Splashed white

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.^[1]^[2]

Characteristics

The splashed white pattern is characterized by the appearance of having been dipped, feet-first, into white paint. Blue eyes are common, but not universal.^[3]^[4]^[5]^[6] The margins of the white markings are crisp, smooth, blocky, and well-defined.^[7]^[6] The head and legs are white, and the tail is often white or white-tipped.^[7]^[5]^[6] The underside of the body is white, and a connected white patch often spreads smoothly up either side of the thorax.^[6] On its own, the splashed white pattern is seldom responsible for white markings that reach the topline, and so it has been categorized as one of the "overo" patterns by Paint horse and Pinto horse registries.^[3]

Minimal splashed whites

The minimal expression of the splashed white pattern can include few or no white markings at all. When only minimal markings are present, other qualities can belie splashed white. Stars and snips may lopsided, off-center, or otherwise strangely placed. Blazes are usually blocky or straight-edged, and bottom-heavy. Splashed white blazes may also be crooked or skewed to one side.^[5] Sabinos with a bold blaze almost invariably have white on the lip or chin, and this is not the case for the facial white of a minimally marked splash; the upper lip typically remains colored.^[5]^[8] Leg markings in a minimally marked splashed white range from hind coronets to high-whites on all four, or marked hindlegs and unmarked forelegs. While sabino leg markings are often tapering with distal patches, the leg markings on a splashed white usually have crisp borders.^[3]

The most reliable identifier of the splashed white pattern is one or more blue, or parti-colored eyes.^[7]^[3]^[5]

Classification

Some breed registries make a distinction between pinto horses and solid horses which may have white markings.

Breed registries for which minimum or maximum white markings are a factor in registration have created imaginary lines to simplify the selection process:

From the ear, to the eye, to the corner of the mouth, to the chin groove,
the knee on the foreleg, and
the hock on the hindleg.

White markings extending past these lines are considered "pinto", "paint" or "colored" while white markings which do not cross these lines are not considered to suggest these traits. However, horses without "excessive white markings" can still have the potential to produce "high white" or distinctly spotted offspring. Splashed white horses have produced generations of "solid" horses, followed by an errant, classically marked splashed white. Discreetly marked splashed whites are responsible for some families of cropout American Quarter Horses.^[3]^[4]

Splashed white mimics

As sabino-type markings also originate on the underside, some splashed whites can be mistaken for cleanly marked sabinos.^[7] Both patterns can be present on the same horse, but splashed white markings are crisp and blocky, and horizontally distributed. In particular, the face markings of splashed whites are straight-edged and bottom heavy, whereas those of sabinos are often tapering or feathered, and often vertical in orientation. Splashed white markings also generally have smoother edges than sabino.^[7] The presence of additional white patterning genes can intensify the amount or obscure the characteristics of splashed white markings.^[4]^[5]

In the Gypsy horse, the pattern called "blagdon" is described as "a solid colour with white "splashed" up from underneath."^[9]

Health concerns

Potential health concerns involving splashed white are deafness and, for some rare variants, embryonic viability.

There is some association between splashed white and congenital deafness in horses that are very white, though many or most splashed white horses have normal hearing.^[4]^[3]^[8]^[10] This type of deafness is probably similar to instances of deafness in white or piebald, blue-eyed examples in other species. In white-coated cats and dogs with deafness, an absence of melanocytes in the inner ear leads to death of the hair cells, which are necessary for perceiving sound.^[10] The presence of pigment around the outside of the ears - which most splash horses have - does not prove the presence of that necessary pigment in the inner ear. There must be pigment inside the inner ear to prevent this problem.^[2] Domestic horses often cope well with deafness, and deaf horses may go undiagnosed. Some deaf horses are more skittish than normal, while others are distinctly calmer. Deafness in horses can be diagnosed by brainstem auditory evoked potential (BAEP), which is minimally invasive and requires no sedation and minimal restraint.^[10] While blue eyes and a white face are often associated with deafness in other species, apron-faced, non-splash horses are not known to be deaf. Nor is the presence of one blue, one normal eye indicative of unilateral deafness. The case horse in Hardland's 2006 case study had one blue eye, while the other was parti-colored, but the horse was bilaterally deaf.^[10]

Some variants of splashed white may produce non-viable embryos if homozygous. SW1 and SW2 have been found in homozygous form, but SW3, SW4, SW5, and SW6 may or may not be homozygous lethal.^[4]^[1]^[2] At present, this cannot be ruled in or out.

In other species, changes to PAX3 (where SW2 and SW4 are found) can cause cleft palates and other congenital defects.^[11]

Inheritance

This American Paint Horse has a white tail and a large, connected area of white across the lower part of her body, suggesting splash genes. The jagged edges are possibly due to sabino traits.

DNA tests exist for six forms of splashed white across two different genes. SW1, SW3, SW5, and SW6 are alleles of MITF, while SW2 and SW4 are alleles of PAX3.^[1]

SW1 is the most common known variant. It is incomplete dominant, meaning it has different effects depending on whether one copy or two copies are present. Horses heterozygous for SW1 (one copy) may have a minimal splash pattern, or may be completely solid. Horses homozygous for SW1 (two copies) have the more extensive classic splashed white pattern.^[5]^[12]

The splashed white pattern was first studied in Finnish Drafts and Welsh ponies by Klemola in 1933.^[13] He divided pinto patterns into a dominant "piebald", describing patterns now known to come from tobiano, frame overo, and sabino, and a recessive "splashed white". His splashed white appears to be describing only the more extensive form of the pattern seen when two copies of SW1 are present.

Originally believed to be very rare outside of Europe, splash is turning out to be more common than previously thought, possibly due to the pattern's tendency to masquerade as modest markings. For example, minimally marked splashed whites have been responsible for cropouts among American Quarter Horses.^[3]

There may be additional forms not yet mapped. In addition to breeds identified as carrying the SW1, -2 or -3 alleles, color patterns described as splashed white but not yet verified via genetic testing have also been identified in American Saddlebreds, Shire horse, Clydesdales, and the Irish Tinker or Gypsy horse.^[3]^[4]^[5]

It is likely that many splashed whites go unidentified, whether because their markings are too minimal to register as "pinto" with breeders, confusion with sabino, or the confounding effect of multiple white spotting patterns.^[3]^[5] Just as the presence of additional white patterning genes can increase the amount of white, other genes may be responsible for restricting the expression of splash.^[5]

The genetic mechanisms behind certain traits, such as blue eyes in a few Arabian horses with otherwise minimal markings, have yet to be identified, though there is speculation that splash genes could be involved.^[14]

Alleles

Informal name	Technical name	Gene
SW1	MITF^prom1	MITF
SW2	PAX3^C70Y	PAX3
SW3	MITF^C280Sfs*20	MITF
Macchiato	MITF^N310S	MITF
SW4	PAX3:p.Pro32Arg	PAX3
SW5	MITF^del	MITF
SW6	SW6	MITF
SW7

^[15]^[16]^[17]^[12]

Prevalence

SW1 is the most prevalent. It appears to be several hundred years old and predates the development of modern breeds. SW1 has been identified in the American Quarter Horse, American Paint Horse, Icelandic horse, Miniature horse, Morgan horse, Shetland pony, Trakehner, and Finnhorse.^[15]^[18]^[19]

SW2 is a PAX3 mutation mistakenly thought to originate with a single Quarter Horse mare foaled in 1987. However, it seems it is actually older, and it has been found in Quarter Horses and Paint horses as well as Noriker and Lipizzan horses.^[18]^[20]^[21]

SW3 has been exclusively found in certain lines of Quarter horses and Paints, and is very rare.^[2]^[4]^[18]

Another allele of splash informally called "Macchiato" has only been found in a single Franche-Montagne horse, probably a spontaneous mutation, and that individual was sterile.^[4]^[15]

SW4 was found in a family of Appaloosa horses.^[16]

SW5 was found in a family of American Paint Horses.^[17]

SW6 is thought to originate with a dual registered Paint horse and Quarter horse stallion.^[12]

Some horses of Quarter Horse lineage carry more than one splash mutation.^[4]^[15]

References

^ ^a ^b ^c "UC Davis Splashed White". Retrieved 2019-05-31.
^ ^a ^b ^c ^d Splashed White Overo (SW1, SW2, SW3)
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ "Splashed white". Genetic Equation. American Paint Horse Association. Archived from the original on 2014-08-22.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Bailey, Ernest Frank; Brooks, Samantha A. (2013). Horse Genetics. CABI. pp. 73–76. ISBN 9781780643298. Retrieved 7 December 2014.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Laura Behning. "Splashed White". Morgan Colors. Retrieved 2009-01-10.
^ ^a ^b ^c ^d Vrotsos, Paul D.; Elizabeth M. Santschi; James R. Mickelson (2001). "The Impact of the Mutation Causing Overo Lethal White Syndrome on White Patterning in Horses" (PDF). AAEP Proceedings. 47. American Association of Equine Practitioners: 385–391.
^ ^a ^b ^c ^d ^e Brooks, S. Studies of genetic variation at the KIT locus and white spotting patterns in the horse. Ph.D. dissertation, University of Kentucky, 2006. [1]
^ ^a ^b Dan Phillip Sponenberg (2003). "5 Patterns Characterized by Patches of White". Equine Color Genetics (2 ed.). Blackwell Publishing. pp. 85–6. ISBN 0-8138-0759-X.
^ "Gypsy Vanner Breed Standard" (PDF). Gypsy Vanner Horse Society [GVHS]. Archived from the original (PDF) on 2012-12-24. Retrieved 2013-02-28.
^ ^a ^b ^c ^d Harland, Malte M.; Allison J. Stewart; Arvle E. Marshall; Ellen B. Belknap (2006). "Diagnosis of deafness in a horse by brainstem auditory evoked potential". Canadian Veterinary Journal. 47 (2): 151–4. PMC 1345730. PMID 16579041.
^ Wu, Meilin; Li, Jun; Engleka, Kurt A.; Zhou, Bo; Lu, Min Min; Plotkin, Joshua B.; Epstein, Jonathan A. (2008). "Persistent expression of Pax3 in the neural crest causes cleft palate and defective osteogenesis in mice". Journal of Clinical Investigation. doi:10.1172/JCI33715. PMC 2381747. PMID 18483623.
^ ^a ^b ^c Bellone, Rebecca R.; Tanaka, Jocelyn; Magdesian, K Gary (2020). "A De Novo MITF Deletion Explains a Novel Splashed White Phenotype in an American Paint Horse". Journal of Heredity. 111 (3): 287–293. doi:10.1093/jhered/esaa009. PMC 7238438. PMID 32242630.
^ Klemola, V (1933). "The "pied" and "splashed white" patterns in horses and ponies". Journal of Heredity. 24 (2): 65–69. doi:10.1093/oxfordjournals.jhered.a103713.
^ "Should dominant white be considered as sabino white?". Horsegenetics.com. Retrieved 27 December 2014.
^ ^a ^b ^c ^d Hauswirth, Regula; Haase, Bianca; et al. (April 12, 2012). "Mutations in MITF and PAX3 Cause "Splashed White" and Other White Spotting Phenotypes in Horses". PLOS Genetics. 8 (4): e1002653. doi:10.1371/journal.pgen.1002653. PMC 3325211. PMID 22511888.
^ ^a ^b Hauswirth, Regula; Jude, Rony; Haase, Bianca; Bellone, Rebecca R.; Archer, Sheila; Holl, Heather; Brooks, Samantha A.; Tozaki, Teruaki; Penedo, Maria Cecilia T.; Rieder, Stefan; Leeb, Tosso (2013). "Novel variants in the KIT and PAX3 genes in horses with white-spotted coat colour phenotypes". Animal Genetics. 44 (6): 763–765. doi:10.1111/age.12057. PMID 23659293.
^ ^a ^b Henkel, J.; Lafayette, C.; Brooks, S. A.; Martin, K.; Patterson-Rosa, L.; Cook, D.; Jagannathan, V.; Leeb, T. (2019). "Whole-genome sequencing reveals a large deletion in the MITF gene in horses with white spotted coat colour and increased risk of deafness" (PDF). Animal Genetics. 50 (2): 172–174. doi:10.1111/age.12762. PMID 30644113. S2CID 58656314.
^ ^a ^b ^c "Splashed-Whte". Veterinary Genetics Laboratory. University of California-Davis. Retrieved 7 December 2014.
^ Alerini, Leena: Uusia värihevosia, uusia testiohjeita Archived 2013-07-02 at the Wayback Machine, 27.6.2013
^ Hauswirth, Regula; Haase, Bianca; Blatter, Marlis; Brooks, Samantha A.; Burger, Dominik; Drögemüller, Cord; Gerber, Vincent; Henke, Diana; Janda, Jozef; Jude, Rony; Magdesian, K. Gary; Matthews, Jacqueline M.; Poncet, Pierre-André; Svansson, Vilhjálmur; Tozaki, Teruaki; Wilkinson-White, Lorna; Penedo, M. Cecilia T.; Rieder, Stefan; Leeb, Tosso (Aug 19, 2019). "Correction: Mutations in MITF and PAX3 Cause "Splashed White" and Other White Spotting Phenotypes in Horses". PLOS Genetics. 15 (8): e1008321. doi:10.1371/journal.pgen.1008321. PMC 6677290. PMID 31374075.
^ Druml, T.; Grilz-Seger, G.; Neuditschko, M.; Horna, M.; Ricard, A.; Pausch, H.; Brem, G. (2018). "Novel insights into Sabino1 and splashed white coat color patterns in horses". Animal Genetics. 49 (3): 249–253. doi:10.1111/age.12657. PMC 6001536. PMID 29635692.

External links

Deaf Horse Association Official Deaf Horse Association's webpage

[UCDavisSplash-1] "UC Davis Splashed White". Retrieved 2019-05-31.

[AmGen-2] Splashed White Overo (SW1, SW2, SW3)

[aphageneticeqsplash-3] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ "Splashed white". Genetic Equation. American Paint Horse Association. Archived from the original on 2014-08-22.

[Bailey-4] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Bailey, Ernest Frank; Brooks, Samantha A. (2013). Horse Genetics. CABI. pp. 73–76. ISBN 9781780643298. Retrieved 7 December 2014.

[morgansplash-5] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Laura Behning. "Splashed White". Morgan Colors. Retrieved 2009-01-10.

[vrotsos2001-6] Vrotsos, Paul D.; Elizabeth M. Santschi; James R. Mickelson (2001). "The Impact of the Mutation Causing Overo Lethal White Syndrome on White Patterning in Horses" (PDF). AAEP Proceedings. 47. American Association of Equine Practitioners: 385–391.

[brooksdissertation-7] Brooks, S. Studies of genetic variation at the KIT locus and white spotting patterns in the horse. Ph.D. dissertation, University of Kentucky, 2006. [1]

[sponenberg2003-8] Dan Phillip Sponenberg (2003). "5 Patterns Characterized by Patches of White". Equine Color Genetics (2 ed.). Blackwell Publishing. pp. 85–6. ISBN 0-8138-0759-X.

[Vanners_Standard-9] "Gypsy Vanner Breed Standard" (PDF). Gypsy Vanner Horse Society [GVHS]. Archived from the original (PDF) on 2012-12-24. Retrieved 2013-02-28.

[harland2006-10] Harland, Malte M.; Allison J. Stewart; Arvle E. Marshall; Ellen B. Belknap (2006). "Diagnosis of deafness in a horse by brainstem auditory evoked potential". Canadian Veterinary Journal. 47 (2): 151–4. PMC 1345730. PMID 16579041.

[Wu2008-11] Wu, Meilin; Li, Jun; Engleka, Kurt A.; Zhou, Bo; Lu, Min Min; Plotkin, Joshua B.; Epstein, Jonathan A. (2008). "Persistent expression of Pax3 in the neural crest causes cleft palate and defective osteogenesis in mice". Journal of Clinical Investigation. doi:10.1172/JCI33715. PMC 2381747. PMID 18483623.

[Magdesian2020-12] Bellone, Rebecca R.; Tanaka, Jocelyn; Magdesian, K Gary (2020). "A De Novo MITF Deletion Explains a Novel Splashed White Phenotype in an American Paint Horse". Journal of Heredity. 111 (3): 287–293. doi:10.1093/jhered/esaa009. PMC 7238438. PMID 32242630.

[klemola-13] Klemola, V (1933). "The "pied" and "splashed white" patterns in horses and ponies". Journal of Heredity. 24 (2): 65–69. doi:10.1093/oxfordjournals.jhered.a103713.

[Horsegenetics-14] "Should dominant white be considered as sabino white?". Horsegenetics.com. Retrieved 27 December 2014.

[Hauswirth-15] Hauswirth, Regula; Haase, Bianca; et al. (April 12, 2012). "Mutations in MITF and PAX3 Cause "Splashed White" and Other White Spotting Phenotypes in Horses". PLOS Genetics. 8 (4): e1002653. doi:10.1371/journal.pgen.1002653. PMC 3325211. PMID 22511888.

[Hauswirth2013-16] Hauswirth, Regula; Jude, Rony; Haase, Bianca; Bellone, Rebecca R.; Archer, Sheila; Holl, Heather; Brooks, Samantha A.; Tozaki, Teruaki; Penedo, Maria Cecilia T.; Rieder, Stefan; Leeb, Tosso (2013). "Novel variants in the KIT and PAX3 genes in horses with white-spotted coat colour phenotypes". Animal Genetics. 44 (6): 763–765. doi:10.1111/age.12057. PMID 23659293.

[Henkel2018-17] Henkel, J.; Lafayette, C.; Brooks, S. A.; Martin, K.; Patterson-Rosa, L.; Cook, D.; Jagannathan, V.; Leeb, T. (2019). "Whole-genome sequencing reveals a large deletion in the MITF gene in horses with white spotted coat colour and increased risk of deafness" (PDF). Animal Genetics. 50 (2): 172–174. doi:10.1111/age.12762. PMID 30644113. S2CID 58656314.

[UCD-18] "Splashed-Whte". Veterinary Genetics Laboratory. University of California-Davis. Retrieved 7 December 2014.

[uusia-19] Alerini, Leena: Uusia värihevosia, uusia testiohjeita Archived 2013-07-02 at the Wayback Machine, 27.6.2013

[HauswirthCorrection-20] Hauswirth, Regula; Haase, Bianca; Blatter, Marlis; Brooks, Samantha A.; Burger, Dominik; Drögemüller, Cord; Gerber, Vincent; Henke, Diana; Janda, Jozef; Jude, Rony; Magdesian, K. Gary; Matthews, Jacqueline M.; Poncet, Pierre-André; Svansson, Vilhjálmur; Tozaki, Teruaki; Wilkinson-White, Lorna; Penedo, M. Cecilia T.; Rieder, Stefan; Leeb, Tosso (Aug 19, 2019). "Correction: Mutations in MITF and PAX3 Cause "Splashed White" and Other White Spotting Phenotypes in Horses". PLOS Genetics. 15 (8): e1008321. doi:10.1371/journal.pgen.1008321. PMC 6677290. PMID 31374075.

[Druml2018-21] Druml, T.; Grilz-Seger, G.; Neuditschko, M.; Horna, M.; Ricard, A.; Pausch, H.; Brem, G. (2018). "Novel insights into Sabino1 and splashed white coat color patterns in horses". Animal Genetics. 49 (3): 249–253. doi:10.1111/age.12657. PMC 6001536. PMID 29635692.

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Splashed white

Characteristics

Minimal splashed whites

Classification

Splashed white mimics

Health concerns

Inheritance

Alleles

Prevalence

See also

References

External links

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