Donald C. Dunbar

Stance dependence of automatic postural adjustments in humans

SummaryThis study investigated the effect of initial stance configuration on automatic postural responses in humans. Subjects were tested in both bipedal and quadrupedal stance postures. The postural responses to horizontal translations of the supporting surface were measured in terms of the forces at the ground, movement of the body segments, and electromyographic (EMG) activity. Postural responses to the same perturbations changed with initial stance posture; these responses were biomechanically appropriate for restoring centre of mass. A change in stance configuration prior to platform movement led to a change in both the spatial and temporal organization of evoked muscle activation. Specifically, for the same direction of platform movement, during bipedal stance muscles on one side of the lower limb were activated in a distal to proximal sequence; during quadrupedal stance, muscles on the opposite side of the lower limb were activated and in a proximal to distal sequence. The most significant finding was an asymmetry in the use of the upper limbs and the lower limbs during postural corrections in quadrupedal stance. Whereas antagonists of the upper limb were either co-activated or co-inhibited, depending on the direction of translation, lower limb antagonists were reciprocally activated and inhibited. Human subjects in a quadrupedal stance posture used the lower limbs as levers, protracting or retracting the hips in order to propel the trunk back to its original position with respect to the hands and feet. Postural responses of the subjects during quadrupedal stance were remarkably similar to those of cats subjected to similar perturbations of the supporting surface. Furthermore, the same predominance of lower limb correction is characteristic of both species, suggesting that the standing cat is a good model for studying postural control in humans.

Locomotion and Posture During Terminal Branch Feeding

We investigated locomotor and postural behavior during terminal branch feeding in order to gain a better understanding of the motor capabilities of primates. We videotaped wild, juvenile bonnet macaques (Macaca radiata) in India as they fed on flower nectar in a simal tree (Bombax malabaricum). Kinematic analysis revealed that they select specific support surfaces and movements that, for their body design and postures, maximize lateral stability and minimize the chances of falling. These choices are made even though the distance and duration of travel to a selected target are frequently increased. Our discussion focuses on particular concepts of how primates contend with balance problems arboreally, potential reasons for changes in footfall patterns, and how the tail contributes to arboreal locomotion and posture. We concluded that balance problems due to the ratio of body size to branch size are usually avoided, at least among juvenile bonnet macaques, by placing the hands and feet on branches extending laterally from the central support branch and not on the central branch itself. The lateral branches permit a wide base of support, which increases lateral stability. Second, juvenile bonnet macaques have a striking ability to rapidly and repeatedly adapt their gait patterns to changing substrate design with minimal interruption to overall progression. Third, primate tails that are not morphologically specialized for prehension nevertheless have important prehensile and sensory functions in arboreal locomotion and posture.

Development of Posture and Locomotion in Free-ranging Primates

Postural and locomotor development is described for free-ranging groups of three monkey species: Macaca mulatta (rhesus macaque), M. radiata (bonnet macaque) and Presbytis entellus (hanuman langur). Behaviors are discussed in terms of infant-1, infant-2, juvenile, adult and elderly age groups. Analysis is based on high-speed motion pictures and videos. Infant-1 monkeys continually cling to their mothers ventral surface with strong hand and foot grasps. Independent motor coordination develops during the infant-2 period. The greatest postural and locomotor activity, both qualitatively and quantitatively, is achieved during the juvenile period. The adult and elderly periods are characterized by a progressive reduction in the variety and, eventually, quality of these motor abilities. Macaques and langurs differ, however, in the course of their behavioral development. These findings have implications for concepts of underlying neural mechanisms, indicate possible species differences in the development of these mechanisms, and demonstrate that the spectrum of volitional behaviors practiced by different age groups of free-ranging animals is broader than that observed in the laboratory setting.

Postural responses in the cat to unexpected rotations of the supporting surface: evidence for a centrally generated synergic organization

SummaryPostural reactions to disruptions of stance are rapid and automatic in both quadrupeds and bipeds. Current evidence suggests that these postural responses are generated by the central nervous system as patterns involving muscle synergies. This study attempted to test this hypothesis of a centrally generated postural mechanism by determining whether the same postural response could be evoked in the freely-standing cat under two different biomechanical conditions. The present work is an extension of previous experiments in which the stance of cats was perturbed by a horizontal translation of the supporting surface in the anterior and posterior directions (Rushmer et al. 1983). We now tested whether simple rotation of the metacarpo- and metatarsophalangeal (M-P) joints that mimics the digit rotation occurring during platform translation, was sufficient to evoke the translation postural response. The rotational perturbations were biomechanically different from translations in that the rotation did not cause displacement of the centre of mass of the animal, nor did it result in any significant movement about any but the M-P joints. Even so, rotational perturbations did evoke the appropriate translational muscle synergies in all four animals. Both plantar flexion rotation and headward translation activated the posterior hindlimb synergy (which included gluteus medius, semitendinosus and lateral gastrocnemius). Similarly, dorsiflexion rotation and tailward translation both activated the same anterior hindlimb synergy (iliopsoas, vastus lateralis and tibialis anterior) together with the forelimb synergy. The postural responses elicited by rotational perturbations were biomechanically inappropriate, and caused the animal to displace its own centre of mass away from the stable, control position. The most striking finding was that the group of muscles in which the medium latency postural response was evoked was different than the group from which short latency reflex responses were elicited. These data support the hypothesis that postural reactions are not merely reflex responses to local sensory inputs associated with the perturbation but, instead, represent a centrally generated response, with the muscle synergy being the controlled unit.

Stabilization and mobility of the head, neck and trunk in horses during overground locomotion: comparisons with humans and other primates

SUMMARY Segmental kinematics were investigated in horses during overground locomotion and compared with published reports on humans and other primates to determine the impact of a large neck on rotational mobility (>20 deg.) and stability (≤20 deg.) of the head and trunk. Three adult horses (Equus caballus) performing walks, trots and canters were videotaped in lateral view. Data analysis included locomotor velocity, segmental positions, pitch and linear displacements and velocities, and head displacement frequencies. Equine, human and monkey skulls and cervical spines were measured to estimate eye and vestibular arc length during head pitch displacements. Horses stabilized all three segments in all planes during all three gaits, unlike monkeys and humans who make large head pitch and yaw rotations during walks, and monkeys that make large trunk pitch rotations during gallops. Equine head angular displacements and velocities, with some exceptions during walks, were smaller than in humans and other primates. Nevertheless, owing to greater off-axis distances, orbital and vestibular arc lengths remained larger in horses, with the exception of head–neck axial pitch during trots, in which equine arc lengths were smaller than in running humans. Unlike monkeys and humans, equine head peak-frequency ranges fell within the estimated range in which inertia has a compensatory stabilizing effect. This inertial effect was typically over-ridden, however, by muscular or ligamentous intervention. Thus, equine head pitch was not consistently compensatory, as reported in humans. The equine neck isolated the head from the trunk enabling both segments to provide a spatial reference frame.

Automatic Postural Responses in the Cat: Responses to Headward and Tailward Translation*

SummaryEMG responses, vertical and A-P shear forces and kinematics of “automatic postural responses” to unexpected translational perturbations in the headward and tailward directions were studied in cats. Muscles acting on the major joints of the forelimbs and hindlimbs were studied. Movement of the animals in response to perturbation were highly stereotyped and consisted of two phases: (1) motion of the feet during platform movement while the trunk remained relatively stationary followed by (2) active correction of posture by movement of the trunk in the direction of perturbation.Vertical force changes occurred after the perturbation was well underway (latency 65 ms) and were related to the displacement of the center of mass and active correction of trunk position. Shear forces showed both passive (inertial) and active components and suggested that the majority of the torque necessary for po,stural correction was generated by the hindlimb.EMG responses in forelimb and shoulder muscles were most correlated with increase in vertical force, showing a generalized co-contraction in tailward translation (when these limbs were loaded) and little activity when the forelimbs were unloaded.EMG responses in hindlimb showed reciprocal activation of agonists and antagonists during perturbation with strong synergies of thigh and foot flexors in tailward translation and thigh and foot extensors in headward translation. The forelimb EMG patterns were most consistent with the conclusion that the forelimb is used primarily for vertical support during perturbation.It was concluded that hindlimb EMG responses were appropriate for both vertical support and performance of the postural correction. The hindlimb muscle synergies observed during translation are the “mirror image” of those observed in humans by other workers.

Cardiac Oxidative Stress Is Elevated at the Onset of Dilated Cardiomyopathy in Streptozotocin-Diabetic Rats

The association between nitric oxide synthase (eNOS and iNOS) status, oxidative stress, and cardiac function was evaluated in streptozotocin (STZ)-diabetic rats to understand the etiology of diabetic cardiomyopathy. Cardiac function was determined by echocardiography. eNOS and iNOS status and superoxide production were assessed by immunohistochemistry and chemiluminescence, respectively. In STZ-diabetic rats, stroke volume, cardiac output, and left ventricular ejection fraction were significantly lower than in controls (CT, P < .05), whereas left ventricular end-systolic volume was higher. Cardiac NOS activity increased from 161 ± 18 cpm/mg tissue in CT rats to 286 ± 20 cpm/mg tissue (P < .001) in STZ-diabetic rats. Furthermore, superoxide production and cardiac eNOS and iNOS levels were higher in STZ-diabetic rats than in CT rats (P < .05). An increased activation of cardiac eNOS and iNOS is observed concomitantly with decreased cardiac function. Thus, increased oxidative stress in the heart may be implicated in the development of dilated cardiomyopathy in STZ-diabetic rats.

Stabilization and mobility of the head and trunk in wild monkeys during terrestrial and flat-surface walks and gallops.

SUMMARY This study investigated the patterns of rotational mobility (>20°) and stability (≤20°) of the head and trunk in wild Indian monkeys during natural locomotion on the ground and on the flat-topped surfaces of walls. Adult hanuman langurs (Semnopithecus entellus) and bonnet macaques (Macaca radiata) of either gender were cine filmed in lateral view. Whole-body horizontal linear displacement, head and trunk pitch displacement relative to space (earth horizontal), and vertical head displacement were measured from the cine films. Head-to-trunk pitch angle was calculated from the head-to-space and trunk-to-space measurements. Locomotor velocities, cycle durations, angular segmental velocities, mean segmental positions and mean peak frequencies of vertical and angular head displacements were then calculated from the displacement data. Yaw rotations were observed qualitatively. During quadrupedal walks by both species, the head was free to rotate in the pitch and yaw planes on a stabilized trunk. By contrast, during quadrupedal gallops by both species, the trunk pitched on a stabilized head. During both gaits in both species, head and trunk pitch rotations were symmetrical about comparable mean positions in both gaits, with mean head position aligning the horizontal semicircular canals near earth horizontal. Head pitch direction countered head vertical displacement direction to varying degrees during walks and only intermittently during gallops, providing evidence that correctional head pitch rotations are not essential for gaze stabilization. Head-to-space pitch velocities were below 350 deg. s–1, the threshold above which, at least among humans, the vestibulo-ocular reflex (VOR) becomes saturated. Mean peak frequencies of vertical translations and pitch rotations of the head ranged from 1 Hz to 2 Hz, a lower frequency range than that in which inertia is predicted to be the major stabilizer of the head in these species. Some variables, which were common to both walks and gallops in both species, are likely to reflect constraints in sensorimotor control. Other variables, which differed between the two gaits in both species, are likely to reflect kinematic differences, whereas variables that differed between the two species are attributed primarily to morphological and behavioural differences. It is concluded that either the head or the trunk can provide the nervous system with a reference frame for spatial orientation and that the segment providing that reference can change, depending upon the kinematic characteristics of the chosen gait.

Stabilization and mobility of the head and trunk in vervet monkeys(Cercopithecus aethiops) during treadmill walks and gallops

SUMMARY The brain requires internal or external reference frames to determine body orientation in space. These frames may change, however, to meet changing conditions. During quadrupedal overground locomotion by monkeys, the head rotates on a stabilized trunk during walking, but the trunk rotates on a stabilized head during galloping. Do the same movement patterns occur during in-place locomotion? Head and trunk pitch rotations were measured, and yaw and roll rotations estimated from cine films of three adult vervet monkeys (Cercopithecus aethiops L. 1758) walking and galloping quadrupedally on a treadmill. Head and trunk rotational patterns during treadmill walks were comparable to the patterns found during overground walks. The rotational velocities of these segments during both treadmill walks and gallops were also comparable to the velocities found during natural locomotion. By contrast, whereas head and trunk rotational patterns during treadmill gallops did occur that were comparable to the patterns practiced during overground gallops, a significantly different pattern involving large and simultaneous head and trunk rotations was more commonly observed. Simultaneous head and trunk rotations may be possible during treadmill gallops because the fixed visual surround is providing an adequate spatial reference frame. Alternatively, or in addition to this visual information, a re-weighting in other sensory modalities may be occurring. Specifically, the vestibular inputs used during overground locomotion to reference gravity or a gravity-derived vector may become less important than proprioceptive inputs that are using the treadmill belt surface as a reference. Regardless, the spatial reference frame being used, blinks that occur at specific times during the largest head yaw rotations may be necessary to avoid the initiation of unwanted and potentially destabilizing lateral sway brought on by sudden increases in optic flow velocity.

Automatic postural responses in the cat: responses of proximal and distal hindlimb muscles to drop of support from a single hind- or forelimb

SummaryCats respond to drop of the support from beneath a single limb with the “diagonal stance response” (Coulmance et al. 1979). They load the limbs on the diagonal opposite to the one containing the dropped limb and unload the third supporting limb in the diagonal containing the dropped limb. Characteristic biomechanical delays in limb motion and in vertical force changes imposed upon the limbs are observed. These delays range from 30 to 45 ms, depending upon the location of the dropped limb. This study describes the kinematics of the “diagonal stance response” and the activation of selected agonist-antagonist muscle pairs acting on the joints of the hindlimb during the response. Proximal and distal hindlimb muscles respond to perturbations in groups that are appropriate to the vertical forces imposed upon the limb. When the hindlimb containing the recording electrodes is loaded by drop of the contralateral hindlimb or the ipsilateral forelimb medium latency (25–45 ms) EMG responses occur in the extensors. This response serves to stiffen the limb against the increased vertical force of loading. A similar response is observed when the hindlimb is reloaded after being dropped. In this case, however, short latency responses precede the medium latency responses in muscles that are passively stretched by the limb drop. When drop of the diagonal forelimb unloads the hindlimb containing the electrodes, medium latency responses are observed in the distal hindlimb flexors, which indicates that the unloading is evoked in part by active lifting of the limb. In most cases, the medium latency responses precede or are coincident with the changes in force imposed on the limb, suggesting that the observed responses are centrally programmed.