In the current chapter, I examined the relationship between the cerebellum, emotion, and morality with evidence from large-scale neuroimaging data analysis. Although the aforementioned relationship has not been well studied in neuroscience, recent studies have shown that the cerebellum is closely associated with emotional and social processes at the neural level. Also, debates in the field of moral philosophy, psychology, and neuroscience have supported the importance of emotion in moral functioning. Thus, I explored the potentially important but less-studies topic with (...) NeuroSynth, a tool for large-scale brain image analysis, while addressing issues associated with reverse inference. The result from analysis demonstrated that brain regions in the cerebellum, the right Crus I and Crus II in particular, were specifically associated with morality in general. I discussed the potential implications of the finding based on clinical and functional neuroimaging studies of the cerebellum, emotional functioning, and neural networks for diverse psychological processes. (shrink)

Three neuropsychological experiments on a group of 16 cerebellar patients and 16 age- and education-matched controls investigated the effects of damage to the cerebellum on English grammatical morphology across production, comprehension, and grammaticality judgment tasks. In Experiment 1, participants described a series of pictures previously used in studies of cortical aphasic patients. The cerebellar patients did not differ significantly from the controls in the total number of words produced or in the proportion of closed-class words. They did differ to a (...) marginally significant extent in the production of required articles. In Experiment 2, participants identified the agent in a series of aurally presented sentences in which three agency cues (subject–verb agreement, word order, and noun animacy) were manipulated. The cerebellar patients were less affected than the controls were by the manipulation of subject–verb agreement to a marginally significant extent. In Experiment 3, participants performed a grammaticality judgment task on a series of aurally presented sentences. The cerebellar patients were significantly less able to discriminate grammatical and ungrammatical sentences than the controls were, particularly when the error was of subject–verb agreement as opposed to word order. The results suggest that damage to the cerebellum can result in subtle impairments in the use of grammatical morphology, and are discussed in light of hypothesized roles for the cerebellum in language. (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].

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].

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.

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].

Mormyrid teleosts have Purkinje cells with palisade dendrites, which probably represent coincidence detectors of parallel fiber activity. Their existence strongly supports the ideas of Braitenberg et al. on cerebellar function. However, the organization of mormyrid granule cells and parallel fibers suggests that a key to cerebellar function is not in interactions within one wave, but between twoopposite tidal waves.

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)

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)

An increasing number of studies demonstrated the involvement of the cerebellum in sequence processing. The current preliminary study is the first to investigate the causal involvement of the cerebellum in sequence generation, using low-frequency repetitive transcranial magnetic stimulation. By targeting the posterior cerebellum, we hypothesized that the induced neuro-excitability modulation would lead to altered performance on a Picture and Story sequencing task, which involve the generation of the correct chronological order of various social and non-social stories depicted in cartoons or (...) sentences. Our results indicate that participants receiving LF-rTMS over the cerebellum, as compared to sham participants, showed a stronger learning effect from pre to post stimulation for both tasks and for all types of sequences. No differences between sequence types were observed. Our results suggest a positive effect of LF-rTMS on sequence generation. We conclude that the cerebellum is causally involved in the generation of sequences of social and nonsocial events. Our discussion focuses on recommendations for future studies. (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.

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)

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].

Magnetic resonance imaging (MRI) can provide a number of measurements relevant to sport-related concussion (SRC) symptoms; however, most studies to date have used a single MRI modality and whole-brain exploratory analyses in attempts to localize concussion injury. This has resulted in highly variable findings across studies due to wide ranging symptomology, severity and nature of injury within studies. A multimodal MRI, symptom-guided region-of-interest (ROI) approach is likely to yield more consistent results. The functions of the cerebellum and basal ganglia transcend (...) many common concussion symptoms, and thus these regions, plus the white matter tracts that connect or project from them, constitute plausible ROIs for MRI analysis. We performed diffusion tensor imaging (DTI), resting-state functional MRI, quantitative susceptibility mapping (QSM), and cerebral blood flow (CBF) imaging using arterial spin labeling (ASL), in youth aged 12-18 years following SRC, with a focus on the cerebellum, basal ganglia and white matter tracts. Compared to controls similar in age, sex and sport (N = 20), recent SRC youth (N = 29; MRI at 8 ± 3 days post injury) exhibited increased susceptibility in the cerebellum (p = 0.032), decreased functional connectivity between the caudate and each of the pallidum (p = 0.035) and thalamus (p = 0.021), and decreased diffusivity in the mid-posterior corpus callosum (p < 0.038); no changes were observed in recovered asymptomatic youth (N = 16; 41 ± 16 days post injury). For recent symptomatic-only SRC youth (N = 24), symptom severity was associated with increased susceptibility in the superior cerebellar peduncles (p = 0.011) and reduced activity in the cerebellum (p = 0.013). Fewer days between injury and MRI were associated with reduced cerebellar-parietal functional connectivity (p < 0.014), reduced activity of the pallidum (p = 0.002), increased CBF in the caudate (p = 0.005), and reduced diffusivity in the central corpus callosum (p < 0.05). Youth SRC is associated with acute cerebellar inflammation accompanied by reduced cerebellar activity and cerebellar-parietal connectivity, as well as structural changes of the middle regions of the corpus callosum accompanied by functional changes of the caudate, all of which resolve with recovery. Early MRI post-injury is important to establish objective MRI-based indicators for concussion diagnosis, recovery assessment and prediction of outcome. (shrink)

We review evidence from neuropsychological studies of patients with damage to the cerebellum that suggests cerebellar involvement in four general categories of cognition: (1) speech and language; (2) temporal processing; (3) implicit learning and memory; (4) visuospatial processing and attention. A relatively strong case can be made for cerebellar contributions to language (including speech perception, lexical retrieval, and working memory) and to temporal processing. However, the evidence concerning cerebellar involvement in non-motor implicit learning and visuospatial processing is more equivocal. We (...) argue that cerebellar contributions to cognition are computationally plausible, given its reciprocal connectivity with the cerebral cortex, and suggest that this function of the cerebellum may be an example of an evolutionary process by which mechanisms originally evolved for one function (in this case, motor control) are adapted to other functions (cognition). (shrink)

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.

Functional imaging studies of neurologically intact adults have demonstrated that the right posterior cerebellum is activated during verb generation, semantic processing, sentence processing, and verbal fluency. Studies of patients with cerebellar damage converge to show that the cerebellum supports sentence processing and verbal fluency. However, to date there are no patient studies that investigated the specific importance of the right posterior cerebellum in language processing, because: case studies presented patients with lesions affecting the anterior cerebellum, and group studies combined patients (...) with lesions to different cerebellar regions, without specifically reporting the effects of right posterior cerebellar damage. Here we investigated whether damage to the right posterior cerebellum is critical for sentence processing and verbal fluency in four patients with focal stroke damage to different parts of the right posterior cerebellum. We examined detailed lesion location by going beyond common anatomical definitions of cerebellar anatomy, and employed a recently proposed functional parcellation of the cerebellum. All four patients experienced language difficulties that persisted for at least a month after stroke but three performed in the normal range within a year. In contrast, one patient with more damage to lobule IX than the other patients had profound long-lasting impairments in the comprehension and repetition of sentences, and the production of spoken sentences during picture description. Spoken and written word comprehension and visual recognition memory were also impaired, however, verbal fluency was within the normal range, together with object naming, visual perception and verbal short-term memory. This is the first study to show that focal damage to the right posterior cerebellum leads to language difficulties after stroke; and that processing impairments persisted in the case with most damage to lobule IX. We discuss these results in relation to current theories of cerebellar contribution to language processing. Overall, our study highlights the need for longitudinal studies of language function in patients with focal damage to different cerebellar regions, with functional imaging to understand the mechanisms that support recovery. (shrink)

Precise timing of muscle contractions is an important prerequisite for motor control and one to which the cerebellum contributes. Braitenberg et al.'s detailed timing hypotheses relate only to a subset of the known features of the organisation of the cerebellum. However, the cerebellar architecture clearly supports the that are central to the authors' proposal and such tidal waves are very likely to contribute to its functions.

Ten cerebellar patients were compared to 10 control subjects on a verbal working memory task in which the phonological similarity of the words to be remembered and their modality of presentation were manipulated. Cerebellar patients demonstrated a reduction of the phonological similarity effect relative to controls. Further, this reduction did not depend systematically upon the presentation modality. These results first document that qualitative differences in verbal working memory may be observed following cerebellar damage, indicating altered cognitive processing, even though behavioral (...) output as measured by the digit span may be within normal limits. However, the results also present problems for the hypothesis that the cerebellar role is specifically associated with articulatory rehearsal as conceptualized in the Baddeley–Hitch model of working memory. (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.

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)

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)