植物科学最前線 4: 3( 2013)  !)#(-,0."*+/   !) Ďò Ź 1,2
1. ĬÞäÞäǵƖ¿Ĝ²ŵƀź153-8902 ŭȋ³ȉÕ 3-8-1
2. źäĒưĖƩĸĵ  
332-0012 Óśůòºõĩţ 4-1-8

Light-responsive strategy in cyanobacteria-Light-responsive system regulating chromatic acclimation, phototaxis and cell aggregationKey words: cyanobacteria; photoreceptor; cyanobacteriochrome; two-component system; c-di-GMP
Rei Narikawa1,2
1. Department of Life Sciences, University of Tokyo
3-8-1 Komaba, Meguro, Tokyo 153-8902, Japan
2. PRESTO, Japan Science and Technology Agency
4-1-8, Honcho, Kawaguchi, Saitama 332-0012, Japan
1
 M>^_GVs>(Ķ%ŠčƋ& !ȏzĺŠšƟ#!Ǫƶ%ǩƍŨŠÐ¿Ď<Ư µİŠŗ"8Ňŀȏŉŀ"%ȏŴő%$'ŖÌÚ3Â2ǶÎȏȊōȏōȏÖÒĈş×%$'ĳǴş×&ŠƠ83'4ȏ±ŗȏıŗȏƬȂ%$#¡Š83'37ȏÎŝ|'Ķ%ş×
&ŠƠ!8/ȏ8Ž'M>^_GVs>(ş×Û±&ćƇ!ȏƐơ<¦²8)ȏAnabaena, Nostoc %$'M>^_GVs>(ƁƍȄȆĪ}"ȏfVvMOX#Æ)98ŦǉƐơ<ƌ 10 & 1 '®¿"ýĎȏ'Ɛơ"ƁƍËæ<îǮŪ&Ư 5 &%8'5 &Ķ%ş×&ǡćÝĶ%Ž ãÍ82ȏM>^_GVs>&(ÝĶȏȏȊù%ş×ćƇĸĵ

;!8# ĦĀ98% "3ȏ(¿ĎŠŗ&#!ĤǪƶ%ĉÔ# ȏȊù%
ćƇĸĵ'ãÍ ŸÇ98çǸȏM>^_GVs>"(ȏ» 6Ý'ćƇŜǆ Ų69!
8'ĝÁ&Áȏ8(ȏǕ8ǋĈȏǉ&ćƇ!ėǹSw`Gǉ'Ǭľ<Û

8Ƴƫȇ²ȏǉ&ćƇ!ėǹSw`Gǉ 6'B]tFyǞ°Ś<Û 8OVyXǢ
Ż%$Ķ%ćƇŜǆ ãÍ!8Û±8ş×&ȇ²ȏ°Ś5¿Ď<Ư 2&ȏM>^_GVs>& !96'ćƇŜǆ ĵƉ9#ƞ 698 zĝȏ1996 ÷&  DNA ŵƀđ&5!ȏ´ƐơĈM>^_GVs> Synechocystis sp. PCC 6803
(Synechocystis)' J^lǦ¨ Łæ9'<ū§7& (Kaneko et al. 1996)ȏŜÍ/"& 40 Ž|'

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M>^_GVs>'J^lǦ¨ Łæ9!8/ȏÝ'Ž"ýǉǑĚĒư ŷƂ9!8
2ȏJ^lĉÔ<ÒŬ#ǣâŶÙ&58ƱŜÐƺĭ Ɗ¯Ū&Ư;9!8'ƕĮȏĶ%ćƇŜǆ& !ȏ¦âugt 6Ɛơxugt/"'ĸĵ'ƾƐ Ġ6 #%78ȌÊ 1ȍ
ĩž"(ȏ9/"&ƭƟ6 ƺĭ<Ư!Ƴƫȇ²ȏǋĈȏƐơ£ǹ<¬Ą8ćƇMOVl& !'ŲƷ<Ƒ8Ř&ȏ¹é ¹é8ǉ#¬Ą8ćƇŜǆ#'ŮǱǱ& !ȏǡćŪċƝ# ƹŒ"Ǆǃ

Ê 1 M>^_GVs>'¹éMOVl'Ĵť

2   9/"&ȏ40 Ž|'M>^_GVs>'J^lƺǁ å ! 7ȏJ^lĉÔ 6¹é'ãÍ<ęŎ8# ¼Ƣ#%!8çǸ&J^lĉÔ<ǂ+8#ȏŉŀ8(ǶŠ'M>^_GVs>& !ȏ¹é ǅë&ãÍ8# ¦ ! (Ashby and Houmard 2006,
Ohmori et al. 2001)9/"&ŠŗŤ"Ų69!8¹é(ȏuTZytƕ¿ÐȌvYeMwȍ
ȏdrbwƕ¿ÐȌLOV Sw`GǉȏBLUF Sw`GǉȏGseXGvlȍ
ȏǯşVXrcvytƕ¿ÐȌd?XGvlȏM>^_GVsCGvlȍ
ȏGktǩƕ¿ÐȌPYPȍ&Þª98M>^_GVs>"(ȏGktǩƕ¿ÐÜ' !'S@e'¹é Ʒ

!8Ř&ȏLOV Sw`Gǉȏ

GseXGvlȏd?XGvlxM>^_GVsCGvl(,#=$'ŉŀȏǶŠ'M>^_GVs>&ãÍ!8¹é"8 (Brudler et al. 2003, Ikeuchi and Ishizuka 2008, Narikawa et al. 2006, Yeh
et al. 1997)zĝȏBLUF Sw`Gǉ4vYeMw(Ǵ69Ž'M>^_GVs>&'0ãÍ8(Jung et al. 2003, Okajima et al. 2005)drbwƕ¿Ð¹é(ȏdrbw Ǽƫ<Ã·82ȏǼƫ¹é#!ĸƢ8zĝȏǯşVXrcvyt<ƕ¿8d?XGvlxM>^_GVs
CGvl'Õ¿ȏÝĶ%ǉ'¹é ÔÅ9!896'ÝĶ%ǉ'¹é(ȏƕ¿!
8ƫƍŽ'ǟ4¹éĸĵ'ǟ&5!çŜ9!8# ¦ 7

8d?XGvl"(ȏǊƫ#ǠǊƫ'ǰ"'¼ǖŪÛĚȏM>^_GVsCGvl'Õ¿ȏǼȌƏȍƫ#Ɨƫ'ǰ"'¼ǖŪÛĚȏƗƫ#Ǌƫ'ǰ"'¼ǖŪÛĚ%$ ÔÅ9!8 (Ikeuchi and

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Ishizuka 2008)

3-,0   !)  M>^_GVs>'¿Ď& !(ȏǯşVXrcvyt¦â<ƕ¿ǍóÞƵ¿d?Kb
sRyl ȏ¶ćą'Gvvd?t Ã·"%<ėǹȏ'B]tFy<¶ćą&Ǟ
8 (Tandeau de Marsac 2003)d?KbsRyl(&²äƋ II &B]tFy<Ǟ! 7ȏş× Û±8#"ȏ²äƋ I &3B]tFy<Ǟ8OVyXǢŻ<Ÿd?Kbs
Ryl(vUY#K>&¦69ȏvUY 6K>ȏK> 6¶ćą*#B]tFy<Ǟ!
8ȌÊ 2Aȍ'Ǟǝż"(ȏųłǭ'<Ã·8ƫƍSw`Gǉ 6ȏǭłǭ'<Ã·8ƫƍSw`Gǉ*#B]tFy Ǟ98K>& !(ȏd?KM>^bsw#Æ)98
ǯşVXrcvytƫƍ ƕ¿>vd?KM>[w# Sw`Gǉ Ǌƫ'Ã·<Ĕ!
8vUY"(ȏd?KM>^bsw ƕ¿d?KM>[wSw`Gǉ ĆãÍȏķƫ'<Ã·!89Ü&ȏŽ&5!(ȏ6&ųłǭ'Ȍ&Ɨƫȍ<Ã·8ƫƍSw`Gǉ ƕ¿8#"ȏÝĶ%ǉ<¿Ď&«Ţ!8% "3ȏd?KBsOsw#ƫƍSw`Gǉ(ȏd?KBsOvbsw# Ɨƫ<Ã·8ƫƍ<ƕ¿!88Ž'M>^_GVs>(ȏş×'ǉ&ć!ȏd?KM>[w#d?KBsOsw'Ǭľ<Û±8
ȌƳƫȇ²ȍ
'Ƴƫȇ²(ȑ 'S@e&¦Ȃ98 (Tandeau de Marsac 1977)ȌÊ 2Aȍ
I Ð(d?KM>[w#d?KBsOsw'ĝ<ĕȏǉ&ć!'Ǭľ<Û±%II Ð(d?KM>[w'Ǭ(Û &ȏd?KBsOsw'Ǭ<Û±8 /7ȏǊƫ}"(ȏd?K
BsOsw<Ō6#"ȏƐơ(Ɨƫ<Ä8zĝȏƗƫ}"(ȏd?KBsOsw'Ǭ<Ø
4#"ȏƐơ(ȋ.ƫ<Ä8III Ð(d?KM>[w#d?KBsOsw'ĝ'Ǭ<Û±8 /7ȏǊƫ}"(ȏd?KBsOsw<Ō6zĝ"d?KM>[w<Ø4ȏƐơ(Ɨƫ<Ä8Ɨƫ}"(ȏd?M>[w<Ō6zĝ"d?KBsOsw<Ø4ȏƐơ(Ǌƫ<Ä8'5 %Ƴƫȇ²Ŝǆ(ȏǳ7ŃßǷ'ǉ Û±7ȏ'¿ĎŠŗ#ƅ¿78ş×'}"ȏ°Ś5¿Ď<Ư 2&ǡćŪ"8# 8
 96'Ƴƫȇ²Ŝǆ(» 6Ų69! ȏIII ÐƳƫȇ²<Ÿ Fremyella diplosiphon &!ȏƳƫȇ²<¬Ą8¹é1996 ÷& Kehoe 6&5!©2!ǣŪ&Àæ9 (Kehoeand Grossman 1996)'¹é<KyY8ǣâ rcaEŶÙ9į"(ȏǊƫĪ}"3ȏƗƫĪ}"3ȏd?KBsOsw#d?KM>[w'ǬľÛ²% rcaE

KyY

8Sw`Gǉ(M>^_GVsCGvlÐ'ƫƍƕ¿Ym@w<ĕ # 6ȏ¹é"8# ŸÇ9! ȏ¹é#!'¦ŘĈ(ǭ6~Ġ" ď'ŵƀHtye(Ǔ÷ȏSynechocystis # Nostoc punctiforme & !ȏRcaE #!ǽ%8M>^_GVsCGvlÐ¹é CcaS <Ʒȏ'¦ŘĈ<ǂ+ (Hirose et al. 2010, Hirose et al.
2008)'ƕĮȏCcaS (Ɨƫ#Ǌƫ'ǰ"¼ǖŪ&ÛĚȌÊ 1ȍ
CcaS (>AXeUX#
!ȏ_GVs>'Ď¦¬ĄƋ#!Ų698aOTNwEZyQYm@w<ĕȏ'Ʀôswǩ²ŅĈ(Ɨƫ&5!ǀð9ȏCcaS

ƗƫPwLy#!ĸƢ!8# ŸÇ9

ȌÊ 2Bȍ
Synechocystis (d?KBsOsw<ĕȏƳƫȇ²<Ÿ% ȏSynechocystis "(ȏǗö'd?KbsRyl&(%Řļ%d?KbsRylǣâȌcpcG2ȍ'Ǒ¢<¬Ą!8#R. Narikawa - 3
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Ġ6 #%!8 (Kondo et al. 2007)zĝȏNostoc ( II Ð'Ƴƫȇ²<ŸȏccaS ǣâŶÙį"(ȏƗƫ"ǀð98d?KBsOsw'Ʈſ ǌ6ȏǫŠį& !Ɨƫ"ǀð98
Ǭ#Ǌƫ"ē¬98Ǭ'ǰŪ%Ǭ'Ʈſ ȏǊȐƗƫĪ}"Ɔƹì996'
#(ȏCcaSƗƫ}"(d?KBsOsw'Ʈſ<ǀðȏǊƫ}"(d?KBsOsw'Ʈſ<ē¬!8#<ŸÇ!8CcaS '}ņ&(ȏCcaR # swǩ<¹¸!ŅĈ²
8uOjwOuFpuySyÐǑ¢Éâ ãÍ8' ccaR ǣâŶÙį& !(ȏƗȐǊƫĪ}"#3&d?KBsOsw'Ʈſǌ6% çǸȏd?KBsOsw&Ǳ;8ǣâƜ'Ǒ¢ugt<ǫŠį4 ccaS, ccaR ǣâŶÙį& !ǂ+8#ȏd?KBsOsw'ƮſǬ#ŮǱǑ¢Ǭ ƹì996'ç 6ȏCcaS

¹é#!Ǌƫ#Ɨƫ'Ǭľ<ČŲȏCcaR 'Ǒ¢ǂƈ<!ȏ'ş×}"°Śƪ¿Ď<Ư 2&d?KBsOswǬ<ǂƈ!8# Ÿ9 Fremyella diplosiphon & !(ȏRcaE '¦ŘĈ(Ġ6 &9!% ȏǣäŪƺĭ 6ȏ
aOTNwEZyQ"8 RcaE

6swǩ<¹¸8 RcaFȏRcaF

Ĉ²8uOjwOuFpuySyÐǑ¢Éâ RcaC #

3

66&swǩ<¹¸!Ņ

'Sw`Gǉǰ"'MHZtǞ<Ɣ!ȏǊƫĪ}"d?KM>[w'Ǒ¢ŅĈ<ǀðȏd?KBsOsw'Ǒ¢ŅĈ<ē¬8
# ŸÇ9!8 (Gutu and Kehoe 2012)Ǔ÷'J^lƺĭ 6ȏd?KBsOsw4d?KB
sOswÀĶƗƫ>wVZ"8d?KBsOvM>[w<ĕ M>^_GVs>& !ȏRcaE
4 CcaSƷ9!8 ȏ96 ¬Ą8#Ŏ98ǣâ ŽĽ&Ŧ%!8#Ċ9896'ǟ(99'Šŗ'ŠƠş×'ǟ&ǌÉ8 39%/ȏ99'¬Ąĸĵ&Ɛ %ǟ(8 ȏÒĩŪ&Ɨƫ#Ǌƫ'Ǭľ&ć!Ɨƫ>wVZ#Ǌ
ƫ>wVZ'Ǭľ<Û±8Ƴƫȇ²Ŝǆ(ȏ¹é8ǉ#¿Ď&«Ţ8ǉ zƨ!
7ȏǽö&ǡćŪ%Ŝǆ"8#(Ġũ"8

Ê 2 M>^_GVs>'Ƴƫȇ²#'¬ĄA: d?KBsOswǬ'0 Û±8 II ÐƳƫȇ²#d?KM>[w#d?KBsOsw'ĝ'Ǭ Û±8 III ÐƳƫȇ²B: Ď¦¬ĄƋ CcaS,
CcaR &58 II ÐƳƫȇ²'¬ĄnWt

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植物科学最前線 4: 7( 2013) » 6Ķ%M>^_GVs>"ǋĈ(Ų69! 7ȏŏ&Á !Ż±8Ļ'ǋĈ(ȏ¿Ď<57ŅŨ&Ư 2&Ǫƶ%ŠŞŜǆ# 8Ǔ÷ȏ& Synechocystis & !ȏǋĈ'Ɠ0 ¦âugt"Şƺ9

8M>^_GVs>'ǜ±&(ȏǛń4ŗƱǾ<Ż±

8Őǋǜ±Ȍ,-zĝÁ*zæǘù"±ȍ4ȏtwitching ǜ±ȌĝÁ<Û % 6ǰĹŪ&±ȍ
Ų69!8 ȏÞƤƬ%$'ǜ±Ʋƛ#!Ų69!8ǿĿ(ĕ%';7ȏÝ'ŽŧµƐƬ' IV ÐƘĿ#5ƘĿ'ǣâ<ĕ Synechocystis & !ȏ96'ǣâ<ŶÙ8#ȏŘæ'ƘĿ4 twitching ǜ±ȏƦœýǉǑĚƢ ňà8# 6ȏƘĿ<ǜ±ȏÜ
ī DNA '¸7ǒ0 ƞ 69!8 (Yoshihara et al. 2001)'5 %ÿ®'ǣâ ğ&ÝěÀæ
9! 7ȏƘĿĩ"%ȏ'ýĎ&Ćƶ%ǣâ3Àæ9!8 (Bhaya et al. 2000,
Okamoto and Ohmori 2002)
 įãĞƼ 6¸7ê Synechocystis 'Ƹį&(ȏũƫŏ&í!Ļ'ǋĈ<ŸƐơ#Ǉ'ǋĈ<ŸƐơ ŋÍ!2ȏ99<´ǻ8#"ȏPCC-P įȏPCC-N į#
96'į(ǣŪ&Ïz"%#ƞ 69! ȏĤǓȏĺMyIwLy<Ţ SNP ƺĭ&57ȏçǸ&Ƶě'ÛŦ ŷƿ9!8 (Kanesaki et al. 2012)ď'ŵƀHtye(& PCC-Pį<Ì½#!ȏŶÙį'ǋĈ Û²8ǣâ<ĘƎ!'ƕĮȏÝ'ǣâƜ Àæ9!8"3ȏ¹é<Â1 3'ChvwǣâƜ pixG GrOSyȏpixD GrOSyȏ

pixA GrOSy Àæ9!8 (Narikawa et al. 2011, Okajima et al. 2005, Yoshihara et al. 2000)
 pixG GrOSy(ȏÞƤƬƆ''_GVs>& !ǋ²Ĉ<¬Ą8ǣâGrOSy#'ŮÀĈ 6Űŭ9GrOSy"896'ǣâ<99 PCC-P į& !ŶÙ8#ȏǇ
'ǋĈ*#ǑĚ8 (Yoshihara et al. 2000)ǋ²Ĉ¬Ą& !²äŗǉ<ČŲ8#ƞ 69!8MCP Sw`Gǉ#ŮÀĈ 8Sw`Gǉ

PixJ1 "8
'Sw`Gǉ(>AXeUX#!ȏ

ǋ²Ĉ¬ĄSw`Gǉ#ÀĶȏMCP Ym@w<ĕȏ6&M>^_GVsCGvlÐƫƍƕ¿Ym@w<ĕ ' PixJ1 Sw`Gǉ<Ɗƴ!'¦ŘĈ<ǂ+#:ȏǼƫ#Ɨƫ&58¼ǖŪÛĚ<Ÿ (Yoshihara et al. 2004)ȌÊ 1ȍ
'ǣâGrOSy',#=$(ȏǋ²Ĉ<¬Ą8_GVs>'ǣâƜ#ŮÀǱ&8 ȏpixG '0q[yG%ǣâ"9(
PatA Ð'uOjwOuFpuySy# M>^_GVs>&ŘŦŪ%Sw`Gǉ<KyY!
 pixD GrOSy( pixD # pixE #  'ǣâ"ĵĎ9ȏpixD <ŶÙ8#ȏ4(7Ļ'ǋĈ Ǉ&ǖǑ8 (Okajima et al. 2005)PixD (drbwƕ¿Ð BLUF Sw`Gǉ"7ȏǼƫŔ
ï&57ȏdrbw'Ã·

10 nm ,$ǭłǭMdX8ȌÊ 1ȍ
/ȏPixD (ģĪ}" PixE #Sw`GǉƵ¿<ýĎȏǼƫŔï&57'Ƶ¿ ƺǻ8#3¦ !8
 pixA GrOSy( pixAȏnixBȏnixC # { 'ǣâ"ĵĎ9!8PCC-P į& !ȏ96'ǣâ<ŶÙ8#ȏpixA ǣâŶÙį& !(ȏĻ'ǋĈ Ǉ&ǖǑ8'&íȏnixBȏ
nixC ǣâŶÙį& !(ȏĝÁĈ'ǖǑ(ǌ6ȏ1:Ļ'ǋĈ'Ż±ǎǻ ǫŠį&ľ+!ĥċ&Þ%! (Narikawa et al. 2011)zĝȏǇ'ǋĈ<Ÿ PCC-N į& !ȏ96'ǣâ<ŶÙ8#ȏnixBȏnixC ǣâŶÙį& !(ȏǇ'ǋĈ Ļ&ǖǑ8'&íȏpixA ǣâŶÙį& !(ȏĝÁĈ'ǖǑ(ǌ6ȏ1:Ǉ'ǋĈ'Ż±ǎǻ ǫŠį&ľ+!ĥċ&Þ%!PixA Sw`Gǉ(M>^_GVsCGvlÐƫƍƕ¿Ym@w#

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aOTNwEZyQYm@w<ĥ8'ƫƍƕ¿Ym@w'¦ŘĈ<ǂ+8#ȏƏƫ#Ɨƫ
'ǰ"¼ǖŪÛĚ<ŸȌÊ 1ȍ
NixB # NixC (99uOjwOuFpuySy"7ȏ96'ǣâƜ GrOSy<ýĎ!8# 6ȏzǙ'MHZtǞƋǏ"ƚ !8¼
ƢĈ Ȋ ȏpixA # nixB, nixC ǣâŶÙį"('ƱŜÐ ǖ&%!8NixBȏNixC

ƣs

wǩ²řč"ŅĈ²ȏswǩ<¹¸8#"~ŅĈ²!8' 39%
 Ǿũ#& pixE # nixC ( pixG ÀĶȏPatA Ð'uOjwOuFpuySy<KyY8`rvH
ǣâ"2ȏ96'ǋĈ¬ĄGrOSyƜ& !ȏPixGȏPixEȏNixC

5 %ý

"ǋĈ'ĝÁĈ<¬Ą8ĤƒÉâ"8¼ƢĈ ŸÇ9899'¹é'Ĉǉ 6ȏ$'¬ĄƋ3ƏǼƫ&ćƇ8# ƞ 698 % 6ȏǫŠį& !´ƫŔï'ǋĈ*'þȀ<ǂ+#:ȏSynechocystis (ķǊƫ&í!Ĥ3ûĻ'ǋĈ<Ÿ#¦ !896'# 6ȏħÀæ9!%ķǊƫ&ćƇ8m@w'ǋĈ¬ĄƋ ãÍȏ'¬ĄƋ'O@UT#!ȏ96'GrOSy Ǳ!8# ŸÇ98
ȌÊ 3ȍ
96'¹é!ƏǼƫ"8#'ǡćŪ%ċƝ<ƞìƏǼƫ(ƏÜ&Ǔȏ¿Ď& 8ǲè<úǌ4"82ȏƏǼƫ ãÍ8Ģ&ȏķǊƫ&í8Ļ'ǋĈ<ē¬8¬ĄƋ#!ÿƂ!8¼ƢĈ 8Ă99'
¹é'ÝǪÛŦį<ŢȏƵě'ǉ<Ɠ0¿;Ȍ )ȏǼƫȎǊƫȏƗƫȎǊƫ%$ȍ'Ŕï ǋĈ& 8þȀ<ǂ+8#"ȏ96'¼ƢĈ Ĳƽ"8#ƞ !8
 'M>^_GVs>'J^l<ǂ+8#ȏ
pixG GrOSy(Ý'M>^_GVs>&
!ãÍ!8 ȏpixD GrOSy(zǥ'M>^_GVs>& ãÍȏpixA GrOSy( Synechocystis ' 0 & ãÍ%

Synechocystis (Ř&ƵǺ%ǋĈ¬ĄMOVl<ŨǞ¼ƢĈ ŸÇ98/ȏPixJ in
vH<ǂ+8#ȏfVvMOX<ýĎ8M>^_GVs>' PixJ Ɯ'0ȏƫƍƕ¿Ym@w'Ǧ¨ Û;!çǸ& Anabaena sp. PCC 7120
& 8 PixJ invH AnPixJ '¦ŘĈ<ǂ+
8#ȏǼȐƗƫ¼ǖŪÛĚ"(%ȏƗȐǊƫ¼ǖŪÛĚ<Ÿ (Narikawa et al. 2008)
fVvMOXýĎM>^_GVs>& !(ȏpixG GrOSy&58ǋĈ¬ĄƋ #(ŦÊ 3 Synechocystis & 8ǋĈ'¬Ą
nWt

%ý&%!8# ŸÇ98

5*+/   !)  ¿Ď(&5!Ȉ±98¶ć#&ã%¶ć#&¦698­Ɵ(&āǘȏĂƟ(ǨƍŅĈ&āǘ82ȏōĪ"(ȏǨƍŅĈ ŮíŪ&} 7ȏ¿ĎŠŗ(ûĪ#5 %OXuO<¹ȏǲè<úǌáŕĈM>^_GVs> Thermosynechococcus

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植物科学最前線 4: 9( 2013)

vulcanus RKN (T. vulcanus) (ȏƧǡŠƠōù

57°C 6"7ȏ45°C "(Ǘö'ŠƠ<ŸT.

vulcanus < 31°C &Ż#ȏƐơ( 24 Ģǰ,$'Ģǰ< !£ǹ<úǌ (Kawano et al. 2011)
ōĪ}"'Ɛơ£ǹ(ȏzƐơü7'ŔïǬ<Ō6#&%8'"ȏ 6'ÈǤ#ſĳŪ%ş×ȇ²#0%# "8'£ǹƐơ&í!ȏ¾Ž'Ǩƍ¤ŞȌPtryQȏ
HtDZyQȏevV>yQ%$ȍ<Ư #ȏPtryQ¤Ş&5!ŘŦŪ&£ǹ ƺň9/ȏ45°C # 31°C "ÑȅƐơ99'ƮſPtvyO<æǬ8#ȏ 31°C "ĥċ&PtvyOƮſ ǀð9!/ȏ45°C

6 31°C *Ɛơ<Ż! 6'PtvyOǬ'S@l

KyO<ƺĭ8#ȏ24 Ģǰ/"'ǰ&ă&Ʈſ!8# ¦ 96'# 6ȏT.
vulcanus & 8ōĪ}"'Ɛơ£ǹ&(PtvyO'Ʈſ Ćƶ"8# ¦  T. vulcanus 'J^lƺĭ(Ư;9!% ȏT. vulcanus #ǓƙȌÖÒugt" 99%'Ǧ¨ȂĈȍ% T. elongatus BP-1 ' J^lǦ¨(Łæ9!8 (Nakamura et al. 2002)T. elongatus 'J^l&(ȏ3'PtvyO¿ĎǨƍĶǣâ ãÍ! 7ȏ96'invH

T. vulcanus &3 !ã

Í!"ȏ96'ǣâ'ŶÙį<ƴ#:ȏTvtll0007 # ǣâ'ŶÙį&
!ȏōĪ}"'Ɛơ£ǹ4PtvyOƮſ Ĳ¥9% 96'# 6ȏ
Tvtll0007

KyY8PtvyO¿ĎǨƍ TvTll0007

ōĪ}"PtvyO<¿ĎȏƐơ£ǹ<úǌ!8# ŸÇ9 (Kawano et al. 2011)TvTll0007 (ƥǈǗYm@w#ŅĈąY
m@wÜ&ȏC ĨƄ& PilZ Ym@w<ĕ ' PilZ Ym@w(ȏc-di-GMP # şř\Gu
CTY<ƕ¿8Ym@w"8c-di-GMP (Ǔ÷_GVs>& !ȏ_@Cd?tlýĎ%$<¬Ą8PDwYmUPwNoy¦â#!Ńŭ<ǹ2!8 (Mills et al. 2011)ǧǩƬ4ÞƤƬ&!ȏçǸ& c-di-GMP

PtvyO¿Ď<¬Ą!8#3ÔÅ9!896'# 6ȏ

T. vulcanus & !3ȏc-di-GMP

PilZ Ym@w&ƕ¿8#"ȏPtvyO¿Ď<¬Ą!8¼ƢĈ ƞ 698Ǔ÷'ƺĭ&
57ȏǼƫ&5!ŅĈ²9!
c-di-GMP <¿Ď8¹é 'Ɛơ£ǹ<¬Ą!8# ŸÇ
9!8'¹é(Ǽƫ#Ɨƫ'ǰ"¼ǖŪ&ÛĚȏǼƫŔï&5!"8ƗƫÃ·Ðc-di-GMP <¿Ď8
6&ȏ

'¹é<KyY8ǣâ<ŶÙ8#ȏǼƫãŪ&ǌ8Ɛơ£ǹǌ6%%

96'# 6ȏ'¹é 6ĉÔǞ<Ɣ!ȏPtvyO¿ĎǨƍ

Ê4 T. vulcanus & 8ōĪ}'Ɛơ£ǹ'¬'ŅĈ²<Ɛơ£ǹ'zǙ'ĄnWt

ņ9 ŸÇ9ȌÊ 4ȍ '¹é Ǽƫ"ŅĈ²98 ȏǉ#Ɛơ£ǹ#'ǱǙĈ<ƞì4 ƃ"3ƻǔ5 &ȏǼƫ(ƏÜ&Ǔ"7ȏB]tFy3ûȏǲè<úǌ4"

R. Narikawa - 7
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植物科学最前線 4: 10( 2013)8'Ŝǆ ō# ȏǲè<ǌ4Ī}"'Ŝǆ"8#3ƞ 8#ȏǼƫ'¹é&57ȏ<ÈǤ8£ǹŜǆ<ǌ#(ǡćŪ"8# 8/ȏÈ( T. vulcanus& 8ƺĭ"8 ȏM>^_GVs>'J^l"ȏ¹éYm@w# c-di-GMP ¿ĎǨƍYm@w<ĕ ¹é<øĘƎ8#ȏ'Ý(drbw<ƕ¿!Ǽƫ<¹é8 LOV Ð
¹é4ǼȐƗǰ"ÛĚ8M>^_GVsCGvlÐ¹é ,#=$"96'ç(ȏÝ'M>^_GVs>& !ȏǼƫ&ćƇ!Ɛơ£ǹ<úǌŜǆ ãÍ8
#<ŸÇ8 39%

6%
 M>^_GVs>& 8ćƇŜǆ<¦âugt"ƾƐ&Şƺ82&ȏǓ÷(ýǉǑĚ ¼
Ƣ%nWtŠŗ<ŢǖǣäŪƺĭ " % 6ȏĺMyIwLy%$'ȃǐŪ%Ēưǚñ&57ȏǽnWtŠŗ& 8q[yG%ćƇŜǆ<ǖǣäŪ&ƺĭ8#­573éġ&%!8J^lĉÔ<ÒŬ#!ȏM>^_GVs>'ÝĶ%ćƇŜǆ<ȁȐǖǣäŪ&ƺĭ8#&57ȏM>^_GVs>'ćƇŜǆ<Ɛơugt"űŪ&Şƺ"8: zĝȏÝĶ%¹é'¹éǉ<Š²äȏ¦äŪ&ƺĭ8#3Ɗ¯Ū&Ư;9!8 (Rockwell et al. 2011, Rockwell et al. 2012)96 'ŲƷ Ʈſ!#"ȏM>^_GVs>'ćƇĐť<ȏ¦âugt 6Ɛơugt/"57ŊŞƺ"8#ĦĀ98

'$&
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