The Role of the Human Cerebellum for Learning from and Processing of External Feedback in Non-Motor Learning: A Systematic Review.

This review aimed to systematically identify and comprehensively review the role of the cerebellum in performance monitoring, focusing on learning from and on processing of external feedback in non-motor learning. While 1078 articles were screened for eligibility, ultimately 36 studies were included in which external feedback was delivered in cognitive tasks and which referenced the cerebellum. These included studies in patient populations with cerebellar damage and studies in healthy subjects applying neuroimaging. Learning performance in patients with different cerebellar diseases was heterogeneous, with only about half of all patients showing alterations. One patient study using EEG demonstrated that damage to the cerebellum was associated with altered neural processing of external feedback. Studies assessing brain activity with task-based fMRI or PET and one resting-state functional imaging study that investigated connectivity changes following feedback-based learning in healthy participants revealed involvement particularly of lateral and posterior cerebellar regions in processing of and learning from external feedback. Cerebellar involvement was found at different stages, e.g., during feedback anticipation and following the onset of the feedback stimuli, substantiating the cerebellum's relevance for different aspects of performance monitoring such as feedback prediction. Future research will need to further elucidate precisely how, where, and when the cerebellum modulates the prediction and processing of external feedback information, which cerebellar subregions are particularly relevant, and to what extent cerebellar diseases alter these processes.

Handedness in schizophrenia and affective disorders: a large-scale cross-disorder study.

While most people are right-handed, a minority are left-handed or mixed-handed. It has been suggested that mental and developmental disorders are associated with increased prevalence of left-handedness and mixed-handedness. However, substantial heterogeneity exists across disorders, indicating that not all disorders are associated with a considerable shift away from right-handedness. Increased frequencies in left- and mixed-handedness have also been associated with more severe clinical symptoms, indicating that symptom severity rather than diagnosis explains the high prevalence of non-right-handedness in mental disorders. To address this issue, the present study investigated the association between handedness and measures of stress reactivity, depression, mania, anxiety, and positive and negative symptoms in a large sample of 994 healthy controls and 1213 patients with DSM IV affective disorders, schizoaffective disorders, or schizophrenia. A series of complementary analyses revealed lower lateralization and a higher percentage of mixed-handedness in patients with major depression (14.9%) and schizophrenia (24.0%) compared to healthy controls (12%). For patients with schizophrenia, higher symptom severity was associated with an increasing tendency towards left-handedness. No associations were found for patients diagnosed with major depression, bipolar disorder, or schizoaffective disorder. In healthy controls, no association between hand preference and symptoms was evident. Taken together, these findings suggest that both diagnosis and symptom severity are relevant for the shift away from right-handedness in mental disorders like schizophrenia and major depression.

Emotional cues reduce Pavlovian interference in feedback-based go and nogo learning.

It is easier to execute a response in the promise of a reward and withhold a response in the promise of a punishment than vice versa, due to a conflict between cue-related Pavlovian and outcome-related instrumental action tendencies in the reverse conditions. This robust learning asymmetry in go and nogo learning is referred to as the Pavlovian bias. Interestingly, it is similar to motivational tendencies reported for affective facial expressions, i.e., facilitation of approach to a smile and withdrawal from a frown. The present study investigated whether and how learning from emotional faces instead of abstract stimuli modulates the Pavlovian bias in reinforcement learning. To this end, 137 healthy adult participants performed an orthogonalized Go/Nogo task that fully decoupled action (go/nogo) and outcome valence (win points/avoid losing points). Three groups of participants were tested with either emotional facial cues whose affective valence was either congruent (CON) or incongruent (INC) to the required instrumental response, or with neutral facial cues (NEU). Relative to NEU, the Pavlovian bias was reduced in both CON and INC, though still present under all learning conditions. Importantly, only for CON, the reduction of the Pavlovian bias effect was adaptive by improving learning performance in one of the conflict conditions. In contrast, the reduction of the Pavlovian bias in INC was completely driven by decreased learning performance in non-conflict conditions. These results suggest a potential role of arousal/salience in Pavlovian-instrumental regulation and cue-action congruency in the adaptability of goal-directed behavior. Implications for clinical application are discussed.

Oligodendrocytes matter: a review of animal studies on early adversity.

Exposure to adversities in early life appears to affect the development of white matter, especially oligodendrocytes. Furthermore, altered myelination is present in regions subjected to maturation during the developmental time when early adversities are experienced. In this review, studies applying two well-established animal models of early life adversity, namely maternal separation and maternal immune activation, focusing on oligodendrocyte alterations and resulting implications for psychiatric disorders are discussed. Studies revealed that myelination is reduced as a result of altered oligodendrocyte expression. Furthermore, early adversity is associated with increased cell death, a simpler morphology, and inhibited oligodendrocyte maturation. However, these effects seem to be region- specific as some brain regions show increased expression while others show decreased expression of oligodendroglia-related genes, and they occur especially in regions of ongoing development. Some studies furthermore suggest that early adversity leads to premature differentiation of oligodendrocytes. Importantly, especially early exposure results in stronger oligodendrocyte-related impairments. However, resulting alterations are not restricted to exposure during the early pre- and postnatal days as social isolation after weaning leads to fewer internodes and branches and shorter processes of oligodendrocytes in adulthood. Eventually, the found alterations may lead to dysfunction and long-lasting alterations in structural brain development associated with psychiatric disorders. To date, only few preclinical studies have focused on the effects of early adversity on oligodendrocytes. More studies including several developmental stages are needed to further disentangle the role of oligodendrocytes in the development of psychiatric disorders.

The role of the cerebellum in internet gaming disorder-A systematic review.

Recent studies increasingly highlight involvement of the cerebellum in drug craving and addiction. However, its exact role, that is, whether the cerebellum is a critical component of a brain network underlying addictive behaviour, or whether it rather is a facilitator or mediator, is still unclear. Findings concerning the newly recognized internet gaming disorder (IGD) suggest that changes in cerebellar connectivity and functioning are associated with behavioural/non-substance addiction. Here, we systematically review the literature on IGD and cerebellar involvement following the PRISMA guidelines. A total of 13 neuroimaging studies met the inclusion criteria. Studies utilized a broad range of diagnostic instruments and resulting cut-off criteria, rendering it difficult to compare findings. Results on altered cerebro-cerebellar connectivity in patients with IGD are mixed; most studies report altered or increased functional connectivity. Moreover, decreased cerebellar grey matter volume is reported. Studies have further indicated that differential activation patterns in the cerebellum may enable discrimination between healthy subjects and subjects with IGD, even allowing for prediction of treatment outcomes. Given the strong connectivity between the cerebellum and cerebral regions, the cerebellum may act as an intermediary between regions involved in craving and addiction and consequently affect symptoms of IGD. Results suggest differential involvement of the cerebellar lobes, emphasizing a need for high-resolution parcellation of the cerebellum in future studies. However, the studies included in the present review have small sample sizes and include mostly male participants. Thus, results may have limited generalizability yet highlight a crucial role of the cerebellum in IGD that needs further investigation.

The impact of social anxiety on feedback-based go and nogo learning.

The term "Pavlovian" bias describes the phenomenon that learning to execute a response to obtain a reward or to inhibit a response to avoid punishment is much easier than learning the reverse. The present study investigated the interplay between this learning bias and individual levels of social anxiety. Since avoidance behavior is a hallmark feature of social anxiety and high levels of social anxiety have been associated with better learning from negative feedback, it is conceivable that the Pavlovian bias is altered in individuals with high social anxiety, with a strong tendency to avoid negative feedback, especially (but not only) in a nogo context. In addition, learning may be modulated by the individual propensity to learn from positive or negative feedback, which can be assessed as a trait-like feature. A sample of 84 healthy university students completed an orthogonalized go/nogo task that decoupled action type (go/nogo) and outcome valence (win/avoid) and a probabilistic selection task based upon which the individual propensity to learn from positive and negative feedback was determined. Self-reported social anxiety and learning propensity were used as predictors in linear mixed-effect model analysis of performance accuracy in the go/nogo task. Results revealed that high socially anxious subjects with a propensity to learn better from negative feedback showed particularly pronounced learning for nogo to avoid while lacking significant learning for nogo to win as well as go to avoid. This result pattern suggests that high levels of social anxiety in concert with negative learning propensity hamper the overcoming of Pavlovian bias in a win context while facilitating response inhibition in an avoidance context. The present data confirm the robust Pavlovian bias in feedback-based learning and add to a growing body of evidence for modulation of feedback learning by individual factors, such as personality traits. Specifically, results show that social anxiety is associated with altered Pavlovian bias, and might suggest that this effect could be driven by altered basal ganglia function primarily affecting the nogo pathway.

Selective Devaluation Affects the Processing of Preferred Rewards.

The present study investigated whether the representation of subjective preferences in the event-related potential is manipulable through selective devaluation, i.e., the consumption of a specific food item until satiety. Thirty-four participants completed a gambling task in which they chose between virtual doors to find one of three snack items, representing a high, medium, or low preference outcome as defined by individual desire-to-eat ratings. In one of two test sessions, they underwent selective devaluation of the high preference outcome. In the other, they completed the task on an empty stomach. Consistent with previous findings, averaged across sessions, amplitudes were increased for more preferred rewards in the time windows of P2, late FRN, and P300. As hypothesised, we also found a selective devaluation effect for the high preference outcome in the P300 time window, reflected in a decrease in amplitude. The present results provide evidence for modulations of reward processing not only by individual factors, such as subjective preferences, but also by the current motivational state. Importantly, the present data suggest that selective devaluation effects in the P300 may be a promising tool to further characterise altered valuation of food rewards in the context of eating disorders and obesity.

Asymmetric coupling of action and outcome valence in active and observational feedback learning.

Learning to execute a response to obtain a reward or to inhibit a response to avoid punishment is much easier than learning the reverse, which has been referred to as "Pavlovian" biases. Despite a growing body of research into similarities and differences between active and observational learning, it is as yet unclear if Pavlovian learning biases are specific for active task performance, i.e., learning from feedback provided for one's own actions, or if they persist also when learning by observing another person's actions and subsequent outcomes. The present study, therefore, investigated the influence of action and outcome valence in active and observational feedback learning. Healthy adult volunteers completed a go/nogo task that decoupled outcome valence (win/loss) and action (execution/inhibition) either actively or by observing a virtual co-player's responses and subsequent feedback. Moreover, in a more naturalistic follow-up experiment, pairs of subjects were tested with the same task, with one subject as active learner and the other as observational learner. The results revealed Pavlovian learning biases both in active and in observational learning, with learning of go responses facilitated in the context of reward obtainment, and learning of nogo responses facilitated in the context of loss avoidance. Although the neural correlates of active and observational feedback learning have been shown to differ to some extent, these findings suggest similar mechanisms to underlie both types of learning with respect to the influence of Pavlovian biases. Moreover, performance levels and result patterns were similar in those observational learners who had observed a virtual co-player and those who had completed the task together with an active learner, suggesting that inclusion of a virtual co-player in a computerized task provides an effective manipulation of agency.

Response inhibition to emotional faces is modulated by functional hemispheric asymmetries linked to handedness.

Effective response inhibition requires efficient bottom-up perceptual processing and effective top-down inhibitory control. To investigate the role of hemispheric asymmetries in these processes, 49 right- and 50 left-handers completed a tachistoscopic Go/Nogo task with positive and negative emotional faces while ERPs were recorded. Frontal resting state EEG asymmetry was assessed as a marker of individual differences in prefrontal inhibitory networks. Results supported a dependency of inhibitory processing on early lateralized processes. As expected, right-handers showed a stronger N170 over the right hemisphere, and better response inhibition when faces were projected to the right hemisphere. Left-handers showed a stronger N170 over the left hemisphere, and no behavioural asymmetry. Asymmetries in response inhibition were also valence-dependent, with better inhibition of responses to negative faces when projected to the right, and better inhibition of responses to positive faces when projected to the left hemisphere. Frontal asymmetry was not related to handedness, but did modulate response inhibition depending on valence. Consistent with the asymmetric inhibition model (Grimshaw & Carmel, 2014), greater right frontal activity was associated with better response inhibition to positive than to negative faces; subjects with greater left frontal activity showed an opposite trend. These findings highlight the interplay between bottom-up and top-down processes in explaining hemispheric asymmetries in response inhibition.

Asymmetries in social touch-motor and emotional biases on lateral preferences in embracing, cradling and kissing.

In human social interaction, affective touch plays an integral role to communicate intentions and emotions. Three of the most important forms of social touch are embracing, cradling and kissing. These behaviours have been demonstrated to be lateralized, but the underlying mechanisms are still not well understood. Both motor and emotive biases have been suggested to affect laterality of social touch. We aimed to systematically investigate how motor preferences and emotive biases influence the lateralization of embracing, cradling and kissing within the same sample. Participants performed all three forms of social touch in neutral, positive and negative emotional conditions. Like a previous study, we found a rightward bias for embracing that was modulated by both motor preferences and the emotional content of the situation. Kissing and cradling were not influenced by motor preferences. In general, a negative emotional connotation of the situation led to a reduction of lateral biases in social touch, independent of the individual direction.

Paw preferences in cats and dogs: Meta-analysis.

Predator-prey relationships have been suggested to be one of the primary evolutionary factors driving the development of functional hemispheric asymmetries. However, lateralization in many predator species is not well understood and existing studies often are statistically underpowered due to small sample sizes and they moreover show conflicting results. Here, we statistically integrated findings on paw preferences in cats and dogs, two predator species within the Carnivora order that are commonly kept as pets in many societies around the globe. For both species, there were significantly more lateralized than non-lateralized animals. We found that 78% of cats and 68% of dogs showed either left- or right-sided paw preference. Unlike humans, neither dogs nor cats showed a rightward paw preference on the population level. For cats, but not dogs, we found a significant sex difference, with female animals having greater odds of being right-lateralized compared to male animals.

Beyond frontal alpha: investigating hemispheric asymmetries over the EEG frequency spectrum as a function of sex and handedness.

Frontal alpha EEG asymmetry, an indirect marker of asymmetries in relative frontal brain activity, are widely used in research on lateralization of emotional processing. While most authors focus on frontal electrode pairs (e.g., F3/F4 or F7/F8), several recent studies have indicated that EEG asymmetries can also be observed outside the frontal lobe and in frequency bands other than alpha. Because the focus of most EEG asymmetry research is on the correlations between asymmetry and other traits, much less is known about the distribution of patterns of asymmetry at the population level. To systematically assess these asymmetries in a representative sample, we determined EEG asymmetries across the head in the alpha, beta, delta and theta frequency bands in 235 healthy adults. We found significant asymmetries in all four frequency bands and across several brain areas, indicating that EEG asymmetries are not limited to frontal alpha. Asymmetries were not modulated by sex. They were modulated by direction of hand preference, with stronger right-handedness predicting greater right (relative to left) alpha power, or greater left (relative to right) activity. Taken together, the present results show that EEG asymmetries other than frontal alpha represent markers of asymmetric brain function that should be explored further.

In vivo measurement of T1 and T2 relaxation times in awake pigeon and rat brains at 7T.

PURPOSE: Establishment of regional longitudinal (T1 ) and transverse (T2 ) relaxation times in awake pigeons and rats at 7T field strength. Regional differences in relaxation times between species and between two different pigeon breeds (homing pigeons and Figurita pigeons) were investigated. METHODS: T1 and T2 relaxation times were determined for nine functionally equivalent brain regions in awake pigeons and rats using a multiple spin-echo saturation recovery method with variable repetition time and a multi-slice/multi-echo sequence, respectively. Optimized head fixation and habituation protocols were applied to accustom animals to the scanning conditions and to minimize movement. RESULTS: The habituation protocol successfully limited movement of the awake animals to a negligible minimum, allowing reliable measurement of T1 and T2 values within all regions of interest. Significant differences in relaxation times were found between rats and pigeons but not between different pigeon breeds. CONCLUSION: The obtained T1 and T2 values for awake pigeons and rats and the optimized habituation protocol will augment future MRI studies with awake animals. The differences in relaxation times observed between species underline the importance of the acquisition of T1 /T2 values as reference points for specific experiments. Magn Reson Med 79:1090-1100, 2018. © 2017 International Society for Magnetic Resonance in Medicine.

The role of the cerebellum for feedback processing and behavioral switching in a reversal-learning task.

Previous studies have reported cerebellar activations during error and reward processing. The present study investigated if the cerebellum differentially processes feedback depending on changes in response strategy during reversal learning, as is conceivable given its internal models for movement and thought. Negative relative to positive feedback in an fMRI-based reversal learning task was hypothesized to be associated with increased cerebellar activations. Moreover, increased activations were expected for negative feedback followed by a change in response strategy compared to negative feedback not followed by such a change, and for first positive feedback after compared to final negative feedback before a change, due to updating of internal models. As predicted, activation in lobules VI and VIIa/Crus I was increased for negative relative to positive feedback, and for final negative feedback before a change in response strategy relative to negative feedback not associated with a change. Moreover, activation was increased for first positive feedback after relative to final negative feedback before a change. These findings are consistent with updating of cerebellar internal models to accommodate new behavioral strategies. Recruitment of posterior regions in reversal learning is in line with the cerebellar functional topography, with posterior regions involved in complex motor and cognitive functions.

The Association Between Eye Movements and Cerebellar Activation in a Verbal Working Memory Task.

It has been argued that cerebellar activations during cognitive tasks may masquerade as cognition, while actually reflecting processes related to movement planning or motor learning. The present study investigated whether the cerebellar load effect for verbal working memory, that is, increased activations in lobule VI/Crus I and lobule VIIB/VIIIA, is related to eye movements and oculomotor processing. Fifteen participants performed an fMRI-based Sternberg verbal working memory task. Oculomotor and cognitive task demands were manipulated by using closely and widely spaced stimuli, and high and low cognitive load. Trial-based quantitative eye movement parameters were obtained from concurrent eye tracking. Conventional MRI analysis replicated the cerebellar load effect in lobules VI and VIIB/VIIIa. With quantitative eye movement parameters as regressors, analysis yielded very similar activation patterns. While load effect and eye regressor generally recruited spatially distinct neocortical and cerebellar regions, conjunction analysis showed that a small subset of prefrontal areas implicated in the load effect also responded to the eye regressor. The present results indicate that cognitive load-dependent activations in lateral superior and posteroinferior cerebellar regions in the Sternberg task are independent of eye movements occurring during stimulus encoding. This is inconsistent with the notion that cognitive load-dependent cerebellar activations merely reflect oculomotor processing.

Look who's judging-Feedback source modulates brain activation to performance feedback in social anxiety.

It is as yet unknown if behavioral and neural correlates of performance monitoring in socially anxious individuals are affected by whether feedback is provided by a person or a computer. This fMRI study investigated modulation of feedback processing by feedback source (person vs. computer) in participants with high (HSA) (N=16) and low social anxiety (LSA) (N=16). Subjects performed a choice task in which they were informed that they would receive positive or negative feedback from a person or the computer. Subjective ratings indicated increased arousal and anxiety in HSA versus LSA, most pronounced for social and negative feedback. FMRI analyses yielded hyperactivation in ventral medial prefrontal cortex (vmPFC)/anterior cingulate cortex (ACC) and insula for social relative to computer feedback, and in mPFC/ventral ACC for positive relative to negative feedback in HSA as compared to LSA. These activation patterns are consistent with increased interoception and self-referential processing in social anxiety, especially during processing of positive feedback. Increased ACC activation in HSA to positive feedback may link to unexpectedness of (social) praise as posited in social anxiety disorder (SAD) psychopathology. Activation in rostral ACC showed a reversed pattern, with decreased activation to positive feedback in HSA, possibly indicating altered action values depending on feedback source and valence. The present findings corroborate a crucial role of mPFC for performance monitoring in social anxiety.

Abnormal brain activation and connectivity to standardized disorder-related visual scenes in social anxiety disorder.

Our understanding of altered emotional processing in social anxiety disorder (SAD) is hampered by a heterogeneity of findings, which is probably due to the vastly different methods and materials used so far. This is why the present functional magnetic resonance imaging (fMRI) study investigated immediate disorder-related threat processing in 30 SAD patients and 30 healthy controls (HC) with a novel, standardized set of highly ecologically valid, disorder-related complex visual scenes. SAD patients rated disorder-related as compared with neutral scenes as more unpleasant, arousing and anxiety-inducing than HC. On the neural level, disorder-related as compared with neutral scenes evoked differential responses in SAD patients in a widespread emotion processing network including (para-)limbic structures (e.g. amygdala, insula, thalamus, globus pallidus) and cortical regions (e.g. dorsomedial prefrontal cortex (dmPFC), posterior cingulate cortex (PCC), and precuneus). Functional connectivity analysis yielded an altered interplay between PCC/precuneus and paralimbic (insula) as well as cortical regions (dmPFC, precuneus) in SAD patients, which emphasizes a central role for PCC/precuneus in disorder-related scene processing. Hyperconnectivity of globus pallidus with amygdala, anterior cingulate cortex (ACC) and medial prefrontal cortex (mPFC) additionally underlines the relevance of this region in socially anxious threat processing. Our findings stress the importance of specific disorder-related stimuli for the investigation of altered emotion processing in SAD. Disorder-related threat processing in SAD reveals anomalies at multiple stages of emotion processing which may be linked to increased anxiety and to dysfunctionally elevated levels of self-referential processing reported in previous studies.

Neural correlates of anticipation and processing of performance feedback in social anxiety.

Fear of negative evaluation, such as negative social performance feedback, is the core symptom of social anxiety. The present study investigated the neural correlates of anticipation and perception of social performance feedback in social anxiety. High (HSA) and low (LSA) socially anxious individuals were asked to give a speech on a personally relevant topic and received standardized but appropriate expert performance feedback in a succeeding experimental session in which neural activity was measured during anticipation and presentation of negative and positive performance feedback concerning the speech performance, or a neutral feedback-unrelated control condition. HSA compared to LSA subjects reported greater anxiety during anticipation of negative feedback. Functional magnetic resonance imaging results showed deactivation of medial prefrontal brain areas during anticipation of negative feedback relative to the control and the positive condition, and medial prefrontal and insular hyperactivation during presentation of negative as well as positive feedback in HSA compared to LSA subjects. The results indicate distinct processes underlying feedback processing during anticipation and presentation of feedback in HSA as compared to LSA individuals. In line with the role of the medial prefrontal cortex in self-referential information processing and the insula in interoception, social anxiety seems to be associated with lower self-monitoring during feedback anticipation, and an increased self-focus and interoception during feedback presentation, regardless of feedback valence.

Disrupted executive cerebro-cerebellar functional connectivity in alcohol use disorder.

BACKGROUND: Alcohol use disorder (AUD) affects 283 million people worldwide and its prevalence is increasing. Despite the role of the cerebellum in executive control and its sensitivity to alcohol, few studies have assessed its involvement in AUD-relevant functional networks. The goal of this study is to compare resting-state functional connectivity (FC) patterns in abstinent adults with a history of AUD and controls (CTL). We hypothesized that group differences in cerebro-cerebellar FC would be present, particularly within the frontoparietal/executive control network (FPN). METHODS: Twenty-eight participants completed a resting-state functional magnetic resonance imaging (rsfMRI) study. CTL participants had no history of AUD, comorbid psychological conditions, or recent heavy drinking and/or drug use. AUD participants had a history of AUD, with sobriety for at least 30 days prior to data collection. Multivariate pattern analysis, an agnostic, whole-brain approach, was used to identify regions with significant differences in FC between groups. Seed-based analyses were then conducted to determine the directionality and extent of these FC differences. Associations between FC strength and executive function were assessed using correlations with Wisconsin Card Sorting Test (WCST) performance. RESULTS: There were significant group differences in FC in nodes of the FPN, ventral attention network, and default mode network. Post hoc analyses predominantly identified FC differences within the cerebro-cerebellar FPN, with AUD showing significantly less FC within the FPN. In AUD, FC strength between FPN clusters identified in the multivariate pattern analysis (MVPA) analysis (Left Crus II, Right Frontal Cortex) was positively associated with performance on the WCST. CONCLUSIONS: Our results show less engagement of the FPN in individuals with AUD than in CTL. FC strength within this network was positively associated with performance on the WCST. These findings suggest that long-term heavy drinking alters cerebro-cerebellar FC, particularly within networks that are involved in executive function.

Cortical processing of saccade-related efference copy signals in patients with cerebellar lesion.

The updating of visual space across saccades is thought to rely on efference copies of motor commands. In humans, thalamic lesions impair performance on a saccadic double-step task, which requires the use of efference copy information, and the altering of saccade-related efference copy processing. This deficit is attributed to disruption of a pathway from the superior colliculus to the frontal eye field. However, the cerebellum is probably also involved in efference copy processing, due to its pivotal role for predictive motor control. The present study investigated the processing of efference copy information in eight patients with focal cerebellar lesions and 22 healthy controls by means of a saccadic double-step task with simultaneous event-related potential recording. Despite intact behavioural performance, a positive event-related potential component between 150 and 450 ms after first saccade onset in the updating condition, which has been interpreted in terms of the integration of efference copy signals with motor intentions for a subsequent saccade, was markedly reduced in the patients. These findings suggest that the cerebellum contributes to on-line saccade monitoring, and that cerebellar lesions alter saccade-related efference copy processing. However, given the intact behavioural performance, the reduced positivity in the patients may indicate that cerebellar damage is accounted for by either exploiting reduced saccade-related information, or making use of compensatory strategies to circumvent a deficit in using efference copy information procured by the cerebellum. The present study extends previous findings on the neural underpinnings of saccadic updating and further elucidates the mechanisms underlying cerebellar predictive motor control.