The role of the anterior insular cortex in self-monitoring: A novel study protocol with electrical stimulation mapping and functional magnetic resonance imaging.

Becoming aware of one's own states is a fundamental aspect for self-monitoring, allowing us to adjust our beliefs of the world to the changing context. Previous evidence points out to the key role of the anterior insular cortex (aIC) in evaluating the consequences of our own actions, especially whenever an error has occurred. In the present study, we propose a new multimodal protocol combining electrical stimulation mapping (ESM) and functional magnetic resonance imaging (fMRI) to explore the functional role of the aIC for self-monitoring in patients undergoing awake brain surgery. Our results using a modified version of the Stroop task tackling metacognitive abilities revealed new direct evidence of the involvement of the aIC in monitoring our performance, showing increased difficulties in detecting action-outcome mismatches when stimulating a cortical site located at the most posterior part of the aIC as well as significant BOLD activations at this region during outcome incongruences for self-made actions. Based on these preliminary results, we highlight the importance of assessing the aIC's functioning during tumor resection involving this region to evaluate metacognitive awareness of the self in patients undergoing awake brain surgery. In a similar vein, a better understanding of the aIC's role during self-monitoring may help shed light on action/outcome processing abnormalities reported in several neuropsychiatric disorders such as schizophrenia, anosognosia for hemiplegia or major depression.

Early response competition over the motor cortex underlies proactive control of error correction.

Response inhibition is a fundamental brain function that must be flexible enough to incorporate proactive goal-directed demands, along with reactive, automatic and well consolidated behaviors. However, whether proactive inhibitory processes can be explained by response competition, rather than by active top-down inhibitory control, remains still unclear. Using a modified version of the Eriksen flanker task, we examined the behavioral and electrophysiological correlates elicited by manipulating the degree of inhibitory control in a task that involved the fast amendment of errors. We observed that restraining or encouraging the correction of errors did not affect the behavioral and neural correlates associated to reactive inhibition. We rather found that an early, sustained and bilateral activation, of both the correct and the incorrect response, was required for an effective proactive inhibitory control. Selective unilateral patterns of response preparation were instead associated with defective response suppression. Our results provide behavioral and electrophysiological evidence of a simultaneous dual pre-activation of two motor commands, likely underlying a global operating mechanism suggesting competition or lateral inhibition to govern the amendment of errors. These findings are consistent with the response inhibitory processes already observed in speed-accuracy tradeoff studies, and hint at a decisive role of early response competition to determine the success of multiple-choice action selection.

Statistical learning and prosodic bootstrapping differentially affect neural synchronization during speech segmentation.

Neural oscillations constitute an intrinsic property of functional brain organization that facilitates the tracking of linguistic units at multiple time scales through brain-to-stimulus alignment. This ubiquitous neural principle has been shown to facilitate speech segmentation and word learning based on statistical regularities. However, there is no common agreement yet on whether speech segmentation is mediated by a transition of neural synchronization from syllable to word rate, or whether the two time scales are concurrently tracked. Furthermore, it is currently unknown whether syllable transition probability contributes to speech segmentation when lexical stress cues can be directly used to extract word forms. Using Inter-Trial Coherence (ITC) analyses in combinations with Event-Related Potentials (ERPs), we showed that speech segmentation based on both statistical regularities and lexical stress cues was accompanied by concurrent neural synchronization to syllables and words. In particular, ITC at the word rate was generally higher in structured compared to random sequences, and this effect was particularly pronounced in the flat condition. Furthermore, ITC at the syllable rate dynamically increased across the blocks of the flat condition, whereas a similar modulation was not observed in the stressed condition. Notably, in the flat condition ITC at both time scales correlated with each other, and changes in neural synchronization were accompanied by a rapid reconfiguration of the P200 and N400 components with a close relationship between ITC and ERPs. These results highlight distinct computational principles governing neural synchronization to pertinent linguistic units while segmenting speech under different listening conditions.

Food is special by itself: Neither valence, arousal, food appeal, nor caloric content modulate the attentional bias induced by food images.

When food cues appear in a visual context, such information is likely to influence eating behavior by enhancing attention for food cues. We investigated whether active but task-irrelevant information could modulate the attentional bias for food stimuli using a novel paradigm in which participants were purposely deceived by being enrolled in a memory experiment. A set of images were first held in working memory and then used as task-irrelevant distractors in a subsequent single target rapid serial visual presentation (RSVP) task, allowing us to investigate the attentional blink (AB) effect elicited by those images. In Experiment 1, the results revealed that food images elicited a larger AB effect than nonfood images. In three follow-up experiments, we investigated whether valence or arousal (Experiment 2), food preparation (Experiment 3), or food caloric content (Experiment 4) were factors related to the attentional bias for food. Overall, our results demonstrated that when held in working memory, food images can easily capture attention, even in circumstances in which the information retained in memory is irrelevant to solve the task, as indicated by the strong correlation found between items that were recognized in the RSVP task and the AB effect. Nonetheless, none of the food-related properties we examined were found to be associated with this attentional bias for food.

Hand Position and Response Assignment Modulate the Activation of the Valence-Space Conceptual Metaphor.

Conceptual metaphor is ubiquitous in language and thought, as we usually reason and talk about abstract concepts in terms of more concrete ones via metaphorical mappings that are hypothesized to arise from our embodied experience. One pervasive example is the conceptual projection of valence onto space, which flexibly recruits the vertical and lateral spatial frames to gain structure (e.g., good is up-bad is down and good is right-bad is left). In the current study, we used a valence judgment task to explore the role that exogenous bodily cues (namely response hand positions) play in the allocation of spatial attention and the modulation of conceptual congruency effects. Experiment 1 showed that congruency effects along the vertical axis are weakened when task conditions (i.e., the use of vertical visual cues, on the one hand, and the horizontal alignment of responses, on the other) draw attention to both the vertical and lateral axes making them simultaneously salient. Experiment 2 evidenced that the vertical alignment of participants' hands while responding to the task-regardless of the location of their dominant hand-facilitates the judgment of positive and negative-valence words, as long as participants respond in a metaphor-congruent manner (i.e., up responses are good and down responses are bad). Overall, these results support the claim that source domain representations are dynamically activated in response to the context and that bodily states are an integral part of that context.

Neurophysiological evidence for the interplay of speech segmentation and word-referent mapping during novel word learning.

Learning a new language requires the identification of word units from continuous speech (the speech segmentation problem) and mapping them onto conceptual representation (the word to world mapping problem). Recent behavioral studies have revealed that the statistical properties found within and across modalities can serve as cues for both processes. However, segmentation and mapping have been largely studied separately, and thus it remains unclear whether both processes can be accomplished at the same time and if they share common neurophysiological features. To address this question, we recorded EEG of 20 adult participants during both an audio alone speech segmentation task and an audiovisual word-to-picture association task. The participants were tested for both the implicit detection of online mismatches (structural auditory and visual semantic violations) as well as for the explicit recognition of words and word-to-picture associations. The ERP results from the learning phase revealed a delayed learning-related fronto-central negativity (FN400) in the audiovisual condition compared to the audio alone condition. Interestingly, while online structural auditory violations elicited clear MMN/N200 components in the audio alone condition, visual-semantic violations induced meaning-related N400 modulations in the audiovisual condition. The present results support the idea that speech segmentation and meaning mapping can take place in parallel and act in synergy to enhance novel word learning.

Headstart for speech segmentation: a neural signature for the anchor word effect.

Learning a new language is an incremental process that builds upon previously acquired information. To shed light on the mechanisms of this incremental process, we studied the on-line neurophysiological correlates of the so-called anchor word effect where newly learned words facilitate segmentation of novel words from continuous speech. Higher segmentation performance was observed for speech streams embedded with newly learned anchor words. The anchor words elicited an enhanced Stimulus-preceding negativity (SPN) component considered to be an index of expectation for incoming relevant information. Moreover, we confirmed a previously reported N400 amplitude increase for the to-be-segmented novel words, indicating a bottom-up learning process whereby new memory representations for the novel words emerge. We propose that the anchor word effect indexed by SPN reflects an expectation for an incoming novel word at the offset of the anchor word, thus facilitating the segmentation process.

The right inferior frontal cortex in response inhibition: A tDCS-ERP co-registration study.

In any given common situation, when an individual controls him/herself or obeys and stops a current action when asked to do, it is because the brain executes an inhibitory process. This ability is essential for adaptive behaviour, and it is also a requirement for accurate performance in daily life. It has been suggested that there are two main inhibitory functions related to behaviour, as inhibition is observed to affect behaviour at different time intervals. Proactive inhibition permits the subject to control his behavioural response over time by creating a response tendency, while reactive inhibition is considered to be a process that usually inhibits an already initiated response. In this context, it has been established that inhibitory function is implemented by specific fronto-basal-ganglia circuits. In the present study, we investigated the role of the right inferior frontal cortex (rIFC) in response inhibition by combining into a single task the Go-NoGo task and the Stop-Signal task. Concurrently, we applied transcranial direct current stimulation (tDCS) over the IFC and recorded electroencephalography (EEG). Thus, we obtained online EEG measurements of the tDCS-induced modifications in the IFC together with the participant's performance in a response inhibition task. We found that applying bilateral tDCS on the IFC (right anodal/left cathodal) significantly increased proactive inhibition, although the behavioural parameters indicative of reactive inhibition were unaffected by the stimulation. Finally, the inhibitory-P3 component reflected a similar modulation under both inhibitory conditions induced by the stimulation. Our data indicates that an online tDCS-ERP approach is achievable, but that a tDCS bilateral montage may not be the most efficient one for modulating the rIFC.

Neurophysiological correlates of cognitive flexibility and feedback processing in violent juvenile offenders.

The persistence of aggressive criminal behavior is recurrently observed in offenders despite being previously advised on the negative consequences of their actions. One possible explanation for the continuation of aggressive behaviors could be that they are the consequence of either possible deficits in cognitive flexibility (set-shifting) or in altered feedback processing. Event-related brain potentials (ERPs) were used to investigate both processes in non-psychopathic violent juvenile offenders. A modified version of the Wisconsin Card Sorting Test (WCST) was used to disentangle the ERP components associated with cognitive set-switching processes (P3) from feedback processing (Frontal-Related Negativity, FRN; P3). The results showed a reduction in the amplitude of the P3 component for the presentation of switch informative signals, related to set-switching processes, in the offender group. Interestingly, a larger amplitude of the P3 related to feedback processing as well as the FRN was observed in this population, probably indicating increased reliance on external feedback processing. At the behavioral level, the offender group presented a larger amount of issues with failures in implementing the new sorting rule. This behavioral pattern could be related to deficits in the ability to switch to another behavior and an altered pattern in processing the feedback information related to the precision of their performance. These observations highlight the possible role of cognitive set-switching and reward sensibility in the maintenance of harmful behaviors in juvenile offenders.

A simultaneous modulation of reactive and proactive inhibition processes by anodal tDCS on the right inferior frontal cortex.

Proactive and reactive inhibitory processes are a fundamental part of executive functions, allowing a person to stop inappropriate responses when necessary and to adjust performance in in a long term in accordance to the goals of a task. In the current study, we manipulate, in a single task, both reactive and proactive inhibition mechanisms, and we investigate the within-subjects effect of increasing, by means of anodal transcranial direct current stimulation (tDCS), the involvement of the right inferior frontal cortex (rIFC). Our results show a simultaneous enhancement of these two cognitive mechanisms when modulating the neural activity of rIFC. Thus, the application of anodal tDCS increased reaction times on Go trials, indicating a possible increase in proactive inhibition. Concurrently, the stop-signal reaction time, as a covert index of the inhibitory process, was reduced, demonstrating an improvement in reactive inhibition. In summary, the current pattern of results validates the engagement of the rIFC in these two forms of inhibitory processes, proactive and reactive inhibition and it provides evidence that both processes can operate concurrently in the brain.

Neurophysiological correlates of error monitoring and inhibitory processing in juvenile violent offenders.

Performance monitoring is crucial for well-adapted behavior. Offenders typically have a pervasive repetition of harmful-impulsive behaviors, despite an awareness of the negative consequences of their actions. However, the link between performance monitoring and aggressive behavior in juvenile offenders has not been closely investigated. Event-related brain potentials (ERPs) were used to investigate performance monitoring in juvenile non-psychopathic violent offenders compared with a well-matched control group. Two ERP components associated with error monitoring, error-related negativity (ERN) and error-positivity (Pe), and two components related to inhibitory processing, the stop-N2 and stop-P3 components, were evaluated using a combined flanker-stop-signal task. The results showed that the amplitudes of the ERN, the stop-N2, the stop-P3, and the standard P3 components were clearly reduced in the offenders group. Remarkably, no differences were observed for the Pe. At the behavioral level, slower stop-signal reaction times were identified for offenders, which indicated diminished inhibitory processing. The present results suggest that the monitoring of one's own behavior is affected in juvenile violent offenders. Specifically, we determined that different aspects of executive function were affected in the studied offenders, including error processing (reduced ERN) and response inhibition (reduced N2 and P3). However, error awareness and compensatory post-error adjustment processes (error correction) were unaffected. The current pattern of results highlights the role of performance monitoring in the acquisition and maintenance of externalizing harmful behavior that is frequently observed in juvenile offenders.

Differences in word recognition between early bilinguals and monolinguals: behavioral and ERP evidence.

We investigated the behavioral and brain responses (ERPs) of bilingual word recognition to three fundamental psycholinguistic factors, frequency, morphology, and lexicality, in early bilinguals vs. monolinguals. Earlier behavioral studies have reported larger frequency effects in bilinguals' nondominant vs. dominant language and in some studies also when compared to corresponding monolinguals. In ERPs, language processing differences between bilinguals vs. monolinguals have typically been found in the N400 component. In the present study, highly proficient Finnish-Swedish bilinguals who had acquired both languages during childhood were compared to Finnish monolinguals during a visual lexical decision task and simultaneous ERP recordings. Behaviorally, we found that the response latencies were overall longer in bilinguals than monolinguals, and that the effects for all three factors, frequency, morphology, and lexicality were also larger in bilinguals even though they had acquired both languages early and were highly proficient in them. In line with this, the N400 effects induced by frequency, morphology, and lexicality were larger for bilinguals than monolinguals. Furthermore, the ERP results also suggest that while most inflected Finnish words are decomposed into stem and suffix, only monolinguals have encountered high frequency inflected word forms often enough to develop full-form representations for them. Larger behavioral and neural effects in bilinguals in these factors likely reflect lower amount of exposure to words compared to monolinguals, as the language input of bilinguals is divided between two languages.

Brain oscillatory activity associated with task switching and feedback processing.

In this study, we sought to dissociate event-related potentials (ERPs) and the oscillatory activity associated with signals indicating feedback about performance (outcome-based behavioral adjustment) and the signals indicating the need to change or maintain a task set (rule-based behavioral adjustment). With this purpose in mind, we noninvasively recorded electroencephalographic signals, using a modified version of the Wisconsin card sorting task, in which feedback processing and task switching could be studied separately. A similar late positive component was observed for the switch and correct feedback signals on the first trials of a series, but feedback-related negativity was observed only for incorrect feedback. Moreover, whereas theta power showed a significant increase after a switch cue and after the first positive feedback of a new series, a selective frontal beta-gamma increase was observed exclusively in the first positive feedback (i.e., after the selection of the new rule). Importantly, for the switch cue, beta-alpha activity was suppressed rather than increased. This clear dissociation between the cue and feedback stimuli in task switching emphasizes the need to accurately study brain oscillatory activity to disentangle the role of different cognitive control processes.

Language learning under working memory constraints correlates with microstructural differences in the ventral language pathway.

The present study combined behavioral measures and diffusion tensor imaging to investigate the neuroanatomical basis of language learning in relation to phonological working memory (WM). Participants were exposed to simplified artificial languages under WM constraints. The results underscore the role of the rehearsal subcomponent of WM in successful speech segmentation and rule learning. Moreover, when rehearsal was blocked task performance was correlated to the white matter microstructure of the left ventral pathway connecting frontal and temporal language-related cortical areas through the extreme/external capsule. This ventral pathway may therefore play an important additional role in language learning when the main dorsal pathway-dependent rehearsal mechanisms are not available.

Words as anchors: known words facilitate statistical learning.

Can even a handful of newly learned words help to find further word candidates in a novel spoken language? This study shows that the statistical segmentation of words from speech stream by adults is facilitated by the presence of known words in the stream. This facilitatory effect is immediate as the known words were acquired only minutes before the onset of the speech stream. Our results demonstrate an interplay between top-down lexical segmentation and bottom-up statistical learning, in line with infant research suggesting that integration of multiple cues facilitates early language learning. The ability to simultaneously benefit from both types of word segmentation cues appears to be present through adulthood and can thus contribute to second language learning.

Neurophysiological markers of novelty processing are modulated by COMT and DRD4 genotypes.

Humans are faced with the dilemma to maintain a stable cognitive set on the one hand and to be able to redirect and switch attention to novel stimuli of potential importance. The dopaminergic system has been implicated in the balance between these two antagonistic constraints and in particular in novelty processing. Here we studied the impact of two polymorphisms affecting dopaminergic functioning (COMT Val108/158Met and DRD4 SNP -521) on neurophysiological correlates of novelty processing. Recording event-related potentials (ERPs) and oscillatory activity in a modified oddball task that featured infrequent but task-irrelevant novel sounds in addition to frequent standard and rare target tones, we examined participants homozygous for the Met or Val variant of COMT as well as homozygous for the C or T variant of DRD4. We found effects mainly on the P3a component to novel stimuli. A greater P3a amplitude was found for the COMT-ValVal group relative to MetMet. There was a tendency for DRD4-TT participants to show greater P3a amplitude and shorter P3a latency. Finally, DRD4-TT and COMT-ValVal participants showed the greatest increase of theta-power to novel stimuli. By contrast, the P3b component to target stimuli showed little influence of the studied polymorphism. Individual differences in dopaminergic genes explain part of the interindividual variance in the neural correlates of novelty but not target processing.

Speech segmentation is facilitated by visual cues.

Evidence from infant studies indicates that language learning can be facilitated by multimodal cues. We extended this observation to adult language learning by studying the effects of simultaneous visual cues (nonassociated object images) on speech segmentation performance. Our results indicate that segmentation of new words from a continuous speech stream is facilitated by simultaneous visual input that it is presented at or near syllables that exhibit the low transitional probability indicative of word boundaries. This indicates that temporal audio-visual contiguity helps in directing attention to word boundaries at the earliest stages of language learning. Off-boundary or arrhythmic picture sequences did not affect segmentation performance, suggesting that the language learning system can effectively disregard noninformative visual information. Detection of temporal contiguity between multimodal stimuli may be useful in both infants and second-language learners not only for facilitating speech segmentation, but also for detecting word-object relationships in natural environments.

The effects of COMT (Val108/158Met) and DRD4 (SNP -521) dopamine genotypes on brain activations related to valence and magnitude of rewards.

People's sensitivity to reinforcing stimuli such as monetary gains and losses shows a wide interindividual variation that might in part be determined by genetic differences. Because of the established role of the dopaminergic system in the neural encoding of rewards and negative events, we investigated young healthy volunteers being homozygous for either the Valine or Methionine variant of the catechol-O-methyltransferase (COMT) codon 158 polymorphism as well as homozygous for the C or T variant of the SNP -521 polymorphism of the dopamine D4 receptor. Participants took part in a gambling paradigm featuring unexpectedly high monetary gains and losses in addition to standard gains/losses of expected magnitude while undergoing functional magnetic resonance imaging at 3 T. Valence-related brain activations were seen in the ventral striatum, the anterior cingulate cortex, and the inferior parietal cortex. These activations were modulated by the COMT polymorphism with greater effects for valine/valine participants but not by the D4 receptor polymorphism. By contrast, magnitude-related effects in the anterior insula and the cingulate cortex were modulated by the D4 receptor polymorphism with larger responses for the CC variant. These findings emphasize the differential contribution of genetic variants in the dopaminergic system to various aspects of reward processing.

ADHD candidate gene (DRD4 exon III) affects inhibitory control in a healthy sample.

BACKGROUND: Dopamine is believed to be a key neurotransmitter in the development of attention-deficit/hyperactivity disorder (ADHD). Several recent studies point to an association of the dopamine D4 receptor (DRD4) gene and this condition. More specifically, the 7 repeat variant of a variable number of tandem repeats (VNTR) polymorphism in exon III of this gene is suggested to bear a higher risk for ADHD. In the present study, we investigated the role of this polymorphism in the modulation of neurophysiological correlates of response inhibition (Go/Nogo task) in a healthy, high-functioning sample. RESULTS: Homozygous 7 repeat carriers showed a tendency for more accurate behavior in the Go/Nogo task compared to homozygous 4 repeat carriers. Moreover, 7 repeat carriers presented an increased nogo-related theta band response together with a reduced go-related beta decrease. CONCLUSIONS: These data point to improved cognitive functions and prefrontal control in the 7 repeat carriers, probably due to the D4 receptor's modulatory role in prefrontal areas. The results are discussed with respect to previous behavioral data on this polymorphism and animal studies on the impact of the D4 receptor on cognitive functions.

Neurophysiological mechanisms involved in language learning in adults.

Little is known about the brain mechanisms involved in word learning during infancy and in second language acquisition and about the way these new words become stable representations that sustain language processing. In several studies we have adopted the human simulation perspective, studying the effects of brain-lesions and combining different neuroimaging techniques such as event-related potentials and functional magnetic resonance imaging in order to examine the language learning (LL) process. In the present article, we review this evidence focusing on how different brain signatures relate to (i) the extraction of words from speech, (ii) the discovery of their embedded grammatical structure, and (iii) how meaning derived from verbal contexts can inform us about the cognitive mechanisms underlying the learning process. We compile these findings and frame them into an integrative neurophysiological model that tries to delineate the major neural networks that might be involved in the initial stages of LL. Finally, we propose that LL simulations can help us to understand natural language processing and how the recovery from language disorders in infants and adults can be accomplished.