EEG-power and -coherence changes in a unimodal and a crossmodal working memory task with visual and kinesthetic stimuli.

We investigated EEG-power and EEG-coherence changes in a unimodal and a crossmodal matching-to-sample working memory task with either visual or kinesthetic stimuli. Angle-shaped trajectories were used as stimuli presented either as a moving dot on a screen or as a passive movement of a haptic device. Effects were evaluated during the different phases of encoding, maintenance, and recognition. Alpha power was modulated during encoding by the stimulus modality, and in crossmodal conditions during encoding and maintenance by the expected modality of the upcoming test stimulus. These power modulations were observed over modality-specific cortex regions. Systematic changes of coherence for crossmodal compared to unimodal tasks were not observed during encoding and maintenance but only during recognition. There, coherence in the theta-band increased between electrode sites over left central and occipital cortex areas in the crossmodal compared to the unimodal conditions. The results underline the importance of modality-specific representations and processes in unimodal and crossmodal working memory tasks. Crossmodal recognition of visually and kinesthetically presented object features seems to be related to a direct interaction of somatosensory/motor and visual cortex regions by means of long-range synchronization in the theta-band and such interactions seem to take place at the beginning of the recognition phase, i.e. when crossmodal transfer is actually necessary.

Control processes during selective long-term memory retrieval.

In our daily life, we often need to selectively remember information related to the same retrieval cue in a consecutive manner (e.g., ingredients from a recipe). To investigate such selection processes during cued long-term memory (LTM) retrieval, we used a paradigm in which the retrieval demands were systematically varied from trial to trial and analyzed, by means of behavior and slow cortical EEG potentials (SCPs), the retrieval processes in the current trial depending on those of the previous trial. We varied whether the retrieval cue, the type of to-be-retrieved association (feature), or retrieval load was repeated or changed from trial to trial. The behavioral data revealed a benefit of feature repetition, probably due to trial-by-trial feature priming. SCPs further showed an effect of cue change with a mid-frontal maximum, suggesting increased control demands when the cue was repeated, as well as a parietal effect of retrieval-load change, indicating increased activation of posterior neural resources when focusing on a single association after all learned associations had been activated previously, compared to staying with single associations across trials. These effects suggest the existence of two distinct types of dynamic (trial-by-trial) control processes during LTM retrieval: (1) medial frontal processes that monitor or regulate interference within a set of activated associations, and (2) posterior processes regulating attention to LTM representations. The present study demonstrates that processes mediating selective LTM retrieval can be successfully studied by manipulating the history of processing demands in trial sequences.

Online action-to-perception transfer: only percept-dependent action affects perception.

Perception self-evidently affects action, but under which conditions does action in turn influence perception? To answer this question we ask observers to view an ambiguous stimulus that is alternatingly perceived as rotating clockwise or counterclockwise. When observers report the perceived direction by rotating a manipulandum, opposing directions between report and percept ('incongruent') destabilize the percept, whereas equal directions ('congruent') stabilize it. In contrast, when observers report their percept by key presses while performing a predefined movement, we find no effect of congruency. Consequently, our findings suggest that only percept-dependent action directly influences perceptual experience.

Nonvisual motor learning improves visual motion perception: evidence from violating the two-thirds power law.

Few studies have reported direct effects of motor learning on visual perception, especially when using novel movements for the motor system. Atypical motor behaviors that violate movement constraints provide an excellent opportunity to study action-to-perception transfer. In our study, we passively trained blindfolded participants on movements violating the 2/3 power law. Before and after motor training, participants performed a visual discrimination task in which they decided whether two consecutive movements were same or different. For motor training, we randomly assigned the participants to two motor training groups or a control group. The motor training group experienced either a weak or a strong elliptic velocity profile on a circular trajectory that matched one of the visual test stimuli. The control group was presented with linear trajectories unrelated to the viewed movements. After each training session, participants actively reproduced the movement to assess motor learning. The group trained on the strong elliptic velocity profile reproduced movements with increasing elliptic velocity profiles while circular geometry remained constant. Furthermore, both training groups improved in visual discrimination ability for the learned movement as well as for highly similar movements. Participants in the control group, however, did not show any improvements in the visual discrimination task nor did participants who did not acquire the trained movement. The present results provide evidence for a transfer from action to perception which generalizes to highly related movements and depends on the success of motor learning. Moreover, under specific conditions, it seems to be possible to acquire movements deviating from the 2/3 power law.

fMRI data predict individual differences of behavioral effects of nicotine: a partial least square analysis.

Reorienting of visuospatial attention can be investigated by comparing reaction times to validly and invalidly cued targets ("validity effect"). The cholinergic agonist nicotine reduces the validity effect and neural activity in the posterior parietal cortex. Behavioral effects of nicotine in nonsmokers are weak and it has been suggested that differences in baseline behavior before nicotine exposure may influence the effect of nicotine. This study investigates whether individual differences in reorienting-related neural activity under placebo may be used to predict individual nicotine effects. Individual nicotine effects are defined as the behavioral effects under nicotine that cannot be predicted by the behavioral data under placebo. Fifteen nonsmoking subjects were given either placebo or nicotine gum (2 mg) prior to performing a cued target detection task inside a magnetic resonance imaging scanner. The results of a partial least square analysis suggest that neural data under placebo can be used to predict individual behavioral effects of nicotine. Neural activity in the left posterior cingulate cortex, the right superior parietal cortex, the right dorsal medial prefrontal cortex, and the left ventral medial prefrontal cortex significantly contributes to that prediction. We conclude that nicotine effects on reorienting attention depend on individual differences in reorienting-related neural activity under placebo and suggest that functional magnetic resonance imaging data can contribute to the prediction of individual drug effects.

The modulatory effects of nicotine on parietal cortex activity in a cued target detection task depend on cue reliability.

This functional magnetic resonance imaging study investigates the effects of nicotine in a cued target detection task when changing cue reliability. Fifteen non-smoking volunteers were studied under placebo and nicotine (Nicorette polacrilex gum 1 and 2 mg). Validly and invalidly cued trials were arranged in blocks with high, middle and low cue reliability. Two effects of nicotine were investigated: its influence on i) parietal cortex activity underlying the processing of invalid vs. valid trials (i.e. validity effect) and ii) neural activity in the context of low, middle and high informative value of the cue (i.e. cue reliability effect). Nicotine did not affect behavioral performance. However, nicotine reduced the difference in the blood oxygenation level dependent (BOLD) signal between invalid and valid trials in the right intraparietal sulcus. The reduction of parietal activity in invalid trials was smaller in the low cue reliability condition. The same posterior parietal region exhibited a nicotinic modulation of BOLD activity in valid trials which was dependent on cue reliability: Nicotine specifically enhanced the neural activity during valid trials in the context of low cue reliability, i.e. when subjects are already in a state of low certainty. We speculate that the right intraparietal sulcus might be part of two networks working in parallel: one responsible for reorienting attention and the other for the cholinergic modulation of cue reliability. By reducing the use of the cue, nicotine modulates parietal activity related to reorienting attention in conditions with higher cue certainty. On the other hand, nicotine increases parietal activity in states of low certainty. This enhanced activation might influence brain regions, such as the posterior cingulate, directly involved in the processing of cue reliability.

Event-related brain potentials evoked by verbs and nouns in a primed lexical decision task.

We investigated whether verbs and nouns evoke comparable behavioral and N400 effects in a primed lexical decision task. Twenty-nine students were tested, 13 in a pilot study in which only response times and error rates were collected and 16 in a study in which ERPs were recorded from 124 scalp electrodes. Stimuli were noun-noun and verb-verb pairs with the targets bearing either a strong, a moderate, or no semantic association to the prime or being a pseudoword. Behavioral data revealed comparable priming effects for both word categories. These proved to be independent from the SOA (250 and 800 ms) and they followed the well-known pattern of decreasing response times and error rates with increasing relatedness between target and prime. ERPs revealed pronounced N400 effects for both word categories with a larger amplitude for noun than for verb pairs. A systematic analysis of topographic differences between noun- and verb-evoked ERPs and N400 effects, respectively, gave no convincing support to the hypothesis that the two word categories activate distinct neuronal networks.

Auditory memory in congenitally blind adults: a behavioral-electrophysiological investigation.

Blind people must rely more than sighted people on auditory input in order to acquire information about the world. The present study was designed to test the hypothesis that blind people have better memory than sighted individuals for auditory verbal material and specifically to determine whether memory encoding and/or retrieval are improved in blind adults. An incidental memory paradigm was employed in which 11 congenitally blind people and 11 matched sighted controls first listened to 80 sentences which ended either with a semantically appropriate or inappropriate word. Immediately following, the recognition phase occurred, in which all sentence terminal words were presented again randomly intermixed with the same number of new words. Participants indicated whether or not they had heard the word in the initial study phase. Event-related brain potentials (ERPs) were recorded from 28 electrode positions during both the encoding and the retrieval phase. Blind participants' memory performance was superior to that of sighted controls. In addition, during the recognition phase, previously presented words elicited ERPs with larger positive amplitudes than new words, particularly over the right hemisphere. During the study phase, words that would subsequently be recognized elicited a more pronounced late positive potential than words that were not subsequently recognized. These effects were reliable in the congenitally blind participants but could only be obtained in the subgroup of sighted participants who had the highest memory performance. These results imply that blind people encode auditory verbal material more efficiently than matched sighted controls and that this in turn allows them to recognize these items with a higher probability.

Individual cortical current density reconstructions of the semantic N400 effect: using a generalized minimum norm model with different constraints (L1 and L2 norm).

Event-related brain potentials were recorded to study whether verbs and nouns activate topographically distinct cortical generators. Fifteen subjects performed a primed lexical decision task with verb/verb and noun/noun pairs. The relatedness between prime and target items was varied in three steps (unrelated, moderately, and strongly related) and the EEG was recorded from 124 scalp electrodes. The topography of cortical sources of the N400 effect was evaluated by standardized differences scores and by cortical current source estimates which were constrained by the individual MRI-determined cortex anatomy. A behavioral priming effect and a substantial N400 effect was found for both word categories. However, the topography of the grand average N400 effect of verbs and nouns did not differ, neither for raw nor for standardized amplitudes. Cortical current source estimates of the N400 effect revealed a very broad and scattered distribution of active locations with pronounced interindividual differences. Cortical current source estimates obtained with the L1-norm and L2-norm model, respectively, differed in the distribution of sources over the cortex but converged on the same "hot spots." The data give no indication that the N400 effect is generated by word category-specific networks which have a different topography. The marked individual differences are discussed with respect to the involved processes and the current source estimation procedures.

Inhibition of return and oculomotor control in the blind.

Our attentional systems orient reflexively to novel environmental stimuli. Such attentive orienting is typically followed by a prolonged period of inhibition, known as inhibition of return (IOR), thought to be linked to the eye movement system. It is widely believed that IOR may provide a tagging mechanism that prevents perseveration, and thus facilitates attentional search. Using a tactile variant of the peripheral spatial cuing paradigm, we show IOR in congenitally blind adults and in an individual who had no eyes. These results demonstrate for the first time that spatial IOR can occur in the absence of oculomotor control.

Event-related potentials during auditory language processing in congenitally blind and sighted people.

While behavioral studies have documented delayed language acquisition in blind children, other studies have revealed better speech discrimination abilities for blind than sighted adults. Several brain imaging studies have provided evidence for cortical reorganization due to visual deprivation but the cerebral organization of language in blind humans is not known yet. We hypothesized that the increasing specialization of language systems normally observed during development may not take place to the same degree in blind individuals since posterior visual areas do not receive their adequate input. On the other hand, we hypothesized that blind people, due to their greater reliance upon the auditory language signal, may process speech faster than sighted people. To test these assumptions, event-related potentials were recorded while 11 congenitally blind and 11 sighted adults matched in age, gender, handedness and education were engaged in a language task. Participants listened to sentences in order to decide after each sentence if it was meaningful or not. Incongruous sentence-final words elicited an N400 effect in both groups. The N400 effect had a left-lateralized fronto-central scalp distribution in the sighted but a symmetric and broad topography in the blind. Furthermore, the N400 effect started earlier in the blind than in the sighted. Closed class compared to open class sentence middle words elicited a more pronounced late negativity in the blind than in the sighted. These results suggest that blind people process auditory language stimuli faster than sighted people and that some language functions may be reorganized in the blind.

Implicit and explicit learning of event sequences: evidence for distinct coding of perceptual and motor representations.

Event-related brain potentials (ERPs) of 21 subjects were recorded in a choice reaction time task with a repeating eight-element long stimulus sequence. The regular event sequence was sometimes interrupted by 'perceptual' or by 'motor deviants' which both replaced an expected stimulus but either preserved or violated the sequence of motor responses. Response times confirmed that all subjects had acquired some knowledge of the sequential dependencies. By means of a post-experimental free recall and recognition test, subjects were classified as having either explicit or implicit knowledge of the event sequence. The ERPs showed different effects for different types of stimuli and the two groups. In the group of explicit learners, a larger N200 component was evoked by both types of deviants and a larger P300 by motor deviants only. In the group of implicit learners these 'perceptual components' remained unaffected. In contrast, in both groups of subjects the lateralized readiness potential (LRP) which accompanied motor deviants revealed a partial activation of the to be expected but incorrect response, i.e. motor learning. These results suggest that explicit learners acquire knowledge about both, stimulus and response dependencies while implicit learners acquire knowledge about response dependencies only.

Topography of brain electrical activity dissociates the sequential order transformation of verbal versus spatial information in humans.

Slow event related potentials (ERPs) of the electroencephalogram were recorded while subjects performed a short-term memory task. They had to mentally transform the order of sequentially presented words (verbal condition) or positions within a grid (spatial condition). The difficulty of the transformation process was varied. The amplitudes of the slow ERPs elicited during the transformation process varied with task difficulty. The topography of this amplitude modulation turned out to be information specific: The maximal effect during transformation of the sequential order of verbal information was found over the left-frontal cortex and during the transformation of spatial information over the parietal cortex. These results suggest that the manipulation of short-term memory contents seems to take place in information specific cortical areas which are thought to be also involved during on-line processing and long-term storage of these informations.

Event-related responses to pronoun and proper name anaphors in parallel and nonparallel discourse structures.

The present study investigated event-related potential (ERP) effects of pronoun and proper name anaphors in both parallel and nonparallel discourse structures. Thirty-seven students processed 400 semantically different text passages. Each trial consisted of two sentences and a comprehension question. The first sentence introduced a protagonist who was referred to by an anaphoric word in the second sentence. The anaphoric word was either a pronoun or a repetition of the proper name of the protagonist and had either the same or a different syntactic role as its antecedent (subject or object). The sentences were presented word by word as rapid serial visual display. Event-related potentials were recorded from 61 scalp electrodes. In agreement with the parallel function strategy, nonparallel discourse structures required longer decision times and exhibited higher error rates than parallel structures. The ERPs revealed two effects: First, pronoun anaphors evoked a more pronounced negativity than proper name anaphors between 270 and 420 ms latency over the frontal cortex electrodes. Another relative negativity occurred between 510 and 600 ms over the parietal cortex electrodes. Second, anaphors in nonparallel positions were accompanied by a more pronounced negativity over the parietal cortex. These data support the idea that an anaphor in nonparallel position triggers extra processing steps, probably search processes in working memory which integrate currently encountered information with previously activated representations.

Improved auditory spatial tuning in blind humans.

Despite reports of improved auditory discrimination capabilities in blind humans and visually deprived animals, there is no general agreement as to the nature or pervasiveness of such compensatory sensory enhancements. Neuroimaging studies have pointed out differences in cerebral organization between blind and sighted humans, but the relationship between these altered cortical activation patterns and auditory sensory acuity remains unclear. Here we compare behavioural and electrophysiological indices of spatial tuning within central and peripheral auditory space in congenitally blind and normally sighted but blindfolded adults to test the hypothesis (raised by earlier studies of the effects of auditory deprivation on visual processing) that the effects of visual deprivation might be more pronounced for processing peripheral sounds. We find that blind participants displayed localization abilities that were superior to those of sighted controls, but only when attending to sounds in peripheral auditory space. Electrophysiological recordings obtained at the same time revealed sharper tuning of early spatial attention mechanisms in the blind subjects. Differences in the scalp distribution of brain electrical activity between the two groups suggest a compensatory reorganization of brain areas in the blind that may contribute to the improved spatial resolution for peripheral sound sources.

Processing of incongruous mental calculation problems: evidence for an arithmetic N400 effect.

We investigated if incongruent solutions of simple multiplication problems would elicit similar event-related brain potentials as inappropriate words in sentences. In Experiment I, 12 subjects verified the appropriateness of solutions of multiplication problems or of final words in short sentences. Both incongruent solutions and incongruent words evoked a phasic negative shift between 300 and 500 ms having a similar topography. In Experiment II, we tested with another sample of 13 subjects if the amplitude of this arithmetic N400 effect was affected differently by different stimulus onset asynchronies (SOA = 200 and 500 ms) and by errors that were either table-related or table-unrelated to the preceding operands. Again, incorrect solutions elicited an arithmetic N400 effect whose amplitude depended on both the relatedness of the solution and the SOA. The ascending part of the N400 effect was always larger for unrelated than for related errors independently of the SOA, whereas the maximum of the N400 effect was larger for unrelated errors in case of a long SOA only. This pattern of effects was similar to that observed with semantic material varying lexical associations. These results suggest that arithmetic incongruencies are handled by the system in a manner functionally similar to that of semantic incongruencies.

On separating processes of event categorization, task preparation, and mental rotation proper in a handedness recognition task.

We investigated the nature of event-related potential (ERP) effects in a handedness recognition task requiring mental rotation. Thirty subjects were tested with rotated and sometimes reflected alphanumeric characters while ERPs were recorded from 18 electrodes. On each trial, a cue provided valid information about the angular displacement of the following probe. This design allowed a distinction between three processing episodes: evaluation of the difficulty of the forthcoming task, preparation for the task, and the mental rotation task itself. The three episodes were accompanied by distinct ERP effects having distinct polarities, a different rank order of amplitudes for different probe orientations, and a different topography. These data confirm previous findings showing that mental rotation is accompanied by a parietal negativity. However, they also suggest that the rotation-related negativity found after tilted stimuli in standard mental rotation tasks is most likely overlapping with another, simultaneously triggered ERP effect functionally related to an evaluation of task difficulty.

Effects of interstimulus interval on auditory event-related potentials in congenitally blind and normally sighted humans.

To test the hypothesis of auditory compensation after early visual deprivation, congenitally blind and sighted adults performed an auditory discrimination task. They had to detect a rare target tone among frequent standard tones. Stimuli were presented with different interstimulus intervals (ISIs) (200, 1000, 2000 ms) and the auditory-event related potentials to all tones and reaction times to targets were recorded. Increasing ISIs resulted in an increasing amplitude of the vertex response (N1-P2) in both groups, but this amplitude recovery was more pronounced in the blind. Furthermore, targets elicited larger and more posteriorly distributed N2 responses in the blind than in the sighted. Since target detection times were shorter in the blind as well, these findings imply compensatory adaptations within the auditory modality in humans blind from birth.

What is improved if a mental rotation task is repeated--the efficiency of memory access, or the speed of a transformation routine?

The present study contrasts 3 theories which provide explanations for performance improvement in mental rotation tasks. Wallace and Hofelich conjectured that the process as such may be executed more rapidly after training, while Bethell-Fox and Shepard attributed practice effects to the fact that images may be transformed first elementwise, but later as a Gestalt. In contrast, Tarr and Pinker assumed that a transformation of an image will no longer be computed after training but simply be retrieved from memory. Thirty-seven subjects participated in 3 test sessions in which they had to decide on the parity of 3-D block figures presented from different perspectives. Experimental group subjects underwent 4 additional practice sessions in which a subset of the figures and a subset of perspective views were used. Tests adapted to the predictions of the 3 theories revealed specific learning effects but no transfer, neither to old objects presented in new perspectives nor to new objects. This supports an instance-based explanation of practice effects which states that objects are represented in multiple perspective views.

Different cortical activation patterns in blind and sighted humans during encoding and transformation of haptic images.

In this study, we investigated whether the occipital cortex of blind humans is activated during haptic perception and/or transformation of a haptic image. Slow event-related brain potentials were monitored from 18 electrodes in 12 sighted and 15 congenitally blind participants while they were engaged in a haptic mental rotation task. In both groups, slow negative shifts appeared over (a) the frontal cortex at the beginning of each processing episode, (b) the left-central to parietal cortex during encoding and maintaining of a haptic image, and (c) the central to parietal cortex during image transformation. A pronounced slow negative potential over the occipital cortex emerged only in the blind individuals and was time-locked to the processing epochs. Its amplitude increased with the amount of processing load. The slow wave effects observed in the blind individuals could indicate that occipital areas participate in specific, nonvisual functions or they could reflect a coactivation of these areas whenever the activation level of task-specific processing modules located elsewhere in the cortex is raised by nonspecific thalamocortical input.