Is the Clavicle of Apes Long? An Investigation of Clavicular Length in Relation to Body Mass and Upper Thoracic Width

An elongated clavicle is one of the distinct features of apes and humans. It plays an important role in providing mobility as well as stability for the shoulder joints. The relative length of the clavicle is an especially important factor in limiting the range of shoulder joint excursion. It is said that among primates, Asian apes, i.e., gibbons and orang-utans, have very long clavicles. At the same time, they also have a wide upper thoracic cage, which may diminish the effective length of the clavicle. To clarify the length of the clavicle in apes, from the standpoint of the functional anatomy of the shoulder girdle, we examined clavicular length in 15 anthropoid species exhibiting various positional behaviors. The results confirm that clavicle length in Asian apes is long, and chimpanzees have a short clavicle like that of Old and New World monkeys, when scaled to body mass. The clavicular length of chimpanzees, however, is intermediate between Old World monkeys and Asian apes when scaled against thoracic width. Therefore, living apes can be grouped together, albeit just barely, by possession of a relatively long clavicle for their thoracic cage size. Interestingly, New World monkeys tend to exhibit a longer clavicle than Old World monkeys of equivalent body mass or thoracic cage width. Although it is unclear whether the ancestral condition of clavicular length in anthropoids was similar to that of living Old or New World monkeys, an elongation of clavicle was an important step toward evolution of the modern body plan of hominoids.     

Technical note: Virtual reconstruction of a fragmentary clavicle

We report a procedure for the virtual reconstruction of incomplete human bones applicable to skeletal remains from archaeological excavations or to reconstructive and prosthetic surgery. To test the procedure, we reconstructed a fragmented left clavicle on the basis of the contralateral clavicle. The procedure involved 3‐D laser scanner acquisition of the left clavicle (complete but broken into two parts), the same manually reconstructed bone, and the intact right clavicle, which was mirror‐imaged and used as a reference for the reconstruction of the whole left clavicle. Because it was not possible to recognize homologous anatomical landmarks, on the two reference models (a mirror‐image copy of the right clavicle and the main fragment of the left), we identified three grids with an increasing number of corresponding landmarks, which constituted the framework of the deformation process. The three reconstructed digital models of the clavicle closely approximated the model of the original clavicle. They also showed that an increasing number of landmarks did not significantly improve the reconstructed model. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

Influence of geometrical personalization on the simulation of clavicle fractures

Finite element body models enable the evaluation of car occupant protection. In general, these models represent average males and inter-individual geometry variability is not taken into account. As the most frequent shoulder injury during car lateral accidents is a clavicle fracture, the purpose of this study is to investigate whether clavicle geometry has an influence on bone response until failure, and whether geometrical personalization of clavicle models is required. Eighteen clavicles from 9 subjects (5 males and 4 females, mean age: 76 +/- 12 years) were harvested. Six clavicles were scanned, enabling the development of subject-specific models and the quantification of geometrical features defining shape and cortical thickness. Bone mineral densities (BMD) were measured through double X-ray absorptiometry. Then, the general clavicle responses to dynamic compression until failure were studied. Simulations of the compression tests were carried out with the subject-specific models to assess the sensitivity of force-deflection clavicle responses to geometrical features. Clavicle fractures occurred at an average velocity of 1.41 +/- 0.4 ms(-1), with a fracture force of 1.48 +/- 0.46 kN and a deflection of 5.4 +/- 1.1 mm. A significant difference was found between male and female clavicle force values at rupture although their BMDs were not significantly different. Simulations with subject-specific models led to the conclusion that cortical bone thickness and bone shape have large effects on bone responses until failure and on fracture location. This study highlights the need for a geometrical personalization of clavicle models in order to take into account both gender discrepancies concerning clavicle shape and aging effects affecting cortical thickness.     

Morphometric affinities of the human shoulder

To analyze differences between apes and monkeys and the affinities of man, we have studied the shoulder girdle of 327 specimens of anthropoid primates. The scapula, clavicle and humerus are viewed as an integrated functional complex on the basis of 18 measurements. Several varieties of multivariate analysis show that man is clearly closer to other hominoids than to the included monkey taxa (whether terrestrial or arboreal, Old World or New World). The marked shoulder differences between apes and monkeys and similarities between apes and man correlate with the muscular anatomy, which in hominoids allows the motions involved in their locomotion and feeding behavior. As the hominid-pongid correspondence in shoulder morphology is especially detailed regarding the functionally important joint surfaces, it is consistent with a fairly recent period of common ancestry and behavior. No hypothetical evolutionary pathway or ancestral form of the human shoulder need look far beyond the model afforded by extant pongids. In contrast with previous studies on the primate shoulder, these results agree with information accumulating from other systems—comparative anatomy, primate behavior, and molecular biology — in suggesting very close relationship between man and extant African pongids.     

A New Interpretation of the Crocodile Forelimb Morphological Features as Adaptation to Parasagittal Quadrupedal Locomotion on the Ground

We describe the crocodile forelimb features that distinguish them from other reptiles. Reduction of the clavicle and a change in the coracoid shape seem to be another way of maintaining the efficient step length, while the antebrachium and manus transformations create peculiar oblique manus position on the ground to promote the forelimb parasagittalization.     

贵州兴义猫猫洞出土的人类化石

本文主要是对贵州兴义猫猫洞遗址出土的人化石进行的初步研究,并对哺乳动物化石、地层、堆积物作叙述。该遗址的人类化石包括下颌骨4件,股骨3段,是中国南方洞穴遗址中发现的数量较多,系统分类地位较清楚的晚期智人的化石。人类化石同层的鹿牙作铀系法测年结果为14500±1200 BP,相当于更新世的末期。     

鸽子洞的人类化石

鸽子洞遗址经过1973和1975发掘,发现大量石制品、用火遗迹和动物化石。笔者在鸽子洞所出的动物碎骨中拣出三件人类化石,它们的形态特征及尺寸都和现代人没有区别,属发展阶段的晚期智人,地质时代为晚更新世晚期。     

Preliminary analysis of<Emphasis Type="Italic"> Nacholapithecus</Emphasis> scapula and clavicle from Nachola,Kenya

The Miocene ape Nacholapithecus is known from rather complete skeletons; some of them preserve the shoulder joint, identified by three scapulae and one clavicle. Comparisons made with other Miocene and living apes (Proconsul, Equatorius, Ugandapithecus) suggest that the mobility of the scapulohumeral joint was important, and scapular features such as the morphology and position of the spine and the morphology of the acromion and axillary border resemble those of climbing arboreal primates except for chimpanzees, gorillas, or orang-utans. From the size of the scapula (male Nasalis size), it is clear that the animal is smaller than an adult chimpanzee, but the clavicle is almost as relatively long as those of chimpanzees. Some features closer to colobine morphology reinforce the hypothesis that Nacholapithecus was probably a good climber and was definitely adapted for an arboreal life.     

Evolution of the human shoulder: some possible pathways

Osteometric data, apparently reflecting functional parameters of the shoulder in the Anthropoidea, have been examined by a combination of multivariate techniques in an attempt to define minimum pathways possibly followed in the evolution of the human shoulder. Principal components analysis has been used to gauge size-related shape effects. The combination of D² and canonical analysis has suggested that the shoulder in man is unlikely to have evolved (a) from one similar to that of any extant monkey whether arboreal or terrestrial, (b) from one similar to that of any terrestrial ape (like present-day Pan and Gorilla), or (c) from one similar to that of a highly specialised ricochetal armswinging ape (such as present day Hylobates and Symphalangus). The analyses suggest positively that the minimum evolutionary pathway may well have been from the shoulder of a totally arboreal ape, presumably genetically related to Pan and Gorilla, but functionally similar to that evolved in parallel in the highly arboreal orang-utan, Pongo. Information from fragmentary fossils (the scapula from Sterkfontein and the clavicle from Olduvai) supports these conclusions.     

The development of the clavicle in man

The development of the human clavicle was studied in 50 to 60 d old human embryos. Our findings are summarized as follows: The whole clavicle develops from a cartilaginous anlage. In the middle part of the clavicle, an osseus cuff develops very early by the ossification in the perichondrium. In the lumen of this cuff, a cartilaginous cork persists which is resorbed and replaced by bone and marrow later than in other bones. It is possible that cartilaginous nests may persist in the middle part of the clavicle. In both extremities of the clavicle, the normal enchondral ossification exists as it is described in other anlages. It is difficult to explicate the syndrome of the cleido-facial and cleido-cranial dysostoses only as disturbances of the endesmal ossification.     

Brief communication: histological age estimation using rib and clavicle.

Histological methods for estimating age at death using osteon population densities for the rib, clavicle, and rib and clavicle combined are presented. Predicting formulas were generated from a sample of 40 individuals of known age, sex, and race. Independent samples of 12 ribs and 7 clavicles were used to test the formulas. Mean differences between known and predicted ages were 1.1 years, 2.6 years, and 3.4 years for the clavicle, rib and clavicle combined, and rib formulas respectively. An analysis of variance found no significant differences among the means for predicted and known ages. Since the formula based upon rib and clavicle combined has the higher standard error and r2, and includes data from different bones, it should provide better overall accuracy and reliability, and is recommended whenever both bones are available.     

Sequence of a yeast DNA fragment containing a chromosomal replicator and a tRNA Glu 3 gene.

The sequence of a 1.9 kb Bam x Hind III fragment from yeast has been determined. This fragment is part of a yeast 6.7 kb Hind III segment cloned into pBR322 (pY20). The fragment carries a single gene for a glutamate tRNA which has no intron. According to genetic analyses [1] this fragment also contains a yeast chromosomal replicator. We have analyzed the sequence for potential open reading frames and for several structural features which are thought to be involved in the initiation of DNA replication. Hybridization studies have revealed that portions of this sequence are repeated within the yeast genome.     

Evaluation of regression-based 3-D shoulder rhythms

The movements of the humerus, the clavicle, and the scapula are not completely independent. The coupled pattern of movement of these bones is called the shoulder rhythm. To date, multiple studies have focused on providing regression-based 3-D shoulder rhythms, in which the orientations of the clavicle and the scapula are estimated by the orientation of the humerus. In this study, six existing regression-based shoulder rhythms were evaluated by an independent dataset in terms of their predictability. The datasets include the measured orientations of the humerus, the clavicle, and the scapula of 14 participants over 118 different upper arm postures. The predicted orientations of the clavicle and the scapula were derived from applying those regression-based shoulder rhythms to the humerus orientation. The results indicated that none of those regression-based shoulder rhythms provides consistently more accurate results than the others. For all the joint angles and all the shoulder rhythms, the RMSE are all greater than 5°. Among those shoulder rhythms, the scapula lateral/medial rotation has the strongest correlation between the predicted and the measured angles, while the other thoracoclavicular and thoracoscapular bone orientation angles only showed a weak to moderate correlation. Since the regression-based shoulder rhythm has been adopted for shoulder biomechanical models to estimate shoulder muscle activities and structure loads, there needs to be further investigation on how the predicted error from the shoulder rhythm affects the output of the biomechanical model.     

Patterns of clavicular bilateral asymmetry in relation to the humerus: variation among humans

Studies of directional asymmetry in the human upper limb have extensively examined bones of the arm, forearm, and hand, but have rarely considered the clavicle. Physiologically, the clavicle is an integrated element of the upper limb, transmitting loads to the axial skeleton and supporting the distal bones. However, clavicles develop in a manner that is unique among the bones of the upper limb. Previous studies have indicated that the clavicle has a right-biased asymmetry in diaphyseal breadth, as in humeri, radii, ulnae, and metacarpals, but unlike these other elements, a left-biased length asymmetry. Few studies have assessed how clavicular asymmetry relates to these other bones of the upper limb. Bilateral directional asymmetry of the clavicle is examined in relation to the humerus in a large, geographically diverse human sample, comparing lengths and diaphyseal breadths. Dimensions were converted into percentage directional (%DA) and absolute (%AA) asymmetries. Results indicate that humans have same-side %DA bias in the clavicles and humeri, and contralateral length %DA between these elements. Diaphyseal breadths in both clavicles and humeri are more asymmetric-both in direction and amount-than lengths. Differences in diaphyseal asymmetry are shown to relate to variation in physical activities among groups, but a relationship between activity and length asymmetry is not supported. This further supports previous research, which suggests different degrees of sensitivity to loading between diaphyseal breadths and maximum lengths of long bones. Differences in lateralized behavior and the potential effects of different bone development are examined as possible influences on the patterns observed among human groups.     

Growth of the clavicle and development of clavicular secondary cartilage in the embryonic mouse

Whether secondary cartilage develops in the mammalian clavicle has been a matter of controversy. This study documents, in the embryonic mouse: (a) the onset of clavicular osteogenesis at 14 days of gestation (Theiler stage 22); (b) the appearance of secondary cartilage at 16 days of gestation (Theiler stage 24) and its persistence as a prominent cartilage until 18 days of gestation; (c) that the relative growth rate of the clavicle is much higher (0.097 mg/g body weight/day) between 16 and 17 days of gestation than at later ages (mean of 0.005 mg/g/day between 17 days of gestation and 4 days postnatally), and (d) that secondary cartilage failed to form in clavicles from 15-day-old embryos maintained in vitro. We conclude that secondary cartilage is a feature of the developing mouse clavicle, that it arises when the relative growth rate of the clavicle is highest, and that the most likely stimulus for differentiation of this cartilage is mechanical, muscle-based and associated with rapid relative clavicular growth.     

A DNA fragment from the human X chromosome short arm which detects a partially homologous sequence on the Y chromosomes long arm.

An X linked human DNA fragment (named DXS31 ) which detects partially homologous sequences on the Y chromosome has been isolated. Regional localisation of the two sex linked sequences was determined using a panel of rodent-human somatic cell hybrids. The X specific sequence is located at the tip of the short arm ( Xp22 .3-pter), i.e. within or close to the region which pairs with the Y chromosome short arm at meiosis. However the Y specific sequence is located in the heterochromatic region of the long arm ( Yq11 -qter) and lies outside from the pairing region. DNAs from several XX male subjects were probed with DXS31 and in all cases a double dose of the X linked fragment was found, and the Y specific fragment was absent. DXS31 detects in chimpanzee a male-female differential pattern identical to that found in man. However results obtained in a more distantly related species, the brown lemur, suggest that the sequences detected by DXS31 in this species might be autosomally coded. The features observed with these X-Y related sequences do not fit with that expected from current hypotheses of homology between the pairing regions of the two sex chromosomes, nor with the pattern observed with other X-Y homologous sequences recently characterized. Our results suggest also that the rule of conservation of X linkage in mammals might not apply to sequences present on the tip of the X chromosome short arm, in bearing with the controversial issue of steroid sulfatase localisation in mouse.     

Structural comparison of two yeast tRNA Glu 3 genes.

DNA sequences in a 1.7 kb Pst fragment from yeast have been determined. This fragment is part of a yeast 7.4 kb Hind III segment cloned ino pBR322 (pY 5). The fragment carries a single gene for a glutamate tRNA. The coding portion of this gene is identical in sequence to that of the tRNA Glu 3 gene from pY 20 [1]. The flanking regions differ in their sequences, but possible secondary structures within the 5'-flanking regions bear similar features. Sequence homologies between pY 5 and pY 20 were detected far outside the tRNA genes. More surprisingly, extended sequence homologies were seen between the flanking regions of the pY 20 tRNA Glu 3 gene and a tRNA Ser gene [2,3]. We have also checked the known tRNA genes for structural similarities. Hybridization studies indicate that portions of the Pst fragment are repeated within the yeast genome.     

Altered alignment of the shoulder girdle and cervical spine in patients with insidious onset neck pain and whiplash-associated disorder

Clinical theory suggests that altered alignment of the shoulder girdle has the potential to create or sustain symptomatic mechanical dysfunction in the cervical and thoracic spine. The alignment of the shoulder girdle is described by two clavicle rotations, i.e, elevation and retraction, and by three scapular rotations, i.e., upward rotation, internal rotation, and anterior tilt. Elevation and retraction have until now been assessed only in patients with neck pain. The aim of the study was to determine whether there is a pattern of altered alignment of the shoulder girdle and the cervical and thoracic spine in patients with neck pain. A three-dimensional device measured clavicle and scapular orientation, and cervical and thoracic alignment in patients with insidious onset neck pain (IONP) and whiplash-associated disorder (WAD). An asymptomatic control group was selected for baseline measurements. The symptomatic groups revealed a significantly reduced clavicle retraction and scapular upward rotation as well as decreased cranial angle. A difference was found between the symptomatic groups on the left side, whereas the WAD group revealed an increased scapular anterior tilt and the IONP group a decreased clavicle elevation. These changes may be an important mechanism for maintenance and recurrence or exacerbation of symptoms in patients with neck pain.