Considerable progress has been made in developing models of cerebellar function in sensorimotor control, as well as in identifying key problems that are the focus of current investigation. In this consensus paper, we discuss the literature on the role of the cerebellar circuitry in motor control, bringing together a range of different viewpoints. The following topics are covered: oculomotor control, classical conditioning (evidence in animals and in humans), cerebellar control of motor speech, control of grip forces, control of voluntary limb movements, timing, sensorimotor synchronization, control of corticomotor excitability, control of movement-related sensory data acquisition, cerebro-cerebellar interaction in visuokinesthetic perception of hand movement, functional neuroimaging studies, and magnetoencephalographic mapping of cortico-cerebellar dynamics. While the field has yet to reach a consensus on the precise role played by the cerebellum in movement control, the literature has witnessed the emergence of broad proposals that address cerebellar function at multiple levels of analysis. This paper highlights the diversity of current opinion, providing a framework for debate and discussion on the role of this quintessential vertebrate structure.

We investigate whether imagery of voluntary movements of different body parts activates somatotopical sections of the human motor cortices. We used functional magnetic resonance imaging to detect the cortical activity when 7 healthy subjects imagine performing repetitive (0.5-Hz) flexion/extension movements of the right fingers or right toes, or horizontal movements of the tongue. We also collected functional images when the subjects actually executed these movements and used these data to define somatotopical representations in the motor areas. In this study, we relate the functional activation maps to cytoarchitectural population maps of areas 4a, 4p, and 6 in the same standard anatomical space. The important novel findings are 1). that imagery of hand movements specifically activates the hand sections of the contralateral primary motor cortex (area 4a) and the contralateral dorsal premotor cortex (area 6) and a hand representation located in the caudal cingulate motor area and the most ventral part of the supplementary motor area; 2). that when imagining making foot movements, the foot zones of the posterior part of the contralateral supplementary motor area (area 6) and the contralateral primary motor cortex (area 4a) are active; and 3). that imagery of tongue movements activates the tongue region of the primary motor cortex and the premotor cortex bilaterally (areas 4a, 4p, and 6). These results demonstrate that imagery of action engages the somatotopically organized sections of the primary motor cortex in a systematic manner as well as activating some body-part-specific representations in the nonprimary motor areas. Thus the content of the mental motor image, in this case the body part, is reflected in the pattern of motor cortical activation.

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We have previously shown that motor areas are engaged when subjects experience illusory limb movements elicited by tendon vibration. However, traditionally cytoarchitectonic area 2 is held responsible for kinesthesia. Here we use functional magnetic resonance imaging and cytoarchitectural mapping to examine whether area 2 is engaged in kinesthesia, whether it is engaged bilaterally because area 2 in non-human primates has strong callosal connections, which other areas are active members of the network for kinesthesia, and if there is a dominance for the right hemisphere in kinesthesia as has been suggested. Ten right-handed blindfolded healthy subjects participated. The tendon of the extensor carpi ulnaris muscles of the right or left hand was vibrated at 80 Hz, which elicited illusory palmar flexion in an immobile hand (illusion). As control we applied identical stimuli to the skin over the processus styloideus ulnae, which did not elicit any illusions (vibration). We found robust activations in cortical motor areas [areas 4a, 4p, 6; dorsal premotor cortex (PMD) and bilateral supplementary motor area (SMA)] and ipsilateral cerebellum during kinesthetic illusions (illusion-vibration). The illusions also activated contralateral area 2 and right area 2 was active in common irrespective of illusions of right or left hand. Right areas 44, 45, anterior part of intraparietal region (IP1) and caudo-lateral part of parietal opercular region (OP1), cortex rostral to PMD, anterior insula and superior temporal gyrus were also activated in common during illusions of right or left hand. These right-sided areas were significantly more activated than the corresponding areas in the left hemisphere. The present data, together with our previous results, suggest that human kinesthesia is associated with a network of active brain areas that consists of motor areas, cerebellum, and the right fronto-parietal areas including high-order somatosensory areas. Furthermore, our results provide evidence for a right hemisphere dominance for perception of limb movement.