Multiple alleles, incomplete dominance,

and codominance
In the real world, genes often come in many versions (alleles). Alleles aren't always fully dominant or
recessive to one another, but may instead display codominance or incomplete dominance.
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Introduction
Gregor Mendel knew how to keep things simple. In Mendel's work on pea
plants, each gene came in just two different versions, or alleles, and these
alleles had a nice, clear-cut dominance relationship (with the dominant allele
fully overriding the recessive allele to determine the plant's appearance).

Today, we know that not all alleles behave quite as straightforwardly as in

Mendel’s experiments. For example, in real life:

 Allele pairs may have a variety of dominance relationships (that is, one allele
of the pair may not completely “hide” the other in the heterozygote).

 There are often many different alleles of a gene in a population.

In these cases, an organism's genotype, or set of alleles, still determines

its phenotype, or observable features. However, a variety of alleles may
interact with one another in different ways to specify phenotype.

As a side note, we're probably lucky that Mendel's pea genes didn't show
these complexities. If they had, it’s possible that Mendel would not have
understood his results, and wouldn't have figured out the core principles of
inheritance—which are key in helping us understand the special cases!

Incomplete dominance
Mendel’s results were groundbreaking partly because they contradicted the
(then-popular) idea that parents' traits were permanently blended in their
offspring. In some cases, however, the phenotype of a heterozygous
organism can actually be a blend between the phenotypes of its homozygous
parents.

For example, in the snapdragon, Antirrhinum majus, a cross between a

homozygous white-flowered plant (C^WC^WCWCWC, start superscript, W,
end superscript, C, start superscript, W, end superscript) and a homozygous
red-flowered plant (C^RC^RCRCRC, start superscript, R, end superscript, C,
start superscript, R, end superscript) will produce offspring with pink flowers
(C^RC^WCRCWC, start superscript, R, end superscript, C, start superscript,
W, end superscript). This type of relationship between alleles, with a
heterozygote phenotype intermediate between the two homozygote
phenotypes, is called incomplete dominance.
In Japanese four o'clock plants red (R) color is incompletely dominant over white (r)
flowers, and the heterozygous condition (Rr) results in plants with pink flowers. For
each of the following construct a punnett square and give phenotypic and genotype
ratios of the offspring.
a) a red plant and a white plant }
b) a red plant and a pink plant }
c) a white plant and a pink plant
d) two pink plants

2. In some cats the gene for tail length shows incomplete dominance. Cats with long
tails and cats with no tails are homozygous for their respective alleles. Cats with one
long tail allele and one no tail allele have short tails. For each of the following
construct a punnett square and give phenotypic and genotype ratios of the offspring.
a) a long tail cat and a cat with no tail
b) a long tail cat and a short tail cat
c) a short tail cat and a cat with no tail
d) two short tail cats.

Incomplete Dominance Solutions

1. red (R), white (r), pink (Rr)
a) a red plant and a white plant
RR X rr, gametes for RR (R), gametes for rr (r), resulting phenotype ratio: 100% pink, resulting
genotype ratio: 100% Rr
b) a red plant and a pink plant
RR X Rr, gametes for RR (R), gametes for Rr (R & r), resulting phenotype ratio: 50% red / 50%
pink, resulting genotype ratio: 50% RR / 50% Rr
c) a white plant and a pink plant
rr X Rr, gametes for rr (r), gametes for Rr (R & r), resulting phenotype ratio: 50% pink / 50%
white, resulting genotype ratio: 50% Rr / 50% rr (Thanks Sarah L.)
d) two pink plants
Rr X Rr, gametes for Rr (R & r), gametes for Rr (R & r), resulting phenotype ratio: 25% red /
50% pink / 25% white, resulting genotype ratio: 25% RR / 50% Rr / 25% rr

2. long tails (L), no tails (ll), short tails (Ll).

a) a long tail cat and a cat with no tail
LL X ll, gametes for LL (L), gametes for ll (l), resulting phenotype ratio: 100% short tails,
resulting genotype ratio: 100% Ll
b) a long tail cat and a short tail cat
LL X Ll, gametes for LL (L), gametes for Ll (L & l), resulting phenotype ratio: 50% long tails /
50% short tails, resulting genotype ratio: 50% LL / 50% Ll
c) a short tail cat and a cat with no tail
Ll X ll, gametes for Ll (L & l), gametes for ll (l), resulting phenotype ratio: 50% short tails / 50%
no tails, resulting genotype ratio: 50% Ll / 50% ll
d) two short tail cats
Ll X Ll, gametes for Ll (L& l), gametes for Ll (L & l), resulting phenotype ratio: 25% Long
Tailed, 50% short tailed, and 25% no tailed, resulting genotype ratio: 25% LL / 50% Ll / 25% ll

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 Diagram of a cross between C^WC^WCWCWC, start superscript, W, end
superscript, C, start superscript, W, end superscript(red) and C^RC^RCRCRC,
 start superscript, R, end superscript, C, start superscript, R, end
superscript(white) snapdragon plants. The F1 plants are pink and of
genotype C^RC^WCRCWC, start superscript, R, end superscript, C, start
superscript, W, end superscript.

We can still use Mendel's model to predict the results of crosses for alleles
that show incomplete dominance. For example, self-fertilization of a pink
plant would produce a genotype ratio of 111 C^RC^RCRCRC, start
superscript, R, end superscript, C, start superscript, R, end
superscript ::colon 222 C^RC^WCRCWC, start superscript, R, end superscript,
C, start superscript, W, end superscript ::colon 111 C^WC^WCWCWC, start
superscript, W, end superscript, C, start superscript, W, end superscript and a
phenotype ratio of 1:2:11:2:11, colon, 2, colon, 1 red:pink:white. Alleles are
still inherited according to Mendel's basic rules, even when they show
incomplete dominance.
 Self-fertilization of pink C^RC^WCRCWC, start superscript, R, end
superscript, C, start superscript, W, end superscriptplants produce red, pink,
and white offspring in a ratio of 1:2:1.

Codominance
Closely related to incomplete dominance is codominance, in which both
alleles are simultaneously expressed in the heterozygote.

We can see an example of codominance in the MN blood groups of humans

(less famous than the ABO blood groups, but still important!). A person's
MN blood type is determined by his or her alleles of a certain gene.
An L^MLML, start superscript, M, end superscript allele specifies production
of an M marker displayed on the surface of red blood cells, while
an L^NLNL, start superscript, N, end superscript allele specifies production of
a slighly different N marker.

Homozygotes (L^ML^MLMLML, start superscript, M, end superscript, L, start

superscript, M, end superscript and L^NL^NLNLNL, start superscript, N, end
superscript, L, start superscript, N, end superscript) have only M or an N
markers, respectively, on the surface of their red blood cells. However,
heterozygotes (L^ML^NLMLNL, start superscript, M, end superscript, L, start
superscript, N, end superscript) have both types of markers in equal numbers
on the cell surface.

As for incomplete dominance, we can still use Mendel's rules to predict

inheritance of codominant alleles. For example, if two people
with L^ML^NLMLNL, start superscript, M, end superscript, L, start
superscript, N, end superscriptgenotypes had children, we would expect to
see M, MN, and N blood types and L^ML^MLMLML, start superscript, M,
 end superscript, L, start superscript, M, end superscript, L^ML^NLMLNL,
start superscript, M, end superscript, L, start superscript, N, end superscript,
and L^NL^NLNLNL, start superscript, N, end superscript, L, start superscript,
N, end superscript genotypes in their children in a 1:2:11:2:11, colon, 2,
colon, 1 ratio (if they had enough children for us to determine ratios
accurately!)

Multiple alleles
Mendel's work suggested that just two alleles existed for each gene. Today,
we know that's not always, or even usually, the case! Although individual
humans (and all diploid organisms) can only have two alleles for a given
gene, multiple alleles may exist in a population level, and different
individuals in the population may have different pairs of these alleles.

As an example, let’s consider a gene that specifies coat color in rabbits,

called the CCC gene. The CCC gene comes in four common
alleles: CCC, c^{ch}cchc, start superscript, c, h, end superscript, c^hchc, start
superscript, h, end superscript, and ccc:
 A CCCCC, C rabbit has black or brown fur
 A c^{ch}cchc, start superscript, c, h, end superscriptc^{ch}cchc, start
superscript, c, h, end superscript rabbit has chinchilla coloration (grayish fur).
 A c^hc^hchchc, start superscript, h, end superscript, c, start superscript, h,
end superscript rabbit has Himalayan (color-point) patterning, with a white
body and dark ears, face, feet, and tail
 A ccccc, c rabbit is albino, with a pure white coat.
 Allelic series of the color gene C in rabbits.

 A CCCCC, Crabbit has black fur.

 A c^{ch}cchc, start superscript, c, h, end superscriptc^{ch}cchc, start
superscript, c, h, end superscriptrabbit has chinchilla coloration (grayish fur).
 A c^hc^hchchc, start superscript, h, end superscript, c, start superscript, h, end
superscriptrabbit has Himalayan (color-point) patterning, with a white body
and dark extremities.
 A ccccc, crabbit is albino, with a pure white coat.
Image credit: "Characteristics and traits: Figure 5," by OpenStax College, Biology (CC BY 3.0).

Multiple alleles makes for many possible dominance relationships. In this

case, the black CCC allele is completely dominant to all the others; the
chinchilla c^{ch}cchc, start superscript, c, h, end superscriptallele is
incompletely dominant to the Himalayan c^hchc, start superscript, h, end
superscript and albino ccc alleles; and the Himalayan c^hchc, start
superscript, h, end superscript allele is completely dominant to the
albino ccc allele.

Rabbit breeders figured out these relationships by crossing different rabbits

of different genotypes and observing the phenotypes of the heterozygous kits
(baby bunnies).