The influence of climate and population structure on East Asian skeletal morphology.

Recent studies have shown that global variation in body proportions is more complex than previously thought as some traits formerly associated with climate adaptation are better explained by geographic proximity and neutral evolutionary forces. While the recent incorporation of quantitative genetic methodologies has improved understanding of patterns related to climate in Africa, Europe, and the Americas, Asia remains underrepresented in recent and historic studies of body form. As ecogeographic studies tend to focus on male morphology, potential sex differences in features influenced by climate remain largely unexplored. Skeletal measurements encompassing the dimensions of the skull, pelvis, limbs, hands, and feet were collected from male (n = 459) and female (n = 442) remains curated in 13 collections across seven countries in East Asia (n = 901). Osteological data were analyzed with sex and minimum temperature as covariates adjusted by autosomal single-nucleotide polymorphism population genetic distance using univariate Bayesian linear mixed models, and credible intervals were calculated for each trait. Analysis supports a relationship between specific traits and climate as well as providing the magnitude of response in both sexes. After accounting for genetic distance between populations, greater association between climate and morphology was found in postcranial traits, with the relationship between climate and the skull limited primarily to breadth measurements. Larger body size is associated with colder climates with most measurements increasing with decreased temperature. The same traits were not always associated with climate for males and females nor correlated with the same intensity for both sexes. The varied directional association with climate for different regions of the skeleton and between the sexes underscores the necessity of future ecogeographic research to holistically evaluate body form and to look for sex-specific patterns to better understand population responses to environmental stresses.

Biomechanical implications of the onset of walking.

Changes in long bone strength associated with the onset of bipedal walking in humans have been previously documented in a longitudinal growth sample. However, it is unclear if this transition can be detected using archaeological, cross-sectional data, which likely encompass more cultural and biological variation than a single dataset of living children. Focusing on variation in cross-sectional polar second moment of area, we evaluate the ratios of femoral, tibial, and humeral strength in seven temporally diverse samples of individuals from birth to the age of eighteen years (n = 501), with subsequent comparisons to immature Late Pleistocene fossils. Using these samples, we determine whether changes related to the developmental onset of bipedality can be detected in a large, multi-population sample, test for differences in long bone strength ratios among Holocene groups that may indicate developmental differences in the onset of walking, and determine whether immature Late Pleistocene samples follow the same patterns as modern humans. Despite great variation within the Holocene sample, clear changes in these ratios are apparent around the age of the onset of walking. Humeral-to-femoral strength increases briefly prior to the age of one, with a sharp decline in relative humeral strength thereafter until age four. A similar pattern is apparent in the ratio of humeral/tibial and femoral/tibial strength. While the general pattern is consistent across all human groups sampled, these ratios vary by skeletal population, which seems to be closely related to variation in tibial length among samples. Although the extremely small fossil sample makes differences difficult to interpret, Neandertals also differ from both Late Pleistocene and Holocene modern humans in their strength ratios. Further research in this area may provide additional information about the skeletal impact of the onset of walking in the past and in additional fossil taxa.

Femoral neck-shaft angle and climate-induced body proportions.

OBJECTIVES: Declination in femoral neck-shaft angle (NSA) is commonly linked to an increased level of physical activity during life. More recently, however, research suggests that lower NSA might also be explained, in part, as the mechanical consequence of differences in ecogeographic body proportions. This study tests the proposed link between NSA and climatic-induced body proportions, using relative body mass (RBM), throughout the course of development. MATERIALS AND METHODS: NSA and RBM were collected for 445 immature remains from five geographic locations. NSA and RBM were standardized for age-effects. ANOVA was used to examine when population differences emerged in both NSA and RBM. Regression analyses were used to examine the pattern of relationship between NSA and RBM. RESULTS: Populations differ significantly in NSA and RBM before skeletal maturity, and these differences occur early in life. While both NSA and RBM change over the course of development, no significant relationship was found between NSA and RBM for any sample, or any age category (p = .244). DISCUSSION: Individuals who have relatively greater relative body mass do not necessarily have lower NSA. Population differences in NSA were found to be variable, while differences in RBM remained consistent across the developmental span. Taken together, these results suggest that regardless of body proportions, the degree of declination of NSA is presumed to be similar among individuals with similar gait and ambulatory behaviors. Conversely, populations differ in RBM from birth, and these differences are consistent throughout development. These two measures likely are responsive to diffing stimuli, and any potential relationship is likely complex and multifactorial.

The path less traveled: Using structural equation modeling to investigate factors influencing bone functional morphology.

OBJECTIVES: The relationship between an organism's mechanical environment and its bone strength has been long established by experimental research. Multiple intrinsic and extrinsic factors, including body mass, muscle strength, genetic background, and nutritional and/or hormonal status, are likely to influence bone deposition and resorption throughout the lifespan, complicating this relationship. Structural equation modeling (SEM) is uniquely positioned to parse this complex set of influences. MATERIALS AND METHODS: Data from the Third National Health and Nutrition Examination Survey, including sex, total body mass, lean body mass, exercise frequency, peak body mass, and age, were analyzed using SEM to determine how they affect bone strength both individually and combined. RESULTS: Body mass is typically the driver of cross-sectional area, but body mass and lean mass have similar effects on the polar moment of area (J). Peak body mass had a strong direct effect on J, despite decreasing strongly with increases in lean mass. Exercise also did not confer a large direct effect on cross-sectional area or J but did modify body mass and lean mass. In females, intentional weight loss was associated with decreased exercise levels. DISCUSSION: SEM is a useful tool for parsing complex systems in bone functional morphology and has the potential to uncover causal links in the study of skeletal remodeling, including factors like weight loss or exercise that may have secondary effects.

Developmental variation in ecogeographic body proportions.

While ecogeographic variation in adult human body proportions has been extensively explored, relatively less attention has been paid to the effect of Bergmann's and Allen's rules on human body shape during growth. The relationship between climate and immature body form is particularly important, as immature mortality is high, mechanisms of thermoregulation differ between young and mature humans, and immature body proportions fluctuate due to basic parameters of growth. This study explores changes in immature ecogeographic body proportions via analyses of anthropometric data from children included in Eveleth and Tanner's (1976) Worldwide Variation in Human Growth, as well as limb proportion measurements in eight different skeletal samples. Moderate to strong correlations exist between climatic data and immature stature, weight, BMI, and bi-iliac breadth; these relationships are as strong, if not stronger, in immature individuals as they are in adults. Correlations between climate and trunk height relative to stature are weak or nonexistent. Altitude also has significant effects on immature body form, with children from higher altitudes displaying smaller statures and lower body weights. Brachial and crural indices remain constant over the course of growth and display consistent, moderate correlations with latitude across ontogeny that are just as high as those detected in adults. The results of this study suggest that while some features of immature body form, such as bi-iliac breadth and intralimb indices, are strongly dictated by ecogeographic principles, other characteristics of immature body proportions are influenced by intrinsic and extrinsic factors such as nutrition and basic constraints of growth.

Ontogeny of limb proportions in late through final Jomon period foragers.

This study reports on developmental patterning in the intralimb indices of Late/Final Jomon period (4000-2300 BP) people. Jomon foragers represent the descendants of migrants from Northeast Asia, who arrived in the Japanese Islands around 20,000 BP. Among adults, Jomon brachial indices are elevated and similar to warm adapted, low latitude people, while crural indices are intermediate and similar to people from moderate latitudes. Two hypotheses regarding the development of intralimb indices among Jomon period foragers are tested: (1) intralimb indices of Jomon people maintain predicted ecogeographic relationships over ontogeny; (2) greater evolvability will be observed in the brachial index, while greater developmental constraint will be observed in the crural index. Changes in intralimb proportions in a Jomon skeletal growth series are compared to those in two contrasting samples: Inuit from Point Hope (cold adapted) and Nubians from Kulubnarti (warm adapted). A quadratic equation best describes the ontogeny of brachial and crural indices, with high indices in infancy followed by a decline in childhood and an increase in adolescence. Despite these shifts, ecogeographically predicted differences and similarities in the indices are maintained between samples throughout ontogeny. In addition, radial relative to humeral length is significantly less correlated than tibial relative to femoral length. These results suggest genetic conservation of intralimb indices over the course of development. However, radial and humeral lengths are less correlated than tibial and femoral lengths among Jomon subadults and adults, potentially suggesting greater evolvability of the brachial index and more developmental constraint on the crural index.

The ontogeny of Holocene and Late Pleistocene human postcranial strength.

While a wide variety of studies have focused on population variation in adult cross-sectional properties, relatively little is known about population variation in postcranial robusticity in immature individuals. Furthermore, the age at which the population differences readily detected in adults manifest during growth is also unknown. This research addresses these gaps in our current understanding through the analysis of immature humeral and femoral long bone strength. Cross-sectional geometry was used to compare the developmental trajectories of diaphyseal strength in Late Pleistocene Neandertal and modern human subadults to a sample of immature humans from seven geographically diverse Holocene populations. Population differences in size-standardized cross-sectional properties appear to be systemic and develop very early in ontogeny in the Holocene sample. In many cases, these differences are present before one year of age. In general, the Late Pleistocene fossil samples fit within the range of recent human variation in long bone strength. Population differences detected here are likely related to a combination of factors including activity patterns, genetic propensities, and nutritional status. These results highlight the complex mosaic of processes that result in adult postcranial robusticity, and suggest that further exploration of the developmental interplay between intrinsic and extrinsic influences on skeletal robusticity will likely enhance our understanding of adult postcranial morphology.

Determination of sex from the hyoid bone.

This article explores size differences related to sex in the hyoid bones from the Robert J. Terry Anatomical Collection. A series of measurements were taken from 398 hyoids, both fused and unfused. The inclusion of unfused hyoids in the study provides the opportunity to investigate previously unknown size differences between sexes as well as to determine their utility in determining sex. Two-way ANOVA was used to explore differences in hyoid size as related to ancestry and sex. Discriminant function analysis was employed to test the ability of the hyoids to be classified by sex. Six discriminant function equations ranging in accuracy from 82% to 85% are provided, each of which is more accurate than many of the discriminant functions developed in past hyoid research, are simple to use, and can be used to estimate the sex of a hyoid regardless of its state of fusion. In addition to providing further information about the morphological form of the hyoid, these analyses provide a method that can be easily employed to assess sex of the individual from the hyoid bone.

Waddling and toddling: the biomechanical effects of an immature gait.

Femoral shape changes during the course of human growth, transitioning from a subcircular tube to a teardrop-shaped diaphysis with a posterior pilaster. Differences between immature and mature bipedalism and body shape may generate different loads, which, in turn, may influence femoral modeling and remodeling during the course of the human lifespan. This study uses two different approaches to evaluate the hypotheses that differences in gait between young and mature walkers result in differences in ground reaction forces (GRFs) and that the differences in loading regimes between young children and adults will be reflected in the geometric structure of the midshaft femur. The results of this analysis indicate that GRFs differ between young walkers and adults in that normalized mediolateral (ML) forces are significantly higher in younger age groups. In addition, these differences between children and adults in the relative level of ML bending force are reflected in changes in femoral geometry during growth. During the earlier stages of human development, immature femoral diaphyses are heavily reinforced in approximately ML plane. The differences in gait between mature and immature walkers, and hence the differences in femoral shape, are likely partially a product of a minimal bicondylar angle and relatively broad body in young children.

Humeral torsion revisited: a functional and ontogenetic model for populational variation.

Anthropological interest in humeral torsion has a long history, and several functional explanations for observed variation in the orientation of the humeral head have been proposed. Recent clinical studies have revived this topic by linking patterns of humeral torsion to habitual activities such as overhand throwing. However, the precise functional implications and ontogenetic history of humeral torsion remain unclear. This study examines the ontogeny of humeral torsion in a large sample of primarily immature remains from six different skeletal collections (n = 407). The results of this research confirm that humeral torsion displays consistent developmental variation within all populations of growing children; neonates display relatively posteriorly oriented humeral heads, and the level of torsion declines steadily into adulthood. As in adults, variation in the angle of humeral torsion in immature individuals varies by population, and these differences arise early in development. However, when examined in the context of the developing muscles of the shoulder complex, it becomes apparent that variation in the angle of humeral torsion is not necessarily related to specific habitual activities. Variability in this feature is more likely caused by a generalized functional imbalance between muscles of medial and lateral rotation that can be produced by a wide variety of upper limb activity patterns during growth.

Exploring ancestral variation of the hyoid.

This study utilizes metric analysis to examine size and shape variation between hyoids of Africans and Europeans in the Robert J. Terry Anatomical Collection. A total of 200 fused and unfused hyoids were measured and three statistical methods were employed to explore variation between ancestries. First, independent sample t-tests showed that some significant size differences do occur between ancestries. Second, to examine shape variation, skeletal measurements were regressed on the geometric mean using least squares linear regression with the residuals used to evaluate size-corrected shape differences. Finally, discriminant function analysis was used to develop two functions for ancestry prediction with overall accuracies of 73% and 77%. Results of the analyses suggest hyoid size and shape differences do occur between ancestries, notably that European hyoids are broader than African hyoids, while the African hyoid is longer than Europeans.

Shanidar 10: a Middle Paleolithic immature distal lower limb from Shanidar Cave, Iraqi Kurdistan.

The analysis of the faunal remains from Shanidar Cave has identified an incomplete immature human distal leg and foot from the deepest levels of the Middle Paleolithic of Shanidar Cave, Iraq. The distal tibia, fibula, first metatarsal, and two tarsals, designated Shanidar 10, derive from a 1-2-year-old infant. The tibia exhibits a transverse line from a stress episode during the last quarter of its first year postnatal. The cross-sectional geometry of the tibial midshaft reveals modest cortical thickening and a level of diaphyseal robusticity similar to those of recent human infants of a similar developmental age.