Stretch reflexes in human masseter.

The reflex response to stretch in most contracting human muscles includes both a short-latency, probably monosynaptic, excitatory component, and a longer-latency, polysynaptic excitation. However, it has been claimed that stretch of the jaw-closing muscles evokes only the short-latency response in masseter. This question was re-examined, using controlled stretches of varied rates and durations. Very brief, rapid stretches analogous to the stimuli used to investigate the 'jaw-jerk' reflex in earlier studies evoked a prominent excitatory peak in the electromyogram at monosynaptic latency excitation, but little or no longer-latency excitation. This response could be produced even by stimuli that were barely detectable by the subject. However, this prominent electrical response did not produce a measurable increase in biting force. In contrast, slower stretches evoked both a short- and a longer-latency excitatory response in the surface electromyogram, as in most limb muscles. It is shown that the absence of a long-latency excitatory response in earlier studies can be explained by the powerful reflex disfacilitation of the motoneurones that occurred at the end of the brief stretches used. Depending on the duration of the stretch, this disfacilitation is often sufficient to mask or abolish the long-latency reflex. The reflex response to stretches was not markedly affected by blocking the activation of mechanoreceptors around the teeth with local anaesthetic, indicating that receptors around the teeth cannot be playing more than a minor role in the response. The stretch-induced increase in force became greater as the velocity of the stretch decreased.     

Responses of Single Motor Units in Human Masseter to Transcranial Magnetic Stimulation of Either Hemisphere

The corticobulbar inputs to single masseter motoneurons from the contra- and ipsilateral motor cortex were examined using focal transcranial magnetic stimulation (TMS) with a figure-of-eight stimulating coil. Fine-wire electrodes were inserted into the masseter muscle of six subjects, and the responses of 30 motor units were examined. All were tested with contralateral TMS, and 87 % showed a short-latency excitation in the peristimulus time histogram at 7.0 ± 0.3 ms. The response was a single peak of 1.5 ± 0.2 ms duration, consistent with monosynaptic excitation via a single D- or I₁-wave volley elicited by the stimulus. Increased TMS intensity produced a higher response probability (   n = 13  , paired t test,   P < 0.05  ) but did not affect response latency. Of the remaining motor units tested with contralateral TMS, 7 % did not respond at intensities tested, and 7 % had reduced firing probability without any preceding excitation. Sixteen of these motor units were also tested with ipsilateral TMS and four (25 %) showed short-latency excitation at 6.7 ± 0.6 ms, with a duration of 1.5 ± 0.3 ms. Latency and duration of excitatory peaks for these four motor units did not differ significantly with ipsilateral vs . contralateral TMS (paired t tests,   P > 0.05  ). Of the motor units tested with ipsilateral TMS, 56 % responded with a reduced firing probability without a preceding excitation, and 19 % did not respond. These data suggest that masseter motoneurons receive monosynaptic input from the motor cortex that is asymmetrical from each hemisphere, with most low threshold motoneurons receiving short-latency excitatory input from the contralateral hemisphere only.     

Excitation and inhibition of trigeminal motoneurons by palatal stimulation

Summary Excitation and inhibition of jaw-closing motoneurons (Masseteric and Temporal Motoneurons, Mass. and Temp. Mns) during transient jaw closing, the so-called jaw-closing reflex, and prolonged jaw opening elicited by palatal stimulation were studied. By pressing the anterior palatal surfaces sustained jaw opening was elicited, suggesting that sustained jaw opening results from inhibition of tonic background activity of jaw-closing motoneurons by inhibitory postsynaptic potentials (IPSPs) elicited by mechanical stimulation of the anterior palatal mucosa. Recordings showed that the onset of IPSPs was 80 ms earlier than the onset of jaw opening. Application of diffuse pressure stimulation to the posterior palatal surfaces elicited bursts of spikes triggered on excitatory postsynaptic potentials (EPSPs), suggesting that mechanosensory receptors from the posterior palatal mucosa send excitatory synaptic inputs to jaw-closing motoneurons. Furthermore, it is suggested that mechanosensory inputs from the posterior palatal mucosa may excite neurons in the central pattern generator and provide the motor patterns responsible for jaw closure during the jaw-closing reflex. We have demonstrated that excitation of Mass. Mns innervating the deep masseter muscle mainly contributed to maintaining the occlusal phase of jaw closure during the jaw-closing reflex. However, the onset of EPSPs was 100 to 160 ms (n = 27) earlier than the onset of jaw closure. In studies on spontaneously occurring jaw closure it was demonstrated that there was a proportional increase in the number of spikes of the Temp. Mn and the mechanical response (jaw closure).     

Reflex responses induced by tooth unloading

The reflex response of the masseter muscle to the rapid unloading of a single maxillary incisor tooth was studied. Unloading of a static force of 2 N in the horizontal direction resulted in a short-latency excitation, inhibition, and long-latency excitation of masseter muscle activity occurring at latencies of approximately 13, 20, and 40 ms, respectively, with a corresponding change in bite force occurring slightly later in each case. Following the blocking of periodontal input by the injection of local anesthetic around the stimulated tooth, inhibitory responses were abolished. Therefore, it is concluded that the observed masseteric inhibition was caused by the unloading of periodontal mechanoreceptors and thus that these receptors may contribute to the jaw unloading reflex.     

Response of human jaw muscles to axial stimulation of the incisor

The role of periodontal mechanoreceptors (PMRs) in the reflex control of the jaw muscles has thus far been mainly derived from animal studies. To date, the work that has been done on humans has been limited and confined to orthogonal stimulation of the labial surface of the tooth. The purpose of this study was to investigate the response of the masseter and digastric muscles in humans to controlled axial stimulation of the upper left central incisor, both before and during a local anaesthetic block of the PMRs. Ten neurologically normal young adult females were tested, each on two separate occasions to confirm the reproducibility of the results. It was found that the reflex response in the masseter was modulated by the rate of rise of the stimulus used and, to a lesser degree, the level of background muscle activity. There was little detectable change in the activity of the digastric muscle under the tested conditions and what was found could be attributed to cross-talk with the masseter. The reflex responses obtained were significantly different between subjects; however retesting the same subject on a different occasion yielded similar results. The results indicate that the most common response of the masseter muscle to brisk axial stimulation of the incisor is a reflex inhibition at 20 ms, followed by a late excitation at 44 ms. However, it is possible that this late excitation could be due to delayed action potentials and hence be artefactual. As the application of a local anaesthetic block removed or significantly reduced both of these responses, it was concluded that they originated from the PMRs. Unlike during orthogonal stimulation, slowly rising stimuli did not produce any excitatory reflex activity. This indicated a difference in jaw reflexes to forces applied in different directions, possibly due to the activation of different receptor types when stimulating the tooth in either the orthogonal or axial directions.     

Control of trigeminal motoneurons from the cerebellar interpositus nucleus of the guinea pig.

1. Effects of stimulation of the cerebellar interpositus nucleus (IPN) on jaw reflexes and trigeminal motoneurons were studied along with the route responsible for the effects in ketamine-anesthetized guinea pigs. 2. Stimulation in the IPN evoked a bilateral, ipsilaterally dominant, short-latency reciprocal effect on jaw reflexes: a depression of the jaw-closing masseteric reflex and a facilitation of the jaw-opening digastric reflex. This reciprocal phase was followed by a non-reciprocal facilitatory phase. 3. Stimulation in the IPN evoked a short-latency, strychnine-sensitive inhibitory postsynaptic potential (IPSP) followed by a rebound depolarizing potential in the jaw-closing masseter motoneurons (MAMNs) and an excitatory postsynaptic potential (EPSP) in the jaw-opening anterior digastric motoneurons (ADMNs). The time course of the intracellular response of MAMNs to the IPN stimulation was similar to that of the IPN-induced effects on the jaw-closing reflex. In contrast, the duration of the IPN-induced EPSP in ADMNs was shorter than the IPN-induced facilitation of the jaw-opening reflex. 4. After the IPN neurons were lost by injection of kainic acid into the nucleus, the reciprocal effect of the IPN stimulation on the jaw reflexes could not be seen, even though the projection fibers from the trigeminal sensory nucleus to the IPN remained essentially intact. 5. Stimulation in the superior cerebellar peduncle (SP) induced the same reciprocal effect on the jaw reflexes as the IPN stimulation. A lesion of the SP virtually abolished the reciprocal effect on the jaw reflexes of stimulation in the IPN ipsilateral with reference to the SP lesion. 6. Transection of the brain stem at the level immediately caudal to the red nucleus did not affect the reciprocal effect of the IPN stimulation on the jaw reflexes. 7. We conclude that the IPN output bilaterally induces an inhibition of jaw-closing MAMNs and an excitation of jaw-opening ADMNs, oligosynaptically by way of a direct cerebelloreticular projection system via the SP.     

Excitatory and inhibitory controls of the masseter and temporal muscles elicited from teeth in the rat.

The periodontal mechanism that controls the jaw reflexes was examined in lightly anesthetized rats. Motor-unit activity in the masseter and temporal muscles was recorded electromyographically and pressure stimulation was applied to either an upper incisor or an upper molar. Reflex effects of dental stimulation varied depending on the level of ongoing activity (background activity, BGA) in each motor unit. Incisal or molar stimulation elicited excitatory reflexes in both the masseter and temporal motor units at a low BGA, but inhibitory reflexes in both types of motor unit at a higher BGA. In contrast to these synergistic reflex actions, the reciprocal reflex actions of the two motor units that belonged to the respective muscles occurred when the BGA was intermediate. The results suggest that different patterns of periodontal jaw reflexes may be elicited, depending on the different levels of BGAs. Furthermore, the present reflexes were modified with the site of a stimulated tooth within the dentition. Incisal stimulation produced greater excitation in the masseter motor unit than in the temporal one, and the opposite type of response occurred during molar stimulation. Moreover, smaller-amplitude motor units with a low reflex threshold and larger-amplitude motor units with a higher reflex threshold tended to exhibit excitatory and inhibitory reflexes, respectively.     

Is the long-latency stretch reflex in human masseter transcortical?

A long-latency stretch reflex (LLSR) has been described in the human masseter muscle, but its pathway remains uncertain. To investigate this, the excitability of corticomotoneuronal (CM) cells projecting to masseter motoneurons during the LLSR was assessed with transcranial magnetic stimulation (TMS). A facilitated response to TMS would be evidence of a LLSR pathway that traverses the motor cortex. Surface electromyogram electrodes were placed over the left or right masseter, and subjects (n=10) bit on bars with their incisor teeth at 10% of maximal electromyographic activity (EMG). Servo-controlled displacements were imposed on the lower jaw to evoke a short- and long-latency stretch reflex in masseter. TMS intensity was just suprathreshold for a response in contralateral masseter. Trials consisted of: (1) stretch alone, (2) TMS alone, and (3) TMS with a preceding conditioning stretch at varied conditioning-testing (C-T) intervals chosen to combine TMS with the short-latency stretch reflex (3 ms, 5 ms) and the LLSR (23–41 ms). Masseter EMG was rectified and averaged. With TMS alone, mean (± SE) MEP area above baseline was 56±9%. The area of masseter MEPs above baseline in the C-T trials was calculated from each EMG average following subtraction of the response to stretch alone. Conditioning muscle stretch had no significant effect on masseter MEPs evoked by TMS with any C-T interval (ANOVA; P=0.90). In addition, subjects were unable to modify the SLSR or LLSR by voluntary command. It is concluded that the long-latency stretch reflex in the masseter does not involve the motor cortex and is not influenced by "motor set". Electronic Publication     

Distribution of recurrent inhibition within a motor nucleus. I. Contribution from slow and fast motor units to the excitation of Renshaw cells

The relation between the size of a monosynaptic reflex (MSR) to triceps surae and the resulting Renshaw cell discharge was used to evaluate the contribution from slow and fast motor units to the excitation of Renshaw cells. It is, however, difficult to interpret these results in terms of excitation contributed by slow and fast motor units because of the following reasons. First, the size of the MSR recorded in ventral roots is not linearly related to the number of recruited motor units, since larger motor axons contribute more to the size of the MSR than smaller ones. Second, the number of spikes evoked in a Renshaw cell burst is not linearly related to the excitatory input because Renshaw cell discharge saturates in the case of large responses. The contribution of small, early-recruited motoneurones to Renshaw cell excitation is consequently overestimated. Procedures were introduced to deal with these problems. It is concluded that the last-recruited motor units (probably 'fast twitch, fast fatiguing') on average contribute four times as much excitation to Renshaw cells as the first recruited ('slow twitch') motor units.     

Synaptic connections from large afferents of wrist flexor and extensor muscles to synergistic motoneurones in man

 Short-latency excitatory Ia reflex connections were determined between pairs of human wrist flexor and extensor muscles. Spindle Ia afferents were stimulated by either tendon tap or electrical stimulation. The activity of voluntarily activated single motor units was recorded intramuscularly from pairs of wrist flexor or extensor muscles. Cross-correlation between stimuli and the discharge of the motor units provided a measure of the homonymous or heteronymous excitatory input to a motoneurone. Homonymous motoneurone facilitation was generally stronger than that of the heteronymous motoneurones. The principal wrist flexors, flexor carpi radialis (FCR) and flexor carpi ulnaris (FCU), were tightly connected through a bidirectional short-latency reflex pathway. In contrast, the extensor carpi ulnaris (ECU) and the extensor carpi radialis (ECR) did not have similar connections. ECU motoneurones received no short-latency excitatory Ia input from the ECR. ECR motoneurones did receive excitatory Ia input from ECU Ia afferents; however, its latency was delayed by several milliseconds compared with other heteronymous Ia excitatory effects observed. The wrist and finger extensors were linked through heteronymous Ia excitatory reflexes. The reflex connections observed in humans are largely similar to those observed in the cat, with the exception of heteronymous effects from the ECU to the ECR and from the extensor digitorum communis (EDC) to the ECU, which are present only in humans. The differences in the reflex organization of the wrist flexors versus the extensors probably reflects the importance of grasping. Received: 19 August 1996 / Accepted: 6 March 1997     

Jaw reflexes evoked by mechanical stimulation of teeth in humans.

Jaw reflexes evoked by mechanical stimulation of teeth in humans. The reflex response of jaw muscles to mechanical stimulation of an upper incisor tooth was investigated using the surface electromyogram (SEMG) of the masseter muscle and the bite force. With a slowly rising stimulus, the reflex response obtained on the masseter SEMG showed three different patterns of reflex responses; sole excitation, sole inhibition, and inhibition followed by excitation. Simultaneously recorded bite force, however, exhibited mainly one reflex response pattern, a decrease followed by an increase in the net closing force. A rapidly rising stimulus also induced several different patterns of reflex responses in the masseter SEMG. When the simultaneously recorded bite force was analyzed, however, there was only one reflex response pattern, a decrease in the net closing force. Therefore, the reflex change in the masseter muscle is not a good representative of the net reflex response of all jaw muscles to mechanical tooth stimulation. The net response is best expressed by the averaged bite force. The averaged bite force records showed that when the stimulus force was developing rapidly, the periodontal reflex could reduce the bite force and hence protect the teeth and supporting tissues from damaging forces. It also can increase the bite force; this might help keep food between the teeth if the change in force rate is slow, especially when the initial bite force is low.     

Vibration-induced discharge patterns of single motor units in the masseter muscle in man.

Single motor unit potentials were recorded with small bipolar wires from intact masseter muscles in the adult man and a detailed parametric analysis of the effects of muscle vibration on motor unit discharges was carried out. 2. When the vibration amplitude was kept constant, each unit started firing at a definite threshold of vibration frequency. With higher frequencies the rate of firing rapidly reached a maximum. Units recruited at higher frequencies presented a lower maximum rate of firing. 3. When the vibration frequency was kept constant, each masseter unit discharged at a definite threshold of vibration amplitude. With higher amplitudes the unit quickly reached a maximum rate of discharge. Units with a higher frequency threshold tended to also present a higher amplitude threshold. Motor unit "excitability" curves could be plotted using the combined threshold conditions for frequency and amplitude of applied vibrations. 4. With a given parametric set of vibration, the units only started firing at a given delay after the onset of vibration. The delay was quite different for different units and it increased considerably, sometimes by several seconds, when the vibration amplitude was made smaller. 5. In all the experimental conditions tested, and even when the unit discharge did not start until several seconds after vibration onset, the unit potential presented a close and highly consistent temporal relation to the vibration cycles. The slow recruitment process is thought to involve a polysynaptic excitatory mechanism which progressively depolarizes the masseter motoneurones close to their threshold, the actual firing being triggered by monosynaptic excitatory post-synaptic potentials from I(a) afferents, hence the small latency jitter recorded. This special pattern of tonic vibration reflex in jaw-closing muscles in man may result from the lack of reciprocal inhibition from the jaw-opening muscles.     

Lateralized phrenic nerve responses to stimulating respiratory afferents in the cat.

We stimulated electrically pharyngeal branch of both glossopharyngeal nerves (PGLN), internal branch of superior laryngeal nerves (ISLN), and carotid sinus nerves (CSN) in anesthetized cats. We recorded simultaneously, averaged, and compared bilaterally evoked phrenic nerve (PHR) activity. Our objective was to demonstrate a short-latency evoked response in the PHR contralateral to the stimulus. Low-intensity stimulation of PGLN and ISLN during inspiration evoked a short-latency contralateral excitation with a latency of 5.2 ms +/- 0.2 SE (16 cats) for PGLN, and 3.8 ms +/- 0.1 SE (13 cats) for ISLN. This excitation could follow stimuli delivered at 100 Hz. Stimulation during expiration did not result in a lateralized excitation. The excitation is followed by bilateral inhibition. Neither strychnine nor picrotoxin prevented either the lateralized response or the inhibition, though strychnine diminished a delayed bilateral excitation following PGLN stimulation. This dalayed (latency 18.7 ms +/- 0.7 SE) bilateral excitation corresponds to the sniff reflex. CSN stimulation did not result in lateralized excitation. We suggest that the lateralized evoked response results from a gated paucisynaptic reflex pathway involving the PGLN and ISLN, ipsilateral inspiratory neurons, and contralateral PHR motoneurons.     

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.     

Discharge of spindle afferents from jaw-closing muscles during chewing in alert monkeys.

The discharge of muscle spindle afferents from monkey spindle afferents from monkey jaw-closing muscles was studied during mastication of natural foods by extracellular recording from the fibers or cell bodies of the tract and mesencephalic nucleus of the fifth nerve. In all, 39 muscle afferents were studied. The spindle associated with 18 of the afferents was positively identified by the afferent's response to gentle, localized palpation of either the temporalis or masseter muscle. Discharge patterns were observed during mastication, and in the majority of cases the qualitative passive response characteristics of the spindle afferent were determined. During steady chewing spindle afferent discharge typically paused briefly during the initial rapid upward part of the chewing cycle. Firing generally began as the jaw slowed its upward movement, and firing rates during the slow grinding portion of the upward movement were within the range of 50-80 spikes/s. All spindles exhibited a brisk discharge during the opening movement, typically within the range of 100-150 spikes/s. One-third of the spindle afferents exhibited a brief, high-frequency burst of firing at the very beginning of the opening movement, presumably as a result of stretch applied to a spindle just previously subjects to fusimotor excitation. Although the results of the study make it clear that spindles in jaw-closing muscles are coactived along with the extrafusal muscle fibers, the fusimotor bias does not seem capable of sustaining discharge in the face of rapid shortening of the muscle. Furthermore, the fact that discharge rate during opening, when the jaw-closing motoneurons are quiescent, is much higher than at any part of the closing cycle, when the motoneurons are active, suggests that the muscle spindles cannot provide the primary excitatory drive to the motoneurons.     

Modulation of stretch-evoked reflexes in single motor units in human masseter muscle by experimental pain

The interaction between muscle pain and motor function of the jaw has been examined in recent years, but the nature of the modulation of the short-latency stretch reflex by pain is not fully understood. In this study, the reflex responses to stretch were measured in single low-threshold motor units that were kept discharging at a constant frequency, before, during and after the induction of experimental pain in one masseter muscle by controlled infusion of hypertonic saline. The probability of evoking a reflex response in individual motor units in the painful muscle at near-monosynaptic latency was reduced by a mean of about 20%. However, the overall reflex response in the surface electromyogram of both the ipsi- and contralateral masseter muscles was greater during pain. This was apparently a secondary response to the pain-induced increase in pre-stimulus activity in the motoneurone pools of both muscles, because increased motoneurone excitability may facilitate stretch reflexes. It is concluded that the most likely explanation for the reduced reflex response of low-threshold masseter motor units during experimental pain is a tonic reduction in the fusimotor drive to the masseter spindles.     

Increases in reflex excitability of monkey masseter motoneurons before a jaw-bite reaction-time response.

Reflex excitability of the motoneurons innervating the masseter muscle of monkeys was tested before a phasic voluntary activation of the jaw-closing muscles (a RT bite response). Single test shocks were delivered to the Mes V which supplies a monosynaptic excitatory input to the jaw-muscle motoneurons. Changes in reflex excitability were assessed by measuring the amplitude of the synchronous muscle potential evoked by the test shock. Amplitudes of the muscle potentials evoked by shocks which occurred just before the beginning of the voluntary EMG response, as judged by the onset of EMG activity of the masseter muscle contralateral to the test shock were many times larger than potentials evoked immediately following the visual RT stimulus. Curves relating the average amplitude of the evoked response to its time before the beginning of the voluntary response suggest that the reflex excitability of the motoneuron pool begins to increase 25-45 ms before the first detectable EMG activity occurs. These results suggest that inputs arrive at the motoneurons of agonist muscles used in rapid RT tasks substantially before changes in the EMG of the muscle are noted. These results, in part, would account for the time interval noted between the beginning of neural activity in suprasegmental structures which presumably excites spinal motoneurons, and the first EMG activity of muscles which are innervated by these motoneurons.     

Jaw muscle response to stimulation of type II somatosensory afferents of limbs in the rat

Convergence of various afferent inputs onto brainstem neurones may play an important role in the regulation of trigeminal motor activity. In particular, previous studies suggest that, besides sensory inputs arising from the orofacial region, extratrigeminal information may modulate jaw muscle function. In the present study the actions exerted on masseter and digastric muscles by the activation of somatosensory afferents coming from fore- and hind limbs were examined. The electromyographic activity (EMG) of masseter and digastric muscles was recorded in 20 anaesthetised rats, and EMG responses to single and paired electrical stimulation of common radial and sciatic nerves, at a threshold intensity for the activation of group II afferent fibres, were studied. The stimulation induced an excitatory response in both masseter and digastric muscles bilaterally. Ipsi- and contralateral radial nerve stimulation evoked masseter responses at latencies of 13.8±2.4 ms and of 18.0±2.6 ms, respectively, and digastric responses 1.6±0.4 ms later. Ipsi- and contralateral sciatic nerve stimulation elicited masseter responses at latencies of 21.4±2.6 ms and of 23.3±2.0 ms, respectively, and digastric responses 2.0±0.2 ms later. The same masseter and digastric motor units were excited by both radial and sciatic nerve stimulation; this suggests a convergence of somatosensory inputs arising from fore- and hind limbs on the same pool of masseter and digastric motoneurones. Paired stimulation of the two nerves did not induce any summation of the responses; this finding suggests that the two inputs, reaching a common relay station, could give rise either to occlusion or to inhibitory interactions. Spinotrigeminal relationship evidenced in this study may be involved in the coordination of jaw and limb movements. Electronic Publication     

Intracellular analysis of reflex pathways underlying the stumbling corrective reaction during fictive locomotion in the cat

In cat and humans, contact between an obstacle and the dorsum of the foot evokes the stumbling corrective reaction (reflex) that lifts the foot to avoid falling. This reflex can also be evoked by short trains of stimuli to the cutaneous superficial peroneal (SP) nerve in decerebrate cats during the flexion phase of fictive locomotion. Here we examine intracellular events in hindlimb motoneurons accompanying stumbling correction. SP stimulation delivered during the flexion phase excites knee flexor motoneurons at short latency [minimum excitatory postsynaptic potential (EPSP) latency 1.8 ms; mean 2.7 ms]. Although a similar short latency excitation occurs in ankle extensors (mean latency, 2.8 ms), recruitment is delayed until successive shocks in the stimulus train overcome the locomotor-related hyperpolarization of ankle extensors. In ankle flexor motoneurons, SP stimulation evokes an inhibition (mean latency, 2.7 ms) that briefly reduces or stops their firing during the flexion phase. There is a phase-dependent modulation of SP-evoked EPSP amplitude as well as latency during locomotion. However, the more obvious change in SP reflex pathways with the onset of fictive locomotion is the reduced inhibition of ankle extensor motoneurons and the increased inhibition of ankle flexors. These results show that the characteristic pattern of hindlimb motoneuron activation during SP nerve-evoked stumbling correction results from 1) di- and trisynaptic excitation of knee flexor and ankle extensor motoneurons; 2) increased inhibitory postsynaptic potentials in ankle flexors and a suppression of inhibition in extensors, 3) sculpting of the short-latency SP postsynaptic effects by motoneuron membrane potential, and 4) longer latency excitatory effects that are likely evoked by lumbar interneurons involved in the generation of fictive locomotion.     

Response of human jaw muscles to axial stimulation of a molar tooth

The reflexes of the main jaw-closer muscles (masseter and anterior temporalis) on both sides of the jaw were investigated using surface electromyography to observe reflex activity following mechanical stimulation of the 1st right upper-molar tooth at various forces under a number of levels of jaw-muscle activity. As with analogous studies performed on the incisor, three distinct reflex events were identified in the EMG before the earliest conscious subject reaction: early excitation, inhibition and late excitation. However, contrary to observations found during studies on the incisor, excitation, not inhibition was the primary reflex response. The application of a local anaesthetic block around the stimulated molar showed that the primary agents in eliciting the observed reflexes were not contained within the periodontium of the stimulated tooth. A diminished representation of periodontal mechanoreceptors around the molar teeth and more elaborate root structures, hence a more solid connection to the jaw and consequently less tooth movement, were deemed the likely reason for the distinction between the reflex responses of the incisal and molar regions. In addition to the reflex studies, the minimum reaction time of a number of subjects was determined to permit the distinction of a reflex event and an event that could be a conscious subject reaction. It was found that the reaction time of the temporalis muscles was significantly shorter than those of the masseter, while no significant difference was found between the left and right sides. Overall, the data showed that the presence or absence of background muscle activity and subject variability were the main causes of changes in the reflex response, provided the level of the stimulus was greater than 3 N. The application of local anaesthetic had no impact on the reflexes evoked.