Dimensions and moment arms of the hind- and forelimb muscles of common chimpanzees (Pan troglodytes).

This paper supplies quantitative data on the hind- and forelimb musculature of common chimpanzees (Pan troglodytes) and calculates maximum joint moments of force as a contribution to a better understanding of the differences between chimpanzee and human locomotion. We dissected three chimpanzees, and recorded muscle mass, fascicle length, and physiological cross-sectional area (PCSA). We also obtained flexion/extension moment arms of the major muscles about the limb joints. We find that in the hindlimb, chimpanzees possess longer fascicles in most muscles but smaller PCSAs than are predicted for humans of equal body mass, suggesting that the adaptive emphasis in chimpanzees is on joint mobility at the expense of tension production. In common chimpanzee bipedalism, both hips and knees are significantly more flexed than in humans, necessitating muscles capable of exerting larger moments at the joints for the same ground force. However, we find that when subject to the same stresses, chimpanzee hindlimb muscles provide far smaller moments at the joints than humans, particularly the quadriceps and plantar flexors. In contrast, all forelimb muscle masses, fascicle lengths, and PCSAs are smaller in humans than in chimpanzees, reflecting the use of the forelimbs in chimpanzee, but not human, locomotion. When subject to the same stresses, chimpanzee forelimb muscles provide larger moments at the joints than humans, presumably because of the demands on the forelimbs during locomotion. These differences in muscle architecture and function help to explain why chimpanzees are restricted in their ability to walk, and particularly to run bipedally.     

Kinematic analysis of bipedal locomotion of a Japanese macaque that lost its forearms due to congenital malformation

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.     

Arboreal bipedalism in wild chimpanzees: implications for the evolution of hominid posture and locomotion

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.     

Muscle mechanical advantage of human walking and running: implications for energy cost.

Muscular forces generated during locomotion depend on an animal's speed, gait, and size and underlie the energy demand to power locomotion. Changes in limb posture affect muscle forces by altering the mechanical advantage of the ground reaction force (R) and therefore the effective mechanical advantage (EMA = r/R, where r is the muscle mechanical advantage) for muscle force production. We used inverse dynamics based on force plate and kinematic recordings of humans as they walked and ran at steady speeds to examine how changes in muscle EMA affect muscle force-generating requirements at these gaits. We found a 68% decrease in knee extensor EMA when humans changed gait from a walk to a run compared with an 18% increase in hip extensor EMA and a 23% increase in ankle extensor EMA. Whereas the knee joint was extended (154-176 degrees) during much of the support phase of walking, its flexed position (134-164 degrees) during running resulted in a 5.2-fold increase in quadriceps impulse (time-integrated force during stance) needed to support body weight on the ground. This increase was associated with a 4.9-fold increase in the ground reaction force moment about the knee. In contrast, extensor impulse decreased 37% (P < 0.05) at the hip and did not change at the ankle when subjects switched from a walk to a run. We conclude that the decrease in limb mechanical advantage (mean limb extensor EMA) and increase in knee extensor impulse during running likely contribute to the higher metabolic cost of transport in running than in walking. The low mechanical advantage in running humans may also explain previous observations of a greater metabolic cost of transport for running humans compared with trotting and galloping quadrupeds of similar size.     

Back muscle function during bipedal walking in chimpanzee and gibbon: implications for the evolution of human locomotion

The evolution of erect posture and locomotion continues to be a major focus of interest among paleoanthropologists and functional morphologists. To date, virtually all of our knowledge about the functional role of the back muscles in the evolution of bipedalism is based on human experimental data. In order to broaden our evolutionary perspective on the vertebral region, we have undertaken an electromyographic (EMG) analysis of three deep back muscles (multifidus, longissimus thoracis, iliocostalis lumborum) in the chimpanzee (Pan troglodytes) and gibbon (Hylobates lar) during bipedal walking. The recruitment patterns of these three muscles seen in the chimpanzee closely parallel those observed in the gibbon. The activity patterns of multifidus and longissimus are more similar to each other than either is to iliocostalis. Iliocostalis recruitment is clearly related to contact by the contralateral limb during bipedal walking in both species. It is suggested that in both the chimpanzee and gibbon, multifidus controls trunk movement primarily in the sagittal plane, iliocostalis responds to and adjusts movement in the frontal plane, while longissimus contributes to both of these functions. In many respects, the activity patterns shared by the chimpanzee and gibbon are quite consistent with recent human experimental data. This suggests a basic similarity in the mechanical constraints placed on the back during bipedalism among these three hominoids. Thus, the acquisition of habitual bipedalism in humans probably involved not so much a major change in back muscle action or function, but rather an improvement in the mechanical advantages and architecture of these muscles.     

Center of mass mechanics of chimpanzee bipedal walking

Center of mass (CoM) oscillations were documented for 81 bipedal walking strides of three chimpanzees. Full‐stride ground reaction forces were recorded as well as kinematic data to synchronize force to gait events and to determine speed. Despite being a bent‐hip, bent‐knee (BHBK) gait, chimpanzee walking uses pendulum‐like motion with vertical oscillations of the CoM that are similar in pattern and relative magnitude to those of humans. Maximum height is achieved during single support and minimum height during double support. The mediolateral oscillations of the CoM are more pronounced relative to stature than in human walking when compared at the same Froude speed. Despite the pendular nature of chimpanzee bipedalism, energy recoveries from exchanges of kinetic and potential energies are low on average and highly variable. This variability is probably related to the poor phasic coordination of energy fluctuations in these facultatively bipedal animals. The work on the CoM per unit mass and distance (mechanical cost of transport) is higher than that in humans, but lower than that in bipedally walking monkeys and gibbons. The pronounced side sway is not passive, but constitutes 10% of the total work of lifting and accelerating the CoM. CoM oscillations of bipedally walking chimpanzees are distinctly different from those of BHBK gait of humans with a flat trajectory, but this is often described as “chimpanzee‐like” walking. Human BHBK gait is a poor model for chimpanzee bipedal walking and offers limited insights for reconstructing early hominin gait evolution. Am J Phys Anthropol 156:422–433, 2015. © 2014 Wiley Periodicals, Inc.     

Muscle architecture of the common chimpanzee (Pan troglodytes): perspectives for investigating chimpanzee behavior

Thorpe et al. (Am J Phys Anthropol 110:179–199, 1999) quantified chimpanzee (Pan troglodytes) muscle architecture and joint moment arms to determine whether they functionally compensated for structural differences between chimpanzees and humans. They observed enough distinction to conclude that musculoskeletal properties were not compensatory and suggested that chimpanzees and humans do not exhibit dynamically similar movements. These investigators based their assessment on unilateral limb musculatures from three male chimpanzees, of which they called one non-adult representative. Factors such as age, sex, and behavioral lateralization may be responsible for variation in chimpanzee muscle architecture, but this is presently unknown. While the full extent of variation in chimpanzee muscle architecture due to such factors cannot be evaluated with data presently available, the present study expands the chimpanzee dataset and provides a preliminary glimpse of the potential relevance of these factors. Thirty-seven forelimb and 36 hind limb muscles were assessed in two chimpanzee cadavers: one unilaterally (right limbs), and one bilaterally. Mass, fiber length, and physiological cross-sectional area (PCSA) are reported for individual muscles and muscle groups. The musculature of an adult female is more similar in architectural patterns to a young male chimpanzee than to humans, particularly when comparing muscle groups. Age- and sex-related intraspecific differences do not obscure chimpanzee-human interspecific differences. Side asymmetry in one chimpanzee, despite consistent forelimb directional asymmetry, also does not exceed the magnitude of chimpanzee-human differences. Left forelimb muscles, on average, usually had higher masses and longer fiber lengths than right, while right forelimb muscles, on average, usually had greater PCSAs than left. Most muscle groups from the left forelimb exhibited greater masses than right groups, but group asymmetry was significant only for the manual digital muscles. The hind limb exhibited less asymmetry than the forelimb in most comparisons. Examination of additional chimpanzees would clarify the full range of inter- and intra-individual variation.     

Twitch contractile properties of plantar flexor muscles in power and endurance trained athletes

This study compared twitch contractile properties of plantar flexor muscles among three groups of 12 subjects each: endurance and power trained athletes and untrained subjects. The posterior tibial nerve was stimulated by supramaximal square wave pulses of 1-ms duration. Power trained athletes had higher twitch maximal force, maximal rates of force development and relaxation and also maximal voluntary contraction (MVC) force. The trained subjects had a smaller twitch maximal force: MVC force ratio and shorter twitch contraction and half-relaxation times than the untrained subjects with no significant differences between the two groups. Thus, the short time for evoked twitches in the athletes compared to the untrained subjects would seem unrelated to the type of training. It is concluded that power training induces a more evident increase of muscle force-generating capacity and speed of contraction and relaxation than endurance training. Accepted: 24 April 1999     

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.     

Chimpanzee thumb muscle cross sections, moment arms and potential torques, and comparisons with humans.

This study investigates the morphological basis of differences between humans and chimpanzees in the kinematical and dynamical parameters of the musculature of the thumb. It is partly intended to test an hypothesis that human thumb muscles can exert significantly greater torques, due to larger muscle cross-sectional areas or to longer tendon moment arms or to both. We focus on the estimation of the potentials of thumb muscles to exert torques about joint axes in a sample of eight chimpanzee cadaver hands. The potential torque of a muscle is estimated by taking the product of a muscle's physiological cross-sectional area (an estimator of force) with its dynamical moment arm (derived from the slope of tendon excursion versus joint angular displacement, obtained during passive movements of cadaver thumb joints). Comparison of our results with similar data obtained for humans at the same Mayo Clinic laboratory shows significant differences between humans and chimpanzees in potential torque of most thumb muscles, those of humans generally exhibiting larger values. The primary reason for the larger torques in humans is that their average moment arms are significantly longer, permitting greater torque for a given muscle size. An additional finding is that chimpanzees and humans differ in the direction of secondary thumb metacarpal movements elicited by contraction of some muscles, as shown by differences in moment arm signs for a given movement in the same muscle. The differences appear to be related to differences in the musculo-skeletal structures of the trapeziometacarpal joint.     

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 muscles 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.     

Bipedal tool use strengthens chimpanzee hand preferences

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.     

Muscle force production during bent-knee,bent-hip walking in humans

Researchers have long debated the locomotor posture used by the earliest bipeds. While many agree that by 3–4 Ma (millions of years ago), hominins walked with an extended-limb human style of bipedalism, researchers are still divided over whether the earliest bipeds walked like modern humans, or walked with a more bent-knee, bent-hip (BKBH) ape-like form of locomotion. Since more flexed postures are associated with higher energy costs, reconstructing early bipedal mechanics has implications for the selection pressures that led to upright walking. The purpose of this study is to determine how modern human anatomy functions in BKBH walking to clarify the links between morphology and energy costs in different mechanical regimes. Using inverse dynamics, we calculated muscle force production at the major limb joints in humans walking in two modes, both with extended limbs and BKBH. We found that in BKBH walking, humans must produce large muscle forces at the knee to support body weight, leading to higher estimated energy costs. However, muscle forces at the hip remained similar in BKBH and extended limb walking, suggesting that anatomical adaptations for hip extension in humans do not necessarily diminish the effective mechanical advantage at the hip in more flexed postures. We conclude that the key adaptations for economical walking, regardless of joint posture, seem to center on maintaining low muscle forces at the hip, primarily by keeping low external moments at the hip. We explore the implications of these results for interpreting locomotor energetics in early hominins, including australopithecines and Ardipithecus ramidus.     

Electromyography of back muscles during quadrupedal and bipedal walking in primates

Despite the extensive electromyographic research that has addressed limb muscle function during primate quadrupedalism, the role of the back muscles in this locomotor behavior has remained undocumented. We report here the results of an electromyographic (EMG) analysis of three intrinsic back muscles (multifidus, longissimus, and iliocostalis) in the baboon (Papio anubis), chimpanzee (Pan troglodytes), and orangutan (Pongo pygmaeus) during quadrupedal walking. The recruitment patterns of these three back muscles are compared to those reported for the same muscles during nonprimate quadrupedalism. In addition, the function of the back muscles during quadrupedalism and bipedalism in the two hominoids is compared. Results indicate that the back muscles restrict trunk movements during quadrupedalism by contracting with the touchdown of one or both feet, with more consistent activity associated with touchdown of the contralateral foot. Moreover, despite reported differences in their gait preferences and forelimb muscle EMG patterns, primates and nonprimate mammals recruit their back muscles in an essentially similar fashion during quadrupedal walking. These quadrupedal EMG patterns also resemble those reported for chimpanzees, gibbons and humans (but not orangutans) walking bipedally. The fundamental similarity in back muscle function across species and locomotor behaviors is consistent with other data pointing to conservatism in the evolution of the neural control of tetrapod limb movement, but does not preclude the suggestion (based on forelimb muscle EMG and spinal lesion studies) that some aspects of primate neural circuitry are unique. © 1994 Wiley-Liss, Inc.     

Changes in sEMG parameters among trunk and thigh muscles during a fatiguing bilateral isometric multi-joint task in trained and untrained subjects

This study aimed to explore changes in the electrical activity distribution among synergist muscles involved in the maintenance of this bilateral multi-joint task. It also tested relations between changes in surface electromyographic (sEMG) parameters with endurance time. Eighteen subjects, trained and untrained in hiking, performed a submaximal (50% of maximal contraction) isometric hiking test until exhaustion. The electrical activity of main superficial muscles implicated in this posture was recorded bilaterally. Trained subjects sustained the hiking position for 315 ± 82 s, versus 225 ± 68 s for untrained subjects. Patterns of electrical activity and mean power frequency (MPF) were different between populations. MPF shift in abdominal muscles was higher than in other synergists for both groups. Although typical changes in sEMG parameters were observed, few relations with endurance time were found, and for untrained subjects only. Changes in the relative contribution among synergists were observed, mainly for trained subjects. It is hypothesized that the task (a complex multi-joint posture involving numerous joints and muscles) may allow some variability in the contribution of synergist muscles during fatigue especially for the trained group. This probably explains the absence of relationship between endurance time and sEMG changes for trained subjects.     

The Laetoli footprints and early hominin locomotor kinematics

A critical question in human evolution is whether the earliest bipeds walked with a bent-hip, bent-knee gait or on more extended hindlimbs. The differences between these gaits are not trivial, because the adoption of either has important implications for the evolution of bipedalism. In this study, we re-examined the Laetoli footprints to determine whether they can provide information on the locomotor posture of early hominins. Previous researchers have suggested that the stride lengths of Laetoli hominins fall within the range of modern human stride lengths and therefore, Laetoli hominins walked with modern-human-like kinematics. Using a dynamic-similarity analysis, we compared Laetoli hominin stride lengths with those of both modern humans and chimpanzees. Our results indicate that Laetoli hominins could have used either a bent-hip, bent-knee gait, similar to a chimpanzee, or an extended-hindlimb gait, similar to a human. In fact, our data suggest that the Laetoli hominins could have walked near their preferred speeds using either limb posture. This result contrasts with most previous studies, which suggest relatively slow walking speeds for these early bipeds. Despite the many attempts to discern limb-joint kinematics from Laetoli stride lengths, our study concludes that stride lengths alone do not resolve the debate over early hominin locomotor postures.     

Development of the visual preference of chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) for photographs of primates: effect of social experience

In a study by Tanaka (2003) five captive chimpanzees preferred photographs of humans to those of chimpanzees. All the subjects were raised by humans and lived in captivity for many years. This suggests their preference might have developed through social experience. In this study examined this hypothesis by using three young chimpanzees raised by their mothers in a captive chimpanzee community. The young chimpanzees were tested four times before six years of age. I also tested eight adult chimpanzees that had been in captivity for more than 20 years. Each subject was presented with digitized color photographs of different species of primates on a touch-sensitive screen. The subjects received a food reward when they touched a photograph, irrespective of which photograph they touched. All the adult chimpanzees touched photographs of humans more frequently than those of any other species of primate. Two of the young chimpanzees showed no species preference before reaching 5 years of age, when they started to show preference for humans. The remaining young chimpanzee consistently preferred chimpanzees. These results suggest that development of visual preference of chimpanzees is affected by social experience during infancy.     

A model-based study of passive joint properties on muscle effort during static stance

This study examined the impact of lower extremity joint stiffnesses and simulated joint contractures on the muscle effort required to maintain static standing postures after a spinal cord injury (SCI). Static inverse computer simulations were performed with a three-dimensional 15 degree of freedom musculoskeletal model placed in 1600 different standing postures. The required lower extremity muscle forces were calculated through an optimization routine that minimized the sum of the muscle stresses squared, which was used as an index of the muscle effort required for each standing posture. Joint stiffnesses were increased and decreased by 100 percent of their nominal values, and contractures were simulated to determine their effects on the muscle effort for each posture. Nominal muscle and passive properties for an individual with a SCI determined the baseline muscle effort for comparisons. Stiffness changes for the ankle plantar flexion/dorsiflexion, hip flexion/extension, and hip abduction/adduction directions had the largest effect on reducing muscle effort by more than 5 percent, while changes in ankle inversion/eversion and knee flexion/extension had the least effect. For erect standing, muscle effort was reduced by more than 5 percent when stiffness was decreased at the ankle plantar flexion/dorsiflexion joint or hip flexion/extension joint. With simulated joint contractures, the postural workspace area decreased and muscle effort was not reduced by more than 5 percent for any posture. Using this knowledge, methods can be developed through the use of orthoses, physical therapy, surgery or other means to appropriately augment or diminish these passive moments during standing with a neuroprosthesis.     

The evolutionary history of human and chimpanzee Y-chromosome gene loss

Recent studies have suggested that gene gain and loss may contribute significantly to the divergence between humans and chimpanzees. Initial comparisons of the human and chimpanzee Y-chromosomes indicate that chimpanzees have a disproportionate loss of Y-chromosome genes, which may have implications for the adaptive evolution of sex-specific as well as reproductive traits, especially because one of the genes lost in chimpanzees is critically involved in spermatogenesis in humans. Here we have characterized Y-chromosome sequences in gorilla, bonobo, and several chimpanzee subspecies for 7 chimpanzee gene-disruptive mutations. Our analyses show that 6 of these gene-disruptive mutations predate chimpanzee-bonobo divergence at approximately 1.8 MYA, which indicates significant Y-chromosome change in the chimpanzee lineage relatively early in the evolutionary divergence of humans and chimpanzees.     

Crouched postures reduce the capacity of muscles to extend the hip and knee during the single-limb stance phase of gait

Many children with cerebral palsy walk in a crouch gait that progressively worsens over time, decreasing walking efficiency and leading to joint degeneration. This study examined the effect of crouched postures on the capacity of muscles to extend the hip and knee joints and the joint flexions induced by gravity during the single-limb stance phase of gait. We first characterized representative mild, moderate, and severe crouch gait kinematics based on a large group of subjects with cerebral palsy (N=316). We then used a three-dimensional model of the musculoskeletal system and its associated equations of motion to determine the effect of these crouched gait postures on (1) the capacity of individual muscles to extend the hip and knee joints, which we defined as the angular accelerations of the joints, towards extension, that resulted from applying a 1N muscle force to the model, and (2) the angular acceleration of the joints induced by gravity. Our analysis showed that the capacities of almost all the major hip and knee extensors were markedly reduced in a crouched gait posture, with the exception of the hamstrings muscle group, whose extension capacity was maintained in a crouched posture. Crouch gait also increased the flexion accelerations induced by gravity at the hip and knee throughout single-limb stance. These findings help explain the increased energy requirements and progressive nature of crouch gait in patients with cerebral palsy.