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The visual system is the physiological basis of visual perception (the ability to ). The system detects, transduces and interprets information concerning within the to construct an and build a of the surrounding environment. The visual system is associated with the and functionally divided into the system (including and ) and the system (including the and ).

The visual system performs a number of complex tasks based on the image forming functionality of the eye, including the formation of monocular images, the neural mechanisms underlying and assessment of distances to () and between objects, motion perception, pattern recognition, accurate motor coordination under visual guidance, and . Together, these facilitate higher order tasks, such as object identification. The neuropsychological side of visual information processing is known as visual perception, an abnormality of which is called visual impairment, and a complete absence of which is called . The visual system also has several non-image forming visual functions, independent of visual perception, including the pupillary light reflex and circadian photoentrainment.

This article describes the human visual system, which is representative of , and to a lesser extent the visual system.


System overview

Optical
Together, the and lens refract light into a small image and shine it on the . The retina transduces this image into electrical pulses using rods and cones. The then carries these pulses through the . Upon reaching the the nerve fibers decussate (left becomes right). The fibers then branch and terminate in three places."How the Human Eye Sees." . Ed. Alan Kozarsky. WebMD, 3 October 2015. Web. 27 March 2016.Than, Ker. "How the Human Eye Works." . TechMedia Network, 10 February 2010. Web. 27 March 2016."How the Human Eye Works | Cornea Layers/Role | Light Rays." NKCF. The Gavin Herbert Eye Institute. Web. 27 March 2016.Albertine, Kurt. Barron's Anatomy Flash CardsTillotson, Joanne. McCann, Stephanie. Kaplan's Medical Flashcards. April 2, 2013."Optic Chiasma." Optic Chiasm Function, Anatomy & Definition. Healthline Medical Team, 9 March 2015. Web. 27 March 2016.Jefferey, G., and M. M. Neveu. "Chiasm Formation in Man Is Fundamentally Different from That in the Mouse." Nature.com. Nature Publishing Group, 21 March 2007. Web. 27 March 2016.


Neural
Most of the optic nerve fibers end in the lateral geniculate nucleus (LGN). Before the LGN forwards the pulses to V1 of the visual cortex (primary) it gauges the range of objects and tags every major object with a velocity tag. These tags predict object movement.

The LGN also sends some fibers to V2 and V3.Card, J. Patrick, and Robert Y. Moore. "Organization of Lateral Geniculate-hypothalamic Connections in the Rat." Wiley Online Library. 1 June. 1989. Web. 27 March 2016.

V1 performs edge-detection to understand spatial organization (initially, 40 milliseconds in, focusing on even small spatial and color changes. Then, 100 milliseconds in, upon receiving the translated LGN, V2, and V3 info, also begins focusing on global organization). V1 also creates a bottom-up to guide attention or .

(2014). 9780199564668, Oxford University Press. .

V2 both forwards (direct and via ) pulses to V1 and receives them. Pulvinar is responsible for and visual attention. V2 serves much the same function as V1, however, it also handles illusory contours, determining depth by comparing left and right pulses (2D images), and foreground distinguishment. V2 connects to V1 - V5.

V3 helps process '' (direction and speed) of objects. V3 connects to V1 (weak), V2, and the inferior temporal cortex.Catani, Marco, and Derek K. Jones. "Brain." Occipito‐temporal Connections in the Human Brain. 23 June 2003. Web. 27 March 2016.

V4 recognizes simple shapes, and gets input from V1 (strong), V2, V3, LGN, and pulvinar. V5's outputs include V4 and its surrounding area, and eye-movement motor cortices (frontal eye-field and lateral intraparietal area).

V5's functionality is similar to that of the other V's, however, it integrates local object motion into global motion on a complex level. V6 works in conjunction with V5 on motion analysis. V5 analyzes self-motion, whereas V6 analyzes motion of objects relative to the background. V6's primary input is V1, with V5 additions. V6 houses the for vision. V6 outputs to the region directly around it (V6A). V6A has direct connections to arm-moving cortices, including the .

The inferior temporal gyrus recognizes complex shapes, objects, and faces or, in conjunction with the , creates new .Moser, May-Britt, and Edvard I. Moser. "Functional Differentiation in the Hippocampus." Wiley Online Library. 1998. Web. 27 March 2016. The is seven unique nuclei. Anterior, posterior and medial pretectal nuclei inhibit pain (indirectly), aid in REM, and aid the accommodation reflex, respectively. The Edinger-Westphal nucleus moderates and aids (since it provides parasympathetic fibers) in convergence of the eyes and lens adjustment.Reiner, Anton, and Harvey J. Karten. "Parasympathetic Ocular Control — Functional Subdivisions and Circuitry of the Avian Nucleus of Edinger-Westphal."Science Direct. 1983. Web. 27 March 2016. Nuclei of the optic tract are involved in smooth pursuit eye movement and the accommodation reflex, as well as REM.

The suprachiasmatic nucleus is the region of the that halts production of (indirectly) at first light.


Structure
  • The eye, especially the retina
  • The optic nerve
  • The optic chiasma
  • The optic tract
  • The lateral geniculate body
  • The optic radiation
  • The
  • The visual association cortex.

These are components of the visual pathway, also called the optic pathway, that can be divided into anterior and posterior visual pathways. The anterior visual pathway refers to structures involved in vision before the lateral geniculate nucleus. The posterior visual pathway refers to structures after this point.


Eye
Light entering the eye is as it passes through the . It then passes through the (controlled by the iris) and is further refracted by the lens. The cornea and lens act together as a compound lens to project an inverted image onto the retina.


Retina
The retina consists of many photoreceptor cells which contain particular called . In humans, two types of opsins are involved in conscious vision: and . (A third type, in some retinal ganglion cells (RGC), part of the mechanism, is probably not involved in conscious vision, as these RGC do not project to the lateral geniculate nucleus but to the .) An opsin absorbs a (a particle of light) and transmits a signal to the cell through a signal transduction pathway, resulting in hyper-polarization of the photoreceptor.

Rods and cones differ in function. Rods are found primarily in the periphery of the retina and are used to see at low levels of light. Each human eye contains 120 million rods. Cones are found primarily in the center (or ) of the retina. There are three types of cones that differ in the of light they absorb; they are usually called short or blue, middle or green, and long or red. Cones mediate day vision and can distinguish and other features of the visual world at medium and high light levels. Cones are larger and much less numerous than rods (there are 6-7 million of them in each human eye).

In the retina, the photoreceptors directly onto bipolar cells, which in turn synapse onto ganglion cells of the outermost layer, which then conduct action potentials to the . A significant amount of visual processing arises from the patterns of communication between in the retina. About 130 million photo-receptors absorb light, yet roughly 1.2 million of ganglion cells transmit information from the retina to the brain. The processing in the retina includes the formation of center-surround of bipolar and ganglion cells in the retina, as well as convergence and divergence from photoreceptor to bipolar cell. In addition, other neurons in the retina, particularly and , transmit information laterally (from a neuron in one layer to an adjacent neuron in the same layer), resulting in more complex receptive fields that can be either indifferent to color and sensitive to motion or sensitive to color and indifferent to motion.


Mechanism of generating visual signals
The retina adapts to change in light through the use of the rods. In the dark, the has a bent shape called cis-retinal (referring to a cis conformation in one of the double bonds). When light interacts with the retinal, it changes conformation to a straight form called trans-retinal and breaks away from the opsin. This is called bleaching because the purified changes from violet to colorless in the light. At baseline in the dark, the rhodopsin absorbs no light and releases , which inhibits the bipolar cell. This inhibits the release of neurotransmitters from the bipolar cells to the ganglion cell. When there is light present, glutamate secretion ceases, thus no longer inhibiting the bipolar cell from releasing neurotransmitters to the ganglion cell and therefore an image can be detected.Saladin, Kenneth D. Anatomy & Physiology: The Unity of Form and Function. 5th ed. New York: McGraw-Hill, 2010.

The final result of all this processing is five different populations of ganglion cells that send visual (image-forming and non-image-forming) information to the brain:

  1. M cells, with large center-surround receptive fields that are sensitive to , indifferent to color, and rapidly adapt to a stimulus;
  2. P cells, with smaller center-surround receptive fields that are sensitive to color and ;
  3. K cells, with very large center-only receptive fields that are sensitive to color and indifferent to shape or depth;
  4. another population that is intrinsically photosensitive; and
  5. a final population that is used for eye movements.

A 2006 University of Pennsylvania study calculated the approximate bandwidth of human retinas to be about 8,960 per second, whereas retinas transfer at about 875 kilobits.

In 2007 Zaidi and co-researchers on both sides of the Atlantic studying patients without rods and cones, discovered that the novel photoreceptive ganglion cell in humans also has a role in conscious and unconscious visual perception. The peak spectral sensitivity was 481 nm. This shows that there are two pathways for vision in the retina – one based on classic photoreceptors (rods and cones) and the other, newly discovered, based on photo-receptive ganglion cells which act as rudimentary visual brightness detectors.


Photochemistry
The functioning of a is often compared with the workings of the eye, mostly since both focus light from external objects in the field of view onto a light-sensitive medium. In the case of the camera, this medium is film or an electronic sensor; in the case of the eye, it is an array of visual receptors. With this simple geometrical similarity, based on the laws of optics, the eye functions as a , as does a CCD camera.

In the visual system, retinal, technically called 1 or "retinaldehyde", is a light-sensitive molecule found in the rods and cones of the . Retinal is the fundamental structure involved in the transduction of into visual signals, i.e. nerve impulses in the ocular system of the central nervous system. In the presence of light, the retinal molecule changes configuration and as a result, a is generated.


Optic nerve
(top), crossing at the , joining left and right eye information in the , and layering left and right visual stimuli in the lateral geniculate nucleus. V1 in red at bottom of image. (1543 image from ' Fabrica)]] The information about the image via the eye is transmitted to the brain along the . Different populations of ganglion cells in the retina send information to the brain through the optic nerve. About 90% of the in the optic nerve go to the lateral geniculate nucleus in the . These axons originate from the M, P, and K ganglion cells in the retina, see above. This parallel processing is important for reconstructing the visual world; each type of information will go through a different route to . Another population sends information to the superior colliculus in the , which assists in controlling eye movements ()
(2025). 9780323013208, Mosby.
as well as other motor responses.

A final population of photosensitive ganglion cells, containing for , sends information via the retinohypothalamic tract to the (), to several structures involved in the control of circadian rhythms and such as the suprachiasmatic nucleus (the biological clock), and to the ventrolateral preoptic nucleus (a region involved in ). A recently discovered role for photoreceptive ganglion cells is that they mediate conscious and unconscious vision – acting as rudimentary visual brightness detectors as shown in rodless coneless eyes.


Optic chiasm
The optic nerves from both eyes meet and cross at the optic chiasm,
(1997). 9780292781498, University of Texas Press.
at the base of the of the brain. At this point, the information coming from both eyes is combined and then splits according to the . The corresponding halves of the field of view (right and left) are sent to the left and right halves of the brain, respectively, to be processed. That is, the right side of primary visual cortex deals with the left half of the field of view from both eyes, and similarly for the left brain. A small region in the center of the field of view is processed redundantly by both halves of the brain.


Optic tract
Information from the right visual field (now on the left side of the brain) travels in the left optic tract. Information from the left visual field travels in the right optic tract. Each optic tract terminates in the lateral geniculate nucleus (LGN) in the thalamus.


Lateral geniculate nucleus

The lateral geniculate nucleus (LGN) is a sensory relay nucleus in the thalamus of the brain. The LGN consists of six layers in and other starting from , including and . Layers 1, 4, and 6 correspond to information from the contralateral (crossed) fibers of the nasal retina (temporal visual field); layers 2, 3, and 5 correspond to from the ipsilateral (uncrossed) fibers of the temporal retina (nasal visual field).

Layer one contains M cells, which correspond to the M (magnocellular) cells of the optic nerve of the opposite eye and are concerned with depth or motion. Layers four and six of the LGN also connect to the opposite eye, but to the P cells (color and edges) of the optic nerve. By contrast, layers two, three and five of the LGN connect to the M cells and P (parvocellular) cells of the optic nerve for the same side of the brain as its respective LGN.

Spread out, the six layers of the LGN are the area of a and about three times its thickness. The LGN is rolled up into two about the size and shape of two small birds' eggs. In between the six layers are smaller cells that receive information from the K cells (color) in the retina. The neurons of the LGN then relay the visual image to the primary visual cortex (V1) which is located at the back of the brain (posterior end) in the in and close to the . The LGN is not just a simple relay station, but it is also a center for processing; it receives reciprocal input from the and subcortical layers and reciprocal innervation from the visual cortex.


Optic radiation
The optic radiations, one on each side of the brain, carry information from the thalamic lateral geniculate nucleus to layer 4 of the . The P layer neurons of the LGN relay to V1 layer 4C β. The M layer neurons relay to V1 layer 4C α. The K layer neurons in the LGN relay to large neurons called blobs in layers 2 and 3 of V1.

There is a direct correspondence from an angular position in the of the eye, all the way through the optic tract to a nerve position in V1 up to V4, i.e. the primary visual areas. After that, the visual pathway is roughly separated into a ventral and dorsal pathway.


Visual cortex
The visual cortex is responsible for processing the visual image. It lies at the rear of the brain (highlighted in the image), above the . The region that receives information directly from the LGN is called the primary visual cortex (also called V1 and striate cortex). It creates a bottom-up saliency map of the visual field to guide attention or eye gaze to salient visual locations. Hence selection of visual input information by attention starts at V1 along the visual pathway.

Visual information then flows through a cortical hierarchy. These areas include V2, V3, V4 and area V5/MT. (The exact connectivity depends on the species of the animal.) These secondary visual areas (collectively termed the extrastriate visual cortex) process a wide variety of visual primitives. Neurons in V1 and V2 respond selectively to bars of specific orientations, or combinations of bars. These are believed to support edge and corner detection. Similarly, basic information about color and motion is processed here.

(2025). 9780838577011, McGraw-Hill. .

Heider, et al. (2002) found that neurons involving V1, V2, and V3 can detect stereoscopic illusory contours; they found that stereoscopic stimuli subtending up to 8° can activate these neurons. Heider, Barbara; Spillmann, Lothar; Peterhans, Esther (2002) "Stereoscopic Illusory Contours— Cortical Neuron Responses and Human Perception" J. Cognitive Neuroscience 14:7 pp.1018-29 accessdate=2014-05-18


Visual association cortex
As visual information passes forward through the visual hierarchy, the complexity of the neural representations increases. Whereas a V1 neuron may respond selectively to a line segment of a particular orientation in a particular location, neurons in the lateral occipital complex respond selectively to a complete object (e.g., a figure drawing), and neurons in the visual association cortex may respond selectively to human faces, or to a particular object.

Along with this increasing complexity of neural representation may come a level of specialization of processing into two distinct pathways: the and the (the Two Streams hypothesis, first proposed by Ungerleider and Mishkin in 1982). The dorsal stream, commonly referred to as the "where" stream, is involved in spatial attention (covert and overt), and communicates with regions that control eye movements and hand movements. More recently, this area has been called the "how" stream to emphasize its role in guiding behaviors to spatial locations. The ventral stream, commonly referred to as the "what" stream, is involved in the recognition, identification and categorization of visual stimuli.

However, there is still much debate about the degree of specialization within these two pathways, since they are in fact heavily interconnected.

proposed the efficient coding hypothesis in 1961 as a theoretical model of sensory coding in the .Barlow, H. (1961) "Possible principles underlying the transformation of sensory messages" in Sensory Communication, MIT Press Limitations in the applicability of this theory in the primary visual cortex (V1) motivated the V1 Saliency Hypothesis that V1 creates a bottom-up saliency map to guide attention exogenously. With attentional selection as a center stage, vision is seen as composed of encoding, selection, and decoding stages.

(2025). 9780198829362, Oxford University Press.

The default mode network is a network of brain regions that are active when an individual is awake and at rest. The visual system's default mode can be monitored during resting state fMRI: Fox, et al. (2005) found that " the human brain is intrinsically organized into dynamic, anticorrelated functional networks", in which the visual system switches from resting state to attention.

In the , the lateral and ventral intraparietal cortex are involved in visual attention and saccadic eye movements. These regions are in the intraparietal sulcus (marked in red in the adjacent image).


Development

Infancy
Newborn infants have limited .
(2025). 9781556429569, SLACK. .
One study found that 74% of newborns can distinguish red, 36% green, 25% yellow, and 14% blue. After one month, performance "improved somewhat." Infant's eyes do not have the ability to accommodate. Pediatricians are able to perform non-verbal testing to assess of a newborn, detect and , and evaluate the eye teaming and alignment. Visual acuity improves from about 20/400 at birth to approximately 20/25 at 6 months of age. This happens because the nerve cells in the and brain that control vision are not fully developed.


Childhood and adolescence
, focus, tracking and other aspects of vision continue to develop throughout early and middle childhood. From recent studies in the and there is some evidence that the amount of time school aged children spend outdoors, in natural light, may have some impact on whether they develop . The condition tends to get somewhat worse through childhood and adolescence, but stabilizes in adulthood. More prominent myopia (nearsightedness) and astigmatism are thought to be inherited. Children with this condition may need to wear glasses.


Adulthood
Vision is often one of the first senses affected by aging. A number of changes occur with aging:
  • Over time, the lens becomes yellowed and may eventually become brown, a condition known as brunescence or brunescent . Although many factors contribute to yellowing, lifetime exposure to ultraviolet light and are two main causes.
  • The lens becomes less flexible, diminishing the ability to accommodate ().
  • While a healthy adult pupil typically has a size range of 2–8 mm, with age the range gets smaller, trending towards a moderately small diameter.
  • On average declines with age. However, there are a number of age-related conditions that can cause excessive tearing.


Other functions

Balance
Along with and vestibular function, the visual system plays an important role in the ability of an individual to control balance and maintain an upright posture. When these three conditions are isolated and balance is tested, it has been found that vision is the most significant contributor to balance, playing a bigger role than either of the two other intrinsic mechanisms. The clarity with which an individual can see his environment, as well as the size of the visual field, the susceptibility of the individual to light and glare, and poor depth perception play important roles in providing a feedback loop to the brain on the body's movement through the environment. Anything that affects any of these variables can have a negative effect on balance and maintaining posture. This effect has been seen in research involving elderly subjects when compared to young controls, in patients compared to age matched controls, patients pre and post surgery, and even something as simple as wearing safety goggles. (one eyed vision) has also been shown to negatively impact balance, which was seen in the previously referenced cataract and glaucoma studies, as well as in healthy children and adults.

According to Pollock et al. (2010) is the main cause of specific visual impairment, most frequently visual field loss (homonymous hemianopia, a visual field defect). Nevertheless, evidence for the efficacy of cost-effective interventions aimed at these visual field defects is still inconsistent.


Clinical significance
'''
From top to bottom:
1. Complete loss of vision, right eye
2. Bitemporal hemianopia
3. Homonymous hemianopsia
4.
5&6. Quadrantanopia with ]]

Proper function of the visual system is required for sensing, processing, and understanding the surrounding environment. Difficulty in sensing, processing and understanding light input has the potential to adversely impact an individual's ability to communicate, learn and effectively complete routine tasks on a daily basis.

In children, early diagnosis and treatment of impaired visual system function is an important factor in ensuring that key social, academic and speech/language developmental milestones are met.

is clouding of the lens, which in turn affects vision. Although it may be accompanied by yellowing, clouding and yellowing can occur separately. This is typically a result of ageing, disease, or drug use.

is a visual condition that causes . The eye's lens becomes too inflexible to accommodate to normal reading distance, focus tending to remain fixed at long distance.

is a type of blindness that begins at the edge of the visual field and progresses inward. It may result in . This typically involves the outer layers of the optic nerve, sometimes as a result of buildup of fluid and excessive pressure in the eye.

(2025). 9781935555162, Harvard Health Publications. .

is a type of blindness that produces a small blind spot in the visual field typically caused by injury in the primary visual cortex.

Homonymous hemianopia is a type of blindness that destroys one entire side of the visual field typically caused by injury in the primary visual cortex.

is a type of blindness that destroys only a part of the visual field typically caused by partial injury in the primary visual cortex. This is very similar to homonymous hemianopia, but to a lesser degree.

, or face blindness, is a brain disorder that produces an inability to recognize faces. This disorder often arises after damage to the fusiform face area.

, or visual-form agnosia, is a brain disorder that produces an inability to recognize objects. This disorder often arises after damage to the .


Other animals
Different are able to see different parts of the ; for example, can see into the , while can accurately target prey with their , which are sensitive to infrared radiation. The possesses arguably the most complex visual system of any species. The eye of the mantis shrimp holds 16 color receptive cones, whereas humans only have three. The variety of cones enables them to perceive an enhanced array of colors as a mechanism for mate selection, avoidance of predators, and detection of prey. Swordfish also possess an impressive visual system. The eye of a can generate to better cope with detecting their at depths of 2000 feet.David Fleshler(10-15-2012) South Florida Sun-Sentinel ,

Many , such as Acromegalomma interruptum which live in tubes on the sea floor of the Great Barrier Reef, have evolved compound eyes on their tentacles, which they use to detect encroaching movement. If movement is detected, the fan worms will rapidly withdraw their tentacles. Bok, et al., have discovered opsins and in the fan worm's eyes, which were previously only seen in simple photoreceptors in the brains of some , as opposed to the receptors in the eyes of most invertebrates. cited by Evolution of fan worm eyes (August 1, 2017) Phys.org

Only (African) and apes (, , ) have the same kind of three-cone photoreceptor color vision humans have, while lower primate (South American) monkeys (, , ) have a two-cone photoreceptor kind of color vision.

(2025). 9780810995543, Abrams.

Biologists have determined that humans have extremely good vision compared to the overwhelming majority of animals, particularly in daylight, surpassed only by a few large species of predatory birds. Other animals such as are thought to rely more on senses other than vision, which in turn may be better developed than in humans.


History
In the second half of the 19th century, many motifs of the nervous system were identified such as the neuron doctrine and brain localization, which related to the being the basic unit of the nervous system and functional localisation in the brain, respectively. These would become tenets of the fledgling and would support further understanding of the visual system.

The notion that the is divided into functionally distinct cortices now known to be responsible for capacities such as (somatosensory cortex), movement (), and vision (), was first proposed by Franz Joseph Gall in 1810. Evidence for functionally distinct areas of the brain (and, specifically, of the cerebral cortex) mounted throughout the 19th century with discoveries by of the (1861), and and of the motor cortex (1871). Based on selective damage to parts of the brain and the functional effects of the resulting , proposed that visual function was localized to the of the brain in 1876. In 1881, more accurately located vision in the , where the primary visual cortex is now known to be.

In 2014, a textbook "Understanding vision: theory, models, and data" illustrates how to link neurobiological data and visual behavior/psychological data through theoretical principles and computational models.


See also

Further reading
  • Heiting, G., (2011). Your infant's vision Development. Retrieved February 27, 2012 from http://www.allaboutvision.com/parents/infants.htm
  • (1995). 9780716760092, Scientific American Library.
  • (2025). 9781429242288, Worth Publishers.
  • (2025). 9780262514620, The MIT Press.
  • . (H.D. Steklis and J. Erwin, editors.) pp. 203–278.
  • Https://web.archive.org/web/20111117045917/http://www.realage.com/check-your-health/eye-health/aging-eye
  • (2025). 9780521883191, Cambridge University Press.
  • .


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