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Light-dependent transcriptional regulation in chloroplasts

Key words: chloroplasts, Cyanidioschyzon merolae, light response, transcriptional regulation, twocomponent system.
Mitsumasa Hanaoka
Graduate School of Horticulture, Chiba University
Matsudo 648, Matsudo, Chiba 271-8510, Japan('  Şŏ%Ƥ§ÄĪ"ĮćŇŔ#J<]^EUl<$Ĵĸ ŇŔĤ7 űĤ
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2004, Nozaki et al. 2007ƣ$G]g%ūų8 6Ƥ>pWop , ;!"ƤƉÅ$Ǝŧ ĉ0"!$§ÄĪ"ĝæ G]gċƂ 51ĳ8ƤdLWG]gİĺ$iVm
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Allen, J.F. 2005. Photosynthesis: the processing of redox signals in chloroplasts. Curr. Biol. 15: R929-932.
Allen, J.F., Santabarbara, S., Allen, C.A., & Puthiyaveetil, S. 2011. Discrete redox signaling pathways
regulate photosynthetic light-harvesting and chloroplast gene transcription. PLoS One 6: e26372.
Ashby, M.K., Houmard, J., & Mullineaux, C.W. 2002. The ycf27 genes from cyanobacteria and eukaryotic
algae: distribution and implications for chloroplast evolution. FEMS Microbiol. Lett. 214: 25-30.
Ashby, M.K., & Mullineaux C.W. 1999. Cyanobacterial ycf27 gene products regulate energy transfer fromphycobilisomes to photosystems I and II. FEMS Microbiol. Lett. 181: 253-260.
Enami, K., Ozawa, T., Motohashi, N., Nakamura, M., Tanaka K., & Hanaoka, M. 2011. Plastid-to-nucleus
retrograde signals are essential for expression of nuclear starch biosynthesis genes during amyloplastdifferentiation in tobacco BY-2 cultured cells. Plant Physiol. 157: 518-530.
M. Hanaoka - 5

BSJ- Revi ew 4: 17( 2013)

植物科学最前線 4: 18( 2013)

Fernández, A.P., & Strand, A. 2008. Retrograde signaling and plant stress: plastid signals initiate cellularstress responses. Curr. Opin. Plant Biol. 11: 509-513.
Fujiwara, T., Kuroiwa, H., Yagisawa, F., Ohnuma, M., Yoshida, Y., Yoshida, M., Nishida, K., Misumi, O.,
Watanabe, S., Tanaka, K., & Kuroiwa, T. 2010. The coiled-coil protein VIG1 is essential for tethering
vacuoles to mitochondria in vacuole inheritance of Cyanidioschyzon merolae. Plant Cell 22: 772-781.
Glöckner, G., Rosenthal, A., & Valentin, K. 2000. The structure and gene repertoire of an ancient red algal
plastid genome. J. Mol. Evol. 51: 382-390.
Goldschmidt-Clermont, M. 1998. Coordination of nuclear and chloroplast gene expression in plant cells. Int.
Rev. Cytol. 177: 115-180.
Hanaoka, M., Kanamaru, K., Takahashi, H., & Tanaka, K. (2003) Molecular genetic analysis of chloroplast
gene promoters dependent on SIG2, a nucleus-encoded sigma factor for the plastid-encoded RNA
polymerase, in Arabidopsis thaliana. Nucleic Acids Res. 31: 7090-7098.
Hanaoka, M., Kawakami, T., & Tanaka, K. 2010. Chloroplast isolation and run-on transcription assay in
Cyanidioschyzon merolae. Endocytobiosis Cell Res. 20: 45-52.
Hanaoka, M., & Tanaka, K. 2008. Dynamics of RpaB-promoter interaction during high light stress, revealed
by chromatin immunoprecipitation (ChIP) analysis in Synechococcus elongatus PCC 7942. Plant J. 56:
327-335.
Imamura, S., Terashita, M., Ohnuma, M., Maruyama, S., Minoda, A., Weber, A.P., Inouye, T., Sekine, Y.,
Fujita, Y., Omata, T., & Tanaka, K. 2010. Nitrate assimilatory genes and their transcriptional regulation ina unicellular red alga Cyanidioschyzon merolae: genetic evidence for nitrite reduction by a sulfite
reductase-like enzyme. Plant Cell Physiol. 51: 707-717.
Ishizaki, Y., Tsunoyama, Y., Hatano, K., Ando, K., Kato, K., Shinmyo, A., Kobori, M., Takeba, G., Nakahira,Y., & Shiina, T. 2005. A nuclear-encoded sigma factor, Arabidopsis SIG6, recognizes sigma-70 typechloroplast promoters and regulates early chloroplast development in cotyledons. Plant J. 42: 133-144.
Kanamaru, K., Nagashima, A., Fujiwara, M., Shimada, H., Shirano, Y., Nakabayashi, K., Shibata, D.,
Tanaka, K., & Takahashi, H. 2001. An Arabidopsis sigma factor (SIG2)-dependent expression of plastidencoded tRNAs in chloroplasts. Plant Cell Physiol. 42: 1034-1043.
Kappel, A.D., & van Waasbergen, L.G. 2007. The response regulator RpaB binds the high light regulatory 1
sequence upstream of the high-light-inducible hliB gene from the cyanobacterium Synechocystis PCC
6803. Arch. Microbiol. 187: 337-342.
Kleine, T., Voigt, C., & Leister, D. 2009. Plastid signalling to the nucleus: messengers still lost in the mists?
Trends Genet. 25: 185-92.
Kobayashi, Y., Imamura, S., Hanaoka, M., & Tanaka, K. 2011. A tetrapyrrole-regulated ubiquitin ligase
controls algal nuclear DNA replication. Nat. Cell Biol. 13: 483-487.
Kobayashi, Y., Kanesaki, Y., Tanaka, A., Kuroiwa, H., Kuroiwa, T., & Tanaka, K. 2009. Tetrapyrrole signal
as a cell cycle coordinator from organelle to nuclear DNA replication in plant cells. Proc. Natl. Acad. Sci.
USA 106: 803-807.
Kuroiwa, T. 2000. The discovery of the division apparatus of plastids and mitochondria. J. Electron Microsc.
M. Hanaoka - 6

BSJ- Revi ew 4: 18( 2013)

植物科学最前線 4: 19( 2013)

49: 123-134.
Leister, D. 2003. Chloroplast research in the genomic age. Trends Genet. 19: 47-56.
Matsuzaki, M. et al. 2004. Genome sequence of the ultrasmall unicellular red alga Cyanidioschyzon merolae
10D. Nature 428: 653-657.
Minoda, A., Nagasawa, K., Hanaoka, M., Horiuchi, M., Takahashi, H., & Tanaka, K. 2005. Microarray
profiling of plastid gene expression in a unicellular red alga, Cyanidioschyzon merolae. Plant Mol. Biol.
59: 375-385.
Minoda, A., Weber, A.P., Tanaka, K., & Miyagishima, S.Y. 2010. Nucleus-independent control of therubisco operon by the plastid-encoded transcription factor Ycf30 in the red alga Cyanidioschyzon merolae.
Plant Physiol. 154: 1532-40.
Misumi, O., Matsuzaki, M., Nozaki, H., Miyagishima, S., Mori, T., Nishida, K., Yagisawa, F., Yoshida, Y.,
Kuroiwa, H., & Kuroiwa, T. 2005. Cyanidioschyzon merolae genome. A tool for facilitating comparable
studies on organelle biogenesis in photosynthetic eukaryotes. Plant Physiol. 137: 567-585.
Nozaki, H., Takano, H., Misumi, O., Terasawa, K., Matsuzaki, M., Maruyama, S., Nishida, K., Yagisawa, F.,
Yoshida, Y., Fujiwara, T., Takio, S., Tamura, K., Chung, S.J., Nakamura, S., Kuroiwa, H., Tanaka, K.,
Sato, N., & Kuroiwa T. 2007. A 100%-complete sequence reveals unusually simple genomic features in
the hot-spring red alga Cyanidioschyzon merolae. BMC Biol. 5: 28.
Ohnuma, M., Misumi, O., Fujiwara, T., Watanabe, S., Tanaka, K., & Kuroiwa T. 2009. Transient genesuppression in a red alga, Cyanidioschyzon merolae 10D. Protoplasma 236: 107-112.
Ohnuma, M., Yokoyama, T., Inouye, T., Sekine, Y., & Tanaka K. 2008. Polyethylene glycol (PEG)-mediatedtransient gene expression in a red alga, Cyanidioschyzon merolae 10D. Plant Cell Physiol. 49: 117-120.
Ohta, N., Matsuzaki, M., Misumi, O., Miyagishima, S.Y., Nozaki, H., Tanaka, K., Shin-I, T., Kohara, Y., &Kuroiwa, T. 2003. Complete sequence and analysis of the plastid genome of the unicellular red alga
Cyanidioschyzon merolae. DNA Res. 10: 67-77.
Ohta, N., Sato, N., & Kuroiwa, T. 1998. Structure and organization of the mitochondrial genome of theunicellular red alga Cyanidioschyzon merolae deduced from the complete nucleotide sequence. Nucleic
Acids Res. 26: 5190-5198.
Pfannschmidt, T., Nilsson, A., & Allen, J.F. 1999 Photosynthetic control of chloroplast gene expression.
Nature 397: 625-628.
Puthiyaveetil, S., Ibrahim, I.M., Jelicić, B., Tomasić, A., Fulgosi, H., & Allen, J.F. 2010. Transcriptional
control of photosynthesis genes: the evolutionarily conserved regulatory mechanism in plastid genome
function. Genome Biol. Evol. 2: 888-896.
Puthiyaveetil, S., Kavanagh, T.A., Cain, P., Sullivan, J.A., Newell, C.A., Gray, J.C., Robinson, C., van derGiezen, M., Rogers, M.B., & Allen. J.F. 2008. The ancestral symbiont sensor kinase CSK links
photosynthesis with gene expression in chloroplasts. Proc. Natl. Acad. Sci. USA 105: 10061-10066.
Reith, M., & Munholland, J. 1995. Complete nucleotide sequence of the Porphyra purpurea chloroplast
genome. Plant Mol. Biol. Rep. 13: 333-335.
Seki, A., Hanaoka, M., Akimoto, Y., Masuda, S., Iwasaki, H., & Tanaka, K. 2007. Induction of a group 2
M. Hanaoka - 7

BSJ- Revi ew 4: 19( 2013)

植物科学最前線 4: 20( 2013)

sigma factor, RPOD3, by high light and the underlying mechanism in Synechococcus elongatus
PCC7942. J. Biol. Chem. 282: 36887-36894.
Takai, N., Nakajima, M., Oyama, T., Kito, R., Sugita, C., Sugita, M., Kondo, T., & Iwasaki, H. 2006. A
KaiC-associating SasA-RpaA two-component regulatory system as a major circadian timing mediator incyanobacteria. Proc. Natl. Acad. Sci. USA 103: 12109-12114.
Woodson, J.D., & Chory, J. 2008. Coordination of gene expression between organellar and nuclear genomes.
Nat. Rev. Genet. 9: 383-395.

M. Hanaoka - 8

BSJ- Revi ew 4: 20( 2013)