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植 物 科 学 最 前 線 3:58 (2012) 606-8502 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 606-8502, Japan 19 Evo Devo Evo Eco 2008 - Marchantia polymorpha L. 4 Qiu et al. 2006 2012 T. Kohchi & K. Ishizaki-1 BSJ-Review 3:58 (2012)

植 物 科 学 最 前 線 3:59 (2012) - 1 Kny 1890 WEB http://www.geheugenvannederland.nl/ 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)

植 物 科 学 最 前 線 3:60 (2012) - 8 X Y 2012 Wann 1925 Lloyd & Steinmetz 1937 Chiyoda et al. 2008 2012 2012 2 2. T. Kohchi & K. Ishizaki-3 BSJ-Review 3:60 (2012)

植 物 科 学 最 前 線 3:61 (2012) 30 Spore Sporelings 2-3 Gemma Gemmalings Gemmalings Sporelings 1 2012 Kubota et al. 2012 1 1 3 - T. Kohchi & K. Ishizaki-4 BSJ-Review 3:61 (2012)

植 物 科 学 最 前 線 3:62 (2012) 3. Barnes & Land (1908) 2012 3 DNA - n=8+ 2012 T. Kohchi & K. Ishizaki-5 BSJ-Review 3:62 (2012)

植 物 科 学 最 前 線 3:63 (2012) 18 2012 19 20 2012 1943 Burgeff Genetische Studien an Marchantia Burgeff 1943 70 1970 Ono 1976 Ohyama et al. 1986, Oda et al. 1992 Takaragaike-1 Takaragaike-2 (Okada et al. 2000) Marchantia polymorpha ssp. ruderalis Marchantia polymorpha Takaragaike-1 Takaragaike-2 PAC Okada et al. 2000 Okada et al. 2001, Ishizaki et al. 2002, Yamato et al. 2007 Takaragaike-1 Takaragaike-2 2006 Takaragaike-1 Takaragaike-2 Joint Genome Institute http://www.jgi.doe.gov/sequencing/why/99191.html, 2012 Kitashirakawa Takaragaike-1 8 8 F1 Recombinant Inbred - T. Kohchi & K. Ishizaki-6 BSJ-Review 3:63 (2012)

植 物 科 学 最 前 線 3:64 (2012) 4. Sporelings Ishizaki et al. 2008 5 Sporelings 2 4 1 1,000 2008 Sporelings T-DNA T-DNA T-DNA 70,000 90% 20 T-DNA T-DNA 20 10 TAIL-PCR ARF+ 2 Sporelings T. Kohchi & K. Ishizaki-7 BSJ-Review 3:64 (2012)

植 物 科 学 最 前 線 3:65 (2012) Sandy Floyd T1 T2 T3 T1 T1 gemma G1 G1 G1 G2 5 5. T. Kohchi & K. Ishizaki-8 BSJ-Review 3:65 (2012)

植 物 科 学 最 前 線 3:66 (2012) - Terada et al. 2007 DTA T-DNA T-DNA 2011 20 PCR 2 200 DNA PCR - 120 DNA Boynton et al. 1984 Chiyoda et al. 2007, Chiyoda et al. 2008, 2008 Ueda et al. 2012 DNA DNA Sporelings Chiyoda et al. 2008 EST Nishiyama et al. 2003 Rensing et al. 2008, Banks et al. 2011 2012 T. Kohchi & K. Ishizaki-9 BSJ-Review 3:66 (2012)

植 物 科 学 最 前 線 3:67 (2012) PIF Leivar et al. 2008 PIF Burgeff (1943) EMS phot T-DNA 2012 T. Kohchi & K. Ishizaki-10 BSJ-Review 3:67 (2012)

植 物 科 学 最 前 線 3:68 (2012) 2012, 2012 accession Joint Genome Institute Monash John Bowman ( ) 2012. LEAFY. BSJ Review. Banks, J. A., Nishiyama, T., Hasebe, M. et al. 2011. The Selaginella genome identifies genetic changes associated with the evolution of vascular plants. Science 332: 960-963. Barnes, C. R., & Land, W. J. G. 1908. The origin of the cupule of Marchantia. Bot. Gazette 46: 401-409. 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. 1988. Chloroplast transformation in Chlamydomonas with high velocity microprojectiles. Science 240: 1534-1538. Burgeff, H. 1943. Genetische Studien an Marchantia. Verlag von Gustav Fischer, Jena. Chiyoda, S., Linley, P. J., Yamato, K. T., Fukuzawa, H., Yokota, A., & Kohchi, T. 2007. 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)

植 物 科 学 最 前 線 3:69 (2012) 16: 41-49. Chiyoda, S., Ishizaki, K., Kataoka, H., Yamato, K. T., & Kohchi, T. 2008. Direct transformation of the liverwort Marchantia polymorpha L. by particle bombardment using immature thalli developing from spores. Plant Cell Rep. 27: 1467-1473. 2008. 5-3b. 67: 601-606. Ishizaki, K., Shimizu-Ueda, Y., Okada, S., Yamamoto, M., Fujisawa, M., Yamato, K.T., Fukuzawa, H., & Ohyama, K. 2002. Multicopy genes uniquely amplified in the Y chromosome-specific repeats of the liverwort Marchantia polymorpha. Nucl. Acids Res. 30: 4675-4681. Ishizaki, K., Chiyoda, S., Yamato, K. T., & Kohchi, T. 2008. Agrobacterium-mediated transformation of the haploid liverwort Marchantia polymorpha L., an emerging model for plant biology. Plant Cell Physiol. 49: 1084-1091. 2008. 5-3a. 67: 597-600. 2012. BSJ Review 3: 114-121. Kny, L. 1890. Bau und Entwickelung von Marchantia polymorpha L.In Botanische Wandtafeln mit Erläuterndem Text pp. 364-401. Verlag von Wiegandt, Hempel & Parey, Berlin. Kubata, A., Ishizaki, K., Hosaka, M., & Kohchi, T. 2012. 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. 2008. Multiple phytochrome-interacting bhlh transcription factors repress premature photomorphogenesis during early seedling development in darkness. Current Biol. 18: 1815-1823. Lloyd, C.E., & Steinmetz, F.H. 1937. Temperature as a factor influencing the sexual response of Marchantia. Americ. J. Bot. 24: 423-425. 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. 2003. Comparative genomics of Physcomitrella patens gametophytic transcriptome and Arabidopsis thaliana: implication for land plant evolution. Proc. Natl. Acad. Sci. USA. 100: 8007-8012. 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. 1986. Chloroplast gene organization deduced from complete sequence of liverwort Marchantia polymorpha chloroplast DNA. Nature 322: 572-574. 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. 2000. Construction of male and female PAC genomic libraries suitable for identification of Y-chromosome-specific clones from the liverwort, Marchantia polymorpha, Plant J. 24: 421-428. T. Kohchi & K. Ishizaki-12 BSJ-Review 3:69 (2012)

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. 2001. The Y chromosome in the liverwort Marchantia polymorpha has accumulated unique repeat sequences harboring a male-specific gene. Proc. Natl. Acad. Sci. USA. 98: 9454-9459. Ono, K. 1976. Cytological observations of calluses and restored thalluses in Marchantia polymorpha. Jpn. J. Genet. 51: 11-18. 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. 2006. The deepest divergences in land plants inferred from phylogenomic evidence. Proc. Natl. Acad. Sci. USA 103: 15511-15516. Rensing, S. A., Lang, D., Zimmer, A. D. et al. 2008. The Physcomitrella genome reveals evolutionary insights into the conquest of land by plants. Science 319: 64-69. Sakakibara, K., Nishiyama, T., Deguchi, H., & Hasebe, M. 2008. Class 1 KNOX genes are not involved in shoot development in the moss Physcomitrella patens but do function in sporophyte development. Evol. Dev. 10: 555-566. Sano, R., Juárez, C.M., Hass, B., Sakakibara, K., Ito, M., Banks, J.A., & Hasebe, M. 2005. KNOX homeobox genes potentially have similar function in both diploid unicellular and multicellular meristems, but not in haploid meristems. Evol. Dev. 7: 69-78. 2012. MpCLE1. BSJ Review 3: 129-133. 2012.. BSJ Review 3: 84-113. Terada, R., Johzuka-Hisatomi, Y., Saitoh, M., Asao, H. & Iida, S. 2007. Gene targeting by homologous recombination as a biotechnological tool for rice functional genomics. Plant Physiol. 144: 846-856. Ueda, M., Kuniyoshi, T., Yamamoto, H., Sugimoto K., Ishizaki, K., Kohchi, T., Nishimura, Y., & Shikanai, T. 2012. Composition and physiological function of the chloroplast NADH dehydrogenase-like complex in Marchantia polymorpha. Plant J. 72: 683-693. Wann, F.B. 1925. Some of the factors involved in the sexual reproduction of Marchantia polymorpha. Americ. J. Bot. 12: 307-318. 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. 2007. Gene organization of the liverwort Y chromosome reveals distinct sex chromosome evolution in a haploid system. Proc. Natl. Acad. Sci. USA. 104: 6472-6477. 2008. 1-4. 67: 23-30. 2012.. BSJ Review 3: 71-83. T. Kohchi & K. Ishizaki-13 BSJ-Review 3:70 (2012)