[show abstract][hide abstract] ABSTRACT: Little is known about how the hemispheres interact in processing of stimuli presented at vertical midline. Processing might be mutually independent or cooperative. Here we measured target identification and visually evoked EEG potentials while stimulus streams containing two targets, T1 and T2, were either presented at vertical midline above and below fixation, or laterally, left and right. With left and right streams, potentials evoked by filler stimuli and by T2 were earlier at the right than the left visual cortex, and T2 was better identified left than right, confirming earlier results and suggesting better capabilities of the right hemisphere in this task. With streams above and below fixation, EEG potentials evoked by filler stimuli and by T2 were likewise earlier at the right than the left hemisphere, and T2 was generally identified as well as, but not better than left T2, in one target constellation even worse (T2 in lower stream preceded by T1 in upper stream). These results suggest right-hemisphere preference for this task even with stimuli at vertical midline, and no added value through hemispheric cooperation. Lacking asymmetry for T1 amidst asymmetries for filler stimuli and for T2 might indicate alternating access of the hemispheres to midline stimuli as one means of hemispheric division of labor.
PLoS ONE 01/2013; 8(2):e57421. · 3.73 Impact Factor
[show abstract][hide abstract] ABSTRACT: The right hemisphere has been shown to play a dominant role in processing of visuo-spatial information. Recently, this role has been studied in the two-stream rapid serial visual presentation task. In this task, two alphanumerical targets are embedded in left and right simultaneous streams of rapidly changing letters. The second target (T2) is identified better in the left than in the right visual field. This difference has been interpreted as advantage of the right hemisphere (RH). However, a disadvantage of the left hemisphere (LH) could not be excluded so far. The LH, specialized for processing of verbal stimuli, might be overloaded due to constant input of letters from both visual fields. In the present study, this overload hypothesis was tested by reducing demands on verbal processing (Experiment 1), and by overloading the RH with non-verbal stimuli: faces (Experiment 2) and irregular shapes (Experiment 3). The left visual field advantage proved to be largely independent from the level of verbal load and from stimulus type. Therefore, although not entirely disproving the overload hypothesis, these results suggest as the most parsimonious explanation this asymmetry reflects a RH advantage, presumably in perceptual and attentional processing, rather than a LH disadvantage caused by verbal overload.
[show abstract][hide abstract] ABSTRACT: In many rapid serial presentation tasks, two targets ("T1" and "T2") have to be distinguished from background stimuli. Here, digits ("lures") were interspersed among the background letters, differing from the T2 digit by occurring before rather than after T1. The resulting inhibitory effects on T2 identification may either be evoked directly by the lures or be triggered by T1, interfering with positive priming of lures on T2. To distinguish between these two alternatives, lures, T1, and T2 were presented in two different simultaneously running streams, T2 was or was not the same digit as a lure, and EEG potentials related to lures, T1, and T2 were recorded. Effects on T2 identification better fit the view that lures exerted positive priming interrupted by T1. Recurrence of lures in the trial led to abridged duration of the lure-evoked N2pc, and T2-evoked N2pc was reduced after lures. Also these N2pc effects may reflect positive priming.
[show abstract][hide abstract] ABSTRACT: A most sensitive and specific electrophysiological indicator of selective processing of visual stimuli is the N2pc component. N2pc is a negative EEG potential peaking 250 ms after stimulus onset, recorded from posterior sites contralateral to relevant stimuli. Additional deflections preceding or following N2pc have been obtained in previous studies, possibly produced by specific stimulus features or specific prime-target sequences. To clarify the entire time-course of the contralateral- ipsilateral (C-I) difference recorded from the scalp above visual cortex in response to left-right pairs of targets and distracters, C-I differences were here compared between two types of stimuli and between stimuli that were or were not preceded by masked neutral primes. The C-I difference waveform consisted of several peaks, termed here P1pc (60-100 ms after target onset), N1pc (120-160 ms), N2pc (220-280 ms), and N3pc (360-400 ms). Being markedly enhanced when stimuli were preceded by the neutral primes, P1pc may indicate a response to stimulus change. Also, when stimuli were primed, N2pc reached its peak earlier, thereby tending to merge with N1pc. N3pc seemed to increase when target discrimination was difficult. N1pc, N2pc, and N3pc appear as three periods of one process. N3pc probably corresponds to L400 or SPCN as described in other studies. These observations suggest that the neurophysiological basis of stimulus-driven focusing of attention on target stimuli is a process that lasts for hundreds of milliseconds, with the relevant hemisphere being activated in an oscillating manner as long as required by the task.
Advances in Cognitive Psychology 01/2012; 8(1):19-28.
[show abstract][hide abstract] ABSTRACT: The number reduction task (NRT) allows us to study the transition from implicit knowledge of hidden task regularities to explicit insight into these regularities. To identify sleep-associated neurophysiological indicators of this restructuring of knowledge representations, we measured frequency-specific power of EEG while participants slept during the night between two sessions of the NRT. Alpha (8-12 Hz) EEG power during slow wave sleep (SWS) emerged as a specific marker of the transformation of presleep implicit knowledge to postsleep explicit knowledge (ExK). Beta power during SWS was increased whenever ExK was attained after sleep, irrespective of presleep knowledge. No such EEG predictors of insight were found during Sleep Stage 2 and rapid eye movement sleep. These results support the view that it is neuronal memory reprocessing during sleep, in particular during SWS, that lays the foundations for restructuring those task-related representations in the brain that are necessary for promoting the gain of ExK.
Journal of Cognitive Neuroscience 08/2011; 24(1):119-32. · 4.49 Impact Factor
[show abstract][hide abstract] ABSTRACT: In patients with the callosal type of anarchic-hand syndrome, the left hand often does not act as intended and counteracts the right hand. Reports are scarce about the underlying neurophysiological mechanisms. We report the case G.H. who developed the syndrome after infarction of the left arteria pericallosa. It has been suggested that the syndrome arises out of lacking inhibition from the dominant left hemisphere on the right hemisphere. Yet, in tests of spatial intelligence G.H. performed much better with his "anarchic" left hand than with his dominant right hand, similar to observations commonly reported in split-brain patients. Left-right manual choice responses and event-related EEG potentials to laterally presented stimuli were measured. Asymmetries were evident in G.H.'s behavior and EEG potentials, different from age-matched healthy participants (n=11). His right-hand responses were fast and unaffected by incompatibility with stimulus location, whereas his left-hand responses were variable and accompanied by a large negative central-midline EEG potential, probably reflecting efforts in initiating the response. G.H.'s visual N1 component peaked earlier and was larger at the right than the left side of the scalp, and the P3 component was distinctly reduced at the right side. Both features occurred independent of side of stimulus presentation and side of responding hand. The effort indicated by the midline negativity and the asymmetrically reduced P3 might directly reflect G.H.'s lack of control on his right hemisphere's processing. The faster visual processing of the right hemisphere suggested by the N1 asymmetry might contribute to "anarchic" processing, making the right hemisphere process stimuli before control impulses exert their effect. These neurophysiological results tend to support the split-brain account which assumes that the syndrome arises by the lack of communication between hemispheres that act according to their respective competences.
Brain and Cognition 06/2011; 77(1):138-50. · 2.82 Impact Factor
[show abstract][hide abstract] ABSTRACT: When two letter streams containing two targets (T1 and T2) are presented left and right, T2 is better identified in the left hemifield. This study on 16 healthy participants used evidence from ERPs to decide whether this advantage is due to better processing in the right hemisphere or to overload of the left one. N2pc and P3 components evoked by T1 and T2 were measured, as well as the VEPs evoked by the stream of distractor stimuli. Already at the onset of the stream, these VEPs peaked earlier at the right than at the left hemisphere. N2pc was evoked earlier and P3 amplitudes were larger with left than with right T2. Previously reported side differences in T1-evoked N2pc were no longer obtained after correcting for constant hemispheric differences. The faster VEP latencies at the right hemisphere from the very beginning of the stimulus series may reflect an advantage in structuring fast sequences, which may cause the left visual-field advantage.
[show abstract][hide abstract] ABSTRACT: Previous research has indicated that information acquired before sleep gets consolidated during sleep. This process of consolidation might be reflected after sleep in changed extent and topography of cortical activation during retrieval of information. Here, we designed an experiment to measure those changes by means of slow event-related EEG potentials (SPs). Retrieval of newly learnt verbal or spatial associations was tested both immediately after learning and two days later. In the night directly following immediate recall, participants either slept or stayed awake. In line with previous studies, SPs measured during retrieval from memory had parietal or left-frontal foci depending on whether the retrieved associations were spatial or verbal. However, contrary to our expectations, sleep-related consolidation did not further accentuate these content-specific topographic profiles. Rather, sleep modified SPs independently of the spatial or verbal type of learned association: SPs were reduced more after sleep than after waking specifically for those stimulus configurations that had been presented in the same combination at retrieval before sleep. The association-independent stimulus-specific effect might generally form a major component of sleep-related effects on memory.
[show abstract][hide abstract] ABSTRACT: In the dual-stream Rapid Serial Visual Presentation task, a stream of stimuli containing two target stimuli is rapidly presented left and right. In previous studies, the second target was better identified in the left than in the right hemifield. In all those studies, alphanumeric stimuli were used both as targets and distracters. We examined to what extent this left visual-field advantage is dependent on reading-direction. The task was performed by Germans (with Latin characters), Israelis (with Latin and Hebrew characters) and Taiwanese (with Latin and Chinese characters). If caused by overlearnt associative links between Latin characters and left-to-right reading, the prominent left visual-field bias should be reversed in Hebrew and disappear in Chinese. Furthermore, if caused by direction of reading in the participant's native language, the left visual-field advantage in Latin conditions should be larger in Germans than in Israelis and Taiwanese. A left visual-field advantage was always observed, though slightly smaller in Hebrew and in Chinese, and there was no difference in the Latin conditions between the three nations. Therefore, it seems that the left visual-field advantage in speeded target identification is not primarily caused by the left-to-right reading-direction, but may be a combined effect resulting from the asymmetric organization of general mechanisms of visual processing and from stimulus-induced preferences.
[show abstract][hide abstract] ABSTRACT: In the present task, series of visual stimuli are rapidly presented left and right, containing two target stimuli, T1 and T2. In previous studies, T2 was better identified in the left than in the right visual field. This advantage of the left visual field might reflect dominance exerted by the right over the left hemisphere. If so, then repetitive transcranial magnetic stimulation (rTMS) to the right parietal cortex might release the left hemisphere from right-hemispheric control, thereby improving T2 identification in the right visual field. Alternatively or additionally, the asymmetry in T2 identification might reflect capacity limitations of the left hemisphere, which might be aggravated by rTMS to the left parietal cortex. Therefore, rTMS pulses were applied during each trial, beginning simultaneously with T1 presentation. rTMS was directed either to P4 or to P3 (right or left parietal cortex) either as effective or as sham stimulation. In two experiments, either one of these two factors, hemisphere and effectiveness of rTMS, was varied within or between participants. Again, T2 was much better identified in the left than in the right visual field. This advantage of the left visual field was indeed modified by rTMS, being further increased by rTMS to the left hemisphere rather than being reduced by rTMS to the right. It may be concluded that superiority of the right hemisphere in this task implies that this hemisphere is less irritable by external interference than the left hemisphere.
Experimental Brain Research 06/2010; 203(2):355-65. · 2.22 Impact Factor
[show abstract][hide abstract] ABSTRACT: Solving a task with insight has been associated with occipital and right-hemisphere activations. The present study tested the hypothesis if sleep-related alterations in functional activation states modulate the probability of insight into a hidden abstract regularity of a task.
State-dependent functional activation was measured by beta and alpha electroencephalographic (EEG) activity and spatial synchronization. Task-dependent functional activation was assessed by slow cortical potentials (SPs). EEG parameters during the performance of the Number Reduction Task (NRT) were compared between before sleep and after sleep sessions. In two different groups, the relevant sleep occurred either in the first or in the second half of the night, dominated by slow wave sleep (SWS) or by rapid eye movement (REM) sleep.
Changes in EEG parameters only occurred in the early-night group, not in the late-night group and indicated occipital and right-hemisphere functional alterations. These changes were associated with off-line consolidation of implicit task representations and with the amount of SWS but they did not predict subsequent insight. The gain of insight was, however, independently associated with changes of spectral beta and alpha measures only in those subjects from the two sleep groups who would subsequently comprehend the hidden regularity of the task. Insight-related enhancement of right frontal asymmetry after sleep did not depend on sleep stages.
It is concluded that off-line restructuring of implicit information during sleep is accompanied by alterations of functional activation states after sleep. This mechanism is promoted by SWS but not by REM sleep and may contribute to attaining insight after sleep. Original neurophysiologic evidence is provided for alterations of the functional activation brain states after sleep. These alterations are associated with a decrease in controlled processing within the visual system and with an increase in the functional connectivity of the right hemisphere, and are supported by SWS in the first half of the night.
PLoS ONE 01/2010; 5(2):e9442. · 3.73 Impact Factor
[show abstract][hide abstract] ABSTRACT: The objective of the present study was to evaluate patterns of implicit processing in a task where the acquisition of explicit and implicit knowledge occurs simultaneously. The number reduction task (NRT) was used as having two levels of organization, overt and covert, where the covert level of processing is associated with implicit associative and implicit procedural learning. One aim was to compare these two types of implicit processes in the NRT when sleep was or was not introduced between initial formation of task representations and subsequent NRT processing. To assess the effects of different sleep stages, two sleep groups (early- and late-night groups) were used where initial training of the task was separated from subsequent retest by 3 h full of predominantly slow wave sleep (SWS) or rapid eye movement (REM) sleep. In two no-sleep groups, no interval was introduced between initial and subsequent NRT performance. A second aim was to evaluate the interaction between procedural and associative implicit learning in the NRT. Implicit associative learning was measured by the difference between the speed of responses that could or could not be predicted by the covert abstract regularity of the task. Implicit procedural on-line learning was measured by the practice-based increased speed of performance with time on task. Major results indicated that late-night sleep produced a substantial facilitation of implicit associations without modifying individual ability for explicit knowledge generation or for procedural on-line learning. This was evidenced by the higher rate of subjects who gained implicit knowledge of abstract task structure in the late-night group relative to the early-night and no-sleep groups. Independently of sleep, gain of implicit associative knowledge was accompanied by a relative slowing of responses to unpredictable items suggesting reciprocal interactions between associative and motor procedural processes within the implicit system. These observations provide evidence for the separability and interactions of different patterns of processing within implicit memory.
Journal of Psychophysiology - J PSYCHOPHYSIOL. 01/2010; 24(2):91-101.
[show abstract][hide abstract] ABSTRACT: Patients with Parkinson's disease (PD) are more sensitive than healthy controls to response-triggering by irrelevant flanking stimuli in speeded choice-response tasks. This increased responsiveness may either indicate a lack of executive control or reflect compensatory efforts to cope with the reduced internal motor drive. Of interest in this context is whether responsiveness is already enhanced in the presymptomatic stage of PD. To address these questions, we studied a group of non-manifesting carriers of heterozygous Parkin and PINK1 mutations while they performed a choice-response task with response-compatible or incompatible flankers. These mutation carriers may be considered a model for pre-clinical PD because the mutant allele leads to a latent nigrostriatal dysfunction and may increase the risk for PD. For comparison, we studied groups of medicated patients with idiopathic PD and of healthy persons age-matched to the mutation carriers and to the patients. Measurements of reaction time, error rate, and the lateralized readiness potential of the EEG provided converging evidence that the mutation carriers were less responsive to distracting flankers than their healthy control group. In contrast, PD patients were more distractible by flankers than their control group, which replicated previous results. Mutation carriers also showed a smaller N2 component of the event-related EEG potential in trials with incompatible flankers relative to their control group, which might indicate reduced inhibitory control. We hypothesize that faulty executive control is the primary deficit, reflected by the reduced N2 component in the mutation carriers. To compensate for this deficit, mutation carriers change their strategy of speed-accuracy trade-off, in order to dampen the excitability of their lateral motor system. Disease progression might prevent symptomatic PD patients from using this compensatory mechanism, leading to increased disinhibition of their lateral motor system.
[show abstract][hide abstract] ABSTRACT: An important aspect of human motor control is the ability to resolve conflicting response tendencies. Here we used single-pulse transcranial magnetic stimulation (TMS) to track the time course of excitability changes in the primary motor hand areas (M1(HAND)) while the motor system resolved response conflicts. Healthy volunteers had to respond fast with their right and left index fingers to right- and left-pointing arrows. These central target stimuli were preceded by flanking arrows, inducing premature response tendencies which competed with correct response activation. The time point of maximum premature activation was individually measured as peak latency of the lateralized readiness potential (LRP) in the EEG. In the subsequent TMS experiment, single pulses were applied to left or right M1(HAND) during the same flanker task. The amplitude of the motor evoked potentials in the contralateral first dorsal interosseus muscle was taken as an index of corticospinal excitability. Guided by the previous LRP measurement, magnetic stimuli were applied 0-90 ms after the individual LRP peak, to cover the epoch of conflict resolution. When flankers were incompatible with the target, excitability of the prematurely activated M1(HAND) gradually decreased during this 90 ms period. This decrease was paralleled by a mirror-symmetrical increase in excitability in the other M1(HAND). These results show that the inappropriate response tendency is cancelled in one M1(HAND) simultaneously with activation of the correct response in the other. This integrated implementation of response activation and cancellation at the level of the M1(HAND) presumably represents a downstream effect orchestrated by premotor brain regions.
European Journal of Neuroscience 08/2009; 30(2):318-26. · 3.75 Impact Factor
[show abstract][hide abstract] ABSTRACT: There is evidence that slow wave sleep (SWS) promotes the consolidation of memories that are subserved by mediotemporal- and hippocampo-cortical neural networks. In contrast to implicit memories, explicit memories are accompanied by conscious (attentive and controlled) processing. Awareness at pre-sleep encoding has been recognized as critical for the off-line memory consolidation. The present study elucidated the role of task-dependent cortical activation guided by attentional control at pre-sleep encoding for the consolidation of hippocampus-dependent memories during sleep.
A task with a hidden regularity was used (Number Reduction Task, NRT), in which the responses that can be implicitly predicted by the hidden regularity activate hippocampo-cortical networks more strongly than responses that cannot be predicted. Task performance was evaluated before and after early-night sleep, rich in SWS, and late-night sleep, rich in rapid eye movement (REM) sleep. In implicit conditions, slow cortical potentials (SPs) were analyzed to reflect the amount of controlled processing and the localization of activated neural task representations.
During implicit learning before sleep, the amount of controlled processing did not differ between unpredictable and predictable responses, nor between early- and late-night sleep groups. A topographic re-distribution of SPs indicating a spatial reorganization occurred only after early, not after late sleep, and only for predictable responses. These SP changes correlated with the amount of SWS and were covert because off-line RT decrease did not differentiate response types or sleep groups.
It is concluded that SWS promotes the neural reorganization of task representations that rely on the hippocampal system despite absence of conscious access to these representations.
Original neurophysiologic evidence is provided for the role of SWS in the consolidation of memories encoded with hippocampo-cortical interaction before sleep. It is demonstrated that this SWS-mediated mechanism does not depend critically on explicitness at learning nor on the amount of controlled executive processing during pre-sleep encoding.
PLoS ONE 02/2009; 4(5):e5675. · 3.73 Impact Factor
[show abstract][hide abstract] ABSTRACT: The present study assessed the effects of awareness at encoding on off-line learning during sleep. A new framework is suggested according to which two aspects of awareness are distinguished: awareness of task information, and awareness of task processing. The number reduction task (NRT) was employed because it has two levels of organization, an overt one based on explicit knowledge of task instructions, and a covert one based on hidden abstract regularities of task structure (implicit knowledge). Each level can be processed consciously (explicitly) or non-consciously (implicitly). Different performance parameters were defined to evaluate changes between two sessions for each of the four conditions of awareness arising from whether explicit or implicit task information was processed explicitly or implicitly. In two groups of subjects, the interval between the pre-sleep and post-sleep sessions was filled either with early-night sleep, rich in slow wave sleep (SWS), or late-night sleep, rich in rapid eye movement (REM) sleep. Results show that implicit processing of explicit information was improved in the post-sleep relative to the pre-sleep session only in the early-night group. Independently of sleep stage, changes between sessions occurred for explicit processing of implicit information only in those subjects who gained insight into the task regularity after sleep. It is concluded that SWS but not REM sleep specifically supports gains in computational skills for the processing of information that was accessible by consciousness before sleep.
Frontiers in Human Neuroscience 01/2009; 3:40. · 2.91 Impact Factor
[show abstract][hide abstract] ABSTRACT: When simultaneous series of stimuli are rapidly presented left and right, containing two target stimuli T1 and T2, T2 is much better identified when presented in the left than in the right hemifield. Here, this effect was replicated, even when shifts of gaze were controlled, and was only partially compensated when T1 side provided the cue where to expect T2. Electrophysiological measurement revealed earlier latencies of T1- and T2-evoked N2(pc) peaks at the right than at the left visual cortex, and larger right-hemisphere T2-evoked N2(pc) amplitudes when T2 closely followed T1. These findings suggest that the right hemisphere was better able to single out the targets in time. Further, sustained contralateral slow shifts remained active after T1 for longer time at the right than at the left visual cortex, and developed more consistently at the right visual cortex when expecting T2 on the contralateral side. These findings might reflect better capacity of right-hemisphere visual working memory. These findings about the neurophysiological underpinnings of the large right-hemisphere advantage in this complex visual task might help elucidating the mechanisms responsible for the severe disturbance of hemineglect following damage to the right hemisphere.
Journal of Cognitive Neuroscience 07/2008; 21(3):474-88. · 4.49 Impact Factor
[show abstract][hide abstract] ABSTRACT: Sleep has proven to support the memory consolidation in many tasks including learning of perceptual skills. Explicit, conscious types of memory have been demonstrated to benefit particularly from slow-wave sleep (SWS), implicit, non-conscious types particularly from rapid eye movement (REM) sleep. By comparing the effects of early-night sleep, rich in SWS, and late-night sleep, rich in REM sleep, we aimed to separate the contribution of these two sleep stages in a metacontrast masking paradigm in which explicit and implicit aspects in perceptual learning could be assessed separately by stimulus identification and priming, respectively. We assumed that early sleep intervening between two sessions of task performance would specifically support stimulus identification, while late sleep would specifically support priming. Apart from overt behavior, event-related EEG potentials (ERPs) were measured to record the cortical mechanisms associated with behavioral changes across sleep. In contrast to our hypothesis, late-night sleep appeared to be more important for changes of behavior, both for stimulus identification, which tended to improve across late-night sleep, and for priming, with the increase of errors induced by masked stimuli correlating with the duration of REM sleep. ERP components proved sensitive to presence of target shapes in the masked stimuli and to their priming effects. Of these components, the N2 component, indicating processing of conflict, became larger across early-night sleep and was related to the duration of S4 sleep, the deepest substage of SWS containing particularly high portions of EEG slow waves. These findings suggest that sleep promotes perceptual learning primarily by its REM sleep portion, but indirectly also by way of improved action monitoring supported by deep slow-wave sleep.
Brain and Cognition 07/2008; 68(2):180-92. · 2.82 Impact Factor