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Ch9. Breeding by interspecific or intergeneric hybridization 1. interspecific or intergenerichybridization 2. Cross incompatibility 3. Breeding Methodology
1. 종속간 교잡 (interspecific or intergeneric hybridization) (1) Definition (Chang TT & Vaughan DA. 1991. Conservation and potentials of rice genetic resources. In Biotechnology in Agric. & Forestry 14 (Rice), Bajaj YPS ed. Springer-Verlag publ. pp. 531-552) (2) Various uses of wild species <1> - Introgression breeding : resistance, quality, yield, ... - The induction of haploids through pseudogamy in potato, alfalfa, cotton. popular, and via chromosome elimination in barley and wheat (Nitzsche W & Wenzel G, 1977. Haploids in Plant breeding, Verlag Paul Parey publ. 101p.) - The (re)synthesis of allopolyploid crops, e.g., in Brassicaceae and Gramineae. - The induction of cytoplasmic male sterility, e.g., in wheat, rice, potato, tobacco, sunflower, Brassica crops
(3) Choice of wild species - close to cultivated species : to overcome cross cross-incompatibility and/or sterility problems - diploids preferred : in order to easily eliminate undesirable traits of wild species (4) Examples A. Amphidiploids - Triticum (amphidiploid, allohexaploid) Triticumaestivum AABBDD 1립계 2립계 X DD Triticumtauschii AABB Triticumturgidum 1립계 BB AA X Triticummonoccum Agropyron? - Triticasecale: AABBDDRR, AABBRR X RR AABBDD Secalecereale Triticumaestivum T. turgidum ABDR wheat(42) rye(14) X RR AABBDD Chromosome doubling Triticale(56) AABBDDRR ABR 2n=56 Chromosome doubling AABBRR 2n=42
B. Interspecific/intergenerichybridization grouped by genome and chromosome number i) Same genome, same chromosome number N. alta (n=9)<A> x N. sanderae (n=9) <A> N. tomentosa (n=12) <T> x N. tomentosiformis (n=12) <T> T. aestivum(n=21) <ABD> x T. spelta (n=21) <ABD> O. sativa (n=12) <A> x O. nivara (n=12) <A> ii) Partially same genome, different chromosome number N. rustica (n=24) <PU> x N. paniculata (n=12) <P> T. aestivum (n=21) <ABD> x T. monocum (n=7) <A> T. dicoccum (n=14) <AB> x T. spelta (n=21) <ABD> iii) different genome, same chromosome number O. sativa (n=12) <A> x O. punctata (n=12) <B> iv) different genome, different chromosome number N. suaveolens (n=16) <Su> x N. glutinosa (n=12) <Gu> T. aestivum (n=21) <ABD> x Secalecereale (n=7) <R> O. sativa (n=12) <A> x O. grandiglumis (n=24) <CCDD> Success rate in Interspecific /intergeneric crossing - I > ii > iii > iv - 2~3 genomes x 1 genome (♀) (♂) - chromosome number higher (♀) x lower (♂) - Self compatible x S. incompatible (♀) (♂)
2. Cross incompatibility incompatibility : a mechanism that "prevents or disturbs the functioning of the pollen-pistil relationship --- a precise and specific reaction to the negative effects of inbreeding active inhibition incongruity: the incompleteness of the relationship as a byproduct of evolutionary divergence passive rejection Williams EG. 1987. Interspecific hybridization in pasture legumes. PBR 5. pp.237-305
(1) Pre-fertilization barriers --- similar mechanism to self-incompatibility - Arrest of pollen tube growth, - Lack of fertilization * Heslop-Harrison. 1982. Pollen-stigma interaction and cross-incompatibility in the grasses. Science 215: 1358-1364
* Evans.1962. Euphytica 11: 164-176, 256-262 : Trifolium에서 relationship between cross-compatibility andgrafting compatibility --- related to anatomical similarity among species
(2) Post-fertilization barriers A. Chromosome elimination <1> H. vulgare x H. bulbosum T. aestivum x H. bulbosum H. vulgare x H. marinum Wheat x sorghum " x maize: chromosome-eliminated B. Hybrid embryo breakdown - delayed cell division in embryo (Roupakias,1986. Euphytica. 35:175-183) : Vicia fava x V.narbonensis F1 embryo는 2 weeks after crossing 200 cells parental embryo는 “ “ 2000-5000 cells - breakdown of embryo --- possibly due to endosperm abnormalities * Abboet al. 1991. Bot. Gaz. 152(3): 316-320 water and nutrient supply to hybrid embryo were blocked in Medicago의 interspecific crossing aborted * Rabakoarihanta외 1979. TAG 54: 55-59 Williams 외 1980. Bot.Gaz.141: 252-257 - failure of the endosperm ( followed by starvation of the hybrid embryo) reason: ' endosperm genome의 genetic unbalance ' marternal tissue와 zygote의 genetic composition의 부조화 * EBN(endosperm balance number) hypothesis --- Solanum, Trifolium(클로버), Avena등. Parrott & Smith. 1985. Evidence for the existence of endosperm balance number ~ ~Can. J. Genet. & Cytol. 28: 581-586. : Each species has an effective endosperm ploidylevel(termed EBN) that determines its crossing behavior. For a cross to be successful, the endosperm must have a ratio of two EBNs from the female parent to one EBN from the male parent. 즉 same EBN successful development of endosperm in hybrid seeds
(3) Genetics for interspecific incompatibility ○ Nettancourt. 1977. Incompatibility in angiosperms * Incongruity hypothesis
○ Lange 외. 1976. Euphytica 25: 609-620 * dominant alleles in wheat-rye crosses : Kr1, Kr2 --> crossability genes ' additive effect ' Kr1의 effect --- bigger wheat(T.aestivum) Hope(Kr1Kr1Kr2Kr2) x rye ↓ X (no zygote) Chinese spring x rye (kr1kr1kr2kr2) ↓ O C.S./Hope 5B (chr.substut. line) x rye (Kr1Kr1kr2kr2) ↓ poor crossability * Kr1은 located on 5B Hope에서 5B가 eliminated된 plant Kr2Kr2 alone x rye intermediate crossability ( 5A? -- not confirmed)
○ Kr1 on 5BL, Kr2 on 5AL, Kr3 on 5D and Kr4 on 1A, with Kr1 having the largest effect on crossability(http://www.jic.ac.uk/staff/graham-moore/index.htm) * Chr. 5B contains Ph1 locus. ○ Manickavelu et al 2009. Plant Syst. Evol. 278: 125-131 - CS (kr1kr2) x Mara 5B (Kr1 kr2) 의 progeny RILs 들을 crossed with rye to know the effect of Kr1 - cDNA-AFLP 방법으로 identified candidate genes for Kr1 ○ Alfares et al. 2009 Genetics 183: 469–481. Fine Mapping and Marker Development for the crossabilitygene SKr on chromosome 5BS of hexaploidwheat. - There was a SKr on chromosome 5BS, besides Kr1 on 5BL - Fine-mapped and Identified candidate gene for SKr
○ Laurie and Bennett (1987) TAG 73: 403-409 - The effect of the crossability loci Krland Kr2on fertilization frequency in hexaploid wheat x maize crosses - Highbury = Kr1 Kr2 CS (5B) = Kr1 kr2 CS = kr1 kr2 ==> - Even Highbury was compatible with maize - No significant difference between CS(5B) and CS. Why?? ===> The effect of Kr1 and Kr2 may differ along species.
(4) Methods for overcoming interspecific incompatibility * Khush & Brar. 1992.
a. Application of growth substances : Auxins, GA3 b. In vitro fertilization or stump pollination stigmatic pollination styler “ placental “ intra-ovarian “ : injection of pollen soln directly into ovary stump pollination : on intact plants without ovary culture,particularly whenstyle length of two species are different--- Solanum, Nicotiana등 c. Irradiation of flower buds --- X-ray : removal of interspecific incompatibility by means of induced mutations d. Use of mentor pollen irradiation freezing and thawing on mentor pollen + incompatible pollen methanol treatment --- effective in Populas, Sesamum * incompatibility within style tissue was maintained. e. Treatment of stigma or pollen with organic solvents : n-hexane, ethyl acetate 등, treatment onstigma before pollination or pollination after pollenwashing with chemicals --- Populas등 f. Somatic hybridization
g. Use of immunosuppressors : breakdown of immune system --- injection treatment of amino-n-caproicacid(EACA), salicylic acid, acriflavin, etc during megasporogenesis effective in crosses of wheat x barley, maize x sorghum h. Bridge cross N. rependa (RR) N. sylvestris (SS) N. tabacum (TT) compatible: RR x SS, SS x TT incompatible : RR x TT i. Embryo culture j. Backcross k. Others * bud pollination * crossing at low temp. condition ( 20℃ in rice interspecific cross) * crossing after chromosome doubling Rice Genetics II p.149 Fig.1
3. Breeding Methodology (1) Introgression
(2) Chromosome substitution or addition Chu. 1982. Anther culture of rice and its significance in distant hybridization. In Rice Tissue Culture Planning Conf., IRRI: 47-53 (3) Amphidiploid: AABBDD, AABBRR 등 (4) Others -- haploid breeding, CMS