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Retina

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6793: 1869:, located on the blind spot or optic disk. This organ is extremely rich in blood vessels and is thought to supply nutrition and oxygen to the bird retina by diffusion through the vitreous body. The pecten is highly rich in alkaline phosphatase activity and polarized cells in its bridge portion – both befitting its secretory role. Pecten cells are packed with dark melanin granules, which have been theorized to keep this organ warm with the absorption of stray light falling on the pecten. This is considered to enhance metabolic rate of the pecten, thereby exporting more nutritive molecules to meet the stringent energy requirements of the retina during long periods of exposure to light. 2818: 2559:, showing that the therapy was both safe and effective. There were no serious adverse events, and patients in all three studies showed improvement in their visual function as measured by a number of methods. The methods used varied among the three trials, but included both functional methods such as visual acuity and functional mobility as well as objective measures that are less susceptible to bias, such as the pupil's ability to respond to light and improvements on functional MRI. Improvements were sustained over the long-term, with patients continuing to do well after more than 1.5 years. 7635: 1815: 2806: 580: 2830: 6792: 1892:
are obtained from the DRIVE dataset. In addition, the classes of vessels of the DRIVE dataset have also been identified, and an automated method for accurate extraction of these bifurcations is also available. Changes in retinal blood circulation are seen with aging and exposure to air pollution, and may indicate cardiovascular diseases such as hypertension and atherosclerosis. Determining the equivalent width of arterioles and venules near the optic disc is also a widely used technique to identify cardiovascular risks.
924: 932: 1708: 2249: 542: 502:(RPE). Although their photoreceptors contain a protein, retinochrome, that recycles retinal and replicates one of the functions of the vertebrate RPE, cephalopod photoreceptors are likely not maintained as well as in vertebrates, and that as a result, the useful lifetime of photoreceptors in invertebrates is much shorter than in vertebrates. Having easily replaced stalk eyes (some lobsters) or retinae (some spiders, such as 1744: 1668: 513:, but have evolved separately. From an evolutionary perspective, a more complex structure such as the inverted retina can generally come about as a consequence of two alternate processes - an advantageous "good" compromise between competing functional limitations, or as a historical maladaptive relic of the convoluted path of organ evolution and transformation. Vision is an important adaptation in higher vertebrates. 916: 2623:), rAAV-mediated expression of PEDF, angiostatin, and the soluble VEGF receptor sFlt-1, which are all antiangiogenic proteins, have been shown to reduce aberrant vessel formation in animal models. Since specific gene therapies cannot readily be used to treat a significant fraction of patients with retinal dystrophy, there is a major interest in developing a more generally applicable survival factor therapy. 2397:. The neural retina occasionally detaches from the pigment epithelium. In some instances, the cause of such detachment is injury to the eyeball that allows fluid or blood to collect between the neural retina and the pigment epithelium. Detachment is occasionally caused by contracture of fine collagenous fibrils in the vitreous humor, which pull areas of the retina toward the interior of the globe. 2849: 876:, is reached, 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 bipolar cells. Since its cones have a much lesser convergence of signals, the fovea allows for the sharpest vision the eye can attain. 571: 42: 2405:: Night blindness occurs in any person with severe vitamin A deficiency. The reason for this is that without vitamin A, the amounts of retinal and rhodopsin that can be formed are severely depressed. This condition is called night blindness because the amount of light available at night is too little to permit adequate vision in vitamin A–deficient persons. 1976:(CNGs). Thus the cell is hyperpolarised. 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. 534: 688:– single layer of cuboidal epithelial cells (with extrusions not shown in diagram). This layer is closest to the choroid, and provides nourishment and supportive functions to the neural retina, The black pigment melanin in the pigment layer prevents light reflection throughout the globe of the eyeball; this is extremely important for clear vision. 2130:
multiplied by the +1 weight factor. The surrounding photoreceptors are the "nearest neighbors" to the centre and are multiplied by the −1/8 value. The sum of all nine of these elements is finally calculated. This summation is repeated for every photoreceptor in the image by shifting left to the end of a row and then down to the next line.
1941:, while most other mammals lack cones with red sensitive pigment and therefore have poorer dichromatic colour vision. However, some animals have four spectral subtypes, e.g. the trout adds an ultraviolet subgroup to short, medium, and long subtypes that are similar to humans. Some fish are sensitive to the polarization of light as well. 2536:(rAAV) vectors possess a number of features that render them ideally suited for retinal gene therapy, including a lack of pathogenicity, minimal immunogenicity, and the ability to transduce postmitotic cells in a stable and efficient manner. rAAV vectors are increasingly utilized for their ability to mediate efficient transduction of 2224: 562:(purple layer), then to the optic nerve fibres. The signals are processed in these layers. First, the signals start as raw outputs of points in the rod and cone cells. Then, the nerve layers identify simple shapes, such as bright points surrounded by dark points, edges, and movement. (Based on a drawing by 2129:
that implements edge detection. This matrix is the computer equivalent to the centre–surround structure. In this example, each box (element) within this matrix would be connected to one photoreceptor. The photoreceptor in the centre is the current receptor being processed. The centre photoreceptor is
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and for early detection of the onset of disease. The mapping of vascular bifurcations is one of the basic steps in biometric identification. Results of such analyses of retinal blood vessel structure can be evaluated against the ground truth data of vascular bifurcations of retinal fundus images that
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When light strikes 11-cis-retinal (in the disks in the rods and cones), 11-cis-retinal changes to all-trans-retinal which then triggers changes in the opsins. Now, the outer segments do not regenerate the retinal back into the cis- form once it has been changed by light. Instead the retinal is pumped
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Rods, cones, and nerve layers in the retina: The front (anterior) of the eye is on the left. Light (from the left) passes through several transparent nerve layers to reach the rods and cones (far right). Chemical changes in the rods and cones send a signal back to the nerves. The signal goes first to
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have the ability to modulate neuronal growth during development to maintain existing cells and to allow recovery of injured neuronal populations in the eye. AAV encoding neurotrophic factors such as fibroblast growth factor (FGF) family members and GDNF either protected photoreceptors from apoptosis
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algorithms. 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
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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. In ON cells, an increment in light intensity in the centre of the receptive
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separate the subretinal space from the blood supply, thus protecting it from microbes and most immune-mediated damage, and enhancing its potential to respond to vector-mediated therapies. The highly compartmentalized anatomy of the eye facilitates accurate delivery of therapeutic vector suspensions
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and concludes with production of the Muller glia. Although each cell type differentiates from the RPCs in a sequential order, there is considerable overlap in the timing of when individual cell types differentiate. The cues that determine a RPC daughter cell fate are coded by multiple transcription
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antennae, wings, and legs. The optic vesicle gives rise to three structures: the neural retina, the retinal pigmented epithelium, and the optic stalk. The neural retina contains the retinal progenitor cells (RPCs) that give rise to the seven cell types of the retina. Differentiation begins with the
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possess one fovea, as opposed to certain bird species, such as hawks, that are bifoviate, and dogs and cats, that 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 farthest edge of the
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The response of cones to various wavelengths of light is called their spectral sensitivity. In normal human vision, the spectral sensitivity of a cone falls into one of three subtypes, often called blue, green, and red, but more accurately known as short, medium, and long wavelength-sensitive cone
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The energy requirements of the retina are even greater than that of the brain. This is due to the additional energy needed to continuously renew the photoreceptor outer segments, of which 10% are shed daily. Energy demands are greatest during dark adaptation when its sensitivity is most enhanced.
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1:posterior segment 2:ora serrata 3:ciliary muscle 4:ciliary zonules 5:Schlemm's canal 6:pupil 7:anterior chamber 8:cornea 9:iris 10:lens cortex 11:lens nucleus 12:ciliary process 13:conjunctiva 14:inferior oblique muscule 15:inferior rectus muscule 16:medial rectus muscle 17:retinal arteries and
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algorithms used by computer programmers to extract or enhance the edges in a digital photograph. Thus, the retina performs operations on the image-representing impulses to enhance the edges of objects within its visual field. For example, in a picture of a dog, a cat and a car, it is the edges of
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Although the inverted retina of vertebrates appears counter-intuitive, it is necessary for the proper functioning of the retina. The photoreceptor layer must be embedded in the retinal pigment epithelium (RPE), which performs at least seven vital functions, one of the most obvious being to supply
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These centre–surround structures are not physical apparent, in the sense that one cannot see them by staining samples of tissue and examining the retina's anatomy. The centre–surround structures are logical (i.e., mathematically abstract) in the sense that they depend on the connection strengths
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A recent study on the evolutionary purpose for the inverted retina structure from the APS (American Physical Society) says that "The directional of glial cells helps increase the clarity of human vision. But we also noticed something rather curious: the colours that best passed through the glial
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The central retina predominantly contains cones, while the peripheral retina predominantly contains rods. In total, the retina has about seven million cones and a hundred million rods. At the centre of the macula is the foveal pit where the cones are narrow and long, and arranged in a hexagonal
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A third view of the "inverted" vertebrate eye is that it combines two benefits - the maintenance of the photoreceptors mentioned above, and the reduction in light intensity necessary to avoid blinding the photoreceptors, which are based on the extremely sensitive eyes of the ancestors of modern
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in the sense that the light-sensing cells are in the back of the retina, so that light has to pass through layers of neurons and capillaries before it reaches the photosensitive sections of the rods and cones. The ganglion cells, whose axons form the optic nerve, are at the front of the retina;
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to the brain, and the blood vessels that supply the retina. The ganglion cells lie innermost in the eye while the photoreceptive cells lie beyond. Because of this counter-intuitive arrangement, light must first pass through and around the ganglion cells and through the thickness of the retina,
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Further computer simulations showed that green and red are concentrated five to ten times more by the glial cells, and into their respective cones, than blue light. Instead, excess blue light gets scattered to the surrounding rods. This optimization is such that color vision during the day is
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expression in the retinal pigmented epithelium (RPE), photoreceptors, or ganglion cells for long periods of time after a single treatment. In addition, the eye and the visual system can be routinely and easily monitored for visual function and retinal structural changes after injections with
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The total sum of this matrix is zero, if all the inputs from the nine photoreceptors are of the same value. The zero result indicates the image was uniform (non-changing) within this small patch. Negative or positive sums mean the image was varying (changing) within this small patch of nine
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In adult humans, the entire retina is about 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
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The bird retina is devoid of blood vessels, perhaps to give unobscured passage of light for forming images, thus giving better resolution. It is, therefore, a considered view that the bird retina depends for nutrition and oxygen supply on a specialized organ, called the "pecten" or
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out to the surrounding RPE where it is regenerated and transported back into the outer segments of the photoreceptors. This recycling function of the RPE protects the photoreceptors against photo-oxidative damage and allows the photoreceptor cells to have decades-long useful lives.
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Bartsch, U.; Oriyakhel, W.; Kenna, P. F.; Linke, S.; Richard, G.; Petrowitz, B.; Humphries, P.; Farrar, G. J.; Ader, M. (2008). "Retinal cells integrate into the outer nuclear layer and differentiate into mature photoreceptors after subretinal transplantation into adult mice".
2696:, the theory that sight occurs from rays emitted by the eyes. However, Ibn Al-Haytham decided that the retina could not be responsible for the beginnings of vision because the image formed on it was inverted. Instead he decided it must begin at the surface of the lens. 2118:
these objects that contain the most information. In order for higher functions in the brain (or in a computer for that matter) to extract and classify objects such as a dog and a cat, the retina is the first step to separating out the various objects within the scene.
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There are two types of centre–surround structures in the retina – on-centres and off-centres. On-centres have a positively weighted centre and a negatively weighted surround. Off-centres are just the opposite. Positive weighting is more commonly known as
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could be transplanted successfully in the mouse retina if donor cells were at a critical developmental stage. Recently Ader and colleagues in Dublin showed, using the electron microscope, that transplanted photoreceptors formed synaptic connections.
2047:), about 10% of axons in the optic nerve are devoted to the fovea. The resolution limit of the fovea has been determined to be around 10,000 points. The information capacity is estimated at 500,000 bits per second (for more information on bits, see 2462:
is a non-invasive method to examine the small arteries and veins in the retina which allows to draw conclusions about the morphology and the function of small vessels elsewhere in the human body. It has been established as a predictor of
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Chapman, N.; Dell'omo, G.; Sartini, M. S.; Witt, N.; Hughes, A.; Thom, S.; Pedrinelli, R. (1 August 2002). "Peripheral vascular disease is associated with abnormal arteriolar diameter relationships at bifurcations in the human retina".
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Wong TY, Knudtson MD, Klein R, Klein BE, Meuer SM, Hubbard LD (2004). "Computer assisted measurement of retinal vessel diameters in the Beaver Dam Eye Study: methodology, correlation between eyes, and effect of refractive errors".
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The above example is called "balanced". The term balanced means that the sum of the negative weights is equal to the sum of the positive weights so that they cancel out perfectly. Retinal ganglion cells are almost never perfectly
1921:). The rod responses are saturated at daylight levels and do not contribute to pattern vision. However, rods do respond to dim light and mediate lower-resolution, monochromatic vision under very low levels of illumination (called 2073:
When the retina sends neural impulses representing an image to the brain, it spatially encodes (compresses) those impulses to fit the limited capacity of the optic nerve. Compression is necessary because there are 100 times more
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has been used to image individual rods and cones in the living human retina, and a company based in Scotland has engineered technology that allows physicians to observe the complete retina without any discomfort to patients.
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can allow one area of the retina to control another (e.g. one stimulus inhibiting another). This inhibition is key to lessening the sum of messages sent to the higher regions of the brain. In some lower vertebrates (e.g. the
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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.
1999:, which modify the synaptic signal before it reaches the ganglion cells, the neural signals being intermixed and combined. Of the retina's nerve cells, only the retinal ganglion cells and few amacrine cells create 6829:
veins 18:optic disc 19:dura mater 20:central retinal artery 21:central retinal vein 22:optic nerve 23:vorticose vein 24:bulbar sheath 25:macula 26:fovea 27:sclera 28:choroid 29:superior rectus muscle 30:retina
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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
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bifurcates and supplies the retina via two distinct vascular networks: the choroidal network, which supplies the choroid and the outer retina, and the retinal network, which supplies the retina's inner layer.
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worked out the optics of the eye and decided that the retina must be where sight begins. He left it up to other scientists to reconcile the inverted retinal image with our perception of the world as upright.
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In addition, the retina has been described as a "window" into the brain and body, given that abnormalities detected through an examination of the retina can discover both neurological and systemic diseases.
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The retina is stratified into distinct layers, each containing specific cell types or cellular compartments that have metabolisms with different nutritional requirements. To satisfy these requirements, the
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In addition to guiding cell fate determination, cues exist in the retina to determine the dorsal-ventral (D-V) and nasal-temporal (N-T) axes. The D-V axis is established by a ventral to dorsal gradient of
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Illustration of the distribution of cone cells in the fovea of an individual with normal colour vision (left), and a colourblind (protanopic) retina. The center of the fovea holds very few blue-sensitive
1929:. At mesopic light levels, both the rods and cones are actively contributing pattern information. What contribution the rod information makes to pattern vision under these circumstances is unclear. 2645:, 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. 2900: 2204:
The retina performs all these calculations in parallel while the computer operates on each pixel one at a time. The retina performs no repeated summations and shifting as would a computer.
1937:. These individuals are not blind to objects of a particular colour, but are unable to distinguish between colours that can be distinguished by people with normal vision. Humans have this 900:), control of messages is "centrifugal" – 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. 380:. Rods function mainly in dim light and provide monochromatic vision. Cones function in well-lit conditions and are responsible for the perception of colour through the use of a range of 887:
is not direct. Since about 150 million receptors and only 1 million optic nerve fibres exist, convergence and thus mixing of signals must occur. Moreover, the horizontal action of the
2829: 2237:). The exact function of the LGN is unknown at this time. The output of the LGN is then sent to the back of the brain. Specifically, the output of the LGN "radiates" out to the V1 5100:
Patton, N.; Aslam, T.; MacGillivray, T.; Deary, I.; Dhillon, B.; Eikelboom, R.; Yogesan, K.; Constable, I. (2006). "Retinal image analysis: Concepts, applications and potential".
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proteins. The role of Pax6 in eye development was elegantly demonstrated by Walter Gehring and colleagues, who showed that ectopic expression of Pax6 can lead to eye formation on
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holds promise as a potential avenue to cure a wide range of retinal diseases. This involves using a non-infectious virus to shuttle a gene into a part of the retina. Recombinant
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The bifurcations and other physical characteristics of the inner retinal vascular network are known to vary among individuals, and these individual variances have been used for
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game where players trace neurons in the retina. The goals of the EyeWire project are to identify specific cell types within the known broad classes of retinal cells, and to
653:– projections of rods and cones ending in the rod spherule and cone pedicle, respectively, these make synapses with dendrites of bipolar cells and horizontal cells. In the 5743:"Treatment of Leber Congenital Amaurosis Due to RPE65Mutations by Ocular Subretinal Injection of Adeno-Associated Virus Gene Vector: Short-Term Results of a Phase I Trial" 2571:
to specific tissues under direct visualization using microsurgical techniques. In the sheltered environment of the retina, AAV vectors are able to maintain high levels of
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The retina translates an optical image into neural impulses starting with the patterned excitation of the colour-sensitive pigments of its rods and cones, the retina's
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The cephalopod retina does not originate as an outgrowth of the brain, as the vertebrate one does. This difference suggests that vertebrate and cephalopod eyes are not
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Once the image is spatially encoded by the centre–surround structures, the signal is sent out along the optic nerve (via the axons of the ganglion cells) through the
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Light striking the retina initiates a cascade of chemical and electrical events that ultimately trigger nerve impulses that are sent to various visual centres of the
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den Hollander, Anneke I.; Roepman, Ronald; Koenekoop, Robert K.; Cremers, Frans P.M. (2008). "Leber congenital amaurosis: Genes, proteins and disease mechanisms".
4923:"Air Pollution and the microvasculature: a crosssectional assessment of in vivo retinal images in the population based multiethnic study of atherosclerosis (MESA)" 1232: 6812: 1437: 255: 3345:
Drazen, J. C.; Yeh, J.; Friedman, J.; Condon, N. (June 2011). "Metabolism and enzyme activities of hagfish from shallow and deep water of the Pacific Ocean".
6205: 3431: 770:, where the optic-nerve fibres leave the eye. It appears as an oval white area of 3 mm. Temporal (in the direction of the temples) to this disc is the 6809: 6144: 2817: 7072: 2491: 1564: 385: 6810: 6174: 1283: 1008: 6815: 5425:
Dinculescu Astra; Glushakova Lyudmyla; Seok-Hong Min; Hauswirth William W (2005). "Adeno-associated virus-vectored gene therapy for retinal disease".
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cells were green to red, which the eye needs most for daytime vision. The eye usually receives too much blue—and thus has fewer blue-sensitive cones.
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retina, the photoreceptors are in front, with processing neurons and capillaries behind them. Because of this, cephalopods do not have a blind spot.
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Yu, DY; Yu, PK; Cringle, SJ; Kang, MH; Su, EN (May 2014). "Functional and morphological characteristics of the retinal and choroidal vasculature".
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Bringmann A, Syrbe S, Görner K, Kacza J, Francke M, Wiedemann P, Reichenbach A (2018). "The primate fovea: Structure, function and development".
1972:. This in turn causes the Ga-subunit of the protein to activate a phosphodiesterase (PDE6), which degrades cGMP, resulting in the closing of Na+ 491:, is avascular (does not have blood vessels), and has minimal neural tissue in front of the photoreceptors, thereby minimizing light scattering. 6811: 2442:
measure the retina's electrical activity, which is affected by certain diseases. A relatively new technology, now becoming widely available, is
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Sabra, A. I. (Ed.). (1011–1021/1989). The optics of Ibn Al-Haytham: Books I-III: On direct vision (A. I. Sabra, Trans.). The Warburg Institute.
778:, a pit that is responsible for sharp central vision, but is actually less sensitive to light because of its lack of rods. Human and non-human 6796: 4829:
Qureshi, T. A.; Habib, M.; Hunter, A.; Al-Diri, B. (June 2013). "A manually-labeled, artery/Vein classified benchmark for the DRIVE dataset".
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field causes the firing rate to increase. In OFF cells, it makes it decrease. In a linear model, this response profile is well described by a
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two-dimensional image of the visual world on the retina, which then processes that image within the retina and sends nerve impulses along the
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subtypes. It is a lack of one or more of the cone subtypes that causes individuals to have deficiencies in colour vision or various kinds of
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Halder, G; Callaerts, P; Gehring, WJ (24 March 1995). "Induction of ectopic eyes by targeted expression of the eyeless gene in Drosophila".
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This strategy is effective against a number of retinal diseases that have been studied, including neovascular diseases that are features of
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from the retina, this is the space created between the posterior cortical vitreous face and the internal limiting membrane of the retina.
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does occur. Some vertebrates, including humans, have an area of the central retina adapted for high-acuity vision. This area, termed the
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therefore, the optic nerve must cross through the retina en route to the brain. No photoreceptors are in this region, giving rise to the
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published numerous experiments demonstrating that sight occurs from light reflecting from objects into the eye. This is consistent with
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Letelier J, Bovolenta P, Martínez-Morales JR (2017). "The pigmented epithelium, a bright partner against photoreceptor degeneration".
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In the photoreceptors, exposure to light hyperpolarizes the membrane in a series of graded shifts. The outer cell segment contains a
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between bipolar and ganglion cells. It is believed that the connection strength between cells is caused by the number and types of
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Additional structures, not directly associated with vision, are found as outgrowths of the retina in some vertebrate groups. In
6800: 682:– inner segments and outer segments of rods and cones, the outer segments contain a highly specialized light-sensing apparatus. 6819: 6818: 5465:
Cideciyan A. V.; Hauswirth W. W.; Aleman T. S.; Kaushal S.; Schwartz S. B.; Boye S. L.; Windsor E. A. M.; et al. (2009).
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Qureshi, T. A.; Hunter, A.; Al-Diri, B. (June 2014). "A Bayesian Framework for the Local Configuration of Retinal Junctions".
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in front of the photoreceptors can be perceived as tiny bright moving dots when looking into blue light. This is known as the
6672: 6598: 6461: 5220: 4897: 4846: 4746: 4337: 3760: 3721: 3696: 2939: 2779: 2734: 2638: 6799: 4796: 3848:(2014). "Proposed Lexicon for Anatomic Landmarks in Normal Posterior Segment Spectral-Domain Optical Coherence Tomography". 2374: 6798: 5674:
Bainbridge J. W. B.; Smith A. J.; Barker S. S.; Robbie S.; Henderson R.; Balaggan K.; Viswanathan A.; et al. (2008).
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Franze K, Grosche J, Skatchkov SN, Schinkinger S, Foja C, Schild D, Uckermann O, Travis K, Reichenbach A, Guck J (2007).
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describes a group of diseases characterized by loss of central vision because of death or impairment of the cells in the
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showing the blood vessels in a normal human retina. Veins are darker and slightly wider than corresponding arteries. The
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Ashtari M.; Cyckowski L. L.; Monroe J. F.; Marshall K. A.; Chung D. C.; Auricchio A.; Simonelli F.; et al. (2011).
5467:"Human RPE65 gene therapy for Leber congenital amaurosis: persistence of early visual improvements and safety at 1 year" 4769: 1754: 368:
and is supported by an outer layer of pigmented epithelial cells. The primary light-sensing cells in the retina are the
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Hauswirth W. W.; Aleman T. S.; Kaushal S.; Cideciyan A. V.; Schwartz S. B.; Wang L.; Conlon T. J.; et al. (2008).
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Cepko, Connie (September 2014). "Intrinsically different retinal progenitor cells produce specific types of progeny".
3075: 2086:", which is carried out by the "centre–surround structures", which are implemented by the bipolar and ganglion cells. 1917:
The cones respond to bright light and mediate high-resolution colour vision during daylight illumination (also called
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and seems to have, according to a study published in 2019, potential in the early detection of Alzheimer's disease.
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Simonelli F.; Maguire A. M.; Testa F.; Pierce E. A.; Mingozzi F.; Bennicelli J. L.; Rossi S.; et al. (2010).
5368:"Functional and morphological changes of the retinal vessels in Alzheimer's disease and mild cognitive impairment" 3428: 2418:
A number of different instruments are available for the diagnosis of diseases and disorders affecting the retina.
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Maguire A. M.; Simonelli F.; Pierce E. A.; Pugh E. N.; Mingozzi F.; Bennicelli J.; Banfi S.; et al. (2008).
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Sherman, J (June 2009). "Photoreceptor integrity line joins the nerve fiber layer as key to clinical diagnosis".
2967:"Sensory Reception: Human Vision: Structure and function of the Human Eye" vol. 27, Encyclopædia Britannica, 1987 2556: 5625:
Maguire A. M.; High K. A.; Auricchio A.; Wright J. F.; Pierce E. A.; Testa F.; Mingozzi F.; et al. (2009).
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is a group of genetic diseases that affect the retina and cause the loss of night vision and peripheral vision.
1949: 834: 4281:"Single-cell transcriptional logic of cell-fate specification and axon guidance in early-born retinal neurons" 2607:. Since the regulation of vascularization in the mature retina involves a balance between endogenous positive 2259:
There are many inherited and acquired diseases or disorders that may affect the retina. Some of them include:
6595:, edited by William Krause, Dept. Pathology and Anatomical science, University of Missouri School of Medicine 3319: 2642: 2386: 1020: 430:, the retina and the optic nerve originate as outgrowths of the developing brain, specifically the embryonic 411:. Neural signals from the rods and cones undergo processing by other neurons, whose output takes the form of 250: 6821: 6502:"The Retinome – Defining a reference transcriptome of the adult mammalian retina/retinal pigment epithelium" 3952:
Turgut, Burak; University, Fırat; Medicine, School of; Ophthalmology, Department of; Elazig; Turkey (2017).
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Zone at the outer choroid with a marked change in texture, in which large circular or ovoid profiles abut a
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Azzopardi G.; Petkov N. (2011). "Detection of Retinal Vascular Bifurcations by Trainable V4-Like Filters".
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produces the most accurate information. Despite occupying about 0.01% of the visual field (less than 2° of
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The unique architecture of the retina and its relatively immune-privileged environment help this process.
814:(including its capillary vessels, not shown) before reaching the rods and cones. Light is absorbed by the 613:– contains nuclei of ganglion cells, the axons of which become the optic nerve fibres, and some displaced 7216: 7185: 7177: 6857: 2370: 2174:
The above matrix is only an approximation to what really happens inside the retina. The differences are:
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bodies (a thin layer of Müller cell footplates exists between this layer and the inner limiting membrane)
389: 6357: 6166: 6091: 384:, as well as high-acuity vision used for tasks such as reading. A third type of light-sensing cell, the 7955: 7915: 7367: 7012: 7002: 6823: 6331: 4550:"Diagrammatic representation of disc shedding and phagosome retrieval into the pigment epithelial cell" 3954:"Past and Present Terminology for the Retinal and Choroidal Structures in Optical Coherence Tomography" 2741: 2604: 2537: 2306: 2234: 1765: 1510: 1499: 1481: 1471: 1443: 1276: 1043: 815: 685: 673: 583:
Distribution of rods and cones along a line passing through the fovea and the blind spot of a human eye
499: 6804: 4974:"Retinal Microvascular Responses to Short-Term Changes in Particulate Air Pollution in Healthy Adults" 4725: 2835:
Illustration of image as 'seen' by the retina independent of optic nerve and striate cortex processing
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The choroid supplies about 75% of these nutrients to the retina and the retinal vasculature only 25%.
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Thick layer of round or ovalshaped hyperreflective profiles, with hyporeflective cores in mid-choroid
7865: 7564: 7394: 6665: 6614: 4461:"Watching the human retina breath in real time and the slowing of mitochondrial respiration with age" 2759: 2194: 6822: 6423:
Rodieck RW (1965). "Quantitative analysis of cat retinal ganglion cell response to visual stimuli".
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Hatakeyama, J; Kageyama, R (February 2004). "Retinal cell fate determination and bHLH factors".
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Thick layer of oval-shaped hyperreflective profiles, with hyporeflective cores in outer choroid
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Hildebrand, Göran Darius; Fielder, Alistair R. (2011). "Anatomy and Physiology of the Retina".
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The vertebrate retina has 10 distinct layers. From closest to farthest from the vitreous body:
551: 435: 397: 262: 238: 226: 190: 6801: 6628: 676:– layer that separates the inner segment portions of the photoreceptors from their cell nuclei 7323: 7128: 7052: 6990: 6978: 6933: 5321: 4655: 2910: 2745: 2567: 2548:. Each cell type can be specifically targeted by choosing the appropriate combination of AAV 2501: 2238: 1988: 1952:(cGMP) keep the Na+ channel open, and thus in the resting state the cell is depolarised. The 1719: 1626: 1321: 1196: 1124: 912:(OCT), 18 layers can be identified in the retina. The layers and anatomical correlation are: 884: 853: 849: 841: 807: 697: 692:
These layers can be grouped into four main processing stages—photoreception; transmission to
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Louwies, Tijs; Panis, Luc Int; Kicinski, Michal; Boever, Patrick De; Nawrot, Tim S. (2013).
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Lo Giudice, Quentin; Leleu, Marion; La Manno, Gioele; Fabre, Pierre J. (1 September 2019).
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auto-fluorescence (FAF), dark-adapted visual thresholds, vascular diameters, pupillometry,
2577: 2345: 2310: 2269: 2263: 2122: 892: 709: 614: 555: 41: 3383: 2282:(CORD) describes a number of diseases where vision loss is caused by deterioration of the 1646:, whereas the N-T axis is coordinated by expression of the forkhead transcription factors 438:(CNS) and is actually brain tissue. It is the only part of the CNS that can be visualized 8: 7614: 7474: 7469: 7434: 7338: 7044: 6995: 6983: 6973: 6928: 6862: 6776: 6723: 6107: 3681: 2767: 2689: 2585: 2541: 2435: 1938: 1925:). The illumination in most office settings falls between these two levels and is called 1514: 1258: 1210: 1167: 1156: 1131: 1096: 857: 728: 693: 667: 641: 633: 623: 610: 547: 465: 7624: 6650: 6568: 6412: 6247: 5792:"The human visual cortex responds to gene therapy-mediated recovery of retinal function" 5385: 4501: 4476: 4460: 4160: 3399: 3151: 1814: 787:. The distance from one ora to the other (or macula), the most sensitive area along the 7634: 7504: 7479: 7459: 7308: 7088: 7084: 7080: 7028: 6867: 6751: 6718: 6608: 6277: 6058: 6025: 5954: 5911: 5816: 5791: 5767: 5742: 5651: 5626: 5599: 5572: 5545: 5520: 5491: 5466: 5402: 5372: 5367: 5343: 5316: 5270: 5245: 5125: 4998: 4973: 4949: 4922: 4903: 4852: 4802: 4701: 4676: 4549: 4386: 4223: 4124: 4099: 4073: 3661: 3613: 3588: 3547: 3520: 3496: 3471: 3296: 3263: 3244: 3170: 3135: 3048: 3013: 2423: 2393: 2381: 2302: 2294: 2279: 2244:
Simplified signal flow: Photoreceptors → Bipolar → Ganglion → Chiasm → LGN → V1 cortex
2075: 2048: 1911: 1547: 1415: 1409: 1398: 1367: 1314: 1265: 1238: 1206: 510: 495: 369: 7655: 6624:
Jeremy Nathans's Seminars: "The Vertebrate Retina: Structure, Function, and Evolution"
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Tso, Mark O.M.; Jampol, Lee M. (1982). "Pathophysiology of Hypertensive Retinopathy".
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Proceedings of the 26th IEEE International Symposium on Computer-Based Medical Systems
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Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology
7802: 7709: 7529: 7464: 7348: 7064: 6968: 6885: 6756: 6682: 6537: 6488: 6457: 6440: 6436: 6313: 6269: 6111: 6063: 6045: 6006: 5998: 5946: 5942: 5903: 5862: 5821: 5772: 5716: 5656: 5604: 5550: 5496: 5442: 5407: 5348: 5275: 5226: 5216: 5194: 5184: 5161: 5117: 5113: 5082: 5074: 5038: 5003: 4954: 4893: 4842: 4742: 4706: 4639: 4619: 4596: 4592: 4506: 4488: 4378: 4374: 4343: 4333: 4310: 4302: 4258: 4215: 4207: 4172: 4129: 4039: 3932: 3898: 3888: 3865: 3845: 3793: 3717: 3692: 3653: 3618: 3552: 3521:"Photoreceptor signaling: supporting vision across a wide range of light intensities" 3501: 3487: 3411: 3364: 3301: 3283: 3236: 3232: 3175: 3083: 3053: 3035: 2935: 2366: 2337: 1835: 1818: 1073: 600: 346: 342: 5958: 5740: 5464: 4907: 4576: 4390: 4227: 2182:
The table is square while the centre–surround structures in the retina are circular.
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Retinal development begins with the establishment of the eye fields mediated by the
856:. In the outer neuropil layer, the rods and cones connect to the vertically running 498:
to the eyes of many vertebrates. Squid eyes do not have an analog of the vertebrate
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Several clinical trials have already reported positive results using rAAV to treat
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to join with axons from the temporal half of the other eye before passing into the
2000: 1934: 1901: 1529: 1214: 826: 579: 484: 412: 393: 365: 314: 310: 145: 111: 105: 72: 6645: 6484: 3882: 3649: 1243:(obliquely orientated fibres; not present in mid-peripheral or peripheral retina) 7724: 7604: 7599: 7584: 7499: 7419: 7246: 7205: 7201: 7123: 5977:"Recombinant AAV-mediated gene transfer to the retina: gene therapy perspectives" 5898: 5881: 5299: 5157: 4939: 4807: 4738: 4254: 3861: 3779: 3435: 2854: 2787: 2775: 2755: 2700: 2670: 2427: 2401: 2208: 2201:. (The computer pixel is basically the equivalent of a biological photoreceptor.) 2060: 2040: 2007: 1992: 1922: 1918: 794:
In section, the retina is no more than 0.5 mm thick. It has three layers of
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Although the overlying neural tissue is partly transparent, and the accompanying
446:. The retina is the part of the body with the greatest continuous energy demand. 442:. Like most of the brain, the retina is isolated from the vascular system by the 219: 66: 6471:
Wässle H, Boycott BB (1991). "Functional architecture of the mammalian retina".
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In the retinal ganglion cells there are two types of response, depending on the
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oxygen and other necessary nutrients needed for the photoreceptors to function.
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Time-Domain OCT of the macular area of a retina at 800 nm, axial resolution 3 μm
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is a vascular structure of complex shape that projects from the retina into the
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Kaynezhad, Pardis; Tachtsidis, Ilias; Sivaprasad, Sobha; Jeffery, Glen (2023).
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Receptive field, for figures and more information on centre–surround structures
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is a central tract of many axons of ganglion cells connecting primarily to the
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Adar SD, Klein R, Klein BE, Szpiro AA, Cotch MF, Wong TY, et al. (2010).
4838: 4770:"Retinal fundus images – Ground truth of vascular bifurcations and crossovers" 3030: 2794:
between neurons in the retina, which will help to determine how vision works.
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can cause damage to the tiny blood vessels that supply the retina, leading to
2309:. There are several modern treatment methods for fixing a retinal detachment: 1177: 1050: 923: 345:. The retina serves a function which is in many ways analogous to that of the 7944: 7882: 7853: 7494: 7439: 7276: 7211: 7048: 6509: 6049: 6002: 5841:"Immune response following intraocular delivery of recombinant viral vectors" 5518: 5198: 5078: 4492: 4347: 4306: 4211: 3902: 3287: 3039: 2885: 2791: 2730: 2219:
Here is an example of an input image and how edge detection would modify it.
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proteins, with subsequent development of the optic vesicles regulated by the
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From innermost to outermost, the layers identifiable by OCT are as follows:
7897: 7829: 7519: 7489: 7359: 7236: 7060: 6895: 6877: 6555: 6541: 6522: 6317: 6273: 6067: 6010: 5950: 5907: 5866: 5825: 5776: 5720: 5676:"Effect of gene therapy on visual function in Leber's congenital amaurosis" 5660: 5608: 5554: 5500: 5446: 5411: 5352: 5279: 5165: 5121: 5086: 5007: 4958: 4524: 4510: 4382: 4314: 4262: 4219: 4133: 4043: 3869: 3797: 3657: 3622: 3556: 3505: 3415: 3368: 3240: 3179: 3087: 3057: 2890: 2616: 2581: 2529: 2487: 2333: 2298: 2230: 2044: 1984: 1945: 1888: 1878: 1866: 1826: 1371: 1336: 1331: 1053: 954: 758:; it supplies oxygen and nutrients to the eye, and may also aid in vision. 751: 724: 431: 350: 173: 6492: 6444: 6115: 5993: 5976: 5857: 5840: 5704: 5590: 5042: 4889: 4710: 4643: 4600: 4176: 2385:
is a white appearance of the retina, and can occur by lipid deposition in
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play a significant role in this process, but that is not represented here.
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When thus excited by light, the photoceptor sends a proportional response
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identified the retina from dissections of cadaver eyes. He called it the
2641:, The University of Southern California, RWTH Aachen University, and the 2483:
The following are commonly modalities of management for retinal disease:
2318: 2186: 2103: 1687: 1086: 830: 784: 716: 595: 408: 334: 6255: 5712: 5536: 5215:(4th ed.). Philadelphia: Elsevier/Mosby. 2006. pp. 2013–2015. 4989: 3711: 1707: 7812: 7662: 7579: 7524: 7409: 7142: 6900: 5573:"Safety and efficacy of gene transfer for Leber's congenital amaurosis" 4733:. Lecture Notes in Computer Science. Vol. 6854. pp. 451–459. 4452: 4297: 4280: 3788: 3759:
Cuenca, Nicolás; Ortuño-Lizarán, Isabel; Pinilla, Isabel (March 2018).
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List of xanthoma variants associated with hyperlipoproteinemia subtypes
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which bridge the inner and outer segments of the photoreceptor cells.
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is used to describe a separation of the neurosensory retina from the
2283: 2126: 2051:) without colour or around 600,000 bits per second including colour. 1969: 1961: 1326: 873: 636:– contains the nuclei and surrounding cell bodies (perikarya) of the 377: 47: 6227:
MacLaren, RE; Pearson, RA; MacNeil, A; et al. (November 2006).
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The centre–surround structures are mathematically equivalent to the
1872: 903: 7846: 7819: 7782: 7714: 7414: 7036: 2549: 2451: 2287: 2223: 1635: 1202: 1171: 1135: 845: 704:; and transmission along the optic nerve. At each synaptic stage, 494:
The cephalopods have a non-inverted retina, which is comparable in
373: 326: 4677:"On connecting large vessels to small – the meaning of murray law" 3193:
Baker, Oliver (23 April 2010). "Focus: Eye Cells as Light Pipes".
2065: 7735: 7288: 7189: 7133: 6849: 6604: 5879: 3843: 3635: 3136:"Muller cells are living optical fibers in the vertebrate retina" 3133: 2783: 2711:
published the first major characterization of retinal neurons in
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Spectral domain optical coherence tomography in macular diseases
915: 848:, where synaptic contacts are made. The neuropil layers are the 7649: 7193: 6733: 6705: 6229:"Retinal repair by transplantation of photoreceptor precursors" 4872:
2014 IEEE Conference on Computer Vision and Pattern Recognition
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Meyer, Carsten H.; Saxena, Sandeep; Sadda, Srinivas R. (2017).
2355: 2273: 1953: 897: 771: 654: 361: 354: 185: 139: 5055: 3716:. Philadelphia, PA: Holt-Saunders International. p. 465. 3472:"Molecular basis for photoreceptor outer segment architecture" 2475:
Treatment depends upon the nature of the disease or disorder.
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channels to transport photons directly to the photoreceptors,
126: 7792: 6923: 4146: 3568: 3566: 2978:"Penn Researchers Calculate How Much the Eye Tells the Brain" 2823:
Another view of the eye and the structures of the eye labeled
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Boston, Marco A. Bonini Filho, MD, and Andre J. Witkin, MD.
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of retinas has been attempted, but without much success. At
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cross-sectional view; eyes vary significantly among animals.
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Webvision: The neural organization of the vertebrate retina
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Staurenghi, Giovanni; Sadda, Srinivas; Chakravarthy, Usha;
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Adeno associated virus and gene therapy of the human retina
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Adeno associated virus and gene therapy of the human retina
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noninvasive advanced technology, such as visual acuities,
2446:(OCT). This non-invasive technique allows one to obtain a 533: 6680: 6417:
Histologie du Système Nerveux de l'Homme et des Vertébrés
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Virginia–Maryland Regional College of Veterinary Medicine
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Modification of systemic risk factors for retinal disease
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of human retinas is 8.75 megabits per second, whereas a
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enhanced, while night-time vision suffers very little".
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Sherman, Jerome; Epshtein, Daniel (15 September 2012).
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Krause's Essential Human Histology for Medical Students
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of sorts, transmission from receptors, to bipolars, to
6636: – Histology Learning System at Boston University 6383: 6226: 5882:"Versatility of AAV vectors for retinal gene transfer" 5142: 3589:"The retinal pigment epithelium in health and disease" 3469: 454: 6599:
Eye, Brain, and Vision – online book – by David Hubel
4797:"DRIVE: Digital Retinal Images for Vessel Extraction" 4723: 4574: 4240: 3909: 2752:'s retinal transfer rate is 875 kilobits per second. 2426:
have long been used to examine the retina. Recently,
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may contain excessive or inappropriate references to
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Thin layer of moderate reflectivity in inner choroid
727:(the rear of the forebrain). It also projects to the 148: 90: 6500:
Schulz HL, Goetz T, Kaschkoetoe J, Weber BH (2004).
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Clinical anatomy and physiology of the visual system
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came to refer to a layer around the brain; the term
2297:, the retina detaches from the back of the eyeball. 2023:
In the transfer of visual signals to the brain, the
123: 117: 84: 78: 4059:"Outer Retinal Layers as Predictors of Vision Loss" 3986:"Outer Retinal Layers as Predictors of Vision Loss" 3712:Romer, Alfred Sherwood; Parsons, Thomas S. (1977). 3012:Viegas, Filipe O.; Neuhauss, Stephan C. F. (2021). 1513:mitochondria zone + Junction between the RPE & 739:. It passes through the other layers, creating the 517:hagfish (fish that live in very deep, dark water). 114: 75: 6603:Kolb, H., Fernandez, E., & Nelson, R. (2003). 5179:Chen, Janglin; Cranton, Wayne; Fihn, Mark (2016). 2669:, from its resemblance to a casting net. The term 6451: 5514: 5512: 5510: 5460: 5458: 5456: 3918: 3747: 3586: 3384:"Retinal Glial Cells Enhance Human Vision Acuity" 3007: 3005: 3003: 2665:layer, from its resemblance to a spider web, and 1873:Biometric identification and diagnosis of disease 904:Layers imagable with optical coherence tomography 7942: 3923:. Springer, Berlin, Heidelberg. pp. 39–65. 3678: 2513: 1968:. This causes the receptor to activate multiple 360:The neural retina consists of several layers of 6795: 6607:. Salt Lake City, Utah: John Moran Eye Center, 5667: 5620: 5618: 5566: 5564: 5315:Seidelmann, SB; et al. (1 November 2016). 4360: 4097: 4002: 3261: 3214: 3212: 844:layer and the rods and cones are two layers of 6129: 5970: 5968: 5734: 5507: 5453: 5244:Frith, Peggy; Mehta, Arpan R (November 2021). 3687:. New York: Oxford University Press. pp.  3518: 3011: 3000: 2478: 2450:volumetric or high resolution cross-sectional 1991:. The photoreceptors are also cross-linked by 7751: 7375: 6666: 6470: 5922: 5183:(2nd ed.). Cham, Switzerland: Springer. 4717: 3735: 3733: 3262:Sperling, L.; Hubbard, R. (1 February 1975). 2737:for their scientific research on the retina. 2619:, such as pigment epithelium-derived factor ( 2197:that is essentially constant from each input 927:Spectral-Domain OCT macula cross-section scan 762:have a similar, but much simpler, structure. 434:; thus, the retina is considered part of the 7389: 6030:Proceedings of the Royal Society of Medicine 5880:Curace Enrico M.; Auricchio Alberto (2008). 5783: 5615: 5561: 5366:Querques, G; et al. (11 January 2019). 4660:: CS1 maint: DOI inactive as of June 2024 ( 4243:Seminars in Cell & Developmental Biology 4078:: CS1 maint: multiple names: authors list ( 3470:Goldberg AF, Moritz OL, Williams DS (2016). 3382:Labin, A. M.; Ribak, E. N. (16 April 2010). 3209: 2552:, promoter, and intraocular injection site. 5965: 5049: 4863: 4822: 4617: 3127: 2454:of the fine structures of the retina, with 1696:Learn how and when to remove these messages 1588:homogenous region of variable reflectivity 7758: 7744: 7382: 7368: 6673: 6659: 6619:United States National Library of Medicine 6422: 5314: 5243: 4620:"Structure and function of vulture pecten" 4104:Cold Spring Harbor Perspectives in Biology 4093: 4091: 4089: 3730: 3381: 2301:is an outdated treatment method. The term 2247: 2222: 1452:Previously: "cone outer segment tips line" 40: 19:For the electronics screen trademark, see 6576: 6531: 6521: 6263: 6092:"Kepler's Discovery of the Retinal Image" 6057: 5992: 5897: 5856: 5838: 5815: 5766: 5694: 5650: 5598: 5544: 5490: 5401: 5342: 5269: 5020: 4997: 4948: 4938: 4879: 4700: 4668: 4500: 4327: 4296: 4123: 3969: 3787: 3612: 3546: 3536: 3495: 3295: 3169: 3159: 3047: 3029: 1802:Learn how and when to remove this message 1784:Learn how and when to remove this message 1446:cells which encase part of the cone OSs. 700:, which also contain photoreceptors, the 6456:. Sunderland, Mass: Sinauer Associates. 5365: 5093: 4727:Computer Analysis of Images and Patterns 4098:Heavner, W; Pevny, L (1 December 2012). 2934:. Boca Raton, FL: Universal Publishers. 2877:– "the father of modern retinal surgery" 2254: 2110:between the bipolar and ganglion cells. 2069:On-centres and off-centres of the retina 2064: 1813: 930: 922: 914: 578: 569: 540: 532: 6548: 6137:"Santiago Ramón y Cajal – Biographical" 6089: 5974: 5246:"The retina as a window into the brain" 4965: 4674: 4410:. Morgan & Claypool Life Sciences. 4086: 4021: 1948:. Inside the cell the normal levels of 7943: 6023: 5205: 4556:from the original on 21 September 2012 3839: 3837: 3835: 3833: 3831: 3829: 3827: 3683:The Synaptic Organization of the Brain 3587:Sparrow JR, Hicks D, Hamel CP (2010). 3326:from the original on 12 September 2017 3069: 3067: 2744:study calculated that the approximate 1846: 1058:(unclear if it can be observed on OCT) 7739: 7363: 6654: 5796:The Journal of Clinical Investigation 5181:Handbook of visual display technology 4274: 4272: 4189: 3825: 3823: 3821: 3819: 3817: 3815: 3813: 3811: 3809: 3807: 3192: 2735:Nobel Prize in Physiology or Medicine 1764:by removing references to unreliable 1422:, the molecule that absorbs photons. 6108:10.1001/archopht.1973.01000040061014 5931:Progress in Retinal and Eye Research 5730:from the original on 11 August 2017. 5102:Progress in Retinal and Eye Research 4607:from the original on 9 October 2014. 4525:"LIGHT-INDUCED DAMAGE to the RETINA" 4414:from the original on 5 December 2017 4363:Progress in Retinal and Eye Research 3073: 2963: 2961: 2959: 2957: 2955: 2953: 2951: 2797: 2375:sudden acquired retinal degeneration 1974:cyclic nucleotide-gated ion channels 1768:where they are used inappropriately. 1737: 1702: 1661: 396:and reflexive responses such as the 6147:from the original on 6 October 2015 5683:The New England Journal of Medicine 5578:The New England Journal of Medicine 4758:from the original on 9 August 2017. 4432: 4100:"Eye development and retinogenesis" 3064: 3018:Frontiers in Molecular Neuroscience 2054: 680:Inner segment / outer segment layer 622:– contains the synapse between the 455:Inverted versus non-inverted retina 13: 7765: 6791: 6406: 6390:from the original on 24 April 2012 6195: 4650:from the original on 14 July 2015. 4618:Bawa, S.R.; YashRoy, R.C. (1974). 4269: 4056: 3804: 3705: 2613:vascular endothelial growth factor 1393:Photoreceptor integrity line (PIL) 594:– basement membrane elaborated by 14: 7967: 6586: 6177:from the original on 30 June 2017 5335:10.1161/CIRCULATIONAHA.116.023425 4978:Environmental Health Perspectives 3268:The Journal of General Physiology 2948: 2929: 2811:The structures of the eye labeled 1960:bound to the receptor protein to 1677:This article has multiple issues. 1416:photoreceptor outer segments (OS) 1410:Photoreceptor outer segments (OS) 528: 329:. The optics of the eye create a 7633: 6208:from the original on 31 May 2015 6198:"Calculating the speed of sight" 5943:10.1016/j.preteyeres.2008.05.003 5114:10.1016/j.preteyeres.2005.07.001 4575:Bawa S.R.; YashRoy R.C. (1972). 4435:"The retinal pigment epithelium" 4403: 4375:10.1016/j.preteyeres.2014.02.001 3488:10.1016/j.preteyeres.2016.05.003 3233:10.1016/j.preteyeres.2018.03.006 3107: 2847: 2828: 2816: 2804: 2766:in London, in 2006, showed that 2758:& Pearson and colleagues at 2597:age-related macular degeneration 2193:. Computers operate on a single 1742: 1706: 1666: 1449:Poorly distinguishable from RPE. 1418:which contain disks filled with 1368:photoreceptor inner segment (IS) 1315:photoreceptor inner segment (IS) 860:, and the horizontally oriented 723:, a visual relay station in the 144: 110: 71: 6629:Retina – Cell Centered Database 6376: 6350: 6324: 6288: 6220: 6189: 6159: 6083: 6074: 6017: 5873: 5832: 5418: 5359: 5308: 5286: 5237: 5172: 5136: 5014: 4914: 4789: 4762: 4611: 4568: 4542: 4517: 4426: 4397: 4354: 4321: 4234: 4183: 4140: 4050: 4015: 3996: 3978: 3945: 3876: 3672: 3629: 3580: 3519:Arshavsky VY, Burns ME (2012). 3512: 3463: 3441: 3422: 3375: 3338: 3312: 3255: 1685:or discuss these issues on the 1657: 879:Though the rod and cones are a 670:– cell bodies of rods and cones 626:axons and the dendrites of the 3408:10.1103/PhysRevLett.104.158102 3186: 3101: 3076:"Simple Anatomy of the Retina" 2984:. 26 July 2006. Archived from 2970: 2923: 2506:Use of nutritional supplements 2125:is at the heart of a computer 1950:cyclic guanosine monophosphate 1630:factor families including the 1599: 948:OCT Layer / Conventional Label 835:blue field entoptic phenomenon 806:. The optic nerve carries the 712:also are laterally connected. 296: 1: 6485:10.1152/physrev.1991.71.2.447 5643:10.1016/S0140-6736(09)61836-5 5262:10.1016/S1474-4422(21)00332-X 5035:10.1016/s0161-6420(82)34663-1 4681:Journal of General Physiology 3650:10.1080/01677063.2017.1395876 3197:. Vol. 25, no. 15. 2916: 2643:University of New South Wales 2514:Uncommon treatment modalities 2387:lipoprotein lipase deficiency 2121:As an example, the following 1366:The outermost portion of the 1313:The innermost portion of the 702:photosensitive ganglion cells 657:region, this is known as the 7319:Optical coherence tomography 7073:Photosensitive ganglion cell 6437:10.1016/0042-6989(65)90033-7 6024:Dobson, J. F. (March 1925). 5899:10.1016/j.visres.2007.07.027 5158:10.1016/j.ophtha.2003.09.039 4940:10.1371/journal.pmed.1000372 4739:10.1007/978-3-642-23672-3_55 4593:10.1016/0014-4835(72)90129-7 4255:10.1016/j.semcdb.2003.09.005 3862:10.1016/j.ophtha.2014.02.023 3780:10.1016/j.ophtha.2017.09.016 3140:Proc. Natl. Acad. Sci. U.S.A 2590:optical coherence tomography 2557:Leber's congenital amaurosis 2470: 2444:optical coherence tomography 2413: 2094:, and negative weighting as 1292:photoreceptor inner segments 989:Posterior cortical vitreous 910:optical coherence tomography 837:(or Scheerer's phenomenon). 822:(both of which are opaque). 554:(yellow layer), then to the 449: 386:photosensitive ganglion cell 7: 7069:Giant retina ganglion cells 6858:Capillary lamina of choroid 4328:Remington, Lee Ann (2012). 4192:Nature Reviews Neuroscience 4116:10.1101/cshperspect.a008391 3929:10.1007/978-3-642-12041-1_2 3320:"How spiders see the world" 2840: 2628:or slowed down cell death. 2479:Common treatment modalities 2371:progressive retinal atrophy 1895: 1859: 1105:bodies (and some displaced 985:Posterior cortical vitreous 864:connect to ganglion cells. 10: 7972: 7013:Retinal pigment epithelium 7003:External limiting membrane 6452:Wandell, Brian A. (1995). 6310:10.1016/j.exer.2008.01.018 6042:10.1177/003591572501801704 6026:"Herophilus of Alexandria" 5394:10.1038/s41598-018-37271-6 4801:Image Sciences Institute, 4485:10.1038/s41598-023-32897-7 4036:10.1016/j.optm.2008.12.006 3971:10.17925/eor.2017.11.01.59 3958:European Ophthalmic Review 3741:Guyton and Hall Physiology 3605:10.2174/156652410793937813 3574:Guyton and Hall Physiology 3361:10.1016/j.cbpa.2011.02.018 3203:10.1103/physrevfocus.25.15 2930:J, Krause William (2005). 2860: 2742:University of Pennsylvania 2684:Between 1011 and 1021 CE, 2648: 2605:retinopathy of prematurity 2588:(ERG), multifocal ERG and 2538:retinal pigment epithelium 2522: 2307:retinal pigment epithelium 2235:lateral geniculate nucleus 2189:traveling down nerve cell 2058: 1899: 1876: 1582:Choroidal-scleral juncture 1277:External limiting membrane 1044:Internal limiting membrane 816:retinal pigment epithelium 737:nucleus of the optic tract 686:Retinal pigment epithelium 674:External limiting membrane 500:retinal pigment epithelium 479:have been shown to act as 407:through the fibres of the 372:, which are of two types: 208:rēte, tunica interna bulbi 18: 7906: 7873: 7866:Peripheral nervous system 7864: 7773: 7697: 7642: 7631: 7402: 7301: 7262: 7176: 7167: 7101: 7021: 6956: 6946: 6909: 6876: 6848: 6836: 6789: 6732: 6704: 6693: 6634:Histology image: 07901loa 6611:. Retrieved 22 July 2014. 6578:10.4249/scholarpedia.3487 6358:"Retina << EyeWire" 6298:Experimental Eye Research 6096:Archives of Ophthalmology 4839:10.1109/cbms.2013.6627847 4638:(inactive 12 June 2024). 4581:Experimental Eye Research 3679:Shepherd, Gordon (2004). 3031:10.3389/fnmol.2021.656000 2760:University College London 2717:The Retina of Vertebrates 2615:(VEGF) and inhibitors of 1987:which in turn signal the 1585: 1568: 1551: 1533: 1494: 1491: 1470: 1467: 1438:Interdigitation zone (IZ) 1429: 1383: 1380: 1375: 1357: 1252: 1249: 1246: 1233:Henle's nerve fiber layer 1227: 1068: 1065: 1062: 1039: 783:retina is defined by the 774:, at whose centre is the 459:The vertebrate retina is 261: 249: 237: 225: 213: 201: 196: 184: 172: 162: 60: 55: 39: 34: 7671:Ascending and Descending 6641:MedlinePlus Encyclopedia 6036:(Sect_Hist_Med): 19–32. 5298:12 February 2012 at the 2342:hypertensive retinopathy 1889:biometric identification 7008:Layer of rods and cones 6964:Inner limiting membrane 6617:NeuroScience 2nd Ed at 6419:, Maloine, Paris, 1911. 6090:Fishman, R. S. (1973). 5439:10.1089/hum.2005.16.649 4775:University of Groningen 4169:10.1126/science.7892602 3990:Review of Ophthalmology 3887:. New Delhi: Springer. 3538:10.1074/jbc.R111.305243 3434:3 December 2013 at the 3388:Physical Review Letters 3161:10.1073/pnas.0611180104 2764:Moorfields Eye Hospital 2713:Retina der Wirbelthiere 2460:Retinal vessel analysis 2033:lateral geniculate body 2013:difference of Gaussians 1085:travelling towards the 1019:has fully or partially 802:, including the unique 791:, is about 32 mm. 733:suprachiasmatic nucleus 721:lateral geniculate body 592:Inner limiting membrane 25:Retina (disambiguation) 7775:Central nervous system 6830: 6549:Dowling, John (2007). 6523:10.1186/1471-2164-5-50 5293:Seeing into the Future 4874:. pp. 3105–3110. 2896:Retinal vein occlusion 2863:anatomical terminology 2778:and his laboratory at 2727:Haldan Keffer Hartline 2709:Santiago Ramón y Cajal 2546:retinal ganglion cells 2534:adeno-associated virus 2465:cardiovascular disease 2323:laser photocoagulation 2070: 1989:retinal ganglion cells 1830: 1755:self-published sources 1627:retinal ganglion cells 1342:(rarely: mitochondria) 936: 935:Macula histology (OCT) 928: 920: 584: 576: 567: 538: 468:. In contrast, in the 436:central nervous system 423:form the optic nerve. 417:retinal ganglion cells 398:pupillary light reflex 289: 263:Anatomical terminology 191:Central retinal artery 23:. For other uses, see 7324:Eye care professional 7129:Foveal avascular zone 6991:Outer plexiform layer 6979:Inner plexiform layer 6934:Iris sphincter muscle 6827: 6615:Retinal layers image. 6454:Foundations of vision 5994:10.1038/sj.gt.3302366 5858:10.1038/sj.gt.3302030 5705:10.1056/NEJMoa0802268 5591:10.1056/NEJMoa0802315 4890:10.1109/cvpr.2014.397 3429:Foundations of Vision 3322:. Australian Museum. 3195:Physical Review Focus 2911:Persistence of vision 2677:came to refer to the 2632:Organ transplantation 2568:blood retinal barrier 2502:Vitreoretinal surgery 2255:Clinical significance 2239:primary visual cortex 2195:floating-point number 2068: 2059:Further information: 2015:and is the basis for 1825:is at right, and the 1817: 1520:Very Hyper-reflective 1488:Very Hyper-reflective 1377:Very Hyper-reflective 1322:endoplasmic reticulum 1197:Outer plexiform layer 1125:Inner plexiform layer 951:Anatomical Correlate 934: 926: 918: 854:inner plexiform layer 850:outer plexiform layer 651:Outer plexiform layer 620:Inner plexiform layer 582: 573: 544: 536: 428:embryonic development 7705:Accidental viewpoint 7344:Physiological Optics 7314:Ocular immune system 7053:Retina ganglion cell 6593:Histology of the Eye 5975:Rolling, F. (2004). 5759:10.1089/hum.2008.107 5483:10.1089/hum.2009.086 5250:The Lancet Neurology 4833:. pp. 485–488. 4693:10.1085/jgp.78.4.431 4624:Cells Tissues Organs 3280:10.1085/jgp.65.2.235 3264:"Squid retinochrome" 3074:Kolb, Helga (1995). 2881:Evolution of the eye 2625:Neurotrophic factors 2601:diabetic retinopathy 2578:contrast sensitivity 2519:Retinal gene therapy 2490:medication, such as 2361:Retinal diseases in 2346:diabetic retinopathy 2311:pneumatic retinopexy 2270:Macular degeneration 2264:Retinitis pigmentosa 309:) is the innermost, 7610:Vertical–horizontal 7168:Anatomical regions 7029:Photoreceptor cells 6996:Outer nuclear layer 6984:Inner nuclear layer 6974:Ganglion cell layer 6929:Iris dilator muscle 6724:Trabecular meshwork 6569:2007SchpJ...2.3487D 6338:on 13 February 2012 6256:10.1038/nature05161 6248:2006Natur.444..203M 5637:(9701): 1597–1605. 5537:10.1038/mt.2009.277 5386:2019NatSR...9...63Q 4990:10.1289/ehp.1205721 4675:Sherman, T (1981). 4477:2023NatSR..13.6445K 4161:1995Sci...267.1788H 4155:(5205): 1788–1792. 4009:Review of Optometry 3714:The Vertebrate Body 3449:"The Retinal Tunic" 3400:2010PhRvL.104o8102L 3152:2007PNAS..104.8287F 2792:map the connections 2768:photoreceptor cells 2690:intromission theory 2586:electroretinography 2542:photoreceptor cells 2185:Neurons operate on 2082:. This is done by " 2076:photoreceptor cells 1939:trichromatic vision 1912:photoreceptor cells 1847:Energy requirements 1829:is near the centre. 1362:Ellipsoid zone (EZ) 1259:Outer nuclear layer 1157:Inner nuclear layer 1097:Ganglion cell layer 789:horizontal meridian 729:superior colliculus 668:Outer nuclear layer 634:Inner nuclear layer 611:Ganglion cell layer 444:blood–brain barrier 388:, is important for 370:photoreceptor cells 7710:Auditory illusions 7505:Impossible trident 6831: 6609:University of Utah 6141:www.nobelprize.org 5839:Bennett J (2003). 5747:Human Gene Therapy 5471:Human Gene Therapy 5427:Human Gene Therapy 5373:Scientific Reports 4803:Utrecht University 4465:Scientific Reports 4298:10.1242/dev.178103 3846:Spaide, Richard F. 3476:Prog Retin Eye Res 3438:, Brian A. Wandell 3221:Prog Retin Eye Res 2861:This article uses 2424:fundus photography 2394:Retinal Detachment 2382:Lipaemia retinalis 2303:retinal detachment 2295:retinal separation 2280:Cone-rod dystrophy 2071: 2049:information theory 1831: 1718:. You can help by 1397:The photoreceptor 1015:In eyes where the 937: 929: 921: 696:; transmission to 630:and amacrine cells 585: 577: 568: 539: 364:interconnected by 7956:Human eye anatomy 7938: 7937: 7934: 7933: 7733: 7732: 7725:Temporal illusion 7720:Tactile illusions 7690:(2015 photograph) 7391:Optical illusions 7357: 7356: 7349:Visual perception 7297: 7296: 7264:Posterior segment 7232:Posterior chamber 7163: 7162: 7065:Bistratified cell 6969:Nerve fiber layer 6942: 6941: 6886:Ciliary processes 6787: 6786: 6512:for eye colour). 6463:978-0-87893-853-7 6242:(7116): 203–207. 6196:Reilly, Michael. 5689:(21): 2231–2239. 5585:(21): 2240–2248. 5525:Molecular Therapy 5329:(18): 1328–1338. 5222:978-0-323-02598-0 5071:10.1042/cs1030111 5029:(10): 1132–1145. 4899:978-1-4799-5118-5 4848:978-1-4799-1053-3 4748:978-3-642-23671-6 4636:10.1159/000144308 4529:photobiology.info 4404:Kiel, Jeffrey W. 4339:978-1-4377-1926-0 4291:(17): dev178103. 4005:"The ABCs of OCT" 3723:978-0-03-910284-5 3698:978-0-19-515956-1 3146:(20): 8287–8292. 2980:(Press release). 2941:978-1-58112-468-2 2798:Additional images 2436:electroretinogram 2367:retinal dysplasia 2338:diabetes mellitus 2168: 2167: 2001:action potentials 1836:ophthalmic artery 1819:Fundus photograph 1812: 1811: 1804: 1794: 1793: 1786: 1736: 1735: 1700: 1597: 1596: 1460:Hyper-reflective 1320:Smooth and rough 1296:Hyper-reflective 1219:Hyper-reflective 1144:Hyper-reflective 1130:Synapses between 1074:Nerve fiber layer 1063:Hyper-reflective 992:Hyper-reflective 827:white blood cells 798:cells and two of 601:Nerve fibre layer 537:Section of retina 506:) rarely occurs. 413:action potentials 394:circadian rhythms 343:visual perception 277: 276: 272: 7963: 7871: 7870: 7760: 7753: 7746: 7737: 7736: 7637: 7590:Schroeder stairs 7565:Peripheral drift 7560:Penrose triangle 7384: 7377: 7370: 7361: 7360: 7334:Refractive error 7272:Vitreous chamber 7217:Anterior chamber 7178:Anterior segment 7174: 7173: 6954: 6953: 6863:Bruch's membrane 6846: 6845: 6838:Uvea / vascular 6794: 6714:Episcleral layer 6702: 6701: 6675: 6668: 6661: 6652: 6651: 6582: 6580: 6545: 6535: 6525: 6496: 6467: 6448: 6413:S. Ramón y Cajal 6400: 6399: 6397: 6395: 6380: 6374: 6373: 6371: 6369: 6364:on 24 March 2012 6360:. Archived from 6354: 6348: 6347: 6345: 6343: 6334:. 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862:horizontal cells 646:horizontal cells 552:horizontal cells 485:light scattering 304: 298: 269:edit on Wikidata 266: 158: 157: 154: 153: 150: 136: 135: 132: 131: 128: 125: 122: 119: 116: 109: 100: 99: 96: 95: 92: 89: 86: 83: 80: 77: 70: 44: 32: 31: 7971: 7970: 7966: 7965: 7964: 7962: 7961: 7960: 7941: 7940: 7939: 7930: 7921:Parasympathetic 7902: 7860: 7769: 7764: 7734: 7729: 7693: 7643:Popular culture 7638: 7629: 7600:Spinning dancer 7420:Ambiguous image 7398: 7388: 7358: 7353: 7293: 7258: 7247:Capsule of lens 7202:Lacrimal system 7169: 7159: 7119:Parafoveal area 7114:Perifoveal area 7097: 7041:Horizontal cell 7017: 6938: 6905: 6872: 6868:Sattler's layer 6839: 6832: 6826: 6783: 6728: 6719:Schlemm's canal 6697: 6689: 6681:Anatomy of the 6679: 6589: 6464: 6431:(11): 583–601. 6409: 6407:Further reading 6404: 6403: 6393: 6391: 6382: 6381: 6377: 6367: 6365: 6356: 6355: 6351: 6341: 6339: 6330: 6329: 6325: 6293: 6289: 6231: 6225: 6221: 6211: 6209: 6194: 6190: 6180: 6178: 6165: 6164: 6160: 6150: 6148: 6135: 6134: 6130: 6120: 6118: 6088: 6084: 6079: 6075: 6022: 6018: 5987:(S1): S26–S32. 5973: 5966: 5927: 5923: 5886:Vision Research 5878: 5874: 5851:(11): 977–982. 5837: 5833: 5788: 5784: 5753:(10): 979–990. 5739: 5735: 5727: 5696:10.1.1.574.4003 5678: 5672: 5668: 5623: 5616: 5569: 5562: 5517: 5508: 5477:(9): 999–1004. 5463: 5454: 5423: 5419: 5364: 5360: 5313: 5309: 5300:Wayback Machine 5291: 5287: 5242: 5238: 5223: 5211: 5210: 5206: 5191: 5177: 5173: 5141: 5137: 5098: 5094: 5054: 5050: 5019: 5015: 4970: 4966: 4919: 4915: 4900: 4881:10.1.1.1026.949 4868: 4864: 4849: 4827: 4823: 4813: 4811: 4795: 4794: 4790: 4780: 4778: 4768: 4767: 4763: 4755: 4749: 4730: 4722: 4718: 4673: 4669: 4653: 4652: 4616: 4612: 4573: 4569: 4559: 4557: 4548: 4547: 4543: 4533: 4531: 4523: 4522: 4518: 4457: 4453: 4443: 4441: 4433:Strauss, Olaf. 4431: 4427: 4417: 4415: 4402: 4398: 4359: 4355: 4340: 4326: 4322: 4277: 4270: 4239: 4235: 4204:10.1038/nrn3767 4188: 4184: 4145: 4141: 4110:(12): a008391. 4096: 4087: 4071: 4070: 4063: 4061: 4055: 4051: 4020: 4016: 4001: 3997: 3984: 3983: 3979: 3950: 3946: 3939: 3917: 3910: 3895: 3881: 3877: 3842: 3805: 3763: 3757: 3748: 3739: 3738: 3731: 3724: 3710: 3706: 3699: 3677: 3673: 3634: 3630: 3585: 3581: 3572: 3571: 3564: 3517: 3513: 3468: 3464: 3459:on 18 May 2007. 3447: 3446: 3442: 3436:Wayback Machine 3427: 3423: 3380: 3376: 3343: 3339: 3329: 3327: 3318: 3317: 3313: 3260: 3256: 3217: 3210: 3191: 3187: 3132: 3128: 3118: 3116: 3106: 3102: 3092: 3090: 3072: 3065: 3010: 3001: 2991: 2989: 2976: 2975: 2971: 2966: 2949: 2942: 2928: 2924: 2919: 2866: 2855:Medicine portal 2853: 2848: 2846: 2843: 2836: 2833: 2824: 2821: 2812: 2809: 2800: 2788:Citizen science 2776:Sebastian Seung 2701:Johannes Kepler 2694:emission theory 2651: 2635:Transplantation 2564:Tight junctions 2527: 2516: 2481: 2473: 2428:adaptive optics 2416: 2402:Night blindness 2325:and pars plana 2257: 2063: 2057: 2008:receptive field 1923:scotopic vision 1919:photopic vision 1908: 1898: 1885: 1875: 1862: 1849: 1808: 1797: 1796: 1795: 1790: 1779: 1773: 1770: 1759: 1747: 1743: 1732: 1726: 1723: 1716:needs expansion 1671: 1667: 1660: 1602: 1548:Sattler's layer 1484:phagosome zone 1309:Myoid zone (MZ) 1028:Hypo-reflective 906: 756:vitreous humour 659:Fiber layer of 603:– axons of the 531: 496:resolving power 489:fovea centralis 457: 452: 311:light-sensitive 299: 273: 147: 143: 138: 113: 104: 103: 102: 74: 65: 64: 51: 28: 17: 16:Part of the eye 12: 11: 5: 7969: 7959: 7958: 7953: 7936: 7935: 7932: 7931: 7929: 7928: 7923: 7918: 7912: 7910: 7904: 7903: 7901: 7900: 7895: 7893:Cranial nerves 7890: 7885: 7879: 7877: 7868: 7862: 7861: 7859: 7858: 7857: 7856: 7851: 7850: 7849: 7844: 7843: 7842: 7837: 7822: 7817: 7816: 7815: 7810: 7805: 7790: 7785: 7779: 7777: 7771: 7770: 7767:Nervous system 7763: 7762: 7755: 7748: 7740: 7731: 7730: 7728: 7727: 7722: 7717: 7712: 7707: 7701: 7699: 7695: 7694: 7692: 7691: 7683: 7682:(1961 drawing) 7675: 7674:(1960 drawing) 7667: 7659: 7652: 7646: 7644: 7640: 7639: 7632: 7630: 7628: 7627: 7622: 7617: 7612: 7607: 7602: 7597: 7595:Shepard tables 7592: 7587: 7582: 7577: 7572: 7567: 7562: 7557: 7555:Penrose stairs 7552: 7547: 7542: 7537: 7532: 7527: 7522: 7517: 7512: 7507: 7502: 7497: 7492: 7487: 7482: 7477: 7472: 7467: 7462: 7457: 7452: 7450:Checker shadow 7447: 7442: 7437: 7432: 7430:Autostereogram 7427: 7422: 7417: 7412: 7406: 7404: 7400: 7399: 7387: 7386: 7379: 7372: 7364: 7355: 7354: 7352: 7351: 7346: 7341: 7336: 7331: 7326: 7321: 7316: 7311: 7305: 7303: 7299: 7298: 7295: 7294: 7292: 7291: 7286: 7281: 7280: 7279: 7268: 7266: 7260: 7259: 7257: 7256: 7255: 7254: 7252:Zonule of Zinn 7249: 7239: 7234: 7229: 7224: 7222:Aqueous humour 7219: 7214: 7209: 7182: 7180: 7171: 7165: 7164: 7161: 7160: 7158: 7157: 7152: 7151: 7150: 7140: 7139: 7138: 7137: 7136: 7131: 7121: 7116: 7105: 7103: 7099: 7098: 7096: 7095: 7025: 7023: 7019: 7018: 7016: 7015: 7010: 7005: 6999: 6998: 6993: 6987: 6986: 6981: 6976: 6971: 6966: 6960: 6958: 6951: 6944: 6943: 6940: 6939: 6937: 6936: 6931: 6926: 6921: 6915: 6913: 6907: 6906: 6904: 6903: 6898: 6893: 6891:Ciliary muscle 6888: 6882: 6880: 6874: 6873: 6871: 6870: 6865: 6860: 6854: 6852: 6843: 6834: 6833: 6790: 6788: 6785: 6784: 6782: 6781: 6780: 6779: 6774: 6769: 6764: 6759: 6754: 6744: 6738: 6736: 6730: 6729: 6727: 6726: 6721: 6716: 6710: 6708: 6699: 6691: 6690: 6678: 6677: 6670: 6663: 6655: 6649: 6648: 6637: 6631: 6626: 6621: 6612: 6601: 6596: 6588: 6587:External links 6585: 6584: 6583: 6546: 6497: 6479:(2): 447–480. 6468: 6462: 6449: 6420: 6408: 6405: 6402: 6401: 6375: 6349: 6323: 6304:(4): 691–700. 6287: 6219: 6188: 6171:nobelprize.org 6158: 6128: 6082: 6073: 6016: 5964: 5937:(4): 391–419. 5921: 5892:(3): 353–359. 5872: 5831: 5782: 5733: 5666: 5614: 5560: 5531:(3): 643–650. 5506: 5452: 5433:(6): 649–663. 5417: 5358: 5307: 5285: 5236: 5221: 5204: 5189: 5171: 5135: 5092: 5065:(2): 111–116. 5048: 5013: 4964: 4913: 4898: 4862: 4847: 4821: 4788: 4761: 4747: 4716: 4687:(4): 431–453. 4667: 4630:(3): 473–480. 4610: 4567: 4541: 4516: 4451: 4425: 4396: 4353: 4338: 4320: 4268: 4233: 4198:(9): 615–627. 4182: 4139: 4085: 4049: 4030:(6): 277–278. 4014: 3995: 3977: 3944: 3938:978-3642120404 3937: 3908: 3894:978-8132236108 3893: 3875: 3803: 3774:(3): 407–422. 3746: 3743:. p. 609. 3729: 3722: 3704: 3697: 3671: 3644:(4): 203–215. 3628: 3599:(9): 802–823. 3579: 3576:. p. 612. 3562: 3511: 3462: 3440: 3421: 3394:(15): 158102. 3374: 3355:(2): 182–187. 3337: 3311: 3274:(2): 235–251. 3254: 3208: 3185: 3126: 3100: 3063: 2999: 2969: 2947: 2940: 2921: 2920: 2918: 2915: 2914: 2913: 2908: 2903: 2898: 2893: 2888: 2883: 2878: 2872: 2859: 2858: 2842: 2839: 2838: 2837: 2834: 2827: 2825: 2822: 2815: 2813: 2810: 2803: 2799: 2796: 2686:Ibn Al-Haytham 2650: 2647: 2609:growth factors 2566:that form the 2523:Main article: 2515: 2512: 2511: 2510: 2507: 2504: 2499: 2496:corticosteroid 2480: 2477: 2472: 2469: 2440:non-invasively 2420:Ophthalmoscopy 2415: 2412: 2407: 2406: 2398: 2390: 2378: 2359: 2358:of the retina. 2352:Retinoblastoma 2349: 2330: 2315:scleral buckle 2291: 2290:in the retina. 2277: 2267: 2256: 2253: 2217: 2216: 2213:amacrine cells 2205: 2202: 2183: 2180: 2166: 2165: 2162: 2159: 2155: 2154: 2151: 2148: 2144: 2143: 2140: 2137: 2115:edge detection 2080:ganglion cells 2056: 2053: 2025:visual pathway 2017:edge detection 1997:amacrine cells 1927:mesopic vision 1897: 1894: 1874: 1871: 1861: 1858: 1848: 1845: 1810: 1809: 1792: 1791: 1750: 1748: 1741: 1734: 1733: 1713: 1711: 1701: 1675: 1674: 1672: 1665: 1659: 1656: 1601: 1598: 1595: 1594: 1592: 1590: 1584: 1579: 1575: 1574: 1572: 1570: 1567: 1565:Haller's layer 1562: 1558: 1557: 1555: 1553: 1550: 1545: 1541: 1540: 1538: 1535: 1532: 1527: 1523: 1522: 1517: 1507: 1506: 1503: 1496: 1495: 1493: 1490: 1485: 1479: 1469: 1465: 1464: 1461: 1458: 1442:Apices of the 1440: 1435: 1431: 1430: 1428: 1426: 1423: 1412: 1407: 1403: 1402: 1395: 1389:IS/OS junction 1385: 1384: 1382: 1379: 1374: 1364: 1359: 1355: 1354: 1352: 1349: 1346: 1345: 1344: 1339: 1334: 1329: 1324: 1311: 1306: 1302: 1301: 1299: 1297: 1294: 1280: 1274: 1270: 1269: 1262: 1254: 1253: 1251: 1248: 1245: 1236: 1229: 1225: 1224: 1222: 1220: 1217: 1200: 1194: 1190: 1189: 1187: 1185: 1182: 1160: 1154: 1150: 1149: 1147: 1145: 1142: 1140:ganglion cells 1128: 1122: 1118: 1117: 1115: 1113: 1110: 1107:amacrine cells 1100: 1094: 1090: 1089: 1077: 1070: 1069: 1067: 1064: 1061: 1047: 1041: 1037: 1036: 1032: 1030: 1025: 1012: 1005: 999: 998: 996: 993: 990: 987: 982: 978: 977: 971: 962: 952: 949: 946: 905: 902: 893:amacrine cells 885:ganglion cells 804:ribbon synapse 710:amacrine cells 698:ganglion cells 690: 689: 683: 677: 671: 665: 648: 638:amacrine cells 631: 617: 615:amacrine cells 608: 598: 560:ganglion cells 556:amacrine cells 530: 529:Retinal layers 527: 456: 453: 451: 448: 426:In vertebrate 275: 274: 265: 259: 258: 253: 247: 246: 241: 235: 234: 229: 223: 222: 217: 211: 210: 205: 199: 198: 194: 193: 188: 182: 181: 176: 170: 169: 164: 160: 159: 62: 58: 57: 53: 52: 45: 37: 36: 21:Retina display 15: 9: 6: 4: 3: 2: 7968: 7957: 7954: 7952: 7951:Visual system 7949: 7948: 7946: 7927: 7924: 7922: 7919: 7917: 7914: 7913: 7911: 7909: 7905: 7899: 7896: 7894: 7891: 7889: 7886: 7884: 7883:Sensory nerve 7881: 7880: 7878: 7876: 7872: 7869: 7867: 7863: 7855: 7854:Limbic system 7852: 7848: 7845: 7841: 7838: 7836: 7833: 7832: 7831: 7828: 7827: 7826: 7823: 7821: 7818: 7814: 7811: 7809: 7806: 7804: 7801: 7800: 7799: 7796: 7795: 7794: 7791: 7789: 7786: 7784: 7781: 7780: 7778: 7776: 7772: 7768: 7761: 7756: 7754: 7749: 7747: 7742: 7741: 7738: 7726: 7723: 7721: 7718: 7716: 7713: 7711: 7708: 7706: 7703: 7702: 7700: 7696: 7689: 7688: 7684: 7681: 7680: 7676: 7673: 7672: 7668: 7665: 7664: 7660: 7658: 7657: 7653: 7651: 7648: 7647: 7645: 7641: 7636: 7626: 7623: 7621: 7618: 7616: 7613: 7611: 7608: 7606: 7603: 7601: 7598: 7596: 7593: 7591: 7588: 7586: 7583: 7581: 7578: 7576: 7573: 7571: 7568: 7566: 7563: 7561: 7558: 7556: 7553: 7551: 7548: 7546: 7543: 7541: 7538: 7536: 7533: 7531: 7528: 7526: 7523: 7521: 7518: 7516: 7513: 7511: 7508: 7506: 7503: 7501: 7498: 7496: 7493: 7491: 7488: 7486: 7485:Fraser spiral 7483: 7481: 7478: 7476: 7473: 7471: 7468: 7466: 7463: 7461: 7458: 7456: 7453: 7451: 7448: 7446: 7443: 7441: 7438: 7436: 7433: 7431: 7428: 7426: 7423: 7421: 7418: 7416: 7413: 7411: 7408: 7407: 7405: 7401: 7396: 7392: 7385: 7380: 7378: 7373: 7371: 7366: 7365: 7362: 7350: 7347: 7345: 7342: 7340: 7339:Accommodation 7337: 7335: 7332: 7330: 7327: 7325: 7322: 7320: 7317: 7315: 7312: 7310: 7307: 7306: 7304: 7300: 7290: 7287: 7285: 7282: 7278: 7277:Vitreous body 7275: 7274: 7273: 7270: 7269: 7267: 7265: 7261: 7253: 7250: 7248: 7245: 7244: 7243: 7240: 7238: 7235: 7233: 7230: 7228: 7225: 7223: 7220: 7218: 7215: 7213: 7212:Fibrous tunic 7210: 7207: 7203: 7199: 7195: 7191: 7187: 7184: 7183: 7181: 7179: 7175: 7172: 7166: 7156: 7153: 7149: 7146: 7145: 7144: 7141: 7135: 7132: 7130: 7127: 7126: 7125: 7122: 7120: 7117: 7115: 7112: 7111: 7110: 7107: 7106: 7104: 7100: 7094: 7090: 7086: 7082: 7078: 7074: 7070: 7066: 7062: 7058: 7054: 7050: 7049:Amacrine cell 7046: 7042: 7038: 7034: 7030: 7027: 7026: 7024: 7020: 7014: 7011: 7009: 7006: 7004: 7001: 7000: 6997: 6994: 6992: 6989: 6988: 6985: 6982: 6980: 6977: 6975: 6972: 6970: 6967: 6965: 6962: 6961: 6959: 6955: 6952: 6949: 6945: 6935: 6932: 6930: 6927: 6925: 6922: 6920: 6917: 6916: 6914: 6912: 6908: 6902: 6899: 6897: 6894: 6892: 6889: 6887: 6884: 6883: 6881: 6879: 6875: 6869: 6866: 6864: 6861: 6859: 6856: 6855: 6853: 6851: 6847: 6844: 6841: 6835: 6778: 6775: 6773: 6770: 6768: 6765: 6763: 6760: 6758: 6755: 6753: 6750: 6749: 6748: 6745: 6743: 6740: 6739: 6737: 6735: 6731: 6725: 6722: 6720: 6717: 6715: 6712: 6711: 6709: 6707: 6703: 6700: 6696: 6695:Fibrous tunic 6692: 6688: 6684: 6676: 6671: 6669: 6664: 6662: 6657: 6656: 6653: 6647: 6643: 6642: 6638: 6635: 6632: 6630: 6627: 6625: 6622: 6620: 6616: 6613: 6610: 6606: 6602: 6600: 6597: 6594: 6591: 6590: 6579: 6574: 6570: 6566: 6562: 6558: 6557: 6552: 6547: 6543: 6539: 6534: 6529: 6524: 6519: 6515: 6511: 6510:transcriptome 6507: 6503: 6498: 6494: 6490: 6486: 6482: 6478: 6474: 6469: 6465: 6459: 6455: 6450: 6446: 6442: 6438: 6434: 6430: 6426: 6421: 6418: 6414: 6411: 6410: 6389: 6385: 6379: 6363: 6359: 6353: 6337: 6333: 6327: 6319: 6315: 6311: 6307: 6303: 6299: 6291: 6283: 6279: 6275: 6271: 6266: 6265:2027.42/62596 6261: 6257: 6253: 6249: 6245: 6241: 6237: 6230: 6223: 6207: 6203: 6202:New Scientist 6199: 6192: 6176: 6172: 6168: 6162: 6146: 6142: 6138: 6132: 6117: 6113: 6109: 6105: 6101: 6097: 6093: 6086: 6077: 6069: 6065: 6060: 6055: 6051: 6047: 6043: 6039: 6035: 6031: 6027: 6020: 6012: 6008: 6004: 6000: 5995: 5990: 5986: 5982: 5978: 5971: 5969: 5960: 5956: 5952: 5948: 5944: 5940: 5936: 5932: 5925: 5917: 5913: 5909: 5905: 5900: 5895: 5891: 5887: 5883: 5876: 5868: 5864: 5859: 5854: 5850: 5846: 5842: 5835: 5827: 5823: 5818: 5813: 5809: 5805: 5801: 5797: 5793: 5786: 5778: 5774: 5769: 5764: 5760: 5756: 5752: 5748: 5744: 5737: 5726: 5722: 5718: 5714: 5710: 5706: 5702: 5697: 5692: 5688: 5684: 5677: 5670: 5662: 5658: 5653: 5648: 5644: 5640: 5636: 5632: 5628: 5621: 5619: 5610: 5606: 5601: 5596: 5592: 5588: 5584: 5580: 5579: 5574: 5567: 5565: 5556: 5552: 5547: 5542: 5538: 5534: 5530: 5526: 5522: 5515: 5513: 5511: 5502: 5498: 5493: 5488: 5484: 5480: 5476: 5472: 5468: 5461: 5459: 5457: 5448: 5444: 5440: 5436: 5432: 5428: 5421: 5413: 5409: 5404: 5399: 5395: 5391: 5387: 5383: 5379: 5375: 5374: 5369: 5362: 5354: 5350: 5345: 5340: 5336: 5332: 5328: 5324: 5323: 5318: 5311: 5304: 5301: 5297: 5294: 5289: 5281: 5277: 5272: 5267: 5263: 5259: 5255: 5251: 5247: 5240: 5232: 5228: 5224: 5218: 5214: 5208: 5200: 5196: 5192: 5190:9783319143460 5186: 5182: 5175: 5167: 5163: 5159: 5155: 5151: 5147: 5146:Ophthalmology 5139: 5131: 5127: 5123: 5119: 5115: 5111: 5108:(1): 99–127. 5107: 5103: 5096: 5088: 5084: 5080: 5076: 5072: 5068: 5064: 5060: 5052: 5044: 5040: 5036: 5032: 5028: 5024: 5023:Ophthalmology 5017: 5009: 5005: 5000: 4995: 4991: 4987: 4983: 4979: 4975: 4968: 4960: 4956: 4951: 4946: 4941: 4936: 4932: 4928: 4924: 4917: 4909: 4905: 4901: 4895: 4891: 4887: 4882: 4877: 4873: 4866: 4858: 4854: 4850: 4844: 4840: 4836: 4832: 4825: 4809: 4805: 4804: 4798: 4792: 4777: 4776: 4771: 4765: 4754: 4750: 4744: 4740: 4736: 4729: 4728: 4720: 4712: 4708: 4703: 4698: 4694: 4690: 4686: 4682: 4678: 4671: 4663: 4657: 4649: 4645: 4641: 4637: 4633: 4629: 4625: 4621: 4614: 4606: 4602: 4598: 4594: 4590: 4586: 4582: 4578: 4571: 4555: 4551: 4545: 4530: 4526: 4520: 4512: 4508: 4503: 4498: 4494: 4490: 4486: 4482: 4478: 4474: 4470: 4466: 4462: 4455: 4440: 4436: 4429: 4413: 4409: 4408: 4400: 4392: 4388: 4384: 4380: 4376: 4372: 4368: 4364: 4357: 4349: 4345: 4341: 4335: 4331: 4324: 4316: 4312: 4308: 4304: 4299: 4294: 4290: 4286: 4282: 4275: 4273: 4264: 4260: 4256: 4252: 4248: 4244: 4237: 4229: 4225: 4221: 4217: 4213: 4209: 4205: 4201: 4197: 4193: 4186: 4178: 4174: 4170: 4166: 4162: 4158: 4154: 4150: 4143: 4135: 4131: 4126: 4121: 4117: 4113: 4109: 4105: 4101: 4094: 4092: 4090: 4081: 4075: 4060: 4053: 4045: 4041: 4037: 4033: 4029: 4025: 4018: 4010: 4006: 3999: 3991: 3987: 3981: 3972: 3967: 3963: 3959: 3955: 3948: 3940: 3934: 3930: 3926: 3922: 3915: 3913: 3904: 3900: 3896: 3890: 3886: 3879: 3871: 3867: 3863: 3859: 3855: 3851: 3850:Ophthalmology 3847: 3840: 3838: 3836: 3834: 3832: 3830: 3828: 3826: 3824: 3822: 3820: 3818: 3816: 3814: 3812: 3810: 3808: 3799: 3795: 3790: 3785: 3781: 3777: 3773: 3769: 3768:Ophthalmology 3762: 3755: 3753: 3751: 3742: 3736: 3734: 3725: 3719: 3715: 3708: 3700: 3694: 3690: 3685: 3684: 3675: 3667: 3663: 3659: 3655: 3651: 3647: 3643: 3639: 3632: 3624: 3620: 3615: 3610: 3606: 3602: 3598: 3594: 3590: 3583: 3575: 3569: 3567: 3558: 3554: 3549: 3544: 3539: 3534: 3530: 3526: 3522: 3515: 3507: 3503: 3498: 3493: 3489: 3485: 3481: 3477: 3473: 3466: 3458: 3454: 3450: 3444: 3437: 3433: 3430: 3425: 3417: 3413: 3409: 3405: 3401: 3397: 3393: 3389: 3385: 3378: 3370: 3366: 3362: 3358: 3354: 3350: 3349: 3341: 3325: 3321: 3315: 3307: 3303: 3298: 3293: 3289: 3285: 3281: 3277: 3273: 3269: 3265: 3258: 3250: 3246: 3242: 3238: 3234: 3230: 3226: 3222: 3215: 3213: 3204: 3200: 3196: 3189: 3181: 3177: 3172: 3167: 3162: 3157: 3153: 3149: 3145: 3141: 3137: 3130: 3115: 3111: 3108:Kolb, Helga. 3104: 3089: 3085: 3081: 3077: 3070: 3068: 3059: 3055: 3050: 3045: 3041: 3037: 3032: 3027: 3023: 3019: 3015: 3008: 3006: 3004: 2987: 2983: 2982:PENN Medicine 2979: 2973: 2964: 2962: 2960: 2958: 2956: 2954: 2952: 2943: 2937: 2933: 2926: 2922: 2912: 2909: 2907: 2904: 2902: 2899: 2897: 2894: 2892: 2889: 2887: 2886:Duplex retina 2884: 2882: 2879: 2876: 2873: 2871: 2868: 2867: 2864: 2856: 2845: 2831: 2826: 2819: 2814: 2807: 2802: 2801: 2795: 2793: 2789: 2785: 2781: 2777: 2772: 2769: 2765: 2761: 2757: 2753: 2751: 2747: 2743: 2738: 2736: 2733:won the 1967 2732: 2731:Ragnar Granit 2728: 2724: 2720: 2718: 2714: 2710: 2705: 2702: 2697: 2695: 2691: 2687: 2682: 2680: 2676: 2672: 2668: 2664: 2660: 2656: 2646: 2644: 2640: 2636: 2633: 2629: 2626: 2622: 2618: 2614: 2610: 2606: 2602: 2598: 2593: 2591: 2587: 2583: 2579: 2574: 2569: 2565: 2560: 2558: 2553: 2551: 2547: 2543: 2539: 2535: 2531: 2526: 2521: 2520: 2508: 2505: 2503: 2500: 2497: 2493: 2489: 2486: 2485: 2484: 2476: 2468: 2466: 2461: 2457: 2453: 2449: 2445: 2441: 2437: 2432: 2429: 2425: 2421: 2411: 2404: 2403: 2399: 2396: 2395: 2391: 2388: 2384: 2383: 2379: 2376: 2372: 2368: 2364: 2360: 2357: 2353: 2350: 2347: 2343: 2339: 2335: 2331: 2328: 2324: 2320: 2316: 2312: 2308: 2304: 2300: 2296: 2292: 2289: 2285: 2281: 2278: 2275: 2271: 2268: 2265: 2262: 2261: 2260: 2252: 2250: 2245: 2242: 2240: 2236: 2232: 2227: 2225: 2220: 2214: 2210: 2207:Finally, the 2206: 2203: 2200: 2196: 2192: 2188: 2184: 2181: 2177: 2176: 2175: 2172: 2163: 2160: 2157: 2156: 2152: 2149: 2146: 2145: 2141: 2138: 2135: 2134: 2131: 2128: 2124: 2119: 2116: 2111: 2109: 2105: 2099: 2097: 2093: 2087: 2085: 2084:decorrelation 2081: 2077: 2067: 2062: 2052: 2050: 2046: 2042: 2036: 2034: 2030: 2029:optic chiasma 2026: 2021: 2018: 2014: 2009: 2004: 2002: 1998: 1994: 1990: 1986: 1985:bipolar cells 1982: 1977: 1975: 1971: 1967: 1966:trans-retinal 1963: 1959: 1955: 1951: 1947: 1942: 1940: 1936: 1930: 1928: 1924: 1920: 1915: 1913: 1907: 1906:Visual acuity 1903: 1893: 1890: 1884: 1880: 1870: 1868: 1857: 1853: 1844: 1840: 1837: 1828: 1824: 1820: 1816: 1806: 1803: 1788: 1785: 1777: 1767: 1763: 1757: 1756: 1751:This section 1749: 1740: 1739: 1730: 1721: 1717: 1714:This section 1712: 1709: 1705: 1704: 1699: 1697: 1690: 1689: 1684: 1683: 1678: 1673: 1664: 1663: 1655: 1653: 1649: 1645: 1639: 1637: 1633: 1628: 1623: 1619: 1615: 1611: 1607: 1593: 1591: 1589: 1583: 1580: 1577: 1576: 1573: 1571: 1566: 1563: 1560: 1559: 1556: 1554: 1549: 1546: 1543: 1542: 1539: 1536: 1531: 1528: 1525: 1524: 1521: 1518: 1516: 1512: 1509: 1508: 1504: 1501: 1498: 1497: 1489: 1486: 1483: 1480: 1477: 1473: 1466: 1462: 1459: 1457: 1456: 1453: 1450: 1445: 1441: 1439: 1436: 1433: 1432: 1427: 1424: 1421: 1417: 1413: 1411: 1408: 1405: 1404: 1400: 1396: 1394: 1390: 1387: 1386: 1378: 1373: 1369: 1365: 1363: 1360: 1356: 1353: 1350: 1347: 1343: 1340: 1338: 1335: 1333: 1330: 1328: 1325: 1323: 1319: 1318: 1316: 1312: 1310: 1307: 1304: 1303: 1300: 1298: 1295: 1293: 1289: 1285: 1281: 1278: 1275: 1272: 1271: 1267: 1266:photoreceptor 1263: 1260: 1257:(Outer half) 1256: 1255: 1244: 1240: 1239:Photoreceptor 1237: 1234: 1231:(Inner half) 1230: 1226: 1223: 1221: 1218: 1216: 1212: 1208: 1207:photoreceptor 1204: 1201: 1198: 1195: 1192: 1191: 1188: 1186: 1183: 1181: 1179: 1173: 1169: 1165: 1161: 1158: 1155: 1152: 1151: 1148: 1146: 1143: 1141: 1137: 1133: 1129: 1126: 1123: 1120: 1119: 1116: 1114: 1111: 1108: 1104: 1103:Ganglion cell 1101: 1098: 1095: 1092: 1091: 1088: 1084: 1081: 1080:Ganglion cell 1078: 1075: 1072: 1071: 1060: 1059: 1055: 1052: 1048: 1045: 1042: 1038: 1035: 1033: 1031: 1029: 1026: 1024: 1022: 1018: 1013: 1011: 1010: 1006: 1004: 1001: 1000: 997: 994: 991: 988: 986: 983: 980: 979: 976: 972: 970: 967: 963: 961: 960: 956: 953: 950: 947: 944: 943: 940: 933: 925: 917: 913: 911: 901: 899: 894: 890: 886: 882: 877: 875: 871: 865: 863: 859: 858:bipolar cells 855: 851: 847: 843: 842:ganglion-cell 838: 836: 832: 828: 823: 821: 817: 812: 809: 808:ganglion-cell 805: 801: 797: 792: 790: 786: 781: 777: 773: 769: 768:optic papilla 763: 761: 757: 753: 749: 744: 743:in primates. 742: 738: 734: 730: 726: 722: 718: 713: 711: 707: 703: 699: 695: 694:bipolar cells 687: 684: 681: 678: 675: 672: 669: 666: 663: 662: 656: 652: 649: 647: 643: 642:bipolar cells 639: 635: 632: 629: 625: 621: 618: 616: 612: 609: 606: 605:ganglion cell 602: 599: 597: 593: 590: 589: 588: 581: 572: 565: 564:Ramón y Cajal 561: 557: 553: 549: 543: 535: 526: 522: 518: 514: 512: 507: 505: 501: 497: 492: 490: 486: 482: 478: 473: 471: 467: 462: 447: 445: 441: 440:noninvasively 437: 433: 429: 424: 422: 418: 414: 410: 406: 401: 399: 395: 391: 387: 383: 379: 375: 371: 367: 363: 358: 356: 352: 348: 344: 340: 339:visual cortex 336: 332: 328: 324: 320: 316: 312: 308: 303: 295: 'net'; 294: 293: 292: 286: 282: 270: 264: 260: 257: 254: 252: 248: 245: 242: 240: 236: 233: 230: 228: 224: 221: 218: 216: 212: 209: 206: 204: 200: 195: 192: 189: 187: 183: 180: 179:Visual system 177: 175: 171: 168: 165: 161: 156: 141: 134: 107: 98: 68: 63: 61:Pronunciation 59: 54: 49: 43: 38: 33: 30: 26: 22: 7898:Spinal nerve 7834: 7830:Diencephalon 7685: 7677: 7669: 7661: 7656:Trompe-l'œil 7654: 7520:Lilac chaser 7490:Gravity hill 7343: 7283: 7237:Ciliary body 7077:Diencephalon 7076: 7061:Parasol cell 7045:Bipolar cell 6947: 6896:Pars plicata 6878:Ciliary body 6746: 6639: 6563:(12): 3487. 6560: 6556:Scholarpedia 6554: 6513: 6506:BMC Genomics 6505: 6476: 6473:Physiol. Rev 6472: 6453: 6428: 6424: 6416: 6392:. Retrieved 6378: 6366:. Retrieved 6362:the original 6352: 6340:. Retrieved 6336:the original 6326: 6301: 6297: 6290: 6239: 6235: 6222: 6210:. Retrieved 6201: 6191: 6179:. Retrieved 6170: 6161: 6149:. Retrieved 6140: 6131: 6119:. Retrieved 6102:(1): 59–61. 6099: 6095: 6085: 6076: 6033: 6029: 6019: 5984: 5981:Gene Therapy 5980: 5934: 5930: 5924: 5889: 5885: 5875: 5848: 5845:Gene Therapy 5844: 5834: 5799: 5795: 5785: 5750: 5746: 5736: 5713:10261/271174 5686: 5682: 5669: 5634: 5630: 5582: 5576: 5528: 5524: 5474: 5470: 5430: 5426: 5420: 5377: 5371: 5361: 5326: 5320: 5310: 5305:, March 2007 5302: 5288: 5253: 5249: 5239: 5212: 5207: 5180: 5174: 5149: 5145: 5138: 5105: 5101: 5095: 5062: 5058: 5051: 5026: 5022: 5016: 4981: 4977: 4967: 4930: 4926: 4916: 4871: 4865: 4830: 4824: 4812:. Retrieved 4808:the original 4800: 4791: 4779:. Retrieved 4773: 4764: 4726: 4719: 4684: 4680: 4670: 4656:cite journal 4627: 4623: 4613: 4587:(1): 92–97. 4584: 4580: 4570: 4558:. Retrieved 4544: 4532:. Retrieved 4528: 4519: 4468: 4464: 4454: 4442:. Retrieved 4438: 4428: 4416:. Retrieved 4406: 4399: 4366: 4362: 4356: 4329: 4323: 4288: 4284: 4249:(1): 83–89. 4246: 4242: 4236: 4195: 4191: 4185: 4152: 4148: 4142: 4107: 4103: 4062:. Retrieved 4052: 4027: 4023: 4017: 4008: 3998: 3989: 3980: 3961: 3957: 3947: 3920: 3884: 3878: 3853: 3849: 3771: 3767: 3740: 3713: 3707: 3682: 3674: 3641: 3638:J Neurogenet 3637: 3631: 3596: 3593:Curr Mol Med 3592: 3582: 3573: 3528: 3524: 3514: 3479: 3475: 3465: 3457:the original 3452: 3443: 3424: 3391: 3387: 3377: 3352: 3346: 3340: 3328:. Retrieved 3314: 3271: 3267: 3257: 3224: 3220: 3194: 3188: 3143: 3139: 3129: 3117:. Retrieved 3113: 3103: 3091:. Retrieved 3079: 3021: 3017: 2990:. Retrieved 2986:the original 2972: 2931: 2925: 2891:Retinal scan 2786:, an online 2773: 2754: 2739: 2721: 2716: 2712: 2706: 2698: 2692:and against 2683: 2678: 2674: 2666: 2662: 2652: 2631: 2630: 2617:angiogenesis 2594: 2561: 2554: 2530:Gene therapy 2528: 2518: 2517: 2488:Intravitreal 2482: 2474: 2433: 2417: 2408: 2400: 2392: 2380: 2334:hypertension 2299:Ignipuncture 2258: 2246: 2243: 2233:to the LGN ( 2231:optic chiasm 2228: 2221: 2218: 2187:spike trains 2173: 2169: 2120: 2112: 2104:ion channels 2100: 2088: 2072: 2045:visual angle 2037: 2022: 2005: 1981:synaptically 1978: 1946:photopigment 1943: 1931: 1916: 1909: 1886: 1879:Retinal scan 1867:pecten oculi 1863: 1854: 1850: 1841: 1832: 1827:macula lutea 1798: 1780: 1774:October 2016 1771: 1760:Please help 1752: 1727:October 2016 1724: 1720:adding to it 1715: 1693: 1686: 1680: 1679:Please help 1676: 1658:Blood supply 1640: 1603: 1587: 1519: 1487: 1454: 1451: 1448: 1447: 1376: 1372:mitochondria 1370:packed with 1341: 1337:Microtubules 1332:Golgi bodies 1317:containing: 1288:Müller cells 1268:cell bodies 1242: 1175: 1174:cell bodies 1057: 1056: 1034: 1027: 1014: 1007: 1002: 974: 969:boundaries? 968: 965: 957: 955:Reflectivity 938: 907: 878: 866: 840:Between the 839: 824: 793: 764: 745: 725:diencephalon 714: 691: 658: 624:bipolar cell 596:Müller cells 586: 523: 519: 515: 508: 503: 493: 474: 460: 458: 432:diencephalon 425: 402: 359: 351:image sensor 306: 301: 288: 280: 278: 232:A15.2.04.002 207: 29: 7916:Sympathetic 7888:Motor nerve 7840:Optic nerve 7788:Spinal cord 7666:(1864 book) 7570:Poggendorff 7545:Oppel-Kundt 7540:Necker cube 7535:Müller-Lyer 7510:Irradiation 7329:Eye disease 7309:Keratocytes 7198:Conjunctiva 7155:Ora serrata 7093:Muller glia 7057:Midget cell 6777:Endothelium 6767:Dua's layer 6121:23 February 5322:Circulation 5256:(11): 892. 4534:23 February 4471:(1): 6445. 4285:Development 3789:10045/74474 3525:J Biol Chem 2723:George Wald 2653:Around 300 2438:is used to 2319:cryotherapy 1956:causes the 1636:homeodomain 1600:Development 1178:Müller cell 1087:optic nerve 1051:Müller cell 975:references 973:Additional 966:anatomical 831:capillaries 785:ora serrata 717:optic nerve 481:fibre-optic 477:glial cells 409:optic nerve 390:entrainment 335:optic nerve 323:vertebrates 197:Identifiers 7945:Categories 7813:Cerebellum 7663:Spectropia 7580:Rubin vase 7530:McCollough 7525:Mach bands 7475:Ehrenstein 7470:Ebbinghaus 7435:Barberpole 7410:Afterimage 7170:of the eye 7143:Optic disc 6901:Pars plana 6772:Descemet's 6752:Epithelium 6425:Vision Res 6212:5 December 6181:5 December 6151:20 October 5380:(63): 63. 3330:5 December 3119:11 January 2917:References 2750:guinea pig 2659:Herophilos 2611:, such as 2456:histologic 2327:vitrectomy 2209:horizontal 2096:inhibitory 2092:excitatory 1970:G-proteins 1900:See also: 1883:Biometrics 1877:See also: 1823:optic disc 1762:improve it 1682:improve it 1622:Drosophila 1164:Horizontal 1049:Formed by 889:horizontal 741:optic disc 735:, and the 706:horizontal 511:homologous 470:cephalopod 466:blind spot 341:to create 283:(from 7908:Autonomic 7825:Forebrain 7798:Hindbrain 7715:Illusions 7687:The dress 7679:Waterfall 7480:Flash lag 7460:Cornsweet 7445:Café wall 7425:Ames room 7403:Illusions 7148:Optic cup 7033:Cone cell 6687:human eye 6516:(1): 50. 6508:(about a 6384:"EyeWire" 6050:0035-9157 6003:0969-7128 5691:CiteSeerX 5199:962009228 5079:0143-5221 4876:CiteSeerX 4493:2045-2322 4444:1 January 4439:Webvision 4369:: 53–93. 4348:745905738 4307:0950-1991 4212:1471-003X 4074:cite news 4024:Optometry 3964:(1): 59. 3903:964379175 3482:: 52–81. 3288:0022-1295 3227:: 49–84. 3114:Webvision 3093:1 January 3080:Webvision 3040:1662-5099 2906:Rhodopsin 2782:launched 2774:In 2012, 2746:bandwidth 2740:A recent 2707:In 1894, 2699:In 1604, 2671:arachnoid 2663:arachnoid 2573:transgene 2492:anti-VEGF 2471:Treatment 2458:quality. 2414:Diagnosis 2179:balanced. 2127:algorithm 1962:isomerise 1688:talk page 1638:factors. 1327:Ribosomes 964:Specific 874:parafovea 846:neuropils 450:Structure 325:and some 313:layer of 48:human eye 7847:Cerebrum 7820:Midbrain 7783:Meninges 7465:Delboeuf 7415:Ambigram 7037:Rod cell 6842:(middle) 6757:Bowman's 6551:"Retina" 6542:15283859 6394:27 March 6388:Archived 6368:27 March 6342:26 March 6318:18329018 6274:17093405 6206:Archived 6175:Archived 6145:Archived 6068:19984605 6011:15454954 5959:30202286 5951:18632300 5908:17923143 5867:12756418 5826:21606598 5777:18774912 5725:Archived 5721:18441371 5661:19854499 5609:18441370 5555:19953081 5501:19583479 5447:15960597 5412:30635610 5353:27682886 5296:Archived 5280:34687632 5231:62034580 5166:15177969 5122:16154379 5087:12149100 5008:23777785 4959:21152417 4927:PLOS Med 4908:14654500 4814:20 April 4781:20 April 4753:Archived 4648:Archived 4605:Archived 4560:22 April 4554:Archived 4511:37081065 4502:10119193 4418:17 April 4412:Archived 4391:21312546 4383:24583621 4315:31399471 4263:15036211 4228:15038502 4220:25096185 4134:23071378 4044:19465337 3870:24755005 3798:29037595 3658:29113536 3623:21091424 3557:22074925 3506:27260426 3432:Archived 3416:20482021 3369:21356325 3324:Archived 3241:29609042 3180:17485670 3088:21413391 3058:33935647 2992:22 April 2841:See also 2756:MacLaren 2675:retiform 2667:retiform 2550:serotype 2452:tomogram 2365:include 2108:synapses 1896:Function 1860:In birds 1478:complex 1286:between 1282:Made of 1205:between 1203:Synapses 1172:amacrine 1136:amacrine 1021:detached 1017:vitreous 852:and the 800:synapses 780:primates 760:Reptiles 628:ganglion 504:Deinopis 461:inverted 366:synapses 327:molluscs 321:of most 142:retinae 7926:Enteric 7875:Somatic 7803:Medulla 7698:Related 7625:Zöllner 7615:White's 7550:Orbison 7515:Jastrow 7289:Choroid 7190:Eyebrow 7134:Foveola 6950:(inner) 6850:Choroid 6698:(outer) 6685:of the 6565:Bibcode 6493:2006220 6445:5862581 6282:4415311 6244:Bibcode 6116:4567856 6059:2201994 5916:9926758 5817:3104779 5768:2940541 5652:4492302 5600:2829748 5546:2839440 5492:2829287 5403:6329813 5382:Bibcode 5344:5219936 5303:Ingenia 5271:7611980 5130:7434103 5043:7155524 4999:3764070 4950:2994677 4857:7705121 4711:7288393 4702:2228620 4644:4428954 4601:5060117 4473:Bibcode 4407:Anatomy 4177:7892602 4157:Bibcode 4149:Science 4125:3504437 4064:7 April 3666:1351539 3614:4120883 3548:3265842 3497:5112118 3396:Bibcode 3297:2214869 3249:5045660 3171:1895942 3148:Bibcode 3049:8081888 2784:EyeWire 2649:History 2592:(OCT). 2540:(RPE), 2286:and/or 1958:retinal 1766:sources 1476:Bruch's 1211:bipolar 1180:nuclei 1168:bipolar 1132:bipolar 1054:endfeet 829:in the 820:choroid 818:or the 655:macular 566:, 1911) 548:bipolar 362:neurons 337:to the 331:focused 317:of the 307:retinas 302:retinae 300:  220:D012160 163:Part of 56:Details 7835:Retina 7650:Op art 7605:Ternus 7585:Sander 7500:Hering 7440:Bezold 7284:Retina 7194:Eyelid 7186:Adnexa 7109:Macula 7089:K cell 7085:M cell 7081:P cell 6957:Layers 6948:Retina 6919:Stroma 6762:Stroma 6747:layers 6742:Limbus 6734:Cornea 6706:Sclera 6646:002291 6540:  6533:512282 6530:  6491:  6460:  6443:  6316:  6280:  6272:  6236:Nature 6114:  6066:  6056:  6048:  6009:  6001:  5957:  5949:  5914:  5906:  5865:  5824:  5814:  5775:  5765:  5719:  5693:  5659:  5649:  5631:Lancet 5607:  5597:  5553:  5543:  5499:  5489:  5445:  5410:  5400:  5351:  5341:  5278:  5268:  5229:  5219:  5213:Retina 5197:  5187:  5164:  5128:  5120:  5085:  5077:  5041:  5006:  4996:  4957:  4947:  4906:  4896:  4878:  4855:  4845:  4745:  4709:  4699:  4642:  4599:  4509:  4499:  4491:  4389:  4381:  4346:  4336:  4313:  4305:  4261:  4226:  4218:  4210:  4175:  4132:  4122:  4042:  3935:  3901:  3891:  3868:  3796:  3720:  3695:  3691:–225. 3664:  3656:  3621:  3611:  3555:  3545:  3504:  3494:  3414:  3367:  3306:235007 3304:  3294:  3286:  3247:  3239:  3178:  3168:  3086:  3056:  3046:  3038:  2938:  2729:, and 2679:retina 2582:fundus 2498:agents 2373:, and 2356:cancer 2274:macula 2123:matrix 1954:photon 1455:(COST) 1279:(ELM) 1261:(ONL) 1241:axons 1076:(NFL) 959:on OCT 908:Using 898:pigeon 881:mosaic 870:mosaic 772:macula 752:pecten 750:, the 731:, the 644:, and 575:cones. 419:whose 382:opsins 355:camera 315:tissue 281:retina 186:Artery 174:System 46:Right 35:Retina 7793:Brain 7620:Wundt 7575:Ponzo 7455:Chubb 7302:Other 7206:Orbit 7124:Fovea 7102:Other 7039:) → ( 7022:Cells 6924:Pupil 6840:tunic 6683:globe 6278:S2CID 6232:(PDF) 5955:S2CID 5912:S2CID 5728:(PDF) 5679:(PDF) 5126:S2CID 4904:S2CID 4853:S2CID 4756:(PDF) 4731:(PDF) 4387:S2CID 4224:S2CID 3764:(PDF) 3662:S2CID 3245:S2CID 2354:is a 2332:Both 2284:cones 2199:pixel 2191:axons 2164:-1/8 2153:-1/8 2142:-1/8 2078:than 2041:fovea 1652:FOXG1 1648:FOXD1 1420:opsin 1235:(HL) 1199:(OPL) 1159:(INL) 1127:(IPL) 1099:(GCL) 1083:axons 1046:(ILM) 811:axons 796:nerve 776:fovea 748:birds 661:Henle 421:axons 405:brain 378:cones 353:in a 287: 285:Latin 267:[ 256:58301 203:Latin 7808:Pons 7495:Grid 7395:list 7242:Lens 7227:Iris 7075:) → 7051:) → 7043:) → 6911:Iris 6538:PMID 6489:PMID 6458:ISBN 6441:PMID 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