Symphyseal fusion and jaw-adductor muscle force: an EMG study.
01 Aug 2000-American Journal of Physical Anthropology (Am J Phys Anthropol)-Vol. 112, Iss: 4, pp 469-492
TL;DR: Both the analysis of the W/B EMG ratios and the muscle firing pattern data support the hypothesis that symphyseal fusion and transversely-directed muscle force in anthropoids are functionally linked, which supports the hypotheses that the evolution of symphySEal fusion in anthropoid is an adaptation to strengthen the symphysis so as to counter increased wishboning stress during forceful unilateral mastication.
Abstract: The purpose of this study is to test various hypotheses about balancing-side jaw muscle recruitment patterns during mastication, with a major focus on testing the hypothesis that symphyseal fusion in anthropoids is due mainly to vertically- and/or transversely-directed jaw muscle forces. Furthermore, as the balancing-side deep masseter has been shown to play an important role in wishboning of the macaque mandibular symphysis, we test the hypothesis that primates possessing a highly mobile mandibular symphysis do not exhibit the balancing-side deep masseter firing pattern that causes wishboning of the anthropoid mandible. Finally, we also test the hypothesis that balancing-side muscle recruitment patterns are importantly related to allometric constraints associated with the evolution of increasing body size. Electromyographic (EMG) activity of the left and right superficial and deep masseters were recorded and analyzed in baboons, macaques, owl monkeys, and thick-tailed galagos. The masseter was chosen for analysis because in the frontal projection its superficial portion exerts force primarily in the vertical (dorsoventral) direction, whereas its deep portion has a relatively larger component of force in the transverse direction. The symphyseal fusion-muscle recruitment hypothesis predicts that unlike anthropoids, galagos develop bite force with relatively little contribution from their balancing-side jaw muscles. Thus, compared to galagos, anthropoids recruit a larger percentage of force from their balancing-side muscles. If true, this means that during forceful mastication, galagos should have working-side/balancing-side (W/B) EMG ratios that are relatively large, whereas anthropoids should have W/B ratios that are relatively small. The EMG data indicate that galagos do indeed have the largest average W/B ratios for both the superficial and deep masseters (2.2 and 4.4, respectively). Among the anthropoids, the average W/B ratios for the superficial and deep masseters are 1.9 and 1.0 for baboons, 1.4 and 1.0 for macaques, and both values are 1.4 for owl monkeys. Of these ratios, however, the only significant difference between thick-tailed galagos and anthropoids are those associated with the deep masseter. Furthermore, the analysis of masseter firing patterns indicates that whereas baboons, macaques and owl monkeys exhibit the deep masseter firing pattern associated with wishboning of the macaque mandibular symphysis, galagos do not exhibit this firing pattern. The allometric constraint-muscle recruitment hypothesis predicts that larger primates must recruit relatively larger amounts of balancing-side muscle force so as to develop equivalent amounts of bite force. Operationally this means that during forceful mastication, the W/B EMG ratios for the superficial and deep masseters should be negatively correlated with body size. Our analysis clearly refutes this hypothesis. As already noted, the average W/B ratios for both the superficial and deep masseter are largest in thick-tailed galagos, and not, as predicted by the allometric constraint hypothesis, in owl monkeys, an anthropoid whose body size is smaller than that of thick-tailed galagos. Our analysis also indicates that owl monkeys have W/B ratios that are small and more similar to those of the much larger-sized baboons and macaques. Thus, both the analysis of the W/B EMG ratios and the muscle firing pattern data support the hypothesis that symphyseal fusion and transversely-directed muscle force in anthropoids are functionally linked. This in turn supports the hypothesis that the evolution of symphyseal fusion in anthropoids is an adaptation to strengthen the symphysis so as to counter increased wishboning stress during forceful unilateral mastication. (ABSTRACT TRUNCATED)
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...Second, dorsoventral shear forces at the symphysis are due to the different forces of rotation caused by muscles from the balancing or working sides of the mandible (Walker, 1978; Hylander et al., 2000)....
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TL;DR: 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.
Abstract: 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.
392 citations
TL;DR: The data suggest that during the power stroke of mastication, the macaque symphysis is predominately sheared dorsoventrally and/or twisted about a transverse axis and bent by lateral transverse bending of the mandibular corpora.
Abstract: The primary purpose of this study was to test various hypotheses about symphyseal stress in primates. First, those patterns of symphyseal strain that would be associated with various hypothetical patterns of symphyseal stress were formulated. Then these hypothetical patterns of stress and strain were tested by comparing the formulated bone strain pattern with actual in vivo symphyseal bone strain patterns. Patterns of in vivo symphyseal bone strain were determined by bonding rosette and/or single-element strain gages to the midline of the middle and lower third of the labial aspect of the symphysis of six adult Macaca fascicularis. Following recovery from the anesthetic, bone strain was recorded during mastication, incision, and isometric biting. Symphyseal bone strain was also recorded during yawning, licking, and threat behaviors. The data suggest that during the power stroke of mastication, the macaque symphysis is predominately sheared dorsoventrally and/or twisted about a transverse axis and bent by lateral transverse bending of the mandibular corpora. During lateral transverse bending of the mandibular corpora, the labial aspect of the macaque symphysis experiences compressive bending stress, while the lingual aspect experiences tensile bending stress. During the opening stroke of mastication and during other jaw opening behaviors, the macaque symphysis is bent by medial transverse bending of the mandibular corpora. At this time the labial aspect of the symphysis experiences tensile bending stress, while its lingual aspect experiences compressive bending stress. During both the power and opening strokes of mastication, the macaque mandible is bent in the plane of its curvature, and therefore the mandible acts as a curved beam. This is important because it results in elevated levels of stress along the lingual aspect of the macaque symphysis, particularly during the power stroke of mastication. During the power stroke of incision, the local effects of the bite force are unknown; however, at this time the lower half of the macaque symphysis is both sheared dorsoventrally and bent due to twisting of the mandibular corpora about their long axes. The results of this stress analysis have implications for understanding the mechanical attributes of symphyseal structure. In order to counter dorsoventral shear, the most important symphyseal attribute is to have adequate cross-sectional area of bone in the plane of the applied stress. In contrast, both the cross-sectional area of bone and symphyseal shape is important in order to counter stress effectively during symphyseal torsion and the three symphyseal bending regimes.(ABSTRACT TRUNCATED AT 400 WORDS)
364 citations
TL;DR: Single‐element and/or rosette strain gages were bonded to mandibular cortical bone in Galago crassicaudatus and Macaca fascicularis to record bone strain during transducer biting and during mastication and ingestion of food objects.
Abstract: 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.
345 citations
TL;DR: The stress analysis and an allometric analysis of mandibular dimensions in female cercopithecine (Old World) monkeys indicates that allometric changes in the symphysis are readily understood if the mandible is modelled as a curved beam.
Abstract: Patterns of stress were analyzed in the mandibular symphysis of Macaca fascicularis using rosette strain gages. During jaw opening, the mandibular symphysis is bent due to medial transverse bending of the mandibular corpora. Levels of stress and strain are relatively low at this time, and the source of this stress is the medially-directed component of force from the lateral pterygoid muscles. During the power stroke of mastication, the symphysis is maximally stressed. At this time the symphysis experiences dorsoventral shear and bending due to lateral transverse bending of the mandibular corpora, i.e., “wishboning.” The dorsoventral shear is due to the vertical component of the balancingside adductor muscle force; the “wishboning” is due to the laterally-directed components of the bite and jaw adductor muscle forces. Unlike dorsoventral shear, “wishboning” results in considerable levels of stress and strain, particularly along the most lingual aspect of the symphysis. The most effective way to counter this stress is to increase the thickness of the symphysis in the labio-lingual direction. The stress analysis and an allometric analysis of mandibular dimensions in female cercopithecine (Old World) monkeys indicates that allometric changes in the symphysis are readily understood if the mandible is modelled as a curved beam. With increasing body size, symphyseal thickness in cercopithecines must increase in a positively allometric fashion so as to prevent the occurrence of dangerously high levels of stress along the most lingual aspect of the symphysis. This is because increasing body size is associated with three factors thathave important consequences within the context of the biomechanics of curved beams: (1) jaw length is positively allometric to body size, (2) mandibular-arch width is negatively allometric to body size, and (3) there is a tendency to use relatively greater amounts of balancing-side jaw muscle force with increased body size because of dietary changes and allometricconstraints on total jaw muscle force.
326 citations
TL;DR: 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.
Abstract: 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)
258 citations