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      Inactivation of Cerebellar Cortical Crus II Disrupts Temporal Processing of Absolute Timing but not Relative Timing in Voluntary Movements

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          Abstract

          Several recent studies have demonstrated that the cerebellum plays an important role in temporal processing at the scale of milliseconds. However, it is not clear whether intrinsic cerebellar function involves the temporal processing of discrete or continuous events. Temporal processing during discrete events functions by counting absolute time like a stopwatch, while during continuous events it measures events at intervals. During the temporal processing of continuous events, animals might respond to rhythmic timing of sequential responses rather than to the absolute durations of intervals. Here, we tested the contribution of the cerebellar cortex to temporal processing of absolute and relative timings in voluntary movements. We injected muscimol and baclofen to a part of the cerebellar cortex of rats. We then tested the accuracy of their absolute or relative timing prediction using two timing tasks requiring almost identical reaching movements. Inactivation of the cerebellar cortex disrupted accurate temporal prediction in the absolute timing task. The rats formed two groups based on the changes to their timing accuracy following one of two distinct patterns which can be described as longer or shorter declines in the accuracy of learned intervals. However, a part of the cerebellar cortical inactivation did not affect the rats’ performance of relative timing tasks. We concluded that a part of the cerebellar cortex, Crus II, contributes to the accurate temporal prediction of absolute timing and that the entire cerebellar cortex may be unnecessary in cases in which accurately knowing the absolute duration of an interval is not required for temporal prediction.

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          Most cited references31

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          Control of mental activities by internal models in the cerebellum.

          Masao ITO (2008)
          The intricate neuronal circuitry of the cerebellum is thought to encode internal models that reproduce the dynamic properties of body parts. These models are essential for controlling the movement of these body parts: they allow the brain to precisely control the movement without the need for sensory feedback. It is thought that the cerebellum might also encode internal models that reproduce the essential properties of mental representations in the cerebral cortex. This hypothesis suggests a possible mechanism by which intuition and implicit thought might function and explains some of the symptoms that are exhibited by psychiatric patients. This article examines the conceptual bases and experimental evidence for this hypothesis.
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            Timing functions of the cerebellum.

            This study investigated the effects of different types of neurological deficits on timing functions. The performance of Parkinson, cerebellar, cortical, and peripheral neuropathy patients was compared to age-matched control subjects on two separate measures of timing functions. The first task involved the production of timed intervals in which the subjects attempted to maintain a simple rhythm. The second task measured the subjects' perceptual ability to discriminate between small differences in the duration of two intervals. The primacy of the cerebellum in timing functions was demonstrated by the finding that these were the only patients who showed a deficit in both the production and perception of timing tasks. The cerebellar group was found to have increased variability in performing rhythmic tapping and they were less accurate than the other groups in making perceptual discriminations regarding small differences in duration. Critically, this perceptual deficit appears to be specific to the perception of time since the cerebellar patients were unaffected in a control task measuring the perception of loudness. It is argued that the operation of a timing mechanism can be conceptualized as an isolable component of the motor control system. Furthermore, the results suggest that the domain of the cerebellar timing process is not limited to the motor system, but is employed by other perceptual and cognitive systems when temporally predictive computations are needed.
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              The evolution of brain activation during temporal processing.

              Timing is crucial to many aspects of human performance. To better understand its neural underpinnings, we used event-related fMRI to examine the time course of activation associated with different components of a time perception task. We distinguished systems associated with encoding time intervals from those related to comparing intervals and implementing a response. Activation in the basal ganglia occurred early, and was uniquely associated with encoding time intervals, whereas cerebellar activation unfolded late, suggesting an involvement in processes other than explicit timing. Early cortical activation associated with encoding of time intervals was observed in the right inferior parietal cortex and bilateral premotor cortex, implicating these systems in attention and temporary maintenance of intervals. Late activation in the right dorsolateral prefrontal cortex emerged during comparison of time intervals. Our results illustrate a dynamic network of cortical-subcortical activation associated with different components of temporal information processing.
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                Author and article information

                Contributors
                Journal
                Front Syst Neurosci
                Front Syst Neurosci
                Front. Syst. Neurosci.
                Frontiers in Systems Neuroscience
                Frontiers Media S.A.
                1662-5137
                24 February 2016
                2016
                : 10
                : 16
                Affiliations
                [1] 1Department of Psychology, Graduate School of Letters, Kyoto University Kyoto, Japan
                [2] 2Japan Society for the Promotion of Science Tokyo, Japan
                [3] 3Laboratory of Neural Information, Graduate School of Brain Science, Doshisha University Kyotanabe, Japan
                Author notes

                Edited by: Detlef H. Heck, University of Tennessee Health Science Center, USA

                Reviewed by: Fahad Sultan, University of Tübingen, Germany; Krystal Lynn Parker, University of Iowa, USA

                *Correspondence: Kenji Yamaguchi, yamaguchi.kenji.38u@ 123456kyoto-u.jp
                Article
                10.3389/fnsys.2016.00016
                4764692
                26941621
                7f8d93ac-835b-441a-8680-59a20a60b30b
                Copyright © 2016 Yamaguchi and Sakurai.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 15 November 2015
                : 08 February 2016
                Page count
                Figures: 7, Tables: 2, Equations: 0, References: 37, Pages: 12, Words: 0
                Categories
                Neuroscience
                Original Research

                Neurosciences
                temporal processing,cerebellum,pharmacological inactivation,voluntary movement,rats
                Neurosciences
                temporal processing, cerebellum, pharmacological inactivation, voluntary movement, rats

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