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Visual cortex

The visual cortex is the primary cortical region of the brain responsible for receiving, integrating, and processing visual information relayed from the retinas via the lateral geniculate nucleus (LGN) of the thalamus.^[1] Located predominantly in the occipital lobe at the posterior end of the cerebral cortex, it enables the perception of visual stimuli such as shapes, colors, motion, and depth through a hierarchical organization of specialized areas.^[1] This processing occurs unconsciously and rapidly, supporting higher-level functions like object recognition while segregating visual data into parallel streams for different attributes.^[1] The visual cortex is anatomically divided into multiple areas, with the primary visual cortex (V1, or striate cortex) serving as the initial entry point for visual signals in Brodmann area 17.^[2] V1 features a six-layered structure, where layer 4 receives direct afferents from the LGN's magnocellular (for motion and low-contrast detection), parvocellular (for form and color), and koniocellular (for blue-yellow color opponency) pathways.^[3] It is retinotopically organized, meaning neurons respond to specific locations in the visual field, with a disproportionate representation of the central (foveal) vision due to higher acuity needs.^[2] Functional columns within V1 include orientation columns, where adjacent neurons prefer similar edge orientations, and ocular dominance columns that segregate inputs from each eye.^[3] Beyond V1, the visual cortex encompasses extrastriate areas such as V2, V3, V4, and V5 (also known as MT), each contributing to increasingly complex analyses.^[1] V2 processes more integrated features like color boundaries and spatial frequencies, while V4 specializes in color constancy and form; V5 focuses on motion direction and speed.^[1] These areas form two major processing streams: the dorsal stream ("where" pathway), projecting to the parietal lobe for spatial awareness and action guidance, and the ventral stream ("what" pathway), extending to the temporal lobe for object identification and recognition.^[2] Feedback connections from higher areas refine V1 processing, enhancing contrast and attention.^[3] Damage to the visual cortex, often from stroke or trauma, can result in cortical blindness, where patients lose conscious vision despite intact eyes and pathways, or specific deficits like achromatopsia (color blindness) from V4 lesions or akinetopsia (motion blindness) from V5 damage.^[1] The right visual cortex primarily processes the left visual field, and vice versa, due to contralateral organization.^[1] Blood supply derives from branches of the posterior cerebral artery, making it vulnerable during posterior circulation events.^[1]

Overview

Definition and Role

The visual cortex refers to the specialized regions of the cerebral cortex in the occipital lobe that receive, integrate, and process visual information relayed from the retinas via the lateral geniculate nucleus (LGN) of the thalamus.^[1]^[4] These areas transform raw sensory signals into meaningful perceptual representations, enabling the detection and interpretation of visual stimuli such as shapes, colors, and movements. In humans, the visual cortex encompasses multiple interconnected regions, collectively occupying a substantial portion of the neocortex and containing an estimated 4-6 billion neurons dedicated to visual processing.^[5] The primary roles of the visual cortex involve initial feature detection, where neurons respond selectively to basic elements like edges and orientations, as demonstrated in seminal studies on cortical receptive fields. This early processing segments visual input into components, followed by integration across regions to form coherent scenes and contribute to visual awareness.^[6] Through hierarchical computations, the visual cortex bridges raw retinal input to higher-level perception, supporting object recognition and spatial understanding without relying on conscious effort for basic operations. Evolutionarily, the visual cortex derives from ancient vertebrate brain structures, with conserved functions for visual processing observed across mammals, reflecting adaptations for enhanced environmental navigation.^[7] Recent findings highlight interactions with subcortical areas like the superior colliculus, which provide primitive, radar-like saliency detection that modulates cortical responses and influences early visual function.^[8]^[9] These subcortical inputs underscore the visual cortex's role in a multimodal network, blending ancient reflexive mechanisms with advanced cortical elaboration.

Location and Inputs

The visual cortex is primarily located in the occipital lobe of the cerebral cortex, with the primary visual cortex (V1) situated along the banks of the calcarine sulcus on the medial surface of the occipital lobe.^[10] This region extends posteriorly toward the occipital pole and receives input corresponding to the contralateral visual field, forming a bilateral structure where the left visual cortex processes the right hemifield and vice versa.^[1] Higher visual areas project beyond the occipital lobe into the parietal and temporal lobes, enabling integration with sensory and associative functions.^[11] Visual inputs to the cortex are relayed primarily through the lateral geniculate nucleus (LGN) of the thalamus, which receives fibers from the optic tract originating in retinal ganglion cells.^[2] The LGN segregates inputs into magnocellular layers, which convey information about motion and depth via large, fast-conducting axons, and parvocellular layers, which transmit details on color and fine form through smaller, slower pathways.^[12] From the LGN, geniculocalcarine fibers form the optic radiations, which traverse the temporal and parietal lobes before terminating in layer 4 of V1, establishing the main afferent pathway for conscious visual perception.^[13] Additional subcortical inputs arrive from the superior colliculus via thalamic relays, such as the pulvinar nucleus, supporting rapid orienting responses to salient stimuli outside the primary thalamocortical route.^[2]^[14] These pathways also incorporate feedback projections from higher cortical areas, modulating early sensory input.^[12] The visual cortex exhibits hemispheric asymmetries, with the right hemisphere showing a bias toward global processing of visual scenes, such as overall configuration and spatial relations, while the left hemisphere favors local processing of fine details and features.^[15] This specialization arises from differential connectivity and attentional mechanisms, influencing how visual information from the contralateral hemifield is integrated.^[16]

Structural Organization

Cytoarchitecture and Layers

The visual cortex, as a neocortical region, exhibits a characteristic six-layered cytoarchitecture that underpins its role in visual processing. This laminar organization is evident across primary (V1) and secondary (V2) visual areas, with layers I through VI varying in thickness, cell density, and connectivity to support segregated input, integration, and output pathways. Layer I, the molecular layer, is relatively cell-sparse and rich in apical dendrites from deeper pyramidal neurons, facilitating diffuse modulation. Layer II (external granular) and Layer III (external pyramidal) are densely packed with small pyramidal cells and interneurons, enabling local intracortical connections. Layer IV (internal granular), the primary recipient of thalamic afferents, is notably thick in visual areas and subdivided into sublayers that segregate inputs from the lateral geniculate nucleus (LGN). Layer V (internal pyramidal) contains large pyramidal neurons that project to subcortical targets, while Layer VI (multiform) provides feedback to thalamic nuclei.^[1]^[17] In V1 (Brodmann area 17), Layer IV is particularly prominent and divided into sublayers 4A, 4B, 4Cα, and 4Cβ, which receive segregated thalamic inputs from the LGN's parvocellular (to 4Cβ, emphasizing color and fine detail), magnocellular (to 4Cα, prioritizing motion and low-contrast forms), and koniocellular (to 4A and parts of Layers II/III) pathways. Layers II/III in V1 integrate these inputs through horizontal connections among pyramidal neurons, forming the basis for feature convergence. Outputs originate from Layer V to structures like the superior colliculus and pons, and from Layer VI back to the LGN for modulation. This striate cortex is distinguished by the prominent "line of Gennari," a myelinated band in Layer IV visible macroscopically due to dense geniculocortical afferents, which defines its cytoarchitectonic boundary.^[17]^[18]^[1] Extrastriate areas like V2 show variations in this laminar pattern, with overall thicker Layers II/III and V for enhanced integration of V1 outputs, and a lack of the line of Gennari. V2's Layer IV is less subdivided but receives direct LGN inputs alongside dense V1 terminations, supporting its role in binding features across visual fields; its cytoarchitecture includes cytochrome oxidase (CO)-dense stripes (thin, pale, thick) that align with functional modules. These structural differences reflect V2's transitional role in processing.^[17]^[19] The cellular composition of these layers includes excitatory pyramidal neurons, which predominate (about 80% of neurons) and form the core of feedforward and feedback circuits via glutamatergic synapses, with morphologies varying by layer—e.g., spiny stellate cells in Layer IV for thalamic relay. Inhibitory interneurons, comprising the remainder, include basket cells (parvalbumin-positive, targeting pyramidal somata for perisomatic inhibition to sharpen responses) and other subtypes like chandelier and Martinotti cells, providing GABAergic control to prevent overexcitation. Recent transcriptomic studies reveal continuous diversification of these cell types during development, driven by spatiotemporal progenitor origins (e.g., medial vs. caudal ganglionic eminence for PV+ vs. SST+ interneurons) and activity-dependent refinement, resulting in over 20 molecularly distinct subtypes in visual cortex by maturity.^[1]^[20]^[21] This layer-specific cytoarchitecture enables parallel processing channels in the visual cortex, where thalamic inputs to Layer IV initiate segregated streams (e.g., magnocellular for coarse features via 4Cα), intracortical elaboration in Layers II/III builds complexity, and descending layers handle efferent signaling, collectively supporting efficient feature extraction while maintaining retinotopic mapping.^[17]^[18]

Retinotopic Organization

Retinotopy refers to the topographic mapping of the visual field onto the visual cortex, where neighboring points on the retina project to neighboring neurons in the cortex, preserving spatial relationships from the retinal input. This point-to-point correspondence ensures that the layout of the visual scene is maintained across multiple stages of visual processing, beginning in the primary visual cortex (V1). A key feature of this organization is foveal magnification, whereby the central portion of the visual field, corresponding to the fovea, is represented by a disproportionately larger area of cortical tissue compared to the periphery; this allocation supports higher visual acuity in central vision, as the fovea contains a higher density of photoreceptors.^[22] In V1, retinotopic maps are organized along dimensions of polar angle and eccentricity. The representation of polar angle follows a systematic progression around the cortical surface, with the vertical meridian—separating the left and right visual hemifields—aligning along the borders of V1 and adjacent areas, often positioned at the calcarine fissure in the occipital lobe. Eccentricity, or distance from the fovea, increases progressively from the foveal representation near the posterior pole of the occipital lobe to peripheral representations more anteriorly. These maps can be precisely delineated in humans using phase-encoded functional magnetic resonance imaging (fMRI) techniques, where rotating wedge stimuli encode polar angle and expanding ring stimuli encode eccentricity, producing traveling waves of activation that reveal the topographic layout.^[23]^[24] The cortical magnification factor (MF) quantifies how the size of cortical representation scales with retinal eccentricity, reflecting the nonlinear allocation of neural resources. It is commonly modeled as

\text{MF} = \frac{k}{r^{\alpha}}

where r is the retinal eccentricity in degrees, k is a constant, and \alpha \approx 0.8 - 1.0; this results in a higher MF near the fovea, where receptive fields are smaller and more numerous to match the demands of fine spatial resolution. For example, in the human V1, the MF can exceed 10 mm/degree near the fovea but drops sharply in the periphery, emphasizing central vision processing.^[22] Recent research highlights distortions and adaptations in retinotopic organization following injury, such as occipital lobe infarction, where affected areas like V1 and V2 show initial shrinkage but subsequent expansion and remapping over months, with shifts in the center of mass of visual field representations up to several millimeters. In models of visual deprivation, such as prolonged darkness followed by light exposure, initial disorganized maps in V1 and downstream areas undergo topological reorganization, with population receptive fields sharpening and aligning to restore stability through network-level plasticity. These adaptations demonstrate the visual cortex's capacity for functional recovery, though retinotopic stability can vary based on the extent of damage and rehabilitation.^[25]^[26]

Primary Visual Areas

Primary Visual Cortex (V1)

The primary visual cortex, also known as the striate cortex or area V1, is the initial cortical stage for processing visual information, receiving the majority of its afferent input from the lateral geniculate nucleus (LGN) of the thalamus.^[27] This input primarily terminates in layer 4, where geniculocortical axons form synapses with spiny stellate and pyramidal neurons. V1 exhibits a columnar organization, characterized by ocular dominance columns approximately 500 μm wide, which segregate inputs from the two eyes, and orientation columns that group neurons selective for similar stimulus orientations, forming functional hypercolumns roughly 1 mm in diameter.^[28] These columns enable parallel processing of monocular and orientation-specific features across the cortical surface.^[29] Neurons in V1 perform basic feature detection, including edge and orientation selectivity, as described in the Hubel-Wiesel model, which distinguishes simple cells with phase-specific receptive fields from complex cells that respond to oriented stimuli regardless of precise position. Simple cells typically exhibit elongated receptive fields with excitatory and inhibitory subregions aligned along a preferred orientation, while complex cells integrate inputs from multiple simple cells to achieve broader spatial invariance. V1 also supports binocular integration, where many neurons receive convergent inputs from both eyes, facilitating disparity tuning and stereopsis. Additionally, V1 neurons display tuning for contrast and spatial frequency, with preferred frequencies often peaking at 1-4 cycles per degree, allowing detection of luminance variations and fine spatial details.^[30] Receptive fields in V1 build upon the center-surround organization inherited from LGN afferents but elaborate into more complex forms, such as end-stopped cells that respond optimally to line segments or corners but are inhibited by longer contours, aiding in the detection of line endings and curvature. Computationally, these orientation-selective receptive fields are well-modeled by Gabor filters, which combine sinusoidal gratings with Gaussian envelopes to capture both spatial frequency and orientation preferences observed in V1 neurons.^[31] V1 maintains a retinotopic map of the visual field, with the fovea represented in a disproportionately large cortical area.^[12] Lesions to V1 typically result in contralateral visual field deficits, such as hemianopia or localized scotomas, leading to cortical blindness in the affected region without impairing higher-level functions like object recognition in intact areas.^[32] These effects underscore V1's role as a gateway for conscious visual perception, though subcortical pathways may mediate residual reflexive behaviors.^[33]

Visual Area V2

Visual Area V2 surrounds the primary visual cortex (V1) in the occipital lobe and occupies approximately 10% of the cortical surface area in macaques, with an average size of about 944 mm². It is divided into repeating cytochrome oxidase (CO)-stained stripes—thick, thin, and pale—that run parallel to the V1/V2 border and reflect compartmentalized processing, with thin stripes receiving inputs from V1 color-sensitive blobs, while thick and pale stripes receive from interblob regions. V2 neurons lack spiny stellate cells in layer 4, instead featuring predominantly pyramidal neurons with short apical dendrites, resulting in a neuron density of around 130,000 per mm³.^[34] V2 receives its primary input from V1, accounting for 76.4% of interareal connections, primarily targeting supragranular layers from V1 layers 4C and 4B, with sparse contributions from the koniocellular layers of the lateral geniculate nucleus (LGN). Outputs from V2 project back to V1 (73.2% of efferents, providing feedback), to area V5/MT (11.9%), and to other higher areas like V3 and V4, enabling relay and refinement of visual signals. Layer 6 neurons in V2 also send projections back to the LGN, supporting recurrent processing.^[34]^[35] Functionally, V2 integrates elemental features from V1 into more complex patterns, with larger receptive fields than V1—averaging twice the size, with a classical receptive field radius of 0.74° and a surround radius of 3.56°. Its columnar organization features interdigitated modules for orientation and disparity selectivity, often described as cobweb-like due to the intertwined representation of these features across stripes. In pale stripes, 80% of neurons show strong orientation selectivity, supporting contour integration by combining local edges into longer boundaries. Thick stripes emphasize disparity processing, with 68% of neurons tuned to binocular depth cues, while thin stripes handle color processing in 63% of cells, facilitating color-form binding. Additionally, 63% of V2 neurons are sensitive to natural texture statistics, aiding texture segmentation by detecting boundaries between regions of differing statistical properties.^[36]^[37] V2 responses are modulated by feedback from higher areas such as V4 and V5/MT, which enhance selectivity for relevant features and suppress irrelevant ones, thereby refining the integration of contours, textures, disparity, and color-form associations before signals propagate further in the visual hierarchy.^[38]

Higher Visual Areas

Visual Area V3

The visual area V3, a higher-order region in the primate visual cortex, is divided into a dorsal component (V3d) and a ventral component (often termed VP or V3v), which together form a horseshoe-shaped band surrounding the anterior border of area V2.^[39] This structure positions V3 as a transitional zone between early and more specialized visual areas, with V3d located above the calcarine sulcus representing the lower visual field and VP below it representing the upper visual field.^[40] V3 exhibits a retinotopic organization that is continuous and mirror-symmetric with V2, forming a second-order representation of the visual field where the horizontal meridian is represented at the anterior border and the vertical meridian along the shared boundary with V2.^[39] Receptive fields in V3 are generally larger than those in V2, facilitating broader spatial integration, and the area includes representations sensitive to axes of rotation in the visual field, particularly in its dorsal portion (V3A in humans), which supports processing of rotational motion patterns.^[40] Additionally, neural activity in V3 is modulated by spatial attention, with top-down signals enhancing responses to attended stimuli across its retinotopic map.^[41] Functionally, V3 contributes to dynamic form processing, including the detection of illusory contours formed by motion or alignment cues, which aids in perceiving coherent shapes without explicit edges.^[42] Neurons in V3 also exhibit speed tuning, with preferred speeds typically ranging from 8 to 16 degrees per second, enabling the analysis of motion velocity in both local and global contexts.^[43] Furthermore, V3 processes binocular disparity, encoding absolute and relative depth cues to support surface segmentation and the perception of three-dimensional structure from stereoscopic information.^[42] In terms of connectivity, V3 receives major inputs from layers 4B of V1 and from V2, integrating orientation, color, and motion signals from these earlier areas.^[39] Its outputs project to area V5 (MT) for advanced motion analysis and to parietal regions, such as the intraparietal sulcus, contributing to visuospatial processing in the dorsal stream.^[40]

Visual Area V4

The visual area V4 is located in the ventral occipitotemporal cortex, spanning the prelunate gyrus, lunate sulcus, superior temporal sulcus, and temporal-occipital gyrus in macaques, with analogous regions in humans near the collateral sulcus.^[44] Unlike earlier visual areas, V4 does not form a single continuous retinotopic map but consists of scattered foci, including color-sensitive clusters such as V4α in humans, which exhibit moderate columnar clustering of neurons with similar color preferences rather than discrete blobs.^[45] These clusters, often termed "globs" for color domains (~500 μm in size), interdigitate with orientation-selective regions, supporting integrated feature processing.^[44] V4 maintains a coarse retinotopic organization, bounded posteriorly by V3 and anteriorly by areas like V4A, with representations of the superior (ventral V4) and inferior (dorsal V4) visual fields.^[44] Neurons in V4 have broad receptive fields, typically spanning 8-10 degrees of visual angle, allowing integration of information across larger spatial scales compared to V1 or V2.^[46] Some V4 neurons demonstrate partial invariance to stimulus position and size, responding consistently to contour fragments like curved shapes regardless of exact location within the field, which facilitates robust object feature representation. V4 plays a key role in color processing within the ventral stream, where neurons exhibit opponent-color tuning and contribute to color constancy by adjusting responses to illumination changes, ensuring stable perception of hues across varying lighting conditions.^[44] For shape processing, V4 neurons show complex selectivity for contours and forms, particularly curvature and angular junctions, encoding object boundaries in a manner that bridges simple orientations from V2 toward higher-level object recognition. Attentional mechanisms enhance V4 activity, with feature-based attention amplifying responses to relevant colors or shapes while suppressing irrelevant ones, thereby prioritizing salient object features in cluttered scenes.^[44] Recent analyses of category selectivity in V4 and adjacent ventral areas reveal a gradient of responses to object classes (e.g., faces, bodies) rather than discrete, modular regions, suggesting a more distributed encoding that aligns with naturalistic visual processing.^[47] Lesions to V4 produce deficits in color perception, including cerebral achromatopsia characterized by impaired hue discrimination and color constancy, as seen in human cases with bilateral damage to the V4 region.^[48] Such damage also leads to particular deficits in 3D shape discrimination and attentional feature selection, underscoring V4's role in integrating form and color for object identification.^[44]

Middle Temporal Area (V5/MT)

The Middle Temporal area (MT), also known as V5, is situated in the posterior bank of the superior temporal sulcus in primates, forming a distinct retinotopically organized region that maps the contralateral visual hemifield, with the central 15° of vision occupying over half its surface area.^[49] This area exhibits a columnar organization where neurons are grouped by their preferred direction of motion, facilitating specialized processing within compact modules.^[49] MT receives primary inputs from layer 4B of V1, particularly from the magnocellular pathway, along with contributions from the thick stripes of V2 and area V3, enabling convergence of motion-related signals from earlier visual stages.^[50] MT plays a pivotal role in motion perception, with the majority of its neurons demonstrating robust selectivity for the direction and speed of visual stimuli, often tuned to velocities around 30°/s.^[51] These neurons integrate local motion signals into coherent global patterns, such as those in random dot kinematograms, supporting the analysis of complex moving scenes.^[49] Adjacent to MT, the medial superior temporal area (MST) extends this processing to optic flow patterns, aiding in the perception of self-motion and heading direction during locomotion.^[52] Functionally, MT neurons display directional tuning curves that peak sharply for preferred directions, and the area shows some segregation in responses to real motion versus apparent motion, with stronger activation often for veridical stimuli in cluttered environments.^[53] MT projects outputs to dorsal stream regions, including the parietal areas such as the ventral intraparietal (VIP) and lateral intraparietal (LIP) areas, as well as frontal regions like the frontal eye fields (FEF), and the superior colliculus, integrating motion information for visuomotor control and attention.^[49] Lesions to bilateral MT result in akinetopsia, a rare condition characterized by motion blindness where fast-moving objects appear trailing or stationary, while slow motion remains perceptible, as documented in patient L.M.^[54]

Visual Area V6

Visual Area V6 is situated in the dorsal occipital cortex, within the depths of the parieto-occipital sulcus. This region features large receptive fields that predominantly encompass the peripheral visual field, enabling broad spatial coverage beyond central vision.^[55] Its organization is retinotopic, mapping the entire contralateral visual hemifield in a systematic manner, though with an emphasis on wide-field representations that extend across extensive portions of the visual periphery. In both macaques and humans, this topographic layout supports processing of expansive visual scenes rather than fine central details.^[56] Neurons in V6 exhibit pronounced sensitivity to translational motion across the visual field and to radial motion components inherent in optic flow patterns generated by self-movement.^[57]^[58] This area also demonstrates saccade-related remapping, where receptive fields shift in anticipation of eye movements to maintain perceptual stability during gaze changes.^[59] Furthermore, V6 integrates extra-retinal cues (such as efference copy and proprioception) with visual signals, particularly during head movements, to form a unified representation of self-motion.^[60]^[61] V6 receives major afferent projections from primary visual areas V1 and V2, as well as from the motion-sensitive middle temporal area MT, allowing it to synthesize early visual and dynamic motion information.^[62] Its efferents project prominently to parietal regions, including area V6A and the superior parietal lobule, facilitating visuomotor transformations essential for action guidance.^[63] Unlike MT, which shows a stronger bias toward foveal motion processing, V6 maintains a distinct peripheral emphasis, prioritizing wide-field dynamics over central stimuli.^[64] In terms of function, V6 plays a critical role in egocentric spatial perception, supporting navigation through environments by analyzing self-motion cues and optic flow for heading direction estimation.^[65] This contributes to real-time visuomotor control, such as in obstacle avoidance during locomotion, where peripheral motion signals inform immediate environmental interactions.^[66] Recent investigations highlight how developmental spontaneous activity patterns help refine motion tuning in extrastriate areas like V6, establishing precise selectivity through early network dynamics.^[67] While V6 builds on motion direction signals from V5, its broader field integration distinguishes it for egocentric tasks.^[68]

Processing Pathways

Ventral Stream

The ventral stream, commonly known as the "what" pathway, is a hierarchical processing route in the primate visual cortex dedicated to object recognition and form perception, originating in the primary visual cortex (V1) and progressing through V2, V4, and culminating in the inferotemporal cortex (IT). This pathway enables the analysis of visual stimuli for identification, independent of spatial location, by integrating progressively complex features along its occipitotemporal course. Seminal lesion studies in monkeys demonstrated that damage to this ventral route impairs object discrimination while preserving spatial abilities, distinguishing it from the dorsal stream.^[69]^[70] Within the ventral stream, parallel channels handle specific attributes: V4 specializes in color processing, contributing to object segmentation and surface representation, while the IT cortex focuses on form and shape invariance, allowing recognition of objects across variations in viewpoint, size, and illumination. Neurons in the IT cortex exhibit viewpoint-invariant responses to complex objects, supporting robust identification through distributed representations that tolerate transformations. Building on edge and orientation selectivity in V1, these areas refine features for holistic object perception. The stream's functions extend to category-specific processing, with fusiform gyrus regions showing enhanced selectivity for faces in the fusiform face area (FFA) and for written words in the visual word form area (VWFA), facilitating rapid expert-level discrimination.^[71]^[72]^[73]^[74] Category hierarchies in the ventral stream adapt to experience, as outlined in Gauthier's expertise model, where fusiform activation increases for subordinate-level categorization in domains of proficiency, such as distinguishing bird species or car models, reflecting plasticity in perceptual tuning.^[75] The pathway features reciprocal connectivity with prefrontal cortex regions, enabling memory integration for contextual object recognition and decision-making during visual tasks. Recent connectome analyses highlight experience-dependent plasticity that refines functional gradients along the ventral stream, with higher-order areas showing greater adaptability to learned visual categories.^[76] Lesions disrupting this pathway, particularly in occipitotemporal regions, result in visual agnosia, characterized by profound deficits in object and face recognition despite intact low-level vision and intellect.^[77]

Dorsal Stream

The dorsal stream, often referred to as the "where" or "how" pathway, originates in the primary visual cortex (V1) and proceeds through secondary areas V2 and V3, then to motion-sensitive regions V5 (also known as MT) and V6, before terminating in the posterior parietal cortex (PPC). This hierarchical progression enables the integration of visual inputs for spatial and action-oriented processing, distinct from object recognition pathways. The stream is predominantly driven by magnocellular (M) inputs from the lateral geniculate nucleus, which facilitate rapid transmission of low-contrast, high-speed signals essential for detecting motion and coarse spatial features. Key functions of the dorsal stream include directing visuospatial attention to relevant locations in the visual field and guiding visuomotor actions, such as planning grasping trajectories based on object orientation and size. It performs critical coordinate transformations, converting retinocentric (eye-centered) representations into allocentric (object- or world-centered) frames to support flexible navigation and manipulation in dynamic environments. These capabilities allow for real-time adjustments during tasks like reaching or avoiding obstacles, emphasizing the stream's role in "vision for action" rather than conscious perception. The PPC within the dorsal stream maintains extensive connectivity with motor-related areas, including reciprocal projections to the frontal eye fields (FEF) that coordinate saccadic eye movements for scanning and fixating targets. Top-down influences from frontal regions modulate dorsal stream activity, introducing variability based on cortical states such as attention or task demands, as evidenced by causal streams along the dorsal attention network that enhance voluntary orienting. Lesions in the PPC, particularly following right-hemisphere strokes, disrupt dorsal stream functions and result in hemispatial neglect syndrome, where individuals fail to detect or respond to stimuli in the contralesional (typically left) visual space due to impaired spatial attention. This deficit highlights the stream's essential role in orienting behavior, with recovery often partial and linked to network reorganization.

Models of Visual Processing

Hierarchical Processing Model

The hierarchical processing model describes visual information flow in the cortex as a series of successive stages, where low-level features detected in early areas are progressively combined into more abstract, complex representations in higher areas, ultimately enabling object recognition. This framework, exemplified by the Riesenhuber-Poggio model, posits that processing begins in V1 with detection of basic elements like edges and orientations, advances to V2 and V4 for integration into shapes and forms, and culminates in the inferotemporal (IT) cortex with representations of whole objects that exhibit tolerance to variations in position, size, and viewpoint. The model emphasizes a feedforward architecture where each stage builds greater invariance through nonlinear operations, allowing the system to generalize across transformations while maintaining selectivity for specific stimuli.^[78] Central to this model are convergence-divergence zones, where inputs from multiple lower-level areas converge onto neurons in higher areas, enabling the synthesis of complex features from simpler ones, as mapped in the primate visual hierarchy. Nonlinear pooling mechanisms, such as max pooling, contribute to transformation tolerance by selecting the strongest matching features across variations, a process simulated computationally in the HMAX model, which replicates physiological responses through alternating layers of selectivity and invariance operations. These zones and pooling operations ensure that higher-level representations are robust to retinal changes, supporting efficient object identification despite shifts in gaze or illumination. Empirical support comes from single-unit recordings in primates, which reveal a gradient of increasing receptive field size and stimulus complexity along the hierarchy: V1 neurons respond to simple bars and edges, while IT neurons selectively activate for complex objects like faces or hands, with clustered responses indicating feature conjunctions. Functional MRI adaptation studies further demonstrate this buildup of invariance, showing reduced BOLD responses in higher ventral areas (e.g., lateral occipital complex and fusiform face area) to repeated presentations of the same object across size or viewpoint changes, but not to dissimilar objects, confirming shared neural representations tolerant to transformations. Although primarily feedforward, the model incorporates feedback mechanisms to refine processing, as in predictive coding frameworks where higher areas send top-down predictions to lower levels to suppress expected inputs and highlight discrepancies, enhancing efficiency and contextual integration in the visual hierarchy. This interplay addresses limitations of pure feedforward accounts by accounting for modulatory effects observed in cortical responses.

Ventral-Dorsal Distinction

The ventral-dorsal distinction, also known as the two-streams hypothesis, posits that visual processing in the primate brain is divided into two functionally distinct pathways: the ventral stream for conscious perception and object recognition, and the dorsal stream for unconscious visuomotor guidance and action. This model originated from studies of patient DF, who suffered bilateral damage to the lateral occipital complex in the ventral stream following anoxia, resulting in visual form agnosia where she could not consciously identify or discriminate object shapes but could accurately grasp objects using visually guided actions, such as scaling her grip aperture to object size. In contrast, patients with dorsal stream damage, like those with optic ataxia, exhibit impaired action guidance despite intact perceptual abilities, supporting the idea that the ventral stream constructs perceptual representations for identification ("what" pathway), while the dorsal stream transforms visual inputs into sensorimotor coordinates for action ("where" or "how" pathway).^[79] Psychologically, this distinction manifests in dissociations between perception and action, particularly with visual illusions that exploit ventral stream processing but spare dorsal stream computations. For instance, in the Ebbinghaus illusion, where a central target circle appears smaller when surrounded by larger circles, healthy individuals misperceive the target's size (ventral influence) but adjust their grip apertures accurately to the actual size during grasping (dorsal calibration), demonstrating how perceptual biases do not affect online action control. Attention plays a modulating role, with dorsal stream mechanisms prioritizing salient stimuli for rapid orienting and action, while ventral processing integrates attentional feedback for detailed object analysis, highlighting a functional interplay that supports adaptive behavior. Recent updates to the model challenge rigid categorizations, proposing instead that category selectivity in the ventral stream operates as continuous gradients rather than discrete modules, allowing flexible representation of visual features across object types like faces, bodies, and scenes.^[80] Ancient attention mechanisms, which emerged over 500 million years ago in vertebrate ancestors, are evidenced by recurrent circuits in the superior colliculus (a dorsal-like structure) that perform center-surround computations for contrast detection and stimulus prioritization independently of cortical input.^[81] These evolutionary roots suggest that dorsal attention functions evolved to support survival-oriented actions, with ventral perception building upon this foundation for more elaborate cognitive processing. Supporting evidence from neuroimaging reveals pathway-specific activations: functional MRI studies show dorsal stream areas like the anterior intraparietal sulcus activating during visually guided grasping tasks without concurrent ventral stream engagement in perceptual areas like the lateral occipital complex. Similarly, AI-based digital twin models, such as deep learning architectures simulating both streams (e.g., VeDo-Net), accurately predict neural responses to visual stimuli, reproducing dissociations in object recognition versus spatial processing by training separate ventral networks on recognition tasks and dorsal networks on transformation metrics. These models confirm the hypothesis by forecasting pathway-specific behaviors with high fidelity to empirical data.^[82] Within the broader hierarchical processing framework, this distinction underscores parallel functional specializations across visual stages.

References

[1]
Neuroanatomy, Visual Cortex - StatPearls - NCBI Bookshelf
The primary purpose of the visual cortex is to receive, segment, and integrate visual information. The processed information from the visual cortex is ...Introduction · Structure and Function · Surgical Considerations
[2]
Chapter 15: Visual Processing: Cortical Pathways
The primary visual cortex, V1, sends input to extrastriate cortex and to visual association cortex. The information from the “color”, “shape/form ...
[3]
The Primary Visual Cortex by Matthew Schmolesky - Webvision
Jun 14, 2007 · Anatomical hierarchical models place the visual cortical areas into a multi-level processing model based upon the pattern of feedforward, ...
[4]
Visual System: Central Processing – Introduction to Neuroscience
The primary visual cortex (also called striate cortex or V1) is located in the posterior region of the human brain within the occipital lobes. In this image the ...
[5]
Visual Cortex in Humans | Request PDF - ResearchGate
Human visual cortex comprises 4-6. billion neurons that are organized into more than a dozen distinct functional areas. These areas include the gray matter ...
[6]
[PDF] PRIMARY VISUAL CORTEX AND VISUAL AWARENESS
Perceptual binding refers to the problem of how the brain integrates diverse information about colour, ori- entation, motion, form and so on into a single ...
[7]
Evolution of neural processing for visual perception in vertebrates
This review compares, across classes of vertebrates, the functional and anatomical characteristics of (a) the neural pathways that process visual information ...
[8]
Functional modulation of primary visual cortex by the superior ...
Sep 25, 2018 · The superior colliculus could potentially also influence the input to the primary visual cortex through its projection to the shell of the dLGN.
[9]
Superior colliculus encodes visual saliency before the ... - PNAS
Aug 14, 2017 · Our results show that neurons in the superficial visual layers of the superior colliculus (SCs) encoded saliency earlier and more robustly than V1 neurons.
[10]
Neuroanatomy, Occipital Lobe - StatPearls - NCBI Bookshelf - NIH
The primary visual cortex, also known as V1 or Brodmann area 17, surrounds the calcarine sulcus on the occipital lobe's medial aspect. It receives the visual ...Structure and Function · Blood Supply and Lymphatics · Surgical Considerations
[11]
[PDF] Visual Cortex in Humans - Stanford University
Regions of cortex that respond powerfully to reti- nal stimulation are called visual cortex. In the human brain, the visual cortex is located in the occipi-.
[12]
The Primary Visual Cortex - Webvision - NCBI Bookshelf
May 1, 2005 · Figure 8. Visual input to the brain goes from eye to LGN and then to primary visual cortex, or area V1, which is located in the posterior of ...
[13]
Neuroanatomy, Optic Tract - StatPearls - NCBI Bookshelf
The left and right LGN serve as points of neural origin for optic radiations (Meyer's loop, central bundle, and Baum's loop) that travel to the primary ...
[14]
Hemispheric specialization for global and local processing - Journals
Neuropsychological evidence indicates that the global aspect of complex visual scenes is preferentially processed by the right hemisphere, and local aspects ...
[15]
Hemispheric differences in global versus local processing of ...
Hemispheric differences in global versus local processing of hierarchical visual stimuli by normal subjects: New data and a meta-analysis of previous studies.
[16]
Functional connectomics spanning multiple areas of mouse ... - Nature
Apr 9, 2025 · Here we introduce the MICrONS functional connectomics dataset with dense calcium imaging of around 75,000 neurons in primary visual cortex (VISp) ...
[17]
https://www.annualreviews.org/doi/10.1146/annurev.neuro.28.061604.135731
[18]
Lamination, Borders, and Thalamic Projections of the Primary Visual ...
Laminar Organization of V1 in Humans and NHPs. According to Brodmann, V1 in humans and NHPs possesses a thick and laminated layer 4 (L4; termed inner ...
[19]
Ventral visual cortex in humans: Cytoarchitectonic mapping of two ...
For early visual areas (V1, V2, V3), anatomical and functional homology between humans and nonhuman primates can be considered as largely proven (Conway and ...
[20]
Layer 4 of mouse neocortex differs in cell types and circuit ... - Nature
Sep 13, 2019 · In line with this, it has been shown that excitatory cells in L4 of mouse and rat primary visual cortex (V1) are pyramidal, in contrast to L4 in ...Missing: diversification | Show results with:diversification
[21]
Development and functional diversification of cortical interneurons
PV+ basket cells are the most abundant type of interneuron in the neocortex. They have highly torturous axons that form synapses on the soma and proximal ...
[22]
Borders of Multiple Visual Areas in Humans Revealed by Functional ...
Retinotopically organized visual areas in humans extend anteriorly to overlap several areas previously shown to be activated by written words. Formats available.
[23]
Borders of multiple visual areas in humans revealed by ... - PubMed
The borders of human visual areas V1, V2, VP, V3, and V4 were precisely and noninvasively determined. Functional magnetic resonance images were recorded.<|control11|><|separator|>
[24]
Reorganization of Retinotopic Maps After Occipital Lobe Infarction
We studied patient JS who had a right occipital infarct that encroached on visual areas V1, V2v and VP. When tested psychophysically, he was very impaired ...Missing: network | Show results with:network
[25]
Extensive topographic remapping and functional sharpening in the ...
Our findings reveal that visual experience following VDM reshapes both structure and function of the visual system and shifts the stability/plasticity balance ...
[26]
The Lateral Geniculate Nucleus - SpringerLink
LGN neurons send axonal projections to the visual cortex via the optic radiations. ... Most axonal output from the LGN terminates in the primary visual cortex (V1) ...
[27]
The Columnar Organization of the Striate Cortex - NCBI - NIH
The striate cortex is composed of repeating units, or modules, that contain all the neuronal machinery necessary to analyze a small region of visual space.
[28]
Complete Pattern of Ocular Dominance Columns in Human Primary ...
The width of the ocular dominance columns was fairly uniform across the visual field representation in striate cortex, except near the monocular crescent ...
[29]
Binocular integration and disparity selectivity in mouse primary ...
We demonstrate that mouse V1 neurons are tuned for binocular disparities, or spatial differences, between the inputs from each eye, thus extracting signals ...
[30]
Integration of Multiple Spatial Frequency Channels in Disparity ...
Jul 8, 2015 · The primary visual cortex is the first stage of this processing, and neurons there are tuned to a limited range of spatial frequencies (SFs).
[31]
Orientation Tuning Depends on Spatial Frequency in Mouse Visual ...
Sep 21, 2016 · In order to explain this dependence between orientation selectivity and SF, we used the common Gabor model (Gabor, 1946) to predict the neuronal ...
[32]
Cortical Blindness - StatPearls - NCBI Bookshelf - NIH
Cortical blindness is an important cause of blindness due to damage to the occipital cortex. It is commonly associated with posterior circulation stroke.
[33]
Primary visual cortex, awareness and blindsight - PubMed Central
Following such lesions, blindsight allows for some residual visual responses to stimuli presented to the scotoma (blind portion of the visual field).
[34]
Anatomy and Physiology of Macaque Visual Cortical Areas V1, V2 ...
Here we provide an overview of over 350 published studies of these three areas in the genus Macaca, whose visual system provides the closest model for human ...
[35]
[PDF] Divided by Cytochrome Oxidase: A Map of the Projections from V1 to ...
Contrary to prior reports, the thin, pale, and thick stripes in V2 received ... Sincich, J. C. Horton, J. Comp. Neurol., in press. (2002). 23. Experiments ...
[36]
The functional organization of area V2, I - Cambridge University Press
Jul 2, 2002 · We used an initial 8-way stripe classification (4 stripes, and 4 “marginal” zones at interstripes boundaries), and a 9-way layer classification ...
[37]
[PDF] A functional and perceptual signature of the second visual area in ...
May 19, 2013 · V1 neurons responded similarly to both stimulus types, whereas. V2 neurons often responded more vigorously to naturalistic textures than to ...
[38]
The Organization of Connections between Areas V5 and V2 in ...
The return projection from V5 to V2 is more widespread than the origin of the forward projection. It extends not only between the clusters of V5-efferent cells ...
[39]
The case for primate V3 | Proceedings of the Royal Society B
Dec 14, 2011 · The visual system in primates is represented by a remarkably large expanse of the cerebral cortex. While more precise investigative studies ...
[40]
Topographic organization of areas V3 and V4 and its relation to ...
We propose that V3 and V4 play a critical role in this supra-areal organization by filtering information about the visual environment along parallel pathways ...Missing: paper | Show results with:paper
[41]
Coding of Stereoscopic Depth Information in Visual Areas V3 and V3A
... illusory contour-defined shapes in V3A, suggesting that V3A may be involved in surface segmentation. It remains to be seen if single neurons in V3A process ...
[42]
Global Motion Processing in Human Visual Cortical Areas V2 and V3
The results suggest the surprising conclusion that global motion processing is a key function of cortical visual areas V2 and V3.
[43]
https://pmc.ncbi.nlm.nih.gov/articles/PMC6705536/
[44]
Organization of Color-Selective Neurons in Macaque Visual Area V4
We conclude that neurons in area V4 are moderately clustered according to their color selectivity and that this weak clustering is columnar in structure.
[45]
Central V4 Receptive Fields Are Scaled by the V1 Cortical ...
May 6, 2009 · This asymmetry is completely determined by the transformation of visual space according to the cortical magnification factor for primary visual ...
[46]
Full article: Rethinking category-selectivity in human visual cortex
Aug 20, 2025 · First, minimally a neural response or region of the brain is 'selective' if it responds differentially and preferentially to a particular ...
[47]
On the role of cortical area V4 in the discrimination of hue and ...
Sep 1, 1987 · The visual disorder following removal of visual area V4 strikingly resembles the clinical disorder of mild cerebral achromatopsia with ...
[48]
The neurological basis of conscious color perception in a blind patient
Lesions restricted to V4 lead to a specific loss of conscious color vision (cerebral achromatopsia). Although color blind, achromatopsic patients can ...
[49]
https://doi.org/10.1146/annurev.neuro.26.041002.131052
[50]
https://doi.org/10.1111/j.1460-9568.1989.tb00798.x
[51]
https://doi.org/10.1152/jn.1983.49.5.1127
[52]
https://doi.org/10.1038/81860
[53]
https://doi.org/10.1152/jn.1971.34.4.1011
[54]
https://doi.org/10.1093/brain/106.2.313
[55]
Wide-Field Retinotopy Defines Human Cortical Visual Area V6 - PMC
Macaque V6 is located in the depths of the parieto-occipital sulcus (POS) and contains a retinotopic map of the entire contralateral hemifield. In contrast to ...
[56]
The cortical visual area V6 in macaque and human brains - PubMed
Area V6 was first described in the macaque monkey and then, recently, in the human. In both primates, it is located in the medial parieto-occipital region of ...
[57]
The functional role of the medial motion area V6 - Frontiers
V6 is a motion sensitive area that largely represents the peripheral part of the visual field and whose cells are very sensitive to translational motion.Missing: saccade | Show results with:saccade
[58]
Sensitivity of human visual cortical area V6 to stereoscopic depth ...
Visual area V6, in the parieto-occipital sulcus, showed a unique pattern of results, responding well to all optic flow patterns but much more strongly when ...Missing: translational | Show results with:translational
[59]
Spatiotopic coding and remapping in humans - PMC - NIH
Neurons in many visual areas show clear saccade-related changes to their RF properties. In some areas, such as areas V6 [3] and VIP [4] there exist neurons ...
[60]
Cortical Connections of Area V6Av in the Macaque: A Visual-Input ...
Feb 2, 2011 · V6 is an extrastriate area that is likely to exist in the brains of most, if not all, primates (Galletti et al., 1999b; Pitzalis et al., 2006; ...
[61]
A new neural framework for visuospatial processing - PubMed Central
The visual information from area V6 reaches the parietal lobe through two main channels: one projecting medially to the bimodal (visual and somatosensory) area ...
[62]
The functional role of the medial motion area V6 - PubMed Central
V6 is a motion sensitive area that largely represents the peripheral part of the visual field and whose cells are very sensitive to translational motion.Missing: bias | Show results with:bias
[63]
Activation of human visual area V6 during egocentric navigation with ...
Apr 10, 2023 · V6 is a retinotopic area located in the dorsal visual stream that integrates eye movements with retinal and visuo-motor signals.
[64]
Visual Neuropsychology in Development: Anatomo-Functional Brain ...
The present review summarizes the available neuropsychological evidence on the development of visual competences, with a particular focus on the associated ...
[65]
Adaptation of spontaneous activity in the developing visual cortex
Spontaneous activity drives the establishment of appropriate connectivity in different circuits during brain development.Missing: V6 obstacle avoidance
[66]
Human V6: The Medial Motion Area | Cerebral Cortex
Jun 5, 2009 · Area V6 is highly selective for coherently moving fields of dots, both at individual and group levels and even with a visual stimulus of ...
[67]
[PDF] Object vision and spatial vision:two cortical p hways - MRI Questions
The two cortical visual pathways are schematized in Fig. t. One of them consists of a multisynaptic occipitotemporal projec- tion system that follows the course ...
[68]
[PDF] Distributed Hierarchical Processing in the Primate Cerebral Cortex
The original version of the cortical hierarchy in the ma- caque visual pathway spanned 6 levels and was based on 36 distinct pathways among 13 identified ...
[69]
Toward a Unified Theory of Visual Area V4 - PMC - PubMed Central
Visual area V4 is a midtier cortical area in the ventral visual pathway. It is crucial for visual object recognition and has been a focus of many studies on ...Missing: scattered | Show results with:scattered
[70]
How does the brain solve visual object recognition? - PubMed Central
Neuronal selectivities to complex object features in the ventral visual pathway of the macaque cerebral cortex. ... Inferotemporal cortex and object vision.
[71]
The Fusiform Face Area: A Module in Human Extrastriate Cortex ...
Jun 1, 1997 · We found an area in the fusiform gyrus in 12 of the 15 subjects tested that was significantly more active when the subjects viewed faces than when they viewed ...
[72]
The visual word form area: expertise for reading in the fusiform gyrus
This brain specialization is essential to rapid reading ability because it enhances perception of words by becoming specifically tuned to recurring properties ...
[73]
Expertise for cars and birds recruits brain areas involved in face ...
Expertise with unfamiliar objects ('greebles') recruits face-selective areas in the fusiform gyrus (FFA) and occipital lobe (OFA).
[74]
Plasticity versus stability across the human cortical visual connectome
Jul 18, 2019 · We quantify plasticity as the complement of the heritability of resting-state functional connectivity and thereby demonstrate a non-monotonic relationship.Missing: stream | Show results with:stream
[75]
Visual agnosia - PubMed
This condition is caused by lesions in the visual association cortex, sparing primary visual cortex. There are 2 main pathways that process visual information: ...
[76]
(PDF) Riesenhuber, M. & Poggio, T. Hierarchical models of object ...
Aug 10, 2025 · Visual processing in cortex is classically modeled as a hierarchy of increasingly sophisticated representations, naturally extending the ...Missing: Tsao | Show results with:Tsao
[77]
Rethinking category-selectivity in human visual cortex - arXiv
Nov 12, 2024 · Here, we argue that while valuable insights have been gained through the lens of category-selectivity, a more complete view of visual function ...
[78]
A deep learning model of dorsal and ventral visual streams for DVSD
Nov 10, 2024 · In this paper, we present a quantitatively accurate model for the visual system. Specifically, we propose a VeDo-Net model that comprises both ventral and ...Missing: digital twin