Electrogenesis of muscle fibers of the rat masseter muscle

It was shown by intracellular recording of resting membrane potentials (RMP) and action potentials that the superficial layers of the rat masseter muscle contain chiefly fibers with a high MPP and small overshoot, whereas the deep layers contain mainly fibers with a low MPP but a high overshoot. The excitability of the cytoplasmic membrane of muscle fibers with different MPP levels was found to be similar with respect to its electrical parameters. It is suggested that the rat masseter muscle contains a high proportion of fast phasic fibers in its superficial layers and slow phasic fibers in its deep layers.Department of Pathological Physiology, N. A. Semashko Moscow Medical Stomatologic Institute. (Presented by A. D. Ado.) Translated from Byulleten' Éksperimental'noi Biologii i Meditsiny, Vol. 88, No. 9, pp. 265–267, September, 1979.     

Reflex responses of motor units in human masseter muscle to electrical stimulation of the lip

Summary The reflex responses of single motor units in human masseter muscle to electrical stimulation of the lip were recorded. The subject maintained a contant mean level of pre-stimulus excitation of the parent motor neurone by biting in such a way that the unit fired at either 10 or 15 Hz during each trial. When firing at 10 Hz, most units were reflexly inhibited for up to 90 ms by electrical stimuli at intensities that were perceived to be mildly uncomfortable. In many units, the inhibition consisted of 2 phases which were separated from each other by a few spikes occurring about 30 ms after the stimulus. It was occasionally possible to evoke only the later phase (latency about 40 ms) with stimuli at intensities near the response threshold. In these instances, the inhibitory response became biphasic at higher stimulus intensities with the emergence of a shorter (10–15 ms) component. Still higher intensities caused the 2 phases of inhibition to merge, giving the appearance of a single, prolonged, inhibitory response. When the pre-stimulus firing frequency of the unit was changed from 10 Hz to 15 Hz, the inhibitory responses to the same stimuli were decreased, with the longer-latency component usually surviving beyond the shorter-latency phase. The pattern of reflex responses observed can be explained by a model based on information derived from intracellular recordings in animal experiments.     

Functional organization of the human masseter muscle

The authors carried out an anatomic and magnetic resonance imaging study of the architecture of the elevator muscles of the mandible in 169 cadavers. The aim of this study was to define the architectural organization of the human masseter muscle, temporalis and pterygoid muscles. Layered dissections and anatomic sections in different spatial planes showed that the masseter muscle exhibited a typical pennate structure consisting of a succession of alternating musculoaponeurotic layers. The muscle had three well-differentiated parts the superficial, intermediate and deep masseter muscles. The same pattern was constantly found 1) for the superficial masseter, two alternate musculoaponeurotic layers oriented at 60∘ in relation to the plane of occlusion, 2) for the intermediate masseter, a single musculo-aponeurotic layer oriented at 90∘ in relation to the occlusal plane, 3) for the deep masseter, three musculoaponeurotic layers whose general orientation was at 90∘ for the bounding layers and 110∘ for the intermediate layer. The MRI study confirmed the reality of this architectural arrangement.     

Evidence for parasympathetic vasodilator fibres in the rat masseter muscle

The present study was designed to examine (1) whether there are vasodilator fibres in the masseter muscle, and (2) if there are, to establish the neural pathways mediating these responses in urethane-anaesthetized rats. Electrical stimulation of the central cut end of the lingual nerve (LN) elicited intensity- and frequency-dependent increases of the blood flow in the masseter muscle (MBF) and lower lip (LBF). Increases in both the MBF and LBF evoked by the LN stimulation were reduced by hexamethonium in a dose-dependent manner (1–10 mg kg⁻¹). Pretreatment with phentolamine or propranolol at a dose of 100 μg kg⁻¹ had no effect on the increases in either MBF or LBF evoked by LN stimulation. Pretreatment with atropine (100 μg kg⁻¹) significantly reduced the MBF increase induced by LN stimulation, but not that in the LBF. The sectioning of the superior cervical sympathetic trunk did not affect the responses. MBF increases occurred with electrical stimulation of the trigeminal ganglion, and these increases were significantly reduced by the administration of hexamethonium and atropine. Lidocaine microinjection into the trigeminal spinal nucleus or salivatory nuclei caused a significant attenuation of the LN-induced MBF increases. When wheat germ agglutinin–horseradish peroxidase (WGA–HRP) was injected into the masseter muscle, labelled neurones were abundantly observed in the otic ganglion. The present study indicates that there are parasympathetic cholinergic and noncholinergic vasodilator fibres originating from cell bodies in the otic ganglion in the rat masseter muscle. The MBF increase evoked by activation of the parasympathetic fibres occurred via the trigeminal mediated reflex, suggesting that the novel parasympathetic vasodilator response may play an important role in the regulation of the haemodynamics of jaw muscles.     

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.     

Inflammation increases the excitability of masseter muscle afferents

Temporomandibular disorder is a major health problem associated with chronic orofacial pain in the masticatory muscles and/or temporomandibular joint. Evidence suggests that changes in primary afferents innervating the muscles of mastication may contribute to temporomandibular disorder. However, there has been little systematic study of the mechanisms controlling the excitability of these muscle afferents, nor their response to inflammation. In the present study, we tested the hypotheses that inflammation increases the excitability of sensory neurons innervating the masseter muscle of the rat and that the ionic mechanisms underlying these changes are unique to these neurons. We examined inflammation-induced changes in the excitability of trigeminal ganglia muscle neurons following intramuscular injections of complete Freund's adjuvant. Three days after complete Freund's adjuvant injection acutely dissociated, retrogradely labeled trigeminal ganglia neurons were studied using whole cell patch clamp techniques. Complete Freund's adjuvant-induced inflammation was associated with an increase in neuronal excitability marked by a significant decrease in rheobase and increase in the slope of the stimulus response function assessed with depolarizing current injection. The increase in excitability was associated with significant decreases in the rate of action potential fall and the duration of the action potential afterhyperpolarization. These changes in excitability and action potential waveform were associated with significant shifts in the voltage-dependence of activation and steady-state availability of voltage-gated K(+) current as well as significant decreases in the density of voltage-gated K(+) current subject to steady-state inactivation. These data suggest that K(+) channel subtypes may provide novel targets for the treatment of pain arising from inflamed muscle. These results also support the hypothesis that the underlying mechanisms of pain arising from specific regions of the body are unique suggesting that it may be possible, if not necessary to treat pain originating from different parts of the body with specific therapeutic interventions.     

Control of masseter muscle tension during sleep

This study evaluated whether muscle tension could be controlled during sleep. Twelve volunteers were assigned to an experimental condition or to a no treatment control group. Experimental subjects were instructed to reduce the tension in their masseter (jaw) muscle when signalled with an auditory tone. These subjects demonstrated cue-controlled tension reductions during awake training sessions. Although both groups evidenced muscle tension increases in response to the tones presented during sleep, experimental subjects produced significantly smaller increases in response to tone stimuli than did control subjects. More importantly, experimental subjects sustained lower tonic levels of muscle tension throughout the sleep sessions than did control subjects. Sleep was moderately disrupted for subjects in both groups. However, experimental subjects evidenced the greatest tension reductions during those responses in which they shifted to a lighter stage of sleep rather than fully awakening. These data suggest that muscle tension can be moderated during sleep.     

Arterial supply to the masseter muscle in the rat

Eighteen rats (thirty-six sides) were injected with red latex into the peripheral arteries through the left ventricle in the heart and fixed in 10% formalin to demonstrate the arterial architecture. According to the method of Yoshikawa et al. who proposed the lamination theory of this muscle, the latex injected specimens were dissected under a stereoscopic microscope. The masseter muscle in the rat was distributed by the masseteric branches of the facial, external carotid and dorsal branch of the infraorbital arteries as well as the transverse facial, masseteric and buccal arteries. This finding was essentially the same as observed on other species which included the dog, cat, crab-eating monkey, rabbit, cow and horse. However, the origin, course and distribution of the posterior deep temporal and masseteric arteries in the rat were considerably different from those of other species. Furthermore, since the way of development of the arteries and the subdivided muscles of the masseter muscle varies among species, the relationships between these arteries and the subdivided muscles seem to differ to some extent from species to species. Outline of the arterial system of the lateral aspect in the rat's head was shown in Fig. 1. The arteries and masseteric branches which were distributed to the subdivided muscles of the masseter muscle in the rat were as follows. 1) The first and second superficial and intermediate masseter muscles were distributed by the masseteric branches of the facial and external carotid arteries as well as the transverse facial, buccal and masseteric arteries. 2) The anterior portion of the deep masseter muscle was supplied by the masseteric branch of the facial, the masseteric and the buccal arteries. 3) The posterior portion of the deep masseter muscle received only the masseteric artery. 4) The maxillomandibular muscle was vascularized by the masseteric branch of the dorsal branch of the infraorbital and the buccal arteries. 5) the zygomaticomandibular muscle included only the masseteric artery.     

Sex-related differences in NMDA-evoked rat masseter muscle afferent discharge result from estrogen-mediated modulation of peripheral NMDA receptor activity

In the present study, the hypothesis that sex-related differences in glutamate-evoked rat masseter muscle afferent discharge may result from estrogen-related modulation of peripheral N-methyl-d-aspartate (NMDA) receptor activity and/or expression was tested by examining afferent fiber discharge in response to masseter injection of NMDA and the expression of NR2A/B subunits by masseter ganglion neurons in male and female rats. The results showed that injection of NMDA into the masseter muscle evoked discharges in putative mechanonociceptive afferent fibers and increased blood pressure that was concentration-dependent, however, a systemic action of NMDA appeared responsible for increased blood pressure. NMDA-evoked afferent discharge was significantly greater in female than in male rats, was positively correlated with plasma estrogen levels in females and was significantly greater in ovariectomized female rats treated with a high dose (5 mug/day) compared with a low dose (0.5 mug/day) of estrogen. Pre-treatment of high dose estrogen-treated-ovariectomized female rats with the Src tyrosine kinase inhibitor PP2 did not affect NMDA-evoked afferent discharge. NMDA-evoked afferent discharge was attenuated by the antagonists ketamine and ifenprodil, which is selective for NR2B containing NMDA receptors. Fewer masseter ganglion neurons expressed the NR2A (16%) subunit as compared with the NR2B subunit (38%), which was expressed at higher frequencies in intact female (46%) and high dose estrogen-treated ovariectomized female (60%) rats than in male (31%) rats. Taken together, these results suggest that sex-related differences in NMDA-evoked masseter afferent discharge are due, at least in part, to an estrogen-mediated increase in expression of peripheral NMDA receptors by masseter ganglion neurons in female rats.     

Inhibition of muscle spindle afferent activity during masseter muscle fatigue in the rat

The influence of muscle fatigue on the jaw-closing muscle spindle activity has been investigated by analyzing: (1) the field potentials evoked in the trigeminal motor nucleus (Vmot) by trigeminal mesencephalic nucleus (Vmes) stimulation, (2) the orthodromic and antidromic responses evoked in the Vmes by stimulation of the peripheral and central axons of the muscle proprioceptive afferents, and (3) the extracellular unitary discharge of masseter muscle spindles recorded in the Vmes. The masseter muscle was fatigued by prolonged tetanic masseter nerve electrical stimulation. Pre- and postsynaptic components of the potentials evoked in the Vmot showed a significant reduction in amplitude following muscle fatigue. Orthodromic and antidromic potentials recorded in the Vmes also showed a similar amplitude decrease. Furthermore, muscle fatigue caused a decrease of the discharge frequency of masseter muscle spindle afferents in most of the examined units. The inhibition of the potential amplitude and discharge frequency was strictly correlated with the extent of muscle fatigue and was mediated by the group III and IV afferent muscle fibers activated by fatigue. In fact, the inhibitory effect was abolished by capsaicin injection in the masseter muscle that provokes selective degeneration of small afferent muscle fibers containing neurokinins. We concluded that fatigue signals originating from the muscle and traveling through capsaicin-sensitive fibers are able to diminish the proprioceptive input by a central presynaptic influence. In the second part of the study, we examined the central projection of the masseter small afferents sensitive to capsaicin at the electron-microscopic level. Fiber degeneration was induced by injecting capsaicin into the masseter muscle. Degenerating terminals were found on the soma and stem process in Vmes and on the dendritic tree of neurons in Vmot. This suggests that small muscle afferents may influence the muscle spindle activity through direct synapses on somata in Vmes and on dendrites of neurons in Vmot.     

Distribution of slow muscle fiber of muscle spindle in postnatal rat masseter muscle

We investigated the properties of the muscle spindle in the masseter muscle at an immunohistochemical level in rats fed for 6 weeks. Slow myosin heavy chain (MyHC) isoforms were measured and intrafusal fibers in the muscle spindle were studied to determine the relationship between the superficial and deep regions of rat masseter muscle after alternated feeding pattern. However, muscle spindles were found in both regions, mainly in the deep region of the posterior superficial region of masseter muscle. The total number of the slow fiber in the intrafusal fiber and number of muscle spindle in the deep region were high from 5 to 8 weeks old in spite of various dimensions of data such as diameter and the compositions of the intrafusal fiber. The relationship of the protein expression of slow MyHC in the two regions at 5 weeks old reversed five weeks later (10 weeks old). This period is an important stage because the mastication system in masseter muscle with muscle spindle may be changed during the alternated feeding pattern of suckling to mastication. The changes may be a marker of the feeding system and of the control by the tension receptor of muscle spindle in this stage of masseter muscle after postnatal development.     

咬肌的基础与临床研究现状

咬肌是咀嚼肌群的重要组成部分。该肌收缩时产生强而有力的上提下颌骨的作用 ,使上、下颌牙列咬合而接触 ,并参与下颌前伸、侧向及后退运动和维持下颌姿势位。临床上由于神经损伤、外伤等各种原因所致咬肌功能丧失、缺损或畸形 ,都将影响病人的进食及面部美容。为了满足临床的需要 ,本文对有关咬肌的基础研究和临床应用方面的资料进行了综述 ,为临床医生提供参考资料。     

Sex differences in rabbit masseter muscle function

The proportions of fibers of different phenotypes in the rabbit masseter muscle differ strikingly in adult males and females. Muscles from females contain similar proportions of small fibers that express both the slow/beta and cardiac alpha myosin heavy chain (MyHC) isoforms and larger fibers containing the IIa MyHC isoform. In muscles from males, nearly 80% of fibers are of the IIa phenotype. To evaluate the functional significance of these sex differences, we used finely graded intramuscular microstimulation to study the contractile properties of masseter motor units in >6-month-old male and female rabbits. Twitch forces and torques in males were significantly greater in magnitude than those of females. Greater proportions of units that produced larger forces/torques were encountered in the males. The same motor units produced force or torque more rapidly in males than in females, principally because units in which twitch rise times were >22 ms were found only in females. The forces applied to the mandible and the torques generated about the right mandibular condyle were studied during cortically evoked rhythmic activation of the masticatory muscles. The overall range of torque rise times and the magnitudes of the peak torques did not differ between sexes. The mean rise time was significantly shorter and the mean peak torque was significantly greater in males. We conclude that sex differences in fiber phenotype proportions are reflected in sex differences in motor unit properties and in the function of these motor units during rhythmic activation.     

Glutamate-induced sensitization of rat masseter muscle fibers.

In rats, intradermal or intraarticular injection of glutamate or selective excitatory amino acid receptor agonists acting at peripheral excitatory amino acid receptors can decrease the intensity of mechanical stimulation required to evoke nocifensive behaviors, an indication of hyperalgesia. Since excitatory amino acid receptors have been found on the terminal ends of cutaneous primary afferent fibers, it has been suggested that increased tissue glutamate levels may have a direct sensitizing effect on primary afferent fibers, in particular skin nociceptors. However, less is known about the effects of glutamate on deep tissue afferent fibers. In the present study, a series of experiments were undertaken to investigate the effect of intramuscular injection of glutamate on the excitability and mechanical threshold of masseter muscle afferent fibers in anesthetized rats of both sexes.Injection of 1.0 M, but not 0.1 M glutamate evoked masseter muscle afferent activity that was significantly greater than that evoked by isotonic saline. The mechanical threshold of masseter muscle afferent fibers, which was assessed with a Von Frey hair, was reduced by approximately 50% for a period of 30 min after injection of 1.0 M glutamate, but was unaffected by injections of 0.1 M glutamate or isotonic saline. Injection of 25% dextrose, which has the same osmotic strength as 1.0 M glutamate, did not evoke significant activity in or decrease the mechanical threshold of masseter muscle afferent fibers. Magnetic resonance imaging experiments confirmed that injection of 25% dextrose and 1.0 M glutamate produced similar edema volumes in the masseter muscle tissue. Co-injection of 0.1 M kynurenate, an excitatory amino acid receptor antagonist, and 1.0 M glutamate attenuated glutamate-evoked afferent activity and prevented glutamate-induced mechanical sensitization. When male and female rats were compared, no difference in the baseline mechanical threshold or in the magnitude of glutamate-induced mechanical sensitization of masseter muscle afferent fibers was observed; however, the afferent fiber activity evoked by injection of 1.0 M glutamate into the masseter muscle was greater in female rats.The results of the present experiments show that intramuscular injection of 1.0 M glutamate excites and sensitizes rat masseter muscle afferent fibers through activation of peripheral excitatory amino acid receptors and that glutamate-evoked afferent fiber activity, but not sensitization, is greater in female than male rats.     

Variability in electromyographic surface recordings of the human masseter muscle

Serial registrations of integrated unilateral surface masseter EMG were performed in eight healthy subjects in the rest position, during controlled gentle biting and during maximal clenching at five different sessions in order to analyse the variability. The recording conditions were standardised. The EMG activity for the group did not differ significantly between the sessions at any level but individual variations were great in the rest position both within and between days. Good consistency was found at controlled force levels. Temporally stable measurements of masseter activity which permit quantitative comparisons thus seem possible.     

Age changes in neuromuscular junctions of masseter muscle

There is little information regarding the relationship between aging and ensuing morphological changes in the neuromuscular junction (NMJ) of the masticatory muscle. The masseter of C57BL/6J male mice at three different ages (young adult, 6 months; mature adult, 12 months; and old, 30 months) was studied. Morphological measurements were taken from zinc iodide osmium stained NMJ. Camera lucida drawings were superimposed on a computer monitor via a video camera and traced using a digitizer. The data were treated statistically using multivariate analysis of variance (MANOVA). No difference was found between 6 and 12 months of age, but a significant decrease in morphological parameters in NMJ from 30-month-old animals was found when compared with mature adult animals. Nerve terminal areas, perimeters, longitudinal extent lengths, and fiber diameters were reduced by 24%, 21%, 15.5% and 23%, respectively. Nerve terminal branches and incidence of sprouts were significantly increased in older animals. Age changes in the NMJ morphology are probably associated with altered balance between degeneration and regeneration of nerve terminals. This view is supported by increased terminal sprouting in old mice which is indicative of the plasticity or remodelling of NMJ during aging. © 1993 Wiley-Liss Inc.     

Anticipatory modulation of neck muscle reflex responses induced by mechanical perturbations of the human forehead

The aim of this study was to test whether anticipation of upcoming head blow stimuli, which elicit reflex responses in the neck muscle, makes the reflex responses greater or not. In nine healthy subjects the reflex responses were elicited in the sternocleidomastoid (SCM) muscle in the eyes-open (EO) and eyes-closed (EC) conditions, which corresponded to the predictable and unpredictable conditions, respectively. The subjects were instructed not to resist the perturbations after the impact. The results demonstrated that the reflex response of the SCM muscle was significantly smaller in the predictable EO condition than in the unpredictable EC condition (P < 0.05), although no significant differences were observed in either the background EMG activities or the head accelerations. Further, this effect of anticipation was observed only in the later reflex EMG component, which most probably mediated the stretch reflex pathway. In contrast, no significant difference was observed in the early component, which was presumed to be the vestibular-collic reflex. The reduced stretch reflex response was suggested to be functionally relevant to the task requirement, i.e., to let the neck extension movement occur, and not to resist after the impact of the head blow. It was concluded that the anticipation has an effect on reducing the stretch reflex responses in the neck muscle, but does not have any effect on the presumed vestibular-collic reflex under the present experimental paradigm. It is suggested that the gain of the stretch reflex pathway is modulated by anticipatory information of upcoming mechanical event.     

Length changes in the human masseter muscle after jaw movement

The human masseter is a multilayered, complex muscle contributing to jaw motion. Because variations in stretch may cause muscle fibers to function over different portions of their length-tension curves, the aim of this study was to determine how parts of the masseter lengthen or shorten during voluntary jaw movements made by living subjects. Magnetic resonance (MR) imaging and optically-based jaw-tracking were used to measure muscle-insertion positions for four parts of the muscle with six degrees of freedom (DOF), before and after maximum-opening, jaw protrusion and laterotrusion in four adult males. Muscle part lengths and intramuscular tendon lengths were calculated, and these data, with fiber-tendon ratios published previously, were used to estimate putative changes in fiber-length. During maximum jaw-opening, the largest increases in muscle length (34-83%) occurred in the medial part of the deep masseter, whereas the smallest changes occurred in the posterior-most, superficial masseter (2-19%). Smaller changes were found during movement to the ipsilateral side, than during protrusion and movement to the contralateral side. On maximum opening, putative fibers in the deep masseter lengthened up to 83%, whereas those of the superficial masseter stretched up to 72%. The masseter muscle does not stretch uniformly for major jaw movement. Jaw motion to the ipsilateral side causes little length change in any part, and the effect of tendon-stretch on estimated fiber lengths is not substantial. The stretch that occurs infers there are task-related changes in the active and passive tensions produced by different muscle regions.