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植物科学最前線 3: 58 ( 2012)6 K"$OV w WCǭÎĳ ȰŔËŦ
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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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T. Kohchi & K. Ishizaki-1

BSJ- Revi ew 3: 58 ( 2012)

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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T. Kohchi & K. Ishizaki-2

BSJ- Revi ew 3: 59 ( 2012)

植物科学最前線 3: 60 ( 2012)Ļ˼ȘȌĴ*ĻʯȢ(Ȃ%8̄ˑǥ('*´AƖ=4%8(̄ɁȄĩȞ®AƌÔıˋ¨Ĵȱɀ) $̄íƦ®Ańʪ&=ſɬ+Ĭ íƦ®£%+̄âźĩȞ%!$8ĩȞʎȔĜũ£%ʘŃ% =&ÙȂ =

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BSJ- Revi ew 3: 60 ( 2012)

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BSJ- Revi ew 3: 61 ( 2012)

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ɖɵÔʐAʘŃ=&!Ļ˼ ÿɶ%=ī*SQǒȌĂǗ)̄[gTQ+ȃźɿ&}gO(ĦǣŪƂA8"[gTQ*ȃźɿ+̄ăÜ*ʽ<K`rǖʿAǆȎ®*ţˎ)̄ƻźȘǣAə=&()Ūƅ>=̂ŕǂ 2012̃ǆȎ®*ȠȘ˅ȼAĕ 3 )ȵȃźɿ+ţˎ*ɖɵAʱǻ&$̄ɖɵª˙̄ŏÔʐ):=ğˎ*ǋɖɵ&ÇɄÀɖɵ0*Ôʐ̄ÇɄÀɖɵ ˀ*ɖɵÔʐAɨ<ʹ̄īɖɵè=&%Ūƅ
>=%ȍ)ǯȥ&+̄¼*ȃźɿ+ǥ®*ʅȎ®&Ăˋ¨ıĜA8"&̄(%8̅ɖɵ)țǅ=O%=&&%=ȃźɿ+Ćǩ(>,¦ȫ=źʯ =*%̄ȠȘǤˤ*Ɲ!Ě(ǁƩAʤʒ=*)Ùșÿɶ%=ȃźɿ+åȓȢ(ĦǣŪƂ%=&&8)̄Ⱦ*¦ȫȎƂ)=7XdX)ń=ƑƎź8˾Ů)ȉȃźɿ9̄łĭĞę)ºĲȃźɿ+Ŋ(&8ƦŞ˜*ºĲ ÿɶ%=
[gTQ*Ïźɶ*˾+̄ŪʯʳƞĻ˼̂ŭʺ̃)8Ș >$=ɼȢ)ǒȌ*Ūʯʳƞ%+̄ɖɵAKX&$Ħǣ DNA AňÉ̄v{*ʤɊ):!$ÏÔè=&Ğ˻ƍʍ Ŵʔ&>=>)ń$̄[gTQ+ȍǡ(ǒȌv{ÒȖA=&(̄ʈÝɯźAȒůɖɵ ;ǒȌ®Aľư)ÏȘ=& % =z-~y! q+=PE
 [gTQ+̄]F˸*ğǀȢ(ǌɾ®Ʀ n=8+źǌɾ®A8"ǧʵȢïɒ(®ÚAúƱ$̄ˋ¨ı*Ð˙ź ȾıǒȌ)ǧ1$­&ȍŲ =*ÃĄ+̄ʳÑʤɊĒı9]mOʯ˒èˑɔ&!ÚŰɏĒı%˷ʆ%=ˋ¨Ȣ(Ð˙ź+ɁȄĩȞ®AƌÔıˋ¨ĴȢ(ʙǈ%+˦Ľ&(=& ī ̄[gTQ+*ÿɶź ­ƌ9ȘȌȾ&ʚ =ʟ+̄Ĭċ̂2012̃A÷ȅ>
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植物科学最前線 3: 63 ( 2012){yQScf[gTQ*Ô˸ĴȢȱɀ+ 18 ɐ( ,)ʱǻA8" ̄ȾÎÔ˸ɫ8ĲĘ̄×ƽ)+ʜʶă)
844(Ƕ ʖ;>̂ŕǂ 2012̃ėÎ%8 19 ɐŭí ; 20 ɐ) $ǫ˔;):=[gTQ*ɎıŪƅ&!ȱɀ´ = ̂ʃƾ 2012̃ ̄ũƲ+ǘǼɏɛ&Ƞž+4<(!:%=1943 Ş) Burgeff +̄Genetische Studien an Marchantiâ[gTQŎ*ˋ¨Ĵȱɀ̃&ƅƷAÓȋ̂Burgeff 1943̃
̄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) $[gTQĞ˻ɖɵ Ǚɂ>̂Ono 1976̃
*ɖɵ+̄ÈȏɂǍ˻Ǆ¥*Ğ˻ ÿɶ%=&9Ħǣ ǔ7$ȣB%=& ;̄ʅɡ®Rjy9xdSeD
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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植物科学最前線 3: 64 ( 2012)

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
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IiNȨ Joint Genome InstitutēJXd~D Monash ĬĴ John Bowman ðĨ̄ˋ¨ĴȱɀƉAų&ƫĴʍ˴ĝȱɀRjyơƟPr̄ʸȟĬĴĬċçşðĨ;&*ÌĂȱɀ%=ȧĂźɘƞ ):=ˋ¨ıȲħ+̄ˮőȩɂĬĴ˺ȚǽðĨ9ĸģ(Ł)żðĨ&*ÌĂȱɀ%=
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ʜ$ƁʦAZIX
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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, Japan
2
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:

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 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
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x@Ȕ¬=:*kE_cN+ =6ĨŻǸ¥)ƪŶɠ20 N+ PAC}Ep}~:Ł)ȱ,=$ JGI %ȎŸ=ƍ;* 10 N)" $+ö¼*ǀǍǱà*ǵé)ü# $ȱ-ɠ==ÁŌ%ȎŸ= kE_cUPUO*ǣŹ@Ȃɡ)ǃðȎŸ PAC N*ETcT
EX*áȐ+Ǜ 3.3 Mb % ;ɠ=+Ǜ 280 Mb &Ȋǌ6:=$ <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;ɠZ
fSPQix*ȎȘ)ơƏ*óɎ+( &ÅŠ=ēɇɠǒǱ:6àN*U|_cKb]@CYp$3ɠćŜ*ȣǈñČ@æ4 1 N@Ʌ $+ǷĊ(ǣŹĶ:=$ <

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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<

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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 =ZfS

PŽQix)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$6v_q<&%:)ɠJGI %ÜĶ EST *Ǜ 90%6v_q%&:ɠðĶ:=CYp~+Qi
x*ĈÕ@Jj$ <&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EgeX
e*6*9;Ĉ(<´ä) <=+ɠZfSPQix) <Ȳ¡ČėĭUEge
XeQix9;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$*ÚɑɠȷÃ 9-ƈǳə¢ȇŜ*Ȳ¡Č*Ď÷+ɠ=:*ƈǳȎŸ*Ĉ(Ɉĕ&(<ª ,ɠſƠ*ȉ(·ÝĖ¥% <oDcNx+ɠ
UEgeXe)+ 5 ÁČǊĎ÷<ɠ
== ) ȵƯ(<ƈǳ@ÁŌ""6ɠȹȇƵ(ƈǳ6Ȗ5:=<ɞôɤɟĵ!$ɠȹ
ȇ<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¡

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

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植物科学最前線 3: 79 ( 2012)dq /5+) CV["kDL9$:=AZh

)$+ X ŻǸ¥6Ǣŗ @Ƞ( *%ɠY ŻǸ¥&âƃ*´äɞȲ¡Č*Ɖĉɠ 9-c}Wu\7ÚĸȷÃ*Ǿǌɟ@ǃ&ƛ=<
 ZfSP X ŻǸ¥*ȷÃŀý@Ķ<5ɠ=2%)Öɍ X ŻǸ¥ȭȽvJ@ƨ $ɠ
JGI CYp~ ver. 0.6 9;áȐ 3.9 Mb *ȷÃ@ŋÀY ŻǸ¥)Ȋ = 64 °*Ȳ¡Č*

ɠĝ(&6 20 °*szO X ŻǸ¥ȷÃ)Ȋ =ɞôɥɟ
=+ɠZfSP X ŻǸ¥& Y ŻǸ¥âŻǸ¥)ƪŶ<&@ś5$Śŏ$ <ɠ==*Ȳ¡Č*ŻǸ¥%*£ǭɁ­+0&A'®Ď=$ :ɠX ŻǸ¥& Y ŻǸ¥ÁĠ$ŶɠǱ+ŻǸ¥r*½Ǫ@Ǭ;ȧ$&ŕƛ=<2ɠZfSP Y ŻǸ¥)+ɠlc (Egydio de Carvalho et al. 2002) ɠvFW (Lorenzetti et al. 2004)

9-ǨǿN}wdzeW(Ikeda et al. 2007) %ɒƑıń)Ļȉ&=<Ȳ¡Č7ɠſƠ 9- ȵ*ÏƠ%®Ď=$ <ɉ²ÝǘɁȭȲ¡Č (Hirai et al. 2008, Mori et al. 2006) *szO('Ď÷<ɠ=:*ɉȲ¡Č*Ʊȡ+*ŅɋQix)Ȋ"!$ ( ĴɠX ŻǸ¥*ȷÃŦ:)==,ɠY ŻǸ¥ 9-ĨŻǸ¥&*ƐȤ@Ȫ$ɠĿƔĒȲ¡Č7ÕŜ¥ƧƠ) <ĿŻǸ¥*ńǎȯǉŦ:)(!$<6*&Ůĳ=<

tmBPMT ZfSP*Qixb]+À> "" ;ɠ=@Çƨ%<ı)$Ŗ«<*ľÐ%<JGI %*CYpǡ=,ɠȫ7)CiaU|@Ȁ ɠâŨ)=2%)Ǿ
ǌc}WN~qcxb]7ĚƟǙǥ% < Kitashirakawa-2 żƪŶ*b]*ǥá6ƷŐJGI %*ƄƜƵ(b]ºȿƨu])Ì ɠȲ¡ĐǀǍŅ*Ŵ® ×Ą:&»)9;© Ñņ*9 ťŲƲb]rW*Ƃǖ@ȐƬ$ <( ɠQixb] 9-c}
WN~qcxb]*ć+ƥ÷Űºȿ% <ɠə¢ĿƀǞ*~NHWc)+°Æ)ĚĽ$ <*%ɠǵé <ȘǱ+ȭǤ= 

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植物科学最前線 3: 80 ( 2012) ƥ÷2%)ć*ſƠǊ)" $*QixŦ:)=$ɠÇƨ%<ŀýȺžƏ)Ă &%ɠɆſƠ)Ȋ:=<ƃ(ƥȜ@ÁČ7Ȳ¡Č)ǣ-"<qYW+Ėŧ)(!*> Qix*ćƃĿ7ȇɌ*ŢÉɑ)À$5)ɠē+ɆſƠQix*(; Ȋ )(;ɠȲ¡Čr%*36Ȋ )(!$2!$<*%+( > ƃ(QixÀ>!$ɠZfSPQix*ƢƖ@Ňǁ<!@ $=<&ǽǱ:+¯$ <gi Ųǋ%Ü;QixqVGNc+ɠJGIɠMonash ĈĐ* John Bowman ×Ą 9- Sandra
Floyd ×Ą&*öɇ»âǀǍ% <ö¼*ǀǍ)" $+ɠŞȵǇĐƺ 9-Đȁőǵ *ǇĐǀǍȆÍȻ*Íń@Ý$ <kE_cN*ȎŸ+ɠãÞĞĈĐ*ɏůř×ĄɠȶĈĐ*ǻůġ×ĄɠŷĈĐ*ƩȞļ×Ą&*»âǀǍ% <đ¹Ⱦ cDNA *ȎŸ+ɠȶĈĐ*ǅƝǆì×Ą 9-QixơĒŚŘƤ&*»âǀǍ% <b]ȎŸ%+ɠȲ¡ĐǀǍŅ*ȾĢǺƇ×Ą 9-Ŵ® ×Ą9;ćĈ(<ŚŘ@ɓ Ȳ¡øô+ɠȶĈĐ*ÙȻĘëƒ)9<6*% <ūĴ)ɠŲqVGNc)Ɂ?!$ƿĢºĮ×Ą@ċ5&<ǒǱ:*ǀǍĔɞȶĈĐ 9-ȦưĈĐɟ*yj)ƚȚ Jr

UcF  UcF^IkDLHN

X ŻǸ¥& Y ŻǸ¥*ĚĽ<Ȳ¡Č@Ƹǩ%ǣA

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植物科学最前線 3: 84 ( 2012)= ūǜȀǻŹŨŀōŀō̯ɁōɣɲɪɃȹɪōśƶ    ǡŹŨů̧ť 
An introduction to the taxonomy and morphology of Marchantia polymorpha
Key words: bryophytes; Marchantia polymorpha; morphology; taxonomy
Masaki Shimamura
Department of Biology, Graduate School of Science, Hiroshima University
Kagamiyama 1-3-1, Higashi-Hiroshima 739-8526, Japan
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cr~kh (Preissia quadrata) ?̷ģƢ͘ŹŨŀōȷǰɠ©ȊƴƇ͙r~khɪ@ 1 Ť?ȋŔYO6̵ģƢ̷͚ģ
ƢYO6̷ģź@Ñʺ˒:ʽʰYO6e. r~khɖokhɪ͚okh (Conocephalum conicum) ?̵ģźo
khɪ@ 1 Ť?ȋŔYO6̵ģź@ȴǧ̷ģƢǧ@ʪňƹŰ?̴=ƕȮ=¿̨-UÑʺ˒!<ȴƖʻģ!<f. r~khɖodlkhɪ͚odlkh (Reboulia hemisphaerica ssp. orientalis) ?̷ģƢ:̵ģźodlkhɪ@ȋŔ! 13 Ť=ɏ̕-UȋŔ=ėûʁ!<®ČèǦ3%9<#͚ǱȹÄ?ʯ́ S?´ĭɒ<èǦOˏȴƖʻģ!<g. r~khɖov_khɪ͚xdthov_kh (Sauteria yatsuensis)?̷ģƢ͘ȷǰɠ©
ȊƴƇ͙ov_khɪ?ȋŔ@ 23 ŤȋŔ=ėûʁ!<̷ģƢ@˃ȻÄ?̋§=6#̵ģƢ@< ͚ȴǧȋŔŉ?̪đ̛!ǉƆȴƖʻģ!<h. r~khɖ_ekhɪ͚_ekh (Riccia fluitans)_ekhɪ@ȋŔ! 23 Ť=ɏ̕-U̵ģź:̷ģź!<#͚̍Ćģ:̍ʀģ@˃ȻÄâ̛=ȎɃʪňÄ=ǧ!<Ȍ§=Ș̑
+8Ƀʦ-UɮOUi. r~khɖhr~khɪ͚hr~kh (Dumortiera hirsuta) ?̵ģƢhr~khɪ@ȋŔPʯ͕ȷ!ɍ˸ɒ̵ģƢ̷͚ģƢYO6!̵ģƢǧ@̀ų=ɡȴƖʻģ!<

M. Shimamura-4
BSJ- Revi ew 3: 87 ( 2012)

植物科学最前線 3: 88 ( 2012)C    =./0%

 r~kh?è͌ōɒɣɲ@͚Marchantia polymorpha L.!1753Ÿ=Linnaeus=R58ˢ˾)V2':=̗U (Linnaeus1753)M. polymorpha L.͘Źʠ?r~kh͙=@ɃʦĮPɃʦɂĸ=R58͚
Ɔƚ=ĿǷƖ!U':!͚Linnaeusʳ˻P0?Ƌ?ɣɲ9˦NSV8T͚ƺĿ#?ɮâè͌
ʟ͘°ɮPĽɮ͙!ˢ˾)V8"2Burgeff (1943)@0VSYM. polymorpha (Ƚʠ)͚M. aquatica͚
M. alpestris?36?ȼɶ+2ɮ:+8ƻɁ+2͛0?Ƌ͚Burgeff (1943)?L:N23è͌ʟ@͚Ŀ#?ɣɲʣ=M. polymorpha?ɮâè͌ʟ:+8Đ%ÜVSV (Schuster 1992)͚ȿĭ@͞6?°ɮ
?LY˦NU?!ʷɒ:<58U (Bischler & Boisselier-Dubayle 1991, Perold 1999, Paton 1999)
è͌ōɒŒʠ!Ǐǋ952͚M. polymorpha L. (Linnaeus 1753)=̬+8@͚Bischler & Boisselier-Dubayle (1991)=RT͚Ĉˢ˾9ƂɅ)V8Uĩȸ (Dillenius 1741−1742)!f|t^:+8ƪŒ)V20?ƆƚɒȺƑ@͚ƍǟ͚M. aquatica:)V82Ǳȹ=ʴ-U͛ ȿĭ˦NSV8U͞6?°ɮ?4M. polymorpha ssp. polymorpha͘ƍǟ͚M. aquatica:)V
82O?͙@͚˃ȻÄ§ł̛=ȋŔ!èû/.͚ʯ́Ïʌʜ?ʹ!̉%8˙U2N=§ł̛?͗ʗ!̎ʓɒ9Ǉɞ9U͛)S=ʯ͕ȷ?¶ţȹ!Ýʙ9U':͚ɃȅģőPȴƖʻģ
YH:Z;6%<':<;9ȺƑ7%SVU¨=ťĮ?Ȍ́=èŰ+͚²ȳɒ<ɂĸ=@Ƀ
ʦ+<ǚ°ɮ@ǅǚ=OèŰ+͘üŬ 1987͙
͚vr~kh:RAV8Uĩ4=ŹŨŀōǱȹǸǚžƝˈ? Ÿ=ʟ͐əšȰhĈ9Ư̶)V2ǸǚYɧ-M. polymorpha ssp.
montivagans (ƍǟ͚M. alpestris:)V82O?)@͚˃ȻÄ?ŵ!2 cm=̕-U':OUŀİ?Ǳȹ9͚˃ȻÄ?§ł̛?͗ʗ!LSV<w9@͚ťĮY§ƒ=è?
˯Ř<ɂĸ=Ƀʦ+͘șŧ̃#?ķƖȤĮ9?Ư̶ˢ̦OU͙
͚²ȳɒ<ɂĸ=@Ƀʦ+<
ǚ°ɮ@'VK9?Ɲ͚ǅǚ9@èŰ!ɥ˦)V8<M. polymorpha ssp.ruderalis ͘ƍǟ͚
Ƚʠ?M. polymorpha:)V82O?͙@͚˃ȻÄ?§ł̛?͗ʗ͘ȋŔ!ɏ̕+<̛è͙!£̎ʓ9͚ʯ͕ȷ?¶ţȹ=̤ȁ!U':͚ȴƖʻģPǔƖɃȅģőYǆɔ=6%U<;?
ȺƑ!U²ȳɒ<ɂĸY§ƒ=üĀɀÝı=Ź#èŰ+͚āĀɀ=O̜ư̊Y§ƒ=ɬÜ!
LSVUǅǚ9O²ŕ?Ğ́9Ǎ̌=Ƀʦ+͚“r~kh”?ġĘ9ĠAV͚Ŀ#?ɣɲŔ9ɣɲǛƾ:+8ɅSV8U?@'?°ɮ9U Burgeff (1943) ?±̸œ͒@¡̅?3°ɮ?̙½ɒ̬ËYʢU¡9͚ʵğȝʎǥYɧ+8
U0V=RU:vr~kh:M. alpestris (ȿĭ͚ssp. montivagans:)V8UO?)?̫9@͚
H:Z;±̸!˵'S<!͚(#ű=Ƀ,2̸ɮ=r~kh=À2ƆƚYO6O?!Uǂ9͚r~kh@͚vr~kh:M. alpestris0V1V:?̫9Ŗǈ=±̸-U0VQ͚Ɖ@r~kh!͚vr~kh:M. alpestris?̸ɮ=˵Ĉ-U:ʢ2Schuster (1983, 1992)O0?ʢYƵƩ+͚r~kh?Ƀʦ!²ȳɒɂĸ=̮SVU': Sr~kh@͚ÚĔǌ·=Ƀ,2̸ɮ=˵Ĉ-Ù͚ų=ǁ+ɮ:ʢ20?ƋˏXV2\m^DNAĿİ?˞ǣY̌
,8͚ 6?°ɮ!̙½ɒ=Oýí9"U':@ǇS =<52!͚r~kh!µ?2°ɮ?̸ɮ
=˵ȥ-U:ˣƧ@ƎSV8< (Boisselier-Dubayle & Bischler 1989, Boisselier-Dubayle etal. 1995)ǨʹÄƺ@.V?°ɮO9ǚ (n=9̷̵͚?ƖǨʹÄYěM) 9U':!è 58
U (Bischler 1986).

M. Shimamura-5
BSJ- Revi ew 3: 88 ( 2012)

植物科学最前線 3: 89 ( 2012)ĩ ͛vr~kh(M. polymorpha ssp. polymorpha)?Ǹǚ ˃ȻÄ̷͘ǭ͙?ľƆ͛˃ȻÄ@ʉ̨#͚ŵ@  ͛èǦ?˝ż!r~kh:ȈG̣˝Ɯȵ+2̷ģƢ͘ɟą͙@˃ȻÄ?Ĳ̛̃#=Áʞ-U̷ģź@Ş)!͚ǧ@ :̨#¿CU'?Ǹǚ@ȴƖʻģY6%8<͛˃ȻÄ§ł̛¶̃͛§ł̛=ȋŔ!Ý#èû/.͚
§ł̛?͗ʗ!̎ʓɒ9Ǉɞ͘ɟą͙ʯ͕ȷ?¶ţȹ̤͛ȁ!<#͚Ş)<ʉʪ9ʙďSV͚Ýʙ͛͘ĩ  ?r~kh?ʯ͕ȷ?¶ţȹ:Ȉ˽/R͙Specim. exam. Japan. Pref. Gunma: Ozegahara, 1400 m alt. H. Ando 9143 (HIRO).

D

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 r~kh?ɃȕĔ?ǵɉYĩ 5 =ɧ-͛r~kh@µ?Ǳȹ:ė,#͚Ǯɘ?ĽûY¾¤·±·Yˏµ?jhǱȹ:ėǷ͚Ƀȕ?¨Ä:<UǱȹÄ͘˃ȻÄ͙@Ăɘ?̝ÐÄ9U
ǱȹÄ@̷̵ɋǭ9̵͚ǭ=@̍ʀģY6%U̵ģƢ̷͚ǭ=@̍ĆģY6%U̷ģƢ!ƆƜ)VUĿ#?t^͌9@͚Ƀȅģő!Ÿ̫?̮SV2ǌǖ?LƆƜ)VU!͚r~kh@Çľɒ
9̡͚ľ=8OåǖY̰8̨͚ǖ̫=X2Tǿ:̵ģƢ:̷ģƢ!ƆƜ)VU̡ľ?
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M. Shimamura-6
BSJ- Revi ew 3: 89 ( 2012)

植物科学最前線 3: 90 ( 2012))VUʪňÄ@̷ǭ?̷ģź?¢̛=ƛį+8Ɯ̨-U͛
ʪňÄ?â̛9@ȟƺè˓!'T͚Ăɘ?ʪň!ƆƜ)VUʪňÄ SƷç)V2ʪň!ɏʻ+͚ĈʁÄ:ĠAVUÄðYʍ8Ǳ
ȹÄ!ƆƜ)VUǔƖɃȅY´+2ʪňƆƜ3%9<#͚ǱȹÄ?ȴƖʻģ?â̛=ƆƜ)VUȴƖʻ=R58Oǆɔ<ȴƖɃȅYˏ¸¢͚ɃȕĔ?ĕȆ̲9˙SVUƆƚ=68͚RTˤ+#˞˨-Uĩ 5͛r~kh?Ƀȕɂ

E33-4
 r~kh?ʪň@ 10–15 µm ɭż?ɀƆ9͚ʪňÄ SƷç)V2:͚̘ƅ<ɂĸǞº!ƻ
A»ɛ-U':<#͚234=ɏʻ-Uɏʻ@ÛÉŋɒ9͚ʪň!ÛĖƜYˏ':!̠˗
9U':!è 58U (Inoue 1960, Nakazato et al. 1999)ʪň@ɏʻ?̴=͚ŀŞ͝6?ʉʪ
YɃ,U£ɹè˓Yˏ͘ĩ 6a͙
'?4Ş)ʉʪ S@ǒë?¹ǯ͘ŷȪ¹ǯ͙!¿̨+͚
ŀ"ʉʪ@ƺżè˓-U':9͚ƺʉʪ!ê=̎<T͚Ǚɷ=¹ǯYO6ĈʁÄ:<U(Inoue 1960)͘ĩ 6b͙
0?Ƌ£˚ñ<ʉʪè˓Yˏ͚ĵȻ?ľƆYʍ8͘ĩ 6c͙͚Uɭż?
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M. Shimamura-7
BSJ- Revi ew 3: 90 ( 2012)

植物科学最前線 3: 91 ( 2012)ĩ 6͛r~kh?ĈʁÄa. ʪň?ǒë?ʉʪè˓9Ƀ,2Ş)ʉʪ S¹ǯ!¿̨-U͘ɟą͙b. ŷȪ¹ǯ͘ɟą͙YǙɷ=O6͚ƺʉʪ!̎<52ĈʁÄc. ëǖ?ĈʁÄ@ȺŒ?ʉʪè˓tYO2.͚Ƿ<ĵȻ?ľƆ
YO6d. ĈʁÄ¡=Ƀ,2ͅɷʉʪ͘ɟą͙
e. ͅɷʉʪ!Ƀ,U:͚0'!ȣƯ̈͘wv͙:<58͘ɟą͙͚ŷ́ɒ<˃ȻÄ?ƆƜ!ŇKUF$(>5)? Ɯȵ+2r~kh?ǱȹÄ͘˃ȻÄ͙@ŵ 8 mm

S 15 mm ɭż͚§ł̛?ćL@ 0.3 mm

S 0.6 mm ɭż9͚®ČȻ=èǦYʝT̄+8Ɯ̨-Ur~khɖ?˃ȻÄ?ǀ́@˖̸=èû+2ŤȻǶ̍:<58T͚ʌʜèû?Ăʆ<xkhʕ?˃ȻƖt^͌?Äð!Ăʆ˃ȻÄ (simple thalloid) :RAVU?=Ś+͚˖̸˃ȻÄ (complex thalloid) :RAVU͘ĩ 7͙˃ȻÄ?ͅɷ̛@ƒʲİ=ȣÜ+8T͚0?¶̃!ʉʪè˓?ɔZ<Ɯ̨Ȳ:<58UƜ̨ȲYǶƜ-Uʉʪʟ?§ł=@͚͜6?ͅɷʉʪ (apical cell) !L:NSVUͅɷʉʪ@Ğ́?
ʉʪ=ȈG8ŀİ9͚ʉʪ˳=ŘZ9T͚¦Ťè˓=R58Ƀ,2ʉʪ=R58Śɫ=ďTĨ
KV8U':9͚µ?ʉʪ:ýí9"Uͅɷʉʪ?Ğ́9@ȕɏ<ʉʪè˓!˵'58Uǂ9͚ͅɷʉʪʳÄ?è˓͊ż@Â#͚ʉʪYéTç-ǂęYĉŗ=ðƏ-U':9͚ǱȹÄ?ƆƚƆƜ=̠˗<ƈóYƦ58UjhǱȹ9@ͅɷʉʪ SéTç)V2͜6?ʉʪ͘qg|͙=Ɇǟ-UʉʪʟY [^|:RC͚
ǱȹÄYǶƜ-Uʌʜ!˚ñɒ=èû-U
jhǱȹ?ǱȹÄ@ [^|!ɰL̠<52Ƕ̍9T͚ [^|@ǱȹÄ?ʌʜèû

M. Shimamura-8
BSJ- Revi ew 3: 91 ( 2012)

植物科学最前線 3: 92 ( 2012)?ĲǚĂÁ:<58U:Uͅɷʉʪ?Ɔ͚ͅɷʉʪ?ʉʪè˓ǂę?ðƏ͚ [^|§9?ʉʪè˓t?˚ñƖ@͚ĕè͌ʟ?Ĳǚɒ<ƆƚƆƜYƵ̝+8Ur~kh?ͅɷʉʪ@ 4 6?ʉʪè˓́YO6ǳƆ9U͘ĩ 8͙ͅɷʉʪ?ʳɆ́͘ľɇ=ư-Ú͙
@Ǳȹ?ʯ́Ï=Õ8̝ʞ+8T͚ʯ́Ï Sɏ̕-UɿȜȉ:ʯ͕ȷ=R58Íˮ)V8
Uͅɷʉʪ SǱȹÄ?ʧ́Ï=éTç)V2qg|=Ɇǟ-Uʧ́ [^|͚ʯ́Ï=éTç)V2qg|=Ɇǟ-Uʯ́ [^|@͚˃ȻÄ?§ł̛YǶƜ-Uʌʜ
͘ːɓ:͓˳͙:<Uͅɷʉʪ?Ïǂ=éTç)V2qg|=Ɇǟ-UÏǂ [^|S@͚˃ȻÄYǶƜ-UÝ8?ʌʜ!èû-U͘ĩ 9, 10͙ [^|§9 4 Ħ?ʉʪè˓!'T͚Ïǂ [^|! 5 ʉʪ:<52ǌȲ9͚ʧ́ːɓʌʜ͘ːɓȋŔ͙͚ʧ͓́˳ʌ
ʜ͚ʡ̛͘˃ȻÄ?ʙ͙
͚ʯ́ːɓʌʜ͘ːɓʯ͕ȷ¹ǯ<;͙͚ʯ͓́˳ʌʜF:èû-U
ʉʪ!ȍŒ-U (Crandall-Stotler 1981)r~kh?ĴĖ͚Ïǂ [^|?ëǖ?è˓t
@͚ͅɷʉʪ:Ý#ė,9T͚®ǿͅɷʉʪ (secondary apical cell) :ORGUƭUʶYU2N͚ͅɷʉʪ:Ïǂ [^|?˫íYʌʜéȷ× Sìǀ-U?@̺+r~kh@Ă?ͅɷʉʪYO2<:˞̟+2̔Ċ?ɣɲÇOĿ#U+ +̎ʓéȷYÆ˕+͚ͅɷʉʪɗ¢?ʯ́ [^|!ʯ͕ȷ?ĈĲYƆƜ+<':YƪǸ=-VAͅɷʉʪYėŒ9"U ˃ȻÄ?ʧ́Ï@͚ÝÄ!fvf9˘XV2ːɓ!ɏ̕+8U0?ɗ¢=@͚ȋŔ(airchamber)!èû-UÎ?ȋŔ@ 6 ˝Ɔ?ŞŔYƆƜ+8U2N͚˃ȻÄYʧ́ SɜNU:͡˝Ɔ?ǹǷ!LUȋŔ@§ł=Áʞ-UȋŔŉ (air pore) =R58ľɇ:̌,8UȋŔŉ@͟ê?ʉʪ=ɂȻ=ďTĨKV2ʉʪ̫̳9T͚ʚǀ́9@Ǽİ=̝ʞ+2ʉʪʟ!
˜ř)VU͘ĩ 11͙ʷɒ=@r~khȋŔŉ@͚ʔɻǝǱȹ?ȋŉ?R<̪̩̒÷@+<:ˢ̅)V8U+ +r~kh͌?#6 ?ɮ͌9@ȋŔŉYĨMǼİ=̝ʞ+2ʉʪ?4͚ǒOŻ̛?ʉʪ!͚ʱĬ?Ľû=RTĽƆ+͚ȋŔŉ?̌ȋƖY˩ƻ+8U:ʢS
VUÇ!˜ř)V8U (Walker & Pennington 1939)ȋŔ?â̛=@˃ʖÄYĿ#ěMʉʪ͘ėûʁ; assimilatory filament͙!èû+͚ÛĖƜYƦ58U͘ĩ 11͙ȋŔŉ:ʉʪ̫̳YÔ2
ȋŔ@͚ȋŉYÔ2ʔɻǝǱȹ?˃ʤʌʜ:ėǷ=͚ÛĖƜʌʜY«ȶ SŎT66͚ƨƹ+P-ȋɘ?®̞ûȱʇYǱȹÄâ=ŝ#':=ƈɶ58U(Meyer et al. 2008)ȋŔŉ?̪̩
̒÷!< ͚£ŐÝ<O?9U:-VA͚ʔɻǝǱȹ?ȋŉ:ȈG8ˆƹ=RUȌ?ƲŃ!
ŀ":­Ȣ)VU+ +͚'?ˆƹÆɅ!͚Į́=ư+2˃ȻÄ¢́ S?Ȍ?ĜĎYŝ
8U? O+V<r~kh͌?ȋŔ?ɏɃ˵ȥ=̬+8@͚ʧ́?ǒľŤ?ʉʪê: 2 Ɋɖ?ʉʪê?̫=Ƀ,2ʉʪ̫̳=˵ȥ-U˨͘âɃ˵ȥ; Hofmeister 1851 <;͙:ʧ́ǒľŤ?
ʉʪėļ?ʉʪ̫̳=˵ȥ-U˨͘ľɃ˵ȥ; Leitgeb1880 <;͙?͝6!52!͚ tr
~khYɅ2 Apostolakos & Galatis (1982) ?ˤʉ<ʌʜōɒɣɲ@͚ȋŔ!ǇS =ľɃ˵ȥ
9U':Yɧ+8Ur~kh=8O͚ǱȹÄ?ͅɷ¶̃?ʌʜéȷY˜ř+͚ȋŔ?
ƨŀ̔ɭY̆:͚ǒľŤ?ːɓʉʪėļ=̳̫!Ƀ,͚0V!ƨŀ+8ȋŔ:<58UǷň!˜ř9"U͘ĩ 12a–c͙ ȋŔ?¢̛?˃ȻÄ?͓˳(ǩʌʜ; parenchymatous tissue)@˱ˈʌʜ:<58Uʉʪ@µ?
ʌʜ=ȈGŀ"#͚ʉʪ̫̳!ɏ̕/.͚˃ʖÄ@µ?ʌʜRTOŞ)#͚ȭɽɾ!ɏ̕+8

M. Shimamura-9
BSJ- Revi ew 3: 92 ( 2012)

植物科学最前線 3: 93 ( 2012)

U˱ˈʌʜ?ʉʪĺ=@ʉʪ?̨˼ǂę=ɗ±-Uǂę=̉Ǉ<ʥć!LSVU(ĩ 13)â̛=ŀ"<ȐÄ (oil body) YěMʉʪ͘ȐÄʉʪ; idioblast, oil cell͙@͚˃ȻÄ?ĕʌʜ9Ɲ=
èû-UĂʰʂbd9UȐÄ@͚^s^}ûĖȹ?ĖƜ͚ˇɰ?Ĵ:<58
U͘Suire et al. 2000͙ ˃ȻÄ?ʯ́Ï͘Į́=ư-UÏ͙=@͚ʯ͕ȷ (ventral scale) :¹ǯ (rhizoid) !èû-U(ĩ13)¹ǯ@˃ȻÄʯ́?ːɓʉʪ!è˓Y¾X.¿̨+8Ƀ,2O?9͚͜ʉʪ S<U¹ǯ=@͚ƽȲȻ͚ʗȻ?âɃʥć=RUʅǷYO6ǔʅ¹ǯ (pegged rhizoid K2@ tuberculate
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Schiffner1909, üŬ 1977)ǔʅ¹ǯ@Ł) 10-20 µm ɭż?O?!Ŀ#͚ʯ́ÝÄ SɃ,8͚˃ȻÄ?§ł̛Yɖƪ-R=ʯ́=ȑ58¿C8U(ĩ 14a, b)ǔʅ¹ǯ@͚ʥćYƆƜ-U̔ɭ9ʉʪ˳YŃȃʉʪ:<U!͚ǝû-U':9͚0V1V?̫=Ö#ȉʉɻȿ˰YîɅ+͚˃ȻÄʯ́ÝÄ=ȌYÈʏ-Uľ̛̌ŝ?ǽʬYƦ58U (McConaha 1941)ǂ͚ŷȪ
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=ŘZ9U23+ʉʪĺ!ˉ2N͚˃ȻÄ?Ē̛è9@6EV2ƆȻ:<58U':
OĿŷȪ¹ǯ@͚¢ǂ=¿̨+͚˃ȻÄYĲȹ=ĪŒ+͚Ĳȹ SȌèP͏èYŝÜ-Uʍ
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r~kh͌?Ŀ#?ɮ9ˁǯYƆƜ-Uˁ͌!Ĳȹ=?CUŷȪ¹
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M. polymorpha ssp. montivagans ?ǱȹÄ=@͚µ?r~kh͌:ėǷ͚ßɃˁ!LSVU!͚r~kh:vr~kh=@LSV< (Ligrone et al. 2007)ɵʇè?ĿĴƝ=ńZ9Ƀʦ-Uè͌ʟ9@ˁ͌:?ßɃ!Ɠ˗< ͚ˁ͌:?ßɃ=̬-UǽʬYŃ58+K52? O+V<r~kh?ǱȹÄː́=@͚vfz_!ɃƗ+8T͚Ǳȹ?Ƀ̨YÌ̏+8
U:ĳĝOU (Kutschera et al. 2007)
 ʯ͕ȷ@͚ʯ́ÝÄY˘R=ɏ̕-U͜ʉʪć?͕ȷȻ?ʌʜ9͚0?ƆP̝ê@͚r~kh͌?ĕè͌ʟ9ȺƑ!T͚ɮYėŒ-U̴? 1 6?ɖŏ:<Ur~kh9@˃ȻÄ?§
łYƬZ9͞ê.6͚ˠ͡ê=̝ʞ+͘ĩ 15a, b͙͚§ł?͝ê=@µ?͟ê=@<ɴ˵Ȼ?¶ţȹ!U͘ĩ 15c͙'?¶ţȹ@͚O:O:˃ȻÄ?ͅɷ¶̃9Ɯ̨ȲYʯ́Ï Sʧ́Ï=˘R=ɏ̕+8Íˮ+8U¶ţȹ@͚˃ȻÄ!Ɯ̨+ʯ͕ȷ?¶ɝÁʞ!Ɯ̨Ȳ S̹
VU=ƍ͚ʯ́Ï=Ƃ"̂KV8ʯ͕ȷ?òǂĞʙ̛?ɴ˵:+8ȄU͘ĩ 15d͙§ł?͝ê
?ʯ͕ȷ@˃ȻÄ?§ł̛=ȑ58̨#¢į-ƯèYO4͚ŷȪ¹ǯY¥Ï Sǝ>U͂Ȼ?
Ƕ̍YƆƜ+8T͚öȾ?ʸȌè̀̈?̌˹:<58U (McConaha 1941, üŬ 1977)͘ĩ 14b͙ǅǚɄr~khţ?µ?ɮ͌9@ʯ͕ȷ@ 4 ê=¦C͚r~kh?R=˃ȻÄ?ʙ Sŀ"#
ľ=@Lç-':@<2N̡͚ľ9ɮYìí-U2N?ƪǸ=<U͘ĩ 15d͙

M. Shimamura-10
BSJ- Revi ew 3: 93 ( 2012)

植物科学最前線 3: 94 ( 2012)ĩ 7͛r~kh˃ȻÄ?Ǻǀ́?Ƕ̍ʧ́Ï?ːɓʌʜ=@ȋŔŉ9ľɇ=̌,2
ȋŔ!èû+͚0?¢=èćǩʌʜ͓͘
˳͙!èû-Uʯ́ːɓ Sʯ͕ȷ:¹
ǯ!èû-Uǔʅ¹ǯ!ʯ͕ȷ=ȑR
=˃ȻÄ?ʯ́ÝÄ=èŰ-UŷȪ¹
ǯ@Ĳȹ=ę 58¿C8U˃ȻÄ?
Ɲ=ȐÄʉʪ!èû-U

ĩ 8. ͅɷʉʪ͘ǉą͙YěMʌʜéȷa. ˃ȻÄ?ʧʯ́=ŷˏ<éȷb. ʧʯ́=įɗ 6Ɯ̨˼=ŷˏ<éȷc. ʧʯ́=įɗ 6Ɯ̨˼YǺǀ-Uéȷd. ͅɷʉʪ:Ğ́? [^|?è˓t?ǹƁĩ

M. Shimamura-11
BSJ- Revi ew 3: 94 ( 2012)

植物科学最前線 3: 95 ( 2012)ĩ 9͛t^͌?è͌ʟ(:? [^|?è˓t:͚ʌʜ?ɏɃ˵ȥ:<Uʉʪ?̖a. jvkhʕj
vkh͘ʿ˃Ɩ͙
͜6? [^| S͜Ǥ?˃!ƆƜ)VUb͛jvkhʕ|^kh͘ùú-Uʿ˃Ɩ͙
͜6? [^| Sŀ"<Ï˃:Ş)<ʧ˃!ƆƜ)VUc.xkh°ʕxkhɖ͘ʿ˃Ɩ͙
͜6? [^| S͝6=éV̂Z3͚U@͝6=ƤTɌKV2͜Ǥ?˃!ƆƜ)VUd. r~kh°ʕ͘˃ȻƖ͙jvkhʕ:ė,è˓tYɧ-!͚jvkh9˃YƆƜ-Uʉʪ!͚˃ȻÄ?ʙ͘ʡ̛͙:<Uĩ r~khÏǂ [^|?ʚǀéȷʡ̛?ƟƆ?
ʉʪ

SéTç)V2ʉʪ?

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ʡ̛?ƟƆ?ʉʪ

SéTç)V2͜ʉʪ=Ɇǟ-U

M. Shimamura-12
BSJ- Revi ew 3: 95 ( 2012)

植物科学最前線 3: 96 ( 2012)ĩ r~kh?ȋŔ:ȋŔŉ ˃ȻÄʧ́ SL2ȋŔŉ͠͡˝Ɔ?Î?ȋŔ?§ł̛=ȋŔŉ!̪đ-U͘ɟą͙˃ȻÄ§ł̛=@ȋŔ!èû+<͇ı͘ɟ͈͙!£̎ʓ=èŰ-UȋŔŉ?ǀ́ȋŔŉY:TK#
Ǽİ=̝ʞ+2ʉʪ@͚ȋŔŉ=́-UÏ!ʥć-UȋŔâ̛=@˃ʖÄ=ŘZ3ėûʁ!UːɓŤ=@˃ʖÄ!èû+<ʧ́Ï S˙2ȋŔŉ?ǒ¡̛˜ř?2N=ėûʁY̰Ċ+2ȋŔŉ͘ǉą͙@ãƆ?̪đ̛Y
O6ȋŔŉ?Ż̛ʥć+2  6?ʉʪ!/Tç+þŊƆ=̪đ+8Uĩ ȋŔ?ƆƜ̔ɭ ʾːɓʌʜ?ǀ́ȋŔŉ?˵ȥ:<Uːɓʉʪ̫=Ƀ,U-"̫͘ɟą͙:ƨŀ+66UȋŔ͘ɟ͈͙
ɏ̕§?ȋŔǒë=-"̫!Ƀ,2Áʞ͘ɟą͙9¡¢ǂę=ʉʪè˓!'U':9͚ȋŔŉĞ́?Ǽİ?ʉʪ̝ʞ!6#SVU͘ɟ͈͙ėûʁYÔ2ȋŔĩ ǩʌʜ͓͘˳͙?ʉʪ ǩʌʜ?˃ʖÄ@ėûʁ?O?:ȈG8Ş)#͚{ɾ!ɏ̕+8U§ł
?ʉʪ@ȐÄʉʪǩʌʜ?ʉʪ=@̉Ǉ<ŲȻ?ʥć!U

M. Shimamura-13
BSJ- Revi ew 3: 96 ( 2012)

植物科学最前線 3: 97 ( 2012)ĩ 14. ǔʅ¹ǯ:ŷȪ¹ǯa. ˃ȻÄ?ʯ́=ȑ58¿CUǔʅ¹ǯƽȲȻ͚ɂȻ?ʥćYO6b. ˃ȻÄʯ́§ł?ǔʅ¹ǯ?ǝ¥Ï Sʯ͕ȷ=˘XV8Uc. Ĳȹ=ę 58¿CUŷȪ¹ǯĩ 15. ʯ͕ȷ?Ɔƚa. ˃ȻÄ?ʯ́ʯ͕ȷ͘ɟ͈͙!ʯ́ÝÄY˘R=èŰ-UâÏ?ʯ͕ȷ
=@µ?ʯ͕ȷ=@<¶ţȹ͘ɟą͙!Ub. ʯ͕ȷ?èŰ?ǹƁĩ¥Ï= 3 ê.6͚Ėˠ 6 ê=̝ê-Uc. ǒâê?ʯ͕ȷ?¶ţȹĞʙ=̤ȁ!U͘ɟ͈͙d. ʧ́ S˙2˃ȻÄͅɷ̛ͅɷʉ
ʪ!Uwv?̛è@ʯ́Ï S/Tç+2ʯ͕ȷ9Íˮ)V8U͘ɟą͙ǒľê?ʯ͕ȷ@˃ȻÄ?ʙ S@Lç+8U͘ɟ͈͙G'4'4 ˃ȻÄ?ʧ́§ł=̛=ȑ58͚ȴƖʻģ͘ǠȻÄ; gemma cup͙!ƆƜ)VU͘ĩ 15a͙ȴƖʻģ?ʙ=éV̂L!T͚ľÏ=ª͈Ȼ?ɴ˵!U':@͚|lr~kh<;ŷȪ<ȴƖʻ

M. Shimamura-14
BSJ- Revi ew 3: 97 ( 2012)

植物科学最前線 3: 98 ( 2012)ģYO6̃ʙɮ:?˫íȲ? 1 6=<U͘ĩ 16a, b͙ȴƖʻģ?â̛9@͚Ż̛?ʉʪY˵ȥ:+8͚ȴƖʻ!ɶ4¡!UƆ9ǿ:ƆƜ)VU͘ĩ 16c͙ȴƖʻ@Ăʉʪ?ǧ:͝6?ȣƯ̈
YO4͚§ł̛=ćL?U͚ãƆ?ŷ2ƆȻY+8U͘ĩ 16d͙ȴƖʻ!Ɯ̨-U̔ɭ9͚ȣƯ̈YÍˮ-UɿȜȉ!ÆSVUȐÄʉʪP¹ǯ?ĈĲ:<UʉʪOèû-U!͚ȴƖʻģ?â̛=ɈK58U4@ȋŔPʯ͕ȷ@èû/.͚Ɔƚɒ=ʧʯƖYƩ2<ʉʪè˓@Ğʙ̛ÝÄ9ė˩ɒ='U2N͚ȴƖʻ@˃ȻÄ?R<ǂęƖ?U¿̨Ɯ̨Y/.͚ãƆ?ľƆYÍ52KKƜ̨-Uǿ:ÆSVƜ̨-UȴƖʻ@͚¯=ƥ+Ė':9ǧ?̛è!è̹+͚ȴƖʻģ?â̛=ˇɰ-Uˇɰ+2ȴƖʻ@͚ȴƖʻģ?ʙ!ƿN¡ǂ=Ź!52Ƕ̍ (splash cup)=<58U2N͚ȴƖʻģâ=˂42̻ɾ=R58͚@>:A)V8Ğ́=͎CƹUȴƖʻ!ȴƖʻģ S 120 cm ?˷̹=K9˺Z3:ˢ̦OU (Equihua 1987)ȴƖʻ!Į́=˂4U:͚Į́=ư-UÏ=¹ǯ!èû+͚ȣƯ̈?ͅɷʉʪ!ȕ÷YŇN͚ʧʯ
Ɩ?Uʌʜèû!ŇKUĩ 16. ȴƖʻģ:ȴƖʻa. ȴƖʻģb. ȴƖʻģ?ʙʙ:0?ľÏ=ɴ˵!Ƀ,Uc. ȴƖʻģ?Ż̛?ƆƜ̋¡?ȴƖʻȴƖʻ@ĂʉʪƖ?ǧ͘ɟą͙YO6d. ǧ Sè̹+2ȴƖʻǧ!è̹+2ɍ͘ɟ͈͙:͝6?wv͘ɟą͙H:!;!
 ̵ǭ:̷ǭ@0V1V̵͚ģƢ (antheridiophore)̷͚ģƢ (archegoniophore) :RAVU͚ÓȻ?ɃȅǦYèû-U͘ĩ 17, 18͙̵ģƢ?¡̛?ãɕȻ?Ƕ̍9U̵ģź (antheridial receptacle)͚
=@̍ʀģ (antheridium) !èû-U̵ģź@˃ȻÄ=R#À2Ƕ̍YƩ58T͚QUP =M. Shimamura-15
BSJ- Revi ew 3: 98 ( 2012)

植物科学最前線 3: 99 ( 2012)

8 ˓-UO?!Ŀ͘ĩ 17 a, c͙̵ģź!Ɯ̨-U̔ɭ9͚ʡ̛ Sʧ́Ï=ȋŔP̍ʀģ͚ʯ́Ï=ʯ͕ȷ:¹ǯ!èû-U͘ĩ 17b͙̵ģƢ?ǧ?ǀ́Y˜ř-U:͚˃ȻÄʯ́ S̎ʓ
-Uǔʅ¹ǯ?ǝY 2 êÔ8U':!è U͘ĩ 17c͙Ɯȵ+2̵ģź@͚ŵ?Źʡ̛Y
O62N̵͚ģź?¡̛=ȌY2NP-Ƕ̍:<58U͘ĩ 17b, d͙̷ģƢ¡̛?ÓȻ?̷
ģź (archegonial receptacle) @͚ȝ# 8-10 ˓+͚ƪȻɴ˵ (digitate ray) :RAVUǶ̍YO6͘ĩ
17a͙
Î?ƪȻɴ˵?̫=@͚ͅɷʉʪ!T̷͚ģź?Ɯ̨̔ɭ9̍Ćģ (archegonium) !
ǿ:ƆƜ)VUƪȻɴ˵?ɏ̕!ǘȵ<ʾ̷ģƢ9@͚̍Ćģ?̛͉!̷ģź?¡ǂPÏ
ǂ=ę8U!͚0?Ƌ̷͚ģź@ʌʜÝÄ!âǐ+<!SƜ̨-U2N͚̍Ćģ@˙ %¡͚ƛį+2Áʞ=ƆƜ)VUR=<U͘ĩ 18b)̷ģƢ?ǧ?ǀ́Y˜ř-U:̵͚ģƢ:ėǷ͚˃ȻÄʯ́ S̎ʓ-Uǔʅ¹ǯ?ǝY 2 êÔ8U (ĩ 18c)̷ģƢ?ǧ9@͚ȋŔPʯ͕ȷOèû-U̵ģƢ̷͚ģƢ?ǧ@Ɔƚɒ=@ʉ̨#ƌ̨+2˃ȻÄ:˞̟9"UK2̵͚
ģź̷͚ģźO¡̛=ȋŔ!èû+͚¢̛=@͕ȷǷ?Ƕ̍!èû-U<;͚Ĳǚɒ<Ƕ̍@˃ȻÄ:ė,9U̷ģź?ƪȻɴ˵?ǀ́Y˜ř-U:͚˃ȻÄ!âǐ+͚ɺȻ=<52O?
9U':!X U͘ĩ 18d͙

ĩ 17. ̵ģź?Ƕ̍a. ̵ģƢľƆb. ̵ģź?ʚǀ́c. ǧ?Ǻǀ́d. ̵ģź?ʯ́ʡ̛Y̰"͚ŀ̛è!ʯ
͕ȷ9˘XVU

M. Shimamura-16
BSJ- Revi ew 3: 99 ( 2012)

植物科学最前線 3: 100 ( 2012)ĩ 18. ̷ģƢ?Ƕ̍a. ̷ģƢľƆb. ̷ģƢ?ʚǀ́͘¢̛?ǧ?̃#͙
c. ǧ?Ǻǀ́d. ƪȻɴ˵?Ǻǀ́I818 ̍ʀģ@̵ģź?ʡ̛?ǳƆ?ʉʪ SéTç)V2ːɓ? 1 ʉʪ=˵Ĉ-U̍ʀģ!Ɯ̨-U̔ɭ9͚0VYďTĨMR=͚ȋŔ=R#À2͚ pjȻ?ʭƝ̍͘ʀģʭ͙!ƆƜ)VU͘ĩ 19a͙̍ʀģʭ?¡̛PĞ́=U̍ʀģYěK<ȋŔ:@ɋ<T͚̍ʀģʭ?â̛=@͚ėûʁ!èû/.͚ĂʉʪƖ?ɿȜȉ?L!şƺɃ,U͘ĩ 19b͙̍ʀģʭ?¡̛@͚Ş)ŉ9ľ̛:̌,8U̍ʀģ@ɡǧYO4͚1 ʉʪć?ohw|ʉʪ?â̛@ʀĈʌʜ (spermatogenous tissue, androgonial tissue) 9UʀĈʌʜ@Ŝǟʀň:<UʀĈʉʪ(spermatogenous cell) ?L S<UʀĈʉʪ@ʉʪ?¿̨Ɯ̨YH:Z;/.͚è˓ǂę!ɗ±-Uʉʪè˓YʝT̄-2N͚ʀĈʌʜ@Ŀƺ?Ş)<ǂƆ?ʉʪ S<U͘ĩ 19c͙̍ʀģâ
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O6͘ĩ 19d͙ t^͌?ǱȹÄ?Äʉʪè˓9@§ƒÄ?R<ɀȻ?ƐŞɻƆƜ§ƒ͘ǴƆƜÄ; polar

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BSJ- Revi ew 3: 100 ( 2012)

植物科学最前線 3: 101 ( 2012)

organizer͙ SƐŞɻƆƜ!'U!͚0?â̛=@§ƒŞÄ!< (Brown & Lemmon 1990,
Shimamura et al. 2004)+ +͚ʀȇʉʪ?è˓9@͚1 Ś?§ƒŞÄ (centriole) YO52§ƒÄ(centrosome)!çȿ+͚ʈ̥Ä?ƐŞɻƆƜ§ƒ:<U (Carothers & Kreitner 1968)§ƒŞÄ@͚ʀʉʪ!ʀňFĽƚ-U̔ɭ9̓ȉ?Ĳ̛˔ʞ͘ĲŻŞÄ; basal body͙F:Ľû-U͘Moser &Kreitner 1970͙
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!ŀ̛èYĄNŲʫÄ̛YO4͚Ǯ=ȑR=p^:RAVUƐŞɻ?ǝ!U
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\!ȷ˛̙½-U':!ɧ)V8T (Pacak & Szweykowska-Kulińska 2003, McDaniel et al.
2007, Jankowiak-Siuda et al. 2008)͚µ?Ŀ#?̱¡Ǳȹ:ėǷ=͚jhǱȹ9Oʉʪ˳bd
!ȇƖ̙½-U2N?ǽǶ!ŋĭ-U:ʢSVUjhǱȹ?̓ȉ?ǀ́@͚Ŀ#?ɚǮ
Ƀȹ=ß̌? 9+2 Ƕ̍?ƐŞɻʟ̓͘ȉ˼ʁ͙ S<U!͚̓ȉ˼ʁ=u^~ľʮYO2<?!ȺƑ9Ur~khYěN͚jhǱȹ?ŐƜ+2ʀň͚U@Ľƚ̋§?ʀʉʪ=%UǶ̍?Ɔƚōɒɣɲ=@̉̔İ̼ň͋Ɛ̧YɅ2Ŀ#?ɣɲÇ!T͚p^YǶƜUƐŞɻ?ƺP͚ĲŻŞÄ?̝ʞ<;͚è͌ʟ(:=ȺƑɒ<Ɔ˳!˙3)V8U (Carothers& Duckett 1980, Renzaglia & Garbary 2001)
 ̍ĆģO̷͚ģź?ʡ̛?ːɓʉʪ=˵ȥ-U!̷͚ģź@͚ŷ́ɒ=Ɯ̨-U̵ģź:̖5
8͚˄+#âǐ+8Ɯ̨-U2N͚UɭżƜ̨+2̷ģź9@͚ǁ2<̍Ćģ@̷ģź?¢̛9ƆƜ)VU̍Ćģ@͜ʉʪ=˵ȥ+͚ʌʜ?âľYè%Uʉʪè˓=R58ǒë=â̛=Ƀ
,2ʉʪ!͚ĆĈʉʪ:<U͘ĩ 21a͙ĆĈʉʪ@̍Ćģ!Ɯ̨-U=ƍè˓+͚Ŝǟ̍Ćģ?̪đ̛:<UǬʉʪ (cover cell)͚4 Î¸¡?͉ȧʉʪ (neck canal cell)͚Ćʉʪ=ư-Uʯȧʉʪ͘ventral canal cell͙͚Ćʉʪ (egg) YƆƜ-U(ĩ 21b)Ǭʉʪ:͉ȧʉʪ!͚̍Ćģ?Ɯȵ̔ɭ9ŪĻ-U':9͚ʀň!ĆʉʪFï̕-U2N?̌˹!̪#͘ĩ 21c, d͙ʯȧʉʪ@Ş)#<U!͚ĐʀǌK9Ȅŋ-U':!Ur~khʕ9@̛͉YǶƜ-U͉ʉʪ!͡ê=¦C͘Ǻǀ́9͉ȧʉʪ?ĞTY 6 Î?ʉʪ!ďTŮ#͙
͚0?â̛!͚Đʀ?̴=ʀň!ĆF:ï̕-U
̌˹:<U͘ĩ 21d͙xkhʕ9@͉ʉʪ! 5 ê=¦E?!áİ9UŐƜ+2̍Ćģ@pjƆ9͚ʱSZ3ʯ̛ (venter) :̨̛͉ (neck) YO4͚ʯ̛?â̛=@͜Î?Ćʉʪ!U̍Ćģ?Ĳ̛@ȗãɺȻ?ʌʜ=ďTĨKV8U͘ĩ 21e͙'?ãɺȻ?ʌʜ@Đʀ!'U:͚̍ĆģYøL̂MˑȻ?ʌʜ=ɏ̕+͚Ñʺ˒(pseudoperianth):RAVU

M. Shimamura-18
BSJ- Revi ew 3: 101 ( 2012)

植物科学最前線 3: 102 ( 2012)ĩ  ͛̍ʀģ?Ƕ̍ ʾ̵ģƢ?ʡ̛?ʚǀ́̍ʀģ?ʚǀ́ʀĈʌʜǀ́ǂƆ?Ş)ʉʪ9ĄN
SVUʀĈʌʜǀ́͘ʀňĽƚǖ͙ʀʉʪ@ ˝Ɔ?ľƆYɧ-ĩ 20. ʀň?Ƕ̍:ƆƜ̔ɭa.ʀȇʉʪ?ƿNǂę?ʉʪè˓b. ̓ȉ͚p^ƐŞɻ?ƆƜc. ʉʪ˳?̇û:Ǯ?æ̶d. p^=ȑ52Ǯ?¿̨e. ʀň?ǹƁĩf. ƐèŶȞ͋Ɛ̧9˜ř+2r~khʀňa–e.
Carothers & Kreitner (1968), Shimamura et al. (1999)Yċʢ=Æĩ

M. Shimamura-19
BSJ- Revi ew 3: 102 ( 2012)

植物科学最前線 3: 103 ( 2012)ĩ 21. ̍Ćģ?ƆƜ̔ɭa. ʾ̍Ćģ:â̛?ĆĈʉʪ͘ɟ͈͙?ʚǀ́b. ̍Ćģ͘ĆʉʪƆƜɗƋ͙?ʚǀ́c. ̍Ćģʯ̛?ʚǀ́͘ʯȧʉʪ:͉ȧʉʪ?̇û͙
d. ̍Ćģ̛͉?Ǻǀ́͉đ͘ɟ͈͙@ 6 ʉʪê9ĨKVUe. ŐƜ+2̍Ćģ

A@1 ̵ģź=Ȍ!˟V8̵ģź?ĕʌʜ?ʉʪ!ʱƃ-U:͚0?Ĭô9̍ʀģ!ŪĻ+͚̍ʀģʭ?¡̛?ŉY̌,8ʀň!̵ģź¡=2K52Ȍ§FĤç-U͘ĩ 22a, b͙'?ŉ?ʚǀ́@\vİ9͚çđ!Ƚ#<58T͚Ǽİ?ʚǀ́Yɧ-̌ų?ȋŔŉ:@͚ƆȻ!ɋ<58
U͘ĩ 22c͙r~khɖ=@̵ģź=ǧ!<ɮOĿ!͚0VS@̵͚ģź?̍ʀģʭ?ŉS͚ɳȋ§=ʀňY 20 cm ̃#?͔)K9̽Ȼ=Ĥç+͚͍=RUʀň?ƹŰYˏ(Shimamura etal. 2008) ³?:'W͚'?R<ɳȋ§F?ʬ÷ɒ<ʀňĤç@r~kh9@ɠSV8<
r~kh@ʷɒ=@̵ģź?¡=ȦK52͚ʀňYěMȜȫ!̻Ȍ=R58@>:A)V͚Ğ
́=͎CƹU':9ʀň!ƹŰ+8U:ʢSV8UDuckett & Pressel (2009)!v
9ɝʹ+2ȜȫY̵ģƢ?¡=ȫ¢+2œ͒=RU:̵͚ģź?¡9˺>2Ȝȫ@ǒŀ 30
cm ?˷̹=͎Cƹ52ǂ9̵ģƢ SĜĎ)V2vȨȜ!͚͜ǌ̫¸â=͚ʄ
10 cm ?ŀ")?j~?ʯ́ÏÝÄ=͚ǔʅ¹ǯ?ǝ=ȑ58Ź!U':Oè 52K2͚
̷ǭYɝʹ+2Ȝȫ=Ț+2:'W͚3060 è9̷ģƢ?¡̛=K9ɝʹ+2Ȝ!ï̕+2'VS?ʎǥ@͚˃ȻÄ?ʯ́P̷ģƢ̵͚ģƢ=ŋĭ-Uǔʅ¹ǯ?ǝ!͚ǱȹÄÝÄ=ȌèY
ˏ"ȠS/U3%9<#͚Đʀ=8Oʀň?ɬ÷ʍ˹:+8ǽʬ+8UēʬƖYɧ+8

M. Shimamura-20
BSJ- Revi ew 3: 103 ( 2012)

植物科学最前線 3: 104 ( 2012)U̵ǭ?̃#9Ƀʦ-U̷ǭ?̷ģź9@͚)K*K<ɏɃȆ̲?ʪňÄ!LSV͚ǧ=̃
Ï=ǁ2=ÆSV2̍Ćģ9@͚Đʀ+2A T?O?OLSVU̍Ćģ@̷ģƢǧ!ɏ̕+8̷ģź!Ʃ4¡&SV2:Oʒʓɒ=ƆƜ)V͚Đʀ@̷ģź!Į¡ Sɶ4¡!52Ȼƚ
9Oʒʓɒ="8U':!ɧĢ)VU (Parihar 1956)ɸʣOœ̴=̷ģź?ǀ́Y˜ř+8
L2:'W̷͚ģƢ?ǧ?¹ǯǝ@̷ģź9Ǧè V+͚Î?ƪȻɴ˵â̛?¹ǯǝ:ʓ"=<58U':!è 52+2!58Î?ƪȻɴ˵?Ĳ̛?̫=Áʞ-U̍Ćģʟ@͚¥ÏY¹ǯǝ=ďTĨKV8U͘ĩ 23͙ʀň@̷ǭ?˃ȻÄ?ʙ=)ï̕-VA͚̍Ćģ?
(#̃ÒK9͚¹ǯǝ?ȉʉɻȿ˰=RTĐ÷ɒ=̒AVU?9@<3W ʀň!̍Ćģ
S?ÅS ?˧ħȹ˳=˧XV8Ȕ$Ɠ˗!U:-VA͚1 mm ¸¢?(#ɡ˷̹ O+V<ĩ 22. ʀň?Ʒça. ̵ģź?¡?Ȍ§=Ĥç+2ʀň(ɑȯ+8Ưè)b. ̍ʀģʭ?¡̛?ʀňĤçđ͘ɟą͙:Ğ́?ȋŔŉ͘ɟ͈͙̍ʀģʭ=@ėûʁ!ɏ̕+<2N͚¡̛ SLU:ȋŔ=ĨKV2͗5J͇ı:+8L:NSVUc. ʀň?Ĥçđ͘ɟą͙:ȋŔ?ʚǀ́͘ɟ͈͙
Ĥçđ?ǀ́@\vİ9͚Ǽİ?̪đ̛YO6ȋŔ
:@ƆȻ!ɋ<U

M. Shimamura-21
BSJ- Revi ew 3: 104 ( 2012)

植物科学最前線 3: 105 ( 2012)ĩ 23. ̷ģź?ƪȻɴ˵Ĳ̛?Ǻǀ́̷ģƢ?â̛=@˃ȻÄʯ́ Sʓ"=<52ǔʅ¹ǯ?ǝ!ÜT̂Z9
U̍Ćģʟ@ƪȻɴ˵F:ʓ#ǔʅ¹ǯ?ǝ=ďTĨKV8U':!è Uʀň!¹ǯǝYʍɆ+8ɬ÷-UĴĖ͚ƱŒ)VUĐʀ?ʍ˹Yɟą9ɧ-

AA3 Đʀ+2Ć@͚̍Ćģ?â̛9ʉʪè˓Y̪Ň-UK.͚¡¢ǂę=è˓+͚¡¢͝6?ʉ
ʪ!ƆƜ)VUʓ80V=ɗ±-UR<ę"9è˓!'U͘ĩ 24a͙̛͉=̃Ï?ʉʪʟ9͚¦Ťè˓!'T͚ľɇ=ư+<âÏ?ʉʪ!Ƀ,2:"=͘ĩ 24b͙͚ľÏY\
]zn_͘amphithecium)͚âÏYa}zn_ (endothecium) :RC͚t^͌9@\ ]zn_ S˅ĺ?L!Ƀ,͚a}zn_ S@ʪĈʌʜ͘ʪň:Ƅʁ͙?L!Ƀ,Ux
kh͌?ŀ̛èP͚q͌?̛9@a}zn_S͚˅?§ƒ̛?˼§͘j;columella͙:RAVUƵƩʌʜ?L!èû+͚ʪĈʌʜ@\ ]zn_=˵ȥ-U ò̅+2R=͚Đʀ!'52̍Ćģ?Ĳ̛ S@Î?̍ĆģYøL̂MR=͚Ñʺ˒(pseudoperianth)!ɏ̕-U)S=0?ľÏ S͚ʽʰ (̷ʽʰ; female involucre) 9Íˮ)VUʽʰ@͚Î?ƪȻɴ˵?Ĳ̛?̫=Áʞ-U̍ĆģʟYďTĨMR=ɏ̕-U (Parihar 1956)
̍Ćģ?ʯ̛@͚ʪňÄ?Ɯ̨=ė˩+8Ɯ̨+͚ʪňÄYɗưøMc| (calyptra) :<M. Shimamura-22
BSJ- Revi ew 3: 105 ( 2012)

植物科学最前線 3: 106 ( 2012)U6KT͚ʪňÄ@âÏ S͆=͚c|͚Ñʺ˒͚ʽʰ=RT 3 ̠=Íˮ)V8Ɯ̨U(ĩ 24c, d, 26a)c|:Ñʺ˒@͚Đʀ!'S<̮Tɏ̕+<ʿ:˃?èû!Ǉɞ<ʿ˃Ɩt^͌9@͚̍ĆģYÍˮ-U2N?˃ɒʌʜYʽ˃ (bract) :RC͚0?â̛=
U͚ĞĨ?˃!ɎĖ+8ʪňÄYøL̂Z3R=LUˑȻ?ÍˮʌʜYʺ˒ (perianth):RZ
9Ur~kh?R=͚˙ %¡˃!<˃ȻƖt^͌9@͚ėǷ?ʌʜ!ʿ˃Ɩt^
͌:@ɋ<Uɋ<UĘɫ9RAVU!͚ʽ˃:ʽʰ͚ʺ˒:Ñʺ˒@͚0V1Vɘė<ʌʜ:ʢSVU jhǱȹ9@̌ų͚ʪňÄ@¡ǂ=ę 58Ɯ̨-U!͚r~khɖ?Ŀ#?ɮ9@̍Ćģ!
̷ģź?¢̛=¢ę"=68U/9͚ʪňÄ@ƛį+͚¢ǂ=ę 58Ɯ̨-UʪňÄ@µ?t^͌:ėǷ͚˶͚ǧ͚˅͘ʪňĥ͙?͞6?̛è S9"8U͘ĩ 24d͙˶ (foot) @͚̝ÐÄ:?̎ʎ̛è9͚̝ÐÄ SʪňÄF?ǫ͏̀̈YƦ58U˶:̝ÐÄʌʜ!ư-ƯÁYʨŽ (placenta) :RC͚ʨŽ9@͚̝ÐÄ͚ʪňÄčǂ?Ï?ʉʪ!˖̸<ʥćYƆƜU'?ʥć@̝ÐÄ–ʪňÄ̫?ȹ˳̀̈:̬̎!U:ʢSV8Uǧ@˶:˅YưʓƯè9͚ʪňƹŰ?̴=¿̨Ɯ̨-U':9͚˅YÍˮģő?ľ=ƥ+ç-˅@͚͜ʉʪć?˅ĺ:0?â̛?ʪĈʌʜ (sporogenous tissue)

S<UʪĈʌʜ=@͚Ƅʁȇʉʪ (elatermother cell, elaterocyte) :ʪĈʉʪ͘sporogenous cell͙!Ś=<58èû-Ur~kh9@Ƅʁȇʉʪ@H:Z;ʉʪè˓YˏX.͚¿̨Ƀ̨?LYˏ͚Ƅʁʉʪ (elater cell) :<U͘okhɪ<;9@ƄʁȇʉʪOè˓+8Ĺõ-U͙ʪĈʉʪ@ 4 Ħ S 5 Ħ?ʉʪè˓Yˏ͚1 6?Ƅʁʉʪ=Ś+ 16−32 Î=Ĺõ+͚ȟƺè˓Yˏʪňȇʉʪ (spore mother cell, sporocyte):<U͘ĩ 24e͙r~khɖ9@͚r~khYěN8ȟƺè˓ǌ?bd?̝èPʉʪè˓˔ʞ?Ɔƚ!R#˩GSV8U(Brown et al. 2007, 2010, Shimamura et al. 2012)r~kh9@ʪňȇʉʪ@23͜6?ʹʇÄ͘˃ʖÄ͙YO4͘ĩ 25a͙͚Ǯ?è˓=Úɶ58͟6=è˓+͚Ŝǟ?
ʪň:<Uʉʪ˳͇ı=ɬ÷-UǮ?è˓@˃ʖÄ?̝è=R58S ,NȍŒ)V2ʉʪ
ǴƖ=ƍ58̏ˏ-U':!è 58U͘ĩ 25b͙t^͌9@˅?§ƒ̛=Ƅʁǝ:RAVU͚ʪňYèû/.Ƅʁ!ǝȻ=èû-U͇ı!ŋĭ-U':!U!͚r~kh?˅?â̛@ʪň:Ƅʁ!Ƿ=èŰ+͚Ƅʁǝ@˦NSV<͘ĩ 26a͙ʪň@͜6?ʪňÄ2T 300,000 Î¸¡͚͜6?̷ģź S 7,000,000 Î¸¡ɄɃ)VU:?ƱŒ!U (O’Hanlon 1926)͜6?̷
ģƢ@Ǌ

Sɩ=

%8ǒŀ 100 Îɭż?ʪňÄYʒʓɒ=ƆƜ-U:ĳĝ (Duckett &Pressel 2009) OU?9͚œ̴?͜6?̷ģƢ2T?ʪň?ɄɃ̢@O5:Ŀ O+V<ȟƺè˓ʋ¬Ƌ͚˅ĺ@ɂȻ?ʥćYƆƜ+͚ǒʋɒ=ʉʪ˳YŃ͘ĩ 26b͙Ƅʁʉʪ@ŭŮ"?ˎǃȻ?ʥćYƆƜ+͚ǒʋɒ=@ʉʪ˳YŃ͚ȃʉʪ9UƄʁ (elater) :<U͘ĩ 26c͙ʪňÄ@͚̍Ćģ=Ɇǟ-Uc|=øKV2Ȼƚ9Ɯ̨-U!͚ʪňYƹŰ-U̴=@͚ǧ!¿̨+͚˅@c|YɤT͚)S=c|Y˘Ñʺ˒:ʽʰY',̪%8͚ɳ§=̾ç-U͘ĩ 26d͙
0?Ƌ͚˅ĺ!ʪňÄ?Úɷ̛ S£˚ñ=ɤV͚Ƅʁ!«Ȥ̒÷-U':9ʪň!ɳ§=Ʒç)VU

M. Shimamura-23
BSJ- Revi ew 3: 106 ( 2012)

植物科学最前線 3: 107 ( 2012)ĩ 24. ʪňÄ?ɏɃ̔ɭa. ʩ͘4 ʉʪǖ͙?̉̔×ĐʀĆ?ǒë?è˓́͘ɟ͈͙=Ś+͚ɗ±-UʉʪǢ!ƆƜ
)VUb. a}zn_!Ƀ,2ʩĐʀĆ?ǒë?è˓́͘ɟ͈͙RT̛͉=̃Ï S˅!èû-Uc. ʩ?Ǻ
ǀ́ʩ@͚̍Ćģ?ʯ̛=Ɇǟ-Uc|͚̍Ćģ?Ĳ̛ Sɏ̕-UÑʺ˒=ďTĨKV8Ɯ̨-Ud. ˶͚ǧ͚ʪňĥ͘˅ĺ:ʪĈʌʜ͙?͞6?͇ı!èû+2ʪňÄ?ʚǀ́e. ȟƺè˓ɗò?ʪĈʌʜ?ʚǀ́Ƅʁʉʪ:ʪňȇʉʪʟ!±¯=̝ʞ-Uĩ 25. ȟƺè˓a. ʪňȇʉʪ?̉̔İ̼ň͋Ɛ̧=RU˜řP: ʹʇÄ͚N: Ǯb. ȟƺè˓ǌ?ʹʇÄ͘ʖ͙:Ǯ͙̿͘?ƫ÷ Ǯ?è˓=Úɶ4ʹʇÄ!è˓-UʹʇÄ?ː́=@-v͖͘ʹ͙!ŋĭ+͚ȟƺè˓
ǖ?ƐŞɻ͘˴͙?ƆƜ?˶Ĵ:<U

M. Shimamura-24
BSJ- Revi ew 3: 107 ( 2012)

植物科学最前線 3: 108 ( 2012)ĩ 26. ŐƜ+2ʪňÄ?Ɔƚa. ȟƺè˓!ʋ¬+2ʪňÄ?ʚǀ́b. ˅ĺ?ɂȻʥćc. ʪň:ƄʁƄʁ@ 2
̠ˎǃ?ʥćYƆƜ-Uˎǃ?ę"@Ý8ŭŮ"d. ǧ!¿̨+͚ʽʰ?ľ=̾ç+2˅AB

=. ./0#6< ǚɯ9ʊ´+2 M. polymorpha (Źʠ?r~kh)?è͌ōɒɣɲ@¨=w=èŰ-U
ǱȹYÙ=ˏXV8"2ʍʘ!T͚\o\Įı?Ǳȹ=68@͚ˤʉ<è͌ōɣɲ͚̙½ɒ˞ǣ@<)V8<ǅǚ9@͚üŬ (1987)!ʟ͐əšȰhĈ9Ư̶)V2ǸǚYO:=͚
vr~kh͘M. polymorpha var. aquatica :+8ėŒȿĭ? ssp. polymorpha =ɘƅ͙Yĳĝ+2':¸ľ=͚Źʠ?r~kh?ɮâè͌ʟ=̬-UɣɲÇ!<r~kh@è͌ōʣPƯ̶ŕ
=:58͚KT=O˻̃<͑O?9+ <2Nȓƙ!ơXV8'< 52:Ȳ@ĚN<)S=ǅǚ=@͚ǨʹÄ¡?ǮŞÄƆƜ̛Á (nucleolar organizer) ?Áʞ!wɄ
?ĕ°ɮ:@ɋ<UÎÄʟ!U':!ĳĝ)V8T (Bischler 1986)͚iǶ̍?̖!
U':!ɧĢ)V8U M. polymorpha ssp. montivagans =̬+8@͚w¸ľ9O͚\ c (Perold 1999)͚üɼ͘Schuster 1992͙
͚n\ (Konstantinova & Bakalin 2009)͚̈́ī (Choi et al. 2010)͚<;͔ʘżU
@͔Ǹ͔?ĮıY§ƒ=¤ɇĕĮ SɃʦ!ĳĝ)V8T͚ǅǚīâ9Oǁ2<˩ǪPǸǚ?äǲˡ!ǕKVU͞6?°ɮ̫?0V1V?§̫ɒ<Ɔƚ?Ǳȹ!ŋĭ-U':@͚è͌ōɒ<˞̟!ɣɲʣ=R58ɋ<UĈħ:O<58"2ÇAȤȬ<ɂĸ9Ƀʦ+8Ur~kh@vr~kh=À2ƆƚYɧ-':!U͘üŬ 1987, Schuster 1992͙
³Ƌ?˞ǣ9͞6?
°ɮP0?§̫İ?Ǳȹ?̙½ɒœÄ͚Ɔƚ?ēĶƖ?ŵ!ǇS =<U':YǖƊ+2

M. Shimamura-25
BSJ- Revi ew 3: 108 ( 2012)

植物科学最前線 3: 109 ( 2012)

 r~kh?ľ̛Ɔƚ=̬+8@ 200 Ÿ¸¡=X2Uɣɲ?ȂĔ?§9͚ˤʉ=˜řˢ̅)V
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ƆƚƆ˳:+8?Lȓɖ)V82jhǱȹ?ľ̛Ɔƚ?ĿǷƖ=68͚àÄɒ<ʌʜ?ɘėƖY̙½ň
ɏȿ9ǲˣ+͚̏ûɏɃōɒ˜Ȳ9ɣɲY̏NU':!9"U O+V<,"*
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X_TR³ºª§ďƣ¸ãz~ŝǈċİÝæÝæǭřæƂůŸŴśŒŴæðĠ
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Cell biology in liverwort
Key words: Cytoskeleton, Membrane traffic, Organelle, Rab5, VAMP727
Atsuko Era & Takashi Ueda
Department of Biological Sciences, Graduate School of Science, The University of Tokyo
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fpZtHbMK7DaGVfd%,űǀ=$ ( +&:ȃ+ǦĎǥ«+Ĝ¤,ȃǰ~ķŒ+Ǜµ+ǝŵ)$ȃOWķŒªõą)ŶãķŒ+§ǘų¢%Ǌ!6+&č?=$ ȃT`MK+ EST ƺı)9;ȃǦĎĜ¤ǥ«@ĩ<
MpVAMP727 T`MK)äÐ<&Ħ:&(!+&:ȃVAMP72 /+ǦĎǥ«+Ĝ¤,ȃLKķŒ+ªõ±)Ǌ!&ȃfpZtHbMK7OW,ŀŨ) VAMP727 Ò
VAMP72 @ß!&ŲÇ=<%, MpVAMP727 & VAMP727 ,9 (ǐǕƈǌ%ĿƜ
$ <+ȃ=,ą+ůŸǃǹ% <ÖX_TR@JPZk¸¶ ķŒƅƛ)$ȃFuHbs+³@¯ć$ <+, )BIXyƒƎ% <& ?=$ <2ȃƅƛ+ĀĒĀēȃƅƛªƲȃ¢żēǩ('ȃBIXyƒƎ+ǫ?<Řǆ,Üõ)ŋ;ȃBIX
y+ě³ƅƛ+śÅŇ³) <āǶ,ǵø)Ý (Hasezawa & Kumagai 2002, Hussey et al. 2006,
Tijs Ketelaar 2001)Ĕ,ȃLifeact (Riedl et al. 2008)&

iw{h@Ŝ $ȃT`MK+BIXyƒ

Ǝ+sDhDp{PyJ@ƾ3 (Era et al. 2009)+ƉĲȃBIXyƒƎĮ+Þ)Č$Ǚ
ü@Ú ""ō;ǜ³$ <ļãƹî=ȁÍ 4Ȃǜ³Ǚü,ȃƅ BIXyƒƎ1'Ý!¿ļ+BIXyƒƎ+Þ)9<çéµ,ȃin vitro Ƃ7OwDa_R_+ƚǎƅƛ)ÕÂ=$ < (Michelot et al. 2007, Staiger et al. 2009)ķŒƅƛ)$6$ȃBIXyƒƎ,nFOy)äņė ǜ³@<&ÕÂ=$ ;ȃT`MK%6+ņė ǜ³ƹî=

A. Era & T. Ueda-3
BSJ- Revi ew 3: 116 ( 2012)

植物科学最前線 3: 117 ( 2012) $%# >CE{Áy2G8X_TR@JPZk¸¶+|A  ¯/1>3
8¬qI6G7GÃÒ·ÒÏ;¿®3Ì?}H58B@+Bar =20 µm
ÐÑJPZk¸¶@¥E
Éx+Bar = 2 µmdMU
k # ÍwÐ ÑÒ²Ì}ÍwÐ"'(ÑÒ²ywÐ$&Ñs©Ò  |<~ >¿®+Bar = 5µm

ņė ǜ³&¿ļȃō;ǜ³62nFOy)ä$ <&ȃ2ȃĈñž@ŰÙȃ2,çéµ$6ȃō;ǜ³Ł2<&,(ȃ4>ǜ³ǙüØ&6Ħ:&(!ȁEra et al. inpressȂ
+T`MKr`{I(BIXyƒƎ+ě³,ȃT`MKŕƠ)ŧǞƅƛ¨Ň³¯ćOQ[o(+6=( ×aL]Ofghl ,X_TRMiN`g, Ĕ,ŘÐȃ½ŶFuHbsm{G{@Ŝ ȃT`MK+ļ(FuHbs@¼Ƹµ<&@ƾ
3$ <( ƹî,ȃķƌą 5 ĥŪ+ǩăƃ 5 mm +ƨŔ¥@QsD^JsQ)ȃ+ư
ũƅƛ@ïǆ&$ <+ƉĲ+ "@)Ɔ$Ƿ 

×*Óºp<TiVp ñƛ+¼Ƹµ),ȃƭ£
VyeIǉ)OJ_ukiX^&ñƛóÐOJ_u@6+ȁSP-GFP-HDELȂ@Ŝ ñƛ,ȃ+³ķŒ¿ļ)O{]ŔĻǚ&Ə
ŪŔĻǚ%Ļē=$ <2ȃĵƟ:Ļǚ6ƹî
=ñƛ¥ň=<9 )³ $ ;ȃƋ +ľļ,Úµ$ Mu
P@ syalyl transferase (ST)@Ŝ $¼Ƹµ&>ȃ^Y]Ŕ+ĻǚñƛǑ)ƹî=Bar = 20 µm

A. Era & T. Ueda-4
BSJ- Revi ew 3: 117 ( 2012)

植物科学最前線 3: 118 ( 2012)×*ÔTiVp<]gkWTiV`[]jlP
 TGN +¼Ƹµ),ȃ
OwDa_R_)$TGN )óÐ<&ŭ:=$ <Q-SNAREȃ
SYP61+ǥ«@6&)T`MK+ EST @ĸƄȃĆ:= MpSYP6 )ƭ£VyeIǉ@Ʈ¾6+@ŜST )9;¼Ƹµ=MuP&ȃTGN % <&ĪĄ=< MpSYP6 Lye{]py],ȃ
ǲĝ""ūŜ<9 (³@ƷBar = 20 µm

×*Õ³º¼ OwDa_R_)$ƅƛƟ)óÐ<&ŭ:=$ <ȃQ-SNAREȃSYP11ȃSYP12ȃ
SYP13 +ǥ«@6&)T`MK+ EST @ĸƄȃĆ:= MpSYP1 )ƭ£VyeIǉ@Ʈ¾ȃT`MK+ƅƛƟ@¼ƸµBar = 20µm

×*Ö£º¼
 OwDa_R_)$ŉƛƟ)óÐ<&ŭ:=$< Q-SNAREȃSYP2 +ǥ«@6&)ȃ
T`MK+ EST @ĸƄȃĆ:= MpSYP2 )ƭ£Vye
Iǉ@Ʈ¾ȃT`MK+ŉƛƟ@¼ƸµT`MK+
ŉƛ,ƨƐ+Ǒ2%ƅ
ƛ !. )û!$ <ŉA. Era & T. Ueda-5
BSJ- Revi ew 3: 118 ( 2012)

植物科学最前線 3: 119 ( 2012)

ƛ@Ǉ¹ĀǉƁ,ƹî=( 2ȃƅƛưôǑ+ŉƛƟ,ǵø)ƴǴ(ĀŔ@ĩ$ <Bar = 20 µm


×*×JPZk¸¶<£º¼ ¢+ Lifeact )9;BIXyƒƎ¼Ƹµ=ƅƛ)$ȃFM1-43 &Ey^ND]{OQ+ǔǋƾƪ)9<ŉƛƟ@ĳƤŉƛƟȃƅƛƟȃƨƐ+Ǫ+Ŗ ŹǪ)BIXyƒƎþ;÷:=$ <ļãª<Bar = 20 µm

×*ØJPZk¸¶<d]Sk^hJ ¢+ Lifeact )9;BIXyƒƎ¼Ƹµ=ƅƛ) ï  $ ȃ Mito
Tracker )9<n]
Ly^tB+ĳƤ@Ư!BIXyƒƎ)Ņ!ǥÀ@Ųn]Ly^tBÜĢƹî=Bar = 20 µmØÎ )X_TR³ºª§@n)
 ÏƥìǞ:+â?=ƘȃT`MKĔ+ůŸë)$6ȃOwDa_R_+Ǩ) "+Ǫ)

T`MKś $ <(A$&6 <ǡȀǡȀ&º;ǮǱ)ȃǪǟ!$OwDa_R_+Ĥ@Ę $2 9 (Ǐ,ňŮ) ( ĀĒ 2;)6Ť(!$ <:śŒ+ĀĒ&,"2;ȃśŒ+Ôĭ·% <ƅƛŷ3ǧ(!$Ū)Ʒ $ <Ā% <ġ)ƅƛ¿%
6ŷ3ǧ(;ĤŤ(=-ȃĀĒ6Ť(!$<ƽ% <êǱ),ĀĒ+Ť(<­Ŷ+śŒ%,ȃ+ƅƛ+ļã2%6Ť(!$ <6+@"<ƅƛ@:)Āǚ<ǠãĐÕŤ

A. Era & T. Ueda-6
BSJ- Revi ew 3: 119 ( 2012)

植物科学最前線 3: 120 ( 2012)(<+:ȃ=,ÿő(+¹ĵƅƛ&Ŭĵƅƛ+ǟ 7ȃ³Œƅƛ&ķŒƅƛ+ǟ ȃ< ,§ǘŏ,ǂ6ŭ<&>> %,T`MK&OwDa_R_+ƅƛ+ǟ ,ȅ4 ¡ù~±)äÐ§ǘų¢:ȃƘ),'+9 (ÜļĎś2=$ !+> $+ÜļĎ,)ǊÊȃ'+9 (vju+ÜļĎ/&Ɠ!$ +2ȃƫǺ&T`MK+ƅƛ&+ǟ ,ȅT`MKƅƛ+'K&!œĉķŒ+ǰ~µ)ĩ®),: +T`MǛµŨ)ƢÃŊ śŒ@Ŝ ƅƛśŒæ+ŸĹŪĽ&,ȃ"2<&>

Èǹ@ƺĦ&% <&č!$ < ƅƛśŒæ+Ôĭ,ƅƛ@ƹ<&% <Ĕ,T`MKƅƛ+FuHbs7ƅƛǾĶ@ƭ£V
yeIǉ)9;¼Ƹµ$ƹî@Ư!$ <T`MK6ķŒ% <:ȃÿő(:ķŒƅƛ+Ě"0FuHbs@¥$$ <ñƛ7MuPȃTGN ('+FuHbs,ȃOwDa_R_+=:&ĺ*¿ļ+ĀĒ@&!$ <9 % <ŘÐ2%+Ĕ+ƹî)9;¬ĦT`M
Kƅƛ+ĨÝ+œĉ,ȃƅƛ !. )û!ŉƛƟ)9;Ŗ5:=ŹǪ@ȃǵø)r`{I(³%ǼÉ<BIXyƒƎ> !œĉ,īƼǓ+ƀÌ@ƞ$ ( ȃ+BI

XyƒƎ+³+Ă²@ą,ŭ; 6+% < ¢)Ǔ09 )ȃFuHbs+BD\y[C[C{,Ɵǘ)ǫ?<ªã%Ń2<8 )śŒ
+Ǡã@Ǆ0ȃĕ<|Ɩ+śŒĚ"Ɵǘ¯ćªã@Ʒ"&:ȃ+śŒƖ)œĉ
Ũ(Ɵǘƈǌ7FuHbsĿƜ <> &Ō%<ƽ% <Ĕ,ǰ~ķŒ)ûä=$ < ARA6 Ò RAB5 +ĿƜ@OwDa_R_&T`MK%Ǆ0ȃǰ~ķŒœŤŨ(Ɵǘƈǌ@
ŧƷ%<+%,( &Ɨ ȃ' 6T`MK+ ARA6 Ò RAB5 ,OwDa_R_ ARA6&,Ť(<ƈǌ%,: $ <9 % < 6ûä=$ < RAB (+%ȃǰ~ķŒ§ǘ+ǧƵ(Ă²@ę!$ <&č!$ 6+:ȃ=,ǽ% <ARA6 Ò RAB5 +ĿƜķŒ+½ƂƊ%'+ŵüÜļµ$ <+,īĦ% ;ȃǵø)ƢÃŊ Èǹ% < ¿ļ+Ɨ Ĥ:ȃŶãķŒ)äÐ( &č?=$  VAMP727 ,ȃŶãķŒœŤŨ(Ɵǘƈǌ%,: $ <> &ĞŌ=$ +ŶãķŒ)äÐ( &,ȃfpZtHbMK7DaGVfd,Ě!$ ( &±ğĐÕ)Ô#6+%ȃÉĔT`

MK+ EST @Ǆ0ƉĲȃT`MK), VAMP727 ļªãäÐ<&¬Ħȃ+±ğ,ƶ!
fpZtHbMK)Ő :& !$ȃLKķŒ)Ő &ƻ!$,(:( &!$ȃOwDa_R_)$ VAMP727 ,ȃŉƛ±¶Š&

&@č ŭ

FuHbs&ŉƛ7ƅƛƟ&+Ʈ¾ħ),:ȃŶãĀē7Q]vQƙĎ)ǫ$ <&Ħ:&(!$ < (Ebine et al.
2011, Ebine et al. 2008)T`MK+ VAMP727 ,OwDa_R_+ VAMP727 &Ť(<ĿƜ@Ě!$<+> T`MK,Ŷã@"( +%ȃÿőǿŀĿƜŧŘ) <Ă²,Ť(!$<>ȃƅƛśŒæŨvju%,' > ŘÐůŸ% < ŘÐĔº;ƇA% <T`MKƅƛśŒæ,ȃ'=622ůŸǗ) <=%6Ɵǘƈǌ7ƅƛǾĶ³Ē+ÜļĎƍ&Ħ:&(!$=:T`MKƅƛ%Ʒ:=<œĉ
T`MK+ĀĒ7śĒ)'+9 )í$ <+@źŁ5:==-ȃǰ~ķŒ+śäĖţ,'+9 )Üļµ$ !+& !ŦÈƺŃ/+Ý(fy]&(<> 

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植物科学最前線 3: 121 ( 2012)ÂÇ ĭůŸ@Ǜ5<) ;ȃ ƿ)(;2ǤÝæ+ń¨å¢śȃŮö¦ý¢śȃǑťÝæ+ÝÆ´ú¢ś)+Ö@;$đǅş~2«¨
Dacks, J.B., & Field, M. C. 2007. Evolution of the eukaryotic membrane-trafficking system: origin, tempo andmode. J Cell Sci. 120: 2977-2985.
Ebine, K., Fujimoto, M., Okatani, Y., Nishiyama, T., Goh, T., Ito, E., Dainobu, T., Nishitani, A., Uemura, T., Sato,
M.H., Thordal-Christensen, H., Tsutsumi, N., Nakano, A., & Ueda, T. 2011. A membrane trafficking pathway
regulated by the plant-specific RAB GTPase ARA6. Nat Cell Biol. 13: 853-859.
Ebine, K., Okatani, Y., Uemura, T., Goh, T., Shoda, K., Niihama, M., Morita, M. T., Spitzer, C., Otegui, M. S.,
Nakano, A., & Ueda, T. 2008. A SNARE complex unique to seed plants is required for protein storage vacuole
biogenesis and seed development of Arabidopsis thaliana. Plant Cell. 20: 3006-3021.
Era, A., Kutsuna, N., Higaki, T., Hasezawa, S., Nakano, A., & Ueda, T. 2012. Microtubule stability affects theunique motility of F-actin in Marchantia polymorpha. J. Plant Res. in press.
Era, A., Tominaga, M., Ebine, K., Awai, C., Saito, C., Ishizaki, K., Yamato, K. T., Kohchi, T., Nakano, A., & Ueda,
T. 2009. Application of Lifeact reveals F-actin dynamics in Arabidopsis thaliana and the liverwort, Marchantia
polymorpha. Plant Cell Physiol. 50: 1041-1048.
Grosshans, B. L., Ortiz, D., & Novick, P. 2006. Rabs and their effectors: Achieving specificity in membrane traffic.
Proc Natl Acad Sci U S A. 103: 11821-11827.
Hasezawa, S., & Kumagai, F. 2002. Dynamic changes and the role of the cytoskeleton during the cell cycle inhigher plant cells. Int Rev Cytol. 214: 161-191.
Hussey, P. J., Ketelaar, T., & Deeks, M. J. 2006. Control of the actin cytoskeleton in plant cell growth. Annu Rev
Plant Biol. 57: 109-125.
Michelot, A., Berro, J., Guerin, C., Boujemaa-Paterski, R., Staiger, C. J., Martiel, J. L., & Blanchoin, L. 2007.
Actin-filament stochastic dynamics mediated by ADF/cofilin. Curr Biol. 17: 825-833.
Riedl, J., Crevenna, A. H., Kessenbrock, K., Yu J. H., Neukirchen, D., Bista, M., Bradke, F., Jenne, D., Holak, T. A.,
Werb, Z., Sixt, M., & Wedlich-Soldner, R. 2008. Lifeact: a versatile marker to visualize F-actin. Nat Methods. 5:605-607.
Staiger, C. J., Sheahan, M. B., Khurana, P., Wang, X., McCurdy, D. W. & Blanchoin, L. 2009. Actin filament
dynamics are dominated by rapid growth and severing activity in the Arabidopsis cortical array. J Cell Biol. 184:269-280.
Tijs Ketelaar, A. M. C. E. 2001. The cytoskeleton in plant cell growth: lessons from root hairs. New Phytologist.
152: 409-418.
Ueda, T., Yamaguchi, M., Uchimiya, H., & Nakano, A. 2001. Ara6, a plant-unique novel type Rab GTPase,functions in the endocytic pathway of Arabidopsis thaliana. EMBO J. 20: 4730-4741.

A. Era & T. Ueda-8
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植物科学最前線 3: 122 ( 2012)29 %1.  7 #ůł×ư
Îō×ß×ßǏŏïßšūŦ{ŐÐŦßšūSzU| 2ðĭ³×ǢǥǥThe role of abscisic acid in stress response and dormancy of liverworts
Key words: abscisic acid; desiccation stress; protein phosphatase 2C.
Daisuke Takezawa
Graduate School of Science and Engineering; Institute for Environmental Science andTechnology, Saitama University
Sakura-ku, Saitama, 338-8570, Japan;:  BhPPzǁǞABAǟ+SQGZwkz²»ň%ǠĮň%+8ş9<;Įňlwrz*}#% ;ũÛĮň) $+ǠABA ǋ; #*ŉĀŗ(ŎƦ :ǠũÛ*Ş7
ĹQ\xQ)ĂŲĸÜǈǆ.*ǋ('*% ;ƲòǠ\DryMP7PyC`^
R^*ÔŔĪƝħ9ǠABA ő»ĉŹƬ7Ǡ¹ã&ĄÏƺ)ǋ>;¥ÛqE_Rp*ƝěƷ@% ;ǞCutler et al. 2010ǟ
ABA +Įň)ŉĞ*²»ň%+(ǠƑƒ7ŽƒǘǠƌǘǠB
fMzixIOǠýő°ň)6į¤<Ǡ22(őŏŒ?Ď#&ŤÃ<$;ǞHartung 2010ǟǠ<9*őň¼ ABA ő»ĉŹƬ7¹ãĴĲ?Ď"$ ;'+ě% ; "1 ǠMKǠPV?¾422(ǐ~ĮňǞEmbryophytaǟ*Jw|i) $ǠABA Ň7ĿƴÇQ\xQĂŲ) $Ĩû¬ě9&("$:ǠABA
ĂŲ)ǋ>;ǐ~ĮňƵ*qE_Rp*ÝÈŤÃ<;ƲòǠeqXvFaNKǞPhyscomitrella patensǟ7T_NKǞMarchantia polymorphaǟ*¥ÛƝħ&ƼÛŢÒċƖ*ǉŖƷ4)#<Ǡǐ~Įň)Ý<; ABA ĂŲ)ǋ>;ƼÛſ&ǠĮň*ƻĂƷ²)
;<9*û¬ě9)(;&ğü<$ ;ǞTakezawa et al. 2011ǟ<:() 86% 



#

 22(ǐ~Įň) $ ABA ǋ>;őŏŗiySQ+Öí)>;Ǡ*)+ũÛ*ĉņǠŞǠĸÜǈǆǠƈƉùĉǍâ('ǠǝűĮň)ŉĀŗ(iySQ&Ǡŷƅ*¥Ƙ{ǇǍâ7Ł{ŇƃăƷ('MKĮň?¾5ǐ~Įň)Ƶ*iySQÝÈ;Ɠǘ7Ɗǘ('*MKĮň+ǠőÁľ°)āǕ(Ĺ*¨Œ)ǋ$ǠżųģĮň&+Ŕ(;qE_Rp)8:ŐÐ)ƻĂ$ ;ĹķƯŗé( ŐÐ) $ǠżųģĮň+Ĭ9¿¸Ĺ?ĸÜ9*ƏĖ?©ÿ;&)8:Ĺ*ēØ?ǌ<&ķ/$ǠƗŘŵ?Ď(MKĮň*ĩǙŸž+ǠĹ*ēØ?ǌ&%( >:)Ǡŷƅxjw%*ǝ ƇĹƃă?Ď!ǠŇŊć) $6ŷƅ*â?Äƽ;ĴĲ?Ď#&Ɓ9<;MKĮň**8

D. Takezawa-1
BSJ- Revi ew 3: 122 ( 2012)

植物科学最前線 3: 123 ( 2012)(ăƪ+ťŗĮň*ǐ~%*ƻĂ)ǂƚ(ùƪ% "&Ɓ9<ǞOliver et al. 2005ǟ
Ǡǐ~Įň*Ʒ²ƹŨ%ĝ6§ğ)¥íJw|i% ;MKĮňǝ Ňƃă?Ď#&á*&?ĕĎ$ ;ǞÆǡǟ ŰƂ+MKĮň*Ň{Ňƃă); ABA *û¬?ě9);&)8:ǠĮň*ǐ~ƻĂ)āǕ% " ABA ĂŲ*·ÊŗĴĲ?ƝěǠ*ý*Ʒ²ƹŨ%Öĳ²żųģ
Įň* ABA ĂŲĴĲ)# $6ŏƝ?ŀ5 &Ɓ$ ;%Ǡr[wĮň&$*¥ÛƝħÍřė<## ;eqXvFaNK7T_NK?Œ ǠABA )8:÷ƫ<;ƃă*Ô²&ǠABA ĂŲ*¥ÛĴĲ)ǋ$Ɲħ?ƕ("$ ;

=:()

 %1 MKĮň* ABA +Ǡ)rbIy|^wč?Œ Ċļ)8:ƓǘǠƊǘ8-XbNK
ǘ*Ŕ(;ũ9į¤<$ ;ǞHartung et al. 1987, Werner et al. 1991ǟ
ƅÛ)ĸÜ?Ď#Ɠ
ǘ&XbNKǘ%+Ǡ
ABA ĸÜ*ǈǆ?Ʒ;&etDUzNKǞFunaria hygrometricaǟ
7^FOGXbNKǞAnthoceros punctatusǟ) $ÏÀ<$ ;ǞGarner and Paolillo 1973,
Hartung et al. 1987ǟ
2ǠƓǘ)# $+Ǡ·Ŵ) ABA ;őŏŒ)# $ǠķƯŗÖ*šūÏÀ ;ǞÆǢǟ
Ɠǘ*·Ŵŷƅ? ABA £ŏ;&Ǡ*ĉǇČ©<ǠŅăőĶùć% ; brood cell Ƣç<;2ǠABA +·Ŵ9Ƌƍ.*¥²?Ǎâ;
&6ÏÀ<$ ;ǞValadon and Mummery 1971, Menon and Lal 1974, Goode et al. 1993, Chopra
and Kapur 1989ǟ
etDUzNK%+ǠABA £ŏ·Ŵŷƅ*Ňƃă?Ƣç;&ě
9&("$:ǠŇQ\xQ ő ABA Ǆ?Ñ®;ǞWerner et al. 1991ǟ
¼etDUzNKŦ)ì;eqXvFaNK) $6Ǡ~Ɵ*8 ( ABA £ŏ)8;ùćÔ²7Ň{¢źQ\xQƃă*Ñ×ÏÀ<$ ;ǞGoode et al. 1993, Minami et al. 2003ǟ
Lbp
Ɲƣ<eqXvFaNK%+Ǡ9) ABA ĂŲƼÛ*ŖŎiygACwƝħ7ǠĂŲ
)ǋ>;PJ^wÅÛ*ĴƆƝħƕ(><$ ;ǞCuming et al. 2007, Komatsu et al. 2009,

D. Takezawa-2
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植物科学最前線 3: 124 ( 2012)

Chater et al. 2011ǟ <)æǠƊǘ%+ǠABA )8"$÷ƫ<;őŏĂŲ)ǋ$þ9<$ ;şƛ+é( , Hellwege 8- Hartung 9*Ǜŗ(šū)8:ǠT_NKŚǞMarchatialesǟ)ì;ÙŇăƊǘ Exormotheca holstii &Ĺő*ƊǘDGNKǞRiccia fluitansǟ) $Ǡ ő
ABA ƐŪ&Ňƃă*ǋƶě9)<$ ;ǞHellwege et al. 1994, 1996ǟ
2ǠDGNK)$+ǠABA £ŏŉŔŗ)Ƣç<; LEA ĳUzdIƪ)#$6ÏÀ<$;

ǞHellwege et al. 1996ǟ
T_NKŚÕ%6ǠgUnUNKŚ)ì;scRNKǞPallavicinia
lyelliiǟ)æǠABA +Ňƃă?Ƣç;&ÏÀ<$ ;ǞPence et al. 2005ǟ
}Ě%ǠT_NK)æ$+ǠABA +Ňƃă?Ƣç( &*źĨ6þ9<$ ;ǞPence et al. 2005ǟ

PryceǞ1972ǟ+ǠƊǘ) $ ABA ¤<;é( &9ǠƊǘ) $ ABA *û¬+ǎàŗ% :Ǡw`uvzǁǞLNAǟǞÆǣǟ*8 (Ɗǘ)ô¥ñ;²»ň ABA
ĳ*?$ ;ºƆă?ďĔǞPryce 1972ǟ
LNA +oEYGT_NKǞLunularia cruciataǟ
*ŞƢç&ǋƶ$ƐŪ;ňƪ&$µǑ<ǠT_NKŅăƉ7xUQ*ƄƮĉǇǠIx
QũÛ*ŖƉ?Č©;&ÏÀ<$ ;0ǠŽƒIyxu*ƃ¢ă?Ñ×;&
ÏÀ6 ;ǞValio and Schwabe 1970ǟǠLNA  ABA &¼ĳ*ŒĴĲ)8:Įň)
Œ$ ;' )# $+ě% ;

D. Takezawa-3
BSJ- Revi ew 3: 124 ( 2012)

植物科学最前線 3: 125 ( 2012)=:,  %1 5+) -/0 ŰƂ9+ǠƊǘ ABA ĂŲ*¥ÛĴĲ?ě9);5ǠT_NK? ABA ĂŲšū*r[
wŵ&$ţŭ;&?Śďáǜ?Ʒ5$ŅƌÌǙĤ) $ ABA *úǔ?Ƥ/&=ǠABA +T_NKŅăƉ*ĉǇ?Č©Ǡ¢ź7Ň)æ;ƃă*Ñ×?Ʒ;
&ě9&("ǞTougane et al. 2010, Akter et al. 2011ǟ
2ǠT_NKÌǙŷƅ) $ABA +MpHũÛ* ABA ƢçăƼÛ Em *iyr|U|©ÿ) ; GUS ƼÛǞEm-GUSǟ
?ŃõÝŗ)ľă²*&9ǠŰƂ9+T_NK)6ǝűĮň&Ƶ* ABA ĂŲ)āƚ(ŷƅ ÅÛÝÈ;&ƁǠƿ×ß*ĺ ǠŠîǠƲŕ×ß*×Â9*´­?þ$
T_NK ABA ĂŲǋƶƼÛ*ĐŶ?Ơ3
 ŰƂ9+ǠT_NK EST ǀ¦ĄÏ9PyC`^R^ ABSCISIC ACID INSENSITIVE1 (ABI1)*
lryJ?ƛ

ǠMpABI1 &½PyC`^R^ ABI1 +iyZCzlQgAU|T2CǞPP2Cǟ?M|]ǠŉŔŗ( Gly  Asp )Ô² abi1-1 ÔŔ+ ABA ǒĆ¹ăƗŎÊ?
Ť&ş9<$ ;PyC`^R^)+ ABI1 ?¾4ǦƼÛJw|i A PP2C ?M|]Ǡ<9 PP2C + ABA PJ^wƺ*Ƨ*©ÿÅÛ% ;&Ɓ9<$ ;ĝƲǠJ
w|i A PP2C ŷƅ

ABA ¹ã PYR/PYL/RCAR &śđŜŒǠiyZCzG^|T

SnRK2 *Ƈvzǁ²?$ ABA PJ^w?Ƨ)©ÿ$ ;&ě9&("ǞPark et
al. 2009ǟ
 T_NK MpABI1 +PyC`^R^*Jw|i A PP2C & C ġŮ*ƞÚ]qCz) $Ŝ¼
ă?Ď!ǞÆǤǟ
ǠŵŻƝħ96Jw|i A )ì;&Ɓ9< "1 ǠN ġŮ*]q
Cz+PyC`^R^ PP2C &*Ŝ¼ă+("ǠīŧƕPJ^w?Ď"$:ǠGFP Ɣ»UzdIƪ)8;êÈƝħáǜ96 MpABI1 *īêÈ)ä;&Ť<
 T_NKŷƅ*}ƹŗŖŎBWSC%+ǠMpABI1 *ƹ«ŖŎ ABA ĂŲŗ( Em-GUS ƼÛ*ŖŎ?øČ©2ǠMpABI1 ƼÛ?ƹ«ŖŎ;eqXvFaNKùƪƭĒ?
¤&=Ǡ*·Ŵŷƅ+ǗƎ( ABA ǒĆ¹ă?Ť(>!ǠǃőĪ) $ABA )8:Ōþ<;¢źƃă7ǝĿƴÇƃă MpABI1 ƹ«ŖŎĪ%+Ɯå<ǠABA Ƣçŗ( LEA UzdIƪ*ƐŪ60&@'ƛ9<( &ě9&("ǞTougane et al. 2010ǟ


D. Takezawa-4
BSJ- Revi ew 3: 125 ( 2012)

植物科学最前線 3: 126 ( 2012) 2ǠMpABI1 ƹ«ŖŎĪ+ ABA ĂŲ %(Łǚ²Ɔ)6ĵē ;&ě9&(:ǠABA ĄÏƺŹƬĮň*ŁĂŲ)ǋ$ ;&ŤÃ<ǞBhyan et al. 2012ǟ
<9*źĨ9ǠƊǘ)6 PP2C ? ABA ĂŲ©ÿĴĲÝÈǠABA ĂŲ)ǋ>;Í
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ABAƇĹ7Ň('*Q\xQƃă) $ĴƆ;ǐ~Įň)ĜƸŗ(²»ň% ;&ŤÃ<Ɗǘ* ABA ĂŲ)ǋ>;ƭ¡ÅÛ7ƃăǋƶƼÛ*ŖŎ©ÿĴĲ)# $*ơŷ+Ġ ě% :ǠªƳ*}ƹŗŖŎŵ7ǠIshizaki (2008)9)8;ùƪƭĒļ)8:Ǡšū*
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ħ;&%<,ǠABA *őŏßŗąƀ?ě9);&&6)ǠABA ĂŲ*·ÊŗĴĲ*ıƚ?ě9)%;&Ɓ$ ;

?:34
 Ģšū+ǠĘƾŦßŝŦßšūƩƙ¯ǅ{Ířšū(C)8-ęßƖǖËšūĮňŐÐŬŢ
­ *¯ĉ)8:Ʒ59<T_NK*ÌǙ8-ƼÛƝħ)# $+ƿ×ß*ĺ Þ¶Ó&Šîö¶ÓǠƲŕ×ß*×Â±ó¶Ó)8;Ǔŗ(Om|\?þ}ƹŗŖŎŵ)8; PP2C *ĴƆƝħ+ĥƱİ×ß*ÉœĽ}¶ÓǠèĦĈĻ¶Ó*ƨŋ)8:§5$ºƆ)(")ŀĆƥ;

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and desiccation tolerance in the liverwort Marchantia polymorpha. Cryobiol. Cryotechnol. 57: 83-86.
Bhyan, S. B., Minami, A., Kaneko, Y., Suzuki, S., Arakawa, K., Sakata, Y., & Takezawa, D. 2012. Coldacclimation in the moss Physcomitrella patens involves abscisic acid-dependent signaling. J. Plant
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An analysis of MpCLE genes in Marchantia polymorpha
Key words: CLE, Hormone, Peptide, Development, Marchantia polymorpha
Shinichiro Sawa, Hiroaki Honda, Ryo Tabata
Kumamoto University
Kurokami 2-39-1, Kumamoto, 860-8555, Japan
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Betsuyaku, S., Sawa, S., & Yamada, M. 2011a. The function of the CLE peptide in plant development
and symbiosis. The Arabidopsis Book. 9: e0149
Betsuyaku, S., Takahashi, F., Kinoshita, A., Miwa, H., Shinozaki, K., Fukuda, H., & Sawa, S.2011b.
Mitogen-Activated Protein Kinase regulated by the CLAVATA receptors contributes to the shootapical meristem homeostasis. Plant Cell Physiol. 52: 14-29.
Ito, Y., Nakanomyo, I., Motose, H., Iwamoto, K., Sawa, S., Dohmae, N., & Fukuda, H. 2006.
Dodeca-CLE peptides as suppressors of plant stem cell. Science 313: 842-845.
Kinoshita, A., Betsuyaku, S., Osakabe, Y., Mizuno, S., Nagawa, S., Stahl, Y., Simon, R.,
Yamaguchi-Shinozaki, K., Fukuda, H., & Sawa, S. 2010. RPK2/TOAD2 is an essential
receptor-like kinase transmitting the CLV3 signalling in Arabidopsis. Development 137:
3911-3920.
Miwa, H., Kinoshita, A., Fukuda, H., & Sawa S. 2009a. Plant meristems: CLV3/ESR-related signaling
in the shoot apical meristem and the root apical meristem. J. Plant Res. 122: 31-39.
Miwa, H., Tamaki, T., Fukuda, H., & Sawa, S. 2009b. Evolution of CLE signaling. Plant Signaling &Behavior 4: 477-481.

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植物科学最前線 3: 133 ( 2012)

Nardmann, J., Reisewitz, P., & Werr, W. 2009. Discrete Shoot and Root Stem Cell-Promoting
WUS/WOX5 Functions Are an Evolutionary Innovation of Angiosperms. Mol Biol Evol. 26:
1745-1755.
Oelkers, K., Goffard, N., Weiller, G., F., Gresshoff, P., M., Mathesius, U., & Frickey, Y. 2008.
Bioinformatic analysis of the CLE signaling peptide family. BMC Plant Biology 8:1.
doi:10.1186/1471-2229-8-1
Ohyama, K., Shinohara, H., Ohnishi, M. O., & Matsubayashi, Y. 2009. A Glycopeptide regulatingstem cell fate in Arabidopsis thaliana. Nature Chem. Biol. 5: 578-580.
Sawa, S., Kinoshita, A., Betsuyaku, S., & Fukuda, H. 2008. A large family of genes that sharehomology with CLE domain in Arabidopsis and rice. Plant Signaling & Behavior 3: 1-3.

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植物科学最前線 3: 134 ( 2012)

ioOgBMS  .5+*g^tl̑ȓ Ƌ
±ΈŔũɾħʪũʠʵʪ˚ęɾħʪũŻǻ
606-8501 ±ΈƖƒ±ąĚʁ̣ͫʃ
Plant-specific transcription factor LEAFY and its possible role in gametophyte development inMarchantia polymorpha
Key words: gametophyte, LEAFY, Marchantia polymorpha
Takashi Araki
Division of Integrated Life Science, Graduate School of Biostudies, Kyoto University
Yoshida-Konoe-cho, Sakyo-ku, Kyoto 606-8501, Japan!
ft_\ψMarchantia polymorphaω2, ΞȨɯψȑ˺Ȩɯ [Embryophyta]ω1ɹɾ1ˍ˚1¢*2,
ȏAðȒ.ìƉ#+˵ FIȞμL¶̥HpȨɯ* HψȖ˟͆Υ¢1, Ɇá+ʟƌ, ŔĨ+Ɇá, ƎȘ1Ę˟͆Lčɩω
ʿ˶1ʠʵŴ*2, ɾɎɽ1Ͷă.źHΙƽF, 2007 ƝFft
_\Lʀ #ʠʵLͶ@)#!1¢*, KIKI2, (1) ̦ŤȨɯ. )!1ƲøEʠʵI)G, !1ʦÚʏȳ˽.̇ĥǡ#IH, (2) ΞȨɯ1Ͷăͼʯ*΃¼ŤΎ̭;+M,͙&)F, Ug[ΙËȉʣ.ʣŰ*H, +ϋ'1̲ɦF, Ȩɯĺȑ1͠æĵŤ LEAFY(LFY) .ʚʔ, ȏð1̴țź͒15+'+) H
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ioOgBMS 

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thalianaω+XYgRψAntirrhinum majusω. ), ̊ơì̧˗˩1ÒȆ.ʌɾHì̧˗˩, ̊̋ì̧˗˩+)1ƆǃLɵƷHͼʯ*, Ύ̯-ƲøLȜ#͠æĵŤ+)ěŰI#ˏ 350
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Moyroud et al. 2010, Siriwardana and Lamb 2012aω͠æĵŤ+)1
LFY 1ȳ˽.' )2, DNA ˙ęrQ* H C rQψƶͭ1Lčɩω1ʹÂȭʹɄŰI, DNA +1ʖ­ÅʀB¬

T. Araki-1
BSJ- Revi ew 3: 134 ( 2012)

植物科学最前線 3: 135 ( 2012)ΐÂă.ƾ̯-OwΌȹŅɰŰI) HψHamès et al. 2008ω=#, 1˙Ȝ.Ņ( ), LFY
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LFY ΃¼Ť- ΃¼Ť~NLƯǍH.̆F-&#.' )2͍* H+IH=#,Ͷă1͙ɜ.' )A, DNA ˙ęrQψC rQω1ȭʹ+ DNA 91˙ęȮƨ,'

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A1ψ,IŞŜ˯* H2ʣŰ) - E * HIF1ˠ̜μ.ΙHč˵Ȏ+)2,
Smith 1955a, Graham 1996 [1993], ¯ 2007 Lčɩω.2Ug[ŧĽHĖ˽ǃ G, ʠʵƴ#IH̬ŤȨɯǡ'ϋ'1 LFY ʖě΃¼Ť, LFY Ug[+ NEEDLY (NLY), .' )2, ɺȜμψΒ̕Ȳμω1˻ŤĲʳ.Hʌɹzi1̴ț̭ǿʱ*-I)G, I.Ņ( ), M.
Frohlich .EG̦ŤȨɯ1̊1͙ɜ.ΙḢĥɖ ¸͆ǰĬI#ψƶͭ1LčɩωʱŤȨɯ·Œ1ΞȨɯ*2, ̞μ1{mWv_\ψPhyscomitrella patensω1ϋ'1 LFY ΃¼Ť
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2005ƶͭ1Lčɩω, ͠æĵŤ+!IõƸHʌɾͼʯ1Ͷă+̲ɦḞĥɖ˵ŹA-I) HψMaizel et al. 2005ω


T. Araki-2
BSJ- Revi ew 3: 135 ( 2012)

植物科学最前線 3: 136 ( 2012)@ioH{~$ Sio ̦ŤȨɯ.HLFY.' )2, Moyroud et al. (2009) .EHȏͫ1˟͆ÝÂLȬ̲H1.Ʋʹ'ψȖȁ¢1Ʃʀȁɴ1ȁɴcqF1ɠI2ȿ.-HωaQusds+XYgR·Œ1̦ŤȨɯ*2, qqψSolanum esculentum [ǌʀĜ+)2, Lycopersicon esculentum], FALSIFLORA[FA]ω, ktOψPetunia × hybrida, ABERRANT LEAF AND FLOWER [ALF]ω, iy^ψNicotiana
tabacum, Nicotiana FLO/LFY [NFL]ω, TrRψPisum sativun, UNIFOLIATA [UNI]ω, ^[`ψLotus
japonicus, LjLFYω, iR_aψMedicago truncatula, SINGLE LEAFLET1 [SGL1]ω, QvψOryza
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IF1Ȩɯʱ1̴țF̶ H+2, ;+M,1ňę, LFY2̊
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΃¼Ť1ͼ÷ʌɹ2I+ &#̥ɹŁ1őăL5- ̇ĥɖ +., ʪ1όʱ1ň
ę.2, ̊1ƯǋƯǍ.ü , ̕1ƯǋƯǍ.AΎ̯-ƲøLȜ#)G, ȳ˽ȴǵőʇÂ*2 IA̭̕ĉ̕ăHʯƣ2ƪ A11, ěȮ1̥ɹŁ2qq1ňę.A̰FIHI2, LFYɧΜǍΕǃψindeterminacyω1˜ǡ. )Ύ̯-ƲøLA'+LʤĪHA1+̴΍I) HěȮ.), Qv1 RFL ΃¼Ť1ʌɹǘõÂ*2, ̊Ǎ͸Ʀ++A.̊ơ1ìȝ1ȫʼ-ɗſ̲ŹIH+F, RFL ΃¼Ť2̊ơì̧˗˩1ɧΜǍΕǃ1˜ǡ.ΙKH+˵ FIḢĥɖ +., RFL ʌɹǘõÂ*2, ƯǍI#̊2ȶƚ-Ưǋ+ʰǃLǡ&)G, RLF ΃¼Ť2̊̋ƯǋƯǍ1£̯-õƸĵŤ*2- +IH ·.ͭ:#E ., ̦ŤȨɯ.HLFY1Ʋø2, ȳ˽̴ț-I#ΜFI#ǿ1ʱL>)
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E ., ΃¼ŤΎ̭.EG, LFYZr1΃¼Ť+NLYZr1΃¼Ťϊ^}'ŧĽH+ȉF.-&#̦ŤȨɯ+[vmƆ*2, NLY Zr1΃¼ŤŗKI#+˵ FI) H

T. Araki-3
BSJ- Revi ew 3: 136 ( 2012)

植物科学最前線 3: 137 ( 2012)

ψFrohlich and Parker 2000, Frohlich 2003, p. 71NLčɩω pPOimψPinus radiataω*, ȏð.ϋ'1΃¼Ť, NEEDLY (NLY) +Pinus radiata
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Siriwardana and Lamb 2012aω̦ŤȨɯ1̊1͙ɜ.ΙHʇ-H¸͆+)2, MADS lZc΃¼Ť1ʠʵ˶* H G. Theissen F.EHA1 HψMelzer et al. 2010ω:5<8()3,0ioW ajȨɯ*2, ̚Ĳajμ1krd|ψCeartopteris richardiiωFϋ΃¼Ť, ʘĲajμ1R|iQψAngiopteris lygodiifoliaωFό΃¼ŤZt[I) H;, myψPsilotum nudumω, cYsψEquisetum arvenseωFA!I"Iϊ΃¼ŤƷFI) HψHimi etal. 2001ω
1 %, krd|1΃¼ŤψCrLFY1, CrLFY2ω.' )2, RNAlq.
EHʌɹ̴ț-I) H ΃¼Ť+A˻ŤÂ¶. )2, ȢξaqE6ɾȺaq
1̏β+̙ƕɱǋ1˻Ť̕*ʌɹ)G, ɰ.ɾȺaq1̏β. )Ƭ ʌɹ̲ŹI#Ȇ, ΉÓÂ¶ψö̕Âω*2, ʌɹ2̓@FIHA11, ʹˋİE6ʹċİƯǍIHȌȒLĠ@), !1ʌɹ2Á A1* H+>-I) HψHimi et al. 2001ω ž̕μ*2, {dtψIsoetes asiaticaωF·ìΉïŇģI) HψHimi et al. 2001ω;,]w̴͇˖¨#QuVi{yψSelaginella moellendorffiiω. )Aĉ^}1ŧĽʣ̓*
HSiriwardana and Lamb (2012a) *2, ΉïOQq1¢. SelLFY +)ĒGFI) Hž̕μ*2=&#ʌɹ͉:FI)F, ̴țƴ#IH
26+* ^\Ȩɯ2, ̌μ, ̞μ, mw_\μ1ό˯ψΖ phylumω.ìFIHψVanderpoorten and Goffinet
2009, ƎȘ 2012 Lčɩω=, mw_\μψˏ150ʱ, Renzaglia et al. 2009ωF2³1+J LFY U
g[1Ňģ2- , ŧĽH+2ʋ - 1*, Ňģƴ#IH+J* H ̞μF2, {Ri_\ʔ1{mWv_\Fϋ'1΃¼ŤψPpLFY1, PpLFY2ω, cY_\ʔ1ik_\Ɔ1ϊʱψAtrichum angustatumωFϋ'1΃¼Ť1ÝΕ.ͫ ·ìȄɭψAtranFlo1,
{
AtranFlo2ω, !I"IZt[I) HψTanahashi et al. 2005, Frohlich and Estabrook 2000ω
Ri_\ʔ1{Ri_\ψFunaria hygrometricaω*2, qcZq̴ț.EGΉÓ

T. Araki-4
BSJ- Revi ew 3: 137 ( 2012)

植物科学最前線 3: 138 ( 2012)Â+˻ŤÂ1΃¼Ťʌɹ1~NQ1Ƚͣ-KI) HψSzövényi et al. 2011ω, ȹǁ-F, LFY Ug[2pic.ʍΓI) - d_\ƆψSphagnumωF2, CrQ
á1ˏ30OwΌȹŅ.źǀH·ìΉïŔΐ.pic.ʍΓI) H, ÝΕ- 2!I.ͫ Ε1Ήï1Ňģ2- ̞μψˏ13,000ʱω1¢*, d_\Ɔ2, ̞μ1Ͷă1ðȒ.
ìƉ#d_\˝., cY_\ʔ+{Ri_\ʔ2, F.ƶ.ìƉ#+cY_\˝+_\˝., !I"IĠ=IHψGoffinet et al. 2009, p. 71NLčɩωƶͭHψLčɩωE ., ̌μLĠ@#µ1ˍ˚+Ƚ:), ̞μ*2 LFY ΃¼Ťɰȸă) H+©ǈIH, !1ȩ̽
.2, Ɵ˄-`[ƾ̯* H ͭ1E ., {mWv_\. )2, ϋ'1΃¼ŤψPpLFY1, PpLFY2ω1GUSiLʀ #ʌɹzi1̴ț-I, ΃¼Ťʡŏ.EGȳ˽ȉF.I) HψTanahashi et al.
2005ω ΃¼Ť+A., ΉÓÂ+˻ŤÂ1 Ȇ*ʌɹ) H1 %ΉÓÂ*2, aqβ*ʌɹ̰FIH;, ʹċİ.Hʌɹ̲ŹI#, ʹˋİ. )2, ,%F1΃¼Ť1ʌɹ
A>+@FI-&#Ȇ, ˻ŤÂ1ʌɾͼʯ*2, 2@˻ŤÂÝÂ*>+@FI#ʌɹ, ˻ŤĲ, 蒴Ƞψsetaω, ͛ψfootω1ìăƶ2, ˻ŤĲ+͛.ΜŰI#PpLFY1 PpLFY2 ¬Ύʡŏȣ.
)2, ΉÓÂ.2ʇƚ2̲ŹI-&#, ˻ŤÂ1ʌɾϊ˔˻ȒEGÚ.2Ͷ̡- +ȉF.-&#1+2,

΃¼Ťēˋċ1ʾì̧.ƾε* H+LǉĥH+̴΍I)

H¬Ύʡŏȣ1ċ.ΏɾŁ1ˋŤLēˋ#ňę.2, ȶƚ-˻ŤÂ1ʌɾ̰FI, ȶƚ-ʌ̋˽+ìΧȽLʤ˻ŤƯǍI#+F, ΉÓÂ.HċƯǍ!1A1.2ʇƚ2- ēˋċʕ%.ɗǿì̧, Â˔˻ì̧L˪Gͬ+*, ˺ʌɾL˘)˻ŤÂƯǍL- +2,ΞȨɯψȑ˺ȨɯωLɰƻ(HΎ̯-Ư͕* HψBower 1930, Smith 1955bω1̲ɦF, ¬Ύʡŏȣ1̥ɹŁ2ȫ@)̇ĥɖ {mWv_\11ʠʵ2, ̦ŤȨɯ·Œ1Ȩɯ.Hī
1ȳ˽̴ț1Æ* G, LFY1ʦÚʏ-ȳ˽1ǫ˓.ź)ʤĪ++A., Ŕ-͈κL HA1+-&) H
.5+*'J  ̌μ2, ˏ400Ɔˏ5,000ʱLǸ, ό'1˝ψ^k_\˝, ft_\˝, m _\˝ω.ŔòI
HψCrandall-Stotler et al. 2009, ƎȘ 2012 Lčɩʱǿ.Ι)2, 7,500ʱ+

ȏͫ1̰ʴAG

H [Konrad et al. 2010]ω
LFY Ug[1Ňģ H12, ft_\˝1ft_\ψFrohlich andEstabrook 2000ω+RX_\ψRiccia fluitansωψMaizel et al. 2005ω1ϋʱ1>* Hψ IA·ìΉïωƶ˶F2, allelic variants +IHϋʱμ1ΉïψCrQ1·ωʍΓI) H̞μ·.,ʝ̰͛) H+̶ H  #ɱɇL͞= ), ʿ˶F2, ft_\FÝΕcDNA+iLĠ?ˏ20 kbp1]wζŃLZă#aQusds+Ƚͣ#ňę, QqΕ, ɰ.ʾQq2 6
kbp ͫ Ε* &#ψͩ¯ʂο;,ȈȖȨɯũ»ʾ72ĴŔ»,2008Ɲωft_\.ü , ˮĨʨ˼ĊŐψɹaXŔũω.ȗȂ.ΙHćúL¹, b_\ψConocephalum conicumω+{b_\ψC. japonicumωFA, CrQ.Ʈ#H·ìΉïLƷ) Hψͩ¯ʂο;, Ȕʌ̥ω ƶͭHʿ˶F1̴țψLčɩωLΝ3, I=*1+J, ʌɹBȳ˽.ΙH̴ț2̞μ+1Ƚͣ1̇ĥɖ ĭκ.' )2, ȵδ*͋H

T. Araki-5
BSJ- Revi ew 3: 138 ( 2012)

植物科学最前線 3: 139 ( 2012)49)= }I49)=j$pkE  1åη*ͭ:#E ., CrQ2ˏ160OwΌȹŅF-HDNA˙ęrQ* G, ʹÂȭʹɄŰI) HψHamès et al. 2008ω ϋÐ1βaqE6ʞ *͵˙I#ϏÐ1α9lZcF-H^zZq-ǙGʈ>ȭʹ*, DNA1zrΉï1Ćì+˙ęHψ#&), ¬ΐÂ*DNA.˙ęHωϏÐ1 %, 9lZcα2 +α3 9lZc-i-9lZcȭʹLĒG, α3 DNA1£ɞá.Àˬ), DNA+1ʖ­Åʀ1ŔĆLǝ&) H Maizel et al. (2005) 2, aQusds1LFYi1õƸ*{mWv_\1 PpLFY1
H 2 PpLFY2 Lʌɹ#ňę., lfy őʇÂ.H̥ɹŁ1ʖ̪+ɏ΃¼Ť1ʌɹͅŽÝ̰FI- +.ʚʔ, ==-ʱΘ*OwΌΉï1ȽͣL-&#!1˙Ȝ, {mW
v_\1LFY*2, CrQá1ϋǑ1OwΌȹŅµ1Ȩɯʱ1LFY+ʇ-H+L̰ $#^e`cΉï*2 394ʆʔ+ 427ʆʔψHamès et al. (2008) *2, 312ʆʔ+345ʆʔ.Ʈ#Hω
1ȹŅ, µ1ʱ*2!I"I, {ckbψHω+OYtψRω* H1.ź, {mWv_\*2, OczYΌψDω+acoQψCω+-&) #ěȮ1ˬǱψ·*2, DCŁ+̹=#, µ1Ȩɯʱ1LFYLHRŁ+̹ωik_\Ɔ1ϊʱ1ϋ'1LFY*A̰FIH+F, ̞μ.ɰȑ1ˬǱ* H+ǭ͋) HǺő PpLFY1 ΃¼Ť+Ư͕͠ǱÂLʀ #̴țF, ϋ'1ˬ
Ǳ1 %, HD 1ˬǱ˙ęΉï1őăLA#F, lfy őʇÂ1ʖ̪˽LȴČĵ* H+ʤI) HHamès et al. (2008) 2, H3129lZcα31·LǍ, α31Nȕ.Ʈ#HP3081%D,ϊ9lZc.ÀˬH+, !1'ö1K307 -2 E6 +2 1Àˬ1[OtψGω+ʕǬʖ­ÅʀH+F, {mWv_\.H HDE6 RCψα31.Ύ-Hα5 á.ÀˬHω1ˬǱ2, 9lZcα31Ήĝ.ƱΰL HĖ˽ǃ H+) H Maizel et al. (2005) 1ʠʵ1Ȍɦ*·ìΉïʝFI) #ϋʱ1̌μ, ft_\+RX_\1LFY
2,

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ʱ. )Aĉ^}+˵ FI,

IAHRŁ* H#&), I=*1+J̌μft_\˝1ύʱ2:)HRŁ* H-, d_\Ɔ1·ìΉïpic.Ŕΐ.ʍΓI)
HψLčɩω,IAĭκ1˂ǑEGAǿȹŅNȕÒ1·ì* G, ̞μ1Ņ·ͫ*ìƉ#1̇ĥɖ [HRŁ, DCŁ1 I1LFYLǡ'.' )2³1+Jȉ*H ̇ĥɖ +., CrQá1ϋǑ1ˬǱ.ü , {mWv_\+ik_\*2, NrQ
+CrQ1Θ1ζŃ72 76OwΌȹŅ+ʞ ψµ1ʱ*2ˏ100OwΌ1A1œ #$̌μ*2ft_\ǇŇ- ω
F., {mWv_\1ϋ'1΃¼Ť1TXgQqȭʹψTanahashi et al. 2005ω2, aQusdsBQuVi{yψaccession: EFJ20684.1ω, ft_\ψͩ¯ʂο;,ȈȖȨɯũ»ʾ72ĴŔ», 2008Ɲω+2ʇ-G, NrQEGNȕÒL^rHζŃ., µ1ʱ.2̰FI- Ε QqLǡ'ik_\Ɔ1ϊʱ1ϋ'1΃¼ŤψAtranFlo1,
AtranFlo2ω1Ήïψaccession: AB286054.1, AB286055.1ωL{mWv_\+ȽͣH+, 1ʱ.
 )A, 1Qq1ŧĽ©ǈIHIFF, ̌μLĠ?µ1ˍ˚+Ƚ:), ̞μ*2
LFY1ɰȸă͙) H+˵ FIHψN1ɦˡȞáω1ɰȸă̞μ1Ͷă1,1ȻΠ*͙
#LǭŰH#@.2, ̞μ1Ņ·.ÀˬHd_\ƆBsbb_\ƆψTakakiaω,T. Araki-6
BSJ- Revi ew 3: 139 ( 2012)

植物科学最前線 3: 140 ( 2012)

N@ioExcU%$ QĚμψ͗ω+̞μψΫω.' )2£̯-ˍ˚Lʤ#ĝ1̳Ư2Qqψ͗̳Ư2̞μ1ϋʱ.1>ŧĽHω1ǥÜÀˬLʤϋȁŦ1O~Nlq2, LFY ĉΧI) H¶̥ʏ-ʱψΫŦ2΃¼ŤȭʹK&) HA1ωLʤϐft_\ψMpω, RX_\ψRfω, b
_\ψCcω, {b_\ψCjω, ik_\1ϊʱψAaω, {mWv_\ψPpω, QuVi{yψSmω{dtψIaω, cYsψEaω, myψPnω, krd|ψCrωʡˡȞ
á2, ̞μ*͙#+˵ FIHőă

T. Araki-7
BSJ- Revi ew 3: 140 ( 2012)

植物科学最前線 3: 141 ( 2012)

̌μ1ft_\˝·Œ1ˍ˚ψ^k_\˝Bm _\˝ωF1LFY1ĉΧƴ#IH
tms"vr7/?=
 ·*, Ęˍ˚1LFY.' )Ȭ͆#, ȏƶ., ʌɹzi.ΙHʝ̰L, ɾɎɽ+1Ι͵*=+@)+.#  =, ̦ŤȨɯ*2, LFY ΃¼Ť1ʌɹ2˻ŤÂψ2nω¶.ΜŰIHE * HaQusd
s*2, ˋ˔˻ψBorges et al. 2008ω+˺Ĳ1ċ˔˻, ý˔˻, ¢Ŗ˔˻ψWuest et al. 2010ω*qcZq̴ț-I) H,

I. )A LFY ΃¼Ť1ʌɹ2>+@FI-&##&), ΉÓÂ*2ʌɹ- +˵ FIH ̬ŤȨɯ. )2, I=*.-I#ʌɹ̴ț2˻ŤÂ1>.ΜFI) HpPOim
1 NLY ΃¼Ť1Æ*2, ̊ˈȼ˔˻ψ2nω*ʌɹH+ŇģI) H, !I·Λ1cob.H̲Ź2-I) - ψMouradov et al. 1998ω
1;, qR{ƆψPiceaω1ÆψCarlsbecker
et al. 2004ωB, ”Mostly Male theory” 1Ǖñʏ-ȩ̽Lǁη.ʌɹ̴ț-I#qR{ƆBXƆ
ψPodocarpusω, QkQƆψTaxusω1 LFY ΃¼Ť+NLY ΃¼Ť1ňę. )A, ˺ɸBž˻ŤĲ.
Hʌɹ2ʣ̓I) H, ˻ŤBΉÓÂ.Hʌɹ2͉:FI) - ψVázquez-Lobo et al.
2007ω
[vmƆ1ňęAěȮ* HψShindo et al. 2001ω ʱŤȨɯ+2ʇ-G, ajȨɯ.Hī1̴țÆ* Hkrd|*2, ˻ŤÂ.
ü ), ΉÓÂ. )Aƪ ʌɹȩëI) H, ʹˋİ, ʹċİ*ʌɹH1+ɦLĠ@), ́˔-ʶΘʏ-ʌɹzi2ȉ* HψHimi et al. 2001ω
ž̕μAĠ@#ʌɹ̴ț
ƴ#IHHimi et al. (2001) 2, 1ʌɹȮǋMADS lZc΃¼Ť1A1+2ʇ-H+F,
MIKCCŁ1MADS lZc΃¼Ť1LFY.EHõƸ2, ajȨɯ. )2=$ʣʹ) -&#+˵Ź) HB2G^\Ȩɯ.Hī1̴țÆ* H{mWv_\*2, ͭψ
Lčɩω1E ., ˻ŤÂ.ü , ΉÓÂ. )AaqβBʹċİ.Hɰʇʏ-ʌɹ̲ŹI) Ḣĥɖ +., PpLFY1, PpLFY2 +A., ʹˋİ. )2ʌɹ>+@FI-&#ψTanahashi et al. 2005ω ·F, ^\Ȩɯ. )2ΉÓÂ, ˻ŤÂ1 Ȇ*ʌɹ) #A1, ajȨɯ*2˻ŤÂψaqβω.HʌɹØĀ+-G, ʱŤȨɯ*2˻ŤÂψaqβω1>.ʌɹΜŰI
HE .-G, ɾȺǍΕ.ΙKHE .-&#, +asU˵ FIHΉÓÂ.H˗˩ɰʇʏ-ʌɹ̰FIH{mWv_\. )A, !1ȳ˽ʏ-Ύ̯ǃ*.˻ŤÂ.ΜFI)
HE .̰ H+2# 9Ṁĥɖ  IF1+Lǁη. ), KIKI2, ft_\1ΉÓÂE6˻ŤÂ.HMpLFY ΃¼Ť
1ʌɹzi1̴țL, RT-PCR, in situ RNA hybridization, MpLFYpro (5.3):GUS Ư͕͠ǱÂ-,.EGͶ@)#ψΊ¯ďƘŵ˙̤;,ȈȖȨɯũ»ʾ74ĴŔ», 2010Ɲ, ȈȖȨɯũ»ʾ75ĴŔ», 2011Ɲω
!1˙Ȝ, {mWv_\+ěȮ., ΉÓÂ+˻ŤÂ1 Ȇ*ʌɹH+ȉF.-&#, ΉÓÂ.Hʌɹ1Ȯǋ2{mWv_\1ňę+2 '1Ύ̯-ɦ*ʇ-H+AK&#!1', ΤǃɾȺİů1ʌɾͼʯ.Hʌɹ* HψΊ¯ďƘŵ˙̤;, ȈȖȨɯũ»ʾ75ĴŔ», 2011Ɲω1ʌɹų΢.ȳ˽ʏ-ǉĥLǡ'Ė˽ǃσ +2, ʌɹǘõ
Â1̥ɹŁ-,FAǹǡI) HψΊ¯ďƘŵ˙̤;, ȈȖȨɯũ»ʾ75ĴŔ», 2011Ɲω1ɦA, PpLFY1 PpLFY2 ¬Ύʡŏȣ1ΉÓÂʇƚLʤ- ++2źɩʏ* H

T. Araki-8
BSJ- Revi ew 3: 141 ( 2012)

植物科学最前線 3: 142 ( 2012).5+*$^tlKDEg^tlx ^\Ȩɯ2, ϋȖ1ίȾLǡ'ˋŤψspermatozoidω+ċ+.E&)ȑǃɾȺL- IF1ΉÓŤƯǍIHɾȺİů2œ˔˻ǃ* G, !I"I, ʹˋİψantheridiumω, ʹċİψarchegoniumω+Ħ3I, b\lqψjacketω+ϊƇ1˔˻ƇΉÓŤ˔˻LĂ>, ΉÓŤĲL-ɾȺİůœ˔˻ǃ* H+2, ˠ̜μ.2- , ^\ȨɯLĠ?ΞȨɯ1ɰƻ* HψBower 1930, Smith
1955b, Ĩȁ*2, ĄƏ 1989 .ƜȊ-̴͆ Hωb\lq2ΉÓŤL§ɫFŬHǉ˰Lǡ%,ΞɽŌ91΁ǀ* H+I) HI.ü ), ʹˋİψΤǃΉÓŤĲω1ňę.2, ƶͭH
E ., ft_\LĠ?·1̌μ*2, ˋ˔˻ŉLŒ·.ǨëH΢.Aʴȫʏ-ƲøLȜ#+ʝFI) HψBergdolt 1926, ƶͭ1Lčɩω ^\Ȩɯ1ȑǃɾȺͼʯ.' )2, ĄƏ (1990b) ., ɾȺİů1ƯǍ, ΦΤǃ, ēˋ-,Ý̈.K#H̴͆ G, ft_\1ɾɎɽ1̴͆ψĄƏ 1990aω1¢.A, ȑǃɾȺͼʯ1̹ͭĠ=IH^\Ȩɯ1ˋŤēˋ.' )2, ŔĨ (2012) 1˟͆1¢.I=*1ʠʵˇɡ.=+@FI)H=#, Ȗ˟͆Υ1¢*A, ƎȘ (2012) ft_\1ȑǃɾȺ.Ι), œǿ1ĹL= )́˔.
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Hͼʯ.ΙKHĖ˽ǃ H+F, *2, ft_\.HΤǃɾȺİ1ìă+ˋŤƯǍ,ēˋ.' ), ŔĨ (2012) E6ƎȘ (2012) +1Ύ̭L΄'', I=*1ʠʵ1ȷėB!1ͼʯ*ƷFI#ʝ̰, Ȕ̴Ʉ1ĭκL¢ƽ.̹ͭH+.# -, ·*2, ĕ ʠʵLƩʀ
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NKa>KZ\_`x ¡: ΤİǓ1ȭʹȠψStω+ΤİƠψRecωFǍH¢: ΤİǓ+ΤİƠ1ǍΕͼʯȻ2έF, Ȼ2ÒέF>#A1ÊŲ, ώ'1cob.ì) Hcobϊ1ΤİǓ2ʎʜƂ1Ú. Hcobύ*, !I=*éέ$&#ΤİƠ1έƜF.-G, ˤĐH+*,ɐ ʑɱ.-Hcobώ2ǍɪȻΠ¢1c\y2, 0.5 mmæʘ+cobì2, ŵ˙̤+Ί¯ďƘ.EH

T. Araki-9
BSJ- Revi ew 3: 142 ( 2012)

植物科学最前線 3: 143 ( 2012)ǓLʌ;Hɦ*, ft_\ʔ1¢*Aɰʇ-ŧĽ+IHψSmith 1955bωΤİƠ1έ2, ΤİǓ1ìăðȒ.2rɱL) H, ǍɪH+, ˤĝ.ǡ%G, ɐ êέ1ʑɱ+-HψN¢ωΤİƠ1·.2ʹˋİ́ψGoebel 1930 *2, Antheridiengrubeω+Ħ3IH~c^ɱ1ʸɍ G, !1¢.ʹˋİđ=&) Hʹˋİ2, Ƞ+b\lqψ˔˻Ƈω.Ă=I#ˋČ˗˩ψspermatogenous tissueωFǍHψp. 12 1N¢Lčɩωʹˋİ́2, ˔ ͳ͝L˘), ΤİƠ1έ.ΗĔ)G, ƶͭHψLčɩωˋ˔˻1Ǽë1΢.2, 1ͳ͝Lͳ&)ˋ˔
˻ŉ1ǨëH ·1ΤǃɾȺİů1ʌɾͼʯ.HMpLFY1ƲøLȉF.H+, Ʈέ1Ύ̯-͈κ*HψE6Lčɩωg^tlxwyGft_\LĠ@), ^\Ȩɯ1ȑǃɾȺͼʯ.Ι)2, ̞̌μũ H 2^\Ȩɯ1ɾɯũ.Ι
Hȏͫ1ǽʪȎǍȎ*2;+M,ˑέøI) - ψSchofield 1985, Vanderpoorten and Goffinet
2009, Goffinet and Shaw 2009ω
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Matthias
Jacob Schleiden+Theodor Schwann .EH˔˻͆ɾȺBʌɾ.źH̰ȆLŔőή'' HȆ*, ˋŤɾȺ. )Ȝ#Ʋø.' ) , =$ßͳ1ɻ̴1ʣʹ.2;,Ϳ&#Ȩɯ.
Μ&)>)A, ̦ŤȨɯ·Œ1Ȩɯψ!1ƮȌ1 KCHΣ̊Ȩɯ Cryptogamiaω2ȑǃɾȺLK- +̰ȆǹΉʏ* G, !1£Ĭ˶2;-F/ Schleiden * &#ψƳ2, ̦ŤȨɯ1ēˋ.Ι)2, ̊ˈ˃F˺ƯǍIH+

͆LǰĬ) #ωψFarley 1982ω
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E6 Farley (1982) .A+(Bower (1930) .EH+, ft_\1Ưǋ+ʌɾͼʯ1̹ͤ2, ~c.HλƺΔ1Ȩ
ɯƯǋũ1şʦ* G, Ȩɯ1ʌɾͼʯψ!1ƮȌ, history of development +Ħ3I) #ω.ʚʔ#ʠʵ1Úπʏ-ŧĽ* H Charles François Brisseau de Mirbel  1835 Ɲ.ʌ̥#ϋ'1͵Ň͋ȁψMirbel 1835a, Mirbel 1835bω.ş=H1 % Mirbel (1835b) 2, ʹˋİ.ʚʔ#ʠʵ* G, ~c^ɱ1ʹˋİ́1¢.đ=&)ŧĽHʹˋİ-,1̰«-ửĹL½&) HMirbel 2, ʹˋİ+!11Ƞ1·ìLęK), ̦ŤȨɯ1Τ̝.Ʈ#HA1+), étamine (stamen) +Ħ6,ʹˋİL anthère (anther), !1¢1ˋČ˔˻L pollen +#1͋ȁ*2, ̦ŤȨɯ+)V k
ψCucurbita pepoωL΂6, ̗1ƯǋLft_\+Ƚͣ) HMirbel 2, ΤİƠ.ź)ƪ ļúLü H+, ʹˋİđ=&) H\lq1ΗĔ·Fˊǃ1ɕÂë)H+

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Schmidel (1747) ψʿ˶2Ṵ̏ω1̲ŹLƩʀ, ̅͟*AěȮ1ˊǃ1ɕÂψliquere visqueuseωL̲Ź#+Ňģ) HF., 1ɕÂ2ÁÑɶ1*2¦ʎ̉.̰ H, ˏ 600 Ñ1λƺΔ*2, œǿ1ιˉB̊ˈˉψgrains entiers de pollenωψˋČ˔˻LǢ+˵ FIHωͱȉɕÂ¢.ɔ&) H1̓@FI#+LŇģ) H, ˋŤ1ʌ̰.2̆F-&#E* HĹɮ VII, ʾ 55 Ĺ FñȄH+, ˋŤŮǍö1ˋČ˔˻LĠ?ʹˋİL̲Ź.ʀ ##@+ǭəIHft_\1ˋŤ2, 1ƶ., ;1̌μ1ˋŤ++A., Franz Unger (1837)

T. Araki-10
BSJ- Revi ew 3: 143 ( 2012)

植物科学最前線 3: 144 ( 2012)ψʿ˶2Ṵ̏ω.E&)ʌ̰I, ϋȖ1ίȾLǡ'+A̲ŹI#Unger 2Iȶƚ-ɱǋ* H+2˵ -&#E * HψHofmeister 1862 [1851] .EHωWilhelm Hofmeister 2, Schleiden 1̖ȎFɲũ*ȅ ȨɯũLũ6, ̦ŤȨɯ1ēˋ+˺ʌɾ1
ʠʵB,

KCHΣ̊Ȩɯ1ȨɯÂ+ɾȺİů1ʌɾ.ΙHƟ˄-̹ͤʏʠʵL-&#˔˻͆.ʹ˿# Hofmeister 2, ̌μ̞μF̦ŤȨɯ.̆HΞȨɯ1ȨɯÂΘ, ɾȺİůΘ1Ȗ͕ʏ-źǀΙËL̰ǚ+*, Ȩɯ1ɾɎɽLΉÓÂ+˻ŤÂ1¶°¶ψGenerationswechselω+)Ǧ HÝȅ ̰ȆLǰĬ#!I=*1͋ª*Ϳ1 #Ǌ &#ψSchleiden A!1ĵ*HωȨɯ.HȑǃɾȺ+ɧǃɾȺ1Ȗ͕ʏ-ɻ̴2, Hofmeister .E&), 2@)ǍͷFI#+

+*HψGoebel 1926 [1924], Farley 1982ω
), Hofmeister 2, !1Ύ̯-̖Ȏ* HσˀΣ̊Ȩɯ1Ƚͣʠʵ
ψHofmeister 1862 [1851]ω1¢*, ̕ɱÂ̌μ.' )Aϊ'1ʻÝÂLƮ)) HƳʠʵ#12, ft_\1;., b
_\ψFegatella conica [= Conocephalum conicum]ω, bW`_\ψRebouillia hemisphaericaω, VnX
ft_\ψLunularia vulgaris [= Lunularia cruciata]ω, x[ft_\ψTargionia hypophyllaω1ύʱ
* &#ft_\.' )2, Ƴ̅͟1̲Ź1̹ͤ++A., öë1 Mirbel L2@+HÚ̡ʠ
ʵ1Ǖñʏ˕´A-&) HΦǃɾȺİů1ʌɾ+˺ʌɾ.Ƚ:H+, ΤǃɾȺİů1ʌɾ.
øI#ˑέ2ΜFI) H, ʹˋİ, ̕ɱÂőƯ)*#ȭʹψΤİƠω1έ., ~c^ɱ1́ψʹˋİ́ω1¢.đ=&)ŧĽH+B, ΤİǓΤİƠ1ƯǍðȒBşČ˔˻F1
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Ä͊* H, Hofmeister 1σˀΣ̊Ȩɯ1Ƚͣʠʵ
ψ1851 ƝîȖ².E&)Ŕƛ.ō̪Ǻ̷
I#̼̎ 1862 Ɲ.î̡I#ω2, 19 ˎ1Ȩɯũ.HȏAΎ̯*ήȅʏ-̖Å1'+
IHψSachs 1906 [1875], Goebel 1926 [1924], Farley 1982ω
!1áŷ2ƼƢ), ̲Ź«ų1́˔-̹ͤ+Ú̡ʠʵ1Ǖñʏ̾È* HK.ȏƶ1ʻψ̼̎*2ʾ 16 ʻω.ʞ ÝÂ1˟Ǡ̈ăG, ϋ'1ʇ-H¶ψΉÓÂ¶+˻ŤÂ¶ω1Θ1¶°¶+Ȭǁ, -GíG'@##%*ǰʤI) H!1¢.2, ̬ŤȨɯ.ˋŤŧĽHĖ˽ǃ.' )1̶ĎA̰ $
HƜɥÅ®Ά++A.̬ŤȨɯ1ˋŤLʌ̰#ɃΏǍΆ2, Annals of Botany ͂FQkR+gom1ˋŤʌ̰.' )1ǗΓLɂ@FI#΢., 1©̶1+L̹) HψIkeno and Hirase
1897ωȶ̱1Ŕũǽ˸Lē-&# Hofmeister 2 19 ˎ1rQm1OVpd. &)2, ȫ@)ʇ̉1˘ȷ1ǡ%£* H!1Ȫ˦+¼̹2, Ζ1 Karl von Goebel +¬Ś1 Constanze .E&)=+@FI) HψGoebel 1926 [1924]ω
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̲ɦFǦ ʕ# Frederick Orpen Bower ŕŝ.EHω
Constanze Ʃʀ) HȇÚFŝ.ű)#ǒˑL̰H+, ̅Ū*ȧń) Hft_\L§ #ɱǋ.Í'E .+ʤB, m _\ψJungermanniaωL 20 ȖA̰'#+ǢE -, ̌μ1ǪΥB̲Ź.' )1̹ͭAǾ̰IH), ɾȺİů1ʌɾ.Ι)2, Hofmeister .˛ ), ƶ.Ȩɯũ1ǽʪȎ1¶Ĝ̿+-H Eduard
Strasburger (1870) ̹ͤʏ-ʠʵL- , ʑɱLǍΤİƠ1ƯɱψNω1ȳ˽ʏ-ǉ˰ψLčɩω.' )A͋) Hft_\1Ưǋʌɾ1Ĺ͆+)ʝFIH Kny (1890) 2, ·
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T. Araki-11
BSJ- Revi ew 3: 144 ( 2012)

植物科学最前線 3: 145 ( 2012)* HˋŤ1Ưǋψʾ 86 Ĺɮ 13 Ĺω1E ., ³Ȉ1ʔFH+ȶʣ-ĹA H, ΤİƠ1
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ŤÂ1ʌɾ+˻ŤƯǍ1ͼʯ.Ι), ³Ȉ*Aȑʀ-ʠʵ+)2, Durand (1908) .EHA1
HDurand 2, I=*1ʠʵ1ɏILē), şČ˔˻F1ʹˋİ1ʌɾͼʯĹ˔.̹ͭĹʤ) HȆ, ˋŤƯǍͼʯ.Ι)2, ɃΏǍΆ.EHʠʵψIkeno 1903ω G, Durand (1908)
B^\Ȩɯ1ǽʪȎψCampbell 1928, Parihar 1962ω.ƩʀI) HzKvtx Durand (1908) 1̹ͭLA+., ʹˋİ1ʌɾͼʯ2·1E .=+@H+*HψNω=, ̍ ΤİƠ1˳·1̥ʐ˔˻FϊÐ1ʹˋİşČ˔˻ψantheridium initial cell, AICωɾHʹˋİşČ˔˻ψAICω2, β·Ò1ʹˋİȖÂȼ˔˻ψantheridium proper mother cell, APMCω+Ņ·Ò1Ƞȼ˔˻ψstalk mother cell, StMCω+.ì̧HʹˋİȖÂȼ˔˻ψAPMCω2, ȱì̧+˥ì
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cell, WCωLɾHˋČ˔˻ψSCω+Ŏ˔˻ψWCω2, ì̧L˪Gͬ, !I"I, ˋČ˗˩+b\lqLƯǍH+.-HˋČ˔˻2, ˋȼ˔˻ψspermatid mother cell, SMCω+), ȃ@ȆĝNzK\_vtx¡: ʹˋİ1ʌɾͼʯ¢: ˋŤƯǋƯǍö1ʹˋİȠE6b\lqψɀ̉ω+ˋČ˗˩ψχ̉ωF-HΉÓŤĲLʤ£: ʹˋİ1ʌɾͼʯ.H˔˻ˍ͐ōȺì̧1Ĵǿ2ȶʤ)- ́˔2ȖȁLčɩ

T. Araki-12
BSJ- Revi ew 3: 145 ( 2012)

植物科学最前線 3: 146 ( 2012)1ȏ˖ì̧L- ϋ'1Ţ˔˻ψˋ˔˻ [spermatid, Spd]ω2ˋŤψspermatozoid, Spω9+őǋψƯǋƯǍωHψspermiogenesisωʹˋİƯǍIHͼʯ*, Ĥĸ1˔˻1ì̧.EG, ʹˋİ
đ=H́ψʹˋİ́ωƯÅFIHzS\` Ȩɯ1ˋŤÝ̈.Ι)2, ɛɐ (1969) B Renzaglia and Garbary (2001) .=+@FI)G, ɰ.
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ƷFI) #ʝ̰L=+@) H!1ƶ1ʠʵǍȜLĒGÜI, Renzaglia and Garbary (2001) 2, 
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H+̶ E  ft_\LĠ?ΞȨɯ1ˋŤ1ίȾ2, œ1ʘȤɾɯ.ßͳ1 “9+2” ȭʹ1ίȾ͡ˌLǡ', jQtŒ̂Lȴ ) HψHyams and Campbell 1985, Carvalho-Santos et al. 2011ωI2, ^\ȨɯBajȨɯ1ˋŤƶͯ͹ɋ˽Lǡ#- +1Čĵ+˵ FIHψŔĨ 2012 .EHωίȾ1ȭǍ̟ʎ͕+!1΃¼Ť.Ι)2, ˠ̜ZrscL¢ƽ.ʠʵͶ@FI)G, œ1ʝ̰̘ʴ) HψMerchant et al. 2007, Witman 2008, Inaba 2011ω
!1˙Ȝ, ίȾ.2 600 ʱL͚ H̟ʎ͕ŧĽH+ȉF.-&) HψWitman 2008ωʿ˶F2, Zrsc1ǇŇLA+., œ1Ȇ1ý̶LƷ'', ft_\1 EST E6]wǇŇψ±ΈŔũɾħʪũʠʵʪɆáŨ¤ǽǧ, ʟƌÞƥĊŐ1ýú.EHωF, jQtŒ̂·Œ1ίȾ͡ˌψjQtá̂, pPO
cZ, ¢ƽź̨ˬω1ȭǍ̟ʎ͕΃¼ŤB, ŅƢžÂ, ¢ƽÂ, ίȾáͦͰˍ-,1ȭǍ̟ʎ͕΃¼ŤLǜëHÅȪLͶ@)G, I=*., ƷFI#A11·.' ), ʹˋİɰʇʏʌɹLʣ̓) HψΊ¯ďƘ;, Ȕʌ̥ω #Ok+2ò., ǃȡ̉ÂψY ȡ̉Âω.Ƥ¥H΃¼ŤψYamato et al. 2009ω1¢FˋŤBʹˋİ1ƯǍ+ȳ˽.ΙKHĖ˽ǃ H΃¼
ŤǜëI) HψYamato et al. 2009, ͫʉŔũŔĨÿƞĊŐ, ʩÎω!1¢.2, RcψMus
musculusω1 Parkin co-regulated gene (PACRG) 1Ug[1E -̇ĥɖ A1Ġ=I) H

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ψYamato et al. 2009, Supporting Information 1 Table 5ωRc1 PACRG ΃¼Ť2ˋƑá*2ˋČ˗˩
*1>ʌɹ)G, 1΃¼ŤLĠ?ȡ̉ÂζŃLȴŗ#őʇÂψquakerviableω*2, Ύˆ-ˋŤȴǵʰ̰FI, PACRG ΃¼Ť1äŽÜ.EGIĴƹH+ʤI) HψLorenzetti et al.
2004, Yanagimachi et al. 2004ω
Zrsc*2, PACRG ̟ʎ͕2, ίȾ͡ˌÝΕ.K#&)ŧĽ,ŒÒ1jlqƺž˃1ϑ˃+ϒ˃1Θ.ƃĽH+̲ŹI) HψIkeda et al. 2007ω ·1ίȾΙ͵̟ʎ͕1;.A, ˋŤ.2, ==-¶͎ˍ1΋˒B, ͘ăǃBΦΤΘ1̓͏
.ΙKH̟ʎ͕-,, !1ƯǍ+ȳ˽.Ύ̯-ƲøLA'̟ʎ͕œǿŧĽHψȾó+ȋ 2006ω

ft_\1ňęAěȮ* G, ίȾΙ͵̟ʎ͕1;., ˋŤɰʇʏ-ŋŅǃȤ̟ʎ͕ψReynolds andWolfe 1978ω, ʹˋİFˋ˔˻ŉLǨëH#@1ȳȭψLčɩω.ΙKH̟ʎ͕ψ˔˻ŎǍ
ì1¶͎΋˒-,ω, ǨëI#ˋ˔˻LĳǾHȳȭψLčɩω.ΙKH¶͎ɿɯ1ęǍ
.ΙKH΋˒, ˋŤ1ͺþLΗşHȳȭ.ΙKH̟ʎ͕, ͺþLĖ˽.H¶͎.ΙKH΋˒,͘ăǃ.ΙKH̟ʎ͕, ΦΤΘ1̓͏.ΙKH̟ʎ͕, + &#A11ŧĽ©ǈIHIF1̟
ʎ͕1΃¼Ť2, ʹˋİˋŤ1ìăͼʯ1ɰŰ1ȻΠ*ʌɹHƾ̯ H+˵ FIHI=*1ƺ˔Ưǋũʏ-ʠʵ.ü ),  #, ΃¼Ť̟ʎ͕+!1ʌɹõƸȳȭ.ΙHʠʵͶ@FIH+.EG, ft_\2ˋŤƯǍˋŤȳ˽LʠʵH#@1ØI#pɾɯ+-HĖ˽ǃLʫ@) HψŔĨÿƞĊŐ, ȈȖȨɯɾɻũ» 2012 ƝƝ»ωz|PbC
 ^\Ȩɯ1ēˋ.2ɀƾ̯* G, ʹˋİFʹċİ.̆H˘͝ɀ1'-G.E&)˙6'
-I3-F- +2ƟʝFI) Hēˋ. #&)ʹˋİFˋŤɋë+ȎIH
+œ , d_\LΝ ), ų΢.ǼëIH12, ˋ˔˻ψ̚ ˔˻Ŏ.Ă=I#ˋŤω1ŉ* HψĄƏ 1990bω
1åη.ͭ:#E ., ̞̌μũ H 2^\Ȩɯ1ɾɯũ.ΙHȏͫ1ǽʪȎ.21ͼʯ.ΙH̹ͭ=&#-, ĕ ǽʪȎμψGoebel 1905 [1898], Campbell
1928, Goebel 1930, Smith 1955, Parihar 1962, Watson 1971ω.΀H+*, E B̹ͭ̰'HI
F1ǽʪȎƩʀ) Hʠʵ2 IA, 1930 Ɲγ=*1A1* G, ȏͫ1b_\1ʠʵ
ÆψShimamura et al. 2008ωLò.I3, ͫƝ1ʠʵ2;+M,- E * HɹĽ*2;+M,
ɊʔIH+1- ͼʯ+ǂKIH1*, ·.ʠʵ1ȷėAĠ@), ſ́˕´H ̌μ.HʹˋİF1ˋ˔˻ŉ1Ǽëͼʯ2, `QxQ_\Ɔ1ϊʱψAstrella californicaω+b_\*, ɬʌʏ-ıëψforcible discharge, explosive dischargeωŇģI#+ψPeirce 1902, Cavers
1903ωLŘȳ., Ιƽǡ#IHE .-&#E * Hb_\1ÆLŇģ# Cavers 2˱Ɲ, b_\1ˋ˔˻ŉ1ıë2 50 Ɲͫ·ö.*. Thuret (1856) .EGŇģI) #+L͋ȁ1̹ͮ+)Þ̥) HψCavers 1904ω
I2, ~c1 E. Bornet F1ʩÎ.EHA1*, Bornet.EH+ Thuret (1856) 2!I=*.A;+M,ƩʀIH+-&#+ų΢, Gustav Thuret1̲Ź2, ɷʹaS̅ɨʪũć»͂ʾ 4 ƕ1¢1 “Analyse des travaux de la Société” +δʔ1¢.łɅHE .˕´I)G, ʔȵ.Aͤ&) - I*2̰'ë1AΨ 
Cavers .1Ňģ1ŧĽLʝF# Bornet 2++, ĭκ1δʔ1ö+ƶ1α., !I"I̜

μ+ľ̤μ1͋ȁLͤ) ), ʋĭ2Ɂ̴H%->., ̩ʧ-OkOȨɯũ˶* &#Thuret 2, A+A+2ɉƵŶ* G, ^cikw.H~cŒ°Ķ1ĩ* &#+A H+

, Ȩɯũ.ŻǁH#@.ȖȪLͧ@, ,1Ŕũ.AǑƆH+-,

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OkOȨɯũ˶+)ʠʵL˛, ̜μ1ɾȺ-,.' )ØI#Ȫ˦Lȹ#{yiψFucusωBabZψCharaω. )ˋŤLʌ̰#12, Thuret * HψFarley 1982ω +J*, ƶͭ1 Belgdolt (1928) 2, 1747 Ɲ., Schmidel ̌μ1ʹˋİΤǃɾȺİů* H+Lʌ̰#΢., *.ʹˋİ1áŷɯ1ǼëL̲Ź) H+ͭ:, F., Thuret (1851) 1mW
i\R^ft_\ψFossombronia pusillaω1ʹˋİ1ΗĔ.ΙH̲Ź+ Moore (1874)ψṴ̏ω1
VnXft_\.Hıëψ̹ͭFñȄH+b_\1ňę+¿) Hω1̲ŹÆLƩʀ)
HIFFñȄH+, Thuret (1856) ·ö1̲ŹŇģ 'ŧĽHE * H ), ft_\1ˋ˔˻ŉǼë.Ι)2, ƎȘ (2012) æʘLĠ@)̴͆) Höë1
Mirbel (1835b) 2, Johannes Hedwig ̞μ+̌μ1dft_\ψJungermannia epiphylla [= Pellia
epiphylla]ω*̊ˈ1ɬʌψl’explosion du pollenωLŇģ) H+.̵I, Hedwig BƳ.Ú̡
H Schmidel [1747] A, ft_\*2ěȮ1ɹ͒L̰H+-&#+, F. Mirbel ̅͟A 2 Ɲ.E8̲ŹȒΘ1¢*, ft_\*!1E -ɹ͒L̰Hƞͺ.2ǆ=I-&#+L̹ͭ)Hěö.˕´# Hofmeister (1862 [1851]) 2, ʹˋİǍɪH+, βʼ·1˔˻ΧI3I.-G, á·1ˋ˔˻ʹˋİ́1ɳ ΗĔ·Lͳ&)Œ.ǛëI, ΤİƠ1έ.-G1Ŕ.ɕɟ+-&)ɹIH+L̲Ź) HIF1̲ŹBƎȘ (2012) 1̹ͭ. HE .,ft_\1ňę.2, b_\+2ʇ-G, ɬʌʏ-ıë*2- 1+2, ʿ˶1ʠʵŴ.
H ʱ1̲Ź*Aʣ@) H, Cavers (1904) 2, b_\+¿#ɬʌʏ-ıëψexplosive dischargeω
2, ft_\1;, bW`_\BOVft_\Ɔ1ϊʱψPreissia commutataω. )A̲ŹH+*, ft_\ʔ.H̈ʏ-ɹ͒* HĖ˽ǃ H+) H
ψSmith 1955 [p.52] .AƩʀI) HωIFH+, ˋ˔˻1Ǽë1Ȯǋ.Ι)2, ft_\1ľŃΥĶΘ*ÃF1ìăɾ) H1AI-  ̌μ. ), ʹˋİ1b\lqβ·ΗĔ, ¢Fˋ˔˻Ǽë- 2ıëIHȳȭ.
Ι)2, Goebel (1898) 1ʠʵÚπ1E * HψBergdolt 1926.EHω
Goebel (1898) 2, ̞μ++A., R^ft_\Ɔ1ϊʱψFossombronia dumortieri [= Fossombronia foveolata]ω+xd_\ψBlyttia
lyellii [= Pallavicinia lyellii]ω.' )̹ͭ) H=#, Goebel (1905 [1898]) .2, ft_\ʪLĠ?
̌μ.' )1ʝ̰=+@FI) HI.˛ ), b_\.Hıëȳȭ.' )2,
Cavers (1904) ̹ͭ)Gψȹǁ-FĹ2- ω, ft_\+!1ěƆʱ.' )2, Bergdolt(1926) ft_\*, Andersen (1931)  Marchantia domingensis *!I"I1̲ŹLĹ++A.Ňģ
) H1 %, Bergdolt (1926) 1Ĺ2, Goebel (1930) 1ǽʪȎ.äΓI) H ȳȭ2̹1όʱ*Ȭ0ßͳ) HBergdolt (1926) 2, ʹˋİá1ˋ˔˻1Ǽë.ΙKHA1+), ψ1ωˋ˔˻LĂ?b\lqψBergdolt 1926 *2, ʹˋİŎ Antheridienwand +IHω1˔˻
1Åʀ, ψ2ωʹˋİ́Ŏψě Kammerwandω1˔˻1Åʀ, ψ3ωʹˋİψá1˔˻ω!1A1
ǡ'̄ɣǃ1ɯ͕1Åʀ, ό'ψ1˗>ęKωLǤ) Hb_\*2, ʹˋİ́1Ŏ˔˻1̄
ɣ1ŸAſ-- E * H, ft_\1ňę.2, ʹˋİ1b\lq˔˻1̄ɣ.EHŸ
Ŕ, ʹˋİ́1Ŏ˔˻2Ǽë.2ΙKF- +IHBergdolt (1926) 2, ʹˋİ1Ǎɪͼʯ*, b\lq˔˻ǟŔH+, =#!1ǟŔ2˔˻͕1ǟŔ.EH1*2-, ˔˻Œ.ˊɕ
͕̘ʴIH+.EH+̹ͭ) Hġɀ.½ , 1ˊɕ͕̄ɣH+.EG, b\lq˔˻ʹˋİ1áÒ.ĝ&)̄F>, ¢. Hˋ˔˻ǨëIHψNωAndersen (1931) .E

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H M. domingensis .H̲ŹAěȮ* G, Ǽëö1b\lq˔˻1˔˻Ŏ.2œΐ1 mucilage
Ġ=IH+, =#, Ǽë.2, ψ1ωb
\lq˔˻1ġɀ.½ áÒ.ĝ̄ɣ, ψ2ωˋ˔˻1˔˻Ŏ1̄ɣ, ψ3ω
ʹˋİ́1Ŏ. #H, ̉˒ψOqaOtωL̘ʴ#˔˻Ď<ļψI.
EGb\lq˔˻1̄ɣʹˋİ1á
ÒȆĝ.ĝ ω, ψ4ωʹˋİ́Ƣ·.
HˊɕȾψparaphysis; ƎȘ 2012 .ĹʤI) Hω1̄ɣ.EHʹˋİ1ʕʹŠĀ1˜ǡ, -,ΙKH, ȏAΎ̯-1
2ψ1ω* H+̲ŹI) H
Andersen (1931) 2=#, ʹˋİ1˔˻1˗˩ăũʏ-̴țÀ>) H, .2́ͭ- I.' )2, ɹĽ1ȅ ǒɉǖ̢.EH̴țƾ̯-+2͋L=#- N zK`nxz|PdCx$-;*13|\_QE¡: Ǎɪ#ʹˋİ¢: ˋŤ˔˻ŉǼëƶ1ʹˋİβ·1˔˻2ƍŏ) Hɀ̉*Ŋ&#·ìb\lq˔˻χ̉1·ì2ˋ˔˻LĠ?·ìψ˔·2ʗʅωAa: ȔǍɪ-ʹˋİ1b\lq˔˻Ab: Ǎɪ#ʹˋİ1b\lq˔˻Ba: ˋŤ˔˻ŉǼëƶ1ʹˋİ1b\lq˔˻̐̉̚*Ŋ&#·ì2˔˻͕Bergdolt (1926) LA+.ÅĹ ʹˋİFˋ˔˻ǼëIH΢.2, ʹˋİ́1ΗĔ·1.ÀˬHb\lq1β·1˔˻ƍŏHψN¢Lčɩω
̞μ1ňę.2, β·.ÀˬH 1 ÐA2ȉʛ.ąòIHſǿ1
˔˻I.ΙKH, ̞μ*Ad_\B̌μ1ňę.2, !1E -ɰò1˔˻2- +I)
HψGoebel 1905 [1898], Goebel 1930ω
1ɦ.' )A, iqlBTx`qlˍ˚-,Lʀ ), b\lqá1Àˬ.ǀ#˔˻1ìăŧĽH, Lȩ̸)>Hƾ̯ J  ʹˋİF1ˋ˔˻ŉ1Ǽëȳȭ.Ι), ȹHĭκ2, ͵1ͼʯ1ƩΑ+-Hɀ,1E.)ʹˋİ́1á·.ÜH+ɦ* HGoebel (1905 [1898]) 2, ʹˋİ́1ɳ ΗĔ·Lͳ)ɀŒFá·.ÜHȳȭ2ȉ* H+''A, ʹˋİ́Ƣ·1ˊɕȾìɈ, ΗĔ·.
̘ʴ)á·L§ɫFÍ͑) H mucilage ġɀ1ƲøAȜ#1$J +ǭə) H30 Ɲ
ƶ1ěȖ1ʾ 3 ɮ. )A, ʹˋİ́1ΗĔ·2, ſ-+AʹˋİǍɪHͼʯ1Θ2, ȿŴť+ěȮ.ɳ #@, ɀ1ÉÜL̻- $J +, ·ö+ěȮ1ǭ͋LƅΗ) HψGoebel
1930ω
1E ., ġɀ1ȳȭ.Ι)Aȉ1ɦȹ&)G, ʠʵ1Äľ H+̶ ! *H-, KIKI1̲Ź*2, ɀ.ǀˁ#ΤİƠ1őă.2, ƃǑʏ-˔˻1˔˻Ŏ̄ɣ.ʄ=F- , İůÝÂ.K#HƯǋőă.½ İůͺþ HE .AǂKIHψΊ¯ďƘ, Ȕʌ̥ωI.' )A, ³ƶ, ȩ̸)>Hƾ̯ H+˵ ) HT. Araki-16
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植物科学最前線 3: 150 ( 2012)bC%z|LezSeY>Fz .̰#ͼʯ.EG, ΤİƠέ1ʹˋİ́ΗĔFǨëI#ˋ˔˻ŉ2, Ð1˔˻.ìΧĳǾ)ɀέ.ƟG, ˋŤ¢Fë)H1ìΧĳǾ1ͼʯ2, ˋ˔˻ŉ.Ġ=IHſΐ1˾˷̥έƫúLɗſH+.EH+H͆ HE * HψĄƏ 1990b .EHωų΢.,
Muggoch and Walton (1942) 2, ǨëI#ˋ˔˻ŉLcQr[c.+, ƍŏ), ˋ˔˻ɀέ.ɒ6, ǿì1 %.ˋŤˋ˔˻F̀ëH+, sQʢΌŋȡ̉Bcj
Ⅲ ȡ̉.E&), ˋ˔˻ŉ.2˾˷ŧĽH+˵ FIH+LŇģ) Hˋ˔˻F,1E.)ˋŤ̀ëH1.' )2ȉ* Hˋ˔˻1˔˻Ŏ2̚ +I)GψĄƏ
1990bω, ˋ˔˻á.HˋŤ1ƯǋƯǍǍɪ1ͼʯ*, ˔˻Ŏ1őă͙, ̀ëLĖ˽.)H+ǈÖIH Ǎɪ#ΤİƠ1·2ʑɱ1êέLǍ)G, ɀLƷ)ˋ˔˻ŉǨëI#ƶ2, ˋ˔˻2!F̀ë#ˋŤLĠ?ɀLɚ #ɐ VZo[c+ȮʖLĢH1E -ɱǋ1+J.Ωɟ1E -ɀɟ̔%H+, ˋŤLĠ?ɀɟĤG.ƭν3I, ΧI#ňǑ.
ˋŤǾƗIH+.-HψStrasburger 1870, Goebel 1905 [1898]ω
ɧǃ̋1ǾƗ1ňę.AʝFI
HclaVlψsplash cupω+Ħ3IH>* HψWatson 1971, ĄƏ 1990aω
I2, ƾAɀ+

ţ͕1͵˛ǃLƾ̯+- ¼ǶȆɉ* HclaVlȆƨ.EHˋŤ1ǾƗLĖ˽.H#@., ˋ˔˻ŉ΁í.ƍŏ, ˋ˔˻ĳǾ)ɀέ.ƟH+2ɰ.Ύ
̯* H!1#@, ˋ˔˻1ìăͼʯ*ęǍI, ˔˻Œ.ìɈIH˾˷Ό-,1ʊɀǃăęɯ1̴ț2, ˋŤ1̀ëLĖ˽.Hˋ˔˻Ŏ1őă1̴ț++A., ³ƶ1Ύ̯-ʠʵ͈κ15+'+)̇ĥɖ  ), 1E .)Ĥĸ.ǾƗI#ˋŤ1 HA12, Φȣ1ʹċİ1ΗĔ·.#,GʚILĖ˽.H#@1ΦİǓΦİƠψarchegoniophore, [archegonial] receptacle 1Ưǋ1ɰƻ.')2*2ͭ:- ψGoebel 1930, ƎȘ 2012 Lčɩωų΢., ˋŤ,1E .)ʹċİ1Η
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植物科学最前線 3: 152 ( 2012)

 LFY .Ι)2, ̦ŤȨɯ1ΜFI#ʱψω+̞μ1{mWv_\ψωLΝ ), ʌɾͼʯ.HƲøLȉF.Hʠʵ2;+M,-I) - ̬ŤȨɯLĠ@, ʌɹzi+̦ŤȨɯ1ʝ̰.A+( ), ÚÜ̲LA&)ƲøǭəI) H.ͼ- ψÆ 3,
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+*#ʾϋʻ2, ƈĔʥŤĊŐF1ǿœ1Ƨ̺ʏ-^q.EG,'1Ύ̯-ɦLǺĮH+*#=#, ŔĨÿƞĊŐF2, ^\ȨɯajȨɯ1ēˋ.ΙH˟͆ČʲL̰) #$, ȏ˖ʲ1č˵.H+*#IF1Ȇ.ɰ.Ǌ͎# [ufq
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