Overo refers to several genetically unrelated pinto coloration patterns of white-over-dark body markings in horses, and is a term used by the American Paint Horse Association to classify a set of pinto patterns that are not tobiano. Overo is a Spanish word, originally meaning "like an egg". [1] The most common usage refers to frame overo , but splashed white and sabino are also considered "overo". [2] [3] A horse with both tobiano and overo patterns is called tovero.
Frame overo, splashed white, and sabino are three separate pinto patterns, genetically unrelated, that are grouped together under the name "overo".
The frame overo pattern tends to have white spots on the flanks and face, surrounded by a dark "frame" of color. If two horses with the frame overo gene are bred together, there is a 25% chance the foal will have lethal white syndrome. [4]
Splashed white or splash overo is a group of patterns that tend to have white on the underside, as if a horse ran through white paint with its head lowered. So far 6 alleles on two genes have been discovered to cause splashed white patterns. There is a correlation between the splashed white pattern and deafness. [5]
Sabino or sabino overo is a group of patterns that typically have white on the legs, face, and sometimes also belly, often with white ticking or roaning. Several alleles on the KIT gene are known to cause sabino or sabino-like patterns. [6] [7]
The frame overo pattern is the most common of the three types of overo patterns recognized in the American Paint Horse breed. [1] A frame overo horse appears to be any solid base color (bay, black, chestnut, etc.) with white irregular patches added, usually with a horizontal orientation. Markings are often of jagged shape rather than rounded, the white rarely crosses the back, the lower legs tend to be dark, and the tail is one color, usually dark. The head is often white or bald-faced, and blue eyes are not uncommon. The frame overo pattern usually behaves like a dominant gene, in that when frame overo horses are crossed on nonspotted horses, about half of the foals come out spotted. [2] There are records of frame overos being produced by two nonspotted parents. There is a theory, however, that these "solid" horses simply may be horses with very minimal expression of overo genetics. [2]
Frame coloring is controversial because it is associated with lethal white syndrome (OLWS or LWS), [8] the equine version of Hirschprung disease. [9] LWS occurs when a foal is homozygous for the Ile118Lys EDNRB mutation, which is considered by many researchers to be "usually responsible" for the frame overo phenotype. [10] However, other researchers emphasize that overo spotting patterns are phenotypically and genetically heterogeneous, that is, may have multiple sources. [11]
The frame overo gene can be masked by other white patterns, particularly tobiano, which is a dominant gene and epistatic to overo. [9] Epistatic means that when both genes are present, this is the one expressed.
In addition, some carriers of the LWS allele appear to be solid. One theory holds that such horses carry the frame gene, but so minimal in expression that they appear solid. [2] Either way, all LWS foals have horses with frame overo patterning in their pedigrees, and horses carrying a frame allele may not necessarily have a visible expression of the frame overo color. [8]
Foals which are homozygous for frame and thus have lethal white syndrome (LWS) are not albinos. Albinism does not exist in horses. [12] LWS foals are born almost or completely white with pink skin, but have blue eyes, not red ones. The lethal trait is that the nerves of the foal's digestive system are undeveloped and the bowel cannot move food along. All lethal white foals die within 72 hours after birth, and are typically euthanized sooner for humane reasons. A horse can be a carrier of the trait when it carries only one copy of the gene, (that is, heterozygous) and carriers are healthy and show no clinical signs of lethal white syndrome. [1]
Thus, it is recommended by organizations such as the American Paint Horse Association that horses of any pattern with overo ancestors be tested to verify their carrier state before being bred. There is a DNA test for the gene which can be used to plan breedings and avoid producing affected foals. [2] The mutation has not been found in solid-colored horses from breeds without frame patterning. [9] Likewise, a sick foal that happens to be white can be tested to avoid inadvertent euthanization of a non-lethal foal who has a simple case of colic.
Lethal white horses should not be confused with non-lethal forms of white, such as dominant white, which may be completely white with pink skin. Blue-eyed, pink-skinned cremellos and other horses carrying the cream gene do not carry the LWS allele unless they also have an ancestor with overo genetics. The splash white and sabino genetics are not associated with lethal white, though such horses could also be carriers of the frame allele.
A splash or splashed white overo pattern appears like a solid-colored horse who has been dipped in white paint, and the color splashed up from the bottom. It is the least common of the overo patterns. [1] The legs and bottom portion of the body are usually white, as is the head, and blue eyes are common. Edges where white and dark color meet are usually crisp and sharp. Recent studies suggest that splashed white may be caused by a dominant gene. [1] Splash overos are more prone to being deaf than other horses. [13]
Outside of the United States, the term "splash" is sometimes applied to horses of apparent Sabino patterning.
Sabino is listed a type of overo coloring by the American Paint Horse Association. [14] This terminology, outdated in terms of modern genetics, creates confusion, because other breeds, including many that will not register "pinto" or "paint" horses, may have individuals with sabino genetics. One reason for the terminology confusion is that "overo" was used by Paint horse breeders in the United States to refer to "anything that is not tobiano." [1] Another reason for this terminology is that the term "overo", when used outside of the US, particularly in Spanish-speaking countries, refers to horses with the speckled roaning patterns typical of horses called sabino in the USA. In addition, strongly marked sabino horses may mimic either overo or tobiano coloring. [1]
In general, sabino is a generic description for a group of similar white spotting patterns. [11] These include high white stockings on all four feet, often extending up the legs with jagged edges, a fairly white head, with markings often extending past the eyes, roaning at the edges of white markings, lip spots, "lacing," and white patches on the lower barrel that may extend to the flanks. [1] A minimal sabino may only have one of the traits associated with sabino horses. [1] On the other hand, a "sabino-white" is a near-white or completely white horse, and sabino genetics are not linked to lethal white. [11]
Tobiano coloring is the inverse of overo spotting. Tobianos have a vertical spotting pattern, large, rounded spots, white that crosses the topline, dark heads, but mostly white legs and white or multi-colored tail. Typical examples tend to have more white than dark, whereas typical frame overo horses tend to more dark than white, though there are many exceptions in both cases. It is created by a different gene from any of the overo patterns and is a dominant gene. All tobianos must have at least one tobiano parent. When a tobiano allele is present, it is epistatic and often masks other patterns. [9]
A tovero horse has pinto spotting patterns that show characteristics of both overo and tobiano and probably carries genes for both. For example, a tovero might have tobiano body spotting with rounded edges and white across the back, yet have irregular facial markings and blue eyes.
Horses may also have a combination of sabino and frame overo genetics or any other combination of genes, resulting patterns that are difficult to smoothly classify into any one group.
Roan is a coat color found in many animals, including horses, cattle, antelope, cat and dogs. It is defined generally as an even mixture of white and pigmented hairs that do not "gray out" or fade as the animal ages. There are a variety of genetic conditions which produce the colors described as "roan" in various species.
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 Horse and Thoroughbred bloodlines, the American Paint Horse Association (APHA) breed registry is now one of the largest in North America. The registry allows some non-spotted animals to be registered as "Solid Paint Bred" and considers the American Paint Horse to be a horse breed with distinct characteristics, not merely a color breed.
Lethal white syndrome (LWS), also called overo lethal white syndrome (OLWS), lethal white overo (LWO), and overo lethal white foal syndrome (OLWFS), is an autosomal genetic disorder most prevalent in the American Paint Horse. Affected foals are born after the full 11-month gestation and externally appear normal, though they have all-white or nearly all-white coats and blue eyes. However, internally, these foals have a nonfunctioning colon. Within a few hours, signs of colic appear; affected foals die within a few days. Because the death is often painful, such foals are often humanely euthanized once identified. The disease is particularly devastating because foals are born seemingly healthy after being carried to full term.
A piebald or pied animal is one that has a pattern of unpigmented spots (white) on a pigmented background of hair, feathers or scales. Thus a piebald black and white dog is a black dog with white spots. The animal's skin under the white background is not pigmented.
Skewbald is a colour pattern of horses. A skewbald horse has a coat made up of white patches on a non-black base coat, such as chestnut, bay, or any colour besides black coat. Skewbald horses which are bay and white are sometimes called tricoloured. These horses usually have pink skin under white markings and dark skin under non-white areas. Other than colour, it is similar in appearance to the piebald pattern. Some animals also exhibit colouration of the irises of the eye that match the surrounding skin. The underlying genetic cause is related to a condition known as leucism. The term is also used to describe spotting patterns in various other animals, such as goats.
Tri-coloured refers to a horse with three different coat colours in a pinto spotting pattern of large white and dark patches, usually bay and white. This colouring is also commonly called skewbald. In modern usage in British English, skewbald and piebald horses are collectively referred to as coloured, while in North American English, the term pinto is used to describe the colour pattern.
Bay is a hair coat color of horses, characterized by a reddish-brown or brown body color with a black point coloration on the mane, tail, ear edges, and lower legs. Bay is one of the most common coat colors in many horse breeds.
A pinto horse has a coat color that consists of large patches of white and any other color. Pinto coloration is also called paint, particolored, or in nations that use British English, simply coloured. Pinto horses have been around since shortly after the domestication of the horse.
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 change throughout the horse's lifetime, unless the horse also carries the gray gene. It is a dominant gene, so any tobiano horse must have at least one parent who carries the tobiano gene.
The Tovero coloration is a mix of tobiano and overo colorations in Pinto horses and American Paint Horses. The genetics of pinto coloration are not always fully understood, and some horses have a combination of patterns that does not fit cleanly in either category.
Horses exhibit a diverse array of coat colors and distinctive markings. A specialized vocabulary has evolved to describe them.
A white horse is born predominantly white and stays white throughout its life. A white horse has mostly pink skin under its hair coat, and may have brown, blue, or hazel eyes. "True white" horses, especially those that carry one of the dominant white (W) genes, are rare. Most horses that are commonly referred to as "white" are actually "gray" horses whose hair coats are completely white. Gray horses may be born of any color and their hairs gradually turn white as time goes by and take on a white appearance. Nearly all gray horses have dark skin, except under any white markings present at birth. Skin color is the most common method for an observer to distinguish between mature white and gray horses.
Sabino describes a distinct pattern of white spotting in horses. In general, Sabino patterning is visually recognized by roaning or irregular edges of white markings, belly spots, white extending past the eyes or onto the chin, white above the knees or hocks, and "splash" or "lacy" marks anywhere on the body. Some sabinos have patches of roan patterning on part of the body, especially the barrel and flanks. Some sabinos may have a dark leg or two, but many have four white legs. Sabino patterns may range from slightly bold face or leg white markings—as little as white on the chin or lower lip—to horses that are fully white.
The American Paint Horse Association (APHA) is a breed registry for the American Paint Horse. It is currently headquartered in Fort Worth, Texas. It was founded in 1965 with the merging of two different color breed registries that had been formed to register pinto-colored horses of Quarter Horse bloodlines. One of these organizations was the American Paint Quarter Horse Association and the other was the American Paint Stock Horse Association.
A cropout, crop-out or crop out is a horse with body spots, including pinto or leopard complex spotting, or "high white" horse markings, with a sire and dam who both appeared to have been solid-colored. There are several variations in the definition, depending on the breed registry involved. There are multiple genetic reasons that may cause a horse to be a cropout. Sometimes solid-colored horses throw cropouts because some spotting patterns are recessive genes that are not necessarily expressed unless the relevant allele is inherited from both parents. Other times a gene may be a dominant or incomplete dominant but so minimally expressed that the horse looks solid but can pass flashy color on to its offspring.
Splashed white or splash is a horse coat color pattern in the "overo" group of spotting patterns that produces pink-skinned, white markings. Many splashed whites have very modest markings, while others have the distinctive "dipped in white paint" pattern. Blue eyes are a hallmark of the pattern, and splash may account for otherwise "solid" blue-eyed horses. Splashed white occurs in a variety of geographically divergent breeds, from Morgans in North America to Kathiawari horses in India. The splashed white pattern is also associated with congenital deafness, though most splashed whites have normal hearing. Splashed white can be caused by multiple variants across two different genes, for which genetic testing is available.
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 to the discovery of the W allelic series, many of these patterns were described by the term sabino, which is still used by some breed registries.
The Pinto Horse Association of America (PtHA) registers horses, utility horses, ponies and miniature horses of various pedigrees with certain kinds of pinto coat colors. The word pinto is Spanish for "paint." In general terms, pinto can apply to any horse marked with unpigmented pink-skinned, white-haired areas on its coat. The Pinto Horse Association of America provides the owners and riders of pintos with a show circuit and a breed organization. The primary requirement for PtHA registration is coat color; the pinto is not a true breed, but a color breed.
Roan is a horse coat color 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 breeds. While the specific mutation responsible for roan has not been exactly identified, a DNA test can determine zygosity 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 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.
Ann Trommershausen Bowling was an American scientist who was one of the world's leading geneticists in the study of horses, conducting research in the areas of molecular genetics and cytogenetics. She was a major figure in the development of testing to determine animal parentage, first with blood typing in the 1980s and then DNA testing in the 1990s. She later became known for her studies of hereditary diseases in horses and equine coat color genetics, as well as research on horse evolution and the development of horse breeds. She studied the population genetics of feral horses, did considerable work to help preserve the Przewalski's horse, and was one of the founding members of the international project to map the horse genome. She was an adjunct professor at the University of California, Davis (UCD), and at the time of her death in 2000 was the executive associate director of the Veterinary Genetics Laboratory (VGL) there. Her unexpected death on December 8, 2000, at age 57 was attributed to a massive stroke.
No true albino mutation of the color gene is known among horses, though several varieties of white horse are popularly known as albinos.