Ino Budgerigar Mutation
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The Ino budgerigar mutation is one of approximately 30 mutations affecting the colour of
budgerigar The budgerigar ( ; ''Melopsittacus undulatus''), also known as the common parakeet, shell parakeet or budgie ( ), is a small, long-tailed, seed-eating parrot native to Australia. Naturally the species is green and yellow with black, scallop ...
s. It is the underlying mutation of the Albino and Lutino varieties and, with
Cinnamon Cinnamon is a spice obtained from the inner bark of several tree species from the genus ''Cinnamomum''. Cinnamon is used mainly as an aromatic condiment and flavouring additive in a wide variety of cuisines, sweet and savoury dishes, biscuits, b ...
, a constituent mutation of the Lacewing variety.


Appearance

In the green series the Ino is known as the Lutino, with pure yellow
contour feathers The pennaceous feather is a type of feather present in most modern birds and in some other species of maniraptoriform dinosaurs. Description A pennaceous feather has a stalk or quill. Its basal part, called a ''calamus'', is embedded in the ski ...
, white or pale yellow
flight feather Flight feathers (''Pennae volatus'') are the long, stiff, asymmetrically shaped, but symmetrically paired pennaceous feathers on the wings or tail of a bird; those on the wings are called remiges (), singular remex (), while those on the tai ...
s and tail feathers and silvery-white cheek patches. In some lights the body can show a very pale green sheen.Watmough, W (1951) p48 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. The eyes of both the Lutino and Albino are red at all ages with white irides when adult, the beak is orange and the feet and legs are pink. The
cere The beak, bill, or Rostrum (anatomy), rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for pecking, wikt:grasp#Verb, grasping, and holding (in wikt:probe ...
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 LLC (stylized as PANTONE) is an American limited liability company headquartered in Carlstadt, New Jersey, and best known for its Pantone Matching System (PMS), a proprietary color order system used in a variety of industries, notably gr ...
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 Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
centre line are late to develop. The Ino gene masks the effect of virtually all other mutations, including
Opaline The opalines are a small group of peculiar heterokonts, currently assigned to the family Opalinidae, in the order Slopalinida. Their name is derived from the opalescent appearance of these microscopic organisms when illuminated with full sunlig ...
,
Dark Darkness is the condition resulting from a lack of illumination, or an absence of visible light. Human vision is unable to distinguish colors in conditions of very low luminance because the hue-sensitive photoreceptor cells on the retina are ...
, Dominant Grey, Dilute, and Clearwing. These genes, when present in an Ino in either
heterozygous 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. Mos ...
or
homozygous 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. Mos ...
form, cause no change in the appearance of the Ino. But the Ino gene does not entirely mask
Cinnamon Cinnamon is a spice obtained from the inner bark of several tree species from the genus ''Cinnamomum''. Cinnamon is used mainly as an aromatic condiment and flavouring additive in a wide variety of cuisines, sweet and savoury dishes, biscuits, b ...
. 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 is a spice obtained from the inner bark of several tree species from the genus ''Cinnamomum''. Cinnamon is used mainly as an aromatic condiment and flavouring additive in a wide variety of cuisines, sweet and savoury dishes, biscuits, b ...
Dilute German Fallows, 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. File:Melopsittacus undulatus -albino in cage-3.jpg, Albino hen File:Melopsittacus undulatus - albino and lutino.jpg, Albino hen and Lutino cock File:White domesticated budgerigar.jpg, Albino hen


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 ull citation? 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 is one of the three primary colours in the RYB color model, RYB colour model (traditional colour theory), as well as in the RGB color model, RGB (additive) colour model. It lies between Violet (color), violet and cyan on the optical spe ...
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 the Netherlands around 1885, while in England Mr C P Arthur of
Melksham Melksham () is a town and civil parish on the Bristol Avon, River Avon in Wiltshire, England, about northeast of Trowbridge and south of Chippenham. The parish population was 18,113 at the 2021 census. History Early history Excavations in ...
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, near
Rugeley Rugeley ( ) is a market town and civil parish in the Cannock Chase District, in Staffordshire, England. It lies on the north-eastern edge of Cannock Chase next to the River Trent; it is north of Lichfield, southeast of Stafford, northeast of ...
in Staffordshire, and Mrs Huntington of
Warwick Warwick ( ) is a market town, civil parish and the county town of Warwickshire in the Warwick District in England, adjacent to the River Avon, Warwickshire, River Avon. It is south of Coventry, and south-east of Birmingham. It is adjoined wit ...
. In August 1932 Mr F J Mullis of
Horsham Horsham () is a market town on the upper reaches of the River Arun on the fringe of the Weald in West Sussex, England. The town is south south-west of London, north-west of Brighton and north-east of the county town of Chichester. Nearby to ...
, Sussex, bred an Albino hen. Watmough (1951) p 47 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 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 Hemmingen () is a town in the district of Hanover, in Lower Saxony, Germany. It is situated approximately 6 km south of Hanover. Until December 2004, Hemmingen belonged to the Regierungsbezirk Hannover, which was dissolved in January 2005. ...
über
Hanover Hanover ( ; ; ) is the capital and largest city of the States of Germany, German state of Lower Saxony. Its population of 535,932 (2021) makes it the List of cities in Germany by population, 13th-largest city in Germany as well as the fourth-l ...
, 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 Sex linkage describes the sex-specific patterns of inheritance and expression when a gene is present on a sex chromosome (allosome) rather than a non-sex chromosome (autosome). Genes situated on the X-chromosome are thus termed X-linked, and a ...
recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and ...
at the ino locus on the
Z chromosome The ZW sex-determination system is a chromosomal system that determines the sex of offspring in birds, some fish and crustaceans such as the giant river prawn, some insects (including butterflies and moths), the schistosome family of flatworms ...
. 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 (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes. There are ...
pigment which is normally present in all feather
barb Barb or the BARBs or ''variation'' may refer to: People * Barb (name), a list of people and fictional characters with the given name or surname * Barb, a term used by fans of Nicki Minaj to refer to themselves * The Barbs, a band Places * Barb, ...
s 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 is a phenomenon in genetics in which the effect of a gene mutation is dependent on the presence or absence of mutations in one or more other genes, respectively termed modifier genes. In other words, the effect of the mutation is depe ...
, and Ino is phenotypically epistatic over many other mutations, including
Dark Darkness is the condition resulting from a lack of illumination, or an absence of visible light. Human vision is unable to distinguish colors in conditions of very low luminance because the hue-sensitive photoreceptor cells on the retina are ...
,
Grey Grey (more frequent in British English) or gray (more frequent in American English) is an intermediate color between black and white. It is a neutral or achromatic color, meaning that it has no chroma. It is the color of a cloud-covered s ...
,
Opaline The opalines are a small group of peculiar heterokonts, currently assigned to the family Opalinidae, in the order Slopalinida. Their name is derived from the opalescent appearance of these microscopic organisms when illuminated with full sunlig ...
, and the Dilute series. It is not epistatic over the
Blue Blue is one of the three primary colours in the RYB color model, RYB colour model (traditional colour theory), as well as in the RGB color model, RGB (additive) colour model. It lies between Violet (color), violet and cyan on the optical spe ...
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. In birds, the cock has two Z chromosomes and the hen has one Z and one W chromosome. So in hens whichever allele is present on the single Z chromosome is fully expressed in the
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
. 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 Z chromosomes (
homozygous 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. Mos ...
) to be expressed in the phenotype. Cocks which are
heterozygous 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. Mos ...
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.Taylor, T G; Warner, C (1986) p 45-46 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 mutation. For details of the genetic relationship, see the
Genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
section in the Sex-linked Clearbody article. The Ino gene is linked to other genes located on the Z chromosome, i.e. to the genes of other sex-linked mutations. In addition to the
Cinnamon Cinnamon is a spice obtained from the inner bark of several tree species from the genus ''Cinnamomum''. Cinnamon is used mainly as an aromatic condiment and flavouring additive in a wide variety of cuisines, sweet and savoury dishes, biscuits, b ...
mutation, these mutations include
Opaline The opalines are a small group of peculiar heterokonts, currently assigned to the family Opalinidae, in the order Slopalinida. Their name is derived from the opalescent appearance of these microscopic organisms when illuminated with full sunlig ...
and
Slate Slate is a fine-grained, foliated, homogeneous, metamorphic rock derived from an original shale-type sedimentary rock composed of clay or volcanic ash through low-grade, regional metamorphism. It is the finest-grained foliated metamorphic ro ...
. The cross-over or recombination 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 Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
in
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. When combined with the Yellowface II and Clearwing mutations the Rainbo ...
), 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 A chromatid (Greek ''khrōmat-'' 'color' + ''-id'') is one half of a duplicated chromosome. Before replication, one chromosome is composed of one DNA molecule. In replication, the DNA molecule is copied, and the two molecules are known as chrom ...
, symbolised as ''cin+''-''ino+''/''cin-ino''. Geneticists call this '
coupling A coupling is a device used to connect two shafts together at their ends for the purpose of transmitting power. The primary purpose of couplings is to join two pieces of rotating equipment while permitting some degree of misalignment or end mo ...
' 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' 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-Ino (Lacewing) and 2% Normal. Hens cannot be split for any sex-linked gene, so only cocks exist in Type I and Type II form.


Notes


References

* *


External links


World Budgerigar Organisation (WBO)


{{DEFAULTSORT:Ino Budgerigar Mutation Budgerigar colour mutations Albino animals