Long-term trends in human body size track regional variation in subsistence transitions and growth acceleration linked to dairying.

Evidence for a reduction in stature between Mesolithic foragers and Neolithic farmers has been interpreted as reflective of declines in health, however, our current understanding of this trend fails to account for the complexity of cultural and dietary transitions or the possible causes of phenotypic change. The agricultural transition was extended in primary centers of domestication and abrupt in regions characterized by demic diffusion. In regions such as Northern Europe where foreign domesticates were difficult to establish, there is strong evidence for natural selection for lactase persistence in relation to dairying. We employ broad-scale analyses of diachronic variation in stature and body mass in the Levant, Europe, the Nile Valley, South Asia, and China, to test three hypotheses about the timing of subsistence shifts and human body size, that: 1) the adoption of agriculture led to a decrease in stature, 2) there were different trajectories in regions of in situ domestication or cultural diffusion of agriculture; and 3) increases in stature and body mass are observed in regions with evidence for selection for lactase persistence. Our results demonstrate that 1) decreases in stature preceded the origins of agriculture in some regions; 2) the Levant and China, regions of in situ domestication of species and an extended period of mixed foraging and agricultural subsistence, had stable stature and body mass over time; and 3) stature and body mass increases in Central and Northern Europe coincide with the timing of selective sweeps for lactase persistence, providing support for the "Lactase Growth Hypothesis."

Trabecular bone ontogeny tracks neural development and life history among humans and non-human primates.

Trabecular bone-the spongy bone inside marrow cavities-adapts to its mechanical environment during growth and development. Trabecular structure can therefore be interpreted as a functional record of locomotor behavior in extinct vertebrates. In this paper, we expand upon traditional links between form and function by situating ontogenetic trajectories of trabecular bone in four primate species into the broader developmental context of neural development, locomotor control, and ultimately life history. Our aim is to show that trabecular bone structure provides insights into ontogenetic variation in locomotor loading conditions as the product of interactions between increases in body mass and neuromuscular maturation. Our results demonstrate that age-related changes in trabecular bone volume fraction (BV/TV) are strongly and linearly associated with ontogenetic changes in locomotor kinetics. Age-related variation in locomotor kinetics and BV/TV is in turn strongly associated with brain and body size growth in all species. These results imply that age-related variation in BV/TV is a strong proxy for both locomotor kinetics and neuromuscular maturation. Finally, we show that distinct changes in the slope of age-related variation in bone volume fraction correspond to the age of the onset of locomotion and the age of locomotor maturity. Our findings compliment previous studies linking bone development to locomotor mechanics by providing a fundamental link to brain development and life history. This implies that trabecular structure of fossil subadults can be a proxy for the rate of neuromuscular maturation and major life history events like locomotor onset and the achievement of adult-like locomotor repertoires.

The evolution and changing ecology of the African hominid oral microbiome.

The oral microbiome plays key roles in human biology, health, and disease, but little is known about the global diversity, variation, or evolution of this microbial community. To better understand the evolution and changing ecology of the human oral microbiome, we analyzed 124 dental biofilm metagenomes from humans, including Neanderthals and Late Pleistocene to present-day modern humans, chimpanzees, and gorillas, as well as New World howler monkeys for comparison. We find that a core microbiome of primarily biofilm structural taxa has been maintained throughout African hominid evolution, and these microbial groups are also shared with howler monkeys, suggesting that they have been important oral members since before the catarrhine-platyrrhine split ca. 40 Mya. However, community structure and individual microbial phylogenies do not closely reflect host relationships, and the dental biofilms of Homo and chimpanzees are distinguished by major taxonomic and functional differences. Reconstructing oral metagenomes from up to 100 thousand years ago, we show that the microbial profiles of both Neanderthals and modern humans are highly similar, sharing functional adaptations in nutrient metabolism. These include an apparent Homo-specific acquisition of salivary amylase-binding capability by oral streptococci, suggesting microbial coadaptation with host diet. We additionally find evidence of shared genetic diversity in the oral bacteria of Neanderthal and Upper Paleolithic modern humans that is not observed in later modern human populations. Differences in the oral microbiomes of African hominids provide insights into human evolution, the ancestral state of the human microbiome, and a temporal framework for understanding microbial health and disease.

Patterns of energy allocation during energetic scarcity; evolutionary insights from ultra-endurance events.

Exercise physiologists and evolutionary biologists share a research interest in determining patterns of energy allocation during times of acute or chronic energetic scarcity. Within sport and exercise science, this information has important implications for athlete health and performance. For evolutionary biologists, this would shed new light on our adaptive capabilities as a phenotypically plastic species. In recent years, evolutionary biologists have begun recruiting athletes as study participants and using contemporary sports as a model for studying evolution. This approach, known as human athletic palaeobiology, has identified ultra-endurance events as a valuable experimental model to investigate patterns of energy allocation during conditions of elevated energy demand, which are generally accompanied by an energy deficit. This energetic stress provokes detectable functional trade-offs in energy allocation between physiological processes. Early results from this modelsuggest thatlimited resources are preferentially allocated to processes which could be considered to confer the greatest immediate survival advantage (including immune and cognitive function). This aligns with evolutionary perspectives regarding energetic trade-offs during periods of acute and chronic energetic scarcity. Here, we discuss energy allocation patterns during periods of energetic stress as an area of shared interest between exercise physiology and evolutionary biology. We propose that, by addressing the ultimate "why" questions, namely why certain traits were selected for during the human evolutionary journey, an evolutionary perspective can complement the exercise physiology literature and provide a deeper insight of the reasons underpinning the body's physiological response to conditions of energetic stress.

Growth and development of trabecular structure in the calcaneus of Japanese macaques (Macaca fuscata) reflects locomotor behavior, life history, and neuromuscular development.

Bone structure dynamically adapts to its mechanical environment throughout ontogeny by altering the structure of trabecular bone, the three-dimensional mesh-like structure found underneath joint surfaces. Trabecular structure, then, can provide a record of variation in loading directions and magnitude; and in ontogenetic samples, it can potentially be used to track developmental shifts in limb posture. We aim to broaden the analysis of trabecular bone ontogeny by incorporating interactions between ontogenetic variation in locomotor repertoire, neuromuscular maturation, and life history. We examine the associations between these variables and age-related variation in trabecular structure in the calcaneus of Japanese macaques (Macaca fuscata). We used high-resolution micro-computed tomography scanning to image the calcaneus in a cross-sectional sample of 34 juvenile M. fuscata aged between 0 and 7 years old at the Primate Research Institute, Japan. We calculated whole bone averages of standard trabecular properties and generated whole-bone morphometric maps of bone volume fraction and Young's modulus. Trabecular structure becomes increasingly heterogeneous in older individuals. Bone volume fraction (BV/total volume [TV]) decreases during the first month of life and increases afterward, coinciding with the onset of independent locomotion in M. fuscata. At birth, primary Young's modulus is oriented orthogonal to the ossification center, but after locomotor onset bone structure becomes stiffest in the direction of joint surfaces and muscle attachments. Age-related variation in bone volume fraction is best predicted by an interaction between the estimated percentage of adult brain size, body mass, and locomotor onset. To explain our findings, we propose a model where interactions between age-related increases in body weight and maturation of the neuromuscular system alter the loading environment of the calcaneus, to which the internal trabecular structure dynamically adapts. This model cannot be directly tested based on our cross-sectional data. However, confirmation of the model by longitudinal experiments and in multiple species would show that trabecular structure can be used both to infer behavior from fossil morphology and serve as a valuable proxy for neuromuscular maturation and life history events like locomotor onset and the achievement of an adult-like gait. This approach could significantly expand our knowledge of the biology and behavior of fossil species.

Evaluation of dual-energy X-ray absorptiometry compared to magnetic resonance imaging for collecting measurements of the human bony pelvis.

OBJECTIVES: Imaging methods to measure the human pelvis in vivo provide opportunities to better understand pelvic variation and adaptation. Magnetic resonance imaging (MRI) provides high-resolution images, but is more expensive than dual-energy X-ray absorptiometry (DXA). We sought to compare pelvic breadth measurements collected from the same individuals using both methods, to investigate if there are systematic differences in pelvic measurement between these imaging methods. METHODS: Three pelvic breadth dimensions (bi-iliac breadth, bi-acetabular breadth, medio-lateral inlet breadth) were collected from MRI and DXA scans of a cross-sectional sample of healthy, nulliparous adult women of South Asian ancestry (n = 63). Measurements of MRI and DXA pelvic dimensions were collected four times in total, with one baseline data collection session and three replications. Data collected from these sessions were averaged, used to calculate technical error of measurement and entered into a Bland-Altman analysis. Linear regression models were fitted with a given MRI pelvic measurement regressed on the same measurement collected from DXA scans, as well as MRI mean bias regressed on DXA mean bias. RESULTS: Technical error of measurement was higher in DXA measurements of bi-iliac breadth and medio-lateral pelvic inlet breadth and higher for MRI measurements of bi-acetabular breadth. Bland Altman analyses showed no statistically significant relationship between the mean bias of MRI and DXA, and the differences between MRI and DXA pelvic measurements. CONCLUSIONS: DXA measurements of pelvic breadth are comparable to MRI measurements of pelvic breadth. DXA is a less costly imaging technique than MRI and can be used to collect measurements of skeletal elements in living people.

Unique foot posture in Neanderthals reflects their body mass and high mechanical stress.

Neanderthal foot bone proportions and morphology are mostly indistinguishable from those of Homo sapiens, with the exception of several distinct Neanderthal features in the talus. The biomechanical implications of these distinct talar features remain contentious, fueling debate around the adaptive meaning of this distinctiveness. With the aim of clarifying this controversy, we test phylogenetic and behavioral factors as possible contributors, comparing tali of 10 Neanderthals and 81 H. sapiens (Upper Paleolithic and Holocene hunter-gatherers, agriculturalists, and postindustrial group) along with the Clark Howell talus (Omo, Ethiopia). Variation in external talar structures was assessed through geometric morphometric methods, while bone volume fraction and degree of anisotropy were quantified in a subsample (n = 45). Finally, covariation between point clouds of site-specific trabecular variables and surface landmark coordinates was assessed. Our results show that although Neanderthal talar external and internal morphologies were distinct from those of H. sapiens groups, shape did not significantly covary with either bone volume fraction or degree of anisotropy, suggesting limited covariation between external and internal talar structures. Neanderthal external talar morphology reflects ancestral retentions, along with various adaptations to high levels of mobility correlated to their presumably unshod hunter-gatherer lifestyle. This pairs with their high site-specific trabecular bone volume fraction and anisotropy, suggesting intense and consistently oriented locomotor loading, respectively. Relative to H.sapiens, Neanderthals exhibit differences in the talocrural joint that are potentially attributable to cultural and locomotor behavior dissimilarity, a talonavicular joint that mixes ancestral and functional traits, and a derived subtalar joint that suggests a predisposition for a pronated foot during stance phase. Overall, Neanderthal talar variation is attributable to mobility strategy and phylogenesis, while H. sapiens talar variation results from the same factors plus footwear. Our results suggest that greater Neanderthal body mass and/or higher mechanical stress uniquely led to their habitually pronated foot posture.

Fluctuating asymmetry, a marker of poor growth quality, is associated with adult male metabolic rate.

OBJECTIVES: Life history theory, a branch of evolutionary theory, predicts the existence of trade-offs in energetic allocation between competing physiological functions. The core metabolic cost of self-maintenance, measured by resting metabolic rate (RMR), represents a large component of human daily energy expenditure. Despite strong selective pressures for energetic frugality and high observed interindividual variation in RMR, the link between RMR and energetic allocation to life-history traits remains understudied in humans. MATERIALS: In a sample of 105 (m = 57, f = 48), we investigated the relationship between adult RMR and investment in growth quality, as measured by fluctuating asymmetry (FA). RESULTS: Measurement of RMR and FA in university rowers revealed a significant positive correlation amongst males (n = 57, r = 0.344, p = 0.005, 1-tailed; standardized 95% CI, 0.090 to 0.598). Convincing evidence for a correlation among females was not found (n = 48, r = 0.142, p = 0.169, 1-tailed, standardized 95% CI, -0.152 to 0.435). DISCUSSION: The data suggest that low-quality asymmetrical growth is associated with later-life metabolic inefficiencies in males. Energetic investment in processes (likely concerning the stress-response) unrelated to growth during childhood may thereby trade-off against adult metabolic efficiency. We suggest that the presence of a relationship between RMR and FA in males but not females may be explained by the additional metabolic strain associated with larger body size and increased male muscularity, which may amplify the inefficiencies arising from low-quality growth.

Combinations of trabecular and cortical bone properties distinguish various loading modalities between athletes and controls.

OBJECTIVES: Variation in trabecular and cortical bone properties is often used to infer habitual behavior in the past. However, the structures of both types of bone are rarely considered together and may even contradict each other in functional interpretations. We examine trabecular and cortical bone properties in various athletes and sedentary controls to clarify the associations between combinations of cortical and trabecular bone properties and various loading modalities. MATERIALS AND METHODS: We compare trabecular and cortical bone properties using peripheral quantitative computed tomography scans of the tibia between groups of 83 male athletes (running, hockey, swimming, cricket) and sedentary controls using Bayesian multilevel models. We quantify midshaft cortical bone rigidity and area (J, CA), midshaft shape index (Imax/Imin), and mean trabecular bone mineral density (BMD) in the distal tibia. RESULTS: All groups show unique combinations of biomechanical properties. Cortical bone rigidity is high in sports that involve impact loading (cricket, running, hockey) and low in nonimpact loaded swimmers and controls. Runners have more anteroposteriorly elliptical midshafts compared to other groups. Interestingly, all athletes have greater trabecular BMD compared to controls, but do not differ credibly among each other. DISCUSSION: Results suggest that cortical midshaft hypertrophy is associated with impact loading while trabecular BMD is positively associated with both impact and nonimpact loading. Midshaft shape is associated with directionality of loading. Individuals from the different categories overlap substantially, but group means differ credibly, suggesting that nuanced group-level inferences of habitual behavior are possible when combinations of trabecular and cortical bone are analyzed.

Automated resolution independent method for comparing in vivo and dry trabecular bone.

OBJECTIVES: Variation in human trabecular bone morphology can be linked to habitual behavior, but it is difficult to investigate in vivo due to the radiation required at high resolution. Consequently, functional interpretations of trabecular morphology remain inferential. Here we introduce a method to link low- and high-resolution CT data from dry and fresh bone, enabling bone functional adaptation to be studied in vivo and results compared to the fossil and archaeological record. MATERIALS AND METHODS: We examine 51 human dry bone distal tibiae from Nile Valley and UK and two pig tibiae containing soft tissues. We compare low-resolution peripheral quantitative computed tomography (pQCT) parameters and high-resolution micro CT (µCT) in homologous single slices at 4% bone length and compare results to our novel Bone Ratio Predictor (BRP) method. RESULTS: Regression slopes between linear attenuation coefficients of low-resolution pQCT images and bone area/total area (BA/TA) of high-resolution µCT scans differ substantially between geographical subsamples, presumably due to diagenesis. BRP accurately predicts BA/TA (R2 = .97) and eliminates the geographic clustering. BRP accurately estimates BA/TA in pigs containing soft tissues (R2 = 0.98) without requiring knowledge of true density or phantom calibration of the scans. DISCUSSION: BRP allows automated comparison of image data from different image modalities (pQCT, µCT) using different energy settings, in archeological bone and wet specimens. The method enables low-resolution data generated in vivo to be compared with the fossil and archaeological record. Such experimental approaches would substantially improve behavioral inferences based on trabecular bone microstructure.

Obstetric dimensions of the female pelvis are less integrated than locomotor dimensions and show protective scaling patterns: Implications for the obstetrical dilemma.

OBJECTIVES: The "obstetrical dilemma" hypothesis assumes that the modern human female pelvis serves two discrete functions: obstetrics and locomotion. We investigate whether these differing functions create observable patterns of morphological covariation and whether those patterns differ by height, weight, and age. This allows evaluation of evidence for canalization and phenotypic plasticity relevant to obstetric and locomotor function among a living female population. METHODS: Landmarks (N = 86) were collected and inter-landmark distances were calculated (N = 36) on the pelvis and proximal femur of CT scans of living women aged 20 to 90 years (M = 93) receiving a routine CT scan. Partial least squares and relative SD of eigenvalues analyses were used to evaluate integration overall and within locomotor and obstetric modules, respectively. Ordinary Least Squared regression was used to evaluate scaling relationships between inter-landmark distances and height, weight, and age. RESULTS: The obstetric pelvis was significantly less internally integrated than the locomotor pelvis. Many obstetric measurements were constrained in absolute terms relative to height; shorter women had relatively larger birth canal dimensions, and several key obstetric dimensions showed relative freedom from height. Lower weight women had some relatively larger obstetric and locomotor dimensions. Regarding age, younger women showed a few relatively larger outlet dimensions. CONCLUSIONS: This study suggests that the obstetric pelvis and the locomotor pelvis function are morphologically distinct, with the obstetric pelvis showing relatively greater flexibility. These relationships between relative constraints support the hypothesis that the modern female pelvis shows evidence of both canalization and phenotypic plasticity in obstetric and locomotor structures.

Using point clouds to investigate the relationship between trabecular bone phenotype and behavior: An example utilizing the human calcaneus.

OBJECTIVES: The objective of this study is to demonstrate a new method for analyzing trabecular bone volume fraction and degree of anisotropy in three dimensions. METHODS: We use a combination of automatic mesh registration, point-cloud correspondence registration, and P-value corrected univariate statistical tests to compare bone volume fraction and degree of anisotropy on a point by point basis across the entire calcaneus of two human groups with different subsistence strategies. RESULTS: We found that the patterns of high and low bone volume fraction and degree of anisotropy distribution between the Black Earth (hunter-gatherers) and Norris Farms (mixed-strategy agriculturalists) are very similar, but differ in magnitude. The hunter-gatherers exhibit higher levels of bone volume fraction and less anisotropic trabecular bone organization. Additionally, patterns of bone volume fraction and degree of anisotropy in the calcaneus correspond well with biomechanical expectations of relative forces experienced during walking and running. CONCLUSIONS: We conclude that comparing site-specific, localized differences in trabecular bone variables such as bone volume fraction and degree of anisotropy in three-dimensions is a powerful analytical tool. This method makes it possible to determine where similarities and differences between groups are located within the whole skeletal element of interest. The visualization of multiple variables also provides a way for researchers to see how the trabecular bone variables interact within the morphology, and allows for a more nuanced understanding of how they relate to one another and the broader mechanical environment.

Baby steps towards linking calcaneal trabecular bone ontogeny and the development of bipedal human gait.

Trabecular bone structure in adulthood is a product of a process of modelling during ontogeny and remodelling throughout life. Insight into ontogeny is essential to understand the functional significance of trabecular bone structural variation observed in adults. The complex shape and loading of the human calcaneus provides a natural experiment to test the relationship between trabecular morphology and locomotor development. We investigated the relationship between calcaneal trabecular bone structure and predicted changes in loading related to development of gait and body size in growing children. We sampled three main trabecular regions of the calcanei using micro-computed tomography scans of 35 individuals aged between neonate to adult from the Norris Farms #36 site (1300 AD, USA) and from Cambridge (1200-1500 AD, UK). Trabecular properties were calculated in volumes of interest placed beneath the calcaneocuboid joint, plantar ligaments, and posterior talar facet. At birth, thin trabecular struts are arranged in a dense and relatively isotropic structure. Bone volume fraction strongly decreases in the first year of life, whereas anisotropy and mean trabecular thickness increase. Dorsal compressive trabecular bands appear around the onset of bipedal walking, although plantar tensile bands develop prior to predicted propulsive toe-off. Bone volume fraction and anisotropy increase until the age of 8, when gait has largely matured. Connectivity density gradually reduces, whereas trabeculae gradually thicken from birth until adulthood. This study demonstrates that three different regions of the calcaneus develop into distinct adult morphologies through varying developmental trajectories. These results are similar to previous reports of ontogeny in human long bones and are suggestive of a relationship between the mechanical environment and trabecular bone architecture in the human calcaneus during growth. However, controlled experiments combined with more detailed biomechanical models of gait maturation are necessary to establish skeletal markers linking growth to loading. This has the potential to be a novel source of information for understanding loading levels, activity patterns, and perhaps life history in the fossil record.

Muscle force interacts with stature to influence functionally related polar second moments of area in the lower limb among adult women.

OBJECTIVES: We sought to determine the relationships between muscle size, function, and polar second moments of area (J) at the midshaft femur, proximal tibia, and midshaft tibia. MATERIALS AND METHODS: We used peripheral quantitative computed tomography to quantify right femoral and tibial J and soft tissue cross-sectional areas, and force plate mechanography to quantify peak power output and maximum force of the right limb, among athletic women and control subjects. RESULTS: Lower limb bone J exhibited strong relationships with estimated force but not power between both groups. Among controls, the strongest relationships between force and J were found at the midshaft femur. Among athletes, these relationships shifted to the tibia, regardless of body size, likely reflecting functional strain related to the major knee extensors and ankle plantarflexors. Together, muscle force and stature explained as much as 82 and 48% of the variance in lower limb bone J among controls and athletes, respectively. DISCUSSION: Results highlight the importance of considering relevant muscle function variables (e.g., force and lever arm lengths) when interpreting behavioral signatures from skeletal remains. Future work to improve the estimation of muscle force from skeletal remains, and incorporate it with lever arm length into analyses, is warranted. Results also suggest that, in doing so, functional relationships between a given section location and musculature should be considered.

Human athletic paleobiology; using sport as a model to investigate human evolutionary adaptation.

The use of sport as a conceptual framework offers unprecedented opportunities to improve our understanding of what the body does, shedding new light on our evolutionary trajectory, our capacity for adaptation, and the underlying biological mechanisms. This approach has gained traction over recent years. To date, sport has facilitated exploration not only of the evolutionary history of our species as a whole, but also of human variation and adaptation at the interindividual and intraindividual levels. At the species level, analysis of lower and upper limb biomechanics and energetics with respect to walking, running and throwing have led to significant advances in the understanding of human adaptations relative to other hominins. From an interindividual perspective, investigation of physical activity patterns and endurance running performance is affording greater understanding of evolved constraints of energy expenditure, thermoregulatory energetics, signaling theory, and morphological variation. Furthermore, ultra-endurance challenges provoke functional trade-offs, allowing new ground to be broken in the study of life history trade-offs and human adaptability. Human athletic paleobiology-the recruitment of athletes as study participants and the use of contemporary sports as a model for studying evolutionary theory-has great potential. Here, we draw from examples in the literature to provide a review of how the use of athletes as a model system is enhancing understanding of human evolutionary adaptation.

Intrapopulation variation in lower limb trabecular architecture.

OBJECTIVES: Trabecular structure is frequently used to differentiate between highly divergent mechanical environments. Less is known regarding the response of the structural properties to more subtle behavioral differences, as the range of intrapopulation variation in trabecular architecture is rarely studied. Examining the extent to which lower limb trabecular architecture varies when inferred mobility levels and environment are consistent between groups within a relatively homogenous population may aid in the contextualization of interpopulation differences, improve detectability of sexual dimorphism in trabecular structure, and improve our understanding of trabecular bone functional adaptation. MATERIALS AND METHODS: The study sample was composed of adult individuals from three high/late medieval cemeteries from Cambridge (10th-16th c.), a hospital (n = 57), a parish cemetery (n = 44) and a friary (n = 14). Trabecular architecture was quantified in the epiphyses of the femur and tibia, using high resolution computed tomography. RESULTS: The parish individuals had the lowest bone volume fraction and trabecular thickness in most regions. Multiple sex differences were observed, but the patterns were not consistent across volumes of interest. DISCUSSION: Differences between the three groups highlight the great variability of trabecular bone architecture, even within a single sedentary population. This indicates that trabecular bone may be used in interpreting subtle behavioral differences, and suggests that multiple archaeological sites need to be studied to characterize structural variation on a population level. Variation in sex and group differences across anatomical locations further demonstrates the site-specificity in trabecular bone functional adaptation, which might explain why little consistent sexual dimorphism has been reported previously.

Trabecular bone structure scales allometrically in the foot of four human groups.

The human foot is highly derived relative to that of other hominoids and therefore a topic of intense research in paleoanthropology. While trabecular bone is thought to be highly plastic in response to habitual behavior, knowledge of how trabecular structure scale with body size is essential for making functional inferences from trabecular bone morphology. Trabecular bone properties scale with negative allometry in interspecific studies that includes a wide range of body size; however, intraspecific scaling patterns often differ from interspecific trends. In this paper we examine patterns of trabecular bone scaling in the calcaneus, talus, and first metatarsal of four human populations with different subsistence strategies and associated levels of terrestrial mobility. We report Bayesian linear regressions between the natural logarithms of femoral head diameter and five standard trabecular variables calculated in five spherical volumes of interest. We additionally report regressions on population-specific z-scores of femoral head diameter and trabecular variables as a way of placing the four populations on a common scale. Results show that with increasing body size there is no change in bone volume fraction (BV/TV) and trabecular thickness (Tb.Th), a slight increase in trabecular spacing (Tb.Sp), and a sharp decrease in connectivity density (Conn.D). Degree of anisotropy was found to scale with positive allometry in the calcaneus, negative allometry in the talar trochlea, and shows no relationship with femoral head diameter in the talar and first metatarsal heads. These results show that scaling of the degree of anisotropy can vary substantially within and between bones. Degree of anisotropy is often used as a proxy for directionality in joint loading when interpreting variation in trabecular structures of fossils and extant primates. Body size should therefore be an important consideration when trabecular bone structure is used to interpret function from fossil morphology.

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.

Trabecular bone functional adaptation and sexual dimorphism in the human foot.

OBJECTIVES: Trabecular bone adapts to the strains placed upon the skeleton during life. Anthropological research has largely focused on linking variation in primate trabecular bone to locomotor mode, to provide a context for interpreting fossil morphology. However, intraspecific variation and its underlying mechanisms are still poorly understood. Trabecular bone is influenced by a variety of factors including body mass, age, diet, temperature, genetics, sex, and behavior. Before trabecular structure can be used to infer habitual behavior in the past, the effects of these factors need to be understood. In this article, we examine variation in trabecular structure in the human foot in four archaeological groups in relation to inferred levels of terrestrial mobility and sex. MATERIALS AND METHODS: We use high-resolution µCT scanning to examine variation in trabecular structure in the human calcaneus, talus, and first metatarsal in two relatively mobile and two relatively sedentary archaeological groups. RESULTS: The four population samples show similar patterns of trabecular variation throughout the foot, influenced by mechanical loading. Greater inferred terrestrial mobility is associated with greater bone volume fraction and thicker, more widely spaced, and less interconnected trabeculae. However, contrary to diaphyseal rigidity, only limited sexual dimorphism was found in trabecular structure. DISCUSSION: This work demonstrates that trabecular bone may serve as a useful proxy of habitual behavior in the fossil and archaeological record when other factors are carefully considered. However, the mechanisms underlying sexual dimorphism are not well understood. As such, inferring sex differences in habitual behavior is currently challenging.

Complex variation of trabecular bone structure in the proximal humerus and femur of five modern human populations.

OBJECTIVE: This project investigates trabecular bone structural variation in the proximal humerus and femur of hunter-gatherer, mixed-strategy agricultural, medieval, and human groups to address three questions: (a) What is the extent of trabecular bone structural variation in the humerus and femur between populations with different inferred activity levels? (b) How does variation in the proximal humerus relate to variation in the proximal femur? (c) Are trabecular bone microstructural variables sexually dimorphic? METHODS: The proximal humerus and femur of 73 adults from five human groups with distinct subsistence strategies were scanned using a micro-computed tomography system. Centralized volumes of interest within the humeral and femoral heads were extracted and analyzed to quantify bone volume fraction, trabecular thickness, trabecular separation, connectivity density, degree of anisotropy, and bone surface density. RESULTS: In the humerus and femur, groups with the highest inferred activity levels have higher bone volume fraction and trabecular thickness, and lower bone surface density than those with lower inferred activity levels. However, the humeral pattern does not exactly mirror that of the femur, which demonstrates a steeper gradient of difference between subsistence groups. No significant differences were identified in trabecular separation. No consistent patterns of sexual dimorphism were present in the humerus or femur. CONCLUSIONS: Reduced skeletal robusticity of proximal humeral and femoral trabecular bone corresponds with reduced activity level inferred from subsistence strategy. However, human trabecular bone structural variation is complex and future work should explore how other factors (diet, climate, genetics, disease load, etc.), in addition to activity, influence bone structural variation.