植物科学最前線

6: 63 ( 2015)-5#+42 67/.ĉđķÚĒËśÎìƋŸŲûƈƑĲ °ŸWYcvƈƑǫǵƩÏƈƑõûǼõû ŸØƍûƈƑƍ ìƋŸŲûĊĻ ĭƇƅĐđēłõćŻùǆǬ 
A mechanism of reprogramming cell fate during early nodule development
Key words: endoreduplication, Lotus japonicus, nodule development, root nodule symbiosis
Takuya Suzaki, Masayoshi Kawaguchi
Division of Symbiotic Systems, National Institute for Basic Biology
Department of Basic Biology, School of Life Sciences, Graduate School for Advances Studies
Nishigonaka 38, Myodaiji, Okazaki 444-8585, Aichi, Japan:9  ô.twƍœŲ*ǫ.œŲȌhƍ.m}YshE,+ȍ/ȎƔƠãÿƲCņ?å
ñơƻ)?őƜƻ*Ƅ Ź?*-<>Ȏő-őƜ*×0@?°ŸÞþCĝİ?Ʋ
ÀCņ(? őƜC(ȎœŲ*őƜƻ/őƜ°Ÿ*×0@?ƔƠ*ūƠ.ŏȊ.;>*>Cìň*#Ƅº°ŸǸ¤Cƛ*)? œŲ*ĤŸŲ.°ŸřŖCƈƑ?)ȎőƜ°Ÿ/!.xdR Y*,?*/ô.ƈƑ-<>ƌ@(?*)?ȎœŲ.
ĝįĝİ.ƈƑ*(:ȂĔ-ƷÖťŵǖÓ6@(?*ı /Ƭ(? *.:ȎőƜ¸ŇžŸǧƎ)/ȎőƜƻ.ĮŎ<1őƜƻµŢ?~sN`K~Uƞ.Ə)?
Nod factor e~M *,>ȎĄœŲő.@6)µÃ(#ƀď.ơƱ.ǫƳµÃȎơƱµǃǐċ@ȎőƜÇìĝİ@?=)?(Brewin 1991, Yang et al. 1994) #&(Ȏ.žŸǧƎ)/Ȏóſ¼ũȌőƜƻĮŎȍ-Ĩƙ#ľ#,ÞþĝİȌőƜĝİȍ-Ò#µÃơƱ.ǦØǙĺȌ~qQ}uQȍǘ?*ǌĺ?*)?(Crespi & Frugier
2008, Suzaki & Kawaguchi 2014) őƜĝİ.ƈƑ/Ȏtwƍ.xdœŲ)?uzTQVȌLotus
japonicusȍ*_HtUzWȌMedicago truncatulaȍCŹ(ż)ƝÀſ-ƈƑǥ9=@(
? _HtUzW.őƜ/ŭǻèőƜ*×0@ȎőƜ.¬Ɩ-őƜw~YcvúçȎŠȅſ-őƜŸǴ?.-ĈȎuzTQV-ĝİ@?őƜ/łƊ,őƜw~YcvŖǣ/7=@ȎņǻèőƜ*×0@?(Brewin 1991, Ferguson et al. 2010) 6#ȎŭǻèőƜ)/Ȏ²§.
ƀďȌƸſ-twƍœŲ)/ƀď/ 3-4 ď=İ?ȍ*²ȃ.ơƱµǃǐċ@ȎőƜCŖ
İ?ơƱ*,?*Ƭ=@(?(Xiao et al. 2014) ĿȎņǻèőƜ)/Ċ=ó§.ƀďơƱ.ơƱµǃǐċ@ȎőƜCŖİ?ơƱ2*µÃ?*Ƭ=@(?(Szczyglowski et al.
1998) .<-ȎőƜ.ĝį;źŊ?ơƱ-ǩ/?ȎőƜƻ.ĮŎ£®řŖ;őƜ
žŸ.ìňſ,wLhZv/uzTQV*_HtUzW/¥ú@(?*ô.ƈƑ<>ƌ@(>Ȏ@6)Ȏ. 2 '.œŲ.ƈƑƭ.C½?*-<>őƜĝİ.ƈ
T. Suzaki & M. Kawaguchi-1
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6: 64 ( 2015)ƑžĎ(# ı ƈƑĈǖ*(?uzTQV/Ȏıäċ)Skvǭ·.ǋǒ;ƈƑ~] Y.Ľ
¨ǀB@(#twƍ.ƺň)?Ȍâ 1ȍ(Sato et al. 2008) SkvVGZ/ 470 Mb 4+)ȎGj*45ÐƘ)>Ȏƺ/ 30cm 4+)Ȏā².ƿ­Ū)üÿ#ŒëÍƲ)? 6#ȎŃǷŁJT_GqȌMG-20ȍCŹ@0Ȏ2 3 Ņ4+)C6B*)
? =-ȎtwƍœŲ)/Ȁ*@(#ĝǗǙĺĳǁŁ=Ɗƕ(?Ŭ:ųĥ)? ǝĖ)/²çĪ.e}Ys^(LORE1)CŹ#_Q}G.Ľ¨«Ā((>ȎǠǪûſ,ƈƑCǀŷï:Ľ''?(Fukai et al. 2012, Urbanski et al. 2012) uzTQ
V.°Ÿm eg )?uzTQVőƜƻȌMesorhizobium lotiȍ:ıäċ)SkvǋǒȎ
~] Y.Ľ¨ǀB@(?(Kaneko et al. 2000) °ŸǸ¤-?ŸŲ/ŭļ-?Ȏ°ŸǸ¤-?ŸŲ.ĿȌuzTQVȎőƜƻȍ.Skvǋǒ@Ȏ6#!@"@.ŸŲƏ-(ŗ ,Ǫſ~] Y;µøb Ľ&(?¡/4*D+, őƜ°Ÿ/ȎĄ-őƜƻ.ǏǕȎőƜƻ.ĄơƱ2.£®Ȏő
Ɯ²-?°ŸƔƠãÿ,+œŲőƜƻ!@"@.ßø-<&(ąĹ-Ƅ Ź,=Te @(>Ȏı -*&(/õòƷÖťŵǖŨǛ.ƈƑĈǖ)?Ȏ)/ȎőƜ¸ŇžŸ-ŮŬCĜ((ȎœŲ§.ƈƑ<>Ģ=@#ńǝ.Ƈǈ;ı .ƈƑİŌC

â 1ȏuzTQVȌAȍ*uzTQV.őƜ(B)ȏ

ƢƬĆ?

YR l : 10 cm (A), 1 mm (B)ȏ;9-5#+42   (% œŲ.ĝįĝİ)/Ȏ2 '.œŲrxȎVGeLGh*K NWơƱðŜµÃCTe ?ōſ,řƲCĶ*ĘƇ=@(? őƜžŸ-(:¡ó)/,ȎVGeLGh*K NW.őƜžŸ-?řƲCǔ3?ŸŶûſ,ƈƑÌ=ǀ
B@(#(Suzaki et al. 2013) ǝĖȎuzTQV*_HtUzW-(ȎVGeLGh.
Êăųÿ@ȎőƜžŸ-?VGeLGh.ǪſřƲ.ŶǋõǥD$(Gonzalez-Rizzo et al. 2006, Murray et al. 2007, Tirichine et al. 2007, Frugier et al. 2008) uzTQV.
LOTUS HISTIDINE KINASE 1 (LHK1)/WGigZg. ARABIDOPSIS HISTIDINE KINASE 4(AHK4) .K]Q)>ȎVGeLGh.Êă*(řƲ? LHK1 .řƲÝöòŽ (lhk1) )/őƜžŸǺĂ@?*=ȎLHK1 /őƜžŸCś-»ģ?©C:'*ƌ@#(Murray et al. 2007) !.Ŀ)Ȏlhk1 òŽ)/ȎőƜĝİƲý¯-/Ĵæ@ȎČļ.őƜĝİ@?*Ƈ=@(# ńǝȎuzTQV)/ LHK1A * LHK3
*Ñ=@#.VGeLGhÊă LHK1 *ǰǅ#řƲC:'*ł=-@#(Held et al. 2014) ĀǾ-Ȏlhk1 lhk1a lhk3 . 3 ǰòŽ)/őƜžŸ¯ǘ=, !.Ŀ)ȎLHK1 .ªĪòŽ*(ȎVGeLGh.Êă-ǸB?fwG.ŬƓůòŽ-<>Ȏ
VGeLGhWQg~QŖİſ-ţĪÃ@? spontaneous nodule formation 2 (snf2) òŽ
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6: 65 ( 2015)Æǿ@(? . snf2 òŽCőƜƻȂúç)Ư(?*ȎƀďơƱµǃƴžſ-ǐċ
@Ȏ“ƴžſőƜ”*×0@?ȎĝįǡĔ.őƜ*ȉ#Ŗǣĝİ@?*B&#(Tirichine et al. 2007) .<-ȎVGeLGhÊă.ÂĪ<1ªĪòŽ.ǋŋ=Ȏ
VGeLGhWQg~Q.ţĪÃőƜžŸ.e~M *(ħǇ'Åµ)?*B
&(# ńǝ)/ȎuzTQV.ő-VGeLGhC?$)ȎŉŚƗ)őƜŗŖ
ǣ.ĝİǐċ@?*:ƌ@(?(Heckmann et al. 2011) VGeLGh=.īíǨ/nY`X-EYm}O.~ǯÃ~ ƟŹ=@(>Ȏ.řŖ/ơƻ;ǮŞ.İµ»ģƟ*ȉ(?*Ƈ=@(?(Heyl & Schmulling 2003) VGeLGhīíǨ)/Ȏńƣſ-/ DNA ƥÏţĪC:' B-type response regulators (RRs)!._ SaeǪø.
žŵCǔƚ?*Ƈ=@(? _HtUzWCĦ-(ȎőƜžŸǧƎ-(žŵǐċ@? RRs ǪøÐÿ@''?(Gonzalez-Rizzo et al. 2006, Ariel et al. 2012)Ȏǎơ
,řƲǋł-/Ƶ&(, K NW:VGeLGh*Ðŗ-ȎőƜžŸ-?ǸÌ=ƈƑ@(#œŲrx. 1 ')?(Suzaki et al. 2013) ǝĖȎı .ƈƑ-<>ȎK NWĨƙƀďơƱµǃŃ-ǐċ@?*B&#(Suzaki et al. 2012) 6#Ȏsnf2 òŽ-?ƴžſőƜĝ
İǧƎ-(ȎK NWĨƙǐċ@?*:B&# =-Ȏ_HtUzW-(ȎK NW.ŕĪǜǟCǺĂ?*ȎőƜƻȂúç)őƜŗ.ŖǣµÃ?*;ȎV
GeLGhWQg~Q/ȎK NWĸ´Ny~E CT f? PIN Ǫø.žŵCTe ?*:ƌ@#(Plet et al. 2011, Rightmyer & Long 2011) @=.ƇǈCƦÏ?*ȎőƜžŸ-?VGeLGhWQg~Q. 1 '.ğ¿*(ȎK NWCőƜžŸÿȇê-ƾƐ?©CĶ&(?ÍƲĪƌÛ@? @6).*AȎ
K NW.ŸÏİȎ
WQg~Q ǜǟ-ǸB?Ǫø.őƜžŸ-?řƲCƌ#íÔ¡/,#9ȎőƜ¸ŇžŸ-?K NW.ğ¿/6,Ŭô ġȎuzTQVȎ_HtUzWÈ
Ŀ-(ǠǪûſ,ƈƑÁǢ?*)Ȏ!.ǎơł=-,?*ŇĠ@?

<9  0 -5#+42$)/.
 @6).uzTQV*_HtUzWCĦ*#ƈƑ-<>ȎőƜĝİ-ǸB?ô.Ǫ
øÐÿ@(? őƜĝİ)/Ȏ“ǂƀ-?őƜƻĮŎ-¢ú#WQg~Q”*“ƀď-?őƜžŸ”.Ǘſ-Ž,? 2 '.»ģƟȎƒǷſ-ǿ@#Ƥƫ-(ȎǤƧſ'ǫÐǔſ-ǥǀ?*Ƈ=@(>Ȏ. 2 '.»ģƟ.+%=Ŀ):Ɖƨ?*Ȏ
ìňſ-/őƜĝİ@, @6).ƈƑ)/Ȏ. 2 '.»ģƟC6Äµ(ƈƑ?*/àȀ)&#Ȏńǝ.ƤƫųŽſqx _ CŹ#ƈƑ-<>Ȏ!.ÜȈ/ƪÙ@''?(Rival et al. 2012, Hayashi et al. 2014) =-ȎƴžſőƜŵǖ.žǈ-<>Ȏ
őƜžŸ-ŮŬCĜ(#ƈƑÍƲ-,&# snf2 òŽ-<>ǐċ@?ƴžſőƜĝİȎ
Ə .ŀƇ.őƜĝİòŽ-<&(ĞȄCÊ?ǔ3#ƈƑ-<> GRAS _Gq.Ǚ³ßø
NODULATION SIGNALING PATHWAY 1 (NSP1) <1 NSP2ȎRWP-RK _Gq.Ǚ³ßø
NODULE INCEPTION (NIN) VGeLGhÊă.Ť-(ȎőƜžŸ.ś.»ģ-Ǹ?*ł=-,&#(Tirichine et al. 2007) NIN /őƜĝİ-ǸB?Ǫø*(ń¸T. Suzaki & M. Kawaguchi-3
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6: 66 ( 2015)

Ðÿ@#œŲ§.Ǫø)>ȎőƜĝİǧƎ)ųŽſ-žŵǐċ@?Ǫø)?*Ƈ=@(#(Schauser et al. 1999) ńǝȎNIN / NUCLEAR FACTOR-Y (NF-Y) ǪøCƃĹ.
Řſ*?*ł=-,>Ȏ=- NIN ; NF-Y ǪøCŖİſ-žŵ ?*ȎőƜƻȂúç)ƀďơƱµǃǐċ@?*:ƌ@#Ȍ#$Ȏsnf2 .ƴžſőƜ*Ž,>ȎőƜŗŖǣ.ĝİ6)-/Ƶ=,ȍ(Soyano et al. 2013) VGeLGhCĵ#ő)/ NIN .ž
ŵǐċ@?*Ȏ
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VGeLGhWQg~Q/ NIN .žŵǐċ-ħǇ)?*Ƭ=@(?(Heckmann et al.
2011, Soyano et al. 2014)=9,"!&-542$)8 ı /VGeLGhWQg~Q.Ť-(őƜ¸ŇžŸ-Ǹ?µøřŖCł=
-?*CƂſ-Ȏ@6) snf2 .ĴæòŽ.YP~ hQCǀȎÆǿ#òŽ.Ç
ßǪø.ǋŋCǀ&(# uzTQV. vagrant infection thread 1 (vag1) / snf2 ¢úſ,ƴžſőƜĝİCĴæ?òŽ*(Æǿ@#(Suzaki et al., 2014) vag1 .ý¯,řƲÝöòŽ*ĩB@?E~)/ȎǡĔ.őƜĝİý¯-Ĵæ@?*=ȎVAG1 /őƜžŸ-ħȆ,Ǫø*Ƭ=@? sXW|gP hQ-<>Ȏ!.ÇßǪøCųÿ#*AȎ
VAG1 /WGigZg-( DNA esG]w} \ VI .Ŗİßø*(Ƈ=@(?
ROOT HAIRLESS 1 (RHL1)/HYPOCOTYL 7 .K]Q)?*?*¹ł#(Sugimoto-Shirasu et al. 2005, Suzaki et al. 2014) DNA esG]w} \ VI /Ő²¦Á*×0@?
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2007) WGigZg. RHL1 C
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řƲ*(:ȉ(?

â 2. ǱŸè(A, C, D)* vag1 òŽ(B, E, F).őƜ¶űȏ*ƌÛ@#

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Ő²¦Á#ĮŎơƱ)é9'@(?(A,C)ȏvag1 òŽ)/Į''?ơƱ.Ő²¦ÁǘŎơƱ.ļ*VGZŦČ(B,E)ȎĮŎơƱ.¾ŝǽ.ơƱȌp}R

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T. Suzaki & M. Kawaguchi-4
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6: 67 ( 2015)ƔƠãÿCǀơƱȌĮŎơƱȍ2*µÃ?*Ƈ=@(?(Foucher & Kondorosi 2000,
Vinardell et al. 2003, Kondorosi & Kondorosi 2004,) vag1 òŽ.ĚE~)/őƜĝİƲý¯-/öB@(=ȎČļ.őƜCĝİ? ĝİ@#őƜ.VGZ/ǱŸè*õ,ǩ
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WĨƙ/VGeLGhWQg~Q.»ģ-@(?*Ƭ=@(? vag1 òŽ)/ƀďơƱµǃ/ǐċ@,:..K NWĨƙ/7=@?*=ȎK NWĨƙ6)/śĔ-řƲ(?*ĩB@? ǌĺ?*ȎƀďơƱ.Ő²¦Á/VGeLGh*K
NWWQg~Q.Ť-(ǐċ@?ÍƲĪƬ=@? .žǈ/ȎJfH.
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ł=-@(?(Yoon et al.
2014) DNA esG]w} \ VI
. A Vp{haeCT f?
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Ƥ7úç?ÍƲĪƬ=@?

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Ƹſ-Ƭ=@(# !.Ŀ)ȎňƈƑ-<>ǈ´@#őƜ¸ŇžŸ-?ƀďơƱ.Ő²¦Á/ơƱVGZ.łƊ,ðõCB, ŵç.*AȎőƜ¸ŇžŸ-(Ő²¦T. Suzaki & M. Kawaguchi-5
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