The Scottish Fold is a breed of domestic cat with a natural dominant gene mutation that affects cartilage throughout the body, causing the ears to "fold", bending forward and down towards the front of the head, which gives the cat what is often described as an "owl-like" appearance.[1]

Scottish Fold
Lilac coloured adult Scottish Fold, displaying round face, round eyes and forward-folded ears
Other namesScot Fold
Origin Scotland
Breed standards
Not recognised by FIFe and GCCF
Domestic cat (Felis catus)

Originally called lop-eared or lops after the lop-eared rabbit, Scottish Fold became the breed's name in 1966.[1] Depending on registries, longhaired Scottish Folds are varyingly known as Highland Fold, Scottish Fold Longhair, Longhair Fold and Coupari.

All Fold cats are affected by osteochondrodysplasia (OCD) a developmental abnormality that affects cartilage and bone development throughout the body. This condition causes the ear fold in the breed and studies point to all Fold cats being affected by it. Fold cats therefore have malformed bone structures and can develop severe painful degenerative joint diseases at an early age.[2] Due to these health conditions, breeding Fold cats is prohibited in several countries and some major cat registries do not recognise the cat breed.

History Edit

Origin Edit

The original Scottish Fold was a white barn cat named Susie, who was found at a farm near Coupar Angus in Perthshire, Scotland, in 1961. Susie's ears had an unusual fold in their middle, making her resemble an owl. When Susie had kittens, two of them were born with folded ears, and one was acquired by William Ross, a neighbouring farmer and cat-fancier.[3] Ross registered the breed with the Governing Council of the Cat Fancy (GCCF) in the United Kingdom in 1966 and started to breed Scottish Fold kittens with the help of geneticist Pat Turner.[4] The breeding programme produced 76 kittens in the first three years – 42 with folded ears and 34 with straight ears. The conclusion from this was that the ear mutation is due to a simple dominant gene.[5]

Susie's only reproducing offspring was a female Fold named Snooks who was also white; a second kitten was neutered shortly after birth. Three months after Snooks' birth, Susie was killed by a car. All Scottish Fold cats share a common ancestry to Susie.

Acceptance Edit

The breed was not accepted for showing in Europe and the GCCF withdrew registrations in 1971 due to crippling deformity of the limbs and tail in some cats and concerns about genetic difficulties and ear problems such as infection, mites, and deafness, but the Folds were exported to America and the breed continued to be established using crosses with British Shorthairs and American Shorthairs. Since the initial concerns were brought, the Fold breed has not had the mite and infection problems, though wax buildup in the ears may be greater than in other cats.[1] The concerns about deformities may have been caused by osteochondrodysplasia, which causes abnormalities in bone and cartilage throughout the body.

Popularity Edit

The rare distinctive physical traits of the breed, combined with their reputation as unusually loving companions, make Folds highly sought-after pets, with Fold kittens typically costing considerably more than kittens of more common breeds.[6] Scottish folds are also popular among celebrities, one of them being American singer Taylor Swift, who owns two Scottish fold cats named Meredith Grey (the titular character of the medical drama series Grey's Anatomy), and Olivia Benson (the protagonist of the police drama series Law & Order: Special Victims Unit).[7]

Breeding ban Edit

In order to protect animal welfare, several countries and states have prohibited breeding with Scottish folds, including the Netherlands in 2014,[8] Austria in 2020,[9][10] Flanders (Belgium) in 2021,[11] Victoria (Australia),[12] Norway in 2023,[13] and even the birthplace of the breed, Scotland.[14][15] Some countries have also banned selling[11] of Scottish fold cats, or the breeding with any cat that bears the gene mutation resulting in osteochondrodysplasia, so even breeding with some of the Scottish straights.[8] Potential parent cats can be tested for this osteochondrodysplasia mutation before breeding.[16]

Furthermore, some of the major cat registries, such as the GCCF[17] and FIFé,[18] do not recognise, nor allow for the registry of, Scottish folds due to their health issues.

Characteristics Edit

Scottish Fold longhair kitten

Ears Edit

Scottish Fold with straight ears

Scottish Fold kittens that do not develop folded ears are known as Scottish Straights.[1] The original cats had only one fold in their ears, but due to selective breeding, breeders have increased the fold to a double or triple crease that causes the ear to lie totally flat against the head.[citation needed]

The breed's distinctive folded ears are produced by an incompletely dominant gene that affects the cartilage of the ears, causing the ears to fold forward and downward, giving a cap-like appearance to the head. Smaller, tightly folded ears set in a cap-like fashion are preferred to a loose fold and larger ear. The large, round eyes and rounded head, cheeks, and whisker pads add to the overall rounded appearance. Despite the folded ears, folds still use their aural appendages to express themselves—the ears swivel to listen, lie back in anger and prick up when the treat bag rustles.[6]

Body Edit

The Scottish Fold is a medium to large sized cat, which can come in any colour, even calico. Males typically weigh 4–6 kg (8.8–13.2 lb), and females weigh 2.7–4 kg (6.0–8.8 lb). The Fold's entire body structure, especially the head and face, is generally rounded, and the eyes large and round. The nose is short with a gentle curve, and the cat's body is well-rounded with a padded look and medium-to-short legs. The head is domed at the top, and the neck very short. The broadly-spaced eyes give the Scottish Fold a "sweet expression".[5] The Scottish Fold's ears are folded hence the name "Scottish Fold".

Coat Edit

Scottish Folds can be either long- or short-haired, and they may have nearly any coat colour or combination of colours (including white).[5] Shorthair Scottish Folds have thick and soft fur, with longhair Folds having longer and exceptionally dense fur around their upper thighs, toes, ears, and tail.[19]

Temperament Edit

Scottish Folds, whether with folded ears or with normal ears, are typically good-natured and placid and adjust to other animals within a household extremely well. They tend to become very attached to their human caregivers and are by nature quite affectionate. Folds also receive high marks for playfulness, grooming and intelligence. Scottish Folds like to be outdoors and enjoy outdoor games and activities. Loneliness is something they heavily dislike.

Habits Edit

Folds are also known for sleeping on their backs.[5] Scottish Folds typically have soft voices and display a complex repertoire of meows and purrs not found in better-known breeds. Folds are also known for sitting with their legs stretched out and their paws on their belly. This position is called the "Buddha Position".[20]

Genetics Edit

An early study suggested that the fold is inherited as an autosomal dominant trait.[21] A later study suggested an incomplete dominance.[22] A cat with folded ears may have either one (heterozygous) or two copies (homozygous) of the dominant fold gene (Fd). A cat with normal ears should have two copies of the normal gene (fd).

Homozygous fold
Fd Fd
Homozygous fold Fd Fd Fd Fd Fd
Fd Fd Fd Fd Fd
Heterozygous fold
Fd fd
Homozygous fold Fd Fd Fd Fd fd
Fd Fd Fd fd Fd
fd fd
Homozygous fold Fd Fd fd Fd fd
Fd Fd fd Fd fd

Mating a homozygous fold with any cat will produce all folds, but because homozygous folds are prone to severe health issues, breeding for them is generally considered unethical. A homozygous to normal mating will produce only heterozygous folds but presumably in ethical breeding programs, there will be no homozygous cats available to breed from.

fd fd
Heterozygous fold Fd Fd fd Fd fd
fd fd fd fd fd

The only generally accepted breeding gives a 50% chance of producing heterozygous folds and 50% chance of producing progeny with normal genes.

Heterozygous fold
Fd fd
Heterozygous fold Fd Fd Fd Fd fd
fd fd Fd fd fd

There is suspicion that some non-fold litters are genetically heterozygous folds but because of very low expression of the gene, appear to be straight-eared. Such kittens may develop folded ears initially which then straighten back out. Because of this there are suggestions by some breeders to avoid mating Folds with straight-eared Scottish Folds but only use British Shorthairs (BSH) as outcross.[23][24] If Scottish Shorthairs are to be used, they should be test mated to a BSH to make sure that they are not genetically folds.[23] If such apparent straight-eared cats are mated with a fold, there is a 75% chance of folds (25% homozygous folds, 50% heterozygous folds) and 25% chance of straight ears.

In 2016 the genetic mutation responsible for the folded ears and the osteochondrodysplasia (OCD) was identified. It was found in a gene encoding a calcium permeable ion channel, transient receptor potential cation channel, subfamily V, member 4 (TRPV4).[25] The mutation is a V342F substitution (c.1024G>T) in the fifth ankyrin repeat within the N-terminal cytoplasmic domain. It was also found in a human patient with metatropic dysplasia.[26]

Health Edit

The typical lifespan of a Scottish Fold is 15 years.[5]

Scottish folds are susceptible to polycystic kidney disease (PKD)[27] and cardiomyopathy.[28] Scottish folds are also prone to degenerative joint disease (a type of arthritis), most commonly affecting the tail, ankles, and knees which can result in reduced range of motion.[19]

Osteochondrodysplasia Edit

Osteochondrodysplasia (OCD) is a developmental abnormality that affects cartilage and bone development throughout the body. This condition causes the ear fold in the breed and, in studies conducted so far, all Fold cats are affected by it.[2] Homozygous Folds are affected by malformed bone structures and develop severe painful degenerative joint diseases at an early age. Some breeders claim that this condition also affects heterozygous Folds, but usually to a much lesser extent and at a later age. Some will be asymptomatic.[citation needed]. There's no scientific proof to this claim.

In a study of Rorden [29] four radiologists, blinded to the ear phenotype, assessed radiographs of 22 Scottish Fold/Straight cats. All cats were genotyped showing the heterozygous mutation in all folded ear cats but not in straight cats. Each reviewer gave on average the folded ear cats a worse "severity score", however the images showed much milder signs than previously published. The authors state that the severity of OCD in heterozygous cats is very variable and subtle. This could be due to other modifier genes or nurture (climate, diet, exercise). So it was shown that the least affected folded ear cat was given identical or less score than the highest rated straight ear cat.

In a case study of Takanosu [30] two Scottish Fold mixed cats with severe exostosis in the hind leg are described. Interestingly both cats were homozygous for the TrpV4 mutation, assuming the parental cats had both the c.1024G>T mutation in the TrpV4 gene. This reinforces the hypothesis that mostly homozygous Scottish Folds are severely affected.[31] On the other hand it is concerning that still Scottish Fold cats are bred with each other, also breeding with other cat strains with skeletal abnormalities (Munchkin, American Curl) should be avoided.

While ethical breeders breed Fold/non-Fold and not Fold/Fold (in the same way Munchkins are bred) to avoid producing homozygous Folds,[5] because heterozygous Folds can also develop progressive arthritis of varying severity, some researchers recommend abandoning the breeding of Fold cats entirely.[21][32] For this reason, the breed is not accepted by either the Governing Council of the Cat Fancy[33] or the Fédération Internationale Féline (FIFé).[34] CFA breeders have stated that using only Fold to non-Fold breeding has eliminated problems with stiff tails, shortened tails and bone lesions.[1] In the FIFé discussion, the representative for British breeders claimed that they were not seeing the problem in their cats, and that the study which showed that all heterozygous also have the condition had a small sample size. An offer of free X-ray radiography was presented to 300 breeders to find a Fold cat with healthy hind legs, but it was never taken up.[35][36] A similar offer was set up by the World Cat Federation together with researcher Leslie Lyons but there was also no response. FIFé stated that they will not consider recognizing Scottish Folds if breeders will not allow their breed to be scrutinized.[37]

In a report on Scottish Folds, the Breed Standards Advisory Council (BSAC) for New Zealand Cat Fancy (NZCF) states that "Breeders may not have appreciated the strength of the evidence that heterozygous cats can and do develop [feline] OCD." While research shows that all heterozygous Folds develop OCD, and anecdotal evidence shows that heterozygous Folds can and do develop OCD, they do not show whether mildly affected parents are more likely to have mildly affected offspring. They also do not show what percentage of Folds are severely affected. The report states that there is not enough information to justify banning Scottish Fold matings, but enough to justify a level of concern.[23] Recommended guidelines include:

  • A requirement for periodic vet examination of breeding cats for any evidence of lameness, stiffness, or pain—breeding cats with signs to be desexed.
  • A requirement for periodic X-rays of breeding cats and comparison of X-ray evidence with clinical symptoms, possibly leading to a requirement that cats with a specified degree of skeletal change to be desexed.
  • Requesting the agreement of pet owners to be periodically contacted by the NZCF or by a researcher, to provide reports about the health of their cat.
  • All information to be reported/submitted to the BSAC to allow information to be collated to give an overall picture of FOCD in Scottish Folds in NZ.
  • Requirements to be in place for a minimum of 5 years to enable tracking of the health of Folds over time.

The Cat Who Went to Paris Edit

The short novel The Cat Who Went to Paris by Peter Gethers features "the most famous Scottish Fold" according to Grace Sutton of The Cat Fanciers' Association.[1] The book documents the life of Gethers and his Fold, Norton, from their first meeting to Norton's eventual death and Gethers' experiences after the loss.

See also Edit

References Edit

  1. ^ a b c d e f Grace Sutton (31 May 1999). "Breed article: Scottish Folds". Cat Fanciers' Association. Archived from the original on 29 April 2019. Retrieved 29 January 2012.
  2. ^ a b "Genetic welfare problems of companion animals: Scottish Fold: Osteochondrodysplasia". Universities Federation for Animal Welfare. 2011. Archived from the original on 19 March 2012. Retrieved 2 February 2012.
  3. ^ "Scottish Fold Cat Breed Information, Pictures, Characteristics & Facts". CatTime. Retrieved 7 July 2018.
  4. ^ "Scottish Fold Profile - History, Appearance & Temperament - Cat World". 26 February 2017. Retrieved 7 July 2018.
  5. ^ a b c d e f "Scottish Fold FAQ". 30 November 1971. Retrieved 3 October 2009.
  6. ^ a b "Choosing a Scottish Fold". Pet Place. Retrieved 3 October 2009.
  7. ^ Dailey, Hannah. "Taylor Swift's Cat Olivia Benson Is Reportedly Worth $97 Million". Billboard. Penske Media Corporation. Retrieved 31 January 2023.
  8. ^ a b Netherlands Food and Consumer Product Safety Authority (NVWA) (31 October 2022). "Fokken met katten - Honden en katten". (in Dutch). Retrieved 1 April 2023.
  9. ^ Tierschutzombudsstelle Wien. "Tierschutzombudsstelle Wien - Qualzucht bei Katzen". (in German). Retrieved 1 April 2023.
  10. ^ "RIS - Tierschutzgesetz - Bundesrecht konsolidiert, Fassung vom 01.04.2023". Retrieved 1 April 2023.
  11. ^ a b "Verbod op de kweek en het verhandelen van fold katten". (in Dutch). Retrieved 1 April 2023.
  12. ^ Department of Jobs, Precincts and Regions (6 July 2020). "Code of Practice for the Breeding of Animals with Heritable Defects that Cause Disease - Agriculture". Agriculture Victoria. Retrieved 1 April 2023.
  13. ^ Løkkevik, Ole (4 July 2023). "Mattilsynet: Avl på katterasen Scottish Fold er i strid med dyrevelferdsloven". VG (in Norwegian). Retrieved 4 July 2023.
  14. ^ Schofield, Claire (2019). "Scottish fold cats: why the adorable breed is banned in Scotland but is huge in the US and Japan". The Scotsman.
  15. ^ The Scottish Parliament (2018). "Question reference: S5W-15391". Retrieved 1 April 2023.
  16. ^ "Scottish Fold | Veterinary Genetics Laboratory". Retrieved 1 April 2023.
  17. ^ Governing Council of the Cat Fancy (2021). "Cat Breeds not recognised by GCCF but with some recognition by WCC members" (PDF).
  18. ^ Fédération Internationale Féline (2023). "Breed standards". Retrieved 1 April 2023.
  19. ^ a b "Scottish Fold Cat Breed Information, Pictures, Characteristics & Facts." CatTime, Accessed 26 September 2017.
  20. ^ "Scottish Fold".
  21. ^ a b Malik, R.; Allan, G.S.; Howlett, C.R.; Thompson, D.E.; James, G.; McWhirter, C.; Kendall, K. (February 1999). "Osteochondrodysplasia in Scottish Fold cats". Aust. Vet. J. Vol. 77, no. 2. pp. 85–92. PMID 10078353.
  22. ^ Takanosum M, Takanosu T, Suzuki H, Suzuki K (April 2008). "Incomplete dominant osteochondrodysplasia in heterozygous Scottish Fold cats". The Journal of Small Animal Practice. 49 (4): 197–9. doi:10.1111/j.1748-5827.2008.00561.x. PMID 18339089.
  23. ^ a b c "Minutes of Executive Council Meeting No: 28 (doc)". New Zealand Cat Fancy. 24–25 September 2011. Retrieved 3 February 2012.
  24. ^ Sutton, Grace. "Breed Article: Scottish Folds". The Cat Fanciers’ Association. Retrieved 7 December 2013.
  25. ^ Gandolfi, B.; Alamri, S.; Darby, W. G.; et al. (2016). "A dominant TRPV4 variant underlies osteochondrodysplasia in Scottish fold cats". Osteoarthritis and Cartilage. 24 (8): 1441–1450. doi:10.1016/j.joca.2016.03.019. PMID 27063440.
  26. ^ Dai, J.; Kim, O. H.; Cho, T. J.; et al. (2010). "Novel and recurrent TRPV4 mutations and their association with distinct phenotypes within the TRPV4 dysplasia family" (PDF). Journal of Medical Genetics. 47 (10): 704–709. doi:10.1136/jmg.2009.075358. PMID 20577006. S2CID 925054.
  27. ^ Shirley Little. "Polycystic Kidney Disease (PKD)". Retrieved 3 October 2009.
  28. ^ Shirley Little. "Cardiomyopathy". Retrieved 3 October 2009.
  29. ^ Rorden, Chris; Griswold, Marilee C.; Moses, Nan; Berry, Clifford R.; Keller, G. Gregory; Rivas, Rudy; Flores-Smith, Helen; Shaffer, Lisa G.; Malik, Richard (2021). "Radiographical Survey of Osteochondrodysplasia in Scottish Fold Cats caused by the TRPV4 gene variant". Human Genetics. 140 (11): 1525–1534. doi:10.1007/s00439-021-02337-5. PMID 34406467. S2CID 237198301.
  30. ^ Takanosu, Masamine; Hattori, Yuki (2020). "Osteochondrodysplasia in Scottish Fold cross-breed cats". Journal of Veterinary Medical Science. 82 (12): 1769–1772. doi:10.1292/jvms.20-0299. PMC 7804039. PMID 33162427.
  31. ^ Takanosu, M.; Takanosu, T.; Suzuki, H.; Suzuki, K. (2008). "Incomplete dominant osteochondrodysplasia in heterozygous Scottish Fold cats". The Journal of Small Animal Practice. 49 (4): 197–199. doi:10.1111/j.1748-5827.2008.00561.x. PMID 18339089.
  32. ^ Malik, Richard (2003). "Genetic Disorders of Cats". World Small Animal Veterinary Association World Congress Proceedings, 2003. Retrieved 20 December 2009.
  33. ^ "The GCCF says Health Comes First" Archived 14 May 2007 at the Wayback Machine, the Governing Council of the Cat Fancy
  34. ^ "Breeding and Registration Rules (date of issue: 01.01.2008)" — as described in §2.7.3 "Genetic Diseases" Archived 20 August 2008 at the Wayback Machine, Fédération Internationale Feline
  35. ^ "FIFe decision not to accept Scottish Fold breed in May 2003". Retrieved 2 February 2012.
  36. ^ "Email from ScottishFoldandScottishShorthairGroup". 2008. Retrieved 2 February 2012.
  37. ^ "Minutes of the Business Meeting of the World Cat Congress: Arnhem, the Netherlands: 14th April. 2009". Retrieved 2 February 2012.

Reference 13 needs a complete link to PubMed

Dai J, Kim OH, Cho TJ, Schmidt-Rimpler M, Tonoki H, Takikawa K,et a. Novel and recurrent TRPV4 mutations and their association with distinct phenotypes within the TRPV4 dysplasia family. J Med Genet. 2010 Oct;47(10):704-9. doi: 10.1136/jmg.2009.075358. Epub 2010 Jun 24. PMID: 20577006.

External links Edit