Visual field and task influence illusory figure responses

In normal viewing conditions, many objects are often hidden or occluded by others, therefore restricting the information that enters the eye. One ability that the human visual system has developed to compensate for this visual limitation is to relate the surrounding elements to globally interpret the whole scene. The appearance of illusory figures (IF) based on surrounding elements also relies on this similar function. In the present study, we hypothesized that different mechanisms may be used by the brain to process IF from the center and periphery of the visual field. We compared magnetoencephalographic responses to IFs presented at different parts of the visual field under three task loads. For central presentation, IF specific responses peaked first in V1/V2 (96-101 ms), and then in the lateral occipital complex (LOC; 132-141 ms), independent of task. For peripheral presentation, the relative modulation towards IF was markedly reduced in V1/V2 and LOC while prominent activation peaks now shifted to the Fusiform Gyrus (from 200 ms onwards). Additionally, the type of task influenced processing at early stages beginning in V1/V2 (87 ms). Our results show that retinal eccentricity plays a crucial role in IF processing: figural completion at the center of the visual field is achieved in an 'automatic' and seemingly effortless fashion whereas peripheral stimulus locations necessitate higher-order object completion stages which rely more heavily on attentional demands.     

Response latencies of neurons in visual areas MT and MST of monkeys with striate cortex lesions

Cortical area, MT (middle temporal area) is specialized for the visual analysis of stimulus motion in the brain. It has been suggested [Brain 118 (1995) 1375] that motion signals reach area MT via two dissociable routes, namely a 'direct' route which bypasses primary visual cortex (area, striate cortex (V1)) and is specialized for processing 'fast' motion (defined as faster than 6 degrees/s) with a relatively short latency, and an 'indirect' route via area V1 for processing 'slow' motion (slower than 6 degrees/s) with a relatively long latency. We tested this proposal by measuring the effects of unilateral V1 lesions on the magnitudes and latencies of responses to fast- and slow-motion (depicted by random dot kinematograms (RDK) ) of single neurons in areas MT and medial superior temporal area (MST) of anaesthetized macaque monkeys. In the unlesioned hemisphere contralateral to a V1 lesion, response magnitudes and latencies of MT neurons were similar to those previously reported from MT neurons in normal monkeys, and there was no significant association between slow movement and long response latency (>100 ms), or between fast movement and short latency (< or =100 ms). V1 lesions led to diminished response magnitudes and increased latencies in area MT of the lesioned hemisphere, but did not selectively abolish MT responses to slow moving stimuli, or abolish long-latency responses to either slow- or fast-moving stimuli. Response magnitudes and latencies in area MST, which receives visual inputs directly from area MT and is also specialized for visual analysis of motion, were unaffected by V1 lesions (though we have shown elsewhere that directionally-selective responses in both areas were impaired by V1 lesions). Overall, the results are incompatible with the hypothesis that there are dissociable routes to MT specialized for processing separately fast and slow motion.     

Neural correlates of visual motion processing without awareness in patients with striate cortex and pulvinar lesions

Patients with striate cortex lesions experience visual perception loss in the contralateral visual field. In few patients, however, stimuli within the blind field can lead to unconscious (blindsight) or even conscious perception when the stimuli are moving (Riddoch syndrome). Using functional magnetic resonance imaging (fMRI), we investigated the neural responses elicited by motion stimulation in the sighted and blind visual fields of eight patients with lesions of the striate cortex. Importantly, repeated testing ensured that none of the patients exhibited blindsight or a Riddoch syndrome. Three patients had additional lesions in the ipsilesional pulvinar. For blind visual field stimulation, great care was given that the moving stimulus was precisely presented within the borders of the scotoma. In six of eight patients, the stimulation within the scotoma elicited hemodynamic activity in area human middle temporal (hMT) while no activity was observed within the ipsilateral lesioned area of the striate cortex. One of the two patients in whom no ipsilesional activity was observed had an extensive lesion including massive subcortical damage. The other patient had an additional focal lesion within the lateral inferior pulvinar. Fiber‐tracking based on anatomical and functional markers (hMT and Pulvinar) on individual diffusion tensor imaging (DTI) data from each patient revealed the structural integrity of subcortical pathways in all but the patient with the extensive subcortical lesion. These results provide clear evidence for the robustness of direct subcortical pathways from the pulvinar to area hMT in patients with striate cortex lesions and demonstrate that ipsilesional activity in area hMT is completely independent of conscious perception. Hum Brain Mapp 36:1585–1594, 2015. © 2014 Wiley Periodicals, Inc.     

Atypical organisation of the auditory cortex in dyslexia as revealed by MEG

Neuroanatomical and -radiological studies have converged to suggest an atypical organisation in the temporal bank of the left-hemispheric Sylvian fissure for dyslexia. Against the background of this finding, we applied high temporal resolution magnetoencephalography (MEG) to investigate functional aspects of the left-hemispheric auditory cortex in 11 right-handed dyslexic children (aged 8–13 years) and nine matched normal subjects (aged 8–14 years). Event-related field components during a passive oddball paradigm with pure tones and consonant–vowel syllables were evaluated. The first major peak of the auditory evoked response, the M80, showed identical topographical distributions in both groups. In contrast, the generating brain structures of the later M210 component were located more anterior to the earlier response in children with dyslexia only. Control children exhibited the expected activation of more posterior source locations of the component that appeared later in the processing stream. Since the group difference in the relative location of the M210 source seemed to be independent of stimulus category, it is concluded that dyslexics and normally literate children differ as to the organisation of their left-hemispheric auditory cortex.     

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.     

Topographic organization of cortical input to striate cortex in the Cebus monkey: a fluorescent tracer study

Cortical afferents to area V1 were studied in seven Cebus monkeys by means of retrograde fluorescent tracers. Injections were placed in V1, under electrophysiological guidance, in the regions of representation of both the upper and lower visual quadrants, at eccentricities that ranged from 0.5 to 64 degrees. In all cases retrogradely filled neurons were found in retinotopically corresponding portions of areas V2 and MT, as defined electrophysiologically (Rosa et al: J. Comp. Neurol. 275:326, 1988; Fiorani et al: J Comp Neurol 287:98, 1989). The results also revealed two other visual zones located anterior to V2 here named third and fourth visual areas. A topographical organization of the connections was observed in these areas, with upper quadrant located ventrally and lower quadrant located dorsally. A clear central-peripheral gradient, from the lateral to the medial cortical surface, was also observed in these areas. Lower field injections revealed crude topographic organization in area DZ and a diffuse projecting zone in the annectent gyrus. Peripheral injections in V1 revealed a clear upper and lower field segregation in areas PO and prostriata as well as a complex topography in MST. In addition, another region of labeling revealed the presence of an area, the temporal ventral posterior region, with an organized topographic representation of the upper field, with a central to peripheral gradient, from the lateral to the medial cortical surface. Three groups of cortical areas were distinguished according to the laminar distribution of neurons labeled from V1. In the first group, which is characterized by dense infra- and supragranular labeling, only V2 was included. The second group consists of areas V3, MT, and PO. These areas show dense labeling in the infragranular layers and occasionally sparse labeling in the supragranular layers. Finally, V4 and the other projecting areas, which are characterized by exclusive labeling of the infragranular layers were included in the third group.     

Neuromagnetic evidence for early semantic access in word recognition

Magnetic brain responses recorded in the human magnetoencephalogram (MEG) distinguished between words with different semantics but carefully matched for frequency and length. Multiple recordings from a single subject showed that 100 ms following stimulus onset, significantly stronger neuromagnetic responses were elicited by words with strong multimodal semantic associations than by other word material. At this early processing step, there was a highly significant correlation (0.80) between the magnitude of brain responses to individual words recorded over parieto-occipital areas and their semantic association strengths. Subsequent to this early difference related to word meaning, additional differences in MEG responses emerged for words from different grammatical categories. Together, these results suggest that word meaning can be reflected by early neuromagnetic brain responses and before the grammatical information about the word is encoded.     

A double dissociation between striate and extrastriate visual cortex for pattern motion perception revealed using rTMS

The neural mechanisms underlying the integration and segregation of motion signals are often studied using plaid stimuli. These stimuli consist of two spatially coincident dynamic gratings of differing orientations, which are either perceived to move in two unique directions or are integrated by the visual system to elicit the percept of a checkerboard moving in a single direction. Computations pertaining to the motion of the individual component gratings are thought to take place in striate cortex (V1) whereas motion integration is thought to involve neurons in dorsal stream extrastriate visual areas, particularly V5/MT. By combining a psychophysical task that employed plaid stimuli with 1 Hz offline repetitive transcranial magnetic stimulation (rTMS), we demonstrated a double dissociation between striate and extrastriate visual cortex in terms of their contributions to motion integration. rTMS over striate cortex increased coherent motion percepts whereas rTMS over extrastriate cortex had the opposite effect. These effects were robust directly after the stimulation administration and gradually returned to baseline within 15 minutes. This double dissociation is consistent with previous patient data and the recent hypothesis that both coherent and transparent motion percepts are supported by the visual system simultaneously and compete for perceptual dominance. Hum Brain Mapp 2009. © 2009 Wiley‐Liss, Inc.     

Attentional load modifies early activity in human primary visual cortex

Recent theories of selective attention assume that the more attention is required by a task, the earlier are irrelevant stimuli filtered during perceptual processing. Previous functional MRI studies have demonstrated that primary visual cortex (V1) activation by peripheral distractors is reduced by higher task difficulty at fixation, but it remains unknown whether such changes affect initial processing in V1 or subsequent feedback. Here we manipulated attentional load at fixation while recording peripheral visual responses with high-density EEG in 28 healthy volunteers, which allowed us to track the exact time course of attention-related effects on V1. Our results show a modulation of the earliest component of the visual evoked potential (C1) as a function of attentional load. Additional topographic and source localization analyses corroborated this finding, with significant load-related differences observed throughout the first 100 ms post-stimulus. However, this effect was observed only when stimuli were presented in the upper visual field (VF), but not for symmetrical positions in the lower VF. Our findings demonstrate early filtering of irrelevant information under increased attentional demands, thus supporting models that assume a flexible mechanism of attentional selection, but reveal important functional asymmetries across the VF.     

Unidirectional visual motion adaptation induces reciprocal inhibition of human early visual cortex excitability

ObjectivesBehavioural observations provided by the waterfall illusion suggest that motion perception is mediated by a comparison of responsiveness of directional selective neurones. These are proposed to be optimally tuned for motion detection in different directions. Critically however, despite the behavioural observations, direct evidence of this relationship at a cortical level in humans is lacking. By utilising the state dependant properties of transcranial magnetic stimulation (TMS), one can probe the excitability of specific neuronal populations using the perceptual phenomenon of phosphenes.MethodWe exposed subjects to unidirectional visual motion adaptation and subsequently simultaneously measured early visual cortex (V1) excitability whilst viewing motion in the adapted and non-adapted direction.ResultFollowing adaptation, the probability of perceiving a phosphene whilst viewing motion in the adapted direction was diminished reflecting a reduction in V1 excitability. Conversely, V1 excitability was enhanced whilst viewing motion in the opposite direction to that used for adaptation.ConclusionOur results provide support that in humans a process of reciprocal inhibition between oppositely tuned directionally selective neurones in V1 facilitates motion perception.SignificanceThis paradigm affords a unique opportunity to investigate changes in cortical excitability following peripheral vestibular disorders.     

Effects of visual deprivation and alterations in binocular competition on responses of striate cortex neurons in the cat

Electrophysiological recordings were made from single neurons in striate cortex of normally reared kittens (group N), kittens raised with binocular lid-suture (group BD), and kittens raised with one eye lid-sutured and the other eye removed (group MD-E). The MD-E group represents a condition in which inputs from the deprived eye have been placed at a competitive advantage over those from the other eye. In agreement with previous studies, fewer cells were responsive to visual stimulation in BD kittens than in N kittens. Among the responsive cells, fewer were direction selective, fewer were orientation selective, and more had inconsistent or fast-adapting responses than in normals. The responsiveness and receptive field properties of striate cortex neurons in the MD-E kittens were less affected by the visual deprivation than in BD kittens; however, they still were abnormal in comparison to normal kittens. Comparison of the ocular dominance distributions for cells in N and BD kittens showed a marked reduction in binocularly driven cells in BD kittens. In addition, in BD kittens, a larger proportion of monocularly driven cells had orientation selective receptive fields than did binocularly driven cells. This difference was not found in normally reared kittens. The results of this study suggest that abnormal binocular interactions contribute to the effects of visual deprivation following binocular lid-suture, probably due to asynchronous light-dark inputs through the closed lids. Removing the other eye and placing inputs from the deprived eye at a competitive advantage during development results in decreased effects on striate cortex neurons. Nevertheless, visual deprivation still produces abnormalities in striate cortex independent of asynchronous or uncorrelated visual stimulation of the two eyes.     

Effects of visual cortex lesions on perceptual grouping in rats

Neural mechanisms mediating perceptual grouping serve to enhance associations among stimulus elements, thereby establishing unified forms. The goals of the present study were to identify cortical areas necessary to perceptually group spatially isolated elements, and to determine if these areas are distinct from regions necessary for the discrimination of simple, solid forms. Rats were trained to discriminate horizontal and vertical lines that were either solid or composed of disjunct elements in which discrimination required perceptual grouping by proximity. Psychophysical procedures established the limits at which proximity served as a cue for grouping. Following perceptual measurements, ablations were made to selective sites within visual cortex. Lesions within area 17 or area 18A, including their interface, produced nearly complete impairment of solid line discrimination as well as perceptual grouping at all levels of proximity, whereas lesions to areas 18 or the far lateral extent of area 18A no effect on these perceptual capacities. These results indicate that grouping by proximity requires early visual processing areas, and shares cortical areas necessary for simple pattern discrimination. These results suggest that mechanisms of grouping modify primary stimulus representations, constructing a pattern of activity functional similar to that elicited by solid forms.     

Effects of attentional load on early visual processing depend on stimulus timing

A growing number of studies suggest that early visual processing is not only affected by low-level perceptual attributes but also by higher order cognitive factors such as attention or emotion. Using high-density electroencephalography, we recently demonstrated that attentional load of a task at fixation reduces the response of primary visual cortex to irrelevant peripheral stimuli, as indexed by the C1 component. In the latter study, peripheral stimuli were always presented during intervals without task-relevant stimuli. Here, we use a similar paradigm but present central task stimuli and irrelevant peripheral stimuli simultaneously while keeping all other stimulus characteristics constant. Results show that rather than to suppress responses to peripheral stimulation, high attentional load elicits higher C1 amplitudes under these conditions. These findings suggest that stimulus timing can profoundly alter the effects of attentional load on the earliest stages of processing in human visual cortex.     

Modeling direct effects of neural current on MRI

We investigate the effect of the magnetic field generated by neural activity on the magnitude and phase of the MRI signal in terms of a phenomenological parameter with the dimensions of length; it involves the product of the strength and duration of these currents. We obtain an analytic approximation to the MRI signal when the neuromagnetically induced phase is small inside the MRI voxel. The phase shift is the average of the MRI phase over the voxel, and therefore first order in that phase; and the reduction in the signal magnitude is one half the square of the standard deviation of the MRI phase, which is second order. The analytic approximation is compared with numerical simulations. For weak currents the agreement is excellent, and the magnitude change is generally much smaller than the phase shift. Using MEG data as a weak constraint on the current strength we find that for a net dipole moment of 10 nAm, a typical value for an evoked response, the reduction in the magnitude of the MRI signal is two parts in 10(5), and the maximum value of the overall phase shift is approximately 4 x 10(-3), obtained when the MRI voxel is displaced 2/3 the size of the neuronal activity. We also show signal changes over a large range of values of the net dipole moment. We compare these results with others in the literature. Our model overestimates the effect on the MRI signal.     

Intersubject variability of functional areas in the human visual cortex

Intersubject variability of striate and extrastriate areas was mapped by conjoined use of positron emission tomography (PET) and magnetic resonance imaging (MRI). We used two dynamic bowtie-shaped random-dot patterns centered symmetrically around the vertical- and horizontal-meridian, respectively, presented during sequential PET scans in 11 subjects. Control condition was simple fixation on a central dot in absence of a surrounding random dot pattern. V1, V2, VP, V3, V3a, V4, V5, and “wordform” areas were identified. After spatial normalization to Talairach atlas space, mean locations and standard deviations about these mean locations for x-, y-, and z-axes were calculated for each area in both hemispheres and compared for differences. The mean standard deviation for all axes across all areas tested was found to be small (4.9 mm). No significant differences were found in the mean standard deviations for the x-, y-, and z-axes in the left hemisphere vs. their counterparts in the right hemisphere. However, when mean standard deviations in both hemispheres were polled together by axis, the mean standard deviation for the y-axis (5.3 mm) was found to be significantly different from the mean standard deviation for the x-axis (4.3 mm). Furthermore, in the left hemisphere, the mean standard deviation for the z-axis (5.7 mm) was significantly greater than the mean standard deviation for the x-axis (3.9 mm). The values reported in this study for mean location and standard deviation of visual areas can be used to establish confidence intervals for distinguishing normal variations from pathology and consequent brain reorganization. Hum. Brain Mapping 6:301–315, 1998. © 1998 Wiley-Liss, Inc.     

Functional decoupling of BOLD and gamma-band amplitudes in human primary visual cortex

Although functional magnetic resonance imaging is an important tool for measuring brain activity, the hemodynamic blood oxygenation level dependent (BOLD) response is only an indirect measure of neuronal activity. Converging evidence obtained from simultaneous recording of hemodynamic and electrical measures suggest that the best correlate of the BOLD response in primary visual cortex is gamma-band oscillations ( approximately 40 Hz). Here, we examined the coupling between BOLD and gamma-band amplitudes measured with magntoencephalography (MEG) in human primary visual cortex in 10 participants. In Experiment A, participants were exposed to grating stimuli at two contrast levels and two spatial frequencies and in Experiment B square and sine wave stimuli at two spatial frequencies. The amplitudes of both gamma-band oscillations and BOLD showed tuning with stimulus contrast and stimulus type; however, gamma-band oscillations showed a 300% increase across two spatial frequencies, whereas BOLD exhibited no change. This functional decoupling demonstrates that increased amplitude of gamma-band oscillations as measured with MEG is not sufficient to drive the subsequent BOLD response.     

Effects of on quisqualate-stimulated phosphoinositide hydrolysis in slices of kitten striate cortex

Stimulation of phosphoinositide (PI) hydrolysis by excitatory amino acids (EAAs) was studied in coronal slices of kitten visual cortex. Coincubation with (NMDA) markedly reduced the stimulation by quisqualate, however, this inhibition developed with a latency of > 10 min and occurred even when the NMDA exposure preceded, but did not overlap with,incubation in quisqualate. This time-course of NMDA inhibition of EAA-stimulated P1 turnover places new constraints on its possible mechanism of inhibition.