Monochromats
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Monochromacy (from
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
''mono'', meaning "one" and ''chromo'', meaning "color") is the ability of organisms to perceive only light intensity without respect to spectral composition. Organisms with monochromacy lack
color vision Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different frequencies independently of light intensity. Color perception is a part of the larger visual system and is mediated by a co ...
and can only see in shades of grey ranging from black to white. Organisms with monochromacy are called monochromats. Many mammals, such as
cetacean Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
s, the
owl monkey Night monkeys, also known as owl monkeys or douroucoulis (), are nocturnal New World monkeys of the genus ''Aotus'', the only member of the family Aotidae (). The genus comprises eleven species which are found across Panama and much of South Am ...
and the
Australian sea lion The Australian sea lion (''Neophoca cinerea''), also known as the Australian sea-lion or Australian sealion, is a species of sea lion that is the only endemic pinniped in Australia. It is currently Monotypic taxon, monotypic in the genus ''Neopho ...
are monochromats. In
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s, monochromacy is one among several other symptoms of severe inherited or acquired diseases, including
achromatopsia Achromatopsia, also known as rod monochromacy, is a medical syndrome that exhibits symptoms relating to five conditions, most notably monochromacy. Historically, the name referred to monochromacy in general, but now typically refers only to an aut ...
or
blue cone monochromacy 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 re ...
, together affecting about 1 in 30,000 people.


Humans

Human vision relies on a
duplex retina A duplex retina is a retina consisting of both rod cells and cone cells, which are the photoreceptor cells for two parallel but mostly separate visual systems. The rods enable the scotopic visual system, which is active in dim light. The cones en ...
, comprising two types of
photoreceptor 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 (visible electromagnetic radiation ...
s. Rods are primarily responsible for dim-light
scotopic vision In the study of visual perception, scotopic vision (or scotopia) 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, c ...
and
cone In geometry, a cone is a three-dimensional figure that tapers smoothly from a flat base (typically a circle) to a point not contained in the base, called the '' apex'' or '' vertex''. A cone is formed by a set of line segments, half-lines ...
s are primarily responsible for day-light
photopic vision Photopic vision is the vision of the eye under well-lit conditions (luminance levels from 10 to 108  cd/m2). In humans and many other animals, photopic vision allows color perception, mediated by cone cells, and a significantly higher vis ...
. For all known
vertebrates Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
, scotopic vision is monochromatic, since there is typically only one class of rod cell. However, the presence of multiple cone classes contributing to photopic vision enables
color vision Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different frequencies independently of light intensity. Color perception is a part of the larger visual system and is mediated by a co ...
during daytime conditions. Most humans have three classes of cones, each with a different class of
opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become retinylidene proteins, but are usually still called opsins regardless. Most pro ...
. These three opsins have different spectral sensitivities, which is a prerequisite for
trichromacy 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. The normal expl ...
. An alteration of any of these three cone opsins can lead to
colorblindness Color blindness, color vision deficiency (CVD) or color deficiency 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 bl ...
. #
Anomalous trichromacy Color blindness, color vision deficiency (CVD) or color deficiency 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 bl ...
, when all three cones are functional, but one or more is altered in its spectral sensitivity. #
Dichromacy Dichromacy (from Greek ''di'', meaning "two" and ''chromo'', meaning "color") is the state of having two types of functioning photoreceptors, called cone cells, in the eyes. Organisms with dichromacy are called dichromats. Dichromats requir ...
, when one of the cones is non-functional and one of the red-green or blue-yellow opponent channels are fully disabled. # Cone monochromacy, when two of the cones are non-functional and both chromatic opponent channels are disabled. Vision is reduced to blacks, whites, and greys. # Rod monochromacy (
Achromatopsia Achromatopsia, also known as rod monochromacy, is a medical syndrome that exhibits symptoms relating to five conditions, most notably monochromacy. Historically, the name referred to monochromacy in general, but now typically refers only to an aut ...
), when all three of the cones are non-functional and therefore photopic vision (and therefore color vision) is disabled. Monochromacy of photopic vision is a symptom of both Cone Monochromacy and Rod Monochromacy, so these two conditions are typically referred to collectively as monochromacy.


Rod monochromacy

Rod monochromacy (RM), also called congenital complete achromatopsia or total color blindness, is a rare and extremely severe form of an
autosomal recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the Phenotype, effect of a different variant of the same gene on Homologous chromosome, the other copy of the chromosome. The firs ...
ly inherited retinal disorder resulting in severe visual handicap. People with RM have a reduced visual acuity, (usually about 0.1 or 20/200), have total color blindness, photo-aversion and
nystagmus Nystagmus is a condition of involuntary (or voluntary, in some cases) Eye movement (sensory), eye movement. People can be born with it but more commonly acquire it in infancy or later in life. In many cases it may result in visual impairment, re ...
. The nystagmus and photo-aversion usually are present during the first months of life, and the prevalence of the disease is estimated to be 1 in 30,000 worldwide. Since patients with RM have no cone function, they lack photopic vision, relying entirely on their rods and scotopic vision, which is necessarily monochromatic. They therefore cannot see any color but only shades of grey.


Cone monochromacy

Cone monochromacy (CM) is a condition defined by the exhibition of only one class of cones. A cone monochromat can have good pattern vision at normal daylight levels, but will not be able to distinguish hues. As humans typically exhibit three classes of cones, cone monochromats can hypothetically derive their photopic vision from any one of them, leading to three categories of cone monochromats: #
Blue cone monochromacy 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 re ...
(BCM), also known as S-cone monochromacy, is an X-linked cone disease. It is a rare congenital stationary cone dysfunction syndrome, affecting less than 1 in 100,000 individuals, and is characterized by the absence of L- and M-cone function. BCM results from mutations in a single red or red–green hybrid
opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become retinylidene proteins, but are usually still called opsins regardless. Most pro ...
gene, mutations in both the red and the green opsin genes or deletions within the adjacent LCR (
locus control region A locus control region (LCR) is a long-range cis-regulatory element that enhances expression of linked genes at distal chromatin sites. It functions in a copy number-dependent manner and is tissue-specific, as seen in the selective expression of ...
) on the X chromosome. #Green cone monochromacy (GCM), also known as M-cone monochromacy, is a condition where the blue and red cones are absent in the
fovea Fovea () (Latin for "pit"; plural foveae ) is a term in anatomy. It refers to a pit or depression in a structure. Human anatomy *Fovea centralis of the retina * Fovea buccalis or dimple * Fovea of the femoral head * Trochlear fovea of the fr ...
. The prevalence of this type of monochromacy is estimated to be less than 1 in 1 million. #Red cone monochromacy (RCM), also known as L-cone monochromacy, is a condition where the blue and green cones are absent in the fovea. Like GCM, the prevalence of RCM is also estimated at less than 1 in 1 million. Cone Monochromats with normal rod function can sometimes exhibit mild color vision due to conditional dichromacy. In mesopic conditions, both rods and cones are active and opponent interactions between the cones and rods can afford slight color vision. According to
Jay Neitz Jay Neitz (born 1953) is an American professor of ophthalmology and a color vision researcher at the University of Washington in Seattle, Washington. Education and career Neitz grew up in Montana. He attended San Jose State University for his u ...
, a color vision researcher at the
University of Washington The University of Washington (UW and informally U-Dub or U Dub) is a public research university in Seattle, Washington, United States. Founded in 1861, the University of Washington is one of the oldest universities on the West Coast of the Uni ...
, each of the three standard color-detecting cones in the retina of
trichromat 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. The normal expl ...
s can detect approximately 100 gradations of color. The brain can process the combinations of these three values so that the average human can distinguish about one million colors. Therefore, a monochromat would be able to distinguish about 100 colors.


Mammals

Until the 1960s, popular belief held that most
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s outside of
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s were monochromats. In the last half-century, however, a focus on behavioral and
genetic testing Genetic testing, also known as DNA testing, is used to identify changes in DNA sequence or chromosome structure. Genetic testing can also include measuring the results of genetic changes, such as RNA analysis as an output of gene expression, or ...
of mammals has accumulated extensive evidence of at least
dichromat Dichromacy (from Greek ''di'', meaning "two" and ''chromo'', meaning "color") is the state of having two types of functioning photoreceptors, called cone cells, in the eyes. Organisms with dichromacy are called dichromats. Dichromats require ...
ic color vision in a number of mammalian
orders Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * H ...
. Mammals are now usually assumed to be dichromats (possessing S- and L-cones), with monochromats viewed as the exceptions. Two mammalian orders containing
marine mammal Marine mammals are mammals that rely on marine ecosystems for their existence. They include animals such as cetaceans, pinnipeds, sirenians, sea otters and polar bears. They are an informal group, unified only by their reliance on marine enviro ...
s exhibit monochromatic vision: *
Pinniped Pinnipeds (pronounced ), commonly known as seals, are a widely range (biology), distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammals. They comprise the extant taxon, extant families Odobenidae (whose onl ...
s (including seals, sea lions and walruses) *
Cetacean Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
s (including dolphins and whales) Unlike the trichromacy exhibited in most primates,
Owl monkey Night monkeys, also known as owl monkeys or douroucoulis (), are nocturnal New World monkeys of the genus ''Aotus'', the only member of the family Aotidae (). The genus comprises eleven species which are found across Panama and much of South Am ...
s (genus ''Aotus'') are also monochromats. Several members of the family
Procyonidae Procyonidae ( ) is a New World family of the order Carnivora. It includes the raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of environments and are generally omnivorous. Ch ...
(
raccoon The raccoon ( or , ''Procyon lotor''), sometimes called the North American, northern or common raccoon (also spelled racoon) to distinguish it from Procyonina, other species of raccoon, is a mammal native to North America. It is the largest ...
,
crab-eating raccoon The crab-eating raccoon, southern raccoon, or South American raccoon (''Procyon cancrivorus'') is a species of raccoon native to marshy and jungle areas of Central and South America (including Trinidad and Tobago). It is found from Costa Rica so ...
and
kinkajou The kinkajou ( /ˈkɪŋkədʒuː/ ''KING-kə-joo''; ''Potos flavus'') is a tropical rainforest mammal of the family Procyonidae related to olingos, coatis, raccoons, and the ringtail and cacomistle. It is the only member of the genus ''Pot ...
) and a few
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s have been demonstrated as cone monochromats, having lost functionality of the S-cone (retaining the L-cone). The light available in an animal's habitat is a significant determiner of a mammal's color vision. Marine,
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
or
burrow file:Chipmunk-burrow (exits).jpg, An eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of Animal lo ...
ing mammals, which experience less light, have less
evolutionary pressure Evolutionary pressure, selective pressure or selection pressure is exerted by factors that reduce or increase reproductive success in a portion of a population, driving natural selection. It is a quantitative description of the amount of change o ...
to preserve dichromacy, so often evolve monochromacy. A recent study using through PCR analysis of genes
OPN1SW Blue-sensitive opsin is a protein that in humans is encoded by the ''OPN1SW'' gene. The OPN1SW gene provides instructions for making a protein that is essential for normal color vision. This protein is found in the retina, which is the light-sensi ...
,
OPN1LW 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 vis ...
, and
PDE6C Cone cGMP-specific 3',5'-cyclic phosphodiesterase subunit alpha' is an enzyme that in humans is encoded by the ''PDE6C'' gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is ...
determined that all mammals in the cohort
Xenarthra Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a superorder and major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and ...
(representing sloths, anteaters and armadillos) developed rod monochromacy through a stem ancestor.


See also

*
Achromatopsia Achromatopsia, also known as rod monochromacy, is a medical syndrome that exhibits symptoms relating to five conditions, most notably monochromacy. Historically, the name referred to monochromacy in general, but now typically refers only to an aut ...
*
Blue cone monochromacy 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 re ...
*
Cone dystrophy A cone dystrophy is an inherited ocular disorder characterized by the loss of cone cells, the photoreceptors responsible for both central and color vision. Presentation The most common symptoms of cone dystrophy are vision loss (age of onset ...
*
Dichromacy Dichromacy (from Greek ''di'', meaning "two" and ''chromo'', meaning "color") is the state of having two types of functioning photoreceptors, called cone cells, in the eyes. Organisms with dichromacy are called dichromats. Dichromats requir ...
*
Trichromacy 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. The normal expl ...
*
Tetrachromacy Tetrachromacy (from Ancient Greek ''tetra'', meaning "four" and ''chroma'', meaning "color") is the condition of possessing four independent channels for conveying color information, or possessing four types of cone cell in the eye. Organisms ...


References


External links

* * {{Color topics Visual disturbances and blindness Color vision