Temporal changes in human temporomandibular joint size and shape

Measurements approximating the size of the temporomandibular joint were taken on a series of genetically homogeneous populations from early Nubia, which span almost 10,000 years and embody a shift from a primarily hunting and gathering adaptation to a completely agricultural lifeway. A generalized trend of reduction of temporomandibular joint size was observed. In addition, a decrease in sexual dimorphism was apparent for all measures of joint size; a change primarily mitigated by reductions in male dimensions. The observed variation in size and form of the temporomandibular joint is most likely the result of the reduction in masticatory muscle robusticity and resultant changes in craniofacial form which have been documented for the transition from a hunting and gathering to an agricultural subsistence in Nubia.     

Constraints on masticatory system evolution in anthropoid primates

It is well established that some observed patterns of force production in the primate masticatory system match those predicted by a simplified lever model. This model is also commonly invoked in adaptive explanations of craniodental diversity. However, systematic studies of the predictive power of this model are missing, leaving open the possibility that factors not traditionally included in the model alter the function and evolution of the masticatory system. One such factor was proposed for mammals generally by Greaves ([1978] J. Zool. (Lond.) 184:271-285), who argued that the temporomandibular joint (TMJ) was poorly suited to being pulled apart. In this constrained lever model, the avoidance of joint distraction leads to limitations on masticatory system form and function. The goal of the present study was to quantify masticatory system diversity in anthropoid primates for comparison with these predictions. Results indicate that all sampled taxa exhibit a form that is consistent with selection against regular distraction of the TMJ. Also apparent from observed patterns of scaling is a regular interaction among a limited set of cranial and dental dimensions, in accordance with the constrained model. However, the data indicate that specific positional relationships among the muscles, joints, and teeth differ from those predicted by Greaves (1978). The pattern of deviation suggests that selection has favored a conservative masticatory system configuration that safeguards the TMJ from distraction during the dynamic processing of irregular foods. The resulting buffered model leads to alternative hypotheses regarding the response of the masticatory system to dietary selection pressures. It may, therefore, improve our understanding of the adaptive significance of primate craniofacial form.     

新疆罗布泊小河墓地居民的口腔健康与饮食

小河墓地是新疆罗布泊地区一处重要的早期青铜时代墓地。本文主要对该墓地出土颅骨所附牙齿的磨耗程度及牙结石沉积状况进行了观察、量化统计和分析,同时也对该人群其他的口腔疾病如根尖脓肿、颞下颌关节病变、生前牙齿脱落等做了简单的统计,以期从古病理学的角度获取当时居民的口腔健康、食物类型和饮食习惯等信息。本研究发现:1)小河人群的牙齿磨耗程度远远高于对比组的古代居民,其上腭圆枕及颞下颌关节炎出现率较高,存在牙齿崩裂现象,且其前后部牙齿磨耗差异不大。这一方面说明其食物加工技术比较落后,食物粗糙坚硬;另一方面小河居民的经济生活方式和食物构成都比较复杂,不同的食物对前部和后部牙齿磨耗的程度造成了不同的影响;此外,小河人群风沙肆虐的生活环境也对其严重牙齿磨耗的形成产生了一定影响。2)小河人群异常严重的牙结石沉积归功于其高蛋白质和碳水化合物的饮食,以及生活用水的水质。3)统计分析发现两性存在上、下颌犬齿的磨耗差异,而这一情况可能暗示了在家庭手工业方面存在男女分工的现象。     

Biomechanical association of dental and temporomandibular pathology in a medieval Nubian population

An analysis of the relationship between oral pathology and degenerative change at the temporomandibular joint (TMJ) was undertaken on an archaeological sample of 122 adult crania from the Medieval site of Kulubnarti in Sudanese Nubia. The crania were sorted into 2 groups: those demonstrating clearly visible bony changes at the joint (TMJ+) and those without visible change (TMJ-). These groups were compared according to 1) age; 2) sex; 3) active dental pathologies (abscesses, caries, partial socket resorption); 4) tooth loss with complete socket resorption; and 5) dental attrition. No statistically significant association was evident between degenerative change at the TMJ and age, active dental pathologies, or dental attrition; however, sex differences and posterior tooth loss with complete socket resorption revealed a significant correspondence to degenerative TMJ changes. Both of these factors agree with the clinical literature and with biomechanical models (most notably that of Hylander) based upon modern populations. Furthermore, the results support the contention that paleopathological conditions can be analyzed from a clinical and functional biomechanical perspective.     

Influences of osteoclast deficiency on craniofacial growth in osteopetrotic (op/op) mice

It is well known that the defect in bone resorption in osteopetrotic (op/op) mice brings about deformation of the cranium and failure of tooth eruption. However, the influences on longitudinal growth of the craniofacial skeleton have not been elucidated. This study was thus conducted to examine craniofacial morphology and longitudinal changes in the op/op mice by means of morphometric analysis with lateral cephalograms. Lateral cephalograms, taken every 10 days from 10- to 90-day-old mice, were analyzed on a personal computer for 11 measurement items. For the nasal bone region, the most prominent differences were found between the op/op and normal mice. The anterior cranial base and occipital bone height presented almost equivalent growth changes in both the op/op and normal mice. The size of mandible, meanwhile, was significantly smaller in the op/op mice than in the normal controls. The gonial angle was also significantly larger in the op/op mice than in the normal mice throughout the experimental period. Thus, substantial differences in craniofacial growth were demonstrated in various areas of the craniofacial complex, which are assumed essentially due to the lack of osteoclastic bone resorption during growing period. Since the difference became more prominent in the anatomic regions relevant to the masticatory functions, it would be a reasonable assumption that reduced masticatory function is also a key determinant for the less-developed craniofacial skeleton in the op/op mouse.     

Tooth wear in eastern grey kangaroos (Macropus giganteus) and western grey kangaroos (Macropus fuliginosus), and its potential influence on diet selection, digestion and population parameters

As teeth wear in eastern and western grey kangaroos, the pattern of enamel ridges changes. This affects the amount of available cutting edges and consequently the masticatory efficiency. These kangaroos also have molar progression. There is evidence for the independence of rates of tooth wear and molar progression, and variation is found in the maximum amount of wear attained by teeth both within and between populations. This variation has the potential to affect diet selection, digestion and population parameters.     

Functional implications of primate enamel thickness.

Recent evolutionary interpretations of Hominoidea have postulated functional relationships between tooth form, diet and masticatory biomechanics. A major consideration is the durability of the tooth under certain dietary conditions. Teeth with low cusps and thicker enamel are able to withstand heavy mastication of abrasive food bolus for a longer period. When comparisons are made between species of higher primates the variables of tooth size, cusp morphology, and enamel thickness appear to be related but until now no systematic analysis has been made to determine the functional relevance of several dental dimensions. This study provides data gained from comparisons of dentition of nine species of primates. Histological sections were made of the post canine teeth and 21 dimensions were compared. The relevant dimensions identified serve to withstand dental wear. The distribution of thicker enamel corresponded to the observed wear planes. Humans had thicker enamel than pongids while the macaque had the thinnest. These preliminary results tend to support theories which explain low, thick, enameled cusps in hominids.     

Evolving Teeth Within a Stable Masticatory Apparatus in Orkney Mice

Mice from the Orkney archipelago exhibit an important diversity regarding molar shape. While on some islands mice display a usual dental pattern, teeth from other islands display additional cusplets and unusual phenotypes that may constitute case studies for evaluating the potential functional relevance of dental changes. We developed a multifaceted approach combining 2D and 3D geometric morphometrics, dental topography, dental wear, biomechanics, estimations of masticatory muscles force, and in vivo bite force on wild-derived lab descendants exemplifying the two extreme dental morphologies. The two strains differed in the geometry of the upper and lower tooth rows, and in the topography of the upper row only. Surprisingly, the most unusual tooth morphology appeared as the least complex because tooth simplification overwhelmed the signal provided by the occurrence of additional cusplets. No difference in bite force nor muscle force was evidenced, showing that the important change in dental morphology was accommodated without major changes in the rest of the masticatory apparatus. The evolution of unusual dental phenotypes was possibly fueled by drift and inbreeding in small and isolated populations on remote islands of the archipelago. No functional counter-selection impeded this diversification, since the unusual dental phenotypes did not disrupt occlusion and mastication.     

Size and shape of the mandibular condyle in primates

The relationships between the size of the articular surface of the mandibular condyle and masticatory muscle size, tooth size, diet, and biomechanical variables associated with mastication were studied by taking 12 measurements on skulls of 253 adult female anthropoid primates, including three to ten specimens from each of 32 species. In regressions of condylar length, width, or area against body weight, logarithmic transformations substantially improve the fit of the equations compared with untransformed data. There is a strong relationship between condylar measurements and body weight, with all correlations being .94 or higher. The slopes of the allometric regressions of length, width, and area of the condylar head indicate slight positive allometry with body size. Folivorous primates have smaller condyles than frugivorous primates, and colobines have smaller condyles than cebids, cercopithecines, or hominoids. When colobines are eliminated, the differences between frugivores and folivores are not significant. However, the two species with the relatively largest condyles are Pongo pygmaeus and Cercocebus torquatus, suggesting that there may be a relationship between unusually large condylar dimensions and the ability to crack hard nuts between the teeth. Cranial features having strong positive correlations with condylar dimensions include facial prognathism, maxillary incisor size, maxillary postcanine area, mandibular ramus breadth, and temporal fossa area. These data are interpreted as indicating that relatively large condyles are associated with relatively large masticatory muscles, relatively inefficient mandibular biomechanics, and a large dentition. These relationships support the growing evidence that the temporomandibular joint is a stress-bearing joint in normal function.     

The functional significance of morphological variation of the human mandible and masticatory muscles

A review is given of what is known about the functional significance of variation of the morphology of the human mandible and jaw muscles. First, the mandible is a lever transferring muscular forces to the teeth. The angle between corpus and ramus and the width of the ramus are particularly relevant in this respect as they determine the mechanical advantage of the lever system and the capacity for sagittal (open-close) movement. The stability of the mandible in asymmetric bites is especially affected by the ratio between the intermolar and intercondylar distances. The repertoire of bite forces that can be generated at any tooth and the loading pattern of the temporomandibular joint are strongly dependent on the relative size of the masseter, temporalis and medial pterygoid muscles. Second, executing its function as a lever, the mandible is subjected to shearing, bending and torsional forces. The bony parts harbouring the teeth, joints and muscle attachments serve to counter these forces; additional strength is needed in three areas i.e. in the symphysis, the condylar neck and in the transition area between corpus and ramus. In human populations there are clear-cut patterns of correlation between some facial skeletal traits, jaw joint morphology and strength and line of action of the jaw muscles. As a result, facial morphologies can be distinguished with marked differences in mechanical performance of their masticatory apparatus. It is suggested that they emerge as a result of diverging environmental influences during postnatal growth.     

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.     

Relationships between attrition and lingual tilting in human teeth

Associations between occlusal dental attrition and the lingual tilting of human teeth were investigated in two aboriginal California populations. A literature survey suggests lingual tilting is related geographically and temporally to the helicoidal occlusal plane in humans; if true, lingual tilting should be expected throughout the Homo osteological record. Buccal alveolar abscesses, exposed pulp chambers, extent of tooth attrition, angle of lingual tilt, and amount of overjet were observed for lingually tilted teeth. These attributes were analyzed statistically, leading to a conclusion that lingual tilting is slight and infrequent unless tooth attrition is pronounced. It is suggested that lingual tilting is due in part to masticatory stress. A feedback model considers lingual tilting as one conspicuous manifestation of a more complex “severe attrition syndrome,” the initial stimulus for which may derive from heavy occlusal tooth wear. Untested possibilities should be examined–e.g., age-related changes, the impact of the helicoidal occlusal plane, and the role of masticatory forces.     

Tooth wear and compensatory modification of the anterior dentoalveolar complex in humans

In populations living in environments where teeth wear severely, some compensatory modification of the dentoalveolar complex is thought to occur during life whereby functional occlusion is maintained as tooth substance is lost by wear. This study investigates one aspect of this modification process: Changes in the anterior dentoalveolar complex that are accompanied with wear were examined in a series of Japanese skeletal samples. In the prehistoric Japanese hunter-gatherer population heavy wear occurs over the entire dentition. The following changes were demonstrated to have occurred in the anterior segment of the dentition accompanied by wear on the anterior teeth: The anterior teeth tip lingually with wear up to a nearly upright position to fill in interproximal spaces that would have been generated by wear, and to maintain contact relations between adjacent teeth. At the same time, the anterior surface of the maxillary alveolar process also inclines lingually to a certain extent. The amount of lingual tipping is greater in the maxillary anterior teeth than in their mandibular antagonists. It is because of this discrepancy that, with age, the horizontal component of the overlap between maxillary and mandibular anterior teeth decreases, and their bite form changes from scissor bite to edge-to-edge bite. Lesser degrees of lingual tipping of the anterior teeth were also detected in the prehistoric agriculturists and historic Japanese populations. The variation in the degree of lingual tipping observed among the samples is explained by inter-population variation in severity and pattern of tooth wear. This and other evidence suggests that mechanisms that compensate for wear in the anterior dentition may be characteristic of all living human populations, independently of the degree of wear severity endured in their environments.     

Differences in loading of the temporomandibular joint during opening and closing of the jaw

Kinematics of the human masticatory system during opening and closing of the jaw have been reported widely. Evidence has been provided that the opening and closing movement of the jaw differ from one another. However, different approaches of movement registration yield divergent expectations with regard to a difference in loading of the temporomandibular joint between these movements. Because of these diverging expectations, it was hypothesized that joint loading is equal during opening and closing. This hypothesis was tested by predicting loading of the temporomandibular joint during an unloaded opening and closing movement of the jaw by means of a three-dimensional biomechanical model of the human masticatory system. Model predictions showed that the joint reaction forces were markedly higher during opening than during closing. The predicted opening trace of the centre of the mandibular condyle was located cranially of the closing trace, with a maximum difference between the traces of 0.45 mm. The hypothesis, postulating similarity of joint loading during unloaded opening and closing of the jaw, therefore, was rejected. Sensitivity analysis showed that the reported differences were not affected in a qualitative sense by muscular activation levels, the thickness of the cartilaginous layers within the temporomandibular joint or the gross morphology of the model. Our predictions indicate that the TMJ is loaded more heavily during unloaded jaw opening than during unloaded jaw closing.     

Dental topography and molar wear in Alouatta palliata from Costa Rica

Paleoprimatologists depend on relationships between form and function of teeth to reconstruct the diets of fossil species. Most of this work has been limited to studies of unworn teeth. A new approach, dental topographic analysis, allows the characterization and comparison of worn primate teeth. Variably worn museum specimens have been used to construct species-specific wear sequences so that measurements can be compared by wear stage among taxa with known differences in diet. This assumes that individuals in a species tend to wear their molar teeth in similar ways, a supposition that has yet to be tested. Here we evaluate this assumption with a longitudinal study of changes in tooth form over time in primates. Fourteen individual mantled howling monkeys (Alouatta palliata) were captured and then recaptured after 2, 4, and 7 years when possible at Hacienda La Pacifica in Costa Rica between 1989-1999. Dental impressions were taken each time, and molar casts were produced and analyzed using dental topographic analysis. Results showed consistent decreases in crown slope and occlusal relief. In contrast, crown angularity, a measure of surface jaggedness, remained fairly constant except with extreme wear. There were no evident differences between specimens collected in different microhabitats. These results suggest that different individual mantled howling monkeys wear their teeth down in similar ways, evidently following a species-specific wear sequence. Dental topographic analysis may therefore be used to compare morphology among similarly worn individuals from different species.     

Tool use in animals: Cognition and ecology. Edited by Crickette M. Sanz,Josep Call,and Christophe Boesch. New York,NJ: Cambridge University Press. 2013. 313 pp. ISBN 978‐1‐107–01119‐9. $110 (hardcover).

Enamel thickness has been linked to functional aspects of masticatory biomechanics and has been demonstrated to be an evolutionary plastic trait, selectively responsive to dietary changes, wear and tooth fracture. European Late Paleolithic and Mesolithic hunter‐gatherers mainly show a flat wear pattern, while oblique molar wear has been reported as characteristic of Neolithic agriculturalists. We investigate the relationships between enamel thickness distribution and molar wear pattern in two Neolithic and medieval populations. Under the assumption that dietary and/or non‐dietary constraints result in directional selective pressure leading to variations in enamel thickness, we test the hypothesis that these two populations will exhibit significant differences in wear and enamel thickness patterns. Occlusal wear patterns were scored in upper permanent second molars (UM2) of 64 Neolithic and 311 medieval subadult and adult individuals. Enamel thickness was evaluated by microtomography in subsamples of 17 Neolithic and 25 medieval individuals. Eight variables describing enamel thickness were assessed. The results show that oblique molar wear is dominant in the Neolithic sample (87%), while oblique wear affects only a minority (42%) of the medieval sample. Moreover, in the Neolithic molars, where buccolingually directed oblique wear is dominant and greatest enamel lost occurs in the distolingual quadrant, thickest enamel is found where occlusal stresses are the most important—on the distolingual cusp. These results reveal a correlation between molar wear pattern and enamel thickness that has been associated to dietary changes. In particular, relatively thicker molar enamel may have evolved as a plastic response to resist wear. Am J Phys Anthropol 155:162–172, 2014. © 2014 Wiley Periodicals, Inc.     

Comparison of Habit Reversal and a Behaviorally-Modified Dental Treatment for Temporomandibular Disorders: A Pilot Investigation

This study tested the hypothesis that a habit reversal program emphasizing awareness and reduction of masticatory muscle activity would significantly reduce pain in patients diagnosed with chronic temporomandibular disorder (TMD) and would be a competitive alternative to a behaviorally-modified dental intervention. Eight individuals diagnosed with TMD were randomly assigned to a splint therapy or habit reversal group. Patients in the splint group received an interocclusal appliance (splint) fabricated from acrylic and were instructed to wear the splint day and night up to a maximum of 20 h per day. Patients in the habit reversal group were given a pager and instructed to check tooth position and masticatory muscle tension when paged. Paging occurred approximately once every 2 h during the day, but not at night. Both groups were instructed to avoid tooth contact and relax the masticatory muscles during the 4 weeks of active treatment. Outcome data were collected at 1 month and 1 year post-treatment intervals. Pain decreased significantly for both groups and did not differ between groups. Habit reversal may be as effective as a behaviorally-modified splint therapy for TMD-related pain.     

Dietary correlates of temporomandibular joint morphology in New World primates

Previous analyses of the masticatory apparatus have demonstrated that the shape of the temporomandibular joint (TMJ) is functionally and adaptively linked to variation in feeding behavior and diet in primates. Building on previous research, this study presents an analysis of the link between diet and TMJ morphology in the context of functional and dietary differences among New World primates. To evaluate this proposed relationship, I used three-dimensional morphometric methods to quantify TMJ shape across a sample of 13 platyrrhine species. A broad interspecific analysis of this sample found strong relationships among TMJ size, TMJ shape, and diet, suggesting that both size and diet are significant factors influencing TMJ morphology in New World primates. However, it is likely that at least some of these differences are related to a division of dietary categories along clade lines.A series of hypotheses related to load resistance capabilities and range of motion in the TMJ were then tested among small groups of closely related taxa with documented dietary differences. These pairwise analyses indicate that some aspects of TMJ morphology can be used to differentiate among closely related species with different diets. However, not all of my predictions were upheld. The anteroposterior dimensions of the TMJ were most strongly consistent with hypothesized differences in ingestive/masticatory behaviors and jaw gape, whereas the predictions generated for variation in entoglenoid and articular tubercle height were not upheld. These results imply that while some features can be reliably associated with increased load resistance and facilitation of wider jaw gapes in the masticatory apparatus, other features are less strongly correlated with masticatory function.     

Morphology and mechanics of the teeth and jaws of white-spotted bamboo sharks (Chiloscyllium plagiosum)

The teeth of white-spotted bamboo sharks (Chiloscyllium plagiosum) are used to clutch soft-bodied prey and crush hard prey; however, the dual function is not evident from tooth morphology alone. Teeth exhibit characteristics that are in agreement with a clutching-type tooth morphology that is well suited for grasping and holding soft-bodied prey, but not for crushing hard prey. The dual role of this single tooth morphology is facilitated by features of the dental ligament and jaw joint. Tooth attachment is flexible and elastic, allowing movement in both sagittal and frontal planes. During prey capture spike-like tooth cusps pierce the flesh of soft prey, thereby preventing escape. When processing prey harder than the teeth can pierce the teeth passively depress, rotating inward towards the oral cavity such that the broader labial faces of the teeth are nearly parallel to the surface of the jaws and form a crushing surface. Movement into the depressed position increases the tooth surface area contacting prey and decreases the total stress applied to the tooth, thereby decreasing the risk of structural failure. This action is aided by a jaw joint that is ventrally offset from the occlusal planes of the jaws. The offset joint position allows many teeth to contact prey simultaneously and orients force vectors at contact points between the jaws and prey in a manner that shears or rolls prey between the jaws during a bite, thus, aiding in processing while reducing forward slip of hard prey from the mouth. Together the teeth, dental ligament, and jaws form an integrated system that may be beneficial to the feeding ecology of C. plagiosum, allowing for a diet that includes prey of varying hardness and elusiveness.     

Effect of craniofacial genotype on the relationship between morphology and feeding performance in cichlid fishes

The relationship between morphology and performance is complex, but important for understanding the adaptive nature of morphological variation. Recent studies have sought to better understand this system by illuminating the interconnectedness of different functional systems; however, the role of genetics is often overlooked. In this study, we attempt to gain insights into this relationship by examining the effect of genotypic variation at putative craniofacial loci on the relationship between morphology and feeding performance in cichlids. We studied two morphologically disparate species, as well as a morphologically intermediate hybrid population. We assessed feeding performance, jaw protrusion, and general facial morphology for each fish. We also genotyped hybrid animals at six previously identified craniofacial loci. Cichlid species were found to differ in facial geometry, kinematic morphology, and performance. Significant correlations were also noted between these variables; however, the explanatory power of facial geometry in predicting performance was relatively poor. Notably, when hybrids were grouped by genotype, the relationship between shape and performance improved. This relationship was especially robust in animals with the specialist allele at sox9b, a well‐characterized regulator of craniofacial development. These data suggest a novel role for genotype in influencing complex relationships between form and function.