Very-Long Chain Fatty Acid Biosynthesis
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1 Very-Long Chain Fatty Acid Biosynthesis Objectives: 1. Review information on the isolation of mutants deficient in VLCFA biosynthesis 2. Generate hypotheses to explain the absence of mutants with lesions in the 2 reductase and the dehydrase genes 3. Generate a model for the organization and control of VLCFA biosynthesis in plants 4. Design experiments to test the model
2 Organization of FAS Enzymes TYPE I (Animals and Fungi) Location: Cytosol Organization: A complex of 2 large subunits (250 kda each) catalyzing all the reactions of FAS KAS ER KR DH TYPE II (Plants and Bacteria) Location: Plastid stroma Organization: Enzyme activities reside on individual proteins this reflects prokaryotic origin of plastids KAS ER KR DH
3 Fatty Acid Synthesis ACC KASIII TA KAS I KR ER DH Ohlrogge and Browse (1995) Plant Cell 7:795
4 Fatty Acid Synthesis FAS Chloroplast lipids AT FATB 16:0 16:0-ACP AT KASII 18:0-ACP FATB DES AT FATA 18:1 18:1-ACP 16:0-CoA LACS 18:0 18:0-CoA LACS 18:1-CoA LACS FAS = fatty acid synthesis FAT = acyl-acp thioesterase LACS = acyl-coa synthetase AT = acyl transferase DES = stearoyl-acp desaturase Main products of FAS are 16:0 and 18:1
5 Main products of FAS are 16:0 and 18:1 AT FAT Lipid synthesis in the chloroplast Export to the ER How do we know that? From ( 14 C)-acetate labeling experiments of isolated plastids in vitro C16 and C18 fatty acyl chains get radioactively labeled What happens when we add isolated microsomes to the radioactively labeled C16 and C18 fatty acids in chloroplasts?
6 Microsomes = small ER derived vesicles nm in diameter
7 SER RER
8 Microsomes = small ER derived vesicles What happens when we add isolated microsomes to the labeled C16 and C18 fatty acids in chloroplasts?
9 TLC analysis of fatty acid elongation products after addition of leek microsomes Radioactively labeled very long chain fatty acids (>C18) get made, such as C20, C22 Evenson and Post-Beitenmiller,1995
10 Very-Long Chain Fatty Acid Biosynthesis Hypothesis: VLCFA synthesis beyond C 18 occurs in which the ER. cellular compartment? Prediction: Each step of fatty acyl chain elongation during VLCFA synthesis requires at least? reactions.
11 Fatty Acid Synthesis Ohlrogge and Browse (1995) Plant Cell 7:795
12 Very-Long Chain Fatty Acid Biosynthesis Hypothesis: VLCFA synthesis beyond C 18 occurs in which the ER. cellular compartment? Prediction: Each step of fatty acyl chain elongation during VLCFA synthesis requires at least 4 reactions.
13 Model of Very-Long Chain Fatty Acid PLASTID Biosynthesis Fatty Acid Synthase (FAS) ER C16-ACP C18-ACP REDUCTION C16-CoA C18-CoA CONDENSATION Fatty Acid Elongase (FAE) DEHYDRATION REDUCTION VLCFAs C16 C18 C20 C22 C24 C26 C28 C30 C32 C34
14 PLASTID Fatty Acid Synthase (FAS) C16-ACP C18-ACP Model of Very-Long Chain Fatty Acid Biosynthesis REDUCTION C16-CoA C18-CoA Malonyl-CoA KCS CONDENSATION Fatty Acid Elongase (FAE) REDUCTION DEHYDRATION VLCFAs All cells Seeds Root C24 C20, C22 C20 to C32 Sphingolipids Triacylglycerols Suberin Epidermis C24 to C34 Waxes
15 Testing the Model of Very-Long Chain Fatty Acid Biosynthesis How? Isolation of mutants. What kind of phenotype would you look for? Absence of VLCFA in lipids. Which lipids?
16 Testing the Model of Very-Long Chain Fatty Acid Biosynthesis How? Isolation of mutants. What kind of phenotype would you look for? Absence of VLCFA in lipids. Which lipids? Seed lipids. How? By gas chromatograph with a flame ionization detector (FID).
17 Preparation of Fatty Acid Samples for Gas Chromatography 1. Place seeds from each sample to a glass tube. 2. Add 0.5 ml 1 N methanolic-hcl (Supelco) to each tube and cap tubes TIGHTLY. 3. Place samples at 80 o C for 2-3 hrs. 4. Cool down (5 minutes). 5. Add 0.5 ml NaCl (0.9%) and 300 µl hexane. 6. Recap and vortex vigorously to extract fatty acid methyl esters into hexane. 7. Carefully draw µl hexane (top phase) and inject into gas chromatograph
18 Gas Liquid Chromatography The mixture to be analyzed is injected into the stream of carrier gas. As it passes along the column, it gets separated into the different compounds. Compounds with a greater affinity for the mobile (gas) phase reach the detector at the end of the column more quickly. Compounds with a greater affinity for the stationary (liquid) phase move more slowly through the column.
19 Analyzing Gas Chromatograms Three main pieces of information can be obtained from a gas chromatogram: the number of compounds in the mixture - represented by the number of peaks how much of each compound is present - represented by the height of the peak (higher = more) the retention time - indicated by the position of the peak (can be compared to reference standard)
20 Analyzing Gas Chromatograms This gas chromatogram shows that: substance A was present in the smallest quantity (it has the smallest peak) substance A had the shortest retention time (shortest carbon chain, no double bonds, eg. 16:0) substances B and C were present in equal amounts substance F had the longest retention time (eg. 18:3) substance F was present in the greatest quantity (it has the largest peak) substance F had the greatest affinity for the stationary phase (and therefore longest retention time)
21 Fae1 Phenotype Fae1 WT
22 Fae1 Phenotype Fatty acid Wild type Fae1 mutant 16: : : : : : : : Values represent Mol % 18:1-CoA à20:1-CoA à22:1-CoA VLCFA = Chain length > C20
23 Summary of the Screen for Mutants With Reduced Levels of Seed Lipid VLCFAs ~10,000 mutagenized seed samples (M 3 ) analyzed by GC-FID Why were M 3 seeds used for the screen? M 1 seeds M 1 plants M 2 seeds M 2 plants M 3 seed families WT seeds mutagen Use seeds from each M 3 seed family for mutant isolation by GC-FID analysis Modified from: Harvest from individual M 2 plants
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