Stressed out: masticatory forces and primate circumorbital form.
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Animals↗
Biomedical subjects
W L Hylander
Publications and source records attributed to W L Hylander.
Electromyography of the anterior temporalis and masseter muscles of owl monkeys (Aotus trivirgatus) and the function of the postorbital septum.
Anthropoids and tarsiers are distinguished from all other vertebrates by the possession of a postorbital septum, which is formed by the frontal, alisphenoid, and zygomatic bones. Cartmill [(1980) In: Evolutionary Biology of the New World Monkeys and Continental Drift. New York: Plenum, p 243-274] suggested that the postorbital septum evolved in the stem lineage of tarsiers and anthropoids to insulate the eye from movements arising in the temporal fossa. Ross [(1996) Am J Phys Anthropol 91:305-324] suggested that the septum insulates the orbital contents from incursions by the line of action of the anterior temporal muscles caused by the unique combination of high degrees of orbital frontation and convergence. Both of these hypotheses must explain why insulation of the orbital contents could not be achieved by decreasing the size of the anterior temporal musculature with a corresponding increase in size of the remaining jaw adductors, rather than evolving a postorbital septum. One possibility is that the anterior temporalis is an important contributor to vertically directed bite forces during all biting and chewing activities. Another possibility is that reduction in anterior temporal musculature would compromise the ability to produce powerful bite forces, either at the incisors or along the postcanine toothrow. To evaluate these hypotheses, electromyographic (EMG) recordings were made from the masseter muscle and the anterior and posterior portions of the temporalis muscles of two owl monkeys, Aotus trivirgatus. The EMG data indicate that anterior temporalis activity relative to that of the superficial masseter is lower during incision than mastication. In addition, activity of the anterior temporalis is not consistently higher than the posterior temporalis during incision. The data indicate relatively greater activity of anterior temporalis compared to other muscles during isometric biting on the postcanine toothrow. This may be due to decreased activity in superficial masseter and posterior temporalis, rather than elevated anterior temporalis activity. The anterior temporalis is not consistently less variable in activity than the superficial masseter and posterior temporalis. The EMG data gathered here indicate no reason for suggesting that the anterior temporal muscles in anthropoids are utilized especially for incisal preparation of hard fruits. Maintenance of relatively high EMG activity in anterior temporalis across a wide range of biting behaviors is to be expected in a vertically oriented and rostrally positioned muscle such as this because, compared to the posterior temporalis, superficial masseter and medial pterygoid, it can contribute relatively larger vertical components of force to bites along the postcanine toothrow. The in vivo data do not support this hypothesis, possibly because of effects of bite point and bite force orientation.
Animals↗
Symphyseal fusion and jaw-adductor muscle force: an EMG study.
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)
Animals↗
Experimental observation, theoretical models, and biomechanical inference in the study of mandibular form.
Experimental studies and mathematical models are disparate approaches for inferring the stress and strain environment in mammalian jaws. Experimental designs offer accurate, although limited, characterization of biomechanical behavior, while mathematical approaches (finite element modeling in particular) offer unparalleled precision in depiction of strain magnitudes, directions, and gradients throughout the mandible. Because the empirical (experimental) and theoretical (mathematical) perspectives differ in their initial assumptions and their proximate goals, the two methods can yield divergent conclusions about how masticatory stresses are distributed in the dentary. These different sources of inference may, therefore, tangibly influence subsequent biological interpretation. In vitro observation of bone strain in primate mandibles under controlled loading conditions offers a test of finite element model predictions. Two issues which have been addressed by both finite element models and experimental approaches are: (1) the distribution of torsional shear strains in anthropoid jaws and (2) the dissipation of bite forces in the human alveolar process. Not surprisingly, the experimental data and mathematical models agree on some issues, but on others exhibit discordance. Achieving congruence between these methods is critical if the nature of the relationship of masticatory stress to mandibular form is to be intelligently assessed. A case study of functional/mechanical significance of gnathic morphology in the hominid genus Paranthropus offers insight into the potential benefit of combining theoretical and experimental approaches. Certain finite element analyses claim to have identified a biomechanical problem unrecognized in previous comparative work, which, in essence, is that the enlarged transverse dimensions of the postcanine corpus may have a less important role in resisting torsional stresses than previously thought. Experimental data have identified subperiosteal cortical thinning as a culprit in diminishing the role of cross-sectional geometry in conditioning the strain environment. These observations raise questions concerning the biomechanical significance of mandibular form in early hominids, fueling persistent arguments over whether gnathic morphology can be related to dietary specialization in the "robust" australopithecines. Nonmechanical explanations (e.g., tooth size or body size) for Paranthropus mandibular dimensions, however, are not compelling as competing hypotheses. Both theoretical and experimental models are in need of refinement before it is possible to conclude that the jaws of the "robust" australopithecines are not functionally linked to elevated masticatory loads.
Animals↗
Elastic properties and masticatory bone stress in the macaque mandible.
One important limitation of mechanical analyses with strain gages is the difficulty in directly estimating patterns of stress or loading in skeletal elements from strain measurements. Because of the inherent anisotropy in cortical bone, orientation of principal strains and stresses do not necessarily coincide, and it has been demonstrated theoretically that such differences may be as great as 45 degrees (Cowin and Hart, 1990). Likewise, relative proportions of stress and strain magnitudes may differ. This investigation measured the elastic properties of a region of cortical bone on both the buccal and lingual surfaces of the lower border of the macaque mandible. The elastic property data was then combined with macaque mandibular strain data from published and a new in vivo strain gage experiment to determine directions and magnitudes of maximum and minimum principal stresses. The goal was to compare the stresses and strains and assess the differences in orientation and relative magnitude between them. The main question was whether these differences might lead to different interpretations of mandibular function. Elastic and shear moduli, and Poisson's ratios were measured using an ultrasonic technique from buccal and lingual cortical surfaces in 12 macaque mandibles. Mandibular strain gage data were taken from a published set of experiments (Hylander, 1979), and from a new experiment in which rosette strain gauges were fixed to the buccal and lingual cortices of the mandibular corpus of an adult female Macaca fascicularis, after which bone strain was recorded during mastication. Averaged elastic properties were combined with strain data to calculate an estimate of stresses in the mandibular corpus. The elastic properties were similar to those of the human mandibular cortex. Near its lower border, the macaque mandible was most stiff in a longitudinal direction, less stiff in an inferosuperior direction, and least stiff in a direction normal to the bone's surface. The lingual aspect of the mandible was slightly stiffer than the buccal aspect. Magnitudes of stresses calculated from average strains ranged from a compressive stress of -16.00 GPa to a tensile stress of 8.84 GPa. The orientation of the principal stresses depended on whether the strain gage site was on the working or balancing side. On the balancing side of the mandibles, maximum principal stresses were oriented nearly perpendicular to the lower border of the mandible. On the working side of the mandibles, the orientation of the maximum principal stresses was more variable than on the balancing side, indicating a larger range of possible mechanisms of loading. Near the lower border of the mandible, differences between the orientation of stresses and strains were 12 degrees or less. Compared to ratios between maximum and minimum strains, ratios between maximum and minimum stresses were more divergent from a ratio of 1.0. Results did not provide any major reinterpretations of mandibular function in macaques, but rather confirmed and extended existing work. The differences between stresses and strains on the balancing side of the mandible generally supported the view that during the power stroke the mandible was bent and slightly twisted both during mastication and transducer biting. The calculated stresses served to de-emphasize the relative importance of torsion. On the working side, the greater range of variability in the stress analysis compared to the strain analysis suggested that a more detailed examination of loadings and stress patterns in each individual experiment would be useful to interpret the results. Torsion was evident on the working side; but in a number of experiments, further information was needed to interpret other superimposed regional loading patterns, which may have included parasagittal bending and reverse parasagittal bending.
Animals↗
Strain in the galago facial skull.
Little experimental work has been directed at understanding the distribution of stresses along the facial skull during routine masticatory behaviors. Such information is important for understanding the functional significance of the mammalian circumorbital region. In this study, bone strain was recorded along the dorsal interorbit, postorbital bar, and mandibular corpus in Otolemur garnettii and O. crassicaudatus (greater galagos) during molar chewing and biting. We determined principal-strain magnitudes and directions, compared peak shear-strain magnitudes between various regions of the face, and compared galago strain patterns with similar experimental data for anthropoids. This suite of analyses were used to test the facial torsion model (Greaves [1985] J Zool (Lond) 207:125-136; [1991] Zool J Linn Soc 101:121-129; [1995] Functional morphology in vertebrate paleontology. Cambridge: Cambridge University Press, p 99-115). A comparison of galago circumorbital and mandibular peak strains during powerful mastication indicates that circumorbital strains are very low in magnitude. This demonstrates that, as in anthropoids, the strepsirhine circumorbital region is highly overbuilt for countering routine masticatory loads. The fact that circumorbital peak-strain magnitudes are uniformly low in both primate suborders undermines any model that emphasizes the importance of masticatory stresses as a determinant of circumorbital form, function, and evolution. Preliminary data also suggest that the difference between mandibular and circumorbital strains is greater in larger-bodied primates. This pattern is interpreted to mean that sufficient cortical bone must exist in the circumorbital region to prevent structural failure due to nonmasticatory traumatic forces. During unilateral mastication, the direction of epsilon(1) at the galago dorsal interorbit indicates the presence of facial torsion combined with bending in the frontal plane. Postorbital bar principal-strain directions during mastication are oriented, on average, very close to 45 degrees relative to the skull's long axis, much as predicted by the facial torsion model. When chewing shifts from one side of the face to the other, there is a characteristic reversal or flip-flop in principal-strain directions for both the interorbit and postorbital bar. Although anthropoids also exhibit an interorbital reversal pattern, peak-strain directions for this clade are opposite those for galagos. The presence of such variation may be due to suborder differences in relative balancing-side jaw-muscle force recruitment. Most importantly, although the strain-direction data for the galago circumorbital region offer support for the occurrence of facial torsion, the low magnitude of these strains suggests that this loading pattern may not be an important determinant of circumorbital morphology.
Animals↗
Masticatory stress, orbital orientation and the evolution of the primate postorbital bar.
A postorbital bar is one of a suite of derived features which distinguishes basal primates from their putative sister taxon, plesiadapiforms. Two hypotheses have been put forward to explain postorbital bar development and variation in circumorbital form: the facial torsion model and visual predation hypothesis. To test the facial torsion model, we employ strain data on circumorbital and mandibular loading patterns in representative primates with a postorbital bar and masticatory apparatus similar to basal primates. To examine the visual predation hypothesis, we employ metric data on orbit orientation in Paleocene and Eocene primates, as well as several clades of visual predators and foragers that vary interspecifically in postorbital bar formation.A comparison of galago circumorbital and mandibular peak strains during powerful mastication demonstrates that circumorbital strains are quite low. This indicates that, as in anthropoids, the strepsirhine circumorbital region is excessively overbuilt for countering routine masticatory loads. The fact that circumorbital peak-strain levels are uniformly low in both primate suborders undermines any model which posits that masticatory stresses are determinants of circumorbital form, function and evolution. This is interpreted to mean that sufficient cortical bone must exist to prevent structural failure due to non-masticatory traumatic forces. Preliminary data also indicate that the difference between circumorbital and mandibular strains is greater in larger taxa.Comparative analyses of several extant analogs suggest that the postorbital bar apparently provides rigidity to the lateral orbital margins to ensure a high level of visual acuity during chewing and biting. The origin of the primate postorbital bar is linked to changes in orbital convergence and frontation at smaller sizes due to nocturnal visual predation and increased encephalization. By incorporating in vivo and fossil data, we reformulate the visual predation hypothesis of primate origins and thus offer new insights into major adaptive transformations in the primate skull.
Animals↗
A comment on: the instantaneous center of rotation during human jaw opening and its significance in interpreting the functional meaning of condylar translation (Chen, x., 1998, Am J phys anthropol 106:35-46)
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Biomechanical Phenomena↗
Biomechanics of torsion in the human mandible.
Comparative investigations of mandibular function among primates have relied upon elementary structural models to estimate states of masticatory stress and strain. In these studies, mandibular corpus morphology is idealized as a homogeneous, isotropic symmetrical body of invariant geometry, and this morphological abstraction is used to infer relative levels of stress and strain in the jaw. In reality, none of the limiting conditions assumed by these models is satisfied; consequently, it is prudent to ask whether this "textbook" engineering approach is valid for the inference of biomechanical behavior. In this study, the predictions of various geometric representations of the mandibular corpus are evaluated against strains recorded in a sample of human jaws loaded in torsion. Symmetrical geometrical models (including various "robusticity" shape indices), although convenient, are probably not consistently reliable for predicting the distribution of strains in the corpus. The experimental data suggest that variations in cortical thickness within sections play a significant role in determining the profile of relative strains. For comparative applications, characterization of the corpus as an asymmetrical hollow ellipse (i.e., with differing thickness of medial and lateral cortical plates) may provide a reasonable portrayal of relative strains.
Aged↗
Mandibular corpus strain in primates: further evidence for a functional link between symphyseal fusion and jaw-adductor muscle force.
Previous work indicates that compared to adult thick-tailed galagos, adult long-tailed macaques have much more bone strain on the balancing-side mandibular corpus during unilateral isometric molar biting (Hylander [1979a] J. Morphol. 159:253-296). Recently we have confirmed in these same two species the presence of similar differences in bone-strain patterns during forceful mastication. Moreover, we have also recorded mandibular bone strain patterns in adult owl monkeys, which are slightly smaller than the galago subjects. The owl monkey data indicate the presence of a strain pattern very similar to that recorded for macaques, and quite unlike that recorded for galagos. We interpret these bone-strain pattern differences to be importantly related to differences in balancing-side jaw-adductor muscle force recruitment patterns. That is, compared to galagos, macaques and owl monkeys recruit relatively more balancing-side jaw-adductor muscle force during forceful mastication. Unlike an earlier study (Hylander [1979b] J. Morphol. 160:223-240), we are unable to estimate the actual amount of working-side muscle force relative to balancing-side muscle force (i.e., the W/B muscle force ratio) in these species because we have no reliable estimate of magnitude, direction, and precise location of the bite force during mastication. A comparison of the mastication data with the earlier data recorded during isometric molar biting, however, supports the hypothesis that the two anthropoids have a small W/B jaw-adductor muscle force ratio in comparison to thick-tailed galagos. These data also support the hypothesis that increased recruitment of balancing-side jaw-adductor muscle force in anthropoids is functionally linked to the evolution of symphyseal fusion or strengthening. Moreover, these data refute the hypothesis that the recruitment pattern differences between macaques and thick-tailed galagos are due to allometric factors. Finally, although the evolution of symphyseal fusion in primates may be linked to increased stress associated with increased balancing-side muscle force, it is currently unclear as to whether the increased force is predominately vertically directed, transversely directed, or is a near equal combination of these two force components (cf. Ravosa and Hylander [1994] In Fleagle and Kay [eds.]: Anthropoid Origins. New York: Plenum, pp. 447-468).
Animals↗
EMG of the human flexor pollicis longus muscle: implications for the evolution of hominid tool use.
Modern humans possess a distinct and well-developed flexor pollicis longus muscle, an extrinsic thumb flexor which is "either rudimentary or absent" in great apes (Straus, 1942, p. 228). Previous workers (e.g., Napier, 1962; Susman, 1988) have related the origin of a well-developed flexor pollicis longus muscle to the acquisition of precision grasping and stone tool making capabilities in early hominids. The proposed functional association between flexor pollicis longus activity, precision grasping, and stone tool manufacture has, however, never been tested experimentally. This study uses electromyographic techniques (EMG) to investigate the role of flexor pollicis longus during a variety of tool making, tool using, and manipulatory behaviors in order to determine the functional and evolutionary significance of the human flexor pollicis longus muscle. Our results indicate that flexor pollicis longus is recruited during forceful tool using and stone tool making behaviors, regardless of the power or precision grip used to hold the tool. In particular, both stone tool use and stone tool making employing three- and four-jaw chuck precision grips elicit consistently high levels of FPL activity. Flexor pollicis longus activity increases most when resistance is increased to the thumb's volar pad during these hammering, cutting, and knapping behaviors. In contrast, we observed relatively low levels of flexor pollicis longus activity during the fine manipulation of food items, the making of slender wooden probes, and the use of these probes as tools. The paleontological, archaeological, and experimental data suggest that a well-developed flexor pollicis longus muscle functioned initially in the hominid lineage to stabilize the terminal pollical phalanx against loads applied to the thumb's apical pad during the frequent and forceful use of unmodified stones as tools.
Animals↗
In vivo bone strain patterns in the zygomatic arch of macaques and the significance of these patterns for functional interpretations of craniofacial form.
It has been proposed that the mammalian facial skeleton is optimized for countering or dissipating masticatory stress. As optimized load-bearing structures by definition exhibit maximum strength with a minimum amount of material, this hypothesis predicts that during chewing and biting there should be relatively high and near uniform amounts of strain throughout the facial skeleton. If levels of strain in certain areas of the facial skeleton are relatively low during these behaviors, this indicates that the amount of bone mass in these areas could be significantly reduced without resulting in the danger of structural failure due to repeated masticatory loads. Furthermore, and by definition, this indicates that these areas are not optimized for countering masticatory stress, and instead their overall morphology and concentration of bone mass has most likely been selected or influenced mainly by factors unrelated to the dissipation or countering of chewing and biting forces. An analysis of in vivo bone strain along the lateral aspect of the zygomatic arch of macaques indicates the clear absence of a high and near uniform strain environment throughout its extent. Instead, there is a steep strain gradient along the zygomatic arch, with the highest strains along its anterior portion, intermediate strains along its middle portion, and the lowest strains along its posterior portion. These data, in combination with earlier published data (Hylander et al., 1991), indicate that levels of functional strains during chewing and biting are highly variable from one region of the face to the next, and therefore it is unlikely that all facial bones are especially designed so as to minimize bone tissue and maximize strength for countering masticatory loads. Thus, the functional significance of the morphology of certain facial bones need not necessarily bear any important or special relationship to routine and habitual cyclical mechanical loads associated with chewing or biting. Furthermore, the presence of these steep strain gradients within the facial skeleton suggests that the amount of bone mass in the low-strain areas may be largely determined by factors unrelated to processes frequently referred to as "functional adaptation," or conversely, that the "optimal strain environment" of bone varies enormously throughout the facial skeleton (cf., Rubin et al., 1994). Based solely on anatomical considerations, it is likely that the zygomatic arch is bent in both the parasagittal and transverse planes and twisted about its long axis. Due to constraints on rosette position, the strain data are incapable of determining if one or more of these loading conditions predominate. Instead, the strain data simply provide limited support for the possible presence of all of these loading regimes. Finally, as the masseter muscle is concentrated along the anterior portion of the zygomatic arch and as the arch has fixed ends, the largest shearing forces and the largest bending and twisting moments are located along its anterior portion. This in turn explains why the largest strains are found along the anterior portion of the zygomatic arch.
Animals↗
Occlusal forces and mandibular bone strain: is the primate jaw "overdesigned"?
Finite element modelling of the function of the periodontium and surrounding alveolar bone suggests these tissues are subjected to unusually large strains in comparison with the bone of the basal mandibular corpus. These studies, in addition to certain experimental investigations, have led to the suggestion that the strains experienced in the basal mandibular corpus are not functionally important. Under this view, size and shape of the basal corpus are not functionally linked to masticatory forces. Since previous comparative investigations have been premised on the assumption that masticatory strains in the basal corpus are functionally important, the assertion that masticatory stresses are concentrated primarily in the alveolar process undermines the credibility of this body of work. The hypothesis that the biomechanical effects of masticatory forces are localized in the alveolar process can be evaluated by reference to a number of bone strain investigations, as well as through consideration of current understanding of bone biology and behavior. Experimental studies indicate that the effects of occlusal forces during mastication are quite apparent in alveolar bone, although relatively large strains are also observed in regions well-removed from a loaded alveolus. It is also apparent that both alveolar and basal mandibular bone are subject to bending and twisting strains associated not only with occlusal forces, but also with muscular and condylar reaction forces. The result is that strain levels in alveolar vs. basal bone may be roughly similar, in contradiction to some published theoretical models. Based on empirical evidence and theoretical considerations, it is premature to conclude that mandibular corpus size and shape are not functionally linked to the biomechanics of chewing and biting.
Animals↗
In vivo and in vitro bone strain in the owl monkey circumorbital region and the function of the postorbital septum.
Anthropoids and tarsiers are the only vertebrates possessing a postorbital septum. This septum, formed by the frontal, alisphenoid, and zygomatic bones, separates the orbital contents from the temporal muscles. Three hypotheses suggest that the postorbital septum evolved to resist stresses acting on the skull during mastication or incision. The facial-torsion hypothesis posits that the septum resists twisting of the face about a rostrocaudal axis during unilateral mastication; the transverse-bending hypothesis argues that the septum resists caudally directed forces acting at the lateral orbital margin during mastication or incision; and the tension hypothesis suggests that the septum resists ventrally directed components of masseter muscle force during mastication and incision. This study evaluates these hypotheses using in vitro and in vivo bone strain data recorded from the circumorbital region of owl monkeys. Incisor loading of an owl monkey skull in vitro bends the face upward in the sagittal plane, compressing the interorbital region rostrocaudally and "buckling" the lateral orbital walls. Unilateral loading of the toothrow in vitro also bends the face in the sagittal plane, compressing the interorbital region rostrocaudally and buckling the working side lateral orbital wall. When the lateral orbital wall is partially cut, so as to reduce the width of its attachment to the braincase, the following changes in circumorbital bone strain patterns occur. During loading of the incisors, lower bone strain magnitudes are recorded in the interorbital region and lateral orbital walls. In contrast, during unilateral loading of the P3, higher bone strain magnitudes are observed in the interorbital region, and generally lower bone strain magnitudes are observed in the lateral orbital walls. During unilateral loading of the M2, higher bone strain magnitudes are observed in both the interorbital region and in the lateral orbital wall ipsilateral to the loaded molar. Comparisons of the in vitro results with data gathered in vivo suggest that, during incision and unilateral mastication, the face is subjected to upward bending in the sagittal plane resulting in rostrocaudal compression of the interorbital region. Modeling the lateral orbital walls as curved plates suggests that during mastication the working side wall is buckled due to the dorsally directed component of the maxillary bite force which causes upward bending of the face in the sagittal plane. The balancing side lateral orbital wall may also be buckled due to upward bending of the face in the sagittal plane as well as being twisted by the caudoventrally directed components of the superficial masseter muscle force. The in vivo data do not exclude the possibility that the postorbital septum functions to improve the structural integrity of the postorbital bar during mastication. However, there is no reason to believe that a more robust postorbital bar could not also perform this function. Hypotheses stating that the postorbital septum originally evolved to reinforce the skull against routine masticatory loads must explain why, rather than evolving a postorbital septum, the stem anthropoids did not simply enlarge their postorbital bars.
Animals↗
Albert A. Dahlberg (1908-1993).
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Anthropology, Physical↗
The mechanical or metabolic function of secondary osteonal bone in the monkey Macaca fascicularis.
Secondary osteonal bone is believed by many to serve a mechanical function, altering the properties and/or orientation of bone in response to fluctuating mechanical demands or in the prevention and/or repair of fatigue microdamage. Based on this belief, secondary osteons should be concentrated mainly in regions experiencing high peak-strain conditions. Others contend that secondary osteonal bone functions primarily in meeting the body's calcium needs, and should be expected to form principally in low peak-strain regions so as to avoid compromising the mechanical strength of the bone. These two hypotheses were tested by examining the distribution of secondary osteonal bone in both relatively high- and low-strain regions of the macaque face. Previous strain-gauge studies have demonstrated a steep strain gradient in the macaque face, with relatively high peak strains in the anterior portion of the zygomatic arch and in the mandibular corpus. Relatively low peak strains have been found in the posterior portion of the zygomatic arch and supraorbital bar. Results presented here show that in the mature macaques, there is no consistent relation between newly forming secondary osteons (i.e. those labelled with fluorescent dyes) and peak strain levels. From these data it is concluded that, in the non-perturbed adult, either mechanical and metabolic factors contribute equally to the observed pattern or that metabolically driven remodelling is initiated without regard to strain levels. In immature macaques, however, the relation between peak strain levels and secondary osteon density is positive, with a significantly higher density of labelled osteons in the high strain regions. From these data it is concluded that, in immature individuals, mechanical factors are predominantly responsible for the initiation of secondary osteonal remodelling.
Age Factors↗
Jaw muscle function and wishboning of the mandible during mastication in macaques and baboons.
An analysis of in vivo bone strain indicates that the mandibular symphysis of macaques experiences lateral transverse bending or "wishboning" during the power stroke of mastication, and this loading regime results in relatively intense concentrations of stress along the lingual aspect of the symphysis (Hylander 1984, 1985). It has been hypothesized that peak wishboning of the macaque mandible, which probably occurs at the very end of the power stroke, that is, after the initial occurrence of maximum intercuspation, is associated with the late peak activity of the balancing-side deep masseter muscle coupled with the rapid decline in the activity of the balancing-side medial pterygoid and superficial masseter muscles (Hylander et al., 1987). The main purpose of this study is to do a detailed analysis of the electromyographic (EMG) activity of the deep and superficial masseter and medial pterygoid muscles so as to provide a better understanding of the external forces associated with wishboning. This was done by recording and analyzing EMG activity from the superficial masseter, deep masseter, and medial pterygoid muscles in macaques and baboons. EMG activity was recorded from bipolar fine-wire electrodes, and the data were quantified and analyzed using digital techniques. The EMG data clearly support our original observation that the balancing-side posterior deep masseter exhibits peak EMG activity relatively late in the power stroke at a time when activity is rapidly decreasing in the balancing-side medial pterygoid and superficial masseter muscles. Moreover, peak activity of the balancing-side deep masseter occurs at a time when the activity of the working-side deep and superficial masseter and medial pterygoid muscles are also decreasing. Thus, these data are consistent with the hypothesis that wishboning of the mandible is influenced significantly by the late occurrence of force from the balancing-side deep masseter. The EMG data also indicate that residual force from the relaxing superficial masseter may contribute significantly to wishboning. Finally, patterns of wishboning are not fully accounted for by our EMG analysis of the deep and superficial masseter and medial pterygoid muscles. This is probably because the lateral ptergyoids, which can either counter increased wishboning or actually cause reverse wishboning, were not included in our EMG analysis. The EMG data demonstrate that jaw-closing muscle recruitment patterns for macaques and baboons differ from those of humans. Nevertheless, in spite of these differences, it appears, based on previously published EMG data, that the human symphysis may also experience wishboning.(ABSTRACT TRUNCATED AT 400 WORDS)
Animals↗
Functional significance of an ossified mandibular symphysis: a reply.
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Animals↗