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Application of Mutation Breeding in Crop Improvement:

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Chapter - 19

Application of Mutation Breeding in Crop

Improvement: Success Stories and Future Perspectives

1
Dikshita Hazarika*, 2Amarjeet Singh Bhogal and 3Himadri Saikia
1
MSc. (Agri.),Plant Breeding and Genetics, Assam Agricultural University
2
PhD Scholar, Plant Breeding and Genetics, Assam Agricultural University
3
PhD Scholar, Agronomy, Assam Agricultural University
*Corresponding email: dikshita.haz04@gmail.com

Abstract
The purposeful introduction of mutations in plant breeding is known as mutation
breeding. It offers prospective chances to domesticate potentially underutilized
wild species for agricultural or horticultural uses as well as to improve the ability of
already introduced crops to adapt to adverse circumstances. The primary objective
of mutant breeding is to increase the frequency and variety of mutations, as well as
the prevalence of viable mutations. Improvements in cereals, legumes, pulses, fruits
vegetables, flowers etc. have been carried out using this strategy. Yenhsing-1, the
first rice variant, was produced by cross-breeding with a mutant. Two early-ripening,
fragrant mutation-derived rice cultivars, “PNR- 381” and “PNR- 102,” were widely
cultivated in Haryana and Uttar Pradesh. Various biotic and abiotic stress resistance
cultivars have been developed in different crops in IARI, Pusa utilizing mutation
breeding.This book chapter provides insightinto the overall idea of mutation breeding
in various major crops, depicting present andfuture scenario.
Keywords: Mutation, mutation breeding, mutagens, oligogenes, polygenes
344

Introduction
The world’s population is anticipated to grow by 30% by 2050, according to the Food
and Agriculture Organisation of the United Nations (FAO). Therefore, there must
be 70% more food produced to fulfill the need of the growing population, which
will place a tremendous strain on the current climate and agricultural productivity
(Zakir2018). Agriculture outputs, which are already endangered by the environment
and limited resources, are under unprecedented pressure. There is an urgent need
of develop varieties that are high yielder, genetically diverse and tolerant to various
biotic and abiotic stresses. For the purpose of improving crops, plant breeding
needs genetic variety of beneficial features. But the needed diversity is frequently
absent. The use of mutagenic substances, like as radiation and certain chemicals, can
cause mutations and produce genetic variants from which desirable mutants can
be chosen. A tried-and-true method of introducing diversity within a crop variety
is mutation induction.
Mutation breeding is the deliberate application of mutations in the field of
plant breeding. It provides potential opportunities for the domestication of promising
underutilized wild species for agricultural or horticultural applications, as well as for
enhancing the adaption of previously introduced crops to unfavorable conditions.
Due to its ease of application, technical and financial sustainability, capacity to be
applied to all plant species, and ability to be used on small or big sizes, mutagenesis
has maintained its popularity for over a century. In 1900 Hugo de Vires coined
the term mutation and defined it as sudden heritable phenotypic alteration in an
individual. Till date over 2,000 plant varieties with induced mutations have been
formally approved for cultivation as new varieties or as parents for the creation of
new varieties without being subject to the same regulatory limitations as genetically
modified organisms (Waugh et al. 2006).
Thus, combining mutagenesis with cutting-edge molecular biology techniques
and in vitro culture technologies has improved crop improvement and breeding
programmes, especially in light of the global climate change ( Jain 2000).

Types of Mutation and Mutagenesis

Mutagenesis is the process of an organism’s genetic information changing owing
to mutagen exposure, resulting in mutation. It can happen naturally or be created
in a lab.. Mutations can cause structural changes in an encoded protein, as well as a
reduction or full loss of its expression. Since a change in the DNA sequence affects
all copies of the encoded protein, mutations may be extremely detrimental to a cell
or organism.
 345

Spontaneous Mutation:
Mutations that occur in the nature in absence of a known mutagen treatment
are known as spontaneous mutation. Spontaneous mutation occurs at a very low
frequency generally at the range of 10-5 to 10-8 in a cell. The origins of spontaneous
mutations are many and include endogenous DNA lesions, mutagenic nucleotide
substrates, and mistakes in the replication of intact template DNA. These sources vary
in frequency and the mutations they cause, and they are each impacted differently
by DNA transactions, DNA sequences, and cellular metabolism.

Induced Mutation:
When mutation is produced artificially by different physical and chemical agents it
is called induced mutations. By applying a mutagen directly to the plant or plant
components like seeds, pollen, ovules, or stem cuttings, mutation can be intentionally
produced. Radiation, chemical reactions, and transposon insertion are the three
main methods for producing mutations. Through the use of X-rays, the first induced
mutations were produced in Drosophila by Muller. In addition to X-rays, gamma
rays and rapid neutron bombardment are other radiation treatments that have been
effective. These procedures can result in deletions (loss of a chromosomal region) or
point mutations (changes in a single nucleotide).

Insertion Mutagenesis:
This kind of mutagenesis, also known as transposon mutagenesis or transposition
mutagenesis, is caused by DNA insertions, either by genetic transformation and
insertion of T-DNA (T-DNA insertion mutagenesis) or by activation of transposon
elements. The resultant mutants are transposon mutants or insertion mutants. Scientists
can examine and comprehend a variety of characteristics of gene and chromosomal
function through the use of insertional mutagenesis.

Site Directed Mutagenesis:

One of the most current molecular crop enhancement technologies is site-directed
mutagenesis, which gives researchers new methods to change the DNA sequence at
a specific region. SDM mutations may be carried either in vitro or in vivo, allowing
scientists to change the DNA of plants at a single or several places. SDM is carried
out using three different techniques: PCR-based, nucleus-based, and vector-based.

Targeted Mutagenesis:
Through the use of either induced or natural mutations, targeted mutagenesis uses
cutting-edge biotechnological techniques to breed new mutant crops. The creation
346

of novel crop varieties using targeted mutagenesis may be more practicable given the
regulatory obstacles that New Breeding Techniques (NBT) like transgenic technology
face. Targeted mutagenesis has the extra benefit of being more economically feasible
than the transgenic technique for developing traits. The time required for breeding
programmes, a fundamental bottleneck of conventional plant breeding projects,
is considerably reduced by the use of genomics in targeted mutagenesis. Targeted
mutagenesis uses genomic technologies like Next Generation Sequencing (NGS).
Targeting Induced Local Lesions in Genomes (TILLING) and Eco TILLING
are two genomic techniques that have made it possible to more effectively search
for mutants in populations.

Classification of Mutation
On the basis of phenotypic changes caused by them mutations are classified into
two categories-

a) Macromutation-These produce large changes in the traits of an or-

ganism and can be detected without the help of any instrument and
can be detected at the level of individual plant.
b) Micromutation- These produce minute or invisible phenotypic
changes and are very difficult to detect. These are of great impor-
tance in plant breeding programmes.

Molecular Basis of Gene Mutation

Presently the term mutation is used to describe those changes which alter the
chemical structure of the gene at molecular level. These changes are mostly referred
to as point mutations. Point mutations occur when a single base pair of a gene is
changed. They are divided into the following three classes based on the molecular
changes associated with them.
» Base Substitution:It is a mutation that replaces one chemical base with
another. It inserts the incorrect nucleotide in the improper location. A silent
mutation could occur from it. In such a mutation, a codon is changed to
another that encodes the equivalent amino acid, leading in no change in the
protein that is created, or it might transform an amino-acid coding codon
to a single-halt codon, resulting in the generation of an incomplete protein
that may not be functional. Additionally, it can result in a change in a codon
that specifies another amino acid, which would result in a little modification
to the protein that is produced. They can be of two type-
 347

a. Transitions- When a purine is replaced by another purine or a pyrimidine

is replaced by another pyrimidime it is known a transition.
b. Transversion- When a purine is changed into a pyrimidine or a pyrimidine
is changed into a purine, a transversion occurs.
» Base Addition: By introducing a DNA fragment, it modifies the number of
DNA bases in a gene. The resulting protein might not be useful as a result.
» Base Deletion: DNA deletion results in a change in the number of nucleotides
in the molecule. Nucleotides may be removed or deleted in any number.
Small deletions may result in the removal of a few base pairs from a gene,
but bigger deletions may result in the removal of a whole gene or nearby
genes, changing the functionality of the proteins that are produced.
» Frameshift Mutation: It occurs when a gene’s reading frame is changed as
a result of a DNA base loss or addition. In a reading frame, there are three
groups of base pairs, each of which codes for one amino acid. This kind of
mutation alters the coding for amino acids and causes a change in how
these bases are arranged. As a result, non-functional proteins are produced.
Frameshift mutations might result from duplications, deletions, or insertions.

Mutagens Used in Crop Improvement Programme

1. Physical Mutagens- The most common physical mutagens employed to cause
plant mutations include X-rays, gamma rays, beta particles, fast neutrons, slow
neutrons, and UV radiation. Ionising radiations such as X-rays, gamma rays,
neutrons, alpha and beta particles and non-ionizing radiations such as UV rays
are two categories used to classify these physical mutagens. Physical mutagens
are more advantageous than chemical mutagens since physical mutagens do not
require washing off or treating of mutagens after use, and physical mutagens do
not form waste. Scientists initially utilised physical mutagens in the 1920s when
they experimented with radium on fruit flies and learned about its mutagenic
properties. Since Muller and Stadler discovered that ionising radiation had
hereditary consequences, more than 70% of mutant types identified to date
have been created using physical mutagens. Muller demonstrated the impact of
X-ray on mutation in 1927 using fruit flies. Lewis John Stadler demonstrated
the genetic impact of X-rays in maize and barley in 1928. Physical mutagens
are now widely used for mutant breeding as a result of these two findings.
» X Ray- Scientists have been employing this in mutant breeding projects
since high doses of X-rays can result in plant mutations. Since electrons
are known to be the source of X-rays, the radiation is created by electrically
348

accelerating the electrons in a high vacuum before they are directed onto a
target, such as a tungsten, gold, or molybdenum barrier. Shorter wavelength
X-rays have a better penetration capacity, making them ideal for inducing
mutations in crops.
» Gamma Rays: Gamma irradiation has emerged as the most common
physical mutagen since the 1950s. Its widespread availability and versatility
in usage for a variety of reasons, including mutant breeding, food irradiation,
and medicinal applications, account for its widespread use. Since gamma
rays are the physical mutagens with the shortest wavelengths, they are
more energetic than other physical mutagens. There are several commercial
sources for gamma rays. Radioisotopes of cobalt-60and caesium-137 are
two examples of these sources. A naturally occurring isotope of potassium
releases gamma rays into the atmosphere.
» Alpha Particles: Alpha particles are made up of two protons and two
neutrons and therefore they posses double positive charge. These particles,
unlike gamma or X-rays, have a mass and are therefore regarded as particles.
They are lesser penetrating than X-rays and gamma rays however, they are
densely ionizing.
» Fast and Thermal Neutrons: The only place where neutrons are stable is
inside the atomic nucleus. Numerous kinetic energy will be released as a
result of neutrons separating from their nuclei. Depending on the energy
emitted, these neutrons can be classified as slow (thermal), moderate, or
rapid. Neutrons were employed to cause mutations in the 1960s and 1970s.
In contrast to existing induced mutagenesis techniques, rapid neutron
irradiation offers a novel approach since it may alter chromosomes in addition
to removing a few to several million DNA bases.
» UV Radiation: Mercury vapour lamps or tubes are used to produce UV
radiations and they are non-ionizing in nature. UV radiation’s ability to
cause mutations was originally identified by Attenberg in 1934. Since that
time, several studies of all sorts have been conducted to determine how UV
light affects crops.
2. Chemical Mutagens- For decades throughout the last century, several scientists
experimented with chemically induced mutations.Thom and Steinberger’s use
of nitrous acid to cause mutations in Aspergillus in 1939 was the first conclusive
finding of chemical mutagenesis. In 1946, Auerbach and Robson made the
conclusive discovery that mustard gas is clearly mutagenic. True gene mutations
may easily be caused by chemical mutagens. The most commonly used chemical
 349

mutagens can be grouped into four classes and they are- 1) alkylating agents 2)
base analogues 3) acridine dyes 4) deamination agent
» Alkylating agents:Since they are the most effective chemical mutagens for
producing novel plant mutants, alkylating compounds have been utilised as
plant chemical mutagens for a long time (Fujimoto and Yamagata, 1982).
Ethylenimine (EI), ethyl methanesulfonate (EMS), methyl methanesulfonate
(MMS), and mustard gas are four examples of frequently used alkylating
chemicals that can be utilised in induced mutagenesis.
» Base Analogues: These are chemically equivalent to nitrogen bases. When
they are integrated into the DNA during replication, they can result in
incorrect base pairing, which causes mutations. Two examples of base
analogues that are frequently utilized as mutagens are 2-amino purine and
5-bromo uracil.
» Acridine Dyes: Chemicals that are introduced between two DNA bases
during replication result in either the insertion of a base pair or the deletion
of one or more base pairs are known as acridine dyes. Examples include
Proflavin and Acriflavin (Arshad et al., 2006). These intercalating agents are
frequently utilised as fluorescent markers for DNA visualisation. Acridine
dyes may stretch the DNA strand in addition to acting as fluorescent markers,
enabling the removal or addition of nucleotide bases.
» Deamination agent- Acts through deamination, the replacement of cytosine
by uracil, which can pair with adenine and thus through subsequent cycles
of replication lead to transitions (Oladosu et al. 2016)

Mechanism of Physical Mutagens

Ionizing radiation - Ionizing radiations are of two types
1. Particulate Radiation- These are produced by radioactive decay and consist
of high energy. Beta rays are produced by radioactive decay of 32P, 35S etc
and are negatively charged. Alpha rays are positively charged consist of two
protons and two neutrons and produced energy through excitation.
2. Non-particulate radiation- They consist of X- rays and gamma rays and are
high energy radiation. X rays and gamma rays are produced by X rays tubes
and radioactive isotope decay of 14C, 60Co respectively. They cause ionization
and excitation and also produce free radicals.
Non-ionizing radiation- UV rays are produced using mercury vapour lamps. They
are absorbed by purines and pyrimidines and forms purine-dimers, pyrimidine-
350

dimers and pyrimidine-hydrates. Mutagenic effect of UV rays is increased with the

presence of large amount of oxygen.
Radiation mostly affects DNA and causes change in the bases such as loss
of base, breakage of hydrogen bonds in DNA, deamination and double or single
stranded breaks in the DNA structure.

Mechanism of Chemical Mutagens

Alkylating agents adds alkyl groups to the nucleotide and causes point mutation.
The effect of alkylation depends of the position and and type of alkyl group added
to the nucleotide. They form cross link between the two strands of DNA and
blocks replication. Base analogues on the other hand get incorporated during DNA
replication as they are similar to that of the bases of DNA. Acridine dyes causes
point mutation by increasing the distance between adjacent base pairs.

Mutation Breeding
Mutation breeding, also known as “variation breeding,” is the strategy of treating
seeds to radiation, chemicals, and or enzyme in order to generate mutations with
desirable traits which is developed as a new genotype. Depending on whether a
plant species is reproduced vegetatively or by seed, the naming of populations is
based on this information. The M0 population, sometimes referred to as generation
zero, refers to seeds before mutation treatment; the M1 generation refers to the seed
batch after mutation treatment. The cells that make up this M1 are chimeric with
cells from a tissue that have various mutations. The outcome is a plant with several
tissues containing various alleles, making it challenging to impossible to measure
heritable phenotypic variations. The M2 population, which is the offspring of the
M1, is the first generation in which it is feasible to screen for mutations. Therefore,
in the M2, single plants can be phenotypically screened and chosen. Following
M3 families may also be the subject of selection; this has the benefit that it allows
for an evaluation of the segregation of mutant features within the family and the
determination of whether the family is homozygous for the mutant allele. M4 and
other succeeding generations may likewise be subject to selection. The detection
of uncommon phenotypes caused by particular mutant combinations can be done
with the help of later generations. Usually, self-pollinating plants like barley, rice,
and wheat use this selection method. Crops that are propagated vegetatively need a
slightly different nomenclature. The beginning material is still known as M0, but after
being subjected to mutagenic treatment, it is known as M1V1. This population carries
chimaeras, similar to the M1 of seed crops, which makes it difficult to determine
the heritability of phenotypes. Multiple rounds of vegetative propagation, such as
 351

M1V2, M1V3, up to M1Vn, are frequently used to address these problems. According
to recent research, chimaera breakdown in bananas can happen quickly and may
reduce the number of propagation cycles ( Jankowicz-Cieslaket. al.,. 2012). Different
classifications for the types of mutations are given by different scientists. Three major
categories are provided by the most recent classification established by Lundqvistet.
al.,. (2012): Chromosome, genome, and gene mutation. The number of genomes
(ploidy) and the number of chromosomes (aneuploidy) within a genome can both
alter as a result of genome mutations. Chromosome deficits, extra chromosomes,
chromosome replacements, and chromosome rearrangements are all examples of
neuploids. They can be created by breeding, particularly in combinations when
the parents each contribute a different amount of chromosomes or genomes. The
random loss of chromosomes caused by radiation can result in aneuploidy. Single
nucleotide modifications or tiny indels are examples of the minor changes that
frequently occur in gene mutations. Such changes could produce a new allele and
be functional. They are therefore intriguing for plant breeding because they offer
beneficial novel diversity ( Badoet. al.,., 2015).

Breeding for Oligogenes

In M1 a mutagen is applied to seedlings before they are planted. Typically, the
quantity of treated seeds is adjusted to produce 500 viable M1 plants at harvest.
Outcrossing should be avoided at all costs; this can be done by planting the M1
population separately or by bagging the blossoms of M1 plants or even the entire M1
population. Because of the heterozygous mutations, M1 plants will be chimaeras. To
grow the M2 progeny rows, 20 to 25 seeds are independently extracted from each
M1 spike. In M2The number of progeny rows grown is about 2,000. The M2 rows are
carefully and frequently observed. However, as the data are based on single plants,
only distinctive mutations are found in M2. To grow unique plant progenies in M3,
all the plants in M2 rows believed to have novel mutations are harvested separately.
If the mutant is unique, it is chosen for testing and subsequent multiplication. The
majority of the modifications, in the meantime, won’t help crops in any way. Only
1-3% of M2 rows might be anticipated to have advantageous mutations. Additionally,
M2 can be cultivated in large quantities through the addition of one or more seeds
from each M1 spike, fruit, or branch. After that, individual plants are chosen in M2,
and their offspring are cultivated in M3. In M3, progeny rows from specifically chosen
plants are raised. Rows that are poor or substandard are omitted. If the mutated
offspring are homogeneous, it is possible to bulk up two or more M3 progenies with
the same mutation. Bulk harvests of mutant M3 rows are made for an initial yield
test in M4. With a satisfactory check, a preliminary yield experiment is carried out
in M4 and promising mutant lines are chosen for replicated multilocation testing.
352

Replicated multilocation yield studies are carried out in M5-M7. The exceptional
line might be made available as a new variety. However, it is best to keep the low-
yielding mutant lines for use in hybridization studies.

Breeding for Polygenes

The same techniques used to grow M1 and M2 for oligogenic characteristics are used
here. M2 plants that are healthy, productive, and appear normal are chosen, and their
seeds are extracted separately to grow individual plant offspring rows in M3. From
specifically chosen individual plants, progeny rows are raised. M3 rows are carefully
scrutinised for any slight phenotypic divergences from the parent variety. Inferior
rows are eliminated. Few rows might be uniform and would be collected in large
quantities. Selection is carried out in M3 rows with segregation; segregation would be
present in most M3 rows. Identification of mutants with changed quantitative features,
such as partial or horizontal disease resistance, is made possible by a thorough and
in-depth analysis of a large number of M3 progeny rows. Such mutants have high
frequencies that are close to 1% or even higher, making it relatively inexpensive to
isolate them. In M4 In a preliminary yield trial, bulk seed from homogenous M3 rows
may be planted with a proper check; superior progenies are chosen for replicated
multi-location yield trials. Critical observation is made of specific M3 plant offspring.
Only if the progeny exhibits segregation and is promising may it be subjected to
selection. For initial yield experiments in M5, superior homogenous progenies are
harvested in large quantities. In M5-M8 In accordance with the stage at which the
progenies become homogeneous, preliminary yield trials and/or multi-location trials
are done. The best offspring may be released as new varieties.

Achievements in Mutation Breeding

There are numerous instances of new and useful character changes in plants that
have greatly boosted the production potential of particular crops over the roughly
80-year history of induced mutations. The main goal of mutation breeding is to
boost the prevalence of viable mutations, as well as the frequency and spectrum of
mutations (Wani et. al. 2011, Khan and Siddiqui 1996). Improvements in cereals
and legumes, which are significant food crops, have long been a top priority for
plant breeders. In China, the first rice varieties, KT 20-74 and SH 30-21, which
were created through induced mutation, were introduced in 1957. The first variety,
Yenhsing-1, was created through a cross-breeding technique using a mutant. Japan
witnessed the release of the semi-dwarf mutant Reimei, whose lodging resistance
greatly enhanced yield. Semi-dwarf rice cultivars with short and stiff straws, such as
Calrose 76 and Basmati 370 have changed the production of rice in the USA and
Pakistan, respectively. A new variety known as “Kashmir Basmati” was developed in
 353

Pakistan from an induced mutation in Basmati 370. It grows earlier, can withstand
cold temperatures, and keeps the parent variety’s aroma and cooking qualities.
‘PNR- 381’ and ‘PNR- 102’, two early ripening and aromatic mutation-derived
rice varieties were popular for cultivation in Haryana and Uttar Pradesh. Over
the course of ten years, a rice mutant known as ‘Zhefu 802’ was grown on more
than 10.6 million acres in China. In Thailand, gamma ray irradiation hastened the
emergence of the aromatic indica rice variety ‘RD6’ in 1977. It was grown widely on
2.4 million acres in 1994-95 (Futsuhara, 1968; Awan, 1991; Kharkwal and Shu,2009).
Centenario, a high yielding, high protein content, early maturity, and yellow rust
resistant barley cultivar developed in 2006, contributes greatly to the country’s food
security. The gamma ray generated mutant ‘Luther’ showed a 20% increase in yield,
better tillering, and lodging resistance, whereas ‘Pennrad’ had winter hardiness,
greater lodging resistance, and early ripening. The ‘TG 26’ mutant peanut variety
produced at the Bhabha Atomic Research Centre in Bombay has a semi-dwarf
plant habit, early maturity, compact pod setting, increased pod bearing, a higher
harvest index and field tolerance to main diseases(Kale et. al.,1997). Pusa - 408
(Ajay), Pusa - 413 (Atul), Pusa - 417 (Girnar), and Pusa - 547 are High Yielding
and Wilt Disease Resistant Chickpea Mutants produced at I.A.R.I., New Delhi, are
based on the direct application of induced micro-mutants in a legume crop in the
globe. Pusa - 547, a mutant variety produced in 2006, features thin testa, appealing
bold seeds, superior cooking quality, and high yield performance under late seeded
conditions in India’s North-Western area ( Kharkwal et al,2005; Kozgar& Khan,
2009). According to the FAO/IAEA mutant variety database, nearly all of the 465
mutations developed in vegetatively propagated plants were in floricultural plants.
These comprised chrysanthemum, dahlia, bougainvillaea, rose, carnation, and others.
Mutants were also developed in various fruit crops like apple, pear, banana, etc. to
increase resistance against diseases and insect pests.

Future Prospects
A new cultivar or variation may only be created from wild species using the traditional
breeding approach after several years. To enhance crops’ quality and quantitative yield
features, induced mutagenesis and its breeding methods are powerful tools that are
widely utilised in agriculture.To generate desirable agronomical features, high yield,
stress tolerance qualities, and resistance ability in different crops, mutagenic induction
is much simpler and less expensive to apply to crops. With the advancement of
breeding practises, genomics, molecular manipulations, and genetic engineering, the
development of genetically new germplasm becomes more viable. For agricultural
enhancement, it would be crucial to use new mutation-related technologies efficiently
and with skilled labour.The mutant plant species may be quickly chosen using certain
354

standard, easy screening methods, PCR, and non-PCR based procedures. It should
thus be used on a variety of crops. Mutated genes may now be identified, pyramided
into a single elite breeding line, and tracked in subsequent breeding programmes.
There are a lot of opportunities in the genomics age, especially given the importance
of conventionally produced mutations in functional genomics (Roychowdhury and
Tah 2013).

Conclusion
To begin a crop improvement programme, the availability of genetic diversity is
crucial. Plant breeding programmes that includes induced mutagenesis has proven
to be effective and reliable. One of the most effective breeding techniques for
generating novel genetic diversity and quickening the trait selection process is
induced mutagenesis. Mutation breeding has produced several mutants with enhanced
agro-economic features of various plant species during the past few decades. The
method has been effectively used in the production of feed crops, medicinal plants,
horticultural plants, and cereals. These mutant cultivars with specific characteristics/
traits like as high yield, tolerance to biotic and abiotic stresses, have been grown
globally, delivering a substantial positive economic effect and contributing to
global food and nutritional security. Despite the abundant mutant resources, issues
remain in feeding an ever-increasing population. To increase agricultural output,
mutant resources for various crop plants must be developed, which may be utilised
to generate novel mutant cultivars that are high yielding and resistant to biotic and
abiotic stresses.

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