光合成研究ξ 27 Ȑ ξ 1 ŷԩҦȐ 78 ŷԪ2017 ȝ 4 ˃
NEWS LETTER Vol. 27 NO. 1 April 2017
THE JAPANESE SOCIETY OF PHOTOSYNTHESIS RESEARCH
集会案内ξ 8 ƘʦˌĢŹɞǘðȝð»Ħӛº©u¬ӛėU3ΗiD

2

トピックス ˬ͢5ǜɅ@NĢŹɞO6k¹ԼPS I oĢĘǥ4iѮk P700 Ӆŗ¬ԼȇȈӐ̟ ÁǙчȭԩΦɡǁԪ

4

研究紹介 Ξŗ̖ϜoӹǓāÅùPBkӹǓñҳϕҹñǓU΁ͫћǵԸ     Ҏįľȿº~ѿ SqrR UŻǠPFUΞŗ̖ϜɈτ˻˳
研究紹介  
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31

序文

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解説 X ϯГͳӹǓ·¼

Ⱦ̫ өǃҿԩưͯΛԪ ˜ȁ Τ½ҿԩΩÛȋˮǁԪ
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解説 Photosystem II S3:k̖ϜϤŹ¹¼~oã@H£¸ºαŐʝЬƥßԩ˕ÞǁԪ39

解説 ˬ͢UĢŗǘϕ IԭӲĢɎrºпŹù I ҄пŹùUϤʹ˳ҨPw¶}¼ñҳU˳ҨưΊТĒǬԩȂȁǁԪ48

ŧͲ ØˆԩÌͺϏǁԪ55
報告記事 ξ 2 ƘĢŹɞϟУ¹¼~³£ӛėƵƁ    

集会案内 ξ 73 ƘЬŧ«¼ “The 4 International Conference “Molecular Life of Diatoms”th˖Ͳ ΤãԩӝрǘӣǁԪ

57

集会案内 ξ 8 ƘƟӮðѭ “Photosynthesis and Hydrogen Energy Research for Sustainability -2017”Ԁ ҳÑԩ˕ÞͭέǁԪ

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集会案内 ξ 25 Ƙ)ĢŹɞ«¼2017ԸŮɈÉɅPЛϜϕUǀ˴Ɏ*άˌ ѝɇԩΦɡǁԪ59˲ˌ ğԩ˕ÞǁԪ

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報告記事 МɥUðUȟ×UÝçU=ƵƁ»ʢðӘ4iU=/0>L̮Û ʺȠԩμƅԗǁԪ×őǻ4iU3ΗiD

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トピックス
植物が安心して光合成できるワケ～PS I を光傷害から護る P700 酸化システム～神戸大学大学院 農学研究科 生命機能科学専攻
嶋川 銀河, 三宅 親弘*
本稿では光合成生物において普遍的に見出される生理現象「P700 酸化」について、その生理的意義および P700 酸化を支える多様な分子メカニズムを紹介する。光合成生物は強光や低 CO2 といった過剰光環境にさらされると光化学系 I (PSI) の反応中心クロロフィルである P700 を酸化する。この事は植物生葉において 90 年代から見出されており、庭先に生えている草木などで簡単に観察できる。なぜ光合成生物は P700 を酸化する必要があるのか？どのようにして P700 は酸化されているのか？その長い歴史の中でいつから光合成生物は P700 を酸化するようになったのか？近年私たちの研究グループは
P700 酸化に関するこれらの問いに対する答えを探ってきた。豊富な「光」と「O2」に囲まれた地球上において、光合成では過剰な光によって有毒な活性酸素が生じる恐れがあり、PSI 内部で生じた活性酸素は PSI に酸化傷害をもたらしてしまう。しかしながら、ダイナミックに変動する地球環境の中において、光合成生物は P700 を酸化させる事によって PSI での活性酸素生成を未然に防いでいる。この P700 酸化は PSI のドナー側およびアクセプター側から多様な分子メカニズムによって制御されており、私たちはこれらを P700 酸化システムと定義した。P700 酸化システムは酸素発生型光合成の始原生物であるシアノバクテリアの時点で既に多様化しており、またその長い進化の歴史の中で変化してきた事が明らかとなった。一見すると何も考えずのんびり光を浴びているだけの植物、そんな彼らの壮絶な生き様に関して、一人でも多くの方々に興味をもっていただければ幸いである。

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V 3 ȤUǢԜUȜƨđP˵́ўȏShKNΡ>lk' (C) ª¹µͯХS3:kĢȰȤP P700 Ӆŗ
Uӝą. FlGlƝ 2A 3hYƝ 2B OΡ@H P700 ӅŗP̋Ǖ PSI ̬Ɏo£¸@H'¼VĥNSejima et al. (2014)2)4iȬͱ@H'ɗЁ5 PSI ĬӀS3:k̬ɎӅϜͯɞUÖӟB

RnJĢĘǥSǰBk PSI ӟȿO/k×oĹc
NΡ@H (Ɲ 2) 2)'ʻɣS30Hª¹µͯХSԔƆ¶ĢoҩϪ΅S͛ǳBkPǢԜǣ·¥¶OǧʭS PSI ĢĘǥoшǭBk×5O6k5&<UʴSԔƆ¶ĢPŻʴS½ǠȰȤUǠȘ
Ģo͛ǳBkP PSI ĢĘǥVɪľ>lk 2)'dJmpǠȘĢUȰȤ5Â5kSLlN P700 5Ӆŗ@N085 (Ɲ 2A) &Ӿ΃0×S P700 5ӅŗBkSLlN PSI ĢĘǥVтilR8Rj (Ɲ2B) &Ϥ˚P@NԔƆ¶Ģ͛ǳʴU P700 Ӆ6ĢŹɞΛθ 27 (1) 2017
Ɲ 3. P700 Ӆŗ¬U®¶(A) FLV ĂǕƪU P700 Ӆŗ¬®¶. ǀ8Ur~µreϮЭRQ
5<JiS/NV`kPЄ2ilk' (B)ĢƄƀĂǕƪU P700 Ӆŗ¬®¶. ǀ8UиǓˬ͢P8S C3 ˬ͢5<J
iS/NV`k'3 LU¶¢VFlGlw¶}¼&ӹǓ&£¸º¡´~UǜĥȪoΡB: NPQ, ӽĢŗǘ΅̴Ģ;ge−(f), ~¸¬ b6/f пŹùƂҘS3:kw·~¸ºº~º; gH +, ATPŹɞӄϜS3:k£¸ºº~º'FlGl NPQ P ge−(f) Vҁ͖ϯOΡ> l k h 1 S ΔpH e £ ´   |  º(PQ) ӅŗһğoəűBk'`H&PQ һğS҃ƙBk ge −(f)UöÃV RISE P@NǠЁ>lk 30)'Y(II)&Y(I)VFlGl PSII
3hY PSI S3:kĢŗǘŮɈUǢŏӋǓůͨoΡ@&VH +V ATP ŹɞӄϜS3:k£¸º̭ĴҧȤoΡB'ƝV
Shimakawa et al. (2017)29) oŪЄS@Nü
ɞ@H'PЄ2ilk'Ɲ 3 OV&~¸¬ b6/f пŹùŗPǆ̬@R4KH PSI PUӜSʬΟR΍ӝ5

ailH (Ɲ 2C) '<U×4i P700 ӅŗUͯͭS3:kӹǓṶleB>o ge−(f)P@NжB'

2)

΅ɗЁV PSI UӟȿO/j&PSI ĬӀS3:k̬~¸¬ b6/f пŹùS30N¢² f UɎӅϜͯɞoɪľBkHcS P700 5Ӆŗ>lkһğҧȤV´tВ¶¼­ºĬUӅɎŗS

×5ĵ4k'ӅŗƪU P700 5ƺŊBk×V&ưòKNԍЦSöÃBkHc 32)&ĢŹɞͯ͢V ΔpHȡͥɚU P700 5̻ǷBk×oɗƃ@N0k'L

oľȿ@N PSI \UӹṶ̌ĤS¢·¼|o4:ǓUāϦ̀5̻ǷBkPPdS& P700 ͯɞUµ

ɞVӹǓñҳ̬ɎSĂǕBkHc 33)&ĢŹɞUԛ

`j P700 5Ӆŗ>lk×ShKN&O2 \̼kӹ

k×Shj P700 oӅŗO6k'ưˌ΅S ΔpH ȳ

3~5ϲƆ>l&FUϤ˚P@N O2−e 1O2 Uͯɞ5ɪľ>lk

19)ŐUaRiCȼҠBk O2 ĂǕ΅çˁ΅ӹǓñҳeɁͮ΅ӹǓñҳ (Cyclic electron flow, CEF) V'¶¼­ºĬUӅɎŗSǩÅBk×5ˈȺ>lk
4. P700 Ӆŗoʒ2kĵǓ­z¬29), 34), 35)z¬ShKNпŹ΅SȬ6҃<>lk'ˌεВƿ\U£¸º̭ĴҧȤ (VH+) 5ɪľ>l¸ºº~º (gH+) 5öÃ@&´t ĢŹɞͯ͢S30N P700 ӅŗV˴(RĵǓ­k×ShKNd ΔpH VÂʪBk 36)'>iS´

OV<liĵǓ­z¬o)P700 Ӆŗ¬*PƄ[ (Ɲ 3)'`H½ʣO&ATP ŹɞӄϜS3:k£'P700 UӅŗһğV PSItВÂSǕƤBktyº¯¶ K+-efﬂux

14, 29, 31)

antiporter 5¶¼­ºĬ4iU£¸º͈ĴoѢUӹǓUĤŲԩ¼ĔԪPĴŲԩr~£¼

ʚBk×O ΔpH ȳɞSȶμL×dΡƌ>lN0

ĔԪS3:kӹǓñҳŮɈU´ºShKN̛k 37)'FU^4ӦÂˬ͢eϮЭS30NV proton

Ǡ>lk'èÃOV&FlGl¼Ĕ&r~£¼ĔSĵ:NԆS P700 Ӆŗ¬oϠã

gradient regulation 5 (PGR5) 5 ΔpH UľȿSǁ6

Bk'RȶŇodL<P5ƵƁ>lN0k 12, 38)'

 PSI U¼ĔS30N˂d˄ňR P700 Ӆŗ ~¸¬ b6/f пŹùƂҘS3:kӹǓñҳŮɈV ΔpH UaRiC&£´|º£¼¶¬V&~¸¬ b6/f пŹùS3:k£¸UұһğShKNdɪľ>lk'<U P700 ӅŗºŕӃ (ΔpH) ĂǕ΅RӹǓñҳŮɈUɪľ

7ĢŹɞΛθ 27 (1) 2017·~¸ºº~V C3 ˬ͢U P700 ӅŗSǁ68¬Vr~µrOтĴ>lreduction-induced suppression of electron transport(RISE) P@NƵƁ>lH

30, 39)ѵͦ@N3j&ͣSĢŹɞ5ɪľ>lkö CO2'RISE 5ͯAkӮ

ͮƹ3hYFlodHiBȰĢ&Ô͟·ÃSV&£´|º£¼¶UұһğSÈб@N

RQOV&½ǠUĢƄƀ̬Ɏ5R:lWˬ͢V~¸¬ f 3hY P700&NADPH UӅŗ5љc

P700 oӅŗBk×5O6R0 45)'

il&Ӿ΃0×S<liӹǓñҳϕÃ̭UӅŗ5& ĢƄƀUåSǁ6Rw·~¸ºº~Ѝo

ΔpH SǩÅBkPЄ2ilk O2 ĂǕ΅çˁ΅dLçˁ΅ӹǓñҳP@N Flavodiiron º~ѿ (FLV `HV FDP) Shk O2 һğŮɈ5ɺ;ӹǓñҳU˿ʋˢS30Nb@mԍЦSтil
H'ɕǠ>lk RISE UĵǓ­z¬P@N&

ilk 46)'FLV V¡´º®~·yϤӅŗƪ£´|ºUÄ҇Shk Q-t~¶Uɪľ5ʇƎ>lN3j

30)

ŹtP 2 ӏº¼o˄@H¡´ºº&âȼVӦÂˬ͢RQ

~ѿO/j&ĢŹɞͯ͢UÉOVr~ΑˣĢŹɞͯ͢dſcHіϟRΛθ5ˈȺ>lµr&ϮЭ&&&нǓˬ͢S§®¸5k'тĴ>lN0k 47)'FLV V PSI Ur~£¼

ĔS30NӹǓoű:ŰKN O2 o H2O S 4 ӹǓ

 ~¸¸¡s¶ЯĢ̾ǠShKNǠЁ>lk
PSII

O U Ģ w  ¶ } ¼ U ͞ ʔ ʗһğBkPЄ2ilk5 14, 46, 48, 49)&FUŮɈUͯ

(Non-photochemical quenching, NPQ) d`H P700

̬ͭɎVӽȘSԟ8r~µrOVĢŹ

ӅŗSǩÅBk×5ˈȺ>lk'ϮЭ~´«®

ɞ̬ɎSřʛBk^QUӹǓñҳ̬Ɏ5Οљ>OV ΔpH ĂǕ΅R NPQ (qE) SӝÅBklk 46, 50)'r~µrU FLV ˿ʋˢV&

ʋShKN PSI ĢĘǥ5̶ĿŗBk×5тĴ>P700 oӅŗO6C&PSI ĢĘǥoиKN@`1'

ƽŐĢ 51)eö CO214)P0KHұņĢͮƹS30N

light-harvesting complex stress-related protein 3 U˿lN3j&NPQ Shk P700 ӅŗUŴЍɎ5Ρƌ

`HҝȝOVӦÂSNĢŹɞoǍcHˬ͢'`HȠЁU NPQ P@N PSII/PSIU 29)e­µ{ 52)S30Nd

e PSII ĢĘǥ (qI) RQdɺ;ilk5&FlG

SѵͦBk×5ʬi4SRKN0k'ͣSȇȈi

>lN0k

40)

ӜUĢw¶}¼ĵӃƽŗ (¼ҷα, qT)

FLV 5 P700 Ӆŗ¬U½ƉP@N PSI ӟȿlŻ˴S P700 Ӆŗ\UǩÅ5Ρƌ>lN0k40-42)

29)

V&̖ӮͮƹOUͯǕ504SŤӧRdU4o'<liUϤ˚V)PSII 4iUӹǓñҳ*5

oͱ0NĹcNΡ@H'FLV o˿ʋ@P700 UӅŗһğ·¥¶Sǁ6RȵԂodL<P

HV̖̝BkPҧe4SĢĘǥUŤӧoкæ:N0k'

S>i>lk 29)'<U×VӦÂˬ͢Oę8ĢƄƀ

 ½ʣO PSI Ur~£¼ĔS30NV&ұņ5̖ÉOV˻Ѝ@R0×oΡ@N3j&`HӦS

SѵͦBk'ǀ8Uçˁ΅ӹǓñҳŮɈS30N

5ÄŴ˿IKH×oΡ@N0k'FLV 5 P700 Ӆ

Â5KHW4jUSPKN FLV UǕƤ

RӹǓUʗҬoб1çˁ΅ӹǓñҳ5 P700 Ӆŗ˂ϡ΅RӹǓUr~£¼V O2 O/j&<Uŗ¬P@Nę0N0H4i<F&ˬ͢UӦ

×VĢŹɞͯ͢5 O2 oҽɓRЃȨ͢P@NɐlÂҫĴ5҃<j2HUIm1'

kI:OR8˄·R)ӹǓUű:Ųԩw·~¸
5. P700 Ӆŗ¬Uǀ˴ɎPҫŗ΅ƽҷºº~Ԫ*P@NѾ8ļͱ@N0k×oɗƃBk'Ā2W C3 ˬ͢V&ĢƄƀçѫUÉO O2 oļ łԅOV˴(R P700 Ӆŗ¬oϠã@HͱBk×OұņRĢw¶}¼oÂɥ8̴ѸB

5&FlGlĢŹɞͯ͢ShKNQU¬okPŻʴSµ¢¸¼ 1,5-µºӅUĮͯɞļͱBkU4VǊ6Ǒ05/k'r~µoбKN0k43)

rV3hF 30 ĜȝłSͫlH˂ųUӅϜ΁ͯƪ

'C3 ˬ͢UĢƄƀ̬ɎVӽȘS

ԟ8&FUw·~¸ºº~ЍShKN CO2

ĢŹɞͯ͢O/k5&ȷiUÉS30NʥS
P700 ӅŗɠͻUγǀ˴Ɏ5ailk'˂dǀ8мĝ͖S30NdĢŹɞSřʛBkβUӹǓñҳ̬Ɏ5шǭ>lk

44)

ΛθˏʠP@Nɧnlk Synechocystis sp. PCC'ĢƄƀShkçˁ΅Rw

8ĢŹɞΛθ 27 (1) 2017
6803 (S. 6803) e Synechococcus elongatus PCCӹǓñҳ̬ɎoΡ@ 55)&FU˿ʋˢOVö CO2

7002 (S. 7002) VFlGlпʙU FLV rt

7002 S3:k FLV ˿ʋˢV&ö CO2 ͮƹS3:7942 (S. 7942) &̳̩ɎU Synechococcus sp. PCC

ͮƹO P700 Ӆŗ5ailR0 14)'Ϥ˚P@N S.

k 2 ʴӜUΙ0Ģ͛ǳ (290 µmol photons m−2 s−1)t¬oɷJ&FlSĂǕ@Hçˁ΅ӹǓñҳ̬ɎoΡB5&Ÿ(UγS3:k FLV U P700 ӅŗSǰBkǩÅȤVĥ8ͽRk (Ɲ 4)

14)

ShKN PSI ĢĘǥShkĢŹɞ̬ɎUǝĥR'S. 6803 V 2ǆ̬oΡ@H 14)'L`j S. 7002 VåU 2 γPV

U FLV ¤¸tª¼ (FLV1/3 3hYͽRj PSI ӟȿo^PpQ FLV UaSĂǕ@N

FLV2/4) o˄@&ö CO2 ͮƹS30N FLV2/4 o0k'<liȇȈ&ӇiUΛθϤ˚Shj&ӅϜ

49, 50, 53,΁ͯƪĢŹɞU¶¼oɰ1r~µr'½ʣO S. 7942 V FLV2/4 oɷHC&ö CO2

S30N)Ģ*5ˌѿ΅SdLŤӧɎ3hY P700

ͮƹOVĢŹɞPPdSӹǓñҳŮɈoɪľBӅŗUɆсɎ5ʥSΟμ@N0H×5Æ͸OĹ'@4@R5iӾ΃0×S&<li 2 γUcNΡ>lHPŻʴS&PSI S3:k̬ɎӅϜͯԟ΁ͫBk×Oçˁ΅ӹǓñҳoԛŐBk
54)

k

50)

ɞɪľ­z¬U̅ù5 P700 ӅŗO/kP@r~µrV FLV 3hYFUçˁ΅ӹǓñҳ̬ɎSӝąR8ö CO2 ͮƹO P700 oӅŗBkH͒ȄiUéџ 2)5Ǣё>lH'

×5O6k 14, 30)'<lV¼ĔS30N RISE ĢŹɞͯ͢V&FUҫŗỦŵUÉOFlGl

Є2il&Ϥ˚΅S<li 2 γUӊͯˢ3hY>DN6H'r~£¼ĔU P700 Ӆŗ

UͯǕͮƹSŹnDN P700 Ӆŗ¬oƽŗRQĻU P700 Ӆŗ¬5˻ЍBkHcIP¬O/kĢƄƀP FLV oĀSɺ;H0 (Ɲ 5) 'FLV ˿ʋˢVö CO2 ͮƹS>i>lNd PSI ĢĘǥoиiR0

14)'½ʣO S. 7002 Vö CO2 ͮƹĢƄƀV C3 ˬ͢S30NӽȘSǁ6Rçˁ΅ӹ

S30N FLV1/3 SĂǕ@HӽȘSǁ6Rçˁ΅Ǔñҳ̬Ɏoɰ15 44)&r~µreϮЭƝ 4. ͽRk 3 γUr~µrS3:kö CO2 ͮƹÃOU P700 ӅŗP PSI ĢĘǥ(A−C) ĢŹɞ CO 2 ͑ȤȻҧˈ\UαбS3:k P700 ӅŗUɈτ. r~µrͯϟЌoſbŮɈ̵̈́(HCO3−o̸Ŋ@R0) SĢo͛ǳBkP&A6S̵̈́ͮƹÉU CO2 5ĢŹɞShKNԒ0ǹ8>lN@`0&ĢŹɞ5ɪľ>lk'<UͥɚoΫHJVĢŹɞU CO2 ͑ȤȻҧˈPǠЁ@N3j 49, 50)&<UP6ĢŹɞVǁ
̕É4ḯ̵É\U CO2 ɶʗShKNȻҧ>lN0k'Ɲ 4A−C OV&CO2 ͑ȤȻҧˈ\Uαб5ͯAN0kʴӜo͖ϯΖţOΡB'ĢȰȤ 180 µmol photons m −2 s−1 UŎ҃Ģoͱ0H'ԦÊ5ӊͯˢ&΃Ê5 FLV ƽͽùo
ΡB' (D−F) ö CO2 ͮƹÃOͯAk PSI ĢĘǥ. ĢȰȤ 290 µmol photons m−2 s−1 UŎ҃Ģoͱ0N CO2 ͑ȤȻҧˈO 2 ʴӜUĢ͛ǳoб0&FUȼ 1 ʴӜUʻԆŗoɻpŐǕ PSI ̬Ɏo̾Ǡ@H'ƝÉŰǕ PSI ̬ɎVĢӅŗŴЍR P700 UĥӋ (Pm) oɗƃBk'¼VĥN Shimakawa et al. (2016) 14)4iȬͱ@H'9ĢŹɞΛθ 27 (1) 2017

Ɲ 5. P700 Ӆŗ¬O/kĢƄƀ3hY FLV Uҫŗ΅ƽҷCyanophyta, r~µr;
Chlorophyta, ϮЭ; Bryophyta, ˬ ͢ ; Pteridophyta,  ˬ
͢ ; Gymnosperm, н Ǔ ˬ ͢ ,Angiosperm (C 3) , C3 ˬ͢'~´«®S30NVFU̬ɎVǶ>8&L`j̖ÉͮƹOVĢƄƀ5w·~¸ººP700 ӅŗSǩÅ@N0kPVЄ2Mi0'~P@N˻ЍO6R0×oΡ@N0k'HI@&

çnjS&<lir~µreϮЭOVĢƄƀçѫFUdUVr~µrUʴ͖

FLV 5ËсRw·~¸ºº~P@N P700 ӅOʥSͧȾ>lN3j&ǀ8UЭԏOͯЊSÄŴ

14, 46, 51, 55)

49, 50, 55)'ϬϾ΅

˿RçѫϕP@N˻Ѝ@N0k×Şɗ@R:RĢƄƀ̬ɎUђă4i&нǓˬ͢eˬ͢&lWRiR0 60, 61)'r~µreϮЭ&ͪ

F@Nˬ͢S30NVĢƄƀ5ŜĵRw·ЭS30NĢƄƀV P700 Ӆŗ¬OVR8&~¸ºº~Ѝo˄@N0k×5ʬi4SĻUͯͭ΅ȶŇoɰKN0kUIm1'Ӿ΃0<ŗSѵͦ@N0kPЄ2ilk

RKN0k

PS&ŻAӦÂˬ͢Od C4 ˬ͢V^PpQĢƄ

56)'½ʣOӦÂˬ͢PVĻUϕϧOҫƀ̬ɎoΡ>R0×4i C3 ˬ͢PVͽRk P700

ŗ@N6HͪЭ Phaeodactylum tricornutum OVĢ
Ƅƀ5w·~¸ºº~P@Nę0N0R

Ӆŗ¬o΁ҳ>DN0kPЄ2ilk 44,

4KH 57)'̖ÉOV O2 UɶʗҧȤ5ǁ̕ÉP̒

56)]Nϗ 10,000 ĵU 1 S`OöÃBk

'ǁ̕ȜдʴS3:k̖ÉU O2 ͑Ȥ5ϗ 250 µM O/k̬ɎUǕƤ5Ρƌ>lH 57)'Г͙͸SVΫHJU
ΗiR0 P700 Ӆŗ¬Uǀ˴Ɏ5`I`I

×oЄɛBkP&ĢƄƀUĹ΁ŮɈO/k¶Uy|¼ŮɈ5ΡB O2 \U Km đV

250~450 µM P/`jSdԟ0' ` H Э ԏ U É O d ² ¼  ·  Э Euglena

gracilis S30NVĢƄƀShkçˁ΅ӹǓñҳ58)

ΒKN0F1O/k'
 PSI ¼ĔU P700 Ӆŗ¬Sӝ@Nd&59)'<lioЄɛr~µrOтĴ>lH RISE 5ˬ͢\UBkPĢŹɞͯ͢V&ǁ̕ÉUѱǫR O2 SыlNĹcNĢƄƀow·~¸ºº~P@NļҫŗoϣHȼd̋>lHU4Vĵ4KN0R0'

ͱ@Ǎc&ӦÂҫĴ5ҫbSLlNˌ˥΅SĢƄϮЭeӦÂˬ͢S30N ΔpH UľȿSѵͦBk

PЄ2ilN0k PGR5 Vr~µrO

ƀo P700 Ӆŗ¬P@NԛŐBkh1SR

j&Ϥ˚P@N FLV 5ÄɆсSRKN@`KH

VŻ˴U˻ЍoɷHR0 51)'<li¼ĔU

PʅǭBk×dŴЍO/k'FLV Shk O2 һğ

P700 Ӆŗ¬5ĢŹɞͯ͢ỦŵUÉOQ

&Uh1Rҫŗ΅ƽҷoҚKN6HU4Зƃ̶0'ŮɈOU Km đV3hF 1 µM O/kHc

48, 49)r~µreϮЭ&F@N̖ӮSͯɑBk
6. P700 Ӆŗ¬V&àȋ΅ƽŐĢͮƹOd

RQSPKNVӂŹ5К4KHU4d
@lR0

29)

'ǢӮ& FLV ˿ʋˢV̖̝

BkP PSII ӹǓñҳ̬ɎUԍЦRöÃoΡB

29)

Hi>lkĢĘǥoͭъBkHcUĵǓưΊO'/k ͒Ȅi 2)5Ομ@H)̬ɎӅϜShk PSI ĢĘ<lVĢƄƀUĹ΁ŮɈO/k¶Uy|ǥUћǵ̥*OV&ԔƆ¶Ģ (10 ~ 1,000 ms)

¼ŮɈ5̖ÉOŜĵSԛŐ@N0R0&

10ĢŹɞΛθ 27 (1) 2017
photons m−2 s−1 (ǃӨĢȰȤUϗ 8 ŇβȤ) P͒Ȅ͛ǳShj&½ΕS@NĢŹɞӹǓñҳϕoӹǓO̿HB<P5O6k'<lV&ˬ͢5ͮƹi 2)Uʣ̥ (20,000 µmol photons m−2 s−1) S̒]·S>i>lHӮUұņĢͮƹo˶ĐBkύ

NǶ>0×5ɺ;ilk'͒Ȅi5ͱ0HĢȰȤĄRʣ̥O/k'<Uʣ̥OV&ĢŹɞUԛŐS

OV&ϗ 1 ʴӜUΙ0ӜO PSI ĢĘǥ5ͯAk'

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Received March 01, 2017; Accepted March 14, 2017;UɎѿъʬS˄ŏO/k'Published April 30, 2017

7. 3njSԸP700 ӅŗPĢŹɞŪЄʜͦ
1. Asada, K. (2006) Production and scavenging ofreactive oxygen species in chloroplasts and theirfunctions. Plant Physiol. 141, 391−396.
2. Sejima, T., Takagi, D., Fukayama, H., Makino, A. andMiyake, C. (2014) Repetitive short-pulse light mainly
inactivates photosystem I in sunflower leaves. Plant
Cell Physiol. 55, 1184−1193.
3. Bowler, C., Van Montagu, M. and Inzé, D. (1992)
Superoxide dismutase and stress tolerance. Annu. Rev.
Plant Physiol. Plant Mol. Biol. 43, 83−116.
4. Kok, B. (1956) On the inhibition of photosynthesis byintense light. Biochim. Biophys. Acta 21, 234−244.
5. Murata, N., Takahashi, S., Nishiyama, Y. andAllakhverdiev, S.I. (2007) Photoinhibition ofphotosystem II under environmental stress. Biochim.
Biophys. Acta Bioenerg. 1767, 414−421.
6. Sundby, C., Chow, W.S. and Anderson, J.M. (1993)
Effects on photosystem II function, photoinhibition, andplant performance of the spontaneous mutation ofserine-264 in the photosystem II reaction center D1
protein in triazine-resistant Brassica napus L. Plant
Physiol. 103, 105−113.
7. Neidhardt, J., Benemann, J.R., Zhang, L. and Melis, A.
(1998) Photosystem-II repair and chloroplast recovery
from irradiance stress: relationship between chronic
photoinhibition, light-harvesting chlorophyll antenna

 RE P700 VӅŗ>lkɆс5/kU4&FlVĢŹɞͯ͢5)ǜɅ@N*ĢŹɞoб1HcO/kPΫHJVЄ2k'P700 ӅŗVĢŹɞSΌ
ʄȵԂ@R0'ĢŹɞ̬Ɏԩ/k0VӹǓñҳ̬
ɎԪP P700 ӅŗUӜSVʬΟR΍ӝVR8 31)&FUёɵSr~µrOV&ö CO2 ͮƹS

>i>lNŻAh1SӹǓñҳ̬Ɏ5öÃ@N
08ÉOd P700 oӅŗBkϟЌPӅŗ@R0ϟ

Ќ5üĴ>lk 14)'dJmp P700 Ӆŗ¬o˿ʋ@NdĢŹɞ̬ɎFUdU5öÃBk×
VR0 14)'ΫHJUΛθ¶¼£V P700 ӅŗP
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N P700 ӅŗUͯͭ΅RɗЁM:oбKH 2)'F
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ĬӀS3:k̬ɎӅϜͯɞoˊ͙Sӟ9HcS
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Miyake, C. (2016) Superoxide and singlet oxygen
produced within the thylakoid membranes both cause
photosystem I photoinhibition. Plant Physiol. 171,1626−1634.
20. Mehler, A.H. (1951) Studies on reactions of illuminatedchloroplasts. Arch. Biochem. Biophys. 33, 65−77.
21. Youngman, R.J. and Elstner, E.F. (1981) Oxygen
species in paraquat toxicity: the crypto-OH radical.
FEBS Lett. 129, 265−268.
22. Polm, M. and Brettel, K. (1998) Secondary pair charge
recombination in photosystem I under strongly
reducing conditions: temperature dependence andsuggested mechanism. Biophys. J. 74, 3173−3181.
23. Cazzaniga, S., Li, Z., Niyogi, K.K., Bassi, R. andDall’Osto, L. (2012) The Arabidopsis szl1 mutant
reveals a critical role of β-carotene in photosystem I
photoprotection. Plant Physiol. 159, 1745−1758.
24. Sonoike, K., Terashima, I., Iwaki, M. and Itoh, S.
(1995) Destruction of photosystem I iron-sulfur centersin leaves of Cucumis sativus L. by weak illumination atchilling temperatures. FEBS Lett. 362, 235−238.
25. Zhang, S. and Scheller, H.V. (2004) Photoinhibition ofphotosystem I at chilling temperature and subsequent
recovery in Arabidopsis thaliana. Plant Cell Physiol.
45, 1595−1602.
26. Klughammer, C. and Schreiber, U. (1994) An improved
method, using saturating light pulses, for thedetermination of photosystem I quantum yield via
P700+-absorbance changes at 830 nm. Planta 192,261−268.
27. Laisk, A. and Oja, V. (1994) Range of photosynthetic
control of postillumination P700+ reduction rate insunflower leaves. Photosynth. Res. 39, 39−50.
28. Miyake, C., Miyata, M., Shinzaki, Y. and Tomizawa, K.
(2005) CO2 response of cyclic electron flow around PSI(CEF-PSI) in tobacco leaves-relative electron fluxes
through PSI and PSII determine the magnitude of
non-photochemical quenching (NPQ) of Chl
fluorescence. Plant Cell Physiol. 46, 629−637.
29. Shimakawa, G., Ishizaki, K., Tsukamoto, S., Tanaka,
M., Sejima, T. and Miyake, C. (2017) The liverwort,
Marchantia, drives alternative electron flow using aflavodiiron protein to protect PSI. Plant Physiol. doi:
10.1104/pp.16.01038.
30. Shaku, K., Shimakawa, G., Hashiguchi, M. and Miyake,
C. (2016) Reduction-induced suppression of electron
flow (RISE) in the photosynthetic electron transportsystem of Synechococcus elongatus PCC 7942. Plant
Cell Physiol. 57, 1443−1453.
31. Takagi, D., Ishizaki, K., Hanawa, H., Mabuchi, T.,
Shimakawa, G., Yamamoto, H. and Miyake, C. (2017)
Diversity of strategies for escaping reactive oxygen

size and photosynthetic productivity in Dunaliellasalina (green algae). Photosynth. Res. 56, 175−184.
8. Nishiyama, Y., Allakhverdiev, S.I., Yamamoto, H.,
Hayashi, H. and Murata, N. (2004) Singlet oxygen
inhibits the repair of photosystem II by suppressing thetranslation elongation of the D1 protein in Synechocystis
sp. PCC 6803. Biochemistry 43, 11321−11330.
9. Tyystjärvi, E. (2008) Photoinhibition of photosystem IIand photodamage of the oxygen evolving manganesecluster. Coord. Chem. Rev. 252, 361−376.
10. Krause, G.H., Köster, S. and Wong, S.C. (1985)
Photoinhibition of photosynthesis under anaerobic
conditions studied with leaves and chloroplasts ofSpinacia oleracea L. Planta 165, 430−438.
11. Terashima, I., Funayama, S. and Sonoike, K. (1994) The
site of photoinhibition in leaves of Cucumis sativus L. at
low temperatures is photosystem I, not photosystem II.
Planta 193, 300−306.
12. Munekage, Y., Hojo, M., Meurer, J., Endo, T., Tasaka,
M. and Shikanai, T. (2002) PGR5 is involved in cyclic
electron flow around photosystem I and is essential forphotoprotection in Arabidopsis. Cell 110, 361−371.
13. Suorsa, M., Jarvi, S., Grieco, M., Nurmi, M.,
Pietrzykowska, M., Rantala, M., Kangasjarvi, S.,
Paakkarinen, V., Tikkanen, M., Jansson, S. and Aro,
E.M. (2012) PROTON GRADIENT REGULATION5
is essential for proper acclimation of Arabidopsis
photosystem I to naturally and artificially fluctuating
light conditions. Plant Cell 24, 2934−2948.
14. Shimakawa, G., Shaku, K. and Miyake, C. (2016)
Oxidation of P700 in photosystem I is essential for thegrowth of cyanobacteria. Plant Physiol. 172,1443−1450.
15. Kudoh, H. and Sonoike, K. (2002) Irreversible damage
to photosystem I by chilling in the light: cause of thedegradation of chlorophyll after returning to normal
growth temperature. Planta 215, 541−548.
16. Zivcak, M., Brestic, M., Kunderlikova, K., Sytar, O.
and Allakhverdiev, S.I. (2015) Repetitive lightpulse-induced photoinhibition of photosystem I
severely affects CO2 assimilation and photoprotectionin wheat leaves. Photosynth. Res. 126, 449−463.
17. Satoh, K. (1970) Mechanism of photoinactivation inphotosynthetic systems II. The occurrence andproperties of two different types of photoinactivation.
Plant Cell Physiol. 11, 29−38.
18. Sonoike, K. (1996) Degradation of psaB gene product,
the reaction center subunit of photosystem I, is caused
during photoinhibition of photosystem I: possible
involvement of active oxygen species. Plant Sci. 115,157−164.
19. Takagi, D., Takumi, S., Hashiguchi, M., Sejima, T. and13ĢŹɞΛθ 27 (1) 2017
S. and Aro, E.M. (2012) Steady-state phosphorylationof light-harvesting complex II proteins preserves
photosystem I under fluctuating white light. Plant
Physiol. 160, 1896−1910.
42. Huang, W., Yang, Y.-J., Hu, H. and Zhang, S.-B. (2016)
Moderate photoinhibition of photosystem II protects
photosystem I from photodamage at chilling stress intobacco leaves. Front. Plant. Sci. 7, 182.
43. Ruuska, S., Andrews, T.J., Badger, M.R., Hudson, G.S.,
Laisk, A., Price, G.D. and von Caemmerer, S. (1999)
The interplay between limiting processes in C3
photosynthesis studied by rapid-response gas exchangeusing transgenic tobacco impaired in photosynthesis.
Funct. Plant Biol. 25, 859−870.
44. Sejima, T., Hanawa, H., Shimakawa, G., Takagi, D.,
Suzuki, Y., Fukayama, H., Makino, A. and Miyake, C.
(2016) Post-illumination transient O2-uptake is driven
by photorespiration in tobacco leaves. Physiol. Plant.
156, 227−238.
45. Takagi, D., Hashiguchi, M., Sejima, T., Makino, A. andMiyake, C. (2016) Photorespiration provides the
chance of cyclic electron flow to operate for theredox-regulation of P700 in photosynthetic electrontransport system of sunflower leaves. Photosynth. Res.
129, 279−290.
46. Helman, Y., Tchernov, D., Reinhold, L., Shibata, M.,
Ogawa, T., Schwarz, R., Ohad, I. and Kaplan, A. (2003)
Genes encoding A-type flavoproteins are essential forphotoreduction of O2 in cyanobacteria. Curr. Biol. 13,
230−235.
47. Yamamoto, H., Takahashi, S., Badger, M.R. andShikanai, T. (2016) Artificial remodelling of alternativeelectron flow by flavodiiron proteins in Arabidopsis.
Nat. Plants 2, 16012.
48. Vicente, J.B., Gomes, C.M., Wasserfallen, A. andTeixeira, M. (2002) Module fusion in an A-type
flavoprotein from the cyanobacterium Synechocystis
condenses a multiple-component pathway in a single
polypeptide chain. Biochem. Biophys. Res. Commun.
294, 82−87.
49. Shimakawa, G., Shaku, K., Nishi, A., Hayashi, R.,
Yamamoto, H., Sakamoto, K., Makino, A. and Miyake,
C. (2015) FLAVODIIRON2 and FLAVODIIRON4
proteins mediate an oxygen-dependent alternativeelectron flow in Synechocystis sp. PCC 6803 underCO2-limited conditions. Plant Physiol. 167, 472−480.
50. Hayashi, R., Shimakawa, G., Shaku, K., Shimizu, S.,
Akimoto, S., Yamamoto, H., Amako, K., Sugimoto, T.,
Tamoi, M., Makino, A. and Miyake, C. (2014)
O2-dependent large electron flow functioned as anelectron sink, replacing the steady-state electron flux inphotosynthesis in the cyanobacterium Synechocystis sp.

species production within photosystem I among land
plants. Physiol. Plant. In press.
32. Nishio, J.N. and Whitmarsh, J. (1993) Dissipation of
the proton electrochemical potential in intact
chloroplasts (II. The pH gradient monitored bycytochrome f reduction kinetics). Plant Physiol. 101,
89−96.
33. Avenson, T.J., Cruz, J.A., Kanazawa, A. and Kramer,
D.M. (2005) Regulating the proton budget of higher
plant photosynthesis. Proc. Natl. Acad. Sci. U. S. A. 102,9709−9713.
34. Livingston, A.K., Cruz, J.A., Kohzuma, K., Dhingra, A.
and Kramer, D.M. (2010) An Arabidopsis mutant withhigh cyclic electron flow around photosystem I (hcef)
involving the NADPH dehydrogenase complex. Plant
Cell 22, 221−233.
35. Schreiber, U. and Neubauer, C. (1990) O2-dependent
electron flow, membrane energization and themechanism of non-photochemical quenching ofchlorophyll fluorescence. Photosynth. Res. 25,
279−293.
36. Takizawa, K., Kanazawa, A. and Kramer, D.M. (2008)
DepletionofstromalPi
induceshigh‘energy-dependent’ antenna exciton quenching (qE) by
decreasing proton conductivity at CFO-CF1 ATPsynthase. Plant Cell Environ. 31, 235−243.
37. Armbruster, U., Carrillo, L.R., Venema, K., Pavlovic,
L., Schmidtmann, E., Kornfeld, A., Jahns, P., Berry,
J.A., Kramer, D.M. and Jonikas, M.C. (2014) Ion
antiport accelerates photosynthetic acclimation influctuating light environments. Nat. Commun. 5, 5439.
38. Johnson, X., Steinbeck, J., Dent, R.M., Takahashi, H.,
Richaud, P., Ozawa, S.-I., Houille-Vernes, L.,
Petroutsos, D., Rappaport, F., Grossman, A.R., Niyogi,
K.K., Hippler, M. and Alric, J. (2014) Proton gradient
regulation 5-mediated cyclic electron flow under ATPor redox-limited conditions: A study of ΔATPase pgr5
and ΔrbcL pgr5 mutants in the green alga
Chlamydomonas reinhardtii. Plant Physiol. 165,
438−452.
39. Shaku, K. and Miyake, C. (2016) ĢŹɞP O₂ UΛθ4iт2N6H, ʢHRӹǓñҳľȿ­z¬UǕƤ. ĢŹɞΛθ 26, 10−22.
40. Bergner, S.V., Scholz, M., Trompelt, K., Barth, J.,
Gäbelein, P., Steinbeck, J., Xue, H., Clowez, S., Fucile,G., Goldschmidt-Clermont, M., Fufezan, C. andHippler, M. (2015) STATE TRANSITION7-dependent
phosphorylationismodulatedbychanging
environmental conditions, and its absence triggers
remodeling of photosynthetic protein complexes. Plant
Physiol. 168, 615−634.
41. Grieco, M., Tikkanen, M., Paakkarinen, V., Kangasjarvi,

14ĢŹɞΛθ 27 (1) 2017
Plant. In press.
57. Shimakawa, G., Matsuda, Y., Nakajima, K., Tamoi, M.,
Shigeoka, S. and Miyake, C. (2017) Diverse strategiesof O2 usage for preventing photo-oxidative damage
under CO2 limitation during algal photosynthesis. Sci.
Rep. 7, 41022.
58. Raven, J.A., Osborne, B.A. and Johnston, A.M. (1985)
Uptake of CO2 by aquatic vegetation. Plant Cell
Environ. 8, 417−425.
59. Jordan, D.B. and Ogren, W.L. (1981) Species variationin the specificity of ribulose biphosphatecarboxylase/oxygenase. Nature 291, 513−515.
60. Eisenhut, M., Ruth, W., Haimovich, M., Bauwe, H.,
Kaplan, A. and Hagemann, M. (2008) The
photorespiratory glycolate metabolism is essential forcyanobacteria and might have been conveyed
endosymbiontically to plants. Proc. Natl. Acad. Sci. U.
S. A. 105, 17199−17204.
61. Rademacher,
N.,
Kern,
R.,
Fujiwara,
T.,
Mettler-Altmann, T., Miyagishima, S., Hagemann, M.,
Eisenhut,M.
andWeber,
A.P.M.
(2016)
Photorespiratory glycolate oxidase is essential for thesurvival of the red alga Cyanidioschyzon merolae underambient CO2 conditions. J. Exp. Bot. 67, 3165−3175.
62. Asada, K. (2009) r~µrUӅϜҶɈP̬ɎӅϜҶɈ. ĢŹɞΛθ 19, 75−80.
63. Miyake, C. and Makino, A. (2003) Water-Water Ƙ҉UҫŗPĢŹɞ¼¼P@NUӅϜļͱ. ŗǘPͯ͢ 41, 478−487.
64. Takagi, D. and Miyake, C. (2014) <l`O˿:N0
H&ҧȤѥ΅ђăSưM8&y¶s¢»w·~¸º»¡¸¼̬ɎU̒ҐPĢŹɞS3:k
O₂UȶŇ: ͯͭ΅Rɗƃ5т2k&ˌÊ\ɹb. ĢŹɞΛθ 24, 97−110.

PCC 6803, but not in the cyanobacterium
Synechococcus sp. PCC 7942. Biosci. Biotechnol.
Biochem. 78, 384−393.
51. Allahverdiyeva, Y., Mustila, H., Ermakova, M.,
Bersanini, L., Richaud, P., Ajlani, G., Battchikova, N.,
Cournac, L. and Aro, E.M. (2013) Flavodiiron proteins
Flv1 and Flv3 enable cyanobacterial growth andphotosynthesis under fluctuating light. Proc. Natl. Acad.
Sci. U. S. A. 110, 4111−4116.
52. Gerotto, C., Alboresi, A., Meneghesso, A., Jokel, M.,
Suorsa, M., Aro, E.-M. and Morosinotto, T. (2016)
Flavodiiron proteins act as safety valve for electrons inPhyscomitrella patens. Proc. Natl. Acad. Sci. U. S. A.
113, 12322−12327.
53. Zhang, P., Allahverdiyeva, Y., Eisenhut, M. and Aro,
E.M. (2009) Flavodiiron proteins in oxygenic
photosyntheticorganisms:
photoprotectionofphotosystem II by Flv2 and Flv4 in Synechocystis sp.
PCC 6803. PLoS One 4, e5331.
54. Shimakawa, G. (2015) r~µrUĢŹɞS3:kӅϜļͱ. ĢŹɞΛθ 25, 16−21.
55. Shimakawa, G., Akimoto, S., Ueno, Y., Wada, A.,
Shaku, K., Takahashi, Y. and Miyake, C. (2016)
Diversity in photosynthetic electron transport under[CO2]-limitation: the cyanobacterium Synechococcus sp.
PCC 7002 and green alga Chlamydomonas reinhardtii
drive an O2-dependent alternative electron flow andnon-photochemical
quenchingofchlorophyll
fluorescence during CO2-limited photosynthesis.
Photosynth. Res. 130, 293−305.
56. Hanawa, H., Takagi, D., Shimakawa, G., Nohira, K.,
Sejima, T., Shaku, K., Makino, A. and Miyake, C.
(2017) Land plants drive photorespiration as higher
electron-sink: Comparative study of post-illuminationtransient O2-uptake rates from liverworts toangiosperms through ferns and gymnosperms. Physiol.

P700 oxidation system protects PSI against photo-oxidative damage: Why are plants safe in
the light?
Ginga Shimakawa and Chikahiro Miyake*
Graduate School of Agricultural Science, Kobe University15ĢŹɞΛθ 27 (1) 2017

研究紹介
硫化水素を電子供与体とする電子伝達系遺伝子の発現誘導：
転写制御タンパク質 SqrR の同定とその硫化水素応答機構§
東京工業大学

大学院生命理工学研究科
清水 隆之*

誕生初期の光合成では硫化水素を電子供与体としていた可能性が高い。現存する酸素非発生型光合成細菌でも硫化水素などのイオウ化合物を電子供与体として光合成を行うものが多く存在する。私は硫化水素に応答したシグナル制御機構を解明するために、紅色光合成細菌 Rhodobacter capsulatus から
硫化水素応答性の転写因子 SqrR を同定し、その分子機構を解析した。その結果、SqrR は、硫化水素
由来の活性イオウ分子種によって 2 つの Cys 残基の間で分子内テトラスルフィド結合が形成されることで、オペレーター領域への DNA 結合能が弱まることがわかった。これより、硫化水素依存的な光合成の転写制御の分子機構が明らかとなった。

1ԮVAcS12

ilN0k'FUHc&Ξŗ̖ϜUɈτ˻˳V& ĢŹɞ5ƦͬÂSњͯ@HĹˈǓӪOV&̖ĹˈUĢŹɞUҫŗSPKNӉсIKHPЄ2

hjdӅŗ5ǧʭRΞŗ̖ϜRQUtyuŗŹ

ilk'͢oӹǓāÅùP@NĢŹɞ5бnlN0HP

 ΫV&Ξŗ̖ϜUɈτ˻˳oĵǓ·¥¶Oъʬ

Є2ilN0k'ǢӮ&Ξŗ̖ϜoӹǓāÅùP

BkHcS&R. capsulatus oͱ0N&Ξŗ̖ϜɈτɎUҎįƙǓ SqrR oŻǠ@H'ˌεOV&SqrR@NĢŹɞoб1ϟУγ5ͫƤdǀʙŻǠ>lU DNA ϤŹЍPҎįľȿ̬Ɏ5 Cys ̋ưUN0k'ĢŹɞӹǓñҳϕOΞŗ̖ϜUӅŗSӝnkӄϜV&Sulfide:quinone oxidoreductaseԩSQRԪ
1)& Flavocytochromecsulfide´¶¡sϤŹȳɞShKNƽŗBk<Po

dehydrogenaseΡB 6)'

ԩFccABԪ2)&Sox ӄϜЀ 3)5ΗilN3j&ͣS

2ԮΞŗ̖ϜɈτɎҎįƙǓ SqrR UŻǠ

SQR 5ËсRȶŇoɰKN0kPЄ2ilN0
4)k '<liӄϜUΛθ4i&Ξŗ̖ϜĂǕ΅R

 Ξŗ̖ϜɈτɎUľȿƙǓUŻǠo&ҹñǓʑĢŹɞӹǓñҳϕV4Rjʬi4SRKN6H'

ü5ǧʭR R. capsulatus oͱ0HԆҹñǘ΅ɥΞŗ̖ϜUӅŗSӝnkӄϜU΁ͫV&Ξŗ̖Ϝ̥ShKNбKH 7)'Ξŗ̖ϜO΁ͫћǵoű:͑ȤSɈANľȿ>lN0kPЄ2ilk'ǢӮ&

k sqr ҹñǓU΁ͫƽŗoɸ˵SBk<PO&ΞϘЛĢŹɞϟУ R. capsulatus S30N&SQR U΁ŗ̖ϜӽəűɎUƽͽˢUšӴoѓaH'ƽͽˢͫ5Ξŗ̖ϜĳͭOћǵ>lk<P5ƵƁ>lUҸɮUHcS&sqr £¸®¼¼UÃ̭S lacZ5)

N0k '@4@&Ξŗ̖ϜɈτɎUľȿƙǓ5ҹñǓoаŹ@H£´«ԩsqr::lacZ-fusionԪ

ŻǠ>lN3iC&і@0ľȿ˻˳VˊIʬi4oüɞ@H'sqr £¸®¼¼5Ξŗ̖ϜtyºOR0'ɍ͏Rͯ͢ҫŗSV&ҹñǓ˳ҨUƽŗ

ԩHS-Ԫͣͽ΅SɈτBk<PoΟљBkHcS&

hjdҹñǓľȿϕUƽŗUʣ5ӉсIPЄ2

sqr::lacZ-fusion oĈ˄Bk R. capsulatus o&Ǌ̕`HVǑ̕ƯԖ˒ëOΞŗµu¬ԩNa2SԪĳ

ͭ@&ŸϟЌU β {´~¼̬Ɏoɸ˵S&ξ 7 ƘʦˌĢŹɞǘðº©u¬ ©¼΁жѽűѽѥʜ*ҩϥġ   

§sqr £¸®¼¼̬ɎUʴӜƽŗo̾Ǡ@H'F
16

ĢŹɞΛθ 27 (1) 2017

Ɲ 1. sqr-lacZ fusionShkβ{´~¼̬Ɏ
Ǌ̕ƯԖԩAԪd@8VĢǑ̕ƯԖԩBԪO log ˈÉˈ`OͯЊ>D&úd̸Ŋ@R0ԩ΃ƗщԪd@8Vϡ͑Ȥ 0.6 mM Na2Sԩ΃ÁщԪo̸Ŋ@HP6o 0
ĵP@N&>iS 0.5&1&2&4 ʴӜƯԖ@HP6U
β {´~¼̬Ɏo̾Ǡ@H'

  



Ɲ 2. ŻǠ@HΞŗ̖ϜɈτɎUľȿƙǓSqrR U
˿ʋˢS3:ksqr£¸®¼¼̬Ɏ
Control V sqr::lacZ-fusion oĈ˄Bkӊͯˢ'ћǵ
o4:N0R0ϟЌԩ΃ЛԪPϡ͑Ȥ 0.6 mM Na2S
Shkћǵo 4 ʴӜ4:HϟЌԩ͔ЛԪOU̾ǠϤ˚oΡB'

UϤ˚&sqr ҹñǓVΞŗ̖ϜtyºԩNa2S V̵̖̈́ÉOVËP@NΞŗ̖ϜtyºP@NǕƤ
BkԪSɈτ@NҎį5ћǵ>lk<P5n4K

 ˂ҝ&ԥЛ¢uͬУ Staphylococcus aureus S

HԩƝ 1Ԫ'30N&Ξŗ̖ϜɈτɎUҎįƙǓ CstR 5ŻǠ

 R. capsulatus ӊͯˢԩWTԪV β-{´~¼>lH 8)'CstR V CsoR/RcnR ¡q«µ¼Sǿ@&ԩlacZ ҹñǓͰ͢ԪodHR0Hc´~¼âƘŻǠ@H SqrR V ArsR/SmtB ¡q«µ¼Sǿ

oƍ½U͕Ϝ̀P@HƯƦOVͯЊBk<P5

BkHc&CstR P SqrR Vĥ8ͽRkº~ѿ

O6R0'½ʣ&sqr £¸®¼¼ľȿÃS lacZ

O/k'R. capsulatus d cstR §®¸o 2 LɷLoҩϤ@H sqr::lacZ-fusion Ĉ˄ˢV&Ξŗ̖ϜǕ

5&FlGl formaldehyde əűɎµ£·¼

ƤÃO lacZ U΁ͫ5ћǵ>lkHc´~¼

FrmR P¶əűɎµ£·¼RcnR Sҝo͕Ϝ̀P@NͯЊ5ŴЍPRk'R. capsulatus

ϴO/k<P5ϕϧъ˗hjn4KH 9,10)'ǢӮ

U sqr::lacZ-fusion Ĉ˄ˢo EMS OĳͭBk<P

S&2 LU cstR §®¸Uš½˿ʋ3hYØӉ˿

Oκ͙ƽͽoћ΁@Hȼ&Ξŗ̖ϜӽǕƤÃO´

ʋˢV&ǝĥRΞŗ̖ϜɈτЍoĈɷ@N0H'~¼oƍ½U͕Ϝ̀P@HƯƦoͱ0NǊ<lhj&R. capsulatus S30N&SqrR 5ËсR̕ƯԖ@H'<U˒ëOVΞŗ̖ϜӽǕƤÃOd
lacZ 5΁ͫBkh1SRKHϟЌԩΞŗ͢ӽəűΞŗ̖Ϝº¼O/kPЄ2ilH'ɎˢԪUaͯЊ5ŴЍO/k'<U~µ¼º3ԮSqrR UΞŗ̖Ϝ˭Η˻˳Uъ˗

Shj&κ͙ƽͽoћ΁@H 108 ČUϟЌ4i

ϗ 30 ˢUΞŗ̖ϜӽəűɎƽͽˢ5šӴO6H'
ȾilHƽͽˢU1J 3 ˢSL0N&

 SqrR 5Ξŗ̖ϜSɈτBkĵǓ˻˳oъʬB

kHcS&ǁБУOǁӋ΁ͫ>DH R. capsulatus
SqrR Uµººº~ѿoϒо@&F

sqr::lacZ-fusion U sqr £¸®¼¼ԉƭSƽͽ5Uͯŗǘ΅Ɏѿoъ˗@H'DNase I ¡£µR0<PoΟљ@Hȼ&¬ DNA Uµ¼ºShKNƽͽφɣUŻǠoбKH'FUϤ˚&пʙUĎмҹñǓ5ŻǠ>lH'ĎмҹñǓoFºrtShKN sqr £¸®¼¼ԉƭS3:k SqrR UϤŹԉƭơǠ@HP<m&-10 PҎįӛǍ͖oſbԉƭSϤŹBk<P5n4j&lGl˿ʋ>DHϤ˚&˿ʋShKNΞŗ̖Ϝӽ

SqrR 5µ£·¼O/k<P5ʬ΃PRKH'ǕƤÃOU sqr U΁ͫ5ԟ8RkҹñǓ5 1 LŻ

SqrR VΞŗ̖ϜSɈANy¦·¼¼\U DNA

Ǡ>lHԩƝ 2Ԫ'˿ʋShKNɪľЍ5ǆnlϤŹчƆɎoƽŗ>DkPЄ2ilHHc&̛Ǡ

k<P4i&<UҹñǓV sqr ҹñǓUµ£·@Hy¦·¼¼ԉƭSǰBk DNA ϤŹчƆɎ

¼O/kPЄ2il&sqrRԩsqr repressorԪPƅo  ¶  ¡  r   t S h K N ̾ Ǡ @ H ' S.

ż@H'

17

ĢŹɞΛθ 27 (1) 2017
Cys107ԪoɷJ&FU1JU 2 LԩCys41 P Cys107Ԫ

aureus CstR VΞŗ̖ϜtyºSVɈτDC&̬

VͽRkϟУγU SqrR §®¸ӜSĈǕ>lNɎtyuĵǓγԩRSS; Reactive sulfur speciesԪS

0H'<li Cys ̋ư5 RSS ɈτSӝnk4˭

8)

ɈτBkPƵƁ>lN0k 'RSS V&Ӆŗ>lёBkHcS&Ÿ Cys ̋ưo Ser ̋ưSϽʈ@H

HӽȘSŮɈɎUԟ0tyuſ˄ĵǓγO&ͯù͖ƽͽ SqrR oͱ0NŻ˴Uъ˗oбKH'FU

ĬOVΞŗ̖ϜГҌd@8VΞŗ̖ϜPĻUĵǓPUŮɈShKNͰͯ>lk

11,12)'ˌΛθOV&Ϥ˚&C9S SqrR V DNA PUϤŹчƆɎVĥùP@Nǁ68öÃ@H5&RSS SǰBkɈτVĈɷ¶yºPΞŗ̖ϜUŮɈShKNͰͯ>@N0HԩƝ 3BԪ'½ʣ&C41S SqrR P C107S SqrRlk RSS O/k¶yº¼¶¡s
ԩGSSHԪo in vitro ǢԜSͱ0H'Ǒ̕˒ëÃO&V RSS ĳͭoű:Nd DNA ϤŹчƆɎVԟ0`

¼\U DNA ϤŹчƆɎo̾Ǡ@HP<m&ˢoͱ0H in vivo ъ˗Ϥ˚Pd½Е@HԩƝ 4Ԫ'`IKHԩƝ 3C, DԪ'<UϤ˚V&sqrR ͖ƽͽһğƪ SqrR P GSSH ĳͭ SqrR U sqr y¦·¼

C9S V WT Ż˴SΞŗ̖ϜtyºSɈτ@Nɪ

GSSH ĳͭShKNϤŹчƆɎ5˄ɗSöÃ@ľ5ъӤ>lH5&C41S P C107S VΞŗ̖ϜtHԩƝ 3AԪ'<U<PV¼¶¡sǕƤÃO SqrR 5y¦·¼¼4iъӴ@eB8Rk<yºǕƤÃOdɪľͥɚ5ϫɷ>lN0H'<l
hj&SqrR UΞŗ̖ϜɈτSV&RSS Shk Cys

PoΡƌ@N0k'
 RSS Shk¶ñҳSV Cys ̋ưU SH ư̋ưUċԕ5ӉсO/k<P5n4KH'
 GSSH Shk SqrR U Cys ̋ưUŗǘċԕoŻU ċ ԕ 5 ӝ Å @ N 0 k P Є 2 i l k ' R.

ǠBkHcS&LC-ESI-MS ShKN GSSH ĳͭ

capsulatus SqrR V 3 LU Cys ̋ưԩCys9, Cys41,

Ɲ 3. ¶¡rtShkSqrR Usqr£¸®¼¼\UDNAϤŹчƆɎǑ̕˒ëÃOUһğƪ SqrRԩ΃ÊԪP GSSH ĳͭ SqrRԩԦÊԪUϤŹʾϯ'A-D VFlGl&WT SqrR, C9S
SqrR, C41S SqrR, C107S SqrR UϤŹʾϯoΡB'18ĢŹɞΛθ 27 (1) 2017
H'һğƪ C9S SqrR OV&ʅǠĵǓӋ 12,296 Da
PǝĥS½ЕBk¶5ȾilH<P4i&ċԕoű:N3iC&Cys ̋ưVǝĥSһğ>l
N0k<P5кæ:ilH'>iS&GSH&GSSG&Na2S Oĳͭ@HƶŹdƽŗ5R4KH'½ʣ&

GSSH Oĳͭ@HƶŹSV&+62 Da (Mr 12,358 Da)UĵǓӋ¡5шǭ>lH'<Uƽŗԩ+62Ԫ
V&ΞԥŧǓ 2 ČUæŊԩ+64ԪP̖ϜŧǓ 2 ČU̴ǆԩ–2ԪOχǠ>lkđPh8½ЕBkHc&
Cys41 P Cys107 UӜOĵǓĬ´¶¡sϤŹoȳɞ@N0k<PoΡƌBk'Ŋ2N&+30
Da (Mr 12,326 Da)UǶ>Rƽŗdшǭ>l&ĵǓĬµ¶¡sϤŹdΡƌ>lH'ESI-MS/MSOµ£ºĳͭ@H GSSH ĳͭ C9S SqrR oĵƝ 4. sqrRŸ͖ƽͽˢS3:ksqr£¸®¼¼̬Ɏ
R. capsulatus U   ¬ Â U sqrR ҹ ñ Ǔ U 3´ ˋ ν S
FLAGo¼BkӃĸoæŊ@H'Ÿ͖ƽͽˢV
sqrR:FLAGƽͽˢodPSüɞ@H'Ɲ2PŻ˴Uɥ
ԆO̾Ǡ@H'˗@HP<m&Ż½ĵǓĬU Cys41 P Cys107 U
ӜOµ¶¡sP´¶¡sϤŹoȳɞ@N0k<P5Ρ>lH'Ż˴UϤ˚V WTSqrR OdȾilH'

 Cys41 P Cys107 5Ξŗ̖ϜĳͭShKNċԕ@HP6U C9S SqrR UĵǓӋƽŗo̾Ǡ@Hoű:k<Po in vivo OΟ4ckHcS&y¼ԩƝ 5Ԫ'<U̾ǠOV&һğƪP GSSH ĳͭU¶ưċԕѓЫU AMS Oĳͭ@Hϭº~ɯĴ^4S&һğƪ¶yºԩGSHԪ&Ӆŗƪ

̵oͱ0Nuvº¢¸sºoбKH'¶yºԩGSSGԪ&Na2S Oĳͭ@H C9S SqrR´¶¡sϤŹoȳɞ@N0k Cys ̋ưUъ˗dбKH'C9S SqrR VϟЌĬOΞŗ̖ϜV AMS OVċԕ>lR0Hc&AMS ċԕShkɈτ̬Ɏo WT Ż˴SɷLԩƝ 4Ԫ'F<Oˌъӹ̨̕ŐOUαŐȤUƽŗV&¡µ¼Uy¼¶

˗OV&SqrR UӅŗ΅Ø̀ŗǘŮɈUȵԂo˂ưUʙSUaSĂǕBk'ǰ͛U SqrR:FLAG S

ǶӢSɪ2kHcS&C9S SqrR oˌъ˗Sͱ0ǰBk Na2S ͗ĳͭº£¶UĵǓӋƺŊԩ2.4

kDaԪVĳͭº£¶UĵǓӋƺŊԩ0.8 kDaԪU
3hF 3 čO/k<P5шǭ>lHԩƝ 6Ԫ'<lhj&Ξŗ̖ϜtyºĳͭShKN¡µ¼U
y¼¶ư5 3 L4i 1 LSRkPЄ2ilH'<

Ɲ 6. in vivoOUSqrR UCys̋ưUċԕU˭ĴCys ̋ưU SH ư5 AMS ċԕo1:k<POͯAk
ӹ̨̕ŐȤ¡4i&SqrR U Cys ̋ưU¡µ¼U
SH ưʙoǠӋ@H'ϟЌoΞŗ̖ϜOĳͭBk<P
O&2.4 kDa ĵU¡5 0.8 kDa ĵU¡S̻Ƿ@H'

Ɲ 5. C9S SqrR ULC-ESI-MSъ˗šӋù C9S SqrR (12,196 Da)5 GSSH ĳͭèƿOшǭ
>lH'GSSH ĳͭOVËS´¶¡sϤŹȳɞ C9S SqrR (12,358 Da)5шǭ>lH'ǷӋԩ≤ 5%Ԫ
Uµ¶¡qtϤŹȳɞ C9S SqrR dшǭ>lH'

19ĢŹɞΛθ 27 (1) 2017ѫҖ ˌΛθo`PckS/Hj&˕ÞȋˮǁǘUƺͲΑØıʖʁSVɥŦ0=ɸǵoѼj`@H'̶
8əѫ0H@`B'`H&ˌΛθV&tºsrǁǘU David P. Giedroc ʖʁ&Carl E. Bauer ʖ

ʁ&Jiangchuan Shen ŢƼPUĨŻΛθShKNбƝ 7. SqrR ShkΞŗ̖ϜɈτɎҹñǓЀԩSulfide-responsive genes; SRGsԪUҎįľȿ˻˳U®¶

0`@H'əѫ0H@`B'
Received February 14, 2017; Accepted February 24, 2017;UǢԜ4iV&Cys41 P Cys107 U˛˺VŻǠO

Published April 30, 2017tº̋ưUӜO GSSH Shk˛˺5ȳɞ>lŪЄʜͦ

6R05&in vitro UǢԜ4i<li 2 LU1.

k<P5Ρ>lN0kHc&in vivo OdĬƤɎUΞŗ̖ϜShKNͯɞ>lH RSS ShKN SqrR

V 2 LUtº̋ưUӜO˛˺ŮɈo҃<2.

BPЄ2ilH' Ξŗ̖ϜɈτɎҎįƙǓ SqrR V&Ξŗ̖Ϝӽ

Schutz, M., Shahak, Y., Padan, E. and Hauska, G.

(1997) Sulfide-quinone reductase from Rhodobacter
capsulatus. J. Biol. Chem. 272, 9890–9894.

Chen, Z., Koh, M., Driessche, G.V., Beeumen, J.J.V.,
Bartsch, R.G., Meyer, T.E., Cusanovich, M.A. andMathews,

F.S.

(1994)

The

structureofflavocytochrome c sulfide dehydrogenase from aǕƤÃOV&һğƪP@Ny¦·¼¼ԉƭSϤpurple phototrophic bacterium. Science 266, 430–

Ź@NҎįoɪľBk'Ξŗ̖ϜǕƤÃOV&F
lSͳ˓Bk RSS ShKNĈǕ>lH Cys ̋ư

3.

ԩCys41 P Cys107ԪUӜOĵǓĬ´¶¡sϤŹ5ȳɞ>l&y¦·¼¼ԉƭPUϤŹч

432.

Ogawa, T., Furusawa, T., Shiga, M., Seo, D. andInoue,

K.

(2010)

Biochemical

studiesofasoxF-encoded monomeric flavoprotein purified fromthe green sulfur bacterium Chlorobaculum tepidumƆɎ5öÃ@&ӉпBk£¸®¼¼ԉƭ\URNA ©µ­´¼ϤŹ5ŴЍPRjҎį5̬Ɏ

that stimulates in vitro thiosulfate oxidation. Biosci.

ŗ>lk'<Uh1RĵǓ˻˳O&Ξŗ̖ϜSɈ

4.

τ@HҎįľȿoбKN0k<P5ʬi4P
RKHԩƝ 7Ԫ'

Biotechnol. Biochem. 74, 771–780.

Sander, J. and Dahl, C. (2009) Metabolism of
inorganic sulfur compounds in purple bacteria, in The

purple phototrophic bacteria (Hunter, C.N., Daldal,F., Thurnaier, M.C. and Beatty, J.Y., Eds.) pp 595–

4. 3njS5.

 ˌΛθOV&Ξŗ̖ϜĂǕ΅RĢŹɞUľȿS
ӝnkƙǓUŻǠPFUĵǓ˻˳Uъ˗oбK622, Springer, Dordrecht, the Netherlands.

Griesbeck, C., Schutz, M., Schodl, T., Bathe, S.,

Nausch, L., Mederer, N., Vielreicher, M. and Hauska,
G. (2002) Mechanism of sulfide-quinone reductase

H'FUϤ˚&Ξŗ̖ϜɈτɎUҎįƙǓ SqrR

investigated using site-directed mutagenesis andoŻǠ@&RSS ShKNȳɞ>lkĵǓĬ´¶¡sϤŹ5ҎįѢωSӉсO/k<Po

6.

ʬi4S@H'@4@&SqrR Shkľȿ˻˳5&ͮƹÉOUΞŗ̖Ϝ͑ȤƽŐSǰɈBkHcS

sulfur analysis. Biochemistry 41, 11552–11565.

Shimizu, T., Shen, J., Fang, M., Zhang, Y., Hori, K.,
Trinidad, J. C., Bauer C. E., Giedroc, D. P. andMasuda,S.

(2017)

The

sulfide-responsive

transcriptional repressor SqrR functions as a master

QU8i0ӉсO/k4Vn4iR0'ҝȝ&ͮregulator of sulfide-dependent photosynthesis. Proc.

ƹÉUɂͯ͢ЀӲS3:k­ъ˗5ɍҧSҫ

Natl.

pO0k'<liUɩвPˌΛθUͯͭǘ΅Rɞ
˚oŹnDk<PO&ͮƹƽŗSŹnDHĢŹɞ

7.

UҫŗUъʬSLR5kPˈȺ@N0k'


20

Acad.

Sci.

USA.

in

press

(doi:

www.pnas.org/cgi/doi/10.1073/pnas.1614133114)

Sganga, M.W. and Bauer, C.E. (1992) Regulatory
factors controlling photosynthetic reaction center andĢŹɞΛθ 27 (1) 201711.

light-harvesting gene expression in Rhodobacter
capsulatus. Cell 68, 945–954.

8.

S., Matsunaga, T., Yamamoto, M., Ono, K.,

CsoR-like sulfurtransferase repressor (CstR) is a

Akaike, T. (2014) Reactive cysteine persulfides andDevarie-Baez, N.O., Xian, M., Fukuto, J.M. andpersulfide sensor in Staphylococcus aureus. Mol.

S-polythiolation regulate oxidative stress and redox

Microbiol. 94, 1343–1360.

signaling. Proc. Natl. Acad. Sci. U. S. A. 111, 7606–

Osman, D., Piergentili, C., Chen, J., Sayer, L., Uson,

12.

I., Huggins, T., Robinson, N. and Pohl, E. (2016) The
effectors

andformaldehyde-responsive

sensory

regulator

sites

FrmR

of

7611

Nishida, M., Sawa, T., Kitajima, N., Ono, K., Inoue,

H., Ihara, H., Motohashi, H., Yamamoto, M.,

Suematsu, M., Kurose, H., van der Vliet, A., Freeman,andmetal-sensing variant. J. Biol. Chem. 291, 19502–
10.

Y., Kumagai, Y., Suematsu, M., Motohashi, H., Fujii,

Luebke, J.L., Shen, J., Bruce, K.E., Kehl-Fie, T.E.,
Peng, H., Skaar, E.P. and Giedroc, D.P. (2014) The

9.

Ida, T., Sawa, T., Ihara, H., Tsuchiya, Y., Watanabe,

B.A., Shibata, T., Uchida, K., Kumagai, Y. and

19516.

Akaike, T. (2012) Hydrogen sulfide anion regulates

protein allostery in the CsoR/RcnR family of

Chem. Biol. 8, 714–724.

Higgins, K.A. and Giedroc, D.P. (2014) Insights into

redox signaling via electrophile sulfhydration,” Nat.

transcriptional repressors. Chem. Lett. 43, 20–25.

The induction of an electron transport chain involved in sulfide oxidation: The identificationand sulfide-responsive mechanism of transcriptional factor SqrR
Takayuki Shimizu*
Graduate School of Bioscience & Biotechnology, Tokyo Institute of Technology

21ĢŹɞΛθ 27 (1) 2017

研究紹介
光応答素子のための光化学系 I とカーボンナノチューブの複合体形成§
1

東京理科大学大学院 理学研究科, 2 東京大学大学院 理学系研究科,
3

東京理科大学 工学部,4 東京理科大学 理学部

二井 大輔 1,*, 宮地 麻里子 2 嶋田 友一郎 3, 野沢 陽佑 4, 伊藤 雅浩 1, 本間 芳和 1,
池平 秀 2, 山野井 慶徳 2, 西原 寛 2, 鞆 達也 1
近年、光化学系 I (PS I) の光合成反応を活用したデバイス作製が盛んに行われている。これは光合成
反応の初期過程において、PS I 電子伝達反応の量子収率が 100%に近いことを利用するものである。
本研究では、導電性に優れた単層カーボンナノチューブ （SWNT） と PS I の複合体を作製し、PS I
を励起することによって生じた電子を SWNT に伝達し、光エネルギーを電流に変換する素子を作製することを目的とした。本研究で用いた PS I は、還元側に遺伝子組換えにより炭素材料に特異的に結合する配列を挿入し、配向性をもたせることにより、効率よく電子を取り出すことを可能にしている。
野生株の PS I との比較により、遺伝子改変した PS I は SWNT と複合体を形成し、光応答素子として機能することを紹介する。

1ԮVAcS34eº~ѿUh1RͯùĵǓPпŹùoȳɞ ͕ϜˏʠV&̀ÆçϜǓU˳ɞсϜP@N˄ͱ

O6k<P5ΗilN0k 3-8)'SWNT жӾPRŴЍɎoίcN0k'͕ϜˏʠP@NV&´º~ѿӜUϤŹ­z¬SL0NV08L¡qt&¡´¼·º&t°®º&z¼¨º

4ƵƁ5/k'`Cŗǘ΅RϤŹUĀP@NV±¼¢ (CNT) F@N´¡vºπ5Ǖº~ѿĔӖPжӾoċԕ@H SWNT ӜU΍ÙƤBk5&ˌΛθOVšȀz¼¨º±¼¢

üͱShkdU5/k 9,10)'`HåUĀP@N&(Single-Walled Carbon Nano Tube: SWNT) oͱ0

SWNT жӾPº~ѿUͿ̖Ɏr«ӅĔӖ˳Ҩo@N3j&FU˳ҨVzt´¶ɸʙShj@R5i&Ϳ̖Ɏ΍Ùüͱo̬ͱBkʣ̥V&̛Ǡ>l&ӌǿɎPŞǵùɎUQJi4UɎѿo

SWNT жӾPº~ѿUƀΓʣŽoº

H'SWNT U˳ҨV&½˙U´¡vºoÊcHӜUͿ̖Ɏ΍ÙüͱShkdU5/k 11,12)'@4

ɷL'SWNT UͣɃP@N&¡·|¢¶OȰӿ¸¼¶Bk<P5Ĵ˓R0ƏԌ͖5/k'пŹùŗUĻUr£¸¼P@N&SWNT жӾSNͯù

O/j&`HǵӹˏʠP@N½Й΅SÿnlN0kӑP̒Ґ@N&ϗ 1000 čUԟ0ӹ̭ǪȤІɎĵǓӜUȰ0΍ÙüͱoļͱBkʣ̥5/k'<ԩԟǵӹɎԪ o˄@N0kHc&º¼eӹ

Uʣ̥U½LP@N&biotin–streptavidin UĵǓљǓϜǓπ\UɈͱ5Є2ilN0k'@4@R5

ѬЍShkŗǘ΅RŮɈo̬ͱ@HĀ5/k 13)'i&͕ϜˏʠVÄ̈́ɎO/j&<U͖5ɈͱUӮ Ģŗǘϕ (PS) I Vr~µr&Эԏ&UƏԌ͖O/KH'FUÉO&ҝȝͯùĵǓoԟπˬ͢ÉSǕƤBkВº~ѿпŹùO/

SWNT UĵʗńP@Nͱ0k<POŴ̈́ŗBkj&ĢԛŐShkӹǓñҳoб1'F@N'`H SWNT V DNA

plastocyanin&d@8V cytochrome c6 4iӹǓoξ 7 ƘʦˌĢŹɞǘðº©u¬ ©¼΁жѽűѽѥʜ*ҩϥġ   

L'<liUͣɎ4i&tUĢӹƽʈϜ

ǀ8UΛθƵƁ5/k

1,2)

ű:Űj ferredoxin \ŏͨh8ñҳBk˻˳od§

22ĢŹɞΛθ 27 (1) 2017
Ser–Ala–Leu 4iRk 12 r«ӅӃĸoŻǠ@ǓU˳ɞсϜP@NUҶŹɎ5тĴ>lN0k
14-17)

H'<U CNTBP Ӄĸo Synechocystis sp. PCC 6803'

U PsaE ¶¼£Ӏõ Gly55–Val56 UӜSɽĤ@H

 PS I пŹùVr~µr4iÁӋùPƽͽùo΍ŻϢaʈ2ShjȾH 27) ԩƝ 1aԪ'

@NšӴ>lkUSǰ@N&ԟπˬ͢4iVšӋ

DNA Uµ¼³ºV&PsaE UÇĔU£´

ùP@NšӴ>lk'Ǌ͞Ɏr~µr
Thermosynechococcus elongatus F@Nԟπˬ͢U

tª¼oſpI PCR ̥hjΟљ@H (data notԟĵъЍOƵƁ>lN0k<P4i&ĵǓ·¥¶ º¸¼¶ŬYҹñǓʓƽ@H

shown)'

Pea sativum 4išӴ>lH PS I UϤʹ˳ҨV&
UŮɈͭъ3hYʑü5ŴЍO/k

18-20)

Synechocystis sp. PCC 6803 ˢV 8 L ¨¶SN'ҝȝU

BG11 O Ư Ԗ o б K H ' Ư Ԗ ͮ ƹ V 20 µmolҹñǓȋǘ3hY~¸¼U΁ǾShphotons m−2 s−1 UЯĢ͓ÃSNǣ̽Oб0&ι̕

j&PS I V΃ӌ&ӌϐǓ/k0Vyº£Po¡s¶¼ (Millex filter, Millipore, USA) oҦ0KHåUˏʠPпŹùŗ>Dk<PSh&Ā2W&ӹ

@NāϦ@H'FlGlUˢ4iU PS I пŹùǓñҳϜǓ/k0V̖Ϝͯɞ\UɈͱ5ˈȺ>UšӴϒоV&´tВoŴ̈́ŗȼ&ӥtyjϜǓP@N̬ͱ>lN3j
lN0k

14-17)

15,21,22)ºÝʈ~¸ª´¡s¼oͱ0NбKH 28)'`' ˌΛθOͱ0H&PS I P SWNT UпŹùŗV&C&ϟЌo 10 mM MgCl2, 5 mM CaCl2 F@N 25%

ͽ΅SϤŹBkӃĸoã@Nȳɞ>lk

ethanesulfonic acid (MES)-NaOH buffer (pH 6.0)(w/v) glycerol oſb 50 mM 2-(N-morpholino)

Phage display ̥ShjŻǠ@H CNT Sǰ@Nͣ
(CNTBP: CNT Binding Peptide)'ˌΛθOV&<

ÉSNɝ͐@&FUȼ{´¼oͱ0NUӃĸo PS I UһğĔSɽĤ@H'<U PS I SΝΜ@H'´tВVҵɅĵӴShjƘů@&FUȼ&2% n-dodecyl-β-D-maltoside (DM) oͱ0ɽĤ@HӃĸV&CNT Sǰ@Nԟ0чƆɎoN&277 K O 10 ĵӜŴ̈́ŗoбKH'ҵɅĵӴ

ɷKN0kHc&SWNT жӾSӃŽɎo˄@NϤ

ȼ&FǗ̵UÂ̹o 50 mM Tris-HCl (pH 7.2),

ŹBkPˈȺ>lk'ˌΛθOV&ǢԜˏʠP@

0.04% DM, 30 mM NaCl SNȜдŗ@HӥtyºNr~µr Synechocystis sp. PCC 6803 oÝʈz´¬Sā@H (DEAE Toyopearl, TOSOH,

ÿͱ@&CNTBP V PsaE ÉǅæҝU¶¼£ӀõS

ɽĤ@H'<UҸǠͭͳV&PsaE ҝĕS¡v·

Japan)'šӴ PS I V&NaCl (30–400 mM) ͑Ȥŕ|º5ϤŹ@ӹǓʁÅSӝnKN0k<P

ӃShjz´¬4ḯĴ@H'5Ϥʹ˳Ҩ4iΡƌ>lN0kHcO/k'ȽK Ɲ 1b Vϒо>lH PS I U SDS-PAGE oΡ@N0k'ƝÉU Lane 1 Vº¸¼¶ PS I& Lane

N&SWNT VĢ͛ǳShj PsaE SӰʄ@H PsaC

2 V CNTBP o PsaE SɽĤ@H PS I (èȼ PS

SϤŹ@N0k Fe-S ~´¼4iӹǓoΌʄI-CNTBP PƄ[) oFlGlΡB'FUϤ˚&PS

ű:Űk<P5ˈȺO6k'

I-CNTBP S30N PsaE Uº5º¸¼¶ ӹ˯ÂSƞǠŗ>lH PS I 4iUĢӹ̭̾Ǡ

PS I P̒]r£¡o@N3j&PsaE Sͣ

V&˴(RĢŹɞͯ͢eˏʠS30Nǀ8UΛθ
5R>lN6H

23-25)

ͽ΅RϤŹӃĸ5ɽĤ>lN0k<P5'ˌΛθOVӃŽɎoɷHD

SDS-PAGE 4idΟљO6H'

H PS I o SWNT ӹ˯SϤŹ>D&Ģ͛ǳShjӹǓṶlo˭ĴBk<PSɞŉ@H

26)'
3Ԯƀů¦~¶̾ǠShk SWNT P PSI Uп

2. z¼¨º±¼¢ϤŹӃĸU΁ͫP PS I

ŹùŗUΟљ

 PS I P SWNT UпŹùŗUΟљV&ƀů

\UɽĤ ѥʜĨЦЅUȇͲiV&phage display ̥oͱ0¦~¶U̾Ǡ&ҤұƪӹǓԍɂӗшǭ(TEM)&
SWNT U¡x¶«ºShjђă@H'

N 10 ČU phage 4i CNT Sǰ@Nԟ0ϤŹɎ
9ƀů¦~¶Shk̾ǠV&PS I P SWNT U

oΡ@H His–Met–Gly–Leu–Thr–Lys–Ile–His–Tyr–23ĢŹɞΛθ 27 (1) 2017

Ɲ 1. PS I \Uz¼¨º±¼¢ϤŹӃĸ (CNTBP) UɽĤӀõ (a) P SDS-PAGE Shkϒо@H PS I
UϢɞUΟљ (b) Lane 1: º¸¼¶ PS I&Lane 2: PS I-CNTBP̷Ź̵̈́oǪȤŕӃҵɅĵӴShj&PS I PŹȼ&PS I P SWNT Ǘ̵oʻɣO 277 K SN 1

SWNT 5пŹùȳɞ@N0kdUPпŹùȳɞʴӜtº|±¥¼oBk<PShj&пŹùȳ@N0R0 PS I oĵӴBk<POђă@H'

ɞoбKH' ˌΛθOͱ0H SWNT V&HiPco ̥ԩԟƣ½Ӆ

 ǪȤŕӃҵɅĵӴV&ӽtyºɎUǪȤŕӃ̈́

SCIENCE, INC. (Tokyo, Japan) 4iҀĤ@H'<

oÿͱ@&ŕӃVҵɅĵӴ±¼¢ĬSNϗ 0.8–1.2 nm O/k<PoΟљ@N0k'SWNTU 60 % (w/v) iodixanol oҵɅ±¼¢UȡSŊŗ ͕ Ϝ ̥ Ԫ O ü о > l H d U O / j & OPTO

̵ (OptiPrep: iodixanol ̵̖̈́, Sigma-Aldrich)29)

U SWNT UΌȹV&Raman ĵĢ̾Ǡ4iΌȹ5£´wº̥SNüо@H'`C&200 µLɝ̵͐üоʣ̥VèÃUҦjO/k'1.0 mg U

2&FUÂS 20 % (w/v) iodixanol o 1.8 mL oŊ

HiPco SWNT o 2% (w/v) Triton X-100 ̵̈́ 1 mL

2H'FUȼ&üо@HŕӃ̵̈́UÂȀS 100 µL

off 1 s) oбKH'FUȼ҄ԁ̦ĳͭ@Ḧ̵́o(iodixanol ͗@) oŊ2H'

U PS I d@8V PS I P SWNT U̷Ź̵̈́P̷Ź@&̗̖ÉO 90 ĵӜ҄ԁ̦ĳͭ (on 1 s,
1 ʴӜҵɅĵӴ (18,000 × g) @&̜̍oӤ0HÂ ƀů¦~¶̾ǠͱUº£¶üоɥԆVèÃUҦjO/k'º¸¼¶P@NU PS I

̹oĵŰ@H'<U SWNT ɝ̵͐oĥNUǢԜ

UaǗ̵P&º¸¼¶ PS I d@8V PS

Oͱ0H'

I-CNTBP PU SWNT ̷Ź̵̈́o 30 ĵӜ 80,000 ×

 PS I P SWNT UϤŹV&PS I P SWNT ɝ̵͐o phosphate buffer [10 mM phosphate (pH 7.4),

g SNҵɅĵӴ (Himac CS100GX II, Hitachi Koki,(pH 6.0), 0.04% DM] ÉSN̷Ź@üо@H'ˌ

ºoĵŰ@&ǣ̽SNĵĢĢȤэ (JASCO

Tokyo, Japan) @H'ҵɅĵӴȼ&ÂȀӀU PS I

0.04% DM]&d@8V MES buffer [10 mM MES

V-660, Tokyo, Japan) oͱ0Nƀů¦~¶oǢԜOͱ0Hº£¶U̷ŹȼU PS I ˂ϡ͑Ȥ

V 0, 10, 20, 30, 50, 70, 100 µg Chl mL O/j&
−1

̾Ǡ@H'

SWNT U˂ϡ͑ȤV 0, 50, 100 µg mL P@H'̷−124

ĢŹɞΛθ 27 (1) 2017
 PS I U SWNT \UƀΓђăP@N&PS I P

0R0 PS I oΡBÂȀӀUº5åǗ̵U(Cm) Pº¸¼¶ PS I UaUƶŹU͑Ȥ (Cp)ƀΓ@ǀ85̜̍@N0k<P5Ρƌ>lk'

ƶŹP̒]Щ`KN3j&SWNT жӾS PS I 5

SWNT ̷Ź̵oͱ0HP6U PSI ºU͑Ȥ Ɲ 3 V&ĵŰ@HÂȀӀU PS I ºUƀĢ

o&FlGlUº£¶UƀĢȤ4i̛Ǡ@H'
PS I P SWNT PUȜд̵̈́S3:kҮӴ PS I U

Ȥ̾Ǡ4iχĴ@Hº¸¼¶ PS I P PS

Ĵ@H'

Phosphate buffer (pH 7.4) UƶŹԩƝ 3aԪ&Ӄĸ

I-CNTBP U SWNT жӾ\UƀΓͣɎO/k'͑Ȥ (Ec) P PS I ƀΓӋ (A)Vȫ (1), (2) SNχ

5͗0ƶŹOdnC4RƀΓVтilk5&PS

 Ec = Ic × Cm/Cp

I-CNTBP U͑ȤĂǕUƀΓ5Ρ>lH'ȽKN&(1)

 A = (Ic − Ec) × V/AsPS I SɽĤ@H CNTBP 5˄ŏS˻Ѝ@N0k<(2)

P5Ρƌ>lk'½ʣO&MES buffer (pH 6.0) UƶŹOV&Phosphate buffer (pH 7.4) UƶŹPŻ˴

Ic V PS I UĹˈ͑Ȥ&V Vº£¶Uùη&F@

N AS V SWNT U̸ŊӋO/k'S PS I U͑ȤĂǕUƀΓ5Ⱦil&`HƀΓӋ¶ PS I d@8V PS I-CNTBP (50, 70, 100 µgChl¶ PS I UƶŹdƀΓӋ5ƺŊ@&`HƀΓӋS

5ƺŊ@HԩƝ 3bԪ'@4@R5i&º¸¼

 Ɲ 2 V&MES buffer (pH 6.0) ÉSNº¸¼mL ) P SWNT (50 µg mL ) o̷Ź@HP6U−1−1

ǁ6RWiL65ȾilHHc&pH ƽŗSPd

ҵɅĵӴȼUįΑO/k'ÂҠ@HҵɅĵӴ˒ë

R0жӾӹС5ƽŗBk<POӽͣͽ΅RƀΓOV&ҵɅĵӴȼS SWNT V̜̍Bk'ȽKN&

5ͯAH<P5Ρƌ>lH'<Uӽͣͽ΅RƀΓ

d@ PS I P SWNT 5пŹùŗ@N0kRiW&

UƺŊV&ϜǓP@NɈͱBkӮSƏԌPRk<

PS I V̜̍Bk<P5ˈȺ>lk'Ɲ 2c U PS

P5Є2ilk'

I-CNTBP P SWNT U̷Ź̵̈́UƶŹ&ƀΓ@N     

Ɲ 2. ǪȤŕӃҵɅĵӴShk SWNT SƀΓ@
N0R0 PS I UĵӴ
a: PS I, b: SWNTԬº¸¼¶ PS I, c: SWNTԬPS I-CNTBP. Ÿ·¼ºU PS I U͑ȤV&Ȍ4i
50, 70, 100 µgChl mL −1 O/k'

Ɲ 3. PS I U SWNT \U (a) Phosphate (pH 7.4)
ŬY (b) MES buffer (pH 6.0) ÉOUƀΓͣɎҁЛ: º¸¼¶ PS I&ӻЛ: PS I-CNTBP

25

ĢŹɞΛθ 27 (1) 20174ԮҤұƪӹǓԍɂӗшǭShk SWNT жӾ\U

 ¡ x  ¶ «    º  ̾ Ǡ V & NIR

 TEM шǭV&Hitachi HF-2000 oͱ0NбKH'

system) oͱ0NбKH'Ŏ̦҃ӘV 400–500 nm

PS I UƀΓђăphotoluminescence spectrometer (Shimadzu NIR-PL
P@&1000–1600 nm UԉƭU¡x¶«ÿͱ@HµV&º£¶͇ÃłSч̖ŗĳͭoбKHdUoͱ0H'º£¶V buffer SNºo̾Ǡ@H'¡x¶«ºV SWNT

100 čȓӆ@HȼSµS͇Ã@&FUȼ 1

U²¼~RͣɃU½LO/j&Ǘμĵʗ>lHʴӜΑιȬ6oбKH'ͷĚVŊҧӹƣ 75 kV S

SWNT V&ҝҁƿԉƭSN΁ĢoΡB' Ɲ 5 V&̵̈́o MES buffer (pH 6.0)P@&SWNT

NȾH'
Ɲ 4 V&̵̈́o Phosphate buffer (pH 7.4) P@H

ɝ̵͐ (100 µg mL−1) PFǗ̵Sº¸¼

ƶŹU SWNT (a) P PS I-CNTBP P SWNT Uп

¶ PS I d@8V PS I-CNTBP (100 µgChl mL−1) oŹù (b) U TEM ͷĚO/k'ͷĚP@N SWNT

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Published April 30, 2017º¸¼¶ PS I V&SWNT ӹ˯ÂSƀΓ@NŪЄʜͦ

ɽĤ>lN0k PS I UһğĔ5ӹ˯SƀΓ@&

1.

0R0&d@8VӃŽ@N0R0<P5Ρƌ>lk'Zheng, M., Jagota, A., Semke, E.D., Diner, B.A.,
McLean, R.S., Lustig, S.R., Richardson, R.E. and Ɲ 6c V&üо@HϜǓSſ`lk PS I ӹǓñ

Tassi, N.G. (2003) DNA-assisted dispersion andҳɞĵ&3hYһğ­sw¼¼P@NU

342.

separation of carbon nanotubes. Nat. Mater. 2, 338–

PMS UһğӹõPϜǓĬӀUӹǓñҳϣ҉oΡ

2.

B'SWNT ӹ˯UӅŗһğӹõUđVŪЄʜͦoŪ͛@H 33-35)'PMS 4iUӹǓV&PS I UŮɈÉ

Nakashima, N., Okuzono, S., Murakami, H., Nakai, T.
andYoshikawa,K.

(2003)

DNA

dissolves

single-walled carbon nanotubes in water. Chem. Lett.

Ʌ4i A0, A1, Fx, FA/FB oϣͳ@N Fe-S cluster S

3.

ñҳ>lk'FUȼ&PS I Uһğ>lH Fe-Scluster Ĕ5 SWNT ӹ˯PƀΓ@N0k<P4i&
SWNT ӹ˯SΌʄӹǓ5ñҳ>lk'SWNT ӹ

4.

˯SĽҳ@HӹǓV Pt/ITO ӹ˯ñҳ>l&F@
N Pt/ITO ӹ˯UӹǓShj PS I SӹǓoāÅ@32, 456–457.

Saifuddin, N., Raziah, A.Z. and Junizah, A.R. (2012)
Carbon nanotubes: a review on structure and theirinteraction with proteins. J. Chem. 2013.

Ito, M., Yajima, H. and Homma, Y. (2016) Strain

effect of cellulose-wrapped single-walled carbon

nanotubes measured by photoluminescence andRaman scattering spectroscopy. Jpn. J. Appl. Phys.
55, 075101.

28ĢŹɞΛθ 27 (1) 2017
5.

6.

Kam, N.W.S. and Dai, H. (2005) Carbon nanotubes

molecular wire into photosystem I: reconstitution of

biological functionality. J. Am. Chem. Soc. 127,

electrode. Angew. Chem. Int. Ed. 48, 1585–1587.

as intracellular protein transporters: generality andthe photoelectric conversion system on a gold

6021–6026.

16.

Yang, W., Thordarson, P., Gooding, J.J., Ringer, S.P.

and Braet, F. (2007) Carbon nanotubes for biological
412001.

17.

Nii, D., Hayashida, T., Yamaguchi, Y., Ikawa, S.,

Shibata, T. and Umemura K. (2014) Selective

9.

Tasis, D., Tagmatarchis, N., Bianco, A. and Prato, M.

10.

18.

carbon nanotubes. Colloids Surf. B, 121, 325–330.

carbonnanotubes

for

19.

(2006) Chemistry of carbon nanotubes. Chem. Rev.
106, 1105–1136.

20.

Qi, H., Ling, C., Huang, R., Qiu, X., Shangguan, L.,

Gao, Q. and Zhang, C. (2012) Functionalization ofassensing

platformforsensitized

21.

electrochemical immunoassay. Electrochim. Acta. 63,
11.

76–82.

single-walled
12.

sidewallcarbonfunctionalization

nanotubes

for

of

22.

proteinimmobilization. J. Am. Chem. Soc. 123, 3838–3839.

insingle-walled

23.

502.

Liu, Z., Galli, F., Janssen, K.G., Jiang, L., van derand Abrahams, J.P. (2010) Stable single-walled

14.

complexbiorecognition. J Phys Chem C. 114, 4345–4352.

24.

for

995.

Kaniber, S.M., Brandstetter, M., Simmel, F.C.,

Carmeli, I. and Holleitner, A.W. (2010) On-chipofcarbonnanotubes

withphotosystem I. J. Am. Chem. Soc. 132, 2872–2873.

Miyachi, M., Yamanoi, Y., Shibata, Y., Matsumoto,H., Nakazato, K., Konno, M., Ito, K., Inoue, Y. andH.

(2010)A

photosensing

systemCommun. 46, 2557–2559.

Nguyen, K. and Bruce, B.D. (2014) Growing green

electricity: progress and strategies for use of

1566.

Ravi, S.K. and Tan, S.C. (2015) Progress andperspectives

in

exploiting

photosynthetic

biomolecules for solar energy harnessing. Energy

Ciesielski, P.N., Scott, A.M., Faulkner, C.J., Berron,

B.J., Cliffel, D.E. and Jennings, G.K. (2008)

25.

Functionalized nanoporous gold leaf electrode films

for the immobilization of photosystem I. ACS Nano 2,
15.

Structural basis for energy transfer pathways in theconversion. Biochim. Biophys. Acta. 1837, 1553–

Olsthoorn, R.C., Oosterkamp, T.H., Hankemeier, T.
nanotube–streptavidin

Qin, X., Suga, M., Kuang, T. and Shen, J.R. (2015)

photosystem I for sustainable photovoltaic energy

Linden, H.J., de Geus, D.C., Voskamp, P., Kuil, M.E.,

carbonat 2.8 Å resolution. eLife 4, e07433.

nanoparticle, and double surfactants in water. Chem.

carbonnanotube dispersions. Chem. Phys. Lett. 429, 497–

13.

The structure of plant photosystem I super-complex

composed of photosystem I, molecular wire, gold

Yumura, M. and Iijima, S. (2006) Selectivity ofproteins

resolution. Nature 411, 909–917.

Mazor, Y., Borovikova, A. and Nelson, N. (2015)

Nishihara,

Matsuura, K., Saito, T., Okazaki, T., Ohshima, S.,
water-soluble

Saenger, W. and Krauß, N. (2001) Three-dimensional

functionalization

Chen, R.J., Zhang, Y., Wang, D. and Dai, H. (2001)
Noncovalent

TiO2 and ZnO. Sci. Rep. 2, 234.

Jordan, P., Fromme, P., Witt, H.T., Klukas, O.,

plant PSI–LHCI supercomplex. Science 348, 989–

single-walled carbon nanotubes with protein by click
chemistry

Vaughn, M., Nazeeruddin, M.K., Bruce, B.D.,

structure of cyanobacterial photosystem I at 2.5 Å

nanobiotechnology.

Nanomaterials 5, 321–350.

Mershin, A., Matsumoto, K., Kaiser, L., Yu, D.,photosystem-I biophotovoltaics on nanostructured

onto DNA molecules adsorbed on single-walled

Umemura, K. (2015) Hybrids of nucleic acids andproduction. Nat. Nanotechnol. 5, 73–79.

Graetzel, M. and Zhang, S. (2012) Self-assembled

binding of single-stranded DNA-binding proteins

8.

Frymier, P. and Bruce, B.D. (2010) Self-organizedphotosynthetic nanoparticle for cell-free hydrogenand biomedical applications, Nanotechnology 18,

7.

Iwuchukwu, I.J., Vaughn, M., Myers, N., O’Neill, H.,

Environ. Sci. 8, 2551–2573.

Miyachi, M., Yamanoi, Y., Tomo, T. and Nishihara,
H. (2016) Cross-sectional TEM analysis of an ITO

coated with photosystem I and molecular wires. J.

2465–2472.

Terasaki, N., Yamamoto, N., Hiraga, T., Yamanoi, Y.,

26.

Yonezawa, T., Nishihara, H., Ohmori, T., Sakai, M.,

Fujii, M., Tohri, A., Iwai, M., Inoue, Y., Yoneyama,

Inorg. Organomet. Polym. 26, 1309–1312.

Nii, D., Miyachi, M., Shimada, Y., Nozawa, Y., Ito,

M, Homma, Y., Ikehira, S., Yamanoi, Y., Nishihara,
H. and Tomo, T. (2017) Conjugates between

S., Minakata, M. and Enami, I. (2009) Plugging a29ĢŹɞΛθ 27 (1) 2017
photosystem I and a carbon nanotube for a phototransition energies in single-walled carbon nanotubes:

10.1007/s11120-016-0324-0).

Phys. Stat. Sol. b 244, 4002–4005.

response27.

device.

Photosynth.

(doi

Res.

Dependence on environmental dielectric constant.

Vermaas, W. (1996) Molecular genetics of the
cyanobacterium

Synechocystis

sp.

PCC

32.

6803:

principles and possible biotechnology applications. J.

28.

Tomo, T., Akimoto, S., Tsuchiya, T., Fukuya, M.,

33.

Tanaka, K. and Mimuro, M. (2008) Isolation and

Arnold, M.S., Green, A.A., Hulvat, J.F., Stupp, S.I.

34.

and Hersam, M.C. (2006) Sorting carbon nanotubes
by electronic structure using density differentiation.
Nat. Nanotechnol. 1, 60–65.

transitionenergy35.

ofsingle-walled carbon nanotubes on surrounding31.

Ed. 48, 7655–7659.

Paolucci, D., Franco, M.M., Iurlo, M., Marcaccio, M.,
Prato, M., Zerbetto, F., Pénicaud A. and Paolucci, F.
SinglingouttheElectrochemistryof

Individual Single-Walled Carbon Nanotubes inS., Mizutani, T., Jiang, J. and Maruyama, S. (2007)
excitonS. and Nakashima, N. (2009) Experimentally(2008)

Miyauchi, Y., Saito, R., Sato, K., Ohno, Y., Iwasaki,of

Tanaka, Y., Hirana, Y., Niidome, Y., Kato, K., Saito,

Single-Walled Carbon Nanotubes. Angew. Chem. Int.

Res. 98, 293–302.

Dependence

Nanotubes. Nano Lett. 8, 3097–3101.

Determined Redox Potentials of Individual (n, m)

center from Synechocystis sp. PCC 6803. Photosynth.

30.

Homma, Y. (2008) Influence of Gas Adsorption on

Optical Transition Energies of Single-Walled Carbon

Appl. Phycol. 8, 263–273.

spectral characterization of photosystem II reaction

29.

Chiashi, S., Watanabe, S., Hanashima, T. andSolution. J. Am. Chem. Soc. 130, 7393–7399.

Kim, K.K., Yoon, S.-M., Park, H.K., Shin, H.-J., Kim,
S.M., Bae, J.J., Cui, Y., Kim, J.M., Choi, J.-Y. and

Lee, Y.H. Doping Strategy of Carbon Nanotubes with

dielectric materials. Chem. Phys. lett. 442, 394–399.

Redox Chemistry. New J. Chem. 2010, 34, 2183–

Ohno, Y., Iwasaki, S., Murakami, Y., Kishimoto, S.,

2183.

Maruyama, S. and Mizutani, T. (2007) Excitonic

Complex formation between photosystem I and a carbon nanotube for a photo-conversion
device
1

2,

3

4

1

Daisuke Nii *, Mariko Miyachi Yuichiro Shimada , Yosuke Nozawa , Masahiro Ito , Yoshikazu
1

2

2

2

Homma , Shu Ikehira , Yoshinori Yamanoi , Hiroshi Nishihara , Tatsuya Tomo1

1

2

Department of Physics, Graduate School of Science, Tokyo University of Science, Department of Chemistry,3

School of Science, The University of Tokyo, Department of Industrial Chemistry, Faculty of Engineering, Tokyo
4

University of Science, Department of Physics, Faculty of Science, Tokyo University of Science30ĢŹɞΛθ 27 (1) 2017

解説特集

光化学系の構造・モデル計算から見えてきた
光合成反応の妙

Editors:

得津 隆太郎（基礎生物学研究所 環境光生物学研究部門）
柏山 祐一郎（福井工業大学 環境情報学部）

序文
解説 X ϯГͳӹǓ·¼Ⱦ̫ өǃҿԩưͯΛԪ ˜ȁ Τ½ҿԩΩÛȋˮǁԪ

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解説 Photosystem II S3:k̖ϜϤŹ¹¼~oã@H£¸ºαŐʝЬƥßԩ˕ÞǁԪ39

解説 ˬ͢UĢŗǘϕ IԭӲĢɎrºпŹù I ҄пŹùUϤʹ˳ҨPw¶}¼ñҳU˳ҨưΊТĒǬԩȂȁǁԪ

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解説特集
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12

ҩϥġ E-mail: tokutsu@nibb.ac.jp
ҩϥġ E-mail: ykas8787@gst.ritsumei.ac.jp32ĢŹɞΛθ 27 (1) 2017

解説
X 線自由電子レーザーを利用したタンパク質高分解能無損傷構造解析‡
理化学研究所・放射光科学総合研究センター、JST/CREST
平田 邦生*
ここ数年で X 線を用いたタンパク質の結晶構造解析は著しい変化を迎えたといえる。それは X 線自由電子レーザー（以下 XFEL）の利用開始による種々の技術革新によるものと言って良いだろう。XFELという新しい X 線プローブがタンパク質結晶構造解析に新しい価値を与え始めている。我々もSPring-8 に併設された XFEL 施設である SACLA(SPring-8 Angstrom Compact Free Electron Laser)を用い、金属含有タンパク質であるウシ心筋シトクロム酸化酵素の「あるがままの構造をより詳細に」捉えることに成功した。従来のタンパク質結晶構造研究と、XFEL 施設を用いたそれの違いを比較しな
がら、タンパク質結晶構造解析に XFEL を用いる利点について紹介する。

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36ĢŹɞΛθ 27 (1) 2017ъŮɈUӛǍͥɚԩS1 ͥɚԪOVӅϜ΁ͯÉɅoҩϥԃ:`@Hiǁƽǒ@0OB'

Mn3B 5+4&Mn4A 5+3 O/k<P&̖ĵъŮɈ

Received March 27, 2017; Accepted April 5, 2017;

˳ɞBk Mn UăʙV Mn1D 5+3&Mn2C 5+4&

S3:kưѿĵǓ O5 UŗǘγV OH O/k<P&–

Published April 30, 2017

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1.

Tukihara, T., Aoyama, H., Yamashita, E., Tomizuka,T., Yamaguchi, H., Itoh, K., Nakashima, R., Yaono, R.
and Yoshikawa, S. (1995) Structures of metal sites of

6. 3njS

oxidized bovine heart cytochrome c oxidase at 2.8 Å.

 ʢ@0Ģ̀ XFEL oͱ0N&Ƚ˓Uº~ѿU X ϯϤʹ˳Ҩъ˗S3:kǁ6R·ºªo2.

ъ̛Bk<P5O6H'ĵǓӋ 40 ¿҄Uº

Science 269, 1069–1074.

Muramoto. K., Hirata, K., Shinzawa-Itoh, K., Yoko-o,S., Yamashita, E., Aoyama, H., Tsukihara, T. andYoshikawa S. (2007) A histidine residue acting as a~ѿпŹùO/k~¸¬ӅŗӄϜ»Ģŗǘϕ

controlling site for dioxygen reduction and protonII пŹùUԟĵъЍ4L͗ʋĘ˳ҨơǠ@Hpumping by cytochrome c oxidase. Proc. Natl. Acad.

<PO&˻ЍoͭъBkHcSӉсR˳ҨɔƵoȾk<P5O6H'XFEL UļͱVº~ѿU3.

Ϥʹ˳Ҩъ˗SʢHRŴЍɎoÅ2&ɟ(VFUͣɃU½Lo̬ͱ@˳Ҩͯ͢ǘUǁ6R½̇oSci. U.S.A. 104, 7881–7886.

Muramoto, K., Ohta, K., Shinzawa-Itoh, K., Kanda,

K., Taniguchi, M., Nabekura H, Yamashita, E.,
Tsukihara, T. and Yoshikawa, S. (2010) Bovine
cytochrome c oxidase structures enable O2 reduction

ҊaĴ@H'

with minimization of reactive oxygens and provide a
proton-pumping gate. Proc. Natl. Acad. Sci. U.S.A.

ѫҖ ˌ˳Ҩъ˗V JST/X ϯГͳӹǓ·¼

¼ʤѐ

4.

Ӊ͖ɠͻѡԌʅҫ×ˮ)͗ʋĘ»Ő΅Ϥʹ˳Ҩъ
˗Shkͯùw¶}¼ƽʈұβUŴфŗ*3h107, 7740–7745.

Aoyama, H., Muramoto, K., Shinzawa-Itoh, K.,

Hirata, K., Yamashita, E., Tsukihara, T., Ogura, T.

and Yoshikawa, S. (2009) A peroxide bridge between

Y JST/CREST)«ºµrƄƀӖU˳Ҩͯ

Fe and Cu ions in the O2 reduction site of fully

oxidized cytochrome c oxidase could suppress theƅέǘ―˳Ҩ5dHiB̅Ο>*Uʒʉoű:Ǣ

proton pump. Proc. Natl. Acad. Sci. U.S.A. 106,ʤ@`@H'ˌΛθoPdSбKHͭΛ»ʔǳĢέǘϭŹΛθº¼UŻě3hYĩȦΐμǁ
5.

UѦġͯʣ»ǘͯUΆ˴&`Hɩвʒʉoԃ0H

2165–2169.

Hirata, K., Yamashita, E., Aoyama, H., Muramoto,

JASRI&RSC wºrµº¼¬UΆ˴oǍK., Yoshikawa, S. and Tsukihara, T. (2004) Scaling

cPBkӝąŸõS̶8əѫ͵@Â;`B'`H

set.,” J. Synchrotron Rad. 11, 60–63.

of one-shot oscillation images with a reference data

ˌεSӝBkǢԜV SACLA ļͱǢԜѡԌԩѡԌ

6.

ͼŷԸ2012A8011& 2012B8040& 2013A8047 and2013B8052ԪO¼ůӲoбKHɞ˚OB'

Physical and chemical considerations of damage

induced in protein crystals by synchrotron radiation: aradiation chemical perspective. J. Synchrotron Rad. 9, `HâƘ&ĢŹɞǘðSɳȺ@N8I>KH
y¼{tO'Neill, P., Stevens, D.L. and Garman, E.F. (2002)¼UġͯʣS̶8əѫ0H@`B'

7.

ĢŹɞǘðјP01<PO PSII SӝBkŧεóĖBk]64PdɌKHUOB5&ΫГҌV X

329–332.

Neutze, R., Wouts, R., van der Spoel, D., Weckert, E.

and Hajdu, J. (2000) Potential for biomolecularimaging with femtosecond X-ray pulses. Nature 406,

ϯϤʹ˳Ҩъ˗Uɥ̥ӛ΁oɰȲ@`@HUO&
XFEL P01ʢĢ̀oQ1Є2NQ1ļͱ@H

8.

4S͖͘oȲNNŧεoˀ4DNԃ6`@H'âƘU΁жSӝҩBkrtsre=ɗтRQ=

9.

37

752–757.

Schlichting,I. (2015)

Structural

biology:

Photosynthetic complex in close-up. Nature 517, 26–

27.

Hirata, K., Shinzawa-Itoh, K., Yano, N., Takemura,ĢŹɞΛθ 27 (1) 2017
S., Kato, K., Hatanaka, M., Muramoto, K., Kawahara,

10.

T., Tsukihara, T., Yamashita, E., Tono, K., Ueno, G.,

Hikima, T., Murakami, H., Inubushi, Y., Yabashi, M.,

Suga, M., Akita, F., Hirata, K., Ueno, G., Murakami,
H., Nakajima, Y., Shimizu, T., Yamashita, K.,

Yamamoto, M., Ago, H. and Shen, J.R. (2015) Native

Ishikawa, T., Yamamoto, M., Ogura, T., Sugimoto,structure of photosystem II at 1.95 Å resolution

H., Shen, J.R., Yoshikawa, S. and Ago, H. (2014)

viewed by femtosecond X-ray pulses. Nature 517,

Determination of damage-free crystal structure of an

99–103.

X-ray–sensitive protein using an XFEL. Nat. Meth.
11, 734–736.

High resolution damage-free structure analyses of supra-molecular complexes with SF-ROX
Kunio Hirata*
RIKEN SPring-8 Center, JST/CREST

38ĢŹɞΛθ 27 (1) 2017

解説
Photosystem II における水素結合ネットワークを介したプロトン移動‡
東京大学

先端科学技術研究センター
斉藤 圭亮*

蛋白質中のプロトン移動は水分子が作る水素結合ネットワーク上で起こる。Photosystem II (PSII) の結晶構造では、蛋白質中の水分子が見えているので、その構造情報を利用することで、PSII 内で起こるプロトン移動のしくみを理論的に調べることができる。本稿では、蛋白質がいかに水素結合ネット
ワークをうまく利用してプロトン移動を行っているのかを、(1) PSII の水分解における第一段階のプロトン排出と、(2) 水分解反応に直接関わらないが酸化還元活性をもつ TyrD からのプロトン排出という２つの具体例を通じて紹介する。

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0'`H&ϣ҉PRk̖ĵǓƂƜUЮ΃ѿͮƹU

0 D2 ĔSõϽ@N0k5&D1 ĔUǰɈBkƶ

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45ĢŹɞΛθ 27 (1) 2017
FourierŹ¹¼~oҦKNʂĴ>lk4d@lR
0'

10.

ѫҖ

 ˌΛθV˕Þǁǘ»ΚŘǅʖʁUdP&ŻΛθ

11.

ǣUċƼѡβUƧÃǸϖ̓PPdSбKHΛθ

12.

Received March 28, 2017; Accepted March 31, 2017;
Published April 30, 2017

2.

Agmon, N. (1995) The Grotthuss mechanism. Chem.

13.

Edition. John Wiley & Sons Inc, Hoboken, NJ.
Phys. Lett. 244, 456–462.

14.

Pomes, R. and Roux, B. (1996) Structure andPal, R., Negre, C.F., Vogt, L., Pokhrel, R., Ertem, M.
Z., Brudvig, G.W. and Batista, V.S. (2013) S-stateoftheoxygen-evolving

complexphotosystem II. Biochemistry 52, 7703–7706.

of

Siegbahn, P. E. (2013) Water oxidation mechanism inphotosystem II, including oxidations, proton release
Biochim. Biophys. Acta 1827, 1003–1019.

Kulik, L. V., Epel, B., Lubitz, W. and Messinger, J.

(2007) Electronic structure of the Mn4OxCa cluster in

Umena, Y., Kawakami, K., Shen, J.R. and Kamiya, N.

13435.

EPR spectroscopy. J. Am. Chem. Soc. 129, 13421–

gramicidin a channel. Biophys. J. 71, 19–39.

oxygen-evolving

15.

Cox, N., Pantazis, D.A., Neese, F. and Lubitz, W.

Suga, M., Akita, F., Hirata, K., Ueno, G., Murakami,

16.

Saito, K., Rutherford, A. W. and Ishikita, H. (2015)(2011)

Crystalstructureof

55–60.

H., Nakajima, Y., Shimizu, T., Yamashita, K.,

17.

ground and excited-state potential surfaces of
1.

Formulation

18.

and

parametrization. J. Am. Chem. Soc. 94, 5612–5625.

Warshel, A. and Levitt, M. (1976) Theoretical studies

19.

of enzymic reactions: dielectric, electrostatic andsteric stabilization of the carbonium ion in theRenger, G. (2001) Photosynthetic water oxidation tooxygen:apparatusandBiochim. Biophys. Acta 1503, 210–228.

Energetics of proton release on the first oxidation
8488.

Petrek, M., Otyepka, M., Banas, P., Kosinova, P.,
Koca, J. and Damborsky, J. (2006) CAVER: a newcavities. BMC Bioinformatics 7, 316.

Takaoka, T., Sakashita, N., Saito, K. and Ishikita, H.
(2016) pKa of a proton-conducting water chain inphotosystem II. J. Phys. Chem. Lett. 7, 1925–1932.

Bricker, T. M., Roose, J. L., Fagerlund, R. D., Frankel,
L. K., and Eaton-Rye, J. J. (2012) The extrinsic
proteins of Photosystem II. Biochim. Biophys. Acta

reaction of lysozyme. J. Mol. Biol. 103, 227–249.
molecular

46, 1588–1596.

tool to explore routes from protein clefts, pockets andWarshel, A. and Karplus, M. (1972) Calculation ofmolecules.

(2013) Biological water oxidation. Acc. Chem. Res.

step in the water-oxidizing enzyme. Nat. Commun. 6,

99–103.

conjugated9.

photosystem II. J. Am. Chem. Soc. 134, 1504–1512.

of photosystem II based on pulse 55Mn-ENDOR andviewed by femtosecond X-ray pulses. Nature 517,

8.

Myers, W. K., Chen, P. Y., Whittaker, J. W. andtranslocation along the single-file water chain in the

structure of photosystem II at 1.95 Å resolution

7.

McConnell, I.L., Grigoryants, V M., Scholes, C.P.,

the S0 and S2 states of the oxygen-evolving complex

dynamics of a proton wire: A theoretical study of H+

Yamamoto, M., Ago, H. and Shen, J.R. (2015) Native

6.

reactions. Biochim. Biophys. Acta 1411, 86–91.

pathways, O-O bond formation and O2 release.

photosystem II at a resolution of 1.9 Å. Nature 473,

5.

water oxidation: a simplex-scheme of its partial

model

ŪЄʜͦ

4.

Haumann, M. and Junge, W. (1999) Photosynthetic

relevance to the oxygen-evolving complex of@Â;`B'

3.

Biochemistry 44, 1708–1718.

spectroscopy.

of (17O, 1H, 2H) water in manganese catalase and its

ɸʎoқҧS8I>KH˝ѠЅUʣSdəѫ͵

Voet, D. and Voet, J.G. (2011) Biochemistry, 4th

infrared

Brudvig, G.W. (2012) EPR-ENDOR characterization

O/k'<<Sѫɗoж@`B'`H&ҶĶR=1.

transform

20.

mechanism.

Suzuki, H., Sugiura, M. and Noguchi, T. (2005) pH

21.

dependence of the flash-induced S-state transitions in

the oxygen-evolving center of photosystem II from

1817, 121–142.

Ifuku, K. and Noguchi, T. (2016) Structural couplingof extrinsic proteins with the oxygen-evolving centerin photosystem II. Front. Plant. Sci. 7, 84.

Wei, X. P., Su, X. D., Cao, P., Liu, X. Y., Chang, W.
R., Li, M., Zhang, X.Z. and Liu, Z.F. (2016) Structureof spinach photosystem II-LHCII supercomplex at

Thermosynechoccocus elongatus as revealed by46ĢŹɞΛθ 27 (1) 201722.

3.2 Å resolution. Nature 534, 69–74.

26.

Sakashita, N., Watanabe, H.C., Ikeda, T. and Ishikita,H.

(2017)

Structurally

conserved

channelsincyanobacterial and plant photosystem II. Photosynth.
23.

27.

Res., 1–11.

Saito, K., Shen, J.R., Ishida, T. and Ishikita, H. (2011)

Biochim. Biophys. Acta 1655, 222–230.

Babcock, G.T., Barry, B.A., Debus, R.J., Hoganson,
C. W., Atamian, M., McIntosh, L., Sithole, I. and
II: from radical chemistry to multielectron chemistry.

structure. Biochemistry 50, 9836–9844.

28.

Ahlbrink, R., Haumann, M., Cherepanov, D.,
Bogershausen, O., Mulkidjanian, A. and Junge, W.

Biochemistry 28, 9557–9565.

Diner, B.A. and Babcock, G.T. (1996). in Oxygenic

Photosynthesis: The Light Reactions (Ort, D. R., andYocum, C. F., Eds.), pp 213–247, Kluwer Academic(1998) Function of tyrosine Z in water oxidation byphotosystem II: electrostatical promotor instead of25.

The stable tyrosyl radical in photosystem II: why D?

Yocum, C. F. (1989) Water oxidation in photosystem

Short hydrogen-bond between redox-active tyrosine

YZ and D1-His190 in the photosystem II crystal

24.

Rutherford, A.W., Boussac, A. and Faller, P. (2004)

29.

hydrogen abstractor. Biochemistry 37, 1131–1142.

Nakamura, S. and Noguchi, T. (2015) Infrared

detection of a proton released from tyrosine YD to the30.

bulk upon its photo-oxidation in photosystem II.
Biochemistry 54, 5045–5053.

Publishers, Dordrecht, Netherlands.

Saito, K., Sakashita, N. and Ishikita, H. (2016)
Energetics of the proton transfer pathway for tyrosine

D in photosystem II. Aust. J. Chem. 69, 991–998.

Saito, K., Rutherford, A.W. and Ishikita, H. (2013)

Mechanism of tyrosine D oxidation in Photosystem II.
Proc. Natl. Acad. Sci. U.S.A. 110, 7690–7695.

Proton Transfer via Hydrogen Bond Networks in Photosystem II
Keisuke Saito *Research Center for Advanced Science and Technology, The University of Tokyo47ĢŹɞΛθ 27 (1) 2017

解説
植物の光化学系 I－集光性アンテナ複合体 I 超複合体の結晶構造とエネルギー伝達の構造基盤‡
岡山大学異分野基礎科学研究所
菅 倫寛*
光化学系 I 複合体は光駆動による電子移動と酸化還元反応を行い、糖の合成に必要な還元力を供給している。植物の光化学系 I 複合体は反応中心コアと光捕集アンテナ I 複合体が光化学系 I-光捕集アンテナ
I 超複合体を形成し、光エネルギーをほぼ 100%の効率で伝達および変換している。光合成における高い効率での光エネルギー利用の基本原理を理解することを目的とし、筆者らは高等植物マメ由来の光化学系 I-光捕集アンテナ I 超複合体の結晶構造を分解能 2.8 Å で決定した。これにより、分子量が 60万に及ぶ超複合体の詳細な構造、とりわけ、光捕集アンテナ I を構成する 4 つの Lhca サブユニットにおいて秩序正しく配置されたクロロフィル、カロテノイド、脂質分子、水分子などによる構造基盤を明らかにし、光エネルギーの捕集と伝達の経路、および、光阻害に対する防御機構を提唱した。

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Received March 3, 2017; Accepted March 29, 2017;ĔSʵ>lN0kӅɎr«ӅUʙ5ǀ0'<liU˳Ҩ΅ȏͽ4i Lhca2/4 U BC ¶¼£Uʣ5Published April 30, 2017˳Ҩƽŗ@ʭ8&`H¶¼­ºĔUӅɎŗoəΗ@ ʭ 0 < P 5 Ρ ƌ > l k U O Lhca2/4 O VŪЄʜͦ

Lhca1/3 P̒]N|º¡s¶Ƙ҉V̬΁O/

1.

Ⱦ̫ өǃҿ (2016) ĢŹɞS3:kȰĢԆŗVy´|ººPU̖ϜϤŹU˴ȫSdҴ

2.

Jordan, P., Fromme, P., Witt, H.T., Klukas, O.,

kPʅǭ>lk'>iS&Lhca2/4 P Lhca1/3 O
05тilk'y´|ººU¶¼­ºĔU­z¬ΛθUʢǾӛ ĢŹɞΛθ 26, 36−42.

Saenger, W., and Kraus, N. (2001) Three-dimentional
structure of cyanobacterial photosystem I at 2.5 Å

νӀV Lhca2/4 OV Trp 127ԩLhca2Ԫ&Trp 126
3.

ԩLhca4ԪP̖ϜϤŹ@N0kUSǰ@&Lhca1/3
OV Trp 101ԩLhca1ԪP Trp 130ԩLhca3ԪSŊ2,
Gln 105ԩLhca1Ԫd@8V Thr 133ԩLhca3ԪPF4.

lGl̖ϜϤŹoȳɞ@N0k'<lV Lhca2/4
U L2 ӀõUʣ5y´|ººPUчƆɎ5

ö0<P5ʅǭ>l&<U<P4id LHCI UÉ

5.

ǅU¢²S30N|º¡s¶Ƙ҉5̬΁O/k<P5Ρƌ>lk 5)'

resolution. Nature 411, 909−917.

Ben-Shem, A., Frolow, F. and Nelson, N. (2003)
Crystal structure of plant photosystem I. Nature 426,
630−635.

Qin, X., Suga, M., Kuang, T. and Shen, J.R. (2015)

Structural basis for energy transfer pathways in theplant PSI-LHCI supercomplex Science 348, 989−995.

Suga, M., Qin, X., Kuang, T. and Shen J.R. (2016)
Structure and energy transfer pathways of the plant
photosystem I-LHCI supercomplex. Curr. Opin.

6. 3njS

6.

 ˌΛθShj&ԟπˬ͢U PSI-LHCI ҄пŹùUіϟR˳Ҩ5ʬi4SRKH'μù˳ҨSdP

7.

M6&QUh1S@NȎǁR҄пŹù5ȳɞ>l
k4&F@NŎ҃>lHw¶}¼VQUϣ҉oStruc. Biol. 39, 46−53.

Amunts, A., Drory, O. and Nelson, N. (2007) The

structure of a plant photosystem I supercomplex at
3.4 Å resolution. Nature 447, 58−63.

Amunts, A., Toporik, H., Borovikova, A. and Nelson.

N. (2010) Structure determination and improved
model of plant photosystem I. J. Biol. Chem. 285,ҦKNŮɈÉɅrӀĵ\Pǵ4lk4&RQ҄
8.

пŹùSӝBkӘȝU΀ƏSτ2HI:OR8&
L2 ӀõSϤŹBky´|ºº5|º3478−3486.

Kuhlbrandt, W., Wang, D.N. and Fujiyoshi, Y.
(1994) Atomic model of plant light-harvesting

complex by electron crystallography. Nature 367,¡s¶Ƙ҉ShjӽĢŗǘ΅̴ĢUȶŇoɰK
N0kP01ʢHRʇ˨dбKH'<liU˳Ҩ9.

ưΊV&Ģw¶}¼ohjŏͨК8ļͱBk<
PUO6kʢуUw¶}¼ƽʈˏʠU®¶614−621.

Liu, Z., Yan, H., Wang, K., Kuang, T., Zhang, J., Gui,
L., An, X. and Chang W. (2004) Crystal structure ofspinach major light-harvesting complex at 2.72 Åº£·¼PdRk<P5ˈȺ>lk'10.

ѫҖ

resolution. Nature 428, 287−292.

Pan, X., Li, M., Wan, T., Wang, L., Jia, C., Hou, Z.,
Zhao, X., Zhang, J. and Chang, W. (2011) Structural
insights into energy regulation of light-harvesting ˌΛθVȂȁǁǘͽĵӊưΠέǘΛθɣU̜

complex CP29 from spinach. Nat. Struct. Mol. Biol.

ȩá&ÉƟέǘӣˬ͢ΛθɣU Xiaochun Qin&

Tingyun Kuang Ѧ̓PUĨŻΛθO/k'X ϯϤ

11.

ʹ˳Ҩъ˗¼V SPring-8 U BL41XU 3hY
BL44XU SN̾Ǡ@H'<UƶoďjN3΢͵@12.

Â;k'`HˌΛθVέǘΛθѸмŌӌπUŌɞ
oű:NбKH'

18, 309−315.

Nelson, N. (2009) Plant Photosystem I − The Most
Efficient Nano-Photochemical Machine. J. Nanosci.
Nanotechno. 9, 1709−1713.

Mazor, Y., Nataf, D., Toporik, H. and Nelson, N.

(2014) Crystal structure of virus-like photosystem I
complexes from the mesophilic cyanobacterium
Synechocystis PCC 6803. eLife 3, e01496.

53ĢŹɞΛθ 27 (1) 2017
13.

14.

Mazor, Y., Borovikova, A. and Nelson, N. (2015)15.

The structure of plant photosystem I super-complex
at 2.8 Å resolution. eLife 4, e07433.

photosystem I light harvesting by zeaxanthin. Proc.

(1998) Arabidopsis mutants define a central role forxanthophyll

cycle

in

the

regulation

Ahn, T.K., Petrozza, A., Polli, D., Fleming G.R.,

Cerullo, G. and Bassi, R. (2014) Regulation of

Niyogi, K.K., Grossman, A.R. and Bjorkman, O.
theBallottari, M., Alcocer, M.J., D’Andrea, C., Viola, D.,

Natl. Acad. Sci. U.S.A. 111, E2431−2438.

ofphotosynthetic energy conversion. Plant Cell 10,1121−1134.

Crystal structure and energy transfer pathways of the plant photosystem I-light harvesting
complex I supercomplex
Michihiro Suga*
Research Institute for Interdisciplinary Science, Okayama University54ĢŹɞΛθ 27 (1) 2017

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久留米大学医学部
原田 二朗
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第 73 回 藤原セミナー
The 4th International Conference “Molecular Life of Diatoms” 開催のお知らせʦʴԸ2017 ȝ 7 ˃ 9 ʦԩʦԪ15:00 Լ 7 ˃ 13 ʦԩˉԪ15:00
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57ĢŹɞΛθ 27 (1) 2017

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8th International Conference “Photosynthesis and Hydrogen Energy Research for
Sustainability -2017”の開催案内
期日： 2017 年 10 月 30-11 月 4 日
場所：インド、ハイデラバード大学
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第 25 回「光合成セミナー2017：反応中心と色素系の多様性」の開催案内
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立命館大学 生命科学部 生命情報学科
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60

ĢŹɞΛθ 27 (1) 2017

若手の会新会長からのごあいさつ
東京大学 大学院総合文化研究科 広域科学専攻
榎本 元
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1,500 ĭ ԩðј&Λθð&¹¼~³£RQU˨ĬoſbԪ

50,000 ĭ ԩÂяPðј\UȠƁʠoſbԪԩɼҙÖǠʦԸȜɞ  ȝ  ˃  ʦԪԩðƉѺ˥V԰˃԰ʦԼ԰Ա˃Բ԰ʦošõP@`BԪԫ пʙȝĵUðѸoġɦ0Oɼjҙb<PdŴЍOB'FUƶŹ&Ҧĉ˾SԩúȝȤԼúȝȤĵԪP3ˀ6Ã>0'ҩϥġ-700-8530 ȂȁΐȂȁȑŘŚ̫ȄÉ 3-1-1ȂȁǁǘͽĵӊưΠέǘΛθɣԟ˺л½ҿ ΛθǣĬʦˌĢŹɞǘð
TEL : 086-251-7861
FAX : 086-251-7876§¼¬¦¼: http://photosyn.jpӁĄɼˁŲȥ ŊĤЅżԸʦˌĢŹɞǘð ŲȥͼŷԸ00140-3-730290
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ξԱ˒ ΋΅ˌðVĢŹɞUưΠ3hYɈͱĵӊUΛθ΁ǾoĆҫ@&ΛθЅ΍ÙUÝ̭o̶ck<Po΋΅PBk'ξԲ˒ ×ˮˌðVł˒U΋΅oҳɞBkHcS&º©u¬ӛėRQU×ˮoб1'ξԳ˒ ðƉ԰ԮǠЁˌðU΋΅SѻŻBkČàV&΂ӔɥϪoϣNðƉSRk<P5O6k'`H&ƚù&˻ӝV&ѻŌðƉSRk<P5O6k'
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ԲԮðѸ
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ìȟ×ðSĴȖBk<P5O6k'
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ʦˌĢŹɞǘðUȶƉҸĴSӝBk͵@ŹnDȜɞ 27 ȝ 5 ˃ 27 ʦ ȟ×ð԰ԮҸɺψͭǎƉðˌðUҸɺoĦ̅SǢʤBkHc&ҸɺψͭǎƉðoϽ8'ҸɺψͭǎƉ 2 żVȘìȟ×ð5ȟ×ð
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「光合成研究」編集委員会
ϱӲӘîΩ ēǃҿԩÞӂǁǘԪϱӲǎƉϑÛ Ģ½ҿԩӼȂǁǘԪϱӲǎƉɞȈ ΢ԩӼȂǁǘԪϱӲǎƉΖǛ Иԩ˕ÞǁǘԪ

日本光合成学会

2017年度役員ðӘԟ˺ л½ҿԩȂȁǁǘԪ×őǻӘơ̚ Ħ̎ԩʩδͲǁǘԪȘìȟ×ͲÉ ̇ԩŘ̳ҲǁǘԪ

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