Weight support and balance during perturbed stance in the chronic spinal cat

The intact cat maintains balance during unexpected disturbances of stance through automatic postural responses that are stereotyped and rapid. The extent to which the chronic spinal cat can maintain balance during stance is unclear, and there have been no quantitative studies that examined this question directly. This study examined whether the isolated lumbosacral cord of the chronic spinal cat can generate automatic postural responses in the hindlimbs during translation of the support surface. Responses to 16 directions of linear translation in the horizontal plane were quantified before and after spinalization at the T(6) level in terms of forces exerted by each paw against the support, motion of the body segments (kinematics), and electromyographic (EMG) activity. After spinalization, the cats were trained on a daily basis to stand on the force platform, and all four cats were able to support their full body weight. The cats usually required assistance for balance or stability in the horizontal plane, which was provided by an experimenter exerting gentle lateral force at the level of the hips. Three of the four animals could maintain independent stance for a brief period (10 s) after the experimenter stabilized them. The fourth cat maintained weight support but always required assistance with balance. Perturbations were delivered during the periods of independent stance in three cats and during assisted stance in the fourth. A response to translation in the spinal cats was observed only in those muscles that were tonically active to maintain stance and never in the flexors. Moreover, latencies were increased and amplitudes of activation were diminished compared with control. Nevertheless, flexors and extensors were recruited easily during behaviors such as paw shake and stepping. It is concluded that centers above the lumbosacral cord are required for the full elaboration of automatic postural responses. Although the spinal cat can achieve good weight support, it cannot maintain balance during stance except for brief periods and within narrow limits. This limited stability is probably achieved through spinal reflex mechanisms and the stiffness characteristics of the tonically active extensors.     

Bilateral labyrinthectomy in the cat: motor behaviour and quiet stance parameters

Summary The purpose of this study was to examine the effect of bilateral labyrinthectomy on quiet stance in the freely-standing cat. Since loss of the vestibular end organs produces marked deficits in motor behaviour, including ataxia and problems with balance, we hypothesized that labyrinthectomized animals would show impairment in quantitative measures of stance. Stance was quantified in terms of the ground reaction forces under each limb and the tonic electromyographic (EMG) activity of selected muscles. Animals were labyrinthectomized by drilling into the vestibule and removing the vestibular epithelium. Following lesion, animals were able to stand unsupported on the force platform within 2 days. To our surprise, the lesioned animals showed little change in stance parameters from the control, pre-lesion state. Thus, our hypothesis of changes in stance parameters was not supported. There was no change in the distribution of vertical forces under the limbs and no increase in sway, as measured by the area of excursion of the centre of pressure over time. The horizontal plane forces, which were diagonally directed prior to lesion, became more laterally directed and larger in amplitude. The change in direction persisted even after the animals had fully compensated for the lesion, but the force amplitudes returned to control values within 10–12 days. The change in horizontal force direction was similar to that observed in normal animals that were required to stand with their paws closer than preferred in the sagittal plane (unpublished observations). EMG activity changed in some muscles but not others, and usually transiently. One limb extensor showed decreases in tonic activity (gluteus medius), but other extensors showed increases (vastus medialis, soleus). It is likely that the changes in EMG levels were due to the biomechanics of the stance configuration related to the changes in direction of the horizontal plane forces. Kinematic recordings are needed to address this issue further.     

Vesico-somatic reflexes in the chronic spinal cat

1. In cats anaesthetized with chloralose whose spinal cords were transected 6-7 weeks previously, spontaneous contractions of the bladder had no effect on mono- or polysynaptic reflexes recorded electrically from the cut lumbar ventral spinal roots.2. In the same cats contractions of the bladder decreased hind-limb extensor and flexor reflexes recorded myographically.3. The effects of (a) pelvic nerve stimulation and distension of the bladder on mono- and polysynaptic reflexes and (b) the effects of bladder contractions on hind-limb reflexes recorded myographically in acute and chronic spinal, decerebrate and anaesthetized cats are discussed.4. The sites of interaction between vesical and somatic reflexes are discussed and further evidence is presented that bladder contractions affect the gamma-loop.     

Joint coordination during quiet stance: effects of vision

Stabilization of the center of mass (CM) is an important goal of the postural control system. Coordination of several joints along the human pendulum is required to achieve this goal. We studied the coordination among body segments with respect to horizontal CM stabilization during a quiet stance task and the effects of vision on CM stability. Subjects were asked to stand quietly on a narrow wooden block supporting only the mid-foot, with either open (EO) or closed (EC) eyes on separate trials. Instant equilibrium points (IEPs) in the center of pressure (CP) trajectory were determined when the horizontal component of the ground reaction force was zero and the CP data were decomposed into their rambling and trembling components. The joint angle, CM and CP data were divided into short cycles (time-normalized to 100 data points) or longer segments (time-normalized to 1000 data points) of equal length beginning and ending in an IEP. Motor abundance with respect to patterns of joint coordination was evaluated using the uncontrolled manifold (UCM) approach. Here, a UCM is a subspace spanning all joint combinations resulting in a given CM position. All combinations of joint angles that lie within this subspace are equivalent with respect to that CM position while joint angle combinations lying in a subspace orthogonal to the UCM lead to deviation from that CM position. UCM analysis was performed on data organized either across time within longer segments or at each point in time across multiple segments or across multiple cycles. Regardless of method of analysis, most of the variance in joint space was constrained to be within the UCM, preserving the mean CM position in both the EO and EC conditions. Joint configuration variance was significantly higher in the EC than in the EO condition although this increase occurred primarily within the UCM rather than in the orthogonal subspace that would have led to variation of the CM position. These results demonstrate the ability of the control system to selectively channel motor variability into directions in joint space that stabilize the CM position. This effect was enhanced when the task was made more challenging in the absence of vision. There was also a significant relationship between joint variance that led to a change in the CM position and, in particular, the rambling component of the CP path, lending some support to the idea that the CNS prescribes a certain stable trajectory of the CP during quiet stance that leads to a small controlled movement of the CM.     

Intracellular recording from spinal cord motoneurons in the chronic cat

This report describes chronic intracellular techniques for identifying spinal cord motoneurons and recording from them for prolonged periods of time. The preparation is the intact, unanesthetized, undrugged, normally respiring cat. Comparisons are made between data obtained in the chronic state with that reported in the “acute” preparation, and standards are suggested for intracellular recording in the chronic preparation. Various traditionally used tests of neuronal functioning are outlined in the context of determining basic electrophysiological properties of motoneurons as a function of state-dependent influences.     

Center of mass control and multi-segment coordination in children during quiet stance

This study aimed to apply an uncontrolled manifold (UCM) approach to investigate how children utilize the variability of multiple body segment movement to facilitate the center of mass (COM) control during quiet stance. Three groups of participants were included in this study: younger children (YC, mean age 6.3 years), older children (OC, mean age 10.3 years), and young adults (YA, mean age 20.5 years). Participants stood on a force platform with their hands on the iliac crests for 40 s in each trial. Two visual conditions were examined including eyes-open and eyes-closed and three trials were collected for each condition. Results showed that all three groups partitioned more variability of multi-segment movement into the UCM subspace (maintaining the mean COM position) than into the ORT subspace (a subspace orthogonal to the UCM subspace, causing the deviation of the COM from its mean position) in both eyes-open and eyes-closed conditions. Furthermore, both the YC and OC groups partitioned a significantly higher percentage of variability into the UCM subspace than the YA group regardless of visual condition. In addition, results of conventional COM variables indicated that only the YC group produced significantly faster sway velocity and greater standard deviation than the YA group. All the results together suggest that children at 6–10 years of age use a similar variability-partitioning strategy (a greater V _UCM and a smaller V _ORT) like young adults in quiet stance to facilitate the COM control, but it takes more than 10 years for children to refine this strategy and achieve an adult-like variability-partitioning capability (i.e., UCM ratio). It also suggests that postural development may include two phases in which children learn to regulate the position and movement of multiple body segments and the COM first and gain an adult-like variability-partitioning capability later.     

Influence of virtual reality on postural stability during movements of quiet stance

Introduction: Balance problems during virtual reality (VR) have been mentioned in the literature but seldom investigated despite the increased use of VR systems as a training or rehabilitation tool. We examined the influence of VR on body sway under different stance conditions. Methods: Seventeen young subjects performed four tasks (standing with feet close together or tandem stance on firm and foam surfaces for 60 s) under three visual conditions: eyes open without VR, eyes closed, or while viewing a virtual reality scene which moved with body movements. Angular velocity transducers mounted on the shoulder provided measures of body sway in the roll and pitch plane. Results: VR caused increased pitch and roll angles and angular velocities compared to EO. The effects of VR were, for the most part, indistinguishable from eyes closed conditions. Use of a foam surface increased sway compared to a firm surface under eyes closed and VR conditions. Conclusion: During the movements of quiet stance, VR causes an increase in postural sway in amplitude similar to that caused by closing the eyes. This increased sway was present irrespective of stance surface, but was greatest on foam.     

Importance of body sway velocity information in controlling ankle extensor activities during quiet stance

In literature, it has been suggested that the CNS anticipates spontaneous change in body position during quiet stance and continuously modulates ankle extensor muscle activity to compensate for the change. The purpose of this study was to investigate whether velocity feedback contributes by modulating ankle extensor activities in an anticipatory fashion, facilitating effective control of quiet stance. Both theoretical analysis and experiments were carried out to investigate to what extent velocity feedback contributes to controlling quiet stance. The experiments were carried out with 16 healthy subjects who were asked to stand quietly with their eyes open or closed. During the experiments, the center of pressure (COP) displacement (COPdis), the center of mass (COM) displacement (COMdis), and COM velocity (COMvel) in the anteroposterior direction were measured. Rectified electromyograms (EMGs) were used to measure muscle activity in the right soleus muscle, the medial gastrocnemius muscle, and the lateral gastrocnemius muscle. The simulations were performed using an inverted pendulum model that described the anteroposterior kinematics and dynamics of quiet stance. In the simulations, an assumption was made that the COMdis of the body would be regulated using a proportional-derivative (PD) controller. Two different PD controllers were evaluated in these simulations: 1) a controller with the high-derivative/velocity gain (HDG) and 2) a controller with the low-derivative/velocity gain (LDG). Cross-correlation analysis was applied to investigate the relationships between time series obtained in experiments 1) COMdis and EMGs and 2) COMvel and EMGs. Identical cross-correlation analysis was applied to investigate the relationships between time series obtained in simulations 3) COMdis and ankle torque and 4) COMvel and ankle torque. The results of these analyses showed that the COMdis was positively correlated with all three EMGs and that the EMGs temporally preceded the COMdis. These findings agree with the previously published studies in which it was shown that the lateral gastrocnemius muscle is actively modulated in anticipation of the body's COM position change. The COMvel and all three EMGs were also correlated and the cross-correlation function (CCF) had two peaks: one that was positive and another that was negative. The positive peaks were statistically significant, unlike the negative ones; they were larger than the negative peaks; and their time shifts were much shorter compared with the time shifts of the negative peaks. When these results were compared with the CCF results obtained for simulated time series, it was discovered that the cross-correlation results for the HDG controller closely matched cross-correlation results for the experimental time series. On the other hand, the simulation result obtained for LDG controller did not match the experimental results. These findings suggest that the actual postural control system during quiet stance adopts a control strategy that relies notably on velocity information and that such a controller can modulate muscle activity in anticipatory manner without using a feed-forward mechanism.     

Changes in segmental reflexes following chronic spinal cord hemisection in the cat:

The effect of a chronic spinal cord hemisection on segmental reflex transmission was studied in cats. Recordings of ventral root responses were made after a terminal transection below the initial lesion to eliminate descending influence through the intact spinal half. Procedures to ensure comparability between sides, that are lacking in earlier work on this experimental model, were introduced in the present work. It was demonstrated that there was an increase in reflex size on the lesioned (left) side, relative to the other side, of both mono-and polysynaptic reflexes. The reflex changes were found at all survival times studied (from 2 to 515 days).In control animals mono-and polysynaptic reflexes were found to be larger on the right side. It is discussed that side symmetry of reflex size is not to be presupposed even in a normal population. Detailed clinical examinations of lesioned animals were not performed, but a tendency for enhancement of the ipsilateral knee-jerk was found. There was no syndrome of spasticity and general motor recovery was very good.     

Post-effect of forward and backward locomotion on body orientation in space during quiet stance

Neural circuits responsible for stance control serve other motor tasks as well. We investigated the effect of prior locomotor tasks on stance, hypothesizing that postural post-effects of walking are dependent on walking direction. Subjects walked forward (WF) and backward (WB) on a treadmill. Prior to and after walking they maintained quiet stance. Ground reaction forces and centre of foot pressure (CoP), ankle and hip angles, and trunk inclination were measured during locomotion and stance. In WF compared to WB, joint angle changes were reversed, trunk was more flexed, and movement of CoP along the foot sole during the support phase of walking was opposite. During subsequent standing tasks, WB induced ankle extension, hip flexion, trunk backward leaning; WF induced ankle flexion and hip extension. The body CoP was displaced backward post-WB and forward post-WF. The post-effects are walking-direction dependent, and possibly related to foot-sole stimulation pattern and trunk inclination during walking.     

Asymmetric balance control between legs for quiet but not for perturbed stance

Interlateral performance asymmetry in upright balance control was evaluated in this investigation by comparing unipedal stance on the right versus the left leg. Participants were healthy young adults, hand–foot congruent preference for the right body side. Balance performance was evaluated in unperturbed quiet stance and in the recovery of balance stability following a mechanical perturbation induced by unexpected load release. Evaluation was made under availability of full sensory information, and under deprivation of vision combined with distortion of sensory inputs from the feet soles. Results from perturbed posture revealed that muscular response latency and postural sway were symmetric between the legs. Unipedal stance was more stable when the body was supported on the right as compared with the left leg. No interaction was found between leg and sensory condition. Our findings are interpreted as resulting from specialization of the sensorimotor system controlling the right leg for continuous low-magnitude postural adjustments, while corrections to large-scale stance sway are symmetrically controlled between body sides.     

Behaviour of the urethral striated sphincter and of the bladder in the chronic spinal cat

Nine cats were spinalized at the thoraco-lumbar junction (T12-L7) and the subsequent behaviour of the bladder and urethral striated sphincter was observed during periods of up to 27 days after spinalization by means of bladder manometry and of urethral electromyography. On the day following operation, the urethral sphincter responds to stimulation of its intact motor nerve, the pudic nerve by reflex (R) and direct (M) responses analogous to those of the intact animal anaesthetized with chloralose. The ratio R/M lies between 1 and 0.6 in the chronic spinal cat whereas it is generally less than 0.5 in the intact chloralose-anaesthetized cat. The tonic activity of the sphincter is weak or not present. The continence, however, is well maintained. The bladder activity appears only 4 to 8 days after spinalization. The bladder can thus void urine during brief contractions. These micturitions are always incomplete. The urethral reflex activity, either spontaneous or triggered by stimulation of the pudic nerve, may be inhibited, i: to a moderate degree by passive bladder distension; ii: almost completely by activation of vesicomotor neurones which provoke the bladder contraction. The first inhibition is seen the day after spinalization and is probably a protective reflex against vesical hypertension. The second inhibition develops progressively and in parallel to the functional recovery of vesical preganglionic neurones. It takes place on a background of antagnostic equilibrium of bladder and of urethral sphincter activities during micturition.     

Experimental knee pain impairs postural stability during quiet stance but not after perturbations

The objective of this study was to examine the effect of experimental knee-related pain on postural control. Twelve healthy subjects stood as quietly as possible on a movable force platform (that measured the centre of pressure and provided fast perturbations) before, during, and after experimental knee-related pain. Lower limb electromyographic (EMG) activity and joint angles were measured. Experimental pain was induced by injecting hypertonic saline into the infrapatellar fat pad (unilateral and bilateral) and isotonic saline was used for control sessions. Compared with the baseline and control sessions, unilateral and bilateral knee-related pain during quiet standing evoked (1) an increased sway displacement in the anterior-posterior direction (P < 0.05), (2) larger knee flexion (P < 0.05), and (3) larger EMG changes. Bilateral pain also induced (1) larger medial-lateral sway displacement and speed (P < 0.05) and (2) larger left hip flexion (P < 0.05). During forward perturbations, subjects leaned forward during both painful conditions when compared with baseline (P < 0.05). The additional impairment by bilateral pain suggests that the non-painful limb in unilateral pain conditions compensates for the impaired postural control. These results show that knee-related pain impairs postural stability during quiet standing, indicating the vulnerability of patients with knee pain to falls. This measure could potentially help clinicians who seek to assess how pain responses may contribute to patient's postural control and stability during quiet standing.