The attentional 'spotlight's' penumbra: center-surround modulation in striate cortex

By enhancing neural activity in respective retinotopic cortical representations attention increases the efficiency with which visual information at a selected location is processed. Behavioral data also suggest that information from the vicinity of the attended region is actively suppressed. In search for a physiological correlate of this 'spotlight's penumbra' we assessed neural responses in retinotopic representations of an attended location and of locations at different distances to it. Relative to passive viewing we found suppressed striate activity for the nearby but not for the far locations. This attention-driven center-surround distribution of neural activity may enhance the contrast between attended and non-attended objects. We relate the different behavior of extrastriate areas to their lower spatial resolution, i.e. larger receptive fields.     

Luminance,but not chromatic visual pathways,mediate amplification of conditioned danger signals in human visual cortex

Complex organisms rely on experience to optimize the function of perceptual and motor systems in situations relevant to survival. It is well established that visual cues reliably paired with danger are processed more efficiently than neutral cues, and that such facilitated sensory processing extends to low levels of the visual system. The neurophysiological mechanisms mediating biased sensory processing, however, are not well understood. Here we used grating stimuli specifically designed to engage luminance or chromatic pathways of the human visual system in a differential classical conditioning paradigm. Behavioral ratings and visual electroencephalographic steady‐state potentials were recorded in healthy human participants. Our findings indicate that the visuocortical response to high‐spatial‐frequency isoluminant (red–green) grating stimuli was not modulated by fear conditioning, but low‐contrast, low‐spatial‐frequency reversal of grayscale gratings resulted in pronounced conditioning effects. We conclude that sensory input conducted via the chromatic pathways into retinotopic visual cortex has limited access to the bi‐directional connectivity with brain networks mediating the acquisition and expression of fear, such as the amygdaloid complex. Conversely, luminance information is necessary to establish amplification of learned danger signals in hierarchically early regions of the visual system.     

Developmental aspects of fear generalization – A MEG study on neurocognitive correlates in adolescents versus adults

BackgroundFear generalization is pivotal for the survival-promoting avoidance of potential danger, but, if too pronounced, it promotes pathological anxiety. Similar to adult patients with anxiety disorders, healthy children tend to show overgeneralized fear responses.ObjectiveThis study aims to investigate neuro-developmental aspects of fear generalization in adolescence – a critical age for the development of anxiety disorders.MethodsWe compared healthy adolescents (14–17 years) with healthy adults (19–34 years) regarding their fear responses towards tilted Gabor gratings (conditioned stimuli, CS; and slightly differently titled generalization stimuli, GS). In the conditioning phase, CS were paired (CS+) or remained unpaired (CS-) with an aversive stimulus (unconditioned stimuli, US). In the test phase, behavioral, peripheral and neural responses to CS and GS were captured by fear- and UCS expectancy ratings, a perceptual discrimination task, pupil dilation and source estimations of event-related magnetic fields.ResultsClosely resembling adults, adolescents showed robust generalization gradients of fear ratings, pupil dilation, and estimated neural source activity. However, in the UCS expectancy ratings, adolescents revealed shallower generalization gradients indicating overgeneralization. Moreover, adolescents showed stronger visual cortical activity after as compared to before conditioning to all stimuli.ConclusionVarious aspects of fear learning and generalization appear to be mature in healthy adolescents. Yet, cognitive aspects might show a slower course of development.     

Defensive engagement and perceptual enhancement

We tested whether visual cortical sensitivity to external cues in the context of an acute defensive reaction is heightened or attenuated. A strong cardiac defense (fear) response was elicited by presenting an abrupt, loud acoustic stimulus following a 10-min period of quiescence. Electrocortical responses to aversive and neutral pictures following defensive stimulus onset were measured using dense-array EEG. Pictures were flickered at 12.5 Hz to evoke steady-state visual evoked potentials (ssVEP), which can be reliably extracted on the basis of single trials. Visual cortical activity indexing perceptual processing was substantially heightened when pictures were shown in temporal proximity to (i.e., 5 s after) the defense stimulus. Replicating previous studies, aversive visual stimuli were associated with enhanced ssVEP amplitude, compared to neutral stimuli. Acute defense facilitates visual perception of external cues and preserves accurate discrimination between threatening and safe cues.     

Aversive conditioning of visual screening with aromatic ammonia for treating aggressive and disruptive behavior in a developmentally disabled child

This study investigated the usefulness of aversive conditioning of a mild punishment procedure that had ceased to suppress target behaviors to clinically acceptable levels. A 6.8-year-old severely retarded boy with high levels of aggressive, destructive, and loud screeching behaviors was readmitted to a psychiatric facility to adjust behavioral programs and rule out seizure disorder. A design combining withdrawal and multiple baselines across behaviors tested for effects of aversive conditioning of visual screening with ammonia and the impact of anticonvulsant medication (carbamazepine, phenytoin). Data indicated that aversively conditioned visual screening temporarily lowered rates of responding. It is tentatively concluded that aversive conditioning might be a useful mechanism to strengthen suppressive effects of mild punishment procedures and a promising approach for alleviating placement problems due to maladaptive behaviors in mentally retarded persons. Maintenance of treatment effects, community placement, and the mandate to choose the least restrictive aversive treatment alternative are discussed. Anticonvulsant medication showed no beneficial effect for the target behaviors investigated.     

Anxiety and spatial attention moderate the electrocortical response to aversive pictures

Aversive stimuli capture attention and elicit increased neural activity, as indexed by behavioral, electrocortical and hemodynamic measures; moreover, individual differences in anxiety relate to a further increased sensitivity to threatening stimuli. Evidence has been mixed, however, as to whether aversive pictures elicit increased neural response when presented in unattended spatial locations. In the current study, ERP and behavioral data were recorded from 49 participants as aversive and neutral pictures were simultaneously presented in spatially attended and unattended locations; on each trial, participants made same/different judgments about pictures presented in attended locations. Aversive images presented in unattended locations resulted in increased error rate and reaction time. The late positive potential (LPP) component of the ERP was only larger when aversive images were presented in attended locations, and this increase was positively correlated with self-reported state anxiety. Findings are discussed in regard to the sensitivity of ERP and behavioral responses to aversive distracters, and in terms of increased neural processing of threatening stimuli in anxiety.     

Attention and sensory gain control: a peripheral visual process?

Attention-related sensory gain control in human extrastriate cortex is believed to improve the acuity of visual perception. Yet given wide variance in the spatial resolution of vision across the retina, it remains unclear whether sensory gain operates homogenously between foveal and nonfoveal retinotopic locations. To address this issue, we used event-related potentials (ERPs) in a variant of the canonical spatial attention task. Participants were cued to expect targets at either fixation (foveal targets) or at a location several degrees above fixation (parafoveal targets). At both target locations, manual reaction times were shorter for cued relative to uncued targets, indicating that attention was consistently oriented to the cued location. Nevertheless, attention-related increases in sensory-evoked cortical activity were only observed at the parafoveal target location, as measured by the amplitude of the lateral occipital P1 ERP component. A second experiment replicated this data pattern using targets with lower stimulus contrast, indicating that the absence of a P1 effect for foveal targets could not be attributed to a saturated P1 response under higher-contrast stimulus conditions. When considered in light of retinogeniculate projections to cortex showing systematic changes in their physiological organization beginning within a degree of visual angle of the fovea, our findings support the proposal that the strategic functions of visual attention may vary with the retinotopic location involved.     

PKMζ maintains 1‐day‐ and 6‐day‐old long‐term object location but not object identity memory in dorsal hippocampus

Continuous activity of the atypical protein kinase C isoform M zeta (PKMζ) is necessary for maintaining long‐term memory acquired in aversively or appetitively motivated associative learning tasks, such as active avoidance, aversive taste conditioning, auditory and contextual fear conditioning, radial arm maze, and watermaze. Whether unreinforced, nonassociative memory will also require PKMζ for long‐term maintenance is not known. Using recognition memory for object location and object identity, we found that inactivating PKMζ in dorsal hippocampus abolishes 1‐day and 6‐day‐old long‐term recognition memory for object location, while recognition memory for object identity was not affected by this treatment. Memory for object location persisted for no more than 35 days after training. These results suggest that the dorsal hippocampus mediates long‐term memory for where, but not what things have been encountered, and that PKMζ maintains this type of spatial knowledge as long as the memory exists. © 2009 Wiley‐Liss, Inc.     

The eyes remember it: oculography and pupillometry during recollection in three amnesic patients

Two patients (TC and SS) with lesions that included the hippocampal regions (predominantly on the left side) were severely impaired in their recall of simple, verbally stated facts. However, both patients remembered spatial information that was temporally associated with semantic information. Specifically, TC and SS could not recall explicitly the content of an episode, but their spontaneous oculomotor behavior showed that they retained some information about the event as their gaze automatically returned to the locations on the computer screen where visual information had been paired to verbally presented information. Thus, this spatial information is implicit, automatically retrieved, and eye-based, as when one patient (TC) was asked to point with the finger to the same positions he was impaired. In addition, in an old/new recognition task, TC and SS and an additional patient, OB, showed significant changes in eye pupil diameter when viewing novel visual stimuli compared to stimuli that they had previously seen, also when they (incorrectly) declared with confidence that an old item was new. The spared memory of these patients, despite severe amnesia for the learning episodes, is characterized by a re-enactment of previous eye fixations that were associated with each (forgotten) episode and physiological responses (as indexed by pupillometry) to previously seen stimuli. Such spared memory can be seen as a type of "snapshot" memory, which automatically processes eye-based spatial information and whose content remains implicit. Finally, we surmise on the basis of the neuroanatomical findings of these patients, that neural substrates in the spared (right) hemisphere might support both the eye fixations' re-enactment and implicit visual pattern recognition.     

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.     

Source estimates for MEG/EEG visual evoked responses constrained by multiple, retinotopically-mapped stimulus locations

Studying the human visual system with high temporal resolution is a significant challenge due to the limitations of the available, noninvasive measurement tools. MEG and EEG provide the millisecond temporal resolution necessary for answering questions about intracortical communication involved in visual processing, but source estimation is ill-posed and unreliable when multiple; simultaneously active areas are located close together. To address this problem, we have developed a retinotopy-constrained source estimation method to calculate the time courses of activation in multiple visual areas. Source estimation was disambiguated by: (1) fixing MEG/EEG generator locations and orientations based on fMRI retinotopy and surface tessellations constructed from high-resolution MRI images; and (2) solving for many visual field locations simultaneously in MEG/EEG responses, assuming source current amplitudes to be constant or varying smoothly across the visual field. Because of these constraints on the solutions, estimated source waveforms become less sensitive to sensor noise or random errors in the specification of the retinotopic dipole models. We demonstrate the feasibility of this method and discuss future applications such as studying the timing of attentional modulation in individual visual areas.     

Spatial specificity of alpha oscillations in the human visual system

Alpha oscillations are strongly modulated by spatial attention. To what extent, the generators of cortical alpha oscillations are spatially distributed and have selectivity that can be related to retinotopic organization is a matter of continuous scientific debate. In the present report, neuromagnetic activity was quantified by means of spatial location tuning functions from 30 participants engaged in a visuospatial attention task. A cue presented briefly in one of 16 locations directing covert spatial attention resulted in a robust modulation of posterior alpha oscillations. The distribution of the alpha sources approximated the retinotopic organization of the human visual system known from hemodynamic studies. Better performance in terms of target identification was associated with a more spatially constrained alpha modulation. The present findings demonstrate that the generators of posterior alpha oscillations are retinotopically organized when modulated by spatial attention.     

Conscious and unconscious processing of fear after right amygdala damage: A single case ERP-study

In this study, we describe a 58-year-old male patient (FZ) with a right-amygdala lesion after temporal lobe infarction. FZ is unable to recognize fearful facial expressions. Instead, he consistently misinterprets expressions of fear for expressions of surprise. Employing EEG/ERP measures, we investigated whether presentation of fearful and surprised facial expressions would lead to different response patterns. We also measured ERPs to aversively conditioned and unconditioned fearful faces.

We compared ERPs elicited by supraliminally and subliminally presented conditioned fearful faces (CS+), unconditioned fearful faces (CS–) and surprised faces. Despite FZ's inability to recognize fearful facial expressions in emotion recognition tasks, ERP components showed different response patterns to pictures of surprised and fearful facial expressions, indicating that covert or implicit recognition of fear is still intact.

Differences between ERPs to CS+ and CS– were only found when these stimuli were presented subliminally. This indicates that intact right amygdala function is not necessary for aversive conditioning.

Previous studies have stressed the importance of the right amygdala for discriminating facial emotional expressions and for classical conditioning. Our study suggests that the right amygdala is necessary for explicit recognition of fear, while implicit recognition of fear and classical conditioning may still occur following lesion of the right amygdala.     

Cortical activation during Pavlovian fear conditioning depends on heart rate response patterns: an MEG study

In the present study, we examined stimulus-driven neuromagnetic activity in a delayed Pavlovian aversive conditioning paradigm using steady state visual evoked fields (SSVEF). Subjects showing an accelerative heart rate (HR) component to the CS+ during learning trials exhibited an increased activation in sensory and parietal cortex due to CS+ depiction in the extinction block. This was accompanied by a selective orientation response (OR) to the CS+ during extinction as indexed by HR deceleration. However, they did not show any differential cortical activation patterns during acquisition. In contrast, subjects not showing an accelerative HR component but rather unspecific HR changes during learning were characterized by greater activity in left orbito-frontal brain regions in the acquisition block but did not show differential SSVEF patterns during extinction. The results suggest that participants expressing different HR responses also differ in their stimulus-driven neuromagnetic response pattern to an aversively conditioned stimulus.     

Eye movements and the perceived location of phosphenes generated by intracranial primary visual cortex stimulation in the blind

BackgroundRestoring sight for the blind using electrical stimulation of the visual pathways is feasible but demands an understanding of the spatial mapping of the visual world at the site of targeted stimulation, whether in the retina, thalamus, or cortex. While a visual cortex stimulator can bypass the eye and create visual percepts, there is an inherent dissociation between this stimulation and eye movements. It is unknown whether and how robustly the brain maintains the oculomotor circuitry in patients with bare- or no-light perception.ObjectiveTo critically and quantitatively evaluate the effect of eye movements have on phosphene locations elicited by cortical stimulation that bypasses the eyes in order to restore sight in blind subjects.MethodsThe NeuroPace Responsive Neurostimulator (RNS) and the Orion visual cortical prosthesis devices were used to electrically stimulate the visual cortex of blind subjects with bare or no light perception. Eye positions were recorded synchronized with stimulation and the location of the percepts were measured using a handheld marker.ResultsThe locations of cortical stimulation-evoked percepts are shifted based on the eye position at the time of stimulation. Measured responses can be remapped based on measured eye positions to determine the retinotopic locations associated with the implanted electrodes, with remapped responses having variance limited by pointing error.ConclusionsEye movements dominate the perceived location of cortical stimulation-evoked phosphenes, even after years of blindness. By accounting for eye positions, we can mimic retinal mapping as in natural sight.     

Depersonalization disorder: thinking without feeling

Patients with depersonalization disorder (DP) experience a detachment from their own senses and surrounding events, as if they were outside observers. A particularly common symptom is emotional detachment from the surroundings. Using functional magnetic resonance imaging (fMRI), we compared neural responses to emotionally salient stimuli in DP patients, and in psychiatric and healthy control subjects. Six patients with DP, 10 with obsessive–compulsive disorder (OCD), and six volunteers were scanned whilst viewing standardized pictures of aversive and neutral scenes, matched for visual complexity. Pictures were then rated for emotional content. Both control groups rated aversive pictures as much more emotive, and demonstrated in response to these scenes significantly greater activation in regions important for disgust perception, the insula and occipito-temporal cortex, than DP patients (covarying for age, years of education and total extent of brain activation). In DP patients, aversive scenes activated the right ventral prefrontal cortex. The insula was activated only by neutral scenes in this group. Our findings indicate that a core phenomenon of depersonalization — absent subjective experience of emotion — is associated with reduced neural responses in emotion-sensitive regions, and increased responses in regions associated with emotion regulation.     

Saccades to a remembered location elicit spatially specific activation in human retinotopic visual cortex

The possible impact upon human visual cortex from saccades to remembered target locations was investigated using functional magnetic resonance imaging (fMRI). A specific location in the upper-right or upper-left visual quadrant served as the saccadic target. After a delay of 2,400 msec, an auditory signal indicated whether to execute a saccade to that location (go trial) or to cancel the saccade and remain centrally fixated (no-go). Group fMRI analysis revealed activation specific to the remembered target location for executed saccades, in the contralateral lingual gyrus. No-go trials produced similar, albeit significantly reduced, effects. Individual retinotopic mapping confirmed that on go trials, quadrant-specific activations arose in those parts of ventral V1, V2, and V3 that coded the target location for the saccade, whereas on no-go trials, only the corresponding parts of V2 and V3 were significantly activated. These results indicate that a spatial-motor saccadic task (i.e., making an eye movement to a remembered location) is sufficient to activate retinotopic visual cortex spatially corresponding to the target location, and that this activation is also present (though reduced) when no saccade is executed. We discuss the implications of finding that saccades to remembered locations can affect early visual cortex, not just those structures conventionally associated with eye movements, in relation to recent ideas about attention, spatial working memory, and the notion that recently activated representations can be "refreshed" when needed.     

Human Sensory Cortex Contributes to the Long-Term Storage of Aversive Conditioning

Growing animal data evince a critical role of the sensory cortex in the long-term storage of aversive conditioning, following acquisition and consolidation in the amygdala. Whether and how this function is conserved in the human sensory cortex is nonetheless unclear. We interrogated this question in a human aversive conditioning study using multidimensional assessments of conditioning and long-term (15 d) retention. Conditioned stimuli (CSs; Gabor patches) were calibrated to differentially activate the parvocellular (P) and magnocellular (M) visual pathways, further elucidating cortical versus subcortical mechanisms. Full-blown conditioning and long-term retention emerged for M-biased CS (vs limited effects for P-biased CS), especially among anxious individuals, in all four dimensions assessed: threat appraisal (threat ratings), physiological arousal (skin conductance response), perceptual learning [discrimination sensitivity (d′) and response speed], and cortical plasticity [visual evoked potentials (VEPs) and cortical current density]. Interestingly, while behavioral, physiological, and VEP effects were comparable at immediate and delayed assessments, the cortical substrates evolved markedly over time, transferring from high-order cortices [inferotemporal/fusiform cortex and orbitofrontal cortex (OFC)] immediately to the primary and secondary visual cortex after the delay. In sum, the contrast between P- and M-biased conditioning confirms privileged conditioning acquisition via the subcortical pathway while the immediate cortical plasticity lends credence to the triadic amygdala–OFC–fusiform network thought to underlie threat processing. Importantly, long-term retention of conditioning in the basic sensory cortices supports the conserved role of the human sensory cortex in the long-term storage of aversive conditioning.SIGNIFICANCE STATEMENT A growing network of neural substrates has been identified in threat learning and memory. The sensory cortex plays a key role in long-term threat memory in animals, but such a function in humans remains unclear. To explore this problem, we conducted multidimensional assessments of immediate and delayed (15 d) effects of human aversive conditioning. Behavioral, physiological, and scalp electrophysiological data demonstrated conditioning effects and long-term retention. High-density EEG intracranial source analysis further revealed the cortical underpinnings, implicating high-order cortices immediately and primary and secondary visual cortices after the long delay. Therefore, while high-order cortices support aversive conditioning acquisition (i.e., threat learning), the human sensory cortex (akin to the animal homolog) underpins long-term storage of conditioning (i.e., long-term threat memory).     

Spatial resolution of EEG cortical source imaging revealed by localization of retinotopic organization in human primary visual cortex

The aim of the present study is to investigate the spatial resolution of electroencephalography (EEG) cortical source imaging by localizing the retinotopic organization in the human primary visual cortex (V1). Retinotopic characteristics in V1 obtained from functional magnetic resonance imaging (fMRI) study were used as reference to assess the spatial resolution of EEG since fMRI can discriminate small changes in activation in visual field. It is well known that the activation of the early C1 component in the visual evoked potential (VEP) elicited by pattern onset stimuli coincides well with the activation in the striate cortex localized by fMRI. In the present experiments, we moved small circular checkerboard stimuli along horizontal meridian and compared the activations localized by EEG cortical source imaging with those from fMRI. Both fMRI and EEG cortical source imaging identified spatially correlated activity within V1 in each subject studied. The mean location error, between the fMRI-determined activation centers in V1 and the EEG source imaging activation peak estimated at equivalent C1 components (peak latency: 74.8+/-10.6 ms), was 7 mm (25% and 75% percentiles are 6.45 mm and 8.4 mm, respectively), which is less than the change in fMRI activation map by a 3 degrees visual field change (7.8 mm). Moreover, the source estimates at the earliest major VEP component showed statistically good correlation with those obtained by fMRI. The present results suggest that the spatial resolution of the EEG cortical source imaging can correctly discriminate cortical activation changes in V1 corresponding to less than 3 degrees visual field changes.     

From knowing what to knowing where: modeling object-based attention with feedback disinhibition of activation

We propose a neural model of visual object-based attention in which the identity of an object is used to select its location in an array of objects. The model is based on neural activity observed in visual search tasks performed by monkeys. In the model, the identity of the object (target) is selected in the higher areas of the ventral stream by means of a cue. Feedback activation from these higher areas carries information about the identity of the target to the (lower) retinotopic areas of the ventral stream. In these areas, the feedback activation interacts with feedforward activation produced by the object array. The interaction occurs in local microcircuits, and results in a selective activation on locations in the retinotopic areas of the visual stream that correspond to the location of the target in the object array. The selective activation consists of a form of gain control, produced by disinhibition. Transmitted to the dorsal stream, this activation directs spatial attention to the location of the target. In this way, an action directed at the target can be generated.