Individual differences in nonverbal number discrimination correlate with event-related potentials and measures of probabilistic reasoning

The current study investigated the neural activity patterns associated with numerical sensitivity in adults. Event-related potentials (ERPs) were recorded while adults observed sequentially presented display arrays (S1 and S2) of non-symbolic numerical stimuli (dots) and made same/different judgments of these stimuli by pressing a button only when numerosities were the same (target trials). The main goals were to contrast the effects of numerical distance (close, medium, and far) and change direction (increasing, decreasing) between S1 and S2, both in terms of behavior and brain activity, and to examine the influence of individual differences in numeracy on the effects of these manipulations. Neural effects of distance were found to be significant between 360 and 600 ms after the onset of S2 (greater negativity-wave activity for closer numerical distances), while direction effects were found between 320 and 440 ms (greater negativity for decreasing direction). ERP change direction effects did not interact with numerical distance, suggesting that the two types of information are processed independently. Importantly, subjects’ behavioral Weber fractions (w) for the same/different discrimination task correlated with distance-related ERP-activity amplitudes. Moreover, w also correlated with a separate objective measure of mathematical ability. Results thus draw a clear link between brain and behavior measures of number discrimination, while also providing support for the relationship between nonverbal magnitude discrimination and symbolic numerical processing.     

Electrophysiological analysis of cortical mechanisms of selective attention to high and low spatial frequencies.

OBJECTIVES: This study investigated whether short-latency (<100ms) event-related potential (ERP) components were modulated during attention to spatial frequency (SF) cues. METHODS: Sinusoidally modulated checkerboard stimuli having high (5 cycles per degree (cpd)) or low (0.8cpd) SF content were presented in random order at intervals of 400-650ms. Subjects attended to either the high or low SF stimuli, with the task of detecting targets of slightly higher or lower SF, respectively, than the above standards. ERPs were recorded from 42 scalp sites during task performance and spatio-temporal analyses were carried out on sensory-evoked and attention-related components. RESULTS: Attended high SF stimuli elicited an early negative difference potential (ND120) starting at about 100ms, whereas attended low SF stimuli elicited a positivity (PD130) in the same latency range. The neural sources of both effects were estimated with dipole modeling to lie in dorsal, extrastriate occipital areas. Earlier evoked components evoked at 60-100ms that were modeled with striate and extrastriate cortical sources were not affected by attention to SF. Starting at 150ms, attended stimuli of both SFs elicited a broad selection negativity (SN) that was localized to ventral extrastriate visual cortex. The SN was larger over the left/right cerebral hemisphere for attended stimuli of high/low SF. CONCLUSIONS: These results support the view that attention to SF does not involve a mechanism of amplitude modulation of early-evoked components prior to 100ms. Attention to high and low SF information involves qualitatively different and hemispherically specialized neural processing operations.     

Processing the universal quantifier during sentence comprehension: ERP evidence

This study investigates the temporal neural dynamics of processing the Chinese universal quantifier dou during Chinese sentence comprehension using the event-related potential (ERP) technique. Universal quantifier violations were created when the universal quantifier dou (all, every) was misplaced either after a singular object noun phrase (NP) in a Subject–Object–Verb (SOV) sentence (Experiments 1 and 3) or after a singular subject NP in a SVO sentence (Experiment 2). Participants were asked to make semantic plausibility judgment (Experiments 1 and 2) or to comprehend sentences real time followed by a sentence recognition test at the end of the experiment (Experiment 3). Experiment 1 found that quantifier violations elicited a sustained positivity from 400 to 1100 ms post-onset of the quantifier and a sustained negativity from 300 to 800 ms post-onset of the following verb. Experiment 2 varied the distance between dou and the following verb by the presence or absence of an adverb between them. Again, the sustained positivity was observed on the mismatching quantifier; in addition, a sustained negativity was observed on the word immediately following the quantifier, regardless of whether this word was a verb or adverb. Experiment 3 used the same stimuli as Experiment 1 but with a different task. The quantifier violation elicited anteriorly distributed negativities over different time intervals post-onset of the quantifier. The sustained positivity is interpreted as being associated with an integration process that links the universal quantifier with the preceding entity. The sustained negativity is attributed to a second-pass process to reinterpret the sentence. Other functional interpretations of the ERP components were discussed and ruled out.     

Predictive value of novel stimuli modifies visual event-related potentials and behavior.

OBJECTIVE: We examined how behavioral context influences novelty processing by varying the degree that a novel event predicted the occurrence of a subsequent target stimulus. METHODS: Visual event-related potentials (ERPs) and reaction times (RTs) were recorded in 3 detection experiments (23 subjects). The predictive value of a novel stimulus on the occurrence of a subsequent target was varied as was novel-target pairing intervals (200-900 ms). In Experiment 1, novel stimuli always preceded a target, in Experiment 2, 40% of novel stimuli were followed by a target, and in Experiment 3, novel stimuli occurred randomly. RESULTS: In Experiment 1, RTs following 100% predictive novels were shortened for targets at all spatial locations and novel-target pairing intervals. Novel stimuli predicting a target generated a central negativity peaking at 300 ms and reduced P3a and P3b ERPs. In Experiments 2 and 3, target RTs were prolonged only when novel and target stimuli were presented in the same spatial location at short ISIs (200 ms). The central novel N2 was smaller in amplitude in comparison to Experiment 1, and novelty P3a and target extrastriate N2 and posterior scalp P3b ERPs were enhanced. CONCLUSIONS: The enhanced N2 for 100% predictive novel stimuli appears to index an alerting system facilitating behavioral detection. The same novel stimuli with no predictive value distract attention and generate a different ERP pattern characterized by increased novelty P3a and target P3b responses. The results indicate that behavioral context determines how novel stimuli are processed and influence behavior.     

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.     

Cueing effects on semantic and perceptual categorization: ERPs reveal differential effects of validity as a function of processing stage

Valid cueing has been shown to accelerate target identification and improve decision accuracy. However, the precise nature and extent to which biasing influences the successive stages of target processing remain unclear. The present event-related potential (ERP) study used a "hybrid" task that combined features of standard cued-attention and task-switching paradigms in order to explore the effects of expectation on both identification and categorization of centrally presented stimuli. Subjects made semantic judgments (living/nonliving) on word targets ("bunny"), and perceptual judgments (right/left) on arrow targets ("<"). Target expectancy was manipulated using cues that were valid (60 percent of trials), invalid (10 percent), or neutral (30 percent). Invalidly cued targets required task-set switching before categorization could commence, and resulted in RT costs relative to validly or neutrally cued targets. Additional benefits from valid-cueing were only observed for word targets. Invalid cueing of both arrow and word targets modulated early posterior visual potentials (P1/N1) and elicited a subsequent anterior P3a (270 ms). The temporal relationship of these effects suggests that the P3a indexed domain-general task-set switching processes recruited in response to the detection of unexpected perceptual information. Subsequent to the P3a and immediately preceding the behavioral response, validly cued targets elicited enhanced stimulus-specific waveforms (arrows: parietal positivity [P290], words: inferior temporal negativity [late ITN: 400-600 ms]). The degree of neural enhancement relative to the invalid and neutral conditions mirrored the magnitude of corresponding RT benefits, suggesting that these waveforms indexed categorization, decision processes or both. Together, these results suggest that valid cueing increases the neural efficiency of initial stimulus identification, facilitating transmission of information to subsequent categorization stages, where increased neural activity leads to behavioral benefits.     

Effects of spatial separation and stimulus probability on the event-related potentials elicited by occasional changes in sound location

The ability to extract information about the spatial location of sounds plays an important role in auditory scene analysis. The present study examined the effects of spatial separation and stimulus probability on auditory event-related potentials (ERPs) to changes in sound location. In Experiment 1, we found that difference waves between ERPs elicited by standard and deviant stimuli showed a biphasic negative-positive response peaking around 126 and 226 ms after deviant onset. The amplitude of both responses increased with decreasing deviant stimulus probability, and increasing stimulus deviance. When the same stimuli were presented with equal probability for all locations (Experiment 2), there were no significant differences in the ERP amplitude and latency. These results suggest that the data reported in Experiment 1 are the result of contextual changes, rather than changes in simple acoustic features. Brain electrical source analyses are consistent with generators located in auditory cortices posterior to Heschel's gyrus. Although occasional changes in sound location elicit earlier peaks than the mismatch negativity (MMN) response reported for other types of deviation, their topographical distribution and behavior are consistent with MMN. The early latency of MMN for changes in sound location is interpreted in the context of an early-warning system to alert the organism to new sound sources in the environment.     

An electrophysiological investigation of non-symbolic magnitude processing: Numerical distance effects in children with and without mathematical learning disabilities

IntroductionThe aim of the present study was to probe electrophysiological effects of non-symbolic numerical processing in 20 children with mathematical learning disabilities (mean age = 99.2 months) compared to a group of 20 typically developing matched controls (mean age = 98.4 months).MethodsEEG data were obtained while children were tested with a standard non-symbolic numerical comparison paradigm that allowed us to investigate the effects of numerical distance manipulations for different set sizes, i.e., the classical subitizing, counting and estimation ranges. Effects of numerical distance manipulations on event-related potential (ERP) amplitudes as well as activation patterns of underlying current sources were analyzed.ResultsIn typically developing children, the amplitudes of a late parietal positive-going ERP component showed systematic numerical distance effects that did not depend on set size. For the group of children with mathematical learning disabilities, ERP distance effects were found only for stimuli within the subitizing range. Current source density analysis of distance-related group effects suggested that areas in right inferior parietal regions are involved in the generation of the parietal ERP amplitude differences.ConclusionOur results suggest that right inferior parietal regions are recruited differentially by controls compared to children with mathematical learning disabilities in response to non-symbolic numerical magnitude processing tasks, but only for stimuli with set sizes that exceed the subitizing range.     

Demonstration of mismatch negativity in the monkey.

In humans, deviant auditory stimuli elicit an event-related potential (ERP) component, termed "mismatch negativity" (MMN), that reflects the operation of a cortical detector of infrequent stimulus change. Epidural auditory ERPs were recorded from 3 cynomolgous monkeys in response to soft and loud clicks. "Oddball" loud or soft stimuli elicited a long-duration frontocentral negativity, peaking at approximately 85 msec, that was superimposed upon cortically generated obligatory ERP components. These data suggest that monkeys might serve as a heuristically valuable system in which to study the neurochemical and neuroanatomical substrates of early context-dependent ERP generation.     

Demonstration of mismatch negativity in the monkey

In humans, deviant auditory stimuli elicit an event-related potential (ERP) component, termed “mismatch negativity” (MMN), that reflects the operation of a cortical detector of infrequent stimulus change. Epidural auditory ERPs were recorded from 3 cynomolgous monkeys in response to soft and loud clicks. “Oddball” loud or soft stimuli elicited a long-duration frontocentral negativity, peaking at approximately 85 msec, that was superimposed upon cortically generated obligatory ERP components. These data suggest that monkeys might serve as a heuristically valuable system in which to study the neurochemical and neuroanatomical substrates of early context-dependent ERP generation.     

The orienting of visuospatial attention: an event-related brain potential study

This study investigated the electrophysiological correlates of shifting, maintaining, and relaxing the focus of attention, using a symbolic cuing task. Cues and imperative stimuli were presented in rapid succession, and the ADJAR procedure was used to remove the contribution of event-related potential (ERP) activity associated with the imperative stimulus from the cue-related ERP waveforms. Initial analyses, comparing left and right attention-directing cues, replicated previous findings of early directing attention negativity (EDAN) and anterior directing attention negativity (ADAN) effects. To isolate ERP activity that is common to leftward and rightward attention shifts, the combined ERP activity elicited by attention-directing cues was compared to the ERP activity elicited by non-informative cues. This analysis revealed a strong and broadly distributed early positivity followed by a sustained central negativity, possibly reflecting the controlled orienting and subsequent maintenance of attentional focus. Finally, imperative stimuli preceded by non-informative cues were characterized by an enhanced posterior P2 effect, with a scalp distribution indicative of generators in visual areas. This result suggests a relatively late (re)activation in visual areas associated with the processing of stimuli that had not been cued in advance.     

Auditory cortical tuning to statistical regularities in phonology

OBJECTIVE: Ample behavioral evidence suggests that distributional properties of the language environment influence the processing of speech. Yet, how these characteristics are reflected in neural processes remains largely unknown. The present ERP study investigates neurophysiological correlates of phonotactic probability: the distributional frequency of phoneme combinations. METHODS: We employed an ERP measure indicative of experience-dependent auditory memory traces, the mismatch negativity (MMN). We presented pairs of non-words that differed by the degree of phonotactic probability in a modified passive oddball design that minimizes the contribution of acoustic processes. RESULTS: In Experiment 1 the non-word with high phonotactic probability (notsel) elicited a significantly enhanced MMN as compared to the non-word with low phonotactic probability (notkel). In Experiment 2 this finding was replicated with a non-word pair with a smaller acoustic difference (notsel-notfel). An MMN enhancement was not observed in a third acoustic control experiment with stimuli having comparable phonotactic probability (so-fo). CONCLUSIONS: Our data suggest that auditory cortical responses to phoneme clusters are modulated by statistical regularities of phoneme combinations. SIGNIFICANCE: This study indicates that the language environment is relevant in shaping the neural processing of speech. Furthermore, it provides a potentially useful design for investigating implicit phonological processing in children with anomalous language functions like dyslexia.     

Online processing of moral transgressions: ERP evidence for spontaneous evaluation

Experimental studies using fictional moral dilemmas indicate that both automatic emotional processes and controlled cognitive processes contribute to moral judgments. However, not much is known about how people process socio-normative violations that are more common to their everyday life nor the time-course of these processes. Thus, we recorded participants’ electrical brain activity while they were reading vignettes that either contained morally acceptable vs unacceptable information or text materials that contained information which was either consistent or inconsistent with their general world knowledge. A first event-related brain potential (ERP) positivity peaking at ∼200 ms after critical word onset (P200) was larger when this word involved a socio-normative or knowledge-based violation. Subsequently, knowledge-inconsistent words triggered a larger centroparietal ERP negativity at ∼320 ms (N400), indicating an influence on meaning construction. In contrast, a larger ERP positivity (larger late positivity), which also started at ∼320 ms after critical word onset, was elicited by morally unacceptable compared with acceptable words. We take this ERP positivity to reflect an implicit evaluative (good–bad) categorization process that is engaged during the online processing of moral transgressions.     

The location of preceding stimuli affects selective processing in a sustained attention situation

Event-related potentials (ERPs) were recorded to visual and auditory stimuli in a situation where subjects were required to attend selectively to the left or right side for an entire experimental block and to detect occasional target stimuli at attended locations. Stimuli were presented randomly at attended and unattended locations. In exp. 1, visual and auditory stimuli were presented in separate blocks, while in exp. 2, they were presented together and subjects had to detect visual targets at attended locations. Stimuli at attended positions elicited enlarged sensory-evoked potentials and an enhanced negativity at midline electrodes as compared with unattended stimuli. The latter effect was, however, modulated by the location of the preceding stimulus. At frontocentral electrodes, it was larger for stimuli that were preceded by stimuli at the contralateral side as compared with stimuli preceded by stimuli at the same location. It is argued that this effect may be due to a different amount of processing required for the preceding stimulus. When the predecessor is at a to-be-attended location, it has to be processed more intensively which may interfere with the processing of the next stimulus.     

Neural correlates of perceptual priming of visual motion

In two experiments, the temporal dynamics of neural activity underlying perceptual priming of visual motion was examined using event-related potentials (ERPs) during directional judgments of the apparent motion of two-dimensional sine-wave gratings. Compared to perceptually ambiguous motion, unambiguous left- or rightward motion was associated with enhanced ERP activity about 300 ms after the onset of apparent motion. In the second experiment, ERPs were recorded to two successive motion jumps in which an unambiguous motion jump served as a prime for a subsequent target motion that was ambiguous. The prime-target time interval was varied between 200, 400, and 1000 ms. In a control (motion reversal) condition, the two motion jumps were both unambiguous but in opposite directions. Compared to the motion reversal condition, motion priming was associated with an enhancement of ERP amplitudes at 100 ms and 350 ms following target stimulus onset. ERP enhancement was greatest at a short prime-target interval of 200 ms, which was also associated behaviorally with the strongest priming. The ERP enhancement and behavioral priming were both eliminated at the long 1000 ms prime-target interval. Functional magnetic resonance imaging (fMRI) data from a subset of subjects supported the view that motion priming involves modulation of neural responses both in early visual cortex and in later stages of visual processing.     

Visual temporal window of integration as revealed by the visual mismatch negativity event-related potential to stimulus omissions

We studied whether, similarly to the auditory modality, short-period temporal integration processes occur in vision. Event-related potentials (ERP) were recorded for occasional stimulus omissions from sequences of patterned visual stimuli. A posterior negative component emerged only when the constant stimulus onset asynchrony (SOA) was shorter than 150 ms. This upper limit is comparable with the duration of the temporal window of integration observed in the auditory modality (including experiments studying the effects of stimulus omissions). Parameters of the posterior negativity were highly similar irrespective of whether the stimuli were task-relevant or not (Experiment 1). Thus, we identified this potential as the visual mismatch negativity (vMMN) component, which reflects task-independent detection of violating regularities of the stimulation. vMMN was followed by an anterior positivity (the P3a), indicating attentional shifts induced by the stimulus omissions. In Experiment 2, a posterior negativity similar to that observed in Experiment 1 emerged after the termination of short trains of stimuli, again only when the SOA was shorter than 150 ms. These results support the notion of a temporal integration window in the visual modality, the duration of which is between 150 and 180 ms.     

Electric brain responses to inappropriate harmonies during listening to expressive music.

OBJECTIVES: Recent studies with event-related brain potentials (ERPs) investigating music processing found (early) negativities with right-hemispheric predominance as a response to inappropriate harmonies within sequences of chords. The stimuli used in those studies were fairly artificial in order to control the experimental factors (e.g. variations in tempo and loudness were eliminated). This raises the question of whether these ERPs can also be elicited during listening to more naturalistic stimuli.METHODS: Excerpts from classical piano sonatas were taken from commercial CDs and presented to the participants while recording the continuous electroencephalogram. Expected chords and unexpected (transposed) chords were presented at the end of chord-sequences.RESULTS: Unexpected chords elicited a negativity which was maximal around 250 ms, visible over both hemispheres, and preponderant over right temporal leads.CONCLUSIONS: The found negativity is strongly reminiscent to both early right anterior negativity and right anterior-temporal negativity, suggesting that cognitive processes underlying these ERP components are not only elicited with fairly artificial experimental stimuli but also when listening to expressive music.     

His or mine? The time course of self-other discrimination in emotion processing

This electroencephalography (EEG) study investigated at which temporal processing stages self-other discrimination in emotion processing occurs. EEG was recorded in 23 healthy participants during silent reading of unpleasant, pleasant, and neutral pronoun-noun and article--noun expressions that were related to the participants themselves, related to an unknown third person, or had no self-other reference at all. Self- and other-related pronoun--noun pairs elicited larger cortical negativity relative to the processing of article--noun pairs at left posterior electrodes as early as 200 ms after stimulus onset. In the same time windows (from 200 ms to 300 ms and 300 ms to 400 ms) the emotionality of the words enhanced event-related brain potential (ERP) amplitudes at parieto-occipital electrodes. From 350 ms onwards, processing of self-related unpleasant words elicited larger frontal negativity compared to unpleasant words that were related to the other or that had no reference at all. In addition, processing of pleasant words vs. neutral or unpleasant words elicited larger positive amplitudes over parietal electrodes from 450 ms after stimulus onset, in particular when words were self-related. Our findings demonstrate that for verbal emotional stimuli, self--other discrimination first occurs at higher-order, cortical processing stages. This is consistent with the view that categorization of information according to certain stimulus aspects (self--other reference, emotionality) occurs before its meaning is integrated.     

Extracting rules: early and late mismatch negativity to tone patterns

The auditory processing of physical stimulus features can be measured by the mismatch negativity. Past studies have shown that higher-order stimulus features also elicit a mismatch negativity. In some studies, a second component, termed late mismatch negativity, has been observed; yet the functional significance of this component remains unclear. We tested two-tone-pattern stimuli following an abstract rule in healthy adults. As expected, the tone pattern elicited a significant mismatch negativity peaking at 146 ms but a significant late mismatch negativity at around 340 ms was also observed. These findings show that the violation of an abstract rule elicits an early and late mismatch negativity. The late mismatch negativity might be triggered on the basis of auditory rule extraction processes and reflect a transfer of rules to the long-term memory.     

His or mine? The time course of self–other discrimination in emotion processing

This electroencephalography (EEG) study investigated at which temporal processing stages self–other discrimination in emotion processing occurs. EEG was recorded in 23 healthy participants during silent reading of unpleasant, pleasant, and neutral pronoun–noun and article–noun expressions that were related to the participants themselves, related to an unknown third person, or had no self–other reference at all. Self- and other-related pronoun–noun pairs elicited larger cortical negativity relative to the processing of article–noun pairs at left posterior electrodes as early as 200 ms after stimulus onset. In the same time windows (from 200 ms to 300 ms and 300 ms to 400 ms) the emotionality of the words enhanced event-related brain potential (ERP) amplitudes at parieto-occipital electrodes. From 350 ms onwards, processing of self-related unpleasant words elicited larger frontal negativity compared to unpleasant words that were related to the other or that had no reference at all. In addition, processing of pleasant words vs. neutral or unpleasant words elicited larger positive amplitudes over parietal electrodes from 450 ms after stimulus onset, in particular when words were self-related. Our findings demonstrate that for verbal emotional stimuli, self–other discrimination first occurs at higher-order, cortical processing stages. This is consistent with the view that categorization of information according to certain stimulus aspects (self–other reference, emotionality) occurs before its meaning is integrated.