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1 Embryonic stem cells 1

2 Stem cells possess the remarkable ability of extensive self-renewal and differentiation into specific cell lineages. Stem cells play essential roles in development and adult tissue homeostasis. Due to their critical importance in normal physiology and the promise for use in regenerative medicine to treat a variety of diseases, stem cells have attracted extensive research interest in recent years. (Cell Res. 17:1-2, 2007) 2

3 Isolation of embryonic stem (ES) cells. 3

4 Immunosurgery on mouse blastocysts a. Mouse blastocysts; b. Blastocysts after exposure to anitserum and then to complement; c. Isolated inner cell mass cleared off dead trophoblastic cells; d. Inner cell mass after 24 h in cultureattachment and outgrowth of cells containing vacuoles; e. Two of several cell types observed in culture of inner cell masses 6 days after plating; f. Embryoid body derived from single inner cell mass after 4 days in culture. (Proc. Natl. Acad. Sci. USA. 72: , 1975) 4

5 A C B A. In vivo derived rabbit blastocysts; B. Blastocyts after removal of the zona pellucida using pronase digestion. (The inner cell mass, ICM, arrow); C. The ICMs were separated from the trophoblast cells by immunosurgery with some lysed or vacuolized cells (arrow head) attached to the border of the clump (arrow). ( 江, 2004) 5

6 A C B D A. Morphology of the rabbit ES cell colonies. The rabbit ES cell colony after seeding 2 days; B. The rabbit ES cell colony after seeding 3 days; C. The colony was losing the typical ES cell morphology; D. The ES cells differentiated into fibroblast-like cells; E. The flattened and differentiated ES cells; F. AP expression in undifferentiated rabbit ES cells. E F ( 歐, 2007) 6

7 Human IVF blastocyst Isolated ICM attached on MEF Primary hes cell colony on feeder cells Derivation of hes cells. (a) Day 8 (IVF=day 0) hatching blastocyst derived after in vitro fertilization. (b) ICM isolated by immunosurgery or mechanical isolation, and attached to the mouse feeder cells (MEF). (c) Thirteen-day-old primary hes cell colony grown on MEF. (d) Undifferentiated hes-ncl1 colony grown on human feeder. (e) Spontaneous differentiation of hes- NCL1cells with neuronal-precursors morphology. Induced differentiation of pluripotent hes cells could be achieved after addition of specific factor. Undifferentiated hes cells on feeder cells Differentiated hes cells (Reproduction, 128: , 2004) 7

8 Immunosurgery of a human blastocyst for the derivation of human ES cell line. (J. Anat , 2002) 8

9 Human ES cells. (a) grown on mouse feeder cells; (b) stained with specific cell-surface marker (TRA1-60); (c) When hes cells are transferred to non-adherent culture plates, they spontaneously form differentiated structures termed embryoid bodies. (Reproduction, 128: , 2004) 9

10 (Eur. J. Cancer 42: , 2006) 10

11 The morphology of undifferentiated ES cell colonies and ES single cells grown in the feeder layer-free culture system (A C) and on MEF feeder layer (D-F). Note the spaces between the cells (A and B) and the high nucleus:cytoplasm ratio typical of hes cells (C and F). (Biol. Reprod , 2004) 11

12 Fluorescent immunostaining of hes cells grown in the feeder layer-free culture system (A-C) and on MEF feeder layer (D-F). (A, D) anti-ssea4 antibody (B, E) anti-tra-1-60 antibody (C, F) anti-tra-1-81 (Biol. Reprod , 2004) 12

13 EB Epithelium ball like structure tubulin actin CD-31 In vitro differentiation of hes cells grown in the feeder layer-free culture system. (Biol. Reprod , 2004) 13

14 Myelinated nerve (ectoderm) Details of hyaline cartilage (mesoderm) Secretory epithelium rich in goblet cells (endoderm) In vivo differentiation of ES cell lines in teratomas. (Biol. Reprod , 2004) 14

15 The LIF/ERK signalling pathway. (Eur. J. Cancer 42: , 2006) 15

16 The TGF signalling pathway. (Eur. J. Cancer 42: , 2006) 16

17 The BMP signalling pathway. (Eur. J. Cancer 42: , 2006) 17

18 BMP signal transduction pathways. (Dev. Biol. 284:1-11, 2005) 18

19 The Wnt signalling pathway. (Eur. J. Cancer 42: , 2006) 19

20 Possible crosstalk between BMP, FGF/PI3K/AKT and Wnt signalling pathways. (Eur. J. Cancer 42: , 2006) 20

21 Table 1. Expression of self-renewal and pluripotent markers in the ES cells Antigens Human Monkey Mouse Rabbit SSEA-1 SSEA-3 SSEA-4 TRA-1-60 TRA Alkaline phosphatase (AP) Nanog Oct-4 Sox-2 (Mitalipov et al., 2006; Stewart et al., 2006; Wang et al., 2007) 21

22 Cloned mice and ntes cells. (Nat. Biotechnol , 2004) 22

23 Steps of the therapeutic cloning technique. (Reproduction, 128: , 2004) 23

24 nuclear transfer of human somatic nuclei into rabbit oocytes Micrographs of nt-units derived from human somatic cells (A) nt-units derived from fibroblasts of a 42-year old donor at 4-cell stage; ( B) morula stage; (C) early blastocyst stage; and (D) hatching blastocyst stage. The arrow points to a visible crack in the zona pellucida. (E-G) An nt-unit going through M-phase during the first mitotic division (6 h after activation) with a generally normal spindle structure. (H) An nt-unit with an unincorporated somatic nucleus. (Bars = 20 μ m) (Cell Res., 13: , 2003) 24

25 Table Somatic cells from donors at different ages formed blastocysts with comparable efficiency. (Cell Res., 13: , 2003) 25

26 Alu probes hybridize specifically to nuclei of primate species. (A) Cultured human, (B) monkey (Rhesus macaque), (C) rabbit and (D) mouse fibroblasts and paraffin sections of (E) human liver, (F) sheep liver, (G) rabbit ovary and (H) mouse ovary were hybridized to Alu probes. Only human and monkey nuclei showed positive signals above the background. (Cell Res., 13: , 2003) 26

27 Results from in situ hybridization demonstrating that probes for rabbit mitochondrial DNA (red) are specie specific. (I, M) Rabbit fibroblasts (J, N) human fibroblasts (K, O) a mixed population of rabbit and human fibroblasts (L, P) Rabbit fibroblasts were hybridized without probes to serve as negative controls (Cell Res., 13: , 2003) 27

28 nt-units at the blastocyst stage contain the primate genome and rabbit mitochondrial DNA. (Q, R, U, V) Paraffin sections of nt-units at the blastocyst stage, (S, W) rabbit parthenogenotes and (T, X) human fibroblasts were hybridized to either (Q, R, S, T) Alu probes or (U, V, W, X) rabbit mitochondrial DNA probes. nt-units at the blastocyst stage were stained positively by both (Q, R) Alu and (U, V) rabbit mitochondrial DNA probes. (W) Rabbit parthenogenotes stained positively by only rabbit mitochondrial DNA probes, and (T) human fibroblasts by only Alu probes. (Cell Res., 13: , 2003) 28

29 Morphology and marker expression in ntes cells (A) An ntes cell colony derived from a 42-year-old somatic cell donor. (B) Higher magnification of (A). (C) A colony derived from a 52-year-old somatic cell donor. (D) Alkaline phosphatase activity in ntes cells after 26 passages. The ntes cells express epitopes recognized by antibodies against (E) SSEA-3, (F) SSEA-4, (G) TRA-1-10, and (H) TRA The ntes cells failed to stain for SSEA-1 (data not shown). (Cell Res., 13: , 2003) 29

30 Somatic differentiation of ntes cells (A) An EB. (B) Outgrowth from an EB. (C-J) Cryostat sections of EBs stained by antibodies against (C) nestin, (D) neuron specific enolase, (E) myoglobin, (F) a-smooth muscle actin, (G) VEGF receptor-2, (H) Tie-2, (I) a-fetoprotein, (J) a- 1-antitrypsin. (Cell Res., 13: , 2003) 30

31 Multilineage differentiation of the ntes cells that have grown out from an EB. (K) and (L) Cells containing fat droplets stained by Oil Red O. (M) Muscle cells. (N) Muscle fibers stained by antibodies against myoglobin. Marker expression by cell groups in the outgrowth: (O) -fetoprotein, (P) - 1-antitrypsin, (Q) VEGF recptor-2, (R) Tie-2, (S) von Willebrand Factor, (T) nestin, (U) myoglobin, and (V) MyoD1. (W-Z) Neuron formation. (W) Neuronal network formed by ntes cells. Neurons stained by (X) neurofilament-h, (Y) neuron specific enolase, and (Z) nestin, respectively. (Cell Res., 13: , 2003) 31

32 Homologous recombination: the targeting vector contains two regions of homology with the endogenous genomic locus to be targeted. (Adv. Drug Deliv. Rev. 57: , 2005) 32

33 Generation of mutant mice from targeted ES cells. (Anat. Histol. Embryol , 2002) 33

34 Generation of mutant mice from targeted ES cells. (Anat. Histol. Embryol , 2002) 34

35 Utilization of the Cre recombinase system for generation of conditional knock-outs. (Anat. Histol. Embryol , 2002) 35

36 Therapeutic cloning as a method of providing autologous cell transplants. (Anat. Histol. Embryol , 2002) 36

37 Methods to reprogram the somatic cells Somatic cells can be reprogrammed by transferring their nuclear contents into oocytes or by fusion with ES cells. Unfertilized eggs and ES cells contain factors that can confer totipotency or pluripotency to somatic cells. 37

38 Different approaches for studying nuclear reprogramming. (Nature, , 2006) 38

39 The developmental potential of ES hybrid cells in vivo. An experimental scheme for generating ES hybrid cells and making chimeric embryos. (Curr. Biol. 11: , 2001) 39

40 Chimeric embryos (E7.5) with ES hybrid cells. Ect, ectoderm; Mes, mesoderm; End, endoderm. (Curr. Biol. 11: , 2001) 40

41 Reprogramming of human cells by Xenopus egg extract. (A) Diagram of the experiment. Top, preparation of egg extract; bottom, preparation of human cells. (Curr. Biol. 14: , 2004) 41

42 Comparative RT-PCR analysis of reprogrammed 293T cell spheres harvested at day 5 and the TERA-1(human embryonal carcinoma cell line). (Curr. Biol. 14: , 2004) 42

43 (8 cell) (16 cell)(late blastula) Properties of Xenopus reprogramming extracts analyzed by RT-PCR. (Curr. Biol. 14: , 2004) 43

44 Differentiation and cloning efficiency. (Nature, , 2006) 44

45 In vitro reprogramming of fibroblasts into apluripotent ES-cell-like state- Oct4, Sox2, c-myc and Klf4 Targeting strategy to generate an Oct4-IRES-GFPneo allele. (Nature, 448: , 2007) 45

46 Phase-contrast micrograph of Oct4 ips cells. b, (clone 18), exhibited strong alkaline phosphatase activity (c) antibodies against SSEA1 (d, e) and Nanog (f, g). h, One example of an ES-like colony 16 days after infection (left).non- ES-like cells (right) (Nature, 448: , 2007) 46

47 Paraffin sections of a teratoma 26 days after subcutaneous injection of Oct4 ips clone 18 cells into SCID mice. H&E: haematoxylin and eosin. e-f: expression was confined to undifferentiated cell types as indicated an immunohistochemical analysis. e: Nanog f :Oct4 (Nature, 448: , 2007) 47

48 Two live pups after 2N blastocyst injection. (c) of the TTF-derived Oct4 ips cell line TT-O25, which had been GFP-labelled with a lentiviral ubiquitin-egfp vector. d, All ips cell embryos were generated by injection of ips cells into 4N blastocysts. Live E12.5 embryos generated from Oct4 ips line O6 (left), from Nanog ips line N14 (middle) and from V.6.5 ES cells (right) are shown. e, A normally developed E14.5 embryo was derived from Oct4 ips cell line O4-16. (Nature, 448: , 2007) 48

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