Simulating Adaptive Human Bipedal Locomotion Based on Phase Resetting Using Foot-Contact Information

Humans generate bipedal walking by cooperatively manipulating their complicated and redundant musculoskeletal systems to produce adaptive behaviors in diverse environments. To elucidate the mechanisms that generate adaptive human bipedal locomotion, we conduct numerical simulations based on a musculoskeletal model and a locomotor controller constructed from anatomical and physiological findings. In particular, we focus on the adaptive mechanism using phase resetting based on the foot-contact information that modulates the walking behavior. For that purpose, we first reconstruct walking behavior from the measured kinematic data. Next, we examine the roles of phase resetting on the generation of stable locomotion by disturbing the walking model. Our results indicate that phase resetting increases the robustness of the walking behavior against perturbations, suggesting that this mechanism contributes to the generation of adaptive human bipedal locomotion

New femoral remains of <i>Nacholapithecus kerioi</i>: 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)

Contributions of phase resetting and interlimb coordination to the adaptive control of hindlimb obstacle avoidance during locomotion in rats: a simulation study.

Obstacle avoidance during locomotion is essential for safe, smooth locomotion. Physiological studies regarding muscle synergy have shown that the combination of a small number of basic patterns produces the large part of muscle activities during locomotion and the addition of another pattern explains muscle activities for obstacle avoidance. Furthermore, central pattern generators in the spinal cord are thought to manage the timing to produce such basic patterns. In the present study, we investigated sensory-motor coordination for obstacle avoidance by the hindlimbs of the rat using a neuromusculoskeletal model. We constructed the musculoskeletal part of the model based on empirical anatomical data of the rat and the nervous system model based on the aforementioned physiological findings of central pattern generators and muscle synergy. To verify the dynamic simulation by the constructed model, we compared the simulation results with kinematic and electromyographic data measured during actual locomotion in rats. In addition, we incorporated sensory regulation models based on physiological evidence of phase resetting and interlimb coordination and examined their functional roles in stepping over an obstacle during locomotion. Our results show that the phase regulation based on interlimb coordination contributes to stepping over a higher obstacle and that based on phase resetting contributes to quick recovery after stepping over the obstacle. These results suggest the importance of sensory regulation in generating successful obstacle avoidance during locomotion

Human shoulder development is adapted to obstetrical constraints

ヒトは小さく生まれて大きく育つ --その秘密は鎖骨にあり--. 京都大学プレスリリース. 2022-04-13.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

チーター（Acinonyx jubatus）における半規管の配置と直交性について

The purpose of this study was to clarify the spatial orientation of the semicircular canals in the cheetah (Acinonyx jubatus) using computed tomography (CT). CT scan was performed on the skull of a cheetah. Scanning data were reconstructed into the cross-sectional images, and then the landmark coordinates were determined to define the planes of the semicircular canals and the cross-sections of the skull. In the cheetah, the lateral semicircular canal was parallel to the base of the skull. The angles between the anterior and posterior semicircular canals was lager than 90 degrees, and each canal showed the similar angle with the sagittal cross-section of the skull, although these planar directions were different. One limitation of this study is that here we provided only one specimen, and thus the present results need to be confirmed by examining a larger number of cases in future studies. However, the method outlined in this study provided quantitative descriptions of orientations and orthogonalities of the semicircular canals of the cheetah. This is potentially useful for indentifying the morphological features of the semicircular canals in felids

Sensory regulation of stance-to-swing transition in generation of adaptive human walking: A simulation study

In this paper, we investigated sensory mechanisms to regulate the transition from the stance to swing phases in the generation of adaptive human bipedal walking based on a neuromusculoskeletal model. We examined the contributions of the sensory information from the force-sensitive afferents in the ankle extensor muscle and from the position-sensitive afferents from the hip, inspired by a neuro-mechanical simulation for the stepping of the hind legs of cats. Our simulation results showed that the sensory signals related to the force in the ankle extensor muscle make a larger contribution than sensory signals related to the joint angle at the hip to produce robust walking against disturbances, as observed in the simulation results of cat locomotion. This suggests that such a sensorimotor mechanism is a general property and is also embedded in the neuro-control system of human bipedal walking

Stabilizing Function of the Musculoskeletal System for Periodic Motion

Animals generate various motions by cooperatively manipulating their complicated and redundant musculoskeletal systems controlled by the nervous system. To reveal the mechanism with which a steady movement is generated, much research has been performed that shows that musculoskeletal properties themselves can stabilze motion without a sensory feedback system if the muscle architecture and the parallel elastic elements within a muscle are tuned appropriately. In this paper, we show that the condition of stability is greatly eased by limiting the targeted movement to the periodic motion. The force-velocity relationship, which is one of the most famous properties of muscle, plays a decisive role in realizing the self-stabilizing ability of the musculoskeletal system

Evaluating functional roles of phase resetting in generation of adaptive human bipedal walking with a physiologically based model of the spinal pattern generator.

The central pattern generators (CPGs) in the spinal cord strongly contribute to locomotor behavior. To achieve adaptive locomotion, locomotor rhythm generated by the CPGs is suggested to be functionally modulated by phase resetting based on sensory afferent or perturbations. Although phase resetting has been investigated during fictive locomotion in cats, its functional roles in actual locomotion have not been clarified. Recently, simulation studies have been conducted to examine the roles of phase resetting during human bipedal walking, assuming that locomotion is generated based on prescribed kinematics and feedback control. However, such kinematically based modeling cannot be used to fully elucidate the mechanisms of adaptation. In this article we proposed a more physiologically based mathematical model of the neural system for locomotion and investigated the functional roles of phase resetting. We constructed a locomotor CPG model based on a two-layered hierarchical network model of the rhythm generator (RG) and pattern formation (PF) networks. The RG model produces rhythm information using phase oscillators and regulates it by phase resetting based on foot-contact information. The PF model creates feedforward command signals based on rhythm information, which consists of the combination of five rectangular pulses based on previous analyses of muscle synergy. Simulation results showed that our model establishes adaptive walking against perturbing forces and variations in the environment, with phase resetting playing important roles in increasing the robustness of responses, suggesting that this mechanism of regulation may contribute to the generation of adaptive human bipedal locomotion