role of mutation in crop improvement

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Introduction: Mutation : A sudden and heritable change in the character of an organism, which does not arise due to segregation or recombination. Mutation Breeding : The genetic improvement of crop plants for various characters through the use of induced mutation. Some other terms: Mutation Changes in genes and chromosomes Mutated Altered genes Mutant New organism with a mutated gene or rearranged chromosomes

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Landmarks in Mutation Breeding Research Scientist Year Contribution Charles Darwin 1859 Recognized (in his book Origin of Species ) that mutant forms arose suddenly and spontaneously, but he was doubtful about their role in evolution. Hugo De Vries 1900 Credited with the discovery of mutations and its role in evolution of species. He studied the variant forms arising in the garden plant, evening primrose ( Oenothera lamarkiana ) and coined the term “Mutation.” H.J. Muller 1927 First to induce genetic mutations in fruit fly (Drosophila melanogaster ) with x-rays.

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Landmarks in Mutation Breeding Research Scientist Year Contribution L.J. Stadler 1928 Described the mutagenic effects of x-rays and gamma rays in barley and maize which became the foundation stone for mutation breeding in crop plants. J.A. Rapoport & others 1945 Discovered mutagenic activity in formaldehyde, diethylsulfate and other chemical compounds. C. Auerbach & J.M. Robson 1946 Found that mutation could be induced by nitrogen and sulfur mustard in Drosophila malanogaster .

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Landmarks in Mutation Breeding Research 1944 Barley mutant cultivar “ Jutta ” was the first cereal crop var. developed by x-rays and was released in Germany in 1955. 1948 “ M.A. 9” world’s first cotton mutant variety induced by x-rays released in India.

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Characteristics of mutation Mutations are – 1)Mostly recessive 2)Harmful 3)Random 4)Recurrent 5)Having Pleiotropic effects 6)Occurs with low frequency. Effects of mutation :- 1) Lethal 2) Sublethal 3) Subvital 4) Vital

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Classification of mutations Based on – Direction: 1)Forward 2)Reverse mutation Tissue: 1)Somatic 2) Germinal Sites: 1) Nuclear 2) Cytoplasmic mutation Character: 1)Morphological 2)Biochemical Visibility: 1)Macro-mutation 2)Micro-mutation Source: 1)Spontaneous 2)Induced mutation

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Induced mutation Mutation which are produced by the use of mutagenic agents(mutagens) Mutagens:- Agent which are used for induction of mutation are known as mutagens S.N. Type of radiation Main properties Mode of action or changes caused 1. X-rays S.I., penetrating and non-particulate. Induce mutations by forming free radicals and ions. Cause addition, deletion, transitions and transversions . 2. Gamma- rays S.I., very penetrating and non particulate. Induce mutations by ejecting atoms from the tissues. Cause addition, deletion, transitions and transversions . 3. Alpha-particles D.I., particulate, less penetrating and positively charged. Act by ionization and excitation. Cause chromosomal and gene mutations. 5. Fast & thermal neutrons D.I., particulate, neutral particles, highly penetrating. Cause chromosomal breakage and gene mutations. 6. Ultra violet rays Non- ionizing, low penetrating. Cause chromosomal breakage and gene mutations.

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Chemical agents Examples Mode of action or changes caused 1)Alkylating agents 2)Base analogues 3)Acridin dyes 4) Others EMS,EI, MMS,NM, etc. 5Bromo uracil, 2Amino purine. Acriflavin ,proflavin Nitrous acid, Hydroxylamine Sodium azide AT↔GC Transition Transition AT↔GC Transition Framshift mutation (addition, deletion of nucleotides) GC↔AT Transitions

Genetic structure changes:

Genetic structure changes Gene (point mutation) Chromosome Genome Gene (point) mutation → a change in specific sequence of nucleotides in DNA molecules leading to the formation of a new type of protein or preventing that of the normally protein → take place at the molecular or sub-microscopic level → Such change may be accompanied by the emergence of a new trait inherited in accordance with Mendel’s Laws

Chromosomal mutation:

Chromosomal mutation Mutation associated with splitting and subsequent changes in the structure of the chromosomes The end of the split chromosomes may fuse to form structure again, but the new chromosomes are not always exactly what the used to be The microscopic structures of chromosomes may be characterized by deletion or deficiency (loss of a chromosomal segment), duplication (doubling of a chromosomal segment), inversion (rearrangement of a group of genes in a chromosomal segment in a such a way that their order is reversed; rearrangement of genetic material in a chromosome results from loss of segment, its rotation by 180°, and fusion of the separated ends) and translocation (change in a position of a chromosome or more often exchange of segments between different chromosomes)

Genome mutation:

Genome mutation Changes in sets of chromosomes Remarks: Breeders are more interested in gene mutation, because chromosomal rearrangement usually produce negative results, such as lower fertility of the offspring Mutant are often of great value for breeding as sources of new, previously unknown useful characters Mutagenesis may be instrumental in obviating the technical difficulties arising in the crossing of such a small flowered crops such as milled

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Mutation breeding The utilization of induced mutation for crop improvement is known as mutation breeding. Steps in mutation breeding :- 1) Objective of programme 2) Selection of variety 3) Plant part treated 4) Dose of mutagen (LD 50) 5) Giving mutagen treatment: a) Irradiation - Immediately planted to raise M1 b) Chemical - Seeds are presoaked and exposed to desired mutagen. 6)Handling of mutagen treated population

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Procedure of mutation breeding The mutation breeding process consists of four important steps: ● Choice of material : The best adopted variety of a crop should be chosen for mutagenesis, because such variety has all acceptable agronomic features. ● Choice of mutagen : The choice of mutagen depends upon the plant parts to be treated. Generally, chemical mutagens are more preferred for seed treatment and radiation for treatment of vegetative parts. ● Mutagenic treatment : The procedure of mutagenic treatment takes three things into account : viz. plant species, dose of mutagen and duration of treatment. In seed propagated species, generally, seed are treated. In vegetatively propagated species, buds, suckers or cuttings are used for mutagenic treatment. LD50 refers to a dose of mutagen that kills 50% of the treated individuals. the duration of treatment depends on the intensity of radiations or concentration of chemical mutagens. ● Handling of treated material : The handling of mutagen treated material differs in seed propagated species and vegetatively propagated species.

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Applications 1) Induction of desirable mutant allele, which is not present in germplasm . 2) Improving specific character of variety without change in genetic make up. 3) Used to improve various quantitative character. 4) Induction of male sterility 5) Production of haploids by irradiating pollens. 6) Overcoming self incompatibility(Lewis-1954)

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Limitations 1) Low frequency 2) Need to screen large number of population. 3) Pleiotropic effect 4) Most of the mutations are recessive. 5) Identification of micro mutation is very difficult.

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Region No. of mutant cultivars % of total Asia 1582 57 Europe 896 32.29 N. America 182 6.56 Africa 60 2.16 L. America 45 1.62 Australia 10 0.37 Total 2775 100.0 Mutant cultivars released in various regions of the world Source: I.A.E.A.,2008

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Country No. of mutants cultivars Rank China 638 1 India 272 2 Japan 232 3 Russia +USSR 214 4 Netherlands 176 5 Germany 176 6 USA 128 7 Other countries include : France (39), Czech. Rep. (36), Canada (35), Italy (35), Pakistan (32), U.K. (32), Bulgaria (30) & Poland (30) Mutant cultivars released in top seven countries of the world Source : IAEA,2009

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Rediscovery of Induced Mutations Why? Major economic successes in USA (rice, barley, sunflower, grape fruit, peppermint: Pakistan (cotton); India (blackgram); Australia & Canada (linseed); Japan (pear); China (rice) Development in biotechnology, especially efficient in vitro culture and molecular markers methods, facilitated application of mutation techniques in breeding and basic research

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Creation of mutants for breeding, germplasm enhancement and biodiversity : e.g. new desired genes for stress resistance (drought, salinity, diseases) or improved quality Induction of genetic variation necessary to exploit the full power of structural and functional genomics for gene isolation directed at the improvement of crops: mutational analysis and plant function and structure What for? Rediscovery of Induced Mutations

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Group No. of mutants % of total Ornamentals 552 24.5 Crops 1700 75.5 Cereals 1072 47.6 Legumes 311 13.8 Industrial 81 3.6 Vegetables 66 2.9 Oil crops 59 2.6 Others 111 4.9 Total 2382 (175 crop species) Mutant cultivars released in different crops in the world Source : IAEA,2000

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Group Species No. of mutants % of total Cereals 7 69 22.3 Legumes 11 52 16.8 Oil crops 5 30 9.7 Industrial 6 24 7.8 Vegetables 8 11 3.6 Ornamentals 9 103 33.3 Other 10 19 6.1 Total 56 309 100.0 Mutant cultivars released in India Source : IAEA,2008

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Mutagens / origin No. of mutants Physical mutagens gamma rays 169 x-rays 26 neutrons 7 gamma EMS 1 other 1 Total 204 Mutagens used in development of mutant cultivars released in India

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Mutagens / origin No. of mutants Chemical Mutagens Ethyl methane sulphonate (EMS) 15 Dimethyl sulphate (DMS) 4 Ethylene imine (EI) 2 Sodium azide (NaN 3 ) 2 Diethyl sulphate (DES) 1 Methyl methane sulphonate (MMS) 1 Colchicine 1 Ethelene oxide 1 Thalium oxide (TO) 1 other 6 Total 34 Mutagens used in development of mutant cultivars released in India

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Main improved characters reported for mutant cultivars in India Main attribute No. of occurrence High yield 86 Early maturity 65 Disease resistance 57 Quality characters 39 Grain quality 67 Abiotic stress resistance 65 Improved plant type 181 Other 9

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Crop Species No. of mutants Mungbean Vigna radiata 13 Chickpea Cicer arietinum 7 Cowpea Vigna unguiculata 7 Blackgram Vigna mungo 7 Pigeonpea Cajanus cajan 5 Moth bean Vigna aconitifolia 5 Lentil Lens culinaris 3 Hyacinth bean Dolichos lablab 2 Pea Pisum sativum 1 French bean Phaseolus vulgaris 1 Cluster bean Cyamopsis tetragonoloba 1 Total 52 Mutant cultivars released in legume crops in India Source: BARC,2008

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Crop Species No. of mutants Groundnut Arachis hypogaea 14 Mustard Brassica juncea 6 Castor Ricinus communis 4 Soybean Glycine max 3 Sesame Sesamum indicum 3 Total 30 Mutant cultivars released in oilseed crops in India

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Variety Release year Main improved attributes & authors Pusa-408 ( Ajay ) 1985, MCK Resistant to Ascochyta blight, high yield, profuse branching, semi erect, maturity 140-155 days Pusa-413 ( Atul ) 1985, MCK Resistant to Fusarium wilt, stunt virus & foot rot, high yield, profuse branching, semi erect, maturity 130-140 days Pusa-417 ( Girnar ) 1985, MCK High resistance to Fusarium wilt & moderate resistance to Ascochyta blight, stunt virus, high yield, profuse branching, maturity 110-130 days Examples of popular mutant cultivars of chickpea developed in India Kharkwal M. C . et al (1985)

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The three chickpea mutants, namely, Pusa-408 (Ajay), Pusa-413 ( Atul ) and Pusa-417 ( Girnar ) developed by the author at I.A.R.I, New Delhi & released by the Govt. of India for commercial cultivation in 1985, are the first ever examples of direct use of induced micro-mutants as improved cultivars in a food legume crop in the world. First examples of mutant cultivars of chickpea developed in the world

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Variety Release year Main improved attributes & authors Pant moong-2 1983, D.P. Singh Resistant to mungbean yellow mosaic virus, higher yield TAP-7 1983, S.E. Pawar Early maturity, tolerant to powdery mildew and leaf spot disease, 23% higher yield over var. Kopergaon TARM-1 1995, S.E. Pawar High yield and resistant to mungbean yellow mosaic virus Examples of popular mutant cultivars of mungbean developed in India

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Variety Release year & Main improved authors attributes TAU-1 1985, S.E. Pawar Early maturity, tolerant to powdery mildew, higher yield TPU-4 1992, S.E. Pawar High yield and moderately resistant to YMV TAU-2 1993, S.E. Pawar High yield and moderately resistant to YMV Some examples of popular mutant cultivars of blackgram developed in India

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Variety Release year Main improved attributes & authors TG-1 ( Vikram ) 1973, S.H. Patil Large kernels (TKW 880g), 47-48% oil, maturity 130-135 days, high yield (4 t/ha) TG-3 1987, S.H. Patil High yield especially under rainfed conditions, more pods per plant TAG-24 1992, S.H. Patil High yield and short plants with higher harvest index. Drought tolerant. Examples of popular mutant cultivars of groundnut developed in India

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Crop & species Mutant var. Release yr. Mutagen Cotton (Gossypium sp.) M.A.-9 1948 gamma rays Wheat (Triticum aestivum) NP-836 1961 x-rays Sugarcane (S. officinarum) Co-6608 1966 gamma rays Castor (Ricinus communis) Aruna 1969 th. neutrons Mulberry (Morus alba) S-54 1974 EMS Bougainvillea (Bougainvillea sp.) Arjuna 1976 gamma rays Blackgram (Vigna mungo) Co-4 1978 MMS Hibiscus (Hibiscus sp.) Purnima 1979 gamma rays Sesame (Sesamum indicum) Kalika 1980 EMS White jute (Corchorus capsularis) Shyamali 1980 x-rays Lentil (Lens culinaris) S-256 1981 radiation Cowpea (vigna unguiculata) V-16 (Amba) 1981 DMS Mungbean (Vigna radiata) TAP-7 1981 gamma rays Chickpea (Cicer arietinum) Pusa-408 1985 gamma rays World’s first mutant cultivars released in India

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Mutant var. Release year Mutagen Pigeonpea ( Cajanus cajan ) Co-3 1977 gamma rays T.Visakha-1 1982 fast neutrons TAT-5 1984 fast neutrons Co-5 1984 gamma rays TAT-10 1985 fast neutrons World’s first and also the only mutant cultivars released in India

Most recently registered mutant varieties :

Most recently registered mutant varieties 1. NIAB-846 Gossypium sp 2. CM2008 Cicer arieatinim 3. Binachinabadam- Arachis hypogea 4. TG 39 Arachis hypogea 5. TG 51 Arachis hypogea (Anon., 2009)

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Conclusion: A major contribution of mutagenesis to plant breeding progress, which can not be over estimated, is its use for the advancement of genetics. The future applications of genetic engineering technology will rely heavily upon mutagenesis on gene pool top localize useful genes, to have involve markers and to provide the desired genetic diversity in molecular gene bank. It can be expected that modern mutation breeding technologies in combination with refined selection methods applicable to large populations will continue to be of great potential for the improvement of both sexually and vegetative propagated crop plants.

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