Mandibular function in Galago crassicaudatus and Macaca fascicularis: an in vivo approach to stress analysis of the mandible.

Single-element and/or rosette strain gages were bonded to mandibular cortical bone in Galago crassicaudatus and Macaca fascicularis. Five galago and eleven macaque bone strain experiments were performed and analyzed. In vivo bone strain was recorded from the lateral surface of the mandibular corpus below the postcanine tooth row during transducer biting and during mastication and ingestion of food objects. In macaques and galagos, the mandibular corpus on the balancing side is primarily bent in the sagittal plane during mastication and is both twisted about its long axis and bent in the sagittal plane during transducer biting. On the working side, it is primarily twisted about its long axis and directly sheared perpendicular to its long axis, and portions of it are bent in the sagittal plane during mastication and molar transducer biting. In macaques, the mandibular corpus on each side is primarily bent in the sagittal plane and twisted during incisal transducer biting and ingestion of food objects, and it is transversely bent and slightly twisted during jaw opening. Since galagos usually refused to bite the transducer or food objects with their incisors, an adequate characterization of mandibular stress patterns during these behaviors was not possible. In galagos the mandibular corpus experiences very little transverse bending stress during jaw opening, perhaps in part due to its unfused mandibular symphysis. Marked differences in the patterns of mandibular bone strain were present between galagos and macaques during the masticatory power stroke and during transducer biting. Galagos consistently had much more strain on the working side of the mandibular corpus than on the balancing side. These experiments support the hypothesis that galagos, in contrast to macaques, employ a larger amount of working-side muscle force relative to the balancing-side muscle force during unilateral biting and mastication, and that the fused mandibular symphysis is an adaption to use a maximal amount of balancing-side muscle force during unilateral biting and mastication. These experiments also demonstrate the effects that rosette position, bite force magnitudes, and types of food eaten have on recorded mandibular strain patterns.     

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.     

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)     

Aspects of masticatory form and function in common tree shrews,Tupaia glis

Tree shrews have relatively primitive tribosphenic molars that are apparently similar to those of basal eutherians; thus, these animals have been used as a model to describe mastication in early mammals. In this study the gross morphology of the bony skull, joints, dentition, and muscles of mastication are related to potential jaw movements and cuspal relationships. Potential for complex mandibular movements is indicated by a mobile mandibular symphysis, shallow mandibular fossa that is large compared to its resident condyle, and relatively loose temporomandibular joint ligaments. Abrasive tooth wear is noticeable, and is most marked at the first molars and buccal aspects of the upper cheek teeth distal to P². Muscle morphology is basically similar to that previously described for Tupaia minor and Ptilocercus lowii. However, in T. glis, an intraorbital part of deep temporalis has the potential for inducing lingual translation of its dentary, and the large medial pterygoid has extended its origin anteriorly to the floor of the orbit, which would enhance protrusion. The importance of the tongue and hyoid muscles during mastication is suggested by broadly expanded anterior bellies of digastrics, which may assist mylohyoids in tensing the floor of the mouth during forceful tongue actions, and by preliminary electromyography, which suggests that masticatory muscles alone cannot fully account for jaw movements in this species.     

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.     

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.     

Biomechanics of prehensile oral mucosa

Large dogs are able to deliver a powerful bite that generates considerable stress in the anterior, prehensile part of the jaws. In the upper jaw most of the biting force is borne by the anterior teeth. The palatal mucosa provides little resistance to deformation. It is easily compressed and rather mobile. In the lower jaw, the mucosa covering the upper surface of the symphysis receives a sizeable portion of the biting force. It is firmly attached to the underlying bone and possesses special connective tissue arrangements that enable it to transduce locally applied pressure to tension distributed over a broad area.     

Kinematics of the pharyngeal jaws during feeding in Oreochromis niloticus (Pisces,Perciformes)

Cichlids possess a complex pharyngeal jaw apparatus, the osteological components of which are two upper pharyngeal jaws, articulating with the neurocranial base, and a single lower pharyngeal jaw. Quantitative cinera-diography revealed that pharyngeal food processing in Oreochromis niloticus involves transport, mastication, and swallowing, effected by cyclical pharyngeal jaw movements. Transport and swallowing occur by simultaneous retractions of both upper pharyngeal jaws. Food reduction (mastication) is effected by lower jaw elevation (compression) and protraction (shear) during upper jaw retraction. Each movement cycle contains a transport, reduction, and swallowing component, although their relative importance may vary within a feeding sequence. The upper and lower pharyngeal jaws show opposite anteroposterior movements during most of the cycle. Variations in the amplitudes and the durations of the different movement components reflect the consistency and the size of the food.     

Experimental analysis of temporomandibular joint reaction force in macaques

Mandibular bone strain in the region immediately below the temporomandibular ligament was analyzed in adult and sub-adult Macaca fascicularis and Macaca mulatta. Following recovery from the general anesthetic, the monkeys were presented food objects, a wooden rod, or a specially designed bite-force transducer. Bone strain was recorded during incisal biting and mastication of food, and also during isometric biting of the rod and/or the transducer. The bone strain data suggest the following: The macaque TMJ is loaded by a compressive reaction force during the power stroke of mastication and incision of food, and during isometric molar and incisor biting. TMJ reaction forces are larger on the contralateral side during both mastication and isometric molar biting. Patterns of ipsilateral TMJ reaction force in macaques during isometric biting vary markedly in response to the position of the bite point. During biting along the premolars or first two molars a compressive reaction force acts about the ipsilateral TMJ; however, when the bite point is positioned along the M3, the ipsilateral TMJ has either very little compressive stress, no stress, or it is loaded in tension.     

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.     

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.     

Incisal biting in the mountain beaver (Aplodontia rufa) and woodchuck (Marmota monax)

Analysis of synchronously recorded cine-radiographs and electromyograms in two rodents (Aplodontia rufa and Marmota monax) demonstrates that jaw movements and muscle activiteis during incisal functions are distinctly different from those found during mastication. Movements during incisal biting are primarily along the midline, accompanied by symmetrical activity of the jaw adductor muscles. Most biting cycles do not end in contact between upper and lower incisors. When contact does occur, the lower incisors are dragged along the lingual surfaces of the upper incisors. Cropping, or tip-to-tip occlusion of upper and lower incisors, was not observed. Sharpening of the lower incisors, a behavior which may be unique to the Rodentia, was recorded in both A. rufa and M. monax. During sharpening, the lingual surface of the lower incisor is dragged across the tip of the upper incisor producing a lingual wear facet. Like incisal biting, sharpening movements are primarily confined to the midline, although there may be lateral movements in some sharpening cycles. Sharpening cycles are among the most rapid cyclic movements recorded in mammals, as the mean frequencies of sharpening are 11 cycles/s in A. rufa and 8 cycles/s in M. monax. © 1995 Wiley-Liss, Inc.     

The functional significance of primate mandibular form.

A stress analysis of the primate mandible suggests that vertically deep jaws in the molar region are usually an adaptation to counter increased sagittal bending stress about the balancing-side mandibular corpus during unilateral mastication. This increased bending stress about the balancing side is caused by an increase in the amount of balancing-side muscle force. Furthermore, this increased muscle force will also cause an increase in dorso-ventral shear stress along the mandibular symphysis. Since increased symphyseal stress can be countered by symphyseal fusion and as increased bending stress can be countered by a deeper jaw, deep jaws and symphyseal fusion are often part of the same functional pattern. In some primates (e.g., Cercocebus albigena), deep jaws are an adaptation to counter bending in the sagittal plane during powerful incisor biting, rather than during unilateral mastication. The stress analysis of the primate mandible also suggests that jaws which are transversely thick in the molar region are an adaptation to counter increased torsion about the long axis of the working-side mandibular corpus during unilateral mastication. Increased torsion of the mandibular corpus can be caused by an increase in masticatory muscle force, an increase in the transverse component of the postcanine bite force and/or an increase in premolar use during mastication. Patterns of masticatory muscle force were estimated for galagos and macaques, demonstrating that the ratio of working-side muscle force to balancing-side muscle force is approximately 1.5:1 in macaques and 3.5:1 in galagos during unilateral isometric molar biting. These data support the hypothesis that mandibular symphyseal fusion is an adaptative response to maximize unilateral molar bite force by utilizing a greater percentage of balancing-side muscle force.     

Functional significance of the mandibular symphysis.

The purpose of this investigation was to relate the morphology of connective tissues in the mandibular symphysis to the behavioral and experimental evidence for mobility and mechanical stress at the symphysis. The anatomy of the symphysis was examined histologically in 6 mammalian orders encompassing 22 species. Behavioral and experimental evidence of stress during the power stroke of the chewing cycle correspond with stresses at the symphysis implied by the location and orientation of symphyseal connective tissues. These stresses are: (1) dorsoventral shear of the symphysis due to the transfer of force from balancing to chewing sides, (2) bending of the symphysis causing tension along the inferior and compression along superior borders due to torsion on the dentaries from the jaw closing muscles, and (3) antero-posterior shear of the symphysis due to an anteriorly directed stress on the chewing side. Interspecific comparisons suggest that leaf eaters can resist greater dorsoventral shear than fruit or insect eaters, but no correlations exist between diet and bending or antero-posterior shear. This suggests that chewing leaves requires larger biting forces.     

The ontogeny of premolar dental wear in Cercocebus albigena (cercopithecidae)

The orientation of striated wear facets on primate teeth serves as a useful guide for reconstructing jaw movements during mastication. Most wear facets on the molars are formed during one of the two well-documented movements, Phase I or Phase II, of the power stroke. Another jaw movement direction, “orthal retraction” (OR) has been proposed to account for a third set of facets occasionally present on the pointed tips of premolars and molars. Evidence advanced here indicates that OR facets on pointed anterior premolars (P₃) of cercopithecoids are actually Phase I facets that have become reoriented as a result of a rotation of this tooth during its eruption. “Orthal retraction” probably does not exist as a discrete masticatory phase.     

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.     

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.     

Transverse masticatory movements, occlusal orientation, and symphyseal fusion in selenodont artiodactyls

Based on extensive experimental work on primates, two masticatory loading regimes have emerged as the likely determinants of mandibular symphyseal fusion-dorsoventral shear and lateral transverse bending (wishboning) (Ravosa and Hylander, 1994; Hylander et al., 1998, 2000). Recently, however, it has been argued that, rather than functioning to strengthen the symphysis during mastication, fusion serves to stiffen the symphyseal joint so as to facilitate increased transverse jaw movements during occlusion (Lieberman and Crompton, 2000). As part of this transverse stiffness model, it has been suggested that taxa with fused symphyses should also exhibit more horizontally oriented occlusal wear facets. Using a series of univariate and bivariate analyses, we test predictions of these three models in a sample of 44 species of selenodont artiodactyls. Consistent with the wishboning and transverse stiffness models, taxa with fused symphyses (camelids) have more horizontally oriented M(2) and M(2) occlusal wear facets, anteroposteriorly (AP) elongate symphyses, and relatively wider corpora. Contrary to the dorsoventral shear model, camelids do not have relatively deeper corpora (due to greater parasagittal bending). While taxa with ossified symphyses have relatively larger symphysis cross-sectional areas, this appears to be the byproduct of an increase in AP symphysis length due to greater lateral transverse bending of the mandible. Theoretical consideration of the biomechanics of mastication further suggests that strength, not stiffness, is the critical factor in determining symphyseal ossification. Thus, like anthropoid primates, fusion in selenodont artiodactyls appears to function in resisting increased wishboning stresses arising from an emphasis on transverse occlusal/mandibular movements and loads.     

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.