植物科学最前線 7:288 (2016)ⴥࡢ㐍໬Ⓨ⏕Ꮫⓗ◊✲࡟ࡘ࠸࡚⪃࠼ࡿᕷᶫ Ὀ⠊⌮໬Ꮫ◊✲ᡤ ⎔ቃ㈨※⛉Ꮫ◊✲ࢭࣥࢱ࣮
ࠛ230-0045 ⚄ዉᕝ┴ᶓ὾ᕷ㭯ぢ༊ᮎᗈ⏫ 1-7-22
Yasunori Ichihashi
Evolutionary developmental studies of leaf shape
Key words: Evo-Devo, Gene expression, Leaf, Network analysis
RIKEN Center for Sustainable Resource Science, Yokohama, Kanagawa, 230-0045 Japanࡣࡌࡵ࡟ ࡝࠺ࡋ࡚⏕≀ࡢᙧࡣࡇࢇ࡞࡟ࡶከᵝ࡞ࡢ࠿㸽⏕≀ࡢᙧែ㐍໬ࡣ㸪㐍໬⏕≀Ꮫ⪅ࢲ࣮࢘࢕ࣥ
ࡢ᫬௦࠿ࡽ⏕≀Ꮫศ㔝࡟࠾ࡅࡿ㔜኱࡞ㄢ㢟ࡢ୍ࡘ࡛࠶ࡿࠋ㏆ᖺ㸪඲⏕≀࡛ඹ㏻ࡍࡿ DNA ศᏊ
ࢆ㏻ࡋ࡚⏕≀ࡢᙧែ㐍໬ࢆ⌮ゎࡍࡿ㐍໬Ⓨ⏕Ꮫࡀὀ┠ࢆᾎࡧ࡚࠸ࡿࠋࡇࡢศ㔝ࡣ㸪Ⓨ⏕ࢆไᚚࡍࡿศᏊ࣓࢝ࢽࢬ࣒ࢆ✀㛫࡛ẚ㍑ゎᯒࡍࡿࡇ࡜࡛㸪࡝ࡢࡼ࠺࡟ከᵝ࡞ᙧែࡀ㐍໬ࡋ࡚ࡁࡓ
ࡢ࠿ศᏊ࡛ࣞ࣋ࣝ᫂ࡽ࠿࡟ࡍࡿࡇ࡜ࢆ┠ⓗ࡜ࡍࡿࠋ⌧ᅾࡲ࡛࡟㸪ື᳜≀ࢆၥࢃࡎᖜᗈ࠸⏕≀✀࡛㐍໬Ⓨ⏕Ꮫⓗ◊✲ࡀ⾜ࢃࢀ࡚࠾ࡾ㸪ከࡃࡢࢣ࣮ࢫ࡛᪂つ㑇ఏᏊࡢ⋓ᚓࡼࡾࡶ㑇ఏᏊⓎ⌧ࡢኚ໬ࡀᙧែ㐍໬࡟኱ࡁࡃ㈉⊩ࡋ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟ࡉࢀ࡚ࡁࡓ㸦Blein et al. 2008, Kimura
et al. 2008, Rebeiz et al. 2009, Loehlin and Werren 2012㸧ࠋࡑࡢࡓࡵ㸪Ⓨ⏕ࢆྖࡿ㑇ఏᏊไᚚࢿࢵ
ࢺ࣮࣡ࢡࡢࡘ࡞ࡂ᥮࠼ࡀᙧែ㐍໬ࢆᘬࡁ㉳ࡇࡍ㥑ືຊ࡛࠶ࡿ࡜ᥦ᱌ࡉࢀ࡚࠸ࡿ㸦Peter andDavidson 2011㸧ࠋ 㐍໬ࡢ㐣⛬࡟࠾࠸࡚⿕Ꮚ᳜≀ࡣ㸪⎔ቃࡢኚ໬࡟㐺ᛂࡍࡿࡓࡵ࡟ⴥࡢᙧࢆከᵝ໬ࡉࡏ࡚ࡁࡓࠋ
ࡓ࡜࠼ࡤࢸࣥࢪࢡ࢔࢜࢖ᒓ㸦Pelargonium㸧࡟࠾࠸࡚ࡣ㸪ྠᒓෆ࡜࠸࠺㏆⦕✀㛫࡛ࡶ༢ⴥ࠿ࡽ
」ⴥࡲ࡛ከᵝᛶ࡟ᐩࢇࡔⴥࡢᙧែࢆほᐹࡍࡿࡇ࡜ࡀ࡛ࡁࡿ㸦Nicotra et al. 2011㸧ࠋࡲࡓ῱ὶἢ
࠸᳜≀ࡣ⣽࠸ⴥࢆⓎ⏕ࡉࡏ࡚⎔ቃ࡟㐺ᛂࡋ࡚࠾ࡾ㸪࢖ࢾ࢞ࣛࢩᒓࡢ Rorippa aquatica ࡣྠ୍ಶయ࡛ࡶ⎔ቃ࡟ᛂࡌ࡚ⴥࡢᙧࢆኚ໬ࡉࡏࡿ㸦Fassett 1940, van Steenis 1981㸧ࠋࡇࡢࡼ࠺࡟ⴥࡀ♧ࡍᙧែࡢከᵝᛶࡣ㸪᳜≀ࡢᙧែ㐍໬ࢆᨭ࠼ࡿไᚚ࣓࢝ࢽࢬ࣒ࢆ⌮ゎࡍࡿୖ࡛Ⰻ࠸◊✲ᑐ㇟࡜࡞ࡿࡓࡵ㸪⌧ᅾࡲ࡛࡟ࡉ࠿ࢇ࡟◊✲ࡉࢀ࡚ࡁࡓࠋࡑࡇ࡛ᮏ⥲ㄝ࡛ࡣ㸪ⴥࡢ㐍໬Ⓨ⏕Ꮫⓗ◊✲ࢆᴫほࡍࡿࡇ࡜࡛㸪᳜≀ࡢᙧែ㐍໬࡟࠾࠸࡚ㄽࡌࡿࠋศᏊ࡛ࣞ࣋ࣝࡢẚ㍑ᙧែᏛⓗ࡞࢔ࣉ࣮ࣟࢳ࠿ࡽጞࡲࡾ㸪」ᩘࡢࣔࢹ᳜ࣝ≀ࢆᑐ㇟࡜ࡋࡓศᏊ㑇ఏᏛⓗ◊✲ࢆಠ▔ࡋ࡚㸪ⴭ⪅ࡽࡀ⾜ࡗࡓḟୡ௦ࢩ࣮ࢣࣥࢫᢏ⾡ࢆά⏝ࡋࡓ㐍໬Ⓨ⏕Ꮫⓗ◊✲ࢆ⤂௓ࡍࡿࠋࡇࡢࡼ࠺࡟ⴥࡢ㐍໬Ⓨ⏕Ꮫ࡜࠸࠺୍◊✲ศ㔝ࡢṔྐࢆ᣺ࡾ㏉ࡿࡇ࡜࡛㸪᳜≀ࡢ㐍໬ࢆ⌮ゎࡋࡼ࠺࡜ࡍࡿ◊✲ࡢ௒ᚋࡢ᪉ྥᛶ࡟ࡘ࠸࡚㆟ㄽࡋࡓ࠸ࠋ

Y. Ichihashi -1
BSJ-Review 7:288 (2016)

植物科学最前線 7:289 (2016)ⴥࡢⓎ⏕Ꮫ ᳜≀ࡢ࣎ࢸ࢕ࣉࣛࣥࡣ㸪ࣇ࢓࢖ࢺ࣐࣮࡜࿧ࡤࢀࡿᇶᮏ༢఩ࡢ⧞ࡾ㏉ࡋᵓ㐀࡛ᙧసࡽࢀ࡚࠾
ࡾ㸪ࣇ࢓࢖ࢺ࣐࣮ࡣⴥ㸪ⱼ㸪ࡑࡢ㛫ࡢ⭆ⱆ࡛ᵓᡂࡉࢀ࡚࠸ࡿࠋࡑࡢ୰࡛ࡶⴥࡀ᭱ࡶᙧែⓗ࡟ከᵝ໬ࡋ࡚࠾ࡾ㸪ⰼჾᐁࡶⴥ࠿ࡽὴ⏕ࡋࡓࡶࡢ࡛࠶ࡿࡇ࡜ࢆ⪃࠼ࡿ࡜㸪ⴥࡢⓎ⏕ࢆ◊✲ࡍࡿࡇ࡜ࡣ㸪⮬↛⏺࡛ࡳࡽࢀࡿ᳜≀ࡢᙧែࡢከᵝᛶࡢ⌮ゎ࡟ࡘ࡞ࡀࡿ㸦Tsukaya 2002㸧ࠋ ௚ࡢከࡃࡢ⏕≀⌧㇟࡟࠾ࡅࡿ◊✲࡜ྠᵝ࡟㸪ⴥࡢⓎ⏕࡟㛵㐃ࡍࡿ◊✲ࡶࣔࢹ᳜ࣝ≀ࡢኚ␗య࠿ࡽከࡃࡢ㑇ఏᏛⓗ࡞▱ぢࡀᚓࡽࢀࡓ㸦ᅗ 1㸧ࠋࢡࣛࢫ I KNOTTED-LIKE HOMEOBOX㸦௨㝆 ࡣ KNOX ࡜ ␎ ࡍ 㸧 㑇 ఏ Ꮚ ࡣ ⱼ 㡬 ศ ⿣ ⤌ ⧊ 㸦 SAM 㸧 ࡢ ↓ 㝈 ᡂ 㛗 ᛶ ࢆ ⥔ ᣢ ࡋ ࡚ ࠾ ࡾ 㸪
BLADE-ON-PETIOLE㸦BOP㸧㑇ఏᏊ࡜ࡑࢀ࡟ࡼࡗ࡚┤᥋ไᚚࡉࢀࡿ ASYMMETRIC LEAVES2
㸦AS2㸧ཬࡧࡑࡢ┦஫స⏝ᅉᏊ AS1 ࡣ KNOX 㑇ఏᏊࢆᢚไࡍࡿࡇ࡜࡟ࡼࡾ㸪SAM ࠿ࡽࡢ୍⩌ࡢ⣽⬊ࢆⴥࡢ⣽⬊࡜ࡋ࡚㐠࿨௜ࡅࡿ㸦Semiarti et al. 2001, Byrne et al. 2002, Iwakawa et al. 2002,
Ha et al. 2003, Lin et al. 2003, Xu et al. 2003, Ha et al. 2004, Hepworth et al. 2005, Norberg et al.
2005, Zgurski et al. 2005, Fu et al. 2007, Ha et al. 2007, Iwakawa et al. 2007, Ueno et al. 2007, Guo et
al. 2008, Ha et al. 2010, Jun et al. 2010, Ichihashi et al. 2011㸧ࠋຍ࠼࡚㸪ࡇࡢ BOP ࠾ࡼࡧ AS1/2 㑇ఏᏊࡣⴥࡢ୕ḟඖ඲࡚ࡢ㍈㸪ࡍ࡞ࢃࡕඛ➃íᇶ㒊㍈,୰ኸíഃ᪉㍈㸪ྥ⫼㍈ࡢᙧᡂ࡟ࡶ㛵୚ࡋ࡚࠸ࡿ㸦Semiarti et al. 2001, Ha et al. 2003, Lin et al. 2003, Xu et al. 2003, Fu et al. 2007, Ha et al. 2007,
Ikezaki et al. 2010, Ichihashi et al. 2011, Kojima et al. 2011, Ishibashi et al. 2012, Chen et al. 2013㸧ࠋྥ⫼㍈ࡢᙧᡂ࡟㛵ࡋ࡚ࡣ㸪௚࡟ࡶከࡃࡢᅉᏊࡀ᫂ࡽ࠿࡟ࡉࢀ࡚࠾ࡾ㸪ྥ㍈ᙧᡂ࡟㛵୚ࡍࡿ㑇ఏᏊ࡜⫼㍈ᙧᡂ࡟㛵୚ࡍࡿ㑇ఏᏊ࡜ࡀ➇ྜࡋ࡚ไᚚࡋྜ࠺ࡇ࡜࡛ⴥࡢᖹ㠃ᡂ㛗ࡀ⥔ᣢࡉࢀ࡚࠸ࡿ㸦Fukushima and Hasebe 2014㸧ࠋⴥࡀ♧ࡍ᭷㝈ᡂ㛗ᛶࡣ㸪microRNA JAW ࡟ࡼࡾไᚚࡉࢀ
࡚࠸ࡿࢡࣛࢫ II TEOSINTE BRANCHED1/CYCLOIDEA/PCF㸦TCP㸧㑇ఏᏊࡀⴥࡢᡂ⇍ࢫࢣࢪ࣮ࣗࣝࡢ㐍⾜ࢆᢚไࡍࡿࡇ࡜࡛ไᚚࡉࢀ࡚࠸ࡿ㸦Nath et al. 2003, Palatnik et al. 2003, Efroni et al.
2008㸧ࠋࡇࡢ TCP 㑇ఏᏊࡢୗὶ࡟࠾࠸࡚㸪microRNA164 ࡀ CUP-SHAPED COTYLEDON㸦CUC㸧
㑇ఏᏊࡢⓎ⌧ሙᡤࢆᚤㄪᩚࡍࡿࡇ࡜࡟ࡼࡾ㸪ⴥࡢ࿘⦕㒊ࡢᙧࡶไᚚࡉࢀ࡚࠸ࡿ㸦Nikovics etal. 2006, Koyama et al. 2007, Kawamura et al.
2010, Koyama et al. 2010㸧ࠋຍ࠼࡚ CUC ࡜ KNOXࡣ┦஫࡟Ⓨ⌧ࢆୖ᪼ࡉࡏࡿṇࡢࣇ࢕࣮ࢻࣂࢵ
ࢡ࣮ࣝࣉ࡟ࡼࡾᙧែᙧᡂࢆᘬࡁ㉳ࡇࡍ࡜⪃࠼ࡽ࠼࡚࠸ࡿ㸦Takada et al. 2001, Blein et al. 2008,
Kawamura et al. 2010㸧ࠋࡇࡢࡼ࠺࡟㸪ࡇࢀࡽࡢไᚚᅉᏊࡀⴥࡢⓎ⏕ࢆ⢭ᐦ࡟ࢥࣥࢺ࣮ࣟࣝࡋ㸪㐍໬ࡢ㐣⛬࡛ࡇࢀࡽࡢᅉᏊࡢไᚚࡀኚ໬ࡍࡿ
࡜㸪ⴥࡢᙧែࡣࢲ࢖ࢼ࣑ࢵࢡ࡟ኚ໬ࡍࡿ࡜⪃࠼ࡽࢀࡿࠋᅗ㸯ⴥࡢⓎ⏕࡟㛵୚ࡍࡿ୺せ࡞㑇ఏᏊ

Y. Ichihashi -2
BSJ-Review 7:289 (2016)

植物科学最前線 7:290 (2016)ⴥࡢ㐍໬Ⓨ⏕Ꮫⓗ◊✲ࡢࡉࡁࡀࡅ ୖグࡋࡓࡼ࠺࡞ከᵝ࡞ⴥࡢᙧࢆ♧ࡍኚ␗యࡢ◊✲࡟ࡼࡾ㸪⮬↛⏺࡛ほᐹ࡛ࡁࡿⴥࡢከᵝᛶࢆ㑇ఏᏊ࡛ࣞ࣋ࣝ⌮ゎࡍࡿ✺◚ཱྀࡀ㛤ࡅࡓࠋࡑࡢ୰࡛ࡶ㸪ⴥࡢᙧែࢆ๻ⓗ࡟ኚ໬ࡉࡏࡿ KNOX
㑇ఏᏊࡀࡲࡎ╔┠ࡉࢀࡓࠋᵝࠎ࡞ⴥࡢᙧࢆ♧ࡍ᳜≀⩌࡟ࡘ࠸࡚ⴥཎᇶ࡟࠾ࡅࡿ KNOX 㑇ఏᏊࡢⓎ⌧ࡀẚ㍑ࡉࢀ㸪KNOX 㑇ఏᏊⓎ⌧࡜ⴥཎᇶࡢᙧែ࡟┦㛵ࡋ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ
㸦Bharathan et al. 2002㸧
ࠋ୍᪉࡛㸪」㞧࡞ⴥཎᇶ࡛࠶ࡗ࡚ࡶ᭱⤊ⓗ࡟༢ⴥࢆⓎ㐩ࡉࡏࡿ✀ࡶ࠶
ࡿࡓࡵ㸦Bharathan et al. 2002㸧㸪ⴥࡢ᭱⤊ᙧែࡔࡅ࡛࡞ࡃⴥࡢึᮇⓎ⏕࡟࠾ࡅࡿ㑇ఏᏊⓎ⌧ࢆ
㋃ࡲ࠼࡞ࡅࢀࡤ㸪┿ࡢ┦ྠᛶࡀ᫂ࡽ࠿࡟࡞ࡽ࡞࠸㸦ᅗ 2㸧ࠋࡇࢀࢆᶵ࡟㸪ẚ㍑ᙧែᏛⓗ࡞࢔ࣉ࣮ࣟࢳ࡟㑇ఏᏊⓎ⌧ࢆẚ㍑ࡍࡿ࡜࠸࠺᪂ᶵ㍈ࢆຍ࠼ࡓ㐍໬Ⓨ⏕Ꮫⓗ◊✲ࡀࡉ࠿ࢇ࡟⾜ࢃࢀ㸪ᙧែ㐍໬࡟࠾ࡅࡿ┦ྠᛶࢆ῝ࡃ⌮ゎ࡛ࡁࡿࡼ࠺࡟࡞ࡗࡓࠋ ༢ⴥ࡜」ⴥ Ѹ  ᳜≀ࡢⴥࡣ༢ⴥ࡜」ⴥ࡟
኱ูࡉࢀ㸪༢ⴥ࡜」ⴥ࡟࠾ࡅࡿჾᐁࡢ┦ྠᛶࡣ㛗࠸㛫㆟ㄽࡉࢀ࡚ࡁࡓࠋࡍ࡞ࢃࡕ」ⴥ඲య
ࡀ༢ⴥ࡜┦ྠ࡞ࡢ࠿㸪」ⴥࢆᵓᡂࡍࡿಶࠎࡢ
ᑠⴥࡀ༢ⴥ࡜┦ྠ࡞ࡢ࠿㸪ᙧែࡢࡳࡢẚ㍑࠿
ࡽࡔࡅ࡛ࡣ᫂ࡽ࠿࡛࡞࠿ࡗࡓ㸦Champagne andSinha 2004㸧ࠋࢺ࣐ࢺࡢ」ⴥࢆ⏝࠸ࡓ୍㐃ࡢⓎ⏕Ꮫⓗゎᯒ࠿ࡽ㸪༢ⴥᙧᡂ࡟㔜せ࡞ KNOX 㑇ఏᏊࡢไᚚ㸪᳜≀࣍ࣝࣔࣥࡢ࣮࢜࢟ࢩࣥ㸪CUC
㑇ఏᏊ࡞࡝ࡢ㌿෗ᅉᏊࡢไᚚࡀ」ⴥࡢᑠⴥᙧᡂ࡟ࡶ㔜せ࡛࠶ࡿࡇ࡜ࡀࢃ࠿ࡗࡓ㸦Ori et al.
2007, Berger et al. 2009, David-Schwartz et al.
2009, Koenig et al. 2009㸧ࠋࡇࢀࡽࡢ▱ぢࢆࡶ࡜࡟」ⴥ࡜༢ⴥࡢ┦ྠᛶ࡟ࡘ࠸࡚⪃࠼ࡿ࡜㸪ୖグࡢ 2 ࡘࡢ௬ㄝࡣ┦཯ࡍࡿࡶࡢ࡛ࡣ࡞ࡃ㸪
KNOX ࡸ CUC ࡞࡝ࡢไᚚᅉᏊࡣ᳜≀ࡢᙧែࣃࢱ࣮ࣥࡢ㦵᱁࡜࡞ࡿࠕᯞศ࠿ࢀࡢࣃࢱ࣮ࣥ
ᙧᡂࠖࢆ♽ඛ✀࡟࠾࠸࡚ไᚚࡋ࡚࠾ࡾ㸪㐍໬ࡢ㐣⛬࡟࠾࠸࡚ࡇࡢไᚚ⣔ࡀⴥࡢⓎ⏕ࣉࣟࢢ࣒ࣛ࡟෌฼⏝ࡉࢀࡓࡓࡵ㸪༢ⴥ࠿ࡽ」ⴥࡀ

ᅗ㸰ᡂ⇍ⴥ࡜ⴥཎᇶࡢ㐪࠸
ⴥࡢึᮇⓎ⏕࡜㑇ఏᏊⓎ⌧ࢆㄪ࡭ࡿࡇ࡜࡛┿ࡢ┦ྠᛶࢆ⌮ゎ࡛ࡁࡿࠋKNOX ࡢ㑇ఏᏊⓎ⌧㒊఩ࢆ㉥࡛♧ࡍࠋ

㐍໬ࡋࡓ࡜⪃࠼ࡽࢀࡿ㸦Koenig and Sinha 2010㸧ࠋ ୧㠃ⴥ࡜༢㠃ⴥ Ѹ ⴥࡣ㏻ᖖ㸪⾲ഃ࡜⿬ഃࡢᛶ㉁ࢆࡶࡘᖹࡓ࠸ᙧ࡟࡞ࡿࠕ୧㠃ⴥ࡛ࠖ࠶ࡿ
ࡀ㸪࢔࣓ࣖࡸࢿࢠ࡜࠸ࡗࡓ୍㒊ࡢ᳜≀ࡣ㸪⿬ഃࡢᛶ㉁ࡋ࠿ᣢࡓ࡞࠸ࠕ༢㠃ⴥࠖࢆࡘࡃࡿࠋ࢖
ࢢࢧᒓ㸦Juncus㸧ࢆ⏝࠸ࡓẚ㍑㑇ఏᏊⓎ⌧ゎᯒ࡟ࡼࡾ㸪༢㠃ⴥ࡛ࡣⴥࡢ⿬ഃࡢᛶ㉁ࢆỴࡵࡿ
AUXIN RESPONSE FACTOR3/ETTIN 㑇ఏᏊࡀⴥཎᇶ඲య࡛ാࡃࡇ࡜࡛㸪⿬ഃࡢᛶ㉁ࡋ࠿ᣢࡓ࡞

Y. Ichihashi -3
BSJ-Review 7:290 (2016)

植物科学最前線 7:291 (2016)ࡃ࡞ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ㸦Yamaguchi et al. 2010㸧ࠋ୍᪉࡛༢㠃ⴥ࡛ࡶᖹࡓ࠸ᙧࡢⴥࢆࡘࡃ
ࡿ✀㸦ⱼ㡬࡟ᑐࡋ࡚ⴥࡀᒎ㛤ࡍࡿ᪉ྥࡀ୧㠃ⴥ࡜ 90 ᗘ㐪࠺㸧ࡀᏑᅾࡋ࡚࠾ࡾ㸪DROOPING
LEAF 㑇ఏᏊࡢⓎ⌧ኚ໬ࡀྥ⫼㍈᪉ྥ࡟ἢࡗࡓ⣽⬊ቑṪࢆಁ㐍ࡍࡿࡇ࡜࡛ᖹࡓ࠸ᵓ㐀ࢆࡘࡃࡿࠋࡇࡢࡇ࡜࠿ࡽ㸪ⴥࡢᖹࡓࡉ࡜࠸࠺ᙧែࡣ୧㠃ⴥ࡜༢㠃ⴥ࡛┦ྠ࡛ࡣ࡞ࡃ㸪඲ࡃ␗࡞ࡿ࣓
࢝ࢽࢬ࣒࡛ࡘࡃࡽࢀࡿࡇ࡜ࡀࢃ࠿ࡗࡓ㸦Yamaguchi et al. 2010㸧ࠋ ⴥ࡜௬ⴥᯞ Ѹ ௬ⴥᯞ࡜ࡣⓎ⏕ࡍࡿ఩⨨࠿ࡽᯞ࡟┦ᙜࡍࡿࡀ㸪ⴥࡢࡼ࠺࡞ᙧែࢆ♧ࡍჾᐁ࡛࠶ࡿࠋ࢔ࢫࣃࣛ࢞ࢫᒓ㸦Asparagus㸧ࡢ௬ⴥᯞᙧᡂ࡟࠾ࡅࡿヲ⣽࡞㑇ఏᏊⓎ⌧ゎᯒ࡟ࡼࡾ㸪௬ⴥᯞ࡛ࡣᯞ࡛ᶵ⬟ࡍࡿ KNOX 㑇ఏᏊ࡜㸪ⴥ࡛ᶵ⬟ࡍࡿ AS1 㑇ఏᏊࡢ୧᪉ࡀⓎ⌧ࡋ࡚࠸ࡿࡇ
࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ㸦Nakayama et al. 2012㸧ࠋຍ࠼࡚㸪௬ⴥᯞ࡛ぢࡽࢀࡓ AS1 ࡢ㑇ఏᏊⓎ⌧ࣃࢱ࣮ࣥࡣ㸪ⴥ࡛ほᐹࡉࢀࡿࣃࢱ࣮ࣥ࡜㠀ᖖ࡟ࡼࡃఝ࡚࠸ࡿࡇ࡜࠿ࡽ㸪௬ⴥᯞࡣᯞ࡟ⴥࡢᛶ㉁ࡀ௜ຍࡉࢀ࡚⏕ࡌࡓჾᐁ࡛࠶ࡿࡇ࡜ࡀ᫂ࡽ࠿࡜࡞ࡗࡓ㸦Nakayama et al. 2012㸧ࠋ ୖグ࡛♧ࡋࡓࡼ࠺࡟㸪ึᮇⓎ⏕࡜㑇ఏᏊⓎ⌧ࢆヲ⣽࡟ㄪ࡭ࡿࡇ࡜࡛ჾᐁࡢ┦ྠᛶ࡜ࡑࡢศ
Ꮚ࣓࢝ࢽࢬ࣒ࡢ୍➃ࡀ᫂ࡽ࠿࡟࡞ࡿࠋ୍᪉࡛㸪ᙧែ㐍໬࡟┦㛵ࡋࡓⓎ⌧ࢆ♧ࡍ㑇ఏᏊࡀᐇ㝿࡟ᙧែ㐍໬࡟㈉⊩ࡋࡓ࠿㸪ࡍ࡞ࢃࡕཎᅉ࡞ࡢ࠿⤖ᯝ࡞ࡢ࠿ࢆⓎ⌧ゎᯒࡢࡳ࠿ࡽ࡛ࡣ⤖ㄽ௜ࡅࡿࡇ࡜ࡣ࡛ࡁ࡞࠸ࠋࡑࡢࡓࡵᙧែ㐍໬ࡢ⫼ᬒ࡟࠶ࡿ࣓࢝ࢽࢬ࣒ࢆ㑇ఏᏊ࡛ࣞ࣋ࣝ⌮ゎࡍࡿࡓࡵ࡟ࡣ㸪ᙧ㉁㌿᥮࡞࡝ࢆ⏝࠸ࡓ㑇ఏᏊࡢ᧯సᐇ㦂࡟ࡼࡿᐇドࡀᚲせ࡟࡞ࡿࠋ㑇ఏᏊࡢᶵ⬟ゎᯒ࡟ࡼࡿⴥࡢᙧែ㐍໬ࡢᐇド◊✲ ࡑࡇ࡛ࣔࢹ᳜ࣝ≀࡜㏆⦕࡛㸪ⴥࡢᙧែࡀከᵝ࡞᳜≀✀࡬࡜◊✲ᑐ㇟ࡀᗈࡀࡾ㸪㑇ఏᏊ᧯స࡟ࡼࡿศᏊ㑇ఏᏛⓗゎᯒࡀ⾜ࢃࢀࡿࡼ࠺࡟࡞ࡗࡓࠋ ≉࡟ࣔࢹ᳜ࣝ≀ࢩࣟ࢖ࢾࢼࢬࢼ࡜㏆⦕࡛」ⴥࢆᣢࡘ࣑ࢳࢱࢿࢶࢣࣂࢼࡸ㸪◊✲ࣜࢯ࣮ࢫࡀ඘ᐇࡋࡓࢺ࣐ࢺ࡜㏆⦕✀ࢆᑐ㇟࡟ࡋࡓ◊✲ࡀඛ⾜ࡋ࡚ከࡃࡢ▱ぢࢆࡶࡓࡽࡋࡓ㸦ᅗ 3㸧
ࠋ๓㏙ࡋࡓ KNOX 㑇ఏᏊ࠾ࡼࡧࡑࢀࢆไᚚࡍࡿ
AS1 㑇ఏᏊࡢไᚚ࣓࢝ࢽࢬ࣒ࡀ࣑ࢳࢱࢿࢶࢣࣂࢼ࡜ࢩࣟ࢖ࢾࢼࢬࢼࡢ㛫࡛ẚ㍑ゎᯒࡉࢀࡓࠋ

ᅗ㸱ࣔࢹ᳜ࣝ≀࡜ࡑࡢ㏆⦕✀ࡢᡂ⇍ⴥࢩࣟ࢖ࢾࢼࢬࢼ㸦A. thaliana㸧࡜㏆⦕࡞࣑ࢳࢱࢿࢶࢣࣂࢼ㸦C. hirsuta㸧㸪ࢺ࣐ࢺ㸦S. lycopersicum㸧࡜㏆⦕࡞ S. pennellii ࡜ S. habrochaites ࡣከᵝ࡞ⴥࡢᙧࢆ♧ࡍࠋ

Y. Ichihashi -4
BSJ-Review 7:291 (2016)

植物科学最前線 7:292 (2016)ࡑࡢ⤖ᯝ㸪AS1 㑇ఏᏊࡢᶵ⬟ࡣಖᏑࡉࢀ࡚࠸ࡿࡶࡢࡢ KNOX 㑇ఏᏊࡢୖὶ࡟࠶ࡿไᚚ㓄ิࡢ㐪࠸࡟ࡼࡾ AS1-KNOX ไᚚ⣔ࡀ 2 ✀㛫࡛␗࡞ࡾⴥࡢᙧែࡢ㐪࠸ࡀ⏕ࡌ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟
࡞ࡗࡓ㸦Hay and Tsiantis 2006㸧ࠋຍ࠼࡚ࢩࣟ࢖ࢾࢼࢬࢼᒓෆ࡛ࡳࡽࢀࡿⴥࡢᙧែࡢከᵝᛶࡢ୍㒊ࡶ KNOX 㑇ఏᏊࡢୖὶ࡟࠶ࡿไᚚࡢኚ໬࡛ㄝ࡛᫂ࡁࡿࡇ࡜ࡀࢃ࠿ࡗࡓ㸦Piazza et al. 2010㸧ࠋ ୍᪉࡛㑇ఏᏊࡢⓎ⌧ไᚚ⣔ࡢ㐪࠸ࡔࡅ࡛࡞ࡃ㸪ࢱࣥࣃࢡ㉁࡛ࣞ࣋ࣝࡢไᚚ⣔ࡢ㐪࠸ࡀ✀㛫ࡢⴥࡢ㐪࠸ࢆ⏕ࡳฟࡍࢣ࣮ࢫࡀࢺ࣐ࢺ࡜㏆⦕✀ Solanum galapagense ࡜ࡢゎᯒ࡛᫂ࡽ࠿࡟࡞ࡗ
ࡓࠋKNOX ࢱࣥࣃࢡ㉁ࡣ BEL-LIKE HOMEODOMAIN ࢱࣥࣃࢡ㉁࡜」ྜయࢆᙧᡂࡋ࡚ᶵ⬟ࡍࡿࡀ㸪࣓࣍࢜ࢻ࣓࢖ࣥࢆᣢࡓ࡞࠸ KNOX ࢱࣥࣃࢡ㉁ PETROSELINUM 㸦PTS㸧 / KANTM ࡣࡑࡢ」ྜయᙧᡂࢆ㜼ᐖࡍࡿ㸦Kimura et al. 2008, Magnani and Hake 2008㸧ࠋS. galapagense ࡛ࡣ
PTS ࡢⓎ⌧ࡀ㐣๫࡜࡞ࡿ㑇ఏⓗኚ␗ࢆᣢࡕ㸪ࡑࡢࡓࡵ KNOX ࡜ BELL 」ྜయᙧᡂࡀ㜼ᐖࡉࢀୗὶࡢไᚚࡀኚ໬ࡍࡿࡇ࡜࡛㸪ⴥࡢᙧែࡀ」㞧໬ࡋࡓࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ㸦Kimura et al.
2008㸧ࠋ ࡲࡓ CUC 㑇ఏᏊࡣᵝࠎ࡞᳜≀✀࡟࠾ࡅࡿⴥࡢᙧែ࡟࠾࠸࡚ಖᏑࡉࢀࡓᶵ⬟ࢆᣢࡘࡇ࡜ࡀ
ᙧ㉁㌿᥮ᐇ㦂࡛᫂ࡽ࠿ࡉࢀ࡚࠾ࡾ㸪㑇ఏᏊ᧯సᐇ㦂࡟ࡼࡾࣔࢹ᳜ࣝ≀࡛᫂ࡽ࠿࡟ࡉࢀ࡚࠸ࡓ
ⴥࡢᙧែࡢไᚚᅉᏊ࡟ࡘ࠸࡚㐍໬ⓗࢫࢣ࣮࡛ࣝࡢᶵ⬟ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ㸦Blein et al. 2008㸧ࠋຍ࠼࡚㸪㏆⦕✀ࡢ㡰㑇ఏᏛⓗ࢔ࣉ࣮ࣟࢳ࡟ࡼࡾ㸪ࣔࢹ᳜ࣝ≀࡛ࡣḞኻࡋࡓ㑇ఏᏊࡀᙧែ㐍໬࡬㛵୚ࡋ࡚࠸ࡿࡇ࡜ࡶ♧ࡉࢀࡓ㸦Sicard et al. 2014, Vlad et al. 2014㸧ࠋ ௨ୖࡢࡼ࠺࡟㸪ⴥࡢᙧែ㐍໬࡟㈉⊩ࡋࡓ㑇ఏᏊࡉࡽ࡟ࡣ✀㛫ࡢᙧែⓗ┦㐪ࢆㄝ᫂ࡍࡿ㑇ఏⓗኚ␗ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋࡇࡢ▱ぢࡣ㸪㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡࡢࡘ࡞ࡂ᥮࠼ࡶࡋࡃࡣ㒊ศⓗ࡞ᨵኚࡀᙧែ㐍໬ࢆᘬࡁ㉳ࡇࡍ࡜࠸࠺௬ㄝࢆᨭᣢࡋࡓࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ಶࠎࡢ㑇ఏᏊࡢᶵ⬟ゎᯒ࠿ࡽࡔࡅ࡛ࡣ㸪Ⓨ⏕ࢆྖࡿ㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡࡀࢩࢫࢸ࣒࡜ࡋ࡚࡝ࡢࡼ࠺࡟ኚ໬
ࡋ࡚ᙧែ㐍໬ࢆᘬࡁ㉳ࡇࡋࡓࡢ࠿㆟ㄽ࡛ࡁ࡞࠸ࠋ

ࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒࠿ࡽࡳࡓⴥࡢᙧែ㐍໬ࡢ඲యീ ࡑࢀ࡛ࡣ୍య㸪࡝ࡢࡼ࠺࡞㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡࡢኚ໬ࡀ⮬↛⏺࡛ほᐹࡉࢀࡿࡼ࠺࡞ከ
ᵝ࡞ᙧࡢⴥࢆ㐍໬ࡉࡏࡿࡢࡔࢁ࠺࠿㸽ࡇࡢၥ㢟ゎỴ࡟࠾࠸࡚㸪㏆ᖺࡢḟୡ௦ࢩ࣮ࢣࣥࢧ࣮ࢆ
౑ࡗࡓࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒㸦RNA-seq㸧ࡀ᭷ຠ࡛࠶ࡿ㸦Ichihashi and Sinha 2014, Ichihashi et al.
2015㸧ࠋࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒ࡟ࡼࡾࢤࣀ࣒࣡࢖ࢻ࡛㑇ఏᏊⓎ⌧ࢹ࣮ࢱࢆᚓࡿࡇ࡜࡛㸪Ⓨ⏕⌧㇟࡞࡝ࡢ」㞧࡞⏕≀Ꮫⓗ࡞⌧㇟ࢆ⌮ゎࡍࡿࡇ࡜ࡀ࡛ࡁ㸪≉࡟ḟୡ௦ࢩ࣮ࢣࣥࢩࣥࢢᢏ⾡࡟ࡼࡾࢤࣀ࣒ࡀゎㄞࡉࢀ࡚࠸࡞࠸㠀ࣔࢹࣝ⏕≀࡟࠾࠸࡚ࡶࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒࢆ㐺⏝࡛ࡁࡿࡇ࡜ࡣ኱ࡁ࡞ࣈࣞ࢖ࢡࢫ࣮࡛ࣝ࠶ࡿ㸦Wang et al. 2009㸧ࠋຍ࠼࡚㸪ᩘᏛࡢ୍ศ㔝࡛࠶ࡿࢢࣛࣇ⌮ㄽࢆࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒ࡟ᑟධࡍࡿࡇ࡜࡟ࡼࡾ㸪ಶࠎࡢ㑇ఏᏊࡢ㛵ಀࢆࢩࢫࢸ࣒࡛ࣞ࣋ࣝ⌮ゎࡍࡿ㸪ࡍ࡞ࢃࡕ㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡࢆ᥎ᐃࡍࡿࡇ࡜ࡀ࡛ࡁࡿࡼ࠺࡟࡞
ࡗࡓ㸦Marbach et al. 2012㸧ࠋ ࡑࡇ࡛ⴭ⪅ࡽࡣ RNA-seq ࢆ⏝࠸࡚㸪ࢺ࣐ࢺ࡜㏆⦕✀㸦S. pennellii ࡜ S. habrochaites㸧㛫ࡢⴥ

Y. Ichihashi -5
BSJ-Review 7:292 (2016)

植物科学最前線 7:293 (2016)

ࡢⓎ⏕࡟ࡘ࠸࡚㐍໬Ⓨ⏕Ꮫⓗゎᯒࢆ⾜࠸㸪ከᵝ࡞ⴥࡢᙧࢆ㐍໬ࡉࡏࡓ㑇ఏᏊไᚚࢿࢵࢺ࣮࣡
ࢡୖࡢኚ໬ࢆ᫂ࡽ࠿࡟ࡋࡓ㸦Ichihashi et al. 2014㸧ࠋࡇࢀࡽࡢ᳜≀✀࡟࠾ࡅࡿⴥࡢⓎ⏕ࢫࢸ࣮ࢪูࡢ RNA-seq ࢹ࣮ࢱࢆ⏝࠸ࡓඹⓎ⌧ࢿࢵࢺ࣮࣡ࢡゎᯒ࡟ࡼࡾ㸪ࢺ࣐ࢺ✀࡟࠾ࡅࡿⴥࡢⓎ⏕ࡢ㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡࡢண

ࢆヨࡳࡓࠋࣂ࢖࢜࢖ࣥࣇ࢛࣐ࢸ࢕ࢵࢡࢫゎᯒ࡟ຍ࠼࡚⏕໬Ꮫᐇ㦂ࢆ⾜ࡗࡓ⤖ᯝ㸪ࢿࢵࢺ࣮࣡ࢡࡢ࿘⦕㒊ศ࡟ⴥࡢᙧࡢ」㞧ᛶࢆไᚚࡍࡿ PTS 㑇ఏᏊࢆྵࡴ
㑇ఏᏊࣔࢪ࣮ࣗࣝࡀ࠶ࡾ㸪ࡇࡢࣔࢪ࣮ࣗࣝෆ࡛ BOP ࡜ LSH ࡢࢱࣥࣃࢡ㉁」ྜయࡀ PTS ࡢⓎ⌧ࢆ┤᥋ไᚚࡋ㸪PTS ࡀ➇ྜࡍࡿ KNOX ࢱࣥࣃࢡ㉁㸦ࢺ࣐ࢺ࡛ࡣ LeT6㸧ࡢ㔞࡟ࡼࡗ࡚㸪ⴥࡢ⣽⬊ቑṪ㛵㐃ᅉᏊࢆྵࡴࢿࢵࢺ࣮࣡ࢡୖࡢከࡃࡢ㑇ఏᏊⓎ⌧ࡀไᚚࡉࢀ࡚࠸ࡿࡇ࡜ࡀࢃ࠿
ࡗࡓ㸦Kimura et al. 2008, Ichihashi et al. 2014㸧ࠋ⥆࠸࡚ࢺ࣐ࢺ✀㛫࡛ⴥࡢⓎ⏕ࡢ㑇ఏᏊไᚚࢿࢵ
ࢺ࣮࣡ࢡࡀ࡝ࡢࡼ࠺࡟ኚ໬ࡋ࡚࠸ࡿࡢ࠿᫂ࡽ࠿࡟ࡍࡿࡓࡵ࡟㸪ࣈ࣮ࢺࢫࢺࣛࢵࣉἲࢆ⏝࠸࡚ࢿࢵࢺ࣮࣡ࢡ⤫ィ㔞ࢆゎᯒࡍࡿࡇ࡜࡛ⴥࡢⓎ⏕ไᚚࢿࢵࢺ࣮࣡ࢡࢆ✀㛫࡛ẚ㍑ࡋࡓࠋຍ࠼࡚
஺㞧ಶయࢆ⏝࠸࡚୧ぶ✀ࡢ࢔ࣞࣝࢆ༊ูࡋࡓ㑇ఏᏊⓎ⌧ゎᯒࢆ⾜࠺ࡇ࡜࡟ࡼࡾ㸪Ⓨ⌧ࡢไᚚᵝᘧࢆㄪ࡭ࡓࠋ௨ୖࡼࡾ㸪BOP ࡢⓎ⌧ࢆไᚚࡍࡿ cis ᅉᏊୖࡢኚ␗ࡀ BOP-LSH-PTS ࡢ㑇ఏᏊࣔࢪ࣮ࣗࣝࡢⓎ⌧ࢆኚ໬ࡉࡏࡓࡓࡵ㸪✀㛫࡛ⴥࡢⓎ⏕ࡢ㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡ࡛኱つᶍ࡞Ⓨ⌧ࡢኚ໬ࡀ㉳ࡁ࡚࠸ࡿࡇ࡜ࡀ♧၀ࡉࢀࡓ㸦Ichihashi et al. 2014㸧ࠋ ࢺ࣐ࢺࡣࡍ࡛࡟ࢤࣀ࣒ࡀゎㄞࡉࢀ㸪ࡑࡢ㏆⦕✀࡟࠾࠸࡚ࡶศᏊ㑇ఏᏛⓗゎᯒࡀྍ⬟࡛࠶ࡿ
㸦Ranjan et al. 2012, Ichihashi and Sinha 2014㸧ࠋࡑࡇ࡛ᐇ㝿࡟ BOP-LSH-PTS ࡢ㑇ఏᏊࣔࢪ࣮ࣗ
ࣝࡢⓎ⌧ኚ໬ࡀᙧែ㐍໬ࢆᘬࡁ㉳ࡇࡋࡓ࠿᳨ドࡍࡿࡓࡵ㸪ୖὶᅉᏊࡢ BOP ࡜ LSH
㑇ఏᏊࡢⓎ⌧ࢆ᧯సࡋࡓᙧ㉁㌿᥮యࢆゎᯒࡍࡿ࡜㸪ࡑࢀࡽࡢ㑇ఏᏊࡢⓎ⌧㔞࡟ࡼࡗ࡚㸪ࢺ࣐ࢺ࡜ࡑࡢ㏆⦕✀࡟࠾ࡅࡿከᵝ
࡞ⴥࡢᙧ㉁ࡢ୍㒊ࡀ෌⌧ࡉࢀࡓ㸦Ichihashi
et al. 2014㸧ࠋࡇࡢ⤖ᯝࡣ BOP-LSH-PTS ࡢ㑇ఏᏊࣔࢪ࣮ࣗࣝࡀᘬࡁ㉳ࡇࡍࢿࢵࢺ࣡
࣮ࢡࡢ኱つᶍ࡞ࡘ࡞ࡂ᭰࠼ࡀⴥࡢᙧࡢከ
ᵝᛶࢆ⏕ࡳฟࡋࡓࡇ࡜ࢆ♧ࡋ㸪௒ࡲ࡛ࡢ㐍໬Ⓨ⏕Ꮫ࡛ᥦၐࡉࢀ࡚࠸ࡿ㸪ࠕ㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡ࡟࠾ࡅࡿ኱つᶍ࡞ࡘ࡞ࡂ
᭰࠼ࡀᙧែ㐍໬ࢆᘬࡁ㉳ࡇࡍࠖ࡜࠸࠺௬ㄝ ࢆ ᐇ 㦂 ࢹ ࣮ ࢱ ࡛ ᨭ ᣢ ࡋ ࡓ 㸦 Peter andDavidson 2011㸧ࠋࡲࡓ⌮ㄽࢿࢵࢺ࣮࣡ࢡ⏕≀Ꮫⓗ▱ぢ࠿ࡽ㸪㑇ఏᏊไᚚࢿࢵࢺ࣮࣡
ࢡࡢ࿘⦕࡟఩⨨ࡍࡿ㑇ఏᏊ⩌ࡢኚ␗ࡢ᪉
ࡀࢿࢵࢺ࣮࣡ࢡࡢࣁࣈ㑇ఏᏊ⩌ࡢኚ␗ࡼᅗ㸲ࢿࢵࢺ࣮࣡ࢡ࠿ࡽࡳࡓᙧែ㐍໬ࡢࣔࢹࣝ
ࡑࢀࡒࢀࡢ㑇ఏᏊࡣ㸪㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡୖ
ࡢ఩⨨࡟ࡼࡗ࡚ᙧែ㐍໬࡬ࡢᐤ୚ࡀ㐪࠺࡜⪃࠼ࡽࢀࡿࠋ

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植物科学最前線 7:294 (2016)ࡾࡶ㐍໬ⓗ࡞ᐤ୚ࡀ኱ࡁ࠸࡜ࡉࢀ࡚࠸ࡓࡀ㸦Vidal et al. 2011㸧㸪௒ᅇࡢ◊✲࡟ࡼࡾึࡵ࡚ࡑࡢ
ணࡀᐇ㦂ⓗ࡟ᨭᣢࡉࢀࡓࠋ๓㏙ࡋࡓࡼ࠺࡟㸪ᖜᗈ࠸᳜≀✀࡟࠾࠸࡚ KNOX 㑇ఏᏊࡢไᚚ࡟

࠾ࡅࡿኚ␗ࡀᙧែ㐍໬࡟⧞ࡾ㏉ࡋ㛵୚ࡍࡿࡇ࡜ࡣ㸦Bharathan et al. 2002, Hay and Tsiantis 2010,
Katayama et al. 2010, Piazza et al. 2010, Nakayama et al. 2012, Nakayama et al. 2014, Sicard et al.
2014, Vlad et al. 2014㸧㸪KNOX 㑇ఏᏊࡀᙧࡸ఩⨨᝟ሗࢆไᚚࡍࡿࢿࢵࢺ࣮࣡ࢡࣔࢪ࣮ࣗࣝ࡜⣽⬊ቑṪ࡟㛵୚ࡍࡿࢿࢵࢺ࣮࣡ࢡࣔࢪ࣮ࣗࣝࢆᶫΏࡍࡿ࣎ࢺࣝࢿࢵࢡୖ࡟࠶ࡿࡓࡵ࠿ࡶࡋࢀ࡞
࠸㸦ᅗ 4㸧
ࠋ௒ᚋࡇࡢࡼ࠺࡞ࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒࢆ㥑౑ࡋࡓ㐍໬Ⓨ⏕Ꮫⓗ◊✲ࡀ௚ࡢ᳜≀࡛ࡢᙧែ㐍໬࡟ᛂ⏝ࡉࢀࡿࡇ࡜࡟࡞ࢀࡤ㸪ࡇࡢ࢔࢖ࢹ࢔ࡢ୍⯡ᛶࡀ᳨ドࡉࢀ࡚࠸ࡃࡔࢁ࠺ࠋ

࠾ࢃࡾ࡟ ௒ᚋ㸪᳜≀ࡢ㐍໬Ⓨ⏕Ꮫศ㔝࡛ࡣ㸪ࢩ࣮ࢣࣥࢫᢏ⾡ࡸ㑇ఏᏊ᧯సᢏ⾡࡞࡝ࡢ㐍Ṍࡢࡓࡵ㸪ࡼࡾヲ⣽࡛ࢫࢣ࣮ࣝࡀ኱ࡁ࡞◊✲ࡀ୺ὶ࡟࡞ࡿࡇ࡜ࡣ᝿ീ࡟㞴ࡃ࡞࠸ࠋࡓ࡜࠼ࡤ㸪኱つᶍ࡞RNA-seq ࢆྍ⬟࡜ࡍࡿᢏ⾡࡟ࡼࡾ㸦Kumar et al. 2012, Townsley et al. 2015㸧㸪ࢿࢵࢺ࣮࣡ࢡࡢື
ⓗኚ໬ࢆ㏣࠺ࡇ࡜ࡀᐜ᫆࡜࡞ࡿࠋຍ࠼࡚㸪ࢿࢵࢺ࣮࣡ࢡࡢᵓ㐀ࢆẚ㍑ࡍࡿࣂ࢖࢜࢖ࣥࣇ࢛࣐
ࢸ࢕ࢵࢡࢫ࡟ࡼࡾ㸦Ideker and Krogan 2012, Fukushima 2013㸧㸪ಶࠎࡢ㑇ఏᏊ࡛ࣞ࣋ࣝ࡞ࡃࠕ㑇ఏᏊ࡜㑇ఏᏊࡢ┦஫స⏝࡛ࣞ࣋ࣝࠖᙧែ㐍໬ࢆゎࡁ᫂࠿ࡍࡇ࡜ࡀ࡛ࡁࡿࠋࡉࡽ࡟ゎᯒ࠿ࡽᚓࡽࢀࡓ⤖ᯝ࡟ᇶ࡙࠸࡚㸪CRISPR/Cas9 ࡞࡝ࡢࢤࣀ࣒⦅㞟ᢏ⾡ࢆ౑ࡗ࡚ேᕤⓗ࡟㑇ఏᏊ㛫ࡢ┦஫స⏝ࢆ᧯సࡍࡿࡇ࡜࡟ࡼࡾ㸦Kumar and Jain 2015㸧㸪㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡୖࡢಶࠎࡢ┦஫స⏝ࡢኚ໬ࡀ࡝ࡢࡼ࠺࡟ᙧែ㐍໬࡟⤖ࡧࡘࡃࡢ࠿᫂ࡽ࠿࡟࡞ࡿࠋ ࡲࡓᢏ⾡㐍Ṍࡔࡅ࡛࡞ࡃ㸪◊✲ᑐ㇟ࡶࡉࡽ࡟ᗈࡀࡿࡔࢁ࠺ࠋࡓ࡜࠼ࡤ㸪๓㏙ࡋࡓ༢㠃ⴥࡸ௬ⴥᯞࡢࡼ࠺࡟㸪≉ᐃࡢ✀ࡶࡋࡃࡣࢢ࣮ࣝࣉ࡟≉␗ⓗ࡞᪂つᙧ㉁ࡢ㐍໬⌧㇟࡟ᑐࡋ࡚㸪ࡼࡾ῝࠸⌮ゎࡀᚓࡽࢀࡿ࡜ண᝿ࡉࢀࡿࠋ≉࡟㣗⹸᳜≀ࡢᤕ⹸ⴥࡸᐤ⏕᳜≀ࡢ྾ჾ࡞࡝ࣘࢽ࣮ࢡ࡞
ᙧែ㐍໬࡟ࡘ࠸࡚㑇ఏᏊ࠾ࡼࡧ㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡ࡛ࣞ࣋ࣝ⌮ゎ࡛ࡁࢀࡤ㸪ࡑࡢࡼ࠺࡞
᳜≀ࡀ┿࡟≉ู࡞㑇ఏⓗ⫼ᬒࢆᣢࡗ࡚࠸ࡿࡢ࠿㸪ࡶࡋࡃࡣࡍ࡛࡟ಖᣢࡋ࡚࠸ࡿ㑇ఏᏊࢭࢵࢺ
ࢆ౑࠸ᅇࡍࡇ࡜࡛᪂つᙧ㉁ࢆ㐍໬ࡉࡏ࠺ࡿࡢ࠿᫂ࡽ࠿࡟࡞ࡿࠋࡉࡽ࡟ಶࠎࡢ᳜≀⣔⤫࡟࠾ࡅࡿ㐍໬ࡔࡅ࡛࡞ࡃ㸪⣔⤫ࢆ㉸࠼ࡓ࡛ࣞ࣋ࣝࡢ኱㐍໬ࡸ཰ᩡ㐍໬࡬ࡢ㐍໬Ⓨ⏕Ꮫⓗ࢔ࣉ࣮ࣟࢳ
ࡶ௒ᚋᮇᚅࡉࢀࡿࠋ ᭱ᚋ࡟ࡇࡢࡼ࠺࡞㐍໬Ⓨ⏕Ꮫⓗ◊✲࠿ࡽᚓࡽࢀࡓ▱ぢࡣ㸪ᇶ♏⛉Ꮫศ㔝࡟࡜࡝ࡲࡽࡎ㸪㎰స≀࡞࡝ࡢᨵⰋ࡟㈉⊩ࡍࡿࡇ࡜ࡀ࡛ࡁࡿ࠿ࡶࡋࢀ࡞࠸ࠋࡓ࡜࠼ࡤ㸪㐍໬ࡢ㐣⛬࡛㑅ᢥࡉࢀࡿ㑇ఏⓗኚ␗ࡣ㸪༢࡟≉ᐃࡢᙧ㉁ࡢࡳࡀ⎔ቃ࡟㐺ᛂࡋࡓ࡜ࡣ⪃࠼࡙ࡽࡃ㸪௚ࡢᙧ㉁࡟࠾ࡅࡿไ
ᚚ⣔࡜ࣂࣛࣥࢫࡢ࡜ࢀࡓ㑇ఏᏊไᚚࢿࢵࢺ࣮࣡ࢡࣔࢪ࣮ࣗࣝࡀ⮬↛⏺࡛㑅ᢥࡉࢀࡿྍ⬟ᛶࡀ༑ศ࡟⪃࠼ࡽࢀࡿࠋࡑࡇ࡛㸪⫱✀࡟࠾࠸࡚㎰ᴗ฼⏝ୖ౯್࠶ࡿ㑇ఏⓗᙧ㉁ࡢࡳ࡟╔┠ࡍࡿࡢ࡛ࡣ࡞ࡃ㸪㐍໬ⓗ⫼ᬒࢆ㋃ࡲ࠼ࡓ㸪ࢩࢫࢸ࣒࡜ࡋ࡚ࡼࡾ㡹ᙉ࡞᳜≀ᙧែࢹࢨ࢖ࣥࢆ┠ᣦࡍ࡭ࡁ࡛࠶ࢁ࠺ࠋࡑࡢ┠ⓗ㐩ᡂ࡟㐍໬Ⓨ⏕Ꮫⓗ◊✲࠿ࡽᚓࡽࢀࡿ▱ぢࡀ㈉⊩࡛ࡁࡿ࡜ᮇᚅࡉࢀࡿࠋ

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植物科学最前線 7:295 (2016)ㅰ㎡
ᮏ✏࡛⤂௓ࡋࡓⴭ⪅ࡽࡢ◊✲ࡢ୍㒊ࡣ㸪⌮໬Ꮫ◊✲ᡤ࣭ᇶ♏⛉Ꮫ≉ู◊✲ဨไᗘ࠾ࡼࡧ⛉Ꮫ◊✲⿵ຓ㔠࣭ⱝᡭ◊✲ B㸦15K18589㸧ࡢᨭ᥼ࢆᚓ࡚㐙⾜ࡋࡓࠋࡲࡓᮏ◊✲ࢆ㐍ࡵࡿ࡟࠶ࡓࡾ㸪ᩘࠎࡢࢧ࣏࣮ࢺࢆ㡬࠸ࡓ㸪࢝ࣜࣇ࢛ࣝࢽ࢔኱Ꮫࢹ࣮ࣅࢫᰯ࣭Prof. Neelima Sinha㸪Prof. Julin
Maloof㸪Dr. José Antonio Aguilar-Martínez㸪Dr. Moran Farhi㸪Dr. Ravi Kumar㸪Dr. Lee Millon㸪
Dr. Jie Peng㸪ࢻࢼࣝࢻࢲࣥࣇ࢛࣮ࢫ◊✲ᡤ࣭Dr. Daniel Chitwood ࡟㸪ࡇࡢሙࢆ೉ࡾ࡚࠾♩⏦ࡋୖࡆࡲࡍࠋᘬ⏝ᩥ⊩
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