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1 植物科学最前線 3:56 (2012) Marchantia polymorpha L International Botanical Congress T. Ueda, S. Sawa & T. Araki - 1 BSJ-Review 3:56 (2012)

2 植物科学最前線 3:57 (2012) T. Ueda, S. Sawa & T. Araki - 2 BSJ-Review 3:57 (2012)

3 植物科学最前線 3:58 (2012) Liverwort, Marchantia polymorpha L., as a reviving model for plant biology Key words: basal land plant, genomics, land plant evolution, Marchantia polymorpha, molecular genetics Takayuki Kohchi and Kimitsune Ishizaki Graduate School of Biostudies, Kyoto University Kyoto , Japan 19 Evo Devo Evo Eco Marchantia polymorpha L. 4 Qiu et al T. Kohchi & K. Ishizaki-1 BSJ-Review 3:58 (2012)

4 植物科学最前線 3:59 (2012) - 1 Kny 1890 WEB Marchantia (2012) (2012) n 2n Nishiyama et al. 2003, Sano et al. 2005, Sakakibara et al., 2008 ) 1. T. Kohchi & K. Ishizaki-2 BSJ-Review 3:59 (2012)

5 植物科学最前線 3:60 (2012) - 8 X Y 2012 Wann 1925 Lloyd & Steinmetz 1937 Chiyoda et al T. Kohchi & K. Ishizaki-3 BSJ-Review 3:60 (2012)

6 植物科学最前線 3:61 (2012) 30 Spore Sporelings 2-3 Gemma Gemmalings Gemmalings Sporelings Kubota et al T. Kohchi & K. Ishizaki-4 BSJ-Review 3:61 (2012)

7 植物科学最前線 3:62 (2012) 3. Barnes & Land (1908) DNA - n= T. Kohchi & K. Ishizaki-5 BSJ-Review 3:62 (2012)

8 植物科学最前線 3:63 (2012) Burgeff Genetische Studien an Marchantia Burgeff Ono 1976 Ohyama et al. 1986, Oda et al Takaragaike-1 Takaragaike-2 (Okada et al. 2000) Marchantia polymorpha ssp. ruderalis Marchantia polymorpha Takaragaike-1 Takaragaike-2 PAC Okada et al Okada et al. 2001, Ishizaki et al. 2002, Yamato et al Takaragaike-1 Takaragaike Takaragaike-1 Takaragaike-2 Joint Genome Institute Kitashirakawa Takaragaike F1 Recombinant Inbred - T. Kohchi & K. Ishizaki-6 BSJ-Review 3:63 (2012)

9 植物科学最前線 3:64 (2012) 4. Sporelings Ishizaki et al Sporelings , Sporelings T-DNA T-DNA T-DNA 70,000 90% 20 T-DNA T-DNA TAIL-PCR ARF+ 2 Sporelings T. Kohchi & K. Ishizaki-7 BSJ-Review 3:64 (2012)

10 植物科学最前線 3:65 (2012) Sandy Floyd T1 T2 T3 T1 T1 gemma G1 G1 G1 G T. Kohchi & K. Ishizaki-8 BSJ-Review 3:65 (2012)

11 植物科学最前線 3:66 (2012) - Terada et al DTA T-DNA T-DNA PCR DNA PCR DNA Boynton et al Chiyoda et al. 2007, Chiyoda et al. 2008, 2008 Ueda et al DNA DNA Sporelings Chiyoda et al EST Nishiyama et al Rensing et al. 2008, Banks et al T. Kohchi & K. Ishizaki-9 BSJ-Review 3:66 (2012)

12 植物科学最前線 3:67 (2012) PIF Leivar et al PIF Burgeff (1943) EMS phot T-DNA 2012 T. Kohchi & K. Ishizaki-10 BSJ-Review 3:67 (2012)

13 植物科学最前線 3:68 (2012) 2012, 2012 accession Joint Genome Institute Monash John Bowman ( ) LEAFY. BSJ Review. Banks, J. A., Nishiyama, T., Hasebe, M. et al The Selaginella genome identifies genetic changes associated with the evolution of vascular plants. Science 332: Barnes, C. R., & Land, W. J. G The origin of the cupule of Marchantia. Bot. Gazette 46: Boynton, J. E., Gillham, N.W., Harris, E. H., Hosler, J. P., Johnson, A. M., Jones, A. R., Randolph-Anderson, B. L., Robertson, D., Klein, T. M., Shark, K. B., & Sanford, J. C Chloroplast transformation in Chlamydomonas with high velocity microprojectiles. Science 240: Burgeff, H Genetische Studien an Marchantia. Verlag von Gustav Fischer, Jena. Chiyoda, S., Linley, P. J., Yamato, K. T., Fukuzawa, H., Yokota, A., & Kohchi, T Simple and efficient plastid transformation system for the liverwort Marchantia polymorpha L. suspension-culture cells. Transgenic Res. T. Kohchi & K. Ishizaki-11 BSJ-Review 3:68 (2012)

14 植物科学最前線 3:69 (2012) 16: Chiyoda, S., Ishizaki, K., Kataoka, H., Yamato, K. T., & Kohchi, T Direct transformation of the liverwort Marchantia polymorpha L. by particle bombardment using immature thalli developing from spores. Plant Cell Rep. 27: b. 67: Ishizaki, K., Shimizu-Ueda, Y., Okada, S., Yamamoto, M., Fujisawa, M., Yamato, K.T., Fukuzawa, H., & Ohyama, K Multicopy genes uniquely amplified in the Y chromosome-specific repeats of the liverwort Marchantia polymorpha. Nucl. Acids Res. 30: Ishizaki, K., Chiyoda, S., Yamato, K. T., & Kohchi, T Agrobacterium-mediated transformation of the haploid liverwort Marchantia polymorpha L., an emerging model for plant biology. Plant Cell Physiol. 49: a. 67: BSJ Review 3: Kny, L Bau und Entwickelung von Marchantia polymorpha L.In Botanische Wandtafeln mit Erläuterndem Text pp Verlag von Wiegandt, Hempel & Parey, Berlin. Kubata, A., Ishizaki, K., Hosaka, M., & Kohchi, T Efficient Agrobacterium-mediated transformation of the liverwort Marchantia polymorpha using regenerating thalli. Biosci. Biotechnol. Biochem. in press. Leivar, P., Monte, E., Oka, Y., Liu, T., Carle, C., Huq, E., & Quail, P.H Multiple phytochrome-interacting bhlh transcription factors repress premature photomorphogenesis during early seedling development in darkness. Current Biol. 18: Lloyd, C.E., & Steinmetz, F.H Temperature as a factor influencing the sexual response of Marchantia. Americ. J. Bot. 24: Nishiyama, T., Fujita, T., Shin-I, T., Seki, M., Nishide, H., Uchiyama, I., Kamiya, A., Carninci, P., Hayashizaki, Y., Shinozaki, K., Kohara, Y., & Hasebe M Comparative genomics of Physcomitrella patens gametophytic transcriptome and Arabidopsis thaliana: implication for land plant evolution. Proc. Natl. Acad. Sci. USA. 100: Oda, K., Yamato, K., Ohta, E., Nakamura, Y., Takemura, M., Nozato, N., Akashi, K., Kanegae, T., Ogura, Y., Kohchi, T., & Ohyama, K. Gene organization deduced from the complete sequence of liverwort Marchantia polymorpha mitochondrial DNA. A primitive form of plant mitochondrial genome. J. Mol. Biol. 223: 1-7. Ohyama, K., Fukuzawa, H., Kohchi, T., Shirai, H., Sano, T., Sano, S., Umesono, K., Shiki, Y., Takeuchi, M., Chang, Z., Aota, S., Inokuchi, H., & Ozeki, H Chloroplast gene organization deduced from complete sequence of liverwort Marchantia polymorpha chloroplast DNA. Nature 322: Okada, S., Fujisawa, M., Sone, T., Nakayama, S., Nishiyama, R., Takenaka, M., Yamaoka, S., Sakaida, M., Kono, K., Takahama, M., Yamato, K.T., Fukuzawa, H., Brennicke, A., & Ohyama, K Construction of male and female PAC genomic libraries suitable for identification of Y-chromosome-specific clones from the liverwort, Marchantia polymorpha, Plant J. 24: T. Kohchi & K. Ishizaki-12 BSJ-Review 3:69 (2012)

15 植物科学最前線 3:70 (2012) Okada, S., Sone, T., Fujisawa, M., Nakayama, S., Takenaka, M., Ishizaki, K., Kono, K., Shimizu-Ueda, Y., Hanajiri, T., Yamato, K.T., Fukuzawa, H., Brennicke, A., & Ohyama, K The Y chromosome in the liverwort Marchantia polymorpha has accumulated unique repeat sequences harboring a male-specific gene. Proc. Natl. Acad. Sci. USA. 98: Ono, K Cytological observations of calluses and restored thalluses in Marchantia polymorpha. Jpn. J. Genet. 51: Qui, Y.L., Li, L., Wang, B., Chen, Z., Knoop, V., Groth-Malonek, M., Dombrovska, O., Lee, J., Kent, L., Rest, J., Estabrook, G.F., Hendry, T.A., Taylor, D.W., Testa, C.M., Ambros, M., Crandall-Stotler, B., Duff, R.J., Stech, M., Frey, W., Quandt, D.. & Davis, C.C The deepest divergences in land plants inferred from phylogenomic evidence. Proc. Natl. Acad. Sci. USA 103: Rensing, S. A., Lang, D., Zimmer, A. D. et al The Physcomitrella genome reveals evolutionary insights into the conquest of land by plants. Science 319: Sakakibara, K., Nishiyama, T., Deguchi, H., & Hasebe, M Class 1 KNOX genes are not involved in shoot development in the moss Physcomitrella patens but do function in sporophyte development. Evol. Dev. 10: Sano, R., Juárez, C.M., Hass, B., Sakakibara, K., Ito, M., Banks, J.A., & Hasebe, M KNOX homeobox genes potentially have similar function in both diploid unicellular and multicellular meristems, but not in haploid meristems. Evol. Dev. 7: MpCLE1. BSJ Review 3: BSJ Review 3: Terada, R., Johzuka-Hisatomi, Y., Saitoh, M., Asao, H. & Iida, S Gene targeting by homologous recombination as a biotechnological tool for rice functional genomics. Plant Physiol. 144: Ueda, M., Kuniyoshi, T., Yamamoto, H., Sugimoto K., Ishizaki, K., Kohchi, T., Nishimura, Y., & Shikanai, T Composition and physiological function of the chloroplast NADH dehydrogenase-like complex in Marchantia polymorpha. Plant J. 72: Wann, F.B Some of the factors involved in the sexual reproduction of Marchantia polymorpha. Americ. J. Bot. 12: Yamato, K. T., Ishizaki, K., Fujisawa, M., Okada, S., Nakayama, S., Fujishita, M., Bando, H., Yodoya, K., Hayashi, K., Bando, T., Hasumi, A., Nishio, T., Sakata, R., Yamamoto, M., Yamaki, A., Kajikawa, M., Yamano, T., Nishide, T., Choi, S. H., Shimizu-Ueda, Y., Hanajiri, T., Sakaida, M., Kono, K., Takenaka, M., Yamaoka, S., Kuriyama, C., Kohzu, Y., Nishida, H., Brennicke, A., Shin-i, T., Kohara, Y., Kohchi, T., Fukuzawa, H. & Ohyama, K Gene organization of the liverwort Y chromosome reveals distinct sex chromosome evolution in a haploid system. Proc. Natl. Acad. Sci. USA. 104: : BSJ Review 3: T. Kohchi & K. Ishizaki-13 BSJ-Review 3:70 (2012)

16 植物科学最前線 3:71 (2012) The sneak preview of the Marchantia polymorpha genome Key words: bryophyte, genetic redundancy, liverwort Katsuyuki T. Yamato 1, Takayuki Kohchi 2 1 Department of Biotechnological Science Kinokawa , Japan 2 Graduate School of Biostudies, Kyoto University Kyoto , Japan Marchantia polymorpha L. (The Arabidopsis Genome Initiative 2000) (International Rice Genome Sequencing Project 2005) (Rensing et al. 2008) (Ohyama et al. 1986) (Shinozaki et al. 1986) 120 kb 160 kb K. T. Yamato & T. Kohchi-1 BSJ-Review 3:71 (2012)

17 植物科学最前線 3:72 (2012) 16 kb (Anderson et al. 1981) 50 kb (Sanger et al. 1982) 1992 (Oda et al. 1992) Y 10 Mb (Yamato et al ) Y (Skaletsky et al. 2003) (Kuroki et al. 2006) X Joint Genome Institute (JGI) Monash John Bowman ( ( 2012) JGI P1 PAC (Okada et al. 2000) PAC JGI 10 PAC 3.3 Mb 280 Mb (Okada et al. 2000) 1% DNA GC 40% PAC kb 1 1 K. T. Yamato & T. Kohchi-2 BSJ-Review 3:72 (2012)

18 植物科学最前線 3:73 (2012) EST 1 10 kb kb 2.3 TAIR10 pmm23-619a2 EST 18,000 20, /10 kb = 20,000 /280 Mb K. T. Yamato & T. Kohchi-3 BSJ-Review 3:73 (2012)

19 植物科学最前線 3:74 (2012) pmm23-619a2 NCBI NR NCBI non-redundant protein sequences Atha, Smol Ppat Mpol EST EST K. T. Yamato & T. Kohchi-4 BSJ-Review 3:74 (2012)

20 植物科学最前線 3:75 (2012) 4 6 trna trna mirna RNA (Floyd and Bowman 2004) rrna 9 X 1 (Fujisawa et al. 2003, Sone et al. 1999) JGI Takaragaike-1 Kitashirakawa-2 ( 2012) 30 PAC 31 EST dcaps (derived cleaved amplified polymorphic sequences) SSR (simple sequence repeats) cm JGI Takaragaike-1 Kitashirakawa-2 F 1 Kitashirakawa-2 JGI Takaragaike-1 Kitashirakawa-2 K. T. Yamato & T. Kohchi-5 BSJ-Review 3:75 (2012)

21 植物科学最前線 3:76 (2012) JGI 2 Fosmid Roche GS-FLX 27x ver. 0.6 ver , Mb X 9 JGI EST 90% cdna 5 /3 (UTR) mrna 5 AUG ATG ATG UTR-ATG 5 UTR K. T. Yamato & T. Kohchi-6 BSJ-Review 3:76 (2012)

22 植物科学最前線 3:77 (2012) 2 10 ATG EST 5 (Strasser et al. 2010) (Leivar and Quail 2011) (Schaefer 2001) (Rensing et al. 2008) (Rensing et al. 2007) LFY 2 ( 2012) PHYTOCHROME INTERACTIG FACTOR (PIF) 1 K. T. Yamato & T. Kohchi-7 BSJ-Review 3:77 (2012)

23 植物科学最前線 3:78 (2012) CLE 32 ( 2012) (Sasaki et al. 2007) ( 2012) 8 X n = 8 + X Y n = 8 + Y Y X Y Y (Yamato et al. 2007) Y 0.1 / 10 kb 0.7 X K. T. Yamato & T. Kohchi-8 BSJ-Review 3:78 (2012)

24 植物科学最前線 3:79 (2012) X Y X X JGI ver Mb Y X X Y X Y Y (Egydio de Carvalho et al. 2002) (Lorenzetti et al. 2004) (Ikeda et al. 2007) (Hirai et al. 2008, Mori et al. 2006) X Y JGI Kitashirakawa-2 JGI K. T. Yamato & T. Kohchi-9 BSJ-Review 3:79 (2012)

25 植物科学最前線 3:80 (2012) JGI Monash John Bowman Sandra Floyd cdna X Y K. T. Yamato & T. Kohchi-10 BSJ-Review 3:80 (2012)

26 植物科学最前線 3:81 (2012) Anderson, S., Bankier, A.T., Barrell, B.G., de Bruijn, M.H., Coulson, A.R., Drouin, J., Eperon, I.C., Nierlich, D.P., Roe, B.A., Sanger, F., Schreier, P.H., Smith, A.J., Staden, R., & Young, I.G Sequence and organization of the human mitochondrial genome. Nature 290: Egydio de Carvalho, C., Tanaka, H., Iguchi, N., Ventela, S., Nojima, H., & Nishimune, Y Molecular cloning and characterization of a complementary DNA encoding sperm tail protein SHIPPO 1. Biol. Reprod. 66: Floyd, S.K., & Bowman, J.L Gene regulation: Ancient microrna target sequences in plants. Nature 428: Fujisawa, M., Nakayama, S., Nishio, T., Fujishita, M., Hayashi, K., Ishizaki, K., Kajikawa, M., Yamato, K.T., Fukuzawa, H., & Ohyama, K Evolution of ribosomal DNA unit on the X chromosome independent of autosomal units in the liverwort Marchantia polymorpha. Chromosome Res. 11: Hirai, M., Arai, M., Mori, T., Miyagishima, S.Y., Kawai, S., Kita, K., Kuroiwa, T., Terenius, O., & Matsuoka, H Male fertility of malaria parasites is determined by GCS1, a plant-type reproduction factor. Curr. Biol. 18: Ikeda, K., Ikeda, T., Morikawa, K., & Kamiya, R Axonemal localization of Chlamydomonas PACRG, a homologue of the human Parkin-coregulated gene product. Cell Motility and the Cytoskeleton 64: International Rice Genome Sequencing Project The map-based sequence of the rice genome. Nature 436: Kuroki, Y., Toyoda, A., Noguchi, H., Taylor, T.D., Itoh, T., Kim, D.S., Kim, D.W., Choi, S.H., Kim, I.C., Choi, H.H., Kim, Y.S., Satta, Y., Saitou, N., Yamada, T., Morishita, S., Hattori, M., Sakaki, Y., Park, H.S., & Fujiyama, A Comparative analysis of chimpanzee and human Y chromosomes unveils complex evolutionary pathway. Nat. Genet. 38: Leivar, P., & Quail, P.H PIFs: pivotal components in a cellular signaling hub. Trends in Plant Science 16: Lorenzetti, D., Bishop, C.E., Justice, & M.J Deletion of the Parkin coregulated gene causes male sterility in the quaking viable mouse mutant. Proc. Natl. Acad. Sci. USA 101: Mori, T., Kuroiwa, H., Higashiyama, T., & Kuroiwa, T GENERATIVE CELL SPECIFIC 1 is essential for angiosperm fertilization. Nature Cell Biol. 8: Oda, K., Yamato, K., Ohta, E., Nakamura, Y., Takemura, M., Nozato, N., Akashi, K., Kanegae, T., Ogura, Y., Kohchi, T., et al Gene organization deduced from the complete sequence of liverwort Marchantia polymorpha mitochondrial DNA. A primitive form of plant mitochondrial genome. J. Mol. Biol. 223: 1-7. Ohyama, K., Fukuzawa, H., Kohchi, T., Shirai, H., Sano, T., Sano, S., Umesono, K., Shiki, K., Takeuchi, M., Chang, Z., Aota, S., Inokuchi, H., & Ozeki, H Chloroplast gene organization deduced from complete sequence of liverwort, Marchantia polymorpha chloroplast DNA. Nature 322: Okada, S., Fujisawa, M., Sone, T., Nakayama, S., Nishiyama, R., Takenaka, M., Yamaoka, S., Sakaida, M., K. T. Yamato & T. Kohchi-11 BSJ-Review 3:81 (2012)

27 植物科学最前線 3:82 (2012) Kono, K., Takahama, M., Yamato, K.T., Fukuzawa, H., Brennicke, A., & Ohyama, K Construction of male and female PAC genomic libraries suitable for identification of Y-chromosome-specific clones from the liverwort, Marchantia polymorpha. Plant J. 24: Rensing, S.A., Ick, J., Fawcett, J.A., Lang, D., Zimmer, A., De Peer, Y.V., & Reski, R An ancient genome duplication contributed to the abundance of metabolic genes in the moss Physcomitrella patens. BMC Evol. Biol. 7: Rensing, S.A., Lang, D., Zimmer, A.D., Terry, A., Salamov, A., Shapiro, H., Nishiyama, T., Perroud, P.F., Lindquist, E.A., Kamisugi, Y., Tanahashi, T., Sakakibara, K., Fujita, T., Oishi, K., Shin-I, T., Kuroki, Y., Toyoda, A., Suzuki, Y., Hashimoto, S., Yamaguchi, K., Sugano, S., Kohara, Y., Fujiyama, A., Anterola, A., Aoki, S., Ashton, N., Barbazuk, W.B., Barker, E., Bennetzen, J.L., Blankenship, R., Cho, S.H., Dutcher, S.K., Estelle, M., Fawcett, J.A., Gundlach, H., Hanada, K., Heyl, A., Hicks, K.A., Hughes, J., Lohr, M., Mayer, K., Melkozernov, A., Murata, T., Nelson, D.R., Pils, B., Prigge, M., Reiss, B., Renner, T., Rombauts, S., Rushton, P.J., Sanderfoot, A., Schween, G., Shiu, S.H., Stueber, K., Theodoulou, F.L., Tu, H., Van de Peer, Y., Verrier, P.J., Waters, E., Wood, A., Yang, L.X., Cove, D., Cuming, A.C., Hasebe, M., Lucas, S., Mishler, B.D., Reski, R., Grigoriev, I.V., Quatrano, R.S., & Boore, J.L The Physcomitrella genome reveals evolutionary insights into the conquest of land by plants. Science 319: Sanger, F., Coulson, A.R., Hong, G.F., Hill, D.F., & Peterse, G.B Nucleotide sequence of bacteriophage λ DNA. J. Mol. Biol. 162: Sasaki, G., Katoh, K., Hirose, N., Suga, H., Kuma, K., Miyata, T., & Su, Z.H Multiple receptor-like kinase cdnas from liverwort Marchantia polymorpha and two charophycean green algae, Closterium ehrenbergii and Nitella axillaris: Extensive gene duplications and gene shufflings in the early evolution of streptophytes. Gene 401: Schaefer, D.G Gene targeting in Physcomitrella patens. Current Opinion in Plant Biology 4: Shinozaki, K., Ohme, M., Tanaka, M., Wakasugi, T., Hayashida, N., Matsubayashi, T., Zaita, N., Chunwongse, J., Obokata, J., Yamaguchishinozaki, K., Ohto, C., Torazawa, K., Meng, B.Y., Sugita, M., Deno, H., Kamogashira, T., Yamada, K., Kusuda, J., Takaiwa, F., Kato, A., Tohdoh, N., Shimada, H., & Sugiura, M The Complete Nucleotide-Sequence of the Tobacco Chloroplast Genome - Its Gene Organization and Expression. EMBO J. 5: Skaletsky, H., Kuroda-Kawaguchi, T., Minx, P.J., Cordum, H.S., Hillier, L., Brown, L.G., Repping, S., Pyntikova, T., Ali, J., Bieri, T., Chinwalla, A., Delehaunty, A., Delehaunty, K., Du, H., Fewell, G., Fulton, L., Fulton, R., Graves, T., Hou, S.F., Latrielle, P., Leonard, S., Mardis, E., Maupin, R., McPherson, J., Miner, T., Nash, W., Nguyen, C., Ozersky, P., Pepin, K., Rock, S., Rohlfing, T., Scott, K., Schultz, B., Strong, C., Tin-Wollam, A., Yang, S.P., Waterston, R.H., Wilson, R.K., Rozen, S., & Page, D.C The male-specific region of the human Y chromosome is a mosaic of discrete sequence classes. Nature 423: Sone, T., Fujisawa, M., Takenaka, M., Nakagawa, S., Yamaoka, S., Sakaida, M., Nishiyama, R., Yamato, K.T., Ohmido, N., Fukui, K., Fukuzawa, H., & Ohyama, K Bryophyte 5S rdna was inserted into 45S rdna repeat units after the divergence from higher land plants. Plant Mol. Biol. 41: K. T. Yamato & T. Kohchi-12 BSJ-Review 3:82 (2012)

28 植物科学最前線 3:83 (2012) Strasser, B., Sanchez-Lamas, M., Yanovsky, M.J., Casal, J.J., & Cerdan, P.D Arabidopsis thaliana life without phytochromes. Proc. Natl. Acad. Sci. U S A 107: The Arabidopsis Genome Initiative Analysis of the genome sequence of the flowering plant Arabidopsis thaliana. Nature 408: Yamato, K.T., Ishizaki, K., Fujisawa, M., Okada, S., Nakayama, S., Fujishita, M., Bando, H., Yodoya, K., Hayashi, K., Bando, T., Hasumi, A., Nishio, T., Sakata, R., Yamamoto, M., Yamaki, A., Kajikawa, M., Yamano, T., Nishide, T., Choi, S.H., Shimizu-Ueda, Y., Hanajiri, T., Sakaida, M., Kono, K., Takenaka, M., Yamaoka, S., Kuriyama, C., Kohzu, Y., Nishida, H., Brennicke, A., Shin-i, T., Kohara, Y., Kohchi, T., Fukuzawa, H., & Ohyama, K Gene organization of the liverwort Y chromosome reveals distinct sex chromosome evolution in a haploid system. Proc. Natl. Acad. Sci. USA 104: BSJ Review 3: LEAFY. BSJ Review 3: Y. 63: MpCLE1. BSJ Review 3: K. T. Yamato & T. Kohchi-13 BSJ-Review 3:83 (2012)

29 植物科学最前線 3:84 (2012) An introduction to the taxonomy and morphology of Marchantia polymorpha Key words: bryophytes; Marchantia polymorpha; morphology; taxonomy Masaki Shimamura Department of Biology, Graduate School of Science, Hiroshima University Kagamiyama 1-3-1, Higashi-Hiroshima , Japan Leopold Kny ( ) (Kny1890) (Stotler 1994) 19 ( 2012) (Marchantiophyta) (Bryophyta) (Anthocerotophyta) M. Shimamura-1 BSJ-Review 3:84 (2012)

30 植物科学最前線 3:85 (2012) (Qiu et al. 2006, 2007) (Crandall-Stotler et al. 2009) (Haplomitriopsida) (Marchantiopsida) (Jungermanniopsida) 20 (simple thalloid) 400 (complex thalloid) (Wellman et al. 2003, Graham et al. 2004) (Walton 1925 Permian Triassic P-T 2 5 (Wheeler 2000) (He-Nygren et al. 2006) (Blasia pusilla) (Cavicularia densa) cm 2 4 (Duckett & Renzaglia 1993, Shimamura et al. 2005, 2006) (Forrest et al. 2006, Shimamura et al. 2012) 400 (Sphaerocarpales) (Neohodgsoniales) (Lunulariales) (Marchantiales) (Long 2006) 2 Marchantiaceae M. Shimamura-2 BSJ-Review 3:85 (2012)

31 植物科学最前線 3:86 (2012) (Marchantiaceae) (Boisselier-Dubayle et al Bucegia (Preissia) (Marchantia) (Marchantia) 40 M. polymorpha ssp. ruderalis M. polymorpha ssp. polymorpha Marchantia paleacea ssp. paleacea Marchantia paleacea. ssp. diptera Marchantia emarginata ssp. tosana Marchantia pinnata (Bishler 1989, 1998, 2012) 2. Shimamura et al. (2012) M. Shimamura-3 BSJ-Review 3:86 (2012)

32 植物科学最前線 3:87 (2012) 3. a. (Cavicularia densa) b. (Lunularia cruciata) 1 c. (Sphaerocarpos donnelli) d. (Preissia quadrata) 1 e. (Conocephalum conicum) 1 f. (Reboulia hemisphaerica ssp. orientalis) 1 3 g. (Sauteria yatsuensis) 2 3 h. (Riccia fluitans) 2 3 i. (Dumortiera hirsuta) M. Shimamura-4 BSJ-Review 3:87 (2012)

33 植物科学最前線 3:88 (2012) Marchantia polymorpha L Linnaeus (Linnaeus1753) M. polymorpha L. Linnaeus Burgeff (1943) M. polymorpha ( ) M. aquatica M. alpestris 3 Burgeff (1943) 3 M. polymorpha (Schuster 1992) (Bischler & Boisselier-Dubayle 1991, Perold 1999, Paton 1999) M. polymorpha L. (Linnaeus 1753) Bischler & Boisselier- -Dubayle (1991) (Dillenius ) M. aquatica M. polymorpha ssp. polymorpha M. aquatica M. polymorpha ssp. montivagans ( M. alpestris ) 2 cm M. polymorpha ssp.ruderalis M. polymorpha Burgeff (1943) 3 M. alpestris ( ssp. montivagans ) M. alpestris M. alpestris Schuster (1983, 1992) DNA 2 (Boisselier-Dubayle & Bischler 1989, Boisselier-Dubayle et al. 1995) 9 (n=9 ) (Bischler 1986). M. Shimamura-5 BSJ-Review 3:88 (2012)

34 植物科学最前線 3:89 (2012) (M. polymorpha ssp. polymorpha) Specim. exam. Japan. Pref. Gunma: Ozegahara, 1400 m alt. H. Ando 9143 (HIRO). 5 M. Shimamura-6 BSJ-Review 3:89 (2012)

35 植物科学最前線 3:90 (2012) µm (Inoue 1960, Nakazato et al. 1999) 6a (Inoue 1960) 6b 6c (Leitgeb 1880) 6d (O'Hanlon 1926) 6e M. Shimamura-7 BSJ-Review 3:90 (2012)

36 植物科学最前線 3:91 (2012) 6 a. b. c. d. e. 8 mm 15 mm 0.3 mm 0.6 mm (simple thalloid) (complex thalloid) 7 (apical cell) M. Shimamura-8 BSJ-Review 3:91 (2012)

37 植物科学最前線 3:92 (2012) 4 8 9, (Crandall-Stotler 1981) (secondary apical cell) (air chamber) 6 (air pore) 11 (Walker & Pennington 1939) ; assimilatory filament 11 (Meyer et al. 2008) 2 ; Hofmeister 1851 ; Leitgeb1880 Apostolakos & Galatis (1982) 12a c ( ; parenchymatous tissue) M. Shimamura-9 BSJ-Review 3:92 (2012)

38 植物科学最前線 3:93 (2012) ( 13) (oil body) ; idioblast, oil cell Suire et al (ventral scale) (rhizoid) ( 13) (pegged rhizoid tuberculate rhizoid) (smooth rhizoid) (Kammerling 1897, Schiffner1909, 1977) µm ( 14a, b) (McConaha 1941) µm 14c 3 M. polymorpha ssp. montivagans (Ligrone et al. 2007) (Kutschera et al. 2007) 1 15a, b 15c 15d (McConaha 1941, 1977) 14b 4 15d M. Shimamura-10 BSJ-Review 3:93 (2012)

39 植物科学最前線 3:94 (2012) 7 8. a. b. c. d. M. Shimamura-11 BSJ-Review 3:94 (2012)

40 植物科学最前線 3:95 (2012) 9 a. b c. d. M. Shimamura-12 BSJ-Review 3:95 (2012)

41 植物科学最前線 3:96 (2012) M. Shimamura-13 BSJ-Review 3:96 (2012)

42 植物科学最前線 3:97 (2012) 14. a. b. c. 15. a. b. 3 6 c. d. ; gemma cup 15a M. Shimamura-14 BSJ-Review 3:97 (2012)

43 植物科学最前線 3:98 (2012) 1 16a, b 16c 16d (splash cup) 120 cm (Equihua 1987) 16. a. b. c. d. (antheridiophore) (archegoniophore) 17, 18 (antheridial receptacle) (antheridium) M. Shimamura-15 BSJ-Review 3:98 (2012)

44 植物科学最前線 3:99 (2012) 8 17 a, c 17b 2 17c 17b, d (archegonial receptacle) 8-10 (digitate ray) 17a (archegonium) 18b) 2 ( 18c) 18d 17. a. b. c. d. M. Shimamura-16 BSJ-Review 3:99 (2012)

45 植物科学最前線 3:100 (2012) 18. a. b. c. d. 1 19a 19b 1 (spermatogenous tissue, androgonial tissue) (spermatogenous cell) 19c (sperm cell) (sperm mother cell) 19d ; polar M. Shimamura-17 BSJ-Review 3:100 (2012)

46 植物科学最前線 3:101 (2012) organizer (Brown & Lemmon 1990, Shimamura et al. 2004) 1 (centriole) (centrosome) (Carothers & Kreitner 1968) ; basal body Moser & Kreitner a d 2 20e, f (Yuasa 1952) DNA (Izumi & Ono 1999, Shimamura et al. 1999) (Pacak & Szweykowska-Kulińska 2003, McDaniel et al. 2007, Jankowiak-Siuda et al. 2008) 9+2 (Carothers & Duckett 1980, Renzaglia & Garbary 2001) 21a (cover cell) 4 (neck canal cell) ventral canal cell (egg) ( 21b) 21c, d 6 21d 5 (venter) (neck) 21e (pseudoperianth) M. Shimamura-18 BSJ-Review 3:101 (2012)

47 植物科学最前線 3:102 (2012) 20. a. b. c. d. e. f. a e. Carothers & Kreitner (1968), Shimamura et al. (1999) M. Shimamura-19 BSJ-Review 3:102 (2012)

48 植物科学最前線 3:103 (2012) 21. a. b. c. d. 6 e. 22a, b 22c 20 cm (Shimamura et al. 2008) Duckett & Pressel (2009) 30 cm 10 cm M. Shimamura-20 BSJ-Review 3:103 (2012)

49 植物科学最前線 3:104 (2012) (Parihar 1956) 23 1 mm 22. a. ( ) b. c. M. Shimamura-21 BSJ-Review 3:104 (2012)

50 植物科学最前線 3:105 (2012) a 24b amphithecium) (endothecium) ; columella (pseudoperianth) ( ; female involucre) (Parihar 1956) (calyptra) M. Shimamura-22 BSJ-Review 3:105 (2012)

51 植物科学最前線 3:106 (2012) 3 ( 24c, d, 26a) (bract) (perianth) 24d (foot) (placenta) (sporogenous tissue) (elater mother cell, elaterocyte) sporogenous cell (elater cell) (spore mother cell, sporocyte) 24e (Brown et al. 2007, 2010, Shimamura et al. 2012) 25a 25b 26a 300,000 7,000,000 (O Hanlon 1926) 100 (Duckett & Pressel 2009) 26b (elater) 26c 26d M. Shimamura-23 BSJ-Review 3:106 (2012)

52 植物科学最前線 3:107 (2012) 24. a. 4 b. c. d. e. 25. a. P: N: b. - M. Shimamura-24 BSJ-Review 3:107 (2012)

53 植物科学最前線 3:108 (2012) 26. a. b. c. 2 d. M. polymorpha ( ) (1987) M. polymorpha var. aquatica ssp. polymorpha (nucleolar organizer) (Bischler 1986) M. polymorpha ssp. montivagans (Perold 1999) Schuster 1992 (Konstantinova & Bakalin 2009) (Choi et al. 2010) 1987, Schuster 1992 M. Shimamura-25 BSJ-Review 3:108 (2012)

54 植物科学最前線 3:109 (2012) 200 Apostolakos, P., & Galatis, B Studies on the development of the air pores and air chambers of Marchantia paleacea. Protoplasma 128: Bischler, H Marchantia polymorpha L. s. lat. karyotype analysis. J. Hattori Bot. Lab. 60: Bischler, H Marchantia L. The Asiatic and Oceanic taxa. Bryophytorum Bibliotheca 38: Bischler, H Systematics and evolution of the genera of the Marchantiales. Bryophytorum Bibliotheca 51: Bischler-Causse, H., & Boisselier-Dubayle, M. C Lectotypification of Marchantia polymorpha L. J. Bryol. 16: Boisselier-Dubayle, M. C., & Bischler, H Electrophoretic studies in Marchantia polymorpha L. J. Hattori Bot. Lab. 67: Boisselier-Dubayle, M. C., Jubier, M. F., Lejeune, B., & Bischler, H Genetic variability in the three subspecies of Marchantia polymorpha (Hepaticae): isozymes, RFLP and RAPD markers. Taxon 44: Boisselier-Dubayle, M. C., Lambourdiere, J., & Bischler, H Molecular phylogenies support multiple morphological reductions in the liverwort subclass Marchantiidae (Bryophyta). Mol. Phylogenet. Evol. 24: Brown, R. C., & Lemmon, B. E Polar organizers mark division axis prior to preprophase band formation in mitosis of the hepatic Reboulia hemisphaerica (Bryophyta). Protoplasma 156: Brown R. C., Lemmon B. E., & Shimamura, M Transformations of the pleiomorphic plant MTOC during sporogenesis in the hepatic Marchantia polymorpha. J. Integr. Plant Biol. 49: Brown, R. C., Lemmon, B. E., & Shimamura, M Diversity in meiotic spindle origin and determination of cytokinetic planes in sporogenesis of complex thalloid liverworts (Marchantiopsida) J. Plant Res.123: Burgeff, H Genetische Studien an Marchantia. Gustav Fischer, Jena, Germany. Carothers, Z. B., & Duckett, J. G The bryophyte spermatozoid: a source of new phylogenetic information. Bull. Torrey Bot. Club 107: Carothers, Z. B., & Kreitner, L Studies of spermatogenesis in the Hepaticae. II. Blepharoplast M. Shimamura-26 BSJ-Review 3:109 (2012)

55 植物科学最前線 3:110 (2012) Structure in the spermatid of Marchantia. J. Cell Biol. 36: Choi, S. S., Bakalin V. A., & Sun, B. Y Unrecorded liverwort species from Mt. Deogyu, Korea. Kor. J. Plant Tax. 40: Crandall-Stotler, B Morphology/anatomy of hepatics and anthocerotes. In: Schultze-Motel W. (ed.) Advances in Bryology I. pp J. Cramer, Vaduz. Crandall-Stotler, B., & Stotler. R. E. & Long, D. G Phylogeny and Classification of the Marchantiophyta. Edinb. J. Bot. 66: Dillenius, J. J Historia muscorum. Theatro Sheldoniano, Oxford, UK. Duckett, J. G., & Pressel, S Extraordinary features of the reproductive biology of Marchantia at Thursley NNR. Field Bryol. 97: Duckett, J. G.. & Renzaglia, K. S The reproductive biology of the liverwort Blasia pusilla L. J. Bryol. 17: Equihua, C Splash-cup dispersal of gemmae in the liverwort Marchantia polymorpha. Cryptogamie Bryologie-Lichénologie 8: Forrest, L. L., Davis, E. C., Long, D. G., Crandall-Stotler, B. J., Clark, A., & Hollingsworth. M. L Unraveling the evolutionary history of the liverworts (Marchantiophyta) Multiple taxa, genomes, and analyses. Bryologist 109: Graham, L. E., Wilcox, L. W., Cook, M. E., & Gensel, P. G Resistant tissues of modern marchantioid liverworts resemble enigmatic Early Paleozoic microfossils. Proc. Natl. Acad. Sci. USA. 101: He-Nygrén, X., Juslén, A., Ahonen, I., Glenny, D., & Piippo, S Illuminating the evolutionary history of liverworts (Marchantiophyta) towards a natural classification. Cladistics 22: Hofmeister, W Vergleichende Untersuchungen der Keimung, Entfaltung und Fruchtbildung hbherer Kryptogamen und der Samenbildung der Gymnospermen. Leipzig. Inoue, H Studies in spore germination and the earlier stages of gametophyte development in the Marchantiales. J. Hattori Bot. Lab. 23: Izumi, Y., & Ono, K Changes in plastid DNA content during the life cycle of the hornwort Anthoceros punctatus L. Cytologia 64: Jankowiak-Siuda, K., Pacak, A., Odrzykoski, I., Wyatt, R. & Szweykowska-Kulińska, Z Organellar inheritance in the allopolyploid mosses species Plagiomnium medium and Plagiomnium curvatulum. Taxon 57: : : : Kammerling, Z Zur Biologic und Physiologie der Marchantiaceen. Flora 84: Kny, L Bau und Entwicklung von Marchantia polymorpha L. Botanische Wandtafeln 8: Paul Parey, Berlin. BSJ Review 3: M. Shimamura-27 BSJ-Review 3:110 (2012)

56 植物科学最前線 3:111 (2012) Konstantinova, N. A., & Bakali, V. A Checklist of liverworts (Marchantiophyta) of Russia. Arctoa 18: Kutschera, U., Thomas, J., & Hornschuh, M Cluster formation in liverwort- associated methylobacteria and its implications. Naturwissenschaften 94: Leitgeb, H., Die Athemöffnung der Marchantiaceen. Sitz-Ber. d. Kaiserl. Akademie d. Wiss. 81: Linnaeus, C Species plantarum. Ed. 1. Salvius, Stockholm. Ligrone, R., Carafa, A., Lumini, E., Bianciotto, V., Bonfante, P., & Duckett, J. G Glomeromycotean associations in liverworts: a molecular, cellular, and taxonomic analysis. Amer. J. Bot. 94: Long, D. G New higher taxa of complex thalloid liverworts (Marchantiophyta Marchantiopsida). Edinb. J. Bot. 63: McConaha, M Ventral structures effecting capillarity in the Marchantiales. Amer. J. Bot. 28: McDaniel, S. F, Willis, J. H., & Shaw, A. J A linkage map reveals a complex basis for segregation distortion in an interpopulation cross in the moss Ceratodon purpureus. Genetics 176: Meyer, M., Seibt, U. & Griffiths, H To concentrate or ventilate? Carbon acquisition, isotope discrimination and physiological ecology of early land plant life forms. Phil. Trans. R. Soc. B. 363: Moser, J. W., & Kreitner, G. L Centrosome structure in Anthoceros laevis and Marchantia polymorpha. J Cell Biol. 44: Nakazato, T., Kadota, A., & Wada, M Photoinduction of spore germination in Marchantia polymorpha L. is mediated by photosynthesis. Plant Cell Physiol. 40: O'Hanlon, M. E Germination of the spores and early stages in the development of the gametophyte of Marchantia polymorpha. Bot. Gaz. 82: Pacak, A. & Szweykowska-Kulińska, Z Organellar inheritance in liverworts: an example of Pellia borealis. J Mol Evol. 56:11-7. Parihar, N. S An Introduction to Embryophyta. Volume I. Bryophyta, Central Book Depot. Allahabad, India. Paton, J. A The Liverwort Flora of the British Isles. Harley Books, Colchester. UK. Perold, S. M Flora of Southern Africa. Hepatophyta. Part 1: Marchantiopsida. In: O.A.Leistner (ed.), Flora of southern Africa: Botanical Research Institute, Pretoria. Qiu, Y.-L., Li, L., Wang, B., Chen, Z.. Knoop, V.. Groth-Malonek, M., Dombrovska, O., Lee, J., Kent. L., Rest, J., Estabrook, G. F., Hendry, T. A., Taylor, D. W., Testa, C. M., Ambros, M., Crandall-Stotler, B., Duff, R. J., Stech, M., Frey, W., Quandt, D., & Davis C. C The deepest divergences in land plants inferred from phylogenomic evidence. Proc. Natl. Acad. Sci. USA 103: Qiu, Y.-L., Li, L., Wang, B., Chen, Z., Dombrovska, O., Lee, J., Kent, L., Li, R., Jobson, R. W., Hendry, T. A., Taylor, D. W., Testa, C. M., & Ambros, M A nonflowering land plant phylogeny inferred M. Shimamura-28 BSJ-Review 3:111 (2012)

57 植物科学最前線 3:112 (2012) from nucleotide sequences of seven chloroplast, mitochondrial, and nuclear genes. Int. J. Plant Sci. 168: Renzaglia, K.S., & Garbary, D. J Motile gametes of land plants: diversity, development, and evolution. Crit Rev Plant Sci. 20: Schiffner, V Studien über die Rhizoiden der Marchantiales. Ann. Jard. Bot. Buitenz. 2 ser., Suppl. 3: Schuster, R Phytogeography of the Bryophyta. In New manual of bryology. Edited by R.M. Schuster. Hattori Bot. Lab., Nichinan, Japan. Schuster, R. M The Hepaticae and Anthocerotae of North America, Vol. 6. Field Museum of Natural History. Chicago. Shimamura, M., Fukushima, H., Yamaguchi, T., & Deguchi, H Behavior of plastid and plastid DNA during spermiogenesis in Dumortiera hirusta. Bryol. Res. 7: Shimamura, M., Brown, R. C., Lemmon, B. E., Akashi, T., Mizuno, K., Nishihara, N. Tomizawa K. I., Yoshimoto, K., Deguchi, H., Hosoya, H., Horio, T., & Mineyuki, Y γ-tubulin in basal land plants: characterization, localization, and implication in the evolution of acentriolar microtubule organizing centers. Plant Cell 16: Shimamura, M., Furuki, T., & Deguchi, H Sporophyte anatomy of Cavicularia densa Steph. (Blasiaceae). Bryologist. 108: Hikobia 14: Shimamura, M., Yamaguchi, T., & Deguchi, H Airborne sperm of Conocephalum conicum (Conocephalaceae). J. Plant Res. 121: Shimamura, M., Itouga, M., & Tsubota, H Evolution of apolar sporocytes in marchantialean liverworts: implications from molecular phylogeny. J. Plant Res. 125: Stotler, R [Review of the book Marchantia L. The European and African Taxa. Bryophytrum Bibliotheca 45 by Bishler, H.] Bryologist 97: Suire, C., Bouvier, F., Backhaus, R. A., Bégu, D., Bonneu, M., & Camara, B Cellular localization of isoprenoid biosynthetic enzymes in Marchantia polymorpha. uncovering a new role of oil bodies. Plant Physiol. 124: Völker, H, Licht- und elektronenmikroskopische Untersuchungen zur Cytokinese an jungen Brutkörpern vonmarchantia polymorpha L. Protoplasma, 75: Walker, R., & Pennington, W The movements of the air pores of Preissia quadrata (Scop.). New Phytologist 38: Walton, J Carboniferous bryophytes. I. Hepaticae. Ann. Bot. 39: Wellman, C. H., Osterff, P. L., & Mohiuddin, U Fragments of the earliest land plants, Nature 425: Wheeler, J. A Molecular phylogenetic reconstructions of the marchantioid liverwort radiation. Bryologist 103: M. Shimamura-29 BSJ-Review 3:112 (2012)

58 植物科学最前線 3:113 (2012) Yuasa, A Studies in the cytology of Pteridophyta XXX. The behavior of the plastid during the process of fertilization. Sci. Pap. Coll. Gen. Edu. Univ. Tokyo 2: M. Shimamura-30 BSJ-Review 3:113 (2012)

59 植物科学最前線 3:114 (2012) Cell biology in liverwort Key words: Cytoskeleton, Membrane traffic, Organelle, Rab5, VAMP727 Atsuko Era & Takashi Ueda Department of Biological Sciences, Graduate School of Science, The University of Tokyo 7-3-1, Hongo, Bunkyo-ku, Tokyo, , Japan etc. Rab GTPase SNARE 1 Rab GTPase SNARE Rab GTPase SNARE A. Era & T. Ueda-1 BSJ-Review 3:114 (2012)

60 植物科学最前線 3:115 (2012) Rab GTPase SNARE 2 (Dacks & Field 2007) RAB5 Rab GTPase Rab5 Dacks & Field 2007 Rab5 (Grosshans et al. 2006) RHA1 ARA7 ARA6 3 Rab5 RHA1 ARA7 Rab5 RAB5 Rab5 ARA6 Rab5 RAB5 ARA6 Bars = 20 µm RAB5 ARA6 RAB5 ARA6 RAB5 ARA6 multivesicular endosome: MVE (Ueda et al. 2001) RAB5 ARA6 RAB5 ARA6 (Ebine et al. 2011) ARA6 A. Era & T. Ueda-2 BSJ-Review 3:115 (2012)

61 植物科学最前線 3:116 (2012) ARA6 ARA6 ARA6 RAB5 RAB5 MpARA6 MpARA6 ARA6 MpARA6 3a MpARA6 MpARA6 erasome 3b RAB5 SNARE VAMP727 Rab GTPase R-SNARE Q-SNARE SNARE VAMP7 R-SNARE VAMP7 VAMP71 VAMP72 2 VAMP72 VAMP727 VAMP727 longin R-SNARE VAMP727 EST MpVAMP727 VAMP72 VAMP727 VAMP72 MpVAMP727 VAMP727 (Hasezawa & Kumagai 2002, Hussey et al. 2006, Tijs Ketelaar 2001) Lifeact (Riedl et al. 2008) (Era et al. 2009) 4 in vitro (Michelot et al. 2007, Staiger et al. 2009) A. Era & T. Ueda-3 BSJ-Review 3:116 (2012)

62 植物科学最前線 3:117 (2012) Bar = 20 µm Bar = 2 µm Bar = 5 µm Era et al. in press 5 5 mm SP-GFP-HDEL syalyl transferase (ST) Bar = 20 µm A. Era & T. Ueda-4 BSJ-Review 3:117 (2012)

63 植物科学最前線 3:118 (2012) TGN TGN Q-SNARE SYP61 EST MpSYP6 ST TGN MpSYP6 Bar = 20 µm Q-SNARE SYP11 SYP12 SYP13 EST MpSYP1 Bar = 20 µm Q-SNARE SYP2 EST MpSYP2 A. Era & T. Ueda-5 BSJ-Review 3:118 (2012)

64 植物科学最前線 3:119 (2012) Bar = 20 µm Lifeact FM1-43 Bar = 20 µm Lifeact Mito Tracker Bar = 20 µm A. Era & T. Ueda-6 BSJ-Review 3:119 (2012)

65 植物科学最前線 3:120 (2012) 4 TGN ARA6 RAB5 ARA6 RAB5 ARA6 RAB ARA6 RAB5 VAMP727 EST VAMP727 VAMP727 (Ebine et al. 2011, Ebine et al. 2008) VAMP727 VAMP727 A. Era & T. Ueda-7 BSJ-Review 3:120 (2012)

66 植物科学最前線 3:121 (2012) Dacks, J.B., & Field, M. C Evolution of the eukaryotic membrane-trafficking system: origin, tempo and mode. J Cell Sci. 120: Ebine, K., Fujimoto, M., Okatani, Y., Nishiyama, T., Goh, T., Ito, E., Dainobu, T., Nishitani, A., Uemura, T., Sato, M.H., Thordal-Christensen, H., Tsutsumi, N., Nakano, A., & Ueda, T A membrane trafficking pathway regulated by the plant-specific RAB GTPase ARA6. Nat Cell Biol. 13: Ebine, K., Okatani, Y., Uemura, T., Goh, T., Shoda, K., Niihama, M., Morita, M. T., Spitzer, C., Otegui, M. S., Nakano, A., & Ueda, T A SNARE complex unique to seed plants is required for protein storage vacuole biogenesis and seed development of Arabidopsis thaliana. Plant Cell. 20: Era, A., Kutsuna, N., Higaki, T., Hasezawa, S., Nakano, A., & Ueda, T Microtubule stability affects the unique motility of F-actin in Marchantia polymorpha. J. Plant Res. in press. Era, A., Tominaga, M., Ebine, K., Awai, C., Saito, C., Ishizaki, K., Yamato, K. T., Kohchi, T., Nakano, A., & Ueda, T Application of Lifeact reveals F-actin dynamics in Arabidopsis thaliana and the liverwort, Marchantia polymorpha. Plant Cell Physiol. 50: Grosshans, B. L., Ortiz, D., & Novick, P Rabs and their effectors: Achieving specificity in membrane traffic. Proc Natl Acad Sci U S A. 103: Hasezawa, S., & Kumagai, F Dynamic changes and the role of the cytoskeleton during the cell cycle in higher plant cells. Int Rev Cytol. 214: Hussey, P. J., Ketelaar, T., & Deeks, M. J Control of the actin cytoskeleton in plant cell growth. Annu Rev Plant Biol. 57: Michelot, A., Berro, J., Guerin, C., Boujemaa-Paterski, R., Staiger, C. J., Martiel, J. L., & Blanchoin, L Actin-filament stochastic dynamics mediated by ADF/cofilin. Curr Biol. 17: Riedl, J., Crevenna, A. H., Kessenbrock, K., Yu J. H., Neukirchen, D., Bista, M., Bradke, F., Jenne, D., Holak, T. A., Werb, Z., Sixt, M., & Wedlich-Soldner, R Lifeact: a versatile marker to visualize F-actin. Nat Methods. 5: Staiger, C. J., Sheahan, M. B., Khurana, P., Wang, X., McCurdy, D. W. & Blanchoin, L Actin filament dynamics are dominated by rapid growth and severing activity in the Arabidopsis cortical array. J Cell Biol. 184: Tijs Ketelaar, A. M. C. E The cytoskeleton in plant cell growth: lessons from root hairs. New Phytologist. 152: Ueda, T., Yamaguchi, M., Uchimiya, H., & Nakano, A Ara6, a plant-unique novel type Rab GTPase, functions in the endocytic pathway of Arabidopsis thaliana. EMBO J. 20: A. Era & T. Ueda-8 BSJ-Review 3:121 (2012)

67 植物科学最前線 3:122 (2012) The role of abscisic acid in stress response and dormancy of liverworts Key words: abscisic acid; desiccation stress; protein phosphatase 2C. Daisuke Takezawa Graduate School of Science and Engineering; Institute for Environmental Science and Technology, Saitama University Sakura-ku, Saitama, , Japan ABA ABA ABA Cutler et al ABA Hartung 2010 ABA Embryophyta ABA ABA Physcomitrella patens Marchantia polymorpha ABA Takezawa et al ABA D. Takezawa-1 BSJ-Review 3:122 (2012)

68 植物科学最前線 3:123 (2012) Oliver et al ABA ABA ABA ABA ABA ABA Hartung et al. 1987, Werner et al ABA Funaria hygrometrica Anthoceros punctatus Garner and Paolillo 1973, Hartung et al ABA ABA brood cell ABA Valadon and Mummery 1971, Menon and Lal 1974, Goode et al. 1993, Chopra and Kapur 1989 ABA ABA Werner et al ABA Goode et al. 1993, Minami et al ABA Cuming et al. 2007, Komatsu et al. 2009, D. Takezawa-2 BSJ-Review 3:123 (2012)

69 植物科学最前線 3:124 (2012) Chater et al ABA, Hellwege Hartung Marchatiales Exormotheca holstii Riccia fluitans ABA Hellwege et al. 1994, 1996 ABA LEA Hellwege et al Pallavicinia lyellii ABA Pence et al ABA Pence et al Pryce 1972 ABA ABA LNA ABA Pryce 1972 LNA Lunularia cruciata Valio and Schwabe 1970 LNA ABA D. Takezawa-3 BSJ-Review 3:124 (2012)

70 植物科学最前線 3:125 (2012) ABA ABA ABA ABA Tougane et al. 2010, Akter et al ABA ABA Em GUS Em-GUS ABA ABA EST ABSCISIC ACID INSENSITIVE1 (ABI1) MpABI1 ABI1 2C PP2C Gly Asp abi1-1 ABA ABI1 A PP2C PP2C ABA A PP2C ABA PYR/PYL/RCAR SnRK2 ABA Park et al MpABI1 A PP2C C A N PP2C GFP MpABI1 MpABI1 ABA Em-GUS MpABI1 ABA ABA MpABI1 ABA LEA Tougane et al D. Takezawa-4 BSJ-Review 3:125 (2012)

71 植物科学最前線 3:126 (2012) MpABI1 ABA ABA Bhyan et al PP2C ABA ABA ABA ABA ABA Ishizaki (2008) ABA ABA ABA ABA ABA (C) PP2C Akter, K., Tougane, K., Kaneko, M., & Takezawa, D Effect of sugar and abscisic acid on freezing and desiccation tolerance in the liverwort Marchantia polymorpha. Cryobiol. Cryotechnol. 57: Bhyan, S. B., Minami, A., Kaneko, Y., Suzuki, S., Arakawa, K., Sakata, Y., & Takezawa, D Cold acclimation in the moss Physcomitrella patens involves abscisic acid-dependent signaling. J. Plant Physiol. 169: Chater, C., Kamisugi, Y., Movahedi, M., Fleming, A., Cuming, A. C., Gray, J. E., & Beerling, D. J Regulatory mechanism controlling stomatal behavior conserved across 400 million years of land plant evolution. Curr. Biol. 21: Chopra, R. N., & Kapur, A Effect of abscisic acid and kinetin on protonemal differentiation in Timmiella anomala. Plant Sci. 61: Cuming, A. C., Cho, S. H., Kamisugi, Y., Graham, H., & Quatrano, R. S Microarray analysis of transcriptional responses to abscisic acid and osmotic, salt, and drought stress in the moss, Physcomitrella patens. New Phytol. 176: Cutler, S. R., Rodriguez, P. L., Finkelstein, R. R., & Abrams, S. R Abscisic acid: emergence of a D. Takezawa-5 BSJ-Review 3:126 (2012)

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74 植物科学最前線 3:129 (2012) MpCLE An analysis of MpCLE genes in Marchantia polymorpha Key words: CLE, Hormone, Peptide, Development, Marchantia polymorpha Shinichiro Sawa, Hiroaki Honda, Ryo Tabata Kumamoto University Kurokami , Kumamoto, , Japan CLE (CLAVATA3/Embryo-Surrounding-Region), CLE CLAVATA3 (CLV3) tracheary element differentiation inhibitory factor (TDIF; CLE41/CLE44) (Betsuyaku et al. 2011a; Miwa et al. 2009a) CLE 100 N C CLE (Ohyama et al. 2010) CLE Oelkers et al CLE CLE CLE Oelkers et al CLE CLE CLE CLE (Sawa et al. 2008) 15 CLE (Miwa et al. 2009b) CLE CLE CLE S. Sawa - 1 BSJ-Review 3:129 (2012)