The retina is a thin layer of cells at the back of the Eyeball of vertebrates and some cephalopods; it is the part of the eye which converts Light into nervous signals.
The retina contains Photoreceptor cells (rods and cones) which receive the light; the resulting neural signals then undergo complex processing by other neurons of the retina, and are transformed into action potentials in retinal ganglion cells whose axons form the Optic nerve. The retina not only detects light, it also plays a significant part in Visual perception. In embryonal development, the retina and the optic nerve originate as outgrowths of the Brain.
The unique structure of the blood vessels in the retina have been used for biometric identification.
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In section the retina is no more than 0.5 mm thick. It has three layers of Nerve cells and two of synapses. The optic nerve carries the Ganglion cell axons to the brain and the blood vessels that open into the retina. Perhaps as a product of Evolution, the ganglion cells lie innermost in the retina while the photoreceptive cells lie outermost. Because of this arrangement, light must first pass through the thickness of the retina 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 rod 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 six million cones and a hundred and twenty-five 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 cones respond to bright light and mediate high-resolution vison and colour vision. The rods respond to dim light and mediate lower-resolution, black-and-white, night vision. It is a lack of cones sensitive to red, blue, or green light that causes individuals to have deficiencies in colour vision or various kinds of colour blindness. Humans and old world monkeys have three different types of cones (trichromatic vision) while other mammals lack cones with red sensitive pigment and therefore have poorer (dichromatic) colour vision. When light falls on a receptor it sends a proportional response synaptically to bipolar cells which in turn signal the retinal ganglion cells. The receptors are also 'cross-linked' by horizontal cells and amacrine cells, which modify the synaptic signal before the ganglion cells. Rod and cone signals are intermixed and combine, although rods are mostly active in very poorly lit conditions and saturate in broad daylight, while cones are not sensitive enough to work at very low light levels.
Despite all being nerve cells, only the retinal ganglion cells and few amacrine cells create action potentials. In the photoreceptors, exposure to light hyperpolarizes the membrane in a series of graded shifts. The outer cell segment contains a photopigment and the process leads to a change in levels of cyclic GMP, altering the sodium conductance of the membrane. The amount of neurotransmitter released is reduced in bright light and increases as light levels fall. The actual photopigment is bleached away in bright light and only replaced as a chemical process, so in a transition from bright light to darkness the eye can take up to thirty minutes to reach full sensitivity (see dark adaptation).
In the retinal ganglion cells there are two types of response, depending on the Receptive field of the cell. The receptive fields of retinal ganglion cells comprise a central approximately circular area, where light has one effect on the firing of the cell, and an annular surround, where light has the opposite effect on the firing of the cell. In ON cells, an increment in light intensity in the centre of the receptive field causes the firing rate to increase. In OFF cells, it makes it decrease. Beyond this simple difference ganglion cells are also differentiated by chromatic sensitivity and the type of spatial summation. Cells showing linear spatial summation are termed X cells (also called "parvocellular", "P", or "midget" ganglion cells), and those showing non-linear summation are Y cells (also called "magnocellular, "M", or "parasol" retinal ganglion cells), although the correspondence between X and Y cells (in the cat retina) and P and M cells (in the primate retina) is not as simple as it once seemed.
In the transfer of signal to the brain, the visual pathway, the retina is vertically divided in two, a temporal half and a nasal half. The axons from the nasal half cross the brain at the optic chiasma to join with axons from the temporal half of the other eye before passing into the lateral geniculate body.
Although there are more than 130 million retinal receptors, there are only approximately 1.2 million fibres (axons) in the optic nerve so a large amount of pre-processing is performed within the retina. The fovea produces the most accurate information. Despite occupying about 0.01% of the visual field (less than 2° of Visual angle), about 10% of axons in the optic nerve are devoted to the fovea. The resolution limit of the fovea has been determined at around 104 points. The information capacity is estimated at 5 x 105 bits per second (for more information on bits, see Information theory) without colour or around 6 x 105 bits per second including colour.
Transplantation of retinas has been attempted, but without much success.
At MIT and the University of New South Wales, an "artificial retina" is under development: an implant which will bypass the photoreceptors of the retina and stimulate the attached nerve cells directly, with signals from a digital camera.
The cephalopod retina does not originate as an outgrowth of the brain, as the vertebrate one does. This shows that vertebrate and cephalopod eyes are not homologous but have evolved separately.
| Sensory system - Visual system - Eye - Retina | |
| Rod cell - Cone cell - Bipolar cell - Amacrine cell - Horizontal cell - Ganglion cell |
| Sensory system - Visual system - Eye | |
| Optic disc - Retina - Cornea - Iris - Pupil - Lens - Macula - Sclera - Optic fovea - Blind spot - Vitreous humour - Aqueous humour - Choroid - Ciliary body - Conjunctiva - Angle structure - Tapetum lucidum |