Show The rods are most sensitive to light and dark changes, shape and movement and contain only one type of light-sensitive pigment. Rods are not good for color vision. In a dim room, however, we use mainly our rods, but we are "color blind." Rods are more numerous than cones in the periphery of the retina. Next time you want to see a dim star at night, try to look at it with your peripheral vision and use your ROD VISION to see the dim star. There are about 120 million rods in the human retina. "Your vision is best when light falls on the fovea." The "g" in "light" will be clear, but words and letters on either side of the "g" will not be clear. One part of the retina does NOT contain any photoreceptors. This is our "blind spot." Therefore any image that falls on this region will NOT be seen. It is in this region that the optic nerves come together and exit the eye on their way to the brain. To find your blind spot, look at the image below or draw it on a piece of paper: Close your left eye. Hold the image (or place your head from the computer monitor) about 20 inches away. With your right eye, look at the dot. Slowly bring the image (or move your head) closer while looking at the dot. At a certain distance, the + will disappear from sight...this is when the + falls on the blind spot of your retina. Reverse the process. Close your right eye and look at the + with your left eye. Move the image slowly closer to you and the dot should disappear. Here is another image that will help you find your blind spot. For this image, close your right eye. With your left eye, look at the red circle. Slowly move your head closer to the image. At a certain distance, the blue line will not look broken! Did you know? Find out more about blind spots, vision, the retina and photoreceptors. Copyright © 1996-2017, Eric H. Chudler, University of Washington
Functional parts of the rods and cones, which are two of the three types of photosensitive cells in the retina IdentifiersMeSHD010786NeuroLex IDsao226523927FMA85613 86740, 85613Anatomical terms of neuroanatomy[edit on Wikidata] A photoreceptor cell is a specialized type of neuroepithelial cell found in the retina that is capable of visual phototransduction. The great biological importance of photoreceptors is that they convert light (visible electromagnetic radiation) into signals that can stimulate biological processes. To be more specific, photoreceptor proteins in the cell absorb photons, triggering a change in the cell's membrane potential. There are currently three known types of photoreceptor cells in mammalian eyes: rods, cones, and intrinsically photosensitive retinal ganglion cells. The two classic photoreceptor cells are rods and cones, each contributing information used by the visual system to form an image of the environment, sight. Rods primarily mediate scotopic vision (dim conditions) whereas cones primarily mediate to photopic vision (bright conditions), but the processes in each that supports phototransduction is similar.[1] A third class of mammalian photoreceptor cell was discovered during the 1990s:[2] the intrinsically photosensitive retinal ganglion cells. These cells are thought not to contribute to sight directly, but have a role in the entrainment of the circadian rhythm and pupillary reflex. PhotosensitivityEach photoreceptor absorbs light according to its spectral sensitivity (absorptance), which is determined by the photoreceptor proteins expressed in that cell. Humans have three classes of cones (L, M, S) that each differ in spectral sensitivity and 'prefer' photons of different wavelengths (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. In accordance with the principle of univariance, 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 abosrbs 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 color vision. HistologyAnatomy of rods and cones varies slightly. Rod and cone photoreceptors are found on the outermost layer of the retina; they both have the same basic structure. Closest to the visual field (and farthest from the brain) is the axon terminal, which releases a neurotransmitter called glutamate to bipolar cells. Farther back is the cell body, which contains the cell's organelles. Farther back still is the inner segment, a specialized part of the cell full of mitochondria. The chief function of the inner segment is to provide ATP (energy) for the sodium-potassium pump. Finally, closest to the brain (and farthest from the field of view) is the outer segment, the part of the photoreceptor that absorbs light. Outer segments are actually modified cilia[5][6] that contain disks filled with opsin, the molecule that absorbs photons, as well as voltage-gated sodium channels. The membranous photoreceptor protein opsin contains a pigment molecule called retinal. In rod cells, these together are called rhodopsin. In cone cells, there are different types of opsins that combine with retinal to form pigments called photopsins. Three different classes of photopsins in the cones react to different ranges of light frequency, a differentiation that allows the visual system to calculate color. 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 signal transduction. The opsin found in the intrinsically photosensitive ganglion cells of the retina is called melanopsin. These cells are involved in various reflexive responses of the brain and body to the presence of (day)light, such as the regulation of circadian rhythms, pupillary reflex and other non-visual responses to light. Melanopsin functionally resembles invertebrate opsins. Retinal mosaicMost vertebrate photoreceptors are located in the retina. The distribution of rods and cones (and classes thereof) in the retina is called the retinal mosaic. Each human retina has approximately 6 million cones and 120 million rods.[8] At the "center" of the retina (the point directly behind the lens) lies the fovea (or fovea centralis), which contains only cone cells; and is the region capable of producing the highest visual acuity or highest resolution. Across the rest of the retina, rods and cones are intermingled. No photoreceptors are found at the blind spot, the area where ganglion cell fibers are collected into the optic nerve and leave the eye.[9] 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. The number and ratio of rods to cones varies among species, dependent on whether an animal is primarily diurnal or nocturnal. Certain owls, such as the nocturnal tawny owl,[10] have a tremendous number of rods in their retinae. Other vertebrates will also have a different number of cone classes, ranging from monochromats to pentachromats. SignalingThe path of a visual signal is described by the phototransduction cascade, 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 optic nerve. The steps that apply to the phototransductuion pathway from vertebrate rod/cone photoreceptors are:
hyperpolarizationUnlike most sensory receptor cells, photoreceptors actually become hyperpolarized when stimulated; and conversely are depolarized 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 cyclic guanosine 3'-5' monophosphate (cGMP), which opens cGMP-gated ion channels. These channels are nonspecific, allowing movement of both sodium and calcium ions when open. The movement of these positively charged ions into the cell (driven by their respective electrochemical gradient) depolarizes the membrane, and leads to the release of the neurotransmitter glutamate. Unstimulated (in the dark), cyclic-nucleotide gated channels in the outer segment are open because cyclic GMP (cGMP) is bound to them. Hence, positively charged ions (namely sodium ions) enter the photoreceptor, depolarizing it to about −40 mV (resting potential in other nerve cells is usually −65 mV). This depolarization current is often known as dark current. bipolar cellsThe photoreceptors (rods and cones) transmit to the bipolar cells, which transmit then to the retinal ganglion cells. Retinal ganglion cell axons collectively form the optic nerve, via which they project to the brain.[8] 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. Absorption of a photon will hyperpolarize the photoreceptor and therefore result in the release of less glutamate at the presynaptic terminal to the bipolar cell. 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 receptor imbedded in that cell's membrane. When glutamate binds to an ionotropic receptor, the bipolar cell will depolarize (and therefore will hyperpolarize with light as less glutamate is released). On the other hand, binding of glutamate to a metabotropic receptor results in a hyperpolarization, so this bipolar cell will depolarize to light as less glutamate is released. 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 detecting color, contrast, edges, etc. AdvantagesPhototransduction in rods and cones is somewhat unusual in that the stimulus (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:
Difference between rods and conesComparison of human rod and cone cells, from Eric Kandel et al. in Principles of Neural Science.[11]
DevelopmentThe 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 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. Early Notch 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 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, retinitis pigmentosa, macular degeneration or other visual deficits may result.[12] Ganglion cell photoreceptorsThis subset (≈1–3%) of retinal ganglion cells, unlike other retinal ganglion cells, are intrinsically photosensitive due to the presence of melanopsin, a light-sensitive protein. Therefore they constitute a third class of photoreceptors, in addition to rod and cone cells.[13] In humans the ipRGCs contribute to non-image-forming functions like circadian rhythms, behavior and pupillary light reflex.[14] Peak spectral sensitivity of the receptor is between 460 and 482 nm.[14] However, they may also contribute to a rudimentary visual pathway enabling conscious sight and brightness detection.[14] 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 ganglion cells, and the study of the receptor offered potential as a new avenue to explore in trying to find treatments for blindness. ipRGCs were only definitively detected ipRGCs in humans during landmark experiments in 2007 on rodless, coneless humans.[15][16] 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 down patients with rare diseases wiping out classic rod and cone photoreceptor function but preserving ganglion cell function.[15][16] 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. Non-human photoreceptorsRod 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 absence of input from the eyes or neurotransmitters.[17] Invertebrate photoreceptors in organisms such as insects and molluscs are different in both their morphological organization and their underlying biochemical pathways. This article describes human photoreceptors. See also
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