Scholars@Duke

• Background
• 

  Education, Training, & Certifications

  • Hem/Onc Clinical Fellowship, Pediatric, Johns Hopkins University 1984 - 1985
  • Pediatric Internship & Residency, Pediatrics, Johns Hopkins University 1981 - 1984
  • M.D., Johns Hopkins University 1981
• 

  Duke Appointment History

  • Instructor, Temporary in the Department of Pediatrics, Pediatrics, Clinical Science Departments 2007
• Recognition
• 

  In the News

  • NOV 19, 2013

    Meet the Duke-NUS Faculty: David Virshup
• Research
• 

  External Relationships

  • Experimental Therapeutics Centre Singapore, renamed the Experimental Drug Development Centre
  • Guidepoint Clinical
  • VVB
• Publications & Artistic Works
• 

  Selected Publications

  • Academic Articles

    • Madan, Babita, David M. Virshup, W David Nes, and David J. Leaver. “Unearthing the Janus-face cholesterogenesis pathways in cancer.” Biochem Pharmacol, May 16, 2021, 114611. https://doi.org/10.1016/j.bcp.2021.114611.
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    • Zacarías-Fluck, Mariano F., Toni Jauset, Sandra Martínez-Martín, Jastrinjan Kaur, Sílvia Casacuberta-Serra, Daniel Massó-Vallés, Erika Serrano Del Pozo, et al. “The Wnt signaling receptor Fzd9 is essential for Myc-driven tumorigenesis in pancreatic islets.” Life Sci Alliance 4, no. 5 (May 2021). https://doi.org/10.26508/lsa.201900490.
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    • Kaur, Amanpreet, Jun Yi Stanley Lim, Sugunavathi Sepramaniam, Siddhi Patnaik, Nathan Harmston, May Ann Lee, Enrico Petretto, David M. Virshup, and Babita Madan. “WNT inhibition creates a BRCA-like state in Wnt-addicted cancer.” Embo Mol Med 13, no. 4 (April 9, 2021): e13349. https://doi.org/10.15252/emmm.202013349.
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    • Brunt, Lucy, Gediminas Greicius, Sally Rogers, Benjamin D. Evans, David M. Virshup, Kyle C. A. Wedgwood, and Steffen Scholpp. “Vangl2 promotes the formation of long cytonemes to enable distant Wnt/β-catenin signaling.” Nat Commun 12, no. 1 (April 6, 2021): 2058. https://doi.org/10.1038/s41467-021-22393-9.
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    • Stephenson, Ewan M., Laura E. J. Usselmann, Vinay Tergaonkar, David M. Virshup, and Robert Dallmann. “Cancer clocks in tumourigenesis: the p53 pathway and beyond.” Endocr Relat Cancer 28, no. 4 (April 2021): R95–110. https://doi.org/10.1530/ERC-20-0475.
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    • Narasimamurthy, Rajesh, and David M. Virshup. “The phosphorylation switch that regulates ticking of the circadian clock.” Mol Cell 81, no. 6 (March 18, 2021): 1133–46. https://doi.org/10.1016/j.molcel.2021.01.006.
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    • Chua, Kenon, David M. Virshup, Eugene G. Odono, Kenneth Tou En Chang, Nicholas Jin Hong Tan, Susan Swee-Shan Hue, Arthur Yi Loong Sim, and Victor Kwan Min Lee. “YJ5 as an immunohistochemical marker of osteogenic lineage.” Pathology 53, no. 2 (February 2021): 229–38. https://doi.org/10.1016/j.pathol.2020.07.017.
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    • Harmston, Nathan, Jun Yi Stanley Lim, Oriol Arqués, Héctor G. Palmer, Enrico Petretto, David M. Virshup, and Babita Madan. “Widespread Repression of Gene Expression in Cancer by a Wnt/β-Catenin/MAPK Pathway.” Cancer Res 81, no. 2 (January 15, 2021): 464–75. https://doi.org/10.1158/0008-5472.CAN-20-2129.
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    • Nygaard, Rie, Jia Yu, Jonathan Kim, Daniel R. Ross, Giacomo Parisi, Oliver B. Clarke, David M. Virshup, and Filippo Mancia. “Structural Basis of WLS/Evi-Mediated Wnt Transport and Secretion.” Cell 184, no. 1 (January 7, 2021): 194-206.e14. https://doi.org/10.1016/j.cell.2020.11.038.
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    • Yu, Jia, Permeen A Mohamed Yusoff, Daniëlle T. J. Woutersen, Pamela Goh, Nathan Harmston, Ron Smits, David M. Epstein, David M. Virshup, and Babita Madan. “The Functional Landscape of Patient-Derived RNF43 Mutations Predicts Sensitivity to Wnt Inhibition.” Cancer Res 80, no. 24 (December 15, 2020): 5619–32. https://doi.org/10.1158/0008-5472.CAN-20-0957.
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    • Liu, Shiyang, Nathan Harmston, Trudy Lee Glaser, Yunka Wong, Zheng Zhong, Babita Madan, David M. Virshup, and Enrico Petretto. “Wnt-regulated lncRNA discovery enhanced by in vivo identification and CRISPRi functional validation.” Genome Med 12, no. 1 (October 22, 2020): 89. https://doi.org/10.1186/s13073-020-00788-5.
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    • Zhong, Zheng, Jia Yu, David M. Virshup, and Babita Madan. “Wnts and the hallmarks of cancer.” Cancer Metastasis Rev 39, no. 3 (September 2020): 625–45. https://doi.org/10.1007/s10555-020-09887-6.
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    • Masuda, Shusaku, Rajesh Narasimamurthy, Hikari Yoshitane, Jae Kyoung Kim, Yoshitaka Fukada, and David M. Virshup. “Mutation of a PER2 phosphodegron perturbs the circadian phosphoswitch.” Proc Natl Acad Sci U S A 117, no. 20 (May 19, 2020): 10888–96. https://doi.org/10.1073/pnas.2000266117.
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    • Yu, Jia, Permeen Akhtar Bt Mohamed Yuso, Pamela Goh, Nathan Harmston, David Epstein, David Virshup, and Babita Madan. “The functional landscape of patient derived RNF43 mutations predicts Wnt inhibitor sensitivity,” March 25, 2020. https://doi.org/10.1101/2020.03.25.006999.
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    • Liu, Shiyang, Nathan Harmston, Trudy Lee Glaser, Yunka Wong, Zheng Zhong, Babita Madan, David Virshup, and Enrico Petretto. “Wnt-regulated lncRNA discovery enhanced by in vivo identification and CRISPRi functional validation,” February 10, 2020. https://doi.org/10.1101/2020.02.09.941005.
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    • Zhong, Zheng, and David M. Virshup. “Wnt Signaling and Drug Resistance in Cancer.” Mol Pharmacol 97, no. 2 (February 2020): 72–89. https://doi.org/10.1124/mol.119.117978.
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    • Lee, Kyunghee, Mayumi Kitagawa, Pei Ju Liao, David M. Virshup, and Sang Hyun Lee. “A Ras-LSD1 axis activates PI3K signaling through PIK3IP1 suppression.” Oncogenesis 9, no. 1 (January 2, 2020): 2. https://doi.org/10.1038/s41389-019-0185-4.
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    • Masuda, Shusaku, Rajesh Narasimamurthy, Hikari Yoshitane, Jae Kyoung Kim, Yoshitaka Fukada, and David Virshup. “Mutation of a PER2 phosphodegron perturbs the circadian phosphoswitch,” December 17, 2019. https://doi.org/10.1101/2019.12.16.876615.
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    • Idris, Muhammad, Nathan Harmston, Enrico Petretto, Babita Madan, and David M. Virshup. “Broad regulation of gene isoform expression by Wnt signaling in cancer.” Rna 25, no. 12 (December 2019): 1696–1713. https://doi.org/10.1261/rna.071506.119.
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    • Lu, Xiaohan, Nathan P. Rudemiller, Jiafa Ren, Yi Wen, Bo Yang, Robert Griffiths, Jamie R. Privratsky, Babita Madan, David M. Virshup, and Steven D. Crowley. “Opposing actions of renal tubular- and myeloid-derived porcupine in obstruction-induced kidney fibrosis.” Kidney Int 96, no. 6 (December 2019): 1308–19. https://doi.org/10.1016/j.kint.2019.06.020.
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    • Zhong, Zheng, Sugunavathi Sepramaniam, Xin Hui Chew, Kris Wood, May Ann Lee, Babita Madan, and David M. Virshup. “PORCN inhibition synergizes with PI3K/mTOR inhibition in Wnt-addicted cancers.” Oncogene 38, no. 40 (October 2019): 6662–77. https://doi.org/10.1038/s41388-019-0908-1.
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    • Philpott, Jonathan, Rajesh Narasimamurthy, Clarisse Ricci, Alfred Freeberg, Sabrina Hunt, Lauren Yee, Rebecca Pelofsky, Sarvind Tripathi, David Virshup, and Carrie Partch. “Casein Kinase 1 dynamics underlie the PER2 circadian phosphoswitch,” August 14, 2019. https://doi.org/10.1101/734624.
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    • Fadhlullah, Siti Farah Bte, Nurashikin Bte Abdul Halim, Jacqueline Y. T. Yeo, Rachel L. Y. Ho, Phoebe Um, Beng Ti Ang, Carol Tang, Wai H. Ng, David M. Virshup, and Ivy A. W. Ho. “Pathogenic mutations in neurofibromin identifies a leucine-rich domain regulating glioma cell invasiveness.” Oncogene 38, no. 27 (July 2019): 5367–80. https://doi.org/10.1038/s41388-019-0809-3.
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    • Greicius, Gediminas, and David M. Virshup. “Stromal control of intestinal development and the stem cell niche.” Differentiation 108 (July 2019): 8–16. https://doi.org/10.1016/j.diff.2019.01.001.
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    • Moti, Naushad, Jia Yu, Gaelle Boncompain, Franck Perez, and David M. Virshup. “Wnt traffic from endoplasmic reticulum to filopodia.” Plos One 14, no. 2 (2019): e0212711. https://doi.org/10.1371/journal.pone.0212711.
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    • Fu, Nai Yang, Bhupinder Pal, Yunshun Chen, Felicity C. Jackling, Michael Milevskiy, François Vaillant, Bianca D. Capaldo, et al. “Foxp1 Is Indispensable for Ductal Morphogenesis and Controls the Exit of Mammary Stem Cells from Quiescence.” Dev Cell 47, no. 5 (December 3, 2018): 629-644.e8. https://doi.org/10.1016/j.devcel.2018.10.001.
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    • Madan, Babita, Nathan Harmston, Gahyathiri Nallan, Alex Montoya, Peter Faull, Enrico Petretto, and David M. Virshup. “Temporal dynamics of Wnt-dependent transcriptome reveal an oncogenic Wnt/MYC/ribosome axis.” J Clin Invest 128, no. 12 (December 3, 2018): 5620–33. https://doi.org/10.1172/JCI122383.
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    • Chee, Yuemin Celina, Jens Pahnke, Ralph Bunte, Vikrant A. Adsool, Babita Madan, and David M. Virshup. “Intrinsic Xenobiotic Resistance of the Intestinal Stem Cell Niche.” Dev Cell 46, no. 6 (September 24, 2018): 681-695.e5. https://doi.org/10.1016/j.devcel.2018.07.023.
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    • Kabiri, Zahra, Gediminas Greicius, Hamed Zaribafzadeh, Amanda Hemmerich, Christopher M. Counter, and David M. Virshup. “Wnt signaling suppresses MAPK-driven proliferation of intestinal stem cells.” J Clin Invest 128, no. 9 (August 31, 2018): 3806–12. https://doi.org/10.1172/JCI99325.
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    • Mattes, Benjamin, Yonglong Dang, Gediminas Greicius, Lilian Tamara Kaufmann, Benedikt Prunsche, Jakob Rosenbauer, Johannes Stegmaier, et al. “Wnt/PCP controls spreading of Wnt/β-catenin signals by cytonemes in vertebrates.” Elife 7 (July 31, 2018). https://doi.org/10.7554/eLife.36953.
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    • “Correction for Fustin et al., Two Ck1δ transcripts regulated by m6A methylation code for two antagonistic kinases in the control of the circadian clock.” Proc Natl Acad Sci U S A 115, no. 27 (July 3, 2018): E6386. https://doi.org/10.1073/pnas.1809838115.
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    • Fustin, Jean-Michel, Rika Kojima, Kakeru Itoh, Hsin-Yi Chang, Shiqi Ye, Bowen Zhuang, Asami Oji, et al. “Two Ck1δ transcripts regulated by m6A methylation code for two antagonistic kinases in the control of the circadian clock.” Proc Natl Acad Sci U S A 115, no. 23 (June 5, 2018): 5980–85. https://doi.org/10.1073/pnas.1721371115.
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    • Narasimamurthy, Rajesh, Sabrina R. Hunt, Yining Lu, Jean-Michel Fustin, Hitoshi Okamura, Carrie L. Partch, Daniel B. Forger, Jae Kyoung Kim, and David M. Virshup. “CK1δ/ε protein kinase primes the PER2 circadian phosphoswitch.” Proc Natl Acad Sci U S A 115, no. 23 (June 5, 2018): 5986–91. https://doi.org/10.1073/pnas.1721076115.
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    • Madan, Babita, Nathan Harmston, Gahyathiri Nallan, Alex Montoya, Peter Faull, Enrico Petretto, and David Virshup. “Oncogenic Wnt/STOP signaling regulates ribosome biogenesis in vivo,” May 20, 2018. https://doi.org/10.1101/326819.
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    • Moti, Naushad, Jia Yu, Gaelle Boncompain, Franck Perez, and David Virshup. “Visualizing Wnt secretion from Endoplasmic Reticulum to Filopodia,” April 14, 2018. https://doi.org/10.1101/271684.
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    • Greicius, Gediminas, Zahra Kabiri, Kristmundur Sigmundsson, Chao Liang, Ralph Bunte, Manvendra K. Singh, and David M. Virshup. “PDGFRα + pericryptal stromal cells are the critical source of Wnts and RSPO3 for murine intestinal stem cells in vivo.” Proc Natl Acad Sci U S A 115, no. 14 (April 3, 2018): E3173–81. https://doi.org/10.1073/pnas.1713510115.
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    • Zhang, F., D. M. Virshup, and J. K. Cheong. “Oncogenic RAS-induced CK1α drives nuclear FOXO proteolysis.” Oncogene 37, no. 3 (January 18, 2018): 363–76. https://doi.org/10.1038/onc.2017.334.
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    • Lee, Ha Yin, Yoko Itahana, Stefan Schuechner, Masahiro Fukuda, H Shawn Je, Egon Ogris, David M. Virshup, and Koji Itahana. “Ca2+-dependent demethylation of phosphatase PP2Ac promotes glucose deprivation-induced cell death independently of inhibiting glycolysis.” Sci Signal 11, no. 512 (January 9, 2018). https://doi.org/10.1126/scisignal.aam7893.
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    • Madan, Babita, Mitchell J. McDonald, Gabrielle E. Foxa, Cassandra R. Diegel, Bart O. Williams, and David M. Virshup. “Bone loss from Wnt inhibition mitigated by concurrent alendronate therapy.” Bone Res 6 (2018): 17. https://doi.org/10.1038/s41413-018-0017-8.
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    • Ho, Soo Yei, Jenefer Alam, Duraiswamy Athisayamani Jeyaraj, Weiling Wang, Grace Ruiting Lin, Shi Hua Ang, Eldwin Sum Wai Tan, et al. “Scaffold Hopping and Optimization of Maleimide Based Porcupine Inhibitors.” J Med Chem 60, no. 15 (August 10, 2017): 6678–92. https://doi.org/10.1021/acs.jmedchem.7b00662.
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    • Narasimamurthy, Rajesh, and David M. Virshup. “A Flick of the Tail Keeps the Circadian Clock in Line.” Mol Cell 66, no. 4 (May 18, 2017): 437–38. https://doi.org/10.1016/j.molcel.2017.05.008.
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    • Alok, Anshula, Zhengdeng Lei, N Suhas Jagannathan, Simran Kaur, Nathan Harmston, Steven G. Rozen, Lisa Tucker-Kellogg, and David M. Virshup. “Wnt proteins synergize to activate β-catenin signaling.” J Cell Sci 130, no. 9 (May 1, 2017): 1532–44. https://doi.org/10.1242/jcs.198093.
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    • Eng, Gracie Wee Ling, Gracie Wee Ling Edison, and David M. Virshup. “Site-specific phosphorylation of casein kinase 1 δ (CK1δ) regulates its activity towards the circadian regulator PER2.” Plos One 12, no. 5 (2017): e0177834. https://doi.org/10.1371/journal.pone.0177834.
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    • Narasimamurthy, Rajesh, and David M. Virshup. “Molecular Mechanisms Regulating Temperature Compensation of the Circadian Clock.” Front Neurol 8 (2017): 161. https://doi.org/10.3389/fneur.2017.00161.
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    • Cheong, Jit Kong, and David M. Virshup. “CK1δ: a pharmacologically tractable Achilles' heel of Wnt-driven cancers?” Ann Transl Med 4, no. 21 (November 2016): 433. https://doi.org/10.21037/atm.2016.11.07.
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    • Liu, Chang Ching, Dong Liang Ma, Ting-Dong Yan, XiuBo Fan, Zhiyong Poon, Lai-Fong Poon, Su-Ann Goh, et al. “Distinct Responses of Stem Cells to Telomere Uncapping-A Potential Strategy to Improve the Safety of Cell Therapy.” Stem Cells 34, no. 10 (October 2016): 2471–84. https://doi.org/10.1002/stem.2431.
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    • Somarelli, J. A., D. Schaeffer, M. S. Marengo, T. Bepler, D. Rouse, K. E. Ware, A. J. Hish, et al. “Distinct routes to metastasis: plasticity-dependent and plasticity-independent pathways.” Oncogene 35, no. 33 (August 18, 2016): 4302–11. https://doi.org/10.1038/onc.2015.497.
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    • Wang, Jiao, Zhizhi Wang, Tingting Yu, Huan Yang, David M. Virshup, Geert J. P. L. Kops, Sang Hyun Lee, et al. “Crystal structure of a PP2A B56-BubR1 complex and its implications for PP2A substrate recruitment and localization.” Protein Cell 7, no. 7 (July 2016): 516–26. https://doi.org/10.1007/s13238-016-0283-4.
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    • Madan, Babita, Matthew P. Walker, Robert Young, Laura Quick, Kelly A. Orgel, Meagan Ryan, Priti Gupta, et al. “USP6 oncogene promotes Wnt signaling by deubiquitylating Frizzleds.” Proc Natl Acad Sci U S A 113, no. 21 (May 24, 2016): E2945–54. https://doi.org/10.1073/pnas.1605691113.
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    • Cheong, Jit Kong, and David M. Virshup. “Keeping autophagy in cheCK1.” Mol Cell Oncol 3, no. 3 (May 2016): e1045117. https://doi.org/10.1080/23723556.2015.1045117.
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    • Madan, Babita, Mehul B. Patel, Jiandong Zhang, Ralph M. Bunte, Nathan P. Rudemiller, Robert Griffiths, David M. Virshup, and Steven D. Crowley. “Experimental inhibition of porcupine-mediated Wnt O-acylation attenuates kidney fibrosis.” Kidney Int 89, no. 5 (May 2016): 1062–74. https://doi.org/10.1016/j.kint.2016.01.017.
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    • Madan, B., Z. Ke, N. Harmston, S. Y. Ho, A. O. Frois, J. Alam, D. A. Jeyaraj, et al. “Wnt addiction of genetically defined cancers reversed by PORCN inhibition.” Oncogene 35, no. 17 (April 28, 2016): 2197–2207. https://doi.org/10.1038/onc.2015.280.
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    • Lo, Kinyui Alice, Pei Yi Ng, Zahra Kabiri, David Virshup, and Lei Sun. “Wnt inhibition enhances browning of mouse primary white adipocytes.” Adipocyte 5, no. 2 (April 2016): 224–31. https://doi.org/10.1080/21623945.2016.1148834.
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    • Madan, Babita, Zhiyuan Ke, Zheng Deng Lei, Frois Ashley Oliver, Masanobu Oshima, May Ann Lee, Steve Rozen, and David M. Virshup. “NOTUM is a potential pharmacodynamic biomarker of Wnt pathway inhibition.” Oncotarget 7, no. 11 (March 15, 2016): 12386–92. https://doi.org/10.18632/oncotarget.7157.
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    • Xiang, Wei, Jit Kong Cheong, Shi Hui Ang, Bryan Teo, Peng Xu, Kartini Asari, Wen Tian Sun, et al. “Pyrvinium selectively targets blast phase-chronic myeloid leukemia through inhibition of mitochondrial respiration.” Oncotarget 6, no. 32 (October 20, 2015): 33769–80. https://doi.org/10.18632/oncotarget.5615.
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    • Zhou, Min, Jae Kyoung Kim, Gracie Wee Ling Eng, Daniel B. Forger, and David M. Virshup. “A Period2 Phosphoswitch Regulates and Temperature Compensates Circadian Period.” Mol Cell 60, no. 1 (October 1, 2015): 77–88. https://doi.org/10.1016/j.molcel.2015.08.022.
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    • Kabiri, Zahra, Akihiko Numata, Akira Kawasaki, Akira Edison, Daniel G. Tenen, and David M. Virshup. “Wnts are dispensable for differentiation and self-renewal of adult murine hematopoietic stem cells.” Blood 126, no. 9 (August 27, 2015): 1086–94. https://doi.org/10.1182/blood-2014-09-598540.
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    • Duraiswamy, Athisayamani Jeyaraj, May Ann Lee, Babita Madan, Shi Hua Ang, Eldwin Sum Wai Tan, Wei Wen Vivien Cheong, Zhiyuan Ke, et al. “Discovery and Optimization of a Porcupine Inhibitor.” J Med Chem 58, no. 15 (August 13, 2015): 5889–99. https://doi.org/10.1021/acs.jmedchem.5b00507.
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    • Madan, Babita, and David M. Virshup. “Targeting Wnts at the source--new mechanisms, new biomarkers, new drugs.” Mol Cancer Ther 14, no. 5 (May 2015): 1087–94. https://doi.org/10.1158/1535-7163.MCT-14-1038.
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    • Ribi, Sebastian, Daniel Baumhoer, Kristy Lee, Kristy Edison, Audrey S. M. Teo, Babita Madan, Kang Zhang, et al. “TP53 intron 1 hotspot rearrangements are specific to sporadic osteosarcoma and can cause Li-Fraumeni syndrome.” Oncotarget 6, no. 10 (April 10, 2015): 7727–40. https://doi.org/10.18632/oncotarget.3115.
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    • Cheong, Jit Kong, Fuquan Zhang, Pei Jou Chua, Boon Huat Bay, Andrew Thorburn, and David M. Virshup. “Casein kinase 1α-dependent feedback loop controls autophagy in RAS-driven cancers.” J Clin Invest 125, no. 4 (April 2015): 1401–18. https://doi.org/10.1172/JCI78018.
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    • Virshup, David M. “Moving upstream in the war on WNTs.” J Clin Invest 125, no. 3 (March 2, 2015): 975–77. https://doi.org/10.1172/JCI80819.
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    • Stewart, Jonathan, Jacqueline James, Glenn W. McCluggage, Stephen McQuaid, Kenneth Arthur, David Boyle, Paul Mullan, et al. “Analysis of wntless (WLS) expression in gastric, ovarian, and breast cancers reveals a strong association with HER2 overexpression.” Mod Pathol 28, no. 3 (March 2015): 428–36. https://doi.org/10.1038/modpathol.2014.114.
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    • Yim, D. G. R., S. Ghosh, G. R. Guy, and D. M. Virshup. “Casein kinase 1 regulates Sprouty2 in FGF-ERK signaling.” Oncogene 34, no. 4 (January 22, 2015): 474–84. https://doi.org/10.1038/onc.2013.564.
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    • Xu, Peng, David M. Virshup, and Sang Hyun Lee. “B56-PP2A regulates motor dynamics for mitotic chromosome alignment.” J Cell Sci 127, no. Pt 21 (November 1, 2014): 4567–73. https://doi.org/10.1242/jcs.154609.
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    • Yu, Jia, and David M. Virshup. “Updating the Wnt pathways.” Biosci Rep 34, no. 5 (October 17, 2014). https://doi.org/10.1042/BSR20140119.
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    • Low, Ivan Cherh Chiet, Thomas Loh, Yiqing Huang, David M. Virshup, and Shazib Pervaiz. “Ser70 phosphorylation of Bcl-2 by selective tyrosine nitration of PP2A-B56δ stabilizes its antiapoptotic activity.” Blood 124, no. 14 (October 2, 2014): 2223–34. https://doi.org/10.1182/blood-2014-03-563296.
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    • Asad, M., M. K. Wong, T. Z. Tan, M. Choolani, J. Low, S. Mori, D. Virshup, J. P. Thiery, and R. Y. -. J. Huang. “FZD7 drives in vitro aggressiveness in Stem-A subtype of ovarian cancer via regulation of non-canonical Wnt/PCP pathway.” Cell Death Dis 5 (July 17, 2014): e1346. https://doi.org/10.1038/cddis.2014.302.
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    • Zhu, Yunhua, Oleg N. Demidov, Amanda M. Goh, David M. Virshup, David P. Lane, and Dmitry V. Bulavin. “Phosphatase WIP1 regulates adult neurogenesis and WNT signaling during aging.” J Clin Invest 124, no. 7 (July 2014): 3263–73. https://doi.org/10.1172/JCI73015.
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    • MacDonald, Bryan T., Annie Hien, Xinjun Zhang, Oladoyin Iranloye, David M. Virshup, Marian L. Waterman, and Xi He. “Disulfide bond requirements for active Wnt ligands.” J Biol Chem 289, no. 26 (June 27, 2014): 18122–36. https://doi.org/10.1074/jbc.M114.575027.
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    • Kabiri, Zahra, Gediminas Greicius, Babita Madan, Steffen Biechele, Zhendong Zhong, Hamed Zaribafzadeh, Hamed Edison, et al. “Stroma provides an intestinal stem cell niche in the absence of epithelial Wnts.” Development 141, no. 11 (June 2014): 2206–15. https://doi.org/10.1242/dev.104976.
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    • Yu, Jia, Joanne Chia, Claire Ann Canning, C Michael Jones, Frédéric A. Bard, and David M. Virshup. “WLS retrograde transport to the endoplasmic reticulum during Wnt secretion.” Dev Cell 29, no. 3 (May 12, 2014): 277–91. https://doi.org/10.1016/j.devcel.2014.03.016.
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    • Madan, Babita, Zhiyuan Ke, Shermaine Q. Y. Lim, Jenefer Alam, Soo Yei Ho, Duraiswamy A. Jeyaraj, Kakaly Ghosh, et al. “Abstract C248: Novel PORCN inhibitors are safe and effective in the treatment of WNT-dependent cancers.” Therapeutic Agents: Other, November 2013. https://doi.org/10.1158/1535-7163.targ-13-c248.
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    • Yim, Daniel G. R., and David M. Virshup. “Unwinding the Wnt action of casein kinase 1.” Cell Res 23, no. 6 (June 2013): 737–38. https://doi.org/10.1038/cr.2013.51.
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    • Xu, Peng, Elizabeth A. Raetz, Mayumi Kitagawa, David M. Virshup, and Sang Hyun Lee. “BUBR1 recruits PP2A via the B56 family of targeting subunits to promote chromosome congression.” Biol Open 2, no. 5 (May 15, 2013): 479–86. https://doi.org/10.1242/bio.20134051.
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    • Stewart, Jonathan, Jacqueline James, Glenn McCluggage, Stephen McQuaid, David Boyle, Kenneth Arthur, Paul Mullan, et al. “A strong correlation between expression of Wntless and of human epidermal growth factor receptor 2 in gastric, ovarian, and breast cancers suggests a novel-signalling pathway involving NFκB and STAT3.” The Lancet 381 (February 2013): S106–S106. https://doi.org/10.1016/s0140-6736(13)60546-2.
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    • Proffitt, Kyle David, Babita Madan, Zhiyuan Ke, Vishal Pendharkar, Lijun Ding, May Ann Lee, Rami N. Hannoush, and David M. Virshup. “Pharmacological inhibition of the Wnt acyltransferase PORCN prevents growth of WNT-driven mammary cancer.” Cancer Res 73, no. 2 (January 15, 2013): 502–7. https://doi.org/10.1158/0008-5472.CAN-12-2258.
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    • Proffitt, Kyle D., and David M. Virshup. “Precise regulation of porcupine activity is required for physiological Wnt signaling.” J Biol Chem 287, no. 41 (October 5, 2012): 34167–78. https://doi.org/10.1074/jbc.M112.381970.
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    • Najdi, Rani, Kyle Proffitt, Stephanie Sprowl, Simran Kaur, Jia Yu, Tracy M. Covey, David M. Virshup, and Marian L. Waterman. “A uniform human Wnt expression library reveals a shared secretory pathway and unique signaling activities.” Differentiation 84, no. 2 (September 2012): 203–13. https://doi.org/10.1016/j.diff.2012.06.004.
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    • Coombs, G. S., A. A. Schmitt, C. A. Canning, A. Alok, I. C. C. Low, N. Banerjee, S. Kaur, et al. “Modulation of Wnt/β-catenin signaling and proliferation by a ferrous iron chelator with therapeutic efficacy in genetically engineered mouse models of cancer.” Oncogene 31, no. 2 (January 12, 2012): 213–25. https://doi.org/10.1038/onc.2011.228.
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    • Covey, Tracy M., Simran Kaur, Tina Tan Ong, Kyle D. Proffitt, Yonghui Wu, Patrick Tan, and David M. Virshup. “PORCN moonlights in a Wnt-independent pathway that regulates cancer cell proliferation.” Plos One 7, no. 4 (2012): e34532. https://doi.org/10.1371/journal.pone.0034532.
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    • Ooi, Chia Huey, Hue Kian Oh, Hannah Zhu’ai Wang, Angie Lay Keng Tan, Jeanie Wu, Minghui Lee, Sun Young Rha, Hyun Cheol Chung, David Marc Virshup, and Patrick Tan. “A densely interconnected genome-wide network of microRNAs and oncogenic pathways revealed using gene expression signatures.” Plos Genet 7, no. 12 (December 2011): e1002415. https://doi.org/10.1371/journal.pgen.1002415.
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    • Cheong, J. K., T. H. Nguyen, H. Wang, P. Tan, P. M. Voorhoeve, S. H. Lee, and D. M. Virshup. “IC261 induces cell cycle arrest and apoptosis of human cancer cells via CK1δ/ɛ and Wnt/β-catenin independent inhibition of mitotic spindle formation.” Oncogene 30, no. 22 (June 2, 2011): 2558–69. https://doi.org/10.1038/onc.2010.627.
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    • Cheong, Jit Kong, and David M. Virshup. “Casein kinase 1: Complexity in the family.” Int J Biochem Cell Biol 43, no. 4 (April 2011): 465–69. https://doi.org/10.1016/j.biocel.2010.12.004.
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    • Virshup, David M., and Philipp Kaldis. “Cell biology. Enforcing the Greatwall in mitosis.” Science 330, no. 6011 (December 17, 2010): 1638–39. https://doi.org/10.1126/science.1199898.
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    • Covey, Tracy M., Kornelia Edes, Gary S. Coombs, David M. Virshup, and Frank A. Fitzpatrick. “Alkylation of the tumor suppressor PTEN activates Akt and β-catenin signaling: a mechanism linking inflammation and oxidative stress with cancer.” Plos One 5, no. 10 (October 21, 2010): e13545. https://doi.org/10.1371/journal.pone.0013545.
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    • Coombs, Gary S., Jia Yu, Claire A. Canning, Charles A. Veltri, Tracy M. Covey, Jit K. Cheong, Velani Utomo, et al. “WLS-dependent secretion of WNT3A requires Ser209 acylation and vacuolar acidification.” J Cell Sci 123, no. Pt 19 (October 1, 2010): 3357–67. https://doi.org/10.1242/jcs.072132.
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    • McCulloch, Malcolm W. B., Tim S. Bugni, Gisela P. Concepcion, Gary S. Coombs, Mary Kay Harper, Simran Kaur, Gina C. Mangalindan, et al. “Carteriosulfonic acids A-C, GSK-3beta inhibitors from a Carteriospongia sp.” J Nat Prod 72, no. 9 (September 2009): 1651–56. https://doi.org/10.1021/np900336f.
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    • Wang, C., K. C. Chang, G. Somers, D. Virshup, B. T. Ang, C. Tang, F. Yu, and H. Wang. “Erratum: Protein phosphatase 2A regulates self-renewal of Drosophila neural stem cells (Development (2009) vol. 136 (2287-2296)).” Development 136, no. 17 (September 1, 2009): 3031. https://doi.org/10.1242/dev.042432.
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    • Wang, Cheng, Kai Chen Chang, Gregory Somers, David Virshup, Beng Ti Ang, Carol Tang, Fengwei Yu, and Hongyan Wang. “Protein phosphatase 2A regulates self-renewal of Drosophila neural stem cells.” Development 136, no. 13 (July 2009): 2287–96. https://doi.org/10.1242/dev.035758.
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    • Virshup, David M., and Daniel B. Forger. “Keeping the beat in the rising heat.” Cell 137, no. 4 (May 15, 2009): 602–4. https://doi.org/10.1016/j.cell.2009.04.051.
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    • McCulloch, Malcolm W. B., Gary S. Coombs, Nikhil Banerjee, Tim S. Bugni, Kendell M. Cannon, Mary Kay Harper, Charles A. Veltri, David M. Virshup, and Chris M. Ireland. “Psammaplin A as a general activator of cell-based signaling assays via HDAC inhibition and studies on some bromotyrosine derivatives.” Bioorg Med Chem 17, no. 6 (March 15, 2009): 2189–98. https://doi.org/10.1016/j.bmc.2008.10.077.
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    • Virshup, David M., and Shirish Shenolikar. “From promiscuity to precision: protein phosphatases get a makeover.” Mol Cell 33, no. 5 (March 13, 2009): 537–45. https://doi.org/10.1016/j.molcel.2009.02.015.
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    • Coombs, Gary S., Tracy M. Covey, and David M. Virshup. “Wnt signaling in development, disease and translational medicine.” Curr Drug Targets 9, no. 7 (July 2008): 513–31. https://doi.org/10.2174/138945008784911796.
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    • Meng, Qing-Jun, Larisa Logunova, Elizabeth S. Maywood, Monica Gallego, Jake Lebiecki, Timothy M. Brown, Martin Sládek, et al. “Setting clock speed in mammals: the CK1 epsilon tau mutation in mice accelerates circadian pacemakers by selectively destabilizing PERIOD proteins.” Neuron 58, no. 1 (April 10, 2008): 78–88. https://doi.org/10.1016/j.neuron.2008.01.019.
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    • Forester, Craig M., Jessica Maddox, Justin V. Louis, Jozef Goris, and David M. Virshup. “Control of mitotic exit by PP2A regulation of Cdc25C and Cdk1.” Proc Natl Acad Sci U S A 104, no. 50 (December 11, 2007): 19867–72. https://doi.org/10.1073/pnas.0709879104.
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    • Forger, Daniel, Didier Gonze, David Virshup, and David K. Welsh. “Beyond intuitive modeling: combining biophysical models with innovative experiments to move the circadian clock field forward.” J Biol Rhythms 22, no. 3 (June 2007): 200–210. https://doi.org/10.1177/0748730407301823.
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    • Virshup, David M., and Daniel B. Forger. “After hours keeps clock researchers CRYing Overtime.” Cell 129, no. 5 (June 1, 2007): 857–59. https://doi.org/10.1016/j.cell.2007.05.015.
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    • Luo, Wen, Annita Peterson, Benjamin A. Garcia, Gary Coombs, Bente Kofahl, Reinhart Heinrich, Jeffrey Shabanowitz, Donald F. Hunt, H Joseph Yost, and David M. Virshup. “Protein phosphatase 1 regulates assembly and function of the beta-catenin degradation complex.” Embo J 26, no. 6 (March 21, 2007): 1511–21. https://doi.org/10.1038/sj.emboj.7601607.
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    • Tsai, I-Chun, Jeffrey D. Amack, Zhong-Hua Gao, Vimla Band, H Joseph Yost, and David M. Virshup. “A Wnt-CKIvarepsilon-Rap1 pathway regulates gastrulation by modulating SIPA1L1, a Rap GTPase activating protein.” Dev Cell 12, no. 3 (March 2007): 335–47. https://doi.org/10.1016/j.devcel.2007.02.009.
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    • Tsai, I-Chun, Margaret Woolf, Deborah W. Neklason, William W. Branford, H Joseph Yost, Randall W. Burt, and David M. Virshup. “Disease-associated casein kinase I delta mutation may promote adenomatous polyps formation via a Wnt/beta-catenin independent mechanism.” Int J Cancer 120, no. 5 (March 1, 2007): 1005–12. https://doi.org/10.1002/ijc.22368.
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    • Gallego, Monica, and David M. Virshup. “Post-translational modifications regulate the ticking of the circadian clock.” Nat Rev Mol Cell Biol 8, no. 2 (February 2007): 139–48. https://doi.org/10.1038/nrm2106.
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    • Saraf, Amit, David M. Virshup, and Stefan Strack. “Differential expression of the B'beta regulatory subunit of protein phosphatase 2A modulates tyrosine hydroxylase phosphorylation and catecholamine synthesis.” J Biol Chem 282, no. 1 (January 5, 2007): 573–80. https://doi.org/10.1074/jbc.M607407200.
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    • Virshup, D. M., E. J. Eide, D. B. Forger, M. Gallego, and E Vielhaber Harnish. “Reversible protein phosphorylation regulates circadian rhythms.” Cold Spring Harb Symp Quant Biol 72 (2007): 413–20. https://doi.org/10.1101/sqb.2007.72.048.
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    • Margolis, Seth S., Jennifer A. Perry, Craig M. Forester, Leta K. Nutt, Yanxiang Guo, Melanie J. Jardim, Michael J. Thomenius, et al. “Role for the PP2A/B56delta phosphatase in regulating 14-3-3 release from Cdc25 to control mitosis.” Cell 127, no. 4 (November 17, 2006): 759–73. https://doi.org/10.1016/j.cell.2006.10.035.
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    • Gallego, Monica, Heeseog Kang, and David M. Virshup. “Protein phosphatase 1 regulates the stability of the circadian protein PER2.” Biochem J 399, no. 1 (October 1, 2006): 169–75. https://doi.org/10.1042/BJ20060678.
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    • Klimowski, Laura K., Benjamin A. Garcia, Jeffrey Shabanowitz, Donald F. Hunt, and David M. Virshup. “Site-specific casein kinase 1epsilon-dependent phosphorylation of Dishevelled modulates beta-catenin signaling.” Febs J 273, no. 20 (October 2006): 4594–4602. https://doi.org/10.1111/j.1742-4658.2006.05462.x.
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    • Gallego, Monica, Erik J. Eide, Margaret F. Woolf, David M. Virshup, and Daniel B. Forger. “An opposite role for tau in circadian rhythms revealed by mathematical modeling.” Proc Natl Acad Sci U S A 103, no. 28 (July 11, 2006): 10618–23. https://doi.org/10.1073/pnas.0604511103.
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    • Swiatek, Wojciech, Heeseog Kang, Benjamin A. Garcia, Jeffery Shabanowitz, Gary S. Coombs, Donald F. Hunt, and David M. Virshup. “Negative regulation of LRP6 function by casein kinase I epsilon phosphorylation.” J Biol Chem 281, no. 18 (May 5, 2006): 12233–41. https://doi.org/10.1074/jbc.M510580200.
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    • Eide, Erik J., Margaret F. Woolf, Heeseog Kang, Peter Woolf, William Hurst, Fernando Camacho, Erica L. Vielhaber, Andrew Giovanni, and David M. Virshup. “Control of mammalian circadian rhythm by CKIepsilon-regulated proteasome-mediated PER2 degradation.” Mol Cell Biol 25, no. 7 (April 2005): 2795–2807. https://doi.org/10.1128/MCB.25.7.2795-2807.2005.
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    • Firulli, Beth A., Dayana Krawchuk, Victoria E. Centonze, Neil Vargesson, David M. Virshup, Simon J. Conway, Peter Cserjesi, Ed Laufer, and Anthony B. Firulli. “Altered Twist1 and Hand2 dimerization is associated with Saethre-Chotzen syndrome and limb abnormalities.” Nat Genet 37, no. 4 (April 2005): 373–81. https://doi.org/10.1038/ng1525.
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    • Gallego, Monica, and David M. Virshup. “Protein serine/threonine phosphatases: life, death, and sleeping.” Curr Opin Cell Biol 17, no. 2 (April 2005): 197–202. https://doi.org/10.1016/j.ceb.2005.01.002.
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    • Eide, Erik J., Heeseog Kang, Stephanie Crapo, Monica Gallego, and David M. Virshup. “Casein kinase I in the mammalian circadian clock.” Methods Enzymol 393 (2005): 408–18. https://doi.org/10.1016/S0076-6879(05)93019-X.
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    • Okamura, Heidi, Carmen Garcia-Rodriguez, Holly Martinson, Jun Qin, David M. Virshup, and Anjana Rao. “A conserved docking motif for CK1 binding controls the nuclear localization of NFAT1.” Mol Cell Biol 24, no. 10 (May 2004): 4184–95. https://doi.org/10.1128/MCB.24.10.4184-4195.2004.
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    • Swiatek, Wojciech, I-Chun Tsai, Laura Klimowski, Andrea Pepler, Janet Barnette, H Joseph Yost, and David M. Virshup. “Regulation of casein kinase I epsilon activity by Wnt signaling.” J Biol Chem 279, no. 13 (March 26, 2004): 13011–17. https://doi.org/10.1074/jbc.M304682200.
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    • Firulli, Beth A., David M. Virshup, and Anthony B. Firulli. “Phosphopeptide mapping of proteins ectopically expressed in tissue culture cell lines.” Biol Proced Online 6 (2004): 16–22. https://doi.org/10.1251/bpo69.
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    • Firulli, Beth A., Marthe J. Howard, Jennifer R. McDaid, Leanne McIlreavey, Karen M. Dionne, Victoria E. Centonze, Peter Cserjesi, David M. Virshup, and Anthony B. Firulli. “PKA, PKC, and the protein phosphatase 2A influence HAND factor function: a mechanism for tissue-specific transcriptional regulation.” Mol Cell 12, no. 5 (November 2003): 1225–37. https://doi.org/10.1016/s1097-2765(03)00425-8.
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    • Liu, Feng, David M. Virshup, Angus C. Nairn, and Paul Greengard. “Mechanism of regulation of casein kinase I activity by group I metabotropic glutamate receptors.” J Biol Chem 277, no. 47 (November 22, 2002): 45393–99. https://doi.org/10.1074/jbc.M204499200.
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    • Liu, Ying, David M. Virshup, Raymond L. White, and Lih-Ching Hsu. “Regulation of BRCA1 phosphorylation by interaction with protein phosphatase 1alpha.” Cancer Res 62, no. 22 (November 15, 2002): 6357–61.
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    • Raetz, Elizabeth A., Sherrie L. Perkins, Marlee A. Carlson, Kevin P. Schooler, William L. Carroll, and David M. Virshup. “The nucleophosmin-anaplastic lymphoma kinase fusion protein induces c-Myc expression in pediatric anaplastic large cell lymphomas.” Am J Pathol 161, no. 3 (September 2002): 875–83. https://doi.org/10.1016/S0002-9440(10)64248-4.
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    • Li, Xinghai, Anne Scuderi, Anthea Letsou, and David M. Virshup. “B56-associated protein phosphatase 2A is required for survival and protects from apoptosis in Drosophila melanogaster.” Mol Cell Biol 22, no. 11 (June 2002): 3674–84. https://doi.org/10.1128/MCB.22.11.3674-3684.2002.
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    • Eide, Erik J., Erica L. Vielhaber, William A. Hinz, and David M. Virshup. “The circadian regulatory proteins BMAL1 and cryptochromes are substrates of casein kinase Iepsilon.” J Biol Chem 277, no. 19 (May 10, 2002): 17248–54. https://doi.org/10.1074/jbc.M111466200.
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    • Toh, K. L., C. Thulin, Y. H. Fu, L. J. Ptacek, and D. M. Virshup. “Sequential multisite phosphorylation by casein kinase I epsilon (CKI epsilon).” Faseb Journal 16, no. 5 (March 22, 2002): A917–A917.
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    • Slayton, William B., Gerald J. Spangrude, Zhong Chen, Wayne F. Greene, and David Virshup. “Lineage-specific trisomy 21 in a neonate with resolving transient myeloproliferative syndrome.” J Pediatr Hematol Oncol 24, no. 3 (March 2002): 224–26. https://doi.org/10.1097/00043426-200203000-00013.
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    • Gao, Zhong-Hua, Joni M. Seeling, Virginia Hill, April Yochum, and David M. Virshup. “Casein kinase I phosphorylates and destabilizes the beta-catenin degradation complex.” Proc Natl Acad Sci U S A 99, no. 3 (February 5, 2002): 1182–87. https://doi.org/10.1073/pnas.032468199.
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    • Li, Xinghai, and David M. Virshup. “Two conserved domains in regulatory B subunits mediate binding to the A subunit of protein phosphatase 2A.” Eur J Biochem 269, no. 2 (January 2002): 546–52. https://doi.org/10.1046/j.0014-2956.2001.02680.x.
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    • Vielhaber, E. L., D. Duricka, K. S. Ullman, and D. M. Virshup. “Nuclear export of mammalian PERIOD proteins.” J Biol Chem 276, no. 49 (December 7, 2001): 45921–27. https://doi.org/10.1074/jbc.M107726200.
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    • Li, X., H. J. Yost, D. M. Virshup, and J. M. Seeling. “Protein phosphatase 2A and its B56 regulatory subunit inhibit Wnt signaling in Xenopus.” Embo J 20, no. 15 (August 1, 2001): 4122–31. https://doi.org/10.1093/emboj/20.15.4122.
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    • Seeling, J. M., X. Li, H. J. Yost, and D. M. Virshup. “PP2A and its B56 regulatory subunit inhibit Wnt signaling in Xenopus.” Developmental Biology 235, no. 1 (July 1, 2001): 254–254.
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    • Eide, E. J., and D. M. Virshup. “Casein kinase I: another cog in the circadian clockworks.” Chronobiol Int 18, no. 3 (May 2001): 389–98. https://doi.org/10.1081/cbi-100103963.
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    • Toh, K. L., C. R. Jones, Y. He, E. J. Eide, W. A. Hinz, D. M. Virshup, L. J. Ptácek, and Y. H. Fu. “An hPer2 phosphorylation site mutation in familial advanced sleep phase syndrome.” Science 291, no. 5506 (February 9, 2001): 1040–43. https://doi.org/10.1126/science.1057499.
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    • Camacho, F., M. Cilio, Y. Guo, D. M. Virshup, K. Patel, O. Khorkova, S. Styren, B. Morse, Z. Yao, and G. A. Keesler. “Human casein kinase Idelta phosphorylation of human circadian clock proteins period 1 and 2.” Febs Lett 489, no. 2–3 (February 2, 2001): 159–65. https://doi.org/10.1016/s0014-5793(00)02434-0.
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    • Kornberg, A. “Public funding of understanding nature.” Iubmb Life 51, no. 2 (February 2001): 71–72. https://doi.org/10.1080/15216540117461.
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    • Raetz, E. A., S. L. Perkins, M. A. Carlson, K. P. Schooler, and D. M. Virshup. “Pediatric ALK-positive lymphomas coexpress c-Myc.” Blood 96, no. 11 (November 16, 2000): 128A-128A.
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    • Vielhaber, E., E. Eide, A. Rivers, Z. H. Gao, and D. M. Virshup. “Nuclear entry of the circadian regulator mPER1 is controlled by mammalian casein kinase I epsilon.” Mol Cell Biol 20, no. 13 (July 2000): 4888–99. https://doi.org/10.1128/MCB.20.13.4888-4899.2000.
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    • Keesler, G. A., F. Camacho, Y. Guo, D. Virshup, C. Mondadori, and Z. Yao. “Phosphorylation and destabilization of human period I clock protein by human casein kinase I epsilon.” Neuroreport 11, no. 5 (April 7, 2000): 951–55. https://doi.org/10.1097/00001756-200004070-00011.
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    • Virshup, D. M. “Protein phosphatase 2A: a panoply of enzymes.” Curr Opin Cell Biol 12, no. 2 (April 2000): 180–85. https://doi.org/10.1016/s0955-0674(99)00074-5.
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    • Gao, Z. H., J. Metherall, and D. M. Virshup. “Identification of casein kinase I substrates by in vitro expression cloning screening.” Biochem Biophys Res Commun 268, no. 2 (February 16, 2000): 562–66. https://doi.org/10.1006/bbrc.2000.2168.
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    • Gietzen, K. F., and D. M. Virshup. “Identification of inhibitory autophosphorylation sites in casein kinase I epsilon.” J Biol Chem 274, no. 45 (November 5, 1999): 32063–70. https://doi.org/10.1074/jbc.274.45.32063.
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    • Goldsby, R. E., S. L. Perkins, D. M. Virshup, A. R. Brothman, and C. S. Bruggers. “Lymphoblastic lymphoma and excessive toxicity from chemotherapy: an unusual presentation for Fanconi anemia.” J Pediatr Hematol Oncol 21, no. 3 (May 1999): 240–43. https://doi.org/10.1097/00043426-199905000-00014.
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    • Honeggar, M., J. Krall, D. M. Virshup, and M. A. Movsesian. “Expression of multiple PP2A serine/threonine phosphatase B56 (B ') subunit isoforms in human cardiac and vascular smooth muscle myocytes.” Journal of Investigative Medicine 47, no. 4 (April 1, 1999): 194A-194A.
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    • Seeling, J. M., J. R. Miller, R. Gil, R. T. Moon, R. White, and D. M. Virshup. “Regulation of beta-catenin signaling by the B56 subunit of protein phosphatase 2A.” Science 283, no. 5410 (March 26, 1999): 2089–91. https://doi.org/10.1126/science.283.5410.2089.
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    • Honeggar, M., J. Krall, D. M. Virshup, and M. A. Movsesian. “Expression of multiple PP2A serine/threonine phosphatase B56 (B ') subunit isoforms in human cardiac and vascular smooth muscle myocytes.” Faseb Journal 13, no. 4 (March 12, 1999): A468–A468.
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    • Rivers, A., K. F. Gietzen, E. Vielhaber, and D. M. Virshup. “Regulation of casein kinase I epsilon and casein kinase I delta by an in vivo futile phosphorylation cycle.” J Biol Chem 273, no. 26 (June 26, 1998): 15980–84. https://doi.org/10.1074/jbc.273.26.15980.
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    • McDonald, J. M., E. C. Douglass, R. Fisher, C. F. Geiser, C. E. Krill, L. C. Strong, D. Virshup, and V. Huff. “Linkage of familial Wilms' tumor predisposition to chromosome 19 and a two-locus model for the etiology of familial tumors.” Cancer Res 58, no. 7 (April 1, 1998): 1387–90.
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    • Cegielska, A., K. F. Gietzen, A. Rivers, and D. M. Virshup. “Autoinhibition of casein kinase I epsilon (CKI epsilon) is relieved by protein phosphatases and limited proteolysis.” J Biol Chem 273, no. 3 (January 16, 1998): 1357–64. https://doi.org/10.1074/jbc.273.3.1357.
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    • McCright, B., and D. M. Virshup. “Identifying protein phosphatase 2A interacting proteins using the yeast two-hybrid method.” Methods Mol Biol 93 (1998): 263–77. https://doi.org/10.1385/0-89603-468-2:263.
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    • Gietzen, K. J., and D. M. Virshup. “Determinants of casein kinase i epsilon (ckir) autoinhibition.” Faseb Journal 11, no. 9 (December 1, 1997).
    • Virshup, D., K. Oietzen, A. Rivers, and U. Utah. “Regulation of casein kinase i.” Faseb Journal 11, no. 9 (December 1, 1997).
    • Gietzen, K. J., and D. M. Virshup. “Determinants of casein kinase I epsilon (CKI epsilon) autoinhibition.” Faseb Journal 11, no. 9 (July 31, 1997): A1356–A1356.
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    • McCright, B., A. M. Rivers, S. Audlin, and D. M. Virshup. “The B56 family of protein phosphatase 2A (PP2A) regulatory subunits encodes differentiation-induced phosphoproteins that target PP2A to both nucleus and cytoplasm.” J Biol Chem 271, no. 36 (September 6, 1996): 22081–89. https://doi.org/10.1074/jbc.271.36.22081.
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    • McCright, B., A. R. Brothman, and D. M. Virshup. “Assignment of human protein phosphatase 2A regulatory subunit genes b56alpha, b56beta, b56gamma, b56delta, and b56epsilon (PPP2R5A-PPP2R5E), highly expressed in muscle and brain, to chromosome regions 1q41, 11q12, 3p21, 6p21.1, and 7p11.2 --> p12.” Genomics 36, no. 1 (August 15, 1996): 168–70. https://doi.org/10.1006/geno.1996.0438.
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    • McCright, B., and D. M. Virshup. “Identification of a new family of protein phosphatase 2A regulatory subunits.” J Biol Chem 270, no. 44 (November 3, 1995): 26123–28. https://doi.org/10.1074/jbc.270.44.26123.
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    • Fish, K. J., A. Cegielska, M. E. Getman, G. M. Landes, and D. M. Virshup. “Isolation and characterization of human casein kinase I epsilon (CKI), a novel member of the CKI gene family.” J Biol Chem 270, no. 25 (June 23, 1995): 14875–83. https://doi.org/10.1074/jbc.270.25.14875.
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    • Sivak, L. E., and D. M. Virshup. “Occurrence of Henoch-Schönlein purpura in a child with Wilms' tumor.” Med Pediatr Oncol 24, no. 3 (March 1995): 213–14. https://doi.org/10.1002/mpo.2950240314.
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    • Cegielska, A., S. Shaffer, R. Derua, J. Goris, and D. M. Virshup. “Different oligomeric forms of protein phosphatase 2A activate and inhibit simian virus 40 DNA replication.” Mol Cell Biol 14, no. 7 (July 1994): 4616–23. https://doi.org/10.1128/mcb.14.7.4616-4623.1994.
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    • Cegielska, A., I. Moarefi, E. Fanning, and D. M. Virshup. “T-antigen kinase inhibits simian virus 40 DNA replication by phosphorylation of intact T antigen on serines 120 and 123.” J Virol 68, no. 1 (January 1994): 269–75. https://doi.org/10.1128/JVI.68.1.269-275.1994.
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    • Coombs, R. H., and B. B. Virshup. “Enhancing the psychological health of medical students: the student well-being committee.” Med Educ 28, no. 1 (January 1994): 47–54. https://doi.org/10.1111/j.1365-2923.1994.tb02684.x.
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    • Cegielska, A., and D. M. Virshup. “Control of simian virus 40 DNA replication by the HeLa cell nuclear kinase casein kinase I.” Mol Cell Biol 13, no. 2 (February 1993): 1202–11. https://doi.org/10.1128/mcb.13.2.1202-1211.1993.
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    • VIRSHUP, D. M., A. CEGIELSKA, K. FISH, S. SHAFFER, and D. HILLAM. “CONTROL OF SV40 DNA-REPLICATION BY THE RECIPROCAL ACTION OF CASEIN KINASE-I AND PROTEIN PHOSPHATASE 2AC.” Journal of Cellular Biochemistry, January 9, 1993, 305–305.
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    • Virshup, D. M., A. A. Russo, and T. J. Kelly. “Mechanism of activation of simian virus 40 DNA replication by protein phosphatase 2A.” Mol Cell Biol 12, no. 11 (November 1992): 4883–95. https://doi.org/10.1128/mcb.12.11.4883-4895.1992.
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    • Modlin, J. F., R. Dagan, L. E. Berlin, D. M. Virshup, R. H. Yolken, and M. Menegus. “Focal encephalitis with enterovirus infections.” Pediatrics 88, no. 4 (October 1991): 841–45.
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    • Scheidtmann, K. H., D. M. Virshup, and T. J. Kelly. “Protein phosphatase 2A dephosphorylates simian virus 40 large T antigen specifically at residues involved in regulation of DNA-binding activity.” J Virol 65, no. 4 (April 1991): 2098–2101. https://doi.org/10.1128/JVI.65.4.2098-2101.1991.
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    • ERDILE, L. F., K. L. COLLINS, A. RUSSO, P. SIMANCEK, D. SMALL, C. UMBRICHT, D. VIRSHUP, et al. “INITIATION OF SV40 DNA-REPLICATION - MECHANISM AND CONTROL.” Cold Spring Harbor Symposia on Quantitative Biology 56 (January 1, 1991): 303–13.
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    • Erdile, L. F., K. L. Collins, A. Russo, P. Simancek, D. Small, C. Umbricht, D. Virshup, L. Cheng, S. Randall, and D. Weinberg. “Initiation of SV40 DNA replication: mechanism and control.” Cold Spring Harb Symp Quant Biol 56 (1991): 303–13. https://doi.org/10.1101/sqb.1991.056.01.037.
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    • Weinberg, D. H., K. L. Collins, P. Simancek, A. Russo, M. S. Wold, D. M. Virshup, and T. J. Kelly. “Reconstitution of simian virus 40 DNA replication with purified proteins.” Proc Natl Acad Sci U S A 87, no. 22 (November 1990): 8692–96. https://doi.org/10.1073/pnas.87.22.8692.
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    • Virshup, D. M. “DNA replication.” Curr Opin Cell Biol 2, no. 3 (June 1990): 453–60. https://doi.org/10.1016/0955-0674(90)90127-z.
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    • Moo-Penn, W. F., D. C. Swan, T. K. Hine, R. M. Baine, D. L. Jue, J. M. Benson, M. H. Johnson, D. M. Virshup, and W. H. Zinkham. “Hb Catonsville (glutamic acid inserted between Pro-37(C2)alpha and Thr-38(C3)alpha). Nonallelic gene conversion in the globin system?” J Biol Chem 264, no. 36 (December 25, 1989): 21454–57.
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    • Virshup, D. M., M. G. Kauffman, and T. J. Kelly. “Activation of SV40 DNA replication in vitro by cellular protein phosphatase 2A.” Embo J 8, no. 12 (December 1, 1989): 3891–98.
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    • Virshup, D. M., and T. J. Kelly. “Purification of replication protein C, a cellular protein involved in the initial stages of simian virus 40 DNA replication in vitro.” Proc Natl Acad Sci U S A 86, no. 10 (May 1989): 3584–88. https://doi.org/10.1073/pnas.86.10.3584.
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    • Wold, M. S., D. H. Weinberg, D. M. Virshup, J. J. Li, and T. J. Kelly. “Identification of cellular proteins required for simian virus 40 DNA replication.” J Biol Chem 264, no. 5 (February 15, 1989): 2801–9.
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    • Virshup, D. M., and V. Bennett. “Clathrin-coated vesicle assembly polypeptides: physical properties and reconstitution studies with brain membranes.” J Cell Biol 106, no. 1 (January 1988): 39–50. https://doi.org/10.1083/jcb.106.1.39.
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    • Jerdan, M. S., R. S. Shapiro, N. B. Smith, D. M. Virshup, and A. F. Hood. “Cutaneous manifestations of Corynebacterium group JK sepsis.” J Am Acad Dermatol 16, no. 2 Pt 2 (February 1987): 444–47. https://doi.org/10.1016/s0190-9622(87)70058-9.
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    • VIRSHUP, D. M., and V. BENNETT. “CLATHRIN-COATED VESICLE ASSEMBLY POLYPEPTIDES - PHYSICAL-PROPERTIES AND IDENTIFICATION OF A NEW SUBUNIT.” Journal of Cell Biology 103, no. 5 (November 1, 1986): A53–A53.
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    • Agre, P., D. Virshup, and V. Bennett. “Bepridil and cetiedil. Vasodilators which inhibit Ca2+-dependent calmodulin interactions with erythrocyte membranes.” J Clin Invest 74, no. 3 (September 1984): 812–20. https://doi.org/10.1172/JCI111497.
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    • Virshup, D. M., W. H. Zinkham, R. L. Sirota, and W. S. Caughey. “Unique sensitivity of Hb Zürich to oxidative injury by phenazopyridine: reversal of the effects by elevating carboxyhemoglobin levels in vivo and in vitro.” Am J Hematol 14, no. 4 (June 1983): 315–24. https://doi.org/10.1002/ajh.2830140402.
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    • Virshup, D. M., H. Zinkham Wm., R. L. Sirota, and W. S. Caughey. “Unique sensitivity of HB Zurich to oxidative injury by phenazopyridine: Reversal of the effect by elevating carboxyhemoglobin levels in vivo and in vitro.” Pediatric Research 16, no. 4 II (January 1, 1982).
    • Kaur, Amanpreet, Sugunavathi Sepramaniam, Jun Yi Stanley Lim, Siddhi Patnaik, Nathan Harmston, May Ann Lee, Enrico Petretto, David M. Virshup, and Babita Madan. “WNT inhibition creates a BRCA-like state in Wnt-addicted cancer,” n.d. https://doi.org/10.1101/2020.06.17.157024.
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  • Conference Papers

    • Ricci, Clarisse Gravina, Jonathan M. Philpott, Rajesh Narasimamurthy, Alfred M. Freeberg, Sabrina R. Hunt, Lauren E. Yee, Rebecca S. Pelofsky, Sarvind Tripathi, David M. Virshup, and Carrie L. Partch. “Free Energy Landscape of Casein Kinase Delta and its Implications for Circadian Rhythm.” In Biophysical Journal, 118:207A-208A. CELL PRESS, 2020.
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    • Tan, D., M. Ng, V. Subbiah, W. Messersmith, V. Teneggi, V. Diermayr, K. Ethirajulu, et al. “Phase I extension study of ETC-159 an oral PORCN inhibitor administered with bone protective treatment, in patients with advanced solid tumours.” In Annals of Oncology : Official Journal of the European Society for Medical Oncology, 29:ix23–24, 2018. https://doi.org/10.1093/annonc/mdy430.002.
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    • Pendharkar, Vishal, Yun Shan Chew, Vithya Manoharan, Choon Bing Low, Hongqian Esther Ong, Soo Yei Ho, Wei Ling Wang, et al. “Abstract 1172:In vivopharmacokinetic properties and antitumor efficacy of porcupine lead inhibitors in the orthotopic murine MMTV-Wnt1 breast tumor model and the human HPAF-II pancreatic xenograft mouse model.” In Experimental and Molecular Therapeutics. American Association for Cancer Research, 2017. https://doi.org/10.1158/1538-7445.am2017-1172.
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    • Ng, Matthew, David S. P. Tan, Vivek Subbiah, Colin D. Weekes, Vincenzo Teneggi, Veronica Diermayr, Kantharaj Ethirajulu, et al. “First-in-human phase 1 study of ETC-159 an oral PORCN inhbitor in patients with advanced solid tumours.” In Journal of Clinical Oncology, 35:2584–2584. American Society of Clinical Oncology (ASCO), 2017. https://doi.org/10.1200/jco.2017.35.15_suppl.2584.
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    • Teneggi, V., M. Ng, D. S. Tan, V. Subbiah, C. Weekes, V. Diermayr, K. Ethirajulu, et al. “A phase 1, first-in-human dose escalation study of ETC-159 in advanced or metastatic solid tumours.” In Annals of Oncology, Vol. 27. OXFORD UNIV PRESS, 2016.
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    • Herr, Keira, Raymond Serrano, Joanne Ong, Babita Madan, David Virshup, and Ann-Marie Chacko. “Characterization of anti-Frizzled Antibody [Zr-89]18R5 for PET Imaging of Pancreatic Cancer.” In Journal of Nuclear Medicine, Vol. 57, 2016.
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    • Poulsen, Anders, Soo Ho, Weilin Wang, Jenefer Alam, Athisayamani Duraiswamy, Grace Lin, Shi Ang, et al. “Scaffold hopping and optimization of maleimide based porcupine inhibitors.” In Abstracts of Papers of the American Chemical Society, Vol. 251. AMER CHEMICAL SOC, 2016.
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    • Cai, W., J. K. Cheong, E. Edison, A. Banerjee, T. Z. Tan, L. Gaboury, E. M. Yousef, et al. “Abstract P4-08-03: DEAD-box RNA helicase DP103 as a novel regulator of Wnt/β-catenin signaling pathway and promotes cancer stem cell-like behavior in triple negative breast cancers.” In Poster Session Abstracts. American Association for Cancer Research, 2016. https://doi.org/10.1158/1538-7445.sabcs15-p4-08-03.
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    • Asad, Mohammad, Meng Kang Wong, Tuan Zea Tan, David Virshup, Jean Paul Thiery, and Ruby Yun-Ju Huang. “Abstract A30: Frizzled-7 (FZD7)-mediated non-canonical Wnt-Planar Cell Polarity (PCP) signalling pathway as a novel molecular driver for the C5/Proliferative/Stem-A molecular subtype of ovarian cancer.” In Molecular Drivers of Ovarian Cancer (Other Than P53). American Association for Cancer Research, 2016. https://doi.org/10.1158/1557-3265.ovca15-a30.
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    • Ke, Zhiyuan, Babita Madan, Shermaine Q. Y. Lim, Sifang Wang, Jenefer Alam, Soo Yei Ho, Duraiswamy Athisayamani Jeyaraj, et al. “Abstract 4449: A novel Porcupine inhibitor is effective in the treatment of cancers with RNF43 mutations.” In Experimental and Molecular Therapeutics. American Association for Cancer Research, 2015. https://doi.org/10.1158/1538-7445.am2015-4449.
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    • Xiang, Wei, Jit Kong Cheong, Shi Hui Ang, Kartini Asari, Bryan Teo, David Virshup, and Charles Chuah. “Pyrvinium Selectively Targets Blast Phase Chronic Myeloid Leukaemia through Inhibition of Mitochondrial Respiration.” In Blood, Vol. 124, 2014.
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    • Lee, V., K. Chua, Y. Jia, and D. Virshup. “Expression of YJ5 (Wnt Carrier Protein Wntless) in bone tumours.” In Virchows Archiv, 465:S16–S16, 2014.
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    • Xu, P., E. Raetz, M. Kitagawa, D. Virshup, and S. Lee. “Phosphorylation Balance on the Kinetochore: BubR1 recruits B56-PP2A to regulate chromosome alignment.” In Molecular Biology of the Cell, Vol. 24, 2013.
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    • Ke, Z., S. Lim, S. Wang, J. Fulwood, K. Proffitt, B. Madan, M. Choong, H. Flotow, D. Virshup, and M. Lee. “172 Identification of Small Molecule Inhibitors of Wnt Secretion.” In European Journal of Cancer, 48:52–53. Elsevier BV, 2012. https://doi.org/10.1016/s0959-8049(12)71970-6.
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    • Cheong, Jit Kong, and David M. Virshup. “Abstract B264: IC261 induces cell cycle arrest and apoptosis of human cancer cells via a CK1δ/ε independent mechanism.” In Therapeutic Agents  Small Molecule Kinase Inhibitors. American Association for Cancer Research, 2009. https://doi.org/10.1158/1535-7163.targ-09-b264.
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    • Wang, Cheng, Kaichen Chang, Greory Somers, David Virshup, Beng Ti Ang, Carol Tang, Fengwei Yu, and Hongyan Wang. “08-P022 Protein phosphatase 2A regulates self-renewal of Drosophila neural stem cells.” In Mechanisms of Development, 126:S150–S150. Elsevier BV, 2009. https://doi.org/10.1016/j.mod.2009.06.329.
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    • Virshup, D. M., E. J. Eide, D. B. Forger, M. Gallego, E Vielhaber Harnish, and Cold Spring Harbor Lab Press. “Reversible Protein Phosphorylation Regulates Circadian Rhythms.” In Clocks and Rhythms, 72:413–20, 2007.
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    • Laufer, E., B. Firulli, D. Krawchuk, V. Centonze, D. Virshup, S. Conway, P. Cserjesi, and A. Firulli. “Hand2 and Twist1 interact antagonistically in the vertebrate limb.” In Developmental Biology, 271:590–590. ACADEMIC PRESS INC ELSEVIER SCIENCE, 2004.
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    • Tsai, I. C., M. Woolf, D. Neklason, R. Burt, and D. M. Virshup. “A disease-associated casein kinase 1 delta mutation alters its physiological roles and may promote adenomatous polyps via wnt/beta-catenin independent mechanism.” In Faseb Journal, 18:C299–C299. FEDERATION AMER SOC EXP BIOL, 2004.
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    • Fu, Y., C. R. Jones, K. Toh, D. Virshup, and L. J. Ptacek. “An hPer2 phosphorylation site mutation in familial Advanced Sleep-Phase Syndrome.” In American Journal of Human Genetics, 69:597–597. UNIV CHICAGO PRESS, 2001.
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    • Slayton, W. B., G. J. Spangrude, Z. Chen, and D. Virshup. “Residual trisomic cells in a patient with resolving transient myeloproliferative disorder and trisomy 21 mosaicism.” In Blood, 94:291B-291B. AMER SOC HEMATOLOGY, 1999.
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    • Vielhaber, E., and D. Virshup. “Identification of Casein Kinase I epsilon (CKI epsilon) substrates using a membrane based kinase assay.” In Faseb Journal, 11:A1359–A1359. FEDERATION AMER SOC EXP BIOL, 1997.
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    • Virshup, D., K. Gietzen, and A. Rivers. “Regulation of casein kinase I epsilon.” In Faseb Journal, 11:A1359–A1359. FEDERATION AMER SOC EXP BIOL, 1997.
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    • Goldsby, R. E., D. Carlson, E. D. Swalberg, and D. Virshup. “Comparing effects on temperature of early versus late leukodepletion in platelet transfusions.” In Blood, 86:1397–1397. W B SAUNDERS CO, 1995.
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    • VIRSHUP, D. M., M. S. WOLD, D. H. WEINBERG, M. G. KAUFFMAN, and T. J. KELLY. “CELLULAR PROTEINS INVOLVED IN SV40 DNA-REPLICATION INVITRO.” In Molecular Mechanisms in Dna Replication and Recombination, edited by C. C. RICHARDSON and I. R. LEHMAN, 127:303–14. WILEY-LISS, INC, 1990.
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