Heterosis and It’s Exploitation

Syllabus regarding Heterosis

• Concept of heterosis/ hybrid vigour

• Estimation of heterosis
• Inbreeding depression
• Male sterility and self incompatibility
• Principles of hybrid seed production
• Types and use of heterotic hybrids
 Teaching References
1. Chopra, V.L. 2001: Breeding Field Crops, Oxford and IBH Publishing
Co. Pvt. Ltd. New Delhi/ Calcutta.
2. Chopra, V.L. 2005: Plant Breeding Theory and Practice, 2nd Edition,
Oxford & IBH Pub. Co. Pvt. Ltd. New Delhi.
3. Dipak Kr Kar and Soma Halder. 2010. Plant Breeding Biometry
Biotechnology. New Central book Agency(P) Ltd. London, Delhi,Kolkata
4. Sing, P. 2004: Essentials of Plant Breeding. Kalyani Publishers, New
Delhi.
5. Sinha, U. and Sinha, S.1998: Cytogenetics, Plant Breeding and Evolution,
Vikas Publishing House Pvt. Ltd. Delhi.
6. Sing, B.D. 2005: PLANT BREEDING Principles and Methods. Seventh
Revised and Enlarged Edition, Kalyani Publishers, New Delhi.
7. Frankel, R. (ed.) 1983. Heterosis: Reappraisal of Theory and Practice.
Monograph, Theor. Appl. Genet. Vol. 6. Springer, Berlin.
 Objectives and Learning Outcome
Objectives: Study of the phenomenon of hybrid vigor for the production
of heterotic hybrids in different crop plants.

Learning Outcome:
After completing this chapter you should be able to:
 Define and compare between heterosis and inbreeding depression
 Explain how heterosis is manifested
 Describe the causes of heterosis
 Explain the mechanisms of male sterility
 Describe the ways to exploit the phenomenon of male sterility
 Explain the procedure of hybrid seed production in different field crops?
 Annotate the advantages and disadvantages of heterosis breeding?
 Meaning of Heterosis:

• When two homozygous inbreeds (a true breeding line

obtained by continuous inbreeding) of dissimilar
genotype are crossed together, the resulting hybrids
( F1 ) obtained from the cross are usually robust
বলিষ্ঠ, vigorous জোরালো , productive and taller than

the either parents.

• This increased productivity or superiority over the

parents is known as heterosis or hybrid vigor.
Heterosis can be defined as the superiority of
F1 hybrid over both the parents in terms of yield or
some other desirable characters.
 Heterosis: concept
• The term was first used by Shull in
1914
• Heterosis is also known as hybrid
vigor, or outbreeding enhancement.
• It is defined as superiority of ‘F1
hybrid’ over it’s both parents in term of
yield or some other characters
• The adjective derived
from heterosis is heterotic.
 Heterosis: concept

• In the F1, all offspring are heterozygotes that show high degree of
heterosis.

• In the F2, random mating has occurred, reducing the frequency of

heterozygotes, that leads declination in heterosis.

• Some times false heterosis may appears ,called Luxuriance.

• Luxuriance is the increased vigor and size of interspecific hybrids.

• Luxuriance and heterosis : the prime difference is:

• Heterosis is accompanied with an increased fertility ,
• But luxuriance is generally sterile or poorly fertile
 History of Heterosis:
• Heterosis has been known since the art of hybridization came into existence.
• Koelreuter (1763) was the first to report hybrid vigour in the hybrids of tobacco,
Datura etc.
• Mendel (1865) observed this in pea crosses.
• Darwin (1876) also reported that inbreeding in plants results in deterioration of
vigour and the crossing in hybrid vigour.
• Beal (1877-1882) reported that F1 hybrids yield as much as 40 percent more of the
parental varieties.
• From subsequent studies on inter-varietal crosses in maize, it was observed that
some of the hybrids show heterosis.
• Gottingen ( Germany), Dr. G.H. ShuII (1914) proposed the term heterosis (In Greek
heteros = different and osis = condition).
• Poweri (1944, 45) reported that the crossing, however, may result in either weak or
vigorous hybrids as compared to parental inbreeds.
 Directions of Heterosis

Heterosis may manifest in either of two

directions:

Possitive Negative

•Positive heterosis: • Negative heterosis:

• Increase in grain yield • Early maturity
• Tiller number • Dwarfness etc
• Increase in protein •Less unwanted chemical
content, vitamins etc. properties
 Manifestation of Heterosis
Increased yield: More disease resistant:
In terms of grain, fruit, leaf, Hybrids sometimes results into
tuber or any other harvest of development of more resistant
interest. to pathogens
Increase in size and general
vigor: Heterosis results in more Increased reproductive ability:
vigorous growth leading to Hybrids exhibit heterosis by
healthier and faster growing expressing high fertility rate
plant than parents
Better quality:
In most cases heterosis results Increase in growth rate:
higher yield accompanied with In many cases hybrids show
better quality. faster growth than parents

Greater adaptability:
Early flowering and maturity:
Heterotic hybrids are generally
Hybrids may show earliness in
more adapted to environment
flowering and maturity than
than inbred lines due to
parents
 Heterosis in cross- and self-pollinated
species

• Magnitude of heterosis is more

higher in cross-pollinated species
than self-pollinated
• Heterosis, commercially exploited in
cross-pollinated crops- maize,
sorghum, cotton, sunflower, onion
etc.
• Heterosis, commercially exploited in
self-pollinated crops- rice , wheat,
tomato etc.
 Types of heterosis

• Types of heterosis on the basis of two different

criteria are given below
• On the basis of origin and nature

– Euheterosis or true heterosis: Mutational heterosis and

Balanced heterosis
– Pseudoheterosis or luxuriance
• On the basis of types of estimation

– Average or Relative heterosis

– Heterobeltiosis
– Useful or standard or Economic heterosis
 Types of Heterosis: On the basis of origin and
nature
Heterosis is of two types:
1. True heterosis (euheterosis); 2. Pseudo-heterosis.
• True Heterosis:
• It is inherited i.e, from parents to offsprings during reproduction. It can be
further divided into two types:
• (a) Mutational true Heterosis: It is the sheltering or shadowing of the
deleterious, un-favourable, often lethal, recessive mutant genes by their
adaptively superior dominant alleles.
• (b) Balanced true heterosis: It arises out of balanced gene combinations
with better adaptive value and agricultural usefulness.
• Pseudo Heterosis:
• Crossing of the two parental forms brings in an accidental, excessive and
un-adaptable expression of temporary vigor and vegetative overgrowth.
• It is also called luxuriance.
• Such types are not inherited from parents to offsprings during
reproduction
Differences between mutational and balanced
heterosis
 Types of Heterosis: On the basis of estimation
or measurement

Types of heterosis on the basis of estimation :

1.Mid-Parent (MP) heterosis/ average heterosis: F1-MP X 100

(F1 performs better than mean of two parents): MP

2.Better Parent (BP) heterosis/ heterobeltiosis: F1-BP X 100

(F1 performs better than better parent): BP

3. Standard / Economic/ or useful heterosis F1-CK X 100

(F1 performs better than the check variety): CK

* Standard heterosis is the most useful term in commercial crop production

 Inbreeding
• Inbreeding = mating of related individuals/ mating between
close relatives

• Often results in a change in the mean of a trait i.e., decrease

in vigor

• Inbreeding is intentionally practiced to:

– create genetic uniformity of laboratory stocks

– produce stocks for crossing (animal and plant breeding)

• Inbreeding is unintentionally generated:

– by keeping small populations (such as is found at zoos)

– during selection
 Inbreeding Depression
• Inbreeding leads to decrease in fitness due to increase in
homozygosity or decrease in heterozygosity.
• Increase in homozygosity increases expression of
deleterious recessives
• Decrease in heterozygosity reduces fitness
 Inbreeding Depression and
Fitness traits

Inbred Outbred
 Inbreeding and Inbreeding
depression
• Inbreeding consists of any system of mating that leads to an
increase in homozygosity in cross pollinated crops

• Inbreeding in cross pollinated crops may be practiced through

– Self pollination (in case of naturally cross pollinating crops
like corn)
– Sib mating (in crops where seed setting is problem due to
self incompatibility)
1. Half-sib mating: mating between plants that have one
parent in common

2. Full-sib mating: mating between plants within the progeny

of a single plant

• Decline in vigor and size in plants due to inbreeding is referred

to as inbreeding depression
 Basis of heterosis
• Genetic basis: Three possible causes:
1. Dominance:Partial to complete dominance
2. Overdominance
3. Epistasis

The issue for breeders - What is the Ideal genotype?

– Partial to complete dominance – Homozygote ( rice, wheat,
tomato etc)
– Overdominance – Heterozygote (Maize, sorghum etc)

• Epigenetic basis : Physiological causes

 Dominance Hypothesis
The hypothesis was proposed by Davenport (1908) and further
explained by Bruce, Keeblke and Pellew( 1910)

– Hybrid vigor is due to action and interaction of favorable dominant

alleles

– Recessive alleles are deleterious

– The deleterious recessive alleles of one parent are suppressed by the

dominant alleles of another parent
– Conversely, inbreeding depression is due to exposure of these
recessive alleles during inbreeding
– Both the parents differ for dominant genes
 Dominance Hypothesis
Example
• Model AA = Aa > aa - AA=10 ,Aa=10 and aa=0
Parent 1 Parent 2
aaBBccDDee = 20 AAbbCCddEE = 30
F1
AaBbCcDdEe = 50

Also note that AABBCCDDEE = 50

But homozygous plants with all dominant alleles are
practically impossible
 Overdominance
First proposed by Shull (1908) and later expanded by East
(1936) and Hull (1945)
• It states that the heterozygote (Aa) at one or more loci is superior to
either homozygote (AA or aa)
• Model would be Aa > aa or AA
• They recognized importance of dominance, but it alone cannot account
for observed heterosis.

• Overdominance is sometimes called as single gene heterosis,

superdominance, cumulative action of divergent alleles and stimulation
of divergent alleles .
 Epistasis
• Epistasis - interaction among loci, may also contribute to
heterosis

• Types of epistatic interactions:

i) additive x additive
ii) dominance x dominance
iii) additive x dominance

 Out of above three genetic explanations of heterosis, the

dominance hypothesis is most widely accepted
 Three hypotheses to explain the genetic
mechanism of hetero­sis

• Phenotype is the sum of gene effects (A, B, A + B).

• (a) The domi­nance model; dominant alleles (A and B) suppress or complement the recessive alleles (a and b).
• (b) The overdominance model; heterozygo­sity (B 1 /B 2 ) at the key locus contributes to heterosis leading to
superior performance.
• (c) The epistasis model; non-allelic genes (A 2 and B 1 ) inherited from the parental lines interact and contribute
to heterosis.
 Physiological Basis of heterosis
Hybrid vigor, is essentially a physiological manifestation, this may be
explained by the following hypotheses:
• Initial capital and physiological stimulus- large embryo and seed size in
hybrids provide initial advantage to the hybrid during germination and
early growth of seedlings
• Balanced metabolism and heterosis at molecular level-many of the
enzymes of heterotic plants exhibit greater efficiency over their parents
• Mitochondrial complementation and heterosis- mitochondria of
heterotic hybrid absorb more O2 and have high
phosphorylation/oxidation(P/O) ratio than those of inbreds.
 Physiological Basis of heterosis

• Greater ability for uptake and utilization of nutrients-

efficient uptake and assimilation of nutrients by hybrids
seedlings confer the following advantages:
 Large number of leaf primordia

 High carboxylase and photophosphorylation activity

 Greater leaf area and larger number of leaves

 More branches per panicle (Rice) and more grains per branch

 High grain weight

 Fixation of Heterosis
Principal ways of fixation of heterosis are-
 Asexual reproduction-

 vegetative reproduction
 Apomixis- in both the cases sexual gametes production doesn’t required,
that’s why there is no chance of genetic segregation.

 Balanced lethal system-homozygotes in lethal condition (a/a) die. In some

plants balanced lethal (A/a) condition suppresses genetic segregation
leading to fix heterosis. Example: Evening primorse.

 Polyploidy- in some cases heterosis may be fixed by chromosome

doubling, e.g. wheat x rye – hybrid is sterile, but the amphidiploid is
fertile.
 Heterosis breeding

Development of hybrid varieties for genetic improvement of yield is

referred to as heterosis breeding.
The following factors are important to develop commercial hybrids:
• Enough magnitude of heterosis

• High percentage of out crossing

• Floral biology- floral biology should permit large scale production

of hybrid seeds with less expenditure
• Available of male sterile (MS) and self incompatible (SI) lines
 Main features of hybrid varieties
• Productivity: hybrid variety must be Highly productive
• Genetic constitution: population must be heterozygous but
homogeneous
• Adaptability: High adaptability to environment
• Application: Should be applicable both in self- and cross-
pollinated species
• Resistance: Should be resistant to biotic and abiotic stresses
 Hybrid seed production
• Hybrid cultivars?
• Hybrid cultivars are the first generation offspring of a cross
between different inbred line parents
• In self pollinated crops, segregating populations are selfed
for 5/6 generations to attain homozygosity, after which a
pure line is selected and cultivated as cultivar. Prior to 1900
same was practiced for cross pollination. This procedure
was a mild form of inbreeding that led to a reduction in
heterozygosity of breeding lines, and gradual loss in vigor
and productivity.
• GH Shull in 1909 proposed a method for producing hybrid
cultivars of corn. Hybrid cultivar development has now been
 Hybrid Cultivar in cross pollinated crops is produced in 3 steps

• Development of inbred lines (normally by several generations

of inbreeding in a natural or segregating population of a cross
pollinated species).

• Production of single-cross F1 hybrid cultivar with many

heterozygous loci by crossing pairs of unrelated inbred lines

• Seed Increase of the single-cross hybrid cultivar for

distribution to the growers
 An inbred line is a homozygous breeding line
developed and maintained by self pollination
S0

S1 Reduction in vigor and size in corn with

successive generations of inbreeding
S2
S3
S4
S5
S6
 Hybrid Corn

• Development of inbred lines: by selfing of heterozygous

populations and/or by chromosome doubling of haploids
• Selection of productive inbreds: inbreds with good combining
ability are selected
• Use single or double Cross with inbred lines to produce hybrid
varieties :the tassels of female parents are removed and natural
cross pollination is allowed
* seed production is carried out in isolation to prevent crossing with
unwanted genotypes and to maintain high digree of genetic
purity
 “Single Cross” Corn

• problem was that inbred parents not very

productive, so it is difficult to produce enough
seeds for farmer

Solution = “Double Cross” Corn

• start with four inbred lines, make 2 single cross

hybrids, then cross the single cross hybrids to
produce the seed corn -= double cross corn
Hybrid Corn
 Hybrid seed production in self-
pollinated crops
• Development of CMS (cytoplasmic male sterile
lines) called ‘A’ line
• Development of Maintainer line called ‘B’ line
• Multiplication of CMS line
• Selection of fertility restorer line, called ‘R’ line
• Cross between A x R lines to produce hybrid
seeds
What is Hybrid Rice?

The first generation offspring of a rice cross between two genetically

diverse parents
 How Hybrid Rice?

Normal Rice Spikelet Sterile Rice Spikelet Hybrid Seed Production

(self pollinated crop) (Male Sterility) (Male Sterile x Normal Rice)
How Hybrid Rice?
How Hybrid Rice?
CMS and fertility restorer lines in
alternative rows
Why Hybrid Rice?

 Heterosis (Hybrid vigor) Application to Increase:

• Productivity (yield/unit/time, 15-20% of yield advantage), and
• Economic returns
 Heterosis
 A universal phenomenon that F1 generation shows
superiority to both parents in agronomic traits or yield
 It presents in all biological systems and has been exploited
commercially in many agricultural crops.
Brief history of hybrid rice

 1926 - Heterosis in rice reported

 1964 - China started hybrid rice research
 1970 - China discovered a commercially usable genetic tool for
hybrid rice (male sterility in a wild rice = Wide Abortive )
 1973 - PTGMS rice was found in China
 1974 - First commercial three-line rice hybrid released in China
 1976 - Large scale hybrid rice commercialization began in China
 1979 - IRRI revived research on hybrid rice
 Brief history of hybrid rice
 1981 - PTGMS rice genetics and application was confirmed
 1982 - Yield superiority of rice hybrids in the tropics confirmed (IRRI)
 1990s - India and Vietnam started hybrid rice programs with IRRI
 1991 - More than 50% of China’s rice land planted to hybrids
 1994 - First commercial two-line rice hybrid released in China
 1994 - 1998 - Commercial rice hybrids released in India, Philippines
Vietnam

Bangladesh:
• 2001- BRRI hybrid dhan1:season-Boro: Life :155days: yield-8.5 T/ha
• 2008- BRRI hybrid dhan2:season-Boro: Life :145days: yield-8.0 T/ha
• 2009- BRRI hybrid dhan3:season-Boro: Life :145days: yield-9.0 T/ha
• 2010- BRRI hybrid dhan4:season-Aman:Life :118days :yield-6.50 T/ha
Hybrid Rice Area in other Asia Countries

Hybrid Rice Area (1,000 ha)

Country 1997 2001 2004 2005 2006

Bangladesh 15 50 90 150 (Exp)

India 100 200 560 NA

Indonesia 10 NA

Myanmar 2 42 NA

Philippines 13 189 367 300(DS)

Vietnam 188 480 650 NA

Total 288 710 1,445

Male Sterility Systems in Rice

 Male sterility: a condition in which the pollen grain is

unviable or cannot germinate and fertilize normally to set
seeds.
An inability to produce or to release functional pollen as a
result of failure of formation or development of functional
stamens, microspores or gametes.

Plant that do not produce viable, functional pollen grains

 Three types of sterility:
• 1. “Pollen sterility” in which male sterile individuals differ from normal

only in the absence or extreme scarcity of functional pollen grains (the

most common and the only one that has played a major role in plant

breeding)

• 2. “Structural or staminal male sterility” in which male flowers or

stamen are malformed and non functional or completely absent

• 3. “Functional male sterility” in which perfectly good and viable pollen

is trapped in indehiscent anther and thus prevented from functioning

 Type of Male-sterile
 Based on its inheritance or origin
 Cytoplasmic male sterility (CMS) = sterile cytoplasm (S)
Male steril comes about as a result of the combined action of nuclear genes and
genic or structural changes in the cytoplasmic organellar genome
maternally inherited
 Nuclear male sterility (NMS) = Genic, genetic, Mendelian
Male sterility is governed solely by one or more nuclear genes
Nuclear inherited
 Non genetic, chemically induced male sterility
Application of specific chemical (gametocides or chemical hybridizing agents)
 Three lines in hybrid rice seed production
The A-line, B-line and R-line model for hybrid seed production
• A-Line is female or seed producing line. It is recessive non-restorer
gene carrier. Cytoplasmic male sterility is induced through back
crossing
• B-line is exactly alike A-line but has normal cytoplasm instead of
male sterile cytoplasm. It is also called as maintainer line
• R-line is the pollen parent in the cross to produce hybrid seed. It is
also called as fertility restorer line

The function of R-line is

– Pollinate A-Line
– Restore fertility in hybrid seed
– Combine with A-line to produce vigorous hybrid seed
Commercial Systems for Hybrid Rice
Simple hybrid with cms and restoration

C1 C1
CMS line (A-line) N1 x N1 Maintainer line (B-line)
CMS, rfrf N, rfrf

Large amounts C1

x
C2
N1 Male line (C-line)
of CMS line N2
N and RfRf

C1
Fertile F1 hybrid
CMS, Rfrf
 Commercial application of heterosis
• Few examples where the heterosis has been utilised for
improvement of the crop plant are:
• Crop Species:
• Asexually propagated species and also cross pollinated species
like maize, jowar, bajra, sunflower, legume, cotton, castor etc.
• Self pollinated species: Rice.wheat, pigeon pea, jute etc.
• Vegetable Crops:
• Tomato, brinjal, onions, cucurbits, etc.
• Fruits:
• In almost all the fruit trees.