植物科学最前線 3: 114 ( 2012)

X_TR³ºª§ďƣ¸ãz~ŝǈċİÝæÝæǭřæƂůŸŴśŒŴæðĠ113-0033ģ¶ĭǣ 7-3-1
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
7-3-1, Hongo, Bunkyo-ku, Tokyo, 113-0033, JapanÓ¼mÈUW\e ŬĵśŒ+ƅƛ¨),ļ(FuHbsäÐȃ==)Îĩ+VyeIǉóÐ$<FuHbsłø(ĿƜ@ƎĚ;ȃ+ĿƜ@ƅƛ+¨ǢŚ×7ÛǢŚ×z°Ŏ)ČŽ$Úµ<;<5),ȃVyeIǉ½FuHbs7+ŪŨÑ/&łű)ǐ
Ǖ=<ĊƵ <·ƟƂFuHbsȁñƛȃMuPȃ]syQMuPbY]x{Iȃ ¼mÈ@t¡+

Ey^U{oȃŉƛȃƅƛƟȃetc.ȂǪ)$ȃ+łű(ǐǕ@ę!$ <+ƟǘOQ[o% <Ɵǘ,ȃǐǕñƛ+©

ƦȃǐǕñƛ+ĽŨƟ/+ƓŢ&ȃ=)ƍƮ¾+Q[YiƔ;ǒǊ<&)9;ē;Ż!
$ ;ȃ©Ʀ+Q[Yi%,Ʊƶƴ¾ȃƓŢzƮ¾+Q[Yi%, Rab GTPase & SNARE VyeIǉ&ȃǛµŨ)ä=ªãŇǍ<ȁÍ 1ȂŤ(<Ɵǘƈǌ%,Ť(<Ʊƶƴ¾ȃRabGTPaseȃSNARE VyeIǉ+SY],: $ <。2ȃ=:+ªã,Îĩ+FuHbs)
óÐ<&:ȃFuHbs+BD\y[C[C{@Ńé$ <ªã% <&6ƻ < "$¹ĵśŒ:ŬĵśŒǁśȃŘä+ŬĵśŒ/&Ǜµ<ǝŵ%ȃƅƛ¨+ƟĻǚ,+ƴǴ@Ø$FuHbs+ŶǺØ ȃ¿ħ)ǐǕƈǌ6Üļµ,% <ƟǘƈǌƏ+¥$%ȃƱƶƴ¾ȃRab GTPaseȃSNARE +}ŏSY]ĿƜ$ <&:ȃƟǘ+Üļµ),ȃ=:}ŏSY]@L{^$ <Ǡã+ǧƴ&ȃ=)ƍÚŤ+Ʃŷ)9<Ŀ

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植物科学最前線 3: 115 ( 2012)

Ɯ+ÜļµĊƵ% !&ŲÇ
=<"2;ȃ
Ʊƶƴ¾ȃRab GTPaseȃ
SNARE +Üļµ,ȃƟǘƈǌ+Üļ
µ&tyI$ <+% <ȁÍ 2Ȃ
(Dacks & Field 2007)

ÔRAB5 Qilb@¡»@ y Rab GTPase ,ȃǐǕñƛ@ĽŨƟ)ƓŢ<Q[Yi@Ǆſ<ªãQDYX% <++ Rab5 Ju{i,ȃ|Ǣ+Û@ǮŬĵśŒ)ûä
=$ ;ȃEy^U{o)$ <³Œ)$ĿƜ ¼mÈv@ËoÌÀ<¼mÈµÅ@ÀÎy+Dacks & Field 2007 CE$,ȃRab5 ļ(Ey^U{oĿƜ+Ǆſ@Ư!$<&ŭ:=$ < (Grosshans et al.
2006)OwDa_R_),ȃRHA1
ARA7ARA6 &3 "+ Rab5 lqw

J <RHA1 & ARA7 ,ȃ³Œ+Rab5 &9Ļǚ@Ě"ȃŬĵśŒ)ä=VDi+ RAB5ȁäÒRab5Ȃ% <|Ĥ ARA6 ,ȃäÒRab5 &,ǖ)BncĬż)Ɲǉǻ@»<('ȃǵø)œĉŨ(Ļǚ@Ě!$ <。+VDi+ RAB5 ,ǰ~ķŒ)ûä=$ <ȃ³Œ7Ƨ
Ǻ@Á4+ŬĵśŒ+ƂƊ),äÐ
( &:ȃARA6 Ju{i,ķŒǛµ+ǝŵ%ŕƠ)ŗĆķŒœ !  @³ºr+  >CEgci2G8K
k^Yle>3:-F+¢ !  @+ >CEgci2G8£º¼<A­=F =¼z¬
Ð"# Ñ>3:-F+¢  AÒ½Æ@v
4F¼mÈµÅ@´uÊ¦>¾±4F<¹.DG:-F+
Bars = 20 µm

ŤŨ( RAB5 % <&Ɨ :=<ȁmstB¹Ƭ@Á4BgLyivIN+|Ǣ),ȃARA6 & <ŵü+œĉ@§ĩ< RAB5 äÐ<ȃ+Şį,Ħ% <Ȃ$ȃ+ ARA6 ,ȃ|'+9 (Ɵǘƈǌ@¯ć$ <+> OwDa_R_)$+ƅƛ¨óÐ@ƹî$3<&ȃäÒ RAB5 6 ARA6 6ȃÜƛµEy^U{oȁmultivesicular endosome: MVEȂ)óÐ
$ ȃƘ+óÐ,è¥),|ơȃ|Ǣǧƴ""Ť(<Ey^U{oǳË)óÐ$ <&Ų=ȁ+&:ȃEy^U{o)6ļ(ŶǺ <&ª<Ȃ(Ueda et al.
2001)+óÐ+ǟ :ȃäÒ RAB5 & ARA6 ȃŤ(<ĿƜ@ĩ$ <&ŲÇ=$$ǑùȃäÒ RAB5 ,Ey^U{o:ŉƛƟ/+ȃARA6 ,Ey^U{o:ƅƛƟ/+ǐǕƂǌ%,: $ <&ȃ" )Ħ:&(! (Ebine et al. 2011)%,ȃ+ ARA6 óÐ$ <Ey^U{o)9!$ȃƅƛƟ2,ƅƛÛ/'+9 (Œǉǜ-=$ <+% >A. Era & T. Ueda-2BSJ- Revi ew 3: 115 ( 2012)

植物科学最前線 3: 116 ( 2012)=,ī ARA6  (<ǿŀĿƜ+ŧŘ)ǫ?<+&Èǹ&&6)īƺĦ% <ȃ

ARA6 ƈǌò(&6 "+Q]vQ)ï<ƙĎ)ǫ?<&ŭƷĆ:="" < $ȃ±Ǔ+ǘ;ȃ+ ARA6 VDi+ RAB5 ,ȃǰ~ķŒ)ûä=$ <ǰ~ķŒ+ÔǢ)ƕ<T`MK6ȃ+VDi+ RAB5 @Ě!$ <ȁMpARA6Ȃ%,ȃMpARA6 6OwD
a_R_+ ARA6 &¿ļEy^U{o:ƅƛƟ/+ǐǕ@¯ć$ <+% > MpARA6+ƅƛ¨óÐ@Ǆ0&>ȃ7,;Ey^U{o)óÐǀ5:=ȁÍ 3aȂ
%,ȃMpARA6 ,+Ey^U{o&'+FuHbs+Ǫ%+ǐǕ@¯ć$ <+% > ȅŘÐ2%)Ć:=$<ƉĲ,ȃMpARA6 ŉƛƟ&6ƅƛƟ&6Ť(<àá(Ɵ¶ŠȁerasomeȂ/+ǐǕ)ǫ?!$<&@ŲÇ$ <ȁÍ 3bȂ+ƉĲ,ȃT`MK)$ķŒœŤŨ RAB5 OwDa_R_&,Ť(<ĿƜ@ĩ$ <&@Ų$ <T`MK,Ǜµ+ǝŵ%ŕƠ+Ɵǘƈǌ@ŧǞȃš+ĥǯ%ƑĴ<ķŒ&(!+6=( Õ°§09 SNAREÒVAMP727 @@Ä¤AÙ Rab GTPase +,:)9;ǐǕñƛĽŨƟ)ƓŢ=ąȃǐǕñƛ)óÐ< R-SNARE &ĽŨƟ)óÐ< Q-SNARE &ƴ¾@ĀēȃƟƮ¾Ǌ<+ SNARE +)6ȃķŒÎ
ĩ+6+ <VAMP7 , R-SNARE +|Ŷ%ȃķŒ+ VAMP7 , VAMP71 & VAMP72 + 2 Ju{i:ē;Ż!$ <:)ŶãķŒ),ȃVAMP72 +NhJu{i% < VAMP727 äÐ$<VAMP727 ),BncĬż+ longin ^pDy&Ä-=<ǸÓ)œĉŨ(ǦĎĜ¤ǥ«@Ě!$;ȃǺ+ R-SNARE ,ƳãķŒǬ)Ř=ķŒ)ǿü)ä=$ <(:ȃVAMP727 ļ+ªã+äÐ,ȃ
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
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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¸¶@¥EÉx+Bar = 2 µmdMUk # 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

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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%ƅƛ !. )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 MitoTracker )9<n]
Ly^tB+ĳƤ@Ư!BIXyƒƎ)Ņ!ǥÀ@Ųn]Ly^tBÜĢƹî=Bar = 20 µm

Ø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-6BSJ- 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£VyeIǉ)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+BIXyƒƎ+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 ŭFuHbs&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]&(<> A. Era & T. Ueda-7BSJ- Revi ew 3: 120 ( 2012)

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«¨
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