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Following FACS, the sorted cells were pelleted and stored at -20ºC until proteomic sample preparation. Subsequently, sorted T and B cells were resuspended in 50 mM Tris pH 8.5 and 1% sodium deoxycholate and the proteins were denatured at 95°C for 10 min. Subsequently, the protein amount was determined by BCA (Thermo Fisher Scientific) following the manufacturer's protocol. Individual samples were reduced with 5 mM DTT for 1 h followed by alkylation with 15 mM iodoacetamide in the dark for another hour. Samples were then digested with LysC endopeptidase (an enzyme to protein ratio of 1:100 w/w) for 2 h, followed by digestion using the same amount of trypsin overnight. Samples were desalted by StageTip (Thermo Fisher Scientific) following the manufacturer's protocol. Cleaned samples were dried in a speed vac and stored at -80°C. Samples were resuspended in buffer A (2% ACN, 0.1% FA) and injected into an UltiMate 3000 UPLC autosampler (Thermo Fisher Scientific), which was connected to an Orbitrap Fusion Lumos Tribrid mass spectrometer (Thermo Fisher Scientific). The peptides were loaded on a trap column (Acclaim PepMap 100 C18, 100 μm × 2 cm) and subsequently separated on a 50 cm long C18 Easy spray column (Thermo Fisher Scientific). Chromatographic separation of the peptides was archived by the following 105 min gradient: 4-20 % of solvent B (98% ACN and 0.1% FA) in 80 min, to 26 % in 10 min, to 95 % in 3 min where it was kept for another 3 min, before going to 4 % in 1 min and hold for 8 min. During the entire gradient, the mass spectrometer was operating in positive polarity. All spectra were acquired in profile mode using the Orbitrap mass analyzer. An acquisition cycle consisted of one survey mass spectrum acquired at a mass resolution of 120,000 from m/z 395 to 1005, with an AGC target of 1,000,000 charges and a maximal injection time of 150 ms. This was followed by 40 DIA MS/MS events with variable isolation windows. The individual mass spectra were recorded at a resolution of 30,000 with 28 % NCE; stepped NCE was set as 2%, and a maximal injection time of 54 ms. A library of MS/MS spectra was created with a pooled sample. The data acquisition was the same as for the individual samples with the exception that the DIA MS/MS spectra were recorded over a mass range of 120 m/z at a resolution of 60,000 with a maximum injection time of 54 ms, and an overlap between MS/MS windows of 4 m/z units."],"repository":["Pride"],"quantification_method":["Not available"],"modification":[""],"data_protocol":["Acquired .raw files were converted to mzML format by MSConvert (version 3.0.21258) 66 applying peak picking in the mass spectra with the vendor-provided algorithm (Thermo Fisher Scientific). The database search was performed in FragPipe (v18.0) as described in 67 using the mouse Swissprot database (17,248 entries). Trypsin with up to one missed cleavage was set as a digestion enzyme, and oxidation of methionine and acetylation of the N-terminus were set as variable modifications. Carbamidomethylation of cysteine residues was set as a fixed modification. Peptide length was restricted to 7 to 50 amino acids, and molecular mass from 500 to 5000 Da. The resulting peptide-spectrum matches were adjusted to a 1% false discovery rate with Percolator 68 as part of the Philosopher toolkit (v4.4) 69 and converted to an MS/MS spectra library. DIA files were analyzed by DIA-NN 1.8.1 70.  All further data processing was performed in R (version 4.2.2). Known contaminants and decoy proteins were excluded from further analysis. Only proteins quantified in three out of the five replicates in at least one sample group were considered. Protein abundances were normalized by variance stabilization normalization approach and missing values at random were imputed by k-nearest neighbor method, whereas missing values not at random were imputed by drawing values from a downshifted normal distribution 71. All statistical comparisons were performed based on a two-tailed Student's t-test with equal variances; the differences between the comparisons were reported as log2-scaled fold changes."],"omics_type":["Proteomics"],"labhead":["Rikard Holmdahl"],"instrument_platform":[""],"submission_type":["PARTIAL"],"labhead_affiliation":["Medical Inflammation Research, Division of Immunology, Department of Medical Biochemistry and Biophysics, Karolinska Institute, Stockholm, Sweden"],"species":["Mus Musculus (mouse)"],"publication":["40285352 Romero-Castillo L, Pandey RK, Xu B, Beusch CM, Oliveira-Coelho A, Zeqiraj K, Svensson C, Xu Z, Luo H, Sareila O, Sabatier P, Ge C, Cheng L, Urbonaviciute V, Krämer A, Lindgren C, Haag S, Viljanen J, Zubarev RA, Kihlberg J, Linusson A, Burkhardt H, Holmdahl R. Tolerogenic antigen-specific vaccine induces VISTA-enriched regulatory T cells and protects against arthritis in DRB1∗04:01 mice. Mol Ther. 2025 33(8):3528-3545 10.1016/j.ymthe.2025.04.034"],"submitter_mail":["christian.beusch@gmail.com"],"submitter_affiliation":["Emory University"],"submitter_country":["United States"],"pubmed_abstract":["Rheumatoid arthritis (RA) is a chronic autoimmune disease characterized by joint inflammation, cartilage damage, and bone erosion. Despite improvements with the introduction of biological disease-modifying anti-rheumatic drugs (DMARDs), RA remains an incurable life-long disease. Advancements in peptide-based vaccination may open new avenues for treating autoimmune diseases, including RA, by inducing immune tolerance while maintaining normal immune function. We have already demonstrated the efficacy of a potent vaccine against RA, consisting of the mouse major histocompatibility complex class II (A<sup>q</sup>) protein bound to the immunodominant type II collagen peptide COL2<sub>259-273</sub>, which needed to be galactosylated at position 264. To translate the vaccine to humans and to further enhance vaccine efficacy, we modified the glycine residue at position 265 and conjugated it with the human DRB1∗04:01 molecule. Remarkably, this modified vaccine (named DR4-AL179) provided robust effectiveness in suppressing arthritis in DRB1∗04:01-expressing mice without the need for galactosylation at position 264. DR4-AL179 vaccination induces tolerance involving multiple immunoregulatory pathways, including the activation of V-type immunoglobulin domain-containing suppressor of T cell activation (VISTA)-positive nonconventional regulatory T cells, which contribute to a potent suppressive response preventing arthritis development in mice. This modified RA vaccine offers a novel therapeutic potential for human autoimmune diseases."],"pubmed_title":["Tolerogenic antigen-specific vaccine induces VISTA-enriched regulatory T cells and protects against arthritis in DRB1∗04:01 mice."],"pubmed_authors":["Romero-Castillo Laura L, Pandey Rajan Kumar RK, Xu Bingze B, Beusch Christian M CM, Oliveira-Coelho Ana A, Zeqiraj Kejsi K, Svensson Carolin C, Xu Zhongwei Z, Luo Huqiao H, Sareila Outi O, Sabatier Pierre P, Ge Changrong C, Cheng Lei L, Urbonaviciute Vilma V, Krämer Alexander A, Lindgren Cecilia C, Haag Sabrina S, Viljanen Johan J, Zubarev Roman A RA, Kihlberg Jan J, Linusson Anna A, Burkhardt Harald H, Holmdahl Rikard R"],"additional_accession":[]},"is_claimable":false,"name":"A novel non-galactosylated tolerogenic antigen-specific vaccine protects against arthritis via VISTA-expressing regulatory T cells in DRB1*04:01 mice","description":"Rheumatoid arthritis (RA) is a chronic autoimmune disease characterized by joint inflammation, cartilage damage and bone erosion. Despite improvements with the introduction of biological disease-modifying anti-rheumatic drugs (DMARDs), RA remains incurable.  Advancements in peptide-based vaccination may open new avenues for treating autoimmune diseases by inducing immune tolerance while maintaining normal immune function. We have demonstrated a potential vaccine against RA, made by mouse major histocompatibility complex class II (Aq) protein bounding the immunodominant type II collagen peptide COL2259-273, which needed to be galactosylated at position 264. To translate the vaccine to humans and to further enhance and simplify this vaccine, we modified the amino acid residue at position 265 and conjugated it with the human DRB1*04:01 molecule. Remarkably, this modified vaccine proved effective in DRB1*04:01-expressing mice without the need for galactosylation at position 264. The vaccine exerts its protective effect through the activation and expansion of VISTA-positive nonconventional regulatory T cells, leading to a potent dominant suppressive response that prevents arthritis development in mice. The new modified vaccine for RA, offers a novel therapeutic approach with potential clinical implications for human autoimmune diseases.","dates":{"publication":"2026-06-29","submission":"2024-10-15"},"accession":"PXD056818","cross_references":{"TAXONOMY":["NEWT:68239","NEWT:108018","NEWT:377960","NEWT:1131","NEWT:190486","NEWT:2737160","NEWT:2042546","NEWT:148942","NEWT:295546","NEWT:258594","NEWT:1126","NEWT:1129","NEWT:89187","NEWT:309807","NEWT:89184","NEWT:309800","NEWT:156563","NEWT:1323429","NEWT:876138","NEWT:44271","NEWT:193516","NEWT:2315688","NEWT:44270","NEWT:592321","NEWT:1073882","NEWT:6819","NEWT:93217","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:1176302","NEWT:418106","NEWT:10029","NEWT:913645","NEWT:2492962","NEWT:641809","NEWT:44250","NEWT:77151","NEWT:317447","NEWT:698761","NEWT:5530","NEWT:868565","NEWT:135674","NEWT:1562698","NEWT:1177","NEWT:11308","NEWT:12637","NEWT:2164133","NEWT:1180","NEWT:2767846","NEWT:1343074","NEWT:314344","NEWT:108061","NEWT:224308","NEWT:3134131","NEWT:49928","NEWT:160621","NEWT:110368","NEWT:1916073","NEWT:1310161","NEWT:1230383","NEWT:77133","NCBITaxon:5207","NEWT:160620","NCBITaxon:77643","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:37919","NEWT:1072569","NEWT:1072568","NEWT:161934","NEWT:156587","NEWT:1148","NEWT:5508","NEWT:135666","NEWT:419481","NEWT:5507","NEWT:1140","NCBITaxon:2157","NEWT:2066563","NEWT:1143","NEWT:1336794","NEWT:1462472","NEWT:1336795","NEWT:644042","NEWT:709964","NEWT:3710","NEWT:3712","NEWT:3711","NEWT:3714","NEWT:65121","NEWT:3716","NEWT:3715","NEWT:2065263","NEWT:1549675","NEWT:177437","NEWT:9014","NEWT:10891","NEWT:118221","NEWT:34869","NEWT:226126","NEWT:1616117","NEWT:34865","NEWT:28174","NEWT:550537","NEWT:284812","NEWT:284811","NEWT:2902907","NEWT:3701","NEWT:3702","NEWT:1245466","NEWT:3704","NEWT:1246791","NEWT:34875","NEWT:10882","NEWT:1431297","NEWT:9005","NEWT:34871","NEWT:9006","NEWT:33548","NEWT:1883426","NEWT:657321","NEWT:1423687","NEWT:657322","NEWT:10886","NEWT:3707","NEWT:3708","NEWT:536231","NEWT:219813","NEWT:1301262","NEWT:319224","NEWT:2305479","NEWT:215451","NEWT:460519","NEWT:272843","NEWT:272844","NEWT:1348799","NEWT:55784","NEWT:485","NEWT:223074","NEWT:2709072","NEWT:2499525","NEWT:467144","NEWT:142786","NEWT:28196","NEWT:2853422","NEWT:487","NEWT:1679718","NEWT:387344","NEWT:657318","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:422203","NEWT:460523","NEWT:2423","NEWT:85694","NEWT:85692","NEWT:911045","NEWT:3750","NEWT:123606","NEWT:85698","NEWT:291146","NEWT:29466","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:34839","NEWT:476","NEWT:1380096","NEWT:28128","NEWT:470","NEWT:41122","NEWT:1328300","NEWT:748693","NEWT:106299","NCBITaxon:50557","NEWT:63363","NEWT:3735","NEWT:29491","NEWT:911022","NEWT:446","NEWT:153481","NEWT:714485","NEWT:445","NEWT:2003758","NEWT:29486","NEWT:63366","NEWT:64697","NEWT:63367","NEWT:29488","NEWT:215402","NEWT:3726","NEWT:1862567","NEWT:591090","NEWT:30455","NEWT:28141","NEWT:9030","NEWT:9031","NEWT:3728","NEWT:108931","NEWT:219799","NEWT:1325292","NEWT:219798","NEWT:548","NEWT:150475","NEWT:546","NEWT:1431340","NEWT:256737","NEWT:51350","NEWT:8694","NEWT:8692","NEWT:542","NEWT:394503","NEWT:13076","NEWT:219785","NEWT:451515","NEWT:8673","NEWT:550","NEWT:10804","NEWT:52694","NEWT:554","NEWT:451516","NEWT:552","NEWT:1325291","NEWT:42410","NEWT:40686","NEWT:1225786","NEWT:771870","NEWT:8665","NEWT:51338","NEWT:377142","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:595495","NEWT:80833","NEWT:479096","NEWT:28108","NEWT:28104","NEWT:1114957","NEWT:568703","NEWT:568708","NEWT:51315","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:6055","NEWT:13037","NEWT:518","NEWT:519","NEWT:941638","NEWT:511","NEWT:7394","NEWT:6063","NEWT:404330","NEWT:1328388","NEWT:13064","NEWT:529120","NEWT:7375","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:37003","N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