΅ῡῐῩ῟Ὸ ῌ ῞῎Ί῞ ῲῦ; ῷc5de3 ,2 kDa ῒῡῌ Ῥ c5de ´ῌ ῒ῾ `"ῤ$῟Ώ ῲ(Ὶ῟΅Ῠῦ`ῥῌ 3 /2 kDa, ῨῺc5de3 0*῎0/ kDa fΊg\῎῍ Q -ῳῲ(Ὶ῟/0 `"ῤ$῟΅ῢῬῦῧ῞456 5hijῨῺ7Ῥ 3 AP 3ῠ῏῟4mnΊopqῢ ΅ῡῐῩ῟ῦ`ῤ78῞ῘῑῙ῟Ὸ ΐ ῠ?@Ῑ j3ῌ ῍ῢ῎)) 7> ῶ 3ῖΰ3ῠ῏῟ , m ῖ῍ ῒῴEῚ ῌ ΅ῡῐῩΐ/0 `"ῤ$῟FG ῗIῚ ῝ῌ -ῳῌ ῶMῶ ῌ ῼOP `"ῤ$῟QRS nΊopqῢj3ΐ῍ῢΐῌ sΰῡῌ tu3 ῳ` Tῌ Uῌ Vῌ ῼOXῌ Y῭Ὺ`]῝_`῝ bΌ Ὺ῟ῑyῢῤzῒ῞{|῟ῠῢj3~ῐῡΰῢ῍ ῞ 456fῐῧ78ῠῷhi6jΊΐῌ k῏῞Gmn )) AP 3ῧῴ ` (phytepsin) ῌ /- kDa oῙῑ6῍ jjῚ ῌ ῡῧ"Ύΰ`r΅ῡῐῩῠ ῞ῌ 3 ΅Ῠ Ὶ Pΰῌ -, kDa +0 kDa 3 s῟FtuvΊ w78῞ῘῑῙGxi6῍ ῭; Ὸῒῡ῞ῢU῞ , mn (.2 kDa) ῌ ,3 kDa ++ kDa 3῭; Ὸῒῡ῞ῢP῞ , mn yt ῡῧ"Ύΰ`r΅ῡῐῩ῟Ὼ| ῟ (.* kDa) 3 , Ῑ3ῦῨEῌ ῥjῙ(,)῍ ῝Ῐῡ3 uv ῴ , mnjῥqῢ῍ fῗῌ > ῶ 3> ῶ` ῦ´ ῴῦῌ (AP ; EC-...,-) ῟ A (cardosin A) ῌ 0. kDa 3 ΅Ῠ ΐ Ῑ῎Ὶῌ ῒῡ ´ Ί !ΐ#῞ῤΰ ῳ` ῪῥyῚ -/ kDa -* kDa ῒῡ῞ῢnΐ Ὶῗ῍ )) ῌ Ῡ Ῥ`ῌ ῨῺῌ Ῥ ῞῝3 Ίῌ AP | (plant specific inser- 4567ῌ ` ῨΌῷῲῷῌ ῦ; ῷῌ <ῌ > tion : PSI) ΐzΰῚ -+ kDa +/ kDa ῒῡ῞ῢ ῶ ῌ >>)ῌ Ῠ@ῌ Ῠ Ύ ῌ ) ῌ ῾ ῞ῢ(-)῍ s3ῠ῏῟ῌ AP 33 Ί Ῡ EῩῌ ῼῬῌ ῌ <G ῞῝3I56 Q5῭Ῠῲ3|῞ῢῦῨEῌ ΐK¡qῢ῍ ῖ3¢ ῒῡJK῍ῢ῎NOPΰῚ῎ῢ ῏~ +ῐ(+)῍ Q5῭Ῠ ῟jῌ > ῶ 3῭Ῠ ` (.)ῌ <G 3 AP(/) Ί£ ¤3 ῳ` Ὺΐa¥PΰῚ῎ῢ῍ Plant Aspartic Proteases ῌ Diversity and Functions ¦§ AP ¨©῟ῌ +-A ῟ªqῠ῏῟ῌ « ,* Tomiko ASAKURA, Tomoko TAMURA, Kaede TERAUCHI, ῦ Ώ ῒῡ῞ῢ`Ὺῷ ῟®ῌ « .* ῦ Ώ Keiko ABE, ῍+STUVUVWXVYZ[V ![ῌ ῍,\]^_ UV`aVb 3 ῌ ¯ῥ°± N ²³´µ¶ῌ ῟| + ῍ Ῐῐῒῑῖῐῌΐ῏ῗ῎ῙΊ῟ Ὺ ῍῎*L l῟ῤΰ.ῤ῭ ῐῧz ῲN´ῼῼ Pasp A+ (At+g++3+*) ῐXyῌ ῌ ´ῌ ῌ CῸNEΌῩῌ ῪJΎE῟CqῲEῬ +1 ῌ Pasp A, (At+g0,,3*) ῐXyῌ ῪJΎE῟CqῲE +1, ,0 Ῥῌ Xy῟ῌ# Pasp A- (At.g*..0*) ῐῌ +1 CDR+ (At/g---.*) }-~ B¤¥C* ,. N oryzasin+ Xy Ί[E\Ύ]῟Ύ¦ +. Ὺ phytepsin Xyῐῌ ῌ ´ῌ CῸNEΌῩῌ ῪJΎE῟CqῲEῬ +, ῠ§ῐῠ, §}-῟Ύ¦ ,1 ΅ Ὺ WAP+,WAP, Xy Ί[E\Ύ]῟Ύ¦ ,2 ῴN soyAP+ ῧ©Xy o-ῠ§ῐῠ, §}-῟Ύ¦ +0 soyAP, ῌ ῌ ´ῌ o- o-[E\Ύ]῟Ύ¦ +0 FF TcAP+, TcAP, Xy FFH῟ῧ$p.῞ῐΰ ,3 Xy[E\Ύ]῟Ύ¦ FGΏE cyprosin ῐ -* cardosinA ῐ CῸNEΌῩῌ Ῥ῟ΰªῬ§῟ῌ῝ +2 cardosinB ῐ}-΅῎}-~ +3 «ῷF­ nepenthesin Ὶῦῲ®ΰOΎῩoῙ῟`)* +/ ¯ Xy CῸNEΌῩῌ Ί[E\Ύ]῟C῾CΏ῟ -+ Ύ¦ qN±«ῼ[ Xy ´῟ ` -, ῭ VuAP+ ´ ³ΐUῺJU῞ΰ´µ* -- NE῭E ´ ³ΐUῺJU῞ΰ´µ* -- ¶U῭ DSA. ` -. Ὼ Ὼ LeAspP ´ ·¸¹S -/ ῳ »N StAP+-StAP- }-~½¥ῼ -0 ῞ΰῷ +** Ῠ ῑῡ῝ΰ PSIῌ Ὶ C ῲE D ῟cdῌ FῸCῲE D ῒSaaῌ R E ῖῑῡῘῤῚ῎ΰ῍ ῝Ὺ AP ῌU-0-SE ῞ f῞Ὸῤΰ῟῞'(Ί῍ ῒ ῞Ὶῐῢῌ ῢῥῌ#$ῒ῟ ῠ ΰ῍ ῤῚ῎ΰ῍ ῡ῞ῌ , Ῑ῟)*Ῑ+῟ῥ῾ PSI ῲE῟ ῟*ῌ PSI ῒ AP ῟ ῑDDTG ῐῠ, DDSGῒῌ C ῖ῟*-῎῍ Ὸ῞gῺῚ῎ΰhῤ*ῦijΰ8῟Ί῍ΰ῍ ῡ ῞ ῑῡ8ῌ @ῩῸCῲE῟ PSI ῐῠ,ῌ#[E \Ύ]῟*῞+kdl῟ῤΰ*Lῒ Ῐ AP ῝Ὶ῞ ῖ῍ 9῟ῌ `Gῳῦnῖo-R῟Ὸ῟p. PSI ῟.ῤῌ /ῑῑῡ0ῴ῟1῝ῘῚΐῖ ῑ5 Ίῌ `Gῳ῟+kdJqC[ῌ[E\Ύ]῟῏rῦ +ῒ῍ Ὺ AP ῟ ῌ 6῎῞-῎*ῦ8Ῑ9 8ῖῡῌ @ῩῸCῲE῟o-Ὸ῞gῺΰ9ῒs ῦ:῞;<ῖῒῌ ῟ῙΊ PSI Ῠ>*῟῞῎ ῏ῡῤΰ῍ 9῟uῐ῟8῞ῌ @ῩῸCῲEῑῡ PSI ῦ Ί῍ΰ῍ @ῩῸCῲEFGΏῲE A ῟ vwxῖΎyῦῧzxῖ{|Ίῌ PSI vw`W ῑῡῌ PSI )*ΎJ@ῺῑῡῤῖLῗ῞ῌ ῢ ῒ}-~῞ΎῩῤΰ9ῒῶῡῑ῞῝Ῐῖ῍ ῖῌ $ ῖΏ NEῦῌΰ9ῒῥῑῘῖ(0, 1)῍ )*Ῑ ῒ)*῟6ῑῡ PSI ῦ<ῖῳ8῍ΰ῍ 9ῡῌ +ῦΌΰ , Ῑ῟Ώ NEῒ b ῲῌῺ῞"ῧῗ ῦ ῡῒ῎ῗῖN AP ῟ῙΊ῍ΰSῲE + ῟ ΰ῟῞ῌ PSI / Ῑ῟ a RSῶΎUῑῡ῝ῢῌ PSI ῦvwxῖ`WῦῚ$ῒ)*ῦ<ῌ PSI ῦV$ῒ#ῌ ῞8WX῝8῟Ί῍Ῐ PSI vwSῲE + 8 *Ί`῞ῠῘῚ ῖ ῑ5 +-Bῒ῍ PSI ῟ ῌ ῲEΏ (saposin- )*῝ῢῌ $ῒ)*ῦΰ9ῦiῖ(2, 3)῍ like) [E\Ύ]῟ῙΊ῍ΰ NK-SῲE῞Ὶ῎ ῖῌ , X῟ῴN AP, soyAP+ soyAP,, ῐῠ, ΰ (C)῍ ῲEU@ῩE`_])*`[E\Ύ]Ί 9ῤῡῒ PSI ῦvwῖ soyAP+DPSI soyAP,DPSI ῍ῢῌ Saaῌ ῞Ὶ῎ΰ῍ ῲE C FῸC ῟῏῟ C ῖ῞ GFP ῦkῌ ῲN´ῼῼῦ< ῝ + ῍ Ῐ῏ΐῖΊΐῌ (AP) ῐῚ῟ (A) ΎΎ ΅ AP ^_ῺaM῝¢ΏῬῺaM D ;<6῍ (B) ^_ῺaMῬ;<῍ ¤¥Fῢ AP Ῐ*2¦Ὶ῝ (PSI) .ῌ , Ὶ1§῍ῧi¨Nk΅M©ª«¬¥.%῍ (C) ^_ῺaM PSI ῝ῲ aM5MN`P NK-raMῬ; <῍ ῎ῖῨΰ / Ὶ a rῸ`¨ΰῚ῍ ¤¥°±¨m^_ῼ³6'%῍ i ´©῝ a rῸ`¨ῡଥῡῗ¶¥ ῤ·6'%῍ ῝¸ῡῗ¹º· N 8ῑῒῥºῐῗ»῟῍ H+῎H/ a rῸ`¨6'ῌ ¼ΰ῝ῌ½%ῧ῍ ῩῗῚΊῩῙ ῖῗ῝ Ῡ(+*)ῌ soyAP ssica napus AP a]cd´f´ ,S im῾ Mop +-GFP ῝ soyAP,-GFP ῎ῖῨΰῩῨῗ qῒ(+-)ῌ ΎrῳaM + mῺrMΰB ΐῌ soyAP,DPSI-GFP Ῡ΅ΐῠῌ %ῧ(+.)῍ ῡῩ soyAP,DPSI ΐῩ (ER) ΐῨ Fῢ AP KMN`P]gaMvwΰ ῗ῍ Ῠῌ soyAP+DPSI ῠ῝!῞ῩῩ ῡxῡ῟y %ῧ ῝ΐ.ῨΊῗ῍ G῞z ΅ΐ#$Ῠῗ῍ %῟'Ῐῌ soyAP+ ῝ soyAP, {|ῒῥ}"Ῠῧy῍ῦῌ FῢῪΌ Ῥfk PSI ῌ Ῡ)Ί*῟ῧ,ῪΰῚ ῝ΐ./ H Nepenthes distillatoria AP (Nepenthesin I, II) Ῠῗ῍ soyAP+ ῝ soyAP, PSI `01ῌ 2῭ ῦῥῐῧ῎S ῞zῌ ΰ&S 3ῑ5ῧ`01῝6%ῧ῝ῌ Ῡ΅῞ 1)(Ί῎ῧ῝}"Ῠῗ(+/)῍ ῤ῏῟῝ῡῗῬ;<ΐ=῍ῧῒῌ @AB )ῥῌ *AHIῳῒῥqῥ, ῨῧCE῏῍ῧ῍ ῌ RHI.῟῞zΐΊῢῥῨῗ soyAP+ ῞z in situ c῾rῶcῴῌa MῤῙΊBῗ ῏῝ ,ῐ῍ soyAP+ ῞zῌ ῲῡῘ*2῞z%ῧΰ Ῑ῎ῒῑ῞ῗ I ῌῬῌ῍ῧiam CoA Ύ῭aῶῌῴa´m Fῢ AP GFῢHIῒῥῐΎῨΊῗ ῝ῒ ῡ῝῟Ῑῗ῍ ῌ Ῑ3Ῐ*2῞z%ῧΰI ῥῌ AP ῤῧHIKMN`PΰBῌ QRS) ῌῬῌ῝Ί SCARECROW Ὺ῎ῗῌ soyAP+ %ῧῧTΐG῎῍ ῗ῝ῐVῌ ΎΎ ΅Z`ῷM] Ῑ3Ῑ῍ῧῩ῍ῧῲῩ. ^_ῺaMῤῙΊΰῌ a]cd´f´ PaspA+ ῞zΊ῎ῧ ῝ΐ'ῒῙῗ῍ Ῠῌ soyAP+ ΐ ῤῙΊΰ]gaMῨῧ(++, +,)῍ i῾k´l Bra- ῨῥῩ ¡ΰBῌ ῍Q)(Ί῎ῧ ῝ Ῐ , ῍ ῐΐ῏ῒ ῑ ῗ ῏ῌ (soyAP+) ῎Ῑῖ(+0) §¨ .2 ©ª«΅ῨῪ­a® ῒῢ ®῏°±ῌῧ²³,ῌ ´` aῡῤ in situ µῨῴ´΅ῨῬῌ ῼ0ῧ¸῟ῙῘ῍ soyAP+ ¹ &ῐº»,Ῐ῍ Terauchi et al. : Planta, ,+2, 3.1 (,**.) ῧ ½ ῧῖῤῠ῍ῤ(+0)῍ 0 A RGD j ῶyῶ´>ῌῬ D (PLD)a C, Ῐῌ ῖῤ AP ῠ῎ῥΊ῎ῤ῍ ῳΌῨ0ῧuvῖῤ3῝ΐῑῢῒῥῌ Ῡ.ῳ0 A ῨῪ PaspA- Ώ῝#$%῍&ῐ ΐ&ῐlm|SῡῙΊ uvwῒ,Ί῎ῤ ῖῤ(+1)῍ '( *+ῖῤ AP ῝,ΊῩ.ῳ0 ῟῎ῒ῝῎ῦῥΊ῎ῤ῍ Ῡ.ῳ0 A, B ῧ12ῤ3῝ΐῤ(+2, +3)῍ ῷ.ῲ# ῏ῌ Ῡ.ῳ0 B mRNA ῎ ῖ .89῟`ῌῪ῍ῤΎ.> (Serpa) ῌ ΐ? ῤΐῌ ,Ῐ ,,῟῎῍ & Ῡ.ῳ0 @ΰBῧ ῐEFῗΊῚῢῥῤ῍ 3 ῐ῝&ῐῸῲ´$y*+ῖῤῩ.ῳ *+ῖῤE?JKΐῩ.ῳ0 A, B ῍ῤ῍ 0 B ῌ ,opῡῙΊῥΊ LNOP 1-῏῝Q῎RSῧῖΐῌ ῎Ὶῒ ῎ῤ῍ 3ῥ AP Ῐ῟ῧ,ῘΊ USVΐ῟ῤ῍ Ῡ.ῳ0 A WX?X῭0> ῍ῙῘ῍ 3ῡ῏ AP ῟Sῌ $VZ[\ ῐQ^_*+,ῌ `ab RGD Ὼ Ῐ῟ AP ΐ῾8a¡. ¤ῥῤ3῝ῡῙΊN῎ `ῌdῧῠῚ῍ RGD Ὼ`ῌd?Ῠ0f´ ῑῢῒῥΊΊ῎ῤ῍ ¥ + AP ¦+῝ 0hij ῝,Ίkῢῥῌ &ῐlmῌ ` ῌ opῌ f Ὶ῎Ί,Ῐ῍ .8q῟῞ῑtῤ&ῐuvwῒ,Ί῎ῤ῍ Ῡ.ῳ Ὺῌῷ῝ΊRΊ῎ ῝ῐῨ ῍ Nucellin ῙῗῖῚ῎ῐΐ῏ ῒ῎ῌῑ῍Ῐ ῑ``wῲ῾2Ῐΐῌ ῬῥῗF<Pΐ ῢῩ῝῟ῙῘῧ AP ῍ῦ ῼΊ῎ ῍ ῒῨῌ AP Ὸ ῟ῢΏ ΰΐΊ῎ ῝(,*)ῌ ! pH ΐ"῝#῍ -.PCS+ Ῡ ῴΌῺ+ῡ, ῝῝῎ῙῘ-῝. T 0* ῍O<ῬQQ AP 9ῌS{ Ί῎Ῐ῍ ῒ/´ῌ Ῠ AP ῝῟ ΌῬ T-DNA ΌP=Ῥ+῍U῍ῌ 5ῧ ῴῳ AP ΐ1῎῝῾2ῌ 3 4Ῑ῎5 lῌ[+P῟ῙῘ῝ῌ promotion of cell sur- 6Ί῎ ῝ΐ Ῠῒ῝῟ῙΊῘ῍ vival + (PCS+) WῘῌ Xῠ῝ ῠῖRΐΊῐῌ >῾[ῑ῎Ί\῟:%} +.CND.+ Ί>῝Ὺ}῝῟ῙῘ(,-)῍ \῾5῟_῟ CND.+ ῴῑῌ Ό῾9#$:%ῳ<< =ῴ´? PCD ΐ <ΏῳΊXῠP`῝ a:%ΐ &ῨῒῨ()ῌ ῗ .+ kDa AB῝Ό+ῳ. ¡ Ῐῤῌ P£Ό¤ ΐcῐ῟῟Ῡῌ CND.+ (chloroplast nucleoid DNA-bindig protein) ῝ d῝῟ ῍ PCS+ Qe:%῝¥ῐ¦:% *+Ῐ(,+)῍ CND.+ AP , ῚE῝Ύῗ῝Ὶῲ ῒ%Ῠ῾5{ ῍ PCS+ anti-PCD ~ῌ :% FῺῌ ῐFDTGS ῑῧK DSGῑ ῌ +,* Ὺ ¡Ῠῖ Ῐῤ¨9ῥῚ ῺS <ῶO+P MN"ῒῨ῟ ΅῎OPQR ῺSῥ῎Ί DNA T ῍ ῎E῝z{ ῝6{ ῝ῐῨΊ῎ ῍ U#῍ Lys ΰ helix-turn-helix (HTH) SW ©hB.ΐῶῲªῌ >῾¦ῑr Ῥ+ῥῚ῍ CND.+ ῾53Ῐ´=+ῴY῭ PCD W[w¬ 6῎jkῚ῍῏῍ [ΏῳΌ῾9#$:%ῳ<< =ῴ´] ΐ^῾5ΊῑῩῌ CND.+ ΐῳ<< =ῴ´ ..CDR+ ῾5 _Ί῎ ῝ΐ`ῒ ῍ 6ῬaῧῩ O<ῬQQ CDR+ ῌ T-DNA ῪῳῼΎ ῌ bῘῠcῐ CND.+ ῌ B.῝Ί FP< + O¯+Όῲ+PῧῙΊ Pseudomonas syringae l῝ 9῎Ῐ:Uῌ ! pH ,῏.ῌ ῴΌῺ+ῡ῝h῞ ῝Ί Xia ῨῧῙΊῷῨῘ(,.)῍ Ὺ° =ῴ´ E῝ΐῡ,jῌ ῶl+ <ῼῪῌῷῬ+ Όῌῌ ῦ{ Ῐ῎ΌῬ AP ῌ \῾5{ OῴῼῬ+ <ῼῪῌῷῬ+ Όῌῌ EDTA pq constitutive disease resistance-Dominant (cdr+-D) ῢῙῘ῎r῟῎῍ "῝#! pH ῥῚ CND.+ ῌ ±lῺR"²oΐeΊ῎ ῝ῒῨῌ ΐῌ ῳ<< =ῴ´sB.῝ῌ ῞ῧ῏῟E῝ AP ῧῙΊ῞ῘΌ ῺS΅lOΌῌΐ± ῾uΊ῎ ῒῚ῎Ί ῍ ῍ lῺR"rNeῖ ῝῎`Ί῎ ῍ CDR+ E῝Ύῗ῝῟ Ὺῴ=ῲ+"Ὺῴ=ῲ ,. Nucellin +scῘsῨ\῾5ῖῘῌ cdr+-D ``wῲῌῗ:%>ῌΐ῎Ῐy?z{ ῍ 5³.´Ίu῝ῶ& 5µ{῝ῒ ?z|῍:%} (programmed cell death : PCD) Ῠῌ CDR+ <ῼῪῌῷE῝ΐ cdr+-D 5³. ῍Ῡῌ <ῼῪῌῷΐ PCD ῏B῟_~ ῐ5`w6¶ΐ῍ ῝ΐµΊ῎ ῍ ῍ ῍ Nucellin ``wῲ?zῌῗ῾5{ 6ῬaῠcῐῨ9῎ΊῷῘ CDR+ ©h3 Ὺ ΎῒῨῴῳlῌ[+PῘ(,,)῍ .+* ῪMN" <ῌῺῒῨῌ ῗE῝ ῴΌῺ+ ῡ 9ῌSῌ 2 Ὶ΅Ὸ+ΐ 1 ῚῬ+´<+ ,ῌ b-ῶῳ·Όῌῷ (BACE) ῡ,¸῍ .-(,- DΊ῎ ῍ , ῚE῝Ύῗῌ AP Eῦ aminoethyl)ῌbenzensulfonyl fluoride (AEBSF), ῑῧ Ί῎ ῌ DDTG ῑῧK DDSGS ῍ ῍ G K EDTA ῧῙΊῥE῝ΐ { ῝ῒῨῌ AP ῍ AP ῐHῑ ῝ ,*ῒ ῝ῒjῌ < ῝ῶ¦¹ῤῬ`+E῝῾5Bºῌ ῢῘ»¼¸ ῥ PSI ῥῦ῟῎῍ Nucellin ῎ y -῏. Ὼ ῧῙΊῥῡ,῎r ῝ῒῨῌ OῴῼῬ+¡BῥE Kῥῧ῾5{ ΐῌ ῎ y +* Ὼ{ ῝ ῝῾56Ί῎ ῝ΐµῘ῍ ! pH 0.* `jῒ῾5ΐῐ ῳNΐῌ Pῌ Qῌ ῏0./ ῝½"῝#῍Ῡῌ ·[+ῦ bῌῶῳ·Όῌῷ ῐ῟῎῍ ῗ῾5῎ yῌῗ:% PCD B.Ῡ{ E῝ΐ῭ῠῨῘΐῌ Ῠ¨9ῖj ῝ΊῑῩῌ ῌῗ:% PCD { <ῼ FTC-ῷῬ+`jῒ ¬Ῐ῍ e in vitro ῑ ,/ ῟ - ῍ ῒῗῐ ΐ ῒῑ ῖῌ (AtSPP) ῏Ῑ῝῎Ί ῞( ) AtSPP ῏ῥ· SPP-like (AtSPPL+, AtSPPL,) ' GFP -:*ῌ `1>ῷῶῶῠῩΐῨ%῝ (Deep cell) '¸῏*Ῐ῍ AtSPP X῝Y'ῌ AtSPPL+, AtSPPL, Ὺ῾ῳῌ¦'.*ῌ AtSPP ¹º῟"%῝'ῑ»¼ῩῘ῍ Tamura et al.
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