Three-dimensional kinematic analysis of the pectoral girdle during upside-down locomotion of two-toed sloths (Choloepus didactylus, Linné 1758)

Background

Theria (marsupials and placental mammals) are characterized by a highly mobile pectoral girdle in which the scapula has been shown to be an important propulsive element during locomotion. Shoulder function and kinematics are highly conservative during locomotion within quadrupedal therian mammals. In order to gain insight into the functional morphology and evolution of the pectoral girdle of the two-toed sloth we here analyze the anatomy and the three-dimensional (3D) pattern of shoulder kinematics during quadrupedal suspensory ('upside-down') locomotion.

Methods

We use scientific rotoscoping, a new, non-invasive, markerless approach for x-ray reconstruction of moving morphology (XROMM), to quantify in vivo the 3D movements of all constituent skeletal elements of the shoulder girdle. Additionally we use histologic staining to analyze the configuration of the sterno-clavicular articulation (SCA).

Results

Despite the inverse orientation of the body towards gravity, sloths display a 3D kinematic pattern and an orientation of the scapula relative to the thorax similar to pronograde claviculate mammalian species that differs from that of aclaviculate as well as brachiating mammals. Reduction of the relative length of the scapula alters its displacing effect on limb excursions. The configuration of the SCA maximizes mobility at this joint and demonstrates a tensile loading regime between thorax and limbs.

Conclusions

The morphological characteristics of the scapula and the SCA allow maximal mobility of the forelimb to facilitate effective locomotion within a discontinuous habitat. These evolutionary changes associated with the adoption of the suspensory posture emphasized humeral influence on forelimb motion, but allowed the retention of the plesiomorphic 3D kinematic pattern.     

Analysis of the shoulder in brachiating spider monkeys.

Adult spider monkeys (Ateles geoffroyi and A. paniscus) were conditioned to brachiate on a rope mill (an arboreal analogue of a treadmill). The postures and excursions of the shoulder girdle were studied by cineradiography. These data, together with conventional cinematographic and anatomical studies, permit reassessment of some characteristic structural and functional features of the shoulder in brachiators. During the propulsive phase, the shoulder joint moves caudad from fifth cervical to seventh cervical levels; at the same time, the joint moves dorsad (from a frontal plane midway between the first thoracic vertebra and the manubrium, to a frontal plane through the spinous processes) and slightly mediad. Spider monkeys position the scapula principally on the dorsum of the thorax, in contrast to quadrupedal primates which maintain a more lateral position (even in suspended postures). During brachiation, the scapula rotates a total of 35°; most of this rotation (20°) occurs in the non-propulsive phase when the free arm is being elevated to secure a new handhold. The sigmoidal shape, twisting of proximal relative to distal ends, and elongation of the clavicle in spider monkeys and other brachiators appear to be related to the specialized positioning of the shoulder girdle on the dorsum of the thorax. Shoulder and elbow movements contribute to the efficiency of the swing in terms of the dynamics of a pendulum.     

Functional analysis of the shoulder girdle of cats during locomotion

The movements of the shoulder girdle of eight adult cats during overground stepping were studied, using standard slow motion cinematographic techniques. The patterns of activity of shoulder muscles were examined, using simultaneous intramuscular electromyography. Walking, trotting and galloping steps were analyzed from digitized single motion picture frame images. Angular movements of the shoulder girdle consist of biphasic flexion and extension of the shoulder joint and a monophasic flexion-extension alternation of the scapula on the thorax during each step cycle. In addition, the center of the scapula moves craniad during the swing phase and caudad during the stance phase with respect to a fixed reference point on the animal. Similar vertical movements of the center of the scapula also occur in each step cycle. Results of EMG studies of the 17 muscles capable of acting on the shoulder girdle indicate that three overall patterns of activity are found: (1) a pattern typical of extensor muscles, active during all the extension epochs; (2) a pattern typical of flexor muscles, active during the flexion epoch; and (3) a biphasic pattern of activity, active twice in each step. There data are used, along with a re-examination of previous models of the mechanics of the shoulder girdle of carnivores to examine the function and mechanics of shoulder motion. It is concluded that the rotary and translatory movements of the shoulder girdle during stepping combine to enhance step length.     

A mechanical link model of two-toed sloths: no pendular mechanics during suspensory locomotion

Sloths are morphologically specialized in suspensory quadrupedal locomotion and posture. During steady-state locomotion they utilize a trot-like footfall sequence. Contrasting the growing amount of published accounts of the functional morphology and kinematics of sloth locomotion, no study concerned with the dynamics of their quadrupedal suspensory locomotion has been conducted. Brachiating primates have been shown to travel at low mechanical costs using pendular mechanics, but this is associated with considerable dynamic forces exerted onto the support. To test whether sloth locomotion can be described by simple connected pendulum mechanics, we analyzed the dynamics of sloth locomotion with use of a mechanical segment link model. The model integrates the body segment parameters and is driven by kinematic data with both segment parameters and kinematic data obtained from the same sloth individual. No simple pendular mechanics were present. We then used the model to carry out an inverse dynamic analysis. The analysis allowed us to estimate net limb joint torques and substrate reaction forces during the contact phases. Predominant flexing limb joint torque profiles in the shoulder, elbow, hip, and knee are in stark contrast to published dominant extensor torques in the limb joints of pronograde quadrupedal mammals. This dissimilarity likely reflects the inverse orientation of the sloth towards the gravity vector. Nevertheless, scapular pivot and shoulder seem to provide the strongest torque for progression as expected based on unchanged basic kinematic pattern previously described. Our model predicts that sloths actively reduce the dynamical forces and moments that are transmitted onto the support. We conclude that these findings reflect the need to reduce the risk of breaking supports because in this case sloths would likely be unable to react quickly enough to prevent potentially lethal falls. To achieve this, sloths seem to avoid the dynamical consequences of effective pendular mechanics.     

Functional morphology of the lemuriform wrist joints and the relationship between wrist morphology and positional behavior in arboreal primates

A comparative study of carpal joint structure and function in six Malagasy lemuriforms was undertaken to test predicted morphoclines in carpal joint morphology between pronograde and orthograde arboreal primates. Patterns of movement at the wrist during locomotion were observed and described for the lemuriform species Lemur fulvus and Propithecus verreauxi. Lemur fulvus, which assumes a pronograde posture during locomotion, extends and pronates the wrist during the support phase of quadrupedal walking and running stride cycles. Furthermore, the forearm of this species exhibits some transverse movement across the proximal wrist joint during the support phase. In contrast, the indriid Propithecus maintains the hand and wrist in a flexed and partially supinated position during vertical clinging and suspensory postures. Habitual quadrupedal and vertical postures in Malagasy primates are in turn related to very different patterns of carpal joint morphology and articular mechanics. Those lemurs which are predominantly pronograde share a series of structural features related to stabilizing the antebrachiocarpal joint during extension and mediolateral deviation and the midcarpal joint during pronation: an intraarticular labrum is present on the inner portion of the radiocarpal ligament, the radiocarpal articular surface is quite flat dorsoventrally, the capitate-trapezoid embrasure is expanded dorsally, and development of the radial and ulnar styloids is more pronounced. The wrists of Propithecus, Avahi, and Lepilemur (vertical clingers) differ from those of quadrupedal lemuriforms in possessing a suite of morphological features related to stabilizing the wrist during antebrachiocarpal flexion and midcarpal supination: the radiocarpal articular surface is deeply curved and tilted anteriorly, the dorsal radiocarpal ligament is very broad, thick, and fibrous, the hamate's triquetral facet is directed proximodistally, and the capitate-trapezoid embrasure is dorsally constricted and expanded palmarly. These observed contrasts in carpal form and function are used to define further the morphological features related to orthograde posture in several lineages of arboreal primates. © 1996 Wiley-Liss, Inc.     

The range of passive arm circumduction in primates: Do hominoids really have more mobile shoulders?

Hominoids and lorines are assumed to possess greater shoulder mobility than other primates. This assumption is based on morphological characteristics of the shoulder, rather than on empirical data. However, recent studies have shown that the glenohumeral joint of hominoids is not more mobile than that of other primates (Chan LK. 2007. Glenohumeral mobility in primates. Folia Primatol (Basel) 78(1):1–18), and the thoracic shape of hominoids does not necessarily promote shoulder mobility (Chan LK. 2007. Scapular position in primates. Folia Primatol (Basel) 78(1):19–35). Moreover, lorines differ significantly from hominoids in both these features, thus challenging the assumption that both hominoids and lorines have greater shoulder mobility. The present study aims to test this assumption by collecting empirical data on shoulder mobility in 17 primate species. Passive arm circumduction (a combination of glenohumeral and pectoral girdle movement) was performed on sedated subjects (except humans), and the range measured on the video images of the circumduction. The motion differed among primate species mostly in the craniodorsal directions, the directions most relevant to the animal's ability to brachiate and slow climb. Hylobatids possessed the highest craniodorsal mobility among all primate species studied. However, nonhylobatid hominoids did not have greater craniodorsal mobility than arboreal quadrupedal monkeys, and lorines did not have greater craniodorsal mobility than arboreal quadrupedal prosimians. Nonhylobatid hominoids and lorines had similar craniodorsal mobility, but this was due to a longer clavicle, more dorsal scapula, and lower glenohumeral mobility in the former, and a shorter clavicle, less dorsal scapula, and greater glenohumeral mobility in the latter. This study provides evidence for the reexamination of the brachiation, slow climbing, and vertical climbing hypotheses. Am J Phys Anthropol, 2008. © 2008 Wiley‐Liss, Inc.     

A regression-based 3-D shoulder rhythm

In biomechanical modeling of the shoulder, it is important to know the orientation of each bone in the shoulder girdle when estimating the loads on each musculoskeletal element. However, because of the soft tissue overlying the bones, it is difficult to accurately derive the orientation of the clavicle and scapula using surface markers during dynamic movement. The purpose of this study is to develop two regression models which predict the orientation of the clavicle and the scapula. The first regression model uses humerus orientation and individual factors such as age, gender, and anthropometry data as the predictors. The second regression model includes only the humerus orientation as the predictor. Thirty-eight participants performed 118 static postures covering the volume of the right hand reach. The orientation of the thorax, clavicle, scapula and humerus were measured with a motion tracking system. Regression analysis was performed on the Euler angles decomposed from the orientation of each bone from 26 randomly selected participants. The regression models were then validated with the remaining 12 participants. The results indicate that for the first model, the r² of the predicted orientation of the clavicle and the scapula ranged between 0.31 and 0.65, and the RMSE obtained from the validation dataset ranged from 6.92° to 10.39°. For the second model, the r² ranged between 0.19 and 0.57, and the RMSE obtained from the validation dataset ranged from 6.62° and 11.13°. The derived regression-based shoulder rhythm could be useful in future biomechanical modeling of the shoulder.     

The dynamics of quadrupedal locomotion

This paper presents a dynamical analysis of quadrupedal locomotion, with specific reference to an adult Nubian goat. Measurements of ground reaction forces and limb motion are used to assess variations in intersegmental forces, joint moments, and instantaneous power for three discernible gaits: walking, running, and jumping. In each case, inertial effects of the torso are shown to dominate to the extent that lower-extremity contributions may be considered negligible. Footforces generated by the forelimbs exceed those exerted by the hindlimbs; and, in general, ground reactions increase with speed. The shoulder and hip dominate mechanical energy production during walking, while the knee plays a more significant role in running. In both cases, however, the elbow absorbs energy, and by so doing functions primarily as a damping (control) element. As opposed to either walking or running, jumping requires total horizontal retardation of the body's center of mass. In this instance, generating the necessary vertical thrust amounts to energy absorption at all joints of the lower extremities.     

Inefficient use of inverted pendulum mechanism during quadrupedal walking in the Japanese macaque

In animal walking, the gravitational potential and kinetic energy of the center of mass (COM) fluctuates out-of-phase to reduce the energetic cost of locomotion via an inverted pendulum mechanism, and, in canine quadrupedal walking, up to 70% of the mechanical energy can be recovered. However, the rate of energy recovery for quadrupedal walking in primates has been reported to be comparatively lower. The present study analyzed fluctuations in the potential and kinetic energy of the COM during quadrupedal walking in the Japanese macaque to clarify the mechanisms underlying this inefficient utilization of the inverted pendulum mechanism in primates. Monkeys walked on a wooden walkway at a self-selected speed, and ground reaction forces were measured, using a force platform, to calculate patterns of mechanical energy fluctuation and rates of energy recovery. Our results demonstrated that rates of energy recovery for quadrupedal walking in Japanese macaques were approximately 30–50%, much smaller than those reported for dogs. Comparisons of the patterns of mechanical energy fluctuation suggested that the potential and kinetic energies oscillated relatively more in-phase, and amplitudes did not attain near equality during quadrupedal walking in Japanese macaques, possibly because of greater weight support (reaction force) of the hindlimbs and more protracted forelimbs at touchdown in the Japanese macaque, two of the three commonly accepted locomotor characteristics distinguishing primates from non-primate mammals.     

Shoulder movements during quadrupedal locomotion in arboreal primates

The kinematics of scapula and shoulder joint movements were analyzed in three species of arboreal quadrupedal primates using cineradiography. Our findings indicate that scapular movement is highly important for forelimb movement in primates with this ancestral mode of locomotion. Retroversion of the scapula (syn. caudal rotation or extension) during the stance phase contributes more than 40% to the stride length of the forelimb. Lateral forelimb excursions, a general feature for arboreal primates, are based on complex three-dimensional scapular movements guided by the clavicle. Humeral abduction is achieved by scapular abduction and transversal rotation of the scapula about its longitudinal axis, and is therefore strikingly different from humeral abduction in humans. At the same time, the movements of the shoulder joint are limited to flexion and extension only.     

Contributions of the individual muscles of the shoulder to glenohumeral joint stability during abduction

The aim of this study was to determine the relative contributions of the deltoid and rotator cuff muscles to glenohumeral joint stability during arm abduction. A three-dimensional model of the upper limb was used to calculate the muscle and joint-contact forces at the shoulder for abduction in the scapular plane. The joints of the shoulder girdle-sternoclavicular joint, acromioclavicular joint, and glenohumeral joint-were each represented as an ideal three degree-of-freedom ball-and-socket joint. The articulation between the scapula and thorax was modeled using two kinematic constraints. Eighteen muscle bundles were used to represent the lines of action of 11 muscle groups spanning the glenohumeral joint. The three-dimensional positions of the clavicle, scapula, and humerus during abduction were measured using intracortical bone pins implanted into one subject. The measured bone positions were inputted into the model, and an optimization problem was solved to calculate the forces developed by the shoulder muscles for abduction in the scapular plane. The model calculations showed that the rotator cuff muscles (specifically, supraspinatus, subscapularis, and infraspinatus) by virtue of their lines of action are perfectly positioned to apply compressive load across the glenohumeral joint, and that these muscles contribute most significantly to shoulder joint stability during abduction. The middle deltoid provides most of the compressive force acting between the humeral head and the glenoid, but this muscle also creates most of the shear, and so its contribution to joint stability is less than that of any of the rotator cuff muscles.     

A Biomechanical Model of the Scapulothoracic Joint to Accurately Capture Scapular Kinematics during Shoulder Movements

The complexity of shoulder mechanics combined with the movement of skin relative to the scapula makes it difficult to measure shoulder kinematics with sufficient accuracy to distinguish between symptomatic and asymptomatic individuals. Multibody skeletal models can improve motion capture accuracy by reducing the space of possible joint movements, and models are used widely to improve measurement of lower limb kinematics. In this study, we developed a rigid-body model of a scapulothoracic joint to describe the kinematics of the scapula relative to the thorax. This model describes scapular kinematics with four degrees of freedom: 1) elevation and 2) abduction of the scapula on an ellipsoidal thoracic surface, 3) upward rotation of the scapula normal to the thoracic surface, and 4) internal rotation of the scapula to lift the medial border of the scapula off the surface of the thorax. The surface dimensions and joint axes can be customized to match an individual’s anthropometry. We compared the model to “gold standard” bone-pin kinematics collected during three shoulder tasks and found modeled scapular kinematics to be accurate to within 2mm root-mean-squared error for individual bone-pin markers across all markers and movement tasks. As an additional test, we added random and systematic noise to the bone-pin marker data and found that the model reduced kinematic variability due to noise by 65% compared to Euler angles computed without the model. Our scapulothoracic joint model can be used for inverse and forward dynamics analyses and to compute joint reaction loads. The computational performance of the scapulothoracic joint model is well suited for real-time applications; it is freely available for use with OpenSim 3.2, and is customizable and usable with other OpenSim models.     

Shoulder joint kinematics during elevation measured by ultrasound-based measuring system.

In order to analyze shoulder joint movements, the authors use a ZEBRIS CMS-HS ultrasound-based movement analysis system. In essence, the measurement involves the determination of the spatial position of the 16 anatomical points, which are specified on the basis of the coordinates of ultrasound-based triplets positioned on the upper limb, the scapula, and the thorax; their spatial position is measured in the course of motion. Kinematic characteristics of 74 shoulder joints of 50 healthy persons were identified during elevation in the plane of the scapula. Kinematic characteristics of motion were identified by scapulothoracic, glenohumeral, and humeral elevation angles; range of angles; scapulothoracis and glenohumeral rhythm; scapulothoracic, glenohumeral, and scapuloglenoid ratios; and the relative displacement between the rotation centers of the humerus and the scapula. Motion of the humerus and the scapula relative to each other was characterized by their rotation as well as the relative displacement between the rotation centers of scapula and humerus. The biomechanical model of the shoulder joint during elevation can be described by analyzing the results of the measurements performed.     

新型有原始肢突胴甲鱼的发现及胴甲鱼早期演化的初步探讨

本文记述的曲靖始突鱼(Procondylolepis qujingensis gen.et sp.nov.)是近几年在云南曲靖早泥盆世地层中发现的有肢突胴甲鱼一原始类型。它和已知胴甲鱼(包括早泥盆世无肢突的和中、晚泥盆世有“盔”状肢突的)不同的最大特点是在其肩带与胸鳍相接的肩关节处有原始的肢突和简单的关节窝;胸鳍甲近端的关节区很小。它展现出胴甲鱼类这一高度特化、长期使人迷惑不解的肢突,在胴甲鱼演化史上发展变化的梗概,填补了肢突从无到有中间的缺环,使人了解到胸鳍的具体结构。文中主要根据肢突的有无和特化程度等,对胴甲鱼早期演化史作了初步探讨,将胴甲鱼分为无肢突超目(Abrachicondylia)和有肢突超目(Brachicondylia)两大部分。始突鱼则代表有肢突超目一早期成员。     

Treatment of Combined Injuries of the Axillary and Suprascapular Nerves with Scapulothoracic Dissociation

A 20-year-old man suffered the combined axillary and suprascapular nerve palsies associated with scapulothoracic dissociation by motorcycle accident. The dislocated shoulder girdle was reduced and stabilized with osteosynthesis of the fractured clavicle and reattachment of the trapezius avulsed from the scapular spine for removal of continuous traction force to these damaged nerves. Because of no evidence of recovery on manual muscle test and electromyogram, exploration for these nerves was administered 6 weeks after injury. Although neurolysis of both nerves revealed neural continuity, excessive tension still existed on the suprascapular nerve. It was thought that previous operation in which the shoulder girdle had been reduced and stabilized as much as possible could not achieve complete anatomical reduction of the scapula. As an additional treatment, medial walls of the suprascapular and spinoglenoid notches were shaven to relax the suprascapular nerve. After a year, complete recovery of both the axillary and suprascapular nerve was identified. Although scapulothoracic dissociation is commonly recognized as massive injury of the shoulder girdle with poor prognosis because of existence of accompanied severe neurovascular injuries, there are more than a few cases in which partial damage on the infraclavicular brachial plexus is only accompanied. In case of them, there is the possibility of lesions in continuity of the nerves in which good prognosis might be expected with surgical intervention including early reduction of the shoulder girdle for removal of excessive tension to the damaged nerve.     

The developmental reduction of the marsupial coracoid: A case study in Monodelphis domestica

During their embryogenesis, marsupials develop a unique structure, the shoulder arch, which provides the structural and muscle‐attachment support necessary for the newborn's crawl to the teat. One of the most pronounced and important aspects of the shoulder arch is an enlarged coracoid. After marsupial newborns reach the teat, the shoulder arch is remodeled and the coracoid is reduced to a small process on the scapula. Although an understanding of marsupial coracoid reduction has the potential to provide insights into both, marsupial evolution and the origin of mammals, little is known about the morphological and cellular processes controlling this process. To remedy this situation, this study examined the morphological and cellular mechanisms behind coracoid reduction in the gray short‐tailed opossum, Monodelphis domestica. A quantitative, morphometric study of shoulder girdle development revealed that the coracoid is reduced in size relative to other aspects of the shoulder girdle by growing at a slower rate. Using a series of molecular assays for cell death, no evidence was found for programmed cell death playing a role in the reduction of coracoid size in marsupials (in contrast to hypotheses of previous researchers). Although it is likely the case that coracoid growth is reduced through a relatively lower rate of cellular proliferation, differences in proliferative rates in the coracoid and scapula were not great enough to be quantified using standard molecular assays. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Like Father,Like Son: Assessment of the Morphological Affinities of A.L. 288–1 (A. afarensis), Sts 7 (A. africanus) and Omo 119–73–2718 (Australopithecus sp.) through a Three-Dimensional Shape Analysis of the Shoulder Joint

The postcranial evidence for the Australopithecus genus indicates that australopiths were able bipeds; however, the morphology of the forelimbs and particularly that of the shoulder girdle suggests that they were partially adapted to an arboreal lifestyle. The nature of such arboreal adaptations is still unclear, as are the kind of arboreal behaviors in which australopiths might have engaged. In this study we analyzed the shape of the shoulder joint (proximal humerus and glenoid cavity of the scapula) of three australopith specimens: A.L. 288–1 (A. afarensis), Sts 7 (A. africanus) and Omo 119–73–2718 (Australopithecus sp.) with three-dimensional geometric morphometrics. The morphology of the specimens was compared with that of a wide array of living anthropoid taxa and some additional fossil hominins (the Homo erectus specimen KNM-WT 15000 and the H. neanderthalensis specimen Tabun 1). Our results indicate that A.L. 288–1 shows mosaic traits resembling H. sapiens and Pongo, whereas the Sts 7 shoulder is most similar to the arboreal apes and does not present affinities with H. sapiens. Omo 119–73–2718 exhibits morphological affinities with the more arboreal and partially suspensory New World monkey Lagothrix. The shoulder of the australopith specimens thus shows a combination of primitive and derived traits (humeral globularity, enhancement of internal and external rotation of the joint), related to use of the arm in overhead positions. The genus Homo specimens show overall affinities with H. sapiens at the shoulder, indicating full correspondence of these hominin shoulders with the modern human morphotype.     

Geometry parameters for musculoskeletal modelling of the shoulder system.

A dynamical finite-element model of the shoulder mechanism consisting of thorax, clavicula, scapula and humerus is outlined. The parameters needed for the model are obtained in a cadaver experiment consisting of both shoulders of seven cadavers. In this paper, in particular, the derivation of geometry parameters from the measurement data is described. The results for one cadaver are presented as a typical example. Morphological structures are modelled as geometrical forms. Parameters describing this form are estimated from 3-D position coordinates of a large number of datapoints on the morphological structure, using a least-squares criterion. Muscle and ligament attachments are represented as a plane or as a (curved) line. Muscle paths are determined by a geometrical form of the bony contour around which the muscle is wrapped. Muscle architecture is determined by the distribution of muscle bundles over the attachment area, mapping the distribution of the origin to the insertion. Joint rotation centers are derived from articular surfaces. Hence, muscle moment arms can be calculated. The result of this study is a set of parameters for each cadaver, describing very precisely the geometry of the shoulder mechanism. This set allows positioning of muscle force vectors a posteriori, and recalculation of position coordinates and moment arms for any position of the shoulder.     

Distal Forelimb Kinematics in <Emphasis Type="Italic">Erythrocebus patas</Emphasis> and <Emphasis Type="Italic">Papio anubis</Emphasis> During Walking and Galloping

When using symmetrical gaits, terrestrial digitigrade monkeys adopt less digitigrade, i.e., more palmigrade-like, hand postures as they move with faster speeds. Accordingly, it appears that, in contrast to other mammals, digitigrady is unrelated to cursoriality in primates. However, researchers have not documented the effects of speed on distal forelimb kinematics in faster asymmetrical gaits, i.e., galloping, when ground reaction forces are typically increased owing to the decreased number of contact points during a stride, combined with higher speed. Thus, it remains possible that primates use digitigrade hand postures during these higher-speed asymmetrical gaits. We investigated 3D angles in the wrist joint and metacarpophalangeal joint of 2 habitually digitigrade terrestrial monkeys, Erythrocebus patas and Papio anubis, across a large range of walking and galloping speeds on a motorized treadmill. Nonparametric analyses reveal that angles, and therefore hand postures, are not different at the subject’s walk-gallop transition. Regression analyses show that when walking, digitigrade postures are adopted at slow speeds and more palmigrade-like postures are adopted at fast speeds. Contrary to expectations, there is little change in hand postures across galloping speeds; both subjects maintained palmigrade-like hand postures with substantial joint yield and reextension during support. These results indicate that the hands are always less digitigrade at faster speeds because the joints of the distal forelimb cannot resist the higher ground reaction forces that accompany these higher speed gaits.