植物科学最前線 7:241 (2016)ᐤ⏕᳜≀ࢥࢩ࣐࢜࢞ࡢ྾ჾᙧᡂᶵᵓ

ᓫ㞞1,2㸪ྜྷ⏣ ⪽Ꮚ3,4㸪ⓑ㡲 ㈼1,21
ᮾி኱Ꮫ኱Ꮫ㝔 ⌮Ꮫ⣔◊✲⛉ࠛ113-0033 ᮾி㒔ᩥி༊ᮏ㒓 7-3-12⌮໬Ꮫ◊✲ᡤ ⎔ቃ㈨※⛉Ꮫ◊✲ࢭࣥࢱ࣮
ࠛ230-0051 ⚄ዉᕝ┴ᶓ὾ᕷ㭯ぢ༊ᮎᗈ⏫ 1-7-223ዉⰋඛ➃⛉Ꮫᢏ⾡㝔኱Ꮫ㝔኱Ꮫ ࣂ࢖࢜ࢧ࢖࢚ࣥࢫ◊✲⛉ࠛ630-0192 ⏕㥖ᕷ㧗ᒣ⏫ 8916-5
4ዉⰋඛ➃⛉Ꮫᢏ⾡኱Ꮫ㝔኱Ꮫ ◊✲᥎㐍ᶵᵓࠛ630-0192 ⏕㥖ᕷ㧗ᒣ⏫ 8916-5
ⱝ➉Developmental study of haustorium in the parasitic plant Phtheirospermum japonicum
Key words: Auxin, Haustorium, Parasitic plant, Root development
Takanori Wakatake1,2, Satoko Yoshida3, Ken Shirasu1,2
1
Graduate School of Science, The University of Tokyo
7-3-1 Hongo, Bunkyo-ku, Tokyo, 113-0033, Japan
2
RIKEN Center for Sustainable Resource Science
1-7-22 Suehiru-cho, Tsurumi-ku Yokohama 230-0045, Kanagawa, Japan
3
Graduate School of Biological Sciences, Nara Institute of Science and Technology
8916-5 Takayama, Ikoma, Nara 630-0192, Japan
4
Institute for Research Initiatives, Division for Research Strategy, Nara Institute of Scienceand Technology, 8916-5, Takayama-cho, Ikoma, Nara, 630-0192, Japan㸯㸬ࡣࡌࡵ࡟௚ࡢ᳜≀࠿ࡽᰤ㣴ࢆዣ࠺⊂≉࡞⏕Ꮡᡓ␎ࢆ࡜ࡿᐤ⏕᳜≀ࡣ㸪⿕Ꮚ᳜≀ࡢ⣔⤫ᶞୖ࡟ᗈࡃศᕸࡋ࡚࠾ࡾ㸪ᑡ࡞ࡃ࡜ࡶ⊂❧࡟ 11 ᅇ㐍໬ࡋࡓ࡜⪃࠼ࡽࢀ࡚࠸ࡿ㸦Barkman et al., 2007㸧ࠋࡑࡢ୰࡛ࡶ㸪ࣁ࣐࢘ࢶ࣎⛉࡟ᒓࡍࡿ᰿ᐤ⏕᳜≀ࡣ⢭ຊⓗ࡟◊✲ࡀ㐍ࡵࡽࢀ࡚࠸ࡿࠋࡑࡢ⌮⏤ࡢ୍ࡘࡣ㸪㎰ᴗ࡟୚࠼ࡿ⤒῭ⓗ࡞ᦆᐖࡀ㠀ᖖ࡟኱ࡁ࠸࠿ࡽ࡛࠶ࡿࠋࣁ࣐࢘ࢶ࣎⛉࡟ᒓࡍࡿ Strigaᒓ࡜ Orobanche ᒓࡣࡉࡲࡊࡲ࡞㎰ᴗస≀࡟ᐤ⏕ࡋ㸪཰㔞ࢆ኱ᖜ࡟ῶ཰ࡉࡏࡿࠋ࡜ࡾࢃࡅ࢔ࣇ
࡛ࣜ࢝ࡢ Striga ᒓ࡟ࡼࡿ⿕ᐖࡣ῝้࡛㸪25 ࣨᅜྜࢃࡏ࡚ 1 ൨ࡶࡢேࡀᙳ㡪ࢆཷࡅ㸪ࡑࡢ⿕ᐖ㢠ࡣ୍ᖺ࡛ 10 ൨ࢻࣝ࡟ࡶୖࡿ࡜᥎ィࡉࢀ࡚࠸ࡿ㸦reviewed in Spallek et al., 2013㸧ࠋ⌧ᅾ㸪ࡇࢀࡽᐤ⏕᳜≀ࡢຠᯝⓗ࡞㝖ཤᡭẁࡣ࡞ࡃ㸪᪩ᛴ࡞ゎỴ᪉ἲࡢ㛤Ⓨࡀᮃࡲࢀ࡚࠸ࡿࠋࡶ࠺୍ࡘ
ࡢ⌮⏤࡜ࡋ࡚ࡣ㸪ࣁ࣐࢘ࢶ࣎⛉࡟ࡣ␗࡞ࡿᐟ୺౫Ꮡᗘࡢᐤ⏕᳜≀ࡀᒓࡋ࡚࠸ࡿ࡜࠸࠺ࡇ࡜ࡀ

BSJ-Review 7:241 (2016)

植物科学最前線 7:242 (2016)࠶ࡿࠋᐤ⏕᳜≀ࡣ㸪ᐟ୺࡬ࡢ౫Ꮡᗘྜ࠸࡟ࡼࡗ࡚㸪᮲௳ⓗ༙ᐤ⏕㸦ගྜᡂ⬟ࢆᣢࡕ⊂❧ࡋ࡚⏕ά࡛ࡁࡿࡀ㸪ᐟ୺᳜≀ࡀ㏆ഐ࡟࠸ࡿሙྜ࡟ࡣᐤ⏕ࢆ㛤ጞࡍࡿ㸧㸪⤯ᑐ༙ᐤ⏕㸦ගྜᡂ⬟ࢆᣢࡘࡀ㸪⏕ά࡟ࡣᐟ୺ࡢᏑᅾࡀ୙ྍḞ㸧㸪⤯ᑐ඲ᐤ⏕㸦ගྜᡂ⬟ࢆࡶࡓࡎ㸪ᰤ㣴ࡣ᏶඲࡟ᐟ୺౫Ꮡ㸧࡜ศ㢮ࡉࢀࡿࠋࣁ࣐࢘ࢶ࣎⛉࡟ࡣ㸪᮲௳ⓗ༙ᐤ⏕࡟ศ㢮ࡉࢀࡿ Tryphysaria ᒓ 㸪⤯ᑐ༙ᐤ⏕࡟ศ㢮ࡉࢀࡿ Striga ᒓ㸪⤯ᑐ඲ᐤ⏕࡟ศ㢮ࡉࢀࡿ Orobanche ᒓ࡞࡝ࡍ࡭࡚ࡢࢡࣛࢫࡢᐤ⏕᳜≀ࡀࡑࢁࡗ࡚࠸ࡿࠋࡇࢀࡽ࡟ຍ࠼࡚㸪㠀ᐤ⏕᳜≀࡛࠶ࡿ Lindenbergia ᒓࡀ࠶ࡿࡢ࡛㸪ᐤ⏕ᙧ㉁ࡢ⋓ᚓ࠿ࡽගྜᡂ⬟ࢆኻ࠺ࡲ࡛ࡢ㐍໬ࡢ㐣⛬ࢆ㸪ྠࡌ⛉࡟ᒓࡍࡿ⌧⏕✀ࢆ⏝࠸࡚◊✲ࡍࡿࡇ࡜ࡀ࡛ࡁࡿࠋࡇࡢࡇ࡜࠿ࡽ㸪㐍໬ࡢ◊✲ࡢ㠀ᖖ࡟Ⰻ࠸ࣔࢹࣝ࡜࡞ࡗ࡚࠸ࡿࠋ⚾ࡓࡕࡢ◊✲ࢢ࣮ࣝࣉ࡛ᢅࡗ࡚࠸ࡿࡢࡣ୺࡟㸪ࢫࢺࣛ࢖࢞㸦Striga hermonthica㸪Striga asiatica㸧࡜ࢥࢩ࣐࢜࢞㸦Phtheirospermum japonicum㸧 ࡢ 3 ✀࡛㸪࠸ࡎࢀࡶࣁ࣐࢘ࢶ࣎⛉࡟ᒓࡍࡿᐤ⏕᳜≀࡛࠶ࡿࠋࣁ࣐࢘ࢶ࣎⛉᳜≀ࢆྵࡵࡓᐤ⏕᳜≀඲⯡࡟ඹ㏻ࡍࡿࡢࡣ㸪௚᳜≀࡟௜╔ࡋ㸪ࡑࡢ⤌⧊࡟౵ධࡍࡿࡓࡵࡢ≉ู࡞ჾᐁࢆⓎ㐩ࡉࡏ࡚ࡁࡓࡇ࡜࡛࠶ࡿࠋࡇࢀࡽࡢჾᐁࢆ⥲⛠ࡋ࡚ࠕ྾ჾ
㸦haustorium㸧ࠖ࡜࿧ࢇ࡛࠸ࡿࠋࣁ࣐࢘ࢶ࣎⛉ᐤ⏕᳜≀ࡣ㸪⮬ࡽࡢ᰿ࡢ୍㒊ࢆ྾ჾ࡬࡜ኚᙧࡋᐟ୺᳜≀࡟ᐤ⏕ࡍࡿࠋࡇࡢ⥲ㄝ࡛ࡣ᰿⣔ᵓ⠏ࡢ୍౛࡜ࡋ࡚㸪᰿ᐤ⏕᳜≀ࡢ྾ჾᙧᡂ࡟ࡘ࠸࡚㸪ࡇࢀࡲ࡛ࡢ▱ぢࢆᴫㄝࡋࡓᚋ㸪⚾ࡓࡕࡢ◊✲ࢢ࣮ࣝࣉ࡛ࡢ᭱㏆ࡢ◊✲ᡂᯝ࡟ࡘ࠸࡚⤂௓ࡍࡿࠋ㸰㸬ࣁ࣐࢘ࢶ࣎⛉ᐤ⏕᳜≀ࢆ⏝࠸ࡓ྾ჾࡢ◊✲㸰㸫㸯㸬྾ჾㄏᑟ࣓࢝ࢽࢬ࣒ຠ⋡ࡼࡃᐤ⏕ࢆ⾜࠺࡟ࡣᐟ୺᳜≀ࡢᏑᅾࢆ㐺ษ࡟ㄆ㆑ࡍࡿᚲせࡀ࠶ࡿࠋ⤯ᑐᐤ⏕᳜≀࡟࡜ࡗ࡚ᐟ୺ࡣᚲせ୙ྍḞ࡞Ꮡᅾ࡞ࡢ࡛㸪ᐟ୺ࡢᏑᅾࡋ࡞࠸࡜ࡇࢁ࡛Ⓨⱆࡍࡿࡇ࡜ࡣ⮬ẅ⾜Ⅽ࡜࡞ࡗ࡚ࡋࡲ࠺ࠋᐤ⏕᳜≀ࡢᣢࡘᐟ୺᳜≀ㄆ㆑ᶵᵓ࡜ࡋ࡚㸪ᐟ୺᳜≀ࡢศἪࡍࡿࢫࢺࣜࢦࣛࢡ
ࢺࣥࢆ⏝࠸ࡓⓎⱆไᚚࡀ࠶ࡿࠋᯞศ࠿ࢀࢆไᚚࡍࡿ᳜≀࡛࣍ࣝࣔࣥ࠶ࡿࢫࢺࣜࢦࣛࢡࢺࣥࡣ㸪ࡶ࡜ࡶ࡜ࢫࢺࣛ࢖࢞ࡢ✀ᏊࡢⓎⱆࢆㄏᑟࡍࡿ≀㉁㸦Strigol㸧࡜ࡋ࡚㸪࣡ࢱࡢ᰿࠿ࡽྠᐃࡉࢀࡓ㸦Cook et al., 1966㸧ࠋᅵተ୰࡛ࡣ୙Ᏻᐃ࡞ࢫࢺࣜࢦࣛࢡࢺࣥࢆⓎⱆࡢࢩࢢࢼࣝ࡜ࡋ࡚฼⏝ࡍࡿࡇ࡜࡛㸪ᐟ୺᳜≀ࡢࡈࡃ㏆ഐ࡛ࡢⓎⱆࡀྍ⬟࡜࡞ࡗ࡚࠸ࡿࠋྠᵝ࡟㸪௚ࡢ᳜≀ࡀ࠸࡞࠸࡜ࡇࢁ࡟྾ჾࢆᙧᡂࡋ࡚ࡶព࿡ࡀ࡞ࡃ㸪࡝ࡇ࡟௚ࡢ᳜≀ࡀᏑᅾࡍࡿ࠿ࢆ㐺ษ࡟ㄆ㆑ࡍࡿࡇ࡜ࡣ྾ჾᙧᡂ࡟࠾࠸࡚ࡶ㔜せ࡛࠶ࡿࠋᐤ⏕᳜≀ࡀ࡝ࡢࡼ࠺࡟ᐟ୺᳜≀ࢆㄆ㆑ࡋ྾ჾࢆᙧᡂࡍࡿࡢ࠿ㄪ࡭ࡿࡓࡵ࡟㸪᰿ࡢⁱฟᾮࡸᢳฟᾮ࠿ࡽ྾ჾㄏᑟ≀㉁㸦HIF: haustorium inducing factor㸧ࢆ༢㞳㸪ྠᐃࡍࡿ◊✲ࡀ⾜࡞ࢃࢀ࡚ࡁࡓࠋࡑࡢ⤖ᯝ㸪ࢯ࣒ࣝ࢞ࡢ᰿ࡢᢳฟᾮ࠿ࡽ 2,6-dimethoxy-1,4-benzoquinone 㸦DMBQ㸧 ࡀS. asiatica ࡢ྾ჾࢆㄏᑟࡍࡿ≀㉁࡜ࡋ࡚ྠᐃࡉࢀࡓ㸦Chang and Lynn, 1986㸧ࠋ୍⯡࡟࣋ࣥࢰ࢟ࣀࣥ㢮ࡣ㸪᳜≀ෆ࡛ࢩ࣑࢟㓟⤒㊰㸪ࣇ࢙ࣀ࣮ࣝ㓟ࡢ㓟໬ⓗ⬺Ⅳ㓟㸪࣌ࣝ࢜࢟ࢩࢲ࣮ࢮࡸࣛࢵ࣮࢝ࢮ࡟ࡼࡿ⣽⬊ቨࣇ࢙ࣀ࣮ࣝࡢศゎ࡞࡝࡟ࡼࡗ࡚⏕ࡌࡿ㸦Caldwell and Steelink, 1969㸧ࠋࡋ࠿ࡋ㸪DMBQࡀࢯ࣒ࣝ࢞ࡢ᰿࠿ࡽ᳨ฟࡉࢀࡓࡢࡣ㸪ࢯ࣒ࣝ࢞ࡢ᰿ࢆ≀⌮ⓗ࡟☻ࡾ₽ࡋࡓ᫬࠿㸪ࢫࢺࣛ࢖࢞࡜ඹᇵ㣴ࡋࡓ᫬ࡔࡅ࡛࠶ࡗࡓ㸦Chang and Lynn, 1986㸧ࠋࡇࡢࡇ࡜࠿ࡽ㸪ᐤ⏕᳜≀ࡀືⓗ࡟HIFࡢ⏕ᡂࢆไᚚࡍࡿࡇ࡜࡛㸪ᐟ୺᳜≀ࡢ㏆ഐ࡛ࡢ྾ჾᙧᡂࢆྍ⬟࡟ࡋ࡚࠸ࡿࡇ࡜ࡀ⪃࠼ࡽࢀࡓࠋࡑࡢᚋ
ࡢ◊✲࠿ࡽ㸪ࢫࢺࣛ࢖࢞᰿➃࡛⏕ᡂࡉࢀࡓH 2 O 2 ࡀᐟ୺᳜≀ࡢ࣌ࣝ࢜࢟ࢩࢲ࣮ࢮࢆάᛶ໬ࡋ㸪

BSJ-Review 7:242 (2016)

植物科学最前線 7:243 (2016)ࡇࡢ࣌ࣝ࢜࢟ࢩࢲ࣮ࢮࡀ⣽⬊ቨࡢࣇ࢙ࣀ࣮ࣝࢆ㓟໬ࡍࡿࡇ࡜࡛HIFࡀ⏕ᡂࡉࢀࡿ࡜࠸࠺࣓࢝
ࢽࢬ࣒ࡀᥦၐࡉࢀ࡚࠸ࡿ㸦Kim et al., 1998, Keyes et al., 2007㸧ࠋDMBQ ࡀ྾ჾㄏᑟ⬟ࢆࡶࡘ≀㉁࡜ࡋ࡚ྠᐃࡉࢀࡓࡇ࡜࠿ࡽ㸪௚ࡢࣇࣛ࣎ࣀ࢖ࢻࡸ࢟ࣀࣥ࡞࡝ࡢࣇ࢙ࣀ࣮ࣝㄏᑟయࡀྠࡌࡼ࠺࡟྾ჾㄏᑟάᛶࢆᣢࡘ࠿࡝࠺࠿ࢸࢫࢺࡉࢀࡓࠋࡑࡢ⤖ᯝ㸪άᛶ࡟ᕪࡣ࠶ࡿࡶࡢࡢ㸪ࢩࣜࣥ࢞㓟ࡸࣂࢽࣜࣥ㓟ࡸࢡ࣐ࣝ㓟࡞࡝ࡢ༢⣧࡞ᵓ㐀ࢆᣢࡘࣇ࢙ࣀ࣮ࣝ㢮㸪࣌࢜ࢽࢪࣥࡸ࣌ࣛࣝࢦࢽࢪࣥ࡞࡝ࡢࣇࣛ࣎ࣀ࢖ࢻࡶ྾ჾࢆㄏᑟ࡛ࡁࡿࡇ࡜ࡀศ࠿ࡗ
ࡓ㸦Albrecht et al., 1999㸧
ࠋDMBQ ࢆྵࡴ࣋ࣥࢰ࢟ࣀࣥࡢ࢔ࢼࣟࢢࡢ࠺ࡕ㸪྾ჾㄏᑟάᛶࢆᣢࡘࡶࡢࡀ≉ᐃࡢ⠊ᅖࡢ㓟໬㑏ඖ㟁఩ࢆᣢࡘࡇ࡜࠿ࡽ㸪HIF ࢩࢢࢼࣜࣥࢢ࡟ࡣ㓟໬㑏ඖࢧ࢖ࢡࣝࡀ㛵ࢃࡗ࡚࠸ࡿࡇ࡜ࡀ♧၀ࡉࢀ࡚࠸ࡿ㸦Smith et al., 1996㸧ࠋࣁ࣐࢘ࢶ࣎⛉᮲௳ⓗ༙ᐤ⏕᳜≀ࡢ Tryphysaria versicolor ࠿ࡽ༢㞳ࡉࢀࡓ࢟ࣀࣥ㑏ඖ㓝⣲ࢆࢥ࣮ࢻࡍࡿ TvQR1 ࢆࣀࢵࢡࢲ࢘ࣥࡋࡓ㝿㸪ㄏᑟࡉࢀࡿ྾ჾࡢᩘࡀῶᑡࡍࡿࡇ࡜ࡀ♧ࡉࢀࡓ㸦Bandaranayake et al., 2010㸧ࡇ࡜࠿ࡽ㸪྾ჾᙧᡂࢆ㛤ጞࡍࡿࢩࢢࢼࣝ࡜ࡋ࡚㸪㓟໬㑏ඖࢩࢢࢼࣝࡀ㛵ࢃࡗ࡚࠸ࡿࡇ࡜ࡀ⌧ᅾࡢࣔࢹࣝ࡜࡞ࡗ࡚࠸ࡿࠋ㸰㸫㸰㸬྾ჾࡢⓎ⏕࡜⤌⧊ᏛA

ᅗ 1B

Pedicularis sylvatica 㸦ࢩ࣐࢜࢞ࢠࢡᒓ㸧࡟࠾ࡅࡿ྾ჾᶓ᩿ษ∦ࡢᶍᘧᅗ (A)ྑഃ

࡟ྥ࠿ࡗ࡚ఙࡧࡿ྾ჾࡢ඲యീࠋ(B)ᐟ୺࡜ࡢ࢖ࣥࢱ࣮ࣇ࢙࢖ࢫ㒊ศࡢᣑ኱ᅗࠋSablon ML du, 1887
Figure 9, 11 ࡼࡾ෌ᥖࠋ⦤ᶍᵝࢆᣢࡘ⣽⬊ࡣ㐨⟶せ⣲ࢆ⾲ࡍࠋᐤ⏕᳜≀ࡀ྾ჾࢆ௓ࡋ࡚࡝ࡢࡼ࠺࡟ᐟ୺᳜≀ࡢ⤌⧊࡟౵ධࡍࡿ࠿ࢆ⌮ゎࡍࡿࡓࡵ㸪྾ჾࢆᵓᡂࡍࡿ⣽⬊ࡢ≉ᛶࡸᵓ㐀㸪ࡑࡢⓎ⏕࡟ࡘ࠸࡚ࡢ◊✲ࡀ⾜࡞ࢃࢀ࡚ࡁࡓࠋ1887 ᖺ࡟ࡣ᪤࡟ࢩ࣐࢜࢞ࢠࢡᒓࡢ༙ᐤ⏕᳜≀࡟࠾ࡅࡿ྾ჾࡢᵓ㐀ࡀ◊✲ࡉࢀ࡚࠾ࡾ㸪㐨⟶せ⣲ࡀᐤ⏕᳜≀ࡢ⥔⟶᮰࠿ࡽᐟ୺᳜≀ࡢ᪉ྥ࡬࡜㐃࡞ࡗ࡚࠸ࡿᵝᏊࡀᥥ࠿ࢀ࡚࠸ࡿ㸦ᅗ 1A㸧
㸦Sablon ML du, 1887㸧ࠋࡇࡢ྾ჾࡢ୰࡟సࡽࢀࡿ㸪ᐤ⏕᳜≀ࡢ⥔⟶᮰࡜ᐟ୺᳜≀ࡢ⥔⟶᮰ࢆࡘ࡞ࡄᵓ㐀ࡣxylem bridge࡜࿧ࡤࢀ࡚࠾ࡾ㸪௚ࡢ⛉ࡢᐤ⏕᳜≀ࡀᐤ⏕ࡍࡿ㝿࡟ࡶࡋࡤࡋࡤほᐹࡉࢀࡿࠋࡲࡓ㸪xylem bridge࡟ἢࡗ࡚ࢧ࢖ࢬࡢᑠࡉ࠸⣽⬊ࡀศ໬ࡋ࡚࠾ࡾ㸪≉Ṧ࡞⣽⬊✀࡛࠶ࡿྍ⬟ᛶࢆ♧၀ࡋ࡚࠸ࡿࠋᐟ୺᳜≀࡜᥋ࡍࡿ㒊ศ࡟ࡣ㛗ࡃఙࡧࡓ⊂≉࡞ᙧ≧ࢆࡋࡓ⣽⬊ࡀᥥ࠿ࢀ࡚࠸ࡿ㸦ᅗ 1B㸧
㸦SablonML du, 1887㸧ࠋᐟ୺࡜ࡢ࢖ࣥࢱ࣮ࣇ࢙࢖ࢫ㒊ศ࡟ࡣ≉Ṧ࡞⣽⬊ࡀศ໬ࡋ࡚ࡃࡿࡇ࡜ࡀ▱ࡽࢀ

BSJ-Review 7:243 (2016)

植物科学最前線 7:244 (2016)࡚࠾ࡾ㸪ࡇࢀࡽࡢ⣽⬊ࡢ࠺ࡕ୍㒊ࡣᐟ୺ࡢ㐨⟶せ⣲ࡢ⣽⬊ቨࡢࣆࢵࢺ㒊ศࢆ㈏㏻ࡋࡓᚋ࡟㐨⟶せ⣲࡬࡜ศ໬ࡍࡿࡇ࡜࡛㸪㐨⟶せ⣲ྠኈࡢ᏶඲࡞㐃⤖ࢆྍ⬟࡜ࡋ࡚࠸ࡿࡇ࡜ࡀ Strigaasiatica ࡛♧ࡉࢀ࡚࠸ࡿ㸦Dörr, 1997㸧ࠋ྾ჾࡢⓎ⏕ึᮇ࡟ࡘ࠸࡚ࡣ㸪ྠࡌࣁ࣐࢘ࢶ࣎⛉ࡢ Agalinis purpurea ࡜࠸࠺༙ᐤ⏕᳜≀ࢆ౑ࡗ࡚ヲ⣽࡟ㄪ࡭ࡽࢀ࡚࠸ࡿ㸦Baird and Riopel, 1984㸧ࠋHIF ࡟᭱ࡶ㧗࠸ឤཷᛶࢆ♧ࡍࡢࡣ᰿➃ࡢ࣓ࣜࢫࢸ࣒㡿ᇦ࡜ఙ㛗㡿ᇦࡀษࡾ᭰ࢃࡿ㡿ᇦ࡛࠶ࡾ㸪᭱ึ࡟ᙧែୖࡢኚ໬ࡀほᐹࡉࢀࡿࡢࡣෆഃࡢ⓶ᒙ࡛࠶ࡿࠋ⓶ᒙ⣽⬊ࡣᾮ⬊໬ࡋ㸪ᨺᑕ᪉ྥ࡬࡜య✚ࢆቑ኱ࡍࡿࠋ᰿ࡀഃ᪉ྥ࡬࡜⭾ࡽࡴࡢࡀ┠࡛☜ㄆ࡛ࡁࡿࡼ࠺࡟࡞ࡿ㡭㸪⾲⓶⣽⬊࡛ࡣᆶᒙศ⿣ࡀ㉳ࡇࡾ㸪ෆഃࡢ⓶ᒙࡸෆ㠧⣽⬊࡞࡝ࡢ῝㒊࡛ࡣࡲࡎ୪ᒙศ⿣ࡀ㉳ࡇࡿࠋࡑࡢᚋ㸪⣽⬊ศ⿣ࢆ⧞ࡾ㏉ࡋࡇࡪ≧ࡢ྾ჾࢆᙧᡂࡋ࡚࠸ࡃࡀ㸪⣽⬊ᩘࡀከࡃ࡞ࡗ࡚࠸ࡃࡓࡵ㸪୪ᒙศ⿣࠿ᆶᒙศ⿣࠿ࡢ༊ูࡣ㞴ࡋࡃ࡞ࡗ࡚࠸ࡿࠋࡓࡔࡋ㸪⾲⓶ཬࡧእഃࡢ⓶ᒙࡣ HIF ฎ⌮ᚋ 48 ᫬㛫ࡲ࡛ࡣ୪ᒙศ⿣ࢆࡏࡎ㸪ᒙࢆಖࡗ࡚࠸ࡿᵝᏊࡀほᐹࡉࢀ࡚࠸ࡿ㸦Baird and Riopel, 1984㸧ࠋ྾ჾࡢⓎ⏕࡟ࡣ྾ჾẟ(haustorialhair) ࡜࿧ࡤࢀࡿ᰿ẟ⣽⬊࡜ఝࡓᵓ㐀ࢆᣢࡘ⣽⬊ࡢศ໬ࡀక࠺ࠋࡇࡢ⣽⬊ࡣ⢓╔≀㉁ࢆศἪࡋ㸪ᐟ୺᳜≀࡬ࡢᐦ╔࡟ᶵ⬟ࡋ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ㸦Baird and Riopel, 1983, Heidge-Jorgensenand Kujit, 1995㸧
ࠋ᭱㏆࡟࡞ࡗ࡚㸪ࢥࢩ࣐࢜࢞ኚ␗యࡢゎᯒ࠿ࡽ㸪ࡇࡢ྾ჾẟࡢⓎ⏕ࡣ᰿ẟ࡜ྠࡌⓎ⏕ࣉࣟࢢ࣒ࣛࢆ௓ࡋ࡚࠸ࡿࡇ࡜ࡀ㑇ఏᏛⓗ࡟ド᫂ࡉࢀࡓ㸦Cui et al., 2016㸧ࠋ⯆࿡῝࠸ࡇ࡜࡟㸪ࢥࢩ࣐࢜࢞ࡢ᰿ẟኚ␗యࡣ྾ჾẟࢆసࡿࡇ࡜ࡣ࡛ࡁ࡞࠸ࡀ㸪྾ჾࢆᙧᡂࡋᐟ୺ෆ࡟౵ධࡍࡿࡇ࡜ࡀ࡛ࡁࡿࠋኚ␗య࡟࠾ࡅࡿ྾ჾࡢෆ㒊ᵓ㐀ࡣ㔝⏕ᆺ࡜ኚࢃࡾ࡞ࡃ㸪࢖ࣥࢱ࣮ࣇ࢙࢖ࢫ㒊ศࡢఙ㛗ࡋࡓ⣽⬊ᒙࡢᙧᡂࡣ᰿ẟࡢఙ㛗࡜ࡣ␗࡞ࡿࣉࣟࢢ࣒࡛ࣛไᚚࡉࢀ࡚࠸ࡿࡇ
࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋ྾ჾࡢᙧᡂࡣ㸪᪤ᏑࡢⓎ⏕ࣉࣟࢢ࣒ࣛࡢὶ⏝࡜ᐤ⏕᳜≀࡟ࣘࢽ࣮ࢡ࡞㑇ఏⓗࣉࣟࢢ࣒ࣛࡢ⋓ᚓࡀ⤌ࡳྜࢃࡉࢀࡿࡇ࡜࡛ᡂ❧ࡋࡓ࡜⪃࠼ࡽࢀࡿࠋ㸰㸫㸱㸬྾ჾࡢࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒ࢔࣓ࣜ࢝ࡢ◊✲ࢳ࣮࣒ࢆ୰ᚰ࡜ࡋ࡚ Parasitic Plant Genome Project (http://ppgp.huck.psu.edu/)ࡀ❧ࡕୖࡆࡽࢀ࡚࠸ࡿࠋࡇࡢࣉࣟࢪ࢙ࢡࢺ࡛ࡣ㸪ᐤ⏕ᙧ㉁⋓ᚓ࡟ᐤ୚ࡋࡓኚ໬࡜㸪ᐤ⏕ᙧ㉁ࡢ⋓ᚓࡢ⤖ᯝ㉳ࡇࡗࡓኚ໬ࢆࢤࣀ࣒࣡࢖ࢻ࡟᫂ࡽ࠿࡟ࡍࡿࡇ࡜ࢆ┠ⓗ࡜ࡋ࡚㸪ẚ㍑ࢤࣀ࣒ゎᯒࡀ⾜ࢃࢀ࡚࠸ࡿࠋලయⓗ࡟ࡣ㸪ࣁ࣐࢘ࢶ࣎⛉࡟ᒓࡍࡿ㠀ᐤ⏕᳜≀࡛࠶ࡿ Lindenbergiaᒓ㸪᮲௳ⓗ༙ᐤ⏕᳜≀ࡢTriphysariaᒓ㸪⤯ᑐ༙ᐤ⏕᳜≀ࡢStrigaᒓ㸪⤯ᑐ඲ᐤ⏕᳜≀ࡢOrobancheᒓ࡞࡝ࡢࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒࡀ⢭ຊⓗ࡟࡞ࡉࢀ࡚࠸ࡿࠋࡇࡢࣉࣟࢪ࢙ࢡࢺ୰࡛ࡣ྾ჾࡀᐤ⏕ࢆࡍࡿ࠺࠼࡛㔜せ࡞ᙺ๭ࢆᢸ࠺࡜ࡋ㸪྾ჾࡢⓎ⏕ẁ㝵ࡈ࡜ࡢࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ࢹ࣮ࢱࢆᚓࡿࡇ࡜࡟↔Ⅼࢆᙜ࡚࡚࠸ࡿࠋࡉࡲࡊࡲ࡞ᐟ୺౫Ꮡᗘࢆᣢࡘ᳜≀✀ࢆ౑ࡗࡓẚ㍑ࢺࣛ
ࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒ࠿ࡽ㸪ᐟ୺࡜ࡢ᥋ゐᚋ࡟Ⓨ⌧ࡀୖ᪼ࡍࡿ 180 㑇ఏᏊࡀྠᐃࡉࢀ࡚࠾ࡾ㸪ࡇࢀࡽࡢ㑇ఏᏊ⩌࡟ࡣࣉࣟࢸ࢔࣮ࢮ㸪⣽⬊ቨಟ㣭㓝⣲㸪⣽⬊እศἪࢱࣥࣃࢡ㉁ࡀ≉࡟ከࡃྵࡲࢀ࡚࠸ࡿࠋᐟ୺࡜᥋ゐࡍࡿ๓ࡢẁ㝵࡛㸪HIF࡟ᛂ⟅ࡍࡿ 100 㑇ఏᏊࡶྠᐃࡋ࡚࠾ࡾ㸪๓ࡢ
180 㑇ఏᏊ࡜ྜࢃࡏ࡚ࣁ࣐࢘ࢶ࣎⛉ᐤ⏕᳜≀࡟࠾ࡅࡿ“parasitism genes”࡜ᐃ⩏ࡉࢀ࡚࠸ࡿ
㸦Yang et al., 2014㸧ࠋ㸱㸬ࣔࢹ᳜ࣝ≀࡜ࡋ࡚ࡢࢥࢩ࣐࢜࢞

BSJ-Review 7:244 (2016)

植物科学最前線 7:245 (2016)

ABCDᅗ 2 ࢥࢩ࣐࢜࢞㸦Phtheirospermum japonicum㸧(A)᳜≀య㸪(B)(C)ⰼ㸪(D)ᯝᐇ
⚾ࡓࡕࡢ◊✲ࢢ࣮ࣝࣉ࡛ࡣࢥࢩ࣐࢜࢞ࢆࣁ࣐࢘ࢶ࣎⛉ᐤ⏕᳜≀ࡢ྾ჾ◊✲ࡢࣔࢹ᳜ࣝ≀࡜ࡋ࡚࠸ࡿ㸦ᅗ 2㸧ࠋࢥࢩ࣐࢜࢞ࡣࣁ࣐࢘ࢶ࣎⛉ࢥࢩ࣐࢜࢞ᒓࡢ᮲௳ⓗ༙ᐤ⏕᳜≀࡛㸪ᮾ࢔ࢪ࢔࡟⮬⏕ࡍࡿࠋ᪥ᮏᅜෆ࡛ࡶぢࡘࡅࡿࡇ࡜ࡀ࡛ࡁ㸪ⰼࡢⰍࡢ␗࡞ࡿ࢚ࢥࢱ࢖ࣉࡶほᐹࡉࢀ࡚࠸ࡿࠋࢥࢩ࣐࢜࢞ࡣ⮬ᐙཷ⢊ࢆࡍࡿ▷᪥᳜≀࡛㸪ேᕤẼ㇟ჾࡢ୰࡛ᐜ᫆࡟᱂ᇵࡍࡿࡇ࡜ࡀ࡛ࡁ㸪⣙ 3 ࣨ᭶࡛ḟୡ௦ࡢ✀Ꮚࡀᚓࡽࢀࡿࠋேᕤⓗ࡟஺㓄ࡉࡏࡿ᪉ἲࡶ☜❧ࡋ࡚࠾ࡾ㸪㑇ఏᏛ࡟㐺ࡋ࡚࠸ࡿࠋ⚾ࡓࡕࡢ◊✲ࢢ࣮ࣝࣉ࡛ࡣ㸪ᒸᒣ┴࡛᥇ྲྀࡉࢀࡓࢥࢩ࣐࢜࢞ࢆ◊✲ᐊ⎔ቃ࡛ᩘୡ௦⮬Ṫࡉࡏ㸪㔝⏕ᆺ࡜ࡋ࡚◊✲࡟౑⏝ࡋ࡚࠸ࡿࠋࡇࡢ㔝⏕ᆺࢆࡶ࡜࡟㸪EMS ࢆኚ␗ཎ࡜ࡋࡓࢫࢡ࣮ࣜࢽࣥࢢ࠿ࡽ㸪྾ჾ࡟ࡉࡲࡊࡲ࡞⾲⌧ᆺࢆ♧ࡍኚ␗యࡢ༢㞳࡟ࡶᡂຌࡋ࡚࠸ࡿ㸦Cui etal., 2016㸧ࠋᙜ◊✲ࢢ࣮ࣝࣉ࡛ࡣࢤࣀ࣒ࡢゎᯒࡶ㐍ࢇ࡛࠾ࡾ㸪ࢤࣀ࣒ࣜࢩ࣮ࢣࣥࢫ࡟ࡼࡿኚ␗యཎᅉ㑇ఏᏊࡢྠᐃࡶྍ⬟࡜࡞ࡗ࡚࠸ࡿࠋࡲࡓ㸪Agrobacterium rhizogenes ࢆ⏝࠸ࡓᙧ㉁㌿᥮ἲࢆ☜❧ࡋ࡚࠾ࡾ㸪ㄏᑟࡉࢀࡓẟ≧᰿ࢆ⏝࠸ࡓศᏊ⏕≀Ꮫⓗゎᯒࢆ࡜ࡶ࡞ࡗࡓᐤ⏕ᐇ㦂ࡶྍ⬟࡛࠶ࡿ㸦Ishida et al., 2011㸧ࠋࢥࢩ࣐࢜࢞ࡣ T. versicolor ࡜ྠᵝ࡟㸪ᖜᗈ࠸᳜≀✀ࢆᐟ୺࡜ࡍࡿࡇ࡜ࡀ࡛ࡁࡿࠋࡇࢀࡲ࡛ࡢ࡜ࡇࢁ㸪ࢩࣟ࢖ࢾࢼࢬࢼ㸪࢖ࢿ㸪ࢯ࣒ࣝ࢞㸪ࢺ࣐ࢺ㸪ࢧࢧࢤ㇋࡟ᐤ⏕࡛ࡁࡿࡀ㸪࣑ࣖࢥࢢࢧ࡜ࢲ࢖ࢬ࡟ࡣᐤ⏕࡛ࡁ࡞࠸ࡇ࡜ࢆ☜ㄆࡋ࡚࠸ࡿࠋ྾ჾࡢⓎ⏕㐣⛬ࡣ㸪ඛ⾜◊✲ࡢ࠶ࡿࣁ࣐࢘ࢶ࣎⛉᮲௳ⓗ༙ᐤ⏕᳜≀࡛࠶ࡿ A. purpurea ࡸ T. versicolor࡜࡯࡜ࢇ࡝ྠࡌ࡛࠶ࡿࠋᐟ୺᳜≀㸪ࡲࡓࡣ DMBQ ࡞࡝ࡢ HIF ࢆ⏝࠸࡚ in vitro ࡛྾ჾࡢㄏᑟࡀ࡛ࡁࡿ㸦ᅗ 3㸧ࠋ

BA

ᅗ 3

ࢥࢩ࣐࢜࢞྾ჾ(A)ࢩࣟ࢖ࢾࢼࢬࢼ᰿㸦ྑ㸧࡟ᐤ⏕ࡍࡿࢥࢩ࣐࢜࢞྾ჾ㸦ᕥ㸧ࠋࢧࣇࣛࢽࣥᰁⰍ࡛㐨⟶せ⣲ࡀ㉥ࡃᰁࡲࡗ࡚ぢ࠼ࡿࠋ(B)DMBQ ࡟ࡼࡗ࡚ㄏ
ᑟࡉࢀࡓࢥࢩ࣐࢜࢞྾ჾࠋBars =100 µm㸲㸬྾ჾᙧᡂ࡜࣮࢜࢟ࢩࣥ
indole-3-acetic acid (IAA) ࡟௦⾲ࡉࢀࡿ᳜≀࣍ࣝࣔࣥࡢ࣮࢜࢟ࢩࣥࡣ㸪᳜≀࡟࠾ࡅࡿჾᐁⓎBSJ-Review 7:245 (2016)

植物科学最前線 7:246 (2016)⏕࡛㔜せ࡞ᙺ๭ࢆᣢࡘࡇ࡜ࡀ▱ࡽࢀ࡚࠸ࡿࠋ౛࠼ࡤ㸪࣮࢜࢟ࢩࣥ⃰ᗘ໙㓄ࡢᴟ኱Ⅼࡀⱼ㡬ศ⿣⤌⧊࡛ࡣⴥཎᇶࡢ㸪᰿ࡢෆ㠧⣽⬊࡛ࡣഃ᰿ཎᇶࡢⓎ⏕ࣃࢱ࣮ࣥࢆไᚚࡋ࡚࠸ࡿࠋࡑࡋ࡚㸪᳜≀యෆ࡛ࡢ࣮࢜࢟ࢩࣥࡢ⃰ᗘ໙㓄ࡣ㸪⣽⬊㛫㍺㏦࡜⏕ྜᡂࡸ␗໬࡞࡝ࡢ௦ㅰ࡟ࡼࡗ࡚ไᚚࡉࢀ࡚࠸ࡿࠋࣁ࣐࢘ࢶ࣎⛉ᐤ⏕᳜≀ࢆ౑ࡗࡓඛ⾜◊✲࠿ࡽ㸪࣮࢜࢟ࢩࣥࡀ྾ჾࡢᙧᡂ࡟㛵ࢃࡿࡇ࡜ࡀ♧၀ࡉࢀ࡚࠸ࡿࠋ࣮࢜࢟ࢩࣥ㍺㏦㜼ᐖ๣㸪ᢠ࣮࢜࢟ࢩࣥ๣࡜㐣๫㔞ࡢ࣮࢜࢟ࢩࣥࡀᐤ⏕ຠ⋡ࢆῶᑡࡉࡏࡿࡇ࡜ࡀ᏶඲ᐤ⏕᳜≀ࡢ Orobanche aegyptiaca ࡛♧ࡉࢀ࡚࠸ࡿ㸦Bar-Nunet al., 2008㸧
ࠋT. versicolor ࡛ࡣ㸪࣮࢜࢟ࢩࣥ㍺㏦㜼ᐖ๣࡜ᢠ࣮࢜࢟ࢩࣥ๣ࡀ྾ჾࡢⓎ⏕ࢆῶᑡࡉࡏࡿࡇ࡜ࡀሗ࿌ࡉࢀ㸪ࡲࡓ㸪IAA2 ࣮࢜࢟ࢩࣥᛂ⟅ᛶࣉ࣮ࣟࣔࢱ࣮ࡢάᛶࡀ DMBQ ࡟ᛂ⟅ࡋ࡚ୖ᪼ࡍࡿࡇ࡜ࡀ♧ࡉࢀ࡚࠸ࡿ㸦Tomilov et al., 2005㸧ࠋ⚾ࡓࡕࡣࢥࢩ࣐࡛࢜࢞ࡢࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒ࠿ࡽ㸪DMBQ ᛂ⟅㑇ఏᏊ࡜ࡋ࡚
PjYUC3 ࢆྠᐃࡋࡓ㸦Ishida et al., submitted㸧ࠋYUCCA ࣇ࢓࣑࣮ࣜ㑇ఏᏊࡣࣇࣛࣅࣥࣔࣀ࢜࢟
ࢩࢤࢼ࣮ࢮࢆࢥ࣮ࢻࡋ࡚࠸ࡿ㑇ఏᏊ࡛࠶ࡾ㸪࣮࢜࢟ࢩࣥ⏕ྜᡂࡢ୺せ⤒㊰࡛ാࡃ㘽㑇ఏᏊ࡛࠶ࡿ㸦Zhao et al., 2001, Mashiguchi et al., 2011㸧ࠋࢩࣟ࢖ࢾࢼࢬࢼ࡛ࡣ 11 㑇ఏᏊࡀ YUCCA ࣇ࢓࣑࣮ࣜ࡟ᒓࡋ࡚࠾ࡾ㸪3 㔜ኚ␗యࡸ 4 㔜ኚ␗య࡛ࡣⴭࡋ࠸Ⓨ⏕ࡢ㜼ᐖࡀᘬࡁ㉳ࡇࡉࢀࡿ
㸦Cheng et al., 2006㸧ࠋࡇࢀࡽࡢࡇ࡜࠿ࡽ㸪YUCCA ࣇ࢓࣑࣮ࣜ㑇ఏᏊ࡟ࡼࡿ࣮࢜࢟ࢩࣥࡢ᫬✵㛫ⓗ࡞⏕ྜᡂࡢไᚚࡣ㸪Ⓨ⏕㐣⛬࡛㔜せ࡞ᙺ๭ࢆᢸࡗ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿࠋPjYUC3ࢆࢥࢩ࣐࡛࢜࢞㐣๫Ⓨ⌧ࡉࡏࡿ࡜㸪ഃ᰿ࡢᙧᡂࡀಁ㐍ࡉࢀ᰿ࡀ▷ࡃ࡞ࡿ࡜࠸࠺᪸㐍ࡉࢀࡓ࣮࢜࢟ࢩࣥᛂ⟅ࡢ⾲⌧ᆺࡀࡳࡽࢀࡿࡇ࡜࡜㸪ᐇ㝿࡟ IAA ࡢෆ⏕㔞ࡀቑຍࡋ࡚࠸ࡿࡇ࡜࠿ࡽ㸪PjYUC3 ࡣࢩࣟ࢖ࢾࢼࢬࢼ YUCCA ࣇ࢓࣑࣮ࣜ࡜ྠᵝ࡟㸪࣮࢜࢟ࢩࣥ⏕ྜᡂ࡟㛵୚ࡋ࡚࠸ࡿࡇ࡜ࡀ♧၀ࡉࢀࡓࠋPjYUC3 ࣉ࣮ࣟࣔࢱ࣮ࡣ HIF ฎ⌮ࡸᐟ୺᳜≀ࡢ᰿࡟ࡼࡗ࡚྾ჾẟࢆྵࡴ⾲⓶⣽⬊࡛Ⓨ⌧ࡀㄏᑟࡉࢀ㸪྾ჾࡢⓎ⏕ࡀ㐍ࡴ࡜྾ჾࡢ㡬➃㒊ศ࡛᭱ࡶᙉ࠸Ⓨ⌧ࢆ♧ࡋࡓࠋ࣮࢜࢟ࢩࣥᛂ⟅ࡢ࣐࣮࣮࡛࢝࠶ࡿ DR5 ࣉ࣮ࣟࣔࢱ࣮ࡢⓎ⌧ࣃࢱ࣮ࣥࢆゎᯒࡍࡿ࡜㸪PjYUC3࡜ྠᵝ࡟྾ჾࡢ㡬➃㒊ศ࡛ᙉ࠸Ⓨ⌧ࡀࡳࡽࢀࡓࠋ྾ჾ㡬➃࡛సࡽࢀࡓ᪂ࡓ࡞࣮࢜࢟ࢩࣥᛂ⟅ࡢᴟ኱Ⅼࡣ㸪྾ჾ㡬➃ࡀᐟ୺᳜≀ࡢ⤌⧊࡟౵ධࡋ࡚࠸ࡃẁ㝵ࡲ࡛⥔ᣢࡉࢀ࡚࠸ࡿᵝᏊࡀほᐹࡉࢀ࡚࠸ࡿࠋࡲࡓ㸪PjYUC3 ࢆ RNAi ἲ࡛ࣀࢵࢡࢲ࢘ࣥࡍࡿ࡜ᐤ⏕ຠ⋡ࡀῶᑡࡋ㸪PjYUC3 ࢆ⾲⓶≉␗ⓗ࡟Ⓨ⌧ㄏᑟࡍࡿ࡜⾲⓶࡛ࡢ⣽⬊ศ⿣࡜྾ჾẟᵝࡢ⣽⬊ࡢㄏᑟࡀ࡛ࡁ㸪྾ჾⓎ⏕ࡢึᮇ࡟㉳ࡇࡿ࢖࣋ࣥࢺࢆ෌⌧ࡍࡿࡇ࡜ࡀ࡛ࡁࡓࡇ࡜࠿ࡽ㸪PjYUC3 ࡣ࣮࢜࢟ࢩࣥ⏕ྜᡂ࡟㛵ࢃࡿ㑇ఏᏊ࡛㸪྾ჾⓎ⏕ึᮇ≉␗ⓗ࡟ᶵ⬟ࡍࡿ࡜࠸࠺ࡇ࡜ࡀ࠸࠼ࡿࠋࡇࢀࡲ࡛ࡢ࡜ࡇࢁ㸪4ࡘࡢࢥࢩ࣐࢜࢞ YUCCA ࣇ࢓࣑࣮ࣜࡢ㑇ఏᏊࢆྠᐃࡋ࡚࠾ࡾ㸪ࡑࡢ࠺ࡕ PjYUC2 ࡜ PjYUC4ࡣࢩࣟ࢖ࢾࢼࢬࢼࡢ YUCCA ࣇ࢓࣑࣮ࣜࡢ࠺ࡕ᰿࡛Ⓨ⌧ࡀࡳࡽࢀࡿ AtYUC3,5,7,8,9 ࡜㑇ఏᏊ⣔⤫ᶞୖ࡛ྠࡌࢡ࣮ࣞࢻ࡟ᒓࡍࡿࠋPjYUC3 ࡣࡇࡢࢡ࣮ࣞࢻࡢࡕࡻ࠺࡝እഃ࡟఩⨨ࡋ࡚࠸ࡿࠋ
S. asiatica㸪O. aegyptiaca㸪
T. versicolor ࡢ EST 㓄ิ࠿ࡽྠᐃࡋࡓࡑࢀࡒࢀࡢ✀࡟࠾ࡅࡿ PjYUC3ࡢ࣍ࣔࣟࢢࡣ㸪ᆅୖ㒊ࡸ྾ჾㄏᑟ๓ࡢ᰿࡛ࡣⓎ⌧ࡀ࡞ࡃ㸪྾ჾㄏᑟᚋ࡟ࡢࡳⓎ⌧ࡀࡳࡽࢀࡿ
ࡇ࡜࠿ࡽ㸪௚ࡢࣁ࣐࢘ࢶ࣎⛉ᐤ⏕᳜≀࡟࠾࠸࡚ࡶ྾ჾⓎ⏕᫬࡟ྠᵝࡢᶵ⬟ࢆ᭷ࡍࡿࡇ࡜ࡀ♧၀ࡉࢀࡓࠋࢩࣟ࢖ࢾࢼࢬࢼࡢ◊✲࠿ࡽࡣ㸪࣮࢜࢟ࢩࣥᛂ⟅ࡢᴟ኱Ⅼࡢᙧᡂ࡟ࡣ㸪PIN ࣇ࢓࣑
࣮ࣜࡸ AUX/LAX ࣇ࢓࣑࣮ࣜ࡞࡝ࡢ࣮࢜࢟ࢩࣥ㍺㏦య࡟ࡼࡿ⣽⬊㛫ᴟᛶ㍺㏦ࡀ㔜せ࡞ᙺ๭ࢆᣢࡘࡇ࡜ࡀ♧ࡉࢀ࡚࠸ࡿࠋࡑࡢࡓࡵ㸪྾ჾ㡬➃࡛ࡢ࣮࢜࢟ࢩࣥᛂ⟅ࡢᴟ኱Ⅼࡢᙧᡂ࡜⥔ᣢ࡟ࡶ㸪௒ᅇྠᐃࡉࢀࡓ PjYUC3 ࡟ࡼࡿ⏕ྜᡂࡔࡅ࡛࡞ࡃ㸪࣮࢜࢟ࢩࣥ㍺㏦య࡟ࡼࡿᴟᛶ㍺㏦ࡀ

BSJ-Review 7:246 (2016)

植物科学最前線 7:247 (2016)㛵୚ࡋ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿࠋYUCCA ࣇ࢓࣑࣮ࣜ㑇ఏᏊࡸ࣮࢜࢟ࢩࣥ㍺㏦య࡞࡝ࡢ࢜
࣮࢟ࢩࣥ㛵㐃㑇ఏᏊࡢ㐍໬ࡀ㸪྾ჾᙧᡂ⬟ࡢ⋓ᚓ࡟࡝ࢀࡔࡅᐤ୚ࡋ࡚࠸ࡿࡢ࠿⯆࿡῝࠸Ⅼ࡛࠶ࡿࠋ㸳㸬࠾ࢃࡾ࡟ḟୡ௦ࢩ࣮ࢣࣥࢧ࣮ࡢⓏሙ࡟ࡼࡗ࡚㸪ศᏊ㑇ఏᏛࡢᑐ㇟࡜࡞ࡿ⏕≀✀ࡢᖜࡣ㣕㌍ⓗ࡟ᗈࡀࡗࡓࠋ⚾ࡓࡕࡢ◊✲ࢢ࣮ࣝࣉ࡛ࡣ㸪ࣁ࣐࢘ࢶ࣎⛉ࡢ᰿ᐤ⏕᳜≀࡛࠶ࡿࢫࢺࣛ࢖࢞࡜ࢥࢩ࣐࢜࢞࡟ࡘ࠸࡚㸪ࢤࣀ࣒᝟ሗ㸪ࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒᝟ሗ㸪ᙧ㉁㌿᥮ἲࡢ☜❧࡞࡝࡟ࡼࡗ࡚◊✲⎔ቃࢆᩚഛࡋ࡚ࡁࡓࠋࡇࢀࡽࡢࣜࢯ࣮ࢫࢆ⏝࠸ࡓ◊✲࠿ࡽ㸪྾ჾ࡜࠸࠺ᐤ⏕᳜≀࡟ࣘࢽ࣮ࢡ࡞ჾᐁࡢⓎ⏕ࡢ୍➃ࡀ㸪࣮࢜࢟ࢩࣥ࡜࠸࠺୍⯡ⓗ࡞᳜≀࣍ࣝࣔࣥࡢࣘࢽ࣮ࢡ࡞ไᚚ࡟ࡼࡗ࡚㸪㐙⾜ࡉࢀ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡾࡘࡘ࠶ࡿࠋ᰿⣔ࡢᵓ⠏࡜࠸࠺ほⅬ࠿ࡽࡳࡿ࡜㸪྾ჾࡢᙧᡂࡣ⎔ቃ࡟ᛂ⟅ࡋࡓჾᐁⓎ⏕ࡢ୍౛࡜ࡳࡿࡇ࡜ࡀ࡛ࡁࡿ࡛࠶ࢁ࠺ࠋᐤ⏕᳜≀ࡣᐟ୺᳜≀ࡀ㏆ࡃ࡟࠸ࡿ࡜࠸࠺⎔ቃࢆㄆ㆑ࡋ㸪ࡑࢀ࡟ᑐࡍࡿᛂ⟅࡜ࡋ࡚྾ჾࢆᙧᡂࡍࡿࡢ࡛࠶ࡿࠋ௒ࡢ࡜ࡇࢁ㸪HIF ࢆ⏝࠸ࡓᐟ୺᳜≀ㄆ㆑ࢩࢫࢸ࣒࡜㸪࣮࢜࢟ࢩࣥࢆ௓ࡋࡓჾᐁⓎ⏕ࡢࢩࢫࢸ࣒ࡀ࡝ࡢࡼ࠺࡟ࡘ࡞ࡀࡗ࡚࠸ࡿࡢ࠿୙࡛᫂࠶ࡿࠋ௒ᚋࡢ◊✲ࡢ኱ࡁ࡞ㄢ㢟ࡢ୍ࡘ࡛࠶ࢁ࠺ࠋከᵝ࡞᳜≀✀ࢆ◊✲ࡍࡿࡇ࡜࡛㸪ᚑ᮶⾜ࢃࢀ࡚ࡁࡓࣔࢹ᳜ࣝ≀ࡢ◊✲࠿ࡽࡔࡅ࡛ࡣࢃ࠿ࡽ࡞࠸㸪᳜≀ࡀᣢࡘᬑ㐢ᛶࡸ㸪ࡑࢀࡒࢀࡢ✀ࡢᣢࡘ≉Ṧᛶࡀ᫂☜࡟࡞ࡗ࡚࠸ࡃࡢ࡛ࡣ࡞࠸ࡔࢁ࠺࠿ࠋ௒ᚋᐤ⏕࣓࢝ࢽࢬ࣒ࡢヲ⣽ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡅࡤ㸪ᐤ⏕㞧ⲡࡢ㜵㝖ἲࡢ☜❧ࡀྍ⬟࡟࡞ࡿ࠿ࡶࡋࢀ࡞࠸ࠋ㸴㸬ㅰ㎡ᮏ✏࡛⤂௓ࡋࡓⴭ⪅ࡽࡢ◊✲ࡣ㸪JSPS DC1 ࡢᨭ᥼ࢆᚓ࡚㐙⾜ࡋࡓࠋ㸵㸬ᘬ⏝ᩥ⊩
Albrecht, H., Yoder, J.I., & Phillips, D.A. 1999. Flavonoids promote haustoria formation in the rootparasite Tryphysaria. Plant Physiol. 119: 585-591.
Bandaranayake, P.C.G., Filappova, T., Tomilov, A., Tomilova, N.B., Jamison-McClung, D., Ngo, Q.,
Inoue, K., & Yoder, J.I. 2010.A single-electron reducing quinone oxidoreductase is necessary toinduce haustorium development in the root parasitic plant Triphysaria. Plant Cell 22: 1404-1419
Baird, V.WM., & Riopel, J.L. 1983. Experimental studies of the attachment of the parasitic
angiosperm Agalinis purpurea to a host. Protoplasma 118: 206-208
Baird, V.WM., & Riopel, J.L. 1984. Experimental studies of haustorium initiation and early
development in Agalinis purpurea (L.) RAF. (Schrophulariaceae) . Amer. J. Bot. 71㸦6㸧: 803-814Barkman, T.J., McNeal, J.R., Lim, S., Coat, G., Croom, H.B., Young, N.D., & dePamphilis, C.W.
2007. Mitochondrial DNA suggests at least 11 origins of parasitism in angiosperms and reveals
genomic chimerism in parasitic plants. BMC Evol. Biol. 7: 248
Bar-Nun, N., Sachs, T., & Mayer, A.M. 2008. A role for IAA in the infection of Arabidopsis thaliana by
Orobanche aegyptiaca. Ann. Bot. 101: 261–5

BSJ-Review 7:247 (2016)

植物科学最前線 7:248 (2016)

Caldwell, E.S., & Steelink, C. 1969. Phenoxy radical intermediate in the enzymatic degradation oflignin model compounds. Biochim. Biophys. Acta. 184: 420-431.
Chang, M. & Lynn, D.G. 1986. The haustorium and the chemistry of host recognition in parasitic
angiosperms. J. Chem. Ecol. 12㸦2㸧: 561-579.
Cheng, Y., Dai, X., & Zhao, Y. 2006. Auxin biosynthesis by the YUCCA flavin monooxygenases
controls the formation of floral organs and vascular tissues in Arabidopsis. Genes Dev. 20: 1790–9.
Cook, C.E., Whichard, L.P., Turner, B., & Wall, M.E. 1966. Germination of witchweed 㸦Striga lutea
Lour.㸧: isolation and properties of a potent stimulant. Science. 154: 1189-1190.
Cui, S., Wakatake, T., Hashimoto, K., Saucet, S.B., Toyooka, K., Yoshida, S., & Shirasu, K. 2016.
Haustorial hairs are specialized root hairs that support parasitism in the facultative parasitic plant
Phtheirospermum japonicum. Plant Phys. 170: 1492-1503
Dörr, I. 1997 How Striga parasitizes its host: a TEM and SEM study. Annals of Botany 79:463–472.
Heide-Jorgensen, H.S., & Kuijt, J. 1993 Epidermal derivatives as xylem elements and transfer cells: astudy of the host-parasite interface in 2 species of Triphysaria (Scrophulariaceae). Protoplasma
174: 173-183
Ishida, J.K., Yoshida, S., Ito, M., Namba, S., & Shirasu, K. 2011 Agrobacterium rhizogenes-mediatedtransformation of the parasitic plant Phtheirospermum japonicum. PLoS ONE 6: e25802
Keyes, W.J., Palmer, A.G., Erbil, W.K., Taylor, J.V., Apkarian, R.P., Weeks, E.R., & Lynn, D.G.
2007. Sernagenesis and the parasitic angiosperm Striga asiatica. Plant J. 51: 707-716.
Kim, D., Kocz, R., Boone, L., Keyes, W.J., & Lynn, D.G. 1998.On becoming a parasite: Evaluating
the role of wall oxidase in parasitic plant development. Chem. Biol. 5: 103-117.
Mashiguchi, K., Tanaka, K., Sakai, T., Sugawara, S., Kawaide, H., Natsume, M., Hanada, A., Yaeno,T.,
Shirasu, K., Yao, H., McSteen, P., Zhao, Y., Hayashi, K., Kamiya, Y., & Kasahara, H. 2011 The
main auxin biosynthesis pathway in Arabidopsis. Proc. Natl. Acad. Sci. U.S.A. 108:18512-7
Sablon ML du. 1887 Recherches sur les organes d'absorption des ptantes parasites (Rhinanthées etSantalacées). Ann. Sci. Nat. Ser. 7 [Bot] 6:90-117 + 3 plates
Smith, C.E., Ruttledge, T., Zeng, Z., O’Malley, R.C., & Lynn, D.G. 1996. A mechanism for inducing
plant development – the genesis of a specific inhibitor. Proc. Natl. Acad. Sci. U.S.A. 92:
11786-11790.
Spallek, T., Mutuku, J.M., & Shirasu, K. 2003. The genus Striga: a witch profile. Mol. Plant Pathol.
14: 861-869.
Tomilov, A.A., Tomilova, N.B., Abdallah, I., & Yoder, J.I. 2005. Localized Hormone Fluxes and EarlyHaustorium Development in the Hemiparasitic Plant. Plant phys.138: 1469–1480.
Yang,Z., Wafula, E.K., Honaas, L.A., Zhang, H., Das, M., Fernandez-Aparicio, M., Huang, K.,
Bandaranayake, P.C.G., Wu, Biao., Der, J.P., Clarke, C.R., Ralph, P.E., Landherr, L., Altman, N.S.,
Timko, M.P., Yoder, J.I., Westwood, J.H., & dePamphilis, C.W. 2014. Comparative transcriptome
analyses reveal core parasitism genes and suggest gene duplication and repurposing as sources ofstructural novelty. Mol. Biol. Evol. 32(3): 767–790BSJ-Review 7:248 (2016)

植物科学最前線 7:249 (2016)

Zhao, Y., Christensen, S.K., Fankhauser, C., Cashman, J.R., Cohen, J.D., Weigel, D., & Chory, J. 2001.
A role for flavin monooxygenase-like enzymes in auxin biosynthesis. Science 291: 306–9

BSJ-Review 7:249 (2016)