male sterility ans self incompatibility in vegetable hybrid seed pr

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Presented By:- Chanchal rana A- 2013-30-066 USE OF MALE STERILITY AND SELF INCOMPATIBILITY FOR HYBRID SEED PRODUCTION IN VEGETABLE CROPS 1

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Male sterility: The failure of plants to produce functional anthers, pollen, or male gametes. Self incompatibility: functional pollen and the receptive stigma of the same plant are not able to fertilize. Male sterility and self incompatibility are mechanisms preventing selfing, thus reducing the labour and cost incurred for hybrid production to an extent 2

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WHY MALE STERILITY AND SELF INCOMAPATABILITY? Production of F 1 seeds at large scale. Reduced cost of labour for hybrid seed production. Speedup the hybridization programme . Used for commercial exploitation of hybrid vigour . 3

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Why hybrids are adopted by farmers? Greater productivity Longer harvest duration Better adaptability to variable environments Better tolerance to diseases and pests Uniform produce Better market acceptability Better nutritional quality 4

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10 20 30 40 50 60 80 70 90 100 27 60 HYBRID SEEDS VARIETAL SEEDS PERCENTAGE 2007 2008 2009 2010 2011 2012 2013 KEN RESEARCH 5

Vast Domestic and International market trend:

Vast Domestic and International market trend Requirement of the vegetable seed 2001-02 2005 Open pollinated varieties 30550 t 48000 t Hybrid vegetable seed 346.2 t 994 t After 2007, requirement of hybrid seeds became greater than OPVs 6

Contribution (%) of various vegetables to hybrid seed market segmentation (2013):

Contribution (%) of various vegetables to hybrid seed market segmentation (2013) 7

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Estimated share of vegetable hybrids in total cropped area in India (2008-09). Crops Total area (ha) Area under F 1 hybrids (ha) Share of hybrid’s area (%) Egg plant 4,79,095 85,300 17.80 Cabbage 2,42,140 76,000 31.39 Cauliflower 3,04,156 10,000 3.29 Chilli 5,73,529 14,000 2.24 Gourds 4,09,270 10,000 2.44 Melons 1,69,350 6,800 4.02 Okra 3,71,665 20,000 5.38 Tomato 4,85,520 1,53,000 31.51 Total 30,34,725 3,75,100 12.36 Source : Indian Horticultural Database (2009) 8

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The most commonly utilized mechanisms for developing commercial hybrids in vegetables. Mechanism Commercially exploited in: Hand emasculation + HP Tomato, eggplant, sweet pepper, okra, hot pepper Remove staminate flowers + HP Cucurbits (bitter gourd, bottle gourd, etc. ) Male sterility + HP Tomato, hot pepper, sweet pepper Male sterility + NP Onion, cabbage, cauliflower, carrot, radish, muskmelon, hot pepper. Self incompability + NP Most of cole vegetables like broccolis, cabbage etc. Gynoecism + NP Cucumber, muskmelon. Remove staminate flowers + NP Cucurbits including bitter gourd, summer squash, winter squash, etc. HP = Hand pollination NP = Natural pollination 9

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Time(minutes ) required for crossing 50 flower buds on male fertile ‘Ms33 IPA’ (MF) and male sterile ‘ ms 331PA’ ( MS) plants in tomato Dhaliwal and Cheema , 2008, Ludhiana Worker   Emasculation   Pollination on MF Emasculation + pollination on MF Pollination on MS Time saved in MS over MF, % 1 22.0 44.0 66.0 37.7 42.9 2 26.1 34.9 60.5 26.3 56.5 3 38.7 45.1 83.8 33.3 60.3 4 37.3 43.8 75.5 34.2 54.7 5 32.8 41.1 73.9 32.7 55.8 Mean 30.7 41.7 71.9 32.8 54.4 CD at 5% 1.5 1.6 2.0 1.1 -  

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MALE STERILITY 11

Introduction:

Introduction 12

Types of male sterility based on phenotypic expression of genes:

Types of male sterility based on phenotypic expression of genes 13

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A) Genetic/nuclear male sterility( GMS) : - wide occurrence in plants - mostly governed by a single recessive gene , ms - male sterile alleles arise spontaneously or may be artificially induced B ) Cytoplasmic male sterility (CMS) : - determined by the cytoplasm - it is the result of mutation in the mitochondrial genome (mtDNA) - CMS transfer easily to a given strain C ) Cytoplasmic-genetic male sterility (CGMS) : - nuclear gene restores the fertility in the male sterile line - also know as nucleoplasmic male sterility - fertility restorer gene R is required D ) Chemically induced male sterility : - IAA, IBA, Ethrel , etc . Kaul (1988) Types of male sterility based on inheritance 14

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By microspore appearance under 100x magnification (0.5% aceti -carmine stain) Gniffke et al., 2009, Taiwan

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Mutant Description Inheritance Governing by single recessive gene Stamenless Stamens absent Monogenic recessive sl Positional sterility Stigma exerted Monogenic recessive ps Pollen sterility Pollen abortive Monogenic recessive ms series Functional sterility Anthers do not dehisce Monogenic recessive ps-2 Table 6: Different male sterile mutants in tomato. 16 Opena et al., 2009, Taiwan

Genetic male sterility:

Genetic male sterility Controlled by a number of nuclear genes- MALE STERILE GENES . Effectiveness of these genes maybe expressed by- %age of viable pollens. %age of seed set. GMS is controlled by a pair of recessive alleles “ msms ”, Few dominant genes have been reported Inheritance pattern 17

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Transfer of genetic male sterility to new strain 18

Utilization in hybrid production:

Utilization in hybrid production 19

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GMS line Mean percent fruit set on GMS line Natural out crossing Hand pollination ACMS 2 -1-1-1 32.64 13.82 ACMS 2 -1-1-4 30.22 14.08 ACMS 2 -5-1-1 35.99 14.82 ACMS 2 -5-1-5 34.39 14.00 ACMS 2 -6-1-1 30.68 15.34 ACMS 2 -6-1-3 32.82 14.60 S. Em - 0.35 Average of mean percent fruit set on GMS lines 32.79 14.44 Anand Patel et al. (2001) 20 Per cent fruit set on male sterile (GMS lines) plants in chilli .

Constraints:

Constraints Generally , pollen dispersal is poor Closely linked markers are rare to identify the male fertile plants . Identifying the male fertile plants is difficult and time consuming. Hence GMS is not commonly used in hybrid seed production. Rouging of male fertile plants from the female line (male sterile) is costly as a result of which the cost of hybrid seed is higher . Use of temperature or photo sensitive genetic male sterile lines (TGMS or PGMS) eliminates this problem 21

Environmental genic male sterility EGMS :

Environmental genic male sterility EGMS EGMS: T hese GMS lines are conditional mutants In a particular environment male sterile mutant plants turn into male fertile. Usually temperature or photoperiod M ostly temperature sensitive EGMS lines have been reported I t is necessary to identify critical temperature or photoperiod for the fertility/sterility expression in temperature and photoperiod sensitive genetic male sterility, respectively. 22

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Vegetable Mutant Reference Cabbage TGMS , PGMS Rundfeldt , 1961 Brussels sprout TGMS Nieuwhof , 1968 Broccoli TGMS Dickson , 1970 Pepper TGMS , TCMS Daskalov , 1972; Shifriss , 1997* Carrot TGMS Kaul , 1988 Tomato TGMS Rick , 1948; Sawhney , 1983   TGMS–Thermo sensitive genic male sterility PGMS–Photoperiod sensitive genic male sterility TCMS–Thermo sensitive cytoplasmic male sterility 23 Environmental genic male sterility EGMS

Cytoplasmic male sterility:

Cytoplasmic male sterility The expression of male sterility in CMS plants is the result of incompatibility between recessive nuclear gene (called maintainer gene; rf ) and male sterile specific cytoplasmic genome . Expression of male sterility is due to the defect in the cytoplasm ( mt -genome ) Cytoplasmic male sterile plants have also been developed in several vegetables through protoplast fusion (Pelletier et al., 1995 ) Three Ogura based improved cytoplasmic male sterile (CMS) lines of cauliflower (Brassica oleracea var. botrytis L.) viz., Ogu1A, Ogu2A and Ogu3A were developed following seven generations of backcrossing with snowball group ( Dey et.al . 2011) 24

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Flower phenotypes in carrot:

Flower phenotypes in carrot a) Normal (N-cytoplasm, restored CMS plants) b) Brown anther CMS c) Petaloid CMS

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27

Utilization:

U tilization 28

CGMS:

CGMS 29 Once dominant restorer ( Rf ) gene (located in nuclear genome) responsible for pollen fertility of a cytoplasmic male sterile line is identified, it is commonly known as cytoplasmic-genic male sterility ( CGMS ).

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Fertility restorer line development 30

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U tilization 31

Transgenic Male Sterility:

Transgenic Male Sterility All the transgenic male sterile lines developed till date are GMS, since they have been developed through transformation of male sterility causing gene construct(s) inside the nuclear genome . Ex: barnase - barstar system in canola ( Brassica sp.) G enes are expressed in pollen themselves ( gametophytic expression ) or cells and tissues ( sporophytic expression) that directly or indirectly support pollen development, such as tapetum , filament, anther wall 32

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33 Transgenic Male Sterility

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34 Transgenic Male Sterility

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35 Transgenic Male Sterility

Male sterility in vegetables:

Male sterility in vegetables Peterson first observed cytoplasmic genic male sterility in Capsicum annuum L . Onion breeding was started in the early thirties, based on male sterility found in onion in California during 1925 from the cultivar Italian Red 13-53 (Jones and Emsweller , 1937 ). I dentification and exploitation of CMS system in onion (Jones and Davis, 1944 ). In India, Sen and Srivastava ( 1957) attempted , for the first time, to develop F1 hybrids in onion (as early as in 1948) using exotic male sterile lines and Indian local male stocks 36

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Genetic male sterility (GMS) has been exploited commercially in chilli and muskmelon to develop F 1 hybrid seed commercially. Punjab Agricultural University (PAU), Ludhiana, India has released two chilli hybrids (CH-1 and CH-3) O ne muskmelon hybrid (Punjab Hybrid) based on the GMS system. Similarly in tomato, work on GMS lines is in progress at PAU . CMS-based hybrids, i.e Kashi Surkh (CCH-2) and Kashi Early (CCH-3 ). Chilli hybrids based on the CGMS system, i.e. Arka Meghna (MSH-172), MSH-149 and MSH-96 . ( Dhall , R. K . status of male sterility for hybrid evelopment in vegetables 2010.) 37

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In carrot ' Pusa Nayanjyoti ', which is based on petaloid CGMS in carrot by IARI at K atrain . Arka Kirtiman and Arka Lalima , In onion based on the CGMS system Onion (Hybrid-63 and Hybrid-35) based on CGMS by IARI H ybrids in cabbage by utilizing Ogura CMS system have been developed The Ogura-type CMS has been transferred into heat-tolerant Indian cauliflower from kale and broccoli through repeated backcrosses four lines (MS-91, MS-51, MS-11 and MS-110) from the former and five lines (MS-01, MS-04, MS-05, MS-09 and MS-10) from the latter were developed . ( Dhall , R. K . status of male sterility for hybrid development in vegetables 2010 .) 38

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ms ms ms ms ms ms Three Possible genotype Fig. 2 : Genetic male sterility for hybrid seed production in chilli THREE POSSIBLE GENOTYPE 39

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40

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Natural population Artificially induced through mutagenesis Genetic engineering Protoplast fusion SOURCES FOR MALE STERILITY 41

Significance of male sterility:

Significance of male sterility 42 Cytoplasmic male sterile plants have also been developed in several vegetables through protoplast fusion (Pelletier et al., 1995).

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43 Significance of male sterility

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SELF INCOMPATIBILITY 44

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Self incompatibility :- SI refers to the inability of a plant to set seed upon self pollination despite male and female gamete is viable, there is no seed set in self pollination first discussion on self-incompatibility by Darwin (1877 Lewis (1954) has suggested various classifications of self incompatibility Heteromorphic system Pin type Thrum type Homomorphic system Sporophytic self incompatibility (SSI) Gametophytic self incompatibility (GSI) 45

History of self-incompatibility :

History of self-incompatibility First discussion on self-incompatibility by Darwin (1877) The term self incompatibility was given by Stout (1917) Bateman (1952, 1954, 1955) gave explanation on incompatibility in three Brassicas plants namely, Iberis amara L., Raphanus sativus L. and Brassica campestris L . 46

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1. Heteromorphic: different morphology of flowers prevent the self pollination Distyly Tristyly Ex: pin and thrum flowers in petunia 2. Homomorphic : flowers have same morphology a. Gametophytic . SI reaction of a pollen is determined by its own genotype not by the genotype of the plant on which the pollen is produced b. Sporophytic . SI reaction of pollen is governed by the genotype of the plant on which the pollen is produced and not by the genotype of the pollen. McCubbin and Dickinson (1997) Types of self incompatibility 47

Difference between GSI and SSI:

Difference between GSI and SSI Gametophytic SI Sporophytic SI The stigma is smooth and wet The stigma is papilate ( hairy) and dry Pollen tube inhibition in style Pollen tube inhibition take place on the stigmatic surface itself The pollen-pistil interaction govern by h aploid genome of each male gametes and diploid genome of pistil tissue ( Haplo-Diplo ) The pollen-pistil interaction govern by genome of the plant on which the male and female gamete produced ( Diplo-Diplo ) Genetic constitution of pollen controls SI Genotype of sporophyte/plant Sterile, partial sterile and fertile crosses obtained Either fertile or sterile are observed 48 Homozygotes not obtained Homozygotes can be obtained

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Gametophytic self incompatibility Sporophytic self incompatibility 49

Sporophytic self incompatibility:

Sporophytic self incompatibility SI phenotype of the pollen is determined by the genotype of the sporophyte (pollen producing parent) It is controlled by single s-locus with multiple allele but also reported more than one loci, more than 80 s-alleles reported in Brassica family ( On the basis of population) 50

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This system was confirmed :- Kale ( Thompson 1957) Radish ( Sampson 1957) Broccoli ( Sampson 1957 and Odland 1962 ) Cabbage ( Adamson 1965) Cauliflower ( Hoser - krauze 1979) 51

Problems in exploiting self Incompatibility:

Problems in exploiting self Incompatibility Sib-incompatibility is weak in certain inbreds Continuous inbreeding may lead to complete loss of the inbred lines Pseudo-incompatibility Hybrid seeds would be expensive if the self-incompatible lines are difficult to maintain 52

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Self incompatibility and its utilization in hybrid seed production 53

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Number of siliqua setting from open pollination, sib mating, bud pollination and by bagging in Indian Cauliflower . Genotype Siliqua setting upon different pollination systems (%) Open pollination Sib mating Bud pollination Self pollination (Bagging) IIVR-1 70 65 50 00 IIVR-50 80 60 40 00 * kataki Early-29 95 75 25 00 *Hazipur-4 58 75 35 00 Pusi – 4 75 75 60 00 * Agahani JBT -23/60 75 95 15 00 *Late Agahani 60 50 20 00 *Aghani-8 75 80 25 00 Pusi Hazipur 90 95 55 00 Agahani-31 85 70 40 20 Agahani long leaf 75 65 50 00 * Agahani small leaf 70 60 35 00 Kuwari-1 80 50 50 00 Kataki-12 100 70 55 00 CD at 5% 11.86 3.58 4.07 - Singh et al . (2002)

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Basic steps in the use of SSI Identification of self-incompatible plants in diverse population/genotypes Development of homozygous self-incompatible lines Identification of S-alleles in the homozygous self-incompatible lines Establishment of inter-allelic relationships among the S-alleles Identifying the best combining lines Maintenance of parental self-incompatible lines Commercial hybrid seed production 55

Maintenance of homozygous SI inbreds :

Maintenance of homozygous SI inbreds Bud pollination / Sibmating Treatment with CO 2 gas (CO 2 enrichment) ( Jirik 1985) or sodium chloride ( Kucera 1990 ) Tissue culture using meristem Sodium chloride sprays Removal of stigmatic surface or whole stigma, useful in sporophytic system. Exposure to high temperature. Double pollination 56

Other methods:

Other methods Electronic aided pollination (EAP); ( Roggen et al . 1972) Steel brush method ( Roggen and Dijik 1972 ) The pollen washing ( Roggen 1974) Thermally aided pollination (TAP); ( Roggen and Dijik 1976 ) 57

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Lines/varieties Number of seeds per siliqua Natural Cross Pollination Bud pollination Self pollination in freshly opened flowers self pollination (bagging) * Pusa Chetaki 5.31 3.11 0.04 0.00 * Pusa Desi 5.40 4.64 0.03 0.00 **Half Red 6.00 1.80 0.00 0.00 Acc. No. 30205 4.76 3.51 4.40 1.23 **Acc . No. 282 5.83 4.08 0.00 0.00 **Chinese Pink 3.82 2.09 0.00 0.00 *BDJ-689 5.85 4.96 0.06 0.00 Seed set in radish lines under different types of pollination. Kutty and Sirohi (2004) **Fully self incompatible *Highly self incompatibility

Use SI line for hybrid production:

Use SI line for hybrid production 59

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Types of crosses 60

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Stable self incompatibility. High seed set of self pollination at bud stage. Favorable and uniform economic characters. Desirable combination ability. The superior self incompatible lines for hybrid seed production should possess the following characters 61

Self incompatibility in vegetables:

Self incompatibility in vegetables Self-incompatibility (SI) has been used for the Cruciferae family vastly. The study of SI in crucifer crops began in Japan, where it still continues. In 1949, a Chinese cabbage F 1 hybrid variety, " Nagaoka Kohai I Go", was produced by Shojiro Ito, and in 1961 a radish F 1 hybrid variety, " Harumaki Minowase ", was produced by a commercial seed company. SI is governed by a series of multiple alleles (referred to as the S -gene) (Bateman 1955 ) 62

Self incompatibility in vegetables:

63 In brassicaceae incompatibility (SSI) has been best characterized and successfully utilized for the development of commercial hybrids (Pearson, 1983; McCubbin and Dickinson, 1997, Tripathy and Singh 2000, Singh 2000, Singh et al. 2001 ). Sporophytic self-incompatibility (SSI) was first observed in radish (Stout, 1920), and its inheritance pattern was first demonstrated (Bateman, 1955 ). The numbers of S allele at S -locus have been reported to be 34 in Raphanus sativus and 60 in B. oleracea . Self incompatibility in vegetables

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Fig. 4: Some genotypes available in India produced using Male sterility and Self incompatibility Male sterility Self incompatibility TOMATO Ludhiana ms33 IPA ms2 IPA ps2 L 3841 ps2 NS 101 ps2 San Pedro ps2 UC 82-B ms 10 36 ms 45 ms 15 47 CHILLI Anand CCA 4261 CCA 4759 CCA 4758 Ludhiana CCA 4261 ONION Bangalore Ms1 Ms2 ms3 RADISH IARI NEW DELHI Pusa Chetaki Pusa Desi Half Red Acc. No. 30205 Acc. No. 282 Chinese Pink BDJ-689 CAULIFLOWER Varanashi IIVR-1 IIVR-50 kataki Early-29 HAZIPUR-4(BP) Pusi – 4 Agahani JBT -23/60 Late Agahani Aghani-8 Pusi Hazipur Agahani long leaf Agahani small leaf Kuwari-1 Kataki-12 64 64

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CONCLUSION Pollination control mechanism is helpful for making the hybrid seed production profitable by reducing cost. The male sterility and self incompatibility are becoming increasingly important for hybrid seed production in vegetables crops. Male sterile lines are available in many crops useful for developing hybrid vigour . Functional male sterility governed by ps-2 gene in tomato has indisputable advantage. GMS and CGMS are utilized for hybrid seed production in chilli . MS line having S, T and G cytoplasm are utilized for hybrid seed production in onion. Highly self incompatible lines are available in radish and cauliflower for its commercial utilization in hybrid seed production . 65

Future prospects:

Future prospects To fulfill the demand of vegetables in our country, develop more of hybrid varieties for higher yield. To identify the stable SI and CMS lines There is need to acquire deeper knowledge about these two important mechanisms for developing location/ environmental specific SI and MS lines Produce transgenic sterility or incompatibility in the crops in which it is not present Exploit the mechanism in the crops identified. 66

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67 Thank you.

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