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Metabolite Profiling • It is estimated that plants as a group synthesize > 400,000 different compounds • This is similar in magnitude to the compound libraries accumulated by chemical companies • Green chemistry vs. non-renewable chemistry • Metabolic profiling as a way to understand gene action MUPGRET workshop, Columbia, MO, June 2005 (HJ Bohnert, UIUC) [email protected] Scaling of the metabolite problem what is the objective – what is the goal. Genomics … not just genes genome & transcriptome sequences markers & QTLs ATCCGAAGCG CTTGGAAAA protein interaction B maps X Y biochemical genetics expression profiles knock-out sRNA & RNAi A Databases, Integration & Intuition protein localization dynamic metabolite catalogs TP Mal structure analysis information mining, hypotheses, experiment - insight, application, virtual life How (much) will ‘encyclopedic’ approaches lead to better understanding? http://www.genome.jp/kegg/ Soybean Fields in Central Illinois Worldwide supposedly the largest man-made Agro-Ecosytem Millions of acres of soybean-corn rotation Free Air Concentration Environment 16 rings 20m diameter ---------4 ambient 4 high CO2 4 high Ozone 4 CO2 + O3 Modeling how the earth’ atmosphere will be in 2050 SoyFACE, UIUC SoyFACE Developmental & diurnal leaf-sampling schedule - 2004 developmental June 30 Early morning Ambient Plant # 1 Ring # 1 Ring # 2 Plant # 2 Plant # 3 Ring # 3 Elevated ozone July 14 Aug 05 noon Elevated carbon-dioxide early night Aug 24 Sept 9 diurnal Combination Ring # 4 No plant is sampled twice. The three leaf-discs are taken from the same (first fully-open) leaflet (5 sampling days/year) x (3 times/day) x (4 treatments) x (4 rings/treatment) x (3 samples/ring) = 720 samples (3 discs each) Total # of GC-MS injections = 720 x 3 injections/sample = 2,160 injections Total GC-MS time/ year = 2, 160 x 1hr/injection = 270 days (at 8 inj./day) Plus standards Metabolite Analyses Harvest treated plant leaves Store in liquid Nitrogen until analysis Extraction (in methanol/water/chloroform mixture) Polar Phase separation (polar and apolar/lipid phases) Apolar Derivatization (GC: to increase volatility, LC: for UV detectability) GC/MS or LC/MS Comparison with spectral libraries for identification/structure determination LC-MS (for unknown metabolites after GC-MS) Derivatization of metabolites, adds side groups at reactive hydroxy-, carboxy- or keto-residues (for example) to make Them identifiable (count number of additions), and renders metabolites more volatile. The derivatized mix is then heated and separated on (a) column(s) in the GC (gas chromatograph) unit of the instrument …… Derivatization reagents 1 step: Methoxyamine Hydrochloride CH3ONH2 · HCl 2 step: N-Trimethylsilyl-N-methyltrifluoroacetamide (MSTFA) CF3CON(CH3)Si(CH3)3 … & each compound identified by a precise elution time before injection to get molecular masses of degradation products HP5890 gas chromatograph and HP5970 mass selective detector A b u n d a n c e volume/weight Abundance T I C : T H - plants F 2 .D Thellungiella 1 .3 e + 0 7 Species A A typical chromatogram 1 .2 e + 0 7 1 .1 e + 0 7 1 e + 0 7 0 0 0 0 0 0 standard hexoses 0 0 0 0 0 0 complex sugars 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 watch here 0 0 0 0 0 0 0 0 0 0 0 0 acids 0 0 0 0 0 0 0 0 0 0 0 0 1 0 .0 1 0 5 .0 2 0 0 .0 2 0 5 .0 3 0 0 .0 3 0 5 .0 4 0 0 .0 4 0 5 .0 5 0 0 .0 5 0 5 .0 6 0 0 .0 0 T TIME im e > --> Abundance 1 .1 e + 0 7 T I C :plants C O I -F 2 . D Arabidopsis Species B 1e+07 9000000 8000000 standard 7000000 6000000 5000000 4000000 watch here 3000000 2000000 1000000 T im e --> 1 0 . 0 01 5 . 0 02 0 . 0 02 5 . 0 03 0 . 0 03 5 . 0 04 0 . 0 04 5 . 0 05 0 . 0 05 5 . 0 06 0 . 0 0 HPLC may be used to enrich certain regions of an elution spectrum (High Performance Liquid Chromatography) HPLC Waters 2690 system Abundance Thellungiella plants T I C : T H -F 2 . D 1400000 1200000 1000000 800000 Glutamine 1600000 Threonic acid 1800000 Glutamic acid 2000000 Ascorbic acid Pyroglutamic acid 2200000 Citric acid Fructose 2400000 Phenylalanine 2600000 Malic acid 2800000 600000 400000 T im e --> 2 2 . 0203 . 0204 . 0205 . 0206 . 0207 . 0208 . 0209 . 0300 . 0301 . 0302 . 0303 . 0304 . 0 0 Focusing on a single compound get mass spectra Consult existing libraries, think, get (or synthesize) a standard A b u n d a n c e T IC : O X A L A C 2 .D 1 .3 e + 0 7 1 .2 e + 0 7 Oxalacetic acid 1 .1 e + 0 7 1 e + 0 7 0 0 0 0 0 0 0 0 0 0 0 0 Internal standard 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 .0 1 0 5 .0 2 0 0 .0 2 0 5 .0 3 0 0 .0 3 0 5 .0 4 0 0 .0 4 0 5 .0 5 0 0 .0 5 0 5 .0 6 0 0 .0 0 T im e - - > Abundanc e T IC: T R E -6 -P .D 1 .2 e + 0 7 1e+07 Trehalose-6-phosphate 8000000 Internal standard 6000000 4000000 2000000 T ime --> 1 0 .0 01 5 .0 02 0 .0 02 5 .0 03 0 .0 03 5 .0 04 0 .0 04 5 .0 05 0 .0 05 5 .0 06 0 .0 0 Resolution of GC/MS instruments Elution times 18.229 min (malate) vs. 18.249 min (GABA) 18.191 min (unknown) Metabolite Profiling – uses • verifying gene expression analyses • control of transgenic (plant) lines (bacteria, yeast, animals) • monitor the effects of mutant proteins • detection of natural variation in species, ecotypes, breeding lines or individuals • detection of new substances Abiotic stress - accumulating metabolites N+ CH2 OH COO - Glycine betaine S+ COO OH HO H Proline COO - H2 COH OH O OH OH HN CH 3 Trehalose 3-dimethylsulfoniopropionate N O O - H OH D-ononitol (+) H COO- N H CH2OH OH OH OH H Ectoine OH OH OH OH CH3 O H OH OH CH2OH Sorbitol CH2OH OH OH OH OH CH2OH Mannitol Sucrose Sucrose-6P Glucose Fructose UDP UDP F6P HK FK TP Fructose UDP-G PPi Tps1 Trehalose-6P UTP G1P Inps (INO) myo-inositol-1P Tpp Imp Trehalose myo-inositol trehalase Imt1 Glc + Glc-P Glycolysis / Krebs cycle D-ononitol G6P F6P F1,6P2 S6pdh Sorbitol-6P Sorbitol Mtldh Mannitol-1P Mannitol leaf/root ratio, emergence seedling growth, leaf size, branching, flowering, seed set DnaJ, HSP,chaperonines, peptidyl-prolyl cis-trans isomerases Chaperone Action Development & Timing interaction & stabilization, LEAs dehydrins carotenoids, pigments, thioredoxins, ascorbateglutathione cycle SOD, ASX, catalase Cold cuticular wax, trichomes, bladder cells, glands cell wall changes, lignification Radical Scavenging Sensing & Signaling abiotic environmental stress Sensing & Signaling Surface Properties Proteome Remodeling Dehydration Sensing & Signaling Stress Proteins proteases, ubiquitin, protease inhibitors Salinity Heat Sensing & Signaling Pathway Adjustment carbohydrate flux, photosynthate partitioning C/N-adjustments, redox status, growth regulator synthesis Hog1- Osmolyte Synthesis & Turnover type glycine betaine, polyols, Proline and other amino acids, ectoine, trehalose, fructans, raffinose, sucrose Calcineurintype Ion Homeostasis Proton pumps/ pyrophosphatase K+-channels/transporters (AKT,HAK,HKT), Na+/H+-antiporters, Na+/hexose symporters, water channels, general cation/anion channels/transporters, ABC-transporters O2 H2O2 Apoplast O2- Ascorbate Oxidations O2 - O2 Oxidase Cytosol Signals 2H+ SOD Plasma membrane Cell wall Metabolism NADP NADPH Ca2+ Import MDAR O 2- Aquaporin O 2 Channels Ca2+ H2O2 MDHA Ascorbate Electron Transport System NAD(P) NAD(P)H GSSG NADPH GSH NADP MDHA Christine Foyer, 2002 Signals DHA Signals An example: Determine metabolites in a pathway that has been indicated by analyzing gene expression. A prominent role for the CBF cold response pathway in configuring the low-temperature metabolome of Arabidopsis Daniel Cook*†, Sarah Fowler*, Oliver Fiehn‡, and Michael F. Thomashow*§¶ *Michigan State University–Department of Energy Plant Research Laboratory and §Department of Crop and Soil Sciences, Michigan State University, East Lansing, MI 48824; and ‡Max Planck Institute of Molecular Plant Physiology, 14424 Potsdam, Germany Cook et al., 2004 – a summary Using GC/MS to determine genetic variation and responses to the environment MAPMAN http://www.genome.jp/kegg/ http://www.genome.jp/kegg/pathway/map/map00020.html An example: What happens in primary metabolism under sulfur Deficiency? Nikiforova et al., May 2005 Importing data into Pathways – biochemical, developmental, regulatory Mapman The three foundations for a complete (?) understanding of the genotype KEGG is expanding to include pathways in human (inherited) diseases mutant lines different species bacterial – animal – fungal - plant Metabotype? As distinct as the genotype with specific, dynamic reactions Arabidopsis thaliana to pathogen state and environmental conditions