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Goldstein"],"technology_type":["Mass Spectrometry","Bottom-up proteomics"],"software":[""],"submitter_keywords":["Brain","Mouse","Multiomics"],"full_dataset_link":["https://www.ebi.ac.uk/pride/archive/projects/PXD056873"],"tissue":["Brain"],"sample_protocol":["Mouse brains were flash frozen and used for whole brain mass spectrometry analysis. Whole brains were homogenized in 5 mL of protein extraction buffer (100 mM Tris pH 8.5, 8 M Urea, 1 mM CaCl2, 10 mM TCEP, 40 mM Chloroacetamide) using a Polytron® PT 3100 tissue homogenizer at 20,000 r/min for 1 min. Protein extracts were sonicated with a Branson probe sonicator and were then quantified via Bradford assay. 100 μg of protein from each sample was diluted with 100 mM Tris, pH 8.5 buffer to lower the urea concentration to 1 M. Lysate proteins were then digested by the addition of trypsin (Pierce) at a 1:50 ratio (enzyme: protein, w/w) and incubated overnight at 37°C with shaking. Trypsin digestion was terminated with the addition of TFA to below pH 3 and the peptide digests were desalted via reversed-phase C18 columns (Sep-Pak, Waters) with a wash buffer of 0.1% TFA and elution buffer of 60% acetonitrile. The desalted peptides were then quantified with a Quantitative Colorimetric Peptide Assay (Pierce). Each sample comprising 50 μg peptides was TMT-labeled with TMTPro 16plex reagents (ThermoFisher, cat. # A44520) according to the manufacturer’s protocol. Labeled samples were combined and desalted on a C18 column prior to basic reversed-phase fractionation. TMT-labeled peptides were fractionated via basic reversed-phase chromatography on the Agilent 1100 series HPLC instrument equipped with the XBridge Peptide BEH C18 column (130 Å, 3.5 μm, 4.6 mm × 250 mm, Waters Corporation). Prior to loading peptides, the C18 column was washed with 100% methanol and equilibrated with Buffer A (0.1% NH4OH and 2% acetonitrile). Peptides were injected via the autosampler and eluted from the column using a gradient of mobile phase A (2% acetonitrile, 0.1% NH4OH) to mobile phase B (98% acetonitrile, 0.1% NH4OH) over 48 min at a flow rate of 0.4 mL/min. The 96 fractions collected were orthogonally concatenated into 24 pooled fractions."],"repository":["Pride"],"quantification_method":[""],"modification":[""],"data_protocol":["All acquired MS/MS spectra were searched against the complete SwissProt mouse proteome using MaxQuant (Version 2.0.1.0), which integrates the Andromeda search engine. TMT reporter ion quantification was performed using MaxQuant with default settings. Briefly, enzyme specificity was set to trypsin and up to two missed cleavages were allowed. Cysteine carbamidomethylation was specified as fixed modification whereas oxidation of methionine and N-terminal protein acetylation were set as variable modifications. Precursor ions were searched with a maximum mass deviation of 4.5 ppm and fragment ions with a maximum mass deviation of 20 ppm. Peptide and protein identifications were filtered at 1% FDR using the target-decoy database search strategy. Proteins that could not be differentiated based on MS/MS spectra alone were grouped to protein groups (default MaxQuant settings). The MaxQuant output file designated “proteinGroups” was used for data normalization and other statistical analysis using the Omics Notebook analysis pipeline"],"omics_type":["Proteomics"],"labhead":["Camron Bryant"],"instrument_platform":[""],"labhead_affiliation":["Professor and Principal Investigator Laboratory of Addicition Genetics Bouve College of Health Sciences Northeastern University Boston, Massachusetts, USA"],"submission_type":["COMPLETE"],"species":["Mus Musculus (mouse)"],"publication":["39257803 Lynch WB, Miracle SA, Goldstein SI, Beierle JA, Bhandari R, Gerhardt ET, Farnan A, Nguyen BM, Wingfield KK, Kazerani I, Saavedra GA, Averin O, Baskin BM, Ferris MT, Reilly CA, Emili A, Bryant CD. Validation studies and multi-omics analysis of Zhx2 as a candidate quantitative trait gene underlying brain oxycodone metabolite (oxymorphone) levels and behavior. bioRxiv. 2024:2024.08.30.610534 10.1101/2024.08.30.610534","40215834 Lynch WB, Miracle SA, Goldstein SI, Beierle JA, Bhandari R, Gerhardt ET, Farnan A, Nguyen BM, Wingfield KK, Kazerani I, Saavedra GA, Averin O, Baskin BM, Ferris MT, Reilly CA, Emili A, Bryant CD. Validation studies and multiomics analysis of Zhx2 as a candidate quantitative trait gene underlying brain oxycodone metabolite (oxymorphone) levels and behavior. J Pharmacol Exp Ther. 2025 392(5):103557 10.1016/j.jpet.2025.103557"],"submitter_mail":["stangold@bu.edu"],"submitter_affiliation":["Boston University"],"submitter_country":["United States"],"doi":["10.6019/PXD056873"],"pubmed_abstract":["Sensitivity to the subjective reinforcing properties of opioids has a genetic component and can predict addiction liability of opioid compounds. We previously identified Zhx2 as a candidate gene underlying increased brain concentration of the oxycodone (OXY) metabolite oxymorphone (OMOR) in BALB/cJ (J) versus BALB/cByJ (By) females that could increase OXY state-dependent reward. A large structural intronic variant is associated with a robust reduction of Zhx2 expression in J mice, which we hypothesized enhances OMOR levels and OXY addiction-like behaviors. We tested this hypothesis by restoring the Zhx2 loss-of-function in J mice (mouse endogenous retroviral element knockout) and modeling the loss-of-function variant through knocking out the Zhx2 coding exon (exon 3 knockout [E3KO]) in By mice and assessing brain OXY metabolite levels and behavior. Consistent with our hypothesis, Zhx2 E3KO females showed an increase in brain OMOR levels and OXY-induced locomotor activity. However, contrary to our hypothesis, state-dependent expression of OXY conditioned place preference decreased in E3KO females and increased in E3KO males. We also overexpressed Zhx2 in the livers and brains of J mice and observed Zhx2 overexpression in select brain regions that was associated with reduced OXY state-dependent learning. Integrative transcriptomic and proteomic analysis of E3KO mice identified astrocyte function, cell adhesion, extracellular matrix properties, and endothelial cell functions as pathways influencing brain OXY metabolite concentration and behavior. These results support Zhx2 as a quantitative trait gene underlying brain OMOR concentration that is associated with changes in OXY behavior and implicate potential quantitative trait mechanisms that together inform our overall understanding of Zhx2 in brain function. SIGNIFICANCE STATEMENT: This study validated Zhx2 as a gene whose dysfunction increases brain levels of a highly potent and addictive metabolite of oxycodone, oxymorphone, in a female-specific manner. This result has broad implications for understanding the role of oxycodone metabolism and brain oxymorphone levels in the addiction liability of oxycodone (the active ingredient in OxyContin) and highlights the need for the study of sex differences in opioid metabolism as it relates to the addiction liability of opioids and opioid use disorder.","Sensitivity to the subjective reinforcing properties of opioids has a genetic component and can predict addiction liability of opioid compounds. We previously identified <i>Zhx2</i> as a candidate gene underlying increased brain concentration of the oxycodone (<b>OXY</b>) metabolite oxymorphone (<b>OMOR</b>) in BALB/cJ (<b>J</b>) versus BALB/cByJ (<b>By</b>) females that could increase OXY state-dependent reward. A large structural intronic variant is associated with a robust reduction of Zhx2 expression in J mice, which we hypothesized enhances OMOR levels and OXY addiction-like behaviors. We tested this hypothesis by restoring the <i>Zhx2</i> loss-of-function in Js (<b>MVKO</b>) and modeling the loss-of-function variant through knocking out the <i>Zhx2</i> coding exon (<b>E3KO</b>) in Bys and assessing brain OXY metabolite levels and behavior. Consistent with our hypothesis, Zhx2 E3KO females showed an increase in brain OMOR levels and OXY-induced locomotor activity. However, contrary to our hypothesis, state-dependent expression of OXY-CPP was decreased in E3KO females and increased in E3KO males. We also overexpressed Zhx2 in the livers and brains of Js and observed Zhx2 overexpression in select brain regions that was associated with reduced OXY state-dependent learning. Integrative transcriptomic and proteomic analysis of E3KO mice identified astrocyte function, cell adhesion, extracellular matrix properties, and endothelial cell functions as pathways influencing brain OXY metabolite concentration and behavior. These results support <i>Zhx2</i> as a quantitative trait gene underlying brain OMOR concentration that is associated with changes in OXY behavior and implicate potential quantitative trait mechanisms that together inform our overall understanding of <i>Zhx2</i> in brain function."],"pubmed_title":["Validation studies and multiomics analysis of Zhx2 as a candidate quantitative trait gene underlying brain oxycodone metabolite (oxymorphone) levels and behavior.","Validation studies and multi-omics analysis of Zhx2 as a candidate quantitative trait gene underlying brain oxycodone metabolite (oxymorphone) levels and behavior."],"pubmed_authors":["Lynch William B WB, Miracle Sophia A SA, Goldstein Stanley I SI, Beierle Jacob A JA, Bhandari Rhea R, Gerhardt Ethan T ET, Farnan Ava A, Nguyen Binh-Minh BM, Wingfield Kelly K KK, Kazerani Ida I, Saavedra Gabriel A GA, Averin Olga O, Baskin Britahny M BM, Ferris Martin T MT, Reilly Christopher A CA, Emili Andrew A, Bryant Camron D CD"],"additional_accession":[]},"is_claimable":false,"name":"Validation studies and multi-omics analysis of Zhx2 as a candidate quantitative trait gene underlying brain oxycodone metabolite (oxymorphone) levels and behavior","description":"Sensitivity to the subjective reinforcing properties of opioids has a genetic component and can predict addiction liability of opioid compounds. We previously identified Zhx2 as a candidate gene underlying increased brain concentration of the oxycodone (OXY) metabolite oxymorphone (OMOR) in BALB/cJ (J) versus BALB/cByJ (By) females that could increase OXY state-dependent reward. A large structural intronic variant is associated with a robust reduction of Zhx2 expression in J mice, which we hypothesized enhances OMOR levels and OXY addiction-like behaviors. We tested this hypothesis by restoring the Zhx2 loss-of-function in Js (MVKO) and modeling the loss-of-function variant through knocking out the Zhx2 coding exon (E3KO) in Bys and assessing brain OXY metabolite levels and behavior. Consistent with our hypothesis, Zhx2 E3KO females showed an increase in brain OMOR levels and OXY-induced locomotor activity. However, contrary to our hypothesis, state-dependent expression of OXY-CPP was decreased in E3KO females and increased in E3KO males. We also overexpressed Zhx2 in the livers and brains of Js and observed Zhx2 overexpression in select brain regions that was associated with reduced OXY state-dependent learning. Integrative transcriptomic and proteomic analysis of E3KO mice identified astrocyte function, cell adhesion, extracellular matrix properties, and endothelial cell functions as pathways influencing brain OXY metabolite concentration and behavior. These results support Zhx2 as a quantitative trait gene underlying brain OMOR concentration that is associated with changes in OXY behavior and implicate potential quantitative trait mechanisms that together inform our overall understanding of Zhx2 in brain function.","dates":{"publication":"2026-03-30","submission":"2024-10-16"},"accession":"PXD056873","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: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:10029","NEWT:913645","NEWT:641809","NEWT:44250","NEWT:77151","NEWT:317447","NEWT:698761","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: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:1072569","NEWT:1072568","NEWT:161934","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: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:284811","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: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: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:85698","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","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:63367","NEWT:29488","NEWT:215402","NEWT:3726","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: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: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: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","NEWT:1045775","NEWT:510516","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"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