光合成研究Ѩ 27 ɥ Ѩ 2 Ƥ‫֊ت‬ɥ 79 Ƥ‫ث‬2017 ɲ 8 ͅ
NEWS LETTER Vol. 27 NO. 2 August 2017
THE JAPANESE SOCIETY OF PHOTOSYNTHESIS RESEARCH$Ň¬ ̟͐ĿƧˋȝĈ Α͌Ĉׂ֜ˬ]:йsK



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ƿǃ ÉµÂ~¸ÉÆλZruΧȘ CO2 ʩѭԠ͛;sԘŖEvO̗OY͹ϿВǭʩѭ΍΂ؓ΋ɂ ú‫ت‬ÞɞǼ‫ث‬84ֹ͍‫ם‬ç ú‫ت‬ɒ˒Ǽ ú‫ث‬94

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ĿƧˋоѝ 27 (2) 2017

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ƿǃ Rubisco ]̈ǶZru C3 Ǜ͹Ͽ]ĿƧˋ΍Ӆ̈Ә]Ԭjֹ͍‫ם‬ç  ‫خ‬ƴЖˁß  ‫خ‬Ͼִ ư  ‫ ت‬ɒ˒Ǽ/ ͙ƂǼ/ ‫ث‬94ƿǃ É:raÆӅŮ]̈ǶZru͹Ͽ§~µГДӅʎƁЖ֨Ԧѣ  ‫خ‬Жӣë̋ȣ  ‫خ‬ӭάʓ  ‫·خ‬Ж̧њ  ‫ֳخ‬ɑ ˋ  ‫ ت‬րТǼպȝ֨/ ѫμǼ/ ‫ثͨت‬͹Ͽ¦~/ ‫ث‬103

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ʑǘɳΟ‫͙ت‬ƂǼ‫ث‬*ՓăӵҸ (corresponding author)

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ĿƧˋоѝ 27 (2) 2017

ǎ 1. ĿƧˋ‫ר‬ȗĉ֕Ҋէ]΅ʇǎÁ¢|‫ר‬ȗĉ֕Ҋէ (H2O ;s NADP +) ^қиƍ/2 S] PSI ~Á‫ר‬ȗĉ֕Ҋէ^ՠиƍVфEv
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PSI WգŘȗԕƧĔyʐˋGU7u0ÔԕƧĔWm¬Ã Æ (Fd)y‫ר‬ȗĜÑĔWGU®À¥Æ (PQ) y֡ļIu0¼É ~Á‫ר‬ȗĉ֕Ҋէ(H 2O ;s O 2)^‫ש‬иƍVфEvU7u0Water-water ~ÂV^/PSI Zru O2 ֡ļZrtГHuÉ¨É
(O2-)
yÉ¨É }·É (SOD) <ÐǙƁG/ГHO֓֯ƁΨ҄ (H2O 2) y|ÂªÆ֯±ÂÉ<ΨeǶ˼G/ϳ΢ƁIu0Flv ^/|¥§Á|;sԓȗ͹ϿZģȚEvu</ԍȗ͹ϿV^ȁxvU7u0In vivo Z:7U Flv ^ NAD(P)H iO^ Fd y‫ר‬ȗĜÑĔWGU O2 yΨe 4 ‫ר‬ȗ֡ļ
IuWҷ9svU7u0PSI ~Á‫ר‬ȗĉ֕W¼É ~Á‫ר‬ȗĉ֕^ NADPH yГjŖI
CWY?/pmf ʐˋy÷GU ATP ГДZȳÑGU7u0FNR; ¬Ã Æ-NADP +ÃÉ/PC; ®
À|¢Æ0Shikanai and Yamamoto (2017)26)ÿǶ0yҊUŎɻ PSI ZːEvu (ǎ 1)0NADP+֡ļy<ĐÏGOǧƧ/ΔpH ^ǮǼIu0ÂÉ¸Æ֯ċxJZ Cyt b6f ԕƧĔ] Q ~ÂZrt ΔpHʯƁ^/Cyt b6f ԕƧĔςʯ]ÆÃ¼ÃÉ

¾ÆZrt PSII ;s PSI e]‫ר‬ȗĉ֕֋ɻy

ʐˋIuCWV/ATP/NADPH Σ]ԼѲZȳÑGŢ‫׏‬Iu0րɲ/C]*photosynthetic control”W

U7u0PSI ~Á‫ר‬ȗĉ֕^/PSI ;smƲ_vu‫ר‬ȗĉ֕Ţʞ</ʎĿ:raĿʎɻ<]‫ר‬ȗy‫ר‬ȗĉ֕ҊէZːICW;s/‫ר‬ȗÆǶƁIuǶŸĿÏV] PSI ĿĵȮ˙ŢZֳԗY

WGU PSI ;s‫ר‬ȗyֆIȡŃʆ]΍Ӆ^˩

0PSI ĿOY70Ä~£¡¡yƬkԍȗ͹ϿV^/2ĵȮÈ˙Ţ¸¢·]Ա҆Z‫ׇ‬GU^/֓Ɣ]S]ЂѢGO PSI ~Á‫ר‬ȗĉ֕Ҋէ<ʕŬy͝OICW<ǦƯEvU7uԠԸyƕҷZGU‫=׳‬O7

10,11‫ث‬8,9‫ث‬

Ț Ǖ G U 7 u 0 PROTON

0ВǭǶƁoӴ

REGULATIONқĔýՄ]ǶƁZrtԗΪEvu ATP/NADPH

NADH dehydrogenase-like (NDH) ԕƧĔ 14,15)] 2

¬]̈́֙ƁZ/ǎ 1 ZфGOÂ¡}­‫ר‬ȗĉS]¬Ã  Æ -® À    ¥ Æ Ã  É (FQR) ZĝȚGOҊէV6u (ǎ 1)0֕<ȴ˷Z‫ׇ‬xRU7u0

PGR5/PGRL1 ĝȚҊէ^/M]|Æµ~Æ A

2. PSI ~Á‫ר‬ȗĉ֕Zru pmf ʐˋWĿ

ZȼIuʾƝʯ;s Arnon sZrtѾ 50 ɲŧZǦƯEvO Cyclic photophospholylation WƩÊҊƧˋŢʞ Â¡}­‫ר‬ȗĉ֕] 1 SV6u PSI ~էV6uWҷ9svU7u 12,16,17)0in vitro Z:7Á‫ר‬ȗĉ֕^/PSI ]jV‫؍‬ŸEvu0PSI;sσГGO‫ר‬ȗy¬Ã

(PGR5)/PGR5-Like

Photosynthetic Phenotype 1 (PGRL1)ԕƧĔ 12,13)WΣ^ǶŸIu</pmf ԼѲy÷GOĿ‫׊‬ʞWĿƧˋ CO2 Ǐȧẏ9u ATP ГДW]ÃÉ

5

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U/ÁÆª¡Æ PGR5/PGRL1 ԕƧĔӎĔ<

FQR ςʯyфICW;s 18)/ЌǕ] PGR5/PGRL1Æ÷GU®À¥Æ®ÉÂZτG/‫ר‬ȗ^ Cyt b6f ԕƧĔĝȚҊէ¹ÂV^/PSI ;s] 2 ‫ר‬ȗѓŸZċ75

ĿƧˋоѝ 27 (2) 2017
̘/pgr5 ΐ˿ͨZ:7U ECS Ԡ͛ZruÀ

7/Q ~ÂZrt 4H+<ÂÉ¸ÆZѓŸIuWҷ9svU7u(H /e =2)
+

-

4,17)

~

0

Ӌ]®ÄÆĉɃЇ gH+]Ĭ<ִГͨZȼ m8ÊS]ҊէZ‫ׇ‬ÑIuӴқĔ NDH ԕƧĔGUǼ=?YuCW;s/ATP Ƨˋ֮҄]ςʯ(NADH dehydrogenase) ]бƩÆ¨՞V6t/u 27)0iO pgr5 ndh çֳǶСͨV^ĿƧˋ‫ר‬ȗ^/Ʋƭֿ]®ÄÆ´Æ®V6u Complex Iԍȗ͹ϿV^ 2 S] PSI ԕƧĔWбȼŘȗֵ 1

Ɓ< pgr5 ǶСͨ]ԋЌǛ]ƑǊWIuՇԿm6
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NDH ĝȚҊէmĿƧˋȧɭЁʿZ:7UV΍Ӆ

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Zrt Complex I V^ NADH ֯ƁZŇʕGU 4H

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Ŀ:raʎĿÏV^/ִГͨZȼGU pgr5 ǶСͨV^ PSI Ŀ‫׋‬Ȯ<rtʎ?ГHu 9,12,30)0C]

қĔ NDH ĝȚҊէV^/NDH ԕƧĔy÷GO

ƑǊWGU/ATP/NADPH Σ]ĐÏZru PSI Ɲ

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svU7u (H /e =4)
+

-

4,17)

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~Á‫ר‬ȗĉ֕Ҋէ]̘< PGR5/PGRL1

ʥ P700 e‫ר‬ȗ<τłÈϤОIuCW< PSI Ŀ˿ĝȚҊէrtmŵЇЩZ pmf ʐˋZȳÑV=uĵyʈ=բCGU7uWҷ9svU7u 5)0

CWZYu0G;GY<s/ӴқĔ NDH <®ÄÆ´Æ®ςʯy͆Iu;X8;^͎PȪؐЩ
ZԪ̧EvU7Y70
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O pgr5 ǶСͨV^ÂÉ¸Æ֯ʯƁZĝȚIu

NPQ <Ǽ=?ĐÏIu</ndh ǶСͨV^ NPQ
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ǎ 2. pmf ~W pmf ˋŘZȼIuÂ¡}­‫ר‬ȗĉ֕]ŵִ͝Гͨ (WT) /pgr5 ǶСͨ:ra Flv ХЌʐ՞կ˼ͨ (pgr5+35S;Flv)] pmf ~W pmf ˋŘy ECS Ԡ͛ZrtԫĞGO0ECS Ԡ͛]Ŀʎɻy 134 μmol photons m -2 s-1‫ل‬1,051 μmol photons
m-2 s-1 V Ƕ Ɓ E K O 0 Shikanai and Yamamoto(2017) 26)ÿǶ0ǶСͨV^/ʌĿÏZ:7UĿƧˋ‫ר‬ȗĉ֕֋ɻ<˙Ţ/؅ƴIu

12)0Cvs]ԋЌǛ^/pmf ĐÏ Z r u ΔpH ʐ ˋ ‫ ت‬Â É ¸ Æ ֯ ʯ Ɓ ‫ ث‬W
ATP/NADPH ΣŢʞ]ΐ˿ZբǊIuWҷ9s
v/PGR5/PGRL1 ĝȚҊէ<ØԗY PSI ~Á‫ר‬ȗĉ֕ҊէV6uCWyфGU7u0Ê

76

ĿƧˋоѝ 27 (2) 2017
3.  ¼ É

 ~  Á   (pseudocyclic) ‫ ר‬ȗ ĉ

΃

 ¼É

~Á‫ר‬ȗĉ֕^/֯҄y PSI]8Q/Flv1 W Flv3 ]|~¬É·V΂ˋE

֕(Flavodiiron protein (Flv) Zru֯҄֡ļ| Synechocystis sp. PCC 6803 V^/Flv ¬{¶ÁÉvu¯ÄÁµÉ< PSI ƝȱĔİV]Ŀĝ‫ר‬ȗƝȱĔWGOĿƧˋ‫ר‬ȗĉ֕ZĝȚGO֯҄֡ļV6u0PSII V]ΨŘԠVГHO 2 ‫ר‬ȗ]Ț֯҄֡ļZ΍ӅG 37)/ǶŸĿÏZ:7U PSI

ѓŸZċ7/Äµ;s 6 Řȗ] H+<ÂÉ¸ÆİZѓŸG(H /e =3)/À~+-¸~Æ;s΂ˋEvu 39)0|¥§Árt‫ר‬ȗyֆIȡŃʆWGU PSI Ŀ‫׋‬Ȯy˙ŢӋ‫ ׆‬ΔpH y

GU7u 40)0in vitro Z:7U/Class C Flv ^ʐ ˋ I u 0 PSI  ~  Á   ‫ ר‬ȗ ĉ ֕ Ʃ ΃ Z

NAD(P)H y‫ר‬ȗĜÑĔWGU/4 ‫ר‬ȗ֡ļZr

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7u(ǎ 1)0֯҄ХГǛĿƧˋГϿZ:7U/

¨É

yÉ

}·É (SOD)W|Â

Flv1/Flv3 ²¹Ä (FlvA/FlvB) ^|¥§ªÆ֯±ÂÉ (APX)ZrtΨZǶ˼Gϳ΢ƁIu water-water ~Â<йsvu

Á|;sԓȗ͹ϿZiVģȚEvU7u</ԍȗ32)͹ϿV^ȁxvU7u 42,43)0( ǎ 1) 0 PSI  ~  Á   ‫ ר‬ȗ ĉ ֕ Ʃ ΃ /
4. Flv ZruÂ¡}­‫ר‬ȗĉ֕]óϲЩ̈

water-water ~Âm NADP ֡ļyċxJZ
+pmf yʐˋIu0G;G water-water ~Âe]Ƕ‫ר‬ȗ¬À^ PSII ֯҄ХГ] 5%͎ϛV6t/
ATP/NADPH ΣԼѲe]ՑЅ^ķ;V6u

31,33,34)

~Á‫ר‬ȗĉ֕^ςʯ֯҄іy0 ĕ̌/ԍȗ͹Ͽ^֏ƁZ:7U Flv yȁRO];‫ؽ‬ԍȗ͹ϿZ:7U/¼É

~Á‫ר‬ |¥§Á|;s C3 ͹Ͽe]֏Ɓ֓єZ:7U/‫ר‬ȗÆWGU΍ӅIu֯҄ĝȚǛ]Â¡}­‫ר‬ȗĉ֕^/¼É

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EvO 38)0|¥§Á| Flv ^/ĿƧˋГϿZЀ͆] Class C Flv ZŘ‫׿‬Ev/֯҄֡ļZ‫ׇ‬

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ĿƧˋоѝ 27 (2) 2017
pgr5+35S;Flv V^/NPQ ԵɃ<ÖĿɻÏ(100–800ȗĉ֕^ PSI ~Á‫ר‬ȗĉ֕]ý̓Ҋէμmol photons m-2 s-1)V^֨ŘЩZG;бԒEvY

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ϲZ/©¸Á
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uŬƧ<ǮǼGU7O0Cvs]ҋ͝;s/O2)/

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Z:7Um/Flv ĝȚ¼É¤ÂÈÀÆ´ÉÉW֍ŸGUˋŘŢʞy~Á‫ר‬ȗ

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~Á‫ר‬ȗĉ֕<јŸG/ĿƧˋ‫ר‬ȗĉ֕֋ɻyǮŰ
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~Á‫ר‬ȗĉ֕

ǎ 4. ĿƧˋ‫ר‬ȗĉ֕ZȼIu Flv ХЌ]ʔ‫ײ‬ĿƧˋ¨À¸É/(A) ĿƧˋ‫ר‬ȗĉ֕֋ɻ/(B)NPQ ]ĿʎɻĝȚʯ0ĿƧˋ‫ר‬ȗĉ֕֋ɻ=PSIIֵȗƛЇ(Y(II)) Χ Ŀʎɻ(μmol photons m-2 s-1)ZrtΪlsvO0Yamamoto et al (2016) 43)ÿǶ0(H /e =3) ]̘< PGR5/PGRL1 ĝȚ PSI ~+-Á‫ר‬ȗĉ֕ (H+/e-=2) rtؓ7Wҷ9sv
U 7 u 17) 0 G ; G Y < s / ִ Г ͨ Z ȼ G U

78

ĿƧˋоѝ 27 (2) 2017
5. Flv Zru PSI ǶŸĿҹʯûÑ

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EKO͹Ͽ]Ӵ‫׬‬ϖɻyϗȧG/M]ɣŘyƜuԼѲEvu0ʜRUΧȘ^͹Ͽ]Вǭʾй¸¢W/Ӵ‫׬‬ϖɻǶƁyΧȘ‫ׅׄ‬ʩѭʯWGUԫĞI·yԠ̧IuÎVֳԗYǇȥV6uWҷ9s

uCW<V=u‫ت‬ǎ 1‫ث‬0ѩҸs^/[CO2]ǶƁ

vu0G;GY<s/M]ŉĔЩYŘȗ΍΂^͎

ZȼIuӴ‫׬‬ϖɻǶƁy˪΄ZǶСͨÁÉPZÐ̧Yϱ<Ǻ70M]ԗǊ]ÊSWGU֝ĉ¢ÆyԆ7/Ǻ̑] CO2 ‫׫‬ʾƝʯǶСͨyƊ‫ע‬ȝЩYԠ͛<CviVǌ‫ף‬V6ROCW<ˬB

GO 1)0EsZ/Ʃ΃]˘λyִГÄ~£¡

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uOl/M]r8YԠ͛^¦É

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ӷֵ̏<ǮŰIuW/ӷ̏WWmZϹmȈxv/2. CO2 ‫׫‬ʾƝʯǶСͨ]Ԡ͛ZrtƩȧEvO

Ӵ‫׬‬ϖɻ^ĐÏIu0ևZ/ΧȘ<ֿׄIuWӷΧȘŢʞǊȗ

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2.1.  Χ Ș Z : A u CO2 ʩ ѭ ] µ   É Ã  ¼¬}Zrt͹Ͽ]Ӵ‫׬‬ϖɻǶƁyƠԚƁIuC‡ÃÉ HT1

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3,4)

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λZrRUƊ‫ע‬EvO

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4)

O/ÁÆ/Ã¢Æ¡É‫׋‬ȮũZrRU
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ht1-3 ^Đ[CO2]<ԵɃIuΧȘ‫ׅ‬ƞ<Ń?բCs¡É‫׋‬ȮũZrRUςʯ<͆ʼZ˙ŢEvOJ/kGwxJ;ZׄHuW78ƚʩ<Ԙsv/

4)

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3,4)

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U7O 4)0ht1-1 ^¡Éςʯ<ϓɅG/ht1-2

xvY;RO

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EvO carbon dioxide insensitive 3‫ت‬cdi3‫ث‬ǶСĔրZГHOOl/iO/ht1-2/ht1-5 ^ѢĔ΂֌]ƑǊ֝ĉȗ^/S Ǜ‫~ד‬Æº¤Â SLAC1<əǰGOOl/¡Éςʯ<Ҕ˩V=Y?

yÉ

YROOlWҷ9svO 4)0iO/ht1-3 ^/|Â³Æ֯/ÁÆ֯շօĔ]֗Ҥ]ÂÄV

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Ү˼EvOCWZrRUк?YROOl/HT1

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ЌEKuCWVςʯy˩SCW<фEvO 9,10)0 SLAC1 ]ςʯƁZ^ N ͏Ѥ‫׹‬ǞZȚǕIuЀէ<ʛ1Z̧s;ZEvSS6u0ΧȘֿׄZֳ
ԗ Y ʕ Ŭ y ͝ O G U 7 u OPEN STOMATA1

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ˋG/CO2 ʩѭʯ]Ԡ͛yԆROҋ͝/HT1 ^΍

ABA ¡Âĉ֕Ǌȗ]ÊSV6u OST1 ^/NÂƏΠ҆ӄyЕ7O in

CALCIUM-DEPENDENT PROTEIN KINASE 6

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ZrRUςʯƁEvu]ZʦԗY‫׹‬Ǟ<րɲ̧

HT1 W OST1 W]‫׆‬ZX]r8YбéėЕ<6s ; Z Y R O 13) 0 SLAC1 ΍ Ӆ ΐ ˿ Ƕ С Ĕ Z

8Yʔ‫<ײ‬6u];^iP̧s;ZYRU:sN ͏ѤW C ͏Ѥ]Ô̘yΐȁEKOÆÀ

SLAC1 Æ¨՞] C ͏ѤiO^ N ͏Ѥ/iO/u];/iO/Mv< SLAC1 ]ςʯƁZX]r
J/in vivo Z:AuͺԪȪؐ<ʦԗV6u0

yɃłG/CO2 W ABA ZȼIuΧȘʩѭʯ

2.2. Χ Ș ׄ ֿ y ‫ ؍‬Ÿ I u ‫  ~ ד‬Æ  º ¤ Âѭʯ]ΐȁ^бԒEvY;RO< CO2 ʩѭʯ^yԼfOҋ͝/7Jv]ʐ՞կ˼͹Ͽm ABA ʩ

SLAC1 ^СYu֨ďV CO2 W ABA ]¡ÂǉʟGO 13)0iO/N ͏ѤW C ͏Ѥ]Ô̘yΐȁEKO SLAC1 ʐ՞կ˼͹ϿyЕ7U HCO3-iyƝȱIu ΧȘ‫ׅ֒ׄ‬ŸZ:7U/M]‫؍‬ŸŮyʐˋIu

O^ ABA ZrRUГHu S Ǜ]‫~ד‬Æ‫ר‬τy]ZֳԗY^Os=yGU7u]<Șջ҆ӄӋϗȧGOWCw/ABA Zru‫~ד‬Æ‫ר‬τ]Х

ZȚǕIu®ÄÆ´Æ®o~Æº¤ÂVГ^όȁGU7O]ZȼGU/HCO3-Zru‫~ד‬

6u0ΧȘ<ؓ[CO2]o ABA yʾйIuWȘջ҆Æ‫ר‬τ]ХГ^ԞϗEvO 13)0EsZ/SLAC1

ӄ;s‫~ד‬Æ]̊Ŗ<Ȏiu0Cv<ѤҜW]ӋՒ֊‫׹‬ǞZȚǕIu|¶¥֯]Ա҆YԠ͛

YRU/҆ӄǹe]Ψ]τŖ<ГH/ΧȘ<ׄH

ZrRU/CO2 ʩѭZ‫ׇ‬ÑGU7uƠӅʯ]6u

2 S]|¶¥֯<ԘS;RO 13)0Cvs]CW;

u0CviV/‫ר‬ΧГЎȝЩY˒λZrt/ؓ
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ABA ¡ÂyЂѢGUƝĤGU7uCW<̧

u R‫ت‬rapid‫ث‬Ǜ]ç֊t<ͺŖEvuCW<ǦƯ
EvU7u

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ͽ‫׬‬V^áϻoؓ[CO2]ВǭZ֙ʩŮ]6u͆Е

GU7uCW<̧s;ZYRO 17)0Ū҆ӄZ:7
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2.3.  Ӌ ï ֊ Ǌ ȗ y ÷ G O Ș ջ ҆ ӄ È Ū ҆ ӄ ]ˬŸyԞȸIuW/Șջ҆ӄZ:AuMvs]
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¶¼¢É¾ÆZru̗G7ΧȘʩѭ΍΂

proton ATPase translocation control 1‫ت‬patrol1‫ث‬Ԙ;s/Șջ҆ӄWŪ҆ӄ;sYuΧȘԏҮZ^/

ǶСͨ^ ht1 ǶСͨWƩ΃/Đ[CO2]͖ĂÏVm

PATROL1 y÷GOΨ-~ÆºÂշօ·<ȚǕIuWҷ9svu‫ت‬ǎ 2‫ث‬0PATROL1

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ƁZċ8ΧȘ‫ׅׄ‬WƩ̰Z PATROL1 ]Șջ҆ɄӄҪОǊȗW‫׿‬čGU7uCW<˸ȧEvO
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֋ɻZ‫ׇ‬ÑGU7uƠӅʯ<фƹEvO0GFP΄Նy̚GO PATROL1‫ت‬GFP-PATROL1‫ث‬Z:

U7uCW<фƹEvO0EsZ/PI3K ‫׋‬ȮũAu҆ӄŌɈǕyԼfOWCw/Șջ҆ӄӋÎW<ΧȘ]̧È̼ʩѭyЀСЩZ‫׋‬ȮIuCWyԘÆŖGO 18)0C]CW^ΧȘ‫ׅׄ‬ŢʞZ:7U/СÉ·ZЁ]΂֌yWRUȚǕG

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GFP-PATROL1 W RFP-AHA1 yŇХЌEKOʐ

PATROL1-OX ]ǗÎ֨]Гֳֵ^ִГͨrtm՞կ˼͹Ͽ]Șջ҆ӄyԞȸIuW/PATROL1

32%ǮŰGO 14)0iO/͜ɸВǭy̆čЩZŎЌ^Șջ҆ӄӋÎZȚǕIu®ÄÆ´Æ®

GOк̰‫׆‬VʎĿWʌĿ<ϫG?łv̓xuĿ

AHA1 y҆ӄ՞İ;sԐ˔QIur8ZɈǕGВǭÏV^/PATROL1-OX WɭZΧȘ<‫ׅ‬7U

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7u slac1/ost1 ^/̈́ǼĿƧˋ֋ɻZ֕Iui

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VZԗIuԵɃ̰‫ִ^׆‬ГͨWΣfкҧEvU

ׄHuW=Z^M]Ê֨yΜGU҆ӄ՞Ōeǉ

7O 19)0PATROL1-OX ^ִГͨ<ΧȘy‫?ׅ‬ƛIuʕŬyˠRU7uWҷ9svu0~¶ÆVrt֋?/Ǽ=?ΧȘy‫=ׅ‬/ִГͨ

PATROL1 ]͹Ͽ҉ҫZ:AuХЌ֨ďyԼfO<ΧȘyׄHu~¶ÆV^ΖɭZΧȘyׄ

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ĿƧˋоѝ 27 (2) 2017
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ĿƧˋоѝ 27 (2) 2017

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94

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45)

0Y:/Cvs]¨ÀʐˋEvY7CWV6RO0Ê̘/ƙȗӴ͹ϿЗ¸ÉÉ]8Q CO2 e]ǡ՞ЀСʯ Sc/o ZS7U

͗ ] ǧ Ƨ Z ^  §  RBCS W ] LS С і ‫׆‬uWԘћmsvU7u

Rubisco ^ʐˋEv/Ě9_ C4 Ǜ͹Ͽ Flaveria^/ĿƲƭ<Ã͖ĂÏV]֓ū¤ÂÉό՗ZȳÑGU7uƠӅʯ<6uOl

46-48)

bidentis ] RbcL yɃłGO‫ך‬Z^ LS Сі‫׆‬/֓ɻZؓ?Y7̘<ȡŃV6uWmҷ9svu0Ȫ‫ך‬/
Rubisco ֵy˙ŢGO~¤Z:7U^ CO2 ԒĺϱǛ]Ѐʯ<ûÑEvU7O 59)0G;G/ʐˋEv

րĲV] PSI ]֡ļЁʿ<î֏G/ʎ7ÃyƝAU7uЁʿZYRU7O

49)

Rubisco ZؓΣςʯÈĐ CO2 ԝƴʯǛW7RO C4
u LS Сі‫ ׆‬Rubisco ]ֵ<ɅY7CWVĿƧˋ֋ɻ]ĐÏ<ГHuW78ƽ‫<׼‬6RO0CvZ

0Cv^ĿƧˋ

WWmZĿƲƭm˙ŢEvOOlV6u0

ȼIuԠέѮ<րɲԘŖEvSS6u0Rubisco

 ²͹ϿZrRU^/րҤ]і;sΣςʯ²Ä֮҄]|Æ­ÂZ^Řȗº±ÄÆ<:ra CO2 e]ǡ՞ЀСʯ Sc/o <WmZؓ7‫ׇ‬ÑGU:t 60)/Ě9_¹Ä] Rubisco

k

c
cat

ֵ < ϓ Ʌ G O ¶ ¼ É  Æ  V Bundle Sheathm]y֜dCWmƠӅV6t/Ɂ͗ЩYСі

Defective 2 (BSD2) 61)/Rubisco Accumulation Factor

Rubisco ]Ƀłyҷ9OǧƧZm֙GU7u0

1 (RAF1) 62)/Rubisco Accumulation Factor 2 (RAF2)·֍Z:7U^/·W»·]
Rubisco <·] Rubisco ZȼGC]r8YЀ<ԘŖEvU7u0Cvs^ RBCS WÊ֓Щ63)

ʯy˩Q/· Rubisco WҮ˼GOǧƧZ^̈́

YԕƧĔyʐˋG/º±Ä¢ÆV¬ÉÂ}

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50)

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uOl/Σςʯ kcat <ؓ7 Rubisco ]̘<͆ŠVu‫ך‬Z/Ä~£¡¡] RAF1 ֝ĉȗyƩ̰Zc6uW]ҷ9̘;s/C4 ͹ϿoȶœǗoؓɏZ
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37,52,53)ɃłIuCWV/Rubisco ֵ/ĿƧˋ֋ɻ:raĨĔ§~µֵ]ĐÏ<6uєɻҠƴEvO

00C]CW^/Сі RbcL WCvZ֙GOŘȗ64)

EsZ^/|¥§Á|<˩SÂ³
É·oԂ‫˩<׿‬S«Ã¥~

º±ÄÆmƩ̰ZɃłIuCWV Rubisco ֵ]

W7RO CO2 Ϭҧ΍΂yҔѱ͕͹Ͽ]ӴқĔZ Rubisco DWѓ͹
Gr8W]Ԭj<YEvSS6u

54-57)ϓɅy‫@׊‬CW<V=uCWyʼƱIu0OPG/
C]оѝĚV^§] RBCS ZȼGėЕIu0M]úZ

]^Ä~£¡¡] RAF1 WYRU7uOl/m/ȪؐȬÃ°ÂV Rubisco ]֮҄ЩЀʯy֏Ɓ
EKuCW<V=u;W78Ԭjm6u0Ɵ҆Ӯŧփ]r8Yƚʩ΍ė 63) <Ȫ‫ך‬ZėЕGU7u

Methanococcoides burtonii ]‫׫‬ĿƧˋ Rubisco Z;X8;^Ð̧V6u0

ÀÆ·Y|¶¥֯ǶСyГHEKOm]]Ö

 Ҕѱ͕͹Ͽe]Â³É·]ѓ͹Z:;s/6uєɻΣςʯ kcat Z̈Ә<jsvO

7U^/ŧփ]r8Z|¥§Á|] RbcL

0y§ZɃłGUm²Ä֮҄<ʐˋEvY7

c

Rubisco <֜˜EvO

58)

W78ƽ‫<׼‬6RO0CvZȼG/§ӴқĔ4-2) СіГϿЗ͗ Rubisco Ƀł]Ԭj

Z Synechococcus elongatus PCC7942 ]ǼɄ­½

s/²͹Ͽ] RbcL ]Ү˼]Ԭj<֏lsv]֓єVėЕIuº±ÄÆ rbcX 65) yƩ̰Z

 Rubisco ]ςʯÖʥ^ RbcL ZȚǕIuCW;¢֝ĉȗW Rubisco ²Ä֮҄]|Æ­Â

U=U7u (̔Ѕ 53) ƕ϶)0²͹ϿWGU^

ɃłGOWCw/ʐ՞կ˼Ĕ] Rubisco ֵ^ִГӴқĔʐ՞կ˼<ΣղЩǫ̇̄Y§<ØWGǛrtmǼɮZĐÏGOm]]Ƀł֝ĉȗЗ͗

UЕ7svU7u0CviVZƽ‫׼‬ZYRU=O] Rubisco ]j<ӸћGU7O 54)0Y:/rbcX y]^/RbcL <|¥§Á|oѿԂ/ƊȗӴХЌÆÀ;s‫ב‬7OǧƧVmƩєɻ97

ĿƧˋоѝ 27 (2) 2017] ֵ ] Rubisco ² Ä ֮ ҄ < Ӹ ћ G O 55) 0 S.

çֳǶСĔZÀÆ±®elongatus PCC7942 V rbcX y¥|GUm

Rubisco ²Ä֮҄^|Æ­ÂEvU:t66)]m]WҮ˼GOÀ¶y RBCS1A¹¡] RBCS ֝ĉȗyɃłGOWCw/Σςʯ kcatc W CO2 e]ǡ՞

/7Jv]ǧƧVmŌǕʯ RAF1 <²Ä֮҄]|ЀСʯ Sc/o <WmZĐÏGO 57)0C]r8Z/Ƀ
łIu Rubisco ]֮҄Щʯ՞<ɭZ LS Сі‫׆‬Æ­ÂyˠRU7O]V^Y7;WäϗEv

Rubisco Zƚ̪EvuW^‫׏‬sY7r8V6u0

U7u0Â³É·]΂ˋԗ҄]Ʃ̰ХЌ
ZS7Umоѝ<֏zV=U:t/Rubisco ֝ĉ

5. :xtZȗWWmZÂ³É·ʐˋ‫ׅ‬Ȏ̰]
Rubisco ]ֳƧyˠ8 ccmM35 ֝ĉȗ67)

 ÿÎ/Rubisco ]̈ǶZru C3 ǛĿƧˋ]΍ӅyƩ̰ZɃłIuCWV/ӴқĔŌV Rubisco <ֳƧIüǶZ‫ׇ‬Iǘր]ʸǦy҇÷GO0Rubisco ֵ

0EsZ/Â³É]˙ŢZrtؓ CO2 ВǭZ֙GO~¤]ėԔZ·]΂ˋԗ҄]8Q/Ξy΂ˋIum]yƬk7^ˋůGOm]]/ĿƧˋ:raĨĔ§~µ?S;y§È°Æ¶|¡VÊ֓ЩZƩ̰ZГД]̈ƿ^ӵG7m]V^Y;RO0EsYu

ХЌEKOWCw/Â³É·΃]΂֌<̈ƿ]OlZ^/Rubisco ÿǹ]ĿƧˋyʙ֋I

CW<ԞȸEvO

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68)

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uǊȗZֳϱЩZѠ҄yŘ֬Iuʦԗ<6u0ÄµVmÂ³É·<ʐˋV=uƠӅ

Rubisco ֵyǮʎIuǧƧV^/Mvs]Ǌȗy

ʯyфGU7u0OPG/M]OlZ^Ʀ΂ˋԗ

̧s;ZGY7W‫؁‬ժЩYĿƧˋ΍Ӆ̈ƿ^͌҄y֙Ζֵ/֙ΖYֵΣVХЌEKuʦԗ<6uʘV=Y70W˪́EvU7u0

 Rubisco ]֮҄΍Ӆ]̈ӘZ:Auƽ‫^׼‬/Ӵ СіГϿЗ͗] RBCS ]Ƀł^֓ƔZÀ¶қĔʐ՞կ˼yԆ8CW<V=u͹Ͽі<‫׏‬s¹¡Z:7UYEvU:t/²ÃÆ/vU7uW78CWV6t/ЀZėϿZ:AuȪ

Ä~£¡¡/©µÅÁW7ROҔѱ͕͹Ͽ]ؐѼ]‫ׅ‬Х<͊ivu0RbcL mG?^ RBCS ]RBCS VҮ˼GOWCw/Cvs] Rubisco <˩Ү˼Z‫ׇ‬GU^/²͹ϿWСіГϿ]бʯ]SĐΣςʯÈؓ CO2 ǡ՞ЀСʯW7ROʯ՞<ӘEZ‫ׇ‬IuŘȗЩǡЮWGU/‫־‬WYu|¶¥0Ҕѱ͕֯o֨ŘЩ֬Ŝ]ЀȧmʦԗWYu0Rubisco ]͹ϿZ:7UmСіГϿЗ͗] RBCS ]ɃłZȢŃҮ˼mԚִZłvuW/ŌǕʯ] RBCS ֝ĉ

S7U]йԘ<ӸћGSS6u0~¤Z:AuĿȗ]ХЌy7;Z˙ŢIu;<Ի‫׼‬WYu</CƧˋ΍Ӆ̈Ә]OlZ C4 ͹ϿV6uÂ·З]OlZ^¥·ҟ‫˘ן‬ԇ<ʦԗWYu0M]Î

͗]ؓΣςʯÈĐ CO2 ǡ՞ЀСʯ Rubisco ] RBCS

V/͊kr8Y Rubisco yʦԗֵPAХЌEKu֝ĉȗyɃłGOWCw/ŃɄ­½¢]8

Ol]ŘȗЩYǡЮyԠ̧G̅ėGYAv_Y

Q 30-79%<Â

sY70öǉ^ԡvY;RO</Rubisco |}LS Сі‫ ׆‬Rubisco Zƚ̪EvU7O

69)·З͗]m]WYRO҉˼9°Éy÷GO Rubisco ]ςʯƁŢʞ]ƽ‫׼‬mГ

Ĕ<ʝsv/Rubisco ]֮҄Ѐʯ<ؓΣςʯÈĐ
CO2 ǡ՞ЀСʯZǶƁGU7O

52)

0OPG/ĿƧ

Hu0Cvs]Ի‫׼‬yԠέIuiVZ^/iPi

ˋ֋ɻ<ǮŰIuCW^Y;RO0M]ЎЗWG

Pգ9Y?U^YsY77?S;]ǯ<6u0

U / Rubisco ] Σ ς ʯ : r a ֵ ] Ǯ Ű Z r u
Rubisco ]ӅŮ]ǮŰZȼGU‫ר‬ȗĉ֕ӅŮ<ÐՄչ

զGU7OCW<˪́EvU7u0M]Ê̘V/ öǉ҇÷GOоѝĚ]Ê֨^/ѐо՗~¤ZĐϖҹʯ] C3 Ǜ͹ϿV6u¹ÉЗ͗(26450074, JP16H02538) :ra JST-CREST (͹Ͽ]ؓΣςʯÈĐ CO2 ǡ՞ЀСʯ Rubisco ] RBCS‫׸‬įʯ) ;s]ԒŲVԆxvO0CCZƐ?ʞх֝ĉȗyɃłGUm Rubisco ]֮҄Ѐʯ^Ǽ=?
ǶƁIuCW^Y;RO70)ЙGÎBu00iO/Ä~£¡¡VØԗZХЌIu RBCS1A :ra 3B ֝ĉȗ]98

ĿƧˋоѝ 27 (2) 2017
Received May 31, 2017; Accepted June 12, 2017;12.

Published Aug 30, 2017

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Published Aug 30, 2017ƕҷ̔Ѕ

ԆRU7u~Æ

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¹ÉÉyЕ7uæ<ŵ͝ЩV6uCWm̧s

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1,4

1,4

2,4

3,4

Noriaki Tanabe , Masahiro Tamoi , Akira Kikuchi , Akiho Yokota , Shigeru Shigeoka
1

2

1,4

Faculty of Agriculture, Kindai University, Graduate School of Life and Environmental Sciences, University of3

4Tsukuba, Plant High Technology Institute, CREST/JST

109

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ƹGU7u0Hikosaka 11) ^¶¼ÃÉ¾Æy

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Z6ORÜӘƠӅYÉV6uW͌ʘ

v]ďҮVɌ‫ׅ‬G/ĿВǭ^6uєɻǏȧEvU

Evu0

4)

7u]V/Ӵ]‫׆‬V^Ǽԙ΅YѠ҄Ŏկτ^բ
CRU7Y7Wҷ9svu0Ѡ҄]կτ<

Received May 30, 2017; Accepted June 23, 2017;

Ǽ=Yʔ‫ײ‬ymSYs_/ӧ͐ұӳW͍͐ұӳ]

Published Aug 30, 2017‫׆‬ZѠ҄ƀ֬Z֖7<6uW͌ʘEvu</
Hikosaka et al. 12) ]¸Ԡ͛V^Ǽ=Y֖7<
Y;RO0C]CW^/Ѡ҄]կτ<ֳԗƕҷ̔ЅV^Y7CWyфƹGU7u0

1.

 YLȪ‫]ך‬Ѡ҄Ř֬^̈́֙V^Y7];/W78ƽ7^/öʚmՇԿEvuֳԗYУƽV6uW

2.

ҷ9svu0Aerts, R. and Chapin, F.S. III (2000) The mineral
nutrition of wild plants revisited: a re-evaluation ofprocesses and patterns. Adv. Ecol. Res. 30, 1–67.

Hirose T, Werger MJA (1987b) Maximizing daily

canopy photosynthesis with respect to the leaf
nitrogen allocation pattern in the canopy. Oecologia
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relationships in leaves of C3 plants. Oecologia 78, 9–

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integrated

photosynthesis of cells and leaves. Phil. Trans. Roy.
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Hasegawa, T., Usui, Y., Nakamura, H. and Hikosaka,

Crop Sci., in press.

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and Hirasawa T. (2011) Hydraulic conductance as4067–4077.

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ĿƧˋоѝ 27 (2) 2017

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CEA/CNRS/Université Aix Marseille, UMR 7265 Biologie Végétale et Microbiologie
Environnementales, 13108 Saint-Paul- lez-Durance, France
Yusuke.Mizokami@gmail.com

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ØĴ‫ؼ‬ŅЫՐǋλó

їЬՐǋʚ˾‫ؼ‬ñ֫ɺ/ñ֫ɦ/NHKƈ՛(äȧ)‫̟͐ؼ‬ĿƧˋȝĈ/ĨĔұГʿȝĈ/іГϿȝĈ/̟͐ėϿȝĈ/̟͐͹ϿȝĈ/̟͐͹ϿГЎȝĈ/̟͐̑ЎГϿȝĈ/̟͐ГʿȝĈ/̟͐ǗЍʹ̩ѐȝ֍Ƨƽ7ƧxKľ‫ؼ‬їЬՐǋæŹɈ TEL 075-353-7272
URL‫ؼ‬www.inamori-f.or.jp ν‫ث‬ƕŰZ^Йս<ʦԗVI0Ա҆^ÎԦ  ZU:йsK7OGiI0





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