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.     

A partial skeleton of Geotrypus antiquus (Talpidae, Mammalia) from the Late Oligocene of the Enspel fossillagerst?tte in Germany

The partial skeleton of a young adult Geotrypus antiquus (de Blainville 1840) from the Upper Oligocene (MP 28) found in Enspel comprises the skull with both mandibles, distal ends of both scapulae, left clavicula, humeri, ulnae and radii of both sides, various elements of the hand, some vertebrae, ribs, and the left femur. For the first time, the previously postulated association between dentition and postcranial elements can be confirmed. The skeleton exhibits strong adaptations for a subterranean life, similar to modern fossorial moles. The humerus is wide with a large pectoral process. The wing-like greater and lesser tuberosities, teres tubercle, and distal epicondylus are clearly developed. The metacarpals and phalanges are broad and stout. There are several sesamoid bones in the broad digging hand, including a prepollex (os falciforme). The preserved bones allowed the forelimb of G. antiquus to be reconstructed. Previous finds of G. antiquus have mainly been from France, with a few specimens from Switzerland and southern Germany. The specimen from Enspel is the northernmost record. A cladistic analysis, based on the matrix of Sánchez-Villagra et al. (Cladistics 22:59?C88, 2006), confirms the basal position of Geotrypus within the Old World moles (Talpini).     

Isolated absence of human carpal bones.

An unusual instance of congenital isolated absence of carpal bones, in a 38-year-old woman, characterized by total lack of the scaphoid, lunate, and pyramidal and a massive fusion of the distal carpal row is reported. This condition is thought to have been the result of an embryonic error exclusively involving the portion of the autopodic blastema from which the carpus originates.     

Biometric study of ruminant carpal bones and implications for phylogenetic relationships

The mammalian Carpus is a complex of several small bones with multiple interactions during walking. Therefore, it is highly probable that different mammalian families developed distinctive constellations in their Carpi, which could be useful for biometric identification of phylogenetic groupings. The carpal bones of nineteen extant ruminant species (nine bovid, nine cervid, and one moschid) have been investigated to search for biometric traits which are diagnostic for the three families. Additionally, we searched for diverging functional adaptations in the carpal constellations. Therefore, measurements have been taken from the five main carpals, which are carrying the body weight. As a sesamoid bone, Os carpi accessorium was excluded. After transformation of the data into their natural logarithms, multivariate methods of factor analyses and discriminant analyses were performed for each bone. Bivariate plots of the factor scores allowed a clear separation of bovids and cervids. The only one species of the Moschidae (Moschus moschiferus) lies closer to the cervids than to the bovids. The grouping is due to phylogenetic relationships and not due to functional differences in the groups or differing habitat preferences. Generally, the carpals of cervids are more slender and higher in contrast to the bulky and flat carpals in bovids. This approach could be used to assign isolated carpal bones found in fossil sites to their ruminant family.     

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.     

New hand bones of Hadropithecus stenognathus: implications for the paleobiology of the Archaeolemuridae

A partial, associated skeleton of Hadropithecus stenognathus (AHA-I) was discovered in 2003 at Andrahomana Cave in southeastern Madagascar. Among the postcranial elements found were the first hand bones (right scaphoid, right hamate, left first metacarpal, and right and left fifth metacarpals) attributed to this rare subfossil lemur. These hand bones were compared to those of extant strepsirrhines and catarrhines in order to infer the positional adaptations of Hadropithecus, and they were compared to those of Archaeolemur in order to assess variation in hand morphology among archaeolemurids. The scaphoid tubercle does not project palmarly as in suspensory and climbing taxa, and the hamate has no hook at all (just a small tubercle), which also points to a poorly developed carpal tunnel. There is a distinctive, radioulnarly directed "spiral" facet for articulation with the triquetrum that is most similar in orientation to that of more terrestrial primates (i.e., Lemur catta, Papio, and Gorilla). The first metacarpal is very reduced and represents only 48% of the length of metacarpal V, as in Archaeolemur, which suggests that pollical grasping of arboreal supports was not important. Compared to Archaeolemur, the shaft of metacarpal V is gracile, and the head has no dorsal ridge and lacks characteristics functionally associated with digitigrade, extended metacarpophalangeal joint postures. Proximally, the articular facet for the hamate is oriented more dorsally. Thus, the carpometacarpal joint V appears to have a distinctive hyperextended set, which has no analog among living or extinct primates. The carpals of Hadropithecus are diagnostic of a pronograde, arboreal and terrestrial (although not digitigrade) locomotor repertoire that typifies Lemur catta and some Old World monkeys. No clinging, suspensory, or climbing specializations that characterize indriids or lorises can be found in the hand of this subfossil lemur. The hand of Hadropithecus likely had similar ranges of movement at the radiocarpal and midcarpal joints as of those of pronograde primates, such as lemurids, for which the hand is held in a more extended, pronated, and neutral (i.e., showing less ulnar deviation) position during locomotion in comparison to that of vertical clingers or slow climbers. Although highly autapomorphic, the hand of Hadropithecus resembles that of its sister taxon, Archaeolemur, in having a very reduced pollex and an articular facet on the scaphoid for a sizeable prepollex. These unusual hand features reinforce the monophyly of the Archaeolemuridae.     

Applications of radiodermatography to human and primate studies

Using 325-mesh tantalum powder as a contrast medium, it is possible to take high-resolution “radiodermatographs” of man and infra-human primates that combine the advantages of postero-anterior hand radiographs and conventional paper palm-and-finger prints. The radiodermatographs are particularly useful in relating creases, triradii and fingertip ridge patterns to differences in the proportions of the carpals, metacarpals and phalanges.     

Did knuckle walking evolve twice?

Although African great apes share a similar quadrupedal locomotor behaviour, there are marked differences in hand morphology and size between the species. Hence, whilst all three species (two genera) of African ape frequently knuckle walk as adults, debate remains as to whether this behaviour is derived from a common ancestor or whether it evolved in parallel in chimpanzees and gorillas. This exploratory morphometric study of the sub-adult and adult wrist of these two genera aims to contribute to this debate. A total of twenty-seven dimensions of the lunate, triquetral, hamate and capitate of sub-adult and adult Pan troglodytes and Gorilla gorilla were analysed in order to determine whether carpal dimensions are generally ontogenetically scaled, and whether differences in growth trajectories, or length of growth, and adult morphologies can be explained by behavioural differences between the two species. Only 56% of all dimensions studied were ontogenetically scaled in sub-adults and some of these dimensions exhibit differing adult proportions between the two species. In general, the dimensions analysed fell into two categories: Pan and Gorilla either follow the same growth trajectories (Pattern A) or the Pan reduced major axis (RMA) regressions were significantly transposed above those of Gorilla (Pattern B). Additionally, it was found that Gorilla carpals appear to cease growing relatively earlier than those of Pan. While a small number of differences, notably those of the lunate, can be accounted for by differences in behaviour between the species, the majority of differences indicate heterochronic modifications of development during evolution, which correspond to kinematic differences in knuckle walking between the African great apes. In light of morphological, behavioural and ecological data currently available it is parsimonious to suggest that knuckle walking has evolved in parallel in the two lineages.     

Dysharmonic maturation of the hand in the congenital malformation syndromes

In many congenital malformation syndromes the pattern of hand-wrist development does not fit the sequence pictured in the Greulich-Pyle atlas. Not infrequently, there is a difference in maturation level of carpal and phalangeal centers in excess of that found in clinically normal children. Usually, the carpal centers are less developed than the phalangeal centers, whereas overall skeletal maturation is retarded (as in trisomy 18) or advanced (as in cerebral gigantism). In still other conditions, specific carpal centers are disproportionately delayed. By way of example, the capitate is differentially delayed in epiphyseal dysplasia, the lunate in homocystinuria, and the scaphoid in Fanconi's anemia and other radial hypoplasia syndromes. Side to side (i.e., bilateral) asymmetries may also occur in the developing hand, as in paralysis, in conditions involving increased local vascularity (as in hemangioma and rheumatoid arthritis) or in conditions associated with decreased vascularity. In the presence of excessive dysharmonic development or major bilateral asymmetry, with or without agenesis of one or more hand bones, assigning meaningful bone ages in congenital malformation syndromes becomes difficult. On the other hand, the degree and pattern of dysharmonic maturation may be helpful in diagnosis.     

Rapid prototyping of scaphoid and lunate bones

In this study, a novel rapid prototyping technology was used to fabricate scaphoid and lunate bone prostheses, two carpal bones that are prone to avascular necrosis. Carpal prostheses were fabricated with an Envisiontec Perfactory® SXGA stereolithography system using Envisiontec eShell 200 photocurable polymer. Fabrication was guided using 3-D models, which were generated using Mimics software (Materialise NV, Leuven, Belgium) from patient computer tomography data. The prostheses were fabricated in a layer-by-layer manner; ∼ 50-μm thick layers were observed in the prostheses. Hardness and Young's modulus values of polymerized eShell 200 material were 93.8 ± 7.25 MPa and 3050 ± 90 MPa, respectively. The minimum compressive force required for fracture was 1360 N for the scaphoid prosthesis and 1248 N for the lunate prosthesis. Polymerized Envisiontec eShell material exhibited high human neonatal epidermal keratinocyte cell viability rate in an MTT assay. The results of this study indicate that small bone prostheses fabricated by stereolithography using eShell 200 polymer may have suitable geometry, mechanical properties, and cytocompatibility properties for in vivo use.     

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.     

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 dorsal arterial network of the wrist with reference to the blood supply of the carpal bones.

The gross examination of 50 minium injected specimens showed us the various patterns of the posterior arterial network of the wrist. This is chiefly provided by the radial artery which gives off a constant dorsal carpal branch over the posterior aspect of the trapezium. This vessel crosses the carpus transversely to the inner border of the hand and is joined in half of the cases by the interosseous artery; contribution of the posterior carpal branch of the ulnar artery was more rarely found than usually described (25%). From the arch thus formed are given off short branches which run upwards along the intercarpal joints and enter the posterior aspects of the proximal carpal bones whose lower halves seem to be the most favorably supplied. On the contrary, the distal carpal bones receive several short twigs which penetrate their posterior surfaces. Although the arrangement of the pedicles may undergo many a variation, attention must be drawn to the fact that the dorsal arteries are smaller but denser than the volar, except for the hamate. Therefore 2 different patterns of arterial supply can be described at the carpus: the blood supply of the outer and axial bones--scaphoid, trapezium, lunate, capitate--derives from the radial and the interosseous arteries and is probably shared equally by volar and dorsal branches; the inner bones--triquetrum, hamate--are primarily supplied by volar or medial branches coming almost exclusively from the ulnar artery.     

PARALLEL EVOLUTION OF HAND ANATOMY IN KANGAROOS AND VOMBATIFORM MARSUPIALS: FUNCTIONAL AND EVOLUTIONARY IMPLICATIONS

Abstract:  The anatomy of the mammalian hand is exposed to an intriguing interplay between phylogeny and function, and provides insights on phylogenetic affinities as well as locomotory habits of extinct species. Within the marsupial order Diprotodontia, terrestrial plantigrade quadrupedalism evolved twice, in the mostly extinct vombatiforms and in extant macropodoids. To assess the influence of functional and phylogenetic signal on the manus in these two clades, manual anatomy and digital proportions in specimens of eight extinct and three extant vombatiforms were investigated and compared with extant macropodoids and extant possums. The results reveal extensive parallelisms in the carpal region of vombatiforms and macropodoids, including flattened distal metacarpal facets, reduction of the palmar process of the hamatum, reduction of mid-wrist joint curve, extensive hamatum/scaphoid contact, and absence of a lunatum. These transformations appear to be related to stabilization of the wrist for plantigrade locomotion. Vombatiforms are apomorphic in scaphoid and triquetrum anatomy and their metacarpals are much more gracile than in other Diprotodontia. Manual diversity is greater in vombatiforms than in macropodoids, as probably was locomotor diversity. Digital proportions as well as wrist anatomy divide the extinct vombatiforms into species resembling arboreal diprotodontians, whereas others group with terrestrial quadrupedal kangaroos and wombats. The latter is suggested to be owing to plantigrade locomotion and/or large size. Carpal anatomy and digital proportions suggest that a range of earlier diverging vombatiforms may have been arboreal or scansorial. As such, we propose that the ancestor of extant vombatiforms (koalas and wombats) may have been arboreal, an option that deserves consideration in the reconstruction of vombatiform evolution.     

Anatomy of the squirrel wrist: bones, ligaments, and muscles

Anatomical differences among squirrels are usually most evident in the comparison of flying squirrels and nongliding squirrels. This is true of wrist anatomy, probably reflecting the specializations of flying squirrels for the extension of the wing tip and control of it during gliding. In the proximal row of carpals of most squirrels, the pisiform articulates only with the triquetrum, but in flying squirrels there is also a prominent articulation between the pisiform and the scapholunate, providing a more stable base for the styliform cartilage, which supports the wing tip. In the proximal wrist joint, between these carpals and the radius and ulna, differences in curvature of articular surfaces and in the location of ligaments also correlate with differences in degree and kind of movement occurring at this joint, principally reflecting the extreme dorsal flexion and radial deviation of the wrist in flying squirrels when gliding. The distal wrist joint, between the proximal and distal rows of carpals, also shows most variation among flying squirrels, principally in the articulations of the centrale with the other carpal bones, probably causing the distal row of carpal bones to function more like a single unit in some animals. There is little variation in wrist musculature, suggesting only minor evolutionary modification since the tribal radiation of squirrels, probably in the early Oligocene. Variation in the carpal bones, particularly the articulation of the pisiform with the triquetrum and the scapholunate, suggests a different suprageneric grouping of flying squirrels than previously proposed by McKenna (1962) and Mein (1970). J. Morphol. 246:85-102, 2000. Published 2000 Wiley-Liss, Inc.     

Robusticity and osteoarthritis at the trapeziometacarpal joint in a Bronze Age population from Tell Abraq, United Arab Emirates

Osteoarthritis (OA) is a progressive disease of the joints and can cause pain, reduced range of motion and strength, and ultimately loss of function at affected joints. Osteoarthritis often occurs at sites where biomechanical stress is acutely severe or moderate but habitual over the course of a lifetime. Skeletal remains from an Umm an-Nar tomb at Tell Abraq, United Arab Emirates (ca. 2300 BC), were recovered and represented over 300 individuals of all ages. The remains were disarticulated, commingled, and mostly fragmented. An analysis of 650 well-preserved adult metacarpal and carpal bones, from the tomb's western chamber, revealed that over 53% of the trapeziometacarpal joint facets showed signs of OA varying from mild to severe. The first and second metacarpals and trapezium bones were sided and evaluated for OA at the trapeziometacarpal joint articulations. Osteoarthritis was detected on 53% of the first metacarpals, 40% of the second metacarpals, and 57% of the trapezium bones. All specimens appeared enlarged, and the first metacarpals were assessed for sexual identification and robusticity. Eighty-five percent of the bones were probable males, and more than 80% of them had a robusticity index of 60 or higher. A strong correlation was found between OA, sex, and robusticity. High levels of OA and robusticity at the thumb suggest that the people of Tell Abraq were habitually involved in biomechanically challenging work with their hands.     

Experimental investigation of the scaphoid strain during wrist motion

The scaphoid is the most frequently fractured of the carpal bones [Taleisnik, J., The Wrist, Churchill Livingstone, New York (1985)]. This project was undertaken to qualitatively evaluate the strain in the scaphoid during wrist motion using a newly developed strain gage method. Strain gage resettes were mounted within the scaphoid and the range of motion of the hand was monitored using a custom designed electrogoniometer and data acquisition system. Ten specimens were utilized for this study. Results indicated that supination/pronation (S/P) of the forearm did not affect the strain in the scaphoid. A map of the strain in the waist of the scaphoid, as a function of flexion/extension (F/E) and radial/ulnar deviation (R/U), was generated. The contour plot of scaphoid strain vs range of motion (ROM) shows a valley where strains are low. Minimum scaphoid strain was found near neutral F/E and 15° of ulnar deviation.     

New primate carpal bones from Rudabánya (late Miocene, Hungary): taxonomic and functional implications

We describe a scaphoid and two capitates from the late Miocene site of Rudabánya, Hungary using qualitative and quantitative comparisons to a large sample of hominoid, cercopithecoid, and platyrrhine primates. The scaphoid (RUD 202) is not fused to the os centrale and in this way is like most primates other than African apes and humans (hominines). Qualitatively, its morphology is most similar to Pongo, and univariate analyses generally confirm an ape-like morphology with an increased range of mobility. One capitate (RUD 167) is compatible in size to the scaphoid, and its morphology suggests a combination of monkey-like generalized arboreality and ape-like enhanced mobility. RUD 203 is a smaller, fragmentary capitate, about half the size of RUD 167, and preserves only the distal portion of the body with the third metacarpal articular surface. Its morphology is virtually identical to that of RUD 167, and an exact randomization test revealed that it is statistically likely to find two carpal bones of such disparate sizes within one taxon. However, due to morphological similarities with other Miocene hominoids as well as implications for size variation within one taxon and sex, we consider the taxonomic affiliation of RUD 203 to be unresolved. We attribute the scaphoid and RUD 167 capitate to the hominine Rudapithecus hungaricus (formerly Dryopithecus brancoi; see Begun et al., 2008) based on overall morphological similarity to extant apes, particularly Pongo, and not to the pliopithecoid Anapithecus hernyaki, the only other primate known from Rudabánya. The similarities in carpal morphology to suspensory taxa are consistent with previous interpretations of Rudapithecus positional behavior. The scaphoid and the RUD 167 capitate are consistent in size with a partial skeleton including associated postcranial and craniodental specimens from the same level at the locality and may be from the same individual. These are the first carpal bones described from Rudabánya and from this taxon, and they add to our understanding of the evolution of arboreal locomotion in late Miocene apes.     

Coordinate systems for the carpal bones of the wrist

The eight small and complexly shaped carpal bones of the wrist articulate in six degrees of freedom with each other and to some extent with the radius and the metacarpals. With the increasing number and sophistication of studies of the carpus, a standardized definition for a coordinate system for each the carpal bones would aid in the reporting and comparison of findings. This paper presents a method for defining and constructing a coordinate system specific to each of the eight carpal bones based upon the inertial properties of the bone, derived from surface models constructed from three-dimensional (3-D) medical image volumes. Surface models from both wrists of 5 male and 5 female subjects were generated from CT image volumes in two neutral wrist positions (functional and clinical). An automated algorithm found the principal inertial axes and oriented them according to preset conditions in 85% of the bones, the remaining bones were corrected manually. Six of the eight carpal bones were significantly more extended in the functional neutral position than in the clinical neutral position. Gender had no significant effect on carpal bone posture in either wrist position. Correlations between the 3-D carpal posture and the commonly used 2-D clinical radiographic carpal angles are established. 3-D coordinate systems defined by the anatomy of the carpal bone, such as the ones presented here, are necessary to completely describe 3-D changes in the posture of the carpal bones.