ǎȎȿŃƸéɣ 2:1̊2011̋

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ïʌ ɪȷŁ
ʃ2ǨļŁķŃķŃˠ¯˜ƪóìƐȿŃȸɅȿNz]NQaœƣ
112-8610 ƪ®õķİ 2-1-1
ˬƻ Çĺ
ưƽķŃȗȎʴǿȿŃˍ
252-0880 ȽĻşȱʌǫŦ§«˔ 1866
 ƽȏ˨3̌ưƽǎȎŃ¹Ɉ 75 Ěķ¹2_`O ĶǕ0ȎʶIȗ<à í -©Ǜµʭ2ǚɲ.çŽ- 2ÞōIɢʦ1,ĂD;.>&?2-

2_`O3̌Ƹʿ2ǎȎȿŃȸɅ2ƔǮ̌ȓȐ̌°ź2Ś˛1*,Ȩʒʠʰ EɢʦȪƧɰȪ0_`O.,ưƽǎȎŃ¹Ŭĭľĕ¹1B),¸ȞF;& *,̌ǎȎ2©Ǜµʭ3Ƃ˱2Ǜµʭ.3ȣ0ĎǎȎ2ȗǳ13ȯƝƂʗ-30Ȏʶ
2µʭ-E.ʤʮF,;& ̌ʿū2ȸɅƖʏ2˅Ǡ.Ȱ;),̌©ǛµʭȸɅ3˷ʺȪ0Ś˛Iˆ̌©ǛµʭȘȎ2ɄǑȪ0ŸëǰȮIǵ5,;ȗ¿˞Ž2ƔǤ.

,̌ƲʍɊ2ʥěȎʶ.,̌_XoºˇȎʶ.,Ɋ ̌;1̌˚ǡą2¡-ǎȎÏƚ*B10)&ȗłƑȡ1©Ǜµʭ3ķ0ʳȑI,;Ķŷ.ʟʆB¼ĉ©ǛµʭȘȎ2ǔ˂2ĶǕƆ3̌ʕ˩1˅ó&ȗĉƐɓ2ˑɗ1ơCF̌$FȗȎĶ
ǕƆ7.*0),;ƽ_`O-3̌&©ǛµʭȘȎI*D' þòí1ȉȇI,̌
;;0ʜůC©ǛµʭȸɅ1L|h,EʁƔ¡ĬȸɅɯIɑˉ̌ʬȂI˵;& ƽȏ˨3̌ʬȂɯ2ƸƬ2ȸɅƐǄIȭDʾJ'ÞōI;.>;& ̂íȪ0©Ǜµʭn7ȫǕIʥ;ƽȏ˨I÷á1ǒ<&'̌ȫǕ2°ź2ȸɅ2ɒ.,'

F4ŀƅ;

M.Kato & T.Aoki-1
BSJ-Review 2:1 (2011)ǎȎȿŃƸéɣ 2:2̊2011̋

}+2"Wek"> *:}|s|hfʱş Ļǲʽǐ˸ĔȘǐƖʏɢĉȸɅǚǔ ɽȸɅƓ
305-8519 ʂĨȱ*4Ŧʌƽ 2-1
The mechanism of yellow flower coloration of Camellia chrysantha
Key words: aluminum; Camellia chrysantha; flavonols; yellow flower.
Natsu Tanikawa
National Institute of Floricultural Science, National Agriculture and Food Research Organization
2-1 Fujimoto, Tsukuba, Ibaraki 305-8519, Japan 
 juUśǎȎ3̌¡ĠúˍI¡Ɓ1L`Lǂúˍ1Ŭáŧ̌ȓģɕ 250 ɂ¶ȵCF,
E

2(ăCɽ2ʛʵIȮȪ1æșF,&23̌ưƽ2{juŮCamellia

japonica̋̌UjuŮC. rusticana̋]^SC. sasanquǎ¡Ġ2mOjuUC. reticulata-̌FCI¡Ɓ1juU2ġɾĔɂ3Ȩˇ,&(ɋȚ2006) ǜɎ1,ɂ˜˩ɂĔɂɰɂȭJ1ʎHFEB10Ďƨ÷ū3ưƽ-?¡Ġ@}moCŖØF&Ǖ 0ɂIș&ɂ˜­˩ɰɂȭJ10),E(Ċş&Ċş1990) juUġɾĔɂ2ɽɼ3̌ȩ@L
m_Lpɼɗ1BEǌ̏ʷ2ɌĞ1Ďȅ̇ɼɽɼ2?2Ȉ $2&>ȅ̇ɼɽɼĔɂIÂDà .ķ0ɰɂȮǖ2*10),E ̇ɼĔɂÂà2­ˏʚ.,ƺŹF,
E2̌̇ɼɽɼIƹ EjuUśǎȎ-E ̇ɽjuUśǎȎ3̌¡ĠúˍC}moôˍI¡Ɓ1ɕ 50 ɂáŧ,E.ĭĐF,E(ɋȚ2006) 2( 1965 ū1¡Ġúˍ2ŬʖĳƮɸǬõ-ĭĐF&̄@0ȅ̇ɼɽɼ2USh̊Camellia chrysanthǎ˖ɽʃ̋(Hu 1965)̌̇ɼĔɂɰƐ2&>1Ƹ?æșF,EjuUśǎȎ-E
̇ɽjuUśǎȎ2ɽɼ1˝ EȸɅ3̌£1USh1,˅>CF,& USh
2̇ɼɽɼ2Ȩɼ2²ɛ<1*,3˚˜ʰʫIF,& Ķ2ǎȎ-3̌ȅ̇ɼɽɼ
3SksNnɼɗ1B),Ȩɼ,E UVeO@Sr_2B1̌̇ɼzsNn2R@S[1B),Ȩɼ,EǎȎ?E UShɽű13̌3 ɂ˶
2zs̊!F?YchʥŖ¿̋.SksNnłģ E.ĭĐF,&(Miyajima et al. 1985̌Scogin 1986) UShɽű-3̌ʒȬəɱ2Ƿɱˍá̇ɼȨɼ,E.ʛőFE2-̌ǨȁƆóĉȎ2˝ɮCF̌3 ɂ˶2zsNn2(Ǹ̇ɼɼɗ-EYch 7-X[]NnUSh2£ʗ̇ɼɼɗ-E.F,&(Miyajima et al. 1985) ̌Ych 7-X[]Nn3Ǹ̇ɼɼɗ-E&>̌USh2ȅ̇ɼȨɼIƙɼɗ.,3ȥĘǢF,&
 USh3ū1B),̇ɼ2ȅǸ2ȣ0EɽIē"&Ďª1W-E3!2Ɲ

N. Tanikawa-1
BSJ-Review 2:2 (2011)ǎȎȿŃƸéɣ 2:3̊2011̋
ƻʀÉ¿.1̇ɼ2ȅǸ2ȣ0EɽIē"&D E 2.ČUSh2̇ɼȨɼ13ÀńŇ0Ȩɼʗěłģ E.ɮCF& YchʥŖ¿13LpO.˘¿IŵƐ,ȅ̇ɼIȨɼ EƆʶĎUSh.ċ Thea ¬ś1á˶FEh̊C.
sinensišʃ̋(Chang & Bartholomew 1984) 3̌µʒȪ0LpOʊɄǎȎ-E (Chenery 1955̌
Matsumoto et al. 1976) FC2.1ȳȮ̌UShɽű2̇ɼȨɼ1ELpO
2˝1*,ŉ˾Iʎ)&}+2"Wek}XP =,1.
 UShɽű-?̌SksNn3ȗĉƐF,ĎSksNnIʊɄ EəɱÞŗę
ņ2W|amɏŁʛőF,E (Miyajima et al. 1985) ʊɄ˕Ř0.F,E?
22̌SksNn1˝ EÜ¿Ȫ0Ďƹ˕@ƐáɛƐ1*,2ĭĐ3Ř0 $-US
hI3>. E 7 ɂ2̇ɽjuUśǎȎ1*,̌HPLC̊high performance liquid
chromatography̋áǃ1BDɽű2SksNn2Ďƹ˕@Ɛá1*,ʩ8& (ʱşC 2010)7 ɂ2̇ɽjuUśǎȎ2ɽű1Ď;FEɢSksNnĎƹ˕3̌kNɊ˕- 0.8̏11.3µgg-1 FW -)&̊ʒ 1̋ Ǹ̇ɼɽɼ2m[VUOɽű2SksNnĎƹ˕kN
Ɋ˕- 16 nmolg-1 FW̊9 µgg-1 FW 1ȰŴ̋.ĭĐF,D̊Nakayama et al. 2006̋
̌F.
Ǧʼ,!F?ċɁů¶-)& &),FC2̇ɽjuUśǎȎ-3̌SksN
n3ĶŘ3ɽű2̇ɼȨɼ1ŏ,E.ɮCFE?22̌ȅ̇ɼ2Ȩɼ13Õá0ȅů30.ȺF& SksNnƐá.,̌violaxanthiňluteoxanthiň(9Z)-violaxanthiň
antheraxanthiňβ-cryptoxanthiňβ-carotene ǏàF& FC3!F? ɇ1 β ȖI?*̌β̌β-SkʥŖ¿-E ;&̌!F2ɂ?ċǕ0SksNnƐáɛƐ-Ď̇ɽjuU
śǎȎ-3̌ɽű2SksNnȗĉƐɓ@ʊɄɲí1,ɂ˜ţŗ.Ñčʤ>CF&̊ʒ 1̋
ʒ 1̍̇ɽjuUśǎȎ2ɽű2SksNn
2, Violaxanthin; 3, luteoxanthin 4, (9Z)-violaxanthin; 7, antheraxanthin; 8, β-cryptoxanthin; 9, β-carotene
1, 5, 6 3ƼċŇ2SksNnƐá (ʱşC 2010 )"> *:|}|s|hf ċUShɓɞCǅIƜ˨,ƝƻĲǣ&ǈ-D0Čȅ̇ɼɽIē"EÉ
N. Tanikawa-2
BSJ-Review 2:3 (2011)ǎȎȿŃƸéɣ 2:4̊2011̋¿̊ğ 1a̋.Ǹ̇ɼɽIē"EÉ¿̊ğ 1b̋ȗE FC2ɽűIǦʼ E.-̌USh2ȅ̇ɼȨɼ2²ɛ<2ʝƴIʢ<& Œȋ.,ȩ{juŮğ 1c̋2ɽIș& (a)            (b)            (c)ğ 1̍(a) ȅ̇ɼUSh̌(b) Ǹ̇ɼUSh̌(c) ȩɼ{juU (Tanikawa et al. 2008)}XQN(4#-<¯˜3̌B$ 400-500 nm 2ǯ˚2×Iďā EȎʶİɼ.,ʤʮ E Ķ2Sks
Nn3ɕ 440 nm ´ʿ1 2 * 3 *2ďāǑķI»̌400-500 nm 2ǯ˚˲ĩ1ķ0ďāIƚ*&>̌SksNnIÂEɽ-3̄@0ȅ̇ɼ2ɽɼ10E ƭ̌zsNn3ɻ
1ɕ 360 nm ´ʿ1ďāǑķIƚ(̌400-500 nm 2ǯ˚˲ĩ1E×2ďāŗ&>̌zsNn1BEɽɼ3̌Ȉɼ̌E3Ǹ̇ɼ1./;EUShɽű2ďāa~WmIǽŇ&.ǦSksNn2a~Wm.3ȣ0Ď
420 nm ´ʿ1ďāǑķʤ>CF&̊ğ 2̋ ȩ{juU-3̌400-700 nm 2ǯ˚˲ĩ-ďāǑķ3ʤ>CF!̌ďā?USh.Ǧ8,ŗ.ȺF&

Absorption (%)

ğ 2̍UShB5ȩɼ{juUɽű2ďāa~Wm (Tanikawa et al. 2008)

N. Tanikawa-3
BSJ-Review 2:4 (2011)ǎȎȿŃƸéɣ 2:5̊2011̋"> *:}X 3;51.<809 USh2ȅ̇ɼɽ.Ǹ̇ɼɽ̌B5ȩɼ{juU2ɽű1*,̌pȞ£ʗzsNnƐá.LpO2Ďƹ˕Iʩ8& pH 3 Dȃ&ɽű1*,ǽŇ& zsN
n3̌HPLC 1BD 360 nm 2×ǯ˚-Ǐà E.-áǃIʎ)& LpOĎƹ˕3
ICP-AES áǃ̊ʥŖɝĉ|bȨ×á×áǃ̋1BDǽŇ& zsNn13 pH ́
9/̇ɼó EƆʶE ȅ̇ɼUShɽű.Ǹ̇ɼUShɽű2 pH 3̌!F?
ċ 5.8 -)&̊ʒ 2̋ ȅ̇ɼɽű.Ǹ̇ɼɽű2!FC?̌3 ɂ˶2zs̌ Y
ch 3-hs]NňYch 3-X[]NňYch 7-X[]Nn̊ğ 3̋
ǏàF& $F%F2Ďƹ˕3̌USh2£ʗ̇ɼɼɗ.,ĭĐF,EYch
7-X[]NnIĎ>,̌ȅ̇ɼɽű.Ǹ̇ɼɽű-ķ0ˈ3ʤ>CF0)&̊ʒ 2̋ ƭ̌LpOĎƹ˕3̌ȅ̇ɼɽű.Ǹ̇ɼɽű-ķ0ˈʤ>CF& ȅ̇ɼɽű2
LpOĎƹ˕3̌Ǹ̇ɼɽűBD?ɕ 3 Ế)&̊ʒ 2̋ ȩ{juU2ɽű2 pH
3 4.2 -ĎUSh2ɽűBD¾)& {juU?LpOIʊɄ EǎȎ-
E (ŜȚ 1980)̌ƽȸɅ-Ãș&ɽ-3LpOĎƹ˕ȅ̇ɼUShɽű2 21̉.Ř0)& ;& HPLC WmX1,zsNn.ƞŇFEyW3˭ũ1ŗ̌{juU2ɽű-3zsNnI9.J/ʊɄ0.ȺėF&
ʒ 2̍ɽű2 pȞ£ʗzsNnƐá.LpO2Ďƹ˕ (Tanikawa et al. 2008)ğ 3̍UShɽű1Ď;FE 3 ɂ˶2zs
1̌Ych 3-hs]Nn̎2̌Ych 3-X[]Nn̎3̌Ych 7-X[]Nn
N. Tanikawa-4
BSJ-Review 2:5 (2011)ǎȎȿŃƸéɣ 2:6̊2011̋

3;51.V <809Gp
 UShɽű1Ď;FE£ʗzs2*-EYch 3-hs]NnǏU
Shɽű.ċ pH 5.8 2 0.1 M ː˒ɤʑǷ- 50 µM ȅů2ȁǷ1ʩƩ& 2.Ych 3-hs]NnȁǷ39:Ȉɼ-̊ğ 4a-0̋
̌ɕ 350 nm 1ďāǑķĎ400-500 nm 2ǯ˚˲ĩ139.J/ďāʤ>CF0ďāa~WmIȺ&̊ğ 4b-0̋ F1 0.1 M ː˒ɤʑ
Ƿ̊pH 5.8̋1ȁʝ&ıóLpOǏ0 µM C 50 µM 10EB1 5 µM !*Ǻï,̌ȁǷ2ɼ2Ĵó.$F1»ďāa~Wm2ĴóIʩ8& Ǻï&LpO˕
Ĳï E1*F,̌ďāǑķ 350 nm Cɕ 420 nm 1ɀò̌400-500 nm 2ǯ˚˲ĩ1EďāĲï E..?1̊ğ 4b̋̌ȁǷ2̇ɼȅȨɼ&̊ğ 4a̋ UShɽű2ďāa
~Wm1EďāǑķ?ɕ 420 nm -D̊ğ 2̋
̌Ych 3-hs]Nn.LpO
1B),ŵƐFEďāa~Wm2ȏƀ.˶¼,E.ȺF&(b)

Absorbance

(a)







ğ LpOIǺï&Ychhs]NnȁǷ2ɼ.ďāa~Wm(a) 50 µM Ych 3-hs]Nn1 AlCl3 I 0 µM̊Ţɇ̋C 50 µM̊Ćɇ̋;- 5 µM !*Ǻï&.2ȁǷ2ɼ2Ĵó̊0.1 M ː˒ɤʑǷ̌pH 5.8̋ (b) ĈȁǷ2ďāa~Wm a~Wm2Ȣć3(a)2ȁǷ2Ȣć1ŒƄ (Tanikawa et al. 2008)>B^#=7-$;3?p <809Z
 UShɽűĊɼɼɗIʉȠǨ̊DIW̋-Ƙà̌ˍIˢNR­Ơǘɳ1ˁ,̌ƘàǷ¡2LpOIĎ=ˢNR˶2ˡÿIʢ<& DIW ƘàǷ̌SßȕǷ̌S
ßȕǷ1ıóLpOIǺï&ȁǷ̌$F%FI 0.1 M ː˒ɤʑǷ̊pH 5.8̋1ʩƩ̌ȁǷ2ɼ.ďāa~WmIǦʼ& DIW ƘàǷ̊ğ 5a-1̋IˢNR­Ơǘɳ1ˁ .̌ȁǷ2̇ɼǸ0D̊ğ 5a-2̋
̌420 nm ´ʿ2ďāǼŘ&̊ğ 5b-2̋ F1ıóLpO
IǺï E.̇ɼ2ȅĚž E..?1
̊ğ5a-3̋̌
420 nm ´ʿ2ďā?Ěž&̊ğ5b-3̋
DIW ƘàǷ2ďāa~Wm-̌3)D.& 420 nm 2ďāǑķʤ>CF0)&23̌
WQ˒0/2ƹǚ˒˶1?LpONRIUmEƆʶE&> (Hue et al.

1986)̌ȈÐ0Ȑƍ-2ɽű.ȣ0ĎɽűɛɨI D*6&.-ȗEǕ 0ƹǚȎ1B),LpONRUmF&.ě-30.ɮCF&

N. Tanikawa-5
BSJ-Review 2:6 (2011)ǎȎȿŃƸéɣ 2:7̊2011̋(b)

Absorbance(a)ğ 5̍UShɽű2ƘàǷ2ɼ.ďāa~Wm(a)ȁǷ2ɼ 1̌ɽűƘàǷ̎2̌ˢNR­ƠSßȕǷ̎3̌ˢNR­ƠSßȕǷ1 AlCl3
IǺï&?2̊!F? 0.1 M ː˒ɤʑǷ̌pH 5.8̋ (b)ĈȁǷ2ďāa~Wm a~Wm2
Ȣć3(a)2ȁǷ2Ȣć1ŒƄ (Tanikawa et al. 2008)x ¶2ɝǄČUSh2̇ɼɽɼ3ɽű1ʊɄFEzs2YchʥŖ¿.LpO2ȰªÂș1B),Ȩɼ E.ɝʫ& LpONR3̌zs
2 A Ȗ2 3 ½B5 5 ½2Ǩ˒ī2³̌
B Ȗ2 o-`wnU_ī1ɝĉ E (Markham & Mabry1968) UShɽű2 3 ɂ˶2YchʥŖ¿3̌!F? B Ȗ1 o-`wnU_īIƹ,ĎLpO2Umɝĉ1BḊɼ2ȅɼó1ʳȑE
F;-Lm_Lpɼɗ1BEˬɼɽɼ2Ȩɼ1,̌LpO0/2˖śNRIƂʗ. EÄĭĐF,ĎL`]N̊LpO̋
̌jW]̊Xr_O̋
̌X
VW̊˗̌Xr_O̌S_O̋̌]ML̊Xr_O̋(Takeda 2006)̌Y_̊˗̌Xr_O̋(Yoshida et al. 2006)̌hi|2ɽŭˍá̊˗̋(Shoji et al. 2007) 2ĭĐE ˖śNR1B),̇ɼɽɼIȨɼ EǎȎ1*,3F;-Ä0̌UShã
>,2ĭĐ-E.ɮ,E
ʆ˨ 12 Ť 3101 1 ɘ3 Ȇ ?2% ǶȶǓŦ2 Ú÷2ʐ1 ˃7EŁ@ʧ .ǝ
E

ǶȶǓ .3juU2.-E ŧ2ǆɼ1,juU2Ȇɘǆ>1șCF&.Č
2B0ʒȓȗ;F& juU2Ȇá3LpOıĶ,˗ıŘ0&>̌
ɘǆ>2˥1fpʶȨɼ"!̌ɘɼɪȨɼ E2'. (ʴǿǎȎ¨Ý 1949),<E.̌
USh2ɽ3ɸáɸʻILpO1B),ǆ>,E2B-E
USh3 1979 ū1ưƽ1ŖØF̌ɭ 1980 ūCUShIÃ)&­˩ɰɂ˛Ľ
F& (ɋȚ 2006) ¶ǁ 30 ū2˜1Ķ2USh˩ɂĔɂÂàF,E ̌F
C˩ɂĔɂ2ɽɼ3Wɼ03Ǹ̇ɼ2ɌĞ1),̌̄@0ȅ̇ɼɽɼ2Ĕɂ3
ÂàF,0 (ɋȚ 2006̌Hwang et al. 1992̌ʖƽC 2004) F;-ʎHF&­˩ɰɂ-3̌
N. Tanikawa-6
BSJ-Review 2:7 (2011)ǎȎȿŃƸéɣ 2:8̊2011̋
USh2­ˏȰƔ1ȩɽ2{juUɓĔɂIș&ÄĶB-E (ɋȚ 2006) °ĚʩǇ&ȩ{juU-3̌pH 3UShɽűBD¾̌zsNn1*,39.J/ʊɄʤ>CF0)& ȩɽ{juU.USh2­ˏ-żCF&˩ɂźµ-3̌ɽű1zsNnIʊɄ̌ɢzzsĎƹ˕-3USh.ċɊ;&3$F¶2Ďƹ˕Iƹ Eɓɞ?łģ E.ĭĐF,E (ʖƽC 2004) FC3̌pH @ȗĉƐFEzsNn2ɂ˶̌LpO2ʊɄ˕1,̌ȅ̇ɼɽɼIȨɼ Eǀ·IǾ&,02?F0 ɽű2 pH ́̌LpO.2ɝĉ1BḊɼó EY
chfN|2zsNnIʊɄ E˩ɂźµÂàFF4̌BDȅ̇ɼIȨɼ E?2.ɮCFE ;&̌$2̇ɼȨɼ2²ɛ<ČǸ̇ɼ2˩ɂźµɓɞ-),?-E'̇ɼIȅȨɼ",ʛʵ E13̌LpOĎƹ˕2ĶĪĢ1ǎĚE3ĥǎ-)Dɰ,,LpO2ʊɄIŹ*.)&.?ñǄE2-30.ƞŇFEȅ̇ɼɽɼ3SksNn1BDȨɼ̌Ǹ̇ɼɽɼ3zsNn1BDȨɼ,E.ɻè3̌ƶ1ÖØʛ.0Ďʘ&Ȯ2̇ɼ2ȅ1?.+,SksNnzs
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ɽ?Ķłģ E )&ǎȎ1,̌$F%F2óĉȎ2̇ɼȨɼ1EʳȑůIʡÆ E13̌ɽű$2?22ďāa~Wm@ĀŔa~WmIǽŇ,äå E0/̌Ǝ˓0ʰʫƂʗ.ƌ,E ̇ɼɽɼ2Ȩɼ1˖ś˝ EǎȎ³1?łģ E23̌°ź
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Yp_l
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BSJ-Review 2:9 (2011)ǎȎȿŃƸéɣ 2:10̊2011̋

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651-2241 ȽƒŦʖõŊʱ 1-1-1 KNC uNR]hcfDevelopment of efficient functional analysis of plant terpene biosynthetic genes:
mainly of sesquiterpene biosynthetic genes
Key words: cytochrome P450; pathway engineering; sesquiterpene biosynthesis.
Hisashi Harada
Technology Development Dept., Biotechnology & Drug Discovery Div.,KNC Laboratories Co., Ltd.
KNC Bioresearch Center, 1-1-1 Murodani, Nishi-ku, Kobe, 651-2241, Japan 
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H. Harada - 1
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H. Harada - 2
BSJ-Review 2:11 (2011)ǎȎȿŃƸéɣ 2:12̊2011̋

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H. Harada - 3
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al. 2010̋ C12ɓIș̌UlMXåǈ̋2ƾƿC3®ˎķŃ2ȴŞC.
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 caUk~óĉȎ2ǔ˂ĶǕƆ3̌ĉƐˑɗ2ɂ˶¶ĵ1?$F1ɟˑɗ̌ 0H(caUk~2È˹ĀƄIƙÈ˹ˑɗ2ĶǕƆ1ʸě E.Gķ ǎȎ*EǕ0©ǛµʭȘȎ.ċǕ1̌caUk~2ȗĉƐ1? P450 I±&È˹ĀƄ˓ʗ0ŸëIǄ&,E Ä4̌Lk_p2ȗĉƐ1,̌ADS 1B),LzK`QĉƐF&ź̌P450 -E CYP71AV1 I±& 3 Ǥˤ2Ǩ˒óĀƄIɜ,̌¡˜¿2Lk_p˒̊artemisinic acid̋7.ĴƠFE.ȵCF,E̊Arsenault et al. 2008̋ ;&̌ɡɽ̊Gossypium arboreum̋1Ď;FEzKNmLU_-E\_̊gossypol̋2ȗĉƐ1
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et al. 2001̋ P450 2ȗóŃȪ0ǚɲʝǃ3̌ȮȪ P450 Iķɶʅ̌ˑǥ̌Ʋʍəɱ0/-Ȩȓ̌ɑʔ&ˑɗIș,2 in vitro 1BEʝǃƔǮɻȪ1șCF,E̌ǦʼȪńŇ0ɷɝĉħˑɗ-E.ČĂDƕ˫ĮĉĶ $-ʿū-3̌caUk~ĉƐC P450 1BEĴƠ;-2˄2ĀƄI in vivo -2ƭǮ̌ 0H(ķɶʅ@ˑǥ1BEuN
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̏300 mg ¶2Lk_p˒ĉƐÄĭĐF,E̊Arsenault et al. 2008̋ ;& 8-wn
U_-δ-S`r2Įĉ1,?ċǕ1̌CAŠCYP706B1̌NADPH-P450 RU_ngWf
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BSJ-Review 2:14 (2011)ǎȎȿŃƸéɣ 2:15̊2011̋dIŖØ&ķɶʅ1,̌Ī˺Ƿ 1 L Ŵ&D 100 mg Ɂů2 8-wnU_-δ-S`rżCF,
E̊Chang et al. 2007̋ ƸʿɉɯC̊2011̋3̌éˀ&caUk~ĉƐˑɗˊºŁ2ǚɲʝǃɓIƢĴ E.
1BĎñȒȪ1 P450 2ǚɲʝǃĄɲ0_akI˛Ȩ& 2_ak-3̌ u˒ʴóˑɗˊºŁɫ2Ǵ1 P450 ˋÔɓˑɗˊºŁI˄ɝ&Ȩȓ|anIÂʔ̌F
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H. Harada - 6
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É 2µʭ˝˄áŁ'-30$FC2rimWIɮƏ&ȸɅʢ<CF,,EFCĶˮȪ0L|hC2ȸɅ1B),̌°źǕ 0k~óĉȎ2ȗĉƐɜʹƴC
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Yp_l
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H. Harada - 8
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Diversity of enzymes in isoflavonoid pathway
Key words: isoflavonoid; P450; phytoalexin; prenyltransferase; soybean.
Tomoyoshi Akashi
Department of Applied Biological Sciences, Nihon University
Fujisawa, Kanagawa 252–0880, Japan ƹǩǸ1Ŗâő EzsNn3˓Œ1īƾ0ƽʦIƂ*˫FC3!F?zPp
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E ǄɓǾǰaWpXǔ Iƣʟǵ,E (Akashi et al. 2003 and 2005a) 2źǔ3ƹǩcDNAIĬɖɣ@ʵǌ0/2ǺȚȳɒȮ-Ǿǰ"˫ȄȀ2ʵȰǖůIƂ*WIǳôĸȀLicN-þŨ˫ùĖʱ¶ĵIĆļ E?2-Ɠɸ2ʱ¶ĵIþŨ Eˍ1Ǫ1Ƥñ-
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BSJ-Review 2:20 (2011)ƹǩȖĸƣêɁ 2:21˩2011˪
-2NezǳǴʹ3Ŭ!?ˤ0 Iris-IFS 3Ȗ IFS .Ǎʛ E. in vitro -ȯ 20ÊˤǖůIȑ& ˤNezsNnǳǴɓIȑ

Iris-IFS 3˫ƹǩ2fiWQ

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 Glyceollin 3˫|kSvˣƷ1 C5 Ne|øº¯ð˫ǲǫQkIŠŵ&g
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8

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Soybean phytoalexinė 5 gNb2XcR2ǳąŵȷʗ ȢɎC3|kSv72|pĠʚȘüŭm[zPąŵȮ2 homogentisate
phytyltransferase (HPT).üŭ˚¹,E.C˫HPT LsʶʴãI?.1gNb EST l

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BSJ-Review 2:21 (2011)ƹǩȖĸƣêɁ 2:22
˩2011˪
f~aIƺȱ˫50˨±2ĆůIȑ

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˩G4DT˪
-E.H)&(Akashi et al. 2009) $2ŦgNb2Zs ʴãƚC1F&.Clf~aIÞŘƺȱ˫Ɠ&1 3 *2X(PT4, PT5, PT6)Iɷ'& $F%FIʵ
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ĵ(SfN8DT-1, SfN8DT-2)ĆļF,E(Sasaki et al. 2008) ȮȻǁ-3 PT2, PT3, PT4, PT5,
PT6 3 SfN8DT .3æ2Wn1ŉ,E gNb-3ʨô2ʬș-NezsNnIĠ
ʓ1 E|pĠʚȘʵȰǳ˫īƾ0ǄɓIƂ*B10)&.ŲļFE LWc|f2ƽʦIǒ>EfvWʓƽʦ1ɜċƂ&F˫«ŦĄʴã2Ǎʛ@ĩǺńØ1BDƚC1&ė 6 ƹǩ2|pĠʚȘʵȰ2âĵȮȻǁ ȮȻǁ3 ClastalW Iǵʟˎȸąǔ-¾ŵ&kp{˒ƦÆĮ 2009˭Ȗlƹǩ2Zs űġ.uNReaIçǵ&ĭǣǩǳąŵʱ¶ĵ2Ćļ˭zK_L˫45: 881-886.
ȾʲȉƚȌƠɊ 2006. zsNn˫NezsNn˭ŊƫˀʍŌ˩ɂ˪ĭǣǩôĸ
˩ƹǩɂ˪˭pp. 387-398.LNy_áǨʲ˭Ʈ©.
Akashi, T., Sawada, Y., Shimada, N., Sakurai, N., Aoki, T. & Ayabe, S. 2003. cDNA cloning andbiochemical

characterizationofS-adenosyl-L-methionine:

2,7,4'-trihydroxyisoflavanone

4'-O-methyltransferase, a critical enzyme of the legume isoflavonoid phytoalexin pathway. Plant Cell
Physiol. 44: 103-112.
Akashi, T., Aoki, T. & Ayabe, S. 2005a. Molecular and biochemical characterization of
2-hydroxyisoflavanone dehydratase. Involvement of carboxylesterase-like proteins in leguminous
isoflavone biosynthesis. Plant Physiol. 137: 882-891.
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Akashi, T., Ishizaki, M., Aoki, T. & Ayabe, S. 2005b. Isoflavonoid production by adventitious root culturesof Irs germanica (Iridaceae). Plant Biotechnol. 22: 207-215.
Akashi, T., Sasaki, K., Aoki, T., Ayabe, S. & Yazaki, K. 2009. Molecular cloning and characterization of acDNA for pterocarpan 4-dimethylallyltransferase catalyzing the key prenylation step in the biosynthesisof glyceollin, a soybean phytoalexin, Plant Physiol. 149: 683-693.
Ayabe, S. & Akashi, T. 2006. Cytochrome P450s in flavonoid metabolism. Phytochem. Rev. 5(2-3):
271-282.
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and Hung-Wen (Ben) Liu (eds) Comprehensive Natural Products II: Chemistry and Biology, Volume 1:
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929-976, Elsevier.
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Hashim, M.F., Hakamatsuka, T., Ebizuka, Y. & Sankawa, U. 1990. Reaction mechanism of oxidative
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Mackova, Z., Koblovska, R. & Lapcik, O. 2006. Distribution of isoflavonoids in non-leguminous taxa – anupdate. Phytochemistry 67: 849– 855.
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non-leguminous families: an update. Nat. Prod. Rep. 22: 504–515.
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xb+J7"J/&2JS]m¼ Ʀ ¸ĬƮ©ʝōĬĸĬĸˊōĸȍȝˊǳčǄɓȖĸʲˁ
184-8588 Ʈ©ʳŅʺ¨ŐǷ˱ˬ˱˳ˬ˰˴
Anthocyanin modification catalyzed by vacuolar proteins.
Key words: anthocyanin; carnation; delphinium; glucosyltransferase.
Nobuhiro Sasaki
Division of Biotechnology and Life Science, Institute of Engineering, Tokyo University ofAgriculture and Technology
2-24-16 Naka-cho, Koganei, Tokyo 184-8588, Japan
ƹǩ3ɠ˫ɥ˫Ʊȁȣ˫ƾ 0ĕĻȶɅ-ƹǩɟȰIąŵ,E -?˫ƣ?īƾ0ɟšIƂ*ɟȰX|3Lm_Lp-E °ɯȀ0ƹǩ2¦ǆ°ʌǴǩ-ELm_L
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F9/īȚīƾ0ɟšIƂ*23˫FC2ȭ@ƤǄʶ1BEɵː0È˝Ė-E
˩Yoshida et al. 2009˪ F;-2ȍȝ-Lm_Lpâĵ2ǳąŵȷʗƚC.F,&
Ǫ1LX[2ąŵ1*,3 -19:Ù,2üŭǊˌ1˅HEʵȰ@ʱ¶ĵøˑF˫
$FC2üŭǄƽ1*,?ɼƯ0F,E˩Tanaka et al. 2008˪ Lm_Lp2ȭ@˫ƤǄʶ1BEÈ˝üŭ1*,?ī2ȍȝġĊ0F,E Lm_Lpâĵ72ȭʚȘüŭ3 UDP-sugar ÃķȀ1ȭʚȘIɽĴ EʵȰ-E UDP-sugar glucosyltransefase˩UGT˪
1B),ɽĴFE.˫;&˫ƤǄʶʚȘüŭ3L_-CoA Ãķĝ2L_ĠʚȘʵȰ1B),ɽĴFE.ȋCF,E˩Yonekura-Sakakibara et al. 2009˪ FC2ʵȰ3ȳɒʓŇěĝ2ʵȰ.,ȋCF,D˫ɝ13Lm_Lp3$2LX[ąŵF,C˫
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CF,E˩Ozeki et al. 2011˪ ˫ƣʟ10),˫Lm_Lp72ƤǄʶʚȘüŭ@ȭʚȘüŭ2ʲǙɒȘɭĝ2ʵȰ1B),ɽĴF,E.˫ʵȰ~˫ʱ¶ĵ~-ƚC.F**E $-˫Ʃț-3ʟŕşȍȝľ-ġĊ,&ǙɒȘɭĝfvWʓ
1BELm_LpÈ˝ʵȰüŭ1*,ȴ¬ E

N. Sasaki-1
BSJ-Review 2:24 (2011)ƹǩȖĸƣêɁ 2:25
˩2011˪

K4K&DJJ/&2J-FJ%]mʔɟ2Sr_3 pelargonidin 3-glucoside˩Pg3G˪2X[aâĵC1\ʶ1B),È˝F& pelargonidin 3-malylglucoside˩Pg3mG˪I ɟȰ.,Ƃ*.ȋCF,
E Lm_Lpâĵ72ɔɏƕƤǄʶ2ʚȘüŭ3L_-CoA Ãķĝ2L_ĠʚȘʵȰ1B),ɽĴFE.ġĊF,E˩D'Auria 2006˪˫\ʶ2ʚȘüŭ1*,3ġĊǢ)& \ʶ-CoA 3đƻȀ1ʐĨF,0&>˫ɦɎC2X|-3Sr_ɠśC»âĵôąǩIžá˫$2žáǙIL_ĠÂ½.,˫Sr_ɠśCžá&ȫʵȰǙIǵ, Pg3G IL_Ġÿŀ½.,üŭ"&.G˫Pg3mG
2ǳŵȐʆF& Ȭɴ&L_Ġÿŀ½2ʓʹâƯIɭ0)&.G˫\ʶ.X[aɕǐɄą&?2-E.¤ŲF& $-˫\ʶX[aIôĸȀ1ąŵ˫$FL_ĠÂ½.,Ò1*,ƺɿ&.G˫žáǩ.Ćƾ1L_ĠÂ½.,Ò.ȑF& 2.C˫Sr_1,Lm_Lp72\ʶʚȘüŭ3˫L_X[aÃķĝ2L_ĠʚȘʵȰ˩AAT˪1B),ɽĴFE.
ȑF&˩Abe et al. 2008˪ $F;-˫Lm_Lp72L_ĠʚȘ1*,ɡ˟ƕƤǄʶ1*,3 AAT ĝ2?2ġĊF,&˫ɔɏƕƤǄʶ2ʚȘ1*,3ä>,2ġĊ-)& AAT ĝ2L_ĠʚȘʵȰ3ǙɒȘɭĝ-E.ȑF,E˩Hause et al. 2002˪
.C˫Sr_1ELm_Lp\ʶʚȘʵȰ˩AMalT˪?ǙɒȘɭĝE.¤ŲF&

Figure 1 Sr_ɠś1EƆļFELm_Lpąŵȷʗ Lm_Lp
2Á.,\p`fN|2âĵIȑ&K4K&DJJ/&2J 5 P R[]mǟƸɟ2Sr_Ƃ* 0Lm_Lpâĵ3 pelargonidin 2 3 º. 5 ºX[a 1 âĵ!*-È˝F˫C1$FC2X[a 1 âĵ2\ʶ1B),ƲǂF&
cyclicmalylpelargonidin IƂ*.ȋCF,E˩Nakayama et al. 2000˪ Lm_Lp2 5 º72ȭʚȘʵȰ˩anthocyanin 5-glucosyltransferase, A5GT˪1*,3®2ƹǩȚ-3ġĊE
N. Sasaki-2
BSJ-Review 2:25 (2011)ƹǩȖĸƣêɁ 2:26
˩2011˪
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2010˪ Ľˍ12fvWʓǙɒ1Șɭ E1*,ƺɿ E&>1˫AA5GT _Xo|hnI GFP fvWʓ2 N ƨȡ1ɬą EB1ƽȨ&[amWmI˫frVɯȁȳɒ7vkMWunmǔ1B),ʬȀ1ńØ& $2ȸƱ˫GFP 2ɫ×ǙɒÝ1ɻłF&.C˫AA5GT fvWʓǙɒȘɭĝ-EāɓůȑďF&
˩Matsuba et al. 2010˪

.G82>J/&2J 7 P R[]mlzMpO 3$2ǬǪ0˒ɠªǏ2ɠ-E lzMpO ɠś2 ɟȰ3_L
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BSJ-Review 2:26 (2011)ƹǩȖĸƣêɁ 2:27
˩2011˪
ʵȰüŭIɭ0)& üŭǴǩI HPLC Iǵ,ɼƯ&.G˫ȭʚȘ1B),ǳ&.ŮH
FEyWɻǜF& 2ǳŵǩIȬɴ˫ʓʹâƯ0C51 NMR â×ǔ1B),Ȑʆ&ȸƱ˫cyanidinn 3,7-diglucoside˩Cy3,7dG˪-E.åƚ& 2.ClzMpO 1
,Lm_Lp2 7 ºʴȭ½ôüŭ3˫acyl-glucose dependent anthocyanin 7-glucosyltransferase
˩AA7GT˪1B),ɽĴF,E?2.ɍCF& $-˫Sr_C AA5GT Iøˑ E21Àǵ&?2.Ć degenerate primer Iǵ,˫lzMpO ɠśCʊɴ&cDNA Iʻĝ., PCR Iɭ˫AA7GT Ëɲ cDNA ƒǧIǮŨ& C1 5’-, 3’-RACE ǔI
ɭ˫$2Ùˀ cDNA IǮŨ& ǮŨ& AA7GT 2ƆļLsʶʴã13ǙɒȘɭ_Xo.
¤ŲFEʴãķě,&&>˫2˖ĞǅƉ&fvWʓĬɖɣ-ąŵFEB1ƽȨ&|anIĬɖɣ7ńØ& 2ȶƇĬɖɣ-ǳǴ"&fvWʓ˫AA7GTǖůIƂ),&.C˫2 cDNA  AA7GT I[n,E?2.ȸʋ¯&˩Matsuba et
al. 2010˪

¡.G82> J/&2JS]mlzMpO ɠś1Ĉ;FE_LslzM3êʠ&B1Lm_Lp2 7 º2X
[aC1ɵƍ2X[a. p-hydoroxybenzoic acid˩pHBA˪1B),È˝F,E
Lm_Lp2 7 º72X[a2ʚȘüŭǙɒȘɭĝ2ʵȰ1B),ɽĴFE.
C˫$2Ŧ2È˝üŭ?ǙɒȘɭĝ2ʵȰ1B),ɽĴFE.ɍE2Ĳş-E.ŮHF
E F;-1?ɡ˟ƕƤǄʶ2ʚȘüŭǙɒȘɭĝ2ʵȰ1B),L_X[aÃķȀ1ɽĴFE.ġĊF,&˩Gläßgen & Seitz 1992˪.C˫lzMpO 1,?Lm_Lp72 pHBA 2ʚȘüŭL_X[aÃķȀ0ʵȰ1B),ɽĴFE
1*,ƺɿ& Ö2 AA7GT üŭ1B),ǮŨ& Cy3,7dG IL_Ġÿŀ½˫pHBG IL_ĠÂ½.,˫lzMpO ɠśCʊɴ&ȫʵȰǙIǵ,ʵȰüŭIɭ0)&üŭǳŵǩI HPLC Iǵ,Ȑʆ&.G˫Ɠ&0yWȐʆF& 2üŭǳŵǩIȬ
ɴ˫ʓʹâƯ0C51 NMR â×ǔ1B),Ȑʆ&.G˫Cy3,7dG 2 7 º2X[a2 6º1 pHBA Qakȸą&ǩʓ-E.åƚ& C1üŭƜ˄IƍƜ˄CƝ;-Śˀ&.G˫HPLC WmX 1Ɠ&0yW 2 ƩɻǜF& FC2âĵʹIɼƯ&ȸƱ˫Cy3,7dG 1 pHBA . glucose  1 âĵ!*ɕǐɄą&?2.˫2 âĵ2 pHBA
. 1 âĵ2 glucose ɕǐɄą&âĵʹ.ɛ& FC2ȸƱC˫lzMpO 2˒
ɟɠś-2_LslzM2ǳąŵȷʗ3˫delphidin 3-rutinoside ąŵF&Ŧ1˫ǙɒÝ1
, 7 ºʴȭ½ôF˫C1$F1ȼ,˫pHB . glucose  1 âĵ!*ʚȘF,.-ąŵFE?2.ƆļF&˩Fig. 2˪¢
F;-˫Lm_Lpâĵ3ĠƩȀ13ȳɒʓ-ĺŵF&Ŧ1˫Ǚɒ7ʜʡFɧȜ
FE.ɍCF,& ˫lzMpO 1E_LslzM2B0âĵ?˫$2Ŧ÷3ȭ.ƤǄʶ1BEÈ˝üŭ-D˫$FC2ī2üŭǙɒÝ-ɭ0HFEāɓůˤ.ŮHFE ?.?.3L_X[a?¦ǆ°ʌǴǩ2Ț-)&.ŮHFE˫$N. Sasaki-4
BSJ-Review 2:27 (2011)ƹǩȖĸƣêɁ 2:28
˩2011˪
2ɧȜĢŸ-EǙɒÝ-˫$FCIçǵ,Ɠ&0âĵIąŵ E°ʌȷʗIǮŨ&.3
¦ǆ°ʌǴǩ2Ț˚Iĥ@ -Ƥñ0ʨô2źǊ-)&.ɍCFE Ǫ1˫ʠ2l
zMpO 2Á2B1˫1 Ț˚2L_X[aIȭ.L_Ġ2Ɣ2Â½.,çǵ
E.ŷǹ3˓Œ1ɜċƂ&FE «Ŧ˫®2ƹǩȚ1,?Ćƾ2°ʌȷʗIȍȝE.-ɵː0¦ǆ°ʌǴǩ2ąŵȷʗ1*,2ơ0EźDŨCFE?2.ƥŤFE
Ʃț-3ʟŕġĊF&ǙɒȘɭĝ2Lm_LpÈ˝ʵȰ1*,ʠ8,& ˫F;-2ġĊ-3$FC2fvWʓ˫ǙɒȘɭ_XoIƂ),E.˫;&˫ƣȵȀ
1ǙɒÝ1ʜʡF,E.ȑďF,E ˫ǙɒȘɭĝ2ʵȰ1B),ɽĴF
ELm_Lp2È˝üŭĽˍ1ǙɒÝ-ɭ0HF,E2˫E3 pre-vacuole @Ǚɒ72ʜʡʣ2ŅĕĻÝ-ɭ0HF,E21*,3˫C0EƺɿŬɶ-E FC
2Ē˗3˫«Ŧ˫Ƨ'1ǒȊI<,0ȳɒÝ2Lm_Lp2ʜʡȷʗ2Ē˗.¿",ɼƚF,?2.ƥŤFE

Figure 2 lzMpO ˒ɠś1,ƆļFELm_LpɟȰąŵȷʗėkp{Abe, Y., Tera, M., Sasaki, N., Okamura, M., Umemoto, N., Momose, M., Kawahara, N., Kamakura, H.,
Goda, Y., Nagasawa, K. & Ozeki, Y. 2008. Detection of 1-O-malylglucose: pelargonidin
N. Sasaki-5
BSJ-Review 2:28 (2011)ƹǩȖĸƣêɁ 2:29
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3-O-glucose-6''-O-malyltransferase activity in carnation (Dianthus caryophyllus). Biochem. Biophys. Res.
Commun. 373: 473-477.
D'Auria, J.C. 2006. Acyltransferases in plants: a good time to be BAHD. Curr. Opin. Plant Biol. 9:331-340.
Gläßgen, W.E. & Seitz, H.U. 1992. Acylation of anthocyanins with hydroxycinnamic acids via
1-O-acylglucosides by protein preparations from cell cultures of Daucus carota L.

Planta 186:582-585.
Hashimoto, F., Tanaka, M., Maeda, H., Fukuda, S., Shimizu, K. & Sakata, Y. 2002. Changes in flower
coloration and sepal anthocyanins of Cyanic delphinium cultivars during flowering. Biosci. Biotechnol.
Biochem. 66: 1652-1659.
Hause, B., Meyer, K., Viitanen, P.V., Chapple, C. & Strack, D. 2002. Immunolocalization of

1-O-sinapoylglucose:malate sinapoyltransferase in Arabidopsis thaliana. Planta 215: 26-32.
Matsuba, Y., Sasaki, N., Tera, M., Okamura, M., Abe, Y., Okamoto, E., Nakamura, H., Funabashi, H.,
Takatsu, M., Saito, M., Matsuoka, H., Nagasawa, K. & Ozeki, Y. 2010. A novel glucosylation reaction

on anthocyanins catalyzed by acyl-glucose-dependent glucosyltransferase in the petals of
carnation and delphinium. Plant Cell 22: 3374-3389.
Nakayama, M., Koshioka, M., Yoshida, H., Kan, Y., Fukui, Y., Koike, A. & Yamaguchi, M. 2000. Cyclic
malyl anthocyanins in Dianthus caryophyllus. Phytochemistry 55: 937-939.
Ozeki, Y., Matsuba, Y., Abe, Y., Umemoto, N. & Sasaki, N. 2011. Pigment Biosynthesis I. Anthocyanins.
In: Ashihara, H., Crozier, A. & Komamine, A. (eds.) Plant Metabolism and Biotechnology. pp. 321-342.
John Wiley & Sons, Ltd, Chichester.
Tanaka, Y., Sasaki, N. & Ohmiya, A. 2008. Biosynthesis of plant pigments: anthocyanins, betalains andcarotenoids. Plant J. 54: 733-749.
Yonekura-Sakakibara, K., Nakayama, T., Yamazaki, M. & Saito, K. 2009. Modification and Stabilizationof Anthocyanins. In: Kevin, G., Kevin, D. & Chris, W. (eds.) Anthocyanins. pp169-190. Springar
Science+Business, New York.
Yoshida, K., Mori, M. & Kondo, T. 2009. Blue flower color development by anthocyanins: from chemical
structure to cell physiology. Nat. Prod. Rep. 26: 884-915.

N. Sasaki-6
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2$-J}^o<(+KOg£Q2$-JdR9IK-
ŗă ɌǂƩ ˋİɞÖȡȖĸŻɮĬĸˊĬĸ uNR]NQaȍȝȖİɞȈǳˡŐˤŊǷ 
Master genes for nicotine biosynthesis in tobacco; approach using low-nicotine mutants
Key words: alkaloids; jasmonates; nicotine; tobacco; transcription factors
Tsubasa Shoji & Takashi Hashimoto
Graduate School of Biological Sciences, Nara Institute of Science and Technology
Takayama 8916-5, Ikoma, Nara 630-0101, Japan
ƹǩɧȜ EīȚīƾ2¦ǆ°ʌǴǩ3ĀƭBDöɨ, ĔıĎ, ƳƑ0/.,ª˚2ÎÙ0Ȓµǳǖ
1ʏǭ,& ¦ǆ°ʌ1˅ Eȍȝ3, ŧƭ3ĭǣǩôĸ1BEôąǩ2øˑĆļ.ǳąŵȷʗ2Ɔ
ļ, ú5ǳôĸ1BEǳąŵʵȰ2Ǫū¯ Ǘ)& ʟŕ, âĵǳǩĸ2ǾňIɑƟ1, ǳąŵʵȰʱ¶ĵ2WpXȂJ1ɭ0HF, $FCBDƞʫȀ0ǆ°ʌʵȰʱ¶ĵCʨô,&.
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0¦ǆ°ʌǴǩ3, ɰĵƹǩ$F%F2Ȗ@ŉ1âŋ&ʨô2ŦƥǊˌ-ǦǾȀ1ǳ&?
2-D, zsNnǳąŵ.3Ǻ0EǬə2èũǄƽI?*?2.ɍCFE
LSNn3ĈȞȰů-44ģĠůIȑ

¦ǆ°ʌǴǩ2ȿȗ-E

˙ɠƹǩ2ȯ

20˨2Ț1ɷáF, $2ôĸƽʦ3â),E?2'-? 12,000 Ț˚1ú6 (Roberts &Wink 1998) ī2LSNn3, Lsʶ@qWRhn˚CąŵF, $2êˠ½@ǳąŵȷʗ1B),õâFE.ī(Ziegler & Facchini 2008) ksNn@zPp|vsNn2Ģą.3Ǻ0D, Ą 2LSNnX|3Ǭȟ1$2ǳąŵȷʗIȐȟ,&.
F,Efu[ŉƹǩǪƤ2LSNn-Ep[hú5$2˚Ƀôąǩ3, $2ŞǋɪůCƹǩ1.),3ˇũǩʓ.,2ųċI?),E(Baldwin et al. 1998) fu[2ɥĿɪ1BD
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BSJ-Review 2:30 (2011)ƹǩȖĸƣêɁ 2:31˩2011˪ąŵʱ¶ĵɉIƶ2Öȡʲâ-ǖůô E ƶȡąŵF&p[h2Ĭʲâ3ńȥI¶H),Ĝ
ʲ7ʚǗF, Ùɢ1ɧȜE(Shoji & Hashimoto

2011a)˩ė˰˪ p[h3Rph.LavVʶIêˠ½.

Eüŭȷʗ1B),ąŵFE

˩ė˱˪Rph3Ńȗ`L-E|mi_IȷǶ,y`ǲI?* N-hy
pO ShR1, LavVʶ3ɲʵȰ-
E NAD .Ûʤ2ȷʗ(Katoh & Hashimoto 2004)1B),y`ǲI?*p[hʶ1$F%FĩƇFE ȷʗ2ȵȃ3ǲ2ɄąIĈ=ɵƍ2üŭCƽŵFE.F,E$2ʅȳ3Ƨɼƚ-
E(De Boer et al. 2009, Kajikawa et al. 2009, Kajikawa et
al. 2011)

2$-JZlOgj"E(+K[ ERF bWagOg
1930 ŕ°1UuùǴ2fu[ȮȻLS
NnI9.J/Ĉ;0.ǾɷF& 2»p[hȮȻ3ȩęʝóȇ-ʱ¶ĸȀɼƯ0F, ˱*2ʱ¶ĵř NICOTINE1 (NIC).
NICOTINE2 (NIC2)1E÷ÔůĩǺùĖ-E.åƚ&(Legg et al. 1971) 2ĩǺ½
1,9:Ù,2p[hǳąŵʱ¶ĵǾǰżèF,E.(Hibi et al. 1994, Katoh et al.
2007, Shoji et al. 2009, Shoji et al. 2010, Kajikawa et al. 2011)C, NIC1 . NIC2 3èũʱ¶ĵ-E
.ƆļF& EST űġ2ɧȜIɑƟ.,, fu[ cDNA LNIǵ&NWLNɼƯʟŕāɓ.0)& Ŷ 3, ʸǳĝ. nic1nic2 ¦ʷĩǺ½2ƶ2Eʱ¶ĵǾǰIfu
[NWLN-ȽɇȀ1ƺȱ, Ɩȋ2p[hąŵ, ʜʡ1˅HEÙ,2ʱ¶ĵĩǺ½-ǾǰżèF,E.ȐʆFE..?1, APETALA2/ETHYLENE RESPONSE FACTOR(AP2/ERF)zK(Nakano et al. 2006)2X| IXa 1ŉ EʚßĖĵʱ¶ĵ ERF189 2ǾǰĩǺ½-ɷCF0.IƚC.&(ė 3, Shoji et al. 2010) IXa X|2 ERF ʱ¶ĵ3fu[Zs 1ņ0.? 25 ʱ¶ĵķě, $FC3Lxn|_a2 AtERF1,AtERF2 1ʟWn 1 . ERF189 IĈ=Wn 2 1âæFE ERF189 IĈ>Wn 2 2ɵƍ2ʱ¶ĵ2
Ǿǰ3 nic2 ĩǺǪǺȀ1˙ɦ1ǾǰżèF,& Zs ]^. PCR 2ȸƱ, 2ǪǺȀ0
Ǿǰżè3Wn 2 1ŉ E ERF 2(, ɵ¦Ê½fu[(Nicotiana tabacum)2ȓÖȚ2˰*E N. tomentosiformis 1Ƕƭ Eʱ¶ĵ, nic2 ĩǺ½Zs -ǅį,E.1ʕĖ,
D, FCǅįʱ¶ĵ˩ņ0.?˵ʱ¶ĵ˪3ĩǺ2ùĖʱ¶ĵř NIC2 1Şʧʾ,E.â)& Ɣ, Wn 1 @Wn 2 1ŉ,?æ2ȓÖȚ-E N. sylvestris Ƕƭ2 ERF
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T.Shoji & T. Hashimoto-2
BSJ-Review 2:31 (2011)ƹǩȖĸƣêɁ 2:32
˩2011˪

ė 4 1ȑ B1, Wn 2 2
ERF 3, N. tomentosiformis 1Ƕƭ E
NIC2 ʱ¶ĵř.$F.ȆĆ0 N.
sylvestris 1Ƕƭ Eʱ¶ĵř$F%F
1Wafǫ1ķě,D, nic2
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E˩Ù,?3Ĭ÷˪.ɍCF&

ERF Wag2$-J}^oOgZlfu[ŠʓʚƇǎǫƶIǵ&ɼƯ-,Wn 2 2 ERF ʱ¶ĵp[hĈʹIŎĂ EafèũĖĵ-E.ȑF&(Shoji et al. 2010) ERF189 IʬìǾ
ǰ E.p[hĈʹʸǳĝfu[-ȯ
2-3 Ê1, nic1nic2 ĩǺ½-ʸǳĝ<10)& ʢ1, Wn 2 2 ERF IǪǺȀ1
RNA œǛǔ-Ǿǰżè&D, nomżèĝ2 ERF189 @ ERF179 IǾǰ"&D E.p[hąŵżCF& 2ȸƱ3, N. benthamianan 2ȆĆʱ¶ĵ
NbERF1 2Ǿǰżè-ŨCFEȸƱ.?ɛ&(ė 3, Todd et al. 2010)
ERF ʚßĖĵ3p[hǳąŵʱ¶ĵ|fÝ2Ǫļ_aʴã(ė 5)1ȸą, ʚßǖů
ô1Ò˩Shoji et al. 2010, De Boer et al. 2011, Shoji & Hashimoto 2011b, 2011c˪ ERF189 .X[[h[Nnÿŀ½2ɬąfvWʓIfu[ǎǫƶ-Ǿǰ",, lU]fnÃķȀ1
ERF189 Iǖůô"E.,8,2p[hǳąŵʱ¶ĵʚßʇńFE(Shoji et al. 2010) ;T.Shoji & T. Hashimoto-3
BSJ-Review 2:32 (2011)ƹǩȖĸƣêɁ 2:33
˩2011˪
&, &ʇńů2Ĭ÷3_W}U_nķě-?ʕE.C, ®2Ėĵ2ɋʂI¬
!1 ERF189 3ǳąŵʱ¶ĵIǖůô E.â)& ǳąŵʱ¶ĵ2˰*-E|m
i_ N-hĠʚȘʵȰ(PMT)2Ǿǰ13ʚß˃ĳǡ2ʟÐ1E GCC box ʷɶ-E.
ȋCF,&(Xu & Timko 2004) Wn 2 2 ERF 3, 2 PMT |fÝ2 GCC box
7 in vitro -ȸą E..?1, ʬȀǾǰȮIǵ&maLWkM~_ɼƯ-, F
C ERF 3, 2 GCC box I¬, PMT |fIǖůô E.ȑF&(Shoji et al. 2010,
De Boer et al. 2011) Ɣ, Wn 1 2 ERF 3 PMT |f2 GCC ʴã13ȸą"!, |fIǖůô E.30)&(Shoji et al. 2010)

ERF WagH!CIJLwNOgF"GK/¦ǆ°ʌǴǩ2ŖɊ-2ąŵȮ3, êˠ½@°ʌ˄½IÂȺ Eǆ°ʌȷʗ.$F1ȼ
¦ǆ°ʌ1ǪǺȀ0ʲâCƽŵFE p[hąŵ1ǪǺȀ0 PMT 0/.3Ǻ0D, y`ǲąŵ1˅HEUsʶa_ĠʚȘʵȰ(QPT)3, p[hąŵ1Ŭɶ0p[hʶ2ÂȺ.ɭ,, ɲʵȰ.,ʷɶ0 NAD ąŵ1?˅HEǆ°ʌʵȰ-E(Katoh &Hashimoto 2004)
fu[13 QPT I[n E˱*2ʱ¶ĵ QPT1 . QPT2˩!F? N. tomentosiformis 1Ƕƭ
E˪ķě E(Sinclair et al. 2000) QPT1 3 8,2ĕĻ-`aʶàǱ2ƤǢ1H
C!ƽŵȀ1Ǿǰ

E21Ń,,

QPT2 3®2p[hǳąŵʱ¶ĵ.
Ćƾ1`aʶŭȤů, ƶǪǺůIȑ, * NIC ʱ¶ĵ2èũIÿE Ľˍ, ERF189 2ʬìǾǰ@Ǆɓżèĝ ERF189 2Ǿǰ3 QPT2 2Ǿǰ1
2<Ţ˔I&(Shoji & Hashimoto
2011c) QPT2 ʱ¶ĵ2|f˖Ğ13˲*2ǄɓȀ0 ERF189 ȸąʴãķě, !F? ERF189 1BE
QPT2 |f2ǖůô1ŁE., in vitro ȸąɼƯ, ʬȀǾǰȮIǵ&maLWkM~_ɼƯ, ú5, fu[ŠʓʚƇǎǫƶIǵ&|fɼƯCâ)&Ɣ, QPT1 |f3

ERF189 1ȸą"!, ;&$2èũIÿ0)&mm, Lxn|_a0/2Zs
13 QPT ʱ¶ĵ3˰[yķě,0 fu[1,, ʱ¶
T.Shoji & T. Hashimoto-4
BSJ-Review 2:33 (2011)ƹǩȖĸƣêɁ 2:34
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ĵʷɵ1B),ǳ&§1ȆĆ0˱*2 QPT ʱ¶ĵ2(, ˰*2ʱ¶ĵ|f˖Ğ
1ɵƍ2 ERF189 ȸąů_aʴãIǮŨ E.-, ERF èũ Ep[hǳąŵV
1þDʞ;F, p[hąŵ2ǮŨ-Ŭɶ.0)&y`ǲÂȺʹ2ĥð1Ńŭ&.ɍC
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Blechert et a1995, Yukimune et al. 1996) ˤȣƹǩ3ƞʫ
Ȁ0[L.0E`aʶ_XoȷʗIƂ),
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et al. 2010) 2ɵą½3WhƽʦĩƇIÅ,ƿȀʱ¶ĵ2ǾǰIȜƼȀ1żè E.ɍCF,
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āɓ.0E
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1ȸą,ȅƅ1ʚßǖůô E4D-0(ė 7), p[hèũ1˅HEWn 2 2 ERF
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3!F?`aʶŭȤůIȑ (Shoji et al. 2010), $2ŭȤů3 MYC2 1Ãķ E?2.
ɍCFE¢qfyz}~`°ʌƊĩ2&>1, °ʌǗʹ1Ţ˔ EťʥǊˌ2Ɗĩ@Ɠ&0ȷʗ2¯ð0/2źǔF;-1ʄ<CF,&ŵïÁ3ˉCF,E(Yun et al. 1992) īǊˌC0EȷʗÙ½IȻƁȀ1[m EèũĖĵIƊĩ E.-, ƣȵǴǩ2ǳǴůI˛ʘȀ1ĥĬ-EāɓůE(Broun 2004, Grotewold 2008) Ʃț-ȴ¬&p[hǳąŵ2afʚßĖĵ3, »p[hĩǺ2ùĖʱ¶ĵř1[nF,E.C?, LSNnʹIƹǩ½Ý-Ľˍ
1[m,E.ɱ¯F,E ;&, ǳąŵʱ¶ĵ|f2Ǫļʴã
T.Shoji & T. Hashimoto-5
BSJ-Review 2:34 (2011)ƹǩȖĸƣêɁ 2:35
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1ȅƅÒE.?ƚC.F, afʚßĖĵIǖǵ&Ɠ&0°ʌƊĩ2Ʒı2ƿ
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Eǡ3Ǫ1ɜċǚ(ė 3,

Menke et al. 1999, van der Fits & Memelink 2000, Shoji & Hashimoto 2011d) ȆĆĖĵ®2`aʶŭȤů2ƤǵǳǱǖůĭǣǩąŵȮ (Gundlach et al. 1992, Blechert et al. 1995, Yukimune et al.
1996)1?WmF,Eāɓů?Ekp{
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59: 735-769.

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230-0045 ǀǘŐ˥ɷõƨŖǷ 1-7-22
Functional identification of secondary metabolism-related genes by coexpression analysis
Keywords: coexpression analysis; functional genomics;
metabolomics; secondary metabolite; transcriptomics.
Masami Yokota Hirai
Metabolic Systems Research Team, RIKEN Plant Science Center
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2010) əê-maW|m ɼƯIɭĢą13˫NWLN1°HD˫Ɨ [am),,Eǆ°_Y]1BEɼƯĥ,E'G &Żɮ@j2ʨǈ1BD˫F;-;Dź2¯CF,0)&īƾ0¦ǆ°ʌǴǩ2ǳąŵ2ɼƚ

M.Y.Hiraiˬ5
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