Effects of growth and speed on hindlimb joint angular displacement patterns in vervet monkeys (Cercopithecus aethiops)

Hip, knee, and ankle joint displacement patterns are compared across both age and speed for five immature vervet monkeys sampled approximately every 6 months over a 3 year period. The analysis indicated that, as a group, the animals displayed no consistent changes in joint patterns as they grew. However, individual animals showed consistent patterns. There were also no consistent effects of size across animals at the walk-gallop transition. This is contrary to McMahon's prediction (J. Appl. Physiol. 39:619-627, 1975) based upon his elastic-similarity model of animal scaling. With increasing speed, when symmetrical gaits were used, all of the animals tended to show a decrease in the relative positions of the hip, knee, and ankle maximum values. Furthermore, across the walk-gallop transition, the animals tended to show a decrease in the range of ankle and knee movements.     

Early development of locomotor behavior in vervet monkeys

The locomotor development of three vervet infants across approximately the first 2 months of life is described. Fairly normal-looking walking movements (as compared to adults) were seen in all the animals by approximately 1 month of age and galloping was observed by 2 months. Early locomotor footfall patterns were often aberrant and bounding-type gaits were sometimes exhibited. Most of the symmetrical gaits observed were classifiable as lateral sequence. Across the 2-month period the animals showed decreased three- and four-foot support and improvements in joint angular displacement patterns. From their earliest locomotor movements the infants showed significant linear relationship between both cycle duration and swing and stance durations of the limbs. We suggest that locomotor control mechanisms are probably fairly mature at birth but that weight support and postural control problems explain the initial locomotor difficulties exhibited by these infants.     

Kinematics of biophysically asymmetric limbs within rate of velocity development

The purpose of this study was to investigate the movement speed characteristics of 2 intrinsically different limbs. Twenty subjects volunteered to participate (10 men and 10 women). Each subject performed 5 repetitions of concentric knee and elbow extension and flexion movements at 60 through 500 d.s(-1) on an isokinetic dynamometer. Kinematic data were collected at 1,000 Hz and separated into rate of velocity development (RVD) and peak torque. Results demonstrated a significant (p < 0.05) main effect for sex for RVD and peak torque. Significant (p < 0.05) differences were also demonstrated between knee and elbow RVD and between knee and elbow peak torque at every speed tested. Neither knee and elbow RVD nor peak torque demonstrated any significant Pearson correlations at any speed tested (r = -0.17-0.41). These results collectively point to the specificity of limb speed and torque as a result of biophysical differences such as length and mass. Therefore, strength and speed may be modulated by neuromotor patterns that differ based on individual limbs.     

Bipedal locomotion in ratites (Paleognatiform): examples of cursorial birds

The gaits of five Ostriches Struthio camelus , seven Emus Dromaius novaehollandiae, two Greater Rheas Rhea americana, two Southern Cassowaries Casuarius casuarius and one Brown kiwi Apteryx australis were filmed at zoological parks. Locomotor parameters were measured using footprints on sandy tracks and video records. Osteological measurements were made on skeletons of the pelvic limbs. All of these terrestrial birds shift from wallking to running at a relative speed below 1. However, they show two different locomotor patterns: the Brown Kiwi increases its speed by increasing its stride length, mainly by increasing the protraction angle. Its hindlimbs make a flexed jointed chain system, its centre of mass is anterior, its femurs are long and the knees act in yield whereas the distal joints act in propulsion. Other ratites, particularly Ostriches, increase their speed by increasing frequency. Their centre of mass is close to the hip, their hindlimbs have an extended jointed chain system with a short erect femur, maximizing a gravity-powered system.     

Crouched posture and high fulcrum, a principle in the locomotion of small mammals: The example of the rock hyrax (Procavia capensis) (Mammalia: Hyracoidea)

The cineradiographic study of the locomotion of the rock hyrax (Procavia capensis) and the functional interpretation of its locomotory system, reveals that the main action of proximal segments is combined with flexed position and low movements of limb joints. This observation can be applied to the locomotion of other small mammals. In the forelimb, scapular rotation and translation account for more than 60% of step length. Effective shoulder joint movements are mostly restricted to less than 20°, and elbow movements range mainly between 20°-50°. The detachment of the shoulder girdle of therian mammals from the axial skeleton, and development of a supraspinous fossa, are correlated with movements at a high scapular fulcrum. Movements at such a high fulcrum are in interdependency with a crouched posture. Only flexed limbs can act as shock absorbers and prevent vertical changes in the center of gravity. Basic differences in forelimb movements exist between larger primates (humeral retraction) and smaller mammals (scapula retraction). In the hyrax, propulsion is due mainly to hip joint movements in symmetrical gaits, but sagittal lumbar spine movements play the dominant role at in-phase gaits. Joint and muscular anatomy, especially of the shoulder region, are discussed in view of the kinematic data.     

Similarities in the Neural Control of the Shoulder and Elbow Joints Belie Their Structural Differences

Movement of the hand in three dimensional space is primarily controlled by the orientation of the shoulder and elbow complexes. Due to discrepancies in proprioceptive acuity, overlap in motor cortex representation and grossly different anatomies between these joints, we hypothesized that there would be differences in the accuracy of aimed movements between the two joints. Fifteen healthy young adults were tested under four conditions – shoulder motion with the elbow constrained and unconstrained, and elbow motion with the shoulder constrained and unconstrained. End point target locations for each joint were set to coincide with joint excursions of 10, 20 or 30 degrees of either the shoulder or elbow joint. Targets were presented in a virtual reality environment. For the constrained condition, there were no significant differences in angular errors between the two joints, suggesting that the central nervous system represents linked segment models of the limb in planning and controlling movements. For the unconstrained condition, although angle errors were higher, hand position errors remained the same as those of the constrained trials. These results support the idea that the CNS utilizes abundant degrees of freedom to compensate for the potentially different contributions to end-point errors introduced by each joint.     

Symmetrical gaits and center of mass mechanics in small-bodied,primitive mammals

Widely accepted relationships between gaits (footfall patterns) and center of mass mechanics have been formulated from observations for cursorial mammals. However, sparse data on smaller or more generalized forms suggest a fundamentally different relationship. This study explores locomotor dynamics in one eutherian and five metatherian (marsupials) mammals—all small-bodied (<2 kg) with generalized body plans that utilize symmetrical gaits. Across our sample, trials conforming to vaulting mechanics occurred least frequently (<10% of all trials) while bouncing mechanics was obtained most commonly (60%); the remaining trials represented mixed mechanics. Contrary to the common situation in large mammals, there was no evidence for discrete gait switching within symmetrical gaits as speed increased. This was in part due to the common practice of grounded running. The adaptive advantage of different patterns of center-of-mass motion and their putative energy savings remain questionable in small-bodied mammals.     

Peculiarities of Activation of Muscles of the Shoulder Belt in Voluntary Two-Joint Movements of the Upper Limb

We studied coordination of central motor commands (СMCs) coming to muscles of the shoulder and shoulder belt in the course of single-joint and two-joint movements including flexion and extension of the elbow and shoulder joints. Characteristics of rectified and averaged EMGs recorded from a few muscles of the upper limb were considered correlates of the CMC parameters. Special attention was paid to coordination of CMCs coming to two-joint muscles that are able to function as common flexors (m. biceps brachii, caput breve, BBcb) and common extensors (m. triceps brachii, caput longum, TBcl) of the elbow and shoulder joints. Upper limb movements used in the tests included planar shifts of the arm from one spatial point to another resulting from either simultaneous changes in the angles of the shoulder and elbow joints or isolated sequential (two-stage) changes in these joint angles. As was found, shoulder muscles providing movements of the elbow with changes in the angle of the elbow joint, i.e., BBcb and TBcl, were also intensely involved in the performance of single-joint movements in the shoulder joint. The CMCs coming to two-joint muscles in the course of two-joint movements appeared, in the first approximation, as sums of the commands received by these muscles in the course of corresponding single-joint movements in the elbow and shoulder joints. Therefore, if we interpret the isolated forearm movement performed due to a change in the angle of the elbow joint as the main motor event, while the shoulder movement is considered the accessory one, we can conclude that realization of a two-joint movement of the upper-limb distal part is based on superposition of CMCs related to basic movements (main and accessory). Neirofiziologiya/Neurophysiology, Vol. 41, No. 1, pp. 48–56, January–February, 2009.     

Distal Forelimb Kinematics in <Emphasis Type="Italic">Erythrocebus patas</Emphasis> and <Emphasis Type="Italic">Papio anubis</Emphasis> During Walking and Galloping

When using symmetrical gaits, terrestrial digitigrade monkeys adopt less digitigrade, i.e., more palmigrade-like, hand postures as they move with faster speeds. Accordingly, it appears that, in contrast to other mammals, digitigrady is unrelated to cursoriality in primates. However, researchers have not documented the effects of speed on distal forelimb kinematics in faster asymmetrical gaits, i.e., galloping, when ground reaction forces are typically increased owing to the decreased number of contact points during a stride, combined with higher speed. Thus, it remains possible that primates use digitigrade hand postures during these higher-speed asymmetrical gaits. We investigated 3D angles in the wrist joint and metacarpophalangeal joint of 2 habitually digitigrade terrestrial monkeys, Erythrocebus patas and Papio anubis, across a large range of walking and galloping speeds on a motorized treadmill. Nonparametric analyses reveal that angles, and therefore hand postures, are not different at the subject’s walk-gallop transition. Regression analyses show that when walking, digitigrade postures are adopted at slow speeds and more palmigrade-like postures are adopted at fast speeds. Contrary to expectations, there is little change in hand postures across galloping speeds; both subjects maintained palmigrade-like hand postures with substantial joint yield and reextension during support. These results indicate that the hands are always less digitigrade at faster speeds because the joints of the distal forelimb cannot resist the higher ground reaction forces that accompany these higher speed gaits.     

Dependence between the parameters of dynamic and stationary segments of the EMG activity in the elbow joint muscles and the joint angle during realization of targeted movements in cats

In experiments on cats we studied the pattern of EMG activity recorded from the flexors and extensors of the elbow joint and related to realization of flexor targeted operant movements of the forearm. The levels of stationary EMG activity generated by the flexors at a stabilized equilibrium position of the joint demonstrated no correlation with the values of joint angles. We suppose that this feature depends on manifestation of the hysteresis effects of muscle contraction. A target position was attained mostly due to the dynamic phases of muscle activity. The respective patterns of the movement-related activity of synergic muscles significantly differed; separate components related to leaving an equilibrium state with a certain acceleration and attaining a presettled equilibrium joint angle could be differentiated in this activity. Final positions of the forearm could be significantly different at equal levels of the stationary EMG activity generated during stabilization of these positions; they depended on specificities in the time course of dynamic phase of the activity (in particular, on the time of activity decay to a stationary level). We conclude that the movement of a limb link from one equilibrium position to another is mostly controlled via coordination of the dynamic phase of muscle activity.     

Locomotor mechanics of the slender loris (Loris tardigradus)

The quadrupedal walking gaits of most primates can be distinguished from those of most other mammals by the presence of diagonal-sequence (DS) footfall patterns and higher peak vertical forces on the hindlimbs compared to the forelimbs. The walking gait of the woolly opossum (Caluromys philander), a highly arboreal marsupial, is also characterized by diagonal-sequence footfalls and relatively low peak forelimb forces. Among primates, three species--Callithrix, Nycticebus, and Loris--have been reported to frequently use lateral-sequence (LS) gaits and experience relatively higher peak vertical forces on the forelimbs. These patterns among primates and other mammals suggest a strong association between footfall patterns and force distribution on the limbs. However, current data for lorises are limited and the frequency of DS vs. LS walking gaits in Loris is still ambiguous. To test the hypothesis that patterns of footfalls and force distribution on the limbs are functionally linked, kinematic and kinetic data were collected simultaneously for three adult slender lorises (Loris tardigradus) walking on a 1.25 cm horizontal pole. All subjects in this study consistently used diagonal-sequence walking gaits and always had higher peak vertical forces on their forelimbs relative to their hindlimbs. These results call into question the hypothesis that a functional link exists between the presence of diagonal-sequence walking gaits and relatively higher peak vertical forces on the hindlimbs. In addition, this study tested models that explain patterns of force distribution based on limb protraction angle or limb compliance. None of the Loris subjects examined showed kinematic patterns that would support current models proposing that weight distribution can be adjusted by actively shifting weight posteriorly or by changing limb stiffness. These data reveal the complexity of adaptations to arboreal locomotion in primates and indicate that diagonal-sequence walking gaits and relatively low forelimb forces could have evolved independently.     

Hysteresis properties of movement in elbow joint of unanesthetized cats when antagonistic muscles are activated using different methods

Flexing and extending movements of the elbow joint were studied in unanesthetized cats. These movements were evoked by intracortical microstimulation and by vibration of the forepaw. After the animals were anesthetized the same movements were studied in response to direct stimulation of the antagonistic muscles. Interaction of the hysteresis effects in the antagonistic muscles under conditions of cortical-evoked movements in response to stimulation of two points of the cortex, one of which evoked flexion, and the other extension of the elbow joint, was studied using external local disturbance. Coactivation of the antagonists was shown to increase both the joint stiffness and the ambiguity of the equilibrium of the joint angle. This ambiguity was expressed in both the antagonistic actuating disturbances as well as in the change of the sequence for activating the antagonistic muscles. Comparison of the cortical-evoked movements and movements evoked by vibrating the forepaw when tested with an external load disturbance showed that with intracortical microstimulation the myotatic reflexes in the activated muscle are suppressed, but when vibration is used they are well defined in both antagonistic muscles. At the same time, in spite of the significantly different pathways for activating spinal neurons, the ambiguity effects displayed when determining the equilibrium of the joint angle were similar in both cases.A. A. Bogomolets Physiology Institute, Ukrainian Academy of Sciences, Kiev. Translated from Neirofiziologiya, Vol. 24, No. 3, pp. 322–330, May–June, 1991.     

Strategies used to stabilize the elbow joint challenged by inverted pendulum loading

The stiffness of activated muscles may stabilize a loaded joint by preventing perturbations from causing large displacements and injuring the joint. Here the elbow muscle recruitment patterns were compared with the forearm loaded vertically (a potentially unstable inverted pendulum configuration) and with horizontal loading. Eighteen healthy subjects were studied with the forearm vertical and supinated and the elbow flexed approximately 90 degrees. In the first experiment EMG electrodes recorded activity of biceps, triceps, and brachioradialis muscles for joint torques produced (a) by voluntarily exerting a horizontal force isometrically (b) by voluntarily flexing and extending the elbow while the forearm was loaded vertically with 135N. The relationship between the EMG and the torque generated was quantified by the linear regression slope and zero-torque intercept. In a second experiment a vertical load increasing linearly with time up to 300N was applied.In experiment 1 the EMG-torque relationships for biceps and triceps had an intercept about 10% of maximum voluntary effort greater with the vertical compared to the horizontal force, the inverse was found for Brachioradialis, but the EMG-torque slopes for both agonist and antagonistic muscles were not different. In experiment 2 there were 29 trials with minimal elbow displacement and all the three muscles activated on the order of 11% of maximum activation to stabilize the elbow; 19 trials had small elbow extension and 14 trials small flexion requiring altered muscle forces for equilibrium; 7 trials ended in large unstable displacement or early termination of the test. An analysis indicate that the observed levels of muscle activation would only provide stability if the muscles' short-range stiffness was at the high end of the published range, hence the elbow was marginally stable. The stability analysis also indicated that the small elbow extension increased stability and flexion decreased stability.     

Dependence of elbow joint stiffness measurements on speed,angle, and muscle contraction level

Elbow joint stiffness is critical to positioning the hand. Abnormal elbow joint stiffness may affect a person's ability to participate in activities of daily living. In this work, elbow joint stiffness was measured in ten healthy young adults with a device adapted from one previously used to measure stiffness in other joints. Measurements of elbow stiffness involved applying a constant-velocity rotational movement to the elbow and measuring the resultant displacement, torque, and acceleration. Elbow stiffness was then computed using a previously-established model for joint stiffness. Measurements were made at two unique elbow joint angles, two speeds, and two forearm muscle contraction levels. The results indicate that the elbow joint stiffness is significantly affected by both rotational speed and forearm muscle contraction level.     

Reconstruction of equilibrium trajectories and joint stiffness patterns during single-joint voluntary movements under different instructions

 A method for reconstructing joint compliant characteristics during voluntary movements was applied to the analysis of oscillatory and unidirectional elbow flexion movements. In different series, the subjects were given one of the following instructions: (1) do not intervene voluntarily; (2) keep the trajectory; (3) in cases of perturbations, return back to the starting position as quickly as possible (only during unidirectional movements). Under the instruction ‘keep trajectory’, the apparent joint stiffness increased by 50% to 250%. During oscillatory movements, this was accompanied by a decrease in the maximal difference between the actual and equilibrium joint trajectories and, in several cases, led to a change in the phase relation between the two trajectories. The coefficients of correlation between joint torque and angle were very high (commonly, over 0.9) under the ‘do not intervene’ instruction. They dropped to about 0.6 under the ‘keep trajectory’ and to about 0.3 under the ‘return back’ instructions. Under these two instructions, the low values of the coefficients of correlation did not allow reconstruction of segments of equilibrium trajectories and joint stiffness values in all the subjects. The results provide further support for the λ-version of the equilibrium-point hypothesis and for using the instruction ‘do not intervene voluntarily’ to obtain reproducible time patterns of the central motor command. Received: 14 December 1993/Accepted in revised form: 16 April 1994     

A comparison of the effects of agonist and antagonist muscle fatigue on performance of rapid movements

The aim of this study was to investigate the effects of agonist and antagonist muscle fatigue on the performance of rapid, self-terminating movements. Six subjects performed rapid, consecutive elbow flexion and extension movements between two targets prior to and after fatiguing either the elbow flexor or elbow extensor muscles. The experiments demonstrated consistent results. Agonist muscle fatigue was associated with a decrease in peak velocity and peak deceleration, while a decrease in peak acceleration was particularly prominent. Antagonist muscle fatigue, however, was associated with a decrease in peak deceleration, while a decrease in both the peak velocity and peak acceleration was modest and, in some tests, non-significant. The relative acceleration time (i.e. acceleration time as a proportion of the total movement time) increased when agonists were fatigued, but decreased when antagonists were fatigued. Taken together, these results emphasize the mechanical roles of the agonist and antagonist muscles; namely, the fatigue of each muscle group particularly affected the movement phase in which that group accelerated a limb, while changes of the movement kinematics pattern provided more time for action of the fatigued muscles. In addition, the results presented suggest that agonist muscle fatigue affects movement velocity more than antagonist muscle fatigue, even in movements that demonstrate prominently both mechanical and myoelectric activity of the antagonist muscles, such as rapid, self-terminating movements. Accepted: 11 February 1997     

Control of the Elbow Extensor Muscles in Slow Targeted Extensions of the Arm in Humans

Relations between the kinematic parameters of slow (non-ballistic) targeted extension movements in the elbow joint of humans and characteristics of the movement-related EMG activity in the two heads of the m. triceps brachii were analyzed. Test movements were performed under conditions of application of non-inertional external loadings directed toward flexion. It was shown that the movement-related EMG activity of the elbow extensors, similarly to what was observed in the flexors at flexion movements with the same parameters, demonstrates a complex structure and includes dynamic and stationary phases. In the former phase, in turn, initial and main components can be differentiated. The rising edge and decay of the main component of the dynamic extensor EMG phase could be approximated by exponential functions; this component was never split into a few subcomponents. Dependences between the amplitudes of m. triceps brachii EMG phases and the amplitude of the movement (or external loading) were, as a rule, nonlinear but monotonic. An increase in the test movement velocity led to an increase in the rate of rise of the rising edge of the dynamic EMG phase, while an increment in the amplitude was less significant. Under the used test conditions, the activity of the elbow extensors was usually accompanied by some coactivation of the antagonists (m. biceps brachii). It is concluded that motor commands coming to the elbow extensors at performance of the extension test movements differ from motor commands to the flexors at analogous flexion test movements by a simpler structure and more tonic pattern. Biomechanical specificities of fixation of the mentioned muscle groups to the arm bones (stability of the moment for application of the extensor force under conditions of changing the joint angle vs variable moment of the flexor force) are considered one of the main reasons for such specificity of the patterns of the extensor and flexor motor commands.     

A three-dimensional analysis of overarm throwing in experienced handball players

The aim of this study was to investigate the contribution of upper extremity, trunk, and lower extremity movements in overarm throwing in team handball. In total, 11 joint movements during the throw were analyzed. The analysis consists of maximal angles, angles at ball release, and maximal angular velocities of the joint movements and their timing during the throw. Only the elbow angle (extension movement range) and the level of internal rotation velocity of the shoulder at ball release showed a significant relationship with the throwing performance. Also, a significant correlation was found for the timing of the maximal pelvis angle with ball velocity, indicating that better throwers started to rotate their pelvis forward earlier during the throw. No other significant correlations were found, indicating that the role of the trunk and lower limb are of minor importance for team handball players.     

Effects of turn angle and pivot foot on lower extremity kinetics during walk and turn actions

This study examined lower extremity joint moments during walk and turn with different turn angles and pivot feet. Seven young adults (age 21+/-1.3 yrs) were asked to walk at a self-selected speed (1.35+/-0.15 m/s) and to turn to the right using right (spin turn) and left (step turn) pivot feet at turn angles of 0 degrees (walking straight), 45 degrees, and 90 degrees. Video and forceplate systems were employed for kinematic and kinetic data collection. Inverse dynamics approach was used to compute joint moments using segmental kinematics, ground reaction forces, and moments. The participants decreased their forward speed by increasing the ankle plantar flexion moment as the turn angle increased. The peak ankle plantar flexion moment during the braking phase increased with increasing turn angle for both spin and step turns. Ankle invertor moments were observed only in spin turns, suggesting that more ankle muscles are involved in spin turns than in step turns. The turn angle had a significant effect on the transverse plane moment profiles at the different lower extremity joints. The results suggest that the loading patterns of different anatomical structures in the lower extremity are affected by both turn angle and pivot foot during walk and turn actions.