EyeLink EEG / fNIRS / TMS Publications
All EyeLink EEG, fNIRS, and TMS research publications (with concurrent eye tracking) up until 2023 (with early 2024s) are listed below by year. You can search the publications using keywords such as P300, Gamma band, NIRS, etc. You can also search for individual author names. If we missed any EyeLink EEG, fNIRS, or TMS articles, please email us!
2017 |
James Mathew; Alexandre Eusebio; Frederic R. Danion Limited contribution of primary motor cortex in eye-hand coordination: A TMS study Journal Article In: Journal of Neuroscience, vol. 37, no. 40, pp. 9730 –9740, 2017. @article{Mathew2017, The ability to track a moving target with the eye is substantially improved when the target is self-moved compared with when it is moved by an external agent. To account for this observation, it has been postulated that the oculomotor system has access to hand efference copy, thereby allowing to predict the motion of the visual target. Along this scheme, we tested the effect of transcranial magnetic stimulation (TMS) over the hand area of the primary motor cortex (M1) when human participants (50% females) are asked to track with their eyes a visual target whose horizontal motion is driven by their grip force. We reasoned that, if the output of M1 is used by the oculomotor system to keep track of the target, on top of inducing short latency disturbance of grip force, single-pulse TMS should also quickly disrupt ongoing eye motion. For comparison purposes, the effect of TMS over M1 was monitored when subjects tracked an externally moved target (while keeping their hand at rest or not). In both cases, results showed no alterations in smooth pursuit, meaning that its velocity was unaffected within the 25-125 ms epoch that followed TMS. Overall, our results imply that the output of M1 has limited contribution in driving the eye motion during our eye-hand coordination task. This study suggests that, if hand motor signals are accessed by the oculomotor system, this is upstream of M1. |
Rebecca E. Paladini; René M. Müri; Jurka Meichtry; Tobias Nef; Fred W. Mast; Urs P. Mosimann; Thomas Nyffeler; Dario Cazzoli The influence of alertness on the spatial deployment of visual attention is mediated by the excitability of the posterior parietal cortices Journal Article In: Cerebral Cortex, vol. 27, no. 1, pp. 233–243, 2017. @article{Paladini2017, With a reduced level of alertness, healthy individuals typically show a rightward shift when deploying visual attention in space. The impact of alertness on the neural networks governing visuospatial attention is, however, poorly understood. By using a transcranial magnetic stimulation twin-coil approach, the present study aimed at investigating the effects of an alertness manipulation on the excitability of the left and the right posterior parietal cortices (PPCs), crucial nodes of the visuospatial attentional network. Participants' visuospatial attentional deployment was assessed with a free visual exploration task and concurrent eye tracking. Their alertness level was manipulated through the time of the day, that is, by testing chronotypically defined evening types both during their circadian on- and off-peak times. The results revealed an increased excitability of the left compared with the right PPC during low alertness. On the horizontal dimension, these results were accompanied by a significant rightward shift in the center and a bilateral narrowing in the periphery of the visual exploration field, as well as a central upward shift on the vertical dimension. The findings show that the manipulation of non-spatial attentional aspects (i.e., alertness) can affect visuospatial attentional deployment and modulate the excitability of areas subtending spatial attentional control. |
Paul Metzner; Titus Malsburg; Shravan Vasishth; Frank Rösler The importance of reading naturally: Evidence from combined recordings of eye movements and electric brain potentials Journal Article In: Cognitive Science, vol. 41, pp. 1232–1263, 2017. @article{Metzner2017, How important is the ability to freely control eye movements for reading comprehension? And how does the parser make use of this freedom? We investigated these questions using coregistration of eye movements and event-related brain potentials (ERPs) while participants read either freely or in a computer-controlled word-by-word format (also known as RSVP). Word-by-word presentation and natural reading both elicited qualitatively similar ERP effects in response to syntactic and semantic violations (N400 and P600 effects). Comprehension was better in free reading but only in trials in which the eyes regressed to previous material upon encountering the anomaly. A more fine-grained ERP analysis revealed that these regressions were strongly associated with the well-known P600 effect. In trials without regressions, we instead found sustained centro-parietal negativities starting at around 320 ms post-onset; however, these negativities were only found when the violation occurred in sentence-final position. Taken together, these results suggest that the sentence processing system engages in strategic choices: In response to words that don't match built-up expectations, it can either explore alternative interpretations (reflected by regressions, P600 effects, and good com-prehension) or pursue a " good-enough " processing strategy that tolerates a deficient interpretation (reflected by progressive saccades, sustained negativities, and relatively poor comprehension). |
Kelly Miles; Catherine M. McMahon; Isabelle Boisvert; Ronny Ibrahim; Peter Lissa; Petra L. Graham; Björn Lyxell Objective assessment of listening effort: Coregistration of pupillometry and EEG Journal Article In: Trends in Hearing, vol. 21, 2017. @article{Miles2017, Listening to speech in noise is effortful, particularly for people with hearing impairment. While it is known that effort is related to a complex interplay between bottom-up and top-down processes, the cognitive and neurophysiological mechan-isms contributing to effortful listening remain unknown. Therefore, a reliable physiological measure to assess effort remains elusive. This study aimed to determine whether pupil dilation and alpha power change, two physiological measures suggested to index listening effort, assess similar processes. Listening effort was manipulated by parametrically varying spectral reso-lution (16-and 6-channel noise vocoding) and speech reception thresholds (SRT; 50% and 80%) while 19 young, normal-hearing adults performed a speech recognition task in noise. Results of off-line sentence scoring showed discrepancies between the target SRTs and the true performance obtained during the speech recognition task. For example, in the SRT80% condition, participants scored an average of 64.7%. Participants' true performance levels were therefore used for subsequent statistical modelling. Results showed that both measures appeared to be sensitive to changes in spectral reso-lution (channel vocoding), while pupil dilation only was also significantly related to their true performance levels (%) and task accuracy (i.e., whether the response was correctly or partially recalled). The two measures were not correlated, suggesting they each may reflect different cognitive processes involved in listening effort. This combination of findings contributes to a growing body of research aiming to develop an objective measure of listening effort. |
Sorato Minami; Kaoru Amano Illusory jitter perceived at the frequency of alpha oscillations Journal Article In: Current Biology, vol. 27, no. 15, pp. 1–13, 2017. @article{Minami2017, Neural oscillations, such as alpha (8–13 Hz), beta (13–30 Hz), and gamma (30–100 Hz), are widespread across cortical areas, and their possible functional roles include feature binding [1], neuronal communication [2, 3], and memory [1, 4]. The most prominent signal among these neural oscillations is the alpha oscillation. Although accumulating evidence suggests that alpha oscillations correlate with various aspects of visual processing [5–18], the number of studies proving their causal contribution in visual perception is limited [11, 16–18]. Here we report that illusory visual vibrations are consciously experienced at the frequency of intrinsic alpha oscillations. We employed an illusory jitter perception termed the motion-induced spatial conflict [19] that originates from the cyclic interaction between motion and shape processing. Comparison between the perceived frequency of illusory jitter and the peak alpha frequency (PAF) measured using magnetoencephalography (MEG) revealed that the inter- and intra-participant variations of the PAF are mirrored by an illusory jitter perception. More crucially, psychophysical and MEG measurements during amplitude-modulated current stimulation [20] showed that the PAF can be artificially manipulated, which results in a corresponding change in the perceived jitter frequency. These results suggest the causal contribution of neural oscillations at the alpha frequency in creating temporal characteristics of visual perception. Our results suggest that cortical areas, dorsal and ventral visual areas in this case, are interacting at the frequency of alpha oscillations [2, 3, 21–27]. |
Roxane J. Itier; Karly N. Neath-Tavares Effects of task demands on the early neural processing of fearful and happy facial expressions Journal Article In: Brain Research, vol. 1663, pp. 38–50, 2017. @article{Itier2017, Task demands shape how we process environmental stimuli but their impact on the early neural processing of facial expressions remains unclear. In a within-subject design, ERPs were recorded to the same fearful, happy and neutral facial expressions presented during a gender discrimination, an explicit emotion discrimination and an oddball detection tasks, the most studied tasks in the field. Using an eye tracker, fixation on the face nose was enforced using a gaze-contingent presentation. Task demands modulated amplitudes from 200 to 350 ms at occipito-temporal sites spanning the EPN component. Amplitudes were more negative for fearful than neutral expressions starting on N170 from 150 to 350 ms, with a temporo-occipital distribution, whereas no clear effect of happy expressions was seen. Task and emotion effects never interacted in any time window or for the ERP components analyzed (P1, N170, EPN). Thus, whether emotion is explicitly discriminated or irrelevant for the task at hand, neural correlates of fearful and happy facial expressions seem immune to these task demands during the first 350 ms of visual processing. |
Syaheed B. Jabar; Alex Filipowicz; Britt Anderson Tuned by experience: How orientation probability modulates early perceptual processing Journal Article In: Vision Research, vol. 138, pp. 86–96, 2017. @article{Jabar2017a, Probable stimuli are more often and more quickly detected. While stimulus probability is known to affect decision-making, it can also be explained as a perceptual phenomenon. Using spatial gratings, we have previously shown that probable orientations are also more precisely estimated, even while participants remained naive to the manipulation. We conducted an electrophysiological study to investigate the effect that probability has on perception and visual-evoked potentials. In line with previous studies on oddballs and stimulus prevalence, low-probability orientations were associated with a greater late positive ‘P300' component which might be related to either surprise or decision-making. However, the early ‘C1' component, thought to reflect V1 processing, was dampened for high-probability orientations while later P1 and N1 components were unaffected. Exploratory analyses revealed a participant-level correlation between C1 and P300 amplitudes, suggesting a link between perceptual processing and decision-making. We discuss how these probability effects could be indicative of sharpening of neurons preferring the probable orientations, due either to perceptual learning, or to feature-based attention. |
Jianrong Jia; Ling Liu; Fang Fang; Huan Luo Sequential sampling of visual objects during sustained attention Journal Article In: PLoS Biology, vol. 15, no. 6, pp. e2001903, 2017. @article{Jia2017b, In a crowded visual scene, attention must be distributed efficiently and flexibly over time and space to accommodate different contexts. It is well established that selective attention enhances the corresponding neural responses, presumably implying that attention would persistently dwell on the task-relevant item. Meanwhile, recent studies, mostly in divided attentional contexts, suggest that attention does not remain stationary but samples objects alternately over time, suggesting a rhythmic view of attention. However, it remains unknown whether the dynamic mechanism essentially mediates attentional processes at a general level. Importantly, there is also a complete lack of direct neural evidence reflecting whether and how the brain rhythmically samples multiple visual objects during stimulus processing. To address these issues, in this study, we employed electroencephalography (EEG) and a temporal response function (TRF) approach, which can dissociate responses that exclusively represent a single object from the overall neuronal activity, to examine the spatiotemporal characteristics of attention in various attentional contexts. First, attention, which is characterized by inhibitory alpha-band (approximately 10 Hz) activity in TRFs, switches between attended and unattended objects every approximately 200 ms, suggesting a sequential sampling even when attention is required to mostly stay on the attended object. Second, the attentional spatiotemporal pattern is modulated by the task context, such that alpha-mediated switching becomes increasingly prominent as the task requires a more uniform distribution of attention. Finally, the switching pattern correlates with attentional behavioral performance. Our work provides direct neural evidence supporting a generally central role of temporal organization mechanism in attention, such that multiple objects are sequentially sorted according to their priority in attentional contexts. The results suggest that selective attention, in addition to the classically posited attentional “focus,” involves a dynamic mechanism for monitoring all objects outside of the focus. Our findings also suggest that attention implements a space (object)-to-time transformation by acting as a series of concatenating attentional chunks that operate on 1 object at a time. |
Elizabeth L. Johnson; Callum D. Dewar; Anne Kristin Solbakk; Tor Endestad; Torstein R. Meling; Robert T. Knight Bidirectional frontoparietal oscillatory systems support working memory Journal Article In: Current Biology, vol. 27, no. 12, pp. 1829–1835, 2017. @article{Johnson2017a, The ability to represent and select information in working memory provides the neurobiological infrastructure for human cognition. For 80 years, dominant views of working memory have focused on the key role of prefrontal cortex (PFC) [1–8]. However, more recent work has implicated posterior cortical regions [9–12], suggesting that PFC engagement during working memory is dependent on the degree of executive demand. We provide evidence from neurological patients with discrete PFC damage that challenges the dominant models attributing working memory to PFC-dependent systems. We show that neural oscillations, which provide a mechanism for PFC to communicate with posterior cortical regions [13], independently subserve communications both to and from PFC—uncovering parallel oscillatory mechanisms for working memory. Fourteen PFC patients and 20 healthy, age-matched controls performed a working memory task where they encoded, maintained, and actively processed information about pairs of common shapes. In controls, the electroencephalogram (EEG) exhibited oscillatory activity in the low-theta range over PFC and directional connectivity from PFC to parieto-occipital regions commensurate with executive processing demands. Concurrent alpha-beta oscillations were observed over parieto-occipital regions, with directional connectivity from parieto-occipital regions to PFC, regardless of processing demands. Accuracy, PFC low-theta activity, and PFC → parieto-occipital connectivity were attenuated in patients, revealing a PFC-independent, alpha-beta system. The PFC patients still demonstrated task proficiency, which indicates that the posterior alpha-beta system provides sufficient resources for working memory. Taken together, our findings reveal neurologically dissociable PFC and parieto-occipital systems and suggest that parallel, bidirectional oscillatory systems form the basis of working memory. |
Tim C. Kietzmann; Anna L. Gert; Frank Tong; Peter König Representational dynamics of facial viewpoint encoding Journal Article In: Journal of Cognitive Neuroscience, vol. 29, no. 4, pp. 637–651, 2017. @article{Kietzmann2017, Faces provide a wealth of information, including the identity of the seen person and social cues, such as the direction of gaze. Crucially, different aspects of face processing require distinct forms of information encoding. Another person's attentional focus can be derived based on a view-dependent code. In contrast, identification benefits from invariance across all view-points. Different cortical areas have been suggested to subserve these distinct functions. However, little is known about the temporal aspects of differential viewpoint encoding in the human brain. Here, we combine EEG with multivariate data analyses to resolve the dynamics of face processing with high temporal resolution. This revealed a distinct sequence of view-point encoding. Head orientations were encoded first, starting after around 60 msec of processing. Shortly afterward, peaking around 115 msec after stimulus onset, a different encoding scheme emerged. At this latency, mirror-symmetric viewing angles elicited highly similar cortical responses. Finally, about 280 msec after visual onset, EEG response patterns demon-strated a considerable degree of viewpoint invariance across all viewpoints tested, with the noteworthy exception of the front-facing view. Taken together, our results indicate that the processing of facial viewpoints follows a temporal sequence of encoding schemes, potentially mirroring different levels of computational complexity. |
Emmanuelle Kristensen; Anne Guerin-Dugué; Bertrand Rivet Regularization and a general linear model for event-related potential estimation Journal Article In: Behavior Research Methods, vol. 49, no. 6, pp. 2255–2274, 2017. @article{Kristensen2017, The usual event-related potential (ERP) estimation is the average across epochs time-locked on stimuli of interest. These stimuli are repeated several times to improve the signal-to-noise ratio (SNR) and only one evoked potential is estimated inside the temporal window of interest. Consequently, the average estimation does not take into account other neural responses within the same epoch that are due to short inter stimuli intervals. These adjacent neural responses may overlap and distort the evoked potential of interest. This overlapping process is a significant issue for the eye fixation-related potential (EFRP) technique in which the epochs are time-locked on the ocular fixations. The inter fixation intervals are not experimentally controlled and can be shorter than the neural response's latency. To begin, the Tikhonov regularization, applied to the classical average estimation, was introduced to improve the SNR for a given number of trials. The generalized cross validation was chosen to obtain the optimal value of the ridge parameter. Then, to deal with the issue of overlapping, the general linear model (GLM), was used to extract all neural responses inside an epoch. Finally, the regularization was also applied to it. The models (the classical average and the GLM with and without regularization) were compared on both simulated data and real datasets from a visual scene exploration in co-registration with an eye-tracker, and from a P300 Speller experiment. The regularization was found to improve the estimation by average for a given number of trials. The GLM was more robust and efficient, its efficiency actually reinforced by the regularization. |
Emmanuelle Kristensen; Bertrand Rivet; Anne Guérin-Dugué Estimation of overlapped Eye Fixation Related Potentials: The General Linear Model, a more flexible framework than the ADJAR algorithm Journal Article In: Journal of Eye Movement Research, vol. 10, no. 1, pp. 1–27, 2017. @article{Kristensen2017a, The Eye Fixation Related Potential (EFRP) estimation is the average of EEG signals across epochs at ocular fixation onset. Its main limitation is the overlapping issue. Inter Fixation Intervals (IFI) - typically around 300 ms in the case of unrestricted eye movement- depend on participants' oculomotor patterns, and can be shorter than the latency of the components of the evoked potential. If the duration of an epoch is longer than the IFI value, more than one fixation can occur, and some overlapping between adjacent neural responses ensues. The classical average does not take into account either the presence of several fixations during an epoch or overlapping. The Adjacent Response algorithm (ADJAR), which is popular for event-related potential estimation, was compared to the General Linear Model (GLM) on a real dataset from a conjoint EEG and eye-tracking experiment to address the overlapping issue. The results showed that the ADJAR algorithm was based on assumptions that were too restrictive for EFRP estimation. The General Linear Model appeared to be more robust and efficient. Different configurations of this model were compared to estimate the potential elicited at image onset, as well as EFRP at the beginning of exploration. These configurations took into account the overlap between the event-related potential at stimulus presentation and the following EFRP, and the distinction between the potential elicited by the first fixation onset and subsequent ones. The choice of the General Linear Model configuration was a tradeoff between assumptions about expected behavior and the quality of the EFRP estimation: the number of different potentials estimated by a given model must be controlled to avoid erroneous estimations with large variances. |
Qingxiao Liu; Bo Tan; Jing Zhou; Zhong Zheng; Ling Li; Yanchun Yang Pathophysiology of refractory obsessive-compulsive disorder Journal Article In: Medicine, pp. 1–11, 2017. @article{Liu2017d, Based on both functional and structural studies of excessive activity, fronto-striatal-thalamic-cortical and cortico-striatal circuits have been hypothesized to underlie the pathophysiology of obsessive-compulsive disorder (OCD). However, the neurobiological underpinnings of OCD refractory to medication and therapy remain controversial. This study aimed to evaluate neuroanatomical abnormalities of the whole brain and to evaluate visual processing in patients with refractory OCD. This study was comprised of 2 experiments. The neuroanatomical abnormalities of the whole brain were evaluated using a visual search in combination with overactive performance monitoring (Experiment I), and visual processing was evaluated using event- related potentials recorded from subjects during performance of a visual search task. We also examined the amplitudes and latency of the error-related negativity (ERN) using a modified flanker task (Experiment II). Standard low-resolution electromagnetic tomography analysis was applied to determine the special areas. Patients with refractory OCD had a significantly greater number of saccades and prolonged latencies relative to the healthy controls. Scalp map topography confirmed that visual cognitive and executive dysfunction was localized to the fusiform gyrus. Furthermore, we found that during a modified flanker task, ERNs had a greater amplitude and a prolonged latency relative to those of the healthy controls. Further data analysis suggested that cognitive dysfunction and compulsive behavior in OCD patients were linked to abnormalities within the dorsolateral prefrontal cortex (DLPFC). We identified abnormal activities within the fusiform gyrus and DLPFC that likely play important roles in the pathophysiology of OCD. |
Anna Marzecová; Andreas Widmann; Iria SanMiguel; Sonja A. Kotz; Erich Schröger Interrelation of attention and prediction in visual processing: Effects of task-relevance and stimulus probability Journal Article In: Biological Psychology, vol. 125, pp. 76–90, 2017. @article{Marzecova2017, The potentially interactive influence of attention and prediction was investigated by measuring event-related potentials (ERPs) in a spatial cueing task with attention (task-relevant) and prediction (probabilistic) cues. We identified distinct processing stages of this interactive influence. Firstly, in line with the attentional gain hypothesis, a larger amplitude response of the contralateral N1, and Nd1 for attended gratings was observed. Secondly, conforming to the attenuation-by-prediction hypothesis, a smaller negativity in the time window directly following the peak of the N1 component for predicted compared to unpredicted gratings was observed. In line with the hypothesis that attention and prediction interface, unpredicted/unattended stimuli elicited a larger negativity at central-parietal sites, presumably reflecting an increased prediction error signal. Thirdly, larger P3 responses to unpredicted stimuli pointed to the updating of an internal model. Attention and prediction can be considered as differentiated mechanisms that may interact at different processing stages to optimise perception. |
Thomas Meindertsma; Niels A. Kloosterman; Guido Nolte; Andreas K. Engel; Tobias H. Donner Multiple transient signals in human visual cortex associated with an elementary decision Journal Article In: Journal of Neuroscience, vol. 37, no. 23, pp. 5744–5757, 2017. @article{Meindertsma2017, The cerebral cortex continuously undergoes changes in its state, which are manifested in transient modulations of the cortical power spectrum. Cortical state changes also occur at full wakefulness and during rapid cognitive acts, such as perceptual decisions. Previous studies found a global modulation of beta-band (12–30 Hz) activity in human and monkey visual cortex during an elementary visual decision: reporting the appearance or disappearance of salient visual targets surrounded by a distractor. The previous studies disentangled neither the motor action associated with behavioral report nor other secondary processes, such as arousal, from perceptual decision processing per se. Here, we used magnetoencephalography in humans to pinpoint the factors underlying the beta-band modulation.We found that disappearances of a salient target were associated with beta-band suppression, and target reappearances with beta-band enhancement. This was true for both overt behavioral reports (immediate button presses) and silent counting of the perceptual events. This finding indicates that the beta-band modulation was unrelated to the execution of the motor act associated with a behavioral report of the perceptual decision. Further, changes in pupil-linked arousal, fixational eye movements, or gamma-band responses were not necessary for the beta-band modulation. Together, our results suggest that the beta-band modulation was a top-down signal associated with the process of converting graded perceptual signals into a categorical format underlying flexible behavior. This signal may have been fed back from brain regions involved in decision processing to visual cortex, thus enforcing a “decision-consistent” cortical state. |
John M. Egan; Gerard M. Loughnane; Helen Fletcher; Emma Meade; Edmund C. Lalor A gaze independent hybrid-BCI based on visual spatial attention Journal Article In: Journal of Neural Engineering, vol. 14, no. 4, pp. 1–8, 2017. @article{Egan2017, Objective. Brain-computer interfaces (BCI) use measures of brain activity to convey a user's intent without the need for muscle movement. Hybrid designs, which use multiple measures of brain activity, have been shown to increase the accuracy of BCIs, including those based on EEG signals reflecting covert attention. Our study examined whether incorporating a measure of the P3 response improved the performance of a previously reported attention-based BCI design that incorporates measures of steady-state visual evoked potentials (SSVEP) and alpha band modulations. Approach. Subjects viewed stimuli consisting of two bi-laterally located flashing white boxes on a black background. Streams of letters were presented sequentially within the boxes, in random order. Subjects were cued to attend to one of the boxes without moving their eyes, and they were tasked with counting the number of target-letters that appeared within. P3 components evoked by target appearance, SSVEPs evoked by the flashing boxes, and power in the alpha band are modulated by covert attention, and the modulations can be used to classify trials as left-attended or right-attended. Main Results. We showed that classification accuracy was improved by including a P3 feature along with the SSVEP and alpha features (the inclusion of a P3 feature lead to a 9% increase in accuracy compared to the use of SSVEP and Alpha features alone). We also showed that the design improves the robustness of BCI performance to individual subject differences. Significance. These results demonstrate that incorporating multiple neurophysiological indices of covert attention can improve performance in a gaze-independent BCI. |
Joshua J. Foster; Emma M. Bsales; Russell J. Jaffe; Edward Awh Alpha-band activity reveals spontaneous representations of spatial position in visual working memory Journal Article In: Current Biology, vol. 27, no. 20, pp. 3216–3223, 2017. @article{Foster2017, An emerging view suggests that spatial position is an integral component of working memory (WM), such that non-spatial features are bound to locations regardless of whether space is relevant [1, 2]. For instance, past work has shown that stimulus position is spontaneously remembered when non-spatial fea- tures are stored. Item recognition is enhanced when memoranda appear at the same location where they were encoded [3–5], and accessing non-spatial infor- mation elicits shifts of spatial attention to the original position of the stimulus [6, 7]. However, these find- ings do not establish that a persistent, active repre- sentation of stimulus position is maintained in WM because similar effects have also been documented following storage in long-termmemory [8, 9]. Here we show that the spatial position of the memorandum is actively coded by persistent neural activity during a non-spatial WM task. We used a spatial encoding model in conjunction with electroencephalogram (EEG) measurements of oscillatory alpha-band (8– 12 Hz) activity to track active representations of spatial position. The position of the stimulus varied trial to trial but was wholly irrelevant to the tasks. We nevertheless observed active neural representa- tions of the original stimulus position that persisted throughout the retention interval. Further experi- ments established that these spatial representations are dependent on the volitional storage of non- spatial features rather than being a lingering effect of sensory energy or initial encoding demands. These findings provide strong evidence that online spatial representations are spontaneously main- tained in WM—regardless of task relevance—during the storage of non-spatial features. |
Matthew A. Gannon; Stephanie M. Long; Nathan A. Parks Homeostatic plasticity in human extrastriate cortex following a simulated peripheral scotoma Journal Article In: Experimental Brain Research, vol. 235, no. 11, pp. 3391–3401, 2017. @article{Gannon2017, Neuroimaging and patient work over the past decade have indicated that, following retinal deafferenta- tion, the human visual cortex undergoes a large-scale and enduring reorganization of its topography such that the clas- sical retinotopic organization of deafferented visual cortex remaps to represent non-classical regions of visual space. Such long-term visual reorganization is proposed to occur through changes in the functional balance of deafferented visual circuits that engage more lasting changes through activity-dependent neuroplasticity. Here, we investigated the short-term changes in functional balance (short-term plastic- ity; homeostatic plasticity) that occur within deafferented human visual cortices. We recorded electroencephalogram (EEG) while observers were conditioned for 6 s with a simu- lated retinal scotoma (artificial scotoma) positioned 8.0° in the periphery. Visual evoked potentials (VEPs) evoked by the onset of sinusoidal visual probes that varied in their tilt were used to examine changes in cortical excitability within and around cortical representations of the simulated sco- toma. Psychophysical orientation functions obtained from discrimination of visual probe tilt were used to examine alterations in the stimulus selectivity within the scotoma representations. Consistent with a mechanism of homeo- static disinhibition, an early extrastriate component of the VEP (the early phase P1) exhibited increased amplitude fol- lowing the condition with a simulated scotoma relative to a stimulus-matched control condition. This increased visual cortical response was associated with a reduction in the slope of the psychophysical orientation function, suggesting a broader tuning of neural populations within scotoma repre- sentations. Together, these findings support a mechanism of disinhibition in promoting visual plasticity and topographi- cal reorganization. |
Nigel Gebodh; M. Isabel Vanegas; Simon P. Kelly Effects of stimulus size and contrast on the initial primary visual cortical response in humans Journal Article In: Brain Topography, vol. 30, no. 4, pp. 450–460, 2017. @article{Gebodh2017, Decades of intracranial electrophysiological investigation into the primary visual cortex (V1) have produced many fundamental insights into the computations carried out in low-level visual circuits of the brain. Some of the most important work has been simply concerned with the precise measurement of neural response variations as a function of elementary stimulus attributes such as contrast and size. Surprisingly, such simple but fundamental characterization of V1 responses has not been carried out in human electrophysiology. Here we report such a detailed characterization for the initial “C1” component of the scalp-recorded visual evoked potential (VEP). The C1 is known to be dominantly generated by initial afferent activation in V1, but is difficult to record reliably due to interindividual anatomical variability. We used pattern-pulse multifocal VEP mapping to identify a stimulus position that activates the left lower calcarine bank in each individual, and afterwards measured robust negative C1s over posterior midline scalp to gratings presented sequentially at that location. We found clear and systematic increases in C1 peak amplitude and decreases in peak latency with increasing size as well as with increasing contrast. With a sample of 15 subjects and ~180 trials per condition, reliable C1 amplitudes of −0.46 µV were evoked at as low a contrast as 3.13% and as large as −4.82 µV at 100% contrast, using stimuli of 3.33° diameter. A practical implication is that by placing sufficiently-sized stimuli to target favorable calcarine cortical loci, robust V1 responses can be measured at contrasts close to perceptual thresholds, which could greatly facilitate principled studies of early visual perception and attention. |
Edden M. Gerber; Tal Golan; Robert T. Knight; Leon Y. Deouell Cortical representation of persistent visual stimuli Journal Article In: NeuroImage, vol. 161, pp. 67–79, 2017. @article{Gerber2017, Research into visual neural activity has focused almost exclusively on onset- or change-driven responses and little is known about how information is encoded in the brain during sustained periods of visual perception. We used intracranial recordings in humans to determine the degree to which the presence of a visual stimulus is persistently encoded by neural activity. The correspondence between stimulus duration and neural response duration was strongest in early visual cortex and gradually diminished along the visual hierarchy, such that is was weakest in inferior-temporal category-selective regions. A similar posterior-anterior gradient was found within inferior temporal face-selective regions, with posterior but not anterior sites showing persistent face-selective activity. The results suggest that regions that appear uniform in terms of their category selectivity are dissociated by how they temporally represent a stimulus in support of ongoing visual perception, and delineate a large-scale organizing principle of the ventral visual stream. |
Siobhán Harty; Peter R. Murphy; Ian H. Robertson; Redmond G. O'Connell Parsing the neural signatures of reduced error detection in older age Journal Article In: NeuroImage, vol. 161, pp. 43–55, 2017. @article{Harty2017, Recent work has demonstrated that explicit error detection relies on a neural evidence accumulation process that can be traced in the human electroencephalogram (EEG). Here, we sought to establish the impact of natural aging on this process by recording EEG from young (18–35 years) and older adults (65–88 years) during the performance of a Go/No-Go paradigm in which participants were required to overtly signal their errors. Despite performing the task with equivalent accuracy, older adults reported substantially fewer errors, and the timing of their reports were both slower and more variable. These behavioral differences were linked to three key neurophysiological changes reflecting distinct parameters of the error detection decision process: a reduction in medial frontal delta/theta (2–7 Hz) activity, indicating diminished top-down input to the decision process; a slower rate of evidence accumulation as indexed by the rate of rise of a centro-parietal signal, known as the error positivity; and a higher motor execution threshold as indexed by lateralized beta-band (16–30 Hz) activity. Our data provide novel insight into how the natural aging process affects the neural underpinnings of error detection. |
Philipp N. Hesse; Constanze Schmitt; Steffen Klingenhoefer; Frank Bremmer Preattentive processing of numerical visual information Journal Article In: Frontiers in Human Neuroscience, vol. 11, pp. 70, 2017. @article{Hesse2017, Humans can perceive and estimate approximate numerical information, even when accurate counting is impossible e.g. due to short presentation time. If the number of objects to be estimated is small, typically around one to four items, observers are able to give very fast and precise judgments with high confidence – an effect that is called subitizing. Due to its speed and effortless nature subitizing has usually been assumed to be preattentive, putting it into the same category as other low level visual features like color or orientation. More recently, however, a number of studies have suggested that subitizing might be dependent on attentional resources. In our current study we investigated the potentially preattentive nature of visual numerical perception in the subitizing range by means of EEG. We presented peripheral, task irrelevant sequences of stimuli consisting of a certain number of circular patches while participants were engaged in a demanding, non-numerical detection task at the fixation point drawing attention away from the number stimuli. Within a sequence of stimuli of a given number of patches (called ‘standards') we interspersed some stimuli of different numerosity (‘oddballs'). We compared the evoked responses to visually identical stimuli that had been presented in two different conditions, serving as standard in one condition and as oddball in the other. We found significant visual mismatch negativity (vMMN) responses over parieto-occipital electrodes. In addition to the ERP analysis, we performed a time-frequency analysis to investigate whether the vMMN was accompanied by additional oscillatory processes. We found a concurrent increase in evoked theta power of similar strength over both hemispheres. Our results provide clear evidence for a preattentive processing of numerical visual information in the subitizing range. |
2016 |
Lauren R. Godier; Jessica C. Scaife; Sven Braeutigam; Rebecca J. Park Enhanced early neuronal processing of food pictures in Anorexia Nervosa: A magnetoencephalography study Journal Article In: Psychiatry Journal, vol. 2016, pp. 1–13, 2016. @article{Godier2016, Neuroimaging studies in Anorexia Nervosa (AN) have shown increased activation in reward and cognitive control regions in response to food, and a behavioral attentional bias (AB) towards food stimuli is reported. This study aimed to further investigate the neural processing of food using magnetoencephalography (MEG). Participants were 13 females with restricting-type AN, 14 females recovered from restricting-type AN, and 15 female healthy controls. MEG data was acquired whilst participants viewed high- and low-calorie food pictures. Attention was assessed with a reaction time task and eye tracking. Time-series analysis suggested increased neural activity in response to both calorie conditions in the AN groups, consistent with an early AB. Increased activity was observed at 150 ms in the current AN group. Neuronal activity at this latency was at normal level in the recovered group; however, this group exhibited enhanced activity at 320 ms after stimulus. Consistent with previous studies, analysis in source space and behavioral data suggested enhanced attention and cognitive control processes in response to food stimuli in AN. This may enable avoidance of salient food stimuli and maintenance of dietary restraint in AN. A later latency of increased activity in the recovered group may reflect a reversal of this avoidance, with source space and behavioral data indicating increased visual and cognitive processing of food stimuli. |
Tal Golan; Ido Davidesco; Meir Meshulam; David M. Groppe; Pierre Mégevand; Erin M. Yeagle; Matthew S. Goldfinger; Michal Harel; Lucia Melloni; Charles E. Schroeder; D. L. Deouell; Ashesh D. Mehta; Rafael Malach Human intracranial recordings link suppressed transients rather than 'filling-in' to perceptual continuity across blinks Journal Article In: eLife, vol. 5, pp. 1–28, 2016. @article{Golan2016, We hardly notice our eye blinks, yet an externally generated retinal interruption of a similar duration is perceptually salient. We examined the neural correlates of this perceptual distinction using intracranially measured ECoG signals from human visual cortex in 14 patients. In early visual areas (V1 and V2), the disappearance of the stimulus due to either invisible blinks or salient blank video frames ('gaps') led to a similar drop in activity level, followed by a positive overshoot beyond baseline, triggered by stimulus reappearance. Ascending the visual hierarchy, the reappearance-related overshoot gradually subsided for blinks but not for gaps. By contrast, the disappearance-related drop did not follow the perceptual distinction - it was actually slightly more pronounced for blinks than for gaps. These findings suggest that blinks' limited visibility compared with gaps is correlated with suppression of blink-related visual activity transients, rather than with 'filling-in' of the occluded content during blinks. |
Gil Gonen-Yaacovi; Ayelet Arazi; Nitzan Shahar; Anat Karmon; Shlomi Haar; Nachshon Meiran; Ilan Dinstein Increased ongoing neural variability in ADHD Journal Article In: Cortex, vol. 81, pp. 50–63, 2016. @article{GonenYaacovi2016, Attention Deficit Hyperactivity Disorder (ADHD) has been described as a disorder where frequent lapses of attention impair the ability of an individual to focus/attend in a sustained manner, thereby generating abnormally large intra-individual behavioral variability across trials. Indeed, increased reaction time (RT) variability is a fundamental behavioral characteristic of individuals with ADHD found across a large number of cognitive tasks. But what is the underlying neurophysiology that might generate such behavioral instability? Here, we examined trial-by-trial EEG response variability to visual and auditory stimuli while subjects' attention was diverted to an unrelated task at the fixation cross. Comparisons between adult ADHD and control participants revealed that neural response variability was significantly larger in the ADHD group as compared with the control group in both sensory modalities. Importantly, larger trial-by-trial variability in ADHD was apparent before and after stimulus presentation as well as in trials where the stimulus was omitted, suggesting that ongoing (rather than stimulus-evoked) neural activity is continuously more variable (noisier) in ADHD. While the patho-physiological mechanisms causing this increased neural variability remain unknown, they appear to act continuously rather than being tied to a specific sensory or cognitive process. |
Sarah Gregory; Marco Fusca; Geraint Rees; D. Samuel Schwarzkopf; Gareth Barnes Gamma frequency and the spatial tuning of primary visual cortex Journal Article In: PLoS ONE, vol. 11, no. 6, pp. e0157374, 2016. @article{Gregory2016a, Visual stimulation produces oscillatory gamma responses in human primary visual cortex (V1) that also relate to visual perception. We have shown previously that peak gamma frequency positively correlates with central V1 cortical surface area. We hypothesized that people with larger V1 would have smaller receptive fields and that receptive field size, not V1 are, might explain this relationship. Here we set out to test this hypothesis directly by investigating the relationship between fMRI estimated population receptive field (pRF) size and gamma frequency in V1. We stimulated both the near-centre and periphery of the visual field using both large and small stimuli in each location and replicated our previous finding of a positive correlation between V1 surface area and peak gamma frequency. Counter to our expectation, we found that between participants V1 size (and not pRF size) accounted for most of the variability in gamma frequency. Within-participants we found that gamma frequency increased, rather than decreased, with stimulus eccentricity directly contradicting our initial hypothesis. |
Tjerk P. Gutteling; W. Pieter Medendorp Role of alpha-band oscillations in spatial updating across whole body motion Journal Article In: Frontiers in Psychology, vol. 7, pp. 671, 2016. @article{Gutteling2016, When moving around in the world, we have to keep track of important locations in our surroundings. In this process, called spatial updating, we must estimate our body motion and correct representations of memorized spatial locations in accordance with this motion. While the behavioral characteristics of spatial updating across whole body motion have been studied in detail, its neural implementation lacks detailed study. Here we use electro-encephalography (EEG) to distinguish various spectral components of this process. Subjects gazed at a central body-fixed point in otherwise complete darkness, while a target was briefly flashed, either left or right from this point. Subjects had to remember the location of this target as either moving along with the body or remaining fixed in the world while being translated sideways on a passive motion platform. After the motion, subjects had to indicate the remembered target location in the instructed reference frame using a mouse response. While the body motion, as detected by the vestibular system, should not affect the representation of body-fixed targets, it should interact with the representation of a world-centered target to update its location relative to the body. We show that the initial presentation of the visual target induced a reduction of alpha band power in contralateral parieto-occipital areas, which evolved to a sustained increase during the subsequent memory period. Motion of the body led to a reduction of alpha band power in central parietal areas extending to lateral parieto-temporal areas, irrespective of whether the targets had to be memorized relative to world or body. When updating a world-fixed target, its internal representation shifts hemispheres, only when subjects' behavioral responses suggested an update across the body midline. Our results suggest that parietal cortex is involved in both self-motion estimation and the selective application of this motion information to maintaining target locations as fixed in the world or fixed to the body. |
Karin Heidlmayr; Karine Dore-Mazars; Xavier Aparico; Frederic Isel In: PLoS ONE, vol. 11, no. 11, pp. e0165029, 2016. @article{Heidlmayr2016, In the present electroencephalographical study, we asked to which extent executive control processes are shared by both the language and motor domain. The rationale was to examine whether executive control processes whose efficiency is reinforced by the frequent use of a second language can lead to a benefit in the control of eye movements, i.e. a non-linguistic activity. For this purpose, we administrated to 19 highly proficient late French-German bilingual participants and to a control group of 20 French monolingual participants an antisaccade task, i.e. a specific motor task involving control. In this task, an automatic saccade has to be suppressed while a voluntary eye movement in the opposite direction has to be carried out. Here, our main hypothesis is that an advantage in the antisaccade task should be observed in the bilinguals if some properties of the control processes are shared between linguistic and motor domains. ERP data revealed clear differences between bilinguals and monolinguals. Critically, we showed an increased N2 effect size in bilinguals, thought to reflect better efficiency to monitor conflict, combined with reduced effect sizes on markers reflecting inhibitory control, i.e. cue-locked positivity, the target-locked P3 and the saccade-locked presaccadic positivity (PSP). Moreover, effective connectivity analyses (dynamic causal modelling; DCM) on the neuronal source level indicated that bilinguals rely more strongly on ACC-driven control while monolinguals rely on PFC-driven control. Taken together, our combined ERP and effective connectivity findings may reflect a dynamic interplay between strengthened conflict monitoring, associated with subsequently more efficient inhibition in bilinguals. Finally, L2 proficiency and immersion experience constitute relevant factors of the language background that predict efficiency of inhibition. To conclude, the present study provided ERP and effective connectivity evidence for domain-general executive control involvement in handling multiple language use, leading to a control advantage in bilingualism. |
Christoph Huber-Huber; Thomas Ditye; María Marchante Fernández; Ulrich Ansorge Using temporally aligned event-related potentials for the investigation of attention shifts prior to and during saccades Journal Article In: Neuropsychologia, vol. 92, pp. 129–141, 2016. @article{HuberHuber2016, According to the pre-motor theory of attention, attention is shifted to a saccade's landing position before the saccade is executed. Such pre-saccadic attention shifts are usually studied in psychophysical dual-task conditions, with a target-discrimination task before saccade onset. Here, we present a novel approach to investigate pre-saccadic attention shifts with the help of event-related potentials (ERPs). Participants executed one or two saccades to color-defined targets while ERPs and eye-movements were recorded. In single-target blocks participants executed a single saccade. In two-targets blocks participants made either a single saccade to one of the targets, or two successive saccades to both targets. Importantly, in two-targets blocks, targets could appear on the same or on opposite sides of the vertical midline. This allowed us to study contra-to-ipsilateral ERP differences (such as the N2pc or PCN) that reflect attention shifts to the targets, prior to saccade onset and during saccades. If pre-saccadic attention shifts to saccade target locations are necessary for saccade execution and if searched-for saccade targets capture attention, there should be enhanced attentional competition (1) between two targets compared to single targets; (2) between two opposite-sides targets compared to two same-side targets; and (3) in two saccades rather than one saccade conditions: More attentional competition was expected to delay saccade latency and to weaken pre-saccadic laterality effects in ERPs. Hypotheses were tested by means of temporally aligned ERPs that were simultaneously time-locked to stimulus onsets, saccade onsets, and saccade offsets. Predictions (1) and (2) were partly and fully confirmed, respectively, but no evidence was found for (3). We explain the implications of our results for the role of attention during saccade preparation, and we point out how temporally aligned ERPs compare to ICA-based electroencephalogram (EEG) artifact correction procedures and to psychophysical dual-task approaches. |
Andrey R. Nikolaev; Radha Nila Meghanathan; Cees Leeuwen Combining EEG and eye movement recording in free viewing: Pitfalls and possibilities Journal Article In: Brain and Cognition, vol. 107, pp. 55–83, 2016. @article{Nikolaev2016, Co-registration of EEG and eye movement has promise for investigating perceptual processes in free viewing conditions, provided certain methodological challenges can be addressed. Most of these arise from the self-paced character of eye movements in free viewing conditions. Successive eye movements occur within short time intervals. Their evoked activity is likely to distort the EEG signal during fixation. Due to the non-uniform distribution of fixation durations, these distortions are systematic, survive across-trials averaging, and can become a source of confounding. We illustrate this problem with effects of sequential eye movements on the evoked potentials and time-frequency components of EEG and propose a solution based on matching of eye movement characteristics between experimental conditions. The proposal leads to a discussion of which eye movement characteristics are to be matched, depending on the EEG activity of interest. We also compare segmentation of EEG into saccade-related epochs relative to saccade and fixation onsets and discuss the problem of baseline selection and its solution. Further recommendations are given for implementing EEG-eye movement co-registration in free viewing conditions. By resolving some of the methodological problems involved, we aim to facilitate the transition from the traditional stimulus-response paradigm to the study of visual perception in more naturalistic conditions. |
Sven Ohl; Reinhold Kliegl Revealing the time course of signals influencing the generation of secondary saccades using Aalen's additive hazards model Journal Article In: Vision Research, vol. 124, pp. 52–58, 2016. @article{Ohl2016, Saccadic eye movements are frequently followed by smaller secondary saccades which are generally assumed to correct for the error in primary saccade landing position. However, secondary saccades can also occur after accurate primary saccades and they are often as small as microsaccades, therefore raising the need to further scrutinize the processes involved in secondary saccade generation. Following up a previous study, we analyzed secondary saccades using rate analysis which allows us to quantify experimental effects as shifts in distributions, therefore going beyond comparisons of mean differences. We use Aalen's additive hazards model to delineate the time course of key influences on the secondary saccade rate. In addition to the established effect of primary saccade error, we observed a time-varying influence of under- vs. overshooting - with a higher risk of generating secondary saccades following undershoots. Moreover, increasing target eccentricity influenced the programming of secondary saccades, therefore demonstrating that error-unrelated variables co-determine secondary saccade programs. Our results provide new insights into the generative mechanisms of small saccades during postsaccadic fixation that need to be accounted for by secondary saccade models. |
Sven Ohl; Christian Wohltat; Reinhold Kliegl; Olga Pollatos; Ralf Engbert Microsaccades are coupled to heartbeat Journal Article In: Journal of Neuroscience, vol. 36, no. 4, pp. 1237–1241, 2016. @article{Ohl2016a, During visual fixation, the eye generates microsaccades and slower components of fixational eye movements that are part of the visual processing strategy in humans. Here, we show that ongoing heartbeat is coupled to temporal rate variations in the generation of microsaccades. Using coregistration of eye recording and ECG in humans, we tested the hypothesis that microsaccade onsets are coupled to the relative phase of the R-R intervals in heartbeats. We observed significantly more microsaccades during the early phase after the R peak in the ECG. This form of coupling between heartbeat and eye movements was substantiated by the additional finding of a coupling between heart phase and motion activity in slow fixational eye movements; i.e., retinal image slip caused by physiological drift. Our findings therefore demonstrate a coupling of the oculomotor system and ongoing heartbeat, which provides further evidence for bodily influences on visuomotor functioning. |
Rick A. Adams; Markus Bauer; Dimitris Pinotsis; Karl J. Friston Dynamic causal modelling of eye movements during pursuit: Confirming precision-encoding in V1 using MEG Journal Article In: Neuroimage, vol. 132, pp. 175–189, 2016. @article{Adams2016, This paper shows that it is possible to estimate the subjective precision (inverse variance) of Bayesian beliefs during oculomotor pursuit. Subjects viewed a sinusoidal target, with or without random fluctuations in its motion. Eye trajectories and magnetoencephalographic (MEG) data were recorded concurrently. The target was periodically occluded, such that its reappearance caused a visual evoked response field (ERF). Dynamic causal modelling (DCM) was used to fit models of eye trajectories and the ERFs. The DCM for pursuit was based on predictive coding and active inference, and predicts subjects' eye movements based on their (subjective) Bayesian beliefs about target (and eye) motion. The precisions of these hierarchical beliefs can be inferred from behavioural (pursuit) data. The DCM for MEG data used an established biophysical model of neuronal activity that includes parameters for the gain of superficial pyramidal cells, which is thought to encode precision at the neuronal level. Previous studies (using DCM of pursuit data) suggest that noisy target motion increases subjective precision at the sensory level: i.e., subjects attend more to the target's sensory attributes. We compared (noisy motion-induced) changes in the synaptic gain based on the modelling of MEG data to changes in subjective precision estimated using the pursuit data. We demonstrate that imprecise target motion increases the gain of superficial pyramidal cells in V1 (across subjects). Furthermore, increases in sensory precision – inferred by our behavioural DCM – correlate with the increase in gain in V1, across subjects. This is a step towards a fully integrated model of brain computations, cortical responses and behaviour that may provide a useful clinical tool in conditions like schizophrenia. |
Micah Allen; Darya Frank; D. Samuel Schwarzkopf; Francesca Fardo; Joel S. Winston; Tobias U. Hauser; Geraint Rees Unexpected arousal modulates the influence of sensory noise on confidence Journal Article In: eLife, vol. 5, pp. 1–17, 2016. @article{Allen2016, Human perception is invariably accompanied by a graded feeling of confidence that guides metacognitive awareness and decision-making. It is often assumed that this arises solely from the feed-forward encoding of the strength or precision of sensory inputs. In contrast, interoceptive inference models suggest that confidence reflects a weighted integration of sensory precision and expectations about internal states, such as arousal. Here we test this hypothesis using a novel psychophysical paradigm, in which unseen disgust-cues induced unexpected, unconscious arousal just before participants discriminated motion signals of variable precision. Across measures of perceptual bias, uncertainty, and physiological arousal we found that arousing disgust cues modulated the encoding of sensory noise. Furthermore, the degree to which trial-by-trial pupil fluctuations encoded this nonlinear interaction correlated with trial level confidence. Our results suggest that unexpected arousal regulates perceptual precision, such that subjective confidence reflects the integration of both external sensory and internal, embodied states. |
Mariana Babo-Rebelo; Craig G. Richter; Catherine Tallon-Baudry Neural responses to heartbeats in the default network encode the self in spontaneous thoughts Journal Article In: Journal of Neuroscience, vol. 36, no. 30, pp. 7829–7840, 2016. @article{BaboRebelo2016, The default network (DN) has been consistently associated with self-related cognition, but also to bodily state monitoring and autonomic regulation. We hypothesized that these two seemingly disparate functional roles of the DN are functionally coupled, in line with theories proposing that selfhood is grounded in the neural monitoring of internal organs, such as the heart. We measured with magnetoencephalograhy neural responses evoked by heartbeats while human participants freely mind-wandered. When interrupted by a visual stimulus at random intervals, participants scored the self-relatedness of the interrupted thought. They evaluated their involvement as the first-person perspective subject or agent in the thought ("I"), and on another scale to what degree they were thinking about themselves ("Me"). During the interrupted thought, neural responses to heartbeats in two regions of the DN, the ventral precuneus and the ventromedial prefrontal cortex, covaried, respectively, with the "I" and the "Me" dimensions of the self, even at the single-trial level. No covariation between self-relatedness and peripheral autonomic measures (heart rate, heart rate variability, pupil diameter, electrodermal activity, respiration rate, and phase) or alpha power was observed. Our results reveal a direct link between selfhood and neural responses to heartbeats in the DN and thus directly support theories grounding selfhood in the neural monitoring of visceral inputs. More generally, the tight functional coupling between self-related processing and cardiac monitoring observed here implies that, even in the absence of measured changes in peripheral bodily measures, physiological and cognitive functions have to be considered jointly in the DN. |
Indu P. Bodala; Junhua Li; Nitish V. Thakor; Hasan Al-Nashash EEG and eye tracking demonstrate vigilance enhancement with challenge integration Journal Article In: Frontiers in Human Neuroscience, vol. 10, pp. 273, 2016. @article{Bodala2016, Maintaining vigilance is possibly the first requirement for surveillance tasks where personnel are faced with monotonous yet intensive monitoring tasks. Decrement in vigilance in such situations could result in dangerous consequences such as accidents, loss of life and system failure. In this paper, we investigate the possibility to enhance vigilance or sustained attention using ‘challenge integration', a strategy that integrates a primary task with challenging stimuli. A primary surveillance task (identifying an intruder in a simulated factory environment) and a challenge stimulus (periods of rain obscuring the surveillance scene) were employed to test the changes in vigilance levels. The effect of integrating challenging events (resulting from artificially simulated rain) into the task were compared to the initial monotonous phase. EEG and eye tracking data is collected and analyzed for n = 12 subjects. Frontal midline theta power and frontal theta to parietal alpha power ratio which are used as measures of engagement and attention allocation show an increase due to challenge integration (p < 0.05 in each case). Relative delta band power of EEG also shows statistically significant suppression on the frontoparietal and occipital cortices due to challenge integration (p < 0.05). Saccade amplitude, saccade velocity and blink rate obtained from eye tracking data exhibit statistically significant changes during the challenge phase of the experiment (p < 0.05 in each case). From the correlation analysis between the statistically significant measures of eye tracking and EEG, we infer that saccade amplitude and saccade velocity decrease with vigilance decrement along with frontal midline theta and frontal theta to parietal alpha ratio. Conversely, blink rate and relative delta power increase with vigilance decrement. However, these measures exhibit a reverse trend when challenge stimulus appears in the task suggesting vigilance enhancement. Moreover, the mean reaction time is lower for the challenge integrated phase (RT mean = 3.65 ± 1.4 secs) compared to initial monotonous phase without challenge (RT mean = 4.6 ± 2.7 secs). Our work shows that vigilance level, as assessed by response of these vital signs, is enhanced by challenge integration. |
Florence Campana; Ignacio Rebollo; Anne E. Urai; Valentin Wyart; Catherine Tallon-Baudry Conscious vision proceeds from global to local content in goal-directed tasks and spontaneous vision Journal Article In: Journal of Neuroscience, vol. 36, no. 19, pp. 5200–5213, 2016. @article{Campana2016, The reverse hierarchy theory (Hochstein and Ahissar, 2002) makes strong, but so far untested, predictions on conscious vision. In this theory, local details encoded in lower-order visual areas are unconsciously processed before being automatically and rapidly combined into global information in higher-order visual areas, where conscious percepts emerge. Contingent on current goals, local details can afterward be consciously retrieved. This model therefore predicts that (1) global information is perceived faster than local details, (2) global information is computed regardless of task demands during early visual processing, and (3) spontaneous vision is dominated by global percepts. We designed novel textured stimuli that are, as opposed to the classic Navon's letters, truly hierarchical (i.e., where global information is solely defined by local information but where local and global orientations can still be manipulated separately). In line with the predictions, observers were systematically faster reporting global than local properties of those stimuli. Second, global information could be decoded from magneto-encephalographic data during early visual processing regardless of task demands. Last, spontaneous subjective reports were dominated by global information and the frequency and speed of spontaneous global perception correlated with the accuracy and speed in the global task. No such correlation was observed for local information. We therefore show that information at different levels of the visual hierarchy is not equally likely to become conscious; rather, conscious percepts emerge preferentially at a global level. We further show that spontaneous reports can be reliable and are tightly linked to objective performance at the global level. |
Jing Chen; Matteo Valsecchi; Karl R. Gegenfurtner LRP predicts smooth pursuit eye movement onset during the ocular tracking of self-generated movements Journal Article In: Journal of Neurophysiology, vol. 116, no. 1, pp. 18–29, 2016. @article{Chen2016c, Several studies indicated that human observers are very efficient at tracking self-generated hand movements with their gaze, yet it is not clear whether this is simply a byproduct of the predictability of self-generated actions or if it results from a deeper coupling of the somatomotor and oculomotor systems. In a first behavioral experiment we compared pursuit performance as observers either followed their own finger or tracked a dot whose motion was externally generated but mimicked their finger motion. We found that even when the dot motion was completely predictable both in terms of onset time and in terms of kinematics, pursuit was not identical to the one produced as the observers tracked their finger, as evidenced by increased rate of catch-up saccades and by the fact that in the initial phase of the movement gaze was lagging behind the dot, whereas it was ahead of the finger. In a second experiment we recorded EEG in the attempt to find a direct link between the finger motor preparation, indexed by the lateralized readiness potential (LRP), and the latency of smooth pursuit. After taking into account finger movement onset variability, we observed larger LRP amplitudes associated with earlier smooth pursuit onset across trials. The same held across subjects, where average LRP onset correlated with average eye latency. The evidence from both experiments concurs to indicate that a strong coupling exists between the motor systems leading to eye and finger movements and that simple top-down predictive signals are unlikely to support optimal coordination. |
Hak Soo Choi; Shinjung Kim; Donghoon Lee; Chang Seok Kim; Myung Yung Jeong Synchronized tracking of brain cognitive processing using EEG and vision signals Journal Article In: Applied Spectroscopy Reviews, vol. 51, no. 7-9, pp. 592–602, 2016. @article{Choi2016, Many efforts have been made to understand the neural mechanisms of the human brain. However, visualization of human brain processing has been a main challenge in the field. It is still largely unknown how the human brain allocates attention to target objects while excluding unrelated information in a complex visual environment. Using simultaneous electroencephalogram and eye tracking measurements, in this study, we analyzed two brain regions separately to detect the brain wave activity during visual information processing. We observed an activation difference between sensory (P100) and cognitive (P300) processing, and the behavioral response was improved by providing valid cue-target location information. Furthermore, neural processing was evaluated according to the specific area of brain activation and eye movements during cognitive processing. Our results demonstrate the correlation between behavior performance and visual stimuli and suggest an advantage of combined paradigms for efficient visual information processing. |
Marieke E. Nieuwenhuijzen; Eva W. P. Borne; Ole Jensen; Marcel A. J. Gerven Spatiotemporal dynamics of cortical representations during and after stimulus presentation Journal Article In: Frontiers in Systems Neuroscience, vol. 10, pp. 42, 2016. @article{Nieuwenhuijzen2016, Visual perception is a spatiotemporally complex process. In this study, we investigated cortical dynamics during and after stimulus presentation. We observed that visual category information related to the difference between faces and objects became apparent in the occipital lobe after 63 ms. Within the next 110 ms, activation spread out to include the temporal lobe before returning to residing mainly in the occipital lobe again. After stimulus offset, a peak in information was observed, comparable to the peak after stimulus onset. Moreover, similar processes, albeit not identical, seemed to underlie both peaks. Information about the categorical identity of the stimulus remained present until 677 ms after stimulus offset, during which period the stimulus had to be retained in working memory. Activation patterns initially resembled those observed during stimulus presentation. After about 200 ms, however, this representation changed and class-specific activity became more equally distributed over the four lobes. These results show that, although there are common processes underlying stimulus representation both during and after stimulus presentation, these representations change depending on the specific stage of perception and maintenance. |
Rosanne M. Diepen; Lee M. Miller; Ali Mazaheri; Joy J. Geng The role of alpha activity in spatial and feature-based attention. Journal Article In: eNeuro, vol. 3, no. 5, pp. 1–11, 2016. @article{Diepen2016, Modulations in alpha oscillations (?10 Hz) are typically studied in the context of anticipating upcoming stimuli. Alpha power decreases in sensory regions processing upcoming targets compared to regions processing distracting input, thereby likely facilitating processing of relevant information while suppressing irrelevant. In this electroencephalography study using healthy human volunteers, we examined whether modulations in alpha power also occur after the onset of a bilaterally presented target and distractor. Spatial attentionwasmanipulated through spatial cues and feature-based attention through adjusting the color-similarity of distractors to the target. Consistent with previous studies, we found that informative spatial cues induced a relative decrease of pretarget alpha power at occipital electrodes contralateral to the expected target location. Interestingly, this pattern reemerged relatively late (300–750 ms) after stimulus onset, suggesting that lateralized alpha reflects not only preparatory attention, but also ongoing attentive stimulus processing. Uninformative cues (i.e., conveying no information about the spatial location of the target) resulted in an interaction between spatial attention and feature-based attention in post-target alpha lateralization. When the target was paired with a low-similarity distractor, post-target alpha was lateralized (500–900 ms). Crucially, the lateralization was absent when target selection was ambig- uous because the distractor was highly similar to the target. Instead, during this condition, midfrontal theta was increased, indicative of reactive conflict resolution. Behaviorally, the degree of alpha lateralization was negatively correlated with the reaction time distraction cost induced by target–distractor similarity. These results suggest a pivotal role for poststimulus alpha lateralization in protecting sensory processing of target information. |
Lorenzo Vignali; Nicole A. Himmelstoss; Stefan Hawelka; Fabio Richlan; Florian Hutzler Oscillatory brain dynamics during sentence reading: A fixation-related spectral perturbation analysis Journal Article In: Frontiers in Human Neuroscience, vol. 10, pp. 191, 2016. @article{Vignali2016, The present study investigated oscillatory brain dynamics during self-paced sentence-level processing. Participants read fully correct sentences, sentences containing a semantic violation and "sentences" in which the order of the words was randomized. At the target word level, fixations on semantically unrelated words elicited a lower-beta band (13-18 Hz) desynchronization. At the sentence level, gamma power (31-55 Hz) increased linearly for syntactically correct sentences, but not when the order of the words was randomized. In the 300-900 ms time window after sentence onsets, theta power (4-7 Hz) was greater for syntactically correct sentences as compared to sentences where no syntactic structure was preserved (random words condition). We interpret our results as conforming with a recently formulated predictive-coding framework for oscillatory neural dynamics during sentence-level language comprehension. Additionally, we discuss how our results relate to previous findings with serial visual presentation vs. self-paced reading. |
Hyojin Park; Christoph Kayser; Gregor Thut; Joachim Gross Lip movements entrain the observers' low-frequency brain oscillations to facilitate speech intelligibility Journal Article In: eLife, vol. 5, pp. 1–17, 2016. @article{Park2016, During continuous speech, lip movements provide visual temporal signals that facilitate speech processing. Here, using MEG we directly investigated how these visual signals interact with rhythmic brain activity in participants listening to and seeing the speaker. First, we investigated coherence between oscillatory brain activity and speaker's lip movements and demonstrated significant entrainment in visual cortex. We then used partial coherence to remove contributions of the coherent auditory speech signal from the lip-brain coherence. Comparing this synchronization between different attention conditions revealed that attending visual speech enhances the coherence between activity in visual cortex and the speaker's lips. Further, we identified a significant partial coherence between left motor cortex and lip movements and this partial coherence directly predicted comprehension accuracy. Our results emphasize the importance of visually entrained and attention-modulated rhythmic brain activity for the enhancement of audiovisual speech processing. |
Maria Solé Puig; Josep Marco Pallarés; Laura Perez Zapata; Laura Puigcerver; Josep Cañete; Hans Supèr Attentional selection accompanied by eye vergence as revealed by event-related brain potentials Journal Article In: PLoS ONE, vol. 11, no. 12, pp. e0167646, 2016. @article{Puig2016, Neural mechanisms of attention allow selective sensory information processing. Top-down deployment of visual-spatial attention is conveyed by cortical feedback connections from frontal regions to lower sensory areas modulating late stimulus responses. A recent study reported the occurrence of small eye vergence during orienting top-down attention. Here we assessed a possible link between vergence and attention by comparing visual event related potentials (vERPs) to a cue stimulus that induced attention to shift towards the target location to the vERPs to a no-cue stimulus that did not trigger orienting attention. The results replicate the findings of eye vergence responses during orienting attention and show that the strength and time of eye vergence coincide with the onset and strength of the vERPs when subjects oriented attention. Our findings therefore support the idea that eye vergence relates to and possibly has a role in attentional selection. |
Pavan Ramkumar; Bruce C. Hansen; Sebastian Pannasch; Lester C. Loschky Visual information representation and rapid-scene categorization are simultaneous across cortex: An MEG study Journal Article In: NeuroImage, vol. 134, pp. 295–304, 2016. @article{Ramkumar2016, Perceiving the visual world around us requires the brain to represent the features of stimuli and to categorize the stimulus based on these features. Incorrect categorization can result either from errors in visual representation or from errors in processes that lead to categorical choice. To understand the temporal relationship between the neural signatures of such systematic errors, we recorded whole-scalp magnetoencephalography (MEG) data from human subjects performing a rapid-scene categorization task. We built scene category decoders based on (1) spatiotemporally resolved neural activity, (2) spatial envelope (SpEn) image features, and (3) behavioral responses. Using confusion matrices, we tracked how well the pattern of errors from neural decoders could be explained by SpEn decoders and behavioral errors, over time and across cortical areas. Across the visual cortex and the medial temporal lobe, we found that both SpEn and behavioral errors explained unique variance in the errors of neural decoders. Critically, these effects were nearly simultaneous, and most prominent between 100 and 250 ms after stimulus onset. Thus, during rapid-scene categorization, neural processes that ultimately result in behavioral categorization are simultaneous and co-localized with neural processes underlying visual information representation. |
Sameer Saproo; Victor Shih; David C. Jangraw; Paul Sajda Neural mechanisms underlying catastrophic failure in human-machine interaction during aerial navigation Journal Article In: Journal of Neural Engineering, vol. 13, pp. 1–12, 2016. @article{Saproo2016, Objective. We investigated the neural correlates of workload buildup in a fine visuomotor task called the boundary avoidance task (BAT). The BAT has been known to induce naturally occurring failures of human–machine coupling in high performance aircraft that can potentially lead to a crash—these failures are termed pilot induced oscillations (PIOs). Approach. We recorded EEG and pupillometry data from human subjects engaged in a flight BAT simulated within a virtual 3D environment. Main results. We find that workload buildup in a BAT can be successfully decoded from oscillatory features in the electroencephalogram (EEG). Information in delta, theta, alpha, beta, and gamma spectral bands of the EEG all contribute to successful decoding, however gamma band activity with a lateralized somatosensory topography has the highest contribution, while theta band activity with a fronto-central topography has the most robust contribution in terms of real-world usability. We show that the output of the spectral decoder can be used to predict PIO susceptibility. We also find that workload buildup in the task induces pupil dilation, the magnitude of which is significantly correlated with the magnitude of the decoded EEG signals. These results suggest that PIOs may result from the dysregulation of cortical networks such as the locus coeruleus (LC)—anterior cingulate cortex (ACC) circuit. Significance. Our findings may generalize to similar control failures in other cases of tight manmachine coupling where gains and latencies in the control system must be inferred and compensated for by the human operators. A closed-loop intervention using neurophysiological decoding of workload buildup that targets the LC-ACC circuit may positively impact operator performance in such situations. |
Yoshihito Shigihara; Hideyuki Hoshi; Semir Zeki Early visual cortical responses produced by checkerboard pattern stimulation Journal Article In: NeuroImage, vol. 134, pp. 532–539, 2016. @article{Shigihara2016, Visual evoked potentials have been traditionally triggered with flash or reversing checkerboard stimuli and recorded with electroencephalographic techniques, largely but not exclusively in clinical or clinically related settings. They have been crucial in determining the healthy functioning or otherwise of the visual pathways up to and including the cerebral cortex. They have typically given early response latencies of 100 ms, the source of which has been attributed to V1, with the prestriate cortex being secondarily activated somewhat later. On the other hand, magnetoencephalographic studies using stimuli better tailored to the physiology of individual, specialized, visual areas have given early latencies of <. 50 ms with the sources localized in both striate (V1) and prestriate cortex. In this study, we used the reversing checkerboard pattern as a stimulus and recorded cortical visual evoked magnetic fields with magnetoencephalography, to establish whether very early responses can be traced to (estimated) in both striate and prestriate cortex, since such a demonstration would enhance considerably the power of this classical approach in clinical investigations. Our results show that cortical responses evoked by checkerboard patterns can be detected before 50 ms post-stimulus onset and that their sources can be estimated in both striate and prestriate cortex, suggesting a strong parallel input from the sub-cortex to both striate and prestriate divisions of the visual cortex. |
Sergei L. Shishkin; Yuri O. Nuzhdin; Evgeny P. Svirin; Alexander G. Trofimov; Anastasia A. Fedorova; Bogdan L. Kozyrskiy; Boris M. Velichkovsky EEG negativity in fixations used for gaze-based control: Toward converting intentions into actions with an eye-brain-computer interface Journal Article In: Frontiers in Neuroscience, vol. 10, pp. 528, 2016. @article{Shishkin2016, We usually look at an object when we are going to manipulate it. Thus, eye tracking can be used to communicate intended actions. An effective human-machine interface, however, should be able to differentiate intentional and spontaneous eye movements. We report an electroencephalogram (EEG) marker that differentiates gaze fixations used for control from spontaneous fixations involved in visual exploration. Eight healthy participants played a game with their eye movements only. Their gaze-synchronized EEG data (fixation-related potentials, FRPs) were collected during game's control-on and control-off conditions. A slow negative wave with a maximum in the parietooccipital region was present in each participant's averaged FRPs in the control-on conditions and was absent or had much lower amplitude in the control-off condition. This wave was similar but not identical to stimulus-preceding negativity, a slow negative wave that can be observed during feedback expectation. Classification of intentional vs. spontaneous fixations was based on amplitude features from 13 EEG channels using 300 ms length segments free from electrooculogram contamination (200..500 ms relative to the fixation onset). For the first fixations in the fixation triplets required to make moves in the game, classified against control-off data, a committee of greedy classifiers provided 0.90 ± 0.07 specificity and 0.38 ± 0.14 sensitivity. Similar (slightly lower) results were obtained for the shrinkage LDA classifier. The second and third fixations in the triplets were classified at lower rate. We expect that, with improved feature sets and classifiers, a hybrid dwell-based Eye-Brain-Computer Interface (EBCI) can be built using the FRP difference between the intended and spontaneous fixations. If this direction of BCI development will be successful, such a multimodal interface may improve the fluency of interaction and can possibly become the basis for a new input device for paralyzed and healthy users, the EBCI “Wish Mouse”. |
Eelke Spaak; Yvonne Fonken; Ole Jensen; Floris P. Lange The neural mechanisms of prediction in visual search Journal Article In: Cerebral Cortex, vol. 26, no. 11, pp. 4327–4336, 2016. @article{Spaak2016, The speed of visual search depends on bottom-up stimulus features (e.g., we quickly locate a red item among blue distractors), but it is also facilitated by the presence of top-down perceptual predictions about the item. Here, we identify the nature, source, and neuronal substrate of the predictions that speed up resumed visual search. Human subjects were presented with a visual search array that was repeated up to 4 times, while brain activity was recorded using magnetoencephalography (MEG). Behaviorally, we observed a bimodal reaction time distribution for resumed visual search, indicating that subjects were extraordinarily rapid on a proportion of trials. MEG data demonstrated that these rapid-response trials were associated with a prediction of (1) target location, as reflected by alpha-band (8-12 Hz) lateralization; and (2) target identity, as reflected by beta-band (15-30 Hz) lateralization. Moreover, we show that these predictions are likely generated in a network consisting of medial superior frontal cortex and right temporo-parietal junction. These findings underscore the importance and nature of perceptual hypotheses for efficient visual search. |
Heng Ru May Tan; Joachim Gross; P. J. Uhlhaas MEG sensor and source measures of visually induced gamma-band oscillations are highly reliable Journal Article In: NeuroImage, vol. 137, pp. 34–44, 2016. @article{Tan2016, High frequency brain oscillations are associated with numerous cognitive and behavioral processes. Non-invasive measurements using electro-/magnetoencephalography (EEG/MEG) have revealed that high frequency neural signals are heritable and manifest changes with age as well as in neuropsychiatric illnesses. Despite the extensive use of EEG/MEG-measured neural oscillations in basic and clinical research, studies demonstrating test-retest reliability of power and frequency measures of neural signals remain scarce. Here, we evaluated the test-retest reliability of visually induced gamma (30-100 Hz) oscillations derived from sensor and source signals acquired over two MEG sessions. The study required participants (N = 13) to detect the randomly occurring stimulus acceleration while viewing a moving concentric grating. Sensor and source MEG measures of gamma-band activity yielded comparably strong reliability (average intraclass correlation |
Akiko Ikkai; Sangita Dandekar; Clayton E. Curtis Lateralization in alpha-band oscillations predicts the locus and spatial distribution of attention Journal Article In: PLoS ONE, vol. 11, no. 5, pp. e0154796, 2016. @article{Ikkai2016, Attending to a task-relevant location changes how neural activity oscillates in the alpha band (8-13Hz) in posterior visual cortical areas. However, a clear understanding of the relationships between top-down attention, changes in alpha oscillations in visual cortex, and attention performance are still poorly understood. Here, we tested the degree to which the posterior alpha power tracked the locus of attention, the distribution of attention, and how well the topography of alpha could predict the locus of attention. We recorded magnetoencephalographic (MEG) data while subjects performed an attention demanding visual discrimination task that dissociated the direction of attention from the direction of a saccade to indicate choice. On some trials, an endogenous cue predicted the target's location, while on others it contained no spatial information. When the target's location was cued, alpha power decreased in sensors over occipital cortex contralateral to the attended visual field. When the cue did not predict the target's location, alpha power again decreased in sensors over occipital cortex, but bilaterally, and increased in sensors over frontal cortex. Thus, the distribution and the topography of alpha reliably indicated the locus of covert attention. Together, these results suggest that alpha synchronization reflects changes in the excitability of populations of neurons whose receptive fields match the locus of attention. This is consistent with the hypothesis that alpha oscillations reflect the neural mechanisms by which top-down control of attention biases information processing and modulate the activity of neurons in visual cortex. |
P. J. López-Peréz; J. Dampuré; J. A. Hernández-Cabrera; H. A. Barber Semantic parafoveal-on-foveal effects and preview benefits in reading: Evidence from fixation related potentials Journal Article In: Brain and Language, vol. 162, pp. 29–34, 2016. @article{LopezPerez2016, During reading parafoveal information can affect the processing of the word currently fixated (parafovea-on-fovea effect) and words perceived parafoveally can facilitate their subsequent processing when they are fixated on (preview effect). We investigated parafoveal processing by simultaneously recording eye movements and EEG measures. Participants read word pairs that could be semantically associated or not. Additionally, the boundary paradigm allowed us to carry out the same manipulation on parafoveal previews that were displayed until reader's gaze moved to the target words. Event Related Potentials time-locked to the prime-preview presentation showed a parafoveal-on-foveal N400 effect. Fixation Related Potentials time locked to the saccade offset showed an N400 effect related to the prime-target relationship. Furthermore, this later effect interacted with the semantic manipulation of the previews, supporting a semantic preview benefit. These results demonstrate that at least under optimal conditions foveal and parafoveal information can be simultaneously processed and integrated. |
Ran Manor; Liran Mishali; Amir B. Geva Multimodal neural network for rapid serial visual presentation brain computer interface Journal Article In: Frontiers in Computational Neuroscience, vol. 10, pp. 130, 2016. @article{Manor2016, Brain computer interfaces allow users to preform various tasks using only the electrical activity of the brain. BCI applications often present the user a set of stimuli and record the corresponding electrical response. The BCI algorithm will then have to decode the acquired brain response and perform the desired task. In rapid serial visual presentation (RSVP) tasks, the subject is presented with a continuous stream of images containing rare target images among standard images, while the algorithm has to detect brain activity associated with target images. In this work, we suggest a multimodal neural network for RSVP tasks. The network operates on the brain response and on the initiating stimulus simultaneously, providing more information for the BCI application. We present two variants of the multimodal network, a supervised model, for the case when the targets are known in advanced, and a semi-supervised model for when the targets are unknown. We test the neural networks with a RSVP experiment on satellite imagery carried out with two subjects. The multimodal networks achieve a significant performance improvement in classification metrics. We visualize what the networks has learned and discuss the advantages of using neural network models for BCI applications. |
Tom R. Marshall; Sophie Esterer; Jim D. Herring; Til O. Bergmann; Ole Jensen On the relationship between cortical excitability and visual oscillatory responses-A concurrent tDCS-MEG study Journal Article In: NeuroImage, vol. 140, pp. 41–49, 2016. @article{Marshall2016, Neuronal oscillations in the alpha band (8–12 Hz) in visual cortex are considered to instantiate ‘attentional gating' via the inhibition of activity in regions representing task-irrelevant parts of space. In contrast, visual gamma-band activity (40–100 Hz) is regarded as representing a bottom-up drive from incoming visual information, with increased synchronisation producing a stronger feedforward impulse for relevant information. However, little is known about the direct relationship between excitability of the visual cortex and these oscillatory mechanisms. In this study we used transcranial direct current stimulation (tDCS) in an Oz–Cz montage in order to stimulate visual cortex, concurrently recording whole-brain oscillatory activity using magnetoencephalography (MEG) whilst participants performed a visual task known to produce strong modulations of alpha- and gamma-band activity. We found that visual stimuli produced expected modulations of alpha and gamma – presenting a moving annulus stimulus led to a strong gamma increase and alpha decrease – and that this pattern was observable both during active (anodal and cathodal) tDCS and sham tDCS. However, tDCS did not seem to produce systematic alterations of these oscillatory responses. The present study thus demonstrates that concurrent tDCS/MEG of the visual system is a feasible tool for investigating visual neuronal oscillations, and we discuss potential reasons for the apparent inability of tDCS to effectively change the amplitude of visual stimulus induced oscillatory responses in the current study. |
Catherine M. McMahon; Isabelle Boisvert; Peter Lissa; Louise Granger; Ronny Ibrahim; Chi Yhun Lo; Kelly Miles; Petra L. Graham Monitoring alpha oscillations and pupil dilation across a performance-intensity function Journal Article In: Frontiers in Psychology, vol. 7, pp. 745, 2016. @article{McMahon2016, Listening to degraded speech can be challenging and requires a continuous investment of cognitive resources, which is more challenging for those with hearing loss. However, while alpha power (8-12 Hz) and pupil dilation have been suggested as objective correlates of listening effort, it is not clear whether they assess the same cognitive processes involved, or other sensory and/or neurophysiological mechanisms that are associated with the task. Therefore, the aim of this study is to compare alpha power and pupil dilation during a sentence recognition task in 15 randomized levels of noise (-7dB to +7dB SNR) using highly intelligible (16 channel vocoded) and moderately intelligible (6 channel vocoded) speech. Twenty young normal hearing adults participated in the study; however, due to extraneous noise, data from 16 (10 females, 6 males; aged 19-28 years) was used in the EEG analysis and 10 in the pupil analysis. Behavioral testing of perceived effort and speech performance was assessed at 3 fixed SNRs per participant and was comparable to sentence recognition performance assessed in the physiological test session for both 16- and 6-channel vocoded sentences. Results showed a significant interaction between channel vocoding for both the alpha power and the pupil size changes. While both measures significantly decreased with more positive SNRs for the 16-channel vocoding, this was not observed with the 6-channel vocoding. The results of this study suggest that these measures may encode different processes involved in speech perception, which show similar trends for highly intelligible speech, but diverge for more spectrally degraded speech. The results to date suggest that these objective correlates of listening effort, and the cognitive processes involved in listening effort, are not yet sufficiently well understood to be used within a clinical setting. |
Anna Antonia Pape; Markus Siegel Motor cortex activity predicts response alternation during sensorimotor decisions Journal Article In: Nature Communications, vol. 7, pp. 13098, 2016. @article{Pape2016, Our actions are constantly guided by decisions based on sensory information. The motor cortex is traditionally viewed as the final output stage in this process, merely executing motor responses based on these decisions. However, it is not clear if, beyond this role, the motor cortex itself impacts response selection. Here, we report activity fluctuations over motor cortex measured using MEG, which are unrelated to choice content and predict responses to a visuomotor task seconds before decisions are made. These fluctuations are strongly influenced by the previous trial's response and predict a tendency to switch between response alternatives for consecutive decisions. This alternation behaviour depends on the size of neural signals still present from the previous response. Our results uncover a response-alternation bias in sensorimotor decision making. Furthermore, they suggest that motor cortex is more than an output stage and instead shapes response selection during sensorimotor decision making. |
Melanie R. Burke; R. O. Coats Dissociation of the rostral and dorsolateral prefrontal cortex during sequence learning in saccades: A TMS investigation Journal Article In: Experimental Brain Research, vol. 234, no. 2, pp. 597–604, 2016. @article{Burke2016, This experiment sought to find whether differences exist between the dorsolateral prefrontal cortex (DLPFC) and the medial rostral prefrontal cortex (MRPFC) for performing stimulus-independent and stimulus-oriented tasks, respectively. To find a causal relationship in these areas, we employed the use of trans-cranial magnetic stimulation (TMS). Prefrontal areas were stimulated whilst participants performed random or predictable sequence learning tasks at stimulus onset (1st presentation of the sequence only for both Random and Predictable), or during the inter-sequence interval. Overall, we found that during the predictable task a significant decrease in saccade latency, gain and duration was found when compared to the randomised conditions, as expected and observed previously. However, TMS stimulation in DLPFC during the delay in the predictive sequence learning task reduced this predictive ability by delaying the saccadic onset and generating abnormal reductions in saccadic gains during prediction. In contrast, we found that stimulation during a delay in MRPFC reversed the normal effects on peak velocity of the task with the predictive task revealing higher peak velocity than the randomised task. These findings provide causal evidence for independent functions of DLPFC and MRPFC in performing stimulus-independent processing during sequence learning in saccades. |
Sarah C. Krall; Lukas J. Volz; Eileen Oberwelland; Christian Grefkes; Gereon R. Fink; Kerstin Konrad The right temporoparietal junction in attention and social interaction: A transcranial magnetic stimulation study Journal Article In: Human Brain Mapping, vol. 37, no. 2, pp. 796–807, 2016. @article{Krall2016, The right temporoparietal junction (rTPJ) has been associated with the ability to reorient attention to unexpected stimuli and the capacity to understand others' mental states (theory of mind [ToM]/false belief). Using activation likelihood estimation meta-analysis we previously unraveled that the anterior rTPJ is involved in both, reorienting of attention and ToM, possibly indicating a more general role in attention shifting. Here, we used neuronavigated transcranial magnetic stimulation to directly probe the role of the rTPJ across attentional reorienting and false belief. Task performance in a visual cueing paradigm and false belief cartoon task was investigated after application of continuous theta burst stimulation (cTBS) over anterior rTPJ (versus vertex, for control). We found that attentional reorienting was significantly impaired after rTPJ cTBS compared with control. For the false belief task, error rates in trials demanding a shift in mental state significantly increased. Of note, a significant positive correlation indicated a close relation between the stimulation effect on attentional reorienting and false belief trials. Our findings extend previous neuroimaging evidence by indicating an essential overarching role of the anterior rTPJ for both cognitive functions, reorienting of attention and ToM. |
Romain Quentin; Seth E. Frankston; Marine Vernet; Monica N. Toba; Paolo Bartolomeo; Lorena Chanes; Antoni Valero-Cabré In: Cerebral Cortex, vol. 26, no. 6, pp. 2381–2390, 2016. @article{Quentin2016, Behavioral and electrophysiological studies in humans and non-human primates have correlated frontal high-beta activity with the orienting of endogenous attention and shown the ability of the latter function to modulate visual performance. We here combined rhythmic transcranial magnetic stimulation (TMS) and diffusion imaging to study the relation between frontal oscillatory activity and visual performance, and we associated these phenomena to a specific set of white matter pathways that in humans subtend attentional processes. High-beta rhythmic activity on the right frontal eye field (FEF) was induced with TMS and its causal effects on a contrast sensitivity function were recorded to explore its ability to improve visual detection performance across different stimulus contrast levels. Our results show that frequency-specific activity patterns engaged in the right FEF have the ability to induce a leftward shift of the psychometric function. This increase in visual performance across different levels of stimulus contrast is likely mediated by a contrast gain mechanism. Interestingly, microstructural measures of white matter connectivity suggest a strong implication of right fronto-parietal connectivity linking the FEF and the intraparietal sulcus in propagating high-beta rhythmic signals across brain networks and subtending top-down frontal influences on visual performance. |
Hongfang Wang; Eleanor Callaghan; Gerard Gooding-Williams; Craig McAllister; Klaus Kessler Rhythm makes the world go round: An MEG-TMS study on the role of right TPJ theta oscillations in embodied perspective taking Journal Article In: Cortex, vol. 75, pp. 68–81, 2016. @article{Wang2016e, While some aspects of social processing are shared between humans and other species, some aspects are not. The former seems to apply to merely tracking another's visual perspective in the world (i.e., what a conspecific can or cannot perceive), while the latter applies to perspective taking in form of mentally "embodying" another's viewpoint. Our previous behavioural research had indicated that only perspective taking, but not tracking, relies on simulating a body schema rotation into another's viewpoint. In the current study we employed Magnetoencephalography (MEG) and revealed that this mechanism of mental body schema rotation is primarily linked to theta oscillations in a wider brain network of body-schema, somatosensory and motor-related areas, with the right posterior temporo-parietal junction (pTPJ) at its core. The latter was reflected by a convergence of theta oscillatory power in right pTPJ obtained by overlapping the separately localised effects of rotation demands (angular disparity effect), cognitive embodiment (posture congruence effect), and basic body schema involvement (posture relevance effect) during perspective taking in contrast to perspective tracking. In a subsequent experiment we interfered with right pTPJ processing using dual pulse Transcranial Magnetic Stimulation (dpTMS) and observed a significant reduction of embodied processing. We conclude that right TPJ is the crucial network hub for transforming the embodied self into another's viewpoint, body and/or mind, thus, substantiating how conflicting representations between self and other may be resolved and potentially highlighting the embodied origins of high-level social cognition in general. |
Meghan B. Mitchell; Steven D. Shirk; Donald G. McLaren; Jessica S. Dodd; Ali Ezzati; Brandon A. Ally; Alireza Atri Recognition of faces and names: Multimodal physiological correlates of memory and executive function Journal Article In: Brain Imaging and Behavior, vol. 10, no. 2, pp. 408–423, 2016. @article{Mitchell2016, We sought to characterize electrophysiological, eye-tracking and behavioral correlates of face-name recognition memory in healthy younger adults using high-density electroencephalography (EEG), infrared eye-tracking (ET), and neuropsychological measures. Twenty-one participants first studied 40 face-name (FN) pairs; 20 were presented four times (4R) and 20 were shown once (1R). Recognition memory was assessed by asking participants to make old/new judgments for 80 FN pairs, of which half were previously studied items and half were novel FN pairs (N). Simultaneous EEG and ET recording were collected during recognition trials. Comparisons of event-related potentials (ERPs) for correctly identified FN pairs were compared across the three item types revealing classic ERP old/new effects including 1) relative positivity (1R > N) bi-frontally from 300 to 500 ms, reflecting enhanced familiarity, 2) relative positivity (4R > 1R and 4R > N) in parietal areas from 500 to 800 ms, reflecting enhanced recollection, and 3) late frontal effects (1R > N) from 1000 to 1800 ms in right frontal areas, reflecting post-retrieval monitoring. ET analysis also revealed significant differences in eye movements across conditions. Exploration of cross-modality relationships suggested associations between memory and executive function measures and the three ERP effects. Executive function measures were associated with several indicators of saccadic eye movements and fixations, which were also associated with all three ERP effects. This novel characterization of face-name recognition memory performance using simultaneous EEG and ET reproduced classic ERP and ET effects, supports the construct validity of the multimodal FN paradigm, and holds promise as an integrative tool to probe brain networks supporting memory and executive functioning. |
Robert M. Mok; Nicholas E. Myers; George Wallis; Anna C. Nobre Behavioral and neural markers of flexible attention over working memory in aging Journal Article In: Cerebral Cortex, vol. 26, no. 4, pp. 1831–1842, 2016. @article{Mok2016, Working memory (WM) declines as we age and, because of its fundamental role in higher order cognition, this can have highly deleterious effects in daily life. We investigated whether older individuals benefit from flexible orienting of attention within WM to mitigate cognitive decline. We measured magnetoencephalography (MEG) in older adults performing a WM precision task with cues during the maintenance period that retroactively predicted the location of the relevant items for performance (retro-cues). WM performance of older adults significantly benefitted from retro-cues. Whereas WM maintenance declined with age, retro-cues conferred strong attentional benefits. A model-based analysis revealed an increase in the probability of recalling the target, a lowered probability of retrieving incorrect items or guessing, and an improvement in memory precision. MEG recordings showed that retro-cues induced a transient lateralization of alpha (8-14 Hz) and beta (15-30 Hz) oscillatory power. Interestingly, shorter durations of alpha/beta lateralization following retro-cues predicted larger cueing benefits, reinforcing recent ideas about the dynamic nature of access to WM representations. Our results suggest that older adults retain flexible control over WM, but individual differences in control correspond to differences in neural dynamics, possibly reflecting the degree of preservation of control in healthy aging. |
Jana Annina Müller; Dorothea Wendt; Birger Kollmeier; Thomas Brand Comparing eye tracking with electrooculography for measuring individual sentence comprehension duration Journal Article In: PLoS ONE, vol. 11, no. 10, pp. e0164627, 2016. @article{Mueller2016b, The aim of this study was to validate a procedure for performing the audio-visual paradigm introduced by Wendt et al. (2015) with reduced practical challenges. The original paradigm records eye fixations using an eye tracker and calculates the duration of sentence comprehension based on a bootstrap procedure. In order to reduce practical challenges, we first reduced the measurement time by evaluating a smaller measurement set with fewer trials. The results of 16 listeners showed effects comparable to those obtained when testing the original full measurement set on a different collective of listeners. Secondly, we introduced electrooculography as an alternative technique for recording eye movements. The correlation between the results of the two recording techniques (eye tracker and electrooculography) was r = 0.97, indicating that both methods are suitable for estimating the processing duration of individual participants. Similar changes in processing duration arising from sentence complexity were found using the eye tracker and the electrooculography procedure. Thirdly, the time course of eye fixations was estimated with an alternative procedure, growth curve analysis, which is more commonly used in recent studies analyzing eye tracking data. The results of the growth curve analysis were compared with the results of the bootstrap procedure. Both analysis methods show similar processing durations. |
Malik M. Naeem Mannan; Shinjung Kim; Myung Yung Jeong; M. Ahmad Kamran Hybrid EEG—Eye tracker: Automatic identification and removal of eye movement and blink artifacts from electroencephalographic signal Journal Article In: Sensors, vol. 16, pp. 241, 2016. @article{NaeemMannan2016, Contamination of eye movement and blink artifacts in Electroencephalogram (EEG) recording makes the analysis of EEG data more difficult and could result in mislead findings. Efficient removal of these artifacts from EEG data is an essential step in improving classification accuracy to develop the brain-computer interface (BCI). In this paper, we proposed an automatic framework based on independent component analysis (ICA) and system identification to identify and remove ocular artifacts from EEG data by using hybrid EEG and eye tracker system. The performance of the proposed algorithm is illustrated using experimental and standard EEG datasets. The proposed algorithm not only removes the ocular artifacts from artifactual zone but also preserves the neuronal activity related EEG signals in non-artifactual zone. The comparison with the two state-of-the-art techniques namely ADJUST based ICA and REGICA reveals the significant improved performance of the proposed algorithm for removing eye movement and blink artifacts from EEG data. Additionally, results demonstrate that the proposed algorithm can achieve lower relative error and higher mutual information values between corrected EEG and artifact-free EEG data. |
Hamidreza Namazi; Vladimir V. Kulish; Amin Akrami The analysis of the influence of fractal structure of stimuli on fractal dynamics in fixational eye movements and EEG signal Journal Article In: Scientific Reports, vol. 6, pp. 26639, 2016. @article{Namazi2016, One of the major challenges in vision research is to analyze the effect of visual stimuli on human vision. However, no relationship has been yet discovered between the structure of the visual stimulus, and the structure of fixational eye movements. This study reveals the plasticity of human fixational eye movements in relation to the ‘complex' visual stimulus. We demonstrated that the fractal temporal structure of visual dynamics shifts towards the fractal dynamics of the visual stimulus (image). The results showed that images with higher complexity (higher fractality) cause fixational eye movements with lower fractality. Considering the brain, as the main part of nervous system that is engaged in eye movements, we analyzed the governed Electroencephalogram (EEG) signal during fixation. We have found out that there is a coupling between fractality of image, EEG and fixational eye movements. The capability observed in this research can be further investigated and applied for treatment of different vision disorders. |
Karly N. Neath-Tavares; Roxane J. Itier Neural processing of fearful and happy facial expressions during emotion-relevant and emotion-irrelevant tasks: A fixation-to-feature approach Journal Article In: Biological Psychology, vol. 119, pp. 122–140, 2016. @article{NeathTavares2016, Research suggests an important role of the eyes and mouth for discriminating facial expressions of emotion. A gaze-contingent procedure was used to test the impact of fixation to facial features on the neural response to fearful, happy and neutral facial expressions in an emotion discrimination (Exp.1) and an oddball detection (Exp.2) task. The N170 was the only eye-sensitive ERP component, and this sensitivity did not vary across facial expressions. In both tasks, compared to neutral faces, responses to happy expressions were seen as early as 100–120 ms occipitally, while responses to fearful expressions started around 150 ms, on or after the N170, at both occipital and lateral-posterior sites. Analyses of scalp topographies revealed different distributions of these two emotion effects across most of the epoch. Emotion processing interacted with fixation location at different times between tasks. Results suggest a role of both the eyes and mouth in the neural processing of fearful expressions and of the mouth in the processing of happy expressions, before 350 ms. |
2015 |
Hélène Devillez; Nathalie Guyader; Anne Guérin-Dugué An eye fixation–related potentials analysis of the P300 potential for fixations onto a target object when exploring natural scenes Journal Article In: Journal of Vision, vol. 15, no. 13, pp. 20, 2015. @article{Devillez2015, The P300 event-related potential has been extensively studied in electroencephalography with classical paradigms that force observers to not move their eyes. This potential is classically used to infer whether a target or a task-relevant stimulus was presented. Few researches have studied this potential through more ecological paradigms where observers were able to move their eyes. In this study, we examined with an ecological paradigm and an adapted methodology the P300 potential using a visual search task that involves eye movements to actively explore natural scenes and during which eye movements and electroencephalographic activity were coregistered. Averaging the electroencephalography signal time-locked to fixation onsets, a P300 potential was observed for fixations onto the target object but not for other fixations recorded for the same visual search or for fixations recorded during the free viewing without any task. Our approach consists of using control experimental conditions with similar eye movements to ensure that the P300 potential was attributable to the fact that the observer gazed at the target rather than to other factors such as eye movement pattern (the size of the previous saccade) or the ‘‘overlap issue'' between the potentials elicited by two successive fixations. We also proposed to model the time overlap issue of the potentials elicited by consecutive fixations with various durations. Our results show that the P300 potential can be studied in ecological situations without any constraint on the type of visual exploration, with some precautions in the interpretation of results due to the overlap issue. |
Mithun Diwakar; Deborah L. Harrington; Jun Maruta; Jamshid Ghajar; Fady El-Gabalawy; Laura Muzzatti; Maurizio Corbetta; Ming-Xiong X. Huang; Roland R. Lee Filling in the gaps: Anticipatory control of eye movements in chronic mild traumatic brain injury Journal Article In: NeuroImage: Clinical, vol. 8, pp. 210–223, 2015. @article{Diwakar2015, A barrier in the diagnosis of mild traumatic brain injury (mTBI) stems from the lack of measures that are adequately sensitive in detecting mild head injuries. MRI and CT are typically negative in mTBI patients with persistent symptoms of post-concussive syndrome (PCS), and characteristic difficulties in sustaining attention often go undetected on neuropsychological testing, which can be insensitive to momentary lapses in concentration. Conversely, visual tracking strongly depends on sustained attention over time and is impaired in chronic mTBI patients, especially when tracking an occluded target. This finding suggests deficient internal anticipatory control in mTBI, the neural underpinnings of which are poorly understood. The present study investigated the neuronal bases for deficient anticipatory control during visual tracking in 25 chronic mTBI patients with persistent PCS symptoms and 25 healthy control subjects. The task was performed while undergoing magnetoencephalography (MEG), which allowed us to examine whether neural dysfunction associated with anticipatory control deficits was due to altered alpha, beta, and/or gamma activity. Neuropsychological examinations characterized cognition in both groups. During MEG recordings, subjects tracked a predictably moving target that was either continuously visible or randomly occluded (gap condition). MEG source-imaging analyses tested for group differences in alpha, beta, and gamma frequency bands. The results showed executive functioning, information processing speed, and verbal memory deficits in the mTBI group. Visual tracking was impaired in the mTBI group only in the gap condition. Patients showed greater error than controls before and during target occlusion, and were slower to resynchronize with the target when it reappeared. Impaired tracking concurred with abnormal beta activity, which was suppressed in the parietal cortex, especially the right hemisphere, and enhanced in left caudate and frontaloral areas. Regional beta-amplitude demonstrated high classification accuracy (92%) compared to eye-tracking (65%) and neuropsychological variables (80%). These findings show that deficient internal anticipatory control in mTBI is associated with altered beta activity, which is remarkably sensitive given the heterogeneity of injuries. |
Benedikt V. Ehinger; Peter Konig; José P. Ossandón Predictions of visual content across eye movements and their modulation by inferred information Journal Article In: Journal of Neuroscience, vol. 35, no. 19, pp. 7403–7413, 2015. @article{Ehinger2015, The brain is proposed to operate through probabilistic inference, testing and refining predictions about the world. Here, we search for neural activity compatible with the violation of active predictions, learned from the contingencies between actions and the consequent changes in sensory input. We focused on vision, where eye movements produce stimuli shifts that could, in principle, be predicted. We compared, in humans, error signals to saccade-contingent changes of veridical and inferred inputs by contrasting the electroencephalographic activity after saccades to a stimulus presented inside or outside the blind spot. We observed early (<250 ms) and late (>250 ms) error signals after stimulus change, indicating the violation of sensory and associative predictions, respectively. Remarkably, the late response was diminished for blind-spot trials. These results indicate that predictive signals occur across multiple levels of the visual hierarchy, based on generative models that differentiate between signals that originate from the outside world and those that are inferred. |
Mónika Gergelyfi; Benvenuto Jacob; Etienne Olivier; Alexandre Zénon Dissociation between mental fatigue and motivational state during prolonged mental activity Journal Article In: Frontiers in Behavioral Neuroscience, vol. 9, pp. 176, 2015. @article{Gergelyfi2015, Mental fatigue (MF) is commonly observed following prolonged cognitive activity and can have major repercussions on the daily life of patients as well as healthy individuals. Despite its important impact, the cognitive processes involved in MF remain largely unknown. An influential hypothesis states that MF does not arise from a disruption of overused neural processes but, rather, is caused by a progressive decrease in motivation-related task engagement. Here, to test this hypothesis, we measured various neural, autonomic, psychometric and behavioral signatures of MF and motivation (EEG, ECG, pupil size, eye blinks, Skin conductance responses (SCRs), questionnaires and performance in a working memory (WM) task) in healthy volunteers, while MF was induced by Sudoku tasks performed for 120 min. Moreover extrinsic motivation was manipulated by using different levels of monetary reward. We found that, during the course of the experiment, the participants' subjective feeling of fatigue increased and their performance worsened while their blink rate and heart rate variability (HRV) increased. Conversely, reward-induced EEG, pupillometric and skin conductance signal changes, regarded as indicators of task engagement, remained constant during the experiment, and failed to correlate with the indices of MF. In addition, MF did not affect a simple reaction time task, despite the strong influence of extrinsic motivation on this task. Finally, alterations of the motivational state through monetary incentives failed to compensate the effects of MF. These findings indicate that MF in healthy subjects is not caused by an alteration of task engagement but is likely to be the consequence of a decrease in the efficiency, or availability, of cognitive resources. |
Tjerk P. Gutteling; Luc P. J. Selen; W. Pieter Medendorp Parallax-sensitive remapping of visual space in occipito-parietal alpha-band activity during whole-body motion Journal Article In: Journal of Neurophysiology, vol. 113, pp. 1574–1584, 2015. @article{Gutteling2015, Despite the constantly changing retinal image due to eye, head, and body movements, we are able to maintain a stable representation of the visual environment. Various studies on retinal image shifts caused by saccades have suggested that occipital and parietal areas correct for these perturbations by a gaze-centered remapping of the neural image. However, such a uniform, rotational, remapping mechanism cannot work during translations when objects shift on the retina in a more complex, depth-dependent fashion due to motion parallax. Here we tested whether the brain's activity patterns show parallax-sensitive remapping of remembered visual space during whole-body motion. Under continuous recording of electroencephalography (EEG), we passively translated human subjects while they had to remember the location of a world-fixed visual target, briefly presented in front of or behind the eyes' fixation point prior to the motion. Using a psychometric approach we assessed the quality of the memory update, which had to be made based on vestibular feedback and other extraretinal motion cues. All subjects showed a variable amount of parallax-sensitive updating errors, i.e., the direction of the errors depended on the depth of the target relative to fixation. The EEG recordings show a neural correlate of this parallax-sensitive remapping in the alpha-band power at occipito-parietal electrodes. At parietal electrodes, the strength of these alpha-band modulations correlated significantly with updating performance. These results suggest that alpha-band oscillatory activity reflects the time-varying updating of gaze-centered spatial information during parallax-sensitive remapping during whole-body motion. |
Wei He; Jon Brock; Blake W. Johnson Face processing in the brains of pre-school aged children measured with MEG Journal Article In: NeuroImage, vol. 106, pp. 317–327, 2015. @article{He2015a, There are two competing theories concerning the development of face perception: a late maturation account and an early maturation account. Magnetoencephalography (MEG) neuroimaging holds promise for adjudicating between the two opposing accounts by providing objective neurophysiological measures of face processing, with sufficient temporal resolution to isolate face-specific brain responses from those associated with other sensory, cognitive and motor processes. The current study used a customized child MEG system to measure M100 and M170 brain responses in 15 children aged three to six years while they viewed faces, cars and their phase-scrambled counterparts. Compared to adults tested using the same stimuli in a conventional MEG system, children showed significantly larger and later M100 responses. Children's M170 responses, derived by subtracting the responses to phase-scrambled images from the corresponding images (faces or cars) were delayed in latency but otherwise resembled the adult M170. This component has not been obtained in previous studies of young children tested using conventional adult MEG systems. However children did show a markedly reduced M170 response to cars in comparison to adults. This may reflect children's lack of expertise with cars relative to faces. Taken together, these data are in accord with recent behavioural and neuroimaging data that support early maturation of the basic face processing functions. |
Wei He; Marta I. Garrido; Paul F. Sowman; Jon Brock; Blake W. Johnson Development of effective connectivity in the core network for face perception Journal Article In: Human Brain Mapping, vol. 36, no. 6, pp. 2161–2173, 2015. @article{He2015b, This study measured effective connectivity within the core face network in young children using a paediatric magnetoencephalograph (MEG). Dynamic casual modeling (DCM) of brain responses was performed in a group of adults (N = 14) and a group of young children aged from 3 to 6 years (N = 15). Three candidate DCM models were tested, and the fits of the MEG data to the three models were compared at both individual and group levels. The results show that the connectivity structure of the core face network differs significantly between adults and children. Further, the relative strengths of face network connections were differentially modulated by experimental conditions in the two groups. These results support the interpretation that the core face network undergoes significant structural configuration and functional specialization between four years of age and adulthood. |
Jörn M. Horschig; Wouter Oosterheert; Robert Oostenveld; Ole Jensen Modulation of posterior alpha activity by spatial attention allows for controlling a continuous brain–computer interface Journal Article In: Brain Topography, vol. 28, no. 6, pp. 852–864, 2015. @article{Horschig2015, Here we report that the modulation of alpha activity by covert attention can be used as a control signal in an online brain-computer interface, that it is reliable, and that it is robust. Subjects were instructed to orient covert visual attention to the left or right hemifield. We decoded the direction of attention from the magnetoencephalogram by a template matching classifier and provided the classification outcome to the subject in real-time using a novel graphical user interface. Training data for the templates were obtained from a Posner-cueing task conducted just before the BCI task. Eleven subjects participated in four sessions each. Eight of the subjects achieved classification rates significantly above chance level. Subjects were able to significantly increase their performance from the first to the second session. Individual patterns of posterior alpha power remained stable throughout the four sessions and did not change with increased performance. We conclude that posterior alpha power can successfully be used as a control signal in brain-computer interfaces. We also discuss several ideas for further improving the setup and propose future research based on solid hypotheses about behavioral consequences of modulating neuronal oscillations by brain computer interfacing. |
Jamila Andoh; Reiko Matsushita; Robert J. Zatorre Asymmetric interhemispheric transfer in the auditory network: Evidence from TMS, resting-state fMRI, and diffusion imaging Journal Article In: Journal of Neuroscience, vol. 43, no. 43, pp. 14602–14611, 2015. @article{Andoh2015, Hemispheric asymmetries in human auditory cortical function and structure are still highly debated. Brain stimulation approaches can complement correlational techniques by uncovering causal influences. Previous studies have shown asymmetrical effects of transcranial magnetic stimulation (TMS) on task performance, but it is unclear whether these effects are task-specific or reflect intrinsic network properties. To test how modulation of auditory cortex (AC) influences functional networks and whether this influence is asymmetrical, the present study measured resting-state fMRI connectivity networks in 17 healthy volunteers before and immediately after TMS (continuous theta burst stimulation) to the left or right AC, and the vertex as a control. We also examined the relationship between TMS-induced interhemispheric signal propagation and anatomical properties of callosal auditory fibers as measured with diffusion-weighted MRI. We found that TMS to the right AC, but not the left, resulted in widespread connectivity decreases in auditory- and motor-related networks in the resting state. Individual differences in the degree of change in functional connectivity between auditory cortices after TMS applied over the right AC were negatively related to the volume of callosal auditory fibers. The findings show that TMS-induced network modulation occurs, even in the absence of an explicit task, and that the magnitude of the effect differs across individuals as a function of callosal structure, supporting a role for the corpus callosum in mediating functional asymmetry. The findings support theoretical models emphasizing hemispheric differences in network organization and are of practical significance in showing that brain stimulation studies need to take network-level effects into account. |
Dario Cazzoli; René M. Müri; Christopher Kennard; Clive R. Rosenthal The role of the right posterior parietal cortex in letter migration between words Journal Article In: Journal of Cognitive Neuroscience, vol. 27, no. 2, pp. 377–386, 2015. @article{Cazzoli2015a, When briefly presented with pairs of words, skilled readers can sometimes report words with migrated letters (e.g., they report hunt when presented with the words hint and hurt). This and other letter migration phenomena have been often used to investigate factors that influence reading such as letter position coding. However, the neural basis of letter migration is poorly understood. Previous evidence has implicated the right posterior parietal cortex (PPC) in processing visuospatial attributes and lexical properties during word reading. The aim of this study was to assess this putative role by combining an inhibitory TMS protocol with a letter migration paradigm, which was designed to examine the contributions of visuospatial attributes and lexical factors. Temporary interference with the right PPC led to three specific effects on letter migration. First, the number of letter migrations was significantly increased only in the group with active stimulation (vs. a sham stimulation group or a control group without stimulation), and there was no significant effect on other error types. Second, this effect occurred only when letter migration could result in a meaningful word (migration vs. control context). Third, the effect of active stimulation on the number of letter migrations was lateralized to target words presented on the left. Our study thus demonstrates that the right PPC plays a specific and causal role in the phenomenon of letter migration. The nature of this role cannot be explained solely in terms of visuospatial attention, rather it involves an interplay between visuospatial attentional and word reading-specific factors. |
Magdalena Chechlacz; Glyn W. Humphreys; Stamatios N. Sotiropoulos; Christopher Kennard; Dario Cazzoli Structural organization of the corpus callosum predicts attentional shifts after continuous theta burst stimulation Journal Article In: Journal of Neuroscience, vol. 35, no. 46, pp. 15353–15368, 2015. @article{Chechlacz2015, Repetitive transcranial magnetic stimulation (rTMS) applied over the right posterior parietal cortex (PPC) in healthy participants has been shown to trigger a significant rightward shift in the spatial allocation of visual attention, temporarily mimicking spatial deficits observed in neglect. In contrast, rTMS applied over the left PPC triggers a weaker or null attentional shift. However, large interindividual differences in responses to rTMS have been reported. Studies measuring changes in brain activation suggest that the effects of rTMS may depend on both interhemispheric and intrahemispheric interactions between cortical loci controlling visual attention. Here, we investigated whether variability in the structural organization of human white matter pathways subserving visual attention, as assessed by diffusion magnetic resonance imaging and tractography, could explain interindividual differences in the effects of rTMS. Most participants showed a rightward shift in the allocation of spatial attention after rTMS over the right intraparietal sulcus (IPS), but the size of this effect varied largely across participants. Conversely, rTMS over the left IPS resulted in strikingly opposed individual responses, with some participants responding with rightward and some with leftward attentional shifts. We demonstrate that microstructural and macrostructural variability within the corpus callosum, consistent with differential effects on cross-hemispheric interactions, predicts both the extent and the direction of the response to rTMS. Together, our findings suggest that the corpus callosum may have a dual inhibitory and excitatory function in maintaining the interhemispheric dynamics that underlie the allocation of spatial attention. |
Tom A. Graaf; Felix Duecker; Martin H. P. Fernholz; Alexander T. Sack Spatially specific vs. unspecific disruption of visual orientation perception using chronometric pre-stimulus TMS Journal Article In: Frontiers in Behavioral Neuroscience, vol. 9, pp. 5, 2015. @article{Graaf2015, Transcranial magnetic stimulation (TMS) over occipital cortex can impair visual processing. Such ‘TMS masking' has repeatedly been shown at several stimulus onset asynchronies (SOAs), with TMS pulses generally applied after the onset of a visual stimulus. Following increased interest in the neuronal state-dependency of visual processing, we recently explored the efficacy of TMS at ‘negative SOAs', when no visual processing can yet occur. We could reveal pre-stimulus TMS disruption, with results moreover hinting at two separate mechanisms in occipital cortex biasing subsequent orientation perception. Here we extended this work, including a chronometric design to map the temporal dynamics of spatially specific and unspecific mechanisms of state-dependent visual processing, while moreover controlling for TMS-induced pupil covering. TMS pulses applied 60-40 ms prior to a visual stimulus decreased orientation processing independent of stimulus location, while a local suppressive effect was found for TMS applied 30-10 ms pre-stimulus. These results contribute to our understanding of spatiotemporal mechanisms in occipital cortex underlying the state-dependency of visual processing, providing a basis for future work to link pre-stimulus TMS suppression effects to other known visual biasing mechanisms. |
Peter H. Donaldson; Caroline T. Gurvich; Joanne Fielding; Peter G. Enticott Exploring associations between gaze patterns and putative human mirror neuron system activity Journal Article In: Frontiers in Human Neuroscience, vol. 9, pp. 523, 2015. @article{Donaldson2015, The human mirror neuron system (MNS) is hypothesized to be crucial to social cognition. Given that key MNS-input regions such as the superior temporal sulcus are involved in biological motion processing, and mirror neuron activity in monkeys has been shown to vary with visual attention, aberrant MNS function may be partly attributable to atypical visual input. To examine the relationship between gaze pattern and interpersonal motor resonance (IMR; an index of putative MNS activity), healthy right-handed participants aged 18–40 (n = 26) viewed videos of transitive grasping actions or static hands, whilst the left primary motor cortex received transcranial magnetic stimulation. Motor- evoked potentials recorded in contralateral hand muscles were used to determine IMR. Participants also underwent eyetracking analysis to assess gaze patterns whilst viewing the same videos. No relationship was observed between predictive gaze and IMR. However, IMR was positively associated with fixation counts in areas of biological motion in the videos, and negatively associated with object areas. These findings are discussed with reference to visual influences on the MNS, and the possibility that MNS atypicalities might be influenced by visual processes such as aberrant gaze pattern. |
Pankhuri Malik; Joost C. Dessing; J. Douglas Crawford Role of early visual cortex in trans-saccadic memory of object features Journal Article In: Journal of Vision, vol. 15, no. 7, pp. 1–17, 2015. @article{Malik2015, Early visual cortex (EVC) participates in visual feature memory and the updating of remembered locations across saccades, but its role in the trans-saccadic integration of object features is unknown. We hypothesized that if EVC is involved in updating object features relative to gaze, feature memory should be disrupted when saccades remap an object representation into a simultaneously perturbed EVC site. To test this, we applied transcranial magnetic stimulation (TMS) over functional magnetic resonance imaging–localized EVC clusters corresponding to the bottom left/right visual quadrants (VQs). During experiments, these VQs were probed psychophysically by briefly presenting a central object (Gabor patch) while subjects fixated gaze to the right or left (and above). After a short memory interval, participants were required to detect the relative change in orientation of a re-presented test object at the same spatial location. Participants either sustained fixation during the memory interval (fixation task)ormade a horizontal saccade that either maintained or reversed the VQ of the object (saccade task). Three TMS pulses (coinciding with the pre-, peri-, and postsaccade intervals) were applied to the left or right EVC. This had no effect when (a) fixation was maintained, (b) saccades kept the object in the same VQ, or (c) the EVC quadrant corresponding to the first object was stimulated. However, as predicted, TMS reduced performance when saccades (especially larger saccades) crossed the remembered object location and brought it into the VQ corresponding to the TMS site. This suppression effect was statistically significant for leftward saccades and followed a weaker trend for rightward saccades. These causal results are consistent with the idea that EVC is involved in the gaze-centered updating of object features for trans-saccadic memory and perception. |
Tom R. Marshall; Til Ole Bergmann; Ole Jensen Frontoparietal structural connectivity mediates the top-down control of neuronal synchronization associated with selective attention Journal Article In: PLoS Biology, vol. 13, no. 10, pp. e1002272, 2015. @article{Marshall2015, Neuronal synchronization reflected by oscillatory brain activity has been strongly implicated in the mechanisms supporting selective gating. We here aimed at identifying the anatomical pathways in humans supporting the top-down control of neuronal synchronization. We first collected diffusion imaging data using magnetic resonance imaging to identify the medial branch of the superior longitudinal fasciculus (SLF), a white-matter tract connecting frontal control areas to parietal regions. We then quantified the modulations in oscillatory activity using magnetoencephalography in the same subjects performing a spatial attention task. We found that subjects with a stronger SLF volume in the right compared to the left hemi- sphere (or vice versa) also were the subjects who had a better ability to modulate right com- pared to left hemisphere alpha and gamma band synchronization, with the latter also predicting biases in reaction time. Our findings implicate the medial branch of the SLF in mediating top-down control of neuronal synchronization in sensory regions that support selective attention. |
Petra Vetter; Marie-Hélène Grosbras; Lars Muckli TMS over V5 disrupts motion prediction Journal Article In: Cerebral Cortex, vol. 25, no. 4, pp. 1052–1059, 2015. @article{Vetter2015, Given the vast amount of sensory information the brain has to deal with, predicting some of this information based on the current context is a resource-efficient strategy. The framework of predictive coding states that higher-level brain areas generate a predictive model to be communicated via feedback connections to early sensory areas. Here, we directly tested the necessity of a higher-level visual area, V5, in this predictive processing in the context of an apparent motion paradigm. We flashed targets on the apparent motion trace in-time or out-of-time with the predicted illusory motion token. As in previous studies, we found that predictable in-time targets were better detected than unpredictable out-of-time targets. However, when we applied functional magnetic resonance imaging-guided, double-pulse transcranial magnetic stimulation (TMS) over left V5 at 13-53 ms before target onset, the detection advantage of in-time targets was eliminated; this was not the case when TMS was applied over the vertex. Our results are causal evidence that V5 is necessary for a prediction effect, which has been shown to modulate V1 activity (Alink et al. 2010). Thus, our findings suggest that information processing between V5 and V1 is crucial for visual motion prediction, providing experimental support for the predictive coding framework. |
Alexandre Zenon; Mariam Sidibe; Etienne Olivier Disrupting the supplementary motor area makes physical effort appear less effortful Journal Article In: Journal of Neuroscience, vol. 35, no. 23, pp. 8737–8744, 2015. @article{Zenon2015, The perception of physical effort is relatively unaffected by the suppression of sensory afferences, indicating that this function relies mostly on the processing of the central motor command. Neural signals in the supplementary motor area (SMA) correlate with the intensity of effort, suggesting that the motor signal involved in effort perception could originate from this area, but experimental evidence supporting this view is still lacking. Here, we tested this hypothesis by disrupting neural activity in SMA, in primary motor cortex (M1), or in a control site by means of continuous theta-burst transcranial magnetic stimulation, while measuring effort perception during grip forces of different intensities. After each grip force exertion, participants had the opportunity to either accept or refuse to replicate the same effort for varying amounts of reward. In addition to the subjective rating of perceived exertion, effort perception was estimated on the basis of the acceptance rate, the effort replication accuracy, the influence of the effort exerted in trial t on trial t+1, and pupil dilation. We found that disruption of SMA activity, but not of M1, led to a consistent decrease in effort perception, whatever the measure used to assess it. Accordingly, we modeled effort perception in a structural equation model and found that only SMA disruption led to a significant alteration of effort perception. These findings indicate that effort perception relies on the processing of a signal originating from motor-related neural circuits upstream of M1 and that SMA is a key node of this network. |
Paul Metzner; Titus Malsburg; Shravan Vasishth; Frank Rösler Brain responses to world knowledge violations: A comparison of stimulus- and fixation-triggered event-related potentials and neural oscillations Journal Article In: Journal of Cognitive Neuroscience, vol. 27, no. 5, pp. 1017–1028, 2015. @article{Metzner2015, Recent research has shown that brain potentials time-locked to fixations in natural reading can be similar to brain potentials recorded during rapid serial visual presentation (RSVP). We attempted two replications of Hagoort, Hald, Bastiaansen, and Petersson [Hagoort, P., Hald, L., Bastiaansen, M., & Petersson, K. M. Integration of word meaning and world knowledge in language comprehension. Science, 304, 438-441, 2004] to determine whether this correspondence also holds for oscillatory brain responses. Hagoort et al. reported an N400 effect and synchronization in the theta and gamma range following world knowledge violations. Our first experiment (n = 32) used RSVP and replicated both the N400 effect in the ERPs and the power increase in the theta range in the time-frequency domain. In the second experiment (n = 49), participants read the same materials freely while their eye movements and their EEG were monitored. First fixation durations, gaze durations, and regression rates were increased, and the ERP showed an N400 effect. An analysis of time-frequency representations showed synchronization in the delta range (1-3 Hz) and desynchronization in the upper alpha range (11-13 Hz) but no theta or gamma effects. The results suggest that oscillatory EEG changes elicited by world knowledge violations are different in natural reading and RSVP. This may reflect differences in how representations are constructed and retrieved from memory in the two presentation modes. |
Nicholas E. Myers; Gustavo Rohenkohl; Valentin Wyart; Mark W. Woolrich; Anna C. Nobre; Mark G. Stokes Testing sensory evidence against mnemonic templates Journal Article In: eLife, vol. 4, pp. 1–25, 2015. @article{Myers2015, Most perceptual decisions require comparisons between current input and an internal template. Classic studies propose that templates are encoded in sustained activity of sensory neurons. However, stimulus encoding is itself dynamic, tracing a complex trajectory through activity space. Which part of this trajectory is pre-activated to reflect the template? Here we recorded magneto- and electroencephalography during a visual target-detection task, and used pattern analyses to decode template, stimulus, and decision-variable representation. Our findings ran counter to the dominant model of sustained pre-activation. Instead, template information emerged transiently around stimulus onset and quickly subsided. Cross-generalization between stimulus and template coding, indicating a shared neural representation, occurred only briefly. Our results are compatible with the proposal that template representation relies on a matched filter, transforming input into task-appropriate output. This proposal was consistent with a signed difference response at the perceptual decision stage, which can be explained by a simple neural model. |
Nicholas E. Myers; Lena Walther; George Wallis; Mark G. Stokes; Anna C. Nobre In: Journal of Cognitive Neuroscience, vol. 27, no. 3, pp. 492–508, 2015. @article{Myers2015a, Working memory (WM) is strongly influenced by attention. In visual WM tasks, recall performance can be improved by an attention-guiding cue presented before encoding (precue) or during maintenance (retrocue). Although precues and retro- cues recruit a similar frontoparietal control network, the two are likely to exhibit some processing differences, because pre- cues invite anticipation of upcoming information whereas retro- cues may guide prioritization, protection, and selection of information already in mind. Here we explored the behavioral and electrophysiological differences between precueing and retrocueing in a new visual WM task designed to permit a direct comparison between cueing conditions. We found marked differences in ERP profiles between the precue and retrocue conditions. In line with precues primarily generating an anti- cipatory shift of attention toward the location of an upcoming item, we found a robust lateralization in late cue-evoked po- tentials associated with target anticipation. Retrocues elicited a different pattern of ERPs that was compatible with an early selec- tion mechanism, but not with stimulus anticipation. In contrast to the distinct ERP patterns, alpha-band (8–14 Hz) lateralization was indistinguishable between cue types (reflecting, in both conditions, the location of the cued item). We speculate that, whereas alpha-band lateralization after a precue is likely to enable anticipatory attention, lateralization after a retrocue may instead enable the controlled spatiotopic access to recently encoded visual information |
Karly N. Neath; Roxane J. Itier Fixation to features and neural processing of facial expressions in a gender discrimination task Journal Article In: Brain and Cognition, vol. 99, pp. 97–111, 2015. @article{Neath2015, Early face encoding, as reflected by the N170 ERP component, is sensitive to fixation to the eyes. Whether this sensitivity varies with facial expressions of emotion and can also be seen on other ERP components such as P1 and EPN, was investigated. Using eye-tracking to manipulate fixation on facial features, we found the N170 to be the only eye-sensitive component and this was true for fearful, happy and neutral faces. A different effect of fixation to features was seen for the earlier P1 that likely reflected general sensitivity to face position. An early effect of emotion (~120. ms) for happy faces was seen at occipital sites and was sustained until ~350. ms post-stimulus. For fearful faces, an early effect was seen around 80. ms followed by a later effect appearing at ~150. ms until ~300. ms at lateral posterior sites. Results suggests that in this emotion-irrelevant gender discrimination task, processing of fearful and happy expressions occurred early and largely independently of the eye-sensitivity indexed by the N170. Processing of the two emotions involved different underlying brain networks active at different times. |
Valentin Wyart; Nicholas E. Myers; Christopher Summerfield Neural mechanisms of human perceptual choice under focused and divided attention Journal Article In: Journal of Neuroscience, vol. 35, no. 8, pp. 3485–3498, 2015. @article{Wyart2015, Perceptual decisions occur after the evaluation and integration of momentary sensory inputs, and dividing attention between spatially disparate sources of information impairs decision performance. However, it remains unknown whether dividing attention degrades the precision of sensory signals, precludes their conversion into decision signals, or dampens the integration of decision information toward an appropriate response. Here we recorded human electroencephalographic (EEG) activity while participants categorized one of two simultaneous and independent streams of visual gratings according to their average tilt. By analyzing trial-by-trial correlations between EEG activity and the information offered by each sample, we obtained converging behavioral and neural evidence that dividing attention between left and right visual fields does not dampen the encoding of sensory or decision information. Under divided attention, momentary decision information from both visual streams was encoded in slow parietal signals without interference but was lost downstream during their integration as reflected in motor mu- and beta-band (10-30 Hz) signals, resulting in a "leaky" accumulation process that conferred greater behavioral influence to more recent samples. By contrast, sensory inputs that were explicitly cued as irrelevant were not converted into decision signals. These findings reveal that a late cognitive bottleneck on information integration limits decision performance under divided attention, and places new capacity constraints on decision-theoretic models of information integration under cognitive load. |
Jeffrey Weiler; Cameron D. Hassall; Olave E. Krigolson; Matthew Heath The unidirectional prosaccade switch-cost: Electroencephalographic evidence of task-set inertia in oculomotor control Journal Article In: Behavioural Brain Research, vol. 278, pp. 323–329, 2015. @article{Weiler2015, The execution of an antisaccade selectively increases the reaction time (RT) of a subsequent prosaccade (the unidirectional prosaccade switch-cost). To explain this finding, the task-set inertia hypothesis asserts that an antisaccade requires a cognitively mediated non-standard task-set that persists inertially and delays the planning of a subsequent prosaccade. The present study sought to directly test the theoretical tenets of the task-set inertia hypothesis by examining the concurrent behavioural and the event-related brain potential (ERP) data associated with the unidirectional prosaccade switch-cost. Participants pseudo-randomly alternated between pro- and antisaccades while electroencephalography (EEG) data were recorded. As expected, the completion of an antisaccade selectively increased the RT of a subsequent prosaccade, whereas the converse switch did not influence RTs. Thus, the behavioural results demonstrated the unidirectional prosaccade switch-cost. In terms of the ERP findings, we observed a reliable change in the amplitude of the P3 - time-locked to task-instructions - when trials were switched from a prosaccade to an antisaccade; however, no reliable change was observed when switching from an antisaccade to a prosaccade. This is a salient finding because extensive work has shown that the P3 provides a neural index of the task-set required to execute a to-be-completed response. As such, results showing that prosaccades completed after antisaccades exhibited increased RTs in combination with a P3 amplitude comparable to antisaccades provides convergent evidence that the unidirectional prosaccade switch-cost is attributed to the persistent activation of a non-standard antisaccade task-set. |
George Wallis; Mark Stokes; Helena Cousijn; Mark W. Woolrich; Anna C. Nobre Frontoparietal and cingulo-opercular networks play dissociable roles in control of working memory Journal Article In: Journal of Cognitive Neuroscience, vol. 27, pp. 2019–2034, 2015. @article{Wallis2015a, We used magnetoencephalography to characterize the spatiotemporal dynamics of cortical activity during top–down control of working memory (WM). fMRI studies have previously implicated both the frontoparietal and cingulo-opercular networks in control over WM, but their respective contributions are unclear. In our task, spatial cues indicating the relevant item in a WM array occurred either before the memory array or during the maintenance period, providing a direct comparison between prospective and retrospective control of WM. We found that in both cases a frontoparietal network activated following the cue, but following retrocues this activation was transient and was succeeded by a cinguloopercular network activation. We also characterized the time course of top–down modulation of alpha activity in visual/parietal cortex. This modulation was transient following retrocues, occurring in parallel with the frontoparietal network activation. We suggest that the frontoparietal network is responsible for top–down modulation of activity in sensory cortex during both preparatory attention and orienting within memory. In contrast, the cinguloopercular network plays a more downstream role in cognitive control, perhaps associated with output gating of memory |
Virginie Wassenhove; Lukasz Grzeczkowski Visual-induced expectations modulate auditory cortical responses Journal Article In: Frontiers in Neuroscience, vol. 9, pp. 11, 2015. @article{Wassenhove2015, Active sensing has important consequences on multisensory processing (Schroeder et al., 2010). Here, we asked whether in the absence of saccades, the position of the eyes and the timing of transient color changes of visual stimuli could selectively affect the excitability of auditory cortex by predicting the "where" and the "when" of a sound, respectively. Human participants were recorded with magnetoencephalography (MEG) while maintaining the position of their eyes on the left, right, or center of the screen. Participants counted color changes of the fixation cross while neglecting sounds which could be presented to the left, right, or both ears. First, clear alpha power increases were observed in auditory cortices, consistent with participants' attention directed to visual inputs. Second, color changes elicited robust modulations of auditory cortex responses ("when" prediction) seen as ramping activity, early alpha phase-locked responses, and enhanced high-gamma band responses in the contralateral side of sound presentation. Third, no modulations of auditory evoked or oscillatory activity were found to be specific to eye position. Altogether, our results suggest that visual transience can automatically elicit a prediction of "when" a sound will occur by changing the excitability of auditory cortices irrespective of the attended modality, eye position or spatial congruency of auditory and visual events. To the contrary, auditory cortical responses were not significantly affected by eye position suggesting that "where" predictions may require active sensing or saccadic reset to modulate auditory cortex responses, notably in the absence of spatial orientation to sounds. |
Lingling Yang; Howard Leung; Markus Plank; Joe Snider; Howard Poizner EEG activity during movement planning encodes upcoming peak speed and acceleration and improves the accuracy in predicting hand kinematics Journal Article In: IEEE Journal of Biomedical and Health Informatics, vol. 19, no. 1, pp. 22–28, 2015. @article{Yang2015, The relationship between movement kinematics and human brain activity is an important and fundamental question for the development of neural prosthesis. The peak velocity and the peak acceleration could best reflect the feedforward-type movement thus it is worthwhile to investigate them further. Most related studies focused on the correlation between kinematics and brain activity during the movement execution or imagery. However, human movement is the result of the motor planning phase as well as the execution phase and researchers have demonstrated that statistical correlations exist between EEG activity during the motor planning and the peak velocity and the peak acceleration using grand-average analysis. In this paper, we examined whether the correlations were concealed in trial-to-trial decoding from the low signal-to-noise ratio of EEG activity. The alpha and beta powers from the movement planning phase were combined with the alpha and beta powers from the movement execution phase to predict the peak tangential speed and acceleration. The results showed that EEG activity from the motor planning phase could also predict the peak speed and the peak acceleration with a reasonable accuracy. Furthermore, the decoding accuracy of the peak speed and the peak acceleration could both be improved by combining band powers from the motor planning phase with the band powers from the movement execution. |
M. Isabel Vanegas; Annabelle Blangero; Simon P. Kelly Electrophysiological indices of surround suppression in humans Journal Article In: Journal of Neurophysiology, vol. 113, no. 4, pp. 1100–1109, 2015. @article{Vanegas2015, Surround suppression is a well-known example of contextual interaction in visual cortical neurophysiology, whereby the neural response to a stimulus presented within a neuron's classical receptive field is suppressed by surrounding stimuli. Human psychophysical reports present an obvious analog to the effects seen at the single-neuron level: stimuli are perceived as lower-contrast when embedded in a surround. Here we report on a visual paradigm that provides relatively direct, straightforward indices of surround suppression in human electrophysiology, enabling us to reproduce several well-known neurophysiological and psychophysical effects, and to conduct new analyses of temporal trends and retinal location effects. Steady-state visual evoked potentials (SSVEP) elicited by flickering “foreground” stimuli were measured in the context of various static surround patterns. Early visual cortex geometry and retinotopic organization were exploited to enhance SSVEP amplitude. The foreground response was strongly suppressed as a monotonic function of surround contrast. Further- more, suppression was stronger for surrounds of matching orientation than orthogonally-oriented ones, and stronger at peripheral than foveal locations. These patterns were reproduced in psychophysical reports of perceived contrast, and peripheral electrophysiological suppression effects correlated with psychophysical effects across subjects. Temporal analysis of SSVEP amplitude revealed short-term contrast adaptation effects that caused the foreground signal to either fall or grow over time, depending on the relative contrast of the surround, consistent with stronger adaptation of the suppressive drive. This electrophysiology paradigm has clinical potential in indexing not just visual deficits but possibly gain control deficits expressed more widely in the disordered brain. |
Yuka O. Okazaki; Jörn M. Horschig; Lisa Luther; Robert Oostenveld; Ikuya Murakami; Ole Jensen Real-time MEG neurofeedback training of posterior alpha activity modulates subsequent visual detection performance Journal Article In: NeuroImage, vol. 107, pp. 323–332, 2015. @article{Okazaki2015, It has been demonstrated that alpha activity is lateralized when attention is directed to the left or right visual hemifield. We investigated whether real-time neurofeedback training of the alpha lateralization enhances participants' ability to modulate posterior alpha lateralization and causes subsequent short-term changes in visual detection performance. The experiment consisted of three phases: (i) pre-training assessment, (ii) neurofeedback phase and (iii) post-training assessment. In the pre- and post-training phases we measured the threshold to covertly detect a cued faint Gabor stimulus presented in the left or right hemifield. During magnetoencephalography (MEG) neurofeedback, two face stimuli superimposed with noise were presented bilaterally. Participants were cued to attend to one of the hemifields. The transparency of the superimposed noise and thus the visibility of the stimuli were varied according to the momentary degree of hemispheric alpha lateralization. In a double-blind procedure half of the participants were provided with sham feedback. We found that hemispheric alpha lateralization increased with the neurofeedback training; this was mainly driven by an ipsilateral alpha increase. Surprisingly, comparing pre- to post-training, detection performance decreased for a Gabor stimulus presented in the hemifield that was un-attended during neurofeedback. This effect was not observed in the sham group. Thus, neurofeedback training alters alpha lateralization, which in turn decreases performances in the untrained hemifield. Our findings suggest that alpha oscillations play a causal role for the allocation of attention. Furthermore, our neurofeedback protocol serves to reduce the detection of unattended visual information and could therefore be of potential use for training to reduce distractibility in attention deficit patients, but also highlights that neurofeedback paradigms can have negative impact on behavioral performance and should be applied with caution. |
Ryszard Auksztulewicz; Karl J. Friston Attentional enhancement of auditory mismatch responses: A DCM/MEG study Journal Article In: Cerebral Cortex, vol. 25, no. 11, pp. 4273–4283, 2015. @article{Auksztulewicz2015, Despite similar behavioral effects, attention and expectation influence evoked responses differently: Attention typically enhances event-related responses, whereas expectation reduces them. This dissociation has been reconciled under predictive coding, where prediction errors are weighted by precision associated with attentional modulation. Here, we tested the predictive coding account of attention and expectation using magnetoencephalography and modeling. Temporal attention and sensory expectation were orthogonally manipulated in an auditory mismatch paradigm, revealing opposing effects on evoked response amplitude. Mismatch negativity (MMN) was enhanced by attention, speaking against its supposedly pre-attentive nature. This interaction effect was modeled in a canonical microcircuit using dynamic causal modeling, comparing models with modulation of extrinsic and intrinsic connectivity at different levels of the auditory hierarchy. While MMN was explained by recursive interplay of sensory predictions and prediction errors, attention was linked to the gain of inhibitory interneurons, consistent with its modulation of sensory precision. |
Dominik R. Bach; Nicholas Furl; Gareth Barnes; Raymond J. Dolan Sustained magnetic responses in temporal cortex reflect instantaneous significance of approaching and receding sounds Journal Article In: PLoS ONE, vol. 10, no. 7, pp. e0134060, 2015. @article{Bach2015, Rising sound intensity often signals an approaching sound source and can serve as a powerful warning cue, eliciting phasic attention, perception biases and emotional responses. How the evaluation of approaching sounds unfolds over time remains elusive. Here, we capitalised on the temporal resolution of magnetoencephalograpy (MEG) to investigate in humans a dynamic encoding of perceiving approaching and receding sounds. We compared magnetic responses to intensity envelopes of complex sounds to those of white noise sounds, in which intensity change is not perceived as approaching. Sustained magnetic fields over temporal sensors tracked intensity change in complex sounds in an approximately linear fashion, an effect not seen for intensity change in white noise sounds, or for overall intensity. Hence, these fields are likely to track approach/recession, but not the apparent (instantaneous) distance of the sound source, or its intensity as such. As a likely source of this activity, the bilateral inferior temporal gyrus and right temporo-parietal junction emerged. Our results indicate that discrete temporal cortical areas parametrically encode behavioural significance in moving sound sources where the signal unfolded in a manner reminiscent of evidence accumulation. This may help an understanding of how acoustic percepts are evaluated as behaviourally relevant, where our results highlight a crucial role of cortical areas. |
Snigdha Banerjee; Hans Peter Frey; Sophie Molholm; John J. Foxe In: European Journal of Neuroscience, vol. 41, no. 6, pp. 818–834, 2015. @article{Banerjee2015, The voluntary allocation of attention to environmental inputs is a crucial mechanism of healthy cognitive functioning, and is probably influenced by an observer's level of interest in a stimulus. For example, an individual who is passionate about soccer but bored by botany will obviously be more attentive at a soccer match than an orchid show. The influence of monetary rewards on attention has been examined, but the impact of more common motivating factors (i.e. the level of interest in the materials under observation) remains unclear, especially during development. Here, stimulus sets were designed based on survey measures of the level of interest of adolescent participants in several item classes. High-density electroencephalography was recorded during a cued spatial attention task in which stimuli of high or low interest were presented in separate blocks. The motivational impact on performance of a spatial attention task was assessed, along with event-related potential measures of anticipatory top-down attention. As predicted, performance was improved for the spatial target detection of high interest items. Further, the impact of motivation was observed in parieto-occipital processes associated with anticipatory top-down spatial attention. The anticipatory activity over these regions was also increased for high vs. low interest stimuli, irrespective of the direction of spatial attention. The results also showed stronger anticipatory attentional and motivational modulations over the right vs. left parieto-occipital cortex. These data suggest that motivation enhances top-down attentional processes, and can independently shape activations in sensory regions in anticipation of events. They also suggest that attentional functions across hemispheres may not fully mature until late adolescence. |
Brett C. Bays; Kristina M. Visscher; Christophe C. Le Dantec; Aaron R. Seitz Alpha-band EEG activity in perceptual learning Journal Article In: Journal of Vision, vol. 15, no. 10, pp. 1–12, 2015. @article{Bays2015, In studies of perceptual learning (PL), subjects are typically highly trained across many sessions to achieve perceptual benefits on the stimuli in those tasks. There is currently significant debate regarding what sources of brain plasticity underlie these PL-based learning improvements. Here we investigate the hypothesis that PL, among other mechanisms, leads to task automaticity, especially in the presence of the trained stimuli. To investigate this hypothesis, we trained participants for eight sessions to find an oriented target in a field of near-oriented distractors and examined alpha-band activity, which modulates with attention to visual stimuli, as a possible measure of automaticity. Alpha-band activity was acquired via electroencephalogram (EEG), before and after training, as participants performed the task with trained and untrained stimuli. Results show that participants underwent significant learning in this task (as assessed by threshold, accuracy, and reaction time improvements) and that alpha power increased during the pre-stimulus period and then underwent greater desynchronization at the time of stimulus presentation following training. However, these changes in alpha-band activity were not specific to the trained stimuli, with similar patterns of posttraining alpha power for trained and untrained stimuli. These data are consistent with the view that participants were more efficient at focusing resources at the time of stimulus presentation and are consistent with a greater automaticity of task performance. These findings have implications for PL, as transfer effects from trained to untrained stimuli may partially depend on differential effort of the individual at the time of stimulus processing. |
Nathan Caruana; Peter Lissa; Genevieve McArthur The neural time course of evaluating self-initiated joint attention bids Journal Article In: Brain and Cognition, vol. 98, pp. 43–52, 2015. @article{Caruana2015a, Background: During interactions with other people, we constantly evaluate the significance of our social partner's gaze shifts in order to coordinate our behaviour with their perspective. In this study, we used event-related potentials (ERPs) to investigate the neural time course of evaluating gaze shifts that signal the success of self-initiated joint attention bids. Method: Nineteen participants were allocated to a "social" condition, in which they played a cooperative game with an anthropomorphic virtual character whom they believed was controlled by a human partner in a nearby laboratory. Participants were required to initiate joint attention towards a target. In response, the virtual partner shifted his gaze congruently towards the target - thus achieving joint attention - or incongruently towards a different location. Another 19 participants completed the same task in a non-social "control" condition, in which arrows, believed to be controlled by a computer program, pointed at a location that was either congruent or incongruent with the participant's target fixation. Results: In the social condition, ERPs to the virtual partner's incongruent gaze shifts evoked significantly larger P350 and P500 peaks compared to congruent gaze shifts. This P350 and P500 morphology was absent in both the congruent and incongruent control conditions. Discussion: These findings are consistent with previous claims that gaze shifts differing in their social significance modulate central-parietal ERPs 350. ms following the onset of the gaze shift. Our control data highlights the social specificity of the observed P350 effect, ruling out explanations pertaining to attention modulation or error detection. |
Evy Cleeren; Cindy Casteels; Karolien Goffin; Peter Janssen; Wim Van Paesschen Ictal perfusion changes associated with seizure progression in the amygdala kindling model in the rhesus monkey Journal Article In: Epilepsia, vol. 56, no. 9, pp. 1366–1375, 2015. @article{Cleeren2015, OBJECTIVE: Amygdala kindling is a widely used animal model for studying mesial temporal lobe epileptogenesis. In the macaque monkey, electrical amygdala kindling develops slowly and provides an opportunity for investigating ictal perfusion changes during epileptogenesis. METHODS: Two rhesus monkeys were electrically kindled through chronically implanted electrodes in the right amygdala over a period of 16 and 17 months. Ictal perfusion single photon emission computed tomography (SPECT) imaging was performed during each of the four predefined clinical stages. RESULTS: Afterdischarge duration increased slowly over 477 days for monkey K and 515 days for monkey S (18 ± 8 s in stage I; 52 ± 13 s in stage IV). During this time, the animals progressed through four clinical stages ranging from interrupting ongoing behavior to bilateral convulsions. Ictal SPECT perfusion imaging showed well-localized but widely distributed regions of hyperperfusion and hypoperfusion, in both cortical and subcortical structures, at every seizure stage. A large portion of the ictal network was involved in the early stages of epileptogenesis and subsequently expanded over time as seizure severity evolved. SIGNIFICANCE: Our data indicate that the different mesial temporal lobe seizure types occur within a common network affecting several parts of the brain, and that seizure severity may be determined by seizure-induced epileptogenesis within a bihemispheric network that is implicated from the start of the process. |