Intensive terrestrial or marine locomotor strategies are associated with inter- and intra-limb bone functional adaptation in living female athletes.

OBJECTIVES: To systematically characterize intra-limb patterns of skeletal plasticity to loading among living women, in order to better understand regional complexity in structural adaptation within the lower limb and more accurately infer behavior in the past. MATERIALS AND METHODS: We used peripheral quantitative computed tomography imaging of the femur, tibia, first and second metatarsals to quantify bone morphology among female controls and athletes representative of either terrestrial or marine mobility, grouped by loading category (odd-impact, repetitive low-impact, and high-magnitude). Parameters included midshaft bone density, areas, rigidity, and shape, epiphyseal bone densities and areas. We assessed between-group differences and the influence of training history on significant variation among the loading groups. RESULTS: Terrestrial mobility strategies were best distinguished by significant midshaft periosteal hypertrophy across the lower limb/foot relative to controls, and by particularly high midshaft femoral and tibial cortical bone areas relative to rowers. Enhanced midshaft bone area was typically paired with decreased bone density among athlete groups. Sport-specific variation in training duration/timing was significantly correlated with multiple midshaft parameters. DISCUSSION: Results demonstrate characteristic patterns of intra-limb adaptation to terrestrial and marine mobility strategies among active women relative to controls, and highlight components of these patterns that may be shaped in part by differences in loading duration/timing. Additionally, our findings support constraints on skeletal variation in the distal tibia and foot relative to more proximal locations about the knee among living women. For example, metatarsal variation was constrained, but where present reflected sport-specific variation in force distribution in the foot.

Body size and body composition effects on heat loss from the hands during severe cold exposure.

OBJECTIVES: This study investigated the influence of body size and composition on maintaining hand temperature during severe cold exposure. The hand's high surface area-to-volume ratio predisposes the hand to heat loss, increasing the risk of cold injury and even hypothermia, which are major selective pressures in cold environments. While vasoregulation may reduce heat loss from the hand, the effect of body form, tissue thermogenesis, and body insulation on heat loss is unknown. MATERIALS AND METHODS: Thermal imaging was used to determine heat loss during a 3-min ice-water hand immersion test carried out on 114 volunteers (female = 63, male = 51). Established anthropometric measures were used to quantify body size, and bioelectrical impedance analysis determined skeletal muscle and fat mass. RESULTS: Skeletal muscle mass relative to body mass was a highly significant predictor of heat loss, while body mass, fat mass, and stature were not. Body composition and body size had little to no significant influence during rewarming after immersion. DISCUSSION: The thermogenic properties of muscle mass support maintenance of hand temperature during severe cold exposure. The findings here suggest that muscular individuals are less susceptible to heat loss and cold injury, and may be better at manual tasks in cold conditions than nonmuscular individuals.

The thermoregulatory function of the human hand: How do palm and digit proportions affect heat loss?

OBJECTIVES: The current study assessed whether ecogeographical patterns seen in hand proportions correlate with heat loss directly. Using a brief severe cold immersion experiment on the hand, the influence of hand and digit dimensions on heat loss was evaluated. MATERIALS AND METHODS: A sample of 113 living individuals were tested. Two-dimensional and three-dimensional scanning techniques were used to assess hand and digit dimensions. Thermal imaging analysis was used to quantify heat loss during a 3-min ice-water immersion of the hands. RESULTS: When body size was accounted for, hand width and digit length relative to total hand length were significant predictors of heat loss from the hand. DISCUSSION: The current study provides empirical evidence to support the link between thermodynamic principles relating to surface area-to-volume ratio, and ecogeographical patterns associated with temperature.

The influence of digit size and proportions on dexterity during cold exposure.

OBJECTIVES: The current study investigated whether size and proportions of the hands and digits affect dexterity during severe cold exposure. As wide hands are known to lose less heat than narrow hands, and narrow digits are associated with greater dexterity, this study aimed to test whether there was a direct trade-off between dexterity and thermoregulation that shapes hand morphology. METHODS: Participants (25 women, 15 men) carried out the Purdue Pegboard test before and after a 3-min ice-water immersion of the hand. Their hand length, hand width, digit lengths, and digit widths were measured using standard anthropometric methods. RESULTS: Wide first and third digits associated with significantly reduced dexterity after immersion relative to individuals with narrower first and third digits. Second digit width positively correlated with average digit temperature after immersion. Hand length and hand width did not influence dexterity. CONCLUSION: The current study suggests that digit width influences dexterity in cold conditions, reflecting patterns found at room temperature. Hand and digit morphology may be the product of two significant constraints on the hand: dexterity and thermoregulation. In cold conditions, hand morphology appears to be predominantly constrained by thermal stress, at the expense of dexterity. This may have important implications for interpreting the morphology of extinct and extant hominins.

Relationship between body mass, lean mass, fat mass, and limb bone cross-sectional geometry: Implications for estimating body mass and physique from the skeleton.

OBJECTIVES: Estimating body mass from skeletal dimensions is widely practiced, but methods for estimating its components (lean and fat mass) are poorly developed. The ability to estimate these characteristics would offer new insights into the evolution of body composition and its variation relative to past and present health. This study investigates the potential of long bone cross-sectional properties as predictors of body, lean, and fat mass. MATERIALS AND METHODS: Humerus, femur and tibia midshaft cross-sectional properties were measured by peripheral quantitative computed tomography in sample of young adult women (n = 105) characterized by a range of activity levels. Body composition was estimated from bioimpedance analysis. RESULTS: Lean mass correlated most strongly with both upper and lower limb bone properties (r values up to 0.74), while fat mass showed weak correlations (r ≤ 0.29). Estimation equations generated from tibial midshaft properties indicated that lean mass could be estimated relatively reliably, with some improvement using logged data and including bone length in the models (minimum standard error of estimate = 8.9%). Body mass prediction was less reliable and fat mass only poorly predicted (standard errors of estimate ≥11.9% and >33%, respectively). DISCUSSION: Lean mass can be predicted more reliably than body mass from limb bone cross-sectional properties. The results highlight the potential for studying evolutionary trends in lean mass from skeletal remains, and have implications for understanding the relationship between bone morphology and body mass or composition.

Declining tibial curvature parallels ∼6150 years of decreasing mobility in Central European agriculturalists.

Long bones respond to mechanical loading through functional adaptation in a suite of morphological characteristics that together ensure structural competence to in vivo loading. As such, adult bone structure is often used to make inferences about past behavior from archaeological remains. However, such biomechanical approaches often investigate change in just one aspect of morphology, typically cross-sectional morphology or trabecular structure. The relationship between longitudinal bone curvature and mobility patterns is less well understood, particularly in the tibia, and it is unknown how tibial curvature and diaphyseal cross-sectional geometry interact to meet the structural requirements of loading. This study examines tibial curvature and its relationship with diaphyseal cross-sectional geometry (CSG) and body size in preindustrial Central Europeans spanning ∼6150 years following the introduction of agriculture in the region. Anteroposterior centroid displacement from the proximo-distal longitudinal axis was quantified at nine diaphyseal section locations (collectively representative of diaphyseal curvature) in 216 tibial three-dimensional laser scans. Results documented significant and corresponding temporal declines in midshaft centroid displacement and CSG properties. Significant correlations were found between mid-diaphyseal centroid displacement and all mobility-related CSG properties, while the relationship weakened toward the diaphyseal ends. No significant relationship was found between centroid displacement and body size variables with the exception of the most distal section location. Results support a relationship between tibial curvature and cross-sectional geometry among prehistoric Central European agricultural populations, and suggest that changes in mechanical loading may have influenced a suite of morphological features related to bone adaptation in the lower limb.

Periosteal versus true cross-sectional geometry: a comparison along humeral, femoral, and tibial diaphyses.

Cross-sectional geometric (CSG) properties of human long bone diaphyses are typically calculated from both periosteal and endosteal contours. Though quantification of both is desirable, periosteal contours alone have provided accurate predictions of CSG properties at the midshaft in previous studies. The relationship between CSG properties calculated from external contours and "true" (endosteal and periosteal) CSG properties, however, has yet to be examined along the whole diaphysis. Cross-sectional computed tomography scans were taken from 21 locations along humeral, femoral, and tibial diaphyses in 20 adults from a late prehistoric central Illinois Valley cemetery. Mechanical properties calculated from images with (a) artificially filled medullary cavities ("solid") and (b) true unaltered cross-sections were compared at each section location using least squares regression. Results indicate that, in this sample, polar second moments of area (J), polar section moduli (Z(p) ), and cross-sectional shape (I(max) /I(min) ) calculated from periosteal contours correspond strongly with those calculated from cross-sections that include the medullary cavity. Correlations are high throughout most of the humeral diaphysis and throughout large portions of femoral and tibial diaphyses (R(2) = 0.855-0.998, all P < 0.001, %SEE ≤ 8.0, %PE ≤ 5.0), the major exception being the proximal quarter of the tibial diaphysis for J and Z(p). The main source of error was identified as variation in %CA. Results reveal that CSG properties quantified from periosteal contours provide comparable results to (and are likely to detect the same differences among individuals as) true CSG properties along large portions of long bone diaphyses.

Thrifty phenotype versus cold adaptation: trade-offs in upper limb proportions of Himalayan populations of Nepal.

The multi-stress environment of high altitude has been associated with growth deficits in humans, particularly in zeugopod elements (forearm and lower leg). This is consistent with the thrifty phenotype hypothesis, which has been observed in Andeans, but has yet to be tested in other high-altitude populations. In Himalayan populations, other factors, such as cold stress, may shape limb proportions. The current study investigated whether relative upper limb proportions of Himalayan adults (n = 254) differ between highland and lowland populations, and whether cold adaptation or a thrifty phenotype mechanism may be acting here. Height, weight, humerus length, ulna length, hand length and hand width were measured using standard methods. Relative to height, total upper limb and ulna lengths were significantly shorter in highlanders compared with lowlanders in both sexes, while hand and humerus length were not. Hand width did not significantly differ between populations. These results support the thrifty phenotype hypothesis, as hand and humerus proportions are conserved at the expense of the ulna. The reduction in relative ulna length could be attributed to cold adaptation, but the lack of difference between populations in both hand length and width indicates that cold adaptation is not shaping hand proportions in this case.

Maternal investment, maturational rate of the offspring and mechanical competence of the adult female skeleton.

LAY SUMMARY: Girls with a slower life history trajectory build a larger body with larger and mechanically stronger bones. Thus, variation in the emergence of slower versus faster life history trajectories during development can have consequences for bone mechanical competence, and hence fracture risk in adulthood. BACKGROUND AND OBJECTIVES: Variation in life history trajectory, specifically relative investment in growth versus reproduction, has been associated with chronic disease risk among women, but whether this scenario extends to skeletal health and fracture risk is unknown. This study investigates the association of life history traits (proxies for maternal investment and maturational rate) with female bone outcomes in adulthood. METHODOLOGY: Body size variables, regional muscle and fat areas, and cross-sectional bone size and strength outcomes were obtained from 107 pre-menopausal women encompassing a wide range of physical activity levels. Developmental parameters (birth weight, age at menarche) were obtained from questionnaires. RESULTS: High birth weight was significantly associated with a proportionately larger body and larger, mechanically stronger bones, independently of physical activity level. It was also positively but non-significantly associated with age at menarche. Later menarche was significantly associated with larger and mechanically stronger bones and substantially less absolute and relative regional subcutaneous fat. Age at menarche exhibited stronger relationships with adult adiposity than did physical activity. CONCLUSIONS AND IMPLICATIONS: Both larger birth weight and later menarche contribute to a slower life history trajectory, which is associated with greater body size, leanness and bone mechanical competence in early adulthood. In contrast, earlier sexual maturity prioritized energy allocation in adiposity over body size and skeletal strength. Thus, the level of maternal investment and the woman's own life history trajectory shape investment in skeletal properties, with implications for fracture risk later in life.

Prehistoric women's manual labor exceeded that of athletes through the first 5500 years of farming in Central Europe.

The intensification of agriculture is often associated with declining mobility and bone strength through time, although women often exhibit less pronounced trends than men. For example, previous studies of prehistoric Central European agriculturalists (~5300 calibrated years BC to 850 AD) demonstrated a significant reduction in tibial rigidity among men, whereas women were characterized by low tibial rigidity, little temporal change, and high variability. Because of the potential for sex-specific skeletal responses to mechanical loading and a lack of modern comparative data, women's activity in prehistory remains difficult to interpret. This study compares humeral and tibial cross-sectional rigidity, shape, and interlimb loading among prehistoric Central European women agriculturalists and living European women of known behavior (athletes and controls). Prehistoric female tibial rigidity at all time periods was highly variable, but differed little from living sedentary women on average, and was significantly lower than that of living runners and football players. However, humeral rigidity exceeded that of living athletes for the first ~5500 years of farming, with loading intensity biased heavily toward the upper limb. Interlimb strength proportions among Neolithic, Bronze Age, and Iron Age women were most similar to those of living semi-elite rowers. These results suggest that, in contrast to men, rigorous manual labor was a more important component of prehistoric women's behavior than was terrestrial mobility through thousands of years of European agriculture, at levels far exceeding those of modern women.

Early Life Conditions and Physiological Stress following the Transition to Farming in Central/Southeast Europe: Skeletal Growth Impairment and 6000 Years of Gradual Recovery.

Early life conditions play an important role in determining adult body size. In particular, childhood malnutrition and disease can elicit growth delays and affect adult body size if severe or prolonged enough. In the earliest stages of farming, skeletal growth impairment and small adult body size are often documented relative to hunter-gatherer groups, though this pattern is regionally variable. In Central/Southeast Europe, it is unclear how early life stress, growth history, and adult body size were impacted by the introduction of agriculture and ensuing long-term demographic, social, and behavioral change. The current study assesses this impact through the reconstruction and analysis of mean stature, body mass, limb proportion indices, and sexual dimorphism among 407 skeletally mature men and women from foraging and farming populations spanning the Late Mesolithic through Early Medieval periods in Central/Southeast Europe (~7100 calBC to 850 AD). Results document significantly reduced mean stature, body mass, and crural index in Neolithic agriculturalists relative both to Late Mesolithic hunter-gatherer-fishers and to later farming populations. This indication of relative growth impairment in the Neolithic, particularly among women, is supported by existing evidence of high developmental stress, intensive physical activity, and variable access to animal protein in these early agricultural populations. Among subsequent agriculturalists, temporal increases in mean stature, body mass, and crural index were more pronounced among Central European women, driving declines in the magnitude of sexual dimorphism through time. Overall, results suggest that the transition to agriculture in Central/Southeast Europe was challenging for early farming populations, but was followed by gradual amelioration across thousands of years, particularly among Central European women. This sex difference may be indicative, in part, of greater temporal variation in the social status afforded to young girls, in their access to resources during growth, and/or in their health status than was experienced by men.

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.