Scapular morphology of great apes and humans: A three-dimensional computed tomography-based comparative study.

The primate scapula has been studied widely since its shape has been shown to correlate with how the forelimb is used in daily activities. In this study, we expand on the existing literature and use an image-based methodology that was originally developed for orthopaedic practice to quantify and compare the three-dimensional (3D) morphology of the scapula across humans and great apes. We expect that this image-based approach will allow us to identify differences between great apes and humans that can be related to differences in mobility and loading regime of the shoulder. We hypothesize that gorillas and chimpanzees will have a similar scapular morphology, geared towards stability and weight-bearing in knuckle-walking, whilst the scapular morphology of orangutans is expected to be more similar to that of humans given their high glenohumeral mobility associated with their suspensory lifestyle. We made 3D reconstructions of computed tomography scans of 69 scapulae from four hominid genera (Pongo, Gorilla, Pan and Homo). On these 3D bone meshes, the inferior glenoid plane was determined, and subsequently, a set of bony landmarks on the scapular body, coracoid, and acromion were defined. These landmarks allowed us to measure a set of functionally relevant angles which represent acromial overhang, subacromial space and coracoacromial space. The angles that were measured are: the delto-fulcral triangle (DFT), comprising the alpha, beta, and delta angle, the acromion-glenoid angle (AGA), the coracoid-glenoid centre-posterior acromial angle (CGA), the anterior tilt (TA CGA) and the posterior tilt of the CGA (PT CGA). Three observers placed the landmarks on the 3D bone meshes, allowing us to calculate the inter-observer error. The main differences in the DFT were found between humans and the great apes, with small differences between the great apes. The DFT of humans was significantly lower compared to that of the great apes, with the smallest alpha (32.7°), smallest delta (45.7°) and highest beta angle (101.6°) of all genera. The DFT of chimpanzees was significantly higher compared to that of humans (p < 0.01), with a larger alpha (37.6°) and delta angle (54.5°) and smaller beta angle (87.9°). The mean AGA of humans (59.1°) was significantly smaller (p < 0.001) than that of gorillas (68.8°). The mean CGA of humans (110.1°) was significantly higher (p < 0.001) than in orangutans (92.9°). Humans and gorillas showed mainly a posterior tilt of their coracoacromial complex whilst chimpanzees showed mainly an anterior tilt. The coracoacromial complex of the orangutans was not tilted anteriorly or posteriorly. With our image-based method, we were able to identify morphological features of the scapula that differed significantly between hominid genera. However, we did not find an overall dichotomy in scapular morphology geared towards high stability (Pan/Gorilla) or high mobility (Homo/Pongo). Further research is needed to investigate the functional implications of these differences in scapular morphology.

Covariation between wrist bone morphology and maximal range of motion during ulnar deviation and supination in extant nonhuman primate taxa.

This study investigates the maximal range of motion (ROM) during wrist deviation and forearm rotation for five different primate genera and the possible correlation with the shape of the distal ulna, triquetrum and hamate. A two-block phylogenetic partial least square analysis was performed to test this covariation in a phylogenetic context, using shape coordinates and a matrix of maximal ROM data as input data. The results show that gibbons have the highest ROM for both ulnar deviation and supination, whereas Macaca exhibited the lowest ROM for supination, and Pan had the lowest ROM for ulnar deviation. These results can be attributed to differences in locomotor behaviour, as gibbons need a large wrist mobility in all directions for their highly arboreal lifestyle, whereas Macaca and Pan need a stable wrist during terrestrial locomotion. However, we found no correlation between distal ulna/triquetrum/hamate shape and maximal ROM during ulnar deviation and supination in the different primate taxa. A larger dataset, in combination with behavioural and biomechanical studies, is needed to establish form-function relationships of the primate hand, which will aid the functional interpretation of primate fossil remains.

The forearm and hand musculature of semi-terrestrial rhesus macaques (Macaca mulatta) and arboreal gibbons (fam.Hylobatidae). Part II. Quantitative analysis.

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.

The forearm and hand musculature of semi-terrestrial rhesus macaques (Macaca mulatta) and arboreal gibbons (Fam. Hylobatidae). Part I. Description and comparison of the muscle configuration.

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.

The digital human forearm and hand.

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.

Insights into the musculature of the bonobo hand.

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.

Impact of Osteoarthritis and Total Joint Arthroplasty on the Kinematics of the Trapeziometacarpal Joint: A Pilot Study.

PURPOSE: To quantify the effect of osteoarthritis (OA) and total trapeziometacarpal (TMC) joint replacement on thumb kinematics during the primary physiological motions of the thumb. METHODS: We included 4 female patients with stage III TMC OA. A computed tomography-based markerless method was used to quantify the 3-dimensional thumb kinematics in patients before and after TMC joint replacement surgery with the Arpe implant. RESULTS: Trapeziometacarpal OA led to a marked decrease of internal rotation and abduction of the first metacarpal (MC1) during thumb flexion and a decrease of MC1 adduction during thumb adduction. As a compensatory phenomenon, the trapezium displayed increased abduction. The absence of MC1 translation in the ball-and-socket implant seems to induce a decrease of MC1 adduction as well as a decrease of trapezium adduction during thumb adduction, compared with OA and healthy joints. Implant replacement displayed an unchanged MC1 flexion during thumb flexion and seemed to slightly increase MC1 axial rotation during thumb flexion and adduction. Abduction and adduction of the MC1 are limited and compensated by this somewhat increased axial rotation, allowing more efficient thumb opposition. CONCLUSIONS: The study highlights that advanced TMC OA mainly restricts the MC1 mobility. We also showed that, whereas total joint arthroplasty is able to restore thumb function, it cannot fully replicate the kinematics of the healthy TMC joint. CLINICAL RELEVANCE: The quantification of TMC joint kinematics in OA and implanted patients is essential to improve our understanding of TMC OA as well as to enhance the functionality of implant designs.

Ulnar shape of extant primates: Functional signals and covariation with triquetrum shape.

OBJECTIVES: In this study, we investigated the shape differences of the distal ulna in a phylogenetic context among a broad range of primate taxa. Furthermore, we evaluated covariation between ulnar and triquetrum shape and a possible association between ulnar shape and locomotor behavior. MATERIALS AND METHODS: We applied 3D geometric morphometrics on a large dataset comprising the distal ulna of 124 anthropoid primate specimens belonging to 12 different genera. For each species, a mean shape was calculated using 11 Procrustes-aligned surface landmarks on the distal ulna. These mean shapes are used in a bgPCA, pPCA, and PACA and 3D morphs were used to visualize more subtle differences between taxa. A p2B-PLS analysis was performed to test the covariance between distal ulnar and triquetrum shape. RESULTS: The results show that more closely related species exhibit a similar distal ulnar shape. Overall, extant hominid ulnae show a shape shift compared to those of extant monkeys and hylobatids. This includes a shortening of the ulnar styloid process and dorspalmarly widening of the ulnar head, shape characteristics that are independent of phylogeny. Within the hominids, Pongo pygmaeus seem to possess the most plesiomorphic distal ulnar shape, while Gorilla and Homo sapiens display the most derived distal ulna. Cercopithecoids, hylobatids, and P. pygmaeus are characterized by a relatively deep ECU groove, which is a shape trait dependent of phylogeny. Although there was no significant covariation between distal ulnar shape and triquetrum shape, the shape differences of the distal ulna between the different primate taxa reveal a possible link with locomotor behavior. CONCLUSIONS: The comparative analyses of this study reveal different shape trends in a phylogenetic context. Highly arboreal primates, such as hylobatids and Ateles fusciceps, show a distal ulnar morphology that appears to be adapted to tensile and torsional forces. In primates that use their wrist under more compressive conditions, such as quadrupedal cercopithecoids and great apes, the distal ulnar morphology seems to reflect increased compressive forces. In modern humans, the distal ulnar shape can be associated to enhanced manipulative skills and power grips. There was no significant covariation between distal ulnar shape and triquetrum shape, probably due to the variation in the amount of contact between the triquetrum and ulna. In combination with future research on wrist mobility in diverse primate taxa, the results of this study will allow us to establish form-function relationships of the primate wrist and contribute towards an evidence-based interpretation of fossil remains.

The role of hind limb tendons in gibbon locomotion: springs or strings?

Tendon properties have an important effect on the mechanical behaviour of muscles, with compliant tendons allowing near-isometric muscle contraction and facilitating elastic energy storage and recoil. Stiff tendons, in contrast, facilitate rapid force transfer and precise positional control. In humans, the long Achilles tendon contributes to the mechanical efficiency of running via elastic energy storage and recovery, and its presence has been linked to the evolution of habitual bipedalism. Gibbons also possess relatively long hind limb tendons; however, their role is as yet unknown. Based on their large dimensions, and inferring from the situation in humans, we hypothesize that the tendons in the gibbon hind limb will facilitate elastic energy storage and recoil during hind-limb-powered locomotion. To investigate this, we determined the material properties of the gibbon Achilles and patellar tendon in vitro and linked this with available kinematic and kinetic data to evaluate their role in leaping and bipedalism. Tensile tests were conducted on tendon samples using a material testing machine and the load-displacement data were used to calculate stiffness, Young's modulus and hysteresis. In addition, the average stress-in-life and energy absorption capacity of both tendons were estimated. We found a functional difference between the gibbon Achilles and patellar tendon, with the Achilles tendon being more suitable for elastic energy storage and release. The patellar tendon, in contrast, has a relatively high hysteresis, making it less suitable to act as elastic spring. This suggests that the gibbon Achilles tendon might fulfil a similar function as in humans, contributing to reducing the locomotor cost of bipedalism by acting as elastic spring, while the high stiffness of the patellar tendon might favour fast force transfer upon recoil and, possibly, enhance leaping performance.

Principal component and linear discriminant analyses for the classification of hominoid primate specimens based on bone shape data.

In this study, we tested the hypothesis that machine learning methods can accurately classify extant primates based on triquetrum shape data. We then used this classification tool to observe the affinities between extant primates and fossil hominoids. We assessed the discrimination accuracy for an unsupervised and supervised learning pipeline, i.e. with principal component analysis (PCA) and linear discriminant analysis (LDA) feature extraction, when tasked with the classification of extant primates. The trained algorithm is used to classify a sample of known fossil hominoids. For the visualization, PCA and uniform manifold approximation and projection (UMAP) are used. The results show that the discriminant function correctly classified the extant specimens with an F1-score of 0.90 for both PCA and LDA. In addition, the classification of fossil hominoids reflects taxonomy and locomotor behaviour reported in literature. This classification based on shape data using PCA and LDA is a powerful tool that can discriminate between the triquetrum shape of extant primates with high accuracy and quantitatively compare fossil and extant morphology. It can be used to support taxonomic differentiation and aid the further interpretation of fossil remains. Further testing is necessary by including other bones and more species and specimens per species extinct primates.

Functional adaptations in the forelimb muscles of non-human great apes.

The maximum capability of a muscle can be estimated from simple measurements of muscle architecture such as muscle belly mass, fascicle length and physiological cross-sectional area. While the hindlimb anatomy of the non-human apes has been studied in some detail, a comparative study of the forelimb architecture across a number of species has never been undertaken. Here we present data from chimpanzees, bonobos, gorillas and an orangutan to ascertain if, and where, there are functional differences relating to their different locomotor repertoires and habitat usage. We employed a combination of analyses including allometric scaling and ancovas to explore the data, as the sample size was relatively small and heterogeneous (specimens of different sizes, ages and sex). Overall, subject to possible unidentified, confounding factors such as age effects, it appears that the non-human great apes in this sample (the largest assembled to date) do not vary greatly across different muscle architecture parameters, even though they perform different locomotor behaviours at different frequencies. Therefore, it currently appears that the time spent performing a particular behaviour does not necessarily impose a dominating selective influence on the soft-tissue portion of the musculoskeletal system; rather, the overall consistency of muscle architectural properties both between and within the Asian and African apes strengthens the case for the hypothesis of a possible ancient shared evolutionary origin for orthogrady under compressive and/or suspensory loading in the great apes.

The extraordinary athletic performance of leaping gibbons.

The distance that animals leap depends on their take-off angle and velocity. The velocity is generated solely by mechanical work during the push-off phase of standing-start leaps. Gibbons are capable of exceptional leaping performance, crossing gaps in the forest canopy exceeding 10 m, yet possess none of the adaptations possessed by specialist leapers synonymous with maximizing mechanical work. To understand this impressive performance, we recorded leaps of the gibbons exceeding 3.7 m. Gibbons perform more mass-specific work (35.4 J kg(-1)) than reported for any other species to date, accelerating to 8.3 ms(-1) in a single movement and redefining our estimates of work performance by animals. This energy (enough for a 3.5 m vertical leap) is 60 per cent higher than that achieved by galagos, which are renowned for their remarkable leaping performance. The gibbons' unusual morphology facilitates a division of labour among the hind limbs, forelimbs and trunk, resulting in modest power requirements compared with more specialized leapers.

Trapeziometacarpal joint mobility in gibbons (fam. Hylobatidae) and rhesus macaques (Macaca mulatta).

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.

The effect of different grasping types on strain distributions in the trapezium of bonobos (Pan paniscus).

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.

Branching patterns of the vascularization and innervation of the primate forelimb.

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.

The effect of substrate compliance on the biomechanics of gibbon leaps.

The storage and recovery of elastic strain energy in the musculoskeletal systems of locomoting animals has been extensively studied, yet the external environment represents a second potentially useful energy store that has often been neglected. Recent studies have highlighted the ability of orangutans to usefully recover energy from swaying trees to minimise the cost of gap crossing. Although mechanically similar mechanisms have been hypothesised for wild leaping primates, to date no such energy recovery mechanisms have been demonstrated biomechanically in leapers. We used a setup consisting of a forceplate and two high-speed video cameras to conduct a biomechanical analysis of captive gibbons leaping from stiff and compliant poles. We found that the gibbons minimised pole deflection by using different leaping strategies. Two leap types were used: slower orthograde leaps and more rapid pronograde leaps. The slower leaps used a wider hip joint excursion to negate the downward movement of the pole, using more impulse to power the leap, but with no increase in work done on the centre of mass. Greater hip excursion also minimised the effective leap distance during orthograde leaps. The more rapid leaps conversely applied peak force earlier in stance where the pole was effectively stiffer, minimising deflection and potential energy loss. Neither leap type appeared to usefully recover energy from the pole to increase leap performance, but the gibbons demonstrated an ability to best adapt their leap biomechanics to counter the negative effects of the compliant pole.

The effect of orthoses on the kinematics of the trapeziometacarpal, scaphotrapeziotrapezoidal, and radioscaphoid joints.

The in vivo effect of four different types of thumb and thumb-wrist orthoses on the three-dimensional kinematics of the trapeziometacarpal (TMC), scaphotrapeziotrapezoidal (STT) and radioscaphoid joints was quantified using computed tomography (CT). Eighteen healthy female volunteers were recruited. The dominant hand of each subject was scanned in four thumb and wrist positions, each in three conditions: without orthosis, with a thumb orthosis (Push Ortho and immediate fitting, IMF) and with a thumb-wrist orthosis (Ligaflex Manu and IMF). CT images were analyzed and rotations relative to the more proximal bone were expressed in a joint-specific coordinate system. Without orthosis, the largest STT rotations were observed during radioulnar deviation of the wrist and the STT range of motion (ROM) was significantly lower during wrist flexion-extension. All tested orthoses caused a significant reduction of the ROM at each joint compared to free motion. Significant differences in movement reduction were observed between prefabricated and IMF orthoses.The IMF thumb-wrist outperformed the Ligaflex Manu in terms of immobilization of the radioscaphoid joint. In addition, the IMF thumb orthosis immobilized the TMC joint significantly better during thumb abduction and adduction than the Push Ortho. We found that different types of thumb and thumb-wrist orthotics are effective in reducing joint mobility. While this reduction tends to be higher using IMF compared to prefabricated orthoses, this effect is only significant for the radioscaphoid and TMC joint. The finding that thumb movements do not induce large STT rotations suggests that the thumb does not need to be immobilized in case of isolated STT osteoarthritis.

Stress distribution in the bonobo (Pan paniscus) trapeziometacarpal joint during grasping.

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.

Functional signals and covariation in triquetrum and hamate shape of extant primates using 3D geometric morphometrics.

In this study, we want to investigate the covariation in the shape of two carpal bones, the triquetrum and hamate, and the possible association with locomotor behavior in a broad range of primate taxa. We applied 3D Geometric Morphometrics on a large data set comprising 309 anthropoid primates of 12 different genera. Principal component analyses were performed on the covariance matrix of 18 (triquetrum) and 23 (hamate) Procrustes-aligned surface landmarks. A two-block partial least square analysis was done to test the covariance between triquetrum and hamate shape, without relying on the predictive models implicit in regression analyses. The results show that the carpal shape of quadrupedal anthropoids, which mainly use their wrist under compressive conditions, differs from that of suspensory primates as their wrist is possibly subjected to tensile and torsional forces. Within the hominids, differences in shape also distinguish more terrestrial from more arboreal species. Even within the great apes, we are able to capture shape differences between species of the same genus. In combination with behavioral and biomechanical studies, the results of this research can be used to establish form-function relationships of the primate hand which will aid the functional interpretation of primate fossil remains.

Muscle moment arms and function of the siamang forelimb during brachiation.

Moment arms have an important modulating impact on muscle function, as they represent the capacity of the muscle to convert muscle action into limb movements. In the current paper, we provide muscle moment arm data of the forelimb of four siamangs, collected by detailed dissections on unfixed cadavers. The aim of this study is to assess the role of different forelimb muscles during brachiation. Moment arm data are compared with similar published data of non-brachiating primates such as macaques, chimpanzees and humans. Our data show that shoulder adductors and endorotators and the elbow flexors are built for force generation, whereas the shoulder abductors, flexors and exorotators are best suited to gain speed and to change direction. Compared to non-brachiating species, both elbow and wrist flexors are particularly noticeable in terms of moment of force-generating capacity. However, the moment of force-generating capacity of the elbow extensor is not negligible, which indicates that the triceps also plays an active role, especially at the end of the support phase. Except for the elbow flexors, all muscles reach their maximum moment of force-generating capacity during the support phase of brachiation. When brachiating on a more complex setup, the siamang will flex the elbows to angles that induce maximum moment arms as well.