Genetics soon became an active field of research, and exceptions to Mendel’s theory appeared. Dominance was often incomplete, and instead of simply 2 alternate forms there might be 3 or more. But a fertilized egg should get only 2.

Triple allelomorphs were first reported in pigeons and doves by L. J. Cole and his students by 1930. Symbols were a bit of a problem, as well as sex-linkage. Well the third allelomorphs was given the same base letter as the others, and aspecial superscript (example B^A). Next came the pattern series in pigeons with at least 4 allelomorphs, and the almond color series.

About this time geneticists were adopting the term allele as an abbreviation of allelomorphs. Also it was found that using the normal wild type of the species as a standard of reference is very helpful in figuring out the “action” of the genes (phenogenetics). For convenience the wild-type genes (alleles) all got identified with a “+” symbol, usually as a superscript.

Several more multiple-alleles series have been found in pigeons, and more will undoubtedly be discovered. Genetics buffs should have an exciting experience working them out. One that is being played with no is the z (gazzi) series, which now includes the “pencilled” gene, according to some tests by Kerry Hendricks, Robert Mangile, Larry Chesling, and me. Originally the study of this gene by Mme. Francqueville concluded that it is a simple recessive and she gave it the symbol, pc. Later study by our editor indicated partial dominance: he said (1993) “Sometimes a pseudo white-bar is produced by the heterozygous condition”, so he used the symbol Pc. That sort of whitening has been called “flash” by fanciers (see for example discussion in PGNV&C 72 by Lynn Kral and Tim Kvidera in relation to “tail-mark” (all-white except tail). They think tail-mark is a single-gene effect but did not investigate the possibility of allelism with gazzi or penciled, which I think is very probable. And in view of the generally intermediate phenotypes of heterozygotes between gazzi, recessive white, and penciled, I suspect that one more allele, almost like the wild-type allele, is lurking in various breeds, and may be the culprit giving the flash effect which Gibson got in crosses of penciled. I am on its trail, and hope somebody else will join in. Assuming that my hypothesis is right, we have at least 6 alleles in the z-series, and we can set up a chart of their various possible pair-combination as follows.

Intersections give: phenotypes of heterozygotes. Question mark means needs checking.