An in vitro and finite element study of load redistribution in the midfoot

A good knowledge of midfoot biomechanics is important in understanding the biomechanics of the entire foot,but it has never been investigated thoroughly in the literature.This study carried out in vitro experiments and finite element analysis to investigate the midfoot biomechanics.A foot-ankle finite element model simulating the mid-stance phase of the normal gait was developed and the model validated in in vitro experimental tests.Experiments used seven in vitro samples of fresh human cadavers.The simulation found that the first principal stress peaks of all midfoot bones occurred at the navicular bone and that the tensile force of the spring ligament was greater than that of any other ligament.The experiments showed that the longitudinal strain acting on the medial cuneiform bone was-26.2±10.8μ-strain,and the navicular strain was-240.0±169.1μ-strain along the longitudinal direction and 65.1±25.8μ-strain along the transverse direction.The anatomical position and the spring ligament both result in higher shear stress in the navicular bone.The load from the ankle joint to five branches of the forefoot is redistributed among the cuneiforms and cuboid bones.Further studies on the mechanism of loading redistribution will be helpful in understanding the biomechanics of the entire foot.     

Relationship of strain magnitude to morphological variation in the primate skull

In a comparative study of variation in primate skulls, Wood and Lieberman ([ 2001 ] Am. J. Phys. Anthropol. 116:13–25) proposed that a predictable relationship exists between in vivo bone‐strain magnitudes and the extent of morphological variation in skeletal structures. They hypothesized that regions subject to high strains are prone to enhanced levels of variation. Three questions are posed with respect to the plausibility of this hypothesis. First, does the proposed relationship hold at different levels of analysis (e.g., for more restricted anatomical regions in which large strain gradients are present)? Second, is the biomechanical rationale for the hypothesis sound, given the current understanding of bone biology? Third, is the hypothesis obviated by consideration of the functional matrix concept of skull development, in which osseous growth is posited to be governed by surrounding soft tissues (e.g., muscle and tendon) and developing spaces (e.g., the nasal capsule)? The different perspectives explored by these questions suggest that the validity of the hypothesis, despite having a defensible theoretical rationale, is likely to be context‐specific. A direct role for strain magnitude in conditioning morphological variation is difficult to demonstrate either comparatively or theoretically, and it is unlikely that a single strain threshold or interval can be directly associated with elevated variation in the skeleton. The conceptual framework of the functional matrix (which allows for independent growth among different regions of the skull) conceivably contravenes the premise of a uniform relationship of strain magnitude to morphological variability. Am J Phys Anthropol, 2003. © 2003 Wiley‐Liss, Inc.     

Biomechanical analysis of foot with different foot arch heights: a finite element analysis

Clinically, different foot arch heights are associated with different tissue injuries to the foot. To investigate the possible factors contributing to the difference in foot arch heights, previous studies have mostly measured foot pressure in either low-arched or high-arched feet. However, little information exists on stress variation inside the foot with different arch heights. Therefore, this study aimed to implement the finite element (FE) method to analyse the influence of different foot arches. This study established a 3D foot FE model using software ANSYS 11.0. After validating the FE model, this study created low-arched, high-arched and normal-arched foot FE models. The FE analysis found that both the stress and strain on the plantar fascia and metatarsal were higher in the high-arched foot, whereas the stress and strain on the calcaneous, navicular and cuboid were higher in low-arched foot. Additionally, forefoot pressure was increased with an increase in arch height.     

Brief communication: Cineradiographic analysis of the chimpanzee (Pan troglodytes) talonavicular and calcaneocuboid joints

During terrestrial locomotion, chimpanzees exhibit dorsiflexion of the midfoot between midstance and toe‐off of stance phase, a phenomenon that has been called the “midtarsal break.” This motion is generally absent during human bipedalism, and in chimpanzees is associated with more mobile foot joints than in humans. However, the contribution of individual foot joints to overall foot mobility in chimpanzees is poorly understood, particularly on the medial side of the foot. The talonavicular (TN) and calcaneocuboid (CC) joints have both been suggested to contribute significantly to midfoot mobility and to the midtarsal break in chimpanzees. To evaluate the relative magnitude of motion that can occur at these joints, we tracked skeletal motion of the hindfoot and midfoot during passive plantarflexion and dorsiflexion manipulations using cineradiography. The sagittal plane range of motion was 38 ± 10° at the TN joint and 14 ± 8° at the CC joint. This finding indicates that the TN joint is more mobile than the CC joint during ankle plantarflexion–dorsiflexion. We suggest that the larger range of motion at the TN joint during dorsiflexion is associated with a rotation (inversion–eversion) across the transverse tarsal joint, which may occur in addition to sagittal plane motion. Am J Phys Anthropol 154:604–608, 2014. © 2014 Wiley Periodicals, Inc.     

Trade‐offs in primate grooming reciprocation: testing behavioural flexibility and correlated evolution

Primates may trade altruistic behaviours, such as grooming, either for itself or for different rank‐related benefits, such as tolerance or agonistic support. Ecological conditions are expected to affect competition and thus the steepness of dominance hierarchies. This, in turn, may influence the value of the different currencies that primates exchange. Thus, it can be hypothesized that, as the dominance hierarchy becomes steeper, more grooming is directed up the hierarchy (in exchange for tolerance or agonistic support) and less grooming is exchanged for other grooming. We assembled a large database of within‐group grooming distribution in primates (38 social groups belonging to 16 species and eight genera) and tested these hypotheses both within species (i.e. comparing different groups of the same species) and between species (using comparative methods that control for phylogenetic relatedness). We found within‐species evidence that steeper dominance hierarchies were associated with more grooming being directed up the hierarchy, and that a trade‐off occurred between the tendency to groom up the hierarchy and the degree of grooming reciprocation (although, in some analyses, only a nonsignificant trend was observed). By contrast, phylogenetically controlled comparisons between species did not reveal evidence of correlated evolution between the steepness of the dominance hierarchy, the tendency to direct grooming up the hierarchy, and the degree of grooming reciprocation. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 439–446.     

Middle Eocene primate tarsals from China: implications for haplorhine evolution

We describe tarsal remains of primates recovered from the Middle Eocene (approximately 45 mya) Shanghuang fissures in southern Jiangsu Province, China. These tarsals document the existence of four higher-level taxa of haplorhine primates and at least two adapid species. The meager and poorly preserved adapid material exhibits some similarities to European adapines like Adapis. The haplorhine primates are divided into two major groups: a "prosimian group" consisting of Tarsiidae and an unnamed group that is anatomically similar to Omomyidae; and an "anthropoid group" consisting of Eosimiidae and an unnamed group of protoanthropoids. The anthropoid tarsals are morphologically transitional between omomyids (or primitive haplorhines) and extant telanthropoids, providing the first postcranial evidence for primates which bridge the prosimian-anthropoid gap. All of the haplorhines are extremely small (most are between 50-100 g), and the deposits contain the smallest euprimates ever documented. The uniqueness of this fauna is further highlighted by the fact that no modern primate community contains as many tiny primates as does the fauna from Shanghuang.     

Correlates of variation in deer antler stiffness: age, mineral content, intra-antler location, habitat, and phylogeny

To test correlations between the stiffness of deer antler and aspects of cervid ecology, we measured the stiffness (in bending) of antler from white-tailed deer ( Odocoileus virginianus ) and compared our results to previous measurements from antlers of other cervid species. Stiffness of antler specimens did not correlate significantly with mineral content or the location within the antler from which specimens were taken in O. virginianus. However, antler stiffness in white-tailed deer decreased significantly between two- and three-year-old bucks, matching the time in O. virginianus life history when males shift from sparring with mostly larger individuals to sparring with mostly smaller individuals. Stiffer antlers may enable younger, smaller bucks to have a more effective lever through which their smaller muscular forces can be transmitted during sparring with older, larger bucks. The stiffnesses we measured for white-tailed deer antler are similar to values measured previously from other members of the odocoileine lineage, which are lower than those previously measured from the antlers of deer living in tropical habitats. However, confidence limits for maximum likelihood reconstructions of the ancestral stiffness of deer antler span the range of high stiffnesses found among tropical deer; furthermore, parsimony-based reconstructions of ancestral antler stiffness are equivocal. Thus, the high antler stiffnesses of tropical deer may reflect the retention of an ancestral condition, rather than adaptation to year-round antler use.     

In vivo function of the craniofacial haft: the interorbital "pillar"

The craniofacial haft resists forces generated in the face during feeding, but the importance of these forces for the form of the craniofacial haft remains to be determined. In vivo bone strain data were recorded from the medial orbital wall in an owl monkey (Aotus), rhesus macaques (Macaca mulatta), and a galago (Otolemur) during feeding. These data were used to determine whether: the interorbital region can be modeled as a simple beam under bending or shear; the face is twisting on the brain case during unilateral biting or mastication; the interorbital "pillar" is being axially compressed during incisor loading and both axially compressed and laterally bent during mastication; and the interorbital "pillar" transmits axial compressive forces from the toothrow to the braincase. The strain data reveal that the interorbital region cannot be modeled as a anteroposteriorly oriented beam bent superiorly in the sagittal plane during incision or mastication. The strain orientations recorded in the majority of experiments are concordant with those predicted for a short beam under shear, although the anthropoids displayed evidence of multiple loading regimes in the medial orbital wall. Strain orientation data corroborate the hypothesis that the strepsirrhine face is twisted during mastication. The hypothesis that the interorbital region is a member in a rigid frame subjected to axial compression during mastication receives some support. The hypothesis that the interorbital region is a member in a rigid frame subjected to lateral bending during mastication is supported by the epsilon1/absolute value epsilon2 ratio data but not by the strain orientation data. The timing of peak shear strains in the medial orbital wall of anthropoids does not bear a consistent relationship to the timing of peak shear strain in the mandibular corpus, suggesting that bite force is not the only external force influencing the medial orbital wall. Strain orientation data suggest the existence of two distinct loading regimes, possibly associated with masseter or medial pterygoid contraction. Regardless of the loading regime, all taxa showed low strain magnitudes in the medial orbital wall relative to the anterior root of the zygoma and the mandibular corpus. The strain gradients documented here and elsewhere suggest that, in anthropoids at least, local effects of external forces are more important than a single global loading regime. The low strain magnitudes in the medial orbital wall and in other thin bony plates around the orbit suggest that these structures are not optimally designed for resisting feeding forces. It is hypothesized that their function is to provide rigid support and protection for soft-tissue structures such as the nasal epithelium, the brain, meninges, and the eye and its adnexa. In contrast with the face of Otolemur, which appears to be subjected to a single predominant loading regime, anthropoids may experience different loading regimes in different parts of the face. This implies that the anthropoid and strepsirrhine facial skulls might be optimized for different functions.     

Apparent density of the primate calcaneo‐cuboid joint and its association with locomotor mode,foot posture,and the “midtarsal break”

Primates use a range of locomotor modes during which they incorporate various foot postures. Humans are unique compared with other primates in that humans lack a mobile fore‐ and midfoot. Rigidity in the human foot is often attributed to increased propulsive and stability requirements during bipedalism. Conversely, fore‐ and midfoot mobility in nonhuman primates facilitates locomotion in arboreal settings. Here, we evaluated apparent density (AD) in the subchondral bone of human, ape, and monkey calcanei exhibiting different types of foot loading. We used computed tomography osteoabsorptiometry and maximum intensity projection (MIP) maps to visualize AD in subchondral bone at the cuboid articular surface of calcanei. MIPs represent 3D volumes (of subchondral bone) condensed into 2D images by extracting AD maxima from columns of voxels comprising the volumes. False‐color maps are assigned to MIPs by binning pixels in the 2D images according to brightness values. We compared quantities and distributions of AD pixels in the highest bin to test predictions relating AD patterns to habitual locomotor modes and foot posture categories of humans and several nonhuman primates. Nonhuman primates exhibit dorsally positioned high AD concentrations, where maximum compressive loading between the calcaneus and cuboid likely occurs during “midtarsal break” of support. Humans exhibit less widespread areas of high AD, which could reflect reduced fore‐ and midfoot mobility. Analysis of the internal morphology of the tarsus, such as subchondral bone AD, potentially offers new insights for evaluating primate foot function during locomotion. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

The extensibility of the plantar fascia influences the windlass mechanism during human running

The arch of the human foot is unique among hominins as it is compliant at ground contact but sufficiently stiff to enable push-off. These behaviours are partly facilitated by the ligamentous plantar fascia whose role is central to two mechanisms. The ideal windlass mechanism assumes that the plantar fascia has a nearly constant length to directly couple toe dorsiflexion with a change in arch shape. However, the plantar fascia also stretches and then shortens throughout gait as the arch-spring stores and releases elastic energy. We aimed to understand how the extensible plantar fascia could behave as an ideal windlass when it has been shown to strain throughout gait, potentially compromising the one-to-one coupling between toe arc length and arch length. We measured foot bone motion and plantar fascia elongation using high-speed X-ray during running. We discovered that toe plantarflexion delays plantar fascia stretching at foot strike, which probably modifies the distribution of the load through other arch tissues. Through a pure windlass effect in propulsion, a quasi-isometric plantar fascia''s shortening is delayed to later in stance. The plantar fascia then shortens concurrently to the windlass mechanism, likely enhancing arch recoil at push-off.     

Infectious disease,behavioural flexibility and the evolution of culture in primates

Culturally transmitted traits are observed in a wide array of animal species, yet we understand little about the costs of the behavioural patterns that underlie culture, such as innovation and social learning. We propose that infectious diseases are a significant cost associated with cultural transmission. We investigated two hypotheses that may explain such a connection: that social learning and exploratory behaviours (specifically, innovation and extractive foraging) either compensate for existing infection or increase exposure to infectious agents. We used Bayesian comparative methods, controlling for sampling effort, body mass, group size, geographical range size, terrestriality, latitude and phylogenetic uncertainty. Across 127 primate species, we found a positive association between pathogen richness and rates of innovation, extractive foraging and social learning. This relationship was driven by two independent phenomena: socially contagious diseases were positively associated with rates of social learning, and environmentally transmitted diseases were positively associated with rates of exploration. Because higher pathogen burdens can contribute to morbidity and mortality, we propose that parasitism is a significant cost associated with the behavioural patterns that underpin culture, and that increased pathogen exposure is likely to have played an important role in the evolution of culture in both non-human primates and humans.     

Forward dynamic simulation of bipedal walking in the Japanese macaque: investigation of causal relationships among limb kinematics, speed, and energetics of bipedal locomotion in a nonhuman primate

Japanese macaques that have been trained for monkey performances exhibit a remarkable ability to walk bipedally. In this study, we dynamically reconstructed bipedal walking of the Japanese macaque to investigate causal relationships among limb kinematics, speed, and energetics, with a view to understanding the mechanisms underlying the evolution of human bipedalism. We constructed a two-dimensional macaque musculoskeletal model consisting of nine rigid links and eight principal muscles. To generate locomotion, we used a trajectory-tracking control law, the reference trajectories of which were obtained experimentally. Using this framework, we evaluated the effects of changes in cycle duration and gait kinematics on locomotor efficiency. The energetic cost of locomotion was estimated based on the calculation of mechanical energy generated by muscles. Our results demonstrated that the mass-specific metabolic cost of transport decreased as speed increased in bipedal walking of the Japanese macaque. Furthermore, the cost of transport in bipedal walking was reduced when vertical displacement of the hip joint was virtually modified in the simulation to be more humanlike. Human vertical fluctuations in the body's center of mass actually contributed to energy savings via an inverted pendulum mechanism.     

Comparative rates of lower jaw diversification in cichlid adaptive radiations

The lower jaw (LJ) provides an ideal trophic phenotype to compare rates and patterns of macroevolution among cichlid radiations. Using a novel phylogeny of four genes (ND2, dlx2, mitfb, and s7), we examined the evolutionary relationships among two of the most phylogenetically disparate cichlid radiations: (i) the Central America Heroines; and (ii) the East African Lake Malawi flock. To quantify jaw morphology, we measured two LJ lever systems in approximately 40 species from each lineage. Using geologic calibrations, we generated a chronogram for both groups and examined the rates of jaw evolution in the two radiations. The most rapidly evolving components of the LJ differed between the two radiations. However, the Lake Malawi flock exhibited a much faster rate of evolution in several components of the LJ. This rapid rate of divergence is consistent with natural selection, promoting unparalleled trophic diversification in Lake Malawi cichlids.     

The ear ossicles and the evolution of the primate ear: A biomechanical approach

From a morphometric viewpoint the variability of human and other primate ear ossicles appears to be suitable for the study of taxonomic and phylogenetic distinction among Primates. It may also be of interest to determine whether they are useful to show differences in the perception of sound from the environment and from conspecifics. The energy transmitted through the ossicles is mantained by the action of different leverages. These modify the action of the ossicles from relatively wide, low energy, movements of the hammer to the smaller, high energy, movements of the stirrup. It seems that the pongid type of ossicle leverage combination saves more energy, possibly with a certain loss of subtle information, but this may be more useful in the wild than decoding voice modulation. The human type leverage, being less demultiplied, may produce a major loss of energy but, perhaps, a more precise conservation of sound information useful for speech communication.     

Biomechanical analysis of primate bipedal walking by computer simulation

A computer simulation technique was applied to make clear the mechanical characteristics of primate bipedal walking. A primate body and the walking mechanism were modeled mathematically with a set of dynamic equations. Using a digital computer, the following were calculated from these equations by substituting measured displacements and morphological data of each segment of the primate: the acceleration, joint angle, center of gravity, foot force, joint moment, muscular force, transmitted force at the joint, electric activity of the muscle, generated power by the leg and energy expenditure in walking.The model was evaluated by comparing some of the calculated results with the experimental results such as foot force and electromyographic data, and improved in order to obtain the agreement between them.The level bipedal walking of man, chimpanzee and Japanese monkey and several types of synthesized walking were analyzed from the viewpoint of biomechanics.It is concluded that the bipedal walking of chimpanzee is nearer to that of man than to that of the Japanese monkey because of its propulsive mechanism, but it requires large muscular force for supporting the body weight.     

Kinematics and spatiotemporal parameters of infant‐carrying in olive baboons

In the field of biomechanics of quadrupedal locomotion in primates, infant‐carrying has received little attention. This study presents the first biomechanical study of infant‐carrying in captive female olive baboons (Papio anubis). We test whether females carrying infants conform 1) to the Support Polygon Model (Rollinson and Martin: Symp Zool Soc Lond 48 (1981) 377–427) of gait selection, according to which diagonality should decrease when the infant is carried cranially and increase when the infant is carried dorsally and caudally; 2) to Biewener's (Biewener: Science 245 ( 1989 ) 45–48) theory of limb postures, according to which females should extend their hind limbs more due to infant load, especially in the later stages when the infant is not fully autonomous but relatively heavy. This study focuses on the sagittal kinematics of quadrupedal gaits (joint angles and spatiotemporal parameters) of four females with and without infant loads at the CNRS Primatology Station (France). High‐speed video recordings were made using the technical platform “Motion Analysis of Primates” available in the animals' place of life. Regarding diagonality, our results do not fully conform to those predicted by the Support Polygon Model of gait selection; however, the model cannot be rejected at this stage in experiment. With regard to limb posture, our results do not support Biewener's (Biewener: Science 245 ( 1989 ) 45–48) theory: loaded females do not extend their hind limbs more as predicted; on the contrary, the hind limbs tend to be more flexed when the infant they carry is relatively heavy. Am J Phys Anthropol 155:392–404, 2014. © 2014 Wiley Periodicals, Inc.