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Department Of Zoology

Term paper assigment

Course Code: Zol-509 Course Name: GENETICS

Due Date: 27-04-2020 Submission Date: 27-04-2020
Course Teacher: MAM NAILA Student Name: Ambreen
Masood
Student ID: BZOF17M031 Session: 2017-2021
Program: BS Zoology Semester: 6th
Regular/Self-support: Regular Main Campus/ Ex- PPP:
Main

ASSIGNMENT COVER SHEET

Title of the Assignment: HETEROSIS AND INBREEDING

Word count: 3635 (without References)
Marks awarded by teacher:
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TABLE OF CONTENT:
Topic: page number:
Heterosis and its history 3-5
Types of heterosis 5-7
Forms of heterosis 7-8
Positive vs negative heterosis 9
Fixation of heterosis 9-10
Genetic basic of heterosis 10-12
Epigenetics and heterosis 12-13
Physiological and biochemical basics of heterosis 14-15
Types of hybrids 15-17
Example and effects of hybrid vigor 17-20
Interbreeding and interbreeding depression 20-21
Degree of interbreeding depression: 21-22
Effects of interbreeding 22-23
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(A) HETEROSIS:
Heterosis is also called as hybrid vigor or outbreeding enhancement. It is defined as the
superiority of F hybrid in one or more characters like yield, adaptation and vegetative
growth over its parents.

It is a multigenic complex trait (a trait which is affected by or determined by many

genes) and can be extrapolated as the total of many physiological and phenotypic
traits.
When two homozygous individuals (true breeds) are crossed together the resulting
hybrid obtained from crossing is robust, vigorous, productive and taller than both
parents. (Shull 1952; Mackay and Falconer 1996)

(B) HISTORY:
1. Rudolph Camerarius (1665-1721)
A professor at Botanic Gardens at Tubingen, 1688
He explained the function of egg and pollen by studying the dioecious and
monoecious plants. He considered the pollen as male .and gave first modern
understanding of PLANT SEXUALITY.
2. Joesph Gottlieb kolreuter (1733-1806)
He was a German botanist. He cultivated plants for studying fertilization and
development .He performed experiments on Tobacco plant. (Nicotiana paniculata
and N.rustica). He demonstrated that hybrids obtained by crossing resemble both
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parents. He recognized two important agents during pollen transfer in plant

fertilization (1) wind (2) insects. He was the first person to observed heterosis
(hybrid vigor) in plants.

(N.rustica) (N.paniculata)
3. Thomas Andrew knight (1759-1838)
He give the principle of anti-breeding in 1799. He also described the dominance
and segregation in garden pea.
4. Gregor Mendel (1822-1884)
Also known as FATHER OF GENETICS. In 1865 He also observed hybrid
vigor in pea. And give the Law of Genetics concerning the transmission of genetic
material from parents to off springs.
5. Charles Darwin (1809-1882)
He worked on origin of species in 1859. He talked about evolution and variability.
He was the First person to describe heterosis in crop plant. He observed that
maize progeny obtained by cross pollination was 25%taller than the parents.
“I raised closed together two large beds of self-fertilized and cross seedlings from
same plant of Linaria vulgaris .To my surprise, crossed plant when fully grown
were plainly taller and more vigorous than the self-fertilized ones”-Charles
Darwin.
(The effects of cross and self-fertilization in vegetable kingdom 1876)
According to Darwin crossbreeding was more beneficial. More than 60 different
species of plants were documented by Darwin with respect to growth,
development and seed fertility as compared to their parents. (Darwin, 1876)
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(Linaria vulgaris)
6. Beal in 1880 on the bases of his experiment he first published 51% increase in
the yield of offspring (F hybrid) over the parents.
7. Johnson in 1891 in support of heterosis said “crossing give better off springs”
8. McCleur in 1892 said crossing imparted vigor while sterility was imparted due
to inbreeding.
9. East and Hayes in 1912 give the concept of heterozygosis. According to them
crossing produces heterozygosis while selfing leads to Homozygosis.
10. G.H. Shull in 1914 coined the term Heterosis (Greek word hetero means
different and osis: condition) and replaced the older term heterozygosis.
11. Jones in 1918 give practical utilization of Heterosis.
12. Boardon in 2000 explained Heterosis as increase in performance of progeny to
that of purebred parents. (Frankel, 2013)

(C) Difference between Heterosis and hybrid

vigor
According to Shull
“The developed superiority of hybrid is called hybrid vigor.”
‘’The mechanism by which this superiority is developed is called Heterosis’’

Types of Heterosis
Types of Heterosis are classified as
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(A) On the basics of nature and origin

I. Euheterosis or true Heterosis

I. Mutational Heterosis
II. Balanced Heterosis
II. Pseudoheterosis

(B) On the basics of estimation

a. Average or relative Heterosis
b. Heterobeltiosis
c. Economic Heterosis

True Heterosis / euherterosis

It is heritable .and passed from parents to off springs during the process of
reproduction. It has two types.
 MUTATIONAL Heterosis
It is the deletion of unfavorable or recessive or mutant genes by their superior,
non-adaptive alleles. Genes which are shed are mostly lethal. Superiority is
attained only in cross pollinated organisms. (Dobzhanskiy, 1952)
 BALANCED Heterosis
It is arised from the balanced gene combination having better adaptive value and
useful for agriculture. Organisms are more adaptive to environmental conditions.

Pseudoheterosis
This term was coined by Mackay in 1876. It is also known as Luxuriance. When
two parents are crossed. Progeny possess superiority over the parents in
vegetative growth but not in yield and adaption. They are poorly fertile and
mostly sterile.

Types on the basics of estimation.

 Average or relative heterosis
When heterosis is measured by average of 2 parents or mid parental value. It is
known as average or relative heterosis.
 Heterobeltiosis
This term was coined by Blizter in 1968.. It is measured by average of best/better
parents.
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 Standard heterosis
It is also known as useful or economic heterosis. And important for plant
breeding. It is measured/estimated by standard, commercial hybrid. (Rai & Rai,
2006)

Method of calculating mid parental heterosis:

(D ) Forms of heterosis
There are 3 major forms of heterosis.
1. Individual heterosis
By the breeding of crossbred animals this individual heterosis is retained. And it
is related with allele’s probability of alleles when different parents are cross
together.
2. Maternal heterosis
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From the maternal heterosis, F1 females obtained by crossing of truebreedings

will get the benefits. For example crossbred cows have various advantages over
the straight bred cows like high reproductive efficiency, high maternal ability,
increased lifetime productivity, high milk and meat production.

3. Paternal heterosis
By paternal heterosis male of F1 will get the benefits. But it is difficult to measure
because there is large influence of female in the herd .but advantages in
reproductive traits is observed. (Rai & Rai, 2006)(

Positive and negative heterosis

This was given by Powers in 1944.positive heterosis is the hybrid organism
tendency to have the traits superior than traits of either parents. Range of variation
is higher than either of the parents .for example liger which is the hybrid offspring
of lion and tigress is large in size and also stronger than both parents. Negative
heterosis means that hybrid organisms traits are inferior to the traits of either
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parents. For example tigon which is produced by the breeding of lioness and tiger
is smaller in size than parents. (Pakistan Journal of Botany, 2007)

Fixation of heterosis:
Fixation of heterosis is difficult. But up to some extent it can be fixed. Four
methods are generally used. To banana, sugarcane, sweet and potato.
 Asexual reproduction:
Vegetative propagation or asexual reproduction is a method used to fix heterosis.
This method is generally applied
 Polyploidy:
Polyploidization or doubling of chromosome are used to fix heterosis.
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It can be done in intergenic hybrid and interspecific hybrid.

 Apomixis:
To conserve hybrid vigor, diploid cells of mother can give rise to apomixes
progeny. Hybrid produced is identical to its mother. This type of fixation is
possible in rose and citrus.
 Balanced lethal system:
In some types of balanced lethal system, inly heterozygous individual can survive.
Survival of homozygous individuals is not possible. This type of fixation is
possible in Orenother. (Mohanan, 2010)

Genetics basic of heterosis

There was 3 models/ hypothesis presented.
(1) Dominance model/hypothesis
(2) Over dominance model/ hypothesis
(3) Epistasis model/ hypothesis
Dominance Hypothesis:
This hypothesis was put forward by Davenport in 1908 and later by Bruce, Keable
and Pelew in 1910. This hypothesis says that hybrid vigor is formed when
dominant / fitness factors of two organisms interact.it further ads that genes
favorable for growth and vigor are always dominant while lethal alleles are
always recessive. So this superiority of dominant genes over recessive is called
heterosis

Assumptions about Dominance hypothesis

~Heterosis is directly related to number of dominant genes which are contributed
by each parent.
~Heterosis mainly occur due to masking of lethal recessive alleles.
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~Heterosis does not occur due to heterozygosity per se.

~There is superiority of dominant alleles over recessive.

According to the Davenport 1908, Bruce 1910, heterosis is governed by the
cumulative action of many loci containing partially or completely dominant
alleles on them. (Davenport 1908; Bruce 1910; Keeble and Pellow 1910)

Objections
(1)According to dominance hypothesis, inbred isolation containing dominant
alleles is possible. The isolated inbred should be vigorous .but this isolation
practically not possible.
(2) In F2 dominance to recessive ratio is 3:1.according to dominance hypothesis
quantitative characters do not show symmetrical distribution. While in F2
distribution is always symmetrical.

Explanation for the 2 objections.

In 1917 jones give a hypothesis known as Dominance of linked gene hypothesis.
In this hypothesis he suggested that quantitative traits are controlled by many
genes which show linkage .in such a cases dominant alleles present in inbred
cannot be isolated. Linkage is also responsible for the symmetrical distribution of
F2.in 1921 Collins also showed that
symmetrical distribution such as yield is governed by polygenes.

Over dominance hypothesis

It is variously called single locus heterosis or single gene heterosis. .this
hypothesis was given by East and shull in 1908. The variety/ diversity of united
gametes is increased by a physiological stimulus to development.in 1930 fisher
called it superdominance.at more than one locus allelic interaction is there which
results in superior phenotypic expression.
Heterosis is caused by heterozygosity per se
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Homozygotes are always inferior to heterozygotes.

Amount of heterosis is directly related with increase in vigor.
Heterozygotes are superior because of
 Divergent alleles have additive action
 In heterozygote, production of superior hybrid is there.
More divergent alleles more heterosis
Evidence of over-dominance in heterokaryon of Neurospora is haploid which
have single set of chromosome in each nucleus.
Prominence and heterozygosis is not possible within same nucleus
Heterosis came from heterokaryon which formed between strains of
N.Tetrasperma heyday of overdominance.( Shull 1908 ; East 1908)
Objections
 For quantitative character like yield heterosis at single locus is not
good
 Due to linkage basis estimation of heterosis not possible
Pseudo –over dominance:
The concept of pseudo-over dominance was given by power in 1945. Pseudo-
over dominance is the genetic intermediate pf over-dominance and dominance.
Because of tight repulsion phase linkage it is basically complementation of simple
dominance. And it appears as over- dominance.
Epistasis hypothesis
This hypothesis was given by Powell in 1944, William 1959.
Epistasis interaction between non allelic genes bring the process of heterosis.
These non-allelic genes are located at two or more loci
Classical approach towards genetic basis of heterosis
 In 1952, Comstock and Robison used generation mean or diallel
dialyses to measure degree of dominance, but he ignore the effects of
epistasis in the estimation.
 Triple test cross shows the contribution of epistasis in heterosis by a
test.(Henry, 2012)

Epigenetics and Heterosis:

A change in gene expression that are not dependent on DNA sequences and
these changes are heritable either due to mitosis or meiosis.
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Types
a. DNA methylation
b. Histone modification
c. RNA interference siRNA,miRNA
DNA methylation
 Addition of methyl group to cytosine or DNA nucleotide adenine.it
is a biochemical process
 Cytosine is converted to 5-methyl cytosine
 DNA sequence/function is not changed but its expression is
changed.
 Genes containing 5-methyle cytosine in promotor region are
transcriptionally inactive or silent and associated with gene
sequencing
 Heterosis occur because of different allele are present at different
locus.
 Different locus play their role in regulatory hierarchies and in this
way control different quantitative traits
 Different alleles are formed from different methylated DNA.if
homozygosity is present in methylated DNA,regulatory hierarchies
slow down gene expression.
 If heterozygosity is present in methylated DNA than it regulated
regulatory factor for gene expression and results in heterosis
sRNA
 MicroRNA small non-coding RNA and sRNA also 20-24
nucleotide
 SRNA plays a vital role in genome regulation.it has 2 classes
 SiRNA (small interfering RNA) play a role in DNA.methylation
and chromatin modification which are stable. This DNA
methylation is guided by RdDM(RNA directed DNA methylation
complex)
Histone modification
These are covalent changes modification in histone protein which usually
occur at N-terminal tail. These modification include transcriptional
potential, nucleosome rearrangement and chromatin changes. The key
histone modification are methylation, acetylation especially lysine lysine,
methyl transferase, demethylase (HKMTs and HDMs), acetylases and
deacetylases (HATs and HDACs). (Yu et al. 1997; Li et al. 2001)
All these changes leads to epigenetic which is leading cause of heterosis
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Physiological basis of heterosis

Ashby’s hypothesis of greater initial capital
Initial weight of embryo result in heterozygosity
Hybrids are similar to their parents in vegetative growth
In short words maintance of initial advantages in embryo size in hybrid vigor
Stages of physiological manifestation of heterosis
a. Seed and embryo development
There was a relation recorded between hybrid vigor and embryo weight
Copland found that there is high vigor present in hybrid embryo during early
stages of development
Ashbey recorded that the major difference between reciprocal cross and weight of
embryo
b. Early secondary growth
Hybrid have high growth rate period
Ashby found that reserve food material is less important than embryo size.
Growth rate of hybrid is more than inbreed immediately after germination of seed
but later it was not observed.later seedling growth
According to Hoffer that hybrid contains intermediate root dry matter
Hybrid absorb and utilize more nitrogen and phosphorus than parents.
Heterotic hybrids was found to contain more catalase activity.
QTL and heterosis
QTL is also called quantitative trait loci.it is important factor in reporting genetic
basis of heterosis.It is used to identify such genes which are involved in heterosis
Different QTL are mapped for study of heterosis in rice which have dominant
over-dominant and epistasis effect. (Nobuta K, et al. 2014)
Heterosis in rice
National key laboratory of crop genetic improvement performed an experiment by
using rice as model system.
Shenzhen 97 and Minghui 63 was crossed to derive F2 generation. In rice it is
observed that heterosis was occurred due to epistasis or interaction on two louses
forming genetic basis of quantitative traits.
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Over-dominance/pseudo-over dominance causes heterosis of yield, no of grains

and grain weight.
Dominance interaction play a vital role in heterosis of grain weight and number.
For rice linkage map and fragment sequenced and mapped.
Conclusion from QTL according to its different genes affect heterosis. (Noubta.K
et al.2014)

Biochemical basis of heterosis

Bottleneck concept
This concept support the concept of dominance.
According to this concept inferiority of parent of hybrid is more important than
superiority of hybrid
Inferiority in parents is due to incomplete allele.

Types of hybrid
Offsprings which are produced by combination of unlike parents or different
varieties of plants and animals. Hybrid plants include
Sweat corn; easy to grow in garden
Meyer lemon; which is produced by cross of true lemon tree and mandarin orange
tree. Hybrid s sweeter than its parent and easy to cultivate.
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Better boy tomatoes; this hybrid is resistant to the disease and have more weight
than original tomatoes.
7

Single cross hybrid

Offspring which produced from single cross of two genetically unlike parents.
Double cross hybrid
First 2 double cross hybrid were grown in US in 1930 and 1940.Two parents of
inbreed are crossed to each other
Intergeneric hybrid
These hybrid are produced by crossing plants of different genus belonging to
same family
Hybrid will produce through pollination because they are related to each other
enough.
Cypress:-also Leyland cypress is produced by crossing a Monterey cypress
(cupressus manocarpa) and Nootka carpa (chaenaecyparis nootkatensis)
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Leyland cypress:

Cactus:-cactus ferobergia is produced by crossing genus ferocactus and genus

leuchtembergia.

Heterosis in animals
Black blady:
Which is produced by the breeding of Black Angus and Hereford by the process
of heterosis …

Equus acinus and Equus caballus are when crossed produce hybrid called mule.
Mule have high strength and adamant. And grow taller than either parent.
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Hybrid produced by the cross of jersery breed from America and led Sindhi breed
of Indian cattle contain high amount of butter in milk. Estimate is about 30%.
Mixed breed face in dogs also example of heterosis in animals. Hybrid dogs run
faster than their parents.
Commercial broilers are also produced by heterosis when white rock and white
Cornish crossed together. Broilers have large vigor and faster growth rate.
Crimson rosella are produced as the result of crossing between faveolus and
eleganus.Crimson rosella produced are more disease resistant.

Effect of heterosis
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Vigor and size:- hybrid produced are large and grow fast and are more
vigor than parents for example tomato fruit size, maize cob size
Yield:- hybrid produced are more seed,fruit,grain size and grain weight
and thus yield is more,because yield is measured in terms of
grain,seed,fruits.
Quality:- hybrid produced are of better quality for example onion hybrid
Physical effects:- hybrid have greater resistance to disease and other
parasites, adaptability of hybrid is more
Earlier maturity and flowering is seen in hybrid species
Biological effect:-hybrid have more ability to reproduce and
survive(Carena er al, 2010)
Factor affecting heterosis
4 factors affect heterosis
Genetic effect
Agronomic performance
Adaptability
Geographical and genetic diversity

Genetic effect
Parental line genetic base play a vital role in heterosis. High peak of heterosis
in hybrid shows that are present breed genetic base is involved in it
Agronomic performance
When two inbreed who yield is low and produce a hybrid then this hybrid has
high heterosis but yield is low and produce a hybrid then this hybrid has high
heterosis but yield is low
Hybrid vigor and parental performance are closely related to each other
Adaptability
Parents and hybrid has correlation on the basis of adaptability
Geographical and genetic diversity
Genetic diversity of varieties of parents and hybrid performance.(Renming et
al.2008)

Interbreeding
Interbreeding is a process of meeting of individual which are closely related to
each other by ancestry .It is common in small populations only
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In interbreeding process traits which are lethal or recessive can affect

offsprings.Because proportion of homozygosity is more in offspring due to
inbreeding.
Sickle cell anemia, hemophilia, cystic fibrosis are disorders which can be
created by inbreeding
It is opposite to heterosis.
Biological fitness is reduced. (Chen, 1993)

History
In 1700, first observation was made about inbreeding
In 1876, Charles Darwin also talked about self-fertilization and inbreeding
East in 1908 and shull in 1909 also published detail information about
inbreeding in maize.

Inbreeding depression
Reduction in fitness and fertility due to inbreeding is also called
inbreeding depression. Physiological efficiency of an organism is
reduced due to decrease in funicity and size.
In each generation heterozygosity is reduced by a factor half due to
self or interbreeding

Thus degree of inbreeding = degree of homozygosity

Interbreeding occur in humans, plants and animals.
Charles Darwin noticed wicked effect of inbreeding depression by
studying 57 plants species in which process of selfing was there.
Inbreeding depression occur due to presence of recessive genes in hybrid
Hermaphrodite species have lower inbreeding depression that outbreeding
species.( Byerly et al.1985)

Inbreeding co-efficient
It is define as way or method to measure inbreeding depression.

F = (He – Ho)/He
F {inbreeding co-efficient}
He {expected heterozygosity which is expected}
Ho {heterozygosity which is observed}
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Interpretation
F=0 without inbreeding
F=1 complete inbreeding is there

Degree of inbreeding depression

Inbreeding depression can be low or high, on these basis it has four types
1. High inbreeding depression
2. Moderate inbreeding depression
3. Low inbreeding depression
4. Lack of inbreeding depression
High inbreeding depression
It is shown by various plants species like alfalfa (M.Sative), hayfield, and
tarweed.
Plant produced by inbreeding are not able to survive because they contain lethal
allele in them.
Rate of fertility is very low and 3-4 inches are able to survive. These generations
have low yield.

Moderate inbreeding depression

It is shown by maize, bajara and jawar.Due to self-pollination these population are
able to survive up to some extent. But these generations poorly reproduced.50%
of open-pollinated varieties are yield by various inbreed lines.
Low inbreeding depression
Small breeding depression is shown by hemp, onion, sunflower etc.
Lethal or recessive characters are shown by small number of plants.
Loss in fertility is not very large and yield is also not small.
Yield of inbreed lines = open pollinated varieties
No inbreeding depression
Inbreeding depression is not shown by the species which are self-pollinated
Inbreeding depression is not developed in such species because they are produced
by self-fertilization and homozygosity is developed. (Miranda, 2010)

Homozygous and heterozygous balance

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This balance concept was given by Mather

Homozygous balance Heterozygous balance

1 Self-fertilized Cross fertilized

2 No genetic load in these species

Sum of recessive allele form
genetic load for a species
present
In a population
3 Species are vanished from Little value of recessive allele
population due to presence of for a species
recessive alleles
4 Lethal alleles are masked by
dominant ones

Effect of inbreeding
1. Lethal and recessive alleles are present and plants have such characters are
lost from that populations
2. The ability of these population to reproduce is low and lines are lost
3. Vigor is also reduced due to inbreeding and dwarf plant appear in a
population
4. Inbreeding leads to homozygosity in a population and variation in a
population are reduced
5. Inbreeding leads towards poor yield
6. Phenotypically population separate into specific lines, which occur
because of homozygosity which leads to the fixation of alleles. Genotype
and phenotype of these lines is different. ( Michod , September 1985)
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.
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References

 Aarnel R, Minranda F, Carena J.B (2010) Quantitative Genetics

in Maize Breeding: London: Lowa State University Press,
Ames, lowa 50010: p 425-471,477-523
 Botanical SP (2007) Pakistan Journal of Botany vol: 39: pp
388-393

 Chen X (1993) Comparison of inbreeding and outbreeding in

hermaphrodite Arianta arbustorum (land snail). 71(5): 456-461

 Darwin CR (1908) the effects of cross and self-fertilization in

the vegetable kingdom. John Murray, London

 Davenport CB (1908) Degeneration, albinism and inbreeding.

Science 28:454-455.

 Henry RJ (2012) Molecular Markers in Plants. Australia: Wiley

Blackwell Publication p: 120-122

 Mohanan K.V.(2010) Essentials of Plant Breeding: New Delhi:

PHI learning Private Limited New Delhi 110001: p 120-127

 R.frankel. (2013). Heterosis reappraisal theory and practice.

New York: Library of Congress Cataloging in Publication Data

 Rai N, Rai M (2006) Heterosis Breeding in vegetable crops.

New Delhi: New India Publishing Agency

 Shull GH (1908) the composition of a field of maize. Am

Breeders Assoc Rep 4:296-301
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 Shull GH (1952) Beginnings of the heterosis concept. In:

Gowen JW (ed) Heterosis: a record of researches directed
toward explaining and utilizing the vigor of hybrids. Lowa
State College Press, Ames pp 14-48

 Venu RC, Nobuta K, et al. (2014) Identification of Candidate

Genes Associated with Positive and Negative Heterosis
Rice .Plos ONE 9(4): e95178
Doi:10.1371/journal.pone.0095178

 Yinchuan L ,Renaming Z, Zhenglin Y, Guanghua H , Rong X

(2008) Preduiction of hybrid grainyield performances in indiaca
rice (Oryza sativa L.) with the effect –increasing loci.Mol
Breed 22: 467-476
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