Gray (horse)
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A gray horse (or grey horse) has a coat color characterized by progressive depigmentation of the colored hairs of the coat. Most gray horses have black skin and dark eyes; unlike some equine dilution genes and some other genes that lead to depigmentation, gray does not affect skin or eye color. Gray horses may be born any base color, depending on other color genes present. White hairs begin to appear at or shortly after birth and become progressively more prevalent as the horse ages as white hairs become intermingled with hairs of other colors. Graying can occur at different rates—very quickly on one horse and very slowly on another. As adults, most gray horses eventually become completely white, though some retain intermixed light and dark hairs. The stages of graying vary widely. Some horses develop a dappled pattern for a period of time, others resemble a roan with more uniform intermixing of light and dark hairs. As they age, some gray horses, particularly those
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. Mo ...
for the gray gene, may develop pigmented speckles in addition to a white coat, a pattern colloquially called a “fleabitten gray.” Gray horses appear in many breeds, though the color is most commonly seen in
breeds A breed is a specific group of domestic animals having homogeneous appearance (phenotype), homogeneous behavior, and/or other characteristics that distinguish it from other organisms of the same species. In literature, there exist several slight ...
descended from Arabian ancestors. Some breeds that have large numbers of gray-colored horses include the
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 ...
, the Arabian, the American Quarter Horse and the
Welsh pony The Welsh Pony and Cob is a group of four closelyrelated horse breeds including both pony and cob types, which originated in Wales in the United Kingdom. The four sections within the breed society for the Welsh breeds are primarily distingui ...
. Breeds with a very high prevalence of gray include the
Percheron The Percheron is a breed of draft horse that originated in the Huisne river valley in western France, part of the former Perche province from which the breed takes its name. Usually gray or black in color, Percherons are well muscled, and ...
, the Andalusian, and the
Lipizzaner The Lipizzan or Lipizzaner ( hr, Lipicanac, cz, Lipicán, hu, Lipicai, it, Lipizzano, sr, Lipicaner, sl, Lipicanec), is a European breed of riding horse developed in the Habsburg Empire in the sixteenth century. It is of Baroque type, and ...
. People who are unfamiliar with horses may refer to gray horses as "white". However, a gray horse whose hair coat is completely "white" will still have black skin (except under markings that were white at birth) and dark eyes. This is how to discern a gray horse from a white horse. White horses usually have pink skin and sometimes even have blue eyes. Young horses with hair coats consisting of a mixture of colored and gray or white hairs are sometimes confused with roan. Some horses that carry dilution genes may also be confused with white or gray. While gray is classified as a coat color by breed registries, genetically it may be more correct to call it a
depigmentation Depigmentation is the lightening of the skin or loss of pigment. Depigmentation of the skin can be caused by a number of local and systemic conditions. The pigment loss can be partial (injury to the skin) or complete (caused by vitiligo). It can be ...
pattern. It is a
dominant allele In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and t ...
, and thus a horse needs only one copy of the gray
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 ...
, that is,
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. Mo ...
, to be gray in color. A
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 ...
gray horse, one carrying two gray alleles, will always produce gray foals.


Prevalence

Gray is common in many breeds. Today, about one horse in 10 carries the mutation for graying with age. The vast majority of
Lipizzaner The Lipizzan or Lipizzaner ( hr, Lipicanac, cz, Lipicán, hu, Lipicai, it, Lipizzano, sr, Lipicaner, sl, Lipicanec), is a European breed of riding horse developed in the Habsburg Empire in the sixteenth century. It is of Baroque type, and ...
s are gray, as are the majority of Andalusian horses. Many breeds of French draft horse such as the
Percheron The Percheron is a breed of draft horse that originated in the Huisne river valley in western France, part of the former Perche province from which the breed takes its name. Usually gray or black in color, Percherons are well muscled, and ...
and Boulonnais are often gray as well. Gray is also found among Welsh Ponies,
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 ...
s, and American Quarter Horses. All of these breeds have common ancestry in the Arabian horse. In particular, all gray Thoroughbreds descend from a horse named
Alcock's Arabian Alcock's Arabian (foaled about 1700, died about 1733), also known as Pelham Grey Arabian and less certainly as Bloody Buttocks and Ancaster Turk, among other names, is the ancestor of all grey-coloured Thoroughbred horses,Lady Wentworth, ''The S ...
, a gray born in 1700. The gray coat color makes up about 3% of Thoroughbreds. Gray also affects spotting patterns of
pinto Pinto is a Portuguese language, Portuguese, Spanish language, Spanish, Sephardi Jews, Jewish (Sephardic), and Italian language, Italian surname. It is a high-frequency surname in all List of countries and territories where Portuguese is an officia ...
s and
leopard complex The leopard complex is a group of genetically related Equine coat color, coat patterns in horses. These patterns range from progressive increases in interspersed white hair similar to Gray (horse), graying or Roan (horse), roan to distinctive, Da ...
horses such as
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 ...
s. Its effects wash out the contrast of the markings of these patterns, sometimes colloquially described as “ghosting.”


Changes in the color of gray horses

A gray foal may be born any color. However, bay, chestnut, or
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 ...
base colors are most often seen. As the horse matures, it “grays out” as white hairs begin to replace the base or birth color. Usually white hairs are first seen by the muzzle, eyes and flanks, occasionally at birth, and usually by the age of one year. Over time, white hairs replace the birth color. The changing patterns of white and dark hairs have many informal names, such as “rose gray,” “salt and pepper,” “iron gray”, or “dapple gray.” As the horse ages, the coat continues to lighten, often to a pure white. Some horses develop pigmented reddish-brown speckles on an otherwise white hair coat. Such horses are often called “fleabitten gray”. Different breeds, and individuals within each breed, take differing amounts of time to gray out. Thus, graying cannot be used to approximate the age of a horse except in the broadest of terms: a very young horse will never have a white coat (unless it is a true white horse), while a horse in its teens usually is completely grayed out. One must also be careful not to confuse the small amount of gray hairs that may appear on some older horses in their late teens or twenties, which do not reflect the gray gene and never cause a complete graying of the horse. This change in hair color means that the same gray horse will appear to be a different color over time, sometimes resulting in a need to change the color noted on
breed registry A breed registry, also known as a herdbook, studbook or register, in animal husbandry and the hobby of animal fancy, is an official list of animals within a specific breed whose parents are known. Animals are usually registered by their breeder ...
papers. Other times, people traveling with gray horses who have a pure white hair coat have encountered problems with non-horse-oriented officials such as police officers or border guards who are unclear about a horse who has papers saying it is "gray" when the horse in front of them appears white. To further complicate matters, the skin and eyes may be other colors if influenced by other factors such as white markings, certain white spotting patterns or dilution genes.


The genetics of gray

The gray gene (G) is an autosomal
dominant gene In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and t ...
. In simple terms, a horse which has even one copy of the gray
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 ...
, regardless of other coat color genes present, will always become gray. This also means that all gray horses must have at least one gray parent. If a gray horse is
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 ...
(GG), meaning that it has a gray
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 ...
from both parents, it will always produce gray offspring no matter the color genetics of the other parent. However, if a gray horse is
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. Mo ...
(Gg), meaning it inherits one copy of the recessive gene (g), that animal may produce offspring who are not gray, depending on the genetics of the other parent and Mendelian inheritance principles. Genetic testing is now possible to determine whether a horse is homozygous or heterozygous for gray. The gray gene does not affect skin or eye color, so grays typically have dark skin and eyes, as opposed to the unpigmented pink skin of
white horses ''The White Horses'' is a 1965 television series co-produced by RTV Ljubljana (now RTV Slovenija) of Yugoslavia and German TV (''Südwestfunk''). Plotline The story follows the adventures of a teenage girl Julia (Helga Anders) who leaves Belgrad ...
. In 2008, researchers at Uppsala University in Sweden identified the genetic mutation that governs the graying process. The study revealed that all gray horses carry an identical mutation that can be traced back to a common ancestor that lived at least two thousand years ago. The discovery that gray can be linked to a single animal provides an example of how humans have "cherry-picked" attractive mutations in domestic animals. Gray is controlled by a single dominant
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 ...
of a
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 b ...
that regulates specific kinds of stem cells.
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 ...
grays turn white faster, are more likely to develop melanomas, and are less prone to develop the "fleabitten" speckling than
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. Mo ...
grays. Researchers suggest the pigmented speckles of the “fleabitten” gray, as well as more intense reddish-brown colored areas called “blood” markings, may be caused by a loss or inactivation of the gray allele in some of the somatic cells as that would explain why the speckles are more common on heterozygous grays than homozygotes.


Melanoma in gray horses

The identification of the gray mutation is of great interest in of medical research since this mutation also enhances the risk for melanoma in horses: some studies have suggested as many as 80% of grays over 15 years of age have some form of melanoma. Growth rate depends on the type, and many are slow-growing, but over time, many develop into a
malignant melanoma Melanoma, also redundantly known as malignant melanoma, is a type of skin cancer that develops from the pigment-producing cells known as melanocytes. Melanomas typically occur in the skin, but may rarely occur in the mouth, intestines, or eye ...
. The study of gray genetics has pointed to a molecular pathway that may lead to tumour development. Both STX17 and the neighboring NR4A3 gene are overexpressed in melanomas from gray horses, and those carrying a loss-of-function mutation in ASIP (agouti signaling protein) had a higher incidence of melanoma, implying that increased melanocortin-1 receptor signaling promotes melanoma development in Gray horses. Some studies indicate as many as 66% of melanomas become malignant, though other studies have found much lower rates, and in one case, zero.


Horse coat colors sometimes confused with gray


White horses

Many people who are unfamiliar with horses refer to a gray horse as "white". However, most
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 o ...
horses have pink skin and some have blue eyes. A horse with dark skin and dark eyes under a white hair coat is gray. However, a gray horse with an underlying homozygous cream base coat color may be born with rosy-pink skin, blue eyes and near-white hair. In such cases, DNA testing may clarify the genetics of the horse.


Roan

Some grays in intermediate stages of graying may be confused with a roan or a
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 ...
. Some heavily fleabitten grays may also be confused with roans. However, roans are easily distinguishable from grays: roan consists of individual white hairs on a dark base coat, usually with the head and legs of the horse darker than the rest of the body. Rabicanos also have intermixed white hairs primarily on the body with a dark head. With gray horses, the head is often the first area to lighten, especially around the eyes and muzzle. Also, roans do not lighten with age, while grays always do. The
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 ...
is another unusual coloration, sometimes seen in
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 ...
horses, that, like gray, can change with age, but unlike gray, the horse does not become progressively lighter until it is pure white. Varnish roans are created by the action of
leopard complex The leopard complex is a group of genetically related Equine coat color, coat patterns in horses. These patterns range from progressive increases in interspersed white hair similar to Gray (horse), graying or Roan (horse), roan to distinctive, Da ...
within breeds such as the Appaloosa and are seldom seen elsewhere.


Diluted colors

The dilution genes that create dun, cream, pearl, silver dapple and champagne coloring may occasionally result in confusion with gray. Some horses with a particular type of
dun A dun is an ancient or medieval fort. In Ireland and Britain it is mainly a kind of hillfort and also a kind of Atlantic roundhouse. Etymology The term comes from Irish ''dún'' or Scottish Gaelic ''dùn'' (meaning "fort"), and is cognat ...
hair coat known as a "blue dun",
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 ...
, or "mouse" dun appear to be a solid gray. However, this color is caused 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 base coat, and horses who are dun have all hairs the same color; there is no intermingling of white and dark hairs. Also, dun horses do not get lighter as they age. Horses that are a light cream color are also not grays. These are usually cremello, perlino or smoky cream horses, all colors produced by the action of the
cream gene The cream gene is responsible for a number of horse coat colors. Horses that have the cream gene in addition to a base coat color that is chestnut will become palomino if they are heterozygous, having one copy of the cream gene, or cremello, if ...
. However, if a gray parent passes on the gene, the hairs will turn white like any other gray. Another cream-colored dilution, the
pearl gene The Pearl gene, also known as the "Barlink factor", is a dilution gene at the same locus as the cream gene, which somewhat resembles the cream gene and the champagne gene but is unrelated to champagne. It is a somewhat rare dilution gene found in ...
or "barlink factor", may also create very light-coated horses. Similarly, the
champagne gene The champagne gene is a simple dominant allele responsible for a number of rare horse coat colors. The most distinctive traits of horses with the champagne gene are the hazel eyes and pinkish, freckled skin, which are bright blue and bright pink ...
can lighten coat color, often producing dappling or light colors that can be confused with gray. In spite of its name, the
silver dapple gene The silver or silver dapple (''Z'') gene is a dilution gene that affects the black base coat color and is associated with Multiple Congenital Ocular Abnormalities. It will typically dilute a black mane and tail to a silvery gray or flaxen color ...
has nothing to do with graying. It is a dilution gene that acts only 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 ...
coat, diluting the coat to a dark brown and the mane to a flaxen shade. Horses that express the silver dapple gene (and do not have the gray gene) are born that color and it will not lighten. However, again, if one parent passes on the gray gene, the gray gene will again be dominant.


Mythology

Throughout history, both gray and
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 o ...
horses have been mythologized. As part of its legendary dimension, the gray horse in myth has been depicted with seven heads ( Uchaishravas) or eight feet ( Sleipnir), sometimes in groups or singly. There are also mythological tales of divinatory gray horses who prophesy or warn of danger.


See also

*
Albinism Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and pink or blue eyes. Individuals with the condition are referred to as albino. Varied use and interpretation of the term ...
*
Equine 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 ...
*
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 ...


Notes


References


"Introduction to Coat Color Genetics"
''from'' Veterinary Genetics Laboratory, School of Veterinary Medicine,
University of California The University of California (UC) is a public land-grant research university system in the U.S. state of California. The system is composed of the campuses at Berkeley, Davis, Irvine, Los Angeles, Merced, Riverside, San Diego, San Franci ...
, Davis. Web Site accessed January 12, 2008 {{Equine coat colors Horse coat colors