Naomichi Ogihara

Generation of human bipedal locomotion by a bio-mimetic neuro-musculo-skeletal model

Abstract. To emulate the actual neuro-control mechanism of human bipedal locomotion, an anatomically and physiologically based neuro-musculo-skeletal model is developed. The human musculo-skeletal system is constructed as seven rigid links in a sagittal plane, with a total of nine principal muscles. The nervous system consists of an alpha motoneuron and proprioceptors such as a muscle spindle and a Golgi tendon organ for each muscle. At the motoneurons, feedback signals from the proprioceptors are integrated with the signal induced by foot–ground contact and input from the rhythm pattern generator; a muscle activation signal is produced accordingly. Weights of connection in the neural network are optimized using a genetic algorithm, thus maximizing walking distance and minimizing energy consumption. The generated walking pattern is in remarkably good agreement with that of actual human walking, indicating that the locomotory pattern could be generated automatically, according to the musculo-skeletal structures and the connections of the peripheral nervous system, particularly due to the reciprocal innervation in the muscle spindles. Using the proposed model, the flow of sensory-motor information during locomotion is estimated and a possible neuro-control mechanism is discussed.

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.

Human Origins and Environmental Backgrounds.

Hidemi Ishida: 40 Years of Footprints in Japanese Primatology and Paleoanthropology.- Hidemi Ishida: 40 Years of Footprints in Japanese Primatology and Paleoanthropology.- Fossil Hominoids and Paleoenvironments.- Seven Decades of East African Miocene Anthropoid Studies.- Evolution of the Vertebral Column in Miocene Hominoids and Plio-Pleistocene Hominids.- Terrestriality in a Middle Miocene Context: Victoriapithecus from Maboko, Kenya.- Late Cenozoic Mammalian Biostratigraphy And Faunal Change.- The Ages and Geological Backgrounds of Miocene Hominoids Nacholapithecus, Samburupithecus, and Orrorin from Kenya.- Functional Morphology.- Patterns of Vertical Climbing in Primates.- Functional Morphology of the Midcarpal Joint in Knuckle-Walkers and Terrestrial Quadrupeds.- Morphological Adaptation of Rat Femora to Different Mechanical Environments.- A Hallmark of Humankind: The Gluteus Maximus Muscle.- Primates Trained for Bipedal Locomotion as a Model for Studying the Evolution of Bipedal Locomotion.- Locomotor Energetics in Nonhuman Primates.- Computer Simulation of Bipedal Locomotion.- Theoretical Approaches.- Paleoenvironments, Paleoecology, Adaptations, and the Origins of Bipedalism in Hominidae.- Arboreal Origin of Bipedalism.- Neontological Perspectives on East African Middle and Late Miocene Anthropoidea.- The Prehominid Locomotion Reflected: Energetics, Muscles, and Generalized Bipeds.- Evolution of the Social Structure of Hominoids.- Are Human Beings Apes, or are Apes People too?.- Current Thoughts on Terrestrialization in African Apes and the Origin of Human Bipedalism.

Functional craniology and brain evolution: from paleontology to biomedicine.

Anatomical systems are organized through a network of structural and functional relationships among their elements. This network of relationships is the result of evolution, it represents the actual target of selection, and it generates the set of rules orienting and constraining the morphogenetic processes. Understanding the relationship among cranial and cerebral components is necessary to investigate the factors that have influenced and characterized our neuroanatomy, and possible drawbacks associated with the evolution of large brains. The study of the spatial relationships between skull and brain in the human genus has direct relevance in cranial surgery. Geometrical modeling can provide functional perspectives in evolution and brain physiology, like in simulations to investigate metabolic heat production and dissipation in the endocranial form. Analysis of the evolutionary constraints between facial and neural blocks can provide new information on visual impairment. The study of brain form variation in fossil humans can supply a different perspective for interpreting the processes behind neurodegeneration and Alzheimer’s disease. Following these examples, it is apparent that paleontology and biomedicine can exchange relevant information and contribute at the same time to the development of robust evolutionary hypotheses on brain evolution, while offering more comprehensive biological perspectives with regard to the interpretation of pathological processes.

Three-dimensional musculoskeletal kinematics during bipedal locomotion in the Japanese macaque, reconstructed based on an anatomical model-matching method.

Studying the bipedal locomotion of non-human primates is important for clarifying the evolution of habitual bipedalism in the human lineage. However, quantitative descriptions of three-dimensional kinematics of bipedal locomotion in non-human primates are very scarce, due to difficulties associated with measurements. In this study, we performed a kinematic analysis of bipedal locomotion on two highly trained (performing) Japanese macaques walking on a treadmill at different speeds and estimated three-dimensional angular motions of hindlimb and trunk segments, based on a model-based registration method. Our results demonstrated a considerable degree of axial rotation occurring at the trunk and hip joints during bipedal locomotion, suggesting that bipedal locomotion in Japanese macaques is essentially three-dimensional. In addition, ranges of angular motions at the hip and ankle joints were larger and the knee joint was more flexed in the mid-stance phase with increasing speed, indicating that gait kinematics are modulated depending on speed. Furthermore, macaques were confirmed to have actually acquired, at least to some extent, the energy conservation mechanism of walking due to pendular exchange of potential and kinetic energy, but effective utilization of this mutual exchange of energy was found to occur only at comparatively low velocity. Spring-like running mechanics were probably more exploited at higher speed because the duty factor was above 0.5. Fundamental differences in bipedal strategy seem to exist between human and non-human primate bipedal locomotion.

Development of an anatomically based whole-body musculoskeletal model of the Japanese macaque (Macaca fuscata).

We constructed a three-dimensional whole-body musculoskeletal model of the Japanese macaque (Macaca fuscata) based on computed tomography and dissection of a cadaver. The skeleton was modeled as a chain of 20 bone segments connected by joints. Joint centers and rotational axes were estimated by joint morphology based on joint surface approximation using a quadric function. The path of each muscle was defined by a line segment connecting origin to insertion through an intermediary point if necessary. Mass and fascicle length of each were systematically recorded to calculate physiological cross-sectional area to estimate the capacity of each muscle to generate force. Using this anatomically accurate model, muscle moment arms and force vectors generated by individual limb muscles at the foot and hand were calculated to computationally predict muscle functions. Furthermore, three-dimensional whole-body musculoskeletal kinematics of the Japanese macaque was reconstructed from ordinary video sequences based on this model and a model-based matching technique. The results showed that the proposed model can successfully reconstruct and visualize anatomically reasonable, natural musculoskeletal motion of the Japanese macaque during quadrupedal/bipedal locomotion, demonstrating the validity and efficacy of the constructed musculoskeletal model. The present biologically relevant model may serve as a useful tool for comprehensive understanding of the design principles of the musculoskeletal system and the control mechanisms for locomotion in the Japanese macaque and other primates.

Dimensions of forelimb muscles in orangutans and chimpanzees.

Eight forelimbs of three orangutans and four chimpanzees were dissected and the muscle mass, fascicle length and physiological cross‐sectional area (PCSA) of all forelimb muscles were systematically recorded to explore possible interspecies variation in muscle dimensions. Muscle mass and PCSA were divided by the total mass and total PCSA of the entire forelimb muscles for normalization. The results indicate that the mass and PCSA ratios of the monoarticular elbow flexors (M. brachialis and M. brachioradialis) are significantly larger in orangutans. In contrast, the mass ratios of the biarticular muscles in the upper arm (the short head of M. biceps brachii and the long head of M. triceps brachii) are significantly larger in chimpanzees. For the rotator cuff muscles, the force‐generating capacity of M. subscapularis is significantly larger in orangutans, whereas the opposite rotator cuff muscle, M. infraspinatus, is larger in chimpanzees. These differences in forelimb muscle dimensions of the two species may reflect functional specialization for their different positional and locomotor behaviors.

Muscle dimensions in the chimpanzee hand

We dissected the forearms and hands of a female chimpanzee and systematically recorded mass, fiber length, and physiological cross-sectional area (PCSA) of all muscles including those of intrinsic muscles that have not been reported previously. The consistency of our measurements was confirmed by comparison with the published data on chimpanzees. Comparisons of the hand musculature of the measured chimpanzee with corresponding published human data indicated that the chimpanzee has relatively larger forearm flexors but smaller thenar eminence muscles, as observed in previous studies. The interosseous muscles were also confirmed to be relatively larger in the chimpanzee. However, a new finding was that relative PCSA, which reflects a muscle’s capacity to generate force, might have increased slightly in humans as a result of relatively shorter muscle fiber length. This suggests that the human intrinsic muscle architecture is relatively more adapted to dexterous manipulative functions. Shortening of the metacarpals and the intervening interosseous muscles might accordingly be a prerequisite for the evolution of human precision-grip capabilities.

Fetal brain development in chimpanzees versus humans

Summary It is argued that the extraordinary brain enlargement observed in humans is due to not only the human-specific pattern of postnatal brain development, but also to that of prenatal brain development [1,2]. However, the prenatal trajectory of brain development has not been explored in chimpanzees ( Pan troglodytes ), even though they are our closest living relatives. To address this lack of information, we tracked fetal development of the chimpanzee brain from approximately 14 to 34 weeks of gestation (just before birth) in utero using three-dimensional ultrasound imaging. The results were compared with those obtained for the human brain during approximately the same period. We found that the brain volume of chimpanzee fetuses was only half that of human fetuses at 16 weeks of gestation. Moreover, although the growth velocity of brain volume increased until approximately 22 weeks of gestation in both chimpanzees and humans, chimpanzee fetuses did not show the same accelerated increase in brain volume as human fetuses after that time. This suggests that maintenance of fast development of the human brain during intrauterine life has contributed to the remarkable brain enlargement observed in humans.

Muscle architecture of the upper limb in the orangutan

We dissected the left upper limb of a female orangutan and systematically recorded muscle mass, fascicle length, and physiological cross-sectional area (PCSA), in order to quantitatively clarify the unique muscle architecture of the upper limb of the orangutan. Comparisons of the musculature of the dissected orangutan with corresponding published chimpanzee data demonstrated that in the orangutan, the elbow flexors, notably M. brachioradialis, tend to exhibit greater PCSAs. Moreover, the digital II–V flexors in the forearm, such as M. flexor digitorum superficialis and M. flexor digitorum profundus, tend to have smaller PCSA as a result of their relatively longer fascicles. Thus, in the orangutan, the elbow flexors demonstrate a higher potential for force production, whereas the forearm muscles allow a greater range of wrist joint mobility. The differences in the force-generating capacity in the upper limb muscles of the two species might reflect functional specialization of muscle architecture in the upper limb of the orangutan for living in arboreal environments.