植物科学最前線

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÷àą Ǩ 1,2
1. ÜƍǬòƂƋĥ
411-8540 ȀĄžąĉǔŰ 1111
2. ťƍƽİŕƇòĦƾĩƲŋŉ, CREST
332-0012 ãŨžĆÉĉļŲ 4-1-8
Regulation and evolution of the chloroplast division machinery
Shin-ya Miyagishima1,2
1. National Institute of Genetics
1111 Yata, Mishima, Shizuoka 411-8540, Japan
2. JST, CREST
4-1-8 Honcho, Kawaguchi, Saitama 332-0012, Japan
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BSJ- Revi ew 5: 21( 2014)

植物科学最前線(

5: 22 ( 2014)#ķ4Ƥł#9«ÌģŭŢ*ȎƖƻ)ō®ŭűľ)ƹơ=4!Kzk$9 (Archibald 2009)
 Ƙƭ¯®ŭeJ[y?

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ȎĞŸ'Ƙƭ¯®ŭǶ£)Ƅƍ%)Ĕ)EzGbxÀ(ěȆ$9%=ƅ#ê)åÌȎƻȊ)Ƙƭ*Ƙƭ8'ı¦)ƹơ=Ð1ȎúƘƭǀ9%(68ȎƘƭ8)ƹơı±ǀ9¹(ƹơØ±õ(¥:9 $*ȎúņŢƘƭ*&)Ŋ(#ȎQ?ceJ[y?®ŭ(űľ9ƹơ)±ǀ=¸Ę#9)Ȏ0)ŋŉ

&)Ŋ(#ǨÀ#

±ǀ¸Ęŋŉ=ǉĿūǉ93(*Ȏ0ƹơ±ǀŋŉ)ūǉ

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ŭȍȎƹơé)EzGbx=Õì9%(68ê)ſńŭŢ)ƔƝ(Į#Ŧƨ)ņŢųé$ǘ0*ơ$ƅ:9ƔƝ*Ȏ::Ɩƻơƻ)űľ)ōƹơ=4!

S. Miyagishima - 2
BSJ- Revi ew 5: 22( 2014)

植物科学最前線

5: 23 ( 2014)Ɨ=ƴƗ%ǋǡ9

USGC#JKD#J87
 ǤĊȎƹơ*ÞƏ±ǀ(6#æŐ9 ȌÚ 2ȍȎǺņŢ)ǭ)ƛƥ$*Ȏ

ê±ǀ4ǈü:#9 (Miyagishima et al. 2011)ǿðȉęǳǈü(68Ȏƹơ±ǀ*)±ǀȂ(#Ȏ¯¿ư%é¿ưÎĶ(,:9%(68Ǩƽ9%<#9 (Possingham and Lawrence 1983, Kuroiwa et al. 1998)1986 ČȎÃƘƭÄíƖ
ƻ(!

#Ȏƹơ)±ǀȂ(y}KŤ)ŉǦȌƴƗ±ǀy}KȖPD y}Kȍ8 (Mita et al. 1986)ȎďƟȎÎŊ)ŉǦ

ǆ))ƻȊ5ǺņŢ$äÑ: (Hashimoto 1998, Kuroiwa et al. 1998):7)ƜŁ(â"Ȏƹơ±ǀ*:7)y}KŤŉǦ

Æƣ9%(6#ďǙ:9%ƅÔ:)ĔȎ)y}KŤŉǦ)ŉģÙð%)ÒǞ(ñÝ9V}fJǗ)Îõ(Ï#)ƂƋ
Ǩ>)ƜŁȎQ?ceJ[y?űľ)ƱĈǯÌß GTPase $9 FtsZ (Osteryoung
and Vierling 1995, Osteryoung et al. 1998, Mori et al. 2001, Vitha et al. 2001)ȎÅ,ȎúſńƘƭűľ)WA_s}ŊV}fJǗ DRP5B (Miyagishima et al. 2003, Gao et al. 2003)
Ȏy}KÒǞ(ĀÝȎ))V}fJǗƧ%ǃÌȌ±ǀǁƦȍ=đģ#ƹơ
±ǀ(Ƕ

9%

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2012)!08Ȏƹơ±ǀ*Q?ceJ[y?)4ǟ>ŋŉ%ȎúƘƭĳ'ŋŉ)Âǒ(67ƹơ±ǀǁƦ
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ȎƓǲ$)ƻȊ=ǸȎƹơ±ǀǁƦ)ŉģV}fJǗƧ*.#ú)ńMct(N^:#9 (Miyagishima andKabeya 2010)$*Ȏ:0$(Îõ:#9±ǀǁƦ)ŉģV}fJǗƧ(!
#ȎQ?ceJ[y?űľ)4)%Ȏúűľ)4)(<#ƙ9

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ưż$ǯÌ#y}KŤ)ŉǦ=8Ȏ)±ǀ(ěǅ'V}fJǗ=±ǀȂ(ĀÝȎƘƭǗ±ǀ=ďǙ (Harry et al. 2006)1995 ČȎQ?ceJ[y?űľ) ftsZ ǬðȎQ|Aa_T_)ƘƭńMct(ǆ!

1995)Ȏƹơ)±ǀ(ěǅ$9%

8 (Osteryoung and VierlingƄǎ: (Osteryoung et al. 1998))Ĕ FtsZ

S. Miyagishima - 3
BSJ- Revi ew 5: 23( 2014)

植物科学最前線ƹơ±ǀȂ)¯¿ưS]|r§(y}KŤ(ĀÝ9%

5: 24 ( 2014)

ĵ7

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(Kuroiwa et al. 2002)FtsZ *¯¿ưS]|r§(9¯§)ƴƗ±ǀy}K)7(
S]|r§(Ʀ9ȌÚ 2ȍ(Miyagishima et al. 2001)
 !08Ȏ®ŭĔǬð*ú)Ƙƭń(ƈƽ4))ȎFtsZ =Ě%eJ[y?ß)ƘƭǗ±ǀŋŉȎƘƭ¯®ŭĔ4ƹơ$ŋƮ#9%ƅ:)ĔȎQ?ceJ[y?űľ)ńN^V}fJǗȎARC6 (Vitha et al. 2003), MinD (Colletti
9%

et al. 2000), MinE (Itoh et al. 2001)Ə4ƹơ±ǀ(Ƕ)ȎŋƮ*ǒ.7:#'

ȎQ?ceJ[y?) minCȌminD, minE %®( FtsZy}K)ƦŔõ(Ƕ<9ȍ(Ȋ)Ǭðơƻ)ńMct(ǆ!

ǭ)ơƻ)ƹơMct(ȎftsI Å, ftsWȌeJ[y?(yF}Ìģ(Ƕ<9ȍ(Ȋ)Ǭð
ceJ[y?
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#ƘƭǗ±ǀǶǧǬð)ê*ȎƻȊȎņŢ)Mct(ñÝ7ȎƘƭ¯®ŭĔȎeJ[y?ß)±ǀÙð)ê*ì<:%ƪ

7:

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 )6
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)ńN^) FtsZ V}fJǗ

Ƕ

FtsZ %ÎŊ( C ĻƎ)¥ñȇáǶŷǗ%)ƜÌ(Ƕ<9%ǨÀȎ©)V}fJǗ%*ŵ'4')(ý#Ȏƹơ)±ǀ(*ȐƊȊ9šĴ) FtsZȌFtsZ2ȍ(*Q?ceJ[y?
ñÝ9ȌeJ[y?(

ƅ:#9ȍȎ4#))±ǀǶǧšĴ) FtsZȌFtsZ1ȍ(*Ȏ)Ǯ

³ş (Miyagishima et al. 2004))%*Ȏƹơ)ģƍǩƉ$ȎC ĻƎŌƸß)FtsZ°Ū%=ƅ#9FtsZ1 % FtsZ2 *ƹơ±ǀȂ(®ĀÝȎin vitro$ovqyr%l[|qyr)Ĵ=đģ$9%ƅ:#9ǺņŢ(

# FtsZ2 * ARC6 %ƜÌȎFtsZ1 * ARC3 %ƜÌ9ȌÚ 2ȍ(Maple et al. 2005, Maple
et al. 2007)ARC3 4Q?ceJ[y?űľ FtsZ

7ǨÀơƴņŢȌơƻȎǺņŢȍ(Ûĸ)V}fJǗ$8ȎFtsZ Ŋ) N ĻƎǭ±% MORN vXiȌ)V}fJǗ(

#ưƯǗ%ƜÌ9%

Ɓ7:#9ȍ=Ð2 C ĻƎǭ±7'9

(Shimada et al. 2004)ƠƐĽņŢȌQWȎƊðņŢȍ*ťƱ) ARC6 fx|KȌPARC6ȍ
=4ȎARC6

FtsZ y}K)đģ=¤Ǩ9Ĵ$ȎPARC6 *ǣ( FtsZ y}Kđģ

=Ƿø9 (Glynn et al. 2009)0ȎARC6 * PDV2 )ȎPARC6 * PDV1 )±ǀȂ) Ā Ý À ( ě ǅ $  9 Ȍ Ú 2 Ȏ Ú 3 ȍ (Glynn et al. 2008, Glynn et al. 2009) 

S. Miyagishima - 4
BSJ- Revi ew 5: 24( 2014)

植物科学最前線

5: 25 ( 2014)ÚȐ ƹơ±ǀǁƦ)ŉǦ%ŋƮAȕÃƘƭƖƻ Cyanidioschyzon merolae )Ƙ
ƭ¯$±ǀ9ƹơƘƭ(ƹơ*!ñÝ9ȌȍguZyGbOL)Ƙƭ¯$±ǀ9ƹơ!)Ƙƭ(êı)ƹơ

Ð0:9ȌȍSLze*:: 2 µmȌȍ10 µmȌȍBȕÃƘƭƖƻ C. merolae =ůǉĿ$ĵ7%'

±ǀǁƦ)Ĩ¾FtsZ y}KȎ¯§) PD y}KȎé§) PD y}KȎWA_s}y}K)ȅ(õ±ǀƦ(đģ:Ȏ)Ĕ±ǀȂ)Æƣ

Ǚ9±ǀ)ķƚőǻ(#ȎFtsZȎ¯§) PD y}K)ȅ(řìȎ±ǀóĔȎé§) PD y}K* 2 !)ïƹơ)ǵ$ĕ(řìȎDRP5BȌW
A_s}ȍ*šĴ)ïƘƭ(ƀŤġ$řì9Cȕ:0$(Îõ:#
9ŉǦÅ,±ǀǁƦŉģV}fJǗ)ñÝƦ%Ž)Ƕ£Q|Aa_T_)
åÌ=ƅQ?ceJ[y?(űľ9Ùð*ơ$ȎúſńƘƭ(űľ9Ùð*:é)ƴ$ƅ#9MCD1, MinD, MinE, ARC3 *::Žů
9%(68 FtsZ y}K)đģƦ=Ŕõ9

UUS-@5BE?487M& ǋ)6ðƧ(ȎeJ[y?)±ǀŋŉ)ğä(â"#Ȏƹơ)±ǀ(Ƕ<9ǬÎõ:í3'

7ȎQ?ceJ[y?)ƘƭǗ±ǀǬð)ê®ŭĔ(ì<:#9%Ȏǿðȉęǳ$żĪǈü:9ƴƗ±ǀy}K(Ȋ)ŉ

S. Miyagishima - 5
BSJ- Revi ew 5: 25( 2014)

植物科学最前線

Ǧ

Q?ceJ[y?$*ǈü:'%'&:ƹơ±ǀV}fJǗ)ñÝ

7Ȏ®ŭĔ(úƘƭ§Ġ:)6ơ±ǀV}fJǗ%#ķ´(Îõ:)

5: 26 ( 2014)7¼

7'ȎúſńƘƭǙŜ)ƹ

WA_s}ŊV}fJǗȎDRP5B $9 WA_s}*ſńŭŢ(Ûĸ) GTPase $8ȎÈùÝßC}^PA]QS)þƭđģĶ(Ȏđģ)þƭ%Ƙƭư=!'ǭ±)ƘƭǗ§ƿȂ$y}KŤ(ǯÌȎþƭ=Ƙƭư

7,8²9)(ěȆ)V}fJǗ%#ǉĿ

Ǩ>$)ĔȎ

Mctk|RBJ](68ȎſńŭŢ(*Ŋ'WA_s}ȊV}fJǗ
< (Praefcke and McMahon 2004))ƭǗ§ƿȂ(ĀÝȎ±ǀ(Ƕ#9%)Ɗ9%s]N}^y?±ǀȂ)Ƙ

äÑ:7(ȎÃƘƭÄíƖƻ

Cyanidioschyzon merolae )Mctk|RBJ])ƜŁȎ)Ɗ(*WA_s}ŊǬð
2 !

'%

ȊÅ,ņŢ(µĵȎšĴ*s]N}^y?±ǀ$*7ñÝ'V}fJǗ=N^#9%

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µĵĔƫ)V}fJǗȌDRP5Bȍ(!#ǒ.%;Ȏƹơ±ǀȂé¿ư)ƘƭǗ§(ĀÝȎƹơ±ǀ$*7%µĵ (Miyagishima et al. 2003))ĔȎQ|Aa_T_ƹơ±ǀèŵ)ǉĿ7ȎƖƻƹơWA_s})EU|K

4ƹơ±ǀ(Ƕ

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9WA_s}ŊV}fJǗ

(ǙŜ9%ĵ7%'(Miyagishima et al. 2008)
 )ĔȎQ|Aa_T_ƹơ±ǀèŵ)ǉĿ7Ȏé¿ưǕǤV}fJǗ PDV1 Å, PDV2Ȍ(6V}fJǗȍ(Miyagishima et al.
Úȑ ƹơ±ǀ(Ƕ9V}fJǗƧ)ǨÀFtsZȎARC6ȎMinDȎMinE Ə*ƹơ)Ɔª$9Q?ceJ[y?(űľȎǬðƧ*úƘƭ)ń(ƈƽ#9ǭ)ƻȊ(*ȎSepFȎFtsIȎFtsWȎMinC Ə4ŏñ#9Ȏ:7*)

ǃı)ƔƝ$ťƍ(ǃıØì<:#9ĴȎDRP5BȎPDVȎMCD1 *®ŭĔúƘƭ(68¼

7:V}fJǗƧ$8ȎDRP5B *ŝƴƻ)±ăĔȎPDV2 % MCD1*ǺņŢ)®ǤƆª$ŧė:7(ȎFtsZ1ȎPDV1ȎPARC6 Ə)fx|K
FtsZ2ȎPDV2ȎARC6

7ǨÀ

S. Miyagishima - 6
BSJ- Revi ew 5: 26( 2014)

植物科学最前線2006)Ȏ¯¿ưǕǤV}fJǗ MCD1 (Nakanishi et al. 2009)Îõ:

5: 27 ( 2014)Ȏ:7)V}fJǗ*ǺņŢ(ñÝ'3ȎƹơģƍĶ(û$*'ȎƻȊ

ǨÀǩƉ$¼:V}fJǗƧ$9%ƪ

7ǺņŢ±ǀV}fJǗ
7:9ȌÚ 2ȎÚ 3ȍ(Miyagishima 2011)
 ǋ)6(Ŋ'ƹơ±ǀǁƦŉģV}fJǗ

Îõ:#

Ȏ:4Ȏ

ǿðȉęǳ$żĪǈü:9ŉǦȌ¯§%é§) PD y}Kȍ)ŉģÙð$*'Ȏ)ÒǞ(ĀÝ94)$9%µĵķǠȎÍŰ7* PD y}K=Ð2ƹ

ơ±ǀǁƦ)ÃǾ(ģ»Ȏé§) PD y}KKzF})ƢƠĽ$9%=ŶǆȌÚ 2ȍ7(Ȏ)KzF}(* glycogenin ŊV}fJǗȎPDR18ȎPDR1PD y}K)đģ(ěȆ$9%ƜÌ#ƅ: (Yoshida et al. 2010, Yoshida

et al. 2012)PD y}K)ƢƠŤŉǦ*ȎƻȊ%ǺņŢ$čǈü:#

8ȎPDR1)EU|K%Ġ:9V}fJǗ*ǺņŢMct(4N^:#9
'

7ȎơƻȊȎS]xucfAzȌĔǡȍ)Mct(* PDR1 =N^9Ǭðǆ!

7'3ȎPD y}K)KzF}Ìģŋŉ)Ƴĝ(!#*Ĕ)ƂƋ



Ē

:9

UVSGC#JKD'72)+/%)+NO ¯§) PD y}K)ŉģV}fJǗ=Ð3Ȏ0ê)±ǀǁƦŉģV}fJǗ
Ɓ$9%ƪ

7:9

ĺȎŪÝ0$(Îõ:#9V}fJǗƧÅ,ŉǦ)Ĩ¾%

)Ƕ£=$ňǐ9 (Yang et al. 2008, Miyagishima 2011, Yoshida et al. 2012)
 ±ǀ(ªƍ#ķ´(ƹơ±ǀõƦ( FtsZ y}K

Q?ceJ[y?űľ) MinD Å, MinEȎ:(¼#ǺņŢ(

#* ARC3 %9ō(¯§) PD y}KȎFtsZ y}K%¯¿ư)ǵ(đģ:9)Ĕé§) PD y}KȌKzF}ƢƠȍ

đģ:Ȏ

MCD1

FtsZ y}K)đģƦŔõ(Ƕđģ:9)%Ȏ

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S. Miyagishima - 7
BSJ- Revi ew 5: 27( 2014)

植物科学最前線)ĀÝ(ěǅ$9%±#9

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5: 28 ( 2014)

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S. Miyagishima - 8
BSJ- Revi ew 5: 28( 2014)

植物科学最前線

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5: 29 ( 2014)

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9%'½Ł*ǎ37:'3ȎPDVƅ:7(Ȏƹ)ŶǪőǻ(#ȎPDVS. Miyagishima - 9
BSJ- Revi ew 5: 29( 2014)

植物科学最前線

5: 30 ( 2014))ǰ*Ȏƹơ±ǀ)Ź>'±ǀƛƥ5ƵĺŠ'ƹ$êȎëģǴƹ$*ÿ'%µĵĴ$Ȏ/

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S. Miyagishima - 10
BSJ- Revi ew 5: 30( 2014)

植物科学最前線

=4!S]xucfAz(

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S. Miyagishima - 11
BSJ- Revi ew 5: 31( 2014)

植物科学最前線(KzF}ƢƠĽ$9)(!#*ȎĔ)ƂƋ

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S. Miyagishima - 12
BSJ- Revi ew 5: 32( 2014)

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.>3<
Archibald, J.M. 2009. The puzzle of plastid evolution. Curr. Biol. 19: R81-88.
Colletti, K.S., Tattersall, E.A., Pyke, K.A., Froelich, J.E., Stokes, K.D. & Osteryoung, K.W.
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dynamin-like protein from plants, is part of the chloroplast division machinery. Proc. Natl.
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Harry, E., Monahan, L. & Thompson, L. 2006. Bacterial cell division: the mechanism and its
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