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Production and Utilization of Doubled Haploid Lines in Wheat Breeding Programs

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Production and Utilization of Doubled Haploid Lines in Wheat Breeding Programs rì yr.t+.Q¿\ PRODUCTION AND UTILIZATION OF DOUBLED HAPLOID LINES IN WHEAT BREEDING PROGRAMS by GHOLAM ALI RANJBAR B.S. Agronomy (Mashhad Univ.), M.S. Plant Breeding (Isfahan Univ. of Industrial Technology), IRAN Thesis submitted for the degree of Doctor of Philosophy ln The University of Adelaide Faculty of Agricultural and Natural Resource Science Department of Plant Science, Waite Agricultural Research Institute March, 1997 Table of Contents SUMMARY... ,.1 STATEMENT .................... iii ACKNOWLEDGEMENTS .iv CHAPTER 1 General Int¡oduction PART I PRODUCTION OF DOUBLED HAPLOID LINES IN WHEAT 4 CHAPTER2 Review of Literature ............... 5 2.1. DousLrD HAPLoIDS, Ennry H¡srony tNn Appl,lcRtto¡.ts 5 2.2. Appttctno¡¡s op HnpLons 6 2.2. Ì. Rapidly Achieving Homozygosity 6 2.2.2. Cytogenetic Research and Genome Mapping 7 2.3. Hnruon PRoDUcTIoN Sysrel,rs poR Wrs¡,r MoDELLED oN Bnru-sy sysrEM 7 2.3. L In vitro Androgenesis (Anther Culture) 8 2.3.2. Interspecific and Intergeneric Hybridi2ation.s...,............,...... 9 2.3.2.1. Wheat x Hordeum bulbosumHybrids ...... 9 2.3.2.2. Wheat x ka mays Hybrids......... II 2.3.2.3. Wheat x Sor g hum bic o lor Hybids........... t2 2.3.2.4. Wheat x Pennisetumamericanum Hybrids......... ............. 13 2.4. Cgnovosotræ Elttvll.¡ATIoN .......... l5 2.4.1. Time of Elimination of Chromosomes.. 17 2. 4. 2. The M echanism of C h ro mo s ome EI imination ............... t8 2.5. Emcre¡¡cy oF HApLorD PRoDUcroN 20 2.5.L Factors Affecting Seed 9et...... 21 2. 5.2. Factors Affe cting Seed ïurviva| ................ )) 2.5.3. Effects of Rate of Embryo Production, Embryo Size and Regeneration 23 2.5.4. Factors Affecting Frequency of Fertilization. 24 2.5. 5. Effects of.Genorype 25 2.5.6. Effects of Gibberellic Acid (GA j) Treatment, Temperature and Pre-Anthesis Pollination.......... 25 2.5.7. Effects of 2,4-D..... 26 2. 6. DupucnrroN oF CHRoMosoME NuN,lseR....... 29 2.6.1. Spontaneous Chromosome Doubling of Haploids 29 2.6.2. Co lchicine Treatment.... 29 2.6.3. Application of Nitrous Oxide 31 Application of ^ _ ?.6.4. Caffeine.. .....31 2.1-.-pou¡Ls_oHAPLotDs As A TooL IN BARLEy BREEDrNG..... ............. see insert at back of thésii' CHAPTER 3 Producing Doubled Haptoids by a Wheat x Maize System.......... 32 3. l. INTRoDUcrIoN ...,...... 32 3.2. M¡,renrA¡-s .qNo MerHoDS......... 35 3.2. I . Production of Doubled Haploids using Dffirent wheat and Maize Genotypes......................,.. 35 3.2.1.1. Plant Materials 35 Table of Contents 3.2.1.2. Emasculation and PolIination.................... 35 3.2. | .5. Incubation and Regeneration of Haploid Plantlets 3t 3.2.1.6. Chromosome Doubling .......43 3.2.2. Production of Doubled Haploids from F¡ Hvbrid of Trident/Molineux........ ..........43 a) Using maize and sorghum pollen sources.....,., ,.43 b ) U s in g diffe re nt c ult ure me dia..................... 46 3.2.3. Production of Doubled Haploids from 24 Different F t Wheat Crosses.. ..46 3.3. RESULTS .48 3.3.1. Production of Doubled Haploids Using Dffirent Wheat and Maize Genotypes.... .48 3.3. I .l . Female Parent Effects .48 3.3.1.2. Male Parent Effects ,49 3.3.2. Production of Doubled Haploids from Trident/Molineux F ¡ Hybrids ... 51 a) Using maize and sorghum pollen sources 5l b) Using dffirent culture media 52 3.3.3. Production of Doubled Haploids from 24 Different Ft Wheat hybrids 53 3.4. DrscussloN 55 I ) Production of haploid embryos 56 2 ) P lantlet regeneration.... 59 3) Chomosome doubling 60 PARTtr STUDIES ON SEGREGATION PATTERNS IN A WIIEAT DOUBLED HAPLOID POPULATION ..................... .61 CHAPTER 4 Segregation Patterns of Known Major Genes in the Doubled Haptoid Population Derived from a (Trident/lVlolineux) Wheat F1 x Maize Crosses 62 4. l. INrrRooucrroN .......... 62 4.2. M¡renrA¡-s AND METHoDs..... 63 4.2.1. Determination of Endosperm Protein Phenotypes 64 4.2.1.1. Gliadin Analysis 65 4.2.1.2. Glutenin Analysis 65 4.2.1.3. SDS-PAGE Preparation and Electrophoresis 66 4.2.1.4. Staining and drying the gels.......... 66 4.2.1.5.Identifying and Scoring the bands 67 4'2.2. Segregation of Genes Controlling Cereal Cyst Nemøtode Reaction (Heterodera avenae).......... 67 4.2.2.1. Setting up the Nursery 67 4.2.2.2. Scoring............. 68 4.2.3. Measurement of Plant Height and GAj Insensitivity......... 68 4.2.4. Assay for High Soil Boron Tolerance............. 71 4,2.5. Segregation of Genes Controlling Rust Reaction...... 73 4.3. Rpsulrs 75 4.3.1. Determination of Endosperm Protein Phenotypes 75 4.3.2. Segregation of Genes Controlling Cereal Cyst Nematode Reaction.. 78 4.3.3. segregation of Genes controlling Plant Height and GA j Insensitivity. 80 4.3.4. Assay for High Soil Boron Tolerance........... 82 4.3.5. Segregation of Genes Controlling Rust Reaction............. 85 4.4. Dlscusslon 85 CHAPTER 5 Inheritance of Some Agronomic Attributes in the Doubled Haploid Population Derived from Trident/lVlolineux F¡ x Maize Crosses .88 Table of Contcnts 5.I . INTRoDUSTIoN 88 5.2. MnrsruA¡-s AND MSTHoDS................ 88 Statistical procedure ...89 5.2.1. Segregation of Genes Controlling Septoria tritici Blotch Reaction.. 89 5.2.2. Segregation of Genes Controlling Coleoptile Length 92 5.2.3. Segregation of Genes Controlling Leaf Colour .. 93 5.2.4. Segregation of Genes Controlling Heading Date 5.3. R¡su¡-rs a¿ 5.3.1. Segregation of Genes Controlling Septoria tritici Blotch.... ...............96 5.3.2. Segregation of Genes Contolling Plant Height 100 5.3.3. Segregation of Genes Controlling Coleoptile Length 101 5. 3.4. S e g regation of Genes Contro lling Leaf Co lour .................. 105 5.3.5. Segregation of Genes Controlling Heading Date .......... I 06 5.3.6. Segregation of Genes Controlling Yield I 07 5.4. Drscussro¡r 1 08 General conclusion I PART Itr CROSS PREDICTION STUDIES WITH WHEAT DOUBLED HAPLOIDS....... 113 CHAPTER 6 6.1. Comparison Between Doubled Haploid and F5 Lines Derived from DifferentWHEATFr......l 14 6. L l . Introduction............ t14 6.1.2. Materials and Methods t17 6.I .3. Results t22 6. L4. Discussion ........... t29 6.2. Comparison Between Hill- and Row-Plot on Yields of Doubled Haploid and F5 Lines of Wheat..... t32 6.2.1. Introduction t32 6.2.2. Materials and M ethods .............. 132 6.2.2.1. Hill-Plot Experiment.. 32 6.2.2.2. Row-Plot Experiment. 32 6.2.2.3. Field trials I 996 ............. 33 33 6.2.4. Discussion ..... 39 CHAPTER 7 General Discussion t4t Rprrnr¡qcps. 149 Appnxurcns ..171 AppsNolx I 172 AppeNux 2 178 Summary For doubled haploids to become a valuable breeding technique, it is necessary that: 1. Success ofdoubled haploid production is not genotype dependent 2.The protocol used is efficient in time and cost 3. The doubled haploid population produced represents a random sample of gametes For wheat, to date, intergeneric hybridization with muze has been shown to be more efficient than other methods used for other cereals such as anther culture or hybridization with Hordeum bulbosum. The protocol which was established and later used in this project involved maize cultivar Illini Gold which was the most efficient pollen donor of the six maize and two sorghum lines tested. 2,4-D at 150 mgl-' was applied after pollination to increase embryo formation rates. The best culture medium for embryo rescue and regeneration was found to be half strength MS. This protocol was then used to produce 217 doubled haploids from the F, of a cross between Trident and Molineux wheats. This cross was polymorphic for a number of attributes. The genetic control of some of these characters was known and the segregation patterns of these characters were used to test for randomness of haploid production. The inheritance of the other characters which were of economic importance were unknown and the doubled haploids were used to investigate their inheritance. Segregation patterns for high and low molecular weight glutenin subunits (GI1AI, GI¡A3, GluB3), resistance to cereal cyst nematode (Cre), plant height, boron tolerance (Bol) and stem rust resistance (Sr38) were investigated. In all cases these genes showed no distorted segregation ratios. The doubled haploids produced appeared to be from a random sample of wheat female gametes and so could confidently be used in gene mapping and to investigate the inheritance of other attributes. puge i Summan The other characters segregating in the Trident/lvlolineux doubled haploid population included resistance to Septoria tritici, coleoptile length, leaf colou¡, heading date and grain yield. As many of these attributes appeared to be polygenic in nature, the four cumulants of their distributions viz. mean, variance, skewness and kurtosis were calculated. Also, the number of genes (or effective factors) controlling these attributes in this cross were estimated. The highest estimate was for coleoptile length of dwarf lines (10 genes) and semidwarf lines (8 genes), the height variation within the semidwarf group (8 or 9 genes) ) and leaf colour (9 gene). The results from two methods used for scoring Septoria tritici ^ blotch reaction were in agreement with the results of other investigators and it was estimated that2 to 3 genes determine resistance to this disease. Additive epistasis and gene interaction were observed in some biased distributions. Multivariate analyses and associations between some of these characters, for example, coleoptile length with plant height, leaf colour with yield and Septoria reaction with plant height and maturity are presented. In a separate study a limited number of doubled haploids were produced for 24 different F, crosses. These doubled haploids were then compared with F, lines derived from the same crosses made in a conventional pedigree breeding program several years earlier.
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