Approach and avoidance learning in obsessive-compulsive disorder

Background: Current neurobiological models of obsessive–compulsive disorder (OCD) propose a dysfunction of cortico‐striato‐thalamo‐cortical circuits that leads to enhanced activity in frontal and striatal brain regions. In accordance with that, OCD patients show alterations in learning and flexible adaptation to changing task requirements. The purpose of this study was to examine feedback‐based learning and to investigate whether learning from positive and negative feedback is differentially altered in OCD. Methods: In this study, 18 OCD patients and 18 healthy comparison subjects conducted a probabilistic selection task. The task consisted of an acquisition and a test phase and allowed disentangling the extent of learning based on positive and negative feedback. Results: Groups did not differ during probabilistic feedback learning in the acquisition phase. In the test phase, OCD patients showed a negative learning bias in contrast to comparison subjects who showed a positive learning bias. Patients were better at avoiding stimuli that were initially associated with negative outcomes than at approaching stimuli that were associated with positive feedbacks. This interaction was also found for reaction times in that patients were faster in avoiding negative and slower in approaching positive stimuli. Conclusion: Enhanced avoidance learning was found in OCD patients that points to exaggerated anticipation and avoidance of aversive outcomes. Further studies are required to investigate whether neurobiological mechanisms, such as dopaminergic signaling or outcome processing, in the orbitofrontal cortex relate to enhanced negative learning in OCD. Depression and Anxiety, 2011. © 2010 Wiley‐Liss, Inc.     

Altered emotional and BOLD responses to negative,positive and ambiguous performance feedback in OCD

While abnormal processing of performance feedback has been associated with obsessive–compulsive disorder (OCD), neural responses to different kinds of feedback information, especially to ambiguous feedback are widely unknown. Using fMRI and a performance adaptive time-estimation task, we acquired blood oxygenation level-dependant responses and emotional ratings to positive, negative and ambiguous performance feedback in patients and healthy controls. Negative and ambiguous feedback led to increased levels of anxiety, guilt and shame in patients. Both negative and ambiguous feedback, as compared to positive feedback, induced increased activation of the insular cortex in patients. Furthermore, patients showed no differential activation to negative feedback in the putamen and to ambiguous feedback in the ventromedial prefrontal cortex (VMPFC). Finally, negative feedback induced increased activation in the midcingulate cortex in patients compared to controls. Findings indicate that both negative and ambiguous performance feedbacks are associated with abnormal negative emotions and altered brain activation, in particular increased insula activation, while activation in the putamen and VMPFC does not differentiate between feedback types in OCD patients. This suggests a parallel pattern of increased and decreased neural sensitivity to different kinds of feedback information and a general emotional hyperresponsivity to negative and ambiguous performance feedback in OCD.     

Updating of Visual Space Across Horizontal Saccades in Cerebellar and Thalamic Lesion Patients

Efference copies of motor commands are used to update visual space across saccades, ultimately ensuring transsaccadic constancy of space. Thalamic lesions have been shown to impair efference copy-based saccadic updating in an oculomotor context, i.e., when two successive saccades are required. Moreover, the cerebellum has also been discussed as one possible source of saccade-related efference copy signals. The present study aimed to investigate the effects of thalamic and cerebellar lesions on saccadic updating in a perceptual context. To this end, seven patients with focal cerebellar lesions, seven patients with focal thalamic lesions and 11 healthy controls completed a perceptual localisation task in which the position of a target had to be updated across a single horizontal saccade, while saccade-related event-related potentials (ERPs) were recorded. Contrary to the expectations, localisation precision in both patient groups did not differ from the respective controls. A positive ERP component with centroparietal distribution occurring from about 300 to 500 ms after saccade onset in the updating condition was observed equally pronounced in controls and thalamic lesion patients. In cerebellar lesion patients, there was evidence of a reduction of this relative positivity in the updating condition, particularly for leftward saccades. This finding suggests that cerebellar damage altered the neural processes underlying saccadic updating in a perceptual context without causing overt behavioural deficits.     

Impaired context reversal learning, but not cue reversal learning, in patients with amnestic mild cognitive impairment

It has been proposed that reversal learning is impaired following damage to the orbitofrontal and ventromedial frontal cortex (OFC/VMFC) and to the medial temporal lobe (MTL), including the hippocampal formation. However, the exact characteristics of the MTL-associated reversal learning deficit are not known. To investigate this issue, we assessed 30 newly diagnosed patients with amnestic mild cognitive impairment (aMCI) and 30 matched healthy controls. All patients fulfilled the aMCI criteria of the Mayo Clinic Alzheimer's Disease Research Center and underwent head magnetic resonance imaging that confirmed MTL atrophy. Reversal learning was assessed using a novel reinforcement learning task. Participants first acquired and then reversed stimulus-outcome associations based on negative and positive feedback (losing and gaining points). Stimuli consisted of a cue (geometric shapes) and a spatial context (background color or pattern). Neuropsychological assessment included tasks related to the MTL (paired associates learning), dorsolateral prefrontal cortex (DLPFC) (extradimensional shift, One-touch Stockings of Cambridge), and OFC/VMFC (Holiday Apartment Task). Results revealed that, relative to controls, patients with aMCI exhibited a marked reversal learning deficit, which was highly selective for the reversal of context. The acquisition of stimulus-outcome associations and cue reversal learning were spared. Performance on the context reversal learning task significantly correlated with the right hippocampal volume. In addition, patients with aMCI had deficits on tests related to DLPFC but not to OFC/VMFC. However, DLPFC dysfunctions were not associated with context reversal learning. These results suggest that MTL deficits in aMCI selectively affect context reversal learning when OFC/VMFC functions are spared. This deficit is not influenced by the valence of the outcome (positive or negative feedback) and by executive dysfunctions.     

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.     

Altered reward processing in the nucleus accumbens and mesial prefrontal cortex of patients with posttraumatic stress disorder

Posttraumatic stress disorder (PTSD) is known to be associated with altered medial prefrontal activation in response to threatening stimuli and with behavioural deficits in prefrontal functions such as working memory and attention. Given the importance of these areas and processes for decision-making, this functional magnetic resonance imaging study investigated whether decision-making is altered in patients with PTSD. In particular, the neural response to gain and loss feedback was evaluated in a decision-making task in which subjects could maximise their number of points total by learning a particular response pattern. Behaviourally, controls learned the correct response pattern faster than patients. Functionally, patients and controls differed in their neural response to gains, but not in their response to losses. During the processing of gains in the late phase of learning, PTSD patients as compared to controls showed lower activation in the nucleus accumbens and the mesial PFC, critical structures in the reward pathway. This reduced activation was not due to different rates of learning, since it was similarly present in patients with unimpaired learning performance. These findings suggest that positive outcome information lost its salience for patients with PTSD. This may reflect decreasing motivation as the task progressed.     

Cerebellar lesions alter performance monitoring on the antisaccade task--an event-related potentials study

Error processing is associated with distinct event-related potential components (ERPs), i.e. the error-related negativity (ERN) which occurs within approximately 150 ms and is typically more pronounced than the correct-response negativity (CRN), and the error positivity (Pe) emerging from about 200 to 400 ms after an erroneous response. The short latency of the ERN suggests that the internal error monitoring system acts on rapidly available central information such as an efference copy signal rather than slower peripheral feedback. The cerebellum has been linked to an internal forward-model which enables online performance monitoring by predicting the sensory consequences of actions, most probably by making use of efference copies. In the present study it was hypothesized that the cerebellum is involved in the fast evaluation of saccadic response accuracy as reflected by the ERN. Error processing on an antisaccade task was investigated in eight patients with focal vascular lesions to the cerebellum and 22 control subjects using ERPs. While error rates were comparable between groups, saccadic reaction times (SRTs) were enhanced in the patients, and the error-correct difference waveforms showed reduced amplitudes for patients relative to controls in the ERN time window. Notably, this effect was mainly driven by an increased CRN in the patients. In the later Pe time window, the difference signal yielded higher amplitudes in patients compared to controls mainly because of smaller Pe amplitudes on correct trials in patients. The altered ERN/CRN pattern suggests that the cerebellum is critically involved in fast classification of saccadic accuracy. Largely intact performance accuracy together with increased SRTs and the altered Pe pattern may indicate a compensatory mechanism presumably related to slower, more conscious aspects of error processing in the patients.     

Attention to affective pictures in closed head injury: event-related brain potentials and cardiac responses

We examined whether closed head injury patients show altered patterns of selective attention to stimulus categories that naturally evoke differential responses in healthy people. Self-reported rating and electrophysiological (event-related potentials [ERPs], heart rate [HR]) responses to affective pictures were studied in patients with mild head injury (n = 20; CT/MRI negative), in patients with predominantly frontal brain lesions (n = 12; CT/MRI confirmed), and in healthy controls (n = 20). Affective valence similarly modulated HR and ERP responses in all groups, but group differences occurred that were independent of picture valence. The attenuation of P3-slow wave amplitudes in the mild head injury group indicates a reduction in the engagement of attentional resources to the task. In contrast, the general enhancement of ERP amplitudes at occipital sites in the group with primarily frontal brain injury may reflect disinhibition of input at sensory receptive areas, possibly due to a deficit in top-down modulation performed by anterior control systems.     

The role of the orbitofrontal cortex in human discrimination learning

Several lines of evidence implicate the prefrontal cortex in learning but there is little evidence from studies of human lesion patients to demonstrate the critical role of this structure. To this end, we tested patients with lesions of the frontal lobe (n = 36) and healthy controls (n = 35) on two learning tasks: the weather prediction task (WPT), and an eight-pair concurrent visual discrimination task (‘Choose’). Performance of both tasks was previously shown to be disrupted in patients with Parkinson's disease; the Choose deficit was only present when patients were medicated. Patients with damage to the orbitofrontal cortex (OFC) were significantly impaired on Choose, compared to both healthy controls and non-OFC lesion patients. The OFC lesion patients showed a mild deficit on the first 50 trials of the WPT, compared to the control subjects but not non-OFC lesion patients. The selective deficit in the OFC patients on Choose performance could not be attributed to the larger lesion size in this group, and the deficit was not correlated with the volume of damage to adjacent prefrontal subregions (e.g. anterior cingulate cortex). These data support the notion that the OFC play a role in normal discrimination learning, and suggest qualitative similarities in learning performance of patients with OFC damage and medicated PD patients.     

Dissociation of habit-learning in Parkinson's and cerebellar disease

Damage to the medial-temporal region is known to result in declarative (explicit) memory deficits but nondeclarative (implicit) memory is largely unaffected by such lesions. Earlier studies have shown that some forms of implicit learning depend on cerebellar circuits but remain preserved following affections of the basal ganglia circuits. It is unknown which forms of implicit learning persist in patients with cerebellar pathology but are affected after basal ganglia lesions. Therefore, we determined if a test sensitive for habit-learning (probabilistic classification task) resulted in normal values for patients with cerebellar disease but resulted in affected results in patients with Parkinson's disease (PD). To this end, 23 patients with PD, 16 patients with familial or idiopathic cerebellar degeneration (CD), and 20 controls were tested for habit-learning. There was no impairment of patients with CD for the early learning period but there was abnormal learning in the PD group. For a later learning period, the patients with the PD showed improved performance. We conclude that the probabilistic learning task is an implicit, nonmotor learning task which is sensitive for basal ganglia pathology but remains unaffected in the case of cerebellar pathology. Such a test may be of special interest for the detection and possible neurobehavioral treatment of cognitive and motor deficits.     

Interhemispheric Connectivity Characterizes Cortical Reorganization in Motor-Related Networks After Cerebellar Lesions

Although cerebellar-cortical interactions have been studied extensively in animal models and humans using modern neuroimaging techniques, the effects of cerebellar stroke and focal lesions on cerebral cortical processing remain unknown. In the present study, we analyzed the large-scale functional connectivity at the cortical level by combining high-density electroencephalography (EEG) and source imaging techniques to evaluate and quantify the compensatory reorganization of brain networks after cerebellar damage. The experimental protocol comprised a repetitive finger extension task by 10 patients with unilateral focal cerebellar lesions and 10 matched healthy controls. A graph theoretical approach was used to investigate the functional reorganization of cortical networks. Our patients, compared with controls, exhibited significant differences at global and local topological level of their brain networks. An abnormal rise in small-world network efficiency was observed in the gamma band (30–40 Hz) during execution of the task, paralleled by increased long-range connectivity between cortical hemispheres. Our findings show that a pervasive reorganization of the brain network is associated with cerebellar focal damage and support the idea that the cerebellum boosts or refines cortical functions. Clinically, these results suggest that cortical changes after cerebellar damage are achieved through an increase in the interactions between remote cortical areas and that rehabilitation should aim to reshape functional activation patterns. Future studies should determine whether these hypotheses are limited to motor tasks or if they also apply to cerebro-cerebellar dysfunction in general.     

Probabilistic classification learning with corrective feedback is selectively impaired in early Huntington's disease--evidence for the role of the striatum in learning with feedback

In general, declarative learning is associated with the activation of the medial temporal lobes (MTL), while the basal ganglia (BG) are considered the substrate for procedural learning. More recently it has been demonstrated the distinction of these systems may not be as absolute as previously thought and that not only the explicit or implicit nature of the memory task alone is important for the distinction of MTL or BG systems. Nevertheless, patients with BG dysfunction - such as patients with Parkinson's disease (PD) or Huntington's disease (HD) - are considered to be impaired at implicit learning. However, a more recent study demonstrated that one implicit learning task, probabilistic classification learning (examples include the weather prediction (WPT) and Mr. Potato Head tasks) is only impaired in PD when it involves learning with corrective feedback (FB) but not when it involves learning in a paired associate (PA) manner, without feedback. Therefore, it has been argued that the presence of feedback rather than the implicit nature of these tasks determines whether or not the BG are recruited. As patients with HD as well as those with PD, have also been shown to be impaired on the standard FB based version of probabilistic classification learning, the question remains as to whether or not there is a similar selective deficit in FB but not PA based probabilistic classification learning in HD. 18 patients with early HD and 18 healthy controls completed FB and PA versions of the WPT task. Relative to controls, HD patients were selectively impaired at WPT learning with feedback. These findings are consistent with previous evidence from studies of probabilistic classification learning in PD. Unlike PD, selective deficits in WPT learning in HD cannot be attributed to the effects of dopaminergic medication and must be directly related to BG dysfunction; for instance even in early HD, only 50% of the neurons in the medial head of caudate remain. We conclude that the striatum is important for WPT learning with feedback. Our findings are consistent with imaging evidence showing recruitment of the caudate during FB based WPT learning, while the MTL is associated with PA based learning.     

The effect of feedback on non-motor probabilistic classification learning in Parkinson's disease

It has been proposed that procedural learning is mediated by the striatum and, it has been reported that patients with Parkinson's disease (PD) are impaired on the weather prediction task (WPT) which involves probabilistic classification learning with corrective feedback (FB). However, PD patients were not impaired on probabilistic classification learning when it was performed without corrective feedback, in a paired associate (PA) manner; suggesting that the striatum is involved in learning with feedback rather than procedural learning per se. In Experiment 1 we studied FB- and PA-based learning in PD patients and controls and, as an improvement on previous methods, used a more powerful repeated measures design and more equivalent test phases during FB and PA conditions (including altering the FB condition to remove time limits on responding). All participants (16 PD patients, H&Y I-III and 14 matched-controls) completed the WPT under both FB and PA conditions. In contrast to previous results, in Experiment 1 we did not find a selective impairment in the PD group on the FB version of the WPT relative to controls. In Experiment 2 we used a between groups design and studied learning with corrective FB in 11 PD patients (H&Y I.5-IV) and 13 matched controls on a more standard version of the WPT similar to that used in previous studies. With such a between groups design for comparison of FB and PA learning on the WPT in PD, we observed impaired learning in PD patients relative to controls across both the FB and PA versions of the WPT. Most importantly, in Experiment 2 we also failed to find a selective impairment on the FB version of the WPT coupled with normal learning on the PA version in PD patients relative to controls. Our results do not support the proposal that the striatum plays a specific role in probabilistic classification learning with feedback.     

Better or worse than expected? Aging, learning, and the ERN

This study examined age differences in error processing and reinforcement learning. We were interested in whether the electrophysiological correlates of error processing, the error-related negativity (ERN) and the feedback-related negativity (FRN), reflect learning-related changes in younger and older adults. To do so, we applied a probabilistic learning task in which we manipulated the validity of feedback. The results of our study showed that learning-related changes were much more pronounced (a) in a response-locked positivity for correct trials compared to the ERN and (b) in a feedback-locked positivity for positive feedback compared to the FRN. These findings provide an important extension to recent theoretical accounts [Holroyd, C. B., & Coles, M. G. H. (2002). The neural basis of human error processing: Reinforcement learning, dopamine, and the error-related negativity. Psychological Review, 109, 679-709; Nieuwenhuis, S., Ridderinkhof, K. R., Talsma, D., Coles, M. G. H., Holroyd, C. B., Kok, A., et al. (2002). A computational account of altered error processing in older age: Dopamine and the error-related negativity. Cognitive, Affective and Behavioral Neuroscience, 2, 19-36] since they suggest that positive learning signals on correct trials contribute to the reward-related variance in the response- and feedback-locked ERPs. This effect has been overlooked in previous studies that have focused on the role of errors and negative feedback for learning. Importantly, we did not find evidence for an age-related reduction of the ERN, when controlling for performance differences between age groups, which questions the view that older adults are generally impaired in error processing. Finally, we observed a substantial reduction of the FRN in the elderly, which indicates that older adults are less affected by negative feedback and rely more on positive feedback during learning. This finding points to an age-related asymmetry in the processing of feedback valence.     

The feedback-related negativity is modulated by feedback probability in observational learning

The feedback-related negativity (FRN), an event-related potentials (ERPs) component reflecting activity of the anterior cingulate cortex (ACC), has been shown to be modulated by feedback expectancy following active choices in feedback-based learning tasks. A general reduction of FRN amplitude has been described in observational feedback learning, raising the question whether FRN amplitude is modulated in a similar way in this type of learning. The present study investigated whether the FRN and the P300 - a second ERP component related to feedback processing - are modulated by feedback probability in observational learning. Thirty-two subjects participated in the experiment. They observed a virtual person choosing between two symbols and receiving positive or negative feedback. Learning about stimulus-specific feedback probabilities was assessed in active test trials without feedback. In addition, the bias to learn from positive or negative feedback and - in a subsample of 17 subjects - empathy scores were obtained. General FRN and P300 modulations by feedback probability were found across all subjects. Only for the FRN in learners, an interaction between probability and valence was observed. Larger FRN amplitudes for negative relative to positive feedback only emerged for the lowest outcome probability. The results show that feedback expectancy modulates FRN amplitude also in observational learning, suggesting a similar ACC function as in active learning. On the other hand, the modulation is only seen for very low feedback expectancy, which suggests that brain regions other than those of the reward system contribute to feedback processing in an observation setting.     

Telling good from bad news: ADHD differentially affects processing of positive and negative feedback during guessing

Neuroimaging studies on ADHD suggest abnormalities in brain regions associated with decision-making and reward processing such as the anterior cingulate cortex (ACC) and orbitofrontal cortex. Recently, event-related potential (ERP) studies demonstrated that the ACC is involved in processing feedback signals during guessing and gambling. The resulting negative deflection, the 'feedback-related negativity' (FRN) has been interpreted as reflecting an error in reward prediction. In the present study, ERPs elicited by positive and negative feedback were recorded in children with ADHD and normal controls during guessing. 'Correct' and 'incorrect' guesses resulted in respectively monetary gains and losses. The FRN amplitude to losses was more pronounced in the ADHD group than in normal controls. Positive and negative feedback differentially affected long latency components in the ERP waveforms of normal controls, but not ADHD children. These later deflections might be related to further emotional or strategic processing. The present findings suggest an enhanced sensitivity to unfavourable outcomes in children with ADHD, probably due to abnormalities in mesolimbic reward circuits. In addition, further processing, such as affective evaluation and the assessment of future consequences of the feedback signal seems to be altered in ADHD. These results may further help understanding the neural basis of decision-making deficits in ADHD.     

Decoupling of autonomic and cognitive emotional reactions after cerebellar stroke

Emotional blunting can be found after cerebellar lesions. However, the mechanism of such a modification is not clear. We present a patient with emotional flattening and increased risk taking after left cerebellar infarct who had an impaired autonomic reactivity to negative as compared with positive reinforcement. This impairment was demonstrated by the patient's undifferentiated skin conductance responses to negative and positive reinforcement, whereas controls produced larger skin conductance responses after negative feedback. The cooccurrence of emotional flattening and undifferentiated autonomic reactions to positively and negatively valenced stimuli strengthens the role of the cerebellum in the modulation of the autonomic responses.     

Does the cerebellum contribute to specific aspects of attention?

We present data on attentional and neuropsychological functions of 16 patients with focal cerebellar lesions (13 tumours, 3 haematomas) compared to normative test data, and to 11 control subjects matched for age, gender, and years of education. Patients showed distinct deficits in qualitative aspects of a divided attention task, and in a working memory task. Performance in selective attention was unimpaired. The results support the concept that the cerebellum plays a role not only in motor, but also in higher cognitive functions. They are discussed on the basis of the idea that prediction and preparation are fundamental functions of the cerebellum. Therefore, the results confirm the idea that cerebellar lesions lead to reduced performance in specific attention tasks.     

Phonological grouping is specifically affected in cerebellar patients: a verbal fluency study

OBJECTIVES: Recent clinical and functional neuroimaging evidence points towards a cerebellar role in verbal production. At present it is not clear how the cerebellum participates in language production. The aim was to investigate the influence of cerebellar lesions on verbal fluency abilities with specific focus on the verbal searching strategies employed by patients with cerebellar damage. METHODS: Twenty five patients with focal or degenerative cerebellar disease and 14 control subjects were tested in a timed verbal fluency task requiring word production under forced (phonemic or semantic) conditions. To analyse the verbal searching strategy employed, semantic and phonemic cluster analyses were also performed. RESULTS: Performances of cerebellar patients were comparable with those of controls in the semantic task; conversely their performances were significantly impaired when tested in the letter task. Cluster analysis results showed that the verbal fluency impairment is linked to specific damage of phonemically related retrieval strategies. CONCLUSION: Cerebellar damage impairs verbal fluency by specifically affecting phonemic rule performances while sparing semantic rule ones. These findings underline the importance of the cerebellar computing properties in strategy development in the linguistic domain.     

Cerebellar Contribution to Spatial Event Processing: Right/Left Discrimination Abilities in Rats

Recently, we demonstrated the involvement of cerebellar circuits in the procedural components of spatial information processing by testing hemicerebellectomized (HCbed) rats in classical spatial paradigms, such as the Morris Water Maze and the water T-maze. Since procedural components are strongly present in these tests, an impairment also in processing more abstract spatial information, linked to 'where an object is' rather than to 'how to find it', could be hidden by the severe procedural deficits. On this basis, we investigated the influence of cerebellar lesions on spatial abilities strictly reducing procedural variables by employing an active avoidance task, first without and then with a request for righvleft discrimination. In the two-way active avoidance task without spatial requests, controls and cerebellar operated rats developed active avoidance responses which were not statistically different, demonstrating that this kind of associative learning is not significantly affected by hemicerebellectomy (HCb). A second experimental group of cerebellar lesioned rats was tested in a modified version of this basic paradigm in which a rightheft discrimination request was added. This group displayed severe deficits, which even in the last testing sessions prevented them from performing comparably to the control animals. Reversal of the rewarded choice, even if it affected the performances of both controls and operated rats in the first inversion trials, elicited the lowest number of correct responses in HCbed rats throughout the entire spatial reversal learning, suggesting a severe deficit in the ability to change an initially learned behaviour. These results demonstrate that, beside having a marked impairment in facing procedural components of spatial processing, cerebellar lesioned rats are severely defective also in rightlleft discrimination tasks, suggesting a role of cerebellar networks also in the discriminative spatial information processing.