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Lathyrus Sativus) for Low Β-L-ODAP Content Genetic improvement of grass pea (Lathyrus sativus) for low β-L-ODAP content Peter Martin Ferdinand Emmrich John Innes Centre Thesis for the degree of Doctor of Philosophy (PhD) Submitted for examination to the University of East Anglia in September 2016 Final version including minor corrections submitted in June 2017 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 use of any information derived there from must be in accordance with current UK Copyright Law. In addition, any quotation or extracts must include full attribution. ii Table of contents Table of contents ........................................................................................................ ii Abstract .................................................................................................................. viii List of Abbreviations .................................................................................................. ix Acknowledgements .................................................................................................. xii List of figures ........................................................................................................... xiv List of tables .............................................................................................................xix Chapter 1 – General introduction .................................................................................... 1 1.1 Food security in the 21st century ..................................................................... 1 1.1.1 The need for increased diversity of food production .................................... 1 1.1.2 The potential of Lathyrus sativus ................................................................... 2 1.2 History of grass pea cultivation ........................................................................ 4 1.2.1 From Neolithic staple to orphan crop ............................................................ 4 1.2.2 Lathyrism – a neurodegenerative disease caused by grass pea .................... 6 1.3 The neurotoxin β-L-ODAP ................................................................................ 9 1.3.1 Biosynthesis of β-L-ODAP in grass pea ........................................................ 11 1.3.2 The ecophysiological role of β-L-ODAP ........................................................ 14 1.3.3 Mitigation of grass pea toxicity through food processing ........................... 15 1.3.4 Physiological breakdown of β-L-ODAP ........................................................ 16 1.3.5 Potential biomedical applications of β-L-ODAP ........................................... 16 1.4 Low-ODAP genotypes of grass pea ................................................................. 17 1.4.1 Released low-ODAP varieties of grass pea .................................................. 17 iii 1.4.2 Attempts to produce ODAP-free genotypes ................................................ 18 1.4.3 The importance of the genotype x environment interaction for β-L-ODAP production ................................................................................................... 19 1.5 Towards the rapid domestication of grass pea ................................................ 20 1.5.1 The problem of grass pea toxicity has not been fully resolved ................... 20 1.5.2 From minor crop to major industrial product: the case of oilseed rape ..... 22 1.5.3 Approaches taken in this study .................................................................... 23 Chapter 2 – Screening grass pea germplasm for low-ODAP genotypes ........................... 25 2.1 Introduction .................................................................................................. 25 2.1.1 Diversity of grass pea germplasm ................................................................ 25 2.1.2 Breeding of low-ODAP varieties .................................................................. 26 2.1.3 Variation in seed β-L-ODAP concentrations between individual seeds and between growth environments ................................................................... 28 2.1.4 Methods for measuring ODAP concentrations ............................................ 28 2.2 Materials and methods .................................................................................. 32 2.2.1 Germplasm ................................................................................................... 32 2.2.2 Production of seed material for germplasm assays .................................... 34 2.2.3 Comparison of extraction media ................................................................. 34 2.2.4 Sensitivity and linearity testing of the plate-based spectrophotometric assay ............................................................................................................. 35 2.2.5 Spectrophotometric assay for ODAP concentrations in individual seeds ... 36 2.2.6 Spectrophotometric assay for ODAP in bulk seed samples ......................... 37 2.2.7 Calculation of ODAP concentrations from absorbance readings ................ 38 2.3 Results and discussion ................................................................................... 40 2.3.1 Optimisation of the spectrophotometric method ....................................... 40 2.3.2 Seed ODAP variation between batches grown in different conditions ....... 43 2.3.3 ODAP concentrations in seeds of accessions from the IPK population ....... 46 2.3.4 ODAP concentrations in seeds of accessions from the USDA population ... 48 iv 2.3.5 ODAP concentrations in seeds of accessions from the EIAR population ..... 52 2.3.6 Screening for L-2,3-diaminopropionic acid accumulating accessions ......... 59 2.4 Summary....................................................................................................... 60 Chapter 3 – Identification of low-ODAP grass peas from a mutagenised population ....... 61 3.1 Introduction .................................................................................................. 61 3.1.1 Increasing genetic variation by mutagenesis ............................................... 61 3.1.2 Approaches to mutagenesis ........................................................................ 62 3.1.3 Selecting an appropriate screening method................................................ 64 3.1.4 Plant tissue used for the mutant screen and adaptations of the screening method ......................................................................................................... 65 3.1.5 Pathway analysis by mutant screening ........................................................ 67 3.2 Materials and methods .................................................................................. 68 3.2.1 Origin of the mutant population .................................................................. 68 3.2.2 Assessment of mutation density ................................................................. 70 3.2.3 Optimisation of seed scarification ............................................................... 71 3.2.4 Growth conditions and sample preparation for the mutant screen ........... 72 3.2.5 The high-throughput spectrophotometric assay ......................................... 75 3.2.6 Automated data handling ............................................................................ 80 3.2.7 Confirmation of low-ODAP mutants by testing individual seeds ................ 81 3.2.8 Synthesis of 13C-labelled β-L-ODAP .............................................................. 81 3.2.9 Characterisation of low-ODAP mutant lines by mass spectrometry ........... 85 3.2.10 Genetic analysis by crossing ........................................................................ 90 3.3 Results and discussion ................................................................................... 92 3.3.1 Assessment of the mutant population derived from LSWT11 .................... 92 3.3.2 Optimisation of the mutant screening method ........................................... 93 3.3.3 Selection of mutants relative to their plate ................................................. 96 3.3.4 Scarification and priming ............................................................................. 97 3.3.5 Data handling and selection of putative mutants ....................................... 98 v 3.3.6 Low-toxin mutants identified ...................................................................... 98 3.3.7 Confirmation of low-ODAP mutants ............................................................ 99 3.3.8 Synthesis of a heavy-isotope-labelled internal standard for LCMS ........... 105 3.3.9 Confirmation and characterisation of mutants ......................................... 111 3.3.10 Accumulation of L-DAP in low-ODAP mutants .......................................... 116 3.3.11 Genetic analysis ......................................................................................... 117 3.3.12 Other mutant phenotypes ......................................................................... 130 3.4 Summary..................................................................................................... 133 Chapter 4 – Identification of candidate genes encoding metabolic enzymes in the β-L- ODAP biosynthetic pathway ....................................................................................... 134 4.1 Introduction ...............................................................................................
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