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

Evolutionary insights into the molecular mechanism ofday-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໬ࡍࡿࡓࡵࡢ࣓࢝ࢽࢬ࣒ࢆ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࠺ࡕ࡛᭱ࡶ☜ᐇ࡟ண ࡍࡿࡇ࡜ࡀ࡛ࡁ㸪ࡑࢀ࡟ࡼࡾ᮶ࡓࡿᏘ⠇ࢆ▱ࡿࡇ࡜ࡀ࡛ࡁࡿ࠿ࡽ࡛࠶ࡿࠋࡇࡢ࣓࢝ࢽࢬ࣒ࡣ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࢖ࢾࢼࢬࢼ࡛ࡣ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㸧ࠖ࡟ࡼࡗ࡚ࡑࡢࢩࢢࢼࣝࡀఏ㐩ࡉࢀ㸪ⱼ㡬࡛ࡢⰼⱆᙧᡂࡀ㛤ጞࡉࢀࡿ࡜࠸࠺௬ㄝࡀᥦၐࡉࢀࡓ㸦Chailakhyan 1936,S. Yamaoka & T. Kohchi-1
BSJ-Review 7:78 (2016)

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, Jaegerand 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, Wiggeet 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࡜ࡼࡤࢀࡿࢻ࣓࢖ࣥ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㛫࡛LOVS. Yamaoka & T. Kohchi-2
BSJ-Review 7:79 (2016)

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
ࠋLOV 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/LKP2ࢱࣥࣃࢡ㉁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◊✲ࡍࡿୖ࡛ඃࢀࡓࣔࢹ᳜ࣝ≀࡜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Chiyodaet 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࢖ࣥࢺࣟࣥࡢᤄධ఩⨨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)ࡢୡ௦࡛࠶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

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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➹⪅ࡽࡢࢢ࣮ࣝࣉ࡟ࡼࡿࢤࣀ࣒࣭ࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒ࠿ࡽ㸪ࢮࢽࢦࢣ࡟ࡣᑡ࡞ࡃ࡜ࡶ CDF ࡢ┦ྠ㑇ఏᏊࡀᏑᅾࡍࡿࡇ࡜ࡀ᫂ࡽ࠿࡟ࡉࢀ࡚࠸ࡿࠋࡇࡢ㑇ఏᏊࡣ㸪ࣀࢵࢡ࢔࢘ࢺ࡟ࡼࡾ⏕Ṫჾᡸࡢᙧᡂࡀಁ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␗ᰴࡢཎᅉ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ㅰ㎡ᮏ✏࡛㏙࡭ࡓ➹⪅ࡽࡢ◊✲࡟ࡘ࠸࡚㸪ி㒔኱Ꮫ኱Ꮫ㝔⏕࿨⛉Ꮫ◊✲⛉ࡢஂಖ⏣ⲁ㸦⌧࣭࣡ࢩࣥࢺࣥ኱Ꮫ㸧࣭Ọᒣၨኴ㑻୧Ặ࡜㸪ᩥ㒊⛉Ꮫ┬࣭⛉Ꮫ◊✲㈝⿵ຓ㔠ࡢຓᡂ࡟ㅰពࢆ⾲ࡍࡿࠋᘬ⏝ᩥ⊩Abe, M., Kobayashi, Y., Yamamoto, S., Daimon, Y., Yamaguchi, A., Ikeda, Y., Ichinoki, H., Notaguchi, M., Goto,K., & Araki, T. 2005. FD, a bZIP protein mediating signals from the floral pathway integrator FT at the shootapex. Science 309: 1052-1056.
Banks, J.A., et al. 2011. The Selaginella genome identifies genetic changes associated with the evolution ofvascular plants. Science 332: 960-963.
Baudry, A., Ito, S., Song, Y.H., Strait, A.A., Kiba, T., Lu, S., Henriques, R., Pruneda-Paz, J.L., Chua, N.H., Tobin,E.M., Kay, S.A., & Imaizumi, T. 2010. F-box proteins FKF1 and LKP2 act in concert with ZEITLUPE tocontrol Arabidopsis clock progression. Plant Cell 22: 606-622.
Benson-Evans, K. 1961. Environmental factors and bryophytes. Nature 191: 255–260.
Benson-Evans, K. 1964. Physiology of the reproduction of bryophytes. Bryologist 67: 431–445.
Berger, F., Bowman, J.L., & Kohchi, T. 2016. Marchantia. Curr. Biol. 26: R186-R187.
Bowman, J.L., Floyd, S.K., & Sakakibara, K. 2007. Green genes-comparative genomics of the green branch of life.
Cell 129: 229-234.
Chailakhyan, M.K. 1936. New facts in support of the hormonal theory of plant development. C. R. (Dokl.) Acad.
Sci. USSR, 13: 79–83.
Chang, Y. & Graham, S.W. 2011. Inferring the higher-order phylogeny of mosses (Bryophyta) and relatives usinga large, multigene plastid data set. Am. J. Bot. 98: 839–849.
Chiyoda, S., Ishizaki, K., Kataoka, H., Yamato, K.T., & Kohchi, T. 2008. Direct transformation of the liverwortMarchantia polymorpha L. by particle bombardment using immature thalli developing from spores. Plant CellRep. 27: 1467-1473.
Corbesier, L., Vincent, C., Jang, S., Fornara, F., Fan, Q., Searle, I., Giakountis, A., Farrona, S., Gissot, L., Turnbull,C., & Coupland, G. 2007. FT protein movement contributes to long-distance signaling in floral induction ofArabidopsis. Science 316: 1030-1033.

S. Yamaoka & T. Kohchi-6
BSJ-Review 7:83 (2016)

植物科学最前線 7:84 (2016)

Corellou, F., Schwartz, C., Motta, J.P., Djouani-Tahri el, B., Sanchez, F., & Bouget, F.Y. 2009. Clocks in the green
lineage: comparative functional analysis of the circadian architecture of the picoeukaryote Ostreococcus. Plant
Cell 21: 3436-3449.
Edwards, D., & Kenrick, P. 2015. The early evolution of land plants, from fossils to genomics: a commentary onLang (1937) 'On the plant-remains from the Downtonian of England and Wales'. Phil. Trans. R. Soc. B 370:
20140343.
Fornara, F., Panigrahi, K.C., Gissot, L., Sauerbrunn, N., Rühl, M., Jarillo, J.A., & Coupland, G. 2009. ArabidopsisDOF transcription factors act redundantly to reduce CONSTANS expression and are essential for a
photoperiodic flowering response. Dev. Cell 17: 75-86.
Fowler, S., Lee, K., Onouchi, H., Samach, A., Richardson, K., Morris, B., Coupland, G., & Putterill, J. 1999.
GIGANTEA: a circadian clock-controlled gene that regulates photoperiodic flowering in Arabidopsis andencodes a protein with several possible membrane-spanning domains. EMBO J. 18: 4679-4688.
Fujiwara, S., Wang, L., Han, L., Suh, S.-S., Salomé, P.A., McClung, C.R., & Somers, D.E. 2008. Post-translational
regulation of the Arabidopsis circadian clock through selective proteolysis and phosphorylation ofpseudo-response regulator proteins. J. Biol. Chem. 283: 23073-23083.
Garner, W.W. & Allard, H.A. 1920. Effect of the relative length of day and night and other factors of theenvironment on growth and reproduction in plants. J. Agric. Res. 18: 553–606.
Holm, K., Kallman, T., Gyllenstrand, N., Hedman, H., & Lagercrantz, U. 2010. Does the core circadian clock inthe moss Physcomitrella patens (Bryophyta) comprise a single loop? BMC Plant Biol. 10: 109.
Imaizumi, T., Tran, H.G., Swartz, T.E., Briggs, W.R., & Kay, S.A. 2003. FKF1 is essential forphotoperiodic-specific light signalling in Arabidopsis. Nature 426: 302-306.
Imaizumi, T., Schultz, T.F., Harmon, F.G., Ho, L.A., & Kay, S.A. 2005. FKF1 F-box protein mediates cyclicdegradation of a repressor of CONSTANS in Arabidopsis. Science 309: 293-297.
Ishizaki, K., Nishihama, R., Yamato, K.T., & Kohchi, T. 2016. Molecular genetic tools and techniques forMarchantia polymorpha research. Plant Cell Physiol. 57: 262-270.
Ito, S., Song, Y.H., & Imaizumi, T. 2012. LOV domain-containing F-box proteins: light-dependent protein
degradation modules in Arabidopsis. Mol. Plant 5: 573-582.
Jaeger, K.E., & Wigge, P.A. 2007. FT protein acts as a long-range signal in Arabidopsis. Curr. Biol. 17: 1050-1054.
Kardailsky, I., Shukla, V.K., Ahn, J.H., Dagenais, N., Christensen, S.K., Nguyen, J.T., Chory, J., Harrison, M.J., &Weigel, D. 1999. Activation tagging of the floral inducer FT. Science 286: 1962-1965.
Kiba, T., Henriques, R., Sakakibara, H., & Chua, N.H. 2007. Targeted degradation of PSEUDO-RESPONSE
REGULATOR5 by an SCFZTL complex regulates clock function and photomorphogenesis in Arabidopsis
thaliana. Plant Cell 19: 2516-2530.
Kim, J., Geng, R., Gallenstein, R.A., & Somers, D.E. 2013. The F-box protein ZEITLUPE controls stability andnucleocytoplasmic partitioning of GIGANTEA. Development 140: 4060-4069.
Kim, W.Y., Fujiwara, S., Suh, S.-S., Kim, J., Kim, Y., Han, L., David, K., Putterill, J., Nam, H.G., & Somers, D.E.
2007. ZEITLUPE is a circadian photoreceptor stabilized by GIGANTEA in blue light. Nature 449: 356-360.
Knott, J.E. 1934. Effect of a localized photoperiod on spinach. Proc. Amer. Soc. Hort. Sci. 31: 152–154.

S. Yamaoka & T. Kohchi-7
BSJ-Review 7:84 (2016)

植物科学最前線 7:85 (2016)

Kobayashi, Y., Kaya, H., Goto, K., Iwabuchi, M., & Araki, T. 1999. A pair of related genes with antagonistic roles
in mediating flowering signals. Science 286: 1960-1962.
ἙෆᏕஅ࣭▼ᓮබᗤ 2012. ྂࡃ࡚᪂ࡋ࠸ࣔࢹ᳜ࣝ≀࡜ࡋ࡚ࡢࢱ࢖㢮ࢮࢽࢦࢣࡢ≉ᚩ. BSJ-Review 3:58-70.
Kubota, A., Kita, S., Ishizaki, K., Nishihama, R., Yamato, K.T., & Kohchi, T. 2014. Co-option of a photoperiodic
growth-phase transition system during land plant evolution. Nat. Commun. 5: 3668.
Más, P., Kim, W.Y., Somers, D.E., & Kay, S.A. 2003. Targeted degradation of TOC1 by ZTL modulates circadianfunction in Arabidopsis thaliana. Nature 426: 567-570.
Matsuo, T., Okamoto, K., Onai, K., Niwa, Y., Shimogawara, K., & Ishiura, M. 2008. A systematic forward genetic
analysis identified components of the Chlamydomonas circadian system. Genes Dev. 22: 918-930.
Mittag, M., Kiaulehn, S., & Johnson, C.H. 2005. The circadian clock in Chlamydomonas reinhardtii. What is itfor? What is it similar to? Plant Physiol. 137: 399-409.
Nelson, D.C., Lasswell, J., Rogg, L.E., Cohen, M.A., & Bartel, B. 2000. FKF1, a clock-controlled gene that
regulates the transition to flowering in Arabidopsis. Cell 101: 331-340.
Park, D.H., Somers, D.E., Kim, Y.S., Choy, Y.H., Lim, H.K., Soh, M.S., Kim, H.J., Kay, S.A., & Nam, H.G. 1999.
Control of circadian rhythms and photoperiodic flowering by the Arabidopsis GIGANTEA gene. Science 285:
1579-1582.
Qiu, Y.-L., et al. 2006. The deepest divergences in land plants inferred from phylogenomic evidence. Proc. Natl.
Acad. Sci. USA 103: 15511-15516.
Rubinstein, C.V., Gerrienne, P., de la Puente, G.S., Astini, R.A., & Steemans, P. 2010. Early middle Ordovician
evidence for land plants in Argentina (eastern Gondwana). New Phytol. 188: 365-369.
Sawa, M., & Kay, S.A. 2011. GIGANTEA directly activates Flowering Locus T in Arabidopsis thaliana. Proc.
Natl. Acad. Sci. USA 108: 11698-11703.
Sawa, M., Nusinow, D.A., Kay, S.A., & Imaizumi, T. 2007. FKF1 and GIGANTEA complex formation isrequired for day-length measurement in Arabidopsis. Science 318: 261-265.
Schultz, T.F., Kiyosue, T., Yanovsky, M., Wada, M., & Kay, S.A. 2001. A role for LKP2 in the circadian clock ofArabidopsis. Plant Cell 13: 2659-2670.
Somers, D.E., Kim, W.Y., & Geng, R. 2004. The F-box protein ZEITLUPE confers dosage-dependent control on
the circadian clock, photomorphogenesis, and flowering time. Plant Cell 16: 769-782.
Song, Y.H., Smith, R.W., To, B.J., Millar, A.J., & Imaizumi, T. 2012. FKF1 conveys timing information for
CONSTANS stabilization in photoperiodic flowering. Science 336: 1045-1049.
Steemans, P., Hérissé, A.L., Melvin, J., Miller, M.A., Paris, F., Verniers, J., & Wellman, C.H. 2009. Origin andradiation of the earliest vascular land plants. Science 324: 353.
Suárez-López, P., Wheatley, K., Robson, F., Onouchi, H., Valverde, F., & Coupland, G. 2001. CONSTANS
mediates between the circadian clock and the control of flowering in Arabidopsis. Nature 410: 1116-1120.
Suetsugu, N., & Wada, M. 2013. Evolution of three LOV blue light receptor families in green plants andphotosynthetic stramenopiles: phototropin, ZTL/FKF1/LKP2 and aureochrome. Plant Cell Physiol. 54: 8-23.
Takase, T., Nishiyama, Y., Tanihigashi, H., Ogura, Y., Miyazaki, Y., Yamada, Y., & Kiyosue, T. 2011. LOV KELCH

S. Yamaoka & T. Kohchi-8
BSJ-Review 7:85 (2016)

植物科学最前線 7:86 (2016)

PROTEIN2 and ZEITLUPE repress Arabidopsis photoperiodic flowering under non-inductive conditions,dependent on FLAVIN-BINDING KELCH REPEAT F-BOX1. Plant J. 67: 608-621.
Tamaki, S., Matsuo, S., Wong, H.L., Yokoi, S., & Shimamoto, K. 2007. Hd3a protein is a mobile flowering signalin rice. Science 316: 1033-1036.
Taoka, K., Ohki, I., Tsuji, H., Furuita, K., Hayashi, K., Yanase, T., Yamaguchi, M., Nakashima, C., Purwestri, Y.A.,
Tamaki, S., Ogaki, Y., Shimada, C., Nakagawa, A., Kojima, C., & Shimamoto, K. 2011. 14-3-3 proteins act as
intracellular receptors for rice Hd3a florigen. Nature 476: 332-335.
Wellman C.H., Osterloff P.L., & Mohiuddin U. 2003. Fragments of the earliest land plants. Nature 425: 282–285.
Wickett, N.J. et al. 2014. Phylotranscriptomic analysis of the origin and early diversification of land plants. Proc.
Natl. Acad. Sci. USA 111: E4859-E4868.
Wigge, P.A., Kim, M.C., Jaeger, K.E., Busch, W., Schmid, M., Lohmann, J.U., & Weigel, D. 2005. Integration ofspatial and temporal information during floral induction in Arabidopsis. Science 309: 1056-1059.
Yamaoka, S., Takenaka, M., Hanajiri, T., Shimizu-Ueda, Y., Nishida, H., Yamato, K.T., Fukuzawa, H., & Ohyama
K. 2004. A mutant with constitutive sexual organ development in Marchantia polymorpha L. Sex. Plant
Reprod. 16: 253-257.
Yasuhara, M., Mitsui, S., Hirano, H., Takanabe, R., Tokioka, Y., Ihara, N., Komatsu, A., Seki, M., Shinozaki, K., &Kiyosue, T. 2004. Identification of ASK and clock-associated proteins as molecular partners of LKP2 (LOVkelch protein 2) in Arabidopsis. J. Exp. Bot. 55: 2015-2027.

S. Yamaoka & T. Kohchi-9
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