Cone cell

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Cone cells
Cone-fundamentals-with-srgb-spectrum.svg
Normalized responsivity spectra of human cone cells, S, M, and L types
Details
Location Retina of vertebrates
Function Color vision
Identifiers
MeSH D017949
NeuroLex ID sao1103104164
TH H3.11.08.3.01046
FMA 67748
Anatomical terms of neuroanatomy

Cone cells or cones are photoreceptor cells in the retina of the vertebrate eye. Cones are active in daylight conditions and enable Photopic vision, as opposed to rod cells, which are active in dim light and enable Scotopic vision. Most vertebrates (including humans) have several classes of cones, each sensitive to a different part of the visible spectrums of light. The comparison of the responses of different cone cell classes enables color vision. There are about six to seven million cones in a human eye (vs ~92 million rods), with the highest concentration occurring towards the macula and most densely packed in the fovea centralis, a 0.3 mm diameter rod-free area with very thin, densely packed cones. Conversely, like rods, they are absent from the optic disc, contributing to the blind spot. [1]

Contents

Cones are less sensitive to light than the rod cells in the retina (which support vision at low light levels), but allow the perception of color. They are also able to perceive finer detail and more rapid changes in images because their response times to stimuli are faster than those of rods. [2] In humans, cones are normally one of three types: S-cones, M-cones and L-cones, with each type bearing a different opsin: OPN1SW, OPN1MW, and OPN1LW respectively. These cones are sensitive to visible wavelengths of light that correspond to short-wavelength, medium-wavelength and longer-wavelength light respectively. [3] Because humans usually have three kinds of cones with different photopsins, which have different response curves and thus respond to variation in color in different ways, humans have trichromatic vision. Being color blind can change this, and there have been some verified reports of people with four types of cones, giving them tetrachromatic vision. [4] [5] [6] The three pigments responsible for detecting light have been shown to vary in their exact chemical composition due to genetic mutation; different individuals will have cones with different color sensitivity.

Structure

Classes

Most vertebrates have several different classes of cone cells, differentiated primarily by the specific photopsin expressed within. The number of cone classes determines the degree of color vision. Vertebrates with one, two, three or four classes of cones possess monochromacy, dichromacy, trichromacy and tetrachromacy, respectively.

Humans normally have three classes of cones, designated L, M and S for the long, medium and short wavelengths of the visible spectrum to which they are most sensitive. [7] L cones respond most strongly to light of the longer red wavelengths, peaking at about 560 nm. M cones, respond most strongly to yellow to green medium-wavelength light, peaking at 530 nm. S cones respond most strongly to blue short-wavelength light, peaking at 420 nm, and make up only around 2% of the cones in the human retina. The peak wavelengths of L, M, and S cones occur in the ranges of 564–580 nm, 534–545 nm, and 420–440 nm nm, respectively, depending on the individual.[ citation needed ] The typical human photopsins are coded for by the genes OPN1LW, OPN1MW, and OPN1SW. The CIE 1931 color space is an often-used model of spectral sensitivities of the three cells of a typical human. [8] [9]

Histology

The structure of a cone cell Cone cell eng.svg
The structure of a cone cell

Cone cells are shorter but wider than rod cells. They are typically 40–50 μm long, and their diameter varies from 0.5–4.0 μm. They are narrowest at the fovea, where they are the most tightly packed. The S cone spacing is slightly larger than the others. [10]

Like rods, each cone cell has a synaptic terminal, inner and outer segments, as well as an interior nucleus and various mitochondria. The synaptic terminal forms a synapse with a neuron bipolar cell. The inner and outer segments are connected by a cilium. [2] The inner segment contains organelles and the cell's nucleus, while the outer segment contains the light-absorbing photopsins, and is shaped like a cone, giving the cell its name. [2]

The outer segments of cones have invaginations of their cell membranes that create stacks of membranous disks. Photopigments exist as transmembrane proteins within these disks, which provide more surface area for light to affect the pigments. In cones, these disks are attached to the outer membrane, whereas they are pinched off and exist separately in rods. Neither rods nor cones divide, but their membranous disks wear out and are worn off at the end of the outer segment, to be consumed and recycled by phagocytic cells.

Distribution

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 cones. ConeMosaics.jpg
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 cones.
Distribution of rods and cones along a line passing through the fovea and the blind spot of a human eye Human photoreceptor distribution.svg
Distribution of rods and cones along a line passing through the fovea and the blind spot of a human eye

While rods outnumber cones in most parts of the retina, the fovea, responsible for sharp central vision, consists almost entirely of cones. The distribution of photoreceptors in the retina is called the retinal mosaic, which can be determined using photobleaching. This is done by exposing dark-adapted retina to a certain wavelength of light that paralyzes the particular type of cone sensitive to that wavelength for up to thirty minutes from being able to dark-adapt, making it appear white in contrast to the grey dark-adapted cones when a picture of the retina is taken. The results illustrate that S cones are randomly placed and appear much less frequently than the M and L cones. The ratio of M and L cones varies greatly among different people with regular vision (e.g. values of 75.8% L with 20.0% M versus 50.6% L with 44.2% M in two male subjects). [12]

Function

Bird, reptilian, and monotreme cone cells BirdCone.png
Bird, reptilian, and monotreme cone cells

The difference in the signals received from the three cone types allows the brain to perceive a continuous range of colors through the opponent process of color vision. Rod cells have a peak sensitivity at 498 nm, roughly halfway between the peak sensitivities of the S and M cones.

All of the receptors contain the protein photopsin. Variations in its conformation cause differences in the optimum wavelengths absorbed.

The color yellow, for example, is perceived when the L cones are stimulated slightly more than the M cones, and the color red is perceived when the L cones are stimulated significantly more than the M cones. Similarly, blue and violet hues are perceived when the S receptor is stimulated more. S Cones are most sensitive to light at wavelengths around 420 nm. At moderate to bright light levels where the cones function, the eye is more sensitive to yellowish-green light than other colors because this stimulates the two most common (M and L) of the three kinds of cones almost equally. At lower light levels, where only the rod cells function, the sensitivity is greatest at a blueish-green wavelength.

Cones also tend to possess a significantly elevated visual acuity because each cone cell has a lone connection to the optic nerve, therefore, the cones have an easier time telling that two stimuli are isolated. Separate connectivity is established in the inner plexiform layer so that each connection is parallel. [13]

The response of cone cells to light is also directionally nonuniform, peaking at a direction that receives light from the center of the pupil; this effect is known as the Stiles–Crawford effect.

S cones may play a role in the regulation of the circadian system and the secretion of melatonin, but this role is not clear yet. Any potential role of the S cones in the circadian system would be secondary to the better established role of melanopsin (see also Intrinsically photosensitive retinal ganglion cell). [14]

Color afterimage

Sensitivity to a prolonged stimulation tends to decline over time, leading to neural adaptation. An interesting effect occurs when staring at a particular color for a minute or so. Such action leads to an exhaustion of the cone cells that respond to that color – resulting in the afterimage. This vivid color aftereffect can last for a minute or more. [15]

Associated diseases

See also

List of distinct cell types in the adult human body

Related Research Articles

<span class="mw-page-title-main">Color blindness</span> Decreased ability to see color or color differences

Color blindness or color vision deficiency (CVD) is the decreased ability to see color or differences in color. The severity of color blindness ranges from mostly unnoticeable to full absence of color perception. Color blindness is usually an inherited problem or variation in the functionality of one or more of the three classes of cone cells in the retina, which mediate color vision. The most common form is caused by a genetic condition called congenital red–green color blindness, which affects up to 1 in 12 males (8%) and 1 in 200 females (0.5%). The condition is more prevalent in males, because the opsin genes responsible are located on the X chromosome. Rarer genetic conditions causing color blindness include congenital blue–yellow color blindness, blue cone monochromacy, and achromatopsia. Color blindness can also result from physical or chemical damage to the eye, the optic nerve, parts of the brain, or from medication toxicity. Color vision also naturally degrades in old age.

<span class="mw-page-title-main">Visible spectrum</span> Portion of the electromagnetic spectrum that is visible to the human eye

The visible spectrum is the band of the electromagnetic spectrum that is visible to the human eye. Electromagnetic radiation in this range of wavelengths is called visible light . The optical spectrum is sometimes considered to be the same as the visible spectrum, but some authors define the term more broadly, to include the ultraviolet and infrared parts of the electromagnetic spectrum as well, known collectively as optical radiation.

<span class="mw-page-title-main">Color vision</span> Ability to perceive differences in light frequency

Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different frequencies independently of light intensity.

<span class="mw-page-title-main">Photoreceptor cell</span> Type of neuroepithelial cell

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 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.

<span class="mw-page-title-main">Tetrachromacy</span> Type of color vision with four types of cone cells

Tetrachromacy is the condition of possessing four independent channels for conveying color information, or possessing four types of cone cell in the eye. Organisms with tetrachromacy are called tetrachromats.

<span class="mw-page-title-main">Trichromacy</span> Possessing of three independent channels for conveying color information

Trichromacy or trichromatism is the possession of three independent channels for conveying color information, derived from the three different types of cone cells in the eye. Organisms with trichromacy are called trichromats.

<span class="mw-page-title-main">Rod cell</span> Photoreceptor cells that can function in lower light better than cone cells

Rod cells are photoreceptor cells in the retina of the eye that can function in lower light better than the other type of visual photoreceptor, cone cells. Rods are usually found concentrated at the outer edges of the retina and are used in peripheral vision. On average, there are approximately 92 million rod cells in the human retina. Rod cells are more sensitive than cone cells and are almost entirely responsible for night vision. However, rods have little role in color vision, which is the main reason why colors are much less apparent in dim light.

In visual physiology, adaptation is the ability of the retina of the eye to adjust to various levels of light. Natural night vision, or scotopic vision, is the ability to see under low-light conditions. In humans, rod cells are exclusively responsible for night vision, as cone cells are only able to function at higher illumination levels. Night vision is of lower quality than day vision because it is limited in resolution 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 dark adaptation 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 photopigments to increase retinal sensitivity. Light adaptation, in contrast, works very quickly, within seconds.

Dichromacy is the state of having two types of functioning photoreceptors, called cone cells, in the eyes. Organisms with dichromacy are called dichromats. Dichromats require only two primary colors to be able to represent their visible gamut. By comparison, trichromats need three primary colors, and tetrachromats need four. Likewise, every color in a dichromat's gamut can be evoked with monochromatic light. By comparison, every color in a trichromat's gamut can be evoked with a combination of monochromatic light and white light.

<span class="mw-page-title-main">Monochromacy</span> Type of color vision

Monochromacy is the ability of organisms to perceive only light intensity without respect to spectral composition. Organisms with monochromacy lack color vision and can only see in shades of grey ranging from black to white. Organisms with monochromacy are called monochromats. Many mammals, such as cetaceans, the owl monkey and the Australian sea lion are monochromats. In humans, monochromacy is one among several other symptoms of severe inherited or acquired diseases, including achromatopsia or blue cone monochromacy, together affecting about 1 in 30,000 people.

<span class="mw-page-title-main">Fovea centralis</span> Small pit in the retina of the eye responsible for all central vision

The fovea centralis is a small, central pit composed of closely packed cones in the eye. It is located in the center of the macula lutea of the retina.

<span class="mw-page-title-main">Melanopsin</span> Mammalian protein found in Homo sapiens

Melanopsin is a type of photopigment belonging to a larger family of light-sensitive retinal proteins called opsins and encoded by the gene Opn4. In the mammalian retina, there are two additional categories of opsins, both involved in the formation of visual images: rhodopsin and photopsin in the rod and cone photoreceptor cells, respectively.

In the study of visual perception, scotopic vision is the vision of the eye under low-light conditions. The term comes from the Greek skotos, meaning 'darkness', and -opia, meaning 'a condition of sight'. In the human eye, cone cells are nonfunctional in low visible light. Scotopic vision is produced exclusively through rod cells, which are most sensitive to wavelengths of around 498 nm (blue-green) and are insensitive to wavelengths longer than about 640 nm. Under scotopic conditions, light incident on the retina is not encoded in terms of the spectral power distribution. Higher visual perception occurs under scotopic vision as it does under photopic vision.

<span class="mw-page-title-main">OPN1LW</span> Protein-coding gene in humans

OPN1LW is a gene on the X chromosome that encodes for long wave sensitive (LWS) opsin, or red cone photopigment. The OPN1LW gene provides instructions for making an opsin pigment that is more sensitive to light in the yellow/orange part of the visible spectrum. The gene contains 6 exons with variability that induces shifts in the spectral range. OPN1LW is subject to homologous recombination with OPN1MW, as the two have very similar sequences. These recombinations can lead to various vision problems, such as red-green colourblindness and blue monochromacy. The protein encoded is a G-protein coupled receptor with embedded 11-cis-retinal, whose light excitation causes a cis-trans conformational change that begins the process of chemical signalling to the brain.

<span class="mw-page-title-main">Evolution of color vision in primates</span> Loss and regain of colour vision during the evolution of primates

The evolution of color vision in primates is highly unusual compared to most eutherian mammals. A remote vertebrate ancestor of primates possessed tetrachromacy, but nocturnal, warm-blooded, mammalian ancestors lost two of four cones in the retina at the time of dinosaurs. Most teleost fish, reptiles and birds are therefore tetrachromatic while most mammals are strictly dichromats, the exceptions being some primates and marsupials, who are trichromats, and many marine mammals, who are monochromats.

<span class="mw-page-title-main">Mammalian eye</span>

Mammals normally have a pair of eyes. Although mammalian vision is not as excellent as bird vision, it is at least dichromatic for most of mammalian species, with certain families possessing a trichromatic color perception.

G-protein-coupled receptor kinase 7 is a serine/threonine-specific protein kinase involved in phototransduction. This enzyme catalyses the phosphorylation of cone (color) photopsins in retinal cones during high acuity color vision primarily in the fovea.

Blue cone monochromacy (BCM) is an inherited eye disease that causes severe color blindness, poor visual acuity, nystagmus, hemeralopia, and photophobia due to the absence of functional red (L) and green (M) cone photoreceptor cells in the retina. BCM is a recessive X-linked disease and almost exclusively affects XY karyotypes.

<span class="mw-page-title-main">Congenital red–green color blindness</span> Most common genetic condition leading to color blindness

Congenital red–green color blindness is an inherited condition that is the root cause of the majority of cases of color blindness. It has no significant symptoms aside from its minor to moderate effect on color vision. It is caused by variation in the functionality of the red and/or green opsin proteins, which are the photosensitive pigment in the cone cells of the retina, which mediate color vision. Males are more likely to inherit red–green color blindness than females, because the genes for the relevant opsins are on the X chromosome. Screening for congenital red–green color blindness is typically performed with the Ishihara or similar color vision test. It is a lifelong condition, and has no known cure or treatment.

<span class="mw-page-title-main">Vertebrate visual opsin</span>

Vertebrate visual opsins are a subclass of ciliary opsins and mediate vision in vertebrates. They include the opsins in human rod and cone cells. They are often abbreviated to opsin, as they were the first opsins discovered and are still the most widely studied opsins.

References

  1. "The Rods and Cones of the Human Eye". HyperPhysics Concepts - Georgia State University.
  2. 1 2 3 Kandel, E.R.; Schwartz, J.H; Jessell, T. M. (2000). Principles of Neural Science (4th ed.). New York: McGraw-Hill. pp.  507–513. ISBN   9780838577011.
  3. Schacter, Gilbert, Wegner, "Psychology", New York: Worth Publishers,2009.
  4. Jameson, K. A.; Highnote, S. M. & Wasserman, L. M. (2001). "Richer color experience in observers with multiple photopigment opsin genes" (PDF). Psychonomic Bulletin and Review. 8 (2): 244–261. doi: 10.3758/BF03196159 . PMID   11495112. S2CID   2389566.
  5. "You won't believe your eyes: The mysteries of sight revealed". The Independent . 7 March 2007. Archived from the original on 6 July 2008. Retrieved 22 August 2009. Equipped with four receptors instead of three, Mrs M - an English social worker, and the first known human "tetrachromat" - sees rare subtleties of colour.
  6. Mark Roth (September 13, 2006). "Some women may see 100,000,000 colors, thanks to their genes". Pittsburgh Post-Gazette. Archived from the original on November 8, 2006. Retrieved August 22, 2009. A tetrachromat is a woman who can see four distinct ranges of color, instead of the three that most of us live with.
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  8. Wyszecki, Günther; Stiles, W.S. (1981). Color Science: Concepts and Methods, Quantitative Data and Formulae (2nd ed.). New York: Wiley Series in Pure and Applied Optics. ISBN   978-0-471-02106-3.
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  11. Foundations of Vision, Brian A. Wandell
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  13. Strettoi, E; Novelli, E; Mazzoni, F; Barone, I; Damiani, D (Jul 2010). "Complexity of retinal cone bipolar cells". Progress in Retinal and Eye Research. 29 (4): 272–83. doi:10.1016/j.preteyeres.2010.03.005. PMC   2878852 . PMID   20362067.
  14. Soca, R (Feb 13, 2021). "S-cones and the circadian system". Keldik. Archived from the original on 2021-02-14.
  15. Schacter, Daniel L. Psychology: the second edition. Chapter 4.9.
  16. 1 2 3 Aboshiha, Jonathan; Dubis, Adam M; Carroll, Joseph; Hardcastle, Alison J; Michaelides, Michel (January 2016). "The cone dysfunction syndromes: Table 1". British Journal of Ophthalmology. 100 (1): 115–121. doi: 10.1136/bjophthalmol-2014-306505 . PMC   4717370 . PMID   25770143.