Vol. 35, pp , 2007 : embryonic stem, ES. Mash1. in vitro. in vitro. functional MRI

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1 143 Vol. 35, pp , : embryonic stem, ES ES in vitro in vitro 1 2 ES 1 7 ES Mash functional MRI 3 17

ªº»6¼ 144 7%8% o 1 ES! 2 2 g ) ; OP9! 6v*+()! { A, B 2 2 g ) 4 g 5ῌ *+, ( -. /E! =L[ ( ª0«*+y 1 {!

64 µh( I {"5 5Q) ῌD ῌ ῐb OP9! ¹ 5ῌ ES ῌ ES! "#$% & IGFII ' u! 6 IJ"! Bvw x! & y Iz!

- *, 345 I {"B I{" ῌvasculogenesis IZ 6 7 8()9: ;< = [ ῌangiogenesis a( {" } I " >?

$/(), 012 LM5NO ;< PQ =& ῌ RS 012!" F y Iz! = H() TU Mash1! # =7oN! B uz! 6E 34()GH VW XY B IZ[=\E <!$

2 ªº»6¼ 144 7%8% o 1 ES! 2 2 g ) ; OP9! 6v*+()! { A, B 2 2 g ) 4 g 5ῌ *+, ( -. /E! =L[ ( ª0«*+y 1 {! = N 20«ῌ C, D 2 2 g ) 4 OP9! 6v*+ ( 24 K g { 5ῌ 1 {! 20«/E! = < } ª0«{ g=±²³on ῌC ῌ F /E! B3E g /E! 64 µh( I {"5 5Q) ῌD ῌ ῐb OP9! ¹ 5ῌ ES ῌ ES! "#$% & IGFII ' u! 6 IJ"! Bvw x! & y Iz! ῌhemangioblast qo r5 8ῑ10 ῌ % ( ()*+,! - *, 345 I {"B I{" ῌvasculogenesis IZ 6 7 8()9: ;< = [ ῌangiogenesis a( {" } I " >?@5 = AB< CDB ( ~6 01 ῌarteriogenesis 3 E 5 ῌ F GH IJ"! K 34( on ῌ B [ 17 ῌ (./(), 012 LM5NO ;< PQ =& ῌ RS 012!" F y Iz! = H() TU Mash1! # =7oN! B uz! 6E 34()GH VW XY B IZ[=\E <! = I! 6E ῌ F I ]?@ ^ F6=_ 4 ῌ! =$H( I {"5 = ` NO IA I! a b I{" 6E ῌ a IZ[ cde B Ofghig Ajklῌk, B%zAῌ ;<;< Q!qE I&={" mn o B ped B ES! qo r N 01 B ῌ ' () I! a =s t & "~ I {"= N ῌ F ;c I! ῌ B I%[ qo ( E5! & 18

$/01ª9«2 3= 6:;"# 145 # 2 k l 2 3 ): /^ ^6 A, B Z[5\5]^1 4 yz ῌA 2 3>6 OP9 & cd 24 ῌB 6 ]$ $CD31 ῌC & VE-cadherin ῌD 6 ῌ VE-cadherin ] ^ 6ῌ ) < b? h $ ῌ ῌ i /01 ES Z[5\5]^1j @c!$ $/01 ES 0 8ῐZ[5\5]^ - <= 6"# >?@ 1 tucv!cd wx$ & 3 yz{ ῌ A!6 B'CD -@ 13ῒ15 ῌ E }96 '~?$ I6 & JK5LM NO?! b6 ':;??@h 6', PQR S TU VWXY-@ ) ῌ - %-' f :;?

I6 & JK5LM NO?! b6 ':;??@h 6', PQR S TU VWXY-@ ) ῌ - %-' f :;?

3 /01ª9«2 3= 6:;"# 145 # 2 k l 2 3 ): /^ ^6 A, B Z[5\5]^1 4 yz ῌA 2 3>6 OP9 & cd 24 ῌB 6 ] ^ k l 6/^ ^! Flk-1 6 ῌ C, D OP9 & cd 24 6 ] ^ 6/^ ^! CD31 ῌC & VE-cadherin ῌD 6 ῌ VE-cadherin ] ^ 6ῌ ) < b? h $ ῌ ῌ i /01 ES d$ %& k l 2 3 #$ %&'(&) &*+,- ῌ. /012 6"# mῌ n:; op?@qrsp 3456 ES 7, Flk-1 89 ':; cd?/01 ES 0 8ῐZ[5\5]^ - <= 6"# >?@ 1 tucv!cd wx$ & 3 yz{ ῌ A!6 B'CD -@ 13ῒ15 ῌ E }96 '~? 4 yz 1ῒ2 6 7, FG ES 7,H! ῌ= ) ^G 6 ] ^ b $ # 1 A, B ῌ 6"#'CD -@ 16ῒ17 ῌ?7? ES cvje %6 } 6 b 7,"#?I6 & JK5LM NO?! b6 ':;??@h 6', PQR S TU VWXY-@ ) ῌ - %-' f :;? &*+,- 7 ῌ %- opῑop /01 6 1G]^/ ES! OP9! 24 cd$ & ]!" ^6 6 'ῌ ;$ b Z[5\5]^1 ῌMC _ `ab;cde 6 ῌ ;? ' u ) ] ^6 f6 6"# gh_,-@ ῌ b ',- # 1C ῌ FG! 17 y 19

4 146 1 mrna 1D RT-PCR Flk-1 18 ES OP9 1 ES CD34 ES OP9 VE-cadherin Flk-1 CD31 ES nicotinic acethylcholine receptor, nachr a a b g e d 5 a7 nachr 19 nachr a1 7 b1 4 a2 b1 OP9 Est-1 20 Vezf-1 21 Hex 22 Flk-1 Est-1 OP9 erythropoietin producing human hepatocellular carcinoma Eph Ephrin Eph-receptor-interacting protein EphrinB2 23 EphB4 20

5 EphrinB2 EphB 4 Tie2 ES OP9 PDGF PDGF B OP9 asma OP9 Flk-1 OP9 VE-cadherin CD31 2 OP9 3 ES ES ES 1 Chiba S, Ikeda R, Kurokawa MS, Yoshikawa H, Takeno M, Nagafuchi H, Tadokoro M, Sekino H, Hashimoto T and Suzuki N. Anatomical and functional recovery by embryonic stem cell-derived neural tissue of a mouse model of brain damage. J Neurol Sci 2004; 219: Ikeda R, Kurokawa MS, Chiba S, Yoshikawa H, Hashimoto T, Tadokoro M and Suzuki N. Transplantation of motoneurons derived from MASH1-transfected mouse ES cells reconstitutes neural networks and improves motor function in hemiplegic mice. Exp neurol 2004; 189: Ikeda R, Kurokawa MS, Chiba S, Yoshikawa H, Ide M, Tadokoro M, Nito S, Nakatsuji N, Kondoh Y, Nagata K, Hashimoto T and Suzuki N. Transplantation of neural cells derived from retinoic acid-treated cynomolgus monkey embryonic stem cells successfully improved motor function of hemiplegic mice with experimental brain injury. Neurobiol Dis 2005; 20: Hamada M, Yoshikawa H, Ueda Y, Kurokawa MS, Watanabe K, Sakakibara M, Tadokoro M, Akashi K, Aoki H and Suzuki N. Introduction of the MASH1 gene into mouse embryonic stem cells leads to di#erentiation of motoneuron precursors lacking Nogo receptor expression that can be applicable for transplantation to spinal cord injury. Neurobiol Dis 2006; 22: Kamochi H, Kurokawa MS, Yoshikawa H, Ueda Y, Masuda C, Takada E, Watanabe K, Sakakibara M, Natuki Y, Kimura K, Beppu M, Aoki H and Suzuki N. Transplantation of myocyte precursors derived from embryonic 21

6 148 stem cells transfected with IGFII gene in a mouse model of muscle injury. Transplantation 2006; 82: Homma R, Yoshikawa H, Takeno M, Kurokawa MS, Masuda C, Takada E, Tsubota K, Ueno S and Suzuki N. Induction of epithelial progenitors in vitro from mouse embryonic stem cells and application for reconstruction of damaged cornea in mice. Invest Pohthalmol Vis Sci 2004; 45: Miyagi T, Takeno M, Nagafuchi H, Takahashi M and Suzuki N. Flk1 cells derived from mouse embryonic stem cells reconstitute hematopoiesis in vivo in SCID mice. Exp Hematol 2002; 30: Keller G, Kennedy M, Papayannopoulou T and Wiles MV. Hematopoietic commitment during embryonic stem cell di#erentiation in culture. Mol Cell Biol 1993; 13: Kennedy M, Firpo M, Choi K, Wall C, Robertson S, Kabrun N and Keller G. A common precursor for primitive erythropoiesis and definitive haematopoiesis. Nature 1997; 386: Park C, Ma YD and Choi K. Evidence for the hemangioblast. Exp Hematol 2005; 33: Risau W. Mechanisms of angiogenesis. Nature 1997; 386: Buschmann I and Schaper W. The pathophysiology of the collateral circulation arteriogenesis. J Pathol 2000; 190: Hirashima M, Kataoka H, Nishikawa S, Matsuyoshi N and Nishikawa S. Maturation of embryonic stem cells into endothelial cells in an in vitro model of vasculogenesis. Blood 1999; 93: Yamashita J, Itoh H, Hirashima M, Ogawa M, Nishikawa S, Yurugi T, Naito M, Nakao K and Nishikawa S. Flk1-positive cells derived from embryonic stem cells serve as vascular progenitors. Nature 2000; 408: Sone M, Itoh H, Yamashita J, Yurugi- Kobayashi T, Suzuki Y, Kondo Y, Nonoguchi A, Sawada N, Yamahara K, Miyashita K, Park K, Shibuya M, Nito S, Nishikawa S and Nakao K. Di#erent di#erentiation kinetics of vascular progenitor cells in primate and mouse embryonic stem cells. Circulation 2003; 107: Levenberg S, Golub JS, Amit M, Itskovitz- Eldor J and Langer R. Endothelial cells derived from human embryonic stem cells. Proc Natl Acad Sci USA 2002; 99: Gerecht-Nir S, Ziskind A, Cohen S and Itskovitz-Eldor J. Human embryonic stem cells as an in vitro model for human vascular development and the induction of vascular di#erentiation. Lab Invest 2003; 83: Lugus JJ, Park C and Choi K. Developmental relationship between hematopoietic and endothelial cells. Immunol Res 2005; 32: Heeschen C, Weis M, Aicher A, Dimmeler S and Cooke JP. A novel angiogenic pathway mediated by non-neuronal nicotinic acetylcholine receptors. J Clin Invest 2002; 110: Sato Y. Role of ETS family transcription factors in vascular development and angiogenesis. Cell Struct Funct 2001; 26: Miyashita H, Kanemura M, Yamazaki T, Abe M and Sato Y. Vascular endothelial zinc finger 1 is involved in the regulation of angiogenesis: possible contribution of stathmin OP18 as a downstream target gene. Arterioscler Thromb Vasc Biol 2004; 24: Minami T, Murakami T, Horiuchi K, Miura M, Noguchi T, Miyazaki J, Hamakubo T, Aird WC and Kodama T. Interaction between hex and GATA transcription factors in vascular endothelial cells inhibits flk-1 KDR-mediated vascular endothelial growth factor signaling. J Biol Chem 2004; 279: Gerety SS, Wang HU, Chen ZF and Anderson DJ. Symmetrical mutant phenotypes of the receptor EphB4 and its specific transmembrane ligand ephrin-b 2 in cardiovascular development. Mol Cell 1999; 4: Wang HU, Chen ZF and Anderson DJ. Mo- 22

7 149 lecular distinction and angiogenic interaction between embryonic arteries and veins revealed by ephrin-b2 and its receptor Eph-B4. Cell 1998; 93: Zhong TP, Childs S, Leu JP and Fishman MC. Gridlock signalling pathway fashions the first embryonic artery. Nature 2001; 414:

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