΅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 V῱ῌ ῼOXῌ Y῭Ὺ`῰]῝῜_`῝ bΌ Ὺ῟ῑyῢῤzῒ῞{|῟ῠῢj3῜~ῐῡΰῢ῍ ῞ ῕456fῐῧ78ῠῷhi6jΊΐῌ k῏῞Gmn ))  AP 3€ῧῴ ` (phytepsin) ῌ /- kDa oῙῑ6῍ jjῚ ῌ ῡῧ"Ύΰ`r΅ῡῐῩῠ ῞ῌ 3 ‚ ΅Ῠ ῜„Ὶ †Pΰῌ -, kDa ῜ +0 kDa 3 s῟FtuvΊ w78῞ῘῑῙGxi6῍ ῭; Ὸ‰῵ῒῡ῞ῢU῔῞ , mn (.2 kDa) ῜ῌ ,3 kDa ῜ ++ kDa 3῭; Ὸ‰῵ῒῡ῞ῢŠP῞ , mn y῵t ῡῧ"Ύΰ`r΅ῡῐῩ῟Ὼ| ῟ (.* kDa) 3 , Ῑ3ῦῨE€‹ῌ ῥjῙ(,)῍ ῝Ῐῡ3Ž uv ῴ , mnjῥ‘qῢ῍ fῗῌ > ῶ 3> ῶ` ῦ῱´  ῴῦῌ (AP ; EC-...,-) ῟ A (cardosin A) ῌ 0. kDa 3 ‚ ΅Ῠ ΐ Ῑ῎Ὶῌ ῕ῒῡ ´ Ί !ΐ#῞ῤΰ ῳ` Ὺῥ’yῚ -/ kDa ῜ -* kDa ῒῡ῞ῢ“”nΐ Ὶ῔ῗ῍ )) ῌ ῵Ῡ Ῥ`ῌ ῨῺῌ Ῥ ῞῝3 Ί῔ῌ  AP •|–—˜™š (plant specific inser- 4567ῌ ` ῨΌῷῲῷῌ ῦ; ῷῌ <῿ῌ > tion : PSI) ΐzΰῚ -+ kDa ῜ +/ kDa ῒῡ῞ῢ†› ῶ ῌ >>)ῌ Ῠ@῰ῌ Ῠ Ύ ῿ ῌ ) ῰ῌ ῾ œ῜῞ῢ(-)῍ s3ῠ῏῟ῌ  AP 3†›ž3Ÿ Ί Ῡ EῩῌ ῼῬῌ ῵῿῵ῌ <῿G ῞῝3I56 Q5῭Ῠῲ3|῞ῢῦῨE€‹ῌ 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 ™š῟®῔ῌ « .* ῦ Ώ Keiko ABE, ῍+STUVUVWXVYZ[V ![ῌ ῍,\]^_ UV`aVb 3 ™šῌ “¯ῥ°± N ²³´µ¶ῌ ῟•| + ῍ Ῐῐῒ῕ῑῖ῔ῐῌΐ῏ῗ῎ῙΊ ῟

Ὺ῵– ῍῎*L ™ l῟ῤΰ.ῤ῭š ῐῧz—˜

ῲ“N´ῼ‘ῼ Pasp A+ (At+g++3+*) ῐXyῌ ›ῌ ´ῌ œῌ C“ῸNE῿ΌῩῌ †† ῪJΎ῵E῟C“qῲEῬ +1 ῰ ῌ ž Pasp A, (At+g0,,3*) ῐXyῌ ῰ †† ῪJΎ῵E῟C“qῲE +1, ,0 Ῥῌ Xy῟ῌ# Pasp A- (At.g*..0*) ῐžῌ ῰ +1 CDR+ (At/g---.*) }-~ B¤¥C* ,. N oryzasin+ Xy Ί῕[E\Ύ]῟Ύ¦ +. †† Ὺ phytepsin Xyῐ›ῌ œῌ ´ῌ ž C“ῸNE῿ΌῩῌ †† ῪJΎ῵E῟C“qῲEῬ +, ῠ§ῐῠ, §}-῟ƒΎ¦ ,1 ΅ Ὺ WAP+,WAP, Xy Ί῕[E\Ύ]῟Ύ¦ ,2 ῴN‘ soyAP+ ῧ©Xy o-ῠ§ῐῠ, §}-῟ƒΎ¦ +0 soyAP, ›ῌ œῌ ´ῌ žo- o-[E\Ύ]῟Ύ¦ +0 FF† TcAP+, TcAP, Xy FF†H῟ῧ$p.῞ῐ῕ΰ ,3 Xy[E\Ύ]῟Ύ¦ FGΏE cyprosin žῐž -* cardosinA žῐž C“ῸNE῿ΌῩῌ žῬ῟ΰª῜žῬ§῟ῌ῝ +2 cardosinB žῐž}-΅῎}-~ +3 «ῷ῿F‘­ nepenthesin Ὶῦῲ®ΰOΎῩoῙ῟Ž`)* +/ ¯ Xy C“ῸNE῿ΌῩῌ Ί῕[E\Ύ]῟C“῾C῵Ώ῟ -+ Ύ¦ qN±«ῼ[ Xy ´῟ ` -, ῰῰῭ VuAP+ ´ ³ΐUῺJU῞ΰ´µ* -- NE῭E ´ ³ΐUῺJU῞ΰ´µ* -- ¶U῭ DSA. ž῰ ` -. Ὼ Ὼ LeAspP ´ ·¸¹S -/ ῳ »N StAP+-StAP- }-~½¥ˆῼ -0

῞ΰῷ +** Ῠ  ῑῡ῝ΰ PSIῌ Ὶ C  ῲE D ῟cdῌ FῸCῲE D ῒSaaῌ R E ῖῑῡῘῤῚ῎ΰ῍ ῝Ὺ AP  ῌU-0-SE ῞ f῞Ὸ῔ῤΰ῟῞'(Ί῍ ῒ ῞Ὶῐῢῌ ῜ῢῥ῕ῌ#$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]῟*῞+kd῜l῟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῔῞gῺΰ9῜ῒs ῜ῦ:῞;<ῖῒῌ ῟ῙΊ PSI Ῠ>*῟῞῎ ῏ῡῤΰ῍ 9῟uῐ῟8῜῞ῌ @ῩῸCῲEῑῡ PSI ῦ Ί῍ΰ῍ @ῩῸCῲE῜FGΏῲE A ῟ vwxῖΎyῦῧzxῖ{|Ίῌ PSI vw`W ῑῡῌ PSI )*ΎJ@ῺῑῡῤῖLῗ῞ῌ ῢ ῒ}-~῞ΎῩῤΰ9῜ῒῶῡῑ῞῝Ῐῖ῍ ῖῌ $ ῖΏ NEῦῌΰ9῜ῒῥῑῘῖ(0, 1)῍ )*Ῑ ῒ)*῟€6ῑῡ PSI ῦ῜<ῖῳ8῍ΰ῍ 9‚ῡῌ +ῦΌΰ , Ῑ῟Ώ NEῒ b ῲῌῺ῞"ῧῗ ῦ ƒῡῒ„῎ῗῖN AP ῟ῙΊ῍ΰ†S῱ῲE + ῟ ῜ΰ῟῞ῌ PSI  / Ῑ῟ a RSῶΎUῑῡ῝ῢῌ PSI ῦvwxῖ`Wῦˆ‰Ὶ$ῒ)*ῦ῜<ῌ PSI ῦV῔$ῒ#῜ῌ  ῞8WX῝8῟Ί῍Ῐ PSI vw†S῱ῲE + 8 *Š‹ŒΊƒŽ`῞ῠῘῚ ῖ ῑ5 +-Bῒ῍ PSI ῟ ῌ ῰ ῲEΏ (saposin- )*῜῝ῢῌ $ῒ)*ῦ῱ΰ9῜ῦiῖ(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 ῦ῿kῌ ῲ“N´ῼ‘ῼῦ< ῝ + ῍ Ῐ῏῜ΐ῕῔ῖΊΐῌ (AP) ῐῚ῟ (A) ΎΎ ΅ AP ^_Ὼ῿aM῝¢ΏῬῺ῿aM D ;<6῍ (B) ^_Ὼ῿aMῬ;<῍ ¤¥Fῢ AP Ῐ*2¦Ὶ῝῰ (PSI) .ῌ , Ὶ†1῔§῜῍ῧi¨Nk΅M©ª«¬¥῜.%῍ (C) ^_Ὼ῿aM PSI ῝ῲ aM5῵MN`P NK-raMῬ; <῍ ῎ῖῨΰ / Ὶ a rῸ`¨ΰῚ῍ ¤¥°±¨m^_ῼ™³6'%῍ i ´©῝῰῜ a rῸ`¨ῡଥῡῗ¶¥ ῤ·῜6'%῍ ῝῰¸ῡῗ¹º· N 8ῑ›ῒῥºῐῗ»῟῍ H+῎H/  a rῸ`¨6'ῌ ¼῔ΰ῝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΅ΐῠ῜ῌ  %ῧ(+.)῍ ῡῩ῜ soyAP,DPSI ΐῩ (ER) ΐ Ῠ Fῢ AP K῵MN`P῿]gaM῰vwΰ ῗ῍ Ῠῌ soyAP+DPSI ῜ῠ῝!῞Ῡ῜Ῡ ῡxῡ῟y῜ %ῧ ῝ΐ.ῨΊ῔ῗ῍ 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 ˆ‰*AHIῳŠῒῥ῱‹qῥ,Œ ῨῧC῔E῏῜῍ῧ῍ ῌ RHI.῟῞zΐΊῢῥῨῗ soyAP+ ῞z  in situ c῾rῶcῴῌa MῤῙΊB”ῗ ῏῝ ,ῐ῍ soyAP+ ῞zῌ ῲῡῘ*2῞z%ῧΰ Ῑ῎ῒῑ῞ῗ I ῌῬῌ῜῍ῧiam CoA Ύ῭aῶῌῴa῰´m Fῢ AP G῕FῢHIῒῥῐΎῨΊ῔ῗ ῝ῒ ῡ῝—῟Ῑῗ῍ ˜ῌ Ῑ3Ῐ*2῞z%ῧΰI ῥῌ AP ῤῧHIK῵MN`PΰBῌ QRS) ῌῬῌ῝Ί SCARECROW Ὺ῎ῗ™šῌ soyAP+ %ῧῧTΐG῎῍ ῗ῝ῐVῌ ΎΎ ΅Z`ῷM῿] Ῑ3Ῑ›῍ῧœžῩ῜῍ῧῲŸῩ῜. ^_Ὼ῿aMῤῙΊΰῌ a]cd´f´ PaspA+ ῞zΊ῎ῧ ῝ΐ'ῒῙῗ῍ Ῠῌ soyAP+ ΐ ῤῙΊΰ῿]gaM῰Ῠῧ(++, +,)῍ i῾k´l Bra- ῨῥῩ ¡ΰBῌ ž῍žQ)(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῝#$%῍&ῐ ΐ&ῐlm|SῡῙΊ uvwῒ,Ί῎ῤ῜ ῖῤ(+1)῍ '(῕ *+ῖῤ AP ῝,ΊῩ.ῳ0 ῟῎ῒ῝῎ῦῥΊ῎ῤ῍ Ῡ.ῳ0 A, B ῧ12ῤ3῝ΐ῜῔ῤ(+2, +3)῍ ῷ.ῲ# ῿῏ῌ Ῡ.ῳ0 B  mRNA ῎ ‚῜ ῖ .89῟`ῌῪ῜῍ῤΎ.> (Serpa) ῌ ΐ? ῤΐῌ ƒ„ ,Ῐ ‚†,‡῜ ,῟῎῍ & Ῡ.ῳ0 @ΰBῧ ῐEFῗΊῚ῕ῢῥῤ῍ 3 ῐˆ῝‰Š‹Œ&ῐῸῲ´῰$y*+ῖῤῩ.ῳ  *+ῖῤE?JKΐῩ.ῳ0 A, B ῜῍ῤ῍ ῔ 0 B  ῌ †,‡op‘ῡῙΊ’“ῥΊ L῿NOP 1-῏῝Q῎῕RSῧῖΐῌ ῎῕Ὶῒ ῎ῤ῍ 3ῥ ”• AP –Ῐ῟—˜ῧ™š,Ῐ›œΊ U῜SVΐ῟ῤ῍ Ῡ.ῳ0 A WX?X῭0> ῜῍ῙῘ῍ 3ῡ῏”• AP žŸ῟—˜S ῌ – $VZ[\ ῐQ^_῜*+,ῌ `ab RGD Ὼ Ῐ῟ AP ΐ῾8a¡῵.῜ ¤ῥῤ3῝ῡῙΊN῎ `ῌdῧῠῚ῍ RGD Ὼ`ῌd ?῕῜ Ῠ0῱f´ ῑῢῒῥΊ῔Ί῎ῤ῍ ¥ +  AP ¦+῝—˜ 0hij ῝,Ίkῢῥῌ &ῐlmῌ ` ῌ opῌ f Ὶ῎Ί,Ῐ῍ .8q῟῞ῑtῤ&ῐuvwῒ,Ί῎ῤ῍ Ῡ.ῳ Ὺῌῷ῝Ί‘RΊ῎ ῝’ῐῨ ῍ Nucellin ‡ ῙῗῖῚ῎ῐΐ῕῏ ῔ῒ῎ῌῑ῍Ῐ ῑ``wῲ῜῾2Ῐΐῌ Ῥ“῜ῥῗ”F῾[ῑ῎Ί‰\῟:%} +.CND.+ ῔Ί>῝Ὺ}῝῟ῙῘ(,-)῍ ‰\῾5›῜῟_῟ CND.+ ῴῑῌ Ό῾9#$:%ῳ<< =ῴ´? PCD ΐ <ΏῳΊXῠP῜Ÿ`῝ a:%ΐ &ῨῒῨ()ῌ ῗ .+ kDa AB῝Ό+῿ῳ. ῔¡ Ῐῤῌ P£Ό¤ ΐcῐ῜῔῟῕῟Ῡῌ CND.+ (chloroplast nucleoid DNA-bindig protein) ῝ d῝῟ ῍ PCS+ Qe῿:%῝¥ῐ¦Œ:% *+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.+ ῾53῜Ῐ´=+ῴY῭  PCD W῰[῵w¬ 6῎jkῚ῜῍™῏῍ [ΏῳΌ῾9#$:%῜ῳ<< =ῴ´] ΐ^῾5ΊῑῩῌ CND.+ ΐῳ<< =ῴ´ ..CDR+ ῾5 ῜_Ί῎ ῝ΐ`ῒ ῍ 6ῬaῧῩ O<Ῥ–Q῵Q 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"²oΐe῔Ί῎ ῝ῒῨῌ  ΐῌ ῳ<< =ῴ´῜sB.῝ῌ ῞ῧ῏῟E῝ AP ῧῙΊ῞ῘΌ ῺS΅lOΌῌΐ± ῾uΊ῎ ῒῚ῎Ί ῜῍ ῍ lῺR"rNe῔ῖ ῝῎`Ί῎ ῍ CDR+ E῝Ύῗ῝῟ Ὺῴ῿=ῲ+"Ὺῴ῿=ῲ ,. Nucellin +scῘsῨ‰\῾5ῖῘ›῜ῌ cdr+-D  ``wῲῌῗ:%>ῌΐ῎Ῐy?z{ ῍ 5³.῕´Ίu›῝ῶ& 5‚µ{῝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 ῐHƒῑ ῝ ,*ῒ „ ῝ῒjῌ < ῝ῶ¦¹ῤῬ`+E῝῾5Bºῌ ῢῘ»¼¸ ῥ PSI ῥῦ†῟῎῍ Nucellin ῎ y -῏. Ὼ ῧῙΊῥῡ,῎r ῝ῒῨῌ OῴῼῬ+¡BῥE K῜‡ῥῧ῕῾5{ ΐῌ ῎ y +* Ὼˆ‰{ ῝ ῝῾56 Ί῎ ῝ΐµῘ῍ ! pH  0.* `jῒ῾5ΐŠῐ῜῔ ῳN‹ΐῌ Pῌ Qῌ Œ῜ ῏0./ ῝½"῝#῍Ῡῌ ·[+ῦ bῌῶῳ·Όῌῷ Šῐ῟῎῍ ῗ῾5῎ yῌῗ:% PCD B.Ῡ{ E῝ΐ῭ῠῨῘΐῌ Ῠ¨9ῖj ῝ŽΊῑῩῌ ῌῗ:% PCD ῔{ <ῼ FTC-῰ῷῬ+`jῒ ¬Ῐ῍ „e in vitro ῑ ,/ ῟ - ῍ ῒῗῐ ΐ ῒῑ ῖ῕ῌ῔ (AtSPP) ῏῜Ῑ῝῎Ί ῞( ) AtSPP ῏ῥ· SPP-like (AtSPPL+, AtSPPL,) ' GFP -:*ῌ `1>ῷῶ῱ῶ†ῠῩΐῨ%῝ (Deep cell) '¸῏*Ῐ῍ AtSPP X῝Y'ῌ AtSPPL+, AtSPPL, Ὺ῾῵ῳῌ¦'.*ῌ AtSPP ῔Œ¹º‰Š‡῟†"%῝'῔‹ῑ»¼ῩῘ῍ Tamura et al. : FEBS J., ,1/, -. (,**2) ½Ῑ

῔Ῠ ῐῦῌ CDR+ ῎ῌ ΰ ῌ %῝῟' r*ῌ ῶisdEΎdῌ%῝'uv῞῍ ῜῟ῑῩῘ῍ Ῠ῟῕pwῨ῍ /AxῚ SPP  {'῏῎῝ @ῧA9}ῒ'Sΰ*῝῎Ῠ῍ /. ῘῚ AP ~῎῟'ῌ 8ῗ { A῎€ῗ' ῠ῞ !῝ῒῘ AP ῎$Ῡΰῌ %῝῍&' ῥ ´‚1῭ῑ-./Ῠ῍ ƒῦ ῌ `1>ῷῶ῱ῶ *Ῐ+,῞-./Ῠ ῥΏ123ῌῲ῞῍Ῠ῍ *ῐ '῏῎῝†῎ῦ῎‡῎ˆ῎‰Š' SPP ῑ῔‹*ῌ ῌ' *7ῑῌ 8ῗῌ 9ῗῠῩ῟':;/Ῠ<*῎=>Ώ ῔Œ‰Š‡ῤ†"%῝ ῖ'ΐ῔‹/Ῠ Ώ123ῌῲῑ῔?Ῡῌ %῝῍@ῧA`῭ῶEF ῟?῎Ῑ*Ῐ ῏῟ -ῐ῍ ‡ €ῗ“”ῖ/!῝• GHΊ῝῎Ῠ῟ῑῦῐ'Ῡ῝ῒῘ῍ Ῡῦ῟: –'Ῐ /Ῠ•–—ῡ῟ῐῦῌ SPP =ῌ΅šῴ῟ ;J ῖ῿ῑL῟῍'῞ῢNῠῩῌ ῟῍῞O AῨῘŠῤ9}ῒ'#œ'~ῑΰῘῩῨ῍ ῠῘῌ % PQ'ῘR/Ῠ RIP (regulated intramembrane ῝῍. GFP ῟ῡ;=῾ῺῬ῞R *Ῐ῟ proteolysis) 'Sΰ/Ῠ῍  AP T῟*῝U῕ῦ ¡ῌ %῎8ῗ SPP ῟¢ῤ'X῝Y῞῍ΊῘ῍ ῠῘ SPP ῩῨῑ signal peptide peptidase (SPP) ῞῍Ῠ῍ SPP '(£῎ῘŠ ῏SPP ῤῘŠῐ ῑ`1>ῷῶ῱ῶ' N WῑX῝Y῍Z'ῌ C Wῑ%῝['-./Ῠ / ‰-.*ῌ ῖ SPP ῤῘŠ Ὺ῾῵ῳῌ¦'.* 3 \῟]^J῟ῢ_Ῡῌ / \῟ 0 \῟]^ Ῐ(,/)῍ Ῡῦ§¨ €ῗ'῏῎῝©ῢ῝ª?῞῍Ῠ ῍'ῖῩῗῩ YD, GXGD ῖ῿ῑ-./Ῠ῍ SPP ῑῌ ‹.ῌ `1>ῷῶ῱ῶ῟ « Q' ῎>Ὸ῞ΰ ,**, ῑ' MHC Ῥd῰ I fg`῭ῶEῼΏi῵ SPP X῝Y'῏῔Ῠ῔‹®ῐ*῝῏ῧῌ *€ῗ'῏ ῟῍῞ῘR/ῨΏ123ῌῲ῟*῝῜kῩῘ῍ X῝Y ῎῝ΌšῬ°±῾²³´µ῞῍Ῠ῍ ῩῦR /Ῠ ῟῞ SPP 'ῥῧnῧoῩῘ`῭ῶEῼΏi῵pῚ ῟῞€ῗ+ΊQA ῏῍Ῠ῎ ¶€ῗ'ῌ QAῐ SPP  . ῏ ῟ῙῘ Ῐ Ὶ῝ῒῠ ῑ῔῕Ί῞ ῜῟ῗ῕ῌ ῖ Pre : presegment, Pro : proseg- ment, DTG ῎ DSG JῗYD ῎ GXGD : +,PXῸº»K (D) ῝ῒ .*ῦΊ῎ῥPXῸº»Kῌ PSI : plant specific insertion, HTH : helix-turn-helix DNA binding motif, Ser rich : Serine rich ¼ῌ PAL : Pro-Ala-Leu ½ ῌ CND41 : chloroplast Nucleoid DNA-bind- ing protein, NAP : Nepenthesin like AP, CDR : constitutive dis- ease resistance, PCS : promotion of cell survival, SPP : signal pep- tide peptidase, PN : presenilin

῝Ῡ῜ΐῥ ῝ῠῙῧῌ ῐ dase database ; http://merops.sanger.ac.uk/) ῌ ῗ SPP Ῡ῜ΐῥ῝Ί῎ῥ῍ 01/ †Ῥ Tῷ῵`῵῜wῼxῳῌ῱῝ +.- † SPP ῝Ῐ AP ῼ ῰ῶ (preseni- ῍+,῝ˆv*ῦῥ‰῾ Ύῒ‹*ῦΊ῎ῥ῍  lin) ῒ῍ῥ῍ ῦ SPP ῝Ῐῌ 3 (῝ῖ)* ῏Ῑ +*. ŒŽῒ AP ῝Ίῑ*ῦΊ῎ῥ῍ ῔ῑΰ ῦῌ +,῎ ῠῒ.*ῦΊ῎ῥῒῌ /0 ῭ῌ SPP ῝ ‘Žῡ’“ῌ Ὶ῟῞ῑΰῲ ῺῪ8ῒ•–*ῦΊΐῗs 45ΐ῜ῌ N 6ῒ7῔89.:;ῥ῍ SPP ῒῖῦ h AP ῌ Ῥ Tῷ῵`῵῜Ῠ˜ῑ - †῜῍Ὶῗ῍ =>῍ῥ῎ , ῟?Ῡ@AΊ+,Ῡ ;ῥB ῕š῜rsh AP ῝Ί›ῤœῗ CND.+,CDR+, Ίῌ ῼ ῰ῶ ῶC΅/ (nicastrin)ῌ aph-+ ῐ Nucellin, PCS+ ižῠ‰῾ ΎŸῒ ῔ῌ sh ῢG pen-, ῝῏H?Ῡ@AΊ+,῝῟ῥ῍ Jῗῌ K _ῐ AP Ῡ¡¢ῗ ῦΰῑŽt῜£¤¥¦§ῩOA; 8ῠ SPP  N 6ῒ7῔89.:;ῥῲ ῺῪ8 ῥ῝ῌ sh AP  - ‘rsh AP ῑŽt῝῞ῦ OῙ;ῥῒῌ ῼ ῰ῶ ῖ45ΐῲ ῺῪ8 ῗ ¨Ὺ ΅ῲῌῩ@AΊ῎ῥ῍ ªHῌ CND.+ ῩK8῝;ῥ῍ ῼ ῰ῶ P RΌTUῌῌVW Ῐ AP, nucellin Ῐ AP, CDR+ ῟῞῝«¬ῒ­;ῥ AP ῐ8῝῟ῥPX Tῴῲ ῺῪ8ῩZῤ\; ῝ῒ]ΰ tῒῬ Tῷ῵`῵ AP ®Ί῟Ώ°X ῌῩ«A; ῦΊ῎ῥῒῌ _ῐῐ῕ῥabῡ῝OῙΰῑ* ῥ῍ Jῗῌ JῚῗ῔±῎ῲTῼ AP ῜῍ῥ ῦΊ῎῟῎῍ efῌ SPP ῟ΰGῼ ῰ῶ Ῡ SPP ῘῑŽ (AtSPP, AtSPPL+-AtSPPL/) ῝ῼ ῰ῶ ;ῥ ῝῜ῌ _ῐgH AP ῝hijῌ *ΰ Ῐ AP ῠ²῎Ῐ*ῦῗ῍  . Ῡ²ῥ῝ῌ ῖ´ῑ _ῐ?ῳT῵X`nῩop῜ΐῢ῏῍ *ῦῥµ\ AP žῑŽῒ 1* .:;ῥ῍ · ῢ῏rsh AP tῌ uvwῼxῴB; ῌ b-῰Ὺ ῲῌ῱ (BACE) ῝­,῍ῥῑŽῠ. ῥῼ yz Ῥ΅ῩΊῌ ῝Ῡa|;ῥ῾῭῿ :ῌ efῌ ῦΰῑŽtῑΰῘ῏῟῝Ῡ ; ῌῲῌwῼxῴ€ῗΊ῎ῥ,ῒ‚῎῍ ῥῠῒ²῎Ῐ*ῦῥ,‚῎῍ _ῐ AP  ¸῟ JῘJῘ´ῌ ΐJῦΊ῎ῥ῍ ῍ῡῌ῟ῙῘ Ῐ Ὶ῝ῐΐ῔ AP῎ wῼxῳῌ῱„ῌῲ´ῌ΅ MEROPS (the pepti- Ὶ῏ (,**,). +) A. Mutlu & S. Gal : Physiol. Plant., +*/, /03 (+333). +2) M. Ramalho-Santos, J. Pissarra, P. Verissimo, S. Pereira, ,) S. Glathe, J. Kervinen, M. Nimtz, G. H. Li, G.J .Tobin, T. R. Salema, E. Pires & C. Faro : Planta, ,*-, ,*. (+331). D. Copeland, D. A. Ashford, A. Wlodawer & J. Costa : J. +3) M. Vieira, J. Pissarr, P. Verissimo, P. Castanheira, Y. Biol. Chem., ,1-, -+,-* (+332). Costa, E. Pires & C. Faro : Plant Mol. Biol., ./, /,3 (,**+). -) M. Ramalho-Santos, P. Verissimo, L. Cortes, B. Samyn, ,*) T. Asakura, H. Watanabe, K. Abe & S. Arai : Eur. J. J. Van Beeumen, E. Pires & C. Faro : Eur. J. Biochem., Biochem., ,-,, 11 (+33/). ,//, +-- (+332). ,+) T. Nakano, S. Murakami, T. Shoji, S. Yoshida, Y. Yama- .) P. C. White, M. C. Cordeiro, D. Arnold, P. E. Brodelius & da & F. Sato : Plant Cell, 3, 01- (+331). J. Kay : J. Biol. Chem., ,1., +002/ (+333). ,,) F. Chen & M. R. Foolad : Plant Mol. Biol., -/, 2,+ (+331). /) H. Park, I. Kusakabe, Y. Sakakibara & H. Kobayashi : ,-) X. Ge, C. Dietrich, M. Matsuno, G. Li, H. Berg & Y. Xia : Biosci. Biotechnol. Biochem., 0/, 1*, (,**+). EMBO Rep., 0, ,2, (,**/). 0) J. Kervinen, G. J. Tobin, J. Costa, D. S. Waugh, A. Wlo- ,.) Y. Xia, H. Suzuki, J. Borevitz, J. Blount, Z. Guo, K. Patel, dawer & A. Zdanov : EMBO J., +2, -3.1 (+333). R. A. Dixon & C. Lamb : EMBO J., ,-, 32* (,**.). 1) C. Frazao, I. Bento, J. Costa, C. M. Soares, P. Verissimo, ,/) T. Tamura, T. Asakura, T. Uemura, T. Ueda, K. Tera- C. Faro, E. Pires, J. Cooper & M. A. Carrondo : J. Biol. uchi, T. Misaka & K. Abe : FEBS J., ,1/, -. (,**2). Chem., ,1., ,103. (+333). ,0) Y. Kagaya, R. Toyoshima, R. Okuda, H. Usui, A. Yama- 2) T. Asakura, I. Matsumoto, J. Funaki, S. Arai & K. Abe : moto & T. Hattori : Plant Cell Physiol., .0, -33 (,**/). Eur. J. Biochem., ,01, /++/ (,***). ,1) P. Runeberg-Roos & M. Saarma : Plant J., +/, +-3 (+332). 3) T. Asakura, K. Abe & S. Arai : Biosci. Biotechnol. Bio- ,2) T. Tamura, K. Terauchi, T. Kiyosaki, T. Asakura, J. chem., /3, +13- (+33/). Funaki, I. Matsumoto, T. Misaka & K. Abe : J. Plant +*) K. Terauchi, T. Asakura, H. Ueda, T. Tamura, K. Ta- Physiol., +0., .1* (,**1). mura, I. Matsumoto, T. Misaka, I. H. Nishimura & K. ,3) M. Laloi, J. McCarthy, O. Morandi, C. Gysler & P. Bu- Abe : J. Plant Physiol., +0-, 2/0 (,**0). cheli : Planta, ,+/, 1/. (,**,). ++) A. Mutlu, J. E. Pfeil & S. Gal : Phytochemistry, .1, +./- -*) M. C. Cordeiro, Z. T. Xue, M. Pietrzak, M. S. Pais & P. E. (+332). Brodelius : Plant Mol. Biol., ,., 1-- (+33.). +,) P. Runeberg-Roos, J. Kervinen, V. Kovaleva, N. V. Raik- -+) N. Hiraiwa, M. Kondo, M. Nishimura & I. H. Nishimura : hel & S. Gal : Plant Physiol., +*/, -,+ (+33.). Eur. J. Biochem., ,.0, +-- (+331). +-) K. D’Hondt, D. Bosch, J. Van Damme, M. Goethals, J. -,) V. Buchanan-Wollaston & C. Ainsworth : Plant Mol. Vandekerckhove & E. Krebbers : J. Biol. Chem., ,02, Biol., --, 2,+ (+331). ,*22. (+33-). --) M. H. Cruz de Carvalho, A. d’Arcy-Lameta, H. R. Ma- +.) T. Asakura, H. Watanabe, K. Abe & S. Arai : J. Agric. cauley, M. Gareil, H. El Maarouf, A. T. Pham-Thi & Y. Food. Chem., ./, +*1* (+331). Z. Fodil : FEBS Lett., .3,, ,., (,**+). +/) S.B.P. Athauda, K. Matsumoto, S. Rajapakshe, M. Kuri- -.) T. Panavas, A. Pikula, P. D. Reid, B. Rubinstein & E. L. bayashi, M. Kojima, N. K-Yoshida, A. Iwamatsu, C. Walker : Plant Mol. Biol., .*, ,-1 (+333). Shibata, H. Inoue & K. Takahashi : Biochem. J., -2+, ,3/ -/) A. Schaller & C. A. Ryan : Plant Mol. Biol., -+, +*1- (,**.). (+330). +0) K. Terauchi, T. Asakura, N. Nishizawa, I. Matsumoto & -0) J. R. Mendieta, M. R. Pagano, F. F. Munoz, G. R. Daleo & K. Abe : Planta, ,+2, 3.1 (,**.). M. G. Guevara : Microbiology, +/,, ,*-3 (,**0). +1) X. Chen, J. E. Pfeil & S. Gal : Eur. J. Biochem., ,03, .01/

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