Our sensorimotor experience unfolds in sequences over time. We hypothesize that the processing of movement sequences with and without a temporal pause will recruit distinct but cooperating neural processes, including cortico-striatal and cortico-cerebellar networks. We thus compare neural activity during sequence learning in the presence and absence of this pause. Young volunteer participants learned sensorimotor sequences using the discrete sequence production (DSP) task, with Pause, No-Pause and Control sequences of four elements in an event related fMRI protocol. The No-Pause and (...) Pause sequences involved a more complex sequential structure than the Control sequence, while the Pause sequences involved insertion of a temporal pause, relative to the No-Pause sequence. The Pause vs. No-Pause contrast revealed extensive fronto-parietal, striatal, thalamic and cerebellar activations, preferentially for the Pause sequences. ROI analysis indicated that the cerebellum displays an early activation that was attenuated over successive runs, and a significant preference for Pause sequences when compared with caudate. These data support the hypothesis that a cortico-cerebellar circuit plays a specific role in the initial processing of temporal structure, while the basal ganglia play a more general role in acquiring the serial response order of the sequence. (shrink)
Brindley proposed that we initially generate movements , under higher cerebral control. As the movement is practiced, the cerebellum learns to link within itself the context in which the movement is made to the lower level movement generators. Marr and Albus proposed that the linkage is established by a special input from the inferior olive, which plays upon an input-output element within the cerebellum during the period of the learning. When the linkage is complete, the occurrence of the context (represented (...) by a certain input to the cerebellum) will trigger (through the cerebellum) the appropriate motor response. The movement is distinguished from the conscious movement by its now being automatic, rapid, and stereotyped. The idea is still controversial, but has been supported by a variety of animal studies and, as reviewed here, is consistent with the results of a number of human PET and ablation studies. I have added to the idea of context-response linkage what I think is another important variable: novel combinations of downstream elements. With regard to the motor system and the muscles, this could explain how varied combinations of muscles may become active in precise time-amplitude specifications so as to produce coordinated movements appropriate to specific contexts. In this target article, I have further extended this idea to the premotor parts of the brain and their role in cognition. These areas receive influences from the cerebellum; they are active both in planning movements that are to be executed and in thinking about movements that are not to be executed. From recent evidence, the cerebellar output extends even to what has been characterized as the ultimate frontal planning area, the cortex, area 46. The cerebellum thus may be involved in context-response linkage, and response combination even at these higher levels. The implication would be that, through practice, an experiential context would automatically evoke a certain mental action plan. The plan would be in the realm of thought, and could lead to execution. The specific cerebellar contribution would be one of the context linkage and the shaping of the response, through trial and error learning. The prefrontal and premotor areas could still plan without the help of the cerebellum, but not so automatically, rapidly, stereotypically, so precisely linked to context, or so free of error. Nor would their activities improve optimally with mental practice. (shrink)
In this commentary we discuss a predictive sensorimotor illusion, the size-weight illusion, in which the smaller of two objects of equal weight is perceived as heavier. We suggest that Grush's emulation theory can explain this illusion as a mismatch between predicted and actual sensorimotor feedback, and present preliminary data suggesting that the cerebellum may be critical for implementing the emulator.
Editors’ note: These four interrelated discussions of the role of the cerebellum in coordinating emotional and higher cognitive functions developed out of a workshop presented by the four authors for the 2000 Conference of the Cognitive Science Society at the University of Pennsylvania. The four interrelated discussions explore the implications of the recent explosion of cerebellum research suggesting an expanded cerebellar role in higher cognitive functions as well as in the coordination of emotional functions with learning, logical thinking, perceptual consciousness, (...) and action planning. (shrink)
(1) The is not the only interpretation of cerebellar histology worth considering. Therefore, it is not imperative to strive for a theory of cerebellar function which gives it a prominent rôle. (2) The experiments with cannot support the tidal wave theory. (3) The notion that only can excite the cerebellar cortex is burdened with many intrinsic difficulties. (4) The common theoretical claim that the accuracy of skilled movements is due to exact pattern-matching processes in the cerebellum may be most misleading.
This commentary reviews the basic physical principles underlying human single- and multi-joint arm movements. The potential role of the cerebellum in dealing with the physics of movement is discussed in the light of recent physiological findings and the theoretical model of cerebellar detection and generation of input and output sequences put forward by Braitenberg and colleagues.
Braitenberg et al.'s view that the cerebellum contributes to multijoint sequences of movement is too narrow to account adequately for results from new anatomical, neurobehavioral, and neuroimaging studies. A broader view is that the cerebellum modulates attention, sensory, motor, and other neural systems in order to accomplish its prime function, which is to learn to predict and prepare for imminent information acquisition, analysis, or action.
The role of the cerebellum as a comparator of desired motor output and actual performance may be most important during learning of a novel motor task, when movement errors are common and corrective movements are produced to compensate for them. It is suggested that PET and recent fMRI data are compatible with such an interpretation. Increased activity in motor cortical areas during motor learning indicates that these areas also contribute to the learning process, [THACH].
We argue that the function of the cerebellum is more than just an error-detecting mechanism. Rather, the cerebellum plays an important role in all movements. The bias in (re)calibration is an unfortunate restrictive result of a very successful and important experiment, [SMITH, THACH].
This article reviews models of the cerebellum and motor learning, from the landmark papers by Marr and Albus through those of the present time. The unique architecture of the cerebellar cortex is ideally suited for pattern recognition, but how is pattern recognition incorporated into motor control and learning systems? The present analysis begins with a discussion of exactly what the cerebellar cortex needs to regulate through its anatomically defined projections to premotor networks. Next, we examine various models showing how the (...) microcircuitry in the cerebellar cortex could be used to achieve its regulatory functions. Having thus defined what it is that Purkinje cells in the cerebellar cortex must learn, we then evaluate theories of motor learning. We examine current models of synaptic plasticity, credit assignment, and the generation of training information, indicating how they could function cooperatively to guide the processes of motor learning. (shrink)
We suggest that the cerebellum generates sensory or estimates based on outgoing motor commands and sensory feedback. Thus, it is not a motor pattern generator (HOUK et al.) but a predictive system which is intimately involved in motor behavior. This theory may explain the sensitivity of the climbing fibers to both unexpected external events and motor errors (SIMPSON et al.), and we speculate that unusual biophysical properties of the inferior olive might allow the cerebellum to develop multiple asynchronous sensory estimates, (...) [HOUK et al.; SIMPSON et al.; THACH]. (shrink)
Thach's target article presents a remarkable overview and integration of animal and human studies on the functions of the cerebellum and makes clear theoretical predictions for both the normal operation of the cerebellum and for the effects of cerebellar lesions in the mature human. Commentary is provided on three areas, namely, spatial navigation, implicit learning, and cerebellar agenesis to elicit further development of the themes already present in Thach's paper, [THACH].
In the behavioral literature on human movement, a distinction is made between the learning of parameters and the learning of new movement forms or topologies. Whereas the target articles by Thach, Smith, and Houk et al. provide evidence for cerebellar involvement in parametrization learning and adaptation, the evidence in favor of its involvement in the generation of new movement patterns is less straightforward. A case is made for focusing more attention on the latter issue in the future. This would directly (...) help to bridge the gap between current neurophysiological approaches to the role of the cerebellum and the behavioral expressions of human motor learning, [HOUK et al.; SMITH; THACH]. (shrink)
For reasons I have never understood, some students of the cerebellum have been unwilling to accept the now overwhelming evidence that the cerebellum exhibits lasting synaptic plasticity and plays an essential role in some forms of learning and memory. With a few exceptions (e.g., target article by SIMPSON et al.) this is no longer the case, as is clear in the excellent target articles on cerebellar LTD and the excellent target review by HOUK et al. [CRÉPEL et al.; HOUR et (...) al.; KANO; LINDEN; SIMPSON et al.; SMITH; VINCENT]. (shrink)
PET studies using classical conditioning paradigms are reported. It is emphasized that PET studies show and not in learning paradigms. The importance of dissociating motor performance and learning deficits in human lesions studies is demonstrated in two exemplary studies. The different role of the cerebellum in adaptation of postural reflexes and learning of complex voluntary arm movements is discussed, [THACH].
Interest in the role of nitric oxide (NO) in the nervous system began with the demonstration that glutamate receptor activation in cerebellar slices causes the formation of a diffusible messenger with properties similar to those of the endothelium-derived relaxing factor. It is now clear that this is due to the Ca2+/calmodulin-dependent activation of the enzyme NO synthase, which forms NO and citrulline from the amino acid L-arginine. The cerebellum has very high levels of NO synthase, and although it has low (...) levels of guanylyl cyclase, cerebellar cyclic guanosine monophosphate (cGMP) levels are an order of magnitude higher than in other brain regions. A transcellular metabolic pathway is also present in the cerebellar cortex to recycle citrulline back to arginine. The NO formed binds to and activates soluble guanylyl cyclase to elevate cGMP levels in target cells. Studies employing NADPH-diaphorase, a selective histochemical marker for NO synthase, together with immunohistochemistry, in situ hybridization and biochemical studies have indicated that NO production occurs in granule and basket cells in the cerebellar cortex, whereas cGMP formation appears to occur largely in other cells, including Purkinje cells. Given that a long-term depression of AMPA currents can be seen in isolated Purkinje cells, this anatomical localization suggests that NO cannot play an essential role in the induction of this form of synaptic plasticity. (shrink)
How do we know that our own actions belong to us? How are we able to distinguish self-generated sensory events from those that arise externally? In this paper, I will briefly discuss experiments that were designed to investigate these questions. In particularly, I will review psychophysical and neuroimaging studies that have investigated how we recognise the consequences of our own actions, and why patients with delusions of control confuse self-produced and externally produced actions and sensations. Studies investigating the failure of (...) this 'self-monitoring' mechanism in patients with delusions of control will be discussed in the context of the hypothesis that overactivity in the parietal cortex and the cerebellum contribute to the misattribution of an action to an external source (). (shrink)
This study examines how a Purkinje cell receives its appropriate olivary error signal during the learning of compound movements. We suggest that the Purkinje cell only reinforces those target pyramidal cells which already participate in the movement, subsequently reducing any repeated error signal, such as its own climbing fiber input, [simpson et al.; smith].
The antisaccade task is a classic task of oculomotor control that requires participants to inhibit a saccade to a target and instead make a voluntary saccade to the mirror opposite location. By comparison, the prosaccade task requires participants to make a visually-guided saccade to the target. These tasks have been studied extensively using behavioural oculomotor, electrophysiological and neuroimaging in both non-human primates and humans. In humans, the antisaccade task is under active investigation as a potential endophenotype or biomarker for multiple (...) psychiatric and neurological disorders. A large and growing body of literature has used functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) to study the neural correlates of the antisaccade and prosaccade tasks. We present a quantitative meta-analysis of all published voxel-wise fMRI and PET studies (18) of the antisaccade task and show that consistent activation for antisaccades and prosaccades is obtained in a fronto-subcortical-parietal network encompassing frontal and supplementary eye fields, thalamus, striatum and intraparietal cortex. This network is strongly linked to oculomotor control and was activated to a greater extent for antisaccade than prosaccade trials. Antisaccade but not prosaccade trials additionally activated dorsolateral and ventrolateral prefrontal cortices. We also found that a number of additional regions not classically linked to oculomotor control were activated to a greater extent for antisaccade versus prosaccade trials; these regions are often reported in antisaccade studies but rarely commented upon. While the number of studies eligible to be included in this meta-analysis was small, the results of this systematic review reveal that antisaccade and prosaccade trials consistently activate a distributed network of regions both within and outside the classic definition of the oculomotor network. (shrink)
The cellular basis of motor learning in the cerebellum has been attributed mostly to long-term depression (LTD) at excitatory parallel fiber (PF)-Purkinje cell (PC) synapses. LTD is induced when PFs are activated in conjunction with a climbing fiber (CF), the other excitatory input to PCs. Recently, by using whole-cell patch-clamp recording from PCs in cerebellar slices, a new form of synaptic plasticity was discovered. Stimulation of excitatory CFs induced a long-lasting (usually longer than 30 min) of 30 sec) and the (...) durations of RP (>30 min) strongly suggests that some intracellular biochemical machinery is involved. Pharmacological evidence suggests that protein kinases are involved in RP of inhibitory synapses and LTD of excitatory PF synapses. Besides the well-described LTD, RP could be a cellular mechanism that plays an important role in motor learning. (shrink)
The hypothesis that play behavior is more prevalent in larger-brained animals has recently been challenged. It may be, for example, that only certain brain structures are related to play. Here, we analyze social play behavior with regards to the cerebellum: a structure strongly implicated in motor-development, and possibly also in cognitive skills. We present an evolutionary analysis of social play and the cerebellum, using a phylogenetic comparative method. Social play frequency and relative cerebellum size are positively correlated. Hence, there appears (...) to be a link between the evolutionary elaboration of social play and the cerebellum. (shrink)
The current study investigated behavioral and neural effects of motor, mental, and combined motor and mental training on a finger tapping task. The motor or mental training groups trained on a finger-sequence for a total of 72 min over six weeks. The motor and mental training group received 72 min motor training and in addition 72 min mental training. Results showed that all groups increased their tapping performance significantly on the trained sequence. After training fMRI data was collected and indicated (...) training specific increases in ventral pre-motor cortex following motor training, and in fusiform gyrus following mental training. Combined motor and mental training activated both the motor and the visual regions. In addition, motor and mental training showed a significant increase in tapping performance on an untrained sequence (transfer). FMRI scanning indicated that the transfer effect involved the cerebellum. Conclusions were that combined motor and mental training recruited both motor and visual systems, and that combined motor and mental training improves motor flexibility via connections from both motor and cognitive systems to the cerebellum. (shrink)
Age-related declines in processing speed are hypothesized to underlie the widespread changes in cognition experienced by older adults. We used a structural covariance approach to identify putative neural networks that underlie age-related structural changes associated with processing speed for 42 adults ranging in age from 19-79 years. To characterize a mechanism by which age-related gray matter changes lead to slower processing speed, we examined the extent to which cerebral small vessel disease influenced the association between age-related gray matter changes and (...) processing speed. A frontal pattern of gray matter and white matter variation that was related to cerebral small vessel disease, as well as a cerebellar pattern of gray matter and white matter variation were uniquely related to age-related declines in processing speed. These results demonstrate that at least 2 distinct factors affect age-related changes in processing speed, which might be slowed by mitigating cerebral small vessel disease and factors affecting declines in cerebellar morphology. (shrink)
As the world’s population ages, a deeper understanding of the relationship between aging and motor learning will become increasingly relevant in basic research and applied settings. In this context, this review aims to address the effects of age on motor sequence learning (MSL) and motor adaptation (MA) with respect to behavioral, neurological and neuroimaging findings. Previous behavioral research investigating the influence of aging on motor learning has consistently reported the following results. First, the initial acquisition of motor sequences is not (...) altered, except under conditions of increased task complexity. Second, older adults demonstrate deficits in motor sequence memory consolidation. And, third, although older adults demonstrate deficits during the exposure phase of MA paradigms, the aftereffects following removal of the sensorimotor perturbation are similar to young adults, suggesting that the adaptive ability of older adults is relatively intact. This paper will review the potential neural underpinnings of these behavioral results, with a particular emphasis on the influence of age-related dysfunctions in the cortico-striatal system on motor learning. (shrink)
Symptoms of essential tremor (ET) are similar to those of Parkinson’s disease (PD) during their initial stages. Presently, there are few stable biomarkers available on a neuroanatomical level for distinguishing between these two diseases. However, few investigations have directly compared the changes in brain volume and assessed the compensatory effects of a change in the parts of the brain associated with PD and with ET. To determine the compensatory and/or degenerative anatomical changes in the brains of PD and ET patients, (...) the present study tested, via two VBM approaches (Basic vs. Dartel VBM processing), the anatomical brain images of 10 PD and 10 ET patients, as well as of 13 age-matched normal controls, obtained through a 3T magnetic resonance scanner. These findings indicate that PD and ET caused specific patterns of brain volume alterations in the brains examined. In addition, our observations also revealed compensatory effects, or self-reorganization, occurring in the thalamus and the MTG in the PD and ET patients, due perhaps in part to the enhanced thalamocortical sensorimotor interaction and the head-eye position readjustment, respectively, in these PD and ET patients. Such a distinction may lend itself to use as a biomarker for differentiating between these two diseases. (shrink)
A growing number of studies have reported altered functional connectivity in schizophrenia during putatively “task-free” states and during the performance of cognitive tasks. However, there have been few systematic examinations of functional connectivity in schizophrenia across rest and different task states to assess the degree to which altered functional connectivity reflects a stable characteristic or whether connectivity changes vary as a function of task demands. We assessed functional connectivity during rest and during thee working memory loads of an N-back task (...) (0-back, 1-back, 2-back) among: 1) individuals with schizophrenia (N=19); the siblings of individuals with schizophrenia (N=28); 3) healthy controls (N=10) and the siblings of healthy controls (n=17). We examined connectivity within and between four brain networks: 1) frontal-parietal (FP); 2) cingulo-opercular (CO); 3) cerebellar (CER); and 4) default mode (DMN). We found that connectivity within the DMN and FP increased significantly between resting state and 0-back, while connectivity within the CO and CER decreased significantly between resting state and 0-back. Further, the DMN became significantly more “anti-correlated” with the FP, CO and CER networks during 0-back as compared to rest. Additionally, we found that connectivity within both the DMN and FP was further modulated by memory load, and that connectivity between the FP and both CO and CER networks increased with memory load. Individuals with schizophrenia and their siblings showed consistent reductions in connectivity between both the FP and CO networks with the CER network, a finding that was similar in magnitude across rest and all levels of working memory load. These findings are consistent with the hypothesis that altered functional connectivity in schizophrenia reflects a stable characteristic that is present across cognitive states. (shrink)
Several themes can be identified in the commentaries. The first is that the climbing fibers may have more than one function; the second is that the climbing fibers provide sensory rather than motor signals. We accept the possibility that climbing fibers may have more than one function consequence(s)’ in the title. Until we know more about the function of the inhibitory input to the inferior olive from the cerebellar nuclei, which are motor structures, we have to keep open the possibility (...) that the climbing fiber signals can be a combination of sensory and motor signals. (shrink)
This paper outlines the functional capacities of a novel scheme for cognitive representation and computation, and it explores the possible implementation of this scheme in the massively parallel organization of the empirical brain. The suggestion is that the brain represents reality by means of positions in suitably constitutes phase spaces; and the brain performs computations on these representations by means of coordinate transformations from one phase space to another. This scheme may be implemented in the brain in two distinct forms: (...) (1) as a phase-space sandwich, which may explain certain laminar structures, such as cerebral cortex and the superior colliculus; and (2) as a neural matrix, which may explain other structures, such as the beautifully orthogonal architecture of the cerebellum. (shrink)
Fractal time fluctuations of the spectral “1/f” form are universal in natural self-organizing systems. Neurobiology is uniquely infused with fractal fluctuations in the form of statistically self-similar clusters or bursts on all levels of description from molecular events such as protein chain fluctuations, ion channel currents and synaptic processes to the behaviors of neural ensembles or the collective behavior of Internet users. It is the thesis of this essay that the brain self-organizes via a vertical collation of these spontaneous events (...) in order to perceive the world and generate adaptive behaviors. REM sleep, which coalesces from self-similar clusters of burst-within-burst behavior during ontogeny, is essential to cognitive-emotional function, and has recurrent fractal organization. Empirical fMRI observations further support the association of fractal fluctuations in the temporal lobes, brainstem and cerebellum during the expression of emotional memory, spontaneous fluctuations of thought and meditative practice. Cognitive-emotional integration arises as amygdaloid-brainstem-cerebellar systems harmonize the vertical “1/f” symphony of coupled isochronous cortical oscillations in the pursuit of mindfulness. (shrink)
The human self model suggests that the construct of self involves functions such as agency, body-centered spatial perspectivity, and long-term unity. Vogeley, Kurthen, Falkai, and Maieret (1999) suggest that agency is subserved by the prefrontal cortex and other association areas of the cortex, spatial perspectivity by the prefrontal cortex and the parietal lobes, and long-term unity by the prefrontal cortex and the temporal lobes and that all of these functions are impaired in schizophrenia. Exploring the connections between the prefrontal cortex (...) and the construct of self, the present article extends the application of the self model to autism. It suggests that in contrast to schizophrenia, agency and spatial perspectivity are probably preserved in autism, but that, similarly to schizophrenia, long-term unity is probably impaired. This hypothesis is compatible with a model of neuropsychological dysfunction in autism in a neural network including parts of the prefrontal cortex, the temporal lobes, and the cerebellum. (shrink)
Evidence for a dichotomy between the planning of an action and its on-line control in humans is reviewed. This evidence suggests that planning and control each serve a specialized purpose utilizing distinct visual representations. Evidence from behavioral studies suggests that planning is influenced by a large array of visual and cognitive information, whereas control is influenced solely by the spatial characteristics of the target, including such things as its size, shape, orientation, and so forth. Evidence from brain imaging and neuropsychology (...) suggests that planning and control are subserved by separate visual centers in the posterior parietal lobes, each constituting part of a larger network for planning and control. Planning appears to rely on phylogenetically newer regions in the inferior parietal lobe, along with the frontal lobes and basal ganglia, whereas control appears to rely on older regions in the superior parietal lobe, along with the cerebellum. Key Words: action; apraxia; control; illusions; optic ataxia; PET; planning; reaching;. (shrink)
Wynn's model for the evolution of spatial cognition is well supported by fossil evidence from brain endocasts, and from neurological studies of the cerebellum and the posterior parietal region of the cerebral cortex. Wynn's intriguing hypothesis that the spatial skill reflected in artifacts is an index of navigational ability, could be further explored by an analysis of lithic transport patterns.