植物科学最前線 4: 1( 2013)

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Various light-responsive strategies in photosynthetic organisms
Key words: algae; cyanobacteria; light environment; photoreceptor; plant
Rei Narikawa
Department of Life Sciences, University of Tokyo
3-8-1 Komaba, Meguro, Tokyo 153-8902, Japan
Mié Ishikawa
Graduate School of Life Sciences, Tohoku University,
2-1-1, Katahira, Aoba-Ku, Sendai, Miyagi 980-8577, Japan(&' ­³¸ÃbÛ 75 ybĔµ]ĕ"ĖATNB9OhuÊÃ#º!hÜÌí42Ėº!huÊÃ#hÜÇù5Ù·ëÔÙç 6 v#Àç".0Ė°gÚ#ÔÎï2#ATNB9O$;V<H8@V#W^0ĖA6JK>EP6#hÜ5ÔÙ1ĖhuÊÃ#hÜ"fl#ÊÃ#Èò"Ë(/Ė

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R. Narikawa & M. Ishikawa - 1

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R. Narikawa & M. Ishikawa - 2

BSJ- Revi ew 4: 2( 2013)

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
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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
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(Synechocystis)' J^lǦ¨ Łæ9'<ū§7& (Kaneko et al. 1996)ȏŜÍ/"& 40 Ž|'

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植物科学最前線 4: 4( 2013)

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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
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植物科学最前線 4: 5( 2013)

Ishizuka 2008)

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BSJ- Revi ew 4: 5( 2013)

植物科学最前線 4: 6( 2013)

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
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#(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

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CcaR &58 II ÐƳƫȇ²'¬ĄnWt

4."   !) R. Narikawa - 4
BSJ- Revi ew 4: 6( 2013)

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

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Okamoto and Ohmori 2002)
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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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植物科学最前線 4: 8( 2013)

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

'$&
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植物科学最前線 4: 13( 2013) "%  -$!#&ŜÓ ď¤ŞÂÇÂÇƕ¸śÇİĺĵ
271-8510 ¤ŞĭąñÔąñ 648
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<$Ĵĸ ŇŔĤ7 űĤ
œ 58783 Şŏ#%ƤĤ#ĦĄ7ğŖ$G]g DNA$ƉÅĩġ

ƢŻqŒŮƣJLUg 97ĈĜ$éĿðħ%Ƥsř#ć$ƉÅĩġ:}7
1$ Ģū87 ƤÑ" 1ĤÛ# %Şŏ $čŕ:óõÉ87ăĹ%Ƥ¦ŇŔńŠ C. merolae # 7ŞŏƉÅ$Żä#ƒ7įũ#ĊŲƤƋĤà$ģ¾éĿł2ƉÅĩġäł$Ƅ£#œÍ7

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植物科学最前線 4: 20( 2013)

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M. Hanaoka - 8

BSJ- Revi ew 4: 20( 2013)

植物科学最前線 4: 21( 2013)

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Light-responsive strategy in stramenopile alga, Vaucheria-Blue-light receptor, AUREOCHROME, act as a transcription factor-

Key words: Aureochrome; Blue-light receptor; Stramenopiles; transcription factor; Vaucheria


Mié Ishikawa1 & Fumio Takahashi1, 2
1.Graduate School of Life Sciences, Tohoku University
2-1-1, Katahira, Aoba-Ku, Sendai, Miyagi 980-8577, Japan
2. PRESTO, Japan Science and Technology Agency
4-1-8, Honcho, Kawaguchi, Saitama 332-0012, Japan
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植物科学最前線 4: 28( 2013)

Kikuyama, M., Wada, M., & Kataoka, H. 2007. AUREOCHROME, a photoreceptor required forphotomorphogenesis in stramenopiles. Proc. Natl. Acad. Sci. U.S. A. 49:19625-19630

M. Ishikawa & F. Takahashi- 8

BSJ- Revi ew 4: 28( 2013)

植物科学最前線 4: 29( 2013)31#.'-!%!&ÃðÍŽŴƆÝàŌŉťśǋŒàǩŌŉťśǋŒàſŽŴéŋñàÔŧ 
Photomorphogenesis in brown alga
Key words: blue light, genome, Phaeophyceae, Photomorphogenesis, photoreceptor
Shinya Yoshikawa
Department of Marine bioscience, Fukui Prefectural University
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植物科学最前線 4: 33( 2013)

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S.Yoshikawa-8

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606-8502 ǧāĀ»ºŴÿǗ°ű
Phytochrome-mediated photomorphogenesis in basal land plants
Keywords: Marchantia polymorpha; liverwort; nuclear translocation; bryophyte; phytochrome
Kimitsune Ishizaki, Keisuke Inoue, Takayuki Kohchi
Graduate School of Biostudies, Kyoto University
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K. Ishizaki- 8

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812-8581 ƞđƓƞđėőÄƫĒ 6-10-1
Molecular mechanism of chloroplast photorelocation movement in land plants
Key words: actin, Arabidopsis, chloroplast movement, organelle movement, phototropin
Noriyuki Suetsugu & Masamitsu Wada
Department of Biology, Faculty of Sciences, Kyushu University
Hakozaki 6-10-1, Higashi-ku, Fukuoka 812-8581, Japan

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2004, Rensing et al., 2008ȨȩǚŒ7ȯȧPpphotA1, PpphotA2, PpphotB1, PpphotB2Ȩ&
71ȖÖÎī!ǲȁÎī:¹ħ

6 ȩPpphotA V?i ǲȁÎī$Īȝ 6ȧKasahara etal. 2004ȨȧèȯȨĺÖǌȠ%ƢdNCwȧMougeotia scalarisȨ&ȭ%gB[[zf|:ĸ ȩ%!8dNCw %Šǀ&ń4#ȧSuetsugu et al. 2005bȨ
ȧèȯȨ

N. Suetsugu & M. Wada - 3

BSJ- Revi ew 4: 47( 2013)

植物科学最前線 4: 48( 2013)        b{~pt2*+8Z_fXBCFM șǆƗÉ&șǆ©ÎīȩǦǆƗÉ&Ǧǆ©Îī:Ɲāƹ&ǝń75ȩƛƹ&ǝń7# Ôǀĭ ȤdNCw&ĵę#ĖõǉƸ:Ƴƿ­$ĸȩğ©
&ĵę#Ȝ:ªĈ©$êƐ$ȧface-onȨȩġ© &©:ȁ63$ªĈ©$ęǏ$Ù
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Crphot :ƌƃ6!ȩȖÖÎī!ǲȁÎī!1$äĨ% ȧOnodera et al.

2005ȨȩgB[[zf|&ŎǥƎ$ȖÖÎī!ǲȁÎī% Ŀ:¹ħ6Šǀ:ĸ63
 6āȕ$ȩOz?_]R]% phot2 òƊ 1țĘ$ġ© ǲȁÎī ǡĉ76
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ü

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ňǱîÝ7ȧLuesse et al. 2010ȨT^LJ

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ƆŚŽ%ȓƏ$35ųřœ ģĳ76!ȩœĜ$^YX:ū06ŚŽ$!&ȩ35ğ©ƅð 1©:ô ÜÏ63ȩȖÖÎī%©İÑĭ Ȥ #6ĪǕ 

ğ©$ć6İÑĭ  6!ǲȁÎī135ğ© ǡĉ7-ȧ' Aihara
et al. (2008)% phot1 %©Ñă\p?|! phot2 %F]~T\p?|:ĸFpvgB[[zf|Ȩȩǿĝ#©ġĝŐ %©ÜÏÀƂ  6 ȖÖÎī%ǀ½&Ȥ ȩǲȁÎī
*%İÑĭ&©Ñă ĪǕ!#ô %ŚŽ&gB[[zf|:ǔľĸ ȩȀúȆǔĥĿ&1!1!%ȖÖÎī!ǲȁÎī: ĿĸV?i!ť5ȩ1Ŀ&ȖÖ
Îī$žÃȤİĝ%V?i*!òÃ!ƾ476k@v?OW2Oz?_]R]%
phot1 &,!;"ȖÖÎī$žÃV?i 6 ȩdpZwE`LJ% PpphotB &%N. Suetsugu & M. Wada - 4

BSJ- Revi ew 4: 48( 2013)

植物科学最前線 4: 49( 2013)

ȌŦȔ!Ǜ6gB[[zf|%=naȅȃµ$36³úƭƵǚŒ 4ȩgB[[zf|Ȁú%Ȇǔ&77%³Ƞƽ ƀƦ$ǧ 4'ƢúŚŽ% phot1 ! phot2 %Ȁ
úȆǔ&ǓúŚŽ!ǐúŚŽ%³Đ%» ǧ5ȩKJŚŽS|Ƞ &ƀǃ$ photA! photB $³ 7)K=GGRJSK;LPT268~Z^j3[i Ȃ%ĺÖǌȠȧdNCw! Mesotaenium caldariorumȨ
ȩdpZwE`LJȩô %ƃƆOWȠ &șǆ© # Ǧǆ©$31ǉƸǺÁ ǡĉ76ȧSuetsugu & Wada 2005,
2007aȨ
ȧèȮȩȯȨ
gB[[zf|&Ǧǆ©:ÜÏ#% ȩ%ƢȠ%©Ñă%ȍ&ńƍ Ǧǆ©$36ÀŔ&ǾǦǆ©$3ÔǳƎ$ĶŲ76% ȩg>[Hzo ȍ6! Ɲá7 ȩǉƸǺÁ$ȍ6g>[Hzo&șǆ©
Ñă×ŞƳƿǂ$Čé6! Ɲá7 ȩǅƎ$g>[Hzo&ŗ6&Ƴƿǥ$ČéŠǀ6! Ƙ476ŷ Ɩƒ ȧFankhauser & Chen 2008ȨĴ
&k@v?OW 4g>[HzoȀú:Hz~^|I6ǻơ ȩgB[[zf|%=naōƧ¦$g>[Hzo%©ÜÏ\p?|ƴÖFpv©Ñă`CHzo:ƌǗȧNozue et al. 1998Ȩ
ȧèȭȨ
`CHzo&gB[[zf|%Šǀ\p?|:+¢ĸ
% ȩƳƿǂ$ČéǉƸǺÁ:¹ħ6g>[Hzo%ā 6Ôǀĭ ġ Ɲá7āȕ$ȩk@v?OW%Ǧǆ©$36©čĭ!ǉƸǺÁ Ŀ:šĽȧșǆ©Îī&ŤĘ 6ȨòƊ rapȧKadota & Wada 1999Ȩ&ȩǣ+ȏ5+%v?| `CHzoȀú$šĽ ȧKawai et al. 2003Ȩ
4$`CHzoȀú: rap òƊ$ĉª
6!Ǧǆ©$36ǉƸǺÁ äĨ% ȩk@v?OW$`CHzo ǉƸǺÁȧ!©čĭȨ%Ǧǆ©Ñă!Šǀ6! ń4 !#ȧèȯȨ`CHzo&k@v?OW # ȩS?u@COWȧDryopteris filix-masȨ
ȩK@s{veȧOnoclea sensibilisȨ
#"ȩ@vlOƟ:Û/ polypod OWȠ%ƽȧƃƆOW%ȳȫȦ:È06Ȩ 41Ǘ ȩT|m?ȧOsmunda japonicaȨ
ȩD^HMȧLygodium japonicumȨ
ȩŪƆOWƩÊùƎ
#OWȠ 4&- %!8Ǘ #ȧKawai et al. 2003Ȩpolypod OWȠ &©čĭ!ǉƸǺÁ Ǧǆ©$3ǡĉ76 ȩÊùƎ#OW &ǡĉ7#polypod OWȠ&ƯȬ¨Ě»%ƍƮȩǐúŚŽ ųřœ:ģĳŅ$ŻƌƎ$ôŞÃȧSchneider et al. 2004Ȩ% ȩ`CHzo%ƁĦ %å!#Ôǀĭ&ȤȧKawai et al.
2003, Schneider et al. 2004Ȩ Ȣ + !$ȩĴ&ĺÖǌȠdNCw 41`CHzo:ƌǗȧSuetsugu et al.
2005bȨ
dNCw!OW%`CHzo&V|cHǥ%\p?|ŝǶ&3 6 ȩȀú
%AHU|}?|[z|ŝǶ « Ɗ#6OWȠ%`CHzoȀú&- ?|[z|:ĸ# ȩdNCw%%`CHzoȀú&ǔľ%?|[z|:ĸ4$ȩdN
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țĘ$ȤȧèȮȨ
dNCw%`CHzo:k@v?OW% rap òƊ ƌƃ6!ȩǦ
ǆ©$36ǉƸǺÁ äĨȧSuetsugu et al. 2005bȨ
dNCw%Ǧǆ©Îī:¹ħ6g
N. Suetsugu & M. Wada - 5

BSJ- Revi ew 4: 49( 2013)

植物科学最前線 4: 50( 2013)>[Hzo1Ó

4ƳƿǂČé Ɲá76% ȩdNCw 1`CHzo Ǧǆ©$36ǉƸǺÁ:¹ħ6Ôǀĭ ȤȧèȯȨ dpZwE`LJ 1Ǧǆ©$3ǉƸǺÁ ǡĉ76 ȧKadota et al. 2000, Sato etal. 2001Ȩ
ȩ`CHzoȀú&ĸ#ȧRensing et al. 2008ȨȧèȮȨ
%ŚŽ!&Ɗ#
5ņǴ%g>[HzoǉƸǺÁ:ǡĉ6!

Ƙ476ȧMittmann et al. 2004,

Uenaka & Kadota 2007ȨȜƍ!$ȩĴ&gB[[zf| Ǧǆ©$36ǉƸǺÁ:¹
ħ6!:ń4 $ȧKasahara et al. 2004ȨȧèȯȨ %!8gB[[zf|"%3$g>[Hzo!ÆǣŠǀ6 &ń4# {Xu}3<AES|z2Yd,0\rg ÁŽ2ǈȠ$6=HX|ƻƷ$ üCxE`vǺÁ%³úpD^Ro$&$Ǵ5 Ƙ476nCO|$3=HX|ƻƷ%:ŵ6ǺÁ!ȩArp2/3 ǔÖȧ=H
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1üé6ȧSparkes 2010, Deeks & Hussey 2005Ȩ ȩ%!8 Arp2/3 ǔÖ%ŚŽ%CxE`vǺÁ*%ȍ&ń4&#ċ# !1ƢúŚŽ$ȩƔŗƆŽ$¬Ǵ#CxE`vȧn[K|\w=ȩLxPȩjxCFOU~oȩĊƿȨ%ǺÁ&nCO|$ ü6! ňǱń4 $#ȧSparkes 2010Ȩ # 4ǉƸǺÁ&ȩArp2/3 ǔÖ%Šǀ šĽòƊ 1ŤĘ 5ȧKadota et al. 2009Ȩ
ȩ-ǣ+47ȏ5%nCO|%
aYH=@[òƊ 1%CxE`vǺÁ%ƊĘ$1ȍ94ŤĘ ȧSuetsugu et al.
2010aȨ! 4ȩǉƸ&%CxE`v!&Ɗ#6ŞĞ =HX|ƻƷ:ȩǺÁ6
! Ɲá76Ĵ&ȩ=HX|ƴÖV|cHǥȧVw|ȩg>|hw|#"Ȩ!Ǎ
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Ġ5Ĕ47=HX|ƻƷ2%ŏ%ȃƼ2ĹÁ$&©ŹĈ$36õ #òÃ&Ǘ47ȩǉƸǺÁ$ǷÁ63#Áı1Ǘ47# !8 ǉƸÞǮ:ǟƳ$ǙĆƴŔȩǉƸ$Ƴ =HX|ƻƷ Ǘ47ȩǉƸǺÁ$òÃ6!:ƌǗ
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Ƹ&Țţ6  

N. Suetsugu & M. Wada - 6

BSJ- Revi ew 4: 50( 2013)

植物科学最前線 4: 51( 2013)b ^jo2*+8  <AES|z3l\ ǲȁǺÁȋù»%Țţſı & cp-=HX|ƻƷ&ǉƸ%«Þç$üéȩǉƸ&9$Á &6 %ï$!"-6ȧƳÍĿÙƗÉȨǲȁÎī&©%ŹĈȂì­ȧș
ƹ%Ȃ³Ȩ%ǉƸ %.ǧ6% ȩphot2 &ǉƸóÂǂ6&Ƴƿǂ%ǉƸ$ĺ6Ȃ³$6!ƾ476phot2©:Ñă6!ȩcp-=HX|ƻƷ%ǻƎ#Ų÷ǧ5ȩǉƸ Ƴƿǂ 4ó7Áõ

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N. Suetsugu & M. Wada - 7

BSJ- Revi ew 4: 51( 2013)

植物科学最前線 4: 52( 2013)b`^jo2*+8  <AES|z3l\Ƴƿǂ% phot1 ! phot2©:Ñă6!ȩ4 %£Õ ƌƆȩƐĺ© Ģ#ǉƸ$1Įî 96ǲȁÎī!&Ɗ#5 cp-=HX|ƻƷ%ǻƎ#Ų÷&ǧ4ȩǉƸ%©ŹĈȂĿÙ»Å³

cp-=HX|ƻƷ%ȆÖ ǧ5ȩ©$Ù Ȗ-6ȧƗÉȨǉƸ%ƠÁǵĝ&ȩȖÖÎī 1ǲȁÎī 1ȩ»Ƨ¦!ĥƧ¦% cp-=HX|ƻƷ%ȈŨ
$ ü5ȩȈŨ õ#7'ǵĝ&ǵ #6ȧKadota et al. 2009Ȩ
4$ǲȁÎī

&©ġĝ%Ń$»ĥ% cp-=HX|ƻƷ%ȈŨ&ñõȩǉƸ&ǵ Á 74
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1#ġ©ŹĈŅ$Ǘ476 cp-=HX|ƻƷ%ǻƎŲ÷!ż¦*%¥éÃ&ǙĆ7ȧYamada et al. 2011Ȩ
%ƴŔ&ȩcp-=HX|ƻƷ%ż¦*%¥é&ǉƸǺÁ%ƴŔ &# ȩǺÁ%ǧå!#5Ħ6!:Ɲ6 Ƴƿ­ Țţ6ǉƸ&Ƴƿǥ$ű6% &# ȩ=HX|ƻƷ$ üƳƿǂ$ĺƕ6 ȧTakagi et al. 2009Ȩȩ7 1%ï 9 $Á6Ƴƿ%ě
Ƭç$ğ©:ŹĈ6!ȩǉƸ%«Þç

cp-=HX|ƻƷ ñ¾7$ǉƸ%Á 1ğ-ǳ$ġ©ŹĈ & cp-=HX|ƻƷ Ų÷ȩǉƸ&õÁ 3$#

ȧKadota et al. 2009Ȩ
%!&ȩ©ġĝ$ ü cp-=HX|ƻƷ%Ȉ%òÃ Ƴƿ
ǂ*%ĺƕ%ơĝ:ŬĀ6!:Ɲá6
 ©$ ü cp-=HX|ƻƷ%ÁıòÃ!ǉƸ©ĀǺÁ!%ȍǷ&ȩgB[[zf|šĽòƊ:ǚŒ$354$ń4 !#phot1 òƊ & phot2 $ üȖ
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÷!%ĥ%¥éÃ ȇƆë!Ũǭ9 $łV?n|I ǧ5ȩ%ƴŔ!ǉƸ ǲȁù06V?n|I1ł ȧIchikawa et al. 2011Ȩ
phot2 òƊ&ǲȁÎī:š
N. Suetsugu & M. Wada - 8

BSJ- Revi ew 4: 52( 2013)

植物科学最前線 4: 53( 2013)Ľ6% ȩġ© 1ȖÖÎī ǡĉ76ȧJarillo et al. 2001, Kagawa et al. 2001Ȩ%òƊ%ǉƸ$ġ©:ŹĈ6!ȩġ© ŹĈ7Ȃ³ % cp-=HX|ƻƷ%Ų÷&ǧ4ȩǳ$ cp-=HX|ƻƷ%ȆÖ ¡Ǹ7ȩȖÖÎī ǡĉ7ȧIchikawa et al. 2011,
Kadota et al. 2009Ȩ
phot1phot2 òƊ & cp-=HX|ƻƷ%© üƎ#òÃ&« ǧ4ȩǉƸ%©ĀǺÁ1Ƴƿǂ*%ĺƕ%©¹ħ1Ǘ47# ȧIchikawa et al. 2011, Kadota
et al. 2009Ȩ
74%ƴŔ 4ȩgB[[zf|& cp-=HX|ƻƷ%Áı:¹ħ6!
ń4 !#  <AES|z3[i298ac
 Oz?_]R]$6³úȀýƎǚŒ$35ȩ©ÑăgB[[zf| # ǉƸǺÁ%£ÕǼƭ2ǺÁƭ$ȍ96Ş#åú ×Ā7ȧèȲȨĴ&ȩ74%å
ú%òƊ$ȩǉƸǺÁ$ cp-=HX|ƻƷ%Áı:ǟƳ$ǙĆ6!$35ȩ
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cp-=HX|ƻƷ%¹ħ$"%3$ȍ6 :ĻĆèȲȩOz?_]R]$6ǉƸǺÁ%£ÕǼƭ2ǺÁƭ$ȍ96åú) " #$%%#% #$ Ĵ ÇȗòƊ%

chup1 òƊ&ň1ģǥ ġ ȩǉƸ©ĀǺÁ:šĽ6 # ȩǉƸ Ƴƿǂ 4ó7-°ȖȧKasahara et al. 2002, Oikawa et
al. 2003Ȩ
ǉŕƳƿ:ǙĆ6!ȩȇƆë &ºŶ© ŹĈ7#ȏ5ǉƸ&%Ƽ:,!;"ò# ȩchup1 òƊ%ǉƸ&ÊģǥŰÁ$Ŷ Ƴƿ­:ƠÁȧKadota et al. 2009, Suetsugu et al. 2010cȨ
%òƊ &Ƴƿǥ%=HX|ƻƷ%Áı&ȇƆë!ò94# ȩcp-=HX|ƻƷ « .47# ȧOikawa et al. 2003, Kadota et al.
2009Ȩ
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N. Suetsugu & M. Wada - 9

BSJ- Revi ew 4: 53( 2013)

植物科学最前線 4: 54( 2013)$Čé6ȒŚŽ$žƊƎ#V|cHǥ 6ȧOikawa et al. 2003Ȩ
ȧèȲȨ
=naōƧ¦%ƋŪĭȞì&ǉƸóÂǂČé$Īȝ 6ȧOikawa et al. 2003, 2008, Schmidt von Braun &Schleiff 2008Ȩ%ĥ8¦%=naōƧ¦Å³%K?x\K?x\p?|&ȩǉƸ Ƴƿǂ$ĺƕ60$ĪǕ 5ȧOikawa et al. 2008Ȩ
ȩ-Ǟȡƪ­%āȡƴŔ &ȈÃ$1
ĪǕ 6ȧLehmann et al. 2011Ȩ
K?x\K?x%Ţ$&=HX|ƴÖȂ üéȩǞȡƪ­ %=HX|ƻƷƴÖǀ ƜǠ7ȧOikawa et al. 2003Ȩ
DxlFOōƧ¦&izw|
ťí$ą/Ȟì!ȒŚŽ% CHUP1 %Ȍ țĘ$¢üĭ%ȤȞì:Û/ȧOikawa et al.
2003Ȩ
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āȕ$ CHUP1 &izg>w|ƴÖǀ:1ȧSchmidt von Braun &Schleiff 2008Ȩ
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植物科学最前線 4: 55( 2013)ȧSuetsugu et al. 2005aȨ
ȧèȲȨž$C~FOw|!ß'76HvQw|Ċƿ:ǐǖ6V|
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植物科学最前線 4: 56( 2013) ŚŽ Ȓ¶ŊȩȒŚŽ&ƐĺŁ©$476ƅð$ ȩŢƨ$ŋƆOWȩ
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