Age-related differences in enhancement and suppression of neural activity underlying selective attention in matched young and old adults.

Selective attention reflects the top-down control of sensory processing that is mediated by enhancement or inhibition of neural activity. ERPs were used to investigate age-related differences in neural activity in an experiment examining selective attention to color under Attend and Ignore conditions, as well as under a Neutral condition in which color was task-irrelevant. We sought to determine whether differences in neural activity between old and young adult subjects were due to differences in age rather than executive capacity. Old subjects were matched to two groups of young subjects on the basis of neuropsychological test performance: one using age-appropriate norms and the other using test scores not adjusted for age. We found that old and young subject groups did not differ in the overall modulation of selective attention between Attend and Ignore conditions, as indexed by the size of the anterior Selection Positivity. However, in contrast to either young adult group, old subjects did not exhibit reduced neural activity under the Ignore relative to Neutral condition, but showed enhanced activity under the Attend condition. The onset and peak of the Selection Positivity occurred later for old than young subjects. In summary, older adults execute selective attention less efficiently than matched younger subjects, with slowed processing and failed suppression under Ignore. Increased enhancement under Attend may serve as a compensatory mechanism.

Frontal and parietal components of a cerebral network mediating voluntary attention to novel events.

Despite the important role that attending to novel events plays in human behavior, there is limited information about the neuroanatomical underpinnings of this vital activity. This study investigated the relative contributions of the frontal and posterior parietal lobes to the differential processing of novel and target stimuli under an experimental condition in which subjects actively directed attention to novel events. Event-related potentials were recorded from well-matched frontal patients, parietal patients, and non-brain-injured subjects who controlled their viewing duration (by button press) of line drawings that included a frequent, repetitive background stimulus, an infrequent target stimulus, and infrequent, novel visual stimuli. Subjects also responded to target stimuli by pressing a foot pedal. Damage to the frontal cortex resulted in a much greater disruption of response to novel stimuli than to designated targets. Frontal patients exhibited a widely distributed, profound reduction of the novelty P3 response and a marked diminution of the viewing duration of novel events. In contrast, damage to posterior parietal lobes was associated with a substantial reduction of both target P3 and novelty P3 amplitude; however, there was less disruption of the processing of novel than of target stimuli. We conclude that two nodes of the neuroanatomical network for responding to and processing novelty are the prefrontal and posterior parietal regions, which participate in the voluntary allocation of attention to novel events. Injury to this network is indexed by reduced novelty P3 amplitude, which is tightly associated with diminished attention to novel stimuli. The prefrontal cortex may serve as the central node in determining the allocation of attentional resources to novel events, whereas the posterior parietal lobe may provide the neural substrate for the dynamic process of updating one's internal model of the environment to take into account a novel event.

Phonological processing in visual rhyming: a developmental erp study.

We employed a visual rhyming priming paradigm to characterize the development of brain systems important for phonological processing in reading. We studied 109 righthanded, native English speakers within eight age groups: 7-8, 9-10, 11-12, 13-14, 15-16, 17-18, 19-20, and 21-23. Participants decided whether two written words (prime-target) rhymed (JUICE-MOOSE) or not (CHAIR-MOOSE). In similar studies of adults, two main event-related potential (ERP) effects have been described: a negative slow wave to primes, larger over anterior regions of the left hemisphere and hypothesized to index rehearsal of the primes, and a negative deflection to targets, peaking at 400-450 msec, maximal over right temporal-parietal regions, larger for nonrhyming than rhyming targets, and hypothesized index phonological matching. In this study, these two ERP effects were observed in all age groups; however, the two effects showed different developmental timecourses. On the one hand, the frontal asymmetry to primes increased with age; moreover, this asymmetry was correlated with reading and spelling scores, even after controlling for age. On the other hand, the distribution and onset of the more posterior rhyming effect (RE) were stable across age groups, suggesting that phonological matching relied on similar neural systems across these ages. Behaviorally, both reaction times and accuracy improved with age. These results suggest that different aspects of phonological processing rely on different neural systems that have different to developmental timecourses.

Is the donut in front of the car? An electrophysiological study examining spatial reference frame processing.

The process of describing an object's location relative to another object results in ambiguity. How do people handle this ambiguity? The present studies examined spatial language processing when use of different reference frames results in ambiguity. We investigated whether electrophysiological (ERP) measures of cognitive processing may elucidate underlying reference frame processing; in particular, we were interested in semantic integration. ERP results showed a larger N400, peaking between 300 and 375 ms, when the intrinsic frame was not used. Behavioural results mirrored this finding, indicating a reduced cognitive processing requirement for the intrinsic reference frame. Previous work has not definitively tied spatial reference frame processing to specific ERP components and their associated cognitive processes. Although the N400 peak seen in this data is early, additional work supports the N400 interpretation, thereby linking spatial frame processing to semantic integration. Results are discussed within the larger context of spatial reference frame processing.

Pathophysiology underlying diminished attention to novel events in patients with early AD.

BACKGROUND: Patients with mild to moderate AD often are apathetic and fail to attend to novel aspects of their environment. OBJECTIVE: To investigate the mechanisms underlying these changes by studying the novelty P3 response that measures shifts of attention toward novel events. METHODS: While event-related potentials were recorded, mildly impaired AD patients and matched normal controls (NC) viewed line drawings that included a repetitive background stimulus, an infrequent target stimulus, and infrequent novel stimuli. Subjects controlled how long they viewed each stimulus by pressing a button. This served as a measure of their allocation of attention. They also responded to targets by depressing a foot pedal. Patients did not differ from NC in age, education, estimated IQ, or mood but were judged by informants to be more apathetic. RESULTS: P3 amplitude to novel stimuli was significantly smaller for AD patients than NC. However, P3 amplitude to target stimuli did not differ between groups. For NC, P3 response to novel stimuli was much larger than to background stimuli. In contrast, for patients with AD, there was no difference in P3 response to novel vs background stimuli. Although NC spent more time looking at novel than background stimuli, patients with AD distributed their viewing time evenly. Remarkably, for patients with AD, the amplitude of the novelty P3 response powerfully predicted how long they would spend looking at novel stimuli (R2 = 0.52) and inversely correlated with apathy severity. CONCLUSIONS: The decreased attention to novel events exhibited by patients with AD cannot be explained by a nonspecific reduction in their attentional abilities. The novelty P3 response is markedly diminished in mild AD, at a time when the target P3 response is preserved. The disruption of the novelty P3 response predicts diminished attention to novel stimuli and is associated with the apathy exhibited by patients with AD.

An electrophysiological index of stimulus unfamiliarity.

This study investigated the functional significance of the N2 response to novel stimuli. In one condition, background, target, and deviant stimuli were simple geometric figures. In a second condition, all stimulus types were unfamiliar/unusual figures. In a third condition, background and target stimuli were unusual figures and deviant stimuli were simple shapes. Unusual figures, whether they were deviant, target, or background stimuli, evoked larger N2 responses than their simple, familiar counterparts. N2 elicited by an unusual background stimulus was larger than that evoked by simple, deviant stimuli, a pattern opposite that exhibited by the subsequent P3. Deviance from immediate context had limited influence over N2 amplitude. The results suggest that novelty N2 and novelty P3 reflect the processing of different aspects of "novel" visual stimuli. The novelty P3 is particularly sensitive to deviation from immediate context. In contrast, the novelty N2 is sensitive to deviation from long-term context that renders a stimulus unfamiliar and difficult to encode.

The influence of stimulus deviance on electrophysiologic and behavioral responses to novel events.

This study investigated the role of stimulus deviance in determining electrophysiologic and behavioral responses to "novelty." Stimulus deviance was defined in terms of differences either from the immediately preceding context or from long-term experience. Subjects participated in a visual event-related potential (ERP) experiment, in which they controlled the duration of stimulus viewing with a button press, which served as a measure of exploratory behavior. Each of the three experimental conditions included a frequent repetitive background stimulus and infrequent stimuli that deviated from the background stimulus. In one condition, both background and deviant stimuli were simple, easily recognizable geometric figures. In another condition, both background and deviant stimuli were unusual/unfamiliar figures, and in a third condition, the background stimulus was a highly unusual figure, and the deviant stimuli were simple, geometric shapes. Deviant stimuli elicited larger N2-P3 amplitudes and longer viewing durations than the repetitive background stimulus, even when the deviant stimuli were simple, familiar shapes and the background stimulus was a highly unusual figure. Compared to simple, familiar deviant stimuli, unusual deviant stimuli elicited larger N2-P3 amplitudes and longer viewing times. Within subjects, the deviant stimuli that evoked the largest N2-P3 responses also elicited the longest viewing durations. We conclude that deviance from both immediate context and long-term prior experience contribute to the response to novelty, with the combination generating the largest N2-P3 amplitude and the most sustained attention. The amplitude of the N2-P3 may reflect how much "uncertainty" is evoked by a novel visual stimulus and signal the need for further exploration and cognitive processing.

Imaginal, semantic, and surface-level processing of concrete and abstract words: an electrophysiological investigation.

Words representing concrete concepts are processed more quickly and efficiently than words representing abstract concepts. Concreteness effects have also been observed in studies using event-related brain potentials (ERPs). The aim of this study was to examine concrete and abstract words using both reaction time (RT) and ERP measurements to determine (1) at what point in the stream of cognitive processing concreteness effects emerge and (2) how different types of cognitive operations influence these concreteness effects. Three groups of subjects performed a sentence verification task in which the final word of each sentence was concrete or abstract. For each group the truthfulness judgment required either (1) image generation, (2) semantic decision, or (3) evaluation of surface characteristics. Concrete and abstract words produced similar RTs and ERPs in the surface task, suggesting that postlexical semantic processing is necessary to elicit concreteness effects. In both the semantic and imagery tasks, RTs were shorter for concrete than for abstract words. This difference was greatest in the imagery task. Also, in both of these tasks concrete words elicited more negative ERPs than abstract words between 300 and 550 msec (N400). This effect was widespread across the scalp and may reflect activation in a linguistic semantic system common to both concrete and abstract words. ERPs were also more negative for concrete than abstract words between 550 and 800 msec. This effect was more frontally distributed and was most evident in the imagery task. We propose that this later anterior effect represents a distinct ERP component (N700) that is sensitive to the use of mental imagery. The N700 may reflect the a access of specific characteristics of the imaged item or activation in a working memory system specific to mental imagery. These results also support the extended dual-coding hypothesis that superior associative connections and the use of mental imagery both contribute to processing advantages for concrete words over abstract words.

On the locus of the semantic satiation effect: evidence from event-related brain potentials.

The present study sought to determine whether semantic satiation is merely a by-product of adaptation or satiation of upstream, nonsemantic perceptual processes or whether the effect can have a locus in semantic memory. This was done by measuring event-related brain potentials (ERPs) in a semantic word-detection task involving multiple presentations of primes and critical related and unrelated words in three experiments involving visual (Experiment 1) and auditory (Experiments 2A and 2B) stimuli. Primes varied in their type case (Experiment 1) or pitch (Experiment 2B) in order to discourage sensory adaptation. Prime satiation and relatedness of the primes to the critical word had interacting effects on ERP amplitude to critical words, particularly within the time-window of the N400 component. Because numerous studies have indicated a role for the N400 in semantic processing, modulation of the N400 relatedness effect by prime satiation (with little or no contribution from perceptual adaptation) suggests that semantic memory can be directly satiated, rather than the cost to semantic processing necessarily resulting from impoverishment of perceptual inputs.

Disruption of attention to novel events after frontal lobe injury in humans.

OBJECTIVE: To investigate whether frontal lobe damage in humans disrupts the natural tendency to preferentially attend to novel visual events in the environment. METHODS: Nine patients with chronic infarctions in the dorsolateral prefrontal cortex (DLPFC) and 23 matched normal controls participated in a study in which subjects viewed repetitive background stimuli, infrequent target stimuli, and novel visual stimuli (for example, fragmented or "impossible" objects). Subjects controlled viewing duration by a button press that led to the onset of the next stimulus. They also responded to targets by pressing a foot pedal. The amount of time spent looking at the different kinds of stimuli, and the target detection accuracy and speed served as dependent variables. RESULTS: Overall, normal controls spent significantly more time than frontal lobe patients looking at novel stimuli. Analysis of responses across blocks showed that initially frontal lobe patients behaved like normal controls by directing more attention to novel than background stimuli. However, they quickly began to distribute their viewing time evenly between novel and background stimuli, a pattern that was strikingly different from normal controls. By contrast, there were no differences between frontal lobe patients and normal controls for viewing duration devoted to background and target stimuli, target detection accuracy, or reaction time to targets. Frontal lobe patients did not differ from normal controls in terms of age, education, estimated IQ, or mood, but were more apathetic as measured by self report and informants' judgments. Attenuated responses to novel stimuli significantly correlated with degree of apathy. CONCLUSIONS: This study demonstrates that DLPFC injury selectively impairs the natural tendency to seek stimulation from novel and unusual stimuli. These data provide the first quantitative behavioural demonstration that the human frontal lobes play a critical part in directing and sustaining attention to novel events. The impairment of novelty seeking behaviour may contribute to the characteristic apathy found in patients with frontal lobe injury.

Dual-coding, context-availability, and concreteness effects in sentence comprehension: an electrophysiological investigation.

Event-related potentials were recorded in 2 experiments while participants read sentences in a word-by-word congruency judgment task. Sentence final words were either congruent, semantically anomalous (Experiments 1 and 2), or neutral (Experiment 2) with respect to sentence context. Half of all final words referred to concrete and half to abstract concepts. A different scalp distribution of the N400 to concrete and abstract final words was found for anomalous and neutral, but not congruent sentences. Although the interaction of context and concreteness is consistent with the context-availability model, the differential scalp distribution of effects for concrete and abstract words, as well as larger context effects for concrete words, was interpreted as being more consistent with an extended dual-code account of semantic processing.

An electrophysiological investigation of semantic priming with pictures of real objects.

Event-related potentials were recorded using color pictures of real objects. Participants made relatedness judgments for pictures that were highly, moderately, or unrelated to a picture of a preceding prime object (Experiment 1) or object identification decisions for related/easily identified, unrelated/easily identified, and unrelated/unidentifiable objects preceded by prime objects (Experiment 2). Unrelated pictures elicited larger event-related potential negativities between 225 and 500 ms than did related pictures, although the first portion of this epoch had a more frontal distribution than did the later portion. The later epoch differentiated the unrelated from the moderately related and the moderately related from the highly related pictures (Experiment 1), but the early epoch produced differences only between the unrelated and related pictures (Experiments 1 and 2). This pattern supports the existence of two separate components, an anterior, image-specific N300 and a later, central/parietal amodal N400.

Processing syntactic relations in language and music: an event-related potential study.

In order to test the language-specificity of a known neural correlate of syntactic processing [the P600 event-related brain potential (ERP) component], this study directly compared ERPs elicited by syntactic incongruities in language and music. Using principles of phrase structure for language and principles of harmony and key-relatedness for music, sequences were constructed in which an element was either congruous, moderately incongruous, or highly incongruous with the preceding structural context. A within-subjects design using 15 musically educated adults revealed that linguistic and musical structural incongruities elicited positivities that were statistically indistinguishable in a specified latency range. In contrast, a music-specific ERP component was observed that showed antero-temporal right-hemisphere lateralization. The results argue against the language-specificity of the P600 and suggest that language and music can be studied in parallel to address questions of neural specificity in cognitive processing.

Event-related brain potentials elicited during phonological processing differentiate subgroups of reading disabled adolescents.

Visual and auditory rhyme judgment tasks were administered to adolescent dyslexics and normal readers while event-related brain potentials were recorded. Reading disabled subjects were split into two groups based on a median split of scores on a visual non-word decoding test. The better decoders were called Phonetics and the poorer decoders were referred to as Dysphonetics. Single syllable, real word stimuli were used, and both rhyming and non-rhyming targets had a 50% chance for matching orthography. In the visual paradigm the normal readers exhibited a left frontal CNV before targets, a large reduction in frontal N400 for matching orthography (orthographic priming), and a large reduction in parietal N400 for rhyming targets (phonological priming). Dysphonetics had an intact CNV and orthographic priming, but the group's phonological priming was very reduced. Phonetics showed both orthographic and phonological priming but had a marked reduction in their CNV. In the auditory task, controls showed a left parietal N400 priming effect for rhyming targets. Dysphonetics showed a similar bilateral effect. The Phonetics did not show a normal priming effect, but produced evidence for priming at a longer latency. Additionally, the Phonetic group responded more slowly than either of the other two groups, who responded with similar latencies. These results support the separation of the reading disabled into a group that has difficulty translating orthography into phonology, and a group that is slower functioning and has reduced capacity in preparing for a response.

Regulation of attention to novel stimuli by frontal lobes: an event-related potential study.

This study examined the relationship between orienting responses to novel events and subsequent exploratory behavior. The N2-P3 electrophysiologic component of the orienting response was found to be larger for novel than repetitive background stimuli. Across subjects, the amplitude of this N2-P3 response in frontal regions strongly predicted the proportional increase in the duration of viewing directed toward novel compared to background stimuli. Within subjects, larger N2-P3 amplitudes in response to novel stimuli were associated with longer viewing durations on those stimuli. These results suggest that the N2-P3 component of the orienting response reflects the activity of a neural system involving frontal networks that dynamically regulates the subsequent allocation of attentional resources to novel stimuli.

Auditory and visual semantic priming using different stimulus onset asynchronies: an event-related brain potential study.

Semantic priming effects (behavioral and electrophysiological) were compared in the visual and auditory modalities across three stimulus onset asynchronies (SOAs; 0, 200, and 800 ms). When both prime and target were presented in the visual modality (the prime just to the left of a fixation point and the target to the right), there were N400 priming effects present across the three SOAs. However, the N400 in the 0-ms SOA condition extended longer in time (800 vs. 500 ms) than in the other SOAs. When both the prime and target were presented in the auditory modality (the prime to the right ear and the target to the left), the largest priming effects were found for the 800-ms SOA. Moreover, there was a relatively early priming effect present in the 0- and 800-ms SOA conditions but not in the 200-ms condition. The results are discussed in terms of modality differences in the time course of word comprehension processes.

Brain potentials elicited by garden-path sentences: evidence of the application of verb information during parsing.

Event-related potentials were recorded from 13 scalp locations while participants read sentences containing a syntactic ambiguity. In Experiment 1, syntactically disambiguating words that were inconsistent with the "favored" syntactic analysis elicited a positive-going brain potential (P600). Experiment 2 examined whether syntactic ambiguities are resolved by application of a phrase-structure-based minimal attachment principle or by word-specific subcategorization information. P600 amplitude was a function of subcategorization biases rather than syntactic complexity. These findings indicate that such biases exist and can influence the parser under certain conditions and that P600 amplitude is a function of the perceived syntactic well-formedness of the sentence.

Concreteness effects in semantic processing: ERP evidence supporting dual-coding theory.

Dual-coding theory argues that processing advantages for concrete over abstract (verbal) stimuli result from the operation of 2 systems (i.e., imaginal and verbal) for concrete stimuli, rather than just 1 (for abstract stimuli). These verbal and imaginal systems have been linked with the left and right hemispheres of the brain, respectively. Context-availability theory argues that concreteness effects result from processing differences in a single system. The merits of these theories were investigated by examining the topographic distribution of event-related brain potentials in 2 experiments (lexical decision and concrete-abstract classification). The results were most consistent with dual-coding theory. In particular, different scalp distributions of an N400-like negativity were elicited by concrete and abstract words.

Event-related brain potentials reflect semantic priming in an object decision task.

Subjects made speeded object decisions about target line drawings which were preceded by semantically related or unrelated prime line drawings. One hundred of the targets depicted real objects and 50 others were unrecognizable non-objects. Similar to a recent picture-matching study, the ERPs from this study generated a larger negativity for unrelated than for related target pictures, between 325 and 550 msec (N400). Although these differences had a similar time course to those seen for the N400 component in semantic priming lexical decision tasks, they were more frontally distributed and were larger over the left rather than the right hemisphere. Non-objects, which were the picture equivalent of pseudowords, produced an even larger negativity with a somewhat different distribution. The results are discussed with regards to recent claims about amodal conceptual memory processes.