植物科学最前線 7:28 (2016)  /2   E   (5?
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657-8501 ůęāőÇµšÐŢ
From C3 to C4 photosynthesis: Can the introduction of C4 Rubisco alone be effective
for the improvement of photosynthesis in C3 plants?
Key words: C4 photosynthesis; elevated CO2; gene engneering; rice (Oryza sativa);
Rubisco; sorghum (sorghum bicolar)
Hiroshi Fukayama
Graduate School of Agricultural Sciences, Kobe University
Kobe, 657-8501, Japan?> /2  *"8%6 C4 ĺŔ8 C3 ĺŔ6ń<1²ÑĖƏÁǖ!ǚŝǅƳĆBƳ+H8ǚ5*57-Iǟ ²ÑĖ CO2 áø7¼ŦÌĒKƜò'Gy`-1,5-v`ǁT~V^cSV^Znc (Rubisco)8ǚƜòÌĒƳĆļź6Ʒ#3ǚO2 KBæƦ3%1²ØÔK
Ƨ#'#3Eǚ²ÑĖ75ČƳÝó35/1GC4 ĺŔ8ƶÆ7ƸŴ2 CO2 ŐƊ6°! C4 ²ÑĖÜƨKŗĎ%ǚRubisco 7Ɯòƾ¥27 CO2 ŐĆKǖA1²ØÔKĞ¾'G#36ĖÂ%,C4 ²ÑĖÜƨKǓÅ$)G68 ATP ¨¹6đƘ35Gǚ²ØÔ¬
ƶ$HGǖōCć²28ǚC4 ĺŔ7ĩ C3 ĺŔDFBį½35GǛ
 #7D5 C4 ĺŔ7²ÑĖ^`j8ǚC3 ĺŔ7²ÑĖƏÁKĥƓ'G2įÄ3ƍǚţ2 C3 ĺŔ6 C4 ²ÑĖÜƨK'GÍFƅ@ƖJH1,ǘHibberd et al. 2008,
Miyao et al. 2011ǙŰƑƩBƬĻŝŔƥŏŬŷĚ6Ěþ%1,ǋ6ǚC3 ĺŔ7Op6lP
[^ä7 C4 ²ÑĖÜƨKû³'Gz_QXl6ÊÃ%,lP[^ä C4 ²ÑĖÜƨKĽĖ'GǀƁ8ǚæĲŧ6}`}RqwvǁT~V^cǘPEPCǙǚ
NADP-\ǁkumZncǚNADP-\ǁǀƁǚwvǁǁ_VncǘPPDKǙ7 4 03ü5ǚŕ6 PEPC 3 PPDK 8 C4 ĺŔ61ǎĂ6ǖŦř%1FǚC3 ĺŔ¶2Ò{6ǖŦř$)G#38ÞǍ2/,ė8ǚzeǚRVdǚOlǚepeKÓA,lP[^ŠĶ7 PEPC ƽ¤ó3 PPDK ƽ¤ó
K+7??Op6û³'G3ěŇ6DFǚ#HE7ǀƁKlP[^¡6ǖŦř$)G#36ĖÂ%,ǘFukayama et al. 2001, Ku et al. 1999Ǚ+%1ǚC4 ²ÑĖÜƨ6đƘ5    
BSJ-Review 7:28 (2016)

植物科学最前線 7:29 (2016)

4 07ǀƁ´1KǖŦř'G 4 ǂĉƦƪĤOpK©Ė%,ǘTaniguchi et al. 2008ǙPEPC KÉŖ2ǖŦř$),ĉƦƪĤOp28ǚØÔ¬ƶ$HG#3E²ÑĖƳĆ¦%,ǘFukayama et al. 2003Ǚ%%ǚ4 ǂĉƦƪĤOp28 PEPC 7ǖŦř7ÄĸĜ.Ŋ$Hǚ²ÑĖƳĆǎĉƦƪĤOp7{6Üď%,ǘTaniguchi et al. 2008Ǚ#7řƤ47D
5Toa6D/1Ƨ#G78Ĭŭ285ǚü5!3B 4 07ǀƁKǖŦř$),-"28ǚ²ÑĖƏÁKñă6¬ƶ'G=46 C4 ²ÑĖÜƨKǓÅ'G#38253ƍEH,7YzKÓA1Bǚ73#I C4 ²ÑĖÜƨ7û³6DF²ÑĖƏ
Á7ĥƓ6ĖÂ%,ª85Ǜ@> /2    
 C3 ĺŔ8´ŸƁ7ƀ 30%K Rubisco 6ğƥ%1GǚC4 ĺŔ8 10%¡2GǘMakino
et al. 2003, Sage et al. 1987ǙC4 ĺŔ8ǚ5*ü5 Rubisco 2ǖ²ÑĖƏÁKƹĖ2G7-Iǟ Rubisco 7ŀŒ3%1ƜòƳĆ7ǀƁ6ń<1¦#38ī6ư<,ǚƜòƳĆ68ŵǇĀFǚC3 ĺŔ28ŕ6¦!ǚCO2 ŐƊÜƨKġ0 C4 ĺŔ28 2 ­=4ǖ#3ūEH1GǘSage 2002, Seemann et al. 1984ǙC4 ĺŔ8ƜòƳĆ7ǖRubisco KŗĎ'G#36DFǚRubisco ÓǄKŌE'#3ÏƏ35Fǚ²ÑĖ7ŸƁ½
şÄŘKǖA1G3ƍEHGǘà 1Ǚ%%ǚRubisco 7ƜòƳĆ3 CO2 ƚÚĔ7Ǉ68lmSx7ǈ«FǚC3 ĺŔ7 Rubisco 8ƜòƳĆ8Ʒ CO2 6ú'GƚÚĔǘCO2 6ú'GTR`øĦ Kc 8¦Ǚ8ǖǘTcherkez et al. 2006Ǚ#7D5 C3 ĺ
Ŕ7 Rubisco 7ŕĔ8ǚŞĻǏÙ¡¿7 250 ppm ŴĆ7¦ CO2 ŐĆ61²ØÔKĞ¾'GD6ƶÆƻĒ%,Ɔĸ3Gřâ7ãŚŜêKƙG3ǚñŅ CO2 ŐĆ8ëÃ
7ƱK,4/1Fǚ2050 Ą68 600 ppm ŴĆ6?2ëÃ'G3ĕ$H1G+7D5ñŅĵ¢28²ØÔ87(3Ğ¾$HG#3Eǚč8 C3 ĺŔB C4 ĺŔ7D

7ŸƁ7ğƥKŌE'#3ǚ²ÑĖCŝŞĔ7ǖ©Ŕ7ǆŦ605G3ƍEHG

A> /2=1)#     /2%

"&# !
%(  " '$    !%(  kcat: %(5ǖňĔä Rubisco Kġ. Rubisco ;Kc:              kcat : Kc: 

[CO2]    BSJ-Review 7:29 (2016)

植物科学最前線 7:30 (2016) ė8²ÑĖƏÁKĥƓ'G,A6ǚC3 ĺŔ6 C4 ĺŔ7ǖňĔä Rubisco Kû³'G#3Kƞ@,OpǚWǒǚ_UOǚfOa54Ƙ5©Ŕ7ï!8 C3 ĺŔ2Gǚ+72BOp7 Rubisco 7ƜòƳĆ8¦#3J/1,ǘSage 2002Ǚ+#2ė8ǚOpKùǔĳĨ3'G#36ņAǚ+7²ÑĖƏÁ7ĥƓ7,A6ƻ%,ƜòƳĆ7ǖ Rubisco KƶÆŧ6ƯOpűĺŔ72ĢƂ%,ǘIshikawa et al. 2009Ǚƛķ%,Opű C4 ĺŔ7 Rubisco 7ƜòƳĆ8ǚ´1OpDFBǖ/,ǚdU7 Rubisco 8ƜòƳĆǖ!ǚ0 CO2 6ú'GƚÚĔBńƫŧǖ,AǚOp7²ÑĖ7ĥƓ6ĮBįÄ3ƍEH,Rubisco 8 RbcL 3 RbcS 7 2 ŵǒ7etXƦ6D/1ĽĖ$H1FǚƜò©şKŮ' L8S8 ĽƴKÍG?26ïĦ7¹ó^|37ũ©şKÎ"GđƘGǘŋÿ 2013ǙǛD/1û³'Gƽ¤ó8¸ĶGǉFOp7ƽ¤ó3ũÒĔǖ#3İ?%+#2Ɯòƾ¥ôâ'G RbcL 7Mqǁƿ»601BńƫKƖ/,ǚdUKÓAOpűĺŔ¶28ũÒĔǎĂ6ǖ!ǚ47ŵ7ƽ¤óKû³%1
BñĀ85B73ƍEH,ǘIshikawa et al. 2009ǙǛRbcL 8ƕƉ§ZqǚRbcS 8ĹZq6ïǂƽ¤óĪ3%1ôâ'G+#2ǚ RbcL 8tjNXUK½ş%,ũÒƅĤ6DGƕƉ§ĉƦƪĤŇǚRbcS 8MYsXjPŇ6DFOp7Ĺ;7ƽ¤ó&*

68ĖÂ%ǚǕ!<#36ǚƜòÌĒ6
1+H=4ǂƘ3ƍEH15/, RbcS Kû³'G-"2 Rubisco 7ƜòƳĆƀ 50% ǖ?G#3ĬE35/,
(Ishikawa et al. 2011)  ĉ Ʀ ƪ Ĥ O p 7
Rubisco 7ƜòƳĆ8dU6ń<G3¦ǚOp7²ÑĖƏÁ7ĥƓ6È¹į
Ä2G#3ǚ²ÑĖƳĆ7^^EJ/1GB>'79,    !<( - &*'%#)!")  15ĩǚdU RbcS 7ǖŦř   

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û³KƖ/,ǘà 2ǙOp7ƕƉ§ĉƦƪ        + $       Rubisco#GP
RbcL`bU*24RbcS@*24TL<YJS+0 ! 9,6'4RbcS (SbRbcS) #$)73(/5&4_%1A<YJ[\,6'4RbcS#[\ RubiscoO^RZQ="(Ishikawa et al. 2011)   ! Rubisco?N FIPRZQ= : %1RbcS (OsRbcS2-5) V RNAi_ 2.(-&8#HRubiscoCa#WZEM  ;%1`bUDKXB>]!,6'4RbcL (SbRbcL)<YJ[\.: dU RbcS KǖŦř'GĉƦƪĤOp28 Rubsco 7ƜòƳĆëÃ%,ǚ²ÑĖƳ
Ć6847D5ÄĸKD>'7-Iǟ Ńē5EĉƦƪĤOp7²ÑĖƳĆ8ǖ    
BSJ-Review 7:30 (2016)

植物科学最前線 7:31 (2016)

CO2 ĵ¢2Ŏø%,èÑBëÃ'G#385/,ǘIshikawa et al. 2011ǙĉƦƪĤOp28Op7 RbcS KŃ%,??dU RbcS Kû³%1G,A6 Rubsco ƸÀŦř%1FǚRubisco ÓǄëÃ%1,ǖ CO2 ĵ¢28 Rubisco ²ÑĖ7ČƳÝó28572ǚ
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Ɩ/1GD6 Rubisco ÓǄKü5!%1ŸƁ½şÄŘKǖAG#3ǘSage et al. 1987Ǚǚ
²ÑĖƏÁ7ĥƓ605G3ƍEHGùǋ6ǚǖ CO2 ĵ¢28ǎĉƦƪĤOp6
1B Rubisco 7ƏÁ8ƸÀ2FMgb`ĉƦƪĤOpKş,ƛķEǎĉƦƪ
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 %,/1Ł6dU RbcS ǖŦřOp6

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RbcL

ř%15+%1ŃF7 4 ƽ¤ó (OsRbcS2-5)

OsRbcS
SbRbcS7éæƿ»8ǎĂ6ũÒĔǖĩ

B

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35/, Rubisco 6ń<1ǚ$E6ƜòƳĆ
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RbcL RbcS7[1 @S8A
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YUO.)- R; $RbcS
%! /2B8AM6$ L SbRbcS
RbcL &/#ZRubisco7Z:C  I2
B. 2B8AM6$ =CO2D93;?2B8AM6$0 CO2QND
9 2H I2PG<>HPG5J
    
BSJ-Review 7:31 (2016)

植物科学最前線 7:32 (2016)

Ć35/,%%ǚŸƁ,F7²ÑĖ2ƙG3ǚ2 ǂĉƦƪĤOp8ǎĉƦƪĤOpDFBǖ®KŮ%,2 ǂĉƦƪĤOp7 Rubisco ÓǄ8ǎĉƦƪĤOp7 20-35%2FǚƜòƳĆëÃ%,38ïüÓǄü5'G3ƍEHGčǚRNAi 7lUƿ»7ŕťĔKǖAG547ĥƓKƖǚRubisco ÓǄKǎĉƦƪĤOp7 40-50%ŴĆ6ơ
ż'G#32H9ǚDFĬŭ6²ÑĖƏÁKĥƓ2G3ƍ1G

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+  0;$ 

 #H?27Ŭŷ2ǚRbcS  Rubisco 7ƜòƏÁ7ĥƓ6"GǂƘ5eZhl3ĖF
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5Ǜ$E6 OsRbcS1 8ǚ7 RbcS 37ģøMqǁƿ»7ũÒĔ 55.4%3¦ǘOsRbcS2-5 Ǉ8 100%ǙǚƕƉ§ŲƖ^Ynƿ»Ĭŭ547ƒ×ŋŕĐKġ0OsRbcS1 SdY8llǚ[Y]54Op¡î7ĺŔ61Bôâ%1FǚĺŔ61²ÑĖ2°!ƲĂ7 RbcS 3ť5GĿƏKį%1G#3ĕ$H,ė8 OsRbcS1 6ŪŨ%1Ɵƃ5ŦřƛķǚĉƦƪĤ§Kş,ŝśĿƏƛķKƖ/,ǘMorita
et al. 2014Ǚ+7ƆĸǚOsRbcS1 8ùǋ6ƕƉ§ǘz`gmǙ6ýâ%ǚRbcL 3łĂ6
£Ñ%1 Rubisco 7]yohl3%1ĿƏ%1G#3J/,%%ǚOsRbcS18²ÑĖňĔ7GƉƔƅƋ28Ŧř%1E(ǚƕǐCų7ƈŻĴÖƭ2Ŧř%1,
OsRbcS1 KƲĂŦř%15ƕƩ2ǖŦř$)ǚRubisco ňĔ;7ÄĸKƛķ%,3#Iǚ
OsRbcS1 ƅ@Ʈ?H, Rubisco 8ƜòƳĆëÃ'G#3J/,ƜòƳĆ;7Äĸ8dU RbcS 3ÒŴĆ2FǚOsRbcS1 Bǖ CO2 Ŝê27²ÑĖĔƏÁ7ĥƓ6įÄ2G3ƍEH, řâǚOsRbcS1 7ĿƏKĬE6'G,A6ǚRNAi qhXfPĉƦƪĤOpKş,e~ƛķCl`XzlƛķKƶA1GƲĂ7²ÑĖ38ť5GRubisco 7

ƣĿƏ3%1ǚƈŻĴÖƭ27 C4 ²ÑĖŧ5 CO2 áøǘHibberd and Quick,

2002ǙǚƐƦÑĖ2ŝ&, CO2 7·áøǘSchwender et al, 2004ǙçÕ$H1G#7D
5

ƣ6 OsRbcS1 ǈ«%1GÏƏĔBƍEHGǚ#7èÑ8(HBąƌ27²ÑĖ6Ĉ,Fǚ²RpWK½ş'G#32

ƣƶƖ'GƉƔƅƋ2Ŧř%1

5 OsRbcS1 7ŦřteKƍG3ǚOsRbcS1 8?26çÕ75

ƣĿƏKĠ/1GÏƏĔG3ƍ1G    
BSJ-Review 7:32 (2016)

植物科学最前線 7:33 (2016)

D> C3 ĺŔ7 C4 Æ8ǚė²ÑĖŬŷƎ7ð2GƯĄǚC4 Æ7ùř7,A7ĮBǂñ
5ǌì35/1,XiĽƴ7¹Æ6ǈ'G¹óToa7ěFĎEH1GǘSlewinski, 2013Ǚ%%ǚö´5 C4 Æ7ùř68ǚ?-?-ĭǇđƘ3ƍEHG
Rubisco 7ƜòƏÁ7ĥƓBǎĂ6ÞǍ2FǚǅĄ6J,FùşÏƏ5{27ĖÂª5/,ė8 C4 ĺŔ7 RbcS K C3 ĺŔ6û³'G3ńƫŧžÉ5ěŇ6DFǚ
Rubisco 7ƜòŕĔKǖňĔä6íĤ'G#36ĖÂ%,řâǚRubisco ÓǄ7ĮƻÆ6
ÍFƅL2Gǚ+HĖÂ'H9đ(C²ÑĖƏÁ7ĥƓ605GB73ıċ%1
GdU RbcS 8ƽ¤ŧ6ƺËóƕĺŔ68ƻş253ĕ$HGǚC4 ĺ
Ŕ8ąŽß5ĺŔŵ61ŖŹ%1¹Æ%1FǚËóƕĺŔ61Bï!7ŵ6 C4ĺŔôâ'GǘSage et al. 2011ǙOp¡î6BǂƘ5©Ŕ7ï!8 C3 ĺŔ2FǚĮƻ
5 C4 ĺŔ7 RbcS KƼ:#36D/1ľ5©Ŕ7²ÑĖƏÁ7ĥƓ6ǚÜƄ%,ĘŤƻş2GB73ƍ1G ĲƢħ7eOl8C3 ĺŔ7 C4 ÆǞRubisco -"2BĊ6Ź0ǟ2/,ǚŰ8$E6RbcS -"2BĊ6Ź0Ǘ3Ɲ,

&4-3ŋÿŉ 2013Ǜǖ CO2 Ŝê6ƻ%, Rubisco 7û³6DGOp7²ÑĖƏÁ7ĥƓǛ²ÑĖŬŷǘ23Ǚ: 24-32Ǜ
Fukayama, H., Hatch, M.D., Tamai, T., Tsuchida, H., Sudoh, S., Furbank, R.T. & Miyao, M. 2003.
Activity regulation and physiological impacts of maize C4-specific phosphoenolpyruvatecarboxylase overproduced in transgenic rice plants. Photosynth. Res. 77: 227-239.
Fukayama, H., Tsuchida, H., Agarie, S., Nomura, M., Onodera, H., Ono, K., Lee, B.H., Hirose, S.,
Toki, S., Ku, M.S., Makino, A., Matsuoka, M. & Miyao, M. 2001. Significant accumulation ofC4-specific pyruvate, orthophosphate dikinase in a C3 plant, rice. Plant Physiol. 127: 1136-1146.
Hibberd, J.M. & Quick, W.P. 2002. Characteristics of C4 photosynthesis in stems and petioles of C3
flowering plants. Nature 415: 451-454.
Hibberd, J.M., Sheehy, J.E. & Langdale J.A. 2008. Using C4 photosynthesis to increase the yield ofrice-rationale and feasibility. Curr. Opin. Plant Biol. 11: 228-231.
Ishikawa, C., Hatanaka, T., Misoo, S. & Fukayama, H. 2009. Screening of high kcat Rubisco among
Poaceae for improvement of photosynthetic CO2 assimilation in rice. Plant Prod. Sci. 12: 345-350.
Ishikawa, C., Hatanaka, T., Misoo, S., Miyake, C. & Fukayama, H. 2011. Functional incorporation ofsorghum small subunit increases the catalytic turnover rate of Rubisco in transgenic rice. Plant
Physiol. 156: 1603-1611.
Ku, M.S., Agarie, S., Nomura, M., Fukayama, H., Tsuchida, H., Ono, K., Hirose, S., Toki, S., Miyao,     
BSJ-Review 7:33 (2016)

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