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Application of mutation in crop improvement

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 International Journal of Research in Agronomy 2021; 4(2): 01-08

E-ISSN: 2618-0618
P-ISSN: 2618-060X
© Agronomy Application of mutation in crop improvement
[Link]
2021; 4(2): 01-08
Received: 02-04-2021 Temesgen Begna
Accepted: 03-05-2021

Temesgen Begna Abstract

Ethiopian Institute of Agricultural Genetic variation is a source of phenotypic diversity and is a major driver of evolutionary diversification.
Research, Chiro National Sorghum Heritable variation was observed and used thousands of years ago in the domestication of plants and
Research and Training Center, animals. The mechanisms that govern the inheritance of traits were later described by Mendel. Plant
Chiro, Ethiopia breeding requires genetic variation of useful traits for crop improvement. The induction of mutations has
been used to enhance the yield, better nutritional quality and wider adaptability of world’s most important
crops such as wheat, rice, pulses, millets and oilseeds. The total area covered by commercially released
mutant cultivars clearly indicates that they have played a significant role in solving food and nutritional
security problems in many countries. Of all the mutant varieties developed, majority of mutants were
produced through direct mutagenesis of the plant propagules and also there are several reports of mutants
derived by irradiating rooted stem cuttings, which paves the way for in vitro mutagenesis. The
incorporation of desired traits from non-adapted landraces or crop wild resources can speed up crop
improvement. Among the different strategies to enhance crop improvement programs, induced mutagenesis
has contributed immensely by creating mutant varieties with improved and desirable genetic changes in
agronomic ally important traits of the crop plants. Such genetic changes can occur spontaneously naturally
at a very low rate or experimentally induced by physical and chemical mutagens. Conventional mutation
techniques have often been used to improve yield, quality, and disease and pest resistance in crops or to
increase the attractiveness of flowers and ornamental plants. In general, mutation is the main source of
genetic variation, which is the raw material for evolution by natural selection. Recognize that mutations are
the basis of microevolution and that adaptations enhance the survival and reproduction of individuals in a
population. Mutation breeding has greater impact in sustainable crop production by developing new mutant
varieties. With the advances in genomics research and availability of genome sequences, induced mutants
continue to be a genetic resource for elucidating genetic mechanisms and metabolic pathways. Agricultural
sustainability and food security are major challenges facing continued population growth. Integration of
existing and new technologies for the induction and exploitation of genetic diversity towards developing
healthier, nutritious and productive crops is the need of the hour. Mutagenesis is a proven technology for
the development of improved or novel varieties with desirable traits. Several mutant genes have been
successfully explored, either directly or indirectly, to complement crop productivity.

Keywords: Mutation, genetic variation, induced mutation, mutagenesis, transgenic crop

Introduction
Mutation is the alteration in the genetic material of a cell of a living organism or of a virus that
is more or less permanent and that can be transmitted to the cell’s or the virus’s descendants.
The genomes of organisms are all composed of DNA whereas viral genomes can be of DNA
or RNA. Mutation refers to sudden heritable change in the phenotype of an individual. The
term mutation was originally coined by Dutch botanist Hugo De Vries since1848–1935 (Jain,
2010a) [4]. Mutation breeding is defined as the deliberate induction and development of mutant
lines for crop improvement. Mutation is the most commonly used method in asexually
propagated crops and self-pollinated crops for creation of variation that used for improvement.
Induced mutations are considered as an alternative to naturally occurring variation as the source
of germplasm for plant improvement programs and as an alternative to hybridization and
Corresponding Author:
recombination in plant breeding.
Temesgen Begna Mutation is a heritable change in a genetic characteristic of an organism and is a natural process
Ethiopian Institute of Agricultural that creates new variants (alleles) of genes. Mutation is the primary source of all genetic
Research, Chiro National Sorghum variations existing in any organism, including plants. Variation so created by mutation provides
Research and Training Center, the raw material for natural selection and is a driving force in evolution. The standard technique
Chiro, Ethiopia
of creating variability by means of altering genes through induction of mutations by physical or

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chemical mutagens and using the same effectively through depends to a large measure on the degree of genetic divergence.
elaborate methods of selection techniques in various generations Genetic diversity is a key factor for crop improvement.
for improvement of a particular crop species for desired Mutation is the origin of all new genetic diversity, occurring
objectives is called mutation breeding. This is frequently when there are occasional errors in the replication of DNA or
practiced by plant breeders all over the world for crop other elements of the production and packaging of genetic
improvement. information within the cells. Although it implies something
The genetic improvement of crops is a crucial component of the negative, mutations can have positive, neutral, or deleterious
efforts to address pressures on global food security and nutrition impacts. Mutations occur rather slowly but continuously.
(Ronald, 2011) [16]. It is estimated that food production should be Mutations at one level, for example, in the nucleotides that are
at least doubled by the year 2050 in order to meet the needs of a the basis of DNA may not all be expressed at other levels such
continually growing population (Ray et al., 2013) [15]. The as protein differences or observable changes in the appearance
availability of heritable variation is a prerequisite for genetic of a plant. The rate of mutation is useful in determining
improvement of crops. Where sufficient variation does not exist evolutionary relationships. Mutation breeding is a fundamental
naturally, it can be created through either random or targeted and highly successful tool in the global efforts of agriculture to
processes. feed an ever increasing and nutritionally demanding human
Globally, the current human population is increasing day to day population. Plant mutation breeding is the process of exposing
and expected to reach 9 billion by 2050 and that will lead to plant seeds, cuttings or cell cultures to radiation, such as gamma
food scarcity on earth. To overcome this increasing demand for rays and then planting the seed or cultivating the irradiated
food and proper nourishment, an improvement in food material in a sterile medium that generates a plantlet.
production is urgently needed (Ronald, 2014) [17]. The envisaged Mutations are the major source of genetic variability and
in food production is daunting because of limited available artificial mutations can be induced by mutagens (Oladosu et al.,
arable land, depleting water resource and varying climatic 2016) [13]. There are three major tools for mutagenesis:
condition. There are different mechanisms for harnessing the biological agents such as transposons, retrotransposons, and T-
heritable variations encoded in the genetic makeup of existing DNA; physical agents such as ionizing radiations and chemical
crop plants so as to use them in the crop improvement programs. agents such as alkylating agents; and azides (Serrat et al., 2014)
[19]
The incorporation of desired traits from non-adapted landraces . Mutation breeding has resulted in thousands of improved
or crop wild resources can speed up crop improvement. Putative varieties with higher yields and improved tolerance to pests,
parental material can also be induced to mutate so as to obtain diseases and environmental stresses. The genetic diversity of
new genes that control desired traits for new crop variety crop plants is the foundation for the sustainable development of
development (Suprasanna et al., 2009) [23]. improved crop varieties for meeting present and future food
Among the different strategies to enhance crop improvement security challenges. Induced mutations offer numerous benefits
programs, induced mutagenesis has contributed immensely by to crop improvement, especially when conventional breeding
creating mutant varieties with improved and desirable genetic techniques fail for the lack of appropriate genetic variation.
changes in agronomically important traits of the crop plants. Sudden, heritable changes in the genetic material, DNA, are
Mutagenesis has become more efficient in combination with known as mutations. Selection of naturally occurring mutations
advanced molecular biology techniques and in vitro culture in wild, ancestral species helped humans in the domestication
methods that result in enhancement of crop improvement or and further improvement of today's crop plants. Plant breeding
breeding program particularly under the global climate change based on the science of genetics, as practiced over the past 100
(Jain, 2010a) [10]. Such induced mutagenesis also helps in the years, exploited the available genetic variability in the primary
mining of new gene alleles that do not occur in the germplasm gene pool of crop plants and sometimes in related species. This
(Roychowdhury and Tah, 2013) [18]. Mutation is the ultimate approach enlarged the yield potential of crops several fold. It
source of all genetic changes which provide the raw material for also a) improved the stability of yield by incorporating
evolution and it is a valuable approach for improvement of resistance to various biotic and abiotic stresses; b) improved
economic characters of plants. Such genetic changes can occur quality of the produce; and c) altered the adaptability of crop
spontaneously naturally at a very low rate or experimentally species, providing opportunities to grow new crops for food
induced by physical and chemical mutagens (Mba et al., 2007). security outside their traditional range. Genetically improved
The use of diverse genetic resources is important for breeding seed (other planting material) is the most significant input for
crop varieties (Glaszmann et al., 2010) [3]. developing sustainable cropping systems for food security and
Crop species with narrow genetic diversity are susceptible to economic growth. Half of the increased productivity of today's
emerging pathogens or other constraints leading to loss of crop plants comes from genetic improvements. The other half is
productivity and this may lead to a serious decline in the areas of contributed by inputs and management practices.
adaptation (Dyer et al., 2014) [1]. Genetic variability of crops can A mutation is the original source of genetic variation and
be partitioned into variability between crop cultivars and primarily creates genetic diversity and can have positive, neutral
variability within a crop cultivar (the genetic differences within and negative impacts in genetic alteration of crop species.
the population of plants that make up the cultivar). The extent of Mutation is the sudden heritable changes of genetic diversity
variation between cultivars is a function of the number that occurred occasionally through aberration of genetic materials
makes up a significant part of the crop acreage and of the like DNA, RNA and protein within the cells. Mutation has a
genetic differences between them. The largest diversity of the great role in increasing genetic diversity in order to feed the
crop germplasm provides greater opportunities for improvement increasing human population (Smith BD, 1989) [22]. Mutation is
regarding its environmental adaptability and acquiring better primarily causes the alteration of genetic diversity. Genetic
agronomic traits from the crop species. Genetic diversity of diversity is the variation occurred in genetic information, which
plants determines their potential for improved efficiency and depends on frequency and diversity of alleles among individuals
hence their use for breeding, which eventually may result in within a population or a species. Mutation is the driving force
enhanced food production. The success of breeding program for sustainable genetic diversity creation which uses in further

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improvement. Induced mutageneses are broadening the genetic Mutation breeding and plant mutagenesis play a significant role
variation whereas a conventional breeding approach is in increasing the genetic variability for desired traits in various
narrowing genetic variability for improvement for long period of food crops (Kozgar, M.I et al., 2012) [7]. Induced mutagenesis is
time. The utilization of mutation breeding is significantly one of the most efficient tools used for the identification of key
generating crop genetic diversities through improvement of regulatory genes and molecular mechanisms. It is a promising
crops to improve the livelihood of the communities (Oladosu Y approach to develop new varieties with improved agronomic
et al., 2016) [13]. The objective of the paper was to understand characteristics, such as higher stress tolerance potential (biotic
the significance mutation in crop improvement to feed the world and abiotic stress) and bio-fortification. Additionally, various
growing population. mutagenesis approaches have been used to study the
evolutionary relationship as well as for the genetic improvement
2. Role of Mutation in Crop Improvement of many organisms, including microbes, animals, and plants
It is well known that evolution and practical breeding depend on (Sikora, P et al., 2011) [21]. Technological advances in molecular
genetic variation. The variations that are found in nature do not biology have re-augmented the mutation breeding making it
represent the original spectra of spontaneous mutations. Rather, more efficient than ever thought before.
they are the result of genotypes recombining within populations The first step in plant breeding is to identify suitable genotypes
and their continuous interaction with environmental factors containing the desired genes among existing varieties, or to
(Novak FJ and Brunner H, 1992) [12]. Green plants are essential create one if it is not found in nature. In nature, variation occurs
for human existence as a source of food, clothing and energy mainly as a result of mutations and without it and plant breeding
resources. Mutations are the primary source of all genetic would be impossible without variation. In this context, the major
variations existing in any organism, including plants aim in mutation-based breeding is to develop and improve well-
(Kharkwal MC, 2012) [6]. The resulting variation provides the adapted plant varieties by modifying one or two major traits to
raw material for natural selection and is also a driving force in increase their productivity or quality. Both physical and
evolution. Spontaneous mutations are very rare and random in chemical mutagenesis is used in inducing mutations in seeds and
terms of time of occurrence, which makes them more difficult to other planting materials. Then, selection for agronomic traits is
use in plant breeding programmes (Lonnig WE, 2005) [9]. In this done in the first generation, whereby most mutant lines may be
way, mutant forms showing both large and small effects on the discarded. Mutation breeding sometimes referred to as
phenotype arise for all kinds of traits (Kharkwal MC, 2012) [6]. variation breeding.
Mutation breeding involves the development of new varieties by The utilization of induced mutants in crop improvement is called
generating and utilizing genetic variability through chemical and mutation breeding. Mutation may be the change in gene,
physical mutagenesis. It is now a pillar of modern plant chromosome or plasma gene (genetic material inside
breeding, along with recombinant breeding and transgenic mitochondria and chloroplasts). The mutation produced by
breeding (Shu QY et al., 2012) [2]. change in the base sequence of gene is called point mutation or
Mutagenesis is the process whereby sudden heritable changes gene mutation. The concept of induced mutagenesis for crop
occur in the genetic information of an organism not caused by improvement developed dated back to the beginning of 20th
genetic segregation or genetic recombination, but induced by century. During the past 80 years, mutation breeding has been
chemical, physical or biological agents (Roychowdhury R successfully utilized for the improvement of crops as well as to
and Tah J, 2013) [18]. Mutation breeding employs three types of supplement the efforts made using traditional methods of plant
mutagenesis. These are induced mutagenesis, in which breeding. There are two major types of Mutation:
mutations occur as a result of irradiation (gamma rays, X-rays,
ion beam, etc.) or treatment with chemical mutagens; site- 2.1 Spontaneous Mutation
directed mutagenesis, which is the process of creating a Spontaneous mutations arise from a variety of sources including
mutation at a defined site in a DNA molecule; and insertion errors in DNA replication, spontaneous lesions and transposable
mutagenesis, which is due to DNA insertions, either through genetic elements. Mutation occurs in nature are called
genetic transformation and insertion of T-DNA or activation of spontaneous mutation. Spontaneous mutation occurs in the
transposable elements (Forster BP and Shu QY, 2012) [20]. Plant organism without any treatment at low rate in the nature. The
breeding requires genetic variation of useful traits for crop frequency of spontaneous mutation is 10-6 (one in one in
improvements (Novak FJ and Brunner H, 1992) [12]. However, million). Different genes in an organism show different mutation
multiple mutant alleles are the sources of genetic diversity for rate. Rates of spontaneous mutation per genome as measured in
crop breeding as well as functional analysis of the targeted gene the laboratory are remarkably similar within broad groups of
in many cases. The key point in mutation breeding is the process organisms but differ strikingly among groups. Mutation is one of
of identifying individuals with a target mutation, which involves the basic phenomena of life. Without mutation, the gradual
two major steps: mutant screening and mutant confirmation development of life from inorganic material would have been
(Forster BP and Shu QY, 2012) [20]. Mutant screening is a impossible and the evolution of living beings from the first
process involving selection of individuals from a large mutated groups of molecules in which a primitive, information-carrying
population that meet specific selection criteria, e.g. early unit cooperated with an energy gaining device up to the present
flowering, disease resistance as compared to the parent. diversity of highly refined living organisms could not have
However, these selections are often regarded as putative mutants occurred. Spontaneous mutations are alleles of initially unknown
or false mutants. Mutant confirmation, on the other hand, is the genes and are given allele names and symbols based on their
process of re-evaluating the putative mutants under a controlled phenotype. Spontaneous mutations and deletions can be used to
and replicated environment using large samples. Through this identify important functional regions within a gene. Spontaneous
process, many putative mutants are revealed to be false mutants. mutations are the result of errors in natural biological processes,
In general, the mutations that are important in crop improvement while induced mutations are due to agents in the environment
usually involve single bases and may or may not affect protein that cause changes in DNA structure.
synthesis (Mba C, 2013).

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2.2 Induced Mutation male sterility, production of haploids, creation of genetic

Induced mutations are very effective in creating genetic variability and overcoming self-incompatibility.
variability for various economic characters in crop plants. Mutations are induced by physical (gamma radiation, high and
Induced mutations have been used for increasing the range of low energy beams) and chemical (ethyl methane sulfonate,
genetic variability in barley, oats, wheat and many other crops. EMS) mutagen treatment of both seed and vegetative propagated
In asexually propagated crops like sugarcane and potato, somatic crops. The mechanism of mutation induction is that the mutagen
mutations may be useful, because the mutant plant can be treatment breaks the nuclear DNA and during the process of
multiplied as a clone. Mutations induced in an organism by DNA repair mechanism, new mutations occur randomly and are
treatment with physical or chemical mutagen are called induced heritable. The changes can also occur in cytoplasmic organelles
mutations. The agents which are used to induce mutation are and also results in chlorophyll mutations, chromosomal or
called mutagens. Certain genes in an organism promote the genomic mutations that enable plant breeders to select useful
mutation of other genes nearby in the chromosomes. Induced mutants such as abiotic and biotic stresses and others resistant
mutations are useful in crop improvement in several principal trait (Mohan Jain, S and Suprasanna, P, 2011) [11].
ways, viz: Development of improved varieties, induction of

Table 1: Classification and Brief Description of Mutation

Basis of Classification and Type of Mutation Brief Description
1. Based on source
Spontaneous Mutation that occur in nature
Induced Mutation which are produced by the use of mutagenic agent
2. Based on direction
Forward mutation Any change from wild type allele
Reverse mutation Any change from mutant allele to wild type
3. Based on tissue
Somatic mutation A mutation in somatic tissue
Germinal mutation A mutation in germ line cell
4. Based on survival
Lethal A mutation which kills the individual that carries it
Sub-lethal When mortality is more than 50% of the individuals that carry mutation
Sub-vital When mortality is less than 50% of the individuals that carry mutation
Vital When all mutant individuals survive
5. Based on site
Nuclear mutation A mutation in nuclear gene
Cytoplasm mutation A mutation in cytoplasm gene
6. Based on character
Morphological A mutation that alters the morphological characters of individuals
Biochemical A mutation that alters biochemical function of individuals
7. Based on visibility
Micro-mutation Mutation with invisible phenotypic changes. Generally observed in quantitative characters
Mutation with distinct morphological changes in phenotypes. Generally found in qualitative
Macro-mutation
characters

Source: Joint Division of the Food and Agriculture Organization of the United Nations and the International Atomic Energy Agency
Fig 1: Distribution of mutant crop varieties by continents (Accessed on 15th July, 2015).

2.3 Mutagenic Agents 2.3.1 Physical Mutagenesis

Agents of artificial mutations are called mutagens. They are In the past 80 years, physical mutagens, mostly ionizing
generally grouped into two broad categories, namely chemical radiations, have been used widely for inducing hereditary
mutagens and physical mutagens aberrations and more than 70% of mutant varieties were
developed using physical mutagenesis. Radiation is defined as

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energy travelling through a distance in the form of waves or a complementary tool in plant breeding. Gamma rays have a
particles. These are relatively high-energy levels of shorter wave length and therefore, possess more energy than
electromagnetic (EM) spectrum that are capable of dislodging protons and X-rays, which gives them ability to penetrate deeper
electrons from the nuclear orbits of the atoms that they impact into the tissue. Neutrons are hazardous and hence have less
upon. The impacted atoms, therefore, become ions. Hence, the penetrating abilities, but they are known to cause serious
term ionizing radiation. These ionizing components of the EM damage to the chromosomes. They are best used for materials,
include cosmic, gamma (γ) and X-rays. The most commonly such as dry seeds. Various forms of neutrons were also studied
used physical mutagens are shown in Table 1. X-rays were the extensively for their use in mutagenesis in the 1960s and 1970s.
first to be used to induce mutations. Since then, various Though it has been proved to be an effective mutagen,
subatomic particles (neutrons, protons, beta particles and alpha particularly for producing large DNA fragment deletions, the
particles) have been generated using nuclear reactors Wiley- application of neutrons in induced mutagenesis is limited. The
Blackwell, (2006). Gamma radiation from radioactive cobalt mutagenic effect of ultraviolet light was discovered by
(60Co) is widely used. It has high penetrating potential and is Altenburg through irradiation of the polar cap cells of fruit fly
hazardous. However, it can be used for irradiating whole plants eggs. The mutagenic potential of these rays have since been
and delicate materials, such as pollen grains. Various mutants confirmed in many organisms. In those organisms, germ tissue
have been developed through gamma radiation. The mutagenic could be easily exposed to the low-penetrating ultraviolet light
effect results mostly from DNA double-strand breaks. which resulted in covalent dimerization of adjacent pyrimidine
The mutants show higher potential for improving plant (Oladosu, Y et al., 2015) [14]
architecture leading to better crop improvement and are used as

Table 2: Examples of commonly used physical mutagens

Mutagen Source Characteristics Hazard
Dangerous,
X-rays X-ray machine Electromagnetic radiation; penetrates tissues from a few millimeters to many centimeters
penetrating
Gamma Radioisotopes and Electromagnetic radiation produced by radioisotopes and nuclear reactors; very Dangerous, very
rays nuclear reaction penetrating into tissues; sources are 60Co (Cobalt-60) and 137Cs (Caesium-137) penetrating
Nuclear reactors or There are different types (fast, slow, thermal); produced in nuclear reactors; uncharged
Neutrons Very hazardous
accelerators particles; penetrate tissues to many centimeters; source is 235U
Beta Radioactive isotopes or Produced in particle accelerators or from radioisotopes; are electrons; ionize; shallowly
May be dangerous
particles accelerators penetrating; sources include 32P and 14C
Alpha Derived from radioisotopes; a helium nucleus capable of heavy ionization; very shallowly
Radioisotopes Very dangerous
particles penetrating
Nuclear reactors or Produced in nuclear reactors and accelerators; derived from hydrogen nucleus; penetrate
Protons Very dangerous
accelerators tissues up to several centimeters
Produced positively charged ions are accelerated at a high speed (around 20%–80% of the
Ion beam Particle accelerators Dangerous
speed of light) deposit high energy on a target
Source: Oladosu, Yusuff et al. (2015) [14]

2.3.2 Chemical mutagenesis epoxides, ethylene imines, ethylene imides, alkyl

The effect of chemical mutagens on plant materials is generally methanesulfonate, alkylnitrosoureas, alkylnitrosoamines,
considered milder. An advantage of chemical mutagenic agents alkylnitrosoamides, alkyl halides, alkyl sulphates, alkyl
is that they can be applied without complicated equipment or phosphates, chloroethyl sulphides, chloroethylamines, and
facilities. The ratio of mutational to undesirable modifications is diazoalkanes. One of the most effective chemical mutagenic
generally higher for chemical mutagens than for physical. groups is the group of alkylating agents (these react with the
Usually, the material is soaked in a solution of the mutagen to DNA by alkylating the phosphate groups as well as the purines
induce mutations. However, chemical mutagens are generally and pyrimidine. Another group is that of the base analogues
carcinogenic and therefore, extra care must be taken for health (they are closely related to the DNA bases and can be wrongly
protection during the procedure. Material and safety data sheets incorporated during replication.
for the specific chemical mutagen chosen should be carefully
read and the agent should be appropriately inactivated before 2.4 Genetic Transformation of Plants (T-DNA) or Insertion
disposal. Despite the large number of mutagenic compounds, Mutation
only a small number has been tested in plants. For more than two decades, scientists have used Agrobacterium-
Among them, only a very restricted group of alkylating agents mediated genetic transformation to generate transgenic plants.
has found large application in plant experimental mutagenesis An initial technology to introduce genes of interest (goi) into
and plant mutation breeding. Over 80% of the registered new Agrobacterium seems complex microbial genetic methodologies
mutant plant varieties reported in the International Atomic that inserted this gene of interest into the transfer DNA (T-
Energy Association (IAEA) database obtained via chemical DNA) region of large tumor-inducing plasmids (Ti-plasmids).
mutagenesis were induced by alkylating agents. Of these, three However, scientists eventually learned that T-DNA transfer
compounds are significant: ethyl methane sulphonate (EMS), 1- could still be effected if the T-DNA region and the virulence
methyl-1-nitrosourea and 1-ethyl-1-nitrosourea, which account (vir) genes required for T-DNA processing and transfer was split
for 64% of these varieties. (Javed, I et al. 2016) [5] Alkylating into two replicons. This binary system permitted facile
agents can be found among a large array of classes of manipulation of Agrobacterium and opened up the field of plant
compounds, including Sulphur mustards, nitrogen mustards, genetic engineering to numerous laboratories (Lee, L.Y and

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Gelvin, S.B., 2008) [8]. Transfer requires three major elements: chromosome. In the presence of these sugars, vir genes are more
(1) T-DNA border repeat sequences (25 bp) that flank the T- fully induced at lower phenolic concentrations.
DNA in direct orientation and delineate the region that will be (2) Processing T-DNA from the parental Ti- or Ri-plasmid
processed from the Ti/Ri-plasmid (Yadav et al., 1982) [24]; (2) (virD1 and virD2). Together, VirD1 (a helicase) and VirD2 (an
vir genes located on the Ti/Ri-plasmid; and (3) various genes endonuclease) bind to and nick DNA at 25-bp directly repeated
(chromosomal virulence [chv] and other genes) located on the T-DNA border repeat sequences). The VirD2 protein covalently
bacterial chromosomes. These chromosomal genes generally are links to the 5′ end of the processed single-strand DNA (the T-
involved in bacterial exopolysaccharide synthesis, maturation, strand) and leads it out of the bacterium, into the plant cell, and
and secretion. to the plant nucleus. (3) Secreting T-DNA and Vir proteins from
The vir region consists of approximately 10 operons (depending the bacterium via a type IV secretion system (virB operon and
upon the Ti- or Ri-plasmid) that serve four major functions: 1) virD4). The Agrobacterium virB operon contains 11 genes, most
Sensing plant phenolic compounds and transducing this signal to of which form a pore through the bacterial membrane for the
induce expression of vir genes (virA and virG). VirA and VirG transfer of Vir proteins. Currently, we know of five such
compose a two-component system that responds to particular proteins that are secreted through this apparatus: VirD2
phenolic compounds produced by wounded plant cells. Because (unattached or attached to the T-strand), VirD5, VirE2, VirE3,
wounding is important for efficient plant transformation, and VirF VirD4 acts as a coupling factor to link VirD2-T-strand
Agrobacterium can sense a wounded potential host by to the type IV secretion apparatus.
perceiving these phenolic compounds. Activation of VirA by (4) Participating in events within the host cell involving T-DNA
these phenolic inducers initiates a phospho-relay, ultimately cytoplasmic trafficking, nuclear targeting, and integration into
resulting in phosphorylation and activation of the VirG protein. the host genome (virD2, virD5, virE2, virE3, and virF). VirD2
Activated VirG binds to the vir box sequences preceding each and VirE2 may play roles in targeting the T-strand to the
vir gene operon, allowing increased expression of each of these nucleus; In addition, VirE2 likely protects T-strands from
operons. In addition to induction of the vir genes by phenolics, nucleolytic degradation in the plant cell. VirF may play a role in
many sugars serve as co-inducers. These sugars are perceived by stripping proteins off the T-strand prior to T-DNA integration
a protein, ChvE, encoded by a gene on the Agrobacterium (Tzfira et al., 2004).

Source: Genetic Transformation of Plants manual notes

2.5 Impact of induced mutation 2.5.1 Economic impact of a new mutant variety
The Plant Breeding and Genetics Section of the Joint Area planted to the variety and percentage of the area under the
FAO/IAEA Division maintain a Mutant Variety Database crop in the region; increased yield, enhanced quality, reduced
(MVD) which lists more than 2250 cultivars of 175 species that use of pesticides and fungicides (e.g. in varieties resistant to
have been officially released in sixty countries. Among these, diseases and insect pests), savings in water (short duration of
1700 are in the food crops cereals (1072), legumes (311), growth and drought tolerance), increased land use through early
vegetables (66), and oil crops (59); the rest are industrial crops maturity to facilitate crop rotation, improved or intensified
and ornamental plants1. In fact, the MVD does not reveal the cropping systems with changed maturity or response to
full usage of the induced variability because it does not include photoperiod. improved processing quality and value of the
all the derivatives selected after cross breeding. The impact of products (oil, starch, malt, beer and whisky), quality preference
the new cultivars can only be estimated by the acreage in which by the consumer (new flower and foliage color in ornamentals,
they are grown. In the absence of well-organized seed skin and flesh color in root and tuber crops and fruit crops,
production in most developing countries, and given the aroma and glutinous nature in rice, and kernel color in wheat),
prevalence of informal seed sales between farmers, it is very increased nutritive value, highly sine and vitamins, increased oil,
difficult to estimate the real spread of a given derivative. To take shelf life, reduced toxins, increased yield of essential oils, new
one important example, rice is the staple food for nearly half of specialty and designer crops, ease of harvest, threshing, increase
the human population. More than 90% is grown in developing in export earnings. Reduction in imports, often, induced
countries mutations lead to more advantages than a simple desired
phenotypic change.

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Table 2: Economic impact of mutant varieties

Crop Country Mutant variety Basis of value assessment Value or area
Cereals
Rice Thailand RD6 and RD15 Total crop value at farm gate for the period 1989–98 US$ 16.9 billion
China Zhefu 802 Cumulative planted area between 1986–1994 10.6 million ha
Japan 18 varieties Total crop value in 1997 US$ 937 million
India PNR-102 and PNR-381 Annual crop value US$ 1,748 million
60–70% rice growing area in
Australia Amaroo Current annual planted area
Australia
Costa Rica Camago 8 Current annual planted are 30% rice growing area in Costa Rica
Vietnam TNDB100 and THDB Total planted area in 1999 220,000 ha
Myanmar Shwewartun Total planted area in 1993 800,000 ha
Jauhar 78, Soghat 90 and
Bread wheat Pakistan Additional income to farmers during 1991–99 US$ 87.1 million
Kiran 95
Durum wheat Italy Creso Additional incometofarmers during 1983–93 US$ 1.8 billion
Uk-Scot land
Golden promise Diamant
Barley Numerous Crop value (1977–2001) area planted in 1972 2.86 million ha
and derived varieties
European countries
Legumes
Chickpea Pakistan CM 88; CM 98 Additional annual income to the growers US$ 9.6 million
Black gram India, Maharashtra
TAU-1 Value of increased production in season 1998–1999 US$ 64.7 million
(urdbean) State
Oil and industrial crops
NIAB-78, Total value of Total value of crop from 1983–1993,Additional US$ 3 billion
Cotton Pakistan
crop from 1983– 1993 income to growers from 1983 onwards, US$ 486 million
Sunflower USA NuSun® Grown area in 1994 50,000 ha
Fruit trees
Japanese
Japan Gold Nijisseiki Additional annual income to growers US$ 30 million
pear
Grapefruit USA, Texas Rio Star Grown area (year 2000) 7,300 ha (75% of total area)
Source: B.S. Ahloowalia, M. Maluszynski∗ & K. Nichterlein (2004) FAO/IAEA

3. Summary and Conclusion source of genetic variation and primarily create genetic diversity
World agriculture sustainability is threatened by increasing and mutation can have positive, neutral and negative impacts in
human populations, reduced availability of cultivated land and genetic alteration of crop species. Mutation is the sudden
changing climate patterns. Plant mutation breeding is a major heritable changes of genetic diversity occurred occasionally
component in addressing these concerns in developing novel through aberration of genetic materials like DNA, RNA and
germplasm in a relatively short time. Mutation breeding is one protein within the cells. Mutation has a great role in increasing
of the breeding methods used in improving crop plants for genetic diversity in order to feed the increasing human
human benefit. Different types of mutation breeding were used population. Genetic diversity is the variation occurred in genetic
in the improvement of crop plants. Induced mutagenesis has information, which depends on frequency and diversity of alleles
played an important role by creating several mutants in different among individuals within a population or a species. Mutation is
crop plants. These mutant varieties with specific character or the driving force for sustainable genetic diversity creation which
trait such as high yield, resistance to biotic and abiotic stresses, uses in further improvement.
have been grown globally bringing a significant positive At present genetic variability is a narrowed using conventional
economic impact and contribute to global food and nutritional breeding approach for a long period of time whereas induced
security and improved livelihoods. Despite the available mutant mutagenesis is one of the most important approaches for
resources, challenges still lie ahead to feed an ever-increasing broadening the genetic variation and diversity in crops to
population. To speed up crop production, mutant resources for circumvent the bottleneck conditions. Induced mutagenesis can
different crop plants have to be established which can be used to contribute to unleashing the potentials of plant genetic resources
create new mutant cultivars which are high yielding, resistant to and thereby avail plant breeders the raw materials required to
biotic and abiotic stresses, enhanced uptake of specific metal, generate the smart crop varieties. Crop varieties generated
deeper rooting systems and modified oil, starch and protein through the exploitations of mutation breeding are significantly
content that can boost industrial processing. contributing to global food and nutritional security and
Mutations are the heritable change to an individual’s genetic improved livelihoods. Induced mutations are significant as novel
makeup and are one of the driving forces of evolution mutations are being isolated for enhanced nutrition quality of
complementing selection (natural and artificial) and crop plants. Another source of nutrition provision is from the
hybridization. Mutations can create evolutionary advantages or neglected and underutilized crops, and requires more attention
disadvantages leading to a preferential selection or deletion of together with the major crops for enhancing nutrition provision
genotypes from the gene pool. Induced mutagenesis is one of the to the ever-growing human population. Perhaps change of food
most efficient tools that has been utilized extensively to create habits would be required gradually move away from the
genetic variation as well as for identification of key regulatory consumption of major crops and start using underutilized crops
genes for economically important traits toward the crop either singly or in combination of both. Developing genetically
improvement. Mutations are one of the fundamental forces of novel germplasm with increased content of these together with
evolution because they fuel the variability in populations and other health benefit components becomes more feasible
thus enable evolutionary change. Mutations are the original concurrent with the enhancement of breeding techniques,

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