Avian furcula morphology may indicate relationships of flight requirements among birds

This study examined furcula (wishbone) shape relative to flight requirements. The furculae from 53 museum specimens in eight orders were measured: 1) three-dimensional shape (SR) as indicated by the ratio of the direct distance between the synostosis interclavicularis and the ligamentous attachment of one of its clavicles to the actual length of the clavicle between those same two points, and 2) curvature within the primary plane (LR) as indicated by the ratio of the length of the clavicle to the sum of the orthogonal distances between the same points using a projected image. Canonical discriminant analysis of these ratios placed the individuals into a) one of four general flight categories and b) one of eight taxonomic orders. The four flight categories were defined as: i) soaring with no flapping, ii) flapping with no soaring, iii) subaqueous (i.e., all wingbeats taking place under water), and iv) partial subaqueous (i.e., wingbeats used for both aerial and submerged flapping). The error rate for placement of the specimens in flight categories was only 26.4%, about half of the error rate for placement in taxonomic orders (51.3%). Subaqueous fliers (penguins, great auks) have furculae that are the most V-shaped. Partial subaqueous fliers (alcids, storm petrels) have furculae that are more U-shaped than the subaqueous fliers but more V-shaped than the aerial flapping fliers. The partial subaqueous fliers have furculae that are also the most anteriorly curved, possibly increasing protraction capability by changing the angle of applied force and increasing attachment area for the origin of the sternobrachialis pectoralis. The increased protraction capability can counteract profile drag, which is greater in water than in air due to the greater density of water. Soaring birds have furculae that are more U-shaped or circular than those of flapping birds and have the smallest range of variation. These results indicate that the shape of the furcula is functionally related to general differences in flight requirements and may be used to infer relationships of these requirements among birds.     

中国汉族锁骨的性别判定

本文通过对279副中国汉族成人锁骨的测量,其中男性241副,女性38副。使用性别判别分析的方法,建立了锁骨性别判别的分析式,使中国汉族锁骨性别判别的准确率达到了90－94．7％,该文提出的性别判别方法,是锁骨性别判别的准确可靠的方法。     

The theropod furcula

The furcula is a structure formed by the midline fusion of the clavicles. This is the element which is unique to theropods and is important for understanding the link between birds and other theropods. New specimens from basal theropods suggest that the furcula appeared very early in theropod history. We review furcula development, function, and morphology, as well as the anatomical terminology applied to it. Furcular morphology is highly variable in crown‐group avians but is rather conserved among nonavian theropods. Here we review, or describe for the first time, the furculae in many nonavian theropods. Furculae occur in nearly all major clades of theropods, as shown by new theropod specimens from the Early Cretaceous of China and a close inspection of previously collected specimens. Informative phylogenetic characters pertaining to the furcula occur throughout Theropoda, though care should betake to consider taphonomic effects when describing furcular morphology. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

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.     

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.     

中国汉族锁骨性别差异的初步研究

本文通过对中国汉族２７９副成人干燥锁骨的测量（其中男性：２４１副,女性：３８副）,研究了中国汉族锁骨性别判定的方法,研究结果表明,用单一指标进行锁骨的性别,准确率很低,本文作者提出了锁骨表面积指数,锁骨截面积指数,并以此作为锁骨性别判定的依据,性别判别率最高可分别为９０．６８％和８４．２３％。本文提出的方法是骨骼性别判定中一种新的简单而准确的方法。     

Functional morphometric analysis of the furcula in mesozoic birds

The furcula displays enormous morphological and structural diversity. Acting as an important origin for flight muscles involved in the downstroke, the form of this element has been shown to vary with flight mode. This study seeks to clarify the strength of this form-function relationship through the use of eigenshape morphometric analysis coupled with recently developed phylogenetic comparative methods (PCMs), including phylogenetic Flexible Discriminant Analysis (pFDA). Additionally, the morphospace derived from the furculae of extant birds is used to shed light on possible flight adaptations of Mesozoic fossil taxa. While broad conclusions of earlier work are supported (U-shaped furculae are associated with soaring, strong anteroposterior curvature with wing-propelled diving), correlations between form and function do not appear to be so clear-cut, likely due to the significantly larger dataset and wider spectrum of flight modes sampled here. Interclavicular angle is an even more powerful discriminator of flight mode than curvature, and is positively correlated with body size. With the exception of the close relatives of modern birds, the ornithuromorphs, Mesozoic taxa tend to occupy unique regions of morphospace, and thus may have either evolved unfamiliar flight styles or have arrived at similar styles through divergent musculoskeletal configurations.     

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.     

Furculae in the Late Triassic theropod dinosaur<Emphasis Type="Italic">Coelophysis bauri</Emphasis>

Furculae have been identified in many dinosaurs and are synapomorphic in some clades (e.g., dromaeosaurids). All coelophysid dinosaurs exceptCoelophysis bauri have been shown to possess furculae. To date, the oldest well-documented furculae have been those of the Early Jurassic coelophysids,Coelophysis kayentakatae andCoelophysis rhodesiensis. The confirmation of furculae in Apachean-agedC. bauri further documents appearance of these elements in the Late Triassic and shows that furculae are synapomorphic in the Coelophysidae. A total of five furculae have been found in New Mexico Museum of Natural History’s (NMMNH) Ghost Ranch, New Mexico Whitaker Quarry block C-8-82. We describe three furculae in articulated juvenile skeletons; two that are missing fragments but are nearly complete, and one apparently complete, a small fragment of a furcula associated with an adultC. bauri, and one complete but isolated furcula. We access the morphology and allometry of the scapulocoracoid and furcula and show that they grow, at least in juveniles, in isometry with the humerus. The furcula ofC. bauri has a widely opened U shape that subtends an angle of ∼ 120°. All the furculae have groove-like epicleidial facets at the distal ends of the rami and some possess a small centrally located hypocleideal process. We reconstruct the complete Shoulder girdle ofC. bauri with proper spacing and angles between the elements and find that the coracoids are very close together under the center of the furcula.     

The p53 tumor suppressor protein is a critical regulator of hematopoietic stem cell behavior

In response to diverse stresses, the tumor suppressor p53 differentially regulates its target genes, variably inducing cell-cycle arrest, apoptosis or senescence. Emerging evidence indicates that p53 plays an important role in regulating hematopoietic stem cell (HSC) quiescence, self-renewal, apoptosis and aging. The p53 pathway is activated by DNA damage, defects in ribosome biogenesis, oxidative stress and oncogene induced p19ARF upregulation. We present an overview of the current state of knowledge about p53 (and its target genes) in regulating HSC behavior, with the hope that understanding the molecular mechanisms that control p53 activity in HSCs and how p53 mutations affect its role in these events may facilitate the development of therapeutic strategies for eliminating leukemia (and cancer) propagating cells.     

New Insight on FGFR3-Related Chondrodysplasias Molecular Physiopathology Revealed by Human Chondrocyte Gene Expression Profiling

Endochondral ossification is the process by which the appendicular skeleton, facial bones, vertebrae and medial clavicles are formed and relies on the tight control of chondrocyte maturation. Fibroblast growth factor receptor (FGFR)3 plays a role in bone development and maintenance and belongs to a family of proteins which differ in their ligand affinities and tissue distribution. Activating mutations of the FGFR3 gene lead to craniosynostosis and multiple types of skeletal dysplasia with varying degrees of severity: thanatophoric dysplasia (TD), achondroplasia and hypochondroplasia. Despite progress in the characterization of FGFR3-mediated regulation of cartilage development, many aspects remain unclear. The aim and the novelty of our study was to examine whole gene expression differences occurring in primary human chondrocytes isolated from normal cartilage or pathological cartilage from TD-affected fetuses, using Affymetrix technology. The phenotype of the primary cells was confirmed by the high expression of chondrocytic markers. Altered expression of genes associated with many cellular processes was observed, including cell growth and proliferation, cell cycle, cell adhesion, cell motility, metabolic pathways, signal transduction, cell cycle process and cell signaling. Most of the cell cycle process genes were down-regulated and consisted of genes involved in cell cycle progression, DNA biosynthesis, spindle dynamics and cytokinesis. About eight percent of all modulated genes were found to impact extracellular matrix (ECM) structure and turnover, especially glycosaminoglycan (GAG) and proteoglycan biosynthesis and sulfation. Altogether, the gene expression analyses provide new insight into the consequences of FGFR3 mutations in cell cycle regulation, onset of pre-hypertrophic differentiation and concomitant metabolism changes. Moreover, impaired motility and ECM properties may also provide clues about growth plate disorganization. These results also suggest that many signaling pathways may be directly or indirectly altered by FGFR3 and confirm the crucial role of FGFR3 in the control of growth plate development.     

Bone histological correlates of high-frequency flapping flight and body mass in the furculae of birds: a phylogenetic approach

A parsimony optimization of the presence of high-frequency flapping flight onto a phylogeny of 29 species of birds shows that this is a derived character state that has been acquired at least four independent times: by the last common ancestor of Alcidae, that of Podicipedidae, that of Anatidae, and that of Rallidae. Cineradiographic analysis has shown that the furculae of birds underwent extraordinary deformations during the wingbeat cycle. Cyclical deformations are known to produce microfractures in the bone tissue, which may be a stimulus for Haversian remodelling, a mechanism of resorption and reconstruction of bone tissue that may repair bone microdamage. In the present study, we performed a comparative analysis in a phylogenetic context to test the effect of the frequency of cyclical deformations and body mass on the rate of Haversian remodelling in the furculae of birds. A variation partitioning analysis showed that the type of flight (high-frequency flapping flight vs. other kinds of flight of lower wing beat frequency) and body mass explained a significant portion of Haversian bone density (the outcome of Haversian remodelling) and that the phylogeny also explained a significant part of this variation. This phylogenetic signal on Haversian bone density variation may be the outcome of phylogenetic signal on the proximate causes producing Haversian remodelling.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 729–738.     

Development of the mouse mandibles and clavicles in the absence of skeletal myogenesis

In this report we employed double-knock-out mouse embryos and fetuses (designated as Myf5-/-: MyoD-/- that completely lacked striated musculature to study bone development in the absence of mechanical stimuli from the musculature and to distinguish between the effects that static loading and weight-bearing exhibit on embryonic development of skeletal system. We concentrated on development of the mandibles (= dentary) and clavicles because their formation is characterized by intramembranous and endochondral ossification via formation of secondary cartilage that is dependent on mechanical stimuli from the adjacent musculature. We employed morphometry and morphology at different embryonic stages and compared bone development in double-mutant and control embryos and fetuses. Our findings can be summarized as follows: a) the examined mutant bones had significantly altered shape and size that we described morphometrically, b) the effects of muscle absence varied depending on the bone (clavicles being more dependent than mandibles) and even within the same bone (e.g., the mandible), and c) we further supported the notion that, from the evolutionary point of view, mammalian clavicles arise under different influences from those that initiate the furcula (wishbone) in birds. Together, our data show that the development of secondary cartilage, and in turn the development of the final shape and size of the bones, is strongly influenced by mechanical cues from the skeletal musculature.     

A probable homologue of the clavicle in the holostean fish Amia calva

The two pairs of appendages associated with the throat region in Amia are described.
The posterior serrated appcndage of Amia calva is considered a homologue of the clavicle. This theory is based on topographical similarity (a bony element situated in front of the ventral part of the cleithrum and overlapping the anterior margin of the adjoining part of the cleithrum), structural similarity (cellular bone with ridges bearing denticles arranged in distinct patterns), ontogenetic similarity (early ontogenctic ossification with other dements of the dermal shoulder girdle prior to the appearance of scales), and phylogenetic cvidcnce (presence of similar elements in the caturid Furo ).
Although the anterior serrated appendage of A. calva exhibits structural and topographical similarities to both scales and clavicles, the ontogenetic evidence favours an homology with scales.
The ossicles located dorsolateral and lateral to the anteroventral part of the cleithrum in Lepisosteus oculatus share structural and topographical similarities to both scales and clavicles. However, ontogenetic evidence indicates that these ossicles are scales rather than fragmented clavicles.     

Age estimation from cortical index of the human clavicle in northwest Indians

The proportion of the cortical thickness to the total diameter of the bone (cortical index) was calculated in paired clavicles obtained from 128 male and 82 female medicolegal postmortem subjects who were apparently healthy prior to their accidental death. The ages of the subjects varied from 15 to 85 years. The clavicles were cut either horizontally or parasagittally, and measurements were taken at midclavicle. From the ages of 15 to 30 years, the cortical index increased. It decreased steadily thereafter, with an initial sharp decrease in the age group 31-40 years in both sexes. After the age of 40 years, this rapid decrease in the index continued in the females, but became slow and gradual in the males. Differences between left and right sides were statistically insignificant in both sexes (P greater than 0.05). However, the sexual differences were significant (P less than 0.01) in the age groups from 41 years onwards.     

Busulfan-induced central polydactyly, syndactyly and cleft hand or foot: a common mechanism of disruption leads to divergent phenotypes

The prevalence of clinical phenotypes that exhibit combinations of central polydactyly, syndactyly, or cleft hand or foot is higher than would be expected for random independent mutations. We have previously demonstrated that maternal ingestion of a chemotherapeutic agent, busulfan, at embryonic day 11 (E11) induces these defects in various combinations in rat embryo limbs. In an effort to determine the mechanism by which busulfan disrupts digital development, we examined cell death by Nile Blue staining and TdT-mediated dUTP nick end labeling (TUNEL) assays; we also carried out whole mount in situ hybridization for fibroblast growth factor-8 (Fgf8), bone morphogenetic protein-4 (Bmp4), and sonic hedgehog (Shh) to examine developmental pathways linked to these defects. In busulfan-treated embryos, diffuse cell death was evident in both ectoderm and mesoderm, peaking at E13. The increased cell death leads to regression of Fgf8 in the apical ectodermal ridge (AER) and Bmp4 and Shh in the underlying mesoderm. The subsequent pattern of interdigital apoptosis and cartilage condensation was variably disrupted. These results suggest that busulfan manifests its teratogenic effects by inducing cell death of both ectoderm and mesoderm, with an associated reduction in tissue and a disruption in the generation of patterning molecules during critical periods of digit specification.