A Mutant of Arabidopsis Lacking a Chloroplast Specific Lipid. By Browse J McCourt P Somerville CR
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1 A Mutant of Arabidopsis Lacking a Chloroplast Specific Lipid By Browse J McCourt P Somerville CR
2 What are lipids? (Bile Salt)
3 Lipid classification lipids Storage lipids Membrane lipids Fatty acids eg. Palmitic acid Trans C16:1 Triacylglycerols Waxes Glycerol phospholipids eg. Phosphatidyl glycerol Sphingolipids Cholesterol Other functions Nutrition: Vitamins Signaling: Steroid hormones Eicosanoids Emulsification: Bile salt
4 Lipid classification lipids Storage lipids Fatty acids eg. Palmitic acid Trans C16:1 Triacylglycerols Waxes Hydrophobic Fatty acyl chain Hydrophilic Carboxylate Palmitic acid ( C16:0) 3 (Trans- C16:1 Δ3 t)
5 Lipid classification lipids Membrane lipids Glycerol phospholipids eg. Phosphatidyl glycerol Sphingolipids Cholesterol Head group Phosphatidyl glycerol (PG) R1 = saturated FA R2 = saturated/unsaturated FA
6 Phosphatidyl glycerol (PG) CH2 OH CHOH CH2
7 Lipid classification lipids Storage lipids Membrane lipids Fatty acids eg. Palmitic acid Trans C16:1 Triacylglycerols Waxes Glycerol phospholipids eg. Phosphatidyl glycerol Sphingolipids Cholesterol Other functions Nutrition: Vitamins Signaling: Steroid hormones Eicosanoids Emulsification: Bile salt
8 Fatty acids Fatty acids Chain Length Degree of unsaturation Short chain =< 6C Medium chain 8 12C Long chain 14-18C Very long chain >= 20C Saturated - Straight chain Unsaturated double bond (s) ie. Cis vs. Trans H H H H
9 Fatty acids Packed acyl chains, Semicrystalline -High energy required to separate -Less fluidity Less packed, Liquid crystalline -Less energy required to separate -More fluidity
10 Fluid Mosaic Model of Plasma Membrane
11 Unsaturation on plasma membrane Packed acyl chains, Semicrystalline -High energy required to separate -Less fluidity / trans -Less packed, Liquid crystalline Cis -Less energy required to separate -More fluidity
12 Major hypotheses about compositions of membrane lipids before this paper: 1. Lipid unsaturation increases chilling resistance 2. Lipid unsaturation differences in thermal tolerance between species 3. Trans unsaturation photosynthetic functions Chilling (~ 0-12 C) causes: -Reduced growth and development -Phase transition from the liquid crystalline state to the semicrystalline So, elevated level of cisunsaturation to maintain membrane fluidity in some species, such as Arabidopsis.
13 Chromatography background Thin-layer chromatography (TLC) Gas-liquid chromatography (GC)
14 TLC Mobile phase: solvent Stationary phase: a sheet of glass coated with silica gel Less polar More polar Silica gel
15 Thin-layer chromatography Rf value Silica gel
16 Thin-layer chromatography Recycle the component we want Silica gel degreaser
17 Gas-liquid chromatography Mobile phase: gas Stationary phase: a high boiling point liquid absorbed onto a solid
18 Gas-liquid chromatography retention time TB Solubility
19 TLC Advantages: More efficient (time and money) Recycle the sample GC Advantages: Easy to use get the amount of the sample Disadvantage Detector is limited Disadvantage Destroy the sample
20 Mutant What type of mutant? Alterations in leaf membrane fatty acyl composition.
21 Material Arabidopsis thaliana (L.) Heynh
22 0.3% EMS (Ethyl methanesulfonate) 16h X 200 μ E/m2/sec 23 70%humidity GC
23 Gas-liquid chromatography Column: 2m % diethylene glycol succinate Detector: flame ionization detector
24 recessive? heritable? X One gene? X 0.3% EMS 16h X X genetic complementation X 200 μ E/m2/sec 23 70%humidity GC
25
26
27 Result No difference of Morphology No difference of growth rate Fatty acid composition changes JB60: no trans-c16:1
28 A chloroplast-specific mutant Biochemical Phenotypes: - Lacks trans-c16:1 - Elevated level of C16:0 A single nuclear mutation?
29 A chloroplast-specific mutant P: JB60 Wild type No trans-c16:1 X Trans-C16:1 F1: ½ amount of trans-c16:1 -codominant -suggesting single nuclear gene
30 A chloroplast-specific mutant P: JB60 Wild type No trans-c16:1 X Trans-C16:1 F1: ½ amount of trans-c16:1 F2: trans-c16:1 no trans-c16:1 3 : 1 So, Single nuclear mutant -Mendelian ratio for one gene
31 A chloroplast-specific mutant What is unusual about trans-c16:1? 1. Trans-configuration 2.Delta-3 double bond 3. It only attaches to position two of PG!! => compare WT-PG vs. Mut-PG
32 Fig. 1 FA composition of PG Wild type Mutant C16:0 34% C16:0 < 34% Trans -C16:1 20% Trans- C16:1 0%
33 Table 1. FA composition of leaves
34 A chloroplast-specific mutant Proposal: a desaturase specifically converts C16:0 at position two of PG to trans-c16:1 is involved However no gene was cloned by that time Name the locus: fada (=FAD4)
35
36
37 Thylakoid membrane PS II LHC II trimer Appressed membrane of grana Stroma S
38 Is fada a chloroplast-encoded gene? Exp 1. mutant in chloroplast ribosome No trans-c16:1 Exp 2. add chloroplast protein synthesis inhibitor No trans-c16:1 Exp 3. Mendelian segregation => Not supportive evidence
39 Other evidence Trans-C16:1 Location Etiolated tissue Light-induced chloroplast development
40 LHCP Why are we focusing on the association of LHCP (Light-harvesting chlorophyll a/b protein complex) with trans-c16:1? light-induced chloroplast development The structure (trimer): each subunit associates with trans-c16:1-pg
41 The main function of LHCP Contribute to formation of the grana Enhance the capture of light energy
42 the thylakoid ultrastructure Wild type mutant
43 The main function of LHCP Contribute to formation of the grana Enhance the capture of light energy
44 The effects of light intensity PS ΙΙ PS Ι
45 Sensitive fluorescence techniques Aim: Extensive analysis of energy transfer from LHCP to photosystem ΙΙ Result: NO functional difference
46 Since FAD4 (2009) Specials about trans-c16:1 of PG -only found in thylakoid membranes -always esterified specifically to the sn-2 position of the glycerol backbone of PG -found exclusively in eukaryotic, Chl a/b-containing photoautotrophs Functions of trans-c16:1 of PG -phosphatidylglycerol (PG) and its fatty acid composition play a crucial role in stabilizing the oligomeric state of LHCII (1983). -not involved in energy transfer (1985) -the apparent stability of oligomeric LHCII is dependent not on PG content, but rather on the molecular species composition of PG such that oligomeric LHCII was stabilized when thylakoids exhibit high levels of PG 16:0/trans-Δ3-hexadecenoic acid (trans-16:1) relative to PG 16:0/16:0 (1998) Formation of trans-c16:1 of PG -by a trans-desaturase located within the inner recesses of the thylakoid compartment. (1991) Gene cloned
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