A new biomechanical model of the mammal jaw based on load path analysis.

The primary function of the tetrapod jaw is to transmit jaw muscle forces to bite points. The routes of force transfer in the jaw have never been studied but can be quantified using load paths - the shortest, stiffest routes from regions of force application to support constraints. Here, we use load path analysis to map force transfer from muscle attachments to bite point and jaw joint, and to evaluate how different configurations of trabecular and cortical bone affect load paths. We created three models of the mandible of the Virginia opossum, Didelphis virginiana, each with a cortical bone shell, but with different material properties for the internal spaces: (1) a cortical-trabecular model, in which the interior space is modeled with bulk properties of trabecular bone; (2) a cortical-hollow model, in which trabeculae and mandibular canal are modeled as hollow; and (3) a solid-cortical model, in which the interior is modeled as cortical bone. The models were compared with published in vivo bite force and bone strain data, and the load paths calculated for each model. The trabecular model, which is preferred because it most closely approximates the actual morphology, was best validated by in vivo data. In all three models, the load path was confined to cortical bone, although its route within the cortex varied depending on the material properties of the inner model. Our analysis shows that most of the force is transferred through the cortical, rather than trabecular bone, and highlights the potential of load path analysis for understanding form-function relationships in the skeleton.

The biomechanics of chewing and suckling in the infant: A potential mechanism for physiologic metopic suture closure.

Craniosynostosis is a condition with neurologic and aesthetic sequelae requiring invasive surgery. Understanding its pathobiology requires familiarity with the processes underlying physiologic suture closure. Animal studies have shown that cyclical strain from chewing and suckling influences the closure of cranial vault sutures, especially the metopic, an important locus of craniosynostosis. However, there are no human data correlating strain patterns during chewing and suckling with the physiologically early closure pattern of the metopic suture. Furthermore, differences in craniofacial morphology make it challenging to directly extrapolate animal findings to humans. Eight finite-element analysis (FEA) models were built from craniofacial computer tomography (CT) scans at varying stages of metopic suture closure, including two with isolated non-syndromic metopic craniosynostosis. Muscle forces acting on the cranium during chewing and suckling were simulated using subject-specific jaw muscle cross-sectional areas. Chewing and suckling induced tension at the metopic and sagittal sutures, and compressed the coronal, lambdoid, and squamous sutures. Relative to other cranial vault sutures, the metopic suture experienced larger magnitudes of axial strain across the suture and a lower magnitude of shear strain. Strain across the metopic suture decreased during suture closure, but other sutures were unaffected. Strain patterns along the metopic suture mirrored the anterior to posterior sequence of closure: strain magnitudes were highest at the glabella and decreased posteriorly, with minima at the nasion and the anterior fontanelle. In models of physiologic suture closure, increased degree of metopic suture closure correlated with higher maximum principal strains across the frontal bone and mid-face, a strain regime not observed in models of severe metopic craniosynostosis. In summary, our work provides human evidence that bone strain patterns from chewing and suckling correlate with the physiologically early closure pattern of the metopic suture, and that deviations from physiologic strain regimes may contribute to clinically observed craniofacial dysmorphism.

Biomechanical and Cortical Control of Tongue Movements During Chewing and Swallowing.

Tongue function is vital for chewing and swallowing and lingual dysfunction is often associated with dysphagia. Better treatment of dysphagia depends on a better understanding of hyolingual morphology, biomechanics, and neural control in humans and animal models. Recent research has revealed significant variation among animal models in morphology of the hyoid chain and suprahyoid muscles which may be associated with variation in swallowing mechanisms. The recent deployment of XROMM (X-ray Reconstruction of Moving Morphology) to quantify 3D hyolingual kinematics has revealed new details on flexion and roll of the tongue during chewing in animal models, movements similar to those used by humans. XROMM-based studies of swallowing in macaques have falsified traditional hypotheses of mechanisms of tongue base retraction during swallowing, and literature review suggests that other animal models may employ a diversity of mechanisms of tongue base retraction. There is variation among animal models in distribution of hyolingual proprioceptors but how that might be related to lingual mechanics is unknown. In macaque monkeys, tongue kinematics-shape and movement-are strongly encoded in neural activity in orofacial primary motor cortex, giving optimism for development of brain-machine interfaces for assisting recovery of lingual function after stroke. However, more research on hyolingual biomechanics and control is needed for technologies interfacing the nervous system with the hyolingual apparatus to become a reality.

Ontogenetic changes in bite force and gape in tufted capuchins.

Bite force and gape are two important performance metrics of the feeding system, and these metrics are inversely related for a given muscle size because of fundamental constraints in sarcomere length-tension relationships. How these competing performance metrics change in developing primates is largely unknown. Here, we quantified in vivo bite forces and gapes across ontogeny and examined these data in relation to body mass and cranial measurements in captive tufted capuchins, Sapajus spp. Bite force and gape were also compared across geometric and mechanical properties of mechanically challenging foods to investigate relationships between bite force, gape and food accessibility (defined here as the ability to breach shelled nuts). Bite forces at a range of gapes and feeding behavioral data were collected from a cross-sectional ontogenetic series of 20 captive and semi-wild tufted capuchins at the Núcleo de Procriação de Macacos-Prego Research Center in Araçatuba, Brazil. These data were paired with body mass, photogrammetric measures of jaw length and facial width, and food geometric and material properties. Tufted capuchins with larger body masses had absolutely higher in vivo bite forces and gapes, and animals with wider faces had absolutely higher bite forces. Bite forces and gapes were significantly smaller in juveniles compared with subadults and adults. These are the first primate data to empirically demonstrate the gapes at which maximum active bite force is generated and to demonstrate relationships to food accessibility. These data advance our understanding of how primates meet the changing performance demands of the feeding system during development.

Mechanical compensation in the evolution of the early hominin feeding apparatus.

Australopiths, a group of hominins from the Plio-Pleistocene of Africa, are characterized by derived traits in their crania hypothesized to strengthen the facial skeleton against feeding loads and increase the efficiency of bite force production. The crania of robust australopiths are further thought to be stronger and more efficient than those of gracile australopiths. Results of prior mechanical analyses have been broadly consistent with this hypothesis, but here we show that the predictions of the hypothesis with respect to mechanical strength are not met: some gracile australopith crania are as strong as that of a robust australopith, and the strength of gracile australopith crania overlaps substantially with that of chimpanzee crania. We hypothesize that the evolution of cranial traits that increased the efficiency of bite force production in australopiths may have simultaneously weakened the face, leading to the compensatory evolution of additional traits that reinforced the facial skeleton. The evolution of facial form in early hominins can therefore be thought of as an interplay between the need to increase the efficiency of bite force production and the need to maintain the structural integrity of the face.

Morphological disparity and evolutionary transformations in the primate hyoid apparatus.

The hyoid apparatus plays an integral role in swallowing, respiration, and vocalization in mammals. Most placental mammals have a rod-shaped basihyal connected to the basicranium via both soft tissues and a mobile bony chain-the anterior cornu-whereas anthropoid primates have broad, shield-like or even cup-shaped basihyals suspended from the basicranium by soft tissues only. How the unique anthropoid hyoid morphology evolved is unknown, and hyoid morphology of nonanthropoid primates is poorly documented. Here we use phylogenetic comparative methods and linear morphometrics to address knowledge gaps in hyoid evolution among primates and their euarchontan outgroups. We find that dermopterans have variable reduction of cornu elements. Cynocephalus volans are sexually dimorphic in hyoid morphology. Tupaia and all lemuroids except Daubentonia have a fully ossified anterior cornu connecting a rod-shaped basihyal to the basicranium; this is the ancestral mammalian pattern that is also characteristic of the last common ancestor of Primates. Haplorhines exhibit a reduced anterior cornu, and anthropoids underwent further increase in basihyal aspect ratio values and in relative basihyal volume. Convergent with haplorhines, lorisoid strepsirrhines independently evolved a broad basihyal and reduced anterior cornua. While a reduced anterior cornu is hypothesized to facilitate vocal tract lengthening and lower formant frequencies in some mammals, our results suggest vocalization adaptations alone are unlikely to drive the iterative reduction of anterior cornua within Primates. Our new data on euarchontan hyoid evolution provide an anatomical basis for further exploring the form-function relationships of the hyoid across different behaviors, including vocalization, chewing, and swallowing.

Suction feeding biomechanics of Polypterus bichir: investigating linkage mechanisms and the contributions of cranial kinesis to oral cavity volume change.

Many fishes use substantial cranial kinesis to rapidly increase buccal cavity volume, pulling prey into the mouth via suction feeding. Living polypterids are a key lineage for understanding the evolution and biomechanics of suction feeding because of their phylogenetic position and unique morphology. Polypterus bichir have fewer mobile cranial elements compared with teleosts [e.g. immobile (pre)maxillae] but successfully generate suction through dorsal, ventral and lateral oral cavity expansion. However, the relative contributions of these motions to suction feeding success have not been quantified. Additionally, extensive body musculature and lack of opercular jaw opening linkages make P. bichir of interest for examining the role of cranial versus axial muscles in driving mandibular depression. Here, we analyzed the kinematics of buccal expansion during suction feeding in P. bichir using X-ray Reconstruction of Moving Morphology (XROMM) and quantified the contributions of skeletal elements to oral cavity volume expansion and prey capture. Mouth gape peaks early in the strike, followed by maximum cleithral and ceratohyal rotations, and finally by opercular and suspensorial abductions, maintaining the anterior-to-posterior movement of water. Using a new method of quantifying bones' relative contributions to volume change (RCVC), we demonstrate that ceratohyal kinematics are the most significant drivers of oral cavity volume change. All measured cranial bone motions, except abduction of the suspensorium, are correlated with prey motion. Lastly, cleithral retraction is largely concurrent with ceratohyal retraction and jaw depression, while the sternohyoideus maintains constant length, suggesting a central role of the axial muscles, cleithrum and ceratohyal in ventral expansion.

Loss of oral sensation impairs feeding performance and consistency of tongue-jaw coordination.

BACKGROUND: Individuals with impaired oral sensation report difficulty chewing, but little is known about the underlying changes to tongue and jaw kinematics. Methodological challenges impede the measurement of 3D tongue movement and its relationship to the gape cycle. OBJECTIVE: The aim of this study was to quantify the impact of loss of oral somatosensation on feeding performance, 3D tongue kinematics and tongue-jaw coordination. METHODOLOGY: XROMM (X-ray Reconstruction of Moving Morphology) was used to quantify 3D tongue and jaw kinematics during feeding in three rhesus macaques (Macaca mulatta) before and after an oral tactile nerve block. Feeding performance was measured using feeding sequence duration, number of manipulation cycles and swallow frequency. Coordination was measured using event- and correlation-based metrics of jaw pitch, anterior tongue length, width and roll. RESULTS: In the absence of tactile sensation to the tongue and other oral structures, feeding performance decreased, and the fast open phase of the gape cycle became significantly longer, relative to the other phases (p < .05). The tongue made similar shapes in both the control and nerve block conditions, but the pattern of tongue-jaw coordination became significantly more variable after the block (p < .05). CONCLUSION: Disruption of oral somatosensation impacts feeding performance by introducing variability into the typically tight pattern of tongue-jaw coordination.

Comparative cranial biomechanics in two lizard species: impact of variation in cranial design.

Cranial morphology in lepidosaurs is highly disparate and characterised by the frequent loss or reduction of bony elements. In varanids and geckos, the loss of the postorbital bar is associated with changes in skull shape, but the mechanical principles underlying this variation remain poorly understood. Here, we sought to determine how the overall cranial architecture and the presence of the postorbital bar relate to the loading and deformation of the cranial bones during biting in lepidosaurs. Using computer-based simulation techniques, we compared cranial biomechanics in the varanid Varanus niloticus and the teiid Salvator merianae, two large, active foragers. The overall strain magnitude and distribution across the cranium were similar in the two species, despite lower strain gradients in V. niloticus In S. merianae, the postorbital bar is important for resistance of the cranium to feeding loads. The postorbital ligament, which in varanids partially replaces the postorbital bar, does not affect bone strain. Our results suggest that the reduction of the postorbital bar impaired neither biting performance nor the structural resistance of the cranium to feeding loads in V. niloticus Differences in bone strain between the two species might reflect demands imposed by feeding and non-feeding functions on cranial shape. Beyond variation in cranial bone strain related to species-specific morphological differences, our results reveal that similar mechanical behaviour is shared by lizards with distinct cranial shapes. Contrary to the situation in mammals, the morphology of the circumorbital region, calvaria and palate appears to be important for withstanding high feeding loads in these lizards.

Twist and chew: three-dimensional tongue kinematics during chewing in macaque primates.

Three-dimensional (3D) tongue movements are central to performance of feeding functions by mammals and other tetrapods, but 3D tongue kinematics during feeding are poorly understood. Tongue kinematics were recorded during grape chewing by macaque primates using biplanar videoradiography. Complex shape changes in the tongue during chewing are dominated by a combination of flexion in the tongue's sagittal planes and roll about its long axis. As hypothesized for humans, in macaques during tongue retraction, the middle (molar region) of the tongue rolls to the chewing (working) side simultaneous with sagittal flexion, while the tongue tip flexes to the other (balancing) side. Twisting and flexion reach their maxima early in the fast close phase of chewing cycles, positioning the food bolus between the approaching teeth prior to the power stroke. Although 3D tongue kinematics undoubtedly vary with food type, the mechanical role of this movement-placing the food bolus on the post-canine teeth for breakdown-is likely to be a powerful constraint on tongue kinematics during this phase of the chewing cycle. The muscular drivers of these movements are likely to include a combination of intrinsic and extrinsic tongue muscles.

Grit your teeth and chew your food: Implications of food material properties and abrasives for rates of dental microwear formation in laboratory Sapajus apella (Primates).

Dental microwear analysis has been employed in studies of a wide range of modern and fossil animals, yielding insights into the biology/ecology of those taxa. Some researchers have suggested that dental microwear patterns ultimately relate back to the material properties of the foods being consumed, whereas others have suggested that, because exogenous grit is harder than organic materials in food, grit should have an overwhelming impact on dental microwear patterns. To shed light on this issue, laboratory-based feeding experiments were conducted on tufted capuchin monkeys [Sapajus apella] with dental impressions taken before and after consumption of different artificial foods. The foods were (1) brittle custom-made biscuits laced with either of two differently-sized aluminum silicate abrasives, and (2) ductile custom-made "gummies" laced with either of the two same abrasives. In both cases, animals were allowed to feed on the foods for 36 hours before follow-up dental impressions were taken. Resultant casts were analyzed using a scanning electron microscope. We asked five questions: (1) would the animals consume different amounts of each food item, (2) what types of dental microwear would be formed, (3) would rates of dental microwear differ between the consumption of biscuits (i.e., brittle) versus gummies (i.e., ductile), (4) would rates of dental microwear differ between foods including larger- versus smaller-grained abrasives, and (5) would rates of dental microwear differ between molar shearing and crushing facets in the animals in these experiments? Results indicated that (1) fewer biscuits were consumed when laced with larger-grained abrasives (as opposed to smaller-grained abrasives), but no such difference was observed in the consumption of gummies, (2) in all cases, a variety of dental microwear features was formed, (3) rates of dental microwear were higher when biscuits versus gummies were consumed, (4) biscuits laced with larger-grained abrasives caused a higher percentage of new features per item consumed, and (5) the only difference between facets occurred with the processing of biscuits, where crushing facets showed a faster rate of wear than shearing facets. These findings suggest that the impact of exogenous grit on dental microwear is the result of a dynamic, complex interaction between (at the very least) grit size, food material properties, and time spent feeding - which is further evidence of the multifactorial nature of dental microwear formation.

Differences in muscle mechanics underlie divergent optimality criteria between feeding and locomotor systems.

Tetrapod musculoskeletal diversity is usually studied separately in feeding and locomotor systems. However, direct comparisons between these systems promise important insight into how natural selection deploys the same basic musculoskeletal toolkit-connective tissues, bones, nerves, and skeletal muscle-to meet the differing performance criteria of feeding and locomotion. Recent studies using this approach have proposed that the feeding system is optimized for precise application of high forces and the locomotor system is optimized for wide and rapid joint excursions for minimal energetic expenditure. If this hypothesis is correct, then it stands to reason that other anatomical and biomechanical variables within the feeding and locomotor systems should reflect these diverging functions. To test this hypothesis, we compared muscle moment arm lengths, mechanical advantages, and force vector orientations of two jaw elevator muscles (m. temporalis and m. superficial masseter), an elbow flexor (m. brachialis) and extensor (m. triceps- lateral head), and a knee flexor (m. biceps femoris-short head) and extensor (m. vastus lateralis) across 18 species of primates. Our results show that muscles of the feeding system are more orthogonally oriented relative to the resistance arm (mandible) and operate at relatively large moment arms and mechanical advantages. Moreover, these variables show relatively little change across the range of jaw excursion. In contrast, the representative muscles of the locomotor system have much smaller mechanical advantages and, depending on joint position, smaller muscle moment arm lengths and almost parallel orientations relative to the resistance arm. These patterns are consistent regardless of phylogeny, body mass, locomotor mode, and feeding specialization. We argue that these findings reflect fundamental functional dichotomies between tetrapod locomotor and feeding systems. By organizing muscles in a manner such that moment arms and mechanical advantage are relatively small, the locomotor system can produce broad joint excursions and high angular velocities with only small muscular contraction. As such, the anatomical organization of muscles within the limbs allows striding animals to move relatively rapidly and with minimal energetic expenditure. In contrast, the anatomical configuration of muscles in the feeding system, at least m. superficial masseter and m. temporalis, favors their force-producing capacity at the expense of excursion and velocity.

Jaw kinematics and mandibular morphology in humans.

Understanding the influence of feeding behavior on mandibular morphology is necessary for interpreting dietary change in fossil hominins. However, mandibular morphology is also likely to have an effect on feeding behavior, including jaw kinematics. Here we examine the relationships between mandibular morphology and jaw kinematics in humans using landmark-based morphometrics to quantify jaw movement. Three-dimensional movements of reflective markers coupled to the mandible and cranium were used to capture jaw movements while subjects chewed cubes of raw and cooked sweet potato. Geometric morphometric methods were adapted to quantify and analyze gape cycle motion paths. Gape cycles varied significantly across chewing sequences and between raw and cooked sweet potato. Variation in gape cycle size and shape is related to the width (intergonial distance) and length of the mandible. These results underline the fact that jaw kinematic variation within and between taxa is related to and may be influenced by mandibular morphology. Future studies examining kinematic variation should assess the influence of morphological differences on movement.

Biomechanics of the mandible of Macaca mulatta during the power stroke of mastication: Loading, deformation, and strain regimes and the impact of food type.

Mandible morphology has yet to yield definitive information on primate diet, probably because of poor understanding of mandibular loading and strain regimes, and overreliance on simple beam models of mandibular mechanics. We used a finite element model of a macaque mandible to test hypotheses about mandibular loading and strain regimes and relate variation in muscle activity during chewing on different foods to variation in strain regimes. The balancing-side corpus is loaded primarily by sagittal shear forces and sagittal bending moments. On the working side, sagittal bending moments, anteroposterior twisting moments, and lateral transverse bending moments all reach similar maxima below the bite point; sagittal shear is the dominant loading regime behind the bite point; and the corpus is twisted such that the mandibular base is inverted. In the symphyseal region, the predominant loading regimes are lateral transverse bending and negative twisting about a mediolateral axis. Compared with grape and dried fruit chewing, nut chewing is associated with larger sagittal and transverse bending moments acting on balancing- and working-side mandibles, larger sagittal shear on the working side, and larger twisting moments about vertical and transverse axes in the symphyseal region. Nut chewing is also associated with higher minimum principal strain magnitudes in the balancing-side posterior ramus; higher sagittal shear strain magnitudes in the working-side buccal alveolar process and the balancing-side oblique line, recessus mandibulae, and endocondylar ridge; and higher transverse shear strains in the symphyseal region, the balancing-side medial prominence, and the balancing-side endocondylar ridge. The largest food-related differences in maximum principal and transverse shear strain magnitudes are in the transverse tori and in the balancing-side medial prominence, extramolar sulcus, oblique line, and endocondylar ridge. Food effects on the strain regime are most salient in areas not traditionally investigated, suggesting that studies seeking dietary effects on mandible morphology might be looking in the wrong places.

Integrating XMALab and DeepLabCut for high-throughput XROMM.

Marker tracking is a major bottleneck in studies involving X-ray reconstruction of moving morphology (XROMM). Here, we tested whether DeepLabCut, a new deep learning package built for markerless tracking, could be applied to videoradiographic data to improve data processing throughput. Our novel workflow integrates XMALab, the existing XROMM marker tracking software, and DeepLabCut while retaining each program's utility. XMALab is used for generating training datasets, error correction and 3D reconstruction, whereas the majority of marker tracking is transferred to DeepLabCut for automatic batch processing. In the two case studies that involved an in vivo behavior, our workflow achieved a 6 to 13-fold increase in data throughput. In the third case study, which involved an acyclic, post-mortem manipulation, DeepLabCut struggled to generalize to the range of novel poses and did not surpass the throughput of XMALab alone. Deployed in the proper context, this new workflow facilitates large scale XROMM studies that were previously precluded by software constraints.

Variation in limb loading magnitude and timing in tetrapods.

Comparative analyses of locomotion in tetrapods reveal two patterns of stride cycle variability. Tachymetabolic tetrapods (birds and mammals) have lower inter-cycle variation in stride duration than bradymetabolic tetrapods (amphibians, lizards, turtles and crocodilians). This pattern has been linked to the fact that birds and mammals share enlarged cerebella, relatively enlarged and heavily myelinated Ia afferents, and γ-motoneurons to their muscle spindles. Both tachymetabolic tetrapod lineages also possess an encapsulated Golgi tendon morphology, thought to provide more spatially precise information on muscle tension. The functional consequence of this derived Golgi tendon morphology has never been tested. We hypothesized that one advantage of precise information on muscle tension would be lower and more predictable limb bone stresses, achieved in tachymetabolic tetrapods by having less variable substrate reaction forces than bradymetabolic tetrapods. To test this hypothesis, we analyzed hindlimb substrate reaction forces during locomotion of 55 tetrapod species in a phylogenetic comparative framework. Variation in species means of limb loading magnitude and timing confirm that, for most of the variables analyzed, variance in hindlimb loading and timing is significantly lower in species with encapsulated versus unencapsulated Golgi tendon organs. These findings suggest that maintaining predictable limb loading provides a selective advantage for birds and mammals by allowing energy savings during locomotion, lower limb bone safety factors and quicker recovery from perturbations. The importance of variation in other biomechanical variables in explaining these patterns, such as posture, effective mechanical advantage and center-of-mass mechanics, remains to be clarified.

Jaw Elevator Muscle Coordination during Rhythmic Mastication in Primates: Are Triplets Units of Motor Control?

The activity of mammal jaw elevator muscles during chewing has often been described using the concept of the triplet motor pattern, in which triplet I (balancing side superficial masseter and medial pterygoid; working side posterior temporalis) is consistently activated before triplet II (working side superficial masseter and medial pterygoid; balancing side posterior temporalis), and each triplet of muscles is recruited and modulated as a unit. Here, new measures of unison, synchrony, and coordination are used to determine whether in 5 primate species (Propithecus verreauxi, Eulemur fulvus, Papio anubis, Macaca fuscata,and Pan troglodytes)muscles in the same triplet are active more in unison, are more synchronized, and are more highly coordinated than muscles in different triplets. Results show that triplet I muscle pairs are active more in unison than other muscle pairs in Eulemur, Macaca, and Papio,buttriplet muscle pairs are mostly not more tightly synchronized than non-triplet pairs. Triplet muscles are more coordinated during triplet pattern cycles than non-triplet cycles, while non-triplet muscle pairs are more coordinated during non-triplet cycles than triplet cycles. These results suggest that the central nervous system alters patterns of coordination between cycles, recruiting triplet muscles as a coordinated unit during triplet cycles but employing a different pattern of muscle coordination during non-triplet cycles. The triplet motor pattern may simplify modulation of rhythmic mastication by being one possible unit of coordination that can be recruited on a cycle-to-cycle basis.

The dental microwear of hard-object feeding in laboratory Sapajus apella and its implications for dental microwear formation.

OBJECTIVES: This study seeks to determine if (a) consumption of hard food items or a mixture of food items leads to the formation of premolar or molar microwear in laboratory capuchin monkeys (Sapajus apella) in one feeding session and (b) rates of microwear formation are associated with the number of food items consumed. MATERIALS AND METHODS: Five adult male capuchins were used in two experiments, one where they were fed unshelled Brazil nuts, and the other where they were fed a mixture of food items. Dental impressions were taken before and after each feeding session. Epoxy casts made from those impressions then were used in SEM analyses of rates of microwear formation. Upper and lower premolars and molars were analyzed. Qualitative comparisons were made and Spearman's rank-order correlations used to examine the relationship between rates of microwear formation and number of Brazil nuts consumed. RESULTS: Premolars and molars generally showed new microwear in the form of pits and scratches. However, the incidence of those features was low (0-6%). Rates of microwear formation were highest during the consumption of Brazil nuts. DISCUSSION: Variations in the rate of microwear formation on the premolars likely reflected patterns of ingestion whereas consistency in the rate of microwear on the molars likely reflected patterns of chewing. While dental microwear formation seemed to be correlated with the number of hard objects consumed, rates did differ between individuals. Differences in results between the two experiments demonstrate some of the limitations in our knowledge of dental microwear formation.

Joint angular excursions during cyclical behaviors differ between tetrapod feeding and locomotor systems.

Tetrapod musculoskeletal diversity is usually studied separately in feeding and locomotor systems. However, comparisons between these systems promise important insight into how natural selection deploys the same basic musculoskeletal toolkit - connective tissues, bones, nerves and skeletal muscle - to meet the differing performance criteria of feeding and locomotion. In this study, we compare average joint angular excursions during cyclic behaviors - chewing, walking and running - in a phylogenetic context to explore differences in the optimality criteria of these two systems. Across 111 tetrapod species, average limb-joint angular excursions during cyclic locomotion are greater and more evolutionarily labile than those of the jaw joint during cyclic chewing. We argue that these findings reflect fundamental functional dichotomies between tetrapod locomotor and feeding systems. Tetrapod chewing systems are optimized for precise application of force over a narrower, more controlled and predictable range of displacements, the principal aim being to fracture the substrate, the size and mechanical properties of which are controlled at ingestion and further reduced and homogenized, respectively, by the chewing process. In contrast, tetrapod limbed locomotor systems are optimized for fast and energetically efficient application of force over a wider and less predictable range of displacements, the principal aim being to move the organism at varying speeds relative to a substrate whose geometry and mechanical properties need not become more homogeneous as locomotion proceeds. Hence, the evolution of tetrapod locomotor systems has been accompanied by an increasing diversity of limb-joint excursions, as tetrapods have expanded across a range of locomotor substrates and environments.

Taking a big bite: Working together to better understand the evolution of feeding in primates.

The study of adaptation requires the integration of an array of different types of data. A single individual can find such integration daunting, if not impossible. In an effort to clarify the role of diet in the evolution of the primate craniofacial and dental apparatus, we assembled a team of researchers that have various types and degrees of expertise. This interaction has provided a range of insights for all contributors, and this has helped to refine questions, clarify the possibilities and limitations that laboratory and field settings offer, and further explore the ways in which laboratory and field data can be suitably integrated. A complete and accurate picture of dietary adaptation cannot be gained in isolation. Collaboration provides the bridge to a more holistic view of primate biology and evolution.