Melanism
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Melanism is the
congenital A birth defect is an abnormal condition that is present at childbirth, birth, regardless of its cause. Birth defects may result in disability, disabilities that may be physical disability, physical, intellectual disability, intellectual, or dev ...
excess of
melanin Melanin (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes. There are ...
in an organism resulting in dark
pigment A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...
. Pseudomelanism, also called abundism, is another variant of pigmentation, identifiable by dark spots or enlarged stripes, which cover a large part of the body of the animal, making it appear melanistic. The morbid deposition of black matter, often of a malignant character causing pigmented
tumor A neoplasm () is a type of abnormal and excessive growth of tissue. The process that occurs to form or produce a neoplasm is called neoplasia. The growth of a neoplasm is uncoordinated with that of the normal surrounding tissue, and persists ...
s, is called melanosis.


Adaptation

Melanism related to the process of
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
is called adaptive. Most commonly, dark individuals become fitter to survive and reproduce in their environment as they are better camouflaged. This makes some species less conspicuous to predators, while others, such as leopards, use it as a foraging advantage during night hunting. Typically, adaptive melanism is
heritable Heredity, also called inheritance or biological inheritance, is the passing on of Phenotypic trait, traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cell (biology), cells or orga ...
: A dominant
allele An allele is a variant of the sequence of nucleotides at a particular location, or Locus (genetics), locus, on a DNA molecule. Alleles can differ at a single position through Single-nucleotide polymorphism, single nucleotide polymorphisms (SNP), ...
, which is entirely or nearly entirely expressed in the
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
, is responsible for the excessive amount of melanin. By contrast, adaptive melanism associated with Batesian mimicry in Zelandoperla fenestrata stoneflies is controlled by a recessive allele at the ebony locus. Adaptive melanism has been shown to occur in a variety of animals, including mammals such as
squirrel Squirrels are members of the family Sciuridae (), a family that includes small or medium-sized rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrel ...
s, many
cats The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the ...
and
canid Canidae (; from Latin, ''canis'', "dog") is a family (biology), biological family of caniform carnivorans, constituting a clade. A member of this family is also called a canid (). The family includes three subfamily, subfamilies: the Caninae, a ...
s, and
coral snake Coral snakes are a large group of elapid snakes that can be divided into two distinct groups, the Old World coral snakes and New World coral snakes. There are 27 species of Old World coral snakes, in three genera ('' Calliophis'', '' Hemibungar ...
s. Adaptive melanism can lead to the creation of morphs, a notable example being the
peppered moth The peppered moth (''Biston betularia'') is a temperate species of Nocturnality, night-flying moth. It is mostly found in the northern hemisphere in places like Asia, Europe and North America. Peppered moth evolution is an example of populatio ...
, whose evolutionary history in the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Northwestern Europe, off the coast of European mainland, the continental mainland. It comprises England, Scotlan ...
is offered as a classic instructional tool for teaching the principles of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
. A more replicated example of human-induced shifts in melanism has arisen from repeated selection against melanic Zelandoperla fenestrata stonefly phenotypes following widespread deforestation in New Zealand.


Industrial melanism

Industrial melanism is an
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
ary effect in insects such as the peppered moth, ''
Biston betularia The peppered moth (''Biston betularia'') is a temperate species of night-flying moth. It is mostly found in the northern hemisphere in places like Asia, Europe and North America. Peppered moth evolution is an example of population genetics a ...
'' in areas subject to industrial pollution. Darker pigmented individuals are favored by
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
, apparently because they are better
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
d against polluted backgrounds. When pollution was later reduced, lighter forms regained the advantage and melanism became less frequent.Grant, B. S., Wiseman L. L., 2002. Recent history of melanism in American peppered moths. Journal of Heredity 93:86-90.Grant, B. S., Cook, A. D., Clarke, C. A., & Owen, D. F. (1998). Geographic and temporal variation in the incidence of melanism in peppered moth populations in America and Britain. ''Journal of Heredity'', 89(5), 465–471. Other explanations have been proposed, such as that the melanin pigment enhances function of immune defences, or a thermal advantage from the darker coloration.Mikkola, K., Albrecht, A., 1988. The melanism of ''Adalia-bipunctata'' around the Gulf of Finland as an industrial phenomenon (Coleoptera, Coccinellidae). Annales Zoologici Fennici 25:177–85.Muggleton, J., Lonsdale, D., Benham, B. R., 1975. Melanism in ''Adalia-bipunctata L'' (ColCoccinellidae) and its relationship to atmospheric pollution. Journal of Applied Ecology 2:451–464.De Jong, P. W., Verhoog, M. D., Brakefield, P. M., 1992. Sperm competition and melanic polymorphism in the 2-spot ladybird, ''Adalla bipunctata'' (Coleoptera, Coccinellidae). Journal of Heredity 70:172–178.


In cats

Melanistic coat coloration occurs as a common polymorphism in 11 of 37
felid Felidae ( ) is the Family (biology), family of mammals in the Order (biology), order Carnivora colloquially referred to as cats. A member of this family is also called a felid ( ). The 41 extant taxon, extant Felidae species exhibit the gre ...
species and reaches high population frequency in some cases but never achieves complete fixation. The
black panther A black panther is the Melanism, melanistic colour variant of the leopard (''Panthera pardus'') and the jaguar (''Panthera onca''). Black panthers of both species have excess black pigments, but their typical Rosette (zoology), rosettes are al ...
, a melanistic
leopard The leopard (''Panthera pardus'') is one of the five extant cat species in the genus ''Panthera''. It has a pale yellowish to dark golden fur with dark spots grouped in rosettes. Its body is slender and muscular reaching a length of with a ...
, is common in the equatorial rainforest of Malaya and the
tropical rainforest Tropical rainforests are dense and warm rainforests with high rainfall typically found between 10° north and south of the Equator. They are a subset of the tropical forest biome that occurs roughly within the 28° latitudes (in the torrid zo ...
on the slopes of some African mountains, such as Mount Kenya. The
serval The serval (''Leptailurus serval'') is a wild small cat native to Africa. It is widespread in sub-Saharan countries, where it inhabits grasslands, wetlands, moorlands and bamboo thickets. Across its range, it occurs in protected areas, and ...
also has melanistic forms in certain areas of
East Africa East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...
. In the
jaguarundi The jaguarundi (''Herpailurus yagouaroundi''; or ) is a wild felidae, cat native to the Americas. Its range extends from central Argentina in the south to northern Mexico, through Central America, Central and South America east of the Andes. T ...
, coloration varies from dark brown and gray to light reddish. Melanic forms of
jaguar The jaguar (''Panthera onca'') is a large felidae, cat species and the only extant taxon, living member of the genus ''Panthera'' that is native to the Americas. With a body length of up to and a weight of up to , it is the biggest cat spe ...
are common in certain parts of
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. In 1938 and 1940, two melanistic
bobcat The bobcat (''Lynx rufus''), also known as the wildcat, bay lynx, or red lynx, is one of the four extant species within the medium-sized wild cat genus '' Lynx''. Native to North America, it ranges from southern Canada through most of the c ...
s were trapped alive in sub-tropical
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
. In 2003, the dominant mode of inheritance of melanism in jaguars was confirmed by performing
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
-transmission analysis in a 116-individual captive pedigree. Melanistic animals were found to carry at least one copy of a mutant '' MC1R'' sequence
allele An allele is a variant of the sequence of nucleotides at a particular location, or Locus (genetics), locus, on a DNA molecule. Alleles can differ at a single position through Single-nucleotide polymorphism, single nucleotide polymorphisms (SNP), ...
, bearing a 15-
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
inframe deletion. Ten unrelated melanistic jaguars were either
homozygous Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mos ...
or
heterozygous Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mos ...
for this allele. A 24-base pair deletion causes the incompletely dominant allele for melanism in the jaguarundi. Sequencing of the agouti signalling peptide in the
agouti gene Agouti-signaling protein is a protein that in humans is encoded by the ASIP gene. It is responsible for the distribution of melanin pigment in mammals. Agouti interacts with the melanocortin 1 receptor to determine whether the melanocyte (pigment ...
coding region revealed a 2-base pair deletion in black
domestic cat The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small Domestication, domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have sh ...
s. These variants were absent in melanistic individuals of Geoffroy's cat, oncilla, pampas cat and Asian golden cat, suggesting that melanism arose independently at least four times in the cat family. Melanism in leopards is inherited as a Mendelian, monogenic recessive trait relative to the spotted form. Pairings of black animals have a significantly smaller litter size than other possible pairings. Between January 1996 and March 2009, Indochinese leopards were photographed at 16 sites in the
Malay Peninsula The Malay Peninsula is located in Mainland Southeast Asia. The landmass runs approximately north–south, and at its terminus, it is the southernmost point of the Asian continental mainland. The area contains Peninsular Malaysia, Southern Tha ...
in a sampling effort of more than 1000 trap nights. Of 445 photographs of melanistic leopards, 410 were taken south of the
Kra Isthmus The Kra Isthmus (, ; ), also called the Isthmus of Kra in Thailand, is the narrowest part of the Malay Peninsula. The western part of the isthmus belongs to Ranong Province and the eastern part to Chumphon Province, both in Southern Thailan ...
, where the non-melanistic morph was never photographed. These data suggest the near fixation of the dark allele in the region. The expected time to fixation of this recessive allele due to
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
alone ranged from about 1,100 years to about 100,000 years. Melanism in leopards has been hypothesized to be causally associated with a selective advantage for ambush. Other theories are that genes for melanism in felines may provide resistance to viral infections, or a high-altitude adaptation, since black fur absorbs more light for warmth.


In birds

In April 2015, an extremely rare black flamingo was spotted on the Mediterranean island of
Cyprus Cyprus (), officially the Republic of Cyprus, is an island country in the eastern Mediterranean Sea. Situated in West Asia, its cultural identity and geopolitical orientation are overwhelmingly Southeast European. Cyprus is the List of isl ...
. The chicken breeds Silkie and Ayam Cemani commonly exhibit this trait. Ayam Cemani is an uncommon and relatively modern breed of chicken from Indonesia. They have a dominant gene that causes hyperpigmentation (Fibromelanosis), making the chicken entirely black; including feathers, beak, and internal organs. Melanistic common pheasants are sometimes intentionally released as game birds. Melanism in feral
rock dove The rock dove (''Columba livia''), also sometimes known as "rock pigeon" or "common pigeon", is a member of the bird family Columbidae (doves and pigeons). In common usage, it is often simply referred to as the "pigeon", although the rock dov ...
s is quite common, especially if the species is abundant in the area. The amount of pigmentation varies, from a slightly darker pigmentation in the pigeon’s wings, to being almost completely black.


In amphibians

The alpine salamander, ''Salamandra atra'', has one subspecies (''S. atra atra'') that is completely black. The pigment comes from a specific cell called a melanophore, which produce the compound melanin. There are four other subspecies of this salamander, and they have varying levels of melanin pigmentation. The subspecies have yellow spots in different concentrations or proportions. The pigment-producing cells that contribute to the yellow spots of some sub-species are called xanthophores. It appears that the fully-black phenotypes do not ever develop these xanthophores. Alpine salamanders produce a toxin from their skin, and both fully melanistic, black salamanders and spotted individuals produce the compound. Studies done that traced DNA histories have suggested that the original alpine salamander phenotype was black with some yellow spots, meaning that the fully black color evolved over time and was thus selected for over many generations.


In humans

Melanism, meaning a mutation that results in completely dark skin, does not exist in humans. In humans, the amount of melanin is determined by three dominant alleles (AABBCC), and different ethnicities have varying amounts. Melanin is the primary determinant of the degree of skin pigmentation and protects the body from harmful
ultraviolet radiation Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of t ...
. The same ultraviolet radiation is essential for the synthesis of
vitamin D Vitamin D is a group of structurally related, fat-soluble compounds responsible for increasing intestinal absorption of calcium, magnesium, and phosphate, along with numerous other biological functions. In humans, the most important compo ...
in skin, so lighter colored skin – less melanin – is an adaptation related to the prehistoric movement of humans away from equatorial regions, as there is less exposure to sunlight at higher latitudes. People from
Sub-Saharan Africa Sub-Saharan Africa is the area and regions of the continent of Africa that lie south of the Sahara. These include Central Africa, East Africa, Southern Africa, and West Africa. Geopolitically, in addition to the list of sovereign states and ...
,
South Asia South Asia is the southern Subregion#Asia, subregion of Asia that is defined in both geographical and Ethnicity, ethnic-Culture, cultural terms. South Asia, with a population of 2.04 billion, contains a quarter (25%) of the world's populatio ...
,
Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
, Australia, and
Melanesia Melanesia (, ) is a subregion of Oceania in the southwestern Pacific Ocean. It extends from New Guinea in the west to the Fiji Islands in the east, and includes the Arafura Sea. The region includes the four independent countries of Fiji, Vanu ...
may have very dark skin, but this is not melanism.


Peutz–Jeghers syndrome

This rare
genetic disorder A genetic disorder is a health problem caused by one or more abnormalities in the genome. It can be caused by a mutation in a single gene (monogenic) or multiple genes (polygenic) or by a chromosome abnormality. Although polygenic disorders ...
is characterized by the development of macules with
hyperpigmentation Hyperpigmentation, also known as the dark spots or circles on the skin, is the darkening of an area of Human skin, skin or nail (anatomy), nails caused by increased melanin. Causes Hyperpigmentation can be caused by sun damage, inflammation, or ...
on the lips and
oral mucosa The oral mucosa is the mucous membrane lining the inside of the mouth. It comprises stratified squamous epithelium, termed "oral epithelium", and an underlying connective tissue termed '' lamina propria''. The oral cavity has sometimes been des ...
( melanosis), as well as benign polyps in the gastrointestinal tract.


Socio-politics

The term ''melanism'' has been used on
Usenet Usenet (), a portmanteau of User's Network, is a worldwide distributed discussion system available on computers. It was developed from the general-purpose UUCP, Unix-to-Unix Copy (UUCP) dial-up network architecture. Tom Truscott and Jim Elli ...
, internet forums and
blog A blog (a Clipping (morphology), truncation of "weblog") is an informational website consisting of discrete, often informal diary-style text entries also known as posts. Posts are typically displayed in Reverse chronology, reverse chronologic ...
s to mean an
African-American African Americans, also known as Black Americans and formerly also called Afro-Americans, are an American racial and ethnic group that consists of Americans who have total or partial ancestry from any of the Black racial groups of Africa. ...
social movement A social movement is either a loosely or carefully organized effort by a large group of people to achieve a particular goal, typically a Social issue, social or Political movement, political one. This may be to carry out a social change, or to re ...
holding that dark-skinned humans are the original people from which those of other skin color originate. The term ''melanism'' has been used in this context as early as the mid-1990s and was promoted by some Afrocentrists, such as Frances Cress Welsing.


See also

*
Albinism Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and reddish pink or blue eyes. Individuals with the condition are referred to as albinos. Varied use and interpretation of ...
** Albino and white squirrels * Amelanism, lack of melanism * Black squirrel * Erythrism, reddish pigmentation * Isabellinism, lowered melanism * Heterochromia iridum * Leucism, a partial loss of pigmentation that results in animals with pale or white skin, hair and/or feathers * Melanosis, hyperpigmentation via increased melanin ** Ocular melanosis *
Peutz–Jeghers syndrome Peutz–Jeghers syndrome (often abbreviated PJS) is an dominance (genetics), autosomal dominant genetic disorder characterized by the development of benign hamartomatous Polyp (medicine), polyps in the human gastrointestinal tract, gastrointestinal ...
, dark patches on the lips etc. * Piebaldism, patchy absence of melanin-producing cells *
Vitiligo Vitiligo (, ) is a chronic autoimmune disorder that causes patches of skin to lose pigment or color. The cause of vitiligo is unknown, but it may be related to immune system changes, genetic factors, stress, or sun exposure, and susceptibili ...
, a skin condition which causes areas of the skin to lose its colour * Xanthochromism, an unusual yellow colouration in animals * Zelandoperla fenestrata, a stonefly exhibiting a
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, who worked on butt ...
melanic polymorphism


References


Bibliography

* * *
Melanism and disease resistance in insects
* Fryer, G. 2013. How should the history of industrial melanism in moths be interpreted? ''The Linnean''. 29 (2): 15 - 22. {{Portal bar, Environment, Ecology, Earth sciences, Evolutionary biology Genetic disorders with no OMIM Disturbances of pigmentation Dermatologic terminology Animal coat colors