Walking-and Running and Jumping-with Dinosaurs and Their Cousins, Viewed Through the Lens of Evolutionary Biomechanics.

Archosauria diversified throughout the Triassic Period before experiencing two mass extinctions near its end ∼201 Mya, leaving only the crocodile-lineage (Crocodylomorpha) and bird-lineage (Dinosauria) as survivors; along with the pterosaurian flying reptiles. About 50 years ago, the "locomotor superiority hypothesis" (LSH) proposed that dinosaurs ultimately dominated by the Early Jurassic Period because their locomotion was superior to other archosaurs'. This idea has been debated continuously since, with taxonomic and morphological analyses suggesting dinosaurs were "lucky" rather than surviving due to being biologically superior. However, the LSH has never been tested biomechanically. Here we present integration of experimental data from locomotion in extant archosaurs with inverse and predictive simulations of the same behaviours using musculoskeletal models, showing that we can reliably predict how extant archosaurs walk, run and jump. These simulations have been guiding predictive simulations of extinct archosaurs to estimate how they moved, and we show our progress in that endeavour. The musculoskeletal models used in these simulations can also be used for simpler analyses of form and function such as muscle moment arms, which inform us about more basic biomechanical similarities and differences between archosaurs. Placing all these data into an evolutionary and biomechanical context, we take a fresh look at the LSH as part of a critical review of competing hypotheses for why dinosaurs (and a few other archosaur clades) survived the Late Triassic extinctions. Early dinosaurs had some quantifiable differences in locomotor function and performance vs. some other archosaurs, but other derived dinosaurian features (e.g., metabolic or growth rates, ventilatory abilities) are not necessarily mutually exclusive from the LSH; or maybe even an opportunistic replacement hypothesis; in explaining dinosaurs' success.

Predictive Simulations of Musculoskeletal Function and Jumping Performance in a Generalized Bird.

Jumping is a common, but demanding, behavior that many animals employ during everyday activity. In contrast to jump-specialists such as anurans and some primates, jumping biomechanics and the factors that influence performance remains little studied for generalized species that lack marked adaptations for jumping. Computational biomechanical modeling approaches offer a way of addressing this in a rigorous, mechanistic fashion. Here, optimal control theory and musculoskeletal modeling are integrated to generate predictive simulations of maximal height jumping in a small ground-dwelling bird, a tinamou. A three-dimensional musculoskeletal model with 36 actuators per leg is used, and direct collocation is employed to formulate a rapidly solvable optimal control problem involving both liftoff and landing phases. The resulting simulation raises the whole-body center of mass to over double its standing height, and key aspects of the simulated behavior qualitatively replicate empirical observations for other jumping birds. However, quantitative performance is lower, with reduced ground forces, jump heights, and muscle-tendon power. A pronounced countermovement maneuver is used during launch. The use of a countermovement is demonstrated to be critical to the achievement of greater jump heights, and this phenomenon may only need to exploit physical principles alone to be successful; amplification of muscle performance may not necessarily be a proximate reason for the use of this maneuver. Increasing muscle strength or contractile velocity above nominal values greatly improves jump performance, and interestingly has the greatest effect on more distal limb extensor muscles (i.e., those of the ankle), suggesting that the distal limb may be a critical link for jumping behavior. These results warrant a re-evaluation of previous inferences of jumping ability in some extinct species with foreshortened distal limb segments, such as dromaeosaurid dinosaurs. Simulations prédictives de la fonction musculo-squelettique et des performances de saut chez un oiseau généralisé Sauter est un comportement commun, mais exigeant, que de nombreux animaux utilisent au cours de leurs activités quotidiennes. Contrairement aux spécialistes du saut tels que les anoures et certains primates, la biomécanique du saut et les facteurs qui influencent la performance restent peu étudiés pour les espèces généralisées qui n'ont pas d'adaptations marquées pour le saut. Les approches de modélisation biomécanique computationnelle offrent un moyen d'aborder cette question de manière rigoureuse et mécaniste. Ici, la théorie du contrôle optimal et la modélisation musculo-squelettique sont intégrées pour générer des simulations prédictives du saut en hauteur maximal chez un petit oiseau terrestre, le tinamou. Un modèle musculo-squelettique tridimensionnel avec 36 actionneurs par patte est utilisé, et une méthode numérique nommée "direct collocation" est employée pour formuler un problème de contrôle optimal rapidement résoluble impliquant les phases de décollage et d'atterrissage. La simulation qui en résulte élève le centre de masse du corps entier à plus du double de sa hauteur debout, et les aspects clés du comportement simulé reproduisent qualitativement les observations empiriques d'autres oiseaux sauteurs. Cependant, les performances quantitatives sont moindres, avec une réduction des forces au sol, des hauteurs de saut et de la puissance musculo-tendineuse. Une manœuvre de contre-mouvement prononcée est utilisée pendant le lancement. Il a été démontré que l'utilisation d'un contre-mouvement est essentielle à l'obtention de hauteurs de saut plus importantes, et il se peut que ce phénomène doive exploiter uniquement des principes physiques pour réussir; l'amplification de la performance musculaire n'est pas nécessairement une raison immédiate de l'utilisation de cette manœuvre. L'augmentation de la force musculaire ou de la vitesse de contraction au-dessus des valeurs nominales améliore grandement la performance de saut et, fait intéressant, a le plus grand effet sur les muscles extenseurs des membres plus distaux (c'est-à-dire ceux de la cheville), ce qui suggère que le membre distal peut être un lien critique pour le comportement de saut. Ces résultats justifient une réévaluation des déductions précédentes de la capacité de sauter chez certaines espèces éteintes avec des segments de membres distaux raccourcis, comme les dinosaures droméosauridés. Voorspellende simulaties van musculoskeletale functie en springprestaties bij een gegeneraliseerde vogel Springen is een veel voorkomend, maar veeleisend, gedrag dat veel dieren toepassen tijdens hun dagelijkse bezigheden. In tegenstelling tot de springspecialisten zoals de anura en sommige primaten, is de biomechanica van het springen en de factoren die de prestaties beïnvloeden nog weinig bestudeerd voor algemene soorten die geen uitgesproken adaptaties voor het springen hebben. Computationele biomechanische modelbenaderingen bieden een manier om dit op een rigoureuze, mechanistische manier aan te pakken. Hier worden optimale controle theorie en musculoskeletale modellering geïntegreerd om voorspellende simulaties te genereren van maximale hoogtesprong bij een kleine grondbewonende vogel, een tinamou. Een driedimensionaal musculoskeletaal model met 36 actuatoren per poot wordt gebruikt, en directe collocatie wordt toegepast om een snel oplosbaar optimaal controleprobleem te formuleren dat zowel de opstijg-als de landingsfase omvat. De resulterende simulatie verhoogt het lichaamszwaartepunt tot meer dan het dubbele van de stahoogte, en belangrijke aspecten van het gesimuleerde gedrag komen kwalitatief overeen met empirische waarnemingen voor andere springende vogels. De kwantitatieve prestaties zijn echter minder, met verminderde grondkrachten, spronghoogtes en spierpeeskracht. Tijdens de lancering wordt een uitgesproken tegenbewegingsmanoeuvre gebruikt. Aangetoond is dat het gebruik van een tegenbeweging van cruciaal belang is voor het bereiken van grotere spronghoogten, en dit fenomeen hoeft alleen op fysische principes te berusten om succesvol te zijn; versterking van de spierprestaties hoeft niet noodzakelijk een proximate reden te zijn voor het gebruik van deze manoeuvre. Het verhogen van de spierkracht of van de contractiesnelheid boven de nominale waarden verbetert de sprongprestatie aanzienlijk, en heeft interessant genoeg het grootste effect op de meer distale extensoren van de ledematen (d.w.z. die van de enkel), wat suggereert dat de distale ledematen een kritieke schakel kunnen zijn voor het springgedrag. Deze resultaten rechtvaardigen een herevaluatie van eerdere conclusies over springvermogen bij sommige uitgestorven soorten met voorgekorte distale ledematen, zoals dromaeosauride dinosauriërs. Prädiktive Simulationen der muskuloskelettalen Funktion und Sprungleistung bei einem generalisierten Vogel Springen ist ein übliches jedoch anstrengendes Verhalten, das viele Tiere bei ihren täglichen Aktivitäten einsetzen. Im Gegensatz zu Springspezialisten, wie Fröschen und einigen Primaten, sind bei allgemeinen Arten, welche keine ausgeprägten Anpassung für Sprungverhalten aufweisen, die Biomechanik beim Springen und die Faktoren, welche die Leistungsfähigkeit beeinflussen, noch wenig untersucht. Computergestützte biomechanische Modellierungsverfahren bieten hier eine Möglichkeit, dies in einer gründlichen, mechanistischen Weise anzugehen. In dieser Arbeit werden die optimale Steuerungstheorie und Muskel-Skelett-Modellierung zusammen eingesetzt, um die maximale Sprunghöhe eines kleinen bodenlebenden Vogels, eines Perlsteisshuhns, zu simulieren und zu prognostizieren. Es wird ein dreidimensionales Muskel-Skelett-Modell mit 36 Aktuatoren pro Bein verwendet, und durch direkte Kollokation wird ein schnell lösbares optimales Steuerungsproblem formuliert, das sowohl die Abstoss- als auch die Landephase umfasst. Die daraus folgende Simulation bringt den Ganzkörperschwerpunkt auf mehr als das Doppelte seiner Standhöhe und entscheidende Aspekte des simulierten Verhaltens entsprechen qualitativ empirischen Beobachtungen für andere springende Vögel. Allerdings ist die quantitative Leistungsfähigkeit geringer, mit reduzierten Bodenkräften, Sprunghöhen und Muskel-Sehnen-Kräften. Beim Abstossen wird ein ausgeprägtes Gegenbewegungsmanöver durchgeführt. Die Durchführung einer Gegenbewegung ist nachweislich entscheidend für das Erreichen grösserer Sprunghöhen, wobei dieses Phänomen möglicherweise nur physikalische Prinzipien auszuschöpfen braucht, um erfolgreich zu sein. Die Verstärkung der Muskelleistung ist daher möglicherweise nicht zwingend ein unmittelbarer Grund für die Verwendung dieses Manövers. Eine Erhöhung der Muskelkraft oder der Kontraktionsgeschwindigkeit über die Nominalwerte hinaus führt zu einer erheblichen Zunahme der Sprungleistung und hat interessanterweise den grössten Effekt bei den weiter distal gelegenen Streckmuskeln der Beine (d.h. bei denjenigen des Sprunggelenks), was darauf hindeutet, dass die distale Gliedmasse ein entscheidendes Element für das Sprungverhalten sein könnte. Diese Ergebnisse geben Anlass zur Überprüfung früherer Schlussfolgerungen hinsichtlich der Sprungfähigkeit einiger ausgestorbener Arten mit verkürzten distalen Gliedmassen, wie beispielsweise bei dromaeosauriden Dinosauriern.

The evolution of pelvic limb muscle moment arms in bird-line archosaurs.

Bipedal locomotion evolved along the archosaurian lineage to birds, shifting from "hip-based" to "knee-based" mechanisms. However, the roles of individual muscles in these changes and their evolutionary timings remain obscure. Using 13 three-dimensional musculoskeletal models of the hindlimbs of bird-line archosaurs, we quantify how the moment arms (i.e., leverages) of 35 locomotor muscles evolved. Our results support two hypotheses: From early theropod dinosaurs to birds, knee flexors' moment arms decreased relative to knee extensors', and medial long-axis rotator moment arms for the hip increased (trading off with decreased hip abductor moment arms). Our results reveal how, from the Triassic Period, bipedal theropod dinosaurs gradually modified their hindlimb form and function, shifting more from hip-based to knee-based locomotion and hip-abductor to hip-rotator balancing mechanisms inherited by birds. Yet, we also discover unexpected ancestral specializations in larger Jurassic theropods, lost later in the bird-line, complicating the paradigm of gradual transformation.

Evolution of forelimb musculoskeletal function across the fish-to-tetrapod transition.

One of the most intriguing questions in vertebrate evolution is how tetrapods gained the ability to walk on land. Although many hypotheses have been proposed, few have been rigorously tested using the fossil record. Here, we build three-dimensional musculoskeletal models of the pectoral appendage in Eusthenopteron, Acanthostega, and Pederpes and quantitatively examine changes in forelimb function across the fin-to-limb transition. Through comparison with extant fishes and tetrapods, we show that early tetrapods share a suite of characters including restricted mobility in humerus long-axis rotation, increased muscular leverage for humeral retraction, but not depression/adduction, and increased mobility in elbow flexion-extension. We infer that the earliest steps in tetrapod forelimb evolution were related to limb-substrate interactions, whereas specializations for weight support appeared later. Together, these results suggest that competing selective pressures for aquatic and terrestrial environments produced a unique, ancestral "early tetrapod" forelimb locomotor mode unlike that of any extant animal.

Patterns of Limb and Epaxial Muscle Activity During Walking in the Fire Salamander, Salamandra salamandra.

Salamanders and newts (urodeles) are often used as a model system to elucidate the evolution of tetrapod locomotion. Studies range from detailed descriptions of musculoskeletal anatomy and segment kinematics, to bone loading mechanics and inferring central pattern generators. A further area of interest has been in vivo muscle activity patterns, measured through electromyography (EMG). However, most prior EMG work has primarily focused on muscles of the forelimb or hindlimb in specific species or the axial system in others. Here we present data on forelimb, hindlimb, and epaxial muscle activity patterns in one species, Salamandra salamandra, during steady state walking. The data are calibrated to limb stride cycle events (stance phase, swing phase), allowing direct comparisons to homologous muscle activation patterns recorded for other walking tetrapods (e.g., lizards, alligators, turtles, mammals). Results demonstrate that Salamandra has similar walking kinematics and muscle activity patterns to other urodele species, but that interspecies variation does exist. In the forelimb, both the m. dorsalis scapulae and m. latissimus dorsi are active for 80% of the forelimb swing phase, while the m. anconaeus humeralis lateralis is active at the swing-stance phase transition and continues through 86% of the stance phase. In the hindlimb, both the m. puboischiofemoralis internus and m. extensor iliotibialis anterior are active for 30% of the hindlimb swing phase, while the m. caudofemoralis is active 65% through the swing phase and remains active for most of the stance phase. With respect to the axial system, both the anterior and posterior m. dorsalis trunci display two activation bursts, a pattern consistent with stabilization and rotation of the pectoral and pelvic girdles. In support of previous assertions, comparison of Salamandra muscle activity timings to other walking tetrapods revealed broad-scale similarities, potentially indicating conservation of some aspects of neuromuscular function across tetrapods. Our data provide the foundation for building and testing dynamic simulations of fire salamander locomotor biomechanics to better understand musculoskeletal function. They could also be applied to future musculoskeletal simulations of extinct species to explore the evolution of tetrapod locomotion across deep-time.

Padrones de actividad muscular epaxial y apendicular durante la cursorialidad de la salamandra-de-fuego, Salamandra salamandra Las salamandras y los tritones (urodelos) son utilizados con frecuencia como un sistema modelo para dilucidar la evolución de la locomoción en los tetrápodos. Los estudios previos varían de descripciones detalladas de la anatomía musculoesquelética y cinemática de los segmentos del cuerpo, a la mecánica de la capacidad de soporte de carga estructural ósea y la generación de padrones centrales. Otra área de interés ha sido los padrones de actividad muscular in vivo, medidos por electromiografía (EMG). Sin embargo, la mayoría de los trabajos anteriores con EMG se han centrado principalmente en los músculos de los miembros anteriores o posteriores en especies específicas o en el sistema axial de otras. En este trabajo, presentamos datos sobre los padrones de actividad muscular en los músculos de los miembros anteriores, posteriores y de la musculatura epaxial en una especie, Salamandra salamandra, durante la marcha continua. Los datos se calibran para los períodos del ciclo de caminar de los miembros (fase de soporte, fase de movimiento), lo que permite comparaciones directas con padrones de activación muscular homólogos registrados para otros tetrápodos cursoriales (por ejemplo, lagartos, caimanes, tortugas y mamíferos). Los resultados demuestran que Salamandra tiene padrones de cinemática cursorial y actividad muscular similares a otras especies de urodelos, pero con variación interespecífica. En los miembros anteriores, ambos los m. dorsalis scapulae y m. latissimus dorsi están activos en 80% de la fase de movimiento del miembro anterior, mientras que el m. anconaeus humeralis lateralis se activa en la transición de la fase de movimiento-soporte y permanece activo en 86% de la fase de soporte. En los miembros posteriores, ambos m. puboischiofemoralis internus y m. extensor iliotibialis anterior están activos en 30% de la fase de movimiento de los miembros posteriores, mientras que el m. caudofemoralis está activo durante el 65% de la fase de movimiento, permaneciendo activo durante la mayor parte de la fase de soporte. Con respecto al sistema axial, las porciones anterior y posterior del m. dorsalis trunci exhibe dos períodos de activación, un padrón consistente con la estabilización y rotación de la cintura pélvica y pectoral. Como sugirido anteriormente, la comparación de los tiempos de actividad muscular de Salamandra con otros tetrápodos cursoriales reveló similitudes en gran escala, lo que podría indicar la conservación de algunos aspectos de la función neuromuscular entre los tetrápodos. Nuestros datos proporcionan una base para la construcción y prueba de simulaciones dinámicas de la biomecánica locomotora de salamandras-de-fuego para comprender mejor las funciones musculoesqueléticas. Nuestros resultados también se pueden aplicar a futuras simulaciones musculoesqueléticas de especies extintas para explorar la evolución de la locomoción de tetrápodos en el tiempo profundo.

Padrões de atividade muscular epaxial e apendicular durante a cursorialidade da salamandra-de-fogo, Salamandra salamandra Salamandras e tritões (urodelos) são freqüentemente utilizados como um sistema modelo para elucidar a evolução da locomoção em tetrápodes. Estudos anteriores variam de descrições detalhadas da anatomia musculoesquelética e cinemática dos segmentos corporais, a mecânica da capacidade de carga estrutural óssea e geradora de padrões centrais. Uma outra área de interesse tem sido os padrões de atividade muscular in vivo, medidos por eletromiografia (EMG). No entanto, a maioria dos trabalhos anteriores de EMG concentrou-se principalmente nos músculos dos membros anteriores ou posteriores em espécies específicas ou no sistema axial de outras. Nesse trabalho, apresentamos dados sobre os padrões de atividade muscular nos membros anteriores, posteriores e musculatura epaxial em uma espécie, Salamandra salamandra, durante caminhada em modo contínuo. Os dados são calibrados para os períodos do ciclo de caminhada dos membros (fase de apoio, fase de movimento), permitindo comparações diretas com padrões de ativação muscular homólogos registrados para outros tetrápodes cursoriais (por exemplo, lagartos, jacarés, tartarugas e mamíferos). Os resultados demonstram que Salamandra possui padrões de cinemática cursorial e atividade muscular semelhantes à outras espécies de urodelos, mas com variação interespecífica. Nos membros anteriores, ambos os m. dorsalis scapulae e m. latissimus dorsi estão ativos em 80% da fase de movimento do membro anterior, enquanto o m. anconaeus humeralis lateralis é ativado na transição da fase de movimento-apoio e continua ativo em 86% da fase de apoio. Nos membros posteriores, ambos m. puboischiofemoralis internus e m. extensor iliotibialis anterior estão ativos em 30% da fase de movimento dos membros posteriores, enquanto o m. caudofemoralis está ativo por 65% da fase de movimento, permanecendo ativo na maior parte da fase de apoio. No que diz respeito ao sistema axial, as porções anterior e posterior do m. dorsalis trunci exibe dois períodos de ativação, um padrão consistente com a estabilização e rotação da cintura peitoral e pélvica. Como préviamente sugerido, a comparação dos tempos de atividade muscular de Salamandra com outros tetrápodes cursoriais revelou similaridades em larga escala, potencialmente indicando a conservação de alguns aspectos da função neuromuscular entre tetrápodes. Os nossos dados fornecem uma base para a construção e testagem de simulações dinâmicas da biomecânica locomotora de salamandras-de-fogo para se entender melhor as funções músculo-esqueléticas. Nossos resultados também podem ser aplicados a futuras simulações músculo-esqueléticas de espécies extintas para explorar a evolução da locomoção de tetrápodes no tempo profundo.

Three-dimensional visualization as a tool for interpreting locomotion strategies in ophiuroids from the Devonian Hunsrück Slate.

Living brittle stars (Echinodermata: Ophiuroidea) employ a very different locomotion strategy to that of any other metazoan: five or more arms coordinate powerful strides for rapid movement across the ocean floor. This mode of locomotion is reliant on the unique morphology and arrangement of multifaceted skeletal elements and associated muscles and other soft tissues. The skeleton of many Palaeozoic ophiuroids differs markedly from that in living forms, making it difficult to infer their mode of locomotion and, therefore, to resolve the evolutionary history of locomotion in the group. Here, we present three-dimensional digital renderings of specimens of six ophiuroid taxa from the Lower Devonian Hunsrück Slate: four displaying the arm structure typical of Palaeozoic taxa (Encrinaster roemeri, Euzonosoma tischbeinianum, Loriolaster mirabilis, Cheiropteraster giganteus) and two (Furcaster palaeozoicus, Ophiurina lymani) with morphologies more similar to those in living forms. The use of three-dimensional digital visualization allows the structure of the arms of specimens of these taxa to be visualized in situ in the round, to our knowledge for the first time. The lack of joint interfaces necessary for musculoskeletally-driven locomotion supports the interpretation that taxa with offset ambulacrals would not be able to conduct this form of locomotion, and probably used podial walking. This approach promises new insights into the phylogeny, functional morphology and ecological role of Palaeozoic brittle stars.

The influence of speed and size on avian terrestrial locomotor biomechanics: Predicting locomotion in extinct theropod dinosaurs.

How extinct, non-avian theropod dinosaurs moved is a subject of considerable interest and controversy. A better understanding of non-avian theropod locomotion can be achieved by better understanding terrestrial locomotor biomechanics in their modern descendants, birds. Despite much research on the subject, avian terrestrial locomotion remains little explored in regards to how kinematic and kinetic factors vary together with speed and body size. Here, terrestrial locomotion was investigated in twelve species of ground-dwelling bird, spanning a 1,780-fold range in body mass, across almost their entire speed range. Particular attention was devoted to the ground reaction force (GRF), the force that the feet exert upon the ground. Comparable data for the only other extant obligate, striding biped, humans, were also collected and studied. In birds, all kinematic and kinetic parameters examined changed continuously with increasing speed, while in humans all but one of those same parameters changed abruptly at the walk-run transition. This result supports previous studies that show birds to have a highly continuous locomotor repertoire compared to humans, where discrete 'walking' and 'running' gaits are not easily distinguished based on kinematic patterns alone. The influences of speed and body size on kinematic and kinetic factors in birds are developed into a set of predictive relationships that may be applied to extinct, non-avian theropods. The resulting predictive model is able to explain 79-93% of the observed variation in kinematics and 69-83% of the observed variation in GRFs, and also performs well in extrapolation tests. However, this study also found that the location of the whole-body centre of mass may exert an important influence on the nature of the GRF, and hence some caution is warranted, in lieu of further investigation.

Gearing effects of the patella (knee extensor muscle sesamoid) of the helmeted guineafowl during terrestrial locomotion.

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.

Using step width to compare locomotor biomechanics between extinct, non-avian theropod dinosaurs and modern obligate bipeds.

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.

Limb proportions show developmental plasticity in response to embryo movement.

Animals have evolved limb proportions adapted to different environments, but it is not yet clear to what extent these proportions are directly influenced by the environment during prenatal development. The developing skeleton experiences mechanical loading resulting from embryo movement. We tested the hypothesis that environmentally-induced changes in prenatal movement influence embryonic limb growth to alter proportions. We show that incubation temperature influences motility and limb bone growth in West African Dwarf crocodiles, producing altered limb proportions which may, influence post-hatching performance. Pharmacological immobilisation of embryonic chickens revealed that altered motility, independent of temperature, may underpin this growth regulation. Use of the chick also allowed us to merge histological, immunochemical and cell proliferation labelling studies to evaluate changes in growth plate organisation, and unbiased array profiling to identify specific cellular and transcriptional targets of embryo movement. This disclosed that movement alters limb proportions and regulates chondrocyte proliferation in only specific growth plates. This selective targeting is related to intrinsic mTOR (mechanistic target of rapamycin) pathway activity in individual growth plates. Our findings provide new insights into how environmental factors can be integrated to influence cellular activity in growing bones and ultimately gross limb morphology, to generate phenotypic variation during prenatal development.

Secondary osteons scale allometrically in mammalian humerus and femur.

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,p<0.005) and Haversian canal area (negative allometry, exponent 0.31,R2 0.45,p<0.005) are significantly related to body mass, independent of phylogeny. This study is the most comprehensive of its kind to date, and allows us to describe overall trends in the scaling behaviour of secondary osteon dimensions, supporting the inference that the osteonal resorption area may be limited by the need to avoid fracture in smaller mammalian species, but the need to maintain osteocyte viability in larger mammalian species.

Big cat, small cat: reconstructing body size evolution in living and extinct Felidae.

The evolution of body mass is a fundamental topic in evolutionary biology, because it is closely linked to manifold life history and ecological traits and is readily estimable for many extinct taxa. In this study, we examine patterns of body mass evolution in Felidae (Placentalia, Carnivora) to assess the effects of phylogeny, mode of evolution, and the relationship between body mass and prey choice in this charismatic mammalian clade. Our data set includes 39 extant and 26 extinct taxa, with published body mass data supplemented by estimates based on condylobasal length. These data were run through 'SURFACE' and 'bayou' to test for patterns of body mass evolution and convergence between taxa. Body masses of felids are significantly different among prey choice groupings (small, mixed and large). We find that body mass evolution in cats is strongly influenced by phylogeny, but different patterns emerged depending on inclusion of extinct taxa and assumptions about branch lengths. A single Ornstein-Uhlenbeck optimum best explains the distribution of body masses when first-occurrence data were used for the fossil taxa. However, when mean occurrence dates or last known occurrence dates were used, two selective optima for felid body mass were recovered in most analyses: a small optimum around 5 kg and a large one around 100 kg. Across living and extinct cats, we infer repeated evolutionary convergences towards both of these optima, but, likely due to biased extinction of large taxa, our results shift to supporting a Brownian motion model when only extant taxa are included in analyses.

What makes an accurate and reliable subject-specific finite element model? A case study of an elephant femur.

Finite element modelling is well entrenched in comparative vertebrate biomechanics as a tool to assess the mechanical design of skeletal structures and to better comprehend the complex interaction of their form-function relationships. But what makes a reliable subject-specific finite element model? To approach this question, we here present a set of convergence and sensitivity analyses and a validation study as an example, for finite element analysis (FEA) in general, of ways to ensure a reliable model. We detail how choices of element size, type and material properties in FEA influence the results of simulations. We also present an empirical model for estimating heterogeneous material properties throughout an elephant femur (but of broad applicability to FEA). We then use an ex vivo experimental validation test of a cadaveric femur to check our FEA results and find that the heterogeneous model matches the experimental results extremely well, and far better than the homogeneous model. We emphasize how considering heterogeneous material properties in FEA may be critical, so this should become standard practice in comparative FEA studies along with convergence analyses, consideration of element size, type and experimental validation. These steps may be required to obtain accurate models and derive reliable conclusions from them.

Comparative axial morphology in pinnipeds and its correlation with aquatic locomotory behaviour.

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.

A systematic review of early versus delayed treatment for type III supracondylar humeral fractures in children.

We performed a systematic review of early versus delayed treatment for type III Gartland supracondylar humeral fractures in children. We identified five non-randomised retrospective studies that fulfilled our criteria. We performed the analysis on 396 patients who sustained a type III supracondylar humeral fracture of which 243 (61.4%) belonged to the early treatment group and 153 (38.6%) belonged to the delayed treatment group. The planned treatment was closed reduction and percutaneous pin fixation. We found that failure of closed reduction and conversion to open reduction was significantly higher in the delayed treatment group (22.9%) as compared with the early treatment group (11.1%). Our study provides evidence that type III supracondylar humeral fractures in children should be treated early within 12h of injury.

Ontogenetic scaling of foot musculoskeletal anatomy in elephants.

This study quantifies the shape change in elephant manus and pes anatomy with increasing body mass, using computed tomographic scanning. Most manus and pes bones, and manus tendons, maintain their shape, or become more gracile, through ontogeny. Contrary to this, tendons of the pes become significantly more robust, suggesting functional adaptation to increasingly high loads. Ankle tendon cross-sectional area (CSA) scales the highest in the long digital extensor, proportional to body mass(1.08+/-0.21), significantly greater than the highest-scaling wrist tendon (extensor carpi ulnaris, body mass(0.69+/-0.09)). These patterns of shape change relate to the marked anatomical differences between the pillar-like manus and tripod-like pes, consistent with differences in fore- and hindlimb locomotor function. The cartilaginous predigits (prepollux and prehallux) of the manus and pes also become relatively more robust through ontogeny, and their pattern of shape change does not resemble that seen in any of the 10 metacarpals and metatarsals. Their CSAs scale above isometry proportional to body mass(0.73+/-0.09) and body mass(0.82+/-0.07) respectively. We infer a supportive function for these structures, preventing collapse of the foot pad during locomotion.

The structure of the cushions in the feet of African elephants (Loxodonta africana).

The uniquely designed limbs of the African elephant, Loxodonta africana, support the weight of the largest terrestrial animal. Besides other morphological peculiarities, the feet are equipped with large subcutaneous cushions which play an important role in distributing forces during weight bearing and in storing or absorbing mechanical forces. Although the cushions have been discussed in the literature and captive elephants, in particular, are frequently affected by foot disorders, precise morphological data are sparse. The cushions in the feet of African elephants were examined by means of standard anatomical and histological techniques, computed tomography (CT) and magnetic resonance imaging (MRI). In both the forelimb and the hindlimb a 6th ray, the prepollex or prehallux, is present. These cartilaginous rods support the metacarpal or metatarsal compartment of the cushions. None of the rays touches the ground directly. The cushions consist of sheets or strands of fibrous connective tissue forming larger metacarpal/metatarsal and digital compartments and smaller chambers which were filled with adipose tissue. The compartments are situated between tarsal, metatarsal, metacarpal bones, proximal phalanges or other structures of the locomotor apparatus covering the bones palmarly/plantarly and the thick sole skin. Within the cushions, collagen, reticulin and elastic fibres are found. In the main parts, vascular supply is good and numerous nerves course within the entire cushion. Vater-Pacinian corpuscles are embedded within the collagenous tissue of the cushions and within the dermis. Meissner corpuscles are found in the dermal papillae of the foot skin. The micromorphology of elephant feet cushions resembles that of digital cushions in cattle or of the foot pads in humans but not that of digital cushions in horses. Besides their important mechanical properties, foot cushions in elephants seem to be very sensitive structures.

A comparison of bilateral uncemented total knee arthroplasty: simultaneous or staged?

In a series of 1304 patients (1867 knees), the results of simultaneous and staged bilateral total knee arthroplasty were compared with each other and with unilateral total knee arthroplasty. The bilateral procedures had a significantly higher rate of complications than unilateral procedures, almost entirely because of thromboembolic problems. However, this did not correspond to an increase in mortality. If a bilateral procedure was indicated, then a simultaneous procedure had no increased risk over a staged procedure. There was no increase in cardiovascular complications, the rate of deep-vein thrombosis or pulmonary embolism or mortality. The rate of infection was lower with a bilateral procedure and the overall revision rate was less than 1% in all groups. The prosthesis functioned as well in all groups in the medium and longer term periods. We feel that simultaneous bilateral total knee arthroplasty is a safe and successful procedure when compared with a staged bilateral procedure. It also has the added benefit of single anaesthetic, reduced costs and decreased total recovery time when compared to a staged bilateral procedure. For these reasons it should be considered as an option in the presence of bilateral knee joint disease.