Agency Attribution in Infancy: Evidence for a Negativity Bias

Adults tend to attribute agency and intention to the causes of negative outcomes, even if those causes are obviously mechanical. Is this over-attribution of negative agency the result of years of practice with attributing agency to actual conspecifics, or is it a foundational aspect of our agency-detection system, present in the first year of life? Here we present two experiments with 6-month-old infants, in which they attribute agency to a mechanical claw that causes a bad outcome, but not to a claw that causes a good outcome. Control experiments suggest that the attribution stems directly from the negativity of the outcome, rather than from physical cues present in the stimuli. Together, these results provide evidence for striking developmental continuity in the attribution of agency to the causes of negative outcomes.     

An Event-Related fMRI Study of Phonological Verbal Working Memory in Schizophrenia

Background

While much is known about the role of prefrontal cortex (PFC) in working memory (WM) deficits of schizophrenia, the nature of the relationship between cognitive components of WM and brain activation patterns remains unclear. We aimed to elucidate the neural correlates of the maintenance component of verbal WM by examining correct and error trials with event-related fMRI.

Methodology/Findings

Twelve schizophrenia patients (SZ) and thirteen healthy control participants (CO) performed a phonological delayed-matching-to-sample-task in which a memory set of three nonsense words was presented, followed by a 6-seconds delay after which a probe nonsense word appeared. Participants decided whether the probe matched one of the targets, and rated the confidence of their decision. Blood-oxygen-level-dependent (BOLD) activity during WM maintenance was analyzed in relation to performance (correct/error) and confidence ratings. Frontal and parietal regions exhibited increased activation on correct trials for both groups. Correct and error trials were further segregated into true memory, false memory, guess, and true error trials. True memory trials were associated with increased bilateral activation of frontal and parietal regions in both groups but only CO showed deactivation in PFC. There was very little maintenance-related cortical activity during guess trials. False memory was associated with increased left frontal and parietal activation in both groups.

Conclusion

These findings suggest that a wider network of frontal and parietal regions support WM maintenance in correct trials compared with error trials in both groups. Furthermore, a more extensive and dynamic pattern of recruitment of the frontal and parietal networks for true memory was observed in healthy controls compared with schizophrenia patients. These results underscore the value of parsing the sources of memory errors in fMRI studies because of the non-linear nature of the brain-behavior relationship, and suggest that group comparisons need to be interpreted in more specific behavioral contexts.     

The Temporal Lobes Differentiate between the Voices of Famous and Unknown People: An Event-Related fMRI Study on Speaker Recognition

It is widely accepted that the perception of human voices is supported by neural structures located along the superior temporal sulci. However, there is an ongoing discussion to what extent the activations found in fMRI studies are evoked by the vocal features themselves or are the result of phonetic processing. To show that the temporal lobes are indeed engaged in voice processing, short utterances spoken by famous and unknown people were presented to healthy young participants whose task it was to identify the familiar speakers. In two event-related fMRI experiments, the temporal lobes were found to differentiate between familiar and unfamiliar voices such that named voices elicited higher BOLD signal intensities than unfamiliar voices. Yet, the temporal cortices did not only discriminate between familiar and unfamiliar voices. Experiment 2, which required overtly spoken responses and allowed to distinguish between four familiarity grades, revealed that there was a fine-grained differentiation between all of these familiarity levels with higher familiarity being associated with larger BOLD signal amplitudes. Finally, we observed a gradual response change such that the BOLD signal differences between unfamiliar and highly familiar voices increased with the distance of an area from the transverse temporal gyri, especially towards the anterior temporal cortex and the middle temporal gyri. Therefore, the results suggest that (the anterior and non-superior portions of) the temporal lobes participate in voice-specific processing independent from phonetic components also involved in spoken speech material.     

The Attention Modulation on Timing: An Event-Related Potential Study

The present study examined the neural mechanisms of attention modulation on timing using ERP and sLORETA measurements in a dual-task paradigm. We parametrically varied the attention to the durations of a 1000-Hz pure tone and further localized the cortical regions that were sensitive to the attention modulation on timing. Results demonstrated that the attention modulation might happen at early stage, approximately 200 ms after stimulus presentation. The P2 component at frontal area served as an early neural correlate of attention effects on timing. More importantly, the contingent negative variation (CNV) appeared at fronto-central area was sensitive to the attention effect. In addition, the supplementary motor area (SMA) was assumed to be one of the key regions for selectively attending to and estimating time. These findings provide temporal and spatial correlates of attention-modulated time processing and potentially help to investigate the neural mechanisms of patients with time perception deficits.     

Effects of Sound Frequency on Audiovisual Integration: An Event-Related Potential Study

A combination of signals across modalities can facilitate sensory perception. The audiovisual facilitative effect strongly depends on the features of the stimulus. Here, we investigated how sound frequency, which is one of basic features of an auditory signal, modulates audiovisual integration. In this study, the task of the participant was to respond to a visual target stimulus by pressing a key while ignoring auditory stimuli, comprising of tones of different frequencies (0.5, 1, 2.5 and 5 kHz). A significant facilitation of reaction times was obtained following audiovisual stimulation, irrespective of whether the task-irrelevant sounds were low or high frequency. Using event-related potential (ERP), audiovisual integration was found over the occipital area for 0.5 kHz auditory stimuli from 190–210 ms, for 1 kHz stimuli from 170–200 ms, for 2.5 kHz stimuli from 140–200 ms, 5 kHz stimuli from 100–200 ms. These findings suggest that a higher frequency sound signal paired with visual stimuli might be early processed or integrated despite the auditory stimuli being task-irrelevant information. Furthermore, audiovisual integration in late latency (300–340 ms) ERPs with fronto-central topography was found for auditory stimuli of lower frequencies (0.5, 1 and 2.5 kHz). Our results confirmed that audiovisual integration is affected by the frequency of an auditory stimulus. Taken together, the neurophysiological results provide unique insight into how the brain processes a multisensory visual signal and auditory stimuli of different frequencies.     

Perceptual Learning of Motion Direction Discrimination with Suppressed and Unsuppressed MT in Humans: An fMRI Study

The middle temporal area of the extrastriate visual cortex (area MT) is integral to motion perception and is thought to play a key role in the perceptual learning of motion tasks. We have previously found, however, that perceptual learning of a motion discrimination task is possible even when the training stimulus contains locally balanced, motion opponent signals that putatively suppress the response of MT. Assuming at least partial suppression of MT, possible explanations for this learning are that 1) training made MT more responsive by reducing motion opponency, 2) MT remained suppressed and alternative visual areas such as V1 enabled learning and/or 3) suppression of MT increased with training, possibly to reduce noise. Here we used fMRI to test these possibilities. We first confirmed that the motion opponent stimulus did indeed suppress the BOLD response within hMT+ compared to an almost identical stimulus without locally balanced motion signals. We then trained participants on motion opponent or non-opponent stimuli. Training with the motion opponent stimulus reduced the BOLD response within hMT+ and greater reductions in BOLD response were correlated with greater amounts of learning. The opposite relationship between BOLD and behaviour was found at V1 for the group trained on the motion-opponent stimulus and at both V1 and hMT+ for the group trained on the non-opponent motion stimulus. As the average response of many cells within MT to motion opponent stimuli is the same as their response to non-directional flickering noise, the reduced activation of hMT+ after training may reflect noise reduction.     

Sex Differences and Emotion Regulation: An Event-Related Potential Study

Difficulties in emotion regulation have been implicated as a potential mechanism underlying anxiety and mood disorders. It is possible that sex differences in emotion regulation may contribute towards the heightened female prevalence for these disorders. Previous fMRI studies of sex differences in emotion regulation have shown mixed results, possibly due to difficulties in discriminating the component processes of early emotional reactivity and emotion regulation. The present study used event-related potentials (ERPs) to examine sex differences in N1 and N2 components (reflecting early emotional reactivity) and P3 and LPP components (reflecting emotion regulation). N1, N2, P3, and LPP were recorded from 20 men and 23 women who were instructed to “increase,” “decrease,” and “maintain” their emotional response during passive viewing of negative images. Results indicated that women had significantly greater N1 and N2 amplitudes (reflecting early emotional reactivity) to negative stimuli than men, supporting a female negativity bias. LPP amplitudes increased to the “increase” instruction, and women displayed greater LPP amplitudes than men to the “increase” instruction. There were no differences to the “decrease” instruction in women or men. These findings confirm predictions of the female negativity bias hypothesis and suggest that women have greater up-regulation of emotional responses to negative stimuli. This finding is highly significant in light of the female vulnerability for developing anxiety disorders.     

Cortical Response Variation with Different Sound Pressure Levels: A Combined Event-Related Potentials and fMRI Study

Introduction

Simultaneous recording of electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) provides high spatial and temporal resolution. In this study we combined EEG and fMRI to investigate the structures involved in the processing of different sound pressure levels (SPLs).

Methods

EEG data were recorded simultaneously with fMRI from 16 healthy volunteers using MR compatible devices at 3 T. Tones with different SPLs were delivered to the volunteers and the N1/P2 amplitudes were included as covariates in the fMRI data analysis in order to compare the structures activated with high and low SPLs. Analysis of variance (ANOVA) and ROI analysis were also performed. Additionally, source localisation analysis was performed on the EEG data.

Results

The integration of averaged ERP parameters into the fMRI analysis showed an extended map of areas exhibiting covariation with the BOLD signal related to the auditory stimuli. The ANOVA and ROI analyses also revealed additional brain areas other than the primary auditory cortex (PAC) which were active with the auditory stimulation at different SPLs. The source localisation analyses showed additional sources apart from the PAC which were active with the high SPLs.

Discussion

The PAC and the insula play an important role in the processing of different SPLs. In the fMRI analysis, additional activation was found in the anterior cingulate cortex, opercular and orbito-frontal cortices with high SPLs. A strong response of the visual cortex was also found with the high SPLs, suggesting the presence of cross-modal effects.     

The Neural Circuitry of Reward Processing in Complex Social Comparison: Evidence from an Event-Related fMRI Study

In this study, Functional magnetic resonance imaging (fMRI) was conducted to investigate the mechanisms by which the brain activity in a complex social comparison context. One true subject and two pseudo-subjects were asked to complete a simple number estimate task at the same time which including upward and downward comparisons. Two categories of social comparison rewards (fair and unfair rewards distributions) were mainly presented by comparing the true subject with other two pseudo-subjects. Particularly, there were five conditions of unfair distribution when all the three subjects were correct but received different rewards. Behavioral data indicated that the ability to self-regulate was important in satisfaction judgment when the subject perceived an unfair reward distribution. fMRI data indicated that the interaction between the ventral striatum and the prefrontal cortex was important in self-regulation under specific conditions in complex social comparison, especially under condition of reward processing when there were two different reward values and the subject failed to exhibit upward comparison.     

An Event-Related Potential Study on the Effects of Cannabis on Emotion Processing

The effect of cannabis on emotional processing was investigated using event-related potential paradigms (ERPs). ERPs associated with emotional processing of cannabis users, and non-using controls, were recorded and compared during an implicit and explicit emotional expression recognition and empathy task. Comparisons in P3 component mean amplitudes were made between cannabis users and controls. Results showed a significant decrease in the P3 amplitude in cannabis users compared to controls. Specifically, cannabis users showed reduced P3 amplitudes for implicit compared to explicit processing over centro-parietal sites which reversed, and was enhanced, at fronto-central sites. Cannabis users also showed a decreased P3 to happy faces, with an increase to angry faces, compared to controls. These effects appear to increase with those participants that self-reported the highest levels of cannabis consumption. Those cannabis users with the greatest consumption rates showed the largest P3 deficits for explicit processing and negative emotions. These data suggest that there is a complex relationship between cannabis consumption and emotion processing that appears to be modulated by attention.     

Temporal Characteristics of Online Syntactic Sentence Planning: An Event-Related Potential Study

During sentence production, linguistic information (semantics, syntax, phonology) of words is retrieved and assembled into a meaningful utterance. There is still debate on how we assemble single words into more complex syntactic structures such as noun phrases or sentences. In the present study, event-related potentials (ERPs) were used to investigate the time course of syntactic planning. Thirty-three volunteers described visually animated scenes using naming formats varying in syntactic complexity: from simple words (‘W’, e.g., “triangle”, “red”, “square”, “green”, “to fly towards”), to noun phrases (‘NP’, e.g., “the red triangle”, “the green square”, “to fly towards”), to a sentence (‘S’, e.g., “The red triangle flies towards the green square.”). Behaviourally, we observed an increase in errors and corrections with increasing syntactic complexity, indicating a successful experimental manipulation. In the ERPs following scene onset, syntactic complexity variations were found in a P300-like component (‘S’/‘NP’>‘W’) and a fronto-central negativity (linear increase with syntactic complexity). In addition, the scene could display two actions - unpredictable for the participant, as the disambiguation occurred only later in the animation. Time-locked to the moment of visual disambiguation of the action and thus the verb, we observed another P300 component (‘S’>‘NP’/‘W’). The data show for the first time evidence of sensitivity to syntactic planning within the P300 time window, time-locked to visual events critical of syntactic planning. We discuss the findings in the light of current syntactic planning views.     

Individual Differences in the Recognition of Facial Expressions: An Event-Related Potentials Study

Previous studies have shown that early posterior components of event-related potentials (ERPs) are modulated by facial expressions. The goal of the current study was to investigate individual differences in the recognition of facial expressions by examining the relationship between ERP components and the discrimination of facial expressions. Pictures of 3 facial expressions (angry, happy, and neutral) were presented to 36 young adults during ERP recording. Participants were asked to respond with a button press as soon as they recognized the expression depicted. A multiple regression analysis, where ERP components were set as predictor variables, assessed hits and reaction times in response to the facial expressions as dependent variables. The N170 amplitudes significantly predicted for accuracy of angry and happy expressions, and the N170 latencies were predictive for accuracy of neutral expressions. The P2 amplitudes significantly predicted reaction time. The P2 latencies significantly predicted reaction times only for neutral faces. These results suggest that individual differences in the recognition of facial expressions emerge from early components in visual processing.     

Emotion-Motion Interactions in Conversion Disorder: An fMRI Study

Objectives

To evaluate the neural correlates of implicit processing of negative emotions in motor conversion disorder (CD) patients.

Methods

An event related fMRI task was completed by 12 motor CD patients and 14 matched healthy controls using standardised stimuli of faces with fearful and sad emotional expressions in comparison to faces with neutral expressions. Temporal changes in the sensitivity to stimuli were also modelled and tested in the two groups.

Results

We found increased amygdala activation to negative emotions in CD compared to healthy controls in region of interest analyses, which persisted over time consistent with previous findings using emotional paradigms. Furthermore during whole brain analyses we found significantly increased activation in CD patients in areas involved in the ‘freeze response’ to fear (periaqueductal grey matter), and areas involved in self-awareness and motor control (cingulate gyrus and supplementary motor area).

Conclusions

In contrast to healthy controls, CD patients exhibited increased response amplitude to fearful stimuli over time, suggesting abnormal emotional regulation (failure of habituation / sensitization). Patients with CD also activated midbrain and frontal structures that could reflect an abnormal behavioral-motor response to negative including threatening stimuli. This suggests a mechanism linking emotions to motor dysfunction in CD.     

Attentional Bias in Snus Users: An Experimental Study

The use of nicotine in the form of “snus” is substantial and increasing in some geographic areas, in particular among young people. It has previously been suggested that addictions may operate through a mechanism of attentional bias, in which stimuli representative of the dependent substance increase in salience, thus increasing the addictive behavior. However, this hypothesis has not been tested for the case of snus. The current experiment used a modified Stroop task and a dot-probe task to investigate whether 40 snus users show an attentional bias towards snus-relevant stimuli, compared to 40 non-snus users. There were no significant differences between the two groups on reaction times or accuracy on either Stroop or dot-probe task, thus failing to show an attentional bias towards snus-relevant stimuli for snus users. This could imply that other mechanisms may contribute to maintenance of snus use than for other addictions. However, this is the first experimental study investigating attentional bias in snus users, and more research is warranted.     

Effects of Auditory Stimuli in the Horizontal Plane on Audiovisual Integration: An Event-Related Potential Study

This article aims to investigate whether auditory stimuli in the horizontal plane, particularly originating from behind the participant, affect audiovisual integration by using behavioral and event-related potential (ERP) measurements. In this study, visual stimuli were presented directly in front of the participants, auditory stimuli were presented at one location in an equidistant horizontal plane at the front (0°, the fixation point), right (90°), back (180°), or left (270°) of the participants, and audiovisual stimuli that include both visual stimuli and auditory stimuli originating from one of the four locations were simultaneously presented. These stimuli were presented randomly with equal probability; during this time, participants were asked to attend to the visual stimulus and respond promptly only to visual target stimuli (a unimodal visual target stimulus and the visual target of the audiovisual stimulus). A significant facilitation of reaction times and hit rates was obtained following audiovisual stimulation, irrespective of whether the auditory stimuli were presented in the front or back of the participant. However, no significant interactions were found between visual stimuli and auditory stimuli from the right or left. Two main ERP components related to audiovisual integration were found: first, auditory stimuli from the front location produced an ERP reaction over the right temporal area and right occipital area at approximately 160–200 milliseconds; second, auditory stimuli from the back produced a reaction over the parietal and occipital areas at approximately 360–400 milliseconds. Our results confirmed that audiovisual integration was also elicited, even though auditory stimuli were presented behind the participant, but no integration occurred when auditory stimuli were presented in the right or left spaces, suggesting that the human brain might be particularly sensitive to information received from behind than both sides.     

Rejection in Bargaining Situations: An Event-Related Potential Study in Adolescents and Adults

The neural correlates of rejection in bargaining situations when proposing a fair or unfair offer are not yet well understood. We measured neural responses to rejection and acceptance of monetary offers with event-related potentials (ERPs) in mid-adolescents (14–17 years) and early adults (19–24 years). Participants played multiple rounds of the Ultimatum Game as proposers, dividing coins between themselves and a second player (responder) by making a choice between an unfair distribution (7 coins for proposer and 3 for responder; 7/3) and one of two alternatives: a fair distribution (5/5) or a hyperfair distribution (3/7). Participants mostly made fair offers (5/5) when the alternative was unfair (7/3), but made mostly unfair offers (7/3) when the alternative was hyperfair (3/7). When participants’ fair offers (5/5; alternative was 7/3) were rejected this was associated with a larger Medial Frontal Negativity (MFN) compared to acceptance of fair offers and rejection of unfair offers (7/3; alternative was 3/7). Also, the MFN was smaller after acceptance of unfair offers (7/3) compared to rejection. These neural responses did not differ between adults and mid-adolescents, suggesting that the MFN reacts as a neural alarm system to social prediction errors which is already prevalent during adolescence.     

Interplay of Agency and Ownership: The Intentional Binding and Rubber Hand Illusion Paradigm Combined

The sense of agency (SoA) refers to the phenomenal experience of initiating and controlling an action, whereas the sense of ownership (SoO) describes the feeling of myness an agent experiences towards his or her own body parts. SoA has been investigated with intentional binding paradigms, and the sense of ownership (SoO) with the rubber-hand illusion (RHI). We investigated the relationship between SoA and SoO by incorporating intentional binding into the RHI. Explicit and implicit measures of agency (SoA-questionnaire, intentional binding) and ownership (SoO-questionnaire, proprioceptive drift) were used. Artificial hand position (congruent/incongruent) and mode of agent (self-agent/other-agent) were systematically varied. Reported SoO varied mainly with position (higher in congruent conditions), but also with agent (higher in self-agent conditions). Reported SoA was modulated by agent (higher in self-agent conditions), and moderately by position (higher in congruent conditions). Implicit and explicit agency measures were not significantly correlated. Finally, intentional binding tended to be stronger in self-generated than observed voluntary actions. Results provide further evidence for a partial double dissociation between SoA and SoO, empirically distinct agency levels, and moderate intentional binding differences between self-generated and observed voluntary actions.     

Deficits in Error-Monitoring by College Students with Schizotypal Traits: An Event-Related Potential Study

The present study used event-related potentials (ERPs) to investigate deficits in error-monitoring by college students with schizotypal traits. Scores on the Schizotypal Personality Questionnaire (SPQ) were used to categorize the participants into schizotypal-trait (n = 17) and normal control (n = 20) groups. The error-monitoring abilities of the participants were evaluated using the Simon task, which consists of congruent (locations of stimulus and response are the same) and incongruent (locations of stimulus and response are different) conditions. The schizotypal-trait group committed more errors on the Simon task and exhibited smaller error-related negativity (ERN) amplitudes than did the control group. Additionally, ERN amplitude measured at FCz was negatively correlated with the error rate on the Simon task in the schizotypal-trait group but not in the control group. The two groups did not differ in terms of correct-related potentials (CRN), error positivity (Pe) and correct-related positivity (Pc) amplitudes. The present results indicate that individuals with schizotypal traits have deficits in error-monitoring and that reduced ERN amplitudes may represent a biological marker of schizophrenia.     

Brain Networks Responsible for Sense of Agency: An EEG Study

BackgroundSelf-agency (SA) is a person’s feeling that his action was generated by himself. The neural substrates of SA have been investigated in many neuroimaging studies, but the functional connectivity of identified regions has rarely been investigated. The goal of this study is to investigate the neural network related to SA.MethodsSA of hand movements was modulated with virtual reality. We examined the cortical network relating to SA modulation with electroencephalography (EEG) power spectrum and phase coherence of alpha, beta, and gamma frequency bands in 16 right-handed, healthy volunteers.ResultsIn the alpha band, significant relative power changes and phase coherence of alpha band were associated with SA modulation. The relative power decrease over the central, bilateral parietal, and right temporal regions (C4, Pz, P3, P4, T6) became larger as participants more effectively controlled the virtual hand movements. The phase coherence of the alpha band within frontal areas (F7-FP2, F7-Fz) was directly related to changes in SA. The functional connectivity was lower as the participants felt that they could control their virtual hand. In the other frequency bands, significant phase coherences were observed in the frontal (or central) to parietal, temporal, and occipital regions during SA modulation (Fz-O1, F3-O1, Cz-O1, C3-T4L in beta band; FP1-T6, FP1-O2, F7-T4L, F8-Cz in gamma band).ConclusionsOur study suggests that alpha band activity may be the main neural oscillation of SA, which suggests that the neural network within the anterior frontal area may be important in the generation of SA.