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Adam Dupuy, PhD

Associate Professor of Pathology
Associate Professor of Anatomy and Cell Biology

Contact Information

Primary Office
3202 Medical Education Research Facility (MERF)
Iowa City, IA 52242


BA, Biology and Chemistry, Central College
PhD, Molecular, Cellular, Developmental Biology, and Genetics, University of Minnesota
Postdoctoral Fellow, University of Minnesota Cancer Center
Postdoctoral Fellow, Mouse Cancer Genetics, National Cancer Institute

Education/Training Program Affiliations

Department of Cell and Developmental Graduate Program, Interdisciplinary Graduate Program in Genetics, Interdisciplinary Graduate Program in Molecular Medicine, Medical Scientist Training Program

Research Summary

The goal of my current research is to understand the role that somatic mutation plays in all aspects of tumor biology. Much of our work has made use of the Sleeping Beauty (SB) transposon system to engineer mouse cancer models in which somatic mutations are generated by transposon insertions. We and our collaborators have recently described a variety of tumor models using the approach, and these studies have identified many novel candidate cancer genes. The current work in my laboratory focuses on the genetics of hepatocellular carcinoma and T-cell leukemia. In addition, we have ongoing collaborative efforts to develop SB models of ovarian and endometrial cancer. Future work in the lab will focus on using our mouse models to address relevant questions encountered by oncologists in the clinic (e.g. metastasis, drug resistance).


Rogers, L. M., Wang, Z., Mott, S. L., Dupuy, A. J. & Weiner, G. J. (2020). A genetic screen to identify gain- and loss-of-function modifications that enhance T-cell infiltration into tumors. Cancer immunology research. PMID: 32611665.

Feddersen, C. R., Wadsworth, L. S., Zhu, E. Y., Vaughn, H. R., Voigt, A. P., Riordan, J. D. & Dupuy, A. J. (2019). A simplified transposon mutagenesis method to perform phenotypic forward genetic screens in cultured cells. BMC Genomics, 20(1), 497. PMID: 31208320.

Loeb, K. R., Hughes, B. T., Fissel, B. M., Osteen, N. J., Knoblaugh, S. E., Grim, J. E., Drury, T. J., Sarver, A., Dupuy, A. J. & Clurman, B. E. (2019). Insertional mutagenesis using the Sleeping Beauty transposon system identifies drivers of erythroleukemia in mice. Sci Rep, 9(1), 5488. PMID: 30940846.

Merritt, N. M., Fullenkamp, C. A., Hall, S. L., Qian, Q., Desai, C., Thomason, J., Lambertz, A. M., Dupuy, A. J., Darbro, B. & Tanas, M. R. (2018). A comprehensive evaluation of Hippo pathway silencing in sarcomas. Oncotarget, 9(60), 31620-31636. PMID: 30167083.

Riordan, J. D., Feddersen, C. R., Tschida, B. R., Beckmann, P. J., Keng, V. W., Linden, M. A., Amin, K., Stipp, C. S., Largaespada, D. A. & Dupuy, A. J. (2018). Chronic liver injury alters driver mutation profiles in hepatocellular carcinoma in mice. Hepatology, 67(3), 924-939. PMID: 28961327.

Ziyad, S., Riordan, J. D., Cavanaugh, A. M., Su, T., Hernandez, G. E., Hilfenhaus, G., Morselli, M., Huynh, K., Wang, K., Chen, J. N., Dupuy, A. J. & Iruela-Arispe, M. L. (2018). A Forward Genetic Screen Targeting the Endothelium Reveals a Regulatory Role for the Lipid Kinase Pi4ka in Myelo- and Erythropoiesis. Cell Reports, 22(5), 1211-1224. PMID: 29386109.

Rauckhorst, A. J., Gray, L. R., Sheldon, R. D., Fu, X., Pewa, A. D., Feddersen, C. R., Dupuy, A. J., Gibson-Corley, K. N., Cox, J. E., Burgess, S. C. & Taylor, E. B. (2017). The mitochondrial pyruvate carrier mediates high fat diet-induced increases in hepatic TCA cycle capacity. Molecular Metabolism, 6(11), 1468-1479. PMID: 29107293.

Montero-Conde, C., Leandro-Garcia, L. J., Chen, X., Oler, G., Ruiz-Llorente, S., Ryder, M., Landa, I., Sanchez-Vega, F., La, K., Ghossein, R. A., Bajorin, D. F., Knauf, J. A., Riordan, J. D., Dupuy, A. J. & Fagin, J. A. (2017). Transposon mutagenesis identifies chromatin modifiers cooperating with Ras in thyroid tumorigenesis and detects ATXN7 as a cancer gene. Proceedings of the National Academy of Sciences of the United States of America, 114(25), E4951-E4960. PMID: 28584132.

Suárez-Cabrera, C., Quintana, R. M., Bravo, A., Casanova, M. L., Page, A., Alameda, J. P., Paramio, J. M., Maroto, A., Salamanca, J., Dupuy, A. J., Ramírez, A. & Navarro, M. (2017). A transposon-based analysis reveals RASA1 is involved in triple negative breast cancer. Cancer Research, 77(6), 1357-1368. PMID: 28108518.

Sun, X., Yu, W., Sanz Navarro, M., Sweat, M., Eliason, S., Sharp, T., Liu, H., Seidel, K., Zhang, L., Moreno, M., Lynch, T., Holton, N. E., Rogers, L., Neff, T., Goodheart, M. J., Michon, F., Klein, O. D., Chai, Y., Dupuy, A. J., Engelhardt, J. F., Chen, Z. & Amendt, B. A. (2016). Sox2 and Lef-1 interact with Pitx2 to regulate incisor development and stem cell renewal. Development, 143(22), 4115-4126. PMID: 27660324.

Guimarães-Young, A., Neff, T., Dupuy, A. J. & Goodheart, M. J. (2016). Conditional deletion of Sox17 reveals complex effects on uterine adenogenesis and function. Developmental Biology, 414(2), 219-227. PMID: 27102016.

Morrissy, A. S., Garzia, L., Shih, D. J., Zuyderduyn, S., Huang, X., Skowron, P., Remke, M., Cavalli, F. M., Ramaswamy, V., Lindsay, P. E., Jelveh, S., Donovan, L. K., Wang, X., Luu, B., Zayne, K., Li, Y., Mayoh, C., Thiessen, N., Mercier, E., Mungall, K. L., Ma, Y., Tse, K., Zeng, T., Shumansky, K., Roth, A. J., Shah, S., Farooq, H., Kijima, N., Holgado, B. L., Lee, J. J., Matan-Lithwick, S., Liu, J., Mack, S. C., Manno, A., Michealraj, K. A., Nor, C., Peacock, J., Qin, L., Reimand, J., Rolider, A., Thompson, Y. Y., Wu, X., Pugh, T., Ally, A., Bilenky, M., Butterfield, Y. S., Carlsen, R., Cheng, Y., Chuah, E., Corbett, R. D., Dhalla, N., He, A., Lee, D., Li, H. I., Long, W., Mayo, M., Plettner, P., Qian, J. Q., Schein, J. E., Tam, A., Wong, T., Birol, I., Zhao, Y., Faria, C. C., Pimentel, J., Nunes, S., Shalaby, T., Grotzer, M., Pollack, I. F., Hamilton, R. L., Li, X. N., Bendel, A. E., Fults, D. W., Walter, A. W., Kumabe, T., Tominaga, T., Collins, V. P., Cho, Y. J., Hoffman, C., Lyden, D., Wisoff, J. H., Garvin, Jr, J. H., Stearns, D. S., Massimi, L., Schüller, U., Sterba, J., Zitterbart, K., Puget, S., Ayrault, O., Dunn, S. E., Tirapelli, D. P., Carlotti, C. G., Wheeler, H., Hallahan, A. R., Ingram, W., MacDonald, T. J., Olson, J. J., Van Meir, E. G., Lee, J. Y., Wang, K. C., Kim, S. K., Cho, B. K., Pietsch, T., Fleischhack, G., Tippelt, S., Ra, Y. S., Bailey, S., Lindsey, J. C., Clifford, S. C., Eberhart, C. G., Cooper, M. K., Packer, R. J., Massimino, M., Garre, M. L., Bartels, U., Tabori, U., Hawkins, C. E., Dirks, P., Bouffet, E., Rutka, J. T., Wechsler-Reya, R. J., Weiss, W. A., Collier, L. S., Dupuy, A. J., Korshunov, A., Jones, D. T., Kool, M., Northcott, P. A., Pfister, S. M., Largaespada, D. A., Mungall, A. J., Moore, R. A., Jabado, N., Bader, G. D., Jones, S. J., Malkin, D., Marra, M. A. & Taylor, M. D. (2016). Divergent clonal selection dominates medulloblastoma at recurrence. Nature, 529(7586), 351-357. PMID: 26760213.

Temiz, N. A., Moriarity, B. S., Wolf, N. K., Riordan, J. D., Dupuy, A. J., Largaespada, D. A. & Sarver, A. L. (2016). RNA sequencing of Sleeping Beauty transposon-induced tumors detects transposon-RNA fusions in forward genetic cancer screens. Genome Research, 26(1), 119-129. PMID: 26553456.

Smith, R. P., Riordan, J. D., Feddersen, C. R. & Dupuy, A. J. (2015). A Hybrid Adenoviral Vector System Achieves Efficient Long-Term Gene Expression in the Liver via piggyBac Transposition. Human Gene Therapy, 26(6), 377-385. PMID: 25808258.

Thiel, W. H., Thiel, K. W., Flenker, K. S., Bair, T., Dupuy, A. J., McNamara, 2nd, J. O., Miller, F. J. & Giangrande, P. H. (2015). Cell-internalization SELEX: method for identifying cell-internalizing RNA aptamers for delivering siRNAs to target cells. Methods in Molecular Biology, 1218, 187-199. PMID: 25319652.

Riordan, J. D., Drury, L. J., Smith, R. P., Brett, B. T., Rogers, L. M., Scheetz, T. E. & Dupuy, A. J. (2014). Sequencing methods and datasets to improve functional interpretation of sleeping beauty mutagenesis screens. BMC Genomics, 15, 1150. PMID: 25526783.

Zhang, Z., Kim, K., Li, X., Moreno, M., Sharp, T., Goodheart, M. J., Safe, S., Dupuy, A. J. & Amendt, B. A. (2014). MicroRNA-26b represses colon cancer cell proliferation by inhibiting lymphoid enhancer factor 1 expression. Molecular Cancer Therapeutics(American Association for Cancer Research), 13(7), 1942-1951. PMID: 24785257.

Gibson-Corley, K. N., Rogers, L. M., Goeken, A., Dupuy, A. J. & Meyerholz, D. K. (2014). Keratoacanthoma Pathobiology in Mouse Models. Diseases, 2(2), 106-119. PMID: 25309748.

Lim, L., Balakrishnan, A., Huskey, N., Jodari, M., Ng, R., Song, G., Cheung, S., Willenbring, H., Dupuy, A. J., Chen, X. & Goga, A. (2014). MiR-494 within an oncogenic microRNA megacluster regulates G1/S transition liver tumorigenesis through suppression of MCC. Hepatology, 59(1), 202-215. PMID: 23913442.

Bard-Chapeau, E. A., Nguyen, A. T., Rust, A. G., Sayadi, A., Lee, P., Chua, B. Q., New, L. S., de Jong, J., Ward, J. M., Chin, C. K., Chew, V., Toh, H. C., Abastado, J. P., Benoukraf, T., Soong, R., Bard, F. A., Dupuy, A. J., Johnson, R. L., Radda, G. K., Chan, E. C., Wessels, L. F., Adams, D. J., Jenkins, N. A. & Copeland, N. G. (2014). Transposon mutagenesis identifies genes driving hepatocellular carcinoma in a chronic hepatitis B mouse model. Nature Genetics, 46(1), 24-32. PMID: 24316982.

DeZwaan-McCabe, D., Riordon, J. D., Arensdorf, A. M., Chikka, M. R., Icardi, M. S., Dupuy, A. J. & Rutkowski, D. T. (2013). The stress-regulated transcription factor CHOP promotes hepatocellular oncogenesis. PLoS Genet, 9(12), e1003937. PMID: 24367269.

Riordan, J. D., Dupuy, A. J. (2013). Domesticated transposable element gene products in human cancer. Mobile Genetic Elements, 3(5), e26693. PMID: 24251072.

Rogers, L. M., Riordan, J. D., Swick, B. L., Meyerholz, D. K. & Dupuy, A. J. (2013). Ectopic expression of Zmiz1 induces cutaneous squamous cell malignancies in a mouse model of cancer. The Journal of Investigative Dermatology, 133(7), 1863-1869. PMID: 23426136.

Zanesi, N., Balatti, V., Riordan, J., Burch, A., Rizzotto, L., Palamarchuk, A., Cascione, L., Lagana, A., Dupuy, A. J., Croce, C. M. & Pekarsky, Y. (2013). A Sleeping Beauty screen reveals NF-kB activation in CLL mouse model. Blood, 121(21), 4355-4358. PMID: 23591791.

Rogers, L. M., Olivier, A. K., Meyerholz, D. K. & Dupuy, A. J. (2013). Adaptive immunity does not strongly suppress spontaneous tumors in a Sleeping Beauty model of cancer. Journal of Immunology, 190(8), 4393-4399. PMID: 23475219.

Riordan, J. D., Keng, V. W., Tschida, B. R., Scheetz, T. E., Bell, J. B., Podetz-Pedersen, K. M., Moser, C. D., Copeland, N. G., Jenkins, N. A., Roberts, L. R., Largaespada, D. A. & Dupuy, A. J. (2013). Identification of rtl1, a retrotransposon-derived imprinted gene, as a novel driver of hepatocarcinogenesis. PLoS Genetics, 9(4), e1003441. PMID: 23593033.

Quintana, R. M., Dupuy, A. J., Bravo, A., Casanova, M. L., Alameda, J. P., Page, A., Sánchez-Viera, M., Ramírez, A. & Navarro, M. (2013). A transposon-based analysis of gene mutations related to skin cancer development. The Journal of Investigative Dermatology, 133(1), 239-248. PMID: 22832494.

Keng, V. W., Sia, D., Sarver, A. L., Tschida, B. R., Fan, D., Alsinet, C., Solé, M., Lee, W. L., Kuka, T. P., Moriarity, B. S., Villanueva, A., Dupuy, A. J., Riordan, J. D., Bell, J. B., Silverstein, K. A., Llovet, J. M. & Largaespada, D. A. (2013). Sex bias occurrence of hepatocellular carcinoma in Poly7 molecular subclass is associated with EGFR. Hepatology, 57(1), 120-130. PMID: 22899566.

DeZwaan-McCabe, D., Riordan, J. D., Arensdorf, A. M., Icardi, M. S., Dupuy, A. J. & Rutkowski, D. T. (2013). The stress-regulated transcription factor CHOP promotes hepatic inflammatory gene expression, fibrosis, and oncogenesis. PLoS Genetics, 9(12), e1003937. PMID: 24367269.

Mann, K. M., Ward, J. M., Yew, C. C., Kovochich, A., Dawson, D. W., Black, M. A., Brett, B. T., Sheetz, T. E., Dupuy, A. J., Chang, D. K., Biankin, A. V., Waddell, N., Kassahn, K. S., Grimmond, S. M., Rust, A. G., Adams, D. J., Jenkins, N. A. & Copeland, N. G. (2012). Sleeping Beauty mutagenesis reveals cooperating mutations and pathways in pancreatic adenocarcinoma. Proceedings of the National Academy of Sciences of the United States of America, 109(16), 5934-5941. PMID: 22421440.

Wu, X., Northcott, P. A., Dubuc, A., Dupuy, A. J., Shih, D. J., Witt, H., Croul, S., Bouffet, E., Fults, D. W., Eberhart, C. G., Garzia, L., Van Meter, T., Zagzag, D., Jabado, N., Schwartzentruber, J., Majewski, J., Scheetz, T. E., Pfister, S. M., Korshunov, A., Li, X. N., Scherer, S. W., Cho, Y. J., Akagi, K., MacDonald, T. J., Koster, J., McCabe, M. G., Sarver, A. L., Collins, V. P., Weiss, W. A., Largaespada, D. A., Collier, L. S. & Taylor, M. D. (2012). Clonal selection drives genetic divergence of metastatic medulloblastoma. Nature, 482(7386), 529-33. PMID: 22343890.

Berquam-Vrieze, K. E., Swing, D. A., Tessarollo, L. & Dupuy, A. J. (2012). Characterization of transgenic mice expressing cancer-associated variants of human NOTCH1. Genesis, 50(2), 112-118. PMID: 21898766.

Berquam-Vrieze, K. E., Nannapaneni, K., Brett, B. T., Holmfeldt, L., Ma, J., Zagorodna, O., Jenkins, N. A., Copeland, N. G., Meyerholz, D. K., Knudson, C. M., Mullighan, C. G., Scheetz, T. E. & Dupuy, A. J. (2011). Cell of origin strongly influences genetic selection in a mouse model of T-ALL. Blood, 118(17), 4646-56. PMID: 21828136.

Thiel, W. H., Bair, T., Wyatt Thiel, K., Dassie, J. P., Rockey, W. M., Howell, C. A., Liu, X. Y., Dupuy, A. J., Huang, L., Owczarzy, R., Behlke, M. A., McNamara, J. O. & Giangrande, P. H. (2011). Nucleotide bias observed with a short SELEX RNA aptamer library. Nucleic acid therapeutics, 21(4), 253-63. PMID: 21793789.

Starr, T. K., Scott, P. M., Marsh, B. M., Zhao, L., Than, B. L., O'Sullivan, M. G., Sarver, A. L., Dupuy, A. J., Largaespada, D. A. & Cormier, R. T. (2011). A Sleeping Beauty transposon-mediated screen identifies murine susceptibility genes for adenomatous polyposis coli (Apc)-dependent intestinal tumorigenesis. Proceedings of the National Academy of Sciences of the United States of America, 108(14), 5765-70. PMID: 21436051.

Brett, B. T., Berquam-Vrieze, K. E., Nannapaneni, K., Huang, J., Scheetz, T. E. & Dupuy, A. J. (2011). Novel molecular and computational methods improve the accuracy of insertion site analysis in Sleeping Beauty-induced tumors. PloS one, 6(9), e24668. PMID: 21931803.

McGrail, M., Hatler, J. M., Kuang, X., Liao, H. K., Nannapaneni, K., Watt, K. E., Uhl, J. D., Largaespada, D. A., Vollbrecht, E., Scheetz, T. E., Dupuy, A. J., Hostetter, J. M. & Essner, J. J. (2011). Somatic mutagenesis with a Sleeping Beauty transposon system leads to solid tumor formation in zebrafish. PloS one, 6(4), e18826. PMID: 21533036.

Dupuy, A. J. (2010). Transposon-based screens for cancer gene discovery in mouse models. Seminars in cancer biology, 20(4), 261-8. PMID: 20478384.

Wang, D. H., Clemons, N. J., Miyashita, T., Dupuy, A. J., Zhang, W., Szczepny, A., Corcoran-Schwartz, I. M., Wilburn, D. L., Montgomery, E. A., Wang, J. S., Jenkins, N. A., Copeland, N. A., Harmon, J. W., Phillips, W. A. & Watkins, D. N. (2010). Aberrant epithelial-mesenchymal Hedgehog signaling characterizes Barrett's metaplasia. Gastroenterology, 138(5), 1810-22. PMID: 20138038.

Dupuy, A. J. (2010). Current applications of transposons in mouse genetics. Methods in enzymology, 477, 53-70. PMID: 20699136.

Collier, L. S., Adams, D. J., Hackett, C. S., Bendzick, L. E., Akagi, K., Davies, M. N., Diers, M. D., Rodriguez, F. J., Bender, A. M., Tieu, C., Matise, I., Dupuy, A. J., Copeland, N. G., Jenkins, N. A., Hodgson, J. G., Weiss, W. A., Jenkins, R. B. & Largaespada, D. A. (2009). Whole-body sleeping beauty mutagenesis can cause penetrant leukemia/lymphoma and rare high-grade glioma without associated embryonic lethality. Cancer research, 69(21), 8429-37. PMID: 19843846.

Dupuy, A. J., Rogers, L. M., Kim, J., Nannapaneni, K., Starr, T. K., Liu, P., Largaespada, D. A., Scheetz, T. E., Jenkins, N. A. & Copeland, N. G. (2009). A modified sleeping beauty transposon system that can be used to model a wide variety of human cancers in mice. Cancer research, 69(20), 8150-6. PMID: 19808965.

Starr, T. K., Allaei, R., Silverstein, K. A., Staggs, R. A., Sarver, A. L., Bergemann, T. L., Gupta, M., O'Sullivan, M. G., Matise, I., Dupuy, A. J., Collier, L. S., Powers, S., Oberg, A. L., Asmann, Y. W., Thibodeau, S. N., Tessarollo, L., Copeland, N. G., Jenkins, N. A., Cormier, R. T. & Largaespada, D. A. (2009). A transposon-based genetic screen in mice identifies genes altered in colorectal cancer. Science (New York, N.Y.), 323(5922), 1747-50. PMID: 19251594.

Keng, V. W., Villanueva, A., Chiang, D. Y., Dupuy, A. J., Ryan, B. J., Matise, I., Silverstein, K. A., Sarver, A., Starr, T. K., Akagi, K., Tessarollo, L., Collier, L. S., Powers, S., Lowe, S. W., Jenkins, N. A., Copeland, N. G., Llovet, J. M. & Largaespada, D. A. (2009). A conditional transposon-based insertional mutagenesis screen for genes associated with mouse hepatocellular carcinoma. Nature biotechnology, 27(3), 264-74. PMID: 19234449.

Su, Q., Prosser, H. M., Campos, L. S., Ortiz, M., Nakamura, T., Warren, M., Dupuy, A. J., Jenkins, N. A., Copeland, N. G., Bradley, A. & Liu, P. (2008). A DNA transposon-based approach to validate oncogenic mutations in the mouse. Proceedings of the National Academy of Sciences of the United States of America, 105(50), 19904-9. PMID: 19064922.

Dupuy, A. J., Jenkins, N. A. & Copeland, N. G. (2006). Sleeping beauty: a novel cancer gene discovery tool. Human molecular genetics, 15 Spec No 1, R75-9. PMID: 16651372.

Collier, L. S., Carlson, C. M., Ravimohan, S., Dupuy, A. J. & Largaespada, D. A. (2005). Cancer gene discovery in solid tumours using transposon-based somatic mutagenesis in the mouse. Nature, 436(7048), 272-6. PMID: 16015333.

Dupuy, A. J., Akagi, K., Largaespada, D. A., Copeland, N. G. & Jenkins, N. A. (2005). Mammalian mutagenesis using a highly mobile somatic Sleeping Beauty transposon system. Nature, 436(7048), 221-6. PMID: 16015321.

Belur, L. R., Frandsen, J. L., Dupuy, A. J., Ingbar, D. H., Largaespada, D. A., Hackett, P. B. & Scott McIvor, R. (2003). Gene insertion and long-term expression in lung mediated by the Sleeping Beauty transposon system. Molecular therapy : the journal of the American Society of Gene Therapy, 8(3), 501-7. PMID: 12946324.

Carlson, C. M., Dupuy, A. J., Fritz, S., Roberg-Perez, K. J., Fletcher, C. F. & Largaespada, D. A. (2003). Transposon mutagenesis of the mouse germline. Genetics, 165(1), 243-56. PMID: 14504232.

Geurts, A. M., Yang, Y., Clark, K. J., Liu, G., Cui, Z., Dupuy, A. J., Bell, J. B., Largaespada, D. A. & Hackett, P. B. (2003). Gene transfer into genomes of human cells by the sleeping beauty transposon system. Molecular therapy : the journal of the American Society of Gene Therapy, 8(1), 108-17. PMID: 12842434.

Dupuy, A. J., Clark, K., Carlson, C. M., Fritz, S., Davidson, A. E., Markley, K. M., Finley, K., Fletcher, C. F., Ekker, S. C., Hackett, P. B., Horn, S. & Largaespada, D. A. (2002). Mammalian germ-line transgenesis by transposition. Proceedings of the National Academy of Sciences of the United States of America, 99(7), 4495-9. PMID: 11904379.

Dupuy, A. J., Fritz, S. & Largaespada, D. A. (2001). Transposition and gene disruption in the male germline of the mouse. Genesis (New York, N.Y. : 2000), 30(2), 82-8. PMID: 11416868.

Dupuy, A. J., Morgan, K., von Lintig, F. C., Shen, H., Acar, H., Hasz, D. E., Jenkins, N. A., Copeland, N. G., Boss, G. R. & Largaespada, D. A. (2001). Activation of the Rap1 guanine nucleotide exchange gene, CalDAG-GEF I, in BXH-2 murine myeloid leukemia. The Journal of biological chemistry, 276(15), 11804-11. PMID: 11278453.

Li, J., Shen, H., Himmel, K. L., Dupuy, A. J., Largaespada, D. A., Nakamura, T., Shaughnessy, Jr, J. D., Jenkins, N. A. & Copeland, N. G. (1999). Leukaemia disease genes: large-scale cloning and pathway predictions. Nature genetics, 23(3), 348-53. PMID: 10610183.