Actual and Imagined Movements Reveal a Dual Role of the Insular Cortex for Motor Control.

Movements rely on a mixture of feedforward and feedback mechanisms. With experience, the brain builds internal representations of actions in different contexts. Many factors are taken into account in this process among which is the immutable presence of gravity. Any displacement of a massive body in the gravitational field generates forces and torques that must be predicted and compensated by appropriate motor commands. The insular cortex is a key brain area for graviception. However, no attempt has been made to address whether the same internal representation of gravity is shared between feedforward and feedback mechanisms. Here, participants either mentally simulated (only feedforward) or performed (feedforward and feedback) vertical movements of the hand. We found that the posterior part of the insular cortex was engaged when feedback was processed. The anterior insula, however, was activated only in mental simulation of the action. A psychophysical experiment demonstrates participants' ability to integrate the effects of gravity. Our results point toward a dual internal representation of gravity within the insula. We discuss the conceptual link between these two dualities.

Daily modulation of the speed-accuracy trade-off.

Goal-oriented arm movements are characterized by a balance between speed and accuracy. The relation between speed and accuracy has been formalized by Fitts' law and predicts a linear increase in movement duration with task constraints. Up to now this relation has been investigated on a short-time scale only, that is during a single experimental session, although chronobiological studies report that the motor system is shaped by circadian rhythms. Here, we examine whether the speed-accuracy trade-off could vary during the day. Healthy adults carried out arm-pointing movements as accurately and fast as possible toward targets of different sizes at various hours of the day, and variations in Fitts' law parameters were scrutinized. To investigate whether the potential modulation of the speed-accuracy trade-off has peripheral and/or central origins, a motor imagery paradigm was used as well. Results indicated a daily (circadian-like) variation for the durations of both executed and mentally simulated movements, in strictly controlled accuracy conditions. While Fitts' law was held for the whole sessions of the day, the slope of the relation between movement duration and task difficulty expressed a clear modulation, with the lowest values in the afternoon. This variation of the speed-accuracy trade-off in executed and mental movements suggests that, beyond execution parameters, motor planning mechanisms are modulated during the day. Daily update of forward models is discussed as a potential mechanism.

Moving sounds within the peripersonal space modulate the motor system.

Interactions between ourselves and the external world are mediated by a multisensory representation of the space surrounding the body, i.e. the peripersonal space (PPS). In particular, a special interplay is observed among tactile stimuli delivered on a body part, e.g. the hand, and visual or auditory external inputs presented close, but not far, from the same body part, e.g. within hand PPS. This coding of multisensory stimuli as a function of their distance from the hand has a role in upper limb actions. However, it remains unclear whether PPS representation affects the motor system only when stimuli occur specifically at the hand location or when they move within a continuous portion of space where the hand can potentially act. Here, in order to study these two alternatively hypotheses, we assessed the critical distance at which moving sounds have a direct effect on hand corticospinal excitability by using Transcranial Magnetic Stimulation (TMS). Specifically, TMS single pulses were delivered when a sound source was perceived at six different positions in space: from very close to subjects' hand (15 cm) to far away (90 cm). Moreover, sound direction was manipulated to test if stimuli approaching and receding from the hand might have the same relevance for the motor system. MEPs amplitude was enhanced when sounds were delivered within a limited distance from the hand (around 60 cm) as compared to when the sounds were beyond this space. This effect captures the spatial boundaries within which PPS representation modulates hand cortico-motor excitability. This spatially-dependent modulation of corticospinal activity was not further affected by the sound direction. Such findings support a strict link between the multisensory representation of the space around the body and the motor representation of potential approaching or defensive acts within that space.

Motor imagery in unipolar major depression.

BACKGROUND: Motor imagery is a potential tool to investigate action representation, as it can provide insights into the processes of action planning and preparation. Recent studies suggest that depressed patients present specific impairment in mental rotation. The present study was designed to investigate the influence of unipolar depression on motor imagery ability. METHODS: Fourteen right-handed patients meeting DSM-IV criteria for unipolar depression were compared to 14 matched healthy controls. Imagery ability was accessed by the timing correspondence between executed and imagined movements during a pointing task, involving strong spatiotemporal constraints (speed/accuracy trade-off paradigm). RESULTS: Compared to controls, depressed patients showed marked motor slowing on both actual and imagined movements. Furthermore, we observed greater temporal discrepancies between actual and mental movements in depressed patients than in healthy controls. Lastly, depressed patients modulated, to some extent, mental movement durations according to the difficulty of the task, but this modulation was not as strong as that of healthy subjects. CONCLUSION: These results suggest that unipolar depression significantly affects the higher stages of action planning and point out a selective decline of motor prediction.

Training the motor cortex by observing the actions of others during immobilization.

Limb immobilization and nonuse are well-known causes of corticomotor depression. While physical training can drive the recovery from nonuse-dependent corticomotor effects, it remains unclear if it is possible to gain access to motor cortex in alternative ways, such as through motor imagery (MI) or action observation (AO). Transcranial magnetic stimulation was used to study the excitability of the hand left motor cortex in normal subjects immediately before and after 10 h of right arm immobilization. During immobilization, subjects were requested either to imagine to act with their constrained limb or to observe hand actions performed by other individuals. A third group of control subjects watched a nature documentary presented on a computer screen. Hand corticomotor maps and recruitment curves reliably showed that AO, but not MI, prevented the corticomotor depression induced by immobilization. Our results demonstrate the existence of a visuomotor mechanism in humans that links AO and execution which is able to effect cortical plasticity in a beneficial way. This facilitation was not related to the action simulation, because it was not induced by explicit MI.

Prism adaptation by mental practice.

The prediction of our actions and their interaction with the external environment is critical for sensorimotor adaptation. For instance, during prism exposure, which deviates laterally our visual field, we progressively correct movement errors by combining sensory feedback with forward model sensory predictions. However, very often we project our actions to the external environment without physically interacting with it (e.g., mental actions). An intriguing question is whether adaptation will occur if we imagine, instead of executing, an arm movement while wearing prisms. Here, we investigated prism adaptation during mental actions. In the first experiment, participants (n = 54) performed arm pointing movements before and after exposure to the optical device. They were equally divided into six groups according to prism exposure: Prisms-Active, Prisms-Imagery, Prisms-Stationary, Prisms-Stationary-Attention, No Conflict-Prisms-Imagery, No Prisms-Imagery. Adaptation, measured by the difference in pointing errors between pre-test and post-test, occurred only in Prisms-Active and Prisms-Imagery conditions. The second experiment confirmed the results of the first experiment and further showed that sensorimotor adaptation was mainly due to proprioceptive realignment in both Prisms-Active (n = 10) and Prisms-Imagery (n = 10) groups. In both experiments adaptation was greater following actual than imagined pointing movements. The present results are the first demonstration of prism adaptation by mental practice under prism exposure and they are discussed in terms of internal forward models and sensorimotor plasticity.

The relation between geometry and time in mental actions.

Mental imagery is a cognitive tool that helps humans take decisions by simulating past and future events. The hypothesis has been advanced that there is a functional equivalence between actual and mental movements. Yet, we do not know whether there are any limitations to its validity even in terms of some fundamental features of actual movements, such as the relationship between space and time. Although it is impossible to directly measure the spatiotemporal features of mental actions, an indirect investigation can be conducted by taking advantage of the constraints existing in planar drawing movements and described by the two-thirds power law (2/3PL). This kinematic law describes one of the most impressive regularities observed in biological movements: movement speed decreases when curvature increases. Here, we compared the duration of identical actual and mental arm movements by changing the constraints imposed by the 2/3PL. In the first two experiments, the length of the trajectory remained constant, while its curvature (Experiment 1) or its number of inflexions (Experiment 2) was manipulated. The results showed that curvature, but not the number of inflexions, proportionally and similarly affected actual and mental movement duration, as expected from the 2/3PL. Two other control experiments confirmed that the results of Experiment 1 were not attributable to eye movements (Experiment 3) or to the perceived length of the displayed trajectory (Experiment 4). Altogether, our findings suggest that mental movement simulation is tuned to the kinematic laws characterizing actions and that kinematics of actual and mental movements is completely specified by the representation of their geometry.

Aging affects the mental simulation/planning of the "rising from the floor" sequence.

We investigated the effect of aging on the ability to mentally simulate/plan a complex sequential action of the whole body, namely "rising from the floor". Forty-four non-demented elderly people (mean age: 85.2±5.5 years) and 20 young people (mean age: 26.6±4.9 years) were included in the study. They were required to put in order six images representing the main movements necessary to get up from a sitting position on the floor. We showed that older subjects had poorer performance-both in terms of proportion of success and response time-than their younger counterparts. These results are in line with previous findings showing age-related alterations in action simulation/action planning processes, and highlight the fact that elderly people have particular difficulties when the action to mentally simulate is complex.

Aging affects the mental rotation of left and right hands.

BACKGROUND: Normal aging significantly influences motor and cognitive performance. Little is known about age-related changes in action simulation. Here, we investigated the influence of aging on implicit motor imagery. METHODOLOGY/PRINCIPAL FINDINGS: Twenty young (mean age: 23.9+/-2.8 years) and nineteen elderly (mean age: 78.3+/-4.5 years) subjects, all right-handed, were required to determine the laterality of hands presented in various positions. To do so, they mentally rotated their hands to match them with the hand-stimuli. We showed that: (1) elderly subjects were affected in their ability to implicitly simulate movements of the upper limbs, especially those requiring the largest amplitude of displacement and/or with strong biomechanical constraints; (2) this decline was greater for movements of the non-dominant arm than of the dominant arm. CONCLUSIONS/SIGNIFICANCE: These results extend recent findings showing age-related alterations of the explicit side of motor imagery. They suggest that a general decline in action simulation occurs with normal aging, in particular for the non-dominant side of the body.

Imagined and actual arm movements have similar durations when performed under different conditions of direction and mass.

Several experiments have suggested that similar physiological substrates are involved in movement execution and motor imagery, and that the same laws of movement control apply to both processes. Using a mental chronometry paradigm, we examined the effects of movement direction and added mass on the duration of actual and imagined movements. Six subjects executed or imagined arm movements in the sagittal and horizontal plane, in three different loading conditions: without added mass, and with an added mass of 1 and 1.5 kg. The duration of both actual and imagined movements was measured by an electronic stopwatch. The actual movements were significantly increased in duration as a function of mass, for both movement directions. However, direction per se had no effect on duration. The duration of imagined movements was very similar to that of actual movements whatever the subject and mass and direction condition. These results show that both inertial and gravitational constraints are accurately incorporated in the timing of the motor imagery process, which appears therefore to be functionally very close to the process of planning and performing the actual movement.