Cerebrocerebellar networks during articulatory rehearsal and verbal working memory tasks (original) (raw)

Functional activation in the cerebellum during working memory and simple speech tasks

Cortex, 2010

Verbal working memory is the ability to temporarily store and manipulate verbal information. This study tested the predictions of a neuroanatomical model of how the cerebellum contributes to verbal working memory . In this model, a large bilateral region in the superior cerebellum is associated with articulatory rehearsal and a right-lateralized region in the inferior cerebellum is associated with the correction of errors within the working memory system. The model was based on neuroimaging findings using item recognition tasks and comparisons between working memory and covert rehearsal tasks, whereas in this fMRI study we used a delayed serial recall (DSR) task because it relies more heavily on articulatory rehearsal, and our comparison tasks included both overt and covert speech tasks. Our results provide some support for the model. In particular, we found multiple activation foci within the superior and inferior sectors of the cerebellum and evidence that these regions show different patterns of activation across working memory and speech tasks. However, the specific patterns of activation were not fully consistent with those reported by . Namely, our results indicate that activation in the superior sector should be functionally subdivided into a medial focus involved in speech processing and a lateral focus more specific to verbal working memory; the results also indicate that activation in the inferior sector is not uniquely right-lateralized. These complex findings speak to the need for future studies to consider the speech-motor aspects of tasks, to investigate the functional significance of adjacent peaks of activation within large regions of cerebellar activation, and to use analysis procedures that support regional distinctions through direct statistical tests. Such studies would help to refine our understanding of how the cerebellum contributes to speech and verbal working memory.

Modality Specific Cerebro-Cerebellar Activations in Verbal Working Memory: An fMRI Study

Behavioural Neurology, 2010

Verbal working memory (VWM) engages frontal and temporal/parietal circuits subserving the phonological loop, as well as, superior and inferior cerebellar regions which have projections from these neocortical areas. Different cerebro-cerebellar circuits may be engaged for integrating aurally- and visually-presented information for VWM. The present fMRI study investigated load (2, 4, or 6 letters) and modality (auditory and visual) dependent cerebro-cerebellar VWM activation using a Sternberg task. FMRI revealed modality-independent activations in left frontal (BA 6/9/44), insular, cingulate (BA 32), and bilateral inferior parietal/supramarginal (BA 40) regions, as well as in bilateral superior (HVI) and right inferior (HVIII) cerebellar regions. Visual presentation evoked prominent activations in right superior (HVI/CrusI) cerebellum, bilateral occipital (BA19) and left parietal (BA7/40) cortex while auditory presentation showed robust activations predominately in bilateral temporal ...

Load- and practice-dependent increases in cerebro-cerebellar activation in verbal working memory: an fMRI study

NeuroImage, 2005

Load-dependent and practice-related changes in neocortical and cerebellar structures involved in verbal working memory (VWM) were investigated using functional MRI (fMRI) and a two alternative forced choice Sternberg paradigm. Using working memory loads ranging from 2 to 6 letters, regions exhibiting linear and quadratic trends in load-dependent activations were identified. Behaviorally, reaction time measurements revealed significant linear increases with increasing memory load, and significant decreases with increased task practice. Brain activations indicated a preponderance of linear load-dependent responses in both superior (lobule VI/Crus I) and inferior (lobule VIIB/ VIIIA) cerebellar hemispheres, as well as in areas of neocortex including left precentral (BA 6), inferior frontal (BA 47), parahippocampal (BA 35), inferior parietal (BA 40), cingulate (BA 32), and right inferior and middle frontal (BA 46/47) regions. Fewer voxels exhibited quadratic without linear trends with the most prominent of these activations located in left inferior parietal (BA 40), precuneus, and parahippocampal regions. Analysis of load  session interactions revealed that right inferior cerebellar and left inferior parietal activations increased with practice, as did the correlations between activation in each region with reaction time, suggesting that changes in this cerebro-cerebellar network underlie practice-related increases in efficiency of VWM performance. These results replicate and extend our previous findings of fMRI activation in the cerebellum during VWM, and demonstrate predominately linear increases in cerebro-cerebellar activation with increasing memory load as well as changes in network function with increased task proficiency.

Lobular patterns of cerebellar activation in verbal working-memory and finger-tapping tasks as revealed by functional MRI

The Journal of neuroscience : the official journal of the Society for Neuroscience, 1997

The lobular distributions of functional activation of the cerebellum during verbal working-memory and finger movement tasks were investigated using functional magnetic resonance imaging (fMRI). Relative to a rest control, finger tapping of the right hand produced ipsilateral-increased activation in HIV/HV [Roman numeral designations based on Larsell's () nomenclature] and HVI and weaker activation in HVIII that was stronger on the ipsilateral side. For a working-memory task, subjects were asked to remember six (high load) or one (low load) visually presented letters across a brief delay. To assess the motoric aspects of rehearsal in the absence of working memory, we asked the subjects to repeatedly read subvocally six or one letters at a rate that approximated the internally generated rehearsal of working memory (motoric rehearsal task). For both tasks, bilateral regions of the superior cerebellar hemispheres (left superior HVIIA and right HVI) and portions of posterior vermis (...

Temporal dynamics of cerebro-cerebellar network recruitment during a cognitive task

Neuropsychologia, 2005

Previous investigations have demonstrated that two regions in the right cerebellum, one located superiorly in hemispheral lobule VI/Crus I and another located inferiorly in hemispheral lobule VIIB/VIIIA, are activated during verbal working memory performance. On the basis of functional neuroimaging patterns of activation, as well as known cortico-pontine and ponto-cerebellar projections, the superior region has been hypothesized to contribute to the articulatory control system of working memory whereas the inferior region has been linked to the phonological store. The present study used event-related fMRI and individual estimates of hemodynamic response for both the cerebellum and neocortex to test this model and characterize the task phase specific cerebro-cerebellar activations for a Sternberg verbal working memory task. Results demonstrated that the right superior cerebellum showed the strongest activation during the initial encoding phase of the task, and, consistent with predictions, a similar pattern was observed in left opercular inferior frontal and premotor regions. In contrast, the right inferior cerebellum exhibited the greatest activation during the maintenance phase of the task, and as predicted, corresponded with activation in the left inferior parietal lobule. The significance of the results with respect to cerebro-cerebellar models of verbal working memory and to theoretical accounts of cerebellar involvement in cognition is discussed.

Functional topography of the cerebellum for motor and cognitive tasks: An fMRI study

NeuroImage, 2012

Anatomical, clinical and imaging findings suggest that the cerebellum is engaged in cognitive and affective functions as well as motor control. Evidence from converging modalities also indicates that there is a functional topography in the human cerebellum for overt control of movement vs. higher functions, such that the cerebellum can be divided into zones depending on connectivity with sensorimotor vs. multimodal association cortices. Using functional MRI, we show that regions active during overt movement differ from those involved in higher-level language, spatial processing and working memory tasks. Nine healthy participants each completed five tasks in order to determine the relative activation patterns for the different paradigms. Right-handed finger-tapping activated right cerebellar lobules IV-V and VIII, consistent with descriptions of the cerebellar homunculi. Verb generation engaged right cerebellar lobules VI-Crus I and a second cluster in lobules VIIB-VIIIA. Mental rotation activation peaks were localized to medial left cerebellar lobule VII (Crus II). A 2-back working memory task activated bilateral regions of lobules VI-VII. Viewing arousing vs. neutral images did not reliably activate the cerebellum or cerebral limbic areas in this study. The cerebellar functional topography identified in this study reflects the involvement of different cerebro-cerebellar circuits depending on the demands of the task being performed: overt movement activated sensorimotor cortices along with contralateral cerebellar lobules IV-V and VIII, whereas more cognitively demanding tasks engaged prefrontal and parietal cortices along with cerebellar lobules VI and VII. These findings provide further support for a cerebellar role in both motor and cognitive tasks, and better establish the existence of functional subregions in the cerebellum. Future studies are needed to determine the exact contribution of the cerebellumand different cerebro-cerebellar circuitsto task performance.

Disruption of Cerebellar Prediction in Verbal Working Memory

Mounting evidence suggests that the right cerebellum contributes to verbal working memory, but the functional role of this contribution remains unclear. In an established theory of motor control, the cerebellum is thought to predict sensory consequences of movements through an internal "forward model." Here, we hypothesize a similar predictive process can generalize to cerebellar non-motor function, and that the right cerebellum plays a predictive role that is beneficial for rapidly engaging the phonological loop in verbal working memory. To test this hypothesis, double-pulse transcranial magnetic stimulation (TMS) was administered over either the right cerebellum or right occipital lobe (control site), on half the trials, to interrupt the rehearsal of a 6-letter sequence. We found that cerebellar stimulation resulted in greater errors in participants' report of the letter in the current position. Additional analyses revealed that immediately after cerebellar TMS, participants were more likely to use out of date information to predict the next letter in the sequence. This pattern of errors is consistent with TMS causing a temporary disruption of state estimation and cerebellar forward model function, leading to prediction errors in the phonological loop.

Sensorimotor, language, and working memory representation within the human cerebellum

Human Brain Mapping, 2019

The cerebellum is involved in a wide range of behaviours. A key organisational principle from animal studies is that somatotopically corresponding sensory input and motor output reside in the same cerebellar cortical areas. However, compelling evidence for a similar arrangement in humans and whether it extends to cognitive functions is lacking. To address this, we applied cerebellar optimised whole-brain functional MRI in 20 healthy subjects. To assess spatial overlap within the sensorimotor and cognitive domains, we recorded activity to a sensory stimulus (vibrotactile) and a motor task; the Sternberg verbal working memory (VWM) task; and a verb generation paradigm. Consistent with animal data, sensory and motor activity overlapped with a somatotopic arrangement in ipsilateral areas of the anterior and posterior cerebellum. During the maintenance phase of the Sternberg task, a positive linear relationship between VWM load and activity was observed in right Lobule VI, extending into Crus I bilaterally. Articulatory movement gave rise to bilateral activity in medial Lobule VI. A conjunction of two independent language tasks localised activity during verb generation in right Lobule VI-Crus I, which overlapped with activity during VWM. These results demonstrate spatial compartmentalisation of sensorimotor and cognitive function in the human cerebellum, with each area involved in more than one aspect of a given behaviour, consistent with an integrative function. Sensorimotor localisation was uniform across individuals, but the representation of cognitive tasks was more variable, highlighting the importance of individual scans for mapping higher order functions within the cerebellum.