A cineradiographic and electromyographic study of mastication in Tenrec ecaudatus

Regular chewing was studied in the specialized Malagasy insectivore Tenrec ecaudatus with the aid of precisely correlated electromyography of the main adductors, digastrics, and two hyoid muscles and cineradiography for which metallic markers were placed in the mandibles, tongue, and hyoid bone. During the power stroke the body of the mandible moves dorsally and medially. The medially directed component of movement at this time is greatly increased by simultaneous rotation of the mandible about its longitudinal axis. The highly mobile symphysis, spherical dentary condyle, loss of superficial masseter muscle and zygoma, and the simplified zalamnodont molars all appear to be related to the large amount of mandibular rotation that occurs during occlusion. The balancing side lateral pterygoid muscle (inferior head) apparently shifts the working side mandible laterally during the last part of opening and the first part of closing. The working side temporalis and the superficial masseter muscle are both responsible for the shift back to the midline. The temporalis is usually active to the same extent on the working and balancing sides during the power stroke. The level of activity (amplitude) of the temporalis and duration of the power stroke increase with harder foods. Whenever soft foods are chewed, the superficial masseter is only active on the working side; whenever foods of increasing hardness are chewed, its level of activity on the balancing side increases to approach that of the working side. Mandibular rotation is greatly reduced when hard foods are chewed.     

Food transport through the anterior oral cavity in macaques

Intraoral transport, the movement of food or liquid through the oral cavity and oropharynx, is a major component of feeding behavior. Stage I transport, transport through the oral cavity prior to mastication, has been described for several mammals (Franks et al.: Arch. Oral Biol. 30:539, 1985; Hiiemae and Crompton: Hildebrand et al. (eds.): Functional Vertebrate Morphology, Cambridge, MA, Belknap Press, 1985). Previous work (Franks et al.: Am. J. Phys. Anthropol. 65:275, 1984) indicated that this was not a significant behavior in macaques in a laboratory setting, because food was ingested directly to the region of the cheek teeth. Although relatively infrequent in a captive situation, stage I transport does occur in long-tailed macaques through a mechanism similar to other mammals, but also subject to unique aspects of primate anatomy. Transport takes several cycles during which the food moves back and forth in an anterior/posterior direction, due to tongue movements. Because anthropoid primates lack the pronounced rugae that in other mammals prevent the anterior displacement of a bolus, stage I transport uses the rounded arch of the upper, anterior dentition to hold the food during the forward movement of the tongue. During the final cycle of transport, a pronounced twisting of the tongue, along a midline anteroposterior axis helps funnel the food item toward the postcanine teeth for subsequent mastication. This twisting, which was described in humans by Abd-El-Malek (J. Anat. 100:215, 1955) but not within the context of jaw movement, occurs prior to the closing phase of the jaw cycle.     

Twist and chew: three-dimensional tongue kinematics during chewing in macaque primates

Three-dimensional (3D) tongue movements are central to performance of feeding functions by mammals and other tetrapods, but 3D tongue kinematics during feeding are poorly understood. Tongue kinematics were recorded during grape chewing by macaque primates using biplanar videoradiography. Complex shape changes in the tongue during chewing are dominated by a combination of flexion in the tongue''s sagittal planes and roll about its long axis. As hypothesized for humans, in macaques during tongue retraction, the middle (molar region) of the tongue rolls to the chewing (working) side simultaneous with sagittal flexion, while the tongue tip flexes to the other (balancing) side. Twisting and flexion reach their maxima early in the fast close phase of chewing cycles, positioning the food bolus between the approaching teeth prior to the power stroke. Although 3D tongue kinematics undoubtedly vary with food type, the mechanical role of this movement—placing the food bolus on the post-canine teeth for breakdown—is likely to be a powerful constraint on tongue kinematics during this phase of the chewing cycle. The muscular drivers of these movements are likely to include a combination of intrinsic and extrinsic tongue muscles.     

Preweaning feeding mechanisms in the rabbit.

Muscle contraction patterns and mandibular movements of infant rabbits during suckling and chewing were compared. Oral muscle activity was recorded by fine-wire electromyography, while jaw movements and milk bottle pressure were registered. Suckling and mastication have a comparable cycle duration and share a common pattern of oral muscle activity which consists of a succession of a jaw closer burst, during which the jaw closes and undergoes a power stroke (in mastication), a suprahyoid burst with a stationary or slightly opening jaw and a digastric burst with fast jaw opening (the power stroke of suckling). Compared to suckling, mastication shows decreased jaw opener activity, increased jaw closer activity, development of jaw closing activity in the lateral pterygoid, and increased asymmetry in the masseter by development of a new differentiated motor pattern on the working side. The study shows that the suckling motor pattern enables the infant rabbits to change to chewing with just a few modifications.     

Morphological and videofluorographic study of the hyoid apparatus and its function in the rabbit (Oryctolagus cuniculus)

The anatomy of the hyoid apparatus and positional changes of the hyoid bone during mastication and deglutition are described in the New Zealand White rabbit (Oryctolagus cuniculus). A testable model is constructed to predict the range of movement during function of the hyoid, a bone entirely suspended by soft tissue. Frame-by-frame analysis of a videofluorographic tape confirms the accuracy of the prediction through observation of hyoid bone excursion during oral behavior. During chewing, translation of the hyoid bone is diminutive and irregular, lacking a clearly discernible path of excursion. However, some movements of the hyoid occur with regularity. During fast opening, anterodorsal movement of the hyoid is interrupted with an abrupt posteroventral depression when the bolus is moved posteriorly toward the cheek teeth by the tongue. This clockwise rotation (when viewed from the right side) of the hyoid accompanies jaw opening and is reversed (posteroventral movement) for the jaw closing sequence. Lateral movements of the hyoid may be slightly coupled to mandibular rotation in the horizontal plane. The findings suggest that the hyoid bone maintains a relatively static position during the dynamics of chewing. The primary function would be to provide a stable base for the movements of the tongue. Another possible function would be to control the position of the larynx within the pharyngeal cavity. Some characteristic features of the rabbit hyoid apparatus may be consequential to relatively erect posture and a saltatory mode of locomotion.     

Biomechanics of the masticatory apparatus of Pteropus giganteus (Megachiroptera)

Jaw mechanics in Pteropus were studied by means of a three-dimensional model. The model included several parameters of muscle architecture, combined with quantified movement and electromyographical data. Estimates of the nature of the applied forces that act upon the mandible during a chewing cycle, and subsequent estimates of reaction forces at the bite point and joints during the powerstroke, were thus obtained for different food consistencies. The resultant muscle force (relative to the palate) shifts from upward and slightly backward at large gapes to upward and markedly backward at the end of closing. The resultant simultaneously moves anteriorly. During the powerstroke it retains a constant position and orientation along the thickened anterior edge of the coronoid process. The early stages of opening are guided by the slope of the teeth and mandibular fossa; during the remaining part of opening the working line of the resultant crosses the skull behind the joint and thus acquires an opening moment. The bite force has downward and forward components, and a slight transverse component. For a given applied muscular force its magnitude is larger in more posteriorly positioned bite points. Both joints are loaded, the contralateral one more than the ipsilateral. Food consistency affects magnitude and orientation of the applied force, and hence, magnitude and orientation of the bite force and magnitude of the joint reaction forces. The magnitude of masseter activity relative to temporalis activity appears to be the key factor for the orientation of the bite force, and hence for the mechanical optimal position of the food. The adaptive value of the general topography of the masticatory muscles in Pteropus is discussed.     

Mandibular movement and muscle activity during mastication in the guinea pig (Cavia porcellus)

Optoelectronic analysis of mandibular movement and electromyography (EMG) of masticatory muscles in Cavia porcellus indicate bilateral, unilateral, and gnawing cycles. During bilateral and unilateral cycles, the mandibular tip moves forward, lateral, and down during the lingual phase of the power stroke to bring the teeth into occlusion. EMG activity is generally asymmetric, with the exception of activity of the temporalis muscle during bilateral cycles. During gnawing cycles, the mandible moves in an anteroposterior direction that is opposite that during bilateral and unilateral chew cycles. Bilateral and unilateral cycles of pellets were significantly longer than carrot. With the exception of the width of bilateral cycles, the magnitude of cycle width, length, and height during the mastication of carrots was greater than that during the mastication of pellets. Significant differences exist between EMG durations during mastication of pellets and carrots. The lateral pterygoid displays continuous activity during gnawing cycles. Significant differences also exist in the durations of EMG activity between the working and balancing side during all three cycle types. High level activity of balancing side temporalis and anterior belly of digastric (ABD) during bilateral cycles occurs during rotation and depression of the mandible during the power stroke. The temporalis apparently provides a ?braking”? or compensatory role during closing and power strokes. Differences between Cavia masticatory patterns and those shown by Rattus and Mesocricetus are apparently due to differences in dental morphology, occlusal relationships, and, possibly, the poorly developed temporalis in Cavia. The large number and wide diversity of rodent groups afford students of mammalian mastication an opportunity to investigate and compare different masticatory specializations.     

Observations on rhinoceros tongue morphology

The rhinoceros tongue is distinguished among perissodactyl organs by its sagittally divided intermolar eminence crowned by a dextral and a sinistral cluster of circumvallate papillae. These structures were originally noted in the Indian rhinoceros by Owen (1852) but have since remained unmentioned for this, or for any other rhinoceros form. They are here described for specimens of the Indian, African White and African Black rhinoceroses. The structural and functional nature of the intermolar eminence is established and this entity is distinguished from an hitherto unrecognized topographical component of the tongue, designated the pars elliptica linguae. The lingual morphology of the three rhinoceros forms is summarised and certain intergeneric differences are noted.     

A dynamic model of jaw and hyoid biomechanics during chewing

Our understanding of human jaw biomechanics has been enhanced by computational modelling, but comparatively few studies have addressed the dynamics of chewing. Consequently, ambiguities remain regarding predicted jaw-gapes and forces on the mandibular condyles. Here, we used a new platform to simulate unilateral chewing. The model, based on a previous study, included curvilinear articular guidance, a mobile hyoid apparatus, and a compressible food bolus. Muscles were represented by Hill-type actuators with drive profiles tuned to produce target jaw and hyoid movements. The cycle duration was 732 ms. At maximum gape, the lower incisor-point was 20.1mm down, 5.8mm posterior, and 2.3mm lateral to its initial, tooth-contact position. Its maximum laterodeviation to the working-side during closing was 6.1mm, at which time the bolus was struck. The hyoid's movement, completed by the end of jaw-opening, was 3.4mm upward and 1.6mm forward. The mandibular condyles moved asymmetrically. Their compressive loads were low during opening, slightly higher on the working-side at bolus-collapse, and highest bilaterally when the teeth contacted. The model's movements and the directions of its condylar forces were consistent with experimental observations, resolving seeming discordances in previous simulations. Its inclusion of hyoid dynamics is a step towards modelling mastication.     

Models of tongue movement in the walrus (Odobenus rosmarus)

Three hypothetical models of tongue movement of the walrus during suction feeding are examined. These models encompass the entire range of simple tongue retraction movements possible by examining 1) movement of the tongue directly to the rear following the curvature of the palate, 2) to the rear and ventrally in a straight line, and 3) ventrally in a straight line. The percent of muscular force available from the hyoglossus, genioglossus, and styloglossus that could be applied toward retraction as predicted by each model is calculated. The resistance that the tongue would provide during retraction is calculated using projected tongue areas and is combined with the above data from the muscles to provide an estimate of the percent of the total available force that is needed to retract the tongue for each model. A separate examination of the direction of tongue-induced wear striations on the palatal and lingual aspects of the teeth is used to help support or reject the three models. The model where the tongue is moved directly to the rear is supported by studies of both muscle force and tooth wear. In the mammalian groups that were compared to the walrus, there is a great deal of interspecific variation in movements of the tongue during suction feeding; no two groups can be considered to have identical stereotyped tongue movements.     

Mastication in springhares,Pedetes capensis: A cineradiographic study

Analysis of lateral and dorsoventral radiographic films shows that ingestion, transport, and mastication in Pedetes capensis (Rodentia) are cyclic and their movement patterns are essentially similar for the three food types offered. During the ingestion cycle, closing of the mouth is accompanied by a backward translation of the condyles, so that movement is predominantly orthal. During the opening stage, the extent of the anterior condylar translation is smaller. As a result the mandibular incisors move ventrally and posteriorly. During the ingestion cycles, food is transported to the back of the tongue, with the transverse rugae and the folds of the upper lip playing important roles. Springhares show a bilateral masticatory pattern; food is chewed on both sides simultaneously. During chewing, the condyles lie in their most forward position at maximum opening of the mouth. The mouth is closed by rotation of the lower jaw around the temporomandibular joint coupled with posterior condylar translation. At the beginning of the slow-closing stage, the upward rotation of the mandible slows and the jaw slowly shifts forward. During the grinding stage, the mandible is shifted forward with both toothrows in occlusion. During the opening stage, the jaw returns to its starting position. Comparison of kinematic and anatomical data on rodent mastication suggests that some dental characteristics form the most important factors regulating the masticatory pattern and consequently allow reasonably reliable prediction of rodent masticatory patterns.     

Mastication in the tuatara,Sphenodon punctatus (reptilia: Rhynchocephalia): Structure and activity of the motor system

The masticatory pattern of Sphenodon punctatus, the sole remaining rhynchocephalian, now restricted to islands off the coast of New Zealand, has been analyzed by detailed anatomy, cinematography, cinefluoroscopy, and electromyography. Food reduction consists of a closing, crushing bite followed by a propalineal sliding of the dentary row between the maxillary and palatine ones. The large, fleshy tongue can be protruded to pick up small prey, and also plays a major role in prey manipulation. The rotational closing movement of the jaw, supporting the basic crushing movement, is induced by the main adductor musculature. It is followed by a propalineal anterior displacement relying heavily on the action of the M. pterygoideus. The fiber lengths of the several muscles reflect the extent of shortening. The most obvious modification appears in the M. pterygoideus, which contains a central slip of pinnately arranged short fibers that act a period different from that of the rest of the muscle; their action increases the power during the terminal portion of the propalineal phase. This also allows the animal to use its short teeth in an effective shearing bite that cuts fragments off large prey. The action of single cusped dentary teeth acting between the maxillary and palatine tooth rows provides a translational crushing-cutting action that may be an analog of the mammalian molar pattern. However, this strictly fore-aft slide does not incorporate capacity for later development of lateral movement.     

Quantitative assay of electromyograms during mastication in domestic cats (Felis catus)

Mastication has been studied by cinematography with synchronized electromyography (computer quantified and analyzed), while unanesthetized, freely feeding cats (Felis catus) were reducing equivalent-sized chunks of raw and cooked beef and cooked chicken. Cats reduce food on one side at a time, and their chewing cycles show both horizontal and anteroposterior deflections. Food objects are shifted from side to side by lateral jerks of the head and movements of the tongue. During the opening phase, the lower jaw is rotated relatively straight downward, and the digastric muscles are active in bilateral symmetry. Near the end of opening, the head jerks upward, both zygomaticomandibulares start to fire, and opening acceleration of the mandible decreases. Closing starts with horizontal displacement of the mandibular canines toward the working side, accompanied by asymmetrical activities from the working side deep temporalis and the balancing side medial pterygoid, as well as a downward jerk of the head. As closing proceeds, the mandibular canines remain near the working side and the working side zygomaticomandibularis and deep masseter are very active. Near the end of closing, the mandibular canine on the working side moves toward the midline, and adductors, digastrics, and lateral pterygoids of both sides are active. The adductors of the working side are generally more active than those of the balancing side. During a reduction sequence, the number and shape of the masticatory cycles, as well as movements of the head, during a reduction sequence are affected significantly by food type. As reduction proceeds, the duration of bite and the muscular activity (as characterized by number and amplitude of spikes) change significantly among muscles of the working and balancing sides. The adductors of the working side are generally most active when cats chew raw beef, less for cooked beef, and least for cooked chicken. In general, the adductor activity reflects food consistency, whereas that of the digastrics and lateral pterygoids reflects more the vertical and lateral displacements of the mandible. Statistical analysis documents that the methods of electrode insertion and test give repeatable results for particular sites in different animals. Thus, it should be possible to compare these results with those produced while other mammalas are masticating.     

Time analysis of hard and soft bolus processing

Objective: Clinical observations and mathematical models show that dental implants are influenced by the magnitude of loading. Therefore, the knowledge of mandible movement during mastication is important to assess occlusal and masticatory force vectors. The purpose of this study was to detect the path of movement of the lower jaw and to distinguish stages of mastication, duration of bolus processing and peak amplitude of mastication. Method: Motion analysis was used to record three-dimensional mandible movements. Individualized sensors were rigidly attached to the mandible of 51 study participants. At the beginning of the measurement, all subjects were asked to move the mandible in extreme positions (maximal opening and maximal lateral movements). Then, each subject masticated a bite of hard and soft food. Duration of bolus mastication and peak amplitude of mastication movement in mesio-distal, cranio-caudal and vestibulo-oral axes related to peak amplitude of marginal movements were evaluated for each subject. The chewing record of each subject was divided into three phases (chopping, grinding and swallowing), and the duration of mastication and number of closing movements were evaluated. Results: The findings of this pilot study suggest that masticatory movements vary in individuals. Bolus character influences the process duration, but not the frequency of closing movements. Neither gender nor age had any influence on either the time or frequency of bolus processing. Conclusion: Relationships to directions and magnitudes of acting chewing force should be more precisely examined since transversally acted forces during grinding are important factors in tooth/implant overloading.     

A cinefluorographic study of mandibular movement during feeding in the rat (Rattus norvegicus)

The anatomy of the masticatory apparatus, and particularly of the mandibular joints, has led to the view that mandibular movement in the Rodentia is predominantly propalinal, or forwards and backwards in direction. As part of an investigation into the mechanism of function of the mandibular joints in these animals, the feeding behaviour of "August" strain and "Wistar" rats was examined by cinephotography and cinefluorography. The rats were trained to feed on barium sulphate impregnated biscuit and animal cake and to drink radio-opaque liquids. Cinefluorographic recordings of ingestion, mastication, deglutition and of drinking were taken in both the lateral and dorso-ventral projections.
Analysis of the recordings has shown a fundamental separation of ingestive and masticatory activity in the rat, which can be attributed to the morphology of the jaws and particularly to the disparity in the lengths of the mandibular and maxillary diastemas. To bring the incisor teeth into occlusion for ingestion, the mandible is brought forward through the rest position and the condyle into articulation with the anterior part of the fossa. In mastication the condyle is moved backwards to bring the molar teeth into occlusion and the condyle into articulation with the posterior articular facet on the fossa. Once the mandible has been moved into the appropriate position for either ingestion or mastication and deglutition, the movements involved in the separation or chewing of the food are cyclical with combined horizontal and transverse movements as well as the fundamental vertical movement acting to open and close the mouth. The basic movement of ingestion carries the mandibular incisors upwards and forwards across the lingual surfaces of the maxillary incisors, so separating the bite. The grinding stroke of mastication is a horizontal movement carrying the mandibular molars anteriorly across the maxillary.     

Mechanism of intraoral transport in macaques

All mammals have the same divisions of cyclic movement of tongue and hyoid during mastication: a protraction or forward phase that begins at minimum gape, and a retraction or return phase. Nonanthropoid mammals transport food from the oral cavity to the oropharynx during the return phase; food on the dorsal surface of the tongue moves distally while the tongue is retracted. Macaques, however, transport food during the protraction phase of tongue/hyoid movement. Food is squeezed posteriorly by contact between the tongue surface and the palate anterior to the food. This mechanism of transport is occasionally seen in nonanthropoid mammals when they are transporting liquids from the oral cavity to the oropharynx. It has, however, not been seen when these mammals transport solid food. One morphological basis for this difference is the reduction in height of the rugae of the palate of macaques. In most mammals these rugae are pronounced ridges that are able to hold food in place during protraction as the tongue slides forward beneath the food. Anthropoids and other mammals differ in the way they store food prior to swallowing. When macaques transport food to the oropharynx, usually they swallow in the next cycle, but always in the next 2 or 3 cycles. Most mammals transport and store food in the oropharynx for several cycles before a swallow clears that region of food. This behavior is correlated with differences in morphology of the oropharynx; anthropoids have reduced valleculae, the area in which other mammals store food prior to swallowing.     

Biplanar X-ray motion analysis of the lower jaw movement during incisor interaction and mastication in the beaver (Castor fiber L. 1758)

The first biplanar X-ray motion analysis of mastication and food processing for Castor fiber is presented. While particles are chipped off interaction of incisors involves variable movements of the lower mandible and thus incisors. After jaw opening the tip of the lower incisors can reach different positions anteriorly of the upper incisors. Then the mandible moves upwards and backwards and brings the tips of the incisors into contact. The lower incisors slide along the wear facet of the upper to the ledge when the cheek teeth occlude. The glenoid fossa and lower jaw condyle are in close contact during incisor contact and no transverse movements are observed. Mastication involves interaction of the cheek teeth with no contact of the incisors. When the cheek teeth are in occlusal contact the mandible is moved forward and transverse, or mediolateral. In consecutive power strokes the jaw is moved alternately to the right and left side. When the jaw opens it is brought into a more central but not totally centred position. During mastication the condyles are positioned posteriorly to the glenoid allowing lateral movement of the mandible. The lateral movement is particularly noticeable in the anterior part of the mandible. With the lateral movements of the incisors one glenoid has to move posteriorly, the other anteriorly.     

Correspondence between laryngeal vocal fold movement and muscle activity during speech and nonspeech gestures.

To better understand the role of each of the laryngeal muscles in producing vocal fold movement, activation of these muscles was correlated with laryngeal movement during different tasks such as sniff, cough or throat clear, and speech syllable production. Four muscles [the posterior cricoarytenoid, lateral cricoarytenoid, cricothyroid (CT), and thyroarytenoid (TA)] were recorded with bipolar hooked wire electrodes placed bilaterally in four normal subjects. A nasoendoscope was used to record vocal fold movement while simultaneously recording muscle activity. Muscle activation level was correlated with ipsilateral vocal fold angle for vocal fold opening and closing. Pearson correlation coefficients and their statistical significance were computed for each trial. Significant effects of muscle (P < or = 0.0005) and task (P = 0.034) were found on the r (transformed to Fisher's Z') values. All of the posterior cricoarytenoid recordings related significantly with vocal opening, whereas CT activity was significantly correlated with opening only during sniff. The TA and lateral cricoarytenoid activities were significantly correlated with vocal fold closing during cough. During speech, the CT and TA activity correlated with both opening and closing. Laryngeal muscle patterning to produce vocal fold movement differed across tasks; reciprocal muscle activity only occurred on cough, whereas speech and sniff often involved simultaneous contraction of muscle antagonists. In conclusion, different combinations of muscle activation are used for biomechanical control of vocal fold opening and closing movements during respiratory, airway protection, and speech tasks.     

Molar occlusion and mandibular movements during occlusion in the American opossum, Didelphis marsupialis L.

The dentition of the American opossum, Didelphis marsupialis , has been examined using prepared skulls, occlusal casts and, during normal function in chewing, by cinefluorography. The information obtained from these studies has been used in a simulation of the power stroke of mastication on a stereotaxic machine. The directions and amplitudes of the movements used in chewing have been demonstrated.
Two types of power stroke are described; a crushing-puncturing stroke associated with tooth-food-tooth contact occurring in the early stages of mastication, and a shearing stroke with tooth-tooth contact occurring later. The first type produces a rapid flattening of the tips of the cusps, the second a series of striated wear facets on the slopes of the cusps. In the shearing stroke the food is triturated by an anteromedial movement of the paracristid of the lower molars across the metacrista of the upper. Active chewing occurs on one side of the mouth only at any one time and for a considerable number of cycles. The symphysis does not cross the midline during any sequence of cycles. The anteromedially directed power stroke is mediated in part by the mobile symphysis which allows asymmetrical movement of the two dentaries and by a Bennett shift of the mandibular condyles.
The similarity between the molars of Didelphis and those of the early therians, notably Alphadon, Pappotherium and Holoclemensia is recognized. It is suggested that the information obtained from this study may assist in the interpretation of early therian molars.     

Cerebral control of face,jaw, and tongue movements in awake cats: Changes in regional cerebral blood flow during lateral feeding

Mastication is achieved by cooperation among facial, masticatory, and lingual muscles. However, cortical control in cats for the masticatory performance is processed by two systems: facial movement processed by facial SI (the first somatosensory cortex), area C, and area M (motor areas), and jaw and tongue movements performed by intraoral SI, masticatory area, and area P (motor area). In particular, outputs from area P organized in the corticobulbar tract are projected bilaterally in the brainstem. In this present study, the aim is to explore changes in the regional cerebral blood flow (rCBF) in the facial SI, area M, and area P during trained lateral feeding (licking or chewing from the right or left side) of milk, fish paste, and small dry fish. The rCBF in area M showed contralateral dominance, and rCBF in area P during chewing or licking from the right or left side was almost the same value. Furthermore, activities of genioglossus and masseter muscles in the left side showed almost the same values during licking of milk and of fish paste, and chewing of small dry fish during lateral feeding. These findings suggest that the cortical process for facial, jaw, and tongue movements may be regulated by the contralateral dominance of area M and the bilateral one of area P.