Heterosis Inbreeding Depression

Heterosis & inbreeding
depression
Dr. Prabhat Kr. Singh
Assistant Professor,
Department of Genetics and Plant Breeding
MSSSoA, CUTM, Paralakhemundi, Odisha, India
Introduction
❖ Cross pollinated species & species reproducing
asexually are highly heterozygous. When these
species are subjected to selfing or inbreeding they
show severe reduction in vigourand fertility.
❖ This phenomenon is known as inbreeding
depression.
Inbreeding
It is mating between individuals related by descent
or having common ancestry. (Brother- Sister mating
or sib mating). The highest degree of inbreeding is
obtained by selfing.
Heterosis
❖ The term heterosis was first used by Shull in1914.
❖ It is defined as the superiority of F1 hybrid over both
the parents in terms of yield or some other characters.
❖ It may be true heterosis or euheterosis to distinguish
it from luxuriance.
❖ Can be occurred in all types of species i.e. self-
pollinating, cross-pollinating and vegetatively
propagated.
❖ Koelreuter (1673) in tobacco, Darwin (1876) in several
vegetable crops were studied about hybrid vigor.
❖ But our present knowledge on heterosis comes from
the work on maize.
Types of heterosis/ Estimation of
heterosis
1. Average heterosis or Relative heterosis: It is the
heterosis where F1 is superior to mid parent value. In
other words superior to average of two parents.
__ ___
F1 - MP
Average heterosis= x 100
MP
Where, F1= Mean of hybrid, MP = Mid parental value
This type of heterosis is of no use in agriculture since

the superiority is below the better parent value.
2.Heterobeltiosis: Superiority of F1 over the better
parent.
__ ___
F1 - BP
Heterobeltiosis = x 100
BP
Where, BP = Mean of better parent
3.Economic or useful heterosis: Superiority of the F1

compared to the high yielding commercial variety in a
particular crop.
__ ___
F1 - CV
Economic heterosis = x 100
CV
Where, CV = Mean of commercial variety

4. Standard heterosis: Sometime heterosis worked out
over the standard commercial hybrid.
__ ___
F1 - SH
Standard heterosis = x 100
SH
Where, SH = Mean value of standard (local commercial) hybrid
5. Negative heterosis: Performance of F1 inferior to

better parent / mid parent value. – e.g. Duration
(earliness), height in cereals, micronaire value in cotton,
and toxic substances like neurotoxin in Lathyrus sativus.
Heterosis and hybrid vigour
❖ Hybrid vigour is used as synonym of heterosis.
❖ Hybrid vigour only refers to superiority of F1 over better
parent, while heterosis describes other situation as well.
❖ In other words hybrid vigour is manifested effect of
heterosis. Thus the term hybrid vigour is used to distinguish
the F1 superiority from negative heterosis.
Luxuriance vs Heterosis
➢ Luxuriance is the increased vigour and size of interspecific
hybrid.
➢ The principal difference between heterosis and luxuriance lies
in the reproductive ability of the hybrid.
➢ Heterosis is accompanied with an increased fertility, while
luxuriance is expressed by interspecific hybrid. That are
generally sterile or poorly fertile.
Manifestation of heterosis May be
in the following form.
1. Increased yield.
2. Increased reproductive ability.
3. Increase in size and vigour.
4. Better quality
5. Greater adaptability.
6. Earlier flowering and maturity
7. Greater resistant to disease and pest
8. Faster growth rate
9. Increase in the number of a plant part
Genetic basis of heterosis and
inbreeding depression
There are three main theories of heterosis and
inbreeding depression.
1. Dominant hypothesis
2. Over dominance hypothesis.
3. Epistasis hypothesis.
1.Dominant hypothesis
❖ First proposed by Davenport in 1908. It was later on
expanded by Bruce, Keeble and Pellow (1910).
❖ According to this hypothesis at each locus the
dominant allele has favourable effect, while the
recessive allele has unfavourable effect. In
heterozygous state, the deleterious effect of
recessive alleles are masked by their dominant alleles.
Inbreeding depression is produced by the harmful
effects of recessive alleles, which become homozygous
due to inbreeding.
❖ Therefore according to this hypothesis heterosis is
not due to heterozygosity rather it is the result of
prevention of effect of harmful recessive by their
dominant allele.
Objections against the dominant hypothesis
a) Failure of isolation of inbreds as vigorous as hybrids:
According to dominance hypothesis it is possible to isolate
inbreds with all the dominant genes E.g. AA. This inbreed
should be as vigourous as that of hybrid. However in
practice such inbreds were not isolated.
b) Symmetrical distribution in F2
In F2 dominant and recessive characters segregate in
the ratio of 3: 1. Quantitative characters, according to
dominance hypothesis should not show symmetrical
distribution. However, F2 nearly always show
symmetrical distribution.
Explanation for the two objections
In 1917 Jones suggested that since quantitative

characters are governed by many genes, they
are likely to show linkage. In such a case inbreds
containing all dominant genes cannot be isolated.
So also the symmetrical distribution in F2 is due
to linkage. This explanation is often known as
Dominance of Linked Genes Hypothesis.
2. Over dominance hypothesis
❖This hypothesis was independently proposed by East
and Shull in 1908. It is also known as single gene
heterosis or super dominance or cumulative of
divergent alleles or stimulation of divergent alleles.
❖According to this hypothesis, heterozygotes at least
some of the loci are superior to both the homozygotes.
❖Thus heterozygote Aa would be superior to AA and aa.
❖In 1936 East proposed that at each locus there are
several alleles e.g. a1, a2 a3, a4… etc, with
increasingly different functions. So heterozygotes
between more divergent alleles would be more
heterotic E.g. a1 a4 will be superior to a1 a2, a2a3 or
a3 a4.
Evidence for Overdominance hypothesis
There are not many clear-cut cases where the heterozygote
is superior to the two homozygotes; but there is no doubt
that in the case of some oligogenes, heterozygotes are
superior to the homozygotes.
1. In case of maize, gene ma affects maturity. The

heterozygote Ma ma is more vigorous and later in anthesis
and maturity than the homozygotes Ma Ma and ma ma.
2. Gustafsson has reported two chlorophyll mutants in barley
that produce larger and more number of seeds in the
heterozygous state than do their normal homozygotes.
3. Similarly, heterozygotes for the hooded gene in barley
show a higher rate of photosynthesis than the two
homozygotes.
4. In human beings (Homo sapiens), sickle cell anaemia is
produced by a recessive gene s which is lethal in the
homozygous state. In Africa, the heterozygotes Ss are at
a selactive advantage over the normal SS individuals
because they are more resistant to malaria.
5. Another case of heterozygote advantage is reported in
Neurospora crassa (bread mold). Gene pab is concerned
with the synthesis of p-aminobenzoic acid. The
heterozygote pab+ pab is more vigorous and shows a
faster growth rate than the two homozygotes pab pab and
pab+pab+.
However, the superiority of heterozygotes need not be a

result of overdominance. It could more easily be due to
linkage in the repulsion phase or epistatic effects, i.e.,
an interaction between two or more nonalleles.
A comparison between dominance and overdominance
hypotheses of heterosis
3.Epistasis Hypothesis
❖ In 1952, Gowen had suggested that influence of one locus on
the expression of another may be involved in heterosis.
❖ Theoretically, epistatic interactions will lead to the maximum
heterosis when the following two conditions are met with.
(1)First, the epistasis should be predominantly of complementary
type, i.e., the estimates of h (dominance effects) and /
(dominance x dominance interaction effects) have the same
sign so that they do not cancel each other out.
(2)Second, the interacting pairs of genes should be dispersed in
both the parents.
❖ It has been suggested that in the absence of overdominance,
dispersion (between the two parents of hybrids) of genes
showing complementary epistasis seems to be the major cause
of heterosis.
Fixation of heterosis
1. Asexual reproduction: easily conserved in vegetatively
propagated crops.
2. Apomixis: Seeds develops without fertilization.
Apomictic seeds generally developed from maternal
diploid cell and hence identical to mother plants. Citrus,
blackberry, many flwering plants, Hieracium (Hawck
weed) are the examples where heterosis can be fix by
apomixis.
3. Balance Lethal System: Homozygotes are lethal and
hence die, only heterozygotes survive. E.g. Evening
primerose (Oenothera spp.), BLS developed due to
complex translocation.
4. Polyploidy: By chromosome doubling or polyploidy,
especially interspecific and intergeneric hybrid. E.g.
wheat-rye cross

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Heterosis & inbreeding

Where, F1= Mean of hybrid, MP = Mid parental value

This type of heterosis is of no use in agriculture since

Where, BP = Mean of better parent

3.Economic or useful heterosis: Superiority of the F1

Where, CV = Mean of commercial variety

Where, SH = Mean value of standard (local commercial) hybrid

5. Negative heterosis: Performance of F1 inferior to

In 1917 Jones suggested that since quantitative

1. In case of maize, gene ma affects maturity. The

However, the superiority of heterozygotes need not be a

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