EyeLink EEG / fNIRS / TMS Publications
All EyeLink EEG, fNIRS, and TMS research publications (with concurrent eye tracking) up until 2024 (with early 2025s) 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!
2015 |
Nicholas E. Myers; Lena Walther; George Wallis; Mark G. Stokes; Anna C. Nobre In: Journal of Cognitive Neuroscience, vol. 27, no. 3, pp. 492–508, 2015. @article{Myers2015a,Working memory (WM) is strongly influenced by attention. In visual WM tasks, recall performance can be improved by an attention-guiding cue presented before encoding (precue) or during maintenance (retrocue). Although precues and retro- cues recruit a similar frontoparietal control network, the two are likely to exhibit some processing differences, because pre- cues invite anticipation of upcoming information whereas retro- cues may guide prioritization, protection, and selection of information already in mind. Here we explored the behavioral and electrophysiological differences between precueing and retrocueing in a new visual WM task designed to permit a direct comparison between cueing conditions. We found marked differences in ERP profiles between the precue and retrocue conditions. In line with precues primarily generating an anti- cipatory shift of attention toward the location of an upcoming item, we found a robust lateralization in late cue-evoked po- tentials associated with target anticipation. Retrocues elicited a different pattern of ERPs that was compatible with an early selec- tion mechanism, but not with stimulus anticipation. In contrast to the distinct ERP patterns, alpha-band (8–14 Hz) lateralization was indistinguishable between cue types (reflecting, in both conditions, the location of the cued item). We speculate that, whereas alpha-band lateralization after a precue is likely to enable anticipatory attention, lateralization after a retrocue may instead enable the controlled spatiotopic access to recently encoded visual information |
Karly N. Neath; Roxane J. Itier Fixation to features and neural processing of facial expressions in a gender discrimination task Journal Article In: Brain and Cognition, vol. 99, pp. 97–111, 2015. @article{Neath2015,Early face encoding, as reflected by the N170 ERP component, is sensitive to fixation to the eyes. Whether this sensitivity varies with facial expressions of emotion and can also be seen on other ERP components such as P1 and EPN, was investigated. Using eye-tracking to manipulate fixation on facial features, we found the N170 to be the only eye-sensitive component and this was true for fearful, happy and neutral faces. A different effect of fixation to features was seen for the earlier P1 that likely reflected general sensitivity to face position. An early effect of emotion (~120. ms) for happy faces was seen at occipital sites and was sustained until ~350. ms post-stimulus. For fearful faces, an early effect was seen around 80. ms followed by a later effect appearing at ~150. ms until ~300. ms at lateral posterior sites. Results suggests that in this emotion-irrelevant gender discrimination task, processing of fearful and happy expressions occurred early and largely independently of the eye-sensitivity indexed by the N170. Processing of the two emotions involved different underlying brain networks active at different times. |
Yuka O. Okazaki; Jörn M. Horschig; Lisa Luther; Robert Oostenveld; Ikuya Murakami; Ole Jensen Real-time MEG neurofeedback training of posterior alpha activity modulates subsequent visual detection performance Journal Article In: NeuroImage, vol. 107, pp. 323–332, 2015. @article{Okazaki2015,It has been demonstrated that alpha activity is lateralized when attention is directed to the left or right visual hemifield. We investigated whether real-time neurofeedback training of the alpha lateralization enhances participants' ability to modulate posterior alpha lateralization and causes subsequent short-term changes in visual detection performance. The experiment consisted of three phases: (i) pre-training assessment, (ii) neurofeedback phase and (iii) post-training assessment. In the pre- and post-training phases we measured the threshold to covertly detect a cued faint Gabor stimulus presented in the left or right hemifield. During magnetoencephalography (MEG) neurofeedback, two face stimuli superimposed with noise were presented bilaterally. Participants were cued to attend to one of the hemifields. The transparency of the superimposed noise and thus the visibility of the stimuli were varied according to the momentary degree of hemispheric alpha lateralization. In a double-blind procedure half of the participants were provided with sham feedback. We found that hemispheric alpha lateralization increased with the neurofeedback training; this was mainly driven by an ipsilateral alpha increase. Surprisingly, comparing pre- to post-training, detection performance decreased for a Gabor stimulus presented in the hemifield that was un-attended during neurofeedback. This effect was not observed in the sham group. Thus, neurofeedback training alters alpha lateralization, which in turn decreases performances in the untrained hemifield. Our findings suggest that alpha oscillations play a causal role for the allocation of attention. Furthermore, our neurofeedback protocol serves to reduce the detection of unattended visual information and could therefore be of potential use for training to reduce distractibility in attention deficit patients, but also highlights that neurofeedback paradigms can have negative impact on behavioral performance and should be applied with caution. |
Gonçalo Padrão; Borja Rodriguez-Herreros; Laura Pérez Zapata; Antoni Rodriguez-Fornells Exogenous capture of medial-frontal oscillatory mechanisms by unattended conflicting information Journal Article In: Neuropsychologia, vol. 75, pp. 458–468, 2015. @article{Padrao2015,A long-standing debate in psychology and cognitive neuroscience concerns the way in which unattended information is processed and influences goal-directed behavior. Although selective attention allows us to filter out task-irrelevant information, there is a substantial number of unattended, yet relevant, events that must be evaluated in a flexible manner so that appropriate behaviors can succeed. Here we inspected the extent to which unattended conflicting visual information, which cannot be consciously identified, influences behavior and activates medial prefrontal cortex (mPFC) mechanisms of action-monitoring and regulation, traditionally associated with conscious control processes.To that end, we performed two experiments using a novel variant of the Eriksen flanker task in which spatial attention was manipulated, preventing the conscious identification of unattended visual events. The first behavioral experiment was conducted to validate the efficacy of the novel paradigm. In the second experiment, we evaluated electrophysiological correlates of mPFC activity (a frontocentral negative ERP component and medial-frontal theta oscillations) in response to attended and unattended conflicting events. The results of both experiments demonstrated that attended and unattended conflicting stimuli altered subjects' behavior in a similar fashion, i.e. slowing down their reaction times and increasing their error rates. Importantly, the results of the EEG experiment showed that unattended conflicting stimuli, similarly to attended conflicting stimuli, led to an increase in theta-related frontocentral ERP activity and medial-frontal theta power, irrespective of the degree of conscious representation of the sources of conflict. This study provides evidence that medial-frontal theta oscillations represent a neural mechanism through which the mPFC may suppress and regulate potentially inappropriate actions that are automatically triggered by conflicting environmental stimuli to which we are oblivious. |
David R. Painter; Paul E. Dux; Jason B. Mattingley Causal involvement of visual area MT in global feature-based enhancement but not contingent attentional capture Journal Article In: NeuroImage, vol. 118, pp. 90–102, 2015. @article{Painter2015,When visual attention is set for a particular target feature, such as color or shape, neural responses to that feature are enhanced across the visual field. This global feature-based enhancement is hypothesized to underlie the contingent attentional capture effect, in which task-irrelevant items with the target feature capture spatial attention. In humans, however, different cortical regions have been implicated in global feature-based enhancement and contingent capture. Here, we applied intermittent theta-burst stimulation (iTBS) to assess the causal roles of two regions of extrastriate cortex - right area MT and the right temporoparietal junction (TPJ) - in both global feature-based enhancement and contingent capture. We recorded cortical activity using EEG while participants monitored centrally for targets defined by color and ignored peripheral checkerboards that matched the distractor or target color. In central vision, targets were preceded by colored cues designed to capture attention. Stimuli flickered at unique frequencies, evoking distinct cortical oscillations. Analyses of these oscillations and behavioral performance revealed contingent capture in central vision and global feature-based enhancement in the periphery. Stimulation of right area MT selectively increased global feature-based enhancement, but did not influence contingent attentional capture. By contrast, stimulation of the right TPJ left both processes unaffected. Our results reveal a causal role for the right area MT in feature-based attention, and suggest that global feature-based enhancement does not underlie the contingent capture effect. |
2014 |
Dan Nemrodov; Thomas Anderson; Frank F. Preston; Roxane J. Itier Early sensitivity for eyes within faces: A new neuronal account of holistic and featural processing Journal Article In: NeuroImage, vol. 97, pp. 81–94, 2014. @article{Nemrodov2014,Eyes are central to face processing however their role in early face encoding as reflected by the N170 ERP component is unclear. Using eye tracking to enforce fixation on specific facial features, we found that the N170 was larger for fixation on the eyes compared to fixation on the forehead, nasion, nose or mouth, which all yielded similar amplitudes. This eye sensitivity was seen in both upright and inverted faces and was lost in eyeless faces, demonstrating it was due to the presence of eyes at fovea. Upright eyeless faces elicited largest N170 at nose fixation. Importantly, the N170 face inversion effect (FIE) was strongly attenuated in eyeless faces when fixation was on the eyes but was less attenuated for nose fixation and was normal when fixation was on the mouth. These results suggest the impact of eye removal on the N170 FIE is a function of the angular distance between the fixated feature and the eye location. We propose the Lateral Inhibition, Face Template and Eye Detector based (LIFTED) model which accounts for all the present N170 results including the FIE and its interaction with eye removal. Although eyes elicit the largest N170 response, reflecting the activity of an eye detector, the processing of upright faces is holistic and entails an inhibitory mechanism from neurons coding parafoveal information onto neurons coding foveal information. The LIFTED model provides a neuronal account of holistic and featural processing involved in upright and inverted faces and offers precise predictions for further testing. |
Haijing Niu; Hao Li; Li Sun; Yongming Su; Jing Huang; Yan Song Visual learning alters the spontaneous activity of the resting human brain: An fNIRS study Journal Article In: BioMed Research International, pp. 1–9, 2014. @article{Niu2014,Resting-state functional connectivity (RSFC) has been widely used to investigate spontaneous brain activity that exhibits correlated fluctuations. RSFC has been found to be changed along the developmental course and after learning. Here, we investigated whether and how visual learning modified the resting oxygenated hemoglobin (HbO) functional brain connectivity by using functional near-infrared spectroscopy (fNIRS). We demonstrate that after five days of training on an orientation discrimination task constrained to the right visual field, resting HbO functional connectivity and directed mutual interaction between high-level visual cortex and frontal/central areas involved in the top-down control were significantly modified. Moreover, these changes, which correlated with the degree of perceptual learning, were not limited to the trained left visual cortex. We conclude that the resting oxygenated hemoglobin functional connectivity could be used as a predictor of visual learning, supporting the involvement of high-level visual cortex and the involvement of frontal/central cortex during visual perceptual learning. |
Cheolsoo Park; Markus Plank; Joseph Snider; Sanggyun Kim; He Crane Huang; Sergei Gepshtein; Todd P. Coleman; Howard Poizner In: IEEE Transactions on Neural Systems and Rehabilitation Engineering, vol. 22, no. 5, pp. 1083–1096, 2014. @article{Park2014,The neural dynamics underlying the coordination of spatially-directed limb and eye movements in humans is not well understood. Part of the difficulty has been a lack of signal processing tools suitable for the analysis of non-stationary electroencephalographic (EEG) signals. Here we use multivariate empirical mode decomposition (MEMD), a data-driven approach that does not employ predefined basis functions. High-density EEG, and arm and eye movements were synchronously recorded in 10 subjects performing time-constrained reaching and/or eye movements. Subjects were allowed to move both the hand and the eyes, only the hand, or only the eyes following a 500-700 ms delay interval where the hand and gaze remained on a central fixation cross. An additional condition involved a non-spatially-directed "lift" movement of the hand. The neural activity during a 500 ms delay interval was decomposed into intrinsic mode functions (IMFs) usingMEMD. Classification analysis revealed that gamma band (30 Hz <) IMFs produced more classifiable features differentiating the EEG according to the different upcoming movements. A benchmark test using conventional algorithms demonstrated that MEMD was the best algorithm for extracting oscillatory bands from EEG, yielding the best classification of the different movement conditions. The gamma rhythm decomposed using MEMD showed a higher correlation with the eventual movement accuracy than any other band rhythm and than any other algorithm. |
Hyeong Dong Park; Stéphanie Correia; Antoine Ducorps; Catherine Tallon-Baudry Spontaneous fluctuations in neural responses to heartbeats predict visual detection Journal Article In: Nature Neuroscience, vol. 17, no. 4, pp. 612–618, 2014. @article{Park2014a,Spontaneous fluctuations of ongoing neural activity substantially affect sensory and cognitive performance. Because bodily signals are constantly relayed up to the neocortex, neural responses to bodily signals are likely to shape ongoing activity. Here, using magnetoencephalography, we show that in humans, neural events locked to heartbeats before stimulus onset predict the detection of a faint visual grating in the posterior right inferior parietal lobule and the ventral anterior cingulate cortex, two regions that have multiple functional correlates and that belong to the same resting-state network. Neither fluctuations in measured bodily parameters nor overall cortical excitability could account for this finding. Neural events locked to heartbeats therefore shape visual conscious experience, potentially by contributing to the neural maps of the organism that might underlie subjectivity. Beyond conscious vision, our results show that neural events locked to a basic physiological input such as heartbeats underlie behaviorally relevant differential activation in multifunctional cortical areas. |
Rafael Polanía; Ian Krajbich; Marcus Grueschow; Christian C. Ruff Neural oscillations and synchronization differentially support evidence accumulation in perceptual and value-based decision making Journal Article In: Neuron, vol. 82, no. 3, pp. 709–720, 2014. @article{Polania2014,Organisms make two types of decisions on a regular basis. Perceptual decisions are determined by objective states of the world (e.g., melons are bigger than apples), whereas value-based decisions are determined by subjective preferences (e.g., I prefer apples to melons). Theoretical accounts suggest that both types of choice involve neural computations accumulating evidence for the choice alternatives; however, little is known about the overlap or differences in the processes underlying perceptual versus value-based decisions. We analyzed EEG recordings during a paradigm where perceptual- and value-based choices were based on identical stimuli. For both types of choice, evidence accumulation was evident in parietal gamma-frequency oscillations, whereas a similar frontal signal was unique for value-based decisions. Fronto-parietal synchronization of these signals predicted value-based choice accuracy. These findings uncover how decisions emerge from topographic- and frequency-specificoscillations that accumulate distinct aspects of evidence, with large-scale synchronization as a mechanism integrating these spatially distributed signals. |
Liina Pylkkänen; Douglas K. Bemis; Estibaliz Blanco Elorrieta Building phrases in language production: An MEG study of simple composition Journal Article In: Cognition, vol. 133, no. 2, pp. 371–384, 2014. @article{Pylkkaenen2014,Although research on language production has developed detailed maps of the brain basis of single word production in both time and space, little is known about the spatiotemporal dynamics of the processes that combine individual words into larger representations during production. Studying composition in production is challenging due to difficulties both in controlling produced utterances and in measuring the associated brain responses. Here, we circumvent both problems using a minimal composition paradigm combined with the high temporal resolution of magnetoencephalography (MEG). With MEG, we measured the planning stages of simple adjective-noun phrases ('red tree'), matched list controls ('red, blue'), and individual nouns ('tree') and adjectives ('red'), with results indicating combinatorial processing in the ventro-medial prefrontal cortex (vmPFC) and left anterior temporal lobe (LATL), two regions previously implicated for the comprehension of similar phrases. These effects began relatively quickly (~180 ms) after the presentation of a production prompt, suggesting that combination commences with initial lexical access. Further, while in comprehension, vmPFC effects have followed LATL effects, in this production paradigm vmPFC effects occurred mostly in parallel with LATL effects, suggesting that a late process in comprehension is an early process in production. Thus, our results provide a novel neural bridge between psycholinguistic models of comprehension and production that posit functionally similar combinatorial mechanisms operating in reversed order. |
Lily Riggs; Takako Fujioka; Jessica Chan; Douglas A. McQuiggan; Adam K. Anderson; Jennifer D. Ryan Association with emotional information alters subsequent processing of neutral faces Journal Article In: Frontiers in Human Neuroscience, vol. 8, pp. 1001, 2014. @article{Riggs2014,The processing of emotional as compared to neutral information is associated with different patterns in eye movement and neural activity. However, the 'emotionality' of a stimulus can be conveyed not only by its physical properties, but also by the information that is presented with it. There is very limited work examining the how emotional information may influence the immediate perceptual processing of otherwise neutral information. We examined how presenting an emotion label for a neutral face may influence subsequent processing by using eye movement monitoring (EMM) and magnetoencephalography (MEG) simultaneously. Participants viewed a series of faces with neutral expressions. Each face was followed by a unique negative or neutral sentence to describe that person, and then the same face was presented in isolation again. Viewing of faces paired with a negative sentence was associated with increased early viewing of the eye region and increased neural activity between 600 and 1200 ms in emotion processing regions such as the cingulate, medial prefrontal cortex, and amygdala, as well as posterior regions such as the precuneus and occipital cortex. Viewing of faces paired with a neutral sentence was associated with increased activity in the parahippocampal gyrus during the same time window. By monitoring behavior and neural activity within the same paradigm, these findings demonstrate that emotional information alters subsequent visual scanning and the neural systems that are presumably invoked to maintain a representation of the neutral information along with its emotional details. |
Lillian M. Rigoli; Daniel Holman; Michael J. Spivey; Christopher T. Kello In: Frontiers in Human Neuroscience, vol. 8, pp. 713, 2014. @article{Rigoli2014,When humans perform a response task or timing task repeatedly, fluctuations in measures of timing from one action to the next exhibit long-range correlations known as 1/f noise. The origins of 1/f noise in timing have been debated for over 20 years, with one common explanation serving as a default: humans are composed of physiological processes throughout the brain and body that operate over a wide range of timescales, and these processes combine to be expressed as a general source of 1/f noise. To test this explanation, the present study investigated the coupling vs. independence of 1/f noise in timing deviations, key-press durations, pupil dilations, and heartbeat intervals while tapping to an audiovisual metronome. All four dependent measures exhibited clear 1/f noise, regardless of whether tapping was synchronized or syncopated. 1/f spectra for timing deviations were found to match those for key-press durations on an individual basis, and 1/f spectra for pupil dilations matched those in heartbeat intervals. Results indicate a complex, multiscale relationship among 1/f noises arising from common sources, such as those arising from timing functions vs. those arising from autonomic nervous system (ANS) functions. Results also provide further evidence against the default hypothesis that 1/f noise in human timing is just the additive combination of processes throughout the brain and body. Our findings are better accommodated by theories of complexity matching that begin to formalize multiscale coordination as a foundation of human behavior. |
Jason Satel; Matthew D. Hilchey; Zhiguo Wang; Caroline S. Reiss; Raymond M. Klein In search of a reliable electrophysiological marker of oculomotor inhibition of return Journal Article In: Psychophysiology, vol. 51, no. 10, pp. 1037–1045, 2014. @article{Satel2014,Inhibition of return (IOR) operationalizes a behavioral phenomenon characterized by slower responding to cued, relative to uncued, targets. Two independent forms of IOR have been theorized: input-based IOR occurs when the oculomotor system is quiescent, while output-based IOR occurs when the oculomotor system is engaged. EEG studies forbidding eye movements have demonstrated that reductions of target-elicited P1 components are correlated with IOR magnitude, but when eye movements occur, P1 effects bear no relationship to behavior. We expand on this work by adapting the cueing paradigm and recording event-related potentials: IOR is caused by oculomotor responses to central arrows or peripheral onsets and measured by key presses to peripheral targets. Behavioral IOR is observed in both conditions, but P1 reductions are absent in the central arrow condition. By contrast, arrow and peripheral cues enhance Nd, especially over contralateral electrode sites. |
Lysianne Beynel; Alan Chauvin; Nathalie Guyader; Sylvain Harquel; Thierry Bougerol; Christian Marendaz; David Szekely What saccadic eye movements tell us about TMS-induced neuromodulation of the DLPFC and mood changes: A pilot study in bipolar disorders Journal Article In: Frontiers in Integrative Neuroscience, vol. 8, pp. 65, 2014. @article{Beynel2014,The study assumed that the antisaccade (AS) task is a relevant psychophysical tool to assess (i) short-term neuromodulation of the dorsolateral prefrontal cortex (DLPFC) induced by intermittent theta burst stimulation (iTBS); and (ii) mood change occurring during the course of the treatment. Saccadic inhibition is known to strongly involve the DLPFC, whose neuromodulation with iTBS requires less stimulation time and lower stimulation intensity, as well as results in longer aftereffects than the conventional repetitive transcranial magnetic stimulation (rTMS). Active or sham iTBS was applied every day for 3 weeks over the left DLPFC of 12 drug-resistant bipolar depressed patients. To assess the iTBS-induced short-term neuromodulation, the saccadic task was performed just before (S1) and just after (S2) the iTBS session, the first day of each week. Mood was evaluated through Montgomery and Asberg Depression Rating Scale (MADRS) scores and the difference in scores between the beginning and the end of treatment was correlated with AS performance change between these two periods. As expected, only patients from the active group improved their performance from S1 to S2 and mood improvement was significantly correlated with AS performance improvement. In addition, the AS task also discriminated depressive bipolar patients from healthy control subjects. Therefore, the AS task could be a relevant and useful tool for clinicians to assess if the Transcranial magnetic stimulation (TMS)-induced short-term neuromodulation of the DLPFC occurs as well as a “trait vs. state” objective marker of depressive mood disorder. |
Marc R. Kamke; Alexander E. Ryan; Martin V. Sale; Megan E. J. Campbell; Stephan Riek; Timothy J. Carroll; Jason B. Mattingley Visual spatial attention has opposite effects on bidirectional plasticity in the human motor cortex Journal Article In: Journal of Neuroscience, vol. 34, no. 4, pp. 1475–1480, 2014. @article{Kamke2014,Long-term potentiation (LTP) and long-term depression (LTD) are key mechanisms of synaptic plasticity that are thought to act in concert to shape neural connections. Here we investigated the influence of visual spatial attention on LTP-like and LTD-like plasticity in the human motor cortex. Plasticity was induced using paired associative stimulation (PAS), which involves repeated pairing of peripheral nerve stimulation and transcranial magnetic stimulation to alter functional responses in the thumb area of the primary motor cortex. PAS-induced changes in cortical excitability were assessed using motor-evoked potentials. During plasticity induction, participants directed their attention to one of two visual stimulus streams located adjacent to each hand. When participants attended to visual stimuli located near the left thumb, which was targeted by PAS, LTP-like increases in excitability were significantly enhanced, and LTD-like decreases in excitability reduced, relative to when they attended instead to stimuli located near the right thumb. These differential effects on (bidirectional) LTP-like and LTD-like plasticity suggest that voluntary visual attention can exert an important influence on the functional organization of the motor cortex. Specifically, attention acts to both enhance the strengthening and suppress the weakening of neural connections representing events that fall within the focus of attention. |
Kohitij Kar; Bart Krekelberg Transcranial alternating current stimulation attenuates visual motion adaptation Journal Article In: Journal of Neuroscience, vol. 34, no. 21, pp. 7334–7340, 2014. @article{Kar2014,Transcranial alternating current stimulation (tACS) is used in clinical applications and basic neuroscience research. Although its behavioral effects are evident from prior reports, current understanding of the mechanisms that underlie these effects is limited. We used motion perception, a percept with relatively well known properties and underlying neural mechanisms to investigate tACS mechanisms. Healthy human volunteers showed a surprising improvement in motion sensitivity when visual stimuli were paired with 10 Hz tACS. In addition, tACS reduced the motion-after effect, and this reduction was correlated with the improvement in motion sensitivity. Electrical stimulation had no consistent effect when applied before presenting a visual stimulus or during recovery from motion adaptation. Together, these findings suggest that perceptual effects of tACS result from an attenuation of adaptation. Important consequences for the practical use of tACS follow from our work. First, because this mechanism interferes only with adaptation, this suggests that tACS can be targeted at subsets of neurons (by adapting them), even when the applied currents spread widely throughout the brain. Second, by interfering with adaptation, this mechanism provides a means by which electrical stimulation can generate behavioral effects that outlast the stimulation. |
Benjamin D. Lester; Paul Dassonville The role of the right superior parietal lobule in processing visual context for the establishment of the egocentric reference frame Journal Article In: Journal of Cognitive Neuroscience, vol. 26, no. 10, pp. 2201–2209, 2014. @article{Lester2014,Visual cues contribute to the creation of an observerʼs ego- centric reference frame, within which the locations and orien- tations of objects can be judged. However, these cues can also be misleading. In the rod-and-frame illusion, for example, a large tilted frame distorts the observerʼs sense of vertical, caus- ing an enclosed rod to appear tilted in the opposite direction. To determine the brain region responsible for processing these spatial cues, we used TMS to suppress neural activity in the superior parietal lobule of healthy observers. Stimulation of the right hemisphere, but not the left, caused a significant reduc- tion in rod-and-frame susceptibility. In contrast, a tilt illusion caused by a mechanism that does not involve a distortion of the observerʼs egocentric reference frame was unaffected. These results demonstrate that the right superior parietal lobule is actively involved in processing the contextual cues that contribute to our perception of egocentric space. |
Indra T. Mahayana; Chia-Lun Liu; Chi Fu Chang; Daisy L. Hung; Ovid J. L. Tzeng; Chi-Hung Juan; Neil G. Muggleton Far-space neglect in conjunction but not feature search following transcranial magnetic stimulation over right posterior parietal cortex Journal Article In: Journal of Neurophysiology, vol. 111, no. 4, pp. 705–714, 2014. @article{Mahayana2014,Near- and far-space coding in the human brain is a dynamic process. Areas in dorsal, as well as ventral visual association cortex, including right posterior parietal cortex (rPPC), right frontal eye field (rFEF), and right ventral occipital cortex (rVO), have been shown to be important in visuospatial processing, but the involvement of these areas when the information is in near or far space remains unclear. There is a need for investigations of these representations to help explain the pathophysiology of hemispatial neglect, and the role of near and far space is crucial to this. We used a conjunction visual search task using an elliptical array to investigate the effects of transcranial magnetic stimulation delivered over rFEF, rPPC, and rVO on the processing of targets in near and far space and at a range of horizontal eccentricities. As in previous studies, we found that rVO was involved in far-space search, and rFEF was involved regardless of the distance to the array. It was found that rPPC was involved in search only in far space, with a neglect-like effect when the target was located in the most eccentric locations. No effects were seen for any site for a feature search task. As the search arrays had higher predictability with respect to target location than is often the case, these data may form a basis for clarifying both the role of PPC in visual search and its contribution to neglect, as well as the importance of near and far space in these. |
Muriel T. N. Panouillères; Ouazna Habchi; Peggy Gerardin; Roméo Salemme; Christian Urquizar; Alessandro Farnè; Denis Pélisson A role for the parietal cortex in sensorimotor adaptation of saccades Journal Article In: Cerebral Cortex, vol. 24, no. 2, pp. 304–314, 2014. @article{Panouilleres2014,Sensorimotor adaptation ensures movement accuracy despite continuously changing environment and body. Adaptation of saccadic eye movements is a classical model of sensorimotor adaptation. Beside the well-established role of the brainstem-cerebellum in the adaptation of reactive saccades (RSs), the cerebral cortex has been suggested to be involved in the adaptation of voluntary saccades (VSs). Here, we provide direct evidence for a causal involvement of the parietal cortex in saccadic adaptation. First, the posterior intraparietal sulcus (pIPS) was identified in each subject using functional magnetic resonance imaging (fMRI). Then, a saccadic adaptation paradigm was used to progressively reduce the amplitude of RSs and VSs, while single-pulse transcranial magnetic stimulation (spTMS) was applied over the right pIPS. The perturbations of pIPS resulted in impairment for the adaptation of VSs, selectively when spTMS was applied 60 ms after saccade onset. In contrast, the adaptation of RSs was facilitated by spTMS applied 90 ms after saccade initiation. The differential effect of spTMS relative to saccade types suggests a direct interference with pIPS activity for the VS adaptation and a remote interference with brainstem-cerebellum activity for the RS adaptation. These results support the hypothesis that the adaptation of VSs and RSs involves different neuronal substrates. |
L. L. Tanaka; J. C. Dessing; Pankhuri Malik; S. L. Prime; J. Douglas Crawford The effects of TMS over dorsolateral prefrontal cortex on trans-saccadic memory of multiple objects Journal Article In: Neuropsychologia, vol. 63, pp. 185–193, 2014. @article{Tanaka2014,Humans typically make several rapid eye movements (saccades) per second. It is thought that visual working memory can retain and spatially integrate three to four objects or features across each saccade but little is known about this neural mechanism. Previously we showed that transcranial magnetic stimulation (TMS) to the posterior parietal cortex and frontal eye fields degrade trans-saccadic memory of multiple object features (Prime, Vesia, & Crawford, 2008, Journal of Neuroscience, 28(27), 6938-6949; Prime, Vesia, & Crawford, 2010, Cerebral Cortex, 20(4), 759-772.). Here, we used a similar protocol to investigate whether dorsolateral prefrontal cortex (DLPFC), an area involved in spatial working memory, is also involved in trans-saccadic memory. Subjects were required to report changes in stimulus orientation with (saccade task) or without (fixation task) an eye movement in the intervening memory interval. We applied single-pulse TMS to left and right DLPFC during the memory delay, timed at three intervals to arrive approximately 100. ms before, 100. ms after, or at saccade onset. In the fixation task, left DLPFC TMS produced inconsistent results, whereas right DLPFC TMS disrupted performance at all three intervals (significantly for presaccadic TMS). In contrast, in the saccade task, TMS consistently facilitated performance (significantly for left DLPFC/. perisaccadic TMS and right DLPFC/. postsaccadic TMS) suggesting a dis-inhibition of trans-saccadic processing. These results are consistent with a neural circuit of trans-saccadic memory that overlaps and interacts with, but is partially separate from the circuit for visual working memory during sustained fixation. |
Lin-Yuan Tseng; Philip Tseng; Wei-Kuang Liang; Daisy L. Hung; Ovid J. L. Tzeng; Neil G. Muggleton; Chi-Hung Juan The role of superior temporal sulcus in the control of irrelevant emotional face processing: A transcranial direct current stimulation study Journal Article In: Neuropsychologia, vol. 64, pp. 124–133, 2014. @article{Tseng2014a,Emotional faces are often salient cues of threats or other important contexts, and may therefore have a large effect on cognitive processes of the visual environment. Indeed, many behavioral studies have demonstrated that emotional information can modulate visual attention and eye movements. The aim of the present study was to investigate (1) how irrelevant emotional face distractors affect saccadic behaviors and (2) whether such emotional effects reflect a specific neural mechanism or merely biased selective attention. We combined a visual search paradigm that incorporated manipulation of different types of distractor (fearful faces or scrambled faces) and delivered anodal transcranial direct current stimulation (tDCS) over the superior temporal sulcus and the frontal eye field to investigate the functional roles of these areas in processing facial expressions and eye movements. Our behavioral data suggest that irrelevant emotional distractors can modulate saccadic behaviors. The tDCS results showed that while rFEF played a more general role in controlling saccadic behavior, rSTS is mainly involved in facial expression processing. Furthermore, rSTS played a critical role in processing facial expressions even when such expressions were not relevant to the task goal, implying that facial expressions and processing may be automatic irrespective of the task goal. |
Jessica M. Wright; Bart Krekelberg Transcranial direct current stimulation over posterior parietal cortex modulates visuospatial localization Journal Article In: Journal of Vision, vol. 14, no. 9, pp. 5–5, 2014. @article{Wright2014a,Visual localization is based on the complex interplay of bottom-up and top-down processing. Based on previous work, the posterior parietal cortex (PPC) is assumed to play an essential role in this interplay. In this study, we investigated the causal role of the PPC in visual localization. Specifically, our goal was to determine whether modulation of the PPC via transcranial direct current stimulation (tDCS) could induce visual mislocalization similar to that induced by an exogenous attentional cue (Wright, Morris, & Krekelberg, 2011). We placed one stimulation electrode over the right PPC and the other over the left PPC (dual tDCS) and varied the polarity of the stimulation. We found that this manipulation altered visual localization; this supports the causal involvement of the PPC in visual localization. Notably, mislocalization was more rightward when the cathode was placed over the right PPC than when the anode was placed over the right PPC. This mislocalization was found within a few minutes of stimulation onset, it dissipated during stimulation, but then resurfaced after stimulation offset and lasted for another 10-15 min. On the assumption that excitability is reduced beneath the cathode and increased beneath the anode, these findings support the view that each hemisphere biases processing to the contralateral hemifield and that the balance of activation between the hemispheres contributes to position perception (Kinsbourne, 1977; Szczepanski, Konen, & Kastner, 2010). |
Daniel Baldauf; Robert Desimone Neural mechanisms of object-based attention Journal Article In: Science, vol. 344, no. 6182, pp. 424–427, 2014. @article{Baldauf2014,How we attend to objects and their features that cannot be separated by location is not understood. We presented two temporally and spatially overlapping streams of objects, faces versus houses, and used magnetoencephalography and functional magnetic resonance imaging to separate neuronal responses to attended and unattended objects. Attention to faces versus houses enhanced the sensory responses in the fusiform face area (FFA) and parahippocampal place area (PPA), respectively. The increases in sensory responses were accompanied by induced gamma synchrony between the inferior frontal junction, IFJ, and either FFA or PPA, depending on which object was attended. The IFJ appeared to be the driver of the synchrony, as gamma phases were advanced by 20 ms in IFJ compared to FFA or PPA. Thus, the IFJ may direct the flow of visual processing during object-based attention, at least in part through coupled oscillations with specialized areas such as FFA and PPA. W |
Marta Castellano; Michael Plöchl; Raul Vicente; Gordon Pipa Neuronal oscillations form parietal/frontal networks during contour integration Journal Article In: Frontiers in Integrative Neuroscience, vol. 8, pp. 64, 2014. @article{Castellano2014,The ability to integrate visual features into a global coherent percept that can be further categorized and manipulated are fundamental abilities of the neural system. While the processing of visual information involves activation of early visual cortices, the recruitment of parietal and frontal cortices has been shown to be crucial for perceptual processes. Yet is it not clear how both cortical and long-range oscillatory activity leads to the integration of visual features into a coherent percept. Here, we will investigate perceptual grouping through the analysis of a contour categorization task, where the local elements that form contour must be linked into a coherent structure, which is then further processed and manipulated to perform the categorization task. The contour formation in our visual stimulus is a dynamic process where, for the first time, visual perception of contours is disentangled from the onset of visual stimulation or from motor preparation, cognitive processes that until now have been behaviorally attached to perceptual processes. Our main finding is that, while local and long-range synchronization at several frequencies seem to be an ongoing phenomena, categorization of a contour could only be predicted through local oscillatory activity within parietal/frontal sources, which in turn, would synchronize at gamma (>30 Hz) frequency. Simultaneously, fronto-parietal beta (13-30 Hz) phase locking forms a network spanning across neural sources that are not category specific. Both long range networks, i.e., the gamma network that is category specific, and the beta network that is not category specific, are functionally distinct but spatially overlapping. Altogether, we show that a critical mechanism underlying contour categorization involves oscillatory activity within parietal/frontal cortices, as well as its synchronization across distal cortical sites. |
Peter Lissa; Genevieve McArthur; Stefan Hawelka; Romina Palermo; Yatin Mahajan; Florian Hutzler Fixation location on upright and inverted faces modulates the N170 Journal Article In: Neuropsychologia, vol. 57, no. 1, pp. 1–11, 2014. @article{Lissa2014,The current study used event-related potentials (ERP) in combination with a variable viewing position paradigm (VVPP) to direct fixations to specific face parts (eyes or mouths) in upright or inverted whole faces. The N170 elicited by the VVPP was greater to faces than to non-face objects (wristwatches), and was delayed and enhanced in response to face inversion. A larger N170 response was elicited when the participants[U+05F3] fixation was directed to the eyes than when directed to the mouths of both upright and inverted faces, an effect that was also modulated by the spatial location of the face in the visual field. The N170 face inversion effect (upright minus inverted) was greater when fixations were directed to the mouth than when directed to the eyes, suggesting that the point of fixation within a face modulates brain potentials due to contributions from the features themselves, as well as their relative location in the visual field. |
Adele Diederich; Annette Schomburg; Marieke K. Vugt Fronto-central theta oscillations are related to oscillations in saccadic response times (SRT): An EEG and behavioral data analysis Journal Article In: PLoS ONE, vol. 9, no. 11, pp. e112974, 2014. @article{Diederich2014,The phase reset hypothesis states that the phase of an ongoing neural oscillation, reflecting periodic fluctuations in neural activity between states of high and low excitability, can be shifted by the occurrence of a sensory stimulus so that the phase value become highly constant across trials (Schroeder et al., 2008). From EEG/MEG studies it has been hypothesized that coupled oscillatory activity in primary sensory cortices regulates multi sensory processing (Senkowski et al. 2008). We follow up on a study in which evidence of phase reset was found using a purely behavioral paradigm by including also EEG measures. In this paradigm, presentation of an auditory accessory stimulus was followed by a visual target with a stimulus-onset asynchrony (SOA) across a range from 0 to 404 ms in steps of 4 ms. This fine-grained stimulus presentation allowed us to do a spectral analysis on the mean SRT as a function of the SOA, which revealed distinct peak spectral components within a frequency range of 6 to 11 Hz with a modus of 7 Hz. The EEG analysis showed that the auditory stimulus caused a phase reset in 7-Hz brain oscillations in a widespread set of channels. Moreover, there was a significant difference in the average phase at which the visual target stimulus appeared between slow and fast SRT trials. This effect was evident in three different analyses, and occurred primarily in frontal and central electrodes. |
Tomer Fekete; Felix D. C. C. Beacher; Jiook Cha; Denis Rubin; Lilianne R. Mujica-Parodi Small-world network properties in prefrontal cortex correlate with predictors of psychopathology risk in young children: A NIRS study Journal Article In: NeuroImage, vol. 85, pp. 345–353, 2014. @article{Fekete2014,Near infrared spectroscopy (NIRS) is an emerging imaging technique that is relatively inexpensive, portable, and particularly well suited for collecting data in ecological settings. Therefore, it holds promise as a potential neurodiagnostic for young children. We set out to explore whether NIRS could be utilized in assessing the risk of developmental psychopathology in young children. A growing body of work indicates that temperament at young age is associated with vulnerability to psychopathology later on in life. In particular, it has been shown that low effortful control (EC), which includes the focusing and shifting of attention, inhibitory control, perceptual sensitivity, and a low threshold for pleasure, is linked to conditions such as anxiety, depression and attention deficit hyperactivity disorder (ADHD). Physiologically, EC has been linked to a control network spanning among other sites the prefrontal cortex. Several psychopathologies, such as depression and ADHD, have been shown to result in compromised small-world network properties. Therefore we set out to explore the relationship between EC and the small-world properties of PFC using NIRS. NIRS data were collected from 44 toddlers, ages 3-5, while watching naturalistic stimuli (movie clips). Derived complex network measures were then correlated to EC as derived from the Children's Behavior Questionnaire (CBQ). We found that reduced levels of EC were associated with compromised small-world properties of the prefrontal network. Our results suggest that the longitudinal NIRS studies of complex network properties in young children hold promise in furthering our understanding of developmental psychopathology. |
Ruth Filik; Hartmut Leuthold; Katie Wallington; Jemma Page Testing theories of irony processing using eye-tracking and ERPs Journal Article In: Journal of Experimental Psychology: Learning, Memory, and Cognition, vol. 40, no. 3, pp. 811–828, 2014. @article{Filik2014,Not much is known about how people comprehend ironic utterances, and to date, most studies have simply compared processing of ironic versus non-ironic statements. A key aspect of the graded salience hypothesis, distinguishing it from other accounts (such as the standard pragmatic view and direct access view), is that it predicts differences between processing of familiar and unfamiliar ironies. Specifically, if an ironic utterance is familiar, then the ironic interpretation should be available without the need for extra inferential processes, whereas for unfamiliar ironies, the literal interpretation would be computed first, and a mismatch with context would lead to a re-interpretation of the statement as being ironic. We recorded participants' eye movements while they were reading (Experiment 1), and electrical brain activity while they were listening to (Experiment 2), familiar and unfamiliar ironies compared to non-ironic controls. Results show disruption to eye movements and an N400-like effect for unfamiliar ironies only, supporting the predictions of the graded salience hypothesis. In addition, in Experiment 2, a late positivity was found for both familiar and unfamiliar ironic materials, compared to non-ironic controls. We interpret this positivity as reflecting ongoing conflict between the literal and ironic interpretations of the utterance. |
Min Suk Kang; Geoffrey F. Woodman The neurophysiological index of visual working memory maintenance is not due to load dependent eye movements Journal Article In: Neuropsychologia, vol. 56, no. 1, pp. 63–72, 2014. @article{Kang2014,The Contralateral Delayed Activity (CDA) is slow negative potential found during a variety of tasks, providing an important measure of the representation of information in visual working memory. However, it is studied using stimulus arrays in which the to-be-remembered objects are shown in the periphery of the left or the right visual field. Our goal was to determine whether fixational eye movements in the direction of the memoranda might underlie the CDA. We found that subjects' gaze was shifted toward the visual field of the memoranda during the retention interval, with its magnitude increasing with the set size. However, the CDA was clearly observed even when the subjects' gaze shifts were absent. In addition, the magnitude of the subjects' gaze shifts was unrelated to their visual working memory capacity measured with behavioral data, unlike the CDA. Finally, the onset latency of the set size dependent eye movements followed the onset of the set size dependent CDA. Thus, our findings clearly show that the CDA does not represent a simple inability to maintain fixation during visual working memory maintenance, but that this neural index of representation in working memory appears to induce eye movements toward the locations of the objects being remembered. |
Lisandro N. Kaunitz; Juan E. Kamienkowski; Alexander Varatharajah; Mariano Sigman; Rodrigo Quian Quiroga; Matias J. Ison Looking for a face in the crowd: Fixation-related potentials in an eye-movement visual search task Journal Article In: NeuroImage, vol. 89, pp. 297–305, 2014. @article{Kaunitz2014,Despite the compelling contribution of the study of event related potentials (ERPs) and eye movements to cognitive neuroscience, these two approaches have largely evolved independently. We designed an eye-movement visual search paradigm that allowed us to concurrently record EEG and eye movements while subjects were asked to find a hidden target face in a crowded scene with distractor faces. Fixation event-related potentials (fERPs) to target and distractor stimuli showed the emergence of robust sensory components associated with the perception of stimuli and cognitive components associated with the detection of target faces. We compared those components with the ones obtained in a control task at fixation: qualitative similarities as well as differences in terms of scalp topography and latency emerged between the two. By using single trial analyses, fixations to target and distractors could be decoded from the EEG signals above chance level in 11 out of 12 subjects. Our results show that EEG signatures related to cognitive behavior develop across spatially unconstrained exploration of natural scenes and provide a first step towards understanding the mechanisms of target detection during natural search. |
Christof Körner; Verena Braunstein; Matthias Stangl; Alois Schlögl; Christa Neuper; Anja Ischebeck In: Psychophysiology, vol. 51, no. 4, pp. 385–395, 2014. @article{Koerner2014,To search for a target in a complex environment is an everyday behavior that ends with finding the target. When we search for two identical targets, however, we must continue the search after finding the first target and memorize its location. We used fixation-related potentials to investigate the neural correlates of different stages of the search, that is, before and after finding the first target. Having found the first target influenced subsequent distractor processing. Compared to distractor fixations before the first target fixation, a negative shift was observed for three subsequent distractor fixations. These results suggest that processing a target in continued search modulates the brain's response, either transiently by reflecting temporary working memory processes or permanently by reflecting working memory retention. |
Timothy Leffel; Miriam Lauter; Masha Westerlund; Liina Pylkkänen Restrictive vs. non-restrictive composition: A magnetoencephalography study Journal Article In: Language, Cognition and Neuroscience, vol. 29, no. 10, pp. 1191–1204, 2014. @article{Leffel2014,Recent research on the brain mechanisms underlying language processing has implicated the left anterior temporal lobe (LATL) as a central region for the composition of simple phrases. Because these studies typically present their critical stimuli without contextual information, the sensitivity of LATL responses to contextual factors is unknown. In this magnetoencephalography (MEG) study, we employed a simple question-answer paradigm to manipulate whether a prenominal adjective or determiner is interpreted restrictively, i.e., as limiting the set of entities under discussion. Our results show that the LATL is sensitive to restriction, with restrictive composition eliciting higher responses than non-restrictive composition. However, this effect was only observed when the restricting element was a determiner, adjectival stimuli showing the opposite pattern, which we hypothesise to be driven by the special pragmatic properties of non-restrictive adjectives. Overall, our results demonstrate a robust sensitivity of the LATL to high level contextual and potentially also pragmatic factors. |
Heath Matheson; Aaron J. Newman; Jason Satel; Patricia A. McMullen Handles of manipulable objects attract covert visual attention: ERP evidence Journal Article In: Brain and Cognition, vol. 86, no. 1, pp. 17–23, 2014. @article{Matheson2014,Previous research has demonstrated that people are faster at making a manual response with the hand that is aligned with the handle of a manipulable object compared to its functional end. According to theories of embodied cognition (ETC), the presentation of a manipulable object automatically elicits sensorimotor simulations of the respective hand and these simulations facilitate the response. However, an alternative interpretation of these data is that handles preferentially attract visual attention, since attended stimuli and locations typically elicit faster responses. We investigated attentional biases elicited by manipulable and non-manipulable objects using event-related-potentials (ERPs). On each trial, a picture of a manipulable object was followed by a target dot that participants had to make a button-press to. The dot was located at either the handle or functional end of the object. Consistent with previous attentional cuing paradigms, we showed that the P1 ERP component was greater in response to targets cued by handles than by functional ends. These results suggest that object handles automatically bias covert attentional processes. These attentional biases may account for earlier behavioural findings, without any recourse to ETC. |
Stéphanie M. Morand; Monika Harvey; Marie-Hélène Grosbras Parieto-occipital cortex shows early target selection to faces in a reflexive orienting task Journal Article In: Cerebral Cortex, vol. 24, no. 4, pp. 898–907, 2014. @article{Morand2014,It is well established that human faces induce stronger involuntary orienting responses than other visual objects. Yet, the timing of this preferential orienting response at the neural level is still unknown. Here, we used an antisaccade paradigm to investigate the neural dynamics preceding the onset of reflexive and voluntary saccades elicited by human faces and nonface visual objects, normalized for their global low-level visual properties. High-density event-related potentials (ERPs) were recorded in observers as they performed interleaved pro- and antisaccades toward a lateralized target. For reflexive saccades, we report an ERP modulation specific to faces as early as 40–60 ms following stimulus onset over parieto-occipital sites, further predicting the speed of saccade execution. This was not linked to differences in the programming of the saccadic eye movements, as it occurred early in time. For the first time, we present electrophysiological evidence of early target selection to faces in reflexive orienting responses over parieto-occipital cortex that facilitates the triggering of saccades toward faces. We argue for a 2-stage process in the representation of a face in involuntary spatial orienting with an initial, rapid implicit processing of the visual properties of a face, followed by subsequent stimulus categorization depicted by the N170 component. |
Hans Peter Frey; Anita M. Schmid; Jeremy W. Murphy; Sophie Molholm; Edmund C. Lalor; John J. Foxe Modulation of early cortical processing during divided attention to non-contiguous locations Journal Article In: European Journal of Neuroscience, vol. 39, no. 9, pp. 1499–1507, 2014. @article{Frey2014,We often face the challenge of simultaneously attending to multiple non-contiguous regions of space. There is ongoing debate as to how spatial attention is divided under these situations. Whereas, for several years, the predominant view was that humans could divide the attentional spotlight, several recent studies argue in favor of a unitary spotlight that rhythmically samples relevant locations. Here, this issue was addressed by the use of high-density electrophysiology in concert with the multifocal m-sequence technique to examine visual evoked responses to multiple simultaneous streams of stimulation. Concurrently, we assayed the topographic distribution of alpha-band oscillatory mechanisms, a measure of attentional suppression. Participants performed a difficult detection task that required simultaneous attention to two stimuli in contiguous (undivided) or non-contiguous parts of space. In the undivided condition, the classic pattern of attentional modulation was observed, with increased amplitude of the early visual evoked response and increased alpha amplitude ipsilateral to the attended hemifield. For the divided condition, early visual responses to attended stimuli were also enhanced, and the observed multifocal topographic distribution of alpha suppression was in line with the divided attention hypothesis. These results support the existence of divided attentional spotlights, providing evidence that the corresponding modulation occurs during initial sensory processing time-frames in hierarchically early visual regions, and that suppressive mechanisms of visual attention selectively target distracter locations during divided spatial attention. |
Galit Fuhrmann Alpert; Ran Manor; Assaf B. Spanier; Leon Y. Deouell; Amir B. Geva Spatiotemporal representations of rapid visual target detection: A single-trial EEG classification algorithm Journal Article In: IEEE Transactions on Biomedical Engineering, vol. 61, no. 8, pp. 2290–2303, 2014. @article{FuhrmannAlpert2014a,Brain computer interface applications, developed for both healthy and clinical populations, critically depend on decoding brain activity in single trials. The goal of the present study was to detect distinctive spatiotemporal brain patterns within a set of event related responses. We introduce a novel classification algorithm, the spatially weighted FLD-PCA (SWFP), which is based on a two-step linear classification of event-related responses, using fisher linear discriminant (FLD) classifier and principal component analysis (PCA) for dimensionality reduction. As a benchmark algorithm, we consider the hierarchical discriminant component Analysis (HDCA), introduced by Parra, et al. 2007. We also consider a modified version of the HDCA, namely the hierarchical discriminant principal component analysis algorithm (HDPCA). We compare single-trial classification accuracies of all the three algorithms, each applied to detect target images within a rapid serial visual presentation (RSVP, 10 Hz) of images from five different object categories, based on single-trial brain responses. We find a systematic superiority of our classification algorithm in the tested paradigm. Additionally, HDPCA significantly increases classification accuracies compared to the HDCA. Finally, we show that presenting several repetitions of the same image exemplars improve accuracy, and thus may be important in cases where high accuracy is crucial. |
Galit Fuhrmann Alpert; Ran Manor; Assaf B. Spanier; Leon Y. Deouell; Amir B. Geva Spatio-temporal representations of rapid visual target detection: A single trial EEG classification Journal Article In: IEEE Transactions on Biomedical Engineering, vol. 61, no. 8, pp. 2290–2303, 2014. @article{FuhrmannAlpert2014,Brain computer interface applications, developed for both healthy and clinical populations, critically depend on decod- ing brain activity in single trials. The goal of the present study was to detect distinctive spatiotemporal brain patterns within a set of event related responses. We introduce a novel classification algo- rithm, the spatially weighted FLD-PCA (SWFP), which is based on a two-step linear classification of event-related responses, using fisher linear discriminant (FLD) classifier and principal compo- nent analysis (PCA) for dimensionality reduction. As a benchmark algorithm, we consider the hierarchical discriminant component Analysis (HDCA), introduced by Parra, et al. 2007. We also con- sider a modified version of the HDCA, namely the hierarchical discriminant principal component analysis algorithm (HDPCA). We compare single-trial classification accuracies of all the three algorithms, each applied to detect target images within a rapid serial visual presentation (RSVP, 10 Hz) of images from five dif- ferent object categories, based on single-trial brain responses. We find a systematic superiority of our classification algorithm in the tested paradigm.Additionally, HDPCA significantly increases clas- sification accuracies compared to the HDCA. Finally, we show that presenting several repetitions of the same image exemplars im- prove accuracy, and thus may be important in cases where high accuracy is crucial. |
Carlos M. Hamamé; Juan R. Vidal; Marcela Perrone-Bertolotti; Tomás Ossandón; Karim Jerbi; Philippe Kahane; Olivier Bertrand; Jean Philippe Lachaux Functional selectivity in the human occipitotemporal cortex during natural vision: Evidence from combined intracranial EEG and eye-tracking Journal Article In: NeuroImage, vol. 95, pp. 276–286, 2014. @article{Hamame2014,Eye movements are a constant and essential component of natural vision, yet, most of our knowledge about the human visual system comes from experiments that restrict them. This experimental constraint is mostly in place to control visual stimuli presentation and to avoid artifacts in non-invasive measures of brain activity, however, this limitation can be overcome with intracranial EEG (iEEG) recorded from epilepsy patients. Moreover, the high-frequency components of the iEEG signal (between about 50 and 150. Hz) can provide a proxy of population-level spiking activity in any cortical area during free-viewing. We combined iEEG with high precision eye-tracking to study fine temporal dynamics and functional specificity in the fusiform face (FFA) and visual word form area (VWFA) while patients inspected natural pictures containing faces and text. We defined the first local measure of visual (electrophysiological) responsiveness adapted to free-viewing in humans: amplitude modulations in the high-frequency activity range (50-150. Hz) following fixations (fixation-related high-frequency response). We showed that despite the large size of receptive fields in the ventral occipito-temporal cortex, neural activity during natural vision of realistic cluttered scenes is mostly dependent upon the category of the foveated stimulus - suggesting that category-specificity is preserved during free-viewing and that attention mechanisms might filter out the influence of objects surrounding the fovea. |
Wei He; Jon Brock; Blake W. Johnson Face-sensitive brain responses measured from a four-year-old child with a custom-sized child MEG system Journal Article In: Journal of Neuroscience Methods, vol. 222, pp. 213–217, 2014. @article{He2014,Background: Previous magnetoencephalography (MEG) studies have failed to find a facesensitive, brain response-M170 in children. If this is the case, this suggests that the developmental trajectory of the M170 is different from that of its electrical equivalent, the N170. We investigated the alternative possibility that the child M170 may not be detectable in conventional adult-sized MEG systems. New method: Brain responses to pictures of faces and well controlled stimuli were measured from the same four-year-old child with a custom child MEG system and an adult-sized MEG system. Results: The goodness of fit of the child's head was about the same over the occipital head surface in both systems, but was much worse over all other parts of the head surface in the adult MEG system compared to the child MEG system. The face-sensitive M170 was measured from the child in both MEG systems, but was larger in amplitude, clearer in morphology, and had a more accurate source localization when measured in the child MEG system. Comparison with existing method: The custom-sized child MEG system is superior for measuring the face-sensitive M170 brain response in children than the conventional adult MEG system. Conclusions: The present results show that the face-sensitive M170 brain response can be elicited in a four-year-old child. This provides new evidence for early maturation of face processing brain mechanisms in humans, and offers new opportunities for the study of neurodevelopmental disorders that show atypical face processing capabilities, such as autism spectrum disorder. |
Randolph F. Helfrich; Hannah Knepper; Guido Nolte; Daniel Strüber; Stefan Rach; Christoph S. Herrmann; Till R. Schneider; Andreas K. Engel Selective modulation of interhemispheric functional connectivity by HD-tACS shapes perception Journal Article In: PLoS Biology, vol. 12, no. 12, pp. 1–15, 2014. @article{Helfrich2014,Oscillatory neuronal synchronization between cortical areas has been suggested to constitute a flexible mechanism to coordinate information flow in the human cerebral cortex. However, it remains unclear whether synchronized neuronal activity merely represents an epiphenomenon or whether it is causally involved in the selective gating of information. Here, we combined bilateral high-density transcranial alternating current stimulation (HD-tACS) at 40 Hz with simultaneous electroencephalographic (EEG) recordings to study immediate electrophysiological effects during the selective entrainment of oscillatory gamma-band signatures. We found that interhemispheric functional connectivity was modulated in a predictable, phase-specific way: In-phase stimulation enhanced synchronization, anti-phase stimulation impaired functional coupling. Perceptual correlates of these connectivity changes were found in an ambiguous motion task, which strongly support the functional relevance of long-range neuronal coupling. Additionally, our results revealed a decrease in oscillatory alpha power in response to the entrainment of gamma band signatures. This finding provides causal evidence for the antagonistic role of alpha and gamma oscillations in the parieto-occipital cortex and confirms that the observed gamma band modulations were physiological in nature. Our results demonstrate that synchronized cortical network activity across several spatiotemporal scales is essential for conscious perception and cognition. |
Kasey S. Hemington; James N. Reynolds In: Clinical Neurophysiology, vol. 125, no. 12, pp. 2364–2371, 2014. @article{Hemington2014,Objective: Children with Fetal Alcohol Spectrum Disorder (FASD) exhibit cognitive deficits that can be probed using eye movement tasks. We employed a recently developed, single-sensor electroencephalographic (EEG) recording device in measuring EEG activity during the performance of an eye movement task probing working memory in this population. Methods: Children with FASD (n= 18) and typically developing children (n= 19) performed a memory-guided saccade task requiring the participant to remember the spatial location of one, two or three stimuli. We hypothesized that children with FASD would (i) exhibit performance deficits, particularly at greater mnemonic loads; and (ii) display differences in theta (4-8 Hz) and alpha (8-12 Hz) frequency band power compared with controls. Results: Children with FASD failed to perform the task correctly more often than controls when presented with two or three stimuli, and demonstrated related reductions in alpha and theta power. Conclusion: These data suggest that the memory-guided task is sensitive to working memory deficits in children with FASD. Significance: Simultaneous recording of EEG activity suggest differing patterns of underlying neural recruitment in the clinical group, consistent with previous literature indicating more cognitive resources are required by children with FASD in order to complete complex tasks correctly. |
Linbi Hong; Jennifer M. Walz; Paul Sajda Your eyes give you away: Prestimulus changes in pupil diameter correlate with poststimulus task-related EEG dynamics Journal Article In: PLoS ONE, vol. 9, no. 3, pp. e91321, 2014. @article{Hong2014,Pupillary measures have been linked to arousal and attention as well as activity in the brainstem's locus coeruleus norepinephrine (LC-NE) system. Similarly, there is evidence that evoked EEG responses, such as the P3, might have LC-NE activity as their basis. Since it is not feasible to record electrophysiological data directly from the LC in humans due to its location in the brainstem, an open question has been whether pupillary measures and EEG variability can be linked in a meaningful way to shed light on the nature of the LC-NE role in attention and arousal. We used an auditory oddball task with a data-driven approach to learn task-relevant projections of the EEG, for windows of data spanning the entire trial. We investigated linear and quadratic relationships between the evoked EEG along these projections and both prestimulus (baseline) and poststimulus (evoked dilation) pupil diameter measurements. We found that baseline pupil diameter correlates with early (175-200 ms) and late (350-400 ms) EEG component variability, suggesting a linear relationship between baseline (tonic) LC-NE activity and evoked EEG. We found no relationships between evoked EEG and evoked pupil dilation, which is often associated with evoked (phasic) LC activity. After regressing out reaction time (RT), the correlation between EEG variability and baseline pupil diameter remained, suggesting that such correlation is not explainable by RT variability. We also investigated the relationship between these pupil measures and prestimulus EEG alpha activity, which has been reported as a marker of attentional state, and found a negative linear relationship with evoked pupil dilation. In summary, our results demonstrate significant relationships between prestimulus and poststimulus neural and pupillary measures, and they provide further evidence for tight coupling between attentional state and evoked neural activity and for the role of cortical and subcortical networks underlying the process of target detection. |
Jörn M. Horschig; Ole Jensen; Martine R. Schouwenburg; Roshan Cools; Mathilde Bonnefond Alpha activity reflects individual abilities to adapt to the environment Journal Article In: NeuroImage, vol. 89, pp. 235–243, 2014. @article{Horschig2014,Recent findings suggest that oscillatory alpha activity (7-13. Hz) is associated with functional inhibition of sensory regions by filtering incoming information. Accordingly the alpha power in visual regions varies in anticipation of upcoming, predictable stimuli which has consequences for visual processing and subsequent behavior. In covert spatial attention studies it has been demonstrated that performance correlates with the adaptation of alpha power in response to explicit spatial cueing. However it remains unknown whether such an adaptation also occurs in response to implicit statistical properties of a task. In a covert attention switching paradigm, we here show evidence that individuals differ on how they adapt to implicit statistical properties of the task. Subjects whose behavioral performance reflects the implicit change in switch trial likelihood show strong adjustment of anticipatory alpha power lateralization. Most importantly, the stronger the behavioral adjustment to the switch trial likelihood was, the stronger the adjustment of anticipatory posterior alpha lateralization. We conclude that anticipatory spatial attention is reflected in the distribution of posterior alpha band power which is predictive of individual detection performance in response to the implicit statistical properties of the task. |
Leyla Isik; Ethan M. Meyers; Joel Z. Leibo; Tomaso Poggio The dynamics of invariant object recognition in the human visual system Journal Article In: Journal of Neurophysiology, vol. 111, no. 1, pp. 91–102, 2014. @article{Isik2014,The human visual system can rapidly recognize objects despite transformations that alter their appearance. The precise timing of when the brain computes neural representations that are invariant to particular transformations, however, has not been mapped in humans. Here we employ magnetoencephalography decoding analysis to measure the dynamics of size- and position-invariant visual information development in the ventral visual stream. With this method we can read out the identity of objects beginning as early as 60 ms. Size- and position-invariant visual information appear around 125 ms and 150 ms, respectively, and both develop in stages, with invariance to smaller transformations arising before invariance to larger transformations. Additionally, the magnetoencephalography sensor activity localizes to neural sources that are in the most posterior occipital regions at the early decoding times and then move temporally as invariant information develops. These results provide previously unknown latencies for key stages of human-invariant object recognition, as well as new and compelling evidence for a feed-forward hierarchical model of invariant object recognition where invariance increases at each successive visual area along the ventral stream. |
David C. Jangraw; Jun Wang; Brent J. Lance; Shih Fu Chang; Paul Sajda Neurally and ocularly informed graph-based models for searching 3D environments Journal Article In: Journal of Neural Engineering, vol. 11, no. 4, pp. 1–13, 2014. @article{Jangraw2014a,OBJECTIVE: As we move through an environment, we are constantly making assessments, judgments and decisions about the things we encounter. Some are acted upon immediately, but many more become mental notes or fleeting impressions-our implicit 'labeling' of the world. In this paper, we use physiological correlates of this labeling to construct a hybrid brain-computer interface (hBCI) system for efficient navigation of a 3D environment. APPROACH: First, we record electroencephalographic (EEG), saccadic and pupillary data from subjects as they move through a small part of a 3D virtual city under free-viewing conditions. Using machine learning, we integrate the neural and ocular signals evoked by the objects they encounter to infer which ones are of subjective interest to them. These inferred labels are propagated through a large computer vision graph of objects in the city, using semi-supervised learning to identify other, unseen objects that are visually similar to the labeled ones. Finally, the system plots an efficient route to help the subjects visit the 'similar' objects it identifies. MAIN RESULTS: We show that by exploiting the subjects' implicit labeling to find objects of interest instead of exploring naively, the median search precision is increased from 25% to 97%, and the median subject need only travel 40% of the distance to see 84% of the objects of interest. We also find that the neural and ocular signals contribute in a complementary fashion to the classifiers' inference of subjects' implicit labeling. SIGNIFICANCE: In summary, we show that neural and ocular signals reflecting subjective assessment of objects in a 3D environment can be used to inform a graph-based learning model of that environment, resulting in an hBCI system that improves navigation and information delivery specific to the user's interests. |
Yoshihito Shigihara; Semir Zeki Parallel processing of face and house stimuli by V1 and specialized visual areas: A magnetoencephalographic (MEG) study Journal Article In: Frontiers in Human Neuroscience, vol. 8, pp. 901, 2014. @article{Shigihara2014,We used easily distinguishable stimuli of faces and houses constituted from straight lines, with the aim of learning whether they activate V1 on the one hand, and the specialized areas that are critical for the processing of faces and houses on the other, with similar latencies. Eighteen subjects took part in the experiment, which used magnetoencephalography (MEG) coupled to analytical methods to detect the time course of the earliest responses which these stimuli provoke in these cortical areas. Both categories of stimuli activated V1 and areas of the visual cortex outside it at around 40 ms after stimulus onset, and the amplitude elicited by face stimuli was significantly larger than that elicited by house stimuli. These results suggest that "low-level" and "high-level" features of form stimuli are processed in parallel by V1 and visual areas outside it. Taken together with our previous results on the processing of simple geometric forms (Shgihara and Zeki, 2013; Shigihara and Zeki, 2014), the present ones reinforce the conclusion that parallel processing is an important component in the strategy used by the brain to process and construct forms. |
Eelke Spaak; Floris P. Lange; Ole Jensen Local entrainment of alpha oscillations by visual stimuli causes cyclic modulation of perception Journal Article In: Journal of Neuroscience, vol. 34, no. 10, pp. 3536–3544, 2014. @article{Spaak2014,Prestimulus oscillatory neural activity in the visual cortex has large consequences for perception and can be influenced by top-down control from higher-order brain regions. Making a causal claim about the mechanistic role of oscillatory activity requires that oscillations be directly manipulated independently of cognitive instructions. There are indications that a direct manipulation, or entrainment, of visual alpha activity is possible through visual stimulation. However, three important questions remain: (1) Can the entrained alpha activity be endogenously maintained in the absence of continuous stimulation?; (2) Does entrainment of alpha activity reflect a global or a local process?; and (3) Does the entrained alpha activity influence perception? To address these questions, we presented human subjects with rhythmic stimuli in one visual hemifield, and arhythmic stimuli in the other. After rhythmic entrainment, we found a periodic pattern in detection performance of near-threshold targets specific to the entrained hemifield. Using magnetoencephalograhy to measure ongoing brain activity, we observed strong alpha activity contralateral to the rhythmic stimulation outlasting the stimulation by several cycles. This entrained alpha activity was produced locally in early visual cortex, as revealed by source analysis. Importantly, stronger alpha entrainment predicted a stronger phasic modulation of detection performance in the entrained hemifield. These findings argue for a cortically focal entrainment of ongoing alpha oscillations by visual stimulation, with concomitant consequences for perception. Our results support the notion that oscillatory brain activity in the alpha band provides a causal mechanism for the temporal organization of visual perception. |
Masha Westerlund; Liina Pylkkänen The role of the left anterior temporal lobe in semantic composition vs. semantic memory Journal Article In: Neuropsychologia, vol. 57, no. 1, pp. 59–70, 2014. @article{Westerlund2014,The left anterior temporal lobe (LATL) is robustly implicated in semantic processing by a growing body of literature. However, these results have emerged from two distinct bodies of work, addressing two different processing levels. On the one hand, the LATL has been characterized as a 'semantic hub׳ that binds features of concepts across a distributed network, based on results from semantic dementia and hemodynamic findings on the categorization of specific compared to basic exemplars. On the other, the LATL has been implicated in combinatorial operations in language, as shown by increased activity in this region associated with the processing of sentences and of basic phrases. The present work aimed to reconcile these two literatures by independently manipulating combination and concept specificity within a minimal MEG paradigm. Participants viewed simple nouns that denoted either low specificity (fish) or high specificity categories (trout) presented in either combinatorial (spotted fish/trout) or non-combinatorial contexts (xhsl fish/trout). By combining these paradigms from the two literatures, we directly compared the engagement of the LATL in semantic memory vs. semantic composition. Our results indicate that although noun specificity subtly modulates the LATL activity elicited by single nouns, it most robustly affects the size of the composition effect when these nouns are adjectivally modified, with low specificity nouns eliciting a much larger effect. We conclude that these findings are compatible with an account in which the specificity and composition effects arise from a shared mechanism of meaning specification. |
2013 |
Yoshihito Shigihara; Semir Zeki Parallelism in the brain's visual form system Journal Article In: European Journal of Neuroscience, vol. 38, no. 12, pp. 3712–3720, 2013. @article{Shigihara2013,We used magnetoencephalography (MEG) to determine whether increasingly complex forms constituted from the same elements (lines) activate visual cortex with the same or different latencies. Twenty right-handed healthy adult volunteers viewed two different forms, lines and rhomboids, representing two levels of complexity. Our results showed that the earliest responses produced by lines and rhomboids in both striate and prestriate cortex had similar peak latencies (40 ms) although lines produced stronger responses than rhomboids. Dynamic causal modeling (DCM) showed that a parallel multiple input model to striate and prestriate cortex accounts best for the MEG response data. These results lead us to conclude that the perceptual hierarchy between lines and rhomboids is not mirrored by a temporal hierarchy in latency of activation and thus that a strategy of parallel processing appears to be used to construct forms, without implying that a hierarchical strategy may not be used in separate visual areas, in parallel. |
Julia M. Stephen; Brian A. Coffman; David B. Stone; Piyadasa Kodituwakku Differences in MEG gamma oscillatory power during performance of a prosaccade task in adolescents with FASD Journal Article In: Frontiers in Human Neuroscience, vol. 7, pp. 900, 2013. @article{Stephen2013,Fetal alcohol spectrum disorder (FASD) is characterized by a broad range of behavioral and cognitive deficits that impact the long-term quality of life for affected individuals. However, the underlying changes in brain structure and function associated with these cognitive impairments are not well-understood. Previous studies identified deficits in behavioral performance of prosaccade tasks in children with FASD. In this study, we investigated group differences in gamma oscillations during performance of a prosaccade task. We collected magnetoencephalography (MEG) data from 15 adolescents with FASD and 20 age-matched healthy controls (HC) with a mean age of 15.9 ± 0.4 years during performance of a prosaccade task. Eye movement was recorded and synchronized to the MEG data using an MEG compatible eye-tracker. The MEG data were analyzed relative to the onset of the visual saccade. Time-frequency analysis was performed using Fieldtrip with a focus on group differences in gamma-band oscillations. Following left target presentation, we identified four clusters over right frontal, right parietal, and left temporal/occipital cortex, with significantly different gamma-band (30-50 Hz) power between FASD and HC. Furthermore, visual M100 latencies described in Coffman etal. (2012) corresponded with increased gamma power over right central cortex in FASD only. Gamma-band differences were not identified for stimulus-averaged responses implying that these gamma-band differences were related to differences in saccade network functioning. These differences in gamma-band power may provide indications of atypical development of cortical networks in individuals with FASD. |
Durk Talsma; Brian J. White; Sebastiaan Mathôt; Douglas P. Munoz; Jan Theeuwes A retinotopic attentional trace after saccadic eye movements: Evidence from event-related potentials Journal Article In: Journal of Cognitive Neuroscience, vol. 25, no. 9, pp. 1563–1577, 2013. @article{Talsma2013,Saccadic eye movements are a major source of disruption to visual stability, yet we experience little of this disruption. We can keep track of the same object across multiple saccades. It is generally assumed that visual stability is due to the process of remapping, in which retinotopically organized maps are updated to compensate for the retinal shifts caused by eye movements. Recent behavioral and ERP evidence suggests that visual attention is also remapped, but that it may still leave a residual retinotopic trace immediately after a saccade. The current study was designed to further examine electrophysiological evidence for such a retinotopic trace by recording ERPs elicited by stimuli that were presented immediately after a saccade (80 msec SOA). Participants were required to maintain attention at a specific location (and to memorize this location) while making a saccadic eye movement. Immediately after the saccade, a visual stimulus was briefly presented at either the attended location (the same spatiotopic location), a location that matched the attended location retinotopically (the same retinotopic location), or one of two control locations. ERP data revealed an enhanced P1 amplitude for the stimulus presented at the retinotopically matched location, but a significant attenuation for probes presented at the original attended location. These results are consistent with the hypothesis that visuospatial attention lingers in retinotopic coordinates immediately following gaze shifts. |
Heng Ru May Tan; Hartmut Leuthold; Joachim Gross Gearing up for action: Attentive tracking dynamically tunes sensory and motor oscillations in the alpha and beta band Journal Article In: NeuroImage, vol. 82, pp. 634–644, 2013. @article{Tan2013,Allocation of attention during goal-directed behavior entails simultaneous processing of relevant and attenuation of irrelevant information. How the brain delegates such processes when confronted with dynamic (biological motion) stimuli and harnesses relevant sensory information for sculpting prospective responses remains unclear. We analyzed neuromagnetic signals that were recorded while participants attentively tracked an actor's pointing movement that ended at the location where subsequently the response-cue indicated the required response. We found the observers' spatial allocation of attention to be dynamically reflected in lateralized parieto-occipital alpha (8-12. Hz) activity and to have a lasting influence on motor preparation. Specifically, beta (16-25. Hz) power modulation reflected observers' tendency to selectively prepare for a spatially compatible response even before knowing the required one. We discuss the observed frequency-specific and temporally evolving neural activity within a framework of integrated visuomotor processing and point towards possible implications about the mechanisms involved in action observation. |
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. @article{Nieuwenhuijzen2013,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. @article{Broeke2013,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. @article{VanPelt2013,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. |
Daniel Belyusar; Adam C. Snyder; Hans Peter Frey; Mark R. Harwood; Josh Wallman; John J. Foxe Oscillatory alpha-band suppression mechanisms during the rapid attentional shifts required to perform an anti-saccade task Journal Article In: NeuroImage, vol. 65, pp. 395–407, 2013. @article{Belyusar2013,Neuroimaging has demonstrated anatomical overlap between covert and overt attention systems, although behavioral and electrophysiological studies have suggested that the two systems do not rely on entirely identical circuits or mechanisms. In a parallel line of research, topographically-specific modulations of alpha-band power (~. 8-14. Hz) have been consistently correlated with anticipatory states during tasks requiring covert attention shifts. These tasks, however, typically employ cue-target-interval paradigms where attentional processes are examined across relatively protracted periods of time and not at the rapid timescales implicated during overt attention tasks. The anti-saccade task, where one must first covertly attend for a peripheral target, before executing a rapid overt attention shift (i.e. a saccade) to the opposite side of space, is particularly well-suited for examining the rapid dynamics of overt attentional deployments. Here, we asked whether alpha-band oscillatory mechanisms would also be associated with these very rapid overt shifts, potentially representing a common neural mechanism across overt and covert attention systems. High-density electroencephalography in conjunction with infra-red eye-tracking was recorded while participants engaged in both pro- and anti-saccade task blocks. Alpha power, time-locked to saccade onset, showed three distinct phases of significantly lateralized topographic shifts, all occurring within a period of less than 1. s, closely reflecting the temporal dynamics of anti-saccade performance. Only two such phases were observed during the pro-saccade task. These data point to substantially more rapid temporal dynamics of alpha-band suppressive mechanisms than previously established, and implicate oscillatory alpha-band activity as a common mechanism across both overt and covert attentional deployments. |
Harriet R. Brown; Karl J. Friston The functional anatomy of attention: A DCM study Journal Article In: Frontiers in Human Neuroscience, vol. 7, pp. 784, 2013. @article{Brown2013,Recent formulations of attention—in terms of predictive coding—associate attentional gain with the expected precision of sensory information. Formal models of the Posner paradigm suggest that validity effects can be explained in a principled (Bayes optimal) fashion in terms of a cue-dependent setting of precision or gain on the sensory channels reporting anticipated target locations, which is updated selectively by invalid targets. This normative model is equipped with a biologically plausible process theory in the form of predictive coding, where precision is encoded by the gain of superficial pyramidal cells reporting prediction error. We used dynamic causal modeling to assess the evidence in magnetoencephalographic responses for cue-dependent and top-down updating of superficial pyramidal cell gain. Bayesian model comparison suggested that it is almost certain that differences in superficial pyramidal cells gain—and its top-down modulation—contribute to observed responses; and we could be more than 80% certain that anticipatory effects on post-synaptic gain are limited to visual (extrastriate) sources. These empirical results speak to the role of attention in optimizing perceptual inference and its formulation in terms of predictive coding. |
Almudena Capilla; Pascal Belin; Joachim Gross The early spatio-temporal correlates and task independence of cerebral voice processing studied with MEG Journal Article In: Cerebral Cortex, vol. 23, no. 6, pp. 1388–1395, 2013. @article{Capilla2013,Functional magnetic resonance imaging studies have repeatedly provided evidence for temporal voice areas (TVAs) with particular sensitivity to human voices along bilateral mid/anterior superior temporal sulci and superior temporal gyri (STS/STG). In contrast, electrophysiological studies of the spatio-temporal correlates of cerebral voice processing have yielded contradictory results, finding the earliest correlates either at ∼300-400 ms, or earlier at ∼200 ms ("fronto-temporal positivity to voice", FTPV). These contradictory results are likely the consequence of different stimulus sets and attentional demands. Here, we recorded magnetoencephalography activity while participants listened to diverse types of vocal and non-vocal sounds and performed different tasks varying in attentional demands. Our results confirm the existence of an early voice-preferential magnetic response (FTPVm, the magnetic counterpart of the FTPV) peaking at about 220 ms and distinguishing between vocal and non-vocal sounds as early as 150 ms after stimulus onset. The sources underlying the FTPVm were localized along bilateral mid-STS/STG, largely overlapping with the TVAs. The FTPVm was consistently observed across different stimulus subcategories, including speech and non-speech vocal sounds, and across different tasks. These results demonstrate the early, largely automatic recruitment of focal, voice-selective cerebral mechanisms with a time-course comparable to that of face processing. |
Brian A. Coffman; Piyadasa Kodituwakku; Elizabeth L. Kodituwakku; Lucinda Romero; Nirupama Muniswamy Sharadamma; David Stone; Julia M. Stephen Primary visual response (M100) delays in adolescents with FASD as measured with MEG Journal Article In: Human Brain Mapping, vol. 34, no. 11, pp. 2852–2862, 2013. @article{Coffman2013,Fetal alcohol spectrum disorders (FASD) are debilitating, with effects of prenatal alcohol exposure persisting into adolescence and adulthood. Complete characterization of FASD is crucial for the development of diagnostic tools and intervention techniques to decrease the high cost to individual families and society of this disorder. In this experiment, we investigated visual system deficits in adolescents (12-21 years) diagnosed with an FASD by measuring the latency of patients' primary visual M100 responses using MEG. We hypothesized that patients with FASD would demonstrate delayed primary visual responses compared to controls. M100 latencies were assessed both for FASD patients and age-matched healthy controls for stimuli presented at the fovea (central stimulus) and at the periphery (peripheral stimuli; left or right of the central stimulus) in a saccade task requiring participants to direct their attention and gaze to these stimuli. Source modeling was performed on visual responses to the central and peripheral stimuli and the latency of the first prominent peak (M100) in the occipital source timecourse was identified. The peak latency of the M100 responses were delayed in FASD patients for both stimulus types (central and peripheral), but the difference in latency of primary visual responses to central vs. peripheral stimuli was significant only in FASD patients, indicating that, while FASD patients' visual systems are impaired in general, this impairment is more pronounced in the periphery. These results suggest that basic sensory deficits in this population may contribute to sensorimotor integration deficits described previously in this disorder. |
Ido Davidesco; Michal Harel; Michal Ramot; Uri Kramer; Svetlana Kipervasser; Fani Andelman; Miri Y. Neufeld; Gadi Goelman; Itzhak Fried; Rafael Malach Spatial and object-based attention modulates broadband high-frequency responses across the human visual cortical hierarchy Journal Article In: Journal of Neuroscience, vol. 33, no. 3, pp. 1228–1240, 2013. @article{Davidesco2013,One of the puzzling aspects in the visual attention literature is the discrepancy between electrophysiological and fMRI findings: whereas fMRI studies reveal strong attentional modulation in the earliest visual areas, single-unit and local field potential studies yielded mixed results. In addition, it is not clear to what extent spatial attention effects extend from early to high-order visual areas. Here we addressed these issues using electrocorticography recordings in epileptic patients. The patients performed a task that allowed simultaneous manipulation ofboth spatial and object-based attention. They were presented with composite stimuli, consisting ofa small object (face or house) superimposed on a large one, and in separate blocks, were instructed to attend one ofthe objects. We found a consistent increase in broadband high-frequency (30–90Hz) power, but not in visual evoked potentials, associated with spatial attention starting withV1/V2 and continuing throughout the visual hierarchy. The magnitude ofthe attentional modulation was correlated with the spatial selectivity of each electrode and its distance from the occipital pole. Interestingly, the latency of the attentional modulation showed a significant decrease along the visual hierarchy. In addition, electrodes placed over high-order visual areas (e.g., fusiform gyrus) showed both effects of spatial and object-based attention. Overall, our results help to reconcile previous observations of discrepancy between fMRI and electrophysiology. They also imply that spatial attention effects can be found both in early and high-order visual cortical areas, in parallel with their stimulus tuning properties. |
Joao C. Dias; Paul Sajda; J. P. Dmochowski; Lucas C. Parra EEG precursors of detected and missed targets during free-viewing search Journal Article In: Journal of Vision, vol. 13, no. 13, pp. 1–19, 2013. @article{Dias2013,When scanning a scene, the target of our search may be in plain sight and yet remain unperceived. Conversely, at other times the target may be perceived in the periphery prior to fixation. There is ample behavioral and neurophysiological evidence to suggest that in some constrained visual-search tasks, targets are detected prior to fixational eye movements. However, limited human data are available during unconstrained search to determine the time course of detection, the brain areas involved, and the neural correlates of failures to detect a foveated target. Here, we recorded and analyzed electroencephalographic (EEG) activity during free-viewing visual search, varying the task difficulty to compare neural signatures for detected and unreported ("missed") targets. When carefully controlled to remove eye-movement-related potentials, saccade-locked EEG shows that: (a) "Easy" targets may be detected as early as 150 ms prior to foveation, as indicated by a premotor potential associated with a button response; (b) object-discriminating occipital activity emerges during the saccade to target; and (c) success and failures to detect a target are accompanied by a modulation in alpha-band power over fronto-central areas as well as altered saccade dynamics. Taken together, these data suggest that target detection during free viewing can begin prior to and continue during a saccade, with failure or success in reporting a target possibly resulting from inhibition or activation of fronto-central processing areas associated with saccade control. |
Ian C. Fiebelkorn; Adam C. Snyder; Manuel R. Mercier; John S. Butler; S. Molholm; John J. Foxe Cortical cross-frequency coupling predicts perceptual outcomes Journal Article In: NeuroImage, vol. 69, pp. 126–137, 2013. @article{Fiebelkorn2013,Functional networks are comprised of neuronal ensembles bound through synchronization across multiple intrinsic oscillatory frequencies. Various coupled interactions between brain oscillators have been described (e.g., phase-amplitude coupling), but with little evidence that these interactions actually influence perceptual sensitivity. Here, electroencephalographic (EEG) recordings were made during a sustained-attention task to demonstrate that cross-frequency coupling has significant consequences for perceptual outcomes (i.e., whether participants detect a near-threshold visual target). The data reveal that phase-detection relationships at higher frequencies are dependent on the phase of lower frequencies, such that higher frequencies alternate between periods when their phase is either strongly or weakly predictive of visual-target detection. Moreover, the specific higher frequencies and scalp topographies linked to visual-target detection also alternate as a function of lower-frequency phase. Cross-frequency coupling between lower (i.e., delta and theta) and higher frequencies (e.g., low- and high-beta) thus results in dramatic fluctuations of visual-target detection. |
Thomas Fischer; Sven-Thomas Graupner; Boris M. Velichkovsky; Sebastian Pannasch Attentional dynamics during free picture viewing: Evidence from oculomotor behavior and electrocortical activity Journal Article In: Frontiers in Systems Neuroscience, vol. 7, pp. 17, 2013. @article{Fischer2013,Most empirical evidence on attentional control is based on brief presentations of rather abstract stimuli. Results revealed indications for a dynamic interplay between bottom-up and top-down attentional mechanisms. Here we used a more naturalistic task to examine temporal signatures of attentional mechanisms on fine and coarse time scales. Subjects had to inspect digitized copies of 60 paintings, each shown for 40 s. We simultaneously measured oculomotor behavior and electrophysiological correlates of brain activity to compare early and late intervals (1) of inspection time of each picture (picture viewing) and (2) of the full experiment (time on task). For picture viewing, we found an increase in fixation duration and a decrease of saccadic amplitude while these parameters did not change with time on task. Furthermore, early in picture viewing we observed higher spatial and temporal similarity of gaze behavior. Analyzing electrical brain activity revealed changes in three components (C1, N1 and P2) of the eye fixation-related potential (EFRP); during picture viewing; no variation was obtained for the power in the frontal beta- and in the theta activity. Time on task analyses demonstrated no effects on the EFRP amplitudes but an increase of power in the frontal theta and beta band activity. Thus, behavioral and electrophysiological measures similarly show characteristic changes during picture viewing, indicating a shifting balance of its underlying (bottom-up and top-down) attentional mechanisms. Time on task also modulated top-down attention but probably represents a different attentional mechanism. |
Chie Nakatani; Mojtaba Chehelcheraghi; Behnaz Jarrahi; Hironori Nakatani; Cees Leeuwen Cross-frequency phase synchrony around the saccade period as a correlate of perceiver's internal state Journal Article In: Frontiers in Systems Neuroscience, vol. 7, pp. 18, 2013. @article{Nakatani2013a,In active vision, eye-movements depend on perceivers' internal state. We investigated peri-fixation brain activity for internal state-specific tagging. Human participants performed a task, in which a visual object was presented for identification in lateral visual field, to which they moved their eyes as soon as possible from a central fixation point. Next, a phrase appeared in the same location; the phrase could either be an easy or hard question about the object, answered by pressing one of two alternative response buttons, or it could be an instruction to simply press one of these two buttons. Depending on whether these messages were blocked or randomly mixed, one of two different internal states was induced: either the task was known in advance or it wasn't. Eye movements and electroencephalogram (EEG) were recorded simultaneously during task performance. Using eye-event-time-locked averaging and independent component analysis, saccade- and fixation-related components were identified. Coss-frequency phase-synchrony was observed between the alpha/beta1 ranges of fixation-related and beta2/gamma1 ranges of saccade-related activity 50 ms prior to fixation onset in the mixed-phrase condition only. We interpreted this result as evidence for internal state-specific tagging. |
Hironori Nakatani; Cees Leeuwen Antecedent occipital alpha band activity predicts the impact of oculomotor events in perceptual switching Journal Article In: Frontiers in Systems Neuroscience, vol. 7, pp. 19, 2013. @article{Nakatani2013,Oculomotor events such as blinks and saccades transiently interrupt the visual input and, even though this mostly goes undetected, these brief interruptions could still influence the percept. In particular, both blinking and saccades facilitate switching in ambiguous figures such as the Necker cube. To investigate the neural state antecedent to these oculomotor events during the perception of an ambiguous figure, we measured the human scalp electroencephalogram (EEG). When blinking led to perceptual switching, antecedent occipital alpha band activity exhibited a transient increase in amplitude. When a saccade led to switching, a series of transient increases and decreases in amplitude was observed in the antecedent occipital alpha band activity. Our results suggest that the state of occipital alpha band activity predicts the impact of oculomotor events on the percept. |
Andrey R. Nikolaev; Peter Jurica; Chie Nakatani; Gijs Plomp; Cees Leeuwen Visual encoding and fixation target selection in free viewing: Presaccadic brain potentials Journal Article In: Frontiers in Systems Neuroscience, vol. 7, pp. 26, 2013. @article{Nikolaev2013,In scrutinizing a scene, the eyes alternate between fixations and saccades. During a fixation, two component processes can be distinguished: visual encoding and selection of the next fixation target. We aimed to distinguish the neural correlates of these processes in the electrical brain activity prior to a saccade onset. Participants viewed color photographs of natural scenes, in preparation for a change detection task. Then, for each participant and each scene we computed an image heat map, with temperature representing the duration and density of fixations. The temperature difference between the start and end points of saccades was taken as a measure of the expected task-relevance of the information concentrated in specific regions of a scene. Visual encoding was evaluated according to whether subsequent change was correctly detected. Saccades with larger temperature difference were more likely to be followed by correct detection than ones with smaller temperature differences. The amplitude of presaccadic activity over anterior brain areas was larger for correct detection than for detection failure. This difference was observed for short "scrutinizing" but not for long "explorative" saccades, suggesting that presaccadic activity reflects top-down saccade guidance. Thus, successful encoding requires local scanning of scene regions which are expected to be task-relevant. Next, we evaluated fixation target selection. Saccades "moving up" in temperature were preceded by presaccadic activity of higher amplitude than those "moving down". This finding suggests that presaccadic activity reflects attention deployed to the following fixation location. Our findings illustrate how presaccadic activity can elucidate concurrent brain processes related to the immediate goal of planning the next saccade and the larger-scale goal of constructing a robust representation of the visual scene. |
Pavan Ramkumar; Mainak Jas; Sebastian Pannasch; Riitta Hari; Lauri Parkkonen Feature-specific information processing precedes concerted activation in human visual cortex Journal Article In: Journal of Neuroscience, vol. 33, no. 18, pp. 7691–7699, 2013. @article{Ramkumar2013,Current knowledge about the precise timing of visual input to the cortex relies largely on spike timings in monkeys and evoked-response latencies in humans. However, quantifying the activation onset does not unambiguously describe the timing of stimulus-feature-specific information processing. Here, we investigated the information content of the early human visual cortical activity by decoding low-level visual features from single-trial magnetoencephalographic (MEG) responses. MEG was measured from nine healthy subjects as they viewed annular sinusoidal gratings (spanning the visual field from 2 to 10° for a duration of 1 s), characterized by spatial frequency (0.33 cycles/degree or 1.33 cycles/degree) and orientation (45° or 135°); gratings were either static or rotated clockwise or anticlockwise from 0 to 180°. Time-resolved classifiers using a 20 ms moving window exceeded chance level at 51 ms (the later edge of the window) for spatial frequency, 65 ms for orientation, and 98 ms for rotation direction. Decoding accuracies of spatial frequency and orientation peaked at 70 and 90 ms, respectively, coinciding with the peaks of the onset evoked responses. Within-subject time-insensitive pattern classifiers decoded spatial frequency and orientation simultaneously (mean accuracy 64%, chance 25%) and rotation direction (mean 82%, chance 50%). Classifiers trained on data from other subjects decoded the spatial frequency (73%), but not the orientation, nor the rotation direction. Our results indicate that unaveraged brain responses contain decodable information about low-level visual features already at the time of the earliest cortical evoked responses, and that representations of spatial frequency are highly robust across individuals. |
Linsey Roijendijk; Jason Farquhar; Marcel A. J. Gerven; Ole Jensen; Stan Gielen In: PLoS ONE, vol. 8, no. 12, pp. e80489, 2013. @article{Roijendijk2013,OBJECTIVE: Covert visual spatial attention is a relatively new task used in brain computer interfaces (BCIs) and little is known about the characteristics which may affect performance in BCI tasks. We investigated whether eccentricity and task difficulty affect alpha lateralization and BCI performance. APPROACH: We conducted a magnetoencephalography study with 14 participants who performed a covert orientation discrimination task at an easy or difficult stimulus contrast at either a near (3.5°) or far (7°) eccentricity. Task difficulty was manipulated block wise and subjects were aware of the difficulty level of each block. MAIN RESULTS: Grand average analyses revealed a significantly larger hemispheric lateralization of posterior alpha power in the difficult condition than in the easy condition, while surprisingly no difference was found for eccentricity. The difference between task difficulty levels was significant in the interval between 1.85 s and 2.25 s after cue onset and originated from a stronger decrease in the contralateral hemisphere. No significant effect of eccentricity was found. Additionally, single-trial classification analysis revealed a higher classification rate in the difficult (65.9%) than in the easy task condition (61.1%). No effect of eccentricity was found in classification rate. SIGNIFICANCE: Our results indicate that manipulating the difficulty of a task gives rise to variations in alpha lateralization and that using a more difficult task improves covert visual spatial attention BCI performance. The variations in the alpha lateralization could be caused by different factors such as an increased mental effort or a higher visual attentional demand. Further research is necessary to discriminate between them. We did not discover any effect of eccentricity in contrast to results of previous research. |
Jason Satel; Matthew D. Hilchey; Zhiguo Wang; Ross Story; Raymond M. Klein The effects of ignored versus foveated cues upon inhibition of return: An event-related potential study Journal Article In: Attention, Perception, & Psychophysics, vol. 75, no. 1, pp. 29–40, 2013. @article{Satel2013,Taylor and Klein (Journal of Experimental Psychology: Human Perception and Performance 26:1639-1656, 2000) discovered two mutually exclusive "flavors" of inhibition of return (IOR): When the oculomotor system is "actively suppressed," IOR affects input processes (the perception/attention flavor), whereas when the oculomotor system is "engaged," IOR affects output processes (the motor flavor). Studies of brain activity with ignored cues have typically reported that IOR reduces an early sensory event-related potential (ERP) component (i.e., the P1 component) of the brain's response to the target. Since eye movements were discouraged in these experiments, the P1 reduction might be a reflection of the perception/attention flavor of IOR. If, instead of ignoring the cue, participants made a prosaccade to the cue (and then returned to fixation) before responding to the target, the motor flavor of IOR should then be generated. We compared these two conditions while monitoring eye position and recording ERPs to the targets. If the P1 modulation is related to the perceptual/attentional flavor of IOR, we hypothesized that it might be absent when the motoric flavor of IOR was generated by a prosaccade to the cue. Our results demonstrated that target-related P1 reductions and behavioral IOR were similar, and significant, in both conditions. However, P1 modulations were significantly correlated with behavioral IOR only when the oculomotor system was actively suppressed, suggesting that P1 modulations may only affect behaviorally exhibited IOR when the attentional/perceptual flavor of IOR is recruited. |
Steven W. Savage; Douglas D. Potter; Benjamin W. Tatler Does preoccupation impair hazard perception? A simultaneous EEG and eye tracking study Journal Article In: Transportation Research Part F: Traffic Psychology and Behaviour, vol. 17, pp. 52–62, 2013. @article{Savage2013,The aim of this current study was to test the hypothesis that contemplating a recent mobile telephone conversation has a detrimental effect on measures of attentional processing in a driving situation. In this within-subjects design, hazard perception performance was compared between high and no cognitive load conditions (with or without a puzzle to solve). We tested 17 participants, all of whom were required to be in possession of a DVLA approved driving license and had completed the hazard perception portion of the British driving test. A novel dual-task paradigm, which did not require subjects to process or produce verbal information during the primary task, was employed to increase participants' cognitive load. Participants were assessed on three categories of performance measures: behavioural, eye movements and cortical activity between both high and no cognitive load conditions whilst watching 20 clips from a hazard perception test. This study was run in a laboratory of the Psychology Research Wing at the University of Dundee. Behavioural findings from the hazard perception test indicate significantly increased reaction times to hazardous stimuli and significantly increased false alarm rates to non-hazardous stimuli in the high cognitive load condition (when contemplating a previous conversation). Analyses of eye movements indicated significant increases in blink frequencies, higher saccade peak velocities and a significant reduction in the spread of fixations along the horizontal axis. Results from EEG recordings showed a significant increase in frontal and a significant decrease in occipital theta activity within the high load condition. Findings were interpreted within the framework of Corbetta, Patel and Schulmann's (2008) networks model of attention control. Our findings suggest that preoccupation with a recent conversation negatively influences the modulatory effect of the central executive on both the stimulus as well as goal-driven networks of the brain. |
S. E. Bosch; Sebastiaan F. W. Neggers; Stefan Van der Stigchel The role of the frontal eye fields in oculomotor competition: Image-guided TMS enhances contralateral target selection Journal Article In: Cerebral Cortex, vol. 23, no. 4, pp. 824–832, 2013. @article{Bosch2013,In order to execute a correct eye movement to a target in a search display, a saccade program toward the target element must be activated, while saccade programs toward distracting elements must be inhibited. The aim of the present study was to elucidate the role of the frontal eye fields (FEFs) in oculomotor competition. Functional magnetic resonance imaging-guided single-pulse transcranial magnetic stimulation (TMS) was administered over either the left FEF, the right FEF, or the vertex (control site) at 3 time intervals after target presentation, while subjects performed an oculomotor capture task. When TMS was applied over the FEF contralateral to the visual field where a target was presented, there was less interference of an ipsilateral distractor compared with FEF stimulation ipsilateral to the target's visual field or TMS over vertex. Furthermore, TMS over the FEFs decreased latencies of saccades to the contralateral visual field, irrespective of whether the saccade was directed to the target or to the distractor. These findings show that single-pulse TMS over the FEFs enhances the selection of a target in the contralateral visual field and decreases saccade latencies to the contralateral visual field. |
Joost C. Dessing; Michael Vesia; J. Douglas Crawford The role of areas MT+/V5 and SPOC in spatial and temporal control of manual interception: An rTMS study Journal Article In: Frontiers in Behavioral Neuroscience, vol. 7, pp. 15, 2013. @article{Dessing2013,Manual interception, such as catching or hitting an approaching ball, requires the hand to contact a moving object at the right location and at the right time. Many studies have examined the neural mechanisms underlying the spatial aspects of goal-directed reaching, but the neural basis of the spatial and temporal aspects of manual interception are largely unknown. Here, we used repetitive transcranial magnetic stimulation (rTMS) to investigate the role of the human middle temporal visual motion area (MT+/V5) and superior parieto-occipital cortex (SPOC) in the spatial and temporal control of manual interception. Participants were required to reach-to-intercept a downward moving visual target that followed an unpredictably curved trajectory, presented on a screen in the vertical plane. We found that rTMS to MT+/V5 influenced interceptive timing and positioning, whereas rTMS to SPOC only tended to increase the spatial variance in reach end points for selected target trajectories. These findings are consistent with theories arguing that distinct neural mechanisms contribute to spatial, temporal, and spatiotemporal control of manual interception. |
Ada Le; Matthias Niemeier Left visual field preference for a bimanual grasping task with ecologically valid object sizes Journal Article In: Experimental Brain Research, vol. 230, pp. 187–196, 2013. @article{Le2013a,Grasping using two forelimbs in opposition to one another is evolutionary older than the hand with an opposable thumb (Whishaw and Coles in Behav Brain Res 77:135–148, 1996); yet, the mechanisms for bimanual grasps remain unclear. Similar to unimanual grasping, the localization of matching stable grasp points on an object is computationally expensive and so it makes sense for the signals to converge in a single cortical hemisphere. Indeed, bimanual grasps are faster and more accurate in the left visual field, and are disrupted if there is transcra- nial stimulation of the right hemisphere (Le and Niemeier in Exp Brain Res 224:263–273, 2013; Le et al. in Cereb Cortex. doi:10.1093/cercor/bht115, 2013). However, research so far has tested the right hemisphere dominance based on small objects only, which are usually grasped with one hand, whereas bimanual grasping is more com- monly used for objects that are too big for a single hand. Because grasping large objects might involve different neural circuits than grasping small objects (Grol et al. in J Neurosci 27:11877–11887, 2007), here we tested whether a left visual field/right hemisphere dominance for biman- ual grasping exists with large and thus more ecologically valid objects or whether the right hemisphere dominance is a function of object size. We asked participants to fixate to the left or right of an object and to grasp the object with the index and middle fingers of both hands. Consistent with previous observations, we found that for objects in the left visual field, the maximum grip apertures were scaled closer to the object width and were smaller and less variable, than for objects in the right visual field. Our results demonstrate that bimanual grasping is predominantly controlled by the right hemisphere, even in the context of grasping larger objects. |
K. M. Sharika; Sebastiaan F. W. Neggers; Tjerk P. Gutteling; Stefan Van der Stigchel; Hendrik Chris Dijkerman; A. Murthy Proactive control of sequential saccades in the human supplementary eye field Journal Article In: Proceedings of the National Academy of Sciences, vol. 110, no. 14, pp. E1311–E1320, 2013. @article{Sharika2013,Our ability to regulate behavior based on past experience has thus far been examined using single movements. However, natural behavior typically involves a sequence of movements. Here, we examined the effect of previous trial type on the concurrent planning of sequential saccades using a unique paradigm. The task consisted of two trial types: no-shift trials, which implicitly encouraged the concurrent preparation of the second saccade in a subsequent trial; and target-shift trials, which implicitly discouraged the same in the next trial. Using the intersaccadic interval as an index of concurrent planning, we found evidence for context-based preparation of sequential saccades. We also used functional MRI-guided, single-pulse, transcranial magnetic stimulation on human subjects to test the role of the supplementary eye field (SEF) in the proactive control of sequential eye movements. Results showed that (i) stimulating the SEF in the previous trial disrupted the previous trial type-based preparation of the second saccade in the nonstimulated current trial, (ii) stimulating the SEF in the current trial rectified the disruptive effect caused by stimulation in the previous trial, and (iii) stimulating the SEF facilitated the preparation of second saccades based on previous trial type even when the previous trial was not stimulated. Taken together, we show how the human SEF is causally involved in proactive preparation of sequential saccades. |
Aline Frey; Gelu Ionescu; Benoît Lemaire; Francisco López-Orozco; Thierry Baccino; Anne Guérin-Dugué Decision-making in information seeking on texts: an eye-fixation-related potentials investigation Journal Article In: Frontiers in Systems Neuroscience, vol. 7, pp. 39, 2013. @article{Frey2013,Reading on a web page is known to be not linear and people need to make fast decisions about whether they have to stop or not reading. In such context, reading, and decision-making processes are intertwined and this experiment attempts to separate them through electrophysiological patterns provided by the Eye-Fixation-Related Potentials technique (EFRPs). We conducted an experiment in which EFRPs were recorded while participants read blocks of text that were semantically highly related, moderately related, and unrelated to a given goal. Participants had to decide as fast as possible whether the text was related or not to the semantic goal given at a prior stage. Decision making (stopping information search) may occur when the paragraph is highly related to the goal (positive decision) or when it is unrelated to the goal (negative decision). EFRPs were analyzed on and around typical eye fixations: either on words belonging to the goal (target), subjected to a high rate of positive decisions, or on low frequency unrelated words (incongruent), subjected to a high rate of negative decisions. In both cases, we found EFRPs specific patterns (amplitude peaking between 51 to 120 ms after fixation onset) spreading out on the next words following the goal word and the second fixation after an incongruent word, in parietal and occipital areas. We interpreted these results as delayed late components (P3b and N400), reflecting the decision to stop information searching. Indeed, we show a clear spill-over effect showing that the effect on word N spread out on word N + 1 and N + 2. |
Hans Peter Frey; Sophie Molholm; Edmund C. Lalor; Natalie N. Russo; John J. Foxe Atypical cortical representation of peripheral visual space in children with an autism spectrum disorder Journal Article In: European Journal of Neuroscience, vol. 38, no. 1, pp. 2125–2138, 2013. @article{Frey2013a,A key feature of early visual cortical regions is that they contain discretely organized retinotopic maps. Titration of these maps must occur through experience, and the fidelity of their spatial tuning will depend on the consistency and accuracy of the eye movement system. Anomalies in fixation patterns and the ballistics of eye movements are well documented in autism spectrum disorder (ASD), with off-center fixations a hallmark of the phenotype. We hypothesized that these atypicalities might affect the development of visuo-spatial maps and specifically that peripheral inputs might receive altered processing in ASD. Using high-density recordings of visual evoked potentials (VEPs) and a novel system-identification approach known as VESPA (visual evoked spread spectrum analysis), we assessed sensory responses to centrally and peripherally presented stimuli. Additionally, input luminance was varied to bias responsiveness to the magnocellular system, given previous suggestions of magnocellular-specific deficits in ASD. Participants were 22 ASD children (7-17 years of age) and 31 age- and performance-IQ-matched neurotypical controls. Both VEP and VESPA responses to central presentations were indistinguishable between groups. In contrast, peripheral presentations resulted in significantly greater early VEP and VESPA amplitudes in the ASD cohort. We found no evidence that anomalous enhancement was restricted to magnocellular-biased responses. The extent of peripheral response enhancement was related to the severity of stereotyped behaviors and restricted interests, cardinal symptoms of ASD. The current results point to differential visuo-spatial cortical mapping in ASD, shedding light on the consequences of peculiarities in gaze and stereotyped visual behaviors often reported by clinicians working with this population. |
M. Guitart-Masip; G. R. Barnes; A. Horner; Markus Bauer; Raymond J. Dolan; E. Duzel Synchronization of medial temporal lobe and prefrontal rhythms in human decision making Journal Article In: Journal of Neuroscience, vol. 33, no. 2, pp. 442–451, 2013. @article{GuitartMasip2013,Optimal decision making requires that we integrate mnemonic information regarding previous decisions with value signals that entail likely rewards and punishments. The fact that memory and value signals appear to be coded by segregated brain regions, the hippocampus in the case of memory and sectors of prefrontal cortex in the case of value, raises the question as to how they are integrated during human decision making. Using magnetoencephalography to study healthy human participants, we show increased theta oscillations over frontal and temporal sensors during nonspatial decisions based on memories from previous trials. Using source reconstruction we found that the medial temporal lobe (MTL), in a location compatible with the anterior hippocampus, and the anterior cingulate cortex in the medial wall of the frontal lobe are the source of this increased theta power. Moreover, we observed a correlation between theta power in the MTL source and behavioral performance in decision making, supporting a role for MTL theta oscillations in decision-making performance. These MTL theta oscillations were synchronized with several prefrontal sources, including lateral superior frontal gyrus, dorsal anterior cingulate gyrus, and medial frontopolar cortex. There was no relationship between the strength of synchronization and the expected value of choices. Our results indicate a mnemonic guidance of human decision making, beyond anticipation of expected reward, is supported by hippocampal-prefrontal theta synchronization. |
Uwe Hassler; Uwe Friese; Ulla Martens; Nelson Trujillo-Barreto; Thomas Gruber Repetition priming effects dissociate between miniature eye movements and induced gamma-band responses in the human electroencephalogram Journal Article In: European Journal of Neuroscience, vol. 38, no. 3, pp. 2425–2433, 2013. @article{Hassler2013,The role of induced gamma-band responses (iGBRs) in the human electroencephalogram (EEG) is a controversial topic. On the one hand, iGBRs have been associated with neuronal activity reflecting the (re-)activation of cortical object representations. On the other hand, it was shown that miniature saccades (MSs) lead to high-frequency artifacts in the EEG that can mimic cortical iGBRs. We recorded EEG and eye movements simultaneously while participants were engaged in a combined repetition priming and object recognition experiment. MS rates were mainly modulated by object familiarity in a time window from 100 to 300 ms after stimulus onset. In contrast, artifact-corrected iGBRs were sensitive to object repetition and object familiarity in a prolonged time window. EEG source analyses revealed that stimulus repetitions modulated iGBRs in temporal and occipital cortex regions while familiarity was associated with activity in parieto-occipital regions. These results are in line with neuroimaging studies employing functional magnetic resonance imaging or magnetoencephalography. We conclude that MSs reflect early mechanisms of visual perception while iGBRs mirror the activation of cortical networks representing a perceived object. |
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. @article{Henderson2013,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. @article{Hutzler2013,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. |
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. @article{Zerouali2013,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. |
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. @article{Kosilo2013,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. @article{Kretzschmar2013,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. @article{Lao2013,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. @article{Lee2013b,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. |
2012 |
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. @article{MacNamara2012,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. @article{Murphy2012,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. @article{Ploechl2012,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. @article{Raimondo2012,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. @article{Reinhart2012,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. |
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. @article{Vaden2012,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. |
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. @article{Dandekar2012,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. |
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. @article{Xu2012,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. |
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. @article{Davare2012,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. @article{Kamke2012,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. @article{Kamke2012a,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. @article{Kanai2012,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. @article{Kar2012,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. @article{Liang2012,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. |