EyeLink EEG / fNIRS / TMS Publications
All EyeLink EEG, fNIRS, and TMS research publications (with concurrent eye tracking) up until 2022 (with early 2023s) 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!
Marieke E. Nieuwenhuijzen; A. R. Backus; A. Bahramisharif; Christian F. Doeller; Ole Jensen; Marcel A. J. Gerven
MEG-based decoding of the spatiotemporal dynamics of visual category perception Journal Article
In: NeuroImage, vol. 83, pp. 1063–1073, 2013.
Visual processing is a complex task which is best investigated using sensitive multivariate analysis methods that can capture representation-specific brain activity over both time and space. In this study, we applied a multivariate decoding algorithm to MEG data of subjects engaged in passive viewing of images of faces, scenes, bodies and tools. We used reconstructed source-space time courses as input to the algorithm in order to localize brain regions involved in optimal image discrimination. Applying this method to the interval of 115 to 315. ms after stimulus onset, we show a focal localization of regression coefficients in the inferior occipital, middle occipital, and lingual gyrus that drive decoding of the different perceived image categories. Classifier accuracy was highest (over 90% correctly classified trials, compared to a chance level accuracy of 50%) when dissociating the perception of faces from perception of other object categories. Furthermore, we applied this method to each single time point to extract the temporal evolution of visual perception. This allowed for the detection of differences in visual category perception as early as 85. ms after stimulus onset. Furthermore, localizing the corresponding regression coefficients of each time point allowed us to capture the spatiotemporal dynamics of visual category perception. This revealed initial involvement of sources in the inferior occipital, inferior temporal and superior occipital gyrus. During sustained stimulation additional sources in the anterior inferior temporal gyrus and superior parietal gyrus became involved. We conclude that decoding of source-space MEG data provides a suitable method to investigate the spatiotemporal dynamics of ongoing cognitive processing.
Emanuel N. Broeke; D. M. Hartgerink; J. Butler; Julien Lambert; André Mouraux
Central sensitization increases the pupil dilation elicited by mechanical pinprick stimulation Journal Article
In: Journal of Neurophysiology, vol. 53, no. 9, pp. 1689–1699, 2013.
High frequency electrical stimulation (HFS) of skin nociceptors triggers central sensitization (CS), manifested as increased pinprick sensitivity of the skin surrounding the site of HFS. Our aim was to assess the effect of CS on pinprick‐evoked pupil dilation responses (PDRs) and pinprick‐evoked brain potentials (PEPs). We hypothesized that the increase in the positive wave of PEPs following HFS would result from an enhanced pinprick‐evoked phasic response of the locus coeruleus‐noradrenergic system (LC‐NS), indicated by enhanced PDRs. In fourteen healthy volunteers, 64 and 96 mN pinprick stimuli were delivered to the left and right forearms, before and twenty minutes after applying HFS to one of the two forearms. Both PEPs and pinprick‐evoked PDRs were recorded. After HFS, pinprick stimuli were perceived as more intense at the HFS treated arm compared to baseline and control site, and this increase was similar for both stimulation intensities. Importantly, the pinprick‐ evoked PDR was also increased and the increase was stronger for 64 as compared to 96 mN stimulation. This is in line with our previous results showing a stronger increase of the PEP positivity at 64 vs. 96 mN stimulation and suggests that the increase in PEP positivity observed in previous studies could relate, at least in part, to enhance LC‐NS activity. However, there was no increase of the PEP positivity in the present study, indicating that enhanced LC‐NS activity is not the only determinant of the HFS‐induced enhancement of PEPs. Altogether, our results indicate that PDRs are more sensitive for detecting CS than PEPs.
Stan Van Pelt; Pascal Fries
Visual stimulus eccentricity affects human gamma peak frequency Journal Article
In: NeuroImage, vol. 78, pp. 439–447, 2013.
The peak frequency of neuronal gamma-band synchronization has received much attention in recent years. Gamma peak frequency shifts to higher frequency values for higher contrast, faster moving, and attended stimuli. In monkey V1, gamma peak frequency for a drifting grating is higher for a parafoveal as compared to an eccentric stimulus (Lima et al., 2010). This effect might be due to the cortical magnification factor: the higher cortical magnification for parafoveal stimuli increases the velocity with which the cortical representations of the moving grating stripes move across the cortical surface. Since faster moving stimuli lead to higher gamma frequency, a faster moving cortical representation might do the same. This explanation predicts that the eccentricity effect on gamma peak frequency is absent for stationary stimuli. To test this, we investigated the effect of eccentricity on gamma peak frequency by recording magnetoencephalography in human subjects while they viewed moving or stationary gratings. We found that both the moving and the stationary stimuli induced lower peak frequencies for larger eccentricities, arguing against an explanation based on the cortical magnification factor. We further investigated whether this eccentricity effect was explained by differences in the size or the spatial frequency of the expected cortical activation. Neither of those explained the eccentricity effect. We propose that the different stimulus and top-down factors leading to higher gamma peak frequency all result in higher stimulus salience, that salience is translated into gamma peak frequency, and that gamma peak frequency might subserve the preferential processing of neuronal activity induced by salient stimuli.
Younes Zerouali; Jean Marc Lina; Boutheina Jemel
Optimal eye-gaze fixation position for face-related neural responses Journal Article
In: PLoS ONE, vol. 8, no. 6, pp. e60128, 2013.
It is generally agreed that some features of a face, namely the eyes, are more salient than others as indexed by behavioral diagnosticity, gaze-fixation patterns and evoked-neural responses. However, because previous studies used unnatural stimuli, there is no evidence so far that the early encoding of a whole face in the human brain is based on the eyes or other facial features. To address this issue, scalp electroencephalogram (EEG) and eye gaze-fixations were recorded simultaneously in a gaze-contingent paradigm while observers viewed faces. We found that the N170 indexing the earliest face-sensitive response in the human brain was the largest when the fixation position is located around the nasion. Interestingly, for inverted faces, this optimal fixation position was more variable, but mainly clustered in the upper part of the visual field (around the mouth). These observations extend the findings of recent behavioral studies, suggesting that the early encoding of a face, as indexed by the N170, is not driven by the eyes per se, but rather arises from a general perceptual setting (upper-visual field advantage) coupled with the alignment of a face stimulus to a stored face template.
John M. Henderson; Steven G. Luke; Joseph Schmidt; John E. Richards
Co-registration of eye movements and event-related potentials in connected-text paragraph reading Journal Article
In: Frontiers in Systems Neuroscience, vol. 7, pp. 28, 2013.
Eyetracking during reading has provided a critical source of on-line behavioral data informing basic theory in language processing. Similarly, event-related potentials (ERPs) have provided an important on-line measure of the neural correlates of language processing. Recently there has been strong interest in co-registering eyetracking and ERPs from simultaneous recording to capitalize on the strengths of both techniques, but a challenge has been devising approaches for controlling artifacts produced by eye movements in the EEG waveform. In this paper we describe our approach to correcting for eye movements in EEG and demonstrate its applicability to reading. The method is based on independent components analysis, and uses three criteria for identifying components tied to saccades: (1) component loadings on the surface of the head are consistent with eye movements; (2) source analysis localizes component activity to the eyes, and (3) the temporal activation of the component occurred at the time of the eye movement and differed for right and left eye movements. We demonstrate this method's applicability to reading by comparing ERPs time-locked to fixation onset in two reading conditions. In the text-reading condition, participants read paragraphs of text. In the pseudo-reading control condition, participants moved their eyes through spatially similar pseudo-text that preserved word locations, word shapes, and paragraph spatial structure, but eliminated meaning. The corrected EEG, time-locked to fixation onsets, showed effects of reading condition in early ERP components. The results indicate that co-registration of eyetracking and EEG in connected-text paragraph reading is possible, and has the potential to become an important tool for investigating the cognitive and neural bases of on-line language processing in reading.
Florian Hutzler; Isabella Fuchs; Benjamin Gagl; Sarah Schuster; Fabio Richlan; Mario Braun; Stefan Hawelka
Parafoveal X-masks interfere with foveal word recognition: Evidence from fixation-related brain potentials Journal Article
In: Frontiers in Systems Neuroscience, vol. 7, pp. 33, 2013.
The boundary paradigm, in combination with parafoveal masks, is the main technique for studying parafoveal preprocessing during reading. The rationale is that the masks (e.g., strings of X's) prevent parafoveal preprocessing, but do not interfere with foveal processing. A recent study, however, raised doubts about the neutrality of parafoveal masks. In the present study, we explored this issue by means of fixation-related brain potentials (FRPs). Two FRP conditions presented rows of five words. The task of the participant was to judge whether the final word of a list was a "new" word, or whether it was a repeated (i.e., "old") word. The critical manipulation was that the final word was X-masked during parafoveal preview in one condition, whereas another condition presented a valid preview of the word. In two additional event-related brain potential (ERP) conditions, the words were presented serially with no parafoveal preview available; in one of the conditions with a fixed timing, in the other word presentation was self-paced by the participants. Expectedly, the valid-preview FRP condition elicited the shortest processing times. Processing times did not differ between the two ERP conditions indicating that "cognitive readiness" during self-paced processing can be ruled out as an alternative explanation for differences in processing times between the ERP and the FRP conditions. The longest processing times were found in the X-mask FRP condition indicating that parafoveal X-masks interfere with foveal word recognition.
Maciej Kosilo; Sophie M. Wuerger; Matt Craddock; Ben J. Jennings; Amelia R. Hunt; Jasna Martinovic
Low-level and high-level modulations of fixational saccades and high frequency oscillatory brain activity in a visual object classification task Journal Article
In: Frontiers in Psychology, vol. 4, pp. 948, 2013.
Until recently induced gamma-band activity (GBA) was considered a neural marker of cortical object representation. However, induced GBA in the electroencephalogram (EEG) is susceptible to artifacts caused by miniature fixational saccades. Recent studies have demonstrated that fixational saccades also reflect high-level representational processes. Do high-level as opposed to low-level factors influence fixational saccades? What is the effect of these factors on artifact-free GBA? To investigate this, we conducted separate eye tracking and EEG experiments using identical designs. Participants classified line drawings as objects or non-objects. To introduce low-level differences, contours were defined along different directions in cardinal color space: S-cone-isolating, intermediate isoluminant, or a full-color stimulus, the latter containing an additional achromatic component. Prior to the classification task, object discrimination thresholds were measured and stimuli were scaled to matching suprathreshold levels for each participant. In both experiments, behavioral performance was best for full-color stimuli and worst for S-cone isolating stimuli. Saccade rates 200-700 ms after stimulus onset were modulated independently by low and high-level factors, being higher for full-color stimuli than for S-cone isolating stimuli and higher for objects. Low-amplitude evoked GBA and total GBA were observed in very few conditions, showing that paradigms with isoluminant stimuli may not be ideal for eliciting such responses. We conclude that cortical loops involved in the processing of objects are preferentially excited by stimuli that contain achromatic information. Their activation can lead to relatively early exploratory eye movements even for foveally-presented stimuli.
Franziska Kretzschmar; Dominique Pleimling; Jana Hosemann; Stephan Füssel; Ina Bornkessel-Schlesewsky; Matthias Schlesewsky
Subjective impressions do not mirror online reading effort: Concurrent EEG-eyetracking evidence from the reading of books and digital media Journal Article
In: PLoS ONE, vol. 8, no. 2, pp. e56178, 2013.
In the rapidly changing circumstances of our increasingly digital world, reading is also becoming an increasingly digital experience: electronic books (e-books) are now outselling print books in the United States and the United Kingdom. Nevertheless, many readers still view e-books as less readable than print books. The present study thus used combined EEG and eyetracking measures in order to test whether reading from digital media requires higher cognitive effort than reading conventional books. Young and elderly adults read short texts on three different reading devices: a paper page, an e-reader and a tablet computer and answered comprehension questions about them while their eye movements and EEG were recorded. The results of a debriefing questionnaire replicated previous findings in that participants overwhelmingly chose the paper page over the two electronic devices as their preferred reading medium. Online measures, by contrast, showed shorter mean fixation durations and lower EEG theta band voltage density--known to covary with memory encoding and retrieval--for the older adults when reading from a tablet computer in comparison to the other two devices. Young adults showed comparable fixation durations and theta activity for all three devices. Comprehension accuracy did not differ across the three media for either group. We argue that these results can be explained in terms of the better text discriminability (higher contrast) produced by the backlit display of the tablet computer. Contrast sensitivity decreases with age and degraded contrast conditions lead to longer reading times, thus supporting the conclusion that older readers may benefit particularly from the enhanced contrast of the tablet. Our findings thus indicate that people's subjective evaluation of digital reading media must be dissociated from the cognitive and neural effort expended in online information processing while reading from such devices.
Junpeng Lao; Luca Vizioli; Roberto Caldara
Culture modulates the temporal dynamics of global/local processing Journal Article
In: Culture and Brain, vol. 1, no. 2-4, pp. 158–174, 2013.
Cultural differences in the way individuals from Western Caucasian (WC) and East Asian (EA) societies perceive and attend to visual information have been consistently reported in recent years. WC observers favor and perceive most efficiently the salient, local visual information by directing attention to focal objects. In contrast, EA observers show a bias towards global information, by preferentially attending elements in the background. However, the underlying neural mechanisms and the temporal dynamics of this striking cultural contrast have yet to be clarified. The combination of Navon figures, which contain both global and local features, and the measurement of neural adaptation constitute an ideal way to probe this issue. We recorded the electrophysiological signals of WC and EA observers while they actively matched culturally neutral geometric Navon shapes. In each trial, participants sequentially viewed and categorized an adapter shape followed by a target shape, as being either: identical; global congruent; local congruent; and different. We quantified the repetition suppression, a reduction in neural activity in stimulus sensitive regions following stimulus repetition, using a single-trial approach. A robust data-driven spatio-temporal analysis revealed at 80 ms a significant interaction between the culture of the observers and shape adaptation. EA observers showed sensitivity to global congruency on the attentional P1 component, whereas WC observers showed discrimination for global shapes at later stages. Our data revealed an early sensitivity to global and local shape cate- gorization, which is modulated by culture. This neural tuning could underlie more complex behavioral differences observed across human populations.
Adrian K. C. Lee; Siddharth Rajaram; Jing Xia; Hari Bharadwaj; Eric D. Larson; Matti S. Hämäläinen; Barbara G. Shinn-Cunningham
Auditory selective attention reveals preparatory activity in different cortical regions for selection based on source location and source pitch Journal Article
In: Frontiers in Neuroscience, vol. 6, pp. 190, 2013.
In order to extract information in a rich environment, we focus on different features that allow us to direct attention to whatever source is of interest. The cortical network deployed during spatial attention, especially in vision, is well characterized. For example, visuospatial attention engages a frontoparietal network including the frontal eye fields (FEFs), which modulate activity in visual sensory areas to enhance the representation of an attended visual object. However, relatively little is known about the neural circuitry controlling attention directed to non-spatial features, or to auditory objects or features (either spatial or non-spatial). Here, using combined magnetoencephalography (MEG) and anatomical information obtained from MRI, we contrasted cortical activity when observers attended to different auditory features given the same acoustic mixture of two simultaneous spoken digits. Leveraging the fine temporal resolution of MEG, we establish that activity in left FEF is enhanced both prior to and throughout the auditory stimulus when listeners direct auditory attention to target location compared to when they focus on target pitch. In contrast, activity in the left posterior superior temporal sulcus (STS), a region previously associated with auditory pitch categorization, is greater when listeners direct attention to target pitch rather than target location. This differential enhancement is only significant after observers are instructed which cue to attend, but before the acoustic stimuli begin. We therefore argue that left FEF participates more strongly in directing auditory spatial attention, while the left STS aids auditory object selection based on the non-spatial acoustic feature of pitch.
Juan E. Kamienkowski; Matias J. Ison; Rodrigo Quian Quiroga; Mariano Sigman
Fixation-related potentials in visual search: A combined EEG and eye tracking study Journal Article
In: Journal of Vision, vol. 12, no. 7, pp. 1–20, 2012.
We report a study of concurrent eye movements and electroencephalographic (EEG) recordings while subjects freely explored a search array looking for hidden targets. We describe a sequence of fixation-event related potentials (fERPs) that unfolds during ; 400 ms following each fixation. This sequence highly resembles the event-related responses in a replay experiment, in which subjects kept fixation while a sequence of images occurred around the fovea simulating the spatial and temporal patterns during the free viewing experiment. Similar responses were also observed in a second control experiment where the appearance of stimuli was controlled by the experimenters and presented at the center of the screen. We also observed a relatively early component (;150 ms) that distinguished between targets and distractors only in the freeviewing condition. We present a novel approach to match the critical properties of two conditions (targets/distractors), which can be readily adapted to other paradigms to investigate EEG components during free eye-movements.
Annmarie MacNamara; Joseph Schmidt; Gregory J. Zelinsky; Greg Hajcak
Electrocortical and ocular indices of attention to fearful and neutral faces presented under high and low working memory load Journal Article
In: Biological Psychology, vol. 91, no. 3, pp. 349–356, 2012.
Working memory load reduces the late positive potential (LPP), consistent with the notion that functional activation of the DLPFC attenuates neural indices of sustained attention. Visual attention also modulates the LPP. In the present study, we sought to determine whether working memory load might exert its influence on ERPs by reducing fixations to arousing picture regions. We simultaneously recorded eye-tracking and EEG while participants performed a working memory task interspersed with the presentation of task-irrelevant fearful and neutral faces. As expected, fearful compared to neutral faces elicited larger N170 and LPP amplitudes; in addition, working memory load reduced the N170 and the LPP. Participants made more fixations to arousing regions of neutral faces and faces presented under high working memory load. Therefore, working memory load did not induce avoidance of arousing picture regions and visual attention cannot explain load effects on the N170 and LPP.
Peter R. Murphy; Ian H. Robertson; Darren Allen; Robert Hester; Redmond G. O'Connell
An electrophysiological signal that precisely tracks the emergence of error awareness Journal Article
In: Frontiers in Human Neuroscience, vol. 6, pp. 65, 2012.
Recent electrophysiological research has sought to elucidate the neural mechanisms necessary for the conscious awareness of action errors. Much of this work has focussed on the error positivity (Pe), a neural signal that is specifically elicited by errors that have been consciously perceived. While awareness appears to be an essential prerequisite for eliciting the Pe, the precise functional role of this component has not been identified. Twenty-nine participants performed a novel variant of the Go/No-go Error Awareness Task (EAT) in which awareness of commission errors was indicated via a separate speeded manual response. Independent component analysis (ICA) was used to isolate the Pe from other stimulus- and response-evoked signals. Single-trial analysis revealed that Pe peak latency was highly correlated with the latency at which awareness was indicated. Furthermore, the Pe was more closely related to the timing of awareness than it was to the initial erroneous response. This finding was confirmed in a separate study which derived IC weights from a control condition in which no indication of awareness was required, thus ruling out motor confounds. A receiver-operating-characteristic (ROC) curve analysis showed that the Pe could reliably predict whether an error would be consciously perceived up to 400ms before the average awareness response. Finally, Pe latency and amplitude were found to be significantly correlated with overall error awareness levels between subjects. Our data show for the first time that the temporal dynamics of the Pe trace the emergence of error awareness. These findings have important implications for interpreting the results of clinical EEG studies of error processing.
Michael Plöchl; José P. Ossandón; Peter König
Combining EEG and eye tracking: Identification, characterization, and correction of eye movement artifacts in electroencephalographic data Journal Article
In: Frontiers in Human Neuroscience, vol. 6, pp. 278, 2012.
Eye movements introduce large artifacts to electroencephalographic recordings (EEG) and thus render data analysis difficult or even impossible. Trials contaminated by eye movement and blink artifacts have to be discarded, hence in standard EEG-paradigms subjects are required to fixate on the screen. To overcome this restriction, several correction methods including regression and blind source separation have been proposed. Yet, there is no automated standard procedure established. By simultaneously recording eye movements and 64-channel-EEG during a guided eye movement paradigm, we investigate and review the properties of eye movement artifacts, including corneo-retinal dipole changes, saccadic spike potentials and eyelid artifacts, and study their interrelations during different types of eye- and eyelid movements. In concordance with earlier studies our results confirm that these artifacts arise from different independent sources and that depending on electrode site, gaze direction, and choice of reference these sources contribute differently to the measured signal. We assess the respective implications for artifact correction methods and therefore compare the performance of two prominent approaches, namely linear regression and independent component analysis (ICA). We show and discuss that due to the independence of eye artifact sources, regression-based correction methods inevitably over- or under-correct individual artifact components, while ICA is in principle suited to address such mixtures of different types of artifacts. Finally, we propose an algorithm, which uses eye tracker information to objectively identify eye-artifact related ICA-components (ICs) in an automated manner. In the data presented here, the algorithm performed very similar to human experts when those were given both, the topographies of the ICs and their respective activations in a large amount of trials. Moreover it performed more reliable and almost twice as effective than human experts when those had to base their decision on IC topographies only. Furthermore, a receiver operating characteristic (ROC) analysis demonstrated an optimal balance of false positive and false negative at an area under curve (AUC) of more than 0.99. Removing the automatically detected ICs from the data resulted in removal or substantial suppression of ocular artifacts including microsaccadic spike potentials, while the relevant neural signal remained unaffected. In conclusion the present work aims at a better understanding of individual eye movement artifacts, their interrelations and the respective implications for eye artifact correction. Additionally, the proposed ICA-procedure provides a tool for optimized detection and correction of eye movement-related artifact components.
Federico Raimondo; Juan E. Kamienkowski; Mariano Sigman; Diego Fernandez Slezak
CUDAICA: GPU optimization of infomax-ICA EEG analysis Journal Article
In: Computational Intelligence and Neuroscience, vol. 2012, pp. 1–8, 2012.
In recent years, Independent Component Analysis (ICA) has become a standard to identify relevant dimensions of the data in neuroscience. ICA is a very reliable method to analyze data but it is, computationally, very costly. The use of ICA for online analysis of the data, used in brain computing interfaces, results are almost completely prohibitive. We show an increase with almost no cost (a rapid video card) of speed of ICA by about 25 fold. The EEG data, which is a repetition of many independent signals in multiple channels, is very suitable for processing using the vector processors included in the graphical units. We profiled the implementation of this algorithm and detected two main types of operations responsible of the processing bottleneck and taking almost 80% of computing time: vector-matrix and matrix-matrix multiplications. By replacing function calls to basic linear algebra functions to the standard CUBLAS routines provided by GPU manufacturers, it does not increase performance due to CUDA kernel launch overhead. Instead, we developed a GPU-based solution that, comparing with the original BLAS and CUBLAS versions, obtains a 25x increase of performance for the ICA calculation.
Robert M. G. Reinhart; Richard P. Heitz; Braden A. Purcell; Pauline K. Weigand; Jeffrey D. Schall; Geoffrey F. Woodman
Homologous mechanisms of visuospatial working memory maintenance in macaque and human: Properties and sources Journal Article
In: Journal of Neuroscience, vol. 32, no. 22, pp. 7711–7722, 2012.
Although areas of frontal cortex are thought to be critical for maintaining information in visuospatial working memory, the event-related potential (ERP) index of maintenance is found over posterior cortex in humans. In the present study, we reconcile these seemingly contradictory findings. Here, we show that macaque monkeys and humans exhibit the same posterior ERP signature of working memory maintenance that predicts the precision of the memory-based behavioral responses. In addition, we show that the specific pattern of rhythmic oscillations in the alpha band, recently demonstrated to underlie the human visual working memory ERP component, is also present in monkeys. Next, we concurrently recorded intracranial local field potentials from two prefrontal and another frontal cortical area to determine their contribution to the surface potential indexing maintenance. The local fields in the two prefrontal areas, but not the cortex immediately posterior, exhibited amplitude modulations, timing, and relationships to behavior indicating that they contribute to the generation of the surface ERP component measured from the distal posterior electrodes. Rhythmic neural activity in the theta and gamma bands during maintenance provided converging support for the engagement of the same brain regions. These findings demonstrate that nonhuman primates have homologous electrophysiological signatures of visuospatial working memory to those of humans and that a distributed neural network, including frontal areas, underlies the posterior ERP index of visuospatial working memory maintenance.
Marco Davare; A. Zénon; Gilles Pourtois; Michel Desmurget; Etienne Olivier
Role of the medial part of the intraparietal sulcus in implementing movement direction Journal Article
In: Cerebral Cortex, vol. 22, no. 6, pp. 1382–1394, 2012.
The contribution of the posterior parietal cortex (PPC) to visually guided movements has been originally inferred from observations made in patients suffering from optic ataxia. Subsequent electrophysiological studies in monkeys and functional imaging data in humans have corroborated the key role played by the PPC in sensorimotor transformations underlying goal-directed movements, although the exact contribution of this structure remains debated. Here, we used transcranial magnetic stimulation (TMS) to interfere transiently with the function of the left or right medial part of the intraparietal sulcus (mIPS) in healthy volunteers performing visually guided movements with the right hand. We found that a "virtual lesion" of either mIPS increased the scattering in initial movement direction (DIR), leading to longer trajectory and prolonged movement time, but only when TMS was delivered 100-160 ms before movement onset and for movements directed toward contralateral targets. Control experiments showed that deficits in DIR consequent to mIPS virtual lesions resulted from an inappropriate implementation of the motor command underlying the forthcoming movement and not from an inaccurate computation of the target localization. The present study indicates that mIPS plays a causal role in implementing specifically the direction vector of visually guided movements toward objects situated in the contralateral hemifield.
Marc R. Kamke; Michelle G. Hall; Harley F. Lye; Martin V. Sale; Laura R. Fenlon; Timothy J. Carroll; Stephan Riek; Jason B. Mattingley
Visual attentional load influences plasticity in the human motor cortex Journal Article
In: Journal of Neuroscience, vol. 32, no. 20, pp. 7001–7008, 2012.
Neural plasticity plays a critical role in learning, memory, and recovery from injury to the nervous system. Although much is known about the physical and physiological determinants of plasticity, little is known about the influence of cognitive factors. In this study, we investigated whether selective attention plays a role in modifying changes in neural excitability reflecting long-term potentiation (LTP)-like plasticity. We induced LTP-like effects in the hand area of the human motor cortex using transcranial magnetic stimulation (TMS). During the induction of plasticity, participants engaged in a visual detection task with either low or high attentional demands. Changes in neural excitability were assessed by measuring motor-evoked potentials in a small hand muscle before and after the TMS procedures. In separate experiments plasticity was induced either by paired associative stimulation (PAS) or intermittent theta-burst stimulation (iTBS). Because these procedures induce different forms of LTP-like effects, they allowed us to investigate the generality of any attentional influence on plasticity. In both experiments reliable changes in motor cortex excitability were evident under low-load conditions, but this effect was eliminated under high-attentional load. In a third experiment we investigated whether the attentional task was associated with ongoing changes in the excitability of motor cortex, but found no difference in evoked potentials across the levels of attentional load. Our findings indicate that in addition to their role in modifying sensory processing, mechanisms of attention can also be a potent modulator of cortical plasticity.
Marc R. Kamke; Harrison E. Vieth; David Cottrell; Jason B. Mattingley
Parietal disruption alters audiovisual binding in the sound-induced flash illusion Journal Article
In: NeuroImage, vol. 62, no. 3, pp. 1334–1341, 2012.
Selective attention and multisensory integration are fundamental to perception, but little is known about whether, or under what circumstances, these processes interact to shape conscious awareness. Here, we used transcranial magnetic stimulation (TMS) to investigate the causal role of attention-related brain networks in multisensory integration between visual and auditory stimuli in the sound-induced flash illusion. The flash illusion is a widely studied multisensory phenomenon in which a single flash of light is falsely perceived as multiple flashes in the presence of irrelevant sounds. We investigated the hypothesis that extrastriate regions involved in selective attention, specifically within the right parietal cortex, exert an influence on the multisensory integrative processes that cause the flash illusion. We found that disruption of the right angular gyrus, but not of the adjacent supramarginal gyrus or of a sensory control site, enhanced participants' veridical perception of the multisensory events, thereby reducing their susceptibility to the illusion. Our findings suggest that the same parietal networks that normally act to enhance perception of attended events also play a role in the binding of auditory and visual stimuli in the sound-induced flash illusion.
Ryota Kanai; Neil G. Muggleton; Vincent Walsh
Transcranial direct current stimulation of the frontal eye fields during pro- and antisaccade tasks Journal Article
In: Frontiers in Psychiatry, vol. 3, pp. 45, 2012.
Transcranial direct current stimulation (tDCS) has been successfully applied to cortical areas such as the motor cortex and visual cortex. In the present study, we examined whether tDCS can reach and selectively modulate the excitability of the frontal eye field (FEF). In order to assess potential effects of tDCS, we measured saccade latency, landing point, and its variability in a simple prosaccade task and in an antisaccade task. In the prosaccade task, we found that anodal tDCS shortened the latency of saccades to a contralateral visual cue. However, cathodal tDCS did not show a significant modulation of saccade latency. In the antisaccade task, on the other hand, we found that the latency for ipisilateral antisaccades was prolonged during the stimulation, whereas anodal stimulation did not modulate the latency of antisaccades. In addition, anodal tDCS reduced the erroneous saccades toward the contralateral visual cue. These results in the antisaccade task suggest that tDCS modulates the function of FEF to suppress reflexive saccades to the contralateral visual cue. Both in the prosaccade and antisaccade tasks, we did not find any effect of tDCS on saccade landing point or its variability. Our present study is the first to show effects of tDCS over FEF and opens the possibility of applying tDCS for studying the functions of FEF in oculomotor and attentional performance.
Kohitij Kar; Bart Krekelberg
Transcranial electrical stimulation over visual cortex evokes phosphenes with a retinal origin Journal Article
In: Journal of Neurophysiology, vol. 108, no. 8, pp. 2173–2178, 2012.
Transcranial electrical stimulation (tES) is a promising therapeutic tool for a range of neurological diseases. Understanding how the small currents used in tES spread across the scalp and penetrate the brain will be important for the rational design of tES therapies. Alternating currents applied transcranially above visual cortex induce the perception of flashes of light (phosphenes). This makes the visual system a useful model to study tES. One hypothesis is that tES generates phosphenes by direct stimulation of the cortex underneath the transcranial electrode. Here, we provide evidence for the alternative hypothesis that phosphenes are generated in the retina by current spread from the occipital electrode. Building on the existing literature, we first confirm that phosphenes are induced at lower currents when electrodes are placed farther away from visual cortex and closer to the eye. Second, we explain the temporal frequency tuning of phosphenes based on the well-known response properties of primate retinal ganglion cells. Third, we show that there is no difference in the time it takes to evoke phosphenes in the retina or by stimulation above visual cortex. Together, these findings suggest that phosphenes induced by tES over visual cortex originate in the retina. From this, we infer that tES currents spread well beyond the area of stimulation and are unlikely to lead to focal neural activation. Novel stimulation protocols that optimize current distributions are needed to overcome these limitations of tES.
Wei-Kuang Liang; Chi-Hung Juan
Modulation of motor control in saccadic behaviors by TMS over the posterior parietal cortex Journal Article
In: Journal of Neurophysiology, vol. 108, no. 3, pp. 741–752, 2012.
The right posterior parietal cortex (rPPC) has been found to be critical in shaping visual selection and distractor-induced saccade curvature in the context of predictive as well as nonpredictive visual cues by means of transcranial magnetic stimulation (TMS) interference. However, the dynamic details of how distractor-induced saccade curvatures are affected by rPPC TMS have not yet been investigated. This study aimed to elucidate the key dynamic properties that cause saccades to curve away from distractors with different degrees of curvature in various TMS and target predictability conditions. Stochastic optimal feedback control theory was used to model the dynamics of the TMS saccade data. This allowed estimation of torques, which was used to identify the critical dynamic mechanisms producing saccade curvature. The critical mechanisms of distractor-induced saccade curvatures were found to be the motor commands and torques in the transverse direction. When an unpredictable saccade target occurred with rPPC TMS, there was an initial period of greater distractor-induced torque toward the side opposite the distractor in the transverse direction, immediately followed by a relatively long period of recovery torque that brought the deviated trace back toward the target. The results imply that the mechanisms of distractor-induced saccade curvature may be comprised of two mechanisms: the first causing the initial deviation and the second bringing the deviated trace back toward the target. The pattern of the initial torque in the transverse direction revealed the former mechanism. Conversely, the later mechanism could be well explained as a consequence of the control policy in this model. To summarize, rPPC TMS increased the initial torque away from the distractor as well as the recovery torque toward the target.
José P. Ossandón; Selim Onat; Dario Cazzoli; Thomas Nyffeler; René M. Müri; Peter König
Unmasking the contribution of low-level features to the guidance of attention Journal Article
In: Neuropsychologia, vol. 50, no. 14, pp. 3478–3487, 2012.
The role of low-level stimulus-driven control in the guidance of overt visual attention has been difficult to establish because low- and high-level visual content are spatially correlated within natural visual stimuli. Here we show that impairment of parietal cortical areas, either permanently by a lesion or reversibly by repetitive transcranial magnetic stimulation (rTMS), leads to fixation of locations with higher values of low-level features as compared to control subjects or in a no-rTMS condition. Moreover, this unmasking of stimulus-driven control crucially depends on the intrahemispheric balance between top-down and bottom-up cortical areas. This result suggests that although in normal behavior high-level features might exert a strong influence, low-level features do contribute to guide visual selection during the exploration of complex natural stimuli.
Sangita Dandekar; Jian Ding; Claudio M. Privitera; Thom Carney; Stanley A. Klein
The fixation and saccade P3 Journal Article
In: PLoS ONE, vol. 7, no. 11, pp. e48761, 2012.
Although most instances of object recognition during natural viewing occur in the presence of saccades, the neural correlates of objection recognition have almost exclusively been examined during fixation. Recent studies have indicated that there are post-saccadic modulations of neural activity immediately following eye movement landing; however, whether post-saccadic modulations affect relatively late occurring cognitive components such as the P3 has not been explored. The P3 as conventionally measured at fixation is commonly used in brain computer interfaces, hence characterizing the post-saccadic P3 could aid in the development of improved brain computer interfaces that allow for eye movements. In this study, the P3 observed after saccadic landing was compared to the P3 measured at fixation. No significant differences in P3 start time, temporal persistence, or amplitude were found between fixation and saccade trials. Importantly, sensory neural responses canceled in the target minus distracter comparisons used to identify the P3. Our results indicate that relatively late occurring cognitive neural components such as the P3 are likely less sensitive to post saccadic modulations than sensory neural components and other neural activity occurring shortly after eye movement landing. Furthermore, due to the similarity of the fixation and saccade P3, we conclude that the P3 following saccadic landing could possibly be used as a viable signal in brain computer interfaces allowing for eye movements.
Steven L. Franconeri; Jason M. Scimeca; Jessica C. Roth; Sarah A. Helseth; Lauren E. Kahn
Flexible visual processing of spatial relationships Journal Article
In: Cognition, vol. 122, no. 2, pp. 210–227, 2012.
Visual processing breaks the world into parts and objects, allowing us not only to examine the pieces individually, but also to perceive the relationships among them. There is work exploring how we perceive spatial relationships within structures with existing representations, such as faces, common objects, or prototypical scenes. But strikingly, there is little work on the perceptual mechanisms that allow us to flexibly represent arbitrary spatial relationships, e.g., between objects in a novel room, or the elements within a map, graph or diagram. We describe two classes of mechanism that might allow such judgments. In the simultaneous class, both objects are selected concurrently. In contrast, we propose a sequential class, where objects are selected individually over time. We argue that this latter mechanism is more plausible even though it violates our intuitions. We demonstrate that shifts of selection do occur during spatial relationship judgments that feel simultaneous, by tracking selection with an electrophysiological correlate. We speculate that static structure across space may be encoded as a dynamic sequence across time. Flexible visual spatial relationship processing may serve as a case study of more general visual relation processing beyond space, to other dimensions such as size or numerosity.
Ryan J. Vaden; Nathan L. Hutcheson; Lesley A. McCollum; Jonathan Kentros; Kristina M. Visscher
Older adults, unlike younger adults, do not modulate alpha power to suppress irrelevant information Journal Article
In: NeuroImage, vol. 63, no. 3, pp. 1127–1133, 2012.
This study examines the neural mechanisms through which younger and older adults ignore irrelevant information, a process that is necessary to effectively encode new memories. Some age-related memory deficits have been linked to a diminished ability to dynamically gate sensory input, resulting in problems inhibiting the processing of distracting stimuli. Whereas oscillatory power in the alpha band (8-12. Hz) over visual cortical areas is thought to dynamically gate sensory input in younger adults, it is not known whether older adults use the same mechanism to gate out sensory input. Here we identified a task in which both older and younger adults could suppress the processing of irrelevant sensory stimuli, allowing us to use electroencephalography (EEG) to explore the neural activity associated with suppression of visual processing. As expected, we found that the younger adults' suppression of visual processing was correlated with robust modulation of alpha oscillatory power. However, older adults did not modulate alpha power to suppress processing of visual information. These results demonstrate that suppression of alpha power is not necessary to inhibit the processing of distracting stimuli in older adults, suggesting the existence of alternative strategies for suppressing irrelevant, potentially distracting information.
Yangqing Xu; Steven L. Franconeri
The head of the table: Marking the "front" of an object is tightly linked with selection Journal Article
In: Journal of Neuroscience, vol. 32, no. 4, pp. 1408–1412, 2012.
Objects in the world do not have a surface that can be objectively labeled the "front." We impose this designation on one surface of an object according to several cues, including which surface is associated with the most task-relevant information or the direction of motion of an object. However, when these cues are competing, weak, or absent, we can also flexibly assign one surface as the front. One possibility is that this assignment is guided by the location of the "spotlight" of selection, where the selected region becomes the front. Here we used an electrophysiological correlate to show a direct temporal link between object structure assignments and the spatial locus of selection. We found that when human participants viewed a shape whose front and back surfaces were ambiguous, seeing a given surface as front was associated with selectively attending to that location. In Experiment 1, this pattern occurred during directed rapid (every 1 s) switches in structural percepts. In Experiment 2, this pattern occurred during spontaneous reversals, from 900 ms before to 600 ms after the reported percept. These results suggest that the distribution of selective attention might guide the organization of object structure.
Genevieve Z. Steiner; Robert J. Barry
Pupillary responses and event-related potentials as indices of the orienting reflex Journal Article
In: Psychophysiology, vol. 48, no. 12, pp. 1648–1655, 2011.
This study examined skin conductance responses, the late positive complex of the event-related potential, and pupillary dilation responses as autonomic and central correlates of the orienting reflex (OR) in the context of indifferent and significant stimuli. In particular, we aimed to clarify the inconsistencies surrounding the pupillary dilation response as an OR index. An auditory dishabituation paradigm was employed, and physiological measures were recorded from 24 participants. Response decrement to a repeated stimulus, response recovery to a change stimulus, and subsequent dishabituation were assessed. Findings confirmed expectations that the skin conductance response and the late positive complex are indices of the OR. The pupillary dilation response, however, demonstrated an unexpected sensitivity to stimulus novelty only, while the prestimulus measure of tonic pupil diameter showed the significance effect that was expected of the phasic measure. Together, these findings argue against the suggestion that the pupillary dilation response is an OR index. The diverse results obtained from this experiment contribute to our understanding of the OR, and provide impetus for further research with a variety of paradigm manipulations.
Shlomit Yuval-Greenberg; Leon Y. Deouell
Scalp-recorded induced gamma-band responses to auditory stimulation and its correlations with saccadic muscle-activity Journal Article
In: Brain Topography, vol. 24, no. 1, pp. 30–39, 2011.
We previously showed that the transient broadband induced gamma-band response in EEG (iGBRtb) appearing around 200-300 ms following a visual stimulus reflects the contraction of extra-ocular muscles involved in the execution of saccades, rather than neural oscillations. Several previous studies reported induced gamma-band responses also following auditory stimulation. It is still an open question whether, similarly to visual paradigms, such auditory paradigms are also sensitive to the saccadic confound. In the current study we address this question using simultaneous eye-tracking and EEG recordings during an auditory oddball paradigm. Subjects were instructed to respond to a rare target defined by sound source location, while fixating on a central screen. Results show that, similar to what was found in visual paradigms, saccadic rate displayed typical temporal dynamics including a post-stimulus decrease followed by an increase. This increase was more moderate, had a longer latency, and was less consistent across subjects than was found in the visual case. Crucially, the temporal dynamics of the induced gamma response were similar to those of saccadic-rate modulation. This suggests that the auditory induced gamma-band responses recorded on the scalp may also be affected by saccadic muscle activity.
Chang-Mao Chao; Philip Tseng; Tzu-Yu Hsu; Jia-Han Su; Ovid J. L. Tzeng; Daisy L. Hung; Neil G. Muggleton; Chi-Hung Juan
Predictability of saccadic behaviors is modified by transcranial magnetic stimulation over human posterior parietal cortex Journal Article
In: Human Brain Mapping, vol. 32, no. 11, pp. 1961–1972, 2011.
Predictability in the visual environment provides a powerful cue for efficient processing of scenes and objects. Recently, studies have suggested that the directionality and magnitude of saccade curvature can be informative as to how the visual system processes predictive information. The pres-ent study investigated the role of the right posterior parietal cortex (rPPC) in shaping saccade curva-tures in the context of predictive and non-predictive visual cues. We used an orienting paradigm that incorporated manipulation of target location predictability and delivered transcranial magnetic stimulation (TMS) over rPPC. Participants were presented with either an informative or uninforma-tive cue to upcoming target locations. Our results showed that rPPC TMS generally increased sac-cade latency and saccade error rates. Intriguingly, rPPC TMS increased curvatures away from the distractor only when the target location was unpredictable and decreased saccadic errors towards the distractor. These effects on curvature and accuracy were not present when the target location was predictable. These results dissociate the strong contingency between saccade latency and saccade curvature and also indicate that rPPC plays an important role in allocating and suppressing attention to distractors when the target demands visual disambiguation. Furthermore, the present study sug-gests that, like the frontal eye fields, rPPC is critically involved in determining saccade curvature and the generation of saccadic behaviors under conditions of differing target predictability.
Zoï Kapoula; Qing Yang; Norman Sabbah; Marine Vernet
Different effects of double-pulse TMS of the posterior parietal cortex on reflexive and voluntary saccades Journal Article
In: Frontiers in Human Neuroscience, vol. 5, pp. 114, 2011.
Gap and overlap tasks are widely used to promote automatic versus controlled saccades. This study examines the hypothesis that the right posterior parietal cortex (PPC) is differently involved in the two tasks. Twelve healthy students participated in the experiment. We used double-pulse transcranial magnetic stimulation (dTMS) on the right PPC, the first pulse delivered at the target onset and the second 65 or 80 ms later. Each subject performed several blocks of gap or overlap task with or without dTMS. Eye movements were recorded with an Eyelink device. The results show an increase of latency of saccades after dTMS of the right PPC for both tasks but for different time windows (0-80 ms for the gap task, 0-65 ms for the overlap task). Moreover, for rightward saccades the coefficient of variation of latency increased in the gap task but decreased in the overlap task. Finally, in the gap task and for leftward saccades only, dTMS at 0-80 ms decreased the amplitude and the speed of saccades. Although the study is preliminary and needs further investigation in detail, the results support the hypothesis that the right PPC is involved differently in the initiation of the saccades for the two tasks: in the gap task the PPC controls saccade triggering while in the overlap task it could be a relay to the Frontal Eye Fields which is known to control voluntary saccades, e.g., memory-guided and perhaps the controlled saccades in the overlap task The results have theoretical and clinical significance as gap-overlap tasks are easy to perform even in advanced age and in patients with neurodegenerative diseases.
Neil G. Muggleton; Roger Kalla; Chi-Hung Juan; Vincent Walsh
Dissociating the contributions of human frontal eye fields and posterior parietal cortex to visual search Journal Article
In: Journal of Neurophysiology, vol. 105, no. 6, pp. 2891–2896, 2011.
Imaging, lesion, and transcranial magnetic stimulation (TMS) studies have implicated a number of regions of the brain in searching for a target defined by a combination of attributes. The necessity of both frontal eye fields (FEF) and posterior parietal cortex (PPC) in task performance has been shown by the application of TMS over these regions. The effects of stimulation over these two areas have, thus far, proved to be remarkably similar and the only dissociation reported being in the timing of their involvement. We tested the hypotheses that 1) FEF contributes to performance in terms of visual target detection (possibly by modulation of activity in extrastriate areas with respect to the target), and 2) PPC is involved in translation of visual information for action. We used a task where the presence (and location) of the target was indicated by an eye movement. Task disruption was seen with FEF TMS (with reduced accuracy on the task) but not with PPC stimulation. When a search task requiring a manual response was presented, disruption with PPC TMS was seen. These results show dissociation of FEF and PPC contributions to visual search performance and that PPC involvement seems to be dependent on the response required by the task, whereas this is not the case for FEF. This supports the idea of FEF involvement in visual processes in a manner that might not depend on the required response, whereas PPC seems to be involved when a manual motor response to a stimulus is required.
Marine Vernet; Qing Yang; Zoï Kapoula
Guiding binocular saccades during reading: A TMS study of the PPC Journal Article
In: Frontiers in Human Neuroscience, vol. 5, pp. 14, 2011.
Reading is an activity based on complex sequences of binocular saccades and fixations. During saccades, the eyes do not move together perfectly: saccades could end with a misalignment, compromising fused vision. During fixations, small disconjugate drift can partly reduce this misalignment. We hypothesized that maintaining eye alignment during reading involves active monitoring from posterior parietal cortex (PPC); this goes against traditional views considering only downstream binocular control. Nine young adults read a text; transcranial magnetic stimulation (TMS) was applied over the PPC every 5 ± 0.2 s. Eye movements were recorded binocularly with Eyelink II. Stimulation had three major effects: (1) disturbance of eye alignment during fixation; (2) increase of saccade disconjugacy leading to eye misalignment; (3) decrease of eye alignment reduction during fixation drift. The effects depend on the side; the right PPC was more involved in maintaining alignment over the motor sequence. Thus, the PPC is actively involved in the control of binocular eye alignment during reading, allowing clear vision. Cortical activation during reading is related to linguistic processes and motor control per se. The study might be of interest for the understanding of deficits of binocular coordination, encountered in several populations, e.g., in children with dyslexia.
Minnan Xu-Wilson; Jing Tian; Reza Shadmehr; David S. Zee
TMS perturbs saccade trajectories and unmasks an internal feedback controller for saccades Journal Article
In: Journal of Neuroscience, vol. 31, no. 32, pp. 11537–11546, 2011.
When we applied a single pulse of transcranial magnetic stimulation (TMS) to any part of the human head during a saccadic eye movement, the ongoing eye velocity was reduced as early as 45 ms after the TMS, and lasted ∼32 ms. The perturbation to the saccade trajectory was not due to a mechanical effect of the lid on the eye (e.g., from blinks). When the saccade involved coordinated movements of both the eyes and the lids, e.g., in vertical saccades, TMS produced a synchronized inhibition of the motor commands to both eye and lid muscles. The TMS-induced perturbation of the eye trajectory did not show habituation with repetition, and was present in both pro-saccades and anti-saccades. Despite the perturbation, the eye trajectory was corrected within the same saccade with compensatory motor commands that guided the eyes to the target. This within-saccade correction did not rely on visual input, suggesting that the brain monitored the oculomotor commands as the saccade unfolded, maintained a real-time estimate of the position of the eyes, and corrected for the perturbation. TMS disrupted saccades regardless of the location of the coil on the head, suggesting that the coil discharge engages a nonhabituating startle-like reflex system. This system affects ongoing motor commands upstream of the oculomotor neurons, possibly at the level of the superior colliculus or omnipause neurons. Therefore, a TMS pulse centrally perturbs saccadic motor commands, which are monitored possibly via efference copy and are corrected via internal feedback.
Qing Yang; Zoï Kapoula
Distinct control of initiation and metrics of memory-guided saccades and vergence by the FEF: A TMS study Journal Article
In: PLoS ONE, vol. 6, no. 5, pp. e20322, 2011.
BACKGROUND: The initiation of memory guided saccades is known to be controlled by the frontal eye field (FEF). Recent physiological studies showed the existence of an area close to FEF that controls also vergence initiation and execution. This study is to explore the effect of transcranial magnetic simulation (TMS) over FEF on the control of memory-guided saccade-vergence eye movements. METHODOLOGY/PRINCIPAL FINDINGS: Subjects had to make an eye movement in dark towards a target flashed 1 sec earlier (memory delay); the location of the target relative to fixation point was such as to require either a vergence along the median plane, or a saccade, or a saccade with vergence; trials were interleaved. Single pulse TMS was applied on the left or right FEF; it was delivered at 100 ms after the end of memory delay, i.e. extinction of fixation LED that was the "go" signal. Twelve healthy subjects participated in the study. TMS of left or right FEF prolonged the latency of all types of eye movements; the increase varied from 21 to 56 ms and was particularly strong for the divergence movements. This indicates that FEF is involved in the initiation of all types of memory guided movement in the 3D space. TMS of the FEF also altered the accuracy but only for leftward saccades combined with either convergence or divergence; intrasaccadic vergence also increased after TMS of the FEF. CONCLUSIONS/SIGNIFICANCE: The results suggest anisotropy in the quality of space memory and are discussed in the context of other known perceptual motor anisotropies.
Snigdha Banerjee; Adam C. Snyder; Sophie Molholm; John J. Foxe
Oscillatory alpha-band mechanisms and the deployment of spatial attention to anticipated auditory and visual target locations: Supramodal or sensory-specific control mechanisms? Journal Article
In: Journal of Neuroscience, vol. 31, no. 27, pp. 9923–9932, 2011.
Oscillatory alpha-band activity (8-15 Hz) over parieto-occipital cortex in humans plays an important role in suppression of processing for inputs at to-be-ignored regions of space, with increased alpha-band power observed over cortex contralateral to locations expected to contain distractors. It is unclear whether similar processes operate during deployment of spatial attention in other sensory modalities. Evidence from lesion patients suggests that parietal regions house supramodal representations of space. The parietal lobes are prominent generators of alpha oscillations, raising the possibility that alpha is a neural signature of supramodal spatial attention. Furthermore, when spatial attention is deployed within vision, processing of task-irrelevant auditory inputs at attended locations is also enhanced, pointing to automatic links between spatial deployments across senses. Here, we asked whether lateralized alpha-band activity is also evident in a purely auditory spatial-cueing task and whether it had the same underlying generator configuration as in a purely visuospatial task. If common to both sensory systems, this would provide strong support for "supramodal" attention theory. Alternately, alpha-band differences between auditory and visual tasks would support a sensory-specific account. Lateralized shifts in alpha-band activity were indeed observed during a purely auditory spatial task. Crucially, there were clear differences in scalp topographies of this alpha activity depending on the sensory system within which spatial attention was deployed. Findings suggest that parietally generated alpha-band mechanisms are central to attentional deployments across modalities but that they are invoked in a sensory-specific manner. The data support an "interactivity account," whereby a supramodal system interacts with sensory-specific control systems during deployment of spatial attention.
Jacob Jolij; H. Steven Scholte; Simon Gaal; Timothy L. Hodgson; Victor A. F. Lamme
Act quickly, decide later: Long-latency visual processing underlies perceptual decisions but not reflexive behavior Journal Article
In: Journal of Cognitive Neuroscience, vol. 23, no. 12, pp. 3734–3745, 2011.
Humans largely guide their behavior by their visual representation of the world. Recent studies have shown that visual information can trigger behavior within 150 msec, suggesting that visually guided responses to external events, in fact, precede conscious awareness of those events. However, is such a view correct? By using a texture discrimination task, we show that the brain relies on long-latency visual processing in order to guide perceptual decisions. Decreasing stimulus saliency leads to selective changes in long-latency visually evoked potential components reflecting scene segmentation. These latency changes are accompanied by almost equal changes in simple RTs and points of subjective simultaneity. Furthermore, we find a strong correlation between individual RTs and the latencies of scene segmentation related components in the visually evoked potentials, showing that the processes underlying these late brain potentials are critical in triggering a response. However, using the same texture stimuli in an antisaccade task, we found that reflexive, but erroneous, prosaccades, but not antisaccades, can be triggered by earlier visual processes. In other words: The brain can act quickly, but decides late. Differences between our study and earlier findings suggesting that action precedes conscious awareness can be explained by assuming that task demands determine whether a fast and unconscious, or a slower and conscious, representation is used to initiate a visually guided response.
Yosuke Kita; Atsuko Gunji; Yuki Inoue; Takaaki Goto; Kotoe Sakihara; Makiko Kaga; Masumi Inagaki; Toru Hosokawa
Self-face recognition in children with autism spectrum disorders: A near-infrared spectroscopy study Journal Article
In: Brain and Development, vol. 33, no. 6, pp. 494–503, 2011.
It is assumed that children with autism spectrum disorders (ASD) have specificities for self-face recognition, which is known to be a basic cognitive ability for social development. In the present study, we investigated neurological substrates and potentially influential factors for self-face recognition of ASD patients using near-infrared spectroscopy (NIRS). The subjects were 11 healthy adult men, 13 normally developing boys, and 10 boys with ASD. Their hemodynamic activities in the frontal area and their scanning strategies (eye-movement) were examined during self-face recognition. Other factors such as ASD severities and self-consciousness were also evaluated by parents and patients, respectively. Oxygenated hemoglobin levels were higher in the regions corresponding to the right inferior frontal gyrus than in those corresponding to the left inferior frontal gyrus. In two groups of children these activities reflected ASD severities, such that the more serious ASD characteristics corresponded with lower activity levels. Moreover, higher levels of public self-consciousness intensified the activities, which were not influenced by the scanning strategies. These findings suggest that dysfunction in the right inferior frontal gyrus areas responsible for self-face recognition is one of the crucial neural substrates underlying ASD characteristics, which could potentially be used to evaluate psychological aspects such as public self-consciousness.
Christian Kluge; Markus Bauer; Alexander P. Leff; Hans-Jochen Heinze; Raymond J. Dolan; Jon Driver; Alexander Paul
Plasticity of human auditory-evoked fields induced by shock conditioning and contingency reversal Journal Article
In: Proceedings of the National Academy of Sciences, vol. 108, no. 30, pp. 12545–12550, 2011.
We used magnetoencephalography (MEG) to assess plasticity of human auditory cortex induced by classical conditioning and contingency reversal. Participants listened to random sequences of high or low tones. A first baseline phase presented these without further associations. In phase 2, one of the frequencies (CS(+)) was paired with shock on half its occurrences, whereas the other frequency (CS(-)) was not. In phase 3, the contingency assigning CS(+) and CS(-) was reversed. Conditioned pupil dilation was observed in phase 2 but extinguished in phase 3. MEG revealed that, during phase-2 initial conditioning, the P1m, N1m, and P2m auditory components, measured from sensors over auditory temporal cortex, came to distinguish between CS(+) and CS(-). After contingency reversal in phase 3, the later P2m component rapidly reversed its selectivity (unlike the pupil response) but the earlier P1m did not, whereas N1m showed some new learning but not reversal. These results confirm plasticity of human auditory responses due to classical conditioning, but go further in revealing distinct constraints on different levels of the auditory hierarchy. The later P2m component can reverse affiliation immediately in accord with an updated expectancy after contingency reversal, whereas the earlier auditory components cannot. These findings indicate distinct cognitive and emotional influences on auditory processing.
Ali Mazaheri; Nicholas E. DiQuattro; Jesse Bengson; Joy J. Geng
Pre-stimulus activity predicts the winner of top-down vs. bottom-up attentional selection Journal Article
In: PLoS ONE, vol. 6, no. 2, pp. e16243, 2011.
Our ability to process visual information is fundamentally limited. This leads to competition between sensory information that is relevant for top-down goals and sensory information that is perceptually salient, but task-irrelevant. The aim of the present study was to identify, from EEG recordings, pre-stimulus and pre-saccadic neural activity that could predict whether top-down or bottom-up processes would win the competition for attention on a trial-by-trial basis. We employed a visual search paradigm in which a lateralized low contrast target appeared alone, or with a low (i.e., non-salient) or high contrast (i.e., salient) distractor. Trials with a salient distractor were of primary interest due to the strong competition between top-down knowledge and bottom-up attentional capture. Our results demonstrated that 1) in the 1-sec pre-stimulus interval, frontal alpha (8-12 Hz) activity was higher on trials where the salient distractor captured attention and the first saccade (bottom-up win); and 2) there was a transient pre-saccadic increase in posterior-parietal alpha (7-8 Hz) activity on trials where the first saccade went to the target (top-down win). We propose that the high frontal alpha reflects a disengagement of attentional control whereas the transient posterior alpha time-locked to the saccade indicates sensory inhibition of the salient distractor and suppression of bottom-up oculomotor capture.
Hironori Nakatani; Nicoletta Orlandi; Cees Van Leeuwen
Precisely timed oculomotor and parietal EEG activity in perceptual switching Journal Article
In: Cognitive Neurodynamics, vol. 5, no. 4, pp. 399–409, 2011.
Blinks and saccades cause transient interruptions of visual input. To investigate how such effects influence our perceptual state, we analyzed the time courses of blink and saccade rates in relation to perceptual switching in the Necker cube. Both time courses of blink and saccade rates showed peaks at different moments along the switching process. A peak in blinking rate appeared 1,000 ms prior to the switching responses. Blinks occurring around this peak were associated with subsequent switching to the preferred interpretation of the Necker cube. Saccade rates showed a peak 150 ms prior to the switching response. The direction of saccades around this peak was predictive of the perceived orientation of the Necker cube afterwards. Peak blinks were followed and peak saccades were preceded by transient parietal theta band activity indicating the changing of the perceptual interpretation. Precisely-timed blinks, therefore, can initiate perceptual switching, and precisely-timed saccades can facilitate an ongoing change of interpretation. textcopyright 2011 The Author(s).
Nicole Naue; Daniel Strüber; Ingo Fründ; Jeanette Schadow; Daniel Lenz; Stefan Rach; Ursula Körner; Christoph S. Herrmann
Gamma in motion: Pattern reversal elicits stronger gamma-band responses than motion Journal Article
In: NeuroImage, vol. 55, no. 2, pp. 808–817, 2011.
Previous studies showed higher gamma-band responses (GBRs, ≈ 40. Hz) of the electroencephalogram (EEG) for moving compared to stationary stimuli. However, it is unclear whether this modulation by motion reflects a special responsiveness of the GBR to the stimulus feature ''motion,'' or whether GBR enhancements of similar magnitude can be elicited also by a salient change within a static stimulus that does not include motion.Therefore, we measured the EEG of healthy subjects watching stationary square wave gratings of high contrast that either started to move or reversed their black and white pattern shortly after their onset. The strong contrast change of the pattern reversal represented a salient but motionless change within the grating that was compared to the onset of the stationary grating and the motion onset. Induced and evoked GBRs were analyzed for all three display conditions. In order to assess the influenceof fixational eye movements on the induced GBRs, we also examined the time courses of microsaccade rates during the three display conditions. Amplitudes of both evoked and induced GBRs were stronger for pattern reversal than for motion onset. There was no significant amplitude difference between the onsets of the stationary and moving gratings. However, mean frequencies of the induced GBR were $sim$10. Hz higher in response to the onsets of moving compared to stationary gratings. Furthermore, the modulations of the induced GBR did not parallel the modulations of microsaccade rate, indicating that our induced GBRs reflect neuronal processes. These results suggest that, within the gamma-band range, the encoding of moving gratings in early visual cortex is primarily based on an upward frequency shift, whereas contrast changes within static gratings are reflected by amplitude enhancement.
Andrey R. Nikolaev; Chie Nakatani; Gijs Plomp; Peter Jurica; Cees Leeuwen
Eye fixation-related potentials in free viewing identify encoding failures in change detection Journal Article
In: NeuroImage, vol. 56, no. 3, pp. 1598–1607, 2011.
We considered the hypothesis that spontaneous dissociation between the direction of attention and eye movement causes encoding failure in change detection. We tested this hypothesis by analyzing eye fixation-related potentials (EFRP) at the encoding stage of a change blindness task; when participants freely inspect a scene containing an unmarked target region, in which a change will occur in a subsequent presentation. We measured EFRP amplitude prior to the execution of a saccade, depending on its starting or landing position relative to the target region. For those landings inside the target region, we found a difference in EFRP between correct detection and failure. Overall, correspondence between EFRP amplitude and the size of the saccade predicted successful detection of change; lack of correspondence was followed by change blindness. By contrast, saccade sizes and fixation durations around the target region were unrelated to subsequent change detection. Since correspondence between EFRP and eye movement indicates that overt attention was given to the target region, we concluded that overt attention is needed for successful encoding and that dissociation between eye movement and attention leads to change blindness.
Sangita Dandekar; Claudio M. Privitera; Thom Carney; Stanley A. Klein
Neural saccadic response estimation during natural viewing Sangita Journal Article
In: Journal of Neurophysiology, vol. 107, no. 4, pp. 1776–1790, 2011.
Studying neural activity during natural viewing conditions is not often attempted. Isolating the neural response of a single saccade is necessary to study neural activity during natural viewing; however, the close temporal spacing of saccades that occurs during natural viewing makes it difficult to determine the response to a single saccade. Herein, a general linear model (GLM) approach is applied to estimate the EEG neural saccadic response for different segments of the saccadic main sequence separately. It is determined that, in visual search conditions, neural responses estimated by conventional event-related averaging are significantly and systematically distorted relative to GLM estimates due to the close temporal spacing of saccades during visual search. Before the GLM is applied, analyses are applied that demonstrate that saccades during visual search with intersaccadic spacings as low as 100-150 ms do not exhibit significant refractory effects. Therefore, saccades displaying different intersaccadic spacings during visual search can be modeled using the same regressor in a GLM. With the use of the GLM approach, neural responses were separately estimated for five different ranges of saccade amplitudes during visual search. Occipital responses time locked to the onsets of saccades during visual search were found to account for, on average, 79 percent of the variance of EEG activity in a window 90-200 ms after the onsets of saccades for all five saccade amplitude ranges that spanned a range of 0.2-6.0 degrees. A GLM approach was also used to examine the lateralized ocular artifacts associated with saccades. Possible extensions of the methods presented here to account for the superposition of microsaccades in event-related EEG studies conducted in nominal fixation conditions are discussed.
Sven-Thomas Graupner; Sebastian Pannasch; Boris M. Velichkovsky
Saccadic context indicates information processing within visual fixations: Evidence from event-related potentials and eye-movements analysis of the distractor effect Journal Article
In: International Journal of Psychophysiology, vol. 80, no. 1, pp. 54–62, 2011.
Attention, visual information processing, and oculomotor control are integrated functions of closely related brain mechanisms. Recently, it was shown that the processing of visual distractors appearing during a fixation is modulated by the amplitude of its preceding saccade (Pannasch & Velichkovsky, 2009). So far, this was demonstrated only at the behavioral level in terms of saccadic inhibition. The present study investigated distractor-related brain activity with cortical eye fixation-related potentials (EFRPs). Moreover, the following saccade was included as an additional classification criterion. Eye movements and EFRPs were recorded during free visual exploration of paintings. During some of the fixations, a visual distractor was shown as an annulus around the fixation position, 100. ms after the fixation onset. The saccadic context of a fixation was classified by its preceding and following saccade amplitudes with the cut-off criterion set to 4° of visual angle. The prolongation of fixation duration induced by distractors was largest for fixations preceded and followed by short saccades. EFRP data revealed a difference in distractor-related P2 amplitude between the saccadic context conditions, following the same trend as in eye movements. Furthermore, influences of the following saccade amplitude on the latency of the saccadic inhibition and on the N1 amplitude were found. The EFRP results cannot be explained by the influence of saccades per se since this bias was removed by subtracting the baseline from the distractor EFRP. Rather, the data suggest that saccadic context indicates differences in how information is processed within single visual fixations.
Visual target selection and motor planning define attentional enhancement at perceptual processing stages Journal Article
In: Frontiers in Human Neuroscience, vol. 4, pp. 14, 2010.
Extracting information from the visual field can be achieved by covertly orienting attention to different regions, or by making saccades to bring areas of interest onto the fovea. While much research has shown a link between covert attention and saccade preparation, the nature of that link remains a matter of dispute. Covert presaccadic orienting could result from target selection or from planning a motor act toward an object. We examined the contribution of visual target selection and motor preparation to attentional orienting in humans by dissociating these two habitually aligned processes with saccadic adaptation. Adaptation introduces a discrepancy between the visual target evoking a saccade and the motor metrics of that saccade, which, unbeknownst to the participant, brings the eyes to a different spatial location. We examined attentional orienting by recording event-related potentials (ERPs) to task-irrelevant visual probes flashed during saccade preparation at four equidistant locations including the visual target location and the upcoming motor endpoint. ERPs as early as 130-170 ms post-probe were modulated by attention at both the visual target and motor endpoint locations. These results indicate that both target selection and motor preparation determine the focus of spatial attention, resulting in enhanced processing of stimuli at early visual-perceptual stages.
Hans Peter Frey; Shane P. Kelly; Edmund C. Lalor; John J. Foxe
Early spatial attentional modulation of inputs to the fovea Journal Article
In: Journal of Neuroscience, vol. 30, no. 13, pp. 4547–4551, 2010.
Attending to a specific spatial location modulates responsivity of neurons with receptive fields processing that part of the environment. A major outstanding question is whether attentional modulation operates differently for the foveal (central) representation of the visual field than it does for the periphery. Indeed, recent animal electrophysiological recordings suggest that attention differentially affects spatial integration for central and peripheral receptive fields in primary visual cortex. In human electroencephalographic recordings, spatial attention to peripheral locations robustly modulates activity in early visual regions, but it has been claimed that this mechanism does not operate in foveal vision. Here, however, we show clear early attentional modulation of foveal stimulation with the same timing and cortical sources as seen for peripheral stimuli, demonstrating that attentional gain control operates similarly across the entire field of view. These results imply that covertly attending away from the center of gaze, which is a common paradigm in behavioral and electrophysiological studies of attention, results in a precisely timed push–pull mechanism. While the amplitude of the initial response to stimulation at attended peripheral locations is significantly increased beginning at 80 ms, the amplitude of the response to foveal stimulation begins to be attenuated.
Tjerk P. Gutteling; Helene M. Ettinger-Veenstra; J. Leon Kenemans; Sebastiaan F. W. Neggers
Lateralized frontal eye field activity precedes occipital activity shortly before saccades: Evidence for cortico-cortical feedback as a mechanism underlying covert attention shifts Journal Article
In: Journal of Cognitive Neuroscience, vol. 22, no. 9, pp. 1931–1943, 2010.
When an eye movement is prepared, attention is shifted toward the saccade end-goal. This coupling of eye movements and spatial attention is thought to be mediated by cortical connections between the FEFs and the visual cortex. Here, we present evidence for the existence of these connections. A visual discrimination task was performed while recording the EEG. Discrimination performance was significantly improved when the discrimination target and the saccade target matched. EEG results show that frontal activity precedes occipital activity contralateral to saccade direction when the saccade is prepared but not yet executed; these effects were absent in fixation conditions. This is consistent with the idea that the FEF exerts a direct modulatory influence on the visual cortex and enhances perception at the saccade end-goal.
Simon P. Kelly; John J. Foxe; Geoffrey Newman; Jay A. Edelman
Prepare for conflict: EEG correlates of the anticipation of target competition during overt and covert shifts of visual attention Journal Article
In: European Journal of Neuroscience, vol. 31, no. 9, pp. 1690–1700, 2010.
When preparing to make a saccadic eye movement in a cued direction, perception of stimuli at the target location is enhanced, just as it is when attention is covertly deployed there. Accordingly, the timing and anatomical sources of preparatory brain activity accompanying shifts of covert attention and saccade preparation tend to exhibit a large degree of overlap. However, there is evidence that preparatory processes are modulated by the foreknowledge of visual distractor competition during covert attention, and it is unknown whether eye movement preparation undergoes equivalent modulation. Here we examine preparatory processes in the electroencephalogram of human participants during four blocked versions of a spatial cueing task, requiring either covert detection or saccade execution, and either containing a distractor or not. As in previous work, a typical pattern of spatially selective occipital, parietal and frontal activity was seen in all task versions. However, whereas distractor presence called on an enhancement of spatially selective visual cortical modulation during covert attention, it instead called on increased activity over frontomedial oculomotor areas in the case of overt saccade preparation. We conclude that, although advance orienting signals may be similar in character during overt and covert conditions, the pattern by which these signals are modulated to ameliorate the behavioral costs of distractor competition is highly distinct, pointing to a degree of separability between the overt and covert systems.
Alon S. Keren; Shlomit Yuval-Greenberg; Leon Y. Deouell
Saccadic spike potentials in gamma-band EEG : Characterization , detection and suppression Journal Article
In: NeuroImage, vol. 49, no. 3, pp. 2248–2263, 2010.
Analysis of high-frequency (gamma-band) neural activity by means of non-invasive EEG is gaining increasing interest. However, we have recently shown that a saccade-related spike potential (SP) seriously confounds the analysis of EEG induced gamma-band responses (iGBR), as the SP eludes traditional EEG artifact rejection methods. Here we provide a comprehensive profile of the SP and evaluate methods for its detection and suppression, aiming to unveil true cerebral gamma-band activity. The SP appears consistently as a sharp biphasic deflection of about 22 ms starting at the saccade onset, with a frequency band of ∼ 20-90 Hz. On the average, larger saccades elicit higher SP amplitudes. The SP amplitude gradually changes from the extra-ocular channels towards posterior sites with the steepest gradients around the eyes, indicating its ocular source. Although the amplitude and the sign of the SP depend on the choice of reference channel, the potential gradients remain the same and non-zero for all references. The scalp topography is modulated almost exclusively by the direction of saccades, with steeper gradients ipsilateral to the saccade target. We discuss how the above characteristics impede attempts to remove these SPs from the EEG by common temporal filtering, choice of different references, or rejection of contaminated trials. We examine the extent to which SPs can be reliably detected without an eye tracker, assess the degree to which scalp current density derivation attenuates the effect of the SP, and propose a tailored ICA procedure for minimizing the effect of the SP.
Yosuke Kita; Atsuko Gunji; Kotoe Sakihara; Masumi Inagaki; Makiko Kaga; Eiji Nakagawa; Toru Hosokawa
Scanning strategies do not modulate face identification: Eye-tracking and near-infrared spectroscopy study Journal Article
In: PLoS ONE, vol. 5, no. 6, pp. e11050, 2010.
BACKGROUND: During face identification in humans, facial information is sampled (seeing) and handled (processing) in ways that are influenced by the kind of facial image type, such as a self-image or an image of another face. However, the relationship between seeing and information processing is seldom considered. In this study, we aimed to reveal this relationship using simultaneous eye-tracking measurements and near-infrared spectroscopy (NIRS) in face identification tasks. METHODOLOGY/PRINCIPAL FINDINGS: 22 healthy adult subjects (8 males and 14 females) were shown facial morphing movies in which an initial facial image gradually changed into another facial image (that is, the subject's own face was changed to a familiar face). The fixation patterns on facial features were recorded, along with changes in oxyhemoglobin (oxyHb) levels in the frontal lobe, while the subjects identified several faces. In the self-face condition (self-face as the initial image), hemodynamic activity around the right inferior frontal gyrus (IFG) was significantly greater than in the familiar-face condition. On the other hand, the scanning strategy was similar in almost all conditions with more fixations on the eyes and nose than on other areas. Fixation time on the eye area did not correlate with changes in oxyHb levels, and none of the scanning strategy indices could estimate the hemodynamic changes. CONCLUSIONS/SIGNIFICANCE: We conclude that hemodynamic activity, i.e., the means of processing facial information, is not always modulated by the face-scanning strategy, i.e., the way of seeing, and that the right IFG plays important roles in both self-other facial discrimination and self-evaluation.
Keith J. Yoder; Matthew K. Belmonte
Combining computer game-based behavioural experiments with high-density EEG and infrared gaze tracking Journal Article
In: Journal of Visualized Experiments, vol. 46, pp. 1–10, 2010.
Experimental paradigms are valuable insofar as the timing and other parameters of their stimuli are well specified and controlled, and insofar as they yield data relevant to the cognitive processing that occurs under ecologically valid conditions. These two goals often are at odds, since well controlled stimuli often are too repetitive to sustain subjects' motivation. Studies employing electroencephalography (EEG) are often especially sensitive to this dilemma between ecological validity and experimental control: attaining sufficient signal-to-noise in physiological averages demands large numbers of repeated trials within lengthy recording sessions, limiting the subject pool to individuals with the ability and patience to perform a set task over and over again. This constraint severely limits researchers' ability to investigate younger populations as well as clinical populations associated with heightened anxiety or attentional abnormalities. Even adult, non-clinical subjects may not be able to achieve their typical levels of performance or cognitive engagement: an unmotivated subject for whom an experimental task is little more than a chore is not the same, behaviourally, cognitively, or neurally, as a subject who is intrinsically motivated and engaged with the task. A growing body of literature demonstrates that embedding experiments within video games may provide a way between the horns of this dilemma between experimental control and ecological validity. The narrative of a game provides a more realistic context in which tasks occur, enhancing their ecological validity (Chaytor & Schmitter-Edgecombe, 2003). Moreover, this context provides motivation to complete tasks. In our game, subjects perform various missions to collect resources, fend off pirates, intercept communications or facilitate diplomatic relations. In so doing, they also perform an array of cognitive tasks, including a Posner attention-shifting paradigm (Posner, 1980), a go/no-go test of motor inhibition, a psychophysical motion coherence threshold task, the Embedded Figures Test (Witkin, 1950, 1954) and a theory-of-mind (Wimmer & Perner, 1983) task. The game software automatically registers game stimuli and subjects' actions and responses in a log file, and sends event codes to synchronise with physiological data recorders. Thus the game can be combined with physiological measures such as EEG or fMRI, and with moment-to-moment tracking of gaze. Gaze tracking can verify subjects' compliance with behavioural tasks (e.g. fixation) and overt attention to experimental stimuli, and also physiological arousal as reflected in pupil dilation (Bradley et al., 2008). At great enough sampling frequencies, gaze tracking may also help assess covert attention as reflected in microsaccades - eye movements that are too small to foveate a new object, but are as rapid in onset and have the same relationship between angular distance and peak velocity as do saccades that traverse greater distances. The distribution of directions of microsaccades correlates with the (otherwise) covert direction of attention (Hafed & Clark, 2002).
Ben M. Harvey; O. J. Braddick; A. Cowey
Similar effects of repetitive transcranial magnetic stimulation of MT+ and a dorsomedial extrastriate site including V3A on pattern detection and position discrimination of rotating and radial motion patterns Journal Article
In: Journal of Vision, vol. 10, no. 5, pp. 1–15, 2010.
Our recent psychophysical experiments have identified differences in the spatial summation characteristics of pattern detection and position discrimination tasks performed with rotating, expanding, and contracting stimuli. Areas MT and MST are well established to be involved in processing these stimuli. fMRI results have shown retinotopic activation of area V3A depending on the location of the center of radial motion in vision. This suggests the possibility that V3A may be involved in position discrimination tasks with these motion patterns. Here we use repetitive transcranial magnetic stimulation (rTMS) over MT+ and a dorsomedial extrastriate region including V3A to try to distinguish between TMS effects on pattern detection and position discrimination tasks. If V3A were involved in position discrimination, we would expect to see effects on position discrimination tasks, but not pattern detection tasks, with rTMS over this dorsomedial extrastriate region. In fact, we could not dissociate TMS effects on the two tasks, suggesting that they are performed by the same extrastriate area, in MT+.
Steven L. Prime; Michael Vesia; J. Douglas Crawford
TMS over human frontal eye fields disrupts trans-saccadic memory of multiple objects Journal Article
In: Cerebral Cortex, vol. 20, no. 4, pp. 759–772, 2010.
We recently showed that transcranial magnetic stimulation (TMS) over the right parietal eye fields disrupts memory of object features and locations across saccades. We applied TMS over the frontal eye fields (FEF) as subjects compared the feature details of visual targets presented either within a single eye fixation (Fixation Task) or across a saccade (Saccade Task). TMS pulses were randomly delivered at one of 3 time intervals around the time of the saccade, or at equivalent times in the Fixation Task. A No-TMS control confirmed that subjects could normally retain approximately 3 visual features. TMS in the Fixation Task had no effect compared with No-TMS, but differences among TMS times were found during right FEF stimulation. TMS over either the right or left FEF disrupted memory performance in the Saccade Task when stimulation coincided most closely with the saccade. The capacity to compare pre-and postsaccadic features was reduced to 1-2 objects, as expected if the spatial aspect of memory was disrupted. These findings suggest that the FEF plays a role in the spatial processing involved in trans-saccadic memory of visual features. We propose that this process employs saccade-related feedback signals similar to those observed in spatial updating.
Michael Vesia; Steven L. Prime; Xiaogang Yan; Lauren E. Sergio; J. Douglas Crawford
Specificity of human parietal saccade and reach regions during transcranial magnetic stimulation Journal Article
In: Journal of Neuroscience, vol. 30, no. 39, pp. 13053–13065, 2010.
Single-unit recordings in macaque monkeys have identified effector-specific regions in posterior parietal cortex (PPC), but functional neuroimaging in the human has yielded controversial results. Here we used on-line repetitive transcranial magnetic stimulation (rTMS) to determine saccade and reach specificity in human PPC. A short train of three TMS pulses (separated by an interval of 100 ms) was delivered to superior parieto-occipital cortex (SPOC), a region over the midposterior intraparietal sulcus (mIPS), and a site close to caudal IPS situated over the angular gyrus (AG) during a brief memory interval while subjects planned either a saccade or reach with the left or right hand. Behavioral measures then were compared to controls without rTMS. Stimulation of mIPS and AG produced similar patterns: increased end-point variability for reaches and decreased saccade accuracy for contralateral targets. In contrast, stimulation of SPOC deviated reach end points toward visual fixation and had no effect on saccades. Contralateral-limb specificity was highest for AG and lowest for SPOC. Visual feedback of the hand negated rTMS-induced disruptions of the reach plan for mIPS and AG, but not SPOC. These results suggest that human SPOC is specialized for encoding retinally peripheral reach goals, whereas more anterior-lateral regions (mIPS and AG) along the IPS possess overlapping maps for saccade and reach planning and are more closely involved in motor details (i.e., planning the reach vector for a specific hand). This work provides the first causal evidence for functional specificity of these parietal regions in healthy humans.
Michael L. Waterston; Christopher C. Pack
Improved discrimination of visual stimuli following repetitive transcranial magnetic stimulation Journal Article
In: PLoS ONE, vol. 5, no. 4, pp. e10354, 2010.
Repetitive transcranial magnetic stimulation (rTMS) at certain frequencies increases thresholds for motor-evoked potentials and phosphenes following stimulation of cortex. Consequently rTMS is often assumed to introduce a “virtual lesion” in stimulated brain regions, with correspondingly diminished behavioral performance. Here we investigated the effects of rTMS to visual cortex on subjects' ability to perform visual psychophysical tasks. Contrary to expectations of a visual deficit, we find that rTMS often improves the discrimination of visual features. For coarse orientation tasks, discrimination of a static stimulus improved consistently following theta-burst stimulation of the occipital lobe. Using a reaction-time task, we found that these improvements occurred throughout the visual field and lasted beyond one hour post-rTMS. Low-frequency (1 Hz) stimulation yielded similar improvements. In contrast, we did not find consistent effects of rTMS on performance in a fine orientation discrimination task. Overall our results suggest that rTMS generally improves or has no effect on visual acuity, with the nature of the effect depending on the type of stimulation and the task. We interpret our results in the context of an ideal-observer model of visual perception.
José P. Ossandón; Andrea Helo; Rodrigo Montefusco-Siegmund; Pedro E. Maldonado
Superposition model predicts EEG occipital activity during free viewing of natural scenes Journal Article
In: Journal of Neuroscience, vol. 30, no. 13, pp. 4787–4795, 2010.
Visual event-related potentials (ERPs) produced by a stimulus are thought to reflect either an increase of synchronized activity or a phase realignment of ongoing oscillatory activity, with both mechanisms sharing the assumption that ERPs are independent of the current state of the brain at the time of stimulation. In natural viewing, however, visual inputs occur one after another at specific subject-paced intervals through unconstrained eye movements. We conjecture that during natural viewing, ERPs generated after each fixation are better explained by a superposition of ongoing oscillatory activity related to the processing of previous fixations, with new activity elicited by the visual input at the current fixation. We examined the electroencephalography (EEG) signals that occur in humans at the onset of each visual fixation, both while subjects freely viewed natural scenes and while they viewed a black or gray background. We found that the fixation ERPs show visual components that are absent when subjects move their eyes on a homogeneous gray or black screen. Single-trial EEG signals that comprise the ERP are predicted more accurately by a model of superposition than by either phase resetting or the addition of evoked responses and stimulus-independent noise. The superposition of ongoing oscillatory activity and the visually evoked response results in a modification of the ongoing oscillation phase. The results presented suggest that the observed EEG signals reflect changes occurring in a common neuronal substrate rather than a simple summation at the scalp of signals from independent sources.
Pia Rämä; Thierry Baccino
Eye fixation-related potentials (EFRPs) during object identification Journal Article
In: Visual Neuroscience, vol. 27, no. 5-6, pp. 187–192, 2010.
Eye fixation-related potential (EFRP) measures electrical brain activity in response to eye fixations. The aim of the current study was to investigate whether the EFRPs vary during consecutive eye fixations while subjects were performing an object identification task. Eye fixations evoked P1 and N1 components at the occipital and parietal recording sites. The latency of P1 component increased during consecutive fixations. The amplitude of P1 increased and the amplitude of N1 decreased during consecutive fixations. The results indicate that EFRPs are modulated during consecutive fixations, suggesting that the current technique may provide a useful tool to study temporal dynamics of visual perception and processes underlying object identification.
Victor Sander; Brian Soper; Stefan Everling
Nonhuman primate event-related potentials associated with pro- and anti-saccades Journal Article
In: NeuroImage, vol. 49, no. 2, pp. 1650–1658, 2010.
Non-invasive event-related potential (ERP) recordings have become a popular technique to study neural activity associated with saccades in humans. To date, it is not known whether nonhuman primates exhibit similar saccade-related ERPs. Here, we recorded ERPs associated with the performance of randomly interleaved pro- and anti-saccades in macaque monkeys. Stimulus-aligned ERPs showed short-latency visual component with more negative P2 and N2 peak amplitudes on anti- than on pro-saccade trials. Saccade-aligned ERPs showed a larger presaccadic negativity on anti- than pro-saccade trials, and a presaccadic positivity on pro-saccade trials, which was attenuated or absent on anti-saccade trials. This was followed by sharp negative spike potential immediately prior to the movement. Overall, these findings demonstrate that macaque monkeys, like humans, exhibit task-related differences of visual ERPs associated with pro- and anti-saccades and furthermore share presaccadic positivity as well as a spike potential prior to these tasks. We suggest that the presaccadic positivity on pro-saccade trials is generated by a source in the contralateral frontal eye fields and that the more negative voltage on anti-saccade trials is the result of additional sources of opposite polarity in neighboring frontal areas.
Ming Qian; Mario Aguilar; Karen N. Zachery; Claudio M. Privitera; Stanley A. Klein; Thom Carney; Loren W. Nolte
Decision-level fusion of EEG and pupil features for single-trial visual detection analysis Journal Article
In: IEEE Transactions on Biomedical Engineering, vol. 56, no. 7, pp. 1929–1937, 2009.
Several recent studies have reported success in applying EEG-based signal analysis to achieve accurate single-trial classification of responses to visual target detection. Pupil responses are proposed as a complementary modality that can support improved accuracy of single-trial signal analysis. We develop a pupillary response feature-extraction and -selection procedure that helps to improve the classification performance of a system based only on EEG signal analysis. We apply a two-level linear classifier to obtain cognitive-task-related analysis of EEG and pupil responses. The classification results based on the two modalities are then fused at the decision level. Here, the goal is to support increased classification confidence through the inherent modality complementarities. The fusion results show significant improvement over classification performance based on a single modality.
Naoyuki Sato; Yoko Yamaguchi
A computational predictor of human episodic memory based on a theta phase precession network Journal Article
In: PLoS ONE, vol. 4, no. 10, pp. e7536, 2009.
In the rodent hippocampus, a phase precession phenomena of place cell firing with the local field potential (LFP) theta is called "theta phase precession" and is considered to contribute to memory formation with spike time dependent plasticity (STDP). On the other hand, in the primate hippocampus, the existence of theta phase precession is unclear. Our computational studies have demonstrated that theta phase precession dynamics could contribute to primate-hippocampal dependent memory formation, such as object-place association memory. In this paper, we evaluate human theta phase precession by using a theory-experiment combined analysis. Human memory recall of object-place associations was analyzed by an individual hippocampal network simulated by theta phase precession dynamics of human eye movement and EEG data during memory encoding. It was found that the computational recall of the resultant network is significantly correlated with human memory recall performance, while other computational predictors without theta phase precession are not significantly correlated with subsequent memory recall. Moreover the correlation is larger than the correlation between human recall and traditional experimental predictors. These results indicate that theta phase precession dynamics are necessary for the better prediction of human recall performance with eye movement and EEG data. In this analysis, theta phase precession dynamics appear useful for the extraction of memory-dependent components from the spatio-temporal pattern of eye movement and EEG data as an associative network. Theta phase precession may be a common neural dynamic between rodents and humans for the formation of environmental memories.
Rolf Verleger; Andreas Sprenger; Sina Gebauer; Michaela Fritzmannova; Monique Friedrich; Stefanie Kraft; Piotr Jaśkowski; Piotr Jas
On why left events are the right ones: Neural mechanisms underlying the left-hemifield advantage in rapid serial visual presentation Journal Article
In: Journal of Cognitive Neuroscience, vol. 21, no. 3, pp. 474–488, 2009.
When simultaneous series of stimuli are rapidly presented left and right, containing two target stimuli T1 and T2, T2 is much better identified when presented in the left than in the right hemifield. Here, this effect was replicated, even when shifts of gaze were controlled, and was only partially compensated when T1 side provided the cue where to expect T2. Electrophysiological measurement revealed earlier latencies of T1- and T2-evoked N2(pc) peaks at the right than at the left visual cortex, and larger right-hemisphere T2-evoked N2(pc) amplitudes when T2 closely followed T1. These findings suggest that the right hemisphere was better able to single out the targets in time. Further, sustained contralateral slow shifts remained active after T1 for longer time at the right than at the left visual cortex, and developed more consistently at the right visual cortex when expecting T2 on the contralateral side. These findings might reflect better capacity of right-hemisphere visual working memory. These findings about the neurophysiological underpinnings of the large right-hemisphere advantage in this complex visual task might help elucidating the mechanisms responsible for the severe disturbance of hemineglect following damage to the right hemisphere.
Franziska Kretzschmar; Ina Bornkessel-Schlesewsky; Matthias Schlesewsky
Parafoveal versus foveal N400s dissociate spreading activation from contextual fit Journal Article
In: NeuroReport, vol. 20, no. 18, pp. 1613–1618, 2009.
Using concurrent electroencephalogram and eye movement measures to track natural reading, this study shows that N400 effects reflecting predictability are dissociable from those owing to spreading activation. In comparing predicted sentence endings with related and unrelated unpredicted endings in antonym constructions ('the opposite of black is white/yellow/nice'), fixation-related potentials at the critical word revealed a predictability-based N400 effect (unpredicted vs. predicted words). By contrast, event-related potentials time locked to the last fixation before the critical word showed an N400 only for the nonrelated unpredicted condition (nice). This effect is attributed to a parafoveal mismatch between the critical word and preactivated lexical features (i.e. features of the predicted word and its associates). In addition to providing the first demonstration of a parafoveally induced N400 effect, our results support the view that the N400 is best viewed as a component family.
Roger Kalla; Neil G. Muggleton; Alan Cowey; Vincent Walsh
Human dorsolateral prefrontal cortex is involved in visual search for conjunctions but not features: A theta TMS study Journal Article
In: Cortex, vol. 45, no. 9, pp. 1085–1090, 2009.
Functional neuroimaging studies have shown that the detection of a target defined by more than one feature (for example, a conjunction of colour and orientation) amongst distractors is associated with the activation of a network of brain areas. Dorsolateral prefrontal cortex (DLPFC), along with areas such as the frontal eye fields (FEF) and posterior parietal cortex (PPC), is a component of this network. While transcranial magnetic stimulation (TMS) had shown that both FEF and PPC are necessary for, and not just correlated with, successful conjunction search, this is not the case for DLPFC. To test the hypothesis that this area is also necessary for efficient conjunction search, TMS was applied over DLPFC and the effects on conjunction and feature (in this case colour) search performance compared with those when TMS was delivered over area MT/V5 and a vertex control stimulation condition. DLPFC TMS impaired performance on the conjunction search task but was without effect on feature search, similar to findings when TMS is delivered over PPC or FEF. Vertex TMS had no effects whereas MT/V5 TMS significantly improved performance with a time course that may indicate that this was due to modulation of V4 activity. These findings illustrate that, like FEF and PPC, DLPFC is necessary for fully effective conjunction visual search performance.
Ulrich Nuding; Roger Kalla; Neil G. Muggleton; Ulrich Büttner; Vincent Walsh; Stefan Glasauer
TMS evidence for smooth pursuit gain control by the frontal eye fields Journal Article
In: Cerebral Cortex, vol. 19, no. 5, pp. 1144–1150, 2009.
Smooth pursuit eye movements are used to continuously track slowly moving visual objects. A peculiar property of the smooth pursuit system is the nonlinear increase in sensitivity to changes in target motion with increasing pursuit velocities. We investigated the role of the frontal eye fields (FEFs) in this dynamic gain control mechanism by application of transcranial magnetic stimulation. Subjects were required to pursue a slowly moving visual target whose motion consisted of 2 components: a constant velocity component at 4 different velocities (0, 8, 16, and 24 deg/s) and a superimposed high-frequency sinusoidal oscillation (4 Hz, +/-8 deg/s). Magnetic stimulation of the FEFs reduced not only the overall gain of the system, but also the efficacy of the dynamic gain control. We thus provide the first direct evidence that the FEF population is significantly involved in the nonlinear computation necessary for continuously adjusting the feedforward gain of the pursuit system. We discuss this with relation to current models of smooth pursuit.
Clive R. Rosenthal; Emma E. Roche-Kelly; Masud Husain; Christopher Kennard
Response-dependent contributions of human primary motor cortex and angular gyrus to manual and perceptual sequence learning Journal Article
In: Journal of Neuroscience, vol. 29, no. 48, pp. 15115–15125, 2009.
Motor sequence learning on the serial reaction time task involves the integration of response-, stimulus-, and effector-based information. Human primary motor cortex (M1) and the inferior parietal lobule (IPL) have been identified with supporting the learning of effector-dependent and -independent information, respectively. Current neurocognitive data are, however, exclusively based on learning complex sequence information via perceptual-motor responses. Here, we investigated the effects of continuous theta-burst transcranial magnetic stimulation (cTBS)-induced disruption of M1 and the angular gyrus (AG) of the IPL on learning a probabilistic sequence via sequential perceptual-motor responses (experiment 1) or covert orienting of visuospatial attention (experiment 2). Functional effects on manual sequence learning were evident during 75% of training trials in the cTBS M1 condition, whereas cTBS over the AG resulted in interference confined to a midpoint during the training phase. Posttraining direct (declarative) tests of sequence knowledge revealed that cTBS over M1 modulated the availability of newly acquired sequence knowledge, whereby sequence knowledge was implicit in the cTBS M1 condition but was available to conscious awareness in the cTBS AG and control conditions. In contrast, perceptual sequence learning was abolished in the perceptual cTBS AG condition, whereas learning was intact and available to conscious awareness in the cTBS M1 and control conditions. These results show that the right AG had a critical role in perceptual sequence learning, whereas M1 had a causal role in developing experience-dependent functional attributes relevant to conscious knowledge on manual but not perceptual sequence learning.
Helene M. Ettinger-Veenstra; W. Huijbers; Tjerk P. Gutteling; M. Vink; J. Leon Kenemans; Sebastiaan F. W. Neggers
fMRI-guided TMS on cortical eye fields: The frontal but not intraparietal eye fields regulate the coupling between visuospatial attention and eye movements Journal Article
In: Journal of Neurophysiology, vol. 102, no. 6, pp. 3469–3480, 2009.
It is well known that parts of a visual scene are prioritized for visual processing, depending on the current situation. How the CNS moves this focus of attention across the visual image is largely unknown, although there is substantial evidence that preparation of an action is a key factor. Our results support the view that direct corticocortical feedback connections from frontal oculomotor areas to the visual cortex are responsible for the coupling between eye movements and shifts of visuospatial attention. Functional magnetic resonance imaging (fMRI)-guided transcranial magnetic stimulation (TMS) was applied to the frontal eye fields (FEFs) and intraparietal sulcus (IPS). A single pulse was delivered 60, 30, or 0 ms before a discrimination target was presented at, or next to, the target of a saccade in preparation. Results showed that the known enhancement of discrimination performance specific to locations to which eye movements are being prepared was enhanced by early TMS on the FEF contralateral to eye movement direction, whereas TMS on the IPS resulted in a general performance increase. The current findings indicate that the FEF affects selective visual processing within the visual cortex itself through direct feedback projections.
Robin Walker; Puncharat Techawachirakul; Patrick Haggard
Frontal eye field stimulation modulates the balance of salience between target and distractors Journal Article
In: Brain Research, vol. 1270, pp. 54–63, 2009.
Natural scenes generally include several possible objects that can be the target for a shift of gaze and attention. The oculomotor system may select a single target by boosting neural activation representing the target, and also by inhibiting neural activity associated with competing alternatives (distractors). We examine the role of the frontal eye field (FEF) in these processes through the effects of single-pulse transcranial magnetic stimulation (TMS) on the distractor-related modulation of saccade trajectories. Participants made voluntary saccades to peripheral locations specified by a central arrow-cue. On some trials, visual distractors appeared remote from the target location. The competing distractor produced a deviation of saccade trajectory, away from the distractor location. Single-pulse TMS stimulation of the right frontal eye field increased this distractor-related deviation compared that observed when stimulation was applied to a control site (vertex). The increase in distractor-related deviation of trajectory, following FEF stimulation, was observed for saccades made in both the left and right visual fields and could not be attributed to an effect of TMS on saccade latency. The enhanced distractor-related deviation following FEF stimulation could reflect increased inhibition of the competing distractor, or reduced salience of the endogenous saccade goal. The results are interpreted in light of neurophysiological evidence that the human FEF is involved in the dynamic interaction between competing stimuli for the selection of a candidate target.
Roger Kalla; Neil G. Muggleton; Chi-Hung Juan; Alan Cowey; Vincent Walsh
The timing of the involvement of the frontal eye fields and posterior parietal cortex in visual search Journal Article
In: NeuroReport, vol. 19, no. 10, pp. 1069–1073, 2008.
The frontal eye fields (FEFs) and posterior parietal cortex (PPC) are important for target detection in conjunction visual search but the relative timings of their contribution have not been compared directly. We addressed this using temporally specific double pulse transcranial magnetic stimulation delivered at different times over FEFs and PPC during performance of a visual search task. Disruption of performance was earlier (0/40 ms) with FEF stimulation than with PPC stimulation (120/160 ms), revealing a clear and substantial temporal dissociation of the involvement of these two areas in conjunction visual search. We discuss these timings with reference to the respective roles of FEF and PPC in the modulation of extrastriate visual areas and selection of responses.
Thomas Nyffeler; Dario Cazzoli; Pascal Wurtz; Mathias Lüthi; Roman Von Wartburg; Silvia Chaves; Anouk Déruaz; Christian W. Hess; René M. Müri
Neglect-like visual exploration behaviour after theta burst transcranial magnetic stimulation of the right posterior parietal cortex Journal Article
In: European Journal of Neuroscience, vol. 27, no. 7, pp. 1809–1813, 2008.
The right posterior parietal cortex (PPC) is critically involved in visual exploration behaviour, and damage to this area may lead to neglect of the left hemispace. We investigated whether neglect-like visual exploration behaviour could be induced in healthy subjects using theta burst repetitive transcranial magnetic stimulation (rTMS). To this end, one continuous train of theta burst rTMS was applied over the right PPC in 12 healthy subjects prior to a visual exploration task where colour photographs of real-life scenes were presented on a computer screen. In a control experiment, stimulation was also applied over the vertex. Eye movements were measured, and the distribution of visual fixations in the left and right halves of the screen was analysed. In comparison to the performance of 28 control subjects without stimulation, theta burst rTMS over the right PPC, but not the vertex, significantly decreased cumulative fixation duration in the left screen-half and significantly increased cumulative fixation duration in the right screen-half for a time period of 30 min. These results suggest that theta burst rTMS is a reliable method of inducing transient neglect-like visual exploration behaviour.
Marine Vernet; Qing Yang; Gintautas Daunys; Christophe Orssaud; Thomas Eggert; Zoï Kapoula
How the brain obeys Hering's law: A TMS study of the posterior parietal cortex Journal Article
In: Investigative Ophthalmology & Visual Science, vol. 49, no. 1, pp. 230–237, 2008.
PURPOSE: Human ocular saccades are not perfectly yoked; the origin of this disconjugacy (muscular versus central) remains controversial. The purpose of this study was to test a cortical influence on the binocular coordination of saccades. METHODS: The authors used a gap paradigm to elicit vertical or horizontal saccades of 10 degrees , randomly interleaved; transcranial magnetic stimulation (TMS) was applied on the posterior parietal cortex (PPC) 100 ms after the target onset. RESULTS: TMS of the left or right PPC increased (i) the misalignment of the eyes during the presaccadic fixation period; (ii) the size difference between the saccades of the eyes, called disconjugacy; the increase of disconjugacy was significant for rightward and downward saccades after TMS of the right PPC and for downward saccades after TMS of the left PPC. CONCLUSIONS: The authors conclude that the PPC is actively involved in maintaining eye alignment during fixation and in the control of binocular coordination of saccades.
Marine Vernet; Qing Yang; Gintautas Daunys; Christophe Orssaud; Zoï Kapoula
TMS of the posterior parietal cortex delays the latency of unpredictable saccades but not when they are combined with predictable divergence Journal Article
In: Brain Research Bulletin, vol. 76, no. 1-2, pp. 50–56, 2008.
This study tests the influence of transcranial magnetic stimulation (TMS) of the posterior parietal cortex (PPC) on the initiation of horizontal and vertical saccades, alone or combined with a predictable divergence. A gap paradigm was used; TMS was applied 100 ms after target onset. TMS of the left PPC increased the latency of unpredictable rightward saccades, while TMS of the right PPC increased the latency of unpredictable downward saccades. Yet, when unpredictable saccades were combined with predictable divergence, neither component was affected. We suggest that in the latter case, the initiation of both components was taken in charge by another area, e.g. frontal. Thus, even when one component was predictable, a common mechanism controls the initiation of both components. The results confirm that TMS only modifies the latency when the cortical area stimulated is involved in the triggering of the eye movement.
Shlomit Yuval-Greenberg; Orr Tomer; Alon S. Keren; Israel Nelken; Leon Y. Deouell
Transient induced gamma-band response in EEG as a manifestation of miniature saccades Journal Article
In: Neuron, vol. 58, no. 3, pp. 429–441, 2008.
The induced gamma-band EEG response (iGBR) recorded on the scalp is widely assumed to reflect synchronous neural oscillation associated with object representation, attention, memory, and consciousness. The most commonly reported EEG iGBR is a broadband transient increase in power at the gamma range ∼200-300 ms following stimulus onset. A conspicuous feature of this iGBR is the trial-to-trial poststimulus latency variability, which has been insufficiently addressed. Here, we show, using single-trial analysis of concomitant EEG and eye tracking, that this iGBR is tightly time locked to the onset of involuntary miniature eye movements and reflects a saccadic "spike potential." The time course of the iGBR is related to an increase in the rate of saccades following a period of poststimulus saccadic inhibition. Thus, whereas neuronal gamma-band oscillations were shown conclusively with other methods, the broadband transient iGBR recorded by scalp EEG reflects properties of miniature saccade dynamics rather than neuronal oscillations.
Steffen Gais; Sabine Köster; Andreas Sprenger; Judith Bethke; Wolfgang Heide; Hubert Kimmig
Sleep is required for improving reaction times after training on a procedural visuo-motor task Journal Article
In: Neurobiology of Learning and Memory, vol. 90, no. 4, pp. 610–615, 2008.
Sleep has been found to enhance consolidation of many different forms of memory. However in most procedural tasks, a sleep-independent, fast learning component interacts with slow, sleep-dependent improvements. Here, we show that in humans a visuo-motor saccade learning task shows no improvements during training, but only during a delayed recall testing after a period of sleep. Subjects were trained in a prosaccade task (saccade to a visual target). Performance was tested in the prosaccade and the antisaccade task (saccade to opposite direction of the target) before training, after a night of sleep or sleep deprivation, after a night of recovery sleep, and finally in a follow-up test 4 weeks later. We found no immediate improvement in saccadic reaction time (SRT) during training, but a delayed reduction in SRT, indicating a slow-learning process. This reduction occurred only after a period of sleep, i.e. after the first night in the sleep group and after recovery sleep in the sleep deprivation group. This improvement was stable during the 4-week follow-up. Saccadic training can thus induce covert changes in the saccade generation pathway. During the following sleep period, these changes in turn bring about overt performance improvements, presuming a learning effect based on synaptic tagging.
Michael Dambacher; Reinhold Kliegl
Synchronizing timelines: Relations between fixation durations and N400 amplitudes during sentence reading Journal Article
In: Brain Research, vol. 1155, no. 1, pp. 147–162, 2007.
We examined relations between eye movements (single-fixation durations) and RSVP-based event-related potentials (ERPs; N400s) recorded during reading the same sentences in two independent experiments. Longer fixation durations correlated with larger N400 amplitudes. Word frequency and predictability of the fixated word as well as the predictability of the upcoming word accounted for this covariance in a path-analytic model. Moreover, larger N400 amplitudes entailed longer fixation durations on the next word, a relation accounted for by word frequency. This pattern offers a neurophysiological correlate for the lag-word frequency effect on fixation durations: word processing is reliably expressed not only in fixation durations on currently fixated words, but also in those on subsequently fixated words.
Sven-Thomas Graupner; Boris M. Velichkovsky; Sebastian Pannasch; Johannes Marx
Surprise, surprise: Two distinct components in the visually evoked distractor effect Journal Article
In: Psychophysiology, vol. 44, no. 2, pp. 251–261, 2007.
The distractor effect is an inhibition of saccades shortly after a sudden visual event. It has been explained both as an oculomotor reflex and as a manifestation of the orienting response. To clarify which explanation is more appropriate, we investigated a possible habituation of this effect. Visual and auditory distractors were presented at gaze-contingent intervals during the perception of meaningful pictures. Both reflexlike and modifiable components were present in the visual distractor effect, with latencies of about 110 and 180 ms, respectively. The influence of visual and auditory distractors on saccades preceded the earliest changes in cortical ERPs. Only for long-term habituation in the visual modality was a correlation with ERPs (N1) found.
Samuel B. Hutton; Brendan S. Weekes
Low frequency rTMS over posterior parietal cortex impairs smooth pursuit eye tracking Journal Article
In: Experimental Brain Research, vol. 183, no. 2, pp. 195–200, 2007.
The role of the posterior parietal cortex in smooth pursuit eye movements remains unclear. We used low frequency repetitive transcranial magnetic stimulation (rTMS) to study the cognitive and neural systems involved in the control of smooth pursuit eye movements. Eighteen participants were tested on two separate occasions. On each occasion we measured smooth pursuit eye tracking before and after 6 min of 1 Hz rTMS delivered at 90% of motor threshold. Low frequency rTMS over the posterior parietal cortex led to a significant reduction in smooth pursuit velocity gain, whereas rTMS over the motor cortex had no effect on gain. We conclude that low frequency offline rTMS is a potentially useful tool with which to explore the cortical systems involved in oculomotor control.
Sebastiaan F. W. Neggers; W. Huijbers; C. M. Vrijlandt; Björn N. S. Vlaskamp; D. J. L. G. Schutter; J. Leon Kenemans
TMS pulses on the frontal eye fields break coupling between visuospatial attention and eye movements Journal Article
In: Journal of Neurophysiology, vol. 98, no. 5, pp. 2765–2778, 2007.
While preparing a saccadic eye movement, visual processing of the saccade goal is prioritized. Here, we provide evidence that the frontal eye fields (FEFs) are responsible for this coupling between eye movements and shifts of visuospatial attention. Functional magnetic resonance imaging (fMRI)-guided transcranial magnetic stimulation (TMS) was applied to the FEFs 30 ms before a discrimination target was presented at or next to the target of a saccade in preparation. Results showed that the well-known enhancement of discrimination performance on locations to which eye movements are being prepared was diminished by TMS contralateral to eye movement direction. Based on the present and other reports, we propose that saccade preparatory processes in the FEF affect selective visual processing within the visual cortex through feedback projections, in that way coupling saccade preparation and visuospatial attention.
A. J. Wills; Aureliu Lavric; G. S. Croft; Timothy L. Hodgson
Predictive learning, prediction errors, and attention: evidence from event-related potentials and eye tracking Journal Article
In: Journal of Cognitive Neuroscience, vol. 19, no. 5, pp. 843–854, 2007.
Prediction error ("surprise") affects the rate of learning: We learn more rapidly about cues for which we initially make incorrect predictions than cues for which our initial predictions are correct. The current studies employ electrophysiological measures to reveal early attentional differentiation of events that differ in their previous involvement in errors of predictive judgment. Error-related events attract more attention, as evidenced by features of event-related scalp potentials previously implicated in selective visual attention (selection negativity, augmented anterior N1). The earliest differences detected occurred around 120 msec after stimulus onset, and distributed source localization (LORETA) indicated that the inferior temporal regions were one source of the earliest differences. In addition, stimuli associated with the production of prediction errors show higher dwell times in an eye-tracking procedure. Our data support the view that early attentional processes play a role in human associative learning.
Glenn F. Wilson; John A. Caldwell; Christopher A. Russell
Performance and psychophysiological measures of fatigue effects on aviation related tasks of varying difficulty Journal Article
In: International Journal of Aviation Psychology, vol. 17, no. 2, pp. 219–247, 2007.
Fatigue is a well known stressor in aviation operations and its interaction with mental workload needs to be understood. Performance, psychophysiological, and subjective measures were collected during performance of three tasks of increasing complexity. A psychomotor vigilance task, multi-attribute task battery and an uninhabited air vehicle task were performed five times during one night's sleep loss. EEG, ECG and pupil area were recorded during task performance. Performance decrements were found at the next to last and/or last testing session. The EEG showed concomitant changes. The degree of impairment was at least partially dependent on the task being performed and the performance variable assessed.
Naoyuki Sato; Yoko Yamaguchi
Theta synchronization networks emerge during human object-place memory encoding Journal Article
In: NeuroReport, vol. 18, no. 5, pp. 419–424, 2007.
Recent rodent hippocampus studies have suggested that theta rhythm-dependent neural dynamics ('theta phase precession') is essential for an on-line memory formation. A computational study indicated that the phase precession enables a human object-place association memory with voluntary eye movements, although it is still an open question whether the human brain uses the dynamics. Here we elucidated subsequent memory-correlated activities in human scalp electroencephalography in an object-place association memory designed according the former computational study. Our results successfully demonstrated that subsequent memory recall is characterized by an increase in theta power and coherence, and further, that multiple theta synchronization networks emerge. These findings suggest the human theta dynamics in common with rodents in episodic memory formation.
Transcranial magnetic stimulation of frontal oculomotor regions during smooth pursuit Journal Article
In: Journal of Neuroscience, vol. 26, no. 2, pp. 458–466, 2006.
Both the frontal eye fields (FEFs) and supplementary eye fields (SEFs) are known to be involved in smooth pursuit eye movements. It has been shown recently that stimulation of the smooth-pursuit area of the FEF [frontal pursuit area (FPA)] in monkey increases the pursuit response to unexpected changes in target motion during pursuit. In the current study, we applied transcranial magnetic stimulation (TMS) to the FPA and SEF in humans during sinusoidal pursuit to assess its effects on the pursuit response to predictable, rather than unexpected, changes in target motion. For the FPA, we found that TMS applied immediately before the target reversed direction increased eye velocity in the new direction, whereas TMS applied in mid-cycle, immediately before the target began to slow, decreased eye velocity. For the SEF, TMS applied at target reversal increased eye velocity in the new direction but had no effect on eye velocity when applied at mid-cycle. TMS of the control region (leg region of the somatosensory cortex) did not affect eye velocity at either point. Previous stimulation studies of FPA during pursuit have suggested that this region is involved in controlling the gain of the transformation of visual signals into pursuit motor commands. The current results suggest that the gain of the transformation of predictive signals into motor commands is also controlled by the FPA. The effect of stimulation of the SEF is distinct from that of the FPA and suggests that its role in sinusoidal pursuit is primarily at the target direction reversal.
Christian Bellebaum; Klaus-Peter Hoffmann; Irene Daum
Post-saccadic updating of visual space in the posterior parietal cortex in humans Journal Article
In: Behavioural Brain Research, vol. 163, no. 2, pp. 194–203, 2005.
Updating of visual space takes place in the posterior parietal cortex to guarantee spatial constancy across eye movements. However, the timing of updating with respect to saccadic eye movements remains a matter of debate. In the present study, event-related potentials (ERPs) were recorded in 15 volunteers during a saccadic double-step task to elucidate the time course of the updating process. In the experimental condition updating of visual space was required, because both saccade targets had already disappeared before the first saccade was executed. A similar task without updating requirements served as control condition. ERP analysis revealed a significantly larger slow positive wave in the retino-spatial dissonance condition compared to the control condition, starting between 150 and 200 ms after first saccade onset. Source analysis showed an asymmetry with respect to the direction of the first saccade. Whereas the source was restricted to the right PPC in trials with leftward first saccades, left and right PPC were involved in rightward trials. The results of the present study suggest that updating of visual space in a saccadic double-step task occurs not earlier than 150 ms after the onset of the first saccade. We conclude that extraretinal information about the first saccade is integrated with motor information about the second saccade in the inter-saccade interval.