Lu, Joyce Jean

JLu6x7Associate Professor

[ CV ]




  • Associate Professor, Stem Cell Program, Genomics Research Center, Academia Sinica, Taiwan. 2015-present
  • Assistant Professor, Stem Cell Program, Genomics Research Center, Academia Sinica, Taiwan. 2007-2015
  • Adjunct Assistant Professor, Genomics and System Biology Program, College of Life Science, National Taiwan University, Taiwan. 2010-present
  • Co-PI, National RNAicore Facility Plateform, 2009-present
  • Postdoctoral Fellow/Associate, Molecular, Department of Cellular, and Developmental Biology, Yale University, USA. 2003-2007
  • Postdoctoral Fellow, Graduate Institute of Microbiology, College of Medicine, National Taiwan University, Taiwan. 2001-2003
  • Ph.D. Institute of Microbiology, National Taiwan University, Taipei, Taiwan. 1995-2000
  • M.S. Institute of Molecular Medicine, National Taiwan University, Taipei, Taiwan. 1992-1994
  • B.S. Department of Medical Technology, National Taiwan University, Taipei, Taiwan. 1988-1992
  • Research Assistant, Graduate Institute of Microbiology, College of Medicine, National Taiwan University, Taiwan. 1994-1995


  • Ruth L. Kirschstein National Research Service Award, 2003-2004, NIH
  • Outstanding Paper Award, National Taiwan University, 2002


High throughput functional screens pinpoints key factors for cell fate determination

Lu-scheme  Research Figure/Table/Scheme

Stem cell renewal and differentiation is important for stem cell biology and regenerative medicine. To efficiently pinpoint factors critical for cell fate determination, systematically functional screens were performed in mouse embryonic stem cells (ESCs), human ESCs, and mesenchymal stem cells (MSCs). We established the first shRNA screen in ESCs. With a shRNA library of kinases and phosphatases contains 4796 shRNAs, we identified 244 genes essential for ESC renewal. Among them, Nme6 and Nme7 is essential for 8 stemness genes expressions. With 517 shRNAs target genes are differentially expressed in undifferentiated human ESCs and differentiated cells or are embryonic lethal, we performed a shRNA functional screen and found 86 genes essential for human ESC renewal. And two candidate genes either bridge the oxidation and reduction status of ESCs with stemness signals, or block neural stem cell differentiation are currently studied in-depth. To fulfill the unmet need of bone diseases, a gain-of-function screen was performed with an overexpression library contains 12380 genes. 9 candidate genes are found to be essential and sufficient to promote the osteogenesis of human MSCs. By this method, we found soluble factor that can completely treat and prevent osteoporosis. Thus we found many factors play critical roles for ESC and MSC renewal and differentiation by systematically functional screens.


  • Yang SC, Liu JJ, Wang CK, Lin YT, Tsai SY, Chen WJ, Huang WK, Tu PA, Lin YC, Chang CF, Cheng CL, Lin H, Lai CY, Lin CY, Lee YH, Chiu YC, Hsu CC, Hsu SC, Hsiao M, Schuyler SC, Lu FL*, Lu J*, to appear, “Down-regulation of ATF1 leads to early neuroectoderm differentiation of human embryonic stem cells by increasing the expression level of SOX2.”, The FASEB Journal: official publication of the Federation of American Societies for Experimental Biology, 33(9),10577-10592. (SCIE)
  • Wang, C.H., Ma, N., Lin, Y.T., Wu, C.C., Wu, H. J., Yu, C.C., Hsiao, M., Lu, F.L., Schuyler, S.C., and Lu, J*., accepted, Current Protocols in Stem Cell Biology, 26: 5C.3.1 - 5C.3.x. pages, Hoboken, NJ,USA: Wiley-Blackwell.
  • Lai YP, Kuo LC, Lin BR, Lin HJ, Lin CY, Chen YT, Hsiao PW, Chang HT, Ko PC, Chen HC, Chang HY, Lu J, Ho HN, Wu-Hsieh BA, Kung JT, Chen SC, 2021, “CD28 engagement inhibits CD73-mediated regulatory activity of CD8(+) T cells.”, Communications biology, 4(1), 595. (SCIE)
  • Cheng Chih-Lun, Yang Shang-Chih, Lai Chien-Ying, Wang Cheng-Kai, Chang Ching-Fang, Lin Chun-Yu, Chen Wei-Ju, Lin Po-Yu, Wu Han-Chung, Ma Nianhan, Lu Frank Leigh*, Lu Jean*, 2020, “CXCL14 Maintains hESC Self-Renewal through Binding to IGF-1R and Activation of the IGF-1R Pathway”, Cells, 9(7), 1706. (SCIE)
  • Nguyen Mai-Huong Thi, Lin Chen-Huan, Liu Szu-Mam, Miyashita Azusa, Ihn Hironobu, Lin Hsuan, Ng Chi Hou, Tsai Jen-Chieh, Chen Ming-Hong, Tsai Mu-Shiun, Lin In-Yu, Liu Shu-Chen, Li Long-Yuan, Fukushima Satoshi*, Lu Jean*, Ma Nianhan*, 2020, “miR-524-5p reduces the progression of the BRAF inhibitor-resistant melanoma”, Neoplasia, 22(12), 789-799. (SCIE)
  • Lai Pei‑Lun, Chen Ting‑Chun, Feng Chun‑Yen, Lin Hsuan, Ng Chi‑Hou, Chen Yun, Hsiao Michael, Lu Jean*, Huang Hsiao‑Chun*, 2020, “Selection of a malignant subpopulation from a colorectal cancer cell line”, Oncology Letters, 20(3), 2937-2945. (SCIE)
  • Kuan II, Lee CC, Chen CH, Lu J, Kuo YS, Wu HC, 2019, “The extracellular domain of epithelial cell adhesion molecule (EpCAM) enhances multipotency of mesenchymal stem cells through EGFR-LIN28-LET7 signaling.”, The Journal of biological chemistry, 294(19), 7769-7786. (SCIE)
  • Chang HC, Huang PH, Syu FS, Hsieh CH, Chang SL, Lu J*, Chen HC*, 2018, “Critical involvement of atypical chemokine receptor CXCR7 in allergic airway inflammation.”, Immunology, 154(2), 274-284. (SCIE)
  • Liu SM, Lin CH, Lu J, Lin IY, Tsai MS, Chen MH, Ma N, 2018, “miR-596 Modulates Melanoma Growth by Regulating Cell Survival and Death.”, The Journal of investigative dermatology, 138(4), 911-921. (SCIE)
  • Wang CK, Yang SC, Hsu SC, Chang FP, Lin YT, Chen SF, Cheng CL, Hsiao M, Lu FL, Lu J*, 2017, “CHAC2 is essential for self-renewal and glutathione maintenance in human embryonic stem cells.”, Free radical biology & medicine, 113, 439-451. (SCIE)
  • Lai PL, Lin H, Chen SF, Yang SC, Hung KH, Chang CF, Chang HY, Lu FL, Lee YH, Liu YC, Huang HC*, Lu J*, 2017, “Efficient Generation of Chemically Induced Mesenchymal Stem Cells from Human Dermal Fibroblasts.”, Scientific reports, 7, 44534. (SCIE)
  • Lin YT, Wang CK, Yang SC, Hsu SC, Lin H, Chang FP, Kuo TC, Shen CN, Chiang PM, Hsiao M, Lu FL, Lu J*, 2017, “Elimination of undifferentiated human embryonic stem cells by cardiac glycosides.”, Scientific reports, 7(1), 5289. (SCIE)
  • Kuan II, Liang KH, Wang YP, Kuo TW, Meir YJ, Wu SC, Yang SC, Lu J*, Wu HC*, 2017, “EpEX/EpCAM and Oct4 or Klf4 alone are sufficient to generate induced pluripotent stem cells through STAT3 and HIF2alpha.”, Scientific reports, 7, 41852. (SCIE)
  • Hsieh MH, Chen YT, Chen YT, Lee YH, Lu J, Chien CL, Chen HF, Ho HN, Yu CJ, Wang ZQ, Teng SC, 2017, “PARP1 controls KLF4-mediated telomerase expression in stem cells and cancer cells.”, Nucleic acids research, 45(18), 10492-10503. (SCIE)
  • Chang WF, Hwu YM, Xu J, Lin CJ, Wang SW, Cheng AS, Lu J, Lu CH, Sung LY, 2016, “Derivation of Patient Specific Pluripotent Stem Cells Using Clinically Discarded Cumulus Cells.”, PloS one, 11(11), e0165715. (SCIE)
  • Huang YC, Lin SJ, Lin KM, Chou YC, Lin CW, Yu SC, Chen CL, Shen TL, Chen CK,Lu J, Chen MR, Tsai CH, 2016, “Regulation of EBV LMP1-triggered EphA4 downregulation in EBV-associated B lymphoma and its impact on patients\' survival.”, Blood, 128(12), 1578-89. (SCIE)
  • Chang JS, Su CY, Yu WH, Lee WJ, Liu YP, Lai TC, Jan YH, Yang YF, Shen CN, Shew JY, Lu J, Yang CJ, Huang MS, Lu PJ, Lin YF, Kuo ML, Hua KT, Hsiao M, 2015, “GIT1 promotes lung cancer cell metastasis through modulating Rac1/Cdc42 activity and is associated with poor prognosis.”, Oncotarget, 6(34), 36278-91.
  • Chua HH, Tsuei DJ, Lee PH, Jeng YM, Lu J, Wu JF, Su DS, Chen YH, Chien CS, Kao PC, Lee CN, Hu RH, Ni YH, Chang MH, 2015, “RBMY, a novel inhibitor of glycogen synthase kinase 3beta, increases tumor stemness and predicts poor prognosis of hepatocellular carcinoma.”, Hepatology, 62(5), 1480-96. (SCIE)
  • Wu, C. C., Wu, H.C., Chen, W.J, Wang, C.H, Lin, C.H., Hsu, S.C., Chen, Y.R., Hsiao, M., Schuyler, S.C., Lu, F. L. , Ma, N.H., and Lu, J.*., 2015, “Suppression of Dual leucine zipper-bearing Kinase activity by Akt is required for mouse embryonic stem cell self-renewal”, CELL CYCLE, 14(8), 1207-1217. (SCIE)
  • Chang, C.F., Hsu, K.H., Shen, C.N., Li, C.L.*, and Lu, J.*, 2014, “Enrichment and characterization of two subgroups of committed osteogenic cells in the mouse endosteal bone marrow with expression levels of CD24”, Journal of Bone Research, 2, 144.
  • Huang HN, Chen SY, Hwang SM, Su MW, Mai W, Wang HW, Cheng WC, Schuyler SC, Ma N, Lu FL, Lu, J.*, 2014, “miR-200c and GATA binding protein 4 regulate human embryonic stem cell renewal and differentiation”, Stem Cell Research, 12(2), 338-353. (SCIE)
  • Liu YC, Kao YT, Huang WK, Lin KY, Wu SC, Hsu SC, Schuyler SC, Li LY, Leigh Lu F, Lu J*, 2014, “CCL5/RANTES is important for inducing osteogenesis of human mesenchymal stem cells and is regulated by dexamethasone.”, Bioscience trends, 8(3), 138-143. (SCIE)
  • Liu SM, Lu J, Lee HC, Chung FH, Ma N, 2014, “miR-524-5p suppresses the growth of oncogenic BRAF melanoma by targeting BRAF and ERK2.”, Oncotarget, 5(19), 9444-9459.
  • Yang YF, Jan YH, Liu YP, Yang CJ, Su CY, Chang YC, Lai TC, Chiou J, Tsai HY, Lu J, Shen CN, Shew JY, Lu PJ, Lin YF, Huang MS, Hsiao M, 2014, “Squalene synthase induces tumor necrosis factor receptor 1 enrichment in lipid rafts to promote lung cancer metastasis.”, American journal of respiratory and critical care medicine, 190(6), 675-687. (SCIE)
  • Lin SJ, Lo M, Kuo RL, Shih SR, Ojcius DM, Lu J, Lee CK, Chen HC, Lin MY, Leu CM, Lin CN, Tsai CH, 2014, “The pathological effects of CCR2+ inflammatory monocytes are amplified by an IFNAR1-triggered chemokine feedback loop in highly pathogenic influenza infection.”, Journal of biomedical science, 21, 99-117. (SCIE)
  • Liu YP, Yang CJ, Huang MS, Yeh CT, Wu AT, Lee YC, Lai TC, Lee CH, Hsiao YW, Lu J, Shen CN, Lu PJ, Hsiao M, 2013, “Cisplatin Selects for Multidrug-Resistant CD133+ Cells in Lung Adenocarcinoma by Activating Notch Signaling.”, Cancer Research, 73(1), 406-416. (SCIE)
  • Chen HW, Chen HY, Wang LT, Wang FH, Fang LW, Lai HY, Chen HH, Lu J, Hung MS, Cheng Y, Chen MY, Liu SJ, Chong P, Lee OK, Hsu SC, 2013, “Mesenchymal stem cells tune the development of monocyte-derived dendritic cells toward a myeloid-derived suppressive phenotype through growth-regulated oncogene chemokines.”, Journal of Immunology, 190(10), 5065-5077. (SCIE)
  • Lu FL, Yu CC, Chiu HH, Liu HE, Chen SY, Lin S, Goh TY, Hsu HC, Chien CH, Wu HC, Chen MS, Schuyler SC, Hsieh WS, Wu MH, Lu J*, 2013, “Sonic hedgehog antagonists induce cell death in acute myeloid leukemia cells with the presence of lipopolysaccharides, tumor necrosis factor-alpha, or interferons.”, Investigational New Drugs, 31(4), 823-832. (SCIE)
  • Wang CH, Ma N, Lin YT, Wu CC, Wu HJ, Yu CC, Hsiao M, Lu FL, Schuyler SC, and Lu, J*, 2013, “UNIT 5C.3 Array-based high-throughput screening in mouse embryonic stem cells with shRNAs”, Current Protocols in Stem Cell Biology, 26: 5C.3.1-5C. 3.19.
  • Wang, C. H., Ma, N. H., Lin, Y.T., Wu, C.C., Hsiao M., Lu, F. L., Yu, C.C., Chen, S.Y., and Lu, J.*, 2012, “A shRNA functional screening in embryonic stem cells reveals Nme6 and Nme7 signaling are crucial for stem cell renewal”, Stem Cells, 30(10), 2199-2211. (SCIE)
  • Chou YC, Chen CL, Yeh TH, Lin SJ, Chen MR, Doong SL, Lu J, Tsai CH, 2012, “Involvement of recepteur d'origine nantais receptor tyrosine kinase in Epstein-Barr virus-associated nasopharyngeal carcinoma and its metastasis.”, The American Journal of Pathology, 181(5), 1773-1781. (SCIE)
  • Chan CC, Cheng LY, Lu J, Huang YH, Chiou SH, Tsai PH, Huo TI, Lin HC, Lee FY, 2012, “The Role of Interferon-gamma Inducible Protein-10 in a Mouse Model of Acute Liver Injury Post Induced Pluripotent Stem Cells Transplantation.”, PLoS One, 7(12), e50577. (SCIE)
  • Chou YC, Lin SJ,Lu J, Yeh TH, Chen CL, Weng PL, Lin JH, Yao M, Tsai CH, 2011, “ Requirement for LMP1-induced RON receptor tyrosine kinase in Epstein-Barr virus-mediated B-cell proliferation”, BLOOD, 118(5), 1340-1349. (SCIE)
  • Lu J, Hou R, Booth C J, Yang SH and Snyder M, 2006, “Defined culture conditions of human embryonic stem cells”, Proceedings of the national academy of sciences of the united states of america, 103, 5688-5693. (SCIE)
  • Lu J, Lin WS, Chen SY , Longnecker R, and Tsai CH, 2006, “Epstein-Barr virus latent membrane protein 2A enhances cell mobility in epithelial cells through syk kinase”, J Biol Chem, 281, 8806-8814. (SCIE)
  • Chang Y, Lee HH , Chen YT, Lu J, Wu SY ,Chen CW, Takada K, and Tsai CH , 2006, “Induction of the early growth response 1 gene by Epstein-Barr virus lytic transactivator Zta”, J Virol, 80, 7748-7755. (SCIE)
  • Lu J, Chua HH, Chen SY, Chen JY and Tsai CH, 2003, “Regulation of matrix metalloproteinase-1 by Epstein-Barr virus proteins”, Cancer Res, 63, 256-262. (SCIE)
  • ChenSY, Lu J, Shih YC and Tsai CH, 2002, “Epstein-Barr virus latent membrane protein 2A regulates c-Jun protein through extracellular signal-regulated kinase”, J Virol, 76, 9556-9561. (SCIE)
  • Lu J, Chen SY, Chua HH, Liu YS, Huang YT, Chang Y, Chen JY, Sheen TS, and Tsai CH, 2000, “Upregulation of tyrosine kinase TKT by the Epstein-Barr virus transactivator Zta”, J Virol, 74, 7391-7399. (SCIE)
  • Huang YT, Sheen TS, Chen CL, Lu J, Chang Y, Chen JY, and Tsai CH, 1999, “Profile of cytokine expression in nasopharyngeal carcinomas: a distinct expression of interleukin 1 in tumor and CD4+ T cells”, Cancer Res, 59, 1599-1605. (SCIE)
  • Chang Y, Tung CH, Huang YT, Lu J, Chen JY, and Tsai CH, 1999, “Requirement for cell-to-cell contact in Epstein-Barr virus infection of nasopharyngeal carcinoma cells and keratinocytes”, J Virol, 73, 8857-8866. (SCIE)
  • Chang Y, Sheen TS, Lu J , Huang YT, Chen JY,Yang CS, and Tsai CH, 1998, “Detection of transcripts initiated from two viral promoters (Cp and Wp) in Epstein-Barr virus-infected nasopharyngeal carcinoma cells and biopsies”, Lab Invest, 78, 715-726. (SCIE)
  • Tsai CH, Liu MT, Chen MR,Lu J, Yang HL,Chen JY and Yang CS, 1997, “Characterization of monoclonal antibodies to the Zta and DNase proteins of Epstein-Barr virus”, J Biomed Sci, 4, 69-77. (SCIE)