Jaw-muscle architecture and mandibular morphology influence relative maximum jaw gapes in the sexually dimorphic Macaca fascicularis.

Maximum jaw gape is a performance variable related to feeding and non-feeding oral behaviors, such as canine gape displays, and is influenced by several factors including jaw-muscle fiber architecture, muscle position on the skull, and jaw morphology. Maximum gape, jaw length, and canine height are strongly correlated across catarrhine primates, but relationships between gape and other aspects of masticatory apparatus morphology are less clear. We examine the effects of jaw-adductor fiber architecture, jaw-muscle leverage, and jaw form on gape in an intraspecific sample of sexually dimorphic crab-eating macaques (Macaca fascicularis). As M. fascicularis males have relatively larger maximum gapes than females, we predict that males will have muscle and jaw morphologies that facilitate large gape, but these morphologies may come at some expense to bite force. Male crab-eating macaques have relatively longer jaw-muscle fibers, masseters with decreased leverage, and temporomandibular joint morphologies that facilitate the production of wide gapes. Because relative canine height is correlated with maximum gape in catarrhines, and males have relatively longer canines than females, these results support the hypothesis that male M. fascicularis have experienced selection to increase maximum gape. The sexes do not differ in relative masseter physiologic cross-sectional area (PCSA), but males compensate for a potential trade-off between muscle excursion versus muscle force with increased temporalis weight and PCSA. This musculoskeletal configuration is likely functionally significant for behaviors involving aggressive canine biting and displays in male M. fascicularis and provides additional evidence supporting the multifactorial nature of the catarrhine masticatory apparatus. Our results have implications for the evolution of craniofacial morphology in catarrhine primates and reinforce the importance of evaluating additional factors other than feeding behavior and diet in analyses of masticatory apparatus form, function, and evolution.

Functional links between canine height and jaw gape in catarrhines with special reference to early hominins.

This study tests the hypothesis that decreased canine crown height in catarrhines is linked to (and arguably caused by) decreased jaw gape. Associations are characterized within and between variables such as upper and lower canine height beyond the occlusal plane (canine overlap), maximum jaw gape, and jaw length for 27 adult catarrhine species, including 539 living subjects and 316 museum specimens. The data demonstrate that most adult male catarrhines have relatively larger canine overlap dimensions and gapes than do conspecific females. For example, whereas male baboons open their jaws maximally more than 110% of jaw length, females open about 90%. Humans and hylobatids are the exceptions in that canine overlap is nearly the same in both the sexes and so is relative gape (ca. 65% for humans and 110% for hylobatids). A correlation analysis demonstrates that a large portion of relative gape (maximum gape/projected jaw length) is predicted by relative canine overlap (canine overlap/jaw length). Relative gape is mainly a function of jaw muscle position and/or jaw muscle-fiber length. All things equal, more rostrally positioned jaw muscles and/or shorter muscle fibers decrease gape and increase bite force during the power stroke of mastication, and the net benefit is to increase the mechanical efficiency during chewing. Similarly, more caudally positioned muscles and/or longer muscle fibers increase the amount of gape and decrease bite force. Overall, the data support the hypothesis that canine reduction in early hominins is functionally linked to decreased gape and increased mechanical efficiency of the jaws.

The feeding biomechanics and dietary ecology of Australopithecus africanus.

The African Plio-Pleistocene hominins known as australopiths evolved a distinctive craniofacial morphology that traditionally has been viewed as a dietary adaptation for feeding on either small, hard objects or on large volumes of food. A historically influential interpretation of this morphology hypothesizes that loads applied to the premolars during feeding had a profound influence on the evolution of australopith craniofacial form. Here, we test this hypothesis using finite element analysis in conjunction with comparative, imaging, and experimental methods. We find that the facial skeleton of the Australopithecus type species, A. africanus, is well suited to withstand premolar loads. However, we suggest that the mastication of either small objects or large volumes of food is unlikely to fully explain the evolution of facial form in this species. Rather, key aspects of australopith craniofacial morphology are more likely to be related to the ingestion and initial preparation of large, mechanically protected food objects like large nuts and seeds. These foods may have broadened the diet of these hominins, possibly by being critical resources that australopiths relied on during periods when their preferred dietary items were in short supply. Our analysis reconciles apparent discrepancies between dietary reconstructions based on biomechanics, tooth morphology, and dental microwear.

Functional and evolutionary significance of the recruitment and firing patterns of the jaw adductors during chewing in Verreaux's sifaka (Propithecus verreauxi).

Jaw-muscle electromyographic (EMG) patterns indicate that compared with thick-tailed galagos and ring-tailed lemurs, anthropoids recruit more relative EMG from their balancing-side deep masseter, and that this muscle peaks late in the power stroke. These recruitment and firing patterns in anthropoids are thought to cause the mandibular symphysis to wishbone (lateral transverse bending), resulting in relatively high symphyseal stresses. We test the hypothesis that living strepsirrhines with robust, partially fused symphyses have muscle recruitment and firing patterns more similar to anthropoids, unlike those strepsirrhines with highly mobile unfused symphyses. Electromyographic (EMG) activity of the superficial and deep masseter, anterior and posterior temporalis, and medial pterygoid muscles were recorded in four dentally adult Verreaux's sifakas (Propithecus verreauxi). As predicted, we find that sifaka motor patterns are more similar to anthropoids. For example, among sifakas, recruitment levels of the balancing-side (b-s) deep masseter are high, and the b-s deep masseter fires late during the power stroke. As adult sifakas often exhibit nearly complete symphyseal fusion, these data support the hypothesis that the evolution of symphyseal fusion in primates is functionally linked to wishboning. Furthermore, these data provide compelling evidence for the convergent evolution of the wishboning motor patterns in anthropoids and sifakas.

Mandibular corpus bone strain in goats and alpacas: implications for understanding the biomechanics of mandibular form in selenodont artiodactyls.

The goal of this study is to clarify the functional and biomechanical relationship between jaw morphology and in vivo masticatory loading in selenodont artiodactyls. We compare in vivo strains from the mandibular corpus of goats and alpacas to predicted strain patterns derived from biomechanical models for mandibular corpus loading during mastication. Peak shear strains in both species average 600-700 microepsilon on the working side and approximately 450 microepsilon on the balancing side. Maximum principal tension in goats and alpacas is directed at approximately 30 degrees dorsocaudally relative to the long axis of the corpus on the working side and approximately perpendicular to the long axis on the balancing side. Strain patterns in both species indicate primarily torsion of the working-side corpus about the long axis and parasagittal bending and/or lateral transverse bending of the balancing-side corpus. Interpretation of the strain patterns is consistent with comparative biomechanical analyses of jaw morphology suggesting that in goats, the balancing-side mandibular corpus is parasagittally bent whereas in alpacas it experiences lateral transverse bending. However, in light of higher working-side corpus strains, biomechanical explanations of mandibular form also need to consider that torsion influences relative corpus size and shape. Furthermore, the complex combination of loads that occur along the selenodont artiodactyl mandibular corpus during the power stroke has two implications. First, added clarification of these loading patterns requires in vivo approaches for elucidating biomechanical links between mandibular corpus morphology and masticatory loading. Second, morphometric approaches may be limited in their ability to accurately infer masticatory loading regimes of selenodont artiodactyl jaws.

From movement to models: a tribute to professor Alan G. Hannam.

This tribute article to Professor Alan G. Hannam is based on 7 presentations for him at the July 1, 2008 symposium honoring 3 "giants" in orofacial neuroscience: Professors B. J. Sessle, J. P. Lund, and A. G. Hannam. This tribute to Hannam's outstanding career draws examples from his 40-year academic career and spans topics from human evolution to complex modeling of the craniomandibular system. The first presentation by W. Hylander provides a plausible answer to the functional and evolutionary significance of canine reduction in hominins. The second presentation, by A. McMillan, describes research activities in the field of healthy aging, including findings that intensity-modulated radiotherapy improves the health condition and quality of life of people with nasopharyngeal carcinoma in comparison to conventional radiotherapy. The developments in dental imaging are summarized in the third paper by E. Lam, and an overview of the bite force magnitude and direction while clenching is described in the fourth paper by M. Watanabe. The last 3 contributions by G. Langenbach, I. Staveness, and C. Peck deal with the topic of bone remodeling as well as masticatory system modeling, which was Hannam's main research interest in recent years. These contributions show the considerable advancements that have been made in the last decade under Hannam's drive, in particular the development of an interactive model comprising, in addition to the masticatory system, also the upper airways. The final section of the article includes a final commentary from Professor Hannam.

Regional variation in IIM myosin heavy chain expression in the temporalis muscle of female and male baboons (Papio anubis).

OBJECTIVE: The purpose of this study was to determine whether high amounts of fast/type II myosin heavy chain (MyHC) in the superficial as compared to the deep temporalis muscle of adult female and male baboons (Papio anubis) correlates with published data on muscle function during chewing. Electromyographic (EMG) data show a regional specialization in activation from low to high amplitude activity during hard/tough object chewing cycles in the baboon superficial temporalis.(48,49) A positive correlation between fast/type II MyHC amount and EMG activity will support the high occlusal force hypothesis. DESIGN: Deep anterior temporalis (DAT), superficial anterior temporalis (SAT), and superficial posterior temporalis (SPT) muscle samples were analyzed using SDS-PAGE gel electrophoresis to test the prediction that SAT and SPT will show high amounts of fast/type II MyHC compared to DAT. Serial muscle sections were incubated against NOQ7.5.4D and MY32 antibodies to determine the breadth of slow/type I versus fast/type II expression within each section. RESULTS: Type I and type IIM MyHCs comprise nearly 100% of the MyHCs in the temporalis muscle. IIM MyHC was the overwhelmingly predominant fast MyHC, though there was a small amount of type IIA MyHC (≤5%) in DAT in two individuals. SAT and SPT exhibited a fast/type II phenotype and contained large amounts of IIM MyHC whereas DAT exhibited a type I/type II (hybrid) phenotype and contained a significantly greater proportion of MyHC-I. MyHC-I expression in DAT was sexually dimorphic as it was more abundant in females. CONCLUSIONS: The link between the distribution of IIM MyHC and high relative EMG amplitudes in SAT and SPT during hard/tough object chewing cycles is evidence of regional specialization in fibre type to generate high occlusal forces during chewing. The high proportion of MyHC-I in DAT of females may be related to a high frequency of individual fibre recruitment in comparison to males.

A preliminary analysis of correlations between chewing motor patterns and mandibular morphology across mammals.

The establishment of a publicly-accessible repository of physiological data on feeding in mammals, the Feeding Experiments End-user Database (FEED), along with improvements in reconstruction of mammalian phylogeny, significantly improves our ability to address long-standing questions about the evolution of mammalian feeding. In this study, we use comparative phylogenetic methods to examine correlations between jaw robusticity and both the relative recruitment and the relative time of peak activity for the superficial masseter, deep masseter, and temporalis muscles across 19 mammalian species from six orders. We find little evidence for a relationship between jaw robusticity and electromyographic (EMG) activity for either the superficial masseter or temporalis muscles across mammals. We hypothesize that future analyses may identify significant associations between these physiological and morphological variables within subgroups of mammals that share similar diets, feeding behaviors, and/or phylogenetic histories. Alternatively, the relative peak recruitment and timing of the balancing-side (i.e., non-chewing-side) deep masseter muscle (BDM) is significantly negatively correlated with the relative area of the mandibular symphysis across our mammalian sample. This relationship exists despite BDM activity being associated with different loading regimes in the symphyses of primates compared to ungulates, suggesting a basic association between magnitude of symphyseal loads and symphyseal area among these mammals. Because our sample primarily represents mammals that use significant transverse movements during chewing, future research should address whether the correlations between BDM activity and symphyseal morphology characterize all mammals or should be restricted to this "transverse chewing" group. Finally, the significant correlations observed in this study suggest that physiological parameters are an integrated and evolving component of feeding across mammals.

A preliminary analysis of correlated evolution in Mammalian chewing motor patterns.

Descriptive and quantitative analyses of electromyograms (EMG) from the jaw adductors during feeding in mammals have demonstrated both similarities and differences among species in chewing motor patterns. These observations have led to a number of hypotheses of the evolution of motor patterns, the most comprehensive of which was proposed by Weijs in 1994. Since then, new data have been collected and additional hypotheses for the evolution of motor patterns have been proposed. Here, we take advantage of these new data and a well-resolved species-level phylogeny for mammals to test for the correlated evolution of specific components of mammalian chewing motor patterns. We focus on the evolution of the coordination of working-side (WS) and balancing-side (BS) jaw adductors (i.e., Weijs' Triplets I and II), the evolution of WS and BS muscle recruitment levels, and the evolution of asynchrony between pairs of muscles. We converted existing chewing EMG data into binary traits to incorporate as much data as possible and facilitate robust phylogenetic analyses. We then tested hypotheses of correlated evolution of these traits across our phylogeny using a maximum likelihood method and the Bayesian Markov Chain Monte Carlo method. Both sets of analyses yielded similar results highlighting the evolutionary changes that have occurred across mammals in chewing motor patterns. We find support for the correlated evolution of (1) Triplets I and II, (2) BS deep masseter asynchrony and Triplets I and II, (3) a relative delay in the activity of the BS deep masseter and a decrease in the ratio of WS to BS muscle recruitment levels, and (4) a relative delay in the activity of the BS deep masseter and a delay in the activity of the BS posterior temporalis. In contrast, changes in relative WS and BS activity levels across mammals are not correlated with Triplets I and II. Results from this work can be integrated with dietary and morphological data to better understand how feeding and the masticatory apparatus have evolved across mammals in the context of new masticatory demands.

Patterns of variation across primates in jaw-muscle electromyography during mastication.

Biologists that study mammals continue to discuss the evolution of and functional variation in jaw-muscle activity during chewing. A major barrier to addressing these issues is collecting sufficient in vivo data to adequately capture neuromuscular variation in a clade. We combine data on jaw-muscle electromyography (EMG) collected during mastication from 14 species of primates and one of treeshrews to assess patterns of neuromuscular variation in primates. All data were collected and analyzed using the same methods. We examine the variance components for EMG parameters using a nested ANOVA design across successive hierarchical factors from chewing cycle through species for eight locations in the masseter and temporalis muscles. Variation in jaw-muscle EMGs was not distributed equally across hierarchical levels. The timing of peak EMG activity showed the largest variance components among chewing cycles. Relative levels of recruitment of jaw muscles showed the largest variance components among chewing sequences and cycles. We attribute variation among chewing cycles to (1) changes in food properties throughout the chewing sequence, (2) variation in bite location, and (3) the multiple ways jaw muscles can produce submaximal bite forces. We hypothesize that variation among chewing sequences is primarily related to variation in properties of food. The significant proportion of variation in EMGs potentially linked to food properties suggests that experimental biologists must pay close attention to foods given to research subjects in laboratory-based studies of feeding. The jaw muscles exhibit markedly different variance components among species suggesting that primate jaw muscles have evolved as distinct functional units. The balancing-side deep masseter (BDM) exhibits the most variation among species. This observation supports previous hypotheses linking variation in the timing and activation of the BDM to symphyseal fusion in anthropoid primates and in strepsirrhines with robust symphyses. The working-side anterior temporalis shows a contrasting pattern with little variation in timing and relative activation across primates. The consistent recruitment of this muscle suggests that primates have maintained their ability to produce vertical jaw movements and force in contrast to the evolutionary changes in transverse occlusal forces driven by the varying patterns of activation in the BDM.

Masticatory motor patterns in ungulates: a quantitative assessment of jaw-muscle coordination in goats, alpacas and horses.

We investigated patterns of jaw-muscle coordination during rhythmic mastication in three species of ungulates displaying the marked transverse jaw movements typical of many large mammalian herbivores. In order to quantify consistent motor patterns during chewing, electromyograms were recorded from the superficial masseter, deep masseter, posterior temporalis and medial pterygoid muscles of goats, alpacas and horses. Timing differences between muscle pairs were evaluated in the context of an evolutionary model of jaw-muscle function. In this model, the closing and food reduction phases of mastication are primarily controlled by two distinct muscle groups, triplet I (balancing-side superficial masseter and medial pterygoid and working-side posterior temporalis) and triplet II (working-side superficial masseter and medial pterygoid and balancing-side posterior temporalis), and the asynchronous activity of the working- and balancing-side deep masseters. The three species differ in the extent to which the jaw muscles are coordinated as triplet I and triplet II. Alpacas, and to a lesser extent, goats, exhibit the triplet pattern whereas horses do not. In contrast, all three species show marked asynchrony of the working-side and balancing-side deep masseters, with jaw closing initiated by the working-side muscle and the balancing-side muscle firing much later during closing. However, goats differ from alpacas and horses in the timing of the balancing-side deep masseter relative to the triplet II muscles. This study highlights interspecific differences in the coordination of jaw muscles to influence transverse jaw movements and the production of bite force in herbivorous ungulates.

Modulation of intra-oral processing in mammals and lepidosaurs.

The mammalian masticatory apparatus is distinguished from the intra-oral processing systems of other amniotes by a number of morphological and functional features, including transverse movements of the teeth during the power stroke, precise occlusion, suspension of the teeth in the socket by a periodontal ligament, diphyodonty (reduction to two generations of teeth), a hard palate, and the presence of a single bone (the dentary) in the lower jaw which articulates with the skull at the temporomandibular jaw joint. The evolution of these features is commonly argued to have improved the efficiency of food processing in the oral cavity. The present aricle highlights the existence in mammals of the fusimotor system and afferent fibers from the periodontal ligament through which the CNS modulates the responses by the muscle spindles. Published data suggest that the fusimotor system and the periodontal afferents are important components in feed-forward (or anticipatory) control of chewing behavior. We hypothesize that this feed-forward control is used to maintain relatively constant cycle lengths in mammals in the face of intra-sequence and inter-sequence variation in material properties of the food, and that this enables them to maintain a higher average chewing frequency than that of lizards. These predictions were evaluated using data on mean cycle length and its variance from the literature and from our own files. On average, mammals have less variable cycle lengths than do lizards and shorter cycle lengths than do lizards of similar size. We hypothesize that by decreasing variance in cycle length, presumably close to the natural frequency of their feeding systems, mammals minimize energy expenditure during chewing, allowing them to chew for longer, thereby maintaining the high rates of food intake required for their high metabolic rates.

Modulation of mandibular loading and bite force in mammals during mastication.

Modulation of force during mammalian mastication provides insight into force modulation in rhythmic, cyclic behaviors. This study uses in vivo bone strain data from the mandibular corpus to test two hypotheses regarding bite force modulation during rhythmic mastication in mammals: (1) that bite force is modulated by varying the duration of force production, or (2) that bite force is modulated by varying the rate at which force is produced. The data sample consists of rosette strain data from 40 experiments on 11 species of mammals, including six primate genera and four nonprimate species: goats, pigs, horses and alpacas. Bivariate correlation and multiple regression methods are used to assess relationships between maximum (epsilon(1)) and minimum (epsilon(2)) principal strain magnitudes and the following variables: loading time and mean loading rate from 5% of peak to peak strain, unloading time and mean unloading rate from peak to 5% of peak strain, chew cycle duration, and chew duty factor. Bivariate correlations reveal that in the majority of experiments strain magnitudes are significantly (P<0.001) correlated with strain loading and unloading rates and not with strain loading and unloading times. In those cases when strain magnitudes are also correlated with loading times, strain magnitudes are more highly correlated with loading rate than loading time. Multiple regression analyses reveal that variation in strain magnitude is best explained by variation in loading rate. Loading time and related temporal variables (such as overall chew cycle time and chew duty factor) do not explain significant amounts of additional variance. Few and only weak correlations were found between strain magnitude and chew cycle time and chew duty factor. These data suggest that bite force modulation during rhythmic mastication in mammals is mainly achieved by modulating the rate at which force is generated within a chew cycle, and less so by varying temporal parameters. Rate modulation rather than time modulation may allow rhythmic mastication to proceed at a relatively constant frequency, simplifying motor control computation.

Phase II jaw movements and masseter muscle activity during chewing in Papio anubis.

It was proposed that the power stroke in primates has two distinct periods of occlusal contact, each with a characteristic motion of the mandibular molars relative to the maxillary molars. The two movements are called phase I and phase II, and they occur sequentially in that order (Kay and Hiiemae [1974] Am J. Phys. Anthropol. 40:227-256, Kay and Hiiemae [1974] Prosimian Biology, Pittsburgh: University of Pittsburgh Press, p. 501-530). Phase I movement is said to be associated with shearing along a series of crests, producing planar phase I facets and crushing on surfaces on the basins of the molars. Phase I terminates in centric occlusion. Phase II movement is said to be associated with grinding along the same surfaces that were used for crushing at the termination of phase I. Hylander et al. ([1987] Am J. Phys. Anthropol. 72:287-312; see also Hiiemae [1984] Food Acquisition and Processing, London: Academic Press, p. 257-281; Hylander and Crompton [1980] Am J. Phys. Anthropol. 52:239-251, [1986] Arch. Oral. Biol. 31:841-848) analyzed data on macaques and suggested that phase II movement may not be nearly as significant for food breakdown as phase I movement simply because, based on the magnitude of mandibular bone strain patterns, adductor muscle and occlusal forces are likely negligible during movement out of centric occlusion. Our goal is to better understand the functional significance of phase II movement within the broader context of masticatory kinematics during the power stroke. We analyze vertical and transverse mandibular motion and relative activity of the masseter and temporalis muscles during phase I and II movements in Papio anubis. We test whether significant muscle activity and, by inference, occlusal force occurs during phase II movement. We find that during phase II movement, there is negligible force developed in the superficial and deep masseter and the anterior and posterior temporalis muscles. Furthermore, mandibular movements are small during phase II compared to phase I. These results suggest that grinding during phase II movement is of minimal importance for food breakdown, and that most food breakdown on phase II facets occurs primarily at the end of phase I movement (i.e., crushing during phase I movement). We note, however, that depending on the orientation of phase I facets, significant grinding also occurs along phase I facets during phase I.

Masseter electromyography during chewing in ring-tailed lemurs (Lemur catta).

We examined masseter recruitment and firing patterns during chewing in four adult ring-tailed lemurs (Lemur catta), using electromyography (EMG). During chewing of tougher foods, the working-side superficial masseter tends to show, on average, 1.7 times more scaled EMG activity than the balancing-side superficial masseter. The working-side deep masseter exhibits, on average, 2.4 times the scaled EMG activity of the balancing-side deep masseter. The relatively larger activity in the working-side muscles suggests that ring-tailed lemurs recruit relatively less force from their balancing-side muscles during chewing. The superficial masseter working-to-balancing-side (W/B) ratio for lemurs overlaps with W/B ratios from anthropoid primates. In contrast, the lemur W/B ratio for the deep masseter is more similar to that of greater galagos, while both are significantly larger than W/B ratios of anthropoids. Because ring-tailed lemurs have unfused and hence presumably weaker symphyses, these data are consistent with the symphyseal fusion-muscle recruitment hypothesis stating that symphyseal fusion in anthropoids provides increased strength for resisting forces created by the balancing-side jaw muscles during chewing. Among the masseter muscles of ring-tailed lemurs, the working-side deep masseter peaks first on average, followed in succession by the balancing-side deep masseter, balancing-side superficial masseter, and finally the working-side superficial masseter. Ring-tailed lemurs are similar to greater galagos in that their balancing-side deep masseter peaks well before their working-side superficial masseter. We see the opposite pattern in anthropoids, where the balancing-side deep masseter peaks, on average, after the working-side superficial masseter. This late activity of the balancing-side deep masseter in anthropoids is linked to lateral-transverse bending, or wishboning, of their mandibular symphyses. Subsequently, the stresses incurred during wishboning are hypothesized to be a proximate reason for strengthening, and hence fusion, of the anthropoid symphysis. Thus, the absence of this muscle-firing pattern in ring-tailed lemurs with their weaker, unfused symphyses provides further correlational support for the symphyseal fusion late-acting balancing-side deep masseter hypothesis linking wishboning and symphyseal strengthening in anthropoids. The early peak activity of the working-side deep masseter in ring-tailed lemurs is unlike galagos and most similar to the pattern seen in macaques and baboons. We hypothesize that this early activity of the working-side deep masseter moves the lower jaw both laterally toward the working side and vertically upward, to position it for the upcoming power stroke. From an evolutionary perspective, the differences in peak firing times for the working-side deep masseter between ring-tailed lemurs and greater galagos indicate that deep masseter firing patterns are not conserved among strepsirrhines.

Jaw-muscle electromyography during chewing in Belanger's treeshrews (Tupaia belangeri).

We examined masseter and temporalis recruitment and firing patterns during chewing in five male Belanger's treeshrews (Tupaia belangeri), using electromyography (EMG). During chewing, the working-side masseters tend to show almost three times more scaled EMG activity than the balancing-side masseters. Similarly, the working-side temporalis muscles have more than twice the scaled EMG activity of the balancing-side temporalis. The relatively higher activity in the working-side muscles suggests that treeshrews recruit less force from their balancing-side muscles during chewing. Most of the jaw-closing muscles in treeshrews can be sorted into an early-firing or late-firing group, based on occurrence of peak activity during the chewing cycle. Specifically, the first group of jaw-closing muscles to reach peak activity consists of the working-side anterior and posterior temporalis and the balancing-side superficial masseter. The balancing-side anterior and posterior temporalis and the working-side superficial masseter peak later in the power stroke. The working-side deep masseter peaks, on average, slightly before the working-side superficial masseter. The balancing-side deep masseter typically peaks early, at about the same time as the balancing-side superficial masseter. Thus, treeshrews are unlike nonhuman anthropoids that peak their working-side deep masseters early and their balancing-side deep masseters late in the power stroke. Because in anthropoids the late firing of the balancing-side deep masseter contributes to wishboning of the symphysis, the treeshrew EMG data suggest that treeshrews do not routinely wishbone their symphyses during chewing. Based on the treeshrew EMG data, we speculate that during chewing, primitive euprimates 1) recruited more force from the working-side jaw-closing muscles as compared to the balancing-side muscles, 2) fired an early group of jaw-closing muscles followed by a second group of muscles that peaked later in the power stroke, 3) did not fire their working-side deep masseter significantly earlier than their working-side superficial masseter, and 4) did not routinely fire their balancing-side deep masseter after the working-side superficial masseter.

Temporalis function in anthropoids and strepsirrhines: an EMG study.

The major purpose of this study is to analyze anterior and posterior temporalis muscle force recruitment and firing patterns in various anthropoid and strepsirrhine primates. There are two specific goals for this project. First, we test the hypothesis that in addition to transversely directed muscle force, the evolution of symphyseal fusion in primates may also be linked to vertically directed balancing-side muscle force during chewing (Hylander et al. [2000] Am. J. Phys. Anthropol. 112:469-492). Second, we test the hypothesis of whether strepsirrhines retain the hypothesized primitive mammalian condition for the firing of the anterior temporalis, whereas anthropoids have the derived condition (Weijs [1994] Biomechanics of Feeding in Vertebrates; Berlin: Springer-Verlag, p. 282-320). Electromyographic (EMG) activities of the left and right anterior and posterior temporalis muscles were recorded and analyzed in baboons, macaques, owl monkeys, thick-tailed galagos, and ring-tailed lemurs. In addition, as we used the working-side superficial masseter as a reference muscle, we also recorded and analyzed EMG activity of the left and right superficial masseter in these primates. The data for the anterior temporalis provided no support for the hypothesis that symphyseal fusion in primates is linked to vertically directed jaw muscle forces during mastication. Thus, symphyseal fusion in primates is most likely mainly linked to the timing and recruitment of transversely directed forces from the balancing-side deep masseter (Hylander et al. [2000] Am. J. Phys. Anthropol. 112:469-492). In addition, our data demonstrate that the firing patterns for the working- and balancing-side anterior temporalis muscles are near identical in both strepsirrhines and anthropoids. Their working- and balancing-side anterior temporalis muscles fire asynchronously and reach peak activity during the power stroke. Similarly, their working- and balancing-side posterior temporalis muscles also fire asynchronously and reach peak activity during the power stroke. Compared to these strepsirrhines, however, the balancing-side posterior temporalis of anthropoids appears to have a relatively delayed firing pattern. Moreover, based on their smaller W/B ratios, anthropoids demonstrate a relative increase in muscle-force recruitment of the balancing-side posterior temporalis. This in turn suggests that anthropoids may emphasize the duration and magnitude of the power stroke during mastication. This hypothesis, however, requires additional testing. Furthermore, during the latter portion of the power stroke, the late activity of the balancing-side posterior temporalis of anthropoids apparently assists the balancing-side deep masseter in driving the working-side molars through the terminal portion of occlusion.

A biomechanical analysis of skull form in gum-harvesting galagids.

Among primates, some highly gummivorous species habitually gouge trees to elicit exudate flow whereas others scrape the hardened gums from trees. These foraging behaviors are thought to require high external forces at the anterior dentition. In this study, we test whether skull form in gouging and scraping galagids corresponds to this suggested need to produce these higher external forces and to resist increased internal loads in the jaws. We find few consistent morphological patterns linking skull form and the generation of high forces during gouging. However, there is some tendency for gougers and scrapers to show increased load resistance capabilities in their mandibles. Future research on the mechanical properties of trees exploited by these species and on jaw function during gouging and scraping will improve our understanding of the mechanical demands of gum feeding on the galagid skull form.

Comparative functional analysis of skull morphology of tree-gouging primates.

Many primates habitually feed on tree exudates such as gums and saps. Among these exudate feeders, Cebuella pygmaea, Callithrix spp., Phaner furcifer, and most likely Euoticus elegantulus elicit exudate flow by biting into trees with their anterior dentition. We define this behavior as gouging. Beyond the recent publication by Dumont ([1997] Am J Phys Anthropol 102:187-202), there have been few attempts to address whether any aspect of skull form in gouging primates relates to this specialized feeding behavior. However, many researchers have proposed that tree gouging results in larger bite force, larger internal skull loads, and larger jaw gapes in comparison to other chewing and biting behaviors. If true, then we might expect primate gougers to exhibit skull modifications that provide increased abilities to produce bite forces at the incisors, withstand loads in the skull, and/or generate large gapes for gouging. We develop 13 morphological predictions based on the expectation that gouging involves relatively large jaw forces and/or jaw gapes. We compare skull shapes for P. furcifer to five cheirogaleid taxa, E. elegantulus to six galagid species, and C. jacchus to two tamarin species, so as to assess whether gouging primates exhibit these predicted morphological shapes. Our results show little morphological evidence for increased force-production or load-resistance abilities in the skulls of these gouging primates. Conversely, these gougers tend to have skull shapes that are advantageous for creating large gapes. For example, all three gouging species have significantly lower condylar heights relative to the toothrow at a given mandibular length in comparison with closely related, nongouging taxa. Lowering the height of the condyle relative to the mandibular toothrow should reduce the stretching of the masseters and medial pterygoids during jaw opening, as well as position the mandibular incisors more anteriorly at wide jaw gapes. In other words, the lower incisors will follow a more vertical trajectory during both jaw opening and closing. We predict, based on these findings, that tree-gouging primates do not generate unusually large forces, but that they do use relatively large gapes during gouging. Of course, in vivo data on jaw forces and jaw gapes are required to reliably assess skull functions during gouging.