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retina : ウィキペディア英語版
retina

The retina ( , pl. ''retinae'', ; from Latin ''rēte'', meaning "net") is the third and inner coat of the eye which is a light-sensitive layer of tissue. The optics of the eye create an image of the visual world on the retina (through the cornea and lens), which serves much the same function as the film in a camera. Light striking the retina initiates a cascade of chemical and electrical events that ultimately trigger nerve impulses. These are sent to various visual centres of the brain through the fibres of the optic nerve.
In vertebrate embryonic development, the retina and the optic nerve originate as outgrowths of the developing brain, so the retina is considered part of the central nervous system (CNS) and is actually brain tissue.〔"Sensory Reception: Human Vision: Structure and function of the Human Eye" vol. 27, Encyclopaedia Britannica, 1987〕〔http://www.uphs.upenn.edu/news/News_Releases/jul06/retinput.htm〕 It is the only part of the CNS that can be visualized non-invasively.
The retina is a layered structure with several layers of neurons interconnected by synapses. The only neurons that are directly sensitive to light are the photoreceptor cells. These are mainly of two types: the rods and cones. Rods function mainly in dim light and provide black-and-white vision, while cones support daytime vision and the perception of colour. A third, much rarer type of photoreceptor, the intrinsically photosensitive ganglion cell, is important for reflexive responses to bright daylight.
Neural signals from the rods and cones undergo processing by other neurons of the retina. The output takes the form of action potentials in retinal ganglion cells whose axons form the optic nerve. Several important features of visual perception can be traced to the retinal encoding and processing of light.
==Structure==

The vertebrate retina has ten distinct layers.〔(The Retinal Tunic. ) Virginia-Maryland Regional College of Veterinary Medicine〕 From closest to farthest from the vitreous body - that is, from closest to the front exterior of the head towards the interior and back of the head:
# Inner limiting membrane – basement membrane elaborated by Müller cells
# Nerve fibre layer – axons of the ganglion cell nuclei (note that a thin layer of Müller cell footplates exists between this layer and the inner limiting membrane)
# Ganglion cell layer – contains nuclei of ganglion cells, the axons of which become the optic nerve fibres for messages and some displaced amacrine cells
# Inner plexiform layer – contains the synapse between the bipolar cell axons and the dendrites of the ganglion and amacrine cells.〔
# Inner nuclear layer – contains the nuclei and surrounding cell bodies (perikarya) of the amacrine cells, bipolar cells and horizontal cells.〔
# Outer plexiform layer – projections of rods and cones ending in the rod spherule and cone pedicle, respectively. These make synapses with dendrites of bipolar cells.〔 In the macular region, this is known as the ''Fiber layer of Henle''.
# Outer nuclear layer – cell bodies of rods and cones
# External limiting membrane – layer that separates the inner segment portions of the photoreceptors from their cell nucleus
# Layer of rods and cones – layer of rod cells and cone cells
# Retinal pigment epithelium - single layer of cuboidal cells (with extrusions not shown in diagram). This is closest to the choroid.
These can be simplified into 4 main processing stages: photoreception, transmission to bipolar cells, transmission to ganglion cells which also contain photoreceptors, the photosensitive ganglion cells, and transmission along the optic nerve. At each synaptic stage there are also laterally connecting horizontal and amacrine cells.
The optic nerve is a central tract of many axons of ganglion cells connecting primarily to the lateral geniculate body, a visual relay station in the diencephalon (the rear of the forebrain). It also projects to the superior colliculus, the suprachiasmatic nucleus, and the nucleus of the optic tract. It passes through the other layers creating the optic disc in primates.
Additional structures, not directly associated with vision, are found as outgrowths of the retina in some vertebrate groups. In birds, the pecten is a vascular structure of complex shape that projects from the retina into the vitreous humour; it supplies oxygen and nutrients to the eye, and may also aid in vision. Reptiles have a similar, but much simpler, structure.
In adult humans, the entire retina is approximately 72% of a sphere about 22 mm in diameter. The entire retina contains about 7 million cones and 75 to 150 million rods. The optic disc, a part of the retina sometimes called "the blind spot" because it lacks photoreceptors, is located at the optic papilla, a nasal zone where the optic-nerve fibres leave the eye. It appears as an oval white area of 3mm². Temporal (in the direction of the temples) to this disc is the macula. At its centre is the fovea, a pit that is responsible for our sharp central vision but is actually less sensitive to light because of its lack of rods. Human and non-human primates possess one fovea as opposed to certain bird species such as hawks who actually are bifoviate and dogs and cats who possess no fovea but a central band known as the visual streak. Around the fovea extends the central retina for about 6 mm and then the peripheral retina. The edge of the retina is defined by the ora serrata. The length from one ora to the other (or macula), the most sensitive area along the horizontal meridian is about 32 mm.
In section the retina is no more than 0.5 mm thick. It has three layers of nerve cells and two of synapses, including the unique ribbon synapse. The optic nerve carries the ganglion cell axons to the brain and the blood vessels that open into the retina. The ganglion cells lie innermost in the retina while the photoreceptive cells lie outermost. Because of this counter-intuitive arrangement, light must first pass through and around the ganglion cells and through the thickness of the retina, (including its capillary vessels, not shown) before reaching the rods and cones. However it does not pass through the epithelium or the choroid (both of which are opaque).
The white blood cells in the capillaries in front of the photoreceptors can be perceived as tiny bright moving dots when looking into blue light. This is known as the blue field entoptic phenomenon (or Scheerer's phenomenon).
Between the ganglion cell layer and the rods and cones there are two layers of neuropils where synaptic contacts are made. The neuropil layers are the outer plexiform layer and the inner plexiform layer. In the outer the rods and cones connect to the vertically running bipolar cells, and the horizontally oriented horizontal cells connect to ganglion cells.
The central retina is cone-dominated and the peripheral retina is rod-dominated. In total there are about seven million cones and a hundred million rods. At the centre of the macula is the foveal pit where the cones are smallest and in a hexagonal mosaic, the most efficient and highest density. Below the pit the other retina layers are displaced, before building up along the foveal slope until the rim of the fovea or parafovea which is the thickest portion of the retina. The macula has a yellow pigmentation from screening pigments and is known as the macula lutea. The area directly surrounding the fovea has the highest density of rods converging on single bipolars. Since the cones have a much lesser power of merging signals, the fovea allows for the sharpest vision the eye can attain.〔
Though the rod and cones are a mosaic of sorts, transmission from receptors to bipolars to ganglion cells is not direct. Since there are about 150 million receptors and only 1 million optic nerve fibres, there must be convergence and thus mixing of signals. Moreover, the horizontal action of the horizontal and amacrine cells can allow one area of the retina to control another (e.g. one stimulus inhibiting another). This inhibition is key to the sum of messages sent to the higher regions of the brain. In some lower vertebrates (e.g. the pigeon), there is a "centrifugal" control of messages - that is, one layer can control another, or higher regions of the brain can drive the retinal nerve cells, but in primates this does not occur.〔

抄文引用元・出典: フリー百科事典『 ウィキペディア(Wikipedia)
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