ŀƞʈσϡ 28 (1) 2018

光合成研究

Ϩ 28 ɀ Ϩ 1 ƛՖӗɀ 81 ƛ՗2018 Ɋ 4 ˪
NEWS LETTER Vol. 28 NO. 1 April 2018
THE JAPANESE SOCIETY OF PHOTOSYNTHESIS RESEARCH
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トピックス

ŀżǿЁ I ÞӹĖ-½Ó±¡ÖѲƞĖn X МЏˡ̞ә35ͅƞՖéĥΗ՗țǺ, ̊̉ ͥƺՖǦԆǦ Ă՗
解説 ɯЊѢԹlLSоүԵϻљЛĖȧǇґ}Rdgĕѡγ̤Ֆ˺ûǦ՗

研究紹介 Лы̖Άn PSI-PSII ҴѲƞĖ

研究紹介

6

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ŀƞʈσϡ 28 (1) 2018

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ŀƞʈσϡ 28 (1) 2018

トピックス
光化学系 I 三量体-フェレドキシン複合体の X 線結晶構造1

大阪大学蛋白質研究所

2

基礎生物学研究所 環境光生物学研究部門（現所属）河合（久保田）寿子 1,2、栗栖 源嗣 1

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5ŀƞʈσϡ 28 (1) 2018

解説
微細藻類における脂質顆粒葉緑体局在説をめぐって

東京大学

大学院総合文化研究科
佐藤 直樹*

近年の微細藻類を用いたバイオ燃料研究ブームのなかで、クラミドモナスでは脂質顆粒が細胞質だけ
でなく葉緑体にも存在するという説が生まれた。しかし、他の生物で細胞質に存在する脂質顆粒がク
ラミドモナスでだけ葉緑体にも存在するのは奇妙であった。筆者らはこの謎を解明する努力を続けてきたが、ようやく一定の結論に達した。無いものを無いと証明することの難しさを含めて、簡単に解説する。
1. oZ}lÛӭn LD oЊм̍l~ǽǇYg_ņнҕϰ §θ̍ΔΆnǤQoӡŧk̈Ձ°Ñ¡Ò[ƘнɻNʤʻWĂ?Ç°Ö±ÑinÑ¥ÔÚÒ (TAG) nɠhĥǽ[нŪ~dоү°Ð½¬?ʃ̃YaÐÇ¢nǛ̴gLŭS̧нkj~πgI@̖Άn LD le?_nԖ?TAG oоүԵϻ (LD) nɠhЊмүlǽǇ[Ֆǁ՝՗@UUhоүԵϻi҄

IgoˉΊ 4 kjƎͼnUi@

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zkЊмlƤz LD NłΥkjƤ|Ǥ̟kΔΑ̧нlԅUiNπgI 2,3)@ˉΊ 1loM[IǁѽNχWgI@Wl?

*ӚАĿ E-mail: naokisat@bio.c.u-tokyo.ac.jp

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xMl̖ΆnљЛĖlo?Ѓ 0.1 ~ 0.2 µm nγɥlӸΔ̌h~ɞŀ˸ĈhљЛĖl LD Nќϟ[~e¿Ð£°Ô»ÌÚÒ (PG) NH?èlinǗƧNΦѩWa 21)@LD nљЛĖȧǇl¿Ð£°¶Ö?°½ÔÚÒkjn§¿ÓԅYgϩвoɞIΤɸʛO?ŇƠσϡвii~¶±ƤhIN?TAG ~ƤzUiNlЃ 2 ɊԄlaπgI

7,8)

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?Ò

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?PG lȚɶLD nљЛĖȧǇƵԶlԅYg?_nЎРѽґ®Ñh~ȡWkоүƤ˫ԵϻNǽǇYÖkjƤ|UiNπgLbN?_nJlĦ\wO̎ʠoÿniUń10)[ƘнɻNHN?zbҊЊoMkI@I

Y?ΏǇnΈ͉ґ˗YaI@

zniU?¡¶¸®Ñl TAG NǽǇ[iIJυȋk҅ʠokI@

2. ԡǺԷɯӿlоүԵϻ (LD) nѻȗ 2000 ɊĄMǴzdaѢԹΖI¸΀ ӸΔ̌nÐÇ±Ê²£ЊмϤЉԽՂ˸Ĉ

11-13)

næh?ÐÇ±Ê²£l[i?¯Ö¿Öϻi LD NǤӹlќϟY?ӗ

(Chlamydomonas reinhardtii) iƫpЛѢNÊ

ɆnŀǿԷɯӿѻȗӕӡǌԡǺԷɯӿ (TEM)ˊΔΕnσϡ

14,15)@UnxMl?

lҴўŖ΃nѻȗh~?_`NЊмŌn

Nannochloropsis ԹՖθκ͸Ѣ՗ Botryococcus

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brauniiՖЛѢ՗?Dunaliella ԹՖЛѢ՗?Chlorella

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12)@β[p?љЛĖnŌӭiЊмүiѵŕSU

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15,16)TAG ¯Ö¿Öќϟ[UiNπ?

H@ϩвoԀɊ?љЛĖŻтn DNA Џƞ©Ö¹үσϡYgOa 23 ԅĢ~H¯Ö¿Öƞʈl̪ʸnHǡΡĖho TAG ќϟNǛŬYa

17?18)?ŻтnԡǺ

@_nԖ?LD nǤQoЊмүlԷɯӿѻȗlo·̵n͎ʁʑdgOa@˩ӎhǽǇ[N?Ûӭn LD NљЛĖŌӭlǽǇ[o?nŻтnтԄԱǐlÂ¿«±ÑÖN

UiNôenÒÚ¿MǗƧWa

19,20)

ǽǇ[҅ʠ? TEM ĹMɫUilʈūY@U

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9

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Received Feb 23, 2018; Accepted Mar 15, 2018; PublishedR

April 30, 2018ӌbΈʄlk

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IN?Ǥӹn¯Ö¿ÖNӫՍ[Uih?LDƎбˉΊNљЛĖŌӭl͙Q~R

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ӌ|UioԞYIi

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 ϑNΤƵlɹJUinÛeo?LD nљЛĖȧBiochem. 81, 687–714.
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Ǉlԅ[җˉNIQe~ΦѩW?ǤQnσϡ

T. C. (2009) Lipid droplets at a glance. J. Cell Sci.

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ɶΖɴƢNɞQ?̸ӃΪƭMhHz

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ƌYIΑ

Welte, M. A. (2015) Expanding roles for lipid
droplets. Curr. Biol. 25, R470–R481.

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plants: thematic review series: lipid droplet synthesis

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Naoki Sato
Department of Life Sciences, Graduate School of Arts and Sciences, University of Tokyo14

ŀƞʈσϡ 28 (1) 2018

研究紹介
緑色植物の PSI-PSII 超複合体§
1

日本製粉株式会社 イノベーションセンター

2

北海道大学 低温科学研究所

3

北海道大学 大学院理学研究院

4

カリフォルニア大学 バークレー校

5神戸大学 内海域環境教育研究センター

6

神戸大学 大学院理学研究科

横野 牧生 1,2,*、高林 厚史 2、岸本 純子 2、藤田 知道 3、岩井 優和 4、村上 明男 5、秋本 誠志 6、田中 歩 2
全ての酸素発生型光合成生物は、2 種類の異なる光合成反応中心 (PSI と PSII) を連動させて光合成を行う。従来、PSI と PSII は別々の場所に存在すると考えられてきた。これは弱く安定した光の下で光合成を行う、先祖型の酸素発生型光合成生物においては正しいものと思われる。しかし陸上植物においては、約半数の PSII は PSI に緊密に結合し、PSI-PSII 超複合体を形成している。超複合体内部では、PSII から PSI へ励起エネルギーが 20ps で伝達される。
本稿では、シロイヌナズナの PSI-PSII 超複合体において、PSI がエネルギーを熱に変換して捨てる反応を促進していることを報告する。この反応は、突然の強光などで PSII から PSI への電子伝達反応が追いつかないときに自然と効率が高まる。この PSI-PSII 超複合体によるエネルギー調節機構は、植物が変動する光環境に適応するために獲得した重要な環境応答機構であろう。

1՛oZ}l12iYaΓǚnǡżhäвnŮҳµÒÚn¸

 ŀƞʈƒɶæɲo?ŀnµÒÚÔÔÐÖ£Nȶ?΃ˎnƒɶæɲlӡŧkŮҳµ½ÒŕǺŌnԡǺnͳYIӟų (ŮҳµÒÒÚNͧzÚ) lЩOʵK?ľŕkŷI (ԡǆ) âK

gYzJbJ@aԡǺn͑Δ{œ[Uih?ɞŪkӪĽť Ώǽ[˩~ǴƋΪkӵЉΦΔǌŀƞʈΔΆƞʈ[@ӪĽťo?ȃȈkôӵżͷЉlˍaiW¡¶¸®Ñ Gloeobacter violaceuskŇ˫Џƞę]g?Ͼęo?££nȱɢhΦѵW?ǒՁΓǚh~ɯɜœ[nláƘ̪?©Ö¹үтоүNτǞW

hH@̖ΆƤ|ӵЉΦΔǌnŀƞʈΔΆho?

ŀhYMǛ̴YkI@Ûˎ?ǤQn¡¶¸®

PSII NŮҳµÒÚΖIg̼ŕǺMԡǺÑo?ǡųnͳYIŀ˸Ĉlԯɶ[a}?

əOʙO?PSI N_nԡǺlWlŮҳµÒ

̟AkΓǚɶϭ̧̞΋ɫYgOa@_nJcnÚâKUih?Ѓ 1V nԡǆΔ{œY

r i e N ? 2013

NADPH ƞʈYgI@~YÛӚnƒɶʝJ

Phycobilisome-PSII-PSI megacomplex hH 1)@

PSII i PSI NΉϥYgǽǇYgIk?cd

PSII i PSI N ? Û e n Ƚ Ǧ k  Ö ® ²

Ɋ l Φ ѵ W  a(phycobilisome) lƠ˜lЏƞ[Uih?ŮҳµÒÚo PSII i PSI lӦŖkŨƞhŕӱW§

Ϩ 8 ƽˑ˳ŀƞʈǿčɊč Å£©ÚΦѩҭƕҭҗˉ*ӚАĿ %!$ !$!%'%

@UnāЍ{o phycobilisome nǽǇlğǽY

15

ŀƞʈσϡ 28 (1) 2018
?ЄѢoUnāЍ{ƕSДIb 2,3)@YnнŪbShkQ?µÒÚ͗ƍ[ůΌnMY phycobilisome ǪdaЛѢԹnЁВho?

ѻ͸M~ӷѴhH@PSI oȢkQi~ 2 ϚԹśnɠh PSII i PSI nŮҳµÒÚn¸ÐÖnŀԇɬ̧̞ĵKgI@Ûeo zeaxanthin i£iɳѴNHda@

low-energy chlorophyll n Ԅ h ɠ ʈ W  

gLcharge-transfer state ĀYaÖ«ÖhH2՛ЛѢЁВn PSI-PSII ҴѲƞĖ8)?~JÛeo low-energy chlorophyll M P700+ ЛѢԹlLIg͖n͙IiUlΔɽ[ϚvnµÒÚϗųĀYaÖ«ÖhH

ho?ǤQn PSII o PSI ioΉϥYgǽǇYg

 9-12)@low-energy chlorophyll N͙QkiլP700

I (ǁ 1a) @UJYaϚhoլ_`nƒɶvµÒÚĎӣ[ůΌoēà[NլæɲlǤ˅nŀʪԚÖ®²NЏƞ[Uih?

zeaxanthin inԄh charge-transfer state ɠʈ[@W

ůΌ P700+vµÒÚĎӣ[ůΌoՌlUnϚn PSI o?P700 iϧƞYgŮҳ

Qk@YaNdgլ͙I low-energy chlorophyllµÒÚ°Ð¬¿[ low-energy chlorophyllnǽǇoµÒÚ͗ƍ[ůΌՌ}͸ůΌъQŀƞʈѦdgInhHJ

4,5)

6)

(red chlorophyll) ʣakIUiM ?ՌϪ̖ΆhӷѴhH@n PSI  ̖ΆoԏßnƌYIŀΓǚlӦɶYaϚ

~ŀʪԚнŪloļgIiбK

@

hH

 ЛѢnæhԏß̖Άl˩~ӎТkÒÚ¿hŮҳµÒÚŇ˫YgI (ǁ 1c) @UoHҾӁѢho?ń PSII n 3 ŨϙɒN PSI iγ¡Ô´²¤²iƠϙɒnŨƞhH 7)@YMYʱЏƞYgŮҳµÒÚŇ˫YgIJ

low-energy chlorophyll n°Ð¬¿n͙WoҾӁѢhH (ǁ 1b) @Ň˫WaŮҳµÒÚo?

iƠϙɒhH@UnUio?N PSII i PSIPSII n open/close lğǽYgӦŖlŕӱWinŮҳµÒÚn¸ÐÖ£ißho7)?ń PSII nЃ 50%N PSI iγʱЏƞYg

бK @low-energy chlorophyll n°Ð¬¿

PSI-PSII ҴѲƞĖlğǽYgI~nn?ŀԇɬn͙Wo?ӞɖѤŀ£Â°Òn¼ÚĒЩMn ѻ ͸ M  o PSI-PSII Ҵ Ѳ ƞ Ė l L S ŚˌW (ǁ 1) @ՌϪ̖Ά (~740 nm) xjn

PSI-NPQ ąǣņ̞l~ğǽYgIUiχ͙ W h o k I ~ n n ? Ҿ Ӂ Ѣ (~725 nm) oƱYgI 13,14)@NʣeѲ˅nŀԇɬ̧̞nlow-energy chlorophyll  P700 lϧƞ[°Ð¬Ŗ¿iYgʣdgL?ԊȈΪk PSI-NPQ nнŪ ՌϪ̖Άhoń PSII nЃ 50%N PSI iγʱЏ ĥ ʣ Y g I  i б K    @ low-energyƞYgŮҳµÒÚŇ˫YgI (ǁ 1d) 7)@

chlorophyll n°Ð¬¿n͙Wo?°Ð¬¿iYg

low-energy chlorophyll n°Ð¬¿~Лы̖Άnæ˨KlԅYgoÿɨnҔԶhH@ǁ 1. Лы̖ΆnӞɖѤŀ£Â°Ò (~20ns)
PSII ƒɶæɲN͘ĖϤЉ͠ɒhŮҳµÒÚƕSƔi?ÛȈnυΌhԡѓŎЏƞNҳU ?PSII ƒɶæɲlŮҳµÒÚNӞgΦΔ[@UnӞgΦΔYaŮҳµÒÚΘ˹nÔÔ½ÒѤŀn͍Ԁŕɂѽ˼Ya~nNӞɖѤŀ£Â°Òh?PSII ƒɶæɲγʱŮҳYaHinŮҳµÒÚnŕɂπUiNœ˹@PSII N PSI iЏƞYgŮҳµÒÚĎӣYgIiObS?PSII Θ˹nӞɖѤŀN PSI n͍ԀԱǐ (>720nm) Mѻ͡W@16

ŀƞʈσϡ 28 (1) 2018ho˩~͙Q?740nm ăӎlѤŀ̚Ǧχ[@z

LHCII trimer ƠǠnԄhnµÒÚϗųo 150aɞŀàho?ń PSII nЃ 70%N PSI-PSII ҴѲ¼ϔϙɒMMiѵϟ~gIa} 18)?~

ƞĖɠʈ[ 7)@UnUioՌϪ̖ΆN?PSII

Y larger PSI-PSII megacomplex lLIg PSII i

i PSI nŮҳµÒÚn¸ÐÖ£ißh

PSI nԄl LHCII trimer nȳNʦzdgIi?

~?ŀԇɬnѻ͸M~?PSI-PSII ҴѲƞĖlǦѻ͡Wa 20 ¼ϔnµÒÚϗųґ˗hOQğǽYgIUiχƱYgI15)

OkI 7)@UnЏ˿Mϩвo?PSI core@i PSII core NγʱЏƞY?_nƩ

l LHCII

3՛ՌϪ̖Άn PSI-PSII ҴѲƞĖn̞ә

trimer NЏƞ[?direct-type n PSI-PSII ҴѲƞ ՌϪ̖ΆlLIg PSII i PSI NjnJlЏĖ n ǽ Ǉ  ʴ ̒ Y a 7,15) @ U   n Џ ˿ o ?

ƞYgIMleIgo?̟AkÊ¯ÒNʴƴWbridged-type nǽǇƣȈ[~nhokIN?

gOa@_nJcnÛeN?Ѳ˅n LHCII trimer¡Ô´²¤²nљlLIg PSII i PSI nԄnh̞ʈWȳl PSII i PSI NЏƞ[iIJµÒÚϗųN 50 ¼ϔϙɒhѻ͡WgI@LHCII trimer

UiM[i 7)?ȢkQi~¡Ô´²¤²

o PSII i PSI näˎlЏƞ[ɻүʣea}?

lLIgoµÒÚϗųl˅Ω¼ϔMM

UoƕSŃ[IÊ¯ÒhHda@

iñʀW bridged-type NǤQǽǇ[Ƙнɻ YMY_nɨ?¡Ô´²¤²M 2 ϚԹnoēI@

PSI-PSII Ҵ Ѳ ƞ Ė N Ͻ ѱ W  a @ 1 Ϛ Թ β o

 Direct-type n PSI-PSII ҴѲƞĖnǽǇo?Ͻѱ

690kDa ϙɒn¤h?PSI core i PSII monomer

Ya PSI-PSII ҴѲƞĖnԡǺԷɯӿѽ˼nЏ˿h̞ʈW?20 ¼ϔh PSII i PSI nԄhnŮi~ξεYkI@ǁ 2 ßoÔÚ£ʝĖlYҳµÒÚϗųNҳOgIa (smaller PSI-PSII

a native page ldgϽѱWa¡Ô´²¤Ê¯Ò (bridged-type) hH

16,17)

7)

complex) @UnUio?PSI i PSII nƠǠ²n larger PSI-PSII megacomplex nԡǺԷɯӿŏNγʱЏƞ[©¿n PSI-PSII ҴѲƞĖnǽθnĝhH@ɠΈloǤ˅n¸ÑÚ¡ÎÖN

ǇχƱYgI@2 ϚԹβo 2400kDa ąßnǦHN?_n 7 Ũąßo?PSII dimer  2 en

OWh?PSI i PSII N 1:1 n̸ΌhЏƞY?LHCII

PSI-LHCI N 2 ƽӀȚϖhʦ{ӌ{?_nz

M-trimer i LHCII L-trimer NЏƞYgIa (larger

 LHCII trimer NƔ

PSI-PSII megacomplex)7)(ǁ 2 à)@_Yg?Un larger15)

ǀ|Ê¯Òhґ˗NeQ@UnÊ¯Òo?PSII i PSI nԄnՌ

PSI-PSII megacomplex lLIg~?PSII i PSI nӘkŮҳµÒÚϗų 7)?Ѳ˅n LHCII MԄhnµÒÚϗųo 20 ¼ϔhҳOgIa@

PSI vnՌӘkµÒÚϗų 19-21)ґ˗hO@

ǁ 2. ¡ Ô  ´ ² ¤ ² n larger PSI-PSII
megacomplex nԡǺԷɯӿŏθnĝĪAnϻǺnǣɠo¸ÑÚ¡ÎÖlȕhL ?˅ΩĪidgř}gƠZǣɠn~nNѵeMϙɒbN?ԀӋiρӋnѼɒkjlÛȈnѶŠNѵIbWa@Un·ɰo?Ǔ˳ik̞ә (PSII dimer  2 en
PSI-LHCI NȚϖlʦ{ӌ|) nz l
LHCII trimer NăIa ăMkMda [Uihґ˗NeQ@ßoĪAnϻǺnŏθլào_UlÊ¯Òӷmƞ]a~n@̘ōN PSII monomer?ʎǌN PSI-LHCI?ȡ
ōN minor LHCII?Ǧōo LHCII trimer@17

ŀƞʈσϡ 28 (1) 2018
ѩ 1. ¡Ô´²¤²n larger PSI-PSII
megacomplex nѤŀ͞Ѫn˜Ȉ˅ (77K)

4. ՌϪ̖Άn PSI-PSII ҴѲƞĖņн ՌϪ̖Άn PSI-PSII ҴѲƞĖho?LHCII NƕSƔdaµÒÚo[wg PSI zao PSII lĎӣW?PSI i PSII nԄhŮҳµÒÚNɉǊțƬ (ns)͞ѪnϨÛʈŕ (ns)͞ѪnϨôʈŕ (ns)

PSII

P700

2.29

0.23

2.42

N closed lki?ńµÒÚo PSI l°Ð¬

P700+

1.76

0.19

1.95

6:4 nŨƞhŇ˫W 7)@ɞŀkjl

¿W@PSI oȢkQi~ 2 ϚԹnŀԇɬ̧̞ĵKgL

? _ n J c n Û e o low-energy

chlorophyll M P700+vnµÒÚϗųĀYқӉaÖ«ÖhH 9-12)@¡Ô´²¤²M ĿϋǌЛѢnÖ¿ÒʴĞYgQbWda

ϽѱYa PSI-PSII ҴѲƞĖnÔÔ½ÒѤŀūŔǓƇǠlɬψΚYßTz[@˳σϡo?JSTnɉǊțƬo?P700+Nќϟ[˸Ĉho 20%ąCREST nʾʷƕSa~nh[@ß˓Qk (ѩ 1) @UnUio?ՌϪ̖Άn
PSI-PSII ҴѲƞĖN?P700+ĀYgŮҳµÒ

Received Mar 8, 2018; Accepted Mar 22, 2018; PublishedÚͿlǡʵhOUiχYgI@PsbS

April 30, 2018.

kjl PSII nŮҳµÒÚn͗ƍ̧̞o?
PSI vnԡǺĎӣʖŞY P700+nќϟŭSƎбˉΊM?ԄʱΪl PSI-PSII ҴѲƞĖlLSÖ

1.

Liu, H., Zhang, H., Niedzwiedzki, D.M., Prado, M.,«ÖģӜ[Uilk@ϩвnѵϟ~He, G., Gross, M.L. and Blankenship, R.E. (2013)ho?ɞŀàn¡Ô´²¤²ho?˩Ǧ 7 Ũn

PhycobilisomesÔÔ½ÒN PSI-PSII ҴѲƞĖlЏƞYg PSI

Photosystems in a Megacomplex in Cyanobacteria.

Excitations

to

Both

Science 342, 1104–1107.

lŀԇɬƕSUiNhO 15)@«Ð
2.

±тńĖhѵaiOn PSII i PSI nŮҳµÒ

Yokono, M., Murakami, A., and Akimoto, S. (2011)
Excitation energy transfer between Photosystem IIÚn¸ÐÖ£o?PSII N open n˜o 1.4:1?PSII

and Photosystem I in red algae: Larger amounts of

N closed n˜o 1:2.3 lk 15)@Un 3 īҴK

phycobilisome enhance spillover. Biochim. Biophys.

¸ÐÖ£nǡżo?Ûɒ PSI-PSII ҴѲƞĖNɠ

Acta, Bioenerg. 1807, 847–853.

ʈWp PSII n open/closed lğǽYgμ˜lŖSupply3.

˨M?ԏßnǡųnǦOIŀΓǚlo

Kowalczyk, N., Rappaport, F., Boyen, C., Wollman,
F.-A., Collén, J. and Joliot, P. (2013) Photosynthesis
in Chondrus crispus: The contribution of energy

JdgeShHJ@

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19ŀƞʈσϡ 28 (1) 2018

研究紹介
葉緑体の ndhA pre-mRNA のスプライシングに関与する
ペンタトリコペプチドリピート（PPR）タンパク質§
名古屋大学遺伝子実験施設
伊藤 綾花*、杉田護*

葉緑体 RNA が成熟する過程で、核コードの PPR タンパク質が重要な役割をはたしていることがわかってきた。しかし、植物種によっては数百個もある PPR タンパク質のうち、作用する標的 RNA 分子やその相互作用部位が解明されたものはわずかにすぎない。ここでは、葉緑体 NDH 複合体を構成する
NdhA サブユニットの発現に関与する新規の PPR タンパク質を明らかにしたので、その研究成果を紹介する。

1՛oZ}l34xy[wgn PPR ©Ö¹үoÇ°Ö±Ñ ¡¶¸®Ñҳͥi[љЛĖo?̍Ù

MљЛĖlÖÅÚ°W@¡Ô´²¤²nЊмүioΡkΉфn¶ÈiӨĎǺΦΏ¡

450 Ϛn PPR ©Ö¹үnJc?љЛĖhĸQi

£®ÈʣdgI@љЛĖӨĎǺnΦΏoӀŏʲȈWnońĖn 3 ŕn 1Ֆ154 Ϛ՗7)h?U̵ԓn{k\?Ӏŏɨn RNA ÓÁÒhǤ̟knJçнN˗MlWano 40 ϚxjhH

1)ŞɬƕS @љЛĖnǘƞ?ӀŏɨnŞɬN

@ºÉ­Ñµnń PPR ©Ö¹үnLiSӷѴhHN?UʝJɣвnǦƂo

_ƂŕoљЛĖhĸQ~niʲȈWgI̍¶ÈlÚ±WgI@_nænèѴkӀN 8)?UzhļнNѽ˗Wano\M 4ŏɨŞɬƾǺiYg?ÂÖ©°ÑÂ¿«±ÑϚl[PkI@clpP-rps12 pre-mRNA nʈ;i£¼Ú°ՖPPR՗Ê«Ú½~e©Ö¹үN͎β¿Ð¡ÖloaQ PpPPR_38 9)?pre-tRNA

WgI@PPR Ê«Ú½o 2000 ɊlΦѵWn 5´˲ϦɠʈloaQ PpPPR_67 i 104 10)?La©Ö¹үÊ«Ú½h?35 Ç¶ӵMk?

s pre-tRNAIle (GAU)n£¿Ð¡Ölԅâθ̍ΔΆl˞ӠΪlǽǇ[ 2)@PPR ©Ö¹ү[ PpPPR_4 11)h?_ąǣnљЛĖ PPR ©Öo PPR Ê«Ú½˅ĪG30 ĪЧӏYʣc?ԏ

¹үņнoá˗hH@ɟσϡȍhoӔӨĎß̖ΆhoǦOk©Ö¹ү½ÇÑÚ̞ʈǿΪʏ͌ΖIgºÉ­Ñµn PPR ©Ö

3)

YgI @¡Ô´²¤²iµl 450 Ϛąß?

¹үnЗЪΪķнѽ˼ѦdgIN?ϩвºÉ­Ñµl 100 Ϛxj?ЛыѢԹl 10Ֆċѡ՗oƃ̙σϡiYg?5 Ϛn PPR ©Ö¹

4)

G20 ˅Ϛn PPR ©Ö¹үNǽǇ[ @Uүņнѽ˼lƔzhnσϡl?PPR ©Ö¹үN RNA nѽ˼N˩~Ӝb PpPPR_66 nЏ˿leIgϴƆC-to-U ОԚ?Ŗˌ?ȃȈż?£¿Ð¡Ö?а

lЋĀYaI@kLҊЊleIgoӎˑœ΄ñȈ҄kjnљЛĖ mRNA nӀŏɨŞɬlԅdgnҗˉVƎͼQbWI 12)@

Ѝb@UnJc?˳Ϝho

IUiNǗƧWgIՖǁ՝՗5, 6)@PPR ©
Ö¹үn N ˲lÒµÐϗѦӱŘNH?

2՛ºÉ­Ñµn PPR ©Ö¹үņнѽ˼§

Ϩ 8 ƽˑ˳ŀƞʈǿčɊč Å£©ÚΦѩҭƕҭҗˉ*ӚАĿ %!$ !+'&&'/,"(
*,!+&&'/,"( PpPPR_66 o 578 Ç¶ӵMk?11 Īn

PPR Ê«Ú½ʣeљЛĖȧǇ©Ö¹үh?

20

ŀƞʈσϡ 28 (1) 2018

ǁ 1. љ Л Ė nӀ ŏ ɨ ¿ Ô ¥ £  z i}a̜ɸǁљЛĖ DNA MӀŏWa RNA ŕǺo̟Ak¿Ô¥¡ÖƕSgʈ; RNAlk@UnӀŏɨn RNA Şɬl̍Ú±nÂÖ©°ÑÂ¿«±Ñ¼Ú°ՖPPR՗©Ö¹ү½ÇÑÚNԅdg
I@Sugita et al. (2016) 6) ʿǡ@

PpPPR_66 ÃÊÔN¡Ô´²¤²?µ?° љЛĖloÖ°ÔÖ~e 12 Ϛn©Ö¹ÊÔ¡kjl~ǽǇ[N_ņнoá˗үӨĎǺi 6 Ϛn tRNA ӨĎǺNǽǇ[@£¿8)

hHda @_Uh PpPPR_66 ņнѽ˼ѦkÐ¡ÖΡɆn˫͹ϴġlҕwa}?ÖJa}?ӨĎǺÚ±Աǐ_dQ°ÔÖʦ|Ţɨn§ÖlĒЩ[¿Ð·ÔÆ¡ÖгɻӨĎǺ¥¬°iЩʵYaӨĎǺτÆÚΖIg RT-PCR ѽ˼Ѧkda@_nЏ˿?

Ǟ̌ՖKO ̌՗ęœYa@YMY˯ɦlƒYg?

KO ̌ho ndhA n£¿Ð¡ÖNńQLUd

ɫa KO ̌nƋЀĖnΔкLsŀƞʈн

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PpPPR_66 nńԀ cDNA  KO ̌lȠŃYgɫaδѰ̌ho ndhA n£¿Ð¡ÖNӸΔǌÓÁÒzhƽɭYaUiM?ndhA n£¿Ðѩ 1. ºÉ­ÑµƋЀĖÔ³ÚnÔÔ½ÒѤŀ¹ÐÉÚ©Ú
¹ÐÉÚ©Ú

ӸΔ̌

KO ̌ Δ66-2

KO ̌ Δ66-3

fPSII
qP

0.448 ± 0.019

0.413 ± 0.013

0.428± 0.022

0.675 ± 0.057

0.628 ± 0.033

0.630 ± 0.025

Fv/Fm

0.825 ± 0.050

0.828 ± 0.004

0.828 ± 0.008

NPQ

1.453 ± 0.294

1.488 ± 0.340

1.278 ± 0.386

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͗ŀ?Fv /Fm ：PS:n˩ǦÔÔ½ÒѤŀӹǺƓΌ?
NPQ：ԥŀżǿ͗ŀ@͡ȈѭЩեFluorCam 800MFՖPhoton System Insruments ωѱ՗͡Ȉ˸Ĉե̖KДPɨ 3 ӛβnºÉ­ÑµƋЀĖÔ³Ú 10 ŕԄˣʍhԯɶW]aɨh͡ȈYa@FooɜIұы͡Ȉŀ?Fm oԿƭ¹Ò£ŀՖ1400 μmol
photons m–2 s–1 for 0.8 s՗_`ͼȞYg͡ȈYa@Կƭ¹Ò£ŀͼȞɨlұыŮҳŀՖ96 μmol photons m–2 s–1 ՗՝ŕԄͼȞYa@ǁ 2. ՖA՗PpPPR_66 ©Ö¹үnÊ«Ú½̞ʈ@љЛĖϗѦloaQ°ÐÖ¢¬°ӱŘՖTP՗i
11 Īn PPR Ê«Ú½Mk@ՖB՗ӸΔ̌ՖWT՗
i PpPPR_66 KO ̌ՖΔ66-2 i Δ66-3՗ BCD ǒǈՖ˩ȡǒǈ՗i BCDAT ǒǈՖӲςӵ¢ÖÊ³È͛ŬYa BCD ǒǈ՗h 3 ӛԄΔкW]aƋ
ЀĖÔ³Ú@£ÚÒ¸Úo 1 cm ѩ[@Ito etal. (2018) 12) ʿǡ@21

ŀƞʈσϡ 28 (1) 2018
kIl~MM\?_nѩΏǌNÛѵӸΔ̌nѩΏǌiđgIaUiM?љЛĖn NDH ͐ɻlɢԫNœgIƘнɻNбKa@_Uh?
PpPPR_66 KO ̌lLSљЛĖ NDH ͐ɻҕwUilY?ûӰǦǿnՑŌŜ͇˂ʬl͡ȈğԴYa@Mini-PAM (WALZ)ѭЩĜdaÔ
Ô½ÒѤŀɞɒn͡ȈǦǿԋΔnŬѡЭȌ̹lѦdgIabIaiU?ñʀӗљЛĖ

NDH ͐ɻѩ[Ůҳŀ½ՖAL off՗γɨlΏÔÔ½ÒѤŀnÛӡΪß˕N KO ̌hoѻȗWkMdaՖǁաA՗@љЛĖ NDH ͐ɻ N ̗ œ W  k M d ai I J U i o? љ Л Ė
ǁ 3. ndhA n RT-PCR ѽ˼
895 bp ˌ΃oÖ°ÔÖN£¿Ð¡ÖhԌM
aԀWlδɟY?1585 bp ˌ΃oÖ°ÔÖƤ|ԀWlδɟ[@PpPPR_66 KO ̌Ֆ066-2 i0
66-3՗ho 1585 bp ˌ΃N̗œWaN?895bp ˌ΃o̗œWkMda@ÓÚÖΠƛ 1ՙ2?3 o PCRƒɶnÒ˅ 26, 30, 34 _`χ[@Ito etal. (2018) 12) ʿǡ@

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NѦa@ǁաB lχYaJl?¡°ÔÈ
b6/f ѲƞĖn¡°ÔÈ f ©Ö¹үՖCytf՗?
H+-ATPase n β ¾Í³¬°ՖAtpB՗?PSII nƒɶæɲ D1 ©Ö¹үՖPsbA՗oӸΔ̌i KO ̌¡Ö̪ʸN PpPPR_66 ӨĎǺn̪Ǫl~hxyƠÓÁÒќϟYgIanlȚYg?љЛĖnhHUiNυҌWa 12)@

NDH ѲƞĖn̞ʈ¾Í³¬°hH PnsB1 i
NdhM NxijќϟYgIkMda@UnUi

3՛PpPPR_66 KO ̌ho NDH ѲƞĖNɠʈW

o?PpPPR_66 KO ̌hoљЛĖ NDH ѲƞĖNɠkI

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ŀƞʈσϡ 28 (1) 2018
ՖETR՗?ÔÔ½ÒѤŀnԥŀżǿΪ͗ŀՖNPQ՗?ŀżǿΪ͗ŀՖqP՗nI\~NӸΔ
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ndhA n£¿Ð¡ÖNzdaQҳdgIkIUiN
RT-PCR ѽ˼i¶ÚÖѽ˼hѻȗWa 12)@ąßnЏ˿o?PpPPR_66 i AtPPR66L ŅнΪlδƠk PPR ©Ö¹үhHUiχYgI@ љЛĖ ndhA ӨĎǺnÖ°ÔÖo?ЛыѢԹn ndhA ӨĎǺloǽǇYkIN?ҾӁѢԹiԏß ̖ Ά n ndhA Ө Ď Ǻ l ǽ Ǉ [  U i M  ?
PPR66L ~Un̖Άl˞ӠΪlǽǇ[Ui
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a o ɢ ԫ [  © Ö ¹  ү i Y g ?

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Δ66-3՗
 5 ŕԄ͡ȈŀՖAL, 50 µmol photons m -2 s-1՗ͼȞY?͡Ȉŀ½γɨnÔÔ½ÒѤŀnÛӡΪß˕ѻȗYa@PAM ÔÔ½Ò½Ò
ÔÉ°ÑÚΖIgÊ³©ÚYa@(B) KO ̌ho?
NDH ѲƞĖn̞ʈ¾Í³¬°hH PnsB1 i
NdhM N̗œWkMda@Ito et al. (2018) 12) ʿǡ@

CHLOROPLAST RNA SPLICING 2 (CRS2) ?
CRS2-ASSOCIATED FACTOR 1 and 2 (CAF1 i
CAF2)?CHLOROPLAST RNA SPLICING AND
RIBOSOME

MATURATION(CRM)FAMILYMEMBER 2 (CFM2)N°ÊÔ¡¡Ô´²¤²hπgI 14)@Un©Ö¹үo?
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4. ¡Ô´²¤²n AtPPR66L ~љЛĖ ndhA

UlȚYg?PpPPR_66 o ndhA £¿Ð¡Ön£¿Ð¡ÖlԅâYgIl·ΡΪlĸQƾǺiѿKbJ@

 PpPPR_66 nÒ§Ôoԏß̖ΆlɎQǽǇY g I  @ ¡ Ô  ´ ² ¤ ² n At2g35130 o

5՛PpPPR_66 o ndhA Ö°ÔÖlЏƞ[PpPPR_66 iƠZ 11 Īn PPR Ê«Ú½Mk?

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PpPPR_66 nÇ¶ӵӱŘio 44%n identityŘ · Ρ Ϊ l Џ ƞ [  U i N π  g I  5) @Ֆ81%n similarity՗~e@˳Ϝho At2g35130

PpPPR_66 N ndhA Ö°ÔÖn£¿Ð¡ÖġȊΪl AtPPR66L iƫuN?AtPPR66L ~lԅâYgIUiM?ndhA Ö°ÔÖlЏ

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At2g35130 n T-DNA ʨŃǡΡ̌ՖSALK_043507;MkѲԜkՌ̫̞әii SALK_065137՗nÔÔ½ÒѤŀӹǺƓΌ

PpPPR_66 N ndhA Ö°ÔÖlЏƞ[MjJiљЛĖ NDH ͐ɻûӰǦǿnȮ˳ȅƇǠlҕM̗҅[a}?ЍʵK PpPPR_66 ©Ö¹ү

wgIabIa@_nЏ˿?ŀƞʈԡǺĎӣӘɒ

ՖrPPR66՗ΖIa?in vitro n RNA ЏƞȋՊ

23

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IlUiNбK@ÿɨ˗Ml[wOҔԶhH@ ˳ϜhoѾkMdaN?ºÉ­Ñµlo PpPPR_66 n¹ÐÔiYg PpPPR_72 NǽǇ[@äвnδƠɻoՌQ?Ç¶ӵӱŘh 73%
identity H?11 Īn PPR Ê«Ú½n RNA ҌҜÚ±~xyƠZhH@UnUiM?äвoƠZ̧н~eiɟřñʀWa@YMY?˳σϡnЏ˿M PpPPR_72 o PpPPR_66 io̧нΪlΡkdgIƘнɻNχƱWa@ĨǠĨðz

ǁ 6. ºÉ­ÑµnљЛĖ ndhA Ö°ÔÖnʲȈô̫̞әi PpPPR_66 nЏƞԱǐՖͶыnӭŕ՗Ö°ÔÖn±ÉÖ9ՖDI՗G;ՖD;՗i?
Ö°ÔÖŕǺŌnӷѴkδõęΖӭĒՖIBS1,
IBS2, EBS1, EBS2, 1?1´?3?3´՗ǁlχ[@
Ito et al. (2018) 12) ʿǡ@hlo PpPPR_72 ņн˗MlYaI@˳ϜҒzaΫ̟MnVʁѵV˂ʬIab
SpɋIh[@қӉ PpPPR_66 ņнѽ˼o?ûӰǦǿnՑŌŜ͇ɟσϡȍnÛ͵ΒϞNѦda@_nЏ˿?rPPR66

˂ʬiՑŌσϡȍnȮ˳ȅƇǠ?ŬѡЭȌƇǠko ndhA Ö°ÔÖn 5´˲ϦԱǐlЏƞ[UislơƗȪǦǿӨĎǺȋՊˏ҃n˴ΗƀɾǺNŚ˗Ya 12)@PPR Ê«Ú½n 5 Πβi 35 ΠβƇǠ?ơƗȪǦǿ°ÐÖ£½ÚÆ®¾ΔƬŕ

nÇ¶ӵnЍ{ƞ]ՖRNA ҌҜÚ±՗hǺσϡʍՖWPI-ITbM՗nÛ͵ΒϞƇǠinŇƠ

Џƞ[ RNA ǙǓHϙɒñ͡[UiNh

σϡiYgѦa~nh[@UnˎAl͙

O 15, 16)@Un RNA ҌҜÚ±ʏʮMlY

QʃқΚYßTz[@za?ʘĖʴĞYgIa

gЏƞԱǐҕwaN?_nJkӱŘoѵIb

bIaéǕɱ˂ʬՖ˺ûȺ̙Ǧǿ՗?΅ӸƩ˂ʬWkMda@UnUio PpPPR_66 N RNA nՖ˺ŽǦǿ՗?ӥѡŤƇǠՖûӰǦǿ՗lʃқIÛ̫ӱŘhokQ?ÒÚ¿:Ö°ÔÖŌnѲaYz[@˳σϡoˑ˳ǿѧʧчčϓǿσϡҨѰԜkՌ̫̞әҌҜYgIƘнɻNбKŭӺՖ15K14917, 17K08195՗nʾʷƕSa~n

@

h[@

6՛ L l љЛĖ ndhA n£¿Ð¡ÖlԅˍYI

Received February 9, 2018; Accepted February 15, 2018;ƾǺiYgºÉ­Ñµn PpPPR_66 i¡ÔPublished April 30, 2018

´²¤²n AtPPR66L ѵIbYa@YMY?
¡Ô´²¤²n AtPPR66L ºÉ­ÑµƎбˉΊn PpPPR_66 KO ̌lȠŃYgɫaδѰ̌1.

Sugita, M. and Sugiura, M. (1996) Posttranscriptional
regulation of plastid gene expression in higher plants.

N?ñʀlƒYg ndhA £¿Ð¡ÖƽɭY
kIiIJǫǱkЏ˿ѻȗYa 12)@UnUio?
AtPPR66L o¡Ô´²¤²n ndhA £¿Ð¡

Plant Mol. Biol. 32, 315–326.
2.

Small, I.D. and Peeters, N. (2000) The PPR motif – a
TPR-related motif prevalent in plant organellarÖloĸQN?ºÉ­Ñµn ndhA £¿proteins. Trends Biochem. Sci. 25, 46–47.

Ð¡ÖloĸMkIUiχƱYgI@U

3.

Lurin, C., Andrés, C., Aubourg, S., Bellaoui, M.,nΑΘiYg?Ϛ·ΡΪk ndhA Ö°ÔÖ̞әBitton, F., Bruyère, C., Caboche, M., Debast, C.,nӤI?ĀǇ[Ăn£¿Ð¡ÖƾǺnӤ

Gualberto, J., Hoffmann, B., Lecharny, A., Le Ret,
M., Martin-Magniette, ML., Mireau, H., Peeters, N.,
Renou, J-P., Szurek, B., Taconnat, L. and Small, I.

24

ŀƞʈσϡ 28 (1) 2018(2004)

Genome-wide

pentatricopeptide

4.

analysisrepeatofproteins

Arabidopsis
reveals

11.

their

Sugita, M. (2016) P-class pentatricopeptide repeat

essential role in organelle biogenesis. Plant Cell 16,

protein PTSF1 is required for splicing of the plastid

2089–2103.

pre-tRNAIle in Physcomitrella patens. Plant J. 86,

Tourasse, N. J., Choquet, U. and Vallon, O. (2013)

493–503.
12.

PPR protein of green algae. RNA Biol. 10, 1526–
5.

6.

H., Shikanai, T. and Sugita, M. (2018) An

Barkan, A. and Small, I. (2014) Pentatricopeptide

evolutionarily

repeat proteins in plants. Annu. Rev. Plant Biol. 65,

pentatricopeptide repeat protein is required to splice415–442.

theSugita, M., Uchiyama, H., and Ichinose, M. (2016)

Physcomitrella patens and Arabidopsis thaliana.

transcript

in

the

moss

Shikanai, T. (2016) Chloroplast NDH: A different
enzyme with a structure similar to that of respiratory

Colcombet, J., Lopez-Obando, M., Heurtevin, L.,

NADH dehydrogenase. Biochim. Biophys. Acta 1857,
1015–1022.
14.

(2013) Systematic study of subcellular localization of

Schmitz-Linneweber, C., Lampe, M-K., Sultan, L.D.

Arabidopsis PPR proteins confirms a massive

andtargeting to organelles. RNA Biol. 10, 1557–1575.

maturases: A window into the evolution of theOstersetzer-Biran,O.

(2015)

Organellar

spliceosome. Biochim. Biophys. Acta 1847, 798–808.

Sugita, M., Ichinose, M., Ide, M. and Sugita, C.
15.

(2013) Architecture of the PPR gene family in the

Barkan, A., Rojas, M., Fujii, S., Yap, A., Chong, Y.S.,

moss Physcomitrella patens. RNA Biol. 10, 1439–

Bond, C.S. and Small, I. (2012) A combinatorial

1445.

amino

Hattori, M., Miyake, H. and Sugita, M. (2007) A

pentatricopeptide repeat proteins. PLoS Genet. 8,

pentatricopeptide repeat protein is required for RNA

e1002910.
16.

processing of clpP pre-mRNA in moss chloroplasts. J.

acid

code

for

RNA

recognitionbyYagi, Y., Hayashi, S., Kobayashi, K., Hirayama, T.
and Nakamura, T. (2013) Elucidation of the RNA

Biol. Chem. 282, 10773–10782.
10.

ndhA

P-subfamily

27, 41–44.
Bernard, C., Martin, K., Berthomé, R., Lurin, C.

9.

plastidconserved

Plant J. DOI: 10.1111/tpj.13884.
13.

for plastid RNA metabolism. Endocytobiosis Cell Res.

8.

Ito, A., Sugita, C., Ichinose, M., Kato, Y., Yamamoto,

1542.

RRM and PPR proteins network as a global regulator

7.

Goto, S., Kawaguchi, Y., Sugita, C., Ichinose, M. and

Sugita, C., Komura, Y., Tanaka, K., Kometani, K.,

recognition code for pentatricopeptide repeat proteins

Satoh,

involved in organelle RNA editing in plants. PLoS

H.

and

Sugita,M.

(2014)

Molecular

ONE 8, e57286

characterization of three PRORP proteins in the moss
Physcomitrella patens: nuclear PRORP protein is notessential for moss viability. PLoS ONE 9, e108962.

A pentatricopeptide repeat (PPR) protein is involved in splicing of chloroplast ndhA pre-mRNA
Ayaka Ito*, Mamoru Sugita*
Center for Gene Research, Nagoya University25

ŀƞʈσϡ 28 (1) 2018

解説特集

変動する光量への光合成機能の調節

Editor:古本強（龍谷大学 農学部）

序文Ɨ˳ ɞՖՕҟǦ՗

27

解説 ΓǚǡųlȚ[̻ǻiŀƞʈnɶϭ˰˶ өɃ?șȳ ÛӬ?οȂ шՖ˺ûǦ՗29

解説 Ó±¬£ǓαiYaљЛĖņнВɬµ¬°ÕÚƟΗƶüՖ˺ûȺ̙Ǧ՗39ҟƖɋЫՖԅѳǿԋǦ՗

51

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ŀƞʈσϡ 28 (1) 2018

解説特集
序文

‡

龍谷大学

農学部

植物生命科学科古本

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ŀƞʈσϡ 28 (1) 2018

解説
環境変動に対する気孔と光合成の応答‡

東京大学

大学院理学系研究科

生物科学専攻植物生態学研究室

木村 遼希*、寺島 一郎、矢守航

気孔は一対の孔辺細胞からなり、植物体内と外界との間のガス交換バルブの役割を担う。植物はこの気孔から二酸化炭素 (CO2)を体内に取り込むと同時に、体内の水分を蒸散によって失っている。気孔は様々な環境要因に応答して、その開度を調節している。例えば、乾燥時には気孔は閉じ、水分の損失を防ぐ。一方、光合成を活発に行えるような条件では、気孔は開き、CO2 を取り込む。気孔は、陸上植物にとって、光合成の基質である CO2 のほぼ唯一の取り込み口であるため、気孔と光合成との関係は古くから研究されてきた。本稿では、これまでに明らかにされてきた気孔開閉の分子メカニズムに関する知見と、様々な環境条件下の気孔動態と光合成応答との関係性を最新の知見を含め紹介する。
1. oZ}lÑÚÒӵhH@UnÃ£ÃÑÚÒӵ §̖ΆNԏßlӜœ[ԖlγԦYaǦOkƵ

ĄқY?ͷЉƽƓ[ŀƫƥЎҺnՉųlo

Զo?IMl̼ŕĥʣYee?CO2 ƥƓ[ATP ӪĽŪNɳѴhH@YaNdg?Ò»M?iIJUihH@ԏß̖Άo?Un¢ÓÖ£ƩӋn CO2 ʹɒoŀƞʈnůΌlԥɆlǦÆ̎˳Ϊlѽ́[UiohOkMdaN?_

OkɢԫâKgI@nǲƄ͸iYg̻ǻ~elхda@̻ǻo?ѩ љЛĖN CO2 vnżǿҲɻχ[UioƗQ

άnÛȚnǻӋЊмl

̞ʈW?љŌiǣΝMχƱWgI 1)@za?љжЊмlLIg?

in£úʵ¸Ò¾nɣŨ˿a[@Un̻ǻNљЛĖNЊмтnЊмԄԕlԦ[ӭŕlԚƞ

̟AkΓǚ˸ĈlɶϭYgӦŖlԃɒҕϰ[[Ui 2,3)?ƫƥŀƫƥhˀœW CO2

Uih?ԏß̖Άo̼ŕnʸǪLWKee?

ŎǂȈhOJl?ЊмԄԕ[OԄkQǎŀƞʈlɳѴk CO2 љŌlƔ}eQ[ӱЩiUi 4)~ǗƧWgI@U

ӌ|UiNhO@oљЛĖl CO2  ̻ǻMљŌviӜŃYa CO2 o?Ò»£lilɣϥeiбK@̻ǻMљŌlŃda

dgÒ»ÖÒlƔӌzaɨ?«ÐCO2 o?љЛĖzhʡ˃[@̻ǻсMљЛĖ

±тhƞʈWa ATP ӪĽŪldg°zhnʡ˃nY[WљжÖª©Ö£iÑÚ£ÑÖӵik?Ͼ¯Ö¿ÖnƞʈlΖ

ƫs?љжÖª©Ö£o?ï΁?CO2 ʹɒ?

I@YMY?Ò»£n CO2 lȚ[Ѻƭ͠ɒϪlɶϭYgǡż[UiNπgI@ɻo̚}gēI@ŬKg?Ò»£o?CO2/O2

CO2 ɶϭnϙɒo̖ΆϚldgΡk 5,6) N?

ʹɒ̸lɶZgÒÄ¡ÓÚ¡ÎÖƒɶiϧQɶϭ[ϚlLIgo?љŌ CO2 ʹɒNēI

òΪlҳU¡µÚ¡ÎÖƒɶ~ѾǷ[iљжÖª©Ö£oß˕Y?ӔlљŌ CO2

@_nΕΆnÛeN?Ò»ÖÒæn°ʹɒNß˕[iљжÖª©Ö£oēà[ÑÚ£ÑÖӵ§ÉÐÚ¦Ԉȏ[Ã£ÃķƢNH 6-8)@UnUio?љŌ CO2 ʹɒNůΌΪlĞБ[UēIΈ͉àh?љЛĖvn CO2 ĞБůΌNՌ}
‡gIUiχYgI@UnJl?̖Άl

ѽґ·ԚBǡų[ŀӹvnŀƞʈ̧нnҕϰCidgљЛĖ?_YgÒ»£l CO2 ůΌQ*ӚАĿ E-mail: hkimura@bs.s.u-tokyo.ac.jp

29

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QπgI@½°°Ô¼ÖNԣыŀƕȑ

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 ŀƞʈi̻ǻԃɒnԅĢleIgoƗQM²¤²ho?11 ϚԹnЊмтǌ H -ATPase nJ

σϡWgI@·l?ÛȈnɞWnȈɆŀn~c?AHA1 (Arabidopsis H+-ATPase 1)Nԣыŀ¡ihҊЊkσϡNkWgOa@˩ӎo?

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ɶϭlԅ[σϡǗƧ~ǛKgOgL

ƢO K+«ËµÒn͐ɻżЎg?ƥ̼NҳU?ȋԖlӸǣlΔк[̖ΆnȋʄlӎIπѵNɫ?

уǆNß˕Yg̻ǻNԃQ@

Ǵ}gI@˳Ϝho?Uzhl˗Mlkd
gOa̻ǻnΓǚɶϭnŕǺÉ³¤ÈleI2՗CO2 ɶϭgѽґ[ii~l?̻ǻiŀƞʈinԅĢle Ǔ˳Ϊlo?CO2 ʹɒNēSp̻ǻoԃO?

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ŀƞʈσϡ 28 (1) 2018lɶϭYgInM˗Ml[UiNɳ԰

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Stomatal and photosynthetic responses to fluctuations of various environmental factors

Haruki Kimura*, Ichiro Terashima, Wataru Yamori
Department of Biological Sciences, Graduate School of Science, The University of Tokyo38

ŀƞʈσϡ 28 (1) 2018

解説
レドックスを基盤とした葉緑体の機能統御ネットワーク‡

東京工業大学

化学生命科学研究所
吉田 啓亮*

レドックス制御は、標的タンパク質のチオール基の酸化還元状態を制御することにより、そのタンパク質の酵素活性を調節する翻訳後修飾の１つである。近年、葉緑体のレドックス制御システムは、チ
オレドキシンファミリーをはじめとする数多くの還元力伝達タンパク質を巧みに使い分け、多岐にわたる葉緑体機能を精密に操っているという驚くべき実態が見えてきた。その制御ネットワークの全貌解明の重要性については国内外を通じて論を俟たず、今もなお活発な研究の対象となっている。本稿
では、最近の研究成果を中心にこれまで得られてきた知見をまとめ、そこから明らかになってきたレドックス制御ネットワークの分子基盤や生理意義を概説する。
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50ŀƞʈσϡ 28 (1) 2018

解説
Flaveria 属における C4 型光合成の進化過程‡

関西学院大学理工学部

生命科学科
谷口 幸美*

C4 型光合成は大気中の湿度や CO2 濃度の低下といった環境の変化に適応して植物が獲得した収斂進化の一形態であり、C3 型光合成を行う祖先種から C2 型（C3-C4 中間型）を経て C4 型光合成を行う種が発生した可能性が示唆されている。C4 型光合成を行う植物種の多くは葉肉細胞と維管束鞘細胞の二細胞間で分業し光合成を行うが、このためには二種類の細胞の位置を正しく制御し、かつ二細胞間の連携をとることが重要となる。本稿では C3 種、C4 種に加えて C2 種（C3-C4 中間種）、C4-like 種が多数現存するキク科の Flaveria 属植物をモデルとして、段階的に進んだと考えられる C4 型光合成の進化過
程やその過程で起こる形態的変化、遺伝子発現の変化について紹介する。
1. oZ}lΏԱǐNЖϯ˷ԩЊмvԊȈWgL

 §C3 ǌŀƞʈho CO2 nǂȈM¡ÎϾ¯ÖҺl¿Ökjnͷ̼żΆƞʈzhnÛӚnĄқƒɶ

gIa}?RuBisCO n¡²Ú¦ƒɶoèlљжЊмŌhѦ@UlȚYg C4NʖŞW?Ռ͠ï΁kjљŌn CO2 ʹɒNēǌŀƞʈnJcnǦӭŕƈ}ôЊмɻn C4

à[˸Ĉàh~ՌIŀƞʈůΌχ[@°Êǌŀƞʈho?ôϚԹnЊмNƄĸYg C4 ƽҺÔ¡ՖZea mays՗°»ՖSaccharumՉųYÛˎnЊмl CO2 ʹУ[ՖÛЊмŌ

officinarum՗lĄѩW C4 ǌŀƞʈѦJęΆh CO2 ȧʍΪlʹУ[~n~Ȣ˅kNǽ

oՌ͠ï΁˸Ĉàh~˔ήkΔкχYΔΕ

ǇYgI

47, 48)

՗@ôЊмɻn C4 ǌŀƞʈho?

?C4 ƽЖϯ˷ԩЊмŌn CO2 ŕǆNՌQĥaɻNԥɆlՌIUiNπgIN?_oU̻ǻMŃda CO2 oӷͷӵÖlǡʵW

n C4 ǌŀƞʈlΘ˹[ɠүhH@aɨ?љжЊмlLIgͷЉ˅N 4 enÔ ǈΐßl C4 ǌŀƞʈƸ|ԏß̖Άo 19 ϓӳӵlǂȈW?£¹ÐÖӵzaoÑÖӵ

7500 ϚxjNǽǇ[UiNυҌWgL

viǡʵWaɨlЖϯ˷ԩЊмviӈӓW

Uo 62 ЁВՖƆǺљԹ 26 ЁВ?ƑǺљԹ

@Жϯ˷ԩЊмho?рͷӵӴЉnĸOl36 ЁВ՗lŕԹW 38)@za?C4 ƽҺlԅâˀ œ W  a CO2 N Ж ϯ ˷ ԩ Њ м l ȧ Ǉ [ [ӨĎǺЬo C3 Ϛn¶Èl~Ú±Wgribulose-1,

carboxylase/oxygenase

IUiNπgI 42)@UnJl?C3 Ϛ(RuBisCO)ldgŎǂȈWͷ̼żΆƞʈzhn¶Èl C4 ǌŀƞʈlɳѴkӨĎǺNǽǇY

nĄқƒɶNѦ@рͷӵƒɶhΔZa¼Ò

gL»ÖӵՖɨӐ[ NAD-malic enzyme ǌhoÐvnӜżNҳUdaiбKUiM?C4³Ö՗Ã£Ã¶ÚÒ¼Ò»ÖӵoљжЊмlǌŀƞʈoƓˈӜżɠүhH

ʌWŎs˩řn CO2 ǂȈƒɶnǓүikՖC4~i~dgIӨĎǺnΦΏkjǡżW]

ƽҺ՗@C4 ǌŀƞʈѦJϚho RuBisCO nΦ

Uih C4 żNəOҳUWaiбKgI

5-bisphosphate?

?MeǤQnΉϥYaЁВlLIg C4 ǌ
C3 ǌ̖ΆN~i37)@za?C3 ǌi C4 ǌnæԄΪkŀƞʈ·ɻ‡χ[ C2 ǌՖC3-C4 æԄǌ՗lŕԹW̖ΆN 21

ѽґ·ԚBǡų[ŀӹvnŀƞʈ̧нnҕϰCЁВǽǇY?_nJc 14 ЁВo C4 ǌϚiŇӗn*ӚАĿ E-mail: y-taniguchi@kwansei.ac.jp

51

ŀƞʈσϡ 28 (1) 2018
ϋĿ~eUiNǗƧWgIUiM 39)?

Ֆǁ 1՗@ÐÖ­̞әoЖϯ˷~YQoҦ̼Ѝ

C4 ǌŀƞʈo C3 ǌMÛҷԻslΦΔYanhЦnƩǀlӱЩWǦOQзǦżYaЊмՖ

okQ C2 ǌՖC3-C4 æԄǌ՗ЎgΦΔYaiбÐÖ­Њм՗i_nǣĲlӱЩWљжЊмN

KgI@ϓ Flaveria Ȭ̖Άo C4 ϚNэŐՖÑÚ£?±­Ҏh Kranz՗nJl{KƤzЁВnæho̸ӃΪӎɊՖЃ 300 ÜɊŢUiMơăSa@C4 ǌŀƞʈѦJϚ

MΏǇlMSg՗l C4 żNҳUdaiWho CO2 o˩řlљжЊмhƠżW?ÐÖ­38)?ƠȬŌl C3 Ϛ C4 ϚbShkQ 9ЊмlʹУW@˳ϜhЋĀ[ Flaveria Ȭen C2 ϚՖC3-C4 æԄϚ՗?3 en C4-like ϚƤ°ÊÔ¡kj?ǤQn C4 ϚhoÐÖ­Њ

ȬhH
hI

23)

мlδɟ[noЖϯ˷ԩЊмhHՖÛӭÉ£@Flaveria Ȭ̖ΆnҳͥiŕԹleI29)l҂°ÈԩЖϯ˷n̄ЍЦNÐÖ­Њмlδɟ

ӄWgInhƎбlYgԬOaI@Flaveria[Ϛ~ǽǇYgI 5,8)՗@za?IQeMnȬ̖ΆlԅYgŀƞʈ̟ɘΔΑǿΪkπѵN

C4 ϚhoљжЊмiЖϯ˷ԩЊмnԄl~JÛ̸ӃΪǤIUiM?˳ϜhoUÊ¯ÒiȭЊмȭNǽǇ[~n~HN 8,9)?Ǔ˳Ϊl

Yg C4 ǌŀƞʈnӜżӡϙi_nӡϙhҳU

o˩řl CO2  C4 ¢ÒÄÖӵviǂȈ[ЊɠʄΪǡżӨĎǺΦΏnǡżleIgЋĀ[мi_рͷӵYg CO2 ʹУ[ЊмNŕM@gIiUl·ɰNH@C4 ϚnЖϯ˷ԩgo˳ϜhoҊӐYkIN?њвnКґЊмoљжЊмl̸wgǦOQ?љЛĖNќϟY
2. C4 ǌŀƞʈlLS C4 ƽҺiљŌ̞ә

gIN?љЛĖąǣnЊмȡƼȇn˅¤ C4 ƽҺlo?Жϯ˷ԩЊмḩн[рͷӵЊмŌhnĒЩl~ǦOkӤINH@Жϯ˷ӴЉnϚԹldgSa 3 en¾©ԩЊмlLSЊмȡƼȇn·ɰoŜΖYgI¿NH@љЛĖlȧǇ[ NADP-malic enzyme

 C4 ƽҺn¾©¿i~ȔʱlԅӚYgL

ՖNADP-ME՗ldgÑÖӵNрͷӵW

ĝKp NADP-ME ǌhoÐÖ­ЊмnљЛĖNADP-ME ǌ ? Ç °  Ö ± Ñ  l ȧ Ǉ [ lLS PSII N͞ȢYgIN NAD-ME ǌho

NAD-malic enzymeՖNAD-ME՗ldgÑÖӵ_ n  J k U i o ѵ   k I 8,9,12,15) @ z a ?

NрͷӵW NAD-ME ǌ?°¨Òn PEP

NAD-ME ǌhoрͷӵoÇ°Ö±ÑhѦ

carboxykinaseՖPEP-CK՗ldgÔӳӵ

N?NADP-ME ǌhoљЛĖhѦ 14)

NрͷӵW PEPCK ǌhH 18)@C4 ǌ̖Άna}l?NAD-ME ǌnЖϯ˷ԩЊмnÇ°Ö

Un¾©¿vnŕԹo?èlĸQрͷӵ±ÑoӎТn NADP-ME ǌnϚi̸ӃYgǦӴЉldgŕԹWgL?Ѳ˅nЎҺěΖ

OQ˅~ǤQkdgI 8,19,32,50)@C4 ϚnљжЊ

YgI̖ΆϚ~ǽǇ[ 13,25,49,51)@za?ЁВ

мhoӎТn C3 Ϛi̸ӃYgљЛĖn˅NȢkŕԹnßhƠÛЁВlȬ[ C4 ϚlѲ˅n¾

Q?C3 ϚnљЛĖNљжЊмnƩТӭ¸Ú©¿NǽǇ[ĝ~ǗƧWgL[JlǽǇ[nlȚYg C4 Ϛho_nՖPanicoideae lo NADP-ME ǌn Setaria Ȭ?

JkӱЩoѵkI 44, 45)@UnљжЊмlLS

NAD-ME ǌ n Panicum Ȭ ? PEP-CK ǌ nљЛĖn͞ȢiӱЩnǡżldg?°¨

6)?

Megathyrsus ȬNǽǇYgI ՗?C4 ̖ΆNӜÒlǽǇ[ PEP carboxyraseՖPEPC՗N CO2

żnӡϙhI\n C4 ƽҺn¾©¿ŜΖ

ՖHCO3-՗[JlkdanMo?Mk

½Ó¡¾Òh

gI

ʪʩY[QkdgIiбK

39)@

HJhH@ C4 ǌŀƞʈѦJ̖Άnљlo C4 ƽҺn¾

3. C3 ǌM C2 ǌՖC3-C4 æԄǌ՗v©¿l\ÐÖ­̞әipѽţǿ ŢӐYaiL5)ԻslҳUdanhokQ?C2 ǌՖC3-C4 æԄǌ՗

Ϊ·ɰNH ՖÛЊмŌh CO2 ȧʍΪlʹУ[~nloÐÖ­̞әoѵkI

47,48)

C3 ǌM C4 ǌvnӜżoÛҷЎgIiñʀWgI@C2 ǌՖC3-C4 æԄ՗52ŀƞʈσϡ 28 (1) 2018
ǌ՗nŀƞʈho C4 ƽҺhokQŀƫƥld[Џ˿?C2 ϚՖC3-C4 æԄϚ՗o C3 Ϛ
3,20)

~ē

gΔZͷЉ˅ 2 nÑ¡ÖŜΖYaÑ¡

Q C4 ϚÖ¡Ë°ÒՖC2 ƽҺ՗ΖIgЖϯ˷ԩЊмv

ho RuBisCO n¡²Ú¦ƒɶldgΔ

CO2 ʹУYgLՖǁ 1՗?UnUiN”C2 ǌ”

Z 2-Ã£ÃÑÚÒӵNÂÒ¡§ÚÈiIJơϖnΘ˹ikdgI@C2 ƽҺŅнhÑ¡ÖviǡʵW?Жϯ˷ԩЊмnÇ°oՌI CO2 ѰĻ͸χ[@C2 ƽҺ

ǁ 1. Flaveria ȬlLS C3 ǌM C4 ǌvnӜżӡϙn̡ɘǁß̵եƝ̖ΆϚnʈ;љṇˌŖ΃@Лыǀ{oљжЊм?ÓÖ¢ыǀ{oЖϯ˷ԩЊмnĄѩΪkĒЩχYgI@za?ԣыǀ{oЖϯ˷nĒЩχYgI@bar=505m
à̵եƝ̖ΆϚlLSĄқЎҺn̜Οǁ@Лыhχ[ЊмoљжЊм?ÓÖ¢ыhχ[ЊмoЖϯ˷ԩЊм?ԣы̀oЖϯ˷?ԣοƉhχ[ЎҺo C2 ƽҺ?ұοƉhѩ[ЎҺo C4 ƽҺ_`ѩ
YgI@Flaveria robustaՖC3 ՗hoЖϯ˷ԩЊмnÇ°Ö±ÑNЖϯ˷ĲlİdgӱЩW
proto-Kranz ̞әNѵ@Ñ¡ÖեGly?¥ÑÖեSer53

ŀƞʈσϡ 28 (1) 2018
Ö±ÑviӈӓWg_Uh 2 ŕǺnÑ¡΁ΓǚàlH̖ΆnЖϯ˷ȔɒNՌIUii

ÖM CO2 i¥ÑÖΔʈ[@ΔʈWa CO2

ԅӚ[ƘнɻʤʻYgI 40)@o RuBisCO ldgƠżW?~JÛˎn¥ÑÖoÂÒ¡§ÚÈhÑ¥ÑÖӵlǡʵW

4. C2 ǌՖC3-C4 æԄǌ՗M C4-like ǌv

aɨlљЛĖh 3-Ã£ÃÑ¥ÑÖӵik

 Flaveria Ȭn C4-like Ϛho C4 ƽҺŅнYg

C3 ƽҺviƞ͑[@Un C2 ƽҺl CO2 ʹIN 27, 28)?C2 ϚՖC3-C4 æԄϚ՗nǤQo̧нУhiknoÇ°Ö±ÑlȧǇ[Ϊk C4 ƽҺʣakIUiNǗƧWgI 3)@Ñ¡Ö¯ÒÄ¡ÐÚ¦ՖGDC՗nΦΏԱÛˎh? 14CO2 nƔǐhH

ՖC3-C4 æԄϚ՗h~ C4 ¢ÒÄÖӵvn CO2?C3 ǌhoљжЊмiЖϯ˷ԩЊмn

ӌ{ȋՊlLIg C2 Ϛ

jcnÇ°Ö±Ñl~ GDC oǽǇ[N?

nƔ

C2 ǌՖC3-C4 æԄǌ՗hoЖϯ˷ԩЊмnÇ°

i~ǗƧWgI 26)@za?Flaveria Ȭ°ÐÖ

Ö±ÑlԊȈWUihÑ¡ÖNʹɒğ£Ñ¿°ÚÈ̸Ӄѽ˼M Flaveria Ȭ C2 ϚǽΪlЖϯ˷ԩЊмviϗųYôЊмԄhn C2

ՖC3-C4 æԄϚ՗ho C4 ƽҺhĸQӴЉЬhHƽҺNĸQJlkdaiбKgI

10, 40)

ӌ{NēIÓÁÒhoHNѵU@

PEPC?¼Ò»ÖӵÑÖӵ¢²Ú¦ՖPPDK՗?

 za C2 ǌՖC3-C4 æԄǌ՗ho GDC nȧǇbNADP-ME?£¹ÐÖӵÇ¶ǓӀϗӴЉ?

ShkQљŌ̞ә~ C3 ǌi̸ӃYgǦOQǡż

Ð³ÖÇ¶ǓӀϗӴЉkjnΦΏN C3 Ϛi

YgL?Жϯ˷ԩЊмNзǦżYЖϯ˷ԩЊм̸ӃYgՌIUiN˗MlkdgI 22)@UŌnÇ°Ö±ÑiљЛĖnӱЩNЖϯ˷Ĳ

nUio?C2 ϚՖC3-C4 æԄϚ՗lLIg~ C4nЊмǜlİdgI?ř˯nÐÖ­̞әɠĄқӴЉĜdaĄқЎҺNӭŕΪļн[40)ʈYgI Ֆǁ՝՗@UnЖϯ˷ԩЊмŌnЊ

UiʀĹW]@Mallman o?C2 ƽҺldмȡƼȇnӱЩo?Ç°Ö±ÑiљЛĖnĒgЖϯ˷ԩЊмlLIgÑ¡ÖnрͷӵlЩNӎQÇ°Ö±ÑhˀœWa CO2 NŎ

ďdgΔʈWÖÊ³Ąқ[a}lsͮœ[nʖŞYљЛĖlůΌΪl°Ð¬

C4 ĄқӴЉЬNĜgIƘнɻʤʻYg¿WUiƘнlYgIiбK@I 22)@ŬKg?ϩвnѽ˼l

Flaveria ȬƤ} C2 ǌՖC3-C4 æԄǌ՗NΦΔYȬ C2 ϚՖC3-C4 æԄϚ՗Flaveria floridana ho C4aǈǐoùͿɅhH?ŀƫƥ͐ɻNՌQkΔƽҺḩн[ӴЉЬoљжЊмiЖϯ˷ԩЊ

38)?C2 ƽҺl CO2 ʹУ̧к˸ĈhHa}мnäˎhΦΏYgL?Flaveria?ΦΏnЊм·Ρɻo΋̞N˫ŜhHdaiбK@

ɫWgIkIUiN˗MlkdaՖTaniguchi

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QeMn C3 ǌhoЖϯ˷ԩЊмnÇ°Ö±Ñ

n C4 ƽҺ CO2 ʹУo̧нYgIkIniб

NːlЖϯ˷ĲlİdgȧǇYgI

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proto-Kranz ip̞ә~eϚNHUio C2 Ϛ CՖ3-C4 æԄϚ՗h[hlΦΏNß˕YgNǗƧWa40)

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?C4 ƽҺ΋ɫ[a}nɠүǡżo C2 Ϛo?Жϯ˷ԩЊмlLIgŀƫƥldgˀœWՖC3-C4 æԄϚ՗n̵ԓhːlǴzdgIiб CO2 ůΌΪlѰҷ[UiƘнlY?

K@Ûˎ?Flaveria Ȭ C4-like Ϛ Flaveria

C2 ǌՖC3-C4 æԄǌ՗vnӜżnӡϙh GDC N

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F. brownii hoēQ C4 ƽҺŅнYgIUiN̞әn͞ȢoЖϯ˷ԩЊмlLS PSII n͐ɻ

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ATP N՟ŕǺi NADPH

ƽҺN CO2 ʹУЎҺiYģн[Jlkd

N 2 ŕǺɳѴhHN?C4 ƽҺՉųYaЏ˿

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F. brownii hoљжЊмlLIg~ RuBisCO NΦ

ȈHa

7)

ATPաŕǺÙNADPH2 ŕǺNɳѴik

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ho RuBisCO n¡²Ú¦ƒɶ~ҳUanΔΕkYl ATP ΔΕhOɮΓǌԡǺĎӣΏYgL}?C2 ƽҺl CO2 ʹУ~̧нYgIUi
NбKgI

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RuBisCO nΦΏNљжЊмl̳ǽYgIez

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γӾǌԡǺĎ

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6. Жϯ˷ԩЊмiљжЊмnŕż

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ՖC4 ǌ

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5. C4-like ǌM C4 ǌv

jleIgoΏǇ~xijѽ˗WgIkI@ Ӝżn˩Ќ£®¬¿hH C4-like ǌM C4 ǌ

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vnӡϙho?RuBisCO nΦΏԱǐNȄńlЖ

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14, 34)@za?UlgIiIJĆґNʴƴWgL

21)

?ӎɊl

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kdg°ÊÔ¡n Zmscr1 ǡΡĖlLIgЖ

@RuBisCO nΦΏԱǐnԊȈżloӀŏɨŞϯ˷ԩЊмnљЛĖΦӣlΡɆNѻȗWaU?_ņ

iM 43)?Ê¯ÒhʴƴWgI¡²ÒƾǺ

̞iYg mRNA nȃȈżlԅâ[ƾǺ©Ö

o SHORT ROOT(SHR)/SCARECROW(SCR)ho¹үnќϟlԅâ[ƾǺkjNбKg

kIMiʤʻWgI 11)@SHR i SCR o GRASINҊYIUio˗MlkdgIkI@za?

½ÇÑÚlȬ[ӀŏƾǺhH

C4 Ϛho RuBisCO nĂl C4 ƽҺnĄқӴЉЬ~¤²lLIg̎n”Жϯ˷-Ōά-άȭ”iIJˀȞЊм·ΡΪlΦΏNŞɬWgIN?UnЊмΈ¹©ÚÖnɠʈŞɬ[Ֆǁ՞՗@¡Ô´·ΡΪkΦΏŞɬloӀŏŞɬӀŏɨŞɬ?

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̞Nԅâ[UiNχƱWgI

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 SCR iγʱΪlδõęΖ[Uil@UnĂl

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Aե¡Ô´²¤²n̎ṇˌԦn̡ɘǁ@ԣыoæɲ̅ՖЖϯ˷ЍЦ՗?ÓÖ¢ыoŌά?Лыoάȭ?
Ψыoǣά_`χYgI@BեSHR Ls SCR lŞɬ̧̞@SHR oæɲ̅hӀŏÙа҄W?
_nԘnŌάЊмviʡ˃Y_UhΦΏ[ SCR iγʱΪlδõęΖ[@ɠʈWa SHR/SCR ѲƞĖoŌάЊмn̍lȧǇY SCR фҽnӀŏ̯lŞɬ[Uil ½Ú±¸¬ÒÚ¿ɠʈYgL?ŌάЊмnԘՖάȭ՗v SHR Nʡ˃[UiԈȏYgI@ŌάЊмnWlԘՖάȭ՗v SHR Nʡ˃[Ǻ̧̞nѽ˗lo˳љΦӣnř˯MЖϯ˷ԩ

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HiбK@za?ΏǇʉAo Flaveria Ȭ16, 17, 30)@Un£ÚÈљnЖϯ˷ЁlЩOʵ

C4 ϚnǡΡĖѽ˼ C2 Ϛi C4-like ϚnϚԄúԜKgбKi?Жϯ˷hΦΏ[ SHR NЖϯ˷ϚɨĄnѽ˼Ӝ}gL?Unѽ˼NӜ}ԩЊмlʡ˃Y_UhΦΏ[ SCR iδõęΖpŕǺΪk C4 ӜżnӡϙN˗MlkUiNYЖϯ˷ԩЊмhHiIJ·ɰ́Ȉ[i˯ɦW@

i~l?SHR Nʡ˃YgUkMdaЊмoљжЊмikiIJÊ¯ÒlkN?UnÊ¯Òh

Received Mar 1, 2018; Accepted April 4, 2018; PublishedoљжЊмÛȭlѶȈ[UiNhOkI@W

April 30, 2018.

lΏȋo~JȢYѲԜkJh?°ÊÔ¡nЊмϚś°ÐÖ£Ñ¿°ÚÈѽ˼lLIg°ÊÔ¡lѲ˅H SHR ӨĎǺnJcnÛ

ƎбˉΊeoЖϯ˷bShkQЖϯ˷ԩЊмh~ΦΏY

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372, 20160386.

of phosphoenolpyruvate carboxykinase-type C4 leaf

Sage R.F., Sage T.L. and Kocacinar F. (2012)

anatomy: immuno-, cytochemical and ultrastructural

Photorespiration

and

the

evolutionofanalyses. Ann. Bot. 98, 77–91.

C450.

photosynthesis. Annu. Rev. Plant Biol. 63, 19–47.
39.

Voznesenskaya E.V., Franceschi V.R., Kiirats O.,

Voznesenskaya E.V., Koteyeva N.K., Edwards G.E.

Sage R.F., Khoshravesh R. and Sage T.L. (2014)

and Ocampo G. (2010) Revealing diversity in

From proto-Kranz to C4 Kranz: building the bridge tostructural

C4 photosynthesis. J. Exp. Bot. 65, 3341–3356.

photosynthesis and a C3-C4 intermediate in genus

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Portulaca L. (Portulacaceae). J. Exp. Bot. 61, 3647–
51.

52.

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3662.

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Wingler A., Walker R.P., Chen Z. and Leegood R.C.

Deficient photosystem II in agranal bundle sheath(1999)

chloroplasts of C4 Plants. Proc. Natl. Acad. Sci. U.S.A.

Phosphoenolpyruvatecarboxykinaseis67, 18–25.

involved in the decarboxylation of aspartate in thebundle sheath of Maize. Plant Physiol. 120, 539–546.

Evolutionary progression from C3 to C4 photosynthesis in the Flaveria

Yukimi Y. Taniguchi
Department of Bioscience, School of Science and Technology, Kwansei Gakuin University59

ŀƞʈσϡ 28 (1) 2018

特別企画：若手研究者の海外留学レポート！

第三回「スイス滞在記」
ETH Zurich(現東京大学大学院

農学生命科学研究科)
森田 隆太郎

私は神戸大学の深山浩准教授のもとで光合成に応答したイネの澱粉生合成制御機構の研究で学位を取得後、日本学術振興会の「特別研究員に向けた海外渡航支援日本-スイス若手研究者交流事業」に採択され、2017 年 4 月から 10 月までの 6 ヶ月間スイス連邦工科大学チューリッヒ校 (ETH
Zurich) の Samuel C. Zeeman 教授のラボに客員研究員として留学してきた。本稿では短い留学期間ではあるが、著者のスイスでの体験を若手の皆様と共有したい。

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60

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61

ŀƞʈσϡ 28 (1) 2018報告記事
第 14 回若手の会セミナー開催報告

東京大学

大学院総合文化研究科
清水 隆之

 яʏnčhnřn҇{iYg?ÿƽn¥Ç²Úo[wgѐҎhѦzYa@ʟɦҚͯвn 1 ýhH?ûӰǦǿn Hedi Pedajas Wl?¥Ç²ÚlƎŬYaʃʀΦѩYaσϡŌȑkjǗƧ҂óiYgzi}gԬIanh?_c~ě]gVѹQbWI@ ÿƽo?11 ˪ 11 ˑlǔ΍ǦǿnҚЭȍLĮ YgѦIzYa@ԃĶlԖYg?ǔ΍ǦǿnՌ̥ʞǺWiόĥˠɡWlo?ĊΜÙӟƸnɌóiYgǤǦkVƄŪԬOzYa@UnǘĮ gɬψΚYßTz[@¥Ç²ÚnŌȑiYgo?ˑ˳h͐ҼYgIǣǃýσϡвÙΞǿΔ 5 ơՖ˺ȺǦn Imran Pancha W?˺ȺǦn Luu Trinh W?ǓΔσn Eunchul Kim
W?ûǦn Hedi Pedajas W?ǔ΍Ǧn Isura Nagahage W՗i͖ǣlΞǿYgσϡѦdgIaˑ˳ý 2 ơՖόʋǦn̕ΗW?ǔ΍ǦnόĥW՗lҚͯYgԬOzYa@ ļϐkǣǃýσϡвÙΞǿΔNŀƞʈԅӚŕӸl~ǤQIl~ԅ\?˳čloƎŬYgIkIa}ú͑ņčNkIUilʆKaόĥWnΦ̒h˳¥Ç²ÚnԃĶlх z
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̫ƽo?˳čЌðɨl¥Ç²ÚԃĶñȈh[@½ÐÖkčh[nh?L̻ӂlƎŬԬS
piɹIz[@62ŀƞʈσϡ 28 (1) 2018ՖϨ 14 ƽяʏnč¥Ç²Ú ƎŬвM՗

An initiation into the world of PPR protein (presentations)
Laboratory of Plant Molecular Genetics, Department of Botany,
Graduate School of Science, Kyoto University

Hedi Pedajas
Amongst the golden gingko leaves of last fall, I had the pleasure of presenting my work at the 14th
Young Researchers Seminar of Japanese Society of Photosynthesis Research in Saitama, the first (ofmany, I believe) to be held in English. To put this into context, I must admit that I switched to mypresent research field of “Gene expression regulation by PPR proteins” only recently ahead of myentering into a PhD course in April 2018, and have absolutely no experience in presenting my work.
Which roughly translates to “full-scale panic mode!!” However, arriving at the venue despite getting
slightly lost on the way, I was quickly put at ease by the friendly atmosphere the organizers created inour small gathering. Even though my topic is only remotely connected to photosynthesis, I hope aninterest might have been sparked in some people hearing about these fascinating RNA binding
proteins for the first time.
Modular PPR proteins recognize specific RNA sequences following a set of rules termed the PPR
code and are implicated in most processes involving the nucleic acid. For example, proteins from the
RFL family have been shown to be involved in post-transcriptional modification of abnormal andpotentially harmful mitochondrial RNA and are characterized by their distinctive rapid and active
evolution. My research focuses on the collaboration of nuclear-encoded RFL2 and mitochondrial
endonuclease RNAseP (PRORP1) in degrading the transcripts from chimeric orf291 (partial cox2 +sequence of unknown origin) in Arabidopsis mitochondria. To clarify the interaction, I am
reconstructing the reaction in Nicotiana benthamiana by using chloroplast transformation to create
plant lines with the expressed target (orf291). The two protein genes are introduced into thetransplastomic plants transiently by using Agrobacterium infection. This is expected to induce anendonucleolytic cut in the target, observable in Northern Blotting. As stressed in my presentation, thissystem is powerful because it allows fast and relatively easy testing of the effect of mutations in the
PPR or can be modified for other PPR proteins and their targets. Therefore, the benthamiana transient
expression method is versatile and has benefits over creating stable transformants and should beconsidered as an alternative when designing your research system.
Ending on this positive note, I would like to take the opportunity to thank the organizers for
inviting me to give the talk and say: “Until next time!” I mean karaoke, of course. ;)63

ŀƞʈσϡ 28 (1) 2018

集会案内

第 26 回「光合成セミナー2018：反応中心と色素系の多様性」の開催案内
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 2018 ɊՖɉʈ 30 Ɋ՗7 ˪ 6 ˑՖӺ՗ƎŬҨեՖ7 ˪ 21 ˑnʇѺčҨ?7 ˪ 22 ˑn˛ԼĄƤ|՗   Ûщ 5,000 ōՖñȈ՗   ǿΔ 3,000 ōՖñȈ՗ã҈ýեϒ˳ҏɴՖόʋǦǿ՗?ǦȰȅәՖǦԆǦǿ՗?ǦƐɤȁՖђǏǦǿ՗?
    œЮ̶éՖơƗȪȺ̙Ǧǿ՗?̽ȳҮ͇ՖόǬȸǦǿ՗?Ȏ̰ˢɚՖՕҟǦǿ՗ΚYӌ{ÙƵIƞ]Ŀե όʋǦǿǦǿԋ Αǿσϡϓ ϒ˳ҏɴ       (E-mail: photosynthesis@boar.kobe-u.ac.jp, Tel: 078-803-5705)
  ¿ÔÐÈLsÿɨn̒Ōoà҂ÃÚÈÂÚ¢lg?˦ˍɿǗԔ˜?ʳӄIaYz[@
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「光合成研究」編集委員会
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2017年度役員čԀՌ̥ ѯÛӬՖȰȮǦǿ՗óŴȧԀǄ̿ Ņ̶Ֆ˓ϛΗǦǿ՗ɆĉɌóΗæ ̱ՖŽ͖ӢǦǿ՗

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˻Η ϊĀՖԅѳǿԋǦǿ՗ Ɋč 2015ɊɆĉɌó́Ȯ ϊÛӬՖϏ÷Ⱥ̙Ǧǿ՗Ɋč 2016ɊɆĉɌó˴͒ ЫЮՖʂǹǦǿ՗Ɋč 2017ɊɆĉɌóƗ˳ ɞՖՕҟǦǿ՗Ɋč 2017ɊɆĉɌó΅Ӹ ƩՖ˺ŽǦǿ՗Ɋč 2018ɊɆĉɌóȎȦ ŀɾՖ˺ŽǦǿ՗Ɋč 2018ɊɆĉɌóԨ ӣíՖ˺ûΑϓǦǿ՗ŀΔΆǿƄčɆĉɌóςŽ ǩՖ˺ûǦǿ՗ɆĉɌóċϏ ıǧӬՖûӰǦǿ՗čҀΰ̇ѡΗ ϊÛՖơƗȪǦǿ՗ÃÚÈÂÚ¢Ŭѡ ѯĀՖȰȮǦǿ՗ОԚԀŀƞʈσϡ Ϩ 28 ɀ Ϩ 1 ƛ Ֆӗɀ 81 ƛ՗ 2018 Ɋ 4 ˪ 30 ˑΦѦˑ ˳ ŀ ƞ ʈ ǿ č
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