Task load modulates tDCS effects on brain network for phonological processing.

Motor participation in phonological processing can be modulated by task nature across the speech perception to speech production range. The pars opercularis of the left inferior frontal gyrus (LIFG) would be increasingly active across this range, because of changing motor demands. Here, we investigated with simultaneous tDCS and fMRI whether the task load modulation of tDCS effects translates into predictable patterns of functional connectivity. Findings were analysed under the "multi-node framework", according to which task load and the network structure underlying cognitive functions are modulators of tDCS effects. In a within-subject study, participants (N = 20) performed categorical perception, lexical decision and word naming tasks [which differentially recruit the target of stimulation (LIFG)], which were repeatedly administered in three tDCS sessions (anodal, cathodal and sham). The LIFG, left superior temporal gyrus and their right homologues formed the target network subserving phonological processing. C-tDCS inhibition and A-tDCS excitation should increase with task load. Correspondingly, the larger the task load, the larger the relevance of the target for the task and smaller the room for compensation of C-tDCS inhibition by less relevant nodes. Functional connectivity analyses were performed with partial correlations, and network compensation globally inferred by comparing the relative number of significant connections each condition induced relative to sham. Overall, simultaneous tDCS and fMRI was adequate to show that motor participation in phonological processing is modulated by task nature. Network responses induced by C-tDCS across phonological processing tasks matched predictions. A-tDCS effects were attributed to optimisation of network efficiency.

Task load modulates tDCS effects on language performance.

Transcranial direct current stimulation (tDCS) effects in cognition are inconsistent across studies. This study aimed to discuss why typical models might be insufficient to explain these effects, and to investigate a brain state factor, task load, with behavioral experiments on phonological processing. The motor theory of speech perception states that motor codes for articulation take part in speech perception, a view sharpened by neuroimaging findings, which show that the motor role in phonological processing is weighted by the nature of the tasks. Three groups of 20 participants, each under a different tDCS condition (anodal, cathodal, or sham), performed a categorical perception (CP), a lexical decision (LD), and a word naming (WN) task while stimulated on the pars opercularis of the left inferior frontal gyrus, a language area typically involved with the motor role. These tasks were assumed to be subserved by a network of nodes which included the target, believed to be increasingly relevant for performance from speech perception to speech production. A-tDCS facilitation and C-tDCS downregulation should directly increase with the relevance of the target for the task. Downregulation of a low relevance node could result in facilitation by compensation from other nodes. Overall, our brain stimulation findings support the neuroimaging literature in that motor participation in phonological processing depends on task nature and show that tDCS effects are modulated by task load relative to the target. Outcomes such as the improved performance following cathodal tDCS in CP and WN suggest that compensatory mechanisms may take place when the tasks involve more complex neuronal networks.

Cerebellar Transcranial Direct Current Stimulation (ctDCS): A Novel Approach to Understanding Cerebellar Function in Health and Disease.

The cerebellum is critical for both motor and cognitive control. Dysfunction of the cerebellum is a component of multiple neurological disorders. In recent years, interventions have been developed that aim to excite or inhibit the activity and function of the human cerebellum. Transcranial direct current stimulation of the cerebellum (ctDCS) promises to be a powerful tool for the modulation of cerebellar excitability. This technique has gained popularity in recent years as it can be used to investigate human cerebellar function, is easily delivered, is well tolerated, and has not shown serious adverse effects. Importantly, the ability of ctDCS to modify behavior makes it an interesting approach with a potential therapeutic role for neurological patients. Through both electrical and non-electrical effects (vascular, metabolic) ctDCS is thought to modify the activity of the cerebellum and alter the output from cerebellar nuclei. Physiological studies have shown a polarity-specific effect on the modulation of cerebellar-motor cortex connectivity, likely via cerebellar-thalamocortical pathways. Modeling studies that have assessed commonly used electrode montages have shown that the ctDCS-generated electric field reaches the human cerebellum with little diffusion to neighboring structures. The posterior and inferior parts of the cerebellum (i.e., lobules VI-VIII) seem particularly susceptible to modulation by ctDCS. Numerous studies have shown to date that ctDCS can modulate motor learning, and affect cognitive and emotional processes. Importantly, this intervention has a good safety profile; similar to when applied over cerebral areas. Thus, investigations have begun exploring ctDCS as a viable intervention for patients with neurological conditions.

Task-Specific Facilitation of Cognition by Anodal Transcranial Direct Current Stimulation of the Prefrontal Cortex.

We previously speculated that depression of cerebellar excitability using cathodal transcranial direct current stimulation (tDCS) might release extra cognitive resources via the disinhibition of activity in prefrontal cortex. The objective of the present study was to investigate whether anodal tDCS over the prefrontal cortex could similarly improve performance when cognitive demands are high. Sixty-three right-handed participants in 3 separate groups performed the Paced Auditory Serial Addition Task (PASAT) and the more difficult Paced Auditory Serial Subtraction Task (PASST), before and after 20 min of anodal, cathodal, or sham stimulation over the left dorsolateral prefrontal cortex (DLPFC). Performance was assessed in terms of the accuracy, latency, and variability of correct verbal responses. All behavioral measures significantly improved for the PASST after anodal DLPFC stimulation, but not the PASAT. There were smaller practice effects after cathodal and sham stimulation. Subjective ratings of attention and mental fatigue were unchanged by tDCS over time. We conclude that anodal stimulation over the left DLPFC can selectively improve performance on a difficult cognitive task involving arithmetic processing, verbal working memory, and attention. This result might be achieved by focally improving executive functions and/or cognitive capacity when tasks are difficult, rather than by improving levels of arousal/alertness.

Modulating cognition using transcranial direct current stimulation of the cerebellum.

Numerous studies have emerged recently that demonstrate the possibility of modulating, and in some cases enhancing, cognitive processes by exciting brain regions involved in working memory and attention using transcranial electrical brain stimulation. Some researchers now believe the cerebellum supports cognition, possibly via a remote neuromodulatory effect on the prefrontal cortex. This paper describes a procedure for investigating a role for the cerebellum in cognition using transcranial direct current stimulation (tDCS), and a selection of information-processing tasks of varying task difficulty, which have previously been shown to involve working memory, attention and cerebellar functioning. One task is called the Paced Auditory Serial Addition Task (PASAT) and the other a novel variant of this task called the Paced Auditory Serial Subtraction Task (PASST). A verb generation task and its two controls (noun and verb reading) were also investigated. All five tasks were performed by three separate groups of participants, before and after the modulation of cortico-cerebellar connectivity using anodal, cathodal or sham tDCS over the right cerebellar cortex. The procedure demonstrates how performance (accuracy, verbal response latency and variability) could be selectively improved after cathodal stimulation, but only during tasks that the participants rated as difficult, and not easy. Performance was unchanged by anodal or sham stimulation. These findings demonstrate a role for the cerebellum in cognition, whereby activity in the left prefrontal cortex is likely dis-inhibited by cathodal tDCS over the right cerebellar cortex. Transcranial brain stimulation is growing in popularity in various labs and clinics. However, the after-effects of tDCS are inconsistent between individuals and not always polarity-specific, and may even be task- or load-specific, all of which requires further study. Future efforts might also be guided towards neuro-enhancement in cerebellar patients presenting with cognitive impairment once a better understanding of brain stimulation mechanisms has emerged.

Restoring cognitive functions using non-invasive brain stimulation techniques in patients with cerebellar disorders.

Numerous studies have highlighted the possibility of modulating the excitability of cerebro-cerebellar circuits bi-directionally using transcranial electrical brain stimulation, in a manner akin to that observed using magnetic stimulation protocols. It has been proposed that cerebellar stimulation activates Purkinje cells in the cerebellar cortex, leading to inhibition of the dentate nucleus, which exerts a tonic facilitatory drive onto motor and cognitive regions of cortex through a synaptic relay in the ventral-lateral thalamus. Some cerebellar deficits present with cognitive impairments if damage to non-motor regions of the cerebellum disrupts the coupling with cerebral cortical areas for thinking and reasoning. Indeed, white matter changes in the dentato-rubral tract correlate with cognitive assessments in patients with Friedreich ataxia, suggesting that this pathway is one component of the anatomical substrate supporting a cerebellar contribution to cognition. An understanding of the physiology of the cerebro-cerebellar pathway previously helped us to constrain our interpretation of results from two recent studies in which we showed cognitive enhancements in healthy participants during tests of arithmetic after electrical stimulation of the cerebellum, but only when task demands were high. Others studies have also shown how excitation of the prefrontal cortex can enhance performance in a variety of working memory tasks. Thus, future efforts might be guided toward neuro-enhancement in certain patient populations, using what is commonly termed "non-invasive brain stimulation" as a cognitive rehabilitation tool to modulate cerebro-cerebellar circuits, or for stimulation over the cerebral cortex to compensate for decreased cerebellar drive to this region. This article will address these possibilities with a review of the relevant literature covering ataxias and cerebellar cognitive affective disorders, which are characterized by thalamo-cortical disturbances.

Task-specific facilitation of cognition by cathodal transcranial direct current stimulation of the cerebellum.

A role for the cerebellum in cognition is controversial, but it is a view that is becoming increasingly popular. The aim of the current study was to investigate this issue using transcranial Direct Current Stimulation (tDCS) during two cognitive tasks that require comparable motor skills, but different levels of working memory and attention. Three groups of twenty-two participants each performed the Paced Auditory Serial Addition Task (PASAT) and a novel variant of this task called the Paced Auditory Serial Subtraction Task (PASST), together with a verb generation task and its two controls, before and after the modulation of cortico-cerebellar connectivity using anodal or cathodal tDCS over the cerebellum. Participants' performance in the difficult PASST task significantly improved after cathodal stimulation compared to sham or anodal stimulation. Improvement in the easier PASAT was equal across all three stimulation conditions. Improvement in verbal response latencies were also greatest during the PASST task after cathodal stimulation, compared to sham and anodal stimulation, and became less variable. Results for the verb generation task complimented those for the PASST, such that the rate and consistency of participants' verbal responses were facilitated by cathodal stimulation, compared to sham and anodal stimulation. These findings suggest that DC stimulation over the right cerebellum affects working memory and attention differently depending on task difficulty. They support a role for the cerebellum in cognitive aspects of behaviour, whereby activity in the prefrontal cortex is likely dis-inhibited by cathodal tDCS stimulation over the right cerebellar cortex, which normally exerts an overall inhibitory tone on the cerebral cortex. We speculate that the cerebellum is capable of releasing cognitive resources by dis-inhibition of prefrontal regions of cerebral cortex, enhancing performance when tasks become demanding.

Cortical control of muscle relaxation: a lateralized readiness potential (LRP) investigation.

OBJECTIVE: We used the lateralized readiness potential (LRP) to investigate cortical mechanisms underlying the termination of muscle contraction. Active suppression and withdrawal of activation have been proposed as underlying mechanisms in isotonic and isometric relaxation. METHODS: Experiment 1 investigated isotonic wrist extension/release from extension. Experiment 2 investigated isometric activation/relaxation of a pinch grip. Tasks were performed with left and right hands and cued auditorily at variable intervals. EEG was recorded from 128 electrodes and processed to derive the LRP timelocked to the onset and offset of muscle contraction. RESULTS: LRPs for isotonic activation and relaxation were of identical amplitude at electrodes overlying the motor cortex, but differed at frontal locations due to higher amplitude re-afferent activity during activation. The isometric LRP was significantly smaller during relaxation than during activation, without differences in scalp distribution. CONCLUSION: The LRP findings confirm differences between isotonic and isometric relaxation, which may be partly explained by the need to suppress a stretch reflex in the former condition. The presence of an LRP associated with isometric relaxation reveals active preparation in the motor cortex, indicating that muscle relaxation in the isometric task cannot be explained solely by withdrawal of activation. SIGNIFICANCE: High-density LRP recordings isolate different cortical mechanisms underlying the termination of muscle contraction.

Force and time control in the production of rhythmic movement sequences in Parkinson's disease.

This study investigated force and time control in rhythm production in Parkinson's disease. Patients and age-matched controls reproduced rhythms consisting of equal (600 ms) or alternating (400 and 800 ms) intervals, produced with equal (12 N) or alternating (8 and 16 N) forces, under paced (synchronization) and unpaced (continuation) conditions. Performance was assessed in terms of accuracy and variability. Reproduction of force amplitudes was less accurate in patients than in controls when either time or force alternated but did not deteriorate further when force and time alternated. In contrast, interval timing was least accurate in patients compared with controls when rhythm alternated on both dimensions. The ratio between alternating force levels was better maintained by controls than by patients, without influence of timing requirements in either group. In contrast, the ratio between alternating intervals was relatively preserved in patients when force remained constant but was severely compromised when force alternated. Timing variability was greater in the patient group, especially in the more affected side, but there was no difference in timing variability between groups as a function of rhythm, and there were no differences between groups in variability of force. The results show that force and time control in patients are affected in different ways. We interpret the preserved timing of rhythms with alternation on one dimension as evidence of intact ability to organize movements in a sequence. The data are compatible with a basal ganglia role in the coregulation of time and force, but do not support a general timing function of the basal ganglia.