The
visual cortex of the
brain is the part of the
cerebral cortex responsible for processing visual information. This article addresses the ventral/dorsal model of the visual cortex. Another model for the perceptual/conceptual neuropsychological model of the visual cortex was studied by Raftopolous.
[1] In Russian neuropsychology, yet another model was discussed by
Alexander Luria for the anterior/posterior approach to understanding the visual cortex.
[2] The visual cortex is located in the
occipital lobe (one of the four major lobes of the cerebral cortex) which is in turn located at the back of the head or skull. The visual cortex is made up of
Brodmann area 17 (the
primary visual cortex), and
Brodmann areas 18 and
19, the
extrastriate cortical areas.
[citation needed]
The primary (parts of the cortex that receive sensory inputs from the
thalamus) visual cortex is also known as
V1,
Visual area
one, and the
striate cortex. The
extrastriate areas consist of visual areas two (
V2), three (
V3), four (
V4), and five (
V5). Both
hemispheres of a brain contain a visual cortex; the left hemisphere visual cortex receives signals from the right
visual field, and the right visual cortex from the left visual field.
Introduction[edit]
This article addresses the ventral/dorsal model of the visual cortex. Another model for the perceptual/conceptual neuropsychological model of the visual cortex was studied by Raftopolous.
[1] In Russian neurospychology, yet another model was discussed by
Alexander Luria for the anterior/posterior approach to understanding the visual cortex.
[2] The primary visual cortex, V1, is the koniocortex (sensory type)
[citation needed] 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.
Each V1 transmits information to two primary pathways, called the ventral stream and the dorsal stream.
- 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 Visual area MT (middle temporal/ 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
[4] 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. Although these ideas are contentious among some vision scientists and psychologists, the proposed division of labor between
vision-for-perception and
vision-for-action is supported by a broad range of findings from
neuropsychology,
neurophysiology, and
neuroimaging. Some of the most controversial evidence comes from studies of visuomotor behaviour in normal observers. For example, 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] Other work such as the one from Scharnowski and Gegenfurtner
[6] suggests that both the action and perception systems are equally fooled by such illusions. More recent 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]
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.
Current research[edit]
Research on the primary visual cortex can involve recording
action potentials from electrodes within the brain of
cats,
ferrets,
rats,
mice, or
monkeys, or through recording intrinsic optical signals from animals or
EEG,
MEG, or
fMRI signals from human and monkey V1.
One recent discovery concerning the human V1 is that signals measured by
fMRI show very large attentional modulation. This result is consistent with another recent electrophysiology study. The study found that although V1 processes visual information before V2 and V4, the attentional modulation occurs in V4 followed by V2 and then finally in V1 suggesting attentional modulation occurs through
feedback from higher-level visual areas to lower-level visual areas.
[10] Other current work on V1 seeks to fully characterize its tuning properties, and to use it as a model area for the canonical cortical circuit.
Lesions to primary visual cortex usually lead to a
scotoma, or hole in the visual field. Note that patients with scotomas are often able to make use of visual information presented to their scotomas, despite being unable to consciously perceive it. This phenomenon has been partially covered in one aspect of what
Larry Weiskrantz of Oxford university called
blindsight, and is widely studied by scientists who are also interested in the neural correlates of
consciousness.
Primary visual cortex (V1)[edit]
The
primary visual cortex is the best-studied visual area in the
brain. In all mammals studied, it is located in the posterior pole of the occipital cortex (the occipital cortex is responsible for processing
visual stimuli). It 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 functionally defined primary visual cortex is approximately equivalent to the anatomically defined
striate cortex. The name "striate cortex" is derived from the
line of Gennari, a distinctive stripe visible to the naked eye that represents
myelinated axons from the
lateral geniculate bodyterminating in layer 4 of the
gray matter.
The primary visual cortex is divided into six functionally distinct layers, labeled 1 through 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 most
magnocellular input from the LGN, while layer 4Cβ receives input from
parvocellular pathways.
The occipital cortex where the visual cortex resides is the smallest of the four cortices of the human brain, which also includes the
temporal cortex,
parietal cortex, and
frontal cortex. The average number of neurons in the adult human primary visual cortex, in each hemisphere, has been estimated at around 140 million.
[11]
Function[edit]
V1 has a very well-defined map of the spatial information in vision. For example, in humans, the upper bank of the
calcarine sulcusresponds 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 transformation of the visual image from
retina to V1. The correspondence between a given location in V1 and in the subjective visual field is very precise: even the
blind spots 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 in the retina, a large portion of V1 is mapped to the small, central portion of visual field, a phenomenon known as
cortical magnification.
[12] Perhaps for the purpose of accurate spatial encoding, neurons in V1 have the smallest
receptive field size 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. Furthermore, individual V1 neurons in human 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
[citation needed]. 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).
[13] 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 divide 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).
Visual area V2, or
secondary visual cortex, also called
prestriate cortex,
[14] 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,
binocular disparity,
[15] and whether the stimulus is part of the figure or the ground (Qiu and von der Heydt, 2005). 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.
[16] 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).
[17][18] Most of the neurons of this area are tuned to simple visual characteristics such as orientation, spatial frequency, size, color, and shape.
[19][20][21] V2 cells also respond to various complex shape characteristics, such as the orientation of illusory contours
[21] and whether the stimulus is part of the figure or the ground.
[22] 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 a recent 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.
[23]
Third visual complex, including area V3[edit]
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[24] 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.
[25]
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 what the human
homologue of V4 is, and this issue is currently the subject of much scrutiny.
[26]
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 inputs 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.
[27][28]
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 cortexare.
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.
[citation needed] 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, encodes stimulus salience, is gated by signals coming from the
frontal eye fields and shows changes in the spatial profile of its receptive fields with attention.
[citation needed]
Visual area V5, also known as
visual area MT (middle temporal), is a region of extrastriate visual cortex that 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.
[29]
Connections[edit]
MT is connected to a wide array of cortical and subcortical brain areas. Its inputs include the visual cortical areas V1, V2, and dorsal V3 (
dorsomedial area),
[30][31] the
koniocellular regions of the
LGN,
[32] and the
inferior pulvinar.
[33] 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.
[34]
A standard view is that V1 provides the "most important" input to MT.
[29] 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.
[35] 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 outputs 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[edit]
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
[36][37]These results suggested that MT plays a significant role in the processing of visual motion.
Lesion studies have also supported the role of MT in motion perception and eye movements.
[38] 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.
[39][40]
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.
[41] 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.
[42]
There is still much controversy over the exact form of the computations carried out in area MT
[43] and some research suggests that feature motion is in fact already available at lower levels of the visual system such as V1.
[44] [45]
Functional organization[edit]
MT was shown to be organized in direction columns.
[46] DeAngelis argued that MT neurons were also organized based on their tuning for binocular disparity.
[47]
The
dorsomedial area (DM) also known as
V6, appears to respond to visual stimuli associated with self-motion
[48] and wide-field stimulation.
[49] V6, is a subdivision of the visual cortex of primates first described by
John Allman and
Jon Kaas in 1975.
[50] 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. DM contains a topographically organized representation of the entire field of vision. Like the middle temporal area V5, DM receives direct connections from the
primary visual cortex. Also similar to V5, DM is also characterized by high
myelin content, a characteristic that is usually present in brain structures involved in fast transmission of information.
[51] 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 perhaps humans. V6 is also sometimes referred to as the parietooccipital area (PO), although the correspondence is not exact.
[52][53]
Properties[edit]
Neurons in area DM/V6 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.
[54][55] However, in comparison with area MT, a much smaller proportion of DM cells shows selectivity for the direction of motion of visual patterns. Another notable difference 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. In contrast, cells in MT are often strongly responsive to such stimuli. 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.
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.
[56]
Pathways[edit]
The connections and response properties of cells in DM/ V6 suggest that this area is a key node in a sub-set 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
[53] 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.
