Electroencephalographic and reaction time asymmetries to musical chord stimuli

EEG was recorded over left and right hemispheres at temporal leads (T3, T4) referred to the vertex (Cz) in 16 right-handed male subjects. Musical chords were presented randomly in monaural sequence, during a task which required a selective motor response to stimuli presented to one ear. The integrated amplitude of the 8-13 Hz alpha rhythm was measured when subjects listened passively. Under all conditions, a lower mean alpha amplitude was recorded over the right hemisphere than the left, regardless of which ear was stimulated. Alpha suppression over the right temporal area was accentuated when the performance task directed attention to the stimulus. Reaction time to left ear stimulation was shorter than that for the right ear. With monaural stimulus presentation behavioral asymmetry, and various EEG asymmetries can be observed. There is hemispheric asymmetry associated with attention to task relevant stimuli indicated by reduction in the alpha rhythm over the right temporal area and asymmetry in reaction time with greater efficiency of the left ear to muscial chords.     

The processing of sound duration after left hemisphere stroke: Event-related potential and behavioral evidence

The ability of left-hemisphere stroke patients ( n = 8) and healthy control subjects ( n = 8) to process sounds preattentively and attentively was studied by recording auditory event-related potentials (ERPs) and behavioral responses. For the right-ear stimulation, the mismatch negativity (MMN) was significantly smaller in the patients than control subjects over both hemispheres. For the left-ear stimuli, the MMN was significantly smaller in the patient group than in the control group over the left hemisphere, whereas no group differences were obtained over the right hemisphere. In addition, the N1 amplitude was reduced bilaterally for the right-ear stimulation (with the reduction being larger over the left hemisphere), whereas no significant effects on the N1 amplitude were found for the left-ear stimulation. Behaviorally, the patients detected significantly fewer deviant tones than did the control subjects irrespective of the stimulated ear. The present results thus suggest that the long-latency ERPs can be used to probe such auditory processing deficits that are difficult to define with behavioral measures. Especially by recording MMN to monaural stimuli, the discrimination accuracy can be separately determined for the left and right temporal lobes.     

Hemispheric Asymmetry and Human Electrodermal Conditioning: The Dichotic Extinction Paradigm

It was hypothesized that classically conditioned responses to an auditory verbal CS initially presented only to the left cerebral hemisphere would be greater than when the same CS was presented to the right hemisphere. The experiment consisted of three different phases in a dichotic extinction paradigm. During the habituation phase the CS+ and CS? were presented binaurally, and separated. All subjects were treated alike. During the acquisition phase the conditioning subjects had the CS+ but not the CS? followed by the UCS. The CS+ and CS? were presented binaurally and separated. The control subjects received no UCS presentations. During the dichotic extinction phase, half of the conditioned subjects had the CS+ presented in the right ear (i.e., with initial left hemisphere input), with the CS? simultaneously presented in the left ear (i.e., with initial right hemisphere input). The other half had the earphones reversed. The control group was similar to the two conditioning groups. During the acquisition phase, significantly larger electrodermal responses were observed to the CS+ compared to the CS? in the conditioning groups, but not in the control group. During the dichotic extinction phase, significantly more resistance to extinction was observed in the CS+ Right-Ear group compared to the CS+ Left-Ear group, with the control group revealing virtually no responding at all. It was concluded that the present results have demonstrated effects of cerebral asymmetry on rate of extinction of the classically conditioned response.     

Effects of Cigarette Smoking on Bilateral Skin Conductance

Bilateral skin conductance was recorded from 8 male and 8 female cigarette smokers and from 8 male and 8 female nonsmoker control subjects on two different days. On one day smokers smoked a cigarette prior to the recording session and on the other they remained abstinent. Subjects also engaged in tasks designed to differentially involve the cerebral hemispheres. SCR asymmetry was measured using two different indices. During both the resting and task periods smokers had larger SCRs in their right than left hands following smoking. Unlike control subjects during the first recording session, smokers did not show task related changes in SCR asymmetry. During tasks smokers showed smaller SCRs in both hands following smoking than following abstinence. Changes in SCR asymmetry associated with smoking were seen in subjects who smoked more cigarettes, reported more subjective craving, were more introverted, and indicated a preference for cognitive processes and strategies associated with the left hemisphere. Results are discussed in terms of relative involvement of the cerebral hemispheres following smoking and smoking abstinence.     

Visual evoked response correlates of cerebral specialization after human commissurotomy

Visual evoked responses (VERs) were recorded from commissurotomy patients and normal subjects in order to investigate the electrophysiological correlates of cerebral lateralization and independent hemispheric processing. A verbal task consisted of detecting rhyming words and a spatial task consisted of comparing matching shapes. Stimuli were delivered, by means of a computer graphic display, into the right visual field, the left visual field, or both visual fields. Behavioral results and evoked response waveforms showed lateralization for verbal processing in the left hemisphere for the patient group. Analysis of variance of product moment correlation coefficients between VER waveforms indicated significant differences between unilateral and bilateral stimulation for both patients and normals. Commissurotomy and normal VERs showed similar patterns of correlation when verbal stimuli were delivered to the left hemisphere, but differed during right hemisphere stimulation. The results are discussed in terms of VER correlates of visual-verbal processing.     

An investigation into asymmetrical cortical regulation of salivary S-IgA in conscious man using transcranial magnetic stimulation.

Transcranial magnetic stimulation (TMS) can activate discrete areas of the cerebral cortex through the intact skull of healthy conscious volunteers. A magnetic coil generates a brief and focused magnetic field that penetrates the skull to activate the specific area of cerebral cortex beneath. This non-invasive procedure is painless, well tolerated by participants and now widely used to explore brain function. We used TMS to investigate asymmetrical cortical regulatory influences on one aspect of immune function: secretion of the antibody immunoglobulin A (S-IgA) into saliva. The right and left temporo-parietal-occipital cortex of 16 healthy, conscious subjects was stimulated on two different occasions, at least 1 week apart. There was an immediate increase in S-IgA concentration following both right and left stimulation. Saliva volume was reduced immediately post-right but not left stimulation. When secretion (microg/min) of S-IgA was calculated (controlling for changing saliva volume) an increase was apparent following left but not right hemisphere stimulation. Furthermore, a significant difference between the relationship between S-IgA concentration and volume of saliva post-left and right stimulation was observed. We conclude that TMS can be used as a tool to investigate cortical regulation of autonomic and immune function in healthy, conscious human subjects and that secretion of saliva and S-IgA is differentially affected by stimulation of the left and right cerebral cortex.     

Phonological and semantic fluencies are mediated by different regions of the prefrontal cortex

Verbal phonological and semantic fluencies were investigated in 24 patients with unilateral prefrontal lesions and 10 normal control subjects. Lesions were limited to small areas within either the dorsolateral (Brodmann's area 46/9) or ventromedial (posterior part of the gyrus rectus) cortices. In a phonological fluency task, patients with lesions to the left dorsolateral region were impaired. In semantic fluency, not only the left dorsolateral group but also the two right frontal damaged groups performed worse than the control group. In agreement with previous studies, our results show that the phonological fluency is mediated by the left dorsolateral prefrontal cortex. In contrast to this, performance on the semantic fluency task depends on a wider portion of the prefrontal cortex involving the left and right dorsolateral and the right ventromedial areas.     

Fatigue suppresses ipsilateral intracortical facilitation

Experimental data in animals and humans have demonstrated connections between right and left motor cortices. Interactions between these cortical areas can be explored with electrical or magnetic stimulation. In the present study we examined the interhemispheric effect of fatigue on intracortical facilitation (ICF) and inhibition (ICI) using a paired-pulse transcranial magnetic stimulation (TMS) paradigm. Ten healthy subjects performed pinch grips with their left hand with 50% maximum voluntary contraction (MVC) until fatigue occurred. In the control experiment, the same number of pinch grips was performed with 5% MVC without inducing fatigue. Motor evoked potentials (MEP) produced by single and paired pulse TMS over the left motor cortex were recorded from right first dorsal interosseous muscle (FDI) and right abductor digiti minimi muscle (ADM) before and after the tasks. ICF of the right FDI was significantly reduced after fatigue ( P=0.0008). Fifteen minutes after finishing the task ICF had returned to baseline values. There was no change of ICF of right FDI in the control experiment without inducing fatigue. In both experiments the right ADM did not show significant MEP changes. Additional control experiments showed that M-responses and F-waves were unchanged in right FDI after performing the fatigue task with left FDI, and TMS test pulse amplitudes were significantly reduced in left FDI after fatigue. Fatigue caused by pinch grips induces a short-lasting and task-specific suppression of intracortical facilitation in the motor cortex of an homologous contralateral hand muscle. These results indicate interhemispheric interactions between the two motor cortices that are still effective after cessation of movements.     

Single-pulse transcranial magnetic stimulation of parietal and prefrontal areas in a memory delay arm pointing task

Fifteen healthy volunteers performed a memory-pointing task using their right arm while single-pulse transcranial magnetic stimulation (TMS) above motor threshold was applied over the posterior parietal or prefrontal cortex of the left or right hemisphere in four blocks of trials. The stimulation was randomly delivered at one of three time intervals during the 3-s delay period (early: 300 ms, intermediate: 1,500 ms, late: 2,700 ms). A separate block with no stimulation was used as control. Only early left parietal stimulation resulted in an increase in the variance of movement amplitude but not direction for all targets in two-dimensional space (both hemifields). The results point to the significance of the contralateral posterior parietal cortex early on during the memorization of the target for an upcoming movement. Taking into consideration the limitations of TMS and those imposed by the particular task, the lack of specific effects of prefrontal stimulation provides evidence that these areas might not be involved in the performance of simple memorized arm movements.     

Hemispheric specialization in the co-ordination of arm and trunk movements during pointing in patients with unilateral brain damage

During pointing movements involving trunk displacement, healthy subjects perform stereotypically, selecting a strategy in which the movement is initiated with either the hand or trunk, and where the trunk continues after the end of the hand movement. In a previous study, such temporal co-ordination was not found in patients with left-hemispheric brain lesions reaching with either their dominant paretic or with their non-dominant non-paretic arm. This co-ordination deficit may be associated in part with the presence of a lesion in the dominant left hemisphere. If so, then no deficit should be observed in patients with stroke-related damage in their non-dominant right hemisphere moving with their ipsilesional arm. To verify this, 21 right-hand dominant adults (7 who had had a stroke in the right hemisphere, 7 who had had a stroke in the left hemisphere and 7 healthy subjects) pointed to two targets located on a table in front of them in the ipsilateral and contralateral workspace. Pointing was done under three movement conditions: while not moving the trunk, while bending the trunk forward and while bending the trunk backwards. The experiment was repeated with the non-paretic arm of patients with stroke and for the right and left arms of healthy subjects. Kinematic data were recorded (Optotrak). Results showed that, compared to healthy subjects, arm-trunk timing was disrupted in patients with stroke for some conditions. As in patients with lesions in the dominant hemisphere, arm-trunk timing in those with lesions in the non-dominant hemisphere was equally more variable than movements in healthy subjects. However, patients with dominant hemisphere lesions used significantly less trunk displacement than those with non-dominant hemisphere lesions to accomplish the task. The deficit in trunk displacement was not due to problems of trunk control or sitting balance since, in control experiments, all subjects were able to move the trunk the required distance, with and without the added weight of the limb. Results support the hypothesis that the temporal co-ordination of trunk and arm recruitment during pointing movements is mediated bilaterally by each hemisphere. However, the difference in the range of trunk displacement between patients with left and right brain lesions suggests that the left (dominant) hemisphere plays a greater role than the right in the control of movements involving complex co-ordination between the arm and trunk. Electronic Publication     

Planning Ability After Frontal and Temporal Lobe Lesions in Humans: The Effects of Selection Equivocation and Working Memory Load

Twenty-one patients with unilateral frontal neurosurgical lesions (11 left and 10 right) and 38 patients with unilateral temporal lobectomies (19 left and 19 right) were compared to 44 normal control subjects in their performance on a computerised version of the Tower of Hanoi (TOH) test. In order to explore the effect of selection equivocation on performance, two types of problems were administered, determined by whetherthe two mainroutes to achieve the goal weresimilar or dissimilar in length. To explore the effectof working memory load, a task was administered that required the subject to repeat a sequence of Tower of Hanoi moves shown by the computer. For the frontal patients, those with right (RF), but not left (LF) lesions were impaired on TOH, but response equivocation had no effecton performance. The LF group, however, were impaired on the working memory task. Forthetemporal patients, only therightlesion group wereimpaired on TOH, but specifically on the dissimilar problems. This group were also impaired on the working memory task. Selection equivocation and working memory load are not thought to be significant factors in producing a planning deficit following frontal lobe lesions, but may cause impairment in patients who have undergone right temporal lobectomies.     

Asymmetrical hemispheric EEG activation evoked by stimulus position during the Simon task

The Simon effect has been previously shown to be asymmetric at both the behavioral and electrophysiological levels. The present investigation was aimed to clarify whether, during a Simon task, hemispheric asymmetry is also observed in the early phases of stimulus processing. In a group of healthy subjects performing the Simon task, we analyzed scalp potentials evoked by the first lateralized cue (left or right), instead of the classical readiness potential preceding the motor response. ERP results showed a significant left cortical activation to stimuli presented in the right visual field at the 140–160 ms time window. Instead, left stimuli elicited a significant activation of the right versus left hemisphere starting at the next 160–180 ms time interval. We linked this asymmetry to that observed in behavioral data: the Simon effect recorded with left stimuli is smaller than the Simon effect recorded with right stimuli. Results confirm the hypothesis that in right handed subjects, left hemisphere is specialized for motor response selection and is able to process right stimuli faster than the right hemisphere does for left stimuli.     

Asymmetries of vestibular dysfunction in major depression

Depression is characterized by alterations in the circadian secretion of hormones, sleep and motor activity, all of which are regulated by suprachiasmatic nuclei (SCN). The vestibular system in the inner ear registers the amount of motor activity. To test the integrity of this motion sensitive system in depression, we studied the vestibulo-ocular reflex (VOR) in depressive patients who were not taking medication and healthy control subjects, which allowed us to investigate each ear and its corresponding nerve centers. Ocular reflex movement depends on vestibular nuclei activity, and we found that at 30 degrees C stimulation the right vestibular system in depressive patients has approximately half the activity of the left side. Significant asymmetry was not detected in control subjects. We also found a significant decrease in the slow phase (16.92+/-9.13 degrees/s) of the reflex in the depressed group as compared with the control group (43.77+/-16.04 degrees/s). The vestibular nuclei of the right and left sides are hypoactive. Specifically, the right vestibular nucleus is hypoactive in depressed people and can easily be measured using VOR. These results support the abnormal asymmetries hypothesis of depression and suggest that these asymmetries also exist at the level of the brain stem or neuronal centers that are afferents to right vestibular nuclei, like SCN or raphe nuclei.     

The role of the dorsolateral prefrontal cortex in retrieval from long-term memory depends on strategies: a repetitive transcranial magnetic stimulation study

The ability to associate a name to a face is a crucially relevant task in daily life. In this study, we investigated the neuronal basis of face-name retrieval in young subjects using repetitive transcranial magnetic stimulation (rTMS) over the left or right dorsolateral prefrontal cortex (DLPFC). The experimental task was composed of two study phases: an encoding phase and a retrieval phase. During the encoding phase, subjects saw a face (familiar or unfamiliar) followed by a name. During the retrieval phase, they saw the face together with two names and had to choose the name that was correctly associated with the face. rTMS was delivered only during retrieval. In addition, we evaluated the use of memory strategies during the task. Accordingly, subjects were subdivided into two groups: strategy users (SU) and no-strategy users (NSU). No rTMS effects were present for familiar face-name pairs, probably due to a ceiling effect. However, for unfamiliar face-name pairs, the different use of memory strategies resulted in different rTMS effects. The SU group showed a selective interference effect after right DLPFC stimulation, whereas the NSU group showed an effect after left DLPFC stimulation. Importantly, the overall performance of the two groups was comparable. We suggest that during memory retrieval the left DLPFC might be recruited when the subject does not apply deliberately a retrieval strategy whereas there is a shift to the right DLPFC if cognitive control processes that are engaged by strategies are needed to guide episodic retrieval.     

Effects of probability bias in response readiness and response inhibition on reaching movements

It is solidly established that unequal stimulus frequencies lead to faster responses to the more likely stimulus; however, the effect of this probability bias on response inhibition is still debated. To tackle this issue, we administered two versions of the stop-signal task to 18 right-handed healthy subjects. In one version, we manipulated the frequency of right and left targets appearance when subjects were required to produce speeded responses (no-stop trials) with the right arm, whereas stop signals occurred with equal frequencies after right or left targets (no-stop signal bias). In the other version, we manipulated the frequency of appearance of stop signals after right or left targets, whereas no-stop trials toward right or left targets had the same frequency (stop-signal bias). Surprisingly, we found a very modest, if any, increase in response readiness toward the more frequent stimulus. However, the no-stop signal bias had an effect on the speed of inhibitory control, as subjects were always faster to suppress a movement toward the side where targets were less likely to occur. Differently, the stop-signal bias had a much more powerful effect. In fact, subjects were faster to withhold movements toward the side where targets were more frequent, while they exhibited longer reaction times for reaches toward the more likely targets. Overall, these results suggest that action preparation and action inhibition are independent competing processes, but subjects tend to place automatically greater importance on the stop task.     

Comparative study of event-related potentials and positron emission tomography activation during a paired-associate memory paradigm

Event-related potentials (ERP) and regional cerebral blood flow (rCBF) activation using ¹⁵O-labeled water associated with retrieval and retention of episodic memory were studied during a visual paired-association task with delayed response in eight healthy subjects. In both studies, the subjects memorized four pairs of figures during the learning period. They were presented with each cue (S1) and asked to judge whether the following figure (S2) formed one of the memorized pairs. In an attempt to identify brain activity related to memory function, a choice reaction task with delay was used as a behavioral control. The ERP study showed a posterior positive component in the difference waveform, which was obtained by subtracting responses in the choice reaction task from those in the paired association task, between 300 and 850 ms after S1 presentation. It was maximal at the parietal midline electrode and distributed predominantly over the left posterior quadrant of the scalp. The rCBF activation study showed a greater increase in rCBF in the right dorsolateral prefrontal cortex (Brodmann’s area 46), left inferior frontal cortex (Brodmann’s area 44/45), left thalamus, and bilateral cerebellar hemisphere during the paired association task as compared to the choice reaction task, which suggests a possible involvement of cerebello-thalamo-cortical circuit in the memory processing. Additionally, it is suggested that the scalp distribution of the ERP component may not necessarily represent regional cortical activation below the electrodes where such a component is observed and could indirectly represent activation in remote areas such as subcortical regions. It seems that ERP and rCBF activation may provide information about different aspects of higher brain function. Received: 19 July 1996 / Accepted: 22 August 1997     

Hemodynamic evoked response of the sensorimotor cortex measured noninvasively with near-infrared optical imaging

We have performed a noninvasive bilateral optical imaging study of the hemodynamic evoked response to unilateral finger opposition task, finger tactile, and electrical median nerve stimulation in the human sensorimotor cortex. This optical study shows the hemoglobin-evoked response to voluntary and nonvoluntary stimuli. We performed measurements on 10 healthy volunteers using block paradigms for motor, sensory, and electrical stimulations of the right and left hands separately. We analyzed the spatial/temporal features and the amplitude of the optical signal induced by cerebral activation during these three paradigms. We consistently found an increase (decrease) in the cerebral concentration of oxy-hemoglobin (deoxy-hemoglobin) at the cortical side contralateral to the stimulated side. We observed an optical response to activation that was larger in size and amplitude during voluntary motor task compared to the other two stimulations. The ipsilateral response was consistently smaller than the contralateral response, and even reversed (i.e., a decrease in oxy-hemoglobin, and an increase in deoxy-hemoglobin) in the case of the electrical stimulation. We observed a systemic contribution to the optical signal from the increase in the heart rate increase during stimulation, and we made a first attempt to subtract it from the evoked hemoglobin signal. Our findings based on optical imaging are in agreement with results in the literature obtained with positron emission tomography and functional magnetic resonance imaging.     

Cerebral blood flow changes associated with experimental pain stimulation in patients with major depression

BACKGROUND: The clinical relationship between pain and depression has been extensively reported. The purpose of this study was to compare the cerebral blood flow (CBF) of patients with major depressive disorder (MDD) during stimulation with experimental pain tolerance or sham stimulation, before and after 2 weeks of at least partially effective antidepressant treatment (ADT), in order to determine the cerebral regions associated with pain processing in the two clinical states. METHODS: Twenty-four antidepressant-free outpatients diagnosed with MDD (DSM-IV), without any pain complaints and a basal score>or=20 points on the Hamilton Rating Scale for Depression were included. Cerebral SPECTs were performed before and after ADT. Patients were stimulated with pain pressure tolerance (PT) or sham stimulation during the radiotracer cerebral uptake time. RESULTS: The comparison between PT and sham stimulation before ADT showed an increase of CBF of PT stimulated patients in right temporal gyrus, left amygdale, right anterior cingulated cortex, bilateral medial frontal gyrus, bilateral insula, lingual gyrus, right precentral gyrus and left postcentral gyrus. Equal comparison after ADT showed an increase of CBF of PT stimulated patients only in left middle frontal gyrus. LIMITATIONS: The sample includes exclusively outpatients with mild-moderate depression. CONCLUSION: CBF before ADT increases in brain areas related with the affective and cognitive components of pain; in contrast, after ADT increases only in cognitive pain related areas. These results offer new avenues to investigate the cerebral substrate of the common relationship between pain and depression.     

Failure to control prepotent pathways in early stage dementia of the Alzheimer's type: evidence from dichotic listening

The authors examined the right ear advantage in a dichotic listening task in healthy aging and very mild and mild stages of Alzheimer's disease. Subjects were simultaneously presented 3 pairs of digits to the left and right ears (e.g., left ear: 4, 3, 1; right ear: 9, 2, 5) for immediate ordered recall. Four lists of triads were presented, varying in presentation rate between digit pairs within a triad (0.5, 1.0, 1.5, 2.0 s). Results indicated that the very mild and mild Alzheimer's groups showed a larger right ear advantage in free recall compared with the healthy controls, indicating a tendency to respond to the prepotent left hemisphere pathway for language processing. Also, the right ear advantage and proportion of switches made during recall were correlated with psychometric measures of frontal lobe function in the mild Alzheimer's group but not in the very mild or healthy control groups.     

Podokinetic after-rotation following unilateral and bilateral podokinetic stimulation

Previous studies demonstrated an aftereffect of walking on a rotating treadmill, involving inadvertent circular navigation with eyes closed [podokinetic after-rotation (PKAR)]. We compared PKAR following unilateral and bilateral podokinetic (PK) stimulation to determine whether the left and right legs could be independently adapted. Each subject performed two sessions of PK stimulation, stepping in place with one foot on either side of the axis of a rotating disk. Subjects experienced bilateral stimulation (i.e., both left and right feet stepped on the rotating disk) in one session and unilateral stimulation (i.e., the left foot stepped on the rotating disk and the right foot stepped on a stationary surface) in the other. Following stimulation, we recorded foot lift-off and touchdown times and pelvic angular velocity while subjects stepped in place on a stationary surface. PKAR velocity following unilateral stimulation was lower than that following bilateral stimulation. Following bilateral stimulation, pelvic rotation was in the counterclockwise (CCW) direction during single-limb support on both the left and right sides. Immediately following left unilateral stimulation, subjects demonstrated CCW pelvic rotation during left single-limb support but not during right single-limb support. Across the first 13 strides, the difference between left and right sides diminished; pelvic angular velocity was then CCW during single-limb support on both sides. This suggests that both the adapted left and the unadapted right limb influenced the final PKAR response with information from the two limbs being integrated over the first few strides.