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Merle is a genetic pattern that can be in a dog's
coat A coat typically is an outer garment for the upper body as worn by either gender for warmth or fashion. Coats typically have long sleeves and are open down the front and closing by means of buttons, zippers, hook-and-loop fasteners, toggles, a ...
. Merle comes in different colors and patterns and can affect all coat colors. The merle gene creates mottled patches of color in a solid or
piebald 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 backgro ...
coat, blue or odd-colored eyes, and can affect skin
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...
as well. There are two general types of colored patches that will appear in a merle coat: liver (red merle) and black (blue merle). Some breeds that can be affected by this genetic trait are Carea Leonés, Australian Shepherds and Catahoula Leopard Dogs. This genetic trait should be taken seriously when breeding merles. Health issues are more typical and more severe when two merles are bred together, so it is recommended that a merle be bred with a non-merle dog of any other color.


Description

Merle can affect all coat colors. The merle forms of brown and black are usually called "red" (though this is not correct; red and brown are genetically different) and "blue" as patches of blue are formed throughout the coat. Dogs who are recessive red can still be affected by merle, but the patches are either hardly seen or, if the dog is a clear recessive red, are not visible at all. Combinations such as
brindle Brindle is a coat coloring pattern in animals, particularly dogs, cattle, guinea pigs, cats, and, rarely, horses. It is sometimes described as "tiger-striped", although the brindle pattern is more subtle than that of a tiger's coat. Brindle ty ...
and sable merle exist, but are not typically accepted in
breed standards In animal husbandry or animal fancy, a breed standard is a description of the characteristics of a hypothetical or ideal example of a breed. The description may include physical or morphological detail, genetic criteria, or criteria of athletic ...
. In addition to altering base coat color, merle also modifies eye color and coloring on the nose and paw pads. The merle gene modifies the dark pigment in the eyes, occasionally changing dark eyes to blue, or part of the eye to be colored blue. Since merle causes random modifications, however, both dark-eyed, blue-eyed, and odd-colored eyes are possible. Pigmentation on paw pads and nose may be mottled by pink.


Breeds

Merle is a distinguishing marking of several breeds, particularly the
Australian Shepherd The Australian Shepherd is a breed of herding dog from the United States. The name of the breed is technically a misnomer, as it was developed in California in the 19th century; the breed was unknown in Australia at the time. It is claimed that ...
and
Catahoula Leopard Dog The Catahoula Leopard Dog is an American dog breed named after Catahoula Parish, Louisiana. It became the state dog of Louisiana in 1979. It is recognized by the United Kennel Club under the name ''Louisiana Catahoula Leopard Dog'', while the Ame ...
, and appears in others, including the
Koolie The Koolie (also known as the Australian Koolie or by the historic misnomer "German Coolie") is an Australian dog breed. The Koolie is a working or herding dog which has existed in Australia since the early 19th century when it was bred fro ...
in Australia, the
Shetland Sheepdog The Shetland Sheepdog, often known as the Sheltie, is a breed of herding dog that originated in the Shetland Islands of Scotland. The original name was ''Shetland Collie,'' but this caused controversy amongst Rough Collie breeders of the time, so ...
, various
collie Collies form a distinctive type of herding dogs, including many related landraces and standardized breeds. The type originated in Scotland and Northern England. Collies are medium-sized, fairly lightly-built dogs, with pointed snouts. Many ...
breeds, the
Cardigan Welsh Corgi The Cardigan Welsh Corgi (; Welsh for "dwarf dog") is one of two separate dog breeds known as Welsh Corgis that originated in Wales; the other is the Pembroke Welsh Corgi. It is one of the oldest breeds of the British Isles. Cardigan Welsh Co ...
, the
Pyrenean Shepherd The Pyrenean Sheepdog, the Chien de Berger des Pyrénées in French, is a small to medium-sized breed of herding dog from the Pyrenees Mountains region of France. It is found herding flocks of sheep throughout the Pyrenees alongside the much la ...
and the Bergamasco Shepherd. In
Dachshund The dachshund ( or ; German: "badger dog"), also known as the wiener dog, badger dog, and sausage dog, is a short-legged, long-bodied, hound-type dog breed. The dog may be smooth-haired, wire-haired, or long-haired, and comes in a variety of ...
s, the merle pattern is known as "dapple". In
Beauceron The Beauceron is a herding dog breed originating from the plains of Central France. The Beauceron is also known as Berger de Beauce (sheepdog from Beauce) or Bas Rouge (red-stockings). History A French herding breed known for centuries in west ...
s the merle pattern is known as "harlequin". The merle gene also plays a part in producing harlequin
Great Dane The Great Dane is a large sized dog breed originating from Germany. The Great Dane descends from hunting dogs from the Middle Ages used to hunt wild boar and deer, and as guardians of German nobility. It is one of the largest breeds in the world ...
s. Depending on the breed, registry, or country, the pattern may be register-able with a registry but not allowed in conformation, and the pattern may not be recognized by the
breed standard In animal husbandry or animal fancy, a breed standard is a description of the characteristics of a hypothetical or ideal example of a breed. The description may include physical or morphological detail, genetic criteria, or criteria of athletic ...
, disqualifying it from showing. There may also be additional requirements for the pattern such as the dog being allowed to have the pattern but must have completely dark eyes with no blue in them. Merle is sometimes introduced to other
purebred dog A dog breed is a particular strain of dog that was purposefully bred by humans to perform specific tasks, such as herding, hunting, and guarding. Dogs are the most variable mammal on Earth, with artificial selection producing around 450 globally ...
breeds through
crossbreeding A crossbreed is an organism with purebred parents of two different breeds, varieties, or populations. ''Crossbreeding'', sometimes called "designer crossbreeding", is the process of breeding such an organism, While crossbreeding is used to main ...
, but these dogs are not purebred and cannot be registered with any reputable kennel club. In 2020 the UK Kennel Club (the oldest in the World) banned the registration of merle Poodles as it is not and never has been a colour pattern recognised in pure Poodles.


Genetic basis

Merle is actually a
heterozygote 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 ...
of an incompletely dominant
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 ...
. If two such dogs are mated, on the average one quarter of the puppies will be "double merles", which is the common term for dogs homozygous for merle, and a high percentage of these double merle puppies could have eye defects and/or be deaf. Knowledgeable and responsible breeders who want to produce merle puppies mate a merle with a non-merle dog; roughly half the puppies will be merles, and none will have the vision or hearing defects associated with double merle dogs. In January 2006 scientists at
Texas A&M University Texas A&M University (Texas A&M, A&M, or TAMU) is a public university, public, Land-grant university, land-grant, research university in College Station, Texas. It was founded in 1876 and became the flagship institution of the Texas A&M Unive ...
announced the discovery of a mobile genetic unit called a
retrotransposon Retrotransposons (also called Class I transposable elements or transposons via RNA intermediates) are a type of genetic component that copy and paste themselves into different genomic locations (transposon) by converting RNA back into DNA through ...
, responsible for the merle mutation in dogs. Merle can be tested for and identified through DNA. A phantom merle or cryptic merle is one with such small patches of merle—or none at all—that it appears to be a non-merle. This is commonly seen in dogs who are recessive red, clear recessive reds in particular, though patches can still be seen in certain red dogs. In America, a dog with the phantom merle coloring is described as being "cryptic for merle." A cryptic merle bred to a merle may produce merles, cryptic merles, or homozygous merles. Another mutation for cryptic merle has been identified by Dr. Helena Synková, and has been given the working name of "atypical merle". It is recommended that if a breeder is unsure if their dog is merle or not, that they be tested for the merle gene.


Merle modifiers

Certain "modifying genes" work in tandem (co-dominate expressive) with the merle gene to create a completely different look to the pattern. Often mistaken for a "double merle", a harlequin merle (or just harlequin), is a dog that carries both the merle pattern gene and the co-dominate modifying gene (carried on a different locus) for harlequin. This causes most or all of the "blue" to be replaced with white, leaving a dog that is mostly white with black patches. All dogs exhibiting the harlequin pattern are also carriers of the merle gene. Common in
Great Danes The Great Dane is a large sized dog breed originating from Germany. The Great Dane descends from hunting dogs from the Middle Ages used to hunt wild boar and deer, and as guardians of German nobility. It is one of the largest breeds in the world ...
, (and registered as Harlequin) it is less commonly seen in other breeds such as the
Catahoula Leopard Dog The Catahoula Leopard Dog is an American dog breed named after Catahoula Parish, Louisiana. It became the state dog of Louisiana in 1979. It is recognized by the United Kennel Club under the name ''Louisiana Catahoula Leopard Dog'', while the Ame ...
,
Shetland Sheepdog The Shetland Sheepdog, often known as the Sheltie, is a breed of herding dog that originated in the Shetland Islands of Scotland. The original name was ''Shetland Collie,'' but this caused controversy amongst Rough Collie breeders of the time, so ...
and
Collie Collies form a distinctive type of herding dogs, including many related landraces and standardized breeds. The type originated in Scotland and Northern England. Collies are medium-sized, fairly lightly-built dogs, with pointed snouts. Many ...
, where the dogs are registered simply as merle. In 2018, a published paper identified multiple different types of merle that affect coat color differently.


Health issues

Dogs with two copies of the merle gene (homozygous merle or "double merle") have an even higher chance of being deaf and with vision impairments. The UK Kennel Club has acknowledged the health risk associated with homozygous merle and stopped registering puppies produced from merle to merle matings in 2013. Merle to merle mating is currently only forbidden in three breeds. Recent research indicates that the majority of health issues occur in dogs carrying both piebald and merle genes. The suppression of pigment cells (
melanocytes Melanocytes are melanin-producing neural crest-derived cells located in the bottom layer (the stratum basale) of the skin's epidermis, the middle layer of the eye (the uvea), the inner ear, vaginal epithelium, meninges, bones, and heart. Mel ...
) in the iris and in the
stria vascularis The stria vascularis of the cochlear duct is a capillary loop in the upper portion of the spiral ligament (the outer wall of the cochlear duct). It produces endolymph for the scala media in the cochlea. Structure The stria vascularis is part of ...
of the
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus. A core component of the cochlea is the Organ of Corti, the sensory org ...
(inner ear) leads to blue eyes and deafness. An auditory-pigmentation disorder in humans, Waardenberg syndrome, reflects some of the problems associated with heterozygous and homozygous merle dogs and genetic research in dogs has been undertaken with the goal of better understanding the genetic basis of this human condition.


Ocular defects

Dogs who are homozygous for the merle pattern gene often have visual and auditory deficits. These dogs are sometimes referred to as 'double merle' and sometimes incorrectly referred to as 'lethal white'. Ocular defects include
microphthalmia Microphthalmia (Greek: grc, μικρός, mikros, small, label=none, grc, ὀφθαλμός, ophthalmos, eye, label=none, also referred as microphthalmos, is a developmental disorder of the eye in which one (unilateral microphthalmia) or both (b ...
, conditions causing increased ocular pressure, and colobomas, among others. Double merle dogs may be deaf or blind or both, and can carry ocular defects in blue or colored eyes. Currently no studies have been done to prove whether or not the merle gene affects the eyes, causing blindness.


Auditory defects

In one study of 38 dachshunds by a German researcher, partial hearing loss was found in 54.6% of double merles and 36.8% of single merles. 1 out of the 11 (9.1%) double merles was fully deaf while none of the single merles were. Another study done by
Texas A&M University Texas A&M University (Texas A&M, A&M, or TAMU) is a public university, public, Land-grant university, land-grant, research university in College Station, Texas. It was founded in 1876 and became the flagship institution of the Texas A&M Unive ...
found that of 22 double merles, 8 were completely deaf and two were deaf in one ear. Of 48 single merles, only one was deaf in one ear, and none were completely deaf. In another study of 70 dogs, 15 of them Catahoula Curs, 4 of the Catahoulas were deaf, while 86% of the double merles of other breeds were deaf. Deaf, blind, and deaf and blind dogs can have good lives when properly cared for. There are a variety of internet groups dedicated to supporting carers of such dogs. Deaf dogs can compete successfully in agility and there are many anecdotal reports of deaf/blind dogs earning their Canine Good Citizen certification, working as
therapy dogs A therapy dog is a dog that is trained to provide affection, comfort and support to people, often in settings such as hospitals, retirement homes, nursing homes, schools, libraries, hospices, or disaster areas. In contrast to assistance dogs, ...
, and competing in dog sports like tracking or Nosework.


See also

* List of dog coat patterns


References


External links


Merle dog coat color genetics



White Aussies Project


* : Basic, simple-language explanation of Australian Shepherd merle colorings


Border Wars - Double Merle Articles

Banning of Merle 'pattern' Chihuahua in Canada

Catahoula Coat Color Genetics - Introduction and History of Merle
{{DEFAULTSORT:Merle (Coat Colour In Dogs) Dog anatomy Animal hair Animal coat colors https://www.americanspanielclub.org/about-the-breed/merle-color-in-cocker-spaniels/