Magnitude of sex differences in dichotomous ossification sequences of the hand and wrist

Among the dichotomous (present/absent:absent/present) ossinication sequences individually ascertained in 3059 boys and girls with at least one but not more than 27 ossification centers of the hand and wrist, 54 such sequences exhibit statistically-significant sex differences in frequency, 32 of them at the 1% confidence level or better. Analyzed by regions (rows and rays, epiphyses and round bones), ten centers, primarily distals and those of the first digit, account for the majority of the significant sex differences.     

The influence of mass,volume and density on the frequency of recovery of fossil hominid hand and wrist bones

A study of the mass, volume and density of each of the wrist and hand bones of male and female human skeletons was undertaken. It was found that the mass and volume (i.e. size) of the bones are well correlated with the relative frequencies of preservation ofAustralopithecus and earlyHomo wrist and hand bones from fossil hominid sites in Africa. In general, the larger the bone, the greater its preservation frequency. In contrast to findings on bovid bones, the density of hand and wrist bones is not well correlated with the frequency of such bones recovered from these sites. These findings may be explained in terms of the agents of deposition of the bones, the physical nature of the deposit, and the methods of extraction of the fossils from the deposit.     

Associations between the rates of elongation of the short bones of the hand

Data from serial radiographs of 40 children were used to study the rates of total, diaphyseal and epiphyseal elongation of the bones of the hand from 3–13 years. Communality indices were calculated from complete correlation matrices for each sex. These indices tended to be lower for boys than girls. Communality indices for distal phalanges were lower than for the other bones and those for epiphyseal elongation were lower than those for total or diaphyseal elongation. Correlations between groups of bones in their rates of elongation were higher in rows of bones than in rays. There were statistically significant neighborhood effects and a slight tendency to marginal effects in the correlations between rates of elongation in rows of bones; neither of these effects was present in rays.     

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.     

The differential frequency of preservation of early hominid wrist and hand bones

A survey of hominid hand and wrist bones of Plio-Pleistocene fossil hominid sites in Africa was undertaken. There are 101 specimens in total, from 7 sites. Carpals are most rarely preserved, but certain elements such as the capitate tend to be more frequently preserved than others. There is a preservation rate cline from proximal to distal in the hand, proximal elements (metacarpals) being numerically better preserved than the distal elements (proximal, middle and distal phalanges, in that order of preservation). The proportion of complete, or nearly complete hand and wrist bones is greater in the distal than in the proximal elements. There is no statistical difference in the frequency of preservation of left and right sides, or in the frequency of preservation of proximal and distal ends of individual bones, although there is a tendency for proximal ends to occur more frequently than distal ends. The incidence of hand and wrist bones in fossil deposits is low compared with that of other post-cranial skeletal elements (with the exception of foot bones, where the incidence is similar). This could be accounted for by depositional factors, but preparation techniques and differential collection of specimens may play a role.     

Pattern profile analysis of hominid and chimpanzee hand bones

In a study designed to complement morphological research on hominid hand bones, length and width measurements of the thumb, index, and middle rays were obtained from radiographs of modern human hands. These rays are primary in precision-gripping postures and are therefore the ones most relevant for investigating evolutionary changes in fine manipulation. Pattern profile analysis allows individuals or samples to be plotted against a reference sample in standard deviation units, or Z-scores. It provides an indication of how different measurements are from modern human averages, while taking into consideration the degree of variation present within modern human samples. A pattern profile for chimpanzees is clearly distinct from humans but quite similar to that of a bonobo, demonstrating the promise of pattern analysis. Partial pattern profiles of several of the more complete early hominid bones from Hadar, Swartkrans, and Olduvai (O.H. 7) are presented and compared. Hadar bones are long and wide at midshaft relative to articular widths; both body-size effects and functional differences are likely. Thumb distal phalanges from Swartkrans and Olduvai both have relatively small base widths, but they differ in other proportions. Two first metacarpals from Swartkrans show distinct patterns. The profiles of La Ferrassie I and Shanidar IV show the characteristically large Neanderthal distal phalanges. Profiles of Skhūl IV and P?edmost III are alike in some regions with reference to modern North American white males, though they are less similar overall than are those of the two Neanderthals. © 1995 Wiley-Liss, Inc.     

Communalities for rates of diaphyseal elongation of short bones of the hand of children with Down syndrome

Communality indices for rates of elongation of diaphyses of short bones of the hand were computed from serial data for children with Down syndrome, 7 to 14 years of age. Communalities were larger for adjacent than for nonadjacent bones and also larger for bones grouped in rows rather than rays of the hand. This pattern is similar to that reported for normal children. Communality indices for rates of diaphyseal elongation for girls with Down syndrome were lower than those of boys with Down syndrome and normal children.     

The essential characteristics of a rating system

The principles of the Tanner-Whitehouse-Healy system for assessing skeletal maturity will be described and in particular the statistical basis for the scores given to each stage of each bone in the hand and wrist. The uses of the system, particularly in clinical cases, will be discussed together with reliability of the ratings. A revised system will be presented in which a part of the stages are amalgamated and the possibility of dropping some of the bones will be discussed. General discussion will be invited on the logical basis for selecting certain stages of certain bones as superior to others as assessors of maturity.     

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.     

Bone density spatial patterns in the distal radius reflect habitual hand postures adopted by quadrupedal primates

Primates adopt diverse hand postures during terrestrial and above-branch quadrupedal locomotion--knuckle-walking, digitigrady, and palmigrady--that incorporate varying degrees of wrist dorsiflexion (i.e., extension). Although relationships between hand postures, wrist joint range of motion, and the external properties of wrist bones (e.g., surface morphology) have been examined, the relationship between hand postures and the internal properties of wrist bones (e.g., bone density) remains largely unexplored. Because articular joint surfaces transmit mechanical loads between conjoining limb bones, measures of density (e.g., magnitudes and patterns) in the subchondral cortical plate of bone of the distal radius can be used to evaluate load regimes experienced by the wrist joint in different hand postures. We assessed apparent (i.e. optical) density patterns in several extant catarrhine primate taxa partitioned into different hand posture groups: knuckle-walking apes, digitigrade monkeys, and palmigrade monkeys. Computed tomography osteoabsorptiometry (CT-OAM) was used to construct maximum intensity projection (MIP) maps of apparent densities. High apparent density areas were characterized relative to a dorsal-volar reference plane and compared across hand posture groups. All groups had large percentage areas of high apparent density in the dorsal region of the distal radial articular surface. Only knuckle-walking apes, however, had a large percentage area of high apparent density in the volar region of the distal radial articular surface. These patterns are consistent with radiocarpal articulations in specific hand postures as evidenced by available radiographic data and suggest that the different habitual hand postures adopted by monkeys and African apes during quadrupedal locomotion have different stereotypic loading patterns. This has implications for understanding the functional morphology and evolution of knuckle-walking and digitigrade hand postures in primates.     

New wrist bones of the Malagasy giant subfossil lemurs

Recently discovered wrist bones of the Malagasy subfossil lemurs Babakotia radofilai, Palaeopropithecus ingens, Mesopropithecus dolichobrachion, and Megaladapis madagascariensis shed new light on the postcranial morphologies and positional behaviors that characterized these extinct primates. Wrist bones of P. ingens resemble those of certain modern hominoids in having a relatively enlarged ulnar head and dorsally extended articular surface on the hamate, features related to a large range of rotation at the inferior radioulnar and midcarpal joints. The scaphoid of P. ingens is also similar to that of the extant tree sloth Choloepus in having an elongate, palmarly directed tubercle forming a deep radial margin of the carpal tunnel for the passage of large digital flexors. In contrast, wrist remains of Megaladapis edwardsi and M. madagascariensis exhibit traits observed in the hands of extant pronograde, arboreal primates; these include a dorsopalmarly expanded pisiform and well-developed "spiral" facet on the hamate. Moreover, Megaladapis spp. and Mesopropithecus dolichobrachion possess bony tubercles (e.g., scaphoid tubercle and hamate hamulus) forming the carpal tunnel that are relatively similar in length to those of modern pronograde lemurs. Babakotia and Mesopropithecus differ from Megaladapis in exhibiting features of the midcarpal joint related to frequent supination and radioulnar deviation of the hand characteristic of animals that use vertical and quadrumanous climbing in their foraging behaviors. Comparative analysis of subfossil lemur wrist morphology complements and expands upon prior inferences based on other regions of the postcranial skeleton, and suggests a considerable degree of locomotor and postural heterogeneity among these recently extinct primates.     

Correlations between rates of epiphyseal and diaphyseal elongation in the short bones of the hand

Serial annual radiographs of the hand have been used to analyze the rates of elongation of the epiphyses and diaphyses of the metacarpals and phalanges in children at ages from 3 to 13 years. The rates of elongation for many corresponding ephiphyses and diaphyses (i.e., of the same bone) are negatively correlated but to an extent that is not statistically significant for any particular bone. This tendency toward negative correlations is found for most of these bones although the correlation coefficients for most of the metacarpals are positive in each sex. Within rays, the correlation indices between the rates of elongation for corresponding epiphyses and diaphyses (i.e., of the same bone) have larger negative or smaller positive values than for those between either adjacent and non-corresponding or non-adjacent and non-corresponding epiphyses and diaphyses (i.e., not of the same bone but of either adjacent or non-adjacent bones). The communality indices for the ratio between the rates of epiphyseal and diaphyseal elongation in particular bones are more highly correlated in the girls than in the boys and within the rows than within the rays. Some implications of the tendency to negative correlations between the rates of elongation of corresponding epiphyses and diaphyses have been discussed.     

Trabecular bone structure in the primate wrist

Trabecular (or cancellous) bone has been shown to respond to mechanical loading throughout ontogeny and thus can provide unique insight into skeletal function and locomotion in comparative studies of living and fossil mammalian morphology. Trabecular bone of the hand may be particularly functionally informative because the hand has more direct contact with the substrate compared with the remainder of the forelimb during locomotion in quadrupedal mammals. This study investigates the trabecular structure within the wrist across a sample of haplorhine primates that vary in locomotor behaviour (and thus hand use) and body size. High‐resolution microtomographic scans were collected of the lunate, scaphoid, and capitate in 41 individuals and eight genera (Homo, Gorilla, Pan, Papio, Pongo, Symphalangus, Hylobates, and Ateles). We predicted that particular trabecular parameters would 1) vary across suspensory, quadrupedal, and bipedal primates based on differences in hand use and load, and 2) scale with carpal size following similar allometric patterns found previously in other skeletal elements across a larger sample of mammals and primates. Analyses of variance (trabecular parameters analysed separately) and principal component analyses (trabecular parameters analysed together) revealed no clear functional signal in the trabecular structure of any of the three wrist bones. Instead, there was a large degree of variation within suspensory and quadrupedal locomotor groups, as well as high intrageneric variation within some taxa, particularly Pongo and Gorilla. However, as predicted, Homo sapiens, which rarely use their hands for locomotion and weight support, were unique in showing lower relative bone volume (BV/TV) compared with all other taxa. Furthermore, parameters used to quantify trabecular structure within the wrist scale with size generally following similar allometric patterns found in trabeculae of other mammalian skeletal elements. We discuss the challenges associated with quantifying and interpreting trabecular bone within the wrist. J. Morphol. 275:572–585, 2014. © 2013 Wiley Periodicals, Inc.     

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.     

Musculoskeletal Model of the Upper Limb Based on the Visible Human Male Dataset

A mathematical model of Ihe human upper limb was developed based on high-resolution medical images of the muscles and bones obtained from the Visible Human Male ( HM) project. Three-dimensional surfaces of the muscles and bones were reconstructed from Computed Tomography (CT) images and Color Cryosection images obtained from the VHM cadaver. Thirteen degrees of freedom were used to describe the orientations of seven bones in the model: clavicle, scapula, humerus, radius, ulna, carpal bones, and hand. All of the major articulations from the shoulder girdle down to the wrist were included in the model. The model was actuated by 42 muscle bundles, which represented the actions of 26 muscle groups in the upper limb. The paths of the muscles were modeled using a new approach called the Obstacle-set Method (33) The calculated paths of the muscles were verified by comparing the muscle moment arms computed in the model with the results of anatomical studies reported in the literature, In-vivo measurements of maximum isometric muscle torques developed at the shoulder, elbow, and wrist were also used to estimate the architectural properties of each musculotendon actuator in the model. The entire musculoskeletal model can be reconstructed using the data given in this paper, along with information presented in a companion paper which defines the kinematic structure of the model (26)     

Fitting manifold surfaces to three-dimensional point clouds

We present a technique for fitting a smooth, locally parameterized surface model (called the manifold surface model) to unevenly scattered data describing an anatomical structure. These data are acquired from medical imaging modalities such as CT scans or MRI. The manifold surface is useful for problems which require analyzable or parametric surfaces fitted to data acquired from surfaces of arbitrary topology (e.g., entire bones). This surface modeling work is part of a larger project to model and analyze skeletal joints, in particular the complex of small bones within the wrist and hand. To demonstrate the suitability of this model we fit to several different bones in the hand, and to the same bone from multiple people.     

Musculoskeletal model of the upper limb based on the visible human male dataset

A mathematical model of the human upper limb was developed based on high-resolution medical images of the muscles and bones obtained from the Visible Human Male (VHM) project. Three-dimensional surfaces of the muscles and bones were reconstructed from Computed Tomography (CT) images and Color Cryosection images obtained from the VHM cadaver. Thirteen degrees of freedom were used to describe the orientations of seven bones in the model: clavicle, scapula, humerus, radius, ulna, carpal bones, and hand. All of the major articulations from the shoulder girdle down to the wrist were included in the model. The model was actuated by 42 muscle bundles, which represented the actions of 26 muscle groups in the upper limb. The paths of the muscles were modeled using a new approach called the Obstacle-set Method [33]. The calculated paths of the muscles were verified by comparing the muscle moment arms computed in the model with the results of anatomical studies reported in the literature. In-vivo measurements of maximum isometric muscle torques developed at the shoulder, elbow, and wrist were also used to estimate the architectural properties of each musculotendon actuator in the model. The entire musculoskeletal model can be reconstructed using the data given in this paper, along with information presented in a companion paper which defines the kinematic structure of the model [26].     

Knuckle-walking anteater: a convergence test of adaptation for purported knuckle-walking features of African Hominidae

Appeals to synapomorphic features of the wrist and hand in African apes, early hominins, and modern humans as evidence of knuckle-walking ancestry for the hominin lineage rely on accurate interpretations of those features as adaptations to knuckle-walking locomotion. Because Gorilla, Pan, and Homo share a relatively close common ancestor, the interpretation of such features is confounded somewhat by phylogeny. The study presented here examines the evolution of a similar locomotor regime in New World anteaters (order Xenarthra, family Myrmecophagidae) and uses the terrestrial giant anteater (Myrmecophaga tridactyla) as a convergence test of adaptation for purported knuckle-walking features of the Hominidae. During the stance phase of locomotion, Myrmecophaga transmits loads through flexed digits and a vertical manus, with hyperextension occurring at the metacarpophalangeal joints of the weight-bearing rays. This differs from the locomotion of smaller, arboreal anteaters of outgroup genera Tamandua and Cyclopes that employ extended wrist postures during above-branch quadrupedality. A number of features shared by Myrmecophaga and Pan and Gorilla facilitate load transmission or limit extension, thereby stabilizing the wrist and hand during knuckle-walking, and distinguish these taxa from their respective outgroups. These traits are a distally extended dorsal ridge of the distal radius, proximal expansion of the nonarticular surface of the dorsal capitate, a pronounced articular ridge on the dorsal aspects of the load-bearing metacarpal heads, and metacarpal heads that are wider dorsally than volarly. Only the proximal expansion of the nonarticular area of the dorsal capitate distinguishes knuckle-walkers from digitigrade cercopithecids, but features shared with digitigrade primates might be adaptive to the use of a vertical manus of some sort in the stance phase of terrestrial locomotion. The appearance of capitate nonarticular expansion and the dorsal ridge of the distal radius in the hominin lineage might be indicative of a knuckle-walking ancestry for bipedal hominins if interpreted within the biomechanical and phylogenetic context of hominid locomotor evolution.     

Molecular evolution of the mitochondrial 12S rRNA in Ungulata (mammalia)

The complete 12S rRNA gene has been sequenced in 4 Ungulata (hoofed eutherians) and 1 marsupial and compared to 38 available mammalian sequences in order to investigate the molecular evolution of the mitochondrial small-subunit ribosomal RNA molecule. Ungulata were represented by one artiodactyl (the collared peccary, Tayassu tajacu, suborder Suiformes), two perissodactyls (the Grevy's zebra, Equus grevyi, suborder Hippomorpha; the white rhinoceros, Ceratotherium simum, suborder Ceratomorpha), and one hyracoid (the tree hyrax, Dendrohyrax dorsalis). The fifth species was a marsupial, the eastern gray kangaroo (Macropus giganteus). Several transition/transversion biases characterized the pattern of changes between mammalian 12S rRNA molecules. A bias toward transitions was found among 12S rRNA sequences of Ungulata, illustrating the general bias exhibited by ribosomal and protein-encoding genes of the mitochondrial genome. The derivation of a mammalian 12S rRNA secondary structure model from the comparison of 43 eutherian and marsupial sequences evidenced a pronounced bias against transversions in stems. Moreover, transversional compensatory changes were rare events within double-stranded regions of the ribosomal RNA. Evolutionary characteristics of the 12S rRNA were compared with those of the nuclear 18S and 28S rRNAs. From a phylogenetic point of view, transitions, transversions and indels in stems as well as transversional and indels events in loops gave congruent results for comparisons within orders. Some compensatory changes in double-stranded regions and some indels in single-stranded regions also constituted diagnostic events. The 12S rRNA molecule confirmed the monophyly of infraorder Pecora and order Cetacea and demonstrated the monophyly of suborder Suiformes. However, the monophyly of the suborder Ruminantia was not supported, and the branching pattern between Cetacea and the artiodactyl suborders Ruminantia and Suiformes was not established. The monophyly of the order Perissodactyla was evidenced, but the relationships between Artiodactyla, Cetacea, and Perissodactyla remained unresolved. Nevertheless, we found no support for a Perissodactyla + Hyracoidea clade, neither with distance approach, nor with parsimony reconstruction. The 12S rRNA was useful to solve intraordinal relationships among Ungulata, but it seemed to harbor too few informative positions to decipher the bushlike radiation of some Ungulata orders, an event which has most probably occurred in a short span of time between 55 and 70 MYA. Correspondence to: E. Douzery