John R. Hutchinson

BoneJ: Free and extensible bone image analysis in ImageJ

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.

Tyrannosaurus was not a fast runner

The fastest gait and speed of the largest theropod (carnivorous) dinosaurs, such as Tyrannosaurus, is controversial. Some studies contend that Tyrannosaurus was limited to walking, or at best an 11 m s-1 top speed, whereas others argue for at least 20 m s-1 running speeds. We demonstrate a method of gauging running ability by estimating the minimum mass of extensor (supportive) muscle needed for fast running. The models predictions are validated for living alligators and chickens. Applying the method to small dinosaurs corroborates other studies by showing that they could have been competent runners. However, models show that in order to run quickly, an adult Tyrannosaurus would have needed an unreasonably large mass of extensor muscle, even with generous assumptions. Therefore, it is doubtful that Tyrannosaurus and other huge dinosaurs (∼6,000 kg) were capable runners or could reach high speeds.

Adductors, abductors, and the evolution of archosaur locomotion

Abstract Living crocodilians (Crocodylia) and birds (Neornithes) differ in many aspects of hindlimb anatomy and locomotor function. How did this disparity evolve? We integrate information from fossils with functional descriptions of locomotion in living crocodilians and birds, using a phylogenetic perspective. We then outline the major changes in three-dimensional control of the hip joint along the line from the ancestral archosaur to birds. Our analysis reveals that most aspects of hip morphology and function in Alligator are ancestral for Archosauria. Femoral protractors and retractors are located cranial and caudal to the hip, respectively. Similarly, femoral adductors and abductors are located ventral and dorsal to the hip. Transformations of this ancestral pattern on the line to birds involved modifications in osteology, myology, and neural control. In some cases, homologous muscles changed function by acquiring new activity patterns. In others, activity was conserved, but origins and/or insertions were altered. Fossil theropods document the stepwise evolution of a novel mechanism of limb adduction/abduction involving long-axis rotation of the femur. This mechanism accounts for the conspicuous absence of significant musculature ventral and dorsal to the hip joint in extant birds.

Biomechanical modeling and sensitivity analysis of bipedal running ability. II. Extinct taxa

Using an inverse dynamics biomechanical analysis that was previously validated for extant bipeds, I calculated the minimum amount of actively contracting hindlimb extensor muscle that would have been needed for rapid bipedal running in several extinct dinosaur taxa. I analyzed models of nine theropod dinosaurs (including birds) covering over five orders of magnitude in size. My results uphold previous findings that large theropods such as Tyrannosaurus could not run very quickly, whereas smaller theropods (including some extinct birds) were adept runners. Furthermore, my results strengthen the contention that many nonavian theropods, especially larger individuals, used fairly upright limb orientations, which would have reduced required muscular force, and hence muscle mass. Additional sensitivity analysis of muscle fascicle lengths, moment arms, and limb orientation supports these conclusions and points out directions for future research on the musculoskeletal limits on running ability. Although ankle extensor muscle support is shown to have been important for all taxa, the ability of hip extensor muscles to support the body appears to be a crucial limit for running capacity in larger taxa. I discuss what speeds were possible for different theropod dinosaurs, and how running ability evolved in an inverse relationship to body size in archosaurs. J. Morphol. 262:441–461, 2004.

Tyrannosaur Paleobiology: New Research on Ancient Exemplar Organisms

Tyrannosaurs Revisited Tyrannosaurs represent some of the most successful and largest carnivores in Earths 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). Tyrannosaurs, the group of dinosaurian carnivores that includes Tyrannosaurus rex and its closest relatives, are icons of prehistory. They are also the most intensively studied extinct dinosaurs, and thanks to large sample sizes and an influx of new discoveries, have become ancient exemplar organisms used to study many themes in vertebrate paleontology. A phylogeny that includes recently described species shows that tyrannosaurs originated by the Middle Jurassic but remained mostly small and ecologically marginal until the latest Cretaceous. Anatomical, biomechanical, and histological studies of T. rex and other derived tyrannosaurs show that large tyrannosaurs could not run rapidly, were capable of crushing bite forces, had accelerated growth rates and keen senses, and underwent pronounced changes during ontogeny. The biology and evolutionary history of tyrannosaurs provide a foundation for comparison with other dinosaurs and living organisms.

A clinical and pharmacokinetic study of isolated limb perfusion with heat and melphalan for melanoma

The pharmacokinetics of isolated limb perfusion were studied to see what melphalan concentrations were achieved and how effective the isolation was. Twenty‐eight patients received 32 limb perfusions with heat and melphalan for locally recurrent or level V melanoma. Melphalan was given 0.75 mg/kg for axillary/popliteal or 1.2 mg/kg for femoral perfusions with heat (perfusate 42°C, limb 40°C) for 1 hour. Melphalan concentratives were measured by high‐performance liquid chromatography in seven patients. Peak perfusate melphalan concentrations were 6.1 to 115 mg/ml, which was one to two logs higher than peak systemic concentratives of melphalan. Isolation of the perfusate circuit from the systemic circulation was better for axillary and popliteal perfusions than for femoral perfusions (P < 0.05). Complete responses were seen in 81% of evaluable patients; long‐term local control was achieved in most patients, although many developed hematogenous metastases. Toxicity included erythema and edema in all, mild leukopenia in two, neuropathy in two, and amputation was required in one patient. Improvements in surgical technique include regional anesthesia to reduce vasospasms and transcutaneous measurement of fluorescein to measure leak. Perfusion with heat and melphalan remains the treatment of choice for in‐transit metastases from melanoma.

Three-dimensional limb joint mobility in the early tetrapod Ichthyostega

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 move. 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 fishes, it seems that either early tetrapods evolved through an initial stage of restricted shoulder 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 trackways.

Analysis of hindlimb muscle moment arms in Tyrannosaurus rex using a three-dimensional musculoskeletal computer model: implications for stance, gait, and speed

Abstract 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...

Biomechanics: Are fast-moving elephants really running?

It is generally thought that elephants do not run, but there is confusion about how fast they can move across open terrain and what gait they use at top speed. Here we use video analysis to show that Asian elephants (Elephas maximus L.) can move at surprisingly high speeds of up to 6.8 m s−1 (25 km h−1) and that, although their gait might seem to be a walk even at this speed, some features of their locomotion conform to definitions of running.

The locomotor kinematics of Asian and African elephants : changes with speed and size

SUMMARY For 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 <500 kg. At any particular speed they adopt greater relative stride frequencies and relative stride lengths compared to larger elephants. This extends to near-maximal locomotor performance as well - smaller elephants reach greater Froude numbers and smaller duty factors, hence likely reach relatively greater peak loads on their limbs and produce this force more rapidly. A variety of lines of kinematic evidence support the inference that elephants change their mechanics near a Froude number of 1 (if not at slower speeds), at least to using more compliant limbs, if not spring-like whole-body kinetics. In some ways, elephants move similarly to many other quadrupeds, such as increasing speed mainly by increasing stride frequency (except at fast speeds), and they match scaling predictions for many stride parameters. The main difference from most other animals is that elephants never change their footfall pattern to a gait that uses a whole-body aerial phase. Our large dataset establishes what the normal kinematics of elephant locomotion are, and can also be applied to identify gait abnormalities that may signal musculoskeletal pathologies, a matter of great importance to keepers of captive elephants.