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Blood was combined with PBS at a ratio of 1:2, layered on top of lymphoprep (15ml) and centrifuged for 30 minutes at 300g on break 1. The buffy coat was extracted and washed in PBS (50ml). PBMC were counted and stained with Live/Dead viability stain, CD56 (clone HCD56/NCAM16.2) and CD3 (SK7/UCHT1) by staining in FACS buffer (PBS + 2% FC fetal calf serum (FCS)) for 20 mins on ice. Live CD56+CD3- NK cells were purified by flow sorting using a BD FACSAria™ III Cell Sorter.   NK cell proteomics analysis:  • Culture of NK cells for proteomic analysis Pure NK cells were seeded at 5x106 cells/ml in RPMI supplemented with 10% FCS and 1% penicillin/ streptomycin and stimulated with IL2 (500IU/ml). Cells were incubated at 37°C, 5% CO2 for 18 hours. NK cells were harvested, washed in ice cold PBS twice and NK cell pellets were snap frozen using liquid nitrogen.   • Proteomic sample preparation Cells were lysed in 5% SDS and 50mM TEAB. Protein concentrations were determined using a micro-BCA assay. Aliquots containing 100 µg of protein were processed using the S-Trap mini protocol (Protifi) as recommended by the manufacturer, with minor modifications. Disulfide bonds were reduced with 20 mM DTT for 10 min at 95 °C, followed by alkylation with 40 mM IAA for 30 min in the dark. After loading the samples onto the S-Trap mini spin columns, the trapped proteins were washed five times with 500 µl of S-Trap binding buffer.   Proteins were digested with trypsin (Pierce, Thermo Fisher) at a 1:40 enzyme-to-protein ratio in a two-step digestion: first overnight at 37 °C in 160 µl of 50 mM TEAB, and then for an additional 6 h at the same temperature. Peptides were eluted from the S-Trap mini columns by adding 160 µl of 50 mM TEAB, followed by 160 µl of 0.2% aqueous formic acid, and finally 160 µl of 50% acetonitrile containing 0.2% formic acid, with centrifugation at 1,000 × g for 1 min after each addition. Tryptic peptides were dried in a SpeedVac and stored at −20 °C.  • Mass Spectrometry Peptides generated from NK cells were analyzed by Data-Independent Acquisition (DIA) on a Q-Exactive™ plus Mass Spectrometer (Thermo Scientific) coupled to a Dionex Ultimate 3000 RS (Thermo Scientific). LC buffers used are the following:  buffer A (0.1% formic acid in Milli-Q water (v/v)) and buffer B (80% acetonitrile and 0.1% formic acid in Milli-Q water (v/v). A total of 1.5 µg of each sample was loaded at 10 μL/min onto a µPAC trapping C18 column (Pharmafluidics). The trapping column was washed for 6 min at the same flow rate with 0.1% TFA and then switched in-line with a Pharma Fluidics, 200 cm, µPAC nanoLC C18 column. The column was equilibrated at a flow rate of 300nl/min for 30 min. The peptides were eluted from the column at a constant flow rate of 300 nl/min with a linear gradient from 1% buffer B to 3.8% buffer B in 6 min, from 3.8% B to 12.5% buffer B in 40 min, from 12.5% buffer B to 41.3% buffer B within 176 min and then from 41.3 % buffer B to 61.3% buffer B in 14 min. The gradient is finally increased from 61.3% buffer B to 100% buffer B in 1 min, and the column was then washed at 100% buffer B for 10 min.  Two blanks were run between each sample to reduce carry-over.  The column was kept at a constant temperature of 50oC.   Q-Exactive plus was operated positive ionization mode using an easy spray source. The source voltage was set to 2.2 Kv and the capillary temperature was 275oC. Data were acquired in DIA Mode.  A scan cycle comprised a full MS scan (m/z range from 345-1155), resolution was set to 70,000, AGC target 3 x 106, maximum injection time 200 ms.  MS survey scans were followed by DIA scans of dynamic window widths with an overlap of 0.5 Th. DIA spectra were recorded at a resolution of 17,500 at 200 m/z using an automatic gain control target of 3 x 106, a maximum injection time of 55 ms and a first fixed mass of 200 m/z. Normalised collision energy was set to 25 % with a default charge state set at 3. Data for both MS scan and MS/MS DIA scan events were acquired in profile mode.  Proteomic sample preparation and mass spectrometry was conducted by the FingerPrints Proteomics facility at Dundee University."],"repository":["Pride"],"quantification_method":["Not available"],"modification":[""],"data_protocol":["Raw DIA mass spectrometry data files were searched using Spectronaut  (Biognosys) version 19. Raw MS files were searched against a mouse database (Swissprot Trembl, November 2023) with the following parameters: directDIA, false discovery rate set to 1%, protein N-terminal acetylation and methionine oxidation were set as variable modifications, and carbamidomethylation of cysteine residues was selected as a fixed modification. Perseus software (REF Tyanova, S. et al. The Perseus computational platform for comprehensive analysis of (prote)omics data. Nat. Methods13, 731–740 (2016)) was used to estimate protein copy numbers according to the method described in Wisniewski et al. (REF: Wiśniewski, J. R., Hein, M. Y., Cox, J. & Mann, M. A “proteomic ruler” for protein copy number and concentration estimation without spike-in standards. Mol. Cell. Proteomics 13, 3497–3506 (2014))"],"omics_type":["Proteomics"],"labhead":["Professor Clair Gardiner"],"instrument_platform":[""],"submission_type":["PARTIAL"],"labhead_affiliation":["Trinity College Dublin: Dublin, IE"],"species":["Homo Sapiens (human)"],"publication":["Not available"],"submitter_mail":["a.howden@dundee.ac.uk"],"submitter_affiliation":["University of Dundee"],"submitter_country":["United Kingdom"],"additional_accession":[]},"is_claimable":false,"name":"Metabolic Tracing Reveals IL-2 Driven Glutaminolysis and De Novo Pyrimidine Synthesis in Human Natural Killer Cells","description":"Natural Killer (NK) cells are innate lymphocytes that are key to intrinsic cancer immunosurveillance and an important target for cancer immunotherapy. Understanding fundamental human NK cell metabolism provides opportunities for optimising NK cell therapies. Little is known about how glutamine, an important cell nutrient and carbon source, is utilised by human NK cells.  To address this, we performed U13C-glutamine tracing experiments by Liquid Chromatography Mass Spectrometry (LCMS) and Gas Chromatography Mass Spectrometry (GCMS) analysis of human NK cells stimulated with IL-2 for 18 hours to provide a global overview of glutamine usage by these cells.  Our results show that glutamine is taken up by resting NK cells and that this increases further upon IL-2 stimulation. Metabolite labelling analysis identified that IL-2 results in greater conversion of glutamine to glutamate, allowing for anaplerotic flux into the TCA cycle. The fate of the glutamine-derived carbons diverged at oxaloacetate (OAA) allowing both bioenergetic and biosynthetic outcomes - some carbons continued around the TCA cycle while others were exported, converted to aspartate and subsequently used for pyrimidine synthesis. Nucleotide synthesis by IL-2 activated NK cells was found to be essential for expression of the activation marker CD69.  The data indicate that glutamine is a key nutrient taken up by human NK cells, and that IL-2 drives glutaminolysis.  Subsequent glutamate is used to support the TCA cycle, generating energy and providing intermediates for de novo pyrimidine synthesis.","dates":{"publication":"2026-04-01","submission":"2026-04-01"},"accession":"PXD076428","cross_references":{"TAXONOMY":["NEWT:108018","NEWT:377960","NEWT:1131","NEWT:190486","NEWT:2042546","NEWT:148942","NEWT:295546","NEWT:258594","NEWT:1126","NEWT:1129","NEWT:89187","NEWT:309807","NEWT:89184","NEWT:309800","NEWT:1323429","NEWT:876138","NEWT:44271","NEWT:193516","NEWT:2315688","NEWT:44270","NEWT:592321","NEWT:1073882","NEWT:6819","NEWT:428386","NEWT:1117","NEWT:498257","NEWT:10036","NEWT:32264","NEWT:10034","NEWT:661410","NEWT:488003","NEWT:224326","NEWT:376619","NEWT:3765","NEWT:2433","NEWT:3767","NEWT:433727","NEWT:135622","NEWT:672127","NEWT:656061","NEWT:37965","NEWT:656064","NEWT:3760","NEWT:36630","NEWT:381198","NEWT:10029","NEWT:913645","NEWT:641809","NEWT:44250","NEWT:317447","NEWT:5530","NEWT:868565","NEWT:135674","NEWT:1562698","NEWT:11308","NEWT:12637","NEWT:2164133","NEWT:1180","NEWT:2767846","NEWT:1343074","NEWT:314344","NEWT:108061","NEWT:224308","NEWT:49928","NEWT:160621","NEWT:110368","NEWT:1916073","NEWT:1310161","NEWT:1230383","NEWT:77133","NEWT:1310165","NEWT:114727","NEWT:5516","NEWT:498211","NEWT:347256","NEWT:5518","NEWT:398007","NEWT:1527468","NEWT:498217","NEWT:498216","NEWT:11320","NEWT:1410384","NEWT:1310173","NEWT:161934","NEWT:1148","NEWT:5508","NEWT:419481","NEWT:5507","NEWT:1140","NCBITaxon:2157","NEWT:2066563","NEWT:1143","NEWT:1462472","NEWT:1336795","NEWT:644042","NEWT:709964","NEWT:3712","NEWT:3711","NEWT:3714","NEWT:65121","NEWT:3715","NEWT:2065263","NEWT:1549675","NEWT:177437","NEWT:9014","NEWT:10891","NEWT:118221","NEWT:34869","NEWT:226126","NEWT:1616117","NEWT:34865","NEWT:550537","NEWT:284812","NEWT:3701","NEWT:3702","NEWT:1245466","NEWT:3704","NEWT:1246791","NEWT:34875","NEWT:9005","NEWT:34871","NEWT:9006","NEWT:33548","NEWT:1883426","NEWT:657321","NEWT:1423687","NEWT:10886","NEWT:3708","NEWT:536231","NEWT:219813","NEWT:1301262","NEWT:2305479","NEWT:215451","NEWT:460519","NEWT:272843","NEWT:272844","NEWT:1348799","NEWT:55784","NEWT:485","NEWT:2499525","NEWT:467144","NEWT:142786","NEWT:2853422","NEWT:487","NEWT:1679718","NEWT:480","NEWT:67767","NEWT:46835","NEWT:216778","NEWT:376686","NEWT:237610","EFO:0001352","NEWT:219809","NEWT:219808","NEWT:119562","NEWT:1883446","NEWT:1233435","NEWT:118223","NEWT:141469","NEWT:272831","NEWT:987059","NEWT:491","NEWT:460523","NEWT:2423","NEWT:85694","NEWT:85692","NEWT:911045","NEWT:3750","NEWT:291146","NEWT:233271","NEWT:3745","NEWT:1715989","NCBITaxon:4751","NEWT:3747","NEWT:278161","NEWT:415978","NEWT:410289","NEWT:191701","NEWT:9054","NEWT:472","NEWT:29459","NEWT:28128","NEWT:470","NEWT:41122","NEWT:1328300","NEWT:748693","NCBITaxon:50557","NEWT:3735","NEWT:29491","NEWT:911022","NEWT:446","NEWT:153481","NEWT:714485","NEWT:445","NEWT:2003758","NEWT:29486","NEWT:63366","NEWT:63367","NEWT:29488","NEWT:215402","NEWT:3726","NEWT:30455","NEWT:28141","NEWT:9030","NEWT:9031","NEWT:3728","NEWT:108931","NEWT:219799","NEWT:1325292","NEWT:219798","NEWT:548","NEWT:150475","NEWT:256737","NEWT:51350","NEWT:8694","NEWT:8692","NEWT:542","NEWT:394503","NEWT:13076","NEWT:219785","NEWT:451515","NEWT:550","NEWT:10804","NEWT:554","NEWT:451516","NEWT:552","NEWT:1325291","NEWT:42410","NEWT:1225786","NEWT:771870","NEWT:8665","NEWT:51338","NEWT:9995","NEWT:28112","NEWT:49012","NEWT:28110","NEWT:10832","NEWT:311339","NEWT:507601","NEWT:520","NEWT:64657","NEWT:186441","NEWT:2585209","NEWT:643680","NEWT:214092","NEWT:40697","NEWT:13095","NEWT:162425","NEWT:242124","NEWT:9986","NEWT:8654","NEWT:536","NEWT:8658","NEWT:1268063","NEWT:8657","NEWT:9983","NEWT:8655","NEWT:80833","NEWT:479096","NEWT:28108","NEWT:28104","NEWT:1114957","NEWT:568703","NEWT:568708","NEWT:52641","NEWT:353152","NEWT:116597","NEWT:353153","NEWT:116598","NEWT:227377","NEWT:1774284","NEWT:27689","NEWT:207769","NEWT:40674","NEWT:51329","NCBITaxon:1763","NEWT:443906","NEWT:13037","NEWT:519","NEWT:941638","NEWT:7394","NEWT:6063","NEWT:404330","NEWT:1328388","NEWT:13064","NEWT:300641","NEWT:39655","NEWT:38323","NEWT:206411","NEWT:80869","NEWT:214053","NEWT:80863","NEWT:665890","NEWT:52639","NEWT:52638","NEWT:8697","NEWT:861109","NEWT:8695","NEWT:1196866","NEWT:6039","NEWT:1045775","NEWT:1123869","NCBITaxon:1335626","NEWT:7370","NEWT:280406","NEWT:83906","NEWT:99488","NEWT:99487","NEWT:1134506","NEWT:255470","NEWT:762903","NEWT:27697","NEWT:86185","NEWT:9935","NEWT:266940","NEWT:287889","NEWT:9938","NEWT:1028307","NEWT:52603","NEWT:92867","NEWT:92866","NEWT:1344414","NEWT:202950","NEWT:741158","NEWT:1293091","NEWT:202956","NEWT:483179","NEWT:315750","NEWT:99490","NEWT:632957","NEWT:11294","NEWT:9925","NCBITaxon:9606","NEWT:1436183","NEWT:1479235","NEWT:4232","NEWT:9922","NEWT:416348","NEWT:105812","NEWT:11298","NEWT:4233","NEWT:4236","NEWT:398031","NEWT:196627","NEWT:523840","NEWT:487521","NEWT:1076878","NEWT:200308","NEWT:200302","NEWT:9913","NEWT:9915","NEWT:105841","NEWT:1170227","NEWT:2666255","NEWT:999810","NEWT:28995","NEWT:1392998","NEWT:1150621","NEWT:1085791","NEWT:745531","N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