植物科学最前線 5:110 （2014）⳦ᚑᒓᰤ㣴᳜≀ࡢ⳦᰿ඹ⏕⣔ࡢከᵝᛶ㇂ட㧗ᗈ໭ᾏ㐨኱Ꮫ໭᪉⏕≀ᅪࣇ࢕࣮ࣝࢻ⛉Ꮫࢭࣥࢱ࣮᳜≀ᅬࠛ໭ᾏ㐨ᮐᖠᕷ୰ኸ༊໭  ᮲す  ୎┠㸦⌧ᡤᒓ㸸ᅜ❧⛉Ꮫ༤≀㤋⟃Ἴᐇ㦂᳜≀ᅬࠛࡘࡃࡤᕷኳஂಖ 㸧Takahiro YagameDiversity of mycorrhizal symbioses in mycoheterotrophic plants
Key words: arbuscular mycorrhizal fungi, ectomycorrhizal fungi, saprophytic fungi,
symbiotic culture, tripartite symbiosis
Field Science Center for Northern Biosphere Hokkaido University Botanic Garden Kita3
Nishi8, Chuo-ku, Sapporo, Hokkaido 060-0003
Current address: Tsukuba Botanical Garden, National Museum of Nature and Science
4-1-1, Amakubo, Tsukuba, Ibaraki 305-0005, Japan㸯㸬ࡣࡌࡵ࡟ 㝣ୖ᳜≀ࡢከࡃࡣගྜᡂࢆ⾜࠺ࡇ࡜࡛⊂❧ᰤ㣴⏕άࢆ㏦ࡾ㸪⏕⏘⪅࡜ࡋ࡚⏕ែ⣔ࢆᨭ࠼࡚࠸ࡿࠋ࡜ࡇࢁࡀ㸶๭௨ୖࡢ᳜≀✀ࡢ᰿࡟┿⳦㢮ࡢ⳦⣒ࡀᐃ╔ࡋ㸦Trappe 1987, Wang & Qiu
2006㸧㸪≉␗ⓗ࡞ᵓ㐀㸦⳦᰿㸧ࡀᙧᡂࡉࢀࡿࠋ㌟㏆࡞౛࡜ࡋ࡚࣐ࢶࢱࢣ࡜࣐ࢶ⛉᳜≀ࡢ㛵ಀࡀᣲࡆࡽࢀࡿࡔࢁ࠺ࠋ࣐ࢶࢱࢣࡣ࣐ࢶ⛉᳜≀ࡢ᰿࡟࠾࠸࡚⳦᰿ඹ⏕ࡍࡿ⳦㢮㸦⳦᰿⳦㸧࡛㸪࣐ࢶ⛉᳜≀࡟ࡣᅵተ୰ࡢ❅⣲ࡸࣜࣥ➼ࢆ౪⤥ࡋ㸪୍᪉࡛࣐ࢶ⛉᳜≀࠿ࡽࡣගྜᡂ⏘≀ࡀ⳦య࡟౪⤥ࡉࢀ࡚࠸ࡿ㸦㕥ᮌ 2005㸧ࠋࡇࡢࡼ࠺࡞ඹ⏕⣔ࢆ᳜≀࡜⳦㢮ࡀᵓ⠏ࡍࡿࡇ࡜࡛㸪᳜≀ࡣᆅ୰࠿ࡽࡢ㣴ศ྾཰ຠ⋡ࢆ㣕㌍ⓗ࡟㧗ࡵࡿࡇ࡜ࡀ࡛ࡁ㸪⳦᰿⳦ࡣⅣ⣲໬ྜ≀ࢆᏳᐃࡋ࡚⋓ᚓ࡛ࡁࡿࠋࢢ࣒ࣟࢫ㛛㸪ᢸᏊ⳦㛛㸪Ꮚᄞ⳦㛛ࡢ⳦㢮ࡀࡇࡢඹ⏕⣔ࢆᵓ⠏ࡍࡿ㸦Peterson et al. 2004㸧ࠋගྜᡂࢆ⾜࠺᳜≀࡜⳦㢮ࡢඹ⏕⣔ࡣ㸪཮᪉࡟࡜ࡗ࡚᭷┈࡛࠶ࡿࠋࡋ࠿ࡋ㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪⳦᰿⳦࡟ගྜᡂ⏘≀ࢆΏࡍࡇ࡜ࡣ࡛ࡁࡎ㸪୍᪉ⓗ࡟⳦࠿ࡽ㣴ศࢆ཰ዣࡋ⏕⫱ࡍࡿ࡜ࡉࢀࡿࠋ⳦ᚑᒓᰤ㣴᳜≀ࡢ⳦᰿ඹ⏕⣔࡟㛵ࡍࡿ◊✲ࡣ㸪⳦᰿⳦ࡢศᏊྠᐃࡸ㸪Ⅳ⣲࡜❅⣲ࡢᏳᐃྠ఩యศ್ูࡢィ࡟ࡼࡿ⳦ᚑᒓᰤ㣴ᛶࡢホ౯ࡀ☜❧ࡋࡓࡇ࡜࡛㸪㏆ᖺ㸪ᵝࠎ࡞᳜≀✀ࢆᑐ㇟࡟◊✲ࡀ࡞ࡉࢀࡿࡼ࠺࡟࡞ࡗ࡚ࡁࡓ㸦Merckx et al. 2013㸧ࠋ㸰㸬⳦ᚑᒓᰤ㣴᳜≀ࡢ⳦᰿ඹ⏕ ⳦ᚑᒓᰤ㣴᳜≀ࡢ⳦᰿ඹ⏕ࡣ㸪኱ࡁࡃ 3 ࡘࡢࢱ࢖ࣉ࡟ศ㢮ࡉࢀࡿࠋ1 ࡘ┠ࡣᶞᮌ࡟͂እ⏕⳦᰿̓࡜࿧ࡤࢀࡿᵓ㐀ࢆᙧᡂࡋඹ⏕ࡍࡿ͂እ⏕⳦᰿⳦̓࠿ࡽ㣴ศࢆᚓ࡚⏕⫱ࡍࡿࢱ࢖ࣉ࡛࠶ࡿࠋࢶࢶࢪ⛉ࡢࢠࣥࣜࣙ࢘ࢯ࢘ࡸࢩࣕࢡࢪࣙ࢘ࢯ࢘㸪ࣛࣥ⛉ࡢࢧ࢝ࢿࣛࣥ࡞࡝ࡢ᳜≀ࡀࡇࢀ࡟࠶ࡓࡿ㸦ᅗ 1A, B, C㸧ࠋࡇࢀࡽࡢ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪ᶞᮌࡀ⏘⏕ࡍࡿගྜᡂ⏘≀ࢆ㸪⳦᰿⳦

T. Yagame - 1

植物科学最前線 5:111 （2014） ᅗ 1 እ⏕⳦᰿⳦࡜⳦᰿ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࠋA, ࢠࣥࣜࣙ࢘ࢯ࢘; B, ࢩࣕࢡࢪࣙ࢘ࢯ࢘; C, ࢧ࢝ࢿࣛࣥࢆ⤒⏤ࡋ⋓ᚓࡍࡿࡇ࡜࡟ࡼࡗ࡚⏕⫱ࡋ࡚࠸ࡿ㸦Bidartondo & Bruns 2002, Selosse et al. 2002,
McKendrick et al. 2002㸧
ࠋ ⳦ᚑᒓᰤ㣴᳜≀ࡼࡾ᳨ฟࡉࢀࡿእ⏕⳦᰿⳦ࡢ୺࡞ศ㢮⩌࡟ࡣ㸪ᢸᏊ⳦㛛ࡢ࣋ࢽࢱࢣ⛉㸪ࣟ࢘ࢱࢣ⛉㸪࢖࣎ࢱࢣ⛉➼ࡀ࠶ࡿ㸦Smith & Read 2008㸧ࠋࡉࡽ࡟ࢭ࢖ࣚ࢘ࢩࣙ࢘ࣟ⛉ࢭ࢖ࣚ࢘ࢩࣙ࢘ࣟᒓ㸦Tuber㸧ࡢࡼ࠺࡞Ꮚᄞ⳦㛛ࡀ⳦᰿⳦࡜࡞ࡿ౛ࡶ▱ࡽࢀ࡚࠸ࡿ㸦Selosse et al. 2004㸧ࠋᶞᮌࡣ▷ᮇ㛫࡛ᯤࢀࡿࡇ࡜ࡣ࡞࠸ࡓࡵ㸪⳦ᚑᒓᰤ㣴᳜≀ࡣእ⏕⳦᰿⳦ࢆ௓ࡋ࡚ᶞᮌࡢගྜᡂ
⏘≀ࢆᏳᐃࡋ࡚ཷࡅྲྀࡿࡇ࡜ࡀ࡛ࡁࡿࠋ
2 ࡘ┠࡟㸪ᮌᮦࡸⴠⴥⴠᯞࢆⅣ⣲※࡜ࡋ࡚⏕⫱ࡍࡿ⭉⏕⳦㢮࠿ࡽ㣴ศࢆᚓ࡚⏕⫱ࡍࡿࣛࣥ⛉ࡢ࢜ࢽࣀࣖ࢞ࣛ㸪ࢶࢳ࢔ࢣࣅ㸪ࢱࢩࣟࣛࣥ࡞࡝ࡀᣲࡆࡽࢀࡿ㸦Kusano 1911, Terashita &Chuman 1987, Yamato et al.2005㸹ᅗ 2㸧ࠋ࢜ࢽࣀࣖ࢞ࣛࡸࢶࢳ࢔ࢣࣅࡀᢸᏊ⳦㛛ࡢ࢟ࢩ࣓ࢪ⛉ࢼࣛࢱࢣᒓ㸦Armillaria㸧ࡢ⳦✀࡜⳦᰿ඹ⏕ࡍࡿࡇ࡜ࡣ㸪ྂࡃࡼࡾ▱ࡽࢀ࡚࠸ࡿ㸦Kusano 1911,Hamada 1939㸧ࠋ஦ᐇ㸪⮬⏕ᆅ࡛ᆅ୰࠿ࡽ᥀ࡾฟࡍ࡜㸪᳜≀య⾲㠃࡟ࢼࣛࢱࢣᒓ⳦㢮ࡀᙧᡂࡍࡿ㯮Ⰽࡢ⳦⣒᮰㸦᰿≧⳦⣒᮰㸧ࡢ௜╔ࡀほᐹࡉࢀࡿ᫬ࡀ࠶ࡿࠋࡋ࠿ࡋ Umata et al.㸦2013㸧ࡣ㸪Ⓨⱆ┤ᚋࡢࢶࢳ࢔ࢣࣅࡢᐇ⏕ⱑ࡟ࡣࢼࣛࢱࢣᒓࡣඹ⏕ࡋ࡚࠾ࡽࡎ㸪ࡑࡢ௦ࡾᢸᏊ⳦㛛ࢱ࣐ࢳࣙࣞ࢖ࢱࢣ┠㸦Polyporales㸧࡟ྵࡲࢀࡿᮍグ㍕ࡢ⳦✀ࡀඹ⏕ࡋ㸪⏕⫱ࢆಁࡋࡓࡇ࡜ࢆሗ࿌ࡋ࡚࠸ࡿࠋࡇࡢ◊✲࡟ࡼࡗ࡚㸪ࢶࢳ࢔ࢣࣅࡣࢼࣛࢱࢣᒓ⳦࡜ࡢඹ⏕⣔ࡀ☜❧ࡉࢀࡿ௨๓࡟㸪࠶ࡿ⛬ᗘࡢ኱ࡁࡉࡢⱑࡲ࡛⏕㛗ࡍࡿࡲู࡛ࡢ⳦✀࡜⳦᰿ඹ⏕ࡍࡿ஦ᐇࡀ♧ࡉࢀࡓࠋࡇࡢࡼ࠺࡞⏕㛗㐣⛬࡟࠾ࡅࡿ⳦᰿⳦ࡢࢩࣇࢺࡣ࢜ࢽࣀ࡛ࣖ࢞ࣛࡶ☜ㄆࡉࢀ࡚࠾ࡾ㸦Xu & Mu 1990㸧
㸪ఱᨾࡑࡢࡼ࠺࡞⌧㇟ࡀ㉳ࡇࡿࡢ࠿㸪኱ኚ⯆࿡῝࠸ࠋ௚࡟ࡶ࠸ࡃࡘ࠿ࡢ⳦ᚑᒓᰤ㣴ᛶࡢࣛࣥ⛉ࡣࢡࢾࢠࢱࢣ⛉ࡸࢼࣚࢱࢣ⛉࡞࡝ᵝࠎ࡞⭉⏕ᛶࡢ⳦✀࡜ඹ⏕ࡍࡿࡀ㸦኱࿴࣭㇂ட 2009㸧㸪⭉⏕⳦࡜ࡢ⳦᰿ඹ⏕ࡣ௚ࡢ⛉ࡢ⳦ᚑᒓᰤ㣴᳜≀࡟ࡣぢࡽࢀ࡞࠸ࠋࡑࡶࡑࡶ㸪ࣛࣥ⛉᳜≀ࡣ⥳Ⰽⴥࢆ᭷ࡍࡿ✀࡛ࡶ㸪ࢣࣛࢺࣂࢩࢹ࢕࣒࢘⛉, ࢶࣛࢫࢿࣛ⛉, ࣟ࢘ࢱࢣ⛉࡞࡝ࡢ࠸ࢃࡺࡿࣜࢰࢡ
ࢺࢽ࢔࡜࿧ࡤࢀࡿ⭉⏕ᛶࡢ␗ᢸᏊ⳦㢮࡜⳦᰿ඹ⏕ࡍࡿࡇ࡜ࡀ≉ᚩ࡛࠶ࡿ㸦Smith & Read 2008㸧ࠋ

T. Yagame - 2

植物科学最前線 5:112 （2014）ᅗ 2 ⭉⏕⳦࡜⳦᰿ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࠋA, ࢜ࢽࣀࣖ࢞ࣛ; B, ࢶࢳ࢔ࢣࣅ; C, ࢱࢩࣟࣛࣥ3 ࡘ┠ࡣ͂࢔࣮ࣂࢫ࣮࢟ࣗࣛ⳦᰿⳦̓
㸦AM ⳦㸧࡜ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀࡛࠶ࡿࠋࢢ࣒ࣟࢫ㛛࡟ᒓࡍࡿ AM ⳦࡟ࡼࡿඹ⏕⣔ࡣ㸪㝣ୖ᳜≀ࡢᵝࠎ࡞ศ㢮⩌ࡢ⊂❧ᰤ㣴᳜≀ࡀႠࡴ᭱ࡶᬑ㐢ⓗ࡞⳦᰿ඹ⏕⣔࡛࠶ࡿ㸦Peterson et al. 2004㸧ࠋ⥳Ⰽ᳜≀࡜ AM ⳦࡜ࡢඹ⏕⣔ࡣ≉␗ᛶࡀపࡃ㸪1 ✀㢮ࡢ᳜≀࡟ከᩘࡢ⳦✀ࡀᐃ╔ࡍࡿࠋ୍᪉㸪⳦ᚑᒓᰤ㣴᳜≀࠿ࡽࡶ」ᩘ✀ࡢ AM ⳦ࡀ᳨ฟࡉࢀࡿࡀ㸪᳜≀ࡢ✀ࡈ࡜࡟≉ᐃࡢ✀ࡢ AM ⳦ࡀ㧗㢖ᗘ᳨࡛ฟࡉࢀࡿഴྥࡀᙉࡃ㸪ࡑࡢከࡃࡣ
Schwarzott et al. 㸦2001㸧࡟ࡼࡾᐃ⩏ࡉࢀࡓࢢ࣒ࣟࢫ㛛ࡢ୰ࡢࢢ࣮ࣝࣉ A ࡜ࡼࡤࢀࡿ⣔⤫࡟ྵ
ࡲࢀࡿ㸦Bidartondo et al. 2002; Yamato et al. 2011; Ogura-Tsujita et al. 2013㸧
ࠋAM ⳦࡜⳦᰿ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪࣍ࣥࢦ࢘ࢯ࢘⛉㸪ࣄࢼࣀࢩࣕࢡࢪࣙ࢘⛉㸪ࢥࣝࢩ࢔⛉㸪ࣜࣥࢻ࢘⛉㸪ࣄ࣓ࣁࢠ⛉㸪ࢧࢡࣛ࢖ࢯ࢘⛉࡞࡝ᵝࠎ࡞⛉࡟ࢃࡓࡾ㸦Merckx et al. 2013㸧㸪᪥ᮏ࡟ࡣ࣍ࣥࢦ࢘ࢯ࢘㸪ࢩࣟࢩࣕࢡࢪࣙ࢘ࡸࢱࢾ࢟ࣀࢩࣙࢡࢲ࢖࡞࡝ࡀศᕸࡍࡿ㸦ᅗ 3A, B,C㸧ࠋࡲࡓ㸪ࢩࢲ᳜≀ࡢࣁࢼࣖࢫࣜ⛉ࡸࣄ࢝ࢤࣀ࢝ࢬࣛ⛉࡞࡝ࡢᆅୗ࡛⏕⫱ࡍࡿ㓄അయࡣ㸪ගྜᡂⰍ⣲ࢆᣢࡓ࡞࠸ࡇ࡜࠿ࡽ⳦ᚑᒓᰤ㣴࡜᥎ᐃࡉࢀ㸪ࡇࢀࡽࡶࢢ࣒ࣟࢫ㛛ࢢ࣮ࣝࣉ A ࡢ⳦✀࡜⳦᰿ඹ⏕ࡍࡿࡇ࡜ࡀ▱ࡽࢀ࡚࠸ࡿ㸦Winther & Friedman 2007a; Winther & Friedman 2007b㸧ࠋAM ⳦࡜ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪እ⏕⳦᰿⳦࡟౫Ꮡࡍࡿ✀࡜ྠᵝ࡟㸪⥳Ⰽ᳜≀࠿ࡽ AM
⳦ࢆ㏻ࡌ࡚ගྜᡂ⏘≀ࢆ཰ዣࡋ㸪⏕⫱ࡋ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿࠋ㸱㸬⳦ᚑᒓᰤ㣴᳜≀࡜⳦᰿⳦ࡢඹ⏕⣔ࡢ᳨ド㸱Ѹ㸯㸬ඛ⾜◊✲࡟ᑐࡍࡿ␲ၥ⳦ᚑᒓᰤ㣴᳜≀ࡀ࡝ࡢࡼ࠺࡞ศ㢮⩌ࡢ⳦㢮࠿ࡽ㣴ศࢆᚓ࡚⏕⫱ࡍࡿࡢ࠿࡜࠸࠺␲ၥࡣ㸪⳦᰿⳦ࡢศᏊྠᐃࡀྍ⬟࡟࡞ࡗࡓࡇ࡜࡛㸪✀ࡈ࡜࡟ゎ᫂ࡉࢀࡘࡘ࠶ࡿࠋࡋ࠿ࡋ㸪ᆅ୰࡟Ꮡᅾࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࡢ᰿ࡣᵝࠎ࡞⳦㢮ࡀᏑᅾࡍࡿᅵተ࡟᭚ࡉࢀ࡚࠸ࡿࡓࡵ㸪ศᏊ⏕≀Ꮫⓗᡭἲ࡟ࡼࡗ࡚⳦᰿⳦ࢆྠᐃࡍࡿ㝿㸪᰿⾲㠃࡟௜╔ࡋ࡚࠸ࡿ⳦᰿⳦௨እࡢ⳦✀ࡢ DNA ࡀ PCR ἲ࡟ࡼࡗ࡚ቑᖜࡉࢀࡿྍ⬟ᛶࡀ࠶ࡿࠋࡲࡓ㸪᰿࠿ࡽࡣ」ᩘࡢ⳦✀ࡀ᳨ฟࡉࢀࡿࡇ࡜ࡀከࡃ㸪ᐇ㝿࡟࡝ࡢ⳦✀ࡀ⳦ᚑᒓᰤ㣴᳜≀ࡢ⏕⫱࡟ᙳ㡪ࢆཬࡰࡋ࡚࠸ࡿ࠿࡟ࡘ࠸࡚ࡣᢕᥱࡉࢀ࡚࠸࡞࠸ሙ

T. Yagame - 3

植物科学最前線 5:113 （2014）ᅗ 3 AM ⳦᰿⳦࡜⳦᰿ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࠋA, ࣍ࣥࢦ࢘ࢯ࢘; B, ࢩࣟࢩࣕࢡࢪࣙ࢘; C, ࢱࢾ࢟ࣀࢩࣙࢡࢲ࢖

ྜࡀከ࠸ࠋඛ⾜◊✲࡛ࡣ㸪ᐃ㔞ⓗ࡞ゎᯒࢆ⾜࠺ࡇ࡜࡛㸪ࡼࡾከࡃࡢሷᇶ㓄ิࢹ࣮ࢱࡀᚓࡽࢀ
ࡓ⳦✀ࢆ୺せ࡞⳦᰿⳦࡛࠶ࢁ࠺࡜᥎ᐃࡋ࡚࠸ࡿࠋࡉࡽ࡟Ᏻᐃྠ఩య್ࢆホ౯ࡍࡿࡇ࡜࡛㸪⳦᰿⳦ࡢ⏕⌮ⓗ≉ᛶࢆ᥎ᐃࡍࡿሙྜࡶ࠶ࡿ㸦Yagame et al. 2012㸧ࠋࡋ࠿ࡋ㸪ࡇࡢᐇ㦂ᡭἲࢆࡶࡗ࡚ࡋ࡚ࡶศᏊྠᐃࡉࢀࡓ⳦✀ࡀ⳦ᚑᒓᰤ㣴᳜≀ࡢ⏕⫱࡟ཬࡰࡍᙳ㡪ࢆホ౯࡛ࡁ࡞࠸ࠋࡑࡇ࡛ⴭ⪅ࡣ⳦ᚑᒓᰤ㣴᳜≀࡜⳦᰿⳦ࡢඹ⏕⣔ࢆᵓ⠏ࡋ㸪᳜≀యࡢ⏕⫱ࢆほᐹࡍࡿ◊✲ࢆ⾜ࡗ࡚ࡁࡓࠋࡑࡢ⤖ᯝ㸪ศᏊྠᐃ࡛ࡣゎ࡛᫂ࡁ࡞࠸㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ⏕⫱ẁ㝵ࡈ࡜ࡢ⳦ඹ⏕ࡢᐇయࢆゎ᫂ࡍࡿࡇ࡜࡟ᡂຌࡋࡓࠋ㸱Ѹ㸰㸬⭉⏕⳦࡜ࡢඹ⏕⣔ᵓ⠏ࣛࣥ⛉ࢱࢩࣟࣛࣥᒓࢱࢩࣟࣛࣥࡣ㸪㛵ᮾᆅ᪉ࢆ໭㝈࡜ࡋ㸪⇕ᖏ࢔ࢪ࢔࠿ࡽ࢔ࣇࣜ࢝す㒊ࡲ࡛⮬⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀࡛࠶ࡿࠋ᪥ᮏ࡛ࡣ 6 ᭶ୗ᪪࠿ࡽ 7 ᭶ୖ᪪࡟ⰼࢆဏ࠿ࡏ㸪㒔ᚰ㒊ࡢ⚄♫ࡢቃෆ࡞࡝࡛ࡶほᐹࡍࡿࡇ࡜ࡀ࡛ࡁࡿࠋࢱࢩࣟࣛࣥࡼࡾ⳦᰿⳦ࢆศ㞳ࡋศᏊྠᐃࡋࡓ⤖ᯝ㸪ࢼࣚࢱࢣ⛉ࡢ⳦✀࡜⳦᰿ඹ⏕ࡍࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ㸦Yamato et al. 2005㸧ࠋࡋ࠿ࡋ㸪ࡇࡢඹ⏕⣔ࡢᏑᅾࡣᙜึ᠜␲ⓗ࡟ぢࡽࢀ࡚࠸ࡓࠋᙜ᫬ࣛࣥ⛉᳜≀ࡢ⳦᰿⳦࡜࠸࠼ࡤࣜࢰࢡࢺࢽ࢔࡛࠶ࡿ㸪࡜࠸࠺ඛධほࡀᙉ࠿ࡗࡓࡓࡵࡔࠋࡑࡇ࡛㸪ⴭ⪅ࡣࢱࢩࣟࣛࣥ࠿ࡽศ㞳ࡉࢀࡓ⳦ᰴࡢࡦ࡜ࡘ࡛࠶ࡿ ME1-1 ᰴࢆࢱࢩࣟࣛࣥࡢ✀Ꮚ࡟᥋✀ࡋ㸪✀ᏊࡀⓎⱆ࣭⏕⫱ࡍࡿ࠿ࢆㄪᰝࡋࡓࠋME1-1 ᰴࡣ Yamato et al. 㸦2005㸧ࡢศᏊ⣔⤫ゎᯒࡢ⤖ᯝ㸪࢖ࢾࢭࣥ࣎ࣥࢱࢣ㸦Coprinellus disseminatus㸧࡟㏆⦕࡛࠶ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟
ࡉࢀ࡚࠸ࡓࠋࡑࡇ࡛ⴭ⪅ࡣࡲࡎ ME1-1 ࡢ✀࡛ࣞ࣋ࣝࡢྠᐃࢆ┠ⓗ࡜ࡋ㸪Ꮚᐇయᙧᡂᐇ㦂ࢆ⾜ࡗࡓࠋࡑࡢ⤖ᯝ㸪኱㔞ࡢᏊᐇయࡀᙧᡂࡉࢀ㸪ࡑࡢᙧែⓗ≉ᚩ࠿ࡽ࢖ࢾࢭࣥ࣎ࣥࢱࢣ࡜ྠᐃࡋ
ࡓ㸦Yagame et al. 2008㸧ࠋࡇࡢ⳦ᰴࢆࢱࢩࣟࣛࣥࡢ✀Ꮚ࡟᥋✀ࡋࡓ࡜ࡇࢁ㸪6 㐌㛫ᚋ㸪Ⓨⱆࡀ☜ㄆࡉࢀࡓ㸦ᅗ 4A㸧
ࠋࡑࡢᚋࢫࢺࣟࣥ࠾ࡼࡧሢⱼࡢⓎ㐩࡞࡝ࡀ☜ㄆࡉࢀ㸦ᅗ 4B, C㸧㸪᧛✀ᚋ
26 㐌࡛ሢⱼࡢ㡬㒊࡟ⰼⱆࡀᙧᡂࡉࢀ㸦ᅗ 4D, E㸧㸪᧛✀ᚋ 27 㐌࡛㛤ⰼ࡟⮳ࡗࡓ㸦ᅗ 4F㸧ࠋ

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植物科学最前線 5:114 （2014）ᅗ 4 ࢱࢩࣟࣛࣥࡢ⏕㛗㐣⛬ࠋA, ᧛✀ 6 㐌┠ ▮༳ࡣࣉࣟࢺࢥ࣮࣒࡟௜╔ࡍࡿ⳦᰿⳦ ME1-1 ࡢ⳦⣒᮰; B, ᧛✀ 8 㐌┠ ▮༳ࡣࢫࢺࣟࣥࡢཎᇶ; C, ᧛✀ 12 㐌┠ S:ࢫࢺࣟࣥ; D, ᧛✀ 16 㐌┠ S:ࢫࢺࣟࣥ T:ሢⱼ; E, ᧛✀ 18 㐌┠ᡂ⇍ࡋࡓሢⱼ I:ⰼᗎ; F, ᧛✀ 27 㐌┠ 㛤ⰼ ࡇࡢࡇ࡜࠿ࡽ㸪ࢱࢩࣟࣛࣥࡣ✀ᏊⓎⱆ࠿ࡽ㛤ⰼ࡟⮳ࡿࡲ࡛༢୍ࡢࢼࣚࢱࢣ⛉ࡢ⳦✀࡜⳦᰿ඹ⏕ࡋ⏕⫱ࡍࡿࡇ࡜ࡀྍ⬟࡞ࡇ࡜ࡀ᫂ࡽ࠿࡜࡞ࡾ㸪ຍ࠼࡚Ⰻዲ࡞⏕⫱⎔ቃୗ࡛ࡣ✀ᏊⓎⱆ࠿ࡽഹ࠿༙ᖺ࡛㛤ⰼࡍࡿࡇ࡜ࡀ᫂ࡽ࠿࡜࡞ࡗࡓ㸦Yagame et al. 2007㸧ࠋࡉࡽ࡟㸪1 ⢏ࡢ✀Ꮚ࠿ࡽ༙ᖺ࡛኱ᑠ⣙ 80 ಶࡶࡢሢⱼࡀᙧᡂࡉࢀࡓࡇ࡜࠿ࡽ㸪㧗࠸ᰤ㣴⦾Ṫ⬟ຊࢆ᭷ࡍࡿࡇ࡜ࡀ☜ㄆࡉࢀࡓࠋࢱࢩࣟࣛࣥࡢ⏕㛗㐣⛬ࡣ㸪Burgeff 㸦1932㸧ࡸ Van Leeuwen㸦1937㸧ࡽ࡟ࡼࡾᐇ⏕ⱑࡢ≉ᚩࡀሗ࿌ࡉࢀ࡚࠸ࡓࡀ㸪ᮏ◊✲࡟ࡼࡗ࡚Ⓨⱆ࠿ࡽ㛤ⰼ࡟⮳ࡿ⏕㛗㐣⛬ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋࡲࡓ㸪ࣛࣥ⛉᳜≀࡜ࢼࣚࢱࢣ⛉ࡀ⳦᰿ඹ⏕ࡍࡿ஦ᐇࡀึࡵ࡚☜ㄆࡉࢀࡓࠋࡉࡽ࡟ࣛࣥ⛉࢖ࣔࢿࣖ࢞ࣛᒓࡢ⳦ᚑᒓᰤ㣴✀࢖ࣔࢿࣖ࢞ࣛࡀ㸪ࢼࣚࢱࢣ⛉ࡢ⳦✀࡜≉␗ⓗ࡟⳦᰿ඹ⏕ࡍࡿ஦ᐇࡀศᏊྠᐃ࡟ࡼࡾ᫂ࡽ࠿࡟ࡉࢀࡓࡇ࡜࠿ࡽ㸦Ogura-Tsujita & Yukawa 2008㸧㸪ࢼࣚࢱࢣ⛉࡜ࡢඹ⏕ࡀ᳜≀ࡢ⳦ᚑᒓᰤ㣴ᛶ㐍໬࡜㛵㐃ࡍࡿࡇ࡜ࡀ♧၀ࡉࢀࡓࠋ ࡑࢀ࡛ࡣࠊࢼࣚࢱࢣ⛉࡜ඹ⏕ࡍࡿ⥳Ⰽⴥࢆ᭷ࡍࡿࣛࣥࡣᏑᅾࡍࡿࡢࡔࢁ࠺࠿ࠋࢧ࢖ࣁ࢖ࣛࣥࡣ⥳Ⰽⴥࢆ᭷ࡍࡿ✀ࡔࡀ㸪ⷧᬯ࠸ᯘᗋ࡟⏕⫱ࡍࡿࡓࡵ⊂❧ᰤ㣴ࡔࡅ࡛⏕⫱ࡍࡿࡇ࡜ࡀ㞴ࡋ࠸࡜⪃࠼ࡽࢀࡓࠋࢧ࢖ࣁ࢖ࣛࣥࡀⷧᬯ࠸⎔ቃ࡟㐺ᛂ࡛ࡁࡿ⌮⏤ࢆ⳦᰿ඹ⏕࠿ࡽゎ᫂ࡋ࡚ࡳࡼ࠺࡜⪃࠼㸪ࡲࡎ⳦᰿⳦ࢆศᏊྠᐃࡋࡓࠋᆅ୰࡛ࢧࣥࢦ≧࡟ศᯞࡍࡿ᰿ⱼࡼࡾ⳦ࢆศ㞳ᇵ㣴ࡋࡓ࡜ࡇࢁ㸪」ᩘࡢ⳦ᰴࡀࢼࣚࢱࢣ⛉࡜ྠᐃࡉࢀࡓࠋศ㞳⳦ᰴࡢ࠺ࡕ 1 ᰴࡣ㸪Ꮚᐇయࢆᙧᡂࡉࡏࡿࡇ࡜࡟ᡂຌࡋ㸪ࢼࣚࢱࢣ⛉ࡢ Coprinellus domesticus ࡜ྠᐃࡉࢀࡓ㸦Yagame et al. 2013㸧ࠋࡲࡓศᏊ⣔⤫ゎᯒࡢ⤖ᯝ㸪ࡇࡢ⳦ࡣࢱࢩࣟࣛࣥࡼࡾศ㞳ࡉࢀࡓ⳦᰿⳦࡟ᴟࡵ࡚㏆⦕࡛࠶ࡿࡇ࡜ࡶ
᫂ࡽ࠿࡟࡞ࡗࡓ㸦ᅗ 5㸧ࠋࡉࡽ࡟ࠊศ㞳ࡋࡓ 2 ⣔⤫ࡢ Coprinellus ᒓ⳦ࢆࢧ࢖ࣁ࢖ࣛࣥࡢ✀Ꮚ࡟ඹ⏕ࡉࡏࡓ⤖ᯝ㸪✀ᏊࡀⓎⱆࡋ᰿ⱼࡀⓎ㐩ࡋࡓࡇ࡜࠿ࡽ㸪⥳Ⰽⴥࢆ᭷ࡍࡿࣛࣥ⛉᳜≀ࡢ୰࡟ࡶࢼࣚࢱࢣ⛉ࡢ⳦✀࡜⳦᰿ඹ⏕ࡍࡿ✀ࡀ࠶ࡿࡇ࡜ࡀド᫂ࡉࢀࡓࠋࢧ࢖ࣁ࢖ࣛࣥࡀ㸪ࢼࣚࢱࢣ

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植物科学最前線 5:115 （2014）ᅗ 5 ࢧ࢖ࣁ࢖ࣛࣥ㸪ࢱࢩࣟࣛࣥ㸪࢖ࣔࢿࣖ࢞ࣛࡢ⳦᰿⳦
࡜ࡑࡢ㏆⦕✀㸦ࢼࣚࢱࢣ⛉㸧ࡢ rDNA ࡢ ITS 㡿ᇦࡢሷᇶ㓄ิ࡟ᇶ࡙ࡃศᏊ⣔⤫ᶞࠋ࣋࢖ࢬἲ࡟ࡼࡾసᡂࠋPP: ஦ᚋ☜⋡, NJBS:㏆㞄⤖ྜἲ࡟ࡼࡿࣈ࣮ࢺࢫࢺࣛࢵࣉ್㸪MPBS㸸᭱኱⠇⣙ἲ࡟ࡼࡿࣈ࣮ࢺࢫࢺࣛࢵࣉ್⛉ࡢࡼ࠺࡞⳦ᚑᒓᰤ㣴᳜≀ࡢ⏕⫱ࢆᨭ࠼࠺ࡿ⳦㢮࡜ඹ⏕ࡍࡿࡇ࡜࠿ࡽ㸪⥳ⴥࢆᣢࡕ࡞ࡀࡽࡶ⳦᰿⳦࡬Ⅳ⣲ࢆ౫Ꮡࡍࡿ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀㸦ヲࡋࡃࡣ➨㸰❶ཧ↷㸧࡛࠶ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿࠋࢧ࢖ࣁ࢖ࣛࣥࡢࡇࡢࡼ࠺࡞ᛶ㉁ࡀ㸪ⷧᬯ࠸ᯘᗋ⎔ቃ࡟࠾ࡅࡿ⏕⫱ࢆྍ⬟࡟ࡋ࡚࠸ࡿࡢ࠿ࡶࡋࢀ࡞࠸ࠋ㸱Ѹ㸱㸬እ⏕⳦᰿⳦࡜ࡢඹ⏕⣔ᵓ⠏ᶞᮌ㸪እ⏕⳦᰿⳦㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ 3 ⪅ඹ⏕⣔ࢆேᕤⓗ࡟ᵓ⠏ࡋࡓ஦౛ࡣ㸪㐣ཤ࡟ᩘ౛࠶ࡿ࡟㐣ࡂ࡞࠸ࠋWarcup 㸦1985㸧ࡣ㸪࣮࢜ࢫࢺࣛࣜ࢔࡟⮬⏕ࡍࡿࣛࣥ⛉᳜≀ Rhizanthella
gardneri ࡢⱑࢆ 3 ⪅ඹ⏕ୗ࡛ 15 㐌㛫᱂ᇵࡋ㸪㛤ⰼ࡟ᡂຌࡋ࡚࠸ࡿࠋࡲࡓ㸪McKendrick et al.
㸦2000㸧ࡣ࣮ࣘࣛࢩ࢔኱㝣࠿ࡽ໭⡿኱㝣࡟࠿ࡅ࡚ᗈࡃศᕸࡍࡿࣛࣥ⛉᳜≀࡛࠶ࡿ Corallorhizatrifida ࡢᐇ⏕ⱑࢆ 3 ⪅ඹ⏕ୗ࡛ 28 㐌㛫⏕⫱ࡉࡏ㸪14C ࡢィ࡟ࡼࡾᶞᮌ࠿ࡽእ⏕⳦᰿⳦ࢆ⤒⏤ࡋࡓⅣ⣲ࡢ⛣ື࡜⏕⏘㔞ࡢቑຍࢆ☜ㄆࡋ࡚࠸ࡿࠋࡉࡽ࡟㸪Bougoure et al. 㸦2010㸧ࡣ
Rhizanthella gardneri ࢆ 3 ⪅ඹ⏕ୗ࡛⣙ 1 㐌㛫⫱ᡂࡋ㸪13C ࠾ࡼࡧ 15N ࢆ⏝࠸࡚ᐟ୺᳜≀࠿ࡽࡢⅣ⣲౪⤥㸪࠾ࡼࡧᅵተ୰࠿ࡽࡢ❅⣲౪⤥࡟࠿࠿ࡿ᫬㛫ࢆィࡋ࡚࠸ࡿࠋࡋ࠿ࡋ㸪ࡇࢀࡽࡢ◊✲࡛ࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ⏕⫱ࢆ㛗ᮇⓗ࡟ほᐹ࡛ࡁ࡚࠸࡞࠸ࠋࡲࡓ㸪⳦ᚑᒓᰤ㣴ࡢ⛬ᗘࡢ㧗࠸✀ࡢࡳࡀ⏝࠸ࡽࢀ࡚࠾ࡾ㸪⥳Ⰽⴥࢆᣢࡘ㒊ศⓗ⳦ᚑᒓᰤ㣴✀࡟ࡘ࠸࡚ࡣ 3 ⪅ඹ⏕᮲௳ୗ࡛⏕⫱ࡉࡏࡓ஦౛ࡀ࡞࠸ࠋ ࡑࡇ࡛ⴭ⪅ࡣ㒊ศⓗ⳦ᚑᒓᰤ㣴✀࡛࠶ࡿࡇ࡜ࡀᆏᮏࡽ㸦2013㸧࡟ࡼࡾ☜ㄆࡉࢀ࡚࠸ࡿࣛࣥ⛉࢟ࣥࣛࣥ࡟ࡘ࠸࡚ேᕤⓗ࡟ 3 ⪅ඹ⏕⣔ࢆᵓ⠏ࡍࡿࡇ࡜ࢆヨࡳࡓࠋࡲࡎ㸪࢟ࣥࣛࣥࡢ᰿ࡼ
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植物科学最前線 5:116 （2014）ࡾ⳦᰿⳦ࢆ」ᩘ⣔⤫ศ㞳ࡋࡓࠋศᏊྠᐃࡢ⤖ᯝ㸪ศ㞳ࡉࢀࡓ⳦ᰴࡣࡍ࡭࡚࢖࣎ࢱࢣ⛉࡟ྵࡲࢀࡿ⳦✀࡜ྠᐃࡉࢀࡓ㸦Yagameand Yamato 2013㸧ࠋḟ࡟㸪ศ㞳⳦ᰴࢆ⏝࠸࡚ࢥࢼࣛ㸦ࣈࢼ⛉㸧࡜እ⏕⳦᰿ࢆᙧᡂࡉࡏࡓࠋศ㞳⳦ᰴ࡜እ⏕⳦᰿ࢆᙧᡂࡋࡓࢥࢼࣛࡢⱑ࡜㸪↓⳦ᇵ㣴࡛⏕⫱ࡉࡏࡓ࢟ࣥࣛࣥࡢⱑࢆྠࡌ㖊ࡢ୰࡛ 120 㐌᱂ᇵࡋࡓࠋᑐ↷༊࡛ࡣ⳦᰿⳦ࢆ᥋✀ࡋ࡚࠸࡞࠸ࢥࢼࣛࡢⱑ࡜࢟ࣥࣛࣥࡢ↓⳦ᇵ

ᅗ 6 ࢟ࣥࣛࣥࡢ᰿ࡢ⏕⫱(A-C)㸪࠾ࡼࡧࢩ࣮ࣗࢺࡢఙ㛗ࠋA, ↓⳦ᇵ㣴ⱑ;B, 3 ⪅ඹ⏕ᇵ㣴 6 ࣨ᭶ᚋ; C, 3 ⪅ඹ⏕ᇵ㣴 18 ࣨ᭶ᚋ;D, 3 ⪅ඹ⏕ୗࡢ㛤ⰼ㣴ⱑࢆ୍⥴࡟᱂ᇵࡋࡓࠋ3⪅ඹ⏕ᇵ㣴ࡋࡓ࢟ࣥࣛࣥࢆᅗ 6 ࡟♧ࡍࠋ3 ⪅ඹ⏕⣔ࢆᵓ⠏ࡋ࡚ 6 ࣨ᭶ᚋ㸪⳦᰿⳦᥋✀༊࡜ᑐ↷༊࡛ⱑࡢ⏕Ꮡ⋡࡟ᕪࡣࡳࡽࢀ࡞࠿ࡗࡓࠋࡋ࠿ࡋ㸪30 ࣨ᭶ࡀ⤒㐣ࡍࡿ࡜᥋✀༊࡛ࡣ 7 ๭௨ୖࡢⱑࡀ⏕Ꮡࡋ࡚࠸ࡓࡀ㸪ᑐ↷༊࡛ࡣࡍ࡭࡚ࡢⱑࡀᯤṚࡋ࡚࠸ࡓࡇ࡜࠿ࡽ㸪࢟ࣥࣛࣥࡢ⏕⫱࡟ࡣ㸪㸱⪅ඹ⏕⣔ࢆ௓ࡋࡓ⳦᰿⳦࠿ࡽࡢᰤ㣴౪⤥ࡀ୙ྍḞ࡛࠶ࡿࡇ࡜ࡀ☜ㄆࡉࢀࡓࠋࢥࢼࣛࡢගྜᡂ⏘≀ࡀ⳦᰿⳦ࢆ௓ࡋ࡚࢟ࣥࣛࣥ࡬࡜㐠ࡤࢀ࡚࠸ࡓࡢ࡛࠶ࡿࠋࡲࡓ㸪࢟ࣥࣛࣥᒓࡣ㔝እ࡟࠾࠸࡚ẖᖺᚲࡎࡋࡶࢩ࣮ࣗࢺࢆᙧᡂࡍࡿࢃࡅ࡛ࡣ࡞ࡃ㸪≧ἣ࡟ᛂࡌ࡚ᆅୗ㒊ࡢࡳ࡛⏕Ꮡࡋ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ㸦Abadie et al. 2006㸧ࠋᮏ◊✲࡟ࡼࡗ࡚㸪࢟ࣥࣛࣥࡀ⳦᰿⳦࠿ࡽࡢ㣴ศࢆ౑ࡗ࡚᰿ࡢ⏕㛗ࢆ⥆ࡅ࡚࠸ࡿࡇ࡜ࢆ㸪ᇵ㣴ୗ࡛ึࡵ࡚☜ㄆࡍࡿࡇ࡜ࡀ࡛ࡁࡓࠋ
㸱㸫㸲㸬$0 ⳦࡜ࡢඹ⏕⣔ᵓ⠏ࡇࡇࡲ࡛እ⏕⳦᰿⳦ࡸ⭉⏕⳦࡜⳦ᚑᒓᰤ㣴᳜≀࡜ࡢඹ⏕⣔ࢆேᕤⓗ࡟సࡾฟࡋࡓᐇ㦂⣔ࢆ⤂௓ࡋࡓࡀ㸪AM ⳦࡜ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࢆேᕤ⎔ቃୗ࡛⫱ᡂࡉࡏࡓ஦౛ࡣ▱ࡽࢀ࡚࠸࡞࠸ࠋAM ⳦ࡣ༢⊂࡛ᇵ㣴ࡍࡿࡇ࡜ࡀᅔ㞴࡛㸪ࡉࡽ࡟᳜≀࡜ࡢඹ⏕⣔ࡀᵓ⠏ࡉࢀ࡞࠸࡜⬊Ꮚࢆᙧᡂ࡛ࡁ࡞࠸㸦Peterson et al. 2004㸧ࠋᐟ୺᳜≀㸪AM ⳦㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ 3 ⪅ඹ⏕⣔ࢆᵓ⠏ࡍࡿࡓࡵ࡟ࡣ㸪 AM ⳦ࢆ⏕⫱ࡢ᪩࠸ⲡᮏ᳜≀࡛ඹ⏕ࡉࡏ, ࡑࡇ࡟⳦ᚑᒓᰤ㣴᳜≀ࢆ᳜࠼௜ࡅࡿᚲせࡀ࠶ࡿࠋ㸲㸬࠾ࢃࡾ࡟⳦ᚑᒓᰤ㣴᳜≀ࡢ⳦᰿ඹ⏕⣔ゎ᫂࡟㛵ࡍࡿ◊✲ࡣ㸪༢࡟⳦᰿⳦ࡢྠᐃࢆ⾜࠺ẁ㝵࠿ࡽ㸪᳨ฟࡉࢀࡓ⳦᰿⳦ࡢࡑࢀࡒࢀࡀ⳦ᚑᒓᰤ㣴᳜≀ࡢ⏕⫱࡟࡝ࡢࡼ࠺࡞ᙳ㡪ࢆ୚࠼࡚࠸ࡿ࠿ࢆ᳨ドࡍࡿẁ㝵࡟࡞ࡗ࡚ࡁࡓࠋඹ⏕ᇵ㣴ࡀᵝࠎ࡞⳦ᚑᒓᰤ㣴ᛶ✀࡛ᡂຌࡍࢀࡤ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ

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植物科学最前線 5:117 （2014）⏕ែ࣭⏕⌮࡟ࡘ࠸࡚⌮ゎࡀ῝ࡲࡿࡇ࡜ࡀᮇᚅࡉࢀࡿࠋࡲࡓ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ୰࡟ࡣ⤯⁛ࡀ༴᝹ࡉࢀࡿ✀ࡶከࡃ㸪ࡇࢀࡽࡢಖ඲࣭ቑṪ࡟㈨ࡍࡿ▱ぢࡀ㞟✚ࡉࢀࡿࡇ࡜ࡀᮇᚅࡉࢀࡿࠋㅰ㎡ ᮏ◊✲ࢆ⾜࠺࡟࠶ࡓࡾ㸪ᖇி⛉Ꮫ኱Ꮫ⏕࿨⎔ቃᏛ㒊 ᒾ℩ ๛஧ᩍᤵ㸪༓ⴥ኱Ꮫᩍ⫱Ꮫ㒊 ኱࿴ ᨻ⚽ ຓᩍ㸪ᮾி㒔ᕷ኱Ꮫ▱㆑ᕤᏛ㒊 㕥ᮌ ᙲ ≉௵ᩍᤵ㸪༓ⴥ୰ኸ༤≀㤋 ྿᫓ ಇගୖᖍ◊✲ဨ㸪 Montpellier ኱Ꮫ Marc-André Sellose ᩍᤵ㸪ᅜ❧⛉Ꮫ༤≀㤋㐟ᕝ▱ஂ◊✲ဨ࡞࡝ከࡃࡢ᪉ࠎࡼࡾ◊✲ࡢ㝿ࡢࡈᣦᑟࡸࢧࣥࣉࣜࣥࢢࡢ㝿ࡢࡈຓຊࢆ㈷ࡗࡓࠋࡲࡓ㸪ࢱࢩࣟࣛࣥࡢࢧࣥࣉࣜࣥࢢ࡟ࡘ࠸࡚ࡣ᫂἞⚄ᐑ㸪᷃ཎ⚄ᐑ㸪ୗ㬞⚄♫࡟ࡈ༠ຊࢆ࠸ࡓࡔ࠸ࡓࠋ௨ୖࡢⓙᵝ࡟ᚰࡼࡾឤㅰࡢពࢆ⾲ࡋࡓ࠸ࠋᘬ⏝ᩥ⊩
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