Memory deficits during electrical stimulation of the speech cortex in conscious man

Cerebral zones supporting language and recent memory were mapped by electrical stimulation during neurosurgical treatment of epileptic patients. Stimulation of sites within the left posterior temporo-parietal cortex produced transient dysphasia. Continued stimulation of this cortical region also produced a retrograde type of verbal memory disorder, indicating a failure in the mechanism responsible for retrieval of stored information. In contrast, stimulation of the anterior temporal neocortex did not produce anomia and, instead, resulted in an anterograde memory loss, apparently caused by a defect in the verbal storage mechanism. Comparable stimulation of homologous areas on the right hemisphere did not interfere with object naming or immediate verbal recall.     

The role of the human thalamus in language and memory: evidence from electrophysiological studies

The data reviewed here indicate that electrical stimulation of the dominant ventrolateral thalamus can produce deficits in language processing that are not seen after similar stimulation of the nondominant ventrolateral thalamus. The nature of the language deficit produced varies, depending upon the rostrocaudal location of the stimulation site. Stimulation of the anterior left ventrolateral thalamus in right-handed patients resulted in production of a repeated erroneous word, stimulation of the medial ventrolateral thalamus evoked perseveration, and stimulation of the posterior ventrolateral thalamus and anterior pulvinar resulted in misnaming and omissions. Additional studies have examined the effect of electrical thalamic stimulation on verbal and nonverbal short-term memory. Left (but not right) ventrolateral thalamic stimulation during verbal memory input greatly decreased subsequent recall errors, while stimulation during verbal memory retrieval increased recall errors. This finding contrasted with those obtained from studies on nonverbal memory, in which right ventrolateral stimulation during memory input decreased recall errors, while left thalamic stimulation at the same stage increased recall errors. Left pulvinar stimulation disrupted verbal memory processing, while right pulvinar stimulation disrupted nonverbal memory processing. Limited evidence suggests that the effects of thalamic electrical stimulation on verbal memory may persist for several days after the stimulation has ended. The lateralization of thalamic functions also affects the motoric aspects of speech production. Left (but not right) ventrolateral thalamic stimulation disrupted speech articulation and increased the expiratory phase of respiration. The fact that these motor effects were evoked from the same general area of the thalamus that produced the language deficits discussed above raises the possibility that the thalamus is involved in coordinating the cognitive and motoric aspects of language production. A model of thalamic function is discussed in which defined regions of the thalamus operate as a "specific alerting response," increasing the input to memory of category-specific material while simultaneously inhibiting retrieval from memory.     

Common pathways in mental imagery and pain perception: an fMRI study of a subject with an amputated arm

The present paper reviews data from two previous studies in our laboratory, as well as some additional new data, on the neuronal representation of movement and pain imagery in a subject with an amputated right arm. The subject imagined painful and non-painful finger movements in the amputated stump while being in a MRI scanner, acquiring EPI-images for fMRI analysis. In Study I (Ersland et al., 1996) the Subject alternated tapping with his intact left hand fingers and imagining "tapping" with the fingers of his amputated right arm. The results showed increased neuronal activation in the right motor cortex (precentral gyrus) when tapping with the fingers of the left hand, and a corresponding activation in the left motor cortex when imagining tapping with the fingers of the amputated right arm. Finger tappings of the intact left hand fingers also resulted in a larger activated precentral area than imagery "finger tapping" of the amputated right arm fingers. In Study II (Rosen et al., 2001 in press) the same subject imagining painful and pleasurable finger movements, and still positions of the fingers of the amputated arm. The results showed larger activations over the motor cortex for movement imagining versus imagining the hand being in a still position, and larger activations over the sensory cortex when imagining painful experiences. It can therefore be concluded that not only does imagery activate the same motor areas as real finger movements, but also that adding instructions of pain together with imaging moving the fingers intensified the activation compared with adding instructions about non-painful experiences. From these studies, it is clear that areas activated during actual motor execution to a large extent also are activated during mental imagery of the same motor commands. In this respect the present studies add to studies of visual imagery that have shown a similar correspondence in activation between actual object perception and imagery of the same object.     

Right to left hemispheric shift in occipital electroencephalographic responses to repeated Kimura figures

Gordon and Carmon (1976) reported that repeated presentations of an initially novel stimulus were associated with a transfer of cerebral dominance over time (trials) from the right to the left hemisphere. To test the generalizability of these results the proportions of alpha rhythms over the left and right occipital and parietal lobes were measured following the presentation of recurring and nonrecurring complex visual patterns (the Kimura Figures) to the upper or lower, left or right peripheral visual fields. Analysis showed increased electrical activation (as inferred by attenuated proportions of alpha rhythms) of the left occipital lobe but decreased activation of the right occipital lobe. This shift occurred during repeated presentations of the same stimuli but not during single presentations of different novel stimuli. There was no significant shift in this activity over the parietal lobes. These results are consistent with the reports of other researchers who have found a shift of dominant neuroelectrical activity from the right to the left hemisphere as the novelty of a visuospatial stimulus decreases.     

The Use of Non-invasive Brain Stimulation Techniques to Facilitate Recovery from Post-stroke Aphasia

Aphasia is a common symptom after left hemispheric stroke. Neuroimaging techniques over the last 10–15 years have described two general trends: Patients with small left hemisphere strokes tend to recruit perilesional areas, while patients with large left hemisphere lesions recruit mainly homotopic regions in the right hemisphere. Non-invasive brain stimulation techniques such as transcranial magnetic stimulation (TMS) and transcranial direct current stimulation (tDCS) have been employed to facilitate recovery by stimulating lesional and contralesional regions. The majority of these brain stimulation studies have attempted to block homotopic regions in the right posterior inferior frontal gyrus (IFG) to affect a presumed disinhibited right IFG (triangular portion). Other studies have used anodal or excitatory tDCS to stimulate the contralesional (right) fronto-temporal region or parts of the intact left IFG and perilesional regions to improve speech-motor output. It remains unclear whether the interhemispheric disinhibition model, which is the basis for motor cortex stimulation studies, also applies to the language system. Future studies could address a number of issues, including: the effect of lesion location on current density distribution, timing of the intervention with regard to stroke onset, whether brain stimulation should be combined with behavioral therapy, and whether multiple brain sites should be stimulated. A better understanding of the predictors of recovery from natural outcome studies would also help to inform study design, and the selection of clinically meaningful outcome measures in future studies.     

Functional lateralization of the human premotor cortex during sequential movements

A neurological truism is that each side of the brain controls movements on the opposite side of the body. Yet some left hemisphere brain lesions cause bilateral impairment of complex motor function and/or ideomotor apraxia. We report that the left dorsal premotor cortex of normal right-handed people plays a fundamental role in sequential movement of both right and left hands. Subjects performed sequential finger movements during functional magnetic resonance imaging of the motor cortices. In right-handed subjects, the volume of activated dorsal premotor cortex showed a left hemispheric predominance during hand movements. We suggest that the observed left premotor dominance contributes to the lateralization found in lesion studies.     

The cortical representation of centrally presented words: A magnetic stimulation study

The right and left visual fields each project to the contralateral cerebral hemispheres. The current study aimed to investigate the extent of the functional overlap of the two hemifields along the vertical meridian. We applied repetitive transcranial magnetic stimulation (rTMS) over the left and right occipital cortex to investigate whether the foveal representation of words is bilaterally represented or is split between the two hemispheres. Employing a lateralized lexical decision task, we first showed a double dissociation between the stimulated cortical site and performance; right visual field (RVF) but not left visual field (LVF) performance was impaired when the left visual cortex was stimulated, and LVF but not RVF performance was impairred when the right visual cortex was stimulated. Unilateral stimulation also significantly impaired lexical decision latencies to centrally presented words. These findings support the suggestion that foveal representation of words is split. We discuss future strategies for the use of TMS in further tests of the split representation account.     

Failure to develop a conditional reflex by stimulation of the cephalad portion of the severed vagosympathetic trunk: Its significance in terms of the central connections of the vagus

Stimulation of the central stump of either vagosympathetic trunk in the dog, the contralateral nerve remaining intact, regularly provoked deep respiratory movements with forceful expiration, followed by a period of apnea, and a fall in blood pressure, systolic and diastolic, of 20–60 mm/Hg. Stimulation of the cephalad portion of the left nerve provoked brief acceleration of heart rate during the period of hyperventilation, followed by bradycardia; when the stimulus was applied to the central stump of the right nerve heart rate remained relatively unchanged. When a 12-second tone as a conditional stimulus (CS) was reinforced during its last six seconds with such stimulation of the vagosympathetic trunk as an unconditional stimulus (US), despite the striking visceromotor responses elicited by the US, no conditional reflex was established even after more than 3,400 trials in 16 dogs (34–781 trials per animal).     

Dorsolateral prefrontal cortex mediates working memory processes in motor skill learning

Neuroimaging studies have shown that the dorsolateral prefrontal cortex (DLPFC) is recruited during motor skill learning, which suggests the involvement of the DLPFC in working memory (WM) processes, such as selection and integration of motor representations temporarily stored in WM. However, direct evidence linking activation of the DLPFC to WM storage and manipulation during motor skill learning in real-time is rare. In this study, we conducted two experiments to investigate the causal role of DLPFC activity in WM storage and manipulation during motor skill learning under low and high WM-demand conditions. Participants received continuous theta burst stimulation (cTBS) and sham stimulation (crossover design) over the left DLPFC (experiment 1) or right DLPFC (experiment 2). Before and after stimulation, participants in both experiments performed a sequential finger-tapping (SFT) task containing repeated sequence (low-WM demand) and non-repeated sequence (high-WM demand) conditions which are used to study WM processes. The number of correct sequences (NoCS) and reproduction error rate were analyzed. Learning gains in NoCS improved significantly with the practice for both sequence types in the presence of either stimulation type. Compared to sham stimulation, cTBS over the left DLPFC resulted in significantly reduced learning gains in NoCS for non-repeated sequences. These results suggest that the left DLPFC contributes to WM manipulation during motor skill learning.     

Association of Fine Motor Loss and Allodynia in Fibromyalgia: An fNIRS Study

Recent studies showed that fine motor control dysfunction was observed in fibromyalgia (FM) syndrome as well as allodynia. However, brain signatures of this association still remain unclear. In this study, finger tapping task (FTT) and median nerve stimulation (MNS) were applied to both hands of 15 FM patients and healthy controls (HC) to understand this relationship. Hemodynamic activity was measured simultaneously using functional near-infrared spectroscopy (fNIRS). Experiments were analyzed separately by using 2x2 repeated measures ANOVA. Results for the FTT experiment revealed that HC showed higher activity than FM patients in bilateral superior parietal gyrus (SPG), left supramarginal gyrus (SMG) and right somatosensory cortex (SI). Furthermore, right-hand FTT resulted in higher activity than left-hand FTT in left SPG, left SI and right motor cortex (MI). In the MNS experiment, FM patients showed higher activity than HC in bilateral SPG, right SMG, right SI and right middle frontal gyrus (MFG). Negative correlation was observed in left SPG between FTT and MNS activities. Besides, MNS activity in left SPG was negatively correlated with left-hand pain threshold.This study revealed that left SPG might be an important indicator to associate fine motor loss and allodynia in FM.     

Changes in excitability in motor cortex (representation of foreleg) of the dog depending on signal signification of flexor reaction during elaboration of trans switching

Excitability in a cortical representation of a foreleg was determined during elaboration of transswitching in dogs in three variations depending upon the stimulus used as a Conditioned stimulus (CS). If the flexor reaction to direct stimulation of the motor cortex was used as a CS, excitability changes of opposite signs appeared in the cortical representation of the foreleg, the threshold of flexion decreased in the alimentary situation and increased in the defense situation. Such changes in excitability were observed during instrumental conditioned transswitching. If subthreshold stimulation of the motor cortex was used as the CS, the threshold of flexion increased in both situations in approximately equal degree.     

Bilateral Transcranial Direct Stimulation Over the Primary Motor Cortex Alters Motor Modularity of Multiple Muscles

Abstract

Transcranial direct current stimulation (tDCS) over the primary motor cortex (M1) has been demonstrated to modulate the motor performance of both healthy individuals and patients with neuromuscular disorders. However, the effect of tDCS on motor control of multiple muscles, which is a prerequisite to change in motor performance, is currently unknown. Using dimensionality reduction analysis, we investigated whether bilateral tDCS over M1 modulates the coordinated activity of 12 muscles. Fifteen healthy men participated in this randomized, double-blind crossover study. Each participant received a 20-min sham and 2-mA stimulation bilaterally over M1 (anode on the right M1 and cathode on the left M1), with a minimum washout period of 4?days. Muscle activation and end-point kinematics were evaluated during a task where participants reached out to a marked target with non-dominant hand as fast as possible, before and immediately after tDCS application. We found decreased similarity in motor modularity and significant changes in muscle activation in a specific motor module, particularly when reaching out to a target placed within arm’s length and improved smoothness index of movement only following 2-mA stimulation. These findings indicate that clinicians and researchers need to consider the simultaneous effect of bilateral tDCS over M1 on multiple muscles when they establish tDCS protocol to change in motor performance of patients with neuromuscular deficits.     

Lateralization of bilateral transfer of visuomotor information in right-handers and left-handers

The author examined the lateralization of transfer of visuomotor information between the right and left hands during unimanual finger-tapping sequences with visual feedback. The finger-tapping task consisted of a target peak force of 2 N and a target intertap interval of 500 ms. Twenty right-handed and 10 left-handed participants performed the motor task, with 3 transfer trials following 3 practice trials. The author observed positive transfers from the left to the right hand for right-handers but the opposite direction of positive transfers for left-handers. However, left-handers showed a less variable peak force than right-handers did. The author discusses left-handers' interhemispheric information processing.     

Movement as a signal during classical conditioning

Analysis of the available data and that of the author disclosed the peculiarities of motor reaction when used as a conditioned stimulus. The author’s data showed that if signal value is attributed to a motor reaction (passive movement or movement evoked by the direct stimulation of the motor cortex), the changes of excitability in the motor cortex representation of the dog’s leg depend on the biological sign of the reinforcing stimulus during classic conditioning. They also remained the same during instrumental conditioning and were opposite in sign, showed increased excitability in the food situation, and decreased excitability in the defense situation. Using the movement as a conditional stimulus, we managed to uncover the commonality between classic and instrumental conditioning. This enabled us to answer questions, discussed by Pavlov and Guthrie, which, it seems to us, had not been convincingly answered during their time.     

Bimanual coordination and interhemispheric interaction

Bimanual coordination of skilled finger movements requires intense functional coupling of the motor areas of both cerebral hemispheres. This coupling can be measured non-invasively in humans with task-related coherence analysis of multi-channel surface electroencephalography. Since bimanual coordination is a high-level capability that virtually always requires training, this review is focused on changes of interhemispheric coupling associated with different stages of bimanual learning. Evidence is provided that the interaction between hemispheres is of particular importance in the early phase of command integration during acquisition of a novel bimanual task. It is proposed that the dynamic changes in interhemispheric interaction reflect the establishment of efficient bimanual ‘motor routines'. The effects of callosal damage on bimanual coordination and learning are reviewed as well as functional imaging studies related to bimanual movement. There is evidence for an extended cortical network involved in bimanual motor activities which comprises the bilateral primary sensorimotor cortex (SM1), supplementary motor area, cingulate motor area, dorsal premotor cortex and posterior parietal cortex. Current concepts about the functions of these structures in bimanual motor behavior are reviewed.     

Neural correlates of partial transmission of sensorimotor information in the cerebral cortex

Using single neuron recordings in monkey primary motor (MI) cortex, two series of experiments were conducted in order to know whether response preparation can begin before perceptual processing finishes, thus providing evidence for a temporal overlap of perceptual and motor processes.

In Experiment 1, a “left/right, Go/No-Go” reaction time (RT) task was used. One monkey was trained to perform wrist flexion/extension movements to align a pointer with visual targets. The visual display was organized to provide a two-dimensional stimulus: side (an easy discrimination between left and right targets) which determined movement direction, and distance (a difficult discrimination between distal and proximal targets) which determined whether or not the movement was to be made. Changes in neuronal activity, when they were time-locked to the stimulus, were almost similar in the Go and No-Go trials, and when they were time-locked to movement onset, were markedly reduced in No-Go as compared to Go trials.

In Experiment 2, a stimulus-response compatibility (SRC) task was used. Two monkeys were trained to align a pointer with visual targets, on either left or right. In the spatially “compatible” trials, they had to point at the stimulus position, whereas in the “incompatible” trials, they had to point at the target located in the opposite side. For 12.5% of neurons, changes in activity associated with incompatible trials looked like changes in activity associated with movements performed in the opposite direction during compatible trials, thus suggesting the hypothesis of an automatic activation of the congruent, but incorrect response.

Results of both experiments provide evidence for a partial transmission of information from visual to motor cortical areas: that is, in the No-Go trials of the first task, information about movement direction, before the decision to perform or not this movement was made, and, in the incompatible trials of the SRC task, information about the congruent, but incorrect response, before the incongruent, but correct response was programmed.     

Asymmetrical transfer of training between hands: implications for interhemispheric communication in normal brain

Two experiments are presented which investigated claims of asymmetrical transfer of training between the hands/hemispheres. In Experiment 1, 96 right- and left-handed male undergraduates practiced an inverted-reversed printing task with either the right or the left hand. Transfer to the opposite hand was then compared to same-hand transfer, in a between-subject design. In Experiment 2, 176 right-handed boys and girls were tested at ages 7, 9, and 11 years. For right-handed subjects in both experiments, the left hand benefited more from opposite-hand training than did the right. The reverse was true for left-handers in Experiment 1, although one group (who wrote with the "inverted" position) showed little transfer in either direction. Two current models of interhemispheric interaction do not satisfactorily explain these findings. A third model, based on cross-activation, may provide a more effective alternative.     

The effect of bicephalic stimulation of the dorsolateral prefrontal cortex on the attentional bias for threat: A transcranial direct current stimulation study

Previous stimulation studies demonstrated that the dorsolateral prefrontal cortex (DLPFC) is involved in threat processing. According to a model of emotional processing, an unbalance between the two DLPFCs, with a hyperactivation of right frontal areas, is involved in the processing of negative emotions and genesis of anxiety. In the present study, we investigated the role of the right and left DLPFC in threat processing in healthy women who also completed the State-Trait Anxiety Inventory (STAI). We simultaneously modulated the activity of the right and left dorsolateral prefrontal cortex by applying bicephalic transcranial direct current stimulation (tDCS) before participants completed a modified version of the classic Posner task using threatening and nonthreatening stimuli as spatial cues. Anodal stimulation on the right DLPFC with a simultaneous cathodal stimulation over the left side induced a disengagement bias in individuals with low STAI scores and a facilitation bias in individuals with high STAI scores. Anodal stimulation on the left DLPFC with the simultaneous cathodal stimulation over the right side did not affect threat processing. The findings of the present study provided specific support to the hypothesis that unbalanced activation between left and right hemispheres with enhanced activation of the right DLPFC is critical in early top-down threat processing in healthy individuals.     

Movement planning and attentional control of visuospatial working memory: evidence from a grasp-to-place task

In this study, we have investigated the influence of available attentional resources on the dual-task costs of implementing a new action plan and the influence of movement planning on the transfer of information into visuospatial working memory. To approach these two questions, we have used a motor–memory dual-task design in which participants grasped a sphere and planned a placing movement toward a left or right target according to a directional arrow. Subsequently, they encoded a centrally presented memory stimulus (4 × 4 symbol matrix). While maintaining the information in working memory, a visual stay/change cue (presented on the left, center or right) either confirmed or reversed the planned movement direction. That is, participants had to execute either the prepared or the re-planned movement and finally reported the symbols at leisure. The results show that both, shifts of spatial attention required to process the incongruent stay/change cues and movement re-planning, constitute processing bottlenecks as they both reduced visuospatial working memory performance. Importantly, the spatial attention shifts and movement re-planning appeared to be independent of each other. Further, we found that the initial preparation of the placing movement influenced the report pattern of the central working memory stimulus. Preparing a leftward movement resulted in better memory performance for the left stimulus side, while the preparation of a rightward movement resulted in better memory performance for the right stimulus side. Hence, movement planning influenced the transfer of information into the capacity-limited working memory store. Therefore, our results suggest complex interactions in that the processes involved in movement planning, spatial attention and visuospatial working memory are functionally correlated but not linked in a mandatory fashion.     

Transfer of short-term motor learning across the lower limbs as a function of task conception and practice order

Interlimb transfer of motor learning, indicating an improvement in performance with one limb following training with the other, often occurs asymmetrically (i.e., from non-dominant to dominant limb or vice versa, but not both). In the present study, we examined whether interlimb transfer of the same motor task could occur asymmetrically and in opposite directions (i.e., from right to left leg vs. left to right leg) depending on individuals’ conception of the task. Two experimental conditions were tested: In a dynamic control condition, the process of learning was facilitated by providing the subjects with a type of information that forced them to focus on dynamic features of a given task (force impulse); and in a spatial control condition, it was done with another type of information that forced them to focus on visuomotor features of the same task (distance). Both conditions employed the same leg extension task. In addition, a fully-crossed transfer paradigm was used in which one group of subjects initially practiced with the right leg and were tested with the left leg for a transfer test, while the other group used the two legs in the opposite order. The results showed that the direction of interlimb transfer varied depending on the condition, such that the right and the left leg benefited from initial training with the opposite leg only in the spatial and the dynamic condition, respectively. Our finding suggests that manipulating the conception of a leg extension task has a substantial influence on the pattern of interlimb transfer in such a way that the direction of transfer can even be opposite depending on whether the task is conceived as a dynamic or spatial control task.