{"database":"Pride","file_versions":[{"headers":{"Content-Type":["application/json"]},"body":{"files":{"Xlsx":["ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/Metadata.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/AcquisitionParameters.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/SAINTexpressPPIScoring.xlsx"],"Txt":["ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/proteinGroups.txt","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/checksum.txt","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/evidence.txt","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/msmsScans.txt","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/msms.txt","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/parameters.txt"],"Xml":["ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/mqpar.xml"],"Raw":["ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis003187.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis002798.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis003197.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis002790.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis002800.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis003195.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis002794.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis003191.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis003189.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis003193.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis002792.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis003199.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis002796.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/exLewis003185.raw"],"Fasta":["ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/04/PXD071892/uniprot_human_canonical_9606_052523.fasta"]},"type":"primary"},"statusCode":"OK","statusCodeValue":200}],"scores":null,"additional":{"labhead_mail":["alexander.marson@ucsf.edu"],"submitter":["Robyn Kaake"],"technology_type":["Affinity purification coupled with mass spectrometry proteomics","Mass Spectrometry","Bottom-up proteomics"],"disease":["Human Immunodeficiency Virus Infectious Disease"],"software":[""],"submitter_keywords":["Ip-ms","Hiv","Ppi","Pi16","Host factor","Cd4+ t cells"],"full_dataset_link":["https://www.ebi.ac.uk/pride/archive/projects/PXD071892"],"tissue":["T Cell","Blood"],"sample_protocol":["PI16 was overexpressed in primary human CD4+ T cells from two donors via CRISPR activation. Briefly, activated CD4⁺ T cells were transduced with dCas9-blast lentivirus together with either PI16-targeting sgRNA-puro lentivirus or a control-sgRNA-puro lentivirus. Transduced cells were selected with blasticidin and puromycin. PI16 overexpressing cells and control-guide treated cells (undetectable PI16 expression) were centrifuged at 700xg for 10 minutes, washed in cold-PBS three times, pelleted, and flash frozen. Multiple pellets of 10e6 cells per condition were frozen for each condition. Cell pellets were thawed and suspended in 1 mL Lysis Buffer (IP buffer: [50mM Tris-HCl, pH 7.4 at 4°C, 150mM NaCl, 1mM EDTA], supplemented with 0.5% Nonidet P 40 (NP40; Fluka Analytical), and cOmplete mini EDTA-free protease and PhosSTOP phosphatase inhibitor cocktails (Roche). After two freeze-thaw cycles (> 20 min on dry ice and partial thawing at 22°C using a ThermoMixer C: 1100 RPM, 3-5 min), cell lysates were frozen on dry ice and kept at -80°C until immunoprecipitation (IP). Lysis was completed by partially thawing frozen samples at 22°C with a ThermoMixer C before incubating them on a tube rotator at 4°C for 30 min. After removing cell debris at 13,000xg, 4 °C for 15 min, PI16 overexpressing and control clarified lysates were incubated with either 2.5 mg PI16 rabbit antibody (α-PI16; Proteintech, #12267-1-AP) or 1mg rabbit IgG isotype control (IgG ctrl; Proteintech, #501003118). Antibody and lysate were allowed to bind for 2 hrs on a tube rotator at 4°C. 50 μL Pierce Protein A/G magnetic beads (Thermo Fisher, #88802) were equilibrated and washed three times with 1.0 mL Lysis Buffer before adding to the lysate-antibody mixture for an additional 2 hr binding at 4°C. Protein-bound beads were then washed two times with 1.0 ml Wash Buffer (IP Buffer supplemented with 0.05% NP40), followed by another two washes with IP Buffer. Beads were resuspended in 50 μL of 2M Urea Buffer [2M Urea, 50mM Tris-HCl, pH 8.0, 1mM DTT, 3mM Iodoacetamide, HPLC water] for reduction and alkylation. Samples were incubated for 45 min at 22°C while shaking at 700 RPM. The reduction/alkylation reaction was quenched with an additional 3mM DTT. Proteins were then incubated in a 37°C water bath overnight with 4 μl trypsin (0.5 mg/ml; Promega). Beads were removed, and peptides were acidified with formic acid until the samples reached a pH < 2. Peptides were desalted with OMIX C18 zip tips (Agilent Technologies, #A57003100) according to the manufacturer’s instructions with the following adjustments: 1) tips were conditioned once with 100% ACN, and two times with 50% ACN, 0.1% formic acid (FA); 2) samples were equilibrated in 0.1% FA; 3) after binding, samples were washed 3x in 0.1% FA; 4) peptides were eluted 1x with 50% ACN, 0.1% FA and 1x with 90% ACN, 0.1% FA. After desalting, peptides were dried by speedvac vacuum centrifugation and stored at -20ºC until prepared for liquid chromatography-tandem mass spectrometry (LC-MS/MS) analysis. Prior to LC-MS/MS, each of the fourteen IP digested samples (i.e. three IPs: α-PI16 IPs from PI16 overexpressing cells;  four IPs: α-PI16 IPs from control guide treated PI16 non-expressing cells; three IPs: IgG ctrl IPs from PI16 overexpressing cells; four IPs: IgG ctrl IPs from control guide treated PI16-non expressing cells (see Metadata Table)) were re-suspended in 12μL 3% ACN, 2% FA solution, filtered with a Multiscreen® 96 well Plate, hydrophilic PVDF membrane (Millipore Sigma, #MSHVN4510), and filters washed with 8μL 3% ACN, 2% FA). For each sample, 1μL was injected for LC-MS/MS analysis on a Vanquish Neo (Thermo Fisher Scientific) coupled online with an Orbitrap Exploris 480 mass spectrometer (Thermo Fisher Scientific). Peptides were separated on a 150μm x 15cm 1.5μm PepSep column (Bruker) with a linear 40-minute gradient at a flow rate of 600nL/minute (details in Acquisition Parameters Table). Buffer A consisted of 0.1% FA in water, and buffer B was 0.1% FA in 80% acetonitrile. For each cycle, one full MS scan (350-1500 m/z, at 120,000 resolution with a normalized AGC target (%) of 300) was followed by data-dependent MS/MS scans (data-dependent mode=cycle time; cycle time = 2 sec)(details in Acquisition Parameters Table)."],"repository":["Pride"],"quantification_method":["Not available"],"modification":[""],"data_protocol":["All raw MS files were searched using MaxQuant (version 2.4.7.0) against the human proteome (UniProt human canonical sequences; downloaded 25 May 2023). Trypsin/P was selected as the protease, two missed cleavages were allowed, and variable modifications were set for methionine oxidation and N-terminal protein acetylation. One static modification was set for carbamidomethyl cysteine. Label-free quantitation (LFQ) was enabled. All other MaxQuant settings were left at the default. MaxQuant search data were scored for protein-protein interaction (PPI) determination using SAINT express with the α-PI16:control guide condition being used as the control, and the other three IP conditions (α-PI16:PI16 overexpression; IgG ctrl:NTC ctrl; IgG ctrl:PI16 overexpression) being treated as independent baits. Proteins with a minimum average spectral count (AvgSpec) > 2, and BFDR < 0.01 were considered high-confidence PI16 interactions (hits).  The IgG ctrl: WT and IgG ctrl:PI16_OE conditions were used as a second filter, with any protein “hits” found in those IPs being removed from our final list as false positives. The high-confidence PI16 interaction network was visualized using Cytoscape (version 3.8.3) with round nodes depicting high-confidence PI16 interacting proteins. Proteins with known pro- or anti-HIV functions (from the literature review) are shown with colored borders, and proteins found to be either proviral or antiviral in the CRISPRa and CRISPRn screens were depicted as filled color nodes. Two additional human proteins key to HIV infection (co-receptor CXCR4 and an antiviral hit from our screen, CORO1A) were added to the network as filled-in V nodes to highlight connections within the PI16 network to these key HIV host factors. Additional known interactions were extracted from complexes in CORUM (https://pubmed.ncbi.nlm.nih.gov/30357367/ downloaded on March 1, 2024 using the OmniPathR package v3.4.0 https://pubmed.ncbi.nlm.nih.gov/27898060/ ), all pairs in STRING (v11.5 https://pubmed.ncbi.nlm.nih.gov/36370105/)  with [combined confidence score > 0.50][confidence from experimental evidence > 0.50], and pairs from HumanNet (v3.0 https://pubmed.ncbi.nlm.nih.gov/34747468/ downloaded November 8, 2023) [including the entire Gold Standard set ][as well as Functional Network pairs with scores greater than 2.0]. and mapped in the network as dotted line edges."],"omics_type":["Proteomics"],"labhead":["Alexander Marson"],"instrument_platform":[""],"submission_type":["PARTIAL"],"labhead_affiliation":["Gladstone Institure for Immunology; The J David Gladstone Institutes; San Francisco, CA, USA"],"species":["Homo Sapiens (human)"],"submitter_mail":["robyn.kaake@gladstone.ucsf.edu"],"publication":["42013838 Rathore U, Dugan E, Thornton H, Kumar VE, Dajani R, Burdick RC, Young JM, Steinhart Z, Lao R, Delviks-Frankenberry KA, Choi W, Henriques WS, Echeverria I, Dann E, Dureja I, Pathak N, Arce MM, McKetney J, Umhoefer JM, Parulekar S, Schmidt R, Polacco BJ, Neidleman J, Montano M, Nguyen VQ, Sali A, Levy JA, Tenthorey JL, Cheng Y, Roan NR, Swaney DL, Kaake RM, Dodgson SE, Hiatt J, Pathak VK, Malik HS, Krogan NJ, Marson A. Systematic discovery of pro- and anti-HIV host factors in primary human CD4+ T cells. Cell. 2026:S0092-8674(26)00382-X 10.1016/j.cell.2026.03.046"],"submitter_affiliation":["Department of Cellular and Molecular Pharmacology, University of California San Francisco, San Francisco, CA, USA"],"submitter_country":["United States"],"pubmed_abstract":["Host factors that promote or restrict human immunodeficiency virus (HIV) infection in human CD4+ T cells have not been comprehensively identified. We employed orthogonal genome-wide CRISPR activation (CRISPRa) and CRISPR knockout screens in primary CD4+ T cells to discover pro- and anti-HIV host factors systematically. Secondary pooled screens and individual perturbations validated high-confidence hits and revealed diverse mechanisms of action. CRISPRa uncovered multiple potent antiviral factors, including PI16, PPID, SHISA3, and ITM2A. PI16 interacts with host factors involved in HIV fusion and inhibits viral entry, whereas PPID (Cyp40), a paralog of the proviral cyclophilin CypA, binds capsid and reduces nuclear import of the HIV core. Structural modeling, evolutionary analyses, and targeted mutagenesis revealed domains and residues required for PPID-mediated HIV restriction, including non-human primate ortholog substitutions that enhance antiviral activity. Together, these data define the functional HIV-host interaction landscape in primary human T cells and uncover new mechanisms modulating infection."],"pubmed_title":["Systematic discovery of pro- and anti-HIV host factors in primary human CD4+ T cells."],"pubmed_authors":["Rathore Ujjwal U, Dugan Eli E, Thornton Hunter H, Kumar Vigneshwari Easwar VE, Dajani Rama R, Burdick Ryan C RC, Young Janet M JM, Steinhart Zachary Z, Lao Reanna R, Delviks-Frankenberry Krista A KA, Choi Wooyoung W, Henriques William S WS, Echeverria Ignacia I, Dann Emma E, Dureja Ishaan I, Pathak Nandini N, Arce Maya M MM, McKetney Justin J, Umhoefer Jennifer M JM, Parulekar Simrun S, Schmidt Ralf R, Polacco Benjamin J BJ, Neidleman Jason J, Montano Mauricio M, Nguyen Vinh Q VQ, Sali Andrej A, Levy Jay A JA, Tenthorey Jeannette L JL, Cheng Yifan Y, Roan Nadia R NR, Swaney Danielle L DL, Kaake Robyn M RM, Dodgson Stacie E SE, Hiatt Joseph J, Pathak Vinay K VK, Malik Harmit S HS, Krogan Nevan J NJ, Marson Alexander A"],"additional_accession":[]},"is_claimable":false,"name":"Systematic Discovery of Pro- and Anti-HIV Host Factors in Primary Human CD4+ T Cells","description":"Despite extensive efforts, significant gaps remain in our understanding of which host factors promote or restrict human immunodeficiency virus (HIV) infection in human CD4+ T cells. Here, we employed orthogonal genome-wide CRISPR activation (CRISPRa) and CRISPR knockout (CRISPRn) screens in primary CD4+ T cells to discover both pro- and anti-HIV host factors systematically. We used secondary pooled screens and targeted individual CRISPR perturbations to validate high-confidence hits and reveal their diverse mechanisms of action. CRISPRa notably revealed potent antiviral factors, including PI16, PPID, SHISA3, and ITM2A – many of which would have been missed by gene knockout experiments alone. When PI16 is overexpressed in CD4+ T cells, it physically interacts with CD4 and numerous HIV host factors involved in viral fusion, and efficiently inhibits HIV entry. We also discovered another novel antiviral factor, PPID (Cyp40). PPID is a paralog of CypA (or PPIA), which is a proviral factor that  regulates nuclear entry kinetics of HIV cores. PPID also binds to HIV capsid, but unlike CypA, it restricts infection by blocking nuclear import of the viral core. Structural modeling, evolutionary analyses, and targeted mutagenesis identified domains and residues that mediate restriction of HIV by PPID, including specific substitutions from non-human primate orthologs that improve HIV restriction. Together, our data systematically define the functional HIV-host interaction landscape in primary human T cells and reveal new mechanisms to modulate infection.","dates":{"publication":"2026-04-27","submission":"2025-12-11"},"accession":"PXD071892","cross_references":{"TAXONOMY":["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: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:418106","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:135666","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:3716","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:1431297","NEWT:9005","NEWT:34871","NEWT:9006","NEWT:33548","NEWT:1883426","NEWT:657321","NEWT:1423687","NEWT:657322","NEWT:10886","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:2499525","NEWT:467144","NEWT:142786","NEWT:28196","NEWT:2853422","NEWT:487","NEWT:1679718","NEWT:387344","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: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:34839","NEWT:476","NEWT:28128","NEWT:470","NEWT:41122","NEWT:1328300","NEWT:748693","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: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:546","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:554","NEWT:451516","NEWT:552","NEWT:1325291","NEWT:42410","NEWT:40686","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: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:13037","NEWT:519","NEWT:941638","NEWT:511","NEWT:7394","NEWT:6063","NEWT:404330","NEWT:1328388","NEWT:13064","NEWT:529120","NEWT:300641","NEWT:39655","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:265619","NEWT:11294","NEWT:754466","NEWT:9925","NCBITaxon:9606","NEWT:1436183","NEWT:1479235","NEWT:4232","NEWT:9922","NEWT:416348","NEWT:105812","NEWT:11298","NEWT:4233","N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