#METABOLOMICS WORKBENCH juliehaines_20240719_102536 DATATRACK_ID:5026 STUDY_ID:ST003332 ANALYSIS_ID:AN005459 PROJECT_ID:PR002070
VERSION             	1
CREATED_ON             	July 19, 2024, 3:03 pm
#PROJECT
PR:PROJECT_TITLE                 	Increased Cholesterol Synthesis Drives Neurotoxicity in Patient Stem
PR:PROJECT_TITLE                 	Cell-Derived Model of Multiple Sclerosis
PR:PROJECT_SUMMARY               	Senescent neural progenitor cells have been identified in brain lesions of
PR:PROJECT_SUMMARY               	people with progressive multiple sclerosis (PMS). However, their role in disease
PR:PROJECT_SUMMARY               	pathobiology and contribution to the lesion environment remains unclear. By
PR:PROJECT_SUMMARY               	establishing directly induced neural stem/progenitor cell (iNSC) lines from PMS
PR:PROJECT_SUMMARY               	patient fibroblasts, we studied their senescent phenotype in vitro. Senescence
PR:PROJECT_SUMMARY               	was strongly associated with inflammatory signaling, hypermetabolism, and the
PR:PROJECT_SUMMARY               	senescence associated secretory phenotype (SASP). PMS-derived iNSCs displayed
PR:PROJECT_SUMMARY               	increased glucose-dependent fatty acid and cholesterol synthesis, which resulted
PR:PROJECT_SUMMARY               	in the accumulation of cholesteryl ester-enriched lipid droplets. An HMG-CoA
PR:PROJECT_SUMMARY               	reductase-mediated lipogenic state was found to induce secretion of the SASP in
PR:PROJECT_SUMMARY               	PMS iNSC conditioned media via transcriptional regulation by
PR:PROJECT_SUMMARY               	cholesterol-dependent transcription factors. SASP from PMS iNSCs induced
PR:PROJECT_SUMMARY               	neurotoxicity. Chemical targeting of HMG-CoA reductase using the
PR:PROJECT_SUMMARY               	cholesterol-lowering drug simvastatin (SV) reprogrammed the SASP and rescued
PR:PROJECT_SUMMARY               	neurotoxicity. Our findings suggest a disease-associated, cholesterol-related,
PR:PROJECT_SUMMARY               	hypermetabolic phenotype of PMS iNSCs that leads to neurotoxic signaling and is
PR:PROJECT_SUMMARY               	rescuable pharmacologically.
PR:INSTITUTE                     	University of Colorado Denver
PR:LABORATORY                    	Lab of Angelo D'Alessandro in collaboration with lab of Stefano Pluchino (Univ
PR:LABORATORY                    	of Cambridge)
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                   	Increased Cholesterol Synthesis Drives Neurotoxicity in Patient Stem
ST:STUDY_TITLE                   	Cell-Derived Model of Multiple Sclerosis - media metabolomics
ST:STUDY_SUMMARY                 	Fibroblasts from skin biopsies from patients with progressive multiple sclerosis
ST:STUDY_SUMMARY                 	(PMS) and age-matched controls (AMC) were derived and reprogrammed to yield
ST:STUDY_SUMMARY                 	induced neural stem cells (iNSCs). The resulting iNSCs were cultured in the
ST:STUDY_SUMMARY                 	presence of 13C6 glucose for 24h to assess changes in the intracellular
ST:STUDY_SUMMARY                 	metabolome, in particular changes in the flux of glucose-derived carbon into the
ST:STUDY_SUMMARY                 	TCA cycle along with steady state levels of compounds in other areas of the
ST:STUDY_SUMMARY                 	metabolome. The data in this study are from the assessment of the conditioned
ST:STUDY_SUMMARY                 	culture media at the end of the 24h incubation period.
ST:INSTITUTE                     	University of Colorado Denver
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
SU:GENDER                        	Female
#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           	A	AMC A S7	Disease state:AMC | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-25
SUBJECT_SAMPLE_FACTORS           	A	AMC A S8	Disease state:AMC | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-26
SUBJECT_SAMPLE_FACTORS           	A	AMC A S9	Disease state:AMC | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-27
SUBJECT_SAMPLE_FACTORS           	B	AMC B S7	Disease state:AMC | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-28
SUBJECT_SAMPLE_FACTORS           	B	AMC B S8	Disease state:AMC | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-29
SUBJECT_SAMPLE_FACTORS           	B	AMC B S9	Disease state:AMC | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-30
SUBJECT_SAMPLE_FACTORS           	C	AMC D S7	Disease state:AMC | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-34
SUBJECT_SAMPLE_FACTORS           	C	AMC D S8	Disease state:AMC | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-35
SUBJECT_SAMPLE_FACTORS           	C	AMC D S9	Disease state:AMC | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-36
SUBJECT_SAMPLE_FACTORS           	D	PMS A S7	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-37
SUBJECT_SAMPLE_FACTORS           	D	PMS A S8	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-38
SUBJECT_SAMPLE_FACTORS           	D	PMS A S9	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-39
SUBJECT_SAMPLE_FACTORS           	E	PMS B S7	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-40
SUBJECT_SAMPLE_FACTORS           	E	PMS B S8	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-41
SUBJECT_SAMPLE_FACTORS           	E	PMS B S9	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-42
SUBJECT_SAMPLE_FACTORS           	F	PMS C S7	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-43
SUBJECT_SAMPLE_FACTORS           	F	PMS C S8	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-44
SUBJECT_SAMPLE_FACTORS           	F	PMS C S9	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-45
SUBJECT_SAMPLE_FACTORS           	G	PMS D S7	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-46
SUBJECT_SAMPLE_FACTORS           	G	PMS D S8	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-47
SUBJECT_SAMPLE_FACTORS           	G	PMS D S9	Disease state:PMS | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-48
SUBJECT_SAMPLE_FACTORS           	NA	NC 7	Disease state:no cell ctrl | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-49
SUBJECT_SAMPLE_FACTORS           	NA	NC 8	Disease state:no cell ctrl | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-50
SUBJECT_SAMPLE_FACTORS           	NA	NC 9	Disease state:no cell ctrl | Sample source:cell culture supernatant	RAW_FILE_NAME(raw file name)=DN47-51
#COLLECTION
CO:COLLECTION_SUMMARY            	3-5 mm skin biopsies were collected in Biopsy Collection Medium (RPMI 1460
CO:COLLECTION_SUMMARY            	[Thermo Fisher] with 1X Antibiotic-Antimycotic [Thermo Fisher]), cut into
CO:COLLECTION_SUMMARY            	smaller pieces (<1 mm) and plated onto a TC-treated 35 mm dish in Biopsy Plating
CO:COLLECTION_SUMMARY            	Medium, composed by Knockout DMEM (Thermo Fisher), 2 mM GlutaMax (Thermo
CO:COLLECTION_SUMMARY            	Fisher), 0.1 mM non-essential amino acids (Thermo Fisher), 0.1 mM
CO:COLLECTION_SUMMARY            	β-Mercaptoethanol (Thermo Fisher), 10% Fetal Bovine Serum (FBS) (Thermo
CO:COLLECTION_SUMMARY            	Fisher), 1X Penicillin-Streptomycin (P/S) (Thermo Fisher) and 1% Nucleosides
CO:COLLECTION_SUMMARY            	(Millipore). Once the first fibroblasts migrated out of the biopsies, the
CO:COLLECTION_SUMMARY            	cultures were maintained in growth medium (DMEM Glutamax I [Thermo Fisher]
CO:COLLECTION_SUMMARY            	supplemented with 10% fetal bovine serum, 1% non-essential amino acids and 1 mM
CO:COLLECTION_SUMMARY            	sodium pyruvate (Thermo Fisher) at 37°C with 5 % CO2 and fed every 3-4 days.
CO:COLLECTION_SUMMARY            	After reaching 90% confluency the fibroblasts were detached with trypsin-EDTA
CO:COLLECTION_SUMMARY            	0.05% for 5 min followed by neutralization in DMEM and spun down at 300xg for 5
CO:COLLECTION_SUMMARY            	min. They were split 1:4 into growth media. Fibroblasts at passage 3-5 were
CO:COLLECTION_SUMMARY            	reprogrammed to iPSCs using the integration free technology based on
CO:COLLECTION_SUMMARY            	non-modified RNA plus microRNA (kit from REPROCELL, formerly Stemgent),
CO:COLLECTION_SUMMARY            	following manufacturer’s instructions. About 25x10^3 fibroblasts/well were
CO:COLLECTION_SUMMARY            	plated onto Matrigel-coated 12-well plates in culture medium for 24 hours and
CO:COLLECTION_SUMMARY            	then in NuFF-conditioned Pluriton reprogramming medium with B18R. Cells were
CO:COLLECTION_SUMMARY            	transfected for 11 consecutive days using Stemfect as following: day 0 microRNA
CO:COLLECTION_SUMMARY            	only, days 1-3 RNA only, day 4 microRNA plus RNA, days 5-11 RNA only. From day
CO:COLLECTION_SUMMARY            	11, TRA-1-60+ colonies (live stained) were manually picked and re-plated on
CO:COLLECTION_SUMMARY            	mouse embryonic fibroblasts in HUESM medium (Knockout-DMEM, 20% knock-out serum,
CO:COLLECTION_SUMMARY            	glutamax 2mM, NEAA 0.1mM, 1X P/S and β-mercaptoethanol 0.1mM [Thermo Fisher]).
CO:COLLECTION_SUMMARY            	Pluripotent colonies were passaged and adapted to feeder-free conditions with
CO:COLLECTION_SUMMARY            	hESC matrigel (Corning) and mTeSR1 (STEMCELL Technologies) medium. The WIBJ iPSC
CO:COLLECTION_SUMMARY            	line, obtained from Cambridge BioResource as a gift from Alessandra Granada, was
CO:COLLECTION_SUMMARY            	reprogrammed from fibroblasts using the CytoTune 1 non-integrating kit (Thermo
CO:COLLECTION_SUMMARY            	Fisher). iPSC media was changed every day. When confluent, cells were lifted
CO:COLLECTION_SUMMARY            	using accutase (Thermo Fisher), spun at 300xg for 5 min, and split 1:10-1:20
CO:COLLECTION_SUMMARY            	onto hESC-matrigel coated plates with Y-27632 (10 uM) (Thermo Fisher) in mTeSR1
CO:COLLECTION_SUMMARY            	media. To generate directly induced neural stem cells (iNSCs) from fibroblasts
CO:COLLECTION_SUMMARY            	we used a nonintegrating Sendai virus-based direct conversion strategy, as
CO:COLLECTION_SUMMARY            	previously described (https://doi.org/10.3791/52831). Briefly, fibroblasts were
CO:COLLECTION_SUMMARY            	seeded at 75,000/well in fibroblast media in a non-coated 12-well. On the next
CO:COLLECTION_SUMMARY            	day the fibroblasts were transduced using the CytoTune-iPS 2.0 Sendai
CO:COLLECTION_SUMMARY            	Reprogramming Kit (Thermo Fisher) with hKOS (MOI: 3), hc-Myc (MOI: 3), hKlf4
CO:COLLECTION_SUMMARY            	(MOI: 3). The day following transfection, the medium was switched to neural
CO:COLLECTION_SUMMARY            	induction medium (NIM) [DMEM:F12 and Neurobasal (1:1), supplemented with N2
CO:COLLECTION_SUMMARY            	supplement (1x, ThermoFisher), 1% glutamax, B27 supplement (1x, ThermoFisher),
CO:COLLECTION_SUMMARY            	CHIR99021 (3 µM, Cell Guidance Systems), SB-431542 (2 µM, Cayman Chemicals)
CO:COLLECTION_SUMMARY            	and hLIF (10 ng/ml, Cell Signaling Technology)], and cells were moved to 39°C
CO:COLLECTION_SUMMARY            	with 5% CO2 to achieve viral clearance over 14 days. A few samples were
CO:COLLECTION_SUMMARY            	collected here to generate positive controls for quality control assays. Medium
CO:COLLECTION_SUMMARY            	changes were performed every other day. Following 25 days of transfection, iNSC
CO:COLLECTION_SUMMARY            	colonies were manually selected, seeded onto Growth Factor Reduced (GFR)
CO:COLLECTION_SUMMARY            	Matrigel Matrix (1:20 in DMEM/F12) coated plates for expansion, and subjected to
CO:COLLECTION_SUMMARY            	quality control assays. iNSCs were maintained in NIM media until 70% confluent,
CO:COLLECTION_SUMMARY            	then lifted using accutase (ThermoFisher), spun at 300xg for 3 mins, and plated
CO:COLLECTION_SUMMARY            	onto GFR-Matrigel coated plates with Y-27632 (10 µM, Miltenyi Biotec) between
CO:COLLECTION_SUMMARY            	1:3-1:5 in NIM media. Media was changed every second day as needed. Experiments
CO:COLLECTION_SUMMARY            	were performed on cells from passages 20-40.
CO:SAMPLE_TYPE                   	Culture Media
#TREATMENT
TR:TREATMENT_SUMMARY             	iNSCs were seeded in NIM with Y-27632 (10 µM) at a density of 100,000 cells/cm2
TR:TREATMENT_SUMMARY             	GFR-matrigel coated wells, in technical triplicates per line. Medium was changed
TR:TREATMENT_SUMMARY             	the next day. Glucose tracer experiments were run on day 2 after seeding by
TR:TREATMENT_SUMMARY             	removing the culture medium and adding glucose-free NIM [DMEM:F12 w/o glucose
TR:TREATMENT_SUMMARY             	and Neurobasal (1:1), supplemented with N2 supplement (1x, ThermoFisher),
TR:TREATMENT_SUMMARY             	Glutamax (Thermo Fisher, 3.2 mM), pyruvate (Agilent, 0.35 mM), B27 supplement
TR:TREATMENT_SUMMARY             	(1x, ThermoFisher), CHIR99021 (3 µM, Cell Guidance Systems), SB-431542 (2 µM,
TR:TREATMENT_SUMMARY             	Cayman Chemicals) and hLIF (10 ng/ml, Cell Signaling Technology)] supplemented
TR:TREATMENT_SUMMARY             	with D-Glucose-13C6 (ThermoFisher, 20.4 mM). Samples were collected after 24
TR:TREATMENT_SUMMARY             	hours for optimal resolution into TCA cycle and anabolic metabolic pathways.
TR:TREATMENT_SUMMARY             	Sample collection was performed on ice. Conditioned culture medium was
TR:TREATMENT_SUMMARY             	collected, spun at 300 x g for 5 minutes to remove cellular debris and 100 µL
TR:TREATMENT_SUMMARY             	was stored at -80 oC until sample processing.
#SAMPLEPREP
SP:SAMPLEPREP_SUMMARY            	Metabolites from thawed media aliquots (20 uL) were extracted with 480 uL of ice
SP:SAMPLEPREP_SUMMARY            	cold 5:3:2 methanol:acetonitrile:water (v/v/v) with vigorous vortexing at 4
SP:SAMPLEPREP_SUMMARY            	degrees C followed by centrifugation for 10 min at 18,000 g, 4°C and stored at
SP:SAMPLEPREP_SUMMARY            	−80°C until analysis.
SP:PROCESSING_STORAGE_CONDITIONS 	4℃
SP:EXTRACT_STORAGE               	-80℃
#CHROMATOGRAPHY
CH:CHROMATOGRAPHY_SUMMARY        	Positive C18
CH:CHROMATOGRAPHY_TYPE           	Reversed phase
CH:INSTRUMENT_NAME               	Thermo Vanquish
CH:COLUMN_NAME                   	Phenomenex Kinetex C18 (150 x 2.1mm,1.7um)
CH:SOLVENT_A                     	100% water; 0.1% formic acid
CH:SOLVENT_B                     	100% acetonitrile; 0.1% formic acid
CH:FLOW_GRADIENT                 	0-0.5 min 5% B, 0.5-1.1 min 5-95% B, 1.1-2.75 min hold at 95% B, 2.75-3 min
CH:FLOW_GRADIENT                 	95-5% B, 3-5 min hold at 5% B
CH:FLOW_RATE                     	450 uL/min
CH:COLUMN_TEMPERATURE            	45
CH:SAMPLE_INJECTION              	20 uL
#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                   	Resolution 70,000, scan range 65-900 m/z, maximum injection time 200 ms,
MS:MS_COMMENTS                   	microscans 2, automatic gain control (AGC) 3 x 10^6 ions, source voltage 4.0 kV,
MS:MS_COMMENTS                   	capillary temperature 320 C, and sheath gas 45, auxiliary gas 15, and sweep gas
MS:MS_COMMENTS                   	0 (all nitrogen). Data converted to mzXML using RawConverter. Metabolites were
MS:MS_COMMENTS                   	annotated and integrated using Maven in conjunction with the KEGG database.
#MS_METABOLITE_DATA
MS_METABOLITE_DATA:UNITS	peak area
MS_METABOLITE_DATA_START
Samples	AMC A S7	AMC A S8	AMC A S9	AMC B S7	AMC B S8	AMC B S9	AMC D S7	AMC D S8	AMC D S9	PMS A S7	PMS A S8	PMS A S9	PMS B S7	PMS B S8	PMS B S9	PMS C S7	PMS C S8	PMS C S9	PMS D S7	PMS D S8	PMS D S9	NC 7	NC 8	NC 9
Factors	Disease state:AMC | Sample source:cell culture supernatant	Disease state:AMC | Sample source:cell culture supernatant	Disease state:AMC | Sample source:cell culture supernatant	Disease state:AMC | Sample source:cell culture supernatant	Disease state:AMC | Sample source:cell culture supernatant	Disease state:AMC | Sample source:cell culture supernatant	Disease state:AMC | Sample source:cell culture supernatant	Disease state:AMC | Sample source:cell culture supernatant	Disease state:AMC | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:PMS | Sample source:cell culture supernatant	Disease state:no cell ctrl | Sample source:cell culture supernatant	Disease state:no cell ctrl | Sample source:cell culture supernatant	Disease state:no cell ctrl | Sample source:cell culture supernatant
L-alanine	3284612	2216189	2768536	2681530	2229748	3213628	3866803	3438853	3910888	1412645	1915648	2147244	2176144	2046095	2193545	1642490	1472006	1318878	2478558	1583079	1837423	141054	69557	90347
L-arginine	52974636	35023120	41679636	24445790	48068900	33245888	33734604	35302764	47450276	18072616	45439920	45267424	19112290	27542022	40549504	8035442	8659089	13774848	14074679	26255074	15589736	56134192	27794502	26291336
L-asparagine	376470	228135	210098	372097	404294	390539	141965	134770	101657	461430	526322	514977	294981	248907	270579	250551	244479	216074	234182	329934	167142	431188	142428	142431
L-glutamate	197305	208892	158701	207014	169708	225188	142888	175647	129392	213229	148469	145416	262035	167352	126822	214901	209163	220266	191466	252203	163735	148500	243518	158637
L-glutamine	36686076	22700408	31165416	21298830	27076840	23588506	26937974	23002874	24921066	16421100	33033154	31447154	15763577	20395352	20078274	10595136	10020347	15105333	16437600	19952570	13364051	19739046	9892077	6935428
L-histidine	3582886	2477137	3278412	2469673	3408388	3769994	3057738	2841734	4119848	1952420	4127839	4361335	2235334	2607206	3624770	2079013	2156525	2276374	2457002	2996515	2435386	3671083	2192653	2248182
L-leucine	69092256	122082936	51038160	52132812	52110496	65558368	93105336	65153184	75378440	127562816	47097716	47092308	56155056	63079712	56263572	68878256	76094344	89917824	53240172	51645020	46295252	67422864	187278928	69767576
L-lysine	6756737	9230212	8377266	10661327	5458807	11398411	12011945	12024635	11231464	8155295	5394720	6398394	10209153	8450333	7440862	7401596	7917284	5739116	7793686	5820547	7573097	6971644	10390411	12384781
L-methionine	9659761	5376911	4756581	9159017	9009951	7304860	7665083	8904736	6832334	7267710	4593681	3826751	8291178	8181054	6070850	3419674	4020451	3053529	3150267	5940760	5143763	8384578	6815856	11620281
L-proline	27670726	23690432	24883400	18328978	22129806	19537418	14926267	18407558	23025368	21938962	29081920	25007600	26558466	32823610	39222580	52000852	44499616	60742540	45451008	65787180	47375868	24342342	14345309	15290043
L-serine	228449	172682	219104	168636	194701	205931	258814	327400	268882	86093	70653	67600	107158	76613	77392	45515	49669	12186	35882	77152	52325	356040	245883	239944
L-threonine	3613479	2667850	3389222	2248712	3343141	2869691	2799060	2358139	2537234	2475637	3763996	4204774	2345140	3001524	3519408	2053951	1882778	2356908	2897305	2759756	2369608	3285600	1974915	1717495
L-tryptophan	36779460	33267294	35042292	29700290	37298948	23800522	29626720	27690442	25138688	24811408	23028880	22777322	26877226	35056468	31018902	21210844	18014916	21437642	16631276	24108226	21729856	38051904	28960778	32059978
L-tyrosine	33920628	40871356	21927716	25598840	26137050	24663030	16659664	19669568	10855159	18787106	17663358	15050563	18400000	31855422	9797698	10684481	15550435	32496010	13206936	13184515	21523494	26933320	18063870	26116450
L-valine	22968112	20920248	19293360	26546982	18622830	25460570	22407558	23395850	24968826	17023414	18990198	18958006	19296312	21415166	23055200	17234384	20464170	20267938	16198935	22855846	18169242	26099890	21574536	22248422
dAMP	293300	194156	207694	205183	192862	196169	205237	203441	185676	164460	277300	247085	133082	132548	152740	123480	131886	142080	129195	167038	180643	11076	0	0
Cytidine	196206	178790	214451	168321	214453	166692	224222	224759	260440	153228	240665	232006	186686	178704	235072	180344	195530	240238	267206	253767	238875	252007	215148	214118
UMP	1030476	1086871	1066257	1247546	664296	1366680	1356425	1307188	1262065	913117	874898	1026443	1324769	1043040	731970	1464751	1349007	1461753	1377863	1476261	729417	344913	687245	747325
Hypoxanthine	2060307	911266	1313730	1324385	1318771	1440559	2453269	2605014	1151020	574501	426863	423232	300784	269114	304498	147854	115632	195096	195731	241114	201271	2267232	1939226	3010731
Allantoate	435973	462503	437205	791824	547989	619181	266949	262873	409244	925284	649261	776601	1391187	1328726	1045533	2107604	1969008	1945822	1820265	1483588	1780773	141180	177970	205499
(S)(+)-Allantoin	87827	72848	65606	74407	54985	60886	64094	72932	82343	55024	39948	39367	60124	59115	41193	9266	19921	26583	39571	34792	38509	58556	70395	111530
5-Hydroxyisourate	1799395	1164107	1303719	1184280	1449112	1579930	1216020	1318992	1563854	802551	1343894	1325214	862967	883783	1179648	733703	706221	762856	928123	803340	968372	188948	40274	61931
Urate	868000	242643	451334	468829	285313	323159	407659	400687	147207	194437	330537	291376	222531	307661	369576	342242	377450	301177	353652	257702	438668	314695	507883	183658
4-Pyridoxate	5438	2010	3624	10306	29397	15995	86639	43948	18588	225170	482821	530091	418165	564696	1054110	123100	85608	170896	5945	0	4570	5359648	2383594	1902560
Pyridoxamine 5'-phosphate	288259	346634	449229	457195	267755	710465	542317	682841	1000686	360649	393603	653867	426281	269802	297708	386946	289752	206766	585945	269897	450983	319602	381378	460137
Nicotinamide	18469194	11316957	5110811	18202858	17104424	14769300	14781917	15397383	10763603	19557020	11635499	11489256	19963570	17796452	16360827	15505216	17777800	7237914	12298465	19091258	17044430	13861187	12300017	16661657
Nicotinate ribonucleotide	69613	57004	117913	38956	88761	52304	63712	89118	94250	46302	117219	155897	41722	54182	89758	54189	62603	80611	64004	56252	48947	76127	62985	71843
5-Phosphoribosylamine	24707	25112	30028	27469	20304	11859	21439	27621	41651	21092	31151	29717	24812	15802	27926	39826	25363	24248	27166	29132	28621	18014	25339	25947
Phosphate	1309066	856534	937382	578530	692266	347605	759788	815442	2205806	799436	905335	1103307	733013	766310	787213	784204	936611	894775	782355	811673	811687	771112	817875	752859
D-Glucose 6-phosphate	40606	17073	7759	46659	29446	15666	22561	27573	13551	22923	31792	26934	40683	43084	25768	35206	24542	8634	19821	36876	34719	0	3769	0
Ascorbate	1308487	380814	663601	691391	457720	463748	607852	598199	199739	367060	535860	455602	277736	443951	620470	534059	557745	442924	588670	358390	641752	460824	823307	270268
gamma-Glutamyl-Se-methylselenocysteine	225045	204125	196775	343628	227688	251099	112256	111851	148827	369268	353166	371853	604493	609189	507112	844483	909160	987819	797088	693965	776688	74001	70032	98587
5-L-Glutamyl-taurine	52339	23394	56578	21073	64512	48350	70270	58841	107884	22802	59842	84800	15897	35179	49626	41127	9739	38846	40853	65080	18458	27105	14465	42812
S-Adenosyl-L-homocysteine	20341548	14947477	23687594	15855453	21476706	15306456	15030443	17399738	19528834	10084112	26533386	24490944	17668702	17580318	22685242	15347229	17525446	15725789	19438474	14631133	16763632	19336922	17400012	15008355
Ornithine	170036	358565	177801	312156	180027	203942	269989	164096	211470	344300	255781	253290	574735	501288	298187	1036495	801739	756429	853697	604520	733551	158398	476833	168508
Putrescine	167847	219009	147129	256727	168306	155633	169572	176166	325378	220529	174145	162392	187450	147392	217572	232750	238612	246810	170956	194918	221895	195739	143681	192575
Spermine	61273	87874	53350	40361	32887	61998	135931	112701	164991	39476	62513	110806	27553	42385	29433	155883	26629	46844	277736	43581	173873	8521	15355	15084
D-Glucosamine	15878	48288	29140	12540	33599	11538	4180	5176	2623	39954	74069	77317	18679	46829	43901	73140	58609	115346	48257	140848	52229	0	0	0
Pantothenate	20159432	24606250	16406915	10151368	18231110	9546397	10611811	9444185	9637505	10629591	14794332	14160941	10995488	19789018	18401246	10513827	10597324	20511834	10069116	23473040	9390864	16230799	11552039	8526536
L-Methionine S-oxide	390649	293617	285328	186422	325575	158859	126689	186441	151223	133704	355739	376239	218287	261007	445746	183160	162987	344910	199262	461152	202576	444357	258056	189925
3-Methyldioxyindole	22018824	23127914	22582728	15414860	21928880	14863535	18123476	17819750	12616595	14259196	14040996	14414069	16542584	21039926	19766982	12565644	11242429	14537561	10628588	16984924	13202595	19678074	19493978	20268280
Indole	18586910	19576602	18883380	10821281	16053892	10687985	13254221	12746391	8952063	9487115	12205861	11941401	11131625	16944182	16671995	8876478	7646095	13072572	7668597	15401896	9483269	16648329	13893595	14764013
Indole-3-acetaldehyde	1076116	1121234	1793405	1669401	927672	1895009	890274	1495233	623029	1062554	742672	690874	1473978	1015521	1287601	929227	563285	1165003	1314245	897863	1769682	983552	992732	968924
Indole-3-acetate	183850	183816	323680	112489	503024	221800	22628	22690	22542	147823	865505	595603	440957	488904	645979	86748	21181	43332	166363	181252	173599	93729	90735	113823
Indoxyl	2517592	2670345	1878735	2977391	1620219	2598507	2417115	2372774	2462249	3175207	2010629	2732451	3225530	2262732	2150559	3623492	3001186	1615625	3036705	2245498	3188406	2441244	1942304	2383177
g-Oxalo-crotonate	480369	408399	399580	316326	277758	359573	710768	773053	625400	80082	63503	74124	22405	24266	26047	6359	0	0	2546	0	0	536982	559640	595454
Sphingosine	14136	12465	9738	10417	10528	14909	10768	7281	10098	19195	8596	9068	7075	7219	8442	5784	19111	8194	8316	7593	7264	8245	5833	5186
Acetylcholine	390694	399453	482972	259413	531880	336760	261056	282417	331577	262870	375853	334225	315459	426710	424429	443023	438127	424857	380079	467813	363478	448768	360213	259815
Choline	59645312	41257324	61757492	64271796	41889876	82084368	97762912	105288856	143506608	28749162	39585804	51786764	56213156	43441476	41781824	2542880	3546702	3386067	20507146	18517286	24595610	64118580	71104248	74340216
Sphinganine 1-phosphate	676073	5883	10050	96808	5337	237401	76669	60616	6090	12977	9669	11162	3363	75833	10291	31721	4741	7781	2650	3351	77945	55566	164974	40608
L-Carnitine	30679026	30648016	40574004	37494300	32917334	41785252	39335840	35503144	57635292	46199180	62923308	71391288	45130976	28381750	42571532	55147888	60457252	38363284	62521692	37997132	52892900	24910224	27000142	27483642
acetyl-carnitine (acyl-C2)	542680	281077	270953	151441	117155	247086	43787	75874	138334	810286	611390	647855	1118779	887562	733593	2167520	2053744	1499407	2156759	1226936	1982727	0	4889	907
butanoyl-l-carnitine (acyl-C4)	317935	317080	369004	180910	215824	164594	130450	102906	96458	825251	664527	784712	397074	426668	354313	400187	466879	601304	332730	440920	271397	0	2843	4556
L-octanoylcarnitine (acyl-C8)	13262	11949	5926	9914	16441	16208	4637	14252	3699	8806	11996	6734	7155	10340	13706	6411	20986	13212	12833	10308	9443	19542	5270	8730
O-Decenoyl-L-carnitine (acyl-C10:1)	8532	10468	8209	10708	11182	11430	9344	7463	11824	9657	9584	8676	9034	9466	10062	10095	8602	10181	9352	8874	8275	8677	12893	7870
O-dodecenoyl-carnitine (acyl-C12:1)	239866	241641	106214	225946	87737	226927	226537	145030	311700	151183	196943	138227	115885	92045	237114	256508	269362	135432	99335	209078	172423	323424	116412	103567
Tetradecenoyl Carnitine (acyl-C14:1)	5078	4728	4408	3798	4244	16672	4144	4889	3415	5176	3999	5320	4657	4725	4276	4316	18566	5669	4505	3191	8029	13699	3835	3421
MS_METABOLITE_DATA_END
#METABOLITES
METABOLITES_START
metabolite_name	KEGG ID	m/z	r.t.
L-alanine	C00041	90.0547	0.71
L-arginine	C00062	175.1183	0.69
L-asparagine	C00152	133.0592	0.70
L-glutamate	C00025	148.0601	1.70
L-glutamine	C00064	147.0760	0.70
L-histidine	C00135	156.0763	0.69
L-leucine	C00123	132.1015	0.91
L-lysine	C00047	147.1124	0.63
L-methionine	C00073	150.0579	0.87
L-proline	C00148	116.0702	0.71
L-serine	C00065	106.0493	0.75
L-threonine	C00188	120.0652	0.70
L-tryptophan	C00078	205.0965	1.70
L-tyrosine	C00082	182.0806	0.97
L-valine	C00183	118.0858	0.72
dAMP	C00360	332.0792	0.71
Cytidine	C00475	244.0903	1.82
UMP	C00105	325.0371	0.72
Hypoxanthine	C00262	137.0454	0.89
Allantoate	C00499	177.0606	0.66
(S)(+)-Allantoin	C02350	159.0500	0.68
5-Hydroxyisourate	C11821	185.0315	0.70
Urate	C00366	169.0342	1.85
4-Pyridoxate	C00847	184.0592	0.76
Pyridoxamine 5'-phosphate	C00647	249.0602	1.16
Nicotinamide	C00153	123.0549	0.95
Nicotinate ribonucleotide	C01185	337.0515	1.74
5-Phosphoribosylamine	C03090	230.0460	1.81
Phosphate	C00009	98.9839	1.89
D-Glucose 6-phosphate	C02965	261.0374	0.99
Ascorbate	C00072	177.0394	1.98
gamma-Glutamyl-Se-methylselenocysteine	C05695	313.0346	0.66
5-L-Glutamyl-taurine	C05844	255.0644	0.74
S-Adenosyl-L-homocysteine	C00021	385.1280	1.74
Ornithine	C01602	133.0972	0.71
Putrescine	C00134	89.1070	0.64
Spermine	C00750	203.2223	0.71
D-Glucosamine	C00329	180.0869	1.69
Pantothenate	C00864	220.1172	1.69
L-Methionine S-oxide	C02989	166.0528	0.72
3-Methyldioxyindole	C05834	146.0596	1.70
Indole	C00463	118.0647	1.71
Indole-3-acetaldehyde	C00637	160.0752	1.56
Indole-3-acetate	C00954	176.0722	1.72
Indoxyl	C05658	134.0596	0.72
g-Oxalo-crotonate	C03453	159.0274	0.77
Sphingosine	C00319	300.2889	2.64
Acetylcholine	C01996	146.1172	1.25
Choline	C00114	104.1066	0.70
Sphinganine 1-phosphate	C01120	382.2785	1.93
L-Carnitine	C00318	162.1119	0.70
acetyl-carnitine (acyl-C2)	C02571	204.1223	0.72
butanoyl-l-carnitine (acyl-C4)	C02862	232.1534	1.70
L-octanoylcarnitine (acyl-C8)	C02838	288.2157	1.74
O-Decenoyl-L-carnitine (acyl-C10:1)	CID57357170	314.2374	2.34
O-dodecenoyl-carnitine (acyl-C12:1)	HMDB13326	342.2630	1.73
Tetradecenoyl Carnitine (acyl-C14:1)	HMDB13329	370.3000	2.96
METABOLITES_END
#END