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.     

Kinematics of the cercopithecine foot on arboreal and terrestrial substrates with implications for the interpretation of hominid terrestrial adaptations

The stereotyped characterizations of quadrupedal foot postures were tested by examining the kinematics of the cercopithecine foot on arboreal and terrestrial supports. Strictly arboreal species were compared with semi-terrestrial species for Cercopithecus, Cercocebus, Lophocebus, and Papio, in semi-natural or experimental settings. Results indicate that the kinematics of the cercopithecine arboreal quadruped differ in degree from stereotypical expectations for an arboreal quadruped. The relatively extended, adducted limb movements of the cercopithecines and the emphasis on the central digit as the functional axis of the foot suggest convergence with terrestrial mammalian cursors, and differ from the platyrrhine or colobine arboreal quadruped. The characteristics of the quadrupedal terrestrial primate foot contrast with the very unique pattern seen in the hominid foot. These contrasts provide a new perspective from which to interpret the hominid adaptation, in which the functional axis has remained fixed between the first and second digits. This pattern differs from virtually all other terrestrial mammals. The influence of bipedalism on this functional pattern is examined.     

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.     

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.     

Balance of the head in hominid evolution

In primates it is useful to distinguish three basic types of bipedal posture: (1) agonial, with extended hips and knees as in modern humans, (2) monogonial, with flexed hips but extended knees. and (3) digonial, with flexed hips and knees as in pongids. Early hominids retained an ancestral, forwardly inclined posture of the neck and head. Therefore the body posture of australopithednes must have differed from that in modem man, in which the centre of gravity of the head can be aligned with that of the body, other major centra of gravity, and important axes of rotation in a single frontal plane. It is suggested that in australopithednes the gravitational tilt of the head was counterbalanced by bent hips in association with hyperextended knees (monogonial posture). In australopithecines the increase in brain weight would have counteracted an improvement in the balance of the head. After the neck had assumed a more vertical posture as a consequence of shortening of the face, selection for an improved balance system in the bipedal posture favoured an increase in the weight of the postcondylar portion of the head, accentuated by selection for a posterior shift of the superior nuchal line in order to minimise the force of the nuchal muscles. At this stage the evolutionary increase in brain weight may have been largely a by-product of the process towards perfecting the bipedal posture. When the centre of gravity of the head had first become aligned with that of the body, the conditions of balance of the head had become favourable for a dramatic increase of brain size, as a result of selection for greater learning and storage capacity of the brain.     

Placement of the diaphragmatic vertebra in catarrhines: implications for the evolution of dorsostability in hominoids and bipedalism in hominins

A fundamental adaptation to orthograde posture and locomotion amongst living hominoid primates is a numerically reduced lumbar column, which acts to stiffen the lower back and reduce injuries to the intervertebral discs. A related and functionally complementary strategy of spinal stability is a caudal position of the diaphragmatic vertebra relative to the primitive condition found in nonhominoid primates and most other mammals. The diaphragmatic vertebra marks the transition in vertebral articular facet (zygapophysis) orientation, which either resists (prediaphragmatic) or allows (postdiaphragmatic) trunk movement in the sagittal plane (i.e., flexion and extension). Unlike most mammals, which have dorsomobile spines (long lumbar columns and cranially placed diaphragmatic vertebrae) for running and leaping, hominoids possess dorsostable spines (short lumbar columns and caudally placed diaphragmatic vertebrae) adapted to orthogrady and antipronogrady. In contrast to humans and other extant hominoids, all known early hominin partial vertebral columns demonstrate cranial displacement of the diaphragmatic vertebra. To address this difference, variation in diaphragmatic placement is assessed in a large sample of catarrhine primates. I show that while hominoids are characterized by modal common placement of diaphragmatic and last rib-bearing vertebrae in general, interspecific differences in intraspecific patterns of variation exist. In particular, humans and chimpanzees show nearly identical patterns of diaphragmatic placement. A scenario of hominin evolution is proposed in which early hominins evolved cranial displacement from the ancestral hominid condition of common placement to achieve effective lumbar lordosis during the evolution of bipedal locomotion.     

Mediolateral reaction forces and forelimb anatomy in quadrupedal primates: implications for interpreting locomotor behavior in fossil primates

The forelimb joints of terrestrial primate quadrupeds appear better able to resist mediolateral (ML) shear forces than those of arboreal quadrupedal monkeys. These differences in forelimb morphology have been used extensively to infer locomotor behavior in extinct primate quadrupeds. However, the nature of ML substrate reaction forces (SRF) during arboreal and terrestrial quadrupedalism in primates is not known. This study documents ML-SRF magnitude and orientation and forelimb joint angles in six quadrupedal anthropoid species walking across a force platform attached to terrestrial (wooden runway) and arboreal supports (raised horizontal poles). On the ground all subjects applied a lateral force in more than 50% of the steps collected. On horizontal poles, in contrast, all subjects applied a medially directed force to the substrate in more than 75% of the steps collected. In addition, all subjects on arboreal supports combined a lower magnitude peak ML-SRF with a change in the timing of the ML-SRF peak force. As a result, during quadrupedalism on the poles the overall SRF resultant was relatively lower than it was on the runway. Most subjects in this study adduct their humerus while on the poles. The kinetic and kinematic variables combine to minimize the tendency to collapse or translate forelimbs joints in an ML plane in primarily arboreal quadrupedal primates compared to primarily terrestrial quadrupedal ones. These data allow for a more complete understanding of the anatomy of the forelimb in terrestrial vs. arboreal quadrupedal primates. A better understanding of the mechanical basis of morphological differences allows greater confidence in inferences concerning the locomotion of extinct primate quadrupeds.     

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.     

Rates of evolution: Is there a conflict between neo-darwinian evolutionary theory and the fossil record?

Neo-darwinian and population genetics theory assumes that the necessary and sufficient set of conditions for all genetic, therefore evolutionary, change has been identified. Punctuationalists have assumed the opposite and cite the fossil record as evidence for change too rapid to be explained in neo-darwinian theory. Data is given here to provide estimates of the rate of evolution in hominid fossils, in living populations, and of that rate which would qualify as punctuational in the hominid fossil record. Evolution in living populations is orders of magnitude greater than that found in the fossil record and far greater than necessary to create apparently instantaneous saltations in the fossil record. It is suggested that such saltations may not represent more rapid rates of evolution but, rather, the persistence of evolutionary change in a given direction for a longer than normal period.     

Bipedal behavior of olive baboons (Papio anubis) and its relevance to an understanding of the evolution of human bipedalism

The bipedal behavior of a troop of olive baboons (Papio anubis) is described. Bipedalism is relatively rare but nevertheless occurs in a wide variety of situations, although bipedalism during feeding occurs much more frequently than in other situations. The incidence of bipedalism varies between different age-sex classes and between individuals within age-sex classes. This pattern of bipedalism occurred within an overall adaptive response, particularly in feeding behavior, which was similar to that of the gelada baboon (Theropithecus gelada). The data on bipedalism is used together with an existing model of early hominid differentiation based on T. gelada to indicate the types of bipedal behavior which might have occurred in early hominid small object feeders and to suggest how a bipedal pattern of this type might have served as a basis for the action of selection for a more committedly bipedal pattern at later stages of hominid evolution.     

Anterior dental cutting in the Laetolil hominids and the evolution of the bicuspid P3

The younger Laetolil hominids provide evidence of a unique anterior cutting complex with a chisel-like action occurring between the lingual-distal C face and a transverse ridge on the P₃ perpendicular to it, extending between the subequal cusps. An earlier adaptation to more efficient grinding may have resulted in reduced canine projection and the development of the lingual P₃ cusp, raising the ridge between it and the buccal cusp. This development acted to retain the anterior cutting function by shifting it to the top of the premolar.     

Body segments decoupling in sitting: control of body posture from automatic chair adjustments

Background

Individuals who cannot functionally reposition themselves adopt a passive body posture and suffer from physical discomfort in long-term sitting. To regulate body load and to prevent sitting related mobility problems, proper posture control is important. The inability to reposition underlines the importance for seating interventions that control body posture from automatic chair adjustments. We developed an adjustable simulator chair that allows the alignment of the trunk, pelvis and thighs to be controlled independently. This study describes the system for decoupled body segments adjustment and develops a predictive model that computes angular chair configuration for desired body postures.

Methods

Eighteen healthy male subjects participated in this study. The experiment involved a protocol of five trials, each investigating the effect of individual chair segment angle adjustment on body segments rotation. Quasi-static chair adjustments were performed, in which angular chair configuration and body segments orientation were measured using an infrared motion capturing system and an inertia sensor attached on the pelvis.

Results

Linear best-fit equations together with the coefficients of determination were computed. Significant relations have been found between angular chair configuration and body segments orientation leading to an algorithm that predicts chair configuration for desired body posture.

Conclusions

The predictive algorithm seems applicable to compute angular chair configuration for desired body posture when the initial body–chair configuration is known. For clinical application, future experiments must be performed on impaired individuals to validate the algorithm in terms of accuracy.     

Cultural evolution: implications for understanding the human language faculty and its evolution

Human language is unique among the communication systems of the natural world: it is socially learned and, as a consequence of its recursively compositional structure, offers open-ended communicative potential. The structure of this communication system can be explained as a consequence of the evolution of the human biological capacity for language or the cultural evolution of language itself. We argue, supported by a formal model, that an explanatory account that involves some role for cultural evolution has profound implications for our understanding of the biological evolution of the language faculty: under a number of reasonable scenarios, cultural evolution can shield the language faculty from selection, such that strongly constraining language-specific learning biases are unlikely to evolve. We therefore argue that language is best seen as a consequence of cultural evolution in populations with a weak and/or domain-general language faculty.     

Conclusions: implications of the Liang Bua excavations for hominin evolution and biogeography

Excavations at Liang Bua, on the Indonesian island of Flores, have yielded a stratified sequence of stone artifacts and faunal remains spanning the last 95 k.yr., which includes the skeletal remains of two human species, Homo sapiens in the Holocene and Homo floresiensis in the Pleistocene. This paper summarizes and focuses on some of the evidence for Homo floresiensis in context, as presented in this Special Issue edition of the Journal of Human Evolution and elsewhere. Attempts to dismiss the Pleistocene hominins (and the type specimen LB1 in particular) as pathological pygmy humans are not compatible with detailed analyses of the skull, teeth, brain endocast, and postcranium. We initially concluded that H. floresiensis may have evolved by insular dwarfing of a larger-bodied hominin species over 880 k.yr. or more. However, recovery of additional specimens and the numerous primitive morphological traits seen throughout the skeleton suggest instead that it is more likely to be a late representative of a small-bodied lineage that exited Africa before the emergence of Homo erectus sensu lato. Homo floresiensis is clearly not an australopithecine, but does retain many aspects of anatomy (and perhaps behavior) that are probably plesiomorphic for the genus Homo. We also discuss some of the other implications of this tiny, endemic species for early hominin dispersal and evolution (e.g., for the “Out of Africa 1” paradigm and more specifically for colonizing Southeast Asia), and we present options for future research in the region.     

Effects of limb mass distribution on the ontogeny of quadrupedalism in infant baboons (Papio cynocephalus) and implications for the evolution of primate quadrupedalism

Primate quadrupedal kinematics differ from those of other mammals. Several researchers have suggested that primate kinematics are adaptive for safe travel in an arboreal, small-branch niche. This study tests a compatible hypothesis that primate kinematics are related to their limb mass distribution patterns. Primates have more distally concentrated limb mass than most other mammals due to their grasping hands and feet. Experimental studies have shown that increasing distal limb mass by adding weights to the limbs of humans and dogs influences kinematics. Adding weights to distal limb elements increases the natural period of a limb's oscillation, leading to relatively long swing and stride durations. It is therefore possible that primates' distal limb mass is responsible for some of their unique kinematics. This hypothesis was tested using a longitudinal ontogenetic sample of infant baboons (Papio cynocephalus). Because limb mass distribution changes with age in infant primates, this project examined how these changes influence locomotor kinematics within individuals. The baboons in this sample showed a shift in their kinematics as their limb mass distributions changed during ontogeny. When their limb mass was most distally concentrated (at young ages), stride frequencies were relatively low, stride lengths were relatively long, and stance durations were relatively long compared to older ages when limb mass was more proximally concentrated. These results suggest that the evolution of primate quadrupedal kinematics was tied to the evolution of grasping hands and feet.     

New method of three-dimensional analysis of bipedal locomotion for the study of displacements of the body and body-parts centers of mass in man and non-human primates: Evolutionary framework

The current biomechanical interpretation of the chimpanzee's bipedal walking argues that larger lateral and vertical displacements of the body center of mass occur in the chimpanzee's “side-to-side” gait than in the human striding gait. The evolutionary hypothesis underlying this study is the following: during the evolution of human bipedalism one of the necessary changes could have been the progressive reduction of these displacements of the body center of mass. In order to quantitatively test this hypothesis, it is necessary to obtain simultaneously the trajectories of the centers of mass of the whole body and of the different body parts. To solve this problem, a new method of three-dimensional analysis of walking, associated with a volumetric modelling of the body, has been developed based on finite-element modelling. An orthogonal synchrophotographic device yielding four synchronous pictures of the walking subject allows a qualitative analysis of the photographic sequences together with the results of their quantitative analysis. This method was applied to an adult man, a 3-year-old girl and a 9-year-old male chimpanzee. Our results suggest that the trajectory of the body center of mass of the human is distinguished from that of the chimpanzee not by a lower movement amplitude but by the synchronization of the transverse and vertical displacements into two periodic curves in phase with one another. The non-human primate uses its repertoire of arboreal movements in its bipedal terrestrial gait, provisionally referred to as a “rope-walker” gait. We show that the interpretation of a “side-to-side” gait is not applicable to the chimpanzee. We argue that similarly this interpretation and the initial hypothesis presuppose a basic symmetric structure of the gait, in relation to the sagittal plane of progression, similar to the human one. This lateral symmetry of the right and left displacements of the center of gravity, in phase with the right and left single supports of walking, is probably a very derived feature of the human gait. We suggest that low lateral and vertical displacements of the body center of mass are not indicative of a progressive bipedal gait and we discuss the new evolutionary implications of our results. © 1993 Wiley-Liss, Inc.     

Dependence of joint stiffness on the conditions of visual control in upright undisturbed stance in humans

We recorded the sagittal and frontal components of the stabilogram of healthy humans in upright undisturbed stance under five conditions of visual control: (i) open eyes (OE); (ii) closed eyes (CE); (iii) visual inversion (VI); (iv) central vision (CV), and (v) diffused light (DL). Through a low-pass filter of trajectories of the center of pressure of feet (CPF), the vertical projection of the center of gravity (CG) and, consequently, the difference CPF-CG were estimated. The former represents the controlled variable, while the latter is proportional to the horizontal acceleration and assumed to express the resultant joint stiffness (mostly in the ankle joints). The stiffness was characterized through a method based on spectral analysis of the CPF-CG variable and subsequent calculations of the median frequency (MF) and the root mean square (RMS) of the spectra. The median frequencies of the spectra of the CPF-CG variable changed slightly under various visual conditions. At standing on a rigid support, they varied from 0.97 to 0.99 Hz and from 0.93 to 0.97 Hz for the CPF-CG, calculated from the sagittal and frontal components of the stabilogram, respectively. Under conditions of a pliable support, the corresponding frequencies varied within the limits of 0.79–0.83 Hz and 0.74–0.78 Hz. In contrast to the median frequencies, the RMSs demonstrated greater variability depending on different visual conditions. At standing on a rigid support, paired comparisons showed significant differences between the RMSs of the spectra of the CPF-CG variable of the sagittal direction under CE and OE conditions (0.14 ± 0.030 and 0.09 ± 0.020 mm, respectively) and under DL and OE conditions (0.130 ± ± 0.025 and 0.090 ± 0.020 mm, respectively). The RMS of the CPF-CG variable calculated for the frontal stabilogram differed significantly from each other for the VI and OE conditions (0.115 ± 0.020 and 0.075 ± ± 0.015 mm, respectively). In case of standing on a pliable support, a greater variability of visual influences on the CPF-CG variable was found. The RMS for its sagittal motion was the greatest under CE conditions (0.19 ± 0.03 mm); it was significantly greater than the respective values under OE, CV, and DL conditions (0.097 ± ± 0.020, 0.110 ± 0.020, and 0.140 ± 0.030 mm, respectively). The means of RMSs of the spectra of the frontal CPF-CG was also the greatest under CE conditions (0.20 ± 0.03 mm) and the smallest under OE conditions (0.095 ± 0.020 mm). In addition, the value of the RMS fluctuations under CE conditions (0.150 ± 0.025 mm) differed significantly from the respective values under OE conditions (0.095 ± 0.020 mm) and CV conditions (0.110 ± 0.020 mm). Thus, our findings support the statement that the influence of visual conditions on the maintenance of vertical stance is mediated (at least partially) by the mechanisms controlling the ankle joint stiffness. This regulation is mostly manifested in changes of a single parameter, the amplitude of fluctuations of the CPF-CG variable. We also found that the joint stiffness can be modulated by both nonspecific visual influences (which, in particular, reflect the perception of illumination) and specific visual influences, related to information on the position of the body and on its movements with respect to external objects. Neirofiziologiya/Neurophysiology, Vol. 38, No. 2, pp. 157–166, March–April, 2006.