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Photoreceptor cell

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panel of genes being expressed. NRL expression leads to the rod fate. NR2E3 further restricts cells to the rod fate by repressing cone genes. RORbeta is needed for both rod and cone development. TRbeta2 mediates the M cone fate. If any of the previously mentioned factors' functions are ablated, the default photoreceptor is a S cone. These events take place at different time periods for different species and include a complex pattern of activities that bring about a spectrum of phenotypes. If these regulatory networks are disrupted,
1868: 746:, and one PDE will cleave many cGMPs. This amplification means that even the absorption of one photon will affect membrane potential and signal to the brain that light is in the visual field. This is the main feature that differentiates rod photoreceptors from cone photoreceptors. Rods are extremely sensitive and have the capacity of registering a single photon of light, unlike cones. On the other hand, cones are known to have very fast kinetics in terms of rate of amplification of phototransduction, unlike rods. 422: 993:
down patients with rare diseases wiping out classic rod and cone photoreceptor function but preserving ganglion cell function. Despite having no rods or cones the patients continued to exhibit circadian photoentrainment, circadian behavioural patterns, melanopsin suppression, and pupil reactions, with peak spectral sensitivities to environmental and experimental light matching that for the melanopsin photopigment. Their brains could also associate vision with light of this frequency.
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rod or M cone generation. L cones are present in primates, however there is not much known for their developmental program due to use of rodents in research. There are five steps to developing photoreceptors: proliferation of multi-potent retinal progenitor cells (RPCs); restriction of competence of RPCs; cell fate specification; photoreceptor gene expression; and lastly axonal growth, synapse formation and outer segment growth.
204: 485: 47: 257: 1707: 223:(see graph). For example, the peak wavelength of the S-cone's spectral sensitivity is approximately 420 nm (nanometers, a measure of wavelength), so it is more likely to absorb a photon at 420 nm than at any other wavelength. Light of a longer wavelength can also produce the same response from an S-cone, but it would have to be brighter to do so. 413: 248: 230:, a photoreceptor's output signal is proportional only to the number of photons absorbed. The photoreceptors can not measure the wavelength of light that it absorbs and therefore does not detect color on its own. Rather, it is the ratios of responses of the three types of cone cells that can estimate wavelength, and therefore enable 726:
the classic (rod or cone) photoreceptor is depolarized in the dark, which means many sodium ions are flowing into the cell. Thus, the random opening or closing of sodium channels will not affect the membrane potential of the cell; only the closing of a large number of channels, through absorption of
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ipRGCs were only definitively detected ipRGCs in humans during landmark experiments in 2007 on rodless, coneless humans. As had been found in other mammals, the identity of the non-rod non-cone photoreceptor in humans was found to be a ganglion cell in the inner retina. The researchers had tracked
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The key events mediating rod versus S cone versus M cone differentiation are induced by several transcription factors, including RORbeta, OTX2, NRL, CRX, NR2E3 and TRbeta2. The S cone fate represents the default photoreceptor program; however, differential transcriptional activity can bring about
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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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Rod and cone photoreceptors are common to almost all vertebrates. The pineal and parapineal glands are photoreceptive in non-mammalian vertebrates, but not in mammals. Birds have photoactive cerebrospinal fluid (CSF)-contacting neurons within the paraventricular organ that respond to light in the
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sight and brightness detection. Classic photoreceptors (rods and cones) also feed into the novel visual system, which may constribute to color constancy. ipRGCs could be instrumental in understanding many diseases including major causes of blindness worldwide like glaucoma, a disease that affects
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signaling maintains progenitor cycling. Photoreceptor precursors come about through inhibition of Notch signaling and increased activity of various factors including achaete-scute homologue 1. OTX2 activity commits cells to the photoreceptor fate. CRX further defines the photoreceptor specific
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The rod and cone photoreceptors signal their absorption of photons via a decrease in the release of the neurotransmitter glutamate to bipolar cells at its axon terminal. Since the photoreceptor is depolarized in the dark, a high amount of glutamate is being released to bipolar cells in the dark.
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In essence, this property allows for one population of bipolar cells that gets excited by light and another population that gets inhibited by it, even though all photoreceptors show the same response to light. This complexity becomes both important and necessary for
502:, the mechanism by which the energy of a photon signals a mechanism in the cell that leads to its electrical polarization. This polarization ultimately leads to either the transmittance or inhibition of a neural signal that will be fed to the brain via the 453:, the area where ganglion cell fibers are collected into the optic nerve and leave the eye. The distribution of cone classes (L, M, S) are also nonhomogenous, with no S-cones in the fovea, and the ratio of L-cones to M-cones differing between individuals. 722:(in this case, light) reduces the cell's response or firing rate, different from most other sensory systems in which a stimulus increases the cell's response or firing rate. This difference has important functional consequences: 571:
A decrease in the intracellular calcium concentration means that less glutamate is released via calcium-induced exocytosis to the bipolar cell (see below). (The decreased calcium level slows the release of the neurotransmitter
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This change in the cell's membrane potential causes voltage-gated calcium channels to close. This leads to a decrease in the influx of calcium ions into the cell and thus the intracellular calcium ion concentration
379:. The function of the photoreceptor cell is to convert the light information of the photon into a form of information communicable to the nervous system and readily usable to the organism: This conversion is called 587:
ATP provided by the inner segment powers the sodium-potassium pump. This pump is necessary to reset the initial state of the outer segment by taking the sodium ions that are entering the cell and pumping them back
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Illustration of the distribution of cone cells in the fovea of an individual with normal color vision (left), and a color blind (protanopic) retina. Note that the center of the fovea holds very few blue-sensitive
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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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a photon, will affect it and signal that light is in the visual field. This system may have less noise relative to sensory transduction schema that increase rate of neural firing in response to stimulus, like
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The net concentration of intracellular cGMP is reduced (due to its conversion to 5' GMP via PDE), resulting in the closure of cyclic nucleotide-gated Na ion channels located in the photoreceptor outer segment
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when not stimulated. This means that glutamate is released continuously when the cell is unstimulated, and stimulus causes release to stop. In the dark, cells have a relatively high concentration of
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Every rod or cone photoreceptor releases the same neurotransmitter, glutamate. However, the effect of glutamate differs in the bipolar cells, depending upon the type of
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Foster, R.G.; Provencio, I.; Hudson, D.; Fiske, S.; Grip, W.; Menaker, M. (1991). "Circadian photoreception in the retinally degenerate mouse (rd/rd)".
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PDE then catalyzes the hydrolysis of cGMP to 5' GMP. This is the second amplification step, where a single PDE hydrolyses about 1000 cGMP molecules.
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expressed in that cell. Humans have three classes of cones (L, M, S) that each differ in spectral sensitivity and 'prefer' photons of different
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Most vertebrate photoreceptors are located in the retina. The distribution of rods and cones (and classes thereof) in the retina is called the
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ganglion cells, and the study of the receptor offered potential as a new avenue to explore in trying to find treatments for blindness.
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has approximately 6 million cones and 120 million rods. At the "center" of the retina (the point directly behind the lens) lies the
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Richardson, T.M. (1969). "Cytoplasmic and ciliary connections between the inner and outer segments of mammalian visual receptors".
1883: 539:, and activates it. This is the first amplification step – each photoactivated opsin triggers activation of about 100 transducins. 3081: 3011: 107: 1599: 1080:"eye, human." Encyclopædia Britannica. Encyclopædia Britannica Ultimate Reference Suite. Chicago: Encyclopædia Britannica, 2010. 3076: 3006: 2621: 1876: 1580: 1895: 1894: 1624: 31: 1875: 1711: 984:
of the receptor is between 460 and 482 nm. However, they may also contribute to a rudimentary visual pathway enabling
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are different in both their morphological organization and their underlying biochemical pathways. This article describes
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results in a hyperpolarization, so this bipolar cell will depolarize to light as less glutamate is released.
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As a result, sodium ions can no longer enter the cell, and the photoreceptor outer segment membrane becomes
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The number and ratio of rods to cones varies among species, dependent on whether an animal is primarily
346: 319: 216: 145: 1899: 961:, a light-sensitive protein. Therefore they constitute a third class of photoreceptors, in addition to 449:. Across the rest of the retina, rods and cones are intermingled. No photoreceptors are found at the 425:
Distribution of rods and cones along a line passing through the fovea and the blind spot of a human eye
368:. In cone cells, there are different types of opsins that combine with retinal to form pigments called 326:(and farthest from the field of view) is the outer segment, the part of the photoreceptor that absorbs 1880: 506:. The steps that apply to the phototransductuion pathway from vertebrate rod/cone photoreceptors are: 3166: 3063: 2993: 2819: 2674: 2389: 1734: 1533: 981: 929: 925: 728: 212: 129: 1898: 421: 171:. The two classic photoreceptor cells are rods and cones, each contributing information used by the 3206: 3051: 3046: 2581: 2514: 2414: 2120: 2071: 2059: 2049: 2004: 1938: 1901: 1852: 1799: 667:
Absorption of a photon will hyperpolarize the photoreceptor and therefore result in the release of
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Unstimulated (in the dark), cyclic-nucleotide gated channels in the outer segment are open because
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and other non-visual responses to light. Melanopsin functionally resembles invertebrate opsins.
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Normal Responses To Non-visual Effects Of Light Retained By Blind Humans Lacking Rods And Cones
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inside the protein from the cis-form to the trans-form, causing the retinal to change shape.
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in the disc membrane of the outer segment absorbs a photon, changing the configuration of a
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The opsin found in the intrinsically photosensitive ganglion cells of the retina is called
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the ipRGCs contribute to non-image-forming functions like circadian rhythms, behavior and
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This results in a series of unstable intermediates, the last of which binds stronger to a
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This article is about cellular photoreceptors. For other types of photoreceptors, see
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there is a lot of amplification in two stages of classic phototransduction: one
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There are currently three known types of photoreceptor cells in mammalian eyes:
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Normalized human photoreceptor absorbances for different wavelengths of light
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The absorption of light leads to an isomeric change in the retinal molecule.
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Stacks of membrane-enclosed disks are unattached to cell membrane directly
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Human Physiology and Mechanisms of Disease by Arthur C. Guyton (1992)
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Phototransduction in rods and cones is somewhat unusual in that the
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Fast response to light, can perceive more rapid changes in stimuli
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Unlike most sensory receptor cells, photoreceptors actually become
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Have less pigment than rods, require more light to detect images
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Have more pigment than cones, so can detect lower light levels
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in other nerve cells is usually −65 mV). This depolarization
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Swaroop, Anand; Douglas Kim; Douglas Forrest (August 2010).
635:) enter the photoreceptor, depolarizing it to about −40 mV ( 1236:"Cilia in the CNS: The quiet organelle claims center stage" 953:, unlike other retinal ganglion cells, are intrinsically 812:
Not very light sensitive; sensitive only to direct light
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photoreceptors are found on the outermost layer of the
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Kandel, E. R.; Schwartz, J.H.; Jessell, T.M. (2000).
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absence of input from the eyes or neurotransmitters.
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About 120 million rods distributed around the retina
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Intrinsically photosensitive retinal ganglion cells
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intrinsically photosensitive retinal ganglion cells
94: 81: 69: 64: 39: 942:Intrinsically photosensitive retinal ganglion cell 211:Each photoreceptor absorbs light according to its 880:About 6 million cones distributed in each retina 183:(dim conditions) whereas cones primarily mediate 1384:(4th ed.). New York: McGraw-Hill. pp.  853:Slow response to light, stimuli added over time 498:The path of a visual signal is described by the 1871: 1649:Campbell, Neil A. & Reece, Jane B. (2002). 1583:. New Scientist, 26 December 2007, issue 2635. 834:High visual acuity; better spatial resolution 2451: 1742: 1655:. San Francisco: Benjamin Cummings. pp.  1297: 1295: 1293: 1291: 765:Comparison of human rod and cone cells, from 8: 1133:Bowmaker J.K. & Dartnall H.J.A. (1980). 3130: 2850: 2843: 2654: 2549: 2479: 2458: 2444: 2436: 2249: 2029: 1921: 1777: 1749: 1735: 1727: 364:. In rod cells, these together are called 265:Anatomy of rods and cones varies slightly. 215:(absorptance), which is determined by the 45: 1553: 1491: 1434: 1259: 1158: 777: 420: 411: 1308:. New York: Worth Publishers. pp.  1073: 663:, via which they project to the brain. 330:. Outer segments are actually modified 1591: 1589: 1575: 1573: 464:. Certain owls, such as the nocturnal 111: 36: 1581:Blind people 'see' sunrise and sunset 932:or other visual deficits may result. 872:Disks are attached to outer membrane 800:(vision under high light conditions) 175:to form an image of the environment, 7: 793:(vision under low light conditions) 607:cyclic guanosine 3'-5' monophosphate 601:when stimulated; and conversely are 152:, triggering a change in the cell's 1337:(7 ed.). Thomson and Wadswoth. 1092:Journal of Comparative Physiology A 542:Each transducin then activates the 1722:Neuroscience Information Framework 25: 1355:. World Owl Trust. Archived from 949:(ipRGCs) are a subset (≈1–3%) of 751:Difference between rods and cones 576:, which excites the postsynaptic 1705: 1234:Louvi, A.; Grove, E. A. (2011). 742:will activate many molecules of 643:is often known as dark current. 255: 246: 108:Anatomical terms of neuroanatomy 2622:Oligodendrocyte progenitor cell 1718:NIF Search – Photoreceptor Cell 809:; sensitive to scattered light 334:that contain disks filled with 1520:Zaidi FH, et al. (2007). 1466:Do MT, Yau KW (October 2010). 1151:10.1113/jphysiol.1980.sp013097 675:terminal to the bipolar cell. 32:Photoreceptor (disambiguation) 1: 686:. When glutamate binds to an 2395:Optical coherence tomography 2149:Photosensitive ganglion cell 1623:. Birds News. Archived from 1381:Principles of Neural Science 1333:Goldstein, E. Bruce (2007). 1302:Schacter, Daniel L. (2011). 1252:10.1016/j.neuron.2011.03.002 1213:10.1016/0042-6989(69)90010-8 1048:Photosensitive ganglion cell 936:Ganglion cell photoreceptors 772:Principles of Neural Science 302:, which contains the cell's 27:Type of neuroepithelial cell 2732:Postganglionic nerve fibers 2145:Giant retina ganglion cells 1934:Capillary lamina of choroid 1415:Nature Reviews Neuroscience 1004:Invertebrate photoreceptors 286:terminal, which releases a 3303: 2727:Preganglionic nerve fibers 2089:Retinal pigment epithelium 2079:External limiting membrane 1484:10.1152/physrev.00013.2010 1033:G protein-coupled receptor 939: 754: 491: 405: 322:. Finally, closest to the 29: 3237:Olfactory receptor neuron 2901:Neurofibril/neurofilament 1865: 1546:10.1016/j.cub.2007.11.034 1305:Psychology Second Edition 901:Confer achromatic vision 500:phototransduction cascade 179:. Rods primarily mediate 142:electromagnetic radiation 128:is a specialized type of 106: 44: 1335:Sensation and Perception 1028:Visual phototransduction 997:Non-human photoreceptors 615:electrochemical gradient 494:Visual phototransduction 228:principle of univariance 138:visual phototransduction 51:Functional parts of the 2084:Layer of rods and cones 2040:Inner limiting membrane 1674:Freeman, Scott (2002). 957:due to the presence of 611:cGMP-gated ion channels 512:Vertebrate visual opsin 226:In accordance with the 3184:Neuromuscular junction 3047:III or Aδ or fast pain 1906: 978:pupillary light reflex 951:retinal ganglion cells 489: 426: 418: 298:. Farther back is the 217:photoreceptor proteins 208: 146:photoreceptor proteins 2400:Eye care professional 2205:Foveal avascular zone 2067:Outer plexiform layer 2055:Inner plexiform layer 2010:Iris sphincter muscle 1903: 1472:Physiological Reviews 1349:"Owl Eye Information" 1283:Foundations of Vision 1006:in organisms such as 692:metabotropic receptor 487: 424: 415: 347:photoreceptor protein 320:sodium-potassium pump 206: 3202:Meissner's corpuscle 3167:Postsynaptic density 3064:Efferent nerve fiber 3052:IV or C or slow pain 2994:Afferent nerve fiber 2820:Satellite glial cell 2420:Physiological Optics 2390:Ocular immune system 2129:Retina ganglion cell 1714:at Wikimedia Commons 1595:Medical News Today. 982:spectral sensitivity 930:macular degeneration 926:retinitis pigmentosa 904:Confer color vision 651:The photoreceptors ( 609:(cGMP), which opens 213:spectral sensitivity 130:neuroepithelial cell 3287:Photoreceptor cells 3207:Merkel nerve ending 2244:Anatomical regions 2105:Photoreceptor cells 2072:Outer nuclear layer 2060:Inner nuclear layer 2050:Ganglion cell layer 2005:Iris dilator muscle 1800:Trabecular meshwork 1712:Photoreceptor cells 1606:. 14 December 2007. 1538:2007CBio...17.2122Z 1359:on 16 February 2018 688:ionotropic receptor 381:signal transduction 148:in the cell absorb 136:that is capable of 3242:Photoreceptor cell 3212:Pacinian corpuscle 3143:Electrical synapse 3097:Lower motor neuron 3092:Upper motor neuron 2813:Internodal segment 2753:Connective tissues 2723:Autonomic ganglion 1907: 1677:Biological Science 1602:2009-02-06 at the 1285:, Brian A. Wandell 1104:10.1007/BF00198171 896:pigment in humans 831:Low visual acuity 490: 427: 419: 209: 154:membrane potential 126:photoreceptor cell 101:85613 86740, 85613 40:Photoreceptor cell 3277:Human eye anatomy 3264: 3263: 3260: 3259: 3227:Free nerve ending 3194:Sensory receptors 3122: 3121: 3037:Ib or Golgi or Aα 2945: 2944: 2828: 2827: 2705:Ramus communicans 2644: 2643: 2640: 2639: 2510:Commissural fiber 2505:Association fiber 2500:Projection fibers 2433: 2432: 2425:Visual perception 2373: 2372: 2340:Posterior segment 2308:Posterior chamber 2239: 2238: 2141:Bistratified cell 2045:Nerve fiber layer 2018: 2017: 1962:Ciliary processes 1863: 1862: 1710:Media related to 1319:978-1-4292-3719-2 908: 907: 682:imbedded in that 671:glutamate at the 637:resting potential 593:Hyperpolarization 548:phosphodiesterase 392:circadian rhythms 318:(energy) for the 122: 121: 117: 16:(Redirected from 3294: 3157:Synaptic vesicle 3152:Chemical synapse 3131: 2851: 2844: 2655: 2550: 2480: 2460: 2453: 2446: 2437: 2410:Refractive error 2348:Vitreous chamber 2293:Anterior chamber 2254:Anterior segment 2250: 2030: 1939:Bruch's membrane 1922: 1914:Uvea / vascular 1870: 1790:Episcleral layer 1778: 1751: 1744: 1737: 1728: 1709: 1695: 1670: 1637: 1636: 1634: 1632: 1613: 1607: 1593: 1584: 1577: 1568: 1567: 1557: 1517: 1506: 1505: 1495: 1463: 1457: 1456: 1438: 1406: 1400: 1399: 1375: 1369: 1368: 1366: 1364: 1345: 1339: 1338: 1330: 1324: 1323: 1299: 1286: 1280: 1274: 1273: 1263: 1246:(6): 1046–1060. 1231: 1225: 1224: 1196: 1190: 1179: 1173: 1172: 1162: 1130: 1124: 1123: 1087: 1081: 1078: 1018:photoreceptors. 845:Concentrated in 778: 582:horizontal cells 396:pupillary reflex 358:molecule called 288:neurotransmitter 259: 250: 199:Photosensitivity 193:pupillary reflex 189:circadian rhythm 114:edit on Wikidata 49: 37: 21: 3302: 3301: 3297: 3296: 3295: 3293: 3292: 3291: 3267: 3266: 3265: 3256: 3188: 3118: 3067: 3058: 3042:II or Aβ and Aγ 2997: 2988: 2941: 2931:Apical dendrite 2926:Dendritic spine 2905: 2867: 2837: 2824: 2808:Node of Ranvier 2803:Myelin incisure 2775: 2747: 2636: 2627:Oligodendrocyte 2610:Ependymal cells 2591: 2541: 2469: 2464: 2434: 2429: 2369: 2334: 2323:Capsule of lens 2278:Lacrimal system 2245: 2235: 2195:Parafoveal area 2190:Perifoveal area 2173: 2117:Horizontal cell 2093: 2014: 1981: 1948: 1944:Sattler's layer 1915: 1908: 1902: 1859: 1804: 1795:Schlemm's canal 1773: 1765: 1757:Anatomy of the 1755: 1702: 1692: 1673: 1667: 1648: 1645: 1640: 1630: 1628: 1615: 1614: 1610: 1604:Wayback Machine 1594: 1587: 1578: 1571: 1526:Current Biology 1519: 1518: 1509: 1465: 1464: 1460: 1427:10.1038/nrn2880 1408: 1407: 1403: 1396: 1377: 1376: 1372: 1362: 1360: 1347: 1346: 1342: 1332: 1331: 1327: 1320: 1301: 1300: 1289: 1281: 1277: 1233: 1232: 1228: 1201:Vision Research 1198: 1197: 1193: 1180: 1176: 1132: 1131: 1127: 1089: 1088: 1084: 1079: 1075: 1071: 1053:Horizontal cell 1024: 999: 944: 938: 913: 892:Three types of 839:Not present in 825:legal blindness 819:night blindness 798:photopic vision 791:scotopic vision 763: 755:Main articles: 753: 716: 700:detecting color 684:cell's membrane 649: 595: 496: 482: 410: 404: 345:The membranous 340:sodium channels 269: 268: 267: 266: 262: 261: 260: 252: 251: 240: 201: 185:photopic vision 181:scotopic vision 118: 60: 35: 28: 23: 22: 15: 12: 11: 5: 3300: 3298: 3290: 3289: 3284: 3279: 3269: 3268: 3262: 3261: 3258: 3257: 3255: 3254: 3252:Taste receptor 3249: 3244: 3239: 3234: 3229: 3224: 3222:Muscle spindle 3219: 3217:Ruffini ending 3214: 3209: 3204: 3198: 3196: 3190: 3189: 3187: 3186: 3181: 3179:Ribbon synapse 3176: 3171: 3170: 3169: 3164: 3159: 3149: 3139: 3137: 3128: 3124: 3123: 3120: 3119: 3117: 3116: 3115: 3114: 3109: 3104: 3094: 3089: 3084: 3079: 3073: 3071: 3060: 3059: 3057: 3056: 3055: 3054: 3049: 3044: 3039: 3034: 3024: 3019: 3014: 3009: 3003: 3001: 2999:Sensory neuron 2990: 2989: 2987: 2986: 2985: 2984: 2974: 2969: 2967:Pseudounipolar 2964: 2959: 2953: 2951: 2947: 2946: 2943: 2942: 2940: 2939: 2938: 2937: 2935:Basal dendrite 2928: 2923: 2915: 2913: 2907: 2906: 2904: 2903: 2898: 2893: 2888: 2886:Axon terminals 2883: 2877: 2875: 2869: 2868: 2866: 2865: 2859: 2857: 2848: 2841: 2830: 2829: 2826: 2825: 2823: 2822: 2817: 2816: 2815: 2810: 2805: 2800: 2785: 2783: 2777: 2776: 2774: 2773: 2768: 2763: 2757: 2755: 2749: 2748: 2746: 2745: 2740: 2738:Nerve fascicle 2735: 2729: 2720: 2719: 2718: 2713: 2701: 2700: 2699: 2694: 2684: 2683: 2682: 2677: 2672: 2661: 2659: 2652: 2646: 2645: 2642: 2641: 2638: 2637: 2635: 2634: 2629: 2624: 2619: 2618: 2617: 2607: 2601: 2599: 2593: 2592: 2590: 2589: 2584: 2579: 2574: 2569: 2564: 2558: 2556: 2547: 2543: 2542: 2540: 2539: 2534: 2529: 2528: 2527: 2522: 2517: 2512: 2507: 2502: 2492: 2486: 2484: 2477: 2471: 2470: 2467:Nervous tissue 2465: 2463: 2462: 2455: 2448: 2440: 2431: 2430: 2428: 2427: 2422: 2417: 2412: 2407: 2402: 2397: 2392: 2387: 2381: 2379: 2375: 2374: 2371: 2370: 2368: 2367: 2362: 2357: 2356: 2355: 2344: 2342: 2336: 2335: 2333: 2332: 2331: 2330: 2328:Zonule of Zinn 2325: 2315: 2310: 2305: 2300: 2298:Aqueous humour 2295: 2290: 2285: 2258: 2256: 2247: 2241: 2240: 2237: 2236: 2234: 2233: 2228: 2227: 2226: 2216: 2215: 2214: 2213: 2212: 2207: 2197: 2192: 2181: 2179: 2175: 2174: 2172: 2171: 2101: 2099: 2095: 2094: 2092: 2091: 2086: 2081: 2075: 2074: 2069: 2063: 2062: 2057: 2052: 2047: 2042: 2036: 2034: 2027: 2020: 2019: 2016: 2015: 2013: 2012: 2007: 2002: 1997: 1991: 1989: 1983: 1982: 1980: 1979: 1974: 1969: 1967:Ciliary muscle 1964: 1958: 1956: 1950: 1949: 1947: 1946: 1941: 1936: 1930: 1928: 1919: 1910: 1909: 1866: 1864: 1861: 1860: 1858: 1857: 1856: 1855: 1850: 1845: 1840: 1835: 1830: 1820: 1814: 1812: 1806: 1805: 1803: 1802: 1797: 1792: 1786: 1784: 1775: 1767: 1766: 1756: 1754: 1753: 1746: 1739: 1731: 1725: 1724: 1715: 1701: 1700:External links 1698: 1697: 1696: 1690: 1671: 1665: 1644: 1641: 1639: 1638: 1627:on 2 July 2017 1608: 1585: 1569: 1532:(24): 2122–8. 1507: 1478:(4): 1547–81. 1458: 1421:(8): 563–576. 1401: 1394: 1370: 1340: 1325: 1318: 1287: 1275: 1226: 1207:(7): 727–731. 1191: 1174: 1125: 1082: 1072: 1070: 1067: 1066: 1065: 1060: 1055: 1050: 1045: 1043:Photosensitive 1040: 1038:Sensory system 1035: 1030: 1023: 1020: 998: 995: 955:photosensitive 940:Main article: 937: 934: 912: 909: 906: 905: 902: 898: 897: 894:photosensitive 890: 887:photosensitive 882: 881: 878: 874: 873: 870: 866: 865: 862: 858: 857: 854: 850: 849: 843: 836: 835: 832: 828: 827: 821: 814: 813: 810: 802: 801: 794: 786: 785: 782: 752: 749: 748: 747: 736: 715: 712: 648: 645: 599:hyperpolarized 594: 591: 590: 589: 585: 569: 565: 562:hyperpolarized 558: 554: 551: 546:cGMP-specific 540: 525: 492:Main article: 481: 478: 431:retinal mosaic 408:Retinal mosaic 406:Main article: 403: 402:Retinal mosaic 400: 264: 263: 254: 253: 245: 244: 243: 242: 241: 239: 236: 200: 197: 120: 119: 110: 104: 103: 98: 92: 91: 86: 79: 78: 73: 67: 66: 62: 61: 50: 42: 41: 26: 24: 18:Photoreception 14: 13: 10: 9: 6: 4: 3: 2: 3299: 3288: 3285: 3283: 3280: 3278: 3275: 3274: 3272: 3253: 3250: 3248: 3245: 3243: 3240: 3238: 3235: 3233: 3230: 3228: 3225: 3223: 3220: 3218: 3215: 3213: 3210: 3208: 3205: 3203: 3200: 3199: 3197: 3195: 3191: 3185: 3182: 3180: 3177: 3175: 3172: 3168: 3165: 3163: 3160: 3158: 3155: 3154: 3153: 3150: 3148: 3144: 3141: 3140: 3138: 3136: 3132: 3129: 3125: 3113: 3112:γ motorneuron 3110: 3108: 3107:β motorneuron 3105: 3103: 3102:α motorneuron 3100: 3099: 3098: 3095: 3093: 3090: 3088: 3085: 3083: 3080: 3078: 3075: 3074: 3072: 3070: 3065: 3061: 3053: 3050: 3048: 3045: 3043: 3040: 3038: 3035: 3033: 3030: 3029: 3028: 3025: 3023: 3020: 3018: 3015: 3013: 3010: 3008: 3005: 3004: 3002: 3000: 2995: 2991: 2983: 2980: 2979: 2978: 2975: 2973: 2970: 2968: 2965: 2963: 2960: 2958: 2955: 2954: 2952: 2948: 2936: 2932: 2929: 2927: 2924: 2922: 2919: 2918: 2917: 2916: 2914: 2912: 2908: 2902: 2899: 2897: 2894: 2892: 2889: 2887: 2884: 2882: 2879: 2878: 2876: 2874: 2870: 2864: 2861: 2860: 2858: 2856: 2852: 2849: 2845: 2842: 2840: 2835: 2831: 2821: 2818: 2814: 2811: 2809: 2806: 2804: 2801: 2799: 2796: 2795: 2794: 2790: 2787: 2786: 2784: 2782: 2778: 2772: 2769: 2767: 2764: 2762: 2759: 2758: 2756: 2754: 2750: 2744: 2741: 2739: 2736: 2733: 2730: 2728: 2724: 2721: 2717: 2714: 2712: 2709: 2708: 2707: 2706: 2702: 2698: 2695: 2693: 2690: 2689: 2688: 2685: 2681: 2678: 2676: 2673: 2671: 2668: 2667: 2666: 2663: 2662: 2660: 2656: 2653: 2651: 2647: 2633: 2630: 2628: 2625: 2623: 2620: 2616: 2613: 2612: 2611: 2608: 2606: 2603: 2602: 2600: 2598: 2594: 2588: 2585: 2583: 2580: 2578: 2575: 2573: 2570: 2568: 2565: 2563: 2560: 2559: 2557: 2555: 2551: 2548: 2544: 2538: 2535: 2533: 2530: 2526: 2523: 2521: 2518: 2516: 2513: 2511: 2508: 2506: 2503: 2501: 2498: 2497: 2496: 2493: 2491: 2488: 2487: 2485: 2481: 2478: 2476: 2472: 2468: 2461: 2456: 2454: 2449: 2447: 2442: 2441: 2438: 2426: 2423: 2421: 2418: 2416: 2415:Accommodation 2413: 2411: 2408: 2406: 2403: 2401: 2398: 2396: 2393: 2391: 2388: 2386: 2383: 2382: 2380: 2376: 2366: 2363: 2361: 2358: 2354: 2353:Vitreous body 2351: 2350: 2349: 2346: 2345: 2343: 2341: 2337: 2329: 2326: 2324: 2321: 2320: 2319: 2316: 2314: 2311: 2309: 2306: 2304: 2301: 2299: 2296: 2294: 2291: 2289: 2288:Fibrous tunic 2286: 2283: 2279: 2275: 2271: 2267: 2263: 2260: 2259: 2257: 2255: 2251: 2248: 2242: 2232: 2229: 2225: 2222: 2221: 2220: 2217: 2211: 2208: 2206: 2203: 2202: 2201: 2198: 2196: 2193: 2191: 2188: 2187: 2186: 2183: 2182: 2180: 2176: 2170: 2166: 2162: 2158: 2154: 2150: 2146: 2142: 2138: 2134: 2130: 2126: 2125:Amacrine cell 2122: 2118: 2114: 2110: 2106: 2103: 2102: 2100: 2096: 2090: 2087: 2085: 2082: 2080: 2077: 2076: 2073: 2070: 2068: 2065: 2064: 2061: 2058: 2056: 2053: 2051: 2048: 2046: 2043: 2041: 2038: 2037: 2035: 2031: 2028: 2025: 2021: 2011: 2008: 2006: 2003: 2001: 1998: 1996: 1993: 1992: 1990: 1988: 1984: 1978: 1975: 1973: 1970: 1968: 1965: 1963: 1960: 1959: 1957: 1955: 1951: 1945: 1942: 1940: 1937: 1935: 1932: 1931: 1929: 1927: 1923: 1920: 1917: 1911: 1854: 1851: 1849: 1846: 1844: 1841: 1839: 1836: 1834: 1831: 1829: 1826: 1825: 1824: 1821: 1819: 1816: 1815: 1813: 1811: 1807: 1801: 1798: 1796: 1793: 1791: 1788: 1787: 1785: 1783: 1779: 1776: 1772: 1771:Fibrous tunic 1768: 1764: 1760: 1752: 1747: 1745: 1740: 1738: 1733: 1732: 1729: 1723: 1719: 1716: 1713: 1708: 1704: 1703: 1699: 1693: 1691:0-13-140941-7 1687: 1683: 1679: 1678: 1672: 1668: 1666:0-8053-6624-5 1662: 1658: 1654: 1653: 1647: 1646: 1642: 1626: 1622: 1621:birdsnews.com 1618: 1612: 1609: 1605: 1601: 1598: 1592: 1590: 1586: 1582: 1576: 1574: 1570: 1565: 1561: 1556: 1551: 1547: 1543: 1539: 1535: 1531: 1527: 1523: 1516: 1514: 1512: 1508: 1503: 1499: 1494: 1489: 1485: 1481: 1477: 1473: 1469: 1462: 1459: 1454: 1450: 1446: 1442: 1437: 1432: 1428: 1424: 1420: 1416: 1412: 1405: 1402: 1397: 1395:0-8385-7701-6 1391: 1387: 1383: 1382: 1374: 1371: 1358: 1354: 1350: 1344: 1341: 1336: 1329: 1326: 1321: 1315: 1311: 1307: 1306: 1298: 1296: 1294: 1292: 1288: 1284: 1279: 1276: 1271: 1267: 1262: 1257: 1253: 1249: 1245: 1241: 1237: 1230: 1227: 1222: 1218: 1214: 1210: 1206: 1202: 1195: 1192: 1188: 1187:0-7216-3299-8 1184: 1178: 1175: 1170: 1166: 1161: 1156: 1152: 1148: 1144: 1140: 1136: 1129: 1126: 1121: 1117: 1113: 1109: 1105: 1101: 1097: 1093: 1086: 1083: 1077: 1074: 1068: 1064: 1063:Amacrine cell 1061: 1059: 1056: 1054: 1051: 1049: 1046: 1044: 1041: 1039: 1036: 1034: 1031: 1029: 1026: 1025: 1021: 1019: 1017: 1013: 1009: 1005: 996: 994: 990: 987: 983: 979: 975: 970: 968: 964: 960: 956: 952: 948: 943: 935: 933: 931: 927: 922: 917: 910: 903: 900: 899: 895: 891: 888: 884: 883: 879: 876: 875: 871: 868: 867: 863: 860: 859: 855: 852: 851: 848: 844: 842: 838: 837: 833: 830: 829: 826: 822: 820: 816: 815: 811: 808: 804: 803: 799: 795: 792: 788: 787: 783: 780: 779: 776: 774: 773: 768: 762: 758: 750: 745: 741: 737: 734: 730: 725: 724: 723: 721: 713: 711: 709: 705: 701: 695: 693: 689: 685: 681: 676: 674: 670: 664: 662: 658: 654: 647:Bipolar cells 646: 644: 642: 638: 634: 631: 627: 622: 620: 616: 612: 608: 604: 600: 592: 586: 583: 579: 578:bipolar cells 575: 570: 566: 563: 559: 555: 552: 549: 545: 541: 538: 534: 530: 526: 523: 520: 517: 513: 509: 508: 507: 505: 501: 495: 486: 479: 477: 475: 474:pentachromats 471: 467: 463: 459: 454: 452: 448: 444: 443:visual acuity 440: 436: 433:. Each human 432: 423: 414: 409: 401: 399: 397: 393: 389: 384: 382: 378: 375: 371: 367: 363: 362: 357: 353: 352: 348: 343: 341: 337: 333: 329: 325: 321: 317: 313: 309: 305: 301: 297: 296:bipolar cells 293: 289: 285: 281: 277: 273: 258: 249: 237: 235: 233: 229: 224: 222: 218: 214: 205: 198: 196: 194: 190: 186: 182: 178: 174: 173:visual system 170: 166: 162: 157: 155: 151: 147: 143: 139: 135: 132:found in the 131: 127: 115: 109: 105: 102: 99: 97: 93: 90: 87: 84: 80: 77: 74: 72: 68: 63: 58: 54: 48: 43: 38: 33: 19: 3241: 3147:Gap junction 3069:Motor neuron 2863:Axon hillock 2839:nerve fibers 2793:Schwann cell 2703: 2686: 2664: 2582:Medium spiny 2495:White matter 2483:Tissue Types 2419: 2313:Ciliary body 2153:Diencephalon 2152: 2137:Parasol cell 2121:Bipolar cell 2104: 1972:Pars plicata 1954:Ciliary body 1822: 1676: 1651: 1643:Bibliography 1629:. Retrieved 1625:the original 1620: 1611: 1529: 1525: 1475: 1471: 1461: 1418: 1414: 1404: 1380: 1373: 1361:. Retrieved 1357:the original 1352: 1343: 1334: 1328: 1304: 1278: 1243: 1239: 1229: 1204: 1200: 1194: 1177: 1142: 1138: 1128: 1098:(1): 39–50. 1095: 1091: 1085: 1076: 1058:Bipolar cell 1000: 991: 971: 945: 918: 914: 885:One type of 823:Loss causes 817:Loss causes 770: 764: 717: 696: 677: 668: 665: 656: 652: 650: 623: 596: 497: 470:monochromats 455: 430: 428: 385: 359: 349: 344: 312:mitochondria 270: 232:color vision 225: 210: 158: 125: 123: 89:sao226523927 3162:Active zone 3127:Termination 2977:Interneuron 2881:Telodendron 2789:Myelination 2771:Endoneurium 2766:Perineurium 2587:Interneuron 2577:Von Economo 2525:Decussation 2520:Nerve tract 2490:Grey matter 2405:Eye disease 2385:Keratocytes 2274:Conjunctiva 2231:Ora serrata 2169:Muller glia 2133:Midget cell 1853:Endothelium 1843:Dua's layer 1579:Coghlan A. 1145:: 501–511. 911:Development 805:Very light 767:Eric Kandel 673:presynaptic 661:optic nerve 603:depolarized 519:Schiff base 504:optic nerve 445:or highest 354:contains a 221:wavelengths 65:Identifiers 3271:Categories 3232:Nociceptor 2972:Multipolar 2921:Nissl body 2798:Neurilemma 2761:Epineurium 2546:Cell Types 2246:of the eye 2219:Optic disc 1977:Pars plana 1848:Descemet's 1828:Epithelium 1139:J. Physiol 1069:References 967:cone cells 959:melanopsin 769:et al. in 744:transducin 714:Advantages 626:cyclic GMP 537:transducin 451:blind spot 447:resolution 388:melanopsin 370:photopsins 304:organelles 3282:Histology 3247:Hair cell 2781:Neuroglia 2743:Funiculus 2632:Microglia 2605:Astrocyte 2562:Pyramidal 2515:Lemniscus 2224:Optic cup 2109:Cone cell 1763:human eye 1657:1064–1067 986:conscious 807:sensitive 796:Used for 789:Used for 757:Cone cell 733:olfaction 619:glutamate 574:glutamate 557:membrane. 535:, called 529:G protein 480:Signaling 466:tawny owl 462:nocturnal 374:transduce 366:rhodopsin 300:cell body 292:glutamate 238:Histology 3032:Ia or Aα 2962:Unipolar 2911:Dendrite 2896:Axolemma 2891:Axoplasm 2675:Ganglion 2615:Tanycyte 2567:Purkinje 2554:Neuronal 2537:Meninges 2532:Neuropil 2113:Rod cell 1918:(middle) 1833:Bowman's 1720:via the 1600:Archived 1564:18082405 1502:20959623 1445:20648062 1436:11346175 1353:owls.org 1270:21435552 1022:See also 1012:molluscs 889:pigment 761:Rod cell 720:stimulus 704:contrast 680:receptor 533:membrane 522:cofactor 310:full of 191:and the 83:NeuroLex 3174:Autapse 3135:Synapse 2982:Renshaw 2957:Bipolar 2834:Neurons 2687:Ventral 2658:General 2572:Granule 2365:Choroid 2266:Eyebrow 2210:Foveola 2026:(inner) 1926:Choroid 1774:(outer) 1761:of the 1682:835–837 1652:Biology 1631:20 July 1555:2151130 1534:Bibcode 1493:4374737 1453:6034699 1386:507–513 1310:136–137 1261:3070490 1221:4979023 1169:7359434 1160:1279132 1120:1124159 1112:1941717 1008:insects 980:. Peak 740:pigment 710:, etc. 641:current 531:in the 516:retinal 458:diurnal 361:retinal 356:pigment 290:called 150:photons 76:D010786 3027:fibers 2665:Dorsal 2360:Retina 2270:Eyelid 2262:Adnexa 2185:Macula 2165:K cell 2161:M cell 2157:P cell 2033:Layers 2024:Retina 1995:Stroma 1838:Stroma 1823:layers 1818:Limbus 1810:Cornea 1782:Sclera 1688:  1663:  1562:  1552:  1500:  1490:  1451:  1443:  1433:  1392:  1316:  1268:  1258:  1240:Neuron 1219:  1189:p. 373 1185:  1167:  1157:  1118:  1110:  974:humans 919:Early 784:Cones 630:sodium 568:falls. 550:(PDE). 544:enzyme 435:retina 417:cones. 280:retina 167:, and 134:retina 2950:Types 2847:Parts 2716:White 2697:Ramus 2680:Ramus 2597:Glial 2378:Other 2282:Orbit 2200:Fovea 2178:Other 2115:) → ( 2098:Cells 2000:Pupil 1916:tunic 1759:globe 1449:S2CID 1363:1 May 1116:S2CID 1016:human 921:Notch 847:fovea 841:fovea 781:Rods 729:touch 708:edges 657:cones 439:fovea 377:color 351:opsin 336:opsin 332:cilia 328:light 324:brain 177:sight 165:cones 112:[ 57:cones 2873:Axon 2855:Soma 2711:Gray 2692:Root 2670:Root 2318:Lens 2303:Iris 2151:) → 2127:) → 2119:) → 1987:Iris 1686:ISBN 1661:ISBN 1633:2017 1560:PMID 1498:PMID 1441:PMID 1390:ISBN 1365:2017 1314:ISBN 1266:PMID 1217:PMID 1183:ISBN 1165:PMID 1108:PMID 1010:and 965:and 759:and 731:and 669:less 655:and 653:rods 633:ions 588:out. 580:and 510:The 308:cell 284:axon 276:cone 274:and 161:rods 71:MeSH 55:and 53:rods 3087:SVE 3082:GVE 3077:GSE 3022:SVA 3017:SSA 3012:GVA 3007:GSA 2650:PNS 2475:CNS 2123:→ ( 1550:PMC 1542:doi 1488:PMC 1480:doi 1431:PMC 1423:doi 1256:PMC 1248:doi 1209:doi 1155:PMC 1147:doi 1143:298 1100:doi 1096:169 972:In 963:rod 472:to 460:or 316:ATP 294:to 272:Rod 96:FMA 3273:: 2791:: 2280:, 2276:, 2272:, 2268:, 2167:, 2163:, 2159:, 2155:: 2147:, 2143:, 2139:, 2135:, 2111:, 1684:. 1659:. 1619:. 1588:^ 1572:^ 1558:. 1548:. 1540:. 1530:17 1528:. 1524:. 1510:^ 1496:. 1486:. 1476:90 1474:. 1470:. 1447:. 1439:. 1429:. 1419:11 1417:. 1413:. 1388:. 1351:. 1312:. 1290:^ 1264:. 1254:. 1244:69 1242:. 1238:. 1215:. 1203:. 1163:. 1153:. 1141:. 1137:. 1114:. 1106:. 1094:. 969:. 928:, 775:. 706:, 702:, 621:. 584:.) 476:. 394:, 383:. 342:. 234:. 195:. 163:, 156:. 124:A 85:ID 3145:/ 3066:/ 2996:/ 2933:/ 2836:/ 2734:) 2725:( 2459:e 2452:t 2445:v 2284:) 2264:( 2131:( 2107:( 1750:e 1743:t 1736:v 1694:. 1669:. 1635:. 1566:. 1544:: 1536:: 1504:. 1482:: 1455:. 1425:: 1398:. 1367:. 1322:. 1272:. 1250:: 1223:. 1211:: 1205:9 1171:. 1149:: 1122:. 1102:: 735:. 116:] 34:. 20:)

Index

Photoreception
Photoreceptor (disambiguation)

rods
cones
MeSH
D010786
NeuroLex
sao226523927
FMA
85613 86740, 85613
Anatomical terms of neuroanatomy
edit on Wikidata
neuroepithelial cell
retina
visual phototransduction
electromagnetic radiation
photoreceptor proteins
photons
membrane potential
rods
cones
intrinsically photosensitive retinal ganglion cells
visual system
sight
scotopic vision
photopic vision
circadian rhythm
pupillary reflex

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