Visual cortex

The visual cortex of the brain is the area of the cerebral cortex that processes visual
information. It is located in the occipital lobe. Sensory input originating from the eyes travels
through the lateral geniculate nucleus in the thalamus and then reaches the visual cortex. The
area of the visual cortex that receives the sensory input from the lateral geniculate nucleus is the
primary visual cortex, also known as visual area 1 (V1), Brodmann area 17, or the striate cortex.
The extrastriate areas consist of visual areas 2, 3, 4, and 5 (also known as V2, V3, V4, and V5, or
Brodmann area 18 and all Brodmann area 19).[1]
 Visual cortex

View of the brain from behind. Red = Brodmann area 17 (primary visual cortex); orange = area 18; yellow
= area 19

Brain shown from the side, facing left. Above: view from outside, below: cut through the middle. Orange
= Brodmann area 17 (primary visual cortex)

Details

Identifiers

Latin Cortex visualis

MeSH D014793 ([Link]

=D014793)

NeuroLex ID nlx_143552 ([Link]

 552)

FMA 242644 ([Link]

es/FMA/?p=classes&conceptid=http%3A%2F%2Fp
[Link]%2Fsig%2Font%2Ffma%2Ffma242644)

Anatomical terms of neuroanatomy

Both hemispheres of the brain include a visual cortex; the visual cortex in the left hemisphere
receives signals from the right visual field, and the visual cortex in the right hemisphere receives
signals from the left visual field.

Introduction

The primary visual cortex (V1) is located in and around the calcarine fissure in the occipital lobe.
Each hemisphere's V1 receives information directly from its ipsilateral lateral geniculate nucleus
that receives signals from the contralateral visual hemifield.

Neurons in the visual cortex fire action potentials when visual stimuli appear within their
receptive field. By definition, the receptive field is the region within the entire visual field that
elicits an action potential. But, for any given neuron, it may respond best to a subset of stimuli
within its receptive field. This property is called neuronal tuning. In the earlier visual areas,
neurons have simpler tuning. For example, a neuron in V1 may fire to any vertical stimulus in its
receptive field. In the higher visual areas, neurons have complex tuning. For example, in the
inferior temporal cortex (IT), a neuron may fire only when a certain face appears in its receptive
field.

The visual cortex receives its blood supply primarily from the calcarine branch of the posterior
cerebral artery.
 Psychological model of the neural processing of visual
information

Ventral-dorsal model …

The dorsal stream (green) and ventral stream (purple) are shown. They originate from primary visual cortex.

V1 transmits information to two primary pathways, called the ventral stream and the dorsal
stream.[2]

The ventral stream begins with V1, goes through visual area V2, then through visual area V4,
and to the inferior temporal cortex (IT cortex). The ventral stream, sometimes called the "What
 Pathway", is associated with form recognition and object representation. It is also associated
with storage of long-term memory.

The dorsal stream begins with V1, goes through Visual area V2, then to the dorsomedial area
(DM/V6) and medial temporal area (MT/V5) and to the posterior parietal cortex. The dorsal
stream, sometimes called the "Where Pathway" or "How Pathway", is associated with motion,
representation of object locations, and control of the eyes and arms, especially when visual
information is used to guide saccades or reaching.

The what vs. where account of the ventral/dorsal pathways was first described by Ungerleider
and Mishkin.[3]

More recently, Goodale and Milner extended these ideas and suggested that the ventral stream
is critical for visual perception whereas the dorsal stream mediates the visual control of skilled
actions.[4] It has been shown that visual illusions such as the Ebbinghaus illusion distort
judgements of a perceptual nature, but when the subject responds with an action, such as
grasping, no distortion occurs.[5]

Work such as the one from Franz et al[6] suggests that both the action and perception systems
are equally fooled by such illusions. Other studies, however, provide strong support for the idea
that skilled actions such as grasping are not affected by pictorial illusions[7][8] and suggest that
the action/perception dissociation is a useful way to characterize the functional division of labor
between the dorsal and ventral visual pathways in the cerebral cortex.[9]

Primary visual cortex (V1)

Micrograph showing the visual cortex (pink). The pia mater and arachnoid mater
including blood vessels are seen at the top of the image. Subcortical white matter
(blue) is seen at the bottom of the image. HE-LFB stain.

The primary visual cortex is the most studied visual area in the brain. In mammals, it is located
in the posterior pole of the occipital lobe and is the simplest, earliest cortical visual area. It is
highly specialized for processing information about static and moving objects and is excellent in
pattern recognition.

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[10] that represents myelinated axons from the lateral
geniculate body terminating in layer 4 of the gray matter.

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 input from
the LGN, while layer 4Cβ receives input from parvocellular pathways.[11]

The average number of neurons in the adult human primary visual cortex in each hemisphere
has been estimated at around 140 million.[12]
Function
…
The first stage of visual processing in the cortex is called V1.

V1 has a very well-defined map (the retinotopic map) of the spatial information in vision. For
example, in humans, the upper bank of the calcarine sulcus (in the occipital lobe) responds
strongly to the lower half of visual field (below the center), and the lower bank of the calcarine to
the upper half of visual field. In concept, this retinotopic mapping is a projection of the visual
image from retina to V1.[13] The correspondence between a given location in V1 and in the
subjective visual field is very precise: even the 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 animals with a fovea (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.[14]
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 orientations, spatial
frequencies and colors (as in the optical system of a camera obscura, but projected onto retinal
cells of the eye, which are clustered in density and fineness).[13] Each V1 neuron propagates a
signal from a retinal cell, in continuation. Furthermore, individual V1 neurons in humans and
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 columns. David Hubel and Torsten Wiesel proposed the classic ice-cube
organization model of cortical columns for two tuning properties: ocular dominance and
orientation. However, this model cannot accommodate the color, spatial frequency and many
other features to which neurons are tuned. The exact organization of all these cortical columns
within V1 remains a hot topic of current research. The mathematical modeling of this function
has been compared to Gabor transforms.

Later in time (after 100 ms), neurons in V1 are also sensitive to the more global organisation of
the scene (Lamme & Roelfsema, 2000).[15] These response properties probably stem from
recurrent feedback processing (the influence of higher-tier cortical areas on lower-tier cortical
areas) and lateral connections from pyramidal neurons (Hupe et al. 1998). While feedforward
connections are mainly driving, feedback connections are mostly modulatory in their effects
(Angelucci et al., 2003; Hupe et al., 2001). Evidence shows that feedback originating in higher-
level areas such as V4, IT, or MT, with bigger and more complex receptive fields, can modify and
shape V1 responses, accounting for contextual or extra-classical receptive field effects (Guo et
al., 2007; Huang et al., 2007; Sillito et al., 2006).

The visual information relayed to V1 is not coded in terms of spatial (or optical) imagery but
rather are better described as edge detection. As an example, for an image comprising half side
black and half side white, the dividing line between black and white has strongest local contrast
(that is, edge detection) and is encoded, while few neurons code the brightness information
(black or white per se). As information is further relayed to subsequent visual areas, it is coded
as increasingly non-local frequency/phase signals. Note that, at these early stages of cortical
visual processing, spatial location of visual information is well preserved amid the local contrast
encoding (edge detection).

A theoretical explanation of the computational function of the simple cells in the primary visual
cortex has been presented in.[16][17] It is described how receptive field shapes similar to those
found by the biological receptive field measurements performed by DeAngelis et al.[18][19] can be
derived as a consequence of structural properties of the environment in combination with
internal consistency requirements to guarantee consistent image representations over multiple
spatial and temporal scales. It is also described how the characteristic receptive field shapes,
tuned to different scales, orientations and directions in image space, allow the visual system to
compute invariant responses under natural image transformations at higher levels in the visual
hierarchy.[20][17]

In primates, one role of V1 might be to create a saliency map (highlights what is important) from
visual inputs to guide the shifts of attention known as gaze shifts.[21] According to the V1
Saliency Hypothesis, V1 does this by transforming visual inputs to neural firing rates from
millions of neurons, such that the visual location signaled by the highest firing neuron is the
most salient location to attract gaze shift. V1's outputs are received by the superior colliculus (in
the mid-brain), among other locations, which reads out the V1 activities to guide gaze shifts.

Differences in size of V1 also seem to have an effect on the perception of illusions.[22]

V2

Visual area V2, or secondary visual cortex, also called prestriate cortex,[23] is the second major
area in the visual cortex, and the first region within the visual association area. It receives strong
feedforward connections from V1 (direct and via the pulvinar) and sends strong connections to
V3, V4, and V5. It also sends strong feedback connections to V1.
In terms of anatomy, V2 is split into four quadrants, a dorsal and ventral representation in the left
and the right hemispheres. Together, these four regions provide a complete map of the visual
world. V2 has many properties in common with V1: Cells are tuned to simple properties such as
orientation, spatial frequency, and color. The responses of many V2 neurons are also modulated
by more complex properties, such as the orientation of illusory contours,[24][25] binocular
disparity,[26] and whether the stimulus is part of the figure or the ground.[27][28] Recent research
has shown that V2 cells show a small amount of attentional modulation (more than V1, less
than V4), are tuned for moderately complex patterns, and may be driven by multiple orientations
at different subregions within a single receptive field.

It is argued that the entire ventral visual-to-hippocampal stream is important for visual
memory.[29] This theory, unlike the dominant one, predicts that object-recognition memory (ORM)
alterations could result from the manipulation in V2, an area that is highly interconnected within
the ventral stream of visual cortices. In the monkey brain, this area receives strong feedforward
connections from the primary visual cortex (V1) and sends strong projections to other
secondary visual cortices (V3, V4, and V5).[30][31] Most of the neurons of this area in primates are
tuned to simple visual characteristics such as orientation, spatial frequency, size, color, and
shape.[25][32][33] Anatomical studies implicate layer 3 of area V2 in visual-information processing.
In contrast to layer 3, layer 6 of the visual cortex is composed of many types of neurons, and
their response to visual stimuli is more complex.

In one study, the Layer 6 cells of the V2 cortex were found to play a very important role in the
storage of Object Recognition Memory as well as the conversion of short-term object memories
into long-term memories.[34]

Third visual cortex, including area V3

The term third visual complex refers to the region of cortex located immediately in front of V2,
which includes the region named visual area V3 in humans. The "complex" nomenclature is
justified by the fact that some controversy still exists regarding the exact extent of area V3, with
some researchers proposing that the cortex located in front of V2 may include two or three
functional subdivisions. For example, David Van Essen and others (1986) have proposed the
existence of a "dorsal V3" in the upper part of the cerebral hemisphere, which is distinct from the
"ventral V3" (or ventral posterior area, VP) located in the lower part of the brain. Dorsal and
ventral V3 have distinct connections with other parts of the brain, appear different in sections
stained with a variety of methods, and contain neurons that respond to different combinations of
visual stimulus (for example, colour-selective neurons are more common in the ventral V3).
Additional subdivisions, including V3A and V3B have also been reported in humans. These
subdivisions are located near dorsal V3, but do not adjoin V2.

Dorsal V3 is normally considered to be part of the dorsal stream, receiving inputs from V2 and
from the primary visual area and projecting to the posterior parietal cortex. It may be
anatomically located in Brodmann area 19. Braddick using fMRI has suggested that area
V3/V3A may play a role in the processing of global motion[35] Other studies prefer to consider
dorsal V3 as part of a larger area, named the dorsomedial area (DM), which contains a
representation of the entire visual field. Neurons in area DM respond to coherent motion of large
patterns covering extensive portions of the visual field (Lui and collaborators, 2006).

Ventral V3 (VP), has much weaker connections from the primary visual area, and stronger
connections with the inferior temporal cortex. While earlier studies proposed that VP contained
a representation of only the upper part of the visual field (above the point of fixation), more
recent work indicates that this area is more extensive than previously appreciated, and like other
visual areas it may contain a complete visual representation. The revised, more extensive VP is
referred to as the ventrolateral posterior area (VLP) by Rosa and Tweedale.[36]

V4

Visual area V4 is one of the visual areas in the extrastriate visual cortex. In macaques, it is
located anterior to V2 and posterior to posterior inferotemporal area (PIT). It comprises at least
four regions (left and right V4d, left and right V4v), and some groups report that it contains
rostral and caudal subdivisions as well. It is unknown whether the human V4 is as expansive as
that of the macaque homologue which is a subject of debate.[37]

V4 is the third cortical area in the ventral stream, receiving strong feedforward input from V2 and
sending strong connections to the PIT. It also receives direct input from V1, especially for central
space. In addition, it has weaker connections to V5 and dorsal prelunate gyrus (DP).

V4 is the first area in the ventral stream to show strong attentional modulation. Most studies
indicate that selective attention can change firing rates in V4 by about 20%. A seminal paper by
Moran and Desimone characterizing these effects was the first paper to find attention effects
anywhere in the visual cortex.[38]

Like V2, V4 is tuned for orientation, spatial frequency, and color. Unlike V2, V4 is tuned for object
features of intermediate complexity, like simple geometric shapes, although no one has
developed a full parametric description of the tuning space for V4. Visual area V4 is not tuned
for complex objects such as faces, as areas in the inferotemporal cortex are.

The firing properties of V4 were first described by Semir Zeki in the late 1970s, who also named
the area. Before that, V4 was known by its anatomical description, the prelunate gyrus. Originally,
Zeki argued that the purpose of V4 was to process color information. Work in the early 1980s
proved that V4 was as directly involved in form recognition as earlier cortical areas. This
research supported the two-streams hypothesis, first presented by Ungerleider and Mishkin in
1982.

Recent work has shown that V4 exhibits long-term plasticity,[39] encodes stimulus salience, is
gated by signals coming from the frontal eye fields,[40] and shows changes in the spatial profile
of its receptive fields with attention.

Middle temporal visual area (V5)

The middle temporal visual area (MT or V5) is a region of extrastriate visual cortex. In several
species of both New World monkeys and Old World monkeys the MT area contains a high
concentration of direction-selective neurons.[41] The MT in primates is thought to play a major
role in the perception of motion, the integration of local motion signals into global percepts, and
the guidance of some eye movements.[41]

Connections
…
MT is connected to a wide array of cortical and subcortical brain areas. Its input comes from
visual cortical areas V1, V2 and dorsal V3 (dorsomedial area),[42][43] the koniocellular regions of
the LGN,[44] and the inferior pulvinar.[45] The pattern of projections to MT changes somewhat
between the representations of the foveal and peripheral visual fields, with the latter receiving
inputs from areas located in the midline cortex and retrosplenial region.[46]

A standard view is that V1 provides the "most important" input to MT.[41] Nonetheless, several
studies have demonstrated that neurons in MT are capable of responding to visual information,
often in a direction-selective manner, even after V1 has been destroyed or inactivated.[47]
Moreover, research by Semir Zeki and collaborators has suggested that certain types of visual
information may reach MT before it even reaches V1.
MT sends its major output to areas located in the cortex immediately surrounding it, including
areas FST, MST, and V4t (middle temporal crescent). Other projections of MT target the eye
movement-related areas of the frontal and parietal lobes (frontal eye field and lateral
intraparietal area).

Function
…
The first studies of the electrophysiological properties of neurons in MT showed that a large
portion of the cells are tuned to the speed and direction of moving visual stimuli.[48][49]

Lesion studies have also supported the role of MT in motion perception and eye movements.[50]
Neuropsychological studies of a patient unable to see motion, seeing the world in a series of
static 'frames' instead, suggested that V5 in the primate is homologous to MT in the
human.[51][52]

However, since neurons in V1 are also tuned to the direction and speed of motion, these early
results left open the question of precisely what MT could do that V1 could not. Much work has
been carried out on this region, as it appears to integrate local visual motion signals into the
global motion of complex objects.[53] For example, lesion to the V5 leads to deficits in perceiving
motion and processing of complex stimuli. It contains many neurons selective for the motion of
complex visual features (line ends, corners). Microstimulation of a neuron located in the V5
affects the perception of motion. For example, if one finds a neuron with preference for upward
motion in a monkey's V5 and stimulates it with an electrode, then the monkey becomes more
likely to report 'upward' motion when presented with stimuli containing 'left' and 'right' as well as
'upward' components.[54]

There is still much controversy over the exact form of the computations carried out in area
MT[55] and some research suggests that feature motion is in fact already available at lower
levels of the visual system such as V1. [56][57]

Functional organization …
MT was shown to be organized in direction columns.[58] DeAngelis argued that MT neurons were
also organized based on their tuning for binocular disparity.[59]

V6
The dorsomedial area (DM) also known as V6, appears to respond to visual stimuli associated
with self-motion[60] and wide-field stimulation.[61] V6 is a subdivision of the visual cortex of
primates first described by John Allman and Jon Kaas in 1975.[62] V6 is located in the dorsal part
of the extrastriate cortex, near the deep groove through the centre of the brain (medial
longitudinal fissure), and typically also includes portions of the medial cortex, such as the
parieto-occipital sulcus (POS).[61]: 7970  DM contains a topographically organized representation
of the entire field of vision.[61]: 7970 

There are similarities between the visual area V5 and V6 of the common marmoset. Both areas
receive direct connections from the primary visual cortex.[61]: 7971  And both have a high myelin
content, a characteristic that is usually present in brain structures involved in fast transmission
of information.[63]

For many years, it was considered that DM only existed in New World monkeys. However, more
recent research has suggested that DM also exists in Old World monkeys and humans.[61]: 7972 
V6 is also sometimes referred to as the parieto-occipital area (PO), although the correspondence
is not exact.[64][65]

Properties
…
Neurons in area DM/V6 of night monkeys and common marmosets have unique response
properties, including an extremely sharp selectivity for the orientation of visual contours, and
preference for long, uninterrupted lines covering large parts of the visual field.[66][67]

However, in comparison with area MT, a much smaller proportion of DM cells shows selectivity
for the direction of motion of visual patterns.[68] Another notable difference with area MT is that
cells in DM are attuned to low spatial frequency components of an image, and respond poorly to
the motion of textured patterns such as a field of random dots.[68] These response properties
suggest that DM and MT may work in parallel, with the former analyzing self-motion relative to
the environment, and the latter analyzing the motion of individual objects relative to the
background.[68]

Recently, an area responsive to wide-angle flow fields has been identified in the human and is
thought to be a homologue of macaque area V6.[69]

Pathways …
The connections and response properties of cells in DM/ V6 suggest that this area is a key node
in a subset of the 'dorsal stream', referred to by some as the 'dorsomedial pathway'. This
pathway is likely to be important for the control of skeletomotor activity, including postural
reactions and reaching movements towards objects[65] The main 'feedforward' connection of DM
is to the cortex immediately rostral to it, in the interface between the occipital and parietal lobes
(V6A). This region has, in turn, relatively direct connections with the regions of the frontal lobe
that control arm movements, including the premotor cortex.

See also

Cortical area

Cortical blindness

Feature integration theory

List of regions in the human brain

Retinotopy

Visual processing

Visual feature array

Complex cell

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External links

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