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1 pp. ES ips zonation ES ips in vitro Tissue morphogenesis of liver epithelial cells Naoki Tanimizu and Toshihiro Mitaka Department of Tissue Development and Regeneration, Research Institute for Frontier Medicine, Sapporo Medical University School of Medicine, South, West, Chuo-ku, Sapporo, Japan in vitro tight junction adherence junction apical basolateral

2 ph Disse Cre Cre-ERT in vitro

3 zonation ERM ezrin-radixinmoesin radixin MRP multidrug resistance protein, abcc LKB Cre Alb- Cre LKB flox flox LKB BSEP bile salt export pump, abcb LKB radixin radixin Wnt β- Neonatal ichthyosis and sclerosing cholangitis NISCH Claudin Cldn ZO occludin in vitro M OSM OSM K-Ras A Matrigel A HepG A A SMP senescence marker protein- A WIFB WIFBPar b MARK microtubule association related kinase EMK MDCK Par b A Taurocholate TC TC camp EPAC LKB AMPK B Matrigel TC FDA fluorescent diacetate

4 ZO FDA fluorescein diacetate Matrigel HNF α E zonation zonation I Carbamoylphosphate synthetase I, CPSI Phosphoenolpyruvate carboxykinase, PEPCK Glutamine synthetase, GS

5 zonation Wnt β- β- APC Wnt β- zonation zonation Wnt β- β- GS Glut- Wnt adenomatous polyposis coli APC CPSI GS Wnt β- zonation Wnt Notch NICD Notch Notch Notch HNF α HNF β Notch TGFβ tubulogenesis branching morphogenesis ductal plate ductal plate Antoniou Sox

6 Sox HNF α Sox HNF α Zong Notch Antoniou TGFβ Notch lineage tracing ductal plate ductal plate Sox HPPL HPPL HPPL I Matrigel Akt HNF β HPPL Matrigel cyst cyst HPPL Matrigel α β γ HPPL β F α CK α α HPPL cyst HPPL Grhl grainyhead-like Grhl Cldn Cldn Rab Rab Cldn Grhl Cldn Cldn -like b Cldn lb

in vitro Grhl Grhl grainyhead like HPPL Grhl in vivo in vitro in vivo in vitro ips ES Kikuchi, S., Hata, M., Fukumoto, K., Yamane, Y., Matsui, T., Tamura, A., Yonemura, S., Yamagishi, H., Keppler, D.

7 in vitro Grhl Grhl grainyhead like HPPL Grhl in vivo in vitro in vivo in vitro ips ES Kikuchi, S., Hata, M., Fukumoto, K., Yamane, Y., Matsui, T., Tamura, A., Yonemura, S., Yamagishi, H., Keppler, D., & Tsukita, S.Nat. Genet.,,. Woods, A., Heslegrave, A.J., Muckett, P.J., Levene, A.P., Clements, M., Mobberley, M., Ryder, T.A., Abu-Hayyeh, S., Williamson, C., Goldin, R.D., Ashworth, A., Withers, D.J., & Carling, D.Biochem. J.,,. Yeh, T.H., Krauland, L., Singh, V., Zou, B., Devaraj, P., Stolz, D.B., Franks, J., Monga, S.P., Sasatomi, E., & Behari, J. Hepatology,,. Grosse, B., Cassio, D., Yousef, N., Bernardo, C., Jacquemin, E., & Gonzales, E.Hepatology,,. Matsui, T., Kinoshita, T., Morikawa, Y., Tohya, K., Katsuki, M., Ito, Y., Kamiya, A., & Miyajima, A.EMBO J.,,. Ishigami, A., Fujita, T., Inoue, H., Handa, S., Kubo, S., Kondo, Y.,&Maruyama,N.Cell Tissue Res.,,. Cohen, D., Brennwald, P.J., Rodriguez-Boulan, E., & Musch, A.J. Cell Biol.,,. Fu, D., Wakabayashi, Y., Lippincott-Schwartz, J., & Arias, I. M.Proc. Natl. Acad. Sci. USA,,. Mitaka, T., Sato, F., Mizuguchi, T., Yokono, T., & Mochizuki, Y.Hepatology,,. Kasuya, J., Sudo, R., Mitaka, T., Ikeda, M., & Tanishita, K. Tissue Eng. Part A,,. Parviz, F., Matullo, C., Garrison, W.D., Savatski, L., Adamson, J.W., Ning, G., Kaestner, K.H., Rossi, J.M., Zaret, K.S., & Duncan, S.A.Nat. Genet.,,. Sakaguchi, T.F., Sadler, K.C., Crosnier, C., & Stainier, D.Y. Curr. Biol.,,. Sekine, S., Lan, B.Y., Bedolli, M., Feng, S., & Hebrok, M. Hepatology,,. Benhamouche, S., Decaens, T., Godard, C., Chambrey, R., Rickman, D.S., Moinard, C., Vasseur-Cognet, M., Kuo, C.J., Kahn, A., Perret, C., & Colnot, S.Dev. Cell,,. Zong, Y., Panikkar, A., Xu, J., Antoniou, A., Raynaud, P., Lemaigre, F., & Stanger, B.Z.Development,,. Tanimizu,N.&Miyajima,A.J. Cell Sci.,,. Antoniou, A., Raynaud, P., Cordi, S., Zong, Y., Tronche, F., Stanger, B.Z., Jacquemin, P., Pierreux, C.E., Clotman, F., & Lemaigre, F.P.Gastroenterology,,. Carpentier, R., Suner, R.E., van Hul, N., Kopp, J.L., Beaudry, J.B., Cordi, S., Antoniou, A., Raynaud, P., Lepreux, S., Jacquemin, P., Leclercq, I.A., Sander, M., & Lemaigre, F.P. Gastroenterology,,. Tanimizu, N., Saito, H., Mostov, K., & Miyajima, A.J.

8 Cell Sci.,,. Tanimizu, N., Miyajima, A., & Mostov, K.E.Mol. Biol. Cell,,. Tanimizu, N., Miyajima, A., & Mostov, K.E.Mol. Biol. Cell,,. Tanimizu, N., Kikkawa, Y., Mitaka, T., & Miyajima, A. J. Biol. Chem., in press. Senga, K., Mostov, K.E., Mitaka, T., Miyajima, A., & Tanimizu, N.Mol. Biol. Cell, in press. Cheung, I.D., Bagnat, M., Ma, T.P., Datta, A., Evason, K., Moore, J.C., Lawson, N.D., Mostov, K.E., Moens, C.B., & Stainier, D.Y.Dev. Biol.,,. Eiraku, M., Takata, N., Ishibashi, H., Kawada, M., Sakakura, E., Okuda, S., Sekiguchi, K., Adachi, T., & Sasai, Y. Nature,,. Sekiya, S. & Suzuki, A.Nature,,.