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Inbreeding and Heterosis

By Laura Merrick, William Beavis, Jode Edwards, Thomas Lübberstedt, Arden Campbell, Deborah Muenchrath,
Shui-Zhang Fei (ISU)

Except otherwise noted, this work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License.
Introduction

Inbreeding

This module focuses on inbreeding, a type of mating of individuals that is often of particular signi�cance to
plant breeders. Inbreeding is de�ned as the mating of individuals that are related by ancestry. Self-pollination
(mating of an individual to itself) represents the most extreme form of inbreeding. Inbreeding leads to an
increase in homozygosity at the expense of heterozygosity. A key feature of inbreeding is that as homozygosity
increases in a population undergoing inbreeding in the absence of selection, the genotype frequency changes
while the allele frequency stays unchanged. Inbreeding may occur unintentionally as a result of selection or
maintenance of small populations. Inbreeding is also deliberately practiced as a method to create genetic
uniformity in populations of interest for genetic or breeding research, for retaining genotypes of inbred cultivars
of self-pollinated species through many years of production, or for reliable production of inbred lines to be used
in the development of commercial hybrid cultivars.
Inbreeding Depression

A phenomenon known as inbreeding depression—the reduced survival and fertility of offspring of related
individuals—occurs in both plants and animals, showing that variation for heritable �tness traits occurs within
populations. The occurrence of inbreeding depression varies across species. Charles Darwin, British naturalist
famous for his theories of evolution and natural selection was the �rst person to make a distinction between
plants that are outbreeders (species with reproductive mechanisms promoting cross-pollination, who typically
exhibit inbreeding depression and tend to be intolerant of inbreeding, e.g., maize and alfalfa) and inbreeders
(species with reproductive mechanisms promoting self-pollination in which inbreeding depression is minimal,
who tolerate many generations of inbreeding, e.g., wheat and oat). Darwin noticed that:

• outcrossing is more common in nature than self-fertilization

• there are complex reproductive systems that promote outcrossing in plants
• many plant species have evolved systems that prevent self-fertilization.

This module also describes hybrid vigor or heterosis, which is a phenomenon that is functionally the opposite
of inbreeding depression. Heterosis is de�ned as the increased vigor of F1 progeny resulting from the mating of
inbred parents. Generally, the performance of the F1 hybrid exceeds the performance of its inbred parents for
various traits. The expression of hybrid vigor might affect phenotypes under single gene or polygenic control,
e.g., size, growth rate, fertility, and yield. The �rst-generation offspring of crosses from mating between
different pure-line inbreds generally show in a greater amount of desired traits of both parents, but hybrid vigor
decreases in F2 and subsequent sel�ng generations due to inbreeding. The exploitation of hybrid vigor or
heterosis is a key feature in the success of hybrid cultivars.
Cross-Pollination

Several mechanisms promote cross-pollination.

• Emergence or maturity of the staminate and pistillate �owers is asynchronous.

◦ Protandry - anthesis occurs before stigma are receptive.
◦ Protogyny - pistillate �ower matures before the staminate �ower.

• Flowers are monoecious or dioecious.

• Mechanical obstruction between the staminate and pistillate �owers in the same individual prevents self-
pollination. Alfalfa �owers, for example, have a membrane over the stigma that precludes self-pollination.
When a bee lands on the �ower, the keel is tripped, rupturing the membrane and exposing the stigma to
pollen carried by the bee from other plants it has visited, effecting cross-pollination.

• Gametes produced on the same plant or clone are unable to effect fertilization.
◦ Self-sterility - gametes from same individual cannot successfully fuse to form a zygote. Sterility
can be caused by lack of function of pollen (male gametes) or ovules (female gametes). Male
sterility, either genetic or cytoplasmic, occurs because the pollen is not viable. Female sterility
occurs when the ovule is defective or seed development is inhibited.
◦ Self-incompatibility - self-pollination may occur, but fertilization and seed set fail.

Pollen is transported from the staminate �ower to the pistillate �ower by wind, insects, or animals. Occasionally
pollen is transported to receptive stigma of the same individual and self-pollination may occur. For example,
pollen from the tassel of a maize plant may land on and pollinate silks on the same plant, effecting self-
pollination.

Sun�ower is ordinarily cross-pollinated. Bees often carry pollen from one plant and deposit it on other plants.

Fig. 1 Beehives on the edge of a sun�ower �eld. Photo by Fig. 2 Close-up of a sun�ower head. Photo by Iowa State
Iowa State University. University.
Self-Pollination

Several �oral mechanisms enforce self-pollination.

• Flowers do not open, preventing external pollen from reaching the stigma.
• Anthesis occurs before the �ower opens.
• Stigma elongates through the staminal column (�laments and anthers) immediately after anthesis.
• Floral organs may obscure the stigma after the �ower opens.

Although these mechanisms usually enforce self-pollination, a low frequency of cross-pollination may occur.
The frequency of cross-pollination in normally self-pollinating species generally depends on the species and
environmental conditions.

Soybean is an example of a species that is normally self-pollinated. Before the �ower opens, the anthers burst
and pollen grains fall out of the anthers on to the receptive stigma contained in the same �ower-self-pollination
occurs.
Objectives

• Understand the effects of inbreeding.

• Be able to assess the amount of inbreeding by consideration of the inbreeding coe�cient.
• Compare and contrast systems of mating that promote inbreeding—self-pollination, half-sib mating, full-
sib mating, and backcrossing.
• Learn why inbreeding depression occurs and know why it happens more commonly in species that are
predominately cross-pollinated vs. those that are predominately self-pollinated.
• Understand the effects of heterosis and know the difference between the dominance vs. the
overdominance hypotheses for explaining its occurrence.
Genetics of Inbreeding

Inbreeding as a Probability

Inbreeding is characterized by a departure from random

mating and involves preferential mating between relatives.
The most signi�cant effect of inbreeding is that replicates
of a single allele in a common ancestor can come together
through the mating of relatives to produce homozygous
progeny whose alleles are identical by descent. In contrast
in a second type of homozygote, two alleles that are said
to be alike in state are those that have the same function,
but did not derive from replication of a single ancestral
gene. These two types of homozygotes can be understood
by analyzing pedigrees such as the ones shown in Fig.3.
The two copies of the A1 allele in the A1A1 offspring
individual at the bottom of Fig. 3a descend from the same
copy in a common ancestor; this type of genotype is
termed autozygous. In contrast the two copies of the
A1 allele in the A1A1 offspring individual at the bottom in
Fig. 3b are descended from two different copies in
ancestors; this type of genotype is allozygous.

Inbreeding can be estimated directly by studying pedigrees

or lineages of ancestry showing genetic relationships.
Fig. 3 Individuals homozygous by descent or by state.
Measurement of Inbreeding

The probability that two genes are identical by descent is called the coe�cient of inbreeding (denoted as F) and
will be the measure of relationship between mating pairs.

The coe�cient of inbreeding (F) is de�ned as the probability that two alleles at the same locus are identical by
descent. At the population level, F describes the average level of homozygosity. The coe�cient of inbreeding is
always expressed relative to a speci�ed base population. The base population is de�ned to be non-inbred (F=0).
The range of F is 0 to 1, with 0 indicating random mating and no inbreeding, while 1 means prolonged sel�ng.

Consider a base population consisting of N individuals each shedding equal numbers of gametes uniting at
random (although see Fehr, 1987, p.113, with respect to the concept of effective population size, or Ne).
Because the base is de�ned to have F=0, each individual in this population carries genes that are non-identical.
The only way a homozygote that carries genes that are identical by descent can arise is by the mating of a male
and female gamete from the same individual that carry a replication of the same gene. Because there are 2N

gametes the probability that two mating gametes are identical by descent is .
Second Generation

In the second generation there are two ways genes that are identical by descent can be joined: 1) by a new
replication of the same ancestral gene; and 2) by the previous replication that occurred in generation 1. The

probability of a new replication event is . The remaining proportion of zygotes, , carry

genes that are independent in origin from Generation 1, but may have been identical in their origin in Generation
0. The probability the genes are identical by descent from Generation 1 is the inbreeding coe�cient of

Generation 1 . Note that F1, F2, Ft, etc in this section (written in italics) denote inbreeding
coe�cients, and not generations as on pages 1 and 2 of this section and in earlier sections.

Therefore the probability of identical homozygotes in Generation 2 is:

Equation 1

Where F1 and F2 are the inbreeding coe�cients of Generations 1 and 2. The same arguments apply to future
generations, so we can write the recurrence equation:

Equation 2
Generation Inbreeding

The inbreeding of any generation is composed of two components: New inbreeding, which arises from self-
fertilization and the "old" that was already there.

Note that inbreeding is cumulative and the absence of inbreeding in Generation t does not change the fact that
a population had old inbreeding from prior generations.

Through a series of algebraic steps, we can write the inbreeding coe�cient as a function of the number of
generations removed from the base populations:

where,
Genotype Frequencies

The genotype frequencies in a population can then be expressed as:

Table 1 Genotype frequencies for a locus with two alleles. Data from Falconer ad Mackay, 1996.

Original Change due to Origin

Frequencies inbreeding
Independent Identical

AA

Aa

aa

Allozygous genes Autozygous

genes

We can examine inbreeding or the probability of identity of alleles by descent, by looking at the genotype
frequencies in Table 1 under two extremes of the F statistic:

• if F = 0 (random mating; no inbreeding)

◦ the equation for genotypic frequencies reduces to the familiar equation for genotypes in Hardy-
Weinberg proportions: p2 + 2pq + q2

• if F = 1 (alleles identical by descent)

◦ the genotype equation reduces to a ratio of homozygotes to heterozygotes as p:0:q
Inbreeding Coefficient

Thus inbreeding leads to homozygosity (all or nearly all loci homozygous), and almost a complete absence of
heterozygosity (all or nearly all loci heterozygosity). As noted in Table 1, with inbreeding there is a de�cit of
heterozygotes equal to 2pqF and an excess of each homozygous class equal to half the de�ciency of the
heterozygotes.

To illustrate one type of mating that promotes inbreeding, consider the case of a population that reproduces by
self-fertilization so that F = 1.

Another way to express the inbreeding coe�cient, F, is to compare the frequency of heterozygotes in the
population to the frequency expected under random mating:

Equation 3

where:

H = frequency of heterozygotes in the population

H0 = expected frequency under HWE, meaning 2pq

Therefore the inbreeding coe�cient is the proportional reduction in heterozygosity relative to a random mating
population with the same allele frequencies.
Inbreeding via Self-Pollination

In a population that reproduces by self-fertilization the inbreeding coe�cient, F = 1. Let’s assume the population
begins with genotypic frequencies in Hardy-Weinberg proportions (p2 + 2pq + q2). With sel�ng,
each homozygote produces only progeny of the same genotype:

AA × AA ⇒ all AA

aa × aa ⇒ all aa

However only half of the progeny of a heterozygote will be like the parent

Aa × Aa ⇒ 1/4 AA, 1/2 Aa, 1/4 aa

Self-pollination therefore reduces the proportion of heterozygotes in the population by half with each
generation until all genotypes in the population are homozygous (Table 2).

Table 2 Increased over generations in frequency of homozygous in a self-pollinated population starting with p = q =
0.05>>>frequency of homozygotes.

Genotype Frequencies

Generation AA Aa aa

1 1/4 1/2 1/4

2 1/4 + 1/8 = 3/8 1/4 1/4 + 1/8 = 3/8

3 3/8 + 1/16 = 7/16 1/8 3/8 + 1/16 = 7/16

4 7/16 + 1/32 = 15/32 1/16 7/16 + 1/32 = 15/32

1/2 0 1/2
Proportion of Homozygotes

With multiple segregating loci, the proportion of homozygotes in various sel�ng generations can be estimated
using the following formula: [(2m – 1)/2m]n: where m is the number of sel�ng generations(m = 1 for F2; m = 2
for F3 and so on) and n is the number of segregating loci. For example, an F1 plant with four independent
segregating loci will result in the following frequency of homozygous plants in F2, [(21 – 1)/21]4 = 1/16 = 6.25%.
The expected proportion of completely homozygous plants in F2 and later generations of sel�ng for different
numbers of segregating loci are indicated in Table 3. The proportion of homozygotes decreases sharply with
increasing heterozygosity (more segregating loci) in the F1.

Table 3 Frequency of completely homozygous individuals in various sel�ng generations in relation to the number of
segregating loci in F1.

Number of segregating loci in the F1 Frequency of completely homozygous individuals (%)

F2 F4 F6 F8

1 50.00 87.5 96.87 99.22

2 25.00 76.56 93.85 98.44

3 12.50 66.99 90.91 97.67

4 6.25 58.62 88.07 86.91

5 3.13 51.29 85.32 96.15

10 0.01 26.31 72.79 92.45

100 7.89 x 10-31 6.1 x 10-4 4.18 45.64

1000 9.33 x 10-302 1.02 x 10-58 1.63 x 10-58 0.04

Study Questions 1
What is the frequency of susceptible plants in the self-pollinated population without selection in the
F2 and F4 generations assuming disease resistance is controlled by a single resistance gene, R and
that F1 plants are heterozygous for the disease resistance locus?

Enter your answer here.

Show Answer
Consequences of Inbreeding

Increasing Inbreeding

Through increasing homozygosity, inbreeding brings together identical alleles at a locus. Homozygosity permits
the expression of recessive alleles that may have been previously masked by heterozygosity in the parent
generation. If recessive alleles are less favorable than dominant ones, the overall �tness of the individual
decreases. Inbreeding is often detrimental because it increases the appearance of lethal and deleterious
recessive traits. The term inbreeding depression describes the decrease in �tness or performance that often
accompanies inbreeding or random genetic drift. Recall that �tness is the relative ability of an individual to
survive and reproduce to contribute its genes to the next generation. Inbreeding depression is further described
in the next section.

Plant stature, vigor, yield and other traits decline with increasing inbreeding, although signi�cant differences
exist among species for the amount of inbreeding depression expressed—ranging from minimal among self-
pollinated crops such as oat and wheat to severe in cross-pollinated polyploid species such as alfalfa, whereby
homozygous genotypes do not survive (Fig. 4). Table 4 on the next screen summarizes some general
differences between outbreeders and inbreeders.

Fig. 4 The effect of inbreeding on plant stature, vigor, yield and other traits.
Outbreeders vs. Inbreeders

Table 4 General features of reproduction and population genetics pertaining to outbreeders vs. inbreeders. Data from
Simmonds and Smartt, 1999.

Outbreeder Inbreeder

Has crossing mechanism, approaches random mating Closed �owering, approaches regular
sel�ng

Individuals heterozygous at many loci Individuals approach homozygosity

Variability distributed over population Variability mostly between component

lines

Carries deleterious recessives Deleterious recessives tend to be

eliminated

Intolerant of inbreeding Tolerant of inbreeding

Much heterozygote advantage (epistasis, Less heterozygote advantage

overdominance)
Inbreeding Depression

Inbreeding Depression and Homozygosity

Inbreeding results in increased homozygosity. In cross-pollinated species, increased homozygosity results in

inbreeding depression or reduced performance. Symptoms of inbreeding depression may include:

• Reduced plant vigor

• Smaller plant size
• Decline in fertility
• Suppressed seed production
• Decreased pollen production
• Inferior seed quality
• Greater susceptibility to insect or pathogen damage or
• Poorer standability.

Why? With increased homozygosity, expression of deleterious recessive alleles may be revealed by �xation that
had been masked by more favorable dominant alleles, or it can be caused by overdominance in which
homozygotes are less �t, resulting in poorer performance.

The severity of inbreeding depression varies with the species and genotype. If the inbreeding depression is too
severe, it may be di�cult or impractical to maintain or propagate inbred lines by seed. In such cases, some
heterogeneity must be maintained or other propagation means used.
Overdominance Hypothesis

Over-dominance

The phenotype of the heterozygous progeny is greater than either parent.

Mating Systems

The frequency of homozygotes in a population can be increased using any of several mating systems:

• Self-pollination—Repeated self-pollination is routinely used to develop pure lines of self-pollinating

crops. It can also be applied to cross-pollinating species to obtain inbred lines. Like pure-lines in self-
pollinated species, inbred lines are homozygous at nearly all loci.
• Half-sib mating—Plants having one parent in common are mated. The pollen source is random from the
breeding population, but the female plants are identi�able.
• Full-sib mating—Plants having both parents in common are crossed.
• Backcrossing—A method in which a hybrid is mated to one of its parents, the recurrent parent, resulting
in a backcross 1 (BC1) population. The resulting backcross offspring are repeatedly crossed to the
recurrent parent.

The inbreeding coe�cient increases more rapidly

with matings among more closely related
individuals. The �rst three mating systems listed
above are compared in Fig. 5. In each case the
initial value of the inbreeding coe�cient is assumed
to be F0 = 0. Backcrossing—like self-fertilization—
can be one of the most extreme forms of
inbreeding.

Fig. 5 Comparison of homozygosity reached by self-

pollination, full-sib mating, and half-sib mating.
Study Questions 2A & 2B

Interpret the trends shown on the graph in Fig. 5.

Fig. 5 Comparison of homozygosity reached by self-pollination, full-sib mating, and half-sib mating.

Which inbreeding system reaches nearly complete homozygosity �rst?

 Self-pollination

 Half-sib mating

 Full-sib mating

 Check

After the 6th generation, what is the percent homozygosity in the half-sib and in the full-sib mating
systems?

Half-sib mating: 68 %
Full-sib Mating: 87 %

 Check
Study Questions 2C

List the circumstance(s) under which the breeder would develop inbred lines using the half-sib mating or the
full-sib mating system rather than self-pollination.

Use half-sib mating when plants are:

Enter your answer here.

Show Answer

Use full-sib mating when plants are:

Enter your answer here.

Show Answer
Study Questions 3

The challenge for the plant breeder is to develop inbred lines that can be maintained and are su�ciently
homozygous to generate reliably superior and uniform progeny when mated to produce the hybrid cultivar.
Inbred lines can be developed from any heterogeneous population. Heterogeneity is essential to obtain
variability for important traits. Without variability for the characters of interest, the breeder cannot make
selections or breeding progress.

What methods could be used to develop breeding lines in self-incompatible or dioecious species, or
where certain lines are male sterile?

 Sel�ng

 Full-sib mating

 Half-sib mating

 Check
Degree of Relatedness of Individuals

The coe�cient of relatedness (rIJ) is a measure of the degree of relatedness between individuals (Table 5). It is
the proportion of genes shared between two individuals I and J due to common descent (e.g., identical by
descent). It ranges from -1.0 (no genes in common, at least among the genetic markers assessed) to +1.0
(vegetative clones). For example, the value of rIJ for a parent and each of their progeny is 0.5 for a sexually
reproducing species because half of each offspring’s genes come from each parent. In the case of full-siblings
the value of rIJ is also 0.5, but on average because siblings receive half of their genes from each of the same
pair of parents, but the haploid set of genes in each parental gamete is a random sample of half of the parental
genome due to recombination (Conner and Hartl 2004).

Table 5 Values of the coe�cient of relatedness (rIJ ) among relatives. Data from Conner and Hartl, 2004.

Mating Systems rIJ

Self-pollination*, vegetative clones, doubled haploids 1.0

Parents and offspring 0.5

Full siblings 0.5

Half siblings 0.25

*After sel�ng for many generations

Heterosis

Heterosis or Hybrid Vigor

When inbred lines of cross-pollinating species are mated, their progeny are often superior to either or both of
the parents for one or more characters, a phenomenon referred to as hybrid vigor or heterosis. Recall that
genetic drift is random, so that different inbred lines or subpopulations tend to be �xed for different alleles at
various loci; when they are intermated, the F1 population will be highly heterozygous. Heterosis is the opposite
of inbreeding depression—heterosis is commonly expressed as

• Improved plant vigor,

• Greater plant size, and/or
• Increased productivity.

The performance of a hybrid relative to its parents can be described in two ways: mid-parent heterosis is the
performance of a hybrid compared with the average of the performance of its parents; high-parent heterosis is a
comparison of the performance of the hybrid with that of the best parent:

Equation 4

Where F1 = performance of the hybrid

MP = average performance of the parents (Parent 1 + Parent 2)/2

HP = performance of the best parent

Dominance vs. Overdominance Hypothesis

Heterosis tends to be greatest in the progeny of diverse genotypes. There are two common explanations for
this phenomenon:

1. Dominance hypothesis—Heterosis results from the complete or partial dominance of favorable alleles at
various loci.
2. Overdominance hypothesis—Heterosis re�ects the superior performance of the heterozygote over either
homozygote.

The next few pages provide a summary of the dominance and overdominance hypotheses from Charlesworth
and Willis (2009). Note the differences between models for a single locus and those describing multiple loci.
They include a third model that involves deleterious alleles at closely linked loci and focuses on so-called
“pseudo-overdominance”, which will not be further discussed here.
Dominance Hypothesis

Complete dominance

The phenotype of the heterozygous progeny equals the phenotype of the homozygous dominant parent.

Partial (incomplete) dominance

The heterozygous progeny has a phenotypic value greater than that of the midparent value, but less than that of
the homozygous dominant parent.

Overdominance

The phenotype of the heterozygous progeny is greater than either parent.

Main Genetic Hypothesis

Recessive deleterious mutations

Dominance hypothesis

Fig. 6 Summary of the main genetic hypotheses for inbreeding depression and heterosis. Adapted from Charlesworth and
Willis, 2009.

Recessive deleterious mutations at closely linked loci

Pseudo-overdominance
Fig. 7 Summary of the main genetic hypotheses for inbreeding depression and heterosis. Adapted from Charlesworth and
Willis, 2009

Single locus with heterozygous advantage

True overdominance
Fig. 8 Summary of the main genetic hypotheses for inbreeding depression and heterosis. Adapted from Charlesworth and
Willis, 2009.
Heterozygote Advantage

Heterozygote advantage is a term that is frequently confused with heterosis. However, heterozygote advantage
is a synonym of overdominance, and refers to a condition in which the heterozygous genotype has a higher
phenotypic value (especially for �tness) than for either homozygous genotype. Heterosis is usually due to a
number of loci that control quantitatively inherited traits, although depending on the situation it could be due to
phenotypes controlled by either single genes or polygenic traits. In contrast, heterozygote advantage describes
effects at a single locus (Conner and Hartl 2004). Compare the circumstances for these two concepts in Table
6.

Table 6 Fitness effects of high and low frequency of heterozygotes. Data from Donner and Hartl, 2004.

Subpopulations

Small and isolated Crossed with each other

Genotypic Highly homozygous Highly heterozygous

frequencies

Fitness Low High

Fitness effects Inbreeding depression Heterosis or hybrid vigor

Cause for �tness Deleterious recessives expressed or Deleterious recessives masked or

effects loss of heterozygote advantage occurrence of heterozygote advantage
Genetics of Heterosis and Inbreeding Depression

Whatever genetic mechanisms can explain heterosis must also be able to explain inbreeding depression. Let’s
take a look at hypothetical numeric examples of these two hypotheses to show how each can explain both.

Although conclusive evidence to support either of these hypotheses remains elusive, the dominance hypothesis
has been more widely accepted. However, it is recognized that a gene's effect is determined by its effect in
combination with:

• Other alleles at the same locus,

• Itself (when homozygous),
• Genes at other loci (epistasis), and
• Closely linked genes.

Thus, dominance, overdominance, and epistasis, most likely all contribute to heterosis.
Heterosis and Inbreeding Depression

In the following examples we will use a scenario where six genes control a certain quantitative characteristic.

1. Favorable Dominant Alleles Theory (with complete dominance at each locus)

a. Heterosis

The homozygous recessive genotype, aa bb cc dd ee ff has a value of 40. One dominant allele at
any of the six genes increases the value by 10, that is, N_=+10.

The following two homozygous parents are crossed. Each has a value of 70 because of having
three loci with dominant alleles [40+(3)(10)=70].

Note that the F1 has a value of 100 because of having six loci with dominant alleles (40 + (6)(10) =
100).

b. Inbreeding depression

Inbreeding depression can be explained with the accumulation of favorable dominant alleles
mechanism. If we self the Aa Bb Cc Dd Ee Ff individuals that have a value of 100, we will get a
population of individuals as follows:

Table 7

Percentage of individuals Number of loci with at least one dominant allele

17.80% 6

35.60% 5

29.66% 4

13.18% 3

3.30% 2

0.44% 1

0.02% 0
This gives an overall population mean value of 85. This is lower than the value of the Aa Bb Cc Dd Ee Ff
parental individuals, thus showing depression upon inbreeding.
Overdominance Theory

2. Overdominance Theory
a. Heterosis

The genotype aa bb cc dd ee ff has a value of 40.

Homozygous dominant allele at any locus increases the value by 7 over the homozygous recessive
allele, i.e. NN = +7.

Heterozygous at any locus increases the value by 10 over the homozygous recessive, i.e. Nn = +10.

The following homozygous parents are crossed; each has a value of 61 because of having three
loci with homozygous alleles [40 + (3)(7) = 61]. The F1 has a value of 100 because of having six
heterozygous loci.

b. Inbreeding depression

Inbreeding depression can be explained with overdominance. If we self the Aa Bb Cc Dd Ee Ff

individuals that have a value of 100, we will get a population of individuals with a mean value of 85
(see table above). As was the case for the “favorable dominant alleles theory”, this value is lower
than the value of the Aa Bb Cc Dd Ee Ff parental individuals, thus showing depression upon
inbreeding.
Genetic Analysis

Molecular techniques are now being used to try to shed more light on the mechanisms involved in heterosis. In
a study published in 2006, Swanson-Wagner et al. assayed the gene expression of 13,999 maize genes from
seedling plants of two inbred parents and their F1 hybrid. When comparing the three genotypes they found
1,367 of the genes produced signi�cantly different amounts of messenger ribonucleic acid (mRNA), which
performs an essential role in protein synthesis.

Analysis of these genes showed that 78 percent of them had additive gene action, 15 percent showed either
high-parent or low-parent dominance, and 3 percent showed either overdominance or underdominance. (The
other 45 genes showed non-additive gene action but with the statistical analysis used these genes could not be
classi�ed as dominant, overdominant, or underdominant.) The experiment analyzed individual genes and was
not designed to evaluate interactions among the genes, so epistasis could not be measured.

This study involved only two maize inbred lines and their hybrid offspring and the evaluation was based on
tissue from seedling plants. Different genes will be active during different parts of the life cycle of the maize
plant and further studies will most likely show differential gene action among different genes. Likewise
different parents and hybrids and different species may well show very different gene action patterns among
the same genes that were tested in this study. With just this one experiment, where nearly 14,000 genes were
evaluated, we can see that heterosis is complex and that multiple types of gene action are involved. Molecular
methods in the context of hybrid breeding will be discussed in detail in the molecular plant breeding course.

When two genetically different parents are mated, heterosis is observed in their F1 progeny.

• All seed and forage species express some level of heterosis.

• Most cross-pollinated species express greater heterosis than do self-pollinated species.

Because of the expense in producing hybrid cultivars, hybrid cultivars are generally developed only in those
crops in which the level of heterosis results in signi�cantly better performance.
Reflection
The Module Re�ection appears as the last "task" in each module. The purpose of the Re�ection is to enhance
your learning and information retention. The questions are designed to help you re�ect on the module and
obtain instructor feedback on your learning. Submit your answers to the following questions to your instructor.

1. In your own words, write a short summary (< 150 words) for this module.
2. What is the most valuable concept that you learned from the module? Why is this conceptvaluable to
you?
3. What concepts in the module are still unclear/the least clear to you?
References
Charlesworth, D. and J.H. Willis. 2009. The genetics of inbreeding depression. Nature Reviews Genetics 10:
783-796.

Conner, J. K., and D.L. Hartl. 2004. A Primer of Ecological Genetics. Sinauer Associates, Sunderland, MA.

Falconer, D.S. and T.F.C. Mackay. 1996. Introduction to Quantitative Genetics. 4th edition. Longman Pub. Group,
Essex, England.

Pierce, B.A. 2010. Genetics: A Conceptual Approach. 3rd edition. W.H. Freeman, New York, NY.

Simmonds, N.W. and J. Smartt. 1999. Evolution of Crop Plants. 2nd edition. Longman, Essex, UK.

Swanson-Wager, R.A., Y. Jia, R. DeCook, L.A. Borsuk, D. Nettleton, and P.S. Schnable. 2006. All possible modes
of gene action are observed in a global comparison of gene expression in a maize F1 hybrid and its inbred
parents. PNAS 103:6805-6810.
Effective Population Size
The concept of effective population size, denoted by the symbol Ne, was introduced by Sewall Wright as number
of breeding individuals in an idealized (as small as possible, i.e., “redundant” individuals would be eliminated)
population showing the same distribution of allele frequencies under random genetic drift or inbreeding as the
population under consideration. The effective population size is usually smaller than the absolute population
size (N). As noted in pages 112-114 in the Fehr textbook, Ne is a relative measure of the number of parents
mating in a population >>> Ne (see Fehr, 1987, p. 112-114).

The general equation for Ne is the following:

Equation 5

where:

N = number of individuals that are mating

Fp = coe�cient of inbreeding for parents

Ne is a basic parameter in many models in population genetics, and will be explained in more detail in the
Population and Quantitative Genetics for Breeding course.
Acknowledgements
This module was developed as part of the Bill & Melinda Gates Foundation Contract No. 24576 for Plant
Breeding E-Learning in Africa.

Crop Genetics Inbreeding and Heterosis Author: Laura Merrick, William Beavis, Jode Edwards, Thomas
Lübberstedt, Arden Campbell, Deborah Muenchrath, and Shui-Zhang Fei (ISU)

Multimedia Developers: Gretchen Anderson, Todd Hartnell, and Andy Rohrback (ISU)

How to cite this module: Merrick, L., W. Beavis, J. Edwards, T. Lübberstedt, A. Campbell, D. Muenchrath, and S. Fei.
2016. Inbreeding and Heterosis. In Crop Genetics, interactive e-learning courseware. Plant Breeding E-Learning in
Africa. Retrieved from [Link]
Source URL: [Link]
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