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1.
Carrier DR 《PloS one》2011,6(5):e19630

Background

Many quadrupedal species stand bipedally on their hindlimbs to fight. This posture may provide a performance advantage by allowing the forelimbs to strike an opponent with the range of motion that is intrinsic to high-speed running, jumping, rapid braking and turning; the range of motion over which peak force and power can be produced.

Methodology/Principal Findings

To test the hypothesis that bipedal (i.e., orthograde) posture provides a performance advantage when striking with the forelimbs, I measured the force and energy produced when human subjects struck from “quadrupedal” (i.e., pronograde) and bipedal postures. Downward and upward directed striking energy was measured with a custom designed pendulum transducer. Side and forward strikes were measured with a punching bag instrumented with an accelerometer. When subjects struck downward from a bipedal posture the work was 43.70±12.59% (mean ± S.E.) greater than when they struck from a quadrupedal posture. Similarly, 47.49±17.95% more work was produced when subjects struck upward from a bipedal stance compared to a quadrupedal stance. Importantly, subjects did 229.69±44.19% more work in downward than upward directed strikes. During side and forward strikes the force impulses were 30.12±3.68 and 43.04±9.00% greater from a bipedal posture than a quadrupedal posture, respectively.

Conclusions/Significance

These results indicate that bipedal posture does provide a performance advantage for striking with the forelimbs. The mating systems of great apes are characterized by intense male-male competition in which conflict is resolved through force or the threat of force. Great apes often fight from bipedal posture, striking with both the fore- and hindlimbs. These observations, plus the findings of this study, suggest that sexual selection contributed to the evolution of habitual bipedalism in hominins.  相似文献   

2.
We describe segment angles (trunk, thigh, shank, and foot) and joint angles (hip, knee, and ankle) for the hind limbs of bonobos walking bipedally ("bent-hip bent-knee walking," 17 sequences) and quadrupedally (33 sequences). Data were based on video recordings (50 Hz) of nine subjects in a lateral view, walking at voluntary speed. The major differences between bipedal and quadrupedal walking are found in the trunk, thigh, and hip angles. During bipedal walking, the trunk is approximately 33-41 degrees more erect than during quadrupedal locomotion, although it is considerably more bent forward than in normal human locomotion. Moreover, during bipedal walking, the hip has a smaller range of motion (by 12 degrees ) and is more extended (by 20-35 degrees ) than during quadrupedal walking. In general, angle profiles in bonobos are much more variable than in humans. Intralimb phase relationships of subsequent joint angles show that hip-knee coordination is similar for bipedal and quadrupedal walking, and resembles the human pattern. The coordination between knee and ankle differs much more from the human pattern. Based on joint angles observed throughout stance phase and on the estimation of functional leg length, an efficient inverted pendulum mechanism is not expected in bonobos.  相似文献   

3.
Independent stance is one of the most difficult motor milestones to achieve. Newly standing infants exhibit exaggerated body movements and can only stand for a brief amount of time. Given the difficult nature of bipedal stance, these unstable characteristics are slow to improve. However, we demonstrate that infants can increase their stability when engaged in a standing goal-directed task. Infants'' balance was measured while standing and while standing and holding a visually attractive toy. When holding the toy, infants stood for a longer period of time, exhibited less body sway, and more mature postural dynamics. These results demonstrate that even with limited standing experience, infants can stabilize posture to facilitate performance of a concurrent task.  相似文献   

4.
Hemodynamics and orthodynamics were investigated in quadrupeds (dogs) and in bipeds (humans). The subjects were investigated at rest in supine or lateral posture, in quadrupedal and then in bipedal posture, and during locomotion. Quadrupedalism in humans was with subjects on their hands and knees. Bipedalism in dogs was on hindlimbs with the forelimbs held by a technician. Blood flow in the main arteries of the body (aorta, external and internal carotid, subclavian, and femoral) was measured by sonography. Positional variations between the main bones of the body were determined from X-rays. This study investigated the reallocation of blood supply to different regions of the body when it switches from quadrupedal to bipedal posture and locomotion. Compared with resting posture, the principal findings are 1) cardiac output shows a minimal increase for humans in bipedal stance and a noticeable increase for dogs as well as humans in quadrupedal stance; 2) quadrupedal stance in humans and dogs and bipedal stance in dogs require increased blood supply to the muscles of the neck, back, and limbs, while human bipedal stance requires none of these; 3) cerebral blood flow (internal carotid) in humans did not change as a result of bipedal posture or locomotion, but showed a noticeable drop in quadrupedal posture and an even further drop in quadrupedal locomotion. The conclusion is that erect posture and encephalization produced a noticeable readjustment and reallocation of blood flow among the different regions of the body: This consisted in shifting a large volume of blood supply from the musculature to the human brain.  相似文献   

5.
To question the relation between uni-and bipedal postural skills, 21 subjects were required to stand on a force platform through uni- and bipedal conditions. These two protocols are commonly used paradigms to assess the balance capacities of healthy and disabled patients. The recorded displacements of the center of pressure (CP) were decomposed along mediolateral and anteroposterior axes and assessed through variance positions and parameters obtained from fractional Brownian motion (fBm) modeling to determine the nature and the spatiotemporal organization of the successive controlling mechanisms. The variances underline the relative independence of the two tasks. Nevertheless, as highlighted by the fBm framework, postural correction is initiated for the unipedal stance after shorter time delays and longer covered distances. When compared to bipedal standing, one of the main characteristics of unipedal standing is to induce better-controlled CP trajectories, as deduced from the scaling regimes computed from the fBm modeling. Lastly, the control of the CP trajectories during the shortest time intervals along the anteroposterior axis appears identical for both uni- and bipedal conditions. Unipedal and bipedal standing controls should thus be viewed as two complementary tasks, each providing specific and complementary insights into the postural control organization.  相似文献   

6.
The degree to which non-human primate behavior is lateralized, at either individual or population levels, remains controversial. We investigated the relationship between hand preference and posture during tool use in chimpanzees (Pan troglodytes) during bipedal tool use. We experimentally induced tool use in a supported bipedal posture, an unsupported bipedal posture, and a seated posture. Neither bipedal tool use nor these supported conditions have been previously evaluated in apes. The hypotheses tested were 1) bipedal posture will increase the strength of hand preference, and 2) a bipedal stance, without the use of one hand for support, will elicit a right hand preference. Results supported the first, but not the second hypothesis: bipedalism induced the subjects to become more lateralized, but not in any particular direction. Instead, it appears that subtle pre-existing lateral biases, to either the right or left, were emphasized with increasing postural demands. This result has interesting implications for theories of the evolution of tool use and bipedalism, as the combination of bipedalism and tool use may have helped drive extreme lateralization in modern humans, but cannot alone account for the preponderance of right-handedness.  相似文献   

7.
We investigated the energetic costs of quadrupedal and bipedal walking in two Japanese macaques. The subjects were engaged in traditional bipedal performance for years, and are extremely adept bipeds. The experiment was conducted in an airtight chamber with a gas analyzer. The subjects walked quadrupedally and bipedally at fixed velocities (<5 km/hr) on a treadmill in the chamber for 2.5-6 min. We estimated energy consumption from carbon dioxide (CO2) production. While walking bipedally, energetic expenditure increased by 30% relative to quadrupedalism in one subject, and by 20% in another younger subject. Energetic costs increased linearly with velocity in quadrupedalism and bipedalism, with bipedal/quadrupedal ratios remaining almost constant. Our experiments were relatively short in duration, and thus the observed locomotor costs may include presteady-state high values. However, there was no difference in experimental duration between bipedal and quadrupedal trials. Thus, the issue of steady state cannot cancel the difference in energetic costs. Furthermore, we observed that switching of locomotor mode (quadrupedalism to bipedalism) during a session resulted in a significant increase of CO2 production. Taylor and Rowntree ([1973] Science 179:186-187) noted that the energetic costs for bipedal and quadrupedal walking were the same in chimpanzees and capuchin monkeys. Although the reason for this inconsistency is not clear, species-specific differences should be considered regarding bipedal locomotor energetics among nonhuman primates. Extra costs for bipedalism may not be great in these macaques. Indeed, it is known that suspensory locomotion in Ateles consumes 1.3-1.4 times as much energy relative to quadrupedal progression. This excess ratio surpasses the bipedal/quadrupedal energetic ratios in these macaques.  相似文献   

8.
In this study, we examined the kinematics of bipedal walking in macaque monkeys that have been highly trained to stand and walk bipedally, and compared them to the kinematics of bipedal walking in ordinary macaques. The results revealed that the trained macaques walked with longer and less frequent strides than ordinary subjects. In addition, they appear to have used inverted pendulum mechanics during bipedal walking, which resulted in an efficient exchange of potential and kinetic energy. These gait characteristics resulted from the relatively more extended hindlimb joints of the trained macaques. By contrast, the body of the ordinary macaques translated downward during the single-limb stance phase due to more flexed hindlimb joints. This resulted in almost in-phase fluctuations of potential and kinetic energy, which indicated that energy transformation was less efficient in the ordinary macaques. The findings provide two insights into the early stage of the evolution of human bipedalism. First, the finding that training considerably improved bipedal walking a posteriori may explain why the very first bipeds that might not yet have been morphologically adapted to bipedal walking continued to walk bipedally. The evolutionary transition from quadrupedalism to bipedalism might not be as difficult as has been envisioned. In addition, the finding that macaques, which are phylogenetically distant from humans and in which bipedal walking is unlike human walking, could develop humanlike gait characteristics with training, provides strong support for the commonly held but unproven idea that the characteristics of the human gait are advantageous to human bipedalism.  相似文献   

9.
Natural selection for positional behavior (posture and locomotion) has at least partially driven the evolution of anatomical form and function in the order Primates. Examination of bipedal behaviors associated with daily activity patterns, foraging, and terrestrial habitat use in nonhuman primates, particularly those that adopt bipedal postures and use bipedal locomotion, allows us to refine hypotheses concerning the evolution of bipedalism in humans. This study describes the positional behavior of wild bearded capuchins (Sapajus libidinosus), a species that is known for its use of terrestrial substrates and its habitual use of stones as tools. Here, we test the association of terrestrial substrate use with bipedal posture and locomotion, and the influence of sex (which co‐varies with body mass in adults of this species) on positional behavior and substrate use. Behavior and location of 16 wild adult bearded capuchins from two groups were sampled systematically at 15 s intervals for 2 min periods for 1 year (10,244 samples). Despite their different body masses, adult males (average 3.5 kg) and females (average 2.1 kg) in this study did not differ substantially in their positional behaviors, postures, or use of substrates for particular activities. The monkeys used terrestrial substrates in 27% of samples. Bipedal postures and behaviors, while not a prominent feature of their behavior, occurred in different forms on the two substrates. The monkeys crouched bipedally in trees, but did not use other bipedal postures in trees. While on terrestrial substrates, they also crouched bipedally but occasionally stood upright and moved bipedally with orthograde posture. Bearded capuchin monkeys' behavior supports the suggestion from anatomical analysis that S. libidinosus is morphologically better adapted than its congeners to adopt orthograde postures.  相似文献   

10.
Field observations of bipedal posture and locomotion in wild chimpanzees (Pan troglodytes) can serve as key evidence for reconstructing the likely origins of bipedalism in the last prehominid human ancestor. This paper reports on a sample of bipedal bouts, recorded ad libitum, in wild chimpanzees in Bwindi Impenetrable National Park in southwestern Uganda. The Ruhija community of chimpanzees in Bwindi displays a high rate of bipedal posture. In 246.7 hr of observation from 2001-2003, 179 instances of bipedal posture lasting 5 sec or longer were recorded, for a rate of 0.73 bouts per observation hour. Bipedalism was observed only on arboreal substrates, and was almost all postural, and not locomotor. Bipedalism was part of a complex series of positional behaviors related to feeding, which included two-legged standing, one-legged standing with arm support, and other intermediate postures. Ninety-six percent of bipedal bouts occurred in a foraging context, always as a chimpanzee reached to pluck fruit from tree limbs. Bipedalism was seen in both male and female adults, less frequently among juveniles, and rarely in infants. Both the frequency and duration of bipedal bouts showed a significant positive correlation with estimated substrate diameter. Neither fruit size nor nearest-neighbor association patterns were significantly correlated with the occurrence of bipedalism. Bipedalism is seen frequently in the Bwindi chimpanzee community, in part because of the unusual observer conditions at Bwindi. Most observations of bipedalism were made when the animals were in treetops and the observer at eye-level across narrow ravines. This suggests that wild chimpanzees may engage in bipedal behavior more often than is generally appreciated. Models of the likely evolutionary origins of bipedalism are considered in the light of Bwindi bipedalism data. Bipedalism among Bwindi chimpanzees suggests the origin of bipedal posture in hominids to be related to foraging advantages in fruit trees. It suggests important arboreal advantages in upright posture. The origin of postural bipedalism may have preceded and been causally disconnected from locomotor bipedalism.  相似文献   

11.
Spatio-temporal gait characteristics (step and stride length, stride frequency, duty factor) were determined for the hind-limb cycles of nine bonobos (Pan paniscus) walking quadrupedally and bipedally at a range of speeds. The data were recalculated to dimensionless quantities according to the principle of dynamic similarity. Lower leg length was used as the reference length. Interindividual variability in speed modulation strategy of bonobos appears to be low. Compared to quadrupedal walking, bipedal bonobos use smaller steps to attain a given speed (differences increase with speed), resulting in shorter strides at a higher frequency. In the context of the ("hybrid") dynamic pattern approach to locomotion (Latach, 1998) we argue that, despite these absolute differences, intended walking speed is the basic control variable which elicits both quadrupedal and bipedal walking kinematics in a similar way. Differences in the initial status of the dynamic system may be responsible for the differences in step length between both gaits. Comparison with data deduced from the literature shows that the effects of walking speed on stride length and frequency are similar in bonobos, common chimpanzees, and humans. This suggests that (at least) within extant homininae, spatio-temporal gait characteristics are highly comparable, and this in spite of obvious differences in mass distribution and bipedal posture.  相似文献   

12.
Human postural sway, as measured by fluctuations of the center of pressure (COP) under the feet of a quietly standing individual, can be characterized as a stochastic process. The fluctuation-dissipation theorem (FDT) provides a linear relationship between the fluctuations of a quasi-static, stochastic system to the same system's relaxation to equilibrium following a perturbation. We applied a similar linear relationship, based on the FDT, to the human postural control system to explore whether anterior-posterior (AP) fluctuations of the COP during quiet stance can be used to predict the AP response of the postural control system to a weak posteriorly directed mechanical perturbation (tug or pull at the waist). We tested 10 healthy elderly (mean age of 69yr) and 10 healthy young (mean age of 25yr) adult subjects. We found that this linear relationship was applicable to the postural control system of all 10 young and eight of the 10 elderly adult subjects. These results suggest that it is possible to predict an individual's dynamic response to a mild perturbation using quiet-stance data, regardless of age. The existence of this FDT-based linear relationship with respect to the human postural control system suggests that, for a given individual, the postural control system may use the same control mechanisms during quiet stance and mild-perturbation conditions, regardless of age.  相似文献   

13.
与手偏好相比较,脚偏好被认为是研究大脑半球中语言功能偏侧性调控表达的一种更佳的行为预测指标。当前国际科学界对于人类大脑半球功能不对称性和肢体偏好进化起源的关注,有力地推动了非人灵长类物种肢体偏好行为学研究,其中关于树栖灵长类物种的相关研究,对身体姿势在灵长类肢体偏好表达的理解有十分关键作用。川金丝猴(Rhinopithecus roxellana)是我特有濒危灵长类物种,主要营树栖生活。本研究首次关注秦岭川金丝猴自发性非移动双足姿势(双足叠放)的脚偏好。研究发现在个体水平上每个焦点动物均表现出明显的脚偏好,在群组水平上表现出显著的右脚偏好,脚偏好表达无显著性别差异,其研究结果支持“姿势起源理论”。本文首次呈现野生旧大陆猴物种群组水平脚偏好的研究证据。  相似文献   

14.
This research examined the effects of task (reaching vs. Tool use) and posture (quadrupedal vs. bipedal) on hand preference in tufted capuchins (Cebus apella). Regarding direction of hand preference, we found a significant main effect of posture, as the bipedal stance elicited greater use of the right hand than did the quadrupedal stance, and a significant posture × task interaction, as bipedal reaching elicited greater use of the right hand than did other postural and task conditions. Further, we found a significant main effect of task on strength of hand preference, as tool use elicited more consistent use of one hand over the other than did reaching. Our findings indicate that bipedal reaching facilitates a mild right-hand bias in intensely manipulative primates. We speculate that this moderate bias may have been pushed in the direction of nearly exclusive right-hand preference in most humans with the development of complex tool use. Am. J. Primatol. 44:147–153, 1998. © 1998 Wiley-Liss, Inc. This article is a US Government work and, as such, is in the public domain in the United States of America.  相似文献   

15.
Spontaneously acquired bipedal locomotion of an untrained Japanese monkey (Macaca fuscata) is measured and compared with the elaborated bipedal locomotion of highly trained monkeys to assess the natural ability of a quadrupedal primate to walk bipedally. The subject acquired bipedalism by himself because of the loss of his forearms and hands due to congenital malformation. Two other subjects are performing monkeys that have been extensively trained for bipedal posture and locomotion. We videotaped their bipedal locomotion with two cameras in a lateral view and calculated joint angles (hip, knee, and ankle) and inertial angle of the trunk from the digitized joint positions. The results show that all joints are relatively more flexed in the untrained monkey. Moreover, it is noted that the ankle is less plantar flexed and the knee is more flexed in mid-to-late stance phase in the untrained monkey, suggesting that the trunk is not lifted up to store potential energy. In the trained monkeys, the joints are extended to bring the trunk as high as possible in the stance phase, and then stored potential energy is exchanged for kinetic energy to move forward. The efficient inverted pendulum mechanism seems to be absent in the untrained monkeys locomotion, implying that acquisition of such efficient bipedal locomotion is not a spontaneous ability for a Japanese monkey. Rather, it is probably a special skill that can only be acquired through artificial training for an inherently quadrupedal primate.This revised version was published online in April 2005 with corrections to the cover date of the issue.  相似文献   

16.
Even though specific adjustments of the multi-joint control of posture have been observed when posture is challenged, multi-joint coordination on a seesaw device has never been accurately assessed. The current study was conducted in order to investigate the multi-joint coordination when subjects were standing on either a seesaw device or on a stable surface, with the eyes open or closed. Eighteen healthy active subjects were recruited. A principal component analysis and a Self-Organizing Maps analysis were performed on the joint angles in order to detect and characterize dominant coordination patterns. Intermuscular EMG coherence was analysed in order to assess the neurophysiological mechanisms associated with these coordination patterns. The results illustrated a multi-joint organization of posture on both stable ground and on the seesaw, with a higher variability among the individual postural responses observed when standing on the seesaw. These findings challenge the classical assumption of ankle mechanisms as dominating control on seesaw devices and confirm that inter-joint coordination in postural control is strongly modulated by stance conditions. When standing on the seesaw without vision, a decrease in intermuscular coherence was observed without any impact on the joint coordination patterns, likely due to an increase dependence on proprioceptive information.  相似文献   

17.
Although the identification and characterization of limb load asymmetries during quiet standing has not received much research attention, they may greatly extend our understanding of the upright stance stability control. It seems that the limb load asymmetry factor may serve as a veridical measure of postural stability and thus it can be used for early diagnostic of the age-related decline in balance control. The effects of ageing and of vision on limb load asymmetry (LLA) during quiet stance were studied in 43 healthy subjects (22 elderly, mean age 72.3+/-4.0 yr, and 21 young, mean age 23.9+/-4.8 yr). Postural sway and body weight distribution were recorded while the subject was standing on two adjacent force platforms during two 120 s trials: one trial was performed with the eyes open (EO), while the other trial was with the eyes closed (EC). The results indicate that LLA was greater in the old adults when compared with the young control subjects. The LLA values were correlated with the postural sway magnitudes especially in the anteroposterior direction. Eyes closure which destabilized posture resulted in a significant increase of body weight distribution asymmetry in the elderly but not in the young persons. The limb load difference between EO and EC conditions showed a significantly greater effect of vision on LLA in the elderly compared to the young subjects. The observed differences in the LLA may be attributed to the decline of postural stability control in the elderly. Ageing results in the progressive decline of postural control and usually the nervous system requires more time to complete a balance recovery action. To compensate for such a deficiency, different compensatory strategies are developed. One of them, as evidenced in our study, is preparatory limb unload strategy (a stance asymmetry strategy) which could significantly shorten reaction time in balance recovery.  相似文献   

18.
Recent studies have indicated that chimpanzee bipedality is mechanically inefficient and dynamically unlike that of humans, thus undermining the chimpanzee analogy for mechanical aspects of the early evolution of hominid bipedalism. This paper continues this theme by measuring the forces and stresses engendered by the muscles during bipedal locomotion, for an untrained chimpanzee and for data from chimpanzees which have been encouraged to walk bipedally, presented in the literature. Peak stresses in the triceps surae were lower for the untrained chimpanzee than for the trained subjects because during the late stance phase, when peak ankle moments occur, the centre of pressure of the ground reaction force on the foot of the untrained chimpanzee stayed close to the ankle joint. In contrast, for the trained subjects it moved closer to the toes, as in human bipedalism. Quadriceps and hip extensor stresses are approximately 30% larger for the untrained chimpanzee than for the trained subjects, because the trained chimpanzees walked with a more erect posture. These results may reflect the way in which muscles can develop in response to training, since research on humans has shown that muscle physiological cross-sectional area increases as a result of exercise, resulting in smaller stresses for a given muscle force. During a slow walk, untrained chimpanzees were found to exert far greater muscle stresses than humans do when running at moderate speed, particularly in the muscles that extend the hip, because of the bent-hip, bent-knee posture.  相似文献   

19.
Voluntary arm-raising movement performed during the upright human stance position imposes a perturbation to an already unstable bipedal posture characterised by a high body centre of mass (CoM). Inertial forces due to arm acceleration and displacement of the CoM of the arm which alters the CoM position of the whole body represent the two sources of disequilibrium. A current model of postural control explains equilibrium maintenance through the action of anticipatory postural adjustments (APAs) that would offset any destabilising effect of the voluntary movement. The purpose of this paper was to quantify, using computer simulation, the postural perturbation due to arm raising movement. The model incorporated four links, with shoulder, hip, knee and ankle joints constrained by linear viscoelastic elements. The input of the model was a torque applied at the shoulder joint. The simulation described mechanical consequences of the arm-raising movement for different initial conditions. The variables tested were arm inertia, the presence or not of gravity field, the initial standing position and arm movement direction. Simulations showed that the mechanical effect of arm-raising movement was mainly local, that is to say at the level of trunk and lower limbs and produced a slight forward displacement of the CoM (1.5 mm). Backward arm-raising movement had the same effect on the CoM displacement as the forward arm-raising movement. When the mass of the arm was increased, trunk rotation increased producing a CoM displacement in the opposite direction when compared to arm movement performed without load. Postural disturbance was minimised for an initial standing posture with the CoM vertical projection corresponding to the ankle joint axis of rotation. When the model was reduced to two degrees of freedom (ankle and shoulder joints only) the postural perturbation due to arm-raising movement increased compared to the four-joints model. On the basis of these results the classical assumption that APAs stabilise the CoM is challenged.  相似文献   

20.
The effectivity of the compensatory role of visual biofeedback in cases of decreased stability of upright posture has been analysed. The deterioration of stance was modelled by a subject standing on a soft surface and with additional weight load on the body. The influence of visual biofeedback was positive only for the compensation of decreased stability of upright posture caused by artificially increased body weight of the subject. The compensatory effectivity of visual biofeedback in stabilization of upright posture during stance on a soft surface was practically negligible. The results have shown that effective compensation of the destabilizing effect by visual biofeedback in human upright posture was possible only when the activity and efficiency of efferent-action part of the postural system remained unchanged.  相似文献   

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