EyeLink EEG / fNIRS / TMS Publications
All EyeLink EEG, fNIRS, and TMS research publications (with concurrent eye tracking) up until 2023 (with early 2024s) are listed below by year. You can search the publications using keywords such as P300, Gamma band, NIRS, etc. You can also search for individual author names. If we missed any EyeLink EEG, fNIRS, or TMS articles, please email us!
2016 |
Anna-Maria Kasparbauer; Inga Meyhöfer; Maria Steffens; Bernd Weber; Merve Aydin; Veena Kumari; Rene Hurlemann; Ulrich Ettinger Neural effects of methylphenidate and nicotine during smooth pursuit eye movements Journal Article In: NeuroImage, vol. 141, pp. 52–59, 2016. @article{Kasparbauer2016, Introduction: Nicotine and methylphenidate are putative cognitive enhancers in healthy and patient populations. Although they stimulate different neurotransmitter systems, they have been shown to enhance performance on overlapping measures of attention. So far, there has been no direct comparison of the effects of these two stimulants on behavioural performance or brain function in healthy humans. Here, we directly compare the two compounds using a well-established oculomotor biomarker in order to explore common and distinct behavioural and neural effects. Methods: Eighty-two healthy male non-smokers performed a smooth pursuit eye movement task while lying in an fMRI scanner. In a between-subjects, double-blind design, subjects either received placebo (placebo patch and capsule), nicotine (7 mg nicotine patch and placebo capsule), or methylphenidate (placebo patch and 40 mg methylphenidate capsule). Results: There were no significant drug effects on behavioural measures. At the neural level, methylphenidate elicited higher activation in left frontal eye field compared to nicotine, with an intermediate response under placebo. Discussion: The reduced activation of task-related regions under nicotine could be associated with more efficient neural processing, while increased hemodynamic response under methylphenidate is interpretable as enhanced processing of task-relevant networks. Together, these findings suggest dissociable neural effects of these putative cognitive enhancers. |
Tomas Knapen; Jascha D. Swisher; Frank Tong; Patrick Cavanagh Oculomotor remapping of visual information to foveal retinotopic cortex Journal Article In: Frontiers in Systems Neuroscience, vol. 10, pp. 54, 2016. @article{Knapen2016a, Our eyes continually jump around the visual scene to bring the high-resolution, central part of our vision onto objects of interest. We are oblivious to these abrupt shifts, perceiving the visual world to appear reassuringly stable. A process called remapping has been proposed to mediate this perceptual stability for attended objects by shifting their retinotopic representation to compensate for the effects of the upcoming eye movement. In everyday vision, observers make goal-directed eye movements towards items of interest bringing them to the fovea and, for these items, the remapped activity should impinge on foveal regions of the retinotopic maps in visual cortex. Previous research has focused instead on remapping for targets that were not saccade goals, where activity is remapped to a new peripheral location rather than to the foveal representation. We used functional MRI and a phase-encoding design to investigate remapping of spatial patterns of activity towards the fovea/parafovea for saccade targets that were removed prior to completion of the eye movement. We found strong evidence of foveal remapping in retinotopic visual areas, which failed to occur when observers merely attended to the same peripheral target without making eye movements toward it. Significantly, the spatial profile of the remapped response matched the orientation and size of the saccade target, and was appropriately scaled to reflect the retinal extent of the stimulus had it been foveated. We conclude that this remapping of spatially structured information to the fovea may serve as an important mechanism to support our world-centered sense of location across goal-directed eye movements under natural viewing conditions. |
Ulrike Zimmer; M H"ofler; Karl Koschutnig; Anja Ischebeck; Margit Höfler; Karl Koschutnig; Anja Ischebeck Neuronal interactions in areas of spatial attention reflect avoidance of disgust, but orienting to danger Journal Article In: NeuroImage, vol. 134, pp. 94–104, 2016. @article{Zimmer2016, For survival, it is necessary to attend quickly towards dangerous objects, but to turn away from something that is disgusting. We tested whether fear and disgust sounds direct spatial attention differently. Using fMRI, a sound cue (disgust, fear or neutral) was presented to the left or right ear. The cue was followed by a visual target (a small arrow) which was located on the same (valid) or opposite (invalid) side as the cue. Participants were required to decide whether the arrow pointed up- or downwards while ignoring the sound cue. Behaviorally, responses were faster for invalid compared to valid targets when cued by disgust, whereas the opposite pattern was observed for targets after fearful and neutral sound cues. During target presentation, activity in the visual cortex and IPL increased for targets invalidly cued with disgust, but for targets validly cued with fear which indicated a general modulation of activation due to attention. For the TPJ, an interaction in the opposite direction was observed, consistent with its role in detecting targets at unattended positions and in relocating attention. As a whole our results indicate that a disgusting sound directs spatial attention away from its location, in contrast to fearful and neutral sounds. |
Eckart Zimmermann; Ralph Weidner; R. O. Abdollahi; Gereon R. Fink Spatiotopic adaptation in visual areas Journal Article In: Journal of Neuroscience, vol. 36, no. 37, pp. 9526–9534, 2016. @article{Zimmermann2016b, The ability to perceive the visual world around us as spatially stable despite frequent eye movements is one of the long-standing mysteries of neuroscience. The existence of neural mechanisms processing spatiotopic information is indispensable for a successful interaction with the external world. However, how the brain handles spatiotopic information remains a matter of debate. We here combined behavioral and fMRI adaptation to investigate the coding of spatiotopic information in the human brain. Subjects were adapted by a prolonged presentation of a tilted grating. Thereafter, they performed a saccade followed by the brief presentation of a probe. This procedure allowed dissociating adaptation aftereffects at retinal and spatiotopic positions. We found significant behavioral and functional adaptation in both retinal and spatiotopic positions, indicating information transfer into a spatiotopic coordinate system. The brain regions involved were located in ventral visual areas V3, V4, and VO. Our findings suggest that spatiotopic representations involved in maintaining visual stability are constructed by dynamically remapping visual feature information between retinotopic regions within early visual areas. |
Tobias Talanow; Anna-Maria Kasparbauer; Maria Steffens; Inga Meyhöfer; Bernd Weber; Nikolaos Smyrnis; Ulrich Ettinger Facing competition: Neural mechanisms underlying parallel programming of antisaccades and prosaccades Journal Article In: Brain and Cognition, vol. 107, pp. 37–47, 2016. @article{Talanow2016, The antisaccade task is a prominent tool to investigate the response inhibition component of cognitive control. Recent theoretical accounts explain performance in terms of parallel programming of exogenous and endogenous saccades, linked to the horse race metaphor. Previous studies have tested the hypothesis of competing saccade signals at the behavioral level by selectively slowing the programming of endogenous or exogenous processes e.g. by manipulating the probability of antisaccades in an experimental block. To gain a better understanding of inhibitory control processes in parallel saccade programming, we analyzed task-related eye movements and blood oxygenation level dependent (BOLD) responses obtained using functional magnetic resonance imaging (fMRI) at 3T from 16 healthy participants in a mixed antisaccade and prosaccade task. The frequency of antisaccade trials was manipulated across blocks of high (75%) and low (25%) antisaccade frequency. In blocks with high antisaccade frequency, antisaccade latencies were shorter and error rates lower whilst prosaccade latencies were longer and error rates were higher. At the level of BOLD, activations in the task-related saccade network (left inferior parietal lobe, right inferior parietal sulcus, left precentral gyrus reaching into left middle frontal gyrus and inferior frontal junction) and deactivations in components of the default mode network (bilateral temporal cortex, ventromedial prefrontal cortex) compensated increased cognitive control demands. These findings illustrate context dependent mechanisms underlying the coordination of competing decision signals in volitional gaze control. |
Christopher M. Warren; Eran Eldar; Ruud L. Brink; Klodiana-Daphne Tona; Nic J. Wee; Eric J. Giltay; Martijn S. Noorden; Jos A. Bosch; Robert C. Wilson; Jonathan D. Cohen; Sander Nieuwenhuis Catecholamine-mediated increases in gain enhance the precision of cortical representations Journal Article In: Journal of Neuroscience, vol. 36, no. 21, pp. 5699–5708, 2016. @article{Warren2016, Neurophysiological evidence suggests that neuromodulators, such as norepinephrine and dopamine, increase neural gain in target brain areas. Computational models and prominent theoretical frameworks indicate that this should enhance the precision of neural represen- tations, but direct empirical evidence for this hypothesis is lacking. In two functional MRI studies, we examine the effect of baseline catecholamine levels (as indexed by pupil diameter and manipulated pharmacologically) on the precision of object representations in the human ventral temporal cortex using angular dispersion, a powerful, multivariate metric of representational similarity (precision). We first report the results of computational model simulations indicating that increasing catecholaminergic gain should reduce the angular dispersion, and thus increase the precision, of object representations from the same category, as well as reduce the angular dispersion of object representations from distinct categories when distinct-category representations overlap. In Study 1 (N?24), we showthat angular dispersion covaries with pupil diameter, an index of baseline catecholamine levels. In Study 2 (N?24), we manipulate catecholamine levels and neural gain using the norepinephrine transporter blocker atomoxetine and demonstrate consistent, causal effects on angular dispersion and brain-wide functional connectivity. Despite the use of very different methods of examining the effect of baseline catecholamine levels, our results show a striking convergence and demonstrate that catecholamines increase the precision of neural representations. |
Kathleen Thomaes; Iris M. Engelhard; Marit Sijbrandij; Danielle C. Cath; Odile A. Heuvel Degrading traumatic memories with eye movements: A pilot functional MRI study in PTSD Journal Article In: European Journal of Psychotraumatology, vol. 7, no. 1, pp. 1–10, 2016. @article{Thomaes2016, Background: Eye movement desensitization and reprocessing (EMDR) is an effective treatment for post-traumatic stress disorder (PTSD). During EMDR, the patient recalls traumatic memories while making eye movements (EMs). Making EMs during recall is associated with decreased vividness and emotionality of traumatic memories, but the underlying mechanism has been unclear. Recent studies support a ''working-memory'' (WM) theory, which states that the two tasks (recall and EMs) compete for limited capacity of WM resources. However, prior research has mainly relied on self-report measures. Methods: Using functional magnetic resonance imaging, we tested whether ''recall with EMs,''relative to a ''recall-only'' control condition, was associated with reduced activity of primary visual and emotional processing brain regions, associatedwith vividness and emotionality respectively, and increased activity of the dorsolateral prefrontal cortex (DLPFC), associated with working memory. We used a randomized, controlled, crossover experimental design in eight adult patients with a primary diagnosis of PTSD. A script-driven imagery (SDI) procedure was used to measure responsiveness to an audio-script depicting the participant's traumatic memory before and after conditions. Results: SDI activated mainly emotional processing-related brain regions (anterior insula, rostral anterior cingulate cortex (ACC), and dorsomedial prefrontal cortex), WM-related (DLPFC), and visual (association) brain regions before both conditions. Although predicted pre-to post-test decrease in amygdala activation after "recall with EMs" was not significant, SDI activated less right amygdala and rostral ACC activity after "recall with EMs" compared to post-"recall-only." Furthermore, functional connectivity from the right amygdala to the rostral ACC was decreased after "recall with EMs" compared with after "recall-only." Conclusions: These preliminary results in a small sample suggest that making EMs during recall, which is part of the regular EMDR treatment protocol, might reduce activity and connectivity in emotional processing-related areas. This study warrants replication in a larger sample. |
Ruud L. Brink; Thomas Pfeffer; Christopher M. Warren; Peter R. Murphy; Klodiana-Daphne Tona; Nic J. Wee; Eric J. Giltay; Martijn S. Noorden; Serge A. R. B. Rombouts; Tobias H. Donner; Sander Nieuwenhuis Catecholaminergic neuromodulation shapes intrinsic MRI functional connectivity in the human brain Journal Article In: Journal of Neuroscience, vol. 36, no. 30, pp. 7865–7876, 2016. @article{Brink2016, The brain commonly exhibits spontaneous (i.e., in the absence of a task) fluctuations in neural activity that are correlated across brain regions. It has been established that the spatial structure, or topography, of these intrinsic correlations is in part determined by the fixed anatomical connectivity between regions. However, it remains unclear which factors dynamically sculpt this topography as a function of brain state. Potential candidate factors are subcortical catecholaminergic neuromodulatory systems, such as the locus ceruleus-norepinephrine system, which send diffuse projections to most parts of the forebrain. Here, we systematically characterized the effects of endogenous central neuromodulation on correlated fluctuations during rest in the human brain. Using a double-blind placebo-controlled crossover design, we pharmacologically increased synaptic catecholamine levels by administering atomoxetine, an NE transporter blocker, and examined the effects on the strength and spatial structure of resting-state MRI functional connectivity. First, atomoxetine reduced the strength of inter-regional correlations across three levels of spatial organization, indicating that catecholamines reduce the strength of functional interactions during rest. Second, this modulatory effectonintrinsic correlations exhibited a substantial degree of spatial specificity: the decrease in functional connectivity showed an anterior–posterior gradient in the cortex, depended on the strength of baseline functional connectivity, and was strongest for connections between regions belonging to distinct resting-state networks. Thus, catecholamines reduce intrinsic correlations in a spatially heterogeneous fashion. We conclude that neuromodulation is an important factor shaping the topography of intrinsic functional connectivity. |
Jelle A. Dijk; Benjamin Haas; Christina Moutsiana; D. Samuel Schwarzkopf Intersession reliability of population receptive field estimates Journal Article In: NeuroImage, vol. 143, pp. 293–303, 2016. @article{Dijk2016, Population receptive field (pRF) analysis is a popular method to infer spatial selectivity of voxels in visual cortex. However, it remains largely untested how stable pRF estimates are over time. Here we measured the intersession reliability of pRF parameter estimates for the central visual field and near periphery, using a combined wedge and ring stimulus containing natural images. Sixteen healthy human participants completed two scanning sessions separated by 10–114 days. Individual participants showed very similar visual field maps for V1-V4 on both sessions. Intersession reliability for eccentricity and polar angle estimates was close to ceiling for most visual field maps (r>.8 for V1-3). PRF size and cortical magnification (CMF) estimates showed strong but lower overall intersession reliability (r≈.4–.6). Group level results for pRF size and CMF were highly similar between sessions. Additional control experiments confirmed that reliability does not depend on the carrier stimulus used and that reliability for pRF size and CMF is high for sessions acquired on the same day (r>.6). Our results demonstrate that pRF mapping is highly reliable across sessions. |
Anouk Mariette Loon; Johannes J. Fahrenfort; Bauke Velde; Philipp B. Lirk; Nienke C. C. Vulink; Markus W. Hollmann; H. Steven Scholte; Victor A. F. Lamme NMDA receptor antagonist ketamine distorts object recognition by reducing feedback to early visual cortex Journal Article In: Cerebral Cortex, vol. 26, no. 5, pp. 1986–1996, 2016. @article{Loon2016, It is a well-established fact that top-down processes influence neural representations in lower-level visual areas. Electrophysiological recordings in monkeys as well as theoretical models suggest that these top-down processes depend on NMDA receptor functioning. However, this underlying neural mechanism has not been tested in humans. We used fMRI multivoxel pattern analysis to compare the neural representations of ambiguous Mooney images before and after they were recognized with their unambiguous grayscale version. Additionally, we administered ketamine, an NMDA receptor antagonist, to interfere with this process. Our results demonstrate that after recognition, the pattern of brain activation elicited by a Mooney image is more similar to that of its easily recognizable grayscale version than to the pattern evoked by the identical Mooney image before recognition. Moreover, recognition of Mooney images decreased mean response; however, neural representations of separate images became more dissimilar. So from the neural perspective, unrecognizable Mooney images all “look the same”, whereas recognized Mooneys look different. We observed these effects in posterior fusiform part of lateral occipital cortex and in early visual cortex. Ketamine distorted these effects of recognition, but in early visual cortex only. This suggests that top-down processes from higher- to lower-level visual areas might operate via an NMDA pathway. |
Annelinde R. E. Vandenbroucke; Johannes J. Fahrenfort; Julia D. I. Meuwese; H. Steven Scholte; Victor A. F. Lamme Prior knowledge about objects determines neural color representation in human visual cortex Journal Article In: Cerebral Cortex, vol. 26, no. 4, pp. 1401–1408, 2016. @article{Vandenbroucke2016, To create subjective experience, our brain must translate physical stimulus input by incorporating prior knowledge and expectations. For example, we perceive color and not wavelength information, and this in part depends on our past experience with colored objects ( Hansen et al. 2006; Mitterer and de Ruiter 2008). Here, we investigated the influence of object knowledge on the neural substrates underlying subjective color vision. In a functional magnetic resonance imaging experiment, human subjects viewed a color that lay midway between red and green (ambiguous with respect to its distance from red and green) presented on either typical red (e.g., tomato), typical green (e.g., clover), or semantically meaningless (nonsense) objects. Using decoding techniques, we could predict whether subjects viewed the ambiguous color on typical red or typical green objects based on the neural response of veridical red and green. This shift of neural response for the ambiguous color did not occur for nonsense objects. The modulation of neural responses was observed in visual areas (V3, V4, VO1, lateral occipital complex) involved in color and object processing, as well as frontal areas. This demonstrates that object memory influences wavelength information relatively early in the human visual system to produce subjective color vision. |
Renée M. Visser; Michelle I. C. Haan; Tinka Beemsterboer; Pia Haver; Merel Kindt; H. Steven Scholte Quantifying learning-dependent changes in the brain: Single-trial multivoxel pattern analysis requires slow event-related fMRI Journal Article In: Psychophysiology, vol. 53, no. 8, pp. 1117–1127, 2016. @article{Visser2016, Single-trial analysis is particularly useful for assessing cognitive processes that are intrinsically dynamic, such as learning. Studying these processes with fMRI is problematic, as the low signal-to-noise ratio of fMRI requires the averaging over multiple trials, obscuring trial-by-trial changes in neural activation. The superior sensitivity of multivoxel pattern analysis over univariate analyses has opened up new possibilities for single-trial analysis, but this may require different fMRI designs. Here, we measured fMRI and pupil dilation responses during discriminant aversive conditioning, to assess associative learning in a trial-by-trial manner. The impact of design choices was examined by varying trial spacing and trial order in a series of five experiments (total n = 66), while keeping stimulus duration constant (4.5 s). Our outcome measure was the change in similarity between neural response patterns related to two consecutive presentations of the same stimulus (within-stimulus) and between patterns related to pairs of different stimuli (between-stimulus) that shared a specific outcome (electric stimulation vs. no consequence). This trial-by-trial similarity analysis revealed clear single-trial learning curves in conditions with intermediate (8.1-12.6 s) and long (16.5-18.4 s) intervals, with effects being strongest in designs with long intervals and counterbalanced stimulus presentation. No learning curves were observed in designs with shorter intervals (1.6-6.1 s), indicating that rapid event-related designs-at present, the most common designs in fMRI research-are not suited for single-trial pattern analysis. These findings emphasize the importance of deciding on the type of analysis prior to data collection. |
Wuyi Wang; Shivakumar Viswanathan; Taraz Lee; Scott T. Grafton In: PLoS ONE, vol. 11, no. 7, pp. e0158465, 2016. @article{Wang2016g, Cortical theta band oscillations (4-8 Hz) in EEG signals have been shown to be important for a variety of different cognitive control operations in visual attention paradigms. However the synchronization source of these signals as defined by fMRI BOLD activity and the extent to which theta oscillations play a role in multimodal attention remains unknown. Here we investigated the extent to which cross-modal visual and auditory attention impacts theta oscillations. Using a simultaneous EEG-fMRI paradigm, healthy human participants performed an attentional vigilance task with six cross-modal conditions using naturalistic stimuli. To assess supramodal mechanisms, modulation of theta oscillation amplitude for attention to either visual or auditory stimuli was correlated with BOLD activity by conjunction analysis. Negative correlation was localized to cortical regions associated with the default mode network and positively with ventral premotor areas. Modality-associated attention to visual stimuli was marked by a positive correlation of theta and BOLD activity in fronto-parietal area that was not observed in the auditory condition. A positive correlation of theta and BOLD activity was observed in auditory cortex, while a negative correlation of theta and BOLD activity was observed in visual cortex during auditory attention. The data support a supramodal interaction of theta activity with of DMN function, and modality-associated processes within fronto-parietal networks related to top-down theta related cognitive control in cross-modal visual attention. On the other hand, in sensory cortices there are opposing effects of theta activity during cross-modal auditory attention. |
Zheng Zhiwei Wang; Kristina Zeljic; Qinying Jiang; Yong Gu; Wei Wang; Zheng Wang Dynamic network communication in the human functional connectome predicts perceptual variability in visual illusion Journal Article In: Cerebral Cortex, vol. 28, no. 1, pp. 1–15, 2016. @article{Wang2016h, The eukaryotic RNA exosome is an essential, multi-subunit complex that catalyzes RNA turnover, maturation, and quality control processes. Its non-catalytic donut-shaped core includes 9 subunits that associate with the 3' to 5' exoribonucleases Rrp6, and Rrp44/Dis3, a subunit that also catalyzes endoribonuclease activity. Although recent structures and biochemical studies of RNA bound exosomes from S. cerevisiae revealed that the Exo9 central channel guides RNA to either Rrp6 or Rrp44 using partially overlapping and mutually exclusive paths, several issues related to RNA recruitment remain. Here, we identify activities for the highly basic Rrp6 C-terminal tail that we term the 'lasso' because it binds RNA and stimulates ribonuclease activities associated with Rrp44 and Rrp6 within the 11-subunit nuclear exosome. Stimulation is dependent on the Exo9 central channel, and the lasso contributes to degradation and processing activities of exosome substrates in vitro and in vivo. Finally, we present evidence that the Rrp6 lasso may be a conserved feature of the eukaryotic RNA exosome. |
Meytal Wilf; Michal Ramot; Edna Furman-Haran; Anat Arzi; Yechiel Levkovitz; Rafael Malach Diminished auditory responses during NREM sleep correlate with the hierarchy of language processing Journal Article In: PLoS ONE, vol. 11, no. 6, pp. e0157143, 2016. @article{Wilf2016, Natural sleep provides a powerful model system for studying the neuronal correlates of awareness and state changes in the human brain. To quantitatively map the nature of sleep-induced modulations in sensory responses we presented participants with auditory stimuli possessing different levels of linguistic complexity. Ten participants were scanned using functional magnetic resonance imaging (fMRI) during the waking state and after falling asleep. Sleep staging was based on heart rate measures validated independently on 20 participants using concurrent EEG and heart rate measurements and the results were confirmed using permutation analysis. Participants were exposed to three types of auditory stimuli: scrambled sounds, meaningless word sentences and comprehensible sentences. During non-rapid eye movement (NREM) sleep, we found diminishing brain activation along the hierarchy of language processing, more pronounced in higher processing regions. Specifically, the auditory thalamus showed similar activation levels during sleep and waking states, primary auditory cortex remained activated but showed a significant reduction in auditory responses during sleep, and the high order language-related representation in inferior frontal gyrus (IFG) cortex showed a complete abolishment of responses during NREM sleep. In addition to an overall activation decrease in language processing regions in superior temporal gyrus and IFG, those areas manifested a loss of semantic selectivity during NREM sleep. Our results suggest that the decreased awareness to linguistic auditory stimuli during NREM sleep is linked to diminished activity in high order processing stations. |
2015 |
Zvi N. Roth; Ehud Zohary Position and identity information available in fMRI patterns of activity in human visual cortex Journal Article In: Journal of Neuroscience, vol. 35, no. 33, pp. 11559–11571, 2015. @article{Roth2015, Parietal cortex is often implicated in visual processing of actions. Action understanding is essentially abstract, specific to the type or goal of action, but greatly independent of variations in the perceived position of the action. If certain parietal regions are involved in action understanding, then we expect them to show these generalization and selectivity properties. However, additional functions of parietal cortex, such as self-action control, may impose other demands by requiring an accurate representation of the location of graspable objects. Therefore, the dimensions along which responses are modulated may indicate the functional role of specific parietal regions. Here, we studied the degree of position invariance and hand/object specificity during viewing of tool-grasping actions. To that end, we characterize the information available about location, hand, and tool identity in the patterns of fMRI activation in various cortical areas: early visual cortex, posterior intraparietal sulcus, anterior superior parietal lobule, and the ventral object-specific lateral occipital complex. Our results suggest a gradient within the human dorsal stream: along the posterior-anterior axis, position information is gradually lost, whereas hand and tool identity information is enhanced. This may reflect a gradual transformation of visual input from an initial retinotopic representation in early visual areas to an abstract, position-invariant representation of viewed action in anterior parietal cortex. |
Frieder M. Paulus; Sören Krach; Marius Blanke; Christine Roth; Marcus Belke; Jens Sommer; Laura Müller-Pinzler; Katja Menzler; Andreas Jansen; Felix Rosenow; Frank Bremmer; Wolfgang Einhäuser; Susanne Knake Fronto-insula network activity explains emotional dysfunctions in juvenile myoclonic epilepsy: Combined evidence from pupillometry and fMRI Journal Article In: Cortex, vol. 65, pp. 219–231, 2015. @article{Paulus2015, Emotional instability, difficulties in social adjustment, and disinhibited behavior are the most common symptoms of the psychiatric comorbidities in juvenile myoclonic epilepsy (JME). This psychopathology has been associated with dysfunctions of mesial-frontal brain circuits. The present work is a first direct test of this link and adapted a paradigm for probing frontal circuits during empathy for pain. Neural and psychophysiological parameters of pain empathy were assessed by combining functional magnetic resonance imaging (fMRI) with simultaneous pupillometry in 15 JME patients and 15 matched healthy controls. In JME patients, we observed reduced neural activation of the anterior cingulate cortex (ACC), the anterior insula (AI), and the ventrolateral prefrontal cortex (VLPFC). This modulation was paralleled by reduced pupil dilation during empathy for pain in patients. At the same time, pupil dilation was positively related to neural activity of the ACC, AI, and VLPFC. In JME patients, the ACC additionally showed reduced functional connectivity with the primary and secondary somatosensory cortex, areas fundamentally implicated in processing the somatic cause of another's pain. Our results provide first evidence that alterations of mesial-frontal circuits directly affect psychosocial functioning in JME patients and draw a link of pupil dynamics with brain activity during emotional processing. The findings of reduced pain empathy related activation of the ACC and AI and aberrant functional integration of the ACC with somatosensory cortex areas provide further evidence for this network's role in social behavior and helps explaining the JME psychopathology and patients' difficulties in social adjustment. |
Benjamin Peters; Jochen Kaiser; Benjamin Rahm; Christoph Bledowski Activity in human visual and parietal cortex reveals object-based attention in working memory Journal Article In: Journal of Neuroscience, vol. 35, no. 8, pp. 3360–3369, 2015. @article{Peters2015, Visual attention enables observers to select behaviorally relevant information based on spatial locations, features, or objects. Attentional selection is not limited to physically present visual information, but can also operate on internal representations maintained in working memory (WM) in service of higher-order cognition. However, only little is known about whether attention to WM contents follows the same principles as attention to sensory stimuli. To address this question, we investigated in humans whether the typically observed effects of object-based attention in perception are also evident for object-based attentional selection of internal object representations in WM. In full accordance with effects in visual perception, the key behavioral and neuronal characteristics of object-based attention were observed in WM. Specifically, we found that reaction times were shorter when shifting attention to memory positions located on the currently attended object compared with equidistant positions on a different object. Furthermore, functional magnetic resonance imaging and multivariate pattern analysis of visuotopic activity in visual (areas V1-V4) and parietal cortex revealed that directing attention to one position of an object held in WM also enhanced brain activation for other positions on the same object, suggesting that attentional selection in WM activates the entire object. This study demonstrated that all characteristic features of object-based attention are present in WM and thus follows the same principles as in perception. |
Golbarg T. Saber; Franco Pestilli; Clayton E. Curtis Saccade planning evokes topographically specific activity in the dorsal and ventral streams Journal Article In: Journal of Neuroscience, vol. 35, no. 1, pp. 245–252, 2015. @article{Saber2015, Saccade planning may invoke spatially-specific feedback signals that bias early visual activity in favor of top-down goals. We tested this hypothesis by measuring cortical activity at the early stages of the dorsal and ventral visual processing streams. Human subjects maintained saccade plans to (prosaccade) or away (antisaccade) from a spatial location over long memory-delays. Results show that cortical activity persists in early visual cortex at the retinotopic location of upcoming saccade goals. Topographically specific activity persists as early as V1, and activity increases along both dorsal (V3A/B, IPS0) and ventral (hV4, VO1) visual areas. Importantly, activity persists when saccade goals are available only via working memory and when visual targets and saccade goals are spatially disassociated. We conclude that top-down signals elicit retinotopically specific activity in visual cortex both in the dorsal and ventral streams. Such activity may underlie mechanisms that prioritize locations of task-relevant objects. |
Sarah Schuster; Stefan Hawelka; Fabio Richlan; Philipp Ludersdorfer; Florian Hutzler Eyes on words: A fixation-related fMRI study of the left occipito-temporal cortex during self-paced silent reading of words and pseudowords Journal Article In: Scientific Reports, vol. 5, pp. 12686, 2015. @article{Schuster2015, The predominant finding of studies assessing the response of the left ventral occipito-temporal cortex (vOT) to familiar words and to unfamiliar, but pronounceable letter strings (pseudowords) is higher activation for pseudowords. One explanation for this finding is that readers automatically generate predictions about a letter string's identity – pseudowords mismatch these predictions and the higher vOT activation is interpreted as reflecting the resultant prediction errors. The majority of studies, however, administered tasks which imposed demands above and beyond the intrinsic requirements of visual word recognition. The present study assessed the response of the left vOT to words and pseudowords by using the onset of the first fixation on a stimulus as time point for modeling the BOLD signal (fixation-related fMRI). This method allowed us to assess the neural correlates of self-paced silent reading with minimal task demands and natural exposure durations. In contrast to the predominantly reported higher vOT activation for pseudowords, we found higher activation for words. This finding is at odds with the expectation of higher vOT activation for pseudowords due to automatically generated predictions and the accompanying elevation of prediction errors. Our finding conforms to an alternative explanation which considers such top-down processing to be non-automatic and task-dependent. |
Renée M. Visser; Anna E. Kunze; Bianca Westhoff; H. Steven Scholte; Merel Kindt Representational similarity analysis offers a preview of the noradrenergic modulation of long-term fear memory at the time of encoding Journal Article In: Psychoneuroendocrinology, vol. 55, pp. 8–20, 2015. @article{Visser2015, Neuroimaging research on emotional memory has greatly advanced our understanding of the pathogenesis of anxiety disorders. While the behavioral expression of fear at the time of encoding does not predict whether an aversive experience will evolve into long-term fear memory, the application of multi-voxel pattern analysis (MVPA) for the analysis of BOLD-MRI data has recently provided a unique marker for memory formation. Here, we aimed to further investigate the utility of this marker by modulating the strength of fear memory with an α2-adrenoceptor antagonist (yohimbine HCl). Fifty-two healthy participants were randomly assigned to two conditions - either receiving 20. mg yohimbine or a placebo pill (double-blind) - prior to differential fear conditioning and MRI-scanning. We examined the strength of fear associations during acquisition and retention of fear (48. h later) by assessing the similarity of BOLD-MRI patterns and pupil dilation responses. Additionally, participants returned for a follow-up test outside the scanner (2-4 weeks), during which we assessed fear-potentiated startle responses. Replicating our previous findings, neural pattern similarity reflected the development of fear associations over time, and unlike average activation or pupil dilation, predicted the later expression of fear memory (pupil dilation 48. h later). While no effect of yohimbine was observed on markers of autonomic arousal, including salivary α-amylase (sAA), we obtained indirect evidence for the noradrenergic enhancement of fear memory consolidation: sAA levels showed a strong increase prior to fMRI scanning, irrespective of whether participants had received yohimbine, and this increase correlated with the subsequent expression of fear (48. h later). Remarkably, this noradrenergic enhancement of fear was associated with changes in neural response patterns at the time of learning. These findings provide further evidence that representational similarity analysis is a sensitive tool for studying (enhanced) memory formation. |
Simone Vossel; Christoph Mathys; Klaas E. Stephan; Karl J. Friston Cortical coupling reflects Bayesian belief updating in the deployment of spatial attention Journal Article In: Journal of Neuroscience, vol. 35, no. 33, pp. 11532–11542, 2015. @article{Vossel2015, The deployment of visuospatial attention and the programming of saccades are governed by the inferred likelihood of events. In the present study, we combined computational modeling of psychophysical data with fMRI to characterize the computational and neural mechanisms underlying this flexible attentional control. Sixteen healthy human subjects performed a modified version of Posner's location-cueing paradigm in which the percentage of cue validity varied in time and the targets required saccadic responses. Trialwise estimates of the certainty (precision) of the prediction that the target would appear at the cued location were derived from a hierarchical Bayesian model fitted to individual trialwise saccadic response speeds. Trial-specific model parameters then entered analyses of fMRI data as parametric regressors. Moreover, dynamic causal modeling (DCM) was performed to identify the most likely functional architecture of the attentional reorienting network and its modulation by (Bayes-optimal) precision-dependent attention. While the frontal eye fields (FEFs), intraparietal sulcus, and temporoparietal junction (TPJ) of both hemispheres showed higher activity on invalid relative to valid trials, reorienting responses in right FEF, TPJ, and the putamen were significantly modulated by precision-dependent attention. Our DCM results suggested that the precision of predictability underlies the attentional modulation of the coupling of TPJ with FEF and the putamen. Our results shed new light on the computational architecture and neuronal network dynamics underlying the context-sensitive deployment of visuospatial attention.$backslash$n$backslash$nSIGNIFICANCE STATEMENT: Spatial attention and its neural correlates in the human brain have been studied extensively with the help of fMRI and cueing paradigms in which the location of targets is pre-cued on a trial-by-trial basis. One aspect that has so far been neglected concerns the question of how the brain forms attentional expectancies when no a priori probability information is available but needs to be inferred from observations. This study elucidates the computational and neural mechanisms under which probabilistic inference governs attentional deployment. Our results show that Bayesian belief updating explains changes in cortical connectivity; in that directional influences from the temporoparietal junction on the frontal eye fields and the putamen were modulated by (Bayes-optimal) updates. |
Dov Yellin; Aviva Berkovich-Ohana; Rafael Malach Coupling between pupil fluctuations and resting-state fMRI uncovers a slow build-up of antagonistic responses in the human cortex Journal Article In: NeuroImage, vol. 106, pp. 414–427, 2015. @article{Yellin2015, Even in absence of overt tasks, the human cortex manifests rich patterns of spontaneous "resting state" BOLD-fMRI fluctuations. However, the link of these spontaneous fluctuations to behavior is presently unclear. Attempts to directly investigate this link invariably lead to disruptions of the resting state. Here we took advantage of the well-established association between pupil diameter and attentional gain to address this issue by examining the correlation between the resting state BOLD and pupil fluctuations. Our results uncover a spontaneously emerging spatiotemporal pupil-BOLD correlation whereby a slow buildup of activity in default mode areas preceded both pupil dilation and wide-spread BOLD suppression in sensorimotor cortex. Control experiments excluded a role for luminance fluctuations or fixation. Comparing the pupil-correlated patterns to activation maps during visual imagery revealed a substantial overlap. Our results indicate a link between behavior, as indexed by pupil diameter, and resting state BOLD fluctuations. These pupil dilations, assumed to be related to attentional gain, were associated with spontaneously emerging antagonism between fundamental cortical networks. |
Ruben S. Van Bergen; Wei Ji Ma; Michael S. Pratte; Janneke F. M. Jehee Sensory uncertainty decoded from visual cortex predicts behavior Ruben Journal Article In: Nature Neuroscience, vol. 18, no. 12, pp. 1728–1730, 2015. @article{VanBergen2015, Bayesian theories of neural coding propose that sensory uncertainty is represented by a probability distribution encoded in neural population activity, but direct neural evidence supporting this hypothesis is currently lacking. Using fMRI in combination with a generative model-based analysis, we found that probability distributions reflecting sensory uncertainty could reliably be estimated from human visual cortex and, moreover, that observers appeared to use knowledge of this uncertainty in their perceptual decisions. |
J. J. Tramper; W. Pieter Medendorp Parallel updating and weighting of multiple spatial maps for visual stability during whole body motion Journal Article In: Journal of Neurophysiology, vol. 114, no. 6, pp. 3211–3219, 2015. @article{Tramper2015, It is known that the brain uses multiple reference frames to code spatial information, including eye-centered and body-centered frames. When we move our body in space, these internal representations are no longer in register with external space, unless they are actively updated. Whether the brain updates multiple spatial representations in parallel, or whether it restricts its updating mechanisms to a single reference frame from which other representations are constructed, remains an open question. We developed an optimal integration model to simulate the updating of visual space across body motion in multiple or single reference frames. To test this model, we designed an experiment in which participants had to remember the location of a briefly presented target while being translated sideways. The behavioral responses were in agreement with a model that uses a combination of eye- and body-centered representations, weighted according to the reliability in which the target location is stored, and updated in each reference frame. Our findings suggest that the brain simultaneously updates multiple spatial representations across body motion. Because both representations are kept in sync, they can be optimally combined to provide a more precise estimate of visual locations in space than based on single-frame updating mechanisms. |
Inga Meyhöfer; Maria Steffens; Anna-Maria Kasparbauer; Phillip Grant; Bernd Weber; Ulrich Ettinger Neural mechanisms of smooth pursuit eye movements in schizotypy Journal Article In: Human Brain Mapping, vol. 36, pp. 340–353, 2015. @article{Meyhoefer2015, Patients with schizophrenia as well as individuals with high levels of schizotypy are known to have deficits in smooth pursuit eye movements (SPEM). Here, we investigated, for the first time, the neural mechanisms underlying SPEM performance in high schizotypy. Thirty-one healthy participants [N = 19 low schizotypes |
Elizabeth Michael; Vincent De Gardelle; Alejo Nevado-Holgado; Christopher Summerfield Unreliable evidence: 2 sources of uncertainty during perceptual choice Journal Article In: Cerebral Cortex, vol. 25, no. 4, pp. 937–947, 2015. @article{Michael2015, Perceptual decisions often involve integrating evidence from multiple concurrently available sources. Uncertainty arises when the integrated (mean) evidence fails to support one alternative over another. However, evidence heterogeneity (variability) also provokes uncertainty. Here, we asked whether these 2 sources of uncertainty have independent behavioral and neural effects during choice. Human observers undergoing functional neuroimaging judged the average color or shape of a multielement array. The mean and variance of the feature values exerted independent influences on behavior and brain activity. Surprisingly, BOLD signals in the dorsomedial prefrontal cortex (dmPFC) showed polar opposite responses to the 2 sources of uncertainty, with the strongest response to ambiguous tallies of evidence (high mean uncertainty) and to homogenous arrays (low variance uncertainty). These findings present a challenge for models that emphasize the role of the dmPFC in detecting conflict, errors, or surprise. We suggest an alternative explanation, whereby evidence is processed with increased gain near the category boundary. |
L. Müller-Pinzler; V. Gazzola; C. Keysers; Jens Sommer; Andreas Jansen; S. Frässle; Wolfgang Einhäuser; Frieder M. Paulus; Sören Krach Neural pathways of embarrassment and their modulation by social anxiety Journal Article In: NeuroImage, vol. 119, pp. 252–261, 2015. @article{MuellerPinzler2015, While being in the center of attention and exposed to other's evaluations humans are prone to experience embarrassment. To characterize the neural underpinnings of such aversive moments, we induced genuine experiences of embarrassment during person-group interactions in a functional neuroimaging study. Using a mock-up scenario with three confederates, we examined how the presence of an audience affected physiological and neural responses and the reported emotional experiences of failures and achievements. The results indicated that publicity induced activations in mentalizing areas and failures led to activations in arousal processing systems. Mentalizing activity as well as attention towards the audience were increased in socially anxious participants. The converging integration of information from mentalizing areas and arousal processing systems within the ventral anterior insula and amygdala forms the neural pathways of embarrassment. Targeting these neural markers of embarrassment in the (para-)limbic system provides new perspectives for developing treatment strategies for social anxiety disorders. |
Sara Ajina; Christopher Kennard; Geraint Rees; Holly Bridge Motion area V5/MT+ response to global motion in the absence of V1 resembles early visual cortex Journal Article In: Brain, vol. 138, no. 1, pp. 164–178, 2015. @article{Ajina2015, Motion area V5/MT+ shows a variety of characteristic visual responses, often linked to perception, which are heavily influenced by its rich connectivity with the primary visual cortex (V1). This human motion area also receives a number of inputs from other visual regions, including direct subcortical connections and callosal connections with the contralateral hemisphere. Little is currently known about such alternative inputs to V5/MT+ and how they may drive and influence its activity. Using functional magnetic resonance imaging, the response of human V5/MT+ to increasing the proportion of coherent motion was measured in seven patients with unilateral V1 damage acquired during adulthood, and a group of healthy age-matched controls. When V1 was damaged, the typical V5/MT+ response to increasing coherence was lost. Rather, V5/MT+ in patients showed a negative trend with coherence that was similar to coherence-related activity in V1 of healthy control subjects. This shift to a response-pattern more typical of early visual cortex suggests that in the absence of V1, V5/MT+ activity may be shaped by similar direct subcortical input. This is likely to reflect intact residual pathways rather than a change in connectivity, and has important implications for blindsight function. It also confirms predictions that V1 is critically involved in normal V5/MT+ global motion processing, consistent with a convergent model of V1 input to V5/MT+. Historically, most attempts to model cortical visual responses do not consider the contribution of direct subcortical inputs that may bypass striate cortex, such as input to V5/MT+. We have shown that the signal change driven by these non-striate pathways can be measured, and suggest that models of the intact visual system may benefit from considering their contribution. |
Sara Ajina; Geraint Rees; Christopher Kennard; Holly Bridge Abnormal contrast responses in the extrastriate cortex of blindsight patients Journal Article In: Journal of Neuroscience, vol. 35, no. 21, pp. 8201–8213, 2015. @article{Ajina2015b, When the human primary visual cortex (V1) is damaged, the dominant geniculo-striate pathway can no longer convey visual information to the occipital cortex. However, many patients with such damage retain some residual visual function that must rely on an alternative pathway directly to extrastriate occipital regions. This residual vision is most robust for moving stimuli, suggesting a role for motion area hMT+. However, residual vision also requires high-contrast stimuli, which is inconsistent with hMT+ sensitivity to contrast in which even low-contrast levels elicit near-maximal neural activation. We sought to investigate this discrepancy by measuring behavioral and neural responses to increasing contrast in patients with V1 damage. Eight patients underwent behavioral testing and functional magnetic resonance imaging to record contrast sensitivity in hMT+ of their damaged hemisphere, using Gabor stimuli with a spatial frequency of 1 cycle/degrees. The responses from hMT+ of the blind hemisphere were compared with hMT+ and V1 responses in the sighted hemisphere of patients and a group of age-matched controls. Unlike hMT+, neural responses in V1 tend to increase linearly with increasing contrast, likely reflecting a dominant parvocellular channel input. Across all patients, the responses in hMT+ of the blind hemisphere no longer showed early saturation but increased linearly with contrast. Given the spatiotemporal parameters used in this study and the known direct subcortical projections from the koniocellular layers of the lateral geniculate nucleus to hMT+, we propose that this altered contrast sensitivity in hMT+ could be consistent with input from the koniocellular pathway. |
Jamila Andoh; Reiko Matsushita; Robert J. Zatorre Asymmetric interhemispheric transfer in the auditory network: Evidence from TMS, resting-state fMRI, and diffusion imaging Journal Article In: Journal of Neuroscience, vol. 43, no. 43, pp. 14602–14611, 2015. @article{Andoh2015, Hemispheric asymmetries in human auditory cortical function and structure are still highly debated. Brain stimulation approaches can complement correlational techniques by uncovering causal influences. Previous studies have shown asymmetrical effects of transcranial magnetic stimulation (TMS) on task performance, but it is unclear whether these effects are task-specific or reflect intrinsic network properties. To test how modulation of auditory cortex (AC) influences functional networks and whether this influence is asymmetrical, the present study measured resting-state fMRI connectivity networks in 17 healthy volunteers before and immediately after TMS (continuous theta burst stimulation) to the left or right AC, and the vertex as a control. We also examined the relationship between TMS-induced interhemispheric signal propagation and anatomical properties of callosal auditory fibers as measured with diffusion-weighted MRI. We found that TMS to the right AC, but not the left, resulted in widespread connectivity decreases in auditory- and motor-related networks in the resting state. Individual differences in the degree of change in functional connectivity between auditory cortices after TMS applied over the right AC were negatively related to the volume of callosal auditory fibers. The findings show that TMS-induced network modulation occurs, even in the absence of an explicit task, and that the magnitude of the effect differs across individuals as a function of callosal structure, supporting a role for the corpus callosum in mediating functional asymmetry. The findings support theoretical models emphasizing hemispheric differences in network organization and are of practical significance in showing that brain stimulation studies need to take network-level effects into account. |
Pinglei Bao; Christopher J. Purington; Bosco S. Tjan Using an achiasmic human visual system to quantify the relationship between the fMRI BOLD signal and neural response Journal Article In: eLife, vol. 4, no. NOVEMBER2015, pp. 1–21, 2015. @article{Bao2015, Achiasma in humans causes gross mis-wiring of the retinal-fugal projection, resulting in overlapped cortical representations of left and right visual hemifields. We show that in areas V1-V3 this overlap is due to two co-located but non-interacting populations of neurons, each with a receptive field serving only one hemifield. Importantly, the two populations share the same local vascular control, resulting in a unique organization useful for quantifying the relationship between neural and fMRI BOLD responses without direct measurement of neural activity. Specifically, we can non-invasively double local neural responses by stimulating both neuronal populations with identical stimuli presented symmetrically across the vertical meridian to both visual hemifields, versus one population by stimulating in one hemifield. Measurements from a series of such doubling experiments show that the amplitude of BOLD response is proportional to approximately 0.5 power of the underlying neural response. Reanalyzing published data shows that this inferred relationship is general. |
Jan Brascamp; Randolph Blake; Tomas Knapen Negligible fronto-parietal BOLD activity accompanying unreportable switches in bistable perception Journal Article In: Nature Neuroscience, vol. 18, no. 11, pp. 1672–1678, 2015. @article{Brascamp2015, The human brain's executive systems have a vital role in deciding and selecting among actions. Selection among alternatives also occurs in the perceptual domain; for instance, when perception switches between interpretations during perceptual bistability. Whether executive systems also underlie this functionality remains debated, with known fronto-parietal concomitants of perceptual switches being variously interpreted as reflecting the switches' cause or as reflecting their consequences. We developed a procedure in which the two eyes receive different inputs and perception demonstrably switches between these inputs, yet the switches themselves are so inconspicuous as to become unreportable, minimizing their executive consequences. Fronto-parietal fMRI BOLD responses that accompanied perceptual switches were similarly minimized in this procedure, indicating that these reflect the switches' consequences rather than their cause. We conclude that perceptual switches do not always rely on executive brain areas and that processes responsible for selection among alternatives may operate outside the brain's executive systems. |
Korhan Buyukturkoglu; Hans Roettgers; Jens Sommer; Mohit Rana; Leonie Dietzsch; Ezgi Belkis Arikan; Ralf Veit; Rahim Malekshahi; Tilo Kircher; Niels Birbaumer; Ranganatha Sitaram; Sergio Ruiz Self-regulation of anterior insula using real-time fMRI and its behavioral effects in obsessive compulsive disorder: A feasibility study Journal Article In: PLoS ONE, vol. 10, no. 8, pp. e0135872, 2015. @article{Buyukturkoglu2015, Introduction: Obsessive-compulsive disorder (OCD) is a common and chronic condition that can have disabling effects throughout the patient's lifespan. Frequent symptoms among OCD patients include fear of contamination and washing compulsions. Several studies have shown a link between contamination fears, disgust over-reactivity, and insula activation in OCD. In concordance with the role of insula in disgust processing, new neural models based on neuroimaging studies suggest that abnormally high activations of insula could be implicated in OCD psychopathology, at least in the subgroup of patients with contamination fears and washing compulsions. Methods: In the current study, we used a Brain Computer Interface (BCI) based on real-time func- tional magnetic resonance imaging (rtfMRI) to aid OCD patients to achieve down-regula- tion of the Blood Oxygenation Level Dependent (BOLD) signal in anterior insula. Our first aim was to investigate whether patients with contamination obsessions and washing com- pulsions can learn to volitionally decrease (down-regulate) activity in the insula in the pres- ence of disgust/anxiety provoking stimuli. Our second aimwas to evaluate the effect of down-regulation on clinical, behavioural and physiological changes pertaining to OCD symptoms. Hence, several pre- and post-training measures were performed, i.e., con- fronting the patient with a disgust/anxiety inducing real-world object (Ecological Disgust Test), and subjective rating and physiological responses (heart rate, skin conductance level) of disgust towards provoking pictures. Results: Results of this pilot study, performed in 3 patients (2 females), show that OCD patients can gain self-control of the BOLD activity of insula, albeit to different degrees. In two patients positive changes in behaviour in the EDT were observed following the rtfMRI trainings. Behavioural changes were also confirmed by reductions in the negative valence and in the subjective perception of disgust towards symptom provoking images. Conclusion: Although preliminary, results of this study confirmed that insula down-regulation is possible in patients suffering from OCD, and that volitional decreases of insula activation could be used for symptom alleviation in this disorder. |
Nathan Caruana; Jon Brock; Alexandra Woolgar A frontotemporoparietal network common to initiating and responding to joint attention bids Journal Article In: NeuroImage, vol. 108, pp. 34–46, 2015. @article{Caruana2015, Joint attention is a fundamental cognitive ability that supports daily interpersonal relationships and communication. The Parallel Distributed Processing model (PDPM) postulates that responding to (RJA) and initiating (IJA) joint attention are predominantly supported by posterior-parietal and frontal regions respectively. It also argues that these neural networks integrate during development, supporting the parallel processes of self- and other-attention representation during interactions. However, direct evidence for the PDPM is limited due to a lack of ecologically valid experimental paradigms that can capture both RJA and IJA. Building on existing interactive approaches, we developed a virtual reality paradigm where participants engaged in an online interaction to complete a cooperative task. By including tightly controlled baseline conditions to remove activity associated with non-social task demands, we were able to directly contrast the neural correlates of RJA and IJA to determine whether these processes are supported by common brain regions. Both RJA and IJA activated broad frontotemporoparietal networks. Critically, a conjunction analysis identified that a subset of these regions were common to both RJA and IJA. This right-lateralised network included the dorsal portion of the middle frontal gyrus (MFG), inferior frontal gyrus (IFG), middle temporal gyrus (MTG), precentral gyrus, posterior superior temporal sulcus (pSTS), temporoparietal junction (TPJ) and precuneus. Additional activation was observed in this network for IJA relative to RJA at MFG, IFG, TPJ and precuneus. This is the first imaging study to directly investigate the neural correlates common to RJA and IJA engagement, and thus support the assumption that a broad integrated network underlies the parallel aspects of both initiating and responding to joint attention. |
Wonil Choi; John M. Henderson Neural correlates of active vision: An fMRI comparison of natural reading and scene viewing Journal Article In: Neuropsychologia, vol. 75, pp. 109–118, 2015. @article{Choi2015, Theories of eye movement control during active vision tasks such as reading and scene viewing have primarily been developed and tested using data from eye tracking and computational modeling, and little is currently known about the neurocognition of active vision. The current fMRI study was conducted to examine the nature of the cortical networks that are associated with active vision. Subjects were asked to read passages for meaning and view photographs of scenes for a later memory test. The eye movement control network comprising frontal eye field (FEF), supplementary eye fields (SEF), and intraparietal sulcus (IPS), commonly activated during single-saccade eye movement tasks, were also involved in reading and scene viewing, suggesting that a common control network is engaged when eye movements are executed. However, the activated locus of the FEF varied across the two tasks, with medial FEF more activated in scene viewing relative to passage reading and lateral FEF more activated in reading than scene viewing. The results suggest that eye movements during active vision are associated with both domain-general and domain-specific components of the eye movement control network. |
S. Clavagnier; Serge O. Dumoulin; R. F. Hess Is the cortical deficit in amblyopia due to reduced cortical magnification, loss of neural resolution, or neural disorganization? Journal Article In: Journal of Neuroscience, vol. 35, no. 44, pp. 14740–14755, 2015. @article{Clavagnier2015, The neural basis of amblyopia is a matter of debate. The following possibilities have been suggested: loss of foveal cells, reduced cortical magnification, loss of spatial resolution of foveal cells, and topographical disarray in the cellular map. To resolve this we undertook a population receptive field (pRF) functional magnetic resonance imaging analysis in the central field in humans with moderate-to-severe amblyopia. We measured the relationship between averaged pRF size and retinal eccentricity in retinotopic visual areas. Results showed that cortical magnification is normal in the foveal field of strabismic amblyopes. However, the pRF sizes are enlarged for the amblyopic eye. We speculate that the pRF enlargement reflects loss of cellular resolution or an increased cellular positional disarray within the representation of the amblyopic eye. |
Anouk J. Brouwer; Jeroen B. J. Smeets; Tjerk P. Gutteling; I. Toni; W. Pieter Medendorp The Müller-Lyer illusion affects visuomotor updating in the dorsal visual stream Journal Article In: Neuropsychologia, vol. 77, pp. 119–127, 2015. @article{Brouwer2015, To guide our actions, the brain has developed mechanisms to code target locations in egocentric coordinates (i.e., with respect to the observer), and to update these when the observer moves. The latter mechanism, called visuomotor updating, is implemented in the dorsal visual stream of the brain. In contrast, the ventral visual stream is assumed to transform target locations into an allocentric reference frame that is highly sensitive to visual contextual illusions. Here, we tested the effect of the Müller-Lyer illusion on visuomotor updating in a double-step saccade task. Using the same paradigm in a 3T fMRI scanner, we investigated the effect of the illusion on the neural correlate of the updating process. Participants briefly viewed the Brentano version of the Müller-Lyer illusion with a target at its middle vertex, while fixating at one of the two endpoints of the illusion. Shortly after the disappearance of the stimulus, the eyes' fixation point moved to a position outside the illusion. After a delay, participants made a saccade to the remembered position of the target. The landing position of this saccade was systematically displaced in a manner congruent with the perceptual illusion, showing that visuomotor updating is affected by the illusion. fMRI results showed that the BOLD response in the occipito-parietal cortex (area V7) and the intraparietal sulcus related to planning of the saccade to the updated target was also modulated by the configuration of the illusion. This suggests that the dorsal visual stream represents perceived rather than physical locations of remembered saccade targets. |
Abdurahman S. Elkhetali; Ryan J. Vaden; Sean M. Pool; Kristina M. Visscher Early visual cortex reflects initiation and maintenance of task set Journal Article In: NeuroImage, vol. 107, pp. 277–288, 2015. @article{Elkhetali2015, The human brain is able to process information flexibly, depending on a person's task. The mechanisms underlying this ability to initiate and maintain a task set are not well understood, but they are important for understanding the flexibility of human behavior and developing therapies for disorders involving attention. Here we investigate the differential roles of early visual cortical areas in initiating and maintaining a task set.Using functional Magnetic Resonance Imaging (fMRI), we characterized three different components of task set-related, but trial-independent activity in retinotopically mapped areas of early visual cortex, while human participants performed attention demanding visual or auditory tasks. These trial-independent effects reflected: (1) maintenance of attention over a long duration, (2) orienting to a cue, and (3) initiation of a task set. Participants performed tasks that differed in the modality of stimulus to be attended (auditory or visual) and in whether there was a simultaneous distractor (auditory only, visual only, or simultaneous auditory and visual). We found that patterns of trial-independent activity in early visual areas (V1, V2, V3, hV4) depend on attended modality, but not on stimuli. Further, different early visual areas play distinct roles in the initiation of a task set. In addition, activity associated with maintaining a task set tracks with a participant's behavior. These results show that trial-independent activity in early visual cortex reflects initiation and maintenance of a person's task set. |
Joseph C. Griffis; Abdurahman S. Elkhetali; Ryan J. Vaden; Kristina M. Visscher Distinct effects of trial-driven and task set-related control in primary visual cortex Journal Article In: NeuroImage, vol. 120, pp. 285–297, 2015. @article{Griffis2015, Task sets are task-specific configurations of cognitive processes that facilitate task-appropriate reactions to stimuli. While it is established that the trial-by-trial deployment of visual attention to expected stimuli influences neural responses in primary visual cortex (V1) in a retinotopically specific manner, it is not clear whether the mechanisms that help maintain a task set over many trials also operate with similar retinotopic specificity. Here, we address this question by using BOLD fMRI to characterize how portions of V1 that are specialized for different eccentricities respond during distinct components of an attention-demanding discrimination task: cue-driven preparation for a trial, trial-driven processing, task-initiation at the beginning of a block of trials, and task-maintenance throughout a block of trials. Tasks required either unimodal attention to an auditory or a visual stimulus or selective intermodal attention to the visual or auditory component of simultaneously presented visual and auditory stimuli. We found that while the retinotopic patterns of trial-driven and cue-driven activity depended on the attended stimulus, the retinotopic patterns of task-initiation and task-maintenance activity did not. Further, only the retinotopic patterns of trial-driven activity were found to depend on the presence of inter-modal distraction. Participants who performed well on the intermodal selective attention tasks showed strong task-specific modulations of both trial-driven and task-maintenance activity. Importantly, task-related modulations of trial-driven and task-maintenance activity were in opposite directions. Together, these results confirm that there are (at least) two different processes for top-down control of V1: One, working trial-by-trial, differently modulates activity across different eccentricity sectors - portions of V1 corresponding to different visual eccentricities. The second process works across longer epochs of task performance, and does not differ among eccentricity sectors. These results are discussed in the context of previous literature examining top-down control of visual cortical areas. |
Marcus Grueschow; Rafael Polania; Todd A. Hare; Christian C. Ruff Automatic versus choice-dependent value representations in the human brain Journal Article In: Neuron, vol. 85, no. 4, pp. 874–885, 2015. @article{Grueschow2015, The subjective values of choice options can impact on behavior in two fundamentally different types of situations: first, when people explicitly base their actions on such values, and second, when values attract attention despite being irrelevant for current behavior. Here we show with functional magnetic resonance imaging (fMRI) that these two behavioral functions of values are encoded in distinct regions of the human brain. In the medial prefrontal cortex, value-related activity is enhanced when subjective value becomes choice-relevant, and the magnitude of this increase relates directly to the outcome and reliability of the value-based choice. In contrast, activity in the posterior cingulate cortex represents values similarly when they are relevant or irrelevant for the present choice, and the strength of this representation predicts attentional capture by choice-irrelevant values. Our results suggest that distinct components of the brain's valuation network encode value in context-dependent manners that serve fundamentally different behavioral aims. |
Hanna Heikkinen; Fariba Sharifian; Ricardo Vigario; Simo Vanni Feedback to distal dendrites links fMRI signals to neural receptive fields in a spiking network model of the visual cortex Journal Article In: Journal of Neurophysiology, vol. 114, no. 1, pp. 57–69, 2015. @article{Heikkinen2015, The blood oxygenation level-dependent (BOLD) response has been strongly associated with neuronal activity in the brain. However, some neuronal tuning properties are consistently different from the BOLD response. We studied the spatial extent of neural and hemodynamic responses in the primary visual cortex, where the BOLD responses spread and interact over much longer distances than the small receptive fields of individual neurons would predict. Our model shows that a feedforward-feedback loop between V1 and a higher visual area can account for the observed spread of the BOLD response. In particular, anisotropic landing of inputs to compartmental neurons were necessary to account for the BOLD signal spread, while retaining realistic spiking responses. Our work shows that simple dendrites can separate tuning at the synapses and at the action potential output, thus bridging the BOLD signal to the neural receptive fields with high fidelity. |
John M. Henderson; Wonil Choi Neural correlates of fixation duration during real-world scene viewing: Evidence from fixation-related (FIRE) fMRI Journal Article In: Journal of Cognitive Neuroscience, vol. 27, no. 6, pp. 1137–1145, 2015. @article{Henderson2015, During active scene perception, our eyes move from one location to another via saccadic eye movements, with the eyes fixating objects and scene elements for varying amounts of time. Much of the variability in fixation duration is accounted for by attentional, perceptual, and cognitive processes associated with scene analysis and comprehension. For this reason, current theories of active scene viewing attempt to account for the influence of attention and cognition on fixation duration. Yet almost nothing is known about the neurocognitive systems associated with variation in fixation duration during scene viewing. We addressed this topic using fixation-related fMRI, which involves coregistering high-resolution eye tracking and magnetic resonance scanning to conduct event-related fMRI analysis based on characteristics of eye movements. We observed that activation in visual and prefrontal executive control areas was positively correlated with fixation duration, whereas activation in ventral areas associated with scene en- coding and medial superior frontal and paracentral regions associated with changing action plans was negatively correlated with fixation duration. The results suggest that fixation duration in scene viewing is controlled by cognitive processes associated with real-time scene analysis interacting with motor planning, consistent with current computational models of active vision for scene perception. |
John M. Henderson; Wonil Choi; Steven G. Luke; Rutvik H. Desai Neural correlates of fixation duration in natural reading: Evidence from fixation-related fMRI Journal Article In: NeuroImage, vol. 119, pp. 390–397, 2015. @article{Henderson2015a, A key assumption of current theories of natural reading is that fixation duration reflects underlying attentional, language, and cognitive processes associated with text comprehension. The neurocognitive correlates of this relationship are currently unknown. To investigate this relationship, we compared neural activation associated with fixation duration in passage reading and a pseudo-reading control condition. The results showed that fixation duration was associated with activation in oculomotor and language areas during text reading. Fixation duration during pseudo-reading, on the other hand, showed greater involvement of frontal control regions, suggesting flexibility and task dependency of the eye movement network. Consistent with current models, these results provide support for the hypothesis that fixation duration in reading reflects attentional engagement and language processing. The results also demonstrate that fixation-related fMRI provides a method for investigating the neurocognitive bases of natural reading. |
Sharna D. Jamadar; Beth P. Johnson; Meaghan Clough; Gary F. Egan; Joanne Fielding Behavioral and neural plasticity of ocular motor control: Changes in performance and fMRI activity following antisaccade training Journal Article In: Frontiers in Human Neuroscience, vol. 9, no. 653, pp. 1–13, 2015. @article{Jamadar2015, The antisaccade task provides a model paradigm that sets the inhibition of a reflexively driven behavior against the volitional control of a goal-directed behavior. The stability and adaptability of antisaccade performance was investigated in 23 neurologically healthy individuals. Behavior and brain function were measured using functional magnetic resonance imaging (fMRI) prior to and immediately following 2 weeks of daily antisaccade training. Participants performed antisaccade trials faster with no change in directional error rate following 2 weeks of training; however this increased speed came at the cost of the spatial accuracy of the saccade (gain) which became more hypometric following training. Training on the antisaccade task resulted in increases in fMRI activity in the fronto-basal ganglia-parietal-cerebellar ocular motor network. Following training, antisaccade latency was positively associated with fMRI activity in the frontal and supplementary eye fields, anterior cingulate and intraparietal sulcus; antisaccade gain was negatively associated with fMRI activity in supplementary eye fields, anterior cingulate, intraparietal sulcus, and cerebellar vermis. In sum, the results suggest that following training, larger antisaccade latency is associated with larger activity in fronto-parietal-cerebellar ocular motor regions, and smaller antisaccade gain is associated with larger activity in fronto-parietal ocular motor regions. |
2014 |
Rasmus Aamand; Yi-Ching Lynn Ho; Thomas Dalsgaard; Andreas Roepstorff; Torben E. Lund Dietary nitrate facilitates an acetazolamide-induced increase in cerebral blood flow during visual stimulation Journal Article In: Journal of Applied Physiology, vol. 116, no. 3, pp. 267–273, 2014. @article{Aamand2014, The carbonic anhydrase (CA) inhibitor acetazolamide (AZ) is used routinely to estimate cerebrovascular reserve capacity in patients, as it reliably increases cerebral blood flow (CBF). However, the mechanism by which AZ accomplishes this CBF increase is not entirely understood. We recently discovered that CA can produce nitric oxide (NO) from nitrite, and that AZ enhances this NO production in vitro. In fact, this interaction between AZ and CA accounted for a large part of AZ's vasodilatory action, which fits well with the known vasodilatory potency of NO. The present study aimed to assess whether AZ acts similarly in vivo in the human cerebrovascular system. Hence, we increased or minimized the dietary intake of nitrate in 20 healthy male participants, showed them a full-field flickering dartboard, and measured their CBF response to this visual stimulus with arterial spin labeling. Doing so, we found a significant positive interaction between the dietary intake of nitrate and the CBF modulation afforded by AZ during visual stimulation. In addition, but contrary to studies conducted in elderly participants, we report no effect of nitrate intake on resting CBF in healthy human participants. The present study provides in vivo support for an enhancing effect of AZ on the NO production from nitrite catalyzed by CA in the cerebrovascular system. Furthermore, our results, in combination with the results of other groups, indicate that nitrate may have significant importance to vascular function when the cerebrovascular system is challenged by age or disease. |
Julia D. I. Meuwese; H. Steven Scholte; Victor A. F. Lamme Latent memory of unattended stimuli reactivated by practice: An fMRI study on the role of consciousness and attention in learning Journal Article In: PLoS ONE, vol. 9, no. 3, pp. e90098, 2014. @article{Meuwese2014, Although we can only report about what is in the focus of our attention, much more than that is actually processed. And even when attended, stimuli may not always be reportable, for instance when they are masked. A stimulus can thus be unreportable for different reasons: the absence of attention or the absence of a conscious percept. But to what extent does the brain learn from exposure to these unreportable stimuli? In this fMRI experiment subjects were exposed to textured figure-ground stimuli, of which reportability was manipulated either by masking (which only interferes with consciousness) or with an inattention paradigm (which only interferes with attention). One day later learning was assessed neurally and behaviorally. Positive neural learning effects were found for stimuli presented in the inattention paradigm; for attended yet masked stimuli negative adaptation effects were found. Interestingly, these inattentional learning effects only became apparent in a second session after a behavioral detection task had been administered during which performance feedback was provided. This suggests that the memory trace that is formed during inattention is latent until reactivated by behavioral practice. However, no behavioral learning effects were found, therefore we cannot conclude that perceptual learning has taken place for these unattended stimuli. |
R. Chris Miall; Se-Ho Nam; J. Tchalenko The influence of stimulus format on drawing-a functional imaging study of decision making in portrait drawing Journal Article In: NeuroImage, vol. 102, pp. 608–619, 2014. @article{Miall2014, To copy a natural visual image as a line drawing, visual identification and extraction of features in the image must be guided by top-down decisions, and is usually influenced by prior knowledge. In parallel with other behavioral studies testing the relationship between eye and hand movements when drawing, we report here a functional brain imaging study in which we compared drawing of faces and abstract objects: the former can be strongly guided by prior knowledge, the latter less so. To manipulate the difficulty in extracting features to be drawn, each original image was presented in four formats including high contrast line drawings and silhouettes, and as high and low contrast photographic images. We confirmed the detailed eye-hand interaction measures reported in our other behavioral studies by using in-scanner eye-tracking and recording of pen movements with a touch screen. We also show that the brain activation pattern reflects the changes in presentation formats. In particular, by identifying the ventral and lateral occipital areas that were more highly activated during drawing of faces than abstract objects, we found a systematic increase in differential activation for the face-drawing condition, as the presentation format made the decisions more challenging. This study therefore supports theoretical models of how prior knowledge may influence perception in untrained participants, and lead to experience-driven perceptual modulation by trained artists. |
D. A. Barany; V. Della-Maggiore; Shivakumar Viswanathan; M. Cieslak; Scott T. Grafton Feature Iinteractions enable decoding of sensorimotor transformations for goal-directed movement Journal Article In: Journal of Neuroscience, vol. 34, no. 20, pp. 6860–6873, 2014. @article{Barany2014, Neurophysiology and neuroimaging evidence shows that the brain represents multiple environmental and body-related features to compute transformations from sensory input to motor output. However, it is unclear how these features interact during goal-directed movement. To investigate this issue, we examined the representations of sensory and motor features of human hand movements within the left-hemisphere motor network. In a rapid event-related fMRI design, we measured cortical activity as participants performed right-handed movements at the wrist, with either of two postures and two amplitudes, to move a cursor to targets at different locations. Using a multivoxel analysis technique with rigorous generalization tests, we reliably distinguished representations of task-related features (primarily target location, movement direction, and posture) in multiple regions. In particular, we identified an interaction between target location and movement direction in the superior parietal lobule, which may underlie a transformation from the location of the target in space to a movement vector. In addition, we found an influence of posture on primary motor, premotor, and parietal regions. Together, these results reveal the complex interactions between different sensory and motor features that drive the computation of sensorimotor transformations. |
D. J. Bridge; Joel L. Voss Hippocampal binding of novel information with dominant memory traces can support both memory stability and change Journal Article In: Journal of Neuroscience, vol. 34, no. 6, pp. 2203–2213, 2014. @article{Bridge2014, Memory stability and change are considered opposite outcomes. We tested the counterintuitive notion that both depend on one process: hippocampal binding of memory features to associatively novel information, or associative novelty binding (ANB). Building on the idea that dominant memory features, or “traces,” are most susceptible to modification, we hypothesized that ANB would selectively involve dominant traces. Therefore, memory stability versus change should depend on whether the currently dominant trace is old versus updated; in either case, novel information will be bound with it, causing either maintenance (when old) or change (when updated). People in our experiment studied objects at locations within scenes (contexts). During reactivation in a new context, subjects moved studied objects to new locations either via active location recall or by passively dragging objects to predetermined locations. After active reactivation, the new object location became dominant in memory, whereas after passive reactivation, the old object location maintained dominance. In both cases, hippocampal ANB bound the currently dominant object-location memory with a context with which it was not paired previously (i.e., associatively novel). Stability occurred in the passive condition when ANB united the dominant original location trace with an associatively novel newer context. Change occurred in the active condition when ANB united the dominant updated object location with an associatively novel and older context. Hippocampal ANB of the currently dominant trace with associatively novel contextual information thus provides a single mechanism to support memory stability and change, with shifts in trace dominance during reactivation dictating the outcome. |
Wonil Choi; Rutvik H. Desai; John M. Henderson The neural substrates of natural reading: A comparison of normal and nonword text using eyetracking and fMRI Journal Article In: Frontiers in Human Neuroscience, vol. 8, pp. 1024, 2014. @article{Choi2014c, Most previous studies investigating the neural correlates of reading have presented text using serial visual presentation (SVP), which may not fully reflect the underlying processes of natural reading. In the present study, eye movements and BOLD data were collected while subjects either read normal paragraphs naturally or moved their eyes through "paragraphs" of pseudo-text (pronounceable pseudowords or consonant letter strings) in two pseudo-reading conditions. Eye movement data established that subjects were reading and scanning the stimuli normally. A conjunction fMRI analysis across natural- and pseudo-reading showed that a common eye-movement network including frontal eye fields (FEF), supplementary eye fields (SEF), and intraparietal sulci was activated, consistent with previous studies using simpler eye movement tasks. In addition, natural reading versus pseudo-reading showed different patterns of brain activation: normal reading produced activation in a well-established language network that included superior temporal gyrus/sulcus, middle temporal gyrus (MTG), angular gyrus (AG), inferior frontal gyrus, and middle frontal gyrus, whereas pseudo-reading produced activation in an attentional network that included anterior/posterior cingulate and parietal cortex. These results are consistent with results found in previous single-saccade eye movement tasks and SVP reading studies, suggesting that component processes of eye-movement control and language processing observed in past fMRI research generalize to natural reading. The results also suggest that combining eyetracking and fMRI is a suitable method for investigating the component processes of natural reading in fMRI research. |
H. X. Wang; Elisha P. Merriam; Jeremy Freeman; David J. Heeger Motion direction biases and decoding in human visual cortex Journal Article In: Journal of Neuroscience, vol. 34, no. 37, pp. 12601–12615, 2014. @article{Wang2014h, Functional magnetic resonance imaging (fMRI) studies have relied on multivariate analysis methods to decode visual motion direction from measurements of cortical activity. Above-chance decoding has been commonly used to infer the motion-selective response properties of the underlying neural populations. Moreover, patterns of reliable response biases across voxels that underlie decoding have been interpreted to reflect maps of functional architecture. Using fMRI, we identified a direction-selective response bias in human visual cortex that: (1) predicted motion-decoding accuracy; (2) depended on the shape of the stimulus aperture rather than the absolute direction of motion, such that response amplitudes gradually decreased with distance from the stimulus aperture edge corresponding to motion origin; and 3) was present in V1, V2, V3, but not evident in MT+, explaining the higher motion-decoding accuracies reported previously in early visual cortex. These results demonstrate that fMRI-based motion decoding has little or no dependence on the underlying functional organization of motion selectivity. |
Michiel Elk; Michiel Elk The left inferior parietal lobe represents stored hand-postures for object use and action prediction Journal Article In: Frontiers in Psychology, vol. 5, pp. 333, 2014. @article{Elk2014, Action semantics enables us to plan actions with objects and to predict others' object-directed actions as well. Previous studies have suggested that action semantics are represented in a fronto-parietal action network that has also been implicated to play a role in action observation. In the present fMRI study it was investigated how activity within this network changes as a function of the predictability of an action involving multiple objects and requiring the use of action semantics. Participants performed an action prediction task in which they were required to anticipate the use of a centrally presented object that could be moved to an associated target object (e.g., hammer-nail). The availability of actor information (i.e., presenting a hand grasping the central object) and the number of possible target objects (i.e., 0, 1, or 2 target objects) were independently manipulated, resulting in different levels of predictability. It was found that making an action prediction based on actor information resulted in an increased activation in the extrastriate body area (EBA) and the fronto-parietal action observation network (AON). Predicting actions involving a target object resulted in increased activation in the bilateral IPL and frontal motor areas. Within the AON, activity in the left inferior parietal lobe (IPL) and the left premotor cortex (PMC) increased as a function of the level of action predictability. Together these findings suggest that the left IPL represents stored hand-postures that can be used for planning object-directed actions and for predicting other's actions as well. |
Juha M. Lahnakoski; Enrico Glerean; Iiro P. Jääskeläinen; Jukka Hyönä; Riitta Hari; Mikko Sams; Lauri Nummenmaa Synchronous brain activity across individuals underlies shared psychological perspectives Journal Article In: NeuroImage, vol. 100, pp. 316–324, 2014. @article{Lahnakoski2014, For successful communication, we need to understand the external world consistently with others. This task requires sufficiently similar cognitive schemas or psychological perspectives that act as filters to guide the selection, interpretation and storage of sensory information, perceptual objects and events. Here we show that when individuals adopt a similar psychological perspective during natural viewing, their brain activity becomes synchronized in specific brain regions. We measured brain activity with functional magnetic resonance imaging (fMRI) from 33 healthy participants who viewed a 10-min movie twice, assuming once a 'social' (detective) and once a 'non-social' (interior decorator) perspective to the movie events. Pearson's correlation coefficient was used to derive multisubject voxelwise similarity measures (inter-subject correlations; ISCs) of functional MRI data. We used k-nearest-neighbor and support vector machine classifiers as well as a Mantel test on the ISC matrices to reveal brain areas wherein ISC predicted the participants' current perspective. ISC was stronger in several brain regions-most robustly in the parahippocampal gyrus, posterior parietal cortex and lateral occipital cortex-when the participants viewed the movie with similar rather than different perspectives. Synchronization was not explained by differences in visual sampling of the movies, as estimated by eye gaze. We propose that synchronous brain activity across individuals adopting similar psychological perspectives could be an important neural mechanism supporting shared understanding of the environment. |
Rebecca P. Lawson; Ben Seymour; Eleanor Loh; Antoine Lutti; Raymond J. Dolan; Peter Dayan; Nikolaus Weiskopf; Jonathan P. Roiser The habenula encodes negative motivational value associated with primary punishment in humans Journal Article In: Proceedings of the National Academy of Sciences, vol. 111, no. 32, pp. 11858–11863, 2014. @article{Lawson2014, Learning what to approach, and what to avoid, involves assigning value to environmental cues that predict positive and negative events. Studies in animals indicate that the lateral habenula encodes the previously learned negative motivational value of stimuli. However, involvement of the habenula in dynamic trial-by-trial aversive learning has not been assessed, and the functional role of this structure in humans remains poorly characterized, in part, due to its small size. Using high-resolution functional neuroimaging and computational modeling of reinforcement learning, we demonstrate positive habenula responses to the dynamically changing values of cues signaling painful electric shocks, which predict behavioral suppression of responses to those cues across individuals. By contrast, negative habenula responses to monetary reward cue values predict behavioral invigoration. Our findings show that the habenula plays a key role in an online aversive learning system and in generating associated motivated behavior in humans. |
Katerina Lukasova; Jens Sommer; Mariana P. Nucci-Da-Silva; Gilson Vieira; Marius Blanke; Frank Bremmer; João R. Sato; Tilo Kircher; Edson Amaro Test-retest reliability of fMRI activation generated by different saccade tasks Journal Article In: Journal of Magnetic Resonance Imaging, vol. 40, no. 1, pp. 37–46, 2014. @article{Lukasova2014, PURPOSE: To assess the reproducibility of brain-activation and eye-movement patterns in a saccade paradigm when comparing subjects, tasks, and magnetic resonance (MR) systems. MATERIALS AND METHODS: Forty-five healthy adults at two different sites (n = 45) performed saccade tasks with varying levels of target predictability: predictable (PRED), position predictable (pPRED), time predictable (tPRED), and prosaccade (SAC). Eye-movement pattern was tested with a repeated-measures analysis of variance. Activation maps reproducibility were estimated with the cluster overlap Jaccard index and signal variance coefficient of determination for within-subjects test-retest data, and for between-subjects data from the same and different sites. RESULTS: In all groups latencies increased with decreasing target predictability: PRED < pPRED < tPRED < SAC (P < 0,001). Activation overlap was good to fair (>0.40) in all tasks in the within-subjects test-retest comparisons and poor (<0.40) in the tPRED for different subjects. The overlap of the different tasks for within-groups data was higher (0.40-0.68) than for the between-groups data (0.30-0.50). Activation consistency was 60-85% in the same subjects, 50-79% in different subjects, and 50-80% in different sites. In SAC, the activation found in the same and in different subjects was more consistent than in other tasks (50-80%). CONCLUSION: The predictive saccade tasks produced evidence for brain-activation and eye-movement reproducibility |
Ryan T. Maloney; Tamara L. Watson; Colin W. G. Clifford Determinants of motion response anisotropies in human early visual cortex: The role of configuration and eccentricity Journal Article In: NeuroImage, vol. 100, pp. 564–579, 2014. @article{Maloney2014, Anisotropies in the cortical representation of various stimulus parameters can reveal the fundamental mechanisms by which sensory properties are analysed and coded by the brain. One example is the preference for motion radial to the point of fixation (i.e. centripetal or centrifugal) exhibited in mammalian visual cortex. In two experiments, this study used functional magnetic resonance imaging (fMRI) to explore the determinants of these radial biases for motion in functionally-defined areas of human early visual cortex, and in particular their dependence upon eccentricity which has been indicated in recent reports. In one experiment, the cortical response to wide-field random dot kinematograms forming 16 different complex motion patterns (including centrifugal, centripetal, rotational and spiral motion) was measured. The response was analysed according to preferred eccentricity within four different eccentricity ranges. Response anisotropies were characterised by enhanced activity for centripetal or centrifugal patterns that changed systematically with eccentricity in visual areas V1-V3 and hV4 (but not V3A/B or V5/MT. +). Responses evolved from a preference for centrifugal over centripetal patterns close to the fovea, to a preference for centripetal over centrifugal at the most peripheral region stimulated, in agreement with previous work. These effects were strongest in V2 and V3. In a second experiment, the stimuli were restricted to within narrow annuli either close to the fovea (0.75-1.88°) or further in the periphery (4.82-6.28°), in a way that preserved the local motion information available in the first experiment. In this configuration a preference for radial motion (centripetal or centrifugal) persisted but the dependence upon eccentricity disappeared. Again this was clearest in V2 and V3. A novel interpretation of the dependence upon eccentricity of motion anisotropies in early visual cortex is offered that takes into account the spatiotemporal "predictability" of the moving pattern. Such stimulus predictability, and its relationship to models of predictive coding, has found considerable support in recent years in accounting for a number of other perceptual and neural phenomena. |
Ulrich J. Pfeiffer; Leonhard Schilbach; Bert Timmermans; Bojana Kuzmanovic; Alexandra L. Georgescu; Gary Bente; Kai Vogeley Why we interact: On the functional role of the striatum in the subjective experience of social interaction Journal Article In: NeuroImage, vol. 101, pp. 124–137, 2014. @article{Pfeiffer2014a, There is ample evidence that human primates strive for social contact and experience interactions with conspecifics as intrinsically rewarding. Focusing on gaze behavior as a crucial means of human interaction, this study employed a unique combination of neuroimaging, eye-tracking, and computer-animated virtual agents to assess the neural mechanisms underlying this component of behavior. In the interaction task, participants believed that during each interaction the agent's gaze behavior could either be controlled by another participant or by a computer program. Their task was to indicate whether they experienced a given interaction as an interaction with another human participant or the computer program based on the agent's reaction. Unbeknownst to them, the agent was always controlled by a computer to enable a systematic manipulation of gaze reactions by varying the degree to which the agent engaged in joint attention. This allowed creating a tool to distinguish neural activity underlying the subjective experience of being engaged in social and non-social interaction. In contrast to previous research, this allows measuring neural activity while participants experience active engagement in real-time social interactions. Results demonstrate that gaze-based interactions with a perceived human partner are associated with activity in the ventral striatum, a core component of reward-related neurocircuitry. In contrast, interactions with a computer-driven agent activate attention networks. Comparisons of neural activity during interaction with behaviorally naïve and explicitly cooperative partners demonstrate different temporal dynamics of the reward system and indicate that the mere experience of engagement in social interaction is sufficient to recruit this system. |
Fabio Richlan; Benjamin Gagl; Stefan Hawelka; Mario Braun; Matthias Schurz; Martin Kronbichler; Florian Hutzler In: Cerebral Cortex, vol. 24, no. 10, pp. 2647–2656, 2014. @article{Richlan2014, The present study investigated the feasibility of using self-paced eye movements during reading (measured by an eye tracker) as markers for calculating hemodynamic brain responses measured by functional magnetic resonance imaging (fMRI). Specifically, we were interested in whether the fixation-related fMRI analysis approach was sensitive enough to detect activation differences between reading material (words and pseudowords) and nonreading material (line and unfamiliar Hebrew strings). Reliable reading-related activation was identified in left hemisphere superior temporal, middle temporal, and occipito-temporal regions including the visual word form area (VWFA). The results of the present study are encouraging insofar as fixation-related analysis could be used in future fMRI studies to clarify some of the inconsistent findings in the literature regarding the VWFA. Our study is the first step in investigating specific visual word recognition processes during self-paced natural sentence reading via simultaneous eye tracking and fMRI, thus aiming at an ecologically valid measurement of reading processes. We provided the proof of concept and methodological framework for the analysis of fixation-related fMRI activation in the domain of reading research. |
Dana Schneider; Virginia P. Slaughter; Stefanie I. Becker; Paul E. Dux Implicit false-belief processing in the human brain Journal Article In: NeuroImage, vol. 101, pp. 268–275, 2014. @article{Schneider2014a, Eye-movement patterns in 'Sally-Anne' tasks reflect humans' ability to implicitly process the mental states of others, particularly false-beliefs - a key theory of mind (ToM) operation. It has recently been proposed that an efficient ToM system, which operates in the absence of awareness (implicit ToM, iToM), subserves the analysis of belief-like states. This contrasts to consciously available belief processing, performed by the explicit ToM system (eToM). The frontal, temporal and parietal cortices are engaged when humans explicitly 'mentalize' about others' beliefs. However, the neural underpinnings of implicit false-belief processing and the extent to which they draw on networks involved in explicit general-belief processing are unknown. Here, participants watched 'Sally-Anne' movies while fMRI and eye-tracking measures were acquired simultaneously. Participants displayed eye-movements consistent with implicit false-belief processing. After independently localizing the brain areas involved in explicit general-belief processing, only the left anterior superior temporal sulcus and precuneus revealed greater blood-oxygen-level-dependent activity for false- relative to true-belief trials in our iToM paradigm. No such difference was found for the right temporal-parietal junction despite significant activity in this area. These findings fractionate brain regions that are associated with explicit general ToM reasoning and false-belief processing in the absence of awareness. |
John M. Henderson; Wonil Choi; Steven G. Luke Morphology of primary visual cortex predicts individual differences in fixation duration during text reading Journal Article In: Journal of Cognitive Neuroscience, vol. 26, no. 12, pp. 2880–2888, 2014. @article{Henderson2014a, In skilled reading, fixations are brief periods of time in which the eyes settle on words. E-Z Reader, a computational model of dynamic reading, posits that fixation durations are under realtime control of lexical processing. Lexical processing, in turn, requires efficient visual encoding. Here we tested the hypothesis that individual differences in fixation durations are related to individual differences in the efficiency of early visual encoding. To test this hypothesis, we recorded participantsʼ eye movements during reading. We then examined individual differences in fixation duration distributions as a function of individual differences in the morphology of primary visual cortex measured from MRI scans. The results showed that greater gray matter surface area and volume in visual cortex predicted shorter and less variable fixation durations in reading. These results suggest that individual differences in eye movements during skilled reading are related to initial visual encoding, consistent with models such as E-Z Reader that emphasize lexical control over fixation time. |
Nora A. Herweg; Bernd Weber; Anna-Maria Kasparbauer; Inga Meyhöfer; Maria Steffens; Nikolaos Smyrnis; Ulrich Ettinger Functional magnetic resonance imaging of sensorimotor transformations in saccades and antisaccades Journal Article In: NeuroImage, vol. 102, pp. 848–860, 2014. @article{Herweg2014, Saccades to peripheral targets require a direct visuomotor transformation. In contrast, antisaccades, saccades in opposite direction of a peripheral target, require more complex transformation processes due to the inversion of the spatial vector. Here, the differential neural mechanisms underlying sensorimotor control in saccades and antisaccades were investigated using functional magnetic resonance imaging (fMRI) at 3. T field strength in 22 human volunteers. We combined a task factor (prosaccades: look towards target; antisaccades: look away from target) with a parametric factor of transformation demand (single vs. multiple peripheral targets) in a two-factorial block design. Behaviorally, a greater number of peripheral targets resulted in decreased spatial accuracy and increased reaction times in antisaccades. No effects were seen on the percentage of antisaccade direction errors or on any prosaccade measures. Neurally, a greater number of targets led to increased BOLD signal in the posterior parietal cortex (PPC) bilaterally. This effect was partially qualified by an interaction that extended into somatosensory cortex, indicating greater increases during antisaccades than prosaccades. The results implicate the PPC as a sensorimotor interface that is especially important in nonstandard mapping for antisaccades and point to a supportive role of somatosensory areas in antisaccade sensorimotor control, possibly by means of proprioceptive processes. |
2013 |
Rasmus Aamand; Thomas Dalsgaard; Yi-Ching Lynn Ho; Arne Møller; Andreas Roepstorff; Torben E. Lund A NO way to BOLD?: Dietary nitrate alters the hemodynamic response to visual stimulation Journal Article In: NeuroImage, vol. 83, pp. 397–407, 2013. @article{Aamand2013, Neurovascular coupling links neuronal activity to vasodilation. Nitric oxide (NO) is a potent vasodilator, and in neurovascular coupling NO production from NO synthases plays an important role. However, another pathway for NO production also exists, namely the nitrate-nitrite-NO pathway. On this basis, we hypothesized that dietary nitrate (NO3-) could influence the brain's hemodynamic response to neuronal stimulation. In the present study, 20 healthy male participants were given either sodium nitrate (NaNO3) or sodium chloride (NaCl) (saline placebo) in a crossover study and were shown visual stimuli based on the retinotopic characteristics of the visual cortex. Our primary measure of the hemodynamic response was the blood oxygenation level dependent (BOLD) response measured with high-resolution functional magnetic resonance imaging (0.64×0.64×1.8mm) in the visual cortex. From this response, we made a direct estimate of key parameters characterizing the shape of the BOLD response (i.e. lag and amplitude). During elevated nitrate intake, corresponding to the nitrate content of a large plate of salad, both the hemodynamic lag and the BOLD amplitude decreased significantly (7.0±2% and 7.9±4%, respectively), and the variation across activated voxels of both measures decreased (12.3±4% and 15.3±7%, respectively). The baseline cerebral blood flow was not affected by nitrate. Ourexperiments demonstrate, for the first time, that dietary nitrate may modulate the local cerebral hemodynamic response to stimuli. A faster and smaller BOLD response, with less variation across local cortex, is consistent with an enhanced hemodynamic coupling during elevated nitrate intake. These findings suggest that dietary patterns, via the nitrate-nitrite-NO pathway, may be a potential way to affect key properties of neurovascular coupling. This could have major clinical implications, which remain to be explored. |
Elisha P. Merriam; Justin L. Gardner; J. Anthony Movshon; David J. Heeger Modulation of visual responses by gaze direction in human visual cortex Journal Article In: Journal of Neuroscience, vol. 33, no. 24, pp. 9879–9889, 2013. @article{Merriam2013, To locate visual objects, the brain combines information about retinal location and direction of gaze. Studies in monkeys have demon-strated that eye position modulates the gain of visual signals with " gain fields, " so that single neurons represent both retinotopic location and eye position. We wished to know whether eye position and retinotopic stimulus location are both represented in human visual cortex. Using functional magnetic resonance imaging, we measured separately for each of several different gaze positions cortical responses to stimuli that varied periodically in retinal locus. Visually evoked responses were periodic following the periodic retinotopic stimulation. Only the response amplitudes depended on eye position; response phases were indistinguishable across eye positions. We used multi-voxel pattern analysis to decode eye position from the spatial pattern of response amplitudes. The decoder reliably discriminated eye position in five of the early visual cortical areas by taking advantage of a spatially heterogeneous eye position-dependent modulation of cortical activity. We conclude that responses in retinotopically organized visual cortical areas are modulated by gain fields qualitatively similar to those previously observed neurophysiologically. |
Jill X. O'Reilly; Urs Schuffelgen; Steven F. Cuell; Timothy E. J. Behrens; Rogier B. Mars; Matthew F. S. Rushworth Dissociable effects of surprise and model update in parietal and anterior cingulate cortex Journal Article In: Proceedings of the National Academy of Sciences, vol. 110, no. 38, pp. E3660–E3669, 2013. @article{OReilly2013, Brains use predictive models to facilitate the processing of expected stimuli or planned actions. Under a predictive model, surprising (low probability) stimuli or actions necessitate the immediate reallocation of processing resources, but they can also signal the need to update the underlying predictive model to reflect changes in the environment. Surprise and updating are often correlated in experimental paradigms but are, in fact, distinct constructs that can be formally defined as the Shannon information (IS) and Kullback-Leibler divergence (DKL) associated with an observation. In a saccadic planning task, we observed that distinct behaviors and brain regions are associated with surprise/IS and updating/DKL. Although surprise/IS was associated with behavioral reprogramming as indexed by slower reaction times, as well as with activity in the posterior parietal cortex [human lateral intraparietal area (LIP)], the anterior cingulate cortex (ACC) was specifically activated during updating of the predictive model (DKL). A second saccade-sensitive region in the inferior posterior parietal cortex (human 7a), which has connections to both LIP and ACC, was activated by surprise and modulated by updating. Pupillometry revealed a further dissociation between surprise and updating with an early positive effect of surprise and late negative effect of updating on pupil area. These results give a computational account of the roles of the ACC and two parietal saccade regions, LIP and 7a, by which their involvement in diverse tasks can be understood mechanistically. The dissociation of functional roles between regions within the reorienting/reprogramming network may also inform models of neurological phenomena, such as extinction and Balint syndrome, and neglect. |
Julia Bender; Kyeong Jin Tark; Benedikt Reuter; Norbert Kathmann; Clayton E. Curtis Differential roles of the frontal and parietal cortices in the control of saccades Journal Article In: Brain and Cognition, vol. 83, no. 1, pp. 1–9, 2013. @article{Bender2013, Although externally as well as internally-guided eye movements allow us to flexibly explore the visual environment, their differential neural mechanisms remain elusive. A better understanding of these neural mechanisms will help us to understand the control of action and to elucidate the nature of cognitive deficits in certain psychiatric populations (e.g. schizophrenia) that show increased latencies in volitional but not visually-guided saccades. Both the superior precentral sulcus (sPCS) and the intraparietal sulcus (IPS) are implicated in the control of eye movements. However, it remains unknown what differential contributions the two areas make to the programming of visually-guided and internally-guided saccades. In this study we tested the hypotheses that sPCS and IPS distinctly encode internally-guided saccades and visually-guided saccades. We scanned subjects with fMRI while they generated visually-guided and internally-guided delayed saccades. We used multi-voxel pattern analysis to test whether patterns of cue related, preparatory and saccade related activation could be used to predict the direction of the planned eye movement. Results indicate that patterns in the human sPCS predicted internally-guided saccades but not visually-guided saccades in all trial periods and patterns in the IPS predicted internally-guided saccades and visually-guided saccades equally well. The results support the hypothesis that the human sPCS and IPS make distinct contributions to the control of volitional eye movements. |
Renée M. Visser; H. Steven Scholte; Tinka Beemsterboer; Merel Kindt Neural pattern similarity predicts long-term fear memory Journal Article In: Nature Neuroscience, vol. 16, no. 4, pp. 388–390, 2013. @article{Visser2013, Although certain changes in the brain may reflect fear learning, there are no known markers that indicate whether an aversive experience will develop into fear memory. We examined the moment-to-moment dynamics of human fear learning by applying multi-voxel pattern analysis to single-trial blood oxygen level–dependent magnetic resonance imaging data. We found that the long-term behavioral expression of fear memory could be predicted from neural patterns at the time of learning. |
Semir Zeki; Jonathan Stutters Functional specialization and generalization for grouping of stimuli based on colour and motion Journal Article In: NeuroImage, vol. 73, pp. 156–166, 2013. @article{Zeki2013, This study was undertaken to learn whether the principle of functional specialization that is evident at the level of the prestriate visual cortex extends to areas that are involved in grouping visual stimuli according to attribute, and specifically according to colour and motion. Subjects viewed, in an fMRI scanner, visual stimuli composed of moving dots, which could be either coloured or achromatic; in some stimuli the moving coloured dots were randomly distributed or moved in random directions; in others, some of the moving dots were grouped together according to colour or to direction of motion, with the number of groupings varying from 1 to 3. Increased activation was observed in area V4 in response to colour grouping and in V5 in response to motion grouping while both groupings led to activity in separate though contiguous compartments within the intraparietal cortex. The activity in all the above areas was parametrically related to the number of groupings, as was the prominent activity in Crus I of the cerebellum where the activity resulting from the two types of grouping overlapped. This suggests (a) that, the specialized visual areas of the prestriate cortex have functions beyond the processing of visual signals according to attribute, namely that of grouping signals according to colour (V4) or motion (V5); (b) that the functional separation evident in visual cortical areas devoted to motion and colour, respectively, is maintained at the level of parietal cortex, at least as far as grouping according to attribute is concerned; and (c) that, by contrast, this grouping-related functional segregation is not maintained at the level of the cerebellum. |
Kyeong Jin Tark; Clayton E. Curtis Deciding where to look based on visual, auditory, and semantic information Journal Article In: Brain Research, vol. 1525, pp. 26–38, 2013. @article{Tark2013, Neurons in the dorsal frontal and parietal cortex are thought to transform incoming visual signals into the spatial goals of saccades, a process known as target selection. Here, we used functional magnetic resonance imaging (fMRI) to test how target selection may generalize beyond visual transformations when auditory and semantic information is used for selection. We compared activity in the frontal and parietal cortex when subjects made visually, aurally, and semantically guided saccades to one of four differently colored dots. Selection was based on a visual cue (i.e., one of the dots blinked), an auditory cue (i.e., a white noise burst was emitted at one of the dots' location), or a semantic cue (i.e., the color of one of the dots was spoken). Although neural responses in frontal and parietal cortex were robust, they were non-specific with regard to the type of information used for target selection. Decoders, however, trained on the patterns of activity in the intraparietal sulcus could classify both the type of cue used for target selection and the direction of the saccade. Therefore, we find evidence that the posterior parietal cortex is involved in transforming multimodal inputs into general spatial representations that can be used to guide saccades. |
Ryan T. Maloney; Tamara L. Watson; Colin W. G. Clifford Human cortical and behavioral sensitivity to patterns of complex motion at eccentricity Journal Article In: Journal of Neurophysiology, vol. 110, no. 11, pp. 2545–2556, 2013. @article{Maloney2013, Complex patterns of image motion (contracting, expanding, rotating, and spiraling fields) are important in the coordination of visually guided behaviors. Whereas specialized detectors in monkey visual cortex show selectivity for particular patterns of complex motion, their representation in human visual cortex remains unclear. In the present study, functional magnetic resonance imaging (fMRI) was used to investigate the sensitivity of functionally defined regions of human visual cortex to parametrically modulated complex motion trajectories, coupled with complementary psychophysical testing. A unique stimulus design made it possible to disambiguate the neural responses and psychophysical sensitivity to complex motions per se from the distribution of local motions relative to the fovea, which are known to enhance cortical activity when presented radial to fixation. This involved presenting several small, separate motion fields in the periphery in a manner that distinguished them from global optic flow patterns. The patterns were morphed through complex motion space in a systematic time-locked fashion when presented in the scanner. Anisotropies were observed in the fMRI signal, marked by an enhanced response to expanding vs. contracting fields, even in early visual cortex. Anisotropies in the psychophysical sensitivity measures followed a similar pattern that was correlated with activity in areas hV4, V5/MT, and MST. This represents the first systematic examination of complex motion perception at both a behavioral and neural level in human observers. The characteristic processing anisotropy revealed in both data sets can inform models of complex motion processing, particularly with respect to computations performed in early visual cortex. |
Tim J. Preston; Fei Guo; Koel Das; Barry Giesbrecht; Miguel P. Eckstein Neural representations of contextual guidance in visual search of real-world scenes Journal Article In: Journal of Neuroscience, vol. 33, no. 18, pp. 7846–7855, 2013. @article{Preston2013, Exploiting scene context and object– object co-occurrence is critical in guiding eye movements and facilitating visual search, yet the mediating neural mechanisms are unknown. We used functional magnetic resonance imaging while observers searched for target objects in scenes and used multivariate pattern analyses (MVPA) to show that the lateral occipital complex (LOC) can predict the coarse spatial location of observers' expectations about the likely location of 213 different targets absent from the scenes. In addition, we found weaker but significant representations of context location in an area related to the orienting of attention (intraparietal sulcus, IPS) as well as a region related to scene processing (retrosplenial cortex, RSC). Importantly, the degree of agreement among 100 independent raters about the likely location to contain a target object in a scene correlated with LOC's ability to predict the contextual location while weaker but significant effects were found in IPS, RSC, the human motion area, and early visual areas (V1, V3v). When contextual information was made irrelevant to observers' behavioral task, the MVPA analysis of LOC and the other areas' activity ceased to predict the location of context. Thus, our findings suggest that the likely locations of targets in scenes are represented in various visual areas with LOC playing a key role in contextual guidance during visual search of objects in real scenes. |
Chen Song; D. Samuel Schwarzkopf; Antoine Lutti; Baojuan Li; Ryota Kanai; Geraint Rees Effective connectivity within human primary visual cortex predicts interindividual diversity in illusory perception Journal Article In: Journal of Neuroscience, vol. 33, no. 48, pp. 18781–18791, 2013. @article{Song2013c, Visual perception depends strongly on spatial context. A classic example is the tilt illusion where the perceived orientation of a central stimulus differs from its physical orientation when surrounded by tilted spatial contexts. Here we show that such contextual modulation of orientation perception exhibits trait-like interindividual diversity that correlates with interindividual differences in effective connectivity within human primary visual cortex. We found that the degree to which spatial contexts induced illusory orientation perception, namely, the magnitude of the tilt illusion, varied across healthy human adults in a trait-like fashion independent of stimulus size or contrast. Parallel to contextual modulation of orientation perception, the presence of spatial contexts affected effective connectivity within human primary visual cortex between peripheral and foveal representations that responded to spatial context and central stimulus, respectively. Importantly, this effective connectivity from peripheral to foveal primary visual cortex correlated with interindividual differences in the magnitude of the tilt illusion. Moreover, this correlation with illusion perception was observed for effective connectivity under tilted contextual stimulation but not for that under iso-oriented contextual stimulation, suggesting that it reflected the impact of orientation-dependent intra-areal connections. Our findings revealed an interindividual correlation between intra-areal connectivity within primary visual cortex and contextual influence on orientation perception. This neurophysiological-perceptual link provides empirical evidence for theoretical proposals that intra-areal connections in early visual cortices are involved in contextual modulation of visual perception. |
Jamie Ferri; Joseph Schmidt; Greg Hajcak; Turhan Canli Neural correlates of attentional deployment within unpleasant pictures Journal Article In: NeuroImage, vol. 70, pp. 268–277, 2013. @article{Ferri2013, Attentional deployment is an emotion regulation strategy that involves shifting attentional focus towards or away from particular aspects of emotional stimuli. Previous studies have highlighted the prevalence of attentional deployment and demonstrated that it can have a significant impact on brain activity and behavior. However, little is known about the neural correlates of this strategy. The goal of the present studies was to examine the effect of attentional deployment on neural activity by directing attention to more or less arousing portions of unpleasant images. In Studies 1 and 2, participants passively viewed counterbalanced blocks of unpleasant images without a focus, unpleasant images with an arousing focus, unpleasant images with a non-arousing focus, neutral images without a focus, and neutral images with a non-arousing focus for 4000. ms each. In Study 2, eye-tracking data were collected on all participants during image acquisition. In both studies, affect ratings following each block indicated that participants felt significantly less negative affect after viewing unpleasant images with a non-arousing focus compared to unpleasant images with an arousing focus. In both studies, the unpleasant non-arousing focus condition compared to the unpleasant arousing focus condition was associated with increased activity in frontal and parietal regions implicated in inhibitory control and visual attention. In Study 2, the unpleasant non-arousing focus condition compared to the unpleasant arousing focus condition was associated with reduced activity in the amygdala and visual cortex. Collectively these data suggest that attending to a non-arousing portion of an unpleasant image successfully reduces subjective negative affect and recruits fronto-parietal networks implicated in inhibitory control. Moreover, when ensuring task compliance by monitoring eye movements, attentional deployment modulates amygdala activity. |
Jeremy Freeman; David J. Heeger; Elisha P. Merriam Coarse-scale biases for spirals and orientation in human visual cortex Journal Article In: Journal of Neuroscience, vol. 33, no. 50, pp. 19695–19703, 2013. @article{Freeman2013, Multivariate decoding analyses are widely applied to functional magnetic resonance imaging (fMRI) data, but there is controversy over their interpretation. Orientation decoding in primary visual cortex (V1) reflects coarse-scale biases, including an over-representation of radial orientations. But fMRI responses to clockwise and counter-clockwise spirals can also be decoded. Because these stimuli are matched for radial orientation, while differing in local orientation, it has been argued that fine-scale columnar selectivity for orientation contributes to orientation decoding. We measured fMRI responses in human V1 to both oriented gratings and spirals. Responses to oriented gratings exhibited a complex topography, including a radial bias that was most pronounced in the peripheral representation, and a near-vertical bias that was most pronounced near the foveal representation. Responses to clockwise and counter-clockwise spirals also exhibited coarse-scale organization, at the scale of entire visual quadrants. The preference of each voxel for clockwise or counter-clockwise spirals was predicted from the preferences of that voxel for orientation and spatial position (i.e., within the retinotopic map). Our results demonstrate a bias for local stimulus orientation that has a coarse spatial scale, is robust across stimulus classes (spirals and gratings), and suffices to explain decoding from fMRI responses in V1. |
Tsafrir Greenberg; Joshua M. Carlson; Jiook Cha; Greg Hajcak; Lilianne R. Mujica-Parodi Neural reactivity tracks fear generalization gradients Journal Article In: Biological Psychology, vol. 92, no. 1, pp. 2–8, 2013. @article{Greenberg2013, Recent studies on fear generalization have demonstrated that fear-potentiated startle and skin conductance responses to a conditioned stimulus (CS) generalize to similar stimuli, with the strength of the fear response linked to perceptual similarity to the CS. The aim of the present study was to extend this work by examining neural correlates of fear generalization. An initial experiment (N= 8) revealed that insula reactivity tracks the conditioned fear gradient. We then replicated this effect in a larger independent sample (N= 25). Activation in the insula, anterior cingulate, right supplementary motor cortex and caudate increased reactivity as generalization stimuli (GS) were more similar to the CS, consistent with participants' overall ratings of perceived shock likelihood and pupillary response to each stimulus. |
Tsafrir Greenberg; Joshua M. Carlson; Jiook Cha; Greg Hajcak; Lilianne R. Mujica-Parodi Ventromedial prefrontal cortex reactivity is altered in generalized anxiety disorder during fear generalization Journal Article In: Depression and Anxiety, vol. 30, no. 3, pp. 242–250, 2013. @article{Greenberg2013a, BACKGROUND: Fear generalization is thought to contribute to the development and maintenance of anxiety symptoms and accordingly has been the focus of recent research. Previously, we reported that in healthy individuals (N = 25) neural reactivity in the insula, anterior cingulate cortex (ACC), supplementary motor area (SMA), and caudate follow a generalization gradient with a peak response to a conditioned stimulus (CS) that declines with greater perceptual dissimilarity of generalization stimuli (GS) to the CS. In contrast, reactivity in the ventromedial prefrontal cortex (vmPFC), a region linked to fear inhibition, showed an opposite response pattern. The aim of the current study was to examine whether neural responses to fear generalization differ in generalized anxiety disorder (GAD). A second aim was to examine connectivity of primary regions engaged by the generalization task in the GAD group versus healthy group, using psychophysiological interaction analysis. METHODS: Thirty-two women diagnosed with GAD were scanned using the same generalization task as our healthy group. RESULTS: Individuals with GAD exhibited a less discriminant vmPFC response pattern suggestive of deficient recruitment of vmPFC during fear inhibition. Across participants, there was enhanced anterior insula (aINS) coupling with the posterior insula, ACC, SMA, and amygdala during presentation of the CS, consistent with a modulatory role for the aINS in the execution of fear responses. CONCLUSIONS: These findings suggest that deficits in fear regulation, rather than in the excitatory response itself, are more critical to the pathophysiology of GAD in the context of fear generalization. |
2012 |
Nathan Faivre; Sylvain Charron; Paul Roux; Stephane Lehericy; Sid Kouider Nonconscious emotional processing involves distinct neural pathways for pictures and videos Journal Article In: Neuropsychologia, vol. 50, pp. 3736–3744, 2012. @article{Faivre2012a, Facial expressions are known to impact observers' behavior, even when they are not consciously identifiable. Relying on visual crowding, a perceptual phenomenon whereby peripheral faces become undiscriminable, we show that participants exposed to happy vs. neutral crowded faces rated the pleasantness of subsequent neutral targets accordingly to the facial expression's valence. Using functional magnetic resonance imaging (fMRI) along with psychophysiological interaction analysis, we investigated the neural determinants of this nonconscious preference bias, either induced by static (i.e., pictures) or dynamic (i.e., videos) facial expressions. We found that while static expressions activated primarily the ventral visual pathway (including task-related functional connectivity between the fusiform face area and the amygdala), dynamic expressions triggered the dorsal visual pathway (i.e., posterior partietal cortex) and the substantia innominata, a structure that is contiguous with the dorsal amygdala. As temporal cues are known to improve the processing of visible facial expressions, the absence of ventral activation we observed with crowded videos questions the capacity to integrate facial features and facial motions without awareness. Nevertheless, both static and dynamic facial expressions activated the hippocampus and the orbitofrontal cortex, suggesting that nonconscious preference judgments may arise from the evaluation of emotional context and the computation of aesthetic evaluation. |
Scott A. Guerin; Clifford A. Robbins; Adrian W. Gilmore; Daniel L. Schacter Retrieval failure contributes to gist-based false recognition Journal Article In: Journal of Memory and Language, vol. 66, no. 1, pp. 68–78, 2012. @article{Guerin2012, People often falsely recognize items that are similar to previously encountered items. This robust memory error is referred to as gist-based false recognition. A widely held view is that this error occurs because the details fade rapidly from our memory. Contrary to this view, an initial experiment revealed that, following the same encoding conditions that produce high rates of gist-based false recognition, participants overwhelmingly chose the correct target rather than its related foil when given the option to do so. A second experiment showed that this result is due to increased access to stored details provided by reinstatement of the originally encoded photograph, rather than to increased attention to the details. Collectively, these results suggest that details needed for accurate recognition are, to a large extent, still stored in memory and that a critical factor determining whether false recognition will occur is whether these details can be accessed during retrieval. |
Fei Guo; Tim J. Preston; Koel Das; Barry Giesbrecht; Miguel P. Eckstein Feature-independent neural coding of target detection during search of natural scenes Journal Article In: Journal of Neuroscience, vol. 32, no. 28, pp. 9499–9510, 2012. @article{Guo2012, Visual search requires humans to detect a great variety of target objects in scenes cluttered by other objects or the natural environment. It is unknown whether there is a general purpose neural detection mechanism in the brain that codes the presence of a wide variety of categories of objects embedded in natural scenes. We provide evidence for a feature-independent coding mechanism for detecting behaviorally relevant targets in natural scenes in the dorsal frontoparietal network. Pattern classifiers using single-trial fMRI responses in the dorsal frontoparietal network reliably predicted the presence of 368 different target objects and also the observer's choices. Other vision-related areas such as the primary visual cortex, lateral occipital complex, the parahippocampal, and the fusiform gyri did not predict target presence, while high-level association areas related to general purpose decision making, including the dorsolateral prefrontal cortex and anterior cingulate, did. Activity in the intraparietal sulcus, a main area in the dorsal frontoparietal network, correlated with observers' decision confidence and with the task difficulty of individual images. These results cannot be explained by physical differences across images or eye movements. Thus, the dorsal frontoparietal network detects behaviorally relevant targets in natural scenes independent of their defining visual features and may be the human analog of the priority map in monkey lateral intraparietal cortex. |
Trenton A. Jerde; Elisha P. Merriam; Adam C. Riggall; James H. Hedges; Clayton E. Curtis Prioritized maps of space in human frontoparietal cortex Journal Article In: Journal of Neuroscience, vol. 32, no. 48, pp. 17382–17390, 2012. @article{Jerde2012, Priority maps are theorized to be composed of large populations of neurons organized topographically into a map of gaze-centered space whose activity spatially tags salient and behaviorally relevant information. Here, we identified four priority map candidates along human posterior intraparietal sulcus (IPS0-IPS3) and two along the precentral sulcus (PCS) that contained reliable retinotopically organized maps of contralateral visual space. Persistent activity increased from posterior-to-anterior IPS areas and from inferior-to-superior PCS areas during the maintenance of a working memory representation, the maintenance of covert attention, and the maintenance of a saccade plan. Moreover, decoders trained to predict the locations on one task (e.g., working memory) cross-predicted the locations on other tasks (e.g., attention) in superior PCS and IPS2, suggesting that these patterns of maintenance activity may be interchangeable across the tasks. Such properties make these two areas in frontal and parietal cortex viable priority map candidates. |
Zoran Josipovic; Ilan Dinstein; Jochen Weber; David J. Heeger Influence of meditation on anti-correlated networks in the brain Journal Article In: Frontiers in Human Neuroscience, vol. 5, pp. 183, 2012. @article{Josipovic2012, Human experiences can be broadly divided into those that are external and related to interaction with the environment, and experiences that are internal and self-related. The cerebral cortex appears to be divided into two corresponding systems: an "extrinsic" system composed of brain areas that respond more to external stimuli and tasks and an "intrinsic" system composed of brain areas that respond less to external stimuli and tasks. These two broad brain systems seem to compete with each other, such that their activity levels over time is usually anti-correlated, even when subjects are "at rest" and not performing any task. This study used meditation as an experimental manipulation to test whether this competition (anti-correlation) can be modulated by cognitive strategy. Participants either fixated without meditation (fixation), or engaged in non-dual awareness (NDA) or focused attention (FA) meditations. We computed inter-area correlations ("functional connectivity") between pairs of brain regions within each system, and between the entire extrinsic and intrinsic systems. Anti-correlation between extrinsic vs. intrinsic systems was stronger during FA meditation and weaker during NDA meditation in comparison to fixation (without mediation). However, correlation between areas within each system did not change across conditions. These results suggest that the anti-correlation found between extrinsic and intrinsic systems is not an immutable property of brain organization and that practicing different forms of meditation can modulate this gross functional organization in profoundly different ways. |
2011 |
Jeremy Freeman; G. J. Brouwer; David J. Heeger; Elisha P. Merriam Orientation decoding depends on maps, not columns Journal Article In: Journal of Neuroscience, vol. 31, no. 13, pp. 4792–4804, 2011. @article{Freeman2011a, The representation of orientation in primary visual cortex (V1) has been examined at a fine spatial scale corresponding to the columnar architecture. We present functional magnetic resonance imaging (fMRI) measurements providing evidence for a topographic map of orientation preference in human V1 at a much coarser scale, in register with the angular-position component of the retinotopic map of V1. This coarse-scale orientation map provides a parsimonious explanation for why multivariate pattern analysis methods succeed in decoding stimulus orientation from fMRI measurements, challenging the widely held assumption that decoding results reflect sampling of spatial irregularities in the fine-scale columnar architecture. Decoding stimulus attributes and cognitive states from fMRI measurements has proven useful for a number of applications, but our results demonstrate that the interpretation cannot assume decoding reflects or exploits columnar organization. |
2010 |
Katrin Herrmann; Leila Montaser-Kouhsari; Marisa Carrasco; David J. Heeger When size matters: Attention affects performance by contrast or response gain Journal Article In: Nature Neuroscience, vol. 13, no. 12, pp. 1554–1561, 2010. @article{Herrmann2010, Covert attention, the selective processing of visual information in the absence of eye movements, improves behavioral performance. Here, we show that attention, both exogenous (involuntary) and endogenous (voluntary), can affect performance by contrast or response gain changes, depending on the stimulus size and the relative size of the attention field. These two variables were manipulated in a cueing task while varying stimulus contrast. We observed a change in behavioral performance consonant with a change in contrast gain for small stimuli paired with spatial uncertainty, but a change in response gain for large stimuli presented at one location (no uncertainty) and surrounded by irrelevant flanking distracters. A complementary neuroimaging experiment revealed that observers' attention field was wider with than without spatial uncertainty. Our results support key predictions of the normalization model of attention, and reconcile previous, seemingly contradictory, findings on the effects of visual attention. Introduction |
2005 |
Zoe Kourtzi; Lisa R. Betts; Pegah Sarkheil; Andrew E. Welchman Distributed neural plasticity for shape learning in the human visual cortex Journal Article In: PLoS Biology, vol. 3, no. 7, pp. 1317–1327, 2005. @article{Kourtzi2005, Expertise in recognizing objects in cluttered scenes is a critical skill for our interactions in complex environments and is thought to develop with learning. However, the neural implementation of object learning across stages of visual analysis in the human brain remains largely unknown. Using combined psychophysics and functional magnetic resonance imaging (fMRI), we show a link between shape-specific learning in cluttered scenes and distributed neuronal plasticity in the human visual cortex. We report stronger fMRI responses for trained than untrained shapes across early and higher visual areas when observers learned to detect low-salience shapes in noisy backgrounds. However, training with high-salience pop-out targets resulted in lower fMRI responses for trained than untrained shapes in higher occipitotemporal areas. These findings suggest that learning of camouflaged shapes is mediated by increasing neural sensitivity across visual areas to bolster target segmentation and feature integration. In contrast, learning of prominent pop-out shapes is mediated by associations at higher occipitotemporal areas that support sparser coding of the critical features for target recognition. We propose that the human brain learns novel objects in complex scenes by reorganizing shape processing across visual areas, while taking advantage of natural image correlations that determine the distinctiveness of target shapes. |