From orthography to meaning: an electrophysiological investigation of the role of phonology in accessing meaning of Chinese single-character words.

Homophone interference effects in Stroop experiments are often taken as evidence for the hypothesis that semantic access in written Chinese language is mediated by activation of phonological processing. We here aim to test this hypothesis with Chinese single-character words by means of event related potential (ERP) recordings. Using color words, homophones of color words and color-word associates as materials in a Stroop task, we found behavioral Stroop interference effects for all stimulus types and an N450 for incongruent color words and color-word associates. Critically, there was no difference in the ERP waveforms elicited by congruent and incongruent homophones in the N450 time window. However, in a later time window (600-800 ms) the incongruent homophones elicited an apparent positivity over left posterior regions. A similar effect was also observed for incongruent color words. These findings thus indicate that phonology does not play an important role in semantic activation of Chinese single-character words, and that the behavioral Stroop effects for homophones possibly arises at later stage of lexical processing.

Processing of famous faces and medial temporal lobe event-related potentials: a depth electrode study.

The present study aims at analyzing the modulation of two types of event-related potentials originating from the human medial temporal lobe, the rhinal AMTL-N400 and the hippocampal P600 by the processing of famous faces. Therefore, we used a face recognition paradigm in which subjects had to discriminate the faces of famous persons from the faces of non-famous persons. Eleven patients with unilateral medial temporal lobe epilepsy undergoing intrahippocampal depth electrode recording for presurgical evaluation participated in this study. Event-related potentials (ERP) were recorded while a sequence of famous and non-famous faces was presented to the patients. The presentation of each face was repeated. The faces evoked N400-like potentials (anterior medial temporal lobe N400, AMTL-N400) in the rhinal cortex and P600-like potentials in the hippocampus. ERPs elicited by famous faces were contrasted with ERPs elicited by non-famous faces. The first presentation of famous faces elicited an enhanced AMTL-N400 and an enhanced hippocampal P600 in comparison to the second presentations of the famous faces or the (first and second presentation of the) non-famous faces. This findings are discussed in terms of associative semantic memory processes and the retrieval of person-specific information from long-term memory stores triggered by the processing of famous faces.

Recognition of famous faces in the medial temporal lobe: an invasive ERP study.

OBJECTIVE: To investigate the involvement of the rhinal cortex and the hippocampus in the processing of famous faces in contrast to nonfamous faces using intracranial event-related potentials (ERPs), and to analyze repetition effects for famous and nonfamous faces. METHODS: ERPs were elicited by pictures of famous and nonfamous faces and recorded from rhinal and hippocampal sites of intracranial electrodes in 10 presurgical patients with unilateral medial temporal lobe epilepsy. Famous and nonfamous faces were presented twice and mixed with distorted faces serving as targets. There was no instruction for an overt discrimination between famous and nonfamous faces. In contrast to nonfamous faces, famous faces stimulate processes related with access and retrieval of semantic memory. RESULTS: All faces evoked anterior medial temporal lobe N400-like (AMTL-N400) potentials in the rhinal cortex and P600-like potentials in the hippocampus. The AMTL-N400 and the hippocampal P600 amplitudes were larger for famous faces than for nonfamous faces. Mean amplitudes of the first and second presentation of famous faces suggest a repetition effect for the rhinal sites; however, they are significant only in the later signal components. No repetition effect was found for nonfamous faces and for potentials from the hippocampus. CONCLUSION: The anterior medial temporal lobe N400 and the hippocampal P600 may be related to the access and retrieval of person-specific semantic memory.

Dissociable brain mechanisms for inhibitory control: effects of interference content and working memory capacity.

In this study, event-related fMRI was used to examine whether the resolution of interference arising from two different information contents activates the same or different neuronal circuitries. In addition, we examined the extent to which these inhibitory control mechanisms are modulated by individual differences in working memory capacity. Two groups of participants with high and low working memory capacity [high span (HS) and low span (LS) participants, respectively] performed two versions of an item recognition task with familiar letters and abstract objects as stimulus materials. Interference costs were examined by means of the recent negative probe technique with otherwise identical testing conditions across both tasks. While the behavioral interference costs were of similar magnitude in both tasks, the underlying brain activation pattern differed between tasks: The object task interference-effects (higher activation in interference trials than in control trials) were restricted to the anterior intraparietal sulcus (IPS). Interference effects for familiar letters were obtained in the anterior IPS, the left postero-ventral and the right dorsolateral prefrontal cortex (PFC) as well as the precuneus. As the letters were more discernible than the objects, the results suggest that the critical feature for PFC and precuneus involvement in interference resolution is the saliency of stimulus-response mappings. The interference effects in the letter task were modulated by working memory capacity: LS participants showed enhanced activation for interference trials only, whereas for HS participants, who showed better performance and also lower interference costs in the letter task, the above-mentioned neuronal circuitry was activated for interference and control trials, thereby attenuating the interference effects. The latter results support the view that HS individuals allocate more attentional resources for the maintenance of task goals in the face of interfering information from preceding trials with familiar stimulus materials.

Conscious recollection and illusory recognition: an event-related fMRI study.

In this event-related functional magnetic resonance imaging (fMRI) study we examined the neuronal correlates of the subprocesses underlying recognition memory. In an explicit memory task, participants had to discriminate studied ('old') words from semantically related and unrelated 'new' (unstudied) words. We examined whether the correct rejection of semantically related words which were similar to old words, which had elicited correct responses, was based on conscious recollection of study phase information. In this task, false-positive responses to semantically related new words can be assumed to result from the assessment of the semantic similarity between test words and studied words with minimal recollection. For correct identification of old words and correct rejection of new related words we found common activation in a variety of brain areas that have been shown to be involved in conscious recollection, among them the left middle frontal gyrus, the precuneus, the retrosplenial cortex, the left parahippocampal gyrus and the thalamus. For correct responses to old words, the frontomedian wall, the posterior cingulate cortex and the nucleus accumbens were additionally activated, suggesting an emotional contribution to these judgements. Correct rejections of related new words were associated with additional activation of the right middle frontal gyrus, suggesting higher monitoring demands for these more difficult recognition judgements. False-positive responses to semantically related new words were associated with enhanced activation in the frontomedian wall. The results point to an important role of the prefrontal cortex as well as medial temporal and medial parietal regions of the brain in successful memory retrieval and conscious recollection.

Syntactic parsing preferences and their on-line revisions: a spatio-temporal analysis of event-related brain potentials.

The present study investigates the processes involved in the recovery from temporarily ambiguous garden-path sentences. Event-related brain potentials (ERP) were recorded while subjects read German subject-object ambiguous relative and complement clauses. As both clause types are initially analyzed as subject-first structures, object-first structures require a revision which is more difficult for complement than for relative clauses. The hypothesis is tested that the revision process consists of two sub-processes, namely diagnosis and actual reanalysis. Applying a spatio-temporal principal component analysis to the ERP data, distinct positive sub-components presumably reflecting different sub-processes could be identified in the time range of the P300 and P600. It will be argued that the P600 is not a monolithic component, and that different sub-processes may be involved at varying time points depending on the type of garden-path sentence.

Slow cortical potentials during retention of object, spatial, and verbal information.

We used event related potentials (ERPs) to examine both the specificity and the timing of slow cortical scalp potentials (SPs) elicited by the retention of object, spatial, and verbal information in working memory (WM). Participants performed a modified delayed matching task in which a task cue presented in the middle of the delay interval indicated what type of information had to be retained for a subsequent comparison with the test stimulus. The first experiment used nameable objects and spatial locations as stimuli. The retrieval mode (visual vs. verbal) was manipulated by presenting either figural information or printed words as test stimuli. Transient ensembles of frontal and parieto-occipital slow waves with different scalp topographies for object and spatial information were evoked as a function of task cues. When words rather than objects were used as test stimuli highly similar, though more pronounced, fronto-parietal slow wave patterns were obtained. The second experiment using unfamiliar objects and non-nameable spatial locations indicated that neither the left frontal negative SP nor the posterior SPs are exclusively related to verbal working memory operations. The results indicate that a parietal negative SP reflects processes of spatial selective attention whereas a parieto-occipital positive SP indexes the retention of visual object information. Left frontal negative SPs are generated by a compound of higher order frontal control processes and vary as a function of information type.

Repetition related ERP effects in a visual object target detection task.

ERP responses to initial and repeated presentations of possible and impossible objects were recorded from 61 recording sites in a simple target detection task. In Experiment 1, the non-target objects were line drawings of possible and impossible 3-D geometric figures and the targets were line drawings of familiar everyday objects or combinations of parts of everyday objects. In Experiment 2, the non-target objects were everyday objects and the targets were possible and impossible 3-D geometric figures. In both experiments, at frontal sites, the repeated possible and impossible non-target items elicited less negative ERP waveforms relative to first presentations between 250 and 350-400 ms. At parieto-occipital sites, in both experiments, the repeated possible and impossible non-target items elicited less positive ERP waveforms than did first presentations beginning at about 300 ms. The briefly reduced frontal negativity to repeated items is consistent with familiarity arising from a facilitation of access to conceptual, semantic and visuo-spatial representations during object categorization. The polarity of the parieto-occipital effect was the reverse of what is usually found in stimulus repetition tasks, although it is consistent with earlier work using similar visual stimuli. It is interpreted as reflecting the availability of a newly formed representation (i.e., token) of the object just experienced.

Event related brain potentials and illusory memories: the effects of differential encoding.

This study investigates event related potentials (ERP) elicited by true and false recognition using words from different semantic categories. In Experiment 1, ERPs for true and false recognition were more positive than for correctly rejected NEW words starting around 300 ms after test word presentation (old/new ERP effects). ERP waveforms for true and false recognition revealed equal early (300-500 ms) fronto-medial old/new ERP effects, reflecting similar familiarity processes, but smaller parietal old/new ERP effects (500-700 ms) for false relative to true recognition, suggesting less active recollection. Interestingly, a subsequent performance based group comparison showed equivalent old/new ERP effects for true and false recognition for participants with high rates of false recognition. In contrast, false recognition failed to elicit an old/new ERP effect in a group with low false recognition rates. To examine whether this between group difference was driven by the differential use of information that studied words and semantically related non studied test words (LURE) have in common (conceptual similarity), we manipulated encoding strategy in Experiment 2. When encoding focused on conceptual similarity, comparable ERP-effects for true and false recognition were obtained, suggesting that both forms of recognition were equally based on familiarity and recollection processes. Conversely, when encoding was focused on item specific features, differences in brain activity for true and false recognition were obtained. The ERP data indicate that, in addition to the false recognition rate, strategic processes during encoding, such as processing conceptual features, are an important factor in determining electrophysiological differences between true and false recognition.

The loci of oscillatory visual-object priming: a combined electroencephalographic and reaction-time study.

The detection of reaction-times (RTs) to a target Kanizsa-type square (an illusory square defined by the colinear arrangement of 90 degrees corner junctions) within a matrix of distractor junctions are expedited when the target display is preceded by a 40-Hz flickering display of premask crosses presented prior to, and at the locations subsequently occupied by the junctions of the target display. Priming effects were obtained when four crosses (which together matched the Gestalt arrangement of the target) were presented at the display locations subsequently occupied by the junctions forming the target Kanizsa square (Elliott and Müller, 1998, 2000). The present study was conducted with the aim of replicating the 40-Hz RT priming effects, while simultaneously recording the observers EEG in order to establish the presence and location of Gestalt priming in the brain. The statistical pattern obtained in the RT data corresponded well with previous studies and was matched by the pattern of target P300 latencies across bilateral central and posterior electrodes. Planned analyses focused upon the evoked 40-Hz activity that co-occurs with the P300, revealing a more specific pattern of 40-Hz priming over the visual cortex. A subsequent series of cross-correlational analyses examined the cortical distribution and timing of Gestalt-prime generation during and subsequent to premask-display presentation. Correlations were revealed between stimulus related 40-Hz activity over a range of cortical loci, including the right temporal lobe, which is considered important for figure coding. Taken together, these findings not only support the role of a distributed 40-Hz mechanism during Gestalt-figure priming, but also suggest that patterns of oscillatory brain activity may be directly influenced by, and interpretable in terms of equivalent temporal patterns of stimulus activity.

Magnetoencephalographic responses to illusory figures: early evoked gamma is affected by processing of stimulus features.

We examined evoked and induced responses in event-related fields and gamma activity in the magnetoencephalogram (MEG) during a visual classification task. The objective was to investigate the effects of target classification and the different levels of discrimination between certain stimulus features. We performed two experiments, which differed only in the subjects' task while the stimuli were identical. In Experiment 1, subjects responded by a button-press to rare Kanizsa squares (targets) among Kanizsa triangles and non-Kanizsa figures (standards). This task requires the processing of both stimulus features (colinearity and number of inducer disks). In Experiment 2, the four stimuli of Experiment 1 were used as standards and the occurrence of an additional stimulus without any feature overlap with the Kanizsa stimuli (a rare and highly salient red fixation cross) had to be detected. Discrimination of colinearity and number of inducer disks was not necessarily required for task performance. We applied a wavelet-based time-frequency analysis to the data and calculated topographical maps of the 40 Hz activity. The early evoked gamma activity (100-200 ms) in Experiment 1 was higher for targets as compared to standards. In Experiment 2, no significant differences were found in the gamma responses to the Kanizsa figures and non-Kanizsa figures. This pattern of results suggests that early evoked gamma activity in response to visual stimuli is affected by the targetness of a stimulus and the need to discriminate between the features of a stimulus.

What have Klingon letters and faces in common? An fMRI study on content-specific working memory systems.

Neuroimaging studies show that prefrontal, premotor, and parietal cortical regions are part of a working memory network that supports the active retention of information. In two experiments we used fMRI to examine whether prefrontal and posterior cortical areas are organized in a content-specific way for object and spatial working memory. Subjects performed a delayed matching-to-sample task modified to allow the examination of content-specific retention processes, independent of perceptual and decision-related processes. In Experiment 1, either unfamiliar geometrical objects (Klingon letters from an artificial alphabet unknown to the participants) or their spatial locations had to be memorized, whereas in Experiment 2, either unfamiliar faces or biological objects (butterflies) were actively memorized. All tasks activated a similar cortical network including posterior parietal (banks of the intraparietal sulcus), premotor (banks of the inferior precentral sulcus) and prefrontal regions (banks of the inferior frontal sulcus), and the presupplementary motor area (pre-SMA). For geometrical objects and faces for which strategic semantic processing can be assumed, this activation was larger in the left than in the right hemisphere, whereas a bilateral or right dominant distribution was obtained for butterflies and spatial locations. The present results do not support the process-specific or content-specific view of the role of the prefrontal cortex in working memory task. Rather, they suggest that the inferior prefrontal cortex houses nonmemonic strategic processing systems required for response selection and task management that can flexibly be used across a variety of tasks and informational domains.

Transient global ischemia specifically modulates visual P300 scalp distribution.

OBJECTIVE: Latency, amplitude, and scalp topography of the visual P300 component was examined in patients who had suffered from transient global ischemia (TGI) due to cardiac arrest and in age matched clinical and healthy controls in order to investigate the diagnostic value of this component. METHOD: Event-related potentials (ERPs) were recorded from 19 scalp electrodes in a visual oddball paradigm. RESULTS: Mean latency of the P300 component was prolonged in both patient groups. Changes in scalp distribution of the P300, however, appear to be specific to anoxic-ischemic encephalopathy. In particular, a selective reduction of the P300 amplitudes at posterior recording sites was observed in TGI patients. Moreover, examination of the auditory P300 in TGI patients revealed that this selective change seems to be restricted to the visual modality. CONCLUSION: The results are discussed with respect to selective vulnerability of brain tissue to hypoxic-ischemic injury. After TGI a modality-specific subset of P300 generators, probably located in the transitional parieto-occipital and extrastriate occipital cortex, appears to be affected. It is also noted, that the visual P300 component could serve as an additional marker of TGI especially in patients who do not show neuropathological changes in structural brain images.

Interfacing mind and brain: a neurocognitive model of recognition memory.

A variety of processes contribute to successful recognition memory, some of which can be associated with spatiotemporally distinct event-related potential old/new effects. An early frontal and a subsequent parietal old/new effect are correlated with the familiarity and recollection subcomponents of recognition memory, respectively, whereas a late, postretrieval old/new effect seems to reflect an ensemble of evaluation processes that are set by the task context in which retrieval occurs. Both the early frontal and the parietal old/new effects are differentially modulated by the informational content (e.g., object forms and spatial locations) of recognition and seem to rely on brain systems damaged in amnesia. The late frontal effect appears to reflect prefrontal cortex activation. A neurophysiologically based model of recognition memory retrieval is presented and it is shown that coupling recognition memory subprocesses with distinct old/new effects allow examination of the time course of the processes that contribute to correct and to illusory memories. In conjunction with event-related functional magnetic resonance imaging activation patterns the brain systems recruited by various aspects of episodic memory retrieval can be identified.

Attentive and pre-attentive periodicity analysis in auditory memory: an event-related brain potential study.

In an event-related potential (ERP) study we presented human subjects with streams of repetitive white noise (semi-periodic noise) under attend and ignore conditions to investigate whether the perception of the periodicity with short cycle-lengths is due to lower level, pre-attentive sensory memory processing or higher level, attentive working memory processing. The ERPs of both conditions reveal N1-like deflections that are time locked on the semi-periodic noise suggesting that the processing of the periodicity is due to a pre-attentive rather than an attentive process. The topography of the deflections suggests that its generators are located in the supratemporal plane. Additionally, the ERPs elicited by infrequent disruptions in the periodicity show differences between the conditions suggesting that the detection of disruptions in periodicity is facilitated by attention.

Event-related brain potentials dissociate visual working memory processes under categorial and identical comparison conditions.

Event-related potentials (ERPs) have been successfully employed to examine the functional and neuronal characteristics of working memory processes. In the present study, we examined the ERP waveforms in a delayed matching task to examine the cognitive processes underlying category and identity comparison and the effects of stimulus complexity. Subjects had to decide whether two visual stimuli are (a) physically identical (identical comparison condition, IC) or (b) identical, irrespective of their orientation (categorial comparison condition, CC). The stimuli were structured five-point patterns, which varied in complexity. For the ERPs elicited during the 1500 ms retention interval, the following pattern of results was obtained: Stimuli in the CC-condition elicited larger P300 components than in the IC-condition. In the IC-condition, the P300 was followed by a broadly distributed negative slow wave. Moreover, complex patterns elicited a posteriorily distributed negativity at 350 ms (N350), whereas the less complex patterns gave rise to a fronto-centrally distributed slow wave that started around 500 ms. These results suggest that S1 was more elaborately processed in the CC-condition, while the more complex figures were associated with an early classification process during the retention interval.

An electrophysiological test of directed forgetting: the role of retrieval inhibition.

A central issue in the research of directed forgetting is whether the differential memory performance for to-be-remembered (TBR) and to-be-forgotten (TBF) items is solely due to differential encoding or whether retrieval inhibition of TBF items plays an additional role. In this study, recognition-related event-related brain potentials (ERPs) were used to examine this issue. The spatio-temporal distributions of the old/new ERP effects obtained in Experiment 1 that employed a directed forgetting paradigm were compared with those recorded in Experiment 2 in which the level of processing was manipulated. In Experiment 1, participants were instructed to remember or to forget words by means of a cue presented after each word. ERPs recorded in the recognition test revealed early phasic frontal and parietal old/new effects for TBR items, whereas TBF items elicited only a frontal old/new effect. Moreover, a late right-frontal positive slow wave was more pronounced for TBF items, suggesting that those items were associated with a larger amount of post-retrieval processing. In Experiment 2, the same cueing method and the same stimulus materials were used, and memory encoding was manipulated by cueing participants to process the words either deeply or shallowly. Both deeply and shallowly encoded items elicited phasic frontal and parietal old/new effects followed by a late right-frontal positive slow wave. However, in contrast to TBR and TBF items, these effects differed only quantitatively. The results suggest that differential encoding alone cannot account for the effects of directed forgetting. They are more consistent with the view that items followed by an instruction to forget become inhibited and less accessible, and, therefore, more difficult to retrieve.

Functional asymmetry of human prefrontal cortex: encoding and retrieval of verbally and nonverbally coded information.

There are several views about the organization of memory functions in the human prefrontal cortex. One view assumes a process-specific brain lateralization according to different memory subprocesses, that is, encoding and retrieval. An alternative view emphasizes content-specific lateralization of brain systems involved in memory processes. This study addresses this apparent inconsistency between process- and content-specific lateralization of brain activity by investigating the effects of verbal and nonverbal encoding on prefrontal activations during encoding and retrieval of environmental novel sounds using fMRI. An intentional memory task was applied in which subjects were required either to judge the sounds' loudness (nonverbal encoding task) or to indicate whether or not a sound can be verbally described (verbal encoding task). Retrieval processes were examined in a subsequent yes/no recognition test. In the study phase the right posterior dorsolateral prefrontal cortex (PFC) was activated in both tasks. During verbal encoding additional activation of the left dorsolateral PFC was obtained. Retrieval-related fMRI activity varied as a function of encoding task: For the nonverbal task we detected an activation focus in the right posterior dorsolateral PFC whereas an activation in the left dorsolateral PFC was observed for the verbal task. These findings indicate that the right dorsolateral PFC is engaged in encoding of auditory information irrespective of encoding task. The lateralization of PFC activity during retrieval was shown to depend on the availability of verbal codes, with left hemispheric involvement for verbally and right hemispheric activation for nonverbally coded information.

Executive control functions in task switching: evidence from brain injured patients.

Executive mechanisms involved in task switching were studied in 18 brain injured patients. The patients had to rapidly switch back and forth between two visual classification tasks and the analyses focused on switch costs, (i.e., performance differences between switch and no-switch trials), and on interference effects, (i.e., processing costs imposed by the presence of interfering stimulus attributes). The patients were grouped according to the side of the brain lesion. Patients with left brain damage (LBD) showed higher switch costs than patients with right brain damage (RBD). These group differences were attributable to disproportionally high switch costs in patients with LBD and language or speech disorders. This result suggests that the efficiency of suppressing internal interference from a recently activated task set depends on the availability of verbal representations of the upcoming task. Patients with RBD showed higher interference from external task sets. This effect was not affected by language or speech disorders. The overall results argue for a fractionation of executive functions to protect against interference from internal and external sources in task switching.

The functional neuroanatomy of novelty processing: integrating ERP and fMRI results.

Recent research indicates that non-tonal novel events, deviating from an ongoing auditory environment, elicit a positive event-related potential (ERP), the novel P3. Although a variety of studies examined the neural network engaged in novelty detection, there is no complete picture of the underlying brain mechanisms. This experiment investigated these neural mechanisms by combining ERP and functional magnetic resonance imaging (fMRI). Hemodynamic and electrophysiological responses were measured in the same subjects using the same experimental design. The ERP analysis revealed a novel P3, while the fMRI responses showed bilateral foci in the middle part of the superior temporal gyrus. When subjects attended to the novel stimuli only identifiable novel sounds evoked a N4-like negativity. Subjects showing a strong N4-effect had additional fMRI activation in right prefrontal cortex (rPFC) as compared to subjects with a weak N4-effect. This pattern of results suggests that novelty processing not only includes the registration of deviancy but may also lead to a fast access and retrieval of related semantic concepts. The fMRI activation pattern suggests that the superior temporal gyrus is involved in novelty detection, whereas accessing and retrieving semantic concepts related to novel sounds additionally engages the rPFC.