Directed information flow: a model free measure to analyze causal interactions in event related EEG-MEG-experiments

In a study that combined event related potential (ERP) and magnetic field (ERMF) data, we analyzed the timing and direction of information flow between striate (S) and extrastriate (ES) cortex by applying a generalized mutual information measure (DIT for "directed information transfer") during a visual spatial attention task. ERP and ERMF recordings showed that selective attention to stimulus arrays in one visual field enhanced late responses (around 200 ms after the stimulus presentation) that were localized in S (ERMF) and ES (ERP) cortex. The results of the DIT analysis indicate there is a significant attention related increase in the flow of information back from ES to S cortex at around 220 ms, with an associated decrease in the flow of information forward from S cortex to ES cortex. These results support the hypothesis that a feedback mechanism guides attention-related processing in primary visual cortex and provide evidence that DIT can by used to evaluate the direction of information flow between cortical areas.     

Modulation of auditory neural responses by a visual context in human fear conditioning.

Responses to a stimulus signaling danger depend not only on the nature of that stimulus, but also on the context in which it is presented. A large body of work has been conducted in experimental animals investigating the neural correlates of contextual modulation of fear responses. However, much less is known about this process in humans. In this study we used functional MRI in a fear conditioning paradigm to explore this phenomenon. Responses to acoustic conditioned stimuli in auditory cortex were modulated by the presence of a visual context which signaled the likelihood of receiving an aversive unconditioned stimulus. Furthermore, the presence of the aversive visual context was associated with enhanced activity in parietal cortex, which may reflect an increase in attention to the presence of environmental threat stimuli.     

Topography of attention in the primary visual cortex

Previous research suggests that feedback circuits mediate the effect of attention to the primary visual cortex (V1). This inference is mainly based on temporal information of the responses, where late modulation is associated with feedback signals. However, temporal data alone are inconclusive because the anatomical hierarchy between cortical areas differs significantly from the temporal sequence of activation. In the current work, we relied on recent physiological and computational models of V1 network architecture, which have shown that the thalamic feedforward, local horizontal and feedback contribution are reflected in the spatial spread of responses. We used multifocal functional localizer and quantitative analysis in functional magnetic resonance imaging to determine the spatial scales of attention and sensory responses. Representations of 60 visual field regions in V1 were functionally localized and four of these regions were targets in a subsequent attention experiment, where human volunteers fixated centrally and performed a visual discrimination task at the attended location. Attention enhanced the peak amplitudes significantly more in the lower than in the upper visual field. This enhancement by attention spread with a 2.4 times larger radius (approximately 10 mm, assuming an average magnification factor) compared with the unattended response. The corresponding target region of interest was on average 20% stronger than that caused by the afferent sensory stimulation alone. This modulation could not be attributed to eye movements. Given the contemporary view of primate V1 connections, the activation spread along the cortex provides further evidence that the signal enhancement by spatial attention is dependent on feedback circuits.     

Brain mechanisms of involuntary visuospatial attention: an event-related potential study

The brain mechanisms mediating visuospatial attention were investigated by recording event-related potentials (ERPs) during a line-orientation discrimination task. Nonpredictive peripheral cues were used to direct participant's attention involuntarily to a spatial location. The earliest attentional modulation was observed in the P1 component (peak latency about 130 ms), with the valid trials eliciting larger P1 than invalid trials. Moreover, the attentional modulations on both the amplitude and latency of the P1 and N1 components had a different pattern as compared to previous studies with voluntary attention tasks. In contrast, the earliest visual ERP component, C1 (peak latency about 80 ms), was not modulated by attention. Low-resolution brain electromagnetic tomography (LORETA) showed that the earliest attentional modulation occurred in extrastriate cortex (middle occipital gyrus, BA 19) but not in the primary visual cortex. Later attention-related reactivations in the primary visual cortex were found at about 110 ms after stimulus onset. The results suggest that involuntary as well as voluntary attention modulates visual processing at the level of extrastriate cortex; however, at least some different processes are involved by involuntary attention compared to voluntary attention. In addition, the possible feedback from higher visual cortex to the primary visual cortex is faster and occurs earlier in involuntary relative to voluntary attention task.     

Mapping feature-sensitivity and attentional modulation in human auditory cortex with functional magnetic resonance imaging

Feature-specific enhancement refers to the process by which selectively attending to a particular stimulus feature specifically increases the response in the same region of the brain that codes that stimulus property. Whereas there are many demonstrations of this mechanism in the visual system, the evidence is less clear in the auditory system. The present functional magnetic resonance imaging (fMRI) study examined this process for two complex sound features, namely frequency modulation (FM) and spatial motion. The experimental design enabled us to investigate whether selectively attending to FM and spatial motion enhanced activity in those auditory cortical areas that were sensitive to the two features. To control for attentional effort, the difficulty of the target-detection tasks was matched as closely as possible within listeners. Locations of FM-related and motion-related activation were broadly compatible with previous research. The results also confirmed a general enhancement across the auditory cortex when either feature was being attended to, as compared with passive listening. The feature-specific effects of selective attention revealed the novel finding of enhancement for the nonspatial (FM) feature, but not for the spatial (motion) feature. However, attention to spatial features also recruited several areas outside the auditory cortex. Further analyses led us to conclude that feature-specific effects of selective attention are not statistically robust, and appear to be sensitive to the choice of fMRI experimental design and localizer contrast.     

Attentional modulation in visual cortex is modified during perceptual learning

Practicing a visual task commonly results in improved performance. Often the improvement does not transfer well to a new retinal location, suggesting that it is mediated by changes occurring in early visual cortex, and indeed neuroimaging and neurophysiological studies both demonstrate that perceptual learning is associated with altered activity in visual cortex. Theoretical treatments tend to invoke neuroplasticity that refines early sensory processing. An alternative possibility is that performance is improved because of an altered attentional strategy and that the changes in early visual areas reflect locally altered top-down attentional modulation. To test this idea, we have used functional MRI to examine changes in attentional modulation in visual cortex while participants learn an orientation discrimination task. By examining activity in visual cortex during the preparatory period when the participant has been cued to attend to an upcoming stimulus, we isolated the top-down modulatory signal received by the visual cortex. We show that this signal changes as learning progresses, possibly reflecting gradual automation of the task. By manipulating task difficulty, we show that the change mirrors performance, occurring most quickly for easier stimuli. The effects were seen only at the retinal locus of the stimulus, ruling out a generalized change in alertness. The results suggest that spatial attention changes during perceptual learning and that this may account for some of the concomitant changes seen in visual cortex.     

Contrast independence of cardinal preference: stable oblique effect in orientation maps of ferret visual cortex

The oblique effect was first described as enhanced detection and discrimination of cardinal orientations compared with oblique orientations. Such biases in visual processing are believed to originate from a functional adaptation to environmental statistics dominated by cardinal contours. At the neuronal level, the oblique orientation effect corresponds to the numerical overrepresentation and narrower tuning bandwidths of cortical neurons representing the cardinal axes. The anisotropic distribution of orientation preferences over large cortical regions was revealed with optical imaging, providing further evidence for the cortical oblique effect in several mammalian species. Our present study explores whether the dominant representation of cardinal contours persists at different stimulus contrasts. Performing intrinsic optical imaging in the ferret visual cortex and presenting drifting gratings at various orientations and contrasts (100%, 30% and 10%), we found that the overrepresentation of vertical and horizontal contours was invariant across stimulus contrasts. In addition, the responses to cardinal orientations were also more robust and evoked larger modulation depths than responses to oblique orientations. We conclude that orientation maps remain constant across the full range of contrast levels down to detection thresholds. Thus, a stable layout of the functional architecture dedicated to processing oriented edges seems to reflect a fundamental coding strategy of the early visual cortex.     

The parietal cortex and attentional modulations of activities of the visual cortex

We recorded high density event-related brain potentials (ERPs) from a patient with focal left parietal damage in a covert visual orienting task requiring detection of targets in the attended or unattended hemifield. A positivity peaking at 120 ms (P1) to the left visual field stimuli was enlarged when attended than unattended and was localized to the right extrastirate cortex. However, spatial attention did not influence the ERPs to the right visual field stimuli. The leftward cue elicited an enlarged P1 relative to the rightward cue. The results suggest that human parietal cortex is critical for the attentional modulation of the neural activities in the extrastriate cortex associated with stimuli in the contralateral hemifield.     

Temporal dynamics of the attentional spotlight: neuronal correlates of attentional capture and inhibition of return in early visual cortex

A stimulus that suddenly appears in the corner of the eye inevitably captures our attention, and this in turn leads to faster detection of a second stimulus presented at the same position shortly thereafter. After about 250 msec, however, this effect reverses and the second stimulus is detected faster when it appears far away from the first. Here, we report a potential physiological correlate of this time-dependent attentional facilitation and inhibition. We measured the activity in visual cortex representations of the second (target) stimulus' location depending on the stimulus onset asynchrony (SOA) and spatial distance that separated the target from the preceding cue stimulus. At an SOA of 100 msec, the target yielded larger responses when it was presented near to than far away from the cue. At an SOA of 850 msec, however, the response to the target was more pronounced when it appeared far away from the cue. Our data show how the neural substrate of visual orienting is guided by immediately preceding sensory experience and how a fast-reacting brain system modulates sensory processing by briefly increasing and subsequently decreasing responsiveness in parts of the visual cortex. We propose these activity modulations as the neural correlate of the sequence of perceptual facilitation and inhibition after attentional capture.     

Modulation of early cortical processing during divided attention to non‐contiguous locations

We often face the challenge of simultaneously attending to multiple non‐contiguous regions of space. There is ongoing debate as to how spatial attention is divided under these situations. Whereas, for several years, the predominant view was that humans could divide the attentional spotlight, several recent studies argue in favor of a unitary spotlight that rhythmically samples relevant locations. Here, this issue was addressed by the use of high‐density electrophysiology in concert with the multifocal m‐sequence technique to examine visual evoked responses to multiple simultaneous streams of stimulation. Concurrently, we assayed the topographic distribution of alpha‐band oscillatory mechanisms, a measure of attentional suppression. Participants performed a difficult detection task that required simultaneous attention to two stimuli in contiguous (undivided) or non‐contiguous parts of space. In the undivided condition, the classic pattern of attentional modulation was observed, with increased amplitude of the early visual evoked response and increased alpha amplitude ipsilateral to the attended hemifield. For the divided condition, early visual responses to attended stimuli were also enhanced, and the observed multifocal topographic distribution of alpha suppression was in line with the divided attention hypothesis. These results support the existence of divided attentional spotlights, providing evidence that the corresponding modulation occurs during initial sensory processing time‐frames in hierarchically early visual regions, and that suppressive mechanisms of visual attention selectively target distracter locations during divided spatial attention.     

Backward masked fearful faces enhance contralateral occipital cortical activity for visual targets within the spotlight of attention

Spatial attention has been argued to be adaptive by enhancing the processing of visual stimuli within the ‘spotlight of attention’. We previously reported that crude threat cues (backward masked fearful faces) facilitate spatial attention through a network of brain regions consisting of the amygdala, anterior cingulate and contralateral visual cortex. However, results from previous functional magnetic resonance imaging (fMRI) dot-probe studies have been inconclusive regarding a fearful face-elicited contralateral modulation of visual targets. Here, we tested the hypothesis that the capture of spatial attention by crude threat cues would facilitate processing of subsequently presented visual stimuli within the masked fearful face-elicited ‘spotlight of attention’ in the contralateral visual cortex. Participants performed a backward masked fearful face dot-probe task while brain activity was measured with fMRI. Masked fearful face left visual field trials enhanced activity for spatially congruent targets in the right superior occipital gyrus, fusiform gyrus and lateral occipital complex, while masked fearful face right visual field trials enhanced activity in the left middle occipital gyrus. These data indicate that crude threat elicited spatial attention enhances the processing of subsequent visual stimuli in contralateral occipital cortex, which may occur by lowering neural activation thresholds in this retinotopic location.     

Tagging cortical networks in emotion: A topographical analysis

Viewing emotional pictures is associated with heightened perception and attention, indexed by a relative increase in visual cortical activity. Visual cortical modulation by emotion is hypothesized to reflect re‐entrant connectivity originating in higher‐order cortical and/or limbic structures. The present study used dense‐array electroencephalography and individual brain anatomy to investigate functional coupling between the visual cortex and other cortical areas during affective picture viewing. Participants viewed pleasant, neutral, and unpleasant pictures that flickered at a rate of 10 Hz to evoke steady‐state visual evoked potentials (ssVEPs) in the EEG. The spectral power of ssVEPs was quantified using Fourier transform, and cortical sources were estimated using beamformer spatial filters based on individual structural magnetic resonance images. In addition to lower‐tier visual cortex, a network of occipito‐temporal and parietal (bilateral precuneus, inferior parietal lobules) structures showed enhanced ssVEP power when participants viewed emotional (either pleasant or unpleasant), compared to neutral pictures. Functional coupling during emotional processing was enhanced between the bilateral occipital poles and a network of temporal (left middle/inferior temporal gyrus), parietal (bilateral parietal lobules), and frontal (left middle/inferior frontal gyrus) structures. These results converge with findings from hemodynamic analyses of emotional picture viewing and suggest that viewing emotionally engaging stimuli is associated with the formation of functional links between visual cortex and the cortical regions underlying attention modulation and preparation for action. Hum Brain Mapp, 2012. © 2011 Wiley Periodicals, Inc.     

The attentional effects of peripheral cueing as revealed by two event-related potential studies.

OBJECTIVE: The mechanism of visual spatial attention elicited by peripheral cueing was investigated in two studies. METHOD: Event-related potentials (ERPs) were recorded when the subjects were performing a spatial frequency discrimination task and a location discrimination task. Stimuli were randomly flashed in the left or right visual field. Prior to each stimulus a peripheral cue was presented with a validity of 75%. RESULTS: The subjects responded faster to valid trials than to invalid trials. The earliest visual ERP component, C1, was not modulated by the cue validity, suggesting that visual spatial attention elicited by peripheral cueing does not involve striate cortex. Valid trials elicited larger contralateral P1 but a smaller contralateral N1 than invalid trials. The early onsets of these attentional effects show that spatial attention affects stimulus processing at early sensory/perceptual stages. The latencies of contralateral P1 and contralateral N1 were shorter for invalid trials, however. The ipsilateral N1 was enhanced by valid trials in the spatial frequency discrimination task but was not in the location discrimination task, whereas the contralateral N1 was larger for invalid trials than for valid trials in both tasks. CONCLUSION: The results indicate that involuntary allocation of attention involves different mechanisms from voluntary allocation of attention.     

Surround modulation of neuronal responses in V1 is as stable over time as responses to direct stimulation of receptive fields

In the primary visual cortex (V1) the modulation of neuronal responses by surround stimuli displays considerable variability. At present, it is not known whether this variability across neurons is due to temporal instability or to neuron-specific differences. We explored this question in the cat visual cortex by making multi-channel recordings while repeatedly presenting surround gratings of collinear and orthogonal orientation to the centre stimulus for a period of 96 h. Our results indicate that surround modulation is temporally stable to about the same degree as the responses evoked by the centre stimuli. The results support the notion that the mechanisms of surround modulation exhibit a high degree of stability and play an important role in the modulation of cortical responses.     

Aversive Conditioning of Spatial Position Sharpens Neural Population-Level Tuning in Visual Cortex and Selectively Alters Alpha-Band Activity

Processing capabilities for many low-level visual features are experientially malleable, aiding sighted organisms in adapting to dynamic environments. Explicit instructions to attend a specific visual field location influence retinotopic visuocortical activity, amplifying responses to stimuli appearing at cued spatial positions. It remains undetermined both how such prioritization affects surrounding nonprioritized locations, and if a given retinotopic spatial position can attain enhanced cortical representation through experience rather than instruction. The current report examined visuocortical response changes as human observers (N = 51, 19 male) learned, through differential classical conditioning, to associate specific screen locations with aversive outcomes. Using dense-array EEG and pupillometry, we tested the preregistered hypotheses of either sharpening or generalization around an aversively associated location following a single conditioning session. Competing hypotheses tested whether mean response changes would take the form of a Gaussian (generalization) or difference-of-Gaussian (sharpening) distribution over spatial positions, peaking at the viewing location paired with a noxious noise. Occipital 15 Hz steady-state visual evoked potential responses were selectively heightened when viewing aversively paired locations and displayed a nonlinear, difference-of-Gaussian profile across neighboring locations, consistent with suppressive surround modulation of nonprioritized positions. Measures of alpha-band (8–12 Hz) activity were differentially altered in anterior versus posterior locations, while pupil diameter exhibited selectively heightened responses to noise-paired locations but did not evince differences across the nonpaired locations. These results indicate that visuocortical spatial representations are sharpened in response to location-specific aversive conditioning, while top-down influences indexed by alpha-power reduction exhibit posterior generalization and anterior sharpening.SIGNIFICANCE STATEMENT It is increasingly recognized that early visual cortex is not a static processor of physical features, but is instead constantly shaped by perceptual experience. It remains unclear, however, to what extent the cortical representation of many fundamental features, including visual field location, is malleable by experience. Using EEG and an aversive classical conditioning paradigm, we observed sharpening of visuocortical responses to stimuli appearing at aversively associated locations along with location-selective facilitation of response systems indexed by pupil diameter and EEG alpha power. These findings highlight the experience-dependent flexibility of retinotopic spatial representations in visual cortex, opening avenues toward novel treatment targets in disorders of attention and spatial cognition.     

Spatial attention can modulate audiovisual integration at multiple cortical and subcortical sites

The role of attention in multisensory integration (MI) is presently uncertain, with some studies supporting an automatic, pre-attentive process and others suggesting possible modulation through selective attention. The goal of this functional magnetic resonance imaging study was to investigate the role of spatial attention on the processing of congruent audiovisual speech stimuli (here indexing MI). Subjects were presented with two simultaneous visual streams (speaking lips in the left and right visual hemifields) plus a single central audio stream (spoken words). In the selective attention conditions, the auditory stream was congruent with one of the two visual streams. Subjects attended to either the congruent or the incongruent visual stream, allowing the comparison of brain activity for attended vs. unattended MI while the amount of multisensory information in the environment and the overall attentional requirements were held constant. Meridian mapping and a lateralized 'speaking-lips' localizer were used to identify early visual areas and to localize regions responding to contralateral visual stimulations. Results showed that attention to the congruent audiovisual stimulus resulted in increased activation in the superior temporal sulcus, striate and extrastriate retinotopic visual cortex, and superior colliculus. These findings demonstrate that audiovisual integration and spatial attention jointly interact to influence activity in an extensive network of brain areas, including associative regions, early sensory-specific visual cortex and subcortical structures that together contribute to the perception of a fused audiovisual percept.     

A retinal source of spatial contrast gain control

Sensory cortex is able to encode a broad range of stimulus features despite a great variation in signal strength. In cat primary visual cortex (V1), for example, neurons are able to extract stimulus features like orientation or spatial configuration over a wide range of stimulus contrasts. The contrast-invariant spatial tuning found in V1 neuron responses has been modeled as a gain control mechanism, but at which stage of the visual pathway it emerges has remained unclear. Here we describe our findings that contrast-invariant spatial tuning occurs not only in the responses of lateral geniculate nucleus (LGN) relay cells but also in their afferent retinal input. Our evidence suggests that a similar contrast-invariant mechanism is found throughout the stages of the early visual pathway, and that the contrast-invariant spatial selectivity is evident in both retinal ganglion cell and LGN cell responses.     

Feature- and object-based attentional modulation in the human auditory "where" pathway

Attending to a visual stimulus feature, such as color or motion, enhances the processing of that feature in the visual cortex. Moreover, the processing of the attended object's other, unattended, features is also enhanced. Here, we used functional magnetic resonance imaging to show that attentional modulation in the auditory system may also exhibit such feature- and object-specific effects. Specifically, we found that attending to auditory motion increases activity in nonprimary motion-sensitive areas of the auditory cortical "where" pathway. Moreover, activity in these motion-sensitive areas was also increased when attention was directed to a moving rather than a stationary sound object, even when motion was not the attended feature. An analysis of effective connectivity revealed that the motion-specific attentional modulation was brought about by an increase in connectivity between the primary auditory cortex and nonprimary motion-sensitive areas, which, in turn, may have been mediated by the paracingulate cortex in the frontal lobe. The current results indicate that auditory attention can select both objects and features. The finding of feature-based attentional modulation implies that attending to one feature of a sound object does not necessarily entail an exhaustive processing of the object's unattended features.     

Neural correlates of covert orienting of visual spatial attention along vertical and horizontal dimensions

Covert orienting of spatial attention along the horizontal meridian of the visual field is mediated by a fronto-parietal neural network. The neural substrates underlying covert orienting of attention along the vertical meridian, however, are less understood. We recorded hemodynamic responses using functional magnetic resonance imaging (fMRI) from healthy volunteers in covert visual orienting tasks that required to detect targets either at the fixation or at peripheral attended locations on the horizontal or vertical meridian in the left (LVF), right (RVF), upper (UVF), and lower (LoVF) visual fields. We found that, relative to when attention was at the fixation, covert orienting of attention along the horizontal and vertical meridia induced enhanced activities in the superior parietal and frontal lobes bilaterally and the cerebellum. In addition, attention to the LoVF and UVF generated stronger activation in the medial frontal cortex, anterior cingulate, precuneus, and the cerebellum relative to attention along the horizontal meridian. The reversed contrast, however, produced stronger activation in the right lingual gyrus and right premotor cortex. The fMRI results suggest that, while a common neural network is engaged in guiding visual spatial attention along the vertical and horizontal dimensions, unique neural correlates are associated with covert attentional orienting along the vertical and horizontal meridia of the visual field.     

β-Band correlates of the fMRI BOLD response

Oscillatory activity in the β-band (15-30 Hz) has been studied in detail in the sensorimotor cortex. It has been postulated that β-activity acts as a localized gating of cortical activity. Here, the induced oscillatory response in the β-band is measured by magnetoencephalography, and the hemodynamic response is measured by fMRI. We assess the linearity of the responses to stimuli of varying duration in the primary motor cortex and to a sinusoidal drifting grating of varying contrast amplitude and drift frequency in the visual cortex. In this way, we explore the nature of β-oscillations and their relationship with hemodynamic effects. Excellent spatial colocalization of BOLD and β-activity in both central and lateral (MT) visual areas and sensorimotor areas suggests that the two are intimately related. In contrast to the BOLD response, the level of β-desynchronization is not modulated by stimulus contrast or by stimulus duration, consistent with a gating role. The amplitude of β-desynchronization in the central visual area is however modulated by drift frequency, and this seems to parallel the modulation in BOLD amplitude at the same location.