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
Ȗ˟͆*2, = LFY .' )Ȭ͆, ' *, ft_\. ) LFY ΙKHĖ˽ǃσ +K
IKI˵ ) H, ȑǃɾȺ.HΤÒ1Ʋø.' ), ft_\Ȩɯũ1ź͒+)ʠʵ
IHE .-&)FɹĽ.̆H=*1ʝ̰L=+@, ³ƶ1ʠʵ1ȆĝǃL˕´H++A., ͇˶1č˵.Ç# 

ioOgBMS 

LEAFY/FLORICAULA (LFY/FLO)ψ·, ˇÊ1#@, LFY +ʅω2, aQusds
ψArabidopsis
thalianaω+XYgRψAntirrhinum majusω. ), ̊ơì̧˗˩1ÒȆ.ʌɾHì̧˗˩, ̊̋ì̧˗˩+)1ƆǃLɵƷHͼʯ*, Ύ̯-ƲøLȜ#͠æĵŤ+)ěŰI#ˏ 350
430 OwΌȹŅFǍG, N ȕÒ. 76 OwΌȹŅ1 N rQ, C ȕ. 160 OwΌȹŅ1 C r
Q+, +A.EÍŧI#rQLǡ'ψNω£+)aQusds.HʠʵF, LFY 2̊̋ƯǍ.HciõƸĵŤ* H+ȉF.-&) Hψ˟͆+)
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¬

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植物科学最前線 3: 135 ( 2012)ΐÂă.ƾ̯-OwΌȹŅɰŰI) HψHamès et al. 2008ω=#, 1˙Ȝ.Ņ( ), LFY
1 DNA 91˙ę1ɾɯɯɻũpȭ˅I, ]w1 LFY ˙ę`Qq1ȶʣ-©ə-I) HψMoyroud et al. 2011ωɻ̴͸I) # N rQ.Ι)A, ȏͫ.-&), ¬ΐÂƯǍ.
HΎ̯ǃŇģI#ψSiriwardana and Lamb 2012bω
F., ̊ơ+̋ɾ .HõƸȯʏ΃¼Ť1˞˭ʏ-ǫ˓-KI, ̊̋ƯǍΙ͵1΃¼Ť*I=*.õƸȯʏ΃¼Ť+)ěŰI) #A1LĠ@), œǿ1΃¼ŤěŰI#ψWinter et al. 2011, Siriwardana and Lamb 2012aω̋ɾ .HõƸȯʏ΃¼Ť1ěŰF2, I=*©ǈI) -&#ɾɻũʌɾũʏͼʯ.
H LFY 1ɢĽʏƲøɒ6G, ̇ĥ H³ƶ1ʠʵ͈κLǰÇH+.-&#Ȕ$.
ſǿ*2 H, Ύ̯- co-factor A 'ěŰI) HψSiriwardana and Lamb 2012aω

LFY 1ȳ˽1ɻ̴2, 1E ., A&4FaQusds1ʠʵ.EGͶM*#, ̊̋ƯǍ
.HciõƸĵŤ+Àˬ(FșHΙƽF, ʱŤȨɯL¢ƽ.ǿœ1Ug[ěŰI) Hψ2012 Ɲ 6 Ȑ 1 Ȉ1Ȍɦ*, ̦ŤȨɯFˏ 2200, ̬ŤȨɯFˏ 110, ajȨɯF 53, ž̕μF 76, ^\ȨɯFˏ 660ψ#$, ŔĆd_\Ɔ (Sphagnum) 1ʱF1ʞ
A1ω1ΉïʍΓI) Hω #ǫ˓F, LFY ΃¼Ť2, ^\ȨɯLĠ?ΞȨɯ.Ɵ
ŧĽ, ̬ŤȨɯ+·1ˍ˚.̰FIHÑǿÂăBȡ̉ÂζŃ1ẕ̌Ȱ-Ñü.͙ĵH΃¼Ť
Ύ̭LΝ3, Ȭ0ĉ^}΃¼Ť+)ŧĽH+ȉF.-&#I2, ΞȨɯ1Ͷă
1ͼʯ*, ΞȨɯ.ĺȑ1A1AĠ@), œ1͠æĵŤ΃¼Ť, ΃¼ŤΎ̭.EGŔ-~NLƯǍHE .-&)#++2źɩʏ* HψRiechmann et al. 2000, Moyroud et al. 2009ω
LFY ΃¼Ť- ΃¼Ť~NLƯǍH.̆F-&#.' )2͍* H+IH=#,Ͷă1͙ɜ.' )A, DNA ˙ęrQψC rQω1ȭʹ+ DNA 91˙ęȮƨ,'

19lZc-i-9lZc̟ʎ͕ψÆ 3, Tc3A qcfcω1!I+¿) HψHamèset al. 2008ω
+F, qch͙ɜ1Ė˽ǃʤĪI) H,

I1ňę.AΉïʖěǃ

1ƣę 2άƚ.Á, ǭə1ŃLë- ψMoyroud et al. 2009ω*., ]w̴͇Ů¨) Hϋʱ1ˠ̜μ, ZrscψChlamydomonas reinhardtiiω+  lZcψVolvox carteriω.2U
g[ŧĽ- +F, ΞȨɯψȑ˺Ȩɯω.ĺȑ1͠æĵŤ* HĖ˽ǃσ , ˠ̜μ1¢*AabZμψĿƈ 2010ωB^U\oμ, Ǭę̜μ+ &#ΞȨɯ1ŞŜ˯+IH
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
ψPpLFY1, PpLFY2ω.ΙHʠʵ΢ʹ&)ØI#ʠʵ* G, ȏͫ.-&)ʖȵ *î̡I#ϋã1^\Ȩɯ1ǽʪȎ1¢*A, ˑέLø )˕´I) HψCuming 2009, Vanderpoorten and Goffinet2009ω΃¼Ťʌɹzi1́˔-̴ț+, ΃¼ŤʡŏȣLʀ #ȳ˽̴ț-IψTanahashi et al.
2005ƶͭ1Lčɩω, ͠æĵŤ+!IõƸHʌɾͼʯ1Ͷă+̲ɦḞĥɖ˵ŹA-I) HψMaizel et al. 2005ω


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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
japonica, RICE FLO/LFY [RFL]ω, qR^aψZea mays, ZEA FLO/LFY 1,2 [ZFL1,2]ω-,*, LFYU
g[1ȳ˽ȴǵőʇÂ H 2ʌɹǘõÂ, ͼ÷ʌɹÂƷFI, ȳ˽̴ț-KI) HψũĜ1ƶ1Ĝʭ2΃¼ŤĜ [ʅʭ]ω
IF1Ȩɯʱ1̴țF̶ H+2, ;+M,1ňę, LFY2̊
1ȶƚ-ʌɾ.ƾ̯* H, aQusdsBXYgR*ȉF.-&) HE -̊̋ƯǍ
1ciõƸĵŤ+)1ƲøLƾAȜ#) H+2ΜF- +* HÆ 3, qqB
qR^a*2, aQusds1ňę.ͫ ƲøLǝ&) H+>H+*H, ABC΃¼Ť
1 %, AZc΃¼Ť1ʌɹõƸ.ΙKH+Lʤȉʣ-ȥǞ2ƷFI) - =#, aQu
sds1lfy őʇÂȫ@)͢ƺ-̊Ǎ͸Ʀʤ- 1.ź, IF1ʱ1ȳ˽ȴǵőʇÂψq
q1fa+leafy inflorescence (lfi), qR^a1zfl1 zfl2¬ΎőʇÂω2ȉʛ-̊Ǎ͸ƦLʤ-,1Ɠʇ>FIHȆ, ktO1ňę.2, ̊1ʌɾͼʯ.ƾ̯*2 HA11, ̊Ǎ·öF̏βì̧˗˩*Ƭ ʌɹLʤALF2, ̊̋ƯǋƯǍ1#@1õΜ̯ĵ+2-&) - !1#@, ALF
΃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>)
A, Ʉ)Ȯ*2-, ŭȊ-Ĺƨă2΁Ʈ*2- ̊1͙ɜ+Ͷă.źHΙƽF, Ņ·̦ŤȨɯψbasal angiospermωLĠ?œ1ʱ*LFY ΃¼Ť1ʌɹzi̴țI) H, *2
7I- +.HSio ̬ŤȨɯ*2, RSRPlkOψWelwitschia mirabilisω, [vmƆψGnetumω, gomμ1ϊʱψZamia furfuraceaω, QkRψGinkgo bilobaω-,1ˍ˚ψFrohlich and Parker 2000, Shindo et al. 1999 ,ωB, ɺȜμψΒ̕Ȳμω1ǿʱψMouradov et al. 1998, Mellerowicz et al. 1998, Vázquez-Lobo et al. 2007ωF΃¼ŤZt[-I)#!1˙Ȝ, [vmƆLΝ̬ŤȨɯ1]w.2, ͭ1
E ., ΃¼ŤΎ̭.EG, LFYZr1΃¼Ť+NLYZr1΃¼Ťϊ^}'ŧĽH+ȉF.-&#̦ŤȨɯ+[vmƆ*2, NLY Zr1΃¼ŤŗKI#+˵ FI) H

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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
FLO/LFY-like (PrFLL) ĉΧI, ΦǃɺȜψŔ˻ŤĲʳω+ΤǃɺȜψž˻ŤĲʳω.H!I"
I1ʌɹzi̴țI#ψMouradov et al. 1998, Mellerowicz et al. 1998ω
!1˙Ȝ, NLY ΃¼Ť£.ΦǃɺȜá1˺ɸLɾHʱυ*ʌɹHψ#$, Mouradov et al. (1998) .2, ΤǃɺȜ1ž˻Ť̕1̊ˈȼ˔˻*AʌɹH+Aȉ̹I) Hω1.ź, PrFLL ΃¼Ť2Ȕɪ-ΤǃɺȜ*2
ʌɹ̰FI#, ʌɾͲ1ΦǃɺȜ*2;+M,ʌɹ̰FI-&#ψRNA lq̴țω
M. Frohlich 2, IF1˙ȜF, NLY ΃¼Ť2ΦǃɺȜ*, PrFLL ΃¼Ť2ΤǃɺȜ*!I"IʌɹH, +

Ŕ˹-̈ăLƩë, NLYZr1΃¼Ť̦ŤȨɯ.̆Hˍ˚*2ŗKI)H++ęK), ̦ŤȨɯ1̊̊ˈL'HΤǃɾȺİůψž˻ŤĲʳω.ʂșH+
H, ”Mostly Male theory” LǰĬ#ψFrohlich and Parker 2000, Frohlich 2003, Ĩȁ.EH̴͆+)2º̛ (2012) 1A1KGB ω, NLYZr1΃¼Ť2ΦǃɾȺİů, LFYZr1
΃¼Ť2ΤǃɾȺİů+

Ĺƨ2, ͭ1pPOim*AǏɨ+2Ʈ)2=&)F, !

1ƶ.̴țI#ɺȜμLĠ?µ1̬ŤȨɯ1ÆψShindo et al. 2001, Vázquez-Lobo et al. 2007-,ωFAǹǡI- -,, ”Mostly Male theory” .ź)2Ǖñʏ-̰ȆƬ ψVázquez-Lobo et al. 2007,
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

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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,

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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
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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
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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
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2,

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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/?=
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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.
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2007ω
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MIKCCŁ1MADS lZc΃¼Ť1LFY.EHõƸ2, ajȨɯ. )2=$ʣʹ) -&#+˵Ź) HB2G^\Ȩɯ.Hī1̴țÆ* H{mWv_\*2, ͭψ
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HE .-G, ɾȺǍΕ.ΙKHE .-&#, +asU˵ FIHΉÓÂ.H˗˩ɰʇʏ-ʌɹ̰FIH{mWv_\. )A, !1ȳ˽ʏ-Ύ̯ǃ*.˻ŤÂ.ΜFI)
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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
H΢., ^\Ȩɯ1Ĝʭ.' )2, ĨĜψČ̖˶ʀ #ũĜ [= ɹ̡1ũĜ]ω1E .̥̹H
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Parihar 1962ω
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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˔
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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
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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
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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
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* &#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
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I#̼̎ 1862 Ɲ.î̡I#ω2, 19 ˎ1Ȩɯũ.HȏAΎ̯*ήȅʏ-̖Å1'+
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!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
ψ!1̼̎ɮ1˲̼+ˢ˫2, ¶°¶LȨɯ1Ξ91
Ͷë+΁ǀ+

̲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, ·
1E -ǍȜ.Ņ( )ÅǍI#, Ŕũ1ǽŴ.Hǽ˸ʀ1ŎǩĹψWandtafelω1òã̴͆Ȏ

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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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H Goebel 1ǽʪȎψGoebel 1930ω.A, ft_\1ɾȺİů1ʌɾͼʯBȳ˽.ΙH̅͟1ʠʵ.A+(ĹBµ1ʠʵ˶.EHĹœǿǮͤI) H·2, 19 ˎ·Λ.Hʠʵė1Ȭʅ* H, ʹˋİʹċİ1ʌɾͼʯ+, ēˋċF1˻
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

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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
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ƷFI) #ʝ̰L=+@) H!1ƶ1ʠʵǍȜLĒGÜI, Renzaglia and Garbary (2001) 2, 
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ψCarothers 1975, Carothers and Kreitner 1967, Carothers and Kreitner 1968, Carothers and Kreitner 1976,
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1ʠʵ2, ft_\.+,=FI=*=&#-I) - aQusdsBQvL2@+Hœ1pȨɯLǸH̦ŤȨɯ, ίȾLǡ#, ίȾΙ͵1΃¼ŤLŗ&) H+ψMerchant et al. 2007, Carvalho-Santos et al. 2011ω, 1ʠʵ1ʶʎ1Ŕ-Čĵ+˵ FIHΞȨɯ.HȍͻČɻ1ǫɂ+2ʇ-H, ft_\Lʀ #ɲ̅1ʠʵ1Ė˽ǃ.Ɵ&)
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
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植物科学最前線 3: 147 ( 2012)

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
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ì1¶͎΋˒-,ω, ǨëI#ˋ˔˻LĳǾHȳȭψLčɩω.ΙKH¶͎ɿɯ1ęǍ
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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,
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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 
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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

T. Araki-15
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植物科学最前線 3: 149 ( 2012)

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
BSJ- Revi ew 3: 149 ( 2012)

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
Ĕ·.#,Gʚ1.' )2, =$EK&) - E * HƎȘ (2012) 2, ȑː¸ȥ,Φȣ.HˋŤ1ʮþ H 2ͦͰ1˘͝+)ȳ˽HĖ˽ǃLʤĪ) HI2, ³ƶȩ̽:# 9Mτúʏ-¸͆* HGUS B̠Û̟ʎ͕Li+)ʌɹHˋŤLʀ #̴ț-,ƴ#IH ʹċİ1ƯǋBʌɾ.' )2*2ͭ:- ψDurand 1908 .́˔-̹ͤ HƎȘ 2012Ačɩω, ʹċİ1ΗĔ·ͫ.ͪGʚ #ˋŤ2, ʹċİθ·Lͳ&)ċ˔˻.̆Hθ·á·.2
θɞ˔˻ψventral canal cellω, !1ř1ċ˔˻1.2̃ɞ˔˻ψventral canal cellω H, ċ˔˻1Ǎɪ1Ȍɦ*2ƍŏ)˔˻áŷɯLθ·1ͳ͝Œ.Ǽë) HIˋŤ1ͅƩ.ΙK&) HĖ˽ǃ˵ FIHψŔĨ 2012 .EHωˋ˔˻ìăˋŤƯǍͼʯ*2, !1E -ͅƩa[s.źHǀˁˍ1̟ʎ͕1΃¼ŤʌɹH2* H!1ěŰ2ȫ@)Ύ̯-͈κ1'* Hċ˔˻.̆H=*1Ε θ·2, =#, ˋŤΘʽª1ň*A H+IHψĄƏ 1990b .E
Hω, Ɋǉ)# 12, ʇ-HÐÂ.ʂșHˋŤ1ňę2ò+), ě1ΉÓÂÐ
Â+!1ZψĆǿÂ* Hω.ʂșHˋŤΘ.2, ΃¼ʏ-Ɠ2- ψ:)΃¼ʏ.ě* Hω+* Hft_\1E -ʱ. ), ˋŤΘʽªɹųʏ.,1ʯƣH12, =$ȩ̽1Äľ G! * HT. Araki-17
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植物科学最前線 3: 151 ( 2012)g^tl'Vr$ ^\Ȩɯ.HˋŤ1ǾƗ2, Ƚͣʏͫ͜Χ.ΜFI)GψVanderpoorten and Goffinet 2009,
Goffinet and Shaw 2009, ĄƏ 1990b, ƎȘ 2012 Ačɩω, ȑǃɾȺ1ųɹ.2ķΨL½ +œ +˵ FIȞμ*2, ˏϋ/ό1ʱΦΤʇȣ* HψVanderpoorten and Goffinet 2009ω, 1+2, ÐÂΘLΡ)), ΤÐÂFΦÐÂ9+ˋŤLͰGƄHƾ̯Lǉĥ, ȑǃɾȺ1ķΨLF.ō̯ĵ+-&) Hü ), ȅ ɾ˸ňǑ.ͶëH΢.2, ƾA ǃ1ÐÂǯ +2
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NR$.5+*¡: Ǎɪ#˻ŤLA'Φȣæʘ1Ć·ì*χ̉.̰ H1˻ŤʜƂ2*.ȟI) HΤȣ1ΤİƠLʤæʘ1Ć·ì.2, =$̍ Τȣ+ΤİƠ̰ H¢: ¡1ͫ.̰FI#,ēˋ*-&#Φȣ1>1˯̔˻ŤĲ2Ȕʌ;*, Ƞ2¾6, Ǣɱʷ͙2ĝ1==ΦİƠ2
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̲ɦF, ˋŤ1ǾƗ.̆H=*1ͼʯψE6ω2, B2GŔ-Ύ̯ǃLǡ'

A1* H+̶ Hēˋͼʯ.Ι)2, ɾɯũʏ.EĠĥɖ , ˋŤ1͘ăǃBͺþǃ.Ɋʔ
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A]wyXh ·., Ȩɯĺȑ1͠æĵŤ LFY +, ft_\. ) LFY ΙKHĖ˽ǃ HȑǃɾȺͼʯ1ΤÒ1·ì.' )Ȭ͆#ɹĽ H 2IF1ʠʵ͈κ.' )2, Ðò1˂Ǒ*̶Ď#

T. Araki-18
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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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ψ·, ʠʵΗşȌF1ßěʠʵ˶ω, ˮĨʨ˼ĊŐψɹaXŔũω1;, ̮ɑʺĊŐψ±ΈŔũω, ƈΏΟƿĊŐψ±ΈŔũω1ýúL¹ * H=#, ƎȘȶȲĊŐψƟƊŔũωF2œ1ǽʤ+ǹǳL #$ ) H̹)Ǌ͎1ǉL̥# Ȗʲ1ÅǍ. #&)2, ðʲ1ȻΠ*, ̞μ1ìμÂˍ1ǔ Ȇ, ^\Ȩɯ1ĕ ũĜ+ɹ̡1ũĜ1źǀ-,L2@+), ƎȘȶȲĊŐFœ1^qL #$ #I.E&), '1cLȔɨ.Κ
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