植物科学最前線 5:130 （2014）⳦ᚑᒓᰤ㣴᳜≀ࡢ㐍໬࡟క࠺⳦᰿⳦┦ࡢࢩࣇࢺ̾㐍໬ࡢ㐨ࡢࡾ࡛࠾ࡇࡗࡓ⳦᰿ඹ⏕ࡢࢲ࢖ࢼ࣑ࢵࢡ࡞ኚ໬̾䢢㎷⏣᭷⣖ ࣭ᶓᒣ₶ ࣭୕ྜྷ୍ග ࣭㐟ᕝ▱ஂ ᮾ໭኱Ꮫ᳜≀ᅬࠛ௝ྎᕷ㟷ⴥ༊ᕝෆ 㸦⌧ᡤᒓబ㈡኱Ꮫ㎰Ꮫ㒊ᛂ⏝⏕≀⛉Ꮫ⛉ࠛబ㈡ᕷᮏᗉ⏫㸯␒ᆅ㸧ᒣᙧ኱Ꮫ⌮Ꮫ㒊⏕≀Ꮫ⛉ࠛᒣᙧᕷᑠⓑᕝ⏫ ༓ⴥ኱Ꮫ኱Ꮫ㝔ᅬⱁᏛ◊✲⛉ࠛ༓ⴥ┴ᯇᡞᕷᯇᡞ ᅜ❧⛉Ꮫ༤≀㤋⟃Ἴᐇ㦂᳜≀ᅬࠛࡘࡃࡤᕷኳஂಖ 

Yuki Ogura-Tsujita, Jun Yokoyama, Kazumitsu Miyoshi, Tomohisa Yukawa
Shifts in mycorrhizal fungi during the evolution of autotrophy to mycoheterotrophy
Key words: autotrophy, Cymbidium, mixotrophy, mycoheterotrophy, mycorrhizal fungiBotanical Gardens, Tohoku University

12-2 Kawauchi, Aobaku, Sendai, Miyagi 980-0862, Japan
Current address: Department of Applied Biological Sciences, Faculty of Agriculture, SagaUniversity, 1 Honjo-machi, Saga 840-8502, Japan

Department of Biology, Faculty of Science, Yamagata University
1-4-12 Kojirakawa-machi, Yamagata 990-8560, Japan

Graduate School of Horticulture Laboratory of Ornamental Plant Science, Chiba UniversityMatsudo, 271-8510, Japan
Tsukuba Botanical Garden, National Museum of Nature and Science4-1-1, Amakubo, Tsukuba, Ibaraki 305-0005, Japan㸯ࡣࡌࡵ࡟ ගྜᡂࢆࡸࡵ࡚ࡋࡲࡗࡓ᳜≀࡛࠶ࡿ⳦ᚑᒓᰤ㣴᳜≀ࡣ୍య࡝ࡢࡼ࠺࡟㐍໬ࡋ࡚ࡁࡓࡢ࠿㸽ࡑࡢ࢝ࢠࡣ⳦᰿⳦࡜ࡢඹ⏕㛵ಀࡀ≬ࡗ࡚ࡋࡲࡗࡓࡇ࡜࡟࠶ࡿࠋ㏻ᖖ㸪᳜≀ࡣගྜᡂ࡟ࡼࡗ࡚ࡏࡗࡏ࡜⮬ศ࡛஧㓟໬Ⅳ⣲࡜Ỉ࠿ࡽ᭷ᶵⅣ⣲࡛࠶ࡿ⢾ࢆྜᡂࡋ࡞ࡀࡽᬽࡽࡋ࡚࠸ࡿࠋ୍᪉㸪⳦ᚑᒓᰤ㣴᳜≀ࡣගྜᡂ࡛ࡣ࡞ࡃ㸪ඹ⏕ࡍࡿ⳦᰿⳦࠿ࡽ᭷ᶵⅣ⣲ࢆࡶࡽࡗ࡚ᬽࡽࡋ࡚࠸ࡿࠋከࡃࡢගྜᡂ᳜≀࡛ࡣ㸪ගྜᡂ࡛ᚓࡓⅣ⣲ࢆ⳦᰿⳦࡟ཷࡅΏࡍ࠿ࢃࡾ࡟㸪⳦࠿ࡽࣜࣥࡸ❅⣲࡞࡝ࡢᰤ㣴ศࢆࡶࡽ࠺࡜࠸࠺ᣢࡕࡘᣢࡓࢀࡘࡢ㛵ಀࢆಖࡗ࡚࠸ࡿࡀ㸪⳦ᚑᒓᰤ㣴᳜≀࡛ࡣ㸪Ⅳ⣲ࡲ࡛ࡶ⳦࠿ࡽᦢྲྀࡋ࡚ࡋࡲ࠺ࡢࡔࠋാ࠿ࡎࡋ࡚Ⅳ⣲ࢆࡶࡽ࠼ࡿࡢࡔ࠿ࡽ㸪ᙼࡽࡢ⳦᰿ඹ⏕ࡣ᳜≀࡟࡜ࡗ࡚ࡲࡉ࡟ࠕ✲ᴟࠖ࡜࠸࠼ࡿࠋᮏ⥲ㄝ࡛ࡣ㸪⳦ᚑᒓᰤ㣴᳜≀࡟ࡳࡽࢀࡿࠕ✲ᴟࠖࡢ⳦᰿ඹ⏕ࣃࢱ࣮ࣥࢆ⤂௓ࡋ㸪ࣛࣥ⛉᳜≀ࢆࣔࢹࣝ࡟⳦᰿⳦ࡢඹ⏕㛵ಀࡀ࡝ࡢࡼ࠺࡟ኚ໬ࡋ࡚࠸ࡗࡓࡢ࠿㸪ࡑࡢㅦ࡟㏕ࡗࡓ⚾㐩ࡢ◊✲౛ࢆ⤂௓ࡋ࡚㸪᭱ᚋ࡟ᮏ◊✲ศ㔝࡟࠾ࡅࡿ௒ᚋࡢᒎᮃࢆ⪃࠼ࡿࠋ

Y. Ogura-Tsujita - 1

植物科学最前線 5:131 （2014）㸰⳦ᚑᒓᰤ㣴᳜≀ࡢ⳦᰿ඹ⏕ࣃࢱ࣮ࣥ
 1990 ᖺ௦࠿ࡽ㸪DNA ࡢ᝟ሗࢆ฼⏝ࡋ࡚⳦ࡢ✀㢮ࢆ≉ᐃࡍࡿ࠸ࢃࡺࡿศᏊྠᐃἲࡀᬑཬࡋ㸪ୡ⏺୰ࡢ◊✲⪅ࡀ➇ࡗ࡚ࡇࡢ≀⌋ࡋ࠸᳜≀ࡢ⳦᰿⳦ࢆㄪ࡭ୖࡆ࡚࠸ࡗࡓ㸦Cullings et al. 1996,Tayor & Bruns 1997, Selosse et al. 2002 ࡯࠿㸧ࠋ᝟ሗࡢ㞟✚࡟࡜ࡶ࡞ࡗ࡚ぢ࠼࡚ࡁࡓࡢࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ㏆⦕࡞ගྜᡂ᳜≀࡜ࡣ඲ࡃ␗࡞ࡿ≉Ṧ࡞⳦᰿⳦࡜ࡢඹ⏕㛵ಀࢆ⠏ࡁୖࡆ࡚࠸ࡓࡢ࡛࠶ࡿࠋࡉࡽ࡟㸪⊂❧ᰤ㣴࠿ࡽ⳦ᚑᒓᰤ㣴࡬ࡢ㐍໬ࡣ㸪ࢥࢣ᳜≀࠿ࡽ⿕Ꮚ᳜≀ࡲ࡛㝣ୖ᳜≀ࡢᵝࠎ࡞⣔⤫࡛⊂❧࡟ 40 ᅇ௨ୖ࠾ࡇࡗ࡚࠸ࡿࡀ㸦Merckx et al. 2013㸧㸪⳦᰿ඹ⏕ࡢ≉Ṧ໬࡟ࡣ࠸ࡃࡘ࠿ࡢࣃࢱ࣮ࣥࡀ࠶ࡿࡇ࡜ࡀࢃ࠿ࡗ࡚ࡁࡓࠋ ࡲࡎࡣ㸪ࢥࢣ᳜≀ࡢࢱ࢖㢮ࢫࢪࢦࢣ⛉࡛࠾ࡇࡗࡓ⳦᰿ඹ⏕ࡢࢲ࢖ࢼ࣑ࢵࢡ࡞ኚ໬ࢆ⤂௓ࡋࡼ࠺㸦ᅗ㸯㸧ࠋከࡃࡢࢱ࢖㢮ࡣࢢ࣒ࣟࢫ㛛ࡢ⳦㢮࡜࢔࣮ࣂࢫ࣮࢟ࣗࣛ⳦᰿ඹ⏕࡜ࡼࡤࢀࡿඹ⏕⣔ࢆᙧᡂࡋ࡚࠸ࡿࡀ㸪ࡑࡢ୰࠿ࡽ㸪⳦᰿⳦࡜ඹ⏕ࡋ࡞࠸㠀⳦᰿ᛶࡢ⣔⤫ࡀฟ⌧ࡋ㸪ࡉࡽ࡟ࡑࡢ୰࠿ࡽ㸪ᢸᏊ⳦ࡢࢶࣛࢫࢿࣛ࡜࠸࠺௚࡜඲ࡃ␗࡞ࡿ✀㢮ࡢ⳦࡜ඹ⏕ࡍࡿࢫࢪࢦࢣ⛉ࡀฟ⌧ᅗ㸯 ࢥࢣ᳜≀ࢱ࢖㢮࡛ࡢ⳦᰿ඹ⏕ࡢኚ໬㸦Kottke & Nebel 2005 ࢆᨵኚࠋ᳜≀ࡢศ㢮య⣔ࡣDavis (2004)࡟ࡼࡿ㸧ࠋ▮༳ࡣඹ⏕㛵ಀࡀኚ໬ࡋࡓ࣏࢖ࣥࢺࢆ♧ࡍࠋ*ࡣ⌧ᅾ౑ࢃࢀ࡚࠸࡞࠸ศ㢮⩌ྡ࡛㸪⌧ᅾ Lepicoleales ࡣ Lepicoleaceae ࡜ࡋ࡚ࢶ࣑࣎ࢦࢣ┠࡟㸪Radulales ࡣ Radulaceae࡜ࡋ࡚ࢡ࣐ࣛࢦࢣࣔࢻ࢟┠㸦Porellales㸧࡟㸪Monocleales ࡣ Monocleaceae ࡜ࡋ࡚ࢮࢽࢦࢣ┠࡟㸪
Ricciales ࡣ Ricciaceae ࡜ࡋ࡚ࢮࢽࢦࢣ┠࡟ྵࡲࢀ࡚࠸ࡿ㸦ᔱᮧ 2012㸧

Y. Ogura-Tsujita - 2

植物科学最前線 5:132 （2014）

ࡍࡿ㸦Bidartondo et al. 2003, Kottke & Nebel 2005㸧ࠋࡑࡋ࡚ࡑࡢᚋ㸪ᢸᏊ⳦࡜ඹ⏕ࡍࡿ⣔⤫ࡢ
୰࠿ࡽ㸪Aneura mirabilis (Cryptothallus mirabilis)࡜࠸࠺⳦ᚑᒓᰤ㣴᳜≀ࡀㄌ⏕ࡋࡓࠋ᳜≀ࡢ㐍໬ࡢ㐨ࡢࡾ࡛㸪⳦᰿⳦࡜ࡢඹ⏕㛵ಀࡣ኱ࡁࡃኚ໬ࡋ࡚࠸ࡿࠋྠᵝ࡟௚ࡢ⣔⤫࡟࠾࠸࡚ࡶ㸪ࡇࡢࡼ࠺࡞⳦ᚑᒓᰤ㣴໬࡟క࠺ඹ⏕㛵ಀࡢࢲ࢖ࢼ࣑ࢵࢡ࡞ࢩࣇࢺࡀぢࡘ࠿ࡗ࡚࠾ࡾ㸪ࡑࡢࢩࣇࢺࣃࢱ࣮ࣥࡣ཰᮰ࡍࡿ⳦ࡢ✀㢮࡟ࡼࡗ࡚࠾࠾ࡊࡗࡥ࡟㸰Ѹ㸯࠿ࡽ㸰Ѹ㸱ࡢ㸱ࡘ࡟ศࡅࡽࢀࡿࠋࡑࢀࡒࢀࡢඹ⏕ࣃࢱ࣮ࣥࢆࡶࡘ᳜≀ྡࡸ⳦ࡢ✀㢮ࡣ➨㸲❶࡟ヲ㏙ࡉࢀ࡚࠸ࡿࡓࡵ㸪ࡇࡇ࡛ࡣ㸪ඹ⏕㛵ಀࡢࢩࣇࢺࣃࢱ࣮ࣥ࡟ࡘ࠸࡚↔Ⅼࢆࡋࡰࡗ࡚⤂௓ࡍࡿ㸦ࡉࡽ࡟ヲࡋ࠸ෆᐜࡣ
Bidartondo (2005), Smith & Read (2008), Sellosse & Roy 2009, Waterman et al. (2013)ཧ↷㸧ࠋ㸰Ѹ㸯㸬እ⏕⳦᰿⳦࡬ࡢࢩࣇࢺ ୺࡟ࢶࢶࢪ⛉࡜ࣛࣥ⛉ࡢ⳦ᚑᒓᰤ㣴✀࡟ぢࡽࢀ㸪ඹ⏕┦ᡭࡣᮏ᮶ᶞᮌ࡜⳦᰿ඹ⏕ࢆ⾜ࡗ࡚࠸ࡿእ⏕⳦᰿⳦࡛࠶ࡿࠋࡑࡶࡑࡶእ⏕⳦᰿⳦ࡣᶞᮌ࠿ࡽⅣ⣲ࢆࡶࡽࡗ࡚ᬽࡽࡋ࡚࠸ࡿࡢ࡛㸪ࡑࡢእ⏕⳦᰿⳦࠿ࡽⅣ⣲ࢆࡶࡽࡗ࡚ᬽࡽࡋ࡚࠸ࡿ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪⤖ᒁࡣ࿘㎶ࡢᶞᮌࡀගྜᡂ࡛ᚓࡓⅣ⣲ࢆ⳦᰿⳦ࡢ⳦⣒ࢆ௓ࡋ࡚฼⏝ࡋ࡚࠸ࡿࡇ࡜࡟࡞ࡿ㸦ᅗ㸰㸧ࠋࡇࡢࡼ࠺࡟ᶞᮌ࣭⳦᰿⳦࣭⳦ᚑᒓᰤ㣴᳜≀࡜࠸࠺㸱ࡘࡢ⏕ࡁ≀ࡀࡘ࡞ࡀࡗ࡚࠸ࡿ≧ែࡣ㸪㸱⪅ඹ⏕⣔࡜ࡼࡤࢀ࡚࠸ࡿࠋ ࢶࢶࢪ⛉ࡢ୰࡛᭱ࡶྂ࠸⣔⤫࡛࠶ࡿࢻ࢘ࢲࣥࢶࢶࢪࡢ௰㛫ࡣ㸪࢔࣮ࣂࢫ࣮࢟ࣗࣛ⳦᰿ᛶ࡛࠶ࡿ࡜ሗ
࿌ࡉࢀ࡚࠸ࡿࡇ࡜࠿ࡽ㸦Abe 2005, Obase & Matsuda2013㸧㸪ࢶࢶࢪ⛉ࡣࡶ࡜ࡶ࡜࢔࣮ࣂࢫ࣮࢟ࣗࣛ⳦᰿
⳦㸦AM ⳦㸧࡜⳦᰿ඹ⏕ࡋ࡚࠸ࡓྍ⬟ᛶࡀ㧗࠸ࠋ⳦ᅗ㸰 እ⏕⳦᰿⳦࡜ࡢ㸱⪅ඹ⏕⣔ࠋ▮༳ࡣⅣ⣲ࡢ⛣ື᪉ྥࢆ♧ࡍᚑᒓᰤ㣴᳜≀࡛࠶ࡿࢠࣥࣜࣙ࢘ࢯ࢘ࡸࢩࣕࢡࢪࣙ࢘ࢯ࢘ࡢ௰㛫ࡣ㸪ࡇࡢḟ࡟ྂ࠸⣔⤫࡛㸪እ⏕⳦᰿ᛶࡢᢸᏊ⳦㢮࡜ඹ⏕ࡋ࡚࠸ࡿࠋ࠾ࡑࡽࡃࢶࢶࢪ⛉࡛ࡣගྜᡂࢆࡸࡵࡿ㐍໬ࡢ㐣⛬࡛㸪⳦᰿⳦ࡀ AM ⳦࠿ࡽእ⏕⳦᰿⳦࡬ࡍࡾ࠿ࢃࡗ࡚ࡋࡲࡗࡓࡢ࡛ࡣ࡞࠸࠿࡜ண᝿ࡉࢀࡿࠋ ࣛࣥ⛉࡛ࡣ㸪᭱ࡶྂ࠸⣔⤫࡛࠶ࡿࣖࢡࢩ࣐ࣛࣥࡢ௰㛫ࢆྵࡴ࡯࡜ࢇ࡝ࡢ✀ࡀ㸪ࣛࣥᆺ⳦᰿⳦࡜ࡼࡤࢀࡿᢸᏊ⳦ࡢࢢ࣮ࣝࣉ㸦ࢶࣛࢫࢿࣛ⛉㸪ࢣࣛࢺࣂࢩࢹ࢕࣒࢘⛉㸪ࣟ࢘ࢱࢣ⛉ࡢ୍㒊ࢆྵࡴ㸧࡜ඹ⏕ࡋ࡚࠸ࡿࡇ࡜࠿ࡽ㸦Yukawa et al. 2009㸧㸪⳦ᚑᒓᰤ㣴໬࡟కࡗ࡚㸪ඹ⏕ࡍࡿ⳦ࡀࣛࣥᆺ⳦᰿⳦࠿ࡽእ⏕⳦᰿⳦࡬ࢩࣇࢺࡋࡓ࡜⪃࠼ࡽࢀࡿࠋࣛࣥ⛉ࡣ⣙ 25,000 ✀ࢆྵࡴ኱ࡁ࡞⛉࡛࠶ࡿࡀ㸪ࣛࣥ⛉ࡢ୰ࡔࡅ࡛ࡶගྜᡂࢆࡸࡵࡿ㐍໬ࡀ 20 ᅇ࠶ࡲࡾ࠾ࡇࡗ࡚࠸ࡿ࡜࠸ࢃࢀ
࡚࠾ࡾ㸦Molvray et al. 2000㸧㸪እ⏕⳦᰿⳦࡬ࡢ஌ࡾ᥮࠼ࡣࣛࣥ⛉ࡢ୰ࡔࡅ࡛ࡶᗄᗘࡶ⧞ࡾ㏉ࡉࢀ࡚࠸ࡿࠋY. Ogura-Tsujita - 3

植物科学最前線 5:133 （2014）㸰Ѹ㸰㸬࢔࣮ࣂࢫ࣮࢟ࣗࣛ⳦᰿⳦࡜ࡢ㸱⪅ඹ⏕ ࣄࢼࣀࢩࣕࢡࢪࣙ࢘⛉ࡸ࣍ࣥࢦ࢘ࢯ࢘⛉࡞࡝࡟ࡳࡽࢀ㸪୺࡟ࢢ࣒ࣟࢫ㛛ࡢࢢ࣒ࣟࢫ⛉㸦ᪧGlomus group A㸧࡟ᒓࡍࡿ⳦᰿⳦࡜ඹ⏕ࡍࡿࣃࢱ࣮࡛ࣥ࠶ࡿࠋ୍⯡࡟㸪AM ⳦࡜ඹ⏕ࡍࡿ⊂❧ᰤ㣴᳜≀࡛ࡣ᳜≀㸯ಶయ࡟」ᩘࡢ⛉࡟ࡲࡓࡀࡿከᩘࡢ AM ⳦ࡀඹ⏕ࡋ࡚࠸ࡿࡀ㸪⳦ᚑᒓᰤ㣴᳜≀࡛ࡣࢢ࣒ࣟࢫ⛉࡟ᒓࡍࡿ≉ᐃࡢࢢ࣮ࣝࣉࡢ⳦࡜㠀ᖖ࡟≉␗ⓗ࡟ඹ⏕ࡋ࡚࠸ࡿሙྜࡀከ࠸
㸦Bidarotondo et al. 2002, Franke et al. 2006 ࡯࠿, ࡓࡔࡋ㸪᭱㏆࡛ࡣ」ᩘࡢ⛉ࡢ AM ⳦࡜ඹ⏕ࡍࡿ౛እࡶぢࡘ࠿ࡗ࡚࠾ࡾ㸦Merckx et al. 2012㸧㸪௒ᚋぢゎࡀኚࢃࡿྍ⬟ᛶࡀ࠶ࡿ㸧ࠋእ⏕⳦᰿⳦࡜ྠᵝ࡟㸱⪅ඹ⏕⣔ࢆᵓ⠏ࡋ㸪㞄᥋ࡍࡿ⊂❧ᰤ㣴᳜≀࠿ࡽ⳦᰿⳦ࢆ௓ࡋ࡚Ⅳ⣲ࢆࡶࡽࡗ࡚ᬽࡽࡋ࡚࠸ࡿࠋ௚ࡢࣃࢱ࣮࡛ࣥࡣ㸪ගྜᡂࢆࡸࡵࡿ㐣⛬࡛㸪ඹ⏕ࡍࡿ⳦ࡀ඲ࡃ㐪࠺✀㢮࡟ࡍࡾ࠿ࢃࡗ࡚ࡋࡲ࠺ࡇ࡜ࡀከ࠸ࡀ㸪ᮏࣃࢱ࣮࡛ࣥࡣ㸪」ᩘࡢ␗࡞ࡿ⣔⤫ࡢඹ⏕ࣃ࣮ࢺࢼ࣮ࡢ୰࠿ࡽ࠶ࡿ≉ᐃࡢ⣔⤫ࡢ⳦ࡢࡳ࡜ඹ⏕ࡍࡿࡼ࠺࡟㛵ಀࢆ⤠ࡾ㎸ࢇ࡛࠸ࡗࡓ࡜⪃࠼ࡽࢀ࡚࠸ࡿ
㸦Merckx & Bidartondo 2008㸧ࠋ㸰Ѹ㸱㸬⭉⏕⳦࡬ࡢࢩࣇࢺ ⌧ᅾࡢ࡜ࡇࢁࣛࣥ⛉࡛ࡢࡳⓎぢࡉࢀ࡚࠸ࡿࢩࣇࢺࣃࢱ࣮࡛ࣥ㸪ᮌᮦࡸⴠⴥⴠᯞࢆศゎࡍࡿ⭉⏕⳦㢮࡜ඹ⏕ࡋ࡚࠸ࡿࠋඛ࡟⤂௓ࡋࡓ㸰ࡘࡢࣃࢱ࣮࡛ࣥࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ㸱⪅ඹ⏕⣔࡟ࡼࡗ࡚⏕ࡁࡓ᳜≀࠿ࡽⅣ⣲ࢆᦢྲྀࡋ࡚࠸ࡓࡀ㸪ࡇࡢሙྜࡣᮌᮦࡸⴠࡕⴥ࡞࡝᳜≀ࡢṚ㧁ࢆศゎࡍࡿ⳦࠿ࡽⅣ⣲ࢆࡶࡽࡗ࡚ᬽࡽࡋ࡚࠸ࡿ㸦ᅗ㸱㸧ࠋࡘࡲࡾ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ࿘㎶࡟㌿ࡀࡗ࡚࠸ࡿಽᮌࡸⴠࡕⴥ㸪ᯤࢀᯞࡀⅣ⣲※࡟࡞ࡗ࡚࠸ࡿࡢࡔࠋእ⏕⳦᰿⳦ࡸ AM ⳦࡜ࡢඹ⏕ࣃࢱ࣮ࣥࡣ㸪࡝ࡕࡽࡢ⳦ࡶ᳜≀࠿ࡽྲྀࡾฟࡋ࡚⳦ࢆ༢⊂࡛ᇵ㣴ࡍࡿࡇ࡜ࡀ㞴ࡋ࠸ࡓࡵ㸪DNA ᝟ሗࢆ⏝࠸ࡓศᏊྠᐃἲ࡟ࡼࡗ࡚ࡈࡃ᭱㏆᫂ࡽ࠿࡟࡞ࡗ࡚ࡁࡓඹ⏕⣔࡛࠶ࡿࠋ୍᪉㸪⭉⏕⳦ࡣẚ㍑ⓗᐜ᫆࡟ᇵ㣴࡛ࡁࡿࡓࡵ㸦➨㸲❶ཧ↷㸧㸪ᮏඹ⏕ࣃࢱ࣮ࣥࡣ࠿࡞ࡾྂ࠸᫬௦࠿ࡽㄆ㆑ࡉࢀ
࡚࠸ࡓ㸦Kusano 1911, Hamada 1939㸧ࠋ๓㏙ࡋࡓࡼ࠺࡟ࣛࣥ⛉ࡢከࡃࡣࣛࣥᆺ⳦᰿⳦࡜ඹ⏕ࡋ࡚࠸ࡿࡇ࡜࠿ࡽ㸪ගྜᡂࢆࡸࡵࡿ㐍໬࡟కࡗ࡚㸪ඹ⏕ࡍࡿ⳦᰿⳦ࡣࣛࣥᆺ⳦᰿⳦࠿ࡽᮌᮦ⭉ᮙ⳦ࡸⴠⴥศゎ⳦࡬࡜ࢩࣇࢺࡋࡓ࡜⪃࠼ࡽࢀࡿࠋࡲࡓ㸪ࡇࡢࡼ࠺࡞ࢩࣇࢺࡣࣛࣥ⛉ࡢ୰࡛⊂❧࡟」ᩘᅇ㉳ࡇࡗࡓࡇ࡜ࡀࢃ࠿ࡗ࡚࠸ࡿ㸦Molvray et al. 2000㸧ࠋᅗ㸱 ᮌᮦ⭉ᮙ⳦ࡸⴠⴥศゎ⳦࡜ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࡢࣔࢹࣝᅗ
Y. Ogura-Tsujita - 4

植物科学最前線 5:134 （2014）㸱㸬⳦ᚑᒓᰤ㣴໬࡬ࡢࢫࢸࢵࣉ ගྜᡂࢆࡸࡵࡿ㐍໬ࡢ㐨ࡢࡾࡣ㸪㸯ࢫࢸࢵࣉࡢ⌧㇟࡛ࡣ࡞ࡃẁ㝵ⓗ࡛㸪࠸ࡃࡘ࠿ࡢ୰㛫ⓗ࡞ࢫࢸ࣮ࢪࡀ࠶ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿࠋMerckx etal. 㸦2013㸧࡛ࡣ㸪ḟࡢ୰㛫ⓗ࡞㸰ࢫࢸࢵࣉࢆ࠶ࡆ࡚࠸ࡿ㸦ᅗ㸲㸧ࠋࡲࡎࡣࡌࡵ࡟㸪✀Ꮚࡸ⬊ᏊࡢⓎⱆ᫬ࡸᐇ⏕࣭㓄അయୡ௦࡜࠸ࡗࡓ⏕άྐࡢึᮇ࡟ࡢࡳ⳦ᚑᒓᰤ㣴  ᅗ㸲 ⳦ᚑᒓᰤ㣴࡬ࡢࢫࢸࢵࣉ㸦Merckx et al. 2013 ࢆᨵኚ㸧࡜࠸࠺ࢫࢱ࢖ࣝࢆ࡜ࡾ㸪ᡂ⇍ᚋࡣ⊂❧ᰤ㣴࡛࠶ࡿࠕ⏕άྐึᮇᆺ⳦ᚑᒓᰤ㣴ࠖ㸦initial mycoheterotrophy㸧࡬⛣⾜ࡍࡿࠋḟ࡟㸪ᡂ⇍ࡋࡓಶయࡶ⳦
᰿⳦࡬ᰤ㣴ࢆ౫Ꮡࡍࡿࡼ࠺࡟࡞ࡿࡀ㸪ࡲࡔගྜᡂࡶ⾜ࡗ࡚࠸ࡿࠕ㒊ศⓗ⳦ᚑᒓᰤ㣴ࠖ㸦partial
myco- heterotrophy㸧ࡢࢫࢸࢵࣉ࡬⛣ࡾ㸪ࡘ࠸࡟ࡣගྜᡂࢆࡸࡵ࡚ࡋࡲ࠺᏶඲⳦ᚑᒓᰤ㣴㸦full
mycoheterotrphy㸧࡜࡞ࡿࠋࡇࢀࡽࡣࣛࣥ⛉ࢆࣔࢹࣝ࡟⪃࠼ࡽࢀࡓࣃࢱ࣮࡛ࣥ࠶ࡿࡀ㸪࠾ࡑࡽࡃ௚ࡢ⣔⤫࡛ࡶྠᵝࡢࢫࢸࢵࣉࢆ࡬࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿࠋ࡛ࡣ㸪๓㏙ࡋࡓࡼ࠺࡞⳦᰿ඹ⏕ࡢ≉Ṧ໬ࡣ㸪࡝ࡢࢫࢸࢵࣉ࡛࠾ࡇࡗ࡚࠸ࡿࡢࡔࢁ࠺࠿ࠋከࡃࡢሙྜ㸪⏕άྐึᮇᆺ⳦ᚑᒓᰤ㣴ࡸ㒊ศⓗ⳦ᚑᒓᰤ㣴࡜࠸ࡗࡓ୰㛫ⓗ࡞ࢫࢸࢵࣉ࡟✺ධࡋࡓẁ㝵࡛ࡍ࡛࡟ࡑࡢ≉Ṧ໬ࡀ࠾ࡇࡗ࡚࠸ࡿࠋࡘࡲࡾ㸪ࡍ࡭࡚ࡢᘬࡁ㔠ࡣ⳦᰿ඹ⏕ࡢኚ໬࡟࠶ࡿࡢࡔࠋ㸲㸬⳦᰿ඹ⏕ࡢኚ໬㐣⛬ࢆ✀ศ໬ࡢ࡛ࣞ࣋ࣝ ගྜᡂࢆࡸࡵࡿ㐍໬ࡢ㐨ࡢࡾ࡛⳦᰿⳦࡜ࡢඹ⏕㛵ಀࡀ࡝ࡢࡼ࠺࡟ኚ໬ࡋࡓࡢ࠿ࢆ᥎ᐃࡍࡿࡓࡵ࡟ࡣ㸪⳦ᚑᒓᰤ㣴᳜≀࡜ࡑࢀࡽ࡟㏆⦕࡞⊂❧ᰤ㣴᳜≀ࡢ⳦᰿⳦ࢆẚ㍑ࡍࢀࡤࡼ࠸ࠋࡇࡢ
ࡼ࠺࡞ẚ㍑◊✲ࡣ㸪ࢶࢶࢪ⛉ࢆ୰ᚰ࡟ᩘከࡃᏑᅾࡋࡓ㸦Bidartondo & Bruns 2002 ࡯࠿㸧ࠋࡋ࠿ࡋ㸪ከࡃࡢ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪㏆⦕࡞⊂❧ᰤ㣴᳜≀࡜࠸࠼࡝ࡶ㠀ᖖ࡟⦕ࡀ㐲ࡃ㸦౛࠼ࡤ㸪⚾㐩ࡢ㌟㏆࡟Ꮡᅾࡍࡿࢠࣥࣜࣙ࢘ࢯ࢘࡟᭱ࡶ㏆⦕࡞⊂❧ᰤ㣴᳜≀ࡣࢻ࢘ࢲࣥࢶࢶࢪࡢ௰㛫࡛࠶ࡿ㸧㸪୧⪅ࢆẚ㍑ࡋ࡚ࡶṇ☜࡞ኚ໬㐣⛬ࢆ㏣࠺ࡇ࡜ࡣ㞴ࡋ࠸ࠋ ⚾㐩ࡣ㸪ࡇࡇ᪥ᮏ࡟ወ㊧ⓗ࡟ࡈࡃ㏆⦕࡞ࢢ࣮ࣝࣉ㛫࡛⊂❧ᰤ㣴✀࡜⳦ᚑᒓᰤ㣴✀ࡀศ໬ࡋ࡚࠸ࡿ᳜≀⩌ࢆࣔࢹࣝ࡜ࡋ࡚Ⓨぢࡋࡓࠋࡑࢀࡣ㸪ࣛࣥ⛉ࢩࣗࣥࣛࣥᒓ㸦Cymbidium㸧᳜≀࡛࠶ࡿࠋࡇࢀࡽࡣ㸪ⰼᒇ࡛ぢ࠿ࡅࡿࢩࣥࣅࢪ࣒࢘ࡢ௰㛫࡛㸪⊂❧ᰤ㣴✀㸪㒊ศⓗ⳦ᚑᒓᰤ㣴✀㸪⳦ᚑᒓᰤ㣴✀ࡢࡍ࡭࡚ࢆྠ୍ᒓෆ࡟ྵࢇ࡛࠾ࡾ㸪㏆⦕✀㛫࡛ᰤ㣴ᛶࡀ⇿Ⓨⓗ࡟ከᵝ໬ࡋࡓ᳜≀⩌࡛࠶ࡿࠋᖾ㐠࡞ࡇ࡜࡟㸪ࡇࡢ㸱ࡘࡢᰤ㣴✀ࡣࡍ࡭࡚᪥ᮏ࡟⮬⏕ࡋ࡚࠸ࡿࠋࡑࡇ࡛㸪⚾㐩ࡣࡇࡢࢩࣗࣥࣛࣥᒓࢆࣔࢹࣝ࡟㸪᳜≀ࡢ⣔⤫ゎᯒ࡜⳦᰿⳦ࡢㄪᰝࢆ⾜ࡗࡓࠋ
Y. Ogura-Tsujita - 5植物科学最前線 5:135 （2014）㸲Ѹ㸯㸬᳜≀ࡢ⣔⤫ゎᯒࡀ≀ㄒࡿ⳦ᚑᒓᰤ㣴໬ࡢ㐨ࡢࡾ ᳜≀ࡢศᏊ⣔⤫ゎᯒ࡟ࡼࡾ㸪ᅗ㸳࡟♧ࡍࡼ࠺࡟ࢩࣗࣥࣛࣥᒓࡣࡶ࡜ࡶ࡜ᶞୖ࡛ᬽࡽࡍ╔⏕ࡢ⊂❧ᰤ㣴✀࡛࠶ࡾ㸪ࡑࡢ୰࠿ࡽᆅ㠃࡟㝆ࡾ࡚ᬽࡽࡍࡼ࠺࡟࡞ࡗࡓᆅ⏕ࡢ㒊ศⓗ⳦ᚑᒓᰤ㣴✀ࡀฟ⌧ࡋ㸪ࡉࡽ࡟ࡑࡢ୰࠿ࡽ⳦ᚑᒓ  ᅗ㸳 ㄪᰝࡋࡓࢩࣗࣥࣛࣥᒓ㸳✀࡜ࡑࢀࡽࡢ㢮⦕㛵ಀᰤ㣴࡛࠶ࡿ࣐ࣖࣛࣥ࡜ࢧ࣑࢞ࣛࣥࡢ㸰✀ࡀฟ⌧ࡋࡓࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ㸦Yukawa et al. 2002, Motomura et al.
2010; ᅗ㸳㸧ࠋࡘࡲࡾ㸪ࢩࣗࣥࣛࣥᒓ࡛ࡣ㸪⊂❧ᰤ㣴࠿ࡽ㒊ศⓗ⳦ᚑᒓᰤ㣴࡜࠸࠺୰㛫ẁ㝵ࢆ࡬࡚⳦ᚑᒓᰤ㣴࡬࡜㐍໬ࡋ࡚࠸ࡓࡢ࡛࠶ࡿࠋࡲࡓࡇࡢ◊✲࠿ࡽ㸪࣐ࣖࣛࣥ࡜ࢧ࣑࢞ࣛࣥࡣ࠾஫࠸࡟ጜጒ㛵ಀ࡟࠶ࡾ㸪ࡇࢀࡽ㸰✀࡟᭱ࡶ㏆⦕࡞ගྜᡂࢆࡍࡿ᳜≀ࡣࢼࢠ࡛ࣛࣥ࠶ࡿࡇ࡜ࡶ᫂ࡽ࠿࡟࡞ࡗࡓࠋ ࡑࡇ࡛⚾㐩ࡣ㸪ᅗ㸳࡟࠶ࡿ㸳✀ࡢ⳦᰿⳦ࢆẚ㍑ࡋ㸪᳜≀ࡀ⊂❧ᰤ㣴࠿ࡽ㒊ศⓗ⳦ᚑᒓᰤ㣴ࢆ࡬࡚⳦ᚑᒓᰤ㣴࡬࡜㐍໬ࡋ࡚࠸ࡃ㐣⛬࡛㸪⳦᰿⳦࡜ࡢඹ⏕㛵ಀࡀ࡝ࡢࡼ࠺࡟ኚ໬ࡋ࡚࠸ࡃ
ࡢ࠿ࢆ㏣࠺ࡇ࡜࡟ࡋࡓࠋ㸲Ѹ㸰㸬ࢩࣗࣥࣛࣥᒓ࡛࠾ࡇࡗࡓ⳦᰿⳦ࡢኚ໬ ࡑࢀࡒࢀࡢ✀࡟ࡘ࠸࡚」ᩘࡢ㔝⏕㞟ᅋ࠿ࡽ㸪⳦᰿⳦ࡀឤᰁࡋ࡚࠸ࡿ᰿ࡸᆅୗⱼࢆ᥇㞟ࡋ㸪⳦ࡢ DNA ᝟ሗࢆ฼⏝ࡋࡓศᏊྠᐃἲ࡟ࡼࡗ࡚⳦ࡢ✀㢮ࢆ≉ᐃࡋࡓ㸦ᅗ㸴㸹ヲࡋࡃࡣ
Ogura-Tsujita et al. 2012㸧ࠋࡍࡿ࡜㸪ぢࡘ࠿ࡗࡓ⳦ࡣ኱ࡁࡃ㸰ࢢ࣮ࣝࣉ࡟ࢃ࠿ࢀࡿࡇ࡜ࡀࢃ࠿ࡗࡓࠋ㸯ࡘ┠ࡢࢢ࣮ࣝࣉࡣ㸪ከࡃࡢࣛࣥ⛉᳜≀ࡀඹ⏕┦ᡭ࡜ࡋ࡚࠸ࡿࣛࣥᆺ⳦᰿⳦࡛࠶ࡾ㸦௨㝆 A ࢢ࣮ࣝࣉ࡜࿧ࡪ㸧㸪ࡶ࠺୍ࡘࡣ㸪እ⏕⳦᰿⳦ࡢࢢ࣮ࣝࣉ࡛㸪࣋ࢽࢱࢣ⛉ࡸࣟ࢘ࢱࢣ⛉࡞࡝୺࡟ᶞᮌࡢ୺せ࡞እ⏕⳦᰿⳦࡜ࡋ࡚▱ࡽࢀࡿ⳦㢮࡛࠶ࡗࡓ㸦௨㝆 B ࢢ࣮ࣝࣉ࡜࿧ࡪ㸧ࠋᑐ㇟࡜ࡋࡓ㸳✀࡟࠾࠸࡚㸪ぢࡘ࠿ࡗࡓ⳦ࡢෆヂࢆㄪ࡭ࡓ࡜ࡇࢁ㸪⊂❧ᰤ㣴ࡢ࣊ࢶ࢝ࣛࣥࡣ A ࢢ࣮ࣝࣉࡢ⳦ࡢࡳ࡜ඹ⏕ࡍࡿࡀ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴ࡢ㸰✀ࡣ A ࡜ B ୧᪉ࡢࢢ࣮ࣝࣉ࡜ඹ⏕ࡋ࡚࠾ࡾ㸪⳦ᚑᒓᰤ㣴ࡢ㸰✀ࡣ B ࢢ࣮ࣝࣉࡢࡳ࡜ඹ⏕ࡋ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋࡍ࡞ࢃࡕ㸪⊂❧ᰤ㣴Ѝ㒊ศⓗ⳦ᚑᒓᰤ㣴Ѝ⳦ᚑᒓᰤ㣴࡜࠸࠺ᰤ㣴ᛶࡢኚ໬࡟క࠸㸪⳦᰿⳦ࡢ✀㢮ࡀ AЍ
A+BЍB ࡜ኚ໬ࡋ࡚࠾ࡾ㸪⳦᰿⳦ࡢኚ໬ࡣ A ࠿ࡽ B ࡬㸯ࢫࢸࢵࣉ࡛ኚ໬ࡍࡿࡢ࡛ࡣ࡞ࡃ A+B࡜࠸࠺୰㛫ẁ㝵ࢆ࡬࡚ኚ໬ࡋ࡚࠸ࡓࡇ࡜ࡀࢃ࠿ࡗࡓࡢ࡛࠶ࡿࠋ≉࡟㸪㒊ศⓗ⳦ᚑᒓᰤ㣴✀ࢼࢠ࡛ࣛࣥࡢ⤖ᯝࡣ⯆࿡῝ࡃ㸪௒ᅇㄪᰝࡋࡓ㸰✀ࡢ㒊ศⓗ⳦ᚑᒓᰤ㣴✀ࡢ࠺ࡕ㸪⳦ᚑᒓᰤ㣴✀࡟᭱ࡶ㏆⦕࡞ගྜᡂ✀࡛࠶ࡿࢼࢠࣛࣥࡣ㸪ࡶ࠺୍✀ࡢ㒊ศⓗ⳦ᚑᒓᰤ㣴✀࡛࠶ࡿࢩࣗࣥࣛࣥ
ࡼࡾࡶ B ࢢ࣮ࣝࣉ࡬ࡢ౫Ꮡᗘࡀࡎ࠸ࡪࢇ㧗࠿ࡗࡓࡢ࡛࠶ࡿ㸦ᅗ㸴㸧ࠋ
Y. Ogura-Tsujita - 6

植物科学最前線 5:136 （2014）ᅗ㸴 ㄪᰝࡋࡓࢩࣗࣥࣛࣥᒓ㸳✀࡛ぢࡽࢀࡓ⳦᰿⳦ࡢෆヂࠋヲࡋࡃࡣ Ogura-Tsujita et al. (2012)ཧ↷ࡘࡲࡾ㸪A+B ࡜࠸࠺୰㛫ẁ㝵࡟࠾࠸࡚ࡶ࠸ࢁ࠸ࢁ࡞ẁ㝵ࡀ࠶ࡗ࡚㸪㐍໬ࡢ㐨ࡢࡾࡢ୰࡛ᚎࠎ
࡟ B ࡬ࡢ౫Ꮡᗘࢆ㧗ࡵ࡚࠸ࡗࡓ࡜⪃࠼ࡽࢀࡿࠋ⳦᰿⳦ࡀ኱ࡁࡃኚ໬ࡍࡿ㐣⛬࡟ࡇࡢࡼ࠺࡞୰㛫ẁ㝵ࡀ࠶ࡗࡓࡇ࡜ࡣᮏ◊✲ࡢ኱ࡁ࡞Ⓨぢ࡛࠶ࡾ㸪ࡇࡢࡼ࠺࡞୰㛫ᆺࡀ⳦᰿ඹ⏕ࡢ≉Ṧ໬࡟㠀ᖖ࡟኱ࡁ࡞ᙺ๭ࢆᯝࡓࡋ࡚࠸ࡿ࡟㐪࠸࡞࠸ࠋ ࡑࡢᚋ㸪⚾㐩ࡣࢩࣗࣥࣛࣥᒓࡢ୰࡛㸪ගྜᡂࢆ⾜࠺⊂❧ᰤ㣴✀ࡢࢫࣝ࢞ࣛࣥ࡜㸪ගྜᡂࢆࡋ࡞࠸⳦ᚑᒓᰤ㣴✀ࡢ࣐ࣖࣛࣥࡢ㸰✀ࢆ஺㓄ࡋ㸪㞧✀ࢆࡘࡃࡿࡇ࡜࡟ᡂຌࡋࡓ
㸦Ogura-Tsujita et al. 2014㸧ࠋ୧ぶࡢᙧ㉁ࢆ࠶ࢃࡏࡶࡘ㞧✀㞟ᅋࢆ࠺ࡲࡃ౑࠺ࡇ࡜࡟ࡼࡗ࡚㸪⳦᰿⳦ࡢኚ໬ࢆࡣࡌࡵ㸪ගྜᡂࢆࡸࡵࡿ㐍໬ࡢ㐨ࡢࡾ࡟࡝ࡢࡼ࠺࡞㑇ఏᏊࡢኚ໬ࡀ࠶ࡗࡓࡢ࠿ࢆㄪ࡭ࡿࡇ࡜ࡀ࡛ࡁࡿࠋ⌧ᅾ㸪ࡇࡢ㞧✀ࡣࣇࣛࢫࢥࡢ୰࡛⥡㯇࡞ⰼࢆࡉ࠿ࡏ࡚࠾ࡾ㸦ᅗ㸵㸧㸪௒ᚋࡢᒎ㛤ࡀᮇᚅࡉࢀࡿࠋ㸳㸬௒ᚋࡢᒎᮃ ⳦ᚑᒓᰤ㣴᳜≀࡟ࡣࡲࡔࡲࡔㅦࡀከ࠸ࠋ௒ࡶୡ⏺୰࡛◊✲ࡀ㐍⾜ࡋ࡚࠸ࡿࡀ㸪኱㔞ࡢ DNA ᝟ሗࢆ▷᫬㛫࡛ゎᯒ࡛ࡁࡿࢩࢫࢸ࣒࡛࠶ࡿḟୡ௦ࢩ࣮ࢡ࢚ࣥࢧ࣮࡜㸪᳜≀࡜⳦᰿⳦ࢆ୍⥴࡟ヨ㦂⟶ࡢ୰࡛ᇵ㣴ࡍࡿඹ⏕ᇵ㣴⣔ࢆά⏝ࡍࡿࡇ࡜࡟ࡼࡗ࡚㸪௒ᚋࡉࡽ࡟࠾ࡶࡋࢁ࠸ࡇ࡜ࡀࢃ࠿ࡾࡑ࠺ࡔࠋ⳦ᚑᒓᰤ㣴᳜≀ࡢඹ⏕┦ᡭ࡛࠶ࡿ⳦᰿⳦ࡣ㸪࡝ࡇ࡛࡝ࡢࡼ࠺࡟ᬽ

Y. Ogura-Tsujita - 7ᅗ㸵 ヨ㦂⟶ࡢ୰࡛ⰼࢆဏ
࠿ࡏࡿ⊂❧ᰤ㣴✀ࢫࣝ࢞ࣛࣥ࡜⳦ᚑᒓᰤ㣴✀࣐ࣖࣛࣥࡢ㞧✀㸦෗┿ᥦ౪㸹㕥ᮌ࿴ᘯ㸧

植物科学最前線 5:137 （2014）ࡽࡋ࡚࠸ࡿࡢࡔࢁ࠺࠿㸽⳦᰿⳦ࡣ୺࡟ᆅ୰࡛⏕⫱ࡋ࡚࠸ࡿࡓࡵ㸪ࡑࡢศᕸࡸᬽࡽࡋࡪࡾࡣ࡯࡜ࢇ࡝᝟ሗࡀ࡞࠸ࡢࡀ⌧≧࡛࠶ࡿࠋ⌧ᅾ㸪ḟୡ௦ࢩ࣮ࢡ࢚ࣥࢧ࣮ࢆ౑ࡗ࡚㸪᥇ྲྀࡋࡓᅵተ୰࡟࠸ࡿࡍ࡭࡚ࡢ⳦㢮ࢆㄪ࡭ୖࡆࡿࡇ࡜ࡀྍ⬟࡟࡞ࡗ࡚ࡁࡓ㸦Bu«e et al. 2009㸧ࠋ࣓ࢱࢤࣀ࣒࡜
࿧ࡤࢀࡿࡇࡢࡼ࠺࡞ᢏ⾡ࢆ౑ࡗ࡚㸪ㅦࡵ࠸ࡓ⳦᰿⳦ࡢ⏕ែࡢ୍➃ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚ࡃࡿ࡟㐪࠸࡞࠸ࠋࡲࡓ㸪ḟୡ௦ࢩ࣮ࢡ࢚ࣥࢧ࣮ࡢᢏ⾡ࢆ౑࠼ࡤ㸪⳦ᚑᒓᰤ㣴᳜≀࡜⊂❧ᰤ㣴᳜≀ࡢࡶࡘ㛗࠸㛗࠸ DNA ᝟ሗࢆ┤᥋ẚ㍑ࡍࡿࡇ࡜ࡶྍ⬟࡟࡞ࡿࠋࡇࡢࡼ࠺࡞ẚ㍑◊✲࡟ࡼࡗ࡚㸪㐍໬ࡢ㐨ࡢࡾ࡟కࡗ࡚࠾ࡇࡗࡓᵝࠎ࡞ᙧ㉁ኚ໬࡟࡝ࡢࡼ࠺࡞㑇ఏᏊࡀ㛵ࢃࡗࡓࡢ࠿ࡀぢ࠼࡚ࡃࡿࡣࡎࡔࠋ ḟୡ௦ࢩ࣮ࢡ࢚ࣥࢧ࣮ࡢࡼ࠺࡞᭱᪂ᢏ⾡ࡶࡉࡿࡇ࡜࡞ࡀࡽ㸪➨㸲❶࡛⤂௓ࡉࢀ࡚࠸ࡿࡼ࠺࡞ඹ⏕ᇵ㣴⣔ࡶከࡃࡢㅦࢆゎỴࡋ࡚ࡃࢀࡿ࡟㐪࠸࡞࠸ࠋ㏻ᖖ㸪Ⅳ⣲ࡣ᳜≀࠿ࡽ⳦᰿⳦࡬ཷࡅΏࡉࢀࡿࡀ㸪⳦ᚑᒓᰤ㣴᳜≀࡛ࡣⅣ⣲ࡢ⛣ື᪉ྥࡀ㏫㌿ࡋ㸪⳦࠿ࡽ᳜≀࡬⛣ືࡍࡿࠋࡇࡢ≀㉁㍺㏦ࡢヲࡋ࠸࣓࢝ࢽࢬ࣒ࡢゎ᫂ࡣ㸪㔝እ࡛⾜࠺ࡢࡣ㠀ᖖ࡟ᅔ㞴࡛࠶ࡾ㸪ඹ⏕ᇵ㣴⣔ࢆ⏝࠸ࡓ◊✲࡛᫂ࡽ࠿࡟࡞ࡗ࡚ࡃࡿ࡟㐪࠸࡞࠸ࠋࡲࡓ㸪ࡇࡢࡼ࠺࡞ᇵ㣴⣔ࢆ౑ࡗ࡚㸪᳜≀࡜⳦࡜ࡢㄆ㆑ᶵᵓࡀ࡝ࡢࡼ࠺࡟ኚ໬ࡋࡓࡢ࠿࡞࡝⏕⌮ⓗ࡞⌧㇟ࢆヲࡋࡃゎ᫂ࡍࡿࡇ࡜ࡀ࡛ࡁࡿࠋ ගྜᡂࢆࡸࡵࡿ㐍໬ࡢ㐨ࡢࡾࡣ㸪ᐇ࡟ከᵝ࡞ᙧ㉁ࡢኚ໬ࢆక࠺」㞧࡞㐨ࡢࡾ࡛࠶ࡿࠋ᳜≀ࡀ㸪࡞ࡐ㸽࡝࠺ࡸࡗ࡚㸽ගྜᡂࢆࡸࡵࡓࡢ࠿㸪ࡑࡢ␲ၥ࡟⟅࠼ࡿࡓࡵ࡟ࡣ㸪᳜≀⏕⌮Ꮫ㸪ᙧែᏛ㸪⳦㢮Ꮫ㸪⏕ែᏛ࡟⣔⤫ศ㢮Ꮫ࡞࡝㸪ᐇ࡟ከᵝ࡞Ꮫၥࢆࡲࡓ࠸࡛◊✲ࢆ⾜࠺ᚲせࡀ࠶ࡿࠋ⳦ᚑᒓᰤ㣴᳜≀ࡢ▱ྡᗘࡣࡲࡔࡲࡔపࡃ㸪௒ᚋᵝࠎ࡞ศ㔝ࡢ◊✲⪅ࡀᮏ᳜≀࡟⯆࿡ࢆᣢࡕ㸪ከ㠃ⓗ࡞࢔ࣉ࣮ࣟࢳ࡛㐍໬ࡢ㐨ࡢࡾࡀゎ᫂ࡉࢀ࡚⾜ࡃࡇ࡜ࢆᮇᚅࡋࡓ࠸ࠋࡲࡓ㸪⳦࡟㢗ࡿ⏕ࡁ᪉ࡺ࠼㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ⎔ቃࡢኚ໬࡟㠀ᖖ࡟ᙅࡃ㸪ከࡃࡢ✀ࡀ⤯⁛ࡢ༴ᶵ࡟ℜࡋ࡚࠸ࡿࠋ⌧ᅾࡣᙼࡽࢆᏲࡿࡍ࡭ࡀ࡯࡜ࢇ࡝࡞࠸ࡓࡵ㸪ୡ⏺ࡢྛᡤ࡛ࡦࡗࡑࡾ࡜ጼࢆᾘࡋ࡚࠸ࡿࡢࡀ⌧≧ࡔࠋ௒ᚋࡢ◊✲ᡂᯝ࡟ࡼࡗ࡚㸪ᮏ᳜≀ࢆಖ඲ࡍࡿ㐨ࡀ㛤࠿ࢀࡿࡇ࡜ࢆ㢪ࡗ࡚ࡸࡲ࡞࠸ࠋ䢢ㅰ㎡䢢 䢢 ᮏ✏࡛⤂௓ࡋࡓ◊✲ࡢ୍㒊ࡣ᪥ᮏᏛ⾡᣺⯆఍࣭⛉Ꮫ◊✲㈝⿵ຓ㔠㸦21370038, 24370040,2440185㸧ࡢຓᡂࢆཷࡅ࡚⾜ࢃࢀࡓࠋᮏ◊✲ࢆ㐙⾜ࡍࡿ࡟࠶ࡓࡾࡈᣦᑟ࠸ࡓࡔ࠸ࡓඛ⏕᪉㸪ࢧࣥࣉࣝ᥇㞟࡟ࡈ༠ຊ࠸ࡓࡔ࠸ࡓⓙᵝ࡟ཌࡃᚚ♩⏦ࡋୖࡆࡿࠋࡲࡓ㸪ᮏ✏ᇳ➹࡟࠶ࡓࡾࢥࢣ᳜≀ࡢ⣔⤫࡟ࡘ࠸࡚ࡈຓゝ࠸ࡓࡔ࠸ࡓᴬཎᜨᏊẶ࡟῝ㅰࡍࡿࠋ䢢ᘬ⏝ᩥ⊩Abe, J.P. 2005. An arbuscular mycorrhizal genus in the Ericaceae. Abstracts of the Mycological Society ofAmerica/ the Mycological Society of Japan Joint Meeting 2005, Hilo, Hawaii.

Bidartondo, M.I., Redecker, D., Hijri, I., Wiemken, A., Bruns, T.D., Dominguez, L., Sersic, A., Leake,J.R., & Read, D.J. 2002. Epiparasitic plants specialized on arbuscular mycorrhizal fungi. Nature419: 389-392.
Bidartondo, M.I., & Bruns, T.D. 2002. Fine-level mycorrhizal specificity in the Monotropoideae(Ericaceae): specificity for fungal species groups. Mol. Eco. 11: 557-569.
Bidartondo, M.I. 2005. The evolutionary ecology of myco-heterotrophy. New Phytol. 167: 335-352.

Y. Ogura-Tsujita - 8

植物科学最前線 5:138 （2014）

Bidartondo, M.I., Bruns, T.D., Weiss, M., Se´rgio, C., & Read, D.J. 2003. Specialized cheating of theectomycorrhizal symbiosis by an epiparasitic liverwort. Proc. R. Soc. Lond. B 270: 835-842.
Buée, M., Reich, M., Murat, C., Morin, E., Nilsson, R.H., Uroz, S., & Martin, F. 2009. 454 pyrosequencing
analyses of forest soils reveal an unexpectedly high fungal diversity. New Phytol. 184: 449-456.

Cullings, K.W., Szaro, T.M., & Bruns, T.D. 1996. Evolution of extreme specialization within a lineageof ectomycorrhizal epiparasites. Nature 379: 63̽66.
Davis, E.C. 2004. A molecular phylogeny of leafy liverworts ( Jungermannidae: Marchantiophyta). In:Goffinet, B., Hollowell, V., & Magill, M.R. (eds.) Molecular Systematics of Bryophytes (MSB 98). pp.
61-86. Missouri Botanical Garden, St. Louis, MO, USA.
Franke, T., Beenken, L., Döring, M., Kocyan, A., & Agerer, R. 2006. Arbuscular mycorrhizal fungi of theGlomus-group A lineage (Glomerales; Glomeromycota) detected in mycoheterotrophic plants fromtropical Africa. Mycol. Prog. 5: 24-31.
Hamada, M. 1939. Studien uber die Mykorrhiza von Galeola septentrionalis Reichb. f. Ein neuer Fall derMykorrhiza-Bildung durch intraradicale Rhizomorpha. Jpn. J. Bot. 10: 151-211.
Kottke, I., & Nebel, M. 2005. The evolution of mycorrhiza-like associations in liverworts: an update. NewPhytol. 167: 330-334.
Kusano, S. 1911. Gastrodia elata and its symbiotic association with Armillaria mellea. J. Coll. Agric. Univ.
Tokyo 4: 1-66.
Merckx, V., & Bidartondo, M.I. 2008. Breakdown and delayed cospeciation in the arbuscular mycorrhizal
mutualism. Proc. R. Soc. B. 275: 1029-1035.
Merckx, V., Janssens, S.B., Hynson, N.A., Specht, C.D., Bruns, T.D., & Smets, E.F. 2012.
Mycoheterotrophic interactions are not limited to a narrow phylogenetic range of arbuscular mycorrhizal
fungi. Mol. Ecol. 21: 1524-1532.
Merckx, V.S.F.T., Mennes, C.B., Peay, K.G., & Geml, J. 2013. Evolution and diversification. In: Merckx,V.S.F.T. (ed) Mycoheterotrophy: the biology of plants living on fungi. pp. 215-244. Springer, New York.
Molvray, M., Kores, P.J., & Chase, M.W. 2000. Polyphyly of mycoheterotrophic orchids and functional
influences on floral and molecular characters. In: Wilson, K.L. & Morrison, D.A. (eds.) Monocots:systematics and evolution. pp. 441-448. Commonwealth Scientific and Industrial Research Organisation
Publishing, Melbourne, Australia.
Motomura, H., Selosse, M.A., Martos, F., Kagawa, A., & Yukawa, T. 2010. Mycoheterotrophy evolved
from mixotrophic ancestors: Evidence in Cymbidium (Orchidaceae). Ann. Bot. 106: 573-581.
Obase, K., Matsuda, Y., & Ito, S. 2013. Enkianthus campanulatus (Ericaceae) is commonly associated witharbuscular mycorrhizal fungi. Mycorrhiza 23: 199-208.
Ogura-Tsujita, Y., Yokoyama, J., Miyoshi, K., & Yukawa, T. 2012. Shifts in mycorrhizal fungi during theevolution of autotrophy to mycoheterotrophy in Cymbidium (Orchidaceae). Am. J. Bot. 99: 1158-1176.
Ogura-Tsujita, Y., Miyoshi, K., Tsutsumi, C., & Yukawa, T. 2014. First flowering hybrid between
autotrophic and mycoheterotrophic plant species: breakthrough in molecular biology ofmycoheterotrophy. J. Plant Res. 127: 299-305.

Y. Ogura-Tsujita - 9植物科学最前線 5:139 （2014）

Selosse, M.A., Weiß, M., Jany, J.L., & Tillier, A. 2002. Communities and populations of sebacinoid
basidiomycetes associated with the achlorophyllous orchid Neottia nidus-avis (L.) L.C.M. Rich. andneighbouring tree ectomycorrhizae. Mol. Ecol. 11: 1831-1844.
Selosse, M.A., & Roy, M. 2009. Green plants that feed on fungi: facts and questions about mixotrophy.
Trends Plant Sci. 14: 64-70.
ᔱᮧṇᶞ 2012㸬ࢥࢣ᳜≀㸬ᡞ㒊 ༤࣭⏣ᮧ ᐇ (⦅) ᪂ࡋ࠸᳜≀ศ㢮ᏛII. pp. 1-12.ㅮㄯ♫㸬ᮾி㸬Smith, S.E., & Read, D.J. 2008. Mycorrhizal symbiosis, third edition, Academic press, San Diego.
Taylor, D.L., & Bruns, T.D. 1997. Independent, specialized invasions of ectomycorrhizal mutualism
by two nonphotosynthetic orchids. Proc. Natl. Acad. Sci. USA 94: 4510-4515.
Waterman, R.J., Klooster, M.R., Hentrich, H., & Bidartondo, M.I. 2013. In: Merckx, V.S.F.T. (ed)Mycoheterotrophy: the biology of plants living on fungi. pp. 267-296. Springer, New York.
Yukawa, T., Miyoshi, K., & Yokohama, J. 2002. Molecular phylogeny and character evolution ofCymbidium (Orchidaceae). Bull. Natl. Sci. Mus. Tokyo 28: 129-139.
Yukawa, T., Ogura-Tsujita, Y., Shefferson, R.P., & Yokoyama, J. 2009. Mycorrhizal diversity in Apostasia(Orchidaceae) indicates the origin and evolution of orchid mycorrhiza. Am. J. Bot. 96: 1997-2009.

Y. Ogura-Tsujita - 10