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Dominant white or white spotting[1][2] is a group of genetically related coat color conditions in the horse, best known for producing an all-white coat, but also able to produce various forms of white spotting and white markings, several of which are sometimes referred to as sabino.
There are many different forms of dominant white or white spotting; as of 2018 they are labeled W1 through W27[3] and are associated with the KIT gene, along with Sabino 1.[B] The white spotting produced can range from flashy white markings like those made by W20, to patterns similar to Sabino 1, to a fully white or almost fully white horse. White horses are born with unpigmented pink skin and white hair, usually with dark eyes. Under normal conditions, at least one parent must be dominant white to produce dominant white offspring. However, most of the currently-known alleles of dominant white can be linked to a documented spontaneous mutation in a single ancestor. For many of the W alleles, the white coats are, as the name suggests, inherited dominantly,[D] meaning that a horse only needs one copy of the allele to have a white or white spotted coat. Others, such as W20, are incompletely dominant, with horses with two copies having more white than horses with only one. In addition, different alleles which on their own give a white spotted but not completely white horse, such as W5 and W10, can combine to make a horse completely white.
White can occur in any breed, and has been studied in many different breeds. Camarillo White Horses are characterized by their white coats.
White is both genetically and visually distinct from gray and cremello. Dominant white is not the same as lethal white syndrome, nor are white horses "albinos". Tyrosinase negative albinism has never been documented in horses. Some forms of white are thought to result in nonviable embryos when a zygote has two W alleles (is homozygous). However, others, such as W20, are perfectly viable in their homozygous form.
Although the term "dominant white" is typically associated with a pure white coat, such horses may be all-white, near-white, partially white, or exhibit an irregular spotting pattern similar to that of sabino horses.[4] To add to the confusion, at least some horses in each of those groups might be referred to as "dominant white", "white spotted", or "sabino". The amount of white hair depends on which KIT alleles are involved.[5] At birth, most of the white hair is rooted in unpigmented pink skin. The pink skin lacks melanocytes, and appears pink from the underlying network of capillaries. White spotting is not known to affect eye color, and most white horses have brown eyes.[6]
White horses are born with pink skin and a white coat, which they retain throughout their lives.[7] The genetic factors that produce an all-white horse are often also capable of producing a near-white horse, which is mostly white but has some areas that are pigmented normally. Near-white horses most commonly have color in the hair and skin along the topline (dorsal midline) of the horse, in the mane, and on the ears.[4] The color is often interspersed as specks or spots on a white background. In addition, the hooves are usually white, but may have striping if there is pigmented skin on the coronary band just above the hoof.[8][9] In some cases, foals born with residual non-white hair may lose some or all of this pigment with age, without the help of the gray factor.[10]
White spotting from a W allele is difficult to identify visually, as it can range from small white markings in the case of a heterozygous W20 horse all the way to an obvious pinto pattern. In addition, even completely white horses can have genes which by themselves would only give white spotting, such as W20 combined with W22[2] or W5 combined with W10. As such, the only reliable way to find out whether a horse has one of the known white spotting patterns from an allele on KIT is to have it genetically tested.
Dominant white is one of several potential genetic causes for horses with near-white or completely white coats; it may occur through spontaneous mutation, and thus may be found unexpectedly in any breed, even those that discourage excessive white markings. To date, dominant white has been identified in multiple families of Thoroughbreds,[10] American Quarter Horses,[4] Frederiksborg horses,[4] Icelandic horses,[4] Shetland ponies,[10] Franches Montagnes horses,[10] South German Draft horses,[4] and in one family of the Arabian horse.[10] The American White Horse, which is descended primarily from one white stallion crossed on non-white mares, is known for its white coat, as is the Camarillo White Horse.[11][12]
The W locus was mapped to the KIT gene in 2007.[10] KIT is short for "KIT proto-oncogene receptor tyrosine kinase".[13] White spotting is caused by multiple forms, or alleles, of the KIT gene.[10] All horses possess the KIT gene, as it is necessary for survival even at the earliest stages of development. The presence or absence of dominant white is based on the presence of certain altered variants of KIT. Each unique form is called an allele, and for every trait, all animals inherit one allele from each parent. The original or "normal" form of KIT, which is expected in horses without dominant white spotting, is called the "wild type" allele.[A] Thus, a dominant white horse has at least one KIT allele with a mutation associated with dominant white spotting, and either the wild type KIT allele or a second allele on KIT associated with white spotting, which could be one of the ones in the W series or could be Sabino 1.
The KIT gene contains over 2000 base pairs, and a change in any of those base pairs results in a mutant allele.[4] Over forty seven such alleles have been identified by sequencing the KIT genes of various horses.[4] The resultant phenotype of many of these alleles is not yet known, but 29 have been linked to white spotting.[14][5] To date, DNA tests can identify if a horse carries the various identified W alleles, some commercially available.[15]
These alleles do not account for all dominantly inherited white spotting in horses. More KIT alleles are expected to be found with roles in white spotting.[4] Most W alleles each occur within a specific breed or family and arise as spontaneous mutations. The KIT gene itself seems prone to mutation, and so new alleles of W could occur in virtually any breed.[5]
Sabino can refer either specifically to Sabino 1 (SB1) or to a variety of visually similar spotting patterns. SB1 creates a nearly pure white horse when homozygous, which is not true for other sabino patterns. To add to the confusion, white spotting created by several W alleles, such as W5, W15, and W19 creates patterns that historically were called sabino. For that reason, the use of the word "sabino" is evolving. Genetically, Sabino 1 is basically another allele on KIT, [45] and thus is probably able to be classified as a form of dominant white.[46]
In its homozygous form, Sabino 1 can be confused with dominant white alleles such as W1, W2, W3, or W4 that create a white or near-white horse with only one copy. Both dominant white and "Sabino-White" horses are identified by all-white or near-white coats with underlying pink skin and dark eyes, often with residual pigment along the dorsal midline. However, it takes two copies of Sabino 1 to produce a Sabino-white horse, and Sabino 1 is not homozygous lethal.[47]
Initially, dominant white was separated from sabino on the grounds that the former had to be entirely white, while the latter could possess some pigment.[48] However, the 2007 and 2009 studies of dominant white showed that many dominant white alleles produce a range of white phenotypes that include horses with pigmented spots in their hair and skin.[4] Each of the larger families of dominant white studied included pure-white horses, horses described as having "sabino-like" white markings, as well as white horses described as "maximal sabino".[4][10]
More recently, dominant white and sabino were distinguished from one another on the grounds that dominant white alleles produce nonviable embryos in the homozygous state, while Sabino 1 was viable when homozygous.[49] However, not all KIT alleles currently identified as "dominant white" have been proven lethal,[16] and in fact W20 is known to be viable in the homozygous form.[50]
The similarities between Dominant White, Sabino 1, and other forms of sabino may reflect their common molecular origin: The W series and SB1 have both been mapped to KIT. The researchers who mapped Sabino 1 suggested that other sabino-like patterns might also map to KIT.[51] Similarly, major alleles for white leg and facial markings have also been mapped to or near to the KIT gene.[52]
The KIT gene encodes a protein called steel factor receptor, which is critical to the differentiation of stem cells into blood cells, sperm cells, and pigment cells. A process called alternative splicing, which uses the information encoded in the KIT gene to make slightly different proteins (isoforms) for use in different circumstances, may impact whether a mutation on KIT affects blood cells, sperm cells, or pigment cells. Steel factor receptor interacts chemically with steel factor or stem cell factor to relay chemical messages. These messages are used during embryonic development to signal the migration of early melanocytes (pigment cells) from the neural crest tissue to their eventual destinations in the dermal layer. The neural crest is a transient tissue in the embryo that lies along the dorsal line. Melanocytes migrate along the dorsal line to a number of specific sites: near the eye, near the ear, and the top of the head; six sites along each side of the body, and a few along the tail. At these sites, the cells undergo a few rounds of replication and differentiation, and then migrate down and around the body from the dorsal aspect towards the ventral aspect and the limb buds.[53]
The timing of this migration is critical; all white markings, from a small star to a pure white coat, are caused by the failed migration of melanocytes.[54]
A certain degree of the eventual amount of white, and its "design", is completely random. The development of an organism from single-celled to fully formed is a process with many, many steps. Even beginning with identical genomes, as in clones and identical twins, the process is unlikely to occur the same way twice. A process with this element of randomness is called a stochastic process, and cell differentiation is, in part, a stochastic process.[55] The stochastic element of development is partly responsible for the eventual appearance of white on a horse, potentially accounting for nearly a quarter of the phenotype.[56] The research team that studied dominant white cited "subtle variations in the amount of residual KIT protein" as a potential cause for the variability in phenotype of horses with the same allele. They also speculated that variability in the phenotype of horses with W1 might be caused by "different efficacies of [nonsense-mediated decay] in different individuals and in different body regions." That is, some horses destroy more of the mutant KIT protein than others.[4]
Early embryonal lethality, also known as early embryonic death or a non-viable embryo, may occur when the embryo possesses two dominant white alleles, or has the homozygous genotype.[57] The reason for this is that many mutations for W are caused by nonsense mutations, frameshift mutations or DNA deletions, which, if homozygous, would make it impossible to produce a functional KIT protein. However, it is possible that homozygous embryos from alleles of missense and splice site mutations might be viable because they have less effect on gene function.[5] For instance, W1 is a nonsense mutation and it is thought that horses with the genotype W1/W1 would die in utero, while W20 is a missense mutation and living horses with the W20/W20 genotype have been found. A 2013 study also located horses that were compound W5/W20 heterozygotes, almost completely white, essentially with greater depigmentation than could be accounted for by either allele alone.[20]
White horses are potent symbols in many cultures.[58] An array of horse coat colors may be identified as "white", often inaccurately, and many are genetically distinct from "dominant white".
"Albino" horses have never been documented, despite references to so-called "albino" horses.[59][60] Dominant white is caused by the absence of pigment cells (melanocytes), whereas albino animals have a normal distribution of melanocytes.[61] Also, a diagnosis of albinism in humans is based on visual impairment, which has not been described in horses with dominant white nor similar coat colors.[62] In other mammals, the diagnosis of albinism is based on the impairment of tyrosinase production.[63] No mutations of the tyrosinase gene are known in horses, however, cream and pearl colors result from mutations to a protein involved in tyrosinase transport.[64]
Foals with lethal white syndrome (LWS) have two copies of the frame overo gene and are born with white or nearly white coats and pink skin. However, unlike dominant white horses, foals with LWS are born with an underdeveloped colon that is untreatable, and if not euthanized, invariably die of colic within a few days of birth.[75] Horses that carry only one allele of the LWS gene are healthy and typically exhibit the "frame overo" spotting pattern. In cases of "solid" horses with frame overo ancestry, uncertain "overo" (non-tobiano) phenotype, or horses with multiple patterns, the LWS allele can be detected by DNA test.[76]
Mosaicism in horses is thought to account for some spontaneous occurrences of white, near-white, spotted, and roan horses.[77] Mosaicism refers to mutations that occur after the single-cell stage, and therefore affect only a portion of the adult cells.[78] Mosaicism may be one possible cause for the rare occurrence of brindle coloring in horses.[79] Mosaic-white horses would be visually indistinguishable from dominant whites. Mosaicism could produce white or partially white foals if a stem cell in the developing foal underwent a mutation, or change to the DNA, that resulted in unpigmented skin and hair. The cells that descend from the affected stem cell will exhibit the mutation, while the rest of the cells are unaffected.
A mosaic mutation may or may not be inheritable, depending on the cell populations affected.[80] Though this is not always the case, genetic mutations can occur spontaneously in one sex cell of a parent during gametogenesis.[81] In these cases, called germline mutations, the mutation will be present in the single-celled zygote conceived from the affected sperm or egg cell, and the condition can be inherited by the next generation.[77]
Dominant white horses were first described in scientific literature in 1912. Horse breeder William P. Newell described his family of white and near-white horses to researcher A. P. Sturtevant of Columbia University:
"The colour of skin is white or so-called pink, usually with a few small dark specks in skin. Some have a great many dark spots in skin. These latter usually have a few dark stripes in hoofs; otherwise the hoofs are almost invariably white. Those that do not have dark specks in skin usually have glass or watch eyes, otherwise dark eyes ... I have one colt coming one year old that is pure white, not a coloured speck on him, not a coloured hair on him, and with glass [blue] eyes."[9]
Sturtevant and his contemporaries agreed that this colt's blue eyes were inherited separately from his white coat.[82] In 1912, Sturtevant assigned the "white" trait to the White or W locus.[9] At the time there was no means of assigning W to a position on the chromosome, or to a gene.
This family of white horses produced Old King in 1908, a dark-eyed white stallion that was purchased by Caleb R. and Hudson B. Thompson. Old King was bred to Morgan mares to produce a breed of horse known today as the American White Horse.[11] A grandson of Old King, Snow King, was at the center of the first major study of the dominant white coat color in horses, conducted in 1969 by Dr. William L. Pulos of Alfred University and Dr. Frederick B. Hutt of Cornell. They concluded, based on test matings and progeny phenotype ratios, that the white coat was dominantly inherited and embryonic lethal in the homozygous state.[83] Other factors, such as variations in expressivity and the influence of multiple genes, may have influenced the progeny ratios that Pulos and Hutt observed.[84] The white coat of the American White Horse has not yet been mapped.
A 1924 study by C. Wriedt identified a heritable white coat color in the Frederiksborg horse.[85] Wriedt described a range of what he considered to be homozygote phenotypes: all-white, white with pigmented flecks, or weißgraue, which transliterates to "white-gray."[86] The German term for gray horse is schimmel, not weißgraue.[87] Heterozygotes, according to Wriedt, ranged from roaned or diluted to more or less solid white horses. Reviewers, such as Miguel Odriozola, reinterpreted Wriedt's data in successive years, while Pulos and Hutt felt that his work had been "erroneous" because Wriedt never concluded that white was lethal when homozygous.[88]
Other researchers prior to modern DNA analysis developed remarkably prescient theories. The gene itself was first proposed and named W in 1948.[5] In a 1969 work on horse coat colors, A los colores del caballo, Miguel Odriozola suggested that various forms of dominantly inherited white spotting might be arranged sequentially along one chromosome, thus allowing for the
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