The discovery of sex-linkage in pigeons: learning at the example of butterflies

In 1908, Doncaster first experimentally demonstrated the concept of sex-linked inheritance in animals, namely on two varieties of gooseberry moth (Abraxas Lacticolor and Abraxas Grossulariata, 2016 Butterfly of the Year). This moth has the WZ / ZZ sex chromosome system, so that from the combination of a male lightened 'Lacticolor' with ZZ chromosome features and a female darker 'Grossulariata' with WZ features, and thus a shorter W chromosome, only female Lacticolor and male Grossulariata are raised.

Fig. 1: Crossing of an Abraxas Lacticolor and an Abraxas Grossulariata at Doncaster 1908, quoted as an excerpt from Crew (1927).

Pigeon fanciers will hardly be interested in the abraxas. However, knowledge of this inheritance mechanism may have contributed to the fact that shortly afterwards the resolution for some previously mysterious observations in domestic pigeons was found. So why silver chicks occasionally are raised out of blue pairs of pigeons and that these are exclusively females. Darwin had already dealt with this in a footnote in Chapter V 'Pigeons' in 1868 without finding an answer. Bonhote and Smalley showed in 1911 that it is one of the effects of this inheritance observed in the moth. Cole 1912 broadened the view almost simultaneously by the ratio of black to dun colored and from red to yellow pigeons. An observation of far-reaching importance? Not just by using an inheritance mechanism found in one species to explain appearances in another. In the case of pigeons, the scheme proved to be useful for explaining breeding results for hereditary factors such as ash red, brown, stipper, rubella, reduced, lethal web-foot, etc. For Reduced, Fig. 2 shows, in parallel to Fig. 1, an example of the effect of gender-specificity in mating a pigeon with reduced inheritance to a pigeon who does not carry the gene.

Fig. 2: Homozygous reduced cock x black hen and F1, reduced-colored hens and black cocks. Source: Sell, Genetik der Taubenfärbungen, Achim 2015.

You can quickly become familiar with the 'mechanics' of the transmission of sex-linked genes and the results that are important for breeding. With a little practice and mental agility, a breeder can easily transfer the statements from Fig. 1 and Fig. 2 to other combinations: a blue cock will breed blue hens with an ash red hen and heterozygous (for color) ash red cocks. In the diagram we simple replace Reduced by blue and black by ash red. From a dilute blue (silver) cock and a blue hen one obtains hemizygous dilute (silver) hens. The cocks are blue, however, heterozygous for the 'dilution factor' that in homozygous cocks and hemizygous hens is responsible for dilution of the color to silver. A yellow cock and a red hen produce yellow hens and heterozygous red cocks. Combinations of color, dilution, and other gender-related factors can be built in with a little practice. Once you understand the schema, you can apply it to the reverse mating, to the inheritance of heterozygous cocks and to the transmission of genes across generations. Incidentally, the fact that Reduced is inherited in a sex-linked manner was discovered with Carl Graefe by a genetically experienced pigeon fancier, Frosty by Andreas Leiß, who in addition to pigeon genetics was particularly fond of city pigeons. http://www.taubensell.de/reduced_ein_erb factor_geht_um_die_Welt.htm

In the case of Stipper, Faded, Frosty and other factors, there are also some special features due to a color gender dimorphism. Homozygous cocks with these traits differ from hemizygous hens and allow sexing by color. An inheritance scheme, many applications and thus a potentially important element for efficient breeding planning. Because it's not just about explanation. You can use the knowledge for a systematic breeding, which can be shown most easily with the help of the Punnet square as a didactic aid (Sell 2012, 2015, 2019). The young cock on the book cover in Fig. 3 and the couple owes their appearance to the interaction of two sex-related recessive factors, it is pure Rubella and also pure Frosty. If bred true a sex-linked strain similar to Faded and Frosty, but more pronounced. Thus an introduction in another exciting level of observation.


Fig. 3: Homozygous cock for both sex-linked genes Frosty and Rubella at the cover of the book ‘Taubenzucht’, and a homozygous frosty and homozygous rubella cock with a hemizygous frosty and hemizygous rubella hen at the right www.taubensell.de


Bonhote, J.L., und F.W. Smalley (1911), On the Colour and Colour Pattern inheritance in pigeons, Zoölogical Societey of London, Proceedings, pp. 601-619 (quoted from Levi 1969)

Cole, L.J. (1912), A case of sex-linked inheritance in the domestic pigeon. Science 36.

Crew, F.A.E. (1927), The Genetics of Sexuality in Animals, Cambridge at the University Press.

Darwin, Charles (1868), Animals and Plants Under Domestication, Chapter V Pigeons, London.

Doncaster, L. (1908), On sex inheritance in the moth, Abraxas grossulariata. 4th Rep. Evol. Comm., Roy. Soc. London (quoted from Crew 1927).

Levi, W.M. (1969), The Pigeon, first edition 1941, Sumter.

Sell, Axel (2012), Pigeon Genetics. Applied Genetics in the Domestic Pigeon, Achim.

Sell, Axel (2015), Genetik der Taubenfärbungen, Achim.

Sell, Axel (2019), Taubenzucht, Möglichkeiten und Grenzen züchterischer Gestaltung, Achim.