The Form-Function Complex of the Primate Masticatory Apparatus

Craniofacial morphology varies considerably between primate species with many aspects of structural variation occurring within the masticatory apparatus. These variations in masticatory fonn are frequently attributed to differences in diet however elucidating the relationship between masticatory form and function is complex. This study combines the techniques of shape analysis in a comparative study of the primate masticatory apparatus with three-dimensional biomechanical modelling of primate jaw mechanics. The relationship between masticatory form and function is investigated within a subfamily ofprimates, the Cercopithecinae. Species within this group display a range of masticatory forms, inhabit a wide range of environments, have varying diets and share close phylogenetic relationships making them an ideal group to investigate the relationship between structure and function. Using the techniques of geometric morphometrics a shape analysis was conducted in which variations within the masticatory forms of the Cercopithecinae were quantified. Functional predictions of observed shape differences were made and considered in light of known environmental and ecological factors with particular emphasis on dietary specialisations and fall back foods. The results of the shape analysis indicate that differences in body size, strata utilisation, and diet appear to play a major role in structuring adult morphological diversity within and among the Cercopithecinae. Many of the shape differences associated with an increased body size appear to reflect the selective pressures of increased predation risk, including increases in the length ofthe jaw and canine teeth. These morphological traits are associated with the production of a large gape and are hypothesised to be functional adaptations to the use of canines as weapons. Other major shape differences apparent between the Cercopithecinae were those associated with the proportion of leaves and fruit in the diet. The highly frugivorous Mandril/us species possessed features associated with increasing gape and retractile motions of the mandible ideal for incision of large fruits. By contrast the dietary specialist Theropithecus gelada possessed features associated with increasing the efficiency of food breakdown and muscle force production, necessary to process their more abrasive foodstuff (i.e. grasses)

Masticatory biomechanics in the rabbit : a multi-body dynamics analysis

Acknowledgement We thank Sue Taft (University of Hull) for the µCT-scanning of the rabbit specimen used in this study. We also thank Raphaël Cornette, Jacques Bonnin, Laurent Dufresne, and l'Amicale des Chasseurs Trappistes (ACT) for providing permission and helping us capture the rabbits used for the in vivo bite force measurements at la Réserve Naturelle Nationale de St Quentin en Yvelines, France.Peer reviewedPublisher PD

Finite element analysis of the cranium : Validity, sensitivity and future directions

Finite element analysis (FEA) is increasingly applied in skeletal biomechanical research in general, and in fossil studies in particular. Underlying such studies is the principle that FEA provides results that approximate reality. This paper provides further understanding of the reliability of FEA by presenting a validation study in which the deformations experienced by a real cadaveric human cranium are compared to those of an FE model of that cranium under equivalent simulated loading. Furthermore, model sensitivity to simplifications in segmentation and material properties is also assessed. Our results show that absolute deformations are not accurately predicted, but the distribution of the regions of relatively high and low strains, and so the modes of global deformation, are reasonably approximated

Geometric morphometrics and finite elements analysis : Assessing the functional implications of differences in craniofacial form in the hominin fossil record

The study of morphological variation in the hominin fossil record has been transformed in recent years by the advent of high resolution 3D imaging combined with improved geometric morphometric (GM) toolkits. In parallel, increasing numbers of studies have applied finite elements analysis (FEA) to the study of skeletal mechanics in fossil and extant hominoid material. While FEA studies of fossils are becoming ever more popular they are constrained by the difficulties of reconstruction and by the uncertainty that inevitably attaches to the estimation of forces and material properties. Adding to these modelling difficulties it is still unclear how FEA analyses should best deal with species variation.Comparative studies of skeletal form and function can be further advanced by applying tools from the GM toolkit to the inputs and outputs of FEA studies. First they facilitate virtual reconstruction of damaged material and can be used to rapidly create 3D models of skeletal structures. Second, GM methods allow variation to be accounted for in FEA by warping models to represent mean and extreme forms of interest. Third, GM methods can be applied to compare FEA outputs - the ways in which skeletal elements deform when loaded. Model comparisons are hampered by differences in material properties, forces and size among models but how deformations from FEA are impacted by these parameters is increasingly well understood, allowing them to be taken into account in comparing FEA outputs.In this paper we review recent advances in the application of GM in relation to FEA studies of craniofacial form in hominins, providing examples from our recent work and a critical appraisal of the state of the art

The biting performance of Homo sapiens and Homo heidelbergensis

Modern humans have smaller faces relative to Middle and Late Pleistocene members of the genus Homo. While facial reduction and differences in shape have been shown to increase biting efficiency in Homo sapiens relative to these hominins, facial size reduction has also been said to decrease our ability to resist masticatory loads. This study compares crania of Homo heidelbergensis and H. sapiens with respect to mechanical advantages of masticatory muscles, force production efficiency, strains experienced by the cranium and modes of deformation during simulated biting. Analyses utilize X-ray computed tomography (CT) scan-based 3D models of a recent modern human and two H. heidelbergensis. While having muscles of similar cross-sectional area to H. heidelbergensis, our results confirm that the modern human masticatory system is more efficient at converting muscle forces into bite forces. Thus, it can produce higher bite forces than Broken Hill for equal muscle input forces. This difference is the result of alterations in relative in and out-lever arm lengths associated with well-known differences in midfacial prognathism. Apparently at odds with this increased efficiency is the finding that the modern human cranium deforms more, resulting in greater strain magnitudes than Broken Hill when biting at the equivalent tooth. Hence, the facial reduction that characterizes modern humans may not have evolved as a result of selection for force production efficiency. These findings provide further evidence for a degree of uncoupling between form and function in the masticatory system of modern humans. This may reflect the impact of food preparation technologies. These data also support previous suggestions that differences in bite force production efficiency can be considered a spandrel, primarily driven by the midfacial reduction in H. sapiens that occurred for other reasons. Midfacial reduction plausibly resulted in a number of other significant changes in morphology, such as the development of a chin, which has itself been the subject of debate as to whether or not it represents a mechanical adaptation or a spandrel

The Long-Baseline Neutrino Experiment: Exploring Fundamental Symmetries of the Universe

The preponderance of matter over antimatter in the early Universe, the dynamics of the supernova bursts that produced the heavy elements necessary for life and whether protons eventually decay --- these mysteries at the forefront of particle physics and astrophysics are key to understanding the early evolution of our Universe, its current state and its eventual fate. The Long-Baseline Neutrino Experiment (LBNE) represents an extensively developed plan for a world-class experiment dedicated to addressing these questions. LBNE is conceived around three central components: (1) a new, high-intensity neutrino source generated from a megawatt-class proton accelerator at Fermi National Accelerator Laboratory, (2) a near neutrino detector just downstream of the source, and (3) a massive liquid argon time-projection chamber deployed as a far detector deep underground at the Sanford Underground Research Facility. This facility, located at the site of the former Homestake Mine in Lead, South Dakota, is approximately 1,300 km from the neutrino source at Fermilab -- a distance (baseline) that delivers optimal sensitivity to neutrino charge-parity symmetry violation and mass ordering effects. This ambitious yet cost-effective design incorporates scalability and flexibility and can accommodate a variety of upgrades and contributions. With its exceptional combination of experimental configuration, technical capabilities, and potential for transformative discoveries, LBNE promises to be a vital facility for the field of particle physics worldwide, providing physicists from around the globe with opportunities to collaborate in a twenty to thirty year program of exciting science. In this document we provide a comprehensive overview of LBNE's scientific objectives, its place in the landscape of neutrino physics worldwide, the technologies it will incorporate and the capabilities it will possess.Comment: Major update of previous version. This is the reference document for LBNE science program and current status. Chapters 1, 3, and 9 provide a comprehensive overview of LBNE's scientific objectives, its place in the landscape of neutrino physics worldwide, the technologies it will incorporate and the capabilities it will possess. 288 pages, 116 figure

Effectiveness of a national quality improvement programme to improve survival after emergency abdominal surgery (EPOCH): a stepped-wedge cluster-randomised trial

BACKGROUND: Emergency abdominal surgery is associated with poor patient outcomes. We studied the effectiveness of a national quality improvement (QI) programme to implement a care pathway to improve survival for these patients. METHODS: We did a stepped-wedge cluster-randomised trial of patients aged 40 years or older undergoing emergency open major abdominal surgery. Eligible UK National Health Service (NHS) hospitals (those that had an emergency general surgical service, a substantial volume of emergency abdominal surgery cases, and contributed data to the National Emergency Laparotomy Audit) were organised into 15 geographical clusters and commenced the QI programme in a random order, based on a computer-generated random sequence, over an 85-week period with one geographical cluster commencing the intervention every 5 weeks from the second to the 16th time period. Patients were masked to the study group, but it was not possible to mask hospital staff or investigators. The primary outcome measure was mortality within 90 days of surgery. Analyses were done on an intention-to-treat basis. This study is registered with the ISRCTN registry, number ISRCTN80682973. FINDINGS: Treatment took place between March 3, 2014, and Oct 19, 2015. 22 754 patients were assessed for elegibility. Of 15 873 eligible patients from 93 NHS hospitals, primary outcome data were analysed for 8482 patients in the usual care group and 7374 in the QI group. Eight patients in the usual care group and nine patients in the QI group were not included in the analysis because of missing primary outcome data. The primary outcome of 90-day mortality occurred in 1210 (16%) patients in the QI group compared with 1393 (16%) patients in the usual care group (HR 1·11, 0·96-1·28). INTERPRETATION: No survival benefit was observed from this QI programme to implement a care pathway for patients undergoing emergency abdominal surgery. Future QI programmes should ensure that teams have both the time and resources needed to improve patient care. FUNDING: National Institute for Health Research Health Services and Delivery Research Programme

Developing a musculoskeletal model of the primate skull: Predicting muscle activations, bite force, and joint reaction forces using multibody dynamics analysis and advanced optimisation methods

An accurate, dynamic, functional model of the skull that can be used to predict muscle forces, bite forces, and joint reaction forces would have many uses across a broad range of disciplines. One major issue however with musculoskeletal analyses is that of muscle activation pattern indeterminacy. A very large number of possible muscle force combinations will satisfy a particular functional task. This makes predicting physiological muscle recruitment patterns difficult. Here we describe in detail the process of development of a complex multibody computer model of a primate skull (Macaca fascicularis), that aims to predict muscle recruitment patterns during biting. Using optimisation criteria based on minimisation of muscle stress we predict working to balancing side muscle force ratios, peak bite forces, and joint reaction forces during unilateral biting. Validation of such models is problematic; however we have shown comparable working to balancing muscle activity and TMJ reaction ratios during biting to those observed in vivo and that peak predicted bite forces compare well to published experimental data. To our knowledge the complexity of the musculoskeletal model is greater than any previously reported for a primate. This complexity, when compared to more simple representations provides more nuanced insights into the functioning of masticatory muscles. Thus, we have shown muscle activity to vary throughout individual muscle groups, which enables them to function optimally during specific masticatory tasks. This model will be utilised in future studies into the functioning of the masticatory apparatus

The application of muscle wrapping to voxel-based finite element models of skeletal structures

Finite elements analysis (FEA) is now used routinely to interpret skeletal form in terms of function in both medical and biological applications. To produce accurate predictions from FEA models, it is essential that the loading due to muscle action is applied in a physiologically reasonable manner. However, it is common for muscle forces to be represented as simple force vectors applied at a few nodes on the model's surface. It is certainly rare for any wrapping of the muscles to be considered, and yet wrapping not only alters the directions of muscle forces but also applies an additional compressive load from the muscle belly directly to the underlying bone surface. This paper presents a method of applying muscle wrapping to high-resolution voxel-based finite element (FE) models. Such voxel-based models have a number of advantages over standard (geometry-based) FE models, but the increased resolution with which the load can be distributed over a model's surface is particularly advantageous, reflecting more closely how muscle fibre attachments are distributed. In this paper, the development, application and validation of a muscle wrapping method is illustrated using a simple cylinder. The algorithm: (1) calculates the shortest path over the surface of a bone given the points of origin and ultimate attachment of the muscle fibres; (2) fits a Non-Uniform Rational B-Spline (NURBS) curve from the shortest path and calculates its tangent, normal vectors and curvatures so that normal and tangential components of the muscle force can be calculated and applied along the fibre; and (3) automatically distributes the loads between adjacent fibres to cover the bone surface with a fully distributed muscle force, as is observed in vivo. Finally, we present a practical application of this approach to the wrapping of the temporalis muscle around the cranium of a macaque skull

Combining geometric morphometrics and functional simulation: an emerging toolkit for virtual functional analyses

The development of virtual methods for anatomical reconstruction and functional simulation of skeletal structures offers great promise in evolutionary and ontogenetic investigations of form-function relationships. Key developments reviewed here include geometric morphometric methods for the analysis and visualization of variations in form (size and shape), finite element methods for the prediction of mechanical performance of skeletal structures under load and multibody dynamics methods for the simulation and prediction of musculoskeletal function. These techniques are all used in studies of form and function in biology, but only recently have they been combined in novel ways to facilitate biomechanical modelling that takes account of variations in form, can statistically compare performance, and relate performance to form and its covariates. Here we provide several examples that illustrate how these approaches can be combined and we highlight areas that require further investigation and development before we can claim a mature theory and toolkit for a statistical biomechanical framework that unites these methods