Extrastriate Visual Areas Research Articles

Retinal lesions induce fast intrinsic cortical plasticity in adult mouse visual system.

Neuronal activity plays an important role in the development and structural-functional maintenance of the brain as well as in its life-long plastic response to changes in sensory stimulation. We characterized the impact of unilateral 15° laser lesions in the temporal lower visual field of the retina, on visually driven neuronal activity in the afferent visual pathway of adult mice using insitu hybridization for the activity reporter gene zif268. In the first days post-lesion, we detected a discrete zone of reduced zif268 expression in the contralateral hemisphere, spanning the border between the monocular segment of the primary visual cortex (V1) with extrastriate visual area V2M. We could not detect a clear lesion projection zone (LPZ) in areas lateral to V1 whereas medial to V2M, agranular and granular retrosplenial cortex showed decreased zif268 levels over their full extent. All affected areas displayed a return to normal zif268 levels, and this was faster in higher order visual areas than in V1. The lesion did, however, induce a permanent LPZ in the retinorecipient layers of the superior colliculus. We identified a retinotopy-based intrinsic capacity of adult mouse visual cortex to recover from restricted vision loss, with recovery speed reflecting the areal cortical magnification factor. Our observations predict incomplete visual field representations for areas lateral to V1 vs. lack of retinotopic organization for areas medial to V2M. The validation of this mouse model paves the way for future interrogations of cortical region- and cell-type-specific contributions to functional recovery, up to microcircuit level.

Multimap formation in visual cortex.

An extrastriate visual area such as V2 or V4 contains neurons selective for a multitude of complex shapes, all sharing a common topographic organization. Simultaneously developing multiple interdigitated maps--hereafter a "multimap"--is challenging in that neurons must compete to generate a diversity of response types locally, while cooperating with their dispersed same-type neighbors to achieve uniform visual field coverage for their response type at all orientations, scales, etc. Previously proposed map development schemes have relied on smooth spatial interaction functions to establish both topography and columnar organization, but by locally homogenizing cells' response properties, local smoothing mechanisms effectively rule out multimap formation. We found in computer simulations that the key requirements for multimap development are that neurons are enabled for plasticity only within highly active regions of cortex designated "learning eligibility regions" (LERs), but within an LER, each cell's learning rate is determined only by its activity level with no dependence on location. We show that a hybrid developmental rule that combines spatial and activity-dependent learning criteria in this way successfully produces multimaps when the input stream contains multiple distinct feature types, or in the degenerate case of a single feature type, produces a V1-like map with "salt-and-pepper" structure. Our results support the hypothesis that cortical maps containing a fine mixture of different response types, whether in monkey extrastriate cortex, mouse V1 or elsewhere in the cortex, rather than signaling a breakdown of map formation mechanisms at the fine scale, are a product of a generic cortical developmental scheme designed to map cells with a diversity of response properties across a shared topographic space.

Involuntary attentional orienting in the absence of awareness speeds up early sensory processing

A long-standing controversy in the field of human neuroscience has revolved around the question whether attended stimuli are processed more rapidly compared to unattended stimuli. We conducted two event-related potential (ERP) experiments employing a temporal order judgment procedure in order to assess whether involuntary attention accelerates sensory processing, as indicated by latency modulations of early visual ERP components. A non-reportable exogenous cue could precede the first target with equal probability at the same (compatible) or opposite (incompatible) location. The use of non-reportable cues promoted automatic, bottom-up attentional capture, and ensured the elimination of any confounds related to the use of stimulus features that are common to both cue and target. Behavioral results confirmed involuntary exogenous orienting towards the unaware cue. ERP results showed that the N1pc, an electrophysiological measure of attentional orienting, was smaller and peaked earlier in compatible as opposed to incompatible trials, indicating cue-dependent changes in magnitude and speed of first target processing in extrastriate visual areas. Complementary Bayesian analysis confirmed the presence of this effect regardless of whether participants were actively looking for the cue (Experiment 1) or were not informed of it (Experiment 2), indicating purely automatic, stimulus-driven orienting mechanisms.

The selectivity of responses to red-green colour and achromatic contrast in the human visual cortex: an fMRI adaptation study.

There is controversy as to how responses to colour in the human brain are organized within the visual pathways. A key issue is whether there are modular pathways that respond selectively to colour or whether there are common neural substrates for both colour and achromatic (Ach) contrast. We used functional magnetic resonance imaging (fMRI) adaptation to investigate the responses of early and extrastriate visual areas to colour and Ach contrast. High‐contrast red–green (RG) and Ach sinewave rings (0.5 cycles/degree, 2 Hz) were used as both adapting stimuli and test stimuli in a block design. We found robust adaptation to RG or Ach contrast in all visual areas. Cross‐adaptation between RG and Ach contrast occurred in all areas indicating the presence of integrated, colour and Ach responses. Notably, we revealed contrasting trends for the two test stimuli. For the RG test, unselective processing (robust adaptation to both RG and Ach contrast) was most evident in the early visual areas (V1 and V2), but selective responses, revealed as greater adaptation between the same stimuli than cross‐adaptation between different stimuli, emerged in the ventral cortex, in V4 and VO in particular. For the Ach test, unselective responses were again most evident in early visual areas but Ach selectivity emerged in the dorsal cortex (V3a and hMT+). Our findings support a strong presence of integrated mechanisms for colour and Ach contrast across the visual hierarchy, with a progression towards selective processing in extrastriate visual areas.

Extrastriate Visual Areas Integrate Form Features over Space and Time to Construct Representations of Stationary and Rigidly Rotating Objects.

When an object moves behind a bush, for example, its visible fragments are revealed at different times and locations across the visual field. Nonetheless, a whole moving object is perceived. Unlike traditional modal and amodal completion mechanisms known to support spatial form integration when all parts of a stimulus are simultaneously visible, relatively little is known about the neural substrates of the spatiotemporal form integration (STFI) processes involved in generating coherent object representations from a succession visible fragments. We used fMRI to identify brain regions involved in two mechanisms supporting the representation of stationary and rigidly rotating objects whose form features are shown in succession: STFI and position updating. STFI allows past and present form cues to be integrated over space and time into a coherent object even when the object is not visible in any given frame. STFI can occur whether or not the object is moving. Position updating allows us to perceive a moving object, whether rigidly rotating or translating, even when its form features are revealed at different times and locations in space. Our results suggest that STFI is mediated by visual regions beyond V1 and V2. Moreover, although widespread cortical activation has been observed for other motion percepts derived solely from form-based analyses [Tse, P. U. Neural correlates of transformational apparent motion. Neuroimage, 31, 766-773, 2006; Krekelberg, B., Vatakis, A., & Kourtzi, Z. Implied motion from form in the human visual cortex. Journal of Neurophysiology, 94, 4373-4386, 2005], increased responses for the position updating that lead to rigidly rotating object representations were only observed in visual areas KO and possibly hMT+, indicating that this is a distinct and highly specialized type of processing.

Sound characteristics which affect attributes of the synaesthetic visual experience

Although research on synaesthesia has blossomed in the last decades, few things are known about inherited sound-colour synaesthesia and processes underlying synaesthetic perception. In this study we investigated how the auditory pitch, the frequency range, the complexity, as well as the structure of the sound affected the synaesthetic visual attributes of chroma, colour brightness, colour plurality and density. A distinct mapping was observed between sound features and visual characteristics, which suggests that visual stimulation requires sensory auditory processing. We propose that congenital synaesthetic perception involves two stages of processing: initially the sensory auditory information is processed by the primary auditory cortex, and then it is distributed to the secondary and extrastriate visual brain areas, generating visual stimulation. This process does not necessarily involve the participation of brain areas responsible for associative thinking.

The Anatomical and Functional Organization of the Human Visual Pulvinar.

The anatomical organization and basic response properties of the visual pulvinar have been extensively studied in nonhuman primates. Yet, relatively little is known about the functional and anatomical organization of the human pulvinar. Using neuroimaging, we found multiple representations of visual space within the ventral human pulvinar and extensive topographically organized connectivity with visual cortex. This organization is similar to other nonhuman primates and provides additional support that the general organization of the pulvinar is consistent across the primate phylogenetic tree. These results suggest that the human pulvinar, like other primates, is well positioned to regulate corticocortical communication.

Cortical Connections of the Caudal Portion of Posterior Parietal Cortex in Prosimian Galagos.

Posterior parietal cortex (PPC) of prosimian galagos includes a rostral portion (PPCr) where electrical stimulation evokes different classes of complex movements from different subregions, and a caudal portion (PPCc) where such stimulation fails to evoke movements in anesthetized preparations ( Stepniewska, Fang et al. 2009). We placed tracer injections into PPCc to reveal patterns of its cortical connections. There were widespread connections within PPCc as well as connections with PPCr and extrastriate visual areas, including V2 and V3. Weaker connections were with dorsal premotor cortex, and the frontal eye field. The connections of different parts of PPCc with visual areas were roughly retinotopic such that injections to dorsal PPCc labeled more neurons in the dorsal portions of visual areas, representing lower visual quadrant, and injections to ventral PPCc labeled more neurons in ventral portions of these visual areas, representing the upper visual quadrant. We conclude that much of the PPCc contains a crude representation of the contralateral visual hemifield, with inputs largely, but not exclusively, from higher-order visual areas that are considered part of the dorsal visuomotor processing stream. As in galagos, the caudal half of PPC was likely visual in early primates, with the rostral PPC half mediating sensorimotor functions.

Spatial specificity and inheritance of adaptation in human visual cortex.

Adaptation at early stages of sensory processing can be propagated to downstream areas. Such inherited adaptation is a potential confound for functional magnetic resonance imaging (fMRI) techniques that use selectivity of adaptation to infer neuronal selectivity. However, the relative contributions of inherited and intrinsic adaptation at higher cortical stages, and the impact of inherited adaptation on downstream processing, remain unclear. Using fMRI, we investigated how adaptation to visual motion direction and orientation influences visually evoked responses in human V1 and extrastriate visual areas. To dissociate inherited from intrinsic adaptation, we quantified the spatial specificity of adaptation for each visual area as a measure of the receptive field sizes of the area where adaptation originated, predicting that adaptation originating in V1 should be more spatially specific than adaptation intrinsic to extrastriate visual cortex. In most extrastriate visual areas, the spatial specificity of adaptation did not differ from that in V1, suggesting that adaptation originated in V1. Only in one extrastriate area—MT—was the spatial specificity of direction-selective adaptation significantly broader than in V1, consistent with a combination of inherited V1 adaptation and intrinsic MT adaptation. Moreover, inherited adaptation effects could be both facilitatory and suppressive. These results suggest that adaptation at early visual processing stages can have widespread and profound effects on responses in extrastriate visual areas, placing important constraints on the use of fMRI adaptation techniques, while also demonstrating a general experimental strategy for systematically dissociating inherited from intrinsic adaptation by fMRI.

Neural correlates of multiple object tracking strategies

Amazingly, human observers can track four independently moving targets. The present study investigated the neural correlates of multiple-object tracking (MOT). Based on previous work we used a modified MOT-task to which subjects exhibited different behaviors. One half of the subjects showed slower RTs and higher error rates with increasing correspondence between tracked items and a probe consisting of 4 highlighted items presented after the tracking. The other half of the subjects had better performance when the probe fully matched the tracked items. Here we sought to investigate the neural representation of the two divergent behavior types. Using multivariate pattern analysis we observed two partly overlapping functional networks associated with the different behaviors. Subjects that responded fast and accurate to full-congruity trials predominantly showed a functional pattern for the full-congruity condition that was very different from patterns associated with any of the partly congruent conditions. This “deviant” pattern was observed in frontal, parietal and extrastriate visual brain areas. In the group of subjects with decreasing performance for increasing target-probe congruity these same regions exhibited a very different functional relationship, in which increasing congruities were associated with linearly changing neural activity patterns. Early low-tier visual areas exclusively exhibited the linear classification pattern while area LO and the primary motor cortex exclusively showed the deviant pattern across all subjects. The coexistence of both networks in groups with different behaviors provides the neural basis for a flexible behavior that can be flexibly adjusted as a function of the strategy employed in the task.

Functional plasticity of the visual system in multiple sclerosis.

OPINION article Front. Neurol., 08 April 2015Sec. Multiple Sclerosis and Neuroimmunology Volume 6 - 2015 | https://doi.org/10.3389/fneur.2015.00079

Direct geniculo-extrastriate pathways: a review of the literature.

The aim of this paper is to review the literature on direct geniculo-extrastriate pathways with special attention to 3D extrastriate visual areas. A literature review was realized using PubMed and Google Scholar. "Lateral geniculate nucleus", "geniculo-extrastriate pathways" and "white matter fiber tracts" were among the keywords used. Existence of geniculo-extrastriate connections was first hypothesized by the clinical observations of Riddoch's syndrome in patients with striate cortex (primary visual area, V1) lesions. Robust histological proof of geniculo-extrastriate pathways exists in monkeys. In humans, these pathways were tested through functional magnetic resonance imaging (fMRI), electro- and magneto-physiological paradigms. Conversely, only indirect proof of the connection between lateral geniculate nucleus and V5 exists. To our knowledge there were not any anatomical studies of geniculo-extrastriate connections in humans. Few human studies take interest in geniculo-extrastriate pathways. Only indirect proof of geniculo-extrastriate pathways exists in humans.

The modulatory impact of reward and attention on global feature selection in human visual cortex

Reward powerfully influences human behaviour and perception, with reward effects being observed already on the level of basic sensory processing. Although reward-related modulations generally resemble those related to attentional selection, it is debated whether these effects indeed reflect the same selection operations. Here we focus on neuromagnetic indices of global colour-based attention in visual cortex, and ask whether reward elicits the same or separable underlying modulation effects. Observers performed a colour/orientation selection task where colour served to define the target as well as reward prospect. On each trial a target containing the target colour and one other colour was presented in the left visual field (VF) together with a bicoloured distractor in the right VF. Reward was delivered on correctly performed trials when the reward colour appeared in the target but not when it appeared in the distractor. The effect of global colour selection was assessed by comparing the brain response to the distractor depending on whether it contained the target colour, the reward colour, both, or neither. We observed that both the reward and target colour led to similar increases of the neuromagnetic response between ~200–260 ms originating from the same ventral extrastriate visual cortex areas, albeit slightly temporally lagged. Importantly, the response to the target and reward colour alone always added up to match the response size of their combined presentation. These results suggest that while reward and attention recruit the same global feature selection effects in extrastriate visual cortex, they are likely controlled by independent top-down influences.

Brain Activity in Response to Visual Symmetry

A number of studies have explored visual symmetry processing by measuring event related potentials and neural oscillatory activity. There is a sustained posterior negativity (SPN) related to the presence of symmetry. There is also functional magnetic resonance imaging (MRI) activity in extrastriate visual areas and in the lateral occipital complex. We summarise the evidence by answering six questions. (1) Is there an automatic and sustained response to symmetry in visual areas? Answer: Yes, and this suggests automatic processing of symmetry. (2) Which brain areas are involved in symmetry perception? Answer: There is an extended network from extrastriate areas to higher areas. (3) Is reflection special? Answer: Reflection is the optimal stimulus for a more general regularity-sensitive network. (4) Is the response to symmetry independent of view angle? Answer: When people classify patterns as symmetrical or random, the response to symmetry is view-invariant. When people attend to other dimensions, the network responds to residual regularity in the image. (5) How are brain rhythms in the two hemispheres altered during symmetry perception? Answer: Symmetry processing (rather than presence) produces more alpha desynchronization in the right posterior regions. Finally, (6) does symmetry processing produce positive affect? Answer: Not in the strongest sense, but behavioural measures reveal implicit positive evaluation of abstract symmetry.

Contextual modulation and stimulus selectivity in extrastriate cortex.

Contextual modulation is observed throughout the visual system, using techniques ranging from single-neuron recordings to behavioral experiments. Its role in generating feature selectivity within the retina and primary visual cortex has been extensively described in the literature. Here, we describe how similar computations can also elaborate feature selectivity in the extrastriate areas of both the dorsal and ventral streams of the primate visual system. We discuss recent work that makes use of normalization models to test specific roles for contextual modulation in visual cortex function. We suggest that contextual modulation renders neuronal populations more selective for naturalistic stimuli. Specifically, we discuss contextual modulation's role in processing optic flow in areas MT and MST and for representing naturally occurring curvature and contours in areas V4 and IT. We also describe how the circuitry that supports contextual modulation is robust to variations in overall input levels. Finally, we describe how this theory relates to other hypothesized roles for contextual modulation.

Beta oscillation dynamics in extrastriate cortex after removal of primary visual cortex.

The local field potential (LFP) in visual cortex is typically characterized by the following spectral pattern: before the onset of a visual stimulus, low-frequency oscillations (beta, 12-20 Hz) dominate, whereas during the presentation of a stimulus these oscillations diminish and are replaced by fluctuations at higher frequencies (gamma, >30 Hz). The origin of beta oscillations in vivo remains unclear, as is the basis of their suppression during visual stimulation. Here we investigate the contribution of ascending input from primary visual cortex (V1) to beta oscillation dynamics in extrastriate visual area V4 of behaving monkeys. We recorded LFP activity in V4 before and after resecting a portion of V1. After the surgery, the visually induced gamma LFP activity in the lesion projection zone of V4 was markedly reduced, consistent with previously reported spiking responses (Schmid et al., 2013). In the beta LFP range, the lesion had minimal effect on the normal pattern of spontaneous oscillations. However, the lesion led to a surprising and permanent reversal of the normal beta suppression during visual stimulation, with visual stimuli eliciting beta magnitude increases up to 50%, particularly in response to moving stimuli. This reversed beta activity pattern was specific to stimulus locations affected by the V1 lesion. Our results shed light on the mechanisms of beta activity in extrastriate visual cortex: The preserved spontaneous oscillations point to a generation mechanism independent of the geniculostriate pathway, whereas the positive beta responses support the contribution of visual information to V4 via direct thalamo-extrastriate projections.

On the Role of Suppression in Spatial Attention: Evidence from Negative BOLD in Human Subcortical and Cortical Structures

There is clear evidence that spatial attention increases neural responses to attended stimuli in extrastriate visual areas and, to a lesser degree, in earlier visual areas. Other evidence shows that neurons representing unattended locations can also be suppressed. However, the extent to which enhancement and suppression is observed, their stimulus dependence, and the stages of the visual system at which they are expressed remains poorly understood. Using fMRI we set out to characterize both the task and stimulus dependence of neural responses in the lateral geniculate nucleus (LGN), primary visual cortex (V1), and visual motion area (V5) in humans to determine where suppressive and facilitatory effects of spatial attention are expressed. Subjects viewed a lateralized drifting grating stimulus, presented at multiple stimulus contrasts, and performed one of three tasks designed to alter the spatial location of their attention. In retinotopic representations of the stimulus location, we observed increasing attention-dependent facilitation and decreasing dependence on stimulus contrast moving up the visual hierarchy from the LGN to V5. However, in the representations of unattended locations of the LGN and V1, we observed suppression, which was not significantly dependent on the attended stimulus contrast. These suppressive effects were also found in the pulvinar, which has been frequently associated with attention. We provide evidence, therefore, for a spatially selective suppressive mechanism that acts at a subcortical level.

Abnormal response to photic stimulation in juvenile myoclonic epilepsy: an EEG-fMRI study.

Juvenile myoclonic epilepsy (JME) is a young-onset electroclinical syndrome, characterized by myoclonic, generalized tonic-clonic, and possibly typical absence seizures. Interictal electroencephalography (EEG) displays 3-6 Hz spike/polyspike and wave pattern. Photosensitivity is common. Our aim was to explore the blood oxygen level-dependent (BOLD) response evoked by a highly provocative photic stimulus in a cohort of people with JME compared to a group of nonphotosensitive healthy controls, and to investigate the hemodynamic phenomena seen in patients with photosensitive JME. We studied 13 JME patients and 18 healthy controls using EEG-functional magnetic resonance imaging (fMRI) performed during low luminance intermittent photic stimulation (IPS). The BOLD response to IPS was investigated both in JME and control groups. In photosensitive JME subjects, we also performed a dynamic evaluation of BOLD signal changes evoked by the photoparoxysmal response (PPR) in a time frame ranging from 10 s before the onset of the EEG paroxysm up until 10 s afterward. The IPS evoked a positive BOLD response in striate and extrastriate visual areas, which was less in JME patients than in controls. Moreover, people with JME had a reduced positive BOLD response in the frontoparietal areas and putamen but a stronger negative BOLD response in the primary sensorimotor cortex (SM1) and in cortical regions belonging to the default mode network (DMN). In JME, the dynamic evaluation of BOLD signal changes related to PPR revealed an early positive response in the putamen and SM1, followed by BOLD signal decrements in the putamen, caudate nuclei, thalami, and SM1. Our results confirm the hypothesis that people with JME might have an altered interaction between the motor circuit and other neuronal networks, with prominent involvement of basal ganglia circuitry. The PPR could be a final expression of pathogenic phenomena occurring in the striato-thalamocortical system, possibly a core feature of system epilepsy JME.

Multisensory Response Modulation in the Superficial Layers of the Superior Colliculus

The mammalian superior colliculus (SC) is made up of seven distinct layers. Based on overall differences in neuronal morphology, afferent and efferent projection patterns, physiological properties, and presumptive behavioral role, the upper three layers have been classically grouped together as the superficial layers and the remaining four layers collectively make up the deep layers. Although the superficial layers receive their primary inputs from the retina and primary visual cortex, the deep layers receive inputs from extrastriate visual cortical areas and from auditory, somatosensory, and motor-related structures. In contrast, there is no evidence of monosynaptic nonvisual inputs to the superficial layers. However, more recent studies have revealed anatomical connections between the superficial and deep layers, thus providing the substrate for possible communication between these two functional divisions of the SC. In this study, we provide physiological evidence for auditory influences on visual responses in the superficial layers of the SC. Using extracellular recordings of local field potentials (LFPs) and multiunit activity, we demonstrate multisensory effects in the superficial layers of the cat SC such that subthreshold auditory activity (as seen in the LFP) modulates visual responses (reflected in spiking activity) when the two stimuli are presented together. These results have important implications for our understanding of the functional organization of the SC and for the neural basis of multisensory integration in general.

Neural Mechanisms Underlying Visual Short-Term Memory Gain for Temporally Distinct Objects

Recent research has shown that visual short-term memory (VSTM) can substantially be improved when the to-be-remembered objects are split in 2 half-arrays (i.e., sequenced) or the entire array is shown twice (i.e., repeated), rather than presented simultaneously. Here we investigate the hypothesis that sequencing and repeating displays overcomes attentional "bottlenecks" during simultaneous encoding. Using functional magnetic resonance imaging, we show that sequencing and repeating displays increased brain activation in extrastriate and primary visual areas, relative to simultaneous displays (Study 1). Passively viewing identical stimuli did not increase visual activation (Study 2), ruling out a physical confound. Importantly, areas of the frontoparietal attention network showed increased activation in repetition but not in sequential trials. This dissociation suggests that repeating a display increases attentional control by allowing attention to be reallocated in a second encoding episode. In contrast, sequencing the array poses fewer demands on control, with competition from nonattended objects being reduced by the half-arrays. This idea was corroborated by a third study in which we found optimal VSTM for sequential displays minimizing attentional demands. Importantly these results provide support within the same experimental paradigm for the role of stimulus-driven and top-down attentional control aspects of biased competition theory in setting constraints on VSTM.