mes V 1 week before electrical lesioning of this nucleus. In these animals the effects of electrical lesioning of the Mes V could be attributed to the loss of muscle receptor afferents since the neurons in the vicinity of the Mes V were destroyed and replaced by glial cells, whereas the Mes V neurons are resistant to kainic acid. When electrical lesioning of the Mes V and sectioning of the maxillary and inferior alveolar nerves were combined in animals with a kainic acid injection into the Mes V, the response of the masseter muscle to application of the strip was almost completely abolished. From these findings, we conclude that both periodontal receptors and muscle spindles are primarily responsible for the facilitation of jaw-closing muscle activities. Furthermore, it is suggested that the transcortical loop may not be the only path producing this facilitation since similar effects were induced in animals with ablation of the cortical masticatory area (CMA), when the test strip was placed between the molars during rhythmic jaw movements induced by pyramidal tract stimulation.     

Exteroceptive suppression of temporalis and masseter muscle activity is enhanced in offspring of hypertensives

Exteroceptive suppression of temporalis and masseter muscle activity was examined in young men with and without a parental history of hypertension. Recent clinical studies suggest that the second exteroceptive suppression period is attenuated in several chronic pain disorders and that this brainstem reflex may serve as a noninvasive index of endogenous pain control. In the present study, offspring of hypertensives exhibited a significant protraction of the late exteroceptive suppression period for both muscle sites, suggesting that the decreased pain sensitivity previously observed in individuals at risk for hypertension may be related to enhanced central pain modulation.     

Fusimotor effects of midbrain stimulation on jaw muscle spindles of the anaesthetized cat

The effects of electrical stimulation within the midbrain on fusimotor output to the jaw elevator muscles were studied in anaesthetized cats. Muscle spindle afferents recorded in the mesencephalic trigeminal nucleus were categorised as primary or secondary by their responses to succinylcholine during sinusoidal or rampand-hold stretches. Changes in their stretch responses during midbrain stimulation were then assessed by changes in bias and in dynamic sensitivity. Problems were encountered in interpreting changes in sine wave stretch responses of primary afferents, in some of which a very small change in firing pattern produced large changes in estimates of the reponse amplitude. Sine wave testing also sometimes over-estimated static effects and under-estimated dynamic effects relative to ramp responses. On other occasions a small amount of static fusimotor activity caused a marked increase in sine response amplitude, which could be wrongly interpreted as a dynamic effect. Consequently, ramp responses only were used for diagnosing fusimotor changes. The most effective region for producing pure dynamic fusimotor excitation was directly rostral to the red nucleus, extending dorsally and ventrally approximately in the course of the retroflex bundle. Stimulation of regions caudal and dorso-caudal to the red nucleus, previously designated as the mesencephalic area for dynamic fusimotor control of leg muscles, gave static or mixed static and dynamic effects on jaw spindles. The use of midbrain stimulation to identify fusimotor neurones of jaw muscles as static or dynamic would be most reliable with stimulation just rostral to the red nucleus and would require spindle afferent behaviour to be monitored at the same time with ramp stretches.     

Stretch reflexes in human abdominal muscles

Homonymous and heteronymous reflex connections of the abdominal muscles were investigated by the application of a tap to the muscle belly and observation of surface electromyographic responses. Reflex responses of the following abdominal muscles were investigated both ipsilateral and contralateral to the tap: rectus abdominis (RA), external oblique (EO) and internal oblique (IO). Reflexes were evoked in each of the homonymous muscles with latencies and estimated conduction velocities compatible with being evoked by Ia muscle afferents and having a monosynaptic component. Short latency heteronymous excitatory reflex connections were also observed in muscles on both ipsilateral and contralateral sides in response to the same stimulus. The latencies of the crossed responses were only marginally longer than responses evoked in the respective ipsilateral muscle. Moreover, the reflexes evoked in the IO muscle from ipsilateral and contralateral IO muscle afferents were of comparable amplitude, as were those reflexes evoked in ipsilateral and contralateral EO and RA muscles when tapping IO. These similarities in the reflex characteristics on the ipsilateral and contralateral sides suggest that abdominal muscle afferents activate similar pathways to muscles on both sides of the body. It follows that if the homonymous stretch reflex of abdominal muscles have a monosynaptic component, then a similar monosynaptic pathway activates synergistic motoneurones, not only ipsilaterally but also contralaterally.     

Sequential activation of axial muscles during different forms of rhythmic behavior in man

In humans, studies of back muscle activity have mainly addressed the functioning of lumbar muscles during postural adjustments or rhythmic activity, including locomotor tasks. The present study investigated how back muscles are activated along the spine during rhythmical activities in order to gain insights into spinal neuronal organization. Electromyographic recordings of back muscles were performed at various trunk levels, and changes occurring in burst amplitudes and phase relationships were analyzed. Subjects performed several rhythmic behaviors: forward walking (FW), backward walking (BW), amble walking (where the subjects moved their arms in phase with the ipsilateral leg), walking on hands and knees (HK) and walking on hands with the knees on the edge of a treadmill (Hand). In a final task, the subjects were standing and were asked to swing (Swing) only their arms as if they were walking. It was found that axial trunk muscles are sequentially activated by a motor command running along the spinal cord (which we term “motor waves”) during various types of locomotion or other rhythmic motor tasks. The bursting pattern recorded under these conditions can be classified into three categories: (1) double-burst rhythmic activity in a descending (i.e., with a rostro-caudal propagation) motor wave during FW, BW and HK conditions; (2) double-burst rhythmic activity with a stationary motor wave (i.e., occurring in a single phase along the trunk) during the ‘amble’ walk condition; (3) monophasic rhythmic activity with an ascending (i.e., with a caudo-rostral propagation) motor wave during the Swing and Hands conditions. Our results suggest that the networks responsible for the axial propagation of motor activity during locomotion may correspond to those observed in invertebrates or lower vertebrates, and thus may have been partly phylogenetically conserved. Such an organization could support the dynamic control of posture by ensuring fluent movement during locomotion.     

Effects of experimental pain on jaw muscle activity during goal-directed jaw movements in humans

To study the effects of masseter muscle pain on jaw muscle electromyographic (EMG) activity during goal-directed tasks. Mandibular movement was tracked and EMG activity was recorded from bilateral masseter, and right posterior temporalis, anterior digastric, and inferior head of lateral pterygoid muscles in 22 asymptomatic subjects at postural jaw position, and during three tasks: (a) protrusion, (b) contralateral (left), (c) open jaw movement. Tasks were performed during three conditions: control (no infusion), test 1 [continuous infusion into right masseter of 4.5% hypertonic saline to achieve 30–60 mm pain intensity on 100-mm visual analog scale (VAS)], and test 2 (isotonic saline infusion; in 16 subjects only); the sequence of hypertonic and isotonic saline was randomized. The average EMG root-mean-square values at 0.5 mm increments of mid-incisor-point displacement were analysed using linear mixed effects model statistics (significance: P < 0.05). Right masseter hypertonic saline infusion resulted in significantly (P < 0.0005) more pain (mean ± SD VAS 47.3 ± 14.3 mm) than isotonic infusion (12.2 ± 17.3 mm). Although there was evidence of inter-subject variation, the principal EMG findings were that the significant effects of hypertonic saline-induced pain on EMG activity varied with the task in which the muscle participated irrespective of whether the muscle was an agonist or an antagonist in the tasks. The direction of the hypertonic saline-induced pain effect on EMG activity (i.e., whether the hypertonic saline-induced EMG activity was less than or greater than control EMG activity) could change with the magnitude of jaw displacement. Hypertonic saline infusion had no significant effect on postural EMG activity in any of the recorded jaw muscles. The data suggest that under constrained goal-directed tasks, the pattern of pain-induced changes in jaw muscle EMG activity is not clear cut, but can vary with the task performed, jaw displacement magnitude, and the subject being studied.     

A comparison of jaw-opener and jaw-closer muscle activity in humans to overcome an external force counteracting jaw movement

In contrast to the jaw-closer muscles, no or very few spindles are present in the jaw-opening digastric muscle. Therefore sensory feedback to the digastric muscle may be different from feedback to the jaw-closer muscles, resulting in a different reaction when jaw movement is perturbed. This possible difference was investigated by comparing the reaction of the digastric muscle when jaw opening is perturbed, with the reaction of the masseter muscle when jaw closing is perturbed. Subjects made rhythmic, 1-Hz open-close movements of the jaw under control of a metronome. During jaw opening (digastric muscle) or, in the other experiments, during jaw closing (masseter muscle), an external force counteracting jaw movement could appear. Series of movements without the force were unexpectedly alternated by series with the force. In both muscles sensory induced activity started approximately 25 ms after the onset of the force and consisted of two phases. In the masseter muscle the maximum of the first increase was reached significantly sooner (37±2 ms SEM) than in the digastric muscle (54±3 ms). The second increase appeared much sooner in the masseter muscle (73±4 ms) than in the digastric muscle (159±10 ms). When the force was expected, in both muscles an increase in preprogrammed muscle activity was observed. Also an increase in reflex activity, generated before 120 ms after the onset of the force, was observed, compared with when the force appeared unexpectedly. The relative increase in reflex activity was approximately 2 times larger than the relative increase in preprogrammed activity. Therefore, the increase in reflex activity when the force was expected may have been caused not only by an increase in recruitment, but also by an increase in the gain of the reflex loops. Reflex activity relative to preprogrammed activity was on average 4 times larger in the masseter muscle than in the digastric muscle. This indicates that the masseter muscle can react more adequately to disturbances of jaw movement than the digastric muscle. Received: 26 December 1996 / Accepted: 2 July 1997     

Phasic and tonic stretch reflexes in muscles with few muscle spindles: human jaw-opener muscles

 We investigated phasic and tonic stretch reflexes in human jaw-opener muscles, which have few, if any, muscle spindles. Jaw-unloading reflexes were recorded for both opener and closer muscles. Surface electromyographic (EMG) activity was obtained from left and right digastric and superficial masseter muscles, and jaw orientation and torques were recorded. Unloading of jaw-opener muscles elicited a short-latency decrease in EMG activity (averaging 20 ms) followed by a short-duration silent period in these muscles and sometimes a short burst of activity in their antagonists. Similar behavior in response to unloading was observed for spindle-rich jaw-closer muscles, although the latency of the silent period was statistically shorter than that observed for jaw-opener muscles (averaging 13 ms). Control studies suggest that the jaw-opener reflex was not due to inputs from either cutaneous or periodontal mechanoreceptors. In the unloading response of the jaw openers, the tonic level of EMG activity observed after transition to the new jaw orientation was monotonically related to the residual torque and orientation. This is consistent with the idea that the tonic stretch reflex might mediate the change in muscle activation. In addition, the values of the static net joint torque and jaw orientation after the dynamic phase of unloading were related by a monotonic function resembling the invariant characteristic recorded in human limb joints. The torque-angle characteristics associated with different initial jaw orientations were similar in shape but spatially shifted, consistent with the idea that voluntary changes in jaw orientation might be associated with a change in a single parameter, which might be identified as the threshold of the tonic stretch reflex. It is suggested that functionally significant phasic and tonic stretch reflexes might not be mediated exclusively by muscle spindle afferents. Thus, the hypothesis that central modifications in the threshold of the tonic stretch reflex underlie the control of movement may be applied to the jaw system. Received: 11 October 1996 / Accepted: 17 March 1997     

Sensitivity of monosynaptic test reflexes to facilitation and inhibition as a function of the test reflex size: a study in man and the cat

Summary In parallel experiments on humans and in the cat it was investigated how the sensitivity of monosynaptic test reflexes to facilitation and inhibition varies as a function of the size of the control test reflex itself. In man the monosynaptic reflex (the Hoffmann reflex) was evoked in either the soleus muscle (by stimulation of the tibial nerve) or the quadriceps muscle (by stimulation of the femoral nerve). In the decerebrate cat monosynaptic reflexes were recorded from the nerves to soleus and medial gastrocnemius muscles; they were evoked by stimulation of the proximal ends of the sectioned L7 and S1 dorsal roots. Various excitatory and inhibitory spinal reflex pathways were used for conditioning the test reflexes (e.g. monosynaptic Ia excitation, disynaptic reciprocal inhibition, cutaneous inhibition, recurrent inhibition, presynaptic inhibition of the Ia fibres mediating the test reflex). It was shown that the additional number of motoneurones recruited in a monosynaptic test reflex by a constant excitatory conditioning stimulus was very much dependent on the size of the test reflex itself. This dependency had the same characteristic pattern whatever the conditioning stimulus. With increasing size of the test reflex the number of additionally recruited motoneurones first increased, then reached a peak (or plateau) and finally decreased. A similar relation was also seen with inhibitory conditioning stimuli. The basic physiological factors responsible for these findings are discussed. Finally, the implications for the interpretation of experiments in man with the H-reflex technique are considered.     

Task-related behaviour of motor units in the human temporalis muscle

Muscle activity patterns in some complex human jaw muscles appear to be task sensitive. However, it is presently uncertain how changes in motor task affect motor unit (MU) behaviour in the human temporalis muscle. In this study, activity was recorded from 40 MUs in the anterior region of the muscle. The lowest sustainable firing frequency (LSFF) was reached by slow increases and decreases in firing rate, then firing was maintained at the lowest possible rate without significant pauses. An array of consecutive interspike intervals (ISI) were sampled digitally and used to measure the LSFF for each task associated with the MU. In a controlled paradigm, MU reflex inhibition was measured during the performance of different tasks. Single electrical pulses of non-noxious intensity were delivered to the gingiva near the maxillary canine tooth. During continuous MU firing at a controlled firing frequency of 10 Hz, series of pulses were delivered with increasing delays, after preselected spikes. The MUs fired continuously during the performance of 1–4 postural and tooth-contact tasks. There were significant differences in LSFFs between tasks in those MUs associated with multiple tasks. In the reflex study, all MUs were inhibited, but the magnitude of the inhibition was highly task dependent. Thus, both LSFF and reflex inhibition of temporalis MUs appear to vary with the motor task and are sensitive to the position of the jaw and the direction and location of tooth contact along the tooth row. This behaviour most likely reflects task-related changes in output from orofacial and muscle afferents.