!" -6 (*Ùû ƯſěèïèïǵŤïźƆƁ
657-8501 ſěÿř½ªŪÉū 1-1
A strategy for the colonization of land by plants: the evolutionary transition
from charophycean green algae to early land plants
Key words: bryophytes; charophycean green algae; gene evolution; land plants;
origin of sporophyte generations.
Hidetoshi Sakayama
Department of Biology, Graduate School of Science, Kobe University
Rokkodai 1-1, Nada-ku, Kobe 657-8501, Japan
0/  ęİňų*>Ǔǝ)ŀŞ'-Ȋ? )ƬBÑ >ƹíŀŞBđ ő 1>3ȊĬǶBǀ>'SZŀŞ8PNŀŞŦ % =>+; *ǷØBǀŕ-çł)ŀŞŦ % >īȊŖǼB+ǜ6-Ƿ',©Ǩ ŀŦBǀ>'&>ŖĄ*,ƥȄƲBăèÚ+ŀŞ7Ŧ % >Ȋ+ÎǛ*,9<9<'ƐŠ+ƵȁƝư% >Ō*,őǦ>ø)ƵȁƝŇ% >ü8ƇƟ?-+ƸǼ*7
śĦ+ƵȁŶ%>

ę+ų*ǃ?>',İǡ3?&>Ȋ+; )Ƶȁ!Ō&,ć&>
%Ȋ+*ƒ 4.8 ¥āĮ*Ō <ŝǷØ0Ǥ±ȊǷŀŞ0'Ǥ¼ƵȁÕȋ SuTLtȁȈX`pLxSuTLt Chara leptosporaȉ+ĆėȈSakayama et al. 2009, J. Phycol.;=
Wiley-Blackwell +ǅÈBċ%Ǖǘȉ
 a. ǖ*ĺĥöŠ*ǚŦȊǔǖ+; *ǀ >ƲŠ+ǫb. ĺ+Ŀ¦ǫ²+ĞèÕĺ+*ţŠ+ =+; )ƓƧȈĜƲ°ȉ>c. ǖƸǼ+ĞèÕ[v~jŠ+ƓƧȈŰúƓƧȉ¬£+ƓƧB Ǎ>; *C& >d. ĺ+Ǡ* %>ŦŇÓò+ĞèÕŦÄÓȊǢƏÓŦÄÓ¬+ÄƓƧȈÇƏĉ*ġËí*)>ȉ,ļȂƓƧ& Ǎ?% >e. ġËí+ǻíȀčǰ¯ŵŝ*Ƥ <?>Å â&ƿA?% >f. ġËíƸǼBĞèǻíȀčǰ¯ŵƄ*;"%ŃłǨ Ȋ²ȁ>&ĭż)ÞŘ')>ŁǢ>?3&+źƆ+ƖĹ&,ȊSuTLtƵȁ+SuTLtȁ (Õ 1) 'PzHN~_ȁ (Õ 2)ǷŀŞ*Ĳ7ǝƛ 'Ƣ <?% > (Õ 3) (Karol et al. 2001, Delwiche et al. 2002, McCourt et

H. Sakayama - 1

al. 2004, Sakayama et al. 2009)ĵƅ&,ȊSuTLtƵȁ <ǷŀŞ0+Ǥ¼BȊŦŏť'µ* >Ǥ¼*ŜŚBƗ"%Ɣ>ÕȌ PzHN~_ȁ+ĆėȈDelwiche et al.
2002, J. Phycol.;= Wiley-Blackwell +ǅÈBċ%Ǖǘȉ
a, b. PzHN~_}HyiKvwxU (Coleochaete orbicularisȉ
a. ­ŲŠ+ƲŠ+ǫUN~yg~, 100µm
b. ġËí+ĞèÕġËí,ļȂƓƧ&Ǎ?% >UN~yg~, 32µmc, d.
PzHN~_}kyic~YȈColeochaete
pulvinataȉ
c. ĺ² ?ƲŠ+ǫĺ+§Ɗ*ŦÄÓ>UN~yg~,32µmd.PzHN~_ȁBşĎ$>Ŋ+; )ŁǢ+ĞèÕ
UN~yg~,10µm

Õȍ SuTLtƵȁ'ǷŀŞ+ƑƘǴȈMcCourt et al. 2004, Trends Ecol. Evol.BĤåÙû  2009,Biophilia BĤåElsevier 'DbUx~+ǅÈBċ%Ģǘȉ
SuTLtƵȁ*, 6 #My~kȈSuTLt
ȁȊPzHN~_ȁȊmSqb{ȁȊLzjXyq`EGrȁȊL{{KjUȁȊsXU_EMpȉ>
+Õ,Ê*Ǥ52(ǷŀŞ*ǝ 'BĕÏ% =Ȋ
Ĳǝ+źƆ&,SuTLtȁĲ7ǷŀŞ*ǝƛ'Ƣ <?% >ĺ+*,Ǥ¼+ǧƃ&šċȈȇȉ3,ŒéȈǑȉĆėůşĎBŽ%>

H. Sakayama - 2

1/%. -!" +
Ĳ³*ȊSuTLtƵȁ'ǷŀŞ+Ŧŏť'µ+Ǩ BħŤ>ǷŀŞ,
1%Ƨí'Ð-?> 2 ¢ Ȉǁ*ũĶ> 2 ƕ+ĻƫBğ#ȉ'Ǭ¤'Ð-?> 1¢  <)> Bƶ"% >ȈÕ 4ȉƧíȈ2 ¢ȉ <ŔĦ²ƺB%ŦƧíŭƭ%Ǭ¤Ȉ1 ¢ȉ&>ǷŀŞ+Ƨí'Ǭ¤,(!<7çƓƧ&>
PNŀŞ&,Ƨí;=Ǭ¤+īèĆė7ƼǺ&>ȊƹíŀŞ*ǝ$*Ċ"%èǟǕ%Ƨí+īè)=Ćė7ƼǺ*)>īȊSuTLtƵȁ&,Ǭ¤,>Ƨí,)(Õ 4)#3=ȊǬ¤Ȉ1 ¢ȉÄ'ƏíB=ġËíȈ2 ¢ȉ&>Ȋŭƭ>'ȈƓƧ²ƺB*ȉ*ŔĦ²ƺB%3 Ȋ®/Ǭ¤*)=ȊƧ
íȑȒB<) SuTLtƵȁ+Ǭ¤,ÀƓƧ "=çƓƧ "=>SuTLtƵȁ+&ŐŦů)SuTLtȁ+Ǭ¤,ȊƼǺ)Ņ¦ŁǢB% > (Õ 1)

ÕȎ ƚƫŀŞ+Ŧŏť'µ+Ǥ¼ȈÙû  2009, Biophilia ;=DbUx~+ǅÈBċ%Ǖǘȉ
 SuTLtƵȁ&, 1 ¢+ī 2 ¢;=7Ŧŏť*Â6>įǳǱȊµ7è 7+ç ȊƹíŀŞ*ǝ$*Ċ"%èǟǕȊ2 ¢+īè)>3ȊhsqI^Kt'SuTLt+ǳ&1 ¢+çƓƧ¼ǒ=ȊSuTLt'hs]xJeQN+ǳ&2 ¢+çƓƧ¼ǒ"% >

2/!" -+; )Ŧŏť'µ* >Ǩ

<ȊŀŞƈ%ǷĚ,Ņ+; *Ƣ<?% > (Graham 1993)çƓƧ+Ǭ¤ +ŦŏťBğ#SuTLtƵȁȈSuTLt
ȁ3,PzHN~_ȁȉġËíȈ2 ¢ȉ*%ŔĦ²ƺBƶ YFq|MB§Ǟ=*% Ȋ+ĉȊŔĦ²ƺ+¶*ƓƧ²ƺBƶ ; *)=ȊĪ*ƧíBšċ'
Ě&> (Õ 5)
&,ȊŀŞ, " (+; *%ȑŔĦ²ƺ+YFq|MȒ'ȑ2 ¢+ƓƧ²ƺȒBµČ+&@

+ŬÒ*ƍ >6*,ȊŢŦ+Ĳ³+ǷŀŞ&>PNŀŞ'+ž§+SuTLtƵȁBĆė'ǩí+zly&ŉǗ>ďƾ>

H. Sakayama - 3Õȏ ǷŀŞ* >Ƨí+ǒŗȈŕǪ}ß 1996 ;=¬ŨơȆÖ+ǅÈBċ%Ǖ
ǘȉSuTLtȁȊ3,PzHN~_ȁ*ȁçƓƧ+Ǭ¤ +ŦŏťB
ğ#Ƶȁ*%ȊġËí+ŔĦ²ƺ+ǲìǥ?Ȋø)çƓƧ+ 2 ¢ ȈƧíȉĆĘ?ȈĪŦǉ'Ð-?% >ȉ
3/, )
 PNŀŞ* >ƧíȈ2 ¢ȉĆĘǴǣǩí+ȋ#* FLORICAULA (FLO) / LEAFYȈLFYȉ
ǩí> (Tanahashi et al. 2005)FLO/LFY ǩí,ƹíŀŞ+S{FdcVc&,Ƭ+ĆėĆĘ*A> MADS n\LUǩí+ŭŢBǈ÷>Ǖ¯Ôí&>Ĳǝ # +ŦŞ&

Ǥ6<?OfrȈ©ǩíȉǂĸ+ƖĹ&,Ȋĵǩí,ǷŀŞ+4 < î×àÍ?% ) ĵǩí,ȊPNŀŞ+hs]xJeQN*%ȊÇƏÄ+Ƌ²ƺ*ƊBŭ> 2 ¢+ƓƧ²ƺ©ƪBµČ>Ôí&>'ĭ< *)"% > (Tanahashi et al. 2005)(Õ 6)

ÕȐ hs]xJeQN* >ƧíĆĘǧƃȈTanahashi et al. 2005, Development BĤåÙû 2009,Biophilia ;=ǕǘThe Company of Biologists 'DbUx~+ǅÈBċ%Ģǘȉ
a-h. ǢÄÓ+ƜĩǼÕa-e.
ǮŦĽȊf-h. FLO/LFY ǩíB1%ŻãĽȊi. ņĀ)ƧíŷÃȈȉ,ľȊŷ=Ȉ  ȉ,Əía, f. Əí¨¶Ȋ b, g. Əí¨ĉȊc, h. ÇƏĉ*ĆĘ?ġËíȊd. Ĳ³+ƓƧ²ƺǒġËíȊe. 2 ƓƧ+ƦǮŦĽ&,ÇƏĉ*ġËíĆĘ?>'ȊƓƧ²ƺì3=ƦBĆĘ%Ƨí0'ĘǱ>īȊFLO/LFY ǩíB1%ŻãĽ&,ÇƏĉ*ġËí,ĆĘ?>ȊƓƧ²ƺǒ<ņĀ)ƧíĆĘ?) 

H. Sakayama - 4"%Ȋ+Ôí(+; *Ǥ¼%+ Bĭ< *>',Ȋ2 ¢+ƓƧ²ƺ+µČSU_r+Ǥ¼BŤǂ>*7S{FdcVc*&ǭƾ')>'Ĕ?>FLO/LFY ǩíæ%Ʊǽ²ƺƕƞĆĘ*ǭƾ)ć·BĹ%KNOTTED1-LIKE HOMEOBOX (KNOX) ǩí,Ȋ
hs]xJeQN*>LwUȋ

% 2 ¢+ŭŦ*Ǵ

% >'àÍ?% > (Sakakibara et al. 2008)SuTLtƵȁ*%7+; )ǩ

í+ǂĸBƶ ďƾ=ȊŢ×źƆǤ6<?% > (Tanabe et al. 2005, Floyd et al. 2006,Floyd & Bowman 2007, Nishiyama 2007)3 źƆÞŲ,¾²&,) ȊĲǝȊSuTLtƵȁ
+ǫ&,æĶǩí+÷¨7Èƨ*)=##>6ȊĉȊ+; )ǩí+ńƨBȊŦ
Bŧ %ǂĸ>źƆ+ùǲ7ĴĈ?> (Abe et al. 2008)

4/%. )   +; *ȊSuTLtƵȁ*,ǷŀŞǇŦ+ǋBǂĭ Ƃõ+ēàCǹ
?% >"%ȊSuTLtƵȁ* >²íǩïůźƆÞŲ+żƈō6<?%>+6*,Ȋ1) ŦŏťBŸĴǳ*ðƖ&>ÝȂƑ+żƈ2) ĆǐǕģƑ+żƈ
3) OfrȈ©ǩíȉǂǊȈĲǝ,ƎÀ*Ǌ6>; *)=##>ȉ
ďǾ&=ȊƳƣ<+źƆMy~k,+ôŢ*º¸% >?ôŢįȊSuTLtƵȁ,ŀŞǤ¼ï+è)jzFLUy~B7<'ĴĈ&>

5/  ĵƅ&,Æ=%, ) ȊSuTLtƵȁ+&7è)Ǥ¼ #7ǒ"% >ȈÕ 3ȉ
' -ȊSuTLtȁ'PzHN~_ȁ,çƓƧ+Ǭ¤Bğ!ȊÄ'ƏíB>Ȋ?<;=ž§ů)ĒǐBğ#hsqI^Kt)(+ġËƵȁȈmSqb{ȁȉ+ǳ&,ȊǬ
¤,ÀƓƧ3,ÌĆ+ƓƧǣ)"Ơ+; )µ&=Ȋ<?> 2 Ƅȁ+Ǭ¤í,ȊĆė <,Ä'Əí+; *½´>'&)(Ǵĵ 2002)+; *ȊǤ¼ïů)ȅ¸BCğ"SuTLtƵȁ+źƆBĐȊű=%?>Ʈƣ+ŮáBƀ,Ĉ!
ĳC& >
&' ĵƅBÜƌ>ǧƃ&ȊƽûıĭÁäȈǯŎèïïǸƁïôȃW|Y~ȉȊǴĵąÁäȈĬĵëíèïŤïǫȉ
Ȋƴ¦þÁäȈķèïèïǵƙËĨ¼źƆƁȉ <ç+¹ǄBǽ DbUx~+ŹñĂŋ <,ĵƅ+6*ç+Õ'ĨƉ+ǕǘǅÈBǽ ĵƅ&ƔƳƣ<Ţ×ƶ"% >źƆ+ǫ,ȊĬĵïƷĠƩƁïźƆǏƻ¹ǯȊĨǫƁïŴƁïźƆǏƻ¹ǯşóǿÛźƆŉǗOfrȊȈǎȉ.: ƁïĝƷ¿ê»źƆ¹Ę+Ro~aBÇ% >*ȊœĖǌ+ĕBƸ H. Sakayama - 5$ #Abe, J., Hiwatashi, Y., Ito, M., Hasebe, M. & Sekimoto, H. 2008. Expression of exogenous genes under thecontrol of endogenous HSP70 and CAB promoters in the Closterium peracerosum–strigosum–littorale
complex. Plant Cell Physiol. 49: 625-632.
Delwiche, C.F., Karol, K.G. & Cimino, M.T. 2002. Phylogeny of the genus Coleochaete (Coleochaetales,Charophyta) and related taxa inferred by analysis of the chloroplast gene rbcL. J. Phycol. 38: 394-403.
Floyd, S.K. & Bowman, J.L. 2007. The ancestral developmental tool kit of land plants. Int. J. Plant Sci.
168: 1-35.
Floyd, S.K., Zalewski, C.S. & Bowman, J.L. 2006. Evolution of class III homeodomain-leucine zipper
genes in streptophytes. Genetics 173: 373-388.
Graham, L.E. 1993. The origin of land plants. John Wiley & Sons, New York.
Karol, K.G., McCourt, R.M., Cimino, M.T. & Delwiche, C.F. 2001. The closest living relatives of landplants. Science 294: 2351-2353.
McCourt, R.M., Delwiche, C.F. & Karol, K.G. 2004. Charophyte algae and land plant origins. Trends Ecol.
Evol. 19: 661-666.
Nishiyama T. 2007. Evolutionary developmental biology of nonflowering land plants. Int. J. Plant Sci. 168:37-47.
Sakakibara, K., Nishiyama, T., Deguchi, H. & Hasebe, M. 2008. Class 1 KNOX genes are not involved inshoot development in the moss Physcomitrella patens but do function in sporophyte development. Evol.
Dev. 10: 555-566.
Sakayama, H., Kasai, F., Kawachi, M., Watanabe, M.M., Nozaki, H., Nishihiro, J., Washitani, I., Shigyo, M.,
Krienitz, L. & Ito, M. 2009. Taxonomic reexamination of Chara globularis (Charales, Charophyceae)
from Japan based on oospore morphology and rbcL gene sequences, and the description of C. leptospora
sp. nov. J. Phycol. 45: 917-927.
ÙûƯ}ƽûıĭ}Ǵĵą}ƴ¦þ 2009. ǷŀŞ+ž§SuTLtƵȁ+Ǥ¼ïŌ <Ƿ0+Ua~x~. Biophilia 569-572.
Ǵĵą 2002. ƀ+ 6ôȃŦŞ —qI^Kt—. ƓƧýï 21: 1230-1231.
Tanabe, Y., Hasebe, M., Sekimoto, H., Nishiyama, T., Kitani, M., Henschel, K., Münster, T., Theissen, G.,Nozaki, H. & Ito, M. 2005. Characterization of MADS-box genes in charophycean green algae and its
implication for the evolution of MADS-box genes. Proc. Natl. Acad. Sci. USA 102: 2436-2441.
Tanahashi, T., Sumikawa, N., Kato, M. & Hasebe, M. 2005. Diversification of gene function: homologs ofthe floral regulator FLO/LFY control the first zygotic cell division in the moss Physcomitrella patens.
Development 132: 1727-1736.
ŕǪ¡}ßǙȈ«ǆȉ1996. ǷŀŞ+ǒŗ. ¬ŨơȆÖ. ķ.

H. Sakayama - 6