Locomotion In Chimpanzees Research Articles

Changes in limb bone diaphyseal structure in chimpanzees during development.

This study tests if femoral and humeral cross-sectional geometry (CSG) and cross-sectional properties (CSPs) in an ontogenetic series of wild-caught chimpanzees (Pan troglodytes ssp.) reflect locomotor behavior during development. The goal is to clarify the relationship between limb bone structure and locomotor behavior during ontogeny in Pan. The latex cast method was used to reconstruct cross sections at the midshaft femur and mid-distal humerus. Second moments of area (SMAs) (Ix, Iy, Imax, Imin), which are proportional to bending rigidity about a specified axis, and the polar SMA (J), which is proportional to average bending rigidity, were calculated at section locations. Cross-sectional shape (CSS) was assessed from Ix/Iy and Imax/Imin ratios. Juvenile and adult subsamples were compared. Juveniles and adults have significantly greater femoral J compared to humeral J. Mean interlimb proportions of J are not significantly different between the groups. There is an overall decreasing trend in diaphyseal circularity between the juvenile phase of development and adulthood, although significant differences are only found in the humerus. Juvenile chimpanzee locomotion includes forelimb- and hindlimb-biased behaviors. Juveniles and adults preferentially load their hindlimbs relative to their forelimbs. This may indicate similar locomotor behavior, although other explanations including a diversity of hindlimb-biased locomotor behaviors in juveniles cannot be ruled out. Different ontogenetic trends in forelimb and hindlimb CSS are consistent with limb bone CSG reflecting functional adaptation, albeit the complex nature of bone functional adaptation requires cautious interpretations of skeletal functional morphology from biomechanical analyses.

Great ape thorax and shoulder configuration—An adaptation for arboreality or knuckle-walking?

Great apes exhibit a suite of morphological traits of the shoulder and upper thorax that have traditionally been linked to orthograde arborealism. Recently it has been proposed that these traits are instead adaptations for knuckle-walking, and more broadly, that knuckle-walking itself is an adaptation for shock absorption during terrestriality. Here we test several tenets of these hypotheses using kinematic and kinetic data from chimpanzees and macaques, and electromyographic data of shoulder muscle activity in chimpanzees. We collected 3D kinematic data to quantify motion of the acromion and trunk during quadrupedalism and vertical climbing in chimpanzees as well as ground reaction forces to investigate the presence and magnitude of impact transient forces during terrestrial locomotion in chimpanzees and macaques. We also investigated patterns of recruitment of select forelimb musculature (triceps brachii and serratus anterior) using previously collected data in chimpanzees to determine whether these muscles may function to absorb impact transient forces. We found that the acromion is significantly more elevated in vertical climbing than during knuckle-walking, while dorsoventral ranges and magnitudes of motion were similar between gaits. Ground reaction forces indicate that only a minority of strides in either chimpanzees or macaques have transient forces and, when present, these transient forces as well as loading rates are small. Electromyographic results show that activity of the triceps brachii may facilitate energy absorption while serratus anterior likely functions to support the trunk, as in other primates. Our data suggest there is little to no evidence supporting recent hypotheses that the African ape upper thorax and shoulder configuration is an adaptation for knuckle-walking, or more broadly, that knuckle-walking exists as an adaptation to absorb impact shock during terrestriality. We do however find some evidence that shoulder configuration allows greater scapular elevation in chimpanzees during arboreal behaviors (e.g., vertical climbing).

Chimpanzee ankle and foot joint kinematics: Arboreal versus terrestrial locomotion.

Many aspects of chimpanzee ankle and midfoot joint morphology are believed to reflect adaptations for arboreal locomotion. However, terrestrial travel also constitutes a significant component of chimpanzee locomotion, complicating functional interpretations of chimpanzee and fossil hominin foot morphology. Here we tested hypotheses of foot motion and, in keeping with general assumptions, we predicted that chimpanzees would use greater ankle and midfoot joint ranges of motion during travel on arboreal supports than on the ground. We used a high-speed motion capture system to measure three-dimensional kinematics of the ankle and midfoot joints in two male chimpanzees during three locomotor modes: terrestrial quadrupedalism on a flat runway, arboreal quadrupedalism on a horizontally oriented tree trunk, and climbing on a vertically oriented tree trunk. Chimpanzees used relatively high ankle joint dorsiflexion angles during all three locomotor modes, although dorsiflexion was greatest in arboreal modes. They used higher subtalar joint coronal plane ranges of motion during terrestrial and arboreal quadrupedalism than during climbing, due in part to their use of high eversion angles in the former. Finally, they used high midfoot inversion angles during arboreal locomotor modes, but used similar midfoot sagittal plane kinematics across all locomotor modes. The results indicate that chimpanzees use large ranges of motion at their various ankle and midfoot joints during both terrestrial and arboreal locomotion. Therefore, we argue that chimpanzee foot anatomy enables a versatile locomotor repertoire, and urge caution when using foot joint morphology to reconstruct arboreal behavior in fossil hominins.

Bipedal and quadrupedal locomotion in chimpanzees

Chimpanzees (Pan troglodytes) habitually walk both bipedally and quadrupedally, and have been a common point of reference for understanding the evolution of bipedal locomotion in early ape-like hominins. Here we compare the kinematics, kinetics, and energetics of bipedal and quadrupedal walking and running in a sample of five captive chimpanzees. Kinematics were recorded using sagittal-plane digital high-speed video of treadmill trials. Kinetics were recorded via a forceplate. Metabolic energy cost was measured via steady-state oxygen consumption during treadmill trials. Consistent with previous work on chimpanzees and other hominoids, we found that the spatiotemporal characteristics, joint angles, ground reaction forces, and metabolic cost of bipedal and quadrupedal locomotion are similar in chimpanzees. Notable differences include hip and trunk angles, which reflected a more orthograde trunk posture during bipedalism, and mediolateral ground reaction forces, which were larger during bipedal walking. Stride frequencies were also higher (and step lengths shorter) during bipedal trials. Bipedal and quadrupedal walking among chimpanzees was similar to that reported for bonobos, gibbons, and other primates. The similarity in cost between bipedal and quadrupedal trials suggests that the adoption of bipedal walking would have had no effect on walking costs for early ape-like hominins. However, habitual bipedalism may have favored modifications of the hip to allow a more orthograde posture, and of the hind limb abductor mechanisms to efficiently exert mediolateral ground forces.

Locomotor and postural development of wild chimpanzees

Chimpanzees are our closest living relatives and their positional repertoire likely includes elements shared with our common ancestor. Currently, limitations exist in our ability to correlate locomotor anatomy with behavioral function in the wild. Here we provide a detailed description of developmental changes in chimpanzee locomotion and posture. Fieldwork was conducted on wild chimpanzees at Ngogo, Kibale National Park, Uganda. The large size of the Ngogo chimpanzee community permitted cross-sectional analysis of locomotor and postural changes across many individuals. Chimpanzee positional behavior proceeds developmentally through a number of distinct stages, each characterized by its own loading regime. Infants principally used their upper limbs while moving; the loading environment changed to more hindlimb dominated locomotion as infants aged. Infants displayed more diversity in their forms of positional behavior than members of any other age-sex class, engaging in behaviors not habitually exhibited by adults. While the most common locomotor mode for infants was torso-orthograde suspensory locomotion, a large shift toward quadrupedal locomotion during infancy occurred at three years of age, when rates of this behavior increased. Overall, the most dramatic transition in positional behavior occurred during juvenility (at approximately five years), with the advent of complete independent locomotion. Juveniles decreased the amount of time they spent clinging and in torso-orthograde suspensory locomotion and increased their time spent sitting and walking and running quadrupedally compared with younger individuals. Juvenility marked the age at which quadrupedal walking became the most frequent locomotor behavior, but quadrupedal walking did not encompass the majority of locomotor time until individuals reached adolescence. Relative to all younger individuals, adolescent chimpanzees (10–13 years) experienced a further increase in the amount of time they walked quadrupedally. Locomotor behavior did not reach adult form until adolescence, closer to the time of epiphyseal fusion than previously thought. These findings provide new data to make predictions about how behavioral transitions influence skeletal change.

A three-dimensional musculoskeletal model of the chimpanzee (Pan troglodytes) pelvis and hind limb

Musculoskeletal models have become important tools for studying a range of muscle-driven movements. However, most work has been in modern humans, with few applications in other species. Chimpanzees are facultative bipeds and our closest living relatives, and have provided numerous important insights into our own evolution. A chimpanzee musculoskeletal model would allow integration across a wide range of laboratory-based experimental data, providing new insights into the determinants of their locomotor performance capabilities, as well as the origins and evolution of human bipedalism. Here, we described a detailed three-dimensional (3D) musculoskeletal model of the chimpanzee pelvis and hind limb. The model includes geometric representations of bones and joints, as well as 35 muscle-tendon units that were represented using 44 Hill-type muscle models. Muscle architecture data, such as muscle masses, fascicle lengths and pennation angles, were drawn from literature sources. The model permits calculation of 3D muscle moment arms, muscle-tendon lengths and isometric muscle forces over a wide range of joint positions. Muscle-tendon moment arms predicted by the model were generally in good agreement with tendon-excursion estimates from cadaveric specimens. Sensitivity analyses provided information on the parameters that model predictions are most and least sensitive to, which offers important context for interpreting future results obtained with the model. Comparisons with a similar human musculoskeletal model indicate that chimpanzees are better suited for force production over a larger range of joint positions than humans. This study represents an important step in understanding the integrated function of the neuromusculoskeletal systems in chimpanzee locomotion.

Asymmetries in postural control and locomotion in chimpanzees (Pan troglodytes)

Posture and locomotion are two of the most primitive and basic motor manifestations of an organism's behavior. Although the restrictions they impose on other motor functions are evident, few studies have considered the possibility of asymmetries in these behaviors in human and nonhuman primates, and how they might impact other asymmetries at higher functional levels. The aim of the current study was to explore in a group of 10 chimpanzees at the Madrid Zoo-Aquarium the degree of asymmetry in four behaviors related to locomotion (walking, ascending, descending, and brachiating) and four behaviors associated with posture (sitting, lying, hanging, and changing postures). Few subjects showed individual preferences, but significant trends in the group for some of the behaviors were found, including right-hand use when initiating quadruped walking, and left-hand use when descending and hanging. Some significant correlations also emerged: a negative one between walking and descending, and a positive one between walking and brachiating and between sitting and changing postures. No correlations were found between locomotor and postural modes. Although we cannot make generalizations on the population level at this time, these findings highlight the importance of considering postural and locomotion factors when studying motor asymmetries in primates.

The ontogeny of chimpanzee and pygmy chimpanzee locomotor behavior: a case study of paedomorphism and its behavioral correlates

Pygmy chimpanzees ( Pan paniscus) have been described as “paedomorphic” in comparison with common chimpanzees ( Pan troglodytes) because some of their adult proportions and features are present in immature chimpanzees. However, interpretations of what this morphological paedomorphism means behaviorally have varied widely. This paper documents the changes in positional behavior that occur during the ontogeny of pygmy and common chimpanzees so as to assess whether the adult pygmy chimpanzee is more similar in its positional behavior to immature rather than adult chimpanzees. Results of field studies indicate that both pygmy and common chimpanzees show significant and similar patterns of change in locomotor behavior during ontogeny. With increasing age, in both species, there is an increase in the frequency of quadrupedalism. In early infancy, chimpanzee positional behavior is clearly forelimb dominated, and clinging, climbing and armhanging are most common. Aided bipedalism is the earliest form of frequent hindlimb use. Only later is the infant able to coordinate forelimb and hindlimb use in quadrupedalism. As the frequency of quadrupedalism increases, the frequency of bipedalism decreases. By 2 years of age, common chimpanzee locomotion is primarily quadrupedal. In addition to changes in the type and frequency of locomotor activities performed, there are also ontogenetic changes in the type of substrates used and the locomotor activities performed on them. Results also indicate that adult pygmy chimpanzee arboreal locomotor behavior differs from that of adult common chimpanzees and most closely resembles that of infant chimpanzees. Adult pygmy chimpanzees, like immature common chimpanzees, use more suspensory and quadrupedal behavior than adult common chimpanzees. Shea (1981) has demonstrated that heterochrony in African ape evolution has resulted in substantial morphological differentiation. The results of this study complement Shea's findings by demonstrating that behavioral differences are associated with the morphological patterns.

Development of chimpanzee locomotion on level surfaces

The objective of this study was to investigate kinesiologically the development of the unique characteristics of the level locomotion of the chimpanzee. The data were obtained semi-longitudinally from six chimpanzees eleven weeks through nineteen years of age. The posture, footfall order, phase duration, speed and foot force (including the hand force) in level locomotion were observed by means of foot contact switches, a 16 mm cine-camera or a video tape recorder and a force plate. The speed or the pattern of locomotion was not particularly controlled. The infants moved freely without any attachments on the body. The age change in locomotion is described. The particular characteristics of the infant chimpanzees compared with those of the adults were: 1) long stance phase duration, 2) wide variety in the difference in the cycle duration between forward movement of the limbs one after another, 3) wide variety in phase duration, speed and foot force, and 4) the forelimbs of the infant just started to stand quadrupedally to carry the larger part of the body weight than the hindlimbs. The dominance of the hindlimbs in locomotor and weight-bearing characteristics becomes clearly fixed at about one year of age. The wide variety of the locomotion pattern will be one of the characteristics of the chimpanzees of all age groups. The human acquisition of bipedal walking is discussed in connection with chimpanzee locomotion.

幼年チンパンジー四足ロコモーション発達の運動力学

The quadrupedal locomotion of three infant chimpanzees was studies and compared with that of two semi-adults. The posture, footfall order, phase duration, speed and foot force in locomotion were measured by means of the force plate and a 16-mm cine camera or video tape recorder. The infant animals moved freely without any attachments on the body. In ontogenic walking of chimpanzees, no clear trend was observed to change from the usual terrestrial pattern to the primate pattern. The free locomotion of chimpanzees from infancy to the semi-adult stage varied considerably. The ontogeny of quadrupedal locomotion in chimpanzees, starting with the four-limb standing position, to two years of age was divided into the following stages: 1) From two to three months of age: Beginning with the four-limb standing position and progressing to gliding advancement on the hands and feet. The forelimbs carry a larger part of the body weight than the himdlimbs during static standing posture. "Fist" walking is seen. The hip, knee and toe joints were always flexed. 2) From four months of age: The two diagonal limbs were simultaneously in the air while walking. Footfall order started. Stance phases were long. The step length was short. The foot was not placed near the position where the hand on the same side was placed. The cycle duration was long. The hindlimbs' maximum downward peak showed a significantly larger magnitude than the forelimbs'. 3) From six months of age: Constant and steady walking. The relative speed was slower in this stage than in older ones, if comparison was made between walks of the same cadence. When the equal relative speeds were compared, the accelerating maximum value of the hindlimb was larger in this stage than in older ones. 4) From nine months of age: "Knuckle" walking started. The step length became long. The cycle duration was short. The foot was placed on the outer or inner side of the hand on the same side. Stance phases became short. 5) From one year of age: The hindlimbs carry a clearly larger part of the body weight than the hindlimbs during the static standing posture. The relative speed was fast. Walking as well as galloping was included in the repertoire of quadrupedal locomotion. The hip, knee and the toe joints were extended. 6) Around two years of age: The animals were very active. The "crutch" was included in quadrupedal locomotion.