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Law of Inheritance

This document discusses Mendel's laws of inheritance, including: - Mendel's law of segregation which states that alleles segregate equally into gametes such that offspring have an equal chance of inheriting either allele. This explains the 3:1 ratio Mendel observed. - Mendel's law of independent assortment which states that genes do not influence each other and all combinations of alleles are possible. This is illustrated through a dihybrid cross example. - The forked-line method, an alternative to Punnett squares for more complex crosses involving multiple genes, applies the product rule to calculate probabilities.

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102 views

Law of Inheritance

This document discusses Mendel's laws of inheritance, including: - Mendel's law of segregation which states that alleles segregate equally into gametes such that offspring have an equal chance of inheriting either allele. This explains the 3:1 ratio Mendel observed. - Mendel's law of independent assortment which states that genes do not influence each other and all combinations of alleles are possible. This is illustrated through a dihybrid cross example. - The forked-line method, an alternative to Punnett squares for more complex crosses involving multiple genes, applies the product rule to calculate probabilities.

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Laws of Inheritance

LEARNING OBJECTIVES
By the end of this section, you will be able to:

Explain Mendel’s law of segregation and independent assortment in terms of


genetics and the events of meiosis
Use the forked-line method and the probability rules to calculate the probability of
genotypes and phenotypes from multiple gene crosses
Explain the effect of linkage and recombination on gamete genotypes
Explain the phenotypic outcomes of epistatic effects between genes
Mendel generalized the results of his pea-plant experiments into four postulates,
some of which are sometimes called “laws,” that describe the basis of dominant
and recessive inheritance in diploid organisms. As you have learned, more complex
extensions of Mendelism exist that do not exhibit the same F2 phenotypic ratios
(3:1). Nevertheless, these laws summarize the basics of classical genetics.

Pairs of Unit Factors, or Genes


Mendel proposed first that paired unit factors of heredity were transmitted
faithfully from generation to generation by the dissociation and reassociation of
paired factors during gametogenesis and fertilization, respectively. After he crossed
peas with contrasting traits and found that the recessive trait resurfaced in the F2
generation, Mendel deduced that hereditary factors must be inherited as discrete
units. This finding contradicted the belief at that time that parental traits were
blended in the offspring.

Alleles Can Be Dominant or Recessive


Photo shows an albino child with his black mother.
Figure 1. The child in the photo expresses albinism, a recessive trait.

Mendel’s law of dominance states that in a heterozygote, one trait will conceal the
presence of another trait for the same characteristic. Rather than both alleles
contributing to a phenotype, the dominant allele will be expressed exclusively. The
recessive allele will remain “latent” but will be transmitted to offspring by the
same manner in which the dominant allele is transmitted. The recessive trait will
only be expressed by offspring that have two copies of this allele (Figure 1), and
these offspring will breed true when self-crossed.

Since Mendel’s experiments with pea plants, other researchers have found that the
law of dominance does not always hold true. Instead, several different patterns of
inheritance have been found to exist.

Equal Segregation of Alleles


Observing that true-breeding pea plants with contrasting traits gave rise to F1
generations that all expressed the dominant trait and F2 generations that expressed
the dominant and recessive traits in a 3:1 ratio, Mendel proposed the law of
segregation. This law states that paired unit factors (genes) must segregate equally
into gametes such that offspring have an equal likelihood of inheriting either factor.
For the F2 generation of a monohybrid cross, the following three possible
combinations of genotypes could result: homozygous dominant, heterozygous, or
homozygous recessive. Because heterozygotes could arise from two different
pathways (receiving one dominant and one recessive allele from either parent), and
because heterozygotes and homozygous dominant individuals are phenotypically
identical, the law supports Mendel’s observed 3:1 phenotypic ratio. The equal
segregation of alleles is the reason we can apply the Punnett square to accurately
predict the offspring of parents with known genotypes. The physical basis of
Mendel’s law of segregation is the first division of meiosis, in which the
homologous chromosomes with their different versions of each gene are segregated
into daughter nuclei. The role of the meiotic segregation of chromosomes in sexual
reproduction was not understood by the scientific community during Mendel’s
lifetime.

Independent Assortment
Mendel’s law of independent assortment states that genes do not influence each
other with regard to the sorting of alleles into gametes, and every possible
combination of alleles for every gene is equally likely to occur. The independent
assortment of genes can be illustrated by the dihybrid cross, a cross between two
true-breeding parents that express different traits for two characteristics. Consider
the characteristics of seed color and seed texture for two pea plants, one that has
green, wrinkled seeds (yyrr) and another that has yellow, round seeds (YYRR).
Because each parent is homozygous, the law of segregation indicates that the
gametes for the green/wrinkled plant all are yr, and the gametes for the
yellow/round plant are all YR. Therefore, the F1 generation of offspring all are
YyRr (Figure 2).
ART CONNECTION
This illustration shows a dihybrid cross between pea plants. In the P generation, a
plant that has the homozygous dominant phenotype of round, yellow peas is
crossed with a plant with the homozygous recessive phenotype of wrinkled, green
peas. The resulting F_{1} offspring have a heterozygous genotype and round,
yellow peas. Self-pollination of the F_{1} generation results in F_{2} offspring
with a phenotypic ratio of 9:3:3:1 for yellow round, green round, yellow wrinkled
and green wrinkled peas, respectively.
Figure 2. This dihybrid cross of pea plants involves the genes for seed color and
texture.

In pea plants, purple flowers (P) are dominant to white flowers (p) and yellow peas
(Y) are dominant to green peas (y). What are the possible genotypes and
phenotypes for a cross between PpYY and ppYy pea plants? How many squares do
you need to do a Punnett square analysis of this cross?

For the F2 generation, the law of segregation requires that each gamete receive
either an R allele or an r allele along with either a Y allele or a y allele. The law of
independent assortment states that a gamete into which an r allele sorted would be
equally likely to contain either a Y allele or a y allele. Thus, there are four equally
likely gametes that can be formed when the YyRr heterozygote is self-crossed, as
follows: YR, Yr, yR, and yr. Arranging these gametes along the top and left of a 4
× 4 Punnett square (Figure) gives us 16 equally likely genotypic combinations.
From these genotypes, we infer a phenotypic ratio of 9 round/yellow:3
round/green:3 wrinkled/yellow:1 wrinkled/green (Figure 2). These are the
offspring ratios we would expect, assuming we performed the crosses with a large
enough sample size.

Because of independent assortment and dominance, the 9:3:3:1 dihybrid


phenotypic ratio can be collapsed into two 3:1 ratios, characteristic of any
monohybrid cross that follows a dominant and recessive pattern. Ignoring seed
color and considering only seed texture in the above dihybrid cross, we would
expect that three quarters of the F2 generation offspring would be round, and one
quarter would be wrinkled. Similarly, isolating only seed color, we would assume
that three quarters of the F2 offspring would be yellow and one quarter would be
green. The sorting of alleles for texture and color are independent events, so we
can apply the product rule. Therefore, the proportion of round and yellow F2
offspring is expected to be (3/4) × (3/4) = 9/16, and the proportion of wrinkled and
green offspring is expected to be (1/4) × (1/4) = 1/16. These proportions are
identical to those obtained using a Punnett square. Round, green and wrinkled,
yellow offspring can also be calculated using the product rule, as each of these
genotypes includes one dominant and one recessive phenotype. Therefore, the
proportion of each is calculated as (3/4) × (1/4) = 3/16.

The law of independent assortment also indicates that a cross between yellow,
wrinkled (YYrr) and green, round (yyRR) parents would yield the same F1 and F2
offspring as in the YYRR x yyrr cross.

The physical basis for the law of independent assortment also lies in meiosis I, in
which the different homologous pairs line up in random orientations. Each gamete
can contain any combination of paternal and maternal chromosomes (and therefore
the genes on them) because the orientation of tetrads on the metaphase plane is
random.

Forked-Line Method
When more than two genes are being considered, the Punnett-square method
becomes unwieldy. For instance, examining a cross involving four genes would
require a 16 × 16 grid containing 256 boxes. It would be extremely cumbersome to
manually enter each genotype. For more complex crosses, the forked-line and
probability methods are preferred.

To prepare a forked-line diagram for a cross between F1 heterozygotes resulting


from a cross between AABBCC and aabbcc parents, we first create rows equal to
the number of genes being considered, and then segregate the alleles in each row
on forked lines according to the probabilities for individual monohybrid crosses
(Figure 3). We then multiply the values along each forked path to obtain the F2
offspring probabilities. Note that this process is a diagrammatic version of the
product rule. The values along each forked pathway can be multiplied because
each gene assorts independently. For a trihybrid cross, the F2 phenotypic ratio is
27:9:9:9:3:3:3:1.

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