Event related potentials (ERP) reveal a robust response to visual symmetry in unattended visual regions.

Visual symmetry at fixation generates a bilateral Event Related Potential (ERP) called the Sustained Posterior Negativity (SPN). Symmetry presented in the left visual hemifield generates a contralateral SPN over the right hemisphere and vice versa. The current study examined whether the contralateral SPN is modulated by the focus of spatial attention. On each trial there were two dot patterns, one to the left of fixation, and one to the right of fixation. A central arrow cue pointed to one of the patterns and participants discriminated its regularity (symmetry or random). We compared contralateral SPN amplitude generated by symmetry at attended and unattended spatial locations. While the response to attended symmetry was slightly enhanced, the response to unattended symmetry was still substantial. Although visual symmetry detection is a computational challenge, we conclude that the brain processes visual symmetry in unattended parts of the visual field.

Lessons from a catalogue of 6674 brain recordings.

It is now possible for scientists to publicly catalogue all the data they have ever collected on one phenomenon. For a decade, we have been measuring a brain response to visual symmetry called the sustained posterior negativity (SPN). Here we report how we have made a total of 6674 individual SPNs from 2215 participants publicly available, along with data extraction and visualization tools (https://osf.io/2sncj/). We also report how re-analysis of the SPN catalogue has shed light on aspects of the scientific process, such as statistical power and publication bias, and revealed new scientific insights.

Bid outcome processing in Vickrey auctions: An ERP study.

Online retailers often sell products using a socially competitive second-price sealed-bid auction known as a Vickrey auction (VA), an incentivized demand-revealing mechanism used to elicit players' subjective values. The VA presents a situation of risky decision-making, which typically implements value processing and a loss aversion mechanism. Neural outcome processing of VA bids are not known; this study explores this for the first time using EEG. Twenty-eight healthy participants bid on household items against an anonymous, computerized opponent. Bid outcome event-related potentials were predicted to differentiate between three conditions: outbid (no-win), large margin win (bargain), and small margin win (snatch). Individual loss aversion values were evaluated in a separate behavioral experiment offering gains or losses of variable amounts but equal chances against an assured gain. Processing outcomes of VA bids were associated with a feedback-related negativity (FRN) potential with a spatial maximum at the vertex (251-271 ms), where bargain win trials resulted in greater FRN amplitudes than snatch win trials. Additionally, a P300 potential was sensitive to win versus no-win outcomes and to retail price. Individual loss aversion level did not correlate with the strength of FRN or P300. Results show that outcome processing in a VA is associated with FRN that differentiates between relatively advantageous and less advantageous gains, and a P300 that distinguishes between the more and less expensive auction items. Our findings pave the way to an objective exploration of economic decision-making and purchasing behavior involving a widely popular auction.

Electrophysiological evidence of the amodal representation of symmetry in extrastriate areas.

Extrastriate visual areas are strongly activated by image symmetry. Less is known about symmetry representation at object-level rather than image-level. Here we investigated electrophysiological responses to symmetry, generated by amodal completion of partially-occluded polygon shapes. We used a similar paradigm in four experiments (N = 112). A fully-visible abstract shape (either symmetric or asymmetric) was presented for 250 ms (t0). A large rectangle covered it entirely for 250 ms (t1) and then moved to one side to reveal one half of the shape hidden behind (t2, 1000 ms). Note that at t2 no symmetry could be extracted from retinal image information. In half of the trials the shape was the same as previously presented, in the other trials it was replaced by a novel shape. Participants matched shapes similarity (Exp. 1 and Exp. 2), or their colour (Exp. 3) or the orientation of a triangle superimposed to the shapes (Exp. 4). The fully-visible shapes (t0-t1) elicited automatic symmetry-specific ERP responses in all experiments. Importantly, there was an exposure-dependent symmetry-response to the occluded shapes that were recognised as previously seen (t2). Exp. 2 and Exp.4 confirmed this second ERP (t2) did not reflect a reinforcement of a residual carry-over response from t0. We conclude that the extrastriate symmetry-network can achieve amodal representation of symmetry from occluded objects that have been previously experienced as wholes.

Investigating the effect of losses and gains on effortful engagement during an incentivized Go/NoGo task through anticipatory cortical oscillatory changes.

Losses usually have greater subjective value (SV) than gains of equal nominal value but often cause a relative deterioration in effortful performance. Since losses and gains induce differing approach/avoidance behavioral tendencies, we explored whether incentive type interacted with approach/avoidance motor-sets. Alpha- and beta-band event-related desynchronization (ERD) was hypothesized to be weakest when participants expected a loss and prepared an inhibitory motor-set, and strongest when participants expected a gain and prepared an active motor-set. It was also hypothesized that effort would modulate reward and motor-set-related cortical activation patterns. Participants completed a cued Go/NoGo task while expecting a reward (+10p), avoiding a loss (-10p), or receiving no incentive (0p); and while expecting a NoGo cue with a probability of either .75 or .25. Pre-movement alpha- and beta-band EEG power was analyzed using the ERD method, and the SV of effort was evaluated using a cognitive effort discounting task. Gains incentivized faster RTs and stronger preparatory alpha band ERD compared to loss and no incentive conditions, while inhibitory motor-sets resulted in significantly weaker alpha-band ERD. However, there was no interaction between incentive and motor-sets. Participants were more willing to expend effort in losses compared to gain trials, although the SV of effort was not associated with ERD patterns or RTs. Results suggest that incentive and approach/avoidance motor tendencies modulate cortical activations prior to a speeded RT movement independently, and are not associated with the economic value of effort. The present results favor attentional explanations of the effect of incentive modality on effort.

Neural responses to reflection symmetry for shapes defined by binocular disparity, and for shapes perceived as regions of background.

Human perception of symmetry is associated with activation in an extended network of extrastriate visual areas. This activation generates an ERP called the Sustained Posterior Negativity (SPN). In most studies so far, the stimuli have been defined by luminance. We tested whether the SPN is present when stimuli are defined by stereoscopic disparity using random dot stereograms (RDS). In Experiment 1, we compared the SPN signal for contours specified by binocular disparity and contours specified by monocular cues. The SPN was equivalent, suggesting that the type of contour does not alter the SPN signal. In Experiment 2 we exploited the unique property of RDS to provide unambiguous figure-ground arrangements. Psychophysical work has shown that symmetry is more easily detected when it is a property of a single object (i.e., within a figure), compared to a property of a gap between two objects (i.e., the ground). Therefore, the target regions in this experiment could either be foreground or background. The SPN onset was delayed when the symmetry was in a ground region. This may be because object formation interferes with the processing of shape information in the ground region.

Spinning objects and partial occlusion: Smart neural responses to symmetry.

In humans, extrastriate visual areas are strongly activated by symmetry. However, perfect symmetry is rare in natural visual images. Recent findings showed that when parts of a symmetric shape are presented at different points in time the process relies on a perceptual memory buffer. Does this temporal integration need a retinotopic reference frame? For the first time we tested integration of parts both in the temporal and spatial domain, using a non-retinotopic frame of reference. In Experiment 1, an irregular polygonal shape (either symmetric or asymmetric) was partly occluded by a rectangle for 500 ms (T1). The rectangle moved to the opposite side to reveal the other half of the shape, whilst occluding the previously visible half (T2). The reference frame for the object was static: the two parts stimulated retinotopically corresponding receptive fields (revealed over time). A symmetry-specific ERP response from ~300 ms after T2 was observed. In Experiment 2 dynamic occlusion was combined with an additional step at T2: the new half-shape and occluder were rotated by 90°. Therefore, there was a moving frame of reference and the retinal correspondence between the two parts was disrupted. A weaker but significant symmetry-specific response was recorded. This result extends previous findings: global symmetry representation can be achieved in extrastriate areas non-retinotopically, through integration in both temporal and spatial domain.

Electrophysiological priming effects demonstrate independence and overlap of visual regularity representations in the extrastriate cortex.

An Event Related Potential (ERP) component called the Sustained Posterior Negativity (SPN) is generated by regular visual patterns (e.g. vertical reflectional symmetry, horizontal reflectional symmetry or rotational symmetry). Behavioural studies suggest symmetry becomes increasingly salient when the exemplars update rapidly. In line with this, Experiment 1 (N = 48) found that SPN amplitude increased when three different reflectional symmetry patterns were presented sequentially. We call this effect 'SPN priming'. We then exploited SPN priming to investigate independence of different symmetry representations. SPN priming did not survive changes in retinal location (Experiment 2, N = 48) or non-orthogonal changes in axis orientation (Experiment 3, N = 48). However, SPN priming transferred between vertical and horizontal axis orientations (Experiment 4, N = 48) and between reflectional and rotational symmetry (Experiment 5, N = 48). SPN priming is interesting in itself, and a useful new method for identifying functional boundaries of the symmetry response. We conclude that visual regularities at different retinal locations are coded independently. However, there is some overlap between different regularities presented at the same retinal location.

Inhibition of cortical somatosensory processing during and after low frequency peripheral nerve stimulation in humans.

OBJECTIVE: Transcutaneous low-frequency stimulation (LFS) elicits long-term depression-like effects on human pain perception. However, the neural mechanisms underlying LFS are poorly understood. We investigated cortical activation changes occurring during LFS and if changes were associated with reduced nociceptive processing and increased amplitude of spontaneous cortical oscillations post-treatment. METHODS: LFS was applied to the radial nerve of 25 healthy volunteers over two sessions using active (1 Hz) or sham (0.02 Hz) frequencies. Changes in resting electroencephalography (EEG) and laser-evoked potentials (LEPs) were investigated before and after LFS. Somatosensory-evoked potentials were recorded during LFS and source analysis was carried out. RESULTS: Ipsilateral midcingulate and operculo-insular cortex source activity declined linearly during LFS. Active LFS was associated with attenuated long-latency LEP amplitude in ipsilateral frontocentral electrodes and increased resting alpha (8-12 Hz) and beta (16-24 Hz) band power in electrodes overlying operculo-insular, sensorimotor and frontal cortical regions. Reduced ipsilateral operculo-insular cortex source activity during LFS correlated with a smaller post-treatment alpha-band power increase. CONCLUSIONS: LFS attenuated somatosensory processing both during and after stimulation. SIGNIFICANCE: Results further our understanding of the attenuation of somatosensory processing both during and after LFS.

Right lateralized alpha desynchronization increases with the proportion of symmetry in the stimulus.

Research into the neural basis of symmetry perception has intensified in the last two decades; however, the functional role of neural oscillations remains unclear. In previous work Makin et al. (2014, Journal of Vision, 14, 1-12) and Wright et al. (2015, Psychophysiology, 52, 638-647) examined occipital alpha event-related desynchronization (alpha ERD). It was concluded that alpha ERD is right lateralized during active regularity discrimination but not during a secondary task. Furthermore, alpha ERD was unaffected by stimulus properties, such as the type of regularity. These conclusions are refuted by new time-frequency analysis on an electroencephalography (EEG) data set first introduced by Makin et al. (2020, Journal of Cognitive Neuroscience, 32, 353-366). We compared alpha ERD across five tasks. First, we found that right lateralization of alpha ERD was evident in all tasks, not just active regularity discrimination. This was caused by hemispheric differences in alpha power during prestimulus baseline (left < right), which equalized after stimulus onset (left = right). Second, we found that Alpha ERD increased with the proportion of symmetric elements in the image (PSYMM). Sensitivity to PSYMM was stronger on the right. These findings suggest that known extrastriate symmetry activations are accompanied by reduced alpha power.

Source dipole analysis reveals a new brain response to visual symmetry.

Visual regularity activates a network of brain regions in the extrastriate cortex. Previous EEG studies have found that this response scales parametrically with proportion of symmetry in symmetry + noise displays. The parametric symmetry response happens in many tasks, but it is enhanced during active regularity discrimination. However, the origins and time course of this selective enhancement are unclear. Here we answered remaining questions with new source dipole analysis. As assumed, the parametric symmetry response found at the sensor level was generated by a pair of dipoles in the left and right extrastriate cortex. This bilateral activity was itself enhanced during regularity discrimination. However, we identified a third, and later, symmetry response in the posterior cingulate during regularity discrimination. Unlike the extrastriate response, this previously unknown activation only indexes strong, task relevant regularity signals. This clarifies the neural circuits which mediate the perceptual and cognitive aspects of symmetry discrimination.

Electrophysiological priming effects confirm that the extrastriate symmetry network is not gated by luminance polarity.

It is known that the extrastriate cortex is activated by visual symmetry. This activation generates an ERP component called the Sustained Posterior Negativity (SPN). SPN amplitude increases (i.e., becomes more negative) with repeated presentations. We exploited this SPN priming effect to test whether the extrastriate symmetry response is gated by element luminance polarity. On each trial, participants observed three stimuli (patterns of dots) in rapid succession (500 ms. with 200 ms. gaps). The patterns were either symmetrical or random. The dot elements were either black or white on a grey background. The triplet sequences either showed repeated luminance (black > black > black, or white > white > white) or changing luminance (black > white > black, or white > black > white). As predicted, SPN priming was comparable in repeated and changing luminance conditions. Therefore, symmetry with black elements is not processed independently from symmetry with white elements. Source waveform analysis confirmed that this priming happened within the extrastriate symmetry network. We conclude that the network pools information across luminance polarity channels.

The response to symmetry in extrastriate areas and its time course are modulated by selective attention.

Neurophysiological studies have shown a strong activation in visual areas in response to symmetry. Electrophysiological (EEG) studies, in particular, have confirmed that amplitude at posterior electrodes is more negative for symmetrical compared to asymmetrical patterns. This response is present even when observers perform tasks that do not require processing of symmetry. In this sense the activation is automatic. In this study we test this automaticity more directly by presenting stimuli that contain both symmetry and asymmetry, as overlapping patterns of dots of different colour (black and white). Observers were asked to respond to symmetry in only one of the two colours. If feature-based attention has no role the response should depend on properties of the image. If attention fully filters only the relevant colour the response should depend on properties of the relevant colour only. Neither of these models fully explained the data. We conclude that selective attention does modulate the neural response to symmetry, however we also found a significant contribution from the irrelevant pattern.

The cortical oscillatory patterns associated with varying levels of reward during an effortful vigilance task.

We explored how reward and value of effort shapes performance in a sustained vigilance, reaction time (RT) task. It was posited that reward and value would hasten RTs and increase cognitive effort by boosting activation in the sensorimotor cortex and inhibition in the frontal cortex, similar to the horse-race model of motor actions. Participants performed a series of speeded responses while expecting differing monetary rewards (0 pence (p), 1 p, and 10 p) if they responded faster than their median RT. Amplitudes of cortical alpha, beta, and theta oscillations were analysed using the event-related desynchronization method. In experiment 1 (N = 29, with 12 females), reward was consistent within block, while in experiment 2 (N = 17, with 12 females), reward amount was displayed before each trial. Each experiment evaluated the baseline amplitude of cortical oscillations differently. The value of effort was evaluated using a cognitive effort discounting task (COGED). In both experiments, RTs decreased significantly with higher rewards. Reward level sharpened the increased amplitudes of beta oscillations during fast responses in experiment 1. In experiment 2, reward decreased the amplitudes of beta oscillations in the ipsilateral sensorimotor cortex. Individual effort values did not significantly correlate with oscillatory changes in either experiment. Results suggest that reward level and response speed interacted with the task- and baseline-dependent patterns of cortical inhibition in the frontal cortex and with activation in the sensorimotor cortex during the period of motor preparation in a sustained vigilance task. However, neither the shortening of RT with increasing reward nor the value of effort correlated with oscillatory changes. This implies that amplitudes of cortical oscillations may shape upcoming motor responses but do not translate higher-order motivational factors into motor performance.

Neural underpinnings of value-guided choice during auction tasks: An eye-fixation related potentials study.

Values are attributed to goods during free viewing of objects which entails multi- and trans-saccadic cognitive processes. Using electroencephalographic eye-fixation related potentials, the present study investigated how neural signals related to value-guided choice evolved over time when viewing household and office products during an auction task. Participants completed a Becker-DeGroot-Marschak auction task whereby half of the stimuli were presented in either a free or forced bid protocol to obtain willingness-to-pay. Stimuli were assigned to three value categories of low, medium and high value based on subjective willingness-to-pay. Eye fixations were organised into five 800 ms time-bins spanning the objects total viewing time. Independent component analysis was applied to eye-fixation related potentials. One independent component (IC) was found to represent fixations for high value products with increased activation over the left parietal region of the scalp. An IC with a spatial maximum over a fronto-central region of the scalp coded the intermediate values. Finally, one IC displaying activity that extends over the right frontal scalp region responded to intermediate- and low-value items. Each of these components responded early on during viewing an object and remained active over the entire viewing period, both during free and forced bid trials. Results suggest that the subjective value of goods are encoded using sets of brain activation patterns which are tuned to respond uniquely to either low, medium, or high values. Data indicates that the right frontal region of the brain responds to low and the left frontal region to high values. Values of goods are determined at an early point in the decision making process and carried for the duration of the decision period via trans-saccadic processes.

Brain Responses to Emotional Faces in Natural Settings: A Wireless Mobile EEG Recording Study.

The detection of a human face in a visual field and correct reading of emotional expression of faces are important elements in everyday social interactions, decision making and emotional responses. Although brain correlates of face processing have been established in previous fMRI and electroencephalography (EEG)/MEG studies, little is known about how the brain representation of faces and emotional expressions of faces in freely moving humans. The present study aimed to detect brain electrical potentials that occur during the viewing of human faces in natural settings. 64-channel wireless EEG and eye-tracking data were recorded in 19 participants while they moved in a mock art gallery and stopped at times to evaluate pictures hung on the walls. Positive, negative and neutral valence pictures of objects and human faces were displayed. The time instants in which pictures first occurred in the visual field were identified in eye-tracking data and used to reconstruct the triggers in continuous EEG data after synchronizing the time axes of the EEG and eye-tracking device. EEG data showed a clear face-related event-related potential (ERP) in the latency interval ranging from 165 to 210 ms (N170); this component was not seen whilst participants were viewing non-living objects. The face ERP component was stronger during viewing disgusted compared to neutral faces. Source dipole analysis revealed an equivalent current dipole in the right fusiform gyrus (BA37) accounting for N170 potential. Our study demonstrates for the first time the possibility of recording brain responses to human faces and emotional expressions in natural settings. This finding opens new possibilities for clinical, developmental, social, forensic, or marketing research in which information about face processing is of importance.

Neural correlates of economic value and valuation context: an event-related potential study.

The value of environmental cues and internal states is continuously evaluated by the human brain, and it is this subjective value that largely guides decision making. The present study aimed to investigate the initial value attribution process, specifically the spatiotemporal activation patterns associated with values and valuation context, using electroencephalographic event-related potentials (ERPs). Participants completed a stimulus rating task in which everyday household items marketed up to a price of £4 were evaluated with respect to their desirability or material properties. The subjective values of items were evaluated as willingness to pay (WTP) in a Becker-DeGroot-Marschak auction. On the basis of the individual's subjective WTP values, the stimuli were divided into high- and low-value items. Source dipole modeling was applied to estimate the cortical sources underlying ERP components modulated by subjective values (high vs. low WTP) and the evaluation condition (value-relevant vs. value-irrelevant judgments). Low-WTP items and value-relevant judgments both led to a more pronounced N2 visual evoked potential at right frontal scalp electrodes. Source activity in right anterior insula and left orbitofrontal cortex was larger for low vs. high WTP at ∼200 ms. At a similar latency, source activity in right anterior insula and right parahippocampal gyrus was larger for value-relevant vs. value-irrelevant judgments. A stronger response for low- than high-value items in anterior insula and orbitofrontal cortex appears to reflect aversion to low-valued item acquisition, which in an auction experiment would be perceived as a relative loss. This initial low-value bias occurs automatically irrespective of the valuation context. NEW & NOTEWORTHY We demonstrate the spatiotemporal characteristics of the brain valuation process using event-related potentials and willingness to pay as a measure of subjective value. The N2 component resolves values of objects with a bias toward low-value items. The value-related changes of the N2 component are part of an automatic valuation process.

Tracking Economic Value of Products in Natural Settings: A Wireless EEG Study.

Economic decision making refers to the process of individuals translating their preference into subjective value (SV). Little is known about the dynamics of the neural processes that underpin this form of value-based decision making and no studies have investigated these processes outside of controlled laboratory settings. The current study investigated the spatio-temporal dynamics that accompany economic valuation of products using mobile electroencephalography (EEG) and eye tracking techniques. Participants viewed and rated images of household products in a gallery setting while EEG and eye tracking data were collected wirelessly. A Becker-DeGroot-Marschak (BDM) auction task was subsequently used to quantify the individual's willingness to pay (WTP) for each product. WTP was used to classify products into low, low medium, high medium and high economic value conditions. Eye movement related potentials (EMRP) were examined, and independent component analysis (ICA) was used to separate sources of activity from grand averaged EEG data. Four independent components (ICs) of EMRPs were modulated by WTP (i.e., SV) in the latency range of 150-250 ms. Of the four value-sensitive ICs, one IC displayed enhanced amplitude for all value conditions excluding low value, and another IC presented enhanced amplitude for low value products only. The remaining two value-sensitive ICs resolved inter-mediate levels of SV. Our study quantified, for the first time, the neural processes involved in economic value based decisions in a natural setting. Results suggest that multiple spatio-temporal brain activation patterns mediate the attention and aversion of products which could reflect an early valuation system. The EMRP parietal P200 component could reflect an attention allocation mechanism that separates the lowest-value products (IC7) from products of all other value (IC4), suggesting that low-value items are categorized early on as being aversive. While none of the ICs showed linear amplitude changes that parallel SV's of products, results suggest that a combination of multiple components may sub-serve a fine-grained resolution of the SV of products.

Pleasant and unpleasant odour-face combinations influence face and odour perception: An event-related potential study.

Odours alter evaluations of concurrent visual stimuli. However, neural mechanisms underlying the effects of congruent and incongruent odours on facial expression perception are not clear. Moreover, the influence of emotional faces on odour perception is not established. We investigated the effects of one pleasant and one unpleasant odour paired with happy and disgusted faces, on subjective ratings and ERP responses to faces. Participants rated the pleasantness of happy and disgusted faces that appeared during 3s pleasant or unpleasant odour pulses, or without odour. Odour pleasantness and intensity ratings were recorded in each trial. EEG was recorded continuously using a 128-channel system. Happy and disgusted faces paired with pleasant and unpleasant odour were rated as more or less pleasant, respectively, compared to the same faces presented in the other odour conditions. Odours were rated as more pleasant when paired with happy faces, and unpleasant odour was rated more intense when paired with disgusted faces. Unpleasant odour paired with disgusted faces also decreased inspiration. Odour-face interactions were evident in the N200 and N400 components. Our results reveal bi-directional effects of odours and faces, and suggest that odour-face interactions may be represented in ERP components. Pairings of unpleasant odour and disgusted faces resulted in stronger hedonic ratings, ERP changes, increased odour intensity ratings and respiratory adjustment. This finding likely represents heightened adaptive responses to multimodal unpleasant stimuli, prompting appropriate behaviour in the presence of danger.