植物科学最前線 7:47 (2016)

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K. Ishizaki & K. Sakakibara1
BSJ-Review 7:47 (2016)

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K. Ishizaki & K. Sakakibara2
BSJ-Review 7:48 (2016)

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K. Ishizaki & K. Sakakibara3
BSJ-Review 7:49 (2016)

植物科学最前線 7:50 (2016)<3+: )űȖƝƔĄ*ñ$ȡÊ+ȋȆ,ɑ20 ǁĥÍ*=ÀĀȪĄ8ĈɌ
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K. Ishizaki & K. Sakakibara4
BSJ-Review 7:50 (2016)

植物科学最前線 7:51 (2016)h^cL'PNŧƊȭ­¤+ť+: )ƿƌ+ǈǡAġĴ=ȪĀÄĨh^cLɑ
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BSJ-Review 7:51 (2016)

植物科学最前線 7:52 (2016)

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植物科学最前線 7:55 (2016)㌴㍈⸴ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡢࢤࣀ࣒࠿ࡽぢࡓ᳜≀ࡢ㝣ୖ໬
ᇼᏕ୍ 1, ኴ⏣ၨஅ 1,2
1ᮾிᕤᴗ኱Ꮫ࣭኱Ꮫ㝔⏕࿨⌮ᕤᏛ◊✲⛉⌧ᅾࡢᡤᒓ㸸ᮾிᕤᴗ኱Ꮫ࣭⏕࿨⌮ᕤᏛ㝔ࠛ226-8501 ᶓ὾ᕷ⥳༊㛗ὠ⏣⏫ 4259 B-65
2ᮾிᕤᴗ኱Ꮫ࣭ᆅ⌫⏕࿨◊✲ᡤ
ࠛ152-8550 ᮾி㒔┠㯮༊኱ᒸᒣ 2-12-1-IE-1
Koichi Hori1 & Hiroyuki Ohta1,2
Klebsormidium flaccidum genome reveals genome evolution for plant terrestrial adaptation
Key words: charophyte, klebsormidium, genome analysis, land colonization
1

Tokyo Institute of Technology, Department of Biological Sciences.

Present address : Tokyo Institute of Technology, School of Life Science and Technology
2

Tokyo Institute of Technology, Earth-Life Science Institute.

ᆅ⌫࡟࠾ࡅࡿ⏕࿨ࡢṔྐ࡟࠾࠸࡚㸪⏕࿨ࡢ㝣ୖ㐍ฟࡣከᵝ࡞✀ࢆ⏕ࡳฟࡋ㸪⌧ᅾࡢ⏕≀ከᵝᛶࡢ♏࡜࡞ࡗ࡚࠸ࡿࠋ᭱ึࡢ⏕࿨ࡢ㝣ୖ㐍ฟࡢ㐣⛬ࡣ࠸ࡲࡔ୙᫂࡞Ⅼࡀከࡃ㸪᳜≀ࡼࡾࡣࡿ࠿ඛ࡟ࣂࢡࢸࣜ࢔ࡢ㝣ୖ㐍ฟࡀ࠶ࡗࡓ࡜⪃࠼ࡽࢀ࡚࠸ࡿ(Battistuzzi & Hedges 2009)ࠋࡋ࠿ࡋ
࡞ࡀࡽ㸪ື≀ࢆࡣࡌࡵ࡜ࡋࡓ」㞧࡞㝣ୖ⏕≀ࢆⓎ㐩ࡉࡏ㸪௒᪥ࡢᆅ⌫⎔ቃࢆᙧᡂࡍࡿ࡟⮳ࡗࡓ┤᥋ࡢཎືຊ࡜ࡋ࡚㸪ගྜᡂ࡟ࡼࡗ࡚஧㓟໬Ⅳ⣲ࢆᅛᐃࡋ᭷ᶵ≀ࢆྜᡂ࡛ࡁࡿ᳜≀ࡢ㝣ୖ㐍ฟࡣ኱ࡁ࡞ᙺ๭ࢆᯝࡓࡋࡓ஦ࡣ☜࠿࡛࠶ࡿࠋᮏ⥲ㄝ࡛ࡣ᳜≀ࡢ㝣ୖ໬ࢆゎ᫂ࡍࡿ࡟࠶ࡓࡗ࡚ 㔜 せ ࡞ ఩ ⨨ ࡙ ࡅ ࡟ ࠶ ࡿ ㌴ ㍈ ⸴ ᳜ ≀ 㛛 ࡢ ࠺ ࡕ 㸪 ࢡ ࣞ ࣈ ࢯ ࣝ ࣑ ࢹ ࢕ ࢘ ࣒ (Klebsormidium
flaccidum)ࡢࢤࣀ࣒ゎㄞ࡜௚⏕≀࡜ࡢẚ㍑ࡢ⤖ᯝࢆ⤂௓ࡍࡿࠋ㸯㸬᳜≀ࡢ㝣ୖ໬࡟ࡘ࠸࡚
᳜≀ࡢ㝣ୖ㐍ฟࡀ࠸ࡘ㉳ࡁࡓࡢ࠿ࡣᐃ࠿࡛ࡣ࡞ࡃ㸪ศᏊ⣔⤫ゎᯒ࠿ࡽ᭷⬇᳜≀㸦㝣ୖ᳜≀㸧ࡢฟ⌧ࡢ᥎ᐃᖺ௦ࡶㅖㄝ࠶ࡿࡢࡀ⌧≧࡛࠶ࡿࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ศᏊ⣔⤫ゎᯒ࡜⬊Ꮚࡢ໬▼ࡼࡾ⣙ 4 ൨ 7 ༓୓ᖺ๓࡟ࡣࡍ࡛࡟⌧⏕ࡢ㝣ୖ᳜≀ࡢඹ㏻♽ඛࡣㄌ⏕ࡋ࡚࠸ࡓ࡜⪃࠼ࡽࢀ࡚࠸ࡿ
(Rubinstein et al. 2010, Clarke et al. 2011, Magallón et al. 2013, Edwards & Kenrick 2015)ࠋ
㝣ୖࡣ㸪஝⇱ࡣࡶࡕࢁࢇᙉ࠸⣸እ⥺㸪኱ࡁ࡞ᗘኚ໬㸪㔜ຊ㸪ᰤ㣴ࡢḞஈ࡞࡝ᵝࠎ࡞ࢫࢺࣞࢫࡀᏑᅾࡍࡿ㐣㓞࡞⎔ቃ࡛࠶ࡾ㸪᳜≀ࡀ㝣ୖ㐍ฟࡍࡿ࡟࠶ࡓࡗ࡚㸪ࡇࢀࡽࡢࢫࢺࣞࢫ࡟㐺ᛂࡍࡿᚲせࡀ࠶ࡗࡓ࡜ᛮࢃࢀࡿࠋ᳜≀ࡢ㝣ୖ໬ࡣࡇࡢࡼ࠺࡞ᙜ᫬ࡢ㝣ୖ⎔ቃ࡟኱ࡁࡃᙳ㡪ࢆཷࡅࡿ୍᪉㸪㓟⣲⃰ᗘࡢቑ኱(Parnell & Foster 2012)㸪஧㓟໬Ⅳ⣲ᅛᐃ㸪㢼໬స⏝ࡸሁ✚(Scott &Glasspool 2006)࡞࡝ᆅ⌫⎔ቃࡢᙧᡂ࡟኱ࡁࡃᐤ୚ࡋ㸪┦஫࡟῝ࡃᙳ㡪ࢆཬࡰࡋ࠶ࡗࡓ࡜⪃࠼ࡽࢀࡿࠋ

K. Hori & H. Ohta-1

BSJ-Review 7:55 (2016)

植物科学最前線 7:56 (2016)㸰㸬㝣ୖ᳜≀ࡢ㉳※㝣ୖ᳜≀ࡣ⥳Ⰽ⸴㢮ࡢ୍⩌࠿ࡽศᒱࡋ㸪⌧ᅾࡢከᵝ࡞㝣ୖ᳜≀࡬࡜Ⓨᒎࡋ࡚ࡁࡓࡀ㸪࡝ࡢࡼ࠺࡞᳜≀ࡀ㝣ୖ࡟㐍ฟࡋ㸪࡝࠺ࡸࡗ࡚㝣ୖ⎔ቃ࡟㐺ᛂࡋⓎᒎࢆ㐙ࡆ࡚࠸ࡗࡓࡢࡔࢁ࠺࠿ࠋ
ࡑࡢゎ᫂࡟ࡣ㝣ୖ᳜≀࡟㏆࠸⸴㢮ࡢ≉ᛶࢆ᫂ࡽ࠿࡟ࡋ㸪௚ࡢ⸴㢮ࡸ㝣ୖ᳜≀࡜ẚ㍑ࡍࡿࡇ࡜ࡀ㔜せ࡞࢔ࣉ࣮ࣟࢳࡢ୍ࡘ࡜ࡋ࡚ᮇᚅࡉࢀࡿࠋ
ከᵝ࡞⸴㢮ࡀᏑᅾࡍࡿ୰࡛⣽⬊ศ⿣ࡢ≉ᚩࡸ⣔⤫ゎᯒ࠿ࡽ㌴㍈⸴᳜≀㛛(Charophyta)࡟ᒓ
ࡍࡿ⸴㢮ࡀ㝣ୖ᳜≀࡟᭱ࡶ㏆࠸࡜⪃࠼ࡽࢀ࡚࠸ࡿ(Lewis & McCourt 2004, Leliaert et al.,2012) ࠋ
㸦㌴㍈⸴᳜≀ࡣከ⣔⤫⩌࡛࠶ࡾ㸪ศ㢮ୖࡢ⾲グࡣࡲࡔ⤫୍ࡉࢀ࡚ࡣ࡞࠸ࡀ㸪ᮏ✏࡛ࡣ
Lewis & McCourt 2004 ࡢศ㢮࡟ᇶ࡙ࡁ㌴㍈⸴᳜≀ࡢศ㢮ࢆ⾲グࡋࡓࠋ㸧㌴㍈⸴᳜≀㛛ࡣࢡࣟࣟ࢟ࣈࢫ⸴⥘㸪ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘⸴⥘㸪ࢥࣞ࢜ࢣ࣮ࢸ⸴⥘㸪᥋ྜ⸴⥘㸪ࢩࣕࢪࢡ⸴⥘ࡢ 5ࡘࡢ⥘ࡀྵࡲࢀࡿ㸦ᅗ㸯㸧ࠋࡇࡢ࡞࠿࡛ᚋ⪅ࡢ 3 ࡘࡣ≉࡟㝣ୖ᳜≀࡟㏆࠸࡜ࡉࢀ㸪ࡑࡢ࠺ࡕ࡝
ࡢ⥘ࡀ㝣ୖ᳜≀ࡢጜጒ⩌࡛࠶ࡿ࠿ࡣ㛗ࡽࡃ㆟ㄽࡀ⥆࠸࡚ࡁࡓࠋ㏆ᖺ㸪㌿෗⏘≀᝟ሗࡢ⵳✚࡜࡜ࡶ࡟㸪ࡼࡾ⢭ᗘࡢ㧗࠸ゎᯒࡀ⾜ࢃࢀ㸪⌧ᅾࡣ᥋ྜ⸴⥘ࡀ㝣ୖ᳜≀ࡢጜጒ⩌࡜ࡍࡿㄝࡀ᭷ຊ
࡜࡞ࡗ࡚࠸ࡿ (Timme et al. 2012, Wickett et al. 2014) ࠋᐇ㝿㸪31 ✀㢮ࡢಖᏑࡉࢀࡓ㓄ิ࡟ᇶ࡙࠸ࡓᅗ㸯ࡢゎᯒ⤖ᯝࡶࡑࢀࢆᨭᣢࡋ࡚࠸ࡿࠋ
ᡃࠎࡣ㝣ୖ໬࡟࠸ࡓࡿ㐣⛬ࡢࡼࡾึᮇ࡟㸪࡝ࡢࡼ࠺࡞㑇ఏᏊࢆ⋓ᚓࡋࡓࡢ࠿࡜࠸࠺ほⅬ࡛᳜≀ࡢ㝣ୖ㐍ฟ࡟ࡘ࠸࡚◊✲ࢆ㐍ࡵࡿ஦ࢆ⪃࠼㸪ࡇࢀࡽࡢ㌴㍈⸴᳜≀ࡢ࠺ࡕẚ㍑ⓗึᮇ࡟ศᒱࡋ㸪ከ⣽⬊ᛶ࡛࠶ࡿࡀࢩࣥࣉࣝ࡞యไࢆᣢࡘࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡢࢤࣀ࣒ゎㄞࢆ㐍ࡵࡓࠋ
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21 ⏕≀✀࡟ඹ㏻ࡋ࡚ಖᏑࡉࢀࡓ᰾ࢥ࣮ࢻࡢࢱࣥࣃࢡ㉁㓄ิ㸦୍㒊 EST 㓄ิ࠿ࡽ᥎ᐃ㸧ࢆᇶ࡟సᡂࡋࡓ᭱ᑬ⣔⤫ᶞ (Hori et al. 2014. Fig. 2 ࢆᨵኚ)㸱㸬ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘࡜ࡣ
ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡣ⣒≧ᛶࡢከ⣽⬊ࡢ⸴㢮࡛࠶ࡾ㸪㐟㉮Ꮚ࡟ࡼࡿ↓ᛶ⏕Ṫࡣሗ࿌ࡉࢀ
࡚࠸ࡿࡀ㸪⣽⬊ศ໬ࡸ᭷ᛶ⏕Ṫࡣሗ࿌ࡉࢀ࡚࠸࡞࠸㸦ᅗ㸰a㸧ࠋ⏕ᜥ⎔ቃࡣῐỈ࠾ࡼࡧ㝣ୖࡢ‵₶࡞⎔ቃ࡛࠶ࡾ㸪ୡ⏺୰࡟ศᕸࡍࡿࠋ⸴㢮࡛ࡣ࠶ࡿࡀ㸪࠶ࡿ⛬ᗘ㝣ୖ⎔ቃ࡟㐺ᛂࡋࡓẼ⏕⸴㢮࡛࠶ࡾ㸪஝⇱(Morison & Sheath 1985, Elster et al. 2008, Karsten & Holzinger 2012)ࡸ෾⤖K. Hori & H. Ohta-2
BSJ-Review 7:56 (2016)

植物科学最前線 7:57 (2016)(Elster eet al. 2008, Nagao
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ࡀⰋࡃ࡞ࡿ࡜෌ࡧࡧቑṪࡍࡿ஦
஦ࡀ࡛ࡁࡿࠋ
ࠋ㠀ᖖ࡟㌟㌟㏆࡞⸴㢮࡛
࡛ࡶ࠶ࡾ㸪┤┤㐽ගࡢᙜࡓ
ࡓࡽ࡞࠸ࢥࣥࣥࢡ࣮ࣜࢺቨࡸ㊰㊰㠃࡞࡝࡛Ỉ
Ỉᢤࡁࣃ࢖ࣉࡢ⬥࡞࡝࡝‵Ẽࡢከ࠸ᡤᡤ࡟ࡋࡤࡋࡤ
ࡤ⩌㞟ࢆᙧᡂ
ᡂࡋ࡚࠸ࡿ 㸦ᅗ㸰b, c㸧
ࠋࠋ࡝ࡢࡼ࠺࡞
࡞ᶵᵓ࡟ࡼࡾ㸪㝣ୖ⎔⎔ቃ࡬㐺ᛂࡋ࡚࡚࠸ࡿࡢ࠿᫂
᫂ࡽ࠿࡛ࡣ࡞࠸ࡀ㸪㝣㝣ୖ⎔ቃ࡬ࡢ㐺㐺ᛂࢆ᫂ࡽ࠿
࠿࡟ࡋ࡚࠸ࡃ࠺࠼࡛㠀㠀ᖖ࡟⯆࿡῝࠸࠸ࠋࡓࡔࡋ㸪
㸪ࢡࣞࣈࢯࣝ
࣑ࣝࢹ࢕࢘ ࣒࡜㝣ୖ᳜≀≀ࡀศᒱࡋ࡚
࡚࠿ࡽ㸪ࢡࣞࣈࢯ࣑ࣝ ࢹ࢕࣒࢘ࡶ⊂⊂⮬ࡢ㐍໬ࢆ
ࢆ㐙ࡆ࡚࠾ࡾ㸪ࢡࣞࣈࣈࢯ࣑ࣝࢹ࢕࣒࢘࢘࡜㝣ୖ᳜
᳜≀࡟ඹ㏻ࡋ࡚࠸ࡿᙧ
ᙧ㉁ࡀ㸪
┿࡟ඹ㏻
㏻ὴ⏕ᙧ㉁࡛
࡛࠶ࡿࡢ࠿ὀ
ὀពࢆᡶ࠺ ᚲせࡀ࠶ࡿࠋᅗ㸰ࢡࣞࣈࢯ
ᅗ࣑ࣝࢹ࢕࣒࢘
࣒ࡢ㢧ᚤ㙾෗
෗┿a. K.
K flaccidum NIES-2285
Nᰴ 㸦ᅛయᇵᆅ࡟࡚࡚⏕⫱㸧b. ࢡࣞࣈࢯ࣑ࣝ
࣑ࢹ࢕࣒࢘࡜ᛮ
ᛮࢃࢀࡿ⸴㢮
㢮ࡀ⏕⫱ࡋ࡚࠸ࡓࢥࣥࢡ
ࢡ࣮ࣜࢺ㊰㠃
c. (b)࠿ࡽ᥇ྲྀࡋ
ࡋࡓ⸴㢮ࡢ㢧ᚤ
ᚤ㙾෗┿

㸲㸬㌴㍈
㍈⸴᳜≀㛛
㛛 Klebsorm
midium flacccidum㸦ࢡࣞࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘㸧
㸧ࡢࢤࣀ࣒
࣒ゎㄞ
ᡃࠎࡣ
ࡣᅜ❧⎔ቃ◊◊✲ᡤᚤ⏕≀≀⣔⤫ಖᏑᏑ᪋タࡼࡾศ
ศㆡࢆཷࡅࡿ
ࡿࡇ࡜ࡀ࡛ࡁ
ࡁࡿ Klebsorrmidium
flaccidumm NIES-22885 ᰴ㸦௨ୗᮏ✏࡛ࡣ K
K. flaccidum NIES-2285 ᰴࢆࢡࣞࣈ
ࣈࢯ࣑ࣝࢹ࢕
࢕࣒࢘࡜
⾲グࡋࡓ
ࡓࠋ
㸧ࢆ⏝࠸
࠸࡚ࢻࣛࣇࢺ
ࢺࢤࣀ࣒㓄ิ
ิࡢゎㄞࢆ⾜⾜ࡗࡓࠋࢤࣀ
ࣀ࣒ゎㄞ࡟ࡣ
ࡣ 454 GS FLX Tit
anium ࡜ Illumina GAIIx
Gࡢ㉸୪ิࢩ࣮ࢣ ࣥࢧ࣮ࢆ⏝⏝࠸࡚⣙ 6.1 Gb ࡢࢤࣀ ࣒㓄ิ࡜㸪⣙ 578
Mb ࡢ㌿㌿෗⏘≀㓄ิ
ิࢆྲྀᚓࡋࡓ
ࡓࠋࡇࢀࡽࡢ
ࡢ㓄ิࢆ࢔ࢭ
ࢭࣥࣈࣝࡋࡓ
ࡓ⤖ᯝ⣙ 1004 Mbp㸦18812 scaff
olds, ࣆࣆ࣮ࢡ࢝ࣂ࣮
࣮⋡:40 ಸ㸧ࡢࡢ᰾ࢤࣀ࣒㓄
㓄ิ࡜ⴥ⥳⥳యࢤࣀ࣒㸦⣙ 181 kbp㸧
㸧
㸪࣑ࢺࢥࣥ
ࣥࢻࣜ࢔
ࢤࣀ࣒㸦⣙ 106 kbpp㸧
㸪㌿෗⏘≀㸦17,422 ᗙ఩, ⣙ 21 Mb㸧ࢆ෌෌ᵓ⠏ࡍࡿ஦
஦ࡀ࡛ࡁࡓࠋࢡࣞࣈ
ࢯ࣑ࣝࢹࢹ࢕࣒࢘ࡢ᰾
᰾ࢤࣀ࣒ࢧ࢖
࢖ࢬࡣ᰾ࡢ ⺯ගᰁⰍീീ࠿ࡽ⣙ 117 Mbp ⛬ᗘ࡜࡜᥎ᐃࡉࢀ㸪
㸪ࣇ࢛ࢫ࣑ࢻࣛ࢖࢖ࣈ࣮ࣛࣜࡢ
ࡢ➃ㄞࡳࡢ⤖ᯝ 20%ᙉᙉ⛬ᗘࡢ཯᚟
᚟㓄ิ㡿ᇦࡀ
ࡀ࠶ࡿ࡜᥎ᐃ
ᐃࡉࢀࡓࡇ࡜࠿ࡽ㸪㓄ิỴᐃᐃࡀᅔ㞴࡞཯
཯᚟㓄ิ㡿ᇦ
ᇦ࡞࡝ࢆ㝖㝖࠸࡚ࢤࣀ࣒
࣒ࡢ࡯ࡰ඲ᇦ
ᇦࡢゎㄞࡀ᏶
᏶஢ࡋࡓࠋࡘࡘ࠸࡛㌿෗⏘≀᝟᝟ሗࡸ㓄ิゎ
ゎᯒ࡞࡝࠿ࡽࢱࣥࣃࢡ ㉁ࢆࢥ࣮ࢻ
ࢻࡍࡿ᰾ࢤࣀ࣒ࡢ 16,0633 㑇ఏᏊ㸪ⴥ⥳యࡢⴥ
117 㑇ఏ
ఏᏊ㸪࣑ࢺࢥ
ࢥࣥࢻࣜ࢔ࡢ
ࡢ 35 㑇ఏᏊᏊࢆண࡞࡝࠿ࡽࡽ㸪ࡇࢀࡽ 16,215
1
㑇ఏᏊࡢᶵ⬟ணணࡋ㸪᪤▱ࡢᶵ⬟⬟ࡀ᫂ࡽ࠿࡞
࡞㑇ఏᏊ࡜ࡢࡢ㢮ఝᛶࢆ⾜ࡗࡓࡓ(Hori et al. 2014, httpp://www.plan
ntmorphogenesis.bbio.titech.ac..jp/~algae_geenome_projeect/klebsorm
midium/)ࠋ

㸳㸬ࢡࣞࣈࢯ࣑ࣝ
࣑ࢹ࢕࣒࢘࡜௚⏕≀✀✀ࡢ㑇ఏᏊẚ㍑ゎᯒࢤࣀ࣒࣒ゎᯒࡢḟࡢ
ࡢẁ㝵࡜ࡋ࡚㸪ࢡࣞࣈ ࢯ࣑ࣝࢹ࢕࣒࢘ࡢ㑇ఏ
ఏᏊࢆ 5 ✀ࡢࡢ㝣ୖ᳜≀࠾
࠾ࡼࡧ 9✀ࡢ⸴㢮㢮࡜ẚ㍑ࡋࡓ
ࡓࠋࡇࢀࡽࡢ⏕≀ࡢ඲඲ࢱࣥࣃࢡ㉁㉁㓄ิࢆࢡࣛ
ࣛࢫࢱࣜࣥࢢ
ࢢࡋ㸪ẚ㍑⏕⏕≀✀ෆ࡟࠾࠸࡚࡚㸪⸴㢮ࡢࡳ
ࡳ࡟Ꮡᅾࡍࡿࢱࣥࣃࢡ ㉁࠿㝣ୖ᳜᳜≀ࡢࡳ࡟Ꮡ
Ꮡᅾࡍࡿࢱࣥ
ࣥࣃࢡ㉁࠿ศศ㢮ࢆ⾜ࡗࡓ⤖ᯝᯝ㸪ࢡࣞࣈࢯ
ࢯ࣑ࣝࢹ࢕࣒࢘ࡢ 1,2338 ࢱࣥࣃࢡ㉁㸦⣙ 8 %㸧ࡣ㝣ୖ᳜%᳜≀࡟≉␗ⓗⓗ࡛࠶ࡿ஦ࡀ᫂ࡽࡽ࠿࡜࡞ࡗࡓ
ࡓ㸦ᅗ㸱㸧ࠋࡲࡓྛ⏕≀≀✀ෆࡢ㢮ఝ
ఝ㑇ఏᏊࢆ㑇
㑇ఏᏊࣇ࢓࣑
࣑࣮ࣜ࡜ࡋ࡚࡚ࡲ࡜ࡵࡿ࡜㸪㝣㝣ୖ᳜≀ࡣ⸴
⸴㢮ࡼࡾከࡃࡢ㑇ఏᏊᏊࢆಖᣢࡋ࡚
࡚࠸ࡿࡀ㸪୺
୺࡟㑇ఏᏊ㔜
㔜」࡟ࡼࡾ㑇
㑇ఏᏊᩘࡀቑຍࡋࡋ࡚࠸ࡿࡇ࡜ࡀ㸪⥲㑇ఏఏᏊᩘ࡜㑇㑇ఏᏊࣇ࢓࣑࣮ࣜᩘࢆࣉ
ࣉࣟࢵࢺࡍࡿ
ࡿ஦࡛ࡣࡗࡁࡁࡾ࡜ぢ
K. H
Hori & H. Oh
hta-3

BSJ-Review 7:57 (2016)

植物科学最前線 7:58 (2016)ᅗ㸱㸱 15 ⏕≀✀✀㛫࡛ࡢ㑇ఏᏊẚ㍑
15 ⏕⏕≀✀ࡢ඲ࢱࣥ
ࣥࣃࢡ㉁㓄ิࢆ
ࢆࢡࣛࢫࢱࣜࣥ
ࣥࢢࡋ㸪ࢡࣛࢫ
ࢫࢱ࣮ෆࡢ௚⏕⏕≀✀ࡢࢱࣥࣃ
ࣃࢡ㉁ࡀ,⸴㢮㢮࡜㝣ୖ᳜≀ࡢࡢ࡝ࡕࡽ࡟≉᭷᭷࡛࠶ࡿ࠿ࢆᇶ
ᇶ࡟ࡋ࡚ศ㢮ࢆࢆ⾜ࡗࡓࠋ (Hoori et al. 2014. FFig. 3a ࢆᨵኚ)

࡚ྲྀࢀࡿ
ࡿ㸦ᅗ㸲㸧
ࠋࢡࣞࣈࢯࣝ
ࢡ࣑ࢹ࢕࣒࢘ ࡣ௚ࡢ⸴㢮㢮࡜㑇ఏᏊᩘ
ᩘ࠶ࡓࡾࡢ㑇
㑇ఏᏊࣇ࢓࣑
࣑࣮ࣜᩘࡣ࠶ࡲࡾࡾኚࢃࡽ࡞࠿
࠿ࡗࡓࡇ࡜࠿ࡽ㸪㑇ఏఏᏊ㔜」ࡣ᳜
᳜≀ࡀ⤌⧊ศ
ศ໬ࢆ⋓ᚓࡋ
ࡋ㸪㝣ୖࡢ」」㞧࡞⎔ቃ࡟ᛂ⟅⟅ࡋ࡚࠸ࡃ㐣
㐣⛬࡛᭷ຠ࡟ാ࠸ࡓࡢ ࡛ࡣ࡞࠸࠿
࠿࡜⪃࠼ࡽࢀ
ࢀࡿࠋࡋ࠿ࡋ
ࡋ࡞ࡀࡽᇶ㒊
㒊㝣ୖ᳜≀࡛࠶ࡿࡿࢮࢽࢦࢣࡣ
ࡣ㑇ఏᏊࡢ㔜
㔜」ࡀᑡ࡞ ࠸࡜ࡢሗ࿌
࿌ࡶ࠶ࡾ㸦኱
኱࿴࣭Ἑෆ, 22012㸧
㸪㑇ఏ
ఏᏊ㔜」
ࡢ᥎⛣࡜
࡜᳜≀ࡢ㝣ୖ
ୖ໬ࡢ㛵୚ࢆ᫂ࡽ࠿࡟ ࡍࡿࡓࡵ࡟
࡟ࡣࡼࡾከࡃࡢ㌴㍈⸴᳜᳜≀࡜ᇶ㒊㝣
㝣ୖ᳜≀ࡢẚ㍑ゎ
ゎᯒࡀ࣮࣏࢟
࣏࢖ࣥࢺ࡜࡞ࡗ࡚࠸ࡃ ࡛࠶ࢁ࠺ࠋ

ᅗ 4 15 ⏕≀✀ࡢ⥲⥲㑇ఏᏊᩘ࡜
ᩘࡢࣉࣟࢵࢺ
㑇ఏᏊࣇ࢓࣑࣮ࣜᩘ
ྛ⏕≀✀ෆࡢ㢮ఝ㑇㑇ఏᏊࢆ㑇ఏᏊ
ᏊᏊࣇ࢓࣑࣮ࣜ࡜ࡋ࡚ࡲ ࡜ࡵ㸪⥲㑇ఏᏊ࡜㑇ఏᏊࣇ࢓࣑࣮ࣜ
࣮ᩘ࡛ࣉࣟࢵ ࢺ⸴㢮ࡣᅗ㸱࡜ྠ
ྠࡌ✀ࢆ౑⏝ࡋࡋࡓࠋ⸴࡚࠸ࡿࠋࠋⅬ⥺ࡣ⸴㢮ࡢࡢࣉࣟࢵࢺࡢ㏆
㏆
ఝ᭤⥺࡛
࡛࠶ࡿࠋ

ࡲࡓᡃ
ᡃࠎࡣ㑇ఏᏊ
Ꮚࣇ࢓࣑࣮ࣜ
࣮ࡢ࡯࠿㸪 ࢱࣥࣃࢡ㉁㉁ࢆᵓᡂࡍࡿ
ࡿࣃ࣮ࢶ࡜⪃⪃࠼ࡽࢀࡿࢱࢱࣥࣃࢡ㉁ࢻ࣓࢖
࢖ࣥࡢᵓᡂ࡟
࡟ࡘ࠸࡚ࡶẚẚ㍑ゎᯒࢆ ⾜ࡗࡓࠋྛྛ⏕≀✀ࡢࢻ
ࢻ࣓࢖ࣥࡢ✀✀㢮ᩘࡸ㸪ࡑ
ࡑࡢ⤌ࡳྜࢃࡏࣃࣃࢱ࣮ࣥ࡜⥲⥲㑇ఏᏊᩘࢆࣉࣟࢵࢺ ࡋࡓ⤖ᯝ㸪ࢻ࣓࢖ࣥࡢ
ࡢ✀㢮ᩘࡣ㝣
㝣ୖ᳜≀࡛ࡍ
ࡍ࡛࡟㢌ᡴࡕ࡟࡞࡞ࡗ࡚࠸ࡿࡀ
ࡀ㸪ࡑࡢ⤌ࡳ
ࡳྜࢃࡏࣃ ࢱ࣮ࣥࡣ⿕⿕Ꮚ᳜≀࡛ࡉ
ࡉࡽ࡟ቑຍࡋ
ࡋ࡚࠸ࡿࡇ࡜࡜ࡀ᫂ࡽ࠿࡜࡞ࡗࡗࡓ㸦ᅗ㸳㸧ࠋࡲࡓゎᯒ
ᯒࡋࡓ 5 ✀ࡢࡢ㝣ୖ᳜≀࡟
࡟ඹ㏻ࡍࡿࢻ
ࢻ࣓࢖ࣥࡸ㸪ࡑࡢ⤌ࡳྜ
ྜࢃࡏࣃ
ࢱ࣮ࣥࢆ
ࢆ⸴㢮ࡀ࡝ࢀ
ࢀࡔࡅ⋓ᚓࡋ࡚࠸ࡿ࠿
࠿ࢆㄪ࡭ࡓ⤖⤖ᯝ㸪ࢡࣞࣈ
ࣈࢯ࣑ࣝࢹ࢕
࢕࣒࡛࢘ࡣ㝣
㝣ୖ᳜≀
࡟ඹ㏻ࡍ
ࡍࡿ 90.7%ࡢ
ࡢࢻ࣓࢖ࣥ࡜ 84.3%ࡢ ࢻ࣓࢖ࣥࡢ
ࡢ⤌ࡳྜࢃࡏ
ࡏࣃࢱ࣮ࣥࢆ
ࢆ⋓ᚓࡋ࡚࠾࠾ࡾ㸪௚ࡢ⸴㢮ࡼࡼࡾ 2ࠥ3 ๭⛬ᗘ㧗࠸ࡇ๭ࡇ࡜ࡀศ࠿ࡗ
ࡗࡓ㸦ᅗ㸴㸧
ࠋࡇࢀࡽࡢࡢࡇ࡜ࡣࢡࣞ
ࣞࣈࢯ࣑ࣝࢹࢹ࢕࣒࢘ࡀ㝣ୖ᳜᳜≀࡟ඹ㏻ࡢ
ࡢࢱࣥࣃࢡ㉁㉁ࡢᶵ⬟ࢆ సࡿ࠺࠼࡛
࡛ከࡃࡢᇶᮏ
ᮏⓗ࡞ࣃ࣮ࢶ
ࢶࢆࡍ࡛࡟⋓⋓ᚓࡋ࡚
K. H
Hori & H. Oh
hta-4
BSJ-Review 7:58 (2016)

植物科学最前線 7:59 (2016)

଺৕೘੟

10000

३টॖॾॼ६ॼ

ॖॿএউছॻওॖথਯ उेलॻওॖথभੌा়ॎचਯ

8000ॡঞঈ९ঝ঑ॹॕक़঒

6000ॖॾढ़ॱ঄ং

঄ওॶজफ़ॿ०ॣ

৵୴థ

4000

॥ॣ೘੟

ᇖథ
2000

䝗䝯䜲䞁ᩘ 䝗䝯䜲䞁䛾⤌䜏ྜ䜟䛫ᩘ㝣ୖ᳜≀ᅗ㸳 15 ⏕≀✀ࡢ⥲㑇ఏᏊᩘ࡜ࢻ࣓࢖ࣥࡢ✀㢮ᩘ࠾ࡼࡧࢻ࣓࢖
ࣥࡢ⤌ࡳྜࢃࡏᩘࡢࣉࣟࢵࢺ
ྛ⏕≀✀ෆࡢ඲ࢱࣥࣃࢡ㉁ࡢ pfamA࠾ࡼࡧ pfamB ࢆ᳨⣴ࡋ㸪ࢻ࣓࢖ࣥࡢ
✀㢮ᩘ࠾ࡼࡧ⤌ࡳྜࢃࡏࣃࢱ࣮ࣥࢆ
࢝࢘ࣥࢺࡋࡓࠋ⸴㢮ࡣᅗ㸱࡜ྠࡌ✀ࢆ౑⏝ࡋ࡚࠸ࡿࠋከࡃࡢ⸴㢮࡟࠾࠸࡚㸪ࢻ࣓࢖ࣥࡢ✀㢮ᩘࡼࡾࡶࢻ࣓࢖ࣥࡢ
⤌ࡳྜࢃࡏࣃࢱ࣮ࣥࡀᑡ࡞ࡃ࡞ࡗ࡚
࠸ࡿࡢࡣ㸪ࢻ࣓࢖ࣥࡢ⤌ࡳྜࢃࡏࣃࢱ
࣮ࣥࡀᑡ࡞ࡃỴࡲࡗࡓ⤌ྜࡏࡀከ࠸
ࡓࡵ࡛࠶ࡿࠋ

⸴㢮

0
0

10000

20000

30000

40000

50000

੕఩஫৕ਯ

100.0

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80.0
吠
吸
叻 60.0呉⋓ᚓ 40.0叏๭
ྜ 20.0ᅗ㸴 㝣ୖ᳜≀࡟ඹ㏻ࡍࡿ
ࢻ࣓࢖ࣥ࠾ࡼࡧࢻ࣓࢖ࣥࡢ
⤌ࡳྜࢃࡏࡢ⋓ᚓ⋡ࣄ࣓ࢶࣜ࢞ࢿࢦࢣ㸪࢖ࢾ࢝ࢱࣄ
ࣂ㸪࢖ࢿ㸪࣏ࣉࣛ㸪ࢩࣟ࢖ࢾࢼ
ࢬࢼ࡟ඹ㏻ࡍࡿ pfam ࢻ࣓࢖ࣥ(4,894 ࢻ࣓࢖ࣥ)࠾ࡼࡧ㸪ࢻ࣓࢖ࣥࡢ⤌ࡳྜࢃࡏࣃࢱ࣮ࣥ(2,801
ࣃࢱ࣮ࣥ)ࡢ࠺ࡕࡑࢀࡒࢀࡢ⸴㢮࡛⋓ᚓࡋ࡚࠸ࡿ๭ྜࠋ

0.0࠸ࡿࡇ࡜ࢆព࿡ࡍࡿࠋࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡣ㌴㍈⸴᳜≀ࡢ୰࡛᪩࠸࠺ࡕ࡟ศᒱࡋ㸪㠀ᖖ࡟ࢩࣥࣉࣝ࡞యไ࡛࠶ࡿ࡟ࡶ࠿࠿ࢃࡽࡎ㸪㝣ୖ᳜≀≉᭷ࡢᵝࠎ࡞ࢩࢫࢸ࣒ࢆ㸪ཎጞⓗ࡞ᙧ࡛࠶ࡗࡓ࡜ࡋ࡚ࡶண᝿௨ୖ࡟⋓ᚓࡋ࡚࠸ࡿ࠿ࡶࡋࢀ࡞࠸ࠋ
௨ୖࡢ⤖ᯝࢆࡲ࡜ࡵࡿ࡜㸪᳜≀ࡀ㝣ୖ࡟㐺ᛂࡋ࡚࠸ࡃ㐣⛬࡛㑇ఏᏊࡢከᵝᛶࡢ⋓ᚓࡣ㸪ḟ
ࡢ 3 ẁ㝵ࡢࢫࢸࢵࣉ࡟ศࡅࡽࢀࡿ࡜⪃࠼ࡽࢀࡿ㸦ᅗ㸵㸧ࠋi) 㝣ୖ᳜≀ࡢඹ㏻♽ඛ࡛࠶ࡿ⥳⸴࠿ࡽࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡀศᒱࡍࡿࡲ࡛ࡢ㛫ࡣ㑇ఏᏊᩘࡢቑຍࡀ㑇ఏᏊࡢ✀㢮ࡢቑຍࢆࡶࡓࡽࡋࡓ࡜⪃࠼ࡽࢀࡿࠋ
ii) ࢥࢣ㸪ࢩࢲ᳜≀ࡢࡼ࠺࡟㝣ୖ⎔ቃ࡟ࡼࡾ㐺ᛂࡋ㸪⤌⧊ศ໬ࡀᙧᡂࡉࢀࡿ࡟ࡣ㸪ྠ㑇ఏᏊ᪘ࡢ୰࡛ࣂ࢚࣮ࣜࢩࣙࣥࢆቑຍࡉࡏ㸪⣽࠿࡞ᶵ⬟ㄪ⠇ࡸⓎ⌧ㄪ⠇ࢆྍ⬟࡟ࡋࡓ࡜⪃࠼ࡽࢀࡿࠋ
iii) ✀Ꮚ᳜≀ࡢࡼ࠺࡞㧗ᗘ࡞㝣ୖ⎔ቃ࡬ࡢ㐺ᛂ࡜⤌⧊ศ໬ࢆྍ⬟࡟ࡍࡿ࡟ࡣ᪤Ꮡࡢࣃ࣮ࢶࡢ
᪂ࡋ࠸⤌ࡳྜࢃࡏࢆ⏕ࡳฟࡋ㸪᪂ࡋ࠸ᶵ⬟ࡢ㑇ఏᏊࢆ⏕ࡳฟࡋࡓࡇ࡜ࡀ㔜せࡔࡗࡓ࡜⪃࠼ࡽࢀࡿࠋࡇࡢࡼ࠺࡞㐣⛬ࡢ୰࡛ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡢ♽ඛࡣ㸪㝣ୖ᳜≀ࡀከ⣽⬊యࡢᵓ⠏ࡸ㝣ୖ⎔ቃ࡟㐺ᛂࡍࡿࡓࡵ࡟Ⓨ㐩ࡉࡏ࡚࠸ࡗࡓ㑇ఏᏊ㸪࠶ࡿ࠸ࡣࡑࡢࣃ࣮ࢶࡢከࡃࢆࡍ࡛࡟⋓ᚓࡋ࡚࠸ࡓ஦ࡀ᥎ᐃࡉࢀࡓࠋ

K. Hori & H. Ohta-5

BSJ-Review 7:59 (2016)

植物科学最前線 7:60 (2016)

෤ᇖ

ॡঞঈ९
९ঝ঑ॹॕक़क़঒॥ॣ೘೘੟‫ؚ‬৵୴థ
థ

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ከ⣽
⣽⬊໬?
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खःॻওॖথ
থभੌाा়ॎचभඃ
ඃ੭஫৕ர৔भ఩஫
LL ংজग़‫ش‬३ঙথ
থभੜਸ

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L ಮ঱৲पঢ়
఩஫৕भர
ரథभੜਸ
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$
$
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ᅗ㸵 ᳜≀≀ࡢ㝣ୖ໬ࡢ 㐣⛬࡜㑇ఏᏊᏊከᵝᛶࡢ⋓⋓ᚓ㸴㸬ࢡࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘
࣒࡟࠾ࡅࡿ ᳜≀࣍ࣝࣔࣔࣥ㛵㐃ᅉᏊ
ࢡࣞࣈࣈࢯ࣑ࣝࢹ࢕
࢕࣒࢘ࡢ㑇ఏ
ఏᏊ࡜⸴㢮㢮㸪㝣ୖ᳜≀≀ࡢ㑇ఏᏊࢆ
ࢆẚ㍑ࡋࡓ⤖⤖ᯝ㸪᝟ሗఏ
ఏ㐩㸪⎔ቃᛂ⟅㸪⣽⬊ቨྜᡂᡂ㸪᳜≀࣍ࣝ
ࣝࣔࣥ㛵㐃㐃ᅉᏊ࡞࡝࡟
࡟㛵ࢃࡿᅉᏊ
Ꮚ࡟㸪㝣ୖ᳜
᳜≀࡛኱ᖜ࡟
࡟ቑຍࡋ࡚࠸ࡿࡶࡶࡢࡸ㸪㝣ୖ
ୖ᳜≀≉␗ⓗⓗ࡞ࡶࡢࡀ ከ࠸ഴྥࡀ
ࡀ᫂ࡽ࠿࡜࡞
࡞ࡗࡓ(Hori eet al. 2014)ࠋ
୰࡛ࡶࡶ᳜≀࣍ࣝࣔ
ࣔࣥ᝟ሗఏ㐩㐩ࡣ㝣ୖ᳜᳜≀ࡢ⎔ቃᛂ
ᛂ⟅࡟࠾࠸࡚࡚㔜せ࡞ᙺ๭
๭ࢆᢸࡗ࡚࠸
࠸ࡿࠋࡑࡇ࡛ࢡࣞࣞࣈࢯ࣑ࣝࢹ
ࢹ࢕࣒࢘࡟࠾
࠾࠸࡚᳜≀≀࣍ࣝࣔࣥࡢ
ࡢྜᡂ⣔㑇ఏ
ఏᏊࢆ᥈⣴ࡋ
ࡋࡓ⤖ᯝ㸪୺୺せ࡞᳜≀࣍ࣝࣔࣔࣥྜᡂ⣔ࡀ
ࡀᏑᅾࡋ࡚࠸
࠸ࡿࡇ࡜ࡀ ᥎ᐃࡉࢀࡓ
ࡓࠋࡲࡓࢡࣞ
ࣞࣈࢯ࣑ࣝࢹ
ࢹ࢕࣒࢘ࡢ᳜
᳜≀࣍ࣝ
ࣔࣥࡢ

ᐃࢆ⾜ࡗࡓ
ࡓ⤖ᯝ㸪࣮࢜
࣮࢟ࢩࣥ㸪 ࢔ࣈࢩࢪࣥ
ࣥ㓟㸪ࢧ࢖ࢺ
ࢺ࢝࢖ࢽࣥ㸪 ࢪࣕࢫࣔࣥ
ࣥ㓟㸪ࢧ

ࣜࢳࣝ㓟
㓟࡜ከᩘࡢ᳜
᳜≀࣍ࣝࣔࣥࡀࢡࣞࣈ
ࣈࢯ࣑ࣝࢹ࢕࣒࢘࡟Ꮡᅾ
ᅾࡍࡿࡇ࡜ࡀ
ࡀ᫂ࡽ࠿࡜࡞
࡞ࡗࡓࠋࡉࡽ࡟᳜᳜≀࣍ࣝࣔࣥ
ࣥࡢࢩࢢࢼࣝ
ࣝ᝟ሗఏ㐩㐩⣔ࡢ㛵㐃ᅉᅉᏊࡢヲ⣽࡞
࡞ゎᯒࢆ⾜ࡗ
ࡗࡓ⤖ᯝ㸪࠸
࠸ࡃࡘ࠿ࡢཷᐜయయ㸪㍺㏦⣔ࡸ
ࡸ᝟ሗఏ㐩⣔⣔ࡀᏑᅾࡋ ࡚࠾ࡾ㸪ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘࡟
࡟࠾࠸࡚ࡶఱఱࡽ࠿ࡢ᳜≀࣍ࣝࣝࣔࣥᛂ⟅ࡀ
ࡀᏑᅾࡋ࡚࠸
࠸ࡿࡇ࡜ࡀ ♧၀ࡉࢀࡓ
ࡓ㸦ᅗ㸶㸧
ࠋ
ࡑࡢ୍
୍᪉࡛ TIR11㸦࣮࢜࢟ࢩࢩࣥཷᐜయ㸧㸪
㸪COI1㸦ࢪࢪࣕࢫࣔࣥ㓟
㓟ཷᐜయ㸧࡞
࡞࡝ࡢࣘࣅ࢟
࢟ࢳࣥ౫Ꮡ ᛶࢱ ࣥ ࣃࢡ ㉁ศ
ศゎ ࢆ௓ ࡋ ࡓ᝟ ሗఏ 㐩㐩⤒㊰ ࡢከከ ࡃࡀᏑ ᅾࡋࡋ ࡚࠾ࡽ ࡎࡎ㸪 ABA ཷᐜཷ యࡢ
PYR/PYYL/RCAR ࡶᏑᅾࡋ࡚࠸
࠸࡞࠸ࡇ࡜ࡀ
ࡀ᫂ࡽ࠿࡜࡞
࡞ࡗࡓࠋࡇࢀ
ࢀࡽࡢཷᐜయ
యࡣ㸪⌧ᅾࡢࡢ᳜≀࣍ࣝࣔࣥ᝟᝟ሗఏ㐩࡟࠾
࠾࠸࡚୺せ࡞⤒㊰ࢆᢸᢸࡗ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚࡚࠾ࡾ㸪᳜≀≀࣍ࣝࣔࣥ㛫㛫ࡢࢡࣟࢫࢺ࣮ࢡࢡࢆ⏕ࡳฟࡍ
ࡍ⤒㊰࡜ࡋ࡚ࡶ▱ࡽࢀࢀ࡚࠸ࡿࠋࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡢ
ࡢ᳜≀࣍ࣝࣔࣔࣥఏ㐩⤒㊰ࡣ㝣㝣ୖ᳜≀ࡢ୺୺せ࡞᝟ሗఏ
ఏ㐩⤒㊰ࢆ ⋓ᚓࡍࡿ๓๓ẁ㝵࡟࠶ࡿ
ࡿ࡜⪃࠼ࡽࢀ
ࢀ㸪㝣ୖ᳜≀≀ࡢ୺せ࡞⤒㊰࡜࡜ࡣ␗࡞ࡗࡓ
ࡓཎጞⓗ࡞᳜
᳜≀࣍ࣝࣔ ࣥఏ㐩⤒㊰㊰࡛࠶ࡿྍ⬟⬟ᛶࡀ⪃࠼ࡽ
ࡽࢀࡿࠋḟࡢㄢ㢟ࡣ㸪ࢡࣞࣈࢯࢯ࣑ࣝࢹ࢕࢘
࣒࢘ࡢ᳜≀࣍ࣝࣔࣥࡢ స⏝ࡸఏ㐩㐩⤒㊰ࢆ᫂ࡽ
ࡽ࠿࡟ࡋ㸪᳜
᳜≀ࡀ㝣ୖ࡟
࡟㐍ฟࡍࡿ㐣⛬࡛࡛㸪᳜≀࣍ࣝ
ࣝࣔࣥࡢ㉳※※ࡸᙺ๭ࢆ ࡝ࡢࡼ࠺࡟
࡟Ⓨ㐩ࡉࡏ࡚
࡚ࡁࡓ࠿ゎ᫂
᫂ࡋ࡚࠸ࡃࡇ
ࡇ࡜࡛࠶ࡿࠋ
K. H
Hori & H. Oh
hta-6

BSJ-Review 7:60 (2016)

植物科学最前線 7:61 (2016)४ক५ঔথ෷
ABP1 ? ?

ROP

PIN

४ঋঞজথ

ग़ॳঞথ

॔ঈ३३থ෷

१জॳঝ෷

AUX1
ETR1

CRE1,AHK

GTG

CTR1

ड़‫ش‬य़३থ

PYR/PYL/RCAR

?

EIN2
TIR1

COI1

GID

NPR3/4
TypeA
ARR

EBF1
NPR1

SnRK2
IAAs

ARF

JAZ

MYC

DELLA

PIF

AHP

CUL3

PP2C
growthregulation१ॖॺढ़ॖॽথ

EIN3

ASK1

RBX
CUL1

SCF complex
phytochrome

TypeB
ARR

CRFs

AREB/ABF೘੟঍ঝঔথૢ௦఩஫৕भ৅ਠ

ᅗ㸶 ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘࡟᳨ฟࡉࢀࡓ᳜≀࣍ࣝࣔࣥ࡜㢮ఝ㑇ఏᏊࡀぢฟࡉࢀࡓ᝟ሗఏ㐩ᅉᏊ
᳜≀࣍ࣝࣔࣥࡢ࠺ࡕ᳨ฟࡉࢀࡓࡶࡢࡣỈⰍࡢ࣎ࢵࢡࢫ࡛♧ࡋࡓࠋ࢚ࢳࣞࣥࡣᮍ ᐃ࡛࠶ࡿࠋ㝣ୖ᳜≀࡛᫂ࡽ࠿࡜࡞ࡗ࡚࠸ࡿ᝟ሗఏ㐩ᅉᏊࡢ࠺ࡕ㸪ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࡛࢘㢮ఝ㑇ఏᏊࡀᏑᅾࡋ࡚࠸ࡓ
ࡶࡢࢆ⥳࡛♧ࡋࡓࠋⅬ⥺ࡣ㢮ఝ㑇ఏᏊࡀぢࡘ࠿ࡗ࡚࠸࡞࠸ࠋ

㸵㸬ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘࡟࠾ࡅࡿ㌿෗ᅉᏊ᳜≀࣍ࣝࣔࣥࡢ௚࡟㸪㌿෗ᅉᏊࡶ⎔ቃᛂ⟅࡟㛵ࢃࡿ㠀ᖖ࡟㔜せ࡞ᅉᏊ࡛࠶ࡿࠋPlant
Transcription Factor Database v3.0 (Jin et al. 2013)ࡢศ㢮ἲ࡟ᇶ࡙࠸࡚ 58 ✀㢮ࡢ㌿෗ᅉᏊࡢྠᐃ
ࢆ⾜ࡗࡓ⤖ᯝ㸪ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘࠿ࡽ 266 㑇ఏᏊࡢ㌿෗ᅉᏊࡀྠᐃࡉࢀࡓ㸦ᅗ㸷㸧ࠋ௚ࡢ⸴㢮࡜ẚ࡭ࡿ࡜ⱝᖸከ࠸ࡶࡢࡢ㸪㝣ୖ᳜≀࡜ẚ㍑ࡍࡿ࡜ᅽಽⓗ࡟ᑡ࡞࠸ࠋ඲㑇ఏᏊ࡟༨ࡵࡿ
๭ྜࡶ⣙ 1.5%࡜௚ࡢ⸴㢮࡜ྠ⛬ᗘ࡛࠶ࡾ㸪㝣ୖ᳜≀ࡢ඲㑇ఏᏊ࡟༨ࡵࡿ㌿෗ᅉᏊࡢ๭ྜࡼࡾ
ᑡ࡞࠸ࡶࡢ࡛࠶ࡗࡓࠋࡋ࠿ࡋ࡞ࡀࡽࡑࡢ✀㢮ࢆẚ㍑ࡍࡿ࡜㸪௚ࡢ⸴㢮ࡼࡾ᱁ẁ࡟ࣂ࢚࣮ࣜࢩ
ࣙࣥࡀቑ࠼࡚࠸ࡿࡇ࡜ࡀࢃ࠿ࡿ㸦ᅗ㸯㸮a㸧ࠋࡲࡓࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘࡜㝣ୖ᳜≀ࡢඹ㏻♽
ૡ୼౤৕ਯ

0

ᇖథ

३॔ॽॹॕड़३॰থ



॥থॻছ५



३ड़঑ॻট



ইख़ड़ॲॡॳছ঒

ड़५ॺঞड़॥ॵढ़५

঑ॡটঔॼ५



ॡটঞছ



ॡছ঑ॻঔॼ५

঎ঝ঎ॵॡ५

ॡঞঈ९ঝ঑ॹॕक़঒

ಮ঱
೘੟

500

1000

2000

2500

3000



঄ওॶজफ़ॿ०ॣ
ॖॾढ़ॱ঄ং

1500




ॖॿ



এউছ



३টॖॾॼ६ॼᅗ㸷 15 ⏕≀✀࡟࠾࠸᳨࡚ฟࡉࢀࡓ㌿෗ᅉᏊᩘᅗෆࡢᩘ್ࡣ඲㑇ఏᏊ࡟༨ࡵࡿ㌿෗ᅉᏊࡢ๭ྜࢆ♧ࡍࠋ

K. Hori & H. Ohta-7

BSJ-Review 7:61 (2016)

植物科学最前線 7:62 (2016)ඛࡀศᒱࡋࡓᚋ࡟ࡶ㌿෗ᅉᏊࡢ✀㢮ࡢቑຍࡀࡳࡽࢀ㸪᳜≀ࡢ㝣ୖ໬ࡢ๓ᚋ࡛㌿෗ᅉᏊࡢ✀㢮㸪ᩘࡀቑຍࡋ㸪ከᵝ࡞㑇ఏᏊไᚚࢆ㌟࡟╔ࡅࡓࡇ࡜ࡀఛ࠼ࡿࠋࡉࡽ࡟㌿෗ᅉᏊ࡟㝈ࡗࡓࢻ࣓࢖
ࣥࡢ⤌ࡳྜࢃࡏᩘࢆࣉࣟࢵࢺࡋࡓ⤖ᯝ㸪⿕Ꮚ᳜≀࡛ࡣࢻ࣓࢖ࣥࡢ⤌ࡳྜࢃࡏࣃࢱ࣮ࣥࢆቑຍ
ࡉࡏ࡚࠾ࡾ㸪㌿෗ᅉᏊࡢከᵝ໬ࡣ㡯┠㸳࡛㏙࡭ࡓ㑇ఏᏊࡢከᵝᛶࡢ⋓ᚓ㐣⛬ࡢ඾ᆺⓗ࡞౛࡜⪃࠼ࡽࢀࡓ㸦ᅗ㸯㸮b㸧ࠋࡇࡢࡼ࠺࡞㌿෗ᅉᏊࡢ✀㢮ቑຍࡸ㸪⤌ࡳྜࢃࡏࣃࢱ࣮ࣥࡢቑຍࡣ㸪ᛴ⃭࡟㑇ఏᏊࢿࢵࢺ࣮࣡ࢡࢆᣦᩘⓗ࡟」㞧໬ࡉࡏࡓ஦ࡣ㛫㐪࠸࡞࠸࡛࠶ࢁ࠺ࠋࡇࡢࡇ࡜ࡣ㸪㝣ୖ࡟㐍ฟࡋࡓ᳜≀ࡀ㸪ᵝࠎ࡞ࢫࢺࣞࢫ⎔ቃ࡟ᰂ㌾࡟㐺ᛂࡋ㸪ᵝࠎ࡞⤌⧊ࢆศ໬ࡉࡏ㸪ከᵝ࡞⣽⬊ࡢ≧ែࢆᐇ⌧࡛ࡁࡿࡼ࠺࡟࡞ࡗࡓ኱ࡁ࡞せᅉ࡜⪃࠼ࡽࢀࡿࠋ௨ୖࡢ⤖ᯝ࠿ࡽ㸪㝣ୖ᳜≀࡜ࡢ㌴㍈⸴᳜≀ࡢඹ㏻♽ඛࡢ࡞࠿࡛㸪ẚ㍑ⓗ᪩ࡃ࡟ศᒱࡋࡓࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡀࢩࣥ
ࣉࣝ࡞యไࢆᣢࡘ࡟ࡶ࠿࠿ࢃࡽࡎ㸪㝣ୖ᳜≀ࡢ♏࡜࡞ࡿᇶᮏⓗ࡞㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡࢆ
⋓ᚓࡋ࡚࠾ࡾ㸪㝣ୖ⎔ቃ࡟㐺ᛂࡋ࡚࠸ࡃ㐣⛬࡛㑇ఏᏊ㔜」࡜ࢻ࣓࢖ࣥࡢ⤌ࡳྜࢃࡏ᪉ࢆ฼⏝ࡋ࡚㸪᪤Ꮡࡢ㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡࢆ㌿⏝ࡋ࡚Ⓨ㐩ࡉࡏ࡚࠸ࡗࡓ࡜࠸࠺㝣ୖ㐍ฟࡢࡓࡵࡢᡓ␎ࡀぢ࠼࡚ࡃࡿࠋ௒ᚋ᳜≀࣍ࣝࣔࣥࡢⓎ㐩࡜ྠᵝ㸪ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡢ㌿෗ᅉᏊࡀఱ
࡟ᛂ⟅ࡋ㸪ఱࢆไᚚࡋ࡚࠸ࡿࡢ࠿᫂ࡽ࠿࡟ࡍࡿࡇ࡜ࡀ㔜せ࡜࡞ࡗ࡚ࡃࡿ࡜ᛮࢃࢀࡿࠋ

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ᅗ㸯㸮 15 ⏕≀✀ࡢ㌿෗ᅉᏊࡢ✀㢮ᩘ㸪ࢻ࣓࢖ࣥࡢ⤌ࡳྜࢃࡏᩘ࡜⥲㑇ఏᏊᩘࡢ㛵ಀa.
b.

ྛ⏕≀✀ෆࡢ㌿෗ᅉᏊࡢ✀㢮ᩘ࡜⥲㑇ఏᏊᩘࡢࣉࣟࢵࢺ
ྛ⏕≀✀ෆࡢ㌿෗ᅉᏊ࡟࠾ࡅࡿࢻ࣓࢖ࣥࡢ⤌ࡳྜࢃࡏᩘ࡜⥲㑇ఏᏊᩘࡢࣉࣟࢵࢺ

㸶㸬௒ᚋࡢᒎᮃ
ࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࡢࢤࣀ࣒ゎㄞ࡟ࡼࡾ㸪᳜≀ࡢ㝣ୖ㐍ฟ๓ࡢẁ㝵࡛ࡍ࡛࡟㝣ୖ໬ࢆྍ⬟࡜ࡍࡿࡼ࠺࡞ᅉᏊࡀฟ⌧ࡋ࡚࠸ࡿࡇ࡜ࡀศ࠿ࡗ࡚ࡁࡓࠋࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘௨እ࡟ࡶ㌴㍈⸴᳜≀ࡢࢤࣀ࣒ゎㄞࡀ㐍ࢇ࡛࠾ࡾ㸪௒ᚋ㝣ୖ㐍ฟࡢ㐣⛬࡛㉳ࡁࡓࢤࣀ࣒ࡢኚ໬ࢆࡼࡾヲ⣽࡟ㄪ࡭ࡿࡇ࡜ࡀ࡛ࡁࡿࡼ࠺࡟࡞ࡗ࡚࠸ࡃࡔࢁ࠺ࠋ௒ᚋࡉࡽ࡟᳜≀ࡢ㝣ୖ໬ࢆ᫂ࡽ࠿࡟ࡋ࡚࠸
ࡃࡓࡵ࡟ࡣ㸪ࡇࢀࡽࡢᅉᏊࡀ㝣ୖ໬๓ࡢ㌴㍈⸴᳜≀࡟࡝ࡢࡼ࠺࡞ᙳ㡪ࢆ୚࠼ࡓࡢ࠿ゎ᫂ࡍࡿࡇ࡜ࡀḟࡢẁ㝵࡛࠶ࡿ࡜ᛮࢃࢀࡿࠋ㌴㍈⸴᳜≀ࡣῐỈ࡜㝣ୖ࡜ࡢ㝿࡟⏕ᜥࡋ࡚࠸ࡿࡶࡢࡀከ
ࡃ㸪㒊ศⓗ࡞ࡀࡽࡇࢀࡽࡢᅉᏊࡀ༙㝣ୖ≧ែࡢ⏕⫱࡟᭷฼࡛࠶ࡾ㸪ᚎࠎ࡟ࡼࡾཝࡋ࠸㝣ୖ⎔
K. Hori & H. Ohta-8
BSJ-Review 7:62 (2016)

植物科学最前線 7:63 (2016)ቃ࡟⏕⫱ࢆᗈࡆ࡚࠸ࡃ࡟࠶ࡓࡗ࡚㸪ᶵ⬟ࡢከᵝ໬ࡸᙉ໬ࢆᡂࡋ㐙ࡆࡓ஦ࡀ⪃࠼ࡽࢀࡿࠋࡲࡓ
௚ࡢྍ⬟ᛶ࡜ࡋ࡚㸪๓㐺ᛂ࡜ࡋ࡚ඹ㏻♽ඛ࡛ࡣ␗࡞ࡿᶵ⬟ࢆᢸࡗ࡚࠾ࡾ㸪㝣ୖ໬࡟࠶ࡓࡗ࡚
ண᝿እ࡟᭷฼࡟ാ࠸ࡓᅉᏊࡀ࠶ࡿྍ⬟ᛶࡶ⪃࠼ࡽࢀࡿࠋಶࠎࡢලయⓗ࡞ᅉᏊࡢ㐍໬㐣⛬ࢆ᫂ࡽ࠿࡟ࡍࡿࡓࡵ࡟ࡣ㸪ᐇ㦂ⓗ࡟ࡑࡢᶵ⬟ࢆᐇドࡍࡿࡇ࡜ࡀᚲせ࡛࠶ࡾ㸪㑇ఏᏊ᧯స⣔ࢆ☜❧ࡋ㸪ᇵ㣴ἲ㸪ᐇ㦂⣔ࡶࡼࡾᢅ࠸ࡸࡍࡃࡋ࡚࠸ࡃࡇ࡜ࡀᚲ㡲࡛࠶ࡿࠋࡇࡢࡇ࡜ࡣ᳜≀ࡢ㝣ୖ໬ࡢゎ᫂ࡢࡳ࡞ࡽࡎ⸴㢮◊✲ࡢⓎᒎ࡟ࡶ኱ࡁࡃ㈉⊩ࡍࡿ࡛࠶ࢁ࠺ࠋ㌴㍈⸴᳜≀㛛࡛ࡣ᥋ྜ⸴⥘ࡢࣄ࣓࣑࢝ࢷ࢟ࣔ㸪ࢱࢸࣈ࢚ࡀ㸪ࡑࢀࡒࢀࣃ࣮ࢸ࢕ࢡࣝ࣎ࣥࣂ࣮ࢻ࣓ࣥࢺἲ㸪࢔ࢢࣟࣂࢡࢸ
࣒ࣜ࢘ἲ࡟ࡼࡗ࡚ᙧ㉁㌿᥮࡟ᡂຌࡋ࡚࠾ࡾ(Abe et al. 2011, Sørensen et al. 2014)㸪ᡃࠎࡶࢡࣞࣈ
ࢯ࣑ࣝࢹ࢕࣒࡛࢘ᛴࡂᙧ㉁㌿᥮⣔ࡢ☜❧ࢆ㐍ࡵ࡚࠸ࡿࠋ௒ᚋࢡࣞࣈࢯ࣑ࣝࢹ࢕࣒࢘ࢆึࡵ࡜
ࡋ࡚ᵝࠎ࡞㌴㍈⸴᳜≀ࡀࣔࢹࣝ⸴㢮࡜ࡋ࡚☜❧ࡋ㸪᳜≀ࡢ㝣ୖ໬ࢆྵࡵ㸪⏕≀㐍໬ࡢ࠶ࡾ࠿ࡓࡀᇉ㛫ぢ࠼ࡿࡇ࡜ࢆᮇᚅࡋ࡚࠸ࡿࠋ

ㅰ㎡
ᮏ◊✲ࡣ᪥ᮏᏛ⾡᣺⯆఍㸪ᖹᡂ㸰㸯ᖺᗘ㹼ᖹᡂ㸰㸳ᖺᗘࢢ࣮ࣟࣂࣝ COE ࣉࣟࢢ࣒ࣛࠕᆅ⌫࠿ࡽᆅ⌫ࡓࡕ࡬ࠖࡢ୍⎔࡜ࡋ࡚᥎㐍ࡉࢀ㸪ᖹᡂ㸰㸱ᖺᗘ࠿ࡽ⌧ᅾࡲ࡛⛉Ꮫᢏ⾡᣺⯆ᶵᵓ㸪ᡓ␎ⓗ๰㐀◊✲᥎㐍஦ᴗ(CREST) ࠕ᳜≀ᰤ㣴⣽⬊ࢆࣔࢹࣝ࡜ࡋࡓ⸴㢮⬡㉁⏕⏘⣔ࡢᡓ␎ⓗᵓ⠏ࠖࡢ୍⎔࡜ࡋ࡚ຍ㏿ⓗ࡟᥎㐍ࡉࢀ࡚࠸ࡿࠋࡲࡓ୍㒊ࡣᖹᡂ㸰㸲ᖺᗘ࠿ࡽࡢᩥ㒊⛉Ꮫ┬㹕㹎㹇ࣉࣟࢢ࣒ࣛ㸪ᆅ⌫⏕࿨◊✲ᡤ࡟ᘬࡁ⥅ࡀࢀ࡚࠸ࡿࠋ࡞࠾ Klebsormidium flaccidum NIES-2285
ࡣᅜ❧⎔ቃ◊✲ᡤ㸪ᚤ⏕≀⣔⤫ಖᏑ᪋タࡼࡾศㆡ㡬࠸ࡓࠋࡲࡓᮏ◊✲ࡣࢡࣞࣈࢯ࣑ࣝࢹ࢕࢘
࣒ゎᯒࢳ࣮࣒(http://www.plantmorphogenesis.bio.titech.ac.jp/~algae_genome_project/klebsormidiu
m/kf_team.htm)࡟ࡼࡿඹྠ◊✲࡛࠶ࡾ㸪඲ဨࡢከ኱࡞㈉⊩ࡀ࠶ࡗ࡚ࡢ◊✲࡜࡞ࡗࡓࠋࡇࡇ࡟グࡋ࡚῝ࡃឤㅰࡢពࢆ⾲ࡋࡓ࠸ࠋᘬ⏝ᩥ⊩Abe, J., Hori, S., Tsuchikane, Y., Kitao, N., Kato, M. & Sekimoto, H. 2011. Stable nuclear
transformation of the Closterium peracerosum–strigosum–littorale complex. Plant Cell Physiol.
52: 1676-1685.
Battistuzzi, F.U. & Hedges, S.B. 2009. A major clade of prokaryotes with ancient adaptations to life
on land. Mol Biol Evol. 26:335-343
Clarke, J.T., Warnock, R.C.M. & Donoghue, P.C.J. 2011. Establishing a time-scale for plant evolution.
New Phytol. 192: 266-301
Edwards, D. & Kenrick, P. 2015. The early evolution of land plants, from fossils to genomics: acommentary on Lang (1937) 'On the plant-remains from the Downtonian of England and Wales'.
Phil. Trans. R. Soc. B 370: 20140343.
Elster, J., Degma, P., Kováþik, ď., Valentová, L., Šramková, K., & Pereira, A. B. 2008. Freezing anddesiccation injury resistance in the filamentous green alga Klebsormidium from the Antarctic,
Arctic and Slovakia. Biologia 63: 843–851.
Hori, K., Maruyama, F., Fujisawa, T., Togashi, T., Yamamoto, N., Seo, M., Sato, S.,

Yamada, T.,

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Mori, H., Tajima, N., Moriyama, T., Ikeuchi, M., Watanabe, M., Wada, H., Kobayashi, K., Saito,
M., Masuda, T., Sasaki-Sekimoto, Y., Mashiguchi, K., Awai, K., Shimojima1, M., Masuda, S.,
Iwai1, M., Nobusawa, T., Narise, T., Kondo, S., Saito, H., Sato, R., Murakawa, M., Ihara, Y.,
Oshima-Yamada, Y., Ohtaka, K., Satoh, M., Sonobe, K., Ishii, M., Ohtani, R., Kanamori, M.,
Honoki, R., Miyazaki, D., Mochizuki, H., Umetsu, J., Higashi, K., Shibata, D., Kamiya, Y., Sato,N., Nakamura, Y., Tabata, S., Ida, S., Kurokawa, K., Ohta, H. 2014. Klebsormidium flaccidum
genome reveals primary factors for plant terrestrial adaptation. Nat. Commun. 5: 3978.
Jin, J., Zhang, H., Kong, L., Gao, G. & Luo, J. 2013. PlantTFDB 3.0: a portal for the functional andevolutionary study of plant transcription factors. Nucl. Acids Res., 42(D1):D1182-D1187.
Karsten, U. & Holzinger, A. 2012. Light, temperature, and desiccation effects on photosynthetic
activity, and drought-induced ultrastructural changes in the green alga Klebsormidium dissectum(Streptophyta) from a high alpine soil crust. Microb. Ecol. 63: 51–63.
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exhibits cold acclimation that is closely associated with compatible solute accumulation andultrastructural changes. Plant Cell Environ. 31: 872–885.
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Geochem. Trans. 13, 7.
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K. Hori & H. Ohta-10

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植物科学最前線 7:65 (2016)

Shaw, A.J., DeGironimo, L., Stevenson, D.W., Surek, B., Villarreal, J.C., Roure, B., Philippe, H.,
dePamphilis, C.W., Chen, T., Deyholos, M.K., Baucom, R.S., Kutchan, T.M., Augustin, M.M.,
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Acad. Sci. USA 111, E4859-E4868.
኱࿴຾ᖾ㸪ἙෆᏕஅ 2012. ぢ࠼࡚ࡁࡓࢮࢽࢦࢣࢤࣀ࣒ ᳜≀⛉Ꮫ᭱๓⥺ 3:71.

K. Hori & H. Ohta-11

BSJ-Review 7:65 (2016)

植物科学最前線 7:66 (2016)  8(*$)'%&W/9@V5 Y+ACRHRZ5
ňÍĖò 1*}ÑŒÈü 2
1ǦŕíôôǯƁô÷ǽR|V~ 920-0934 ǦŕĆöŰ 13-1

ǿ ũěāȆƈĪíôŪôǠ 171-8501 ĿǡǉăÇƽœƸ 3-34-1Ȁ
2
Max Planck Institute for Plant Breeding Research*Carl-von-Linné-Weg 10, 50829 Cologne Germany
The evolution of TALE homeobox transcription factors contributes to the increase in the complexity ofthe diploid body plan in land plants
Key words: Alternation of generations; Evolution of sporophytes; Gene duplication; KNOX; BELL
Keiko Sakakibara1*, Chihiro Furumizu2,
1

Advanced Science Research Center, Kanazawa University, Kanazawa 920-0934, Japan(*Current address: Rikkyo University, 3-34-1 Nishi-Ikebukuro,Toshima-ku, 171-8501, Tokyo Japan)
2

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C~TzH#(Ç¾98

K. Sakakibara and C. Furumizu- 1
BSJ-Review 7:66 (2016)

植物科学最前線 7:67 (2016) ǭņţ(ŚŒĕƞƭƳǺ'NuOGtƳǺ'ǩ 6ƒ4®8000ĈÀ&ºĂ#ƥ 6
9!8(Lewis and McCourt 2004; Wellman et al. 2003; Wickett et al. 2014)NuOGtƳǺNuOGt#ǭņţ'Ŭŗū<őǔ!08#ȁNuOGt(Ņŷ Êŷ' &ìƔƨ'Ǣ© <¡8 ȁƼ
ŷ' (ÊƔƨ'ĤÔò'0"8į"ȁǭņţ(Ƽŷ' 3ìƔƨ'ƨò <ČĚ8ǭņţ'ǙÆ'½Ĺ&'Ƒƚ 6ºĂ#ƥ 698KIņţ"(Ŭŗū'íÉ(Êŷ'Ǣ©
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[KOAC  8(*$)'%&W/9! W/9XR] W/9 W/9
 nsCoYGPâǓ¸Þò(ŸŅŬţ µǘ&ġǏŞ'ÑǓ¸Þò'"7ȁŸŅŬţ'µǘƀ°& !ȁDNAƙÔƩ<ſnsC_s@|'ƋȂ+wYYGP#ƋȃlwYGP'ǩ&Ȅ>qdǣŏå'ģ²<×1TALEânsCoYGPǟò#Ȅ>qdǣŏå'ģ² %ǳTALEânsCoYGPâǟò&º 9!#ƥ 69!8(Derelle et al. 2007)ņţ"(TALEâǓ¸Þ
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6Êǲ9 BELLǟò# KNOX ǟò'ºòƑƚǁŀ'ƙł (Mukherjee et al. 2009; Sakakibara 2016)&å NWǺ'³Jdrǁǆ(ũàǙƶ"7ȁĐȁBELL/KNOX ǟòī ĵž&%8#Ĺď98

K. Sakakibara and C. Furumizu- 2
BSJ-Review 7:67 (2016)

植物科学最前線 7:68 (2016)èÄ!8KNOXǟò#ÕŊ&, ǭņţ'BELLǟò&!3ǟòǤƼ&57ǟòīèÄ8¬Ö ǃ2698įȁƳǺ'BELLǟò"(ȁƓƳN>a]?CNU|ȁÏ*ƞƳǺ(ǟòǤƼ&5!Ŭȃ§'BELLǟò<ġ ȁNuOGtƳǺGyjTxq]?Ar
Klebsormidium flaccidum"(Ȃ§óà8ºĂĈ'Ñ 6ȁƳǺ'BELLǟò#ǭņţ'BELLǟò'ƑƚǪ¤(ĺĵ6 "(%, 6%8ǁŀ ď98
 BELLǟò#KNOXǟò'ƀ°ų%ŋƩ<ƥú8"Ǥƾ%ȁƞƳǺ'TALEânsCoYG
Pǟò&Ǫ8ŽƆ<ō'Ƿ"ƕ8


[KOACRHGE#2<  8(*$)'%&W/9 ÊƔƨƞƳǺGvq_t`P Chlamydomonas reinhardtii (kvP#p@`P'ȃ'Ǣâ<ġȁŃǼŌĽ"(Êŷ'ŃǼƔƨ Ǣ©ò+#ºÆ8'ǯ&ȁkvPâǢ©ò( BELL ŊV|fGǍ<K~_8 GSP1 (Gamete-specific plus1)ǟò<ȁp@`PâǢ©ò( KNOX V|fGǍ<K~_8 GSM1 (Gamete-specific minus1)ǟò<Ųũ896'V|fGǍ(99'Ǣ©
ò& !(ƔƨǍ&Āà8KNOX V|fGǍ(Ǔ¸Þò#!­ ȁƪǒ(ŅƂƶNH`x<ġȁĤÔĐȁBELL V|fGǍ#l\zW@p~<ČĚ8#"Ņ+#ƂƶȁĤÔòǿƼ
ŷȀŤűų%ǟòŲũ<Ǉƍ8#"Ƽŷ 'ºÆ<¿Ē!8# ſ9(Lee et al.
2008; Zhao et al. 2001)96'÷ǽ4ƞƭņţ& 8 TALE ǟò'Ǣ¼ǁŀ'ƙł&å !ȁ
Lee 6(2008)( KNOX ǟò# BELL ǟò'ǙÆ ǭņţ'Ƽŷ 'ìƔƨÆ&Ǥƾ"# ǅ<ĦÝ!8ō'Ƿ"(ǭņţ'Ŭŗū'ǙÆ& 8 TALE ânsCoYGPǟ
ò'ĎÂ<ƥú8"Ǥƾ%Żƿ<36ȁgsZwEcLI' KNOX ǟòİ# BELL ǟòİ&Ǫ8ŽƆ<ƕ8(Horst et al. 2016; Sakakibara et al. 2013)

ßȄ ÊƔƨƞƳGvq_t`P'Ŭŗū# KNOX ǟò# BELL ǟò'ĎÂ Lee et al. 2008 57ĩë

K. Sakakibara and C. Furumizu- 3
BSJ-Review 7:68 (2016)

植物科学最前線 7:69 (2016)[  W/9 W/9Y+AC,.-.#2<  åǠǭņţ& 8 KNOX2 ǟòİ# BELL ǟòİ'ŋƩ&!(KIņţR|Ǻ'gsZwEcLI Physcomitrella patens <Ůǁŀ Ǚ="8gsZwEcLIJdr&( 2 §' KNOX2 ǟò K~_9! 7ǿßȃȀ
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6Ǣ© ŊƗƣ<ºÆ# 6ȁKNOX2 ǟò Ƽŷ& !Ǣ© 'ŲŬkzHvr<ĝ¿8#" <¿Ē!8# ſ9(Sakakibara et al. 2013)
 įȁgsZwEcLI(ȅ§'BELLǟò<ġ ȁ' '"8PpBELL1ǟò<Ǣ© & !ǛÁŲũ8#ȁŅŷ(Êŷ'//ƨò ŊƗƣ Ǆý9(Horst et al. 2016)'ƙł 6ȁgsZwEcLI'BELLǟò ƨò 'ŲŬkzHvr<¿Ē8#&57ȁ '»ħ &Ǫ8# ſ9/ȁBiFCǁŀ#Ú)98ĜŖ"VeK'ƷŴƔƨ<Ů!V|
fGǍǩ'ŷ¡Ů<ǁŀƙłȁPpBELL1V|fGǍ#ƴ±V|fGǍĭŢ#'ƵÔV|fGǍgsZwEcLI'KNOX1/(KNOX2V|fGǍ#ƴ±V|fGǍĭŢ#'ƵÔV|fGǍ'9#3ŷ¡ŮÒƩ"8# ſ9!8'ƙł(ȁgsZwEcLI"3PpBELL1
V|fGǍ gsZwEcLIKNOXV|fGǍ'9 #ţŪų&ŷ¡Ů! <¿Ē
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6Êǲ9KNOX1 ǟòİ&ā8ǟò"8 Knotted-1 "7(Vollbrecht et al. 1991)ȁKNOX1 ǟòİ&!(ƹòņţ<ŮŽƆ ìīȁæØ9!8(Hay and Tsiantis 2010)^AtzKN' Knotted-1 ǟò'æØ&ƛ!Nz@b`Q` Arabidopsis thaliana 'ƯǵºƺƗƣŌî8ëű

shootmeristemless (stm)'ÍÞǟò#! STM ǟò Êǲ9, KNOX1 V|fGǍ<K~_!8
# ĵ6 &%(Barton and Poethig 1993; Long et al. 1996)/ȁNz@b`Q`'' KNOX1 f
vzH'ŋƩŌĨëű "8 knat1/brevipedicellus (bp)(ƮƯ'ǧ ð698Ʒũâ<ſȁƔƨéÔĚǟòȁŤ&wHa|ÔĚƘǐ'ǟòŲũ Ĵ!8# 6ȁKNAT1/BP ǟò(Ɣƨ'ĺºÆťę<ǝ»&ƜġȁƮƯ'ƔƨºÆ#ǧ8À&wHa|'ŔŹ!/ '<ĝ¿!8#ƥ 69!8(Mele et al. 2003)įȁKNAT1/BP ǟò<Nz@b`Q`"ǛÁŲũ8#ư&űěų&ºƺƗƣ ČĚ98(Chuck et al. 1996)96'÷ 6ȁƹòņţ
KNOX1 ǟò(ƨò 'ƯǵºƺƗƣ'ČĚƜġ#ƔƨºÆ'ĝ¿&ŋƩ!8#ƥ 69!8(Hay and Tsiantis 2010)
 åǠǭņţ"8KIņţgsZwEcLI'ġȄ§' KNOX1 ǟò(ƨò 'ǵƉƔƨ#ºƺƗƣ"ŋƩ8gsZwEcLI'ƨò 'ºƺƗƣ(ƧŲŬ'½Ĺ&ČĚ98 ȁĐ&ºƺƗƣ řîȁȂ§'ƨò' <ºÆ!ƨò 'ŲŬ<Ɩ8gsZwEcLI' KNOX1ǟò(93'ƨò &Ķų&¡698ºƺƗƣ"Ųũ8gsZwEcLI KNOX1 ǟ

K. Sakakibara and C. Furumizu- 4
BSJ-Review 7:69 (2016)

植物科学最前線 7:70 (2016)òǤŋƩŌĨëű & !(ƨò 'ºƺƗƣ ǝ»&Ɯġ9%2ȁƙł#!*%ƨò <ČĚ8'# 6ȁKNOX1 ǟò(gsZwEcLI& !3ƹòņţ'KNOX1 C~TzH#ÕŊ&ºƺƗƣ<Ɯġ8ŋƩ<ğ!8#ƥ 698(Sakakibara et al. 2008)
 Nz@b`Q`'5 %Êư<ČĚ8ƹòņţ"( KNOX1 ǟò(ƯǵºƺƗƣ"Ųũȁư
ÍåČĚ&°ƈ!õưǸã"Ųũ řî89&û!ȁ^p^ Solanum lycopersicum 4Nz@b`Q`'ǖƢƄ'qXVcZIe` Cardamine hirsuta %$Ƽư<ČĚ8ņţ"( KNOX1 ǟò'Ųũ ưÍå'þưÍå"·*ŗĕÆ8#"Ƽư'ŲŬ ¿Ē9!8# Ż69!
8(Blein et al. 2008; Hay and Tsiantis 2010)/ȁƼư<ČĚ8ņţ" KNOX1 ǟò<ǛÁŲũçÔ(Ƽư'ƼǱ Ǚ98(Hareven et al. 1996)96'ŽƆ( KNOX1 ǟò'ŲũŊċ'ëÆ ƹòņţ'ư'ǙÆ#ìŊÆ&ǌŧ#<ſÜ!8'5 &ƹòņţ'ƨò 'Ưǵ"( KNOX1 ǟò'Ųũ(ĶƇǩų&Ɛơ&¿Ē9! 7ȁƯǵºƺƗƣ"ǧĹǩƜġ98į, ŲŬ½Ĺ'ưÍå"(ĝ¿98# ƼǱ%ŲũŊċ<ſ ȁgsZwEcLI'KNOX1 ǟò'Ųũ(ƨò 'ƧŲŬ'½Ĺ'ºƺƗƣ&Ǭõ9!8ƹòņţ'ưÍå"'KNOX1 ǟò'Ųũĝ¿&Ǫ8Ǔ¸Þò#! MYB _s@|<×1 ARP (ASYMMETRICLEAVES1/ROUGH SHEATH2/PHANTASTICA)V|fGǍȁLOB (LATERAL ORGAN BOUNDARIES)_s@|<×1 AS2 (ASYMMETRIC LEAVES2)V|fGǍ'ƼÔ 4 YAB (YABBY) V|fGǍ%$ æØ9!8(Byrne et al. 2000; Hake et al. 2004; Kumaran et al. 2002)96'ǇƍÞò'ǠǿAS1ȁAS2 4 YABȀ'C~TzH(gsZwEcLIJdr&(óà(Rensing et al. 2008;
Sakakibara et al. 2008)ȁǭņţ'ǙÆ'Ǜƃ"ȁƹòņţ<×1Ƒƚ" KNOX1 ǟò'ŲũǇƍ&Ǫ;8Į%^v|PÞò Ũđ9#ĥŝ98Ä !ŲũǇƍ&Ǫ;8NPǸã ǙÆƙłȁKNOX1 ǟò'Ųũ¿ĒcY^{~G ƼǱ&%#"ȁƯǵ"'Ųũ Ɯġ985&%7ȁƹòņţ'Ƒƚ"'ƨò 'ƼǱÆ&% #ƥ 698(榊原恵子 2013)

[P9AC W/9BN Knotted-1ǟò'Êǲ<ƉƟ#!ȁ
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ǟò( KNOX1 ǟò#(ű%7ȁƨò 'ºÆƗƣ"&ŋƩ8# ĥŝ9! ȁ

K. Sakakibara and C. Furumizu- 5
BSJ-Review 7:70 (2016)

植物科学最前線 7:71 (2016)'ǅ<ƻ8ķǖ'ŽƆĚł<ō&ƕ8
 Nz@b`Q` KNOX2 ǟòfvzH'ŏ7'Ȅ§'ÊŦëű (93ǥŬâ#-.ÕŊ'Ʒũâ<ſ ȁknat3 knat4 knat5 Ǥëű (Nz@b`Q` KNOX1 ǟò<ǛÁŲũĶ'
5 &ư »9Ǖ1Ʒũâ<ſ(Furumizu et al. 2015; Serikawa et al. 1997a; Truernit et al. 2006)/ȁ
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ò K~_8 KNOX2KNOX ǟò( KȁKNOX1 ǟò( K1ȁKNOX2 ǟò( K2ȁBELL

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2012)KNOX2 Ǔ¸Þòǟò(KNOX1 ǟòİ#KNOX2 ǟòİ&ºĂ#ĥŝ'V~JY^'Õõ%$'Đ'ǁŀĔƾ"8 ȁ'÷
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K. Sakakibara and C. Furumizu- 6
BSJ-Review 7:71 (2016)

植物科学最前線 7:72 (2016)&¿Ē8ÒƩĕ ſÜ9(ßȅ; Furumizu et al. 2015)
 °&Ƽư<ČĚ8ņţ& ! KNOX1 ǟò Ƽư'ƼǱ<Ǚ8#<ǂÏ ȁq
XVcZIe`& 8ƼưČĚ&( KNOX1 ǟò Ĕƾ"7ȁKNOX1 ǟò'ǛÁŲũ(Ƽư<ŉĚ8þư'Łī<èÄ8(Hay and Tsiantis 2006)įȁqXVcZIe`" KNOX2 ǟò<ǛÁŲũ8#Ƽư'ƼǱ Ŝÿ8# 6ȁNz@b`Q`'áǠ'ČęČĚ#ÕŊ&ȁ
qXVcZIe`'ƼưČĚ& !3 KNOX2 ǟò( KNOX1 ǟò#Ðû'ĎÂ<ł#ſÜ9(Furumizu et al. 2015)96'Żƿ 6ȁƹòņţ&ƫ8Ƒƚ" KNOX1 ǟòİ#KNOX2 ǟòİ'ŋƩ íºÆ#ĥŝ98KNOX1 ǟò# KNOX2 ǟò'ŲũǸ
ã&Ǥ%7 %#, /Ǔ¸Þò#!ŋƩ'ǜ ĥŝ98# 6, ǟò'Ųũ<Ǉƍ8NPǸã#ǟò'ŭţ"8V|fGǍ #3&ëÆ#"ŋƩºÆ&ƫ#ƥ 69
8 /ȁƹòņţ"' KNOX ǟòİ'Ǔ¸Þò'ŋƩ'ëÆ<ï÷&ſ¢#!ȁNz@b`Q`4^p^"Õõ9ȁnsC_s@|<Ō KNOX ŊV|fGǍ KNATM 4 PTS/TKD1

Ģ

69896' DNA ƙÔƩ<ġ% KNOX ŊV|fGǍ(ǘć' KNOX Ǔ¸Þò'ŋƩ<ƊÔų&ǫøȁKNOX Ǔ¸Þò&58V~JY^ǟò'Ųũ¿Ē'ĝ¿Þò#!­#ſÜ9!8(Kimura et al. 2008; Magnani and Hake 2008)

[  W/9 W/9XRP9ACM907+02RZ5TD Nz@b`Q`Jdr&( 13 §' BELL ǟò ×/98
ŋƩŌĨëű 'ǟôų%ǁŀ&5
7, STM ǟò%$' KNOX1 ǟò( PENNYWISE 4 POUND-FOOLISH %$' BELL ǟò##3&ŋƩ8# ſ9!8(Byrne et al. 2003; Rutjens et al. 2009; Smith and Hake 2003)į, ŋƩŌĨëű 'Ʒũâ'Ǻ4ǛÁŲũņţ 'ǁŀ 6 KNOX2 ǟò( BELL1ȁSAWTOOTH1 (SAW1)ȁ
SAW2 %$' BELL ǟò##3&ŋƩ8ÒƩĕ ſÜ9! 7ȁNz@b`Q`' BELL ǟò&(ŋƩ'ºÆ 8# ĥŝ98(Furumizu et al. 2015)96'ŽƆ 6ȁBELL ǟòİ¶"'ǟòǤƼ#'Đ'ŋƩºÆ3 KNOX ǟò#'ŷ¡Ů'Ɨ0Ô;'ìŊÆ&ù#ƥ 698KNOX ǟò# BELL ǟò'Ɨ0Ô;'ìŊÆ 57ǾĊ%Ǔ¸ǇƍcY^{~G'ŉƏ<ÒƩ&ȁ'ƙłȁƹòņţ'áǠ'ƼǱ%o]?kv|'ǙÆ&3ǌŧ'"(% #ƥ 698["
 2014 Ĉ&´Ǩ9NuOGtƳǺGyjTxq]?Ar'Jdrėæ<×2Ƒƚǁŀ&57ȁ
KNOX ǟòİ(ǭņţ'ìŊÆ&°ƈ!ȁKNOX1 # KNOX2 'ǟòİ&º 9!
# ĵ6 #%ǿßȃ; Hori et al. 2014; Sakakibara 2016)NuOGtƳǺ"( (æØ9! 6ȁ 6ȁǭņţ'Ƒƚ"' KNOX1 # KNOX2 'ǟòİ"ĮǀŋƩŨđ

#&57ȁƨò 'ƼǱ%ìƔƨ ¿'žƈ&ŋƩ85 &%#ƥ 698NuOGtƳ
Ǻ& !96' KNOX1 ǟòȁKNOX2 ǟò 99$'5 %ŋƩ<ğ!8 (ƬK. Sakakibara and C. Furumizu- 7
BSJ-Review 7:72 (2016)

植物科学最前線 7:73 (2016)Ù ś  #  :   ǖ Ĉ ȁ N u O G t Ƴ Ǻ n N q _ z Ŷ ' g s q D [ F t Closterium
peracerosum-strigosum-littorale complex #V\jB Penium margaritaceum & !ČǍǓħƑ'žƈ
ŷō"æØ9!

7(Abe et al. 2011; Sørensen et al. 2014)ȁĐȁNuOGtƳǺ"'KNOX1/KNOX2 ǟòİ4 BELL ǟò'ŋƩǁŀ ď98
 /ȁƓƳ 63 KNOX ǟò# BELL ǟò 990 ! 7(Matsuzaki et al. 2004;
Mukherjee et al. 2009)ȁKNOX-BELL Ƙǐ'ǙÆ<ƥ 8"96'ŋƩǁŀ3+=ƬÙś
KNOX ǟòİ# BELL ǟòİ'ǏŞ< '.!08#ȁ96'ǟòİ(ƓƳ#ƞƭņţ'µǘƀ°"'ǟòǤƼ&5!Ŭ#ĥŝ9ȁǏŞ'ÑǟòǤƼ#'Đ'ǙÆ ņţ'o]?kv|4ǭņţ'ŬŗÓ'ǙÆ&í%čǴ< #ƥ 698 Ļƅ"(ŲŬǟôų%ŽƆ ¯÷!8ƞƳǺ#KIņţȁ!ƹòņţ'ŽƆ<ē&ƕ! ȁ'ºǺƤ& 8ǟòŋƩ'ǁŀ&Ö÷ǽƑ'Ĭ«3ǖĈȁǻǑų&Ǚ="8NWņţ& !(ũà´Ǩ9!8þưǺ'@bDVgeSelaginella moellendorffiiJdr
"%ȁƲ' NW"8wXu~_qQ{vh%$'Jdrǁŀ3Ǚ269! 7(Sessa et al.
2014)ȁwXu~_qQ{vh%$"ǟòý² ÒƩ%# æØ9 (Muthukumar et al. 2013;
Plackett et al. 2014)ĐȁNWņţ"'99'ǟòİ¶"'ǟòī ĵ6 #%7ȁ96'ǟò'ŋƩǁŀ Ǚ2698#"ȁƜƌļņţ"'KNOXǟòİ#BELLǟòİ'ǟòǤƼ#ŋƩºÆ<ŐǮų&Ǉ,8# ÒƩ#%8: /ȁũó'KIņţ't]xȁV@Ǻ'SaLIȁR|Ǻ'gsZwEcLI&Ä ȁƋ'KIņţ"8ZdLI't]xÆ3ñ/!
8(Szövényi et al. 2015)Đȁ96'ºǺƤ& 8BELL/KNOXǟò'ŋƩ4ǭņţ"Įǀ&Ũđ#ƥ 698ŋƩ_s@|'ǁŀ Ǚ1#ȁKNOXǟòİÏ*BELLǟòİ'ǟòǤƼ#9&ƛĮǀŋƩŨđ $'5 &ǭņţ'ČęǙÆ4ŬŗÓ'ǙÆ<36' '³
ǋ ĵ6 #%8: SU C~P^vw> Monash íô' John L. Bowman Ëê&ĲǶ 6'ĸŵ% discussion <Ęǈ/
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1

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K. Sakakibara and C. Furumizu- 12
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植物科学最前線 7:78 (2016)᳜≀࡟࠾ࡅࡿ᪥㛗࡟ࡼࡿᡂ㛗┦㌿᥮ไᚚࡢ࣓࢝ࢽࢬ࣒࡜ࡑࡢ㐍໬ᒣᒸᑦᖹ࣭ἙෆᏕஅ
ி㒔኱Ꮫ ኱Ꮫ㝔⏕࿨⛉Ꮫ◊✲⛉ 㑇ఏᏊ≉ᛶᏛศ㔝
ࠛ606-8502 ி㒔ᕷᕥி༊໭ⓑᕝ㏣ศ⏫

Evolutionary insights into the molecular mechanism of
day-length-dependent transition to reproductive growth in plants
Keywords: day length, evolution, growth-phase transition, land plants, Marchantia polymorpha,
reproductive growth
Shohei Yamaoka and Takayuki Kohchi
Graduate School of Biostudies, Kyoto University
Kyoto, 606-8502, Japan㸯㸬ࡣࡌࡵ࡟ ᳜≀ࡣ㸪⣙ 4 ൨ 7 ༓୓ᖺ๓࡟㝣ୖ࡟㐍ฟࡋࡓ
㸦Edwards and Kenrick 2015, Rubinstein et al. 2010, Steemans
et al.,2009, Wellman et al. 2003㸧ࠋ㝣ୖ࡛ࡣ㸪ග㸪 ᗘ㸪஝⇱࣭‵₶ࡀ⤯࠼ࡎኚ໬ࡍࡿࠋᅛ╔ᆺࡢ⏕άࢆႠࡴ㝣ୖ᳜≀ࡣ㸪ࡇ࠺ࡋࡓ⎔ቃࡢኚືࢆ㐺ษ࡟ឤ▱ࡋ㸪⮬㌟ࡢ⏕Ꮡ࡜⏕Ṫࢆ᭱㐺໬ࡍࡿࡓࡵࡢ࣓࢝ࢽ
ࢬ࣒ࢆ㐍໬ࡉࡏ࡚ࡁࡓࠋ࡞࠿࡛ࡶ㸪ᰤ㣴ᡂ㛗࠿ࡽ⏕Ṫᡂ㛗࡬ࡢ㌿᥮㸦ᡂ㛗┦㌿᥮㸧ࢆ࠸ࡘ⾜࠺࠿࡜࠸࠺ࡇ࡜ࡣ㸪᳜≀ࡢ⦾Ṫᡓ␎࡟࡜ࡗ࡚ࡁࢃࡵ࡚㔜せ࡛࠶ࡿࠋ⌧⏕ࡢ㝣ୖ᳜≀ࡢከࡃࡣ㸪୺࡟᪥㛗ࢆឤ▱ࡍࡿࡇ࡜࡛ᡂ㛗┦㌿᥮ࡢࢱ࢖࣑ࣥࢢࢆỴࡵ࡚࠸ࡿࠋ᪥㛗ࡢኚ໬ࡣ㸪ࡉࡲࡊࡲ࡞⎔ቃࡢኚ໬ࡢ࠺ࡕ࡛᭱
ࡶ☜ᐇ࡟ண ࡍࡿࡇ࡜ࡀ࡛ࡁ㸪ࡑࢀ࡟ࡼࡾ᮶ࡓࡿᏘ⠇ࢆ▱ࡿࡇ࡜ࡀ࡛ࡁࡿ࠿ࡽ࡛࠶ࡿࠋࡇࡢ࣓࢝ࢽࢬ
࣒ࡣ㸪᳜≀ࡢ㐍໬ࡢ࡞࠿࡛࠸ࡘ⋓ᚓࡉࢀ㸪ࡑࡢཎᙧ࡜ࡣ࡝ࡢࡼ࠺࡞ࡶࡢࡔࡗࡓࡢࡔࢁ࠺࠿ࠋᮏ⥲ㄝ࡛ࡣ㸪㝣ୖ᳜≀ࡢᇶ㒊⣔⤫ࡢ୍✀࡛࠶ࡿⱏ㢮ࢮࢽࢦࢣ࡟㸪᪥㛗࡟ࡼࡿᡂ㛗┦㌿᥮ࡢ࣓࢝ࢽࢬ࣒ࡢཎᙧࡀ
Ꮡᅾࡍࡿ࡜⪃࠼ࡽࢀࡿࡇ࡜࡟ࡘ࠸࡚ゎㄝࡍࡿࠋ㸰㸬᪥㛗࡟ࡼࡿⰼⱆᙧᡂไᚚࡢ࣓࢝ࢽࢬ࣒ ⿕Ꮚ᳜≀ࡢᡂ㛗┦㌿᥮ࡣ㸪ⰼⱆࡢᙧᡂ࡜ࡋ࡚ほᐹࡉࢀࡿࠋ᪥㛗࡟ࡼࡿⰼⱆᙧᡂไᚚࡢ࣓࢝ࢽࢬ࣒ࡣ㸪㛗᪥᳜≀࡛࠶ࡿࢩࣟ࢖ࢾࢼࢬࢼࢆ౑ࡗࡓศᏊ㑇ఏᏛ◊✲࡟ࡼࡗ࡚኱ࡁࡃ⌮ゎࡀ㐍ࢇࡔࠋࢩࣟ࢖ࢾࢼࢬ
ࢼ࡛ࡣ㸪ⰼⱆᙧᡂࡢࢱ࢖࣑ࣥࢢ࡟␗ᖖࡀ⏕ࡌࡿ࡜ࣟࢮࢵࢺⴥࡢᩘࡀ␗࡞ࡗ࡚ࡃࡿࠋࡇࢀࢆᣦᶆ࡟㸪㛗᪥ࡢୗ࡛㐜ဏࡁ࡜࡞ࡿኚ␗ᰴࡀ༢㞳ࡉࢀ㸪ⰼⱆᙧᡂࢆಁ㐍ࡍࡿᙺ๭ࢆᣢࡘ㑇ఏᏊࡀ᫂ࡽ࠿࡟ࡉࢀࡓࠋ
ࡑࡢ୺࡞ࡶࡢࡣ㸪FT㸪CONSTANS (CO)㸪FLAVIN-BINDING KELCH REPEAT F-BOX1 (FKF1)㸪GIGANTEA(GI)࡛࠶ࡿࠋ ⿕Ꮚ᳜≀࡛ࡣ㸪⣙ 80 ᖺ๓࡟㸪ⴥ࡛᪥㛗ࢆឤ▱ࡋ㸪ࠕࣇࣟࣜࢤࣥ㸦ⰼⱆᙧᡂ࣍ࣝࣔࣥ㸧ࠖ࡟ࡼࡗ࡚ࡑࡢ
ࢩࢢࢼࣝࡀఏ㐩ࡉࢀ㸪ⱼ㡬࡛ࡢⰼⱆᙧᡂࡀ㛤ጞࡉࢀࡿ࡜࠸࠺௬ㄝࡀᥦၐࡉࢀࡓ㸦Chailakhyan 1936,

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植物科学最前線 7:79 (2016)

Garner and Allard 1920, Knott 1934㸧ࠋࡼ࠺ࡸࡃ᭱㏆࡟࡞ࡗ࡚㸪ࡇࡢࠕࣇࣟࣜࢤࣥࠖࡢᐇయࡀ FT ࢱࣥࣃࢡ㉁࡛࠶ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟ࡉࢀࡓࠋ㛗᪥࡟࡞ࡿ࡜㸪㌿෗ᅉᏊ CO ࡀάᛶ໬ࡋ FT 㑇ఏᏊࡢⓎ⌧ࢆಁ㐍ࡍࡿ㸦Kobayashi et al. 1999, Kardailsky et al.1999, Suárez-López et al. 2001㸧
ࠋⴥࡢ⥔⟶᮰ࡢᖌ㒊క⣽⬊࡛ࡘࡃࡽࢀࡓపศᏊ㔞ࡢ FT ࢱࣥࣃࢡ㉁ࡣ㸪ᖌ⟶ࢆ㏻ࡌ࡚ⱼ㡬ศ⿣⤌⧊࡬㐠ࡤࢀࡿ㸦Corbesier et al. 2007, Jaeger
and Wigge 2007, Tamaki et al. 2007㸧ࠋࡑࡇ࡛ FT ࡣ㌿෗ᅉᏊ FD ࡞࡝࡜㌿෗」ྜయࢆᙧᡂࡋ㸪㌿෗ᅉᏊ
APETALA1 ࡞࡝ࡢ㑇ఏᏊࢆⓎ⌧ࡉࡏࡿࡇ࡜࡛ⰼⱆศ⿣⤌⧊ࡢᙧᡂࢆ㛤ጞࡉࡏࡿ㸦Abe et al. 2005, Wigge
et al.,2005, Taoka et al.,2011㸧ࠋࡋࡓࡀࡗ࡚㸪
ⰼⱆᙧᡂࡢ㛤ጞࡣ CO ࡜ FT ࡢάᛶ࡟ࡼࡗ࡚Ỵࡲࡿ࡜࠸࠼ࡿࠋ ࡛ࡣ㸪᪥㛗ࡢኚ໬ࡣ࡝ࡢࡼ࠺࡟ࡋ࡚ CO ࡜ FT ࢆάᛶ໬ࡍࡿࡢࡔࢁ࠺࠿ࠋ᭱㏆ࡢ◊✲࠿ࡽ㸪CO ࡜
FT ࡢάᛶ໬ࡣ㌿෗࣭⩻ヂᚋࡢẁ㝵࡛ከ㔜ࡢㄪ⠇ࢆཷࡅ࡚࠾ࡾ㸪ࡑࡢㄪ⠇࡟ࡣ FKF1 ࡜ GI ࡀ୰ᚰⓗ࡞ᙺ๭ࢆᯝࡓࡋ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚ࡁࡓࠋCO ࡢ㌿෗ࡣᴫ᪥ࣜࢬ࣒ࢆ♧ࡋ㸪㛗᪥ࡢୗ࡛ࡣኤ᪉࡟㌿෗⏘≀ࡢ⵳✚㔞ࡀࣆ࣮ࢡࢆ㏄࠼ࡿࡀ㸪ࡇࡢㄪ⠇࡟㛵ࢃࡿศᏊࡀ FKF1 ࡜ GI ࡛࠶ࡿ㸦Suárez-López etal. 2001, Imaizumi et al. 2003㸧
ࠋFKF1 ࡣ㸪F-box ࡜ࡼࡤࢀࡿ E3 ࣘࣅ࢟ࢳ࣮ࣥࣜ࢞ࢮ」ྜయࡢᙧᡂ࡟㛵ࢃࡿࢻ࣓࢖ࣥࢆࡶࡕ㸪ᶆⓗࢱࣥࣃࢡ㉁ࢆࣉࣟࢸ࢔ࢯ࣮࣒࡟ࡼࡿศゎ࡟ᑟࡃࢱࣥࣃࢡ㉁ࡢ୍✀࡛࠶ࡿ
㸦Nelson et al. 2000, Imaizumi et al. 2003㸧
ࠋࡲࡓ N ᮎ➃ഃ࡟ LOV (Light, Oxgen, and Voltage sensing)࡜ࡼࡤ
ࢀࡿࢻ࣓࢖ࣥࢆࡶࡕ㸪ⓎⰍᅋ࡜ࡋ࡚ࣇࣛࣅ࣭ࣥࣔࣀࢾࢡࣞ࢜ࢳࢻࢆ㠀ඹ᭷⤖ྜࡋ࡚࠾ࡾ㸪㟷Ⰽග࡟ᑐࡍࡿឤཷᛶࢆ♧ࡍ㸦Nelson et al. 2000, Imaizumi et al. 2003㸧
ࠋࡉࡽ࡟ C ᮎ➃ഃ࡟ࡣ kelch ࣜࣆ࣮ࢺ࡜ࡼࡤ
ࢀࡿࢻ࣓࢖ࣥࢆࡶࡕ㸪ࡇࡢ㒊ศࢆ௓ࡋ࡚㌿෗ᢚไᅉᏊ CYCLING DOF FACTOR 1 (CDF1)࡜┦஫స⏝ࡋ㸪ࡑࡢᏳᐃᛶࢆไᚚࡋ࡚࠸ࡿ㸦Imaizumi et al. 2005㸧
ࠋCDF1 ࡣ CO 㑇ఏᏊࡢࣉ࣮ࣟࣔࢱ࣮࡟⤖ྜࡍࡿࡇ࡜
࡛ࡑࡢⓎ⌧ࢆᢚไࡍࡿാࡁࢆࡶࡘ㸦Imaizumi et al. 2005㸧
ࠋCDF1 ࡣࢩࣟ࢖ࢾࢼࢬࢼ࡟ᑡ࡞ࡃ࡜ࡶ 3 ࡘࡢ࣍ࣔࣟࢢࡀᏑᅾࡋ㸪ࡇࢀࡽࡶ CO ࡢⓎ⌧ㄪ⠇࡟㛵ࢃࡗ࡚࠸ࡿ㸦Fornara et al. 2009㸧ࠋ୍᪉㸪GI ࡣ㝣ୖ᳜≀࡛ࡢࡳぢࡽࢀࡿศᏊ㔞⣙ 130 kDa ࡢࢱࣥࣃࢡ㉁࡛࠶ࡿ㸦Fowler et al. 1999, Park et al. 1999㸧
ࠋFKF1 ࡜GI ࡣ㟷Ⰽගࡢୗ࡛Ᏻᐃ࡞ࢱࣥࣃࢡ㉁」ྜయࢆᙧᡂࡋ㸪CDF ࢱࣥࣃࢡ㉁ࢆศゎࡍࡿࡇ࡜࡛ CO ࡢ㌿෗ࢆάᛶ໬ࡍࡿാࡁࢆࡶࡘ㸦Sawa et al. 2007, Fornara et al. 2009㸧
ࠋFKF1 ࡜ GI ࡢ㑇ఏᏊⓎ⌧ࡣᴫ᪥᫬ィ࡟ไᚚࡉࢀ࡚࠾ࡾ㸦Fowler et al. 1999, Park et al. 1999, Nelson et al. 2000㸧㸪㛗᪥ୗ࡛ࡣ FKF1 ࡜ GI ࡢ㑇ఏᏊⓎ⌧ࡢࣆ࣮ࢡࡀኤ᪉࡟୍⮴ࡍࡿࠋࡉࡽ࡟ࡇࡢ࡜ࡁ㟷Ⰽග࡟ࡼࡾ FKF1-GI 」ྜయࡀࡼࡾᏳᐃ໬ࡉࢀࡿࠋࡇ
࠺ࡋࡓࡇ࡜ࡀ༠ㄪⓗ࡟ຠࡃࡇ࡜࡛㸪ኤ᪉࡟ FKF1-GI 」ྜయ࡟ࡼࡿ CDF ࡢศゎࡀ᭱ࡶຠ⋡ⓗ࡟⾜ࢃࢀ㸪CO ࡢ㌿෗άᛶ໬ࡀ㉳ࡇࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ㸦Sawa et al. 2007㸧ࠋࡉࡽ࡟ FKF1 ࡣ⩻ヂᚋㄪ⠇࡟ࡶ㛵ࢃࡗ࡚࠾ࡾ㸪CO ࢱࣥࣃࢡ㉁࡜┤᥋┦஫స⏝ࡋ㸪CO ࢆᏳᐃ໬ࡉࡏࡿࡇ࡜ࡀ᫂ࡽ࠿࡟ࡉࢀ࡚࠸ࡿ㸦Song et al.
2012㸧
ࠋFT ࡢ㌿෗⮬యࡶ FKF1 ࡜ GI ࡟ࡼࡿㄪ⠇ࢆཷࡅ࡚࠸ࡿࠋFT 㑇ఏᏊࡢࣉ࣮ࣟࣔࢱ࣮࡟ࡣ CO ࡜ྠᵝ࡟ CDF ࢱࣥࣃࢡ㉁ࡀ⤖ྜࡋ㸪FKF1-GI 」ྜయ࡟ࡼࡿㄪ⠇ࢆཷࡅࡿ㸦Song et al. 2012㸧ࠋࡲࡓ㸪GIࡣูࡢ」ᩘࡢ㌿෗ᢚไᅉᏊ࡜┦஫స⏝ࡍࡿࡇ࡜࡛ FT ࡢ㌿෗ᢚไ࡟㛵ࢃࡗ࡚࠸ࡿࡇ࡜ࡶ᫂ࡽ࠿࡟ࡉࢀ
࡚࠸ࡿ㸦Sawa and Kay 2011㸧ࠋ㸱㸬ⰼⱆᙧᡂ࡜ᴫ᪥᫬ィࡢไᚚ࡟㛵ࢃࡿ FKF1/ZTL/LKP2 㑇ఏᏊࣇ࢓࣑࣮ࣜ ࢩࣟ࢖ࢾࢼࢬࢼ࡟ࡣ ZEITLUPE (ZTL)㸪LOV KELCH PROTEIN 2 (LKP2)࡜࠸࠺㸰ࡘࡢ FKF1 ┦ྠ㑇ఏᏊࡀᏑᅾࡋ㸪ࡇࢀࡽࡣ㑇ఏᏊࣇ࢓࣑࣮ࣜࢆᙧᡂࡋ࡚࠸ࡿ㸦ᅗ 1㸧
ࠋZTL ࡜ LKP2 ࡢᶵ⬟ࡣ FKF1 ࡜␗࡞ࡾ㸪୺࡟ᴫ᪥᫬ィࡢไᚚ࡟㛵ࢃࡗ࡚࠸ࡿࠋZTL ࡣ㸪ኪ㛫࡟࠾࠸࡚ᴫ᪥ࣜࢬ࣒ࡢ୰ᚰ᣺ືᏊ࡛࠶ࡿ
TIMING OF CAB EXPRESSION 1 (TOC1)࡜PSEUDO RESPONSE REGULATOR 5 (PRR5)࡜ࡢ㛫࡛LOV

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植物科学最前線 7:80 (2016)ࢻ࣓࢖ࣥࢆ௓ࡋ࡚┦஫స⏝ࡋ㸪ࡇࢀࡽࢆࣉࣟࢸ࢔ࢯ࣮࣒࡟ࡼࡿศゎ࡟ᑟࡃ㸦Más et al. 2003, Yasuhara et al.
2004, Kiba et al. 2007, Fujiwara et al. 2008, Baudry et al. 2010㸧ࠋ୍᪉㸪᪥୰࡟࠾࠸࡚ࡣ㸪ZTL ࡣ FKF㸯࡜ྠᵝ࡟ LOV ࢻ࣓࢖ࣥࢆ௓ࡋ࡚ GI ࡜⤖ྜࡋ㸪㟷Ⰽගࡢୗ࡛Ᏻᐃ࡞ࢱࣥࣃࢡ㉁」ྜయࢆᙧᡂࡍࡿࠋGI ࡢ⵳✚ࡣᴫ᪥ࣜࢬ࣒ࢆ♧ࡍࡓࡵ㸪ZTL-GI 」ྜయࡢᙧᡂࡣ༗ᚋ࡟࠶ࡓࡿ᫬㛫࡟ࣆ࣮ࢡࢆ㏄࠼ࡿ㸦Kim et al.
2007㸧ࠋࡇࡢ࡜ࡁᣕᢠⓗ࡟TOC1 ࡜PRR5࡜ࡢ⤖ྜࡀ㜼ᐖࡉࢀࡿࡓࡵ㸪ࡇࢀࡽࡢࢱࣥࣃࢡ㉁ࡣᏳᐃ໬ࡋ㸪ᴫ᪥ࣜࢬ࣒ࡀࡼࡾ᫂░࡟้ࡲࢀࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ㸦Fujiwara et al. 2008㸧
ࠋLKP2 ࡣ ZTL ࡜ẚ࡭࡚ࢩࣟ࢖ࢾࢼࢬࢼ᳜≀య࡛ࡢⓎ⌧㔞ࡀᴟࡵ࡚ప࠸ࡓࡵ㸪ࡑࡢኚ␗ࡔࡅ࡛ࡣᴫ᪥ࣜࢬ࣒࡟␗ᖖࡀぢࡽࢀ࡞࠸㸦Schultz et al. 2001, Baudry et al. 2010㸧ࠋࡋ࠿ࡋ㸪㐣๫Ⓨ⌧ࡉࡏࡿ࡜ᴫ᪥ࣜࢬ࣒ࡀ୙ᩚ࡜࡞ࡾ㸪ZTL ࣉ࣮ࣟࣔࢱ࣮ไᚚୗ࡛Ⓨ⌧ࡉࡏࡿ࡜ ztl ኚ␗ࢆ┦⿵࡛ࡁࡿࡇ࡜࠿ࡽ㸪LKP2 ࡣ ZTL ࡜࡯ࡰྠᵝࡢศᏊᶵ⬟ࢆࡶࡘ࡜⪃࠼ࡽࢀ࡚࠸ࡿ㸦Schultz et al. 2001, Baudry et al. 2010㸧
ࠋ FKF1ࡶࡲࡓ㸪ᴫ᪥᫬ィࡢไᚚ࡟㛵୚ࡋ࡚࠸ࡿࠋ
fkf1ኚ␗ࡔࡅ࡛ࡣᴫ᪥ࣜࢬ࣒࡟ᙳ㡪ࡀぢࡽࢀ࡞࠸ࡀ㸪ztl fkf1 ஧㔜ኚ␗ᰴ࡛ࡣ ztl ኚ␗ᰴ࡜ẚ࡭࡚ኪ㛫࡛ࡢ TOC1 ࡜ PRR5 ࡢศゎࡀࡼࡾᙉࡃ㜼ᐖࡉࢀ㸪ᴫ᪥ࣜࢬ࣒ࡣࡼࡾ㛗࿘ᮇ໬ࡍࡿ㸦Baudry et al. 2010㸧ࠋࡋ࠿ࡋ㸪FKF1 ࢆ ZTL ࣉ࣮ࣟࣔࢱ࣮ไᚚୗ࡛Ⓨ⌧ࡉࡏ࡚ࡶ ztl ኚ␗࡟ࡼࡿᴫ᪥ࣜࢬ࣒␗ᖖࢆ┦⿵࡛ࡁ࡞࠸ࡇ࡜࠿ࡽ㸪ᴫ᪥᫬ィࡢไᚚ࡟࠾ࡅࡿ FKF1 ࡢస⏝ᶵ
ᗎࡣ ZTL࣭LKP2 ࡜␗࡞ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ㸦Baudry et al. 2010㸧ࠋ ୍᪉࡛㸪ZTL ࡜ LKP2 ࡣⰼⱆᙧᡂ࡟ࡶ㛵ࢃࡗ࡚࠸ࡿࠋztl ኚ␗ᰴࡣ▷᪥᮲௳ୗ࡛ᙅ࠸᪩ဏࡁࡢ⾲⌧ᆺࢆ♧ࡋ㸪lkp2 ࡜ࡢ஧㔜ኚ␗࡛ࡼࡾ᪩ဏࡁ࡜࡞ࡾ㸪CO ࡢⓎ⌧ࡣୖ᪼ࡍࡿ㸦Somers et al. 2004, Takase et al.
2011㸧ࠋࡋ࠿ࡋ㸪fkf1 ztl lkp2 ୕㔜ኚ␗ᰴ࡛ࡣࡇࡢ⾲⌧ᆺࡀᢚไࡉࢀ㸪CO ࡢⓎ⌧ࡶᢚไࡉࢀࡿ㸦Fornara etal. 2009, Takase et al. 2011㸧ࠋࡲࡓ㸪ZTL ࡜ LKP2 ࢆࡑࢀࡒࢀ㐣๫Ⓨ⌧ࡉࡏࡿ࡜㸪CO ࡢⓎ⌧ࡀᢚไࡉࢀ㸪
ᙧ㉁㌿᥮ᰴࡣ㛗᪥᮲௳ୗ࡛㐜ဏࡁ࡜࡞ࡿ㸦Schultz et al. 2001, Somers et al. 2004㸧ࠋࡇࡢ⌧㇟ࡢㄝ᫂ࡢࡦ࡜
ࡘ࡜ࡋ࡚㸪FKF1 ࡜ ZTL࣭LKP2 ࡢ⣽⬊ෆᒁᅾࡢ㐪࠸ࡀᣲࡆࡽࢀࡿࠋFKF1 ࡣ᰾࡛ᶵ⬟ࡍࡿࡢ࡟ᑐࡋ㸪
ZTL ࡜ LKP2 ࡣ୺࡟⣽⬊㉁࡟ᒁᅾࡍࡿ
㸦Kim et al. 2007, Sawa et al. 2007, Takase et al. 2011, Kim et al. 2013㸧
ࠋ
LOV ࢻ࣓࢖ࣥࢆྵࡴ ZTL ࢱࣥࣃࢡ㉁ࡢ N ᮎ➃ഃࡣ GI ࡜ࡢ⤖ྜ⬟ࢆࡶࡘࡀ㸪ࡇࡢ㒊ศࡔࡅࢆ㐣๫Ⓨ⌧ࡉࡏࡿ࡜㸪GI ࡢ᰾ᒁᅾࡀ㜼ᐖࡉࢀ㸪ᙧ㉁㌿᥮ᰴࡀ㐜ဏࡁ࡜࡞ࡿ㸦Kim et al. 2013㸧ࠋࡇࡢࡇ࡜࠿ࡽ㸪FKF1
࡜ ZTL࣭LKP2 ࡣ GI ࢆࠕඹ᭷ࠖࡋ࡚࠾ࡾ㸪ࡇࢀࡽࡀṇᖖ࡟ᶵ⬟ࡍࡿࡓࡵ࡟ࡣ㸪᰾࡜⣽⬊㉁࡛ࡢ GI ࡢศᏊᩘࡢࣂࣛࣥࢫࡀ㔜せ࡛࠶ࡿ࡜⪃࠼ࡽࢀࡿࠋZTL ࡢ N ᮎ➃㒊ศࡢ㐣๫Ⓨ⌧ᰴ࡛ࡣ㸪GI ࡀ⣽⬊㉁࡛
ࡼࡾከࡃ౑ࢃࢀ࡚ࡋࡲ࠺ࡓࡵ࡟㸪᰾ෆࡢ GI ࡢศᏊᩘࡀῶࡾ㸪ࡑࡢ⤖ᯝ FKF1 ࡢᶵ⬟ࡀᢚไࡉࢀ࡚㐜ဏࡁ࡟࡞ࡿ࡜⪃࠼ࡽࢀࡿ㸦Kim et al. 2013㸧
ࠋFKF1 ࡜ ZTL࣭LKP2 ࡢస⏝ᶵᗎ࡟ࡘ࠸࡚ࡣ௚ࡢ௬ㄝࡶᥦၐ
ࡉࢀ࡚࠾ࡾ㸪௒ᚋࡢ᳨ドࡀᚅࡓࢀࡿ㸦Ito et al. 2012, Suetsugu and Wada 2013㸧
ࠋ

㸲㸬FKF1/ZTL/LKP2 ࡢ♽ඛᆺࢱࣥࣃࢡ㉁ࡣⱏ㢮ࢮࢽࢦࢣࡢᡂ㛗┦㌿᥮࡟㛵ࢃࡿ
 FKF1/ZTL/LKP2 ࡜ GI ࡣ㸪࡜ࡶ࡟㝣ୖ᳜≀㐍໬ࡢึᮇ࠿ࡽᏑᅾࡋ࡚࠸ࡓ࡜⪃࠼ࡽࢀࡿࠋࢩࢲ᳜≀ࡢ࠺ࡕ᭱ࡶྂ࠸ศ㢮⩌࡟ᒓࡍࡿᑠⴥ㢮࢖ࢾ࢝ࢱࣄࣂ Selaginella moellendorffii ࡛ࡣ㸪FKF1/ZTL/LKP2 ࡜ GIࡢ┦ྠ㑇ఏᏊࡀࡑࢀࡒࢀ㸯ࡘࡎࡘᏑᅾࡍࡿ㸦Holm et al. 2010, Banks et al. 2011㸧ࠋࡋ࠿ࡋࢥࢣ᳜≀ࡢ࠺ࡕ㸪⹊㢮ࣄ࣓ࢶࣜ࢞ࢿࢦࢣ Physcomitrella patens ࡛ࡣ㸪LOV ࢻ࣓࢖ࣥࡔࡅࢆࡶࡘࢱࣥࣃࢡ㉁࡜㸪F-box

S. Yamaoka & T. Kohchi-3
BSJ-Review 7:80 (2016)

植物科学最前線 7:81 (2016)ᅗ㸯. FKF1/ZTL/LKP2 ࡢ㑇ఏᏊ࣭ࢱࣥࣃࢡ㉁ࡢᵓ㐀࡜⣔⤫ᶞ
A㸪ࢮࢽࢦࢣ࠾ࡼࡧࢩࣟ࢖ࢾࢼࢬࢼࡢ GI ࠾ࡼࡧ FKF1 ࣍ࣔࣟࢢࡢ㑇ఏᏊᵓ㐀ࠋ㯮ࡢ࣎ࢵࢡࢫࡣ࢚࢟ࢯࣥ㸪⅊Ⰽࡢ࣎ࢵࢡࢫࡣ㠀⩻ヂ㡿ᇦࢆ♧ࡍࠋᅗࡣ Kubota et al. (2014)ࡼࡾᨵኚࠋB㸪ࢮࢽࢦࢣ࠾ࡼࡧࢩࣟ࢖ࢾࢼࢬࢼࡢ
FKF1/ZTL/LKP2 ࢱࣥࣃࢡ㉁ࡢᵓ㐀ࠋLOV㸪LOV ࢻ࣓࢖ࣥ㸪kelch repeats㸪kelch ࣜࣆ࣮ࢺࠋC㸪FKF1/ZTL/LKP2
ࢱࣥࣃࢡ㉁ࡢ LOV ࢻ࣓࢖ࣥࡢ࢔࣑ࣀ㓟㓄ิࠋ▮ᑼࡣⓎⰍᅋࡢ㠀ඹ᭷⤖ྜ࡟ᚲせ࡞ࢩࢫࢸ࢖ࣥṧᇶࢆ♧ࡍࠋD㸪
FKF1/ZTL/LKP2 ࢱࣥࣃࢡ㉁ࡢ⣔⤫ᶞࠋMp㸸ࢮࢽࢦࢣ㸦Marchantia polymorpha㸧㸪Sm㸸࢖ࢾ࢝ࢱࣄࣂ㸦Selaginella
moellendorffii㸧㸪Ac㸸ࢱ࣐ࢿࢠ㸦Allium cepa㸧㸪Os㸸࢖ࢿ㸦Oryza sativa㸧㸪Ta㸸ࢥ࣒ࢠ㸦Triticum aestivum㸧㸪Gm㸸ࢲ࢖ࢬ㸦Glycine max㸧㸪At㸸ࢩࣟ࢖ࢾࢼࢬࢼ㸦Arabidopsis thaliana㸧㸪Mc㸸࢔࢖ࢫࣉࣛࣥࢺ㸦ࣁ࣐࣑ࢬࢼ⛉Mesembryanthemum crystallinum㸧ࠋᅗࡣ Kubota et al. (2014)ࡼࡾᨵኚࠋ

࡜ kelch ࣜࣆ࣮ࢺࢆྵࡴ㒊ศ㓄ิࢆࡶࡘࢱࣥࣃࢡ㉁ࡢ㑇ఏᏊࡣᏑᅾࡍࡿࡀ㸪FKF1/ZTL/LKP2 ࡢ┦ྠ㑇ఏᏊࡣᏑᅾࡋ࡞࠸ࠋࡲࡓ GI ࡢ┦ྠ㑇ఏᏊࡣぢࡘ࠿ࡗ࡚࠸࡞࠸㸦Holm et al. 2010㸧ࠋࡉࡽ࡟㸪Ỉ⏕ࡢ༢⣽⬊⥳⸴ Chlamydomonas reinhardtii ࡜ Ostreococcus tauri ࡛ࡣ FKF1/ZTL/LKP2࣭GI ࡢ┦ྠ㑇ఏᏊࡣ࠸ࡎࢀࡶぢࡘ࠿ࡗ࡚࠸࡞࠸㸦Mittag et al. 2005, Matsuo et al. 2008, Corellou et al. 2009㸧ࠋࡇࡢࡇ࡜࠿ࡽ㸪ᚑ᮶ࡣ
FKF1/ZTL/LKP2࣭GI ࡟ࡼࡿᡂ㛗┦㌿᥮ࡢ࣓࢝ࢽࢬ࣒ࡣ⥔⟶᮰᳜≀ࡢㄌ⏕௨㝆࡟⋓ᚓࡉࢀࡓࡶࡢ࡜⪃࠼ࡽࢀ࡚࠸ࡓࠋ ⱏ㢮ࢮࢽࢦࢣ Marchantia polymorpha ࡣ㝣ୖ᳜≀㐍໬ࡢ᭱ࡶᇶ㒊࡛ศᒱࡋࡓࢥࢣ᳜≀ࡢ୍✀࡛࠶ࡿ㸦Qiu et al. 2006, Chang and Graham 2011, Wickett et al. 2014㸧ࠋ᭱㏆ࢮࢽࢦࢣࡣศᏊ㑇ఏᏛࡢᇶ┙ࡀᩚഛࡉࢀࠊไᚚ⣔㑇ఏᏊࡢ㐍໬࡜ከᵝᛶࠊ࠶ࡿ࠸ࡣ᳜≀ࡢ㝣ୖ໬ࢆࡶࡓࡽࡋࡓ࣓࢝ࢽࢬ࣒ࢆ◊✲ࡍࡿୖ࡛ඃ
ࢀࡓࣔࢹ᳜ࣝ≀࡜ࡋ࡚ὀ┠ࡉࢀ࡚࠸ࡿ㸦Ishizaki et al. 2016, Berger et al. 2016, Bowman et al. 2007㸧ࠋࢮࢽࢦࢣࡣ㛗᪥᳜≀࡛࠶ࡾ㸪ⴥ≧యࡢඛ➃㒊࡟ച≧ࡢ⏕Ṫჾᡸࢆᙧᡂࡍࡿࡇ࡜࡛ᡂ㛗┦㌿᥮ࢆ⾜࠺㸦Benson-Evans 1961, Benson-Evans 1964㸧ࠋࡇࡢ┦㌿᥮࡟ࡣ㐲㉥Ⰽග࡟ᐩࡴග㉁ࡀᚲせ࡛࠶ࡿ㸦Chiyoda
et al. 2008, Ἑෆ࣭▼ᓮ 2012㸧ࠋ➹⪅ࡽࡢࢢ࣮ࣝࣉࡣ㸪ࢮࢽࢦࢣࡢࢤࣀ࣒࡟ FKF1/ZTL/LKP2 ࡜ GI ࡢ┦ྠ㑇ఏᏊࡀࡑࢀࡒࢀ㸯ಶࡎࡘᏑᅾࡍࡿࡇ࡜ࢆぢฟࡋ㸪ࡇࢀࡽࢆࡑࢀࡒࢀ MpFKF ࡜ MpGI ࡜ྡ௜ࡅࡓ
㸦Kubota et al. 2014㸧ࠋࢩࣟ࢖ࢾࢼࢬࢼࡢ FKF1 ࡜ GI ࡢ㑇ఏᏊ࡜ࡑࡢᵓ㐀ࢆẚ㍑ࡍࡿ࡜㸪࢖ࣥࢺࣟࣥࡢ
ᤄධ఩⨨ࡀࡼࡃ୍⮴ࡋ࡚࠾ࡾ㸪ࡇࢀࡽࡣ㐍໬ⓗ࡟ಖᏑࡉࢀ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡓ㸦ᅗ 1A㸧
ࠋMpFKF ࡣ㸪௚ࡢ FKF1/ZTL/LKP2 ࢱࣥࣃࢡ㉁࡜ྠᵝ㸪LOV ࢻ࣓࢖࣭ࣥF-box࣭kelch ࣜࣆ࣮ࢺࢆࡶࡗ࡚࠸ࡓ㸦ᅗ 1B㸧
ࠋࡲࡓ LOV ࢻ࣓࢖࡛ࣥࡣⓎⰍᅋࡢ㠀ඹ᭷⤖ྜ࡟ᚲせ࡞ࢩࢫࢸ࢖ࣥṧᇶࡀಖᏑࡉࢀ࡚࠸ࡓ㸦ᅗ 1C㸧
ࠋ⣔⤫ゎᯒ࡟ࡼࡿ࡜㸪MpFKF ࡣ࢖ࢾ࢝ࢱࣄࣂࡢ┦ྠ㑇ఏᏊ(SmFKF)࡜᭱ࡶ㏆⦕࡛࠶ࡾ㸪⿕Ꮚ᳜≀ࡢ FKF1㸪
ZTL/LKP2 ࡢࢡ࣮ࣞࢻ࠿ࡽࡣ⊂❧ࡋ࡚࠸ࡓ㸦ᅗ 1D㸧
ࠋࡇࡢࡇ࡜࠿ࡽ㸪FKF1/ZTL/LKP2 ࢱࣥࣃࢡ㉁ࡣ㸪
MpFKF ࢆࡣࡌࡵ࡜ࡍࡿࢥࢣ࣭ࢩࢲ᳜≀ࡢࡶࡢࢆ♽ඛ࡜ࡋ㸪᳜≀ࡢ㐍໬ࡀ㐍ࡴ࡟ࡘࢀ㸪㑇ఏᏊ㔜」ࡢᚋ㸪 

S. Yamaoka & T. Kohchi-4
BSJ-Review 7:81 (2016)

植物科学最前線 7:82 (2016)ᅗ㸰. ࢮࢽࢦࢣ࡜ࢩࣟ࢖ࢾࢼࢬࢼࡢ⏕ά⎔࡜ FKF1-GI 」ྜయࡢస⏝Ⅼn ࡣ༙ᩘయୡ௦ࢆ㸪2n ࡣ㸰ಸయୡ௦ࢆ♧ࡍࠋ

FKF1 ࡜ ZTL/LKP2 ࡟௦⾲ࡉࢀࡿ㸰✀㢮ࡢศᏊ࡟ᶵ⬟ⓗ࡟ศ໬ࡋ࡚࠸ࡗࡓ࡜⪃࠼ࡽࢀࡿ㸦Kubota et al.
2014㸧ࠋ ࢮࢽࢦࢣࡣ㸪㐲㉥Ⰽග࡟ᐩࡴගࡢୗ࡟࠾࠸࡚㸪㛗᪥᮲௳࡛ࡣ⏕Ṫჾᡸࢆᙧᡂࡋ࡚ᡂ㛗┦㌿᥮ࡍࡿࡀ㸪▷᪥᮲௳࡛ࡣⴥ≧యࡢࡲࡲᡂ㛗ࡋ⥆ࡅࡿࠋMpFKF ࡜ MpGI ࡢ㑇ఏᏊࣀࢵࢡ࢔࢘ࢺᰴࡣ㸪࡝ࡕࡽࡶ㛗᪥᮲௳ୗ࡛⏕Ṫჾᡸࡢᙧᡂࡀࡳࡽࢀ࡞࠿ࡗࡓࠋࡲࡓࡑࢀࡒࢀࡢ㐣๫Ⓨ⌧ᰴ࡛ࡣ㸪▷᪥᮲௳ୗ࡛ࡶ㸪㛗᪥᮲௳ୗ࡜࡯࡜ࢇ࡝ྠࡌ᫬ᮇ࡟⏕Ṫჾᡸࢆᙧᡂࡋࡓࠋࡉࡽ࡟㸪MpFKF ࡜ MpGI ࢆྠ᫬࡟㐣๫Ⓨ⌧ࡉࡏࡓᰴ࡛ࡣ㸪༢⊂ࡢ㐣๫Ⓨ⌧ᰴ࡜࡯࡜ࢇ࡝ྠࡌ᫬ᮇ࡟⏕Ṫჾᡸࢆᙧᡂࡋࡓࠋࡲࡓ MpFKF ࡜ MpGI ࡢ
ࢱࣥࣃࢡ㉁ࡣ㸪ᑡ࡞ࡃ࡜ࡶ㛗᪥᮲௳ࡢ᫂ᮇ࡛ࡣ┦஫స⏝ࡋ࠺ࡿࡇ࡜ࡶ᫂ࡽ࠿࡟࡞ࡗࡓ㸦Kubota et al.
2014㸧ࠋࡇࢀࡽࡢࡇ࡜࠿ࡽ㸪MpFKF ࡜ MpGI ࡣࢮࢽࢦࢣࡢᡂ㛗┦㌿᥮࡟ᚲせ࡞ศᏊ࡛࠶ࡾ㸪」ྜయࢆ
ᙧᡂࡋ࡚ಁ㐍ⓗ࡟ാࡃ࡜⪃࠼ࡽࢀࡿࠋ⣔⤫ゎᯒ࡜࠶ࢃࡏ࡚⪃࠼ࡿ࡜㸪ࡇࡢ MpFKF-MpGI 」ྜయ࡟ࡼ
ࡿไᚚࡀ㸪㝣ୖ᳜≀ࡢᡂ㛗┦㌿᥮ࡢ࣓࢝ࢽࢬ࣒ࡢཎᙧ࡛࠶ࡿ࡜⪃࠼ࡽࢀࡿ㸦Kubota et al. 2014㸧ࠋ
MpFKF-MpGI 」ྜయࡀୗὶࡢᡂ㛗┦㌿᥮ࡢᐇ⾜ᅉᏊࢆ┤᥋ไᚚࡋ࡚࠸ࡿࡢ࠿㸪࠶ࡿ࠸ࡣࢩࣟ࢖ࢾࢼࢬࢼ ZTL-GI 」ྜయࡢࡼ࠺࡟ᴫ᪥᫬ィࡢไᚚࢆ௓ࡋ࡚ᡂ㛗┦㌿᥮࡟㛵ࢃࡗ࡚࠸ࡿࡢ࠿㸪௒ᚋࡢ◊✲ࡢ㐍ᒎࡀᮇᚅࡉࢀࡿࠋ

  MpFKF ࡜ MpGI ࡢ⏕⌮ᶵ⬟࡟ࡘ࠸࡚ࡢ▱ぢࡣ㸪᳜≀ࡢ⏕ά⎔ࡢ㐍໬ࢆ⪃࠼ࡿୖ࡛ࡶ♧၀࡟ᐩࢇ࡛࠸ࡿࠋᅗ 2 ࡟♧ࡍࡼ࠺࡟㸪ࢮࢽࢦࢣࡢ⏕ά⎔ࡢ኱㒊ศࡣ㸪༙ᩘయ(n)ࡢ㓄അయୡ௦ࡀ༨ࡵࡿࠋ⏕Ṫჾᡸࡢ୰࡛ཷ⢭࡟ࡼࡾ⬇ࡀ⏕ࡌ㸪ࡑࢀࡀⓎ㐩ࡋ࡚⬊Ꮚయ࡜࡞ࡿࡀ㸪ࡇࡢ㛫ࡢ⤌⧊ࡔࡅࡀ㸰ಸయ(2n)ࡢୡ
௦࡛࠶ࡾ㸪ࡑࡢࢧ࢖ࢬ࣭Ꮡ⥆ࡢᮇ㛫࡜ࡶ࡟ᑠࡉ࠸ࠋࡇࡢ㓄അయୡ௦—⬊Ꮚయୡ௦ࡢ㛵ಀࡣ㸪㝣ୖ᳜≀ࡢ㐍໬ࡀ㐍ࡴ࡟ࡘࢀᚎࠎ࡟㏫㌿ࡍࡿࠋࢩࣟ࢖ࢾࢼࢬࢼ࡞࡝ࡢ⿕Ꮚ᳜≀࡛ࡣ㸪⏕ά⎔ࡢ኱༙ࡣ⬊Ꮚయୡ௦ࡀ༨ࡵࡿࠋ㓄അయୡ௦ࡣ㸪ⰼჾᐁࡢ୰࡛ᙧᡂࡉࢀࡿⰼ⢊࡜⬇ࡢ࠺࡜࠸࠺ᑠࡉ࡞⤌⧊ࡢࡳ࡛࠶ࡾ㸪ࡑࢀࡣཷ⢭࡜࡜ࡶ࡟⤊ࢃࡗ࡚ࡋࡲ࠺ࠋFKF1/ZTL/LKP2 ࡜ GI ࡟ࡼࡿᡂ㛗┦㌿᥮ࡀ⾜ࢃࢀࡿ᫬ᮇࡣ㸪ࢮࢽࢦࢣ࡛ࡣ㓄അయୡ௦࡛࠶ࡾ㸪⿕Ꮚ᳜≀࡛ࡣ⬊Ꮚయୡ௦࡛࠶ࡿ㸦ᅗ 2㸧ࠋࡇࡢࡇ࡜ࡣ㸪㐍໬ࡢ㐣⛬࡛㸪ᮏ᮶ࡣ㓄അయୡ௦࡛ᶵ⬟ࡋ࡚࠸ࡓᡂ㛗┦㌿᥮ࡢ࣓࢝ࢽࢬ࣒ࡀ㸪⬊Ꮚయୡ௦࡛ࡢไᚚ࡟㌿⏝ࡉࢀࡓࡇ࡜ࢆ
♧ࡋ࡚࠸ࡿ㸦Kubota et al. 2014㸧
ࠋ

S. Yamaoka & T. Kohchi-5
BSJ-Review 7:82 (2016)

植物科学最前線 7:83 (2016) ࢮࢽࢦࢣ࡛ࡣ㸪࡝ࡢࡼ࠺࡞ศᏊࡀ MpFKF-MpGI ࡢୗὶ࡛ാࡁ㸪ᡂ㛗┦㌿᥮ࢆᐇ⾜ࡍࡿࡢࡔࢁ࠺࠿ࠋ
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ࡀಁ㐍ࡉࢀࡿࡇ࡜࠿ࡽ㸪ᡂ㛗┦㌿᥮࡟㛵ࢃࡿࡇ࡜ࡀ♧ࡉࢀ࡚࠸ࡿ㸦Ọᒣࡽ ᪥ᮏ᳜≀Ꮫ఍➨ 78 ᅇ኱఍2014, Ἑෆ࣭ᒣᒸࡽ ᮍⓎ⾲㸧ࠋࡲࡓ➹⪅ࡽࡣࡇࢀࡲ࡛࡟㸪㐲㉥Ⰽගࢆྵࡲ࡞࠸ගࡢୗ࡛㸪᪥㛗࡟㛵ࢃࡽࡎ⏕Ṫჾᡸࢆᖖ࡟ᙧᡂࡍࡿࡇ࡜ࡢ࡛ࡁࡿኚ␗ᰴࢆ༢㞳ࡋ࡚࠸ࡿ㸦Yamaoka et al. 2004㸧ࠋࡇࡢኚ␗ᰴࡢ
ཎᅉ㑇ఏᏊࢆྠᐃࡍࡿࡇ࡜࡛㸪ࢮࢽࢦࢣࡢᡂ㛗┦㌿᥮ࡢᐇ⾜ᅉᏊࡀ᫂ࡽ࠿࡟࡞ࡾ㸪᳜≀ࡢᡂ㛗┦㌿᥮ࡢ࣓࢝ࢽࢬ࣒ࡀ࡝ࡢࡼ࠺࡟㐍໬ࡋ࡚ࡁࡓ࠿㸪ࡑࡢ඲ᐜࡢゎ᫂࡟ࡘ࡞ࡀࡿࡇ࡜ࡀᮇᚅࡉࢀࡿࠋ

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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
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RIKEN Center for Sustainable Resource Science,
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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%

=7

´țŦǝ@7"7)), ƾǜĂƘ, ƗƑĂƘ(*33'´țƀ8ƜȦƀāé<=:Ś#ȟÈƘ(&)9 (Ƒ3=#) , =:)ȸ§ĪƜ×Ƙ') , ƜƘ') , (" #*, =3$÷)ȇȄ;,

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)

植物科学最前線 7:93 (2016)ƅ)ŎȘ´țƦ²şƅ)ůȿ$*, VNS nBwȪÿ*pOxƾǜūȀċǝÃ(Ä #ƾǜô©ɏ¾ĤŶĪ@¨ŏ# ÒǝĪɓ țűƾǜ(*ɓ Ƚè<6, 9;țűŦǝ@ɓ6<6(* èÃ(ĴĳĪ)<ƆŬÈƾǜô@£;¹ħǵ<%$*, pOxƾǜūȀċ%ƾǜô©ɏ¾Ĥ%7( VNS %ĭÜ

vW\WE_^)ŷ$¾

Ĥ=<ǀ4ȟÈ#)*ƻģ$<Ć$> )my`KhSP PpVNS )Ǻř :, VNS (9<țűƾǜºÈƍǭ)Ⱦşƅ) :³#(´ț<yJfXx$<%ŋ: %'!ɕXu et al. 2014ɖ
=*, =3$ȟÈƘ'ȸ§Īŋ$!RPşƅ)jEdEd%, ǇƸŖşƅ)ȤƸƾǜ, %7(×nBwȪÿ)®(9!#£:=# <%@ƳƘ(ƥȱǵ'ƠǶ%'!

XURFLMS;

 "=G

Q8:A)&63K?

 PpVNS Ȫÿ@UEgeXe$ȣÁƗƍ<%ƖİƘȤƸƾǜºÈĚȌ=<ɕXu etal. 2014ɖ% :7ŋ: '9 (, VNS ǟȑ)îœƘ'Ȓ¶ãÿŦǝ*Ⱦşƅ) ';ŉ ůȿ$ƣƲ=# ;, Ⱥ, VNS *ȟÈȣƩ$ȞǊ%țűƾǜºÈ(ȸȞĞÂ@ĵ!#%Ĭ¯=<Àɉ$ƿț;, ǰÿşƅ$*, ȤƸƾǜºÈ)RCȒ¶¾ĤWLx%
#, ƴ)vW\WE_^ VNS→ƴ)vW\WE_^ MYB→ȤƸƾǜºÈ)ŷ=
Ƕ¹=#ɕYamaguchi & Demura 2010; ¹ŕøȈ 2015; Nakano et al. 2015ɖǡÜŹ %(,my`KhSP PpVNS7 ȀċƘȣÁƗƍŌ)c WNpcxǺř)ǂŚ, UEgeXe
VNS )ŷȪÿ)szO%ǖ :=<Ȫÿ)÷, my`KhSP$7 PpVNS )ŷ(āé# <%ƥ=ɕXu et al. 2014ɖ
PpVNS ŷ$¨ā=# UEgeXeȤ
ƸƾǜºÈȸȞȪÿ)szOȪÿ%#*, ƴ)vW\WE_^%ǖ

:=#<MYB Ȓ¶ãÿ)szO, pOxƾǜū(ȸ?<UWaE paCZȪÿ XCP1 )szO,9-ƾǜôǵšįįº$<YWÖįȮƼȪÿ$< CesA )szO, nGipliEdÖįǁȏȮƼȪÿǔ'&Ù3=# <'? , VNS ǟ¡)ºÿŦǝ)4':, VNS→MYB % !śǀ@þ6, ȤƸƾǜºÈ$Ƕ:=< VNS (9<Ȓ¶¾ĤWLx)ø3 'ɃĜ, RPşƅ%ǇƸŖşƅ)ŎȘ´țƦ²)Ōƀ$ň(ƣƲ=# %ǖ :=< )ǂŚ*, țűƾǜȟÈ%ǹƀ :Ƕ#7Ɇĕ(ƥßƘ$!%)7, ¥+RPşƅƾǜô(*ǇƸŖşƅŨô(÷Ù3=<nGiĪuv$<Of *Ù3=#' ɕƪ(9!#*Of ǁȏ)ȷƅȋ(<nGȯ@£<%8, nGiĪƅȋô(ǪƬ# <%*º !# <; Ligrone et al. 2008ɖ
(7?:, PpVNS7 @my`KhSP$ȣÁƗƍ<%Of ƑÖį("'<nGfpliEdǁȏ)ȮƼȪÿǔ)Ɨƍ
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$*ƾǜô)Of ÈțűƾǜºÈpOx)ņ'ƼȣƩ%#ǀ4ȗ3=
, %ȣƩ@Ĭą<%$<"3;, ƍé)ǇƸŖşƅ(Ƕ:=<Ǵɂ'țűƾǜºÈȣƩ*, RPşƅ%ǇƸŖşƅ)ŎȘ´țƦ²(<ÌþëȾşƅƋģ# VNS→pOxƾǜūɗƾǜô©ɏ%

WLx@rW(, VNS ŷ)ºÿEr c÷ŢÈǴɂÈ<%$įƲ#%ǖ :=9 ɕäɝɖmy`KhSP*ŭĩ':ǜÿ¡(jEdEd@ºÈ' , ǜÿ¡$Ɨƍ< PpVNS4 n_cɋì)ȒȚƾǜºÈ(ȸ# <%*²(ș0ɕXu et al. 2014ɖRPşƅ7ƪ(9!#*ǜÿ¡)n_c)Ħì(jEdEd@ºÈ<)$, Ì¿Ƙ(*RPşƅ$*ȭ¬¡ǜÿ¡á?, VNS→ƾǜūɗƾǜô©ɏ%´ț)WLx¤?=# <%Ƕ'%$<ƍé)şƅȟÈ

ĂƘ'ǶŇ$*, RPşƅ)Ʀ²ºĒġ(ƍ=U]şƅ%ƪÿşƅ)ŎȘ´țƦ²(#, ǜÿ¡)ŲǆƘ'ƾǜóŮǝÃƋģ=, ǜÿ¡)øëÈ÷ŦǝÈÄȜ%ļż=# <ɕLigrone et al. 2012ɖ öÈ¢ÁƅȋƑƒ@Òǝ(, Of '&)ņ'ŨȆǁȏ)ȟÈ)Ý-ű%'<%*Ĭ¯(Ʌ' Of ÈŨô*țűƾǜ)èÃě
ę@ɎȐƘ(Ŋ, ȵȎɄ)űȕȚ@Òǝ(, Ñ)őƑU])ǎŜ/%ǐ!%ǖ :=# <ɕSperry 2003ɖ
 ǖ <%, ƍƑ)Ⱦşƅ)țűƾǜ)kHU~ *³#, Ŏ½)ȾşƅƋģ VNS Ŧǝ@çÓ%"", ȵ ȟÈ)ȣƩ$ŷºÿEr cƗĎ÷ŢÈ<%$Ƒ4¹=#7)$<, %ŃƏ<%$<

YU

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