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713:, anti-inflammatory, and vasoprotective properties of anthocyanins allow them to demonstrate diverse health effects. In humans, anthocyanins are effective for a variety of health conditions including neurological damage, atherosclerosis, diabetes, as well as visual impairment. Anthocyanins frequently interact with other phytochemicals to potentiate biological effects; therefore, contributions from individual biomolecules remains difficult to decipher. As a result of anthocyanins providing bright colouration to flowers, the plants containing these phytochemicals are naturally successful in attracting pollinators such as birds and bees. The fruits and vegetables produced by such plants are also brightly pigmented attracting animals to eat them and disperse the seeds. Due to this natural mechanism anthocyanin-containing plants are widely abundant in most areas of the world. The high abundance and distribution of anthocyanin-containing plants make it a natural food source for many animals. Through fossil evidence it is known that these compounds were eaten in high amounts by primitive hominins.
540:(RP), and other retinal diseases, with increasing impairment as the diseases progress. AMD is a chronic, progressive disease that causes a part of the retina, called the macula, to slowly deteriorate over time. It is the leading cause of vision loss among people age 50 and older. It is characterized by a breakdown of the RPE/Bruch's membrane complex in the retina, leading to an accumulation of cholesterol deposits in the macula. Eventually, these deposits become clinically visible drusen that affect photoreceptor health, causing inflammation and a predisposition to choroidal neovascularization (CNV). During the AMD disease course, the RPE/Bruch's function continues to deteriorate, hampering nutrient and oxygen transport to the rod and cone photoreceptors. As a side effect of this process, the photoreceptors exhibit impaired dark adaptation because they require these nutrients for replenishment of photopigments and clearance of opsin to regain scotopic sensitivity after light exposure.
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adaptation. In order for dark adaptation to be significantly accelerated an individual should ideally begin this practice 30 minutes prior to entering a low luminescence setting. This practice will allow an individual to maintain their photopic (day) vision whilst preparing for scotopic vision. The insensitivity to red light will prevent the rod cells from further becoming bleached and allow for the rhodopsin photopigment to recharge back to its active conformation. Once an individual enters a dark setting most of their rod cells will already be accommodated to the dark and be able to transmit visual signals to the brain without an accommodation period.
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evolution of mammals from their reptilian ancestors approximately 275 million years ago there was a nocturnal phase in which complex colour vision was lost. Being that these pro-mammals were nocturnal they increased their sensitivity in low luminescence settings and reduced their photopic system from tetrachromatic to dichromatic. The shift to a nocturnal lifestyle would demand more rod photoreceptors to absorb the blue light emitted by the moon during the night. It can be extrapolated that the high ratio of rods to cones present in modern human eyes was retained even after the shift from nocturnal back to diurnal.
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evolution for sensing light because retinal causes a shift in photoreceptor absorbance to the visible light range. This shift in absorbance is especially important for life on Earth because it generally matches the peak irradiance of sunlight on its surface. A second reason why retinal evolved to be vital for human vision is because it undergoes a large conformational change when exposed to light. This conformational change is believed to make it easier for the photoreceptor protein to distinguish between its silent and activated state thus better controlling visual phototransduction.
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wavelengths (green light), or long wavelengths (red light). Rod photoreceptors only contain one type of photopigment, rhodopsin, which has a peak sensitivity at a wavelength of approximately 500 nanometers which corresponds to blue-green light. The distribution of photoreceptor cells across the surface of the retina has important consequences for vision. Cone photoreceptors are concentrated in a depression in the center of the retina known as the
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onions, blueberries, bilberries, red cabbage, and eggplant. The ingestion of any of these food sources will yield a variety of phytochemicals in addition to anthocyanins because they naturally exist together. The daily intake of anthocyanins is estimated to be approximately 200 milligrams in the average adult; however, this value can reach several grams per day if an individual is consuming flavonoid supplements.
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period of dark adaptation was measured prior to and two hours following supplementation in all participants. Results from this experiment indicate that anthocyanins significantly accelerated dark adaptation at merely one dose level compared to the placebo. Observing the data as a whole
Nakaishi et al. concluded that anthocyanins effectively reduced the dark adaptation period in a dose-dependent manner.
709:. This group of approximately 600 bioactive antioxidants carries the strongest physiological effects of any plant compound. These chemicals are also the most visible of the flavonoid phytochemicals because they provide bright blue, red, or purple pigmentation to many plant species. Anthocyanins also serve to protect the photosynthetic tissues from the direct rays of the sun. In addition, the
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structure thereby accelerating its access to the retinal binding pocket. By having a diet rich in anthocyanins an individual is able to generate rhodopsin in shorter periods of time because of the increased affinity of opsin to retinal. Through this mechanism an individual is able to accelerate dark adaptation and achieve night vision in a shorter period of time.
504:, rapid sensitivity fluctuations, is powered by nerve control. The merging of signals by virtue of the diffuse ganglion cells, as well as horizontal and amacrine cells, allow a cumulative effect. Thus that area of stimulation is inversely proportional to intensity of light, a strong stimulus of 100 rods equivalent to a weak stimulus of 1,000 rods.
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function was measured prior to supplementation, 1 day post-treatment, and 75 days post-treatment. It was observed that after merely one day of vitamin A supplementation the recovery kinetics of dark adaptation were significantly accelerated after photoreceptor bleaching. Dark adaptation was further accelerated following 75 days of treatment.
224:(ipRGCs). Rods and cones are responsible for vision and connected to the visual cortex. ipRGCs are more connected to body clock functions and other parts of the brain but not the visual cortex. Rods and cones can be easily distinguished by their structure. Cone photoreceptors are conical in shape and contain cone
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and therefore a lack of vitamin A in the diet. In developed countries night blindness has historically been uncommon due to adequate food availability; however, the incidence is expected to increase as obesity becomes more common. Increased obesity rates correspond to an increased number of bariatric
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is necessary for proper functioning of the human eye. The photopigment rhodopsin found in human rod cells is composed of retinal, a form of vitamin A, bound to an opsin protein. Upon the absorption of light rhodopsin was decomposed into retinal and opsin through bleaching. Retinal could then have one
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and decrease in number towards the periphery of the retina. Conversely, rod photoreceptors are present at high density throughout the most of the retina with a sharp decline in the fovea. Perception in high luminescence settings is dominated by cones despite the fact that they are greatly outnumbered
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In a double-blind, placebo-controlled study conducted by
Nakaishi et al. a powdered anthocyanin concentrate derived from black currants was provided to a number of participants. Participants received one of three doses of anthocyanins to measure if the result occurred in a dose-dependent manner. The
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in primates occurred approximately 55 million years ago when the surface temperature of the planet began to rise. The primates were diurnal rather than nocturnal in nature and therefore required a more precise photopic visual system. A third cone photopigment was necessary to cover the entire visual
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Despite the fact that the resolution of human day vision is far superior to that of night vision, human night vision provides many advantages. Like many predatory animals, humans can use their night vision to prey upon and ambush other animals without their awareness. Furthermore, in the event of an
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adaptation following the intake of blueberry products. In neither study did the blueberry anthocyanin intake affect the length of dark adaptation. From these results Kalt et al. concluded that blueberry anthocyanins provide no significant difference to the dark adaptation component of human vision.
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Brightly coloured fruits and vegetables are rich in anthocyanins. This makes sense intuitively because anthocyanins offer pigmentation to plants. Blackberries are the most anthocyanin-rich foods, containing 89-211 milligrams per 100 grams. Other foods that are rich in this phytochemical include red
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Vitamin A is present in both animal and plant sources as retinoids and carotenoids, respectively. Retinoids can be used immediately by the body upon absorption into the cardiovascular system; however, plant-based carotenoids must be converted to retinol prior to utilization by the body. The highest
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Vitamin A serves many functions in the human body outside of healthy vision. It is vital in maintaining a healthy immune system as well as promoting normal growth and development. The average adult male and female should consume 900 and 700 micrograms of vitamin A per day, respectively. Consumption
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which hydrolyses cGMP to further reduce its concentration. This reduces opening of the cGMP gated Na channels to hyperpolarise the cell, once again making it sensitive to small increases in brightness. Without dark adaptation, the photoreceptor would remain depolarized under scotopic conditions and
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Intensity and duration of the pre-adapting light: By increasing the levels of pre-adapting luminances, the duration of cone mechanism dominance extends, while the rod mechanism switch over is more delayed. In addition the absolute threshold takes longer to reach. The opposite is true for decreasing
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Various studies have been conducted testing the effective of vitamin A supplementation on dark adaptation. In a study by
Cideciyan et al. the length of dark adaptation was measured in a patient with systemic vitamin A deficiency (VAD) before and after vitamin A supplementation. The dark adaptation
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Submarines: Oftentimes submarines are "rigged for red", meaning that the boat is going to be surfacing or coming to periscope depth at night. During such times illumination within certain compartments is switched to red light to allow the eyes of the lookouts and officers to adjust to the darkness
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Aviators commonly wear red lensed glasses or goggles prior to taking off in the dark to ensure that they are able to see outside of the aircraft. Furthermore, throughout flight the cockpit is illuminated with dim red lights. This lighting is to ensure that the pilot is able to read instruments and
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Wavelength of the threshold light: Varying the wavelengths of stimuli also affect the dark adaptation curve. Long wavelengths—such as extreme red—create the absence of a distinct rod/cone break, as the rod and cone cells have similar sensitivities to light of long wavelengths. Conversely, at short
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Anthocyanins accelerate dark adaptation in humans by enhancing the regeneration of the rod photopigment, rhodopsin. Anthocyanins accomplish this by binding directly to opsin upon the degradation of rhodopsin to its individual constituents by light. Once bound to opsin, the anthocyanin changes its
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Although many aspects of the human visual system remain uncertain, the theory of the evolution of rod and cone photopigments is agreed upon by most scientists. It is believed that the earliest visual pigments were those of cone photoreceptors, with rod opsin proteins evolving later. Following the
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and all-trans retinal. Dark adaptation of both rods and cones requires the regeneration of the visual pigment from opsin and 11-cis retinal. Therefore, the time required for dark adaptation and pigment regeneration is largely determined by the local concentration of 11-cis retinal and the rate at
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emergency situation occurring at night, humans can increase their chances of survival if they are able to perceive their surroundings and get to safety. Both of these benefits can be used to explain why humans did not completely lose the ability to see in the dark from their nocturnal ancestors.
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Despite the fact that many scientists believe anthocyanins to be beneficial in accelerating dark adaptation in humans, a study conducted by Kalt et al. in 2014 showed blueberry anthocyanins have no effect. In this study two double-blind, placebo-controlled studies were conducted to examine dark
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In sufficiently bright light, convergence is low, but during dark adaptation, convergence of rod signals boost. This is not due to structural changes, but by a possible shutdown of inhibition that stops convergence of messages in bright light. If only one eye is open, the closed eye must adapt
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Vitamin A-based opsin proteins have been used for sensing light in organisms for most of evolutionary history beginning approximately 3 billion years ago. This feature has been passed from unicellular to multicellular organisms including Homo sapiens. This vitamin was most likely selected by
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As a result of rod cells having a peak sensitivity at a wavelength of 500 nanometers they cannot perceive all colours on the visual spectrum. Because rod cells are insensitive to long wavelengths, the use of red lights and red lens glasses has become a common practice for accelerating dark
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as their visual pigments. There exist three types of cone photoreceptors, each being maximally sensitive to a specific wavelength of light depending on the structure of their opsin photopigment. The various cone cells are maximally sensitive to either short wavelengths (blue light), medium
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Night blindness can be caused by a number of factors the most common of which being vitamin A deficiency. If detected early enough nyctalopia can be reversed and visual function can be regained; however; prolonged vitamin A deficiency can lead to permanent visual loss if left untreated.
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During World Wars I and II British Air Force aviators were known to consume extensive amounts of bilberry jam. The aviators consumed this anthocyanin-rich food due to its many visual benefits, included accelerated dark adaptation, which would be valuable for night bombing missions.
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Measurement of a patient's dark adaptation function is essentially a bioassay of the health of their Bruch's membrane. As such, research has shown that, by measuring dark adaptation, doctors can detect subclinical AMD at least three years earlier than it is clinically evident.
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which it is delivered to the opsin in the bleached rods. The decrease in calcium ion influx after channel closing causes phosphorylation of metarhodopsin II and speeds up the cis-retinal to trans-retinal inactivation. The phosphorylation of activated rhodopsin is mediated by
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This part of the curve is limited by quantal fluctuation in the background. The visual system is usually compared with a theoretical construct called the ideal light detector. To detect the stimulus, the stimulus must sufficiently exceed the fluctuations of the background
141:). However, it takes approximately five minutes for the eye to adapt from darkness to bright sunlight. This is due to cones obtaining more sensitivity when first entering the dark for the first five minutes but the rods taking over after five or more minutes.
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of two fates: it could recombine with opsin to reform rhodopsin or it could be converted into free retinol. The
American scientist George Wald was the first to recognize that the visual system expends vitamin A and is dependent upon diet for its replacement.
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The eye takes approximately 20–30 minutes to fully adapt from bright sunlight to complete darkness and becomes 10,000 to 1,000,000 times more sensitive than at full daylight. In this process, the eye's perception of color changes as well (this is called the
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period of up to two hours in which each eye adjusts from a high to a low luminescence "setting", increasing sensitivity hugely, by many orders of magnitude. This adaptation period is different between rod and cone cells and results from the regeneration of
685:, both of whom had vitamin A deficiency. Within 8 days of oral supplementation of vitamin A both patients had their visual function restored to normal. Furthermore, adaptation kinetics significantly improved in both subjects following supplementation.
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to allow sodium and calcium influx. Sodium influx contributes to depolarization while calcium influx increases local calcium concentrations near the receptor. Calcium binds to a modulatory protein, which is proposed to be
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There are multiple commercial dark adaptometers (AdaptDx, MetroVision MonCvONE, Roland
Consult DARK-Adaptometer). Besides those free-viewing devices, a fundus-tracked workflow has been recently introduced by modifying a
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improves considerably within 5–10 minutes in the dark. Color testing has been used to determine the time at which rod mechanism takes over; when the rod mechanism takes over colored spots appear colorless as only
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The threshold in this portion of the TVI curve is determined by the dark/light level. Sensitivity is limited by neural noise. The background field is relatively low and does not significantly affect threshold.
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With light adaptation, the eye has to quickly adapt to the background illumination to be able to distinguish objects in this background. The process for light adaptation occurs over a period of five minutes.
263:, adjusting the amount of light that reaches the retina very quickly by about a factor of ten. Since it contributes only a tiny fraction of the overall adaptation to light it is not further considered here.
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Nakaishi, H.; Matsumoto, H.; Tominaga, S.; Hirayama, M. (2000). "Effects of black currant anthocyanoside intake on dark adaptation and vdt work-induced transient refractive alteration in healthy humans".
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Visual
Response to Darkness. Cones work at high light levels (during the day but also during driving at night in the headlamp spotlight). Rods take over at twilight and night. The y-axis has logarithmic
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Kalt, Wilhelmina; McDonald, Jane E.; Fillmore, Sherry A. E.; Tremblay, Francois (2014). "Blueberry
Effects on Dark Vision and Recovery after Photobleaching: Placebo-Controlled Crossover Studies".
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Pfau K, Jeffrey BG, Cukras CA (September 2023). "Low-Dose
Supplementation with Retinol Improves Retinal Function in Eyes with Age-Related Macular Degeneration but Without Reticular Pseudodrusen".
800:, the stimulus is increased until the detection threshold is reached against the background. A monophasic or biphasic threshold versus intensity TVI curve is obtained through this method for both
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are more sensitive to light and so take longer to fully adapt to the change in light. Rods, whose photopigments regenerate more slowly, do not reach their maximum sensitivity for about two hours.
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Later independent studies in Sorby fundus dystrophy confirmed the effect of
Vitamin A on dark adaptation. Likewise, Vitamin A was shown to accelerate (to a lesser extent) dark adaptation in AMD.
129:. This means that the brightest and the darkest light signal that the eye can sense are a factor of roughly 1,000,000,000 apart. However, in any given moment of time, the eye can only sense a
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animal-based sources of vitamin A are liver, dairy products, and fish. Fruits and vegetables containing high amounts of carotenoids are dark green, yellow, orange, and red in colour.
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Using increment threshold experiments, light adaptation can be measured clinically. In an increment threshold experiment, a test stimulus is presented on a background of a certain
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of eye function both in isolation and in tandem to adjust the visual system. Changes in the sensitivity of rods and cones in the eye are the major contributors to dark adaptation.
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Ollivier, F. J.; Samuelson, D. A.; Brooks, D. E.; Lewis, P. A.; Kallberg, M. E.; Komaromy, A. M. (2004). "Comparative morphology of the tapetum lucidum (among selected species)".
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Raming K, Gliem M, Charbel Issa P, Birtel J, Herrmann P, Holz FG, Pfau M, Hess K (February 2022). "Visual
Dysfunction and Structural Correlates in Sorsby Fundus Dystrophy".
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are nocturnal most likely because upon reflection of light back through the retina the initial images become blurred. Humans, like their primate relatives, do not possess a
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Owsley, Cynthia; McGwin, Gerald; Clark, Mark E.; Jackson, Gregory R.; Callahan, Michael A.; Kline, Lanning B.; Witherspoon, C. Douglas; Curcio, Christine A. (2016-02-01).
476:. This reduces cGMP production by guanylyl cyclase to lower cGMP concentration during prolonged darkness. Elevated calcium concentration also increases the activity of
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prior to looking outside of the boat. Additionally, compartments on a submarine may be illuminated with red light in order to simulate night conditions for the crew.
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Tirupula, K. C.; Balem, F.; Yanamala, N.; Klein-Seetharaman, J. (2009). "ph-dependent interaction of rhodopsin with cyanidin-3-glucoside. 2. functional aspects".
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Watson, S., & Gorski, K. A. (2011). Invasive cardiology: A manual for cath lab personnel. (3rd ed., pp. 61-62). Sudbury, MA: Jones & Bartlett Learning.
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Dietary Reference Intakes for Vitamin A, Vitamin K, Arsenic, Boron, Chromium, Copper, Iodine, Iron, Manganese, Molybdenum, Nickel, Silicon, Vanadium, and Zinc
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Kemp, Colin M.; Jacobson, Samuel G.; Faulkner, David J.; Walt, Robert W. (1988). "Visual function and rhodopsin levels in humans with vitamin A deficiency".
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Several different methods, with varying levels of evidence, have been purported or demonstrated to increase the rate at which vision can adapt in the dark.
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At saturation, the rod system becomes unable to detect the stimulus. This section of the curve occurs for the cone mechanism under high background levels.
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Structure, function, and pathology of Bruch's membrane. In: Ryan SJ, et al, eds. Retina, Vol 1, Part 2: Basic Science and Translation to Therapy. 5th ed
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Owsley, C.; Jackson, G. R.; White, M.; Feist, R.; Edwards, D. (2001-07-01). "Delays in rod-mediated dark adaptation in early age-related maculopathy".
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Bhatia, K; Jenkins, C; Prasad, M; Koki, G; Lombange, J (1989). "Immunogenetic studies of two recently contacted populations from Papua New Guinea".
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1990:
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Losso, J. N., Shahidi, F., & Bagchi, D. (2007). Anti-angiogenic functional and medicinal foods. Boca Raton, FL: Taylor & Francis Group.
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and colors cannot be discerned; only shades of gray are seen. In order for humans to transition from day to night vision they must undergo a
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wavelengths the rod/cone break is more prominent, because the rod cells are much more sensitive than cones once the rods have dark adapted.
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Stuart JA, Brige RR (1996). "Characterization of the primary photochemical events in bacteriorhodopsin and rhodopsin". In Lee AG (ed.).
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Bartlett NR. Dark and light adaptation. In: Graham CH, editor. Vision and visual perception. New York: John Wiley and Sons, Inc.; 1965.
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Size and location on the retina: The location of the test spot affects the dark adaptation curve because of the distribution of the
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possess high-resolution night vision, allowing them to discriminate objects with high frequencies in low illumination settings. The
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are only able to function at higher illumination levels. Night vision is of lower quality than day vision because it is limited in
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Aguilar M, Stiles WS. Saturation of the rod mechanism of the retina at high levels of stimulation. Opt Acta (Lond) 1954;1:59–65.
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is blind to dim light (due to its cone-only array) and the rods are more sensitive, so a dim star on a moonless night must be
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896:, so it stimulates the rods. This is not due to pupil width since an artificial fixed-width pupil gives the same results.
1036:"Sensory Reception: Human Vision: Structure and function of the Human Eye" vol. 27, p. 179 Encyclopædia Britannica, 1987
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Numerous clinical studies have shown that dark adaptation function is dramatically impaired from the earliest stages of
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are able to regain maximum retinal sensitivity in 9–10 minutes of darkness whereas rods require 30–45 minutes to do so.
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2700:"Reversible night blindness – a reminder of the increasing importance of vitamin a deficiency in the developed world"
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The human eye can function from very dark to very bright levels of light; its sensing capabilities reach across nine
1825:"Delayed Rod-Mediated Dark Adaptation Is a Functional Biomarker for Incident Early Age-Related Macular Degeneration"
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spectrum enabling primates to better discriminate between fruits and detect those of the highest nutritional value.
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take approximately 9–10 minutes to adapt to the dark. Sensitivity to light is modulated by changes in intracellular
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is a reflective structure that is responsible for this superior night vision as it mirrors light back through the
1027:"Sensory Reception: Human Vision: Structure and Function of the Human Eye" Encyclopædia Britannica, vol. 27, 1987
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Ophthalmologists sometimes measure patients' dark adaptation using an instrument known as a dark adaptometer.
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1686:"Establishing Fully-Automated Fundus-Controlled Dark Adaptometry: A Validation and Retest-Reliability Study"
1578:"Cyclic GMP and calcium: The internal messengers of excitation and adaptation in vertebrate photoreceptors"
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Owsley C, McGwin G, Jackson GR, Heimburger DC, Piyathilake CJ, Klein R, White MF, Kallies K (April 2006).
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Kolb, Helga (2013). "Photoreceptors". In Helga Kolb; Ralph Nelson; Eduardo Fernandez; Bryan Jones (eds.).
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Insufficiency of adaptation most commonly presents as insufficient adaptation to dark environment, called
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396:. The regeneration of the photopigments occurs during dark adaptation albeit at markedly different rates.
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289:, as there is not an abrupt transition between the two mechanism. This adaptation forms the basis of the
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Purves, D., Augustine, G. J., & Fitzpatrick, D. (2001). Neuroscience. (2nd ed.). Sinauer Associates.
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2293:"Effect of short-term, high-dose retinol on dark adaptation in aging and early age-related maculopathy"
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1627:"Purification and physiological evaluation of a guanylate cyclase activating protein from retinal rods"
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ratio of 1,000. What enables the wider reach is that the eye adapts its definition of what is black.
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to increase retinal sensitivity. Light adaptation, in contrast, works very quickly, within seconds.
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H Davson. Physiology of the eye. 5th ed. London: Macmillan Academic and Professional Ltd.; 1990.
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H Davson. Physiology of the eye. 5th ed. London: Macmillan Academic and Professional Ltd.; 1990
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2197:"Rod plateaux during dark adaptation in sorsby's fundus dystrophy and vitamin a deficiency".
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Schwab, I. R.; Yuen, C. K.; Buyukmihci, N. C.; Blankenship, T. N.; Fitzgerald, P. G. (2002).
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Inhibition by neurons also affects activation in synapses. Together with the bleaching of a
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Extreme red light used on a ship's bridge at night to aid dark adaptation of the crew's eyes
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Normalised absorption spectra of the three human photopsins and of human rhodopsin (dashed).
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1548:"Human eye | Definition, Anatomy, Diagram, Function, & Facts | Britannica"
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Lamb, T. D.; Pugh Jr, E. N. (2004). "Dark adaptation and the retinoid cycle of vision".
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for radiologists to adapt their eyes to view screens during fluoroscopic procedures.
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2637:"Temporal and spatial summation in human vision at different background intensities"
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level (about 0.03 cd/m), the cone mechanism is involved in mediating vision;
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2143:"Retina, retinol, retinal and the natural history of vitamin a as a light sensor"
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starts with the isomerizing of the pigment chromophore from 11-cis to all-trans
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Proceedings of the National Academy of Sciences of the United States of America
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above 3000 micrograms per day can lead to acute or chronic hypervitaminosis A.
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The concept of red lenses for dark adaptation is based upon experimentation by
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1129:"Light and Dark Adaptation by Michael Kalloniatis and Charles Luu – Webvision"
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Dark adaptation is far quicker and deeper in young people than the elderly.
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A subsequent study by Kemp et al. studied dark adaptation in subjects with
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2109:. Food and Nutrition Board of the Institute of Medicine. pp. 82–161.
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Rhodopsin and G-Protein Linked Receptors, Part A (Vol 2, 1996) (2 Vol Set)
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375:, a biological pigment in the photoreceptors of the retina, immediately
246:
The pupillary light reflex is a quick but minor mechanism of adaptation
1008:
http://research.physics.illinois.edu/QI/Photonics/pdf/PWDec16Holmes.pdf
1006:
Rebecca Holmes, "Seeing single photons". Physics World, December 2016.
903:
Night blindness is especially prominent in developing countries due to
405:
384:
220:
The human eye contains three types of photoreceptors, rods, cones, and
176:
89:
2595:
2159:
869:
Effect of night blindness. Left: good night vision. Right: nightblind.
2006:
Summitt, D. (2004). Tales of a cold war submariner. (1st ed., p. 138)
469:
430:
Three factors affect how quickly the rod mechanism becomes dominant:
331:
31:
1997:, Federal Aviation Administration. (2015). Medical facts for pilots.
885:, that is, inability to see clearly in bright light, is much rarer.
338:
cells to an increased amount of light. Most animals which possess a
285:. The range where two mechanisms are working together is called the
2698:
Clifford, Luke J.; Turnbull, Andrew M.J.; Denning, Anne M. (2013).
2376:"Anthocyanins and Human Health: An In Vitro Investigative Approach"
1684:
Oertli JM, Pfau K, Scholl HP, Jeffrey BG, Pfau M (December 2023).
1479:
Aubert H. Physiologie der Netzhaut. Breslau: E. Morgenstern; 1865.
864:
787:
724:
630:
551:
388:
357:
306:
249:
225:
211:
908:
surgeries, causing malabsorption of vitamin A in the human body.
811:) is taken in isolation it can be seen to possess four sections:
463:
concentration is high in photoreceptors. cGMP binds to and opens
2753:
Light and Dark Adaptation by Michael Kalloniatis and Charles Luu
524:
device that enables testing of patients with unstable fixation.
281:. Below this level, the rod mechanism comes into play providing
46:, is the ability to see under low-light conditions. In humans,
982:
US Department of Defense, Defense Technical Information Center
408:
321:
161:
74:
35:
792:
Schematic of the increment threshold curve of the rod system
508:
separately upon reopening to match the already adapted eye.
481:
so also remain unresponsive to small changes in brightness.
1093:
Webvision: The Organization of the Retina and Visual System
612:
maps while maintaining scotopic vision for looking outside.
577:
and his early work with radiology. In 1916, the scientist
346:
and therefore were predisposed to be a diurnal species.
186:
99:
2141:
Zhong, M.; Kawaguchi, R.; Kassai, M.; Sun, H. (2012).
1190:
Transactions of the American Ophthalmological Society
528:
Using Dark Adaptation Measurement to Diagnose Disease
807:
When the threshold curve for a single system (i.e.,
2027:"The discovery of the visual function of vitamin A"
1227:"Eye shape and the nocturnal bottleneck of mammals"
234:by rods (approximately 4.5 million to 91 million).
222:
intrinsically photosensitive retinal ganglion cells
844:proportional to the square root of the background.
1225:Hall, M. I.; Kamilar, J. M.; Kirk, E. C. (2012).
2199:Investigative Ophthalmology & Visual Science
1500:"The variations in visual threshold measurement"
472:, removing this protein's stimulatory effect on
50:are exclusively responsible for night vision as
1278:. Greenwich, Conn: JAI Press. pp. 33–140.
1892:, Abbott, B. (2012). Sensation and perception.
1795:"Facts About Age-Related Macular Degeneration"
38:to adjust to various levels of light. Natural
2441:, Sterling, M. (2001). What are anthocyanins?
2072:
2070:
2068:
2066:
2064:
2062:
266:In response to varying ambient light levels,
8:
1403:Psychology: The Science of Mind and Behavior
1023:
1021:
1019:
1017:
1015:
978:"Night Vision Manual for the Flight Surgeon"
2693:
2691:
2689:
2427:
2425:
2423:
2421:
1925:"Evolution of colour vision in vertebrates"
1918:
1916:
1914:
1912:
1910:
2584:Journal of Agricultural and Food Chemistry
1625:Gorczyca, W.A.; Gray-Keller, M.P. (1994).
1144:
1142:
1046:Jackson GR, Owsley C, McGwin G Jr (1999).
387:. Then this pigment dissociates into free
2745:at the U.S. National Library of Medicine
2723:
2660:
2401:
2391:
2308:
2168:
2158:
2042:
1940:
1900:
1898:
1878:
1876:
1856:
1711:
1701:
1660:
1650:
1593:
1523:
1456:
1250:
1201:
1063:
200:Learn how and when to remove this message
113:Learn how and when to remove this message
2380:Journal of Biomedicine and Biotechnology
2136:
2134:
2132:
2130:
2128:
2126:
1571:
1569:
1567:
1114:
1112:
1110:
971:
969:
967:
965:
241:
2577:
2575:
2573:
2544:
2542:
2540:
2496:
2494:
2492:
2457:
2455:
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2451:
2449:
2447:
2369:
2367:
2365:
2363:
2216:
2214:
2212:
2020:
2018:
2016:
2014:
2012:
1301:
1299:
1297:
1295:
976:Miller RE, Tredici TJ (1 August 1992).
961:
705:make up the majority of the 4000 known
259:A minor mechanism of adaptation is the
2192:
2190:
2188:
1085:
1083:
435:the levels of pre-adapting luminances.
2078:"Vitamins and Minerals - Unlock Food"
597:It is believed that the emergence of
497:are inhibited, reducing convergence.
16:Response of the eye to light and dark
7:
1335:Progress in Retinal and Eye Research
988:from the original on January 4, 2022
840:Threshold increases with background
1965:"Circadian Rhythm and Human Health"
1576:Pugh, E.N. Jr.; Lamb, T.D. (1990).
2464:"Bioflavonoids - Medicinal Plants"
1231:Proceedings of the Royal Society B
187:improve coverage for other species
100:improve coverage for other species
14:
881:. The opposite problem, known as
556:Astronomer preserves night vision
2515:10.1111/j.1751-1097.2008.00533.x
1347:10.1016/j.preteyeres.2004.03.001
1163:10.1111/j.1463-5224.2004.00318.x
534:Age-related Macular Degeneration
311:Reflection of camera flash from
166:
79:
2503:Photochemistry and Photobiology
773:The photochemical reaction is:
2653:10.1113/jphysiol.1958.sp005978
2462:Wimmer, Richard (2022-11-15).
2101:Institute of Medicine (2001).
1516:10.1113/jphysiol.1969.sp008818
1433:"Shedding Light on Adaptation"
413:cyclic guanosine monophosphate
189:and discuss this issue on the
102:and discuss this issue on the
1:
2235:10.1016/S0014-4835(88)80076-9
1749:10.1016/s0161-6420(01)00580-2
1437:Journal of General Physiology
1065:10.1016/s0042-6989(99)00092-9
2346:10.1097/IAE.0000000000003840
1841:10.1016/j.ophtha.2015.09.041
1595:10.1016/0042-6989(90)90013-b
1431:Hurley, JB (February 2002).
548:Accelerating dark adaptation
2716:10.1016/j.optom.2013.01.002
2552:Alternative Medicine Review
1048:"Aging and dark adaptation"
777:Rhodopsin ⇌ retinal + opsin
581:invented the first pair of
2796:
1186:"Evolution of the tapetum"
858:
695:
624:
459:conditions, intracellular
365:
300:
297:Advantages of night vision
155:
2641:The Journal of Physiology
2393:10.1155/S111072430440401X
2297:Invest Ophthalmol Vis Sci
2270:10.1016/j.ajo.2021.07.032
2223:Experimental Eye Research
1400:Passer and Smith (2008).
738:Effect on dark adaptation
679:primary biliary cirrhosis
512:Measuring Dark Adaptation
181:excessively human-centric
94:excessively human-centric
2747:Medical Subject Headings
2031:The Journal of Nutrition
1406:(4th ed.). p.
1151:Veterinary Ophthalmology
707:flavonoid phytochemicals
493:, merging of signals on
451:Intracellular signalling
381:Visual phototransduction
1993:March 26, 2015, at the
809:just cones or just rods
418:The sensitivity of the
283:scotopic (night) vision
2635:Barlow, H. B. (1958).
2437:April 2, 2015, at the
1690:Transl Vis Sci Technol
1652:10.1073/pnas.91.9.4014
1243:10.1098/rspb.2012.2258
1133:webvision.med.utah.edu
870:
793:
756:Contradictory evidence
730:
636:
583:red adaptation goggles
557:
465:cGMP gated Na channels
379:in response to light.
363:
317:
261:pupillary light reflex
256:
247:
238:Ambient light response
217:
30:is the ability of the
2468:Chinese Herbs Healing
2044:10.1093/jn/131.6.1647
1703:10.1167/tvst.12.12.18
1449:10.1085/jgp.119.2.125
1377:"Clinical Guidelines"
868:
791:
728:
669:Experimental evidence
634:
579:Wilhelm Trendelenburg
565:Red lights and lenses
555:
361:
320:Many animals such as
310:
253:
245:
215:
2704:Journal of Optometry
2310:10.1167/iovs.05-1292
1942:10.1038/eye.1998.143
1096:. University of Utah
924:Dark adaptor goggles
894:viewed from the side
660:Evolutionary context
651:Sources of vitamin A
589:Evolutionary context
538:Retinitis Pigmentosa
2590:(46): 11180–11189.
1778:Curcio, CA (2013).
1643:1994PNAS...91.4014G
1510:(403–419): 403–19.
1498:Hallett PE (1969).
1237:(1749): 4962–4968.
783:Increment threshold
747:Supportive evidence
491:rod or cone pigment
175:This article about
127:orders of magnitude
88:This article about
2743:Adaptation,+Ocular
1888:2018-06-21 at the
1552:www.britannica.com
871:
794:
731:
637:
558:
500:Alpha adaptation,
364:
318:
257:
248:
218:
158:Photoreceptor cell
2768:Animal physiology
2596:10.1021/jf503689c
2374:Lila, MA (2004).
2160:10.3390/nu4122069
2153:(12): 2069–2096.
2082:www.unlockfood.ca
2025:Wolf, G. (2001).
1969:photobiology.info
1588:(12): 1923–1948.
1417:978-0-07-256334-4
1285:978-1-55938-659-3
939:Neural adaptation
729:Blackberry fruits
478:phosphodiesterase
210:
209:
202:
123:
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115:
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2727:
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2459:
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2429:
2416:
2415:
2405:
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2371:
2358:
2357:
2340:(9): 1462–1471.
2329:
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2312:
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2282:
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2247:
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2218:
2207:
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2121:
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2098:
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2074:
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2046:
2037:(6): 1647–1650.
2022:
2007:
2004:
1998:
1985:
1979:
1978:
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1975:
1961:
1955:
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1944:
1920:
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1902:
1893:
1880:
1871:
1870:
1860:
1820:
1814:
1813:
1811:
1810:
1801:. Archived from
1790:
1784:
1783:
1775:
1769:
1768:
1743:(7): 1196–1202.
1732:
1726:
1725:
1715:
1705:
1681:
1675:
1674:
1664:
1654:
1637:(9): 4014–4018.
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1558:
1544:
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1067:
1043:
1037:
1034:
1028:
1025:
1010:
1004:
998:
997:
995:
993:
973:
765:Light adaptation
474:guanylyl cyclase
291:Duplicity Theory
273:Above a certain
205:
198:
194:
170:
169:
162:
118:
111:
107:
83:
82:
75:
2795:
2794:
2788:
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2758:
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2739:
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2697:
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2678:
2634:
2633:
2629:
2624:
2620:
2615:
2611:
2581:
2580:
2571:
2548:
2547:
2538:
2500:
2499:
2490:
2485:
2481:
2472:
2470:
2461:
2460:
2445:
2439:Wayback Machine
2430:
2419:
2373:
2372:
2361:
2331:
2330:
2326:
2290:
2289:
2285:
2258:Am J Ophthalmol
2255:
2254:
2250:
2220:
2219:
2210:
2205:(9): 1786–1794.
2196:
2195:
2186:
2140:
2139:
2124:
2117:
2100:
2099:
2095:
2086:
2084:
2076:
2075:
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2024:
2023:
2010:
2005:
2001:
1995:Wayback Machine
1986:
1982:
1973:
1971:
1963:
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1958:
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1908:
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1890:Wayback Machine
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1582:Vision Research
1575:
1574:
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1546:
1545:
1541:
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1483:
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1140:
1127:
1126:
1122:
1117:
1108:
1099:
1097:
1089:
1088:
1081:
1058:(23): 3975–82.
1045:
1044:
1040:
1035:
1031:
1026:
1013:
1005:
1001:
991:
989:
975:
974:
963:
958:
953:
944:Purkinje effect
919:Adaptive system
914:
875:night blindness
863:
857:
826:Square root law
786:
767:
758:
749:
740:
723:
700:
694:
686:
683:Crohn's disease
671:
662:
653:
635:11-cis-Retinal2
629:
623:
608:
591:
575:Antoine Béclère
567:
560:
550:
530:
514:
487:
453:
370:
368:Purkinje effect
356:
354:Dark adaptation
344:tapetum lucidum
340:tapetum lucidum
327:tapetum lucidum
314:tapetum lucidum
305:
299:
279:photopic vision
240:
231:fovea centralis
206:
195:
184:
171:
167:
160:
154:
139:Purkinje effect
119:
108:
97:
84:
80:
73:
60:dark adaptation
44:scotopic vision
17:
12:
11:
5:
2793:
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2738:
2737:External links
2735:
2732:
2731:
2710:(3): 173–174.
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2647:(2): 337–350.
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2569:
2558:(6): 553–562.
2536:
2509:(2): 463–470.
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2386:(5): 306–313.
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2229:(2): 185–197.
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2008:
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1935:(3): 541–547.
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859:Main article:
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802:cones and rods
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495:ganglion cells
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442:in the retina.
440:rods and cones
436:
427:encode color.
355:
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268:rods and cones
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152:Cones vs. rods
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2303:(4): 1310–8.
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1923:Link (1998).
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1829:Ophthalmology
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1805:on 2015-12-22
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2103:"Vitamin A"
1782:. Elsevier.
1381:www.aoa.org
1196:: 187–200.
883:hemeralopia
836:Weber's law
711:antioxidant
698:Anthocyanin
599:trichromacy
420:rod pathway
2762:Categories
2473:2023-02-08
2087:2023-02-08
1974:2023-02-08
1809:2017-02-04
1696:(12): 18.
1557:2023-02-08
1386:2023-02-08
1100:2023-02-08
1052:Vision Res
956:References
879:nyctalopia
861:Nyctalopia
817:Dark light
696:See also:
625:See also:
485:Inhibition
366:See also:
301:See also:
156:See also:
143:Cone cells
71:Efficiency
56:resolution
52:cone cells
28:adaptation
24:physiology
2147:Nutrients
1849:1549-4713
1757:0161-6420
1504:J Physiol
992:4 January
929:Human eye
842:luminance
798:luminance
640:Vitamin A
627:Vitamin A
621:Vitamin A
394:recoverin
373:Rhodopsin
275:luminance
191:talk page
104:talk page
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912:See also
831:(noise).
457:scotopic
255:scaling.
131:contrast
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406:calcium
385:retinal
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