Comparing and combining sliding semilandmarks and weighted spherical harmonics for shape analysis.

Quantifying morphological variation is critical for conducting anatomical research. Three-dimensional geometric morphometric (3D GM) landmark analyses quantify shape using homologous Cartesian coordinates (landmarks). Setting up a high-density landmark set and placing it on all specimens, however, can be a time-consuming task. Weighted spherical harmonics (SPHARM) provides an alternative method for analyzing the shape of such objects. Here we compare sliding semilandmark and SPHARM analyses of the calcaneus of Gorilla gorilla gorilla (n = 20), Pan troglodytes troglodytes (n = 20), and Homo sapiens (n = 20) to determine whether the SPHARM and sliding semilandmark analyses capture comparable levels of shape variation. We also compare both the sliding semilandmark and SPHARM analyses to a novel combination of the two methods, here termed SPHARM-sliding. In SPHARM-sliding, the vertices of the surface models produced from the SPHARM analysis (that are the same in number and relative location) are used as the starting landmark positions for a sliding semilandmark analysis. Calcaneal shape variation quantified by all three analyses was summarized using separate principal components analyses. Results were compared using the root mean square (RMS) and maximum distance between surface models of species averages scaled (up) to centroid size created from each analysis. The average RMS was 0.23 mm between sliding semilandmark and SPHARM average surface models, 0.19 mm between SPHARM and SPHARM sliding average surface models, and 0.22 mm between sliding semilandmark and SPHARM sliding average surface models. Although results indicate that all three analyses are comparable methods for 3D shape analysis, there are advantages and disadvantages to each. While the SPHARM analysis is less time-intensive, it is unable to capture the same level of detail around the sharp edges of articular facets on average surface models as the sliding semilandmark analysis. The SPHARM analysis also does not allow for individual articular facets to be analyzed in isolation. SPHARM-sliding, however, captures the same level of detail as the sliding semilandmark analysis, and (as in the sliding semilandmark analysis) allows for the evaluation of individual portions of bone. SPHARM is a comparable method to a 3D GM analysis for small, irregularly shaped bones, such as the calcaneus, and SPHARM-sliding allows for an expedited set up process for a sliding semilandmark analysis.

A morphometric analysis of early Eocene Euprimate tarsals from Gujarat, India.

Early Eocene primate postcranial bones from the Vastan lignite mine of Gujarat, India, have proven useful for understanding the haplorhine and strepsirrhine divergence. Previous analyses of material assigned to Asiadapidae supported interpretations that these primates were generalized arboreal quadrupeds, while the omomyid Vastanomys was likely to have been more proficient leaper than asiadapids. More recent examinations of long bone cross-sectional properties and calcaneal elongation have complicated the behavioral interpretations of these fossils. This study examines whole talar and calcaneal morphology of the Vastan material to refine the locomotor reconstructions of these fossils. A comparative sample of extant primate species representing various locomotor behaviors was obtained by accessing surface models from MorphoSource.org. Surface models of fossil specimens attributed to Asiadapis cambayensis, Marcgodinotius indicus, and Vastanomys major were generated from micro-computed tomography scans. A morphological analysis was carried out using weighted spherical harmonics, a Fourier-based method that represents surfaces using coefficients associated with a common set of spherical harmonic functions. The coefficients describing each surface were then used as shape variables in a principal components analysis. Significant differences between locomotor groups were assessed using nonparametric tests. Results from extant comparative samples show that locomotor behavior can be predicted from both talar and calcaneal morphology when phylogenetic relationships are known. Consistent with previous analyses, our results indicate that Asiadapis cambayensis and Marcgodinotius indicus were likely arboreal quadrupeds with some leaping capabilities. Vastanomys major is reconstructed as an arboreal quadruped with greater leaping proficiency than its asiadapid counterparts based on its talar morphology.

Two Late Pleistocene human femora from Trinil, Indonesia: Implications for body size and behavior in Southeast Asia.

Late Pleistocene hominin postcranial specimens from Southeast Asia are relatively rare. Here we describe and place into temporal and geographic context two partial femora from the site of Trinil, Indonesia, which are dated stratigraphically and via Uranium-series direct dating to ca. 37-32 ka. The specimens, designated Trinil 9 and 10, include most of the diaphysis, with Trinil 9 being much better preserved. Microcomputed tomography is used to determine cross-sectional diaphyseal properties, with an emphasis on midshaft anteroposterior to mediolateral bending rigidity (Ix/Iy), which has been shown to relate to both body shape and activity level in modern humans. The body mass of Trinil 9 is estimated from cortical area and reconstructed length using new equations based on a Pleistocene reference sample. Comparisons are carried out with a large sample of Pleistocene and Holocene East Asian, African, and European/West Asian femora. Our results show that Trinil 9 has a high Ix/Iy ratio, most consistent with a relatively narrow-bodied male from a mobile hunting-gathering population. It has an estimated body mass of 55.4 kg and a stature of 156 cm, which are small relative to Late Pleistocene males worldwide, but larger than the penecontemporaneous Deep Skull femur from Niah Cave, Malaysia, which is very likely female. This suggests the presence of small-bodied active hunter-gatherers in Southeast Asia during the later Late Pleistocene. Trinil 9 also contrasts strongly in morphology with earlier partial femora from Trinil dating to the late Early-early Middle Pleistocene (Femora II-V), and to a lesser extent with the well-known complete Femur I, most likely dating to the terminal Middle-early Late Pleistocene. Temporal changes in morphology among femoral specimens from Trinil parallel those observed in Homo throughout the Old World during the Pleistocene and document these differences within a single site.

Calcaneal shape variation in humans, nonhuman primates, and early hominins.

The foot has played a prominent role in evaluating early hominin locomotion. The calcaneus, in particular, plays an important role in weight-bearing. Although the calcanei of early hominins have been previously scrutinized, a three-dimensional analysis of the entire calcaneal shape has not been conducted. Here, we investigate the relationship between external calcaneal shape and locomotion in modern Homo sapiens (n = 130), Gorilla (n = 86), Pan (n = 112), Pongo (n = 31), Papio (n = 28), and hylobatids (Hylobates, Symphalangus; n = 32). We use these results to place the calcanei attributed to Australopithecus sediba, A. africanus, A. afarensis, H. naledi, and Homo habilis/Paranthropus boisei into a locomotor context. Calcanei were scanned using either surface scanning or micro-CT and their external shape analyzed using a three-dimensional geometric morphometric sliding semilandmark analysis. Blomberg's K statistic was used to estimate phylogenetic signal in the shape data. Shape variation was summarized using a principal components analysis. Procrustes distances between all taxa as well as distances between each fossil and the average of each taxon were calculated. Blomberg's K statistic was small (K = 0.1651), indicating weak phylogenetic effects, suggesting variation is driven by factors other than phylogeny (e.g., locomotion or body size). Modern humans have a large calcaneus relative to body size and display a uniquely convex cuboid facet, facilitating a rigid midfoot for bipedalism. More arboreal great apes display relatively deeper cuboid facet pivot regions for increased midfoot mobility. Australopithecus afarensis demonstrates the most human-like calcaneus, consistent with obligate bipedalism. Homo naledi is primarily modern human-like, but with some intermediate traits, suggesting a different form of bipedalism than modern humans. Australopithecus africanus, A. sediba, and H. habilis/P. boisei calcanei all possess unique combinations of human and nonhuman ape-like morphologies, suggesting a combination of bipedal and arboreal behaviors.

Bilateral asymmetry and developmental plasticity of the humerus in modern humans.

OBJECTIVE: This study investigates bilateral asymmetry in the humerus of modern human populations with differing activity patterns to assess the relative plasticity of different bone regions in response to environmental influences, particularly the biomechanical demands of handedness. METHODS: External breadths, cross-sectional properties, and centroid sizes were used to quantify directional and absolute asymmetry of humeral diaphyseal, distal periarticular, and articular regions in six populations with differing subsistence strategies (total n = 244). Geometric section properties were measured using computed tomography at six locations along the distal humerus, while centroid sizes of the distal articular and periarticular regions, as well as eight segments of the diaphysis, were extracted from external landmark data. Bilateral asymmetries were compared between populations and sexes. Each property was also tested for correlation with bilateral asymmetry at 40% of bone length, which has been shown to correlate with handedness. RESULTS: Asymmetry is highest in the diaphysis, but significant through all distal bone regions. Asymmetry increases in the region of the deltoid tuberosity, and progressively declines distally through the shaft and distal periarticular region. Articular asymmetry is higher than periarticular asymmetry, approaching levels seen just proximal to the olecranon fossa, and is weakly but significantly correlated with diaphyseal asymmetry. Hunter-gatherers from Indian Knoll have significantly higher levels of asymmetry than other groups and are more sexually dimorphic, particularly in cross-sectional properties of the diaphysis. CONCLUSIONS: Humeral dimensions throughout the diaphysis, including regions currently used in taxonomic assignments of fossil hominins, likely respond to in vivo use, including population and sex-specific behaviors.

Gorilla calcaneal morphological variation and ecological divergence.

OBJECTIVES: The primate foot has been extensively investigated because of its role in weight-bearing; however, the calcaneus has been relatively understudied. Here we examine entire gorilla calcaneal external shape to understand its relationship with locomotor behavior. MATERIALS AND METHODS: Calcanei of Gorilla gorilla gorilla (n = 43), Gorilla beringei graueri (n = 20), and Gorilla beringei beringei (n = 15) were surface or micro-CT scanned. External shape was analyzed through a three-dimensional geometric morphometric sliding semilandmark analysis. Semilandmarks were slid relative to an updated Procrustes average in order to minimize the bending energy of the thin plate spline interpolation function. Shape variation was summarized using principal components analysis of shape coordinates. Procrustes distances between taxa averages were calculated and resampling statistics run to test pairwise differences. Linear measures were collected and regressed against estimated body mass. RESULTS: All three taxa exhibit statistically different morphologies (p < .001 for pairwise comparisons). G. g. gorilla demonstrates an anteroposteriorly elongated calcaneus with a deeper cuboid pivot region and mediolaterally flatter posterior talar facet. G. b. beringei possesses the flattest cuboid and most medially-angled posterior talar facets. G. b. graueri demonstrates intermediate articular facet morphology, a medially-angled tuberosity, and an elongated peroneal trochlea. DISCUSSION: Articular facet differences separate gorillas along a locomotor gradient. G. g. gorilla is adapted for arboreality with greater joint mobility, while G. b. beringei is adapted for more stereotypical loads associated with terrestriality. G. b. graueri's unique posterolateral morphology may be due to a secondary transition to greater arboreality from a more terrestrial ancestor.

Internal architecture of the mandibular condyle of rabbits is related to dietary resistance during growth.

Although there is considerable evidence that bone responds to the loading environment in which it develops, few analyses have examined phenotypic plasticity or bone functional adaptation in the masticatory apparatus. Prior work suggests that masticatory morphology is sensitive to differences in food mechanical properties during development; however, the importance of the timing/duration of loading and variation in naturalistic diets is less clear. Here, we examined microstructural and macrostructural differences in the mandibular condyle in four groups of white rabbits (Oryctolagus cuniculus) raised for a year on diets that varied in mechanical properties and timing of the introduction of mechanically challenging foods, simulating seasonal variation in diet. We employed sliding semilandmarks to locate multiple volumes of interest deep to the mandibular condyle articular surface, and compared bone volume fraction, trabecular thickness and spacing, and condylar size/shape among experimental groups. The results reveal a shared pattern of bony architecture across the articular surface of all treatment groups, while also demonstrating significant among-group differences. Rabbits raised on mechanically challenging diets have significantly increased bone volume fraction relative to controls fed a less challenging diet. The post-weaning timing of the introduction of mechanically challenging foods also influences architectural properties, suggesting that bone plasticity can extend well into adulthood and that bony responses to changes in loading may be rapid. These findings demonstrate that bony architecture of the mandibular condyle in rabbits responds to variation in mechanical loading during an organism's lifetime and has the potential to track dietary variation within and among species.

Estimating the in vivo location of the talus from external surface landmarks.

OBJECTIVES: Finite element analysis has gained popularity in anthropological research to connect morphological form to measurable function but requires that loads are applied at appropriate anatomical locations. This is challenging for the ankle because the joint surfaces are not easily determined given their deep anatomical location. While the location of the talonavicular and subtalar joints can be directly determined via medical imaging, regression equations are a common, less invasive method to estimate joint locations from surface anatomy. We propose a regression-based method to locate the in vivo positions of the talonavicular and subtalar joints employing three-dimensional (3D) surface markers, such as those used routinely in gait studies. METHODS: Navicular height was measured on weight-bearing radiographs (WBR) and simulated weight-bearing computed tomography (SWCT) scans to ensure SWCT correctly simulated foot weight-bearing configuration. The location of external foot markers and internal locations of the talonavicular and posterior subtalar joint were measured on each SWCT. Stepwise regression analysis was used to select the external markers that best predicted the three internal locations. RESULTS: Navicular heights measured on WBR and SWCT scans were not statistically different (p = .44), indicating that SWCTs recreate the weight-bearing position of the foot. The navicular tubercle and medial and lateral malleoli were the best predictors of subtalar and talonavicular joint locations. These palpable anatomical locations explained more variation in internal joint location (r2 > .79; SEE < 3.0 mm) than other landmarks. DISCUSSION: This study demonstrates that external palpable landmarks can predict the location of the talonavicular and subtalar joints.

Variation in obstetric dimensions of the human bony pelvis in relation to age-at-death and latitude.

OBJECTIVES: Previous studies associate females who died in young adulthood with narrower obstetric pelvic dimensions, presumably in association with obstetric insufficiency (though this causal relationship is unresolved). In this study, we examine whether females within groups living at higher latitudes present this pattern, as high-latitude groups have larger pelvic dimensions than groups previously examined. These patterns are compared with males. We assess whether there is evidence for younger ages-at-death in females to have been in response to natural selection against narrower true pelvis dimensions. METHODS: We measured 14 pelvic dimensions in 327 adults (188 females, 139 males), representing archaeological sites from mid-latitude and high-latitude North America. Individuals were placed into a "young" or "not young" age-at-death category. Latitude, sex, and age-at-death groups were compared using ANOVAs and scaled variance, and evidence for selection was examined with F-tests. RESULTS: Pelvic dimensions were larger in high-latitude females and males. Females but not males who died at younger ages had smaller pelvic canals than older individuals, especially in the mediolateral inlet and anteroposterior outlet dimensions. Variance in all pelvic dimensions is equal between the two female age-at-death groups. CONCLUSIONS: We found narrower obstetrical dimensions in the female pelvis among individuals who died at younger ages. However, statistically equivalent variances in the two female age-at-death groups does not support natural selection on pelvic dimensions as leading to younger ages at death. We instead argue that this difference is result of continued growth due to remodeling in the pelvis occurring in females, but not males, after early adulthood.

Long bone diaphyseal shape follows different ontogenetic trajectories in captive and wild gorillas.

OBJECTIVES: A number of studies have demonstrated the ontogenetic plasticity of long bone diaphyseal structure in response to mechanical loading. Captivity should affect mechanical loading of the limbs, but whether captive apes grow differently than wild apes has been debated. Here, we compare captive and wild juvenile and adult Gorilla to ascertain whether growth trajectories in cross-sectional diaphyseal shape are similar in the two environments. MATERIALS AND METHODS: A sample of young juvenile (n = 4) and adult (n = 10) captive Gorilla gorilla gorilla specimens, with known life histories, were compared with age-matched wild G.g. gorilla (n = 62) and G. beringei beringei (n = 75) in relative anteroposterior to mediolateral bending strength of the femur, tibia, and humerus. Cross sections were obtained using peripheral quantitative CT. RESULTS: Captive and wild adult G.g. gorilla differed in bending strength ratios for all three bones, but these differences were not present in young juvenile G.g. gorilla. In comparisons across taxa, captive juvenile G.g. gorilla were more similar to wild G.g. gorilla than to G.b. beringei, while captive adult G.g. gorilla were more similar in shape to G.b. beringei in the hind limb. DISCUSSION: Captive and wild G. gorilla follow different ontogenetic trajectories in long bone diaphyseal shape, corresponding to environmental differences and subsequent modified locomotor behaviors. Differences related to phylogeny are most evident early in development.

Trabecular mapping: Leveraging geometric morphometrics for analyses of trabecular structure.

OBJECTIVES: Trabecular microstructure of limb bone epiphyses has been used to elucidate the relationship between skeletal form and behavior among mammals. Such studies have often relied on the analysis of a single volume of interest (VOI). Here we present a method for evaluating variation in bone microstructure across articular surfaces by leveraging sliding semilandmarks. METHODS: Two samples were used to demonstrate the proposed methodology and test the hypothesis that microstructural variables are homogeneously distributed: tali from two ape genera (Pan and Pongo, n = 9) and modern human distal femora (n = 10). Sliding semilandmarks were distributed across articular surfaces and used to locate the position of multiple VOIs immediately deep to the cortical shell. Trabecular bone properties were quantified using the BoneJ plugin for ImageJ. Nonparametric MANOVA tests were used to make group comparisons and differences were explored using principal components analysis and visualized using color maps. RESULTS: Tests reveal that trabecular parameters are not distributed homogeneously and identify differences between chimpanzee and orangutan tali with regards to trabecular spacing and degree of anisotropy, with chimpanzee tali being more anisotropic and having more uniformly spaced trabeculae. Human males and females differed in the pattern of trabecular spacing with males having more uniform trabecular spacing across the joint surface. CONCLUSIONS: The proposed procedure quantifies variation in trabecular bone parameters across joint surfaces and allows for meaningful statistical comparisons between groups of interest. Consequently it holds promise to help elucidate links between trabecular bone structure and animal behavior.

The relative position of the human fibula to the tibia influences cross-sectional properties of the tibia.

OBJECTIVES: The fibula transmits loads within the lower limb of hominids. The few studies of variation in the cross-sectional geometric (CSG) properties of the fibula have established differences in its rigidity among groups engaged in distinct habitual loading activities. This study adds to this research by considering the relationship between CSG properties and the anatomical position of the fibula relative to the tibia among groups with differences in documented activity patterns. MATERIAL AND METHODS: We used pQCT scans taken at 50% of the length of the lower leg in 83 healthy young adult collegiate-aged individuals divided into five activity groups: runners, swimmers, cricketers, field hockey players, and non-athletes. We compared variation in calculated CSG properties against the distance between fibular and tibial centroids, as well as the angle of that plane relative to the plane of tibial Imax . RESULTS: Tibial and fibular CSG properties vary with respect to the relative position of the two bones. Tibial CSG properties differ in concert with the relative angle of the fibula to tibial Imax , while fibular CSG properties differ with the distance between the elements. Fibulae are more posterior-medially positioned in groups engaged in terrestrial athletics than among swimmers. DISCUSSION: The tibia and fibula experience different loads. The relative position of the two bones leads to compensatory differences in their CSG properties, perhaps due to increased resistance to bending in fibulae with greater distances from the tibia. Examinations of tibial CSG properties without considering the fibula limits interpretations about activity.

Fluctuating and directional asymmetry in the long bones of captive cotton-top tamarins (Saguinus oedipus).

OBJECTIVES: Skeletal asymmetries reflect developmental stability and mechanical, functional, and physiological influences on bone growth. In humans, researchers have documented the greatest limb bone bilateral asymmetry in diaphyseal breadths, with less asymmetry in articular and maximum length dimensions. However, it remains unclear as to whether the pattern observed for humans is representative of nonhuman primates, wherein bilateral loading may minimize directional asymmetry. This study adds to the small body of asymmetry data on nonhuman primates by investigating patterns of long bone asymmetry in a skeletal sample of Saguinus oedipus (cotton-top tamarin). MATERIALS AND METHODS: Humeri, radii, ulnae, femora, and tibiae of 76 adult captive cotton-top tamarin skeletons (48 males, 28 females) were measured bilaterally. We included maximum length, midshaft diaphyseal breadths, and at least one articular measurement for each bone to assess directional (DA) and fluctuating asymmetry (FA) in each dimension. RESULTS: Most dimensions exhibit significant FA, and very few have significant DA; DA is limited to the lower limb, especially in knee dimensions. Overall, the magnitudes of asymmetry in tamarins have a consistent ranking that follows the same pattern as found in humans. DISCUSSION: This first study of DA and FA among multiple dimensions throughout the limbs of a non-hominoid primate suggests that previously-reported patterns of human bilateral asymmetry are not exclusive to humans. The results further indicate potential underlying differences in constraints on variation within limb bones. While processes shaping variation await further study, our results argue that different long bone dimensions may reflect dissimilar evolutionary processes.

Macroscopic differences in adult human femora are linked to body mass index.

Bone functional adaptation is routinely invoked to interpret skeletal morphology despite ongoing debate regarding the limits of the bone response to mechanical stimuli. The wide variation in human body mass presents an opportunity to explore the relationship between mechanical load and skeletal response in weight-bearing elements. Here, we examine variation in femoral macroscopic morphology as a function of body mass index (BMI), which is used as a metric of load history. A sample of 80 femora (40 female; 40 male) from recent modern humans was selected from the Texas State University Donated Skeletal Collection. Femora were imaged using x-ray computed tomography (voxel size ~0.5 mm), and segmented to produce surface models. Landmark-based geometric morphometric analyses based on the Coherent Point Drift algorithm were conducted to quantify shape. Principal components analyses were used to summarize shape variation, and component scores were regressed on BMI. Within the male sample, increased BMI was associated with a mediolaterally expanded femoral shaft, as well as increased neck-shaft angle and decreased femoral neck anteversion angle. No statistically significant relationships between shape and BMI were found in the female sample. While mechanical stimulus has traditionally been applied to changes in long bong diaphyseal shape it appears that bone functional adaptation may also result in fundamental changes in the shape of skeletal elements.

An Introduction to The Orbital Buttresses.

Facial buttresses are supportive bony structures of the facial skeleton that form a thick, strong, and protective framework for the face. Surgical fixation may be required to restore morphology and function when damage to these buttresses occurs. We sought to determine if, similar to buttresses of the facial skeleton, buttresses of the internal orbit exist. Hence, we analyzed 10 human cadaver skulls imaged by microcomputed tomography (micro-CT). Image processing and thickness/heat mapping were performed using Avizo and ImageJ softwares. After identifying the orbital buttresses, we reviewed CT scans of patients who had orbital fractures across three years to determine the frequency of fracture of the orbital buttresses. We identified 5 buttresses of the internal orbit: superomedial fronto-ethmoidal strut with the deep orbital buttress, inferomedial strut with the posterior ledge, inferior orbital fissure, sphenoid-frontal superolateral strut, and the sphenoid lip. The average threshold orbital buttress thickness was 1.36 (0.25) mm. A total of 1186 orbits of 593 individuals were analyzed for orbital buttress involvement. Orbital buttresses were spared in 770 (65%) orbits. The inferomedial strut with the posterior ledge was the most commonly fractured buttress in 14.4% of orbits (n=171), followed by the sphenoid strut and lip (66 [5.6%]). To our knowledge, this is the first description of the buttresses of the internal orbit. Orbital reconstruction for fracture repair or oncologic purposes requires the support of orbital buttresses. Understanding the anatomy of orbital buttresses is crucial for successful surgical planning, proper implant positioning, and restoration of function and appearance.

Carpal allometry of African apes among mammals.

OBJECTIVES: Morphological variation in African ape carpals has been used to support the idea that Pan and Gorilla evolved knuckle-walking independently. Little work, however, has focused on the effect of body mass on carpal morphology. Here, we compare carpal allometry in Pan and Gorilla to that of other quadrupedal mammals with similar body mass differences. If allometric trends in Pan and Gorilla carpals mirror those of other mammals with similar body mass variation, then body mass differences may provide a more parsimonious explanation for African ape carpal variation than the independent evolution of knuckle-walking. MATERIALS AND METHODS: Three linear measurements were collected on the capitate, hamate, lunate, and scaphoid (or scapholunate) of 39 quadrupedal species from six mammalian families/subfamilies. Relationships between linear measurements and estimated body mass were analyzed using reduced major axis regression. Slopes were compared to 0.33 for isometry. RESULTS: Within Hominidae, higher body mass taxa (Gorilla) have relatively anteroposteriorly wider, mediolaterally wider, and/or proximodistally shorter capitates, hamates, and scaphoids than low body mass taxa (Pan). These allometric relationships are mirrored in most, but not all, mammalian families/subfamilies included in the analysis. CONCLUSIONS: Within most mammalian families/subfamilies, carpals of high body mass taxa are proximodistally shorter, anteroposteriorly wider, and mediolaterally wider than those of low body mass taxa. These distinctions may be caused by the need to accommodate relatively higher forelimb loading associated with greater body mass. Because these trends occur within multiple mammalian families/subfamilies, some carpal variation in Pan and Gorilla is consistent with body mass differences.

Hip width and metabolic energy expenditure of abductor muscles.

Despite a paucity of physiological evidence, simplistic biomechanical analyses have led researchers to assume that humans who have wider hips use more energy to walk. Pitting biomechanical first principles against physiological data has led to little deepening of our understanding of bipedalism and its evolution. Both approaches, however, use proxies for the energy used by muscles. We decided to approach the question directly. Using a musculoskeletal model of the human body that estimates the metabolic energy expenditure of muscle activation for 48 people (23 women), 752 trials were evaluated. Metabolic energy consumption for the abductor muscles was summed over a stride to create total abductor energy expenditure. We calculated the maximum hip joint moment acting in the coronal plane and the functional distance between the hip joint centers. We hypothesize that wider hips would be correlated with both maximum coronal plane hip moment and increased total abductor energy expenditure when mass and velocity were controlled. Linear regressions with multiple independent variables, clustered by participant to control for the non-independence of the data points, were performed in Stata. We found that hip width does not predict total abductor energy expenditure, although mass and velocity combine to predict 61% of the variation (both p<0.001). Maximum hip joint coronal plane moment is predicted by pelvic width (p<0.001) and, in combination with mass and velocity (both p<0.001), explains 79% of the variation. Our results indicate that people use their morphology in ways that limit differences in energy expenditure. Consistent with recent discussion, intraspecific variation might not be useful to understand differences among species.

Morphological variation of the Pan talus relative to that of Gorilla.

OBJECTIVES: Differences in talar articular morphology relative to locomotion have recently been found within Pan and Gorilla. Whole-bone talar morphology within, and shared variation among, Pan and Gorilla (sub)species, however, has yet to be investigated. Here we separately analyze talar external shape within Pan (P. t. troglodytes, P. t. schweinfurthii, P. t. verus, P. paniscus) and Gorilla (G. g. gorilla, G. b. beringei, G. b. graueri) relative to degree of arboreality and body size. Pan and Gorilla are additionally analyzed together to determine if consistent shape differences exist within the genera. MATERIALS AND METHODS: Talar external shape was quantified using a weighted spherical harmonic analysis. Shape variation both within and among Pan and Gorilla was described using principal component analyses. Root mean square distances were calculated between taxon averages, and resampling statistics conducted to test for pairwise differences. RESULTS: P. t. verus (most arboreal Pan) talar shape significantly differs from other Pan taxa (p < 0.05 for pairwise comparisons) driven by more asymmetrical trochlear rims and a medially-set talar head. P. t. troglodytes, P. t. schweinfurthii, and P. paniscus do not significantly differ (p > 0.05 for pairwise comparisons). All gorilla taxa exhibit significantly different talar morphologies (p < 0.007 for pairwise comparisons). The more terrestrial subspecies of G. beringei and P. troglodytes exhibit a superoinferiorly taller talar head/neck complex. DISCUSSION: P. t. verus exhibits talar morphologies that have been previously related to more frequent arboreality. The adaptations in the more terrestrial G. beringei and P. troglodytes subspecies may serve to facilitate load transmission.

Effects of obesity on talar micro- and macro-morphology.

The addition of information regarding obesity status to the forensic anthropological biological profile could significantly contribute to the identification of human skeletal remains since over 40% of the U.S. adult population is currently obese. This study examines the differences in talar shape and trabecular bone structure between obese and non-obese individuals. A sample of 20 obese and 20 non-obese divided evenly by sex was selected from the Texas State University Donated Skeletal Collection. Tali were imaged using x-ray computed tomography (voxel size: 28-38.7 µm). Image stacks were processed to produce binary images as well as trabecular thickness and spacing maps. Landmark-based geometric morphometric analyses were conducted to quantify shape variation. Shape coordinates were used to locate 100 geometrically homologous volumes of interest within each talus. Bone volume fraction, trabecular thickness, and trabecular spacing were extracted at each volume of interest. Within each sex, a one-way ANCOVA was used to determine if significant differences exist between obese and non-obese individuals in trabecular bone after controlling for age. The size of the talus as well as subtle aspects of shape were found to distinguish the sexes. The results further indicate that bone volume fraction significantly differs between obese and non-obese males. In females, bone volume fraction is correlated with age but does not differ between obese and non-obese. The study demonstrates that bone microstructure is a promising approach to estimating body mass or body mass index category but age effects diminish the potential for the talus to be used alone.

Variation in human limb joint articular morphology.

OBJECTIVES: Synovial joints in human limbs strike a balance between mobility, stability, and articular fit, yet little is known about how these conflicting demands pattern intraspecific variation in articular shape. In this study, we use geometric morphometrics to establish the apportionment and magnitude of morphological variance of the articular surfaces of the human shoulder, elbow, hip, and knee. We hypothesize that variances will be comparable between articulating surfaces within a joint and will be larger in joints with smaller ranges of motion, given their plurality of functional demands. MATERIALS AND METHODS: Three-dimensional landmarks were taken on the articular surfaces of the glenohumeral, humeroulnar, acetabulofemoral, and tibiofemoral joints from CT scans of 200 skeletons from the University of Tennessee Donated Skeletal Collection (84 females, 116 males). Root mean-squared distances between articulations calculated from Procrustes shape coordinates were used to determine variance distributions. RESULTS: We found no difference in variances for each articular surface between the sexes or between left and right articular surfaces. A high range of motion is associated with greater morphological variance; however, this pattern is largely driven by the concave articular surfaces of each joint, which consistently exhibit statistically greater variance than their convex counterparts. DISCUSSION: The striking pattern of differential variance between articulating morphologies points to potential disparities in development between them. Consistently higher variance in concave surfaces may relate to chondral modeling theory for the formation of joints. Establishing intraspecific morphological variance patterns is a first step in understanding coordinated evolution among articular features.