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The role of corticospinal (CS) pathways in anticipatory motor actions was evaluated using transcranial magnetic stimulation (TMS) of the primary motor cortex projecting to motoneurons (MNs) of wrist muscles. Preloaded wrist flexors were suddenly unloaded by the experimenter or by the subject using the other hand (self-unloading). After sudden unloading, the wrist joint involuntarily flexed to a new position. In contrast, during self-unloading the wrist remained almost motionless, implying that an anticipatory postural adjustment occurred. In the self-unloading task, anticipation was manifested by a decrease in descending facilitation of pre-activated flexor MNs starting ∼72 ms before changes in the background EMG activity. Descending facilitation of extensor MNs began to increase ∼61 ms later. Conversely, these influences remained unchanged before sudden unloading, implying the absence of anticipation. We also tested TMS responses during EMG silent periods produced by brief muscle shortening, transiently resulting in similar EMG levels before the onset and after the end of self-unloading. We found reduced descending facilitation of flexor MNs after self-unloading. To explain why the wrist excursion was minimized in self-unloading due to these changes in descending influences, we relied on previous demonstrations that descending systems pre-set the threshold positions of body segments at which muscles begin to be activated, thus pre-determining the equilibrium point to which the system is attracted. Based on this notion, a more consistent explanation of the kinematic, EMG and descending patterns in the two types of unloading is proposed compared to the alternative notion of direct pre-programming of kinematic and/or EMG patterns. This article is protected by copyright. All rights reserved

Advances to the equilibrium-point (EP) theory and solutions to several classical problems of action and perception are suggested and discussed. Among them are (1) the posture-movement problem of how movements away from a stable posture can be made without evoking resistance of posture-stabilizing mechanisms resulting from intrinsic muscle and reflex properties; (2) the problem of kinesthesia or why our sense of limb position is fairly accurate despite ambiguous positional information delivered by proprioceptive and cutaneous signals; (3) the redundancy problems in the control of multiple muscles and degrees of freedom. Central to the EP hypothesis is the notion that there are specific neural structures that represent spatial frames of reference (FRs) selected by the brain in a task-specific way from a set of available FRs. The brain is also able to translate or/and rotate the selected FRs by modifying their major attributes-the origin, metrics, and orientation-and thus substantially influence, in a feed-forward manner, action and perception. The brain does not directly solve redundancy problems: it only limits the amount of redundancy by predetermining where, in spatial coordinates, a task-specific action should emerge and allows all motor elements, including the environment, to interact to deliver a unique action, thus solving the redundancy problem (natural selection of action). The EP theory predicts the existence of specific neurons associated with the control of different attributes of FRs and explains the role of mirror neurons in the inferior frontal gyrus and place cells in the hippocampus. WIREs Cogni Sci 2011 2 287-304 DOI: 10.1002/wcs.108 For further resources related to this article, please visit the WIREs website.

It has previously been established that muscles become active in response to deviations from a threshold (referent) position of the body or its segments, and that intentional motor actions result from central shifts in the referent position. We tested the hypothesis that corticospinal pathways are involved in threshold position control during intentional changes in the wrist position in humans. Subjects moved the wrist from an initial extended to a final flexed position (and vice versa). Passive wrist muscle forces were compensated with a torque motor such that wrist muscle activity was equalized at the two positions. It appeared that motoneuronal excitability tested by brief muscle stretches was also similar at these positions. Responses to mechanical perturbations before and after movement showed that the wrist threshold position was reset when voluntary changes in the joint angle were made. Although the excitability of motoneurons was similar at the two positions, the same transcranial magnetic stimulus (TMS) elicited a wrist extensor jerk in the extension position and a flexor jerk in the flexion position. Extensor motor-evoked potentials (MEPs) elicited by TMS at the wrist extension position were substantially bigger compared to those at the flexion position and vice versa for flexor MEPs. MEPs were substantially reduced when subjects fully relaxed wrist muscles and the wrist was held passively in each position. Results suggest that the corticospinal pathway, possibly with other descending pathways, participates in threshold position control, a process that pre-determines the spatial frame of reference in which the neuromuscular periphery is constrained to work. This control strategy would underlie not only intentional changes in the joint position, but also muscle relaxation. The notion that the motor cortex may control motor actions by shifting spatial frames of reference opens a new avenue in the analysis and understanding of brain function.

The role of the basal ganglia in the adaptive control of movement was investigated by unexpectedly perturbing movements in 8 patients with Parkinson's disease (PD) tested off medication and in 6 aged-matched healthy subjects. Subjects performed two movement components simultaneously and without visual feedback: touching the nose with the finger while leaning the trunk forward. Subjects wore a harness connected to an electromagnet, which was attached to a wall. The trunk movement was mechanically blocked in randomly selected trials by engaging the electromagnet. While healthy subjects performed the task equally well in both conditions, PD subjects' hand movements significantly deteriorated in trunk-perturbed compared to trunk-free trials. Deteriorated hand movements were characterized by segmented hand paths, unsmooth velocity profiles, and prolonged movement times. This finding indicated that the relatively local trunk perturbation had a global effect on the hand movement of PD subjects, necessitating them to reinitiate, after some delay, their arm movement in perturbed trials. Thus, the basal ganglia may be a critical node in brain networks mediating the flexibility of responses to altered motor states.

An electromyographical study has been made of carp swimming muscles at various points along the body length during sustained and burst locomotion. During sustained swimming, red muscles show a constant time lag between activation of anterior and posterior segments which is practically independent of the speed of locomotion. In contrast, anterior and posterior homolateral segments are activated simultaneously during bursts of rapid movement. This pattern of co-ordination between body segments differs from that reported for other species.