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Roan is a
horse coat color Horses exhibit a diverse array of coat colors and distinctive markings. A specialized vocabulary has evolved to describe them. While most horses remain the same color throughout life, a few, over the course of several years, will develop a diff ...
pattern characterized by an even mixture of colored and white hairs on the body, while the head and " points"—lower legs, mane, and tail—are mostly solid-colored. Horses with roan coats have white hairs evenly intermingled throughout any other color. The head, legs, mane, and tail have fewer scattered white hairs or none at all. The roan pattern is dominantly inherited, and is found in many
horse breed A horse breed is a selectively bred population of domesticated horses, often with pedigrees recorded in a breed registry. However, the term is sometimes used in a broader sense to define landrace animals of a common phenotype located within a lim ...
s. While the specific
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...
responsible for roan has not been exactly identified, a
DNA test 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, o ...
can determine
zygosity 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. Mo ...
for roan in several breeds. True roan is always present at birth, though it may be hard to see until after the foal coat sheds out. The coat may lighten or darken from winter to summer, but unlike the
gray Grey (more common in British English) or gray (more common in American English) is an intermediate color between black and white. It is a neutral or achromatic color, meaning literally that it is "without color", because it can be composed o ...
coat color, which also begins with intermixed white and colored hairs, roans do not become progressively lighter in color as they age. The silvering effect of mixed white and colored hairs can create coats that look bluish or pinkish.


Identification

Horses with the roan pattern have an even mixture of white and colored hairs in the coat. These interspersed white hairs are more scattered or absent on the horse's head, mane, tail, and lower legs. The unaffected color on the legs often forms a sharp, inverted "V" above the knee and hock, not seen in other roan-like coat patterns. The nonwhite background coat may be any color, as determined by unrelated genetic factors. Often, the background coat color is used in combination with the word "roan" to describe the shade of a roan horse's coat, such as bay roan or gold champagne roan, but colloquial terms also are used for some colors. The most common terms for various roan colors are: * Blue roan is loosely applied to any roan with a dark underlying coat that gives it a bluish cast. In the strictest sense, though, "blue roan" is a common synonym for a roan with a
black Black is a color which results from the absence or complete absorption of visible light. It is an achromatic color, without hue, like white and grey. It is often used symbolically or figuratively to represent darkness. Black and white have o ...
background coat. * Red roan used to include both chestnut and bay roans. In 1999, the
American Paint Horse The American Paint Horse is a breed of horse that combines both the conformational characteristics of a western stock horse with a pinto spotting pattern of white and dark coat colors. Developed from a base of spotted horses with Quarter Hors ...
Association changed its coat color descriptions; roans with a chestnut background coat are registered "red roan", while "bay roan" is its own category. The
American Quarter Horse The American Quarter Horse, or Quarter Horse, is an American breed of horse that excels at sprinting short distances. Its name is derived from its ability to outrun other horse breeds in races of a quarter mile or less; some have been clocked at s ...
Association followed suit in 2003. Previously, the term strawberry roan described the pinkish color of a light chestnut or
sorrel Sorrel (''Rumex acetosa''), also called common sorrel or garden sorrel, is a perennial herbaceous plant in the family Polygonaceae. Other names for sorrel include spinach dock and narrow-leaved dock ('dock' being a common name for the genus '' ...
roan. While less common, the term lilac roan may be applied to a dark chestnut roan, and honey roan to
palomino Palomino is a genetic color in horses, consisting of a gold coat and white mane and tail; the degree of whiteness can vary from bright white to yellow. Genetically, the palomino color is created by a single allele of a dilution gene called t ...
s or the lightest sorrels. *Bay roan replaced red roan as the term for a roan with a bay background coat. Some roan horses have more white hair than others, and even individual horses may look lighter or darker based on the season, and their coats may vary from year to year. While roan is always present at birth, the soft first coat of newborn foals may not show the white hairs well. Some roan horses get darker with age. Generally, roans appear to have more white hair when they have their short summer coats and darker when they have their winter coats. These peculiar tendencies of roans led to the Icelandic word for roan, which translates as "always changing color." Roans have other unusual characteristics. If the skin is damaged by even a very minor scrape, cut, or
brand A brand is a name, term, design, symbol or any other feature that distinguishes one seller's good or service from those of other sellers. Brands are used in business, marketing, and advertising for recognition and, importantly, to create an ...
, the coat grows back in solid-colored without any white hairs. These regions of solid-colored coat are called "corn spots" or "corn marks", and can appear even without the horse having had a visible injury. Another trait is reverse dappling; many horses develop rings of hair that appear slightly different-colored, called dapples, which often indicate good health. Usually dapples are darker than the surrounding coat, but on a roan, the dapples are lighter. File:Cornmarks.jpg, Corn marks or corn spots occur where a roan's skin has been damaged. The hair grows back in without any white. File:Bayroanlegs.JPG, The
foreleg A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial (anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead ...
s of this bay roan show the characteristic inverted "V" of dark hair not affected by roan. File:Kabayos03.jpg, This strawberry roan mare, though very light-colored, is still identifiable as a roan by the dark color of her extremities and the
brand A brand is a name, term, design, symbol or any other feature that distinguishes one seller's good or service from those of other sellers. Brands are used in business, marketing, and advertising for recognition and, importantly, to create an ...
on her hindquarters, which has grown back in without white hairs. File:Reverse-dapples.jpg, Reverse dapples on a bay roan File:900-231 Herzog Carl Alexander.jpg, This dark bay roan, painted in the 18th century carrying the
Duke of Württemberg Duke is a male title either of a monarch ruling over a duchy, or of a member of royalty, or nobility. As rulers, dukes are ranked below emperors, kings, grand princes, grand dukes, and sovereign princes. As royalty or nobility, they are ranke ...
, has dark extremities and corn spots.


Terminology

In the most general sense, the word "roan" refers to any animal with a mixture of white and colored hairs in the coat."''roan'', ''a''. and ''n.1''"
Oxford English Dictionary The ''Oxford English Dictionary'' (''OED'') is the first and foundational historical dictionary of the English language, published by Oxford University Press (OUP). It traces the historical development of the English language, providing a com ...
. 2nd edition 1989. OED Online.
Oxford University Press Oxford University Press (OUP) is the university press of the University of Oxford. It is the largest university press in the world, and its printing history dates back to the 1480s. Having been officially granted the legal right to print books ...
. 3 June 2008. .
Recent research into
equine coat color genetics Equine coat color genetics determine a horse's coat color. Many colors are possible, but all variations are produced by changes in only a few genes. The "base" colors of the horse are determined by the Extension locus, which in recessive form (e ...
indicates that "true roan" or "classic roan" is distinct from several similar patterns that are collectively called "roaning". In studies of the white patterning genotypes of laboratory mice, no fewer than four produced roaning or flecking. Therefore, the existence of other types of roaning conditions not covered by those mentioned here is possible and likely. The patterns identified as "roaning" have particular qualities that can be used to tell them apart from true roan.


Roan mimics


Gray vs. roan

Roans are sometimes mistaken for grays. However, grays lighten with age, while roans do not. Gray is one of the most common coat colors found in horses, and is found in almost all breeds. The defining characteristic of the gray coat is that it becomes progressively lighter over time. Gray
foal A foal is an equine up to one year old; this term is used mainly for horses, but can be used for donkeys. More specific terms are colt for a male foal and filly for a female foal, and are used until the horse is three or four. When the foal i ...
s may be born any color, and there may be no indication of the future gray coat at birth. Mature grays may retain none of their original coat color, and have a "
white White is the lightest color and is achromatic (having no hue). It is the color of objects such as snow, chalk, and milk, and is the opposite of black. White objects fully reflect and scatter all the visible wavelengths of light. White on ...
" coat, while the color of the skin and
eyes Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conve ...
is unchanged. The first white hairs are usually seen around the muzzle and eyes. As a gray may go from entirely colored to entirely white over the course of its life, the process of "graying out" can, at times, closely resemble roan.
Thoroughbred The Thoroughbred is a horse breed best known for its use in horse racing. Although the word ''thoroughbred'' is sometimes used to refer to any breed of purebred horse, it technically refers only to the Thoroughbred breed. Thoroughbreds are c ...
and
Arabian Horse The Arabian or Arab horse ( ar, الحصان العربي , DIN 31635, DMG ''ḥiṣān ʿarabī'') is a horse breed, breed of horse that originated on the Arabian Peninsula. With a distinctive head shape and high tail carriage, the Arabian is ...
s registered as "roan" are often gray, especially
chestnuts The chestnuts are the deciduous trees and shrubs in the genus ''Castanea'', in the beech family Fagaceae. They are native to temperate regions of the Northern Hemisphere. The name also refers to the edible nuts they produce. The unrelated ...
turning gray. Unlike grays, roans do not develop more white hair with age, and without white markings, roans retain colored heads.


Dun vs. roan

Blue dun or
grullo Grulla or grullo, also called blue dun, gray dun or mouse dun, is a color of horses in the dun family, characterized by tan-gray or mouse-colored hairs on the body, often with shoulder and dorsal stripes and black barring on the lower legs. In t ...
(also grulla, mouse dun) coloring is created by the
dun gene The dun gene is a dilution gene that affects both red and black pigments in the coat color of a horse. The dun gene lightens most of the body while leaving the mane, tail, legs, and primitive markings the shade of the undiluted base coat color. ...
acting on a
black Black is a color which results from the absence or complete absorption of visible light. It is an achromatic color, without hue, like white and grey. It is often used symbolically or figuratively to represent darkness. Black and white have o ...
base coat, and is a coat color with a bluish cast and darker points.Sponenberg, Dan Phillip (2003). "''Grullo'' is the Spanish name for the crane bird, and these horses are similar to the bluish color of the sandhill crane." p. 37. Unlike blue roans, grullos are solid color and appear bluish due to low amounts of
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...
in each hair, not interspersed white hairs. Like other dun coat colors, grullos have dark or black
primitive markings Primitive markings are a group of hair coat markings and qualities seen in several equine species, including horses, donkeys, and asses. In horses, they are associated with primitive breeds, though not limited to such breeds. The markings are ...
, always including a stripe down the back.


Rabicano vs. roan

One pattern of roaning is
rabicano Rabicano, sometimes called white ticking, is a horse coat color characterized by limited roaning in a specific pattern: its most minimal form is expressed by white hairs at the top of a horse's tail, often is expressed by additional intersperse ...
, also called white ticking. While true roans have an even intermixture of white hairs throughout the body, except the extremities, the white hairs of a rabicano are densest around the base of the tail and the flank. Rabicano roaning frequently forms rings of white hair around the base of the tail, and in extensively roaned rabicanos, the white hairs may converge to form vertical stripes over the ribcage. Rabicano is found in many breeds, and may account for some "roan"
Arabians The Arabs (singular: Arab; singular ar, عَرَبِيٌّ, DIN 31635: , , plural ar, عَرَب, DIN 31635: , Arabic pronunciation: ), also known as the Arab people, are an ethnic group mainly inhabiting the Arab world in Western Asia, No ...
.


Sabino vs. roan

Roaning is also associated with some of the sabino white spotting patterns. Many patterns in many breeds are called "sabino", and these patterns usually feature irregular, rough-edged patches of white that originate from the lower legs, face, and ventral midline. The borders of these white patches can be heavily roaned, and some sabinos can be mistaken for roans. The roaning of sabinos originates in a white patch, and the roaning is uneven.


Varnish roan vs. roan

The
leopard complex The leopard complex is a group of genetically related coat patterns in horses. These patterns range from progressive increases in interspersed white hair similar to graying or roan to distinctive, Dalmatian-like leopard spots on a white coat. ...
colors, characteristic of the
Appaloosa The Appaloosa is an American horse breed best known for its colorful spotted coat pattern. There is a wide range of body types within the breed, stemming from the influence of multiple breeds of horses throughout its history. Each horse's colo ...
breed, have several manifestations that feature mixtures of white and colored hairs. A
varnish roan Varnish roan describes a horse with coloration similar to roan, but with some changes in color over the years, though not to the extreme of a gray. This type of roaning only occurs in conjunction with the Leopard complex. Varnish roans are born w ...
, one type of leopard complex coat color also called "marble", is an all-over blend of white and colored hairs. Patches of skin that lie close to the bone, such as on the face and legs, and the point of shoulder and point of hip, do not grow as much white hair. These darker patches are called "varnish marks" and are not found in true roans. Varnish roans can also be distinguished from true roans by the presence of leopard complex characteristics, such as the white
sclera The sclera, also known as the white of the eye or, in older literature, as the tunica albuginea oculi, is the opaque, fibrous, protective, outer layer of the human eye containing mainly collagen and some crucial elastic fiber. In humans, and som ...
, finely striped
hooves The hoof (plural: hooves) is the tip of a toe of an ungulate mammal, which is covered and strengthened with a thick and horny keratin covering. Artiodactyls are even-toed ungulates, species whose feet have an even number of digits, yet the rumi ...
, and
mottled Mottle is a pattern of irregular marks, spots, streaks, blotches or patches of different shades or colours. It is commonly used to describe the surface of plants or the skin of animals. In plants, mottling usually consists of yellowish spots o ...
skin around the eyes, muzzle, and genitals.


Prevalence

True or classic roan is common in European
draft Draft, The Draft, or Draught may refer to: Watercraft dimensions * Draft (hull), the distance from waterline to keel of a vessel * Draft (sail), degree of curvature in a sail * Air draft, distance from waterline to the highest point on a vesse ...
breeds such as the
Brabant Brabant is a traditional geographical region (or regions) in the Low Countries of Europe. It may refer to: Place names in Europe * London-Brabant Massif, a geological structure stretching from England to northern Germany Belgium * Province of Bra ...
,
Ardennes The Ardennes (french: Ardenne ; nl, Ardennen ; german: Ardennen; wa, Årdene ; lb, Ardennen ), also known as the Ardennes Forest or Forest of Ardennes, is a region of extensive forests, rough terrain, rolling hills and ridges primarily in Be ...
,
Trait Du Nord The Trait du Nord, previously also known as Ardennais du Nord or Ardennais de type Nord, is a breed of heavy draft horse developed and bred in the area of Hainaut in western Belgium and in northeastern France. Originally considered a subtype o ...
, Italian Heavy Draft, and Rhenish-German Cold-Blood. It is also found regularly in North American breeds such as the
Quarter Horse The American Quarter Horse, or Quarter Horse, is an American breed of horse that excels at sprinting short distances. Its name is derived from its ability to outrun other horse breeds in races of a quarter mile or less; some have been clocked at ...
,
Paint Horse The American Paint Horse is a breed of horse that combines both the conformational characteristics of a western stock horse with a pinto spotting pattern of white and dark coat colors. Developed from a base of spotted horses with Quarter Hors ...
,
Peruvian Paso The Peruvian Horse is a breed of light saddle horse known for its smooth ride. It is distinguished by a natural, four-beat, lateral gait called the ''paso llano.'' This breed is protected by the Peruvian government through Decree number 25919 o ...
,
Paso Fino The Paso Fino is a naturally gaited light horse breed dating back to horses imported to the Caribbean from Spain. Pasos are prized for their smooth, natural, four-beat, lateral ambling gait; they are used in many disciplines, but are especially ...
,
Standardbred The Standardbred is an American horse breed best known for its ability in harness racing, where members of the breed compete at either a trot or pace. Developed in North America, the Standardbred is recognized worldwide, and the breed can trace i ...
,
Spanish Mustang The Spanish Mustang is an American horse breed descended from horses brought from Spain during the early conquest of the Americas. They are classified within the larger grouping of the Colonial Spanish horse, a type that today is rare in Spain. ...
,
Missouri Fox Trotter The Missouri Fox Trotter is a horse breed that originated in the state of Missouri in the United States. It was developed in the Ozark Mountains by settlers in the early 19th century, and quickly developed into a gaited breed appreciated for it ...
and
Tennessee Walking Horse The Tennessee Walking Horse or Tennessee Walker is a breed of gaited horse known for its unique four-beat running-walk and flashy movement. It was originally developed as a riding horse on farms and plantations in the American South. It is ...
. British ponies such as Dales,
Welsh Welsh may refer to: Related to Wales * Welsh, referring or related to Wales * Welsh language, a Brittonic Celtic language spoken in Wales * Welsh people People * Welsh (surname) * Sometimes used as a synonym for the ancient Britons (Celtic peop ...
, Gypsy Cob,
Shetland Shetland, also called the Shetland Islands and formerly Zetland, is a subarctic archipelago in Scotland lying between Orkney, the Faroe Islands and Norway. It is the northernmost region of the United Kingdom. The islands lie about to the no ...
,
Connemara Connemara (; )( ga, Conamara ) is a region on the Atlantic Ocean, Atlantic coast of western County Galway, in the west of Ireland. The area has a strong association with traditional Irish culture and contains much of the Connacht Irish-speak ...
, and
New Forest The New Forest is one of the largest remaining tracts of unenclosed pasture land, heathland and forest in Southern England, covering southwest Hampshire and southeast Wiltshire. It was proclaimed a royal forest by William the Conqueror, featu ...
ponies may be roan. Icelandic Horses and
Miniature Horse A miniature horse is a breed or type of horse characterised by its small size. Usually it has been bred to display in miniature the physical characteristics of a full-sized horse, but to be little over in height, or even less. Although such hor ...
s may also be roan. Some roan
Trakehner Trakehner is a light warmblood breed of horse, originally developed at the East Prussian state stud farm in the town of Trakehnen from which the breed takes its name. The state stud ( :de:Hauptgestüt Trakehnen) was established in 1731 and ...
s and American Saddlebreds are known. The Hokkaido Pony of
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
may also be roan.Overton, Rebecca (2004). "Other studies discovered them in two other equine breeds, the Ardennais, found in France, and the Hokkaido, which live in Japan." To date, only one Thoroughbred family has been genetically verified as true roan, and there are no
Arabians The Arabs (singular: Arab; singular ar, عَرَبِيٌّ, DIN 31635: , , plural ar, عَرَب, DIN 31635: , Arabic pronunciation: ), also known as the Arab people, are an ethnic group mainly inhabiting the Arab world in Western Asia, No ...
that are true roan. A small number of
Thoroughbred The Thoroughbred is a horse breed best known for its use in horse racing. Although the word ''thoroughbred'' is sometimes used to refer to any breed of purebred horse, it technically refers only to the Thoroughbred breed. Thoroughbreds are c ...
offspring of the
brindle Brindle is a coat (animal), coat coloring pattern in animals, particularly dogs, cattle, guinea pigs, Cat, cats, and, rarely, horses. It is sometimes described as "tiger-striped", although the brindle pattern is more subtle than that of a tiger' ...
-patterned stallion Catch A Bird phenotypically appear to be true roan, and though few of those reproduced, some offspring of his daughter Slip Catch, though registered as bay, have apparently been verified as carrying the roan gene. Nonetheless, most Thoroughbreds and Arabians with roan-like coats are most likely related to graying or
rabicano Rabicano, sometimes called white ticking, is a horse coat color characterized by limited roaning in a specific pattern: its most minimal form is expressed by white hairs at the top of a horse's tail, often is expressed by additional intersperse ...
characteristics. Thoroughbred "roans" are described by the
Jockey Club The Jockey Club is the largest commercial horse racing organisation in the United Kingdom. It owns 15 of Britain's famous racecourses, including Aintree, Cheltenham, Epsom Downs and both the Rowley Mile and July Course in Newmarket, amo ...
as a mixture of white with red or brown hairs, but which researchers identify as chestnuts turning gray. The Arabian Horse Association defines "roan" as an even mixture of white and colored, usually chestnut, hairs, but researchers suggest most animals so classified are probably either rabicano, or have a partially-spotted pattern that results from a
dominant white Dominant white (W) is a group of genetically related coat color alleles on the KIT gene of the horse, best known for producing an all-white coat, but also able to produce various forms of white spotting, as well as bold white markings. Prior ...
allele.


Genetics

Roan is a simple dominant trait symbolized by the ''Rn''
allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chro ...
. Traits that are dominantly inherited cannot skip generations, meaning that two nonroan parents cannot produce a roan offspring. In cases where roan has appeared to skip generations, one of the parents is usually discovered to be slightly roaned. A roan can also be born from two seemingly nonroan parents if the roan coat is "masked" by extensive white markings or gray. In some cases, the supposedly roan offspring is not true roan at all, but rabicano, sabino, or influenced by some other genetic factor. The
University of California, Davis The University of California, Davis (UC Davis, UCD, or Davis) is a public land-grant research university near Davis, California. Named a Public Ivy, it is the northernmost of the ten campuses of the University of California system. The institut ...
School of Veterinary Medicine's genetics services has developed a
DNA test 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, o ...
that uses
genetic marker A genetic marker is a gene or DNA sequence with a known location on a chromosome that can be used to identify individuals or species. It can be described as a variation (which may arise due to mutation or alteration in the genomic loci) that can be ...
s to indirectly determine the number of ''Rn'' or ''rn'' alleles a horse has. The
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...
responsible for true roan has not yet been identified exactly, but been assigned to equine
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
3 (ECA3) in the KIT sequence. A roan zygosity test is reliable for American Quarter Horse, American Paint Horse, Welsh Pony, and Gypsy Cob. Until a direct test is developed, the roan zygosity test may enable breeders to produce roans more reliably.


Homozygous roan

A 1979 study of American-bred Belgian draft horses found fewer roan offspring from roan-to-roan matings than expected for a simple dominant trait. Finding neither stillborn nor sickly, short-lived foals from these roan parents, the researchers concluded that in the
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. Mo ...
condition the roan gene was lethal to the
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
or
fetus A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal deve ...
. Other studies at the time were using progeny ratios to identify potential lethality related to other coat colors, such as "
dominant white Dominant white (W) is a group of genetically related coat color alleles on the KIT gene of the horse, best known for producing an all-white coat, but also able to produce various forms of white spotting, as well as bold white markings. Prior ...
" and "
lethal white ''Lethal White'' is the fourth novel in the ''Cormoran Strike'' series, written by J. K. Rowling and published under the pseudonym Robert Galbraith. The novel was released on 18 September 2018. Background ''Lethal White'' is preceded by ...
", so roan was believed to follow a similar pattern. Genetic science in the 1970s could not provide a clear answer, as methods of molecular analysis had not yet been developed."The paper by Hintz and Van Vleck caused all this commotion, but there were earlier mentions of a roan cross being lethal, too," Penedo explained. "But it was never fully established by any molecular analysis that homozygous roan horses would not be produced from matings of two roan parents." in Overton, 2004. The belief in "lethal roan" persisted until recently, when homozygous roan stallions with large numbers of offspring - all roan - were satisfactorily identified. Homozygous roan stallions were identified in both European populations of Belgian horses in 1977, and in North American, in Quarter Horse stallions. In some breeds, homozygous roans can be identified by an indirect
DNA marker A genetic marker is a gene or DNA sequence with a known location on a chromosome that can be used to identify individuals or species. It can be described as a variation (which may arise due to mutation or alteration in the genomic loci) that can be ...
test. Such horses, with the
genotype The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
''Rn/Rn'', produce 100% roan offspring. Homozygous roans and heterozygous roans (''Rn/rn'') are identical in appearance.


Equine linkage group II

During the production of
sex cells Sex is the trait that determines whether a sexually reproducing animal or plant produces male or female gametes. Male plants and animals produce smaller mobile gametes (spermatozoa, sperm, pollen), while females produce larger ones ( ova, of ...
, DNA is "reshuffled" to ensure that the next generation does not inherit ancestral
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s as is, but a mixture of DNA from each parent. Paired chromosomes exchange parallel pieces of DNA, a process called
chromosomal crossover Chromosomal crossover, or crossing over, is the exchange of genetic material during sexual reproduction between two homologous chromosomes' non-sister chromatids that results in recombinant chromosomes. It is one of the final phases of geneti ...
, prior to being passed on to the next generation. When particular
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s are located physically close together on a chromosome, they tend to be exchanged together. The rate at which two genes are passed on together can be used to calculate their
distance Distance is a numerical or occasionally qualitative measurement of how far apart objects or points are. In physics or everyday usage, distance may refer to a physical length or an estimation based on other criteria (e.g. "two counties over"). ...
from each other on the chromosome. Genes that tend to be passed on as a group form what is called a "
linkage group Genetic linkage is the tendency of DNA sequences that are close together on a chromosome to be inherited together during the meiosis phase of sexual reproduction. Two genetic markers that are physically near to each other are unlikely to be separ ...
". Equine linkage group II includes a number of genes. The first linkage was found between a
blood type A blood type (also known as a blood group) is a classification of blood, based on the presence and absence of antibodies and inherited antigenic substances on the surface of red blood cells (RBCs). These antigens may be proteins, carbohydrate ...
marker and
tobiano Tobiano is a spotted color pattern commonly seen in pinto horses, produced by a dominant gene. The tobiano gene produces white-haired, pink-skinned patches on a base coat color. The coloration is almost always present from birth and does not c ...
white spotting in 1978. The same blood type marker,
albumin Albumin is a family of globular proteins, the most common of which are the serum albumins. All the proteins of the albumin family are water-soluble, moderately soluble in concentrated salt solutions, and experience heat denaturation. Albumins ...
, was linked to another blood protein called
vitamin D-binding protein Vitamin D-binding protein (DBP), also/originally known as gc-globulin (group-specific component), is a protein that in humans is encoded by the ''GC'' gene. DBP is genetically the oldest member of the albuminoid family and appeared early in the e ...
. In 1982, a linkage grouping was proposed, including three genes for serum
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s, and three for coat color: tobiano spotting,
chestnut The chestnuts are the deciduous trees and shrubs in the genus ''Castanea'', in the beech family Fagaceae. They are native to temperate regions of the Northern Hemisphere. The name also refers to the edible nuts they produce. The unrelat ...
, and roan. Research since equine linkage group II was defined has identified the exact location of the tobiano allele and the chestnut allele. The region of the chromosome that harbors the roan gene is homologous to parts of chromosomes in other species that also control coat color, even some similar to roan. Presently, the '' KIT'' gene is the primary candidate gene for roan. While a chromosomal inversion of ''KIT'' causes
tobiano Tobiano is a spotted color pattern commonly seen in pinto horses, produced by a dominant gene. The tobiano gene produces white-haired, pink-skinned patches on a base coat color. The coloration is almost always present from birth and does not c ...
white spotting, ''KIT'' also harbors one or more
allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chro ...
s responsible for sabino spotting, no fewer than eleven alleles responsible for
dominant white Dominant white (W) is a group of genetically related coat color alleles on the KIT gene of the horse, best known for producing an all-white coat, but also able to produce various forms of white spotting, as well as bold white markings. Prior ...
spotting, and is thought to be a major contributor to many other forms of less distinctive white markings. The effect of linkage between roan and chestnut is readily observed. If a horse possesses one chromosome with the
wildtype The wild type (WT) is the phenotype of the typical form of a species as it occurs in nature. Originally, the wild type was conceptualized as a product of the standard "normal" allele at a locus, in contrast to that produced by a non-standard, "m ...
nonchestnut allele and the dominant roan allele (''E'' and ''Rn''), while the other chromosome contains the recessive chestnut allele and the recessive nonroan allele (''e'' and ''rn''), it will outwardly appear blue roan, barring the influence of other genes. Normally, the chestnut and roan alleles would be separated during chromosomal crossover, but these two linked genes usually remain together. Such a horse will produce sex cells that are either ''E/Rn'' or ''e/rn''. Mated to chestnut nonroan partners (''e/rn''), the horse would produce primarily blue roans, or chestnut nonroans, but few chestnut roans and few black nonroans. If the recessive ''e'' and dominant ''Rn'' were on the same chromosome, though, the horse would be expected to produce primarily chestnut roans and nonchestnut nonroans with chestnut, nonroan partners.


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{{Equine coat colors Horse coat colors