Two general frameworks have been articulated to describe how the passage of time is perceived. One emphasizes that the judgment of the duration of a stimulus depends on the operation of dedicated neural mechanisms specialized for representing the temporal relationships between events. Alternatively, the representation of duration could be ubiquitous, arising from the intrinsic dynamics of nondedicated neural mechanisms. In such models, duration might be encoded directly through the amount of activation of sensory processes or as spatial patterns of activity in a network of neurons. Although intrinsic models are neurally plausible, we highlight several issues that must be addressed before we dispense with models of duration perception that are based on dedicated processes.

The cerebellum is often active in imaging studies of verbal working memory, consistent with a putative role in articulatory rehearsal. While patients with cerebellar damage occasionally exhibit a mild impairment on standard neuropsychological tests of working memory, these tests are not diagnostic for exploring these processes in detail. The current study was designed to determine whether damage to the cerebellum is associated with impairments on a range of verbal working memory tasks, and if so, under what circumstances. Moreover, we assessed the hypothesis that these impairments are related to impaired rehearsal mechanisms. Patients with damage to the cerebellum (n = 15) exhibited a selective deficit in verbal working memory: spatial forward and backward spans were normal, but forward and backward verbal spans were lower than controls. While the differences were significant, digit spans were relatively preserved, especially in comparison to the dramatic reductions typically observed in classic 'short-term memory' patients with perisylvian brain damage. The patients tended to be more impaired on a verbal version compared to a spatial version of a working memory task with a long delay and this impairment was correlated with overall symptom and dysarthria severity. These results are consistent with a contribution of the cerebellum to rehearsal and suggest that inclusion of a delay before recall is especially detrimental in individuals with cerebellar damage. However, when we examined markers of rehearsal (i.e. word-length and articulatory suppression effects) in an immediate serial recall task, we found that qualitative aspects of the patients' rehearsal strategies were unaffected. We propose that the cerebellum may contribute to verbal working memory during the initial phonological encoding and/or by strengthening memory traces rather than by fundamentally subserving covert articulatory rehearsal.

Schlerf JE, Verstynen TD, Ivry RB, Spencer RM. Evidence of a novel somatopic map in the human neocerebellum during complex actions. J Neurophysiol 103: 3330 -3336, 2010. First published April 14, 2010 doi:10.1152/jn.01117.2009. The human neocerebellum has been hypothesized to contribute to many high-level cognitive processes including attention, language, and working memory. Support for these nonmotor hypotheses comes from evidence demonstrating structural and functional connectivity between the lateral cerebellum and cortical association areas as well as a lack of somatotopy in lobules VI and VII, a hallmark of motor representations in other areas of the cerebellum and cerebral cortex. We set out to test whether somatotopy exists in these lobules by using functional magnetic resonance imaging to measure cerebellar activity while participants produced simple or complex movements, using either fingers or toes. We observed a previously undiscovered somatotopic organization in neocerebellar lobules VI and VIIA that was most prominent when participants executed complex movements. In contrast, activation in the anterior lobe showed a similar somatotopic organization for both simple and complex movements. While the anterior somatotopic representation responded selectively during ipsilateral movements, the new cerebellar map responded during both ipsi-and contralateral movements. The presence of a bilateral, task-dependent somatotopic map in the neocerebellum emphasizes an important role for this region in the control of skilled actions.

A central tenet of motor neuroscience is that the cerebellum learns from sensory prediction errors. Surprisingly, neuroimaging studies have not revealed definitive signatures of error processing in the cerebellum. Furthermore, neurophysiologic studies suggest an asymmetry, such that the cerebellum may encode errors arising from unexpected sensory events, but not errors reflecting the omission of expected stimuli. We conducted an imaging study to compare the cerebellar response to these two types of errors. Participants made fast out-and-back reaching movements, aiming either for an object that delivered a force pulse if intersected or for a gap between two objects, either of which delivered a force pulse if intersected. Errors (missing the target) could therefore be signaled either through the presence or absence of a force pulse. In an initial analysis, the cerebellar BOLD response was smaller on trials with errors compared with trials without errors. However, we also observed an error-related decrease in heart rate. After correcting for variation in heart rate, increased activation during error trials was observed in the hand area of lobules V and VI. This effect was similar for the two error types. The results provide evidence for the encoding of errors resulting from either the unexpected presence or unexpected absence of sensory stimulation in the human cerebellum.

The cerebellum is critically important for error driven adaptive motor learning, as evidenced by the fact that cerebellar patients do not adapt well to sudden predictable perturbations. However, recent work has shown that cerebellar patients adapt much better if the perturbation is gradually introduced. Here we explore physiological mechanisms that underlie this distinction between abrupt and gradual motor adaptation in humans. We used Transcranial Magnetic Stimulation (TMS) to evaluate whether neural mechanisms within the cerebellum contribute to either process during a visuomotor reach adaptation. When a visuomotor rotation was introduced abruptly, cerebellar excitability changed early in learning, and approached baseline levels near the end of the adaptation block. However, we observed no modulation of cerebellar excitability when we presented the visuomotor rotation gradually during learning. Similarly, we did not observe cerebellar modulation during trial-by-trial adaptation to random visuomotor displacements or during reaches without perturbations. This suggests that the cerebellum is most active during the early-phases of adaptation when large perturbations are successfully compensated.

The cerebellum has long been recognized to play an important role in motor adaptation. Individuals with cerebellar ataxia exhibit impaired learning in visuomotor adaptation tasks such as prism adaptation and force field learning. Both types of tasks involve the adjustment of an internal model to compensate for an external perturbation. This updating process is error driven, with the error signal based on the difference between anticipated and actual sensory information. This process may entail a credit assignment problem, with a distinction made between error arising from faulty representation of the environment and error arising from noise in the controller. We hypothesized that people with ataxia may perform poorly at visuomotor adaptation because they attribute a greater proportion of their error to their motor control difficulties. We tested this hypothesis using a computational model based on a Kalman filter. We imposed a 20-deg visuomotor rotation in either a single large step or in a series of smaller 5-deg steps. The ataxic group exhibited a comparable deficit in both conditions. The computational analyses indicate that the patients' deficit cannot be accounted for simply by their increased motor variability. Rather, the patients' deficit in learning may be related to difficulty in estimating the instability in the environment or variability in their motor system.

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Although these results clearly show the limitations of current approaches, they also show the potential of anatomical and functional MRI for the study of the human cerebellum.