Human shoulder development is adapted to obstetrical constraints.

In humans, obstetrical difficulties arise from the large head and broad shoulders of the neonate relative to the maternal birth canal. Various characteristics of human cranial development, such as the relatively small head of neonates compared with adults and the delayed fusion of the metopic suture, have been suggested to reflect developmental adaptations to obstetrical constraints. On the other hand, it remains unknown whether the shoulders of humans also exhibit developmental features reflecting obstetrical adaptation. Here we address this question by tracking the development of shoulder width from fetal to adult stages in humans, chimpanzees, and Japanese macaques. Compared with nonhuman primates, shoulder development in humans follows a different trajectory, exhibiting reduced growth relative to trunk length before birth and enhanced growth after birth. This indicates that the perinatal developmental characteristics of the shoulders likely evolved to ease obstetrical difficulties such as shoulder dystocia in humans.

Skipping without and with hurdles in bipedal macaque: global mechanics.

Macaques trained to perform bipedally used running gaits across a wide range of speeds. At higher speeds they preferred unilateral skipping (galloping). The same asymmetric stepping pattern was used while hurdling across two low obstacles placed at the distance of a stride within our experimental track. In bipedal macaques during skipping, we expected a differential use of the trailing and leading legs. The present study investigated global properties of the effective and virtual leg, the location of the virtual pivot point (VPP), and the energetics of the center of mass (CoM), with the aim of clarifying the differential leg operation during skipping in bipedal macaques. When skipping, macaques displayed minor double support and aerial phases during one stride. Asymmetric leg use was indicated by differences in leg kinematics. Axial damping and tangential leg work did not influence the indifferent peak ground reaction forces and impulses, but resulted in a lift of the CoM during contact of the leading leg. The aerial phase was largely due to the use of the double support. Hurdling amplified the differential leg operation. Here, higher ground reaction forces combined with increased double support provided the vertical impulse to overcome the hurdles. Following CoM dynamics during a stride, skipping and hurdling represented bouncing gaits. The elevation of the VPP of bipedal macaques resembled that of human walking and running in the trailing and leading phases, respectively. Because of anatomical restrictions, macaque unilateral skipping differs from that of humans, and may represent an intermediate gait between grounded and aerial running.

Direct visualization and measurement of the plantar aponeurosis behavior in foot arch deformation via the windlass mechanism.

The plantar aponeurosis (PA) is an elastic longitudinal band that contributes to the generation of a propulsive force in the push-off phase during walking and running through the windlass mechanism. However, the dynamic behavior of the PA remains unclear owing to the lack of direct measurement of the strain it generates. Therefore, this study aimed to visualize and quantify the PA behavior during two distinct foot postures: (i) neutral posture and (ii) windlass posture with midtarsal joint plantarflexion and metatarsophalangeal joint dorsiflexion, using computed tomography scans. Six healthy adult males participated in the experiment, and three-dimensional reconstruction of the PA was conducted to calculate its path length, width, thickness, and cross-sectional area. This study successfully visualized and quantified the morphological changes in the PA induced by the windlass mechanism, providing a precise reference for biomechanical modeling. This study also highlighted the interindividual variability in the PA morphology and stretching patterns. Although the windlass posture was not identical to that observed in the push-off phase during walking, the observed PA behavior provides valuable insights into its mechanics and potential implications for foot disorders.

Peak vertical ground force of hand-knee crawling in human adults.

[Purpose] Fall risk is immanent in humans because they are bipedal. Bipedalism has transited from quadrupedalism in both evolutional and developmental contexts. Past studies have measured the peak vertical ground force of forelimbs and hindlimbs in quadrupedalism; and revealed that load dominancy shifted from forelimbs to hindlimbs during evolution. The dominance of hindlimb peak vertical ground force allows forelimb freedom and is considered important for locomotor evolution toward bipedalism. With this consideration, we hypothesize that hindlimb peak vertical ground force is dominant in human adults when they designedly crawl in a quadrupedal manner. [Participants and Methods] Six healthy human adults crawled on their hands and knees over a pressure platform. We calculated the peak vertical ground force of their hands and knees by integrating the pressure of the contact area of each limb. [Results] The mean knee peak vertical ground force at 0.694 (per body weight) was significantly higher than that of the hand at 0.372 (per body weight). The mean hand/knee peak vertical ground force ratio was 0.536; therefore, it was -0.624 on the natural logarithmic scale. [Conclusions] Our findings on human adults are compatible with existing considerations on locomotor evolution toward bipedalism. Our findings contribute to the comprehensive understanding of human locomotion.

Relationship between flightlessness and brain morphology among Rallidae.

Studies have suggested that the brain morphology and flight ability of Aves are interrelated; however, such a relationship has not been thoroughly investigated. This study aimed to examine whether flight ability, volant or flightless, affects brain morphology (size and shape) in the Rallidae, which has independently evolved to adapt secondary flightlessness multiple times within a single taxonomic group. Brain endocasts were extracted from computed tomography images of the crania, measured by 3D geometric morphometrics, and were analyzed using principal component analysis. The results of phylogenetic ANCOVA showed that flightless rails have brain sizes and shapes that are significantly larger than and different from those of volant rails, even after considering the effects of body mass and brain size respectively. Flightless rails tended to have a wider telencephalon and more inferiorly positioned foramen magnum than volant rails. Although the brain is an organ that requires a large amount of metabolic energy, reduced selective pressure for a lower body weight may have allowed flightless rails to have larger brains. The evolution of flightlessness may have changed the position of the foramen magnum downward, which would have allowed the support of the heavier cranium. The larger brain may have facilitated the acquisition of cognitively advanced behavior, such as tool-using behavior, among rails.

New femoral remains of Nacholapithecus kerioi: Implications for intraspecific variation and Miocene hominoid evolution.

The middle Miocene stem kenyapithecine Nacholapithecus kerioi (16-15 Ma; Nachola, Kenya) is represented by a large number of isolated fossil remains and one of the most complete skeletons in the hominoid fossil record (KNM-BG 35250). Multiple fieldwork seasons performed by Japanese-Kenyan teams during the last part of the 20th century resulted in the discovery of a large sample of Nacholapithecus fossils. Here, we describe the new femoral remains of Nacholapithecus. In well-preserved specimens, we evaluate sex differences and within-species variation using both qualitative and quantitative traits. We use these data to determine whether these specimens are morphologically similar to the species holotype KNM-BG 35250 (which shows some plastic deformation) and to compare Nacholapithecus with other Miocene hominoids and extant anthropoids to evaluate the distinctiveness of its femur. The new fossil evidence reaffirms previously reported descriptions of some distal femoral traits, namely the morphology of the patellar groove. However, results also show that relative femoral head size in Nacholapithecus is smaller, relative neck length is longer, and neck-shaft angle is lower than previously reported for KNM-BG 35250. These traits have a strong functional signal related to the hip joint kinematics, suggesting that the morphology of the proximal femur in Nacholapithecus might be functionally related to quadrupedal-like behaviors instead of more derived antipronograde locomotor modes. Results further demonstrate that other African Miocene apes (with the exception of Turkanapithecus kalakolensis) generally fall within the Nacholapithecus range of variation, whose overall femoral shape resembles that of Ekembo spp. and Equatorius africanus. Our results accord with the previously inferred locomotor repertoire of Nacholapithecus, indicating a combination of generalized arboreal quadrupedalism combined with other antipronograde behaviors (e.g., vertical climbing).

Trunk and leg kinematics of grounded and aerial running in bipedal macaques.

Across a wide range of Froude speeds, non-human primates such as macaques prefer to use grounded and aerial running when locomoting bipedally. Both gaits are characterized by bouncing kinetics of the center of mass. In contrast, a discontinuous change from pendular to bouncing kinetics occurs in human locomotion. To clarify the mechanism underlying these differences in bipedal gait mechanics between humans and non-human primates, we investigated the influence of gait on joint kinematics in the legs and trunk of three macaques crossing an experimental track. The coordination of movement was compared with observations available for primates. Compared with human running, macaque leg retraction cannot merely be produced by hip extension, but needs to be supported by substantial knee flexion. As a result, despite quasi-elastic whole-leg operation, the macaque's knee showed only minor rebound behavior. Ankle extension resembled that observed during human running. Unlike human running and independent of gait, torsion of the trunk represents a rather conservative feature in primates, and pelvic axial rotation added to step length. Pelvic lateral lean during grounded running by macaques (compliant leg) and human walking (stiff leg) depends on gait dynamics at the same Froude speed. The different coordination between the thorax and pelvis in the sagittal plane as compared with human runners indicates different bending modes of the spine. Morphological adaptations in non-human primates to quadrupedal locomotion may prevent human-like operation of the leg and limit exploitation of quasi-elastic leg operation despite running dynamics.

Estimation of the External Knee Adduction Moment during Gait Using an Inertial Measurement Unit in Patients with Knee Osteoarthritis.

Although the external knee adduction moment (KAM) during gait was shown to be a quantitative parameter of medial knee osteoarthritis (OA), it requires expensive equipment and a dedicated large space to measure. Therefore, it becomes a major reason to limit KAM measurement in a clinical environment. The purpose of this study was to estimate KAM using a single inertial measurement unit (IMU) during gait in patients with knee OA. A total of 22 medial knee OA patients (44 knee joints) performed conventional gait analysis using three-dimensional (3D) motion capture system. At the same time, we attached commercial IMUs to six body segments (sternum, pelvis, both thighs, and both shanks), and IMU signals during gait were recorded synchronized with the motion capture system. The peak-to-peak difference of acceleration in the lateral/medial axis immediately after heel contact was defined as the thrust acceleration (TA). We hypothesized that TA would represent the lateral thrust of the knee during the stance phase and correlate with the first peak of KAM. The relationship between the peak KAM and TA of pelvis (R = 0.52, p < 0.001), shanks (R = 0.57, p < 0.001) and thighs (R = 0.49, p = 0.001) showed a significant correlation. The root mean square error (RMSE) of linear regression models of pelvis, shanks, and thighs to estimate KAM were 0.082, 0.079, and 0.084 Nm/(kg·m), respectively. Our newly established parameter TA showed a moderate correlation with conventional KAM. The current study confirmed our hypothesis that a single IMU would predict conventional KAM during gait. Since KAM is known as an indicator for prognosis and severity of knee OA, this new parameter has the potential to become an accessible predictor for medial knee OA instead of KAM.

Morphological variations of the human talus investigated using three-dimensional geometric morphometrics.

INTRODUCTION: The shape of the talus determines the positional and kinematic features of the subtalar, talonavicular, and talocrural joints during walking. Thus, detailed knowledge of the pattern of sexual dimorphism of the human talus may be useful for revealing the pathogenetic mechanism of foot and knee disorders, which are more prevalent in females. The aim of this study was to characterize and visualize the three-dimensional shape variations of the talus in relation to sex and age using geometric morphometrics. MATERIALS AND METHODS: Computed tomography images of 56 feet without talar injuries or disorders were used in this study. Thirty-seven anatomical landmarks were identified on a bone model of the talus to calculate principal components (PCs) of shape variations among specimens. PC scores were compared between sexes, and their correlations with age were also investigated. RESULTS: The female talus had a longer neck and narrower head width than the male talus. The superior trochlea was tilted more laterally in the frontal plane in females. Furthermore, the female talar head was more twisted and was more elongated in the dorsoplantar direction. CONCLUSIONS: Morphological features of the talus in females could alter the subtalar and talonavicular joint kinematics during walking and could be a structural factor in the pathogenetic mechanism underlying foot and knee disorders. This study contributes to the comprehensive understanding of shape variations in the human talus.

Three-dimensional geometric morphometrics of thorax-pelvis covariation and its potential for predicting the thorax morphology: A case study on Kebara 2 Neandertal.

The skeletal torso is a complex structure of outstanding importance in understanding human body shape evolution, but reconstruction usually entails an element of subjectivity as researchers apply their own anatomical expertise to the process. Among different fossil reconstruction methods, 3D geometric morphometric techniques have been increasingly used in the last decades. Two-block partial least squares analysis has shown great potential for predicting missing elements by exploiting the covariation between two structures (blocks) in a reference sample: one block can be predicted from the other one based on the strength of covariation between blocks. The first aim of this study is to test whether this predictive approach can be used for predicting thorax morphologies from pelvis morphologies within adult Homo sapiens reference samples with known covariation between the thorax and the pelvis. The second aim is to apply this method to Kebara 2 Neandertal (Israel, ∼60 ka) to predict its thorax morphology using two different pelvis reconstructions as predictors. We measured 134 true landmarks, 720 curve semilandmarks, and 160 surface semilandmarks on 60 3D virtual torso models segmented from CT scans. We conducted three two-block partial least squares analyses between the thorax (block 1) and the pelvis (block 2) based on the H. sapiens reference samples after performing generalized Procrustes superimposition on each block separately. Comparisons of these predictions in full shape space by means of Procrustes distances show that the male-only predictive model yields the most reliable predictions within modern humans. In addition, Kebara 2 thorax predictions based on this model concur with the thorax morphology proposed for Neandertals. The method presented here does not aim to replace other techniques, but to rather complement them through quantitative prediction of a virtual 'scaffold' to articulate the thoracic fossil elements, thus extending the potential of missing data estimation beyond the methods proposed in previous works.

Low leg compliance permits grounded running at speeds where the inverted pendulum model gets airborne.

Animals typically switch from grounded (no flight phases) to aerial running at dimensionless speeds u^ < 1. But some birds use grounded running far above u^ = 1, which puzzles biologists because the inverted pendulum becomes airborne at this speed. Here, we combine computer experiments using the spring-mass model with locomotion data from small birds, macaques and humans to understand the relationship between leg function (stiffness, angle of attack), locomotion speed and gait. With our model, we found three-humped ground reaction force profiles for slow grounded running speeds. The minimal single-humped grounded running speed is u^ = 0.4. This speed value roughly coincides with the transition speed from vaulting to bouncing mechanics in bipeds. Maximal grounded running speed in the model is not limited. In experiments, animals changed from grounded to aerial running at dimensionless contact time around 1. Considering these real-world contact times reduces the solution space drastically, but experimental data fit well. The model still predicts maximal grounded running speed  u^ > 1 for low stiffness values used by birds but decreases below u^ = 1 for increasing stiffness. For stiffer legs used in human walking and running, periodic grounded running vanishes. At speeds at which birds and macaques change to aerial running, we found periodic aerial running to intersect grounded running. This could explain why animals can alternate between grounded and aerial running at the same speed and identical leg parameters. Compliant legs enable different gaits and speeds with similar leg parameters, stiff legs require parameter adaptations.

Assessing thoraco-pelvic covariation in Homo sapiens and Pan troglodytes: A 3D geometric morphometric approach.

OBJECTIVES: Understanding thoraco-pelvic integration in Homo sapiens and their closest living relatives (genus Pan) is of great importance within the context of human body shape evolution. However, studies assessing thoraco-pelvic covariation across Hominoidea species are scarce, although recent research would suggest shared covariation patterns in humans and chimpanzees but also species-specific features, with sexual dimorphism and allometry influencing thoraco-pelvic covariation in these taxa differently. MATERIAL AND METHODS: N = 30 adult H. sapiens and N = 10 adult Pan troglodytes torso 3D models were analyzed using 3D geometric morphometrics and linear measurements. Effects of sexual dimorphism and allometry on thoraco-pelvic covariation were assessed via regression analyses, and patterns of thoraco-pelvic covariation in humans and chimpanzees were computed via Two-Block Partial Least Squares analyses. RESULTS: Results confirm the existence of common aspects of thoraco-pelvic covariation in humans and chimpanzees, and also species-specific covariation in H. sapiens that is strongly influenced by sexual dimorphism and allometry. Species-specific covariation patterns in chimpanzees could not be confirmed because of the small sample size, but metrics point to a correspondence between the most caudal ribs and iliac crest morphology that would be irrespective of sex. CONCLUSIONS: This study suggests that humans and chimpanzees share common aspects of thoraco-pelvic covariation but might differ in others. In humans, torso integration is strongly influenced by sexual dimorphism and allometry, whilst in chimpanzees it may not be. This study also highlights the importance not only of torso widths but also of torso depths when describing patterns of thoraco-pelvic covariation in primates. Larger samples are necessary to support these interpretations.

Three-Dimensional Morphological Variations of the Human Calcaneus Investigated Using Geometric Morphometrics.

The shape of the calcaneus determines the mechanical interaction of the foot with the ground during the heel-strike in human walking. Detailed knowledge of the pattern of sexual dimorphism of the human calcaneus could help to clarify the pathogenetic mechanism of foot and knee disorders, which are more prevalent in females. Therefore, the aim of this study was to characterize and visualize the three-dimensional shape variations of the calcaneus in relation to sex and age using geometric morphometrics. Computed tomography images of 56 feet without subtalar injuries or disorders were used in this study. Thirty-seven anatomical landmarks were identified on the bone model of the calcaneus to calculate principal components (PCs) of shape variations among specimens. The PC scores were compared between males and females, and their correlations with age were also analyzed. The female calcaneus was longer in length and shorter in height than that of males. The medial process of the calcaneal tuberosity in females was more inferiorly projected and the tuberosity was shifted more laterally. Also, the calcaneus was wider and the sustentaculum tali thickened with aging. Female structural features of the calcaneus alter the kinematics of the foot during walking and could be a structural factor in foot and knee disorders. This study contributes to a comprehensive understanding of shape variations in the human calcaneus. Clin. Anat., 33:751-758, 2020. © 2019 Wiley Periodicals, Inc.

Effect of natural full weight-bearing during standing on the rotation of the first metatarsal bone.

To evaluate the rotational change in the first metatarsal bone (1MT) of the foot during natural standing using an upright computed tomography (CT) scanner with 320-detector rows. A total of 52 feet of 28 asymptomatic subjects (aged 23-39 years) were evaluated in the natural standing position with or without weight-bearing. A foot pressure plate was used to determine the non-weight-bearing (NWB) or single leg full-weight-bearing (s-FWB) conditions. CT examinations were performed using a noise index of 15 for a slice thickness of 5 mm, rotation speed of 0.5 sec, and slice thickness of 0.5 mm. The rotation of the 1MT was measured on the coronal CT image, which cut the sesamoids' bellies in the frontal slide of the first metatarsal and sesamoids perpendicular to the longitudinal bisection of the third metatarsal, and compared between the weight-bearing conditions. Intra- and inter-observer reliabilities of the rotation angle were also evaluated. The intra- and inter-observer correlation coefficients were 0.961 and 0.934, respectively. The 1MT pronation angle was significantly greater in the s-FWB condition than in the NWB condition (15.2° ± 5.4° vs. 12.5° ± 5.3°, P < 0.01). No sex difference was found in the magnitude of the 1MT pronation angle as a result of weight-bearing. This study first demonstrated that pronation of 1MT occurs due to natural full-weight-bearing in asymptomatic feet. The 1MT's rotational movement under weight-bearing conditions may relate to the onset and pathogenesis of the hallux valgus. Clin. Anat. 32:715-721, 2019. © 2019 Wiley Periodicals, Inc.

Multivariate analysis of variations in intrinsic foot musculature among hominoids.

Comparative analysis of the foot muscle architecture among extant great apes is important for understanding the evolution of the human foot and, hence, human habitual bipedal walking. However, to our knowledge, there is no previous report of a quantitative comparison of hominoid intrinsic foot muscle dimensions. In the present study, we quantitatively compared muscle dimensions of the hominoid foot by means of multivariate analysis. The foot muscle mass and physiological cross-sectional area (PCSA) of five chimpanzees, one bonobo, two gorillas, and six orangutans were obtained by our own dissections, and those of humans were taken from published accounts. The muscle mass and PCSA were respectively divided by the total mass and total PCSA of the intrinsic muscles of the entire foot for normalization. Variations in muscle architecture among human and extant great apes were quantified based on principal component analysis. Our results demonstrated that the muscle architecture of the orangutan was the most distinctive, having a larger first dorsal interosseous muscle and smaller abductor hallucis brevis muscle. On the other hand, the gorilla was found to be unique in having a larger abductor digiti minimi muscle. Humans were distinguished from extant great apes by a larger quadratus plantae muscle. The chimpanzee and the bonobo appeared to have very similar muscle architecture, with an intermediate position between the human and the orangutan. These differences (or similarities) in architecture of the intrinsic foot muscles among humans and great apes correspond well to the differences in phylogeny, positional behavior, and locomotion.

Bipedal gait versatility in the Japanese macaque (Macaca fuscata).

It was previously believed that, among primates, only humans run bipedally. However, there is now growing evidence that at least some non-human primates can not only run bipedally but can also generate a running gait with an aerial phase. Japanese macaques trained for bipedal performances have been known to exhibit remarkable bipedal locomotion capabilities, but no aerial-phase running has previously been reported. In the present study, we investigated whether Japanese macaques could run with an aerial phase by collecting bipedal gait sequences from three macaques on a level surface at self-selected speeds (n = 188). During our experiments, body kinematics and ground reaction forces were recorded by a motion-capture system and two force plates installed within a wooden walkway. Our results demonstrated that macaques were able to utilize a variety of bipedal gaits including grounded running, skipping, and even running with an aerial phase. The self-selected bipedal locomotion speed of the macaques was fast, with Froude speed ranging from 0.4 to 1.3. However, based on congruity, no single trial that could be categorized as a pendulum-like walking gait was observed. The parameters describing the temporal, kinematic, and dynamic characteristics of macaque bipedal running gaits follow the patterns previously documented for other non-human primates and terrestrial birds that use running gaits, but are different from those of humans and from birds' walking gaits. The present study confirmed that when a Japanese macaque engages in bipedal locomotion, even without an aerial phase, it generally utilizes a spring-like running mechanism because the animals have a limited ability to stiffen their legs. That limitation is due to anatomical restrictions determined by the morphology and structure of the macaque musculoskeletal system. The general adoption of grounded running in macaques and other non-human primates, along with its absence in human bipedal locomotion, suggests that abandonment of compliant gait was a critical transition in the evolution of human obligatory bipedalism.

Sexual dimorphism of body size in an African fossil ape, Nacholapithecus kerioi.

Sexual size dimorphism in the African fossil ape Proconsul nyanzae (18 million years ago, 18 Ma) has been previously documented. However, additional evidence for sexual dimorphism in Miocene hominoids can provide great insight into the history of extant hominoid mating systems. The present study focused on body mass (BM) sexual dimorphism in Nacholapithecus kerioi from the Middle Miocene (16-15 Ma) in Africa. Bootstrap analysis revealed that P. nyanzae BM sexual dimorphism was lower than that in Pan troglodytes, which exhibits moderate sexual dimorphism, as reported previously. The same simulation revealed that BM sexual dimorphism of N. kerioi was comparable with that in Gorilla spp.; i.e., the males were approximately twice as large as the females. High sexual dimorphism in extant apes is usually indicative of a polygynous social structure (gorilla) or solitary/fission-fusion social system (orangutan). However, because of the high proportion of adult males in this fossil assemblage, the magnitude of dimorphism inferred here cannot be associated with a gorilla-like polygynous or oranguran-like solitary/fission-fusion social structure, and may reflect either taphonomic bias, or some other social structure. Extant hominoids have a long evolutionary history owing to their deep branching, comprising only a few existing members of the original highly successful group. Therefore, it is not surprising that the mating systems of extant hominoids fail to provide fossil apes with a perfect "model". The mating systems of extinct hominoids may have been more diverse than those of extant apes.

Global dynamics of bipedal macaques during grounded and aerial running.

Macaques trained to perform bipedally use grounded running, skipping and aerial running, but avoid walking. The preference for grounded running across a wide range of speeds is substantially different from the locomotion habits observed in humans, which may be the result of differences in leg compliance. In the present study, based on kinematic and dynamic observations of three individuals crossing an experimental track, we investigated global leg properties such as leg stiffness and viscous damping during grounded and aerial running. We found that, in macaques, similar to human and bird bipedal locomotion, the vector of the ground reaction force is directed from the center of pressure (COP) to a virtual pivot point above the center of mass (COM). The visco-elastic leg properties differ for the virtual leg (COM-COP) and the effective leg (hip-COP) because of the position of the anatomical hip with respect to the COM. The effective leg shows damping in the axial direction and positive work in the tangential component. Damping does not prevent the exploration of oscillatory modes. Grounded running is preferred to walking because of leg compliance. The transition from grounded to aerial running is not accompanied by a discontinuous change. With respect to dynamic properties, macaques seem to be well placed between bipedal specialists (humans and birds). We speculate that the losses induced in the effective leg by hip placement and slightly pronograde posture may not pay off by facilitating stabilization, making bipedal locomotion expensive and insecure for macaques.

3D geometric morphometrics of thorax variation and allometry in Hominoidea.

Ever since the seminal papers of Keith and Schultz, hominoid primate ribcages have been described as either "funnel-" or "barrel-shaped." Following this dichotomic typology, it is currently held that Homo sapiens and hylobatids (gibbons and siamangs) share a barrel-shaped ribcage and that they are more similar to each other than to the funnel-shaped thoraces of great apes (Gorilla, Pan, and Pongo). Other researchers hypothesized that thoracic width and the invagination of the thoracic spine into the thorax are related to allometry. However, analyses that take into account the complex three-dimensional (3D) shape of the ribcage are lacking. Here, we address hypotheses about thorax shape and evolution using 3D morphometrics of thoraces in anatomical connection obtained by computed tomography scans of 23 hominoid cadavers and 10 humans and examining thorax compartments composed of seven ribs (1-7 thorax) and of 11 ribs (1-11 thorax). In the 1-7 thorax analyses, the human thorax is uniquely flat because of torsion of the upper and central ribs, differing from all non-human hominoids including hylobatids. In the 1-11 thorax analyses, humans are markedly different from African great apes, with hylobatids and orangutans intermediate. In full shape space analyses, affinities between orangutans and humans on the one hand and between hylobatids and African great apes on the other are evident. Therefore, we reject the hypothesis that humans and hylobatids bear any special affinities in overall 3D thorax shape to each other. We find that larger thoraces are wider and flatter, with a more invaginated spine, supporting the allometric hypothesis. Hominoid thorax variation shows complex interactions between allometry, rib curves, torsion, and declination, and the morphology of the costo-vertebral joint and the thoracic vertebral column. When considering functional specializations alongside phylogenetic relationships, an overly simplistic dichotomy between funnel-shaped and barrel-shaped thoraces is not supported.

Age- and sex-associated morphological variations of metatarsal torsional patterns in humans.

It has been demonstrated that the torsional patterns of the metatarsal heads are associated with the presence or absence of the medial longitudinal arch in hominoid feet. The relatively untwisted second metatarsal is unique in humans, but that of the African apes is much more inverted, suggesting that the torsion of the second metatarsal might represent the overall shape and flatness of the foot. Some clinical studies have recently argued that the onset of foot pathologies such as hallux valgus might be related to the torsional pattern of the metatarsals. However, to date, no studies have systematically investigated the morphological variations of the torsional patterns of human metatarsals. In this study, therefore, the aim was to clarify the age- and sex-associated variations in the torsional patterns of human metatarsals using three-dimensional computed tomography. The torsion angles of the five metatarsals were calculated by defining the dorsopalmar vector of the metatarsal base and the vector corresponding to the rotational axis of the metatarsal head. The present result demonstrated that the second metatarsals of females were significantly more inverted with increasing age. Flat foot is known to be most common in elderly women. Whether there is a cause-effect relationship between second metatarsal torsion and flattening of the medial longitudinal arch has yet to be answered, but this study suggested that torsion of the second metatarsal might possibly be used as an indicator for the early diagnosis of flat foot and associated foot pathologies. Clin. Anat. 30:1058-1063, 2017. © 2017 Wiley Periodicals, Inc.