Ino budgerigar mutation
Encyclopedia
The Ino budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigar
Budgerigar
The Budgerigar , also known as Common Pet Parakeet or Shell Parakeet informally nicknamed the budgie, is a small, long-tailed, seed-eating parrot, and the only species in the Australian genus Melopsittacus...

s. It is the underlying mutation of the Albino and Lutino varieties and, with Cinnamon
Cinnamon budgerigar mutation
The Cinnamon budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is the underlying mutation of the Cinnamon variety and, with Ino, a constituent mutation of the Lacewing variety.- Appearance :...

, a constituent mutation of the Lacewing variety.

Appearance

In the green series the Ino is known as the Lutino, with pure yellow contour feathers, white or pale yellow flight feather
Flight feather
Flight feathers are the long, stiff, asymmetrically shaped, but symmetrically paired feathers on the wings or tail of a bird; those on the wings are called remiges while those on the tail are called rectrices . Their primary function is to aid in the generation of both thrust and lift, thereby...

s and tail feathers and silvery-white cheek patches. In some lights the body can show a very pale green sheen.

In the blue series the Ino is known as the Albino, and is pure white throughout. The cheek patches are almost the same colour as the body, but slightly more silvery. In some lights the body can show a very pale blue sheen.
Variety Pantone Code
Lutino 102


The eyes of both the Lutino and Albino are red at all ages with white irides
Iris (anatomy)
The iris is a thin, circular structure in the eye, responsible for controlling the diameter and size of the pupils and thus the amount of light reaching the retina. "Eye color" is the color of the iris, which can be green, blue, or brown. In some cases it can be hazel , grey, violet, or even pink...

 when adult, the beak is orange and the feet and legs are pink. The cere
Cère
The Cère is a long river in south-western France, left tributary of the Dordogne River. Its source is in the south-western Massif Central, near the mountain Plomb du Cantal...

 of an adult Ino cock is greyish-purple rather than blue.

The World Budgerigar Organisation has established precise standards for some budgerigar body colours using the Pantone
Pantone
Pantone Inc. is a corporation headquartered in Carlstadt, New Jersey, USA. The company is best known for its Pantone Matching System , a proprietary color space...

 Codes, as shown to the right for the Lutino.

The Ino mutation also induces changes in the nestling. The down is white rather than grey and appears only sparsely, never growing down the centre of the back. As the feathers appear, those down the spine and along the ventral centre line are late to develop.

The Ino gene masks the effect of virtually all other mutations, including Opaline
Opaline budgerigar mutation
The Opaline budgerigar mutation is one of approximately 30 mutations affecting the colour or appearance of budgerigars. It is the underlying mutation of the Opaline variety...

, Dark
Dark budgerigar mutation
The Dark budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is part of the genetic constitution of the following recognised varieties: Dark Green and Olive in the green series and Cobalt, Mauve and Violet in the blue series.- Appearance :Budgerigars...

, Dominant Grey
Dominant Grey budgerigar mutation
The Dominant Grey budgerigar mutation, often called the Australian Grey or simply Grey, is one of approximately 30 mutations affecting the colour of budgerigars. It is the basis of the Grey-Green and Grey standard varieties.- Appearance :...

, Dilute
Dilute budgerigar mutation
The Dilute budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is one of the constituent mutations of several recognised varieties: the Light, Dark, Olive, Grey and Suffused Yellows and the Grey and Suffused Whites....

, and Clearwing
Clearwing budgerigar mutation
The Clearwing budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is the underlying mutation of the Clearwing variety, often known as Yellowwings in the green series and Whitewings in the blue series. When combined with the Greywing mutation the...

. These genes, when present in an Ino in either heterozygous or homozygous form, cause no change in the appearance of the Ino. But the Ino gene does not entirely mask Cinnamon
Cinnamon budgerigar mutation
The Cinnamon budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is the underlying mutation of the Cinnamon variety and, with Ino, a constituent mutation of the Lacewing variety.- Appearance :...

.
A Cinnamon Ino, usually called a Lacewing, has pale brown or fawn spots, tail and wing markings. These markings are quite clear, but considerably fainter than the markings of a normal Cinnamon.

The Dark-eyed Clear has a similar body colour to the Ino, but has solid reddish-purple eyes without a white iris.

Cinnamon
Cinnamon budgerigar mutation
The Cinnamon budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is the underlying mutation of the Cinnamon variety and, with Ino, a constituent mutation of the Lacewing variety.- Appearance :...

 Dilute
Dilute budgerigar mutation
The Dilute budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is one of the constituent mutations of several recognised varieties: the Light, Dark, Olive, Grey and Suffused Yellows and the Grey and Suffused Whites....

 German Fallows
German Fallow budgerigar mutation
The German Fallow budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. At least three types of Fallow, the German, English, and Scottish, all named after their country of origin, have been established, although none of these types is common. They are...

, NSL Inos and Inos are all very similar and difficult to distinguish from each other, but the first two are so uncommon difficulties arise rarely in practice.

Historical notes

The first known reference to the Ino mutation in the budgerigar was a report by Mr L van der Snickt, a Belgian fancier, in the German avicultural paper Die Gefiederte Welt (The Feathered World) in 1879. He wrote that he had seen that year nine Lutinos, all hens. (In fact, he called them Albinos, since the name Lutino did not then exist, but from his description and the fact that the Blue
Blue budgerigar mutation
The Blue budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is part of the genetic constitution of the following recognised varieties: Skyblue, Cobalt, Mauve and Violet.- Appearance :...

 mutation was not established until the 1880s it is clear they were Lutinos.) One breeder of these birds was Mr Kessels, also of Belgium, who in 1881 bred 25 Lutinos, all hens.

A coloured picture of a Lutino appeared in the Brussels journal, Acclimatation Illustrée, in 1882, and it is thought they were being bred in Holland around 1885, while in England Mr C P Arthur of Melksham
Melksham
Melksham is a medium-sized English town, lying on the River Avon. It lies in the county of Wiltshire.It is situated southeast of the city of Bath, south of Chippenham, west of Devizes and north of Warminster on the A350 national route. The 2001 UK census cited Melksham as having 20,000...

 in Wiltshire bred what he believed was a pair of Inos around 1887.
After the 1880s no mention seems to have been made in the press of Inos until the 1930s, when interest in budgerigar mutations suddenly increased.

In 1930/31, Lutino hens were owned by both Capt H S Stokes of Longdon
Longdon, Staffordshire
Longdon is a village and civil parish in the District of Lichfield, Staffordshire, England, situated midway between the towns of Rugeley and Lichfield. The parish also includes the nearby villages of Upper Longdon, Longdon Green and Gentleshaw....

, near Rugeley
Rugeley
Rugeley is a historic market town in the county of Staffordshire, England. It lies on the northern edge of Cannock Chase, and is situated roughly midway between the towns of Stafford, Cannock, Lichfield and Uttoxeter...

 in Staffordshire, and Mrs Huntington of Warwick
Warwick
Warwick is the county town of Warwickshire, England. The town lies upon the River Avon, south of Coventry and just west of Leamington Spa and Whitnash with which it is conjoined. As of the 2001 United Kingdom census, it had a population of 23,350...

. In August 1932 Mr F J Mullis of Horsham
Horsham
Horsham is a market town with a population of 55,657 on the upper reaches of the River Arun in the centre of the Weald, West Sussex, in the historic County of Sussex, England. The town is south south-west of London, north-west of Brighton and north-east of the county town of Chichester...

, Sussex, bred an Albino hen.
None of these led to an established strain.

In September 1931, Mr E Böhm of Bawerk in Germany bred, as the last of nine young from a pair of Cobalt split Dilutes,

a snow-white red-eyed hen
- the first recorded Albino. Almost exactly a year later, on 12 September 1932, a second Albino hen was bred by Mr Fischer of Honow
Hönow
Hönow is a village in Brandenburg, Germany, near the border of Berlin. It belongs to the municipality of Hoppegarten, in the district of Märkisch-Oderland; and its population is of 8,300 inhabitants.-Geography:...

 in Germany from a pair of Skyblues. Both of these strains were established by the original breeders and also by others who acquired early stock from them, in particular by Kurt Kokemüller of Arnum über Hanover
Hanover
Hanover or Hannover, on the river Leine, is the capital of the federal state of Lower Saxony , Germany and was once by personal union the family seat of the Hanoverian Kings of Great Britain, under their title as the dukes of Brunswick-Lüneburg...

, and Mr Schrapel, also of Hanover, who performed together the first genetic investigations into the Ino mutation and published the first correct pairing expectations in the German publication Der Wellensittich (The Budgerigar) in November and December 1933.

A third appearance of the Ino mutation occurred in Germany around 1933, when Mr Kuhlewein bred a Lutino hen in an uncontrolled breeding flight.

This strain was also established.

Other Ino mutations also appeared in Europe in the early 1930s, and several British fanciers, including Walter Higham, Scott and Camplin, and Tod Boyd, had imported continental Lutinos by the mid-1930s.
Some of these turned out to be of the non-sex-linked type and the unwitting mixing of the two mutations led to considerable confusion. All British Inos seem to have descended from these imported continental Inos.

In 1976, Dr T Daniels began a controlled programme of pairings to produce a Cinnamon Ino by deliberately crossing Cinnamons to Inos, and to estimate the cross-over value between these two mutations. The first Cinnamon Ino was produced in late 1979 and was identical in appearance to a Lacewing.

Genetics

The Ino mutation is a sex-linked recessive
Recessive
In genetics, the term "recessive gene" refers to an allele that causes a phenotype that is only seen in a homozygous genotype and never in a heterozygous genotype. Every person has two copies of every gene on autosomal chromosomes, one from mother and one from father...

 at the ino locus on the X chromosome
X chromosome
The X chromosome is one of the two sex-determining chromosomes in many animal species, including mammals and is common in both males and females. It is a part of the XY sex-determination system and X0 sex-determination system...

. The wild-type genetic symbol is ino+ and the ino mutant allele has the symbol ino. Its effect is to inhibit the production of the melanin
Melanin
Melanin is a pigment that is ubiquitous in nature, being found in most organisms . In animals melanin pigments are derivatives of the amino acid tyrosine. The most common form of biological melanin is eumelanin, a brown-black polymer of dihydroxyindole carboxylic acids, and their reduced forms...

 pigment which is normally present in all feather barbs in either the medullary or cortical cells or both. The presence of black melanin pigment in the cortex of the barbs is necessary for the production of the black markings and in the medulla of barbs for the production of the blue colouration (which combines with the yellow pigment in birds of the green series to produce the green colouration), so this mutation removes all black and blue colourations resulting in a white bird in the blue series and a yellow bird in the green series.

Because the Ino mutation totally inhibits the production of normal melanin pigment it prevents the visible expression of all the other mutations which depend on the presence of melanin to show their effect. This is called epistasis
Epistasis
In genetics, epistasis is the phenomenon where the effects of one gene are modified by one or several other genes, which are sometimes called modifier genes. The gene whose phenotype is expressed is called epistatic, while the phenotype altered or suppressed is called hypostatic...

, and Ino is phenotypically epistatic over many other mutations, including Dark
Dark budgerigar mutation
The Dark budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is part of the genetic constitution of the following recognised varieties: Dark Green and Olive in the green series and Cobalt, Mauve and Violet in the blue series.- Appearance :Budgerigars...

, Grey
Dominant Grey budgerigar mutation
The Dominant Grey budgerigar mutation, often called the Australian Grey or simply Grey, is one of approximately 30 mutations affecting the colour of budgerigars. It is the basis of the Grey-Green and Grey standard varieties.- Appearance :...

, Opaline
Opaline budgerigar mutation
The Opaline budgerigar mutation is one of approximately 30 mutations affecting the colour or appearance of budgerigars. It is the underlying mutation of the Opaline variety...

, and the Dilute
Dilute budgerigar mutation
The Dilute budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is one of the constituent mutations of several recognised varieties: the Light, Dark, Olive, Grey and Suffused Yellows and the Grey and Suffused Whites....

 series. It is not epistatic over the Blue
Blue budgerigar mutation
The Blue budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is part of the genetic constitution of the following recognised varieties: Skyblue, Cobalt, Mauve and Violet.- Appearance :...

 mutation, so there are two forms of the albino budgerigar, one in the green series called the Lutino and one in the blue series called the Albino. Both these varieties may be masking many other hypostatic mutations, so the genotype of an Albino or Lutino with respect to these mutations cannot be determined visually. Nor is the Ino mutation epistatic over the Cinnamon mutation — see below.
Sex Genotype Phenotype
Cocks ino+/ino+ Normal
ino+/ino Normal (/ino)
ino/ino Ino
Hens ino+/Y Normal
ino/Y Ino


In birds, the cock has two X chromosomes and the hen has one X and one Y chromosome. So in hens whichever allele is present on the single X chromosome is fully expressed in the phenotype
Phenotype
A phenotype is an organism's observable characteristics or traits: such as its morphology, development, biochemical or physiological properties, behavior, and products of behavior...

. Hens cannot be split for Ino (or any other sex-linked mutation). In cocks, because Ino is recessive, the Ino allele must be present on both X chromosomes (homozygous) to be expressed in the phenotype. Cocks which are heterozygous for Ino are identical to the corresponding Normal. Such birds are said to be split for Ino, usually written '/ino'.

The table on the right shows the appearance of all possible genetic combinations involving just the Ino mutation.

The Ino mutation does not mask the Cinnamon mutation, these two genes being neither fully epistatic nor hypostatic to each other. When combined in doubly homozygous form (cin-ino/cin-ino in cocks or cin-ino/Y in hens) the Lacewing phenotype is produced. The Cinnamon markings are clearly visible, although considerably fainter than in a normal Cinnamon. For many years the Lacewing was thought by many to be a separate mutation but it was demonstrated in 1979 that it was simply a Cinnamon Ino when a Lacewing was deliberately produced by combining separate Cinnamon and Ino genes. Once brought together, these two genes are almost always inherited together due to the close linkage between them, giving the impression of being a single gene.

The Ino mutation is a member of a series of multiple alleles at the ino locus. Only one other member is known -- the Sex-linked Clearbody
Sex-linked Clearbody budgerigar mutation
The Sex-linked Clearbody budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is the underlying mutation of the Texas Clearbody variety....

 mutation. For details of the genetic relationship, see the Genetics section in the Sex-linked Clearbody
Sex-linked Clearbody budgerigar mutation
The Sex-linked Clearbody budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is the underlying mutation of the Texas Clearbody variety....

 article.

The Ino gene is linked
Genetic linkage
Genetic linkage is the tendency of certain loci or alleles to be inherited together. Genetic loci that are physically close to one another on the same chromosome tend to stay together during meiosis, and are thus genetically linked.-Background:...

 to other genes located on the X chromosome, i.e. to the genes of other sex-linked mutations. In addition to the Cinnamon
Cinnamon budgerigar mutation
The Cinnamon budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is the underlying mutation of the Cinnamon variety and, with Ino, a constituent mutation of the Lacewing variety.- Appearance :...

 mutation, these mutations include Opaline
Opaline budgerigar mutation
The Opaline budgerigar mutation is one of approximately 30 mutations affecting the colour or appearance of budgerigars. It is the underlying mutation of the Opaline variety...

 and Slate
Slate budgerigar mutation
The Slate budgerigar mutation is one of approximately 30 mutations affecting the colour of budgerigars. It is the underlying mutation of the Slate variety.- Appearance :...

. The cross-over
Genetic linkage
Genetic linkage is the tendency of certain loci or alleles to be inherited together. Genetic loci that are physically close to one another on the same chromosome tend to stay together during meiosis, and are thus genetically linked.-Background:...

 or recombination
Genetic recombination
Genetic recombination is a process by which a molecule of nucleic acid is broken and then joined to a different one. Recombination can occur between similar molecules of DNA, as in homologous recombination, or dissimilar molecules, as in non-homologous end joining. Recombination is a common method...

 values between Ino and these linked genes has not been measured accurately, but some approximate measurements of the cross-over values have been made:
  • Cinnamon-Ino COV: Breeding results collected by C Warner and T Daniels found just 1 crossover in 36 between Cinnamon and Ino. Other measurements found at least 1 cross-over in 18, so combining these the best estimate of the recombination value is ≥4±3%.

  • Opaline-Ino COV: Only one direct measurement of the Opaline-Ino linkage has been reported. This found 3 cross-overs in 10, giving a recombination value of 30±17%. But since the ino locus is very close to the cinnamon locus the COV for Opaline-Ino must be very similar to that for Opaline-Cinnamon. The Opaline-Cinnamon linkage has been measured to be approximately 36±6% (see Genetics in Opaline budgerigar mutation
    Opaline budgerigar mutation
    The Opaline budgerigar mutation is one of approximately 30 mutations affecting the colour or appearance of budgerigars. It is the underlying mutation of the Opaline variety...

    ), so these two results are in agreement within the limited statistics.


Cocks split for both Cinnamon and Ino have one Cinnamon allele and one Ino allele together with one each of the corresponding wild-type alleles. The linkage between the Cinnamon and Ino genes gives rise to two types of split cinnamon-ino cocks, both visually identical.
  • Type I split cinnamon-ino cocks are bred by mating Cinnamon-Inos (Lacewings) to Normals and have the two mutant alleles on the same chromatid
    Chromatid
    A chromatid is one of the two identical copies of DNA making up a duplicated chromosome, which are joined at their centromeres, for the process of cell division . They are called sister chromatids so long as they are joined by the centromeres...

    , symbolised as cin+-ino+/cin-ino. Geneticists call this 'coupling
    Genetic linkage
    Genetic linkage is the tendency of certain loci or alleles to be inherited together. Genetic loci that are physically close to one another on the same chromosome tend to stay together during meiosis, and are thus genetically linked.-Background:...

    ' rather than 'Type I'. Because of the linkage, the Cinnamon and Ino alleles from Type I cocks tend to be inherited together in their progeny. When mated to Normal hens, Type I cocks produce predominantly Cinnamon-Ino (Lacewing) and Normal hens, with Cinnamon and Ino hens resulting extremely rarely from a cross-over. Roughly 48% of the hens will be Cinnamon-Ino (Lacewing), 48% Normal, 2% Cinnamon and 2% Ino.

  • Type II split cinnamon-ino cocks are bred by mating Cinnamons to Inos and have the Cinnamon and Ino mutant alleles on opposite chromatids, symbolised as cin+-ino/cin-ino+. Geneticists call this 'repulsion
    Genetic linkage
    Genetic linkage is the tendency of certain loci or alleles to be inherited together. Genetic loci that are physically close to one another on the same chromosome tend to stay together during meiosis, and are thus genetically linked.-Background:...

    ' rather than 'Type II'. Because of the separation, the Cinnamon and Ino alleles from Type II birds tend to be inherited separately in their progeny. When mated to Normal hens, Type II cocks produce predominantly Cinnamon and Ino hens, with Cinnamon-Ino (Lacewing) and Normal hens resulting extremely rarely from cross-overs. Roughly 48% of the hens will be Cinnamon, 48% Ino, 2% Cinnamon-Opaline (Lacewing) and 2% Normal.


Hens cannot be split for any sex-linked gene, so only cocks exist in Type I and Type II form.

External links

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