Visuotopic Maps Research Articles

Wiring minimization of deep neural networks reveal conditions in which multiple visuotopic areas emerge

The visual system is characterized by multiple mirrored visuotopic maps, with each repetition corresponding to a different visual area. In this work we explore whether such visuotopic organization can emerge as a result of minimizing the total wire length between neurons connected in a deep hierarchical network. In particular we ask, given N neurons with a given connectivity and a 2-d grid with N locations, how will the neurons be placed on the grid such that the total distance between the connected neurons is minimized? This problem is an NP-hard combinatorial problem which we solve using simulated annealing. We first construct multi-layer feedforward hierarchical networks, and examine how different parameters of the network such as filter size, number of channels, and stride affect the placement of the neurons on the grid. By introducing visual input to the network we can visualize the resulting visuotopic organization on the 2-d grid. Our results show that networks with purely feedforward connectivity typically result in a single visuotopic map, and in certain cases no visuotopic map emerges. However, when we modify the network by introducing lateral connections, with sufficient lateral connectivity among neurons within layers, multiple visuotopic maps emerge, where some connectivity motifs yield mirrored alternations of visuotopic maps–a signature of biological visual system areas. These results demonstrate that different connectivity profiles have different emergent organizations under the minimum wiring hypothesis, and highlight that characterizing the large-scale spatial organizing of tuning properties in a biological system might also provide insights into the underlying connectivity.

Multiple Visuotopically Organized Subdivisions of the Lateral Pulvinar/Central Lateral Inferior Pulvinar Project into Thin and Thick Stripe Compartments of V2 in Macaques

Abstract The lateral and central lateral inferior pulvinar (PL/PIcl) of primates has been implicated in playing an important role in visual processing, but its physiological and anatomical characteristics remain to be elucidated. It has been suggested that there are two complete visuotopic maps in the PL/PIcl, each of which sends afferents into V2 and V4 in primates. Given that functionally distinct thin and thick stripes of V2 both receive inputs from the PL/PIcl, this raises the possibility of a presence of parallel segregated pathways within the PL/PIcl. To address this question, we selectively injected three types of retrograde tracers (CTB-488, CTB-555, and BDA) into thin or thick stripes in V2 and examined labeling in the PL/PIcl in macaques. As a result, we found that every cluster of retrograde labeling in the PL/PIcl included all three types of signals next to each other, suggesting that thin stripe– and thick stripe–projecting compartments are not segregated into domains. Unexpectedly, we found at least five topographically organized retrograde labeling clusters in the PL/PIcl, indicating the presence of more than two V2-projecting maps. Our results suggest that the PL/PIcl exhibits greater compartmentalization than previously thought. They may be functionally similar but participate in multiple cortico-pulvinar-cortical loops.

Triple visual hemifield maps in a case of optic chiasm hypoplasia

In humans, each hemisphere comprises an overlay of two visuotopic maps of the contralateral visual field, one from each eye. Is the capacity of the visual cortex limited to these two maps or are plastic mechanisms available to host more maps? We determined the cortical organization of the visual field maps in a rare individual with chiasma hypoplasia, where visual cortex plasticity is challenged to accommodate three hemifield maps. Using high-resolution fMRI at 7T and diffusion-weighted MRI at 3T, we found three hemiretinal inputs, instead of the normal two, to converge onto the left hemisphere. fMRI-based population receptive field mapping of the left V1–V3 at 3T revealed three superimposed hemifield representations in the left visual cortex, i.e. two representations of opposing visual hemifields from the left eye and one right hemifield representation from the right eye. We conclude that developmental plasticity including the re-wiring of local intra- and cortico-cortical connections is pivotal to support the coexistence and functioning of three hemifield maps within one hemisphere.

Effect of Suppression Width in Spontaneous Detection of Numerosity

Number sense is the spontaneous ability of human adults, infants, newborns, and other animals to process approximate numbers. However, how number sense can emerge in the absence of learning and fine-tuning has not been elucidated. On the basis of the treatment of visual pathways as an information processor, we developed two computational models for numerosity detection with single and dual suppression values. The information processor transforms the visual information of numerosities into input information of neurons in the parietal and prefrontal cortex via suppression and visuotopic mapping without the need for the construction of neural networks, training, and fine-tuning. In the single suppression model, the suppression width is a more fundamental determinant than numerical quantity in numerosity detection. Our dual suppression model clarifies the effect of suppression width values in two visual regions on spontaneous numerosity detection. It also predicts innate and acquired training behavior in humans and animals. Results suggest that the range of the lateral inhibition of neurons at successive stages in visual pathways is the neural basis of human number sense that ultimately gives rise to number theory and mathematics.

Müller Glial Cells Participate in Retinal Waves via Glutamate Transporters and AMPA Receptors

SUMMARYRetinal waves, the spontaneous patterned neural activities propagating among developing retinal ganglion cells (RGCs), instruct the activity-dependent refinement of visuotopic maps. Although it is known that the wave is initiated successively by amacrine cells and bipolar cells, the behavior and function of glia in retinal waves remain unclear. Using multiple in vivo methods in larval zebrafish, we found that Müller glial cells (MGCs) display wave-like spontaneous activities, which start at MGC processes within the inner plexiform layer, vertically spread to their somata and endfeet, and horizontally propagate into neighboring MGCs. MGC waves depend on glutamatergic signaling derived from bipolar cells. Moreover, MGCs express both glia-specific glutamate transporters and the AMPA subtype of glutamate receptors. The AMPA receptors mediate MGC calcium activities during retinal waves, whereas the glutamate transporters modulate the occurrence of retinal waves. Thus, MGCs can sense and regulate retinal waves via AMPA receptors and glutamate transporters, respectively.

Online LI-rTMS during a Visual Learning Task: Differential Impacts on Visual Circuit and Behavioral Plasticity in Adult Ephrin-A2A5-/- Mice.

Repetitive transcranial magnetic stimulation (rTMS) induces plasticity in normal and abnormal neural circuitries, an effect that may be influenced by intrinsic brain activity during treatment. Here, we study potential synergistic effects between low-intensity rTMS (LI-rTMS) and concurrent neural activity in promoting circuit reorganization and enhancing visual behavior. We used ephrin-A2A5–/– mice, which are known to possess visuotopic mapping errors that are ameliorated by LI-rTMS, and assessed the impact of stimulation when mice were engaged in a visual learning task. A detachable coil was affixed to each mouse, and animals underwent 2 wk of 10-min daily training in a two-choice visual discrimination task with concurrent LI-rTMS or sham stimulation. No-task controls (+LI-rTMS/sham) were placed in the task arena without visual task training. At the end of the experiment, visuomotor tracking behavior was assessed, and corticotectal and geniculocortical pathway organization was mapped by injections of fluorescent tracers into the primary visual cortex. Consistent with previous results, LI-rTMS alone improved geniculocortical and corticotectal topography, but combining LI-rTMS with the visual learning task prevented beneficial corticotectal reorganization and had no additional effect on geniculocortical topography or visuomotor tracking performance. Unexpectedly, there was a significant increase in the total number of trials completed by task + LI-rTMS mice in the visual learning task. Comparison with wild-type mice revealed that ephrin-A2A5–/– mice had reduced accuracy and response rates, suggesting a goal-directed behavioral deficit, which was improved by LI-rTMS. Our results suggest that concurrent brain activity during behavior interacts with LI-rTMS, altering behavior and different visual circuits in an abnormal system.

Reestablishing the Chemoarchitectural Borders Based on Electrophysiological and Connectivity Data.

In this chapter, we discuss the poor agreement between visuotopic maps described using electrophysiological and connectivity data and the subdivisions of the pulvinar based on chemoarchitecture. We focus on the differences and similarities between New and Old World monkeys to evaluate how this agreement evolved during evolution. There is some agreement in the localization of P1, described using electrophysiological and connectivity data, and the lateral and central portions of the nucleus pulvinaris inferior (PI), defined based on chemoarchitectural criteria. Similarly, there is some colocalization between P3 and the medial portion of PI in both New and Old World monkeys. One difference between primates refers to P2, which is present in the Old World macaque monkey but absent in the New World monkeys. P4, which has not been studied in all primates, shows a partial spatial agreement with the dorsal portion of the chemoarchitecturally defined PL.

Stereotyped initiation of retinal waves by bipolar cells via presynaptic NMDA autoreceptors

Glutamatergic retinal waves, the spontaneous patterned neural activities propagating among developing retinal ganglion cells (RGCs), instruct the activity-dependent refinement of visuotopic maps. However, its initiation and underlying mechanism remain largely elusive. Here using larval zebrafish and multiple in vivo approaches, we discover that bipolar cells (BCs) are responsible for the generation of glutamatergic retinal waves. The wave originates from BC axon terminals (ATs) and propagates laterally to nearby BCs and vertically to downstream RGCs and the optic tectum. Its initiation is triggered by the activation of and consequent glutamate release from BC ATs, and is mediated by the N-methyl-D-aspartate subtype of glutamate receptors (NMDARs) expressed at these ATs. Intercellular asymmetry of NMDAR expression at BC ATs enables the preferential initiation of waves at the temporal retina, where BC ATs express more NMDARs. Thus, our findings indicate that glutamatergic retinal waves are initiated by BCs through a presynaptic NMDA autoreceptor-dependent process.

Auditory Spatial Coding Flexibly Recruits Anterior, but Not Posterior, Visuotopic Parietal Cortex.

Audition and vision both convey spatial information about the environment, but much less is known about mechanisms of auditory spatial cognition than visual spatial cognition. Human cortex contains >20 visuospatial map representations but no reported auditory spatial maps. The intraparietal sulcus (IPS) contains several of these visuospatial maps, which support visuospatial attention and short-term memory (STM). Neuroimaging studies also demonstrate that parietal cortex is activated during auditory spatial attention and working memory tasks, but prior work has not demonstrated that auditory activation occurs within visual spatial maps in parietal cortex. Here, we report both cognitive and anatomical distinctions in the auditory recruitment of visuotopically mapped regions within the superior parietal lobule. An auditory spatial STM task recruited anterior visuotopic maps (IPS2–4, SPL1), but an auditory temporal STM task with equivalent stimuli failed to drive these regions significantly. Behavioral and eye-tracking measures rule out task difficulty and eye movement explanations. Neither auditory task recruited posterior regions IPS0 or IPS1, which appear to be exclusively visual. These findings support the hypothesis of multisensory spatial processing in the anterior, but not posterior, superior parietal lobule and demonstrate that recruitment of these maps depends on auditory task demands.

Spatial characteristics of evoked potentials elicited by a MEMS microelectrode array for suprachoroidal-transretinal stimulation in a rabbit

Suprachoroidal-transretinal stimulation (STS) can potentially restore vision. This study investigated the spatial characteristics of cortical electrical evoked potentials (EEPs) elicited by STS. A 4 × 4 thin-film platinum microelectrode stimulating array (200 μm electrode diameter and 400 μm center-to-center distance) was fabricated by a micro-electro-mechanical systems (MEMS) techniques and implanted into the suprachoroidal space of albino rabbits. The current threshold to elicit reliable EEPs by a single electrode was 41.6 ± 12.6 μA, corresponding to a 66.2 ± 20.1 μC · cm(-2) charge density per phase, which was lower than the reported safety limits. Spatially differentiated cortical responses could be evoked by STS through different rows or columns of electrical stimulation; furthermore, shifts in the location of the maximum cortical activities were consistent with cortical visuotopic maps; increasing the number of simultaneously stimulating electrodes increased the response amplitudes of EEPs and expanded the spatial spread as well. In addition, long-term implantation and electrical stimulation of the MEMS electrode array in suprachoroidal space are necessary to evaluate systematically the safety and biocompatibility of this approach. This study indicates that the STS approach by a MEMS-based platinum electrode array is a feasible alternative for visual restoration, and relatively high spatial discrimination may be achieved.

Cortical connective field estimates from resting state fMRI activity.

One way to study connectivity in visual cortical areas is by examining spontaneous neural activity. In the absence of visual input, such activity remains shaped by the underlying neural architecture and, presumably, may still reflect visuotopic organization. Here, we applied population connective field (CF) modeling to estimate the spatial profile of functional connectivity in the early visual cortex during resting state functional magnetic resonance imaging (RS-fMRI). This model-based analysis estimates the spatial integration between blood-oxygen level dependent (BOLD) signals in distinct cortical visual field maps using fMRI. Just as population receptive field (pRF) mapping predicts the collective neural activity in a voxel as a function of response selectivity to stimulus position in visual space, CF modeling predicts the activity of voxels in one visual area as a function of the aggregate activity in voxels in another visual area. In combination with pRF mapping, CF locations on the cortical surface can be interpreted in visual space, thus enabling reconstruction of visuotopic maps from resting state data. We demonstrate that V1 ➤ V2 and V1 ➤ V3 CF maps estimated from resting state fMRI data show visuotopic organization. Therefore, we conclude that—despite some variability in CF estimates between RS scans—neural properties such as CF maps and CF size can be derived from resting state data.

Intraparietal Sulcus Activity and Functional Connectivity Supporting Spatial Working Memory Manipulation

The intraparietal sulcus (IPS) is recruited during tasks requiring attention, maintenance and manipulation of information in working memory (WM). While WM tasks often show broad bilateral engagement along the IPS, topographic maps of contralateral (CL) visual space have been identified along the IPS, similar to retinotopic maps in visual cortex. In the present study, we asked how these visuotopic IPS regions are differentially involved in the maintenance and manipulation of spatial information in WM. Visuotopic mapping was performed in 26 participants to define regions of interest along the IPS, corresponding to previously described IPS0-4. In a separate task, we showed that while maintaining the location of a briefly flashed target in WM preferentially engaged CL IPS, manipulation of spatial information by mentally rotating the target around a circle engaged bilateral IPS, peaking in IPS1 in most participants. Functional connectivity analyses showed increased interaction between the IPS and prefrontal regions during manipulation, as well as interhemispheric interactions. Two control tasks demonstrated that covert attention shifts, and nonspatial manipulation (arithmetic), engaged patterns of IPS activation and connectivity that were distinct from WM manipulation. These findings add to our understanding of the role of IPS in spatial WM maintenance and manipulation.

Retinotopic maps in the pulvinar of bush baby (otolemur garnettii)

Despite its anatomical prominence, the function of the primate pulvinar is poorly understood. A few electrophysiological studies in simian primates have investigated the functional organization of pulvinar by examining visuotopic maps. Multiple visuotopic maps have been found for all studied simians, with differences in organization reported between New and Old World simians. Given that prosimians are considered closer to the common ancestors of New and Old World primates, we investigated the visuotopic organization of pulvinar in the prosimian bush baby (Otolemur garnettii). Single-electrode extracellular recording was used to find the retinotopic maps in the lateral (PL) and inferior (PI) pulvinar. Based on recordings across cases, a 3D model of the map was constructed. From sections stained for Nissl bodies, myelin, acetylcholinesterase, calbindin, or cytochrome oxidase, we identified three PI chemoarchitectonic subdivisions, lateral central (PIcl), medial central (PIcm), and medial (PIm) inferior pulvinar. Two major retinotopic maps were identified that cover PL and PIcl, the dorsal one in dorsal PL and the ventral one in PIcl and ventral PL. Both maps represent central vision at the posterior end of the border between the maps, the upper visual field in the lateral half and the lower visual field in the medial half. They share many features with the maps reported for the pulvinar of simians, including the location in pulvinar and the representation of the upper-lower and central-peripheral visual field axes. The second-order representation in the lateral map and a laminar organization are likely features specific to Old World simians.

The Retinotopy of a Prosimian (Bush Baby) Pulvinar

The pulvinar nucleus in primates has connections with all known visual cortices, yet little is known about its functions. To clarify the functional organization of pulvinar, a few studies have investigated its visuotopic maps in simians using electrophysiological methods. These studies consistently showed two maps, but the organization of these maps were reported to differ between the New and Old World simians. In this study we investigated the visuotopy of the prosimian bush baby (Otolemur garnettii) pulvinar, as prosimians are considered closer to the common ancestors of New and Old World primates. We used single electrode extracellular recording to determine the retinotopy of the lateral (PL) and inferior (PI) subdivisions of the pulvinar and compared the visuotopic maps to bush baby pulvinar chemoarchitecture. Three subdivisions, the lateral central (PIcl), medial central (PIcm) and medial (PIm) inferior pulvinar, were identified in bush baby PI, using myelin, acetylcholinesterase and cytochrome oxidase staining. We found two visuotopic maps which covered PL and PIcl, a dorsal map in dorsal PL and a ventral map in PIcl and ventral PL. Both maps had a lateral upper field representation and a medial lower field representation. Their central vision representations were found on the postero-medial side of the border between the two maps. The global features of the visuotopic maps, including location, orientation of the central-peripheral and upper-lower field axes, appeared similar between bush baby and the Old World macaque monkey but different from the New World cebus monkey. Using reconstructed 3D models of these maps, however, we found details that differed between bush baby and those reported in macaque monkey. These differences included: 1) the first order representation of the dorsal map, 2) the central vision representation as a point and 3) the vertical meridian representation on a curve in bush baby but not in macaque. Meeting abstract presented at VSS 2013

Visuotopic mapping of the parietal cortex distinguishes areas involved in synesthetic feature binding

Visuotopic mapping of the parietal cortex distinguishes areas involved in synesthetic feature binding

Attention maps in the brain.

Over 20 distinct cerebral cortical areas contain spatial map representations of the visual field. These retinotopic, or visuotopic, cortical areas occur not only in the occipital lobe but also in the parietal, temporal, and frontal lobes. The cognitive influences of visuospatial attention operate via these cortical maps and can support selection of multiple objects at the same time. In early visual cortical areas, spatial attention enhances responses of selected items and diminishes the responses to distracting items. In higher order cortex, the maps support a spatial indexing role, keeping track of the items to be attended. These maps also support visual short-term memory (VSTM) representations. In each hemisphere, all the known maps respond selectively to stimuli presented within the contralateral visual field. However, a hemispheric asymmetry emerges when the attentional or VSTM demands of a task become significant. In the parietal lobe, the right hemisphere visuotopic maps switch from coding only contralateral visual targets to coding memory and attention targets across the entire visual field. This emergent asymmetry has important implications for understanding hemispatial neglect syndrome, and supports a dynamic network form of the representational model of neglect. WIREs Cogn Sci 2013, 4:327-340. doi: 10.1002/wcs.1230 This article is categorized under: Psychology > Attention Neuroscience > Cognition.

Global workspace dynamics: cortical "binding and propagation" enables conscious contents.

A global workspace (GW) is a functional hub of binding and propagation in a population of loosely coupled signaling elements. In computational applications, GW architectures recruit many distributed, specialized agents to cooperate in resolving focal ambiguities. In the brain, conscious experiences may reflect a GW function. For animals, the natural world is full of unpredictable dangers and opportunities, suggesting a general adaptive pressure for brains to resolve focal ambiguities quickly and accurately. GW theory aims to understand the differences between conscious and unconscious brain events. In humans and related species the cortico-thalamic (C-T) core is believed to underlie conscious aspects of perception, thinking, learning, feelings of knowing (FOK), felt emotions, visual imagery, working memory, and executive control. Alternative theoretical perspectives are also discussed. The C-T core has many anatomical hubs, but conscious percepts are unitary and internally consistent at any given moment. Over time, conscious contents constitute a very large, open set. This suggests that a brain-based GW capacity cannot be localized in a single anatomical hub. Rather, it should be sought in a functional hub – a dynamic capacity for binding and propagation of neural signals over multiple task-related networks, a kind of neuronal cloud computing. In this view, conscious contents can arise in any region of the C-T core when multiple input streams settle on a winner-take-all equilibrium. The resulting conscious gestalt may ignite an any-to-many broadcast, lasting ∼100–200 ms, and trigger widespread adaptation in previously established networks. To account for the great range of conscious contents over time, the theory suggests an open repertoire of binding1 coalitions that can broadcast via theta/gamma or alpha/gamma phase coupling, like radio channels competing for a narrow frequency band. Conscious moments are thought to hold only 1–4 unrelated items; this small focal capacity may be the biological price to pay for global access. Visuotopic maps in cortex specialize in features like color, retinal size, motion, object identity, and egocentric/allocentric framing, so that a binding coalition for the sight of a rolling billiard ball in nearby space may resonate among activity maps of LGN, V1-V4, MT, IT, as well as the dorsal stream. Spatiotopic activity maps can bind into coherent gestalts using adaptive resonance (reentry). Single neurons can join a dominant coalition by phase tuning to regional oscillations in the 4–12 Hz range. Sensory percepts may bind and broadcast from posterior cortex, while non-sensory FOKs may involve prefrontal and frontotemporal areas. The anatomy and physiology of the hippocampal complex suggest a GW architecture as well. In the intact brain the hippocampal complex may support conscious event organization as well as episodic memory storage.

Auditory Spatial Attention Representations in the Human Cerebral Cortex

Auditory spatial attention serves important functions in auditory source separation and selection. Although auditory spatial attention mechanisms have been generally investigated, the neural substrates encoding spatial information acted on by attention have not been identified in the human neocortex. We performed functional magnetic resonance imaging experiments to identify cortical regions that support auditory spatial attention and to test 2 hypotheses regarding the coding of auditory spatial attention: 1) auditory spatial attention might recruit the visuospatial maps of the intraparietal sulcus (IPS) to create multimodal spatial attention maps; 2) auditory spatial information might be encoded without explicit cortical maps. We mapped visuotopic IPS regions in individual subjects and measured auditory spatial attention effects within these regions of interest. Contrary to the multimodal map hypothesis, we observed that auditory spatial attentional modulations spared the visuotopic maps of IPS; the parietal regions activated by auditory attention lacked map structure. However, multivoxel pattern analysis revealed that the superior temporal gyrus and the supramarginal gyrus contained significant information about the direction of spatial attention. These findings support the hypothesis that auditory spatial information is coded without a cortical map representation. Our findings suggest that audiospatial and visuospatial attention utilize distinctly different spatial coding schemes.

Evaluation of receptive field size from higher harmonics in visuotopic mapping using continuous stimulation optical imaging

The extraction and detection of specific responses from a large amount of background noise has been the subject of a considerable body of research in brain functional imaging, and more specifically in optical intrinsic signal imaging. Recent work by Kalatsky and Stryker (2003) showed that by combining different conditions and using periodic stimuli, recording times can be reduced. Spectral decomposition is then used to provide amplitude and phase information locked to the stimulus. A drawback of the above method is that by focusing only on a single harmonic, response information is limited. The shape of the hemodynamic response function (HRF) and the temporal variations in the neural responses cannot be assessed. In this work it is argued that additional information about neural responses can be gathered by using higher harmonics. Moving bars were used to generate visuotopic maps on large portions of the cat visual cortex. Up to four simultaneously bars moving repetitively across the visual field at different frequencies were used to sample the HRF in the Fourier domain. The HRF profile obtained with continuous stimulation was spatially homogeneous throughout the cortex and similar to the HRF profile obtained using episodic stimulation. Furthermore, by modeling the optical response as a convolution between HRF and neuronal responses, the ratio of the second harmonic to the first provided an estimation of the receptive field size. This was further validated by measuring spatial frequency selectivity. Therefore, the use of higher harmonics opens new avenues to estimate receptive field size from temporal signals.

Maps of Visual Space in Human Occipital Cortex Are Retinotopic, Not Spatiotopic

We experience the visual world as phenomenally invariant to eye position, but almost all cortical maps of visual space in monkeys use a retinotopic reference frame, that is, the cortical representation of a point in the visual world is different across eye positions. It was recently reported that human cortical area MT (unlike monkey MT) represents stimuli in a reference frame linked to the position of stimuli in space, a "spatiotopic" reference frame. We used visuotopic mapping with blood oxygen level-dependent functional magnetic resonance imaging signals to define 12 human visual cortical areas, and then determined whether the reference frame in each area was spatiotopic or retinotopic. We found that all 12 areas, including MT, represented stimuli in a retinotopic reference frame. Although there were patches of cortex in and around these visual areas that were ostensibly spatiotopic, none of these patches exhibited reliable stimulus-evoked responses. We conclude that the early, visuotopically organized visual cortical areas in the human brain (like their counterparts in the monkey brain) represent stimuli in a retinotopic reference frame.