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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

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AZIX ʃƾœ 2012. ǒȌĖƻ*ʳÑĒıLEAFY&[gTQ*ƻźȘǣ. 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.

植物科学最前線 3: 68 ( 2012)

BSJ- Revi ew 3: 68 ( 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. ì£ȚŇĬ�ǭÎĳ� 2008. ÈāƅȱɀǮ 5-3b [gTQ*Ğ˻ɖɵ&ǒȌ®*r~X_eŪʯʳƞǮ. �� 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.

ȰŔËŦ�ǭÎĳ� 2008. ÈāƅȱɀǮ 5-3a DP�lOb�Gy):=[gTQǐRjyŪʯʳƞǮ. �� 67: 597-600.

żɽóı��Țʭŵ 2012. [gTQɖɵȘȌĴ 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.

植物科学最前線 3: 69 ( 2012)

BSJ- Revi ew 3: 69 ( 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.

Ǿ˂�ˍ�ǀȚɓɃ�Țȝ� 2012. MpCLE1ˋ¨ı*ǚɶʙǈ. BSJ Review 3: 129-133. ŕǂǜǙ 2012. [gTQ*Ô˸Ĵ&ŪƂĴ. 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.

Ĭċçş�ȰŔËŦ�ǭÎĳ� 2008. ÈāƅȱɀǮ1-4 [gTQ*Ğ˻Ǯ. �� 67: 23-30. Ĭċçş�ǭÎĳ� 2012. [gTQRjy. BSJ Review 3: 71-83.

植物科学最前線 3: 70 ( 2012)

BSJ- Revi ew 3: 70 ( 2012)

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