A matter of availability: sharper tuning for memorized than for perceived stimulus features.

Our visual environment is relatively stable over time. An optimized visual system could capitalize on this by devoting less representational resources to objects that are physically present. The vividness of subjective experience, however, suggests that externally available (perceived) information is more strongly represented in neural signals than memorized information. To distinguish between these opposing predictions, we use EEG multivariate pattern analysis to quantify the representational strength of task-relevant features in anticipation of a change-detection task. Perceptual availability was manipulated between experimental blocks by either keeping the stimulus available on the screen during a 2-s delay period (perception) or removing it shortly after its initial presentation (memory). We find that task-relevant (attended) memorized features are more strongly represented than irrelevant (unattended) features. More importantly, we find that task-relevant features evoke significantly weaker representations when they are perceptually available compared with when they are unavailable. These findings demonstrate that, contrary to what subjective experience suggests, vividly perceived stimuli elicit weaker neural representations (in terms of detectable multivariate information) than the same stimuli maintained in visual working memory. We hypothesize that an efficient visual system spends little of its limited resources on the internal representation of information that is externally available anyway.

The Effect of Cognitive Load on Auditory Susceptibility During Automated Driving.

OBJECTIVE: We experimentally test the effect of cognitive load on auditory susceptibility during automated driving. BACKGROUND: In automated vehicles, auditory alerts are frequently used to request human intervention. To ensure safe operation, human drivers need to be susceptible to auditory information. Previous work found reduced susceptibility during manual driving and in a lesser amount during automated driving. However, in practice, drivers also perform nondriving tasks during automated driving, of which the associated cognitive load may further reduce susceptibility to auditory information. We therefore study the effect of cognitive load during automated driving on auditory susceptibility. METHOD: Twenty-four participants were driven in a simulated automated car. Concurrently, they performed a task with two levels of cognitive load: repeat a noun or generate a verb that expresses the use of this noun. Every noun was followed by a probe stimulus to elicit a neurophysiological response: the frontal P3 (fP3), which is a known indicator for the level of auditory susceptibility. RESULTS: The fP3 was significantly lower during automated driving with cognitive load compared with without. The difficulty level of the cognitive task (repeat or generate) showed no effect. CONCLUSION: Engaging in other tasks during automated driving decreases auditory susceptibility as indicated by a reduced fP3. APPLICATION: Nondriving task can create additional cognitive load. Our study shows that performing such tasks during automated driving reduces the susceptibility for auditory alerts. This can inform designers of semi-automated vehicles (SAE levels 3 and 4), where human intervention might be needed.

Annual variation in attentional response after methylphenidate treatment.

Prevalence rates of attention-deficit/hyperactivity disorder (ADHD) differ with geographical areas varying in sunlight intensity. Sun- or daylight reaching the retina establishes entrainment of the circadian clock to daylight. Changes herein, hence, alterations in clock alignment, could be reflected indirectly in inattention via sleep duration. We here studied (1) annual variation in inattention at treatment initiation; (2) annual variation in response to ADHD treatment [methylphenidate (MPH)] by day of treatment initiation; and (3) dose dependence. We predicted least baseline inattention during a period of high sunlight intensity implying more room for improvement (i.e., a better treatment response) when sunlight intensity is low. These hypotheses were not confirmed. High-dose treated patients, however, had significantly better attention after treatment than low-dosed treated patients, only when treated in the period from winter to summer solstice. Change in solar irradiance (SI) during low-dosed treatment period was negatively related to attentional improvement. The above described findings were primarily found in inattention ratings and replicated in omission errors on a continuous performance task. Daylight and inattention have been proposed to be related via mediation of the circadian system. One mechanism of MPH may be to enhance sensitivity to the diurnal entrainment to sunlight and the question can be raised whether appropriate lighting could potentiate the effects of stimulants.

Resting-state theta/beta EEG ratio is associated with reward- and punishment-related reversal learning.

Prior research has shown that the ratio between resting-state theta (4-7 Hz)-beta (13-30 Hz) oscillations in the electroencephalogram (EEG) is associated with reward- and punishment-related feedback learning and risky decision making. However, it remains unclear whether the theta/beta EEG ratio is also an electrophysiological index for poorer behavioral adaptation when reward and punishment contingencies change over time. The aim of the present study was to investigate whether resting-state theta (4-7 Hz)-beta (13-30 Hz) EEG ratio correlated with reversal learning. A 4-min resting-state EEG was recorded and a gambling task with changing reward-punishment contingencies was administered in 128 healthy volunteers. Results showed an inverse relationship between theta/beta EEG ratio and reversal learning. Our findings replicate and extend previous findings by showing that higher midfrontal theta/beta EEG ratios are associated with poorer reversal learning and behavioral adaptive responses under changing environmental demands.

Action preparation shapes processing in early visual cortex.

Preparation for an action, such as grasping an object, is accompanied by an enhanced perception of the object's action-relevant features, such as orientation and size. Cortical feedback from motor planning areas to early visual areas may drive this enhanced perception. To examine whether action preparation modulates activity in early human visual cortex, subjects grasped or pointed to oriented objects while high-resolution fMRI data were acquired. Using multivoxel pattern analysis techniques, we could decode with >70% accuracy whether a grasping or pointing action was prepared from signals in visual cortex as early as V1. These signals in early visual cortex were observed even when actions were only prepared but not executed. Anterior parietal cortex, on the other hand, showed clearest modulation for actual movements. This demonstrates that preparation of actions, even without execution, modulates relevant neuronal populations in early visual areas.

The effect of attenuating noradrenergic neurotransmission by clonidine on brain activity measures of visuospatial attention.

OBJECTIVE: In the current study, we investigated the role of noradrenaline in directing (bias) and disengagement of visuospatial attention. METHODS: We assessed the effect of clonidine on event-related brain potential (ERP) reflections of bias and disengagement in a double-blind placebo-controlled crossover design. An initial dose of 200-µg clonidine was replaced by 100 µg because of marked side effects. Twenty-one healthy male participants performed the visual-spatial cueing task while an electroencephalogram (EEG) was recorded. The behavioral output is the validity effect (benefit of cueing in terms of reaction time to targets). ERP indices for bias were the cue-related early directing attention negativity and late directing attention positivity, and the target-elicited P1 and N1 modulations by validity ('validity-effect'). The ERP index for disengagement was the target-elicited 'late positive deflection' modulation by validity. Behavioral analyses were performed on 16 participants, electrophysiological analyses on a subset (n=9). RESULTS: Clonidine attenuated the N1 effect, albeit in a subsample. Neither cue-elicited ERPs nor the behavioral validity effect were affected. Clonidine-induced blood pressure reduction was correlated with the reduction of the late positive deflection effect under clonidine. CONCLUSION: Clonidine attenuated the result of bias in a subsample and may have a modulating effect on disengagement.

How salience enhances inhibitory control: An analysis of electro-cortical mechanisms.

Stop-signal tasks (SSTs) combined with human electro-cortical recordings (Event-Related Potentials, ERPs) have revealed mechanisms associated with successful stopping (relative to failed), presumably contributing to inhibitory control. The corresponding ERP signatures have been labeled stop N1 (+/- 100-ms latency), stop N2 (200 ms), and stop P3 (160-250 ms), and argued to reflect more sensory-specific (N1) versus more generic (N2, P3) mechanisms. However, stop N1 and stop N2, as well as latencies of stop-P3, appear to be quite inconsistent across studies. The present work addressed the possible influence of stop-signal salience, expecting high salience to induce clear stop N1s but reduced stop N2s, and short-latency stop P3s. Three SST varieties were combined with high-resolution EEG. An imperative visual (go) stimulus was occasionally followed by a subsequent (stop) stimulus that signalled to withhold the just initiated response. Stop-Signal Reaction Times (SSRTs) decreased linearly from visual-low to visual-high-salience to auditory. Auditory Stop N1 was replicated. A C1-like visual evoked potential (latency < 100 ms) was observed only with high salience, but not robustly associated with successful versus failed stops. Using the successful-failed contrast a visual stop-N1 analogue (112-156 ms post-stop-signal) was identified, as was right-frontal stop N2, but neither was sensitive to salience. Stop P3 had shorter latency for high than for low salience, and the extent of the early high-salience stop P3 correlated inversely with SSRT. These results suggest that salience-enhanced inhibitory control as manifest in SSRTs is associated with generic rather than sensory-specific electrocortical mechanisms.

Drift-diffusion modeling reveals that masked faces are preconceived as unfriendly.

During the COVID-19 pandemic, the use of face masks has become a daily routine. Studies have shown that face masks increase the ambiguity of facial expressions which not only affects (the development of) emotion recognition, but also interferes with social interaction and judgement. To disambiguate facial expressions, we rely on perceptual (stimulus-driven) as well as preconceptual (top-down) processes. However, it is unknown which of these two mechanisms accounts for the misinterpretation of masked expressions. To investigate this, we asked participants (N = 136) to decide whether ambiguous (morphed) facial expressions, with or without a mask, were perceived as friendly or unfriendly. To test for the independent effects of perceptual and preconceptual biases we fitted a drift-diffusion model (DDM) to the behavioral data of each participant. Results show that face masks induce a clear loss of information leading to a slight perceptual bias towards friendly choices, but also a clear preconceptual bias towards unfriendly choices for masked faces. These results suggest that, although face masks can increase the perceptual friendliness of faces, people have the prior preconception to interpret masked faces as unfriendly.

Differential functional roles of slow-wave and oscillatory-α activity in visual sensory cortex during anticipatory visual-spatial attention.

Markers of preparatory visual-spatial attention in sensory cortex have been described both as lateralized, slow-wave event-related potential (ERP) components and as lateralized changes in oscillatory-electroencephalography alpha power, but the roles of these markers and their functional relationship are still unclear. Here, 3 versions of a visual-spatial cueing paradigm, differing in perceptual task difficulty and/or response instructions, were used to investigate the functional relationships between posterior oscillatory-alpha changes and our previously reported posterior, slow-wave biasing-related negativity (swBRN) ERP activity. The results indicate that the swBRN reflects spatially specific, pretarget preparatory activity sensitive to the expected perceptual difficulty of the target detection task, correlating in both location and strength with the early sensory-processing N1 ERP to the target, consistent with reflecting a preparatory baseline-shift mechanism. In contrast, contralateral event-related decreases in alpha-band power were relatively insensitive to perceptual difficulty and differed topographically from both the swBRN and target N1. Moreover, when response instructions emphasized making immediate responses to targets, compared with prescribing delayed responses, contralateral alpha-event-related desynchronization activity was particularly strong and correlated with the longer latency target-P3b activity. Thus, in contrast to the apparent perceptual-biasing role of swBRN activity, contralateral posterior alpha activity may represent an attentionally maintained task set linking stimulus-specific information and task-specific response requirements.

Recruitment of a long-term memory supporting neural network during repeated maintenance of a multi-item abstract visual image in working memory.

Humans can flexibly transfer information between different memory systems. Information in visual working memory (VWM) can for instance be stored in long-term memory (LTM). Conversely, information can be retrieved from LTM and temporarily held in WM when needed. It has previously been suggested that a neural transition from parietal- to midfrontal activity during repeated visual search reflects transfer of information from WM to LTM. Whether this neural transition indeed reflects consolidation and is also observed when memorizing a rich visual scene (rather than responding to a single target), is not known. To investigate this, we employed an EEG paradigm, in which abstract six-item colour-arrays were repeatedly memorized and explicitly visualized, or merely attended to. Importantly, we tested the functional significance of a potential neural shift for longer-term consolidation in a subsequent recognition task. Our results show a gradually enhanced- and sustained modulation of the midfrontal P170 component and a decline in parietal CDA, during repeated WM maintenance. Improved recollection/visualization of memoranda upon WM-cueing, was associated with contralateral parietal- and right temporal activity. Importantly, only colour-arrays previously held in WM, induced a greater midfrontal P170-response, together with left temporal- and late centro-parietal activity, upon re-exposure. These findings provide evidence for recruitment of an LTM-supporting neural network which facilitates visual WM maintenance.

Eye tracking unconscious face-to-face confrontations: dominance motives prolong gaze to masked angry faces.

In primates, dominance/submission relationships are generally automatically and nonaggressively established in face-to-face confrontations. Researchers have argued that this process involves an explicit psychological stress-manipulation mechanism: Striding with a threatening expression, while keeping direct eye contact, outstresses rivals so that they submissively avert their gaze. In contrast, researchers have proposed a reflexive and implicit modulation of face-to-face confrontation in humans, on the basis of evidence that dominant and submissive individuals exhibit vigilant and avoidant responses, respectively, to facial anger in masked emotional Stroop tasks. However, these tasks do not provide an ecologically valid index of gaze behavior. Therefore, we directly measured gaze responses to masked angry, happy, and neutral facial expressions with a saccade-latency paradigm and found that increased dominance traits predict a more prolonged gaze to (or reluctance to avert gaze from) masked anger. Furthermore, greater non-dominance-related reward sensitivity predicts more persistent gaze to masked happiness. These results strongly suggest that implicit and reflexive mechanisms underlie dominant and submissive gaze behavior in face-to-face confrontations.

Attentional bias in high- and low-anxious individuals: evidence for threat-induced effects on engagement and disengagement.

Attentional bias to threatening visual stimuli (words or pictures) is commonly present in anxious individuals, but not in non-anxious people. There is evidence to show that attentional bias to threat can be induced in all individuals when threat is imposed by threat not of symbolic nature, but by cues that predict aversive stimulation (loud noise or electric shock). However, it is not known whether attentional bias in such situations is still influenced by individual differences in anxiety. This question was addressed in two experiments using a spatial cuing task in which visual cues predicted the occurrence of an aversive event consisting of a loud human scream. Speeded attentional engagement to threat cues was positively correlated with trait anxiety in Experiment 1. Experiment 2 showed that speeded attentional engagement was present only in participants selected for high anxiety but not in low-anxious participants. In both experiments, slower disengagement from threat cues was found in all participants, irrespective of their trait anxiety levels.

No consistent startle modulation by reward.

Previous studies have not clearly demonstrated whether motivational tendencies during reward feedback are mainly characterized by appetitive responses to a gain or mainly by aversive consequences of reward omission. In the current study this issue was addressed employing a passive head or tails game and using the startle reflex as an index of the appetitive-aversive continuum. A second aim of the current study was to use startle-reflex modulation as a means to compare the subjective value of monetary rewards of varying magnitude. Startle responses after receiving feedback that a potential reward was won or not won were compared with a baseline condition without a potential gain. Furthermore, startle responses during anticipation of no versus potential gain were compared. Consistent with previous studies, startle-reflex magnitudes were significantly potentiated when participants anticipated a reward compared to no reward, which may reflect anticipatory arousal. Specifically for the largest reward (20-cents) startle magnitudes were potentiated when a reward was at stake but not won, compared to a neutral baseline without potential gain. In contrast, startle was not inhibited relative to baseline when a reward was won. This suggests that startle modulation during feedback is better characterized in terms of potentiation when missing out on reward rather than in terms of inhibition as a result of winning. However, neither of these effects were replicated in a more targeted second experiment. The discrepancy between these experiments may be due to differences in motivation to obtain rewards or differences in task engagement. From these experiments it may be concluded that the nature of the processing of reward feedback and reward cues is very sensitive to experimental parameters and settings. These studies show how apparently modest changes in these parameters and settings may lead to quite different modulations of appetitive/aversive motivation. A future experiment may shed more light on the question whether startle-reflex modulation after feedback is indeed mainly characterized by the aversive consequences of reward omission for relatively large rewards.

Neuro-Cardiac-Guided TMS (NCG TMS): A replication and extension study.

Neuro-Cardiac-Guided Transcranial Magnetic Stimulation (NCG-TMS) was studied for its potential to specifically target the frontal-vagal network. Previous research demonstrated that prefrontal stimulation led to significant heartrate slowing. We aimed to replicate these results in a larger sample and extend the findings to investigate dose-response relationships, reproducibility and stimulation frequency (10 Hz and intermittent theta burst (iTBS)). Data of forty-five healthy controls were analyzed, of which 28 received 10 Hz TMS (NCG-TMS) and 27 iTBS (NCG-iTBS; 10 received both protocols) at different stimulation sites according to the 10-20-EEG system. NCG-TMS yielded a relative heartrate deceleration at the F3/4 coil position replicating earlier studies. Both internal consistency and dose-response relationships were found. For NCG-iTBS adverse events were reported and topography for frontal-vagal activation was more lateralised relative to NCG-TMS. These results indicate that we were able to transsynaptically stimulate the frontal-vagal network and that excitability thresholds for the prefrontal cortex may differ relative to motor cortex.

Lateralized frontal eye field activity precedes occipital activity shortly before saccades: evidence for cortico-cortical feedback as a mechanism underlying covert attention shifts.

When an eye movement is prepared, attention is shifted toward the saccade end-goal. This coupling of eye movements and spatial attention is thought to be mediated by cortical connections between the FEFs and the visual cortex. Here, we present evidence for the existence of these connections. A visual discrimination task was performed while recording the EEG. Discrimination performance was significantly improved when the discrimination target and the saccade target matched. EEG results show that frontal activity precedes occipital activity contralateral to saccade direction when the saccade is prepared but not yet executed; these effects were absent in fixation conditions. This is consistent with the idea that the FEF exerts a direct modulatory influence on the visual cortex and enhances perception at the saccade end-goal.

Cannabinoid modulations of resting state EEG θ power and working memory are correlated in humans.

Object representations in working memory depend on neural firing that is phase-locked to oscillations in the theta band (4-8 Hz). Cannabis intake disrupts synchronicity of theta oscillations and interferes with memory performance. Sixteen participants smoked cigarettes containing 0.0, 29.3, 49.1, or 69.4 mg Delta(9)-tetrahydrocannabinol (THC) in a randomized crossover design and performed working memory and general attention tasks. Dose-dependent effects of THC were observed for resting state EEG theta and beta power, working memory (per-item search time), and attentional performance (percent errors and RT). The THC effects on EEG theta power and memory performance were correlated, whereas other EEG and behavioral effects were not. These findings confirm and extend previous results in rodents and humans, and corroborate a neurocomputational model that postulates that temporal aspects of information processing in working memory depend causally on nested oscillations in the theta and gamma (>30 Hz) bands.

Dopaminergic and noradrenergic manipulation of anticipatory reward and probability event-related potentials.

Predicting what will happen in the future in terms of potential reward is essential in daily life. The aim of the current study was to investigate the neurotransmitter systems involved in the anticipation of reward value and probability. We hypothesized that dopaminergic and noradrenergic antagonism would affect anticipation of reward value and probability, respectively. Twenty-three healthy participants were included in a haloperidol (2 mg) × clonidine (0.150 mg) × placebo cross-over design and subjected to a Go/NoGo experimental task during which cues signaled the probability of subsequent target appearance. Reward value (amount of money that could be won for correct and fast responding to the target) as well as probability of target appearance was orthogonally manipulated across four task blocks. Cue-elicited EEG event-related potentials were recorded to assess anticipation of value and probability, respectively. The processing of reward value was affected by dopaminergic antagonism (haloperidol), as evidenced by reduction of the reward-related positivity and P300 to reward cues. This reduction was specifically significant for subjects with high baseline dopamine levels for the P300 and most pronounced for these subjects for the reward-related positivity. In contrast, the processing of reward probability was affected by noradrenergic antagonism (clonidine). In addition, both drugs reduced overall performance (omission rate, response speed variability). We conclude that at least anticipation of reward value and probability, respectively, is specifically affected by dopaminergic versus noradrenergic antagonism.

The influence of cognitive load on susceptibility to audio.

In this study we evaluate how cognitive load affects susceptibility to auditory signals. Previous research has used the frontal P3 (fP3) event related potential response to auditory novel stimuli as an index for susceptibility to auditory signals. This work demonstrated that tasks that induce cognitive load such as visual and manual tasks, reduced susceptibility. It is however unknown whether cognitive load without visual or manual components also reduces susceptibility. To investigate this, we induced cognitive load by means of the verb generation task, in which participants need to think about a verb that matches a noun. The susceptibility to auditory signals was measured by recording the event related potential in response to a successively presented oddball probe stimulus at 3 different inter-stimulus intervals, 0 ms, 200 ms or 400 ms after the offset of the noun from the verb generation task. An additional control baseline condition, in which oddball response was probed without a verb generation task, was also included. Results show that the cognitive load associated with the verb task reduces fP3 response (and associated auditory signal susceptibility) compared to baseline, independent of presentation interval. This suggests that not only visual and motor processing, but also cognitive load without visual or manual components, can reduce susceptibility to auditory signals and alerts.

A frontal-vagal network theory for Major Depressive Disorder: Implications for optimizing neuromodulation techniques.

Major Depressive Disorder (MDD) is a psychiatric disorder characterized by high comorbidity with cardiovascular disease. Furthermore, a combination of high heart rate (HR) and low heart rate variability (HRV) has been frequently reported in depressed patients. The present review proposes a frontal-vagal (brain-heart) network that overlaps with functional nodes of the depression network. Moreover, we summarize neuromodulation studies that have targeted key nodes in this depression network, with subsequent impact on heart rate (HR) or heart-rate-variability (HRV), such as the dorsolateral prefrontal cortex (DLPFC), subgenual anterior cingulate cortex (sgACC), and the vagus nerve (VN). Based on the interplay of this frontal-vagal network, we emphasize the importance of including HR and HRV measurements in human depression studies, in particular those that conduct neuromodulation, in order to obtain a better understanding of the pathways that are affected, and we explore the possibilities of using this frontal-vagal interplay as a method for target engagement in neuromodulation treatments. This frontal-vagal network theory opens-up the possibility for individualizing neuromodulation treatments such as rTMS. A recent development called Neuro-Cardiac-Guided TMS (NCG-TMS), was developed based on this theory, and an individual-participant meta-analysis is presented. Four studies provide consistent and replicable support for NCG-TMS as a target engagement method, with consistent HR deceleration during frontal TMS and HR acceleration during motor strip TMS.

The development of automated access to symbolic and non-symbolic number knowledge in children: an ERP study.

Infants can visually detect changes in numerosity, which suggests that a (non-symbolic) numerosity system is already present early in life. This non-symbolic system is hypothesized to serve as the basis for the later acquired symbolic system. Little is known about the processes underlying the transition from the non-symbolic to symbolic code. In the current study we investigated the development of automatization of symbolic number processing in children from second (6.0 years) and fourth grade (8.0 years) and adults using a symbolic and non-symbolic size congruency task and event-related potentials (ERPs) as a measure. The comparison between symbolic and non-symbolic size congruency effects (SCEs) allowed us to disentangle processes necessary to perform the task from processes specific to numerosity notation. In contrast to previous studies, second graders already revealed a behavioral symbolic SCE similar to that of adults. In addition, the behavioral SCE increased for symbolic and decreased for non-symbolic notation with increasing age. For all age groups, the ERP data showed that the two magnitudes interfered at a level before selective activation of the response system, for both notations. However, only for the second graders distinct processes were recruited to perform the symbolic size comparison task. This shift in recruited processes for the symbolic task only might reflect the functional specialization of the parietal cortex.