Long-axis Rotation Research Articles

Exploring elbow kinematics in the central bearded dragon (Pogona vitticeps) using XROMM: Implications for the role of forearm long-axis rotation in non-avian reptile posture and mobility.

The functional morphology and kinematics of the elbow joint remain relatively understudied in squamates. Previous investigations of lizard elbow morphology and kinematics suggest long-axis rotation (LAR) of the radius and ulna during stance allows the manus to remain pronated during forelimb retraction. Using XROMM (X-ray Reconstruction of Moving Morphology), we explored the range of 3D movements and kinematics of the humerus, radius, and ulna in three adult male Central bearded dragons (Pogona vitticeps) during trackway walking. Our data indicate that the elbow joint of P. vitticeps experiences significant rotations in all three dimensions and that the radius and ulna adduct and rotate laterally on their long axes relative to the elbow joint and to one another during stance. These movements allow the distal ends of the radius and ulna to remain in a configuration necessary for manus pronation. These data support previous inferences that the radius and ulna of lizards move independently at the wrist joint. We suggest that independent LAR of the radius and ulna relative to the elbow joint and to one another may be an ancestral mechanism in lizards and perhaps more broadly across non-avian reptiles.

Real-time guidance by deep learning of experienced operators to improve the standardization of echocardiographic acquisitions.

Impaired standardization of echocardiograms may increase inter-operator variability. This study aimed to determine whether the real-time guidance of experienced sonographers by deep learning (DL) could improve the standardization of apical recordings. Patients (n = 88) in sinus rhythm referred for echocardiography were included. All participants underwent three examinations, whereof two were performed by sonographers and the third by cardiologists. In the first study period (Period 1), the sonographers were instructed to provide echocardiograms for the analyses of the left ventricular function. Subsequently, after brief training, the DL guidance was used in Period 2 by the sonographer performing the second examination. View standardization was quantified retrospectively by a human expert as the primary endpoint and the DL algorithm as the secondary endpoint. All recordings were scored in rotation and tilt both separately and combined and were categorized as standardized or non-standardized. Sonographers using DL guidance had more standardized acquisitions for the combination of rotation and tilt than sonographers without guidance in both periods (all P ≤ 0.05) when evaluated by the human expert and DL [except for the apical two-chamber (A2C) view by DL evaluation]. When rotation and tilt were analysed individually, A2C and apical long-axis rotation and A2C tilt were significantly improved, and the others were numerically improved when evaluated by the echocardiography expert. Furthermore, all, except for A2C rotation, were significantly improved when evaluated by DL (P < 0.01). Real-time guidance by DL improved the standardization of echocardiographic acquisitions by experienced sonographers. Future studies should evaluate the impact with respect to variability of measurements and when used by less-experienced operators. NCT04580095.

Suction feeding of West African lungfish (Protopterus annectens): An XROMM analysis of jaw mechanics, cranial kinesis, and hyoid mobility.

ABSTRACTSuction feeding in fishes is characterized by rapid cranial movements, but extant lungfishes (Sarcopterygii: Dipnoi) exhibit a reduced number and mobility of cranial bones relative to actinopterygian fishes. Despite fusion of cranial elements, lungfishes are proficient at suction feeding, though the impacts of novel cranial morphology and reduced cranial kinesis on feeding remain poorly understood. We used X-ray reconstruction of moving morphology (XROMM) to study the kinematics of seven mobile elements (neurocranium, upper jaw, lower jaw, tongue, ceratohyal, clavicle, and cranial rib) and two muscles (costoclavicular portion of the hypaxialis and rectus cervicis) during the feeding strikes of West African lungfish (Protopterus annectens). We found that feeding by P. annectens on non-evasive prey is relatively slow, with a mean time to peak gape of 273 ms. Lower jaw depression and clavicular rotation were hinge-like, with one degree of freedom, but the ceratohyals rotated in a complex motion involving depression and long-axis rotation. We quantified the relative contributions to oral cavity volume change (RCVC) and found that oral cavity expansion is created primarily by ceratohyal and clavicle motion. P. annectens suction feeds relatively slowly but successfully through muscle shortening of hypaxial and rectus cervicis muscles contributing to hyoid mobility.

A therian mammal with sprawling kinematics? Gait and 3D forelimb X-ray motion analysis in tamanduas.

Therian mammals are known to move their forelimbs in a parasagittal plane, retracting the mobilised scapula during stance phase. Non-cursorial therian mammals often abduct the elbow out of the shoulder-hip parasagittal plane. This is especially prominent in Tamandua (Xenarthra), which suggests they employ aspects of sprawling (e.g. lizard-like) locomotion. Here, we tested whether tamanduas use sprawling forelimb kinematics, i.e. a largely immobile scapula with pronounced lateral spine bending and long-axis rotation of the humerus. We analysed high-speed videos and used X-ray motion analysis of tamanduas walking and balancing on branches of varying inclinations and provide a quantitative characterization of gaits and forelimb kinematics. Tamanduas displayed lateral sequence/lateral couplets on flat ground and horizontal branches, but increased diagonality on steeper inclines and declines, resulting in lateral sequence/diagonal couplets gaits. This result provides further evidence for high diagonality in arboreal species, probably maximising stability in arboreal environments. Further, the results reveal a mosaic of sprawling and parasagittal kinematic characteristics. The abducted elbow results from a constantly internally rotated scapula about its long axis and a retracted humerus. Scapula retraction contributes considerably to stride length. However, lateral rotation in the pectoral region of the spine (range: 21deg) is higher than reported for other therian mammals. Instead, it is similar to that of skinks and alligators, indicating an aspect generally associated with sprawling locomotion is characteristic for forelimb kinematics of tamanduas. Our study contributes to a growing body of evidence of highly variable non-cursorial therian mammal locomotor kinematics.

The Association Between Hip-Shoulder Separation Angles and Technique Characteristics in World-Class High Jumpers.

Hip-shoulder separation (H-Ssep) has been widely researched in many sporting activities (e.g., golf) to provide information on the contribution of torso rotation to performance and injury. Although it is necessary for high jumpers to generate significant long-axis rotation to successfully clear the bar, limited information exists on H-Ssep for high jump athletes. As such, this study aimed to (a) characterize the H-Ssep of world-class high jump athletes during competition, (b) determine if differences exist between male and female athletes and (c) to examine the relationship between H-Ssep and the biomechanical parameters used to describe high jump technique. Twenty-nine world-class high jumpers (17 males, 12 females) were video recorded (frame rate: 120–200 Hz) during the 2017 and 2018 World Athletics Championship finals. H-Ssep was quantified at touchdown (TD) and take-off (TO) following manual digitizing (SIMI motion) and a number of other common biomechanical parameters were computed. The observed levels of H-Ssep at TD (−46±12°) and TO (16 ±11°) were in line with those reported previously for other sports. The magnitude of H-Ssep varied between individuals and showed significant associations with other approach and take-off characteristics. Significant differences in H-Ssep were not evident between male and female athletes despite significant differences in other performance- and technique-related parameters. These findings highlight the divergent take-off characteristics of world-class performers and their reliance on hip-shoulder interactions when generating long axis rotation. Coaches should be mindful of the mechanical and physical consequences of H-Ssep when developing technical models, conditioning interventions and coaching strategies.

Evolutionary Innovations of the Feeding Apparatus in Herbivorous Dinosaurs

Although most studies in feeding biology of non‐avian dinosaurs have investigated feeding function and ecology in carnivorous dinosaurs, the complex diversity of feeding in herbivorous dinosaurs has recently been explored in much greater detail. Herbivorous non‐avian dinosaurs, composed of nearly all sauropodomorphs and ornithischians as well as a few theropod groups (i.e., ornithomimosaurs, therizinosaurs, and oviraptorosaurs), show a wide range of craniomandibular feeding mechanisms, each with unique adaptations for processing various forms of plant‐life. Here, I will review over a century of feeding studies in herbivorous dinosaurs, with particular focus on more recent advancements in our knowledge of their feeding evolution and ecology. These studies have primarily focused on the many components influencing the functional morphology of the head and neck of these animals, including: dental morphology, with studies in dental microwear and macrowear, overall morphological analysis of the skull and cervical vertebrae, reconstruction of cranial musculature and other soft tissues of the head, and biomechanics of the skull, with comparative analyses of mechanical advantage of the jaw system and 3D computer modeling in the form of finite element analyses and multibody dynamics analyses with the use of laser scans and CTs of dinosaur skulls. All these studies are instrumental in our understanding of the evolution and diversity of bite forces as well as feeding mechanisms that range from orthal to palinal feeding motions as well as bilateral hemi‐mandibular long‐axis rotation (especially in a variety of ornithischians). The similarities and differences between herbivorous dinosaurs and the herbivores of today (especially mammalian megaherbivores) show a great deal about how evolutionary opportunities and constraints have made a significant impact on the diverse nature of herbivory and its importance in ecosystems around the world.

Ankle structure of the Tokay gecko (Gekko gecko) and its role in the deployment of the subdigital adhesive system.

The remarkable ability of geckos to adhere to smooth surfaces is often thought of in terms of external structures, including the branching setae that make contact with the surface producing van der Waals forces. Some geckos also exhibit unique movements of the distal segments of the limbs during locomotion and static clinging, including active digital hyperextension and considerable pedal rotation. During static clinging, geckos can exhibit considerable adduction/abduction of the pes while the crus and thigh remain firmly adpressed to the substratum. This decoupling of pedal adduction/abduction from ankle flexion/extension and pedal long-axis rotation is a significant departure from pedal displacements of a typical lizard lacking adhesive ability. The structure of the ankle is likely key to this decoupling, although no detailed comparison of this complex joint between pad-bearing geckos and other lizards is available. Here we compare the configuration of the mesotarsal joint of nongekkotan lizards (Iguana and Pristidactylus) with that of the Tokay gecko (Gekko gecko) using prepared skeletons, scanning electron microscopy, and micro-computed tomographic (µCT) scans. We focus on the structure of the astragalocalcaneum and the fourth distal tarsal. The mesotarsal joint exhibits a suite of modifications that are likely associated with the secondarily symmetrical pes of pad-bearing geckos. For example, the lateral process of the astragalocalcaneum is much more extensive in G. gecko compared with other lizards. The mesotarsal joint exhibits several other differences permitting dissociation of long-axis rotation of the pes from flexion-extension movement, including a reduced ventral peg on the fourth distal tarsal, an articulatory pattern dominated by a well-defined, expansive distomesial notch of the astragalocalcaneum, and an associated broad proximodorsal articulatory facet of the fourth distal tarsal. Pad-bearing geckos are capable of effectively deploying their intricate adhesive system across a broad array of body angles because of this highly modified ankle. Future research should determine whether the differences encountered in G. gecko (and their extent) apply to the Gekkota as a whole and should examine how the elements of the ankle move dynamically during locomotion across a range of taxa.

Alligators employ intermetatarsal reconfiguration to modulate plantigrade ground contact.

ABSTRACTFeet must mediate substrate interactions across an animal's entire range of limb poses used in life. Metatarsals, the ‘bones of the sole’, are the dominant pedal skeletal elements for most tetrapods. In plantigrade species that walk on the entirety of their sole, such as living crocodylians, intermetatarsal mobility offers the potential for a continuum of reconfiguration within the foot itself. Alligator hindlimbs are capable of postural extremes from a belly sprawl to a high walk to sharp turns – how does the foot morphology dynamically accommodate these diverse demands? We implemented a hybrid combination of marker-based and markerless X-ray reconstruction of moving morphology (XROMM) to measure 3D metatarsal kinematics in three juvenile American alligators (Alligator mississippiensis) across their locomotor and maneuvering repertoire on a motorized treadmill and flat-surfaced arena. We found that alligators adaptively conformed their metatarsals to the ground, maintaining plantigrade contact throughout a spectrum of limb placements with non-planar feet. Deformation of the metatarsus as a whole occurred through variable abduction (twofold range of spread) and differential metatarsal pitching (45 deg arc of skew). Internally, metatarsals also underwent up to 65 deg of long-axis rotation. Such reorientation, which correlated with skew, was constrained by the overlapping arrangement of the obliquely expanded metatarsal bases. Such a proximally overlapping metatarsal morphology is shared by fossil archosaurs and archosaur relatives. In these extinct taxa, we suggest that intermetatarsal mobility likely played a significant role in maintaining ground contact across plantigrade postural extremes.

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.

Skeletal kinematics of the hyoid arch in the suction-feeding shark Chiloscyllium plagiosum.

White-spotted bamboo sharks, Chiloscyllium plagiosum, generate strong suction-feeding pressures that rival the highest levels measured in ray-finned fishes. However, the hyostylic jaw suspension of these sharks is fundamentally different from the actinopterygian mechanism, including more mobile hyomandibulae, with the jaws and ceratohyal suspended from the hyomandibulae. Prior studies have proposed skeletal kinematics during feeding in orectolobid sharks from indirect measurements. Here, we tested these hypotheses using XROMM to measure cartilage motions directly. In agreement with prior hypotheses, we found extremely large retraction and depression of the ceratohyal, facilitated by large protraction and depression of the hyomandibula. Somewhat unexpectedly, XROMM also showed tremendous long-axis rotation (LAR) of both the ceratohyal and hyomandibula. This LAR likely increases the range of motion for the hyoid arch by keeping the elements properly articulated through their large arcs of motion. XROMM also confirmed that upper jaw protraction occurs before peak gape, similarly to actinopterygian suction feeders, but different from most other sharks in which jaw protrusion serves primarily to close the mouth. Early jaw protraction results from decoupling the rotations of the hyomandibula, with much of protraction occurring before peak gape with the other rotations lagging behind. In addition, the magnitudes of retraction and protraction of the hyoid elements are independent of the magnitude of depression, varying the shape of the mouth among feeding strikes. Hence, the large variation in suction-feeding behavior and performance may contribute to the wide dietary breadth of bamboo sharks.

Rolling of the jaw is essential for mammalian chewing and tribosphenic molar function

Over the past two centuries, mammalian chewing and related anatomical features have been among the most discussed of all vertebrate evolutionary innovations1-3. Chief among these features are two characters: the dentary-only mandible, and the tribosphenic molar with its triangulated upper cusps and lower talonid basin3-5. The flexible mandibular joint and the unfused symphysis of ancestral mammals-in combination with transformations of the adductor musculature and palate-are thought to have permitted greater mobility of each lower jaw, or hemimandible6,7. Following the appearance of precise dental occlusion near the origin of the mammalian crown8,9, therians evolved a tribosphenic molar with a craggy topography that is presumed to have been used to catch, cut and crush food. Here we describe the ancestral tribosphenic therian chewing stroke, as conserved in the short-tailed opossum Monodelphis domestica: it is a simple symmetrical sequence of lower tooth-row eversion and inversion during jaw opening and closing, respectively, enacted by hemimandibular long-axis rotation. This sequence is coupled with an eversion-inversion rotational grinding stroke. We infer that the ancestral therian chewing stroke relied heavily on long-axis rotation, including symmetrical eversion and inversion (inherited from the first mammaliaforms) as well as a mortar-and-pestle rotational grinding stroke that was inherited from stem therians along with the tribosphenic molar. The yaw-dominated masticatory cycle of primates, ungulates and other bunodont therians is derived; it is necessitated by a secondarily fused jaw symphysis, and permitted by the reduction of high, interlocking cusps10-12. The development of an efficient masticatory system-culminating in the tribosphenic apparatus-allowed early mammals to begin the process of digestion by shearing and crushing food into small boli instead of swallowing larger pieces in the reptilian manner, which necessitates a long, slow and wholly chemical breakdown. The vast diversity of mammalian teeth has emerged from the basic tribosphenic groundplan13.

How moles destroy your lawn: the forelimb kinematics of Eastern moles in loose and compact substrates

The interplay between morphological specialization and kinematic flexibility is important for organisms that move between habitats within different substrates. Burrowing is energetically expensive and requires substantial interaction with soil to dislodge and transport it. True moles (Talpidae) have extraordinary forelimb morphologies and a unique ability to dig in loose as well as compact soils, yet we know little of how moles coordinate their forelimb joint kinematics when digging in soils of different compactness. Using marker-based X-ray Reconstruction of Moving Morphology (XROMM), we tested the hypothesis that moles burrow using different forelimb kinematics in loose and compact substrates. We predicted that moles raise mounds of loose soil by performing powerful compacting strokes mainly with long-axis rotation of the humerus (i.e. pronation/supination), but shear compact soil away by performing scratching strokes involving amplified elbow extension, similar to most scratching diggers. We also predicted that in both types of substrate, moles displace soil rearward like other mammalian diggers. Our results support our hypothesis but not the predictions. Eastern moles (Scalopus aquaticus) move substrates upward using compacting strokes in loose substrates and outward from the body midline using scratching strokes in compact substrates; unlike the digging strokes of most mammalian forelimb diggers, the power-stroke of moles itself does not displace substrates directly rearward. Compacting and scratching strokes involve similar ranges of humeral pronation and retraction at the scapulohumeral (shoulder) joint, yet the movements at the elbow and carpal joints differ. Our results demonstrate that the combination of stereotypic movements of the shoulder joint, where the largest digging muscles are located, and flexibility in the elbow and carpal joints makes moles extremely effective diggers in both loose and compact substrates.

Pectoral girdle and forelimb musculoskeletal function in the echidna ( Tachyglossus aculeatus ): insights into mammalian locomotor evolution

Although evolutionary transformation of the pectoral girdle and forelimb appears to have had a profound impact on mammalian locomotor and ecological diversity, both the sequence of anatomical changes and the functional implications remain unclear. Monotremes can provide insight into an important stage of this evolutionary transformation, due to their phylogenetic position as the sister-group to therian mammals and their mosaic of plesiomorphic and derived features. Here we build a musculoskeletal computer model of the echidna pectoral girdle and forelimb to estimate joint ranges of motion (ROM) and muscle moment arms (MMA)—two fundamental descriptors of biomechanical function. We find that the echidna's skeletal morphology restricts scapulocoracoid mobility and glenohumeral flexion–extension compared with therians. Estimated shoulder ROMs and MMAs for muscles crossing the shoulder indicate that morphology of the echidna pectoral girdle and forelimb is optimized for humeral adduction and internal rotation, consistent with limited in vivo data. Further, more muscles act to produce humeral long-axis rotation in the echidna compared to therians, as a consequence of differences in muscle geometry. Our musculoskeletal model allows correlation of anatomy and function, and can guide hypotheses regarding function in extinct taxa and the morphological and locomotor transformation leading to therian mammals.

Intermittent Treadmill Running Induces Kinematic Compensations to Maintain Soccer Kick Foot Speed Despite No Change in Knee Extensor Strength.

Kicking is a fundamental skill and a primary noncontact mechanism of injury in soccer, with injury incidence increasing during the latter stages of match-play. Ten male professional soccer players completed a 90-minute treadmill protocol based on the velocity profile of soccer match-play. Preexercise, and at 15-minute intervals, players completed a maximal velocity kick subjected to kinematic analysis at 200Hz. Preexercise, and at the end of each half, players also completed isokinetic concentric knee extensor repetitions at 180°·s-1, 300°·s-1, and 60°·s-1. Kicking foot speed was maintained at ∼19m·s-1, with no main effect for exercise duration. In relation to proximal-distal sequencing during the kicking action, there was a significant increase in the duration (but not magnitude) of thigh rotation, with a compensatory decrease in the duration (but not magnitude) of shank rotation during the latter stages of the exercise protocol. In relation to long-axis rotation, pelvic orientation at ball contact was maintained at ∼6°, representing a total pelvic rotation in the order of ∼15° during the kicking action. Peak knee extensor torque at all speeds was also maintained throughout the protocol, such that kinematic modifications are not attributable to a decline in knee extensor strength.

Forelimb muscle and joint actions in Archosauria: insights from Crocodylus johnstoni (Pseudosuchia) and Mussaurus patagonicus (Sauropodomorpha).

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 three-dimensional 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.

3-D range of motion envelopes reveal interacting degrees of freedom in avian hind limb joints.

Measuring range of motion (ROM) is a valuable technique that can link bone morphology to joint function in both extant and extinct taxa. ROM results are commonly presented as tables or graphs of maxima and minima for each rotational degree of freedom. We investigate the interactions among three degrees of freedom using X-ray reconstruction of moving morphology (XROMM) to measure ROM of the main hind limb joints of Helmeted Guineafowl (Numida meleagris). By plotting each rotation on an axis, we generate three-dimensional ROM volumes or envelopes composed of hundreds of extreme joint positions for the hip, knee, and intertarsal joints. We find that the shapes of ROM volumes can be quite complex, and that the contribution of long-axis rotation is often substantial. Plotting invivo poses from individual birds executing different behaviors shows varying use of potential rotational combinations within their ROM envelopes. XROMM can provide unprecedented high-resolution data on the spatial relationship of skeletal elements and thereby illuminate/elucidate the complex ways in which soft and hard tissues interact to produce functional joints. In joints with three rotational degrees of freedom, two-dimensional representations of ROM (flexion/extension and abduction/adduction) are incomplete.

An Investigation of the Locomotor Function of Therian Forearm Pronation Provides Renewed Support for an Arboreal, Chameleon-like Evolutionary Stage

Although investigations of forelimb characteristics are central to therian evolutionary studies, the functional origins of forearm pronation are neglected. However, recent research based on bipedal manipulations strongly suggests that proximal radioulnar joint mobility is highly conserved in tetrapods. This new information calls for a replication of previously published physical simulations of forearm bone movements, to investigate whether active therian pronation/supination evolved from the plesiomorphic mechanism via which locomotor-induced torsion is passively alleviated during forelimb retraction. Preliminary results using representative extant and extinct tetrapod forelimb elements are supportive, and also offer insight into why another overlooked forearm trait, osteological full pronation (mechanically aligned elbow and wrist/finger joints), evolved only in therians and chameleons. During forelimb retraction in tetrapods with unfused radii/ulnae, the radius unexpectedly remains fixed in place as a functional complex with the firmly planted manus/carpus, which the ulnar complex (ulna/humerus) displaces relative to. Therefore, the highly conserved functional morphology of the tetrapod forearm indicates that enhanced therian manual dexterity, which emphasizes isolated radial movements bipedally, was preceded by the locomotor evolution of ulnar supination relative to the radius quadrupedally. This counterintuitive information indicates that the traditional hypothesis, that therian pronation/supination evolved arboreally to amplify radial mobility, requires modification. The authors propose that proximal long-axis rotations of the therian ulnar complex co-evolved with osteological full pronation during a period of arboreal, chameleon-like locomotion, to continue allowing torsion at a reinforced proximal radioulnar joint. These adaptations were later or simultaneously co-opted for object manipulation using active radioulnar pronation/supination.

Forelimb Kinematics of Rats Using XROMM, with Implications for Small Eutherians and Their Fossil Relatives.

The earliest eutherian mammals were small-bodied locomotor generalists with a forelimb morphology that strongly resembles that of extant rats. Understanding the kinematics of the humerus, radius, and ulna of extant rats can inform and constrain hypotheses concerning typical posture and mobility in early eutherian forelimbs. The locomotion of Rattus norvegicus has been extensively studied, but the three-dimensional kinematics of the bones themselves remains under-explored. Here, for the first time, we use markerless XROMM (Scientific Rotoscoping) to explore the three-dimensional long bone movements in Rattus norvegicus during a normal, symmetrical gait (walking). Our data show a basic kinematic profile that agrees with previous studies on rats and other small therians: rats maintain a crouched forelimb posture throughout the step cycle, and the ulna is confined to flexion/extension in a parasagittal plane. However, our three-dimensional data illuminate long-axis rotation (LAR) movements for both the humerus and the radius for the first time. Medial LAR of the humerus throughout stance maintains an adducted elbow with a caudally-facing olecranon process, which in turn maintains a cranially-directed manus orientation (pronation). The radius also shows significant LAR correlated with manus pronation and supination. Moreover, we report that elbow flexion and manus orientation are correlated in R. norvegicus: as the elbow angle becomes more acute, manus supination increases. Our data also suggest that manus pronation and orientation in R. norvegicus rely on a divided system of labor between the ulna and radius. Given that the radius follows the flexion and extension trajectory of the ulna, it must rotate at the elbow (on the capitulum) so that during the stance phase its distal end lies medial to ulna, ensuring that the manus remains pronated while the forelimb is supporting the body. We suggest that forelimb posture and kinematics in Juramaia, Eomaia, and other basal eutherians were grossly similar to those of rats, and that humerus and radius LAR may have always played a significant role in forelimb and manus posture in small eutherian mammals.

Shoulder girdle rotation, forelimb movement and the influence of carapace shape on locomotion in Testudo hermanni (Testudinidae).

Studies into the function of structures are crucial for making connections between morphology and behaviour of organisms, but are still rare for the terrestrial Testudinidae. We investigated the kinematics of shoulder girdle and forelimb motion in Hermann's tortoise Testudo hermanni using biplanar X-ray fluoroscopy with a twofold aim: firstly, to understand how the derived shapes of shoulder girdle and carapace together influence rotation of the girdle; and, secondly, to understand how girdle rotation affects forelimb excursion. The total degree of shoulder rotation in the horizontal plane is similar to a species with a less domed shell, but because of the long and nearly vertically oriented scapular prong, shoulder girdle rotation contributes more than 30% to the horizontal arc of the humerus and nearly 40% to the rotational component of step length. The antebrachium and manus, which act as a functional unit, contribute roughly 50% to this component of the step length because of their large excursion almost parallel to the mid-sagittal plane. This large excursion is the result of the complex interplay between humerus long-axis rotation, counter-rotation of the antebrachium, and elbow flexion and extension. A significant proportion of forelimb step length results from body translation that is due to the propulsive effect of the other limbs during their stance phases. Traits that are similar to other tortoises and terrestrial or semi-aquatic turtles are the overall slow walk because of a low stride frequency, and the lateral-sequence, diagonally coupled footfall pattern with high duty factors. Intraspecific variation of carapace shape and shoulder girdle dimensions has a corresponding effect on forelimb kinematics.

Guineafowl with a twist: asymmetric limb control in steady bipedal locomotion.

In avian bipeds performing steady locomotion, right and left limbs are typically assumed to act out of phase, but with little kinematic disparity. However, outwardly appearing steadiness may harbor previously unrecognized asymmetries. Here, we present marker-based XROMM data showing that guineafowl on a treadmill routinely yaw away from their direction of travel using asymmetrical limb kinematics. Variation is most strongly reflected at the hip joints, where patterns of femoral long-axis rotation closely correlate to degree of yaw divergence. As yaw deviations increase, hip long-axis rotation angles undergo larger excursions and shift from biphasic to monophasic patterns. At large yaw angles, the alternately striding limbs exhibit synchronous external and internal femoral rotations of substantial magnitude. Hip coordination patterns resembling those used during sidestep maneuvers allow birds to asymmetrically modulate their mediolateral limb trajectories and thereby advance using a range of body orientations.