Visual cortex: Difference between revisions

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Research on V1 has also revealed the presence of orientation-selective cells, which respond preferentially to stimuli with a specific orientation, contributing to the perception of edges and contours. The discovery of these orientation-selective cells has been fundamental in shaping our understanding of how V1 processes visual information.
 
Furthermore, V1 exhibits plasticity, allowing it to undergo functional and structural changes in response to sensory experience. Studies have demonstrated that sensory deprivation or exposure to enriched environments can lead to alterations in the organization and responsiveness of V1 neurons, highlighting the dynamic nature of this critical visual processing hub.{{cn|date=October 2024}}
 
The primary visual cortex, which is defined by its function or stage in the visual system, is approximately equivalent to the striate cortex, also known as Brodmann area 17, which is defined by its anatomical ___location. The name "striate cortex" is derived from the line of Gennari, a distinctive stripe visible to the naked eye that represents [[myelin]]ated [[axons]] from the [[lateral geniculate body]] terminating in layer 4 of the [[gray matter]].
 
Brodmann area 17 is just one subdivision of the broader Brodmann areas, which are regions of the cerebral cortex defined based on cytoarchitectural differences. In the case of the striate cortex, the line of Gennari corresponds to a band rich in myelinated nerve fibers, providing a clear marker for the primary visual processing region.{{cn|date=October 2024}}
 
Additionally, the functional significance of the striate cortex extends beyond its role as the primary visual cortex. It serves as a crucial hub for the initial processing of visual information, such as the analysis of basic features like orientation, spatial frequency, and color. The integration of these features in the striate cortex forms the foundation for more complex visual processing carried out in higher-order visual areas. Recent neuroimaging studies have contributed to a deeper understanding of the dynamic interactions within the striate cortex and its connections with other visual and non-visual brain regions, shedding light on the intricate neural circuits that underlie visual perception.<ref>{{cite journal |vauthors=Glickstein M, Rizzolatti G |title=Francesco Gennari and the structure of the cerebral cortex |journal=Trends in Neurosciences |volume=7 |issue=12 |pages=464–467 |date=1 December 1984|doi=10.1016/S0166-2236(84)80255-6 |s2cid=53168851 }}</ref>
 
The primary visual cortex is divided into six functionally distinct layers, labeled 1 to 6. Layer 4, which receives most visual input from the [[lateral geniculate nucleus]] (LGN), is further divided into 4 layers, labelled 4A, 4B, 4Cα, and 4Cβ. Sublamina 4Cα receives mostly [[Magnocellular cell|magnocellular]] input from the LGN, while layer 4Cβ receives input from [[Parvocellular cell|parvocellular]] pathways.<ref>{{cite journal | vauthors = Hubel DH, Wiesel TN | title = Laminar and columnar distribution of geniculo-cortical fibers in the macaque monkey | journal = The Journal of Comparative Neurology | volume = 146 | issue = 4 | pages = 421–450 | date = December 1972 | pmid = 4117368 | doi = 10.1002/cne.901460402 | s2cid = 6478458 }}</ref><ref>{{cite book |last1=Churchland |first1=Patricia Smith |last2=Sejnowski |first2=Terrence Joseph |author1-link=Patricia Churchland |author2-link=Terry Sejnowski |title=[[The Computational Brain]] |publisher=[[MIT Press]] |___location=Cambridge, Massachusetts |year=1992 |isbn=978-0-262-53120-7 |page=149}}</ref>
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{{further|Visual system}}
{{technical|date=September 2016}}
The initial stage of visual processing within the visual cortex, known as V1, plays a fundamental role in shaping our perception of the visual world. V1 possesses a meticulously defined map, referred to as the retinotopic map, which intricately organizes spatial information from the visual field. In humans, the upper bank of the calcarine sulcus in the occipital lobe robustly responds to the lower half of the visual field, while the lower bank responds to the upper half. This retinotopic mapping conceptually represents a projection of the visual image from the retina to V1.
 
The importance of this retinotopic organization lies in its ability to preserve spatial relationships present in the external environment. Neighboring neurons in V1 exhibit responses to adjacent portions of the visual field, creating a systematic representation of the visual scene. This mapping extends both vertically and horizontally, ensuring the conservation of both horizontal and vertical relationships within the visual input.
 
Moreover, the retinotopic map demonstrates a remarkable degree of plasticity, adapting to alterations in visual experience. Studies have revealed that changes in sensory input, such as those induced by visual training or deprivation, can lead to shifts in the retinotopic map. This adaptability underscores the brain's capacity to reorganize in response to varying environmental demands, highlighting the dynamic nature of visual processing.
 
Beyond its spatial processing role, the retinotopic map in V1 establishes intricate connections with other visual areas, forming a network crucial for integrating diverse visual features and constructing a coherent visual percept. This dynamic mapping mechanism is indispensable for our ability to navigate and interpret the visual world effectively.
 
The correspondence between specific locations in V1 and the subjective visual field is exceptionally precise, even extending to map the blind spots of the retina. Evolutionarily, this correspondence is a fundamental feature found in most animals possessing a V1. In humans and other species with a fovea (cones in the retina), a substantial portion of V1 is mapped to the small central portion of the visual field—afield, a phenomenon termed cortical magnification. This magnification reflects an increased representation and processing capacity devoted to the central visual field, essential for detailed visual acuity and high-resolution processing.
 
Notably, neurons in V1 have the smallest receptive field size, signifying the highest resolution, among visual cortex microscopic regions. This specialization equips V1 with the ability to capture fine details and nuances in the visual input, emphasizing its pivotal role as a critical hub in early visual processing and contributing significantly to our intricate and nuanced visual perception.<ref>{{cite journal | vauthors = Wu F, Lu Q, Kong Y, Zhang Z | title = A Comprehensive Overview of the Role of Visual Cortex Malfunction in Depressive Disorders: Opportunities and Challenges | journal = Neuroscience Bulletin | volume = 39 | issue = 9 | pages = 1426–1438 | date = September 2023 | pmid = 36995569 | pmc = 10062279 | doi = 10.1007/s12264-023-01052-7 }}</ref>
 
In addition to its role in spatial processing, the retinotopic map in V1 is intricately connected with other visual areas, forming a network that contributes to the integration of various visual features and the construction of a coherent visual percept. This dynamic mapping mechanism is fundamental to our ability to navigate and interpret the visual world effectively.<ref name= kepler1604 >Johannes Kepler (1604) Paralipomena to Witelo whereby The Optical Part of Astronomy is Treated (Ad Vitellionem Paralipomena, quibus astronomiae pars optica traditvr, 1604), as cited by A.Mark Smith (2015) From Sight to Light. Kepler modeled the eye as a water-filled glass sphere, and discovered that each point of the scene taken in by the eye projects onto a point on the back of the eye (the retina).</ref> The correspondence between a given ___location in V1 and in the subjective visual field is very precise: even the [[Blind spot (vision)|blind spots]] of the retina are mapped into V1. In terms of evolution, this correspondence is very basic and found in most animals that possess a V1. In humans and other animals with a [[Fovea centralis|fovea]] ([[Cone cell|cones]] in the retina), a large portion of V1 is mapped to the small, central portion of visual field, a phenomenon known as [[cortical magnification]]. Perhaps for the purpose of accurate spatial encoding, neurons in V1 have the smallest [[receptive field]] size (that is, the highest resolution) of any visual cortex microscopic regions.
 
The tuning properties of V1 neurons (what the neurons respond to) differ greatly over time. Early in time (40 ms and further) individual V1 neurons have strong tuning to a small set of stimuli. That is, the neuronal responses can discriminate small changes in visual [[Orientation (mental)|orientations]], [[spatial frequencies]] and [[color]]s (as in the optical system of a [[camera obscura]], but projected onto [[retina]]l cells of the eye, which are clustered in density and fineness).<ref name= kepler1604 /> Each V1 neuron propagates a signal from a retinal cell, in continuation. Furthermore, individual V1 neurons in humans and other animals with [[binocular vision]] have ocular dominance, namely tuning to one of the two eyes. In V1, and primary sensory cortex in general, neurons with similar tuning properties tend to cluster together as [[cortical column]]s. [[David Hubel]] and [[Torsten Wiesel]] proposed the classic ice-cube organization model of cortical columns for two tuning properties: [[ocular dominance columns|ocular dominance]] and orientation. However, this model cannot accommodate the color, spatial frequency and many other features to which neurons are tuned {{Citation needed|date=November 2011}}. The exact organization of all these cortical columns within V1 remains a hot topic of current research.