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Deuterium labeling: Deuterium labeling of 1 μL of the equilibration reaction was initiated with an 18-fold dilution into D2O labeling buffer (18 μL, 20 mM HEPES-KOH, 150 mM KCl, 5 mM MgCl2, pD 7.6, 99.9% D2O).  After each labeling time (10 seconds, 1 minute, 10 minutes) at 23 °C (room temperature), the labeling reaction was quenched with the addition of 19 μL of ice-cold quenching buffer (0.8 M guanidinium chloride, 0.8% formic acid pH 2.4, H2O) and analyzed immediately.   LC/MS: Deuterated and control samples were digested online at 15 °C using a AffiPro Nep-2 column (AffiPro, AP-PC-004). The cooling chamber of the HDX system, which housed all the chromatographic elements, was held at 0.0 ± 0.1 °C for the entire time of the measurements. Peptides were trapped and desalted on a VanGuard Pre-Column trap [2.1 mm × 5 mm, ACQUITY UPLC BEH C18, 1.7 μm (Waters, 186002346)] for 3 minutes at 100 μL/min.  Peptides were eluted from the trap using a 5%-35% gradient of acetonitrile with 0.1% formic acid over 6 minutes at a flow rate of 100 μL/min, and separated using an ACQUITY UPLC HSS T3, 1.8 μm, 1.0 mm × 50 mm column (Waters, 186003535).  The back pressure averaged ~12,950 psi at 0 °C and 5% acetonitrile 95% water, 0.1% formic acid.  To eliminate peptide carryover, a wash solution [1.5 M guanidinium chloride, 0.8% formic acid and 4% acetonitrile] was injected over the protease column during each analytical run.  Mass spectra were acquired using a Waters Synapt G2-Si HDMSE mass spectrometer in ion mobility mode.  The mass spectrometer was calibrated with direct infusion of a solution of glu-fibrinopeptide (Sigma, F3261) at 200 femtomole/μL at a flow rate of 5 μL/min prior to data collection.  A conventional electrospray source was used, and the instrument was scanned over the range 50 to 2000 m/z.  The instrument configuration was the following: capillary was 3.2 kV, trap collision energy at 4 V, sampling cone at 40 V, source temperature of 80 °C and desolvation temperature of 175 °C. The error of determining the average deuterium incorporation for each peptide was at or below ± 0.25 Da, based on deuterated peptide standards."],"repository":["Pride"],"modification":[""],"quantification_method":["Not available"],"data_protocol":["Peptides were identified from replicate (as detailed in the Supplemental Datafile) HDMSE analyses of undeuterated control samples using PLGS 3.0.1 (Waters Corporation). Peptide masses were identified from searches using non-specific cleavage of a custom database containing the sequence of BAX (UniProt: Q07812) with mutation F116A, no missed cleavages, no PTMs, a low energy threshold of 135, an elevated energy threshold of 35 and an intensity threshold of 500. No false discovery rate (FDR) control was performed.  The peptides identified in PLGS (excluding all neutral loss and in-source fragmentation identifications) were then filtered in DynamX 3.0 (Waters Corporation) implementing a minimum products per amino acid cut-off of 0.25, at least 1 consecutive product ion (see Supplemental Datafile).  Those peptides meeting the filtering criteria to this point were further processed by DynamX 3.0 (Waters Corporation). The relative amount of deuterium in each peptide was determined by the software by subtracting the centroid mass of the undeuterated form of each peptide from the deuterated form, at each time point, for each condition.  The deuterium uptake values (provided in the Supplemental Datafile) were used to generate all difference maps found in the Figures as well as the Supplemental Datafile.  Deuterium levels were not corrected for back exchange and thus reported as relative [Wales & Engen (2015). Mass Spectrom Rev. 25(1), 158-170]."],"omics_type":["Proteomics"],"labhead":["Thomas E. Wales"],"instrument_platform":[""],"labhead_affiliation":["Northeastern University"],"submission_type":["PARTIAL"],"species":["Mus Musculus (mouse)"],"submitter_mail":["t.wales@northeastern.edu"],"publication":["Not available"],"submitter_affiliation":["Northeastern University"],"submitter_country":["United States"],"additional_accession":[]},"is_claimable":false,"name":"Discovery of an Enantioselective Covalent Inhibitor of BAX for Cytoprotection in Vivo","description":"The pro-apoptotic BCL-2 protein BAX is a central executioner of mitochondrial apoptosis and a driver of acute and chronic cell loss in diverse human diseases. The C126 residue of BAX lies within a key regulatory region and its derivatization by physiologic and pharmacologic ligands can alternatively activate or inhibit BAX function. Here, we report the discovery of an enantioselective covalent BAX inhibitor that confers cytoprotection across multiple cell types and in vivo. Chemoproteomic screening identified tryptoline acrylamide stereoprobes that derivatized BAX at C126 with high selectivity, constraining its conformation, and suppressing apoptosis in a strictly BAX-dependent manner, as confirmed by genetic knockout models. Medicinal chemistry optimization yielded covalent BAX inhibitor 3 (CBI-3), a more stable analog with retained enantioselectivity, improved pharmacokinetics, and dose-responsive cytoprotective activity against ABT-737 and cisplatin-induced apoptosis without intrinsic toxicity. In a murine model of Fas-induced fulminant hepatic failure, CBI-3 significantly reduced hepatocyte apoptosis and preserved liver histology and animal survival, nearly phenocopying Bax deletion. These findings establish enantioselective covalent inhibition of BAX as a viable strategy for cytoprotection with demonstrable in vivo efficacy, opening the door to next-generation therapeutics that directly block pathologic cell death.","dates":{"publication":"2026-06-28","submission":"2025-10-16"},"accession":"PXD069506","cross_references":{"TAXONOMY":["NEWT:330879","NEWT:377960","NEWT:2042546","NEWT:259447","NEWT:295546","NCBITaxon:2719036","NCBITaxon:1280","NEWT:112503","NEWT:1129","NEWT:309807","NEWT:59753","NEWT:89184","NEWT:309800","NEWT:281395","NEWT:1211601","NEWT:876138","NEWT:44271","NEWT:193516","NEWT:1073882","NEWT:6819","NEWT:475174","NEWT:111205","NEWT:1117","NEWT:498257","NEWT:10036","NEWT:1590","NEWT:10034","NEWT:661410","NEWT:638632","NEWT:224326","NEWT:376619","NCBITaxon:79857","NEWT:1096976","NEWT:3765","NEWT:1589","NEWT:135622","NEWT:35786","NEWT:1580","NEWT:399784","NEWT:96731","NEWT:656064","NEWT:1582","NEWT:36630","NEWT:383379","NEWT:418106","NCBITaxon:12721","NEWT:10029","NEWT:913645","NEWT:641809","NEWT:1392488","NEWT:38815","NEWT:317447","NEWT:4688","NEWT:111225","NEWT:7719","NEWT:868565","NEWT:135674","NEWT:79329","NEWT:30069","NEWT:12637","NEWT:59729","NEWT:2164133","NEWT:295105","NEWT:108061","NEWT:60711","NEWT:224308","NEWT:3347","NEWT:160621","NEWT:212790","NEWT:110368","NEWT:1310161","NEWT:77133","NEWT:145481","NEWT:1310165","NEWT:29058","NCBITaxon:79824","NEWT:1912919","NEWT:410658","NEWT:44688","NEWT:44689","NEWT:498211","NEWT:347256","NEWT:5518","NEWT:398007","NEWT:1527468","NEWT:498217","NEWT:498216","NEWT:11320","NEWT:246196","NEWT:246197","NEWT:145458","NEWT:44685","NEWT:161934","NEWT:156587","NEWT:1148","NEWT:5508","NEWT:3329","NEWT:419481","NEWT:5507","NEWT:410661","NEWT:1140","NCBITaxon:2157","NEWT:1143","NEWT:1287689","NEWT:1094343","NEWT:1462472","NEWT:1336795","NEWT:644042","NEWT:1182590","NEWT:3712","NEWT:3711","NEWT:3714","NEWT:270643","NEWT:2065263","NEWT:177437","NEWT:10418","NEWT:118698","NEWT:1616117","NEWT:118696","NEWT:34865","NEWT:52283","NEWT:28174","NEWT:284812","NEWT:8175","NEWT:43330","NEWT:1603293","NEWT:44664","NEWT:43346","NEWT:3702","NEWT:1245466","NEWT:3704","NEWT:244366","NEWT:1246791","NEWT:118694","NEWT:2850","NEWT:118691","NEWT:34871","NEWT:33548","NEWT:52299","NEWT:8165","NEWT:96794","NEWT:3708","NEWT:332648","NEWT:44670","NEWT:536231","NEWT:376219","NEWT:219813","NEWT:1510","NEWT:460519","NEWT:1515","NEWT:572307","NEWT:1432138","NEWT:1424507","NEWT:1194599","NEWT:272844","NEWT:1348799","NEWT:33995","NEWT:55784","NEWT:1303443","NEWT:9483","NEWT:485","NEWT:56636","NEWT:2709072","NEWT:2499525","NEWT:2853422","NEWT:1679718","NEWT:480","NEWT:67767","NEWT:46835","NEWT:109757","NEWT:582580","NEWT:1185650","NEWT:216778","NEWT:294607","NEWT:1502","NEWT:128017","NEWT:376686","NEWT:237610","NEWT:95486","NEWT:1883446","NEWT:1233435","NEWT:118223","NEWT:109760","NEWT:29031","NEWT:317047","NEWT:491","NEWT:460523","NEWT:235443","NEWT:1757312","NEWT:85694","NEWT:109786","NEWT:108458","NEWT:5936","NEWT:320637","NEWT:1078283","NEWT:3750","NEWT:983964","NEWT:11706","NEWT:32644","NEWT:527796","NEWT:499175","NEWT:109779","NEWT:3745","NEWT:1715989","NCBITaxon:4751","NEWT:3747","NEWT:1116234","NEWT:1255228","NEWT:410289","NEWT:373153","NEWT:1640278","NEWT:472","NEWT:940825","NEWT:529507","NEWT:1071661","NEWT:28128","NEWT:470","NEWT:5911","NEWT:748693","NCBITaxon:50557","NEWT:39251","NEWT:29491","NEWT:101841","NEWT:446","NEWT:153481","NEWT:2014887","NEWT:33952","NEWT:445","NEWT:153009","NEWT:261756","NEWT:166313","NEWT:63366","NEWT:63367","NEWT:215402","NEWT:1547","NEWT:906914","NEWT:9031","NEWT:27292","NEWT:1050722","NEWT:3728","NEWT:108931","NEWT:1293497","NEWT:1055524","NEWT:150475","NEWT:267872","NEWT:381666","NEWT:172269","NEWT:9534","NEWT:5180","NEWT:256737","NEWT:9541","NEWT:8694","NEWT:33936","NEWT:8692","NEWT:2903","NEWT:185579","NEWT:13076","NEWT:207340","NEWT:1006581","NEWT:8673","NEWT:33940","NEWT:550","NEWT:554","NEWT:451516","NEWT:552","NEWT:1325291","NEWT:36185","NEWT:1054211","NEWT:1225786","NEWT:771870","NEWT:51338","NEWT:575412","NEWT:28112","NEWT:6493","NEWT:10832","NEWT:6494","NEWT:6491","NEWT:507601","NEWT:520","NEWT:186441","NEWT:643680","NEWT:214092","NCBITaxon:6157","NEWT:13095","NEWT:162425","NEWT:104105","NEWT:216257","NEWT:9986","NEWT:8654","NEWT:8658","NEWT:1268063","NEWT:8657","NEWT:9983","NEWT:8655","NEWT:5147","NEWT:28108","NEWT:28104","NEWT:407821","NEWT:568703","NCBITaxon:2","NEWT:568708","NEWT:1613","NEWT:986","NEWT:52641","NEWT:28532","NEWT:353152","NEWT:298176","NEWT:227377","NEWT:1774284","NEWT:40674","NEWT:1194669","NEWT:51329","NEWT:443906","NEWT:519","NEWT:2510939","NEWT:7394","NEWT:6063","NEWT:1328388","NEWT:105487","NEWT:1548728","NEWT:667127","NEWT:9557","NEWT:377586","NEWT:300641","NEWT:39655","NEWT:9554","NEWT:38323","NEWT:475932","NEWT:256318","NEWT:206411","NCBITaxon:6191","NEWT:229533","NEWT:214053","NEWT:28550","NEWT:80863","NEWT:90675","NEWT:52639","NEWT:52638","NEWT:57075","NEWT:1666905","NEWT:8697","NEWT:8695","NEWT:884204","NEWT:510516","NEWT:1045775","NEWT:1123869","NEWT:9544","NEWT:9545","NEWT:979","NEWT:7370","NEWT:83906","NEWT:99488","NEWT:1134506","NEWT:255470","NEWT:38783","NEWT:6426","NEWT:33090","NEWT:9935","NEWT:287889","NEWT:3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