The postcranial skeleton of modern Homo sapiens is relatively gracile compared with other hominoids and earlier hominins. This gracility predisposes contemporary humans to osteoporosis and increased fracture risk. Explanations for this gracility include reduced levels of physical activity, the dissipation of load through enlarged joint surfaces, and selection for systemic physiological characteristics that differentiate modern humans from other primates. This study considered the skeletal remains of four behaviorally diverse recent human populations and a large sample of extant primates to assess variation in trabecular bone structure in the human hip joint. Proximal femur trabecular bone structure was quantified from microCT data for 229 individuals from 31 extant primate taxa and 59 individuals from four distinct archaeological human populations representing sedentary agriculturalists and mobile foragers. Analyses of mass-corrected trabecular bone variables reveal that the forager populations had significantly higher bone volume fraction, thicker trabeculae, and consequently lower relative bone surface area compared with the two agriculturalist groups. There were no significant differences between the agriculturalist and forager populations for trabecular spacing, number, or degree of anisotropy. These results reveal a correspondence between human behavior and bone structure in the proximal femur, indicating that more highly mobile human populations have trabecular bone structure similar to what would be expected for wild nonhuman primates of the same body mass. These results strongly emphasize the importance of physical activity and exercise for bone health and the attenuation of age-related bone loss. trabecular bone | gracilization | human evolution | biomechanics | mobility C ompared with other hominoids and extinct hominin species, more recent humans possess relatively gracile postcranial skeletons (1-9). One of the consequences of this gracility in contemporary humans is an increased fracture risk associated with age-related bone loss and osteoporosis [hip fractures alone are projected to reach 6.26 million per year globally by 2050 (10)] (11-15). The etiology of this relative gracility remains uncertain, and this uncertainty hinders the development of strategies for mitigating fracture risk and morbidity. The progressive gracilization of the Homo postcranial skeleton was originally detected in cortical bone structure (1, 2), but has now been demonstrated in the trabecular bone microstructure of joints (12,14,(16)(17)(18)(19), where osteoporotic fracture risk is highest (20). Most notably, in an analysis of thoracic vertebral bodies, Cotter et al. (12) found that young adult humans have significantly lower trabecular bone volume fraction (BV/TV) and thinner vertebral shells than similarly sized apes. Griffin et al. (16) also found significantly lower BV/TV in the human first and second metatarsal heads compared with hominoid primates. The results of these studies are corroborated by work on the hominoid...

Variation in upper limb long bone cross-sectional properties may reflect a phenotypically plastic response to habitual loading patterns. Structural differences between limb bones have often been used to infer past behavior from hominin remains; however, few studies have examined direct relationships between behavioral differences and bone structure in humans. To help address this, cross-sectional images (50% length) of the humeri and ulnae of university varsity-level swimmers, cricketers, and controls were captured using peripheral quantitative computed tomography. High levels of humeral robusticity were found in the dominant arms of cricketers, and bilaterally among swimmers, whereas the most gracile humeri were found in both arms of controls, and the nondominant arms of cricketers. In addition, the dominant humeri of cricketers were more circular than controls. The highest levels of ulnar robusticity were also found in the dominant arm of cricketers, and bilaterally amongst swimmers. Bilateral asymmetry in humeral rigidity among cricketers was greater than swimmers and controls, while asymmetry for ulnar rigidity was greater in cricketers than controls. The results suggest that more mechanically loaded upper limb elements--unilaterally or bilaterally--are strengthened relative to less mechanically loaded elements, and that differences in mechanical loading may have a more significant effect on proximal compared to distal limb segments. The more circular humerus in the dominant arm in cricketers may be an adaptation to torsional strain associated with throwing activities. The reported correspondence between habitual activity patterns and upper limb diaphyseal properties may inform future behavioral interpretations involving hominin skeletal remains.

Mobility patterns affect the loads placed on the lower limbs during locomotion and may influence variation in lower limb diaphyseal robusticity and shape. This relationship commonly forms the basis for inferring mobility patterns from hominin fossil and skeletal remains. This study assesses the correspondence between athletic histories, varying by loading intensity, repetition and directionality, measured using a recall questionnaire, and peripheral quantitative computed tomography-derived measurements of tibial diaphysis rigidity and shape. Participants included male university varsity cross-country runners (n = 15), field hockey players (n = 15), and controls (n = 20) [mean age: 22.1 (SD +/- 2.6) years]. Measurements of tibial rigidity (including J, %CA, Imax, Imin, and average cortical thickness) of both runners and field hockey players were greater than controls (P < or = 0.05). Differences in tibial shape (Imax/Imin, P < or = 0.05) between runners and hockey players reflect pronounced maximum plane (Imax) rigidity in runners, and more symmetrical hypertrophy (Imax, Imin) among hockey players. This corresponds with the generally unidirectional locomotor patterns of runners, and the multidirectional patterns of hockey players. These results support the relationship between mobility and tibial diaphysis morphology as it is generally interpreted in the anthropological literature, with greater levels of mobility associated with increased diaphyseal robusticity and shape variation. Although exercise intensity may be the primary influence on these properties, the repetitiveness of the activity also deserves consideration. In conclusion, bone morphological patterns can reflect habitual behaviors, with adaptation to locomotor activities likely contributing to variation in tibial rigidity and shape properties in archaeological and fossil samples.