Copyright  2004 by the Genetics Society of America

Perspectives
Anecdotal, Historical and Critical Commentaries on Genetics
Edited by James F. Crow and William F. Dove

An Alternative Possibility for Seed Coat Color Determinaton

in Mendel’s Experiment

James R. Myers1
Department of Horticulture, Oregon State University, Corvallis, Oregon 97331

I T is possible that in his test of the hypothesis of inde-

pendence among three genes influencing seed char-
acters of pea (cotyledon color and shape and flower or
seed color) Mendel relied on partial dominance of seed
color to classify heterozygotes as well as homozygotes
in his cross. It would make sense for him to do this if
possible, rather than growing progeny to classify flower
color. He indicated that hybrid seed coats “are often
more spotted and the spots sometimes coalesce into
small bluish-purple patches.” Correns also described
partial dominance of seed coat color, but it is not clear
to me whether he used the information in classifying
his populations. Apart from these descriptions, I do not
know of any subsequent researchers who used partial
dominance of seed coat color to classify heterozygotes.
While on average the testa of the heterozygote is more
intensely pigmented, variations among seeds from dif-
ferent plants as well as among seeds of the same plant
are observed. In some crosses, differences between the
pigmented homozygote and heterozygote may be suffi-
ciently discrete for accurate classification. In other
crosses, the lines between heterozygotes and homozy-
gotes may be blurred. If it were possible to recreate
the original crosses, one might be able to determine
whether Mendel would have had the opportunity to
classify his material on the basis of seed characteristics
alone.
Classification is complicated by additional seed coat
color genes of which neither Mendel nor Correns ap-
peared to be aware. Correns’s description of an “orange-
red seed coat, which turns brown on aging” may describe
the M gene. Mendel describes the seed coat as being
“grey, grey-brown, or leather-brown,” the latter of which
might be the M gene. This gene does not appear to be
involved in partial dominance. Rather, it is the intensity
of anthocyanin spotting that gives a clue as to the geno-
type. Anthocyanin spotting is controlled by two loci (F
and Fs), each independent of the other. We do not
know whether one or both were present in the parental
material of Mendel. Mendel does not report indepen-
dent segregation of anthocyanin spotting. If one of these
genes were segregating, it would complicate the classifi-
cation of partial dominance of seed color. Both Mendel
and Correns reported parents that did not show the
purple spotting, but when crossed, a new character (pur-
ple spotting) was observed in the F1 and subsequent
generations. This result could be explained if the par-
ents had the genotype aa FF ⫻ Aa ff [white flowers/
colorless seed coat ⫻ purple flowers/gray (unspotted)
seed coat] (A, purple flowers and gray seed coat; a,
white flowers and colorless seed coat; F, anthocyanin
spotting on seed coat; f, absence of anthocyanin spot-
ting; aa prevents expression of F). Anthocyanin spotting
would appear in the F1: Aa Ff.
But the following would be expected to segregate in
the F2: A-F- (purple flowers/spotted seed); A-ff (purple
flowers/gray but nonspotted seed); aaF- (white flowers/
colorless seed coat); aaff (white flowers/colorless seed
coat). Such segregation would produce an expectation
of 3:1 for the spotted to nonspotted seeds among the
purple flower class instead of the 2:1 ratio expected if
the spotting in the heterozygote could be unambigu-
ously identified. In this issue of Genetics, C. E. Novitski
has performed the customary ␹2 tests and has shown
that both alternatives are statistically in agreement with
Mendel’s data, although Mendel’s 2:1 ratio gives a some-
what better fit (Myers’ ␹2 ⫽ 29.2, P ⫽ 0.30; Mendel’s
␹2 ⫽ 15.3, P ⫽ 0.95).

1
Author e-mail: myersja@science.oregonstate.edu

Genetics 166: 1137 ( March 2004)