Pre-attentive segmentation in the primary visual cortex

The activities of neurons in primary visual cortex have been shown to be significantly influenced by stimuli outside their classical receptive fields. We propose that these contextual influences serve pre-attentive visual segmentation by causing relatively higher neural responses to important or conspicuous image locations, making them more salient for perceptual pop-out. These locations include boundaries between regions, smooth contours, and pop-out targets against backgrounds. The mark of these locations is the breakdown of spatial homogeneity in the input. for instance, at the border between two texture regions of equal mean luminance. This breakdown causes changes in contextual influences, often resulting in higher responses at the border than at surrounding locations. This proposal is implemented in a biologically based model of VI in which contextual influences are mediated by intra-cortical horizontal connections. The behavior of the model is demonstrated using examples of texture segmentation, figure-ground segregation, target-distractor asymmetry, and contour enhancement, and is compared with psychophysical and physiological data. The model predicts (1) how neural responses should be tuned to the orientation of nearby texture borders, (2) a set of qualitative constraints on the structure of the intracortical connections, and (3) stimulus-dependent biases in estimating the locations of the region borders by pre-attentive vision.     

Hemodynamic traveling waves in human visual cortex

Functional MRI (fMRI) experiments rely on precise characterization of the blood oxygen level dependent (BOLD) signal. As the spatial resolution of fMRI reaches the sub-millimeter range, the need for quantitative modelling of spatiotemporal properties of this hemodynamic signal has become pressing. Here, we find that a detailed physiologically-based model of spatiotemporal BOLD responses predicts traveling waves with velocities and spatial ranges in empirically observable ranges. Two measurable parameters, related to physiology, characterize these waves: wave velocity and damping rate. To test these predictions, high-resolution fMRI data are acquired from subjects viewing discrete visual stimuli. Predictions and experiment show strong agreement, in particular confirming BOLD waves propagating for at least 5-10 mm across the cortical surface at speeds of 2-12 mm s-1. These observations enable fundamentally new approaches to fMRI analysis, crucial for fMRI data acquired at high spatial resolution.     

Sounds reset rhythms of visual cortex and corresponding human visual perception

An event in one sensory modality can phase reset brain oscillations concerning another modality. In principle, this may result in stimulus-locked periodicity in behavioral performance. Here we considered this possible cross-modal impact of a sound for one of the best-characterized rhythms arising from the visual system, namely occipital alpha-oscillations (8-14 Hz). We presented brief sounds and concurrently recorded electroencephalography (EEG) and/or probed visual cortex excitability (phosphene perception) through occipital transcranial magnetic stimulation (TMS). In a first, TMS-only experiment, phosphene perception rate against time postsound showed a periodic pattern cycling at ~10 Hz phase-aligned to the sound. In a second, combined TMS-EEG experiment, TMS-trials reproduced the cyclical phosphene pattern and revealed a ~10 Hz pattern also for EEG-derived measures of occipital cortex reactivity to the TMS pulses. Crucially, EEG-data from intermingled trials without TMS established cross-modal phase-locking of occipitoparietal alpha oscillations. These independently recorded variables, i.e., occipital cortex excitability and reactivity and EEG phase dynamics, were significantly correlated. This shows that cross-modal phase locking of oscillatory visual cortex activity can arise in the human brain to affect perceptual and EEG measures of visual processing in a cyclical manner, consistent with occipital alpha oscillations underlying a rapid cycling of neural excitability in visual areas.     

Retinotopic organization of visual cortex in human infants

1. Download : Download high-res image (281KB)
2. Download : Download full-size image

     

Early development of visual cortex in human fetuses

Prenatal development of visual cortex (area 17) was studied in human fetuses of 8-9, 13-15 and 16-18 weeks of gestation, with a view to analyse the early critical events. Under light microscope, five zones of development were seen in all the age groups. The total thickness of cortex of area 17 as well as that of its cortical plate was measured with the help of camera lucida. It was observed that the total thickness of the cortex increased with increase in age. Diversity in the shape, size, staining intensity and arrangement of neurons was noted in the different zones. Most of the cells were found to have a thin rim of cytoplasm and a prominent nucleus with multiple nucleoli. The cells in subventricular zone and cortical plate were regularly arranged in vertical rows while in other zones, they were irregularly scattered. Several mitotic figures were seen in the ventricular zone at 8-9 weeks but later they were also noticed in subventricular and intermediate zones. In the later ages the mitotic figures were observed to be fewer in the ventricular zone. No mitosis was seen in cortical plate at any age period.     

Contrast adaptation and representation in human early visual cortex

The human visual system can distinguish variations in image contrast over a much larger range than measurements of the static relationship between contrast and response in visual cortex would suggest. This discrepancy may be explained if adaptation serves to re-center contrast response functions around the ambient contrast, yet experiments on humans have yet to report such an effect. By using event-related fMRI and a data-driven analysis approach, we found that contrast response functions in V1, V2, and V3 shift to approximately center on the adapting contrast. Furthermore, we discovered that, unlike earlier areas, human V4 (hV4) responds positively to contrast changes, whether increments or decrements, suggesting that hV4 does not faithfully represent contrast, but instead responds to salient changes. These findings suggest that the visual system discounts slow uninformative changes in contrast with adaptation, yet remains exquisitely sensitive to changes that may signal important events in the environment.     

Activation of the human cortex during visual attention and selection

Cortical activation in visual discrimination tasks was estimated by measurement of the CNV (contingent negative variation) and N1-P3 components of visual ERPs in frontal, parietal, occipital and temporal leads recorded in 18 young healthy adults. In all investigated tasks, the maximal values of CNV and ERPa were observed in parietal regions. The estimation of cortical readiness state (CNV) is quite a useful procedure in the attention tasks because amplitude and stability of ERPs depend on preceding cortical excitability. The prevalence of parietal activation in visual attention tasks may be considered as the dominance of occipito-parietal way (stream) in human visual attention system.     

Population receptive field dynamics in human visual cortex

Seminal work in the early nineties revealed that the visual receptive field of neurons in cat primary visual cortex can change in location and size when artificial scotomas are applied. Recent work now suggests that these single neuron receptive field dynamics also pertain to the neuronal population receptive field (pRF) that can be measured in humans with functional magnetic resonance imaging (fMRI). To examine this further, we estimated the pRF in twelve healthy participants while masking the central portion of the visual field. We found that the pRF changes in location and size for two differently sized artificial scotomas, and that these pRF dynamics are most likely due to a combination of the neuronal receptive field position and size scatter as well as modulatory feedback signals from extrastriate visual areas.     

The representation of time course events in visual arts and the development of the concept of time in children: a preliminary study

By means of a careful search we found several representations of dynamic contents of events that show how the depiction of the passage of time in the visual arts has evolved gradually through a series of modifications and adaptations. The general hypothesis we started to investigate is that the evolution of the representation of the time course in visual arts is mirrored in the evolution of the concept of time in children, who, according to Piaget (1946), undergo three stages in their ability to conceptualize time. Crucial for our hypothesis is Stage II, in which children become progressively able to link the different phases of an event, but vacillate between what Piaget termed 'intuitive regulations', not being able to understand all the different aspects of a given situation. We found several pictorial representations - mainly dated back to the 14th to 15th century - that seem to fit within a Stage II of children's comprehension of time. According to our hypothesis, this type of pictorial representations should be immediately understood only by those children who are at Piaget's Stage II of time conceptualization. This implies that children at Stages I and III should not be able to understand the representation of time courses in the aforementioned paintings. An experiment was run to verify the agreement between children's collocation within Piaget's three stages - as indicated by an adaptation of Piaget's original experiment - and their understanding of pictorial representations that should be considered as Stage II type of representations of time courses. Despite the small sample of children examined so far, results seem to support our hypothesis. A follow-up (Experiment 2) on the same children was also run one year later in order to verify other possible explanations. Results from the two experiments suggest that the study of the visual arts can aid our understanding of the development of the concept of time, and it can also help to distinguish between the perceptual and the cognitive constraints (i.e. representational or cultural) in the representation of the succession of events.     

Cue-triggered activity replay in human early visual cortex

The recall of learned temporal sequences by a visual cue is an important form of experience-based neural plasticity. Here we observed such reactivation in awake human visual cortex using intracranial recording. After repeated exposure to a moving dot, a flash of the dot was able to trigger neural reactivation in the downstream receptive field along the motion path. This effect was observed only when the cue appeared near the receptive field. The estimated traveling speed was faster compared to the activation induced by the real motion. We suggest a range-limited, time-compressed reactivation as a result of repeated visual exposure in awake human visual cortex.     

Increased activity in human visual cortex during directed attention in the absence of visual stimulation

When subjects direct attention to a particular location in a visual scene, responses in the visual cortex to stimuli presented at that location are enhanced, and the suppressive influences of nearby distractors are reduced. What is the top-down signal that modulates the response to an attended versus an unattended stimulus? Here, we demonstrate increased activity related to attention in the absence of visual stimulation in extrastriate cortex when subjects covertly directed attention to a peripheral location expecting the onset of visual stimuli. Frontal and parietal areas showed a stronger signal increase during this expectation than did visual areas. The increased activity in visual cortex in the absence of visual stimulation may reflect a top-down bias of neural signals in favor of the attended location, which derives from a fronto-parietal network.     

Multiple spotlights of attentional selection in human visual cortex

Spatially directed attention strongly enhances visual perceptual processing. The metaphor of the "spotlight" has long been used to describe spatial attention; however, there has been considerable debate as to whether spatial attention must be unitary or may be split between discrete regions of space. This question was addressed here through functional MR imaging of human subjects as they performed a task that required simultaneous attention to two briefly displayed and masked targets at locations separated by distractor stimuli. These data reveal retinotopically specific enhanced activation in striate and extrastriate visual cortical representations of the two attended stimuli and no enhancement at the intervening representation of distractor stimuli. This finding of two spotlights was obtained within a single cortical hemisphere and across the two hemispheres. This provides direct evidence that spatial attention can select, in parallel, multiple low-level perceptual representations.     

Predicting the stream of consciousness from activity in human visual cortex

Can the rapid stream of conscious experience be predicted from brain activity alone? Recently, spatial patterns of activity in visual cortex have been successfully used to predict feature-specific stimulus representations for both visible and invisible stimuli. However, because these studies examined only the prediction of static and unchanging perceptual states during extended periods of stimulation, it remains unclear whether activity in early visual cortex can also predict the rapidly and spontaneously changing stream of consciousness. Here, we used binocular rivalry to induce frequent spontaneous and stochastic changes in conscious experience without any corresponding changes in sensory stimulation, while measuring brain activity with fMRI. Using information that was present in the multivariate pattern of responses to stimulus features, we could accurately predict, and therefore track, participants' conscious experience from the fMRI signal alone while it underwent many spontaneous changes. Prediction in primary visual cortex primarily reflected eye-based signals, whereas prediction in higher areas reflected the color of the percept. Furthermore, accurate prediction during binocular rivalry could be established with signals recorded during stable monocular viewing, showing that prediction generalized across viewing conditions and did not require or rely on motor responses. It is therefore possible to predict the dynamically changing time course of subjective experience with only brain activity.     

Temporal course of disinhibition of the visual cortex neurons and their sensitivity to cross-like figures

In acute experiments with narcotized and paralyzed cats, we studied responses of 74 striate neurons to cross-like figure under synchronous and asynchronous presentation of its lines. The aim of the study was to characterize the temporal course of interaction between three RF zones: main excitatory, end-inhibitory, and side disinhibitory ones. Previously we have found that this interaction is responsible for sensitivity to a cross in near 3/4 of cat striate cells with such sensitivity. In neurons with sensitivity to a cross, we found two types of temporal interaction between zones of RF. In the 1st type cells (14/23), the response significantly increased if the disinhibitory and the main excitatory zones of RF were stimulated simultaneously. Neurons of the 2nd type (9/23) revealed opposite temporal function: synchronous activation of RF zones evoked a minimal response. Simulation shows that the 1st type of behavior is connected with disinhibitory mechanism, while that of the 2nd type--with combination of this mechanism with convergence of orientation detectors of the previous functional level.     

Responses to lightness variations in early human visual cortex

Lightness is the apparent reflectance of a surface, and it depends not only on the actual luminance of the surface but also on the context in which the surface is viewed [1-10]. The cortical mechanisms of lightness processing are largely unknown, and the role of early cortical areas is still a matter of debate [11-17]. We studied the cortical responses to lightness variations in early stages of the human visual system with functional magnetic resonance imaging (fMRI) while observers were performing a demanding fixation task. The set of dynamically presented visual stimuli included the rectangular version of the classic Craik-O'Brien stimulus [3, 18, 19] and a variant that led to a weaker lightness effect, as well as a pattern with actual luminance variations. We found that the cortical activity in retinotopic areas, including the primary visual cortex (V1), is correlated with context-dependent lightness variations.     

Feedback contributions to visual awareness in human occipital cortex

It has traditionally been assumed that processing within the visual system proceeds in a bottom-up, feedforward manner from retina to higher cortical areas. In addition to feedforward processing, it is now clear that there are also important contributions to sensory encoding that rely upon top-down, feedback (reentrant) projections from higher visual areas to lower ones. By utilizing transcranial magnetic stimulation (TMS) in a metacontrast masking paradigm, we addressed whether feedback processes in early visual cortex play a role in visual awareness. We show that TMS of visual cortex, when timed to produce visual suppression of an annulus serving as a metacontrast mask, induces recovery of an otherwise imperceptible disk. In addition to producing disk recovery, TMS suppression of an annulus was greater when a disk preceded it than when an annulus was presented alone. This latter result suggests that there are effects of the disk on the perceptibility of the subsequent mask that are additive and are revealed with TMS of the visual cortex. These results demonstrate spatial and temporal interactions of conscious vision in visual cortex and suggest that a prior visual stimulus can influence subsequent perception at early stages of visual encoding via feedback projections.