Loading patterns and jaw movements during mastication in Macaca fascicularis: a bone-strain, electromyographic, and cineradiographic analysis

Rosette strain gage, electromyography (EMG), and cineradiographic techniques were used to analyze loading patterns and jaw movements during mastication in Macaca fascicularis. The cineradiographic data indicate that macaques generally swallow frequently throughout a chewing sequence, and these swallows are intercalated into a chewing cycle towards the end of a power stroke. The bone strain and jaw movement data indicate that during vigorous mastication the transition between fast close and the power stroke is correlated with a sharp increase in masticatory force, and they also show that in most instances the jaws of macaques are maximally loaded prior to maximum intercuspation, i.e. during phase I (buccal phase) occlusal movements. Moreover, these data indicate that loads during phase II (lingual phase) occlusal movements are ordinarily relatively small. The bone strain data also suggest that the duration of unloading of the jaw during the power stroke of mastication is largely a function of the relaxation time of the jaw adductors. This interpretation is based on the finding that the duration from 100% peak strain to 50% peak strain during unloading closely approximates the half-relaxation time of whole adductor jaw muscles of macaques. The EMG data of the masseter and medial pterygoid muscles have important implications for understanding both the biomechanics of the power stroke and the external forces responsible for the "wishboning" effect that takes place along the mandibular symphysis and corpus during the power stroke of mastication. Although both medial pterygoid muscles reach maximum EMG activity during the power stroke, the activity of the working-side medial pterygoid peaks after the balancing-side medial pterygoid. Associated with the simultaneous increase of force of the working-side medial pterygoid and the decrease of force of the balancing-side medial pterygoid is the persistently high level of EMG activity of the balancing-side deep masseter (posterior portion). This pattern is of considerable significance because the direction of force of both the working-side medial pterygoid and the balancing-side deep masseter are well aligned to aid in driving the working-side lower molars across the upper molars in the medial direction during unilateral mastication.(ABSTRACT TRUNCATED AT 400 WORDS)     

The Ontogeny of Mammalian Mastication

Movements of the oral apparatus begin during the fetal periodand develop in a consistent order. Jaw openingappears first,followed by active jaw closure and tongue movements, lip movements,sucking, and finally masticatory movements. The later developingmovements appear prenatally in precocious mammals such as guineapigs and sheep, but are postnatal in altricial mammals suchas rats, hamsters and rabbits. The orderly development of oralbehavior is probably related to the progressive maturation ofthe nervous system and neuromuscular connections. Most newbornmammals feed exclusively bysuckling, a combination of the tongueworking against the nipple and negative pressure at the backof the oral cavity. Thetransition from suckling to masticationis gradual and involves considerable learning. In at least onespecies, the domestic pig, infant animals chew using a somewhatdifferent muscular contraction pattern from that of adults.Age changes in muscle action lines are the most likely explanationfor this difference. After being established in infancy, theprocess of mastication undergoes only minor changes in rateand relative muscle activity during the juvenile period. Throughoutontogeny there is a reciprocal relation between morphology andbehavior. While masticatory performance depends on structureat any given stage, it also has profound effects on furthermusculoskeletal growth and differentiation.     

Ontogeny of oral function in hamsters (Mesocricetus auratus)

The oral apparatus of neonatal and juvenile golden hamsters was investigated by clearing and staining of whole crania, videotaping of behavior, and electromyography of several jaw muscles. Chewing developed during the first postnatal week and matured in the second; however, suckling was still the primary mode of feeding. Micromovements of the jaws occurred early when the osseous skeleton and joints developed. Macromovements correlated well with EMG records and were limited to jaw opening at birth. Muscles of the oral floor generated large bursts of activity during jaw opening and tongue protrusion from 0 days postnatal (dpn), when simple and stereotyped gaping was induced, until 14 dpn, when movements were spontaneous and not stereotyped nor inducible. However, adductor muscle activity was brief, low in amplitude, and primarily involved with jaw stabilization until 4 dpn, when these muscles became active during closing the jaws; closing activity increased in frequency and amplitude until the end of the second week. Development of frequent, coordinated macromovements of chewing was associated with the refinement of joint structure and dental occlusion and with the growth of the craniofacial skeleton. Jaw movements and associated EMG's correlated better with available data on development of neural circuitry than with that for musculoskeletal development.     

Masticatory motor patterns in ungulates: a quantitative assessment of jaw-muscle coordination in goats, alpacas and horses

We investigated patterns of jaw-muscle coordination during rhythmic mastication in three species of ungulates displaying the marked transverse jaw movements typical of many large mammalian herbivores. In order to quantify consistent motor patterns during chewing, electromyograms were recorded from the superficial masseter, deep masseter, posterior temporalis and medial pterygoid muscles of goats, alpacas and horses. Timing differences between muscle pairs were evaluated in the context of an evolutionary model of jaw-muscle function. In this model, the closing and food reduction phases of mastication are primarily controlled by two distinct muscle groups, triplet I (balancing-side superficial masseter and medial pterygoid and working-side posterior temporalis) and triplet II (working-side superficial masseter and medial pterygoid and balancing-side posterior temporalis), and the asynchronous activity of the working- and balancing-side deep masseters. The three species differ in the extent to which the jaw muscles are coordinated as triplet I and triplet II. Alpacas, and to a lesser extent, goats, exhibit the triplet pattern whereas horses do not. In contrast, all three species show marked asynchrony of the working-side and balancing-side deep masseters, with jaw closing initiated by the working-side muscle and the balancing-side muscle firing much later during closing. However, goats differ from alpacas and horses in the timing of the balancing-side deep masseter relative to the triplet II muscles. This study highlights interspecific differences in the coordination of jaw muscles to influence transverse jaw movements and the production of bite force in herbivorous ungulates.     

Mandibular movements of the albino rat during feeding

Jaw movements of albino rats during biting and mastication of relatively hard food were recorded by means of conventional and X-ray cinematography. Mandibular kinetics have been analysed in the context of passive mechanical limits imposed by jaw morphology, particularly of the joints, and by the food itself. Movements have been described in terms of degrees of gape, condylar translation and horizontal rotation of the rami about the symphysis. During biting the condyle remains in the anterior two-thirds of the fossa, moves forward as the jaw opens and the converse. The rami usually spread well apart; the lower incisors are usually approximated. Incised food particles are transported toward the molars by means of coordinated jaw and tongue movements. The prominent palatal rugae of the diastemal region abet this process. In the power stroke of mastication, the mandible shifts forward as the lower toothrows move a little inward; the condyles occupy the posterior two-thirds of the fossa. All movements seen were bilaterally symmetrical. Simultaneous chewing occurred on both sides. It is suggested that the lingual components in the primarily anterior power stroke enhance grinding efficiency. A movable symphysis appears to be of critical importance in facilitating this type of mastication.     

Videofluorographic analysis of tongue movement in the rabbit (Oryctolagus cuniculus)

The movement of the entire tongue and intermolar eminence during mastication is described in the domestic rabbit (Oryctolagus cuniculus). Tongue movement and jaw position were analyzed videofluorographically from separate lateral and dorso-ventral views in six rabbits. Metallic markers were inserted into the tongue so that its movement was visible on the fluorographic image. Frame-by-frame analysis of the videofluorographic tape recordings demonstrates that tongue movement in all animals was identical in direction during each part of the chewing cycle. In the lateral view the forepart of the tongue moves down and forward during the opening stroke, whereas the intermolar eminence moves up and forward to appose the palate. During the closing stroke, as the tip of the tongue moves up and back, the intermolar eminence lowers from the palate and retracts. During the power stroke the forepart of the tongue is at its most elevated and retruded position, while the intermolar eminence is its lowest and most retruded. The dorso-ventral view showed that lateral movement of the tongue and mandible are highly synchronous. The intermolar eminence decreases in width during the power stroke, possibly twisting to place or keep food on the teeth. An anterior to posterior undulating movement of the entire tongue occurs throughout the chewing cycle. As the intermolar eminence elevates to appose the palate during the opening stroke, it may replace the bolus on the teeth on the chewing side. The intermolar eminence also appears to be twisting during the closing and power strokes to place or maintain food on the teeth.     

Phase-linked variations in the amplitude of the digastric nerve jaw-opening reflex response during fictive mastication in the rabbit

The diagastric nerve reflex response to stimulation of the upper lip was studied in urethan-anesthetized rabbits paralysed with pancuronium bromide. Rhythmic bursts of masticatory activity were evoked in the nerve by repetitive electrical stimulation of the motor cortex. The amplitude and latency of the reflex responses during fictive mastication were compared with preceding control values. When stimuli close to threshold were given, the largest and earliest responses occurred during the digastric burst. When intense stimuli were employed, the largest responses were out of phase with the burst, although the latency was still shortest when the motoneurons were rhythmically active. Since the pattern is essentially the same as that seen during normal mastication, we conclude that the cyclical modulation of reflex amplitude and latency is not the result of sensory feedback generated by the movements themselves but is instead governed by the central motor program.     

Electromyographic responses to prescribed mastication.

The aim was to understand between-volunteer differences in Electromyography (EMG) behaviour during chewing. EMG was used to record the electrical activity of the temporal and masseter muscles of volunteers, who carried out mastication movements by operating calibrated springs held between their incisors. The volunteers coordinated their jaw movements with the signal produced by a metronome, at four rates: 30, 60, 90 and 120 beats per minute (bpm). Raw data were analyzed to examine the distributions of the intervals between chews. For the highest prescribed chew rates, the volunteers' distributions were very similar. The distributions varied most for the 30 bpm data, suggesting that volunteers differed in their ability to carry out and maintain this prescribed chewing pattern. The data were Fourier transformed to give power spectra in the frequency domain. The low frequency (<10 Hz) region contained spectral features related to the prescribed chew rate. Principal component analysis of the power spectra revealed that readings from each volunteer clustered together, and the clusters could be largely separated. Such grouping was found irrespective of whether data from each chew rate were analyzed separately or simultaneously. This indicated that within-volunteer variance, arising from the different chew rates as well as between-session variance, is lower than between-volunteer variance; even when individuals are asked to make jaw movements in the same prescribed manner, they can nevertheless be uniquely distinguished by their muscle activity as recorded by EMG.     

EMG of the digastric muscle in gibbon and orangutan: Functional consequences of the loss of the anterior digastric in orangutans

Unlike all other primates, the digastric muscle of the orangutan lacks an anterior belly; the posterior belly, while present, inserts directly onto the mandible. To understand the functional consequences of this morphologic novelty, the EMG activity patterns of the digastric muscle and other potential mandibular depressors were studied in a gibbon and an orangutan. The results suggest a significant degree of functional differentiation between the two digastric bellies. In the gibbon, the recruitment pattern of the posterior digastric during mastication is typically biphasic. It is an important mandibular depressor, active in this role during mastication and wide opening. It also acts with the anterior suprahyoid muscles to move the hyoid prior to jaw opening during mastication. The recruitment patterns of the anterior digastric suggest that it is functionally allied to the geniohyoid and mylohyoid. For example, although it transmits the force of the posterior digastric during mandibular depression, it functions independent of the posterior digastric during swallowing. Of the muscles studied, the posterior digastric was the only muscle to exhibit major differences in recruitment pattern between the two species. The posterior digastric retains its function as a mandibular depressor in orangutans, but is never recruited biphasically, and is not active prior to opening. The unique anatomy of the digastric muscle in orangutans results in decoupling of the mechanisms for hyoid movement and mandibular depression, and during unilateral activity it potentially contributes to substantial transverse movements of the mandible. Hypotheses to explain the loss of the anterior digastric should incorporate these functional conclusions. © 1994 Wiley-Liss, Inc.     

The physiology and ontogeny of daily oral behaviors

In the masticatory system, activities of muscles are the main source of force. The daily activity of the jaw muscle is a measure of the total daily loading of the tissues involved. This article gives an overview on the recent assessments of the physiology and ontogeny of the daily use of the jaw muscles. Variations in the characteristics of daily activity could be linked to differences in the types of fibers composing the muscles as well as to the properties of the underlying bone, although these relationships are not absolute. Experimental decrease of the hardness of foods eaten by rats and rabbits showed a significant decrease in the number of daily bursts of feeding. These reductions in daily muscular activity were accompanied by higher mineralization of bone and by a transition toward "faster" fiber types in the muscles. It was revealed in rabbits that the characteristics of the daily activities of muscles (total duration of activity, number and lengths of bursts) were not altered during the transition from suckling to chewing and remained largely unaffected during further postnatal development. These results suggest that, despite large anatomical and functional changes, the average daily load on the jaw muscles by the masticatory system appears to be established before chewing develops and remains largely unchanged all the way through development. Whenever the daily muscular activity changes, this seems to have a significant effect on the properties of the tissues involved.     

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.     

Three-dimensional finite element modelling of muscle forces during mastication

This paper presents a three-dimensional finite element model of human mastication. Specifically, an anatomically realistic model of the masseter muscles and associated bones is used to investigate the dynamics of chewing. A motion capture system is used to track the jaw motion of a subject chewing standard foods. The three-dimensional nonlinear deformation of the masseter muscles are calculated via the finite element method, using the jaw motion data as boundary conditions. Motion-driven muscle activation patterns and a transversely isotropic material law, defined in a muscle-fibre coordinate system, are used in the calculations. Time-force relationships are presented and analysed with respect to different tasks during mastication, e.g. opening, closing, and biting, and are also compared to a more traditional one-dimensional model. The results strongly suggest that, due to the complex arrangement of muscle force directions, modelling skeletal muscles as conventional one-dimensional lines of action might introduce a significant source of error.     

Effect of sensory input from the tongue on jaw movement in normal feeding in the opossum

Opossums were presented with solid and liquid foods. The movements of the jaw and tongue were recorded cineradiographically together with recordings of the EMG activity in muscles opening the jaw and moving the base of the tongue (hyoid). The jaw opening in each cycle was in two stages--01 and 02; 01 had a constant amplitude irrespective of the food ingested. Ingestion of liquid (which involved continuous accumulation of a liquid bolus in the valleculae prior to swallowing) was associated with cycles of oral movement in which 02 was small; tongue retraction was associated with this opening. In contrast, solid and semisolid food ingestion was associated with large angles of jaw opening in 02 that also coincided with the tongue retraction. In this latter case a characteristic pattern of EMG activity, in which all the muscles moving the hyoid were simultaneously active, was added to the pattern seen in lapping; this additional activity had an EMG pattern that was consistent with a jaw opening reflex. The findings contrast with other reports that the jaw opening reflex is suppressed in mastication. Experimentally induced tongue contact with a variety of solid surfaces during lapping (an activity involving accumulation of a liquid bolus in the valleculae) induced neither increased jaw opening nor the additional EMG pattern. However, in situations when there was no bolus in the valleculae, additional jaw opening activity was elicited when the tongue contracted solids intra- or extra-orally. It is suggested that the ability of sensory input, from the anterior tongue, to elicit a jaw opening reflex and to change the type of jaw/tongue cycle was dependent upon the extent of bolus accumulation in the valleculae and therefore indirectly upon the consistency of the food.     

Sensory feedback in the control of mouthpart movements in the desert locust Schistocerca gregaria

ABSTRACT. When imposed movements were applied to one or both mandibles of the desert locust, Schistocerca gregaria , the other mouthparts moved in synchrony with the mandibles. This occurred in the presence or absence of food, and when the mandibles were driven at a higher or lower frequency than that seen during normal feeding. Electromyogram recordings from the mandibular closer muscles revealed bursts of activity at the same frequency as the imposed movement. This activity occurred during mandibular closing. Burst length was a function of driving wavelength. At low driving frequencies (less than 0.5 Hz), smaller bursts were seen prior to the longer closing burst; a series of similar small bursts was seen when the mandibles were held in the open position. When one mandible was driven, closer muscle activity was largely confined to that side. In the presence of food, however, activity was seen in both closer muscles. A possible mechanism for this is described. After destruction of the campaniform sensilla on the ventral surface of the mandibles, the bursts of activity in the mandibular closers, seen when the mandibles were held open, were replaced by continuous activity. This suggests that the function of these sensilla is to inhibit motor output to the closer muscles when the tension becomes high. When feeding on relatively incompressible food the closer muscle burst length increased, although chewing frequency did not alter. This effect was also produced by loading the mandibles artificially. A model for the feedback control of this behaviour is proposed.     

Control of jaw movements in two species of macropodines (Macropus eugenii and Macropus rufus).

The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.     

Control of jaw movements in two species of macropodines (Macropus eugenii and Macropus rufus)

The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.     

Examination of the relationships between jaw opener and closer rhythmical muscle activity in an in vitro brainstem jaw-attached preparation

An in vitro jaw-attached brainstem preparation was developed to investigate the relationship between jaw opener and closer muscle activity during chemically induced rhythmical jaw movements in neonatal rats. In the majority of preparations examined, where a defined region of brainstem was isolated and the neuronal innervation of the jaw opener and closer muscles was left intact, bath application of the excitatory amino acid agonist N -methyl-D,L-aspartate (NMA, 20-40 muM) in combination with bicuculline (BIC 10 muM), a GABAA antagonist, produced rhythmical electromyogram (EMG) activity in jaw opener and closer muscles, bilaterally, in conjunction with rhythmical jaw movements. Low concentrations of NMA (20 muM) in combination with BIC produced temporally coordinated activity between the jaw opener and closer muscles, ipsilaterally. With higher doses of NMA (40 muM), each muscle group exhibited bursting, but temporal coordination between them was difficult to establish. Similarly, NMA application in combination with the glycine antagonist strychnine (STR, 10 muM), also produced rhythmical EMG activity from both opener and closer muscles, ipsilaterally, but showed no temporal coordination between the antagonist muscle pair. However, coordination of opener and closer muscle discharge could be restored by the addition of BIC to the bath. We suggest that there exist separate, but coordinated, rhythm generator circuits for opener and closer motoneuronal discharge located in close proximity to the trigeminal motor nucleus and under GABAergic control for production of temporal coordination between rhythmogenic circuits.     

Evolution of muscle activity patterns driving motions of the jaw and hyoid during chewing in Gnathostomes

Although chewing has been suggested to be a basal gnathostome trait retained in most major vertebrate lineages, it has not been studied broadly and comparatively across vertebrates. To redress this imbalance, we recorded EMG from muscles powering anteroposterior movement of the hyoid, and dorsoventral movement of the mandibular jaw during chewing. We compared muscle activity patterns (MAP) during chewing in jawed vertebrate taxa belonging to unrelated groups of basal bony fishes and artiodactyl mammals. Our aim was to outline the evolution of coordination in MAP. Comparisons of activity in muscles of the jaw and hyoid that power chewing in closely related artiodactyls using cross-correlation analyses identified reorganizations of jaw and hyoid MAP between herbivores and omnivores. EMG data from basal bony fishes revealed a tighter coordination of jaw and hyoid MAP during chewing than seen in artiodactyls. Across this broad phylogenetic range, there have been major structural reorganizations, including a reduction of the bony hyoid suspension, which is robust in fishes, to the acquisition in a mammalian ancestor of a muscle sling suspending the hyoid. These changes appear to be reflected in a shift in chewing MAP that occurred in an unidentified anamniote stem-lineage. This shift matches observations that, when compared with fishes, the pattern of hyoid motion in tetrapods is reversed and also time-shifted relative to the pattern of jaw movement.     

Temporalis function in anthropoids and strepsirrhines: an EMG study

The major purpose of this study is to analyze anterior and posterior temporalis muscle force recruitment and firing patterns in various anthropoid and strepsirrhine primates. There are two specific goals for this project. First, we test the hypothesis that in addition to transversely directed muscle force, the evolution of symphyseal fusion in primates may also be linked to vertically directed balancing-side muscle force during chewing (Hylander et al. [2000] Am. J. Phys. Anthropol. 112:469-492). Second, we test the hypothesis of whether strepsirrhines retain the hypothesized primitive mammalian condition for the firing of the anterior temporalis, whereas anthropoids have the derived condition (Weijs [1994] Biomechanics of Feeding in Vertebrates; Berlin: Springer-Verlag, p. 282-320). Electromyographic (EMG) activities of the left and right anterior and posterior temporalis muscles were recorded and analyzed in baboons, macaques, owl monkeys, thick-tailed galagos, and ring-tailed lemurs. In addition, as we used the working-side superficial masseter as a reference muscle, we also recorded and analyzed EMG activity of the left and right superficial masseter in these primates. The data for the anterior temporalis provided no support for the hypothesis that symphyseal fusion in primates is linked to vertically directed jaw muscle forces during mastication. Thus, symphyseal fusion in primates is most likely mainly linked to the timing and recruitment of transversely directed forces from the balancing-side deep masseter (Hylander et al. [2000] Am. J. Phys. Anthropol. 112:469-492). In addition, our data demonstrate that the firing patterns for the working- and balancing-side anterior temporalis muscles are near identical in both strepsirrhines and anthropoids. Their working- and balancing-side anterior temporalis muscles fire asynchronously and reach peak activity during the power stroke. Similarly, their working- and balancing-side posterior temporalis muscles also fire asynchronously and reach peak activity during the power stroke. Compared to these strepsirrhines, however, the balancing-side posterior temporalis of anthropoids appears to have a relatively delayed firing pattern. Moreover, based on their smaller W/B ratios, anthropoids demonstrate a relative increase in muscle-force recruitment of the balancing-side posterior temporalis. This in turn suggests that anthropoids may emphasize the duration and magnitude of the power stroke during mastication. This hypothesis, however, requires additional testing. Furthermore, during the latter portion of the power stroke, the late activity of the balancing-side posterior temporalis of anthropoids apparently assists the balancing-side deep masseter in driving the working-side molars through the terminal portion of occlusion.     

Functional links between canine height and jaw gape in catarrhines with special reference to early hominins

This study tests the hypothesis that decreased canine crown height in catarrhines is linked to (and arguably caused by) decreased jaw gape. Associations are characterized within and between variables such as upper and lower canine height beyond the occlusal plane (canine overlap), maximum jaw gape, and jaw length for 27 adult catarrhine species, including 539 living subjects and 316 museum specimens. The data demonstrate that most adult male catarrhines have relatively larger canine overlap dimensions and gapes than do conspecific females. For example, whereas male baboons open their jaws maximally more than 110% of jaw length, females open about 90%. Humans and hylobatids are the exceptions in that canine overlap is nearly the same in both the sexes and so is relative gape (ca. 65% for humans and 110% for hylobatids). A correlation analysis demonstrates that a large portion of relative gape (maximum gape/projected jaw length) is predicted by relative canine overlap (canine overlap/jaw length). Relative gape is mainly a function of jaw muscle position and/or jaw muscle‐fiber length. All things equal, more rostrally positioned jaw muscles and/or shorter muscle fibers decrease gape and increase bite force during the power stroke of mastication, and the net benefit is to increase the mechanical efficiency during chewing. Similarly, more caudally positioned muscles and/or longer muscle fibers increase the amount of gape and decrease bite force. Overall, the data support the hypothesis that canine reduction in early hominins is functionally linked to decreased gape and increased mechanical efficiency of the jaws. Am J Phys Anthropol, 2013. © 2012 Wiley Periodicals, Inc.