Hen feathering in cocks
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Hen feathering in cocks is the occurrence of a genetically conditioned character in domestic fowl (''
Gallus gallus domesticus The chicken (''Gallus gallus domesticus'') is a domesticated junglefowl species, with attributes of wild species such as the grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster or cock is a term for an adult ...
''). Males with this condition develop a female-type plumage, although otherwise look and respond as virile males. Hen-feathering in cocks is one of the typical characteristics of the
Sebright Bantam The Sebright (IPA: ) is a British breed of bantam chicken. It is a true bantam – a miniature bird with no corresponding large version – and is one of the oldest recorded British bantam breeds. It is named after Sir John Saunders Sebrig ...
, a breed established circa 1810, in accordance with the intentions of its creator, Sir John Saunders Sebright.
Sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
in plumage is very common in birds, particularly within Phasianidae where males are bigger and have brighter and more colorful plumage than females among other morphological differences.


Cocks with hen-feathering

Males of most chicken breeds distinguish from their females in having longer, sharp and more scalloped feathers in neck, hackle, saddle and wing bows. But in some breeds, like the fancy breeds
Sebright The Sebright ( IPA: ) is a British breed of bantam chicken. It is a true bantam – a miniature bird with no corresponding large version – and is one of the oldest recorded British bantam breeds. It is named after Sir John Saunders Sebri ...
and Campine and some game breeds like Pettai Madhirione can see males that have a plumage completely similar in all aspects to that of females. This unusual type of feathering called so much the interest of biologists that was studied from different points of view, and as a consequence, the inheritance of this condition has been well understood, while the biochemical basis determining this condition is still under study.


Mode of inheritance

It had been early established that hen-feathering is a trait controlled by a simple autosomic dominant gene, whose expression is limited to the male sex. Somes RG Jr, George FW, Baron J, Noble JF, Wilson JD. Inheritance of the henny-feathering trait of the Sebright bantam chicken. J. Hered. 1984 Mar-Apr;75(2):99-102. The genetic symbol proposed by F. B. Hutt in 1958 to designate this autosomic dominant gene was ''Hf'' (after "hen feathering")Hutt, F.B. Genética Avícola. Salvat Editores, S.A. 1ra.ed. España, 1958 which was accepted by other geneticists. Both homozygous ''Hf/Hf'' and heterozygous ''Hf/hf'' males are of the hen-feathering type, while ''hf/hf'' are of the normal feathering type. Meanwhile, female carriers of ''Hf'' gene can not be identified unless they are submitted to progeny tests. Attempts to demonstrate genetic linkage to several known loci have been unsuccessful.


Expressiveness of hen-feathering in cocks

Hen-feathering in heterozygous ''Hf/hf'' males is sometimes difficult to identify because some of those males may show only a few female feathers in their first adult plumage. But those males reach a complete female plumage after the first moult when they acquire the adult plumage of the second year.Punnet, R. C. and Bailey, P. G. 1921 Genetic studies in poultry. III. Hen-feathered cocks. Journal of Genetics 11,37-57 This happens because hen-feathering requires a masculine hormone and in some cases first adult plumage is completed before testicles work normally. Sometimes, well characterized old males of the hen-feathering type can turn, with age, into normal male plumage due to a subnormal testicular activity. These variations in the expression of the character, even within the same individual, defies common sense and may confuse non-advised breeders who might expect a better mating behaviour in normal male-feathering cocks than in those of the hen-feathering type. Some
Sebright The Sebright ( IPA: ) is a British breed of bantam chicken. It is a true bantam – a miniature bird with no corresponding large version – and is one of the oldest recorded British bantam breeds. It is named after Sir John Saunders Sebri ...
breeders consider hen-feathering to have adverse effect on the fertility of the male and may use cocks that do not carry the trait for breeding purposes, despite their automatic disqualification in shows.


Hormonal dependence of hen-feathering

Hen-feathering in cocks does not manifest itself unless the feathers are developed under the influx of sexual hormones. The effect on feathering is just the same both whether the hormone comes from the own testicles of the male or from an injection of testosterone. Chickens of both sexes develop male-type plumage after gonadectomization. Strictly speaking, they develop a neutral plumage with long acute feathers, like those of male plumage, because ovarian hormone is also necessary to develop female plumage. It is well known that some of the main
estrogens Estrogen or oestrogen is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three major endogenous estrogens that have estrogenic hormonal acti ...
derive from
androgen An androgen (from Greek ''andr-'', the stem of the word meaning "man") is any natural or synthetic steroid hormone that regulates the development and maintenance of male characteristics in vertebrates by binding to androgen receptors. This in ...
s. Androstenedione transforms into
estrone Estrone (E1), also spelled oestrone, is a steroid, a weak estrogen, and a minor female sex hormone. It is one of three major endogenous estrogens, the others being estradiol and estriol. Estrone, as well as the other estrogens, are synthesized ...
, while
testosterone Testosterone is the primary sex hormone and anabolic steroid in males. In humans, testosterone plays a key role in the development of male reproductive tissues such as testes and prostate, as well as promoting secondary sexual characteristi ...
transforms into
estradiol Estradiol (E2), also spelled oestradiol, is an estrogen steroid hormone and the major female sex hormone. It is involved in the regulation of the estrous and menstrual female reproductive cycles. Estradiol is responsible for the development o ...
through the action of the enzyme
aromatase Aromatase (), also called estrogen synthetase or estrogen synthase, is an enzyme responsible for a key step in the biosynthesis of estrogens. It is CYP19A1, a member of the cytochrome P450 superfamily, which are monooxygenases that catalyze many ...
. So this enzyme plays an important role in the biosynthesis of
estrogens Estrogen or oestrogen is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three major endogenous estrogens that have estrogenic hormonal acti ...
.


Enhanced aromatase activity in the skin

Skin of adult chickens carrying ''Hf'' has increased levels of estrogens. When skin grafts taken from females carrying ''Hf'' gene, are put on normal males, the skin of the grafts develop female (shorter and rounded tip) feathers. If the hosts of the grafts are subsequently castrated, all the feathers, including that of the skin graft turned into the male-type. In female chickens of all breeds development of female feathering pattern is mediated by estrogens. In normal ''hf/hf'' individuals rates of estrogen formation are undetectable or low in all control tissues other than ovary. Conversely rate of estrogen formation is high in skin and skin appendages and detectable in many other tissues biopsied from mature ''Hf/Hf'' or ''Hf/hf'' birds. The henny-feathering trait in males is the result of increased conversion of androgen to estrogen in skin. Increased estrogen formation in the skin and other peripheral tissues of chickens with the henny-feathering trait is due to an enhanced activity of the
aromatase Aromatase (), also called estrogen synthetase or estrogen synthase, is an enzyme responsible for a key step in the biosynthesis of estrogens. It is CYP19A1, a member of the cytochrome P450 superfamily, which are monooxygenases that catalyze many ...
complex of enzymes responsible for estrogen synthesis. Estrogen formation is as much as several hundred-fold higher in fibroblasts cultured from skin of chickens carrying the henny feathering trait compared to that observed in fibroblasts from skin of control chickens. The average rate of
aromatase Aromatase (), also called estrogen synthetase or estrogen synthase, is an enzyme responsible for a key step in the biosynthesis of estrogens. It is CYP19A1, a member of the cytochrome P450 superfamily, which are monooxygenases that catalyze many ...
activity, measured using a sensitive isotopic assay, in the skin of heterozygous ''Hf/hf'' chicks is approximately half that of homozygous ''Hf/Hf'' henny chicks. But half maximal level of the enzyme is sufficient to allow full development of female feathering in affected male birds and this explains why hen-feathering in heterozygous ''Hf/hf'' males is sometimes difficult to identify in their first adult plumage: See Expressiveness.... In normal ''hf/hf'' birds the activity of the enzyme is limited in all tissues other than ovary.


Aromatase excess syndrome in humans

There is a curious resemblance of the increased
aromatase Aromatase (), also called estrogen synthetase or estrogen synthase, is an enzyme responsible for a key step in the biosynthesis of estrogens. It is CYP19A1, a member of the cytochrome P450 superfamily, which are monooxygenases that catalyze many ...
activity in chickens, to the
aromatase excess syndrome Aromatase excess syndrome (AES or AEXS) is a rare genetic and endocrine syndrome which is characterized by an overexpression of aromatase, the enzyme responsible for the biosynthesis of the estrogen sex hormones from the androgens, in turn resu ...
in humans, a rather rare syndrome that can lead to
gynecomastia Gynecomastia (also spelled gynaecomastia) is the abnormal non-cancerous enlargement of one or both breasts in males due to the growth of breast tissue as a result of a hormone imbalance between estrogens and androgens. Updated by Brent Wisse ( ...
in boys, and to
precocious puberty In medicine, precocious puberty is puberty occurring at an unusually early age. In most cases, the process is normal in every aspect except the unusually early age and simply represents a variation of normal development. In a minority of childr ...
and
gigantomastia Breast hypertrophy is a rare medical condition of the breast connective tissues in which the breasts become excessively large. The condition is often divided based on the severity into two types, macromastia and gigantomastia. Hypertrophy of the ...
in girls, and which is also inherited in an autosomal dominant fashion.Fukami, M. ''et al'' 2011 "Aromatase excess syndrome: identification of cryptic duplications and deletions leading to gain of function of CYP19A1 and assessment of phenotypic determinants". J. Clin. Endocrinol. Metab. 96 (6): E1035–43.


See also

*
Aromatase Aromatase (), also called estrogen synthetase or estrogen synthase, is an enzyme responsible for a key step in the biosynthesis of estrogens. It is CYP19A1, a member of the cytochrome P450 superfamily, which are monooxygenases that catalyze many ...
*
Aromatase excess syndrome Aromatase excess syndrome (AES or AEXS) is a rare genetic and endocrine syndrome which is characterized by an overexpression of aromatase, the enzyme responsible for the biosynthesis of the estrogen sex hormones from the androgens, in turn resu ...


References

{{Reflist Aviculture Chicken plumage patterns Feathers