Development of a test bench for biomechanical simulation-a preliminary study of mandibular forces.

Purpose: The aim of this study is to develop a test bench, which integrates different complexity levels and enables in that way a flexible and dynamic testing for mid and long term intervals as well as testing of maximum loads till implant failure of different osteosynthesis systems on the mandible. Material and Methods: For this purpose, an analysis of the state of the art regarding existing test benches was combined with interviews of clinical experts to acquire a list of requirements. Based on these requirements a design for a modular test bench was developed. During the implementation of the test stand, functional tests were continuously carried out and improvements made. Depending on the level of complexity, the test bench can be used either as an incorporated variant or as a standalone solution. In order to verify the performance and the degree of fulfilment of the requirements of these two variants of the test bench, preliminary studies were carried out for all levels of complexity. In these preliminary studies, commercially available osteosynthesis and reconstruction plates were investigated for their biomechanical behaviour and compared with data from the literature. Results: In total, fourteen test runs were performed for the different levels of complexity. Firstly, five test runs were executed to test the simplified load scenario in the incorporated variant of the test bench. High forces could be transmitted without failure of the miniplates. Secondly a quasi-static test scenario was examined using the incorporated variant with simplified load insertion. Five experiments with a number of cycles between 40,896 and 100,000 cycles were carried out. In one case the quasi-static testing resulted in a fracture of the tested reconstruction plate with a failure mode similar to the clinical observations of failure. The last four test runs were carried out using the standalone variant of the test bench simulating complex load patterns via the insertion of forces through imitated muscles. During the test runs joint forces were measured and the amplitude and vector of the resulting joint forces were calculated for both temporomandibular joints. Differences in the force transmission depending on the implant system in comparison to the zero sample could be observed. Conclusion: The presented modular test bench showed to be applicable for examination of the biomechanical behavior of the mandible. It is characterized by the adjustability of the complexity regarding the load patterns and enables the subsequent integration of further sensor technologies. Follow-up studies are necessary to further qualify and optimize the test bench.

Determination of muscle strength and function in plesiosaur limbs: finite element structural analyses of Cryptoclidus eurymerus humerus and femur.

Background: The Plesiosauria (Sauropterygia) are secondary marine diapsids. They are the only tetrapods to have evolved hydrofoil fore- and hindflippers. Once this specialization of locomotion had evolved, it remained essentially unchanged for 135 Ma. It is still controversial whether plesiosaurs flew underwater, rowed, or used a mixture of the two modes of locomotion. The long bones of Tetrapoda are functionally loaded by torsion, bending, compression, and tension during locomotion. Superposition of load cases shows that the bones are loaded mainly by compressive stresses. Therefore, it is possible to use finite element structure analysis (FESA) as a test environment for loading hypotheses. These include muscle reconstructions and muscle lines of action (LOA) when the goal is to obtain a homogeneous compressive stress distribution and to minimize bending in the model. Myological reconstruction revealed a muscle-powered flipper twisting mechanism. The flippers of plesiosaurs were twisted along the flipper length axis by extensors and flexors that originated from the humerus and femur as well as further distal locations. Methods: To investigate locomotion in plesiosaurs, the humerus and femur of a mounted skeleton of Cryptoclidus eurymerus (Middle Jurassic Oxford Clay Formation from Britain) were analyzed using FE methods based on the concept of optimization of loading by compression. After limb muscle reconstructions including the flipper twisting muscles, LOA were derived for all humerus and femur muscles of Cryptoclidus by stretching cords along casts of the fore- and hindflippers of the mounted skeleton. LOA and muscle attachments were added to meshed volumetric models of the humerus and femur derived from micro-CT scans. Muscle forces were approximated by stochastic iteration and the compressive stress distribution for the two load cases, "downstroke" and "upstroke", for each bone were calculated by aiming at a homogeneous compressive stress distribution. Results: Humeral and femoral depressors and retractors, which drive underwater flight rather than rowing, were found to exert higher muscle forces than the elevators and protractors. Furthermore, extensors and flexors exert high muscle forces compared to Cheloniidae. This confirms a convergently evolved myological mechanism of flipper twisting in plesiosaurs and complements hydrodynamic studies that showed flipper twisting is critical for efficient plesiosaur underwater flight.

Three-dimensional finite element analysis of the dural folds and the human skull under head acceleration.

Bone and collagen fiber architecture adapt to external mechanical loads. In humans, due to the low insertion of the temporal muscle, mastication does not lead to a physiological loading of the calvaria. Forces applied to the skull by the dural folds can lead to compressive stresses in the calvaria. To investigate the relationship between mechanical loads and form in the skull and its membranes, in a finite element three-dimensional model of the human skull, loads due to head acceleration in daily activities are applied to the falx cerebri and the tentorium cerebelli. The dural folds are modeled as membranes. The stress paths in the dural folds correlate with anatomical fiber direction. Head accelerations of 9 g lead to compressive stress in the calvaria. Finite element analysis of the falx cerebri and the tentorium cerebelli can be used to study the influence of mechanical stresses on the ossification of the dural folds and their impact on calvarial growth. This study presents an example of functional loading of bone by fibrous membranes and describes a possible mechanism by which Wolff's law works on the bone of the calvaria creating evolutionarily beneficial lightweight constructions.

Foreflipper and hindflipper muscle reconstructions of Cryptoclidus eurymerus in comparison to functional analogues: introduction of a myological mechanism for flipper twisting.

BACKGROUND: Plesiosaurs, diapsid crown-group Sauropterygia, inhabited the oceans from the Late Triassic to the Late Cretaceous. Their most exceptional characteristic are four hydrofoil-like flippers. The question whether plesiosaurs employed their four flippers in underwater flight, rowing flight, or rowing has not been settled yet. Plesiosaur locomotory muscles have been reconstructed in the past, but neither the pelvic muscles nor the distal fore- and hindflipper musculature have been reconstructed entirely. METHODS: All plesiosaur locomotory muscles were reconstructed in order to find out whether it is possible to identify muscles that are necessary for underwater flight including those that enable flipper rotation and twisting. Flipper twisting has been proven by hydrodynamic studies to be necessary for efficient underwater flight. So, Cryptoclidus eurymerus fore- and hindflipper muscles and ligaments were reconstructed using the extant phylogenetic bracket (Testudines, Crocodylia, and Lepidosauria) and correlated with osteological features and checked for their functionality. Muscle functions were geometrically derived in relation to the glenoid and acetabulum position. Additionally, myology of functionally analogous Chelonioidea, Spheniscidae, Otariinae, and Cetacea is used to extract general myological adaptations of secondary aquatic tetrapods to inform the phylogenetically inferred muscle reconstructions. RESULTS: A total of 52 plesiosaur fore- and hindflipper muscles were reconstructed. Amongst these are flipper depressors, elevators, retractors, protractors, and rotators. These muscles enable a fore- and hindflipper downstroke and upstroke, the two sequences that represent an underwater flight flipper beat cycle. Additionally, other muscles were capable of twisting fore- and hindflippers along their length axis during down- and upstroke accordingly. A combination of these muscles that actively aid in flipper twisting and intermetacarpal/intermetatarsal and metacarpodigital/metatarsodigital ligament systems, that passively engage the successive digits, could have accomplished fore-and hindflipper length axis twisting in plesiosaurs that is essential for underwater flight. Furthermore, five muscles that could possibly actively adjust the flipper profiles for efficient underwater flight were found, too.

Humerus osteology, myology, and finite element structure analysis of Cheloniidae.

Adaptation of osteology and myology lead to the formation of hydrofoil foreflippers in Cheloniidae (all recent sea turtles except Dermochelys coriacea) which are used mainly for underwater flight. Recent research shows the biomechanical advantages of a complex system of agonistic and antagonistic tension chords that reduce bending stress in bones. Finite element structure analysis (FESA) of a cheloniid humerus is used to provide a better understanding of morphology and microanatomy and to link these with the main flipper function, underwater flight. Dissection of a Caretta caretta gave insights into lines of action, that is, the course that a muscle takes between its origin and insertion, of foreflipper musculature. Lines of action were determined by spanning physical threads on a skeleton of Chelonia mydas. The right humerus of this skeleton was micro-CT scanned. Based on the scans, a finite element (FE) model was built and muscle force vectors were entered. Muscle forces were iteratively approximated until a uniform compressive stress distribution was attained. Two load cases, downstroke and upstroke, were computed. We found that muscle wrappings (m. coracobrachialis magnus and brevis, several extensors, humeral head of m. triceps) are crucial in addition to axial loading to obtain homogenous compressive loading in all bone cross-sections. Detailed knowledge on muscle disposition leads to compressive stress distribution in the FE model which corresponds with the bone microstructure. The FE analysis of the cheloniid humerus shows that bone may be loaded mainly by compression if the bending moments are minimized.

Biomechanical Study of the Development of Long Bones: Finite Element Structure Synthesis of the Human Second Proximal Phalanx Under Growth Conditions.

Torsional loads are a possible mechanical explanation for the architecture of long bone. Finite element structure synthesis (FESS) has previously successfully been used as a deductive technique using Wolff's Law by applying expected loads to an unspecific homogeneous solid and eliminating stress free parts to verify muscle forces. The extended approach presented in this article includes further mechanobiological rules to model the development from a cartilage model to a finger bone. In contrast to former computational models, simulation of processes leading to both external growth and internal differentiation are included. Combined axial and torsional loads synthesize a complete human secondary proximal phalanx model comparable to form and internal structure to that observed in vivo. While the computational model is very sensitive to initial alterations of loads, changes after growth have a minor effect as observed in animal models. Predictions of cartilage growth and ossification during FESS showed significant similarities to ontogeny indicating the importance of mechanical factors for the morphogenesis of bone during growth. Anat Rec, 302:1389-1398, 2019. © 2018 Wiley Periodicals, Inc.

Finite-Element Syntheses of Callus and Bone Remodeling: Biomechanical Study of Fracture Healing in Long Bones.

Computational simulations of fracture healing are a valuable tool to improve fracture treatment and implants. Several finite-element models have been established to predict callus formation due to mechanobiological rules. This work provides a comprehensive simulation for virtual implantation through the combination of callus simulation and finite-element structural synthesis (FESS) of (re-)modeling during and after healing based on Pauwel's theory of histogenesis and Wolff's law. The simulation is based on a linear elastic material model and includes generation of fracture hematoma and initial mesenchymal stem cell concentration out of an unspecified solid, cell proliferation, migration, and differentiation due to mechanical stimuli and time-dependent axial loading. Three nondisplaced femoral shaft fractures with initial interfragmentary movement of 0.2, 0.6, and 1 mm and one fracture with 4 mm translation are modeled. The predictions of interfragmentary movement during fracture healing, healing success, and healing time agree with observed clinical outcome, animal models, and other numerical models. Initial interfragmentary movement between 0.2 and 1 mm leads to healing success, with the fastest healing occurring at 0.2 mm. The model of the dislocated fractures shows no further bending after remodeling and is loaded with physiological stress of -13 MPa. Ideal load-time graphs may give insight into the bone's ability to withstand loads as healing time progresses, and thus holds potential for applications in rehabilitation planning. Better knowledge of the forces present during fracture healing is needed to deploy simulations for surgical planning and manufacturing of patient individualized implants. Anat Rec, 301:2112-2121, 2018. © 2018 Wiley Periodicals, Inc.

Inclusion of periodontal ligament fibres in mandibular finite element models leads to an increase in alveolar bone strains.

Alveolar bone remodelling is vital for the success of dental implants and orthodontic treatments. However, the underlying biomechanical mechanisms, in particular the function of the periodontal ligament (PDL) in bone loading and remodelling, are not well understood. The PDL is a soft fibrous connective tissue that joins the tooth root to the alveolar bone and plays a critical role in the transmission of loads from the tooth to the surrounding bone. However, due to its complex structure, small size and location within the tooth socket it is difficult to study in vivo. Finite element analysis (FEA) is an ideal tool with which to investigate the role of the PDL, however inclusion of the PDL in FE models is complex and time consuming, therefore consideration must be given to how it is included. The aim of this study was to investigate the effects of including the PDL and its fibrous structure in mandibular finite element models. A high-resolution model of a human molar region was created from micro-computed tomography scans. This is the first time that the fibrous structure of the PDL has been included in a model with realistic tooth and bone geometry. The results show that omission of the PDL creates a more rigid model, reducing the strains observed in the mandibular corpus which are of interest when considering mandibular functional morphology. How the PDL is modelled also affects the strains. The inclusion of PDL fibres alters the strains in the mandibular bone, increasing the strains in the tooth socket compared to PDL modelled without fibres. As strains in the alveolar bone are thought to play a key role in bone remodelling during orthodontic tooth movement, future FE analyses aimed at improving our understanding and management of orthodontic treatment should include the fibrous structure of the PDL.

Biomechanical properties of predator-induced body armour in the freshwater crustacean Daphnia.

The freshwater crustacean Daphnia is known for its ability to develop inducible morphological defences that thwart predators. These defences are developed only in the presence of predators and are realized as morphological shape alterations e.g. 'neckteeth' in D. pulex and 'crests' in D. longicephala. Both are discussed to hamper capture, handling or consumption by interfering with the predator's prey capture devices. Additionally, D. pulex and some other daphniids were found to armour-up and develop structural alterations resulting in increased carapace stiffness. We used scanning transmission electron microscopy (STEM) and confocal laser scanning microscopy (CLSM) to identify predator-induced structural and shape alterations. We found species specific structural changes accompanying the known shape alterations. The cuticle becomes highly laminated (i.e. an increased number of layers) in both species during predator exposure. Using nano- and micro-indentation as well as finite element analysis (FEA) we determined both: the structure's and shape's contribution to the carapace's mechanical resistance. From our results we conclude that only structural alterations are responsible for increased carapace stiffness, whereas shape alterations appear to pose handling difficulties during prey capture. Therefore, these defences act independently at different stages during predation.

Skeletal immaturity, rostral sparing, and disparate hip morphologies as biomechanical causes for Legg-Calvé-Perthes' disease.

Legg-Calvé-Perthes' (Perthes') disease is a developmental disease of the hip joint that may result in numerous short and long term problems. The etiology of the disease remains largely unknown, but the mechanism is believed to be vascular and/or biomechanical in nature. There are several anatomical characteristics that tend to be prevalent in children with Perthes' disease, namely: skeletal immaturity, reduced height, and rostral sparing. We present an overview of the literature, summarizing the current understanding of the pathogenesis, particularly related to how the formation of the vasculature to the femoral epiphysis places children aged 5-8 at a higher risk for Perthes' disease, how skeletal immaturity and rostral sparing could increase the probability of developing Perthes' disease, and how animal models have aided our understanding of the disease. In doing so, we also explore why Perthes' disease is correlated to latitude, with populations at higher latitudes having higher incidence rates than populations closer to the Equator. Finally, we present five hypotheses detailing how Perthes' disease could have a biomechanical cause. Clin. Anat. 29:759-772, 2016. © 2016 Wiley Periodicals, Inc.

Structure and function of the septum nasi and the underlying tension chord in crocodylians.

A long rostrum has distinct advantages for prey capture in an aquatic or semi-aquatic environment but at the same time poses severe problems concerning stability during biting. We here investigate the role of the septum nasi of brevirostrine crocodilians for load-absorption during mastication. Histologically, both the septum nasi and the septum interorbitale consist of hyaline cartilage and therefore mainly resist compression. However, we identified a strand of tissue extending longitudinally below the septum nasi that is characterized by a high content of collagenous and elastic fibers and could therefore resist tensile stresses. This strand of tissue is connected with the m. pterygoideus anterior. Two-dimensional finite element modeling shows that minimization of bending in the crocodilian skull can only be achieved if tensile stresses are counteracted by a strand of tissue. We propose that the newly identified strand of tissue acts as an active tension chord necessary for stabilizing the long rostrum of crocodilians during biting by transforming the high bending stress of the rostrum into moderate compressive stress.

The biomechanical function of periodontal ligament fibres in orthodontic tooth movement.

Orthodontic tooth movement occurs as a result of resorption and formation of the alveolar bone due to an applied load, but the stimulus responsible for triggering orthodontic tooth movement remains the subject of debate. It has been suggested that the periodontal ligament (PDL) plays a key role. However, the mechanical function of the PDL in orthodontic tooth movement is not well understood as most mechanical models of the PDL to date have ignored the fibrous structure of the PDL. In this study we use finite element (FE) analysis to investigate the strains in the alveolar bone due to occlusal and orthodontic loads when PDL is modelled as a fibrous structure as compared to modelling PDL as a layer of solid material. The results show that the tension-only nature of the fibres essentially suspends the tooth in the tooth socket and their inclusion in FE models makes a significant difference to both the magnitude and distribution of strains produced in the surrounding bone. The results indicate that the PDL fibres have a very important role in load transfer between the teeth and alveolar bone and should be considered in FE studies investigating the biomechanics of orthodontic tooth movement.

Development and three-dimensional morphology of the zygomaticotemporal suture in primate skulls.

Cranial sutures are an essential part of the growing skull, allowing bones to increase in size during growth, with their morphology widely believed to be dictated by the forces and displacements that they experience. The zygomaticotemporal suture in primates is located in the relatively weak zygomatic arch, and externally it appears a very simple connection. However, large forces are almost certainly transmitted across this suture, suggesting that it requires some level of stability while also allowing controlled movements under high loading. Here we examine the 2- and 3-dimensional (3D) morphology of the zygomaticotemporal suture in an ontogenetic series of Macaca fascicularis skulls. High resolution microcomputed tomography data sets were examined, and virtual and physical 3D replicas were created to assess both structure and general stability. The zygomaticotemporal suture is much more complex than its external appearance suggests, with interlocking facets between the adjacent zygomatic and temporal bones. It appears as if some movement is permitted across the suture in younger animals, but as they approach adulthood the complexity of the suture's interlocking bone facets reaches a level where these movements become minimal.

Quantitative interpretation of tracks for determination of body mass.

To better understand the biology of extinct animals, experimentation with extant animals and innovative numerical approaches have grown in recent years. This research project uses principles of soil mechanics and a neoichnological field experiment with an African elephant to derive a novel concept for calculating the mass (i.e., the weight) of an animal from its footprints. We used the elephant's footprint geometry (i.e., vertical displacements, diameter) in combination with soil mechanical analyses (i.e., soil classification, soil parameter determination in the laboratory, Finite Element Analysis (FEA) and gait analysis) for the back analysis of the elephant's weight from a single footprint. In doing so we validated the first component of a methodology for calculating the weight of extinct dinosaurs. The field experiment was conducted under known boundary conditions at the Zoological Gardens Wuppertal with a female African elephant. The weight of the elephant was measured and the walking area was prepared with sediment in advance. Then the elephant was walked across the test area, leaving a trackway behind. Footprint geometry was obtained by laser scanning. To estimate the dynamic component involved in footprint formation, the velocity the foot reaches when touching the subsoil was determined by the Digital Image Correlation (DIC) technique. Soil parameters were identified by performing experiments on the soil in the laboratory. FEA was then used for the backcalculation of the elephant's weight. With this study, we demonstrate the adaptability of using footprint geometry in combination with theoretical considerations of loading of the subsoil during a walk and soil mechanical methods for prediction of trackmakers weight.

Boundary conditions for heat transfer and evaporative cooling in the trachea and air sac system of the domestic fowl: a two-dimensional CFD analysis.

Various parts of the respiratory system play an important role in temperature control in birds. We create a simplified computational fluid dynamics (CFD) model of heat exchange in the trachea and air sacs of the domestic fowl (Gallus domesticus) in order to investigate the boundary conditions for the convective and evaporative cooling in these parts of the respiratory system. The model is based upon published values for respiratory times, pressures and volumes and upon anatomical data for this species, and the calculated heat exchange is compared with experimentally determined values for the domestic fowl and a closely related, wild species. In addition, we studied the trachea histologically to estimate the thickness of the heat transfer barrier and determine the structure and function of moisture-producing glands. In the transient CFD simulation, the airflow in the trachea of a 2-dimensional model is evoked by changing the volume of the simplified air sac. The heat exchange between the respiratory system and the environment is simulated for different ambient temperatures and humidities, and using two different models of evaporation: constant water vapour concentration model and the droplet injection model. According to the histological results, small mucous glands are numerous but discrete serous glands are lacking on the tracheal surface. The amount of water and heat loss in the simulation is comparable with measured respiratory values previously reported. Tracheal temperature control in the avian respiratory system may be used as a model for extinct or rare animals and could have high relevance for explaining how gigantic, long-necked dinosaurs such as sauropoda might have maintained a high metabolic rate.

The mechanical significance of the temporal fasciae in Macaca fascicularis: an investigation using finite element analysis.

Computational finite element analyses (FEAs) of the skull predict structural deformations under user specified loads and constraints, with results normally presented as stress and strain distributions over the skull's surface. The applied loads are generally a representation of the major adductor musculature, with the skull constrained at bite positions and at the articulating joints. However, virtually all analyses ignore potentially important anatomical structures, such as the fasciae that cover the temporalis muscle and attach onto the zygomatic arch. In vivo experimental studies have shown that removal of the temporal fasciae attachment onto the zygomatic arch in Cebus monkeys results in significant bone adaptation and remodeling in this region, suggesting the fasciae play an important role in stabilising the arch during biting. Here we investigate this potential stabilising role by carrying out FEAs of a macaque skull with and without temporal fasciae included. We explore the extent to which the zygomatic arch might be stabilized during biting by a synchronized tensioning of the temporal fasciae, acting to oppose masseteric contraction forces. According to our models, during temporalis muscle bulging the forces generated within the tensioned temporal fasciae are large enough to oppose the pull of the masseter. Further, a near bending-free state of equilibrium within the arch can be reached, even under forceful biting. We show that it is possible to eliminate the high strain gradients in and around the zygomatic arch that are present in past computational studies, with strains being more uniform in magnitude than previously thought.

Biology of the sauropod dinosaurs: the evolution of gigantism.

The herbivorous sauropod dinosaurs of the Jurassic and Cretaceous periods were the largest terrestrial animals ever, surpassing the largest herbivorous mammals by an order of magnitude in body mass. Several evolutionary lineages among Sauropoda produced giants with body masses in excess of 50 metric tonnes by conservative estimates. With body mass increase driven by the selective advantages of large body size, animal lineages will increase in body size until they reach the limit determined by the interplay of bauplan, biology, and resource availability. There is no evidence, however, that resource availability and global physicochemical parameters were different enough in the Mesozoic to have led to sauropod gigantism. We review the biology of sauropod dinosaurs in detail and posit that sauropod gigantism was made possible by a specific combination of plesiomorphic characters (phylogenetic heritage) and evolutionary innovations at different levels which triggered a remarkable evolutionary cascade. Of these key innovations, the most important probably was the very long neck, the most conspicuous feature of the sauropod bauplan. Compared to other herbivores, the long neck allowed more efficient food uptake than in other large herbivores by covering a much larger feeding envelope and making food accessible that was out of the reach of other herbivores. Sauropods thus must have been able to take up more energy from their environment than other herbivores. The long neck, in turn, could only evolve because of the small head and the extensive pneumatization of the sauropod axial skeleton, lightening the neck. The small head was possible because food was ingested without mastication. Both mastication and a gastric mill would have limited food uptake rate. Scaling relationships between gastrointestinal tract size and basal metabolic rate (BMR) suggest that sauropods compensated for the lack of particle reduction with long retention times, even at high uptake rates. The extensive pneumatization of the axial skeleton resulted from the evolution of an avian-style respiratory system, presumably at the base of Saurischia. An avian-style respiratory system would also have lowered the cost of breathing, reduced specific gravity, and may have been important in removing excess body heat. Another crucial innovation inherited from basal dinosaurs was a high BMR. This is required for fueling the high growth rate necessary for a multi-tonne animal to survive to reproductive maturity. The retention of the plesiomorphic oviparous mode of reproduction appears to have been critical as well, allowing much faster population recovery than in megaherbivore mammals. Sauropods produced numerous but small offspring each season while land mammals show a negative correlation of reproductive output to body size. This permitted lower population densities in sauropods than in megaherbivore mammals but larger individuals. Our work on sauropod dinosaurs thus informs us about evolutionary limits to body size in other groups of herbivorous terrestrial tetrapods. Ectothermic reptiles are strongly limited by their low BMR, remaining small. Mammals are limited by their extensive mastication and their vivipary, while ornithsichian dinosaurs were only limited by their extensive mastication, having greater average body sizes than mammals.

Functional Analysis of the Primate Shoulder.

Studies of the shoulder girdle are in most cases restricted to morphological comparisons and rarely aim at elucidating function in a strictly biomechanical sense. To fill this gap, we investigated the basic functional conditions that occur in the shoulder joint and shoulder girdle of primates by means of mechanics. Because most of nonhuman primate locomotion is essentially quadrupedal walking-although on very variable substrates-our analysis started with quadrupedal postures. We identified the mechanical situation at the beginning, middle, and end of the load-bearing stance phase by constructing force parallelograms in the shoulder joint and the scapulo-thoracal connection. The resulting postulates concerning muscle activities are in agreement with electromyographical data in the literature. We determined the magnitude and directions of the internal forces and explored mechanically optimal shapes of proximal humerus, scapula, and clavicula using the Finite Element Method. Next we considered mechanical functions other than quadrupedal walking, such as suspension and brachiation. Quadrupedal walking entails muscle activities and joint forces that require a long scapula, the cranial margin of which has about the same length as the axillary margin. Loading of the hand in positions above the head and suspensory behaviors lead to force flows along the axillary margin and so necessitate a scapula with an extended axillary and a shorter cranial margin. In all cases, the facies glenoidalis is nearly normal to the calculated joint forces. In anterior view, terrestrial monkeys chose a direction of the ground reaction force requiring (moderate) activity of the abductors of the shoulder joint, whereas more arboreal monkeys prefer postures that necessitate activity of the adductors of the forelimb even when walking along branches. The same adducting and retracting muscles are recruited in various forms of suspension. As a mechanical consequence, the scapula is in a more frontal, rather than parasagittal, position on the thorax. In both forms of locomotion-quadrupedal walking and suspension-the compression-resistant clavicula contributes to keeping the shoulder complex distant from the rib cage. Future studies should consider the consequences for thorax shape. The morphological specializations of all Hominoidea match the functional requirements of suspensory behavior. The knowledge of mechanical functions allows an improved interpretation of fossils beyond morphological similarity.

Principles of determination and verification of muscle forces in the human musculoskeletal system: Muscle forces to minimise bending stress.

While there are a growing number of increasingly complex methodologies available to model geometry and material properties of bones, these models still cannot accurately describe physical behaviour of the skeletal system unless the boundary conditions, especially muscular loading, are correct. Available in vivo measurements of muscle forces are mostly highly invasive and offer no practical way to validate the outcome of any computational model that predicts muscle forces. However, muscle forces can be verified indirectly using the fundamental property of living tissue to functional adaptation and finite element (FE) analysis. Even though the mechanisms of the functional adaptation are not fully understood, its result is clearly seen in the shape and inner structure of bones. The FE method provides a precise tool for analysis of the stress/strain distribution in the bone under given loading conditions. The present work sets principles for the determination of the muscle forces on the basis of the widely accepted view that biological systems are optimized light-weight structures with minimised amount of unloaded/underloaded material and hence evenly distributed loading throughout the structure. Bending loading of bones is avoided/compensated in bones under physiological loading. Thus, bending minimisation provides the basis for the determination of the musculoskeletal system loading. As a result of our approach, the muscle forces for a human femur during normal gait and sitting down (peak hip joint force) are obtained such that the bone is loaded predominantly in compression and the stress distribution in proximal and diaphyseal femur corresponds to the material distribution in bone.

Functional shape of the skull in vertebrates: which forces determine skull morphology in lower primates and ancestral synapsids?

In order to determine the extent to which the shape of the synapsid skull is adapted for resisting the mechanical loads to which it is subjected, block- or simple plate-shaped finite-element models were constructed and loaded with external muscle and bite forces in locations estimated to resemble points of application of these forces. These 2D or 3D finite-element models were iteratively loaded and modified by removing elements that experience only low stresses, and the resulting morphologies of the models were compared with fossil skulls of synapsids and the skulls of extant mammals. The results suggest that the stress flows in these unspecific models are very similar to the arrangement of bone material in real skulls. Morphological differences between taxa depend on a few a priori conditions: length and position of the tooth rows in relation to the braincase, arrangement of muscles, position of the orbits, and position of the nasal opening. Given these initial conditions, finite-element analysis consistently reveals the close similarity between stress flows and real skulls. The major difference between mammal-like reptiles and primates is the size of the braincase. This difference accounts for most of the morphological divergence. The postorbital bar seems to be a constructional element of the skull, rather than a means to protect the eyes. The skull shapes of higher primates are determined mainly by masticatory forces and less by external forces acting on the head. This study demonstrates the utility of finite-element modeling for testing hypotheses regarding relationships between form and function in vertebrate skulls.