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 oq®IϒՁDҡёôQIĻŠԿЊG'.cs®vҀԊIɖŎ
ҩÕ Ͻ áׄ̒ÚǓ áׅ 139
DŽNJ ğŠDZзIҿƇôIĻǬօƇ̸͂Iӊ˦IԺȜ
±¯µ¯¶¶ áׄȡȟǓׅ 148
DŽNJūDz ğƇʊϮЕG˜4$Δͱg[<a6˜ʗҲµ˜ϪӃ 157
Čĵ êЀ ĎǕՌׄÚՑǓׅ 158
DŽNJ ̴ñĦΞΛ֣ɤդG`cğƇʊϮЕI˜Ȝժ ̠ξ Вׄ̒šǓׅ 159 DŽNJ ̰ΩG'.cсѽµфѠ±°CIՈìǬϒδӊ̕
ɇυ˓ͭ áׄÚՑǓáׅ 171
DŽNJ y¤v±v~¬´'`Nl°¶nl´wgν$<
÷ΩѹЋIŖβеָԶŠ ՎǧȢξͭøׄ̒ÚǓׅ 180
ƻƚ0Ɲ Ū̴ȢƹGλѹ6cğƇʊλΩIğձȀIͫէýǯΪɲ
Ȕ̌ϤԀǬ áׄÃǗǓ áׅ 194
ū·Ŵ О ƥĆұ¡´ ԩҩ ɦ̒šǓ 195
ãÓLJ The 1st Asia-Oceania International Congress on PhotosynthesisŵőǁƏ
ĴŎô įŚľőŗׄƼλϮׅ 202
ãÓLJ О1ƥjjµqjjƬտҕʒIëժĒǁƏ 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
Ć Ćǹ ǹDž Dž
®
®Î Îģ ģó ó Ǩ ǨE Eƞ ƞ B B/ / & &( (* *
ĸŁ®ÎģóǨǿőƽǟ ȁāĀêóŸ´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~gժ 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'֤$4W6
M
M_ _N Np p} }Z Zc c0 0Ź Źǘ ǘ, ,ƴ ƴƤ Ƥ¡ ¡5 50 0´ ´Â ÂǗ ǗƊ Ɗ/ / B B P
P} }S Sc cƮ Ʈnj nj0 0ĕ ĕ» »
Ņêóêóǯ ƣÎĵÂƀƏƈ ċáƈóúı
ĸŁîñêó ůóǠ ŪnjűŪƈóƈ
êǭčƒêó ǿƖIJƪĮǕŒŐ
ƶƅŘų±ñåųđÖų¬ûŇď
ľĥ+ŹƱ!&ƼñŌŪ1ñƴƜƬ²/M_Np}Zc,?2CBƴƤ¡0ºǾ¡Eēģ"BM _Np}Zc²/1ƮnjǤÿEq Z/SnIǫ¡.-EƵƍ!&p}x}sbI ȁPLBȂp_}VJd,°/Źǘ"Bñƴ®EÉB,PLB 0ğǓ.ăæ,_}VJdƯ 0ēģAM_Np}Zc1ƴƤ¡5,´Â"BM_Np}Zc,ƴƤ¡0ƮnjƟģ1*
A-'@<P}ScƮnj+ByiP}ScXYHXT~\ ȁMGDGȂ,YP}S cXYHXT~\ ȁDGDGȂEƿŐģģ´,"BŹƱĻ@®Űä+űƪ!&ŌŪ +1C@0P}ScƮnj®Îģ=_}VJdƯ0ēģƴƤ¡0Źǘ/Ǥƿ.ĕ»Eň&"
,Ĺ@/ C*&ľĥŹƱĻ0M_Np}Zc0ēģ= PLB 0Źǘ/B MGDG = DGDG0ǤƿĠ1Ĺ+(&8&M_Np}Zc@ƴƤ¡/´Â"BDZ/BPLB@_
}VJdƯ50^Jev`S.Ưçį/*P}ScƮnj-0?/ǬDB0<´(*
.(&ÚĤ1MGDG=DGDG0Îģƭ¼ !&XJge[eEŲ*M_Np }ZcŹǘĻ/P}ScƮnjĪĕ»EĹ@/!&8&M_Np}Zc@ƴƤ¡50´Â ǗƊ/BMGDG,DGDG0Ǭ/Ǭ!*<ƀƏEǕ;*AĿķ0Ơň/)*ƨø"B
1. JJ55ZZGG
§ƲÃFEI˶ʐCϒҒ4<ҸǬ̰ΩJёŠ8 7ѼԜ)Ң4,îէ4<$f^c§©~ׄ־
ŠҒλ&ׅIɒʃgDc0ID*Ǭҡсѽī CJҡёôIʼnִôD4Boq®)ϒ Ձ4:IīՍGJʑȹF²®zlDD [G²®¦®¡kׄPLBׅD`KdcӄŇ
͋4$̩ǬΫI̸҅Է)ɒʊ2dc0daI҅
GJv²²k°IƇʊÃլôC#c²v
²²k¯ׄPchlideׅD:dgğýǯϖGv
²²k¯ׄChlideׅQDǍˀ6c Pchlide Ջ ĜՓоׄLPORׅ)ңА6cPchlideIǑ,J҅
¼CLPOR'`NNADPHDğͯɲƵI»ѯҿƇ
*ԸчĞE-mail: [email protected]
ôgɒʊ4ğIΠȐgɚ?B$c 1,2)>FX GҸǬ̰ΩäǐḬΩ]Ҫ֥~jv¯
jIǑ,J˶ʐC[ͯɲǵ6cPchlideՋĜՓ оgʨ@<Z˶ʐCv²²k°g÷dF$I JLPOR4(ʨ<F$ҸǬ̰ΩIΪɥC#c3) ˶ʐCϒҒ4<ҸǬ̰ΩI־ŠǬҡ)ğgż .cDğͯɲƵҿƇôÃI Pchlide Jϟ>G
ChlideGǍˀ2d:Iɜ2aGv²²k°Q
DãӰ2dc4)ǓՙGңА4B$<LPORJԶ ](GĻӊ2dc·˝ğŠDZзҿƇô´v ԊgJ5ZD6cğƇʊխԸ´vԊ)Ґо DD[GƇʊ2dcPLB)ɱԶGȥlj6cDƈ ˮG®zl҅IͯϒF̸Ъ)ԍ0bц̘D 4Boq®JҡёôQDĻŠ6c0I ԿЊJLPORG`cPchlide IğǍˀGő&
c
cm m` `S SZ Z
ğƇʊխԸՈìǬIԛığƇʊ´vԊ]Ґ оIƇʊğƇʊҺѣIɒʊDğƇʊŹɫзIͯ
ɲŠ2aGPLB (a®zl҅QI҅Ǎˀ gƍX0daIԿЊ)֏ȶGl¤v(@
ԧԶGԍ0c<ZğƇʊŹɫзI̸Ъ]ҡёô IϒՁ̸͂gӧRc<ZI§°зD4BϚh GϮЕ2dB*<
PLB D®zl҅JW?<,ϊFc̸Էg 4B$c)E>a[ҀԊÑȞg¶G4<
̸҅ԷC#bҋƑͻ$0DG:daIҀԊф ʊ[֏ȶGïB$cׄƪ1ׅ0daI҅ҀԊI ǓՍĻJ̰Ω]~jv¯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$z¶´ƵI Кô̸Էgʨ@<ZūέCJ®¦®̸Էg÷d 7ãfbGt|{° II D`Kdc֏Ñ
Ȟ̸Էg÷c´®¦®ҀԊC#c:dGȌ4 DGDG JӇ͛ɲ֝ՍDό͛ɲȖՍI®´)
`,~¯´¶ƵIКô̸Էgʨ@<Z͛Ί
ÃCÑȞ̸Էgɒʊ6c®¦®ҀԊC#c
7)ׄƪ2ׅ®zl҅IMGDG/DGDG͖J2 Gԩ$ČgDc0D)ϪadB'b0Iфʊ)
®zl҅Ϊ́I̸ԷGӂC#cƀѿɲ) Ϸƚ2dB$cPLB]²®zlIɒʊG
Ȍ4B[MGDG/DGDG ͖)ӂC#cƀѿɲ
)ӌŷ2dB*<)8):IӝсJǺ(CJF$
0dWCIǍϊôFEgν$<ӊ̕(ağθ dž½Cλѹ4<̰ΩGխ4BJs®vҀԊ) ğƇʊ]®zl҅IɒʊҡёôIϒՁG
ӂFɖŎg̘<60D)˦a(G2dB*<)
˶ʐ½CIoq®Iɒʊ]PLBIϒՁ PchlideIƇʊDLPOR DIҿƇôɒʊG'.c Ҁ Ԋ I ӂ ɲ J ¾ ˦ C # ? < Þ ƥ ʋ J MGDG]DGDGIƇʊѿŏ)ñ½4<~²l
gν$Bs®vҀԊ)˶ʐCʢ%ɖŎ gӧR<IC:Iʊ̘gтß6cW<˶ɒʃ ɒʊɜoq®(aҡёôQĻŠ6cԿ ЊGs®vҀԊ)EI`%Gխ¿6cI(Ϊ GPLB (a®zl҅QIl¤vF҅
ǍˀGEI`%FɖŎg̘<6I(G@$B[
ŻbфhC'b:I˿˜Iц̘[ù8Bтß4
<$
2. ooqq®®ϒϒՁՁGG''..ccMGDGIIɖɖŎŎ ̰ΩI MGDG ƇʊJҐоôş҅GȗƱ6c MGDG ƇʊՓоG`?Bұfdc~²l
GJҐоôīş҅GȗƱ6cMGD1 Dǐ ş҅GȗƱ6cMGD2DMGD3I3ЋIMGDG
ƪ
ƪ1. ®®zzll҅҅DDPLBIIҀҀԊԊффʊʊII͖͖ԞԞ m±´m(aūֈ2d<ҡёôI®zl
҅5)Dz¥u־ŠҒλ&(aūֈ2d<o
q®IPLB6)IҀԊфʊÅӂF҅ҀԊI Xgӑԟ
ƪ
ƪ2. MGDGGG `` cc tt || {{ °°II̸̸ ԷԷ DD DGDGGG``ccÑÑȞȞ̸̸ԷԷ
ό͛ɲȖՍGȌ4̴ɲ֝ՍIôАIŭZcŎƇ )Ȕ2$MGDGJz¶´Ƶg4B'bt|
{°IID`Kdc̴ɲ֝ՍgīďGօƇ4<
̸ԷgDb]6$DGDGJ~¯´¶Ƶg4 B'b͛ΊÃCÑȞ̸Էgɒʊ6c
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#b®zl҅Güш2dcs®v
ҀԊIƇʊIǓŧgʢ?B$cMGD2DMGD3 J¯´ͅÈˮGҐоôǐGüш2dc DGDG IƇʊGÅGӺΰ6cDѮ&ad®zl҅
Is®vҀԊIƇʊQIȄ¿J̴ZBճǺϖ
C#c11,12)
MGD1ՈìǬgǶĤGͅ$<vjmǍ ϊôCJs®vҀԊ)TDhEƇʊ2dF$
<Z®zl҅Iɒʊ)'0a7̰ΩôI λѹG[ͻņFɕ֕)λ5c10)Ғλ&I͑ռC Ҋ͎DFc0DGő&MGD1IvjmǍ ϊJѼɒʊ[ɐ,ձȀ4ǬҡIɒʊ¾ĤgɌ*
ԍ06<Z13)0IǍϊôI־ŠҒλ&gν$<
ӊ̕J¾ƀѿC#?<:0CʋJMGD1 mRNA g¶yD4<ÜȪ£lv² RNA
ׄamiR-MGD1ׅgt|¦ ´ׄDEXׅýǯ ϖGϒδ28cзgν$14)MGD1Iϒδʙń) oq®IϒՁGEI`%Fɕ֕g¿&
cI(gӧR<15)
amiR-MGD1ɒԊԛˀôg˶ʐCϒҒɜ4ˢլ
λѹ28<D0eDEX ĸζ4<ćôׄ+DEXׅ C J MGD1 I mRNA ± ° ) ̈ ĸ ζ ć ô
ׄ−DEXׅI35%GWCʙń2d<:0CҀԊ фʊgĻ̕4<D0e+DEX IDGDGƍՙJ
−DEX DTDhEǍfaF(?<)MGDG I ՙJ−DEXI65%GWCñ½4B$<:I<Z
−DEX C 1.5 ЊɃC#?< MGDG/DGDG ͖) +DEXCJTU1ׄMGDGDDGDG)РՙׅG F?<͆G0IMGDGIȕ)oq®
IɒʃG¿&cɕ֕gӧR<+DEX CJ PLB I̩Ǭ̸ԷJf7(GËd<)PLB Iϒ Ձ:I[IGJǓ*Fɕ֕g¿&F(?<ׄƪ 3ׅ0Iц̘(aPLBIɒʊGJMGDG/DGDG
͖IųȅFz´²¶°JɨӂCΝ$0D)ŀ
˦4<PLBI|lJLPORIңАՙGǓ*
,ȫƄ2dc0D)ϪadB$c2)+DEX|´
°CMGDG)ȕ4B[LPOR IңАG`
bPLBI|lJǍfaF(?<DѮ&adc
·˝²®zlIϒՁJ+DEXćôCǓ*
,ձȀ2d<ׄƪ3ׅ+DEXCJ҅IÅӂҀԊ ʊĻC#c MGDG )ȕ4<G[((fa7 PLB JԵȶI|lGԩ$±°WCϒՁ4<
0D(a:Iϲ®zlIɒʊGú&
cs®vҀԊIՙ)¾ԑ4<I([4dF$
ƪ
ƪ3. amiR-MGD1II־־ŠŠǬǬҡҡGG''.. c
cooqq®®IIɒɒʃʃDDPLBII̸̸ Է
Է
ׄA, BׅDEX ̈ĸζz´²¶°
ׄ−DEXׅDׄC, DׅDEXĸζ4MGD1 gʙń4<|´°ׄ+DEXׅׄBׅ
ׄDׅJ:d;dׄAׅׄCׅIPLB IʥǓƪϩŮJ²®zlg Ϸ6x¶°¶J1 μmׄA, Cׅ D0.2 μmׄB, DׅׄFujii et al., 201715ׅ
g ˌ Ǎ www.plantphysiol.org.
Copyright American Society of Plant Biologists.ׅ
͆GMGDGIȕ)PchlideIƇʊ]ңА ҿƇôɒʊGEI`%Fɕ֕g¿&cI(gӧ R<+DEX I־ŠҒλ&CJPchlide Iՙ)
−DEX Iй 60%GWCȕ4B$<: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 ¦ ° 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
ңА²il°) MgMTIǍϊôD`,ïB
$<0D(aMGDGIȕJMgMT)ӋǨ6
cGǓ*Fɕ֕gŷU4<DѮ&ad c
Pchlide IĻ̕(a+DEX CJğͯɲƵI
Pchlide )ΪGȕ4B$c0D[˦a(DF?
<MGDG IȕG`b Pchlide ) LPOR ]
NADPHD%W,ҿƇôg÷dB$F$ƀѿɲ)
Ѯ&adc2aGğͯɲƵI»ѯҿƇôJ҅
¼CօƇ4Ǒՙôgɒʊ6c0D)ϪadB$
c)+DEXI־ŠҒλ&CJ»ѯҿƇôIǑ ՙôŠ[ձȀ2dB$c0D)Ļ(?<0da Iц̘JMGDG)Pchlide-LPOR-NADPH»ѯ ҿƇôIɒʊ]:IǑՙôŠGӂC#c0D gϷ4B$c
ä¼Iц̘(aMGDGIҕȸIȕJPLB IɒʊGJǓ*Fɕ֕g¿&F$)²®z lIɒʊ]PchlideIƇʊğͯɲƵI»ѯҿ ƇôIɒʊgɐ,ձȀ6c0D)˦a(DF?
<15)
3. ooqq®®ϒϒՁՁGG''..ccDGDGIIɖɖŎŎ MGDGƇʊDƈ̹DGDGIƇʊ[Ґоôş
҅CұfdcUDP-Gal (a MGDG Gs®v
ƪ
ƪ4. ALAĸĸζζɜɜIIamiR-MGD1GG''..ccPchlideƇƇʊʊÃÃլլôôIIңңАА
־ŠҒλ&GALAg24ˮլĸζ4<ɜIPchlideƇʊÃլôIңАՙg−DEXD+DEX|´°C͖Ԟ4
< Pchlide I Ƈ ʊ х Ԕ D Ɔ Ã լ ô I ̸ Է g w ® I ½ G Ϸ 4 < ׄFujii et al., 201715)g ˌ Ǎ www.plantphysiol.org. Copyright American Society of Plant Biologists.ׅ
¶1ĻǬgԛЈ6cDGDGƇʊՓоJ~
²lGJDGD1DDGD2I2Ћ)ǯƱ 6cE>a[ҐоôIǐş҅GȗƱ6c)ё ҐфѠIDGDGIǓŧgƇʊ6cIJDGD1C
#c16,17)DGD2JMGD2]MGD3DĦGĕ*
¯´ͅÈˮIDGDG ƇʊGÅGȄ¿6c 11,16) ğθdž½Cѹ?< DGD1 IvjmǍϊô
ׄdgd1ׅCJDGDG ƍՙJλ̦I 10%ä½ Gȕ4ćô)ϬŠ6cFḚΩôIλѹG[
ɐ,ɕ֕6c)ҡёôIϒՁGJҊƓϖFɕ֕
J¿&7v²²k°ՙ]ğƇʊͯɲI#cЊ ɃIȕ]®zl҅)΅˻6cD$?<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$<ʋIϮЕ(aMGDG IҕȸIȕ)PchlideIƇʊ]ңА»ѯҿƇ ôIɒʊGǓ*Fɕ֕g¿&c0D)Ļ(?<
) 15)DGDG )oq®IϒՁG'$B EI`%FɖŎg̘<6I(G@$BJ̈ӊ˦
C#?<:0CʋJ dgd1 Ǎϊôgν$
DGD1 Iͅ˅)oq®IϒՁGEI`
%Fɕ֕g¿&cI(gӧR<19)־ŠҒλ&G '.cDGD1IȄ¿)Ļ(?B$F(?<<Z W7dgd1I־ŠҒλ&IҀԊфʊgӧR<D0 eDGDG Iƍՙ)λ̦I 20%ЊɃGWCñ
½4B$<0Iц̘(aoq®G'
$B[DGD1)ÅӂFDGDGƇʊՓоC#c0 D)˦a(DF?<dgd1 Ioq®g ӈȉ4<D0ePLBJλ̦DƈЊɃIǓ*2 WCϒՁ4B$<)īՍI̩Ǭ̸ԷJǓȷGË dB$<ׄƪ5ׅW<λ̦Ioq®
JǓʟ 1 @IǓ*FĬΫׄǻտGJεΫׅI PLB4(ʨ<F(?<)dgd1CJ͍FPLB) ҿ˕Xadc0D)Ǒ,PLBIϤĬɃ[Ң4, ñ½4B$<ҿ˕X&cPLB JǶĤG˛Φ Š4B$cI(ļΦǐID0eC@F)?B$
cI(JĻ(aF$)PLBIɒʃGϊȶ)λ5 B$c0DJ˦a(C#c0IPLB Iɒʃϊ ȶJīՍI̩Ǭ̸ԷIËdDխĀ6cI([4 dF$W<dgd1 CJ²®zlIɒʊ [TDhEӃadF(?<)amiR-MGD1 I x¶Dƈ5,s®vҀԊIǓŧ)PLB I ɒʊGúfd<<Z²®zlg÷cҀԊ )¾ԑ4<ƀѿɲ)Ѯ&adc
͆GPchlide IңА]ƇʊхԔIΫʃgӧR
ƪ
ƪ5. dgd1II־־ŠŠǬǬҡҡGG''..ccoo q
q®®IIɒɒʃʃDDPLBII̸̸ԷԷ
ׄA, Bׅλ̦ׄwild typeׅDׄC, Dׅ dgd1ׄBׅׄDׅJ:d;dׄAׅ
ׄCׅIPLB IʥǓƪϩŮJ²
®zlgϷ6x¶°¶J1 μmׄA, CׅD0.2 μmׄB, DׅׄFujii
et al., 201819) g ˌ Ǎ
www.plantphysiol.org. Copyright American Society of Plant Biologists.ׅ
<ц̘amiR-MGD1 IǂƇDƈ̹dgd1 CJ
PchlideIңАՙ)ȕF,ALAIͽőǻֶ(a
Mg-Proto IXI§¦°o°ŠI
)ΪGձȀ2dB$c0D)Ļ(?<ׄƪ 6ׅ MgMT Iʢ%Js®vҀԊIȕI ɕ֕gΪGż.]6$DѮ&adc)ĨôϖG EI`%FձȀ)ԍ0cI(G@$BJÞɜI Ӧ֠C#cW<dgd1CJğͯɲƵIPchlide IŎƇ)ȕ4B$<0d[ amiR-MGD1 I +DEX Dƈ̹C#c)Pchlide IƇʊձȀ[ğ
ͯɲƵIȕ[amiR-MGD1I˝)`b֣ҢC
#?<:d;dIҀԊIȕIЊɃׄ+DEX amiR-MGD1CJMGDG)35%Iȕdgd1C J DGDG ) 80%Iȕׅ[ù8BѮ&cD
Pchlide IƇʊ]ğͯɲƵIңАGȌ6cɕ֕J
MGDGI˝)ɐ$DѮ&adcamiR-MGD1I +DEX CJ»ѯҿƇôIǑՙôŠ)ձȀ2dB
$<):I`%FδӷJ dgd1 CJӃadF (?<·˝amiR-MGD1DϊFbdgd1Ǎϊ ôCJPchlideIğǍˀɜIChlide-LPORҿƇô IʫŗGϊȶ)λ5B$<®~ªΠȐɜ
Chlide-LPOR ҿƇôIǑՙôJÑՙôQDӊֈ
4B$,)dgd1 CJ:IԿЊ)Լ,F?B$
<0daIц̘(aMGDGDDGDGJE>
a[ğͯɲƵ»ѯҿƇôIɒʊGӂC#c) :Iխfb˝JϊFcDѮ&adc
4. ooqq®®((aaҡҡёёôôQQIIĻĻŠŠˮˮGG''.. c
css®®vvҀҀԊԊIIɖɖŎŎ
~²lI־ŠҒλ&CJğΠȐɜ1 ˮլIˮΛC PLB I̩Ǭ̸Է)Ǔ*,Ëd4 ˮլɜGJPLB IƉ͏[TDhEӈȉ2dF, Fc0IտPLBIШΫI̸Է)ϟʹʑȹFҥ
̸҅ԷGǍˀ2d²®zlׄ[4,J
®zl҅Iʼnִôׅ)֣ҢGϒՁ6c0D)˦
a(G2dB$c 20)PLB D®zl҅J֏
ȶGï<ҀԊфʊgʨ>E>a[s®vҀԊ gÅʊĻD6c)ׄƪ1ׅ0Il¤vF
҅ǍˀGs®vҀԊ)EI`%FɖŎg̘<
6 I ( J ` , Ļ ( ? B $ F $ : 0 C ʋ J amiR-MGD1 D dgd1 I־ŠҒλ&IёŠԿЊg ӈȉ6c0DC0IԿЊG'.cs®vҀԊ IӂɲgӧR<ׄ̈ϒҶׅ
ёŠˮIҀԊՙIхˮϖFĻ̕gұ?<D0
ƪ
ƪ6. ALAĸĸζζɜɜIIdgd1GG''..ccPchlideƇƇʊʊÃÃլլôôIIңңАА
־ŠҒλ&GALAg24ˮլĸζ4<ɜIPchlideƇʊÃլôIңАՙgλ̦Ddgd1C͖Ԟ4<Pchlide IƇʊхԔDƆÃլôI̸Էgw®I½GϷ4<ׄFujii et al., 201819)gˌǍwww.plantphysiol.org.
Copyright American Society of Plant Biologists.ׅ
eğΠȐʼn(ağΠȐ12ˮլɜWCJҒλ
&Is®vҀԊIƍՙJλ̦G'$B[T DhELJ&F$0D)Ļ(?<0I0D(a PLB (a®zl҅QIǍˀJs®vҀԊ ILJőgɨӂD4F$0D)Ϸ2d<W<
+DEX amiR-MGD1Ddgd1CJ:d;dMGDG DDGDGIՙ)ёŠԿЊgԵ4BȶGñ$±
°Gʙń2dB$<0daḬΩôG'$B ğΠȐɜIǬҡIҐоôI̸ԷǍŠgǬ֣ɤ դCхˮϖGӈȉ4<D0e+DEX amiR-MGD1 CJPLB IǘJ−DEX Dƈ̹GԶ](Gԍ*
<):IɜI®zl҅ɒʊ)ɐ,ձȀ2d
<:dGȌ4dgd1CJPLBI͏φΩ)F(
F(ǘ87:IɜI®zl҅Iɒʊ[ձ Ȁ2dc0D)Ļ(?<E>aḬΩôG'$
B[LPORI±°JğΠȐɜGԶ](Gȕ 4<·˝v²²k°] LHCII IÅӂ´
vԊC#cLHCB1IңАJɐ,ձȀ2d<
F9amiR-MGD1Ddgd1IլCJPLB(a
®zl҅QIǍˀΪG PLB IǘG'$
BǓ*FՂ$)Ӄad<I=e%(@Iƀ ѿɲD4BDGDG )®¦®ҀԊC#cIGȌ 4MGDG JūέCҀԊÑȞgɒʊC*F$
´®¦®ҀԊC#c0D)ʫ/adcׄƪ2ׅ DEX ĸζ4< amiR-MGD1 I־ŠҒλ&CJ MGDG Iƍՙ)ΪϊϖGȕ4B$c<Z DGDG IŎƇ)Եȶ`b[ָ$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®vҀ ԊƇʊIʙń[ь,ICҡёôɒʊJձȀ2d
<WWGFcDʺ2dc·˝dgd1 Ǎϊô CJDGDGIՙ)֣ҢGȕ4B$c<ZPLB CJMGDGIŎƇ)ϊȶGָ$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 Jǘ4®zl҅
)ϒՁ6cin vitro Iǻֶ(a´®¦®Ҁ
ԊIMGDG[LHCIIDϠÓ÷ν6c0DC®¦
®̸Էg÷dc0D)ϪadB$c21)ğθdž½ Cλѹ4<dgd1Jλ̦DƈЊɃGLHCIIgң АC*c0D(a22)ɝGңА4< LHCII F E)MGDGDϠÓ÷ν6c0DC®¦®ҀԊ C#cDGDG gTDhEͅ$B[˿уϖGJ
®zl҅gɒʊC*cI([4dF$
¼ӑIʺ)E0WC͋4$I(Gő&F9 s®vҀԊIͅ˅G`?Bv²²k°Ƈʊ хԔ]ğƇʊ´vԊIңА)ձȀ2dcI (J¾˦C#cW<ʋJv²²k°I ƇʊGő&̧]Ґоôz¶IğƇʊՈìǬI mRNA ңА[s®vҀԊIƇʊձȀG`?B ʙń2dc0DgӃĹ4B$c)12,14)ׄ̈ϒҶׅ
:d)EI`%F¦r¥Cԍ0cI([˦
a(CF$s®vҀԊGő&®zl҅
GJՔɲҀԊC#c i°w¯²¶
°D° t¡~°j~°w¯²¶°) Åӂ̸ʊʊĻD4BƍWdc)oq®
IϒՁ]ҡёôQIĻŠIԿЊG'.c0da IՔɲҀԊIɖŎ[ӊ˦6R*Ӧ֠D4B͏2 dB$cÞɜ2aGϮЕgԺZ0daIύƛ g˦a(G4B$*<$
Ӱ ӰԢԢ
ˢ̊ǟǬǓDZIȔ̖ɷ2hGJǬ֣ɤդӊ
̕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.
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.
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
®
®Â Âó óƙ ƙ I ƾ ƾΠΡ ¡0 0´ ´ñ ñDz DzÎ ÎŒ ŒŐ Ő0 0DŽ DŽĹ Ĺ0 0Ǖ Ǖý ý
‡
āĀêó Ÿ´ǥâƂƈóƀƏĥ hl~Z~ ^ ûŧŽǟǿőƽǟ*
ǣƛŹűà®Îģǵñǘƙ/*®MhR E¸Ų!*p}ZcXHf<!1XcS
wcEǣÂ!nLdQXEǞ¬"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 Ŀǒ0ƀƏEě/Ɲ"B
1. JJ55ZZGG
§͛gՔŠ4NADP+gՋĜ6cՔоϒλƵğƇ ʊǬìՁзJğo°u¶głν4BϟľG
͂ѿ6c@IğŠDZзG`bִŗ2dcğŠ DZзIIׄPSIIׅJɐ$ՔŠŏgɒʊ4͛gĻӊ4
®t´ׄPQׅgՋĜ6c)ğŠDZзI
ׄPSIׅJ®t´ׄPQׅD~v²¥b6f
ׄCyt b6fׅ(aIǬg®~j´ׄPcׅ [4,J~v²¥cׄCyt cׅgß4Bż.Żb
n±t~´ׄFdׅgՋĜ6cՋĜƵFdJ
n±t~´ NADP ՔŠՋĜՓоׄFNRׅG
ǬgìՁ4NADPHgλʊ6c0I`%Fϟ գƵǬìՁзׄ¯jǬìՁзׅG`bՔо )ϒλ4ΚՔƫǺŹɫGɨӂFՋĜŏC#c
NADPHgλʊ6cW<0IŹɫDĦɖ4B
®zl҅gß4<²´ִŗŏׄpmfׅ) ɒʊ2dATPƇʊՓоIĕ*CATP)Ƈʊ2 dc·˝FdgՋĜ4<ǬJCyt b6fQìՁ 2dĮN PSI gՋĜ6cхԔ)#c0dJ
NADPH gλʊ4F$) ATP ƇʊGɨӂF pmf
gɒʊ4ɣθϖǬìՁзׄ|lv¯vǬ ì Ձ з ׅ D ƒ K d B $ c PSI I Ջ Ĝ ď C J
‡~´¢m¥®zl҅ԎҿƇôI̸ԷDl¤v
*ԸчĞE-mail: [email protected]
NADP+äǐG[Ио]ϱ־gՋĜ6cхԔG
Ǭ)ìՁ2d<bҡёôIՓоIͯɲgՋĜŏ G`bӧЩ6c±vńɡGɨӂFǬg üш4<b6c0I`%GPSIIՋĜďJǑ͂
ѿC#bğƇʊǬìՁзIǬIĻՒG'$
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 ǺѩgƪGWDZ<ŹɫÃɧҿƇôׄPSI RCׅ JŹɫÃɧ|«PsaA DPsaB (a̸
ʊ2dc®zl҅gɐ$τͯɲŋׄñ
CñΙɃISDSׅCƀΊŠ6cDŹɫÃɧׄRCׅ
DŽNJ
ƪ
ƪ1. ²²££ďď((aaӃӃ<<PSIҿҿƇƇôôII||««̸̸ʊʊ
PSI RCהŹɫÃɧ|«PsaADPsaB(a̸ʊ2dcPSIŹɫÃɧҿƇôPSI coreהŹɫÃɧҿƇ
ôGáIPSI|«gцƇ6cPSIzjҿƇô²£ďGPsaC/D/E|«)v®¶
̸Էgɒʊ6c½ďGJPsaFDPsaJ)ǯƱ4PsaFJ°¶¦´ďGӇ͛ϖF̸Էg[>®~
j´цƇՍðgɒ ʊ6cPsaF IŹȌďI̴G J PsaH/PsaI/PsaL )v® ¶̸Էgɒ ʊ6c PSI-LHCIה̰ΩCJPSI coreIPsaF/PsaJďG4ՙôIj´ҿƇôLHCIׄLHCI4ׅ)цƇ6cv®¤
§IPSI-LHCIGJ8ՙôILHCIׄLHCI8ׅDPsaGDPsaHIլG2ՙôILHCIׄLHCI2ׅ)ǯƱ 6cPsaGDPsaKJLHCI4'`NLHC8IцƇgǵǺŠ6co´mIPSI-LHCI̸Էׄ5L8RׅgƼG 4B÷ʊ4<
|«Gv²²k°)цƇ4<v²²
k°´vԊׄCPIׅ)ūֈ2dcPSI z jҿƇôׄPSI coreׅJPSI|«ITD hEgцƇ4<ҿƇôC#cՔоϒλƵğƇʊ λΩIPSI coreG10|«ׄPsaA-FPsaI-Lׅ )ăǯ2dcׄҶ1ׅPsaGPsaHPsaNPsaO J̰Ω]ёҪׄv®¤§ׅGǯƱ6c)
~jv¯jGJǯƱ4F$:dGŹ4B PsaM D PsaX JÅG~jv¯jGǯƱ6 cPsaP J̰ΩGǯƱ6c¯´ՔŠ´vԊ C#cPSIzjҿƇôGJй100ĻǬIChl a й20ĻǬIβ-r²´2 ĻǬIt´
ׄk²t´5-§²t~k²t´
[4,J¦t´-4ׅ3@I4Fe-4SƵI՜ϱ
־ÃɧׄFXFAFBׅ)ǯƱ4ğʱօµğŠDZ ŹɫµҝĻֈIǵǺеǬìՁʊĻׄPc ]
Cyt C'`NFdׅIՔеՋĜŹɫgұ%~j
v¯jI®zl҅gƔFτͯɲ ŋCƀΊŠ6cD3ՙôIzjҿƇô)ūֈ2 dc3 ՙô̸ԷIǵǺŠGPsaL)ӂFɖŎ g̘<4B$c·˝̰ΩDёҪIzjҿƇô JūՙôCPsaF/PsaJ |«ďGօğɲ v²²k°´vҿƇôׄLHCIׅgцƇ4 PSI-LHCIԎҿƇôgɒʊ6cLHCIIPSI core QIцƇJǵǺC̰ΩDёҪI®zl҅g
Ɣ F τ ͯ ɲ ŋ C ƀ Ί Š 4 B Ļ σ 6 c D PSI-LHCI)ɠadc
zjҿƇôIÅӂFՍĻI̸ԷJ~jv
¯jD̰ΩC`,ăǯ2dB$cŹɫÃɧҿ ƇôׄPSI RCׅJ҅ӽԵ¯vg11[@2
@IϠƈFŹɫÃɧ|«ׄPsaADPsaBׅ (a̸ʊ2dĿ̆Ǭü¿ôI P700ҿ˕I
ǬżȂôTDhEIj´v²²k°
ׄzjµj´ׅgцƇ6c˿уϖFǬż ȂôC#cFADFBJ²£ďIҶGǯƱ 6cPsaCGцƇ6cPsaCJPsaDDPsaEĦG
²£ďGn±t~´цƇ֜ƹgɒʊ6 c·˝҅ӽԵ¯vg1@[@PsaFJ :I N ̉ďIӇ͛֜ƹg°¶¦´ďGǓ*,ɏ bĹ4°¶¦´ďIҶGцƇ6cPsaNDĦ G®~j´цƇՍðI·Սgɒʊ6c
PsaJ J҅ӽԵ¯vg 1 @[@Ȕ2F|
«CPsaFGւʹ6c
3. PSIҿҿƇƇôôIIĻĻǬǬօօƇƇJJǑǑ͑͑ռռԿԿЊЊCC##cc 0dWCGPSI|«gz¶6cՈ ìǬIԛı]ѭӕńɡIӊ̕JͯϒGԺZad B*<)Ƈʊ2d<|«]ziv
¶)ҿƇôQĻǬօƇ6c̸͂Iӊ˦JԼd B$<PSIҿƇôI̸Է(aƆ̸ʊ|«
J֙ɀ͋4,͑ռϖGĻǬօƇ6cD̆ɚ2 dcҿƇôIÃɧGǯƱ6cŹɫÃɧ|«
)²l£¶C#cPSI RCIɒʊ)
˿ Ŀ I Ļ Ǭ օ Ƈ Կ Њ C # c ҡ ё ô z ¶ I PsaAPsaBPsaC Iͅ˅Ǎϊ̦g:d;dν$
<´vԊI°®°Iǻֶ(a˿ĿG PsaB )®zl҅Gʯģ2d: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,$PSI|«IĻǬօƇgßͽ&
6cƦǬׄĻǬօƇƦǬ[4,JĻǬ~¨²´ׅ
)ǯƱ6cJ7C#cҶ 2 GWDZ<ƦǬ) PSIҿƇôIĻǬօƇGխ¿6cDѮ&adc̨
ʤJׅׄͅ˅̦CJPSIҿƇôIңА)Ǔ*
,ȕ6c)PSI|«gz¶6cՈ ìǬIϒδJɕ֕gTDhEż.F$ׅׄPSI
|«DϠÓ÷ν6cFEC#cΪG ɜѯCJġώͣ͒ͪ]Փ͔¶l¯~
¥ׄYeast Two-Hybrid SystemׅG`cլʹϖ
F ˝ ͪ G ` c Ϡ Ó ÷ ν ) ǁ Ə 2 d B $ c
Ycf31,6,11)Ycf41,4,8,16)PYG7/CGL71/Ycf37̰ׄΩ/
v®¤§/~jv¯jCIƉЇׅ
3,13,17)JϤ̧ğƇʊλΩD~jv¯jGă
ǯ2dB$c<ZΪGӂFɖŎgʢ?B$c Dʺ2dcϤ̧ğƇʊλΩCJYcf3DYcf4 J ҡ ё ô y ¥ G z ¶ 2 d B $ c ) Y3IP1/CGL59̰ׄΩ/v®¤§CIƉЇׅ
J̧y¥Gz¶2dB$c:IáIƦǬ Y3IP14,7)PSA22,15)PSA312)PPD15)J̰ΩDҪ
֥GӃĹ2ḑy¥Gz¶2dB$c Ycf3 D Y3IP1/CGL59 J TPRׄtetratricopeptide
repeatׅ¦l´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~jv¯jG[ăǯ2dB$c:
Iɜ®zl҅IҶƱɲ´vԊC#c Ycf3gz¶6c0D)Ϸ2d<1)ҡёôɒԊ ԛˀG`b ycf3 ՈìǬgvjm4<v®
¤§DzIɒԊԛˀôGJ PSI )Ĥ ,ңА4F$<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@
$BJTDhE¾˦C#?<
: 0 C П ѯ a J HAׄHuman influenza hemagglutininׅwׄYPYDVPDYAׅg C ̉М GүƇ4<Ycf3ׄYcf3-HAׅgҡёôGϒδ28
<v®¤§ҡёôɒԊԛˀôg÷Ĺ4 Ycf3 D Ϡ Ó ÷ ν 6 c ´ v Ԋ g j k
k¶еҾ4<ׄƪ2Aׅɠad<Ycf3-HA̻ ƗGJY3IP1ׄYcf3 Interacting Protein 1ׅ)ӃĹ 2d<Y3IP1JzYcf3D°m´6c
´vԊD4BˡGǁƏ2dB$c4)v®¤
§CJjkk¶еҾG`bָ$м ɃI̻Ɨ)ɠadYcf3DY3IP1 )TUɑՙǯ Ʊ6c0D)˦a(GF?<4(4PSI¢¯
JTDhE̱Ĺ2dF(?<D0e) ʂɃIָ$mn´Ļ̕DԊՙĻ̕gұ%D PSIŹɫÃɧ|«ׄPsaA DPsaBׅ)̱
Ĺ2d<7)
Ĥсѽ´vԊgˍȐɲƈðĜоI35SIǯ Ʊ½Cƻ֭4ϫˮլ̻ӱׄ°®°ׅ4<ɜ сѽgϰϯ4®zl҅gеҾ4:dgτ
ͯ ɲ ŋ C ƀ Ί Š 4 B Ycf3-HA g j k
k¶еҾ4<6cD(Fbɐ,®°2d<
PSI RC|«)̱Ĺ2d˜ӄGƇʊ2
d<PSI RC|«)Ycf3GΪϊϖGцƇ 4B$c0D)Ļ(?<͆GсѽgΝ̻ӱI
32S IǯƱ½CЈ4˜ӄƇʊ2d<´vԊ gԮԓׄnlׅ6cDYcf3HA̻ƗIPSI RC |«I®°JǓ*,ȕ4<
ɟ?BYcf3JY3IP1 D§ª¶°gɒʊ4
˜ӄGƇʊ2d<PSI RC|«g·Կɲ
ϖGцƇ4PSI RCIĻǬօƇgßͽ&6cƦ ǬC#cDцӪ4<7)
°®°Iǻֶ(a Ycf3-HA JԶ,Ƈʊ 2dc)¾ǵǺCY3IP1JԼ,Ƈʊ2dc)ǵ ǺC#c0D[˦a(GF?<Ycf3 IãӰƥ ԛԶɃJYcf3GцƇ6cŹɫÃɧ|«
I:dDTUƈ5C#?<)ãӰƥԛ)Զ$0 DI͂ѿϖFζοJ¾˦C#cY3IP1J҅ӽԵ
֜ƹg[@´vԊC#c)Y3IP1ͅ˅̦C [Ycf3JTU͋ȶGңА4Ycf3I®zl
҅QIцƇIǵǺɲJ͋ȶC#?<ɟ?B
Y3IP1JYcf3I®zl҅QIцƇgǵǺŠ
28cICJF,ãӰƥԛ6cYcf3)®z
l҅¼I͋4$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¯vg[>
Ycf3 `bǵǺG®zl҅GцƇ6c®
zl҅gƔF̐çCƀΊŠ4B~¬жȅɃ ŞՒԎՃɧͪCYcf4gĻֈ6cDӃ(.IԊ
ՙ)1,000kDaЊɃI(FbǓ*F̸Էgɒʊ4
B$c0D)Ļ(?<1):0CҡёôɒԊԛ ƪ
ƪ2. Ycf3-Y3IP1§§ªª¶¶°°JJPSI RCIIĻĻǬǬ օ
օƇƇGGɨɨ֚֚CC##cc
Aהjkk¶еҾ4<Ycf3-HA I´
vԊфʊ1הYcf3-HAgƍY®zl҅gƀ
ΊŠ4BеҾ4<̻Ɨ2הλƵI®zl
҅gƀΊŠ4BеҾ4<̻ƗׄȌΠǻֶׅBה
Ycf3-HAеҾ̻ƗImn´Ļ̕Cה´
vԊI°µnlǻֶĤсѽ´v Ԋg35SC°®°4<ɜ®zl҅'
`NYcf3-HAgеҾ4®°2d<´v
Ԋg̱Ĺ4<ׄPׅƈ̹Gnl4<ɜG
®zl҅'`NYcf3-HAgеҾ4®°2 d<´vԊg̱Ĺ4<ׄCׅ
ƪ
ƪ3. Ycf4§§ªª¶¶°°JJPSI coreDDPSI-LHCI I
IĻĻǬǬօօƇƇGGխխ¿¿66cc
Aהjkk¶еҾ4<HA-Ycf4 I´
vԊфʊ1הHA-Ycf4gƍY®zl҅gƀ
ΊŠ4BеҾ4<̻Ɨ2הλƵI®zl
҅gƀΊŠ4BеҾ4<̻ƗׄȌΠǻֶׅ3ה еҾ4<PSI-LHCI̻ƗPSIDLHCII¢¯
)̱Ĺ2dcBה´vԊI°µ
nlǻֶĤсѽ´vԊg35SC°
®°ׄPׅ[4,JnlׄCׅ4<ɜ
®zl҅ׄTMׅ'`NHA-Ycf4gеҾ4®
°2d<´vԊg̱Ĺ4<
ˀͪG`bYcf4IN̉МGHAwgүƇ28
< HA-Ycf4gǓՙϒδ28ƀΊŠ4<®z l҅(aϫˮլGjkk¶еҾ6cD PSIҿƇôIĻǬօƇÃլôgцƇ4<HA-Ycf4
̻Ɨ)ūֈC*<7):I¢¯фʊgӧ RcDʼnԬI Ycf3-HA ̻ƗG͖RBǯƱ6c PSI|«I˕)(FbǑ(?<ׄƪ3ׅ 0I0DJYcf3-Y3IP1§ª¶°G`cŹɫà ɧҿƇôIĻǬօƇGɌ*ь,ԿЊG Ycf4 §
ª¶°)խ¿4B$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$c0D)Ϸ2d<°µn lǻֶJPSIzj|«)·ԿɲϖG
®°2dc0DgϷ4˜ӄƇʊ2d< PSI
|«) HA-Ycf4QцƇ4B$c0D) ϲ(Zad<4(4LHCI|«J®
°2dF(?<
еҾ4< HA-Ycf4 g~¬жȅɃŞՒԎՃɧͪ
G(.cDPSI coreDPSI-LHCIGϠɑ6cσ ĻGёI´)Ļֈ2d<ׄƪ4ׅ:d;d I¢¯фʊgӊ̕6cDPSIzjGϠ ɑ6c´JPsaGPsaKPsaNPsaOäǐ 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$(FbϫˮլGҿƇôGфXԦW dc)®zl҅G¶°2dB$<ȕՙI LHCI|«)PSI coreGцƇ4<DѮ&
adcPSI ͅ˅̦I®zl҅GJq¯{
£¶ΫʃILHCI)ǵǺGңА6c14)':a, LHCI|«IƇʊJPSI core|«
IƇʊDǶĤGJĦɖ4B'a7ͯϒGğƇ ʊƣǹgƇʊ4B$cҡёôCJLHCI)f7(
CJ#c)¶°2dB$cDѮ&adc
ƪ
ƪ4. Ycf4§§ªª¶¶°°DDццƇƇ66ccPSIĻĻǬǬօօƇƇÃÃլլôô
AהеҾ4<HA-Ycf4̻Ɨ(aĻֈ2d<PSIĻǬօƇÃլôY1DY2Jv²²k°Dr²l
gƍY)Y3JΝҐCHA-Ycf4IXgƍYBהY1-Y3I¢¯g̝Ґ4ḆĹCהY1-Y3IPSI
¢¯gmn´Ļ̕G`ḇĹDהY1-Y3ILHCI¢¯gmn´Ļ̕G`ḇĹ
7. PSIҿҿƇƇôôIIĻĻǬǬօօƇƇ§§°°
0dWCGɠad<ĻǬօƇIӊ̕ц̘(a v®¤§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
ç½CJӂC#c<ZՔо)ձȀ6cĻǬօ ƇԿЊGխf?B$ca4$:IԿЊD4BѮ
&adcIJǬżȂôI՜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)ָ$ŹɫÃɧ|«
gĻǬօƇ4Ǒ˕Iziv¶gцƇ 28c<Z˿[ƨ։FĻǬօƇ͑ռC#b:
I<ZǑ˕IĻǬօƇƦǬ)խ¿4B$cDѮ
&adc·˝PSI RC)Ƈʊ2dcDáI PSI|«IĻǬօƇJ͖ԞϖȂ˧C#c Dʺ2dc)0IԿЊGӂFɖŎg̘<6 I)Ycf4 §ª¶°C#cPSI |«
)PSI RCG͋4,цƇ6cԿЊgßͽ&42 aG¾ǵǺFĻǬօƇÃլôg²j¶F EG`cĻӊ(aǴ?B$cI([4dF$
Ycf4 §ª¶°J LHCI IцƇG[խ¿4B$
cv®¤§IPSI-LHCIJLHCIg10|
ƪ
ƪ5. vv®®¤¤§§IIPSIҿҿƇƇôôII͑͑ռռϖϖĻĻǬǬօօƇƇ
2@IŹɫÃɧ|«PsaADPsaB(aŹɫÃɧҿƇôQIĻǬօƇGYcf3-Y3IP1/CGL59§ª¶
°Jɨ֚IɖŎg̘<60IԿЊGPYG7/CGL71DPSA3)²£ď(aPPD1DPSA2J°¶¦´
ď(aխ¿6cDѮ&adB$cɌ*ь*ŹɫÃɧIƐјGǯƱ6cȔƵI|«)ĻǬօƇ4
PSI core)λʊ2dc~jv¯jIPSI coreҿƇôJ»ՙôgɒʊ4ĻǬօƇԿЊJǶÎ6cv
®¤§CJPSI coreGLHCI8)PsaFďGцƇ4PSI-LHCIIÃլôׄPSI-LHCI subcomplexׅ)ɒ ʊ2dc:IɜPsaGDPsaK)q¯{£¶LHCIIÁМGLHCI2)PsaH/PsaG/PsaBIԩĐGцƇ4
PSI-LHCIIĻǬօƇ)ǶÎ6cPSI core(aPSI-LHCIIĻǬօƇԿЊGJÅGYcf4§ª¶°)խ¿
6c
«цƇ6c10)PSI coreIĻǬօƇDJ έК4BLHCII8ՙôC#cLHCI8gɒʊ4 PSI core I PsaF/PsaJ ď G ц Ƈ 4 PSI-LHCI
subcomplexgɒʊ6cȕF,D[0IĻǬօƇ
͑ռWCYcf4§ª¶°)խ¿6c˿ɜIĻ ǬօƇԿЊJLHCI8IцƇgǵǺŠ6c PsaG DPsaKD͏bI2ՙôC#cLHCI2IцƇC#
c0IԿЊG`bPSI-LHCI)Ƕʊ4Ycf4§
ª¶°(aӊֈ6cPSIҿƇôGJziv
¶)Ǒ˕ǯƱ6c)0IĻǬօƇԿЊI§
°GJziv¶IфXԦXIĻǬ̸͂Jƍ WdB$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
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 .
Editor: Ɔ ¨ëǟȁǢêóűÕƈóƀƏƈȂ
Čĵ êЀ ĎǕՌׄÚՑǓׅ 158
DŽNJ ̴ñĦΞΛ֣ɤդG`cğƇʊϮЕI˜Ȝժ ̠ξ Вׄ̒šǓׅ 159
DŽNJ ̰ΩG'.cсѽµфѠ±°CIՈìǬϒδӊ̕
ɇυ˓ͭ¼̊ɵȹՃҩׄ͞ÚՑǓǚҏĞМǓׅ 171
DŽNJ y¤v±v~¬´'`Nl°¶nl´wgν$<
÷ΩѹЋIŖβеָԶŠ ՎǧȢξͭøׄ̒ÚǓׅ 180
DŽ
DŽNJ NJū ūDz Dz
®ÎģƀƏ/ķ!ŦşE<&@"ķĨƺķžǀ
Čĵ
‡
Ɔ¨ëǟȁǢêóűÕƈóƀƏƈȂ
$@Iˮã[˜<FʗҲ֑˜)ЄDZG˜4$ϪӃD֧ԖϖFϒȜg[<a6ԩȺCJ͆Àã
~¶vo´|¶gν$<ԎÂľՈìǬՒľӊ̕)̹FλΩIy¥ɻǁ]mRNAϒδɻǁIŻɠ ǍϊՈìǬƈǺgƀѿD4Ƽ϶ϮЕIXFa7ŤDZ]ԣDZD$?<ɫνGԩ$ĻG'$B[֑
˜g[<a4<W<TALEN]CRISPR/Cas9GãҶ2dcy¥ѕօʗҲIϒȜJy¥¼CI èɿIՈìǬIˌǍgƀѿD4δãλΩDZI̹ϠDԺ͌I¶g·Ǎ28B$c2aGJv
®lqǬ֣ɤդgν$<ūвǬӊ̕G`c̸Էӊ̕ʗҲJX ѓц˴̸Էӊ̕CJƨ։F´v ԊҿƇô̸Էӊ˦IƀѿɲgǓ*,ȿ/cDD[G̸Էӊ̕:I[II˘Șg½/cDѮ&adc ğƇʊϮЕIĻG'$B[:%4<ʗҲ֑˜IɴɷJǓ*,0dWC̊ӞC[ӑÐD4Bтß 4B*<2aGÞɜIϮЕĻIϒȜI<ZGJʋϮЕѯJ˜4$ʗҲ֑˜QIʪʌDD[
Gժϒ2dc˜ʗҲQIj´gȶGɏ?B',ɨӂ)#c
:0C̊ΪօӑÐCJğƇʊϮЕG˜4$Δͱg[<a6˜ʗҲµ˜ϪӃ D֠4B̹F ĻCͯϒGϮЕ2dB$c3w°¶Gӛ֠ʾüg'֤$4<W7̒šǓDZI̠ξВŬNJGJ 1҈ԗCժϒ2d<̴ñĦΞΛ֣ɤդgν$<˿˜Iʊ̘g1тß֖$<:0CJ¶Ն ЈFEI®zl҅´vԊIl¤vgϟʹӈȉC*cƀѿɲIXFa7ğƇʊ҅´
vԊIūĻǬĻğG`bPSIƐԤIo°u¶ЈŗIl¤vׄ˄a+ׅgӲӪC*c0D) Ϸ2d<W<ǚҏĞМǓIՃҩ͞ŬNJIw°¶(aJфѠµсѽ±°CIՈìǬϒδӊ̕
Iʒͪg̶Ӥ֖$<ՃҩŬNJIw°¶Jέ҈Gժϒ2d<ԧԶфѠĻֈʗҲgν$B0dW CG̰ΩI̶ˢˮӎIфѠΪϊɲg˦a(G2dB*<̊ӑÐCJ̡ΫфѠDͶΫфѠIğƇ ʊG'.c͂ѿĻŠFEğƇʊϮЕѯGֲ̝X]6$ӛ֠(aǤW?B˿˜I 1 сѽ®´v
¯¶¥ʗҲWCӊӤ$<=$<:4B˿ɜG̒ÚǓDZIȢξͭøŬNJIw°¶G÷Ωѹ ЋIָԶŠDŖβŠgƀѿD6c2@IʗҲy¤v±v~¬´ׄGSׅDl°¶
nl´wׄHPׅG@$B:IàфXDδΫg1тß֖$<˿˜IѹЋʗҲIѻ˲G#c ϧ°IеȅFζӪGַ(2dcDD[GGSG'.c̰ΩIθdžɫУI§°Š]HPG'.
cӎ֘ϞµǺʗҲFEG'$BJğƇʊϮЕѯDIz®¡±¶~¬´Iƀѿɲgʂ5<
ä¼ĻǬсѽфѠ:4BćôI±°C:IΪɲgʡĹ4δӷIŲƦgζӊ6c<Z I˜4$ʗҲD:0(aɠadcϪӃgтß4<δ͑ռCJğƇʊϮЕDIխĀJ˦ϲCF$[
I[#c([4dF$)$7d[˜4$ΔͱgλYƀѿɲ)#cDѮ&B$c̊ΪօӑÐ)*?
(.DF?BϊĻլIØͱ]˜4$ĻQIʪʌG@F)dKȻκC#c
˿ɜG̊ΪօIѕօG#<?BJ'ɪ4$ÃƺПѯ̢ӥѯI˝GǓǍ'ÀӛGF?<
ΪGՃҩw°¶Ȣξw°¶IҢѯIϗ̹GJϊĻIևӞ(aIЗΟIý֟G[խfa7 ɭ,ӑÐIƺПgɌ*ż.B֖$<0IǂgċbBĤBIϗ̹Gͻ,ɡϸπ4¼/c
‡ӊӤΪօğƇʊϮЕG˜4$Δͱg[<a6˜ʗҲµ˜ϪӃ
1ԸчĞE-mail: [email protected]
DŽNJūDz
Ŏ
Ŏ Ť Ť° °ũ ũŨ Ũǻ ǻę ęǧ ǧ/ /? ?B B® ®Î Îģ ģƀ ƀƏ Ə0 0ķ ķý ýǪ Ǫ
‡
ŅÃêóêóǯ ůóƀƏƈ ʼnų Ǝ*
ĤǪŹ!*&Ŏ Ť°ũŨǻęǧ/?AĘ@C&Ŀǒ0ģň/)*ãÓ"B80 K+1®Â óƙ IȁPSIȂ®Âóƙ IIȁPSIIȂ0Ʒ®ZrSc)0m S/´Ǵ"B0,E¸Ų!
*S}vdyeZ0®Îģ]lSnjÍģ´0ƜƬ²´ćE®óǻęǧ/?AÌǂÂ"B,/µ
;*ģ½!&Za cǛƉºė/BÔǑHae PSIPSII 0-'@,°üÝ"B0Eō Lj!&8&PSI0Ʒ®Èŭǿ.B80 K/*Synechocystis sp. PCC6803@ÅǴ!&
Å0PSIǦ¡@0Ʒ®ZrSc0ĻǫçÂEťõ"B,/ģ½!&Ʒ®ĒĎm S
Ƨ0İ@ǃť CPSI0Haeƙ0^Jev`S.©ǷEĹ@/!&
1. JJ55ZZGGההğğƇƇʊʊĻĻGG''..cc̴̴ññCCII֣֣ ɤ
ɤդդӈӈȉȉIÍ́ννɲɲ
§̊ӑÐCJПѯa)ժϒgԺZB*<̴ñ
֣ɤդgłν4Bұ?B*<ğƇʊϮЕI˿
ԩIʊ̘G@$Bтß6cäʼn(ağDZ֣ɤ դgν$<ҡёôīՍIw®̸ԷIӈ1, 2)] θdžǍŠGí%w®|lIǍŠIӈ3, 4)F E)ǁƏ2dB*<0%4<ϪӃJ®zl
҅I̸Է҈ô)Ǎŗ4:dGɌ*7adB
®zl҅GƷZԦWd<´vԊԎҿƇô I̸Է[Į̸Ъ2dcD$%l¤vFʽ ĖIɨӂɲgϷƚ4B$c0%4<ǁƏäʼnG [û&K¶ՆЈG'$BJƐԤj´
зC#c LHCIIׄLight-Harvesting Complex of Photosystem IIׅI®zl҅īIЈŗ)ɾǺ2 dB$<bğձȀgż.<ğŠDZз IIׄPSIIׅ Jw®£¶´GԽKdBĆɢgż.cDѮ
&adB$<bD®zl҅īՍI´v ԊԎҿƇôIl¤vFǯƱ̹ɋJȿ,ӟ ӱ2dB*<4(4l¤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
J®zl҅Il¤vgӊ˦6c<Z Iϟʂϖɻǁgʾü6c[IC#b¼ӑIƛ֠
GȌ6c´gʾü6cD̆ɚ2dc ·˝Cv²²k°I҈ȁҮğg̱Ĺ6cҮ ğ֣ɤդIǂƇC[ğŠDZз Iׄphotosystem I:PSIׅD PSII gţŁ4B:d;dIсѽīĻȲ gƀӅŠ6c0DJ$,@(I¯t¶Fʒ
ͪ5, 6)gν$F.dKƨ։C#?<ǽCJPSI
IҮğźβJ֏ȶGñ,W< PSII I¶vD Ļֈ4B$F$0D)ƨ։IÅӂFŲƦC#c ʋ)ժϒ4<̴ñ֣ɤդCJôИоFE IijǨgν$<ijŰG`b80 KЊɃIɃCI
֣ɤդӈȉ)ƀѿC#c7)`,Ϫadc`%G 80 KCJǑ,IğƇʊλΩCPSIDPSIIJ˦ϲ GĻֈ4<Үğv°gϷ60D(aՅɑ FĻğҺѣ)#dKPSIDPSIIgţŁ4Bӈ
6c0D)ƀѿDFcLHCII)PSIDPSIII լgɘ̑4B$cDѮ&adB$c¶Ն ЈFEIδӷgӊ̕6cǂƇGPSIDPSIII
®zl҅īIȗƱgƀӅŠC*c0DJ֏ȶ ǴŖDFc0IӑÐCJПѯa)ұ?B*
DŽNJ
<¶ՆЈG'.c LHCII IсѽīĻȲI ƀӅŠIϮЕ8)gтß6c
[%·@I̴ñ֣ɤդIɫνûD4BğƇ ʊ´vԊgȌӷD4<ū·ĻǬĻğIϮЕ7,
9)gтß6cū·ĻǬ(aIҮğĄƅg̱Ĺ6 c0DJǑ˕IĻǬ(aIĄƅg̱Ĺ6cǂƇ GȹƴŠG`bց2dB4W%ĻǬ͕I®@
*gӈ6c0DgƀѿG6cū·ĻǬ̱ĹG
`bɠadc[%·@IӂFɻǁJĻǬIˮ լGýǯ4<˄a+Gխ6c[IC#cū·I ğƇʊ´vԊҿƇô(aIҮğv°
IхˮǍŠgԮԓ6cDǽC[̴ñC[֞
ўG˄a+)ӈ2dc0D)ǁƏ2dB$c
9-13)˄a+I̱ĹJğƇʊ´vԊIl
¤vFǯƱ̹ɋg˦a(G6c0DGџ) c
2. ̴̴ññ֣֣ɤɤդդIIժժϒϒ
֣ɤդIЖլĻӊѿJȌΩ±´IժŽ˕
ׄNumerical Aperture: NAƪŵΠׅC͢Wc
̾˝ƊGJͫէIŧĻg NA CŎ?<ЊɃI|
lğԜ˝ƊGJͫէgNAIÑÉCŎ?<Њ ɃI|l)ĻӊC*c'`:IճτDFc ğDZ֣ɤդG'$Bָ$NAgɠcGJׅȌ Ω±´D|´°DIԒֈgϫ,6cW<J
ׅ|´°DȌΩ±´IլIЖլgָ$șʛ βIΩԊׄͦFEIôׅCΆ<60D)ɨӂD Fcׅgǻδ6c<ZǽCJșʛβIָ
$ôCȌΩ±´D|´°IլIЖլgΆ
<6ͷ±´)ʷν2dc)̴ñCJÞI D0e0I˝ɋIǁƏJF$ׅgǻδ6c˝
ͪD4Bô¯m¥FEIijǨIF(G|´
°D·ђGΆ<4B4W%˝ͪ)#cΊүϭ җJΤ҆ɏĀ˕)֏ȶGȔ2$ΩԊC#b̴
ñG4B[źќ]Ǎɒ)ԍ0aF$<Z̴ñ
CIúνGѰ&c±´gǻδ6c0D)C
*c14)>FXGô¯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)̴ñC͍h=b±´gդТ GƫǺ6cʹϦŋ[̴ñGѰ&adF(?<
b6c<Z¶°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<̴ñ֣ɤդI̶ψ ƪ'`NǻտIҺѣıϤgϷ6|´° °
DȌΩ±´լIЖլJָϤЖGă<dc<Z Τìȓ)ʙ&adȌΩ±´҈ôJijŰ2dF
$0I<ZҐźȮҽ͋'`NĖ΅˻źȮҽ
͋)˞2d<Plan ApoƵIȌΩ±´gʷν6 c0D)ƀѿDF?<|´°ΊJ̗Iϭ җs®CʬWd<ЖլGȅթ2dcICϤЖ G2a2dcf.CJF$0I˝ɋIʷνG`
bժϒ4<ɑˮGJ̴ñ֣ɤդD4BÀτ˿
ָI NA0.9 gǻδ4ЖլĻӊѿD4B̾˝Ɗ
й350 nmğԜ˝Ɗй1.2 µmgՁʊ4<ǻտ IȌΩ±´JָϤЖÃGѣ(d<տGīՍG Ж͚)͏aF$`%GȱӼƗׄM Plan Apo HR×100NA0.9MitutoyoׅGЖ͚ǭ)ժ.ad
<ΪͬI[Igúν4B$c¡¶°H5ׄBSׅ gƥԛ28c0DCȌΩ±´gƫǺ6cɁ̔
)¼½Gŗ$BΞΛIгӧ˖gұ%îќƀѿF
²¶G`bָϤЖgă?<WWɁ̔I¼½ ŗ)ƀѿDF?B$c|´° °JΤ҆ɏ Ā˕)TU 0 C#cϭҗҾI¶GƫǺ6 c0DCɃǍŠG`c¯gʠ2&B$
c|´° °Dv®lqIz¶°
ׄCHׅJΤìȓ)ָ,̞Ԛɲ[#c
՞¦~ªCџ$C$c0I0DG`bϟѓ ȓ ģ ͂ g ß 4 < | ´ ° ð ѣ I г ŗ ) ƀ ѿ D F?B'bָ$ˊ÷ɲgǻδ4B$c Ɂ̔ÃǗIЙ(a±¶}¶)ȓģ2dȌΩ±
´G`b|´°¼Gօğ2dcs°¤
®¶ȌG`b±¶}¶ðѣIԌ̢gұ%|´
°(aIҮğJƈ5ȌΩ±´G`bօZad
̱ĹзGԽKdcĻğƣGцƇ4<CCDr¦
®G`bҮğv°)Żɠ2dcĻğƣI ģbŽ)´ ¶°GF?B$c0DCĦΞΛ
֣ɤդD4B÷ŗ6cW<|´°IԳԿĖ J֣ɤդ¼ՍGӔѣ4<LED(aIğgν$B CMOS r¦®G`bŻɠC*c`%GF?B$
c
3. ссѽѽīīğğƇƇʊʊ´´vvԊԊIIččƱƱggƀƀӅӅŠŠ ȇ ǔ ƻ Ʋ C ƻ ֭ 4 < v ® ¤ § с ѽ g
iׄpH 7.520 mM tricine50 mM NaClׅ GʉΘ4<ɜ̴ñ֣ɤդI|´° °Q
4<0I͑ռCJ|´°Ƀg23°C Gӧ˖6c15ĻI˶֙ɫɜGPSIğׄ710 nm 170 µmol photons m−2s−1ׅgΠȐ6c0DC
¶QӠȓ6cğΠȐժǤɜ15ĻCijŰ gժǤ6c¶ՆЈӠȓğIΠȐJ|´
°Ƀ)80 KGՁ6cWCIй30Ļэʨ4<
¶ӠȓIտGJPSIIğׄ470 nm120 µmol photons m−2s−1ׅgν$Bƈ̹Iˊ÷gұ%
PSIğPSIIğJƪILED(aIğg´
k°GԵ4Bɠ<֣ɤդCǺ2d
<80 KIсѽIҮğv°GJ#cЊɃI
®@*)Ӄad<)Ǒ˕IсѽCȹƴ6cD
ԵȶIҮğğɃӎG`cǺDTUƈ5v
°)ɠad<8)
ƪ2G80 KCӈ2d<v®¤§с ѽIĦΞΛҮğ֣ɤդĖgϷ6ŔԍͫէJ445 nmC#cƆv°CƪMN GϷ4<`
%FҮğv°)Żɠ2dcʋJĤB Iv°IҮğv°gא@Ismխ
˕IƔCkk´w4<kk´wI О·͑ռCJ10 ćЊɃIсѽIv°g ĤB%W,Įδ6c`%F 5 @Ismխ˕I
®¦¶g͞Zcşʦϖׄglobalׅkk
´wgұ%О·͑ռIkk´wCJƆ smխ˕I¶vðѣDȷJϊFcсѽCƈ 5ČgŻc)ʮȷJϊFcD$%̏љ̐çI
½Cұ%0IО·͑ռIşʦϖkk´w Iц̘5@IsmʊĻI¶vðѣDȷ)͞
Zadcƪ2 MN(aĻ(c`%G¶
G`bҕȸ¶vͫէGՂ$JӃad<)5
@IʊĻJϫͫէ(a֙G680 nm686 nm692 nm712 nm730 nmâԩG¶vgʨ@˿[
ϫͫէI680 nmʊĻׄёGǃb@P2d<¶
vׅgÅGLHC(aIҮğDȵȝ4<686 nm 692 nmI2ʊĻׄ֍Gǃb@P2d<¶vׅ
JеҾ2d<PSIII77 KҮğv°DI ƪ
ƪ1. ̴̴ññ֣֣ɤɤդդII̶̶ψψƪƪׄׄȫȫׅׅDDǻǻտտIIҺҺѣѣııϤϤׄׄƄƄׅׅ
MO: ȌΩ±´PZ: o jvªo¶LM: ϟѓȓģ͂CH: v®lqz¶°
QS: ΊүϭҗҾ|´°¶BS: ΞΛгӧ˖ν¡¶°H5RM/HM: ŹȐդ/ŧԳԿդƄIıϤƄ
½J|´°ǽȫƪJ˙ΰ7`bˌǍ
֥ï(a PSIIIҮğDȵȝ4<էͫէI@
IʊĻׄԋGǃb@P2d<¶vׅgPSIDȵ ȝ4<ОÑ͑ռCJ͢Ǻ2d<smʊĻI
¶vðѣȷgƫǺ4BʮȷIXg¯¶
® ¦ ¶ D 4 B Ĥ v ° I v ° I
kk´wgұ%0%4BLHCPSII PSII3@IʊĻIҮğɐɃmap)ɠadc ƪ2 A"CE"GI"KJ0I`%G4Bɠ ad<ƆʊĻIҮğɐɃIсѽī map C#c
˿¼͑G¶QӠȓ2d<сѽ¼(a
͑ϞG¶QӠȓ2d<сѽIĖgϷ4
<¼(a3͑ϞGJ¶QӠȓ4<) PSIҮğɐɃILJɐ)áIсѽG͖RBťĻCJ F(?<сѽIĖgϷ600CJ0I`%F сѽgՆЈ¾ťĻFсѽ DƒP0DG6c Ǒ,Iсѽgӈȉ6cD·ǺIŎƇCՆЈ¾ť ĻFсѽ)Ӄad<·ωƄIDHLJ:d
;dIсѽIPSII/PSI ͖βmapC#bPSIIҮ ğĖgPSIҮğĖCŎbЧ4Bɠ<¶
GӠȓ2d<сѽIûC֣ҢC#c)LHC '
`N PSIIIҮğĖCJсѽīGс($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$%Ð ɻ)#cDɰfdc®zl҅īG'.c PSIIDPSIIčƱJPSII/PSI͖βmapC`b֣
ҢGӃadcv®¤§CJָР̰ΩI
`%G PSII)ȗƱ6cw®̸Է)ϒՁ4B$
F$0D)Ϫadc)0daIǺ¶)Ϸ 6`%G#cЊɃPSIPSIIJϊFcȗƱgϷ 6`%C#c0I`%Gժϒ4<̴ñ֣ɤ դ)v®¤§G'.cPSID PSIIIč ƱgƀӅŠ6cIGťĻFЖլĻӊѿgʨ@0 D)Ϸ2d<0I0DJä½Iӊ̕gԺZc
¼CIʼnʾDFc
¶ՆЈG'$BEIЊɃLHC)ÁğŠ DZзլgЈŗ4B$cI(g˦a(G6c<Z GJ¶:d;dGӠȓ4<сѽC LHC)PSIPSIIDEIЊɃƈ5ǂʐGǯƱ6c (ׄĦȗƱׅg˦a(G6cɨӂ)#c0I0 DgǺՙϖGӘþ6c<Z·@IсѽG'.c Ɔv°CILHCʊĻIϠȌɐɃgњԜG :Iv°G'.c PSII/PSI Čg̾ԜG4<
Ϡխƪg÷ʊ4<ׄƪ3ׅ0daIϠխƪÃI
·@I¶ΛJ:IсѽIҮğĖI#cv
°G'.cČgҶ4B'bƄGðѣ6c¶
ΛJ͖Ԟϖ PSIIIǑ$сѽīIՍðGȫG ðѣ6c¶ΛJPSIIǑ$ՍðGϠɑ6c ƪ
ƪ2. ¶¶ׄׄA""Dׅׅ¶¶ׄׄE""Hׅׅ G
GӠӠȓȓ22dd<<vv®®¤¤§§ссѽѽII80 KGG''..cc
̴
̴ññ֣֣ɤɤդդσσĖĖ
I"LJ¶ՆЈ)ťĻGԍ0aF(?<сѽ IҮğĖׄAEIׅׄBFJׅׄCG KׅJ:d;dLHCPSIIPSIGȵȝ2d<Ү ğʊĻCĮ̸ʊ4<ĖgϷ6ׄDHLׅJ PSIIIĖgPSIIĖCŎ?BɠadcRatio map
¶J2 µmgҶ6MNJA"D'`NE"H GϷ4<сѽI#cv°G'.cҮğv
°ׄΛׅDא@Ismխ˕IƔC¿&adc
kk´w˻ѓׄԋǻѓׅƆsmʊĻg ǃb@P4<¶vCϷ4<˙ΰ8`bˌǍ
ƪIϠխƪCƄѸ¼)bIĻȲJPSII IǑ$ՍðCLHCҮğ)ɐ$0DgϷ4Ƅ½ )bIĻȲJPSI Үğ)ɐ$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$%ĔƊgϷ 4<0DGFcƪGӈ4<6RBIсѽ IR Čg:IсѽIPSII/PSI ͖ׄсѽÃIĤ
v°CȹƴׅGȌ4B²4<ƪC JƄGðѣ6c¶ΛJ¶Iсѽ ȫGðѣ6c¶ΛJ¶IсѽC#
cԋҐI»ӉJ710 nmIğgΠȐ4B¶
GӠȓ4<сѽI¶C#b֍ҐIÄJ 460 nmIğΠȐG`b¶QӠȓ4<с ѽC#c¶ՆЈӠȓg4<G[խfa7 ՆЈ)¾ťĻC#?<ՆЈ¾ťĻсѽׄͺ$¶
Λׅg·@Iw°¶D4:däǐIсѽg
¶QӠȓ2d<сѽ¶QӠȓ 2d<сѽIw°¶GĻ֥4<¶
w°¶C¶w°¶D͖Ԟ4B R ČJҕȸȔ2,F?B$c)Ǔ*FǍŗCJF (?<:I·˝CՆЈ¾ťĻw°¶Iсѽ CJRČ)́ɿGȔ2,F?<сѽ)$,@(
Ӄad<0daIц̘gӊՕ6c§°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. ƆƆvv°°GG''..ccLHC Ү
ҮğğIIϠϠȌȌɐɐɃɃgg::IIvv°° G
G''..ccPSII/PSIČČGGȌȌ44BB
²
²44<<ϠϠխխƪƪ
¶ׄȫׅ'`N¶
ׄÃǗׅGӠȓ2d<сѽ ƄJՆЈ)¾ťĻ=?<сѽ Ɔw°¶4 @IсѽIц̘g Ϸ4<˙ΰ8`bˌǍ