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Josefine Waldthaler; Alexander Sperlich; Aylin König; Charlotte Stüssel; Frank Bremmer; Lars Timmermann; David Pedrosa
High (130 Hz)- and mid (60 Hz)-frequency deep brain stimulation in the subthalamic nucleus differentially modulate response inhibition: A preliminary combined EEG and eye tracking study Journal Article
In: NeuroImage: Clinical, vol. 37, pp. 1–11, 2023.
While deep brain stimulation (DBS) in the subthalamic nucleus (STN) improves motor functions in Parkinson's disease (PD), it may also increase impulsivity by interfering with the inhibition of reflexive responses. The aim of this study was to investigate if varying the pulse frequency of STN-DBS has a modulating effect on response inhibition and its neural correlates. For this purpose, 14 persons with PD repeated an antisaccade task in three stimulation settings (DBS off, high-frequency DBS (130 Hz), mid-frequency DBS (60 Hz)) in a randomized order, while eye movements and brain activity via high-density EEG were recorded. On a behavioral level, 130 Hz DBS stimulation had no effect on response inhibition measured as antisaccade error rate, while 60 Hz DBS induced a slight but significant reduction of directional errors compared with the DBS-off state and 130 Hz DBS. Further, stimulation with both frequencies decreased the onset latency of correct antisaccades, while increasing the latency of directional errors. Time-frequency domain analysis of the EEG data revealed that 60 Hz DBS was associated with an increase in preparatory theta power over a midfrontal region of interest compared with the off-DBS state which is generally regarded as a marker of increased cognitive control. While no significant differences in brain activity over mid- and lateral prefrontal regions of interest emerged between the 60 Hz and 130 Hz conditions, both stimulation frequencies were associated with a stronger midfrontal beta desynchronization during the mental preparation for correct antisaccades compared with DBS off-state which is discussed in the context of potentially enhanced proactive recruitment of the oculomotor network. Our preliminary findings suggest that mid-frequency STN-DBS may provide beneficial effects on response inhibition, while both 130 Hz- and 60 Hz STN-DBS may promote voluntary actions at the expense of slower reflexive responses.
Sebastian C. Coleman; Zelekha A. Seedat; Anna C. Whittaker; Agatha Lenartowicz; Karen J. Mullinger
Beyond the beta rebound: Post-task responses in oscillatory activity follow cessation of working memory processes Journal Article
In: NeuroImage, vol. 265, pp. 1–11, 2023.
Post-task responses (PTRs) are transitionary responses occurring for several seconds between the end of a stimulus/task and a period of rest. The most well-studied of these are beta band (13 – 30 Hz) PTRs in motor networks following movement, often called post-movement beta rebounds, which have been shown to differ in patients with schizophrenia and autism. Previous studies have proposed that beta PTRs reflect inhibition of task-positive networks to enable a return to resting brain activity, scaling with cognitive demand and reflecting cortical self-regulation. It is unknown whether PTRs are a phenomenon of the motor system, or whether they are a more general self-modulatory property of cortex that occur following cessation of higher cognitive processes as well as movement. To test this, we recorded magnetoencephalography (MEG) responses in 20 healthy participants to a working-memory task, known to recruit cortical networks associated with higher cognition. Our results revealed PTRs in the theta, alpha and beta bands across many regions of the brain, including the dorsal attention network (DAN) and lateral visual regions. These PTRs increased significantly (p < 0.05) in magnitude with working-memory load, an effect which is independent of oscillatory modulations occurring over the task period as well as those following individual stimuli. Furthermore, we showed that PTRs are functionally related to reaction times in left lateral visual (p < 0.05) and left parietal (p < 0.1) regions, while the oscillatory responses measured during the task period are not. Importantly, motor PTRs following button presses did not modulate with task condition, suggesting that PTRs in different networks are driven by different aspects of cognition. Our findings show that PTRs are not limited to motor networks but are widespread in regions which are recruited during the task. We provide evidence that PTRs have unique properties, scaling with cognitive load and correlating significantly with behaviour. Based on the evidence, we suggest that PTRs inhibit task-positive network activity to enable a transition to rest, however, further investigation is required to uncover their role in neuroscience and pathology.
Xianliang Ge; Yunxian Pan; Sujie Wang; Linze Qian; Jingjia Yuan; Jie Xu; Nitish Thakor; Yu Sun
Improving intention detection in single-trial classification through fusion of EEG and eye-tracker data Journal Article
In: IEEE Transactions on Human-Machine Systems, vol. 53, no. 1, pp. 132–141, 2023.
Intention decoding is an indispensable procedure in hands-free human-computer interaction (HCI). Conventional eye-tracking system using single-model fixation duration possibly issues commands ignoring users' real expectation. In the current study, an eye-brain hybrid brain-computer interface (BCI) interaction system was introduced for intention detection through fusion of multi-modal eye-track and ERP (a measurement derived from EEG) features. Eye-track and EEG data were recorded from 64 healthy participants as they performed a 40-min customized free search task of a fixed target icon among 25 icons. The corresponding fixation duration of eye-trackingw and ERP were extracted. Five previously-validated LDA-based classifiers (including RLDA, SWLDA, BLDA, SKLDA, and STDA) and the widely-used CNN method were adopted to verify the efficacy of feature fusion from both offline and pseudo-online analysis, and optimal approach was evaluated through modulating the training set and system response duration. Our study demonstrated that the input of multi-modal eye-track and ERP features achieved superior performance of intention detection in the single trial classification of active search task. And compared with single-model ERP feature, this new strategy also induced congruent accuracy across different classifiers. Moreover, in comparison with other classification methods, we found that the SKLDA exhibited the superior performance when fusing feature in offline test (ACC=0.8783
Zahra Azizi; Reza Ebrahimpour
Explaining integration of evidence separated by temporal gaps with frontoparietal circuit models Journal Article
In: Neuroscience, vol. 509, pp. 74–95, 2023.
Perceptual decisions rely on accumulating sensory evidence over time. However, the accumulation process is complicated in real life when evidence resulted from separated cues over time. Previous studies demonstrate that participants are able to integrate information from two separated cues to improve their performance invariant to an interval between the cues. However, there is no neural model that can account for accuracy and confidence in decisions when there is a time interval in evidence. We used behavioral and EEG datasets from a visual choice task —Random dot motion— with separated evidence to investigate three candid distributed neural networks. We showed that decisions based on evidence accumulation by separated cues over time are best explained by the interplay of recurrent cortical dynamics of centro-parietal and frontal brain areas while an uncertainty-monitoring module included in the model.
Constanze Schmitt; Milosz Krala; Frank Bremmer
Neural signatures of actively controlled self-motion and the subjective encoding of distance Journal Article
In: eNeuro, vol. 9, no. 6, pp. 1–18, 2022.
Navigating through an environment requires knowledge about one's direction of self-motion (heading) and traveled distance. Behavioral studies showed that human participants can actively reproduce a previously observed travel distance purely based on visual information. Here, we employed electroencephalography (EEG) to investigate the underlying neural processes. We measured, in human observers, event-related potentials (ERPs) during visually simulated straight-forward self-motion across a ground plane. The participants' task was to reproduce (active condition) double the distance of a previously seen self-displacement (passive condition) using a gamepad. We recorded the trajectories of self-motion during the active condition and played it back to the participants in a third set of trials (replay condition). We analyzed EEG activity separately for four electrode clusters: frontal (F), central (C), parietal (P), and occipital (O). When aligned to self-motion onset or offset, response modulation of the ERPs was stronger, and several ERP components had different latencies in the passive as compared with the active condition. This result is in line with the concept of predictive coding, which implies modified neural activation for self-induced versus externally induced sensory stimulation. We aligned our data also to the times when subjects passed the (objective) single distance d_obj and the (subjective) single distance d_sub. Remarkably, wavelet-based temporal-frequency analyses revealed enhanced theta-band activation for F, P, and O-clusters shortly before passing d_sub. This enhanced activation could be indicative of a navigation related representation of subjective distance. More generally, our study design allows to investigate subjective perception without interfering neural activation because of the required response action.
Poppy Sharp; Tjerk Gutteling; David Melcher; Clayton Hickey
Spatial attention tunes temporal processing in early visual cortex by speeding and slowing alpha oscillations Journal Article
In: Journal of Neuroscience, vol. 42, no. 41, pp. 7824–7832, 2022.
The perception of dynamic visual stimuli relies on two apparently conflicting perceptual mechanisms: rapid visual input must sometimes be integrated into unitary percepts but at other times must be segregated or parsed into separate objects or events. Though they have opposite effects on our perceptual experience, the deployment of spatial attention benefits both operations. Little is known about the neural mechanisms underlying this impact of spatial attention on temporal perception. Here, we record magnetoencephalography (MEG) in male and female humans to demonstrate that the deployment of spatial attention for the purpose of segregating or integrating visual stimuli impacts prestimulus oscillatory activity in retinotopic visual brain areas where the attended location is represented. Alpha band oscillations contralateral to an attended location are therefore faster than ipsilateral oscillations when stimuli appearing at this location will need to be segregated, but slower in expectation of the need for integration, consistent with the idea that a frequency is linked to perceptual sampling rate. These results demonstrate a novel interaction between temporal visual processing and the allocation of attention in space.
Elio Balestrieri; Niko A. Busch
Spontaneous alpha-band oscillations bias subjective contrast perception Journal Article
In: Journal of Neuroscience, pp. 1–31, 2022.
Perceptual decisions depend both on the features of the incoming stimulus and on the ongoing brain activity at the moment the stimulus is received. Specifically, trial-to-trial fluctuations in cortical excitability have been linked to fluctuations in the amplitude of prestimulus $alpha$ oscillations ($sim$8-13 Hz), which are in turn are associated with fluctuations in subjects' tendency to report the detection of a stimulus. It is currently unknown whether $alpha$ oscillations bias postperceptual decision-making, or even bias subjective perception itself. To answer this question, we used a contrast discrimination task in which both male and female human subjects reported which of two gratings (one in each hemifield) was perceived as having a stronger contrast. Our EEG analysis showed that subjective contrast was reduced for the stimulus in the hemifield represented in the hemisphere with relatively stronger prestimulus $alpha$ amplitude, reflecting reduced cortical excitability. Furthermore, the strength of this spontaneous hemispheric lateralization was strongly correlated with the magnitude of individual subjects' biases, suggesting that the spontaneous patterns of $alpha$ lateralization play a role in explaining the intersubject variability in contrast perception. These results indicate that spontaneous fluctuations in cortical excitability, indicated by patterns of prestimulus $alpha$ amplitude, affect perceptual decisions by altering the phenomenological perception of the visual world.
Angela Radetz; Markus Siegel
Spectral fingerprints of cortical neuromodulation Journal Article
In: Journal of Neuroscience, vol. 42, no. 18, pp. 3836–3846, 2022.
Pupil size has been established as a versatile marker of noradrenergic and cholinergic neuromodulation, which has profound effects on neuronal processing, cognition, and behavior. However, little is known about the cortical control and effects of pupil-linked neuromodulation. Here, we show that pupil dynamics are tightly coupled to temporally, spectrally, and spatially specific modulations of local and large-scale cortical population activity in the human brain. We quantified the dynamics of band-limited cortical population activity in resting human subjects using magnetoencephalography and investigated how neural dynamics were linked to simultaneously recorded pupil dynamics. Our results show that pupil-linked neuromodulation does not merely affect cortical population activity in a stereotypical fashion. Instead, we identified three frontal, precentral, and occipitoparietal networks, in which local population activity with distinct spectral profiles in the theta, beta, and alpha bands temporally preceded and followed changes in pupil size. Furthermore, we found that amplitude coupling at;16 Hz in a large-scale frontoparietal network predicted pupil dynamics. Our results unravel network-specific spectral fingerprints of cortical neuromodulation in the human brain that likely reflect both the causes and effects of neuromodulation.
Kumari Liza; Supratim Ray
Local interactions between steady-state visually evoked potentials at nearby flickering frequencies Journal Article
In: Journal of Neuroscience, vol. 42, no. 19, pp. 3965–3974, 2022.
Steady-state visually evoked potentials (SSVEPs) are widely used to index top-down cognitive processing in human electroencephalogram (EEG) studies. Typically, two stimuli flickering at different temporal frequencies (TFs) are presented, each producing a distinct response in the EEG at its flicker frequency. However, how SSVEP responses in EEGs are modulated in the presence of a competing flickering stimulus just because of sensory interactions is not well understood. We have previously shown in local field potentials (LFPs) recorded from awake monkeys that when two overlapping full-screen gratings are counterphased at different TFs, there is an asymmetric SSVEP response suppression, with greater suppression from lower TFs, which further depends on the relative orientations of the gratings (stronger suppression and asymmetry for parallel compared with orthogonal gratings). Here, we first confirmed these effects in both male and female human EEG recordings. Then, we mapped the response suppression of one stimulus (target) by a competing stimulus (mask) over a much wider range than the previous study. Surprisingly, we found that the suppression was not stronger at low frequencies in general, but systematically varied depending on the target TF, indicating local interactions between the two competing stimuli. These results were confirmed in both human EEG and monkey LFP and electrocorticogram (ECoG) data. Our results show that sensory interactions between multiple SSVEPs are more complex than shown previously and are influenced by both local and global factors, underscoring the need to cautiously interpret the results of studies involving SSVEP paradigms.SIGNIFICANCE STATEMENT Steady-state visually evoked potentials (SSVEPs) are extensively used in human cognitive studies and brain-computer interfacing applications where multiple stimuli flickering at distinct frequencies are concurrently presented in the visual field. We recently characterized interactions between competing flickering stimuli in animal recordings and found that stimuli flickering slowly produce larger suppression. Here, we confirmed these in human EEGs, and further characterized the interactions by using a much wider range of target and competing (mask) frequencies in both human EEGs and invasive animal recordings. These revealed a new "local" component, whereby the suppression increased when competing stimuli flickered at nearby frequencies. Our results highlight the complexity of sensory interactions among multiple SSVEPs and underscore the need to cautiously interpret studies involving SSVEP paradigms.
Arno Libert; Arne Van Den Kerchove; Benjamin Wittevrongel; Marc M. Van Hulle
Analytic beamformer transformation for transfer learning in motion-onset visual evoked potential decoding Journal Article
In: Journal of Neural Engineering, vol. 19, pp. 1–16, 2022.
Objective. While decoders of electroencephalography-based event-related potentials (ERPs) are routinely tailored to the individual user to maximize performance, developing them on populations for individual usage has proven much more challenging. We propose the analytic beamformer transformation (ABT) to extract phase and/or magnitude information from spatiotemporal ERPs in response to motion-onset stimulation. Approach. We have tested ABT on 52 motion-onset visual evoked potential (mVEP) datasets from 26 healthy subjects and compared the classification accuracy of support vector machine (SVM), spatiotemporal beamformer (stBF) and stepwise linear discriminant analysis (SWLDA) when trained on individual subjects and on a population thereof. Main results. When using phase- and combined phase/magnitude information extracted by ABT, we show significant improvements in accuracy of population-trained classifiers applied to individual users (p < 0.001). We also show that 450 epochs are needed for a correct functioning of ABT, which corresponds to 2 min of paradigm stimulation. Significance. We have shown that ABT can be used to create population-trained mVEP classifiers using a limited number of epochs. We expect this to pertain to other ERPs or synchronous stimulation paradigms, allowing for a more effective, population-based training of visual BCIs. Finally, as ABT renders recordings across subjects more structurally invariant, it could be used for transfer learning purposes in view of plug-and-play BCI applications.
Klaartje T. H. Heinen; J. Leon Kenemans; Stefan Stigchel
Recruitment of a long-term memory supporting neural network during repeated maintenance of a multi-item abstract visual image in working memory Journal Article
In: Scientific Reports, vol. 12, pp. 1–17, 2022.
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.
Katherine L. Hermann; Shridhar R. Singh; Isabelle A. Rosenthal; Dimitrios Pantazis; Bevil R. Conway
Temporal dynamics of the neural representation of hue and luminance polarity Journal Article
In: Nature Communications, vol. 13, pp. 1–19, 2022.
Hue and luminance contrast are basic visual features. Here we use multivariate analyses of magnetoencephalography data to investigate the timing of the neural computations that extract them, and whether they depend on common neural circuits. We show that hue and luminance-contrast polarity can be decoded from MEG data and, with lower accuracy, both features can be decoded across changes in the other feature. These results are consistent with the existence of both common and separable neural mechanisms. The decoding time course is earlier and more temporally precise for luminance polarity than hue, a result that does not depend on task, suggesting that luminance contrast is an updating signal that separates visual events. Meanwhile, cross-temporal generalization is slightly greater for representations of hue compared to luminance polarity, providing a neural correlate of the preeminence of hue in perceptual grouping and memory. Finally, decoding of luminance polarity varies depending on the hues used to obtain training and testing data. The pattern of results is consistent with observations that luminance contrast is mediated by both L-M and S cone sub-cortical mechanisms.
Beatriz Herrera; Jacob A. Westerberg; Michelle S. Schall; Alexander Maier; Geoffrey F. Woodman; Jeffrey D. Schall; Jorge J. Riera
Resolving the mesoscopic missing link: Biophysical modeling of EEG from cortical columns in primates Journal Article
In: NeuroImage, vol. 263, pp. 1–14, 2022.
Event-related potentials (ERP) are among the most widely measured indices for studying human cognition. While their timing and magnitude provide valuable insights, their usefulness is limited by our understanding of their neural generators at the circuit level. Inverse source localization offers insights into such generators, but their solutions are not unique. To address this problem, scientists have assumed the source space generating such signals comprises a set of discrete equivalent current dipoles, representing the activity of small cortical regions. Based on this notion, theoretical studies have employed forward modeling of scalp potentials to understand how changes in circuit-level dynamics translate into macroscopic ERPs. However, experimental validation is lacking because it requires in vivo measurements of intracranial brain sources. Laminar local field potentials (LFP) offer a mechanism for estimating intracranial current sources. Yet, a theoretical link between LFPs and intracranial brain sources is missing. Here, we present a forward modeling approach for estimating mesoscopic intracranial brain sources from LFPs and predict their contribution to macroscopic ERPs. We evaluate the accuracy of this LFP-based representation of brain sources utilizing synthetic laminar neurophysiological measurements and then demonstrate the power of the approach in vivo to clarify the source of a representative cognitive ERP component. To that end, LFP was measured across the cortical layers of visual area V4 in macaque monkeys performing an attention demanding task. We show that area V4 generates dipoles through layer-specific transsynaptic currents that biophysically recapitulate the ERP component through the detailed forward modeling. The constraints imposed on EEG production by this method also revealed an important dissociation between computational and biophysical contributors. As such, this approach represents an important bridge between laminar microcircuitry, through the mesoscopic activity of cortical columns to the patterns of EEG we measure at the scalp.
Zhenhong Hu; Immanuel B. H. Samuel; Sreenivasan Meyyappan; Ke Bo; Chandni Rana; Mingzhou Ding
Aftereffects of frontoparietal theta tACS on verbal working memory: Behavioral and neurophysiological analysis Journal Article
In: IBRO Neuroscience Reports, vol. 13, pp. 469–477, 2022.
Verbal working memory is supported by a left-lateralized frontoparietal theta oscillatory (4–8 Hz) network. We tested whether stimulating the left frontoparietal network at theta frequency during verbal working memory can produce observable after-stimulation effects in behavior and neurophysiology. Weak theta-band alternating electric currents were delivered via two 4 × 1 HD electrode arrays centered at F3 and P3. Three stimulation configurations, including in-phase, anti-phase, or sham, were tested on three different days in a cross-over (within-subject) design. On each test day, the subject underwent three experimental sessions: pre-, duringand post-stimulation sessions. In all sessions, the subject performed a Sternberg verbal working memory task with three levels of memory load (load 2, 4 and 6), imposing three levels of cognitive demand. Analyzing behavioral and EEG data from the post-stimulation session, we report two main observations. First, in-phase stimulation improved task performance in subjects with higher working memory capacity (WMC) under higher memory load (load 6). Second, in-phase stimulation enhanced frontoparietal theta synchrony during working memory retention in subjects with higher WMC under higher memory loads (load 4 and load 6), and the enhanced frontoparietal theta synchronization is mainly driven by enhanced frontal→parietal theta Granger causality. These observations suggest that (1) in-phase theta transcranial alternating current stimulation (tACS) during verbal working memory can result in observable behavioral and neurophysiological consequences post stimulation, (2) the short-term plasticity effects are state- and individual-dependent, and (3) enhanced executive control underlies improved behavioral performance ### Competing Interest Statement The authors have declared no competing interest.
Stefano Ioannucci; Guillermo Borragán; Alexandre Zénon
Passive visual stimulation induces fatigue under conditions of high arousal elicited by auditory tasks Journal Article
In: Journal of Experimental Psychology: General, vol. 151, no. 12, pp. 3097–3113, 2022.
Theories of cognitive fatigue disagree on whether performance decrement is caused by motivational or functional alterations. Here, drawing inspiration from the habituation and visual adaptation literature, we tested the assumption that keeping neural networks active for an extensive period of time entails consequences at the subjective and objective level—the defining characteristics of fatigue—when confounds such as motivation, boredom, and level of skill are controlled. In Experiment 1, we revealed that passive visual stimulation affected the performance of a subsequent task that was carried out in the same portion of visual space. While under conditions of low cognitive load and arousal, participants improved their performance in the stimulated quadrant; the reverse was observed under high arousal conditions. This latter performance decrement correlated also with the reported subjective level of fatigue and occurred while neural responses to the saturating stimulus remained constant, as assessed through steady-state EEG. In subsequent experiments, we replicated and further characterized this performance deterioration effect, revealing its specificity to the stimulated eye and stimulus orientation. Across the three experiments, the decrease in performance was correlated with pupil-linked arousal, suggesting its mediating effect in this phenomenon. In sum, we show that repeated stimulation of neural networks under high-arousal conditions leads to their altered functional performance, a mechanism which may play a role in the development of global cognitive fatigue
Gelu Ionescu; Aline Frey; Nathalie Guyader; Emmanuelle Kristensen; Anton Andreev; Anne Guérin-Dugué
Synchronization of acquisition devices in neuroimaging: An application using co-registration of eye movements and electroencephalography Journal Article
In: Behavior Research Methods, vol. 54, pp. 2545–2564, 2022.
Interest in applications for the simultaneous acquisition of data from different devices is growing. In neuroscience for example, co-registration complements and overcomes some of the shortcomings of individual methods. However, precise synchronization of the different data streams involved is required before joint data analysis. Our article presents and evaluates a synchronization method which maximizes the alignment of information across time. Synchronization through common triggers is widely used in all existing methods, because it is very simple and effective. However, this solution has been found to fail in certain practical situations, namely for the spurious detection of triggers and/or when the timestamps of triggers sampled by each acquisition device are not jointly distributed linearly for the entire duration of an experiment. We propose two additional mechanisms, the "Longest Common Subsequence" algorithm and a piecewise linear regression, in order to overcome the limitations of the classical method of synchronizing common triggers. The proposed synchronization method was evaluated using both real and artificial data. Co-registrations of electroencephalographic signals (EEG) and eye move- ments were used for real data. We compared the effectiveness of our method to another open source method implemented using EYE-EEG toolbox. Overall, we show that our method, implemented in C++ as a DOS application, is very fast, robust and fully automatic.
Shanice E. W. Janssens; Sanne Ten Oever; Alexander T. Sack; Tom A. Graaf
“Broadband alpha transcranial alternating current stimulation”: Exploring a new biologically calibrated brain stimulation protocol Journal Article
In: NeuroImage, vol. 253, pp. 1–12, 2022.
Transcranial alternating current stimulation (tACS) can be used to study causal contributions of oscillatory brain mechanisms to cognition and behavior. For instance, individual alpha frequency (IAF) tACS was reported to enhance alpha power and impact visuospatial attention performance. Unfortunately, such results have been inconsistent and difficult to replicate. In tACS, stimulation generally involves one frequency, sometimes individually calibrated to a peak value observed in an M/EEG power spectrum. Yet, the ‘peak' actually observed in such power spectra often contains a broader range of frequencies, raising the question whether a biologically calibrated tACS protocol containing this fuller range of alpha-band frequencies might be more effective. Here, we introduce ‘Broadband-alpha-tACS', a complex individually calibrated electrical stimulation protocol. We band-pass filtered left posterior resting-state EEG data around the IAF (± 2 Hz), and converted that time series into an electrical waveform for tACS stimulation of that same left posterior parietal cortex location. In other words, we stimulated a brain region with a ‘replay' of its own alpha-band frequency content, based on spontaneous activity. Within-subjects (N = 24), we compared to a sham tACS session the effects of broadband-alpha tACS, power-matched spectral inverse (‘alpha-removed') control tACS, and individual alpha frequency (IAF) tACS, on EEG alpha power and performance in an endogenous attention task previously reported to be affected by alpha tACS. Broadband-alpha-tACS significantly modulated attention task performance (i.e., reduced the rightward visuospatial attention bias in trials without distractors, and reduced attention benefits). Alpha-removed tACS also reduced the rightward visuospatial attention bias. IAF-tACS did not significantly modulate attention task performance compared to sham tACS, but also did not statistically significantly differ from broadband-alpha-tACS. This new broadband-alpha-tACS approach seems promising, but should be further explored and validated in future studies.
Jianrong Jia; Ying Fan; Huan Luo
Alpha-band phase modulates bottom-up feature processing Journal Article
In: Cerebral Cortex, vol. 32, no. 6, pp. 1–9, 2022.
Recent studies reveal that attention operates in a rhythmic manner, that is, sampling each location or feature alternatively over time. However, most evidence derives from top-down tasks, and it remains elusive whether bottom-up processing also entails dynamic coordination. Here, we developed a novel feature processing paradigm and combined time-resolved behavioral measurements and electroencephalogram (EEG) recordings to address the question. Specifically, a salient color in a multicolor display serves as a noninformative cue to capture attention and presumably reset the oscillations of feature processing. We then measured the behavioral performance of a probe stimulus associated with either high- or low-salient color at varied temporal lags after the cue. First, the behavioral results (i.e., reaction time) display an alpha-band ($sim$8 Hz) profile with a consistent phase lag between high- and low-salient conditions. Second, simultaneous EEG recordings show that behavioral performance is modulated by the phase of alpha-band neural oscillation at the onset of the probes. Finally, high- and low-salient probes are associated with distinct preferred phases of alpha-band neural oscillations. Taken together, our behavioral and neural results convergingly support a central function of alpha-band rhythms in feature processing, that is, features with varied saliency levels are processed at different phases of alpha neural oscillations.
Lu Jiang; Xiaoyang Li; Weihua Pei; Xiaorong Gao; Yijun Wang
A hybrid brain-computer interface based on visual evoked potential and pupillary response Journal Article
In: Frontiers in Human Neuroscience, vol. 16, pp. 1–16, 2022.
Brain-computer interface (BCI) based on steady-state visual evoked potential (SSVEP) has been widely studied due to the high information transfer rate (ITR), little user training, and wide subject applicability. However, there are also disadvantages such as visual discomfort and “BCI illiteracy.” To address these problems, this study proposes to use low-frequency stimulations (12 classes, 0.8–2.12 Hz with an interval of 0.12 Hz), which can simultaneously elicit visual evoked potential (VEP) and pupillary response (PR) to construct a hybrid BCI (h-BCI) system. Classification accuracy was calculated using supervised and unsupervised methods, respectively, and the hybrid accuracy was obtained using a decision fusion method to combine the information of VEP and PR. Online experimental results from 10 subjects showed that the averaged accuracy was 94.90 ± 2.34% (data length 1.5 s) for the supervised method and 91.88 ± 3.68% (data length 4 s) for the unsupervised method, which correspond to the ITR of 64.35 ± 3.07 bits/min (bpm) and 33.19 ± 2.38 bpm, respectively. Notably, the hybrid method achieved higher accuracy and ITR than that of VEP and PR for most subjects, especially for the short data length. Together with the subjects' feedback on user experience, these results indicate that the proposed h-BCI with the low-frequency stimulation paradigm is more comfortable and favorable than the traditional SSVEP-BCI paradigm using the alpha frequency range.
Richard Johnston; Adam C. Snyder; Rachel S. Schibler; Matthew A. Smith
EEG signals index a global signature of arousal embedded in neuronal population recordings Journal Article
In: eNeuro, vol. 9, no. 3, pp. 1–16, 2022.
Electroencephalography (EEG) has long been used to index brain states, from early studies describing activity in the presence and absence of visual stimulation to modern work employing complex perceptual tasks. These studies have shed light on brain-wide signals but often lack explanatory power at the single neuron level. Similarly, single neuron recordings can suffer from an inability to measure brain-wide signals accessible using EEG. Here, we combined these techniques while monkeys performed a change detection task and discovered a novel link between spontaneous EEG activity and a neural signal embedded in the spiking responses of neuronal populations. This “slow drift” was associated with fluctuations in the subjects' arousal levels over time: decreases in prestimulus a power were accompanied by increases in pupil size and decreases in microsaccade rate. These re- sults show that brain-wide EEG signals can be used to index modes of activity present in single neuron recordings, that in turn reflect global changes in brain state that influence perception and behavior.
Oren Kadosh; Yoram Bonneh
Face familiarity revealed by fixational eye movements and fixation-related potentials in free viewing Journal Article
In: Scientific Reports, vol. 12, no. 1, pp. 1–13, 2022.
Event-related potentials (ERPs) and the oculomotor inhibition (OMI) in response to visual transients are known to be sensitive to stimulus properties, attention, and expectation. We have recently found that the OMI is also sensitive to face familiarity. In natural vision, stimulation of the visual cortex is generated primarily by saccades, and it has been recently suggested that fixation-related potentials (FRPs) share similar components with the ERPs. Here, we investigated whether FRPs and microsaccade inhibition (OMI) in free viewing are sensitive to face familiarity. Observers freely watched a slideshow of seven unfamiliar and one familiar facial images presented randomly for 4-s periods, with multiple images per identity. We measured the occipital fixation-related N1 relative to the P1 magnitude as well as the associated fixation-triggered OMI. We found that the average N1-P1 was significantly smaller and the OMI was shorter for the familiar face, compared with any of the seven unfamiliar faces. Moreover, the P1 was suppressed across saccades for the familiar but not for the unfamiliar faces. Our results highlight the sensitivity of the occipital FRPs to stimulus properties such as face familiarity and advance our understanding of the integration process across successive saccades in natural vision.
Sharif I. Kronemer; Mark Aksen; Julia Z. Ding; Jun Hwan Ryu; Qilong Xin; Zhaoxiong Ding; Jacob S. Prince; Hunki Kwon; Aya Khalaf; Sarit Forman; David S. Jin; Kevin Wang; Kaylie Chen; Claire Hu; Akshar Agarwal; Erik Saberski; Syed Mohammad Adil Wafa; Owen P. Morgan; Jia Wu; Kate L. Christison-Lagay; Nicholas Hasulak; Martha Morrell; Alexandra Urban; R. Todd Constable; Michael Pitts; R. Mark Richardson; Michael J. Crowley; Hal Blumenfeld
Human visual consciousness involves large scale cortical and subcortical networks independent of task report and eye movement activity Journal Article
In: Nature Communications, vol. 13, pp. 1–17, 2022.
The full neural circuits of conscious perception remain unknown. Using a visual perception task, we directly recorded a subcortical thalamic awareness potential (TAP). We also developed a unique paradigm to classify perceived versus not perceived stimuli using eye measurements to remove confounding signals related to reporting on conscious experiences. Using fMRI, we discovered three major brain networks driving conscious visual perception independent of report: first, increases in signal detection regions in visual, fusiform cortex, and frontal eye fields; and in arousal/salience networks involving midbrain, thalamus, nucleus accumbens, anterior cingulate, and anterior insula; second, increases in frontoparietal attention and executive control networks and in the cerebellum; finally, decreases in the default mode network. These results were largely maintained after excluding eye movement-based fMRI changes. Our findings provide evidence that the neurophysiology of consciousness is complex even without overt report, involving multiple cortical and subcortical networks overlapping in space and time.
Louisa Kulke; Lena Brümmer; Arezoo Pooresmaeili; Annekathrin Schacht
Visual competition attenuates emotion effects during overt attention shifts Journal Article
In: Psychophysiology, vol. 59, pp. 1–14, 2022.
Numerous different objects are simultaneously visible in a person's visual field, competing for attention. This competition has been shown to affect eye-movements and early neural responses toward stimuli, while the role of a stimulus' emotional meaning for mechanisms of overt attention shifts under competition is unclear. The current study combined EEG and eye-tracking to investigate effects of competition and emotional content on overt shifts of attention to human face stimuli. Competition prolonged the latency of the P1 component and of saccades, while faces showing emotional expressions elicited an early posterior negativity (EPN). Remarkably, the emotion-related modulation of the EPN was attenuated when two stimuli were competing for attention compared to non-competition. In contrast, no interaction effects of emotional expression and competition were observed on other event-related potentials. This finding indicates that competition can decelerate attention shifts in general and also diminish the emotion-driven attention capture, measured through the smaller effects of emotional expression on EPN amplitude. Reduction of the brain's responsiveness to emotional content in the presence of distractors contradicts models that postulate fully automatic processing of emotions.
Wupadrasta Santosh Kumar; Keerthana Manikandan; Dinavahi V. P. S. Murty; Ranjini Garani Ramesh; Simran Purokayastha; Mahendra Javali; Naren Prahalada Rao; Supratim Ray
Stimulus-induced narrowband gamma oscillations are test–retest reliable in human EEG Journal Article
In: Cerebral Cortex Communications, vol. 3, no. 1, pp. 1–15, 2022.
Visual stimulus-induced gamma oscillations in electroencephalogram (EEG) recordings have been recently shown to be compromised in subjects with preclinical Alzheimer's Disease (AD), suggesting that gamma could be an inexpensive biomarker for AD diagnosis provided its characteristics remain consistent across multiple recordings. Previous magnetoencephalography studies in young subjects have reported consistent gamma power over recordings separated by a few weeks to months. Here, we assessed the consistency of stimulus-induced slow (20–35 Hz) and fast gamma (36–66 Hz) oscillations in subjects (n = 40) (age: 50–88 years) in EEG recordings separated by a year, and tested the consistency in the magnitude of gamma power, its temporal evolution and spectral profile. Gamma had distinct spectral/temporal characteristics across subjects, which remained consistent across recordings (average intraclass correlation of $sim$0.7). Alpha (8–12 Hz) and steady-state-visually evoked-potentials were also reliable. We further tested how EEG features can be used to identify 2 recordings as belonging to the same versus different subjects and found high classifier performance (AUC of $sim$0.89), with temporal evolution of slow gamma and spectral profile being most informative. These results suggest that EEG gamma oscillations are reliable across sessions separated over long durations and can also be a potential tool for subject identification.
Ilmari Kurki; Aapo Hyvärinen; Linda Henriksson
Dynamics of retinotopic spatial attention revealed by multifocal MEG Journal Article
In: NeuroImage, vol. 263, pp. 1–13, 2022.
Visual focal attention is both fast and spatially localized, making it challenging to investigate using human neuroimaging paradigms. Here, we used a new multivariate multifocal mapping method with magnetoencephalography (MEG) to study how focal attention in visual space changes stimulus-evoked responses across the visual field. The observer's task was to detect a color change in the target location, or at the central fixation. Simultaneously, 24 regions in visual space were stimulated in parallel using an orthogonal, multifocal mapping stimulus sequence. First, we used univariate analysis to estimate stimulus-evoked responses in each channel. Then we applied multivariate pattern analysis to look for attentional effects on the responses. We found that attention to a target location causes two spatially and temporally separate effects. Initially, attentional modulation is brief, observed at around 60–130 ms post stimulus, and modulates responses not only at the target location but also in adjacent regions. A later modulation was observed from around 200 ms, which was specific to the location of the attentional target. The results support the idea that focal attention employs several processing stages and suggest that early attentional modulation is less spatially specific than late.
Timo L. Kvamme; Mesud Sarmanlu; Christopher Bailey; Morten Overgaard
Neurofeedback modulation of the sound-induced flash illusion using parietal cortex alpha oscillations reveals dependency on prior multisensory congruency Journal Article
In: Neuroscience, vol. 482, pp. 1–17, 2022.
Spontaneous neural oscillations are key predictors of perceptual decisions to bind multisensory sig- nals into a unified percept. Research links decreased alpha power in the posterior cortices to attention and audio- visual binding in the sound-induced flash illusion (SIFI) paradigm. This suggests that controlling alpha oscillations would be a way of controlling audiovisual binding. In the present feasibility study we used MEG- neurofeedback to train one group of subjects to increase left/right and another to increase right/left alpha power ratios in the parietal cortex. We tested for changes in audiovisual binding in a SIFI paradigm where flashes appeared in both hemifields. Results showed that the neurofeedback induced a significant asymmetry in alpha power for the left/right group, not seen for the right/left group. Corresponding asymmetry changes in audiovisual binding in illusion trials (with 2, 3, and 4 beeps paired with 1 flash) were not apparent. Exploratory analyses showed that neurofeedback training effects were present for illusion trials with the lowest numeric disparity (i.e., 2 beeps and 1 flash trials) only if the previous trial had high congruency (2 beeps and 2 flashes). Our data suggest that the relation between parietal alpha power (an index of attention) and its effect on audiovisual binding is dependent on the learned causal structure in the previous stimulus. The present results suggests that low alpha power biases observers towards audiovisual binding when they have learned that audiovisual signals orig- inate from a common origin, consistent with a Bayesian causal inference account of multisensory perception.
Timo L. Kvamme; Mesud Sarmanlu; Morten Overgaard
Doubting the double-blind: Introducing a questionnaire for awareness of experimental purposes in neurofeedback studies Journal Article
In: Consciousness and Cognition, vol. 104, pp. 1–13, 2022.
Double-blinding subjects to the experiment's purpose is an important standard in neurofeedback studies. However, it is difficult to provide evidence that humans are entirely unaware of certain information. This study used insights from consciousness studies and neurophenomenology to develop a contingency awareness questionnaire for neurofeedback. We assessed whether participants had an awareness of experimental purposes to manipulate their attention and multisensory perception. A subset of subjects (5 out of 20) gained a degree of awareness of experimental purposes as evidenced by their correct guess about the purposes of the experiment to affect their attention and multisensory perceptions specific to their double-blinded group assignment. The results warrant replication before they are applied to clinical neurofeedback studies, given the considerable time taken to perform the questionnaire (∼25 min). We discuss the strengths and limitations of our contingency awareness questionnaire and the growing appeal of the double-blinded standard in clinical neurofeedback studies.
Nan Li; Olaf Dimigen; Werner Sommer; Suiping Wang
Parafoveal words can modulate sentence meaning: Electrophysiological evidence from an RSVP-with-flanker task Journal Article
In: Psychophysiology, vol. 59, pp. 1–18, 2022.
During natural reading, readers can take up some visual information from not-yet-fixated words to the right of the current fixation and it is well-established that this parafoveal preview facilitates the subsequent foveal processing of the word. However, the extraction and integration of word meaning from parafoveal words and their possible influence on the semantic content of the sentence are controversial. In the current study, we recorded event-related potentials (ERPs) in the RSVP-with-flankers paradigm to test whether and how updates of sentential meaning, based only on parafoveal information, may influence the subsequent foveal processing. In Chinese sentences, the congruency of parafoveal and foveal target words with the sentence was orthogonally manipulated. In contrast to previous research, we also controlled for potentially confounding effects of parafoveal-to-foveal repetition priming (identity preview effects) on the N400. Crucially, we found that the classic effect of foveal congruency on the N400 component only appeared when the word in preview had been congruent with sentence meaning; in contrast, there was no N400 as a function of foveal incongruency when the preview word had also been incongruent. These results indicate that sentence meaning rapidly adapts to parafoveal preview, altering the semantic context for the subsequently fixated word. We also show that correct parafoveal preview generally attenuates the N400 once a word is fixated, regardless of congruency. Taken together, our findings underline the highly generative and adaptive framework of language comprehension.
Baiwei Liu; Anna C. Nobre; Freek Ede
Functional but not obligatory link between microsaccades and neural modulation by covert spatial attention Journal Article
In: Nature Communications, vol. 13, pp. 1–10, 2022.
Covert spatial attention is associated with spatial modulation of neural activity as well as with directional biases in fixational eye movements known as microsaccades. We studied how these two ‘fingerprints' of attention are interrelated in humans. We investigated spatial modulation of 8-12 Hz EEG alpha activity and microsaccades when attention is directed internally within the spatial layout of visual working memory. Consistent with a common origin, spatial modulations of alpha activity and microsaccades co-vary: alpha lateralisation is stronger in trials with microsaccades toward versus away from the memorised location of the to-be-attended item and occurs earlier in trials with earlier microsaccades toward this item. Critically, however, trials without attention-driven microsaccades nevertheless show clear spatial modulation of alpha activity – comparable to trials with attention-driven microsaccades. Thus, directional biases in microsaccades correlate with neural signatures of spatial attention, but they are not necessary for neural modulation by spatial attention to be manifest.
Sasu Mäkelä; Jan Kujala; Riitta Salmelin
Removing ocular artifacts from magnetoencephalographic data on naturalistic reading of continuous texts Journal Article
In: Frontiers in Neuroscience, vol. 16, pp. 1–18, 2022.
Naturalistic reading paradigms and stimuli consisting of long continuous texts are essential for characterizing the cortical basis of reading. Due to the highly dynamic nature of the reading process, electrophysiological brain imaging methods with high spatial and temporal resolution, such as magnetoencephalography (MEG), are ideal for tracking them. However, as electrophysiological recordings are sensitive to electromagnetic artifacts, data recorded during naturalistic reading is confounded by ocular artifacts. In this study, we evaluate two different pipelines for removing ocular artifacts from MEG data collected during continuous, naturalistic reading, with the focus on saccades and blinks. Both pipeline alternatives are based on blind source separation methods but differ fundamentally in their approach. The first alternative is a multi-part process, in which saccades are first extracted by applying Second-Order Blind Identification (SOBI) and, subsequently, FastICA is used to extract blinks. The other alternative uses a single powerful method, Adaptive Mixture ICA (AMICA), to remove all artifact types at once. The pipelines were tested, and their effects compared on MEG data recorded from 13 subjects in a naturalistic reading task where the subjects read texts with the length of multiple pages. Both pipelines performed well, extracting the artifacts in a single component per artifact type in most subjects. Signal power was reduced across the whole cortex in all studied frequency bands from 1 to 90 Hz, but especially in the frontal cortex and temporal pole. The results were largely similar for the two pipelines, with the exception that SOBI-FastICA reduced signal in the right frontal cortex in all studied frequency bands more than AMICA. However, there was considerable interindividual variation in the effects of the pipelines. As a holistic conclusion, we choose to recommend AMICA for removing artifacts from MEG data on naturalistic reading but note that the SOBI-FastICA pipeline has also various favorable characteristics.
Soukayna Bekkali; George J. Youssef; Peter H. Donaldson; Jason He; Michael Do; Christian Hyde; Pamela Barhoun; Peter G. Enticott
Do gaze behaviours during action observation predict interpersonal motor resonance? Journal Article
In: Social Cognitive and Affective Neuroscience, vol. 17, no. 1, pp. 61–71, 2022.
Interpersonal motor resonance (IMR) is a common putative index of the mirror neuron system (MNS), a network containing specialised cells that fire during both action execution and observation. Visual content inputs to the MNS, however, it is unclear whether visual behaviours mediate the putative MNS response. We aimed to examine gaze effects on IMR during action observation. Neurotypical adults (N = 99; 60 female) underwent transcranial magnetic stimulation, electromyography, and eye-tracking during the observation of videos of actors performing grasping actions. IMR was measured as a percentage change in motor evoked potentials (MEPs) of the first dorsal interosseous muscle during action observation relative to baseline. MEP facilitation was observed during action observation, indicating IMR (65.43%
Lorenzo Diana; Giulia Scotti; Edoardo N. Aiello; Patrick Pilastro; Aleksandra K. Eberhard-Moscicka; Ren M. Müri; Nadia Bolognini
Conventional and HD-tDCS may (or may not) modulate overt attentional orienting: An integrated spatio-temporal approach and methodological reflection Journal Article
In: Brain Sciences, vol. 12, pp. 1–20, 2022.
Transcranial Direct Current Stimulation (tDCS) has been employed to modulate visuo- spatial attentional asymmetries, however, further investigation is needed to characterize tDCS- associated variability in more ecological settings. In the present research, we tested the effects of offline, anodal conventional tDCS (Experiment 1) and HD-tDCS (Experiment 2) delivered over the posterior parietal cortex (PPC) and Frontal Eye Field (FEF) of the right hemisphere in healthy participants. Attentional asymmetries were measured by means of an eye tracking-based, ecological paradigm, that is, a Free Visual Exploration task of naturalistic pictures. Data were analyzed from a spatiotemporal perspective. In Experiment 1, a pre-post linear mixed model (LMM) indicated a leftward attentional shift after PPC tDCS; this effect was not confirmed when the individual baseline performance was considered. In Experiment 2, FEF HD-tDCS was shown to induce a significant leftward shift of gaze position, which emerged after 6 s of picture exploration and lasted for 200 ms. The present results do not allow us to conclude on a clear efficacy of offline conventional tDCS and HD- tDCS in modulating overt visuospatial attention in an ecological setting. Nonetheless, our findings highlight a complex relationship among stimulated area, focality of stimulation, spatiotemporal aspects of deployment of attention, and the role of individual baseline performance in shaping the effects of tDCS.
Stefano Gallotto; Teresa Schuhmann; Felix Duecker; Marij Middag-van Spanje; Tom A. Graaf; Alexander T. Sack
Concurrent frontal and parietal network TMS for modulating attention Journal Article
In: iScience, vol. 25, pp. 1–14, 2022.
Transcranial magnetic stimulation (TMS) has been applied to frontal eye field (FEF) and intraparietal sulcus (IPS) in isolation, to study their role in attention. However, these nodes closely interact in a “dorsal attention network”. Here, we compared effects of inhibitory TMS applied to individually fMRI-localized FEF or IPS (single-node TMS), to effects of simultaneously inhibiting both regions (“network TMS”), and sham. We assessed attention performance using the lateralized attention network test, which captures multiple facets of attention: spatial orienting, alerting, and executive control. TMS showed no effects on alerting and executive control. For spatial orienting, only network TMS showed a reduction of the orienting effect in the right hemifield compared to the left hemifield, irrespective of the order of TMS application (IPS→FEF or FEF→IPS). Network TMS might prevent compensatory mechanisms within a brain network, which is promising for both research and clinical applications to achieve superior neuromodulation effects.
Adam M. McNeill; Rebecca L. Monk; Adam W. Qureshi; Stergios Makris; Valentina Cazzato; Derek Heim
Elevated ad libitum alcohol consumption following continuous theta burst stimulation to the left-dorsolateral prefrontal cortex is partially mediated by changes in craving Journal Article
In: Cognitive, Affective and Behavioral Neuroscience, vol. 22, pp. 160–170, 2022.
Previous research indicates that following alcohol intoxication, activity in prefrontal cortices is reduced, linking to changes in associated cognitive processes, such as inhibitory control, attentional bias (AB), and craving. While these changes have been implicated in alcohol consumption behaviour, it has yet to be fully illuminated how these frontal regions and cognitive processes interact to govern alcohol consumption behaviour. The current preregistered study applied continuous theta burst transcranial magnetic stimulation (cTBS) to examine directly these relationships while removing the wider pharmacological effects of alcohol. A mixed design was implemented, with cTBS stimulation to right and left dorsolateral prefrontal cortex (DLPFC), the medial orbital frontal cortex (mOFC) and Vertex, with measures of inhibitory control, AB, and craving taken both pre- and post-stimulation. Ad libitum consumption was measured using a bogus taste task. Results suggest that rDLPFC stimulation impaired inhibitory control but did not significantly increase ad libitum consumption. However, lDLPFC stimulation heightened craving and increased consumption, with findings indicating that changes in craving partially mediated the relationship between cTBS stimulation ofprefrontal regions and ad libitum consumption. Medial OFC stimulation and AB findings were inconclusive. Overall, results implicate the left DLPFC in the regulation of craving, which appears to be a prepotent cognitive mechanism by which alcohol consumption is driven and maintained.
Paola Mengotti; Anne Sophie Käsbauer; Gereon R. Fink; Simone Vossel
Combined TMS-fMRI reveals behavior-dependent network effects of right temporoparietal junction neurostimulation in an attentional belief updating task Journal Article
In: Cerebral Cortex, vol. 32, pp. 4698–4714, 2022.
Updating beliefs after unexpected events is fundamental for an optimal adaptation to the environment. Previous findings suggested a causal involvement of the right temporoparietal junction (rTPJ) in belief updating in an attention task. We combined offline continuous theta-burst stimulation (cTBS) over rTPJ with functional magnetic resonance imaging (fMRI) to investigate local and remote stimulation effects within the attention and salience networks. In a sham-controlled, within-subject crossover design, 25 participants performed an attentional cueing task during fMRI with true or false information about cue predictability. By estimating learning rates from response times, we characterized participants' belief updating. Model-derived cue predictability entered the fMRI analysis as a parametric regressor to identify the neural correlates of updating. rTPJ-cTBS effects showed high interindividual variability. The expected learning rate reduction with false cue predictability information by cTBS was only observed in participants showing higher updating in false than in true blocks after sham. cTBS modulated the neural signatures of belief updating, both in rTPJ and in nodes of the attention and salience networks. The interindividual variability of the behavioral cTBS effect was related to differential activity and rTPJ connectivity of the right anterior insula. These results demonstrate a crucial interaction between ventral attention and salience networks for belief updating.
Marij Middag-van Spanje; Felix Duecker; Stefano Gallotto; Tom A. Graaf; Caroline Heugten; Alexander T. Sack; Teresa Schuhmann
Transcranial magnetic stimulation over posterior parietal cortex modulates alerting and executive control processes in attention Journal Article
In: European Journal of Neuroscience, vol. 56, pp. 5853–5868, 2022.
Attention includes three different functional components: generating and maintaining an alert state (alerting), orienting to sensory events (orienting), and resolving conflicts between alternative actions (executive control). Neuroimaging and patient studies suggest that the posterior parietal cortex (PPC) is involved in all three attention components. Transcranial magnetic stimulation (TMS) has repeatedly been applied over the PPC to study its functional role for shifts and maintenance of visuospatial attention. Most TMS-PPC studies used only detection tasks or orienting paradigms to investigate TMS-PPC effects on attention processes, neglecting the alerting and executive control components of attention. The objective of the present study was to investigate the role of PPC in all three functional components of attention: alerting, orienting, and executive control. To this end, we disrupted PPC with TMS (continuous theta-burst stimulation), to modulate subsequent performance on the Lateralized-Attention Network Test, used to assess the three attention components separately. Our results revealed hemifield-specific effects on alerting and executive control functions, but we did not find stimulation effects on orienting performance. While this field of research and associated clinical development have been predominantly focused on orienting performance, our results suggest that parietal cortex and its modulation may affect other aspects of attention as well.
Johannes Rennig; Michael S. Beauchamp
Intelligibility of audiovisual sentences drives multivoxel response patterns in human superior temporal cortex Journal Article
In: NeuroImage, vol. 247, pp. 1–9, 2022.
Regions of the human posterior superior temporal gyrus and sulcus (pSTG/S) respond to the visual mouth movements that constitute visual speech and the auditory vocalizations that constitute auditory speech, and neural responses in pSTG/S may underlie the perceptual benefit of visual speech for the comprehension of noisy auditory speech. We examined this possibility through the lens of multivoxel pattern responses in pSTG/S. BOLD fMRI data was collected from 22 participants presented with speech consisting of English sentences presented in five different formats: visual-only; auditory with and without added auditory noise; and audiovisual with and without auditory noise. Participants reported the intelligibility of each sentence with a button press and trials were sorted post-hoc into those that were more or less intelligible. Response patterns were measured in regions of the pSTG/S identified with an independent localizer. Noisy audiovisual sentences with very similar physical properties evoked very different response patterns depending on their intelligibility. When a noisy audiovisual sentence was reported as intelligible, the pattern was nearly identical to that elicited by clear audiovisual sentences. In contrast, an unintelligible noisy audiovisual sentence evoked a pattern like that of visual-only sentences. This effect was less pronounced for noisy auditory-only sentences, which evoked similar response patterns regardless of intelligibility. The successful integration of visual and auditory speech produces a characteristic neural signature in pSTG/S, highlighting the importance of this region in generating the perceptual benefit of visual speech.
Andreas Alexandersen; Gábor Csifcsák; Josephine Groot; Matthias Mittner
The effect of transcranial direct current stimulation on the interplay between executive control, behavioral variability and mind wandering: A registered report Journal Article
In: Neuroimage: Reports, vol. 2, no. 3, pp. 1–19, 2022.
Mind wandering (MW) is a mental phenomenon humans experience daily. Yet, we lack a complete understanding of the neural basis of this pervasive mental state. Over the past decade there has been an increase in publications using transcranial direct current stimulation (tDCS) to modulate the propensity to mind wander, but findings are diverse, and a satisfactory conclusion is missing. Recently, Boayue et al. (2020) reported successful reduction of mind wandering using high-definition tDCS (HD-tDCS) over the dorsolateral prefrontal cortex, providing preliminary evidence for the efficacy of HD-tDCS in interfering with mind wandering. The current study is a high-powered, pre-registered direct replication attempt of the effect found by Boayue et al. (2020). In addition, we investigated whether the effects of HD-tDCS on mind wandering would be prolonged and assessed the underlying processes of mind wandering using electroencephalography (EEG) and pupillometry during a finger-tapping random sequence generation task that requires the use of executive resources. We failed to find any evidence of the original effect of reduced MW during and after stimulation. When combining our data with the data from Boayue et al. (2020), the original effect of reduced MW caused by HD-tDCS disappeared. In addition, we observed increased occipital alpha power as task duration increased and increased midfrontal theta power preceding response patterns signaling high executive function use. Finally, tonic and phasic pupil size decreased as task duration increased yet, phasic responses were increased, while tonic responses were reduced preceding reports of MW. Additionally phasic pupil size also showed a tendency to be increased during periods of high executive function use. Importantly, none of the EEG or pupil measures were modulated by HD-tDCS. We conclude that HD-tDCS over the dorsolateral prefrontal cortex does not affect MW propensity and its neural signatures. Furthermore, we recommend that previously reported effects of tDCS on mind wandering and other cognitive functions should only be accepted after a successful pre-registered replication.
Kristijan Armeni; Umut Güçlü; Marcel Gerven; Jan-Mathijs Schoffelen
A 10-hour within-participant magnetoencephalography narrative dataset to test models of language comprehension Journal Article
In: Scientific Data, vol. 9, pp. 1–18, 2022.
Recently, cognitive neuroscientists have increasingly studied the brain responses to narratives. At the same time, we are witnessing exciting developments in natural language processing where large-scale neural network models can be used to instantiate cognitive hypotheses in narrative processing. Yet, they learn from text alone and we lack ways of incorporating biological constraints during training. To mitigate this gap, we provide a narrative comprehension magnetoencephalography (MEG) data resource that can be used to train neural network models directly on brain data. We recorded from 3 participants, 10 separate recording hour-long sessions each, while they listened to audiobooks in English. After story listening, participants answered short questions about their experience. To minimize head movement, the participants wore MEG-compatible head casts, which immobilized their head position during recording. We report a basic evoked-response analysis showing that the responses accurately localize to primary auditory areas. The responses are robust and conserved across 10 sessions for every participant. We also provide usage notes and briefly outline possible future uses of the resource.
Alessandro Benedetto; Hao Tam Ho; Maria Concetta Morrone
The readiness potential correlates with action-linked modulation of visual accuracy Journal Article
In: Eneuro, pp. 1–17, 2022.
Visual accuracy is consistently shown to be modulated around the time of the action execution. The neural underpinning of this motor-induced modulation of visual perception is still unclear. Here, we investigate with EEG whether it is related to the readiness potential, an event-related potential linked to motor preparation. Across 18 human participants, the magnitude of visual modulation following a voluntary button press was found to correlate with the readiness potential amplitude measured during visual discrimination. Participants' amplitude of the readiness potential in a purely motor-task was also found to correlate with the extent of the motor-induced modulation of visual perception in the visuomotor task. These results provide strong evidence that perceptual changes close to action execution are associated with motor preparation processes and that this mechanism is independent of task contingencies. Further, our findings suggest that the readiness potential provides a fingerprint of individual visuomotor interaction.
Sebastian Schindler; Niko Busch; Maximilian Bruchmann; Maren-Isabel Wolf; Thomas Straube
Early ERP functions are indexed by lateralized effects to peripherally presented emotional faces and scrambles Journal Article
In: Psychophysiology, vol. 59, no. 2, pp. 1–16, 2022.
A large body of research suggests that early event-related potentials (ERPs), such as the P1 and N1, are potentiated by attention and represent stimulus amplification. However, recent accounts suggest that the P1 is associated with inhibiting the irrelevant visual field evidenced by a pronounced ipsilateral P1 during sustained attention to peripherally presented stimuli. The current EEG study further investigated this issue to reveal how lateralized ERP findings are modulated by face and emotional information. Therefore, participants were asked to fixate the center of the screen and pay sustained attention either to the right or left visual field, where angry or neutral faces or their Fourier phase-scrambled versions were presented. We found a bilateral P1 to all stimuli with relatively increased, but delayed, ipsilateral P1 amplitudes to faces but not to scrambles. Explorative independent component analyses dissociated an earlier lateralized larger contralateral P1 from a later bilateral P1. By contrast, the N170 showed a contralateral enhancement to all stimuli, which was most pronounced for neutral faces attended in the left hemifield. Finally, increased contralateral alpha power was found for both attended hemifields but was not significantly related to poststimulus ERPs. These results provide evidence against a general inhibitory role of the P1 but suggest stimulus-specific relative enhancements of the ipsilateral P1 for the irrelevant visual hemifield. The lateralized N170, however, is associated with stimulus amplification as a function of facial features.
Sebastian Schindler; Theresa Sofie Richter; Maximilian Bruchmann; Niko A. Busch; Thomas Straube
Effects of task load, spatial attention, and trait anxiety on neuronal responses to fearful and neutral faces Journal Article
In: Psychophysiology, vol. 59, pp. 1–14, 2022.
There is an ongoing debate on how different components of the event-related potential (ERP) to threat-related facial expressions are modulated by attentional conditions and interindividual differences in trait anxiety. In the current study (N = 80), we examined ERPs to centrally presented, task-irrelevant fearful and neutral faces, while participants had to solve a face-unrelated visual task, which differed in difficulty and spatial position. Critically, we used a fixation-controlled experimental design and ensured the spatial attention manipulation by spectral analysis of the EEG. Besides the factors emotion, spatial attention, and perceptual load, we also investigated correlations between trait anxiety and ERPs. While P1 emotion effects were insignificant, the N170 was increased to fearful faces regardless of load and spatial attention conditions. During the EPN time window, a significantly increased negativity for fearful faces was observed only during low load and spatial attention to the face. We found no significant relationship between ERPs and trait anxiety, questioning the hypothesis of a general hypersensitivity toward fearful expressions in anxious individuals. These results show a high resistance of the N170 amplitude increase for fearful faces to spatial attention and task load manipulations. By contrast, the EPN modulation by fearful faces index a resource-dependent stage of the ERP, requiring both spatial attention at the location of faces and low load of the face-irrelevant task.
Elena Selezneva; Nicole Wetzel
The impact of probabilistic cues on sound-related pupil dilation and ERP responses in 7–9-year-old children Journal Article
In: Auditory Perception & Cognition, vol. 5, no. 1-2, pp. 86–106, 2022.
Control of involuntary orienting of attention toward new but task-irrelevant events is essential to successfully perform a task. We investigated top-down control of involuntary orienting of attention caused by task-irrelevant novel sounds embedded in a sequence of repeated standard sounds in 7–9-year-old children (N = 30) and in an adult control group (N = 30). The type of sound was announced by visual cues, which were correct in 80% of the trials. We co-registered sound-related pupil dilation responses (PDR), the attention-related component P3a in the EEG and performance. Task-irrelevant novel sounds evoked increased amplitudes of the PDR and the P3a and prolonged reaction times in both age groups. In children only, invalidly cued novel sounds evoked larger PDR amplitudes than validly cued novel sounds, while this cue effect was not observed for standard sounds. In both age groups, P3a amplitudes in the centro-parietal region were reduced to the correctly cued compared to the incorrectly cued novel sounds, indicating top-down control of orienting of attention. The reaction time prolongation to both validly and invalidly cued novel sounds were similar in both age groups. These findings demonstrate that children are capable of reducing the orienting of attention and evaluation triggered by task-irrelevant sounds by using probabilistic cues. Children's pupil results indicate a high sensitivity of pupil dynamics to cue-related top-down influences on novel sound processing, emphasizing the utility of pupillometry in developmental research.
Benjamin J. Stauch; Alina Peter; Isabelle Ehrlich; Zora Nolte; Pascal Fries
Human visual gamma for color stimuli Journal Article
In: eLife, vol. 11, pp. 1–18, 2022.
Strong gamma-band oscillations in primate early visual cortex can be induced by homogeneous color surfaces (Peter et al., 2019; Shirhatti and Ray, 2018). Compared to other hues, particularly strong gamma oscillations have been reported for red stimuli. However, precortical color processing and the resultant strength of input to V1 have often not been fully controlled for. Therefore, stronger responses to red might be due to differences in V1 input strength. We presented stimuli that had equal luminance and cone contrast levels in a color coordinate system based on responses of the lateral geniculate nucleus, the main input source for area V1. With these stimuli, we recorded magnetoencephalography in 30 human participants. We found gamma oscillations in early visual cortex which, contrary to previous reports, did not differ between red and green stimuli of equal L-M cone contrast. Notably, blue stimuli with contrast exclusively on the S-cone axis induced very weak gamma responses, as well as smaller event-related fields and poorer change-detection performance. The strength of human color gamma responses for stimuli on the L-M axis could be well explained by L-M cone contrast and did not show a clear red bias when L-M cone contrast was properly equalized.
Anna Lena Stroh; Konstantin Grin; Frank Rösler; Davide Bottari; José Ossandón; Bruno Rossion; Brigitte Röder
Developmental experiences alter the temporal processing characteristics of the visual cortex: Evidence from deaf and hearing native signers Journal Article
In: European Journal of Neuroscience, vol. 55, no. 6, pp. 1629–1644, 2022.
To date, the extent to which early experience shapes the functional characteristics of neural circuits is still a matter of debate. In the present study, we tested whether congenital deafness and/or the acquisition of a sign language alter the temporal processing characteristics of the visual system. Moreover, we investigated whether, assuming cross-modal plasticity in deaf individuals, the temporal processing characteristics of possibly reorganised auditory areas resemble those of the visual cortex. Steady-state visual evoked potentials (SSVEPs) were recorded in congenitally deaf native signers, hearing native signers, and hearing nonsigners. The luminance of the visual stimuli was periodically modulated at 12, 21, and 40 Hz. For hearing nonsigners, the optimal driving rate was 12 Hz. By contrast, for the group of hearing signers, the optimal driving rate was 12 and 21 Hz, whereas for the group of deaf signers, the optimal driving rate was 21 Hz. We did not observe evidence for cross-modal recruitment of auditory cortex in the group of deaf signers. These results suggest a higher preferred neural processing rate as a consequence of the acquisition of a sign language.
Lina Teichmann; Denise Moerel; Anina N. Rich; Chris I. Baker
The nature of neural object representations during dynamic occlusion Journal Article
In: Cortex, vol. 153, pp. 66–86, 2022.
Objects disappearing briefly from sight due to occlusion is an inevitable occurrence in everyday life. Yet we generally have a strong experience that occluded objects continue to exist, despite the fact that they objectively disappear. This indicates that neural object representations must be maintained during dynamic occlusion. However, it is unclear what the nature of such representation is and in particular whether it is perception-like or more abstract, for example, reflecting limited features such as position or movement direction only. In this study, we address this question by examining how different object features such as object shape, luminance, and position are represented in the brain when a moving object is dynamically occluded. We apply multivariate decoding methods to Magnetoencephalography (MEG) data to track how object representations unfold over time. Our methods allow us to contrast the representations of multiple object features during occlusion and enable us to compare the neural responses evoked by visible and occluded objects. The results show that object position information is represented during occlusion to a limited extent while object identity features are not maintained through the period of occlusion. Together, this suggests that the nature of object representations during dynamic occlusion is different from visual representations during perception.
William Thyer; Kirsten C. S. Adam; Gisella K. Diaz; Itzel N. Velázquez Sánchez; Edward K. Vogel; Edward Awh
Storage in visual working memory recruits a content-independent pointer system Journal Article
In: Psychological Science, vol. 33, no. 10, pp. 1680–1694, 2022.
Past work has shown that storage in working memory elicits stimulus-specific neural activity that tracks the stored content. Here, we present evidence for a distinct class of load-sensitive neural activity that indexes items without representing their contents per se. We recorded electroencephalogram (EEG) activity while adult human subjects stored varying numbers of items in visual working memory. Multivariate analysis of the scalp topography of EEG voltage enabled precise tracking of the number of individuated items stored and robustly predicted individual differences in working memory capacity. Critically, this signature of working memory load generalized across variations in both the type and number of visual features stored about each item, suggesting that it tracked the number of individuated memory representations and not the content of those memories. We hypothesize that these findings reflect the operation of a capacity-limited pointer system that supports on-line storage and attentive tracking.
Anne E. Urai; Tobias H. Donner
Persistent activity in human parietal cortex mediates perceptual choice repetition bias Journal Article
In: Nature Communications, vol. 13, no. 1, pp. 1–15, 2022.
Humans and other animals tend to repeat or alternate their previous choices, even when judging sensory stimuli presented in a random sequence. It is unclear if and how sensory, associative, and motor cortical circuits produce these idiosyncratic behavioral biases. Here, we combined behavioral modeling of a visual perceptual decision with magnetoencephalographic (MEG) analyses of neural dynamics, across multiple regions of the human cerebral cortex. We identified distinct history-dependent neural signals in motor and posterior parietal cortex. Gamma-band activity in parietal cortex tracked previous choices in a sustained fashion, and biased evidence accumulation toward choice repetition; sustained beta-band activity in motor cortex inversely reflected the previous motor action, and biased the accumulation starting point toward alternation. The parietal, not motor, signal mediated the impact of previous on current choice and reflected individual differences in choice repetition. In sum, parietal cortical signals seem to play a key role in shaping choice sequences.
Mats W. J. Es; Tom R. Marshall; Eelke Spaak; Ole Jensen; Jan-Mathijs Schoffelen
Phasic modulation of visual representations during sustained attention Journal Article
In: European Journal of Neuroscience, vol. 55, no. 11-12, pp. 3191–3208, 2022.
Sustained attention has long been thought to benefit perception in a continuous fashion, but recent evidence suggests that it affects perception in a discrete, rhythmic way. Periodic fluctuations in behavioral performance over time, and modulations of behavioral performance by the phase of spontaneous oscillatory brain activity point to an attentional sampling rate in the theta or alpha frequency range. We investigated whether such discrete sampling by attention is reflected in periodic fluctuations in the decodability of visual stimulus orientation from magnetoencephalographic (MEG) brain signals. In this exploratory study, human subjects attended one of the two grating stimuli, while MEG was being recorded. We assessed the strength of the visual representation of the attended stimulus using a support vector machine (SVM) to decode the orientation of the grating (clockwise vs. counterclockwise) from the MEG signal. We tested whether decoder performance depended on the theta/alpha phase of local brain activity. While the phase of ongoing activity in the visual cortex did not modulate decoding performance, theta/alpha phase of activity in the frontal eye fields and parietal cortex, contralateral to the attended stimulus did modulate decoding performance. These findings suggest that phasic modulations of visual stimulus representations in the brain are caused by frequency-specific top-down activity in the frontoparietal attention network, though the behavioral relevance of these effects could not be established.
Dominik Welke; Edward A. Vessel
Naturalistic viewing conditions can increase task engagement and aesthetic preference but have only minimal impact on EEG quality Journal Article
In: NeuroImage, vol. 256, pp. 1–19, 2022.
Free gaze and moving images are typically avoided in EEG experiments due to the expected generation of artifacts and noise. Yet for a growing number of research questions, loosening these rigorous restrictions would be beneficial. Among these is research on visual aesthetic experiences, which often involve open-ended exploration of highly variable stimuli. Here we systematically compare the effect of conservative vs. more liberal experimental settings on various measures of behavior, brain activity and physiology in an aesthetic rating task. Our primary aim was to assess EEG signal quality. 43 participants either maintained fixation or were allowed to gaze freely, and viewed either static images or dynamic (video) stimuli consisting of dance performances or nature scenes. A passive auditory background task (auditory steady-state response; ASSR) was added as a proxy measure for overall EEG recording quality. We recorded EEG, ECG and eye tracking data, and participants rated their aesthetic preference and state of boredom on each trial. Whereas both behavioral ratings and gaze behavior were affected by task and stimulus manipulations, EEG SNR was barely affected and generally robust across all conditions, despite only minimal preprocessing and no trial rejection. In particular, we show that using video stimuli does not necessarily result in lower EEG quality and can, on the contrary, significantly reduce eye movements while increasing both the participants' aesthetic response and general task engagement. We see these as encouraging results indicating that — at least in the lab — more liberal experimental conditions can be adopted without significant loss of signal quality.
Wen Wen; Zhibang Huang; Yin Hou; Sheng Li
Tracking neural markers of template formation and implementation in attentional inhibition under different distractor consistency Journal Article
In: Journal of Neuroscience, vol. 42, no. 24, pp. 4927–4936, 2022.
Performing visual search tasks requires optimal attention deployment to promote targets and inhibit distractors. Rejection templates based on the distractor's feature can be built to constrain the search process. We measured electroencephalography (EEG) of human participants of both sexes when they performed a visual search task in conditions where the distractor cues were constant within a block (fixed-cueing) or changed on a trial-by-trial basis (varied-cueing). In the fixed-cueing condition, sustained decoding of the cued colors could be achieved during the retention interval and the participants with higher decoding accuracy showed larger suppression benefits of the distractor cueing in the search period. In the varied-cueing condition, the cued color could only be transiently decoded after its onset and the higher decoding accuracy was observed from the participants who demonstrated lower suppression benefit. The differential neural representations of the to-be-ignored color in the two cueing conditions as well as their reverse associations with behavioral performance implied that rejection templates were formed in the fixed-cueing condition but not in the varied-cueing condition. Additionally, we observed stronger posterior alpha lateralization and mid-frontal theta/beta power during the retention interval of the varied-cueing condition, indicating the cognitive costs in template formation caused by the trialwise change of distractor colors. Taken together, our findings revealed the neural markers associated with the critical roles of distractor consistency in linking template formation to successful inhibition.
Stephen Whitmarsh; Christophe Gitton; Veikko Jousmäki; Jérôme Sackur; Catherine Tallon-Baudry
Neuronal correlates of the subjective experience of attention Journal Article
In: European Journal of Neuroscience, vol. 55, no. 11-12, pp. 3465–3482, 2022.
The effect of top–down attention on stimulus-evoked responses and alpha oscillations and the association between arousal and pupil diameter are well established. However, the relationship between these indices, and their contribution to the subjective experience of attention, remains largely unknown. Participants performed a sustained (10–30 s) attention task in which rare (10%) targets were detected within continuous tactile stimulation (16 Hz). Trials were followed by attention ratings on an 8-point visual scale. Attention ratings correlated negatively with contralateral somatosensory alpha power and positively with pupil diameter. The effect of pupil diameter on attention ratings extended into the following trial, reflecting a sustained aspect of attention related to vigilance. The effect of alpha power did not carry over to the next trial and furthermore mediated the association between pupil diameter and attention ratings. Variations in steady-state amplitude reflected stimulus processing under the influence of alpha oscillations but were only weakly related to subjective ratings of attention. Together, our results show that both alpha power and pupil diameter are reflected in the subjective experience of attention, albeit on different time spans, while continuous stimulus processing might not contribute to the experience of attention.
Seth B. Winward; James Siklos-Whillans; Roxane J. Itier
Impact of face outline, parafoveal feature number and feature type on early face perception in a gaze-contingent paradigm: A mass-univariate re-analysis of ERP data Seth Journal Article
In: Neuroimage: Reports, vol. 2, no. 4, pp. 1–17, 2022.
Recent ERP research using a gaze-contingent paradigm suggests the face-sensitive N170 component is modulated by the presence of a face outline, the number of parafoveal facial features, and the type of feature in parafovea (Parkington and Itier, 2019). The present study re-analyzed these data using robust mass univariate statistics available through the LIMO toolbox, allowing the examination of the ERP signal across all electrodes and time points. We replicated the finding that the presence of a face outline significantly reduced ERP latencies and amplitudes, suggesting it is an important cue to the prototypical face template. However, we found that this effect began around 114 ms, and was maximal during the P1-N170 and N170-P2 intervals. The number of features present in parafovea also impacted the entire waveform, with systematic reductions in amplitude and latency as the number of features increased. This effect was maximal around 120 ms during the P1-N170 interval and around 170 ms between the N170 and P2. The ERP response was also modulated by feature type; contrary to previous findings this effect was maximal around 200 ms and the P2 peak. Although we provide partial repli- cation of the previous results on the N170, the effects were more temporally distributed in the present analysis. These effects were generally maximal before and after the N170 and were the weakest at the N170 peak itself. This re-analysis demonstrates that classical ERP analysis can obscure important aspects of face processing beyond the N170 peak, and that tools like mass univariate statistics are needed to shed light on the whole time-course of face processing.
Maren-Isabel Wolf; Maximilian Bruchmann; Gilles Pourtois; Sebastian Schindler; Thomas Straube
Top-down modulation of early visual processing in V1: Dissociable neurophysiological effects of spatial attention, attentional load and task-relevance Journal Article
In: Cerebral Cortex, vol. 32, no. 10, pp. 2112–2128, 2022.
Until today, there is an ongoing discussion if attention processes interact with the information processing stream already at the level of the C1, the earliest visual electrophysiological response of the cortex. We used two highly powered experiments (each N = 52) and examined the effects of task relevance, spatial attention, and attentional load on individual C1 amplitudes for the upper or lower visual hemifield. Bayesian models revealed evidence for the absence of load effects but substantial modulations by task-relevance and spatial attention. When the C1-eliciting stimulus was a task-irrelevant, interfering distracter, we observed increased C1 amplitudes for spatially unattended stimuli. For spatially attended stimuli, different effects of task-relevance for the two experiments were found. Follow-up exploratory single-trial analyses revealed that subtle but systematic deviations from the eye-gaze position at stimulus onset between conditions substantially influenced the effects of attention and task relevance on C1 amplitudes, especially for the upper visual field. For the subsequent P1 component, attentional modulations were clearly expressed and remained unaffected by these deviations. Collectively, these results suggest that spatial attention, unlike load or task relevance, can exert dissociable top-down modulatory effects at the C1 and P1 levels.
Wenyuan Yu; Wenhui Sun; Nai Ding
Asymmetrical cross-modal influence on neural encoding of auditory and visual features in natural scenes Journal Article
In: NeuroImage, vol. 255, pp. 1–10, 2022.
Natural scenes contain multi-modal information, which is integrated to form a coherent perception. Previous studies have demonstrated that cross-modal information can modulate neural encoding of low-level sensory features. These studies, however, mostly focus on the processing of single sensory events or rhythmic sensory sequences. Here, we investigate how the neural encoding of basic auditory and visual features is modulated by cross-modal information when the participants watch movie clips primarily composed of non-rhythmic events. We presented audiovisual congruent and audiovisual incongruent movie clips, and since attention can modulate cross-modal interactions, we separately analyzed high- and low-arousal movie clips. We recorded neural responses using electroencephalography (EEG), and employed the temporal response function (TRF) to quantify the neural encoding of auditory and visual features. The neural encoding of sound envelope is enhanced in the audiovisual congruent condition than the incongruent condition, but this effect is only significant for high-arousal movie clips. In contrast, audiovisual congruency does not significantly modulate the neural encoding of visual features, e.g., luminance or visual motion. In summary, our findings demonstrate asymmetrical cross-modal interactions during the processing of natural scenes that lack rhythmicity: Congruent visual information enhances low-level auditory processing, while congruent auditory information does not significantly modulate low-level visual processing.
Marie Zelenina; MacIej Kosilo; Janir Da Cruz; Marília Antunes; Patrícia Figueiredo; Mitul A. Mehta; Diana Prata
Temporal dynamics of intranasal oxytocin in human brain electrophysiology Journal Article
In: Cerebral Cortex, vol. 32, no. 14, pp. 3110–3126, 2022.
Oxytocin (OT) is a key modulator of human social cognition, popular in behavioral neuroscience. To adequately design and interpret intranasal OT (IN-OT) research, it is crucial to know for how long it affects human brain function once administered. However, this has been mostly deduced from peripheral body f luids studies, or uncommonly used dosages. We aimed to characterize IN-OT's effects on human brain function using resting-state EEG microstates across a typical experimental session duration. Nineteen healthy males participated in a double-blind, placebo-controlled, within-subject, cross-over design of 24 IU of IN-OT in 12-min windows 15 min-to1 h 42min after administration. We observed IN-OT effects on all microstates, across the observation span. During eyes-closed, IN-OT increased duration and contribution of A and contribution and occurrence of D, decreased duration and contribution of B and C; and increased transition probability C-to-B and C-to-D. In eyes-open, it increased A-to-C and A-to-D. As microstates A and D have been related to phonological auditory and attentional networks, respectively, we posit IN-OT may tune the brain for reception of external stimuli, particularly of social nature - tentatively supporting current neurocognitive hypotheses of OT. Moreover, we contrast our overall results against a comprehensive literature review of IN-OT time-course effects in the brain, highlighting comparability issues.
TianHong Zhang; YingYu Yang; LiHua Hua Xu; XiaoChen Tang; YeGang Hu; Xin Xiong; YanYan Wei; HuiRu Ru Cui; YingYing Tang; HaiChun Liu; Tao Chen; Zhi Liu; Li Hui; ChunBo Li; XiaoLi Guo; JiJun Wang
Inefficient integration during multiple facial processing in pre-morbid and early phases of psychosis Journal Article
In: The World Journal of Biological Psychiatry, vol. 23, no. 5, pp. 1–13, 2022.
Objectives: We used eye-tracking to evaluate multiple facial context processing and event-related potential (ERP) to evaluate multiple facial recognition in individuals at clinical high risk (CHR) for psychosis. Methods: In total, 173 subjects (83 CHRs and 90 healthy controls [HCs]) were included and their emotion perception performances were accessed. A total of 40 CHRs and 40 well-matched HCs completed an eye-tracking task where they viewed pictures depicting a person in the foreground, presented as context-free, context-compatible, and context-incompatible. During the two-year follow-up, 26 CHRs developed psychosis, including 17 individuals who developed first-episode schizophrenia (FES). Eighteen well-matched HCs were made to complete the face number detection ERP task with image stimuli of one, two, or three faces. Results: Compared to the HC group, the CHR group showed reduced visual attention to contextual processing when viewing multiple faces. With the increasing complexity of contextual faces, the differences in eye-tracking characteristics also increased. In the ERP task, the N170 amplitude decreased with a higher face number in FES patients, while it increased with a higher face number in HCs. Conclusions: Individuals in the very early phase of psychosis showed facial processing deficits with supporting evidence of different scan paths during context processing and disruption of N170 during multiple facial recognition.
Jing Zhu; Shiqing Wei; Xiannian Xie; Changlin Yang; Yizhou Li; Xiaowei Li; Bin Hu
Content-based multiple evidence fusion on EEG and eye movements for mild depression recognition Journal Article
In: Computer Methods and Programs in Biomedicine, vol. 226, pp. 1–11, 2022.
Background and objective: Depression is a serious neurological disorder that has become a major health problem worldwide. The detection of mild depression is important for the diagnosis of depression in early stages. This research seeks to find a more accurate fusion model which can be used for mild depression detection using Electroencephalography and eye movement data. Methods: This study proposes a content-based multiple evidence fusion (CBMEF) method, which fuses EEG and eye movement data at decision level. The method mainly includes two modules, the classification performance matrix module and the dual-weight fusion module. The classification performance matrices of different modalities are estimated by Bayesian rule based on confusion matrix and Mahalanobis distance, and the matrices were used to correct the classification results. Then the relative conflict degree of each modality is calculated, and different weights are assigned to the above modalities at the decision fusion layer according to this conflict degree. Results: The experimental results show that the proposed method outperforms other fusion methods as well as the single modality results. The highest accuracies achieved 91.12%, and sensitivity, specificity and precision were 89.20%, 93.03%, 92.76%. Conclusions: The promising results showed the potential of the proposed approach for the detection of mild depression. The idea of introducing the classification performance matrix and the dual-weight model to multimodal biosignals fusion casts a new light on the researches of depression recognition.
Jing Zhu; Changlin Yang; Xiannian Xie; Shiqing Wei; Yizhou Li; Xiaowei Li; Bin Hu
Mutual Information Based Fusion Model (MIBFM): Mild depression recognition using EEG and pupil area signals Journal Article
In: Journal of LATEX Class Files, vol. 3045, pp. 1–14, 2022.
The detection of mild depression is conducive to the early intervention and treatment of depression. This study explored the fusion of electroencephalography (EEG) and pupil area signals to build an effective and convenient mild depression recognition model. We proposed Mutual Information Based Fusion Model (MIBFM), which innovatively used pupil area signals to select EEG electrodes based on mutual information. Then we extracted features from EEG and pupil area signals in different bands, and fused bimodal features using the denoising autoencoder. Experimental results showed that MIBFM could obtain the highest accuracy of 87.03%. And MIBFM exhibited better performance than other existing methods. Our findings validate the effectiveness of the use of pupil area as signals, which makes eye movement signals can be easily obtained using high resolution camera, and the EEG electrode selection scheme based on mutual information is also proved to be an applicable solution for data dimension reduction and multimodal complementary information screening. This study casts a new light for mild depression recognition using multimodal data of EEG and pupil area signals, and provides a theoretical basis for the development of portable and universal application systems.
Sven Braeutigam; Jessica Clare Scaife; Tipu Aziz; Rebecca J. Park
A longitudinal magnetoencephalographic study of the effects of deep brain stimulation on neuronal dynamics in severe anorexia nervosa Journal Article
In: Frontiers in Behavioral Neuroscience, vol. 16, pp. 1–12, 2022.
Anorexia Nervosa (AN) is a debilitating psychiatric disorder characterized by the relentless pursuit of thinness, leading to severe emaciation. Magnetoencephalography (MEG)was used to record the neuronal response in seven patients with treatment-resistant AN while completing a disorder-relevant food wanting task. The patients underwent a 15-month protocol, where MEG scans were conducted pre-operatively, post-operatively prior to deep brain stimulation (DBS) switch on, twice during a blind on/off month and at protocol end. Electrodes were implanted bilaterally into the nucleus accumbens with stimulation at the anterior limb of the internal capsule using rechargeable implantable pulse generators. Three patients met criteria as responders at 12 months of stimulation, showing reductions of eating disorder psychopathology of over 35%. An increase in alpha power, as well as evoked power at latencies typically associated with visual processing, working memory, and contextual integration was observed in ON compared to OFF sessions across all seven patients. Moreover, an increase in evoked power at P600-like latencies as well as an increase in $gamma$-band phase-locking over anterior-to-posterior regions were observed for high- compared to low-calorie food image only in ON sessions. These findings indicate that DBS modulates neuronal process in regions far outside the stimulation target site and at latencies possibly reflecting task specific processing, thereby providing further evidence that deep brain stimulation can play a role in the treatment of otherwise intractable psychiatric disorders.
Marion Brickwedde; Yulia Bezsudnova; Anna Kowalczyk; Ole Jensen; Alexander Zhigalov
Application of rapid invisible frequency tagging for brain computer interfaces Journal Article
In: Journal of Neuroscience Methods, vol. 382, pp. 1–9, 2022.
Background: Brain-computer interfaces (BCI) based on steady-state visual evoked potentials (SSVEPs/SSVEFs) are among the most commonly used BCI systems. They require participants to covertly attend to visual objects flickering at specified frequencies. The attended location is decoded online by analysing the power of neuronal responses at the flicker frequency. New method: We implemented a novel rapid invisible frequency-tagging technique, utilizing a state-of-the-art projector with refresh rates of up to 1440 Hz. We flickered the luminance of visual objects at 56 and 60 Hz, which was invisible to participants but produced strong neuronal responses measurable with magnetoencephalography (MEG). The direction of covert attention, decoded from frequency-tagging responses, was used to control an online BCI PONG game. Results: Our results show that seven out of eight participants were able to play the pong game controlled by the frequency-tagging signal, with average accuracies exceeding 60 %. Importantly, participants were able to modulate the power of the frequency-tagging response within a 1-second interval, while only seven occipital sensors were required to reliably decode the neuronal response. Comparison with existing methods: In contrast to existing SSVEP-based BCI systems, rapid frequency-tagging does not produce a visible flicker. This extends the time-period participants can use it without fatigue, by avoiding distracting visual input. Furthermore, higher frequencies increase the temporal resolution of decoding, resulting in higher communication rates. Conclusion: Using rapid invisible frequency-tagging opens new avenues for fundamental research and practical applications. In combination with novel optically pumped magnetometers (OPMs), it could facilitate the development of high-speed and mobile next-generation BCI systems.
Maximilian Bruchmann; Sebastian Schindler; Mandana Dinyarian; Thomas Straube
The role of phase and orientation for ERP modulations of spectrum-manipulated fearful and neutral faces Journal Article
In: Psychophysiology, vol. 59, no. 3, pp. 1–13, 2022.
Prioritized processing of fearful compared to neutral faces has been proposed to result from evolutionary adaptation of the contrast sensitivity function (CSF) to the features of emotionally relevant faces and/or vice versa. However, it is unknown whether a stimulus merely has to feature the amplitude spectrum of a fearful face to be prioritized or whether the relevant spatial frequencies have to occur with specific phases and orientations. Prioritized processing is indexed by specific increases of Event-Related Potentials (ERPs) of the EEG and occurs throughout different early processing stages, indexed by emotion-related modulations of the P1, N170, and EPN. In this pre-registered study, we manipulated phase and amplitude properties of the Fourier spectra of neutral and fearful faces to test the effect of phase coherence (PC, face vs. scramble) and orientation coherence (OC, original vs. rotational average) and their interactions with differential emotion processing. We found that differential emotion processing was not present at the level of P1 but strongly affected N170 and EPN. In both cases, intact phase coherence was required for enhanced processing of fearful faces. OC did not interact with emotion. While faces produced the typical N170 effect, we observed a reversed effect for scrambles. Additional exploratory independent component analysis (ICA) suggests that this reversal could signal a mismatch between an early "perceptual hypothesis" and feedback of configural information. In line with our expectations, fearful-neutral differences for the N170 and EPN depend on configural information, i.e., recognizable faces.
Jon Burnsky; Franziska Kretzschmar; Erika Mayer; Adrian Staub
The influence of predictability, visual contrast, and preview validity on eye movements and N400 amplitude: Co-registration evidence that the N400 reflects late processes Journal Article
In: Language, Cognition and Neuroscience, pp. 1–22, 2022.
Two eye movement/EEG co-registration experiments investigated effects of predictability, visual contrast, and parafoveal preview in normal reading. Replicating previous studies, in Experiment 1 contrast and predictability additively influenced fixation durations, and in Experiment 2 invalid preview eliminated the predictability effect on early eye movement measures. In both experiments, predictability influenced the amplitude of the N400 component of the fixation-related potential. In Experiment 1, visual contrast did not influence the N400, and in Experiment 2, the effect of predictability on the N400 was larger with invalid preview, in opposition to the eye movement pattern. The N400 may reflect a late process of accessing conceptual representations while the duration of the eyes' fixation on a word is sensitive to the difficulty of perceptual encoding and early stages of word recognition. The effects of predictability on both fixation duration and the N400 suggest an influence of this variable at two distinct processing stages.
Anna Corriveau; Alexis Kidder; Lina Teichmann; Susan G. Wardle; Chris I. Baker
Sustained neural representations of personally familiar people and places during cued recall Journal Article
In: Cortex, vol. 158, pp. 71–82, 2022.
The recall and visualization of people and places from memory is an everyday occurrence, yet the neural mechanisms underpinning this phenomenon are not well understood. In particular, the temporal characteristics of the internal representations generated by active recall are unclear. Here, we used magnetoencephalography (MEG) and multivariate pattern analysis to measure the evolving neural representation of familiar places and people across the whole brain when human participants engage in active recall. To isolate self- generated imagined representations, we used a retro-cue paradigm in which participants were first presented with two possible labels before being cued to recall either the first or second item. We collected personalized labels for specific locations and people familiar to each participant. Importantly, no visual stimuli were presented during the recall period, and the retro-cue paradigm allowed the dissociation of responses associated with the la- bels from those corresponding to the self-generated representations. First, we found that following the retro-cue it took on average $sim$1000 ms for distinct neural representations of freely recalled people or places to develop. Second, we found distinct representations of personally familiar concepts throughout the 4 s recall period. Finally, we found that these representations were highly stable and generalizable across time. These results suggest that self-generated visualizations and recall of familiar places and people are subserved by a stable neural mechanism that operates relatively slowly when under conscious control.
Prateek Dhamija; Allison Wong; Asaf Gilboa
Early auditory event related potentials distinguish higher-order from first-order aversive conditioning Journal Article
In: Frontiers in Behavioral Neuroscience, vol. 16, pp. 1–14, 2022.
Stimuli in reality rarely co-occur with primary reward or punishment to allow direct associative learning of value. Instead, value is thought to be inferred through complex higher-order associations. Rodent research has demonstrated that the formation and maintenance of first-order and higher-order associations are supported by distinct neural substrates. In this study, we explored whether this pattern of findings held true for humans. Participants underwent first-order and subsequent higher-order conditioning using an aversive burst of white noise or neutral tone as the unconditioned stimuli. Four distinct tones, initially neutral, served as first-order and higher-order conditioned stimuli. Autonomic and neural responses were indexed by pupillometry and evoked response potentials (ERPs) respectively. Conditioned aversive values of first-order and higher-order stimuli led to increased autonomic responses, as indexed by pupil dilation. Distinct temporo-spatial auditory evoked response potentials were elicited by first-order and high-order conditioned stimuli. Conditioned first-order responses peaked around 260 ms and source estimation suggested a primary medial prefrontal and amygdala source. Conversely, conditioned higher-order responses peaked around 120 ms with an estimated source in the medial temporal lobe. Interestingly, pupillometry responses to first-order conditioned stimuli were diminished after higher order training, possibly signifying concomitant incidental extinction, while responses to higher-order stimuli remained. This suggests that once formed, higher order associations are at least partially independent of first order conditioned representations. This experiment demonstrates that first-order and higher-order conditioned associations have distinct neural signatures, and like rodents, the medial temporal lobe may be specifically involved with higher-order conditioning.
Marwa El Zein; Ray J. Dolan; Bahador Bahrami
Shared responsibility decreases the sense of agency in the human brain Journal Article
In: Journal of Cognitive Neuroscience, vol. 34, pp. 2065–2081, 2022.
Sharing responsibility in social decision-making helps individuals use the flexibility of the collective context to benefit them-selves by claiming credit for good outcomes or avoiding the blame for bad outcomes. Using magnetoencephalography, we examined the neuronal basis of the impact that social context has on this flexible sense of responsibility. Participants performed a gambling task in various social contexts and reported feeling less responsibility when playing as a member of a team. A reduced magnetoencephalography outcome processing effect was observed as a function of decreasing responsibility at 200 msec post outcome onset and was centered over parietal, central, and frontal brain regions. Before outcome revelation in socially made decisions, an attenuated motor preparation signature at 500 msec after stimulus onset was found. A boost in reported responsibility for positive outcomes in social contexts was associated with increased activity in regions related to social and reward processing. Together, these results show that sharing responsibility with others reduces agency, influencing pre-outcome motor preparation and post-outcome processing, and provides opportunities to flexibly claim credit for positive outcomes.
Hesham A. Elshafei; Corinne Orlemann; Saskia Haegens
The impact of eye closure on anticipatory $alpha$ activity in a tactile discrimination task Journal Article
In: eNeuro, vol. 9, no. 1, pp. 1–11, 2022.
One of the very first observations made regarding $alpha$ oscillations (8–14 Hz), is that they increase in power over posterior areas when awake participants close their eyes. Recent work, especially in the context of (spatial) attention, suggests that $alpha$ activity reflects a mechanism of functional inhibition. However, it remains unclear how eye closure impacts anticipatory $alpha$ modulation observed in attention paradigms, and how this affects subsequent behavioral performance. Here, we recorded magnetoencephalography (MEG) in 33 human participants performing a tactile discrimination task with their eyes open versus closed. We replicated the hallmarks of previous somatosensory spatial attention studies: $alpha$ lateralization across the somatosensory cortices as well as $alpha$ increase over posterior (visual) regions. Furthermore, we found that eye closure leads to (1) reduced task performance; (2) widespread increase in $alpha$ power; and (3) reduced anticipatory visual $alpha$ modulation (4) with no effect on somatosensory $alpha$ lateralization. Regardless of whether participants had their eyes open or closed, increased visual $alpha$ power and somatosensory $alpha$ lateralization improved their performance. Thus, we provide evidence that eye closure does not alter the impact of anticipatory $alpha$ modulations on behavioral performance. We propose there is an optimal visual $alpha$ level for somatosensory task performance, which can be achieved through a combination of eye closure and top-down anticipatory attention.
Tahnée Engelen; Anne Buot; Julie Grèzes; Catherine Tallon-baudry
Whose emotion is it? Perspective matters to understand brain-body interactions in emotions Journal Article
In: NeuroImage, vol. 268, pp. 1–14, 2022.
Feeling happy, or judging whether someone else is feeling happy are two distinct facets of emotions that nev- ertheless rely on similar physiological and neural activity. Differentiating between these two states, also called Self/Other distinction, is an essential aspect of empathy, but how exactly is it implemented? In non-emotional cognition, the transient neural response evoked at each heartbeat, or heartbeat evoked response (HER), indexes the self and signals Self/Other distinction. Here, using electroencephalography ( n = 32), we probe whether HERs' role in Self/Other distinction extends also to emotion–a domain where brain-body interactions are particularly relevant. We asked participants to rate independently validated affective scenes, reporting either their own emotion (Self) or the emotion expressed by people in the scene (Other). During the visual cue indicating to adopt the Self or Other perspective, before the affective scene, HERs distinguished between the two conditions, in visual cortices as well as in the right frontal operculum. Physiological reactivity (facial electromyogram, skin conduc- tance, heart rate) during affective scene co-varied as expected with valence and arousal ratings, but also with the Self- or Other-perspective adopted. Finally, HERs contributed to the subjective experience of valence in the Self condition, in addition to and independently from physiological reactivity. We thus show that HERs represent a trans-domain marker of Self/Other distinction, here specifically contributing to experienced valence. We propose that HERs represent a form of evidence related to the ‘I' part of the judgement ‘To which extent do I feel happy'. The ‘I' related evidence would be combined with the affective evidence collected during affective scene presentation, accounting at least partly for the difference between feeling an emotion and identifying it in someone else.
João Estiveira; Camila Dias; Diana Costa; João Castelhano; Miguel Castelo-Branco; Teresa Sousa
An action-independent role for midfrontal theta activity prior to error commission Journal Article
In: Frontiers in Human Neuroscience, vol. 16, pp. 1–17, 2022.
Error-related electroencephalographic (EEG) signals have been widely studied concerning the human cognitive capability of differentiating between erroneous and correct actions. Midfrontal error-related negativity (ERN) and theta band oscillations are believed to underlie post-action error monitoring. However, it remains elusive how early monitoring activity is trackable and what are the pre-response brain mechanisms related to performance monitoring. Moreover, it is still unclear how task-specific parameters, such as cognitive demand or motor control, influence these processes. Here, we aimed to test pre- and post-error EEG patterns for different types of motor responses and investigate the neuronal mechanisms leading to erroneous actions. We implemented a go/no-go paradigm based on keypresses and saccades. Participants received an initial instruction about the direction of response to be given based on a facial cue and a subsequent one about the type of action to be performed based on an object cue. The paradigm was tested in 20 healthy volunteers combining EEG and eye tracking. We found significant differences in reaction time, number, and type of errors between the two actions. Saccadic responses reflected a higher number of premature responses and errors compared to the keypress ones. Nevertheless, both led to similar EEG patterns, supporting previous evidence for increased ERN amplitude and midfrontal theta power during error commission. Moreover, we found pre-error decreased theta activity independent of the type of action. Source analysis suggested different origin for such pre- and post-error neuronal patterns, matching the anterior insular cortex and the anterior cingulate cortex, respectively. This opposite pattern supports previous evidence of midfrontal theta not only as a neuronal marker of error commission but also as a predictor of action performance. Midfrontal theta, mostly associated with alert mechanisms triggering behavioral adjustments, also seems to reflect pre-response attentional mechanisms independently of the action to be performed. Our findings also add to the discussion regarding how salience network nodes interact during performance monitoring by suggesting that pre- and post-error patterns have different neuronal sources within this network.
John J. Foxe; Emily J. Knight; Evan J. Myers; Cody Zhewei Cao; Sophie Molholm; Edward G. Freedman
The strength of feedback processing is associated with resistance to visual backward masking during Illusory Contour processing in adult humans Journal Article
In: NeuroImage, vol. 259, pp. 1–10, 2022.
Re-entrant feedback processing is a key mechanism of visual object-recognition, especially under compromised viewing conditions where only sparse information is available and object features must be interpolated. Illusory Contour stimuli are commonly used in conjunction with Visual Evoked Potentials (VEP) to study these filling-in processes, with characteristic modulation of the VEP in the ∼100-150 ms timeframe associated with this re-entrant processing. Substantial inter-individual variability in timing and amplitude of feedback-related VEP modulation is observed, raising the question whether this variability might underlie inter-individual differences in the ability to form strong perceptual gestalts. Backward masking paradig ms have been used to study inter-individual variance in the ability to form robust object perceptions before processing of the mask interferes with object-recognition. Some individuals recognize objects when the time between target object and mask is extremely short, whereas others struggle to do so even at longer target-to-mask intervals. We asked whether timing and amplitude of feedback-related VEP modulations were associated with individual differences in resistance to backward masking. Participants (N=40) showed substantial performance variability in detecting Illusory Contours at intermediate target-to-mask intervals (67 ms and 117 ms), allowing us to use kmeans clustering to divide the population into four performance groups (poor, low-average, high-average, superior). There was a clear relationship between the amplitude (but not the timing) of feedback-related VEP modulation and Illusory Contour detection during backward masking. We conclude that individual differences in the strength of feedback processing in neurotypical humans lead to differences in the ability to quickly establish perceptual awareness of incomplete visual objects.
Anna L. Gert; Benedikt V. Ehinger; Silja Timm; Tim C. Kietzmann; Peter König
WildLab: A naturalistic free viewing experiment reveals previously unknown electroencephalography signatures of face processing Journal Article
In: European Journal of Neuroscience, vol. 56, pp. 6022–6038, 2022.
Neural mechanisms of face perception are predominantly studied in well-controlled experimental settings that involve random stimulus sequences and fixed eye positions. While powerful, the employed paradigms are far from what constitutes natural vision. Here, we demonstrate the feasibility of ecologically more valid experimental paradigms using natural viewing behavior, by combining a free viewing paradigm on natural scenes, free of photographer bias, with advanced data processing techniques that correct for overlap effects and co-varying nonlinear dependencies of multiple eye movement parameters. We validate this approach by replicating classic N170 effects in neural responses, triggered by fixation onsets (fERPs). Importantly, our more natural stimulus paradigm yielded smaller variability between subjects than the classic setup. Moving beyond classic temporal and spatial effect locations, our experiment furthermore revealed previously unknown signatures of face processing: This includes category-specific modulation of the event-related potential (ERP)'s amplitude even before fixation onset, as well as adaptation effects across subsequent fixations depending on their history.
Erin Goddard; Thomas A. Carlson; Alexandra Woolgar
Spatial and feature-selective attention have distinct, interacting effects on population-level tuning Journal Article
In: Journal of Cognitive Neuroscience, vol. 34, no. 2, pp. 290–312, 2022.
Attention can be deployed in different ways: When searching for a taxi in New York City, we can decide where to attend (e.g., to the street) and what to attend to (e.g., yellow cars). Although we use the same word to describe both processes, nonhuman primate data suggest that these produce distinct effects on neural tuning. This has been challenging to assess in humans, but here we used an opportunity afforded by multivariate decoding of MEG data. We found that attending to an object at a particular location and attending to a particular object feature produced effects that interacted multiplicatively. The two types of attention induced distinct patterns of enhancement in occipital cortex, with feature-selective attention producing relatively more enhancement of small feature differences and spatial attention producing relatively larger effects for larger feature differences. An information flow analysis further showed that stimulus representations in occipital cortex were Granger-caused by coding in frontal cortices earlier in time and that the timing of this feedback matched the onset of attention effects. The data suggest that spatial and feature-selective attention rely on distinct neural mechanisms that arise from frontal-occipital information exchange, interacting multiplicatively to selectively enhance task-relevant information.
Mariana M. Gusso; Kate L. Christison-Lagay; David Zuckerman; Ganesh Chandrasekaran; Sharif I. Kronemer; Julia Z. Ding; Noah C. Freedman; Percy Nohama; Hal Blumenfeld
More than a feeling: Scalp EEG and eye signals in conscious tactile perception Journal Article
In: Consciousness and Cognition, vol. 105, pp. 1–14, 2022.
Understanding the neural basis of consciousness is a fundamental goal of neuroscience, and sensory perception is often used as a proxy for consciousness in empirical studies. However, most studies rely on reported perception of visual stimuli. Here we present behavior, high density scalp EEG and eye metric recordings collected simultaneously during a novel tactile threshold perception task. We found significant N80, N140 and P300 event related potentials in perceived trials and in perceived versus not perceived trials. Significance was limited to a P100 and P300 in not perceived trials. We also found an increase in pupil diameter and blink rate and a decrease in microsaccade rate following perceived relative to not perceived tactile stimuli. These findings support the use of eye metrics as a measure of physiological arousal associated with conscious perception. Eye metrics may also represent a novel path toward the creation of tactile no-report tasks in the future.
Tjerk P. Gutteling; Lonieke Sillekens; Nilli Lavie; Ole Jensen
Alpha oscillations reflect suppression of distractors with increased perceptual load Journal Article
In: Progress in Neurobiology, vol. 214, pp. 1–13, 2022.
Attention serves an essential role in cognition and behavior allowing us to focus on behaviorally-relevant objects while ignoring distraction. Perceptual load theory states that attentional resources are allocated according to the requirements of the task, i.e., its ‘load'. The theory predicts that the resources left to process irrelevant, possibly distracting stimuli, are reduced when the perceptual load is high. However, it remains unclear how this allocation of attentional resources specifically relates to neural excitability and suppression mechanisms. In this magnetoencephalography (MEG) study, we show that brain oscillations in the alpha band (8–13 Hz) implemented the suppression of distracting objects when the perceptual load was high. In parallel, high load increased the neuronal excitability for target objects, as reflected by rapid invisible frequency tagging. We suggest that the allocation of resources in tasks with high perceptual load is implemented by a gain increase for targets, complemented by distractor suppression reflected by alpha-band oscillations closing the ‘gate' for interference.
Qing He; Xin Yue Yang; Baoqi Gong; Keyan Bi; Fang Fang
Boosting visual perceptual learning by transcranial alternating current stimulation over the visual cortex at alpha frequency Journal Article
In: Brain Stimulation, vol. 15, no. 3, pp. 546–553, 2022.
Background: Transcranial alternating current stimulation (tACS) has been widely used to alter ongoing brain rhythms in a frequency-specific manner to modulate relevant cognitive functions, including visual functions. Therefore, it is a useful tool for exploring the causal role of neural oscillations in cognition. Visual functions can be improved substantially by training, which is called visual perceptual learning (VPL). However, whether and how tACS can modulate VPL is still unclear. Objective: This work aims to explore how tACS modulates VPL and the role of neural oscillations in VPL. Methods: A between-subjects design was adopted. Subjects were assigned to six groups and undertook five daily training sessions to execute an orientation discrimination task. During training, five groups received occipital tACS stimulation at 6, 10, 20, 40, and sham 10 Hz respectively, and one group was stimulated at the sensorimotor regions by 10 Hz tACS. Results: Compared with the sham stimulation, occipital tACS at 10 Hz, but not at other frequencies, accelerated perceptual learning and increased the performance improvement. However, these modulatory effects were absent when 10 Hz tACS was delivered to the sensorimotor areas. Moreover, the tACS-induced performance improvement lasted at least two months after the end of training. Conclusion: TACS can facilitate orientation discrimination learning in a frequency- and location-specific manner. Our findings provide strong evidence for a pivotal role of alpha oscillations in boosting VPL and shed new light on the design of effective neuromodulation protocols that can facilitate rehabilitation for patients with neuro-ophthalmological disorders.
Jacqueline Katharina Meier; Bernhard P. Staresina; Lars Schwabe
Stress diminishes outcome but enhances response representations during instrumental learning Journal Article
In: eLife, vol. 11, pp. 1–25, 2022.
Stress may shift behavioural control from a goal-directed system that encodes action-outcome relationships to a habitual system that learns stimulus-response associations. Although this shift to habits is highly relevant for stress-related psychopathologies, limitations of existing behavioural paradigms hinder research from answering the fundamental question of whether the stress-induced bias to habits is due to reduced outcome processing or enhanced response processing at the time of stimulus presentation, or both. Here, we used EEG-based multivariate pattern analysis to decode neural outcome representations crucial for goal-directed control, as well as response representations during instrumental learning. We show that stress reduced outcome representations but enhanced response representations. Both were directly associated with a behavioural index of habitual responding. Furthermore, changes in outcome and response representations were uncorrelated, suggesting that these may reflect distinct processes. Our findings indicate that habitual behaviour under stress may be the result of both enhanced stimulus-response processing and diminished outcome processing.
René Michel; Laura Dugué; Niko A. Busch
Distinct contributions of alpha and theta rhythms to perceptual and attentional sampling Journal Article
In: European Journal of Neuroscience, vol. 55, no. 11-12, pp. 3025–3039, 2022.
Accumulating evidence suggests that visual perception operates in an oscillatory fashion at an alpha frequency (around 10 Hz). Moreover, visual attention also seems to operate rhythmically, albeit at a theta frequency (around 5 Hz). Both rhythms are often associated to "perceptual snapshots" taken at the favorable phases of these rhythms. However, less is known about the unfavorable phases: do they constitute "blind gaps," requiring the observer to guess, or is information sampled with reduced precision insufficient for the task demands? As simple detection or discrimination tasks cannot distinguish these options, we applied a continuous report task by asking for the exact orientation of a Landolt ring's gap to estimate separate model parameters for precision and the amount of guessing. We embedded this task in a well-established psychophysical protocol by densely sampling such reports across 20 cue-target stimulus onset asynchronies in a Posner-like cueing paradigm manipulating involuntary spatial attention. Testing the resulting time courses of the guessing and precision parameters for rhythmicities using a fast Fourier transform, we found an alpha rhythm (9.6 Hz) in precision for invalidly cued trials and a theta rhythm (4.8 Hz) in the guess rate across validity conditions. These results suggest distinct roles of the perceptual alpha and the attentional theta rhythm. We speculate that both rhythms result in environmental sampling characterized by fluctuating spatial resolution, speaking against a strict succession of blind gaps and perceptual snapshots.
Sara Milligan; Martín Antúnez; Horacio A. Barber; Elizabeth R. Schotter
Are eye movements and EEG on the same page?: A coregistration study on parafoveal preview and lexical frequency Journal Article
In: Journal of Experimental Psychology: General, pp. 1–23, 2022.
Readers extract visual and linguistic information not only from fixated words but also upcoming parafoveal words to introduce new input efficiently into the language processing pipeline. The lexical frequency of upcoming words and similarity with subsequent foveal information both influence the amount of time people spend once they fixate the word foveally. However, it is unclear from eye movements alone the extent to which parafoveal word processing, and the integration of that word with foveally obtained information, continues after saccade plans have been initiated. To investigate the underlying neural processes involved in word recognition after saccade planning, we coregistered electroencephalogram (EEG) and eye movements during a gaze-contingent display change paradigm. We orthogonally manipulated the frequency of the parafoveal and foveal words and measured fixation related potentials (FRPs) upon foveal fixation. Eye movements showed primarily an effect of preview frequency, suggesting that saccade planning is based on the familiarity of the parafoveal input. FRPs, on the other hand, demonstrated a disruption in downstream processing when parafoveal and foveal input differed, but only when the parafoveal word was high frequency. These findings demonstrate that lexical processing continues after the eyes have moved away from a word and that eye movements and FRPs provide distinct but complementary accounts about oculomotor behavior and neural processing that cannot be obtained from either method in isolation. Furthermore, these findings put constraints on models of reading by suggesting that lexical processes that occur before an eye movement program is initiated are qualitatively different from those that occur afterward.
Michael T. Miuccio; Gregory J. Zelinsky; Joseph Schmidt
Are all real-world objects created equal? Estimating the “set-size” of the search target in visual working memory Journal Article
In: Psychophysiology, vol. 59, pp. 1–13, 2022.
Are all real-world objects created equal? Visual search difficulty increases with the number of targets and as target-related visual working memory (VWM) load increases. Our goal was to investigate the load imposed by individual real-world objects held in VWM in the context of search. Measures of visual clutter attempt to quantify real-world set-size in the context of scenes. We applied one of these measures, the number of proto-objects, to individual real-world objects and used contralateral delay activity (CDA) to measure the resulting VWM load. The current study presented a real-world object as a target cue, followed by a delay where CDA was measured. This was followed by a four-object search array. We compared CDA and later search performance from target cues containing a high or low number of proto-objects. High proto-object target cues resulted in greater CDA, longer search RTs, target dwell times, and reduced search guidance, relative to low proto-object targets. These findings demonstrate that targets with more proto-objects result in a higher VWM load and reduced search performance. This shows that the number of proto-objects contained within individual objects produce set-size like effects in VWM and suggests proto-objects may be a viable unit of measure of real-world VWM load. Importantly, this demonstrates that not all real-world objects are created equal.
Dinavahi V. P. S. Murty; Keerthana Manikandan; Wupadrasta Santosh Kumar; Ranjini Garani Ramesh; Simran Purokayastha; Bhargavi Nagendra; M. L. Abhishek; Aditi Balakrishnan; Mahendra Javali; Naren Prahalada Rao; Supratim Ray
Stimulus-induced gamma rhythms are weaker in human elderly with mild cognitive impairment and Alzheimer's disease Journal Article
In: Bio-protocol, vol. 12, no. 7, pp. 1–13, 2022.
Stimulus-induced narrow-band gamma oscillations (20–70 Hz) are induced in the visual areas of the brain when particular visual stimuli, such as bars, gratings, or full-screen hue, are shown to the subject. Such oscillations are modulated by higher cognitive functions, like attention, and working memory, and have been shown to be abnormal in certain neuropsychiatric disorders, such as schizophrenia, autism, and Alzheimer's disease. However, although electroencephalogram (EEG) remains one of the most non-invasive, inexpensive, and accessible methods to record brain signals, some studies have failed to observe discernable gamma oscillations in human EEG. In this manuscript, we have described in detail a protocol to elicit robust gamma oscillations in human EEG. We believe that our protocol could help in developing non-invasive gamma-based biomarkers in human EEG, for the early detection of neuropsychiatric disorders.
Adam J. Naples; Jennifer H. Foss-Feig; Julie M. Wolf; Vinod H. Srihari; James C. McPartland
Predictability modulates neural response to eye contact in ASD Journal Article
In: Molecular Autism, vol. 13, no. 42, pp. 1–12, 2022.
Background: Deficits in establishing and maintaining eye-contact are early and persistent vulnerabilities of autism spectrum disorder (ASD), and the neural bases of these deficits remain elusive. A promising hypothesis is that social features of autism may reflect difficulties in making predictions about the social world under conditions of uncertainty. However, no research in ASD has examined how predictability impacts the neural processing of eye-contact in naturalistic interpersonal interactions. Method: We used eye tracking to facilitate an interactive social simulation wherein onscreen faces would establish eye-contact when the participant looked at them. In Experiment One, receipt of eye-contact was unpredictable; in Experiment Two, receipt of eye-contact was predictable. Neural response to eye-contact was measured via the N170 and P300 event-related potentials (ERPs). Experiment One included 23 ASD and 46 typically developing (TD) adult participants. Experiment Two included 25 ASD and 43 TD adult participants. Results: When receipt of eye-contact was unpredictable, individuals with ASD showed increased N170 and increased, but non-specific, P300 responses. The magnitude of the N170 responses correlated with measures of sensory and anxiety symptomology, such that increased response to eye-contact was associated with increased symptomology. However, when receipt of eye-contact was predictable, individuals with ASD, relative to controls, exhibited slower N170s and no differences in the amplitude of N170 or P300. Limitations: Our ASD sample was composed of adults with IQ > 70 and included only four autistic women. Thus, further research is needed to evaluate how these results generalize across the spectrum of age, sex, and cognitive ability. Additionally, as analyses were exploratory, some findings failed to survive false-discovery rate adjustment. Conclusions: Neural response to eye-contact in ASD ranged from attenuated to hypersensitive depending on the predictability of the social context. These findings suggest that the vulnerabilities in eye-contact during social interactions in ASD may arise from differences in anticipation and expectation of eye-contact in addition to the perception of gaze alone.
Ádám Nárai; Zsuzsanna Nemecz; Zoltán Vidnyánszky; Béla Weiss
Lateralization of orthographic processing in fixed-gaze and natural reading conditions Journal Article
In: Cortex, vol. 157, pp. 99–116, 2022.
Lateralized processing of orthographic information is a hallmark of proficient reading. However, how this finding obtained for fixed-gaze processing of orthographic stimuli translates to ecologically valid reading conditions remained to be clarified. To address this shortcoming, here we assessed the lateralization of early orthographic processing in fixed-gaze and natural reading conditions using concurrent eye-tracking and EEG data recorded from young adults without reading difficulties. Sensor-space analyses confirmed the well-known left-lateralized negative-going deflection of fixed-gaze EEG activity throughout the period of early orthographic processing. At the same time, fixation-related EEG activity exhibited left-lateralized followed by right-lateralized processing of text stimuli during natural reading. A strong positive relationship was found between the early leftward lateralization in fixed-gaze and natural reading conditions. Using source-space analyses, early left-lateralized brain activity was obtained in lateraloccipital and posterior ventral occipito-temporal cortices reflecting letter-level processing in both conditions. In addition, in the same time interval, left-lateralized source activity was found also in premotor and parietal brain regions during natural reading. While brain activity remained left-lateralized in later stages representing word-level processing in posterior and middle ventral temporal regions in the fixed-gaze condition, fixation-related source activity became stronger in the right hemisphere in medial and more anterior ventral temporal brain regions indicating higher-level processing of orthographic information. Although our results show a strong positive relationship between the lateralization of letter-level processing in the two reading modes and suggest lateralized brain activity as a general marker for processing of orthographic information, they also clearly indicate the need for reading research in ecologically valid conditions to identify the neural basis of visuospatial attentional, oculomotor and higher-level processes specific to natural reading.
M. P. Noonan; A. H. Von Lautz; Y. Bauer; C. Summerfield; M. S. Stokes
Differential modulation of visual responses by distractor or target expectations Journal Article
In: Attention, Perception, and Psychophysics, pp. 1–18, 2022.
Discriminating relevant from irrelevant information in a busy visual scene is supported by statistical regularities in the environment. However, it is unclear to what extent immediate stimulus repetitions and higher order expectations (whether a repetition is statistically probable or not) are supported by the same neural mechanisms. Moreover, it is also unclear whether target and distractor-related processing are mediated by the same or different underlying neural mechanisms. Using a speeded target discrimination task, the present study implicitly cued subjects to the location of the target or the distractor via manipulations in the underlying stimulus predictability. In separate studies, we collected EEG and MEG alongside behavioural data. Results showed that reaction times were reduced with increased expectations for both types of stimuli and that these effects were driven by expected repetitions in both cases. Despite the similar behavioural pattern across target and distractors, neurophysiological measures distinguished the two stimuli. Specifically, the amplitude of the P1 was modulated by stimulus relevance, being reduced for repeated distractors and increased for repeated targets. The P1 was not, however, modulated by higher order stimulus expectations. These expectations were instead reflected in modulations in ERP amplitude and theta power in frontocentral electrodes. Finally, we observed that a single repetition of a distractor was sufficient to reduce decodability of stimulus spatial location and was also accompanied by diminished representation of stimulus features. Our results highlight the unique mechanisms involved in distractor expectation and suppression and underline the importance of studying these processes distinctly from target-related attentional control.
Nir Ofir; Ayelet N. Landau
Neural signatures of evidence accumulation in temporal decisions Journal Article
In: Current Biology, vol. 32, no. 18, pp. 4093–4100, 2022.
Cognitive models of interval timing can be formulated as an accumulation-to-bound process.1–5 However, the physiological manifestation of such processes has not yet been identified. We used electroencephalography (EEG) to measure the neural responses of participants while they performed a temporal bisection task in which they were requested to categorize the duration of visual stimuli as short or long.6 We found that the stimulus-offset and response-locked activity depends on both stimulus duration and the participants' decision. To relate this activity to the underlying cognitive processes, we used a drift-diffusion model.7 The model includes a noisy accumulator starting with the stimulus onset and a decision threshold. According to the model, a stimulus duration will be categorized as “long” if the accumulator reaches the threshold during stimulus presentation. Otherwise, it will be categorized as “short.” We found that at the offset of stimulus presentation, an EEG response marks the distance of the accumulator from the threshold. Therefore, this model offers an accurate description of our behavioral data as well as the EEG response using the same two model parameters. We then replicated this finding in an identical experiment conducted in the tactile domain. We also extended this finding to two different temporal ranges (sub- and supra-second). Taken together, the work provides a new way to study the cognitive processes underlying temporal decisions, using a combination of behavior, EEG, and modeling.
Sergio Osorio; Martín Irani; Javiera Herrada; Francisco Aboitiz
Neural responses to sensory novelty with and without conscious access Journal Article
In: NeuroImage, vol. 262, pp. 1–12, 2022.
Detection of novel stimuli that violate statistical regularities in the sensory scene is of paramount importance for the survival of biological organisms. Event-related potentials, phasic increases in pupil size, and evoked changes in oscillatory power have been proposed as markers of sensory novelty detection. However, how conscious access to novelty modulates these different brain responses is not well understood. Here, we studied the neural responses to sensory novelty in the auditory modality with and without conscious access. We identified individual thresholds for conscious auditory discrimination and presented to our participants sequences of tones, where the last stimulus could be another standard, a subthreshold target or a suprathreshold target. Participants were instructed to report whether the last tone of each sequence was the same or different from those preceding it. Results indicate that attentional orientation to behaviorally relevant stimuli and overt decision-making mechanisms, indexed by the P3 event-related response and reaction times, best predict whether a novel stimulus will be consciously accessed. Theta power and pupil size do not predict conscious access to novelty, but instead reflect information maintenance and unexpected sensory uncertainty. These results highlight the interplay between bottom-up and top-down mechanisms and how the brain weights neural responses to novelty and uncertainty during perception and goal-directed behavior.
Nadia Paraskevoudi; Iria SanMiguel
Sensory suppression and increased neuromodulation during actions disrupt memory encoding of unpredictable self-initiated stimuli Journal Article
In: Psychophysiology, vol. 60, pp. 1–25, 2022.
Actions modulate sensory processing by attenuating responses to self- compared to externally generated inputs, which is traditionally attributed to stimulus-specific motor predictions. Yet, suppression has been also found for stimuli merely coinciding with actions, pointing to unspecific processes that may be driven by neuromodulatory systems. Meanwhile, the differential processing for self-generated stimuli raises the possibility of producing effects also on memory for these stimuli; however, evidence remains mixed as to the direction of the effects. Here, we assessed the effects of actions on sensory processing and memory encoding of concomitant, but unpredictable sounds, using a combination of self-generation and memory recognition task concurrently with EEG and pupil recordings. At encoding, subjects performed button presses that half of the time generated a sound (motor-auditory; MA) and listened to passively presented sounds (auditory-only; A). At retrieval, two sounds were presented and participants had to respond which one was present before. We measured memory bias and memory performance by having sequences where either both or only one of the test sounds were presented at encoding, respectively. Results showed worse memory performance – but no differences in memory bias –, attenuated responses, and larger pupil diameter for MA compared to A sounds. Critically, the larger the sensory attenuation and pupil diameter, the worse the memory performance for MA sounds. Nevertheless, sensory attenuation did not correlate with pupil dilation. Collectively, our findings suggest that sensory attenuation and neuromodulatory processes coexist during actions, and both relate to disrupted memory for concurrent, albeit unpredictable sounds.
Xinzhen Pei; Guiying Xu; Yunhui Zhou; Luna Tao; Xiaozhu Cui; Zhenyu Wang; Bingru Xu; An-Li Wang; Xi Zhao; Haijun Dong; Yan An; Yang Cao; Ruxue Li; Honglin Hu; Yuguo Yu
A simultaneous electroencephalography and eye-tracking dataset in elite athletes during alertness and concentration tasks Journal Article
In: Scientific Data, vol. 9, pp. 1–15, 2022.
The dataset of simultaneous 64-channel electroencephalography (EEG) and high-speed eye-tracking (ET) recordings was collected from 31 professional athletes and 43 college students during alertness behavior task (ABT) and concentration cognitive task (CCT). The CCT experiment lasting 1–2 hours included five sessions for groups of the Shooting, Archery and Modern Pentathlon elite athletes and the controls. Concentration targets included shooting target and combination target with or without 24 different directions of visual distractors and 2 types of music distractors. Meditation and Schulte Grid trainings were done as interventions. Analysis of the dataset aimed to extract effective biological markers of eye movement and EEG that can assess the concentration level of talented athletes compared with same-aged controls. Moreover, this dataset is useful for the research of related visual brain-computer interfaces.
Thomas Pfeffer; Christian Keitel; Daniel S. Kluger; Anne Keitel; Alena Russmann; Gregor Thut; Tobias H. Donner; Joachim Gross
Coupling of pupil-and neuronal population dynamics reveals diverse influences of arousal on cortical processing Journal Article
In: eLife, vol. 11, pp. 1–28, 2022.
Fluctuations in arousal, controlled by subcortical neuromodulatory systems, continuously shape cortical state, with profound consequences for information processing. Yet, how arousal signals influence cortical population activity in detail has so far only been characterized for a few selected brain regions. Traditional accounts conceptualize arousal as a homogeneous modulator of neural population activity across the cerebral cortex. Recent insights, however, point to a higher specificity of arousal effects on different components of neural activity and across cortical regions. Here, we provide a comprehensive account of the relationships between fluctuations in arousal and neuronal population activity across the human brain. Exploiting the established link between pupil size and central arousal systems, we performed concurrent magnetoencephalographic (MEG) and pupillographic recordings in a large number of participants, pooled across three laboratories. We found a cascade of effects relative to the peak timing of spontaneous pupil dilations: Decreases in low-frequency (2–8 Hz) activity in temporal and lateral frontal cortex, followed by increased highfrequency (>64 Hz) activity in mid-frontal regions, followed by monotonic and inverted U relationships with intermediate frequency-range activity (8–32 Hz) in occipito-parietal regions. Pupil-linked arousal also coincided with widespread changes in the structure of the aperiodic component of cortical population activity, indicative of changes in the excitation-inhibition balance in underlying microcircuits. Our results provide a novel basis for studying the arousal modulation of cognitive computations in cortical circuits.
Mattia Pietrelli; Jason Samaha; Bradley R. Postle
Spectral distribution dynamics across different attentional priority states Journal Article
In: Journal of Neuroscience, vol. 42, no. 19, pp. 4026–4041, 2022.
Anticipatory covert spatial attention improves performance on tests of visual detection and discrimination, and shifts are accompanied by decreases and increases of a band power at electroencephalography (EEG) electrodes corresponding to the attended and unattended location, respectively. Although the increase at the unattended location is often interpreted as an active mechanism (e.g., inhibiting processing at the unattended location), most experiments cannot rule out the alternative possibility that it is a secondary consequence of selection elsewhere. To adjudicate between these accounts, we designed a Posner- style visual cueing task in which male and female human participants made orientation judgments of targets appearing at one of four locations: up, down, right, or left. Critically, trials were blocked such that within a block the locations along one meridian alternated in status between attended and unattended, and targets never appeared at the other two, making them irrelevant. Analyses of the concurrently measured EEG signal were conducted on “traditional” narrowband a (8–14 Hz), as well as on two components resulting from the decomposition of this signal: “periodic” a;and the slope of the aperiodic 1/f-like component. Although data from right-left blocks replicated the familiar pattern of lateralized asymmetry in narrowband a power, with neither a signal couldwe findevidence for any difference inthe time course at unattended versus irrelevant locations, an outcome consistent with the secondary-consequence interpretation of attention- related dynamics in the a band. Additionally, 1/f slope was shallower at attended and unattended locations, relative to irrelevant, suggesting a tonic adjustment of physiological state.
Michael Plöchl; Ian Fiebelkorn; Sabine Kastner; Jonas Obleser
Attentional sampling of visual and auditory objects is captured by theta-modulated neural activity Journal Article
In: European Journal of Neuroscience, vol. 55, no. 11-12, pp. 3067–3082, 2022.
Recent evidence suggests that visual attention alternately samples two behaviourally relevant objects at approximately 4 Hz, rhythmically shifting between the objects. Whether similar attentional rhythms exist in other sensory modalities, however, is not yet clear. We therefore adapted and extended an established paradigm to investigate visual and potential auditory attentional rhythms, as well as possible interactions, on both a behavioural (detection performance
Tzvetan Popov; Bart Gips; Nathan Weisz; Ole Jensen
Brain areas associated with visual spatial attention display topographic organization during auditory spatial attention Journal Article
In: Cerebral Cortex, pp. 1–12, 2022.
Spatially selective modulation of alpha power (8–14 Hz) is a robust finding in electrophysiological studies of visual attention, and has been recently generalized to auditory spatial attention. This modulation pattern is interpreted as reflecting a top-down mechanism for suppressing distracting input from unattended directions of sound origin. The present study on auditory spatial attention extends this interpretation by demonstrating that alpha power modulation is closely linked to oculomotor action. We designed an auditory paradigm in which participants were required to attend to upcoming sounds from one of 24 loudspeakers arranged in a circular array around the head. Maintaining the location of an auditory cue was associated with a topographically modulated distribution of posterior alpha power resembling the findings known from visual attention. Multivariate analyses allowed the prediction of the sound location in the horizontal plane. Importantly, this prediction was also possible, when derived from signals capturing saccadic activity. A control experiment on auditory spatial attention confirmed that, in absence of any visual/auditory input, lateralization of alpha power is linked to the lateralized direction of gaze. Attending to an auditory target engages oculomotor and visual cortical areas in a topographic manner akin to the retinotopic organization associated with visual attention.
Frida A. B. Printzlau; Nicholas E. Myers; Sanjay G. Manohar; Mark G. Stokes
Neural reinstatement tracks spread of attention between object features in working memory Journal Article
In: Journal of Cognitive Neuroscience, vol. 34, no. 9, pp. 1681–1701, 2022.
Attention can be allocated in working memory ( WM) to select and privilege relevant content. It is unclear whether attention selects individual features or whole objects in WM. Here, we used behavioral measures, eye-tracking, and EEG to test the hypothesis that attention spreads between an object's features in WM. Twenty-six participants completed a WM task that asked them to recall the angle of one of two oriented, colored bars after a delay while EEG and eye-tracking data were collected. During the delay, an orthogonal “incidental task” cued the color of one item for a match/mismatch judgment. On congruent trials (50%), the cued item was probed for subsequent orientation recall; on incongruent trials (50%), the other memory item was probed. As predicted, selecting the color of an object in WM brought other features of the cued object into an attended state as revealed by EEG decoding, oscillatory $alpha$-power, gaze bias, and improved orientation recall performance. Together, the results show that attentional selection spreads between an object's features in WM, consistent with object-based attentional selection. Analyses of neural processing at recall revealed that the selected object was automatically compared with the probe, whether it was the target for recall or not. This provides a potential mechanism for the observed benefits of nonpredictive cueing in WM, where a selected item is prioritized for subsequent decision-making.
Estelle Raffin; Adrien Witon; Roberto F. Salamanca-Giron; Krystel R. Huxlin; Friedhelm C. Hummel
Functional segregation within the dorsal frontoparietal network: A multimodal dynamic causal modeling study Journal Article
In: Cerebral Cortex, vol. 32, no. 15, pp. 3187–3205, 2022.
Discrimination and integration of motion direction requires the interplay of multiple brain areas. Theoretical accounts of perception suggest that stimulus-related (i.e., exogenous) and decision-related (i.e., endogenous) factors affect distributed neuronal processing at different levels of the visual hierarchy. To test these predictions, we measured brain activity of healthy participants during a motion discrimination task, using electroencephalography (EEG) and functional magnetic resonance imaging (fMRI). We independently modeled the impact of exogenous factors (task demand) and endogenous factors (perceptual decision-making) on the activity of the motion discrimination network and applied Dynamic Causal Modeling (DCM) to both modalities. DCM for event-related potentials (DCM-ERP) revealed that task demand impacted the reciprocal connections between the primary visual cortex (V1) and medial temporal areas (V5). With practice, higher visual areas were increasingly involved, as revealed by DCM-fMRI. Perceptual decision-making modulated higher levels (e.g., V5-to-Frontal Eye Fields, FEF), in a manner predictive of performance. Our data suggest that lower levels of the visual network support early, feature-based selection of responses, especially when learning strategies have not been implemented. In contrast, perceptual decision-making operates at higher levels of the visual hierarchy by integrating sensory information with the internal state of the subject.
Kati Roesmann; Ida Wessing; Sophia Kraß; Elisabeth J. Leehr; Tim Klucken; Thomas Straube; Markus Junghöfer
Developmental aspects of fear generalization – A MEG study on neurocognitive correlates in adolescents versus adults Journal Article
In: Developmental Cognitive Neuroscience, vol. 58, pp. 1–12, 2022.
Background: Fear generalization is pivotal for the survival-promoting avoidance of potential danger, but, if too pronounced, it promotes pathological anxiety. Similar to adult patients with anxiety disorders, healthy children tend to show overgeneralized fear responses. Objective: This study aims to investigate neuro-developmental aspects of fear generalization in adolescence – a critical age for the development of anxiety disorders. Methods: We compared healthy adolescents (14–17 years) with healthy adults (19–34 years) regarding their fear responses towards tilted Gabor gratings (conditioned stimuli, CS; and slightly differently titled generalization stimuli, GS). In the conditioning phase, CS were paired (CS+) or remained unpaired (CS-) with an aversive stimulus (unconditioned stimuli, US). In the test phase, behavioral, peripheral and neural responses to CS and GS were captured by fear- and UCS expectancy ratings, a perceptual discrimination task, pupil dilation and source estimations of event-related magnetic fields. Results: Closely resembling adults, adolescents showed robust generalization gradients of fear ratings, pupil dilation, and estimated neural source activity. However, in the UCS expectancy ratings, adolescents revealed shallower generalization gradients indicating overgeneralization. Moreover, adolescents showed stronger visual cortical activity after as compared to before conditioning to all stimuli. Conclusion: Various aspects of fear learning and generalization appear to be mature in healthy adolescents. Yet, cognitive aspects might show a slower course of development.
Amirsaman Sajad; Steven P. Errington; Jeffrey D. Schall
Functional architecture of executive control and associated event-related potentials in macaques Journal Article
In: Nature Communications, vol. 13, no. 1, pp. 1–19, 2022.
The medial frontal cortex (MFC) enables executive control by monitoring relevant information and using it to adapt behavior. In macaques performing a saccade countermanding (stop-signal) task, we simultaneously recorded electrical potentials over MFC and neural spiking across all layers of the supplementary eye field (SEF). We report the laminar organization of neurons enabling executive control by monitoring the conflict between incompatible responses, the timing of events, and sustaining goal maintenance. These neurons were a mix of narrow-spiking and broad-spiking found in all layers, but those predicting the duration of control and sustaining the task goal until the release of operant control were more commonly narrow-spiking neurons confined to layers 2 and 3 (L2/3). We complement these results with evidence for a monkey homolog of the N2/P3 event-related potential (ERP) complex associated with response inhibition. N2 polarization varied with error-likelihood and P3 polarization varied with the duration of expected control. The amplitude of the N2 and P3 were predicted by the spike rate of different classes of neurons located in L2/3 but not L5/6. These findings reveal features of the cortical microcircuitry supporting executive control and producing associated ERPs.
Delia A. Gheorghe; Muriel T. N. Panouillères; Nicholas D. Walsh
Investigating the effects of cerebellar transcranial direct current stimulation on saccadic adaptation and cortisol response Journal Article
In: Cerebellum and Ataxias, vol. 8, no. 1, pp. 1–11, 2021.
Background: Transcranial Direct Current Stimulation (tDCS) over the prefrontal cortex has been shown to modulate subjective, neuronal and neuroendocrine responses, particularly in the context of stress processing. However, it is currently unknown whether tDCS stimulation over other brain regions, such as the cerebellum, can similarly affect the stress response. Despite increasing evidence linking the cerebellum to stress-related processing, no studies have investigated the hormonal and behavioural effects of cerebellar tDCS. Methods: This study tested the hypothesis of a cerebellar tDCS effect on mood, behaviour and cortisol. To do this we employed a single-blind, sham-controlled design to measure performance on a cerebellar-dependent saccadic adaptation task, together with changes in cortisol output and mood, during online anodal and cathodal stimulation. Forty-five participants were included in the analysis. Stimulation groups were matched on demographic variables, potential confounding factors known to affect cortisol levels, mood and a number of personality characteristics. Results: Results showed that tDCS polarity did not affect cortisol levels or subjective mood, but did affect behaviour. Participants receiving anodal stimulation showed an 8.4% increase in saccadic adaptation, which was significantly larger compared to the cathodal group (1.6%). Conclusion: The stimulation effect on saccadic adaptation contributes to the current body of literature examining the mechanisms of cerebellar stimulation on associated function. We conclude that further studies are needed to understand whether and how cerebellar tDCS may module stress reactivity under challenge conditions.
Guanpeng Chen; Ziyun Zhu; Qing He; Fang Fang
Offline transcranial direct current stimulation improves the ability to perceive crowded targets Journal Article
In: Journal of Vision, vol. 21, no. 2, pp. 1–10, 2021.
The deleterious effect of nearby flankers on target identification in the periphery is known as visual crowding. Studying visual crowding can advance our understanding of the mechanisms of visual awareness and object recognition. Alleviating visual crowding is one of the major ways to improve peripheral vision. The aim of the current study was to examine whether transcranial direct current stimulation (tDCS) was capable of alleviating visual crowding at different visual eccentricities and with different visual tasks. In the present single-blind sham-controlled study, subjects were instructed to perform an orientation discrimination task or a letter identification task with isolated and crowded targets in the periphery, before and after applying 20 minutes of 2 mA anodal tDCS to visual cortex of the hemisphere contralateral or ipsilateral to visual stimuli. Contralateral tDCS significantly alleviated the orientation crowding effect at two different eccentricities and the letter crowding effect. This alleviation was absent after sham or ipsilateral stimulation and could not be fully explained by the performance improvement with the isolated targets. These findings demonstrated that offline tDCS was effective in alleviating visual crowding across different visual eccentricities and tasks, therefore providing a promising way to improve spatial vision rapidly in crowded scenes.
Andra Coldea; Stephanie Morand; Domenica Veniero; Monika Harvey; Gregor Thut
Parietal alpha tACS shows inconsistent effects on visuospatial attention Journal Article
In: PLoS ONE, vol. 16, no. 8, pp. e0255424, 2021.
Transcranial alternating current stimulation (tACS) is a popular technique that has been used for manipulating brain oscillations and inferring causality regarding the brain-behaviour relationship. Although it is a promising tool, the variability of tACS results has raised questions regarding the robustness and reproducibility of its effects. Building on recent research using tACS to modulate visuospatial attention, we here attempted to replicate findings of lateralized parietal tACS at alpha frequency to induce a change in attention bias away from the contra- towards the ipsilateral visual hemifield. 40 healthy participants underwent tACS in two separate sessions where either 10 Hz tACS or sham was applied via a high-density montage over the left parietal cortex at 1.5 mA for 20 min, while performance was assessed in an endogenous attention task. Task and tACS parameters were chosen to match those of previous studies reporting positive effects. Unlike these studies, we did not observe lateralized parietal alpha tACS to affect attention deployment or visual processing across the hemifields as compared to sham. Likewise, additional resting electroencephalography immediately offline to tACS did not reveal any notable effects on individual alpha power or frequency. Our study emphasizes the need for more replication studies and systematic investigations of the factors that drive tACS effects.
Raymundo Machado Azevedo Neto; Andreas Bartels
Disrupting short-term memory maintenance in premotor cortex affects serial dependence in visuomotor integration Journal Article
In: Journal of Neuroscience, vol. 41, no. 45, pp. 9392–9402, 2021.
Human behavior is biased by past experience. For example, when intercepting a moving target, the speed of previous targets will bias responses in future trials. Neural mechanisms underlying this so-called serial dependence are still under debate. Here, we tested the hypothesis that the previous trial leaves a neural trace in brain regions associated with encoding task-relevant information in visual and/or motor regions. We reasoned that injecting noise by means of transcranial magnetic stimulation (TMS) over premotor and visual areas would degrade such memory traces and hence reduce serial dependence. To test this hypothesis, we applied bursts of TMS pulses to right visual motion processing region hV5/MT1 and to left dorsal premotor cortex (PMd) during intertrial intervals of a coincident timing task performed by twenty healthy human participants (15 female). Without TMS, participants presented a bias toward the speed of the previous trial when intercepting moving targets. TMS over PMd decreased serial dependence in comparison to the control Vertex stimulation, whereas TMS applied over hV5/MT1 did not. In addition, TMS seems to have specifically affected the memory trace that leads to serial dependence, as we found no evidence that participants' behavior worsened after applying TMS. These results provide causal evidence that an implicit short-term memory mechanism in premotor cortex keeps information from one trial to the next, and that this information is blended with current trial information so that it biases behavior in a visuomotor integration task with moving objects.
P. J. Hills; G. Arabacı; J. Fagg; L. Canter; C. Thompson; R. Moseley
Low-frequency rTMS to the parietal lobe increases eye-movement carryover and decreases hazard rating Journal Article
In: Neuropsychologia, vol. 158, pp. 107895, 2021.
The persistence of attentional set from one task to a secondary unrelated task, revealed through carryover of eye movements, has been attributed to increased activation in the parietal lobe and decreased activation to the frontal lobe. To directly test this, we adopted a modified version of the Thompson and Crundall (2011) paradigm using low-frequency repetitive TMS to P3 and F3. In each trial, participants viewed letter-strings that were arranged horizontally, vertically, or randomly across the screen before viewing a road image and providing a hazardousness rating for it. The orientation of the letter search influenced eye movements to the road images and this carryover was greater following stimulation to F3 than to P3 (or sham). Furthermore, hazardous ratings were lower following P3 stimulation. These results confirm the involvement of attentional orienting and switching mechanisms in the carryover of eye movements. It is suggested that this “attentional inertia” effect will increase with greater orienting of attentional resources in an initial task and poor inhibition of previously-relevant settings between tasks.