Dog Days of Science ■ 123
and brown offspring that would result if two
heterozygous (Bb) black-coated dogs were
mated. It is important to understand that the
predicted ratios simply give the probability that
a particular offspring will have a certain pheno-
type or genotype; the actual ratio will vary
(see “What Are the Odds?” on page 124).
Peas in a Pod
Mendel’s research on pea seeds led to his second
law, the law of independent assortment.
This law states that when gametes form, the
two alleles of any given gene segregate during
meiosis independently of any two alleles of
other genes. For example, pea seeds can have
a round or wrinkled shape, and they can be
yellow or green. Two different genes control the
two different traits—the R gene, with alleles R
(round) a nd r (wrinkled), controls seed shape,
while the Y gene, with alleles Y (yellow) and
y (green), controls the color of the seed—but
neither gene affects the inheritance of the
other.
Mendel tested the idea of independent assort-
ment in the set of experiments illustrated in
Figure 7.6. He tracked seed shape, the trait
controlled by the R and r alleles, and seed color,
controlled by the Y and y alleles. The test of his
hypothesis came when he examined the pheno-
types of the offspring produced by crossing the
heterozygous F 1 plants (RrYy). As predicted by
the hypothesis, two new phenotypic combina-
tions were found among the F 2 offspring: plants
with round, green seeds (RRyy or Rryy) and
plants with wrinkled, yellow seeds (rrYY or
rrYy). Figure 7.6 summarizes the ratios of the
two parental phenotypes and the two novel,
nonparental phenotypes.
Traits controlled by a single gene and
unaffected by environmental conditions are
called Mendelian traits. But when Mendel
described his laws of inheritance, he had no
idea what genes were made of, where they
were located within a cell, or how they segre-
gated and independently assorted. Today we
know that genes are located on chromosomes
and that these chromosomes are the basis
for all inheritance. We call these assertions
the chromosome theory of inheritance,
which explains the mechanism underlying
P generation
F 1 generation
F 2 generation
Eggs
Sperm
RrYy
RRYY rryy
Phenotypes of
Mendel’s F 2 plants
appear in a ratio
close to 9:3:3:1. The
presence of round,
green and wrinkled,
yellow offspring
showed that the two
traits—seed shape
and seed color—are
inherited
independently.
...and when these
four types of
gametes are used to
produce the F 2
generation, a 9:3:3:1
phenotype ratio
should result.
Assuming the two traits are
inherited independently, RrYy
individuals should produce
equal numbers of RY, Ry, rY,
and ry gametes...
The phenotype of the F 1 hybrids
confirmed that round seed
shape is dominant over
wrinkled, and that yellow seed
color is dominant over green.
Mendel crossed plants with
round, yellow seeds (RRYY) and
plants with the contrasting
phenotype: wrinkled seed shape
and green seed color (rryy).
RRYY RRYy
RRYy RRyy
RrYY RrYy
RrYy Rryy
RrYY RrYy
RrYy Rryy
rrYY rrYy
rrYy rryy
RY Ry
RY Ry rY ry
RY ry
rY ry
RY
Ry
rY
ry
×
Summary of Mendel’s results
9/16 Round, yellow (315 plants)
3/16 Round, green (106 plants)
3/16 Wrinkled, yellow (101 plants)
1/16 Wrinkled, green (32 plants)
Figure 7.6
Independent assortment of pea color and shape
Mendel used two-trait breeding experiments, called dihybrid crosses, to
test the hypothesis that the alleles of two different genes are inherited
independently from each other. M
Q1: List all the possible offspring genotypes and phenotypes.
Q2: What is the offspring phenotype ratio?
Q3: Complete a Punnett square for a genetic cross of two true-
breeding Portuguese water dogs—one with a black, wavy coat
(homozygous dominant, BBWW) and one with a brown, curly coat
(homozygous recessive, bbww). What is the phenotype ratio of
their offspring (F 1 )? Now fill out another Punnett square, crossing
two of the offspring. What is the phenotype ratio of the
F 2 generation?