I have developed "tennis elbow" from lugging this book around the past four weeks, but it is worth the pain, the effort, and the aspirin. It is also worth the (relatively speaking) bargain price. Including appendixes, this book contains 894 pages of text. The entire panorama of the neural sciences is surveyed and examined, and it is comprehensive in its scope, from genomes to social behaviors. The editors explicitly state that the book is designed as "an introductory text for students of biology, behavior, and medicine," but it is hard to imagine any audience, interested in any fragment of neuroscience at any level of sophistication, that would not enjoy this book. The editors have done a masterful job of weaving together the biologic, the behavioral, and the clinical sciences into a single tapestry in which everyone from the molecular biologist to the practicing psychiatrist can find and appreciate his or

Principles of Neural Science

https://www.cs.cmu.edu/~hgeyer/Teaching/R16-899B/Papers/Winter95Gait&Posture.pdf

Human balance and posture control during standing and walking

The basal ganglia: focused selection and inhibition of competing motor programs.

We studied the extent to which automatic postural actions in standing human subjects are organized by a limited repertoire of central motor programs. Subjects stood on support surfaces of various lengths, which forced them to adopt different postural movement strategies to compensate for the same external perturbations. We assessed whether a continuum or a limited set of muscle activation patterns was used to produce different movement patterns and the extent to which movement patterns were influenced by prior experience. Exposing subjects standing on a normal support surface to brief forward and backward horizontal surface perturbations elicited relatively stereotyped patterns of leg and trunk muscle activation with 73- to 110-ms latencies. Activity began in the ankle joint muscles and then radiated in sequence to thigh and then trunk muscles on the same dorsal or ventral aspect of the body. This activation pattern exerted compensatory torques about the ankle joints, which restored equilibrium by moving the body center of mass forward or backward. This pattern has been termed the ankle strategy because it restores equilibrium by moving the body primarily around the ankle joints. To successfully maintain balance while standing on a support surface short in relation to foot length, subjects activated leg and trunk muscles at similar latencies but organized the activity differently. The trunk and thigh muscles antagonistic to those used in the ankle strategy were activated in the opposite proximal-to-distal sequence, whereas the ankle muscles were generally unresponsive. This activation pattern produced a compensatory horizontal shear force against the support surface but little, if any, ankle torque. This pattern has been termed the hip strategy, because the resulting motion is focused primarily about the hip joints. Exposing subjects to horizontal surface perturbations while standing on support surfaces intermediate in length between the shortest and longest elicited more complex postural movements and associated muscle activation patterns that resembled ankle and hip strategies combined in different temporal relations. These complex postural movements were executed with combinations of torque and horizontal shear forces and motions of ankle and hip joints. During the first 5-20 practice trials immediately following changes from one support surface length to another, response latencies were unchanged. The activation patterns, however, were complex and resembled the patterns observed during well-practiced stance on surfaces of intermediate lengths.(ABSTRACT TRUNCATED AT 400 WORDS)

Central programming of postural movements: adaptation to altered support-surface configurations.

It is generally accepted that human bipedal upright stance is achieved by feedback mechanisms that generate an appropriate corrective torque based on body-sway motion detected primarily by visual, vestibular, and proprioceptive sensory systems. Because orientation information from the various senses is not always available (eyes closed) or accurate (compliant support surface), the postural control system must somehow adjust to maintain stance in a wide variety of environmental conditions. This is the sensorimotor integration problem that we investigated by evoking anterior-posterior (AP) body sway using pseudorandom rotation of the visual surround and/or support surface (amplitudes 0.5-8 degrees ) in both normal subjects and subjects with severe bilateral vestibular loss (VL). AP rotation of body center-of-mass (COM) was measured in response to six conditions offering different combinations of available sensory information. Stimulus-response data were analyzed using spectral analysis to compute transfer functions and coherence functions over a frequency range from 0.017 to 2.23 Hz. Stimulus-response data were quite linear for any given condition and amplitude. However, overall behavior in normal subjects was nonlinear because gain decreased and phase functions sometimes changed with increasing stimulus amplitude. "Sensory channel reweighting" could account for this nonlinear behavior with subjects showing increasing reliance on vestibular cues as stimulus amplitudes increased. VL subjects could not perform this reweighting, and their stimulus-response behavior remained quite linear. Transfer function curve fits based on a simple feedback control model provided estimates of postural stiffness, damping, and feedback time delay. There were only small changes in these parameters with increasing visual stimulus amplitude. However, stiffness increased as much as 60% with increasing support surface amplitude. To maintain postural stability and avoid resonant behavior, an increase in stiffness should be accompanied by a corresponding increase in damping. Increased damping was achieved primarily by decreasing the apparent time delay of feedback control rather than by changing the damping coefficient (i.e., corrective torque related to body-sway velocity). In normal subjects, stiffness and damping were highly correlated with body mass and moment of inertia, with stiffness always about 1/3 larger than necessary to resist the destabilizing torque due to gravity. The stiffness parameter in some VL subjects was larger compared with normal subjects, suggesting that they may use increased stiffness to help compensate for their loss. Overall results show that the simple act of standing quietly depends on a remarkably complex sensorimotor control system.

Sensorimotor integration in human postural control

A scheme for understanding the organization of human postural movements is developed in the format of a position paper. The structural characteristics of the body and the geometry of muscular actions are incorporated into a three-dimensional graphical representation of human movement mechanics in the sagittal plane. A series of neural organizational hypotheses limit a theoretically infinite number of combinations of muscle contractions and associated movement trajectories for performing postural corrections: (1) Controls are organized to use the minimum number of muscles; (2) frequently performed movements are organized to require a minimum of neural decision-making.These hypotheses lead to the prediction that postural movements are composed of muscle contractile strategies derived from a limited set of distinct contractile patterns. The imposition of two mechanical constraints related to the configuration of support and to requirements for body stability with respect to gravity predict the conditions under which individual movement strategies will be deployed.A complementary organizational scheme for the senses is developed. We show that organization of postural movements into combinations of distinct strategies simplifies the interpretation of sensory inputs. The fine-tuning of movement strategies can be accomplished by breaking down the complex array of feedback information into a series of scalar quantities related to the parameters of the movement strategies. For example, the magnitude, aim, and curvature of the movement trajectory generated by an individual strategy can be adjusted independently.The second half of the report compares theoretical predictions with a series of actual experimental observations on normal subjects and patients with known sensory and motor disorders. Actual postural movements conform to theoretical predictions about the composition of individual movement strategies and the conditions under which each strategy is used. Observations on patients suggest how breakdowns in individual steps within the logical process of organization can lead to specific movement abnormalities.Discussion focuses on the areas needing further experimentation and on the implications of the proposed organizational scheme. We conclude that although our organizational scheme is not new in demonstrating the need for simplifying the neural control of movement, it is perhaps original in imposing discrete logical control upon a continuous mechanical system. The attraction of the scheme is that it provides a framework compatible with both mechanical and physiological information and amenable to experimental testing.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0140525X00020008

The organization of human postural movements: A formal basis and experimental synthesis

Doubt about the role of stretch reflexes in movement and posture control has remained in part because the questions of reflex “usefulness” and the postural “set” have not been adequately considered in the design of experimental paradigms. The intent of this study was to discover the stabilizing role of stretch reflexes acting upon the ankle musculature while human subjects performed stance tasks requiring several different postural “sets”. Task specific differences of reflex function were investigated by experiments in which the role of stretch reflexes to stabilize sway during stance could be altered to be useful, of no use, or inappropriate. Because the system has available a number of alternate inputs to posture (e.g., vestibular and visual), stretch reflex responses were in themselves not necessary to prevent a loss of balance. Nevertheless, 5 out of 12 subjects in this study used long-latency (120 msec) stretch reflexes to help reduce postural sway. Following an unexpected change in the usefulness of stretch reflexes, the 5 subjects progressively altered reflex gain during the succeeding 3–5 trials. Adaptive changes in gain were always in the sense to reduce sway, and therefore could be attenuating or facilitating the reflex response. Comparing subjects using the reflex with those not doing so, stretch reflex control resulted in less swaying when the task conditions were unchanging. However, the 5 subjects using reflex controls oftentimes swayed more during the first 3–5 trials after a change, when inappropriate responses were elicited. Four patients with clinically diagnosed cerebellar deficits were studied briefly. Among the stance tasks, their performance was similar to normal in some and significantly poorer in others. Their most significant deficit appeared to be the inability to adapt long-latency reflex gain following changes in the stance task. The study concludes with a discussion of the role of stretch reflexes within a hierarchy of controls ranging from muscle stiffness up to centrally initiated responses.

Adapting reflexes controlling the human posture.

1. We have examined rapid postural adjustments associated with a class of voluntary movements that disturb postural equilibrium. In the text that follows, these motor activities are termed associated postural adjustments and voluntary focal movements, respectively. Standing human subjects performed a variety of movement tasks on a hand-held manipulandum, resulting in disturbances to their postural equilibrium. The experimental use of movements that interact with the subject's environment in a relatively simple was permitted a more precise comparison of the postural adjustments with their associated focal movements. 2. Subjects either pulled or pushed on a stiff interface (the handle) or they responded in a predetermined way to handle perturbations. These activities were carried out with various degrees of steady-state postural stability. Prior to and during these movements, support surface and handle forces, electromyographic (EMG) signals, and body sway were monitored. 3. In addition to previously shown postural adjustments associated with reaction-time armed movements, we have demonstrated these postural activities occur in concept with segmental stretch reflexes and self-initiated (untriggered) movements. Postural adjustments were initiated shortly before all focal movements tested except the short-latency component of the biceps stretch reflex (25- to 30-ms latency). However, this reflex component was rarely elicited by handle perturbations in free-standing subjects; therefore, postural adjustments usually preceded any biceps activity under this condition. 4. By varying the degree of steady-state postural equilibrium, a reciprocal gain/threshold relationship between postural and focal components was documented, i.e., when stability was high, postural activity was reduced or absent and focal activity enhanced. Conversely, the biceps stretch reflex was difficult to elicit under any condition where the subjects was not fully supported in the direction of movement and reaction times of focal movements were prolonged. 5. Postural activities associated with focal movements were found to share a number of organizational properties with automatic postural adjustments to support surface movements. Specifically, the postural muscle synergies were equivalent in muscle composition, relative activation magnitudes, and relative temporal sequencing. Furthermore, both types of postural adjustments were highly specific in locus and magnitude to the quality of steady-state postural equilibrium (e.g., postural "set"). 6. A conceptual model is proposed that suggests one simple way in which the reciprocal influence of postural set on postural and focal movement components and their temporal sequencing might be accomplished. Furthermore, we propose in this model a common central organization of postural adjustments associated with focal movements and those elicited by support-surface movements.

Properties of postural adjustments associated with rapid arm movements

This study examines the roles of somatosensory and vestibular information in the coordination of postural responses. The role of somatosensory information was examined by comparing postural responses of healthy control subjects prior to and following somatosensory loss due to hypoxic anesthesia of the feet and ankles. The role of vestibular information was evaluated by comparing the postural responses of control subjects and patients with bilateral vestibular loss. Postural responses were quantified by measuring 1) spatial and temporal characteristics of leg and trunk EMG activation; 2) ankle, knee, and hip joint kinematics, and 3) surface forces in response to anterior and posterior surface translations under different visual and surface conditions. Results showed that neither vestibular nor somatosensory loss resulted in delayed or disorganized postural responses. However, both types of sensory deficits altered the type of postural response selected under a given set of conditions. Somatosensory loss resulted in an increased hip strategy for postural correction, similar to the movement strategy used by control subjects while standing across a shortened surface. Vestibular loss resulted in a normal ankle strategy but lack of a hip strategy, even when required for the task of maintaining equilibrium on a shortened surface. Neither somatosensory nor vestibular loss resulted in difficulty in utilizing remaining sensory information for orientation during quiet stance. These results support the hypothesis that cutaneous and joint somatosensory information from the feet and ankles may play an important role in assuring that the form of postural movements are appropriate for the current biomechanical constraints of the surface and/or foot. The results also suggest that vestibular information is necessary in controlling equilibrium in a task requiring use of the hip strategy. Thus, both somatosensory and vestibular sensory information play important roles in the selection of postural movement strategies appropriate for their environmental contexts.

L. M. Nashner

Papers

Central programming of postural movements: adaptation to altered support-surface configurations.

The organization of human postural movements: A formal basis and experimental synthesis

Adapting reflexes controlling the human posture.

Properties of postural adjustments associated with rapid arm movements

Postural strategies associated with somatosensory and vestibular loss.