The functional morphology of the cercopithecoid wrist and inferior radioulnar joints, and their bearing on some problems in the evolution of the Hominoidea.

The cercopithecoid wrist joint differs from the wrist joints of hominoids in several ways. The distal ulna, the distal radius, the pisiform, the triquetrum, the hamate, and the base of the fifth metacarpal are on the one hand remarkably alike among cercopithecoid genera, and on the other remarkably distinct from homologous bones in the Hominoidea. Functionally, the triquetrum and the pisiform, in conjuction with the ulnar styloid process, check the proximal carpal row during ulnar deviation, and are possibly important in stabilizing the wrist during dorsiflexion as well. The head of the ulna almost certainly betokens a range of radioulnar supination in cercopithecoids that is substantially less than is to be found in any of the hominoid genera. The articulation between the hamate bone and the base of the fifth metacarpal allows for considerable dorsiflexion in the Cercopithecoidea; this potential was not evidenct in any of the hominoids examined. Behaviorally, the cercopithecoid wrist can most profitably be viewed as an adaptation for a quadrupedal life style involving dorsiflexion of the wrist and palmigrade/digitigrade substrate contact. The hominoid wrist joint is not adapted for such a behavioral potential.     

Carpal bone kinematics and ligament lengthening studied for the full range of joint movement

Present data on carpal kinematics and carpal ligament behaviour are limited to flexion and deviation movements of the hand. These motions do not represent all the wrist-joint motions which are important for the activities of daily living. The goal of this project was to obtain insight into carpal kinematics and carpal ligament behaviour during motions of the hand covering the full range of motion of the wrist joint.

The carpals and the ligaments of four wrist-joint specimens were provided with radiopaque markers. These joints were subjected to Röntgenstereophotogrammetric experimentation in a large number of hand positions to determine carpal positions and ligament lengths. The movements of the carpal bones were described by means of finite helical axes (FHA).

It was found that the movements of the carpals in the distal row closely resemble those of the hand. Conversely, the motions of the carpals of the proximal row appeared not to be directly proportional to the hand motions and exhibited clear out-of-plane movements. Furthermore, it could be shown that movements of the hand into the ulnodorsal quadrant of the full range of hand motion corresponds to larger helical rotations and translations for most of the carpals than when the hand was moved into any other quadrant. The maximal ligament length changes determined did not exceed the length changes reported for pure flexion and pure deviation movements of the hand.     

Vascularized fibular graft after excision of giant-cell tumor of the distal radius: wrist arthroplasty versus partial wrist arthrodesis

Several reconstructive procedures have been described for the complete defect of the distal radius that is created after a wide excision of a giant-cell tumor of bone, including hemiarthroplasty using the vascularized fibular head and partial wrist arthrodesis between a vascularized fibula and the scapholunate portion of the proximal carpal row. The objectives of this study are to compare clinical and radiographic results between the partial wrist arthrodesis and the wrist arthroplasty, and to discuss which procedure is superior. Four patients with giant-cell tumors involving the distal end of the radius were treated with en bloc resection and reconstruction with a free vascularized fibular graft. The wrists in two patients were reconstructed with an articular fibular head graft and the remaining two patients underwent partial wrist arthrodesis using a fibular shaft transfer. There was radiographic evidence of bone union at the host-graft junctions in all cases. In the newly reconstructed wrist joint, there was palmar subluxation of the carpal bones and degenerative changes in both patients. Local recurrence was seen in one patient. According to the functional results described by Enneking et al., the mean functional score was 67 percent. The functional scores including wrist/forearm range of motion in the cases with partial wrist arthrodesis were superior to those with wrist arthroplasty. A partial wrist arthrodesis using a vascularized fibular shaft graft appears a more useful and reliable procedure for reconstruction of the wrist after excision of the giant-cell tumor of the distal end of the radius than a wrist arthroplasty using the vascularized fibular head, although our study includes only a small number of patients.     

The effect of tendon loading on in-vitro carpal kinematics of the wrist joint

Measurements of in-vitro carpal kinematics of the wrist provide valuable biomechanical data. Tendon loading is often applied during cadaver experiments to simulate natural stabilizing joint compression in the wrist joint. The purpose of this study was to investigate the effect of tendon loading on carpal kinematics in-vitro.A cyclic movement was imposed on 7 cadaveric forearms while the carpal kinematics were acquired by a 4-dimensional rotational X-ray imaging system. The extensor- and flexor tendons were loaded with constant force springs of 50 N, respectively. The measurements were repeated without a load on the tendons. The effect of loading on the kinematics was tested statistically by using a linear mixed model.During flexion and extension, the proximal carpal bones were more extended with tendon loading. The lunate was on the average 2.0° (p=0.012) more extended. With tendon loading the distal carpal bones were more ulnary deviated at each angle of wrist motion. The capitate was on the average 2.4° (p=0.004) more ulnary deviated.During radioulnar deviation, the proximal carpal bones were more radially deviated with the lunate 0.7° more into radial deviation with tendon loading (p<0.001). Conversely, the bones of distal row were more flexed and supinated with the capitate 1.5° more into flexion (p=0.025) and 1.0° more into supination (p=0.011).In conclusion, the application of a constant load onto the flexor and extensor tendons in cadaver experiments has a small but statistically significant effect on the carpal kinematics during flexion–extension and radioulnar deviation.     

Clinical anatomy of the m. flexor carpi radialis tendon sheath

At the transitional zone from the forearm to the hand the insertion tendon of the m.flexor carpi radialis (FCR) glides on a fibrous and fatty cushion, which is connected dorsally with the joint capsule of the radiocarpal articulation. The tendon distally crosses the palmar side of the scaphoid tubercle and enters the dorsally curved rim of the trapezoid tubercle. At the level of the wrist joint the narrow tendon sheath begins, which extends to the insertion at the metacarpus. Immediately after entering the gliding tunnel the tendon branches off radially as a rule with an accessory fibre strand 8 mm in width to the scaphoid, trapezium and the joint capsule between these two bones. The insertion tendon regularly is attached to the palmar and radial surfaces of the second and third metacarpal bones. The wall of the osteofibrous gliding tunnel can be prominent following trauma, inflammation or arthrosis deformans in the trapezio-scaphoideal joint and may irritate the tendon (tendovaginosis stenosans). Against resistance forces pain will occur in the wrist joint during palmar flexion. The typical point of tenderness is situated at the entering of the tendon in the thenar region. Operative decompression will be effective by opening the radial wall of the tendon sheath from the carpal tunnel.     

Simulation of extension,radial and ulnar deviation of the wrist with a rigid body spring model

A novel computational model of the wrist that predicts carpal bone motion was developed in order to investigate the complex kinematics of the human wrist.This rigid body spring model (RBSM) of the wrist was built using surface models of the eight carpal bones, the bases of the five metacarpal bones, and the distal parts of the ulna and radius, all obtained from computed tomography (CT) scans of a cadaver upper limb. Elastic contact conditions between the rigid bodies modeled the influence of the cartilage layers, and ligamentous structures were constructed using nonlinear, tension-only spring elements. Motion of the wrist was simulated by applying forces to the tendons of the five main wrist muscles modeled.Three wrist motions were simulated: extension, ulnar deviation and radial deviation. The model was tested and tuned by comparing the simulated displacement and orientation of the carpal bones with previously obtained CT-scans of the same cadaver arm in deviated (45°ulnar and 15°radial), and extended (57°) wrist positions. Simulation results for the scaphoid, lunate, capitate, hamate and triquetrum are presented here and provide credible prediction of carpal bone movement. These are the first reported results of such a model. They indicate promise that this model will assist in future wrist kinematics investigations. However, further optimization and validation are required to define and guarantee the validity of results.     

Resting Orientations of Dinosaur Scapulae and Forelimbs: A Numerical Analysis,with Implications for Reconstructions and Museum Mounts

The inclination of the scapular blade and the resting pose of the forelimb in dinosaurs differ among reconstructions and among skeletal mounts. For most dinosaurian taxa, no attempt has previously been made to quantify the correct resting positions of these elements. Here, we used data from skeletons preserved in articulation to quantify the resting orientations of the scapula and forelimb in dinosaurs. Specimens were included in the study only if they were preserved lying on their sides; for each specimen the angle between forelimb bones at a given joint was included in the analysis only if the joint was preserved in articulation. Using correlation analyses of the angles between the long axis of the sacrum, the first dorsal centrum, and the scapular blade in theropods and Eoraptor, we found that vertebral hyperextension does not influence scapular orientation in saurischians. Among examined taxa, the long axis of the scapular blade was found to be most horizontal in bipedal saurischians, most vertical in basal ornithopods, and intermediate in hadrosauroids. We found that in bipedal dinosaurs other than theropods with semilunate carpals, the resting orientation of the elbow is close to a right angle and the resting orientation of the wrist is such that the hand exhibits only slight ulnar deviation from the antebrachium. In theropods with semilunate carpals the elbow and wrist are more flexed at rest, with the elbow at a strongly acute angle and with the wrist approximately at a right angle. The results of our study have important implications for correct orientations of bones in reconstructions and skeletal mounts. Here, we provide recommendations on bone orientations based on our results.     

A novel method to replicate the kinematics of the carpus using a six degree-of-freedom robot

Understanding the kinematics of the carpus is essential to the understanding and treatment of wrist pathologies. However, many of the previous techniques presented are limited by non-functional motion or the interpolation of points from static images at different postures. We present a method that has the capability of replicating the kinematics of the wrist during activities of daily living using a unique mechanical testing system. To quantify the kinematics of the carpal bones, we used bone pin-mounted markers and optical motion capture methods. In this paper, we present a hammering motion as an example of an activity of daily living. However, the method can be applied to a wide variety of movements. Our method showed good accuracy (1.0–2.6°) of in vivo movement reproduction in our ex vivo model. Most carpal motion during wrist flexion–extension occurs at the radiocarpal level while in ulnar deviation the motion is more equally shared between radiocarpal and midcarpal joints, and in radial deviation the motion happens mainly at the midcarpal joint. For all rotations, there was more rotation of the midcarpal row relative to the lunate than relative to the scaphoid or triquetrum. For the functional motion studied (hammering), there was more midcarpal motion in wrist extension compared to pure wrist extension while radioulnar deviation patterns were similar to those observed in pure wrist radioulnar deviation. Finally, it was found that for the amplitudes studied the amount of carpal rotations was proportional to global wrist rotations.     

The lorisiform wrist joint and the evolution of "brachiating" adaptations in the hominoidea.

In lorisines (Loris, Nycticebus, Perodicticus, Arctocebus), the tip of the ulna is reduced to the dimensions of a styloid process, a new and more proximal ulnar head is developed, and the pisiform is displaced distally away from its primitive contact with the ulna. In some Nycticebus, intra-articular tissues separate the ulna from the triquetrum. These traits are not seen in other quadrupedal primates, but they are characteristic of extant hominoids. Among hominoids, these features have been interpreted as adaptations to arm-swinging locomotion. Since hominoid-like features of the wrist joint are found in lorisines, but not in New World monkeys that practice arm-swinging locomotion, these features may have been evolved in both lorisines and large hominoids to enhance wrist mobility for cautious arboreal locomotion involving little or no leaping. Most of the other morphological traits characteristic of modern hominoids can be explained as adaptations to cautious quadrupedalism as well as to brachiation, and may have developed for different reasons in different lineages descended from an unspecialized cautious quadruped resembling Alouatta.     

Functional osteology of the avian wrist and the evolution of flapping flight

The avian wrist is extraordinarily adapted for flight. Its intricate osteology is constructed to perform four very different, but extremely important, flight-related functions. (1) Throughout the downstroke, the cuneiform transmits force from the carpometacarpus to the ulna and prevents the manus from hyperpronating. (2) While gliding or maneuvering, the scapholunar interlocks with the carpometacarpus and prevents the manus from supinating. By employing both carpal bones simultaneously birds can lock the manus into place during flight. (3) Throughout the downstroke-upstroke transition, the articular ridge on the distal extremity of the ulna, in conjuction with the cuneiform, guides the manus from the plane of the wing toward the body. (4) During take-off or landing, the upstroke of some heavy birds exhibits a pronounced flick of the manus. The backward component of this flick is produced by reversing the wrist mechanism that enables the manus to rotate toward the body during the early upstroke. The upward component of the flick is generated by mechanical interplay between the ventral ramus of the cuneiform, the ventral ridge of the carpometacarpus, and the ulnocarpo-metacarpal ligament. Without the highly specialized osteology of the wrist it is doubtful that birds would be able to carry out successfully the wing motions associated with flapping flight. Yet in Archaeopteryx, the wrist displays a very different morphology that lacks all the key features found in the modern avian wrist. Therefore, Archaeopteryx was probably incapable of executing the kinematics of modern avian powered flight.     

A 3D quantitative comparison of trapezium and trapezoid relative articular and nonarticular surface areas in modern humans and great apes

The structure and functions of the modern human hand are critical components of what distinguishes Homo sapiens from the great apes (Gorilla, Pan, and Pongo). In this study, attention is focused on the trapezium and trapezoid, the two most lateral bones of the distal carpal row, in the four extant hominid genera, representing the first time they have been quantified and analyzed together as a morphological-functional complex. Our objective is to quantify the relative articular and nonarticular surface areas of these two bones and to test whether modern humans exhibit significant shape differences from the great apes, as predicted by previous qualitative analyses and the functional demands of differing manipulative and locomotor strategies. Modern humans were predicted to show larger relative first metacarpal and scaphoid surfaces on the trapezium because of the regular recruitment of the thumb during manipulative behaviors; alternatively, great apes were predicted to show larger relative second metacarpal and scaphoid surfaces on the trapezoid because of the functional demands on the hands during locomotor behaviors. Modern humans were also expected to exhibit larger relative mutual joint surfaces between the trapezoid and adjacent carpals than do the great apes because of assumed transverse loads generated by the functional demands of the modern human power grip. Using 3D bone models acquired through laser digitizing, the relative articular and nonarticular areas on each bone are quantified and compared. Multivariate analyses of these data clearly distinguish modern humans from the great apes. In total, the observed differences between modern humans and the great apes support morphological predictions based on the fact that this region of the human wrist is no longer involved in weight-bearing during locomotor behavior and is instead recruited solely for manipulative behaviors. The results provide the beginnings of a 3D comparative standard against which further extant and fossil primate wrist bones can be compared within the contexts of manipulative and locomotor behaviors.     

The mole's thumb -- evolution of the hand skeleton in talpids (Mammalia)

Osteological specimens representing 15 out of the 16 currently recognized talpid genera were examined and scored for seven discrete morphological characters of the hand. The phylogenetic distribution of these characters was studied in the context of alternative hypotheses of talpid relationships, using three species of shrews and a hedgehog as outgroups. All talpids show a similar number and arrangement of carpal bones. The most obvious differences concern the presence of additional sesamoid bones, the relative size of the os falciforme when present, and the degree of fusion of the scaphoid and lunate in the proximal carpal row. Marked differences in the relative length and proportions of the metacarpals also exist. The development of the carpals in Talpa europaea was studied through examination of histological sections of the hand of an embryo and a neonate. Whereas carpal anatomy in the neonate mirrors the arrangement and proportions of the adult, in the embryo the scaphoid and lunate are still separate, there are no signs of the os falciforme, and the size proportions of metacarpals to carpals are obviously different to those of the adult. A prehallux or tibial sesamoid, serial homologue to the os falciforme or prepollex (a radial sesamoid), does not have an obvious functional role, and its presence might be the result of a common epigenetic control in the hand and the foot resulting in a non-adaptive structure in the latter.     

Carpal ontogeny in Monodelphis domestica and Caluromys philander (Marsupialia)

Carpal bones have experienced numerous changes during marsupial evolution, even though their diversity and development remain poorly studied. The aim of this work was to document adult form and the pattern of mesenchymal tissue condensation and formation of chondrification and ossification centers in the hand of two marsupials. Two fundamental questions were asked: whether the loss of embryonic precursors was associated with the loss of adult elements, or whether there were developmental signs of ancestral mammalian elements that have been fused or lost in marsupial taxa. We were also interested to find out whether there is sexual dimorphismus in the carpals, as has been reported for some didelphids. Histological sections, cleared and stained specimens and macerated skeletons representing an ontogenetic series of Monodelphis domestica were used to document carpal development. Comparisons were made with perinatal stages of Caluromys philander and with adult specimens of other marsupials. A prenatal M. domestica in the 13th day after conception has a cell condensation that because of its position is homologized with a centrale, which is at birth already lost or fused. Neonatal M. domestica and C. philander have the number and arrangement of their adult carpal anatomy. Trapezium and trapezoid start ossification later than most other carpals, while pisiform and prepollex are the last to do so. Adult males of M. domestica have relatively larger and more robust pisiforms, compared to other carpals, than females. This sexual dimorphism develops relatively late as it was not recorded in male specimens around 160 days old. An extra sesamoid bone located just distal to the radius and proximo-palmar to the scaphoid was recorded in specimens of C. philander, C. derbianus and Didelphis virginiana.     

New Developmental Evidence Clarifies the Evolution of Wrist Bones in the Dinosaur–Bird Transition

From early dinosaurs with as many as nine wrist bones, modern birds evolved to develop only four ossifications. Their identity is uncertain, with different labels used in palaeontology and developmental biology. We examined embryos of several species and studied chicken embryos in detail through a new technique allowing whole-mount immunofluorescence of the embryonic cartilaginous skeleton. Beyond previous controversy, we establish that the proximal–anterior ossification develops from a composite radiale+intermedium cartilage, consistent with fusion of radiale and intermedium observed in some theropod dinosaurs. Despite previous claims that the development of the distal–anterior ossification does not support the dinosaur–bird link, we found its embryonic precursor shows two distinct regions of both collagen type II and collagen type IX expression, resembling the composite semilunate bone of bird-like dinosaurs (distal carpal 1+distal carpal 2). The distal–posterior ossification develops from a cartilage referred to as “element x,” but its position corresponds to distal carpal 3. The proximal–posterior ossification is perhaps most controversial: It is labelled as the ulnare in palaeontology, but we confirm the embryonic ulnare is lost during development. Re-examination of the fossil evidence reveals the ulnare was actually absent in bird-like dinosaurs. We confirm the proximal–posterior bone is a pisiform in terms of embryonic position and its development as a sesamoid associated to a tendon. However, the pisiform is absent in bird-like dinosaurs, which are known from several articulated specimens. The combined data provide compelling evidence of a remarkable evolutionary reversal: A large, ossified pisiform re-evolved in the lineage leading to birds, after a period in which it was either absent, nonossified, or very small, consistently escaping fossil preservation. The bird wrist provides a modern example of how developmental and paleontological data illuminate each other. Based on all available data, we introduce a new nomenclature for bird wrist ossifications.     

Does gliding when pregnant select for larger females?

For terrestrial vertebrates, gliding imposes unique constraints on the interaction of body mass and structural size, particularly with reference to minimizing wing loading. Females of gliding animals experience increases in wing loading during pregnancy or gravidity, and selection may favour increased structural size to compensate for the added mass. We tested whether pregnant southern flying squirrels Glaucomys volans had similar wing loading as males, and whether females with lower wing loading bore heavier litters, than those with greater wing loading. Males had greater wing loading than females, regardless of the latter's reproductive state (males: 38.4±3.62 N m⁻², pregnant females: 30.7±4.21 N m⁻² and non-pregnant females: 26.8±5.13 N m⁻²). The slope of the linear relationship between planar surface area and body mass was similar between pregnant females and males, however ( F =0.383, P =0.322). Thus female flying squirrels may optimize their litter mass to minimize wing loading during pregnancy. Contrary to our prediction, females with greater wing loading had heavier litters than those with lower wing loading, which suggests reproductive output may be influenced by other ecological factors.     

The attachments of the lateral and medial ends of the extensor retinaculum of the human wrist

The attachments of the lateral and medial ends of the extensor retinaculum of the human wrist were observed in 52 human upper limbs (from 26 cadavers) by dissection. In all the specimens used in this study, the lateral end of the retinaculum was found to be attached to the distal part of the anterior border of the radius and its medial end was attached to the styloid process of the ulna, the pisiform and the triquetrum.     

The asymmetry of the carpal joint and the evolution of wing folding in maniraptoran theropod dinosaurs

In extant birds, the hand is permanently abducted towards the ulna, and the wrist joint can bend extensively in this direction to fold the wing when not in use. Anatomically, this asymmetric mobility of the wrist results from the wedge-like shape of one carpal bone, the radiale, and from the well-developed convexity of the trochlea at the proximal end of the carpometacarpus. Among the theropod precursors of birds, a strongly convex trochlea is characteristic of Coelurosauria, a clade including the highly derived Maniraptora in addition to tyrannosaurs and compsognathids. The shape of the radiale can be quantified using a ‘radiale angle’ between the proximal and distal articular surfaces. Measurement of the radiale angle and reconstruction of ancestral states using squared-change parsimony shows that the angle was small (15°) in primitive coelurosaurs but considerably larger (25°) in primitive maniraptorans, indicating that the radiale was more wedge-shaped and the carpal joint more asymmetric. The radiale angle progressively increased still further within Maniraptora, with concurrent elongation of the forelimb feathers and the forelimb itself. Carpal asymmetry would have permitted avian-like folding of the forelimb in order to protect the plumage, an early advantage of the flexible, asymmetric wrist inherited by birds.     

Locomotor adaptations reflected in the wrist joints of early tertiary primates (adapiformes)

The positional behaviors inferred for early Tertiary adapiform primates have been the subject of considerable debate. Adapiform wrist morphology is analyzed here within the context of extant morphoclines in carpal joint shape in order to reconstruct adapiform positional behavior. Extant vertical clingers, slow climbers, and arboreal quadrupeds differ significantly from one another in length of the m. flexor carpi ulnaris lever arm, shape of the midcarpal joint articular surface, and size and divergence of the pollical carpometacarpal articulation. These morphological differences are functionally related to differential requirements for wrist flexion, midcarpal mobility and stability, and pollical grasping, respectively. Adapis, Notharctus, and Smilodectes share with living arboreal quadrupeds a tall pisiform body, a mediolaterally flat midcarpal joint surface, and a relatively unexpanded thumb joint. Functionally, these features are related to flexing the wrist from extended positions during palmigrade, quadrupedal locomotion, increasing midcarpal joint stability during quadrupedal, weight-bearing postures, and grasping arboreal supports of predominantly horizontal and oblique orientation. The Messel adapiform (genus indet.) shares certain features of the midcarpal and pollical carpometacarpal articulations with extant vertical clingers, suggesting that this taxon used vertical substrates more frequently than other adapiforms. © 1996 Wiley-Liss, Inc.     

The Evolution and Paleobiogeography of Flying Squirrels (Sciuridae, Pteromyini) in Response to Global Environmental Change

Flying squirrels are strictly arboreal squirrels adopting a special gliding form of locomotion. This group of animals has a long history that has mirrored the vicissitude of forests. The discrepancy in the distribution between fossils and extant species indicates a mysterious evolution history requiring further exploration. This study compiles the worldwide fossils of Pteromyini to the species level in order to reproduce the spatiotemporal distribution pattern of flying squirrels and deduce the ancestral distribution according to dispersal-vicariance analysis of a phylogeny of the extant species. In addition, we reconstruct the paleoenvironmental background and find that flying squirrels probably originated in the Oligocene–Miocene transition from Europe and immediately dispersed to Asia and North America. Influenced by glaciation, CO₂ reduction, geologic movements and the Paratethys retreat, the Northern Hemisphere underwent climate deterioration and grassland expansion during the late Miocene, and thus the diversity of Pteromyini dramatically decreased. The uplift of the Tibet Plateau in addition to the strengthened Asian monsoons intensified the aridity in central Asia, but brought sufficient water to the densely forested regions of South and Southeast Asia. These forests are likely both refugia and diversification center for flying squirrels during glacial periods in the Quarternary. The subsequent connection and separation events among these heterogeneous habitats has probably been a driving force in the speciation of flying squirrels. Based on this work, we predict a bleak future for the flying squirrels, one which is closely associated with the fate of forests in Asia.