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Vestibular-neck interaction and transformation of sensory coordinates.

The article considers findings and concepts on vestibular-proprioceptive interaction for self-motion perception and postural control under the form of simple describing models. It points out that vestibular-neck interaction is only a small fraction of an extended mechanism of co-ordinate transformations. This links together the different parts of our bodies, so that sensory information arising in one part of the body can be used for perceptual or motor tasks in other parts. Particular emphasis is put on the problems that arise from imperfect signal transduction in the vestibular semicircular canal systems at low stimulus frequencies/velocities. Also, a "down-and-up-channeling" principle is suggested, by which the body support is linked via coordinate transformations to the internal notion of physical space provided by the vestibular system. Furthermore, the following question is addressed: how does the brain use visual input to overcome the vestibular deficiencies, at the risk of visual self-motion illusions? Finally, a conceptual model of postural control is presented in which a proprioceptive feedback loop that links the body to its support surface is merged with a loop for postural stabilization in space.

Humans↗

Characterization of contralateral torques during static hip efforts in healthy subjects and subjects with hemiparesis.

Contralateral torques exerted at the hip were measured in healthy subjects and subjects with hemiparesis performing unilateral static hip efforts in abduction, adduction, flexion and extension, in a sitting position, at two torque levels. In general, the ipsilateral hip efforts were accompanied by mirrored contralateral torques in both groups of subjects. The directionality of these contralateral torques indicates that their action at the pelvis is mechanically opposite to the ipsilateral efforts, suggesting that they ensure the stabilization of the pelvis. In healthy subjects, analyses of variance showed no difference in the magnitude of the contralateral torques with regard to which limb was used to perform the task. However, a significant increase in magnitude was demonstrated in the contralateral torques concurrent with the increasing level of effort requested ipsilaterally. In hemiparetic subjects, when performing the tasks with their paretic limb, the magnitude of the contralateral torques was significantly increased in the non-paretic limb when compared with those measured in the paretic limb during non-paretic limb efforts. Based on the present results, a model of postural control is presented to explain the relationship between the ipsilateral and contralateral torques. Using this model, it is hypothesized that the increased contralateral torques observed in hemiparetic subjects when performing the tasks with their paretic limb is related to the weakness of the paretic muscles. The clinical importance of exercises used for the re-education of the paretic lower limb in this population, which consist of resisting the non-paretic hip movements in order to strengthen the paretic hip muscles, is discussed in light of these results.

Adult↗

Lumbosacral orthoses reduce trunk muscle activity in a postural control task.

Biomechanical modeling estimated that trunk muscle activity during various tasks could be reduced by 1-14% without the loss of spine stability when a lumbosacral orthosis (LSO) is worn [Cholewicki, J., 2004. The effects of lumbosacral orthoses on spine stability: what changes in EMG can be expected? Journal of Orthopedic Research 22, 1150-1155]. The present study experimentally tested these theoretical predictions in an unstable sitting task. This task required subjects to balance on a seat supported by a plastic hemisphere (slashed circle=30cm) and placed on a force plate that tracked the center of pressure (CoP). The average CoP velocity quantified subjects' performance. Healthy subjects (12 males, 11 females) balanced for 20s in 3 trials performed with and without the LSO in random order. EMG was recorded bilaterally from rectus abdominis (RA), external oblique (EO), thoracic (TES) and lumbar erector spinae (LES), and expressed as the % of maximum voluntary activation (%MVA). There was no difference in the balance performance with and without the LSO (p=0.13). However, EMG averaged across the trials was significantly lower in the LSO, as compared to the No LSO condition, for TES (5.8+/-3.2 vs. 6.4+/-3.7%MVA, p=0.02) and LES (3.7+/-1.5 vs. 5.9+/-3.9%MVA, p=0.01). No significant differences were present in the abdominal muscle activity. These results agree with earlier spine modeling simulations, which predicted the greatest reduction in muscle activity due to LSO to occur in TES and LES. It was hypothesized that such a reduction in muscle co-contraction could benefit patients with low back pain, who exhibit elevated muscular activity during postural tasks such as walking, standing and sitting.

Abdominal Muscles↗

A double-inverted pendulum model for studying the adaptability of postural control to frequency during human stepping in place.

In order to analyze the influence of gravity and body characteristics on the control of center of mass (CM) oscillations in stepping in place, equations of motion in oscillating systems were developed using a double-inverted pendulum model which accounts for both the head-arms-trunk (HAT) segment and the two-legged system. The principal goal of this work is to propose an equivalent model which makes use of the usual anthropometric data for the human body, in order to study the ability of postural control to adapt to the step frequency in this particular paradigm of human gait. This model allows the computation of CM-to-CP amplitude ratios, when the center of foot pressure (CP) oscillates, as a parametric function of the stepping in place frequency, whose parameters are gravity and major body characteristics. Motion analysis from a force plate was used to test the model by comparing experimental and simulated values of variations of the CM-to-CP amplitude ratio in the frontal plane versus the frequency. With data from the literature, the model is used to calculate the intersegmental torque which stabilizes the HAT when the Leg segment is subjected to a harmonic torque with an imposed frequency.

Biomechanical Phenomena↗

The effect of predictable and unpredictable motor tasks on postural control after traumatic brain injury.

This study examined the effects of environmental predictability on postural control during functional reaching while seated in healthy individuals and patients with traumatic brain injury (TBI). The postural perturbation used required reaching to the left versus reaching to the right, while seated, under predictable versus unpredictable conditions. Indexes of postural control--trajectory stability and response reach times--were measured using an electromechanical system in ten patients with TBI and ten healthy subjects. In the TBI group, greater trajectory stability and shorter response reach time were recorded under unpredictable conditions when reaching to both the right and left (d-index 0.57-2.3). In the control group, greater trajectory stability and shorter response time were recorded under predictable and unpredictable environments compared with the TBI group (d-index 0.46-0.95). This study refutes the hierarchical, predictable-to-unpredictable-environment model of postural control evaluation and treatment. The relationship between information processing demands and postural skill appears more complex than a simple linear association. Predictable and unpredictable conditions may be used concurrently, not sequentially, in TBI rehabilitation.

Adult↗

Training affects the development of postural adjustments in sitting infants.

1. The present study addressed the question of whether daily balance training can affect the development of postural adjustments in sitting infants. 2. Postural responses during sitting on a moveable platform were assessed in twenty healthy infants at 5-6, 7-8 and 9-10 months of age. Multiple surface EMGs and kinematics were recorded while the infants were exposed to slow and fast horizontal forward (Fw) and backward (Bw) displacements of the platform. After the first session the parents of nine infants trained their child's sitting balance daily. 3. At the youngest age, when none of the infants could sit independently, the muscle activation patterns were direction specific and showed a large variation. This variation decreased with increasing age, resulting in selection of the most complete responses. Training facilitated response selection both during Fw and Bw translations. This suggests a training effect on the first level of the central pattern generator (CPG) model of postural control. 4. Training also affected the development of response modulation during Fw translations. It accelerated the development of: (1) the ability to modulate EMG amplitude with respect to platform velocity and initial sitting position, (2) antagonist activity and (3) a distal onset of the response. These findings point to a training effect on the second level of the CPG model of postural adjustments.

Age Factors↗

The effect of environmental regulations on postural control after stroke.

OBJECTIVES: The primary objective of this study was to examine the effect of environmental predictability on postural control after stroke. A reaching task for seated subjects was used as the postural perturbation. Trajectory stability (the pathway followed by the subject's body center for pressure with respect to time during the reaching task) was used as the index of postural control. It was hypothesized that trajectory stability would be greater under predictable conditions. METHOD: A specially designed electromechanical system was used to measure the trajectory stability ratios for 100 subjects, 50 with poststroke hemiplegia and 50 who had not had stroke. All subjects completed a task that required reaching to the left versus reaching to the right, under predictable versus unpredictable conditions. Postural control was measured via a trajectory instability ratio in both the anterior-posterior and medial-lateral planes. RESULTS: Although the effect of predictability on postural control was significant, it was not as hypothesized for both groups. There was greater trajectory stability under unpredictable conditions when reaching to the right as measured in both the anterior-posterior and medial-lateral planes and when reaching to the left as measured in the medial-lateral plane. CONCLUSION: These findings refute the assumption of the hierarchical, predictable-to-unpredictable-environment model for postural control evaluation and treatment. The relationship between information processing demands and postural skill is probably more complex than the simple linear association implied. Perhaps the two conditions, predictable and unpredictable, should be worked on concurrently, not sequentially.

Adult↗

Simulating mechanical consequences of voluntary movement upon whole-body equilibrium: the arm-raising paradigm revisited.

Voluntary arm-raising movement performed during the upright human stance position imposes a perturbation to an already unstable bipedal posture characterised by a high body centre of mass (CoM). Inertial forces due to arm acceleration and displacement of the CoM of the arm which alters the CoM position of the whole body represent the two sources of disequilibrium. A current model of postural control explains equilibrium maintenance through the action of anticipatory postural adjustments (APAs) that would offset any destabilising effect of the voluntary movement. The purpose of this paper was to quantify, using computer simulation, the postural perturbation due to arm raising movement. The model incorporated four links, with shoulder, hip, knee and ankle joints constrained by linear viscoelastic elements. The input of the model was a torque applied at the shoulder joint. The simulation described mechanical consequences of the arm-raising movement for different initial conditions. The variables tested were arm inertia, the presence or not of gravity field, the initial standing position and arm movement direction. Simulations showed that the mechanical effect of arm-raising movement was mainly local, that is to say at the level of trunk and lower limbs and produced a slight forward displacement of the CoM (1.5 mm). Backward arm-raising movement had the same effect on the CoM displacement as the forward arm-raising movement. When the mass of the arm was increased, trunk rotation increased producing a CoM displacement in the opposite direction when compared to arm movement performed without load. Postural disturbance was minimised for an initial standing posture with the CoM vertical projection corresponding to the ankle joint axis of rotation. When the model was reduced to two degrees of freedom (ankle and shoulder joints only) the postural perturbation due to arm-raising movement increased compared to the four-joints model. On the basis of these results the classical assumption that APAs stabilise the CoM is challenged.

Arm↗

Postural feedback responses scale with biomechanical constraints in human standing.

We tested whether human postural responses can be described in terms of feedback control gains, and whether these gains are scaled by the central nervous system to accommodate biomechanical constraints. A feedback control model can describe postural responses for a wide range of perturbations, but biomechanical constraints-such as on the torque that can be exerted on the ground-make a single set of feedback gains inappropriate for all perturbations. To observe how postural responses change with perturbation magnitude, we applied fast, backward perturbations of magnitudes 3-15 cm to 13 healthy young volunteers (4 men, 9 women, aged 20-32 years). We used a 3-segment, sagittal-plane biomechanical model and a linear state feedback controller to reproduce the observed postural responses. Optimization was used to identify the best-fit feedback control gains for each trial. Results showed that trajectories of joint angles and joint torques were scaled with perturbation magnitude. This scaling occurred gradually, rather than abruptly changing at magnitudes where biomechanical constraints became active. Feedback gains were found to fit reasonably well with data ( R(2)=0.92) and to be multivariate and heterogenic in character, meaning that the torque produced at any joint is generally a function of motions not only at the same joint, but other joints as well. Hip gains increased and ankle gains decreased nearly linearly with perturbation magnitude, in accordance with biomechanical limitations on ground reaction torque. These results indicate that postural adjustments can be described as a single feedback control scheme, with scalable heterogenic gains that are adjusted according to biomechanical constraints.

Adult↗

Complexities in ETS-domain transcription factor function and regulation: lessons from the TCF (ternary complex factor) subfamily. The Colworth Medal Lecture.

The ETS-domain transcription factor family can be divided into a series of subfamilies. Elk-1 represents the founding member of the ternary complex factor (TCF) subfamily. By focusing on the TCF subfamily, we can demonstrate the complexities that exist in the function and regulation of ETS-domain transcription factors. This article focuses on Elk-1 in detail and summarizes the functions of other TCFs. The key themes covered include the domain structure of the TCFs, the mechanisms of complex formation with serum response factor, regulation of TCFs by mitogen-activated protein kinase cascades, and transcriptional regulatory properties of the TCFs. Finally, the emerging role of the TCFs in vivo is discussed. A picture is developing indicating that, while these proteins exhibit significant sequence and functional conservation, key differences in their structure and regulation are being identified which may relate to unique functions of these proteins in vivo.

Amino Acid Sequence↗

Assessing muscle stiffness from quiet stance in Parkinson's disease.

In previous studies, we developed a postural stiffness measure that is extracted from foot center-of-pressure (COP) trajectories from quietly standing individuals and is based on an analytical mechanical model of posture control. Here we apply this measure to patients with Parkinson's disease (PD). We correlated the postural stiffness measure with different clinical rating scales, obtained from patients. Kendall's rank correlation was highly significant between the stiffness measure and rigidity, bradykinesia, posture impairment, gait, and leg agility, respectively, as rated by the Unified Parkinson's Disease Rating Scale. These results provide further evidence that a higher intrinsic muscle stiffness may contribute to the aforementioned clinically defined symptoms. From a clinical standpoint, this work indicates that the proposed postural stiffness measure may be useful as an assessment tool for the evaluation of PD patients subsequent to pharmacological and surgical treatment.

Disability Evaluation↗

Organization of compensatory postural coordination patterns.

The authors investigated whether compensatory postural coordination patterns are organized according to the same dynamical principles as are nonequilibrium phase transitions. Eight participants were asked to maintain upright balance on a moving platform that was sinusoidally translated in the anterior-posterior direction and was systematically increased and decreased 0.19 Hz as a step function every 10 platform cycles through the frequency range 0.19-1.46 Hz. At low platform frequencies, all participants exhibited small joint angular motions with high variability, and the relative phase between the joint motions exhibited drifting patterns and large fluctuations. As platform frequency increased, the amplitude of joint motion increased systematically and joint-specific oscillatory patterns emerged. The findings provided no evidence for a Hopf bifurcation or hysteresis in the transitions of postural coordination modes, however, or, more generally, a basis for distinguishing the relevance of linear versus nonlinear models of postural control.

Adult↗

Effects of visual and auditory short-term memory tasks on the spatiotemporal dynamics and variability of postural sway.

The authors measured postural sway while participants (N = 20 in each experiment) stood on a rigid or a compliant surface, with their eyes open or closed, and while they did or did not perform a short-term memory (STM) task. In Experiment 1, the STM stimuli were presented visually; in Experiment 2, the stimuli were presented auditorily. In both experiments, fine-scaled, mediolateral postural-sway variability decreased as the cognitive load imposed by the STM task increased. That effect was independent of support surface and vision manipulations. The spatiotemporal profile of postural sway was affected by both visual and auditory STM tasks, but to a greater degree by the auditory task. The authors discuss implications of the results for theories and models of postural control.

Adolescent↗

Resonance in a mathematical model of baroreflex control: arterial blood pressure waves accompanying postural stress.

A mathematical model of the arterial baroreflex was developed and used to assess the stability of the reflex and its potential role in producing the low-frequency arterial blood pressure oscillations called Mayer waves that are commonly seen in humans and animals in response to decreased central blood volume. The model consists of an arrangement of discrete-time filters derived from published physiological studies, which is reduced to a numerical expression for the baroreflex open-loop frequency response. Model stability was assessed for two states: normal and decreased central blood volume. The state of decreased central blood volume was simulated by decreasing baroreflex parasympathetic heart rate gain and by increasing baroreflex sympathetic vaso/venomotor gains as occurs with the unloading of cardiopulmonary baroreceptors. For the normal state, the feedback system was stable by the Nyquist criterion (gain margin = 0.6), but in the hypovolemic state, the gain margin was small (0.07), and the closed-loop frequency response exhibited a sharp peak (gain of 11) at 0.07 Hz, the same frequency as that observed for arterial pressure fluctuations in a group of healthy standing subjects. These findings support the theory that stresses affecting central blood volume, including upright posture, can reduce the stability of the normally stable arterial baroreflex feedback, leading to resonance and low-frequency blood pressure waves.

Adult↗

A predictive model study of the visual contribution to canine postural control.

Dogs were trained to stand on a movable table and their quiet stance was perturbed by osccillation of the table during normal sighted condition and during blindfolded condition. The data formed the frequency response characteristic (describing function) for postural control with and without visual input. A feedback model was tested to assess the effect of visual input during the perturbation of quiet stance. The results of the tests of the model indicate that the effect of a visual input depends on the context of the multiple sensory factors influencing postural control. The effectiveness of the visual input increases if there is a conflict between the visually derived body position cues and the other cues that indicate the orientation of the body.

Animals↗

Biomechanical models for vibration feedthrough to hands and head for a semisupine pilot.

A series of tracking experiments under vibration has been carried out on the AMRL/BBV shaker facilities covering three axes of vibration with sinusoidal and random waveforms and different control stick configurations. Based on this and other data, a lumped-parameter biomechanical model has been evolved to suit the needs of aircraft control system designers for the new generation of low-altitude, high-speed bombers and highly maneuverable fighters. This paper shows that measured vibration feedthrough to hands and head can be adequately described by this model when linearized about the appropriate configuration of display, posture, and control. The model includes effects of: semisupine torso; sliding hip, plus rocking chest supported on a compliant buttocks/seat; head bobbing on an articulated neck; upper arm and forearm links plus grip-interface compliance, driven by an active neuromuscular system; elbow rest (optional); and stick "feel system" dynamics. Examples are given of the model's application to predict effects of: a 65 degrees semisupine seat, apparent impedance increase of a control stick under pilot control, and a sliding arm rest.

Aerospace Medicine↗

Postural dynamics in the standing human.

The purpose of this study was to develop a mathematical model of the linkage dynamics in upright standing, and to use this model to study output principles for postural control. The standing human was modelled in the sagittal plane as a three-segment linkage. Mechanical disturbances were simulated as forces which could be applied at various points in this linkage. An iterative approach was used to find joint torque combinations which would restore balance within 80 ms of these mechanical disturbances. The model predicted that a specific proportional relationship was necessary between the hip, knee and ankle torques in order for balance to be restored. This proportional relationship was shown to be a function of the model structure, but independent of the location, direction and amplitude of the disturbance. These predictions were tested experimentally. A disturbance apparatus was designed to apply an impulsive force to the subjects. The joint torque responses of the subjects were in quantitative agreement with the predictions of the model. The results suggest that a fixed relationship between joint torques may be required to restore balance, and this fixed relationship may make the task of postural control simpler for the nervous system.

Computer Simulation↗