51$ ᖱῦἪࢅ⏕࠷ࡒ㧏➴᳔∸ࡡකྙᠺᶭ⬗ࡡᨭንࡡムࡲ
Application of the RNA-interference technique on the modification of plant
photosynthetic activity௿⚗ ೸ኯ㑳 ா㒌ኬᏕኬᏕ㝌⏍࿤⛁Ꮥ◂✪⛁ ຐᩅAssistant Professor, Kentaro Ifuku: Graduate School of Biostudies, Kyoto University࿰ᩝ こ᪠ ⤄ᥦ࠻ '1$ ᢇ⾙࡞ࡻࡾ᳔∸ᶭ⬗ࡡᨭንࡢࠉᆀ⌣⎌⑧ၡ㢗ࡡゆỬ࡞᭯ຝ࡝ᡥアࡡୌࡗ࡚࠵ࡾࠊ≁࡞᳔∸࠿⾔࠹᭩ࡵ㔔こ࡝໩Ꮥཬᚺ࡚࠵ࡾකྙᠺࡢࠉᆀ⌣୕ࡡ୹ࡒࡾ࢙ࢾ࣭ࣜ࢟″࡚࠵ࡽࠉࡐࡡิⓆア㝭࡚࠵ࡾラฦゆ㓗⣪Ⓠ⏍ཬᚺࡢᶭ⬗ᨭንࡡ㔔こ࡝ᵾⓏ࡚࠵ࡾࠊᮇ◂✪࡚ᠻࠍࡢࠉ51$ ᖱῦἪࢅ⏕࠷࡙කྙᠺ㓗⣪Ⓠ⏍ཬᚺ࡞㛭ࢂࡾ SVE3 㐿ఎᏄࡡⓆ⌟ᢒโ᳔∸మࢅషᠺࡊࠉ᳔∸ࡡラฦゆ㓗⣪Ⓠ⏍ཬᚺࡡெ㎎Ⓩ࡝ᨭን࠿ྊ⬗࡚࠵ࡾࡆ࡛ࢅ
᪺ࡼ࠾࡞ࡊࡒࠊ51$ ᖱῦἪ࡛ࡢ஦ᮇ㙈ࡡ 51$ ฦᏄࢅ⣵⬂࡞ᑙථࡌࡾࡆ࡛࡚ࠉ≁ᏽࡡ㐿ఎ
ᏄࡡⓆ⌟ࢅᢒโࡌࡾᢇ⾙࡚࠵ࡾࠊࡈࡼ࡞ࡆࡡ 51$ ᖱῦἪࢅᚺ⏕ࡊࠉラฦゆ㓗⣪Ⓠ⏍ཬᚺ࡞㛭୙ࡌࡾ࡛⩻࠻ࡼࡿࡾ᪜▩㐿ఎᏄࠉ→ࡦࠉᮅ▩㐿ఎᏄࡡⓆ⌟โᚒࢅ⾔࠷ࠉකྙᠺ㓗⣪Ⓠ⏍ཬᚺㄢ⟿ࡡฦᏄᶭᵋࡡゆ᳔᪺࡛∸ࡡකྙᠺᶭ⬗ࡡྡྷ୕ࢅ┘ᣞࡊࡒࠊAbstract
Our daily life is dependent on photosynthesis that produces oxygen and assimilates carbon
dioxide into organic matter. It determines to a large extent the composition of our atmosphereand provides essential food and fuel. The first step of this process occurs in a multisubunit
pigment-protein complex photosystem II using light energy to split water into oxygen and its
reducing equivalents. The molecular mechanism of the water-splitting reaction is largely
uncharacterized, and intensive studies are needed for the development of this principal
energy-conversion reaction. We showed that the water-splitting reaction could be engineeredusing molecular technique, especially, RNA interference (RNAi) technology, in planta. RNAi isa powerful tool to control the gene expression by introducing double-stranded RNA into plantcells. We further developed RNAi technique to control the water-splitting reaction precisely andto find out novel components that control this reaction.

偏㸣 ◂✪ ┘ Ⓩࡐࡆ࡚ࠉ྘ SVE3 㐿ఎᏄࡡ v㟸⩳ズ㡷ᇡ ᭯⏕᳔∸ࡡ㛜Ⓠ࡞࠵ࡒࡖ࡙ࡢࠉකྙᠺ

ࢅᵾⓏ࡛ࡊࡒ GV51$ Ⓠ⌟࣊ࢠࢰ࣭ࢅ⏕࠷࠿⏍ࡳ࢙ࢾ࣭ࣜ࢟ࡡ㞺⤝ࣁࣚࣤࢪ࠿᳔∸࡙ࠉ4 ࡗࡡ SVE3 㐿ఎᏄ࡞ᑊࡊᕣ␏Ⓩ࡝

మහ࡚㐲ว࡚࠵ࡾࡆ࡛࠿㔔こ࡚࠵ࡾࠊ༳51$L 'LIIHUHQWLDO51$Lࠉ௧ୖࠉG51$Lࡔࠉ᳔∸࡞⎌⑧ࢪࢹࣝࢪ⪇ᛮࠉ⪇⑋ᛮ࡛ࢅ⾔࠷ࠉ⵫✒ࡌࡾ 3VE3 ࢰࣤࣂࢠ㈹㔖࠿ア࠷ࡖࡒ᩺ࡒ࡝ᶭ⬗ࢅ௛ຊࡌࡾࠉࡵࡊࡂࡢ

㝭Ⓩ࡞ን໩ࡊࡒᙟ㈹㌷ᥦ᳔∸ࡡషᠺࢅ⾔ኣ཭ᛮࢅ௛୙ࡌࡾ࡛࠷ࡖࡒ㝷࡞ࡢࠉᚪࡍࡖࡒ ᅒ偏$ ࠊࡐࡡ⤎ᯕࠉක໩Ꮥ⣌ ,, ࡡࡐࡿ࡞ᚺࡋࡒ࢙ࢾ࣭ࣜ࢟౩⤝ࢅ⩻៎ࡎࡠ㔖࡚ࡢ࡝ࡂࠉ⵫✒ࡌࡾ 3VE3 㔖࡛ක໩Ꮥ⣌ࡣ࡝ࡼ࡝࠷ࠊୌ᪁࡚ࠉఴ๨࡝࢙ࢾ࣭ࣜ࢟,, Ὡᛮ࡞᪺ࡼ࠾࡝とࡡ┞㛭࠿࠵ࡾࡆ࡛࠿

࠿ᾐ㈕ࡈࡿ࡝࠷ሔྙ࡞ࡢὩᛮ㓗⣪⛸➴࡞᪺ࡼ࠾࡛࡝ࡖࡒࠊࡐࡆ࡚ḗ࡞ SVE3 㐿ఎᏄࡻࡾ㓗໩ࢪࢹࣝࢪࡡⅴᅄ࡛࡝ࡽࠉ᳔∸ࡡࢅ㐛๨Ⓠ⌟ࡈࡎࡒ᳔∸ࡡషᠺࢅ⾔ࡖࡒࠊ⏍⫩࡞୘ฺࢅ⏍ࡋࡾࡆ࡛࡛࡝ࡾࠊࡆ࠹ࡊ

ࡊ࠾ࡊ࡝࠿ࡼࠉ㏳ᖏࡡ⏍⫩᮪௲ୖ࡞࠽࠷

ࡒ࢙ࢾ࣭ࣜ࢟౩⤝ࢅெ㎎Ⓩ࡞ㄢ⟿ࡌࡾᡋ࡙㔕⏍ᰬ௧୕ࡡකྙᠺὩᛮࢅ♟ࡌ಴మࡢ␆࡛ࡊ࡙ࠉᠻࠍࡢ 51$ ᖱῦἪ 51$L ࢅᚺᚋࡼࡿ࡝࠾ࡖࡒࠊ

⏕ࡊ࡙ࠉラฦゆ㓗⣪Ⓠ⏍ཬᚺ࡞├᥃Ⓩࠉ㛣᥃Ⓩ࡞㛭ࢂࡾࢰࣤࣂࢠ㈹ࢦࣇࣗࢼࢴࢹ

偐—偐 㸣dRNAi ࡞ ࠽ࡄ ࡾᶭ ⬗ ┞⿭ ࠉ→ࢅᨭንࡊࠉ࢙ࢾ࣭ࣜ࢟౩⤝㔖ࢅㄢᩒࡌࡾࡦࠉ ฦᏄ ⨠ ᥦࡆ࡛ࢅ⩻᱄ࡊࡒࠊᐁ㥺࡞ࡢࠉラฦゆ㓗⣪ LQYLWUR ࡡᐁ㥺࡚ࡢࠉ␏⛸᳔∸࡞⏜ᮮⓆ⏍ཬᚺࢅ⾔࠹ක໩Ꮥ⣌ ,, ࢰࣤࣂࢠ㈹々ࡌࡾ 3VE3 ࢅ⤎ྙࡈࡎࡾࡆ࡛࡚ක໩Ꮥ⣌ྙమ࡞࠽࠷࡙ࠉࡐࡡㄢ⟿Ⓩᙲ๪ࢅᢰࡖ࡙,, Ὡᛮࢅᨭን࡚ࡀࡾྊ⬗ᛮ࠿♟ࡈࡿ࡙࠷࠷ ࡾ ࡛ ⩻ ࠻ ࡼ ࡿ ࡙ ࠷ ࡾ SVE3 㐿 ఎ Ꮔ ࢅࡾࠊࡐࡆ࡚ G51$L ࢅ⏕࠷࡙ࠉ␏⛸᳔∸⏜51$L ࡡ୹ࡒࡾᵾⓏ࡛ࡊࡒࠊࡈࡼ࡞ SVE3

ᮮࡡ 3VE3 ࡞ࡻࡾහᅹࢰࣁࢤ 3VE3 ࡡ LQ࡝࡜ࡡ᪜▩㐿ఎᏄ௧አ࡞ࡵࠉラฦゆ㓗⣪

YLYR ࡚ࡡฦᏄ⨠ᥦࢅムࡲࡒࠊරమⓏ࡞ࡢࠉⓆ⏍ཬᚺࡡโᚒ࡞㛭ࢂࡾᮅ⇆ᏽࡡ㐿ఎᏄ๑㏑ࡊࡒࢰࣁࢤ SVE3 㐿ఎᏄࡡ v㟸⩳ズࡡ༟㞫ࢅࠉ࣓ࢸ᳔ࣜ∸࡚࠵ࡾࢨࣞ࢕ࢽࢻ

㡷ᇡࢅᵾⓏ࡛ࡊࡒ GV51$ Ⓠ⌟࣊ࢠࢰ࣭ࢅ

ࢫࢻ⏕࠷࡙⾔ࡖࡒࠊᑙථࡊࡒᙟ㈹㌷ᥦమ࡞ࠉ␏⛸᳔∸⏜ᮮࡡ

3VE3 ࢅᙁโⓆ⌟ࡌࡾࢤࣤࢪࢹࣚࢠࢹࢅ්偐㸣 ◂✪ ⤊ 㐛ᑙථࡊࡒ偆ᅒ偏%假ࠊࡆࡿ࡞ࡻࡽ࣌ࢗࣝࣤ

偐—偏 㸣 Differential RNAi (dRNAi)࡞ࡻ

ࢮࢗࡷ࡛࢞ࣖࢗࣛ࠷ࡖࡒ␏⛸᳔∸⏜ᮮࡡࡾ psbP 㐿ఎ Ꮔࡡ 㐽 ᢝⓏ Ⓠ⌟ ᢒ โ

3VE3 ࢅ୹こ࡞⵫✒ࡌࡾᙟ㈹㌷ᥦࢰࣁࢤࡡ

ᮞᩩ࡞⏕࠷ࡒࢰࣁࢤ偆Nicotiana tabacum假షᠺ࡞ᠺຉࡊࡒࠊࡈࡼ࡞ࡆࡡ㐛⛤࡚ࠉ࡞ࡢ 4 ࡗࡡ SVE3 㐿ఎᏄ࠿Ꮛᅹࡊࠉࡐࡡධሲᇱ࡛࠷࠹㟸ᖏ࡞▯࠷ᵾⓏ㒼า࡚᭯ຝ࡝

࡙࠿ᶭ⬗Ⓩ࡝ࢰࣤࣂࢠ㈹ࢅⓆ⌟ࡊ࡙࠷ࡾࠊ ᢒโຝᯕࢅ♟ࡌࠉ᩺ぜ 51$L ࣊ࢠࢰ࣭⣌ࡡ偐—㸨 㸣᩺ ぜ ක໩ Ꮥ ⣌ II โ ᚒ ᅄᏄ ࡡ᥀㛜Ⓠࢅ⾔ࡖࡒࠊ⣬ ࢣࢿ࣑᝗ሒ࠿ฺ⏕࡚ࡀࡾ࣓ࢸ᳔ࣜ∸࡚࠵ࡾࢨࣞ࢕ࢽࢻࢫࢻࢅᮞᩩ࡞ࠉ⌟᮶ாኬᏕ
ࡡኬᯐに༡⒬ࡼ࠿㛜Ⓠࡊࡒ ATTED-II ࢨࢪࢷ࣑ࢅฺ⏕ࡊ࡙᪜Ꮛࡡ࣏࢕ࢠࣞ࢓ࣝ࢕ࢸ࣭ࢰ࣭࣊ࢪࢅゆᯊࡊࠉラฦゆ㓗⣪Ⓠ⏍ཬᚺࡡโᚒ࡞㛭ࢂࡾᅄᏄࡡ༟㞫ࢅムࡲࡒࠊࡐࡊ࡙㐽ᢜࡈࡿࡒು⿭㐿ఎᏄ࡞㛭ࡊ࡙ࠉ
51$L ᙟ㈹㌷ᥦమࡷ✲↓ን␏మࢅ⏕࠷ࡒᶭ⬗ゆᯊࢅ㐅ࡴࡒࠊ
ᅒ偏㸯 v875 ࢅ⏕࠷ࡒ G51 $L ࠉ࠽ࡻࡦථࡿᡘࡊ࡞ࡻࡾ⨠ᥦ偆┞⿭假ࢰࣁࢤࡡ SVE3 㐿ఎᏄࣆ࢒࣐࣭ࣛࢅᙟᠺࡌࡾࠉ
ࢡ࣭ࣜࣈ , $% ࡛ࢡ࣭ࣜࣈ ,, $)) 㐿ఎᏄ࡞ᑊࡊࠉセ㍉Ⓩ┞⇆ᛮ࠿఩࠷ v㟸⩳ズ㡷ᇡࢅᵾⓏ࡛ࡊࡒ 51$L ࢅ⾔ࡖࡒࠊࡐࡡ⤎ᯕࠉࡐࡿࡑࡿࡡⓆ⌟࣊ࢠࢰ࣭ࡢࠉࢡ࣭ࣜࣈ≁␏Ⓩ࡝㐿ఎᏄⓆ⌟ᢒโࡊࡒࠊ % ࡈࡼ࡞ࠉv㟸⩳ズ㡷ᇡࢅ㝎࠷ࡒᙁโⓆ⌟ࢤࣤࢪࢹࣚࢠࢹࢅ
්ᑙථࡌࡾࡆ࡛࡞ࡻࡽࠉහᅹ 3VE3 ࢅ࢞ࣖࢗࣛ&X 3VE3 ࡝࡜࡚⨠ᥦࡌࡾࠉࡵࡊࡂࡢࠉᢒโᙟ㈹ࢅࢰࣁࢤ 3VE3 ࡚┞⿭ࡌࡾࡆ࡛࠿ྊ⬗࡚࠵ࡖࡒࠊ㸨㸣 ◂✪ ᠺ ᯕ 51$L ࢅ⏕࠷࡙ࠉ㧏➴᳔∸ࡡラฦゆ㓗⣪Ⓠ⏍ཬᚺ࡞࠽ࡄࡾ 3VE3 ࢰࣤࣂࢠ㈹ࡡฦᏄ
ᶭ⬗ࢅ᪺☔࡞ࡊࡒࠊࡈࡼ࡞ G51$L ࡛࠷࠹ᡥἪࢅ᳔∸࡚ิࡴ࡙ᚺ⏕㛜Ⓠࡊࠉ3VE3 ฦᏄࡡ㔖Ⓩ偆㈹Ⓩ假ᨭን࡞ࡻࡾ᳔∸ࡡකྙᠺ⬗ࡡᨭን࠿ LQYLYR ࡚ྊ⬗࡚࠵ࡾࡆ࡛ࢅ♟ࡊࡒࠊࡐࡡ㐛⛤࡚ࠉES ࡛࠷࠹ᴗࡴ࡙▯࠷ᵾⓏ㒼าࢅ⏕࠷ࡾ᩺ぜ 51$L ࣊ࢠࢰ࣭ࡡ㛜Ⓠࡵ⾔ࡖࡒࠊࡆࡡ࣊ࢠࢰ࣭⣌࡞ࡻࡽࠉ᳔∸ 51$L ࣊ࢠࢰ࣭ᵋ⠇࠿ࡻࡽᐖ᪾࡞
࡝ࡾ࡛࡛ࡵ࡞ࠉG51$L ࡞࠽ࡄࡾᵾⓏ㒼าࡡ㐽ᢝࡡᖕ࠿ኬࡀࡂᗀ࠿ࡾࡆ࡛࠿᭿ᙽ࡚ࡀࡾࠊࡱࡒࠉࢨࣞ࢕ࢽࢻࢫࢻ࠾ࡼක໩Ꮥ⣌ᅒ偐㸯G5 1$L ࡞ࡻࡖ࡙షᠺࡈࡿࡒᙟ㈹㌷ᥦ᳔∸ࡡක໩Ꮥ⣌ , , Ὡᛮ ྘ᙟ㈹㌷ᥦ᳔∸ࡡකྙᠺὩᛮ偆ක໩Ꮥ⣌ IIὩᛮ㸯Fv/Fm ೋ假ࢅࢠࣞࣞࣆ࢔ࣜ⺧කฦᯊ࡞ࡻࡖ࡙ฦᯊࡊࡒࠊᕞഁࡢࠉྎഁ෕┷ࡡ᳔∸ࢅࠉක໩Ꮥ⣌Ὡᛮೋ࡚Ⰵฦࡄࡊࡒࡵࡡ࡚࠵ࡾࠊ㉝
࡞㎾࠷⛤Ὡᛮ࠿㧏ࡂࠉ㟯࡞㎾࠷⛤Ὡᛮ࠿఩࠷
ࡆ࡛ࢅ♟ࡌࠊࡆࡡࡻ࠹࡞ࠉPsbP ࡡⓆ⌟㔖ࢅን࠻ࡾࡆ࡛࡚ࠉずࡒ┘࡚ࡢ⇆ࡋ࡞ず࠻࡙ࡵࠉᵕࠍ࡝ක໩Ꮥ⣌ II Ὡᛮࢅᣚࡗ᳔∸ࢅషࡽฝࡌࡆ࡛࠿ྊ⬗࡚࠵ࡖࡒࠊ,, Ὡᛮࡡ᩺ぜ࡝โᚒᅄᏄࡡು⿭ࢅ⇆ᏽࡊࡒ偆ㄵᩝᢖ✇୯假ࠊ㸩㸣 ௑ᚃ ࡡ ㄚ㢗 ࡛Ⓠ ᒈ ᮇ◂✪࡚షᠺࡈࡿࡒᵕࠍ࡝කྙᠺὩᛮࢅᣚࡗᙟ㈹㌷ᥦమࡢࠉᵕࠍ࡝⎌⑧ୖ࡞࠽ࡄࡾ᳔∸మහࡡ࢙ࢾ࣭ࣜ࢟㞺⤝ࣁࣚࣤࢪ

Physiol.139, 1175-1184 (2005).

ࡡᐁឺࢅゆᯊࡌࡾ୕࡚ࠉ㟸ᖏ࡞Ⰳ࠷࣓ࢸ
ࣜ⣌࡛࡝ࡾ࡛⩻࠻ࡼࡿࡾࠊୌ᪁࡚ࠉ51$Lࠉ

偐㸣Ishihara S, Yamamoto Y, Ifuku K, and→ࡦࠉG51$L ࡞ࡻࡖ࡙షࡽฝࡈࡿࡒᙟ㈹㌷

Sato F. Functional analysis of fourᥦ᳔∸ࡡ⏍⫩ࣂࣆ࢚࣭࣏ࣤࢪ࡞㛭ࡊ࡙ࡢࠉmembersࡆࡿࡱ࡚ࡡ࡛ࡆࢀ㔕⏍ᰬࢅ෻㥑ࡌࡾࡵࡡphotosystem II in Nicotiana tabacumࡢᚋࡼࡿ࡙࠷࡝࠷ࠊ௑ᚃࠉᚋࡼࡿࡒᙟ㈹using differential RNA interference.

㌷ᥦమ࡞㛭ࡊ࡙ࠉࡈࡼ࡞␏࡝ࡾ⎌⑧᮪௲

Plant Cell Physiol. 46, 1885-1893ୖ࡞࠽ࡄࡾ⏍⫩セ㍉ࢅ⾔࠹࡛࡛ࡵ࡞ࠉࡱ

(2005).

ofthePsbP

familyinࡓゆᯊ࠿༎ฦ࡞㐅ࢆ࡚࠷࡝࠷ 3VE3 ࢅ㐛๨㸨㸣Ifuku K, Nakatsu T, Shimamoto R,Ⓠ⌟ࡌࡾ᳔∸ࠉ→ࡦࠉ␏⛸᳔∸⏜ᮮࡡ 3VE3

Yamamoto Y, Ishihara S, Kato H, SatoࢅⓆ⌟ࡌࡾ᳔∸࡞㛭ࡊ࡙ࠉࡻࡽリ⣵࡝ゆF. Structure and function of the PsbP

ᯊࢅ㐅ࡴࡾᚪこ࠿࠵ࡾࠊࡱࡒࠉࢨࣞ࢕ࢽ

protein of photosystem II from higher

ࢻࢫࢻ࡚ずฝࡈࡿࡒ᩺ぜ࡝ක໩Ꮥ⣌ ,, โplants. Photosynth. Res. 84, 251-255

ᚒᅄᏄࡡು⿭࡞㛭ࡊ࡙ࡢࠉࡐࡡฦᏄᶭ⬗(2005)

ࡡゆ᪺࠿㟸ᖏ࡞⮾࿝῕࠷ࠊ௑ᚃࡵࡆࡿࡼ

㸩㸣Murakami, R, Ifuku K, Takabayashi A,ࡡ◂✪ࢅ⤽⤾ࡊࠉ᳔∸ᶭ⬗ྡྷ୕࡞࠽ࡄࡾ

Shikanai T, Endo, T, Sato F. Functionalකྙᠺࡡ࢙ࢾ࣭ࣜ࢟౩⤝⬗ㄢ⟿ࡡ㔔こᛮ

dissection of two Arabidopsis PsbO

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proteins: PsbO1 and PsbO2. FEBS J.

ࡖࡒ 51$L ࡡᢇ⾙㛜Ⓠ࡞㛭ࡊ࡙ࡢࠉ᳔∸⛁272, 2165-2175 (2005)

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