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Vehicle door latch performance testing presently utilizes uniaxial quasi-static loading conditions. Current technology enables sophisticated virtual testing of a broad range of systems. Door latch failures have been observed in vehicles under a variety of conditions. Typically, these conditions involve multi-axis loading conditions. The loading conditions presented during rollovers on passenger vehicle side door latches have not been published. Rollover crash test results, rollover crashes, and physical Federal Motor Vehicle Safety Standard (FMVSS) 206 latch testing results are reviewed. The creation and validation of a passenger vehicle door latch model is described. The multi-axis loading conditions observed in virtual rollover testing at the latch location are characterized and applied to the virtual testing of a latch in the secondary latch position. The results are then compared with crash test and real world rollover results for the same latch. The results indicate that a door latch that meets the secondary latch position requirements may fail at loads substantially below the FMVSS 206 uniaxial failure loads. In the side impact mode, risks associated with door handle designs and the potential for inertial release can be considered prior to manufacturing with virtual testing. An example case showing the effects of material and spring selection illustrates the potential issues that can be detected in advance of manufacturing. The findings suggest the need for re-examining the relevance of existing door latch testing practices in light of the prevalence of rollover impacts and other impact conditions in today's vehicle fleet environment.

Many of the major locomotor transitions during the evolution of Archosauria, the lineage including crocodiles and birds as well as extinct Dinosauria, were shifts from quadrupedalism to bipedalism (and vice versa). Those occurred within a continuum between more sprawling and erect modes of locomotion and involved drastic changes of limb anatomy and function in several lineages, including sauropodomorph dinosaurs. We present biomechanical computer models of two locomotor extremes within Archosauria in an analysis of joint ranges of motion and the moment arms of the major forelimb muscles in order to quantify biomechanical differences between more sprawling, pseudosuchian (represented the crocodile Crocodylus johnstoni) and more erect, dinosaurian (represented by the sauropodomorph Mussaurus patagonicus) modes of forelimb function. We compare these two locomotor extremes in terms of the reconstructed musculoskeletal anatomy, ranges of motion of the forelimb joints and the moment arm patterns of muscles across those ranges of joint motion. We reconstructed the threedimensional paths of 30 muscles acting around the shoulder, elbow and wrist joints. We explicitly evaluate how forelimb joint mobility and muscle actions may have changed with postural and anatomical alterations from basal archosaurs to early sauropodomorphs. We thus evaluate in which ways forelimb posture was correlated with muscle leverage, and how such differences fit into a broader evolutionary context (i.e. transition from sprawling quadrupedalism to erect bipedalism and then shifting to graviportal quadrupedalism). Our analysis reveals major differences of muscle actions between the more sprawling and erect models at the shoulder joint. These differences are related not only to the articular surfaces but also to the orientation of the scapula, in which extension/flexion movements in Crocodylus (e.g. protraction of the humerus) correspond to elevation/depression in Mussaurus. Muscle action is highly influenced by limb posture, more so than morphology. Habitual quadrupedalism in Mussaurus is not supported by our analysis of joint range of motion, which indicates that glenohumeral protraction was severely restricted. Additionally, some active pronation of the manus may have been possible in Mussaurus, allowing semi-pronation by a rearranging of the whole antebrachium (not the radius against the ulna, as previously thought) via long-axis rotation at the elbow joint. However, the muscles acting around this joint to actively pronate it may have been too weak to drive or maintain such orientations as opposed to a neutral position in between pronation and supination. Regardless, the origin of quadrupedalism in Sauropoda is not only linked to manus pronation but also to multiple shifts of forelimb morphology, allowing greater flexion movements of the glenohumeral joint and a more columnar forelimb posture., Facultad de Ciencias Naturales y Museo

Background The mouse is one of the most widely used animal models to study neuromuscular diseases and test new therapeutic strategies. However, findings from successful pre-clinical studies using mouse models frequently fail to translate to humans due to various factors. Differences in muscle function between the two species could be crucial but often have been overlooked. The purpose of this study was to evaluate and compare muscle excursions in walking between mice and humans. Methods Recently published musculoskeletal models of the mouse hindlimb and human lower limb were used to simulate muscle-tendon dynamics during mouse and human walking, a key daily activity. Muscle fiber length changes (fiber excursions) of 25 muscle homologs in the two species were calculated from these simulations and then compared. To understand potential causes of differences in fiber excursions in walking, joint excursions and muscle moment arms were also compared across one gait cycle. Results Most muscles (19 out of 25 muscles) of the mouse hindlimb had much smaller fiber excursions as compared to human lower limb muscles during walking. For these muscles, fiber excursions in mice were only 48 ± 19% of those in humans. The differences in fiber excursion between the two species were primarily due to the reduced joint excursions and smaller muscle moment arms in mice as compared to humans. Conclusions Since progressive neuromuscular diseases, such as Duchenne muscular dystrophy, are known to be accelerated by damage accumulated from active muscle lengthening, these results suggest that biomechanical differences in muscle function during walking between mice and humans may impede the translations of knowledge gained from mouse models to humans. This knowledge would add a fresh perspective on how pre-clinical studies on mice might be better designed to improve translation to human clinical trials. Electronic supplementary material The online version of this article (10.1186/s13395-017-0143-9) contains supplementary material, which is available to authorized users.

Intra-cortical bone remodelling is a cell-driven process that replaces existing bone tissue with new bone tissue in the bone cortex, leaving behind histological features called secondary osteons. While the scaling of bone dimensions on a macroscopic scale is well known, less is known about how the spatial dimensions of secondary osteons vary in relation to the adult body size of the species. We measured the cross-sectional area of individual intact secondary osteons and their central Haversian canals in transverse sections from 40 stylopodal bones of 39 mammalian species (body mass 0.3–21 000 kg). Scaling analysis of our data shows that mean osteonal resorption area (negative allometry, exponent 0.23,R2 0.54,pR2 0.45,p

Broiler chickens are increasingly at the forefront of global meat production but the consequences of fast growth and selection for an increase in body mass on bird health are an ongoing concern for industry and consumers. To better understand the implications of selection we evaluated energetics and behaviour over the 6-week hatch-to-slaughter developmental period in a commercial broiler. The effect of posture on resting metabolic rate becomes increasingly significant as broilers grow, as standing became more energetically expensive than sitting. The proportion of overall metabolic rate accounted for by locomotor behaviour decreased over development, corresponding to declining activity levels, mean and peak walking speeds. These data are consistent with the inference that broilers allocate energy to activity within a constrained metabolic budget and that there is a reducing metabolic scope for exercise throughout their development. Comparison with similarly sized galliforms reveals that locomotion is relatively energetically expensive in broilers.

Human patellae (kneecaps) are thought to act as gears, altering the mechanical advantage of knee extensor muscles during running. Similar sesamoids have evolved in the knee extensor tendon independently in birds, but it is unknown if these also affect the mechanical advantage of knee extensors. Here, we examine the mechanics of the patellofemoral joint in the helmeted guineafowl Numida meleagris using a method based on muscle and tendon moment arms taken about the patella's rotation centre around the distal femur. Moment arms were estimated from a computer model representing hindlimb anatomy, using hip, knee and patellar kinematics acquired via marker‐based biplanar fluoroscopy from a subject running at 1.6 ms−1 on a treadmill. Our results support the inference that the patella of Numida does alter knee extensor leverage during running, but with a mechanical advantage generally greater than that seen in humans, implying relatively greater extension force but relatively lesser extension velocity.

How extinct, non-avian theropod dinosaurs locomoted is a subject of considerable interest, as is the manner in which it evolved on the line leading to birds. Fossil footprints provide the most direct evidence for answering these questions. In this study, step width—the mediolateral (transverse) distance between successive footfalls—was investigated with respect to speed (stride length) in non-avian theropod trackways of Late Triassic age. Comparable kinematic data were also collected for humans and 11 species of ground-dwelling birds. Permutation tests of the slope on a plot of step width against stride length showed that step width decreased continuously with increasing speed in the extinct theropods (p< 0.001), as well as the five tallest bird species studied (p< 0.01). Humans, by contrast, showed an abrupt decrease in step width at the walk–run transition. In the modern bipeds, these patterns reflect the use of either a discontinuous locomotor repertoire, characterized by distinct gaits (humans), or a continuous locomotor repertoire, where walking smoothly transitions into running (birds). The non-avian theropods are consequently inferred to have had a continuous locomotor repertoire, possibly including grounded running. Thus, features that characterize avian terrestrial locomotion had begun to evolve early in theropod history.

International audience; Over the past two decades, the development of methods for visualizing and analysing specimens digitally, in three and even four dimensions, has transformed the study of living and fossil organisms. However, the initial promise that the widespread application of such methods would facilitate access to the underlying digital data has not been fully achieved. The underlying datasets for many published studies are not readily or freely available, introducing a barrier to verification and reproducibility, and the reuse of data. There is no current agreement or policy on the amount and type of data that should be made available alongside studies that use, and in some cases are wholly reliant on, digital morphology. Here, we propose a set of recommendations for minimum standards and additional best practice for three-dimensional digital data publication, and review the issues around data storage, management and accessibility.

The hindlimb of theropod dinosaurs changed appreciably in the lineage leading to extant birds, becoming more ‘crouched' in association with changes to body shape and gait dynamics. This postural evolution included anatomical changes of the foot and ankle, altering the moment arms and control of the muscles that manipulated the tarsometatarsus and digits, but the timing of these changes is unknown. Here, we report cellular-level preservation of tendon- and cartilage-like tissues from the lower hindlimb of Early Cretaceous Confuciusornis. The digital flexor tendons passed through cartilages, cartilaginous cristae and ridges on the plantar side of the distal tibiotarsus and proximal tarsometatarsus, as in extant birds. In particular, fibrocartilaginous and cartilaginous structures on the plantar surface of the ankle joint of Confuciusornis may indicate a more crouched hindlimb posture. Recognition of these specialized soft tissues in Confuciusornis is enabled by our combination of imaging and chemical analyses applied to an exceptionally preserved fossil., Birds have a more crouched posture compared to their theropod dinosaur ancestors. Here, Jiang and colleagues describe a lower hindlimb of the Early Cretaceous bird Confuciusornis with soft tissues apparently preserved even as molecules, indicating a somewhat more modern posture in ancient birds.

Foot pathology is a common and important health concern in captive rhinoceroses worldwide, but osteopathologies are rarely diagnosed, partly because of a lack of radiographic protocols. Here, we aimed to develop the first radiographic protocol for rhinoceros feet and describe the radiographic anatomy of the white rhinoceros (Ceratotherium simum) hind foot (pes). Computed tomographic images were obtained of nine cadaver pedes from seven different white rhinoceroses and assessed for pathology. A single foot deemed free of pathology was radiographed using a range of different projections and exposures to determine the best protocol. 3D models were produced from the CT images and were displayed with the real radiographs to describe the normal radiographic anatomy of the white rhinoceros pes. An optimal radiographic projection was determined for each bone in the rhinoceros pes focusing on highlighting areas where pathology has been previously described. The projections deemed to be most useful were D60Pr-PlDiO (digit III), D45Pr45M-PlDiLO (digit II), and D40Pr35L-PlDiLO (digit IV). The primary beam was centered 5-7 cm proximal to the cuticle on the digit of interest. Articular surfaces, ridges, grooves, tubercles, processes and fossae were identified. The radiographic protocol we have developed along with the normal radiographic anatomy we have described will allow for more accessible and effective diagnosis of white rhinoceros foot osteopathologies.

Synopsis Over the past century, various modern analogs have been used to infer the evolution of locomotor performance in stem tetrapods and their fish ancestors, with varying success. Here, we conduct a phylogenetic review of these modern analogs, from chondrichthyans to mammals, highlighting the broad spectrum of vertebrate clades and locomotor behaviors. The pros and cons behind utilizing modern analogs for the early stages of the transition from water to land also are discussed. In particular, it is noted that any hypothesis about locomotion not only must be supported by evidence from living animals but must also be consistent with character transformations in the fossil record. A ‘‘totalevidence’’ approach that emphasizes what extinct taxa could not do, rather than focusing on the specifics of how they functioned, is thus recommended. An example of this approach, which investigates mobility of the limb joints in modern semi-aquatic animals and in the Devonian stem tetrapod Ichthyostega, is detailed. We propose that various locomotion behaviors of modern quadrupeds can be ruled out for Ichthyostega, but that forelimb ‘‘crutching’’ motions, as seen in living mudskippers, may have been possible. The potential for movement in other known Devonian stem tetrapods is assessed through an anatomical comparison of limb joint morphology—and associated mobility—with Ichthyostega, and deemed to have been quite similar. Limbs before terrestriality

Foot problems are a major cause of morbidity and mortality in elephants, but are underreported due to difficulties in diagnosis, particularly of conditions affecting the bones and internal structures. Here we evaluate post-mortem computer tomographic (CT) scans of 52 feet from 21 elephants (seven AfricanLoxodonta africanaand 14 AsianElephas maximus), describing both pathology and variant anatomy (including the appearance of phalangeal and sesamoid bones) that could be mistaken for disease. We found all the elephants in our study to have pathology of some type in at least one foot. The most common pathological changes observed were bone remodelling, enthesopathy, osseous cyst-like lesions, and osteoarthritis, with soft tissue mineralisation, osteitis, infectious osteoarthriti, subluxation, fracture and enostoses observed less frequently. Most feet had multiple categories of pathological change (81% with two or more diagnoses, versus 10% with a single diagnosis, and 9% without significant pathology). Much of the pathological change was focused over the middle/lateral digits, which bear most weight and experience high peak pressures during walking. We found remodelling and osteoarthritis to be correlated with increasing age, more enthesopathy in Asian elephants, and more cyst-like lesions in females. We also observed multipartite, missing and misshapen phalanges as common and apparently incidental findings. The proximal (paired) sesamoids can appear fused or absent, and the predigits (radial/tibial sesamoids) can be variably ossified, though are significantly more ossified in Asian elephants. Our study reinforces the need for regular examination and radiography of elephant feet to monitor for pathology and as a tool for improving welfare.

The question of how tetrapod limbs evolved from fins is one of the great puzzles of evolutionary biology. While palaeontologists, developmental biologists, and geneticists have made great strides in explaining the origin and early evolution of limb skeletal structures, that of the muscles remains largely unknown. The main reason is the lack of consensus about appendicular muscle homology between the closest living relatives of early tetrapods: lobe-finned fish and crown tetrapods. In the light of a recent study of these homologies, we re-examined osteological correlates of muscle attachment in the pectoral girdle, humerus, radius, and ulna of early tetrapods and their close relatives. Twenty-nine extinct and six extant sarcopterygians were included in a meta-analysis using information from the literature and from original specimens, when possible. We analysed these osteological correlates using parsimony-based character optimization in order to reconstruct muscle anatomy in ancestral lobe-finned fish, tetrapodomorph fish, stem tetrapods, and crown tetrapods. Our synthesis revealed that many tetrapod shoulder muscles probably were already present in tetrapodomorph fish, while most of the more-distal appendicular muscles either arose later from largely undifferentiated dorsal and ventral muscle masses or did not leave clear correlates of attachment in these taxa. Based on this review and meta-analysis, we postulate a stepwise sequence of specific appendicular muscle acquisitions, splits, and fusions that led from the ancestral sarcopterygian pectoral fin to the ancestral tetrapod forelimb. This sequence largely agrees with previous hypotheses based on palaeontological and comparative work, but it is much more comprehensive in terms of both muscles and taxa. Combined with existing information about the skeletal system, our new synthesis helps to illuminate the genetic, developmental, morphological, functional, and ecological changes that were key components of the fins-to-limbs transition.

This study probes the underlying locomotion principles of earliest organisms that could both swim and walk. We hypothesize that properly coordinated leg and body movements could have provided a substantial benefit toward locomotion on complex media, such as early crawling on sand. In this extended abstract, we summarize some of our recent advances in integrating biology, physics and robotics to gain insight into tetrapod locomotor coordination and control principles. Here, we observe crawling salamanders as a biological model for studying tetrapod locomotion on sloped granular substrates. Further, geometric mechanics tools are used to provide a theoretical framework predicting efficacious body motions on yielding terrain. Finally, we employ these coordination strategies on a robophysical salamander model traversing a sandy slope. This analysis of salamander-like robotic motion in granular media can be seen as a first application of how tools from geometric mechanics can provide insight into the character and principles of legged locomotion.

In quadrupeds the musculature of the hindlimbs is expected to be responsible for generating most of the propulsive locomotory forces, as well as contributing to body support by generating vertical forces. In supporting the body, postural changes from crouched to upright limbs are often associated with an increase of body mass in terrestrial tetrapods. However, felids do not change their crouched limb posture despite undergoing a 300-fold size increase between the smallest and largest extant species. Here, we test how changes in the muscle architecture (masses and lengths of components of the muscle-tendon units) of the hindlimbs and lumbosacral region are related to body mass, to assess whether there are muscular compensations for the maintenance of a crouched limb posture at larger body sizes. We use regression and principal component analyses to detect allometries in muscle architecture, with and without phylogenetic correction. Of the muscle lengths that scale allometrically, all scale with negative allometry (i.e. relative shortening with increasing body mass), whereas all tendon lengths scale isometrically. Only two muscles' belly masses and two tendons' masses scale with positive allometry (i.e. relatively more massive with increasing body mass). Of the muscles that scale allometrically for physiological cross-sectional area, all scale positively (i.e. relatively greater area with increasing body mass). These muscles are mostly linked to control of hip and thigh movements. When the architecture data are phylogenetically corrected, there are few significant results, and only the strongest signals remain. None of the vertebral muscles scaled significantly differently from isometry. Principal component analysis and manovas showed that neither body size nor locomotor mode separate the felid species in morphospace. Our results support the inference that, despite some positively allometric trends in muscle areas related to thigh movement, larger cats have relatively weaker hindlimb and lumbosacral muscles in general. This decrease in power may be reflected in relative decreases in running speeds and is consistent with prevailing evidence that behavioural changes may be the primary mode of compensation for a consistently crouched limb posture in larger cats.

International audience; Graviportal taxa show an allometric increase of the cross-sectional area of supportive bones and are assumed to display microanatomical changes associated with an increase in bone mass. This evokes osteosclerosis (i.e. an increase in bone compactness observed in some aquatic amniotes). The present study investigates the changes in bones' microanatomical organization associated with graviportality and how comparable they are with aquatically acquired osteosclerosis aiming to better understand the adaptation of bone to the different associated functional requirements. Bones of graviportal taxa show microanatomical changes that are not solely attributable to allometry. They display a thicker cortex and a proportionally smaller medullary cavity, with a wider transition zone between these domains. This inner cancellous structure may enable to better enhance energy absorption and marrow support. Moreover, the cross-sectional geometric parameters indicate increased resistance to stresses engendered by bending and torsion, as well as compression. Adaptation to a graviportal posture should be taken into consideration when analyzing possibly amphibious taxa with a terrestrial-like morphology. This is particularly important for palaeoecological inferences about large extinct tetrapods that might have been amphibious and, more generally, for the study of early stages of adaptation to an aquatic life in amniotes.

Three-dimensional reconstruction and modelling of limb joint mobility in the early tetrapod Ichthyostega is used to provide insights into an important step in vertebrate evolution—the transition from swimming to walking. The locomotion of the early tetrapod Ichthyostega is a puzzle because it had an unusual skeleton — a robust ribcage, a spine adapted for up-and-down rather than lateral movement, and paddle-like limbs that seem unsuited to walking. This paper presents a three-dimensional reconstruction and computer modelling of limb-joint mobility based on Ichthyostega fossils. The models suggest that Ichthyostega was unable move its limbs in a walking pattern. Instead it may have shuffled on land using movements similar to those seen in seals. This implies that the transition from swimming to walking was a gradual process. The origin of tetrapods and the transition from swimming to walking was a pivotal step in the evolution and diversification of terrestrial vertebrates. During this time, modifications of the limbs—particularly the specialization of joints and the structures that guide their motions—fundamentally changed the ways in which early tetrapods could move1,2,3,4. Nonetheless, little is known about the functional consequences of limb anatomy in early tetrapods and how that anatomy influenced locomotion capabilities at this very critical stage in vertebrate evolution. Here we present a three-dimensional reconstruction of the iconic Devonian tetrapod Ichthyostega and a quantitative and comparative analysis of limb mobility in this early tetrapod. We show that Ichthyostega could not have employed typical tetrapod locomotory behaviours, such as lateral sequence walking. In particular, it lacked the necessary rotary motions in its limbs to push the body off the ground and move the limbs in an alternating sequence. Given that long-axis rotation was present in the fins of tetrapodomorph fishes5,6,7, it seems that either early tetrapods evolved through an initial stage of restricted shoulder8,9 and hip joint mobility or that Ichthyostega was unique in this respect. We conclude that early tetrapods with the skeletal morphology and limb mobility of Ichthyostega were unlikely to have made some of the recently described Middle Devonian trackways10.

Birds form the largest extant group of bipedal animals and occupy a broad range of body masses, from grams to hundreds of kilograms. Additionally, birds occupy distinct niches of locomotor behaviour, from totally flightless strong runners such as the ratites (moa, kiwi, ostrich) to birds that may walk, dabble on water or fly. We apply a whole-bone approach to investigate allometric scaling trends in the pelvic limb bones (femur, tibiotarsus, tarsometatarsus) from extant and recently extinct birds of greatly different size, and compare scaling between birds in four locomotor groups; flightless, burst-flying, dabbling and flying. We also compare scaling of birds’ femoral cross-sectional properties to data previously collected from cats. Scaling exponents were not significantly different between the different locomotor style groups, but elevations of the scaling relationships revealed that dabblers (ducks, geese, swans) have particularly short and slender femora compared with other birds of similar body mass. In common with cats, but less pronounced in birds, the proximal and distal extrema of the bones scaled more strongly than the diaphysis, and in larger birds the diaphysis occupied a smaller proportion of bone length than in smaller birds. Cats and birds have similar femoral cross-sectional area (CSA) for the same body mass, yet birds’ bone material is located further from the bone’s long axis, leading to higher second and polar moments of area and a greater inferred resistance to bending and twisting. The discrepancy in the relationship between outer diameter to CSA may underlie birds’ reputation for having ‘light’ bones.

The size of an organism is critical to understanding its biology. A telling example comes from the use of sound to localize a potential predatory or prey: this can be based on interaural time difference, but only if the distance between the ears, and hence the size of the head, is large enough. Or consider vision: the diffraction limit constrains the size of image-forming eyes to a certain minimum. The importance of size as a holistic feature of organisms, with consequences for our understanding at every length scale, was taken firmly on board in the early 20th century.

SUMMARY Foot pressure distributions during locomotion have causal links with the anatomical and structural configurations of the foot tissues and the mechanics of locomotion. Elephant feet have five toes bound in a flexible pad of fibrous tissue (digital cushion). Does this specialized foot design control peak foot pressures in such giant animals? And how does body size, such as during ontogenetic growth, influence foot pressures? We addressed these questions by studying foot pressure distributions in elephant feet and their correlation with body mass and centre of pressure trajectories, using statistical parametric mapping (SPM), a neuro-imaging technology. Our results show a positive correlation between body mass and peak pressures, with the highest pressures dominated by the distal ends of the lateral toes (digits 3, 4 and 5). We also demonstrate that pressure reduction in the elephant digital cushion is a complex interaction of its viscoelastic tissue structure and its centre of pressure trajectories, because there is a tendency to avoid rear ‘heel’ contact as an elephant grows. Using SPM, we present a complete map of pressure distributions in elephant feet during ontogeny by performing statistical analysis at the pixel level across the entire plantar/palmar surface. We hope that our study will build confidence in the potential clinical and scaling applications of mammalian foot pressures, given our findings in support of a link between regional peak pressures and pathogenesis in elephant feet.

Regional variation in the axial skeleton of pinnipeds (seals and walruses) and its correlation with aquatic locomotory behaviour is examined using vertebral functional profiles. The results demonstrate clear morpho-functional differences in the thoracolumbar region of modern pinnipeds (Phocidae, Otariidae, Odobenus) that can be strongly linked to swimming style. Phocid seals have a rigid thoracic region attached to a highly flexible lumbar region with long muscular lever arms providing the necessary mobility and leverage to perform pelvic oscillations. Conversely, otariid seals have extremely flexible inter-vertebral joints along the length of the column which should enhance manoeuvrability and turning performance. They also have greater muscular leverage in the anterior thoracic region to support pectoral oscillations. Odobenus (walrus) shows vertebral characteristics most similar to phocids, but with some otariid qualities, consistent with an intermediate or mixed form of aquatic locomotion, with pelvic oscillation dominating over pectoral oscillation. Comparison of the vertebral functional profiles in the fossil taxon Allodesmus kernensis with those of modern pinniped clades reveals that this extinct pinniped may also have used a combination of pectoral and pelvic oscillatory movements during swimming, but in a manner opposite to that of Odobenus, with pectoral oscillatory movements dominating. This study raises questions about the evolution and diversification of pinniped locomotory behaviours, but also provides the necessary framework to begin to examine axial mechanics and locomotory stages in other fossil pinnipedimorphs and their relatives in more detail.

Some tetrapods hang upside down from tree branches when moving horizontally. The ability to walk in quadrupedal suspension has been acquired independently in at least 14 mammalian lineages. During the stance (supportive) phase of quadrupedal suspension, the elbow joint flexor muscles (not the extensors as in upright vertebrates moving overground) are expected to contract to maintain the flexed limb posture. Therefore muscular control in inverted, suspended quadrupeds may require changes of muscle control, and even morphologies, to conditions opposite to those in upright animals. However, the relationships between musculoskeletal morphologies and elbow joint postures during the stance phase in suspended quadrupeds have not been investigated. Our analysis comparing postures and skeletal morphologies in Choloepus (Pilosa), Pteropus (Chiroptera), Nycticebus (Primates) and Cynocephalus (Dermoptera) revealed that the elbow joints of these animals were kept at flexed angles of 70–100 ° during the stance phase of quadrupedal suspension. At these joint angles the moment arms of the elbow joint flexors were roughly maximized, optimizing that component of antigravity support. Our additional measurements from various mammalian species show that suspended quadrupeds have relatively small extensor/flexor ratios in both muscle masses and maximum moment arms. Thus, in contrast to the pattern in normal terrestrial quadrupeds, suspended quadrupeds emphasize flexor over extensor muscles for body support. This condition has evolved independently multiple times, attendant with a loss or reduction of the ability to move in normal upright postures.

Alpacas are increasingly popular as domesticated companion and commercial animals. Their footfall patterns, however, are not well documented. It would be fascinating to know if artificial selection has changed alpacas' locomotor patterns from the ancestral condition in the vicuna. Some members of the Camelidae pace rather than trot, but the gaits of most species have not been studied quantitatively. Thus, investigating alpacas' gaits might contribute to understanding the factors influencing gait choice and evolution. We aimed to quantify the temporal footfall patterns of alpacas to determine observed gaits and to describe gait parameters as a function of speed. Kinematic data (3D motion capture) of locomotor patterns over a range of speeds were collected from four alpacas. We quantified the influence of speed on stance time, swing time, stride time (and frequency), duty factor and stride length, and created gait diagrams for symmetrical and asymmetrical gaits. Alpacas moved using lateral sequence walks, mainly lateral couplets walks and runs with some lateral sequence, singlefoot footfall patterns. Remarkably, the alpacas never truly paced or trotted. At faster speeds, they switched to asymmetrical gaits, predominantly transverse gallops. With increasing speed, stance and stride times decreased, and stride frequencies and stride length increased. Swing times decreased slightly with speed for symmetrical gaits, and a tendency towards an increasing swing time was evident in asymmetrical gaits. It is still uncertain why quadrupeds choose particular gaits and intriguingly, extant camelids (alpacas, dromedary camels) do not trot. The apparent absence of pacing in alpacas deserves further investigation in an experimental and comparative framework. Yet, this absence seems inherited from their vicuna ancestors, indicating that pacing gaits may not be ancestral or common for Camelidae. Future studies should include kinetic and anatomical data to provide insight into whole-body mechanics, and include other unstudied species such as guanacos and vicunas.

Bone geometry is commonly measured on computed tomographic (CT) and X-ray microtomographic (μCT) images. We obtained hundreds of CT, μCT and synchrotron μCT images of bones from diverse species that needed to be analysed remote from scanning hardware, but found that available software solutions were expensive, inflexible or methodologically opaque. We implemented standard bone measurements in a novel ImageJ plugin, BoneJ, with which we analysed trabecular bone, whole bones and osteocyte lacunae. BoneJ is open source and free for anyone to download, use, modify and distribute.

Tyrannosaurs Revisited Tyrannosaurs represent some of the most successful and largest carnivores in Earth's history. An expanding fossil record has allowed studies of their evolution and behavior that now allow broader comparisons with other groups, not just dinosaurs. Brusatte et al. (p. 1481 ) review the biology and evolutionary history of tyrannosaurs and update their phylogenetic relations to include several new fossils. The analysis suggests that tyrannosaurs remained relatively small (less than about 5 meters long) until the Late Cretaceous (about 80 million years ago).

The structure and motion of elephant limbs are unusual compared with those of other animals. Elephants stand and move with straighter limbs (at least when walking), and have limited speed and gait. We devised novel experiments to examine how the limbs of elephants support and propel their mass and to explore the factors that may constrain locomotor performance in these largest of living land animals. We demonstrate that elephant limbs are remarkably compliant even in walking, which maintains low peak forces. Dogma defines elephant limbs as extremely “columnar” for effective weight support, but we demonstrate that limb effective mechanical advantage (EMA) is roughly one-third of that predicted for their size. EMA in elephants is actually smaller than that in horses, which are only one-tenth their mass; it is comparable to human limb values. EMA drops sharply with speed in elephants, as it does in humans. Muscle forces therefore must increase as the limbs become more flexed, and we show how this flexion translates to greater volumes of muscle recruited for locomotion and hence metabolic cost. Surprisingly, elephants use their forelimbs and hindlimbs in similar braking and propulsive roles, not dividing these functions among limbs as was previously assumed or as in other quadrupeds. Thus, their limb function is analogous to four-wheel-drive vehicles. To achieve the observed limb compliance and low peak forces, elephants synchronize their limb dynamics in the vertical direction, but incur considerable mechanical costs from limbs working against each other horizontally.

Plausible poses were identified for the hind limb of Tyrannosaurus rex and three other non-avian theropod dinosaurs at mid-stance of locomotion using constraint-based exclusion. This new method, validated by analysis of two species of birds, involves applying demonstrably realistic geometric and kinetic (force-based) constraints from extant animals to exclude, rather than include, potential poses. Starting with a “configuration space” of millions of candidate poses, we used a step-wise series of criteria to constrict the volume to a small subset of solutions, which can serve as starting points for reconstructing complete stride cycles. It was found that the maximum relative mid-stance limb force, as well as the relative number of configurations at lower forces, decreased with increasing body size. Constraint-based exclusion restricted Tyrannosaurus to a narrow region of neither very columnar nor very flexed poses that may have allowed relatively slow running, but no reasonable combinations of inp...

As one of the most important daily motor activities, human locomotion has been investigated intensively in recent decades. The locomotor functions and mechanics of human lower limbs have become relatively well understood. However, so far our understanding of the motions and functional contributions of the human spine during locomotion is still very poor and simultaneous in-vivo limb and spinal column motion data are scarce. The objective of this study is to investigate the delicate in-vivo kinematic coupling between different functional regions of the human spinal column during locomotion as a stepping stone to explore the locomotor function of the human spine complex. A novel infrared reflective marker cluster system was constructed using stereophotogrammetry techniques to record the 3D in-vivo geometric shape of the spinal column and the segmental position and orientation of each functional spinal region simultaneously. Gait measurements of normal walking were conducted. The preliminary results show that the spinal column shape changes periodically in the frontal plane during locomotion. The segmental motions of different spinal functional regions appear to be strongly coupled, indicating some synergistic strategy may be employed by the human spinal column to facilitate locomotion. In contrast to traditional medical imaging-based methods, the proposed technique can be used to investigate the dynamic characteristics of the spinal column, hence providing more insight into the functional biomechanics of the human spine.

Owing to their cursorial background, ostriches (Struthio camelus) walk and run with high metabolic economy, can reach very fast running speeds and quickly execute cutting manoeuvres. These capabilities are believed to be a result of their ability to coordinate muscles to take advantage of specialized passive limb structures. This study aimed to infer the functional roles of ostrich pelvic limb muscles during gait. Existing gait data were combined with a newly developed musculoskeletal model to generate simulations of ostrich walking and running that predict muscle excitations, force and mechanical work. Consistent with previous avian electromyography studies, predicted excitation patterns showed that individual muscles tended to be excited primarily during only stance or swing. Work and force estimates show that ostrich gaits are partially hip-driven with the bi-articular hip–knee muscles driving stance mechanics. Conversely, the knee extensors acted as brakes, absorbing energy. The digital extensors generated large amounts of both negative and positive mechanical work, with increased magnitudes during running, providing further evidence that ostriches make extensive use of tendinous elastic energy storage to improve economy. The simulations also highlight the need to carefully consider non-muscular soft tissues that may play a role in ostrich gait.

The colossal size and body plan of sauropod dinosaurs are unparalleled in terrestrial vertebrates. However, to-date there have been only limited attempts to examine temporal and phylogenetic patterns in the sauropod bauplan. Here we combine three-dimensional computational models with phylogenetic reconstructions to quantify the evolution of whole-body shape and body segment properties across the sauropod radiation. Limitations associated with the absence of soft tissue preservation in fossils result in large error bars about mean absolute body shape predictions. However, applying any consistent skeleton:body volume ratio to all taxa does yield changes in body shape that appear concurrent with major macroevolutionary events in sauropod history. A caudad shift in centre-of-mass in Middle Triassic Saurischia, associated with the evolution of bipedalism in various dinosaur lineages, was reversed in Late Triassic sauropodomorphs. A craniad centre-of-mass shift coincided with the evolution of quadrupedalism in the Late Triassic, followed by a more striking craniad shift in Late Jurassic–Cretaceous titanosauriforms, which included the largest sauropods. These craniad CoM shifts are strongly correlated with neck enlargement, a key innovation in sauropod evolution and pivotal to their gigantism. By creating a much larger feeding envelope, neck elongation is thought to have increased feeding efficiency and opened up trophic niches that were inaccessible to other herbivores. However, we find that relative neck size and center-of-mass position are not strongly correlated with inferred feeding habits. Instead the craniad center-of-mass positions of titanosauriforms appear closely linked with locomotion and environmental distributions, potentially contributing to the continued success of this group until the end-Cretaceous, with all other sauropods having gone extinct by the early Late Cretaceous.

Mice are one of the most commonly used laboratory animals, with an extensive array of disease models in existence, including for many neuromuscular diseases. The hindlimb is of particular interest due to several close muscle analogues/homologues to humans and other species. A detailed anatomical study describing the adult morphology is lacking, however. This study describes in detail the musculoskeletal geometry and skeletal muscle architecture of the mouse hindlimb and pelvis, determining the extent to which the muscles are adapted for their function, as inferred from their architecture. Using I2KI enhanced microCT scanning and digital segmentation, it was possible to identify 39 distinct muscles of the hindlimb and pelvis belonging to nine functional groups. The architecture of each of these muscles was determined through microdissections, revealing strong architectural specialisations between the functional groups. The hip extensors and hip adductors showed significantly stronger adaptations towards high contraction velocities and joint control relative to the distal functional groups, which exhibited larger physiological cross sectional areas and longer tendons, adaptations for high force output and elastic energy savings. These results suggest that a proximo-distal gradient in muscle architecture exists in the mouse hindlimb. Such a gradient has been purported to function in aiding locomotor stability and efficiency. The data presented here will be especially valuable to any research with a focus on the architecture or gross anatomy of the mouse hindlimb and pelvis musculature, but also of use to anyone interested in the functional significance of muscle design in relation to quadrupedal locomotion.

We examined whether elephants shift to using bouncing (i.e. running) mechanics at any speed. To do this, we measured the three-dimensional centre of mass (CM) motions and torso rotations of African and Asian elephants using a novel multisensor method. Hundreds of continuous stride cycles were recorded in the field. African and Asian elephants moved very similarly. Near the mechanically and metabolically optimal speed (a Froude number (Fr) of 0.09), an inverted pendulum mechanism predominated. With increasing speed, the locomotor dynamics quickly but continuously became less like vaulting and more like bouncing. Our mechanical energy analysis of the CM suggests that at a surprisingly slow speed (approx. 2.2 m s−1, Fr 0.25), the hindlimbs exhibited bouncing, not vaulting, mechanics during weight support. We infer that a gait transition happens at this relatively slow speed: elephants begin using their compliant hindlimbs like pogo sticks to some extent to drive the body, bouncing over their relatively stiff, vaulting forelimbs. Hence, they are not as rigid limbed as typically characterized for graviportal animals, and use regular walking as well as at least one form of running gait.

We developed a three-dimensional, biomechanical computer model of the 36 major pelvic limb muscle groups in an ostrich (Struthio camelus) to investigate muscle function in this, the largest of extant birds and model organism for many studies of locomotor mechanics, body size, anatomy and evolution. Combined with experimental data, we use this model to test two main hypotheses. We first query whether ostriches use limb orientations (joint angles) that optimize the moment-generating capacities of their muscles during walking or running. Next, we test whether ostriches use limb orientations at mid-stance that keep their extensor muscles near maximal, and flexor muscles near minimal, moment arms. Our two hypotheses relate to the control priorities that a large bipedal animal might evolve under biomechanical constraints to achieve more effective static weight support. We find that ostriches do not use limb orientations to optimize the moment-generating capacities or moment arms of their muscles. We infer that dynamic properties of muscles or tendons might be better candidates for locomotor optimization. Regardless, general principles explaining why species choose particular joint orientations during locomotion are lacking, raising the question of whether such general principles exist or if clades evolve different patterns (e.g. weighting of muscle force-length or force-velocity properties in selecting postures). This leaves theoretical studies of muscle moment arms estimated for extinct animals at an impasse until studies of extant taxa answer these questions. Finally, we compare our model’s results against those of two prior studies of ostrich limb muscle moment arms, finding general agreement for many muscles. Some flexor and extensor muscles exhibit self-stabilization patterns (posture-dependent switches between flexor/extensor action) that ostriches may use to coordinate their locomotion. However, some conspicuous areas of disagreement in our results illustrate some cautionary principles. Importantly, tendon-travel empirical measurements of muscle moment arms must be carefully designed to preserve 3D muscle geometry lest their accuracy suffer relative to that of anatomically realistic models. The dearth of accurate experimental measurements of 3D moment arms of muscles in birds leaves uncertainty regarding the relative accuracy of different modelling or experimental datasets such as in ostriches. Our model, however, provides a comprehensive set of 3D estimates of muscle actions in ostriches for the first time, emphasizing that avian limb mechanics are highly three-dimensional and complex, and how no muscles act purely in the sagittal plane. A comparative synthesis of experiments and models such as ours could provide powerful synthesis into how anatomy, mechanics and control interact during locomotion and how these interactions evolve. Such a framework could remove obstacles impeding the analysis of muscle function in extinct taxa.

Stability during locomotion arises from many complex interactions which are not yet fully understood. Studies into how sensory feedback from muscle spindles contributes to this have used neuromechanical simulations, however their accuracy is limited due to the lack of available physiological data. This presentation outlines the methods used to develop a musculoskeletal model of the mouse hindlimb, which will be used to predict responses to perturbations which occur with or without sensory feedback. The 3D musculoskeletal model, which represents bones, joints and 39 muscles of the mouse hindlimb, was created using SIMM (Musculographics, USA) through a combination of muscle architecture data and I2KI enhanced microCT scanning. Architecture data, such as muscle mass, pennation angle and fibre length were gathered from microdissections, while muscle geometry was determined using I2KI microCT scanning, a non-destructive method of determining muscle paths and attachment points. Muscle moment arms predicted by the model were found to agree with measured data from previous studies. In future work, in vivo optogenetics will be used to characterise changes in gait and muscle forces which occur in free running mice in response to induced neural and mechanical perturbations with or without sensory feedback from hindlimb muscles. These physiological data will be used to update and refine the musculoskeletal model.

SUMMARYFor centuries, elephant locomotion has been a contentious and confusing challenge for locomotion scientists to understand, not only because of technical difficulties but also because elephant locomotion is in some ways atypical of more familiar quadrupedal gaits. We analyzed the locomotor kinematics of over 2400 strides from 14 African and 48 Asian elephant individuals (body mass 116-4632 kg) freely moving over ground at a 17-fold range of speeds, from slow walking at 0.40 m s-1 to the fastest reliably recorded speed for elephants, 6.8 m s-1. These data reveal that African and Asian elephants have some subtle differences in how size-independent kinematic parameters change with speed. Although elephants use a lateral sequence footfall pattern, like many other quadrupeds, they maintain this footfall pattern at all speeds, shifting toward a 25% phase offset between limbs (singlefoot) as they increase speed. The duty factors of elephants are greater for the forelimbs than for the hindlimbs, so an aerial phase for the hindquarters is reached at slower speeds than for the forequarters. This aerial phase occurs at a Froude number of around 1,matching theoretical predictions. At faster speeds, stance and swing phase durations approach asymptotes, with the duty factor beginning to level off,concurrent with an increase in limb compliance that likely keeps peak forces relatively low. This increase of limb compliance is reflected by increased compression of the hindlimbs. Like other tetrapods, smaller elephants are relatively more athletic than larger ones, but still move very similarly to adults even at

Archosaurian reptiles evolved a more erect posture and parasagittal gait early on. Early dinosaurs were the first habitual striding bipeds, a trait retained by living birds. Yet there is much more to archosaur locomotor evolution than these two transitions. I review our understanding of the pattern of locomotor evolution from the first archosaurs to Crocodylia and Neornithes, outlining where transitions of locomotor function evolved. I evaluate current research approaches, advocating more experimental work on extant animals to establish rigorous form–function relationships, and more biomechanical research that is bolstered by validation and sensitivity analysis of its assumptions, methods, and results. To cite this article: J.R. Hutchinson, C. R. Palevol 5 (2006) .

Here we redescribe the holotype and only specimen of Segisaurus halli, a small Early Jurassic dinosaur and the only theropod known from the Navajo Sandstone. Our study highlights several important ...

Muscle moment arms are important determinants of muscle function; however, it is challenging to determine moment arms by inspecting bone specimens alone, as muscles have curvilinear paths that change as joints rotate. The goals of this study were to (1) develop a three-dimensional graphics-based model of the musculoskeletal system of the Cretaceous theropod dinosaur Tyrannosaurus rex that predicts muscle-tendon unit paths, lengths, and moment arms for a range of limb positions; (2) use the model to determine how the T. rex hindlimb muscle moment arms varied between crouched and upright poses; (3) compare the predicted moment arms with previous assessments of muscle function in dinosaurs; (4) evaluate how the magnitudes of these moment arms compare with those in other animals; and (5) integrate these findings with previous biomechanical studies to produce a revised appraisal of stance, gait, and speed in T. rex. The musculoskeletal model includes ten degrees of joint freedom (flexion/extension, ab...

I used a simple mathematical model of the inverse dynamics of locomotion to estimate the minimum muscle masses required to maintain quasi-static equilibrium about the four main limb joints at mid-stance of fast running. Models of 10 extant taxa (a human, a kangaroo, two lizards, an alligator, and five birds) were analyzed in various bipedal poses to examine how anatomy, size, limb orientation, and other model parameters influence running ability. I examined how the muscle masses required for fast running compare to the muscle masses that are actually able to exert moments about the hip, knee, ankle, and toe joints, to see how support ability varies across the limb. I discuss the assumptions and limitations of the models, using sensitivity analysis to see how widely the results differed with feasible parameter input values. Even with a wide range of input values, the models validated the analysis procedure. Animals that are known to run bipedally were calculated as able to preserve quasi-static equilibrium about their hindlimb joints at mid-stance, whereas non-bipedal runners (iguanas and alligators) were recognized as having too little muscle mass to run quickly in bipedal poses. Thus, this modeling approach should be reliable for reconstructing running ability in extinct bipeds such as nonavian dinosaurs. The models also elucidated how key features are important for bipedal running capacity, such as limb orientation, muscle moment arms, muscle fascicle lengths, and body size. None of the animals modeled had extensor muscle masses acting about any one joint that were 7% or more of their body mass, which provides a reasonable limit for how much muscle mass is normally apportioned within a limb to act about a particular joint. The models consistently showed that a key biomechanical limit on running ability is the capacity of ankle extensors to generate sufficiently large joint moments. Additionally, the analysis reveals how large ratite birds remain excellent runners despite their larger size; they have apomorphically large extensor muscles with relatively high effective mechanical advantage. Finally, I reconstructed the evolution of running ability in the clade Reptilia, showing that the ancestors of extant birds likely were quite capable runners, even though they had already reduced key hip extensors such as M. caudofemoralis longus.

Dinosauromorphs evolved a wide diversity of hind limb skeletal morphologies, suggesting highly divergent articular soft tissue anatomies. However, poor preservation of articular soft tissues in fossils has hampered any follow-on functional inferences. We reconstruct the hip joint soft tissue anatomy of non-dinosaurian dinosauromorphs and early dinosaurs using osteological correlates derived from extant sauropsids and infer trends in character transitions along the theropod and sauropodomorph lineagues. Femora and pelves of 107 dinosauromorphs and outgroup taxa were digitized using 3D imaging techniques. Key transitions were estimated using maximum likelihood ancestral state reconstruction. The hips of dinosauromorphs possessed wide a disparity of soft tissue morphologies beyond the types and combinations exhibited by extant archosaurs. Early evolution of the dinosauriform hip joint was characterized by the retention of a prominent femoral hyaline cartilage cone in post-neonatal individuals, with the cartilage cone independently reduced within theropods and sauropodomorphs. The femur of Dinosauriformes possessed a fibrocartilage sleeve on the metaphysis, which surrounded a hyaline core. The acetabulum of Dinosauriformes possessed distinct labrum and antitrochanter structures. In sauropodomorphs, hip congruence was maintained by thick hyaline cartilage on the femoral head, whereas theropods relied on acetabular tissues such as ligaments and articular pads. In particular, the craniolaterally ossified hip capsule of non- Avetheropoda neotheropods permitted mostly parasagittal femoral movements. These data indicate that the dinosauromorph hip underwent mosaic evolution within the saurischian lineage and that sauropodomorphs and theropods underwent both convergence and divergence in articular soft tissues, correlated with transitions in body size, locomotor posture, and joint loading.

Emus (Dromaius novaehollandiae) are exclusively terrestrial, bipedal and cursorial ratites with some similar biomechanical characteristics to humans. Their growth rates are impressive as their body mass increases eighty-fold from hatching to adulthood whilst maintaining the same mode of locomotion throughout life. These ontogenetic characteristics stimulate biomechanical questions about the strategies that allow emus to cope with their rapid growth and locomotion, which can be partly addressed via scaling (allometric) analysis of morphology. In this study we have collected pelvic limb anatomical data (muscle architecture, tendon length, tendon mass and bone lengths) and calculated muscle physiological cross sectional area (PCSA) and average tendon cross sectional area from emus across three ontogenetic stages (n=17, body masses from 3.6 to 42 kg). The data were analysed by reduced major axis regression to determine how these biomechanically relevant aspects of morphology scaled with body mass. Muscle mass and PCSA showed a marked trend towards positive allometry (26 and 27 out of 34 muscles respectively) and fascicle length showed a more mixed scaling pattern. The long tendons of the main digital flexors scaled with positive allometry for all characteristics whilst other tendons demonstrated a less clear scaling pattern. Finally, the two longer bones of the limb (tibiotarsus and tarsometatarsus) also exhibited positive allometry for length and the two others (femur and first phalanx of digit III) had trends towards isometry. These results indicate that emus experience a relative increase in their muscle force-generating capacities, as well as potentially increasing the force-sustaining capacities of their tendons, as they grow. Furthermore, we have clarified anatomical descriptions and provided illustrations of the pelvic limb muscle-tendon units in emus.

The three-dimensional anatomy of the ostrich (Struthio camelus) knee (femorotibial, femorofibular, and femoropatellar) joint has scarcely been studied, and could elucidate certain mechanobiological properties of sesamoid bones. The adult ostrich is unique in that it has double patellae, while another similar ratite bird, the emu, has none. Understanding why these patellae form and what purpose they may serve is dually important for future studies on ratites as well as for understanding the mechanobiological characteristics of sesamoid bone development. For this purpose, we present a three-dimensional anatomical study of the ostrich knee joint, detailing osteology, ligaments and menisci, and myology. We have identified seven muscles which connect to the two patellae and compare our findings to past descriptions. These descriptions can be used to further study the biomechanical loading and implications of the double patella in the ostrich.