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.     

Sexual differences in Neanderthal limb bones

European and Near Eastern Neanderthal postcranial remains have been analyzed to determine the degrees of sexual dimorphism in limb bone size and robusticity present among the Neanderthals. The remains were sexed on the basis of pelvic morphology where possible (seven males and three females) and otherwise on the basis of absolute size employing limb bone lengths and articular dimensions (12 males and 15 females). Neanderthal sexual size dimorphism, both within single site samples and in the total sexable sample, is virtually the same as that of recent human samples. Furthermore, despite a tendency towards more robust limbs, the Neanderthals exhibit sexual dimorphism in limb bone shaft and articular robusticity similar to that of recent human samples. By the time of the Neanderthals, sexual dimorphism in limb bone size and robusticity appears to have reached recent human proportions.     

Bilateral asymmetry of the humerus during growth and development

The development of handedness throughout growth can be investigated by using bilateral asymmetry of the humerus as a proxy for this trait. A large skeletal sample of nonadults from English archaeological sites was examined using standard metric techniques to assess when right-sided asymmetry first appears in the human skeleton. Results of this work indicate a change in directional asymmetry during growth and development, with infants and young children exhibiting no significant asymmetry and older children and adolescents demonstrating right-sidedness. This trend is consistent with what has been observed in previous studies of upper limb asymmetry in skeletal material and behaviorally in living children, adding further strength to the premise that biomechanical forces strongly influence bilateral asymmetry in the upper limb bones. Variability in the magnitude of asymmetry between different features of the humerus was also noted. This characteristic can be explained by differing degrees of genetic canalization, with length and articular dimensions being more strongly canalized than diaphyseal properties.     

Postcranial robusticity in Homo. II: Humeral bilateral asymmetry and bone plasticity

The analysis of humeral asymmetry in Recent human skeletal samples and an extant tennis-player sample documents minimal asymmetry in bone length, little asymmetry in distal humeral articular breadth, but pronounced and variable asymmetry in mid- and distal diaphyseal crosssectional geometric parameters. More specifically, skeletal samples of normal modern Euroamericans, prehistoric and early historic Amerindians, and prehistoric Japanese show moderate (ca. 5–14%) median asymmetry in diaphyseal cross-sectional areas and polar second moments of area, whereas the tennis-player sample, with pronounced unilateral physical activity, exhibits median asymmetries of 28–57% in the same parameters. A sample of Neandertals with nonpathological upper limbs exhibits similarly low articular asymmetry but pronounced diaphyseal asymmetries, averaging 24–57%. In addition, three Neandertals with actual or possible post-traumatic upper limb alterations have the same low articular asymmetry but extremely high diaphyseal asymmetries, averaging 112–215%. These data support those from experimental work on animals, exercise programs of humans, and human clinical contexts in establishing the high degree of diaphyseal plasticity possible for humans, past and present, under changing biomechanical loading conditions. This lends support to activity-related functional interpretations of changing human diaphyseal morphology and robusticity during the Pleistocene. © 1994 Wiley-Liss, Inc.     

Bilateral asymmetry in the limb bones of the chimpanzee (Pan troglodytes)

There is much debate in behavioral primatology on the existence of population-level handedness in chimpanzees. The presence or absence of functional laterality in great apes may shed light on the origins of human handedness and on the evolution of cerebral asymmetry. The plasticity of long bone diaphyses in response to mechanical loading allows the functional interpretation of differences in cross-sectional geometric. While left-right asymmetry in upper limb diaphyseal morphology is a known property in human populations, it remains relatively unexplored in apes. We studied bilateral asymmetry in 64 skeletons of wild-caught chimpanzee using the humerus, second metacarpal, and femur. The total subperiosteal area (TA) of the diaphyses was measured at 40% of maximum humeral length and at the midshaft of the metacarpals and femora using external silicone molds. Overall, the TA values of the left humeri were significantly greater than the right, indicating directional asymmetry. This effect was even greater when the magnitude of difference in TA between each pair of humeri was compared. The right second metacarpals showed a tendency toward greater area than did the left, but this did not reach statistical significance. The lack of asymmetry in the femur serves as a lower limb control, and suggests that the upper limb results are not a product of fluctuating asymmetry. These findings imply behavioral laterality in upper limb function in chimpanzees, and suggest a complementary relationship between precision and power.     

Dental arch form in the cercopithecidae

The dental arches of the major genera of Old World monkeys (superfamily Cercopithecoidea) were studied by morphometric techniques. Bicanine and bimolar breadths and arch lengths were ascertained for maxillary and mandibular arches. This data was then subjected to a variety of statistical tests. Corresponding arch dimensions of the upper and lower dentition showed the highest correlations, while the lowest correlations were generally observed between comparisons of arch dimensions and body size. A new expression was developed, relative male palate size (RMPS), which quantified the degree of sexual dimorphism while correcting for body size. The four hierarchies examined using RMPS values were sexual dimorphism, modes of sexual selection and predator defense and diet. Maxillary bicanine breadth was the only parameter that exhibited sexual dimorphism consistently in each of the four hierarchies, although differences in arch size were identified for diet and predator defense. Species grouped by predator defense showed the most sexual dimorphism in arch parameters.     

Activity patterns in the Sahara Desert: an interpretation based on cross-sectional geometric properties

The Garamantian civilization flourished in modern Fezzan, Libya, between 900 BC and 500 AD, during which the aridification of the Sahara was well established. Study of the archaeological remains suggests a population successful at coping with a harsh environment of high and fluctuating temperatures and reduced water and food resources. This study explores the activity patterns of the Garamantes by means of cross-sectional geometric properties. Long bone diaphyseal shape and rigidity are compared between the Garamantes and populations from Egypt and Sudan, namely from the sites of Kerma, el-Badari, and Jebel Moya, to determine whether the Garamantian daily activities were more strenuous than those of other North African populations. Moreover, sexual dimorphism and bilateral asymmetry are assessed at an intra- and inter-population level. The inter-population comparisons showed the Garamantes not to be more robust than the comparative populations, suggesting that the daily Garamantian activities necessary for survival in the Sahara Desert did not generally impose greater loads than those of other North African populations. Sexual dimorphism and bilateral asymmetry in almost all geometric properties of the long limbs were comparatively low among the Garamantes. Only the lower limbs were significantly stronger among males than females, possibly due to higher levels of mobility associated with herding. The lack of systematic bilateral asymmetry in cross-sectional geometric properties may relate to the involvement of the population in bilaterally intensive activities or the lack of regular repetition of unilateral activities.     

Skeletal indicators of subsistence patterns and activity régime in the Mesolithic sample from Grotta dell’Uzzo (Trapani,Sicily): a case study

The skeletal sample from Grotta dell’Uzzo, Sicily (2 adult females and 5 adult males) was compared to a number of more representative population samples from Western Europe and the Mediterranean Basin. The majority of these were from Italian pre- and protohistoric sites. The research protocol analyzed skeletal indicators of labour activity and sexual division of labour (body size and proportions, sexual dimorphism, limb lateralization, bone robustness, the development of muscular attachments, accessory articular facets, signs of muscular hyperfunction). Sexual dimorphism and limb lateralization showed some regular patterns of possible general significance in all the samples examined here. A general pattern of gracilization and de-specialization of physical activity is observed in the Mesolithic as compared to the Upper Palaeolithic samples. The main features of the Mesolithic samples are: a reduction of body size and bone robustness, a lower degree of sexual dimorphism and limb bone asymmetry, a reduction of the mechanical stress on the lower limbs indicated by less pronounced muscular attachments and reduced talar flattening. This trend is reversed towards the Neolithic period. The main features of these variations are discussed in relation to economic and environmental changes. The Uzzo sample fits well into the general picture of the Western European Mesolithic, although showing some intermediate features between the Mesolithic and the Neolithic samples. This paper is dedicated to the memory of the late Roland Menk, who made significant contribution to our understanding of the Mesolithic transition.     

Bone structural data for the Denver longitudinal growth study

The Denver Longitudinal Growth Study is one of the most comprehensive longitudinal studies of growth carried out to date. Measurements of long bone lengths, cross-sectional diaphyseal properties, and articular breadths were taken from archived radiographs of 20 study subjects measured at an average of 36 time points from early infancy to late adolescence. Anthropometric dimensions for these individuals were also obtained from study archives. These data are now available in an online data set: 10.5061/dryad.xsj3tx9gq .     

Study on long bones: variation in angular traits with sex, age, and laterality

The biological problems considered in this study are sexual dimorphism in angular osteometric traits, changes in these traits with aging and their bilateral asymmetry in the same individual. The sample comprises the right and left long bones of 200 skeletons (100 males and 100 females) of Sardinian adults (Frasetto collection) for whom the sex and age at death are known. As concerns the sexual dimorphism, some angular traits of upper limb (joint axis angle of the ulna in particular) and the lower limb (condylo-diaphyseal angle of the femur and retroversion angle of the tibia) are significantly different between males and females. Torsion angles of long bones (especially the femur) were found to change with age. In sides comparison revealed a clear right-left asymmetry in our sample. The asymmetry, sexual dimorphism and changes with age are discussed in relation to varying stresses of different living activities. Our data indicate that the angular traits of long bones could be very informative in biomechanical interpretations of human activities based on skeletal remains.     

Postcranial adaptations for climbing in Loridae (Primates)

The Loridae are an arboreal family of small primates that are specialized for slow and quiet climbing. This paper examines the relationship between lorid locomotory behaviour and postcranial skeletal morphology. Lorid humeral and femoral diaphyseal geometric cross-sectional properties, articular surface areas, and lengths are compared to those properties in other small primates with less specialized locomotory behaviour. The comparative sample includes both closely related prosimians and more distantly related platyrrhines.
Results indicate that lorids have greater humeral and femoral diaphyseal rigidity than other quadrupedal primates of similar body size, suggesting that lorid limbs are subjected to greater forces. Lorids also have relatively larger humeral and femoral articulations, corresponding to field and laboratory observations which indicate that lorid joints are highly mobilc. In addition, lorids have long humeri relative to femoral length, and compared to humeral length in less specialized prosimians of similar body mass. Long humeral length relative to femoral length is interpreted as a climbing adaptation because similar limb proportions are also seen in many non-primate climbers. Altogether, humeral and femoral diaphyseal cross-sectional properties, articular surface areas, and lengths comprise a suite of characters which have potential for identifying climbing specialists in the fossil record.     

新疆敏麻蜥3个地理种群趾长及趾长比的两性异形

趾长比是指动物不同趾长的比值,常被认为是早期雄激素或雌激素暴露的标记物之一.本文通过测量新疆敏麻蜥Eremias arguta 3个地理种群的雌雄成体各趾长并计算趾长比,探究不同地理种群趾长及趾长比的两性差异.结果表明,伊犁地区雄性后肢的第2趾显著大于雌性(P<0.05);富蕴县雄性前肢、后肢的第2、3、4、5趾长均显...     

Odontometric profile of a prehistoric southeastern Florida population

This study analyzes and compares the dental size variation and sexual dimorphism of the Highland Beach Mound people, a late southeastern prehistoric aboriginal Florida population (ca. 800-1200 AD) with a somewhat earlier midwestern group (ca. 200-800 AD) and an Archaic sample (ca. 4160-2558 BC). The lengths and breadths of maxillary and mandibular permanent teeth of 99 individuals were measured, and crown surface areas were calculated for the molars. Results indicated that there was little sexual dimorphism. Only the mesiodistal dimensions of the upper P2 and lower C were significantly different at the P less than 0.05 level. In terms of dental size, what appears to be a chronologically based reduction in most points of comparison was seen in all three groups, with the exception of the molars, which were larger in the Highland Beach sample than in the earlier Schultz Mound.     

Divergence in Male and Female Manipulative Behaviors with the Intensification of Metallurgy in Central Europe

Humeral morphology has been shown to reflect, in part, habitual manipulative behaviors in humans. Among Central European agricultural populations, long-term social change, increasing task specialization, and technological innovation all had the potential to impact patterns of habitual activity and upper limb asymmetry. However, systematic temporal change in the skeletal morphology of agricultural populations in this region has not been well-characterized. This study investigates diachronic patterns in humeral biomechanical properties and lengths among 174 adult Central European agriculturalists through the first ∼5400 years of farming in the region. Greater asymmetry in biomechanical properties was expected to accompany the introduction of metallurgy, particularly in males, while upper limb loading patterns were expected to be more similar between the Bronze and Iron Ages. Results revealed a divergence in the lateralization of upper limb biomechanical properties by sex between the Early/Middle Neolithic and Early/Middle Bronze Age. Neolithic females had significantly more variable properties than males in both humeri, while Bronze Age female properties became homogeneous and very symmetrical relative to the right-biased lateralization of contemporaneous males. The Bronze Age to Iron Age transition was associated with morphological change among females, with a significant increase in right-biased asymmetry and a concomitant reduction in sexual dimorphism. Relative to biomechanical properties, humeral length variation and asymmetry were low though some significant sexual dimorphism and temporal change was found. It was among females that the lateralization of humeral biomechanical properties, and variation within them, changed most profoundly through time. This suggests that the introduction of the ard and plow, metallurgical innovation, task specialization, and socioeconomic change through ∼5400 years of agriculture impacted upper limb loading in Central European women to a greater extent than men.     

HANDEDNESS IN THE BROWN-THROATED PARAKEET ARATINGA PERTINAX EST RELATION WITH SKELETAL ASYMMETRY

Observations on 56 specimens of Aratinga pertinax when bringing food to the beak prove that 28 birds were right-handed while the other 28 were left-handed. A biometric analysis reveals a slight departure from bilateral symmetry in hindlimb bones in close relationship with handedness. In the right-handed birds, there is a significant predominance in the length of the right hindlimb as a whole and all right limb segments. The opposite holds true in left-handed parrots (except for the femur). Right-handed birds are in general more highly asymmetrical than left-handed ones. There is a positive correlation between asymmetry of homologous segments of a pair of limbs and the asymmetry of the other segments of the same pair of limbs. The absence of significant negative correlations between bilateral differences of the different segments of a limb indicates that departures from bilateral symmetry tend to affect the limb as a whole, and thus is not in agreement with the rule of compensating variations.     

Heat stress, fluctuating asymmetry and prenatal selection in the laboratory rat

Recent studies have demonstrated that stress may increase the fluctuating asymmetry of teeth and limbs in laboratory animals. The present study investigates the effects of heat on such parameters. Pregnant laboratory rats are exposed to temperatures of 33 degrees C during gestation. Increases in fluctuating asymmetry of dental dimensions as well as bone density are found for the pups of such females when compared to unstressed controls. A general increase in limb lengths is also reported for young of heat stressed animals. Prenatal selection with differential survival is suggested as a possible explanation for differences found between the experimental and control animals.     

不同球类运动对大学生上下肢骨密度及其方向性不对称的影响

为探究不同球类运动对上下肢骨密度及其方向性不对称的影响,特选取研究对象共210人,均分为7组,即篮球组、排球组、足球组、羽毛球组、乒乓球组、网球组和对照组,测量其上下肢的骨密度和方向性不对称性(DA)。结果显示,篮球组、排球组、羽毛球组、乒乓球组和网球组的上肢骨密度都是右侧明显高于左侧,下肢骨密度仅足球组表现为右侧高于左侧;不同组间的上下肢骨密度和DA具有统计学差异。结果还显示,左侧上肢骨密度的组间排序为排球组>足球组>第二亚组(乒乓球组、网球组和羽毛球组)>对照组,篮球组>乒乓球组、网球组和羽毛球组>对照组;右侧上肢骨密度的排序为网球组>乒乓球组、排球组和篮球组>足球组>对照组,乒乓球组>羽毛球组>足球组>对照组;左侧下肢骨密度的排序为足球组>篮球组、排球组、羽毛球组、乒乓球组和网球组>对照组;右侧下肢骨密度的排序为足球组>排球组、篮球组、网球组、羽毛球组和乒乓球组>对照组,排球组>羽毛球组和乒乓球组>对照组;DA的排序为网球组>乒乓球组、羽毛球组、篮球组和排球组,乒乓球组&g...     

The dentition of the Indian Knoll skeletal population: odontometrics and cusp number.

Data on the permanent dentition of 153 individuals from the well known Indian Knoll skeletal population are presented. Mesiodistal and buccolingual measurements were taken with a Helios dial caliper. Cusp number of maxillary and mandibular molars are recorded. The Indian Knoll dentition is larger than many modern groups but smaller than Australoid or Mesolithic groups. With the exception of maxillary 12, males have larger teeth than females in both dimensions. The lower canine is the most dimorphic tooth. Through rank order correlation, an association was shown between the sexual dimorphism of the mesiodistal and buccolingual dimensions. Compared to modern groups, the Indian Knoll population displays a moderate degree of sexual dimorphism in tooth size. In general, the coefficients of variation were greater for the more distal teeth within morphological classes. Amounts of size variability did not differ significantly between the sexes; moreover, rank order correlations indicated that patterns of variability in both dimensions were similar for males and females. The predominant cusp number pattern for upper molars is 4-3-3 and for lowers 5-5(4)-5. No sex differences were shown for cusp occurrence or bilateral asymmetry in cusp number.     

The development of asymmetry: the sidedness of drug-induced limb abnormalities is reversed in situs inversus mice

We are studying the development of handedness, in particular the relationships between handed structures with bilateral symmetry, for example the limbs, and those with lateral asymmetry, such as the heart, lungs and gut. Asymmetric (unilateral) developmental limb abnormalities can be induced by chemical treatment of mouse embryos, either in utero by acetazolamide, or in culture by misonidazole. We have examined these effects in mice homozygous for the iv gene. The development of bilateral symmetry in iv/iv mice is normal, but the control of asymmetry appears to be random, that is 50% develop normally (situs solitus), 50% with laterally inverted viscera (situs inversus). We find that the handedness of induced asymmetric limb defects is highly correlated with embryonic visceral situs. Right limb defects are induced in situs solitus embryos, left-sided defects in situs inversus. This suggests that the mechanism of induction of asymmetric defects is not related to any intrinsic difference between the development of left and right limbs, but is connected to visceral asymmetry. In addition, the high correlation of limb defects with situs was observed in culture as well as in utero suggesting that the maternal environment plays no role in the development of asymmetry.     

Body Mass Estimates in Extinct Mammals from Limb Bone Dimensions: the Case of North American Hyaenodontids

The body mass estimation of several limb bone dimensions (shaft cross-sectional properties, articular sizes, and bone lengths) were examined using bivariate linear regression analyses. The sample included taxonomically and behaviourally diverse small to medium-sized Recent carnivorans and carnivorous marsupials. All examined limb bone dimensions indicated low errors (percentage standard error of estimate, 8–13) for the body mass estimations. Among them, humeral and femoral shaft properties correlated best with body weight, while limb bone lengths gave larger errors. Both humeral and femoral head dimensions have relatively large individual variations, and distal humeral articular dimensions seem to be influenced more by phylogenetic differences. The regressions based on each locomotor group gave slightly lower errors than those based on the total pooled sample. The results were then applied to hyaenodontid creodonts from the Eocene–Oligocene of North America. The estimated body masses (kg) are: Arfia , 5.4–9.5; Prototomus , <6.0; Pyrocyon , 2.6; Sinopa , 1.3–1.4; Tritemnodon , 7.6–13; Prolimnocyon , 1.6; Thinocyon , 0.7–2.5; Machaeroides , 12; Limnocyon , 7.8– 16; Hyaenodon , 9.1–43. The various limb bone dimensions give different body mass values, but the variation in estimates is smaller compared to those derived from dental or cranial measurements.