To be successful, precision manipulation of small objects requires a refined coordination of forces excerted on the object by the tips of the fingers and thumb. The present paper deals quantitatively with the regulation of the coordination between the grip force and the vertical lifting force, denoted as the load force, while small objects were lifted, positioned in space and replaced by human subjects using the pinch grip. It was shown that the grip force changed in parallel with the load force generated by the subject to overcome various forces counteracting the intended manipulation. The balance between the two forces was adapted to the friction between the skin and the object providing a relatively small safety margin to prevent slips, i.e. the more slippery the object the higher the grip force at any given load force. Experiments with local anaesthesia indicated that this adaptation was dependent on cutaneous afferent input. Afferent information related to the frictional condition could influence the force coordination already about 0.1 s after the object was initially gripped, i.e. approximately at the time the grip and load forces began to increase in parallel. Further, “secondary”, adjustments of the force balance could occur later in response to small short-lasting slips, revealed as vibrations in the object. The new force balance following slips was maintained, indicating that the relationship between the two forces was set on the basis of a memory trace. Its updating was most likely accounted for by tactile afferent information entering intermittently at inappropriate force coordination, e.g. as during slips. The latencies between the onset of such slips and the appearance of the adjustments (0.06–0.08 s) clearly indicated that the underlying neural mechanisms operated highly automatically.

Roles of glabrous skin receptors and sensorimotor memory in automatic control of precision grip when lifting rougher or more slippery objects

The equilibrium control hypothesis (λ model) is considered with special reference to the following concepts: (a) the length-force invariant characteristic (IC) of the muscle together with central and reflex systems subserving its activity; (b) the tonic stretch reflex threshold (λ) as an independent measure of central commands descending to alpha and gamma motoneurons; (c) the equilibrium point, defined in terms of λ, IC and static load characteristics, which is associated with the notion that posture and movement are controlled by a single mechanism; and (d) the muscle activation area (a reformulation of the “size principle”)— the area of kinematic and command variables in which a rank-ordered recruitment of motor units takes place. The model is used for the interpretation of various motor phenomena, particularly electromyographic patterns. The stretch reflex in the λ model has no mechanism to follow-up a certain muscle length prescribed by central commands. Rather, its task is to bring the system to an ...

Once More on the Equilibrium-Point Hypothesis (λ Model) for Motor Control

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

This three-part paper presents an approach to the control of dynamic interaction between a manipulator and its environment. Part I presented the theoretical reasoning behind impedance control. In Part II the implementation of impedance control is considered. A feedback control algorithm for imposing a desired cartesian impedance on the end-point of a nonlinear manipulator is presented. This algorithm completely eliminates the need to solve the \"inverse kinematics problem\" in robot motion control. The modulation of end-point impedance without using feedback control is also considered, and it is shown that apparently \"redundant\" actuators and degrees of freedom such vs exist in the primate musculoskeletal system may be used to modulate end-point impedance and may play an essential functional role in the control of dynamic interaction.

Impedance Control: An Approach to Manipulation: Part II—Implementation

This paper examines the postulate that an important function of the activity of antagonist muscle groups is to modulate mechanical impedance. Some biomechanical modeling and analyses are presented leading to a prediction of simultaneous activation of antagonist muscles in the maintenance of upright posture of the forearm and hand. An experimental observation of antagonist coactivation in this situation is presented.

Adaptive control of mechanical impedance by coactivation of antagonist muscles

1. The stretch reflex in the elbow flexor musculature was studied in 23 human subjects. The subjects were required to establish an initial force equivalent to 10% maximum at a prescribed initial length; mechanical disturbances delivered at random times increased load force to 15% or reduced it to 5%. We measured arm force, displacement, and EMG (usually biceps); acceleration was calculated from displacement, and average responses from sets of 10 like trials. 2. Modification of the stretch reflex was studied by comparing average responses obtained with different instructions, but with the same disturbance. The usual introductions were "compensate for arm deflection" and "do not intervene voluntarily". The initial response did not depend on instruction; changes in response that depended on instruction began abruptly after a latent period which ranged from 70 to 320 ms (measured from force and acceleration), depending on conditions and subject. The latency became longer (10-50 ms) and more variable when the subject did not know the direction of disturbance in advance. This and other observations indicate that modifications of the stretch reflex are not produced by servo actions. They are produced by triggered reactions, which occur at both short and long latencies and which have properties resembling the movements produced in a reaction-time task. 3. We confirmed that most subjects can suppress triggered reactions when the instruction calls for no intervention, leaving an unmodified reflex response. This response consists of a compliant deflection of the arm in the direction of the disturbance. 4. The compensatory actions associated with unmodified stretch (and unloading) reflexes were assessed from EMG responses of biceps. During a 300-ms transient phase, EMG changes were notably asymmetric when responses to symmetric disturbances were compared. Increased force stretched biceps and produced a prominent increase in EMG, whereas decreased force allowed biceps to shorten and produced either an EMG decrease of smaller magnitude or an actual increase. These asymmetric reflex actions produced quite symmetric mechanical responses (arm displacements and forces), which implies the existence of and compensation for nonlinear muscle mechanical properties. This result is discussed in relation to the hypothesis that the function of the stretch reflex is to compensate for variations in muscle properties, thus maintaining stiffness. 5. Effective control of muscle length or joint position does not result from servo action by the stretch reflex. Errors in position are corrected only when triggered reactions are superimposed on the reflex response.

J. C. Houk

Papers

Regulatory actions of human stretch reflex

Dependence of dynamic response of spindle receptors on muscle length and velocity.

Sampling of total muscle force by tendon organs.