The visual system is the part of the nervous system which allows organisms to see. It interprets the information from visible light to build a representation of the world surrounding the body. The visual system has the complex task of (re)constructing a three dimensional world from a two dimensional projection of that world. The psychological manifestation of visual information is known as visual perception.

Introduction

Optical layout of the eye
The image projected onto the retina is inverted due to the optics of the eye.

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

Different species are able to see different parts of the light spectrum; for example, bees can see into the ultraviolet,^[1] while pit vipers can accurately target prey with their infrared imaging sensors.^[2]

History

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

The notion that the cerebral cortex is divided into functionally distinct cortices now known to be responsible for capacities such as touch (somatosensory cortex), movement (motor cortex), and vision (visual cortex), was first proposed by Franz Joseph Gall in 1810.^[3] Evidence for functionally distinct areas of the brain (and, specifically, of the cerebral cortex) mounted throughout the 19th century with discoveries by Paul Broca of the language centre (1861), and Gustav Fritsch and Edouard Hitzig of the motor cortex (1871).^[3][4] Based on selective damage to parts of the brain and the functional effects this would produce (lesion studies), David Ferrier proposed that visual function was localised to the parietal lobe of the brain in 1876.^[4] In 1881, Hermann Munk more accurately located vision in the occipital lobe, where the primary visual cortex is now known to be.^[4]

Biology of the visual system

Eye

The eye is a complex biological device. The functioning of a camera is often compared with the workings of the eye, mostly since both focus light from external objects in the visual field 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 transducer, as does a CCD camera.

Light entering the eye is refracted as it passes through the cornea. It then passes through the pupil (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.

S. Ramón y Cajal, Structure of the Mammalian Retina, 1900

Retina

The retina consists of a large number of photoreceptor cells which contain a particular protein molecule called an opsin. In humans, two types of opsins are involved in vision: rod opsins and cone opsins. (A third type, melanopsin in some of the retinal ganglion cells, part of the body clock mechanism, is probably not involved in vision.) An opsin absorbs a photon (a particle of light) and transmits a signal to the cell through a signal transduction pathway, resulting in hyperpolarization of the photoreceptor. (For more information, see Photoreceptor cell).

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. Cones are found primarily in the center (or fovea) of the retina. There are three types of cones that differ in the wavelengths of light they absorb; they are usually called short or blue, middle or green, and long or red. Cones are used primarily to distinguish color and other features of the visual world at normal levels of light.

In the retina, the photoreceptors synapse directly onto bipolar cells, which in turn synapse onto ganglion cells of the outermost layer, which will then conduct action potentials to the brain. A significant amount of visual processing arises from the patterns of communication between neurons in the retina. About 130 million photoreceptors absorb light, yet roughly 1.2 million axons of ganglion cells transmit information from the retina to the brain. The processing in the retina includes the formation of center-surround receptive fields 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 horizontal and amacrine cells, 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.

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 depth, indifferent to color, and rapidly adapt to a stimulus;
2. P cells, with smaller center-surround receptive fields that are sensitive to color and shape;
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 8960 kilobits per second, whereas guinea pig retinas transfer at about 875 kilobits. ^[5]

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.^[6][7][8] The peak spectral sensitivity was 481nm. This shows that there are two pathways for sight in the retina - one based on classic photoreceptors (rods and cones) and the other, newly discovered, based on photoreceptive ganglion cells which act as rudimentary visual brightness detectors.

Photochemistry

In the visual system, retinal, technically called retinene₁ or "retinaldehyde", is a light-sensitive retinene molecule found in the rods and cones of the retina. Retinal is the fundamental structure involved in the transduction of light 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 nerve impulse is generated.

Fibers to thalamus

Optic nerve

Information flow from the eyes (top), crossing at the optic chiasma, joining left and right eye information in the optic tract, and layering left and right visual stimuli in the lateral geniculate nucleus. V1 in red at bottom of image. (1543 image from Andreas Vesalius' Fabrica)

A final population of photosensitive ganglion cells, containing melanopsin, sends information via the retinohypothalamic tract (RHT) to the pretectum (pupillary reflex), to several structures involved in the control of circadian rhythms and sleep such as the suprachiasmatic nucleus (SCN, the biological clock), and to the ventrolateral preoptic nucleus (VLPO, a region involved in sleep regulation).^[10] A recently discovered role for photoreceptive ganglion cells is that they mediate conscious and unconscious vision - acting as rudimentary visual brightness detectors shown in rodless coneless eyes.^[6]

Optic chiasm

The optic nerves from both eyes meet and cross at the optic chiasm,^[11][12] at the base of the hypothalamus of the brain. At this point the information coming from both eyes is combined and then splits according to the visual field. 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.^[9] 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.

Six layers in the LGN

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 humans and other primates starting from catarhinians, including cercopithecidae and apes. Layers 1, 4, and 6 correspond to information from the contralateral (crossed) fibers of the nasal visual field; layers 2, 3, and 5 correspond to information from the ipsilateral (uncrossed) fibers of the temporal visual field. Layer one (1) 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 (4 & 6) 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 (2, 3, & 5) 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. The six layers of the LGN are the area of a credit card, but about three times the thickness of a credit card, rolled up into two ellipsoids 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 (caudal end) in the occipital lobe in and close to the calcarine sulcus.

Gray's FIG. 722? Scheme showing central connections of the optic nerves and optic tracts.

Optic radiation

The optic radiations carries information from the thalamic lateral geniculate nucleus to layer 4 of the visual cortex. The P layer neurons of the LGN relay to V1 layer 4C ?. The M layer neurons relay to V1 layer 4C ?. 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 field of view of the eye, all the way through the optic tract to a nerve position in V1. At this juncture in V1, the image path ceases to be straightforward; there is more cross-connection within the visual cortex.

Visual cortex

Visual cortex: V1, V2, V3, V4, V5 (also called MT)

See also

• Dorsal stream
• Echolocation
• Machine vision
• Memory-prediction framework
• Ventral stream
• Visual perception
• Visual modularity

Notes

References

• shows optic nerve 522 years before Vesalius' engraving.
• .
• .
• .
• .
• .
• . (H.D. Steklis and J. Erwin, editors.) pp. 203-278.
• .
• .
• (References, pp.180-198. Index, pp.199-202. 202 pages.)
• .

External links

• "Webvision: The Organization of the Retina and Visual System" - John Moran Eye Center at University of Utah
• VisionScience.com - An online resource for researchers in vision science.
• Journal of Vision - An online, open access journal of vision science.
• Hagfish research has found the ?missing link? in the evolution of the eye. See: ''Nature Reviews Neuroscience. ''

zh-min-nan:S?-kak h?-thóng ca:Sistema visual de:Visuelles System es:Visión eo:Vida sistemo fr:Système visuel id:Sistem penglihatan he:????? ?????? pam:Visual system no:Syn ru:?????????? ?????????? simple:Visual system sv:Syn zh:????