• 検索結果がありません。

О 28 Ȱ Ȱ О О 3 ƅ ƅׄ ׄԵ ԵȰ Ȱ 83 ƅ ƅׅ ׅ 2018 Ⱥ Ⱥ 12 ̀ ̀ NEWS LETTER Vol. 28 NO. 3 December 2018

N/A
N/A
Protected

Academic year: 2021

シェア "О 28 Ȱ Ȱ О О 3 ƅ ƅׄ ׄԵ ԵȰ Ȱ 83 ƅ ƅׅ ׅ 2018 Ⱥ Ⱥ 12 ̀ ̀ NEWS LETTER Vol. 28 NO. 3 December 2018 "

Copied!
78
0
0

読み込み中.... (全文を見る)

全文

(1)

光合成研究

О

О 28 Ȱ Ȱ О О 3 ƅ ƅׄ ׄԵ ԵȰ Ȱ 83 ƅ ƅׅ ׅ 2018 Ⱥ Ⱥ 12 ̀ ̀ NEWS LETTER Vol. 28 NO. 3 December 2018

THE JAPANESE SOCIETY OF PHOTOSYNTHESIS RESEARCH

ĆǹDž ğƇʊDZëëէgу&cG#<?B ָ̀Ҽ·ՌׄȡȟǓׅ 138

cm`SZ oˆqœ®€IϒՁDҡёôQIĻŠԿЊG'.cs®vҀԊIɖŎ

ҩÕ Ͻ áׄ̒ÚǓ áׅ 139

DŽNJ ğŠDZзIҿƇôIĻǬօƇ̸͂Iӊ˦IԺȜ

’±–¯µ€¯¶‡¶ áׄȡȟǓׅ 148

DŽNJūDz ğƇʊϮЕG˜4$Δͱg[<a6˜ʗҲµ˜ϪӃ 157

Čĵ êЀ ĎǕՌׄÚՑǓׅ 158

DŽNJ ̴ñ΂ĦΞΛ֣ɤդG`cğƇʊϮЕI˜Ȝժ ̠ξ Вׄ̒šǓׅ 159 DŽNJ ̰ΩG'.c׌сѽµфѠ±ž°CIՈìǬϒδӊ̕

ɇυ˓ͭ áׄÚՑǓáׅ 171

DŽNJ y“¤‰v‚±v~¬´'`N”l€°¶œ‰šn“†l™´wgν$<

÷ΩѹЋIŖβеָԶŠ ՎǧȢξͭøׄ̒ÚǓׅ 180

ƻƚ0Ɲ— Ū̴֌ȢƹGλѹ6cğƇʊλΩIğձȀIͫէýǯΪɲ

Ȕ̌ϤԀǬ áׄÃǗǓ áׅ 194

ū·šŴ О ƥĆұ¡€´ ԩҩ ɦ̒šǓ 195

ãÓLJ The 1st Asia-Oceania International Congress on PhotosynthesisŵőǁƏ

ĴŎô įŚľőŗׄƼλϮׅ 202

ãÓLJ О1ƥjjµq‚jjƬտҕʒIëժĒǁƏ ϼă ˳ɓׄ̒ÚǓׅ 204 ãÓLJ International Symposium on Photosynthesis and Chloroplast Biogenesis ŵőǁƏ

ϭȧ ӄǬׄÚՑǓׅ 205

ÐŘȗ(aI'Ϫa8 207

ˢ̊ğƇʊDZëëƙģëπԦ˽ 208

ˢ̊ğƇʊDZëëŇ 209

ğƇʊϮЕ ʚЍӄǺ 211

ȼÐëƉЮ 212

ѕօɜӑµӑÐŚօ 213

ğƇʊϮЕ ѕօǥƙµˢ̊ğƇʊDZë2018ȺɃɖƙ 214

ԆœͪÜëƙȿƏ

(2)

Ć Ćǹ ǹDž Dž

®

®Î Îģ ģó óœ œœ œǨ ǨE Eƞ ƞ B B/ / & &( (* *

ĸŁ®ÎģóœœǨ„ǿőƽ†ǟ ȁāĀêó„Ÿ´ǥâƂƈóƀƏĥȂ

Ⱥ(aˢ̊ğƇʊDZëëէg'Ɍ*ż.4DZëIԽƠGˇf?B Ⱥ)<

>W4< ȺIÐŘȗէIּīłͧ2h ȺäղJƮ͡ĥ͓2hGÅFԽ ƠC'ÀӛGFbW4<W< ȺIѕօէIӁȟøǰ2h ȺI ѕօէIêЀĎǕՌ2hGğƇʊϮЕ IѕօCJ'ÀӛGFbW4<2aG ȶèȼÐ]ѕօǥƙIϗ2hI1ũŏG`b͕ȺIȺëD ƥI³¶v~‰œgժ Ē4Ⱥ ƥIëǁğƇʊϮЕ gϒұ4 ȺլIԽƠgǓ*Fƛ֠)ԍ0c0 DF,уfc0D)C*W4<E%[́b։%13$W6ΪGȺëCJ~´¢

m¥¢€†¶ϒҶ¢€†¶Ԉ͂ƣȜϷFEǓ*FDZëIȺëG͖Ѹ6cé σgǻ˞4ğƇʊϮЕIϒȜGȕ4([4dW8h)ӺΰC*<ICJF$(D҈

Ӹ4B$W6W<ëǁ[ȷȿ$ğƇʊϮЕI˿˜IϪӃgȿZcIGǓ*FӺΰ g4B$cDɰ$W60I @IͯŗJˢ̊ğƇʊDZëIǓ*F̟CÞɜ[C*

cճbыь4B$,0D)ɨӂ=Dʂ5B$W6

ëէgɌ*ż.<D*GJҐFͯŗg4`%Dɰ$ʽ$B$<IC6)$,

@(IéσJè̆ÃGу&c0D)C*F(?<0Dg'ӚN4W6ÃC[ğƇʊ IˑЄ˽g÷céσJԼdF)a[ԺұÃC6ICЂIԯèɜGFbW6)Ƕʊ2 8<$Dɰ?B$W6ϗ2hI1ũŏgý֟6c0D)#cDɰ$W6):ID

*J`e4,'֤$Ҋ4W6

̑Ⱥ(aJÚՑǓDZIּīłͧ2h)ëէgɌ*ы+W6ЂG¾ԑ4B$<éσ

ŏDͯŏCˢ̊ğƇʊDZëI˼FcϒȜGӺΰ4B֖.c[IDĄ5B$W6Þɜ

[DZëGȌ6cëƙIϗ2hI1ũŏDА̴ϖFŵőg'֤$4W6

(3)

M

M_ _N Np p} }Z Zc c0 0Ź Źǘ ǘ, ,ƴ ƴƤ Ƥ¡ ¡5 50 0´ ´Â ÂǗ ǗƊ Ɗ/ / B B P

P} }S Sc cƮ Ʈnj nj0 0ĕ ĕ» »

Ņ”êóêóǯ ƣÎĵÂƀƏƈ ċáƈóúı

ĸŁîñêó ůóǠ ŪnjűŪƈóƈ

êǭčƒêó ǿƖIJƪĮǕŒŐ

ƶ’ƅŘų±ñåųđÖų¬ûŇď†

ľĥ+ŹƱ!&ƼñŌŪ1ñƴƜƬ²/M_Np}Zc,?2CBƴƤ¡0ºǾ¡Eēģ"BM _Np}Zc²/1Ʈnj‘ǤÿEq…Z/SnIǫ¡.-EƵƍ!&p}x}sbI ȁPLBȂp_}VJd,°/Źǘ"Bñƴ®EÉB,PLB 0ğǓ.ăæ,_}VJdƯ 0ēģAM_Np}Zc1ƴƤ¡5,´Â"BM_Np}Zc,ƴƤ¡0ƮnjƟģ1ž*

A-'@<P}ScƮnj+ByiP}ScXYHXT~\…ȁMGDGȂ,YP}S cXYHXT~\…ȁDGDGȂEŽƿŐģģ´,"BŹƱĻ@®Űä‰+űƪ!&ŌŪ +1C@0P}ScƮnj®Îģ=_}VJdƯ0ēģƴƤ¡0Źǘ/Ǥƿ.ĕ»Eň&"

,Ĺ@/ C*&ľĥŹƱĻ0M_Np}Zc0ēģ= PLB 0Źǘ/B MGDG = DGDG0ǤƿĠ1ŠĹ+(&8&M_Np}Zc@ƴƤ¡/´Â"BDZ/BPLB@_

}VJdƯ50^Jev`S.Ưçį/*P}ScƮnj-0?/ǬDB0<´(*

.(&–ÚĤ1MGDG=DGDG0Îģƭ¼ ‰!&XJge[eEŲ*M_Np }ZcŹǘĻ/P}ScƮnjĪĕ»EĹ@/!&8&M_Np}Zc@ƴƤ¡50´Â ǗƊ/BMGDG,DGDG0Ǭ‹/Ǭ!*<ƀƏEǕ;*AĿķ0Ơň/)*ƨø"B

1. JJ55ZZGG

§ƲÃFEI˶ʐCϒҒ4<ҸǬ̰ΩJёŠ8 7ѼԜ)Ң4,îէ4<$f^c§©~ׄ־

ŠҒλ&ׅIɒʃgDc0ID*Ǭҡсѽī CJҡёôIʼnִôD4Boˆqœ®€)ϒ Ձ4:IīՍGJʑȹFœ²ˆ®zlŽDD [Gœ²®¦®¡ŒkׄPLBׅD`KdcӄŇ

͋4$̩ǬΫI̸҅Է)ɒʊ2dc0daI҅

GJv²²šk°IƇʊÃլôC#cœ²v

²²šk¯ŽׄPchlideׅD:dgğýǯϖGv

²²šk¯ŽׄChlideׅQDǍˀ6c Pchlide Ջ ĜՓоׄLPORׅ)ңА6cPchlideIǑ,J҅

¼CLPOR'`NNADPHDğͯɲƵI»ѯҿƇ

*ԸчĞE-mail: [email protected]

ôgɒʊ4ğIΠȐgɚ?B$c 1,2)>FX GҸǬ̰ΩäǐḬΩ]Ҫ֥~j“•v‹¯

jIǑ,J˶ʐC[ͯɲǵ6cPchlideՋĜՓ оgʨ@<Z˶ʐCv²²šk°g÷dF$I JLPOR4(ʨ<F$ҸǬ̰ΩIΪɥC#c3) ˶ʐCϒҒ4<ҸǬ̰ΩI־ŠǬҡ)ğgż .cDğͯɲƵҿƇôÃI Pchlide Jϟ>G

ChlideGǍˀ2d:Iɜ2aGv²²šk°Q

DãӰ2dc4)ǓՙGңА4B$<LPORJԶ ](GĻӊ2dc·˝ğŠDZзҿƇô†´–v ԊgJ5ZD6cğƇʊխԸ†´–vԊ)Ґо DD[GƇʊ2dcPLB)ɱԶGȥlj6cDƈ ˮGˆ®zlŽ҅IͯϒF̸Ъ)ԍ0bц̘D 4Boˆqœ®€JҡёôQDĻŠ6c0I ԿЊJLPORG`cPchlide IğǍˀGő&

c

cm m` `S SZ Z

(4)

ğƇʊխԸՈìǬIԛığƇʊ†´–vԊ]Ґ оIƇʊğƇʊҺѣIɒʊDğƇʊŹɫзIͯ

ɲŠ2aGPLB (aˆ®zlŽ҅QI҅Ǎˀ gƍX0daIԿЊ)֏ȶG‡l¤‰v(@

ԧԶGԍ0c<ZğƇʊŹɫзI̸Ъ]ҡёô IϒՁ̸͂gӧRc<ZI§Œ°зD4BϚh GϮЕ2dB*<

PLB Dˆ®zlŽ҅JW?<,ϊFc̸Էg 4B$c)E>a[ҀԊÑ՗Ȟgž¶€G4<

̸҅ԷC#bҋƑͻ$0DG:daIҀԊф ʊ[֏ȶGïB$cׄƪ1ׅ0daI҅ҀԊI ǓՍĻJ̰Ω]~j“•v‹¯jGΪ́IжҀ ԊC#bj~°w¯‚²¶°Gs®v¶€

) 1 ĻǬцƇ4<§“s®v~°j~°w

¯‚²¶°ׄMGDGׅDMGDG Gs®v¶

€)[% 1 ĻǬцƇ4<s®v~°j~

°w¯‚²¶°ׄDGDGׅ):d;dĤôIй 50%D30%gŭZc5,6)MGDGJό͛ɲȖՍG Ȍ4BӇ͛ɲ֝Ս)ϠȌϖGȔ2$z¶´ƵI Кô̸Էgʨ@<ZūέCJ®¦®̸Էg÷d 7ãfbGt|{° II D`Kdc֏Ñ՗

Ȟ̸Էg÷c“´®¦®ҀԊC#c:dGȌ4 DGDG JӇ͛ɲ֝ՍDό͛ɲȖՍI•®´€)

`,~¯´‡¶ƵIКô̸Էgʨ@<Z͛Ί

͹ÃCÑ՗Ȟ̸Էgɒʊ6c®¦®ҀԊC#c

7)ׄƪ2ׅˆ®zlŽ҅IMGDG/DGDG͖J2 Gԩ$ČgDc0D)ϪadB'b0Iфʊ)

ˆ®zlŽ҅Ϊ́I̸ԷG՗ӂC#cƀѿɲ) Ϸƚ2dB$cPLB]œ²ˆ®zlŽIɒʊG

Ȍ4B[MGDG/DGDG ͖)՗ӂC#cƀѿɲ

)ӌŷ2dB*<)8):IӝсJǺ(CJF$

0dWCIǍϊôFEgν$<ӊ̕(ağθ dž½Cλѹ4<̰ΩGխ4BJs®vҀԊ) ğƇʊ]ˆ®zlŽ҅IɒʊҡёôIϒՁG՗

ӂFɖŎg̘<60D)˦a(G2dB*<)

˶ʐ½CIoˆqœ®€Iɒʊ]PLBIϒՁ PchlideIƇʊDLPOR DIҿƇôɒʊG'.c Ҁ Ԋ I ՗ ӂ ɲ J ¾ ˦ C # ? < Þ ƥ ʋ J MGDG]DGDGIƇʊѿŏ)ñ½4<~²l‘

gν$Bs®vҀԊ)˶ʐCʢ%ɖŎ gӧR<IC:Iʊ̘gтß6cW<˶ɒʃ ɒʊɜoˆqœ®€(aҡёôQĻŠ6cԿ ЊGs®vҀԊ)EI`%Gխ¿6cI(Ϊ GPLB (aˆ®zlŽ҅QI‡l¤‰vF҅

ǍˀGEI`%FɖŎg̘<6I(G@$B[

ŻbфhC'b:I˿˜Iц̘[ù8Bтß4

<$

2. ooˆˆqqœœ®®€€ϒϒՁՁGG''..ccMGDGIIɖɖŎŎ ̰ΩI MGDG ƇʊJҐоôş҅GȗƱ6c MGDG ƇʊՓоG`?Bұfdc~²l‘

GJҐоôīş҅GȗƱ6cMGD1 Dǐ ş҅GȗƱ6cMGD2DMGD3I3ЋIMGDG

ƪ

ƪ1. ˆˆ®®zzlŽ҅҅DDPLBIҀԊԊффʊʊI͖ԞԞ  m±´„m(aūֈ2d<ҡёôIˆ®zl

Ž҅5)Dz¥u־ŠҒλ&(aūֈ2d<oˆ

qœ®€IPLB6)IҀԊфʊÅӂF҅ҀԊI Xgӑԟ

ƪ

ƪ2. MGDGGG `` cc  tt || {{  °°II̸̸ ԷԷ DD DGDGGG``cÑ՗՗ȞȞ̸̸ԷԷ

ό͛ɲȖՍGȌ4̴ɲ֝ՍIôАIŭZcŎƇ )Ȕ2$MGDGJz¶´Ƶg4B'bt|

{°IID`Kdc̴ɲ֝ՍgīďGօƇ4<

̸ԷgDb]6$DGDGJ~¯´‡¶Ƶg4 B'b͛Ί͹ÃCÑ՗Ȟ̸Էgɒʊ6c

(5)

Ƈ ʊ Փ о ) ǯ Ʊ 4 ҡ F E I ё Ґ ф Ѡ C J MGDG IǓՍĻJ MGD1 G`?BƇʊ2dc

9,10)MGDGJDGDGƇʊIƼԊD4B[úfd

c0D(aMGD1JDGDGIƇʊG[ɨӂ¾ ƀͅC#bˆ®zlŽ҅Güш2dcs®v

ҀԊIƇʊIǓŧgʢ?B$cMGD2DMGD3 J¯´ͅÈˮGҐоôǐGüш2dc DGDG IƇʊGÅGӺΰ6cDѮ&adˆ®zlŽ҅

Is®vҀԊIƇʊQIȄ¿J̴ZBճǺϖ

C#c11,12)

MGD1ՈìǬgǶĤGͅ$<“‰vjmǍ ϊôCJs®vҀԊ)TDhEƇʊ2dF$

<Zˆ®zlŽ҅Iɒʊ)'0a7̰ΩôI λѹG[ͻņFɕ֕)λ5c10)Ғλ&I͑ռC Ҋ͎DFc0DGő&MGD1I“‰vjmǍ ϊJѼɒʊ[ɐ,ձȀ4ǬҡIɒʊ¾ĤgɌ*

ԍ06<Z13)0IǍϊôI־ŠҒλ&gν$<

ӊ̕J¾ƀѿC#?<:0CʋJMGD1 mRNA g†¶y‰D4<ÜȪ£lv² RNA

ׄamiR-MGD1ׅgŒt|¦†…´ׄDEXׅýǯ ϖGϒδ28cзgν$14)MGD1Iϒδʙń) oˆqœ®€IϒՁGEI`%Fɕ֕g¿&

cI(gӧR<15)

amiR-MGD1ɒԊԛˀôg˶ʐCϒҒɜ4ˢլ

λѹ28<D0eDEX ĸζ4<ćôׄ+DEXׅ C J MGD1 I mRNA ± ž ° ) ̈ ĸ ζ ć ô

ׄ−DEXׅI35%GWCʙń2d<:0CҀԊ фʊgĻ̕4<D0e+DEX IDGDGƍՙJ

−DEX DTDhEǍfaF(?<)MGDG I ՙJ−DEXI65%GWCñ½4B$<:I<Z

−DEX C 1.5 ЊɃC#?< MGDG/DGDG ͖) +DEXCJTU1ׄMGDGDDGDG)РՙׅG F?<͆G0IMGDGI΀ȕ)oˆqœ®

€IɒʃG¿&cɕ֕gӧR<+DEX CJ PLB I̩Ǭ̸ԷJf7(GËd<)PLB Iϒ Ձ:I[IGJǓ*Fɕ֕g¿&F(?<ׄƪ 3ׅ0Iц̘(aPLBIɒʊGJMGDG/DGDG

͖IųȅFz´²¶°JɨӂCΝ$0D)ŀ

˦4<PLBI|lJLPORIңАՙGǓ*

,ȫƄ2dc0D)ϪadB$c2)+DEX|´

œ°CMGDG)΀ȕ4B[LPOR IңАG`

bPLBI|lJǍfaF(?<DѮ&adc

·˝œ²ˆ®zlŽIϒՁJ+DEXćôCǓ*

,ձȀ2d<ׄƪ3ׅ+DEXCJ҅IÅӂҀԊ ʊĻC#c MGDG )΀ȕ4<G[((fa7 PLB JԵȶI|lGԩ$±ž°WCϒՁ4<

0D(a:IϜ²ˆ®zlŽIɒʊGú&

cs®vҀԊIՙ)¾ԑ4<I([4dF$

ƪ

ƪ3. amiR-MGD1I־ŠŠǬǬҡҡGG''.. c

coˆqœ®®€€IɒʃʃDDPLBI̸ Է

Է

ׄA, BׅDEX ̈ĸζz´²¶°

ׄ−DEXׅDׄC, DׅDEXĸζ4MGD1 gʙń4<|´œ°ׄ+DEXׅׄBׅ

ׄDׅJ:d;dׄAׅׄCׅIPLB IʥǓƪϩŮJœ²ˆ®zlŽg Ϸ6€x¶°•¶J1 μmׄA, Cׅ D0.2 μmׄB, DׅׄFujii et al., 201715ׅ

g ˌ Ǎ www.plantphysiol.org.

Copyright American Society of Plant Biologists.ׅ

(6)

͆GMGDGI΀ȕ)PchlideIƇʊ]ңА ҿƇôɒʊGEI`%Fɕ֕g¿&cI(gӧ R<+DEX I־ŠҒλ&CJPchlide Iՙ)

−DEX Iй 60%GWC΀ȕ4B$<:0C PchlideƇʊIɛԶ͑ռIλʊΩC#c 5-j¤“

±›¯´ՔׄALAׅg־ŠҒλ&GԿōG¿&c 0DCPchlideƇʊхԔIΫʃgӧR<−DEX CJALA ĸζG`b˶ʐCI˿уμΩC#c Pchlide)֣ҢGLJő4<)+DEX CJPchlide I ң А ՙ ) ȕ F , ã f b G à լ ô C # c Protoporphyrin IXׄProto IXׅ Mg-Proto IX Mg-proto IX monomethyl esterׄMg-Proto IX MEׅ )ңА4<ׄƪ4ׅ0Iц̘(aProto IXG MggՒð6cMg-chelataseׄMgChׅMg-Proto IX g ¦ ˆ ° o € ‹ ° Š 6 c Mg-proto IX monomethyl esteraseׄMgMTׅ͆G5ωϞIθ

̸Էgȓģ4 divinyl Pchlide g÷c Mg-ProtoIX ME cyclaseׄMgCYׅG`cãӰхԔ)ձȀ2d B $ c 0 D ) Ϸ ƚ 2 d < Ϊ G +DEX C J

Mg-Proto IXIңА)֣ҢC#b:IÃլôI

ңАœ²šil°) MgMTIǍϊôD`,ïB

$<0D(aMGDGI΀ȕJMgMT)ӋǨ6

c€‹‰œGǓ*Fɕ֕gŷU4<DѮ&ad c

Pchlide IĻ̕(a+DEX CJğͯɲƵI

Pchlide )ΪG΀ȕ4B$c0D[˦a(DF?

<MGDG I΀ȕG`b Pchlide ) LPOR ]

NADPHD%W,ҿƇôg÷dB$F$ƀѿɲ)

Ѯ&adc2aGğͯɲƵI»ѯҿƇôJ҅

¼CօƇ4Ǒՙôgɒʊ6c0D)ϪadB$

c)+DEXI־ŠҒλ&CJ»ѯҿƇôIǑ ՙôŠ[ձȀ2dB$c0D)Ļ(?<0da Iц̘JMGDG)Pchlide-LPOR-NADPH»ѯ ҿƇôIɒʊ]:IǑՙôŠG՗ӂC#c0D gϷ4B$c

ä¼Iц̘(aMGDGIҕȸI΀ȕJPLB IɒʊGJǓ*Fɕ֕g¿&F$)œ²ˆ®z lŽIɒʊ]PchlideIƇʊğͯɲƵI»ѯҿ ƇôIɒʊgɐ,ձȀ6c0D)˦a(DF?

<15)

3. ooˆˆqqœœ®®€€ϒϒՁՁGG''..ccDGDGIIɖɖŎŎ MGDGƇʊDƈ̹DGDGIƇʊ[Ґоôş

҅CұfdcUDP-Gal (a MGDG Gs®v

ƪ

ƪ4. ALAĸĸζζɜɜIIamiR-MGD1GG''..ccPchlideƇƇʊʊÃÃլլôôIңАА

־ŠҒλ&GALAg24ˮլĸζ4<ɜIPchlideƇʊÃլôIңАՙg−DEXD+DEX|´œ°C͖Ԟ4

< Pchlide I Ƈ ʊ х Ԕ D Ɔ à լ ô I ̸ Է g w ® š I ½ G Ϸ 4 < ׄFujii et al., 201715)g ˌ Ǎ www.plantphysiol.org. Copyright American Society of Plant Biologists.ׅ

(7)

¶€1ĻǬgԛЈ6cDGDGƇʊՓоJ~

²l‘GJDGD1DDGD2I2Ћ)ǯƱ 6cE>a[ҐоôIǐş҅GȗƱ6c)ё ҐфѠIDGDGIǓŧgƇʊ6cIJDGD1C

#c16,17)DGD2JMGD2]MGD3DĦGĕ*

¯´ͅÈˮIDGDG ƇʊGÅGȄ¿6c 11,16) ğθdž½Cѹ?< DGD1 I“‰vjmǍϊô

ׄdgd1ׅCJDGDG ƍՙJ՘λ̦I 10%ä½ G΀ȕ4ćô)ϬŠ6cFḚΩôIλѹG[

ɐ,ɕ֕6c)ҡёôIϒՁGJҊƓϖFɕ֕

J¿&7v²²šk°ՙ]ğƇʊͯɲI#cЊ ɃI΀ȕ]ˆ®zlŽ҅)΅˻6cD$?<ձ Ȁ)XadcЊɃC#c 17)>FXGdgd1 I ϬŠGխ4BJ¨€§´ՔIԿōλμ)ŲƦ C#c0D)Ļ(?B$c)18)F9DGDGI

΀ȕ)¨€§´ՔIԿōλμgɌ*ԍ06I (G@$BJÞID0e¾˦C#c

amiR-MGD1gν$<ʋIϮЕ(aMGDG IҕȸI΀ȕ)PchlideIƇʊ]ңА»ѯҿƇ ôIɒʊGǓ*Fɕ֕g¿&c0D)Ļ(?<

) 15)DGDG )oˆqœ®€IϒՁG'$B EI`%FɖŎg̘<6I(G@$BJ̈ӊ˦

C#?<:0CʋJ dgd1 Ǎϊôgν$

DGD1 Iͅ˅)oˆqœ®€IϒՁGEI`

%Fɕ֕g¿&cI(gӧR<19)־ŠҒλ&G '.cDGD1IȄ¿)Ļ(?B$F(?<<Z W7dgd1I־ŠҒλ&IҀԊфʊgӧR<D0 eDGDG Iƍՙ)՘λ̦I 20%ЊɃGWCñ

½4B$<0Iц̘(aoˆqœ®€G'

$B[DGD1)ÅӂFDGDGƇʊՓоC#c0 D)˦a(DF?<dgd1 Ioˆqœ®€g ӈȉ4<D0ePLBJ՘λ̦DƈЊɃIǓ*2 WCϒՁ4B$<)īՍI̩Ǭ̸ԷJǓȷGË dB$<ׄƪ5ׅW<՘λ̦Ioˆqœ®€

JǓʟ 1 @IǓ*FĬΫׄǻտGJεΫׅI PLB4(ʨ<F(?<)dgd1CJ͍FPLB) ҿ˕Xadc0D)Ǒ,PLBIϤĬɃ[Ң4, ñ½4B$<ҿ˕X&cPLB JǶĤG˛Φ Š4B$cI(ļΦǐID0eC@F)?B$

cI(JĻ(aF$)PLBIɒʃGϊȶ)λ5 B$c0DJ˦a(C#c0IPLB Iɒʃϊ ȶJīՍI̩Ǭ̸ԷIËdDխĀ6cI([4 dF$W<dgd1 CJœ²ˆ®zlŽIɒʊ [TDhEӃadF(?<)amiR-MGD1 I x¶€Dƈ5,s®vҀԊIǓŧ)PLB I ɒʊGúfd<<Zœ²ˆ®zlŽg÷cҀԊ )¾ԑ4<ƀѿɲ)Ѯ&adc

͆GPchlide IңА]ƇʊхԔIΫʃgӧR

ƪ

ƪ5. dgd1I־ŠŠǬǬҡҡGG''..ccoˆ q

œ®®€€IɒʃʃDDPLBI̸ԷԷ

ׄA, Bׅ՘λ̦ׄwild typeׅDׄC, Dׅ dgd1ׄBׅׄDׅJ:d;dׄAׅ

ׄCׅIPLB IʥǓƪϩŮJœ²

ˆ®zlŽgϷ6€x¶°•¶J1 μmׄA, CׅD0.2 μmׄB, DׅׄFujii

et al., 201819) g ˌ Ǎ

www.plantphysiol.org. Copyright American Society of Plant Biologists.ׅ

(8)

<ц̘amiR-MGD1 IǂƇDƈ̹dgd1 CJ

PchlideIңАՙ)ȕF,ALAIͽőǻֶ(a

Mg-Proto IXI§“¦ˆ°o€‹°ŠI€‹‰œ

)ΪGձȀ2dB$c0D)Ļ(?<ׄƪ 6ׅ MgMT Iʢ%€‹‰œJs®vҀԊI΀ȕI ɕ֕gΪGż.]6$DѮ&adc)ĨôϖG EI`%FձȀ)ԍ0cI(G@$BJÞɜI Ӧ֠C#cW<dgd1CJğͯɲƵIPchlide IŎƇ)΀ȕ4B$<0d[ amiR-MGD1 I +DEX Dƈ̹C#c)Pchlide IƇʊձȀ[ğ

ͯɲƵI΀ȕ[amiR-MGD1I˝)`b֣ҢC

#?<:d;dIҀԊI΀ȕIЊɃׄ+DEX amiR-MGD1CJMGDG)35%I΀ȕdgd1C J DGDG ) 80%I΀ȕׅ[ù8BѮ&cD

Pchlide IƇʊ]ğͯɲƵIңАGȌ6cɕ֕J

MGDGI˝)ɐ$DѮ&adcamiR-MGD1I +DEX CJ»ѯҿƇôIǑՙôŠ)ձȀ2dB

$<):I`%FδӷJ dgd1 CJӃadF (?<·˝amiR-MGD1DϊFbdgd1Ǎϊ ôCJPchlideIğǍˀɜIChlide-LPORҿƇô IʫŗGϊȶ)λ5B$<š®‰~ªΠȐɜ

Chlide-LPOR ҿƇôIǑՙôJÑՙôQDӊֈ

4B$,)dgd1 CJ:IԿЊ)Լ,F?B$

<0daIц̘(aMGDGDDGDGJE>

a[ğͯɲƵ»ѯҿƇôIɒʊG՗ӂC#c) :Iխfb˝JϊFcDѮ&adc

4. ooˆˆqqœœ®®€€((aaҡҡёёôôQQIIĻĻŠŠˮˮGG''.. c

css®®vvҀҀԊԊIIɖɖŎŎ

~²l‘I־ŠҒλ&CJğΠȐɜ1 ˮլIˮΛC PLB I̩Ǭ̸Է)Ǔ*,Ëd4 ˮլɜGJPLB IƉ͏[TDhEӈȉ2dF, Fc0IտPLBIШΫI̸Է)ϟʹʑȹFҥ

̸҅ԷGǍˀ2dœ²ˆ®zlŽׄ[4,Jˆ

®zlŽ҅Iʼnִôׅ)֣ҢGϒՁ6c0D)˦

a(G2dB$c 20)PLB Dˆ®zlŽ҅J֏

ȶGï<ҀԊфʊgʨ>E>a[s®vҀԊ gÅʊĻD6c)ׄƪ1ׅ0I‡l¤‰vF

҅ǍˀGs®vҀԊ)EI`%FɖŎg̘<

6 I ( J ` , Ļ ( ? B $ F $ : 0 C ʋ J amiR-MGD1 D dgd1 I־ŠҒλ&IёŠԿЊg ӈȉ6c0DC0IԿЊG'.cs®vҀԊ I՗ӂɲgӧR<ׄ̈ϒҶׅ

ёŠˮIҀԊՙIхˮϖFĻ̕gұ?<D0

ƪ

ƪ6. ALAĸĸζζɜɜIIdgd1GG''..ccPchlideƇƇʊʊÃÃլլôôIңАА

־ŠҒλ&GALAg24ˮլĸζ4<ɜIPchlideƇʊÃլôIңАՙg՘λ̦Ddgd1C͖Ԟ4<Pchlide IƇʊхԔDƆÃլôI̸Էgw®šI½GϷ4<ׄFujii et al., 201819)gˌǍwww.plantphysiol.org.

Copyright American Society of Plant Biologists.ׅ

(9)

eğΠȐʼn(ağΠȐ12ˮլɜWCJҒλ

&Is®vҀԊIƍՙJ՘λ̦G'$B[T DhELJ&F$0D)Ļ(?<0I0D(a PLB (aˆ®zlŽ҅QIǍˀJs®vҀԊ ILJőgɨӂD4F$0D)Ϸ2d<W<

+DEX amiR-MGD1Ddgd1CJ:d;dMGDG DDGDGIՙ)ёŠԿЊgԵ4BȶGñ$±ž

°Gʙń2dB$<0daḬΩôG'$B ğΠȐɜIǬҡIҐоôI̸ԷǍŠg֋Ǭ֣ɤ դCхˮϖGӈȉ4<D0e+DEX amiR-MGD1 CJPLB I͸ǘJ−DEX Dƈ̹GԶ](Gԍ*

<):IɜIˆ®zlŽ҅ɒʊ)ɐ,ձȀ2d

<:dGȌ4dgd1CJPLBI͏φΩ)F(

F(͸ǘ87:IɜIˆ®zlŽ҅Iɒʊ[ձ Ȁ2dc0D)Ļ(?<E>aḬΩôG'$

B[LPORI±ž°JğΠȐɜGԶ](G΀ȕ 4<·˝v²²šk°] LHCII IÅӂ†´–

vԊC#cLHCB1IңАJɐ,ձȀ2d<

F9amiR-MGD1Ddgd1IլCJPLB(a

ˆ®zlŽ҅QIǍˀΪG PLB I͸ǘG'$

BǓ*FՂ$)Ӄad<I=e%(׌@Iƀ ѿɲD4BDGDG )®¦®ҀԊC#cIGȌ 4MGDG JūέCҀԊÑ՗ȞgɒʊC*F$

“´®¦®ҀԊC#c0D)ʫ/adcׄƪ2ׅ DEX ĸζ4< amiR-MGD1 I־ŠҒλ&CJ MGDG Iƍՙ)ΪϊϖG΀ȕ4B$c<Z DGDG IŎƇ)Եȶ`b[ָ$ΫʃGF?B$

cDGDG JūέC®¦®̸ԷgDc0D)C

*cICğΠȐɜJ LPOR IĻӊGí$ PLB I̩Ǭ̸Է)ȥlj4PLBJ:IWWDGDG] áI®¦®ҀԊG`bʑȹFҥ̸҅ԷGǍˀ2 dcI=e%4(4:IɜJ LHCII FEI ğƇʊ҅†´–vԊ)ңА87W<s®vҀ ԊƇʊIʙń[ь,ICҡёôɒʊJձȀ2d

<WWGFcDʺ΃2dc·˝dgd1 Ǎϊô CJDGDGIՙ)֣ҢG΀ȕ4B$c<ZPLB CJMGDGIŎƇ)ϊȶGָ$ΫʃGF?B$

cğΠȐɜLPOR JĻӊ2d PLB I̩Ǭ̸

ԷJȥdc)͏2d<MGDGJūέC®¦®

̸ԷgDdF$ICPLBg÷?B$<҅J®¦

®GǍˀ2dc0DF,Ķօ4<WWէ,͏c I([4dF$4(4F)adgd1 G'$B

[ˮլIхԿDĦGPLB J͸ǘ4ˆ®zlŽ҅

)ϒՁ6cin vitro Iǻֶ(a“´®¦®Ҁ

ԊIMGDG[LHCIIDϠÓ÷ν6c0DC®¦

®̸Էg÷dc0D)ϪadB$c21)ğθdž½ Cλѹ4<dgd1J՘λ̦DƈЊɃGLHCIIgң АC*c0D(a22)ɝGңА4< LHCII F E)MGDGDϠÓ÷ν6c0DC®¦®ҀԊ C#cDGDG gTDhEͅ$B[˿уϖGJˆ

®zlŽ҅gɒʊC*cI([4dF$

¼ӑIʺ΃)E0WC͋4$I(Gő&F9 s®vҀԊIͅ˅G`?Bv²²šk°Ƈʊ хԔ]ğƇʊ†´–vԊIңА)ձȀ2dcI (J¾˦C#cW<ʋJv²²šk°I ƇʊGő&̧]Ґоôz¶ŽIğƇʊՈìǬI mRNA ңА[s®vҀԊIƇʊձȀG`?B ʙń2dc0DgӃĹ4B$c)12,14)ׄ̈ϒҶׅ

:d)EI`%F¦r¥Cԍ0cI([˦

a(CF$s®vҀԊGő&ˆ®zlŽ҅

GJՔɲҀԊC#c €šiˆ°w¯‚²¶

°D€° t“¡~°j~°w¯‚²¶°) Åӂ̸ʊʊĻD4BƍWdc)oˆqœ®€

IϒՁ]ҡёôQIĻŠIԿЊG'.c0da IՔɲҀԊIɖŎ[ӊ˦6R*Ӧ֠D4B͏2 dB$cÞɜ2aGϮЕgԺZ0daIύƛ g˦a(G4B$*<$

Ӱ ӰԢԢ

ˢ̊ǟǬǓDZIȔ̖ɷ2hGJ֋Ǭ֣ɤդӊ

̕G'$BǑǓFcӺΰg$<=*W4<0I ǂgċbBű,ɡϸπ4¼/W6̊ϮЕJ JSPS ЄϮԂׄ16J10176 2671101626440170ׅ Iœʊgż.BұfdW4<

Received Nov 15, 2018; Accepted Nov 20, 2018; Published Dec 31, 2018.

ŵ ŵѮѮ˙˙ΰΰ

1. Solymosi, K. and Schoefs, B. (2010) Etioplast and etio-chloroplast formation under natural conditions:

The dark side of chlorophyll biosynthesis in angiosperms. Photosynth. Res. 105, 143–166.

(10)

2. Masuda, T. and Takamiya, K. (2004) Novel insights into the enzymology, regulation and physiological functions of light-dependent protochlorophyllide oxidoreductase in angiosperms. Photosynth. Res. 81, 1–29.

3. Fujita, Y. (1996) Protochlorophyllide reduction: a key step in the greening of plants. Plant Cell Physiol.

37, 411–421.

4. Tanaka, R., Kobayashi, K. and Masuda, T. (2011) Tetrapyrrole metabolism in Arabidopsis thaliana. The Arabidopsis Book 9, e0145.

5. Dorne, A., Joyard, J. and Douce, R. (1990) Do thylakoids really contain phosphatidylcholine ? Proc.

Natl. Acad. Sci. U.S.A. 87, 71–74.

6. Selstam, E. and Sandelius, A.S. (1984) A comparison between prolamellar bodies and prothylakoid membranes of etioplasts of dark-grown wheat concerning lipid and polypeptide composition. Plant Physiol. 76, 1036–1040.

7. Shipley, G.G., Green, J.P. and Nichols, B.W. (1973) The phase behavior of monogalactosyl, digalactosyl, and sulphoquinovosyl diglycerides. Biochim. Biophys.

Acta 311, 531–544.

8. Selstam, E. (1998) Development of thylakoid membranes with respect to lipids, in Lipids in Photosynthesis: Structure, Function and Genetics (Siegenthaler, P.A. and Murata, N., Eds.) pp 209–224, Kluwer, Dordrecht, The Netherlands.

9. Kobayashi, K. (2016) Role of membrane glycerolipids in photosynthesis, thylakoid biogenesis and chloroplast development. J. Plant Res. 129, 565–

580.

10. Kobayashi, K., Kondo, M., Fukuda, H., Nishimura, M. and Ohta, H. (2007) Galactolipid synthesis in chloroplast inner envelope is essential for proper thylakoid biogenesis, photosynthesis, and embryogenesis. Proc. Natl. Acad. Sci. U.S.A. 104, 17216–17221.

11. Kobayashi, K., Awai, K., Nakamura, M., Nagatani, A., Masuda, T. and Ohta, H. (2009) Type-B monogalactosyldiacylglycerol synthases are involved in phosphate starvation-induced lipid remodeling, and are crucial for low-phosphate adaptation. Plant J. 57, 322–331.

12. Kobayashi, K., Narise, T., Sonoike, K., Hashimoto, H., Sato, N., Kondo, M., Nishimura, M., Sato, M., Toyooka, K., Sugimoto, K., Wada, H., Masuda, T.

and Ohta, H. (2013) Role of galactolipid biosynthesis in coordinated development of photosynthetic

complexes and thylakoid membranes during chloroplast biogenesis in Arabidopsis. Plant J. 73, 250–261.

13. Kobayashi, K. and Ohta, H. (2008) Possible requirement of galactolipids for embryogenesis, in Photosynthesis. Energy from the Sun (Allen, J.F., Gantt, E., Golbeck, J.H., Osmond, B., Eds.) pp 783–

786, Springer, Dordrecht, The Netherlands.

14. Fujii, S., Kobayashi, K., Nakamura, Y. and Wada, H.

(2014) Inducible knockdown of MONOGALACTOSYLDIACYLGLYCEROL

SYNTHASE1 reveals roles of galactolipids in organelle differentiation in Arabidopsis cotyledons.

Plant Physiol. 166, 1436–1449.

15. Fujii, S., Kobayashi, K., Nagata, N., Masuda, T. and Wada, H. (2017) Monogalactosyldiacylglycerol facilitates synthesis of photoactive protochlorophyllide in etioplasts. Plant Physiol. 174, 2183–2198.

16. Kelly, A.A., Froehlich, J.E. and Dörmann, P. (2003) Disruption of the two digalactosyldiacylglycerol synthase genes DGD1 and DGD2 in Arabidopsis reveals the existence of an additional enzyme of galactolipid synthesis. Plant Cell 15, 2694–2706.

17. Dörmann, P., Hoffmann-Benning, S., Balbo, I. and Benning, C. (1995) Isolation and characterization of an Arabidopsis mutant deficient in the thylakoid lipid digalactosyl diacylglycerol. Plant Cell 7, 1801–1810.

18. Lin, Y., Chen, L., Herrfurth, C., Feussner, I. and Li, H. (2016) Reduced biosynthesis of digalactosyldiacylglycerol, a major chloroplast membrane lipid, leads to oxylipin overproduction and phloem cap lignification in Arabidopsis. Plant Cell 28, 219–232.

19. Fujii, S., Kobayashi, K., Nagata, N., Masuda, T. and Wada, H. (2018) Digalactosyldiacylglycerol is essential for organization of the membrane structure in etioplasts. Plant Physiol. 177, 1487–1497.

20. Kowalewska, Ł.M., Mazur, R., Suski, S., Garstka, M.

and Mostowska, A. (2016) Three-dimensional visualization of the internal plastid membrane network during runner bean chloroplast biogenesis.

Plant Cell 28, 875–891.

21. Simidjiev, I., Stoylova, S., Amenitsch, H., Javorfi, T., Mustardy, L., Laggner, P., Holzenburg, A. and Garab, G. (2000) Self-assembly of large, ordered lamellae from non-bilayer lipids and integral membrane proteins in vitro. Proc. Natl. Acad. Sci. U.S.A. 97, 1473–1476.

(11)

22. Härtel, H., Lokstein, H., Dörmann, P., Grimm, B. and Benning, C. (1997) Changes in the composition of the photosynthetic apparatus in the

galactolipid-deficient dgd1 mutant of Arabidopsis thaliana. Plant Physiol. 115, 1175–1184.

Role of galactolipids in etioplast development and etioplast-to-chloroplast differentiation Sho Fujii1, Noriko Nagata2, Tatsuru Masuda1, Hajime Wada1 and Koichi Kobayashi3,*

1Graduate School of Arts and Sciences, The University of Tokyo, 2Faculty of Science, Japan Women’s University,

3Faculty of Liberal Arts and Sciences, Osaka Prefecture University

(12)

®

®Â Âó óƙ ƙ I ƾ ƾΠΡ ¡0 0´ ´ñ ñDz DzÎ ÎŒ ŒŐ Ő0 0DŽ DŽĹ Ĺ0 0Ǖ Ǖý ý

āĀêó Ÿ´ǥâƂƈóƀƏĥ h€l~„Z~…^…ûŧކǟǿőƽ†ǟ*

ǣƛŹűà®Îģǵñǘƙ/*®MhR…E¸Ų!*p}ZcXHfƒ<!1XcS

wcEǣÂ!nL€dQXƒEǞ¬"B®ÂóƙIȁPSIȂ11115Wo{f`c,100-3000V nGS]…@Őģ CBƾdz.ŐǔE<)ƾΡ+BPSI ƾΡ0Őģģ´0űÂóźDŽņ, ŐǔDŽņ1êǕý!&űÎģ0´ñŒŐ0DŽĹ1ǖC*BPSI ƾΡ0Ƶƍ/Ǥƿ+

B,ƨ@C*&ƴƤ¡V…d0Ȇ)0ÛñȁYcf3,Ycf4ȂPSIƾΡ0Wo{f`c0´ñ DzÎ/Ǭ‹"B,Eƃ"żĭź.Ljīƃ C&ŁDŽNJ+1ƕƩ@0PSIƾΡ0´ñDzÎ0 Ŀǒ0ƀƏEě/Ɲ—"B

1. JJ55ZZGG

§͛gՔŠ4NADP+gՋĜ6cՔоϒλƵğƇ ʊ֋ǬìՁзJğo’°u¶głν4BϟľG

͂ѿ6c׍@IğŠDZзG`bִŗ2dcğŠ DZзIIׄPSIIׅJɐ$ՔŠŏgɒʊ4͛gĻӊ4

œ®€t“´ׄPQׅgՋĜ6c)ğŠDZзI

ׄPSIׅJœ®€t“´ׄPQׅD~v²¥b6f

ׄCyt b6fׅ(aI֋Ǭgœ®€~j´ׄPcׅ [4,J~v²¥cׄCyt cׅgß4Bż.Żb

šn±Žt~´ׄFdׅgՋĜ6cՋĜƵFdJ

šn±Žt~´ NADP ՔŠՋĜՓоׄFNRׅG

֋ǬgìՁ4NADPHgλʊ6c0I`%Fϟ գƵ֋ǬìՁзׄ¯j֋ǬìՁзׅG`bՔо )ϒλ4ΚՔƫǺŹɫGɨӂFՋĜŏC#c

NADPHgλʊ6cW<0IŹɫDĦɖ4B

ˆ®zlŽ҅gß4<œ²´ִŗŏׄpmfׅ) ɒʊ2dATPƇʊՓоIĕ*CATP)Ƈʊ2 dc·˝FdgՋĜ4<֋ǬJCyt b6fQìՁ 2dĮN PSI gՋĜ6cхԔ)#c0dJ

NADPH gλʊ4F$) ATP ƇʊGɨӂF pmf

gɒʊ4ɣθϖ֋ǬìՁзׄ|lv¯‰v֋Ǭ ì Ձ з ׅ D ƒ K d B $ c PSI I Ջ Ĝ ď C J

~´¢m¥ˆ®zlŽ҅ԎҿƇôI̸ԷD‡l¤v€

*ԸчĞE-mail: [email protected]

NADP+äǐG[Ио]ϱ־gՋĜ6cхԔG֋

Ǭ)ìՁ2d<bҡёôIՓоIͯɲgՋĜŏ G`bӧЩ6c±Ž‰v€ńɡGɨӂF֋Ǭg üш4<b6c0I`%GPSIIՋĜďJǑ͂

ѿC#bğƇʊ֋ǬìՁзI֋ǬIĻՒG'$

BÃɧϖF͂ѿg̘<4B$c

ğŠDZзҿƇôIλƇʊIӊ̕JȂ˧CJF

$)PSIIJãӰƥԛ)Զ$<ZĻǬօƇ̸͂

Iӊ̕J͖ԞϖԺhC$<0dGȌ4BPSI JãӰƥԛ)Լ,ҿƇôIλƇʊIӊ̕JȂ˧

CF$4(4PSIҿƇôIλƇʊgʢ%DѮ

&adcƦǬׄĻǬօƇƦǬׅIƈǺ)ԺX0 daIƦǬgͅ˅4<Ǎϊ̦Iӊ̕(a PSI ҿ ƇôIλƇʊIĻǬ̸͂Iӊ˦)ȕ47@ԺȜ 4B$c

2. PSIҿҿƇƇôôII||››««‰‰̸̸ԷԷ

PSIҿƇôJǑ˕I|›«‰(a̸ʊ2d cׄҶ1ׅ|›«‰фʊIϊFcҿƇôI Ǻѩgƪ׌GWDZ<ŹɫÃɧҿƇôׄPSI RCׅ JŹɫÃɧ|›«‰PsaA DPsaB (a̸

ʊ2dcˆ®zlŽ҅gɐ$τ֐ͯɲŋׄñ΂

CñΙɃISDSׅCƀΊŠ6cDŹɫÃɧׄRCׅ

DŽNJ

(13)

ƪ

ƪ1. €€²²££ďď((aӃ<<PSIҿҿƇƇôôII|›««‰‰̸̸ʊʊ

PSI RCהŹɫÃɧ|›«‰PsaADPsaB(a̸ʊ2dcPSIŹɫÃɧҿƇôPSI coreהŹɫÃɧҿƇ

ôGáIPSI|›«‰gцƇ6cPSIzjҿƇô€²£ďGPsaC/D/E|›«‰)v®€†¶

̸Էgɒʊ6c½ďGJPsaFDPsaJ)ǯƱ4PsaFJ°¶¦´ďGӇ͛ϖF̸Էg[>œ®€~

j´цƇՍðgɒ ʊ6cPsaF IŹȌďI̴G J PsaH/PsaI/PsaL )v®€ †¶̸Էgɒ ʊ6c PSI-LHCIה̰ΩCJPSI coreIPsaF/PsaJďG4ՙôIj´‹ҿƇôLHCIׄLHCI4ׅ)цƇ6cv®¤

ާ€IPSI-LHCIGJ8ՙôILHCIׄLHCI8ׅDPsaGDPsaHIլG2ՙôILHCIׄLHCI2ׅ)ǯƱ 6cPsaGDPsaKJLHCI4'`NLHC8IцƇgǵǺŠ6co´ŽmIPSI-LHCI̸Էׄ5L8RׅgƼG 4B÷ʊ4<

(14)

|›«‰Gv²²šk°)цƇ4<v²²

šk°†´–vԊׄCPIׅ)ūֈ2dcPSI z jҿƇôׄPSI coreׅJPSI|›«‰ITD hEgцƇ4<ҿƇôC#cՔоϒλƵğƇʊ λΩIPSI coreG10|›«‰ׄPsaA-FPsaI-Lׅ )ăǯ2dcׄҶ1ׅPsaGPsaHPsaNPsaO J̰Ω]ёҪׄv®¤Ž§€ׅGǯƱ6c)

~j“•v‹¯jGJǯƱ4F$:dGŹ4B PsaM D PsaX JÅG~j“•v‹¯jGǯƱ6 cPsaP J̰ΩGǯƱ6c¯´ՔІ´–vԊ C#cPSIzjҿƇôGJй100ĻǬIChl a й20ĻǬIβ-r²‹´2 ĻǬIšt“´

ׄšk²t“´5-§“—޲t~šk²t“´

[4,J¦t“´-4ׅ3@I4Fe-4SƵI՜ϱ

־ÃɧׄFXFAFBׅ)ǯƱ4ğʱօµğŠDZ Źɫµ֋ҝĻֈIǵǺе֋ǬìՁʊĻׄPc ]

Cyt C'`NFdׅIՔеՋĜŹɫgұ%~j

“•v‹¯jIˆ®zlŽ҅g΂ƔFτ֐ͯɲ ŋCƀΊŠ6cD3ՙôIzjҿƇô)ūֈ2 dc3 ՙô̸ԷIǵǺŠGPsaL)՗ӂFɖŎ g̘<4B$c·˝̰ΩDёҪIzjҿƇô JūՙôCPsaF/PsaJ |›«‰ďGօğɲ v²²šk°†´–vҿƇôׄLHCIׅgцƇ4 PSI-LHCIԎҿƇôgɒʊ6cLHCIIPSI core QIцƇJǵǺC̰ΩDёҪIˆ®zlŽ҅g

΂ Ɣ F τ ֐ ͯ ɲ ŋ C ƀ Ί Š 4 B Ļ σ 6 c D PSI-LHCI)ɠadc

zjҿƇôIÅӂFՍĻI̸ԷJ~j“•v

‹¯jD̰ΩC`,ăǯ2dB$cŹɫÃɧҿ ƇôׄPSI RCׅJ҅ӽԵ¯‰v€g11[@2

@IϠƈFŹɫÃɧ|›«‰ׄPsaADPsaBׅ (a̸ʊ2dĿ̆֋Ǭü¿ôI P700ҿ˕I

֋ǬżȂôTDhEIj´‹v²²šk°

ׄzjµj´‹ׅgцƇ6c˿уϖF֋Ǭż ȂôC#cFADFBJ€²£ďIҶ֐GǯƱ 6cPsaCGцƇ6cPsaCJPsaDDPsaEĦG

€²£ďGšn±Žt~´цƇ֜ƹgɒʊ6 c·˝҅ӽԵ¯‰v€g1@[@PsaFJ :I N ̉ďIӇ͛֜ƹg°¶¦´ďGǓ*,ɏ bĹ4°¶¦´ďIҶ֐GцƇ6cPsaNDĦ Gœ®€~j´цƇՍðI·Սgɒʊ6c

PsaJ J҅ӽԵ¯‰v€g 1 @[@Ȕ2F|›

«‰CPsaFGւʹ6c

3. PSIҿҿƇƇôôIIĻĻǬǬօօƇƇJJǑǑ͑͑ռռԿԿЊЊCC##cc 0dWCGPSI|›«‰gz¶Ž6cՈ ìǬIԛı]ѭӕńɡIӊ̕JͯϒGԺZad B*<)Ƈʊ2d<|›«‰]zšiv

†¶)ҿƇôQĻǬօƇ6c̸͂Iӊ˦JԼd B$<PSIҿƇôI̸Է(aƆ̸ʊ|›«‰

J֙ɀ͋4,͑ռϖGĻǬօƇ6cD̆ɚ2 dcҿƇôIÃɧGǯƱ6cŹɫÃɧ|›«

‰)‹²‡l£¶C#cPSI RCIɒʊ)

˿ Ŀ I Ļ Ǭ օ Ƈ Կ Њ C # c ҡ ё ô z ¶ Ž I PsaAPsaBPsaC Iͅ˅Ǎϊ̦g:d;dν$

<†´–vԊI–°€®ž°Iǻֶ(a˿ĿG PsaB )ˆ®zlŽ҅Gʯģ2d:dģD4 BPsaA )âő2d:IɜGPsaC )цƇ6c DǁƏ2dB$c19)4(4ĻǬօƇԿЊJ̴

ZBԶ,Ժұ6c<ZĻǬօƇÃլôg̱Ĺ6 c0DJ։4$ğƇʊƣǹgͯϒGƇʊ6cȌ

˕LJ͐̆Iсѽgν$<ӊ̕(aPsaGDPsaK g[<F$PSI-LHCI subcomplexCf7(GңА 6c0D)Ϸ2d<9,18)0dJPsaG D PsaK IцƇԶɃ)Լ$<ẔĹ2d<ĻǬօƇÃլ ôC#cPSIҿƇôIĻǬօƇԿЊIӊ̕g˼

GԺZcGJ˜<Fʒͪgν$c0D)ɨӂC

#c

4. PSIҿҿƇƇôôIIĻĻǬǬօօƇƇGGխխ¿¿66ccƦƦǬǬ

Ǒ˕I PSI |›«‰)ҿևFҿƇôG͋

ϲGĻǬօƇ6cԿЊJ҈ϒϖGԍ0cDJѮ

&G,$PSI|›«‰IĻǬօƇgßͽ&

6cƦǬׄĻǬօƇƦǬ[4,JĻǬ~¨Ÿ²´ׅ

)ǯƱ6cJ7C#cҶ 2 GWDZ<ƦǬ) PSIҿƇôIĻǬօƇGխ¿6cDѮ&adc̨

ʤJׄ׌ׅͅ˅̦CJPSIҿƇôIңА)Ǔ*

,΀ȕ6c)PSI|›«‰gz¶Ž6cՈ ìǬIϒδJɕ֕gTDhEż.F$ׄ׍ׅPSI

|›«‰DϠÓ÷ν6cFEC#cΪG ɜѯCJġώͣ͒ͪ]Փ͔ж”l›¯‰Ž~

€‹¥ׄYeast Two-Hybrid SystemׅG`cլʹϖ

(15)

F ˝ ͪ G ` c Ϡ Ó ÷ ν ) ǁ Ə 2 d B $ c

Ycf31,6,11)Ycf41,4,8,16)PYG7/CGL71/Ycf37̰ׄΩ/

v®¤Ž§€/~j“•v‹¯jCIƉЇׅ

3,13,17)JϤ̧ğƇʊλΩD~j“•v‹¯jGă

ǯ2dB$c<ZΪG՗ӂFɖŎgʢ?B$c Dʺ΃2dcϤ̧ğƇʊλΩCJYcf3DYcf4 J ҡ ё ô y “ ¥ G z ¶ Ž 2 d B $ c ) Y3IP1/CGL59̰ׄΩ/v®¤Ž§€CIƉЇׅ

J̧y“¥Gz¶Ž2dB$c:IáIƦǬ Y3IP14,7)PSA22,15)PSA312)PPD15)J̰ΩDҪ

֥GӃĹ2ḑy“¥Gz¶Ž2dB$c Ycf3 D Y3IP1/CGL59 J TPRׄtetratricopeptide

repeatׅަl´g[>áI†´–vԊDϠÓ

÷ν6cƀѿɲ)Ѯ&adc

5. Ycf3DDY3IP1JJ§§ªª¶¶°°ggɒɒʊʊ44ŹŹɫɫÃÃɧɧ I

IĻĻǬǬօօƇƇGGɨɨ֚֚CC##cc

ҡ ё ô y “ ¥ G ǯ Ʊ 6 c ycf3ׄhypothetical chloroplast open reading frame 3ׅJϤ̧ğƇʊ λΩGăǯ2dcӥXŻbׄ̚ORFׅD4BƈǺ 2d~j“•v‹¯jG[ăǯ2dB$c:

Iɜˆ®zlŽ҅IҶƱɲ†´–vԊC#c Ycf3gz¶Ž6c0D)Ϸ2d<1)ҡёôɒԊ ԛˀG`b ycf3 ՈìǬg“‰vjm4<v®

¤Ž§€D†•zIɒԊԛˀôGJ PSI )Ĥ ,ңА4F$<ZYcf3J PSI ҿƇôIĻǬօ ƇGɨ֚IƦǬC#c1,11)Ycf3JՍĻϖGǍɲ 4<PSIŹɫÃɧ|›«‰PsaA]zjҿƇ ô|›«‰IPsaDDϠÓ÷ν6c0DgϷ 6in vitroIӊ̕ц̘)ǁƏ2d<6)†´–v ԊլIϠÓ÷νGխfcDѮ&adB$c TPR

ަl´g[@Ycf3 JPSIҿƇôIĻǬօƇ

Gխfcɨ֚ƦǬC#c0DJϲ(C#c)E I`%G PSI ҿƇôgĻǬօƇ28cI(G@

$BJTDhE¾˦C#?<

: 0 C П ѯ a J HAׄHuman influenza hemagglutininׅ†wׄYPYDVPDYAׅg C ̉М GүƇ4<Ycf3ׄYcf3-HAׅgҡёôGϒδ28

<v®¤Ž§€ҡёôɒԊԛˀôg÷Ĺ4 Ycf3 D Ϡ Ó ÷ ν 6 c † ´ – v Ԋ g j š k 

‹k¶еҾ4<ׄƪ2Aׅɠad<Ycf3-HA̻ ƗGJY3IP1ׄYcf3 Interacting Protein 1ׅ)ӃĹ 2d<Y3IP1J†•zYcf3Dœ°‡m´6c

†´–vԊD4BˡGǁƏ2dB$c4)v®¤

ާ€CJjšk‹k¶еҾG`bָ$м ɃI̻Ɨ)ɠadYcf3DY3IP1 )TUɑՙǯ Ʊ6c0D)˦a(GF?<4(4PSI¢¯

ŸœˆŽJTDhE̱Ĺ2dF(?<D0e) ʂɃIָ$mn€†´Ļ̕DԊՙĻ̕gұ%D PSIŹɫÃɧ|›«‰ׄPsaA DPsaBׅ)̱

Ĺ2d<7)

Ĥсѽ†´–vԊgˍȐɲƈðĜоI35SIǯ Ʊ½Cƻ֭4ϫˮլ̻ӱׄ–°€®ž°ׅ4<ɜ сѽgϰϯ4ˆ®zlŽ҅gеҾ4:dgτ

֐ ͯ ɲ ŋ C ƀ Ί Š 4 B Ycf3-HA g j š k 

‹k¶еҾ4<6cD(Fbɐ,®ž°2d<

PSI RC|›«‰)̱Ĺ2d˜ӄGƇʊ2

d<PSI RC|›«‰)Ycf3GΪϊϖGцƇ 4B$c0D)Ļ(?<͆GсѽgΝ̻ӱI

32S IǯƱ½CЈ4˜ӄƇʊ2d<†´–vԊ gԮԓׄˆnl€ׅ6cDYcf3HA̻ƗIPSI RC |›«‰I®ž°JǓ*,΀ȕ4<

ɟ?BYcf3JY3IP1 D§ª¶°gɒʊ4

˜ӄGƇʊ2d<PSI RC|›«‰g·Կɲ

(16)

ϖGцƇ4PSI RCIĻǬօƇgßͽ&6cƦ ǬC#cDцӪ4<7)

–°€®ž°Iǻֶ(a Ycf3-HA JԶ,Ƈʊ 2dc)¾ǵǺCY3IP1JԼ,Ƈʊ2dc)ǵ ǺC#c0D[˦a(GF?<Ycf3 IãӰƥ ԛԶɃJYcf3GцƇ6cŹɫÃɧ|›«‰

I:dDTUƈ5C#?<)ãӰƥԛ)Զ$0 DI͂ѿϖFζοJ¾˦C#cY3IP1J҅ӽԵ

֜ƹg[@†´–vԊC#c)Y3IP1ͅ˅̦C [Ycf3JTU͋ȶGңА4Ycf3Iˆ®zlŽ

҅QIцƇIǵǺɲJ͋ȶC#?<ɟ?B

Y3IP1JYcf3Iˆ®zlŽ҅QIцƇgǵǺŠ

28cICJF,ãӰƥԛ6cYcf3)ˆ®z

lŽ҅¼I͋4$ðѣGǯƱ28cɖŎg[@

I([4dF$

6. Ycf4§§ªª¶¶°°JJPSIzzjjDDPSI-LHCIҿҿƇƇ ô

ôIIĻĻǬǬօօƇƇGGխխ¿¿66cc

PSI RCIĻǬօƇGɌ*ь*ƐԤՍIzj|

›«‰)ĻǬօƇ2dc0IԿЊGɨӂF I)Ycf4C#c0D)˦a(GF?<Ycf4J N ̉МâԩG׍@I҅ӽԵ¯‰v€g[>

Ycf3 `bǵǺGˆ®zlŽ҅GцƇ6cˆ®

zlŽ҅g΂ƔF̐çCƀΊŠ4B~¬жȅɃ ŞՒԎՃɧͪCYcf4gĻֈ6cDӃ(.IԊ

ՙ)1,000kDaЊɃI(FbǓ*F̸Էgɒʊ4

B$c0D)Ļ(?<1):0CҡёôɒԊԛ ƪ

ƪ2. Ycf3-Y3IP1§§ª¶¶°°JJPSI RCIĻǬǬ օ

օƇƇGɨ֚֚CC##cc

Aהjšk‹k¶еҾ4<Ycf3-HA I†´–

vԊфʊ1הYcf3-HAgƍYˆ®zlŽ҅gƀ

ΊŠ4BеҾ4<̻Ɨ2ה՘λƵIˆ®zlŽ

҅gƀΊŠ4BеҾ4<̻ƗׄȌΠǻֶׅBה

Ycf3-HAеҾ̻ƗImn€†´Ļ̕Cה†´–

vԊI–°€µˆnl€ǻֶĤсѽ†´–v Ԋg35SC–°€®ž°4<ɜˆ®zlŽ҅'

`NYcf3-HAgеҾ4®ž°2d<†´–v

Ԋg̱Ĺ4<ׄPׅƈ̹Gˆnl€4<ɜGˆ

®zlŽ҅'`NYcf3-HAgеҾ4®ž°2 d<†´–vԊg̱Ĺ4<ׄCׅ

ƪ

ƪ3. Ycf4§§ª¶¶°°JJPSI coreDDPSI-LHCI I

ĻǬǬօօƇƇGխ¿¿66cc

Aהjšk‹k¶еҾ4<HA-Ycf4 I†´–

vԊфʊ1הHA-Ycf4gƍYˆ®zlŽ҅gƀ

ΊŠ4BеҾ4<̻Ɨ2ה՘λƵIˆ®zlŽ

҅gƀΊŠ4BеҾ4<̻ƗׄȌΠǻֶׅ3ה еҾ4<PSI-LHCI̻ƗPSIDLHCII¢¯Ÿ

œˆŽ)̱Ĺ2dcBה†´–vԊI–°€µ

ˆnl€ǻֶĤсѽ†´–vԊg35SC–°€

®ž°ׄPׅ[4,Jˆnl€ׄCׅ4<ɜˆ

®zlŽ҅ׄTMׅ'`NHA-Ycf4gеҾ4®

ž°2d<†´–vԊg̱Ĺ4<

(17)

ˀͪG`bYcf4IN̉МGHA†wgүƇ28

< HA-Ycf4gǓՙϒδ28ƀΊŠ4<ˆ®z lŽ҅(aϫˮլGjšk‹k¶еҾ6cD PSIҿƇôIĻǬօƇÃլôgцƇ4<HA-Ycf4

̻Ɨ)ūֈC*<7):I¢¯ŸœˆŽфʊgӧ RcDʼnԬI Ycf3-HA ̻ƗG͖RBǯƱ6c PSI|›«‰I˕)(FbǑ(?<ׄƪ3ׅ 0I0DJYcf3-Y3IP1§ª¶°G`cŹɫà ɧҿƇôIĻǬօƇGɌ*ь,ԿЊG Ycf4 §

ª¶°)խ¿4B$c0DgϷ6еҾ4<

HA-Ycf4 ̻ƗGJǑ,I PSI zj|›«‰

IáGLHCI|›«‰[ǯƱ4<mn€†

´Ļ̕CӧRcDPsaGPsaKPsaNPsaOä ǐIzj|›«‰J6RḆĹ2d<W<

Lhca2DLhca9äǐIגЋILHCI|›«‰

[̱Ĺ2d<ɟ?BYcf4§ª¶°JPSI core IĻǬօƇ=.CF,PSI-LHCI IĻǬօƇG WCխ¿4B$c0D)Ϸ2d<–°€µˆn l€ǻֶJPSIzj|›«‰)·ԿɲϖG

®ž°2dc0DgϷ4˜ӄƇʊ2d< PSI

|›«‰) HA-Ycf4QцƇ4B$c0D) ϲ(Zad<4(4LHCI|›«‰J®

ž°2dF(?<

еҾ4< HA-Ycf4 g~¬жȅɃŞՒԎՃɧͪ

G(.cDPSI coreDPSI-LHCIGϠɑ6cσ ĻGёI•´Ž)Ļֈ2d<ׄƪ4ׅ:d;d I¢¯ŸœˆŽфʊgӊ̕6cDPSIzjGϠ ɑ6c•´ŽJPsaGPsaKPsaNPsaOäǐ Izj|›«‰J6RḆĹ2d<·˝

PSI-LHCIGϠɑ6cσĻGJ¼ӑI PSIzj|

›«‰IáG Lhca2 D Lhca9 äǐIגЋI LHCI|›«‰)̱Ĺ2d<0daIц̘

JHA-Ycf4JPSI coreDPSI-LHCI subcomplex GϠɑ6cĻǬօƇÃլôgцƇ4B$c0D gϷ4B$cPSI coreI|›«‰J†´–

vƇʊGí$(FbϫˮլGҿƇôGфXԦW dc)ˆ®zlŽ҅Gœ¶°2dB$<ȕՙI LHCI|›«‰)PSI coreGцƇ4<DѮ&

adcPSI ͅ˅̦Iˆ®zlŽ҅GJq¯{

£¶ΫʃILHCI)ǵǺGңА6c14)':a, LHCI|›«‰IƇʊJPSI core|›«‰

IƇʊDǶĤGJĦɖ4B'a7ͯϒGğƇ ʊƣǹgƇʊ4B$cҡёôCJLHCI)f7(

CJ#c)œ¶°2dB$cDѮ&adc

ƪ

ƪ4. Ycf4§§ª¶¶°°DцƇƇ66ccPSIĻĻǬǬօօƇƇÃÃլլôô

AהеҾ4<HA-Ycf4̻Ɨ(aĻֈ2d<PSIĻǬօƇÃլôY1DY2Jv²²šk°Dr²‹“lŽ

gƍY)Y3JΝҐCHA-Ycf4IXgƍYBהY1-Y3I¢¯ŸœˆŽg̝Ґ4ḆĹCהY1-Y3IPSI

¢¯ŸœˆŽgmn€†´Ļ̕G`ḇĹDהY1-Y3ILHCI¢¯ŸœˆŽgmn€†´Ļ̕G`ḇĹ

(18)

7. PSIҿҿƇƇôôIIĻĻǬǬօօƇƇ§§ŒŒ°°

0dWCGɠad<ĻǬօƇIӊ̕ц̘(a v®¤Ž§€I PSI ҿƇôIĻǬօƇԿЊg ƪ5GWDZ<ŹɫÃɧ|›«‰PsaAD PsaBJƇʊDƈˮG҅Qʯģ2dPSI RCgɒʊ 6cDѮ&adcYcf3-Y3IP1 §ª¶°J0

IPSI RCĻǬօƇGɨ֚FɖŎg̘<60I

Կ Њ G ' $ B Ź ɫ à ɧ I €  ² £ ď C J CGL71/Pyg7/Ycf373)DPSA312))°¶¦´ďCJ

PPD15)DPSA22,15))ɨ֚CJF$)՗ӂFɖŎg

̘<4B$cΪGPYG7/CGL71/Ycf37Jǡ͚̐

ç½CJ՗ӂC#c<ZՔо)ձȀ6cĻǬօ ƇԿЊGխf?B$ca4$:IԿЊD4BѮ

&adcIJ֋ǬżȂôI՜ϱ־ÃɧIфXԦ XC#cFXJPsaADPsaBGцƇ6c)FA

DFBJPsaCGцƇ6c$7d[՜ϱ־ÃɧJ ūֈ4<PSI core̻ƗCJǵǺC#c)|›«

‰gӊֈ28cDՔоGȌ4B̴ZB¾ǵ ǺGFcɟ?BPYG7/CGL71/Ycf37Jq¯{

£¶gɒʊ4B՜ϱ־ÃɧIƇʊՍð(aՔо gնŴ6cɖŎg̘<4B$cI([4dF$

PSI RC IƇʊJό͛ɲ)ָ$ŹɫÃɧ|›«

‰gĻǬօƇ4Ǒ˕Izšiv†¶gцƇ 28c<Z˿[ƨ։FĻǬօƇ͑ռC#b:

I<ZǑ˕IĻǬօƇƦǬ)խ¿4B$cDѮ

&adc·˝PSI RC)Ƈʊ2dcDáI PSI|›«‰IĻǬօƇJ͖ԞϖȂ˧C#c Dʺ΃2dc)0IԿЊG՗ӂFɖŎg̘<6 I)Ycf4 §ª¶°C#cPSI |›«‰

)PSI RCG͋4,цƇ6cԿЊgßͽ&42 aG¾ǵǺFĻǬօƇÃլôgœ²‹j¶ƒF EG`cĻӊ(aǴ?B$cI([4dF$

Ycf4 §ª¶°J LHCI IцƇG[խ¿4B$

cv®¤Ž§€IPSI-LHCIJLHCIg10|

ƪ

ƪ5. v®¤¤ŽŽ§§€€IIPSIҿҿƇƇôôI͑ռռϖϖĻĻǬǬօօƇƇ

2@IŹɫÃɧ|›«‰PsaADPsaB(aŹɫÃɧҿƇôQIĻǬօƇGYcf3-Y3IP1/CGL59§ª¶

°Jɨ֚IɖŎg̘<60IԿЊGPYG7/CGL71DPSA3)€²£ď(aPPD1DPSA2J°¶¦´

ď(aխ¿6cDѮ&adB$cɌ*ь*ŹɫÃɧIƐјGǯƱ6cȔƵI|›«‰)ĻǬօƇ4

PSI core)λʊ2dc~j“•v‹¯jIPSI coreҿƇôJ»ՙôgɒʊ4ĻǬօƇԿЊJǶÎ6cv

®¤Ž§€CJPSI coreGLHCI8)PsaFďGцƇ4PSI-LHCIIÃլôׄPSI-LHCI subcomplexׅ)ɒ ʊ2dc:IɜPsaGDPsaK)q¯{£¶LHCIIÁМGLHCI2)PsaH/PsaG/PsaBIԩĐGцƇ4

PSI-LHCIIĻǬօƇ)ǶÎ6cPSI core(aPSI-LHCIIĻǬօƇԿЊGJÅGYcf4§ª¶°)խ¿

6c

(19)

›«‰цƇ6c10)PSI coreIĻǬօƇDJ έК4BLHCII8ՙôC#cLHCI8gɒʊ4 PSI core I PsaF/PsaJ ď G ц Ƈ 4 PSI-LHCI

subcomplexgɒʊ6cȕF,D[0IĻǬօƇ

͑ռWCYcf4§ª¶°)խ¿6c˿ɜIĻ ǬօƇԿЊJLHCI8IцƇgǵǺŠ6c PsaG DPsaKD͏bI2ՙôC#cLHCI2IцƇC#

c0IԿЊG`bPSI-LHCI)Ƕʊ4Ycf4§

ª¶°(aӊֈ6cPSIҿƇôGJzšiv

†¶)Ǒ˕ǯƱ6c)0IĻǬօƇԿЊI§Œ

°GJzšiv†¶IфXԦXIĻǬ̸͂Jƍ WdB$F$0IԿЊIǓՍĻJ̈ӊ˦IWW C#b:Iӊ˦JÞɜIϮЕIԺȜĞɚ4<

$

Received Dec 11, 2018; Accepted Dec 13, 2018; Published Aug 31, 2018.

ŵ ŵѮѮ˙˙ΰΰ

1. Boudreau, E., Takahashi, Y., Lemieux, C., Turmel, M. and Rochaix, J.D. (1997) The chloroplast ycf3 and ycf4 open reading frames of Chlamydomonas reinhardtii are required for the accumulation of the photosystem I complex. EMBO J. 16, 6095–6104.

2. Fristedt, R., Williams-Carrier, R., Merchant, S.S. and Barkan, A. (2014) A thylakoid membrane protein harboring a DnaJ-type zinc finger domain is required for photosystem I accumulation in plants. J. Biol.

Chem. 289, 30657–30667.

3. Heinnickel, M. et al. (2016) Tetratricopeptide repeat protein protects photosystem I from oxidative disruption during assembly. Proc. Natl. Acad. Sci.

U.S.A. 113, 2774–2779.

4. Krech, K. et al. (2012) The plastid genome-encoded Ycf4 protein functions as a nonessential assembly factor for photosystem I in higher plants. Plant Physiol. 159, 579–591.

5. Liu, J. et al. (2012) PsbP-domain protein1, a nuclear-encoded thylakoid lumenal protein, is essential for photosystem I assembly in Arabidopsis.

Plant Cell 24, 4992–5006.

6. Naver, H., Boudreau, E. and Rochaix, J.D. (2001) Functional studies of Ycf3: its role in assembly of photosystem I and interactions with some of its subunits. Plant Cell 13, 2731–2745.

7. Nellaepalli, S., Ozawal, S. I., Kuroda, H. and Takahashi, Y. (2018) The photosystem I assembly apparatus consisting of Ycf3-Y3IP1 and Ycf4 modules. Nat. Commun. 9

8. Ozawa, S. et al. (2009) Biochemical and structural studies of the large Ycf4-photosystem I assembly complex of the green alga Chlamydomonas reinhardtii. Plant Cell 21, 2424–2442

9. Ozawa, S., Onishi, T. and Takahashi, Y. (2010) Identification and characterization of an assembly intermediate subcomplex of photosystem I in the green alga Chlamydomonas reinhardtii. J. Biol.

Chem. 285, 20072–20079

10. Ozawa, S. I. et al. (2018) Configuration of Ten Light-Harvesting Chlorophyll a/b Complex I Subunits in Chlamydomonas reinhardtii Photosystem I. Plant Physiol. 178, 583–595

11. Ruf, S., Kossel, H. and Bock, R. (1997) Targeted inactivation of a tobacco intron-containing open reading frame reveals a novel chloroplast-encoded photosystem I-related gene. J. Cell Biol. 139, 95–102 12. Shen, J., Williams-Carrier, R. and Barkan, A. (2017)

PSA3, a Protein on the Stromal Face of the Thylakoid Membrane, Promotes Photosystem I Accumulation in Cooperation with the Assembly Factor PYG7. Plant Physiol. 174, 1850–1862.

13. Stockel, J., Bennewitz, S., Hein, P. and Oelmuller, R.

(2006) The evolutionarily conserved tetratrico peptide repeat protein pale yellow green7 is required for photosystem I accumulation in Arabidopsis and copurifies with the complex. Plant Physiol. 141, 870–

878.

14. Takahashi, Y., Yasui, T.A., Stauber, E.J. and Hippler, M. (2004) Comparison of the subunit compositions of the PSI-LHCI supercomplex and the LHCI in the green alga Chlamydomonas reinhardtii. Biochemistry 43, 7816–7823.

15. Wang, Y.W. et al. (2016) The DnaJ-Like Zinc Finger Domain Protein PSA2 Affects Light Acclimation and Chloroplast Development in Arabidopsis thaliana.

Front. Plant Sci. 7, 360.

16. Wilde, A. et al. (1995) Inactivation of a Synechocystis sp strain PCC 6803 gene with homology to conserved chloroplast open reading frame 184 increases the photosystem II-to-photosystem I ratio. Plant Cell 7, 649–658.

17. Wilde, A., Lunser, K., Ossenbuhl, F., Nickelsen, J.

and Borner, T. (2001) Characterization of the

(20)

cyanobacterial ycf37: mutation decreases the photosystem I content. Biochem. J 357, 211–216.

18. Wittenberg, G. et al. (2017) Identification and characterization of a stable intermediate in photosystem I assembly in tobacco. Plant J. 90, 478–490.

19. Wostrikoff, K., Girard-Bascou, J., Wollman, F.A. and Choquet, Y. (2004) Biogenesis of PSI involves a cascade of translational autoregulation in the chloroplast of Chlamydomonas. EMBO J. 23, 2696–

2705.

Recent Progress in the understanding of the molecular mechanism for the photosystem I complex assembly

Sreedhar Nellaepalli, Shin-ichiro Ozawa, and Yuichiro Takahashi*

Research Institute for Interdisciplinary Science, Okayama University .

(21)

Editor: ›Ɔ ¨ëǟȁ”Ǣêó„űÕƈóƀƏƈȂ

Čĵ êЀ ĎǕՌׄÚՑǓׅ 158

DŽNJ ̴ñ΂ĦΞΛ֣ɤդG`cğƇʊϮЕI˜Ȝժ ̠ξ Вׄ̒šǓׅ 159

DŽNJ ̰ΩG'.c׌сѽµфѠ±ž°CIՈìǬϒδӊ̕

ɇυ˓ͭ¼̊ɵȹՃҩׄ͞ÚՑǓǚҏĞМǓׅ 171

DŽNJ y“¤‰v‚±v~¬´'`N”l€°¶œ‰šn“†l™´wgν$<

÷ΩѹЋIŖβеָԶŠ ՎǧȢξͭøׄ̒ÚǓׅ 180

DŽ

DŽNJ NJū ūDz Dz

®ÎģƀƏ/ķ!ŦşE<&@"ķĨƺ„ķžǀ

(22)

Čĵ

›Ɔ¨ëǟȁ”Ǣêó„űÕƈóƀƏƈȂ

$@Iˮã[˜<FʗҲ֑˜)ЄDZG˜4$ϪӃD֧ԖϖFϒȜg[<a6ԩȺCJ͆Àã

~¶vo´|¶gν$<ԎÂľՈìǬՒľӊ̕)̹FλΩIy“¥ɻǁ]mRNAϒδɻǁIŻɠ ǍϊՈìǬƈǺgƀѿD4Ƽ϶ϮЕIXFa7ŤDZ]ԣDZD$?<ɫνGԩ$Ļ՘G'$B[֑

˜g[<a4<W<TALEN]CRISPR/Cas9GãҶ2dcy“¥ѕօʗҲIϒȜJy“¥¼CI èɿIՈìǬIˌǍgƀѿD4δãλΩDZI̹ϠDԺ͌I€™¶Žg·Ǎ28B$c2aGJv

®lq֋Ǭ֣ɤդgν$<ūвǬӊ̕G`c̸Էӊ̕ʗҲJX ѓц˴̸Էӊ̕CJƨ։F†´–v ԊҿƇô̸Էӊ˦IƀѿɲgǓ*,ȿ/cDD[G̸Էӊ̕:I[II˘Șg½/cDѮ&adc ğƇʊϮЕIĻ՘G'$B[:%4<ʗҲ֑˜IɴɷJǓ*,0dWC̊ӞC[ӑÐD4Bтß 4B*<2aGÞɜIϮЕĻ՘IϒȜI<ZGJʋϮЕѯJ˜4$ʗҲ֑˜QIʪʌDD[

Gժϒ2dc˜ʗҲQIj´‹gȶGɏ?B',ɨӂ)#c

:0C̊ΪօӑÐCJğƇʊϮЕG˜4$Δͱg[<a6˜ʗҲµ˜ϪӃ D֠4B̹F Ļ՘CͯϒGϮЕ2dB$c3w°¶œGӛ֠ʾüg'֤$4<W7̒šǓDZI̠ξВŬNJGJ 1҈ԗCժϒ2d<̴ñ΂ĦΞΛ֣ɤդgν$<˿˜Iʊ̘g1тß֖$<:0CJ€‹¶Ն ЈFEIˆ®zlŽ҅†´–vԊI‡l¤v€gϟʹӈȉC*cƀѿɲIXFa7ğƇʊ҅†´

–vԊIūĻǬĻğG`bPSIƐԤIo’°u¶ЈŗI‡l¤v€ׄ˄a+ׅgӲӪC*c0D) Ϸ2d<W<ǚҏĞМǓIՃҩ͞ŬNJIw°¶œ(aJфѠµсѽ±ž°CIՈìǬϒδӊ̕

Iʒͪg̶Ӥ֖$<ՃҩŬNJIw°¶œJέ҈Gժϒ2d<ԧԶфѠĻֈʗҲgν$B0dW CG̰ΩI̶ˢˮӎIфѠΪϊɲg˦a(G2dB*<̊ӑÐCJ̡ΫфѠDͶ֐ΫфѠIğƇ ʊG'.c͂ѿĻŠFEğƇʊϮЕѯGֲ̝X]6$ӛ֠(aǤW?B˿˜I 1 сѽ®´€v

¯œ¶¥ʗҲWCӊӤ$<=$<:4B˿ɜG̒ÚǓDZIȢξͭøŬNJIw°¶œG÷Ωѹ ЋIָԶŠDŖβŠgƀѿD6c2@IʗҲy“¤‰v‚±‰v~¬´ׄGSׅD”l€°¶œ‰

šn“†l™´wׄHPׅG@$B:IàфXDδΫg1тß֖$<˿˜IѹЋʗҲIѻ˲G#c Ï΃§Œ°IеȅFζӪGַ(2dcDD[GGSG'.c̰ΩIθdžɫУI§Œ°Š]HPG'.

cӎ΃֘Ϟµ΃ǺʗҲFEG'$BJğƇʊϮЕѯDIz®¡±¶~¬´Iƀѿɲgʂ5<

ä¼ĻǬсѽфѠ:4BćôI±ž°C:IΪɲgʡĹ4δӷIŲƦgζӊ6c<Z I˜4$ʗҲD:0(aɠadcϪӃgтß4<δ͑ռCJğƇʊϮЕDIխĀJ˦ϲCF$[

I[#c([4dF$)$7d[˜4$ΔͱgλYƀѿɲ)#cDѮ&B$c̊ΪօӑÐ)*?

(.DF?BϊĻ՘լIØͱ]˜4$Ļ՘QIʪʌG@F)dKȻκC#c

˿ɜG̊ΪօIѕօG#<?BJ'ɪ4$ÃƺПѯ̢ӥѯI˝GǓǍ'ÀӛGF?<

ΪGՃҩw°¶œȢξw°¶œIҢѯIϗ̹GJϊĻ՘IևӞ(aIЗΟIý֟G[խfa7 ɭ,ӑÐIƺПgɌ*ż.B֖$<0IǂgċbBĤBIϗ̹Gͻ,ɡϸπ4¼/c

ӊӤΪօğƇʊϮЕG˜4$Δͱg[<a6˜ʗҲµ˜ϪӃ

1ԸчĞE-mail: [email protected]

DŽNJūDz

(23)

Ŏ

Ŏ   Ť Ť° °ũ ũŨ Ũǻ ǻę ęǧ ǧ/ /? ?B B® ®Î Îģ ģƀ ƀƏ Ə0 0ķ ķý ýǪ Ǫ

ŅÃêóêóǯ ůóƀƏƈ ʼnų Ǝ*

ĤǪŹ!*&Ŏ Ť°ũŨǻęǧ/?AĘ@C&Ŀǒ0ģň/)*ãÓ"B80 K+1®Â óƙ IȁPSIȂ®Âóƙ IIȁPSIIȂ0Ʒ®ZrSc‘)0m…S/´Ǵ"B0,E¸Ų!

*S}vdyeZ0®Îģ]ƒlSnjÍģ´0ƜƬ²´ćE®óǻęǧ/?AÌǂÂ"B,/µ

;*ģ½!&Za…cǛƉºė/BÔǑHƒae PSIPSII 0-'@,°üÝ"B0Eō Lj!&8&PSI0Ʒ®Èŭǿ.B80 K/*Synechocystis sp. PCC6803@ÅǴ!&

ņ0PSI‡Ǧ¡@0Ʒ®ZrSc0ĻǫçÂEťõ"B,/ģ½!&Ʒ®ĒĎm…SŸ

Ƨ0İ@ǃť CPSI0Hƒaeƙ0^Jev`S.©ǷEĹ@/!&

1. JJ55ZZGGההğğƇƇʊʊĻĻ՘՘GG''..cc̴̴ññ΂΂CCII֣֣ ɤ

ɤդդӈӈȉȉIÍ́ννɲɲ

§̊ӑÐCJПѯa)ժϒgԺZB*<̴ñ

΂֣ɤդgłν4Bұ?B*<ğƇʊϮЕI˿

ԩIʊ̘G@$Bтß6cäʼn(ağDZ֣ɤ դgν$<ҡёôīՍIw®̸ԷIӈ΃1, 2)] θdžǍŠGí%w®|lIǍŠIӈ΃3, 4)F E)ǁƏ2dB*<0%4<ϪӃJˆ®zl

Ž҅I̸Է҈ô)Ǎŗ4:dGɌ*7adBˆ

®zlŽ҅GƷZԦWd<†´–vԊԎҿƇô I̸Է[Į̸Ъ2dcD$%‡l¤‰vFʽ ĖIɨӂɲgϷƚ4B$c0%4<ǁƏäʼnG [û&K€‹¶ՆЈG'$BJƐԤj´‹

зC#c LHCIIׄLight-Harvesting Complex of Photosystem IIׅIˆ®zlŽ҅īIЈŗ)ɾǺ2 dB$<bğձȀgż.<ğŠDZз IIׄPSIIׅ Jw®£¶´GԽKdBĆɢgż.cDѮ

&adB$<bDˆ®zlŽ҅īՍI†´–v ԊԎҿƇôI‡l¤‰vFǯƱ̹ɋJȿ,ӟ ӱ2dB*<4(4‡l¤‰vFǯƱ̹

ɋD$%[IIǻʃJEI`%F[I(Dƛ

ӊӤΪօğƇʊϮЕG˜4$Δͱg[<a6˜ʗҲµ˜Ϫ Ӄ

*ԸчĞE-mail: [email protected]

fd<ˮJ?*bD4<У)#cӕCJF$I )δΫCJF$=e%(ğDZ֣ɤդG`cӈ΃

Jˆ®zlŽ҅I‡l¤v€gӊ˦6c<Z Iϟʂϖɻǁgʾü6c[IC#b¼ӑIƛ֠

GȌ6c—´gʾü6cD̆ɚ2dc ·˝Cv²²šk°I҈ȁҮğg̱Ĺ6cҮ ğ֣ɤդIǂƇC[ğŠDZз Iׄphotosystem I:PSIׅD PSII gţŁ4B:d;dIсѽīĻȲ gƀӅŠ6c0DJ$,@(I¯‰t¶Fʒ

ͪ5, 6)gν$F.dKƨ։C#?<ǽ΂CJPSI

IҮğźβJ֏ȶGñ,W< PSII I™¶vD Ļֈ4B$F$0D)ƨ։IÅӂFŲƦC#c ʋ)ժϒ4<̴ñ΂֣ɤդCJ͹ôИоFE IijǨgν$<ijŰG`b80 KЊɃI΂ɃCI

֣ɤդӈȉ)ƀѿC#c7)`,Ϫadc`%G 80 KCJǑ,IğƇʊλΩCPSIDPSIIJ˦ϲ GĻֈ4<Үğ€Ÿv°gϷ60D(aՅɑ FĻğҺѣ)#dKPSIDPSIIgţŁ4Bӈ΃

6c0D)ƀѿDFcLHCII)PSIDPSIII լgɘ̑4B$cDѮ&adB$c€‹¶Ն ЈFEIδӷgӊ̕6cǂƇGPSIDPSIIIˆ

®zlŽ҅īIȗƱgƀӅŠC*c0DJ֏ȶ ǴŖDFc0IӑÐCJПѯa)ұ?B*

DŽNJ

(24)

<€‹¶ՆЈG'.c LHCII IсѽīĻȲI ƀӅŠIϮЕ8)gтß6c

[%·@I̴ñ΂֣ɤդIɫνûD4BğƇ ʊ†´–vԊgȌӷD4<ū·ĻǬĻğIϮЕ7,

9)gтß6cū·ĻǬ(aIҮğĄƅg̱Ĺ6 c0DJǑ˕IĻǬ(aIĄƅg̱Ĺ6cǂƇ GȹƴŠG`bց2dB4W%ĻǬ͕I•®@

*gӈ΃6c0DgƀѿG6cū·ĻǬ̱ĹG

`bɠadc[%·@I՗ӂFɻǁJĻǬIˮ լGýǯ4<˄a+Gխ6c[IC#cū·I ğƇʊ†´–vԊҿƇô(aIҮğ€Ÿv°

IхˮǍŠgԮԓ6cDǽ΂C[̴ñ΂C[֞

ўG˄a+)ӈ΃2dc0D)ǁƏ2dB$c

9-13)˄a+I̱ĹJğƇʊ†´–vԊI‡l

¤‰vFǯƱ̹ɋg˦a(G6c0DGџ) c

2. ̴̴ññ΂΂֣֣ɤɤդդIIժժϒϒ

֣ɤդIЖլĻӊѿJȌΩ±´IժŽ˕

ׄNumerical Aperture: NAƪ׌ŵΠׅC͢Wc

̾˝ƊGJͫէIŧĻg NA CŎ?<ЊɃI|

lğԜ˝ƊGJͫէgNAIÑÉCŎ?<Њ ɃI|l)ĻӊC*c'`:IճτDFc ğDZ֣ɤդG'$Bָ$NAgɠcGJ׌ׅȌ Ω±´D|´œ°DIԒֈgϫ,6cW<J

׍ׅ|´œ°DȌΩ±´IլIЖլgָ$șʛ βIΩԊׄͦFEI͹ôׅCΆ<60D)ɨӂD Fc׍ׅgǻδ6c<Zǽ΂CJșʛβIָ

$͹ôCȌΩ±´D|´œ°IլIЖլgΆ

<6͹ͷ±´)ʷν2dc)̴ñ΂CJÞI D0e0I˝ɋIǁƏJF$׌ׅgǻδ6c˝

ͪD4B͹ô¯m¥FEIijǨIF(G|´

œ°D·ђGΆ<4B4W%˝ͪ)#cΊүϭ җJΤ҆ɏĀ˕)֏ȶGȔ2$ΩԊC#b̴

ñ΂G4B[źќ]Ǎɒ)ԍ0aF$<Z̴ñ

΂CIúνGѰ&c±´gǻδ6c0D)C

*c14)>FXG͹ô¯m¥IșʛβJ1.025 ЊɃCЖ͚DǓȮF,͹ͷȌΩ±´D4BI Ŗ̘JTDhEɠadF$0I˝ɋIƛ֠ΛJ ҐźȮIҽ͋)ƨ։C#c0DG#cșʛβI

ͫէýǯɲ(aλ5cҐźȮIҽ͋gұ%<Z GJҿ˕I±´gфXƇf8<фК±´g

ν$c0D)́ŖDFc)ҿ˕±´gƫǺ6 cդТI՚ȝ)̴ñ΂C͍h=b±´gդТ GƫǺ6cʹϦŋ[̴ñ΂GѰ&adF(?<

b6c<Z”¶Ž°J(Fbָ$1̗IΊү ϭҗҾI±´CҐźȮgʙ&c˝ɋD4B'

̯Ƶg4<ΊүϭҗIīՍŹȐgłν4<ŹȐ ƵI±´)ǁƏ2dB$c15)0IǂƇGJ șʛG`cҐźȮ)(Fbʙ&adB'b́Ŗ ɲ)Ϸ2dB$c

ʋJҿ˕±´(aFcȌΩ±´g˛Τ ϤЖ̺GӔѣ6c˝ɋgʷν4<֣ɤդgժϒ 4B*<7)ƪ1Gժϒ4<̴ñ΂֣ɤդI̶ψ ƪ'`NǻտIҺѣıϤgϷ6|´œ° °‡

DȌΩ±´լIЖլJָϤЖGă<dc<Z Τìȓ)ʙ&adȌΩ±´҈ôJijŰ2dF

$0I<ZҐźȮҽ͋'`NĖ֐΅˻źȮҽ

͋)˞2d<Plan ApoƵIȌΩ±´gʷν6 c0D)ƀѿDF?<|´œ°Ί͹J׍̗Iϭ җs®€CʬWd<ЖլGȅթ2dcICϤЖ G2a2dcf.CJF$0I˝ɋIʷνG`

bժϒ4<ɑˮGJ̴ñ΂֣ɤդD4BÀτ˿

ָI NA0.9 gǻδ4ЖլĻӊѿD4B̾˝Ɗ

й350 nmğԜ˝Ɗй1.2 µmgՁʊ4<ǻտ IȌΩ±´JָϤЖÃGѣ(d<տGīՍG Ж͚)͏aF$`%GȱӼƗׄM Plan Apo HR×100NA0.9MitutoyoׅGЖ͚ǭ)ժ.ad

<ΪͬI[Igúν4B$c¡¶°H5ׄBSׅ gƥԛ28c0DCȌΩ±´gƫǺ6cɁ̔

)¼½Gŗ$BΞΛIгӧ˖gұ%îќƀѿF

ž²¶G`bָϤЖgă?<WWɁ̔I¼½ ŗ)ƀѿDF?B$c|´œ° °‡JΤ҆ɏ Ā˕)TU 0 C#cϭҗҾI€‹¶GƫǺ6 c0DC΂ɃǍŠG`cޝšgʠ2&B$

c|´œ° °‡Dv®lq€†‰Iz¶°

Ž‰ŽׄCHׅJΤìȓ)ָ,̞Ԛɲ[#c

՞¦‰~ªCџ$C$c0I0DG`bϟѓ ȓ ģ ͂ g ß 4 < | ´ œ ° ð ѣ I г ŗ ) ƀ ѿ D F?B'bָ$ˊ÷ɲgǻδ4B$c Ɂ̔ÃǗIЙ(a±¶}¶)ȓģ2dȌΩ±

´G`b|´œ°¼Gօğ2dcs°•“¤

®¶ȌG`b±¶}¶ðѣIԌ̢gұ%|´œ

°(aIҮğJƈ5ȌΩ±´G`bօZad

(25)

̱ĹзGԽKdcĻğƣGцƇ4<CCDr¦

®G`bҮğ€Ÿv°)Żɠ2dcĻğƣI ģbŽ)™´ ¶°GF?B$c0DCĦΞΛ

֣ɤդD4B÷ŗ6cW<|´œ°IԳԿĖ J֣ɤդ¼ՍGӔѣ4<LED(aIğgν$B CMOS r¦®G`bŻɠC*c`%GF?B$

c

3. ссѽѽīīğğƇƇʊʊ††´´––vvԊԊIIččƱƱggƀƀӅӅŠŠ ȇ ǔ ƻ Ʋ C ƻ ֭ 4 < v ® ¤ Ž §  € с ѽ g

•‰šiׄpH 7.520 mM tricine50 mM NaClׅ GʉΘ4<ɜ̴ñ΂֣ɤդI|´œ° °‡Q

‚‰4<0I͑ռCJ|´œ°΂Ƀg23°C Gӧ˖6c15ĻI˶֙ɫɜGPSIğׄ710 nm 170 µmol photons m−2s−1ׅgΠȐ6c0DC€

‹¶׌QӠȓ6cğΠȐժǤɜ15ĻCijŰ gժǤ6c€‹¶ՆЈӠȓğIΠȐJ|´

œ°΂Ƀ)80 KGՁ6cWCIй30Ļэʨ4<

€‹¶׍ӠȓIտGJPSIIğׄ470 nm120 µmol photons m−2s−1ׅgν$Bƈ̹Iˊ÷gұ%

PSIğPSIIğJƪ׌ILED(aIğg•´

Ž–€šk°†GԵ4Bɠ<֣ɤդC΃Ǻ2d

<80 KIсѽIҮğ€Ÿv°GJ#cЊɃI

•®@*)Ӄad<)Ǒ˕IсѽCȹƴ6cD

ԵȶIҮğğɃӎG`c΃ǺDTUƈ5€Ÿv

°)ɠad<8)

ƪ2G80 KCӈ΃2d<v®¤Ž§€с ѽIĦΞΛҮğ֣ɤդĖgϷ6ŔԍͫէJ445 nmC#cƆ™v‚°Cƪ׍MN GϷ4<`

%FҮğ€Ÿv°)Żɠ2dcʋJĤB I™v‚°IҮğ€Ÿv°gא@Ism€խ

˕IƔCšk‰‹k´w4<šk‰‹k´wI О·͑ռCJ10 ćЊɃIсѽI€Ÿv°g ĤB%W,Įδ6c`%F 5 @Ism€խ˕I

–®¦¶†g͞Zcşʦϖׄglobalׅšk‰‹k

´wgұ%О·͑ռIšk‰‹k´wCJƆ sm€խ˕I™¶vðѣDȷJϊFcсѽCƈ 5ČgŻc)ʮȷJϊFcD$%̏љ̐çI

½Cұ%0IО·͑ռIşʦϖšk‰‹k´w Iц̘5@Ism€ʊĻI™¶vðѣDȷ)͞

Zadcƪ2 MN(aĻ(c`%G€‹¶

G`bҕȸ™¶vͫէGՂ$JӃad<)5

@IʊĻJϫͫէ(a֙G680 nm686 nm692 nm712 nm730 nmâԩG™¶vgʨ@˿[

ϫͫէI680 nmʊĻׄёGǃb@P2d<™¶

vׅgÅGLHC(aIҮğDȵȝ4<686 nm 692 nmI2ʊĻׄ֍Gǃb@P2d<™¶vׅ

JеҾ2d<PSIII77 KҮğ€Ÿv°DI ƪ

ƪ1. ̴̴ññ΂΂֣֣ɤɤդդI̶ψψƪƪׄׄȫȫׅׅDǻտտIҺѣѣııϤϤׄׄƄƄׅׅ

MO: ȌΩ±´PZ: ™o…jvˆªo¶†LM: ϟѓȓģ͂CH: v®lq€†‰z¶°Ž‰Ž

QS: ΊүϭҗҾ|´œ°€‹¶BS: ΞΛгӧ˖ν¡¶°H5RM/HM: ŹȐդ/ŧԳԿդƄIıϤƄ

½J|´œ°ǽȫƪJ˙ΰ7`bˌǍ

(26)

֥ï(a PSIIIҮğDȵȝ4<էͫէI׍@

IʊĻׄԋGǃb@P2d<™¶vׅgPSIDȵ ȝ4<ОÑ͑ռCJ͢Ǻ2d<sm€ʊĻI

™¶vðѣȷgƫǺ4BʮȷIXg𝶖

® ¦ ¶ † D 4 B Ĥ ™ v ‚ ° I € Ÿ v  ° I

šk‰‹k´wgұ%0%4BLHCPSII PSII3@IʊĻIҮğɐɃmap)ɠadc ƪ2 A"CE"GI"KJ0I`%G4Bɠ ad<ƆʊĻIҮğɐɃIсѽī map C#c

˿¼͑G€‹¶׌QӠȓ2d<сѽ¼(a׍

͑ϞG€‹¶׍QӠȓ2d<сѽIĖgϷ4

<¼(a3͑ϞGJ€‹¶׍QӠȓ4<) PSIҮğɐɃILJɐ)áIсѽG͖RBťĻCJ F(?<сѽIĖgϷ600CJ0I`%F сѽgՆЈ¾ťĻFсѽ DƒP0DG6c Ǒ,Iсѽgӈȉ6cD·ǺIŎƇCՆЈ¾ť ĻFсѽ)Ӄad<·ωƄIDHLJ:d

;dIсѽIPSII/PSI ͖βmapC#bPSIIҮ ğĖgPSIҮğĖCŎbЧ4Bɠ<€‹¶׌

GӠȓ2d<сѽIûC֣ҢC#c)LHC '

`N PSIIIҮğĖCJсѽīGс($вǬΫI

̸Է)Ӄadc0daIвǬΫ̸ԷJPSII ԎҿƇô)͖ԞϖǑ,ǯƱ6c PSII-rich ֜ƹC

#cDѮ&adc0I`%Fɤс̸Է)Ӄad F$сѽ[#?<)ʋI֣ɤդCJğԜ˝Ɗ IЖլĻӊѿ)ñ$0D(a':a,JсѽI Ɗ*G`?BJɤс̸Է)Ӄ&G,$D$%Ð ɻ)#cDɰfdcˆ®zlŽ҅īG'.c PSIIDPSIIčƱJPSII/PSI͖βmapC`b֣

ҢGӃadcv®¤Ž§€CJָР̰ΩI

`%G PSII)ȗƱ6cw®̸Է)ϒՁ4B$

F$0D)Ϫadc)0daI΃ǺŒ¶†)Ϸ 6`%G#cЊɃPSIPSIIJϊFcȗƱgϷ 6`%C#c0I`%Gժϒ4<̴ñ΂֣ɤ դ)v®¤Ž§€G'.cPSID PSIIIč ƱgƀӅŠ6cIGťĻFЖլĻӊѿgʨ@0 D)Ϸ2d<0I0DJä½Iӊ̕gԺZc

¼CIʼnʾDFc

€‹¶ՆЈG'$BEIЊɃLHC)ÁğŠ DZзլgЈŗ4B$cI(g˦a(G6c<Z GJ€‹¶׌׍:d;dGӠȓ4<сѽC LHC)PSIPSIIDEIЊɃƈ5ǂʐGǯƱ6c (ׄĦȗƱׅg˦a(G6cɨӂ)#c0I0 DgǺՙϖGӘþ6c<Z·@IсѽG'.c Ɔ™v‚°CILHCʊĻIϠȌɐɃgњԜG :I™v‚°G'.c PSII/PSI Čg̾ԜG4<

Ϡխƪg÷ʊ4<ׄƪ3ׅ0daIϠխƪÃI

·@IŒ¶†ΛJ:IсѽIҮğĖI#c™v

‚°G'.cČgҶ4B'bƄGðѣ6cŒ¶

†ΛJ͖Ԟϖ PSIIIǑ$сѽīIՍðGȫG ðѣ6cŒ¶†ΛJPSIIǑ$ՍðGϠɑ6c ƪ

ƪ2. €€‹‹¶¶׌׌ׄׄA""Dׅׅ€€‹‹¶¶׍׍ׄׄE""Hׅׅ G

Ӡȓȓ22dd<<v®¤¤ŽŽ§§€€ссѽѽII80 KGG''..cc

̴

̴ññ΂΂֣֣ɤɤդդσσĖĖ

I"LJ€‹¶ՆЈ)ťĻGԍ0aF(?<сѽ IҮğĖׄAEIׅׄBFJׅׄCG KׅJ:d;dLHCPSIIPSIGȵȝ2d<Ү ğʊĻCĮ̸ʊ4<ĖgϷ6ׄDHLׅJ PSIIIĖgPSIIĖCŎ?BɠadcRatio map

•¶J2 µmgҶ6MNJA"D'`NE"H GϷ4<сѽI#c™v‚°G'.cҮğ€Ÿv

°ׄΛׅDא@Ism€խ˕IƔC¿&adc

šk‰‹k´w˻ѓׄԋǻѓׅƆsm€ʊĻg ǃb@P4<™¶vCϷ4<˙ΰ8`bˌǍ

(27)

ƪ׎IϠխƪCƄѸ¼)bIމĻȲJPSII IǑ$ՍðCLHCҮğ)ɐ$0DgϷ4Ƅ½ )bIމĻȲJPSI Үğ)ɐ$ՍðCLHC Үğ[ɐ$D$%ĔƊgϷƚ6cūмGѮ&d K€‹¶׍CLHCI·Ս)PSIďGЈŗ6 cIC#dKϠխƪIĔ*)Ȕ2,FcĔƊ) ӃadcJ7=DÏɾ4<

ƪ3GϷ2d<ϠխƪgӃcD€‹¶׌Ӡ ȓсѽCJϲ(GƄѸ¼)bIĔ*gϷ6ĔƊ )ɠad<)€‹¶׍ӠȓсѽG'$BJ¼ CԬR<ÏɾԵbGĔ*)Ȕ2,FcD$%Ĕ ƊJӃadF(?<·˝CPSII/PSI ͖)€

‹¶׌׍GӠȓ2d<сѽIČIÃլЊɃI ČgϷ6ՆЈ¾ťĻсѽG'$BϠխƪIĔ*

)́ɿGȔ2,FcD$%ĔƊ)Ӄad<`b ǺՙϖFӘþg6c<Zƪ3IϠխƪG'$B

™j„´IϠխĀ˕RgӎЧ4<RJ1(a1 WCIČgŻc)͋IǓ*F R ČJϠխƪG Ϸ4<Ǎ˕IլG͋IϠխ)#c0DgϷ4Ӹ I R ČJӸIϠխ)#c0DgϷ6@Wb

# c с ѽ I R Č ) Ǔ * . d K : I с ѽ J PSII-rich֜ƹCLHCҮğ)ɐ$D$%ĔƊgϷ 4<0DGFcƪ׏Gӈ΃4<6RBIсѽ IR Čg:IсѽIPSII/PSI ͖ׄсѽÃIĤ

™v‚°CȹƴׅGȌ4Bœ²‰4<ƪ׏C JƄGðѣ6cŒ¶†ΛJ€‹¶׌Iсѽ ȫGðѣ6cŒ¶†ΛJ€‹¶׍IсѽC#

cԋҐI»ӉJ710 nmIğgΠȐ4B€‹¶

׌GӠȓ4<сѽIŒ¶†C#b֍ҐIÄJ 460 nmIğΠȐG`b€‹¶׍QӠȓ4<с ѽC#c€‹¶ՆЈӠȓg4<G[խfa7 ՆЈ)¾ťĻC#?<ՆЈ¾ťĻсѽׄͺ$Œ¶

†Λׅg·@Iw°¶œD4:däǐIсѽg

€‹¶׌QӠȓ2d<сѽ€‹¶׍QӠȓ 2d<сѽIw°¶œGĻ֥4<€‹¶׍

w°¶œC€‹¶׌w°¶œD͖Ԟ4B R ČJҕȸȔ2,F?B$c)Ǔ*FǍŗCJF (?<:I·˝CՆЈ¾ťĻw°¶œIсѽ CJRČ)́ɿGȔ2,F?<сѽ)$,@(

Ӄad<0daIц̘gӊՕ6c§Œ°gƪ׏

ÃGϷ4B$c

€‹¶׌сѽCָ$ R Č)Ӄad<IJ PSIIDLHCIĦȗƱG`c[IDӊՕ4<ׄƪ 4Ƅ¼ׅ€‹¶׍сѽCJPSI(aIҮğ ɐɃ)ɐ,F?B$c<ZLHCI·ՍJPSID цƇ4Bo’°u¶güш4B$cJ7C#c G[((fa7€‹¶׍сѽCָ$ R Č)Ӄ ad<0DgӤ˦6c<ZPSIIƐԤG PSIID

ƪ

ƪ3. ƆƆ™™v‚°°GG''..ccLHC Ү

ҮğğIϠȌȌɐɐɃɃgg::I™v‚°° G

G''..ccPSII/PSIČČGȌ44Bœ

²

²‰‰44<Ϡխխƪƪ

€‹¶׌ׄȫׅ'`N€‹¶

׍ׄÃǗׅGӠȓ2d<сѽ ƄJՆЈ)¾ťĻ=?<сѽ Ɔw°¶œ4 @IсѽIц̘g Ϸ4<˙ΰ8`bˌǍ

参照

関連したドキュメント

In this chapter, we shall introduce light affine phase semantics, which is meant to be a sound and complete semantics for ILAL, and show the finite model property for ILAL.. As

In this paper, Zipf’s law, allometric scaling, and fractal relations will be integrated into the same framework based on hierarchy of cities, and, then, a model of playing cards will

В работе [2] доказана финитная независимость отно- шений вложимости и эпиморфности на любом нетривиальном конгруэнц-дист- рибутивном

Here we shall supply proofs for the estimates of some relevant arithmetic functions that are well-known in the number field case but not necessarily so in our function field case..

(4S) Package ID Vendor ID and packing list number (K) Transit ID Customer's purchase order number (P) Customer Prod ID Customer Part Number. (1P)

I have been visiting The Nippon Foundation, Kashiwa Company, Japan Aerospace Exploration Agency (JAXA), Ariake Water Reclamation Center, Tokyo University of Marine Science and

Estos requisitos difieren de los criterios de clasificación y de la información sobre peligros exigida para las hojas de datos de seguridad y para las etiquetas de manipulación

日髙真吾 企画課長 日髙真吾 園田直子 企画課長 鈴木 紀 丹羽典生 樫永真佐夫 樫永真佐夫 樫永真佐夫 川瀬 慈 齋藤玲子 樫永真佐夫 三島禎子 山中由里子 川瀬