The computational brain: Patricia S. Churchland and Terrence J. Sejnowski (MIT Press, Cambridge, MA, 1992); xi, 544 pages, $39.95

This is a review of the proprioceptive senses generated as a result of our own actions. They include the senses of position and movement of our limbs and trunk, the sense of effort, the sense of force, and the sense of heaviness. Receptors involved in proprioception are located in skin, muscles, and joints. Information about limb position and movement is not generated by individual receptors, but by populations of afferents. Afferent signals generated during a movement are processed to code for endpoint position of a limb. The afferent input is referred to a central body map to determine the location of the limbs in space. Experimental phantom limbs, produced by blocking peripheral nerves, have shown that motor areas in the brain are able to generate conscious sensations of limb displacement and movement in the absence of any sensory input. In the normal limb tendon organs and possibly also muscle spindles contribute to the senses of force and heaviness. Exercise can disturb proprioception, and this has implications for musculoskeletal injuries. Proprioceptive senses, particularly of limb position and movement, deteriorate with age and are associated with an increased risk of falls in the elderly. The more recent information available on proprioception has given a better understanding of the mechanisms underlying these senses as well as providing new insight into a range of clinical conditions.

The Proprioceptive Senses: Their Roles in Signaling Body Shape, Body Position and Movement, and Muscle Force

The motor system may use internal predictive models of the motor apparatus to achieve better control than would be possible by negative feedback. Several theories have proposed that the cerebellum may form these predictive representations. In this article, we review these theories and try to unify them by reference to an engineering control model known as a Smith Predictor. We suggest that the cerebellum forms two types of internal model. One model is a forward predictive model of the motor apparatus (e.g., limb and muscle), providing a rapid prediction of the sensory consequences of each movement. The second model is of the time delays in the control loop (due to receptor and effector delays, axonal conductances, and cognitive processing delays). This model delays a copy of the rapid prediction so that it can be compared in temporal register with actual sensory feedback from the movement. The result of this comparison is used both to correct for errors in performance and as a training signal to learn the first model. We discuss evidence that the cerebellum could form both of these models and suggest that the cerebellum may hold at least two separate Smith Predictors. One, in the lateral cerebellum, would predict the movement outcome in visual, egocentric, or peripersonal coordinates. Another, in the intermediate cerebellum, would predict the consequences in motor coordinates. Generalization of the Smith Predictor theory is discussed in light of cerebellar involvement in nonmotor control systems, including autonomic functions and cognition.

Is the cerebellum a smith predictor

In the brain and heart, rapidly inactivating (A-type) voltage-gated potassium (Kv) currents operate at subthreshold membrane potentials to control the excitability of neurons and cardiac myocytes. Although pore-forming alpha-subunits of the Kv4, or Shal-related, channel family form A-type currents in heterologous cells, these differ significantly from native A-type currents. Here we describe three Kv channel-interacting proteins (KChIPs) that bind to the cytoplasmic amino termini of Kv4 alpha-subunits. We find that expression of KChIP and Kv4 together reconstitutes several features of native A-type currents by modulating the density, inactivation kinetics and rate of recovery from inactivation of Kv4 channels in heterologous cells. All three KChIPs co-localize and co-immunoprecipitate with brain Kv4 alpha-subunits, and are thus integral components of native Kv4 channel complexes. The KChIPs have four EF-hand-like domains and bind calcium ions. As the activity and density of neuronal A-type currents tightly control responses to excitatory synaptic inputs, these KChIPs may regulate A-type currents, and hence neuronal excitability, in response to changes in intracellular calcium.

Modulation of A-type potassium channels by a family of calcium sensors

Whilst exposure to vibration is traditionally regarded as perilous, recent research has focussed on potential benefits. Here, the physical principles of forced oscillations are discussed in relation to vibration as an exercise modality. Acute physiological responses to isolated tendon and muscle vibration and to whole body vibration exercise are reviewed, as well as the training effects upon the musculature, bone mineral density and posture. Possible applications in sports and medicine are discussed. Evidence suggests that acute vibration exercise seems to elicit a specific warm-up effect, and that vibration training seems to improve muscle power, although the potential benefits over traditional forms of resistive exercise are still unclear. Vibration training also seems to improve balance in sub-populations prone to fall, such as frail elderly people. Moreover, literature suggests that vibration is beneficial to reduce chronic lower back pain and other types of pain. Other future indications are perceivable.

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Vibration as an exercise modality: how it may work, and what its potential might be.

1. Previous studies have used tendon vibration to investigate kinesthetic illusions in the isometric limb and end point control in the moving limb. These previous studies have shown that vibration ...

Proprioceptive consequences of tendon vibration during movement

1. Recent studies have shown that the CNS uses proprioceptive information to coordinate multijoint movement sequences; proprioceptive input related to the kinematics of one joint rotation in a move...

Proprioceptive coordination of movement sequences: role of velocity and position information

1. The individual joint rotations of a movement sequence might be controlled either by a central motor plan or by motion-dependent (i.e., kinesthetic) sensory input. Most previous research has focused on how the nervous system uses central motor plans to control movement sequences. This study examined how the nervous system uses kinesthetic input to control a multijoint movement sequence. 2. Human subjects were trained to extend the elbow horizontally at 22 degrees/s and to open the hand as the elbow passed through a 2 degrees-wide target zone. Different distances to the target zone were used to examine a wide range of movement times of the elbow to target zone (i.e., 150-1,500 ms). 3. A hydraulic apparatus simulated a spring resistance to the elbow extension. In some trials, the spring constant was unexpectedly increased or decreased just before the subject initiated the elbow extension, causing the elbow to slow down or speed up. Because these changes in spring constant were randomly imposed and because no visual feedback was available, subjects had to use kinesthetic input to control this motor task. 4. The experimental subjects employed two different strategies for the use of kinesthetic input to control this motor task. In the first strategy, the subjects used kinesthetic input related to the elbow rotation to detect and correct velocity errors caused by the changes in spring constant. The onset of error correction varied between 92 and 196 ms after the appearance of velocity errors. The proportion of the error corrected by the time the elbow reached the target zone varied between 31 and 78%, depending on the movement time to the target zone. However, because this correction for velocity errors was neither instantaneous nor complete, the changes in spring constant caused leads and lags in the time that the elbow reached the target zone. 5. In the second strategy, subjects used kinesthetic input related to the elbow rotation to advance or delay the onset of the hand movement, thereby compensating for leads and lags in the arrival of the elbow at the target zone. These adjustments in the triggering time of the hand movement allowed subjects to open the hand while the elbow was in the target zone. This kinesthetic triggering mechanism was effective for elbow rotations reaching the target zone within 150-1,500 ms. 6. These results suggest that, to fully understand how multijoint movement sequences are controlled by the nervous system, sensory mechanisms must be considered in addition to central mechanisms.

Paul J. Cordo

Papers

Proprioceptive consequences of tendon vibration during movement

Proprioceptive coordination of movement sequences: role of velocity and position information

Kinesthetic control of a multijoint movement sequence

Sensory control of target acquisition

Force and displacement-controlled tendon vibration in humans