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Dual Checkpoint Inhibition with Ipilimumab plus Nivolumab After Progression on Sequential PD-1/PDL-1 Inhibitors Pembrolizumab and Atezolizumab in a Patient with Lynch Syndrome, Metastatic Colon, and Localized Urothelial Cancer.

Winer A, Ghatalia P, Bubes N, Anari F, Varshavsky A, Kasireddy V, Liu Y, El-Deiry WS.

Oncologist. 2019 Aug 23. pii: theoncologist.2018-0686. doi: 10.1634/theoncologist.2018-0686. [Epub ahead of print]


Gid10 as an alternative N-recognin of the Pro/N-degron pathway.

Melnykov A, Chen SJ, Varshavsky A.

Proc Natl Acad Sci U S A. 2019 Aug 6;116(32):15914-15923. doi: 10.1073/pnas.1908304116. Epub 2019 Jul 23.


On the cause of sleep: Protein fragments, the concept of sentinels, and links to epilepsy.

Varshavsky A.

Proc Natl Acad Sci U S A. 2019 May 28;116(22):10773-10782. doi: 10.1073/pnas.1904709116. Epub 2019 May 13.


Clinical Utilization Pattern of Liquid Biopsies (LB) to Detect Actionable Driver Mutations, Guide Treatment Decisions and Monitor Disease Burden During Treatment of 33 Metastatic Colorectal Cancer (mCRC) Patients (pts) at a Fox Chase Cancer Center GI Oncology Subspecialty Clinic.

Ghatalia P, Smith CH, Winer A, Gou J, Kiedrowski LA, Slifker M, Saltzberg PD, Bubes N, Anari FM, Kasireddy V, Varshavsky A, Liu Y, Ross EA, El-Deiry WS.

Front Oncol. 2019 Jan 17;8:652. doi: 10.3389/fonc.2018.00652. eCollection 2018.


N-degron and C-degron pathways of protein degradation.

Varshavsky A.

Proc Natl Acad Sci U S A. 2019 Jan 8;116(2):358-366. doi: 10.1073/pnas.1816596116. Review.


Formyl-methionine as an N-degron of a eukaryotic N-end rule pathway.

Kim JM, Seok OH, Ju S, Heo JE, Yeom J, Kim DS, Yoo JY, Varshavsky A, Lee C, Hwang CS.

Science. 2018 Nov 30;362(6418). pii: eaat0174. doi: 10.1126/science.aat0174. Epub 2018 Nov 8.


Understanding the Pro/N-end rule pathway.

Dougan DA, Varshavsky A.

Nat Chem Biol. 2018 May;14(5):415-416. doi: 10.1038/s41589-018-0045-0. No abstract available.


A reference-based protein degradation assay without global translation inhibitors.

Oh JH, Chen SJ, Varshavsky A.

J Biol Chem. 2017 Dec 29;292(52):21457-21465. doi: 10.1074/jbc.M117.814236. Epub 2017 Nov 9.


The Ubiquitin System, Autophagy, and Regulated Protein Degradation.

Varshavsky A.

Annu Rev Biochem. 2017 Jun 20;86:123-128. doi: 10.1146/annurev-biochem-061516-044859. Review.


Control of Hsp90 chaperone and its clients by N-terminal acetylation and the N-end rule pathway.

Oh JH, Hyun JY, Varshavsky A.

Proc Natl Acad Sci U S A. 2017 May 30;114(22):E4370-E4379. doi: 10.1073/pnas.1705898114. Epub 2017 May 17.


An N-end rule pathway that recognizes proline and destroys gluconeogenic enzymes.

Chen SJ, Wu X, Wadas B, Oh JH, Varshavsky A.

Science. 2017 Jan 27;355(6323). pii: eaal3655. doi: 10.1126/science.aal3655.


Analyzing N-terminal Arginylation through the Use of Peptide Arrays and Degradation Assays.

Wadas B, Piatkov KI, Brower CS, Varshavsky A.

J Biol Chem. 2016 Sep 30;291(40):20976-20992. Epub 2016 Aug 10.


Degradation of Serotonin N-Acetyltransferase, a Circadian Regulator, by the N-end Rule Pathway.

Wadas B, Borjigin J, Huang Z, Oh JH, Hwang CS, Varshavsky A.

J Biol Chem. 2016 Aug 12;291(33):17178-96. doi: 10.1074/jbc.M116.734640. Epub 2016 Jun 23.


Formyl-methionine as a degradation signal at the N-termini of bacterial proteins.

Piatkov KI, Vu TT, Hwang CS, Varshavsky A.

Microb Cell. 2015;2(10):376-393.


Degradation of the Separase-cleaved Rec8, a Meiotic Cohesin Subunit, by the N-end Rule Pathway.

Liu YJ, Liu C, Chang Z, Wadas B, Brower CS, Song ZH, Xu ZL, Shang YL, Liu WX, Wang LN, Dong W, Varshavsky A, Hu RG, Li W.

J Biol Chem. 2016 Apr 1;291(14):7426-38. doi: 10.1074/jbc.M116.714964. Epub 2016 Feb 8.


Control of mammalian G protein signaling by N-terminal acetylation and the N-end rule pathway.

Park SE, Kim JM, Seok OH, Cho H, Wadas B, Kim SY, Varshavsky A, Hwang CS.

Science. 2015 Mar 13;347(6227):1249-1252. doi: 10.1126/science.aaa3844.


Liat1, an arginyltransferase-binding protein whose evolution among primates involved changes in the numbers of its 10-residue repeats.

Brower CS, Rosen CE, Jones RH, Wadas BC, Piatkov KI, Varshavsky A.

Proc Natl Acad Sci U S A. 2014 Nov 18;111(46):E4936-45. doi: 10.1073/pnas.1419587111. Epub 2014 Nov 4.


Discovery of the biology of the ubiquitin system.

Varshavsky A.

JAMA. 2014 May 21;311(19):1969-70. doi: 10.1001/jama.2014.5549. No abstract available.


Calpain-generated natural protein fragments as short-lived substrates of the N-end rule pathway.

Piatkov KI, Oh JH, Liu Y, Varshavsky A.

Proc Natl Acad Sci U S A. 2014 Mar 4;111(9):E817-26. doi: 10.1073/pnas.1401639111. Epub 2014 Feb 18.


The N-terminal methionine of cellular proteins as a degradation signal.

Kim HK, Kim RR, Oh JH, Cho H, Varshavsky A, Hwang CS.

Cell. 2014 Jan 16;156(1-2):158-69. doi: 10.1016/j.cell.2013.11.031. Epub 2013 Dec 19.


Ubiquitin reference technique and its use in ubiquitin-lacking prokaryotes.

Piatkov K, Graciet E, Varshavsky A.

PLoS One. 2013 Jun 25;8(6):e67952. doi: 10.1371/journal.pone.0067952. Print 2013.


Control of protein quality and stoichiometries by N-terminal acetylation and the N-end rule pathway.

Shemorry A, Hwang CS, Varshavsky A.

Mol Cell. 2013 May 23;50(4):540-51. doi: 10.1016/j.molcel.2013.03.018. Epub 2013 Apr 18.


Neurodegeneration-associated protein fragments as short-lived substrates of the N-end rule pathway.

Brower CS, Piatkov KI, Varshavsky A.

Mol Cell. 2013 Apr 25;50(2):161-71. doi: 10.1016/j.molcel.2013.02.009. Epub 2013 Mar 14.


The auto-generated fragment of the Usp1 deubiquitylase is a physiological substrate of the N-end rule pathway.

Piatkov KI, Colnaghi L, Békés M, Varshavsky A, Huang TT.

Mol Cell. 2012 Dec 28;48(6):926-33. doi: 10.1016/j.molcel.2012.10.012. Epub 2012 Nov 15.


The N-end rule pathway counteracts cell death by destroying proapoptotic protein fragments.

Piatkov KI, Brower CS, Varshavsky A.

Proc Natl Acad Sci U S A. 2012 Jul 3;109(27):E1839-47. doi: 10.1073/pnas.1207786109. Epub 2012 Jun 5.


The ubiquitin system, an immense realm.

Varshavsky A.

Annu Rev Biochem. 2012;81:167-76. doi: 10.1146/annurev-biochem-051910-094049. Review. No abstract available.


Three decades of studies to understand the functions of the ubiquitin family.

Varshavsky A.

Methods Mol Biol. 2012;832:1-11. doi: 10.1007/978-1-61779-474-2_1.


Ubiquitin ligases of the N-end rule pathway: assessment of mutations in UBR1 that cause the Johanson-Blizzard syndrome.

Hwang CS, Sukalo M, Batygin O, Addor MC, Brunner H, Aytes AP, Mayerle J, Song HK, Varshavsky A, Zenker M.

PLoS One. 2011;6(9):e24925. doi: 10.1371/journal.pone.0024925. Epub 2011 Sep 13.


Plexin-A4 promotes tumor progression and tumor angiogenesis by enhancement of VEGF and bFGF signaling.

Kigel B, Rabinowicz N, Varshavsky A, Kessler O, Neufeld G.

Blood. 2011 Oct 13;118(15):4285-96. doi: 10.1182/blood-2011-03-341388. Epub 2011 Aug 10.


The N-end rule pathway and regulation by proteolysis.

Varshavsky A.

Protein Sci. 2011 Aug;20(8):1298-345. doi: 10.1002/pro.666. Review.


The N-end rule pathway is mediated by a complex of the RING-type Ubr1 and HECT-type Ufd4 ubiquitin ligases.

Hwang CS, Shemorry A, Auerbach D, Varshavsky A.

Nat Cell Biol. 2010 Dec;12(12):1177-85. doi: 10.1038/ncb2121. Epub 2010 Nov 14.


Mouse Dfa is a repressor of TATA-box promoters and interacts with the Abt1 activator of basal transcription.

Brower CS, Veiga L, Jones RH, Varshavsky A.

J Biol Chem. 2010 May 28;285(22):17218-34. doi: 10.1074/jbc.M110.118638. Epub 2010 Mar 31.


N-terminal acetylation of cellular proteins creates specific degradation signals.

Hwang CS, Shemorry A, Varshavsky A.

Science. 2010 Feb 19;327(5968):973-7. doi: 10.1126/science.1183147. Epub 2010 Jan 28.


The N-end rule pathway controls multiple functions during Arabidopsis shoot and leaf development.

Graciet E, Walter F, Ó'Maoiléidigh DS, Pollmann S, Meyerowitz EM, Varshavsky A, Wellmer F.

Proc Natl Acad Sci U S A. 2009 Aug 11;106(32):13618-23. doi: 10.1073/pnas.0906404106. Epub 2009 Jul 20.


Glutamine-specific N-terminal amidase, a component of the N-end rule pathway.

Wang H, Piatkov KI, Brower CS, Varshavsky A.

Mol Cell. 2009 Jun 26;34(6):686-95. doi: 10.1016/j.molcel.2009.04.032.


Two proteolytic pathways regulate DNA repair by cotargeting the Mgt1 alkylguanine transferase.

Hwang CS, Shemorry A, Varshavsky A.

Proc Natl Acad Sci U S A. 2009 Feb 17;106(7):2142-7. doi: 10.1073/pnas.0812316106. Epub 2009 Jan 21.


The N-end rule at atomic resolution.

Varshavsky A.

Nat Struct Mol Biol. 2008 Dec;15(12):1238-40. doi: 10.1038/nsmb1208-1238. No abstract available.


Regulation of peptide import through phosphorylation of Ubr1, the ubiquitin ligase of the N-end rule pathway.

Hwang CS, Varshavsky A.

Proc Natl Acad Sci U S A. 2008 Dec 9;105(49):19188-93. doi: 10.1073/pnas.0808891105. Epub 2008 Nov 25.


Successful inhibition of tumor development by specific class-3 semaphorins is associated with expression of appropriate semaphorin receptors by tumor cells.

Kigel B, Varshavsky A, Kessler O, Neufeld G.

PLoS One. 2008 Sep 26;3(9):e3287. doi: 10.1371/journal.pone.0003287.


Semaphorin-3B is an angiogenesis inhibitor that is inactivated by furin-like pro-protein convertases.

Varshavsky A, Kessler O, Abramovitch S, Kigel B, Zaffryar S, Akiri G, Neufeld G.

Cancer Res. 2008 Sep 1;68(17):6922-31. doi: 10.1158/0008-5472.CAN-07-5408.


Amino acids induce peptide uptake via accelerated degradation of CUP9, the transcriptional repressor of the PTR2 peptide transporter.

Xia Z, Turner GC, Hwang CS, Byrd C, Varshavsky A.

J Biol Chem. 2008 Oct 24;283(43):28958-68. doi: 10.1074/jbc.M803980200. Epub 2008 Aug 15.


Discovery of cellular regulation by protein degradation.

Varshavsky A.

J Biol Chem. 2008 Dec 12;283(50):34469-89. doi: 10.1074/jbc.X800009200. Epub 2008 Aug 15. No abstract available.


Substrate-binding sites of UBR1, the ubiquitin ligase of the N-end rule pathway.

Xia Z, Webster A, Du F, Piatkov K, Ghislain M, Varshavsky A.

J Biol Chem. 2008 Aug 29;283(35):24011-28. doi: 10.1074/jbc.M802583200. Epub 2008 Jun 19.


Enzymatic N-terminal addition of noncanonical amino acids to peptides and proteins.

Connor RE, Piatkov K, Varshavsky A, Tirrell DA.

Chembiochem. 2008 Feb 15;9(3):366-9. doi: 10.1002/cbic.200700605. No abstract available.


The N-end rule pathway is a sensor of heme.

Hu RG, Wang H, Xia Z, Varshavsky A.

Proc Natl Acad Sci U S A. 2008 Jan 8;105(1):76-81. Epub 2007 Dec 27.


Targeting the absence: homozygous DNA deletions as immutable signposts for cancer therapy.

Varshavsky A.

Proc Natl Acad Sci U S A. 2007 Sep 18;104(38):14935-40. Epub 2007 Sep 10.


Listeriolysin O secreted by Listeria monocytogenes into the host cell cytosol is degraded by the N-end rule pathway.

Schnupf P, Zhou J, Varshavsky A, Portnoy DA.

Infect Immun. 2007 Nov;75(11):5135-47. Epub 2007 Aug 6.


Semaphorin signaling in vascular and tumor biology.

Neufeld G, Lange T, Varshavsky A, Kessler O.

Adv Exp Med Biol. 2007;600:118-31. Review.


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