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.

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.

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 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.

Topographic differences of slow event-related brain potentials in blind and sighted adult human subjects during haptic mental rotation.

Twelve blindfolded sighted, nine congenitally blind, and seven adventitiously blind subjects were tested in a haptic mental rotation task while slow event-related brain potentials in the EEG were recorded from 17 scalp locations. The overall topography of the slow wave pattern which prevailed during the task differed for sighted and for blind, but not for congenitally and adventitiously blind subjects. While the tactile stimuli were encoded, the blind showed a pronounced occipital and the sighted a pronounced frontal activation. The task-specific amplitude increment of a negative slow wave which can be understood as a manifestation of the process of mental rotation proper, showed a different topography for sighted and for blind subjects too. It had its maximum over central to parietal cortical areas in both groups, but it extended more towards occipital regions in the blind. In both groups, the effects were very similar to those observed in former studies with visual versions of the mental rotation task, i.e. the slow wave amplitude over central to parietal areas increased monotonously with an increasing angular disparity of the two stimuli to be compared. These results are discussed with respect to the question of whether visual deprivation in the blind can cause a reorganization of cortical representational maps.

Event-related potentials during auditory and somatosensory discrimination in sighted and blind human subjects.

The objective of the present study was to test if and to what extent phasic and tonic event-related potentials of the human EEG may reflect phenomena of cortical plasticity. In particular, it was tested if the occipital cortex of blind subjects participates in the processing of non-visual stimuli. To this end, 12 blind and 12 blindfolded sighted subjects were tested in an auditory and a somatosensory discrimination task with 2 levels of discrimination difficulty. Slow and fast event-related potentials were recorded from 18 scalp electrodes. In addition to the negative slow waves found in sighted subjects over frontal and central sites during auditory and somatosensory discrimination, a pronounced negative wave was revealed in the blind also over occipital brain areas. These negative shifts were time-locked to the train of stimuli which had to be monitored with sustained attention, i.e. they rised and resolved with the beginning and the end of a 20-s discrimination time epoch. The P300 complex, on the other hand, which is a slow positive deflection over the posterior part of the scalp and which follows rare and task-relevant events 200-800 ms after stimulus onset was significantly smaller at occipital electrodes in the blind than in the sighted subjects. Combined with neurophysiological and neuronanatomical evidence originating from studies with visually deprived animals, these data suggest that the occipital cortex of blind human subjects is coactivated whenever the system is engaged in a task which requires sustained attention and is less effectively inhibited at the end of a perceptual time epoch. In total, the data cast doubt on the hypothesis that the occipital cortex of blind subjects participates in modality-specific non-visual information processing.

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.

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.

Stimulus-induced gamma oscillations: harmonics of alpha activity?

Event-related changes of spectral power of the EEG were studied for each integer frequency between 5 and 100 Hz in three subjects during memory storage and retrieval. Spectra were calculated for successive, overlapping time epochs in seven channels. In one subject a stimulus-locked increase of power was observed at 12 Hz, while in the other two alpha power decreased at the individual peak frequencies of 9 and 11 Hz, respectively. In all subjects corresponding changes of power appeared at frequencies which were integer multiples of the individual dominant alpha frequencies. An analysis of the cross covariance of the alpha, beta and gamma activity revealed high coefficients for harmonic frequencies only while all other covariances were negligibly small. It is argued that event-related gamma activity may be an epiphenomenon of event-related changes within the alpha band.

Topography of brain electrical activity dissociates the retrieval of spatial versus verbal information from episodic long-term memory in humans.

Topography and amplitude of slow event-related potentials (ERPs) of the electroencephalogram (EEG) were studied during acquisition and recall of spatial and verbal associations. Subjects learned associations between line drawings and two types of mediators. The latter were either positions in a grid or concrete nouns. In a cued recall test subjects had to decide whether two drawings were linked to each other or not via an associated position or noun. The topography of slow ERPs 1-4 s after stimulus presentation obtained from 18 scalp electrodes dissociated the memory processes: The maximum potential was found over the parietal cortex with spatial and over the left frontal cortex with verbal information. The same topographic pattern emerged during both anticipation learning and cued recall. Moreover, the amplitude at the topographic maximum increased when more associations had to be retrieved. These results are compatible with the idea that memory representations are reactivated in localized cortical cell assemblies specialized for particular codes.

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.

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.

Imagery-perception interaction depends on the shape of the image: a reply to Farah (1989).

Farah (1989) reported that point threshold stimuli are detected better if they appear at spatial locations in the visual field that are covered by an image. By replicating her experiment with 3 instead of the original 2 images, we found that the effect depends on the shape of the mentally projected image. A second experiment with 9 different shapes revealed that the effect is modulated by the compactness and the size of the image--it is enhanced with increasing compactness and attenuated with increasing size. These findings do not unequivocally support the idea that imagery and perception interact because both processes share the same representational medium. Rather, they suggest that imagery can cause a figure-ground segregation in the visual field and that the shape of the figure may determine the amount of attention that is allocated to different sections in the visual field.

Cortical potential correlates of selective attention in multidimensional scaling.

Cortical evoked potentials were recorded from the vertex of subjects performing a similarity rating task. Proximity data were analysed according to INDSCAL (Carroll and Chang, 1970). Brain activity was averaged separately for different stimulus attributes and different stages of the judgement process. The amplitudes of the two positive deflections of the averaged evoked potential, peaking around 160 and 330 msec after stimulus onset respectively, were systematically influenced by two factors: The weight, which the evoking attribute received in the overall dissimilarity judgement, and the position of the evoking attribute in the stimulus train. The results support the hypothesis that differences in the perception of similarity are due to differences in attentional set. Reference is made to the 'additive-difference model' of scaling behaviour (Tversky and Krantz, 1970) and Broadbent's (1970) distinction between stimulus set and response set attention.

Principal components and varimax-rotated components in event-related potential research: some remarks on their interpretation.

Some properties of principal components analysis (PCA) and simple structure rotation, which are relevant to the study of event-related potentials (ERPs), were examined both in theory and with simulated data. The analysis casts some doubt on whether it is useful and desirable to rotate the loading pattern of a PCA of ERPs to simple structure. The evidence presented is more in favour of the unrotated solution. In particular we were able to demonstrate with simulated data that a solution rotated to simple structure may lead to false conclusions about the functional independence of ERP peaks. Beyond this, the more general question is asked of whether principal components should be accepted as basic waveforms, i.e. as physiologically meaningful entities which, for example, represent different ERP generators.

The double-priming paradigm: a tool for analyzing the functional significance of endogenous event-related brain potentials.

A new paradigm to study the functional significance of 'P300' is presented. Its advantages are: The precise definition and manipulation of cognitive operations which are triggered by the very same events as used for ERP extraction; and a systematic control over the probability of events known to affect endogenous event-related potential components (probabilities of single events, event categories, and event sequences). By employing the paradigm in two experiments with visual stimuli, three subcomponents of 'P300' were identified: P3a; P3b; and positive Slow Wave (pSW). Experimental manipulations revealed that P3b is related to the information processing resources required to alter a perceptual set and pSW to the resources required when abstract information permanently stored in memory must be retrieved. The data further revealed that the same-different disparity in response latency for matching letters has at least two ERP correlates: A difference in P3b latency; and a difference in the amplitude of a negative recess between P3a and P3b.

Slow negative brain potentials as reflections of specific modular resources of cognition.

The paper summarizes a series of experiments in which the topography and the amplitude of slow event-related brain potentials (slow ERPs) was studied in cognitive tasks which imposed different amounts of load on functionally distinct processing modules. The results suggest that the topography of slow ERPs reflects the relative activation/inactivation of distinct cortical cell assemblies while the absolute amplitude of the negative maximum seems to reflect how much a particular cell assembly is activated at a particular time. Translated into the terminology of cognitive resource theories one could say that tasks which evoke distinct slow wave patterns draw on independent resources. Likewise, the amplitude of a slow wave pattern could be related to the construct of resource allocation, i.e. the larger the amplitude of a slow wave pattern the more resources of a particular type are allocated to a task. These findings are discussed with respect to possible generator mechanism of slow waves and in relation to possible causes of capacity limitations of the human information processing system.

Dynamics of activation in long-term memory: the retrieval of verbal, pictorial, spatial, and color information.

Paivio's (1986) dual code theory was tested in 5 experiments with a new paradigm for the FAN effect that enforced genuine memory recall. Subjects had to learn associations between concepts and mediators. The FAN of the concepts in relation to the mediators was varied systematically. Response times (RT) were measured while subjects had to decide whether 2 concepts were linked to each other or not by a common mediator. In Experiment 1 the concepts and mediators were words, whereas in the other experiments the concepts were line drawings. Colors served as mediators in Experiment 2 and spatial locations served as mediators in Experiments 3, 4, and 5. All of the experiments were equivalent with respect to the FAN, the learning procedure, and the retrieval test. In all of the experiments, RT proved to be a linear function of the FAN. These results suggested that the same dynamics hold for all types of information stored in long-term memory.

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.

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.