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In this study, we investigate the branching patterns of the vascularization and innervation of the primate forelimb by performing detailed dissections of five unembalmed nonhuman primate specimens belonging to five different species, that is, rhesus macaque (Macaca mulatta), white-handed gibbon (Hylobates lar), Western gorilla (Gorilla gorilla), chimpanzee (Pan troglodytes), and bonobo (Pan paniscus). Results are compared with five embalmed human specimens (Homo sapiens), and anatomical data of previous studies on nonhuman primates are also included to provide a broader comparative framework. The results show that the overall configuration of the forelimb blood vessels and nerves of the different primate species is similar, although some apparent interspecific differences are found. In all nonhuman primates, in contrast to humans, the superficial vena basilica is absent. Moreover, in gorilla, chimpanzee, and bonobo, the superficial vena cephalica is confined to the forearm. In humans, both an arteria interossea anterior and posterior are present, while in nonhuman primates, only an arteria interossea anterior is present, which migrates to the posterior side at the level of the musculus pronator quadratus. For the innervation, the nervus (n.) medianus and n. ulnaris connect in the forearm of the gorilla and macaque. In the gibbon, the brachial plexus shows some differences in the branching pattern at the fasciculus level compared to the other primates. We conclude that the forelimb innervation branching pattern shows some minor differences between the nonhuman primate species, compared to higher plasticity in the vascularization. However, the exact functional implications of these differences still remain unclear. Therefore, more research on a broader range of primate species and sampling more specimens for each taxon are needed. ispartof: JOURNAL OF MORPHOLOGY vol:283 issue:9 pages:1273-1284 ispartof: location:United States status: published

Nonhuman primates have a highly diverse locomotor repertoire defined by an equally diverse hand use. Based on how primates use their hands during locomotion, we can distinguish between terrestrial and arboreal taxa. The 'arboreal' hand is likely adapted towards high wrist mobility and grasping, whereas the 'terrestrial' hand will show adaptations to loading. While the morphology of the forearm and hand bones have been studied extensively, functional adaptations in the forearm and hand musculature to locomotor behaviour have been documented only scarcely. In this paper, we investigate the forelimb musculature of the highly arboreal gibbons (including Hylobates lar,Hylobates pileatus,Nomascus leucogenys,Nomascus concolor and Symphalangus syndactylus) and compare this with the musculature of the semi-terrestrial rhesus macaques (Macaca mulatta). Anatomical data from previous dissections on knuckle-walking bonobos (Pan paniscus) and bipedal humans (Homo sapiens) are also included to further integrate the analyses in the scope of catarrhine hand adaptation. This study indicates that the overall configuration of the arm and hand musculature of these primates is very similar but there are some apparent differences in relative size which can be linked to differences in forelimb function and which might be related to their specific locomotor behaviour. In macaques, there is a large development of wrist deviators, wrist and digital flexors, and m. triceps brachii, as these muscles are important during the different phases of palmi- and digitigrade quadrupedal walking to stabilize the wrist and elbow. In addition, their m. flexor carpi ulnaris is the most important contributor to the total force-generating capacity of the wrist flexors and deviators, and is needed to counteract the adducting torque at the elbow joint during quadrupedal walking. Gibbons show a relatively high force-generating capacity in their forearm rotators, wrist and digital flexors, which are important muscles in brachiation to actively regulate forward movement of the body. The results also stress the importance of the digital flexors in bonobos, during climbing and clambering, and in humans, which is likely linked to our advanced manipulation skills. ispartof: JOURNAL OF ANATOMY vol:238 issue:2 pages:321-337 ispartof: location:England status: published

OBJECTIVES: The purpose of this study is to investigate the differences in 3D kinematics of the trapeziometacarpal (TMC) joint between gibbons (fam. Hylobatidae) and macaques (Macaca mulatta), two non-human primate groups with a distinct locomotor behavior. Gibbons are highly arboreal species, while macaques are quadrupeds. Here, we investigate the mobility and structural constraints of the TMC joint in both these primates and evaluate the hypothesis that differences in locomotor mode are reflected in joint structure and function. MATERIALS AND METHODS: We have developed an innovative software suite allowing for the quantification of in situ 3D kinematics based on medical imaging of the primate TMC joint using a unique sample of eight gibbons and seven macaques. These analyses are further supported by detailed dissection of the surrounding ligaments. RESULTS: The data demonstrate distinct differences in TMC joint mobility between gibbons and macaques, with wide ranges of motion in the gibbon TMC joint and restricted movement in macaques. Furthermore, the dissections show little dissimilarity in ligament anatomy that could be associated with the differences in TMC joint capabilities. CONCLUSION: We conclude that gibbons possess a highly mobile TMC joint and the ball-and-socket morphology allows for large ranges of motion. This type of morphology, however, does not offer much inherent stabilization. Lack of structural joint reinforcement suggests that gibbons may have difficulty in performing any type of power grasp with high loads. Macaques, on the other hand, are shown to have a considerably reinforced TMC joint, which is likely related to the habitual loading of the thumb during locomotion. ispartof: American Journal Of Physical Anthropology vol:177 issue:4 pages:708-718 ispartof: location:United States status: published

The thumb has played a key role in primate evolution due to its involvement in grasping and manipulation. A large component of this wide functionality is by virtue of the uniquely shaped trapeziometacarpal (TMC) joint. This TMC joint allows for a broad range of functional positions, but how its bone structure is adapted to withstand such a large variety of loading regimes is poorly understood. Here, we outline a novel, integrated finite element - micro finite element (FE-µFE) workflow to analyse strain distributions across the internal bony architecture. We have applied this modelling approach to study functional adaptation in the bonobo thumb. More specifically, the approach allows us to evaluate how strain is distributed through the trapezium upon loading of its distal articular facet. As loading conditions, we use pressure distributions for different types of grasping that were estimated in a previous study. Model evaluation shows that the simulated strain values fall within realistic boundaries of the mechanical response of bone. The results show that the strain distributions between the simulated grasps are highly similar, with dissipation towards the proximo-ulnar cluster of trabeculae regardless of trapezial bone architecture. This study presents an innovative FE-µFE approach to simulating strain distributions, and yields insight in the functional adaptation of the TMC joint in bonobos.

The primate thumb plays a central role in grasping and the basal trapeziometacarpal (TMC) joint is critical to its function. The TMC joint morphology varies across primates, yet little is known about form-function interaction within in the TMC joint. The purpose of this study was to investigate how stress distributions within the joint differ between five grasping types commonly employed by bonobos (Pan paniscus). Five cadaveric bonobo forearms were CT scanned in five standardized positions of the hand as a basis for the generation of parametric finite element models to compare grasps. We have developed a finite element analysis (FEA) approach to investigate stress distribution patterns in the TMC joint associated with each grasp type. We hypothesized that the simulated stress distributions for each position would correspond with the patterns expected from a saddle-shaped joint. However, we also expected differences in stress patterns arising from instraspecific variations in morphology. The models showed a high agreement between simulated and expected stress patterns for each of the five grasps (86% of successful simulations), while partially (52%) and fully (14%) diverging patterns were also encountered. We identified individual variations of key morphological features in the bonobo TMC joint that account for the diverging stress patterns and emphasized the effect of interindividual morphological variation on joint functioning. This study gives unprecedented insight in the form-function interactions in the TMC joint of the bonobo and provides an innovative FEA approach to modelling intra-articular stress distributions, a valuable tool for the study of the primate thumb biomechanics. ispartof: Peerj vol:9 ispartof: location:United States status: published

Primates live in very diverse environments and, as a consequence, show an equally diverse locomotor behaviour. During locomotion, the primate hand interacts with the superstrate and/or substrate and will therefore probably show adaptive signals linked with this locomotor behaviour. Whereas the morphology of the forearm and hand bones have been studied extensively, the functional adaptations in the hand musculature have been documented only scarcely. To evaluate whether there are potential adaptations in forelimb musculature to locomotor behaviour, we investigated the forearm and hand musculature of the highly arboreal gibbons (including Hylobates lar, Hylobates pileatus, Nomascus leucogenys, Nomascus concolor, Symphalangus syndactylus) and compared this with the musculature of the semi-terrestrial rhesus macaques (Macaca mulatta) by performing complete and detailed dissections on a sample of 15 unembalmed specimens. We found that the overall configuration of the upper arm and hand musculature is highly comparable between arboreal gibbons and semi-terrestrial macaques, and follows the general primate condition. Most of the identified differences in muscle configuration are located in the forearm. In macaques, a prominent m. epitrochleoanconeus is present, which potentially helps to extend the forearm and/or stabilize the elbow joint during quadrupedal walking. The m. flexor carpi radialis shows a more radial insertion in gibbons, which might be advantageous during brachiation, as it can aid radial deviation. The fingers of macaques are controlled in pairs by the m. extensor digiti secondi et tertii proprius and the m. extensor digiti quarti et quinti proprius-a similar organization can also be found in their flexors-which might aid in efficient positioning of the hand and fingers on uneven substrates during quadrupedal walking. In contrast, extension of the little finger in gibbons is controlled by a separate m. extensor digiti minimi, whereas digits 2 to 4 are extended by the m. extensor digitorum brevis, suggesting that simultaneous extension of digits 2-4 in gibbons might be important when reaching or grasping an overhead support during brachiation. In conclusion, the overall configuration of the forelimb and hand musculature is very similar in gibbons and macaques, with some peculiarities which can be linked to differences in forelimb function and which might be related to the specific locomotor behaviour of each group. ispartof: Journal Of Anatomy vol:237 issue:4 pages:774-790 ispartof: location:England status: published

The human thumb is specialized for manual tasks as it is no longer typically involved in locomotion. However, members of the genus Pan - the closest extant relatives of modern humans - also have a highly mobile thumb, which allows complex manual tasks such as tool-crafting and use. Here, we investigate the thumb kinematics of bonobos (Pan paniscus) in relation to the morphology of their trapeziometacarpal joint using unembalmed bonobo specimens and compare that with the human condition. We use computed tomography-based models of skeletal elements of the thumb during positions of maximum abduction/adduction and flexion/extension to determine the kinematics of the first metacarpal in bonobos and healthy human volunteers. In addition, the 3D geometry of the trapeziometacarpal joint is quantified and, together with an assessment of the ligaments surrounding the joint, is related to the obtained kinematics of the first metacarpal. Our results show a similar trapezial 3D morphology and similar kinematics of the first metacarpal in bonobos and humans, with a markedly higher extension of the first metacarpal in humans. This study provides an integrated analysis of thumb anatomy and kinematics in a unique sample of bonobo specimens.

How changes in anatomy affect joint biomechanics can be studied using musculoskeletal modelling, making it a valuable tool to explore joint function in healthy and pathological joints. However, gathering the anatomical, geometrical and physiological data necessary to create a model can be challenging. Very few integrated datasets exist and even less raw data is openly available to create new models. Therefore, the goal of the present study is to create an integrated digital forearm and make the raw data available via an open-access database. An un-embalmed cadaveric arm was digitized using 7T MRI and CT scans. 3D geometrical models of bones, cartilage, muscle and muscle pathways were created. After MRI and CT scanning, physiological muscle parameters (e.g. muscle volume, mass, length, pennation angle, physiological cross-sectional area, tendon length) were obtained via detailed dissection. After dissection, muscle biopsies were fixated and confocal microscopy was used to visualize and measure sarcomere lengths. This study provides an integrated anatomical dataset on which complete and accurate musculoskeletal models of the hand can be based. By creating a 3D digital human forearm, including all relevant anatomical parameters, a more realistic musculoskeletal model can be created. Furthermore, open access to the anatomical dataset makes it possible for other researchers to use these data in the development of a musculoskeletal model of the hand. ispartof: Journal of Anatomy vol:233 issue:5 pages:557-566 ispartof: location:England status: published

The human hand is well known for its unique dexterity which is largely facilitated by a highly mobile, long and powerful thumb that enables both tool manufacturing and use, a key component of human evolution. The bonobo (Pan paniscus), the closest extant relative to modern humans together with the chimpanzee (Pan troglodytes), also possesses good manipulative capabilities but with a lower level of dexterity compared with modern humans. Despite the close phylogenetic relationship between bonobos and humans, detailed quantitative data of the bonobo forelimb musculature remains largely lacking. To understand how morphology may influence dexterity, we investigated the functional anatomy of the bonobo hand using a unique sample of eight bonobo cadavers, along with one chimpanzee and one human (Homo sapiens) cadaver. We performed detailed dissections of unembalmed specimens to collect quantitative datasets of the extrinsic and intrinsic hand musculature, in addition to qualitative descriptions of the forelimb muscle configurations, allowing estimation of force-generating capacities for each functional group. Furthermore, we used medical imaging to quantify the articular surface of the trapeziometacarpal joint to estimate the intra-articular pressure. Our results show that the force-generating capacity for most functional groups of the extrinsic and intrinsic hand muscles in bonobos is largely similar to that of humans, with differences in relative importance of the extensors and rotators. The bonobo thumb musculature has a lower force-generating capacity than observed in the human specimen, but the estimated maximal intra-articular pressure is higher in bonobos. Most importantly, bonobos show a higher degree of functional coupling between the muscles of the thumb, index and lateral fingers than observed in humans. It is conceivable that differentiation and individualization of the hand muscles rather than relative muscle development explain the higher level of dexterity of humans compared with that of bonobos. ispartof: Journal of Anatomy vol:233 issue:3 pages:328-340 ispartof: location:England status: published