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Liverwort, Marchantia polymorpha L., as a reviving model for plant biology
Key words: basal land plant, genomics, land plant evolution, Marchantia polymorpha, molecular
genetics
Takayuki Kohchi and Kimitsune Ishizaki
Graduate School of Biostudies, Kyoto University
Kyoto 606-8502, Japan[gTQ+̄19 ɐŭí)ƭ)ȠȘ˅ȼ ʟɖ)ʘŃ>̄ǒȌĴȱɀ*{cǒȌ%!
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˸9ajTQ˸&&8)SQǒȌ&$̅")4&7;>=&8= ̄]F˸̄Z˸̄ajTQ˸+ 4 ÄŞ¤Ü)ÔŐ$=ˣǒȌ*ğˎ*ɏɛ+44(ʨʥ*°ę+=8**̄ǐˋ¨ı9ʅɡ®ˋ¨ı*ːÖ9ǖʿ*Ôıɏɛʙǈ&!Ȟ(=ƊǮAɘ5ā@ɠāȢ(Ôıɏɛʙǈ ;̄]F˸+SQǒȌ*( %8ƹ8þÔŐ̄ˣǒȌ̂ǜȴ)+ƻɴǒȌ̃*ğˎ)¬ɩ=& ȵ>$=̂Qiu et al. 2006̃
[gTQ*Ô˸*¬ɩ#*ʟɖ)"$+̄ŕǂ̂2012̃A÷ȅ>*˂èȢ(¬ɩ#+̄[gTQ*˓ʔ(˿ß*-&"%=˂è)+ˋ¨ı*Ȓů̄ďį̄ǚɶĩƞA© ̄ğǀȢ)ˋ¨ı+ȷÇ ;ýɜ8*%<̄
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植物科学最前線 3: 59 ( 2012)]F˸AĻ˼ǁƩ&Ôıˋ¨ĴȢ(ʙǈ+̄ˣǒȌƅɂ*Ȗʙ*˗&(=&ƽŬ>=ŋ̄ȔȘ*[gTQ+̄ʏıǒȌ*ȷÇ)ũ=ʝ%+(&)+ǯſ Ŵʔ%=

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£%=[gTQ*ȠȘ˅ȼ)"$+̄Knŷ1890̃ Ǘ(ǒȌ*ĕʧ* *̅ɃA[gTQ)$$Óȋ>;*Ɏɣ(F~Xd+̄ȔĘ WEB UFd%8ȥ)=& % =̂http://www.geheugenvannederland.nl/; Marchantia %Ǔɕ̃
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>> ȗĥŶɇAȵ7̄ī*ȱɀ%ːÁ®£Ańʪ)=& % =ːÁ®£ÅñȢ(SQǒȌ&ɵı®£ ÅñȢ(ˣǒȌ*ÚŰǚǖ*ǧʵ+̄ǐȧ&ȘĊȔʪ*˝¸&!Ȃ%ƴ˄ź&īǗźAȮ=%˯Ŝ)ɻĈǵȱɀńʪ&(=ʚƞ =&̄SQǒȌ&ɞɉǃǒȌAǧʵ=&):!$̄ɵı®£*ÚŰǚǖ ːÁ®£%ƅɂÚŰǚǖAʳș*? &!č )ɇ ů;>=̂Nishiyama et al. 2003, Sano et al. 2005, Sakakibara et al.,2008 (')

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BSJ- Revi ew 3: 59 ( 2012)

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Ʈ˙)á $̄­ǹ8ȧʳƞA¹˂=&8Ġć>$=̂Lloyd & Steinmetz 1937̃ũƲ*Ļ˼+˔Ī*L~Xļ9ȡȈˬȕAÈǻ&$ǒȌ ǑĞ>$(( ;̄Ɔ Ļ˼ļ%[gTQAǑĞ=&˙ƮǄ¥9­ǹǄ¥)$8ƅ˙ȧʳƞ ʘŃ>( !*ŭ̄ʋÈȁAÈǻ)$=& ũƲ*Ļ˼&*Ĭ (ˇ%<̄ˈʰɾÈAʑÈ=&%ȧʳƞ ¹˂>=& @ !>):!$̄Ļ˼ļȗĥ%£Ađ& ÿɶ&(!̂Chiyoda et al. 2008̃Șǣƅ˙ȧ)Ȼʌ=&̄˧Ǐ%+ʿɎĐ̄˩Ǐ%+ʿòĐ Ūƅ>=ʿɎĐ ;ƣÓ>Ɏı ʿòĐÎ*òɖɵ)ǰȬ&̄ýɎ ʱ<̄ɵı® Ūƅ>=
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[gTQ*Ïźɶ*˾+̄ŪʯʳƞĻ˼̂ŭʺ̃)8Ș >$=ɼȢ)ǒȌ*Ūʯʳƞ%+̄ɖɵAKX&$Ħǣ DNA AňÉ̄v{*ʤɊ):!$ÏÔè=&Ğ˻ƍʍ Ŵʔ&>=>)ń$̄[gTQ+ȍǡ(ǒȌv{ÒȖA=&(̄ʈÝɯźAȒůɖɵ ;ǒȌ®Aľư)ÏȘ=& % =z-~y! q+=PE
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̄ȠȘ˅ȼ9ŪƂĴȢʘŃ9ɁȄĩȞ®('AīƦû˖ˋ¨ʙǈ*ɚǉ8û˖>$ <̄ɑ 70 ŞÜ)[gTQ*ȱɀ ȣB)ʌ@>$& @ =ɻĈǵʎȔĜAȵɁȄĩȞ®*ʜʺ8ī ̄Ǣŷ( ;Ļ˼ǁƩ&$ȔĘÉƊ=&+Ĕ˫%=
1970 Ş£̄ȆǀĬĴ) $[gTQĞ˻ɖɵ Ǚɂ>̂Ono 1976̃
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Rjy*ʙǈ)Ùș>̂Ohyama et al. 1986, Oda et al. 1992̃
 ̄Ğ˻ɖɵ*Rjy)+Ğ˻):=ǌɾ®t%*ĩè =&9̄ȔĘ*ǘǼɏɛ ;Ǭĸ8*&ǧʵ=&JLi~Rjy)8ʆīĜ ĲĘ=& Ư; )(!$=̂Ĭċ̄ȰŔ̄ǭÎ ƿȠʎc]̃
ŭ̄*[gTQĞ˻ɖɵAĻ˼)Ùș=ġā)+ʜ*Ȃ)ǯſ Ŵʔ%= Ôıˋ¨Ĵȱɀ%+Ļ˼ǁƩ)ǘǼɏɛAĸ7=&):!$̄ȃș(ǶAˌ=& ˓ʗ>$ ȔĘ̄ȱɀș*ǘǼǏ&$ˏśĺǫęê%ƛ˨ɏɛ̄Takaragaike-1̂˧Ǐ̃&
Takaragaike-2̂˩Ǐ̃ Ùș>$= (Okada et al. 2000)>;*ɏɛ+̄Marchantia polymorpha ssp.
ruderaliŝ "$*Ȑɬ* Marchantia polymorphã)Ŏ=Takaragaike-1 Ǐ :. Takaragaike-2 Ǐțǅ* PAC OAș$Rjy~Fq~ ǖɋ>̂Okada et al. 2000̃̄źǌɾ®*ǖʿʙǈʌ@>̂Okada et al. 2001, Ishizaki et al. 2002, Yamato et al. 2007̃ȔĘ̄>;*ɏɛ+̄Rjyr
WHOdA+7̄44(ȱɀ%˔ȘĜǘǼǏ&$ƛș>$=SQǒȌ*ġā̄ːÁ®£%ƛü=7̄ƛüǤˤ%ƭ)ɒɏ%=&ʚ = ̄Takaragaike-1:. Takaragaike-2*˜)+̄źǌɾ®*ˇ%(Ŝǌɾ®)8ĤğīĜ ĲĘ%̄2006 Ş ;ƈːA˂7̄Takaragaike-1 țǅ*Ŝǌɾ®& Takaragaike-2 *˩źǌɾ®A8"˩ǏAůɍėIi
NȨ Joint Genome Institute ) =Sx|gbErP~y)ƛƐ>Rjyʙʢ)+ƈːɏɛ ș;>$=̂http://www.jgi.doe.gov/sequencing/why/99191.html, Ĭċ 2012̃ 4̄ˋ¨ęĕ±ƅ9ˋ¨ıw`oP*7*ńȅɏɛ&$̄ˏĬĴ*ǖÎ ; Kitashirakawa
ǏAƛ˨Takaragaike-1 Ǐ&*˜*īĜAÙș$̄8 ǀ*Ŝǌɾ®Aʎ 8 "*ˀ˘ɫ ;(=ˋ¨ęĕA±ƅȔĘ̄RjyȌȖęĕ&ˋ¨ęĕ*ɛāA˂7$=w`oP) $
8̄íƦ®*ÙȂ =ːŭů;>= F1 ɵı)țǅ=ǒȌ®+̄) Recombinant Inbred ɏɛ*:(w`oP˨ē&$Ùșÿɶ%=ːÁ®£%+sbƜāź8ĲĘ(7̄ˋ¨Ȣ(w`oP+ǔ7$ľư%=

|yq+=?/Fl|-zy% #)N! BnoG
ǒȌ*ŪʯʳƞǮ*( %8̄DPlObD):=Ūʯʳƞ+åȓȢ "Ɍ·%=ũ×+
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9 4. % #!)BnoG
DPlObDAș[gTQ*Ūʯʳƞ+Ĕ˫%!Ļ˼ļ) $ǧʵȢűȘȌǶÉ*Ŋ(ɵı*ƛ˨ ÿɶ&(!&! &(<̄Ūʯʳƞ ľư)(!ɵı ;Ğ˻[gTQ Sporelings +ȣB)Ôʐ$ <̄DPlObD):=Ūʯʳƞ)ˉǁƩ%=̂Ishizaki et al. 2008̃Ǵ®Ğę*( %ɵıAȠɿ̄5 Ʈȥ* Sporelings AǼÂ=)DPlObDAá ̄2 Ʈ˜*ÌĲĞ˻):!$Ɓǌ=ǀǅ̄[gTQ+DPlObD*
ƁǌĿ%+( ̄DZdVT*Ƿá):!$åȓȢ(Ɓǌ ÿɶ&(=ÌĲĞ˻ŭ̄Ǳǳ):!$DPlObDAˢö̄ˊƏș*ʈÝAą6ˊƏĞę%[gTQAĞ˻=*ǔ7$ïɒ%ľư(Ơ±):!$̄īƦ*Ūʯʳƞ® ů;>=̂ĕ 4̃Ɔ*ȱɀļ%+ 1 ɵıƶ; 1,000 ʸƦ*Ūʯʳƞ®Aů$=ü<ƌɵıƦAĦ9,̄XQD`r8ÿɶ%
=ŪʯʳƞĻ˼Ǯ*ʟɖ(ʙʡ+ȰŔ ǭÎ̂2008̃A÷ȅ>
[gTQ* Sporelings AșŪʯʳƞ åȓȢ%=&9̄íƦ®£Ańʪ&$=&Aǲ $̄T-DNA AĩȞô :.ĩȞ*]P&$Ùș=& ÿɶ&(![gTQ*Rj
yUFȲŝĚȢˋ¨ıUFY :. T-DNA *ƙÉƦA8&)ʞɈ&?̄ÊlFDX(T-DNAƙÉ>=ġā̄ɑ 70,000 *ȏɂŪʯʳƞ®%1$*ˋ¨ı ť+Ȳħ>=&ž>4̄90%*ƽŬ¿%Êˋ¨ıAɟɪ=)+̄ɑ 20 ɏɛ*Ūʯʳƞ¼®Aƌ&)(=&ʛɈ>>4%)̄ŪƂȢ(ĩȞ®AʓƦÔ˪̄*( *{cQX&$ǨļA4!Ūƅ(ĩȞ®Aˊ.̄
*ôĒˋ¨ıA T-DNA A]P&$Ăĸ̂ĦȚƳȸ̄
ǩʩƿɰ̄ȰŔËŦ ƿȠʎc]̃
4̄[gTQ ˾Ȁť*JMV%ǊǠ=&AÙș$̄JMV­ƁýźĩȞ®*XOgPAʌ!Ș Ǣ=8*AˊƏ=ŨÚȢ(XOgPƬǮ%>,̄-&<*ȱɀɮ%8ǧʵȢľư)ɟɪȢ( T-DNA ĩȞ®XOgP ÿ
ɶ%=ʛɈ 20 Ūʯʳƞ®A±ƅ̄˾ȀťJMVąƻĞę%ȘĲ=ĩȞ®A 10 ƦɏɛüůTAIL-PCR Aʌ!&?̄ʳÑǲźè]Fr*JMVŶɇʳÑĒı̂ARF+̃ˋ¨ı ȏɂ)Ȳħ> 2 Ǐ)á $̄ɻĈǵˋ¨ı)]P É!Ǘ(ɏɛ ů;>̂ǝȚȪțı̄ȰŔËŦ; ƿȠʎc]̃
Sporelings AǁƩ&=Ūʯʳƞ+åȓȢ%= ̄ȍĸ*ˋ¨ȢɳƵA8"ɏɛ);)Ūʯʳƞ
=)+̄ɵıAǼÂ=7*Ʋ˜Aʔ=&!ʣ˶ !]lSŪʯʳƞ Õƪʅ
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植物科学最前線 3: 65 ( 2012)AÙș=:)[gTQʅȎ®AÕƪ$%+̄åȓȢ(DPlObDŪʯʳƞ+ʱ (!ÈŶɇ*ȱɀ ;ʅȎ®*Õƪ˰) =ÏȘ˅ȼ%+̄È¶ĲȢ)ɖɵÔʐAʠň% =& @ !*Ǥˤ*ɖɵ+Ôʐ ǲȠ%=&)ǯȥ̄ÏȘ*ʅȎ®Ańʪ)DPlObD*ƁǌAʌ!ÕƪÒȖŭ̄ƯǄ¥%ƦƮĞ˻$Ôʐ*ǲȠ(ɖɵAą6ʅȎ®AǁƩ)=&%ĵĸ$Ūʯʳƞ® ů;>ɵıA³!ƬǮ&ǧʵ=&̄ȏɂŪʯʳƞɏɛAīƦ±ƅ=)+Ą %=8**̄Ʀë*Ūʯʳƞ® ů;>>,::(ī*Ļ˼)Ùș% =ĻșȢ(ƬǮ%=̂ºȚʂ̄ȰŔËŦ̄ƿȠʎ̃ȔƲȂ%̄ƹ×)ÙșkFP
wFVAą7$̇"*ˊƏwK [gTQŪʯʳƞ)Ùșÿɶ%=̂ȚĻ̄ȰŔËŦ̄
Sandy Floyd ; ƿȠʎ̃˅ÞȠȔǏ*±ƅ9ȧʑźǓĸAʌġā̄=+ȍĸ*ɁȄĩȞ®9Ūʯʳƞ®)ˋ¨ıAňÉ=ġā*ƬǮ&$ƻș%=
DPlObDA¡[gTQ*Ūʯʳƞũ£*¼®) $+̄ɖɵ Mz~ȎƂ%ĲĘ=&8=:%=ʽŜ*ǒȌŪʯʳƞ%+̄ː):!$ȾıAů$̄Ūʯʳƞũ£* T1; T2̄T3 &£Ađ$̄ʙǈ)ˉv{Ɯā®ɏɛAȴɂ=[gTQ%+̄íƦ®%=&&ȃźɿ ̅ɖɵ)țǅ=O%=7̄T1 ¼® ;ȃźɿAÔ˪$̂ôÛȢ)&"* T1;̅ɏɛÔ˪=̃̄ɒɏAȴɂ% =>+̄ȃźɿ̂gemmã*Ʌ£&

&%̄G1 &ĉB%=̂·Ĺ̄£&ĉ/ ǸƦÔʐ+ə$(&)ǯȥ̃G1 £+ɏɛ&$ɞƔ=&&8)̄-&"* G1 ¼® ;ů;>= G2 £AĚ(ǁƩ&$Ļ˼)µ$
=̂ĕ 5̃
ȃźɿA¡=Ħǣ+ʷʾ%=7̄ȯƽ˜)ˋ¨Ȣ(ȎƂAɞƔ44%ʙǈÿɶ(ȎƂ&(=[gTQ+̄ǧʵȢľư)ɒɏ&$*Ūʯʳƞ®Aů=&% =}gO(Ļ˼ɏ%=&ʚ =

9 5. TDj dc `b

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植物科学最前線 3: 66 ( 2012)|-{y^8eG q+=_; ʼˋ¨ĴȢ(ˋ¨ıǚɶʙǈ)+̄ȧĂɘƞ AÙșˋ¨ı]RbEP Ũß(ƊǤ&(=nzaLiTQ+̄˾˵ť%ȧĂɘƞ Aʱ& Ȯ;>$= ̄[gTQ)+ũ$+4;(:%=> Z˸&]F˸*ˇ&ɼè% = +Ư%=˺Ț;+FiAǁƩ)åȓȢ(ˋ¨ıȲħɏɛ*ˊƏƬǮA˛Ƞ̂Terada et al. 2007̃
>+̄ʈÝɯź*ˊƏwKˋ¨ıĂ"*ȧĂźˎÔ%Ɨ5̄;)ĪÀ)ʫ*ˊƐwK̂*ġā+̄̅ɖɵǦ%= DTA ˋ¨ı̃Aːɩ T-DNA SXd~OdAș=&):<̄ʽŜ* T-DNA *~
^y(ƙÉ):=ŪʯʳƞAƚˢ=&8*%=ǧʵRjyʙǈ)ȧĂźɘƞ +Ŵ˳*a
&ɭ ;>*%̄*ƊǮA[gTQ)Ŷș2011 Ş*í,)̄ƹ×*ƅà´ Ó$ ;̄
*ƊǮ):!$ȔĘ4%)ɑ 20 ˋ¨ı Ȳħ>$=̂ȰŔ; ƿȠʎ̃
PCR ǮAș$ȥȢ*Ūʯʳƞ®AĂĸ=Ŵʔ = ̄ů;>ʈÝɯźǏ*ɑ 2́ ǜˋ¨ıȲħ>8*%=ƊǮȢ)+ƢĎ*°ę+=8**̄200 ¼®2'*¾ʑǏ:< DNA AƒÓ$ PCR Aʌ,̄23ȴĻ)ˋ¨ıȲħǏAĂĸ% =:)(!|-|ykg-! BnoG ǒȌɖɵ)+ǐ*¢)̄ʅɡ®&xdSeD)Rjy ĲĘ=ʅɡ®Rjy+̄ǒȌɖɵ*Èāƅǚɶ)˓ʔ(ˋ¨ıAų) 120 4<*ˋ¨ı Se>=Èāƅ) $+̄ǐR
jy&ʅɡ®Rjy*îʤȢ(ȠȔ ˓ʔ%=ɼȢ)̄ʅɡ® DNA %*ˋ¨ıȲħ+̄ȧĂɘƞ AÙș$=O~xe{fXʅɡ®%ĩȞAȧʑ=&!ˊƏƬǮ%Ġć>
̂Boynton et al. 1984̃ȔĘ%+̄]lS̄nzaLiTQ(') $̄XuO_jwFVɯźˋ¨ı&!ˊƏwKˋ¨ıAƗ6Ū*ȧĂ˴ĝA SXd~OdAmbEOLǮ('*ȌȖȢ(ƊǮ%ɖɵÎ*ʅɡ®)ňÉ$ˊƏ=ƬǮ ȴɂ$={cǒȌVF
hfYf%+ʅɡ®Ūʯʳƞ+ȴɂ$(ĂǗ*ƊǮA[gTQĞ˻ɖɵ :.ǒȌ®%Ŷș&?̄Ūʯʳƞ®AǧʵȢľư)ů=&)ƅà̂Chiyoda et al. 2007, Chiyoda et al. 2008, ì£Ț ǭÎ2008̃Ļ˥)̄
*ƬǮAș$ʅɡ®ˋ¨ı*ǚɶʙǈ ʌ@>$=̂Ueda et al. 2012̃ʅɡ® DNA +īSo%=7̄1$*SoAȲħ DNA )ɩ ƞ v{r~Xdx`O(ɏɛ±ƅ)Ʋ˜Aʔ=* ɼȢ%= [gTQ%+̄Sporelings Ańʪ&$ï˪Ūʯʳƞ® ƹ× ;v{r~Yx`O%=ġā8!̂Chiyoda et al. 2008̃Ô˪ũ×+sb(ȎƂ%!$8ȃźɿ*Ğ˻Aə=&%v{r~Xdx`O(~FAʷʾ)Ô˪=& ÿɶ%=̂̀Î; ȹ»̃}yaU<[- 1u VFhfYf%+īƦ*ɁȄĩȞ® Ô˪>̄Ǘ(ȘĊȔʪ*Ȗʙ)Ĭ ʬȑɁȄĩȞ®*Ô˪+̄ȘĊȔʪ*Ȗʙ)˓ʔ(ƬǮ%=nzaLiTQ* EST ʙǈ ;̄ˣǒȌ+ƅɂũ× ;44(Ⱦ˸*ˋ¨ıAºƔ$=& ȵ>$=̂Nishiyama et al. 2003̃
>+ʸŞ*nzaLiTQ9FhK]nl&!ğˎǒȌ*Rjyʙǈ ;8ơƔ>$=̂Rensing et al. 2008, Banks et al. 2011̃
[gTQ8ʏıǒȌ 8"ĐķA8(8**̄ˋ¨ıt%+ğǀȢ(ˋ¨ıZ`dAºƔ$=& ȵ>$=̂Ĭċçş 2012̃
4̄[gTQ
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植物科学最前線 3: 67 ( 2012)+˂è*˅ȼ%ǌɾ®t%*½èAə˼& (&ž>̄ÚŰĒı) =ˋ¨Ȣ(Ð˙ź nzaLiTQ&ǧ1$8­& ȵ>$=*ÃĄ+ȍ)ÚŰɏˋ¨ı%˷ʆ
%="4<̄[gTQ+þÍȢ(ɁȄĩȞ®*Ô˪)á $̄Rjyʙǈ ;8Ôıˋ¨Ĵ)ˉ
ǁƩ%=& ȵČ>$=VFhfYf%+ˋ¨ı*Ð˙ź*7ïĩȞ®*ʎȔĜ Ưȭ%(´8īƦĠć>$=´ ,̄ÈŪƂŪƅ) =ʫ*ʳÑĒı%= PIF pCx+ʓƦ*ˋ¨ı Ð˙Ȣ)±ș=̂Leivar et al. 2008̃>)ń$̄[gTQ* PIF Ǘˋ¨ı+ïSo%<̄ˋ¨ıAȲħ=&ǜŜ(ÈŪƂŪƅ ʱ;(̂²ȶ; ƿȠʎ̃*´ ȵ:)̄ğǀȢ(ȘĊȔʪ*Ôıǚǖ*Ȗʙ)[gTQ*ɁȄĩȞ® ūɂ"ÿɶź =Üʺ* Burgeff *ƅƷ (1943))+̄ũƲƭ)[gTQŎ*ɁȄĩȞ® Ô˪>$ʙǈ>$=& Ġć>$=ɻĈǵŪƂAȵ8*9ˋ¨ȢɳƵ*8*8ī>;+ʭ˓(ĩȞ®
%= ̄ȔĘ%+ÉƊɶ%=&&˔Ī ;Ô˪>7ˋ¨ȢɳƵ Ư%=* Ǣŷ(&?%= ȔĘ̄Ǆ¥Ȣ)+ƢĎ*°ę+= ̄[gTQ*ɵıALwɢ9 EMS ('*ĩȞô):!$ÒȖ$ĩȞ®Aï˪=& ÿɶ%=̂ȰŔËŦ; ƿȠʎ̃[gTQAș=ƹĬ*ÙȂ+̄Üʺ*ˋ¨Ȣ(Ð˙ź ­&)á $̄ʽŜƌːÁ®£ íƦ®%<̄Åź9âź&!
êØ(ɁȄĩȞ® ũ£%Ô˪% =&%=>4%)̄JMV*Ɓýź ­ĩȞ®9˭ɾÈŌź ­ĩȞ®*Ô˪)ƅà$=ÈŌź*ĩȞ®*( )+̄˭ɾÈýľ® phot )ĩȞA8"8* ą4>$̂ŉǇƀ; ƿȠʎ̃Ūʯʳƞ*˲%ʺ1:)̄D
PlObDA¡=Ūʯʳƞ%ƙÉ>= T-DNA AĩȞ]P&$ĩȞ®AÔ˪=&8ÿ
ɶ&(!$= ŭ+̄īƦ*ĩȞ®Aû˨$̄Ļ˼)ɹț)µ=®ÚA±=& ˓ʔ%=~y@M
 [gTQ)+ˋ¨Ĵȱɀ*˙ǟĀ = ̄Ȕ£Ȣ(Ôıˋ¨Ĵȱɀ+İ4!, <%=
[gTQ Ȫ)Å>{cȘȌ&(=)+̄" *ʣ˶ = ǁƩ&$*ƌ+̄>>*ȘȌȾ)Ėƻ*č˶ =[gTQ+˩˧ȞǏ%=7̄ĩȞʎȔĜʘŃA=˥)+źŚ ¼®Ś)ǯſ Ŵʔ%=4̄[gTQɘɧ+VFhfYf)ǧ1$ȳ̄44(èāȌAʇȿ$=7̄[gTQ ;Ș®ÔıAï˪$ʙǈ=7)+ŘĮ Ŵʔ%=
4̄ɖɵȘȌĴȢ(ʘŃ+ƅǉ  <""= ̂żɽ Ț 2012̃
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 [gTQ*ȠȘ)˝=þÍȢ(ʜʺ+ÆĻ$= ̄ȠȘ*ƕǘ&(=wKˋ¨ı+ʲ$ <̄ɖɵ9Ôı)"$ȬȥȠȘɏʧ*Ȗʙ ʲ$= ŭ*˂èȠȘĴȢ(ȱɀ)Ĭ)ƽŬ̂Ǿ; 2012, ʃƾ 2012̃
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