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ĈëÑĪ 1ƕ¼ď 21ȦưĈĐƧƠƦĤĐȵ649-6493 ëƋğƻǚ*ģĦȈț 9302ȶĈĐĈĐɃƧêǇĐǀǍǇ606-8502 ȶĬȶĦĥÔÓƴģȩÁƫ
The “sneak” preview of the Marchantia polymorpha genome
Key words: bryophyte, genetic redundancy, liverwort
Katsuyuki T. Yamato1, Takayuki Kohchi2
1

Department of Biotechnological Science
Kinokawa 649-6493, Japan2
Graduate School of Biostudies, Kyoto University
Kyoto 606-8502, Japan š zbſƠ&$Ůĳ=$ <üȵɆſƠZfSPɞMarchantia polymorpha L.ɟ*QixȎȘȮA% <=2%UEgeXe7Eh('*ȃČſƠ@+5&<ƃ(ſƠǊ*QixȎȘƐȤ=ɠɆſƠQix*(; Ŧ:)(!$ɠ
ſƠ*ɆȮÀ@ǰ <%ȹȉ(£ǭ) <üȵɆſƠ*Qixŀý+ɠ*&>ly`~KhSP*6*)ɂ:=$ <Ųǧȗ%+ɠZfSPQix*WfNqm{@Ȁ ɠɆſƠQix*(; @ǰę<5*š(ųş@Ŗ« nm/5+)"*7;_` ſƠ%Ä5$*¹QixƔĒ=*+UEgeXe% < (The Arabidopsis Genome
Initiative 2000) *ĴɠEh (International Rice Genome Sequencing Project 2005) ('*§Ơ*QixɔƊŦ:)=ɠ:)+ly`~KhSP (Rensing et al. 2008) & !zbſƠ*Qix)" $6ȎȘ=$ŲǋûǒŨƞ% 30 *ɆſƠǊ*Qixŀýºȿ=$ <ɞhttp://www.phytozome.net/ɠhttp://www.plantgdb.org/('ɟ&@ǰ <&ɠZfSP*Q
ixǀǍ+Ĵā@ōłå5( ɠIKh}Qix@æ4ī Łé%Qix@œþáɠſƠ) <QixǀǍ)$ZfSP+4>¶Ʉ@Â!$ &Ȋ<&6%< 1986 ĩɠZfSP (Ohyama et al. 1986) &]jR (Shinozaki et al. 1986) *ǼǨ¥Qix*¹ĀüȷÃƹƊ %ȎȘ===*TEX+Ǜ 120 kb &Ǜ 160 kb % ;ɠİŨŤ)Ŧ:

K. T. Yamato & T. Kohchi-1

BSJ- Revi ew 3: 71 ( 2012)

植物科学最前線 3: 72 ( 2012))(!$ lc*wcRd~CQix*Ǜ 16 kb (Anderson et al. 1981) 7oBVQix*Ǜ 50 kb (Sanger et al. 1982) &ƐȤ$6žƏ)Ĉ ǼǨ¥QixÄ*ȎȘ+ɠǼǨ¥
6"°*Ȳ¡Č%+(ɠȲ¡ŀý*¹¥µ)ƘƷƞ% ?8<QixǀǍ&âȌƞ)
ǎ!$ &ȏ < 1992 ĩɠſƠ&$+Ä5$ZfSP%wcRd~CQix*¹ĀüȷÃŦ:)(;(Oda et al. 1992) ɠZfSP+ǼǨ¥ 9-wcRd~CŢ*QixȎȘ=Ä*ƧƠǊ&(!*9 )ɠZfSP%+ſƠǟǲ)Ď÷< 3 Ǌ*Qix*

2 ǊūÄ)ȎȘ

=$ ;ɠZfSP@ƨ IKh}QixǀǍ+ſƠQixǀǍ*+;&£ǭ<&%< 2007 ĩɠZfSP6"ŻǸ¥*%+ū6ĜY ŻǸ¥ɞǛ 10 Mbɟ*QixŦ:)(! (Yamato et al. 2007ɠ大和勝幸 2009) =+ſƠĿŻǸ¥&$+Ä5$ɠÕŜ¥ƧƠ*ĿŻǸ¥&$6Ä5$ɠY ŻǸ¥&$6lc (Skaletsky et al. 2003)

9-^kV

(Kuroki et al. 2006) )Ɗ 3 ƮƷ% !
 2008 ĩɠZfSP*ƍ=QixɞĨŻǸ¥ 9- X ŻǸ¥ɟ*ȎȘċ2!=+ǗHhMƺ Joint Genome Institute (JGI)ɠȝ Monash ĈĐ* John Bowman :ɠȶĈĐ*ƕ
¼ď:@RC&<öɇ»âǀǍ% <(http://www.jgi.doe.gov/sequencing/why/99191.html)ƥ÷%+ɠȲ¡ĐǀǍŅ*Ŵ® :6Ì?;ɠ
Qixŀýüƶ*ŝ³)ä$ÏÌȫ$ < Ųǧȗ%+ɠZfSP*zbſƠ&$*ēɚüƶ01ǂǎ=$ <&@Ȣ2(河内孝之石崎公庸 2012) ɠzbſƠ&$*ZfSPQix*ơĹ@QixqVGNc*ȮŔƢƖ7Qix~[W&&6)Ǡ<om8&>12@'>A?!f/5+)*7;
 JGI %+ɠĚȜQix*üŲŀý@Ķ<5ɠĈȋƅUPUO@ȿċ<É)kE_cWP%UPUO@Ȁ!$ <ZfSP*þáɠɉż* P1 ƪŶĤŻǸ¥ɞPACɟ
}Ep}~ (Okada et al. 2000) 9;ȱA 30 N)" $ĀüȷÃ@ƔĒɠZfSPQix@Ȕ¬=:*kE_cN+ =6ĨŻǸ¥)ƪŶɠ20 N+ PAC}Ep}~:Ł)ȱ,=$ JGI %ȎŸ=ƍ;* 10 N)" $+ö¼*ǀǍǱà*ǵé)ü# $ȱ-ɠ==ÁŌ%ȎŸ= kE_cUPUO*ǣŹ@Ȃɡ)ǃðȎŸ PAC N*ETcTEX*áȐ+Ǜ 3.3 Mb % ;ɠ=+Ǜ 280 Mb &Ȋǌ6:=$ <QixTEX (Okada et al.
2000) *Ǜ 1%)ƹİ<ZfSPQix DNA * GC æȺ+ 40%ÉĴ% ;ɠĝ(&6ðȎŸ PAC Nɀ%+Ĉ(,:"+(!2ɠUPUO7CYp~O*Ɉĕ&(<ĈĜ*ÚĸȷÃ60&A'Ȋ:=(!cc}Wu\@ĺ&<ȣǈñČ6 1 N@Ʌ $ĝ(ɠ100 kb ) 1 °ǉĭ*ɖĭ% !*ǣŹ9;ɠZfSPQix*ȎȘ)ơƏ*óɎ+( &ÅŠ=ēɇɠǒǱ:6àN*U|_cKb]@CYp$3ɠćŜ*ȣǈñČ@æ4 1 N@Ʌ $+ǷĊ(ǣŹĶ:=$ <K. T. Yamato & T. Kohchi-2BSJ- Revi ew 3: 72 ( 2012)

植物科学最前線 3: 73 ( 2012)Ȃɡ kE_cUPUO*2&5 Ɗ)ɠ
b]rW)Ƴȼ=$ <CwiȸȷÃ)Ě<ə¢ĿƀǞ 9-ZfSP ESTɞȂɢɟ*v_nO@Ȁ ɠ]kNȟRdɕú@ŕƛćŜ*ȣǈñČ@æ4 1 N
Ć%+ȇŜ*]kNȟȲ¡ČȊ =ɠ*ėĭ+ 10 kb İ;Ǜ 0.7 % !=+UEgeXe* 10 kb İ; 2.3ɞTAIR10 )ü#ɟ9;¤ kE_cN*ɡ"% < pMM23-619A2 *Ȳ¡Čøô@ôɡ)ǃɠȲ¡Č*Áħ772,:% <&?<ZfSP*ǧȲ¡ČŜ+Ŧ:%+( ɠEST b]:+ 18,00020,000 % <&ŕƛ=<=+ɠkE_cN*b]:ŕƛ=<Ȳ¡ČėĭǛ 0.7/10 kbɞ=20,000 °/280 Mbɟ&9 Ǵ<K. T. Yamato & T. Kohchi-3

BSJ- Revi ew 3: 73 ( 2012)

植物科学最前線 3: 74 ( 2012)do=0A-J  8&>12@,A)?,?(]!#   We
Ŝč@Ɔǩ+ pMM23-619A2 *ȷÃ@ǃɠ*ƓǸ*ɕú) <t_NW+ŕĒHL[@ɠŕĒHL[*ÿ;".ȵÁ+Rdɕú@ǃƴǸ*ɕú) <t_NW+*ȷÃ)Ě<ə¢ɕú@ǃɦNCBI NRɠNCBI non-redundant protein sequencesɧAtha, Smol

9- PpatɠUEgeXeɠEgJ]lj 9-ly`~KhSP*Ȳ¡ČzbɧMpol ESTɠZfSP EST

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植物科学最前線 3: 75 ( 2012)

 kE_cN*ɠ4 N%áȐ 6 °* tRNA Ȳ¡ČȊ =ZfSPŽQix)Ď÷< tRNA Ȳ¡Č*ȕǟ)" $+Ĵ*ȎŸĳ=<2ɠȲ¡ČƲƥșǕ)Ɂ?< miRNA ('*¤ÁČ RNA )" $6ɠɆſƠ%®Ď=$ <6* ȵȊ=$ < (Floyd and Bowman 2004) ɠ¹Ė+Ŧ:%+( ( ɠZfSP* rRNA Ȳ¡Č+ɠ*ƼŽƧƠâƃN}W]@ıń$ ;ɠĨŻǸ¥) 9 ǔŅɠX ŻǸ¥) 1 ǔŅĎ÷<&ǃ=$ < (Fujisawa et al. 2003, Sone et al. 1999) pmkDKJ JGI )9!$Ŗ«=<Qixb]*ŝÃÒ7ɠąƯ¥*Ȳ¡Čv_nO@Ȁ 5ɠƄƜǙǥ Takaragaike-1 ż 9- Kitashirakawa-2 ż (河内孝之石崎公庸 2012) )Ȋ:=<ćù@Çƨ$Ȳ¡øô@§ńɞôɢɟ
*Ũɠȑ* 30 °*kE_cN@æ4 PAC N9;§ńćùvJ31 °ɠ$ZfSP ESTdCAPS (derived cleaved amplified polymorphic sequences)9-Ȳ¡Č*ȷÃ9;§ń

9- SSR (simple sequence repeats) v

J78 °@ƨ ɞÙȻĘë:ɠŰƲȂb]ɟáȐ 109 °*vJ+ɠZfSP*ĨŻǸ
¥Ŝ&â% < 8 °*ȭȽǮ)Ûŵɠ=:*¹Ⱦ+Ǜ 900 cM &(! Ȳ¡øô*ǘĭ@<5ɠƥ÷ JGI )$ Takaragaike-1 ż 9- Kitashirakawa-2 ż@ȷ$Ķ:= F1 Ɋò*ĈȋƅUPUOȮȀ% <:)ɠöǎȲ¡ĐǀǍŅ)$ɠ

ĚƟǙǥ% < Kitashirakawa-2 ż*ƊUPT)9<b]ÜĶ=ɠĴȨ< JGI *Qixb]&*ƐȤȮ5:=$ <Jo /5+)kDKJƄƜǙǥ Takaragaike-1 ż 9- Kitashirakawa-2 ż)Ȋ:=<ćù@ÇƨȲ¡øôvJ§ń=$ < ȵ*Ȳ¡Č)" $6øô)ǃ

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植物科学最前線 3: 76 ( 2012)qm3<92*7;
 JGI )9<ĈȋƅUPUO+ɠ) 2 Ǌə*Cq^%ēţ=$ <(? ɠTKƗ)9< Fosmid N*űǐȷÃƔĒɠ$ƊUPT Roche GS-FLX )9<¹QixU|_cK% <=2%)Ǜ 27x *QixJj_V@Ȱńɠ2011 ĩ 12 Ŭ)+ŪĒCYp~ ver. 0.6 Rw{faD¼ɂĒ%~~W=$ <CYp~ ver. 0.6 )<RaDOŜ+Ǜ 8,400 %ɠ*Ⱦ*áȐ+Ǜ 200 Mbɠ(? Qix*Ǜ 70ɝRaDO)9!$Jj=$ <&)(<2ɠȲ¡øô§ń)ƨ := 109 °*vJɠ 9-=2%)Öɍ$ < X ŻǸ¥ȭȽvJ9 °*¹$6v_q<&%:)ɠJGI %ÜĶ EST *Ǜ 90%6v_q%&:ɠðĶ:=CYp~+Qix*ĈÕ@Jj$ <&Ůĳ<&%< đ¹Ⱦ cDNA @ŪĒCYp~)v_nO³Ƶ(ȎŸ)9;ɠZfSP*]kNȟȲ¡Č* ǶƵ(ƂȬȊ $ɞȂɣɟ2ɠHL[EcƂȬ@Ȋ<&ɠȲ¡Čİ;*HL[Ŝ+UEgeXe9;ZfSP*Ţĝ( =)Ɂȭ$ɠZfSP*
HL[+UEgeXe*6*9;77Ⱦ ´ä@ǃ$ <ɠǒǱ:=2%ň
!$Ȳ¡Č@Ȋ<ɂ;ɠZfSP*Ȳ¡Č) <EcŒ¸ȵ£+ɠ*ɆſƠȲ¡Č%6®Ď=$ <*ũȪ% <ȕǟ+Ŧ% <ɠZfSP%+Ec@6( Ȳ¡Č*Êáć ßǳĿ6 < Ţ%ɠEc 9- 5’/3’ɐǯȓɕú (UTR) *Ⱦ
+UEgeXe*6*9;ɘǽ)Ĉɠ*ǣŹZfSP*Ȳ¡Č*¹Ⱦ+UEgeXe*6*9;Ĉ(<´ä) <=+ɠZfSPQix) <Ȳ¡ČėĭUEgeXeQix9;6¤ &* ñ&(!$ <Ƕ)ɠƼŽƧƠ mRNA * 5’²:Ȋ$ūÄ* AUG ȿċRd&(;ɠ:ǯȓȿċ=<ɠZfSP*Ȳ¡Č%+ɠƧƠǊI[O&*ƐȤ:ȿċRd% <&įǃí=< ATG *ƙ)ɠ,,ȇŜ* ATG Ȋ:=<ɞôɣɟ*9 (UTR-ATG )+ŲŶ*Ș3ź) <6*6( 6*6æ2=$ <&:ɠZfSP*ǯȓȅǭ+ƌ ȿċRd@ȱ.5*¦:*3@6"&ǰ :=<ƥŨƞ%+ɠƌ ȿċRdèȥ*»ȪƂȬ@æ5ɠ*3+Ŧ% <*&:ɠZfSP*Ȳ¡Č*ȿċRd@ȒĒ;ɠ5’UTR @æ5ı%ƧƠǊ)ě¸<ɇ)+ƘŁĻȉ% <    dp /5+) ,>&64.4kDLXjZh

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植物科学最前線 3: 77 ( 2012)

Jp "kDL Xj
*Ȳ¡Č*Ǒ 2 HL[)ɠƧƠǊ*I[O&*ə¢Ŀ:ŕĒ=<ȿċRdĎ÷<ɞƽØɟɠŕĒȿċRd*ƙ)+ 10 °* ATG Ď÷<rmGlQEkDLXR =2%)Ķ:=ZfSP EST

9-Qixŀý:Ȋ $&*%ɠzbſƠ&

$ :ū6ȹȉ"ɜËƵ(6*+ɠȲ¡Čȹȇ*ĝ(% <Ȳ¡Čȹȇ+ƧƠ*6"Ȳ¡Čkc~@ŎĈɠ9;ȇɌ%ćƃ(3@ƧƠ)ēȅ$*ÚɑɠȷÃ 9-ƈǳə¢ȇŜ*Ȳ¡Č*Ď÷+ɠ=:*ƈǳȎŸ*Ĉ(Ɉĕ&(<ª ,ɠſƠ*ȉ(·ÝĖ¥% <oDcNx+ɠUEgeXe)+ 5 ÁČǊĎ÷<ɠ== ) ȵƯ(<ƈǳ@ÁŌ""6ɠȹȇƵ(ƈǳ6Ȗ5:=<ɞôɤɟĵ!$ɠȹȇ<ƈǳ)" $+Ö ąƯ¥%+ȍę)ɠ=@ƾ<5)+ćȹąƯ¥@§ń($+(:((Strasser et al. 2010) 6ɠoDcNx&ƹ§ƨ<]kNȟ*Ȳ¡Č*ćȲ¡ČoBw~@ıń$ <5 (Leivar and Quail 2011) ɠoDcNx@
UOe¡ȰǙ+ȇɌ)ÃÒ$ ;ɠ*ȎŸóɎ&(!$ <*9 (Ȳ¡Čȹȇ+ɠȃČſƠ**Ȳ¡Č)" $6 ǶƵ)Ȋ:=<Ţɠly`~KhSP+ɠſƠ%î ɛÎƣ(ƹâǢŗ ßǳ(ēɚǙ% ; (Schaefer2001) ɠ*Qix6Ŧ:)=$ <&: (Rensing et al. 2008) ɠzbſƠ&$īƨ :=$ <ɠYə*Ǚǥ < +ly`~KhSP%*Ȳ¡Čȹȇ <(Rensing et al. 2007) ª ,oDcNxȲ¡Č+ȃČſƠ*6*&+Ư(<Ȳ¡ČoBw~@ıńɠ¾ȾĿɛ(!$ <ɞôɤɟ2ɠȃČſƠ%+Ö Ȳ¡Č% < LFY 6ɠly`~KhSP)+ 2 RnĎ÷< (荒木崇 2012) *5ɠȲ¡ČƈǳȎŸ)Ɂ$+ɠüȵɆſƠ% <ly`~KhSP)$6UEgeXe&âƃ*ïɗ@Ŋ $ <&ȏ< =)ĚɠZfSP)Ď÷<oDcNx+ɠȃČſƠoDcNxoBw~ÁĠ<É)ÁĠ&Ȋ:=<ɡÁČǊ*3% <ɞôɤɟ:)ɠoDcNx@UOe¡Ȱ%ĺƵ(ĲÊ@Ź&ǰ :=$ < PHYTOCHROME INTERACTIG FACTOR(PIF) +ɠZfSP%+ 1 ÁČǊȊ"!$ :ɠȲ¡Čǁă<&·ĽǓƯĨ&(<ɞ¨Ǆ:ɠŰƲȂb]ɟ
ȃČſƠ7ly`~KhSP%+UOe¡Ȱ7ıŃıń)Ɂ?
<Ȳ¡Č*ćȲ¡ČoBw~@ıń$ <ɠZfSPQix)+*ćÖ Ȳ¡

K. T. Yamato & T. Kohchi-7BSJ- Revi ew 3: 77 ( 2012)

植物科学最前線 3: 78 ( 2012)Č&$Ď÷$ <: ɞȂɤ) ȵ@ǃɟ:)ɠÁȄǢǫ*Ǧŏ7ıŃıń)ȹȉ(ĲÊ@Ź CLE Ȳ¡Č+ɠȃČſƠ%+ 32 Ȳ¡Č:(<Ȳ¡ČoBw~@ıń$ <ɠZfSP)+==*Ǚǥ*üȵ%ÁĠI[OɡȲ¡Č"Ȋ"!$ <(澤進一郎 2012) âƃ*´äɠ
*ÈķǙȲ¡Č)" $ýç=$ < (Sasaki et al. 2007) =+ɠÖǜ(¥È@6"ZfSPɆſƠ)»Ȫ)Ȋ:=<3@³ ""6ɠ*
3@Ś <Ȳ¡ČƂńƁ5$ÖǜɠüŲƵ% <ßǳĿ@ǃ"2;ɠZfSP@zb&<&%ɠȃČſƠ)Ȋ:=<ÈķǙ*ƈǳƵȹȇ6+ÃÒ)9<ȇɌ@ðȳ""ɠɆſƠ)»Ȫ<üŲƵ(3@ȎŦ%<ßǳĿ <ƹâǢŗ )9<Ȳ¡Čǁăßǳ)(!ƥ÷ (河内孝之石崎公庸 2012) ɠȲ¡Ƶ¾ȾĿ*¤+ɠZfSP*zbſƠ&$*Ĉ(į3% <Jq 9%2'>;kDLHLabY

smQUcF ZfSP+ɋɉƯżſƠ% ;ɠĨŻǸ¥ 8 Ų)Ì ɠĿŻǸ¥&$ɋż%+ X ŻǸ¥ɞn =8 + Xɟɠɉż%+ Y ŻǸ¥ɞn = 8 + Yɟ@6"Y ŻǸ¥*ĀüȷÃ& X ŻǸ¥*ȵÁȷÃ*ƐȤ:ɠZfSP Y ŻǸ¥6ɠlc7^kV* Y ŻǸ¥&âƃ)ĨŻǸ¥:ÁÒ&ǰ :=< (Yamato et al. 2007) Y ŻǸ¥*Ȳ¡Čėĭ+ 0.1 / 10 kb ǉĭ% ;ɠĨŻǸ¥*ŕ
Ē± 0.7 &Ɛ/$Ŧ:)¤ =+ɠX ŻǸ¥&*Ǣŗ ŉÈ=&% ȵ*Ȳ¡ČƉĉɠ:)cc}Wu\7ÚĸȷÃǾǌ5&ǰ :=<( ɠÕŜ¥

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BSJ- Revi ew 3: 78 ( 2012)

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