#METABOLOMICS WORKBENCH juliehaines_20210702_093259 DATATRACK_ID:2737 STUDY_ID:ST001864 ANALYSIS_ID:AN003022 PROJECT_ID:PR001177
VERSION             	1
CREATED_ON             	July 17, 2021, 11:15 am
#PROJECT
PR:PROJECT_TITLE                 	Targeting host glycolysis as a strategy for antimalarial development
PR:PROJECT_SUMMARY               	Glycolysis controls cellular energy, redox balance, and biosynthesis.
PR:PROJECT_SUMMARY               	Antiglycolytic therapies are under investigation for treatment of obesity,
PR:PROJECT_SUMMARY               	cancer, aging, autoimmunity, and microbial diseases. Interrupting glycolysis is
PR:PROJECT_SUMMARY               	highly valued as a therapeutic strategy, because glycolytic disruption is
PR:PROJECT_SUMMARY               	generally tolerated in mammals. Unfortunately, anemia is a known dose-limiting
PR:PROJECT_SUMMARY               	side effect of these inhibitors and presents a major caveat to development of
PR:PROJECT_SUMMARY               	antiglycolytic therapies. We developed specific inhibitors of enolase – a
PR:PROJECT_SUMMARY               	critical enzyme in glycolysis – and validated their metabolic and cellular
PR:PROJECT_SUMMARY               	effects on human erythrocytes. Enolase inhibition increases erythrocyte
PR:PROJECT_SUMMARY               	susceptibility to oxidative damage and induces rapid and premature erythrocyte
PR:PROJECT_SUMMARY               	senescence, rather than direct hemolysis. We apply our model of red cell
PR:PROJECT_SUMMARY               	toxicity to address questions regarding erythrocyte glycolytic disruption in the
PR:PROJECT_SUMMARY               	context of Plasmodium falciparum malaria pathogenesis. Our study provides a
PR:PROJECT_SUMMARY               	framework for understanding red blood cell homeostasis under normal and disease
PR:PROJECT_SUMMARY               	states and clarifies the importance of erythrocyte reductive capacity in malaria
PR:PROJECT_SUMMARY               	parasite growth.
PR:INSTITUTE                     	University of Colorado Anschutz Medical Campus
PR:LAST_NAME                     	Haines
PR:FIRST_NAME                    	Julie
PR:ADDRESS                       	12801 E 17th Ave, Room 1303, Aurora, Colorado, 80045, USA
PR:EMAIL                         	julie.haines@cuanschutz.edu
PR:PHONE                         	3037243339
#STUDY
ST:STUDY_TITLE                   	Targeting host glycolysis as a strategy for antimalarial development
ST:STUDY_SUMMARY                 	Glycolysis controls cellular energy, redox balance, and biosynthesis.
ST:STUDY_SUMMARY                 	Antiglycolytic therapies are under investigation for treatment of obesity,
ST:STUDY_SUMMARY                 	cancer, aging, autoimmunity, and microbial diseases. Interrupting glycolysis is
ST:STUDY_SUMMARY                 	highly valued as a therapeutic strategy, because glycolytic disruption is
ST:STUDY_SUMMARY                 	generally tolerated in mammals. Unfortunately, anemia is a known dose-limiting
ST:STUDY_SUMMARY                 	side effect of these inhibitors and presents a major caveat to development of
ST:STUDY_SUMMARY                 	antiglycolytic therapies. We developed specific inhibitors of enolase – a
ST:STUDY_SUMMARY                 	critical enzyme in glycolysis – and validated their metabolic and cellular
ST:STUDY_SUMMARY                 	effects on human erythrocytes. Enolase inhibition increases erythrocyte
ST:STUDY_SUMMARY                 	susceptibility to oxidative damage and induces rapid and premature erythrocyte
ST:STUDY_SUMMARY                 	senescence, rather than direct hemolysis. We apply our model of red cell
ST:STUDY_SUMMARY                 	toxicity to address questions regarding erythrocyte glycolytic disruption in the
ST:STUDY_SUMMARY                 	context of Plasmodium falciparum malaria pathogenesis. Our study provides a
ST:STUDY_SUMMARY                 	framework for understanding red blood cell homeostasis under normal and disease
ST:STUDY_SUMMARY                 	states and clarifies the importance of erythrocyte reductive capacity in malaria
ST:STUDY_SUMMARY                 	parasite growth.
ST:INSTITUTE                     	University of Colorado Anschutz Medical Campus
ST:LAST_NAME                     	Haines
ST:FIRST_NAME                    	Julie
ST:ADDRESS                       	12801 E 17th Ave, Room 1303, Aurora, Colorado, 80045, USA
ST:EMAIL                         	julie.haines@cuanschutz.edu
ST:PHONE                         	3037243339
#SUBJECT
SU:SUBJECT_TYPE                  	Cultured cells
SU:SUBJECT_SPECIES               	Homo sapiens
SU:TAXONOMY_ID                   	9606
#SUBJECT_SAMPLE_FACTORS:         	SUBJECT(optional)[tab]SAMPLE[tab]FACTORS(NAME:VALUE pairs separated by |)[tab]Raw file names and additional sample data
SUBJECT_SAMPLE_FACTORS           	-	2	group:Control, 0h	RAW_FILE_NAME=2
SUBJECT_SAMPLE_FACTORS           	-	4	group:Control, 0h	RAW_FILE_NAME=4
SUBJECT_SAMPLE_FACTORS           	-	6	group:Control, 0h	RAW_FILE_NAME=6
SUBJECT_SAMPLE_FACTORS           	-	20	group:POMSF, 0h	RAW_FILE_NAME=20
SUBJECT_SAMPLE_FACTORS           	-	22	group:POMSF, 0h	RAW_FILE_NAME=22
SUBJECT_SAMPLE_FACTORS           	-	24	group:POMSF, 0h	RAW_FILE_NAME=24
SUBJECT_SAMPLE_FACTORS           	-	26	group:POMHEX, 0h	RAW_FILE_NAME=26
SUBJECT_SAMPLE_FACTORS           	-	28	group:POMHEX, 0h	RAW_FILE_NAME=28
SUBJECT_SAMPLE_FACTORS           	-	30	group:POMHEX, 0h	RAW_FILE_NAME=30
SUBJECT_SAMPLE_FACTORS           	-	32	group:Control, 0.5h	RAW_FILE_NAME=32
SUBJECT_SAMPLE_FACTORS           	-	34	group:Control, 0.5h	RAW_FILE_NAME=34
SUBJECT_SAMPLE_FACTORS           	-	36	group:Control, 0.5h	RAW_FILE_NAME=36
SUBJECT_SAMPLE_FACTORS           	-	50	group:POMSF, 0.5h	RAW_FILE_NAME=50
SUBJECT_SAMPLE_FACTORS           	-	52	group:POMSF, 0.5h	RAW_FILE_NAME=52
SUBJECT_SAMPLE_FACTORS           	-	54	group:POMSF, 0.5h	RAW_FILE_NAME=54
SUBJECT_SAMPLE_FACTORS           	-	56	group:POMHEX, 0.5h	RAW_FILE_NAME=56
SUBJECT_SAMPLE_FACTORS           	-	58	group:POMHEX, 0.5h	RAW_FILE_NAME=58
SUBJECT_SAMPLE_FACTORS           	-	60	group:POMHEX, 0.5h	RAW_FILE_NAME=60
SUBJECT_SAMPLE_FACTORS           	-	62	group:Control, 1h	RAW_FILE_NAME=62
SUBJECT_SAMPLE_FACTORS           	-	64	group:Control, 1h	RAW_FILE_NAME=64
SUBJECT_SAMPLE_FACTORS           	-	66	group:Control, 1h	RAW_FILE_NAME=66
SUBJECT_SAMPLE_FACTORS           	-	80	group:POMSF, 1h	RAW_FILE_NAME=80
SUBJECT_SAMPLE_FACTORS           	-	82	group:POMSF, 1h	RAW_FILE_NAME=82
SUBJECT_SAMPLE_FACTORS           	-	84	group:POMSF, 1h	RAW_FILE_NAME=84
SUBJECT_SAMPLE_FACTORS           	-	86	group:POMHEX, 1h	RAW_FILE_NAME=86
SUBJECT_SAMPLE_FACTORS           	-	88	group:POMHEX, 1h	RAW_FILE_NAME=88
SUBJECT_SAMPLE_FACTORS           	-	90	group:POMHEX, 1h	RAW_FILE_NAME=90
SUBJECT_SAMPLE_FACTORS           	-	122	group:Control, 6h	RAW_FILE_NAME=122
SUBJECT_SAMPLE_FACTORS           	-	124	group:Control, 6h	RAW_FILE_NAME=124
SUBJECT_SAMPLE_FACTORS           	-	126	group:Control, 6h	RAW_FILE_NAME=126
SUBJECT_SAMPLE_FACTORS           	-	140	group:POMSF, 6H	RAW_FILE_NAME=140
SUBJECT_SAMPLE_FACTORS           	-	142	group:POMSF, 6H	RAW_FILE_NAME=142
SUBJECT_SAMPLE_FACTORS           	-	144	group:POMSF, 6H	RAW_FILE_NAME=144
SUBJECT_SAMPLE_FACTORS           	-	146	group:POMHEX, 6H	RAW_FILE_NAME=146
SUBJECT_SAMPLE_FACTORS           	-	148	group:POMHEX, 6H	RAW_FILE_NAME=148
SUBJECT_SAMPLE_FACTORS           	-	150	group:POMHEX, 6H	RAW_FILE_NAME=150
#COLLECTION
CO:COLLECTION_SUMMARY            	Banked blood (Saint Louis Children’s Hospital) and fresh blood [healthy
CO:COLLECTION_SUMMARY            	donors] was washed and stored at 50% hematocrit at 4ºC up to 1 month past the
CO:COLLECTION_SUMMARY            	clinical expiration date and collection date, respectively. Parasite strains
CO:COLLECTION_SUMMARY            	provided by the Malaria Research and Reference Reagent Resource Center (MR4) as
CO:COLLECTION_SUMMARY            	follows: 3D7 (MRA-102); K1 (MRA-159); D10 (MRA-201); and IPC-5202 (MRA-1240).
CO:COLLECTION_SUMMARY            	Unless indicated, erythrocytes were cultured as previously described at 2%
CO:COLLECTION_SUMMARY            	hematocrit in complete media (incomplete media supplemented with 27mM sodium
CO:COLLECTION_SUMMARY            	bicarbonate, 11mM glucose, 5mM HEPES, 1mM sodium pyruvate, 0.37mM hypoxanthine,
CO:COLLECTION_SUMMARY            	0.01mM thymidine, 10ug/mL gentamicin, and 0.5% Albumax (Thermo-Fisher) under 5%
CO:COLLECTION_SUMMARY            	O2/5% CO2/90% N2 atmosphere at 37ºC (71).
CO:SAMPLE_TYPE                   	Blood (whole)
#TREATMENT
TR:TREATMENT_SUMMARY             	Freshly collected erythrocytes were washed, resuspended to 10% hematocrit in
TR:TREATMENT_SUMMARY             	buffer [25mM HEPES (pH 7.4), 120mM NaCl, 5.4mM KCl, 1.8mM CaCl2, and 1mM
TR:TREATMENT_SUMMARY             	NaH2PO4], incubated for 1 hour at 37ºC, then centrifuged at 2000xg for 5
TR:TREATMENT_SUMMARY             	minutes at 4ºC, resuspended in fresh wash buffer to 30-40% hematocrit, and
TR:TREATMENT_SUMMARY             	split into 210uL aliquots of packed erythrocytes. Erythrocytes were resuspended
TR:TREATMENT_SUMMARY             	in 253uL of RPMI containing 11.9mM D-[1,2,3-13C] glucose (Millipore-Sigma) and
TR:TREATMENT_SUMMARY             	either treated with POM-SF or POM-HEX at five times the EC50s for MetHb
TR:TREATMENT_SUMMARY             	formation, or an untreated solvent control with the balance volume to 550uL
TR:TREATMENT_SUMMARY             	using washing buffer. All samples were incubated at 37ºC shaking at 500 RPM.
TR:TREATMENT_SUMMARY             	Triplicate samples of supernatants and packed erythrocytes were collected at 0,
TR:TREATMENT_SUMMARY             	0.5, 1, and 6-hour intervals and snap-frozen in liquid N2.
#SAMPLEPREP
SP:SAMPLEPREP_SUMMARY            	RBC and media samples were extracted at 1:10 and 1:20 dilutions, respectively,
SP:SAMPLEPREP_SUMMARY            	in ice-cold 5:3:2 MeOH:MeCN:H2O v/v/v, with vigorous vortexing at 4 degrees C
SP:SAMPLEPREP_SUMMARY            	for 30 min. The extraction buffer was supplemented with 40uM 3-13C lactate to
SP:SAMPLEPREP_SUMMARY            	allow absolute quantification of lactate isotopologues, including lactate
SP:SAMPLEPREP_SUMMARY            	generated via glycolysis ([1,2,3-13C3] enriched) vs. the pentose phosphate
SP:SAMPLEPREP_SUMMARY            	pathway ([2,3-13C2] enriched). Supernatants were clarified via centrifugation
SP:SAMPLEPREP_SUMMARY            	(10,000 g, 10 min, 4 C) then analyzed by UHPLC-MS.
#CHROMATOGRAPHY
CH:CHROMATOGRAPHY_SUMMARY        	10 μl of RBC extracts were injected and separated through a 3 min isocratic
CH:CHROMATOGRAPHY_SUMMARY        	elution on a Kinetex XB-C18 column (150 × 2.1 mm i.d., 1.7 μm particle size
CH:CHROMATOGRAPHY_SUMMARY        	– Phenomenex, Torrance, CA, USA) at 250 μl/min (mobile phase: 5%
CH:CHROMATOGRAPHY_SUMMARY        	acetonitrile, 95% 18 mΩ H2O, 0.1% formic acid; column temperature: 25°C).
CH:CHROMATOGRAPHY_TYPE           	Reversed phase
CH:INSTRUMENT_NAME               	Vanquish
CH:COLUMN_NAME                   	Kinetex XB-C18 2.1 x 150 mm, 1.7 um
CH:FLOW_GRADIENT                 	isocratic
CH:FLOW_RATE                     	250 ul/min
CH:COLUMN_TEMPERATURE            	25
#ANALYSIS
AN:ANALYSIS_TYPE                 	MS
#MS
MS:INSTRUMENT_NAME               	Thermo Q Exactive Orbitrap
MS:INSTRUMENT_TYPE               	Orbitrap
MS:MS_TYPE                       	ESI
MS:ION_MODE                      	POSITIVE
MS:MS_COMMENTS                   	The UHPLC system was coupled online with a QExactive mass spectrometer (Thermo,
MS:MS_COMMENTS                   	San Jose, CA, USA), scanning in Full MS mode (2 μscans) at 70,000 resolution
MS:MS_COMMENTS                   	from 60-900 m/z, with 4 kV spray voltage, 15 sheath gas and 5 auxiliary gas,
MS:MS_COMMENTS                   	operated in positive ion mode. Calibration was performed before each analysis
MS:MS_COMMENTS                   	using a positive calibration mix (Piercenet – Thermo Fisher, Rockford, IL,
MS:MS_COMMENTS                   	USA). Limits of detection (LOD) were characterized by determining the smallest
MS:MS_COMMENTS                   	injected amino acid amount required to provide a signal to noise (S/N) ratio
MS:MS_COMMENTS                   	greater than three using < 5 ppm error on the accurate intact mass. Based on a
MS:MS_COMMENTS                   	conservative definition for Limit of Quantitation (LOQ), these values were
MS:MS_COMMENTS                   	calculated to be three fold higher than determined LODs. MS data acquired from
MS:MS_COMMENTS                   	the QExactive was converted from .raw file format to.mzXML format using
MS:MS_COMMENTS                   	MassMatrix (Cleveland, OH, USA). Amino acid assignments were performed using
MS:MS_COMMENTS                   	MAVEN (Princeton, NJ, USA). The MAVEN software platform provides tools for peak
MS:MS_COMMENTS                   	picking, feature detection and metabolite assignment against the KEGG pathway
MS:MS_COMMENTS                   	database. Assignments were further confirmed using a process for chemical
MS:MS_COMMENTS                   	formula determination using isotopic patterns and accurate intact mass (Clasquin
MS:MS_COMMENTS                   	et al. 2012). Analyte retention times were confirmed by comparison with external
MS:MS_COMMENTS                   	standard retention times, as indicated above.
#MS_METABOLITE_DATA
MS_METABOLITE_DATA:UNITS	peak area
MS_METABOLITE_DATA_START
Samples	2	4	6	20	22	24	26	28	30	32	34	36	50	52	54	56	58	60	62	64	66	80	82	84	86	88	90	122	124	126	140	142	144	146	148	150
Factors	group:Control, 0h	group:Control, 0h	group:Control, 0h	group:POMSF, 0h	group:POMSF, 0h	group:POMSF, 0h	group:POMHEX, 0h	group:POMHEX, 0h	group:POMHEX, 0h	group:Control, 0.5h	group:Control, 0.5h	group:Control, 0.5h	group:POMSF, 0.5h	group:POMSF, 0.5h	group:POMSF, 0.5h	group:POMHEX, 0.5h	group:POMHEX, 0.5h	group:POMHEX, 0.5h	group:Control, 1h	group:Control, 1h	group:Control, 1h	group:POMSF, 1h	group:POMSF, 1h	group:POMSF, 1h	group:POMHEX, 1h	group:POMHEX, 1h	group:POMHEX, 1h	group:Control, 6h	group:Control, 6h	group:Control, 6h	group:POMSF, 6H	group:POMSF, 6H	group:POMSF, 6H	group:POMHEX, 6H	group:POMHEX, 6H	group:POMHEX, 6H
L-alanine	7242486	8912985	7158126	6401872	7707776	9548117	7276286	6965748	9276854	7480886	11017300	8467653	8488872	9986075	10053640	11022420	9016195	11613360	11535820	9231328	11538390	8346966	9138009	11150540	11237570	9082003	10491410	10969840	11443560	11424540	8349172	9189227	8309714	7742060	6765194	7989718
L-arginine	3645470	3443846	9090948	10991800	7604968	2766940	7336580	15088100	4282606	13723380	8684725	19371740	8178514	5942294	8911512	8785005	13727590	10864640	0	13744450	0	10190230	10068300	0	4177597	11876080	3039427	5252256	3209683	0	2946548	0	2669604	4957038	6708256	3717685
L-asparagine	1955497	1718105	1780944	1890954	2563879	2415541	3068902	3142219	2945099	3395971	3729742	3916355	3697817	4013357	4488063	5661364	6039170	5522350	7475794	8221690	7636048	7511366	6724548	7843522	7529444	8128166	7162482	8130102	8022632	7240662	5765030	5602932	5223490	5500968	5875878	6245446
L-aspartate	11091010	13810000	14237900	17090460	22648230	20645570	35189230	32183950	35373890	40054810	31004290	41877040	33760500	36995630	41818210	50653890	46697620	45450190	42812370	34213380	41737220	31242800	32609190	28977270	22781500	22855950	17495780	15131190	16100970	15237630	13088800	12512050	11753010	10782740	11882530	11221290
L-cysteine	164335	15456	21691	753697	333364	73529	410844	441704	192847	372697	617454	876904	561229	255438	157519	389729	984356	384315	839938	1901536	857355	1799691	1833489	569716	256841	1283310	621093	182424	450465	327922	396722	113805	220393	497907	597171	1046938
L-glutamate	11577210	14549840	17442730	18758910	22203190	23167270	39111810	36415790	41567720	47836980	36828700	44181400	32566450	34313870	36771310	36589290	35517710	35454460	37446450	28754190	33860430	31818100	26790660	24948110	21648810	21005850	17337580	15451160	16259690	13287320	7502046	6145646	6743000	5993644	6138418	6664012
L-glutamine	18782980	19098040	20496370	26240550	25889110	22819750	35105340	36487550	27667520	40028650	43453770	53001990	37675980	40447630	51119000	60019230	67716420	70323830	87982600	89650900	87960390	84990440	90075740	82648600	69670420	88069180	65724080	80429540	86487620	75386690	62602440	57636720	54822440	63653390	62558980	63527520
glycine	916092	1229423	1076504	991120	929621	1128718	1292235	1209066	1296391	1330217	1501164	1186888	1559153	1612522	1747246	2207561	2319184	2352487	2332139	2494888	2354494	1774210	2277206	2530172	2528167	2516934	2977812	2173097	1698986	1578259	1742885	1714175	1543826	1509661	1660320	1671999
L-histidine	1033545	1232930	1449159	1619643	1234682	1630573	1769342	1855329	1637418	1876483	1470164	1546089	1391019	1382119	1348738	1306931	1354423	1303530	1393839	1557032	1666253	1586777	1542917	1566890	1220038	1285775	1073028	1638212	1590312	1558378	1455397	1380310	1442633	1601860	1469214	1874259
L-leucine	84864420	98450840	113613900	98293860	121621000	95003460	113213700	129158500	101707700	176780100	145869600	209409200	170340900	183069700	203282700	215707300	281365200	204740400	269772900	174288200	286553600	227143600	253522000	230609800	197049200	284880100	135540700	236095600	182780800	234542100	126729300	158261800	214992900	146707300	146636200	127084400
L-isoleucine	27213850	31808080	18323510	30400470	29503980	22626360	0	27936670	20057970	26536840	18821960	28306620	17875020	16703940	16900170	17042920	15478430	16072940	15411240	9726021	15039160	12721400	12448730	10936780	9186887	13060150	8518843	13770660	14295000	16570240	11665480	13137060	14932190	12783540	14155870	11402200
L-lysine	3740801	3855109	4049775	3758556	3805278	3275392	3764694	3909943	3344321	3449402	2802153	3558222	2697722	2387239	2589775	2545018	2495028	1992931	2587068	2170731	2252804	2401434	2033049	1892300	1462514	1532769	1053325	1685797	1672237	1650726	1431318	1332538	1515191	1470367	1239543	1446493
L-methionine	9200364	11001770	12370150	10420790	12949040	9062012	11956280	12811740	9765501	17111030	13409880	19758990	15214200	16647130	18540080	19536960	25744730	18473870	25072570	16441860	26839330	22400440	23539650	22203540	18061030	29116580	15850100	21473350	19221410	21242210	12699880	14782820	18305190	15566940	16613260	12651970
L-phenylalanine	4000974	4370876	5234788	4951308	5411315	4321240	3891675	5354106	4427160	6541264	5620058	7398644	8078942	8858083	11243140	13667980	17993540	12454130	16507440	11649700	16523760	15787740	14930370	14305940	12185220	15606000	8200482	13037470	11015450	9639966	6889194	7467530	8392055	7055430	6501138	4974045
L-proline	25905060	31503310	37904130	38608690	36174930	32050070	38653320	40424470	27828480	44150680	42732580	61939070	42189960	34470820	39593780	48674620	52245730	38873150	52767020	54651640	76282780	65842720	59039420	53928540	41362770	62461570	46346440	73253990	77275160	85140290	72463830	66349620	67761050	63860800	54607500	59644550
L-serine	1668785	1634940	1541094	1543712	1815681	2047142	2274197	2299246	2071490	2664274	3157992	2801792	2932647	2658852	3792309	4555184	4840247	4454737	5539202	6387166	6063134	4641730	5259884	5997134	6193948	5885706	5910584	7095122	6610254	5793606	4639024	4683864	4281188	4503508	4599316	4861448
L-threonine	2399336	2632629	2622006	2655535	3056179	2691288	3337686	3502021	3234011	4629332	6548448	5786486	5712476	5049590	5980532	6272120	7919814	7084572	9317557	8192858	8955097	8463630	8550655	8853418	7846008	8675493	7029224	10112340	10771460	9588840	6950698	7993730	7078666	8822238	8151734	8771327
L-tryptophan	3875911	2971389	4179942	5640244	3842866	3636155	4610792	3916344	4513774	3918519	5951456	5314482	5933300	3776525	4714087	5024426	4783172	4620149	4634684	5083126	3705227	4996572	4414736	4348400	3997264	3893585	2668211	2242726	2249158	1770914	1426287	1207282	1712412	1850765	1630310	1266923
L-tyrosine	9819055	12384870	13186980	11105300	12673510	8942085	13109670	14429420	10609080	21223430	17103670	24344380	19447910	20923130	21612620	23678360	30120250	23727170	31843290	22086390	34124320	25777940	28736400	27511530	23887520	36199650	22133130	33006780	27436980	30090640	17352420	21653340	24970450	21338880	23044650	16856950
L-valine	94790780	120532000	111798200	91523130	114851800	138536000	124533000	119934100	119285300	130492700	140741900	146753700	133793600	145416600	141754600	170500400	166747300	160342600	154491000	126128600	170886600	153296800	146531900	151010300	159184700	133927300	166485600	143683400	163899700	181770300	173139900	188680000	172792700	159161400	162102000	147152300
AMP	848440	743273	969475	1480470	1217078	1210007	2427259	4082492	2384482	970419	2246575	1143026	2219571	1694654	1241942	2038929	4362031	1738786	506136	866012	921815	2138458	1657126	1771637	1959322	2601012	1218053	399988	294786	274563	502118	408555	585272	372148	497296	488108
Cytosine	50318	41932	36597	44987	51506	127229	85457	238140	474432	468114	353620	577592	561985	581079	575038	636887	552688	672873	584383	473589	416615	367906	333341	322381	475667	342725	266982	129029	96772	48291	80549	41630	51936	0	0	0
Xanthine	353816	331585	325432	284515	363070	338429	386500	378072	391009	337165	337079	288908	366141	357245	328598	339786	331695	309317	273730	191378	224273	230597	191415	189689	187881	142585	148391	120820	114168	93736	82895	98148	92447	78196	72041	83053
Allantoate	215979	291440	395496	591935	419994	147691	362687	439892	225726	384391	359695	572779	271115	203970	255959	252451	384609	239029	433968	494383	499867	665229	503563	314641	127648	405965	101548	555548	781967	678656	544744	305564	473965	578160	591710	852104
5-Hydroxyisourate	3520943	4484736	3342449	2986779	4004029	4097905	4576981	4137426	5159640	5484614	5767334	5158496	5173060	5159936	5684584	6716168	4602454	5109334	3484308	1241391	2658584	1498294	2098099	4214002	4866014	2622047	4928044	3002117	3287091	4557006	4596630	7026874	4569560	4805142	4493092	3269531
Nicotinamide	17695020	22932120	22942710	15687080	23502230	26625030	25644780	26082850	28934970	28186300	29146560	32267610	36853290	39379030	37216570	37370020	42312260	38969670	40452660	30969620	48213360	34616800	37797060	48575360	48045310	50795930	41800080	36360780	28538400	31589450	25434410	31762010	26684450	20164270	21607350	18233590
NAD+	16014	14017	48961	42594	268418	234083	320487	325026	410885	415784	59048	217944	77617	198747	413750	523908	246839	445534	189261	98690	134427	87330	41062	43998	15669	44600	21461	19745	63968	110660	56510	40010	93017	52874	59460	98539
Glutathione	90790380	91277530	105289400	166742000	141376100	96583620	128866300	141526000	145170900	150883600	130681400	184168600	164095600	151838000	144792000	145879400	138207200	122325600	143605500	83234780	124156000	153599500	149582800	91690660	50645600	77735670	57434140	42323210	71690820	67953900	41436760	33306830	33673380	45890000	38679130	43347650
Cys-Gly	1265812	1298274	1476999	2173179	2001217	1458611	1911857	2026439	2129311	2041609	1874078	2805206	2554519	2282035	2127208	2174782	2132634	1970652	2184732	1201590	1961014	2317702	2136714	1316489	757079	1195785	808783	547757	1012563	914813	706261	527098	456492	780780	557222	770028
gamma-Glutamyl-gamma-aminobutyrate	72229	110308	101415	86917	89110	106730	80783	58981	110469	111527	89519	102344	99270	131652	100094	98433	113168	104677	77835	61285	113317	76316	89259	105686	90001	58860	72574	57841	99247	73181	58894	66443	59972	58716	52619	33564
5-L-Glutamyl-taurine	185262	163911	137646	127849	93753	174081	113217	105442	156925	118524	106527	100855	145472	153750	136048	165468	174022	86803	91788	120877	146223	111131	118202	115541	172503	93789	210974	163411	147519	160627	168545	169693	166476	149356	128339	134607
Dimethylglycine	1158315	1119542	1148019	887867	1007648	1451348	1126342	1121192	1220684	1023901	1313678	1124184	1143787	1172701	1056851	1057688	1081250	974504	922661	817410	997550	885609	1019251	895514	1121919	860458	1044726	733963	870545	893963	746152	819752	597643	687291	686462	597891
Phosphoserine	34901	75142	89006	48678	238263	225470	131097	159138	151921	185437	63095	115299	54285	259964	71349	51746	36270	35436	0	0	83926	0	0	34569	0	0	17697	0	0	68283	54102	59064	27359	23248	25682	10410
5-10-Methenyltetrahydrofolate	306445	253138	240917	232563	306028	371155	351552	349097	347523	334051	343353	264651	353605	416941	323701	306631	318900	347029	324880	73862	214135	105857	79596	106137	180627	174125	110128	93634	195933	199820	107596	127336	127304	153752	137489	105432
Dihydrofolate	683383	962222	1366938	2520762	2249420	796018	2172015	2079941	1275623	1771559	1713174	2183656	1241337	672423	1265272	1716039	1969456	2050220	2016746	1704250	1928295	3002905	2068950	984928	693795	1828188	654660	1128063	2025175	1963669	1854582	970433	1180881	1440768	1739643	1870007
Ornithine	2990241	2787284	2895834	2194669	2531277	2486040	2514672	2562544	2473310	3033119	2643493	2824507	2340844	2336708	2322689	2166915	2000352	1645857	2348722	1675975	2047026	1574158	1551047	1569893	1572732	1304313	1157778	2059251	1991110	1916218	1633977	1677024	1772457	1751728	1634916	1640906
L-Citrulline	1767039	1748466	1436237	1147182	1839881	2357700	2184273	2130999	2774189	2587890	3444301	2509326	3175462	3524434	3478369	4163411	4124376	4141121	3855932	3071037	3688271	2741721	3233512	3991063	5239894	3996319	4923732	3510993	3217070	2882749	2425838	3346727	2922783	2707606	3040124	2480167
Putrescine	73032	123540	85455	47345	62064	72670	50371	62643	78725	64359	65002	76024	82958	71188	67992	78162	78667	88250	78102	20499	67097	49616	28683	42348	40436	30431	26254	31370	38905	44683	37752	45487	44557	42404	26967	31081
Spermidine	9669072	10973220	11595560	11591200	9798829	10462710	9432251	10076880	9634518	8995980	8291308	8658556	9648868	9194941	9183828	10982440	8207350	9509229	10787930	6066270	9713023	8368386	7923916	6823004	7322232	6779582	6254818	5520509	6985576	7388376	6370488	6429104	6269272	6663330	5528966	5684254
Spermine	7537668	8315000	9062967	9695125	8727847	7987754	8859790	9580675	9081707	9078657	7557918	11298180	9735713	8895951	10009370	11024820	9130809	8602339	11898880	6319848	10199840	10875730	11047970	7995280	7575438	7096642	6916938	6482126	9569657	9562685	8120150	8005472	7754088	8482888	7611702	6886198
alpha-D-Glucosamine 1-phosphate	1754212	2294460	2059948	1104679	1629167	3066858	1717910	1655192	2323316	1832612	1921680	1978100	2085126	2400531	2178979	2318962	1610935	1743297	1454972	622140	1390976	445390	615148	857838	885785	447527	1206572	556250	483712	891138	854331	1674888	1259018	963190	822304	547638
D-Glucosamine	100599	71465	43228	101862	80725	148024	65635	126447	110609	155612	243234	114112	172754	202140	187680	257111	305071	216461	157444	275806	172217	229347	126798	204138	603082	334369	1047878	387256	240532	315223	284479	282166	201156	333472	252679	270924
Carnosine	292183	368499	318063	173913	221355	349476	222104	172612	252196	243178	248919	175830	264054	293412	291329	265699	153550	251971	223936	122431	204551	106274	94538	202780	208897	171191	189571	130503	138071	155862	167635	225552	166394	145710	152171	115481
Creatine	234414600	318146100	270408200	221715200	262051900	327596100	263908300	263017300	295904900	263065500	302022600	310500600	298583300	328117300	301746400	382087200	313305400	315468600	300368000	203992800	328406900	246046800	226030900	223187200	257064200	212314400	230439100	179072800	180543300	159443500	160293200	166273000	156512900	136700500	141876800	127181700
Creatinine	3432043	4822214	4392838	3206365	3745552	4224918	3898479	3592886	3780629	3547669	3718582	4339918	3458522	3247964	2949527	3989326	3093836	3733526	3069319	2187872	3433499	2765076	2618535	2869582	2384491	2514293	2494148	2233818	2653736	2681164	2213653	2245138	2283341	2180963	2317730	1846364
4-Acetamidobutanoate	482838	605201	515242	500313	569969	792565	704801	681390	853011	805159	863213	716805	793028	854494	872988	1131192	1109252	904544	987170	815404	974823	890785	978190	1036717	1121744	927219	1254033	994681	1057988	1143814	1112732	1240005	1028027	1031214	975830	893186
Pantothenol	368701	281903	343141	350304	466036	286176	272508	350689	334964	289268	337690	207834	268863	246874	208466	274857	189366	158193	200947	178337	188249	234083	240826	190643	193157	132865	188660	126092	191566	182721	224320	254855	264480	282305	218752	217982
Taurine	299653	291009	258042	271240	361676	429164	424303	376886	531447	464208	756434	657622	739618	1034483	1053512	1393084	1120399	1329468	1206362	1526766	1547151	1135181	1164582	1538401	1773471	1312687	2028976	1075145	1189036	1262806	1345898	1220096	977457	964803	907633	1249708
Hypotaurine	0	0	33466	0	0	30993	34868	15679	30015	15560	29297	0	52271	41610	51476	36888	40751	42898	30815	60127	40631	62413	52968	51661	103467	51099	99880	54838	51221	40166	77569	35710	37590	70983	33421	59442
2S-5S-Methionine sulfoximine	25579	46253	41861	30311	36891	33168	36947	36744	42463	38795	35672	32821	34776	57898	31125	48843	36047	28976	33465	34188	63216	34471	36948	47274	57074	27488	38058	57584	18547	62312	68125	137515	68219	58641	68236	51953
L-Selenomethionine	66877	74609	62170	72186	99674	71547	74652	90598	93395	76412	85441	55979	67041	62951	59433	48320	56506	52817	54028	63222	70785	39314	65738	66273	89063	83908	77833	63030	64754	59376	44531	47659	64186	55508	66985	58627
3-Methyleneoxindole	289631	330826	395037	444247	372500	445986	689466	554842	558429	488045	715096	678767	767893	557601	624170	812286	632503	552382	661965	650956	303637	615007	629784	552821	639362	437321	382851	183625	283678	224255	123957	118613	108916	181389	61098	50112
Indole	0	45254	34280	0	0	7879	51743	110000	110694	129912	54871	134092	72125	70760	98240	245698	170484	200535	255474	141892	287735	84224	251938	138234	237965	297789	60205	179564	136399	201487	91036	171349	135763	119936	69563	13546
kynurenine	44778	47750	49273	91755	52583	23225	66973	72882	34793	56844	48807	71503	43185	24759	42470	43153	65784	51345	68870	60470	56763	94770	68226	46634	26487	49009	35126	38857	58373	63085	46718	35945	32710	46369	48343	57205
Anthranilate	2136706	2394115	1993240	1609968	2248408	2823925	2409012	2342804	2495756	2773476	2563938	2833630	2715907	2568390	2830827	3080599	3621297	2837360	2855851	2371713	3720293	2678246	3053952	3113083	3955486	3806074	4460602	3330015	3210149	3300941	3566868	3548797	3472605	3134830	3002063	2686948
L-Adrenaline	450330	684016	689073	768482	753492	533812	608148	768103	706670	595860	553183	663994	791391	625721	740050	763337	598169	620457	460741	352261	650787	531125	581400	535752	396648	439686	218937	303330	354834	367846	389723	346161	294346	362105	283502	363803
N-Methylethanolamine phosphate	49882480	65965330	47142250	28547390	46289080	58875180	46241220	39764690	46690290	46505040	43754470	40622520	39859450	46928930	42756600	41564330	35555920	34358640	24728300	10244570	21553970	12536960	10400600	10270420	19899090	7832400	11939380	7516694	9019291	9632585	8371394	16564570	12875210	10021600	9425862	7505018
L-Carnitine	21211810	31360660	23751390	20900810	21878360	28486170	22589830	20440680	26911520	20644770	22396530	23855580	26751310	28286090	26577670	29186070	22497960	27169700	23243610	15685010	23770330	18625500	18824590	19828420	19317950	14256860	12675050	11624360	12525140	13794300	14206890	14256750	13816500	12795580	11640490	11256240
Triacanthine	22164610	30042780	30998680	27036160	25951270	30706750	26886270	26795550	29032470	25269550	21375150	37561630	34400670	28139860	31260250	37119930	34166360	36829210	48913950	32952940	63466050	42589900	46588470	42131970	39231420	44387530	38153160	37865450	37399170	40524370	31742420	32096140	33392830	27578160	23469700	20526440
2-Methyleneglutarate	154078	186555	146330	81073	138201	272465	303800	277260	215683	298268	424735	225674	413694	345612	471790	555029	651401	370575	442526	463875	377695	400929	379619	409462	1025708	656045	1454474	617559	483273	516957	523402	522880	467619	571287	707295	431368
6-Lactoyl-5-6-7-8-tetrahydropterin	16996110	20905530	26160860	45772780	38416290	16698790	39367820	39099040	25926470	40051310	37203440	54721780	34146880	23333240	33607280	36374720	58450240	55746290	66647440	101763500	83126740	100368300	101032200	53336450	25027300	57319660	18590220	49194560	59179680	48531360	43135110	25993480	32699620	38726790	38461850	51770350
Methylenediurea	2417236	3569764	3002049	2374217	2941613	1786621	2799328	2569735	3234268	2712193	3285142	3337051	1706986	3689554	3217221	1860148	3034831	3408143	3326286	2204778	0	2632574	2339792	2464966	2743227	1639654	2537452	964346	1409285	1713689	718505	0	0	0	273716	1395920
3-Oxalomalate	2244073	2744972	2255156	2192783	2168116	2454102	1932854	1738194	2117245	2039326	1714721	1684613	1605792	1948504	1490777	1067926	1056530	1175766	834036	429826	808154	682818	667434	783731	753229	593127	804119	652252	1063626	1682338	1736982	2345188	2250576	1731914	1814147	1731486
cis-p-Coumarate	2070246	2729943	3061878	2992060	2938005	2030771	3021448	3072764	2393429	5003258	3905261	6058574	4748520	4509256	5415018	5653550	6679384	5152054	7647230	5185008	7913792	6106428	7067380	6952470	5421210	6948648	5240592	7867638	6509826	7041522	4099038	5263296	6045816	5010038	5209538	3974754
Poly-gamma-D-glutamate	13284100	13261030	13833430	17194170	20423520	15869960	22832790	23839590	18601020	26900940	28301590	35631510	24894430	27716280	34294160	39495050	46113520	48594600	58838240	59054840	60230940	57794520	61893020	55385120	45514340	60061630	43103990	54711650	58140170	50556000	42833100	38871560	39133470	44525350	42536610	44643690
N-Acyl-D-aspartate	191192	239223	216349	250239	271767	272112	281773	323992	289207	279018	235901	256854	246531	300573	249991	231176	255874	258130	225203	160350	205986	157384	116413	131125	138665	104645	77190	54150	50213	55369	61930	72225	71820	61607	54953	52563
MS_METABOLITE_DATA_END
#METABOLITES
METABOLITES_START
metabolite_name	CmpdID	parent	medRt
L-alanine	C00041	90.0557	1.16
L-arginine	C00062	175.1191	1.11
L-asparagine	C00152	133.0609	1.17
L-aspartate	C00049	134.0449	1.35
L-cysteine	C00097	122.0273	1.20
L-glutamate	C00025	148.0604	1.28
L-glutamine	C00064	147.0765	1.17
glycine	C00037	76.0402	1.17
L-histidine	C00135	156.0769	1.14
L-leucine	C00123	132.1021	1.22
L-isoleucine	C00407	132.1020	1.63
L-lysine	C00047	147.1129	1.02
L-methionine	C00073	150.0584	1.22
L-phenylalanine	C00079	166.0863	1.24
L-proline	C00148	116.0710	1.20
L-serine	C00065	106.0504	1.17
L-threonine	C00188	120.0658	1.18
L-tryptophan	C00078	205.0973	1.27
L-tyrosine	C00082	182.0811	1.22
L-valine	C00183	118.0866	1.19
AMP	C00020	348.0704	1.82
Cytosine	C00380	112.0510	1.34
Xanthine	C00385	153.0399	1.00
Allantoate	C00499	177.0610	1.08
5-Hydroxyisourate	C11821	185.0322	1.17
Nicotinamide	C00153	123.0556	1.22
NAD+	C00003	664.1161	1.75
Glutathione	C00051	308.0909	1.70
Cys-Gly	C01419	179.0485	1.70
gamma-Glutamyl-gamma-aminobutyrate	C15767	233.1132	2.25
5-L-Glutamyl-taurine	C05844	255.0631	1.13
Dimethylglycine	C01026	104.0711	1.19
Phosphoserine	C02532	186.0165	1.27
5-10-Methenyltetrahydrofolate	C00445	457.1687	1.18
Dihydrofolate	C00415	444.1613	1.18
Ornithine	C01602	133.0974	1.02
L-Citrulline	C00327	176.1031	1.17
Putrescine	C00134	89.1081	1.02
Spermidine	C00315	146.1653	1.03
Spermine	C00750	203.2232	1.03
alpha-D-Glucosamine 1-phosphate	C06156	260.0533	1.20
D-Glucosamine	C00329	180.0867	1.19
Carnosine	C00386	227.1152	1.12
Creatine	C00300	132.0769	1.18
Creatinine	C00791	114.0666	1.18
4-Acetamidobutanoate	C02946	146.0812	1.19
Pantothenol	C00864	220.1181	0.26
Taurine	C00245	126.0223	1.18
Hypotaurine	C00519	110.0275	1.18
2S-5S-Methionine sulfoximine	C03829	181.0626	1.11
L-Selenomethionine	C05335	198.0015	0.98
3-Methyleneoxindole	C02796	146.0602	1.27
Indole	C00463	118.0655	1.24
kynurenine	C00328	209.0903	1.11
Anthranilate	C00108	138.0551	1.19
L-Adrenaline	C00788	184.0946	1.15
N-Methylethanolamine phosphate	C01210	156.0422	1.18
L-Carnitine	C00318	162.1126	1.16
Triacanthine	C08435	204.1231	1.22
2-Methyleneglutarate	C02930	145.0497	1.19
6-Lactoyl-5-6-7-8-tetrahydropterin	C04244	240.1093	1.16
Methylenediurea	C06381	133.0739	1.19
3-Oxalomalate	C01990	207.0142	1.15
cis-p-Coumarate	C06738	165.0547	1.22
Poly-gamma-D-glutamate	C05723	130.0501	1.17
N-Acyl-D-aspartate	C06380	161.0297	0.99
METABOLITES_END
#END