Jaw-closing Movements Research Articles

Masticatory Muscles Activation and TMJ Space During Asymmetrically Loaded Jaw Closing

Masticatory muscle activation and temporomandibular joint (TMJ) load generated during asymmetrically loaded jaw closing are largely unknown. Two different strategies were developed to explain how the central nervous system (CNS) generates muscle activation patterns during motion: minimization of joint load (MJL) vs. minimization of muscle effort (MME). The aim of the present study was to investigate, experimentally, the neuromuscular strategy selected by the CNS to coordinate jaw closing in reaction to the application of an external asymmetric load. Masticatory muscle activation was measured with electromyography (EMG) and the minimum intra-articular distance (MID) was assessed by dynamic stereometry to infer joint loading. Ten healthy subjects performed jaw-closing movements against an asymmetric mandibular load set from 0.0 to 2.0 kg in 0.5-kg steps. Recordings were analyzed by exploratory and graphical statistical tools. Moreover, the observed differences in MID and EMG among the various mandibular loads were tested using non-parametric tests for repeated measures data. The ipsilateral-contralateral differences in MID and EMG of the anterior temporalis showed a significant increase (p < 0.001, p = 0.01) with increasing asymmetrical load with both joints being most heavily loaded at 1 kg. EMG signals of the masseter did not change significantly with increasing load. This study is the first to have analyzed the changes in the TMJ intra-articular space during asymmetrically loaded jaw-closing movements, not only three dimensionally and dynamically, but also combined with EMG. Asymmetrical load affected the TMJ space and masticatory muscle activation patterns, primarily resulting in an increased activation of the anterior temporalis muscle. This might suggest the involvement of a control mechanism to protect the joints from overloading. However, the results do not fully support the hypothesis of MJL nor the MME strategy.

Effects of preventing intercuspation on the precision of jaw movements.

Closing movements are among the jaw's basic physiological motor actions. During functional movements, the jaw changes position continually, which requires appropriate proprioception. However, the significance of the various proprioceptive receptors involved and how they interact is not yet fully clear. This study's main objective was to test whether preventing intercuspation (IC) for 1week would affect the precision of jaw-closing movements into IC and the functional space of habitual chewing movements (HCM). A secondary objective was to compare precision of jaw-closing movements into IC with the precision of movements into a target position (TP) far from IC. Fourteen participants' HCM and jaw-closing movements into IC were recorded on two sessions (T1 and T2) 1week apart. Between sessions, participants wore posterior bite plates to prevent IC. They also received a 10-minute training session at T1 to guide their jaw-closing movements into TP. The precision of the closing movements into IC and TP was analysed. For HCM, the vertical amplitude, lateral width and area of chewing cycles were evaluated. The precision of jaw movements into IC increased as the jaw gap decreased, but precision did not differ significantly between T1 and T2. For HCM, the vertical amplitude and area of chewing cycles increased significantly between T1 and T2. The precision of the closing trajectory into TP increased significantly during the training session. Our results confirm the excellent adaptability of the craniomandibular system, controlled by stringent motor programmes that are supported by continuous peripheral sensory input.

Improvement in three-dimensional facial configuration and jaw motion following surgical orthodontic treatment of a case with jaw deviation

The establishment of normal stomatognathic function and an acceptable facial appearance is considered an important treatment goal in orthodontics. The patient, who demonstrated posterior crossbites and jaw deviation, underwent treatment with a combined orthodontic-surgical approach utilizing maxillary transverse distraction osteogenesis, Le Fort I osteotomy, and bilateral SSRO. The smoothness of the masticatory jaw-closing movement and three-dimensional facial configuration were compared pre- and post-treatment. Correction of the posterior crossbites enabled the patient to perform smoother jaw-closing movements during chewing. Additionally, asymmetry in the morphology of the chin, cheek, and lips was reduced. These favorable changes in masticatory jaw movement kinematics and three-dimensional evaluation of the face justify the application of surgical-orthodontic treatment to improve jaw function and facial appearance.

Precision of jaw‐closing movements for different jaw gaps

Jaw-closing movements are basic components of physiological motor actions precisely achieving intercuspation without significant interference. The main purpose of this study was to test the hypothesis that, despite an imperfect intercuspal position, the precision of jaw-closing movements fluctuates within the range of physiological closing movements indispensable for meeting intercuspation without significant interference. For 35 healthy subjects, condylar and incisal point positions for fast and slow jaw-closing, interrupted at different jaw gaps by the use of frontal occlusal plateaus, were compared with uninterrupted physiological jaw closing, with identical jaw gaps, using a telemetric system for measuring jaw position. Examiner-guided centric relation served as a clinically relevant reference position. For jaw gaps ≤4mm, no significant horizontal or vertical displacement differences were observed for the incisal or condylar points among physiological, fast, and slow jaw-closing. However, the jaw positions under these three closing conditions differed significantly from guided centric relation for nearly all experimental jaw gaps. The findings provide evidence of stringent neuromuscular control of jaw-closing movements in the vicinity of intercuspation. These results might be of clinical relevance to occlusal intervention with different objectives.

Improvements in smoothness of chewing cycles in adults with mandibular prognathism after surgery: a longitudinal study

Although adults with mandibular prognathism are known to show impaired smoothness of masticatory jaw movements, it remains uncertain whether/how the surgical-orthodontic treatment is effective to improve masticatory jaw movements. The aims of this study were (i) to verify whether the surgical-orthodontic treatment of patients with mandibular prognathism improve smoothness of the chewing jaw movements post-operatively and, if so, (ii) to examine whether the improved parameters show similar quantities as those of the control subjects. The chewing jaw movements for 13 patients with mandibular prognathism were recorded at pre- and post-treatment stages. The patient group was divided into two groups: Class III(closed) showed full occlusal contact at the habitual intercuspal position, whereas Class III(open) showed inability of occlusal contact between the upper and lower anterior teeth. The control group (CG) consisted of 52 subjects having acceptably good occlusion. The normalised jerk cost (NJC), movement duration and peak velocity in jaw closing were compared. For both Class III(open) and Class III(closed) groups, statistical comparisons revealed that the NJC and movement duration were decreased after the treatment, whereas the peak velocity was increased (all P < 0·01). For the Class III(open), these quantities at the post-treatment stage did not show any significant differences compared with those for the CG. It is concluded that the smoothness of chewing jaw movements for patient with mandibular prognathism were improved after treatment, and for the patients with anterior open bite at pre-treatment, the post-operatively achieved smoothness of jaw-closing movements did not differ from those of the CG.

Smoothness of chewing jaw movements in adults with mandibular prognathism

Indices such as smoothness, movement time, peak velocity, and symmetry of the velocity profile have been shown to be effective in explaining the degree of skilfulness of human saccadic eye, limb, and jaw motions. We investigated whether adult subjects with mandibular prognathism show impaired smoothness of the masticatory jaw movements. Forty-nine adults with skeletal Class III malocclusions and 52 healthy adults with acceptably good occlusions were selected respectively as Test and Control subjects. Subjects of the Test Group were subdivided into two groups: Class III(closed) showed full occlusal contact between the upper and lower teeth at the habitual intercuspal position, whereas Class III(open) showed inability of occlusal contact between the upper and lower anterior teeth. Each subject was asked to chew a piece of chewing gum. The normalised jerk-cost (NJC), movement duration, and tangential velocity profile during jaw-closing movements were compared between groups. Test Groups showed greater NJC (P < 0·01) with longer movement duration (P < 0·01) and lower peak velocity (P < 0·01) than the Control Group did. Class III(closed) showed greater NJC (P < 0·01) with longer movement duration (P < 0·01) and lower peak velocity (P < 0·01) than Class III(open) did. Results show that the mandibular movements made by the Test Groups exhibit lower skilfulness than those made by the Control Group. The jaw movement skilfulness of the prognathic patients decreases most drastically with existence of malocclusal contact between upper and lower anterior teeth.

Masseter length determines muscle spindle reflex excitability during jaw-closing movements

The masticatory muscles are considered to be important determinants of facial form, but little is known of the muscle spindle reflex characteristics and their relationship, if any, to face height. The aim of this study was to determine whether spindle reflexes, evoked by mechanical stimulation of an incisor and recorded on the masseter muscle, correlated with different facial patterns. Twenty-eight adult volunteers (16 women; ages, 19-38 years) underwent 2-N tap stimuli to their maxillary left central incisor during simulated mastication. Reflexes were recorded during local anesthesia of the stimulated tooth to eliminate the contribution from periodontal mechanoreceptors. Surface electromyograms of the reflex responses of the jaw muscles to these taps were recorded via bipolar electrodes on the masseter muscle and interpreted by using spike-triggered averaging of the surface electromyograms. Lateral cephalometric analysis was carried out with software (version 10.5, Dolphin, Los Angeles, Calif; and Mona Lisa, Canberra, Australia). Two-newton tooth taps produced principally excitatory reflex responses beginning at 17 ms poststimulus. Correlation analysis showed a significant relationship between these muscle spindle reflexes and facial heights: specifically, shorter face heights were associated with stronger spindle reflexes. This correlation was strongest between the derived measure of masseter length and the spindle reflex strength during jaw closure (r= -0.49, P= 0.008). These results suggest that a similar muscle spindle stimulus will generate a stronger reflex activation in the jaw muscles of patients with shorter faces compared with those with longer faces. This finding might help to explain the higher incidence of clenching or bruxism in those with short faces and also might, in the future, influence the design of orthodontic appliances and dental prostheses.

Modulation of cortically induced rhythmic jaw movements in rats by stimulation of the vestibular nuclear complex

We study whether stimulation of the vestibular nuclear (VN) complex can modulate rhythmic jaw movements in rats anesthetized by urethane. Rhythmic jaw movements were induced by repetitive electrical stimulation of the orofacial motor cortex. Stimulation of the medial vestibular nucleus (MVN) during the jaw-closing phase increased the amplitude of the jaw-closing movement. (This is not a movement that continues to closure.) Stimulation of the MVN during the jaw-opening phase disturbed the rhythm of jaw movements and induced a small jaw-closing movement. Stimulation of the superior VN (SVN) and the lateral VN (LVN) during the jaw-closing phase did not affect the amplitude of the jaw-closing movement. Stimulation of the SVN and the LVN during the jaw-opening phase increased the amplitude of the jaw-opening movement, however. Stimulation of the inferior VN during the jaw-closing and the jaw-opening phase, respectively decreased the amplitude of the jaw-closing and the jaw-opening movements. Stimulation applied outside the VN did not modulate the amplitude of the jaw movements. These results imply that the VN is involved in the modulation of rhythmic jaw movements induced by stimulation of the orofacial motor cortex.

Modulation of cortically induced rhythmical jaw movements by stimulation of the red nucleus in the rat

We study whether stimulation of the red nucleus (RN) can modulate rhythmical jaw movements in rats anesthetized by urethane. Rhythmical jaw movements were induced by repetitive electrical stimulation of the two cortical masticatory areas (area A: the orofacial motor cortex; area P: the insular cortex). Stimuli applied to the RN did influence rhythmical jaw movements induced by stimulation of the A-area. Stimuli applied in the jaw-closing phase increased the amplitude of the jaw-closing movement. Stimuli applied in the jaw-opening phase disturbed the rhythm of jaw movements and induced a small jaw-closing movement. Stimuli applied to the RN did not influence rhythmical jaw movements induced by stimulation of the P-area. These results indicate that the RN is involved in the modulation of rhythmical jaw movements induced by stimulation of the A-area.

The riddle of the large loss in bite force after fast jaw-closing movements.

In unloading experiments (in which the resistance to a forceful static bite is suddenly removed), it is shown that the residual bite force (when the jaw system is arrested shortly after the unloading) is remarkably small. For example, of a 100-N initial bite force, only 18 N is left after a jaw travel distance of 5.0 mm. The present experiments were designed to study whether the magnitude of the low residual bite force is dependent on the initial bite force, the initial degree of mouth opening, and the distance of jaw travel. Furthermore, we analyzed whether the low magnitude of the residual force can be attributed to reflex events of the jaw muscles or to the force-length properties of the jaw-closing muscles. It was found that the residual forces are largely dependent on the distance of jaw travel and are barely sensitive to variations in initial mouth-opening. The relative residual forces are independent of the magnitude of the initial bite force. The maximum residual forces are on the order of 25% of the initial bite force after a jaw travel of 4.5 mm. The low values of the residual forces cannot be attributed to reflex events, because it took about 80 ms for the masseter muscles to decrease their force to a 50% level after their excitation was switched off. Furthermore, it was shown that the force-length properties of the jaw-closing muscles are not responsible for the small values of the residual forces, since over the trajectories used in the present experiments, the sarcomere lengths of the jaw-closing muscles were beyond their optimum. It is suggested that the low residual forces are brought about by (1) a non-uniform sarcomere behavior of the jaw-closing muscles when contracting, or (2) a long-lasting change in the myofilament system of the closing muscles induced by the sudden shortening of muscle fibers.

The jaw open-close movements predicted by biomechanical modelling

The aim of this study was to analyse unloaded jaw-opening and jaw-closing movements in humans. For this purpose a dynamical 6-degrees-of-freedom mathematical model of the human masticatory system was developed. It incorporated morphology, muscle architecture and dynamical muscle properties. Various symmetrical jaw-opening and jaw-closing movements were simulated based upon different muscle activation schemes. It was found that the balance between swing and slide of the mandibular condyle at the onset of a jaw-opening movement was predominantly dependent on the level of activation of the digastric and inferior lateral pterygoid museles. The level of activation of the temporalis muscle parts was of critical importance for the jaw-closing movements. The amount of jaw opening was limited by the passive forces of the jaw-closing muscles. In contrast, the influence of the passive forces of the jaw-opening muscles on the jaw-closing movement was neglectable. Throughout the movements the temporomandibular joints remained loaded. The average torques generated by the jaw-opening or jaw-closing muscles with respect to the centre of gravity of the lower jaw had similar orientations and can be considered to be responsible for joint stabilization. The average direction of their lines of action, however, was about opposite, and this can be considered as the major discriminant between a movement in opening or closing direction.

Contribution of the digastric muscles to the control of bite force in man

The contribution of the (co-contracting) digastric muscles to the rapid decline in bite-force magnitude after unloading of a static bite was investigated by asking participants to perform two different biting tasks with sudden unloading, and correlating the degree of co-contraction of the digastrics (as derived from their electromyograms) with the impact force, the impact velocity (as measured after a travel distance of 5 mm), and the residual force when the jaw system was in static conditions again after the impact. Co-contraction of the digastrics was varied by asking participants to perform the biting task while controlling bite force (force-controlled experiments) or jaw position (position-controlled experiments). In half of the experiments, participants co-contracted their digastrics more strongly in the position-controlled than the force-controlled experiments. However, there was no clear relation between the level of co-contraction and the magnitude of the impact force, the impact velocity and the residual force. The results imply that co-contraction of the digastric muscles is not sufficient to explain the reduction in bite force and the low impact velocity after an unexpected jaw-closing movement. Two other possible mechanisms that reduce forces in an unloaded jaw system are: (1) force-velocity properties of the activated jaw muscles in conjunction with creep of the aponeurotic sheets of the jaw muscles, resulting in a slow partial recovery of the biting force after impact; (2) force-length properties of jaw-opening muscles, an activity not recorded here.

Influence of the Dynamical Properties of the Human Masticatory Muscles on Jaw Closing Movements

Influence of the Dynamical Properties of the Human Masticatory Muscles on Jaw Closing Movements

Biomechanical Analysis of Jaw-closing Movements

This study concerns the complex interaction between active muscle forces and passive guiding structures during jaw-closing movements. It is generally accepted that the ligaments of the joint play a major role in condylar guidance during these movements. While these ligaments permit a wide range of motions, it was assumed that they are not primarily involved in force transmission in the joints. Therefore, it was hypothesized that muscle forces and movement constraints caused by the articular surfaces imply a necessary and sufficient condition to generate ordinary jaw-closing movements. This hypothesis was tested by biomechanical analysis. A dynamic six-degrees-of-freedom mathematical model of the human masticatory system has been developed for qualitative analysis of the contributions of the different masticatory muscles to jaw-closing movements, it was found that the normally observed movement, which includes a swing-slide condylar movement along the articular eminence, can be generated by various separate pairs of masticatory muscles, among which the different parts of the masseter as well as the medial pterygoid muscle appeared to be the most suitable to complete this action. The results seem to be in contrast to the general opinion that a muscle with a forward-directed force component may not be suitable for generating jaw movements in which the condyle moves backward. The results can be explained, however, by biomechanical analysis which includes not only muscle and joint forces as used in standard textbooks of anatomy, but also the torques generated by these forces.

The effect of bilateral cold block of the primate face primary somatosensory cortex on the performance of trained tongue-protrusion task and biting tasks.

1. Studies using ablation, intracortical microstimulation (ICMS) and surface stimulation, and single-neuron recordings have suggested that the primate primary somatosensory cortex (SI) may play an important role in movement control. Our aim was to determine whether bilateral inactivation of face SI would indeed interfere with the control of tongue or jaw-closing movements. 2. Effects of reversible inactivation by cooling of face SI was investigated in two monkeys trained to perform both a tongue-protrusion task and a biting task. The cooling experiments were carried out after the orofacial representation within SI was identified by systematically defining the mechanoreceptive field of single neurons recorded in face SI. The deficits in the tongue or jaw-closing movement were evaluated by the success rates for the monkeys' performance of both tasks and by the force and electromyographic (EMG) activity recorded from the masseter, genioglossus, and digastric muscles associated with the tasks. 3. During bilateral cooling of face SI, there was a statistically significant reduction in the success rates for the performance of the tongue-protrusion task in comparison with control series of trials while the thermodes used to cool face SI were kept at 37 degrees C. Detailed analyses of force and EMG activity showed that the principal deficit was the inability of the monkeys to maintain a steady tongue-protrusive force in the force holding period during each trial and to exert a consistent tongue-protrusion force between different trials. The task performance returned to control protocol levels at 4 min after commencement of rewarming. 4. Identical cooling conditions did not significantly affect the success rates for the performance of the biting task. Although the extent of the deficit was not severe enough to cause a significant reduction in successful rates for the biting task, cooling did significantly affect the ability of the monkeys to maintain a steady force in the holding period during each trial and to exert a consistent force between different trials. In one monkey the success rate of the biting task was also not affected by bilaterally cooling of face SI with a pair of larger thermodes placed on the dura over most of the face SI, face primary motor cortex (face MI), and adjacent cortical regions.(ABSTRACT TRUNCATED AT 400 WORDS)

Functional properties of single neurons in the face primary motor cortex of the primate. I. Input and output features of tongue motor cortex.

1. We have recently demonstrated that reversible, cooling-induced inactivation of the face motor cortex results in a severe impairment in the ability of monkeys (Macaca fascicularis) to perform a tongue-protrusion task but produces only relatively minor effects on the performance of a biting task by the same monkeys. To establish a neuronal correlate for these different behavioral relations, the present study has detailed the afferent input and intracortical microstimulation (ICMS)-defined output features of a population of face motor cortical neurons, and in a subsequent study we have documented the activities of the same population of neurons during the performance of the tongue-protrusion and biting tasks. 2. Of the 231 single neurons recorded within the face motor cortex, 163 were located at sites from which ICMS (less than or equal to 20 microA) could evoke tongue movements (i.e., "tongue-MI" sites) at the lowest threshold for eliciting orofacial movements. The remainder were located at sites from which ICMS evoked jaw movements ("jaw-MI" sites), face movements ("face-MI" sites), or at a few sites, tongue movements and, at the same threshold intensity, either a jaw movement or a facial movement. 3. We confirmed the general organizational features of the face motor cortex that have been defined in previous studies, but we documented in detail the organizational features for tongue-MI. Thus we found that tongue movements were well represented, whereas jaw-closing movements were poorly represented; the representations for face, jaw, and tongue movements were overlapped; the same ICMS-evoked tongue movement could be multiply represented within tongue-MI; tongue-MI was characterized by a prominent input from superficial mechanosensory afferents, whereas there was little evidence for deep input; a close spatial match was found between ICMS-defined motor output and somatosensory afferent input for tongue-MI. 4. A variety of tongue movements could be evoked by ICMS at tongue-MI sites and were categorized into protrusion, retrusion, laterally directed, and other types of tongue movement. Low-threshold (i.e., less than or equal to 5 microA) ICMS-defined tongue-MI sites, which were considered to represent "efferent zones" projecting relatively directly to motoneurons, were reconstructed three dimensionally to provide insights into the spatial organization of tongue-MI. Examples of each of the four low-threshold efferent-zone categories were usually found throughout the ICMS-defined tongue-MI without any apparent preferential distribution. Furthermore, different low-threshold efferent-zone categories had close spatial relationships to each other in cortex.(ABSTRACT TRUNCATED AT 400 WORDS)

Functional properties of single neurons in the face primary motor cortex of the primate. II. Relations with trained orofacial motor behavior.

1. The previous paper has described in detail the input and output features of single neurons located at sites within primate face motor cortex from which intracortical microstimulation (ICMS, less than or equal to 20 microA) evoked tongue movements at the lowest threshold ("tongue-MI" sites); for comparative purposes, we also reported on the input and output features of a smaller number of neurons recorded at sites from which ICMS could evoke jaw movements ("jaw-MI" sites), facial movements ("face-MI" sites), or, at a few sites, tongue movements and, at the same threshold intensity, either a jaw movement or a facial movement. 2. Our findings of an extensive and diverse representation of sites within face motor cortex of monkeys for the generation of elemental components of tongue movement, and the relatively few sites from which jaw-closing movements could be evoked, were consistent with our recent observations that reversible, cooling-induced inactivation of the face motor cortex severely impaired the performance by monkeys of a tongue-protrusion task but had only relatively minor effects on the performance of a biting task. In an attempt to establish a neuronal correlate for these different behavioral relations, the present study has documented the task-related activities of those single neurons that were characterized in the previous paper in terms of afferent input and ICMS-defined output features. 3. Each task required the development and maintenance by each monkey of a fixed force level for a minimum period of time to obtain a fruit-juice reward. During one or both of these tasks, we characterized the activities of 231 single face motor cortical neurons that were located at the above-mentioned ICMS-defined sites. Neurons were said to be related to a particular task if they showed statistically significant differences in firing rates during the task in comparison with a control pretrial period (PTP). 4. In tongue-MI, there was a significantly higher proportion of neurons (63% of 156 neurons tested) that were related to the tongue-protrusion task than to the biting task (15% of 65). However, in jaw-MI the proportion of neurons that were biting task-related (63% of 19) was significantly higher than the proportion related to the tongue-protrusion task (11% of 9); the proportion of biting task-related neurons at ICMS-defined jaw-closing sites was also higher than that at jaw-opening sites.(ABSTRACT TRUNCATED AT 400 WORDS)

Effects of reversible inactivation by cooling of the primate face motor cortex on the performance of a trained tongue-protrusion task and a trained biting task

1. Intracortical microstimulation (ICMS) and surface stimulation studies of primate face motor cortex have shown an extensive representation within face motor cortex devoted to movements of the tongue and face; only a very small representation for jaw-closing movements has ever been demonstrated. These data suggest that face motor cortex plays a critical role in the generation of tongue and facial movements but is less important in the generation of jaw-closing movements. Our aim was to determine whether disruption of primate face motor cortical function would indeed interfere with the generation of tongue movements but would not interfere with the generation of jaw-closing movements. 2. The face motor cortex was reversibly inactivated with the use of cooling in two monkeys that were trained to perform both a tongue-protrusion task and a biting task. Recording of single neuronal activity in the cortex beneath the thermode confirmed the reversible inactivation of the cortex. Each task involved a series of trials in which the monkey was required to produce a preset force level for a 0.5-s force holding period; the monkey received a fruit-juice reward if it successfully completed a task trial. Cooling of the ICMS-defined face motor cortex was achieved bilaterally or, in one experiment, unilaterally by circulating coolant through thermodes placed either on intact dura overlying face motor cortex in both monkeys or directly on the exposed pia in one of the monkeys;thermode temperature was lowered to 3-5 degrees C during cooling. Electromyographic (EMG) recordings were also made from masseter, genioglossus, and digastric muscles. 3. During bilateral cooling of the thermodes on the dura overlying the face motor cortex, there was a significant reduction in the success rates for the performance of the tongue-protrusion task in comparison with control series of trials (i.e., precool and postcool) in which the thermodes were kept at 37 degrees C. Quantitative analyses of force and EMG activity showed that the principal deficit was an inability of each monkey to exert sufficient force with its tongue for a sufficient length of time onto the tongue-protrusion task transducer; this deficit was paralleled by a reduction in the level of genioglossus and digastric EMG activity. At 4 min after commencement of rewarming, task performance had returned to control, precool levels.(ABSTRACT TRUNCATED AT 400 WORDS)

Human jaw movement in mastication and speech

The study of jaw movement in humans is a primary source of information about the relationship between voluntary movement and more primitive motor functions. This study focused on the geometric form of the velocity function, as measured by linear voltage displacement transducer. Movement amplitudes, maximum velocities and durations were greater in mastication than in speech. Nevertheless, there were detailed similarities in the shape of the normalized velocity functions. In jaw-closing movements, the normalized functions were similar in form over differences in rate, movement amplitude (speech movements) and the compliance of the bolus (mastication). In opening movements, the functions for mastication and speech were again similar over differences in amplitude and compliance. However, they differed in shape for fast and slow movements. Normalized acceleration and deceleration durations were approximately equal in rapid movements, whereas, for slower movements, deceleration took substantially longer than acceleration.

A hierarchy of neural control of mastication in the rat

In rats anaesthetized with ketamine, rhythmic jaw-opening and jaw-closing movements were induced by palatal stimulation. The two masseter muscles (jaw-closing) and the four digastric muscles (jaw-opening) were fitted with electrodes, which could be used either for electrical stimulation or for recording electromyographic responses. Electrical stimulation of the masseters in the phase when the digastrics were the contracting muscles, caused responses in the digastrics. The amplitude of these responses was dependent on whether the stimulated masseters were active or not. The responses in digastric persisted when contraction of the masseters during stimulation was prevented by dantrolene sodium but they disappeared when the masseteric nerves were blocked with xylocain. The responses in digastric are thus reflexes from stimulating afferent fibres in the masseteric nerves. Likewise, electrical stimulation of the four digastrics in the phase when the masseters were contracting, caused responses in the masseters. The amplitude of these responses, however, was independent of the state of activity of the stimulated digastrics. Furthermore, the responses in masseter disappeared when contraction of the digastrics was prevented by dantrolene sodium; but they persisted when the digastric nerves were blocked with xylocain, provided the digastrics continued to twitch to the electric stimuli. The responses in masseter are thus reflexes in masseter caused by mechanical stretch transmitted from the digastric twitches. In the rhythmic preparation, prevention of contraction of the masseters or digastrics by dantrolene sodium or xylocain leaves the overall frequency and amplitude of the evoked rhythmic activity unchanged. In the quiescent preparation, without rhythmic activity, stimulation of the digastrics still causes the response in the masseters, but the reflex onto digastrics from electrical stimulation of the masseters is inactive. It is proposed that oral activity is controlled by two relatively independent systems. The first “programme-oriented” system generates the basic movements. The second “environment-oriented” system has access to the programme for correcting for perturbations. Muscles poorly supplied with muscle spindles (the digastrics) act supposedly as pure effectors. Corrective alterations in their activity after a perturbation are mainly prompted by muscle receptors in their antagonists.