The semicircular canal system of vertebrates helps coordinate body movements, including stabilization of gaze during locomotion. Quantitative phylogenetically informed analysis of the radius of curvature of the three semicircular canals in 91 extant and recently extinct primate species and 119 other mammalian taxa provide support for the hypothesis that canal size varies in relation to the jerkiness of head motion during locomotion. Primate and other mammalian species studied here that are agile and have fast, jerky locomotion have significantly larger canals relative to body mass than those that move more cautiously.generalized least-squares analysis ͉ mammals ͉ vestibular system

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...

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SummaryA number of methods for measuring anisotropy in trabecular bone using high-resolution X-ray computed tomography exist, which give different answers but have not been compared in detail. In this study, we examine the mean-intercept length (MIL), star volume distribution (SVD) and star length distribution (SLD) methods, their algorithmic implementation for three-dimensional (3D) data, and how their results relate to each other. A uniform ordered sampling scheme for determining which orientations to sample during analysis enhances the reproducibility of anisotropy and principal component direction determinations, with no evident introduction of biasing. This scheme also facilitates the creation of a 3D rose diagram that can be used to gain additional insights from the data. The directed secant algorithm that is frequently used for traversing pixel and voxel grids for these calculations is prone to bias unless a previously unreported normalization is used. This normalization ameliorates the bias present when using cubic voxels, and also permits calculations on data sets in which the slice spacing is not equal to the pixel spacing. Overall, the three methods for quantification of anisotropy give broadly similar results, but there are systematic divergences that can be traced to their differences in data and processing, and which may impact on their relative utility in estimating mechanical properties. Although discussed in the context of computed tomography of trabecular bone, the methods described here may be applied to any 3D data set from which fabric information is desired.

Most analyses of trabecular microarchitecture in mammals have focused on the functional significance of interspecific variation, but they have not effectively considered the influence of body size or phylogeny on bone architecture. The goals of this study were to determine the relationship between trabecular bone and body size in the humeral and femoral heads of extant primates, and to assess the influence of phylogeny on bone microstructure. Using a sample of 235 individuals from 34 primate species, ranging in body size from 0.06 to 130 kg, the relationships between trabecular bone structure and body size were assessed by using conventional and phylogenetic regression analyses. Bone volume fraction, trabecular thickness and trabecular spacing increase with body size, whereas bone surface-area-to-volume ratio decreases. Shape variables such as trabecular number, connectivity density and degree of anisotropy scale inversely with size. Most of these variables scale with significant negative allometry, except bone surface-area-to-volume ratio, which scales with slight positive allometry. Phylogenetic regressions indicate a relatively weak phylogenetic signal in some trabecular bone variables. These data demonstrate that, relative to body size, large primates have thinner and more tightly packed trabeculae than small primates. The relatively thin trabeculae in large primates and other mammals, coupled with constraints on trabecular thickness related to osteocyte function, suggest that increased skeletal loads in the postcranial joints of large mammals are probably mitigated not only through alterations in trabecular microarchitecture, but also through other mechanisms such as changes in cortical bone distribution, limb posture and gait speed.

Understanding the mechanically-mediated response of trabecular bone to locomotion-specific loading patterns would be of great benefit to comparative mammalian evolutionary morphology. Unfortunately, assessments of the correspondence between individual trabecular bone features and inferred behavior patterns have failed to reveal a strong locomotion-specific signal. This study assesses the relationship between inferred locomotor activity and a suite of trabecular bone structural features that characterize bone architecture. High-resolution computed tomography images were collected from the humeral and femoral heads of 115 individuals from eight anthropoid primate genera (Alouatta, Homo, Macaca, Pan, Papio, Pongo, Trachypithecus, Symphalangus). Discriminant function analyses reveal that subarticular trabecular bone in the femoral and humeral heads is significantly different among most locomotor groups. The results indicate that when a suite of femoral head trabecular features is considered, trabecular number and connectivity density, together with fabric anisotropy and the relative proportion of rods and plates, differentiate locomotor groups reasonably well. A similar, yet weaker, relationship is also evident in the trabecular architecture of the humeral head. The application of this multivariate approach to analyses of trabecular bone morphology in recent and fossil primates may enhance our ability to reconstruct locomotor behavior in the fossil record.