Spatiotemporal Dynamics of Attention Networks Revealed by Representational Similarity Analysis of EEG and fMRI.

The fronto-parietal attention networks have been extensively studied with functional magnetic resonance imaging (fMRI), but spatiotemporal dynamics of these networks are not well understood. We measured event-related potentials (ERPs) with electroencephalography (EEG) and collected fMRI data from identical experiments where participants performed visual and auditory discrimination tasks separately or simultaneously and with or without distractors. To overcome the low temporal resolution of fMRI, we used a novel ERP-based application of multivariate representational similarity analysis (RSA) to parse time-averaged fMRI pattern activity into distinct spatial maps that each corresponded, in representational structure, to a short temporal ERP segment. Discriminant analysis of ERP-fMRI correlations revealed 8 cortical networks-2 sensory, 3 attention, and 3 other-segregated by 4 orthogonal, temporally multifaceted and spatially distributed functions. We interpret these functions as 4 spatiotemporal components of attention: modality-dependent and stimulus-driven orienting, top-down control, mode transition, and response preparation, selection and execution.

Gaming is related to enhanced working memory performance and task-related cortical activity.

Gaming experience has been suggested to lead to performance enhancements in a wide variety of working memory tasks. Previous studies have, however, mostly focused on adult expert gamers and have not included measurements of both behavioral performance and brain activity. In the current study, 167 adolescents and young adults (aged 13-24 years) with different amounts of gaming experience performed an n-back working memory task with vowels, with the sensory modality of the vowel stream switching between audition and vision at random intervals. We studied the relationship between self-reported daily gaming activity, working memory (n-back) task performance and related brain activity measured using functional magnetic resonance imaging (fMRI). The results revealed that the extent of daily gaming activity was related to enhancements in both performance accuracy and speed during the most demanding (2-back) level of the working memory task. This improved working memory performance was accompanied by enhanced recruitment of a fronto-parietal cortical network, especially the dorsolateral prefrontal cortex. In contrast, during the less demanding (1-back) level of the task, gaming was associated with decreased activity in the same cortical regions. Our results suggest that a greater degree of daily gaming experience is associated with better working memory functioning and task difficulty-dependent modulation in fronto-parietal brain activity already in adolescence and even when non-expert gamers are studied. The direction of causality within this association cannot be inferred with certainty due to the correlational nature of the current study.

Media multitasking is associated with distractibility and increased prefrontal activity in adolescents and young adults.

The current generation of young people indulges in more media multitasking behavior (e.g., instant messaging while watching videos) in their everyday lives than older generations. Concerns have been raised about how this might affect their attentional functioning, as previous studies have indicated that extensive media multitasking in everyday life may be associated with decreased attentional control. In the current study, 149 adolescents and young adults (aged 13-24years) performed speech-listening and reading tasks that required maintaining attention in the presence of distractor stimuli in the other modality or dividing attention between two concurrent tasks. Brain activity during task performance was measured using functional magnetic resonance imaging (fMRI). We studied the relationship between self-reported daily media multitasking (MMT), task performance and brain activity during task performance. The results showed that in the presence of distractor stimuli, a higher MMT score was associated with worse performance and increased brain activity in right prefrontal regions. The level of performance during divided attention did not depend on MMT. This suggests that daily media multitasking is associated with behavioral distractibility and increased recruitment of brain areas involved in attentional and inhibitory control, and that media multitasking in everyday life does not translate to performance benefits in multitasking in laboratory settings.

Behavioral and electrophysiological indicators of auditory distractibility in children with ADHD and comorbid ODD.

Involuntary switching of attention to distracting sounds was studied by measuring effects of these events on auditory discrimination performance and event-related brain potentials (ERPs) in 6-11-year-old boys with Attention Deficit-Hyperactivity Disorder (ADHD) and comorbid Oppositional Defiant Disorder (ODD) and in age-matched controls. The children were instructed to differentiate between two animal calls by pressing one response button, for example, to a dog bark and another button to a cat mew. These task-relevant sounds were presented from one of two loudspeakers in front of the child, and there were occasional task-irrelevant changes in the sound location, that is, the loudspeaker. In addition, novel sounds (e.g., a sound of hammer, rain, or car horn) unrelated to the task were presented from a loudspeaker behind the child. The percentage of correct responses was lower for target sounds preceded by a novel sound than for targets not preceded by such sound in the ADHD group, but not in the control group. In both groups, a biphasic positive P3a response was observed in ERPs to the novel sounds. The later part of the P3a appeared to continue longer over the frontal scalp areas in the ADHD group than in the controls presumably because a reorienting negativity (RON) ERP response following the P3a was smaller in the ADHD group than in the control group. This suggests that the children with ADHD had problems in reorienting their attention to the current task after a distracting novel sound leading to deterioration of performance in this task. The present study also indicates that children with ADHD and comorbid ODD show same kind of distractibility as found in previous studies for children with ADHD without systematic comorbid ODD.

Working memory resources are shared across sensory modalities.

A common assumption in the working memory literature is that the visual and auditory modalities have separate and independent memory stores. Recent evidence on visual working memory has suggested that resources are shared between representations, and that the precision of representations sets the limit for memory performance. We tested whether memory resources are also shared across sensory modalities. Memory precision for two visual (spatial frequency and orientation) and two auditory (pitch and tone duration) features was measured separately for each feature and for all possible feature combinations. Thus, only the memory load was varied, from one to four features, while keeping the stimuli similar. In Experiment 1, two gratings and two tones-both containing two varying features-were presented simultaneously. In Experiment 2, two gratings and two tones-each containing only one varying feature-were presented sequentially. The memory precision (delayed discrimination threshold) for a single feature was close to the perceptual threshold. However, as the number of features to be remembered was increased, the discrimination thresholds increased more than twofold. Importantly, the decrease in memory precision did not depend on the modality of the other feature(s), or on whether the features were in the same or in separate objects. Hence, simultaneously storing one visual and one auditory feature had an effect on memory precision equal to those of simultaneously storing two visual or two auditory features. The results show that working memory is limited by the precision of the stored representations, and that working memory can be described as a resource pool that is shared across modalities.

The mismatch negativity (MMN) in basic research of central auditory processing: a review.

In the present article, the basic research using the mismatch negativity (MMN) and analogous results obtained by using the magnetoencephalography (MEG) and other brain-imaging technologies is reviewed. This response is elicited by any discriminable change in auditory stimulation but recent studies extended the notion of the MMN even to higher-order cognitive processes such as those involving grammar and semantic meaning. Moreover, MMN data also show the presence of automatic intelligent processes such as stimulus anticipation at the level of auditory cortex. In addition, the MMN enables one to establish the brain processes underlying the initiation of attention switch to, conscious perception of, sound change in an unattended stimulus stream.

Electrophysiological evidence of enhanced distractibility in ADHD children.

Abnormal involuntary attention leading to enhanced distractibility may account for different behavioral and cognitive problems in children with attention deficit hyperactivity disorder (ADHD). This was investigated in the present experiment by recording event-related brain potentials (ERPs) to distracting novel sounds during performance of a visual discrimination task. The overall performance in the visual task was less accurate in the ADHD children than in the control children, and the ADHD children had a higher number of omitted responses following novel sounds. In both groups, the distracting novel sounds elicited a biphasic P3a ERP component and a subsequent frontal Late Negativity (LN). The early phase of P3a (180-240 ms) had significantly smaller amplitudes over the fronto-central left-hemisphere recording sites in the ADHD children than in the control group presumably due to an overlapping enhanced left-hemisphere dominant negative ERP component elicited in the ADHD group. Moreover, the late phase of P3a (300-350 ms) was significantly larger over the left parietal scalp areas in the ADHD children than in the controls. The LN had a smaller amplitude and shorter latency over the frontal scalp in the ADHD group than in the controls. In conclusion, the ERP and behavioral effects caused by the novel sounds reveal deficient control of involuntary attention in ADHD children that may underlie their abnormal distractibility.

Are different kinds of acoustic features processed differently for speech and non-speech sounds?

This study examined how changes in different types of acoustic features are processed in the brain for both speech and non-speech sounds. Event-related potentials (ERPs) were recorded in native Finnish speakers presented with sequences of repetitive vowels (/e/) or complex harmonical tones interspersed with infrequent changes in duration, frequency and either a vowel change (/o/ for vowel sequences) or a double deviant (frequency+duration change for tone sequences). The stimuli were presented monaurally in separate blocks to either the left or right ear. The results showed that speech stimuli were more efficiently processed than harmonical tones as reflected by an enhanced mismatch negativity (MMN) and P3a ERP components. In addition, the duration change in vowels elicited a larger MMN component than the equivalent change in tones. This result might reflect enhanced processing of duration features in the Finnish language in which phoneme duration plays a critical role.

Brain activity index of distractibility in normal school-age children.

Children's attention is easily diverted from a current activity to a new event in the environment. This was indexed in school-age children by diminished performance speed and accuracy in a visual discrimination task caused by task-irrelevant novel sounds. Event-related brain potentials (ERPs) elicited by these distracting sounds showed a prominent positive deflection that was generated by brain processes associated with involuntary switching of attention to novel sounds. Recordings of the magnetoencephalographic (MEG) counterpart of this brain activity revealed a major bilateral generator source in the superior temporal cortex. However, ERP scalp distributions indicated also overlapping brain activity generated in other brain areas involved in involuntary attention switching. Moreover, differences in ERP amplitudes and in their correlations with the reaction times between younger (7-10 years) and older (11-13 years) children indicated developmental changes in attentional brain functions.

Electrical responses reveal the temporal dynamics of brain events during involuntary attention switching.

Surviving in the natural environment requires the rapid switching of attention among potentially relevant stimuli. We studied electrophysiologically the involuntary switching time in humans performing a task designed to study brain mechanisms of involuntary attention and distraction (C. Escera et al., 1998, J. Cogn. Neurosci., 10, 590-604). Ten subjects were instructed to discriminate visual stimuli preceded by a task-irrelevant sound, this being either a repetitive tone (P = 0.8) or a distracting sound, i.e. a slightly higher deviant tone (P = 0.1) or an environmental novel sound (P = 0.1). In different conditions, the sounds preceded the visual stimuli by 245 or 355 ms. Deviant tones and novel sounds prolonged reaction times significantly to subsequent visual stimuli by 7.4 (P < 0.02) and 15.2 ms (P < 0.003), respectively. In addition to a mismatch negativity (MMN) and a positive-polarity, 320-ms latency, P3a event-related potential associated, respectively, with detection of the distracting sound and the subsequent orienting of attention to it, a late frontal negative deflection was observed in distracting trials. The peak latency of this brain response from sound onset was 580 ms in the 245-ms condition and 115 ms longer in the 355-ms condition (P < 0.001), peaking consequently at 340 ms from visual stimulus onset, irrespective of the onset of the distracting sound. We suggest that this late frontal negative response may signal over the scalp the process of reallocating attention back to the original task after momentary distraction, and therefore that recovering from distraction may take a similar shifting time as orienting attention involuntarily towards unexpected novelty.

Cerebral mechanisms underlying orienting of attention towards auditory frequency changes.

Brain mechanisms underlying detection of auditory frequency changes were studied with event-related potentials (ERPs) in 14 human subjects discriminating visual stimuli. Scalp-current density mapping revealed bilateral components of mismatch negativity (MMN) in frontal and auditory cortices. Deviance-related activations in frontal and temporal cortex began to be significant at 94 ms and 154 ms in the right hemisphere, and at 128 ms and 132 ms in the left hemisphere. The magnitude of MMN-neuroelectric currents from the left temporal cortex correlated significantly (r = -0.56, p < 0.05) with distraction caused by MMN-eliciting deviant tones. These results suggest a complex cerebral circuitry involved in frequency change detection and strongly support the role of this circuitry in driving attention involuntarily towards potentially relevant frequency changes in the acoustic environment.

Memory traces for words as revealed by the mismatch negativity.

Brain responses to the same spoken syllable completing a Finnish word or a pseudo-word were studied. Native Finnish-speaking subjects were instructed to ignore the sound stimuli and watch a silent movie while the mismatch negativity (MMN), an automatic index of experience-dependent auditory memory traces, was recorded. The MMN to each syllable was larger when it completed a word than when it completed a pseudo-word. This enhancement, reaching its maximum amplitude at about 150 ms after the word's recognition point, did not occur in foreign subjects who did not know any Finnish. These results provide the first demonstration of the presence of memory traces for individual spoken words in the human brain. Using whole-head magnetoencephalography, the major intracranial source of this word-related MMN was found in the left superior temporal lobe.

Effects of acoustic gradient noise from functional magnetic resonance imaging on auditory processing as reflected by event-related brain potentials.

The processing of sound changes and involuntary attention to them has been widely studied with event-related brain potentials (ERPs). Recently, functional magnetic resonance imaging (fMRI) has been applied to determine the neural mechanisms of involuntary attention and the sources of the corresponding ERP components. The gradient-coil switching noise from the MRI scanner, however, is a challenge to any experimental design using auditory stimuli. In the present study, the effects of MRI noise on ERPs associated with preattentive processing of sound changes and involuntary switching of attention to them were investigated. Auditory stimuli consisted of frequently presented "standard" sounds, infrequent, slightly higher "deviant" sounds, and infrequent natural "novel" sounds. The standard and deviant sounds were either sinusoidal tones or musical chords, in separate stimulus sequences. The mismatch negativity (MMN) ERP associated with preattentive sound change detection was elicited by the deviant and novel sounds and was not affected by the prerecorded background MRI noise (in comparison with the condition with no background noise). The succeeding positive P3a ERP responses associated with involuntary attention switching elicited by novel sounds were also not affected by the MRI noise. However, in ERPs to standard tones and chords, the P1, N1, and P2 peak latencies were significantly prolonged by the MRI noise. Moreover, the amplitude of the subsequent "exogenous" N2 to the standard sounds was significantly attenuated by the presence of MRI noise. In conclusion, the present results suggest that in fMRI the background noise does not interfere with the imaging of auditory processing related to involuntary attention.

Fast vigilance decrement in closed head injury patients as reflected by the mismatch negativity (MMN).

Event-related potentials (ERPs) were measured from 24 chronic closed head injury (CHI) patients and 18 age- and education-matched controls. The oddball paradigm was applied while subjects were watching a silent movie. The standard (p=0.8) sound of 75 ms duration had a basic frequency of 500 Hz with harmonic partials of 1000 Hz and 1500 Hz, whereas these frequencies for the pitch deviant were each 10% higher. The frequencies of the duration deviant matched with those of the standard but was 25 ms in duration. The MMN (mismatch negativity), generated by the brain's automatic auditory change-detector mechanism, was elicited by both deviants. No significant differences in the MMN latency or amplitude for either pitch or duration deviants were found between the groups. However, the MMN amplitude for the pitch deviant decreased in the patient group during the experiment considerably faster than in controls, suggesting a faster vigilance decrement in the patients.

Intracranial identification of an electric frontal-cortex response to auditory stimulus change: a case study.

The aim of the present study was to clarify whether ERPs recorded directly from the human frontal cortex contributed to the auditory N1 and mismatch negativity (MMN) elicited by changes in non-phonetic and phonetic sounds. We examined the role of prefrontal cortex in the processing of stimulus repetition and change in a 6-year-old child undergoing presurgical evaluation for epilepsy. EEG was recorded from three bilateral sub-dural electrode strips located over lateral prefrontal areas during unattended auditory stimulation. EEG epochs were averaged to obtain event-related potentials (ERPs) to repeating (standard) tones and to infrequent (deviant) shorter duration tones and complex sounds (telephone buzz). In another condition, ERPs were recorded to standard and deviant syllables, /ba/ and /da/, respectively. ERPs to vibration stimuli delivered to the fingertips were not observed at any of the sub-dural electrodes, confirming modality specificity of the auditory responses. Focal auditory ERPs consisting of P100 and N150 deflections were recorded to both tones and phonemes over the right lateral prefrontal cortex. These responses were insensitive to the serial position of the repeating sound in the stimulus train. Deviant tones evoked an MMN peaking at around 128 ms. Deviant complex sounds evoked ERPs with a similar onset latency and morphology but with an approximately two-fold increase in peak-to-peak amplitude. We conclude that right lateral prefrontal cortex (Brodmann's area 45) is involved in early stages of processing repeating sounds and sound changes.

Phonological aspects of word recognition as revealed by high-resolution spatio-temporal brain mapping.

We describe, for the first time, the use of high-resolution event-related brain potentials (hrERP) to identify the spatio-temporal characteristics of neural systems involved in phonological analysis. Subjects studied a visual word/non-word that was followed by the brief presentation of a prime letter (e.g. House, M) with the instruction to anticipate the word/non-word formed by replacing the word's first letter with the prime letter. After the prime letter, an auditory target word/non-word was presented that either matched/mismatched expectations (e.g., Mouse/Barn). ERPs were recorded to the onset of the auditory targets and scalp topographical maps were derived for the phonological mismatch negativity (PMN). The PMN reflected phonological analysis and examination of the peak topography revealed that the response was characterized by a prominent frontal, right-asymmetrical distribution. Spatial de-blurring (using current source density maps) indicated that the PMN scalp topography resulted primarily from an active left anterior source. The current results provide the initial evidence for the localization of the intra-cranial generator(s) involved in phonological analysis.

Separate time behaviors of the temporal and frontal mismatch negativity sources.

It has been proposed that mismatch negativity (MMN) is generated by temporal and frontal lobe sources, the former being associated with change detection and the latter with involuntary switching of attention to sound change. If this switching of attention is triggered by the temporal cortex change-detection mechanism, one would expect that the frontal component of MMN is activated later than the temporal one. This was studied by using 64-channel electroencephalography (EEG) and 122-channel magnetoencephalography (MEG) with realistically shaped head models to determine the source current distribution in different lobes as a function of time. Minimum-norm estimation (MNE) was performed, constraining the solution to the reconstructed cortical sheet. The results support the hypothesis that the frontal MMN generator is activated later than the auditory cortex generator.

Involuntary attention and distractibility as evaluated with event-related brain potentials.

This article reviews recent event-related brain potential (ERP) studies of involuntary attention and distractibility in response to novelty and change in the acoustic environment. These studies show that the mismatch negativity, N(1) and P(3a) ERP components elicited by deviant or novel sounds in an unattended sequence of repetitive stimuli index different processes along the course to involuntary attention switch to distracting stimuli. These studies used new auditory-auditory and auditory-visual distraction paradigms, which enable one to assess objectively abnormal distractibility in several clinical patient groups, such as those suffering from closed-head injuries or chronic alcoholism.

Lateralized automatic auditory processing of phonetic versus musical information: a PET study.

Previous positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) studies show that during attentive listening, processing of phonetic information is associated with higher activity in the left auditory cortex than in the right auditory cortex while the opposite is true for musical information. The present PET study determined whether automatically activated neural mechanisms for phonetic and musical information are lateralized. To this end, subjects engaged in a visual word classification task were presented with phonetic sound sequences consisting of frequent (P = 0.8) and infrequent (P = 0.2) phonemes and with musical sound sequences consisting of frequent (P = 0.8) and infrequent (P = 0.2) chords. The phonemes and chords were matched in spectral complexity as well as in the magnitude of frequency difference between the frequent and infrequent sounds (/e/ vs. /o/; A major vs. A minor). In addition, control sequences, consisting of either frequent (/e/; A major) or infrequent sounds (/o/; A minor) were employed in separate blocks. When sound sequences consisted of intermixed frequent and infrequent sounds, automatic phonetic processing was lateralized to the left hemisphere and musical to the right hemisphere. This lateralization, however, did not occur in control blocks with one type of sound (frequent or infrequent). The data thus indicate that automatic activation of lateralized neuronal circuits requires sound comparison based on short-term sound representations.

Dysfunction of the auditory cortex persists in infants with certain cleft types.

Language and learning disabilities occur in almost half of individuals with oral clefts. The characteristics of these cognitive dysfunctions vary according to the cleft type, and the mechanisms underlying the relation between cleft type, cognitive dysfunction, and cleft-caused middle-ear disease are unknown. This study investigates preattentive auditory discrimination, which plays a significant role in language acquisition and usage, in infants with different cleft types. A mismatch negativity (MMN) component of brain evoked potentials, which indexes preconscious sound discrimination, and brain responses to rare sine-wave tones were recorded in 12 healthy infants and 32 infants with oral clefts at the ages of 0 and 6 months. Infants with clefts were subdivided into two categories: those with cleft lip and palate (CLP) (n=11 at birth, n=6 at the age of 6 months) and those with cleft palate only (CPO) (n=17 at birth, n=8 at the age of 6 months). At both ages, brain responses to rare sounds tended to be smaller in both cleft subgroups than in healthy peers. However, in the latency range of 300 to 500 ms, the MMN was significantly smaller in infants with CPO. In infants with CLP, the MMN was comparable to that of healthy infants. Differences in auditory discrimination between infants with CLP and CPO, as reflected by MMN, were detectable at birth and persisted into later infancy. This pattern parallels known behavioural differences between children with these cleft types. Brain responses to rare sounds, in contrast, had no differentiative power with respect to the cleft type.