Body shape and differences between species

The variation of body shape among prosimians is reviewed. Special emphasis is placed on the selective advantages, that is the mechanical reasons, to which variants of the locomotor apparatus can be traced back. There are differences found in the cheiridia, but at present they cannot be explained in terms of mechanics; there is nearly no knowledge about the mechanical meaning of their diversity. Myological characteristics of taxa can be explained mechanically, but this has not yet been done. Well known are variations of body proportions. These discriminate higher taxa, and are largely coincident with the often-used locomotor categories. In spite of this, there are only few sound arguments about the real biomechanic value of characteristic proportions for a given locomotor mode. What is known on this field, is reviewed. Progress can be made only, if the mechanical conditions, set by postural behavior and locomotion, are understood completely. The subtle distinctions between lower taxonomic units can normally be identified only on the basis of detailed and quantified analyses of movements on one hand, and of biometrics on the other. In the few cases in which such studies have been made, the differences of morphology fit to the mechanical requirements of locomotion which also differ only in quantitative details.     

The metabolic cost of walking in humans, chimpanzees, and early hominins

Bipedalism is a defining feature of the hominin lineage, but the nature and efficiency of early hominin walking remains the focus of much debate. Here, we investigate walking cost in early hominins using experimental data from humans and chimpanzees. We use gait and energetics data from humans, and from chimpanzees walking bipedally and quadrupedally, to test a new model linking locomotor anatomy and posture to walking cost. We then use this model to reconstruct locomotor cost for early, ape-like hominins and for the A.L. 288 Australopithecus afarensis specimen. Results of the model indicate that hind limb length, posture (effective mechanical advantage), and muscle fascicle length contribute nearly equally to differences in walking cost between humans and chimpanzees. Further, relatively small changes in these variables would decrease the cost of bipedalism in an early chimpanzee-like biped below that of quadrupedal apes. Estimates of walking cost in A.L. 288, over a range of hypothetical postures from crouched to fully extended, are below those of quadrupedal apes, but above those of modern humans. These results indicate that walking cost in early hominins was likely similar to or below that of their quadrupedal ape-like forebears, and that by the mid-Pliocene, hominin walking was less costly than that of other apes. This supports the hypothesis that locomotor energy economy was an important evolutionary pressure on hominin bipedalism.     

Effect of footwear on intramuscular EMG activity of plantar flexor muscles in walking

One of the purposes of footwear is to assist locomotion, but some footwear types seem to restrict natural foot motion, which may affect the contribution of ankle plantar flexor muscles to propulsion. This study examined the effects of different footwear conditions on the activity of ankle plantar flexors during walking. Ten healthy habitually shod individuals walked overground in shoes, barefoot and in flip-flops while fine-wire electromyography (EMG) activity was recorded from flexor hallucis longus (FHL), soleus (SOL), and medial and lateral gastrocnemius (MG and LG) muscles. EMG signals were peak-normalised and analysed in the stance phase using Statistical Parametric Mapping (SPM). We found highly individual EMG patterns. Although walking with shoes required higher muscle activity for propulsion than walking barefoot or with flip-flops in most participants, this did not result in statistically significant differences in EMG amplitude between footwear conditions in any muscle (p > 0.05). Time to peak activity showed the lowest coefficient of variation in shod walking (3.5, 7.0, 8.0 and 3.4 for FHL, SOL, MG and LG, respectively). Future studies should clarify the sources and consequences of individual EMG responses to different footwear.     

Holocene footprints in Namibia: The influence of substrate on footprint variability

We report a Holocene human and animal footprint site from the Namib Sand Sea, south of Walvis Bay, Namibia. Using these data, we explore intratrail footprint variability associated with small variations in substrate properties using a “whole foot” analytical technique developed for the studies in human ichnology. We demonstrate high levels of intratrail variability as a result of variations in grain size, depositional moisture content, and the degree of sediment disturbance, all of which determine the bearing capacity of the substrate. The two principal trails were examined, which had consistent stride and step lengths, and as such variations in print typology were primarily controlled by substrate rather than locomotor mechanics. Footprint typology varies with bearing capacity such that firm substrates show limited impressions associated with areas of peak plantar pressure, whereas softer substrates are associated with deep prints with narrow heels and reduced medial longitudinal arches. Substrates of medium bearing capacity give displacement rims and proximal movement of sediment, which obscures the true form of the medial longitudinal arch. A simple conceptual model is offered which summarizes these conclusions and is presented as a basis for further investigation into the control of substrate on footprint typology. The method, model, and results presented here are essential in the interpretation of any sites of greater paleoanthropological significance, such as recently reported from Ileret (1.5 Ma, Kenya; Bennett et al.: Science 323 (2009) 1197–1201). Am J Phys Anthropol 151:265–279, 2013. © 2013 Wiley Periodicals, Inc.     

Evolutionary reversals of limb proportions in early hominids? Evidence from KNM-ER 3735 (Homo habilis)

Upper-to-lower limb proportions of Homo habilis are often said to be more ape-like than those of its reputed ancestor, Australopithecus afarensis. Such proportions would either imply multiple evolutionary reversals or parallel development of a relatively short upper limb in A. afarensis and later Homo. However, assessments of limb proportions are complicated by the fragmentary nature of the two known H. habilis skeletons, OH 62 and KNM-ER 3735. Initially, KNM-ER 3735 was compared to A.L. 288-1 (A. afarensis) using a single modern human and chimpanzee as reference. Here, based on a larger comparative sample, we find that the relative size of the distal humerus, radial head, and shaft of both KNM-ER 3735 and A.L. 288-1 lie within the range of variation of modern humans, whereas their sacra are small as is the case for all early hominids. In addition, their manual phalanges are similar in having a gracile base but robust midshaft. Contrary to earlier studies, the fossils are not differentiable from each other statistically with respect to all features listed above. On the other hand, they differ in robusticity of the scapular spine and relative length of the radial neck. An exact randomization test suggests only a very low probability of finding a similar degree of difference within a single species of extant hominoids. In contrast to the consensus view, we conclude that A.L. 288-1 had a short, human-like forearm, whereas KNM-ER 3735 possessed a distinctly longer forearm and more powerful shoulder girdle. This interpretation fits with earlier conclusions that suggested human-like humerofemoral proportions but chimpanzee-like brachial proportions for Homo habilis. Thus, the scenario of a unidirectional, progressive change in limb proportions within the hominid lineage is not supported by our work.     

The relationship between speed, contact time and peak plantar pressure in terrestrial walking of bonobos

The aim of this paper is twofold. Firstly, we investigate whether contact times, as recorded by pedobarographic systems during quadrupedal and bipedal walking of bonobos, can be used to reliably calculate actual velocities, by applying formulae based on lateral-view video recordings. Secondly, we investigate the effect of speed on peak plantar pressures during bipedal and quadrupedal walking of the bonobo. Data were obtained from 4 individuals from a group of bonobos at the Animal Park Planckendael. From our study, we can conclude that both walking speeds calculated from contact times and lower leg length or simply from recorded contact times are good estimators for walking speed, when direct observation of the latter is impossible. Further, it was found that effects of speed on peak plantar pressures and vertical forces are absent or at least subtle in comparison to a large variation in pressure patterns. In bonobos, the same pressure patterns are used at all walking speeds, and, in consequence, we do not expect major changes in foot function.     

Pitch control and speed limitation during overground deceleration in lemurid primates

An animal's fitness is influenced by the ability to move safely through its environment. Recent models have shown that aspects of body geometry, for example, limb length and center of mass (COM) position, appear to set limits for pitch control in cursorial quadrupeds. Models of pitch control predict that the body shape of these and certain other primates, with short forelimbs and posteriorly positioned COM, should allow them to decelerate rapidly while minimizing the risk of pitching forward. We chose to test these models in two non-cursorial lemurs: Lemur catta, the highly terrestrial ring-tailed lemur, and Eulemur fulvus, the highly arboreal brown lemur. We modeled the effects of changes in limb length and COM position on maximum decelerative potential for both species, as well as collecting data on maximal decelerations across whole strides. In both species, maximum measured decelerations fell below the range of pitch-limited deceleration values predicted by the geometric model, with the ring-tailed lemur approaching its pitch limit more closely. Both lemurs showed decelerative potential equivalent to or higher than horses, the only comparative model currently available. These data reinforce the hypothesis that a relatively simple model of body geometry can predict aspects of maximum performance in animals. In this case, it appears that the body geometry of primates is skewed toward avoiding forward pitch in maximal decelerations.     

The calcaneus of Australopithecus afarensis and its implications for the evolution of bipedality

Calcanei from African apes, modern humans, and Australopithecus afarensis are compared to investigate the anatomical and mechanical changes that occurred in this bone as a result of the transition to terrestrial bipedality. Features analyzed include the cross-sectional area and volume of the calcaneal tuber, the geometry and orientation of the articular surfaces, and the surface topography of the calcaneal corpus. Calcaneal morphology is unequivocal in its partitioning of quadrupedal pongids and bipedal hominids.     

The locomotor anatomy of Australopithecus afarensis

The postcranial skeleton of Australopithecus afarensis from the Hadar Formation, Ethiopia, and the footprints from the Laetoli Beds of northern Tanzania, are analyzed with the goal of determining (1) the extent to which this ancient hominid practiced forms of locomotion other than terrestrial bipedality, and (2) whether or not the terrestrial bipedalism of A. afarensis was notably different from that of modern humans. It is demonstrated that A. afarensis possessed anatomic characteristics that indicate a significant adaptation for movement in the trees. Other structural features point to a mode of terrestrial bipedality that involved less extension at the hip and knee than occurs in modern humans, and only limited transfer of weight onto the medial part of the ball of the foot, but such conclusions remain more tentative than that asserting substantive arboreality. A comparison of the specimens representing smaller individuals, presumably female, to those of larger individuals, presumably male, suggests sexual differences in locomotor behavior linked to marked size dimorphism. The males were probably less arboreal and engaged more frequently in terrestrial bipedalism. In our opinion, A. afarensis from Hadar is very close to what can be called a "missing link." We speculate that earlier representatives of the A. afarensis lineage will present not a combination of arboreal and bipedal traits, but rather the anatomy of a generalized ape.     

The relationship between mitochondrial shape and function and the cytoskeleton

Mitochondria are crucial organelles for life and death of the cell. They are prominent players in energy conversion and integrated signaling pathways including regulation of Ca2+ signals and apoptosis. Their functional versatility is matched by their morphological plasticity and by their high mobility, allowing their transport at specialized cellular sites. This transport occurs by interactions with a variety of cytoskeletal proteins that also have the ability to influence shape and function of the organelle. A growing body of evidence suggests that mitochondria use cytoskeletal proteins as tracks for their movement; in turn, mitochondrial morphology and function is regulated via mostly uncharacterized pathways, by the cytoskeleton.     

The relationship between habitat use and body shape in geckos

Geckos are a highly diverse group of lizards, with more than 1,700 species that exhibit a wide range of behaviors, ecologies, and sizes. However, no study has examined links between habitat use and body shape in pad-bearing geckos. We set out to answer a basic question using a data set of pad-bearing geckos (112 species, 103 pad-bearing, 9 padless, 42 genera): Do geckos that occur in different habitats also differ in body shape? Overall, we found that body shape was surprisingly similar among our sample of pad-bearing species, with the exception of the genus Uroplatus, which was clearly distinct from other geckos due to its depressed body and long limbs. However, the padless geckos differed in body shape from the pad-bearing geckos by having longer arms and legs and less rotund bodies. We found that about half of the pad-bearing species primarily inhabit trees, with the other half, divided approximately equally among rocks, the ground, and mixed habitats. We found no significant links between habitat use and body shape, nor any propensity for larger species to occupy different habitats than smaller species. Padless species tend to inhabit rock and ground substrates. Our results indicate that pad-bearing geckos have a relatively uniform body form, which contrasts with to their diversity in color, size, and behavior. Indeed, our data show that the general gecko body shape is suitable for a wide range of habitats, ranging from arboreal to terrestrial. This pattern is a departure from other ecomorphological studies and suggests that geckos may not easily fit into the mold of adaptive radiation, as suggested by prior studies.     

The relationship between domain duplication and recombination

Protein domains represent the basic evolutionary units that form proteins. Domain duplication and shuffling by recombination are probably the most important forces driving protein evolution and hence the complexity of the proteome. While the duplication of whole genes as well as domain-encoding exons increases the abundance of domains in the proteome, domain shuffling increases versatility, i.e. the number of distinct contexts in which a domain can occur. Here, we describe a comprehensive, genome-wide analysis of the relationship between these two processes. We observe a strong and robust correlation between domain versatility and abundance: domains that occur more often also have many different combination partners. This supports the view that domain recombination occurs in a random way. However, we do not observe all the different combinations that are expected from a simple random recombination scenario, and this is due to frequent duplication of specific domain combinations. When we simulate the evolution of the protein repertoire considering stochastic recombination of domains followed by extensive duplication of the combinations, we approximate the observed data well. Our analyses are consistent with a stochastic process that governs domain recombination and thus protein divergence with respect to domains within a polypeptide chain. At the same time, they support a scenario in which domain combinations are formed only once during the evolution of the protein repertoire, and are then duplicated to various extents. The extent of duplication of different combinations varies widely and, in nature, will depend on selection for the domain combination based on its function. Some of the pair-wise domain combinations that are highly duplicated also recur frequently with other partner domains, and thus represent evolutionary units larger than single protein domains, which we term "supra-domains".     

肠道微生物代谢产物—S-雌马酚与 人类健康关系研究进展

大豆异黄酮是一种年销量很大、市场普及率较高的保健品。具有抗氧化、抗肿瘤等功能,对延缓女性衰老和改善更年期综合征有很好效果。但研究表明,大豆异黄酮的保健功能主要归功于其肠道代谢产物S-雌马酚。S-雌马酚是豆类食品在肠道内经特定微生物代谢后产生的一种高度稳定小分子,其与雌激素的结构和功能高度相似,能与β-雌激素(ER-β)受体结合。具有防治更年期综合征、心血管疾病和多种雌素依赖性癌症的功能。到目前为止,雌马酚的功能及其与人类健康之间的关系还有待进一步研究。本文结合国内外最新研究进展,着重介绍雌马酚的生理功能及其与各类疾病的研究进展,展望雌马酚在各类疾病防治中可能发挥的重要作用。     

Optimal muscle fascicle length and tendon stiffness for maximising gastrocnemius efficiency during human walking and running

Muscles generate force to resist gravitational and inertial forces and/or to undertake work, e.g. on the centre of mass. A trade-off in muscle architecture exists in muscles that do both; the fibres should be as short as possible to minimise activation cost but long enough to maintain an appropriate shortening velocity. Energetic cost is also influenced by tendon compliance which modulates the timecourse of muscle mechanical work. Here we use a Hill-type muscle model of the human medial gastrocnemius to determine the muscle fascicle length and Achilles tendon compliance that maximise efficiency during the stance phase of walking (1.2 m/s) and running (3.2 and 3.9 m/s). A broad range of muscle fascicle lengths (ranging from 45 to 70 mm) and tendon stiffness values (150-500 N/mm) can achieve close to optimal efficiency at each speed of locomotion; however, efficient walking requires shorter muscle fascicles and a more compliant tendon than running. The values that maximise efficiency are within the range measured in normal populations. A non-linear toe-region region of the tendon force-length properties may further influence the optimal values, requiring a stiffer tendon with slightly longer muscle fascicles; however, it does not alter the main results. We conclude that muscle fibre length and tendon compliance combinations may be tuned to maximise efficiency under a given gait condition. Efficiency is maximised when the required volume of muscle is minimised, which may also help reduce limb inertia and basal metabolic costs.     

Structural adaptations of the femur and humerus to arboreal and terrestrial environments in three species of macaque

One reason to measure cross-sectional structural properties of primate long bones is to define mechanically relevant complexes of traits that describe the adaptation of bone to different biomechanical environments. This can be effectively accomplished when congeneric species having different postural and locomotor behaviors are compared. This paper compares the cross-sectional geometry of the femur and humerus in three behaviorally different macaque species as a basis for defining such patterns. Cross-sectional moments of inertia in the standard anatomical planes were calculated at five locations along the diaphyses of the femur and humerus in Macaca fascicularis, M. nemestrina, and M. mulatta. The data suggest that the "barrel-shaped" femur is associated with behaviors for which long limbs and small body size are an asset. This may be associated with, but is not restricted to, leaping behaviors. The data also suggest that structural rigidity of the femur and humerus is greater per unit body weight in primates that spend significant amounts of time in terrestrial environments than in those that are more restricted to climbing in arboreal environments.     

The relationship between whole bone stiffness and strength is age and sex dependent

Accurately estimating whole bone strength is critical for identifying individuals that may benefit from prophylactic treatments aimed at reducing fracture risk. Strength is often estimated from stiffness, but it is not known whether the relationship between stiffness and strength varies with age and sex. Cadaveric proximal femurs (44 Male: 18–78 years; 40 Female: 24–95 years) and radial (36 Male: 18–89 years; 19 Female: 24–95 years) and femoral diaphyses (34 Male: 18–89 years; 19 Female: 24–95 years) were loaded to failure to evaluate how the stiffness-strength relationship varies with age and sex. Strength correlated significantly with stiffness at all sites and for both sexes, as expected. However, females exhibited significantly less strength for the proximal femur (58% difference, p < 0.001). Multivariate regressions revealed that stiffness, age and PYD were significant negative independent predictors of strength for the proximal femur (Age: M: p = 0.005, F: p < 0.001, PYD: M: p = 0.022, F: p = 0.025), radial diaphysis (Age: M = 0.055, PYD: F = 0.024), and femoral diaphysis (Age: M: p = 0.014, F: p = 0.097, PYD: M: p = 0.003, F: p = 0.091). These results indicated that older bones tended to be significantly weaker for a given stiffness than younger bones. These results suggested that human bones exhibit diminishing strength relative to stiffness with aging and with decreasing PYD. Incorporating these age- and sex-specific factors may help to improve the accuracy of strength estimates.     

Effect of age and speed on the step-to-step transition phase during walking

Gait is a powerful measurement tool to evaluate the functional decline throughout ageing. Falls in elderly adults happen mainly during the redirection of the center of mass of the body (CoM) in the transition between steps. In young adults, this step–to–step transition begins before the double contact phase (DC) with a simultaneous forward and upward acceleration of the CoM. We hypothesize that, compared to young adults, elderly adults would exhibit unbalanced contribution of the back leg and the front leg during the transition. We calculated the mean vertical push-off done by the back leg (F_BACK) and the mean impact force on the front leg (F_FRONT) during the transition. Eight young (mean ± SD; age: 24 ± 2 y) and 19 elderly (age: 74 ± 6 y) healthy adults walked on a force-measuring treadmill at five selected speeds ranging from 0.56 to 1.67 m·s⁻¹. Results show that, at mid and high speeds, elderly adults exhibit a smaller F_BACK compared to young adults, possibly linked to the decreased plantar flexion of the back foot. As a consequence, F_FRONT is significantly increased and the transition begins lately in the step, at the beginning of DC. Also, elderly adults show an inability to accelerate the CoM upward and forward simultaneously. Our findings show a different adaptation of the step–to–step transition with speed in elderly adults and identify two potential indicators of gait impairment with age: the F_FRONT/F_BACK contribution and the synchronization between the upward and forward acceleration of the CoM during the transition.