Incisal bite force direction in humans and the functional significance of mammalian mandibular translation

Incisal bite force direction was recorded and analyzed in ten human subjects using a specially designed force transducer. In all ten subjects the maxillary incisal bite force was vertically and anteriorly directed both during static biting and during biting associated with simultaneous mandibular translation and rotation. Since the resultant muscle force could not have been equal and opposite to the mandibular bite force, the mandibular condyles must have been loaded. These data demonstrate that the mandible acts as a lever during incisal biting and that there is no consistent relationship between incisal bite force direction and object size. In some individuals the bite force direction was more vertical during biting on a large transducer (30 mm high), while in other subjects it was more vertical during biting on a small transducer (10 mm high).     

Relationships between the size and spatial morphology of human masseter and medial pterygoid muscles, the craniofacial skeleton, and jaw biomechanics

The relationship between human craniofacial morphology and the biomechanical efficiency of bite force generation in widely varying muscular and skeletal types is unknown. To address this problem, we selected 22 subjects with different facial morphologies and used magnetic resonance imaging, cephalometric radiography, and data from dental casts to reconstruct their craniofacial tissues in three dimensions. Conventional cephalometric analyses were carried out, and the cross-sectional sizes of the masseter and medial pterygoid muscles were measured from reconstituted sections. The potential abilities of the muscles to generate bite forces at the molar teeth and mandibular condyles were calculated according to static equilibrium theory using muscle, first molar, and condylar moment arms. On average, the masseter muscle was about 66% larger in cross section than the medial pterygoid and was inclined more anteriorly relative to the functional occlusal plane. There was a significant positive correlation (P less than 0.01) between the cross-sectional areas of the masseter and medial pterygoid muscles (r = 0.75) and between the bizygomatic arch width and masseter cross-sectional area (r = 0.56) and medial pterygoid cross-sectional area (r = 0.69). The masseter muscle was always a more efficient producer of vertically oriented bite force than the medial pterygoid. Putative bite force from the medial pterygoid muscle alone correlated positively with mandibular length and inversely with upper face height. When muscle and tooth moment arms were considered together, a system efficient at producing force on the first molar was statistically associated with a face having a large intergonial width, small intercondylar width, narrow dental arch, forward maxilla, and forward mandible. There was no significant correlation between muscle cross-sectional areas and their respective putative bite forces. This suggests that there is no simple relationship between the tension-generating capacity of the muscles and their mechanical efficiency as described by their spatial arrangement. The study shows that in a modern human population so many combinations of biomechanically relevant variables are possible that subjects cannot easily be placed into ideal or nonideal categories for producing molar force. Our findings also confirm the impression that similar bite-force efficiencies can be found in subjects with disparate facial features.     

In vivo bone strain in the mandible of Galago crassicaudatus.

Single element foil strain gages were bonded to mandibular cortical bone in eight specimens of Galago crassicaudatus. The gage was bonded below the Pm4 or M2 adjacent to the lower border of the mandible. The bonded strain gage was connected to form one arm of a Wheatstone bridge. Following recovery from the general anesthetic, the restrained Galago bit either a piece of wood, a food object, or a bite force transducer. During these biting episodes, mandibular bone strain deformed the strain gage and the resulting change in electrical resistance of the gage caused voltage changes across the Wheatstone bridge. These changes, directly proportional to the amount of bone strain along the gage site, were recovered by a strip chart recorder. Bone strain was measured on both the working and balancing sides of the jaws. Maximum values of bone strain and bite force were 435 microstrain (compression) and 8.2 kilograms respectively. During bending of the mandible, the correlation between bone strain (tension or compression) and bite force ranged from -0.893 (tension) to 0.997 (compression). The experiments reported here demonstrate that only a small percentage of the Galago bite force is due to balancing side muscle force during isometric unilateral molar biting. In addition, these experiments demonstrate that the Galago mandible is bent in a predictable manner during biting. The amount of apparent mandibular bone strain is dependent on (1) the magnitude of the bite force and (2) the position of the bite point.     

The orientation of the force of the jaw muscles and the length of the mandible in mammals

Ungulates generally have large masseter and pterygoid muscles and a necessarily large angular process provides attachment surface on the mandible. The temporalis muscle tends to be small. It has been suggested that this is an adaptation for enhanced control of the lower jaw and reduction of forces at the jaw joint. I suggest an additional reason: because of the geometry of the jaw, the length of that segment of the lower jaw that spans the distance from the jaw joint to the most posterior tooth is significantly reduced when the masseler and pterygoid are the dominant muscles; this region is necessarily much longer when the temporalis is large.     

Sexual size dimorphism in caecilian amphibians: analysis, review and directions for future research

Sexual dimorphism, widespread in the animal kingdom, describes differences between the sexes in size, shape and many other traits. Sexual size dimorphism (SSD) plays a significant role in understanding life history evolution and mating systems. The snakelike morphology of limbless caecilian amphibians lacking obvious secondary sexual characters (in contrast to frogs and salamanders) impedes accurate intrasexual comparisons. In this study, sexual size dimorphism in the oviparous caecilian Ichthyophis cf. kohtaoensis, a phylogenetically basal caecilian, was analysed. Females were larger in all body and head characters tested. However, when adjusted to body size (total length), females differed only in their cloacal shape. Clutch volume was positively correlated to female body size, thus female fecundity increased with body size supporting the hypothesis of a fecundity-selected SSD in the oviparous Ichthyophis cf. kohtaoensis. A review of the present SSD data for caecilians shows that many species are monomorphic for body size but show dimorphism in head size, while other species demonstrate female-biased SSD. Male-biased SSD has not been reported for caecilians. To understand life history evolution in caecilians, further studies on the reproductive biology of other taxa are urgently needed, in particular for rhinatrematids and uraeotyphlids. New data will allow phylogenetically controlled comparative analyses to fully explore the pattern of SSD among caecilian lineages.     

Functional morphology of bite mechanics in the great barracuda (Sphyraena barracuda)

The great barracuda, Sphyraena barracuda, is a voracious marine predator that captures fish with a swift ram feeding strike. While aspects of its ram feeding kinematics have been examined, an unexamined aspect of their feeding strategy is the bite mechanism used to process prey. Barracuda can attack fish larger than the gape of their jaws, and in order to swallow large prey, can sever their prey into pieces with powerful jaws replete with sharp cutting teeth. Our study examines the functional morphology and biomechanics of 'ram-biting' behavior in great barracuda where the posterior portions of the oral jaws are used to slice through prey. Using fresh fish and preserved museum specimens, we examined the jaw mechanism of an ontogenetic series of barracuda ranging from 20 g to 8.2 kg. Jaw functional morphology was described from dissections of fresh specimens and bite mechanics were determined from jaw morphometrics using the software MandibLever (v3.2). High-speed video of barracuda biting (1500 framess(-1)) revealed that prey are impacted at the corner of the mouth during capture in an orthogonal position where rapid repeated bites and short lateral headshakes result in cutting the prey in two. Predicted dynamic force output of the lower jaw nearly doubles from the tip to the corner of the mouth reaching as high as 58 N in large individuals. A robust palatine bone embedded with large dagger-like teeth opposes the mandible at the rear of the jaws providing for a scissor-like bite capable of shearing through the flesh and bone of its prey.     

A dynamic model of mouth closing movements in clariid catfishes: the role of enlarged jaw adductors

Some species of Clariidae (air breathing catfishes) have extremely large (hypertrophied) jaw closure muscles. Besides producing higher bite forces, the enlarged muscles may also cause higher accelerations of the lower jaw during rapid mouth closure. Thus, jaw adductor hypertrophy could potentially also enable faster mouth closure. In this study, a forward dynamic model of jaw closing is developed to evaluate the importance of jaw adductor hypertrophy on the speed of mouth closure. The model includes inertia, pressure, tissue resistance and hydrodynamic drag forces on the lower jaw, which is modelled as a rotating half-ellipse. Simulations are run for four clariid species showing a gradual increase in jaw adductor hypertrophy (Clarias gariepinus, Clariallabes longicauda, Gymnallabes typus and Channallabes apus). The model was validated using data from high-speed videos of prey captures in these species. In general, the kinematic profiles of the fastest mouth closure from each species are reasonably well predicted by the model. The model was also used to compare the four species during standardized mouth closures (same initial gape angle, travel distance and cranial size). These simulations suggest that the species with enlarged jaw adductors have an increased speed of jaw closure (in comparison with the non-hypertrophied C. gariepinus) for short lower jaw rotations and when feeding at high gape angles. Consequently, the jaw system in these species seems well equipped to capture relatively large, evasive prey. For prey captures during which the lower jaw rotates freely over a larger distance before impacting the prey, the higher kinematic efficiency of the C. gariepinus jaw system results in the fastest jaw closures. In all cases, the model predicts that an increase in the physiological cross-sectional area of the jaw muscles does indeed contribute to the speed of jaw closure in clariid fish.     

A biomechanical model for analysis of muscle force,power output and lower jaw motion in fishes

Fish skulls are complex kinetic systems with movable components that are powered by muscles. Cranial muscles for jaw closing pull the mandible around a point of rotation at the jaw joint using a third-order lever mechanism. The present study develops a lever model for the jaw of fishes that uses muscle design and the Hill equation for nonlinear length-tension properties of muscle to calculate dynamic power output. The model uses morphometric data on skeletal dimensions and muscle proportions in order to predict behavior and force transmission mediated by lever action. The computer model calculates a range of dynamic parameters of jaw function including muscle force, torque, effective mechanical advantage, jaw velocity, bite duration, bite force, work and power. A complete list of required morphometrics is presented and a software program (MandibLever 2.0) is available for implementing lever analysis. Results show that simulations yield kinematics and timing profiles similar to actual fish feeding events. Simulation of muscle properties shows that mandibles reach their peak velocity near the start of jaw closing, peak force at the end of jaw closing, and peak power output at about 25% of the closing cycle time. Adductor jaw muscles with different mechanical designs must have different contractile properties and/or different muscle activity patterns to coordinate jaw closing. The effective mechanical advantage calculated by the model is considerably lower than the mechanical advantage estimated from morphological lever ratios, suggesting that previous studies of morphological lever ratios have overestimated force and underestimated velocity transmission to the mandible. A biomechanical model of jaw closing can be used to interpret the mechanics of a wide range of jaw mechanisms and will enable studies of the functional results of developmental and evolutionary changes in skull morphology and physiology.     

The caecilian cardio-vascular system and intra-amphibian relationships

The cardio-vascular system of Hypogeophis is described and compared with that of other caecilians. The extant caecilians are divisible into two geographical groups in which the varying degrees of asymmetry of the arterial arches evolved in parallel. The nomenclature of the cephalic veins is discussed and the cranial circulation of caecilians is compared with that of some urodeles and anurans. The general similarities which exist between the vascular system of caecilians and urodeles may be indicative of a close relationship. However, it is pointed out that more detailed information is required before intra-amphibian relationships can be established with any degree of certainty.     

Drinking behavior and jaw muscle (EMG) activity in the pigeon (Columba livia)

The relation between jaw movements and jaw muscle activity was examined during two different types of drinking in pigeons: tip and rictus drinking. The amplitude and duration of jaw opening is greater for rictus than for tip drinking, but both types involve individual cycles of jaw-opening and closing movements, organized into bouts. Cycle duration increases gradually over the initial portion of the bout and is relatively constant thereafter.Each drinking cycle is composed of an initial rapid jaw-opening component, a sustained opening phase of variable duration and a closing movement. The initial and final phases are related, respectively, to activity in the upper beak levator (protractor) and the jaw closer (adductor, pterygoid) muscles. The amplitude and duration of the sustained phase are correlated with the magnitude and duration of activity in the lower jaw opener (depressor). The kinematic and electromyographic organization of jaw movements during drinking is discussed in relation to the morphology of the jaw apparatus and the functional requirements of the behavior.Abbreviations AMEM adductor mandibulae externus muscle - DM depressor mandibulae muscle - EMG electromyographic - PQP protractor quadrati et pterygoidei muscle - PTP pseudotemporalis profundus muscle - PVL/PVM pterygoideus ventralis muscle, pars lateralis and medialis     

Jaw movement kinematics and jaw muscle (EMG) activity during drinking in the pigeon (Columba livia)

Summary Movements of the maxilla and mandible were recorded during drinking in the head-fixed pigeon and correlated with electromyographic activity in representative jaw muscle groups. During drinking, each jaw exhibits opening and closing movements along both the dorso-ventral and rostro-caudal axes which may be linked with or independent of each other. All subjects showed small but systematic increases in cycle duration over the course of individual drinking bouts. Cyclic jaw movements during drinking were correlated with nearly synchronous activity in the protractor (levator) of the upper jaw and in several jaw closer muscles, as well as with alternating activity in tongue protractor and retractor muscles. No EMG activity was ever recorded in the lower jaw opener muscle, suggesting that lower jaw opening in this preparation is produced, indirectly, by the contraction of other muscles. The results clarify the contribution of the individual jaws to the generation of gape variations during drinking in this species.Abbreviations AMEM adductor mandibulae externus muscle - DM depressor mandibulae muscle - EMG electromyographic - GENIO geniohyoideus muscle - LB lower beak - LED light-emitting diode - PQP protractor quadrati et pterygoidei muscle - PVL pterygoideus ventralis muscle, pars lateralis - SeH/StH serpihyoideus or stylohyoideus muscle - UB upper beak     

The functional relationship between feeding type and jaw and cranial morphology in ungulates

The relationship between jaw and skull morphology and feeding type (grazer, mixed feeder, browser, frugivorous, omnivorous) was analysed in 94 species of extant ungulates. A total of 21 morphological traits of the jaw and skull (17 and 4, respectively) were analysed using analysis of covariance, with body mass as covariate. To take into account the phylogenetic effect, simulations were generated under the Brownian motion model of character evolution. Analysis of covariance was applied to these simulations and the simulated F-ratios were used to assess the signification of the F-ratios for the real values of the traits. The feeding types had a weak effect on ungulate cranial and jaw morphology in comparison with the phylogenetic effect, since, before phylogeny correction, the analysis of covariance showed statistically significant differences associated with feeding type in 15 out of the 21 traits analysed. After controlling for phylogeny, only 2 significant traits remained, the length of the coronoid process and the occipital height. Omnivorous species had shorter coronoid processes than grazers or mixed feeders, and the occipital height was greater in the omnivorous species than in the grazers, mixed feeders or browsers. The coronoid process is involved in the generation of bite force, being the effective moment arm of the temporalis muscle, and occipital height is positively related to the force exerted by the temporalis muscle. This result matches the hypothesis that species with a toughness diet should show higher bite force (“toughness” describes the resistance of a material to being mechanically broken down). When the omnivorous species were excluded from the analysis, no differences in jaw and skull morphology were detected between the rest of the feeding types. Received: 1 September 1998 / Accepted: 2 November 1998     

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.     

Molar occlusion and jaw mechanics of the Eocene primate Adapis

Wear facets on molars of the Eocene primate Adapis magnus are described. Striations on these wear facets indicate three separate directions of mandibular movement during mastication. One direction corresponds to a first stage of mastication involving orthal retraction of the mandible. The remaining two directions correspond to buccal and lingual phases of a second stage of mastication involving a transverse movement of the mandible. The mechanics of jaw adduction are analysed for both the orthal retraction and transverse stages of mastication. During the orthal retraction stage the greatest component of bite force is provided by the temporalis muscles acting directly against the food with the mandible functioning as a link rather than as a lever. A geometrical argument suggests that during the transverse stage of mastication bite force is provided by the temporalis muscles of both sides, the ipsilateral medial and lateral pterygoid muscles, and the contralateral masseter muscle.     

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 assessment of jaw movement direction from dental microwear

In order to resolve several controversies about unusual jaw movements such as thegosis and orthal retraction, information about direction of jaw movement is essential. Ryan (1979a,b) proposed that asymmetry of microwear striations might provide such data. In vivo occurrences of asymmetric striations on chimpanzee molars are presented and analyzed. Results suggest that either thegosis contacts commonly occur on chimpanzee molars or that Ryan's model, as developed in vitro, does not accurately apply to naturally occurring molar microwear.     

角蟾科三亚科蝌蚪角质颌的显微结构比较(两栖纲,无尾目)

采用扫描电镜技术研究了角蟾科8种蝌蚪角质颌的显微结构特征和形态特点,阐述了角质颌对蝌蚪觅食方式的影响。实验结果表明:角蟾科蝌蚪的角质颌属于两个不同的类型。拟髭蟾亚科和掌突蟾亚科的蝌蚪具有相似的显微结构特征:角质颌呈厚重的U型,角质化程度高。颌鞘呈基部宽、顶端尖的圆锥形;角蟾亚科的蝌蚪角质颌呈纤弱的弓型,角质化程度低。颌鞘呈基部窄、长而顶端略弯曲的象牙型。进一步的分析发现,8种蝌蚪的颌鞘直径和密度呈显著负相关。这种显微结构的变化趋势也反映出蝌蚪对栖息环境和觅食方式的适应性。