EMG of chimpanzee shoulder muscles during knuckle-walking: problems of terrestrial locomotion in a suspensory adapted primate

By most accounts, the upper limb of the chimpanzee is primarily adapted to suspensory postures and locomotion. In order to determine how the derived morphology of the chimpanzee forelimb has affected the form of quadrupedal locomotion displayed by these animals, electromyographic activity patterns of 10 shoulder muscles during knuckle-walking in two chimpanzee subjects were analysed and compared to data on the opossum and cat taken from the literature. Telemetered electromyography coupled with simultaneous video recording was employed in order to study unfettered locomotion in the chimpanzee subjects.
Chimpanzees are characterized by a quadrupedal gait in which the hind limb overstrides the ipsilateral forelimb. Forelimb position in the plane of abduction/adduction is significantly affected by whether the hind limb passes inside or outside its ipsilateral forelimb. The degree of abduction adduction of the forelimb, in turn, influences many of the muscle activity patterns. That is, some muscles would be more frequently or less frequently active, depending on whether the arm was relatively abducted or adducted during a stride. Thus, there can be no single motor programme that generates the step cycle in chimpanzees.
While there are some parallels between muscle recruitment patterns for chimpanzee, opossum and cat quadrupedalism, the results of this study also indicate that many aspects of muscle use in chimpanzees have been significantly influenced by factors related to increased mobility of the upper limb. Finally, this study has revealed that moving the arm forward during swing phase of knuckle-walking is not a simple product of muscular elTort. and that other mechanisms must be involved. However, it is unclear at present exactly what these mechanisms may be.     

Shoulder movements during quadrupedal locomotion in arboreal primates

The kinematics of scapula and shoulder joint movements were analyzed in three species of arboreal quadrupedal primates using cineradiography. Our findings indicate that scapular movement is highly important for forelimb movement in primates with this ancestral mode of locomotion. Retroversion of the scapula (syn. caudal rotation or extension) during the stance phase contributes more than 40% to the stride length of the forelimb. Lateral forelimb excursions, a general feature for arboreal primates, are based on complex three-dimensional scapular movements guided by the clavicle. Humeral abduction is achieved by scapular abduction and transversal rotation of the scapula about its longitudinal axis, and is therefore strikingly different from humeral abduction in humans. At the same time, the movements of the shoulder joint are limited to flexion and extension only.     

Mechanics of torque generation during quadrupedal arboreal locomotion

Quadrupedal animals moving on arboreal substrates face unique challenges to maintain stability. The torque generated by the limbs around the long axis of a branch during locomotion may clarify how the animals remain stable on arboreal supports. We sought to determine what strategy gray short-tailed opossums (Monodelphis domestica) use to exert torque and avoid toppling. The opossums moved across a branch trackway about half the diameter of their bodies. Part of the trackway was instrumented to measure substrate reaction forces and torque around the long axis of the branch. Kinematic analysis was used to estimate the center of pressure of the manus and pes; from center of pressure and vertical and mediolateral forces, the torque generated by substrate reaction forces versus muscular effort could be determined. Forelimbs generated significantly greater torque than hindlimbs, which is probably explained by the greater weight-bearing role of the forelimbs. Fore- and hindlimbs generated torque in opposite directions because contralateral fore- and hindlimbs typically contacted the branch. Torque generated by muscular effort, however, was often in the same direction in both fore- and hindlimbs. The muscle-generated torque is likely the result of mediolateral movement of the center of mass caused by mediolateral undulation of the torso. These results bear an important implication for the study of arboreal locomotion: center of mass dynamics are at least as important as static positions. M. domestica is a good representative for a primitive mammal, and comparisons with arboreal specialists will shed light on how proficient arboreal locomotion evolved.     

Relation between object properties and EMG during reaching to grasp

In order to stably grasp an object with an artificial hand, a priori knowledge of the object’s properties is a major advantage, especially to ensure subsequent manipulation of the object held by the hand. This is also true for hand prostheses: pre-shaping of the hand while approaching the object, similar to able-bodied, allows the wearer for a much faster and more intuitive way of handling and grasping an object. For hand prostheses, it would be advantageous to obtain this information about object properties from a surface electromyography (sEMG) signal, which is already present and used to control the active prosthetic hand.We describe experiments in which human subjects grasp different objects at different positions while their muscular activity is recorded through eight sEMG electrodes placed on the forearm. Results show that sEMG data, gathered before the hand is in contact with the object, can be used to obtain relevant information on object properties such as size and weight.     

Scapula form and locomotion in chimpanzee evolution

A number of primatologists have followed Coolidge (Am. J. Phys. Anthropol. 18:1–57, 1933) in suggesting that 1) there are significant shape differences in scapula form between pygmy and common chimpanzees, 2) scapulae of P. paniscus resemble those of hylobatids more than do those of P. troglodytes, and 3) therefore pygmy chimpanzees may exhibit a greater component of arm-swinging and other arboreal behaviors than common chimpanzees. In this paper I utilize a comparative analysis of ontogenetic allometries of linear dimensions to determine shape differences in the scapulae of adult pygmy and common chimpanzees and to clarify size-related changes in shape resulting from ontogenetic scaling, i.e., the differential extension of common patterns of growth allometry. Results demonstrate that the scapulae of adult P. paniscus are relatively narrower (in a direction approximately perpendicular to the scapula spine) than those of P. troglodytes, supporting Coolidge's original claim. The allometric analysis further demonstrates, however, that the two chimpanzee species exhibit ontogenetic scaling for all proportions of the scapula examined. Thus, adult pygmy chimpanzees have the scapula proportions observed in small adult and subadult P. troglodytes of comparable scapula size. The implications of this finding for past claims concerning differences in locomotor behavior between the species are discussed. This work lends additional support to previous studies that have demonstrated a high frequency of ontogenetic scaling within the genus Pan and a pedomorphic or juvenilized morphology in the pygmy chimpanzee.     

EMG activity in hyoid muscles during pig suckling

Infant suckling is a complex behavior that includes cycles of rhythmic sucking as well as intermittent swallows. This behavior has three cycle types: 1) suck cycles, when milk is obtained from the teat and moved posteriorly into the valleculae in the oropharynx; 2) suck-swallow cycles, which include both a rhythmic suck and a pharyngeal swallow, where milk is moved out of the valleculae, past the larynx, and into the esophagus; and 3) postswallow suck cycles, immediately following the suck-swallow cycles. Because muscles controlling these behaviors are active in all three types of cycles, we tested the hypothesis that different patterns of electromyographic (EMG) activity in the mylohyoid, hyoglossus, stylohyoid, and thyrohyoid muscles of the pig characterized each cycle type. Anterior mylohyoid EMG activity occurred regularly in every cycle and was used as a cycle marker. Thyrohyoid activity, indicating the pharyngeal swallow, was immediately preceded by increased stylohyoid and hyoglossus activity; it divided the suck-swallow cycle into two phases. Timed from the onset of the suck-swallow cycle, the first phase had a relatively fixed duration while the duration of the second phase, timed from the thyrohyoid, varied directly with cycle duration. In short-duration cycles, the second phase could have a zero duration so that thyrohyoid activity extended into the postswallow cycle. In such cycles, all swallowing activity that occurred after the thyrohyoid EMG and was associated with bolus passage through the pharynx fell into the postswallow cycle. These data suggest that while the activity of some muscles, innervated by trigeminal and cervical plexus nerves, may be time locked to the cycle onset in swallowing, the cycle period itself is not. The postswallow cycle consequently contains variable amounts of pharyngeal swallowing EMG activity. The results exemplify the complexity of the relationship between rhythmic sucking and the swallow.     

The integrated function of muscles and tendons during locomotion

The mechanical roles of tendon and muscle contractile elements during locomotion are often considered independently, but functionally they are tightly integrated. Tendons can enhance muscle performance for a wide range of locomotor activities because muscle-tendon units shorten and lengthen at velocities that would be mechanically unfavorable for muscle fibers functioning alone. During activities that require little net mechanical power output, such as steady-speed running, tendons reduce muscular work by storing and recovering cyclic changes in the mechanical energy of the body. Tendon stretch and recoil not only reduces muscular work, but also allows muscle fibers to operate nearly isometrically, where, due to the force-velocity relation, skeletal muscle fibers develop high forces. Elastic energy storage and recovery in tendons may also provide a key mechanism to enable individual muscles to alter their mechanical function, from isometric force-producers during steady speed running to actively shortening power-producers during high-power activities like acceleration or uphill running. Evidence from studies of muscle contraction and limb dynamics in turkeys suggests that during running accelerations work is transferred directly from muscle to tendon as tendon stretch early in the step is powered by muscle shortening. The energy stored in the tendon is later released to help power the increase in energy of the body. These tendon length changes redistribute muscle power, enabling contractile elements to shorten at relatively constant velocities and power outputs, independent of the pattern of flexion/extension at a joint. Tendon elastic energy storage and recovery extends the functional range of muscles by uncoupling the pattern of muscle fiber shortening from the pattern of movement of the body.     

EMG activity of hip and trunk muscles during deep-water running

The present study used synchronized motion analysis to investigate the activity of hip and trunk muscles during deep-water running (DWR) relative to land walking (LW) and water walking (WW). Nine healthy men performed each exercise at self-determined slow, moderate, and fast paces, and surface electromyography was used to investigate activity of the adductor longus, gluteus maxima, gluteus medius, rectus abdominis, oblique externus abdominis, and erector spinae. The following kinematic parameters were calculated: the duration of one cycle, range of motion (ROM) of the hip joint, and absolute angles of the pelvis and trunk with respect to the vertical axis in the sagittal plane. The percentages of maximal voluntary contraction (%MVC) of each muscle were higher during DWR than during LW and WW. The %MVC of the erector spinae during WW increased concomitant with the pace increment. The hip joint ROMs were larger in DWR than in LW and WW. Forward inclinations of the trunk were apparent for DWR and fast-paced WW. The pelvis was inclined forward in DWR and WW. In conclusion, the higher-level activities during DWR are affected by greater hip joint motion and body inclinations with an unstable floating situation.     

EMG patterns of rat ankle extensors and flexors during treadmill locomotion and swimming

Intramuscular electromyography (EMG) was used to determine and compare the recruitment patterns of the rat soleus (Sol), tibialis anterior (TA), and a deep and a superficial portion of the medial gastrocnemius (MG) during treadmill locomotion at various speeds and inclines and during swimming. Raw EMG signals for 10-20 step or stroke cycles were rectified, averaged, and processed to determine cycle period (EMG onset of one cycle to EMG onset of the next cycle), EMG burst duration, and integrated area of the rectified burst (IEMG). Mean EMG per burst was calculated as IEMG/burst duration. IEMG/min was calculated as IEMG times the number of bursts (cycles) per minute. Cycle period and burst duration of the extensors decreased hyperbolically, while the TA burst duration was unchanged, with increased treadmill speed. With increased treadmill speed, IEMG was decreased in the Sol and unchanged in the MG and TA, whereas IEMG/min decreased in the Sol and increased in the MG and TA. An elevation in treadmill incline resulted in an increase in the activation levels of the MG but not in the Sol or TA. These data indicate that the additional power required at increased speeds and/or inclines of treadmill locomotion is derived from the recruitment of the fast extensors, e.g., the MG. The mean cycle period during swimming was similar to that observed during the fastest treadmill locomotion. EMG burst durations and amplitudes, however, were higher in the TA, relatively similar in the MG, and lower in the Sol during swimming than treadmill locomotion.(ABSTRACT TRUNCATED AT 250 WORDS)     

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.     

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.     

Effects of temperature and perch diameter on arboreal locomotion in the snake Elaphe guttata

Arboreality is widespread in multiple lineages of snakes and these habitats are important for foraging, escaping predators, and thermoregulation for many species. However, very little is known about factors influencing the arboreal locomotor abilities of snakes. Arboreal performance was assessed in a semi-arboreal snake (Elaphe guttata) using an artificial perch apparatus. Locomotor velocity, body posture, and balance was measured during movement on three perch diameters (3, 6, 10-cm) at three temperatures (10, 20, 30 degrees C). Velocities attained by E. guttata on perches are much slower than those of terrestrial lateral undulation and swimming and somewhat slower than concertina velocities recorded in other species across the same experimental temperatures. At higher temperatures, faster speeds were associated with a more elongated posture. At lower temperatures, snakes displayed a more looped body posture, but still fell more often than at higher temperatures. Our results suggest that temperature has a large influence on both balance and movement by snakes on perches. Although there were no differences in velocities resulting from perch diameter, snakes fell more often from thicker perches. This differs from arboreal velocities attained by limbed vertebrates, which decrease with decreasing perch diameter, suggesting that snakes have a size-relative advantage over limbed animals, such as lizards, when traversing a network of narrow branches. Future studies investigating arboreal locomotion among snakes that vary both phylogenetically and morphologically are needed to assess the potential benefits of limblessness in complex, three-dimensional environments.     

Examination of extrinsic foot muscles during running using mfMRI and EMG.

Over-pronation has been cited as a key contributor to many types of running injuries. However, the roles of the extrinsic foot muscles during running have not been adequately identified. The purpose of this study was to examine the muscle functional (mf) MRI and EMG responses to perturbations of the foot by running in varus, neutral and valgus wedged shoes. Ten males ran at 3.6 m/s in specially constructed shoes for 5 min with T2-weighted mfMRI collected before and after each run. The change in T2 from before to after each run characterized the level of metabolic activity in each of muscle. Kinematic and EMG data were also collected while subjects ran on a treadmill. There were no T2 differences across the three shoe conditions. In contrast, there was significantly less EMG activity in the tibialis anterior and soleus while wearing the neutral shoe. Overall, the results did not support the theory that muscle activity would increase as the degree of eversion increased. It also appears that surface EMG was more sensitive to differences between conditions than mfMRI. However, this study illustrated that mfMRI may be a useful tool for quantifying muscle activity in cases where surface EMG is inadequate.     

The jump as a fast mode of locomotion in arboreal and terrestrial biotopes

The jump is always used for locomotion. For its execution in arboreal and terrestrial biotopes the requirements are of somewhat different nature. In an arboreal biotope the jump is characterized by a rapid progression through discontinuous substrates and the ability to take off from a small area and a secure landing on a spot. This requires well coordinated movements in all phases of the jump. On the ground, the jump is less frequent and often used for crossing obstacles or gaps. In primates both variants can be observed. In order to relate the details of locomotor behaviour to a certain environment, the biomechanics of jumping are analyzed in five primate species: The three mainly arboreal prosimian species Galago moholi, the smallest and most specialized leaper of all, Galago garnettii, a medium-sized bushbaby with some capacities for jumping, and Lemur catta also with some abilities to jump. The two simian species, Macaca fuscata and Homo sapiens, are usually terrestrial and have good jumping capacities, although not in terms of quantity. The investigation is based on high-speed motion analyses (100-500 frames/second) and the synchronized records of a force-plate from which all subjects had to jump off. On the basis of the results two kinds of jumping can be distinguished: standing and running jumps. The three prosimian species perform standing jumps. Dorsiflexion of their tails compensates ventrally oriented rotational moments of the trunk during body extension at take-off. The upward arm swing yields an overall increase in take-off velocity without additional muscular force exerted by the legs. The main difference among the species are the high relative forces in the small Galago moholi (up to 13 times body weight) as compared to the larger G. garnettii (8.5 times body weight) and the even larger Lemur catta (4.5 times body weight). In Homo sapiens the standing jump is characterized by an extensive arm swing backward, which is then followed by a forward and upward movement. The velocity at take-off is much smaller if compared to the prosimians. The running jump in Macaca fuscata is always preceded by at least one gallop cycle. The body assumes a ball shape at the beginning of the actual take-off. This is advantageous for rotating the body into a position in which the trunk axis is in line with the direction of movement. The tail of the Japanese macaque is too short to compensate the trunk's lift exerted on the hip region by the extending hindlimbs.(ABSTRACT TRUNCATED AT 400 WORDS)     

EMG and force production of some human shoulder muscles during isometric abduction

Surface EMG was recorded in four subjects on three different occasions from the three parts of the deltoid, the clavicular part of the pectoralis major and from the infraspinatus muscles at different angles of abduction, in the frontal and scapular plane. The integrated EMG was related to the maximum values found for each muscle or muscle part during test contractions (%EMG). Linear relations can be seen for abduction angle vs %EMG. During abduction in the scapular plane the middle and posterior parts of the deltoid muscle showed significantly less activity than in the frontal plane. A simple two dimensional model to calculate the deltoid force out of total external moment at the shoulder is presented. For the middle part of the deltoid an EMG-force relation is presented. The maximal deltoid forces found during test contractions are compared with the absolute muscle force. Also, the length-force relation for the middle part of the deltoid muscle is given between 30° and 90° of abduction.