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Control of the Size and Composition of the Embryo in Cereals

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Control of the Size and Composition of the Embryo in Cereals Control of the size and composition of the embryo in cereals Frederick R Cook A thesis submitted to the University of East Anglia for the degree of Doctor of Philosophy John Innes Centre February 2011 © This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with the author and that no quotation from the thesis, nor any information derived there from, may be published without the author’s prior consent. I. Abstract The embryo is an important source of lipid, protein, minerals and vitamins in the cereal caryopsis, and modification of embryo size and/or composition has the potential to improve the nutritional value of the grain. In this study, two approaches were taken to investigate the control of embryo size and composition. To investigate whether lesions affecting grain development also have effects on the embryo, a collection of shrunken-grain mutants of barley were studied. Embryo weight was found to be unaffected or to be reduced concomitantly with grain weight, with the exception of mutants with lesions at the Lys3 locus. Four lys3 mutants were shown to have greater embryo weight (as previously described in lys3 mutant Risø1508 [Tallberg, 1977]) and lys3 embryos contained greater amounts of protein and micronutrients. An attempt was made to identify the gene underlying the Lys3 locus by characterising the mutant phenotype. The rice giant embryo (ge) mutant was identified as having a similar phenotype to lys3. The barley orthologue of the rice Ge gene was located on chromosome 2H and it therefore cannot underlie Lys3 on 5H. The possibility that Lys3 and Ge act in the same metabolic pathway is discussed. The role of starch in the embryo was investigated by the identification and characterisation of an ADP-glucose pyrophosphorylase large-subunit mutant of rice (apl3-2), which had a 35 percent reduction in starch in the embryo. There was no discernable difference in embryo size or lipid content in the mature embryo. As such, there is no evidence that the quantity of starch accumulated during development is important for determining final embryo composition or for normal embryo development. The apl3-2 mutant was shown to have greatly reduced starch in the culms which adds to our understanding of the function of this gene. 2 II. Table of Contents 1. General introduction ....................................................................................... 10 1.1. The importance of cereals ................................................................................... 12 1.2. Formation of the cereal caryopsis ...................................................................... 15 1.2.1. Fertilisation ....................................................................................................... 15 1.2.2. Embryo development ....................................................................................... 15 1.2.3. Endosperm development ................................................................................. 17 1.2.4. Assimilate supply to the developing embryo .................................................... 18 1.3. Grain composition ................................................................................................ 20 1.3.1. Starch ............................................................................................................... 22 1.3.2. Protein .............................................................................................................. 24 1.3.3. Lipid .................................................................................................................. 26 1.3.4. Minerals and vitamins....................................................................................... 27 1.4. Cereals for food and feed ..................................................................................... 27 1.4.1. Starch ............................................................................................................... 28 1.4.2. Protein .............................................................................................................. 29 1.4.3. Lipid .................................................................................................................. 29 1.4.4. Minerals and vitamins....................................................................................... 30 1.5. Control of the size of the embryo and endosperm ............................................ 30 1.5.1. Endosperm size ............................................................................................... 31 1.5.2. Embryo size ..................................................................................................... 31 1.6. Embryo composition ............................................................................................ 36 1.7. Research aims ...................................................................................................... 37 2. Materials and methods ................................................................................... 38 2.1. Plant materials ...................................................................................................... 38 2.1.1. Barley ............................................................................................................... 38 2.1.2. Rice .................................................................................................................. 38 2.1.3. Maize ................................................................................................................ 38 2.1.4. Wheat-barley chromosome addition lines ........................................................ 39 2.2. Plant growth .......................................................................................................... 39 2.2.1. Growth conditions and soils ............................................................................. 39 2.2.2. Seed sterilisation .............................................................................................. 40 3 2.2.3. Embryo weights ................................................................................................ 40 2.2.4. Judging the developmental stage of grains ...................................................... 40 2.2.5. Crossing ........................................................................................................... 40 2.3. Biochemical methods ........................................................................................... 41 2.3.1. Carbohydrate extraction ................................................................................... 41 2.3.2. Carbohydrate assay ......................................................................................... 42 2.3.3. Protein extraction and assay ............................................................................ 43 2.3.4. Lipid extraction and measurement ................................................................... 43 2.3.5. ADP-glucose pyrophosphorylase extraction .................................................... 44 2.3.6. ADP-glucose pyrophosphorylase assay ........................................................... 44 2.3.7. Separation of plastidial and cytosolic AGPase isoforms using fast protein liquid chromatography (FPLC) ............................................................................................. 44 2.3.8. SDS-PAGE and immunoblotting ...................................................................... 45 2.4. Molecular methods ............................................................................................... 45 2.4.1. Isolation of RNA from plant tissue .................................................................... 45 2.4.2. Isolation of genomic DNA from plant tissue ..................................................... 46 2.4.3. First strand cDNA synthesis ............................................................................. 46 2.4.4. Polymerase chain reaction (PCR) .................................................................... 46 2.4.5. Primers ............................................................................................................. 46 2.4.6. Visualisation of PCR products .......................................................................... 47 2.4.7. DNA sequencing .............................................................................................. 47 2.5. Light microscopy .................................................................................................. 47 2.5.1. Fixation ............................................................................................................. 47 2.5.2. Visualisation and analysis ................................................................................ 48 2.6. Bioinformatic methods ......................................................................................... 48 2.6.1. Sequence alignment ........................................................................................ 48 2.7. Statistical methods ............................................................................................... 48 3. Endosperm-embryo interactions: embryo size and composition in mutant barley lines with altered starch accumulation in the endosperm ................... 49 3.1. Aim ......................................................................................................................... 49 3.2. Introduction ..........................................................................................................
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