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Objectives: Recent studies have associated subarticular trabecular bone distribution in the extant hominid first metacarpal (Mc1) with observed thumb use, to infer fossil hominin thumb use. Here, we analyze the entire Mc1 to test for interspecific differences in: (1) the absolute volume of trabecular volume fraction, (2) the distribution of the deeper trabecular network, and (3) the distribution of trabeculae in the medullary cavity, especially beneath the Mc1 disto-radial flange., Materials and Methods: Trabecular bone was imaged using micro-computed tomography in a sample of Homo sapiens (n = 11), Pan paniscus (n = 10), Pan troglodytes (n = 11), Gorilla gorilla (n = 10) and Pongo sp., (n = 7). Using Canonical Holistic Morphometric Analysis (cHMA), we tested for interspecific differences in the trabecular bone volume fraction (BV/TV) and its relative distribution (rBV/TV) throughout the Mc1, including within the head, medullary cavity, and base., Results: P. paniscus had the highest, and H. sapiens the lowest, BV/TV relative to other species. rBV/TV distribution statistically distinguished the radial concentrations and lack of medullary trabecular bone in the H. sapiens Mc1 from all other hominids. H. sapiens and, to a lesser extent, G. gorilla also had a significantly higher trabecular volume beneath the disto-radial flange relative to other hominids., Discussion: These results are consistent with differences in observed thumb use in these species and may also reflect systemic differences in bone volume fraction. The trabecular bone extension into the medullary cavity and concentrations beneath the disto-radial flange may represent crucial biomechanical signals that will aid in the inference of fossil hominin thumb use., (© 2023 The Authors. American Journal of Biological Anthropology published by Wiley Periodicals LLC.)

Objectives: Analyses of external bone shape using geometric morphometrics (GM) and cross-sectional geometry (CSG) are frequently employed to investigate bone structural variation and reconstruct activity in the past. However, the association between these methods has not been thoroughly investigated. Here, we analyze whole bone shape and CSG variation of metacarpals 1-5 and test covariation between them., Materials and Methods: We analyzed external metacarpal shape using GM and CSG of the diaphysis at three locations in metacarpals 1-5. The study sample includes three modern human groups: crew from the shipwrecked Mary Rose (n = 35 metacarpals), a Pre-industrial group (n = 50), and a Post-industrial group (n = 31). We tested group differences in metacarpal shape and CSG, as well as correlations between these two aspects of metacarpal bone structure., Results: GM analysis demonstrated metacarpus external shape variation is predominately related to changes in diaphyseal width and articular surface size. Differences in external shape were found between the non-pollical metacarpals of the Mary Rose and Pre-industrial groups and between the third metacarpals of the Pre- and Post-industrial groups. CSG results suggest the Mary Rose and Post-industrial groups have stronger metacarpals than the Pre-industrial group. Correlating CSG and external shape showed significant relationships between increasing external robusticity and biomechanical strength across non-pollical metacarpals (r: 0.815-0.535; p ≤ 0.05)., Discussion: Differences in metacarpal cortical structure and external shape between human groups suggest differences in the type and frequency of manual activities. Combining these results with studies of entheses and kinematics of the hand will improve reconstructions of manual behavior in the past., (© 2023 The Authors. American Journal of Biological Anthropology published by Wiley Periodicals LLC.)

The trabecular bone morphology of adult extant primates has been shown to reflect mechanical loading related to locomotion. However, ontogenetic studies of humans and other mammals suggest an adaptive lag between trabecular bone response and current mechanical loading patterns that could result in adult trabecular bone morphology reflecting juvenile behaviours. This study investigates ontogenetic changes in the trabecular bone structure of the third metacarpal of mountain gorillas (Gorilla beringei beringei; n = 26) and western lowland gorillas (Gorilla gorilla gorilla; n = 26) and its relationship to expected changes in locomotor loading patterns. Results show that trabecular bone reflects predicted mechanical loading throughout ontogeny. Bone volume fraction, trabecular thickness and trabecular number are low at birth and increase with age, although degree of anisotropy remains relatively stable throughout ontogeny. A high concentration of bone volume fraction can be observed in the distopalmar region of the third metacarpal epiphysis in early ontogeny, consistent with the high frequency of climbing, suspensory and other grasping behaviours in young gorillas. High trabecular bone concentration increases dorsally in the epiphysis during the juvenile period as terrestrial knuckle-walking becomes the primary form of locomotion. However, fusion of the epiphysis does not take place until 10-11 years of age, and overall trabecular structure does not fully reflect the adult pattern until 12 years of age, indicating a lag between adult-like behaviours and adult-like trabecular morphology. We found minimal differences in trabecular ontogeny between mountain and western lowland gorillas, despite presumed variation in the frequencies of arboreal locomotor behaviours. Altogether, ontogenetic changes in Gorilla metacarpal trabecular structure reflect overall genus-level changes in locomotor behaviours throughout development, but with some ontogenetic lag that should be considered when drawing functional conclusions from bone structure in extant or fossil adolescent specimens., (© 2022 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

Objectives: The dexterity of fossil hominins is often inferred by assessing the comparative manual anatomy and behaviors of extant hominids, with a focus on the thumb. The aim of this study is to test whether trabecular structure is consistent with what is currently known about habitually loaded thumb postures across extant hominids., Materials and Methods: We analyze first metacarpal (Mc1) subarticular trabecular architecture in humans (Homo sapiens, n = 10), bonobos (Pan paniscus, n = 10), chimpanzees (Pan troglodytes, n = 11), as well as for the first time, gorillas (Gorilla gorilla gorilla, n = 10) and orangutans (Pongo sp., n = 1, Pongo abelii, n = 3 and Pongo pygmaeus, n = 5). Using a combination of subarticular and whole-epiphysis approaches, we test for significant differences in relative trabecular bone volume (RBV/TV) and degree of anisotropy (DA) between species., Results: Humans have significantly greater RBV/TV on the radiopalmar aspects of both the proximal and distal Mc1 subarticular surfaces and greater DA throughout the Mc1 head than other hominids. Nonhuman great apes have greatest RBV/TV on the ulnar aspect of the Mc1 head and the palmar aspect of the Mc1 base. Gorillas possessed significantly lower DA in the Mc1 head than any other taxon in our sample., Discussion: These results are consistent with abduction of the thumb during forceful "pad-to-pad" precision grips in humans and, in nonhuman great apes, a habitually adducted thumb that is typically used in precision and power grips. This comparative context will help infer habitual manipulative and locomotor grips in fossil hominins., (© 2019 Wiley Periodicals, Inc.)

Trabecular bone remodels during life in response to loading and thus should, at least in part, reflect potential variation in the magnitude, frequency and direction of joint loading across different hominid species. Here we analyse the trabecular structure across all non-pollical metacarpal distal heads (Mc2-5) in extant great apes, expanding on previous volume of interest and whole-epiphysis analyses that have largely focused on only the first or third metacarpal. Specifically, we employ both a univariate statistical mapping and a multivariate approach to test for both inter-ray and interspecific differences in relative trabecular bone volume fraction (RBV/TV) and degree of anisotropy (DA) in Mc2-5 subchondral trabecular bone. Results demonstrate that whereas DA values only separate Pongo from African apes (Pan troglodytes, Pan paniscus, Gorilla gorilla), RBV/TV distribution varies with the predicted loading of the metacarpophalangeal (McP) joints during locomotor behaviours in each species. Gorilla exhibits a relatively dorsal distribution of RBV/TV consistent with habitual hyper-extension of the McP joints during knuckle-walking, whereas Pongo has a palmar distribution consistent with flexed McP joints used to grasp arboreal substrates. Both Pan species possess a disto-dorsal distribution of RBV/TV, compatible with multiple hand postures associated with a more varied locomotor regime. Further inter-ray comparisons reveal RBV/TV patterns consistent with varied knuckle-walking postures in Pan species in contrast to higher RBV/TV values toward the midline of the hand in Mc2 and Mc5 of Gorilla, consistent with habitual palm-back knuckle-walking. These patterns of trabecular bone distribution and structure reflect different behavioural signals that could be useful for determining the behaviours of fossil hominins., (© 2019 Anatomical Society.)

Previously, a micro-finite element (micro-FE)-based inverse remodelling method was presented in the literature that reconstructs the loading history of a bone based on its architecture alone. Despite promising preliminary results, it remains unclear whether this method is sensitive enough to detect differences of bone loading related to pathologies or habitual activities. The goal of this study was to test the sensitivity of the inverse remodelling method by predicting joint loading histories of metacarpal bones of species with similar anatomy but clearly distinct habitual hand use. Three groups of habitual hand use were defined using the most representative primate species: manipulation (human), suspensory locomotion (orangutan), and knuckle-walking locomotion (bonobo, chimpanzee, gorilla). Nine to ten micro-computed tomography scans of each species ([Formula: see text] in total) were used to create micro-FE models of the metacarpal head region. The most probable joint loading history was predicted by optimally scaling six load cases representing joint postures ranging from [Formula: see text] (extension) to [Formula: see text] (flexion). Predicted mean joint load directions were significantly different between knuckle-walking and non-knuckle-walking groups ([Formula: see text]) and in line with expected primary hand postures. Mean joint load magnitudes tended to be larger in species using their hands for locomotion compared to species using them for manipulation. In conclusion, this study shows that the micro-FE-based inverse remodelling method is sensitive enough to detect differences of joint loading related to habitual manual activities of primates and might, therefore, be useful for palaeoanthropologists to reconstruct the behaviour of extinct species and for biomedical applications such as detecting pathological joint loading.

Objectives: Humans display an 85-95% cross-cultural right-hand bias in skilled tasks, which is considered a derived behavior because such a high frequency is not reported in wild non-human primates. Handedness is generally considered to be an evolutionary byproduct of selection for manual dexterity and augmented visuo-cognitive capabilities within the context of complex stone tool manufacture/use. Testing this hypothesis requires an understanding of when appreciable levels of right dominant behavior entered the fossil record. Because bone remodels in vivo, skeletal asymmetries are thought to reflect greater mechanical loading on the dominant side, but incomplete preservation of external morphology and ambiguities about past loading environments complicate interpretations. We test if internal trabecular bone is capable of providing additional information by analyzing the thumb of Homo sapiens and Pan., Materials and Methods: We assess trabecular structure at the distal head and proximal base of paired (left/right) first metacarpals using micro-CT scans of Homo sapiens (n = 14) and Pan (n = 9). Throughout each epiphysis we quantify average and local bone volume fraction (BV/TV), degree of anisotropy (DA), and elastic modulus (E) to address bone volume patterning and directional asymmetry., Results: We find a right directional asymmetry in H. sapiens consistent with population-level handedness, but also report a left directional asymmetry in Pan that may be the result of postural and/or locomotor loading., Conclusion: We conclude that trabecular bone is capable of detecting right/left directional asymmetry, but suggest coupling studies of internal structure with analyses of other skeletal elements and cortical bone prior to applications in the fossil record., (© 2016 Wiley Periodicals, Inc.)

Almécija and colleagues claim that we apply a simplified understanding of bone functional adaptation and that our results of human-like hand use in Australopithecus africanus are not novel. We argue that our results speak to actual behavior, rather than potential behaviors, and our functional interpretation is well supported by our methodological approach, comparative sample, and previous experimental data., (Copyright © 2015, American Association for the Advancement of Science.)

The distinctly human ability for forceful precision and power "squeeze" gripping is linked to two key evolutionary transitions in hand use: a reduction in arboreal climbing and the manufacture and use of tools. However, it is unclear when these locomotory and manipulative transitions occurred. Here we show that Australopithecus africanus (~3 to 2 million years ago) and several Pleistocene hominins, traditionally considered not to have engaged in habitual tool manufacture, have a human-like trabecular bone pattern in the metacarpals consistent with forceful opposition of the thumb and fingers typically adopted during tool use. These results support archaeological evidence for stone tool use in australopiths and provide morphological evidence that Pliocene hominins achieved human-like hand postures much earlier and more frequently than previously considered., (Copyright © 2015, American Association for the Advancement of Science.)

Bone is capable of adapting during life in response to stress. Therefore, variation in locomotor and manipulative behaviours across extant hominoids may be reflected in differences in trabecular bone structure. The hand is a promising region for trabecular analysis, as it is the direct contact between the individual and the environment and joint positions at peak loading vary amongst extant hominoids. Building upon traditional volume of interest-based analyses, we apply a whole-epiphysis analytical approach using high-resolution microtomographic scans of the hominoid third metacarpal to investigate whether trabecular structure reflects differences in hand posture and loading in knuckle-walking (Gorilla, Pan), suspensory (Pongo, Hylobates and Symphalangus) and manipulative (Homo) taxa. Additionally, a comparative phylogenetic method was used to analyse rates of evolutionary changes in trabecular parameters. Results demonstrate that trabecular bone volume distribution and regions of greatest stiffness (i.e., Young's modulus) correspond with predicted loading of the hand in each behavioural category. In suspensory and manipulative taxa, regions of high bone volume and greatest stiffness are concentrated on the palmar or distopalmar regions of the metacarpal head, whereas knuckle-walking taxa show greater bone volume and stiffness throughout the head, and particularly in the dorsal region; patterns that correspond with the highest predicted joint reaction forces. Trabecular structure in knuckle-walking taxa is characterised by high bone volume fraction and a high degree of anisotropy in contrast to the suspensory brachiators. Humans, in which the hand is used primarily for manipulation, have a low bone volume fraction and a variable degree of anisotropy. Finally, when trabecular parameters are mapped onto a molecular-based phylogeny, we show that the rates of change in trabecular structure vary across the hominoid clade. Our results support a link between inferred behaviour and trabecular structure in extant hominoids that can be informative for reconstructing behaviour in fossil primates.

Trabecular architecture was assessed by 3D micro-computed tomography from spherical volumes of interest located within the head and base of metacarpals (MC) 1, 2, and 5 from n = 19 adult common chimpanzees. Two subspecies, West African Pan troglodytes verus from the Taï Forest, Côte d'Ivoire (n = 12) and Central African P. t. troglodytes from Cameroon (n = 7), were studied. For the combined sample, the metacarpal head is distinguished by greater bone volume fraction across all metacarpals, though the MC 1 is distinctive in having thicker, more plate-like trabeculae. The architecture in the MC 2 and MC 5 can be related to strains associated with terrestrial knuckle-walking. In particular, the relatively robust MC 5 head architecture may result from functional loading incurred during braking and use of a palm-in hand posture. Examining differences between samples, we found that the Cameroon chimpanzees possess a more robust architecture across all metacarpals in the form of greater bone volume fraction, higher connectivity, and somewhat more plate-like structure. These differences are not explicable in terms of population distinctions in body size or daily travel distance, but possibly reflect a combination of more terrestrial knuckle-walking in the Cameroon sample and more diverse hand postures and precision handling required of nut-cracking in West African chimpanzees., (2010 Wiley-Liss, Inc.)

Micro-computed tomographic analyses of trabecular bone architecture have been used to clarify the link between positional behavior and skeletal anatomy in primates. However, there are methodological decisions associated with quantifying and comparing trabecular anatomy across taxa that vary greatly in body size and morphology that can affect characterizations of trabecular architecture, such as choice of the volume of interest (VOI) size and location. The potential effects of these decisions may be amplified in small, irregular-shaped bones of the hands and feet that have more complex external morphology and more heterogeneous trabecular structure compared to, for example, the spherical epiphysis of the femoral head. In this study we investigate the effects of changes in VOI size and location on standard trabecular parameters in two bones of the hand, the capitate and third metacarpal, in a diverse sample of nonhuman primates that vary greatly in morphology, body mass and positional behavior. Results demonstrate that changes in VOI location and, to a lesser extent, changes in VOI size had a dramatic affect on many trabecular parameters, especially trabecular connectivity and structure (rods vs. plates), degree of anisotropy, and the primary orientation of the trabeculae. Although previous research has shown that some trabecular parameters are susceptible to slight variations in methodology (e.g. VOI location, scan resolution), this study provides a quantification of these effects in hand bones of a diverse sample of primates. An a priori understanding of the inherent biases created by the choice of VOI size and particularly location is critical to robust trabecular analysis and functional interpretation, especially in small bones with complex arthroses., (© 2010 The Authors. Journal of Anatomy © 2010 Anatomical Society of Great Britain and Ireland.)

For comparative 3D microCT studies of trabecular bone, the use of a volume of interest (VOI) scaled to body size may avoid over-sampling the trabecular mass in smaller versus larger-bodied taxa and comparison of regions that are not functionally homologous (Fajardo and Müller: Am J Phys Anthropol 115 (2001) 327-336), though the influence on quantitative analyses using scaled versus nonscaled VOIs remains poorly characterized. We compare trabecular architectural properties reflecting mass, organization, and orientation from three volumes of interest (large, scaled, and small) obtained from the distal first metacarpal in a sample of Homo (n = 10) and Pan (n = 12). We test the null hypotheses that neither absolute VOI size, nor scaling of the VOI to metacarpal size as a proxy for body size, biases intraspecific analyses nor impacts the detection of interspecific differences. These hypotheses were only partially supported. While certain properties (e.g., bone volume fraction or trabecular thickness) were not affected by varying VOI size within taxa, others were significantly impacted (e.g., intersection surface, connectivity, and structure). In comparing large versus scaled VOIs, we found that the large VOI inflated the number and/or magnitude of significant differences between Homo and Pan. In summary, our results support the use of scaled VOIs in studies of trabecular architecture., (2010 Wiley-Liss, Inc.)