植物科学最前線 7:87 (2016)

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1

ûǣ²ƳƨĂĲǮøĂȼøĂ kEITEH WƢƮƨ
630-0192 ûǣƝƑɑĔɓđƕ 8916-52ƏƢ ƐòȊžƨĂƢƮY \

230-0045 ƧûēƝŤŸĔɔǶÉŒĘƕ 1-7-22
Misato Ohtani1,2 & Taku Demura1,2
Evolution of water-conducting cells in land plant;
lessons from VNS family gene encoding NAC transcription factor
Keywords: land plant evolution, NAC transcription factor, VND, VNS, water-conducting cell1

Graduate School of Biological Sciences, Nara Institute of Science and Technology,
Ikoma, Nara, 630-0192 Japan
2

RIKEN Center for Sustainable Resource Science,
Yokohama, Kanagawa, 230-0045 Japan NAC Ȓ¶ãÿ*şƅƆƖƘ'Ȓ¶ãÿǔ$;, ƗƑ8ƾǜºÈ, WcWĨƷ'&ĖĘ şƅƑƏȣƩ(ȸ# <%Ơ:=# <œƫ$*, țűƾǜɕwater-conducting cellɖºÈ)v
W\WE_^%ǖ :=# < NAC Ȓ¶ãÿ VNS nBw)Ŏņ)ƠǶ@7%(, Ⱦşƅ
)tbDp ȟÈ)ȣƩ$şƅƆƖƘ'Ȓ¶ãÿ%)Ȓ¶¾ĤWLx&!ĞÂ@Ś#) (" #ǖĉ  ZRFLM êƎÔ, şƅȾƐò/)ȟ¹%ȨĨ@įȠ%*, ƍé)êƑĮæ("'<ȱǵ'ƴŪ$!%ǖ :=# <=3$ű$ō:# şƅȾÈ<(!#,áɌ%'!ƐòǵƼ*33ǖ :=<, $7Ƒā(ħɊ'ű)ƣ¨ø'ȃɌ%'!
%*Ĭ¯(Ʌ' Ȼ:=űȊž/)ȨĨ)ǂŚ%#, Ⱦşƅ*ű@¡µȕȚ<%(ƆÈƾǜ, țűƾǜ@Ƒ4¹(Ǡ! ƍé)Ⱦşƅ7",

7'țűƾǜ@äɘ(Ĺțűƾǜ(´ț)Ɔĥ*,=7ū

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ƍā)țűƾǜ)$7!%7ȟÈĜ%ǻ <), ǰÿşƅ7"ȤƸƾǜ$<ȤƸƾǜ*ƶƇ)ĜƇ@ƾǜ$, )ƾǜ%"'!
#ƸƇǀǑ@Ĝį<ɕäɘaɖȤƸƾǜºÈ)ȣƩ$*, )ƾǜ%)"'ƚ(ưĀɕA ɖ%Ý+=<ă³(ƾǜôºǺ=ø'ĀƯ, ű@ÅƌƘ(ȕȚ$<9 ('!
# <=(Ċ#, ǲÿşƅ8U]şƅǵ'țűƾǜ%# <)ȤƸ%Ý+=<ƾ
ǜ$<ɕäɘbɖȤƸ*ƽȳĜ)ƾǜ$;, ƾǜ×õȞǂ<%*'Ɂ)ƾǜ%Ļ
M. Ohtani & T. Demura-1
BSJ-Review 7:87 (2016)

植物科学最前線 7:88 (2016)<ɇ(ôĀɕ/ ɖ%Ý+=<ƭ@÷łĜį, )ôĀ@ț
#ű@ȕȚ<ǢƘ(ȤƸƾǜ*ȤƸƾǜ9;7ǩȵɕÈơƪ$*ȵ 3 cm (ȥ<ȤƸƾǜ7Ƕ"!#<;Cichan & Taylor 1992ɖ, ƾǜğ*ȤƸƾǜ)=%Ű0#Č =*, ȤƸ)ñÖ(*űȕȚŦǝ}f_cƾǜÊ $<
6, ƾǜȵȵ ŇțűÅƌ<%%ǖűƏƘ'ƏƔ(9<:=#< ɕ Tyree &Zimmermann 2002ɖ
Ċ#ȤƸ)ñÖ(*, ȤƸƾǜ×õȞǂƸƇǀǑțűŦǝ}f_c%#®6, ğ)ø'ƾǜ@AȞǂ#ȵù Ƹ@£<%$9;țűÅƌ@Ŋ<%ȨĨȟÈȌ!

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*ȵȓŇØ(88ȵ ĜĮ@# <%÷, ɘɖÌĜȋȞǃƔŘ%'!Āāé<ŏĀ\Ep%, əɖĀ*Ĝį=' ƁĀ\Ep, Ơ:=# <Ǥɍ%Ǭɍe V{z V{SPǈ)ȫ)ƪ7")* 7(ɘɖ$;, ǬɍvSPǈ)ȫ*əɖ@Ɨȥ# <%
(9Ɨȥ# <ñÖ(*RPşƅ)țűƾǜɕǀǑɖ*jEdEd%Ý+=<%;, ƉǕ(əɖ)ƁĀ\Ep(Ȼ!#jEdEd%Ý.ñÖ7<ɕLigrone et al. 2000, 2012ɖįƃ
jEdEdƾǜ)TEX*, ƪ(9!#øƖ'!# ;ɕ200~1,500 µm ȵ, 10~25 µm ğɖ,ª¡TEX8¡¾)ƪƆĪ%ȸȞŹ %Ơ:=# <ɕLigrone et al. 2000ɖ Ŏġ(, țűƾǜ)Ɔĥ@3%6<$, ƾǜô©ɏ)áɌ@Ï;#  ǇƸŖşƅ
7"ȤƸȤƸƾǜ)Ɔĥ%#, Ë ŨƾǜôɕŨôɖ@ǪƬ<ƀĹ:=<Ũô*ŨôɕƾǜºǱ)%(Ƒ4¹=<ƾǜô$, ³#)şƅƾǜ7!# <ɖ%ƾǜǞ
)ȷ(ŨƘ(Ĝį=<ï šȝ¡$;, YW, (LU '&)qwYW,
#Of % !kEIuv, Ũô%*Ɩ'!ěå'ɓŨšȝ@%!#ǪƬ#<Ũô*ƾǜ(ě ŦŞƘěę8ǘűĪ@<%$, =, ȤƸȤƸƾǜ( <9;ɓ țűŦǝ)ã$<%ǖ :=# <Ċ#, RPşƅ)țűƾǜ)ƾǜô
*ƪ(9!#øƖ'!# ;, ÛȖ)¡ƾǜ9;7ǫ ƾǜô@7"ñÖ7=+, ŋƟ'Ǜ
Ë@ƥñÖ7<ɕLigrone et al. 2000ɖǛËƾǜôȌ<ñÖ(7, ¥ +WMSP)
M. Ohtani & T. Demura-2
BSJ-Review 7:88 (2016)

植物科学最前線 7:89 (2016)

jEdEd)ñÖ(*, ŨƘ(ǛË<?$*' )$, Ũô%*Ý0' %7ðÚ=# <ɕLigrone et al. 2002ɖ
:(Ā)$Ň(7ø'Ȧ ;, ȤƸƾǜ$*ôĀ*ÌĜ
ȋȞǃ%*ƛĻȸȞĪ', Ũô)ŴƞȹĈ=<%$Ĝį=<Ň$, RPşƅ)ŏĀțűƾǜ)Ā*ÌĜȋȞǃ@)ƔŘ%# <ɕLigrone et al. 2000; Tyree & Zimmermann
2002ɖRPşƅ)țűƾǜ)ñÖ(*, ƪ(9<kHU~ ĖĘ 6Š(*ǻ ' ,
č'%7ƾǜô©ɏ(" #*ǇƸŖşƅ)țűƾǜ%)ƜȦƀ7÷ 9 $< )9 (, ƍǭȾşƅ)țűƾǜ$<jEdEd, ȤƸƾǜ, ȤƸƾǜ*,
țű%

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3(ųƞ" # ' áɌ$*<,ŎȘ, ƵǗȥ)Ȟ)ƢƮ :,  ȟÈƘ«ɇ(ȸ<ƥß@7:ǺřǂŚģ:=#œƫ$*, Ⱥ, ƵǗȥ)ƢƮįŚ@Ħ(țűƾǜºÈ)ºÿ¾ĤŦš)ƠǶ@ƿ"", țűƾǜ)ȟÈ(" #ǖĉ<
VURFLM9;1)*6)$,+Z  0#256
 țűƾǜºÈ(ȸ<ºÿŦš)ƢƮ*, (ȤƸƾǜ@\Q_c%#ǭ?=#ºÿƑƅĂƘƢƮ)Ƴǌ@ȶ )*, m{Nf^[FǨǙƾǜ :)şƅsz źÄ(9<ƛĻƘ'ȤƸƾǜ/)ºÈȒľȀċƺ$;, )ƺ(9!#÷)ȤƸƾǜºÈȸȞȪÿ)×ą8Ŧ
ǝǺřǭ?=ɕFukuda & Komamine 1980; Demura et al. 2002 '&ǿƾ* Fukuda 1996 8 Turner
et al. 2007 )ǋȁ%)(Ě =# <ńƊ@ÍƂɖ3, UEgeXe8up% !zbşƅ)ɎȐƘ'ĆɒƐòŃ­(!#, ȤƸƾǜºÈ8ŨôĜį%ȞÆøǷţc W
Npcxb\ǪƬ, Ȫÿīð)ķ±ǭ?=ɕ¥ + Sterkey et al.
2004; Turner & Somerville
1997 '&ǿƾ* Nakano etal. 2015 '&)ǋȁ8)(Ě =# <ńƊ@ÍƂ)%ɖ
  , 2005 ė, U
EgeXeíɐƾǜ@Ŕ
Ņ%ȤƸƾǜºÈȀċ
ƺ@Ɠ c WNpcxǺř(9!#, 7 ")Ȫÿ:šį=<

NAC Ȓ¶ãÿnBw, VNDɕVASCULAR-RELATED NAC-DOMAINɖȪÿnBw×ą=ɕKubo et al. 2005; äəaɖǿƾ' VND ȪÿŦǝǺř)ǂŚ, VND nBwµ$7%(VND6 % VND7 , ==, ǉƚƇ)Ũô@7"ġƑőȫȤƸ%:AƇ)Ũô@7"ÌƑ
M. Ohtani & T. Demura-3
BSJ-Review 7:89 (2016)

植物科学最前線 7:90 (2016)őȫȤƸ)ºÈ@¾Ĥ<vW\WE_^ɕȴȪÿɖ%#® # <%ŋ: %'!
ɕKubo et al. 2005; Yamaguchi et al. 2008; äəbɖ
VND nBw)×ą)ðÚ :čȡ=#, ȤƸƾǜ%×Ţ(ĜįĐºǱǀǑ :£:=<őȋƾǜɕOf @Ù5Ũƾǜô@Ɨȥ<ƾǜ@őȋƾǜ%Ý.ɖ$<őȫǏǇƾǜ)ºÈ¾Ĥ)vW\WE_^%#, VND nBw)ýüȪÿǔ(ď< NST1 (NAC SECONDARY WALL THICKENING PROMOTING FACTOR1)
9- NST3/SND1 (SECONDARY WALL-ASSOCIATED NAC DOMAIN PROTEIN1) ×ą=
ɕZhong et al. 2006; Mitsuda et al. 2007; äəaɖ
őȫ@šį<ȤƸƾǜ%ǏǇƾǜ*êkEIvW)øȫº@šį<őȋkEIvW)ǵ¦Ǆž$<% :, VND

9- NST/SND nBw

=:)ƾǜ)ºÈWE_^%#×ą=%*, îƤƘ(7ĨƓƘ(7ȱǵ'ƗǶ$
!
 U  E g e X e $ * , VND

9-NST/SND nBw)ýüȪÿ%#,:(ɚ") NAC Ȓ¶ãÿāé# <ǡÜŹ %(, =: SMBɕSOMBREROɖ,
BRN1ɕBEARSKIN1ɖ 9- BRN2 *, U
EgeXe)ŝ·ƾǜ)ºÈįƃ@¾Ĥ
# <%ƥ=ɕWillemsen et al. 2008;
Bennett et al. 2010; äəaɖ:( SMB, BRN1
9- BRN2 Ȫÿ)ȣÁƗƍ*, VND 8
NST/SND nBw)Ȫÿ@ȣÁƗƍñÖ%×Ţ(, ȤƸƾǜŢ)ŨôŴƞ)ƖİƘȀċ@ĚȌ%7º!

ɕBennett et al. 2010ɖ)ǂŚ*, VND,NST/SND, SMB Op :'< NAC Ȓ¶ãÿToOp*´ț#, ȤƸɕŢɖƾǜ)ºÈ@Ȁċ<ſéƘ'ǝÃ@ŏ#<%@ƥ# <'? , şƅȟÈ)ŉ ůȿ$)ToOp)´țƦ²%'< NAC \ lNȋȤƸƾǜºÈ¾Ĥ
ǝÃ@Ƌģ# ;, )ġ, ȟÈȟ5(
Ģ!# VND, NST/SND, SMB ÕOp)ŦǝºÈȌ!%Ĭ¯$< ºÿŦǝ)´țĪ :, ǩǗ *, )To O   p @ VNS ɕ VND, NST/SND,
SMB-related proteinɖnBw%Ý.%@Ľà# <ɕOhtani et al. 2011; Xu et al.
2014; Nakano et al. 2015ɖä 3 (*, QixǺȂǭ?='şƅƪ( < VNS \ lNȋ)ºÿƺǅť@ƥɕNakano et al. 2015ɖ
M. Ohtani & T. Demura-4
BSJ-Review 7:90 (2016)

植物科学最前線 7:91 (2016)

VNS \ lNȋ*Ⱦşƅ(Ę¨ā=# ;, RPU]şƅ) VNS \ lNȋ%)ȸ§Ī'& :ǖ #, şƅȟÈŎ½(¹ƍVNS \ lNȋ*, VND \Ep(Ș ȭ¼@ĸ!#%ļż=<3, ƍé3$(Qixȭ¼ǺȂ=# <ǲÿşƅ(*, NST/SND Op(ď< VNS āé# ' ɕNakano et al. 2015ɖǰÿşƅƆƖƘ( NST/SND \Ep)
VNS ȪÿȟÈ) , < *ǲÿşƅƺǅ$ NST/SND \Ep) VNS Ȫÿŧǧ), ġ)ǿƾǺřħǵ$<, )Ć*ȤƸ%ȤƸƾǜ)Ȧ @ǖ <$7ǡÜŹ
ƀ$< 2010 ėÀġ(*, Ǵł)ƢƮ
Op(9!# VND69-

VND7, SND1/NST3 )]EN
c\Q_cȪÿǔŋ:(=ɕOhashi-Ito et al. 2010;
Zhong et al. 2010; Yamaguchi et al.
2011ɖ=:)ƢƮ(9!#,VNS \ lNȋ(9!#ŨôĜį8ƾǜū(ȸ?<ȪÿƛĻƘ(ũ)Ɨƍ¾Ĥ@Ð#<%×Ō(, MYB @Ħ(÷)Ȓ¶ãÿ VNS )ŷ$Ŧǝ# <%ƥ=ƍéǖ :=# <, UEg
eXe( <ȤƸƾǜ)Ȓ¶¾Ĥh_cNzb@äɛ(ƥUEgeXe, up
, }J, Eh, oLub
DFx, \FvS|U, cF
zRU'&zbǰÿşƅ) VNS Ȫÿ)ŦǝǺř :, =:)şƅ$*3 (ƴ)vW\WE_^ VNS→ƴ)vW\WE_^ MYB→ȤƸƾǜºÈ)ŷ=¨ā=# <%º !# ;, č'%7ǰÿşƅ(#*, )ȫº@țűƾǜºÈ)Ȓ¶¾ĤRCW

Lx%Ƕ'%$ $<:(UEgeXeƢƮ :, VNS )ŷŷ%7(÷)Ȓ¶ãÿāé, =:Ȓ¶ãÿȒ¶r\ lNȋr$Ǵɂ(Ɯ£Ɠ':, Ȩ»'ƾǜºÈ)¾Ĥ(Ŧǝ# <%º ;""<ɕYamaguchi & Demura 2010;
Nakano et al. 2015; Taylor-Teeples et al. 2015ɖțűƾǜ*ūƾǜ$;, ƾǜºÈ<ƩęȟA$3 %ƾǜȢÞ@ö <%*¹Ř' )6, ºÈ)vW\WE_^ VNS )ŶĪ@¾Ĥ
<6(Ǵɂ "ƹǍ'ºÿyJfXxƗȥ# <) 7=' 

WU/4-5%.('  ENOC

9



 #, äɚ)ºÿƺǅťƥ% ;, VNS nBwȪÿ*ÌþƘ'Ⱦşƅ%=<RP
M. Ohtani & T. Demura-5
BSJ-Review 7:91 (2016)

植物科学最前線 7:92 (2016)şƅQix(7āé# <$ǩǗ:*, VNS Ŧǝ(ȸ<ȟÈƘ'ƠǶ@ģ<6, zbRPşƅ$<my`KhSPɕPhyscomitrella patensɖ@ŔŅ%Ǻř@ǭ!#my`KhSPQix(* 8 ") VNS ȪÿɕPpVNS1ɠPpVNS8ɖāé# ;, ºÿƺǅťǺř
:*, =:*RPşƅƺǅµ$ƈƲ#ŀŁ%ƥ=# <ɕXu et al. 2014; äɚɖƗƍǺř)ǂŚ, PpVNS Ȫÿ*Ǩ)ǚ8ǥ)Ħɋì'&, țűƾǜ$<jEdEdāé<ȫº$Ɨƍ# <%ŋ: %'!), Ǩ)ǚ$ěƗƍ< PpVNS1,PpVNS6, PpVNS7 (ȸ<ȱöƖ¡ ppvns1 ppvns6 ppvns7 @£ǳ%>, )öƖ¡$*țűƾǜ$<jEdEd)Ĝį³('<%, 3)6(jEdEd@țű)ȕȚȌ:''<%º !ɕäɜaɖ
:(, ǥ$Ɨƍ< PpVNS4 )öƖ¡$*, ǥ)jEdEdŧú, ǥµȫ)țűŦǝ12ă³(ú?=# ɕäɜbɖ=:)öƖ¡@ƄŗɕƜĊŽę 75%ɖ(ǒ%, ȲƑë(Ű0#ŉ ůȿ$Ǧ=#3 ɕäɜɖ=:)%:, PpVNS RP)țűƾǜĜį(ħɊ)Ȫÿ$<%, :(ǻ +jEdEdmy`
KhSP)¡µțű(#ȱǵ$<%, ½6#ĆɒƘ(ŋ: ('!ɕXu et al. 2014ɖ )ƠǶ*, R
Pşƅ%ǇƸŖşƅ)ŎȘ´țƦ²şƅ)ůȿ$, ň(
VNS țűƾǜºÈ)¾Ĥãÿ%
#)Ŧǝ@ŏ#%@ƥß# <ǡÜŹ %(, PpVNS Ȫÿ@my`KhS
Pµ$ȣÁƗƍ<%, ³ȑƘ'pOxƾǜūȀċ=<ɕXu et al. 2014ɖ
"3;, RPşƅ VNS Ȫÿ)ƴ
ǕƘ'ºÿŦǝ*pOxƾǜūȀċǝ$<%ǺȰ<%$9 ƍé)Ⱦşƅ7"țűƾǜ*îœƘ(*³#ūƾǜ$<%%Ö?#ǖ #7, VNS nBwȪÿȟÈ)Ƴǌ*, ɓ țűŦǝ)6(pOxƾǜū@ƾǜºÈpOx(ǀ4ȗ5ƀ(!)7=' 3, :(ǡÜŹ ƀ%#, ș) ppvns öƖ¡$*, Ǩ)ǚ(āé<WaEdƾǜɕƾǜôǛË, âĄĿĸ(Ŧǝ<%ǖ :=# <RP)Ŀĸƾǜɖ)ƾǜôǛ
ËŻč<ɕppvns1 ppvns6 ppvns7ɟäɜaɖ, < *, ǜÿ¡n_cɋì(£:=<ȒȚƾǜɕƾǜôµ«(Ʊ¹<%$ƾǜǯɇƬ@óø, ƾǜȷ)ƅȋȒȚ@ǭ %ǖ :=# <ƾǜɖ)ºÈƖĕ('<ɕppvns4ɖ%7ƥ=ɕXu et al. 2014ɖ
WaEdƾǜ7c Wn
Bƾǜ7ƾǜô)ø'©ɏȌ<ƾǜ$<% :, PpVNS (*pOxƾǜūȀċǝ
(Ä #ƾǜô©ɏ¾ĤǝÃ<%ǖ :=<my`KhSP)ñÖ(*jEdEd*ËôÈ' , RPşƅ$7ƪ(9!#*Ë ô@7!jEdEdƾǜ£:=<ɕLigrone etal. 2000ɖ)%@ǖ <%, ÌþëȾşƅ) ';ŉ ůȿ, č'%7RPşƅ%ǇƸŖş
M. Ohtani & T. Demura-6
BSJ-Review 7:92 (2016)

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M. Ohtani & T. Demura-7
BSJ-Review 7:93 (2016)

植物科学最前線 7:94 (2016)@Ȁċ<%º ! :$<)% :, RPşƅ)ůȿ$*Of zivÖį(Ǡ<³Ȇǁȏ*Ƌģ=# ' !, )ġ, VNS Ȫÿ)¾ĤÎĨ$<%śǀ*)33(ɕ< *ġ1&ƈƲ#Ƌģ=<Ĝ$ɖOf ƑÖįŦšȟÈ, U]şƅȟÈ)ůȿ$*ƾǜô)Of ÈțűƾǜºÈpOx)ņ'ƼȣƩ%#ǀ4ȗ3=
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M. Ohtani & T. Demura-8
BSJ-Review 7:94 (2016)

植物科学最前線 7:95 (2016)

ãÿǔ, NAC nBwȪÿ ! &)9 (ȟÈ#) , ) :&)9 (VNS Ȫÿƍ=#) , %áɌ7<Șė)ŨUP WĲǮ)ȟĎ*33'Ƒƅƪ)QixīðÏģ@Òǝ(# <,  ŰȔQixĂƘ'Cp^, VNS n
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ǎŜ%, #*ȾƑƅæ)ƗĎ("'!# !) :( țűƾǜ)ȟÈ, şƅ)tbDp )ȟÈ%&)9 (ȸȞ#) ǽ(Ĺ9 '33'ƢƮ)ȟ
Ď(9!#,  şƅțűƾǜȟÈ)Ȥ);)ȅ:(Ǻŋ =<%@ŐĠ @JBI
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