植物科学最前線 5:84 （2014）ගྜᡂࢆࡸࡵࡓ᳜≀࣮⳦ᚑᒓᰤ㣴᳜≀ࡢࡓ࡝ࡗࡓ㐍໬ࡢ㐨ࡢࡾ࣮࢜࢞ࢼ࢖ࢨ࣮
㎷⏣ ᭷⣖ᮾ໭኱Ꮫ᳜≀ᅬࠛ௝ྎᕷ㟷ⴥ༊ᕝෆ 
㸦⌧ᡤᒓబ㈡኱Ꮫ㎰Ꮫ㒊ᛂ⏝⏕≀⛉Ꮫ⛉ࠛబ㈡ᕷᮏᗉ⏫㸯␒ᆅ㸧㐟ᕝ ▱ஂ
ᅜ❧⛉Ꮫ༤≀㤋⟃Ἴᐇ㦂᳜≀ᅬࠛࡘࡃࡤᕷኳஂಖ 
᳜≀ࡢ᳜≀ࡓࡿࡺ࠼ࢇࡣ㸪ගྜᡂ࡟ࡼࡗ࡚⮬ࡽ࢚ࢿࣝࢠ࣮ࢆసࡾฟࡍ⊂❧ᰤ㣴ᛶ࡟࠶ࡾࡲ
ࡍࠋ࡜ࡇࢁࡀගྜᡂࢆࡸࡵ࡚ࡋࡲ࠸㸪ඹ⏕ࡍࡿ⳦㢮࠿ࡽᰤ㣴ࡸỈࢆࡶࡽࡗ࡚ࡢࢇࡁ࡟ᬽࡽࡍ
᳜≀ࡀ㸪ୡ⏺࡟ 880 ✀࡯࡝࠸ࡿࡢ࡛ࡍࠋࡇ࠺ࡋࡓ᪉ἲ࡛⏕ࡁࡿ✀㢮ࢆ⳦ᚑᒓᰤ㣴᳜≀㸦mycoheterotrophic plants㸧࡜ࡼࡧ㸪ࢥࢣ᳜≀࠿ࡽ⿕Ꮚ᳜≀ࡲ࡛ᵝࠎ࡞㝣ୖ᳜≀ࡢ⣔⤫࡛ᗄᗘࡶ㐍໬ࡋ࡚ࡁࡲࡋࡓࠋ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪ࡶ࡜ࡶ࡜ࡣගྜᡂࢆࡍࡿ⊂❧ᰤ㣴᳜≀࡛࠶ࡾ㸪ගྜᡂࢆࡸࡵ࡚࠸ࡃ㐍໬ࡀ࠾ࡇࡗࡓࡇ࡜ࡀࢃ࠿ࡗ࡚࠸ࡲࡍࠋࡲࡓ㸪ࡇࡢගྜᡂࢆࡸࡵࡿ㐍໬ࡢ㐨ࡢࡾ࡛㸪ᐇ࡟ከᵝ࡞ᙧ㉁㐍໬ࡀ࠾ࡇࡿࡇ࡜ࡶࢃ࠿ࡗ࡚ࡁࡲࡋࡓࠋගྜᡂᙧ㉁ࡢኚ໬࡟ຍ࠼㸪ⴥࡀ࡞ࡃ࡞ࡿ㸪✀Ꮚࢧ࢖ࢬࡀ㠀ᖖ࡟ᑠࡉࡃ࡞ࡿ࡞࡝ࡢᴟ➃࡞ᙧែⓗኚ໬㸪ࡉࡽ࡟ࡣ⳦᰿⳦ࡸ
㏦⢊᪻⹸࡞࡝௚ࡢ⏕≀࡜ࡢඹ⏕㛵ಀࡲ࡛ࡶኚ໬ࡍࡿࡓࡵ㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ᳜≀ࡢᵝࠎ࡞ᙧ㉁㐍໬ࢆ◊✲ࡍࡿ㠀ᖖ࡟ࡼ࠸ࣔࢹࣝ࡜࡞ࡾࡲࡍࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ࡇࢀࡲ࡛⳦ᚑᒓᰤ㣴᳜≀ࡣ◊✲ࡢᑐ㇟࡜ࡋ࡚ぢ㐣ࡈࡉࢀ࡚࠸ࡲࡋࡓࠋ
ᮏ⥲ㄝ㞟࡛ࡣ㸪ගྜᡂࢆ࠾ࡇ࡞࠺⊂❧ᰤ㣴࠿ࡽ⳦ᚑᒓᰤ㣴࡬ࡢ㐍໬ࡢ㐨ࡢࡾ࡛࡞࡟ࡀ㉳ࡇࡗࡓ࠿ࢆࡉࡲࡊࡲ࡞どⅬ࡛⤂௓ࡋࡲࡍࠋࡲࡎ➨㸰❶࡛ࡣ⳦ᚑᒓᰤ㣴᳜≀࡟㛵ࡍࡿᇶ♏᝟ሗࢆ⤂௓ࡋ㸪➨㸱❶࡛ཷ⢊ᵝᘧࡢኚ໬㸪ࡑࡋ࡚➨㸲❶࡛ࡣ⳦᰿⳦࡜ࡢඹ⏕⣔ࡢࢲ࢖ࢼ࣑ࢵࢡ࡞ኚ
໬ࡢヰࢆࡈ⤂௓ࡋࡲࡍࠋࡲࡓ㸪⊂❧ᰤ㣴࠿ࡽ⳦ᚑᒓᰤ㣴࡬㐍໬ࡍࡿ㝿ࡢ୰㛫ẁ㝵࡜⪃࠼ࡽࢀ࡚࠸ࡿ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀࡟ࢫ࣏ࢵࢺࢆ࠶࡚㸪➨㸳❶࡛㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢⓎⱆ⏕ែ࡟㛵ࡍࡿ◊✲ࢆ⤂௓ࡋ㸪᭱ᚋ࡟➨㸴❶࡛ࡣ㐍໬ࡢ㐨ࡢࡾ࡛㉳ࡇࡗࡓ࣮࢟࢖࣋ࣥࢺ࡛࠶ࡿ⳦᰿
ඹ⏕ࡢኚ໬࡜௒ᚋࡢᒎᮃࡢヰ࡛⥾ࡵࡃࡃࡾࡲࡍࠋ
᳜≀ࡀ᳜≀ࢆࡸࡵࡿ࡯࡝ࡢ኱⫹࡞ኚ㠉ࢆᐇ⌧ࡋ࡚࠸ࡿ࠿ࡽ࡟ࡣ㸪⳦ᚑᒓᰤ㣴ᛶࢆ⋓ᚓࡍࡿࣉࣟࢭࢫ࡛࠾ࡶࡋࢁ࠸㐍໬ࡀ࠾ࡇࡗ࡚࠸ࡿࡣࡎ࡛ࡍࡀ㸪」㞧࡞㐍໬ࡢ㐨ࡢࡾࢆ⣣ゎࡃ࡟ࡣ㸪
ᐇ࡟ᵝࠎ࡞Ꮫၥศ㔝࠿ࡽከ㠃ⓗ࡞࢔ࣉ࣮ࣟࢳࡀᚲせ࡛ࡍࠋᮏ⥲ㄝࡣ㸪᳜≀Ꮫⓗ࡟኱ኚ࠾ࡶࡋ
ࢁ࠸ᮏ᳜≀ࢆᗈࡃⓙᵝ࡟▱ࡗ࡚࠸ࡓࡔࡁࡓ࠸࡜࠸࠺┠ⓗ࡛௻⏬ࡋࡲࡋࡓࠋࡇࡢࣘࢽ࣮ࢡ࡞⏕≀ᮦᩱ࡟ගࡀ࠶ࡓࡾ㸪ࡉࡲࡊࡲ࡞࢔ࣉ࣮ࣟࢳ࡛◊✲ࡀ㐍ᒎࡍࡿࡇ࡜ࢆ㢪ࡗ࡚ࡸࡳࡲࡏࢇࠋ
ᮏ⥲ㄝࡣ㸪᪥ᮏ᳜≀Ꮫ఍➨ 77 ᅇ኱఍㸦2013 ᖺ 9 ᭶㸧࡛㛤ദࡉࢀࡓࢩ࣏ࣥࢪ࣒࢘ࡢㅮ₇ෆᐜࢆ෌ᵓᡂࡋࡓࡶࡢ࡛ࡍࠋࢩ࣏ࣥࢪ࣒࢘ࡢ㛤ദ࡟࠶ࡓࡗ࡚࠾ୡヰ࡟࡞ࡾࡲࡋࡓ኱఍ᐇ⾜ጤဨࡢඛ⏕᪉㸪ࡑࡋ࡚ࡇࡢࣞࣅ࣮ࣗࢆฟ∧ࡍࡿᶵ఍ࢆ୚࠼࡚ࡃࡔࡉ࠸ࡲࡋࡓᗈሗጤဨࡢඛ⏕᪉࡟ཌࡃ࠾♩⏦ࡋୖࡆࡲࡍࠋ
Y. Ogura-Tsujita & T. Yukawa - 1

植物科学最前線 5:85 （2014）⳦ᚑᒓᰤ㣴᳜≀ࡢ⣔⤫࡜㐍໬㐟ᕝ▱ஂ
ᅜ❧⛉Ꮫ༤≀㤋⟃Ἴᐇ㦂᳜≀ᅬࠛ ࡘࡃࡤᕷኳஂಖ 
Tomohisa Yukawa
Phylogenetics and evolution of mycoheterotrophic plants
Key words: Evolution, mycoheterotrophy, mycorrhizal fungi, nutritional mode, symbiosis
Tsukuba Botanical Garden, National Museum of Nature and Science
4-1-1, Amakubo, Tsukuba, Ibaraki 305-0005, Japan㸯㸬ࡣࡌࡵ࡟ ⳦ᚑᒓᰤ㣴᳜≀࡜ࡣ㸪⏕ࡁࡿࡓࡵ࡟ᚲせ࡞ᰤ㣴ࢆගྜᡂ࡟௦ࢃࡗ࡚ඹ⏕ࡍࡿ⳦࡟౫Ꮡࡍࡿ᳜≀࡛࠶ࡿࠋᆅ⌫ୖࡢ᳜≀ࡢ⣙ 80%ࡢ✀ࡣᆅୗࡢ᰿ࡸⱼ࡛⳦㢮࡜ඹ⏕ࡋ࡚࠾ࡾ㸪ࠕ⳦᰿ඹ⏕ࠖ࡜࿧ࡪࠋ
ࡇࡇ࡛᳜≀ࡣගྜᡂ࡟ࡼࡗ࡚✌࠸ࡔⅣ⣲ࡢ୍㒊ࢆඹ⏕⳦࡟୚࠼㸪⳦ࡣ᳜≀ࡢ⏕Ꮡ࡟Ḟ࠿ࡍࡇ࡜ࡢ
࡛ࡁ࡞࠸❅⣲࡜ࣜࣥࢆ᳜≀࡟୚࠼ࡿ┦฼ඹ⏕ࡀᡂࡾ❧ࡗ࡚࠸ࡿࠋ࡜ࡇࢁࡀࡲࢀ࡟㸪᳜≀ࡀගྜᡂࡢᶵ⬟ࢆኻࡗ࡚⳦࠿ࡽ୍᪉ⓗ࡟ᰤ㣴ࢆዣ࠸ྲྀࡿ㐍໬ࡀ㉳ࡇࡗ࡚࠸ࡿ㸦ᅗ㸯㸧ࠋࡇࢀࡣ᳜≀ࡀ᳜≀ࢆࡸࡵࡿ࡟➼ࡋ࠸㸪ᰤ㣴ᦤྲྀࡢ୍኱࢖ࣀ࣮࣋ࢩࣙࣥ࡜࠸࠼ࡼ࠺ࠋ
⮬ศ࡛ᰤ㣴ࢆసࡽ࡞࠸࡞ࡽ᰿ࡶⴥࡶ࠸ࡽ࡞
࠸ࠋከࡃࡢ⳦ᚑᒓᰤ㣴᳜≀ࡣ⏕ࡁࡿࡓࡵ࡟୙
せ࡞ჾᐁࡀ࡞ࡃ࡞ࡗ࡚ወጁ࡞ጼ࡟ኚࢃࡾᯝ࡚
࡚࠸ࡿࠋ⏕ࡁ≀࡜ࡋ࡚㏥໬ࡋࡓᏑᅾࡢࡼ࠺࡟
ឤࡌࡿࡀ㸪࡯࠿ࡢ⏕≀࠿ࡽᰤ㣴ࢆᶓྲྀࡾࡋ࡚
⏕ࡁࡿࡢࡣ㸪ᐇࡣᕦጁ࡛Ὑ⦎ࡉࢀࡓࡸࡾ᪉࡟
㐪࠸࡞࠸ࠋࡇࡢ⳦ᚑᒓᰤ㣴᳜≀࡟╔┠ࡍࢀࡤ㸪⏕≀ࡢᰤ㣴ᦤྲྀࡸඹ⏕⣔ࡢ㐍໬ࡢゎ᫂࡟᪂ࡓ࡞ගࢆ࠶࡚ࡿࡇ࡜ࡀ࡛ࡁࡿࡣࡎࡔࠋࡇࡇ࡛ࡣ
⳦ᚑᒓᰤ㣴᳜≀ࡢᇶᮏⓗ࡞≉ᛶࢆ⤂௓ࡋࡼ࠺ࠋ㸰㸬ぢ㐣ࡈࡉࢀࡓ࠾࡝ࢁࡁࡢከᵝᛶ㸰Ѹ㸯㸬✀㢮࡜⣔⤫⳦ᚑᒓᰤ㣴᳜≀ࡣ 12 ⛉ 90 ᒓ⣙ 530 ✀ࡀ㸪ୡ⏺ࡢ⇕ᖏࠥளᐮᖏࡢ‵₶࡞ᆅᇦ࡟ศᕸࡍ

ᅗ㸯 ⳦ᚑᒓᰤ㣴᳜≀ࡢᰤ㣴⛣ືࠋࡩࡘ࠺᳜≀ࡣ㸪ගྜᡂ࡛సࡾฟࡋࡓⅣ⣲ࡢ୍㒊ࢆඹ⏕ࡍࡿ⳦࡟୚࠼㸪௦ࢃࡾ࡟⳦ࡣ❅⣲࡜ࣜࣥࢆ᳜≀࡟୚࠼ࡿࠋ࡜ࡇࢁࡀ⳦ᚑ
ᒓᰤ㣴᳜≀ࡣගྜᡂࡍࡿࡇ࡜ࢆࡸࡵ㸪⳦࠿ࡽ୍᪉ⓗ࡟ᰤ㣴ࢆዣ࠸ྲྀࡗ࡚࠸ࡿ

ࡿ㸦Merckx 2013㸧ࠋ᪥ᮏ࡟ࡣ⣙ 70 ✀ࡀศᕸࡋ㸪ୡ⏺࡛ࡶࡗ࡜ࡶከᵝᛶࡢ㧗࠸ᆅᇦࡢࡦ࡜ࡘ࡛࠶ࡿࠋ㌟㏆࡞࡜ࡇࢁ࡛ࡶ㸪ࢠࣥࣜࣙ࢘ࢯ࢘㸦ᅗ㸰㸹ࢶࢶࢪ⛉㸧㸪࣐ࣖࣛࣥ㸦ࣛࣥ⛉㸧࡞࡝⳦ᚑᒓᰤ㣴᳜≀ࡣࡦࡗࡑࡾ࡜ᬽࡽࡋ࡚ࡋ࡚࠸ࡿࠋ

T. Yukawa - 1

植物科学最前線 5:86 （2014）㝣ୖ᳜≀ࡢ኱⣔⤫ࡢ࡛ࣞ࣋ࣝࡳࡿ࡜㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪ⱏ᳜≀㛛㸪ࣄ࢝ࢤࣀ࢝ࢬࣛ⥘㸦ᑠⴥ㢮㸧㸪ࢩࢲ㸦኱ⴥࢩࢲ㸧⥘㸪⌫ᯝ᳜≀⥘㸪⿕Ꮚ᳜≀⥘࡟ฟ⌧ࡋࡓࠋ኱㒊ศࡢ✀ࡣ⿕Ꮚ᳜≀⥘࡟ྵࡲࢀࡿ㸦ᅗ㸱㸧ࠋ⣔⤫ᶞࢆࡼࡾ࡝ࡇࢁ࡟⳦ᚑᒓᰤ㣴ᛶࡢฟ⌧ࡋࡓ࣏࢖ࣥࢺࢆ᥎ᐃࡍࡿ࡜㸪⣙ 50 ᅇࡢ㐍໬ࡀ࠶ࡗࡓ࡜᥎
ᐃ࡛ࡁࡿࠋ⳦ᚑᒓᰤ㣴ᛶࡀࡇࢀࡔࡅࡢᅇᩘࡃࡾ࠿࠼ࡋ㐍໬ࡋࡓ࡜࠸࠺ࡇ࡜ࡣ㸪ࡇࡢᛶ㉁ࢆ⋓ᚓࡍࡿࡇ࡜ࡣ㏵᪉ࡶ࡞ࡃప࠸
☜⋡࡛ࡣ࡞࠸ࡇ࡜ࢆ♧ࡋ࡚࠸ࡿࠋ
ࡲࡓ◊✲ᮦᩱ࡜ࡋ࡚ࡳࢀࡤ㸪⊂❧ᰤ㣴ᛶ࠿ࡽ⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬ࢆ᳨ドࡍࡿ⣔ࡀࡓࡃࡉࢇ
࠶ࡿࢃࡅ࡛㸪ẚ㍑ࡍࡿ࠺࠼࡛ࡓ࠸࡬ࢇ㒔ྜࡀࡼ࠸ࠋ
㸰Ѹ㸰㸬ࡲࡔࡲࡔࡳࡘ࠿ࡿ᪂✀ ୡ⏺ࡢ⳦ᚑᒓᰤ㣴᳜≀ࡢ✀ᩘࡣ㸪ࡇࡢ 20 ᖺ㛫࡛ᛴቑࡋࡓࠋ
᪥ᮏ࡛ࡶḟࠎ࡜᪂✀㸪᪂グ㘓ࡀሗ࿌ࡉࢀ࡚࠸ࡿࠋࡓ࡜࠼ࡤࣛ
ࣥ⛉ࡢࢱࣥࢨ࣡ࢧ࢝ࢿࣛࣥࠋ⚄ዉᕝ┴ࡢ୹ἑᒣ⣔࡛Ⓨぢࡉࢀᅗ㸰 ࢠࣥࣜࣙ࢘ࢯ࢘ࠋ㔛ᒣ࡛ࡼࡃぢ࠿ࡅࡿ⳦ᚑᒓᰤ㣴᳜≀ࠋ඲య
࡟༙㏱᫂ࡢⓑⰍ࡛ගྜᡂࡍࡿࡇ࡜
ࡀ࡛ࡁ࡞࠸ࠋ࣋ࢽࢱࢣ⛉ࡢ⳦࠿ࡽᰤ㣴ࢆࡶࡽࡗ࡚࠸ࡿࡇ࡜ࡀศ࠿ࡗ
࡚࠸ࡿ㸦෗┿ ᮌୗ᫭ᙪ㸧

2008 ᖺ࡟Ⓨ⾲ࡉࢀࡓ㸦Yagame et al. 2008; ᅗ㸲㸧ࠋ㤳㒔ᅪࡢࣁ
࢖࢟ࣥࢢࢥ࣮ࢫ࡛ࡶ࠶ࡾ㸪᫂἞௨᮶ከࡃࡢ᳜≀◊✲⪅ࡀㄪᰝࡋࡓᅵᆅ࡟᪂✀ࡀぢ㏨ࡉࢀ╀ࡗ࡚࠸ࡓࡢ࡛࠶ࡿࠋᾏእ࡟┠ࢆྥࡅࡿ࡜ᒓ࡛ࣞ࣋ࣝ᪂ወ࡞ࡶࡢࡶ࠸ࢁ࠸ࢁⓎぢࡉࢀ࡚࠸ࡿࠋࡓ࡜࠼ࡤ㞝ᅗ㸱 ⳦ᚑᒓᰤ㣴᳜≀ࡢฟ⌧ࡍࡿ⣔⤫ࠋࡓࡔࡋ㸪⌫ᯝ᳜≀⥙࡟ᒓࡍࡿࣃࣛࢩࢱࢡࢫࢫ࣭࢘ࢫࢱ Parasitaxus
usta㸦࣐࢟⛉㸧ࡢ⳦ᚑᒓᰤ㣴ᛶ࡟ࡘ࠸࡚ࡣࡉࡽ࡟◊✲ࡀᚲせ࡜ࡉࢀࡿࠋ㝣ୖ᳜≀ࡢศᏊ⣔⤫ᶞ㸦Soltis et al.
2011 ࡟ࡶ࡜࡙ࡃ㸧ࢆࣜࣇ࢓ࣞࣥࢫ࡜ࡋࡓ㸦ࠗ᪂ࡋ࠸᳜≀ศ㢮Ꮫ I㸦ㅮㄯ♫㸧
࠘p.8 ࡢᅗࢆᨵኚ㸧

T. Yukawa - 2

植物科学最前線 5:87 （2014）ⷁ࡜㞤ⷁࡢ㓄⨨ࡀ㏫㌿ࡋࡓࠕ᥅◚ࡾࠖࡢ᳜≀㸪࣍ࣥࢦ࢘ࢯ࢘
⛉ࡢࣞ࢝ࣥࢻࢽ࢔㸦Lecandonia㸧ࡣዲ౛ࡔࠋ ࡞ࡐከࡃࡢ✀ࡀぢ㏨ࡉࢀ࡚ࡁࡓࡢ࠿㸽⳦ᚑᒓᰤ㣴᳜≀ࡣ㛤ⰼ࣭⤖ᐇᮇࡋ࠿ᆅୖ㒊࡟ฟ࡚࠸࡞࠸ࡇ࡜ࡀ᭱኱ࡢ⌮⏤ࡔࢁ࠺ࠋࡘࡲࡾⓎぢࡢࢳࣕࣥࢫࡀࡁࢃࡵ࡚㝈ࡽࢀ࡚࠸ࡿࡢࡔࠋࡓ࡜࠼
ࡤࣛࣥ⛉ࡢࢱࢩࣟࣛࣥࡣ  ᖺࡢ࠺ࡕᆅୖ㒊࡟ࡏ࠸ࡐ࠸  㐌㛫࡯࡝ࡋ࠿ጼࢆ⌧ࡉ࡞࠸ࠋࡇࢀ࠿ࡽࡶወጁ࡞⳦ᚑᒓᰤ㣴᳜≀ࡀࡲࡔࡲࡔⓎぢࡉࢀࡿࡔࢁ࠺ࠋ
㸰Ѹ㸱㸬࠸ࡲࡔㅦࡵࡃ㢮⦕㛵ಀ ⳦ᚑᒓᰤ㣴᳜≀ࡢ◊✲࡛ࡲࡎࡪࡘ࠿ࡿ㞴ၥࡣ㸪⣔⤫ࡀࡣࡗࡁࡾࡋ࡞࠸ࡇ࡜ࡔࡗࡓࠋ⳦ᚑᒓᰤ㣴ᛶ࡬㐍໬ࡍࡿ࡜ࡉࡲࡊࡲ
࡞ᙧ㉁ࡀࢻࣛࢫࢸ࢕ࢵࢡ࡟ኚ໬ࡍࡿࡓࡵ㸪࠸ࡗࡓ࠸࡝ࡢ᳜≀࡜⦕ࡀ㏆࠸ࡢ࠿ศ࠿ࡽ࡞ࡃ࡞ࡿࠋศ㢮࡟ᅔࡗࡓࡢ࡛㸪⳦ᚑᒓᰤ㣴᳜≀ࡢᒓࡸ⛉ࡀࡓࡃࡉࢇసࡽࢀࡓࠋ
࡜ࡇࢁࡀ  ୡ⣖࡟ධࡗ࡚ከ
ࡃࡢ᳜≀ࡢࢤࣀ࣒ࡢሷᇶ㓄ิ᝟ሗࡀ⵳✚ࡍࡿ࡜࡜ࡶ࡟㸪⳦ᚑᒓᅗ㸲 ࢱࣥࢨ࣡ࢧ࢝ࢿࣛࣥࠋ
⚄ዉᕝ┴ࡢ୹ἑᒣ⣔࡛Ⓨぢࡉࢀ 2008 ᖺ࡟Ⓨ⾲ࡉࢀࡓࠋᱵ㞵᫬ࡢ▷ᮇ㛫ࡔࡅᆅୖ࡟
⌧ࢀࡿࡢ࡛㸪ࡇࢀࡲ࡛ぢ㏨ࡉࢀ࡚࠸ࡓࡢࡔࢁ࠺㸦෗┿ ㇂ட㧗ᗈ㸧ᰤ㣴᳜≀ࡢண᝿ࡉࢀ࡞࠿ࡗࡓ㢮⦕ࡀḟࠎ࡜᫂ࡽ࠿࡟࡞ࡗ࡚ࡁࡓࠋ
ࡵࡊࡲࡋ࠸౛࡜ࡋ࡚⊂❧ᰤ㣴ᛶࡢ࢜ࢮࢯ࢘ᒓ㸦ᅗ㸳㸧࡜⳦ᚑᒓᰤ㣴ᛶࡢࢧࢡࣛ࢖ࢯ࢘ᒓ㸦ᅗ㸴㸧ࡢጜጒ㛵ಀࡢⓎぢࢆ࠶ࡆ࡚࠾ࡇ࠺㸦Fuse & Tamura 2000, Cameron et al. 2003;ᅗ㸵㸧ࠋ࢜ࢮࢯ࢘ᒓࡣ
᪥ᮏᅛ᭷ࡢ࢜ࢮࢯ࢘  ✀ࡢࡳ࡛㸪᪥ᮏࡢࣘࢽ࣮ࢡ࡞⏕≀ᆅ⌮ࢆ㇟ᚩࡍࡿ᳜≀࡜ࡋ࡚ࡼࡃ▱ࡽࢀ࡚ࡁࡓࠋࡋ࠿ࡋ㸪࢔ࢪ࢔࡟ࢃࡎ࠿  ✀ࡀ▱ࡽࢀࡿࢧࢡࣛ࢖ࢯ࢘ᒓ࡜ࡶ࡝ࡶ㸪༢Ꮚⴥ᳜≀ࡢ୰࡛㢮⦕ࡢࡼࡃศ࠿ࡽ࡞࠸᳜≀࡜ࡋ࡚ྲྀࡾṧࡉࢀ࡚࠸ࡓࠋ
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ᅗ㸳 ࢜ࢮࢯ࢘ࠋࡇࢀࡲ࡛㢮⦕ࡀㅦࡔࡗࡓ᪥ᮏᅛ᭷ࡢ᳜≀ࠋ⳦ᚑᒓᰤ㣴ᛶࡢࢧࢡࣛ࢖ࢯ࢘ᒓ࡜ࡶࡗ࡜ࡶ⦕ࡀ㏆࠸ࡇ
࡜ࡀ᭱㏆ࡢ◊✲࡛᫂ࡽ࠿࡟࡞ࡗࡓ


T. Yukawa - 3

ᅗ㸴 ࢧࢡࣛ࢖ࢯ࢘ࠋᅜෆ࡛ࡣ୰㒊ᆅ᪉࡜዆⨾኱ᓥࡢ㝈ࡽࢀࡓሙᡤ࡟ࡋ࠿ศ
ᕸࡋ࡞࠸⌋ࡋ࠸⳦ᚑᒓᰤ㣴᳜≀

植物科学最前線 5:88 （2014）࠸ࡃࡘ࠿ࡢ㑇ఏᏊ㡿ᇦࢆ౑ࡗ࡚༢Ꮚⴥ᳜≀ࡢ㐍໬ࢆ᥈ࡿ◊✲࡛୧⪅ࡢࢹ࣮ࢱࢆຍ࠼ࡿ࡜㸪㦫ࡃ࡭ࡁࡇ࡜࡟ࡶࡗ࡜ࡶ⦕ࡢ㏆࠸ࡶࡢ࡝࠺ࡋ࡟࡞
ࡿࡇ࡜ࡀศ࠿ࡾ㸪࠸ࡲ࡛ࡣࢧࢡࣛ࢖ࢯ࢘⛉࡜࠸࠺ࡦ࡜ࡘࡢ⛉࡟ࡲ࡜ࡵࡽࢀ࡚࠸ࡿࠋࡇ࠺ࡋ࡚⊂❧ᰤ㣴᳜≀࡜⳦ᚑᒓᰤ㣴᳜≀ࡢ࣌࢔ࡀࡣࡗࡁࡾ
ࡍࢀࡤࠊ⊂❧ᰤ㣴ᛶ࠿ࡽ⳦ᚑᒓᰤ㣴ᛶ࡟ኚ໬ࡍ
ࡿࣉࣟࢭࢫ࡛࡝ࢇ࡞㐍໬ࡀ㉳ࡇࡗࡓ࠿᳨ドࡍࡿࡇ࡜ࡀ࡛ࡁࡿࡼ࠺࡟࡞ࡿ㸦Yamato et al. in press㸧ࠋ࡜ࡣ࠸࠼㸪࠸ࡃࡘ࠿ࡢ⳦ᚑᒓᰤ㣴᳜≀࡛ࡣࢤࣀ࣒ࡢሷᇶ⨨᥮㏿ᗘࡀ኱ࡁࡃୖ᪼ࡍࡿࡓࡵ㸪࠸ࡲࡔ࡟☜࠿ࡽࡋ࠸⣔⤫㛵ಀࢆᥥࡃࡇ࡜ࡀ࡛ࡁ࡞
࠸ࠋ⳦ᚑᒓᰤ㣴᳜≀ࡢ⣔⤫ࡢ඲ㇺࡣࡲࡔࡘ࠿ࡵ
࡚࠸࡞࠸ࡢࡀ⌧≧ࡔࠋ㸱㸬ࠕ᰿ࡶⴥࡶ࡞࠸ࠖ㐍໬࡬ࡢ㐨ࡢࡾᅗ㸵 ༢Ꮚⴥ᳜≀ࡢ୰࡛ࡢ࢜ࢮࢯ࢘ᒓ࡜ࢧࢡࣛ
࢖ࢯ࢘ᒓࡢ఩⨨㸦Cameron et al. 2003 ࢆᨵኚ㸧ࠋ㑇ఏᏊࡢ᝟ሗࢆ౑ࡗ࡚ゎᯒࡍࡿࡲ࡛ࠊ୧⪅ࡀጜጒ㛵ಀ࡟࠶ࡿࡇ࡜࡟ࡔࢀࡶẼ࡙࠿࡞࠿ࡗࡓ

㸱Ѹ㸯㸬ࡦ࡜ࡗ࡜ࡧ࡛࡞࠸」㞧࡞㐍໬⳦ᚑᒓᰤ㣴ࡢ✀࡟ࡶࡗ࡜ࡶ⦕ࡢ㏆࠸⊂❧ᰤ㣴✀ࡀࡣࡗࡁࡾࡋࡣࡌࡵࡓࡇ࡜࡛㸪⊂❧ᰤ㣴࠿ࡽ⳦ᚑᒓᰤ㣴࡬ࡢ㐍໬ࡢࣉࣟࢭࢫ࡛ఱࡀ㉳ࡇࡗࡓ࠿ࢆ㧗࠸⢭ᗘ᳨࡛ド࡛ࡁࡿࡼ࠺࡟࡞ࡗࡓࠋ
⊂❧ᰤ㣴✀࡜⳦ᚑᒓᰤ㣴✀ࡢ࣌࢔࣮ࢆ౑ࡗ࡚ẚ㍑ࡋ࡚ࡳࡼ࠺ࠋ⳦࡟ᰤ㣴ࢆ౫Ꮡࡍࡿ㐍໬ࡢ㐣⛬࡛㸪ᬑ㏻ⴥࡢ㏥໬㸪᰿ࡸᆅୗⱼࡢኚ໬㸪✀Ꮚࡢᚤ⣽໬࡜࠸ࡗࡓᙧࡢኚ໬࡟࡜࡝ࡲࡽࡎ㸪␗᫬ᛶࡢⓎ⌧ࡢࡼ࠺࡞Ⓨ⏕࡟㛵ࢃࡿኚ໬㸪⳦᰿ඹ⏕⣔㸦➨㸲❶࡛ヲ㏙㸧ࡸ㏦⢊ඹ⏕⣔㸦➨㸱❶࡛ヲ㏙㸧࡞࡝⏕ែࡢኚ໬㸪ẼᏍࡢ㏥໬㸪ⴥ⥳⣲ࡢ႙ኻ㸪Ⰽ⣲యࢤࣀ࣒ࡢ㏥໬࡞࡝⏕⌮࡟㛵ࢃࡿኚ໬࡜࠸ࡗࡓ
ࡉࡲࡊࡲ࡞ᙧ㉁ࡀ㐍໬ࡋ࡚ࡣࡌࡵ࡚㸪᳜≀ࡣගྜᡂࡏࡎ࡟⏕⫱࡛ࡁࡿࡇ࡜ࡀศ࠿ࡾࡣࡌࡵࡓࠋࡘ
ࡲࡾࠕ᰿ࡶⴥࡶ࡞࠸ࠖ㐍໬ࡣ㸪ᑡᩘࡢ㑇ఏᏊࡢኚ໬࡛㉳ࡇࡿࡼ࠺࡞ࢩࣥࣉࣝ࡞ࡶࡢ࡛࡞࠸ࠋ
ࡉࡽ࡟࠸ࢁ࠸ࢁ࡞⣔⤫ࡢ⳦ᚑᒓᰤ㣴✀ࢆㄪ࡭ࡿ࡜㸪⣔⤫ࡈ࡜࡟␗࡞ࡗࡓᙧ㉁≧ែࡢ⤌ࡳྜࢃࡏ࡛⳦ᚑᒓᰤ㣴ᛶࡀ㐍໬ࡋ࡚࠸ࡿࡇ࡜ࡀศ࠿ࡿ㸦ᅗ㸶㸧ࠋ⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬ࡣࡦ࡜ࡘࡢࣔࢹ࡛ࣝㄝ
࡛᫂ࡁ࡞࠸ࡇ࡜ࡶ᫂ࡽ࠿ࡔࠋᅗ㸶 ⳦ᚑᒓᰤ㣴᳜≀ࡢయࡢࡘࡃࡾࡣ✀࡟ࡼࡗ࡚኱ࡁࡃ␗࡞ࡿࠋࢥࣥࢪ࢟ࣖ
࢞ࣛ㸦ࣛࣥ⛉㸧ࡢᆅୗ㒊㸦ᕥ㸧ࠋ⫧኱ࡋࡓᆅୗⱼ࡛⳦࡜ඹ⏕ࡋ᰿ࢆࡘࡃࡽ࡞
࠸ࠋ࣍ࣥࢦ࢘ࢯ࢘㸦࣍ࣥࢦ࢘ࢯ࢘⛉㸧ࡢᆅୗ㒊㸦ྑ㸧ࠋ㛗ࡃ᰿ࢆࡢࡤࡋ᰿࡛⳦࡜
ඹ⏕ࡍࡿ 㸦෗┿ ᮌୗ᫭ᙪ㸧

T. Yukawa - 4

植物科学最前線 5:89 （2014）㸱Ѹ㸰㸬㐍໬ࡢ୰㛫ẁ㝵࣭㒊ศⓗ⳦ᚑᒓᰤ㣴 ࡇࢀࡲ࡛᳜≀ࡢ⊂❧ᰤ㣴࠿ࡽ⳦ᚑᒓᰤ㣴࡬ࡢ㐍໬ࡣ㸪࣡ࣥࢫࢸࢵࣉࡢ༢⣧࡞஦㇟࡜ࡋ࡚ᢅࢃࢀ
࡚ࡁࡓࠋࡋ࠿ࡋࠕⴥࡢ㏥໬ࠖ࡜࠸ࡗࡓࡦ࡜ࡘࡢᙧ㉁ࡢ㐍໬࡛ࡣ࡞ࡃ㸪ከᩘࡢ⊂❧ࡋࡓᙧ㉁ࡢ㐍໬ࡀ⳦ᚑᒓᰤ㣴ᛶࡢᡂ❧࡟ᚲ㡲࡛࠶ࡿࡇ࡜ࡀࡣࡗࡁࡾࡋࡓࡇ࡜࠿ࡽࡶ㸪┦ᙜ࡞᫬㛫ࢆ࠿ࡅ࡚㐍໬ⓗࠕᐇ㦂ࠖࢆࡋ࡞ࡀࡽ᏶඲࡞⳦ᚑᒓᰤ㣴ᛶࢆᡭ࡟ධࢀࡓ࡟㐪࠸࡞࠸ࠋࡇ࠺⪃࠼࡚⊂❧ᰤ㣴࠿ࡽ⳦ᚑᒓᰤ㣴࡬㐍໬ࡍࡿࣉࣟࢭࢫࢆ࡚࠸ࡡ࠸࡟ㄪ࡭ࡿ࡜㸪᏶඲࡞⳦ᚑᒓᰤ㣴࡬㐍໬ࡍࡿ‽ഛẁ㝵࡜ᛮࢃࢀࡿ≧ែࡀぢ࠼ጞࡵࡓࠋ ࡓ࡜࠼ࡤ⳦ᚑᒓᰤ㣴✀࡜⦕ࡢ㏆࠸⊂❧ᰤ㣴✀ࡢᰤ㣴ᦤྲྀࢆㄪ࡭ࡿ࡜㸪ࡳ࠿ࡅࡣࡩࡘ࠺࡟ගྜᡂ
ࢆࡋ࡚࠸ࡿࡼ࠺࡟ぢ࠼ࡿࡶࡢࡢ㸪⳦࠿ࡽ࠿࡞ࡾࡢᰤ㣴ࢆࡶࡽࡗ࡚࠸ࡿࡇ࡜ࡀศ࠿ࡾࡣࡌࡵࡓࠋࡇ
ࡢඃᰂ୙᩿࡞ᰤ㣴ࡢ࡜ࡾ᪉ࢆࠕ㒊ศⓗ⳦ᚑᒓᰤ㣴ࠖ࡜࠸࠺㸦ᅗ㸷㸧ࠋ⊂❧ᰤ㣴࡜⳦ᚑᒓᰤ㣴࡟⏤᮶ࡍࡿⅣ⣲ࡢయෆ࡛ࡢ๭ྜࡣ✀࡟ࡼࡗ࡚㐪࠺ࡋ㸪⏕άྐࡢẁ㝵㸪⏕⫱⎔ቃ࡟ࡼࡗ࡚ࡶኚࢃࡿࡇ࡜ࡀ
ࣛࣥ⛉ࢩࣗࣥࣛࣥᒓ㸦ᅗ 㸧ࡸࢧ࢝ࢿࣛࣥ㐃ࢆ౑ࡗࡓ◊✲࡞࡝࡛ศ࠿ࡾጞࡵࡓ㸦Motomura et al.
2010 ࡞࡝㸹➨㸳❶࡛ヲ㏙㸧ࠋᅗ㸷 㒊ศⓗ⳦ᚑᒓᰤ㣴ࡢ࢖࣓࣮ࢪࠋ᳜≀యࢆᵓᡂࡍࡿⅣ⣲ࡢ⏤᮶ࡣගྜᡂ⏘≀࡜⳦࡟኱ูࡉࢀࡿࠋᕥ➃ࡣ᏶඲࡞⊂❧ᰤ㣴ࠋྑ➃ࡣ᏶඲࡞⳦ᚑᒓᰤ㣴ࠋ㸰ࡘࡢ㛫࡟ࡉࡲࡊࡲ࡞ࣞ࣋ࣝࡢ㒊ศⓗ⳦ᚑᒓᰤ㣴ࡀᏑᅾࡍࡿ ᳜≀࡟࡜ࡗ࡚⊂❧ᰤ㣴ࢆ᏶඲࡟ᤞ࡚ࡿࠕ࢝࣋ࠖࡣ㧗࠸ࡔࢁ࠺ࠋ㒊ศⓗ⳦ᚑᒓᰤ㣴ࡢẁ㝵࡛ࡉࡲࡊࡲ࡞ᙧ㉁㐍໬ࡢࡺࡽࡂࡀチᐜࡉࢀࡿࣉࣟࢭࢫࡀ㸪⳦ᚑᒓᰤ㣴࡟㐺ࡋࡓᙧ㉁≧ែࡢ⤌ࡳྜࢃࡏࡢ⋓ᚓ࡟㔜せ࡞ࡢ࡛ࡣ࡞࠸ࡔࢁ࠺࠿ࠋ࡜ࡍ
ࢀࡤ㸪⊂❧ᰤ㣴࠿ࡽ⳦ᚑᒓᰤ㣴࡬୍㊊㣕ࡧ࡟ࡣ㐍ࡲࡎ㸪⊂❧ᰤ㣴э㒊ศⓗ⳦ᚑᒓᰤ㣴э⳦ᚑᒓᰤ㣴࡜࠸࠺㐨ࡢࡾࡀࡼࡾᬑ㐢ⓗ࡛࠶ࡿྍ⬟ᛶࡀ㧗࠸ࠋ㸱Ѹ㸱㸬ඹ⏕⳦ࡢከᵝᛶゝ࠺ࡲ࡛ࡶ࡞ࡃ⳦ᚑᒓᰤ㣴᳜≀ࡀ⏕ࡁ࡚࠸ࡃࡓࡵ࡟ࡣ㸪ᰤ㣴ࢆศࡅ࡚ࡃࢀࡿẼ๓ࡢࡼ࠸⳦ࡢࣃ࣮ࢺࢼ࣮ࡀᚲ㡲ࡔࠋࡇࢀࡲ࡛㸪ࡁࢃࡵ࡚㝈ࡽࢀࡓࢢ࣮ࣝࣉࡢ⳦ࡀ⳦ᚑᒓᰤ㣴᳜≀ࡢඹ⏕⪅࡜ぢ࡞ࡉࢀ࡚࠸ࡓࡀ㸪ศᏊྠᐃἲࢆ฼⏝ࡋࡓࢹ࣮ࢱࡢ⵳✚࡛㸪ᢸᏊ⳦㛛㸪Ꮚᄞ⳦㛛㸪ࢢ࣒ࣟࢫ㛛࡟ࡲࡓᅗ  ࢩࣗࣥࣛࣥࠋ㟷ࠎ࡜ࡋࡓⴥࢆࡓࡃࡉࢇ╔ࡅṇᖖ࡞ගྜᡂ⬟ࢆᣢࡘࡀ㸪ಶయ࡟ࡼࡗ࡚ࡣయෆࡢ ⛬ᗘࡢⅣ⣲ࡣඹ⏕ࡍࡿ⳦࠿ࡽዣࡗ࡚࠸ࡿࡇ࡜ࡀศ࠿ࡗࡓ㸦෗┿ ୰ᒣ༤ྐ㸧

T. Yukawa - 5

植物科学最前線 5:90 （2014）ࡀࡿᗈ⠊࡞ศ㢮⩌ࡢ⳦ࡀඹ⏕ࡍࡿࡇ࡜ࡀศ࠿ࡗ
࡚ࡁࡓ㸦➨㸲❶࡛ヲ㏙㸧ࠋ࠾ࡶࡋࢁ࠸ࡇ࡜࡟㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ⊂❧ᰤ㣴᳜≀ࡼࡾࡎࡗ࡜ከᵝ࡞⳦㢮ࡢࢢ࣮ࣝࣉ࡜ඹ⏕㛵ಀࢆྲྀࡾ⤖ࢇ࡛࠸ࡿࠋ⳦ᚑᒓᰤ㣴᳜≀ࡢ⳦᰿⳦ࡢ୰࡟ࡣ⊂❧ᰤ㣴᳜≀࡜ࡣࡅࡗࡋ࡚ඹ⏕ࡋ࡞࠸ࢢ࣮ࣝࣉࡀࡋࡤࡋࡤ⌧ࢀࡿࠋࡓ࡜࠼ࡤปᝏ࡞
᳜≀⑓ཎ⳦࡜࡞ࡿࢼࣛࢱࢣᒓ㸦ᅗ 㸸࢟ࢩ࣓ࢪ⛉㸧ࡸⴠⴥศゎ⳦࡛࠶ࡿ࣍࢘ࣛ࢖ࢱࢣ⛉࡞࡝ࡔ㸦Ogura-Tsujita et al. 2009 ࡞࡝㸧ࠋࡘࡲࡾ⊂❧ᰤ㣴᳜≀ࡣ㸪㸲൨ᖺࡢṔྐࢆ࠿ࡅ࡚⠏ࡁୖࡆ࡚ࡁ
ࡓ⳦࡜ࡢඹ⏕㛵ಀ࡟ᏳఫࡍࡿಖᏲⓗ࡞Ꮡᅾ࡛㸪ᅗ  ࢼࣛࢱࢣࡢᏊᐇయࠋ࠸ࡃࡘ࠿ࡢ⳦ᚑᒓᰤ
㣴᳜≀ࡣ㸪ࢼࣛࢱࢣࡢࡼ࠺࡟ᙉ࠸ᐤ⏕ᛶ㸪⑓ཎᛶࢆᣢࡘ⳦ࡍࡽ㣫࠸࡞ࡽࡋ࡚ඹ⏕ࡍࡿ㸦෗┿ ⣽▮๛㸧ᑐࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࡣࣜࢫࢡࢆෑࡋࡘࡘ㨩ຊⓗ࡞ࣃ࣮ࢺࢼ࣮ࢆồࡵ࡚ࡉࡲࡼ࠺ࢻࣥࣇ࢓࡛ࣥ࠶
ࡿࠋ㸲㸬ࡳ࠼࡚ࡁࡓඹ⏕⣔ࡢᐇయ ୺ᙺࡢ᳜≀࡜⳦ࡢ㢦ࡀぢ࠼࡚ࡁࡓ࡜ࡇࢁ࡛㸪ḟ࡟୧⪅ࡢసࡾฟࡍඹ⏕⣔࡟ὀ┠ࡋ࡚ࡳࡼ࠺ࠋ ࠸ࡲ࠾ヰࡋࡓࡼ࠺࡟⳦ᚑᒓᰤ㣴᳜≀ࡣ඲య࡜ࡋ࡚ࡉࡲࡊࡲ࡞ࢢ࣮ࣝࣉࡢ⳦࡜ඹ⏕ࡍࡿࡀ㸪ࡑࢀࡒࢀࡢ✀࡟╔┠ࡍࡿ࡜㸪㝈ࡽࢀࡓ✀ࡢ⳦ࡔࡅ࡜ඹ⏕ࡍࡿࡇ࡜ࡀࡩࡘ࠺ࡔࠋࡓ࡜࠼ࡤ㞄ࡾྜࡗ࡚ ✀ࡢ⳦ᚑᒓᰤ㣴᳜≀ࡀ⏕࠼࡚࠸࡚ࡶ㸪ྠࡌ⳦࡜ඹ⏕ࡍࡿࢃࡅ࡛ࡣ࡞࠸㸦ᅗ 㸧ࠋࡶࡗ࡜ࡶ⦕ࡢ㏆࠸⊂❧ᰤ㣴᳜≀࡜⳦ᚑᒓᰤ㣴᳜≀ࡢ࣌࢔࡛ࡑࢀࡒࢀࡀඹ⏕ࡍࡿ⳦ࡢከᵝᛶࢆẚ࡭ࡿ࡜㸪⳦ᚑᒓᰤ㣴᳜≀ࡢඹ⏕⳦ࡢከᵝᛶࡣ᱁ẁ࡟పࡃ࡞ࡿ㸦Ogura-Tsujita et al.
2012; Yamato et al, in press㸧ࠋ⳦ᚑᒓᰤ㣴᳜≀ࡣࣃ࣮ࢺࢼ࣮ࡢ
࠼ࡾዲࡳࡀ⃭ࡋ࠸ࡢࡔࠋࡑࡢ⌮⏤࡜ࡋ࡚ᐤ୺࡜ᐤ⏕⪅ࡢ཮᪉࡛㐍໬ࡢࠕ㌷ᣑ➇தࠖࡀ㉳ࡇࡗ࡚࠸ࡿࡇ࡜ࡀ⪃࠼ࡽࢀࡿࡀ㸪ࡲࡔド᫂ࡉࢀ࡚࠸࡞࠸ࠋ
ඹ⏕ࡍࡿ⳦ࡢ✀㢮ࡀ㝈ᐃࡉࢀࢀࡤ㸪᳜≀ࡣ≉ᐃࡢ⳦ࡢ✀㢮࡜࠺ࡲࡃࡸࡗ࡚࠸ࡃࡓࡵࡢ㐺ᛂࡀ㔜せ࡟࡞ࡿࡔࢁ࠺ࠋࡑ
ࡢⱞປࢆ㧟㧮ࡉࡏࡿࡢࡀ㸪⏕άྐࡢࢫࢸ࣮ࢪ࡟ࡼࡗ࡚⳦᰿
⳦ࡀࢩࣇࢺࡍࡿ⌧㇟࡛࠶ࡿ㸦➨㸳❶࡛ヲ㏙㸧ࠋࡓ࡜࠼ࡤ࢜ࢽ
ࣀࣖ࢞ࣛ㸦ᅗ 㸧ࡣࢼࣛࢱࢣ࡜ඹ⏕ࡍࡿࡇ࡜࡛᭷ྡࡔࡀ㸪ࡇࢀࡣ⏕⫱ࡀ㐍ࢇ࡛࠿ࡽࡢࡇ࡜࡛㸪Ⓨⱆ࠿ࡽᐇ⏕ࡢึᮇ࡛ࡣࢡࢾࢠࢱࢣᒓࡢ⳦࡜ࡋ࠿ඹ⏕ࡋ࡞࠸ࡇ࡜ࡀศ࠿ࡗࡓ㸦Xu& Guo 2000㸧
ࠋ᝿ീࢆࡓࡃࡲࡋࡃࡍࢀࡤ㸪᭱ᙉࡢ᳜≀⑓ཎ⳦ࢼࣛࢱࢣࡣ㸪ࡦᙅ࡞ᐇ⏕ࡢࣃ࣮ࢺࢼ࣮࡟ࡣ࡞ࡾ࠼ࡎ㸪ⴠࡕⴥࡢศゎ⪅ࢡࢾࢠࢱࢣ࡟⏕⫱ึᮇࡢ㣴⫱ࢆࡺࡔࡡ࡚࠸ࡿࡼ࠺࡟ࡳ
T. Yukawa - 6

ᅗ  ᖖ⥳ᗈⴥᶞᯘࡢᯘᗋ࡛࡜
࡞ࡾ࠶ࡗ࡚ဏࡃ  ✀ࡢ⳦ᚑᒓᰤ㣴᳜≀㸪ࢩࣕࢡࢪࣙ࢘ࢯ࢘㸦ࢶࢶࢪ⛉㸧࡜࣍ࣥࢦ࢘ࢯ࢘㸦࣍ࣥࢦ࢘ࢯ
࢘⛉㸧ࠋ୧⪅ࡢ⳦᰿⳦ࡣࡲࡗࡓࡃู
ࡢ✀㢮࡛࠶ࡿ

植物科学最前線 5:91 （2014）ᅗ 㸸࢜ࢽࣀࣖ࢞ࣛࠋ㧗ࡉ 1.5m ࡟ࡶ࡞ࡾ㸪⳦ᚑᒓᰤ㣴᳜≀࡜ࡋ࡚ࡣࡎࡤࡠࡅ࡚኱ࡁ࠸ࠋ⏕άྐࡢึᮇ
ࡣࢡࢾࢠࢱࢣᒓ㸪ᚋᮇࡣࢼࣛࢱࢣᒓ࡜ࣃ࣮ࢺࢼ࣮ࢆኚ࠼ࡿࠋࢼࣛࢱࢣࡢᕧ኱࡞ࣂ࢖࣐࢜ࢫ࡟ᨭ࠼ࡽࢀ࡚㸪࢜ࢽࣀࣖ࢞ࣛࡣࡢࡧࡢࡧ࡜⏕⫱ࡋ࡚࠸ࡿࡼ࠺࡟ぢ࠼ࡿ㸦෗┿ ᮌୗ᫭ᙪ㸧࠼ࡿࠋ࠸ࡎࢀ࡟ࡏࡼ㸪᠓ࢁ࡟ࡋ࡚࠸ࡓࣃ࣮ࢺࢼ࣮ࡀኚࢃࡿࡢࡣࠕ࣡ࢣ࠶ࡾࠖ࡞ࡢࡣ☜࠿ࡔࠋ࡛ࡣ᳜≀ࡢ⊂❧ᰤ㣴࠿ࡽ⳦ᚑᒓᰤ㣴࡬ࡢ㐍໬࡜࠸࠺኱ࡁ࡞ࢱ࢖࣒ࢫࢣ࣮࡛ࣝ㸪⳦᰿ඹ⏕⣔ࡣ࡝࠺ኚࢃࡗࡓࡢࡔࢁ࠺ࠋࡉࡁ࡯࡝⊂❧ᰤ㣴э㒊ศⓗ⳦ᚑᒓᰤ㣴э⳦ᚑᒓᰤ㣴࡜࠸࠺㐍໬ࡢࢩࢼࣜ࢜
ࢆ⤂௓ࡋࡓࡀ㸪ࢩࣗࣥࣛࣥᒓࢆ౑ࡗࡓ◊✲࡛ࡇࢀࡽࡢࢫࢸ࣮ࢪࡈ࡜࡟ඹ⏕ࡍࡿ⳦ࡢ✀㢮ࡢ⤌ࡳྜࢃࡏࡀኚࢃࡿࡇ࡜ࡀศ࠿ࡗࡓ㸦Ogura-Tsujita et al. 2012㸧ࠋ㒊ศⓗ⳦ᚑᒓᰤ㣴ࡢẁ㝵࡛᪂ࡋࡃඹ⏕ࢆጞࡵࡓ⳦ࡢ✀㢮ࡀ㸪⳦ᚑᒓᰤ㣴ࡢ㐍໬ࡢᘬࡁ㔠࡟࡞ࡗ࡚࠸ࡿࡇ࡜ࢆ♧ࡍࢹ࣮ࢱࡔࡀ㸪ࡇࡇ࡛ࡶ㒊ศⓗ⳦ᚑᒓᰤ㣴ࡀ⳦ᚑᒓᰤ㣴ࢆ㔊ᡂࡍࡿ⯙ྎ࡟࡞ࡿࡇ࡜ࢆ⿬௜ࡅ࡚࠸ࡿ㸦➨㸴❶࡛ヲ㏙㸧ࠋ㸳㸬ࡇࢀ࠿ࡽࡢ◊✲ࡇࡇࡲ࡛࠾ヰࡋࡋࡓෆᐜࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ࡯ࢇࡢ୍㒊ࡢ⣔⤫࠿ࡽᚓࡽࢀࡓࡶࡢ࡟㐣ࡂ࡞࠸ࠋ᳜≀ࡢ⳦ᚑᒓᰤ㣴ᛶ㐍໬ࡢண᝿ࢆࡣࡿ࠿࡟㉸࠼ࡓ」㞧࡞ᐇయࢆ᫂ࡽ࠿࡟ࡍࡿ࡟ࡣ㸪ከࡃࡢ⣔⤫ࢆ౑ࡗࡓ⥙⨶ⓗ࡞◊✲ࡀ୙ྍḞࡔࠋࡲࡓ⳦ᚑᒓᰤ㣴᳜≀ࡣ⳦᰿ඹ⏕⣔࡟࠸ࡢࡕࢆクࡍࡣ࠿࡞࠸Ꮡᅾ
࡛㸪ከࡃࡢ✀ࡣ⤯⁛ࡢ࠾ࡑࢀࡀ࠶ࡿࠋಖ඲ࢆ㐍ࡵࡿࡓࡵ࡟ࡶ㸪ࡑࢀࡒࢀࡢ✀ࡢ⏕≀Ꮫⓗ࡞≉ᛶࢆ᫂ࡽ࠿࡟ࡋ࡞ࡅࢀࡤ࡞ࡽ࡞࠸ࠋ
'1$ ࡸᏳᐃྠ఩యࢆ౑ࡗࡓゎᯒᡭἲࡢ㐍ᒎࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ◊✲࡟ࣈࣞ࢖ࢡࢫ࣮ࣝࢆࡶࡓ
ࡽࡋࡓࠋ࡜ࡇࢁࡀࡲࡔ኱ࡁ࡞ၥ㢟ࡀṧࡉࢀ࡚࠸ࡿࠋᆅୗࡢືែࡔࠋ⳦ᚑᒓᰤ㣴᳜≀ࡢ࠾ࡶ࡞⏕ά✵㛫ࡣ㸪⳦࡜ඹ⏕ࡍࡿᅵࡢ୰࡟࠶ࡿࠋᆅୖ࡟ฟࡿࡢࡣ㛤ⰼ࣭⤖ᐇࡍࡿᩘ༑᪥࡟ࡍࡂ࡞࠸ࠋ⳦ᚑᒓ
ᰤ㣴᳜≀ࢆࡼࡾࡼࡃ▱ࡿ࡟ࡣ㸪ࡇࡢᆅ୰ࡢࠕࣈࣛࢵࢡ࣎ࢵࢡࢫࠖࡢ㒊ศࡢゎ᫂ࡀ࢝ࢠࡔࢁ࠺ࠋ
ࠕࣈࣛࢵࢡ࣎ࢵࢡࢫࠖࢆྍど໬ࡍࡿ࡟ࡣ㸪⮬⏕ᆅ᧛✀ࡢᢏἲࡀ᭷ຠ࡛࠶ࡿ㸦㎷⏣࣭㐟ᕝ 2008㸧ࠋᅗ  ⮬⏕ᆅ᧛✀ࡢᡭ㡰ࠋ✀Ꮚࢆࢼ࢖࣓ࣟࣥࢵࢩࣗ࡞࡝࡟ධࢀࢫࣛ࢖ࢻࣇ࢕࣒ࣝࡢᯟ࡛ᅛᐃࡍࡿࠋᅵࢆ
ࡃࡎࡉ࡞࠸ࡼ࠺࡟ࢫࣜࢵࢺࢆసࡾ㸪ᆅ୰࡟࠺ࡵࡿࠋ㐺ᙜ࡞᫬ᮇ࡟᥀ࡾୖࡆほᐹࡍࡿ㸦෗┿ᕥ ๓⏣⥤Ꮚ㸪෗┿ྑ ㎷⏣᭷⣖㸧

T. Yukawa - 7

植物科学最前線 5:92 （2014）✀Ꮚࢆࢼ࢖࣓ࣟࣥࢵࢩࣗ࡞࡝࡟ᑒධࡋᆅ୰࡟࠺ࡵࡿࢩࣥࣉࣝ࡞ࡶࡢࡔࡀ㸪࠸ࡘ㸪࡝ࡇ࡛㸪࡝ࡢࡼ
࠺࡟Ⓨⱆࡍࡿ࠿ᢕᥱࡍࡿࡇ࡜ࡀ࡛ࡁ㸪ᆅୗ㒊ࡢᙧែࢆ◚ቯࡏࡎ࡟ほᐹ࡛ࡁࡿ㸦ᅗ 㸹㸳㸲❶ࡶཧ↷㸧ࠋࠕࣈࣛࢵࢡ࣎ࢵࢡࢫࠖゎ᫂ࡢษࡾᮐ࡜ࡶ࠸࠼ࡿ㸪⮬⏕ᆅ᧛✀ࢆ฼⏝ࡋࡓ◊✲ࡀ㐍ࡴࡇ࡜ࢆᮇ
ᚅࡋࡓ࠸ࠋ
ࡇࢀࡲ࡛⳦ᚑᒓᰤ㣴᳜≀ࡢ◊✲ࡣ㸪⳦᰿ඹ⏕⣔ࡢゎ᫂࡟೫ࡗ࡚࠸ࡓࡁࡽ࠸ࡀ࠶ࡿࠋ௒ᚋࡣࡉࡲ
ࡊࡲ࡞◊✲ศ㔝ࡢ▱ぢࢆ⤖㞟ࡋ㸪ከ㠃ⓗ࡞࢔ࣉ࣮ࣟࢳ࡛ࠕ᳜≀ࢆࡸࡵࡓ᳜≀ࠖࡢṇయ࡟㏕ࡾࡓ࠸
ࡶࡢࡔࠋ


ㅰ㎡
 ࡇࡇ࡛⤂௓ࡋࡓ◊✲ࡢ୍㒊ࡣ㸪᪥ᮏᏛ⾡᣺⯆఍࣭⛉Ꮫ◊✲㈝⿵ຓ㔠㸦 ࡢᨭ᥼ࢆᚓ࡚⾜ࢃࢀࡓࠋ

ᘬ⏝ᩥ⊩
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Petrosaviaceae to include Japonolirion. Brittonia 55: 214-225.
Fuse, S., & Tamura, M.N. 2000. A Phylogenetic analysis of the plastid matK gene with emphasis on
Melanthiaceae sensu lato. Plant Biol. 2: 415-427.
Merckx, V.S.F.T. (ed.) 2013. Mycoheterotrophy: The biology of plants living fungi, Springer, New York.
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.
Ogura-Tsujita, Y., Gebauer, G., Hashimoto, T., Umata, H., & Yukawa, T. 2009. Evidence for noveland specialized mycorrhizal parasitism: the orchid Gastrodia confusa gains carbon from saprotro
phic Mycena. Proc. R. Soc. B. 276: 761-767.
Ogura-Tsujita Y., Yokoyama J., Miyoshi K., & Yukawa T. 2012. Shifts in mycorrhizal fungi during
the evolution of autotrophy to mycoheterotrophy in Cymbidium (Orchidaceae). Am. J. Bot. 99:
1158-1176.
Soltis, D.E. et al. 2011. Angiosperm phylogeny: 17 genes, 640 taxa. Am. J. Bot. 98: 704-730.
ᡞ㒊༤㸪⏣ᮧᐇ 2012㸬᪂ࡋ࠸᳜≀ศ㢮Ꮫ I㸬ㅮㄯ♫㸬ᮾி㸬㎷⏣᭷⣖㸪㐟ᕝ▱ஂ 2008. ࣛࣥ⛉᳜≀ࡢ㔝እ᧛✀ヨ㦂ἲ㸬ಖ඲⏕ែᏛ◊✲ 13: 121-127.
Yagame, T., Katsuyama, T., & Yukawa, T. 2008. A New Species of Neottia (Orchidaceae) from the Tanzawa
Mountains, Japan. Acta Phytotax. Geobot. 59: 219-222.
Yamato, M., Ogura-Tsujita, Y., Takahashi, H., & Yukawa, T., in press. Significant difference in mycorrhizal
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T. Yukawa - 8

植物科学最前線 5:93 （2014）⳦ᚑᒓᰤ㣴ᛶࡢ⏕άᵝᘧࢆྍ⬟࡟ࡋࡓᵝࠎ࡞㐺ᛂ㐍໬̿≉࡟㏦⢊ᵝᘧࡢኚ໬࡟ࡘ࠸࡚
䢢䢢ᮎḟ೺ྖ࣭ຍ⸨┿ி㒔኱Ꮫ኱Ꮫ㝔ே㛫࣭⎔ቃᏛ◊✲⛉ࠛி㒔ᕷᕥி༊ྜྷ⏣஧ᮏᯇ⏫Kenji Suetsugu, Makoto Kato
Evolution of life history traits suited to mycoheterotrophic life stylewith special reference to pollination biology
Key words: breeding system, complex adaptive trait, mixotrophy, mycoheterotrophy,
Orchidaceae, reproductive biology, self-pollination
Graduate School of Human and Environmental Studies, Kyoto University
Yoshida Nihonmatsu-cho, Sakyo, Kyoto 606-8501, Japan㸯ࡣࡌࡵ࡟
ከࡃࡢ㝣ୖ᳜≀ࡣ᰿ᅪ࡟࠾࠸࡚⳦᰿⳦࡜ඹ⏕㛵ಀࢆ⤖ࢇ࡛࠾ࡾ㸪᳜≀ࡣගྜᡂ࡛ᚓࡓ
ྠ໬⏘≀ࢆ⳦᰿⳦࡟ᥦ౪ࡋ㸪ࡑࡢぢ㏉ࡾ࡟⳦᰿⳦ࡣỈࡸ❅⣲ࡸࣜࣥ࡞࡝ࡢ↓ᶵሷ㢮ࡢ྾཰ࡸ㜵⾨ࡢᡭຓࡅࢆࡋ࡚࠸ࡿࠋ
ࡋ࠿ࡋ࡞ࡀࡽ㸪᳜≀ࡢ୰࡟ࡣ㸪ගྜᡂ⬟ຊࢆኻ࠸㸪௚ࡢ⏕≀࠿ࡽࡍ࡭࡚ࡢ㣴ศࢆ␎ዣࡍࡿ࡜࠸
࠺≉␗࡞㐍໬ࢆ㐙ࡆࡓᚑᒓᰤ㣴᳜≀ࡀᏑᅾࡍࡿࠋࡇࢀࡽࡢ᳜≀ࡣ㸪௚ࡢ᳜≀࡟ྲྀࡾ௜࠸࡚㣴ศࢆዣ࠺ᐤ⏕᳜≀࡜㸦Suetsugu etal. 2008, 2012a㸧㸪⳦᰿⳦࠿ࡽ㣴ศࢆዣ࠺⳦ᚑᒓᰤ㣴᳜≀࡟኱ูࡉࢀࡿ 㸦Leake 1994, Bidartondo
2005; ᅗ 1㸧ࠋ≉࡟㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪㛤ⰼ㸪⤖ᐇᮇ௨እࡣᆅୖ࡟ጼࢆ⌧ࡉࡎ㸪᳜≀య⮬యࡀぢࡘ࠿ࡾ࡟ࡃ࠸ࡓࡵ㸪᳜≀┦ࡢ◊✲ࡀ㐍ࢇᅗ ᐤ⏕᳜≀࡜⳦ᚑᒓᰤ㣴᳜≀ ୧⪅࡜ࡶගྜᡂ⬟ࢆಖᣢࡋࡓࡶࡢ࡜᏶඲࡟႙ኻࡋࡓࡶࡢࡀ࠶ࡿࠋᐤ⏕᳜≀ࡣ྾ჾ࡛௚ࡢ᳜≀࡟ྲྀࡾ௜ࡃࡇ࡜࡛㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ⳦⣒ࢆ⣽⬊୰࡟ྲྀࡾ㎸ࡴࡇ࡜࡛㸪ᐤ⏕ࢆᡂ❧ࡉࡏࡿ 

K. Suetsugu - 1

植物科学最前線 5:94 （2014）ࡔ᪥ᮏ࡛ࡉ࠼㸪࡯࡜ࢇ࡝ࡢ✀㢮࡟࠾࠸࡚ṇ☜࡞ศᕸ᝟ሗࡣㅦࡢࡲࡲ࡛࠶ࡿ 㸦Suetsugu &Ishida 2011, Suetsugu et al. 2012b, Suetsugu 2013, Suetsugu et al. 2013 ࡞࡝㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ
⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪᪤▱ࡢ✀ᩘࡇࡑࡑࢀ࡯࡝ከࡃ࡞࠸ࡶࡢࡢ㸪ᖜᗈ࠸ศ㢮⩌࡛ぢࡽࢀ㸪⿕Ꮚ᳜≀ࡔࡅ࡛ࡶ㸪ࢶࢶࢪ⛉㸪ࣄ࣓ࣁࢠ⛉㸪ࣜࣥࢻ࢘⛉㸪ࣄࢼࣀࢩࣕࢡࢪࣙ࢘⛉㸪ࢥࣝࢩ࢔⛉㸪࢔࣓ࣖ⛉㸪ࣛࣥ⛉㸪ࢧࢡࣛ࢖ࢯ࢘⛉㸪࣍ࣥࢦ࢘ࢯ࢘⛉࡟࠾࠸࡚㸪ࡑࢀࡒࢀ⊂❧࡟㉳※ࡋ࡚࠸ࡿࡇ࡜ࡀࢃ࠿ࡗ࡚࠸ࡿ 㸦Leake 1994, Bidartondo 2005㸧ࠋࡇࢀࡲ࡛㸪⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪ወ␗࡞ᙧែ࠿ࡽேࠎࡢ㛵ᚰࢆ㞟ࡵ࡚ࡁࡓࡀ㸪Ꮫၥⓗ࡞᥈ồࡣ༑ศ࡛ࡣ࡞࠿
ࡗࡓࠋࡼࡗ࡚㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ࡝ࡢࡼ࠺࡟㉳※ࡋ㸪࡝ࡢࡼ࠺࡞㐺ᛂࢆ㐙ࡆࡓࡢ࠿ࢆ᫂
ࡽ࠿࡟ࡍࡿࡇ࡜ࡣ㸪᳜≀Ꮫ࡟࠾ࡅࡿ㔜せ࡞ㄢ㢟࡛࠶ࡿ࡜ゝ࠼ࡿࡔࢁ࠺ࠋ ㏆ᖺ㸪ศᏊ⏕≀ᏛⓗᡭἲࡸᏳᐃྠ఩యศᯒ࡜࠸ࡗࡓᡭἲࡢⓎ㐩࡟క࠸㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪ࡑࡢ㐍໬ࡢ㐣⛬࡛㸪࡝ࡢࡼ࠺࡞㐺ᛂࢆ㐙ࡆࡓࡢ࠿᫂ࡽ࠿࡟࡞ࡾࡘࡘ࠶ࡿࠋ౛࠼ࡤ㸪⥳ⴥࢆᣢࡘ✀㢮࡜᏶඲࡟ⴥ⥳⣲ࢆᣢࡓ࡞࠸✀ࢆྵࡴ㏆⦕✀ࡢ⣔⤫ᶞࢆసᡂࡍࡿࡇ࡜࡛㸪᏶඲࡟ⴥ⥳⣲ࢆኻࡗࡓ⳦ᚑᒓᰤ㣴ᛶࡢ✀ࡢฟ⌧࡟ඛ⾜ࡋ࡚㸪⥳ⴥࢆᣢࡕ࡞ࡀࡽ㸪⮬㌟ࡢ
ගྜᡂ⏘≀ࡢ୙㊊ࢆ⿵࠺㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶࡢ✀ࡀඛ࡟⌧ࢀࡿሙྜࡀ࠶ࡿࡇ࡜ࡀᣦ᦬
ࡉࢀ࡚࠸ࡿ 㸦Selosse and Roy 2009; ➨㸰❶㸪➨㸴❶ཧ↷㸧ࠋࡲࡓࡑࡢ⣔⤫ᶞୖ࡟㸪⳦࡬ࡢ౫Ꮡᗘࡸඹ⏕⳦ྀ࡜࠸ࡗࡓᙧ㉁ࢆ࣐ࢵࣆࣥࢢࡍࡿࡇ࡜࡛㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡟క࠸㸪
ࡼࡾⅣ⣲※ࢆዣ࠸ࡸࡍ࠸⳦㢮࡬ࡢᐟ୺㌿᥮ࡸ≉Ṧ໬ࡀ㉳ࡇࡿࡇ࡜ࡶ᫂ࡽ࠿࡟࡞ࡗ࡚ࡁ
ࡓ 㸦Selosse & Roy 2009㸧ࠋୖグࡢࡼ࠺࡟⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪ࡑࡢ㐍໬ࡢ㐣⛬࡛⳦࡜ࡢ㛵ࢃࡾࢆኚ໬ࡉࡏ࡚ࡁࡓࡇ
࡜ࡣ㸪㠀ᖖ࡟ࢃ࠿ࡾࡸࡍ࠸ࢫࢺ࣮࣮࡛ࣜ࠶ࡾ㸪ཷࡅධࢀࡸࡍ࠸ࡶࡢ࡛࠶ࡿࠋࡋ࠿ࡋ࡞ࡀ
ࡽ㸪ࠕ⳦ᚑᒓᰤ㣴᳜≀ࡀࡑࡢ≉␗࡞⏕άྐࢆ඲࠺ࡍࡿࡓࡵ࡟࡝ࡢࡼ࠺࡞㐺ᛂࢆ㐙ࡆࡓࡢ࠿ࠖࢆ⌮ゎࡍࡿࡓࡵ࡟ࡣ㸪ᆅୗ㒊࡜ᆅୖ㒊ࡢ୧᪉࡛㸪࡝ࡢࡼ࠺࡞㐺ᛂࡀぢࡽࢀࡿࡢ࠿ࢆ▱ࡿᚲせࡀ࠶ࡿࡣࡎ࡛࠶ࡿࠋᮏ✏࡛ࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪࡝ࡢࡼ࠺࡞㐺ᛂࢆ㐙ࡆ㸪⏕Ꮡࢆྍ⬟࡜ࡋ࡚࠸ࡿࡢ࠿ࢆ㸪ࡇࢀࡲ࡛࠶ࡲࡾὀ┠ࡉࢀ࡚࠸࡞࠿ࡗࡓᆅୖ㒊࡛ࡢ㐺ᛂࢆྵࡵ⤂௓ࡋࡓ࠸ࠋ

㸰⳦ᚑᒓᰤ㣴ᛶࡢ㐺ᛂⓗព⩏ࡲࡎ⳦㢮࠿ࡽ㣴ศࢆ␎ዣࡍࡿ࡜࠸࠺㐍໬࡟㸪࡝ࡢࡼ࠺࡞㐺ᛂⓗព⩏ࡀ࠶ࡿࡢ࠿⪃࠼࡚ࡳࡼ࠺ࠋ୍⯡ⓗ࡟ᗈࡃཷࡅධࢀࡽࢀ࡚࠸ࡿㄝ᫂ࡣ㸪⳦࡟ᐤ⏕ࡍࡿࡇ࡜࡛㸪௚ࡢ➇த┦ᡭࡢᑡ࡞࠸ᬯ࠸ᯘᗋ࡛ࡶ⏕Ꮡྍ⬟࡟࡞ࡗࡓ࡜࠸࠺ࡶࡢ࡛࠶ࡿ 㸦Bidartondo et al. 2004㸧ࠋࡋ࠿ࡋ㸪ᬯ࠸ᯘᗋ࡛ࡢ⏕⫱ࢆྍ⬟࡟ࡍࡿࡇ࡜௨እ࡟ࡶ㸪⳦࡬ࡢ౫Ꮡᛶࢆ㧗ࡵࡿࡇ࡜ࡢ㐺ᛂⓗព⩏ࡀ࠶ࡿྍ⬟ᛶࡣᤞ࡚ࡁࢀ࡞࠸ࠋ ᡃࠎࡣ㸪⳦࡟ᐤ⏕ࡍࡿࡇ࡜ࡢูࡢ㐺ᛂⓗព⩏࡜ࡋ࡚㸪୍ᗘ㸪ᆅୖ㒊ࢆ⌧ࡋࡓᚋ࡛ࡶ㸪ࢫࢺࣞࢫ࡟ࡉࡽࡉࢀࡓ㝿࡟ࡣ㸪⩣ᖺ㸪ᆅୗ࡟࡜࡝ࡲࡾ㸪⳦࠿ࡽ༑ศ࡞Ⅳ⣲※ࢆᚓࡓᚋ㸪෌ᗘ㸪ᆅୖ㒊ࢆ⌧ࡍ࡜࠸࠺ࡇ࡜ࡀ࡛ࡁࡿ࡜࠸࠺฼Ⅼࢆ⪃࠼ࡓࠋ㏻ᖖࡢගྜᡂࡔࡅࢆ⾜ࡗ࡚࠸ࡿ᳜≀࡟࠾࠸࡚ࡶ㸪ከᖺ⏕᳜≀ࡢሙྜ㸪ᆅୖ࡟⌧ࢀ࡞࠸≧ែ࡛Ꮡᅾࡋ⥆ࡅ㸪⩣ᖺ௨

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植物科学最前線 5:95 （2014）㝆㸪෌ᗘ㸪ᆅୖ㒊ࡀ⌧ࢃࢀࡿࡇ࡜ࡀ࠶ࡿࠋࡇ
࠺ࡋࡓ⌧㇟ࡣࠕᆅୖჾᐁࡢఇ╀ 㸦vegetative
dormancy ࡢヂㄒ ௨ୗࡣ㸪༢࡟ఇ╀࡜ࡍࡿ㸧ࠖ࡜࿧ࡤࢀࡿࡀ 㸦Shefferson 2009㸧㸪✀Ꮚࡢఇ╀
࡜ࡣ␗࡞ࡾ㸪᰿ࡢ࿧྾࡞࡝ࡢ௦ㅰάືࡣ೵Ṇࡋ࡞࠸ࠋࡑࡢࡓࡵࡇࢀࡲ࡛࡟ㄪᰝࡉࢀࡓ࡯࡜
ࢇ࡝ࡢ᳜≀࡛㸪⵳✚ࡋࡓ㣴ศࡀᯤῬࡋ࡚ࡋࡲ
࠸㸪ఇ╀ࢆ⤒ࡓಶయࡢ⏕Ꮡ⋡ࡀపୗࡍࡿࡇ࡜
ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ 㸦Gregg & Kéry 2006,
Hutchings 1987, Shefferson et al. 2003, Shefferson& Tali 2007 ࡞࡝㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ⳦࡟ᐤ⏕ࡍࡿ᳜≀ࡢሙྜ㸪ఇ╀୰ࡶ㸪⳦㢮࠿ࡽⅣ⣲※ࢆ⿵࠺ࡇ࡜ࡀ࡛ࡁࡿࡓࡵ㸪ఇ╀ࡢࢥࢫࢺࡀ㍍ῶࡉ
ࢀ㸦ࡴࡋࢁᆅୖ㒊ࢆ⌧ࡉ࡞࠸ࡇ࡜࡛㣗ᐖ࡞࡝ࢆཷࡅࡿ༴㝤ᛶࡀῶࡿࡓࡵ㸧㸪⏕Ꮡ⋡ࢆಖࡘࡇᅗ 㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࢟ࣥࣛࣥ

࡜ࡀ࡛ࡁࡿྍ⬟ᛶࡀ࠶ࡿࠋ ࡑࡇ࡛ᡃࠎࡣ㸪Mark and recapture ἲࢆ⏝࠸㸪㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶࡢ࢟ࣥࣛࣥࡢఇ╀⋡ࡸ⏕Ꮡ⋡ࡀ࡝ࡢ⛬ᗘ࠿㸪ࡉࡽ࡟㣗ᐖࡸ⤖ᐇࡢ᭷↓࡞࡝࡟ࡼࡾ㸪ఇ╀⋡ࡸ⏕Ꮡ⋡ࡀ࡝ࡢࡼ࠺࡟ኚ໬ࡍࡿࡢ࠿ࢆ᳨ウࡋࡓࠋ࢟ࣥࣛࣥࡣ᱂ᇵࡀ㞴ࡋࡃ㸪㛗ᮇ㛫⥔ᣢࡍࡿࡇ࡜ࡣ㞴ࡋ࠸ࡓࡵ㸪⥳ⴥࢆᒎ㛤ࡋ࡚࠿ࡽࡶ⳦࡟౫Ꮡࡋ࡚࠸ࡿྍ⬟ᛶࡀ♧၀ࡉࢀ࡚࠸ࡓࡀ㸪㏆ᖺ⾜ࢃࢀࡓ᱂ᇵᐇ㦂ࡸᏳᐃྠ఩యศᯒ࠿ࡽ㸪ᐇ㝿࡟⳦࡟౫Ꮡࡋ࡚࠸ࡿࡇ࡜ࡀド᫂ࡉࢀ࡚࠸ࡿ᳜≀࡛࠶ࡿ 㸦ᅗ 2; Yagame & Yamato 2013, ᆏᮏࡽ 2013㸧ࠋࡑࡢ⤖ᯝ㸪ᡃࠎࡢண᝿㏻ࡾ㸪㣗ᐖࢆཷࡅࡓ࡜ࡁࡸ⤖ᐇࡋࡓ࡜ࡁ࡞࡝㸪᳜≀ࡢ㈨※ࡀᯤῬࡋࡓ㝿࡟ࡣ㸪ఇ╀ࡀ㉳ࡇࡾࡸࡍࡃ࡞ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋࡲࡓࡇࡢࡼ࠺࡟⏕Ꮡࡀ㞴ࡋࡃ࡞ࡗࡓ≧ែ࡛ఇ╀࡟ධࡿ࡟ࡶ㛵ࢃࡽࡎ㸪ఇ╀ࢆ⤒ࡓಶయࡢ⏕Ꮡ⋡ࡀపୗࡍࡿࡇ࡜ࡣ࡞࠿ࡗࡓ 㸦ᮎḟࡽ ᢞ✏୰㸧ࠋᐇࡣ㸪ඛ⾜◊✲࡛ࡶ㸪౛እⓗ࡟ఇ╀ᚋࡢ⏕Ꮡ⋡ࡀୗࡀࡽ࡞࠸✀㢮ࡀ▱ࡽࢀ࡚࠸ࡿࠋࡇ
ࢀࡽࡣ㸪⳦ᚑᒓᰤ㣴ᛶࡢほⅬ࠿ࡽࡣ㆟ㄽࡉࢀ࡚࠸࡞࠿ࡗࡓࡀ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶࡢ✀㢮࡛」ᩘᅇ㸪ሗ࿌ࡉࢀ࡚࠸ࡿ⌧㇟࡛࠶ࡿ 㸦Shefferson et al. 2005, Jäkäläniemi et al. 2011㸧ࠋୖグࡢࡇ࡜ࢆ⪃࠼ేࡏࡿ࡜㸪ࠕࢫࢺࣞࢫࢆཷࡅࡓ㝿࡟㸪ᆅୖ㒊ࢆᒎ㛤ࡏࡎ࡟ᆅୗ࡟࡜࡝
ࡲࡿࡇ࡜ࡀ࡛ࡁࡿࠖࡇ࡜ࡶ㸪⳦࡟ᐤ⏕ࡍࡿព⩏ࡢ୍ࡘ࡛࠶ࡿ࡜⪃࠼ࡿࡇ࡜ࡀ࡛ࡁࡿࠋ
䢢㸱࢔ࣝࣅࣀ✺↛ኚ␗య࠿ࡽぢ࠼ࡿ↓ⴥ⥳໬ࡢ㞀ቨ
ඛ࡟㏙࡭ࡓࡼ࠺࡟㸪ࣛࣥ⛉ࡸࢶࢶࢪ⛉࡞࡝ࡢ୍㒊ࡢ✀㢮ࡣ㸪Ⓨ㐩ࡋࡓ⥳ⴥࢆᒎ㛤ࡋ㸪୍ぢࡍࡿ࡜ගྜᡂࡔࡅ࡛⏕Ꮡྍ⬟ࡢࡼ࠺࡟ぢ࠼ࡿ࡟ࡶ࠿࠿ࢃࡽࡎ㸪⳦㢮࡟ࡶⅣ⣲ࢆ୍㒊౫Ꮡࡋ࡚࠸ࡿ✀㢮ࡀᏑᅾࡍࡿ 㸦Gebauer & Meyer 2003, Bidartondo et al. 2004, Tedersoo et
al. 2007 ࡞࡝㸧ࠋࡇࡢࡼ࠺࡞㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࢆྵࡴࢢ࣮ࣝࣉ࠿ࡽ㸪ⴥ⥳⣲ࢆኻࡗࡓ

K. Suetsugu - 3

植物科学最前線 5:96 （2014）᏶඲࡟⳦ᚑᒓᰤ㣴ᛶ࡟≉໬ࡋࡓ✀㢮ࡀฟ⌧ࡍࡿࡇ࡜ࡀ࠶ࡿࡀ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴ࡢ✀ᩘ࡟ẚ࡭࡚㸪᏶඲࡟⳦ᚑᒓᰤ㣴ᛶࢆ⋓ᚓࡋࡓ✀ᩘࡣࡁࢃࡵ࡚ᑡ࡞࠸ࠋ๓㡯࡛㏙࡭ࡓࡼ࠺࡟
⳦࡟ᐤ⏕ࡍࡿ࡜࠸࠺⏕άྐࡀ᳜≀࡟࡜ࡗ࡚ᚓ⟇࡛࠶ࡿ࡞ࡽࡤ㸪⳦࠿ࡽࡢ㣴ศ␎ዣࡔࡅ࡛
⏕⫱ࡍࡿ↓ⴥ⥳᳜≀ࡣఱᨾᑡ࡞࠸ࡢࡔࢁ࠺࠿ࠋࡇࡢࡇ࡜ࢆ⪃࠼ࡿࡓࡵ࡟㸪↓ⴥ⥳໬ࡢ⋓ᚓ࡟࡝ࡢࡼ࠺࡞㞀ቨࡀ࠶ࡿࡢ࠿ࢆ⪃࠼࡚ࡳࡼ࠺ࠋ ⯆࿡῝࠸ࡇ࡜࡟㸪㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪᏶඲࡟ⴥ⥳⣲ࢆኻࡗࡓ࢔ࣝࣅࣀ✺↛ኚ␗యࢆ⏕ࡌࡿࡇ࡜ࡀከ࠸ 㸦Selosse et al. 2004, Julou etal. 2005, Tranchida-Lombardo et al. 2010 ࡞࡝㸧ࠋ
㏻ᖖࡢ᳜≀࡛࠶ࢀࡤ㸪ⴥ⥳⣲ࢆኻࡗࡓ࢔ࣝࣅ
ࣀࡣ㸪✀Ꮚ࡟㈓ⶶࡉࢀࡓ㣴ศࢆ౑࠸ᯝࡓࡍ࡜
ᯤࢀ࡚ࡋࡲ࠺ࠋࡋ࠿ࡋ࡞ࡀࡽ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࢔ࣝࣅࣀࡣ㸪⳦࠿ࡽⅣ⣲※ࢆྵࡴ㣴ศࢆ␎ዣࡍࡿࡇ࡜ࡀ࡛ࡁࡿࡓࡵ㸪ⴥ⥳⣲ࢆᣢ
ࡘ㏻ᖖಶయ࡜ྠ⛬ᗘࡲ࡛ᡂ㛗ࡋ㸪ⰼࢆဏ࠿ࡏ
ࡿࡇ࡜ࡀ࡛ࡁࡿࡢ࡛࠶ࡿࠋࡇࡢࡇ࡜ࡣ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪⳦࠿ࡽ␎ዣࡋࡓ㣴ศࡔࡅ࡛⏕Ꮡࡍࡿࡇ࡜ࡀྍ⬟࡞ࡇ࡜ࢆ♧၀ࡋ࡚࠸ࡿࠋ
࡟ࡶ࠿࠿ࢃࡽࡎ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࢔ࣝࣅࣀࡣ⛥࡞Ꮡᅾ࡛࠶ࡾ㸪㞟ᅋෆ࡟ᗈࡀࡿࡇ
࡜ࡣࡵࡗࡓ࡟࡞࠸ࡢ࡛࠶ࡿࠋᅗ ࢟ࣥࣛࣥࡢ࢔ࣝࣅࣀ✺↛ኚ␗య⳦࠿ࡽ㣴ศࢆ␎ዣࡍࡿ⬟ຊࢆ⋓ᚓࡋࡓࢢ࣮ࣝࣉ
ࡢ୰࡛㸪᏶඲࡟↓ⴥ⥳࡟࡞ࡗࡓ✀㢮ࡀᑡ࡞࠸ࡇ࡜ࡸ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࢔ࣝࣅࣀ
ࡀ㸪㞟ᅋෆ࡟ᗈࡀࡽ࡞࠸ࡇ࡜ࡣ㸪⳦࡟ᑐࡍࡿᐤ⏕⬟ຊࡢ⋓ᚓ௨እࡢ㠃࡛ࡶ㸪↓ⴥ⥳໬ࢆ㐩ᡂࡍࡿࡓࡵ࡟ࡣࡉࡽ࡞ࡿ㐺ᛂࡀᚲせ࡛࠶ࡿࡇ࡜ࢆ♧၀ࡋ࡚࠸ࡿࠋ ࢟ࣥࣛࣥࡶ௚ࡢ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ౛࡟ࡶࢀࡎ㸪ࡋࡤࡋࡤ࢔ࣝࣅࣀࢆ⏕ࡌࡿಶయ⩌ࡀぢཷࡅࡽࢀࡿ 㸦ᅗ 3㸧ࠋࡇࡢࡼ࠺࡞࢔ࣝࣅࣀࡀఱᨾ㞟ᅋ୰࡟ᗈࡀࡗ࡚࠸࠿࡞࠸ࡢ࠿
㸦ࡘࡲࡾ㸪↓ⴥ⥳ࡢ㐍໬࡟ࡣ࡝ࡢࡼ࠺࡞㞀ቨࡀ࠶ࡿࡢ࠿㸧 ࢆ㸪࢟ࣥࣛࣥࡢ࢔ࣝࣅࣀ࡜
㏻ᖖಶయࡢ✀Ꮚ⏕⏘ຊࢆ㐺ᛂᗘࡢᣦᶆ࡜ࡍࡿࡇ࡜࡛⪃࠼࡚ࡳࡓ࠸ࠋ࢟ࣥࣛࣥࡣ㸪㏦⢊⪅ࡀ࠸࡞ࡅࢀࡤᯝᐇࢆࡘࡅࡿࡇ࡜ࡀฟ᮶࡞࠸᳜≀࡛㸪ࣄ࣓ࣁࢼࣂࢳࡸࢥࣁࢼࣂࢳ࡜࠸ࡗࡓ
ᑠᆺࡢࣁࢳ࡟ⰼ⢊ࡢ፹௓ࢆクࡋ࡚࠸ࡿ 㸦ᮎḟࡽ ᢞ✏୰㸧ࠋࡼࡗ࡚㸪࢔ࣝࣅࣀࡢ࡯࠺ࡀ
㏦⢊⪅ࡢㄏᘬຊࡀஈࡋࡅࢀࡤ㸪ᯝᐇࢆࡘࡅࡿᩘ㸪ࡦ࠸࡚ࡣ✀Ꮚᩘ࡟㐪࠸ࡀฟࡿ࡜⪃࠼ࡽ
ࢀࡿࠋ㏦⢊⪅ࡢゼࢀࡓᅇᩘࢆṇ☜࡟ᢕᥱࡍࡿࡇ࡜ࡣ㸪㏻ᖖࡢ᳜≀࡟࠾࠸࡚ࡣ㛤ⰼᮇ㛫୰⥅⥆ࡋ࡚ほᐹࡋ࡞࠸㝈ࡾᴟࡵ࡚㞴ࡋ࠸ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ࣖࢡࢩ࣐ࣛࣥள⛉ࢆ㝖ࡃࣛࣥ⛉࡛ࡣ㸪ほᐹࡀᐜ࡛᫆࠶ࡿ 㸦Suetsugu & Fukushima 2013, Suetsugu & Tanaka 2013a, b㸧ࠋఱᨾ࡞ࡽࡤ㸪ᩘ༑୓ࡢⰼ⢊ࡀ஫࠸࡟⤖ྜࡋ࡚ሢ࡟࡞ࡗࡓⰼ⢊ሢ࡜࿧ࡤࢀࡿᵓ㐀࡜࡞ࡗ࡚࠾

K. Suetsugu - 4

植物科学最前線 5:97 （2014）ࡾ㸪ࡑࡢࡓࡵ඲࡚ࡢⰼ⢊ࡣ㸪᪻⹸ࡀ 1 ᅇ㸦ከࡃ࡚ࡶᩘᅇ㸧ゼࢀࡓࡔࡅ࡛ᣢࡕཤࡽࢀࡿ࠿
ࡽ࡛࠶ࡿࠋࡑࡇ࡛ᐇ㝿࡟ⰼ⢊ࡢᣢࡕཤࡾ⋡ࡸཷ⢊⋡ࢆㄪ࡭ࡓ࡜ࡇࢁ㸪࢔ࣝࣅࣀಶయࡶ㏻
ᖖಶయ࡜ྠ⛬ᗘ㸪㏦⢊⪅ࢆ࠾ࡧࡁᐤࡏࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ 㸦ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡘࡲࡾ㸪࢟ࣥࣛࣥࡢ࢔ࣝࣅࣀಶయࡢ㐺ᛂᗘࡀୗࡀࡿ⌮⏤ࡣ㸪㏦⢊⪅ࡢㄏᘬຊࡢ
ᕪ࡛ࡣ࡞࠸ࡢ࡛࠶ࡿࠋࡇࡢᚋࡢ㡯┠࡛ࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪ཷ⢊⏕ែࡢ㠃࡛ࡶไ⣙ࢆཷࡅ࡚࠸ࡿࡇ࡜ࢆ㏙࡭ࡿࡀ㸪ࡇࢀ࡟ࡣᬯᡤ࡬ࡢ㐍ฟࡀ῝ࡃ㛵ࢃࡗ࡚࠸ࡿࠋࡋ࠿ࡋ࡞ࡀࡽ
࢟ࣥࣛࣥࡢ࢔ࣝࣅࣀࡣ㸪㏻ᖖಶయ࡜ྠࡌࡼ࠺࡞ග⎔ቃ࡟⏕⫱ࡋ࡚࠸ࡿࡓࡵ㸪ཷ⢊ࡢ㠃࡛㸪ⴭࡋ࠸୙฼┈ࢆ⿕ࡿࡇ࡜ࡣ࡞࠸ࡢ࡛࠶ࡿࠋ㐺ᛂᗘࡀୗࡀࡿ⌮⏤ࡀ㏦⢊⪅࡜ࡣ↓㛵ಀ࡛࠶ࡿࡇ࡜ࡣ㸪⮬ື⮬ᐙཷ⢊ࡀྍ⬟࡞ࢠࣥࣛࣥࡸࣁ࣐࡛࢝࢟ࣛࣥࡶ࢔ࣝࣅࣀಶయࡶ㞟ᅋෆ࡟ᗈࡀࡗ࡚࠸࡞࠸ࡇ࡜࠿ࡽࡶ♧၀ࡉࢀࡿ 㸦ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋ ࡑࢀ࡛ࡣఱᨾ࢔ࣝࣅࣀࡣ㞟ᅋෆ࡟ᗈࡀࡽ࡞࠸ࡢ࡛࠶ࢁ࠺࠿ࠋ࢔ࣝࣅࣀ࡜㏻ᖖಶయࡢ㑇ఏⓗ࡞Ỵᐃ࣓࢝ࢽࢬ࣒ࡣ᫂ࡽ࠿࡛ࡣ࡞࠸ࡶࡢࡢ㸪࢔ࣝࣅࣀࡀຎໃ࡛࣍ࣔ࠶ࡿྍ⬟ᛶࡣ༑ศ⪃࠼ࡽࢀ㸪ࡇࡢࡇ࡜ࡀ୍ᅉ࡜࡞ࡗ࡚࠸ࡿࡢ࠿ࡶࡋࢀ࡞࠸ࠋࡋ࠿ࡋ࡞ࡀࡽほᐹࢆ⥆ࡅ࡚࠸ࡿ࡜㸪ࡑࡢ௚ࡢ⌮⏤ࡢᏑᅾࡶ♧၀ࡉࢀࡓࠋ୰࡛ࡶ኱ࡁ࡞⌮⏤࡜ࡋ࡚࢔ࣝࣅࣀࡣ㏻ᖖಶ
యࡼࡾࡶ᪩࠸ࢱ࢖࣑ࣥࢢ 㸦ከࡃࡢሙྜ✀Ꮚࡀᡂ⇍ࡍࡿ๓㸧 ࡟ᾘኻࡋ࡚ࡋࡲ࠺ࡓࡵ㸪㏻
ᖖಶయ࡟ẚ࡭ 1%௨ୗࡢ✀Ꮚࡋ࠿ṧࡍࡇ࡜ࡀ࡛ࡁ࡞࠸ࡇ࡜ࡀᣲࡆࡽࢀࡿࠋ⤖ᐇ๓࡟ᯤࢀ
࡚ࡋࡲ࠺ࡢࡣ㸪ගྜᡂࢆ⾜ࢃ࡞࠸࡟ࡶ㛵ࢃࡽࡎ኱ࡁ࡞ⴥࡸẼᏍࢆಖᣢࡋ࡚࠸ࡿࡓࡵ㸪⵨ᩓࡀ࠺ࡲࡃ࠸࠿ࡎ஝⇱ࡋࡸࡍࡃ㏻ᖖಶయࡼࡾ᪩ࡃᯤࢀ࡚ࡋࡲ࠺ࡇ࡜ࡀ୍ᅉࡔ࡜⪃࠼ࡽࢀࡿ 㸦Roy et al. 2013, ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡲࡓ࢔ࣝࣅࣀࡣⴥ⥳⣲ࢆᣢࡘ㏻ᖖ
ಶయ࡜ྠ⛬ᗘࡲ࡛ᡂ㛗ࡍࡿࡶࡢࡢ㸪ගྜᡂ⬟ࢆኻࡗ࡚࠸ࡿศࡔࡅⅣ⣲ไ㝈࡟㝗ࡾࡸࡍ࠸ྍ⬟ᛶࡶ࠶ࡿࠋࡑࡢሙྜ㸪ྠ໬ჾᐁࡢ㏥໬ࡣ㸪⵨ᩓ㐣ከ࡟࡞ࡿࡢࢆ㜵ࡄࡔࡅ࡛࡞ࡃ㸪㈨※๐ῶࡢព࿡࡛ࡶ኱ࡁ࡞ព࿡ࢆᣢࡘ࡜⪃࠼ࡽࢀࡿࠋ௚࡟ࡶ㸪㸦1㸧 ࢔ࣝࣅࣀಶయࡣ㏻ᖖ
ಶయࡼࡾࡶ㸪⳦࡬ࡢ౫Ꮡᗘࡀ㧗࠸ࡓࡵ❅⣲ྵ᭷⋡ࡀ㧗ࡃ㸪᳜㣗ᛶ᪻⹸࡟࡜ࡗ࡚ዲ㐺࡞㣵࡟࡞ࡿࡇ࡜ࡸ㸪㸦2㸧 ཯ᑕ⋡ࡢ㧗࠸ⓑ࠸ⴥࢆᣢࡘࡇ࡜࡛㸪᳜㣗ᛶ᪻⹸࠿ࡽぢࡘࡅࡽࢀࡸ
ࡍࡃ࡞ࡿࡇ࡜࡟ࡼࡿ㣗ᐖ⋡ࡢୖ᪼࡞࡝ࡶ࢔ࣝࣅࣀࡢ⏕⫱࡟ࡣ୙฼࡟ാ࠸࡚࠸ࡿ
㸦Stöckel et al. 2011, Roy et al. 2013, ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡘࡲࡾ↓ⴥ⥳᳜≀࡟࡞ࡿࡓࡵ࡟ࡣ㸪⳦࠿ࡽࡢ㣴ศ␎ዣࢆྍ⬟࡟ࡍࡿ⬟ຊࡢࡳ࡞ࡽࡎ㸪ẼᏍࡢ㏥໬㸪ⴥ࡟௦⾲ࡉࢀࡿྠ໬ჾᐁࡢ㏥໬㸪㛤ⰼ࠿ࡽ⤖ᐇࡲ࡛ࡢᮇ㛫ࡢ▷⦰㸪㠀㛤ⰼ᫬ᮇࡢఇ╀㸪ಖㆤⰍ㸪⦾
Ṫᵝᘧࡢኚ໬ 㸦⦾Ṫᵝᘧࡢኚ໬࡟ࡘ࠸࡚ࡣḟ㡯࡛ヲࡋࡃㄝ᫂ࡍࡿ㸧 ࡞࡝ࡢᵝࠎ࡞㐺ᛂࢆ㐙ࡆࡿᚲせࡀ࠶ࡿ࡜⪃࠼ࡽࢀࡿࠋࡇࢀࡽࡢ㐺ᛂࢆྠ᫬࡟㐙ࡆࡿࡇ࡜ࡀ㞴ࡋ࠸ࡓࡵ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀࡟ẚ࡭㸪ⴥ⥳⣲ࢆኻࡗࡓ⳦ᚑᒓᰤ㣴᳜≀ࡣᑡ࡞࠸ࡢ࠿ࡶࡋࢀ࡞࠸ࠋ

㸲⳦ᚑᒓᰤ㣴᳜≀࡛༟㉺ࡍࡿ⮬ᐙཷ⢊๓㡯࡛ࡣ㸪࢔ࣝࣅࣀࢆ⏝࠸࡚㸪ᙜ௦ࡢ⏕⫱࡟㛵ࡋ࡚㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡟࡝ࡢࡼ࠺
࡞㞀ቨࡀ࠶ࡿࡢ࠿ࢆ⪃࠼ࡓࠋḟ࡟ࠕ✀ࠖ࡜ࡋ࡚ᚑᒓᰤ㣴ᛶࢆ⋓ᚓࡋ࡚࠸ࡿࡶࡢࢆ⏝࠸࡚㸪⦾Ṫ࡟㛵ࡋ࡚㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ⋓ᚓ࡟㸪࡝ࡢࡼ࠺࡞ไ⣙ࡀ࠶ࡿࡢ࠿ࢆ⪃࠼ࡓ࠸ࠋ⳦ᚑ

K. Suetsugu - 5

植物科学最前線 5:98 （2014）ᒓᰤ㣴ᛶࡢ㐍໬ࡢ฼Ⅼ࡜ࡋ࡚ࡼࡃᣲࡆࡽࢀࡿࡶࡢࡣ㸪ගྜᡂࢆࡍࡿᚲせࡀ࡞࠸ࡓࡵ㸪௚ࡢ➇த┦ᡭࡢᑡ࡞࠸ᬯ࠸ᯘᗋ࡛⏕Ꮡྍ⬟࡟࡞ࡿ࡜࠸࠺ࡶࡢ࡛࠶ࡿࠋࡋ࠿ࡋ࡞ࡀࡽᬯ࠸⎔ቃ࡛⏕άࡍࡿࡇ࡜ࡣ㸪⳦࡜ࡢ㛵ࢃࡾ௨እࡢ㠃࡛ไ⣙ࢆཷࡅࡿࡇ࡜ࡣ࡞࠸ࡢࡔࢁ࠺࠿ࠋ౛
࠼ࡤ㸪ᬯ࠸ᯘᗋ࡛ࡢ⏕άࢆᡂࡋ㐙ࡆࡿࡓࡵ࡟ࡣ㸪⦾Ṫࡶᬯ࠸⎔ቃ࡛⾜࠺ᚲせࡀ࠶ࡿࡀ㸪ࡇ࠺ࡋࡓ⎔ቃࡣࣁࢼࣂࢳ࡞࡝ࡢゼⰼᛶ᪻⹸ࡢ࡟ࡂࢃ࠸࡜ࡣ↓⦕ࡢୡ⏺࡛࠶ࡿ 㸦Herrera
1995, Herrera 1997, Lee et al. 2001㸧ࠋࡑࡢࡓࡵ㸪⳦࡟ᐤ⏕ࡍࡿ࡜࠸࠺⏕άྐࡣ㸪ⷧᬯ࠸ᯘᗋ࡛ཷ⢊ࢆ㐩ᡂࡍࡿ࡜࠸࠺ᅔ㞴ࢆ᳜≀࡟ᙉ࠸࡚࠸ࡿࠋࡑࡇ࡛ࡇࡢࡼ࠺࡞ᬯ࠸ሙᡤ࡟⏕⫱ࡍࡿ᳜≀ࡣ㸪࡝ࡢࡼ࠺࡞᪉ἲ࡛ཷ⢊ࢆ㐩ᡂࡋ࡚࠸ࡿࡢ࠿⤂௓ࡋ࡚࠸ࡃࡇ࡜࡟ࡋࡼ࠺ࠋ ⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡜ཷ⢊ᵝᘧࡢ㛫࡟ఱࡽ࠿ࡢ㛵㐃ࡀ࠶ࡿࡢ࡛ࡣ࡞࠸࠿࡜࠸࠺௬ㄝࡣ௨
๓࠿ࡽᏑᅾࡋ࡚࠾ࡾ㸪Bidartondo 㸦2005㸧 ࡣ㸪㐣ཤࡢ⳦ᚑᒓᰤ㣴᳜≀ࡢ㏦⢊ᵝᘧ࡟㛵ࡍࡿሗ࿌ࢆࡶ࡜࡟㸪
ࠕᆅୗ㒊࡛⳦᰿⳦ࢆ㦄ࡋ࡚࠸ࡿ⳦
ᚑᒓᰤ㣴᳜≀ࡣ㸪ᆅୖ㒊࡛⻤ࢆࡔࡉࡎ࡟㏦⢊⪅ࢆ㦄ࡍࡇ࡜ࡣ࡛ࡁࡎ㸪⮬Ṫࢆ᥇⏝ࡋ࡚࠸ࡿࠖ
࡜࠸࠺௬ㄝࢆᥦၐࡋࡓࠋࡋ࠿ࡋ࡞ࡀࡽ⳦ᚑᒓ
ᰤ㣴᳜≀ࡢ㏦⢊ᵝᘧ࡟㛵ࡍࡿ◊✲ࡣ㸪ᾮᾐᶆᮏࢆ⏝࠸࡚㸪ᰕ㢌࡜ⵂࡢ఩⨨㛵ಀ࡞࡝࠿ࡽཷ
⢊ᵝᘧࢆ᥎ᐃࡋࡓ◊✲ࡀ࡯࡜ࢇ࡝࡛
㸦Warming 1901, Oehler 1927㸧㸪ᐇ㝿࡟㔝እほᐹ㸪⿄᥃ࡅฎ⌮࡞࡝ࢆ㏻ࡋ࡚ヲ⣽࡟ཷ⢊ᵝᘧࢆ᳨ウࡋࡓ◊✲ࡣഹ࠿࡛࠶ࡗࡓࠋࡑࡇ࡛ᡃࠎࡣ㸪ࠕ⳦ᚑᒓᰤ㣴ᛶ࡜㏦⢊ᵝᘧࡢኚ໬࡟ࡣᐦ᥋࡞
㛵ࢃࡾࡀ࠶ࡿࠖ࡜࠸࠺௬ㄝࡢୗ㸪᪥ᮏ࡟⏘ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࡢཷ⢊ᵝᘧࢆヲ⣽࡟᳨ウࡋᅗ ⤖ᐇᮇࡢࢶࢳ࢔ࢣࣅⷣࡢ᫬ᮇ࡟⿄᥃ࡅࢆ⾜ࡗࡓಶయ࡜⮬↛≧ែࡢಶయ ྠ⛬ᗘࡢ⤖ᐇࡀぢࡽࢀࡿ ࡓࠋࡑࡢ⤖ᯝ㸪ࢩࣗࣥࣛࣥᒓ㸪࣒ࣚ࢘ࣛࣥᒓ㸪ࢱࢿ࢞ࢩ࣐࣒ࣚ࢘ࣛࣥᒓ㸪࢖ࣔࣛࣥᒓ㸪ࣄ࣓ࣀࣖ࢞ࣛᒓ㸪ࣄࢼࣀࢩࣕࢡࢪࣙ࢘ᒓ㸪࢜ࢽࣀࣖ࢞ࣛᒓ㸪ࢶࢳ࢔ࢣࣅᒓ㸪ࢺࣛ࢟ࢳࣛ
ࣥᒓ࡞࡝ࡢ⳦ᚑᒓᰤ㣴᳜≀ࡢከࡃࡀ㸪⮬ື⮬ᐙཷ⢊ࢆ㐍໬ࡉࡏ࡚࠸ࡓࡇ࡜ࡀ᫂ࡽ࠿࡟࡞
ࡗࡓࠋ ౛࠼ࡤ㸪↓ⴥ⥳ࡢࢶࢳ࢔ࢣࣅࡣ㸪ࣂࢽࣛள⛉㸪ࣂࢽࣛ㐃࡟ᒓࡋ㸪ගྜᡂࢆࡍࡿࣂࢽࣛ
࡜㏆⦕࡛࠶ࡿ 㸦Cameron & Carmen Molina 2006㸧ࠋࣂࢽࣛᒓࡢ᳜≀ࡣ㸪ᑠᆺࡢࣁࢼࣂࢳ
࡟ࡼࡗ࡚㏦⢊ࡀ⾜ࢃࢀࡿࡶࡢࡀ࡯࡜ࢇ࡝࡛㸪᪻⹸ࡢຓࡅ࡞ࡋ࡛ཷ⢊ࡀ㉳ࡇࡿ✀㢮ࡣࡈࡃࢃࡎ࠿࡛࠶ࡿ 㸦Soto Arenas 2003㸧
ࠋࢶࢳ࢔ࢣࣅࡶⰼᙧែࡣ㸪ࣂࢽࣛࡢࡑࢀ࡜㠀ᖖ࡟ࡼࡃఝ࡚࠾ࡾ㸪ࢥࣁࢼࣂࢳ࡞࡝ࡢᑠᆺࡢࣁࢼࣂࢳ࡟ࡼࡗ࡚㏦⢊ࡉࢀࡿࡇ࡜ࡀண᝿ࡉࢀࡿࠋࡋ
࠿ࡋ࡞ࡀࡽὴᡭ࡞ⰼࢆࡋ࡚࠸ࡿ࡟ࡶ࠿࠿ࢃࡽࡎ㸪ࢶࢳ࢔ࢣࣅ࡟ࡘ࠸࡚ࡣ㸪࠿࡞ࡾ᪩࠸ࢱ
࢖࣑ࣥࢢ࡛ 㸦ሙྜ࡟ࡼࡗ࡚ࡣ㛤ⰼࡍࡿ๓࡟㸧㸪ⵂᖗ࡟᱁⣡ࡉࢀࡓⰼ⢊ሢࡀࡍࡄୗࡢᰕ㢌

K. Suetsugu - 6

植物科学最前線 5:99 （2014）࡟⁥ࡾⴠࡕ㸪⮬ື⮬ᐙཷ⢊ࡀ㉳ࡇࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ 㸦Suetsugu 2013㸧ࠋ᪩࠸ࢱ࢖࣑ࣥࢢ࡛⮬ື⮬ᐙཷ⢊ࡀ㉳ࡇࡿࡇ࡜ࢆ⪃࠼ࡿ࡜㸪ࢶࢳ࢔ࢣࣅࡣ୺࡟⮬Ṫ࡛ ✀Ꮚࢆṧࡋ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿ 㸦ᅗ 4㸧ࠋࣂࢽࣛᒓ࡛ࡣ⮬ᐙཷ⢊ࡢ✀㢮ࡀᴟࡵ࡚ᑡ࡞࠸ࡇ࡜ࢆ⪃࠼ࡿ࡜㸪ࡇࡢ஦ࡣ㸪⳦ᚑᒓᰤ㣴ᛶ࡜⮬Ṫࡢ㐍໬࡟㛵㐃ࡀ࠶ࡿྍ⬟ᛶࢆ♧၀ࡋ࡚࠸ࡿࠋ ࡋ࠿ࡋ࡞ࡀࡽ⳦ᚑᒓᰤ㣴ᛶ࡜⮬Ṫࡢ㐍໬࡟㛵㐃ᛶࡀ࠶ࡿ࠿ࢆ᫂ࡽ࠿࡟ࡍࡿࡓࡵ࡟ࡣ㸪ヲ⣽࡞⣔⤫㛵ಀࡀ᫂ࡽ࠿࡜࡞ࡗ࡚࠸ࡿ✀⩌ࢆ⏝࠸ࡿ࡯࠺ࡀ㐺ษ࡛࠶ࡿࠋࡇࢀࡲ࡛ࡢ◊✲࠿ࡽ㸪ࢩࣗࣥࣛࣥᒓ࡛ࡣ㸪↓ⴥ⥳ᛶࡀ 1 ᅇ㐍໬ࡋࡓࡇ࡜㸪↓ⴥ⥳ᛶࡢ࣐ࣖࣛࣥ࡜ࢧ࣑࢞
ࣛࣥࡣ༢⣔⤫⩌࡜࡞ࡿࡇ࡜㸪୧✀ࡢጜጒ⩌࡟ࡣᬑ㏻ⴥࢆ⏕ࡌࡿࢼࢠࣛࣥࡀ఩⨨ࡍࡿࡇ࡜㸪ࡲࡓࡇࢀࡽࡢእ⩌ࡀࢩ࡛ࣗࣥࣛࣥ࠶ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ 㸦Yukawa et al. 2002㸧
ࠋࡲࡓᏳᐃྠ఩యศᯒ࡟ࡼࡗ࡚㸪ᬑ㏻ⴥࢆᒎ㛤ࡍࡿࢩࣗࣥࣛࣥ㸪ࢼࢠࣛࣥࡶ㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶ࡛࠶ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ 㸦Motomura et al. 2010㸧ࠋࡼࡗ࡚ࢩࣗࣥࣛࣥᒓ
ࡣ㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡜᳜≀ࡢᙧ㉁㐍໬ࢆ᳨ウࡍࡿࡢ࡟ࡍࡄࢀࡓ⣔࡛࠶ࡿ࡜࠸࠼ࡿࠋ
ࡇࡢࢩࣗࣥࣛࣥᒓ࡟࠾࠸࡚ࡶ㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡜ཷ⢊ᵝᘧࡢ㛫࡟ࡣ㸪┦㛵ࡀぢࡽࢀࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋࢼࢠࣛࣥࡢእ⩌࡜࡞ࡿࢩࣗࣥࣛࣥ࡜࡛࢝ࣥࣛࣥࡣ㸪࣑ࢶࣂࢳࡢ࣮࣮࣡࢝࡜ࢣࣈ࢝ࣁࢼࣂࢳࡢ㞝࡟ࡑࢀࡒࢀ㏦⢊ࡉࢀ㸪⮬ື⮬ᐙཷ⢊ࡶ⾜ࢃ࡞࠸ 㸦Tsuji& Kato 2010㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ࡼࡾ⳦ᚑᒓᰤ㣴ᛶࢆᙉࡵ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿࢼࢠࣛࣥࡸ
ⴥࢆ⏕ࡌ࡞࠸࣐ࣖࣛࣥࡸࢧ࣑࡛࢞ࣛࣥࡣ㸪୺࡟⮬ື⮬ᐙཷ⢊ࢆ⾜ࡗ࡚࠸ࡿࠋ⯆࿡῝࠸ࡇ࡜࡟㸪୰ᅜࡢࢼࢠࣛࣥࡢಶయ⩌࡛ࡣ㸪⮬ື⮬ᐙཷ⢊ࢆ⾜ࢃ࡞࠸ಶయ⩌ࡢᏑᅾࡶ☜ㄆࡉࢀ࡚࠸ࡿ 㸦Cheng et al. 2007㸧ࠋࢩࣗࣥࣛࣥᒓ࡛ࡣ㸪↓ⴥ⥳ࡢ㐍໬࡟కࡗ࡚⭉⏕⳦࠿ࡽእ⏕⳦᰿⳦࡬࡜ࣃ࣮ࢺࢼ࣮ࡢࢩࣇࢺࡀ㉳ࡇࡗ࡚࠾ࡾ㸪ࡑࡢ㐣⛬࡛୧⪅ࢆಖᣢࡍࡿ୰㛫ẁ㝵ࢆ⤒ࡿࡇ࡜ࡀศ࠿ࡗ࡚࠸ࡿࡀ 㸦Ogura-Tsujita et al. 2012㸧
㸪㏦⢊ᵝᘧࡢ㠃࡛ࡶ㸪ⴥࢆ⏕ࡌ࡞࠸✀⩌ࡢ㐍໬࡟ඛ⾜ࡋ࡚㸪୙᏶඲࡞ᙧ࡛⮬ື⮬ᐙཷ⢊ࡀ㐍໬ࡋ࡚࠸ࡿࡢࡣ⯆࿡῝࠸ࠋ ࡇ࠺ࡋࡓ⮬Ṫࡢ㐍໬ࡣᬯ࠸ᯘᗋ࡛☜ᐇ࡟⦾Ṫࡍࡿࡢ࡟ᙺ❧ࡗࡓ࡜⪃࠼ࡽࢀࡿࠋ⳦ᚑᒓ
ᰤ㣴᳜≀ࡀ⮬ື⮬ᐙཷ⢊ࢆ᥇⏝ࡋࡸࡍ࠸࡜࠸࠺ண

ࡣࡇࢀࡲ࡛࡟ࡶᏑᅾࡋࡓࡀ

㸦Bidartondo 2005㸧㸪⣔⤫㛵ಀࡀ᫂ࡽ࠿࡜࡞ࡗ࡚࠸ࡿ✀⩌࡛㸪㔝እほᐹ࡜⿄᥃ࡅᐇ㦂ࢆ
㏻ࡋ࡚ᐇドࡋࡓ౛࡜ࡋ࡚ᮏ◊✲ࡣ౯್ࡀ࠶ࡿࠋࡲࡓ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ࡞࠿࡛ࡶ㸪⮬Ṫࢆࡲࡗࡓࡃ⾜ࢃ࡞࠸࢝ࣥࣛࣥ࡜ࢩࣗࣥࣛࣥࡣ㸪ẚ㍑ⓗ᫂ࡿ࠸⎔ቃࢆዲࡳ㸪㛤ⰼ᫬ᮇࡶᯘෙࡀ㛢㙐ࡋ࡚ࡋࡲ࠺๓ࡢ෤ࡸ᪩᫓࡛࠶ࡿⅬࡶࡁࢃࡵ࡚⯆࿡῝࠸ࠋ୍⯡ⓗ࡟ࡣ㸪᪻⹸ࡀ࠶ࡲࡾάືࡋ࡚࠸࡞࠸෤ࡸ᪩᫓࡟㛤ⰼࡍࡿࡇ࡜ࡣ୙฼࡜ࡉࢀ㸪ࡇࡢࡼ࠺࡞᫬ᮇ࡟㛤ⰼࡍࡿ᳜≀ࡣ㫽࡞࡝᪻⹸௨እࡢ㏦⢊⪅࡟౫Ꮡࡍࡿሙྜࡀከ࠸ 㸦Kunitake et al. 2004, Fang etal. 2012 ࡞࡝㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪࢝ࣥࣛࣥ࡜ࢩࣗࣥࣛࣥࡣ㸪࡯࠿ࡢ㏆⦕✀࡜ྠᵝ࡟ࣁࢼࣂࢳࢆ㏦⢊⪅࡜ࡋ࡚࠸ࡿ࡟ࡶ㛵ࢃࡽࡎ㸪෤ࡸ᪩᫓࡟㛤ⰼࡍࡿࠋࡇࢀࡽࡢ㛤ⰼ᫬ᮇࡶ㸪ࡼࡾ᫂ࡿ࠸⎔ቃ࡛ཷ⢊ࢆ㐩ᡂࡍࡿࡓࡵࡢ㐺ᛂ࡛࠶ࡿྍ⬟ᛶࡀ࠶ࡿࠋ

㸳⳦ᚑᒓᰤ㣴᳜≀࡟࠾ࡅࡿ㏦⢊⪅ࡢ㔜せᛶ๓㡯࡛㏙࡭ࡓ㏻ࡾ㸪ᬯ࠸ᯘᗋ࡟⏕⫱ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀࡟࡜ࡗ࡚㸪ࢥࣁࢼࣂࢳࡸ࣑ࢶ

K. Suetsugu - 7

植物科学最前線 5:100 （2014）ࣂࢳ࡞࡝ࡢ᫂ࡿ࠸⎔ቃࢆዲࡴ᪻⹸࡟㏦⢊ࢆ㢗ࡿࡇ࡜ࡣ㠀ᖖ࡟㞴ࡋ࠸࡜⪃࠼ࡽࢀࡿࠋࡋ࠿
ࡋ࡞ࡀࡽ㸪ᬯ࠸⎔ቃ࡟㐍ฟྍ⬟࡞᪻⹸࡟㏦⢊ࢆクࡍࡇ࡜ࡀ࡛ࡁࢀࡤ㸪ᬯ࠸ᯘᗋ࡛ࡶ௚Ṫࢆ⾜࠺ࡇ࡜ࡀྍ⬟࡛ࡣ࡞࠸ࡔࢁ࠺࠿ࠋ౛࠼ࡤ࣐ࣝࣁࢼࣂࢳࡣ㸪యㄪ⠇ࡀྍ⬟࡞ࡓࡵ㸪ᬯ࠸⎔ቃ࡛ࡶ᥇㣵ࢆ⾜࠺ഴྥࡀ࠶ࡿࠋࡋࡓࡀࡗ࡚㸪࣐ࣝࣁࢼࣂࢳࢆ࿧ࡧᐤࡏࡿࡇ࡜ࡀ࡛
ࡁࢀࡤ㸪௚Ṫ࡛⏕άྐࢆᅇࡍࡇ࡜ࡶྍ⬟࡛࠶ࡿ࡜⪃࠼ࡽࢀࡿࠋ஦ᐇ㸪ࢠࣥࣜࣙ࢘ࢯ࢘ࡸ
ࣔ࢖࣡ࣛࣥࢆྵࡴ࠸ࡃࡘ࠿ࡢ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪኱㔞ࡢ⻤ࢆሗ㓘࡜ࡍࡿࡇ࡜࡛㸪ࢺ࣐ࣛࣝࣁࢼࣂࢳࢆึࡵ࡜ࡍࡿ࣐ࣝࣁࢼࣂࢳࢆ㏦⢊⪅࡜ࡋ࡚฼⏝ࡋ㸪⮬ື⮬ᐙཷ⢊ࡶ⾜ࢃ࡞࠸㸦Ushimaru & Imamura 2002, ᮎḟ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡲࡓࢩࣙ࢘࢟ࣛࣥࡣ㸪⻤ࢆࡔࡍࡇ࡜
࡞ࡃ㸪㩭ࡸ࠿࡞ⰼࢆࡘࡅ㸪᥇㣵⤒㦂ࡢὸ࠸࣐ࣝࣁࢼࣂࢳ࡟㣵ࡀ࠶ࡿ࡜ᛮࢃࡏࡿࡇ࡜࡛㸪㏦⢊ࢆ㐩ᡂࡋ࡚࠸ࡿ 㸦ᮎḟ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡉࡽ࡟⳦ᚑᒓᰤ㣴᳜≀ࡢ୰࡟ࡣ㸪ࡇࢀࡲ࡛ゼⰼᛶ࡜⪃࠼ࡽࢀ࡚࠸࡞࠿ࡗࡓ᪻⹸ࢆ㏦⢊⪅࡜ࡋ࡚฼⏝ࡍࡿࡶࡢࡲ࡛Ꮡᅾࡍࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡾࡘࡘ࠶ࡿࠋࡇࡢࡼ࠺࡞౛࡜ࡋ࡚㸪ᡃࠎࡣ㸪࢜ࢽࣀࣖ࢞ࣛᒓࣖࢶࢩࣟࣛࣥ⠇ࡢ≉Ṧ࡞ཷ⢊ᵝᘧࢆⓎぢࡋࡓ 㸦ᮎḟࡽ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࢩࣙ࢘ࢪࣙ࢘ࣂ࢚ࡣ㸪⭉ᩋࡀጞࡲࡗࡓᯝᐇࡸ࢟ࣀࢥࢆ᥈ࡋ࡚ᬯ࠸ᯘᗋࢆ㣕ࡧᅇࡿࡀ㸪ࣖࢶࢩࣟࣛࣥ⠇ࡢከࡃࡢ✀ࡣ㸪࢟ࣀࢥ⮯ࡸⓎ㓝⮯࡟ᨃែࡍࡿࡇ࡜࡛㸪ࢩࣙ࢘ࢪࣙ࢘ࣂ࢚࡟㏦⢊ࢆクࡋ࡚࠸ࡿࠋࡲࡓ⯆࿡῝࠸ࡇ࡜࡟ࡇࡢࣖࢶࢩࣟࣛࣥ⠇ࡢ✀⩌࡟ࡣ㸪㏆ࡃ࡟࢟ࣀࢥࡀᏑᅾࡍࡿࡇ࡜࡛㸪ཷ⢊⋡ࡀୖࡀࡿ✀㢮ࡀ࠶ࡿࡇ࡜ࡶ᫂ࡽ࠿࡟࡞ࡗࡓࠋࣖࢶ
ࢩࣟࣛࣥࡣ⭉⏕⳦࡟ᐤ⏕ࡍࡿ≉ᚩ࠿ࡽ㸪ᬯࡃ‵ࡗࡓ⎔ቃ࡟⏕⫱ࡍࡿࡇ࡜ࡀከࡃ㸪஦ᐇ㸪ࣖࢶࢩࣟࣛࣥࡢ⏕࠼ࡿ࿘㎶࡟ࡣ⭉⏕⳦ࡢᏊᐇయࡀฟ⌧ࡍࡿࡇ࡜ࡶከ࠸ࠋࡘࡲࡾࣖࢶࢩࣟ
ࣛࣥࡣ㸪⮬㌟ࡢࡑࡤ࡟ከࡃ⏕࠼ࡿࠕ࢟ࣀࢥࠖ࡟㏦⢊⪅ࢆㄏᘬࡋ࡚ࡶࡽ࠺ࡇ࡜ࡀྍ⬟࡞ࡢ࡛࠶ࡿ 㸦Suetsugu & Kato 2013, ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࣖࢶࢩࣟࣛࣥࡣ㸪࿘㎶࡟࢟ࣀࢥࡀ࡞࠸⎔ቃ࡛ࡶ㸪ཷ⢊ࢆ㐩ᡂࡍࡿࡇ࡜ࡀ࡛ࡁࡿࡓࡵ㸪࢟ࣀࢥ࡟ࡼࡿ㏦⢊⪅ࡢㄏᘬ࡟᏶඲࡟㢗ࡗ࡚࠸ࡿࢃࡅ࡛ࡣ࡞࠸ࡶࡢࡢ㸪ࣖࢶࢩࣟࣛࣥࡣ㸪ᰤ㣴ࡢࡳ࡞ࡽࡎ㏦⢊ࢧ࣮ࣅࢫࡶ㸪࢟ࣀࢥ࡟౫Ꮡࡋ࡚࠸ࡿ࡜࠸࠼ࡿࠋ
 Bidartondo 㸦2005㸧 ࡣ㸪ࠕࡍ࡛࡟ᆅୗ㒊࡛⳦᰿⳦ࢆ㦄ࡋ࡚࠸ࡿ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪ᆅୖ㒊࡛⻤ࢆࡔࡉࡎ࡟㏦⢊⪅ࢆ㦄ࡍࡇ࡜ࡣ࡛ࡁ࡞࠸ࠖ࡜࠸࠺௬ㄝࢆᥦၐࡋࡓࡀ㸪ࢩࣙ࢘࢟ࣛࣥࡶࣖࢶࢩࣟࣛࣥࡶ⻤ࢆฟࡉ࡞࠸ࡓࡵ㸪ࡇࡢ௬ㄝ࡟ࡣヱᙜࡋ࡞࠸౛እࡶ࠶ࡿ࡜ゝ࠼ࡿࠋࡋ࠿ࡋ࡞ࡀࡽ㸪⻤࡞࡝ࡢሗ㓘ࢆ㏦⢊⪅࡟ᥦ౪ࡋ࡞࠸ 㸦௨ୗ↓ሗ㓘ⰼ࡜㏙࡭ࡿ㸧 ⳦ᚑᒓᰤ㣴᳜≀࡛ࡣ㸪≉࡟⮬Ṫࡀ༟㉺ࡋ࡚࠸ࡿࡢࡣ஦ᐇ࡛࠶ࡿࠋࡇࡢࡇ࡜࡟ࡣ㸪↓ሗ㓘ⰼࡣ㏦⢊⪅࡟ሗ㓘ࢆᥦ౪ࡋ࡞࠸ࡓࡵ㸪⊂❧ᰤ㣴ᛶ᳜≀ࡢሙྜ࡛ࡶ㏦⢊⪅ࡀゼࢀࡿ㢖ᗘࡀᴟࡵ࡚
ప࠸ࡇ࡜ࡀ࠾ࡑࡽࡃ㛵ಀࡋ࡚࠸ࡿ 㸦Neiland & Wilcock 1998, Tremblay et al. 2005㸧ࠋࡘࡲࡾඖࠎእ஺㓄ࡢᶵ఍ࡀᑡ࡞࠿ࡗࡓ↓ሗ㓘ⰼࡢ⳦ᚑᒓᰤ㣴᳜≀ࡢ♽ඛ✀ࡣ㸪ᚑᒓᰤ㣴ᛶࢆ⋓ᚓࡋᬯ࠸⎔ቃ࡟㐍ฟࡍࡿ㐣⛬࡛㸪ࡼࡾᙉ࠸㏦⢊⪅ไ㝈ࢆࡇ࠺ࡴࡿࡇ࡜࡟࡞ࡗࡓ࡜⪃࠼ࡽࢀࡿࠋࡑࡢࡼ࠺࡞᮲௳ୗ࡛ࡣ㸪⦾Ṫࡢಖ㞀࡜ࡋ࡚⮬ື⮬ᐙཷ⢊࡛ࡁࡿᛶ㉁ࡀ㸪ࡼࡾ᭷฼࡟ാࡃࡓࡵ㸪⻤ࢆฟࡉ࡞࠸⳦ᚑᒓᰤ㣴᳜≀࡛ࡣ㸪⮬Ṫࡀ༟㉺ࡋ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿࠋ

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植物科学最前線 5:101 （2014） ࡇࢀࡲ࡛⳦ᚑᒓᰤ㣴᳜≀࡛⮬Ṫࡀ༟㉺ࡍࡿせᅉ࡜ࡋ࡚ࡣ㸪ᐤ⏕⏕ά࡟క࠺㈨※ไ㝈ࡀཎᅉ࡜࠸ࢃࢀ࡚ࡁࡓ 㸦Takahashi et al. 1993, Zhang & Saunders 2000㸧
ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪㸦1㸧⮬Ṫࢆ⾜࠺⳦ᚑᒓᰤ㣴᳜≀࡛ࡶ㩭ࡸ࠿࡞ⰼࢆᣢࡘࡶࡢࡀከ࠸ࡇ࡜ 㸦Zhou et al. 2012,
Suetsugu 2013; ᅗ 5㸧 ࡸ㸪
㸦2㸧 ᬯ࠸⎔ቃ࡟⏕ᜥࡍࡿ᪻⹸ࢆ㏦⢊⪅࡟᥇⏝ࡋ࡚࠸ࡿ⳦ᚑ
ᒓᰤ㣴᳜≀࡛ࡣ㸪⮬Ṫࢆ⾜ࢃ࡞࠸ࡇ࡜ࡶከ࠸ࡇ࡜ࢆ⪃࠼ྜࢃࡏࡿ࡜ 㸦Hentrich et al. 2010,
Klooster & Culley 2009㸧
㸪㈨※ไ㝈ࡔࡅ࡛ࡣ࡞ࡃ㸪ග⎔ቃࡶ⳦ᚑᒓᰤ㣴᳜≀ࡢ⦾Ṫᵝᘧ࡟ᙳ㡪ࢆ୚࠼࡚࠸ࡿ࡜࠸࠼ࡿࠋࡇࡢࡇ࡜ࢆ⿬௜ࡅࡿドᣐ࡜ࡋ࡚ࡣ㸪࢜ࢽࣀࣖ࢞ࣛᒓࡢ࢜ࢽ
ࣀࣖ࢞ࣛࡣ㸪ᬯ࠸⎔ቃ࡛ࡣᑓࡽ࢔࣏࣑ࢡࢩࢫ࡟ࡼࡾ✀Ꮚࢆ⏕⏘ࡍࡿࡀ㸪᫂ࡿ࠸ࢠࣕࢵࣉ
࡞࡝࡟⏕⫱ࡍࡿಶయ࡛ࡣ㸪࠿࡞ࡾࡢ๭ྜ࡛ࢥࣁࢼࣂࢳ࡟ࡼࡿ௚Ṫࢆ⾜ࡗ࡚࠸ࡿࡇ࡜ࡀᣲࡆࡽࢀ 㸦ᮎḟ࣭ຍ⸨ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡲࡓᐟ୺࡜࡞ࡿ⳦ࡢࣁࣅࢱࢵࢺ㑅ዲᛶࡶᙳ㡪ࡍࡿࡓࡵ୍ᴫ࡟ࡣゝ࠼࡞࠸ࡀ㸪ࣁࢼࣂࢳ፹ࡢ⳦ᚑᒓᰤ㣴᳜≀ࡣ᫂ࡿ࠸⎔ቃ࡟⏕⫱ࡍࡿሙྜࡀከ࠸ 㸦Suetsugu & Kato 2012, ᮎḟ ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋࡘࡲࡾ⳦ᚑᒓᰤ㣴ᛶࡢ⋓ᚓ࡟ࡼࡾ㸪௚ࡢ➇த┦ᡭࡢᑡ࡞࠸ᬯ࠸ᯘᗋ࡛⏕Ꮡྍ⬟࡟࡞ࡿ࡜ࡉࢀ࡚ࡁࡓࡀ㸪ᐇ㝿ࡢ࡜ࡇࢁࡣ㸪㏦⢊ᵝᘧࡢⅬ࡛ࡶ≉ู࡞㐺ᛂࢆ㐙ࡆ࡞ࡅࢀࡤ㸪ᬯᡤ࡬ࡢ㐍ฟࡣ୙ྍ⬟࡞ࡢ࡛࠶ࡿࠋᅗ 㸬⮬ື⮬ᐙཷ⢊ྍ⬟࡞⳦ᚑᒓᰤ㣴᳜≀㸦㩭ࡸ࠿࡞ⰼࢆࡘࡅࡿࡶࡢࡀከࡃ㸪୰࡟ࡣ኱㔞ࡢⰼ⻤ࢆศἪࡍࡿࡶࡢࡶᏑᅾࡋ㸪௚Ṫࡢྍ⬟ᛶࢆ᏶඲࡟ᨺᲠࡋ࡚࠸ࡿヂ࡛ࡣ࡞࠸ࡇ࡜ࢆ᝿
㉳ࡉࡏࡿ  ୖẁᕥ࠿ࡽ㡰࡟㸪࣍ࢡࣜࢡ࣒ࣚ࢘ࣛࣥ㸪ࣄࢼࣀࢩࣕࢡࢪࣙ࢘㸪ࢩࣟࢩࣕࢡࢪ
ࣙ࢘㸪࣐ࣖࣛࣥ㸪ࢶࢳ࢔ࢣࣅ㸪ࢱࢩࣟࣛࣥ

K. Suetsugu - 9

植物科学最前線 5:102 （2014）ࡲࡓ࢜ࢽࣀࣖ࢞ࣛࡢࡼ࠺࡟㸪⮬Ṫ࡜௚Ṫࢆేࡏᣢࡘ⦾Ṫᵝᘧࡣ㸪ከࡃࡢ⳦ᚑᒓᰤ㣴᳜≀࡟࡜ࡗ࡚㔜せ࡞ᙺ๭ࢆᯝࡓࡋ࡚࠸ࡿࡢ࠿ࡶࡋࢀ࡞࠸ 㸦ᅗ 6㸧ࠋ⳦ᚑᒓᰤ㣴᳜≀࡜ࡑࡢ⳦᰿⳦ࡢ㛵ಀࡣ㸪ᐤ⏕⪅࡜ᐟ୺ࡢ㛵ಀ࡜࡜ࡽ࠼ࡿࡇ࡜ࡀ࡛ࡁࡿࠋࡇࢀࡲ
࡛ࡢ◊✲ࡢከࡃࡀ㸪⳦᰿ඹ⏕࡟࠾ࡅࡿඹ⏕࠿ࡽᐤ⏕࡬ࡢኚ໬࡟ࡣ≉ᐃࡢ⳦࡬ࡢ≉Ṧ໬ࡀᚲせ࡛࠶ࡿࡇ࡜ࢆ ♧ ၀ ࡋ ࡚ ࠸ ࡿ 㸦 Leake 2004,
Bidartondo 2005㸧ࠋࡇࡢඹ⏕⳦ࡢࢩ ᅗ ࢜ࢽࣀࣖ࢞ࣛ࡜ࡑࡢ㏦⢊⪅ࡢࢥࣁࢼࣂࢳ

ࣇࢺࡸ≉Ṧ໬ࡢ㉳ࡇࡿ⌮⏤࡟㛵ࡍࡿ᭷ຊ࡞௬ㄝࡀ㸪࡯࠿ࡢᐤ⏕⪅ʊᐟ୺⣔࡜ྠᵝ࡟㸪⳦ᚑᒓᰤ㣴᳜≀࡜ᐟ୺࡜ࡢ㛫࡛㌷ᣑ➇தࡀാ࠸࡚࠸ࡿ࠿ࡽ࡜࠸࠺ࡶࡢ࡛࠶ࡿ 㸦Leake
2004, Selosse & Roy 2009㸧ࠋࡇࡢࡼ࠺࡞ᩛᑐⓗ࡞┦஫స⏝ࡢሙ࡛ࡣ㸪ྠࡌ㐺ᛂᗘࢆ⥔ᣢࡍࡿࡓࡵ࡟㸪┦ᡭ࡟ᑐࡋ࡚⤯࠼ࡎ㐺ᛂࡋ⥆ࡅ࡞ࡅࢀࡤ࡞ࡽ࡞࠸ࠋࡇࡢࡇ࡜ࡣ㸪ࣝ࢖ࢫ࣭࢟ࣕࣟࣝࡢ❺ヰࠕ㙾ࡢᅜࡢ࢔ࣜࢫࠖ࡟Ⓩሙࡍࡿ㉥ࡢዪ⋤ࡢࠕྠࡌሙᡤ࡟࡜࡝ࡲࡿࡓࡵ࡟
ࡣ㸪඲㏿ຊ࡛㉮ࡾ⥆ࡅ࡞ࡅࢀࡤ࡞ࡽ࡞࠸ࠖ࡜࠸࠺ࢭࣜࣇ࡟࡞ࡒࡽ࠼࡚㉥ࡢዪ⋤௬ㄝ࡜࿧ࡤࢀ࡚࠸ࡿ 㸦Ladle 1992㸧ࠋࡑࡋ࡚㐺ᛂ㐍໬ࡢ㏿ᗘࢆ᪩ࡵࡿࡼ࠺࡞ᛶ㉁ࡀ᭷ᛶ⏕Ṫ࡛࠶ࡾ㸪ᩛᑐⓗ࡞┦஫స⏝ࡢᏑᅾࡇࡑࡀ㸪᭷ᛶ⏕ṪࡢᏑᅾព⩏ࢆᢸಖࡍࡿ㔜せ࡞せᅉ࡜ぢ࡞ࡉࢀ࡚࠸ࡿ 㸦Ladle 1992㸧ࠋᮏ᮶㸪⳦᰿ඹ⏕⣔࡛ࡣ㸪࠾஫࠸ࡀⰋ࠸ࣃ࣮ࢺࢼ࣮࠿࡝࠺࠿ࢆぢศࡅࡿ௙⤌ࡳࡀ࠶ࡾ㸪ࡼࡾከࡃ❅⣲ࡸࣜࣥࢆᥦ౪࡛ࡁࡿ⳦㸪ྠࡌࡃ㸪ࡼࡾከࡃⅣ⣲ࢆ౪⤥࡛ࡁࡿ᳜≀ࡀ㸪┦ᡭ࡟㑅ࡤࢀࡿ࡜ࡉࢀࡿ 㸦Kiers et al. 2011㸧ࠋࡘࡲࡾ᳜≀ࡶ⳦ࡶࣃ࣮ࢺࢼ࣮ࡢၿࡋᝏࡋࢆࡋ
ࡗ࠿ࡾ࡜㆑ูࡋ࡚࠾ࡾ㸪ཷࡅྲྀࡿ㈨※࡟ᛂࡌ࡚㏦ࡾ㎸ࡴ㣴ศࡢ㔞ࢆㄪ⠇ࡋ࡚࠸ࡿࡢ࡛࠶
ࡿࠋࣛࣥ⛉᳜≀࡜ࡑࡢ⳦᰿⳦ࡢ㛫࡛ࡣ㸪ࡇࡢࡼ࠺࡞ࠕᑂၥࠖࡢ௙⤌ࡳࡣ᫂ࡽ࠿࡟࡞ࡗ࡚
࠸࡞࠸ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ࣛࣥ⛉᳜≀࡛ࡶከࡃࡢሙྜࡣ㸪ⓎⱆᙜึࡣⅣ⣲ࢆྵࡴ㣴ศࢆ⳦࡟౫Ꮡࡍࡿࡶࡢࡢ㸪⥳ⴥࢆᒎ㛤ࡋ࡚࠿ࡽⅣ⣲ࢆ⳦࡟㏉㑏ࡍࡿࡇ࡜࡛㸪┦฼ඹ⏕ࢆႠࢇ࡛࠸ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ 㸦Cameron et al. 2006, 2008, Hynson et al. 2009㸧ࠋ࡟ࡶ࠿࠿ࢃࡽࡎ
⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪ᮏ᮶㸪᳜≀ഃࡢᮏ᮶ࡢᥦ౪≀࡛࠶ࡿⅣ⣲ࢆ୚࠼࡞࠸ࡔࡅ࡛ࡣ࡞ࡃ㸪㏫࡟ዣ࠸ྲྀࡗ࡚⏕⫱ࢆྍ⬟࡟ࡋ࡚࠸ࡿࠋࡼࡗ࡚ࡑࡢ⫼ᬒ࡟ࡣ㸪ඹ⏕ࢩࢢࢼࣝࡢᨃែ࡞࡝
ࡢᕦጁ࡞ࢩࢫࢸ࣒ࡀᏑᅾࡍࡿࡇ࡜ࡀண᝿ࡉࢀࡿࠋ㉥ࡢዪ⋤௬ㄝ࡟ೌࡗ࡚⪃࠼࡚ࡳࡿ࡜㸪ᐟ୺࡜࡞ࡿ⳦㢮ࢆ㦄ࡋ⥆ࡅࡿࡓࡵ࡟ࡣ㸪⤯࠼ࡎ㐺ᛂࡋ⥆ࡅ࡞ࡅࢀࡤ࡞ࡽࡎ㸪ᑓࡽ⮬Ṫࡢࡳࢆ⾜࠺⏕άྐࡣᴟࡵ࡚୙฼࡜⪃࠼ࡽࢀࡿࠋࡲࡓ㸪ࡇࡢࡼ࠺࡞ᐤ⏕⪅ʊᐟ୺⣔࡟㛵ࢃࡿၥ㢟Ⅼࢆ⪃࠼ࡎ࡜ࡶ㸪⮬ᐙཷ⢊ࡢࡳ࡟㢗ࡿ⏕άྐ࡟ࡣ㸪㏆஺ᙅໃࢆึࡵ࡜ࡍࡿᵝࠎ࡞୙฼┈ࡀᏑᅾࡍࡿ 㸦Lloyd 1992㸧ࠋࡇࡢࡼ࠺࡞୙฼┈ࢆゎᾘࡍࡿࡢ࡟㸪⛥࡟㉳ࡇࡿ௚Ṫࡀ

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植物科学最前線 5:103 （2014）㔜せ࡞ാࡁࢆᯝࡓࡋ࡚
࠾ࡾ㸪⮬Ṫ࡜௚Ṫࢆేࡏᣢࡘ⦾Ṫᵝᘧࡀ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ⏕άࢫࢱ࢖ࣝ࡟࡜ࡗ࡚ࡣ㐺ࡋ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿࠋ
ࡋ࠿ࡋ࡞ࡀࡽᴟ➃࡟ᬯᡤ࡟㐍ฟࡋࡓ⳦ᚑᒓᰤ
㣴᳜≀ࡣ㸪㛤ⰼࡍࡿࡶࡢ࡛ࡶ௚Ṫࡢᶵ఍ࢆ࡯
ࡰ᏶඲࡟ኻࡗ࡚࠾ࡾ㸪୰࡟ࡣࡘࡰࡳࡢ≧ែ࡛⮬ᐙཷ⢊ࡍࡿ᏶඲⮬Ṫᅗ ᏶඲⮬Ṫᆺࡢⰼ࡛࠶ࡿ㛢㙐ⰼࡢࡳࢆࡘࡅࡿࢶ࣑࣎ࣖࢶࢩࣟࣛࣥ ᕥ ࡜ࢱࢣࢩ࣐ࣖࢶࢩࣟࣛࣥ ྑ ᆺࡢ㛢㙐ⰼࡢࡳࢆࡘࡅࡿ✀㢮ࡶᏑᅾࡍࡿ 㸦Hsu et al. 2012, Suetsugu 2013, Suetsugu et al.
2013; ᅗ 7㸧ࠋࡑࡢࡼ࠺࡞✀㢮ࡣ㸪㐜࠿ࢀ᪩࠿ࢀ㸪⳦ࢆ㦄ࡍࡇ࡜ࡀ࡛ࡁ࡞ࡃ࡞ࡾ㸪⤯⁛ࡍࡿ㐠࿨࡟࠶ࡿࡢ࠿ࡶࡋࢀ࡞࠸ࠋࡶࡋࡑ࠺࡛࠶ࡿ࡞ࡽࡤ㸪⳦࠿ࡽ㣴ศࢆ࠿ࡍࡵྲྀࡿ࡜࠸࠺
ᙼࡽࡢ⏕ࡁ᪉ࡣ㸪୍ぢ㠀ᖖ࡟㈼࠸ࡼ࠺࡟ぢ࠼ࡿࡶࡢࡢ㸪ࡑࡢ௚ࡢ㠃࡛ᩘࠎࡢ୙฼┈ࢆక࠺ࡶࡢ࡛࠶ࡾ㸪㐍໬ࡢ㐣⛬࡛⿄ᑠ㊰࡟㏞࠸㎸ࡳ㸪㛗࠸㐍໬ࡢ㐣⛬࡛ࡣᾘ࠼ࡺࡃ㐠࿨࡟࠶ࡿ࡜࡜ࡽ࠼ࡿࡇ࡜ࡀ࡛ࡁࡿ࠿ࡶࡋࢀ࡞࠸ࠋ

㸴ࡲ࡜ࡵ࡜௒ᚋࡢᒎᮃ
ᡃࠎࡢ◊✲ࢆྵࡴ㏆ᖺࡢ◊✲ࡢ㐍ᒎ࡟ࡼࡾ㸪⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪ࡑࡢ≉␗࡞ᛶ㉁ࢆ⋓ᚓ࣭⥔ᣢࡍࡿࡓࡵ࡟㸪ᐟ୺⳦࡟ᑐࡋ࡚ࡔࡅ࡛࡞ࡃ㸪௚ࡢ⏕≀࡜ࡢ┦஫స⏝࡟࠾࠸࡚ࡶᩘࠎ
ࡢᩘወ࡞㐺ᛂࢆ㐙ࡆ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡾࡘࡘ࠶ࡿࠋ୺࡟⮬Ṫࢆ⾜ࡗ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚ࡁࡓࡓࡵ࠿㸪⳦ᚑᒓᰤ㣴ᛶࡢ✀⩌࡟࠾࠸࡚ࡣ㸪ࠕᕼᑡᛶࠖ㸪ࠕศᕸࠖ㸪ࠕከᵝ໬ࠖ࡞࡝ࡣ୺࡟ᐟ୺⳦࡜ࡢ┦஫స⏝ࡢᩥ⬦࡛ࡢࡳ㆟ㄽࡉࢀ㸪ࡑࡢ௚ࡢせᅉࡀ⪃៖ࡉࢀࡿࡇ࡜ࡣ
ᑡ࡞࠿ࡗࡓ 㸦Taylor et al. 2003a,b, Merckx & Bidartondo 2008, Bougoure et al. 2009, 2010,
Hazard et al. 2012㸧ࠋࡋ࠿ࡋ࡞ࡀࡽ㸪ከࡃࡢ⳦ᚑᒓᰤ㣴ᛶࡢ✀ࢆྵࡴࣛࣥ⛉ࡣ㸪㏦⢊ඹ⏕࡜࠸࠺㠃࡛ࡶ㠀ᖖ࡟≉Ṧ໬ࡋ࡚࠾ࡾ㸪≉ᐃࡢ㏦⢊⪅ࢆ࿧ࡧᐤࡏࡿࡓࡵ࡟㸪⊂≉࡞ⰼᙧែࢆ㐍໬ࡉࡏ࡚࠸ࡿࡶࡢࡀ࡯࡜ࢇ࡝࡛࠶ࡿ 㸦Cozzolino & Widmer 2005, Waterman &Bidartondo 2008, Waterman et al. 2011㸧
ࠋࡲࡓ㏦⢊⪅࡜ᐦ᥋࡞┦஫స⏝ࢆ⠏࠸࡚࠸ࡿࡓࡵ㸪⥳ⴥࢆᣢࡘ✀㢮࡛ࡣ㸪ࡑࡢᕼᑡᛶࡣ㸪㏦⢊⪅ࡢ≉Ṧ໬ࡢ⛬ᗘࡸ㸪㏦⢊⪅ࡢᕼᑡᛶ࡜኱ࡁ
ࡃ㛵ಀࡋ࡚࠸ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ㸦Neiland & Wilcock 1998, Dixon 2009, Phillips
et al. 2011㸧ࠋࡲࡓࣛࣥ⛉ࡢ⇿Ⓨⓗ࡞ከᵝ໬ࡢ⫼ᬒ࡟ࡣ㸪ཷ⢊ᵝᘧࡸ㏦⢊⪅ࡢࢩࣇࢺࡀ኱
ࡁࡃ㛵ࢃࡗ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ 㸦Cozzolino & Widmer 2005㸧ࠋ⳦࠿ࡽ඲࡚ࡢ㣴ศࢆ␎ዣࡋ࡚࠸ࡿ⳦ᚑᒓᰤ㣴᳜≀ࡢࠕᕼᑡᛶࠖ㸪ࠕศᕸࠖ㸪ࠕከᵝ໬ࠖࡀ⳦࡜ࡢ㛵ࢃࡾ࡟ࡼࡾつ

K. Suetsugu - 11

植物科学最前線 5:104 （2014）ᐃࡉࢀ࡚࠸ࡿ࡜࠸࠺⪃࠼ࡣ㠀ᖖ࡟㨩ຊⓗ࡞ࡶࡢ࡛࠶ࡿࡀ㸪㏦⢊ᵝᘧ㸪✀Ꮚᩓᕸᵝᘧ㸪ࡑࡢ௚㠀⏕≀Ꮫⓗ࡞せᅉࢆ⪃៖࡟ධࢀ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢᐇែ࡟㏕ࡿࡇ࡜ࡀ㔜せ࡛࠶ࢁ࠺ࠋㅰ㎡ᮏ◊✲ࡣ㸪᪥ᮏᏛ⾡᣺⯆఍ࡢ≉ู◊✲ဨዡບ㈝ 㸦12J00602㸧 ࡢᨭ᥼ࢆཷࡅ࡚࠸ࡿࠋ
ࡲࡓ◊✲඲⯡㸪ཎ✏ᇳ➹࡟࠶ࡓࡗ࡚㈗㔜࡞࢔ࢻࣂ࢖ࢫࢆ㡬࠸ࡓᕝ໭⠜༤ኈ࡟ᚚ♩⏦ࡋୖࡆࡓ࠸ࠋࡲࡓ࢔ࣝࣅࣀࡢ࢟ࣥࣛࣥࡢ෗┿ࡣ㸪෗┿ᐙࡢᨾ᪂஭࿴ஓẶ࡟ࡈᥦ౪࠸ࡓࡔ࠸ࡓࠋ
ࡉࡽ࡟⤯⁛༴᝹᳜≀ࡀከ࠸㸪⳦ᚑᒓᰤ㣴᳜≀ࢆ◊✲ࡍࡿ࡟࠶ࡓࡾከࡃࡢ᳜≀ឡዲᐙࡢ᪉ࠎ࡟㸪⮬⏕ᆅሙᡤࡢ᱌ෆࡸ᝟ሗᥦ౪࡞࡝ࡢᙧ࡛࠾ୡヰ࡟࡞ࡗ࡚࠸ࡿࠋࡇࡇ࡟ࡍ࡭࡚ࡢ
ேࢆᣲࡆࡿࡇ࡜ࡣ࡛ࡁ࡞࠸ࡀ㸪ከᩘࡢ᪉ࠎࡢᨭ᥼ࢆཷࡅ㸪◊✲ࡀྍ⬟࡜࡞ࡗ࡚࠸ࡿࡇ࡜
ࢆグࡋ࡚ឤㅰࡋࡓ࠸ࠋ

䢢ᘬ⏝ᩥ⊩
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Waterman, R.J., & Bidartondo, M.I. 2008. Deception above, deception below: linking pollinationand mycorrhizal biology of orchids. J. Exp. Bot. 59: 1085-1096.
Yagame, T., & Yamato, M. 2013. Mycoheterotrophic growth of Cephalanthera falcata (Orchidaceae)
in tripartite symbioses with Thelephoraceae fungi and Quercus serrata (Fagaceae) in pot culture
condition. J. Plant Res. 126: 215-222.
Yukawa, T., Miyoshi, K., & Yokohama, J. 2002. Molecular phylogeny and character evolution ofCymbidium (Orchidaceae). Bull. Natl. Sci. Mus. Tokyo 28: 129-139.
Zhang, D., & Saunders, R.M.K. 2000. Reproductive biology of a mycoheterotrophic species,
Burmannia wallichii (Burmanniaceae). Bot. J. Linn. Soc. 132: 359-367.

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Zhou, X., Lin, H., Fan, X.L., & Gao, J.Y. 2012. Autonomous self-pollination and insect visitation ina saprophytic orchid, Epipogium roseum (D.Don) Lindl. Aust. J. Bot. 60: 154-159.

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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, ࢧ࢝ࢿࣛࣥ
ࢆ⤒⏤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࢖ࣚ࢘ࢩ
ࣙ࢘ࣟᒓ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


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植物科学最前線 5:112 （2014）ᅗ 2 ⭉⏕⳦࡜⳦᰿ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀ࠋA, ࢜ࢽࣀࣖ࢞ࣛ; B, ࢶࢳ࢔ࢣࣅ; C, ࢱࢩࣟࣛࣥ

3 ࡘ┠ࡣ͂࢔࣮ࣂࢫ࣮࢟ࣗࣛ⳦᰿⳦̓
㸦AM ⳦㸧࡜ඹ⏕ࡍࡿ⳦ᚑᒓᰤ㣴᳜≀࡛࠶ࡿࠋࢢ࣒ࣟ
ࢫ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୰࡟Ꮡᅾࡍ
ࡿ⳦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, ࢱࢾ࢟ࣀࢩ
ࣙࢡࢲ࢖ྜ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ᡭἲࢆࡶࡗ
࡚ࡋ࡚ࡶศᏊྠᐃࡉࢀࡓ⳦✀ࡀ⳦ᚑᒓᰤ㣴᳜≀ࡢ⏕⫱࡟ཬࡰࡍᙳ㡪ࢆホ౯࡛ࡁ࡞࠸ࠋࡑࡇ࡛
ⴭ⪅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⳦᰿⳦࡜࠸࠼ࡤࣜࢰࢡࢺ
ࢽ࢔࡛࠶ࡿ㸪࡜࠸࠺ඛධほࡀᙉ࠿ࡗࡓࡓࡵࡔࠋ
ࡑࡇ࡛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

T. Yagame - 4

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࠿࡟࡞ࡗࡓࠋ
ࡲࡓ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Ꮚᐇయࢆᙧᡂࡉࡏࡿ
ࡇ࡜࡟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

T. Yagame - 5

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࠸ࡿ
ࡢ࠿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 Corallorhiza
trifida 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࠸ࡿࣛ
ࣥ⛉࢟ࣥࣛࣥ࡟ࡘ࠸࡚ேᕤⓗ࡟ 3 ⪅ඹ⏕⣔ࢆᵓ⠏ࡍࡿࡇ࡜ࢆヨࡳࡓࠋࡲࡎ㸪࢟ࣥࣛࣥࡢ᰿ࡼ
T. Yagame - 6

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Yagame
and Yamato 2013㸧ࠋḟ࡟㸪ศ㞳⳦ᰴࢆ⏝࠸࡚ࢥࢼࣛ
㸦ࣈࢼ⛉㸧࡜እ⏕⳦᰿ࢆᙧᡂࡉࡏࡓࠋศ㞳⳦ᰴ࡜እ⏕⳦᰿ࢆᙧᡂࡋࡓࢥࢼ
ࣛࡢⱑ࡜㸪↓⳦ᇵ㣴࡛⏕⫱ࡉࡏࡓ࢟ࣥࣛࣥࡢⱑࢆ
ྠࡌ㖊ࡢ୰࡛ 120 㐌᱂ᇵࡋࡓࠋᑐ↷༊࡛ࡣ⳦᰿⳦
ࢆ᥋✀ࡋ࡚࠸࡞࠸ࢥࢼࣛ
ࡢⱑ࡜࢟ࣥࣛࣥ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☜ㄆࡉࢀࡓࠋࢥࢼࣛࡢ
ගྜᡂ⏘≀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

T. Yagame - 7

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ࢱࢩࣟࣛࣥࡢ
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植物科学最前線 5:120 （2014）㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢ
Ⓨⱆ⏕ែ࠿ࡽぢࡓ⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬ᶫᮏ 㟹ᖏᗈ␆⏘኱Ꮫ␆⏘⏕࿨⛉Ꮫ◊✲㒊㛛
ࠛ080-8555 ᖏᗈᕷ✄⏣⏫す 2 ⥺ 11 ␒ᆅ
Yasushi Hashimoto
Mycoheterotrophic germination of Pyrola asarifolia dust seeds
Key words: dust seed, forest floor, host specificity, partial mycoheterotrophy, Pyroleae
Department of Life Science and Agriculture, Obihiro University of Agriculture and Veterinary Medicine,
Inada-cho, Obihiro, Hokkaido 080-8555, Japan㸯㸬ࡣࡌࡵ࡟ ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪᳜≀࡜ࡋ࡚ࡢᰤ㣴ᦤྲྀࡢᇶ┙࡛࠶ࡿගྜᡂ⬟ຊࢆ㸪ࡍ࡭࡚࣭ࡶࡋࡃࡣ㒊ศⓗ࡟ᤞ࡚࡚㸪௚ࡢ⏕≀㸪ࡇࡇ࡛ࡣ࢝ࣅࡸࡁࡢࡇ࡜࿧ࡤࢀࡿ⳦㢮࡟౫Ꮡࡋ࡚⏕ࡁࡿ࡜࠸࠺㸪ࡎ࠸ࡪࢇᛮ࠸ษ
ࡗࡓᛶ㉁ࢆ⋓ᚓࡋࡓ᳜≀㐩࡛࠶ࡿࠋᙼࡽࡣࡑࡢ㐍໬ࡢ㐣⛬࡛㸪」ᩘࡢ᳜≀ࡢ⣔⤫࡛」ᩘᅇฟ⌧ࡋ࡚࠸ࡿࡇ࡜ࡀ▱ࡽࢀ࡚࠸ࡿࠋࡇࡢ⳦ᚑᒓᰤ㣴᳜≀ࡣ㸪᳜≀ࡢᰤ㣴ࡢᦤྲྀᶵ⬟ࡸඹ⏕⣔ࡢ㐍໬㐣⛬ࢆゎ᫂ࡍࡿୖ࡛㸪ࡲࡓ㸪᳃ᯘ⏕ែ⣔࡛ࡢ⏕≀㛫┦஫స⏝ࡢ㔜せᛶࢆ⪃࠼ࡿୖ࡛㸪㠀ᖖ࡟⯆࿡῝࠸Ꮡᅾ࡜ࡋ࡚ὀ
┠ࡉࢀ࡚࠸ࡿ㸦Whitfield 2007㸪Selosse & Roy 2009㸧ࠋࡇࡢ⳦ᚑᒓᰤ㣴᳜≀ࡢ࠺ࡕ㸪ගྜᡂ⬟ຊࢆ࠶ࡿ⛬ᗘಖᣢࡋࡓࡲࡲ㸪⳦࡬ࡢᚑᒓᰤ㣴ᛶࢆ♧ࡍ᳜≀ࡶ▱ࡽࢀ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀㸦ΰྜᰤ㣴᳜≀㸧࡜࿧ࡤࢀ࡚࠸ࡿࠋࡇࡢࡼ࠺࡞୰㛫ⓗ࡞ᛶ㉁ࢆᣢࡘ᳜≀ࡢᏑᅾࡣ㸪ᬯ࠸᳃ᯘᯘᗋ⎔ቃୗ࡛᳜≀ࡀ⳦࡬ࡢ౫Ꮡᛶࢆ⋓ᚓࡋ࡚࠸ࡃ㐍໬ࡢ㐣⛬ࡢ㸪๓㐺ᛂẁ㝵࡟ᙜࡓࡿ࡜⪃࠼ࡽࢀࡿ㸦Roy et al. 2013㸧ࠋ ࡲࡓ୍⯡࡟㸪⳦ࡶࡋࡃࡣ௚ࡢ᳜≀࡬ࡢᐤ⏕ᛶࢆ♧ࡍ᳜≀ࡣ㸪ඹ㏻ࡋࡓ≉ᚩⓗ࡞ᙧ㉁ࢆ᭷ࡋ࡚࠸ࡿࡇ࡜
ࡀ▱ࡽࢀࡿࠋࡑࡢ᭱ࡶ≉ᚩⓗ࡞ࡢࡀࠕDust seedࠖ㸦ᇕ✀Ꮚ㸧࡜࿧ࡤࢀࡿᚤ⣽✀Ꮚࡢᙧᡂ࡛࠶ࡿ㸦Eriksson &Kainulainen 2011㸧ࠋࢶࢶࢪ⛉࡟ᒓࡍࡿࢠࣥࣜࣙ࢘ࢯ࢘
ࡸࢩࣕࢡࢪࣙ࢘ࢯ࢘ࡢ௰㛫ࡀసࡿ✀Ꮚࡶ㸪ᮏ✏࡛୺࡟⤂௓ࡍࡿ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡀྵࡲࢀࡿ࢖ࢳࣖ
ࢡࢯ࢘ࡢ௰㛫ࡶ㸪ࡉࡽ࡟㸪ࢶࢶࢪ⛉࡜ࡣศ㢮Ꮫⓗ㢮⦕ᛶࡀ㐲࠸ࣛࣥ⛉᳜≀ࡶ㸪እぢୖ㦫ࡃ࡯࡝㢮ఝࡋࡓ✀Ꮚࢆసࡿ㸦ᅗ 1㸧ࠋࡇࢀࡣ㸪ᚑᒓᰤ㣴ᛶࢆᣢࡗࡓ᳜
≀࡟࠾ࡅࡿ཰ᩡ㐍໬࡜⪃࠼ࡽࢀ࡚࠸ࡿࠋ ගྜᡂ⬟ຊࢆṧࡋࡓࡲࡲ⳦࡬ࡢ౫Ꮡᛶࡶ࠶ࢃࡏᣢ
ࡘ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢ㸪ᚤ⣽✀ᏊⓎⱆ᫬ࡢඹ⏕ᅗ ྠ୍ࡢ᳃ᯘ࠿ࡽ᥇ྲྀࡋࡓᚤ⣽✀Ꮚࡢ౛ࠋᕥୖ࣭ࢩ
ࣕࢡࢪࣙ࢘ࢯ࢘ྑୖ࣭ࢡࣔ࢟ࣜࢯ࢘㸦ࣛࣥ⛉㸧ᕥୗ࣭࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ྑୗ࣭ࢥ࢖ࢳࣖࢡࢯ࢘ࠋ
ࢫࢣ࣮ࣝࡣ PP⳦࡜ࡢ㛵ࢃࡾྜ࠸ࢆ᫂ࡽ࠿࡟ࡍࡿࡇ࡜ࡣ㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬ࡀ࡝ࡢࡼ࠺࡞㐣⛬ࢆ࡬࡚㉳ࡇࡗ࡚ࡁࡓࡢ࠿ࢆ᥎ ࡍࡿ୍ຓ࡟࡞ࡿ࡜⪃࠼ࡽࢀࡿ㸦Bidartondo
Y. Hashimoto - 1

植物科学最前線 5:121 （2014）& Read 2008㸪Selosse & Roy 2009㸧ࠋࡑࡇ࡛ᮏ✏࡛ࡣ㸪ࡇࡢ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀࡜⪃࠼ࡽࢀࡿࢶࢶࢪ
⛉ࡢ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࢆྲྀࡾୖࡆ㸪ࡑࡢ✀ᏊⓎⱆࡢ㝿࡟ぢࡽࢀࡿඹ⏕⳦࡜ࡢ⯆࿡῝࠸㛵ಀᛶࢆ⤂௓ࡍࡿࠋ㸰㸬࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ ⳦ᚑᒓᰤ㣴᳜≀ࡢ௦⾲ⓗ࡞Ꮡᅾ࡜ࡋ࡚㸪ࣛࣥ⛉᳜≀࡟ᒓࡍࡿከࡃ
ࡢ✀ࡀ▱ࡽࢀ࡚࠸ࡿࡀ㸪ࢶࢶࢪ⛉ࢩࣕࢡࢪࣙ࢘ࢯ࢘ள⛉㸦Monotropoideae㸧࡟ᒓࡍࡿࢩࣕࢡࢪࣙ࢘ࢯ࢘ࡸࢠࣥࣜࣙ࢘ࢯ࢘ࡶ㸪ࡼࡃ▱ࡽࢀࡓ⳦ᚑᒓᰤ㣴᳜≀࡛࠶ࡿ㸦ᅗ2㸧ࠋ᳃ᯘࡢᯘᗋ࡟⏕⫱ࡋ㸪ࠕ⭉ⴥᅵࡢୖ࡟⏕࠼ࡿ⭉⏕᳜≀ࠖ࡞࡝࡜ᅗ㚷➼࡟グ㍕ࡉࢀ࡚ࡁࡓࡓ
ࡵ㸪ⴠⴥⴠᯞ࡞࡝ࡢศゎ≀࠿ࡽⅣỈ໬≀ࢆᚓ࡚⏕άࡋ࡚࠸ࡿ࢖࣓࣮ᅗ ࢩࣕࢡࢪࣙ࢘ࢯ࢘

ࢪࢆᣢࡓࢀ࡚࠸ࡓࡀ㸪㏆ᖺࡢDNAゎᯒࢆ⏝࠸ࡓ◊✲࡟ࡼࡗ࡚㸪ᙼࡽࡢ᰿࡟ᐃ
╔ࡋ࡚࠸ࡿ⳦㸦⳦᰿⳦㸧ࡢ✀ྠᐃࡀ࡞ࡉࢀࡓࠋࡑࡢ⤖ᯝ㸪ࡇࢀࡽࡢ᳜≀ࡀ࿘ᅖࡢඃ༨ᶞᮌ✀ࡢ᰿࡟┦฼ඹ⏕ࡋ࡚࠸ࡿࠕእ⏕⳦᰿⳦ࠖ࡜⳦᰿ࢆᙧᡂࡋ࡚࠾
ࡾ㸪ࡇࡢ⳦᰿⳦ࢆ⤒⏤ࡋ࡚ᶞᮌࡢගྜ
ᡂ⏘≀ࢆᚓ࡚࠸ࡿࡇ࡜㸪ࡲࡓ㸪ࡑࡢᐤ⏕ᑐ㇟࡜࡞ࡗ࡚࠸ࡿ⳦ࡢ✀ࡀ㸪≉␗ⓗ࡛࠶ࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸ࡿ
㸦Smith & Read 2008㸧ࠋ᭷ࡾయ࡟ゝ࠼ࡤ㸪ⴥ⥳⣲ࢆᣢࡓ࡞࠸⤯ᑐⓗ⳦ᚑᒓᰤ㣴ᛶࡢࢩࣕࢡࢪࣙ࢘ࢯ࢘㢮ࡣ㸪⮬ࡽࡢ⏕㛗࡜✀Ꮚ⏕⏘࡟ᚲせ࡞≀㉁ࡢࡍ࡭࡚ࢆ㸪᰿࡟ᐃ╔ࡋ࡚࠸ࡿዲࡳࡢࡁࡢࡇࡢ⳦⣒ᅗ ᕥ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ྑୖࢥࣂࣀ࢖ࢳࣖࢡࢯ࢘ྑୗ
ࢥ࢖ࢳࣖࢡࢯ࢘

ࢆ㣗࡭࡚ࡲ࠿࡞ࡗ࡚࠾ࡾ㸪㛫᥋ⓗ࡟࿘ᅖࡢᶞᮌ࡟ᐤ⏕ࡋ࡚⏕ࡁ࡚࠸ࡿࡇ࡜࡟࡞ࡿࠋ ࡇࢀࡽ⤯ᑐⓗ⳦ᚑᒓᰤ㣴ᛶࡢࢩࣕࢡࢪࣙ࢘ࢯ࢘㢮࡟㏆⦕࡞᳜≀࡜ࡋ࡚㸪ྠࡌࢶࢶࢪ⛉ࡢࢩࣕࢡࢪࣙ
࢘ࢯ࢘ள⛉࡟ᒓࡍࡿ࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡀ▱ࡽࢀ࡚࠸ࡿ㸦ᅗ3, 4ᕥ㸧ࠋᙼࡽࡣከᖺ⏕ࡢᖖ⥳ⲡᮏ᳜≀࡛㸪ୡ⏺ⓗ࡟30✀⛬ᗘࡀ▱ࡽࢀ࡚࠾ࡾ㸪໭༙⌫ࡢ ᖏ࠿ࡽᴟᆅ࡟࠿ࡅ࡚ศᕸࡋ㸪ᇶᮏⓗ࡟᳃ᯘෆࡸᯘ⦕࡟
⏕⫱ࡍࡿࠋࡑࡢ࠺ࡕ㸪ᮏ✏࡛୺࡟⤂௓ࡍࡿ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡣ㸪ᯘᗋ࡛ẚ㍑ⓗ኱ࡁ࡞ࢥࣟࢽ࣮ࢆ
సࡗ࡚⩌⏕ࡍࡿ᳜≀࡛࠶ࡿ㸦ᅗ3㸧ࠋ᭱ึ࡟➹⪅ࡀࡇࡢ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘࡟⯆࿡ࢆᣢࡗࡓࡢࡣ㸪ࡲࡔಟኈㄢ⛬ࡢᏛ⏕ࡢ㡭㸪ㄪᰝࡢྜ㛫࡟㸦᫬࡟ㄪᰝࡑࡗࡕࡢࡅ࡛㸧㸪㊃࿡ࡢࡁࡢࡇ⊁ࡾࢆࡋ࡚࠸ࡓ㝿࡛࠶ࡿࠋ⤒㦂ⓗ࡟ᙼࡽࡢࢥࣟࢽ࣮ࡢ୰࡛ࡣ㸪ࡁࡢࡇࡀ࠶ࡲࡾ⏕࠼࡚࠸࡞࠸ࡼ࠺࡟ឤࡌ࡚␲ၥ࡟ᛮࡗ࡚࠸ࡓ
ࡢࡔࡀ㸪ᙜ᫬ࡣ⳦᰿ࡢ◊✲ࡣึࡵ࡚࠾ࡽࡎ㸪ࡇࡢ␲ၥ࡟㛵ࢃࡿ◊✲ࢆฟ᮶ࡿ࡜ࡣᛮࡗ࡚࠸࡞࠿ࡗࡓࠋ
ࡲࡓ㸪ࡑࡢᙜ᫬ࡣࡇࢀࡽ࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡢ⳦᰿ࡸᰤ㣴ᛶ࡟ࡘ࠸࡚ࡢ◊✲ࡣ㸪࡯࡜ࢇ࡝࡞ࡉࢀ࡚࠸
࡞࠿ࡗࡓࠋࡋ࠿ࡋ㏆ᖺ㸪࡟ࢃ࠿࡟ὀ┠ࡀ㞟ࡲࡾጞࡵ㸪࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡢぶಶయࡀసࡿ⳦᰿ࡀ㸪ࢩࣕࢡࢪࣙ࢘ࢯ࢘㢮࡜ྠᵝ࡟㸪࿘ᅖࡢᶞᮌ࡜ඹ⏕ࡋ࡚࠸ࡿእ⏕⳦᰿⳦࡟ࡼࡿࡶࡢ࡛࠶ࡿࡇ࡜㸪ࡲࡓ㸪ࢩ
Y. Hashimoto - 2

植物科学最前線 5:122 （2014）ࣕࢡࢪࣙ࢘ࢯ࢘㢮࡜ࡣ␗࡞ࡾ㸪ࡑࡢእ⏕⳦᰿⳦ࡢ✀ࡀ࠶ࡲࡾ≉␗ⓗ࡛ࡣ࡞࠸ࡇ࡜ࡀ᫂ࡽ࠿࡟࡞
ࡗ࡚ࡁࡓ㸦Tedersoo et al. 2007㸪
Zimmer et al. 2007㸪Hynson &Bruns 2009㸪Hashimoto et al. 2012㸧ࠋࡉࡽ࡟㸪ᙼࡽࡢⴥࡢᏳᐃྠ఩య
13

C್࠿ࡽ⪃࠼ࡿ࡜㸪࢖ࢳࣖࢡࢯ

࢘ࡢ௰㛫ࡣ᮲௳࡟ࡼࡗ࡚㸪࠶ࡿ⛬ᗘࡢⅣỈ໬≀ࢆ⳦᰿࠿ࡽᚓ࡚
࠸ࡿ㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀࡛࠶ࡿྍ⬟ᛶࡀ㧗࠸࡜⪃࠼ࡽࢀ࡚࠸ᅗ ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢࢥࣟࢽ࣮㸦ᕥ㸧࡜⳦᰿㸦ྑୖ㸧࡜⳦᰿ࡢ
ᶓ᩿㠃㸦ྑୗ⾲⓶⣽⬊ෆ࡟⳦⣒ࡀ㐍ධࡋ࡚࠸ࡿ㸧ࡿ㸦Tedersoo et al. 2007㸪Zimmer et
al. 2007㸪Hynson & Bruns 2009㸧ࠋࡍ࡞ࢃࡕ㸪ᙼࡽࡣᚲせ࡜ࡍࡿᰤ㣴ศࡢ୍㒊ࢆ㸪᰿ࡢඹ⏕⳦ࢆ㣗࡭࡚ࡲ࠿࡞ࡗ࡚࠾ࡾ㸪㛫᥋ⓗ࡟ᶞᮌ࡟ᐤ⏕ࡋ࡚࠸ࡿࡇ࡜࡟࡞ࡿࠋࡓࡔࡋ㸪ࡇࢀࡀ࡝ࡢ⛬ᗘ࡞ࡢ࠿ࡣ᫂ࡽ࠿࡛ࡣ࡞࠸ࠋ ⴭ⪅ࡀ⌧ᅾ୺࡞ㄪᰝᆅ࡜ࡋ࡚࠸ࡿ໭ᾏ㐨ࡢᖏᗈᕷ࿘㎶࡛ࡣ㸪࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡢ࠺ࡕ㸪ࡇࡢ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡀ᭱ࡶ┠❧ࡘᏑᅾ࡛࠶ࡿࠋ࡯࠿࡟ࡶࢥࣂࣀ࢖ࢳࣖࢡࢯ࢘㸪ࢪࣥࣚ࢘࢖ࢳࣖࢡࢯ࢘㸪ࢥ࢖ࢳࣖࢡࢯ࢘࡞࡝ࡀᬑ㏻࡟ぢࡽࢀࡿ㸦ᅗ3㸪4㸪7㸧
ࠋࡇࢀࡽࡢ✀ࡣ㸪ᒣ㛫㒊ࡢᶞ㱋ࡢ㧗࠸᳃ᯘ࡛ぢ࠿ࡅࡿࡇ࡜ࡶ࠶ࡿࡀ㸪࣐࢝ࣛࢶࡸࢺࢻ࣐ࢶ࡞࡝ࡢேᕤᯘࡸ㸪ࢩࣛ࢝ࣥࣂ࡞࡝ࡢ஧ḟᯘࡢᯘᗋ࡛㸪ࡑࡇࡀࢧࢧ࡟᏶඲࡟そࢃࢀ࡚࠸࡞࠸㝿ࡢඃ༨✀࡜ࡋ࡚ぢ࠿ࡅࡿࡇ࡜ࡀከ࠸ࠋᙼࡽࡣ㸪ᬯ࠸ᯘᗋࡢ⎔ቃ࡟㐺ᛂࡋ࡚࠾ࡾ㸪ᆅୗⱼࢆ౑ࡗ࡚⏕⫱ሙᡤࢆᗈࡆ㸪᮲௳ࡀྜ࠺࡜㦫ࡃ࡯࡝኱ࡁ࡞ࢥࣟࢽ࣮ࢆᙧᡂࡋ࡚㸪ึኟ࡟ࡣぢ஦࡞ⰼࢆဏ࠿ࡏ࡚࠸ࡿ㸦ᅗ7ྑ㸧ࠋࡕ࡞ࡳ࡟㸪ࡇࢀࡽ࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡀඹ⏕⳦࡜ᙧᡂࡍࡿ⳦
᰿ࡣ㸪௚ࡢከࡃࡢⲡᮏ᳜≀ࡀసࡿࢱ࢖ࣉࡢ⳦᰿㸦࢔࣮ࣂࢫ࣮࢟ࣗࣛ⳦᰿㸧࡜ࡣ␗࡞ࡗࡓᙧ≧ࡢ⳦᰿࡛࠶ࡿ㸦Massicotte et al. 2008㸧ࠋࡑࡢ୰࡛ࡶ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡣẚ㍑ⓗࡋࡗ࠿ࡾࡋࡓ⳦㠧ࢆᣢࡘ⳦᰿ࢆᙧᡂࡍࡿࡇ࡜ࡀከ࠸㸦ᅗ4ྑୖୗ㸧ࠋ㸱㸬ᚤ⣽✀Ꮚ ⳦ᚑᒓᰤ㣴᳜≀ࡢࡘࡃࡿᚤ⣽✀Ꮚࡣ㸪ᩘ༑ಶࡢ⣽⬊࠿ࡽ࡞ࡿᑠࡉ࡞⬇࡜✀⓶ࡢࡳ࡛ᵓᡂࡉࢀ㸪Ⓨⱆࡢ㝿ࡢᰤ㣴※࡜࡞ࡿᏊⴥ࠶ࡿ࠸ࡣ⬇ஙࢆᣢࡗ࡚࠸࡞࠸ࡇ࡜ࡀ≉ᚩ࡜࡞ࡿ㸦ᅗ 1㸧ࠋࡑࡢࡓࡵ㸪ᑡ࡞ࡃ࡜
ࡶⓎⱆࡢึᮇࡣ㸪እ㒊࠿ࡽᰤ㣴※ࢆ☜ಖࡋ࡞ࡅࢀࡤ㸪ࡑࡢⓎⱆ⏕㛗ࡢ㐣⛬ࢆ࠺ࡲࡃ㐍ࡵࡿࡇ࡜ࡀྔࢃ
࡞࠸ࠋࡑࡋ࡚㸪ぶಶయࡀ⥳ࡢⴥࢆᣢࡕ⮬ࡽගྜᡂࢆ⾜ࡗ࡚࠸ࡿ᳜≀✀࡛࠶ࡗࡓ࡜ࡋ࡚ࡶ㸪ᚤ⣽✀Ꮚࢆసࡿ᳜≀ࡣ㸪Ⓨⱆ࠿ࡽⴥࢆ᏶ᡂࡍࡿࡲ࡛ࡢᗂಶయࡢ㛫ࡣ㸪⤯ᑐⓗ࡞⳦ᚑᒓᰤ㣴᳜≀࡜ࡋ࡚⏕άࡋ࡚࠸ࡿࡇ࡜࡟࡞ࡿࠋ ࡇࡢᚤ⣽✀Ꮚࡣ㸪ᰤ㣴※ࢆእ㒊࡟㢗ࡗ࡚࠸ࡿࡇ࡜࠿ࡽ㸪ಶࠎࡢ✀ᏊࡢⓎⱆᡂຌ⋡ࡀప࠸࡜ண᝿ࡉࢀ
ࡿࠋࡋ࠿ࡋぢ᪉ࢆኚ࠼ࡿ࡜㸪ᑗ᮶࡟ࢃࡓࡗ࡚ᰤ㣴ࢆ౫Ꮡࡍࡿᐤ⏕ᑐ㇟ࢆ㸪ึᮇࡢ⏕㛗㛤ጞ᫬Ⅼ࡛㸪☜ᐇ࡟☜ಖࡋ࡚࠸ࡿ࡜ࡶ⪃࠼ࡽࢀࡿࠋࡉࡽ࡟㸪ᚤ⣽✀Ꮚࡢ⏕⏘ࡣ㸪㸯⢏࠶ࡓࡾࡢࢥࢫࢺࡀ᭱ᑠ㝈࡟ᢚ࠼ࡽࢀࡿࡇ࡜࠿ࡽ㸪ྠࡌࢥࢫࢺ࡛኱㔞ࡢ✀Ꮚࡢ⏕⏘ࡀྍ⬟࡜࡞ࡾ㸪ᩘ༓࠿ࡽ୓༢఩ࡢ✀Ꮚࡀぶ᳜≀㸯ಶY. Hashimoto - 3

植物科学最前線 5:123 （2014）య࠿ࡽ⏕⏘ࡉࢀࡿࠋࡇࢀࡽࡢᚤ⣽✀Ꮚࡀぶ࠿ࡽᨺฟࡉࢀࡓᚋࡣ㸪ࡑࡢ㍍ࡉࢆ⏕࠿ࡋ࡚㸪㢼ࡸỈࡢὶࢀ࡟஌ࡗ࡚ᗈࡃศᩓࡍࡿࡇ࡜ࡀྍ⬟࡛࠶ࡿࠋࡲࡓ㸪ࡇࢀࡽࡢ✀Ꮚࡣ㸪ᅵተ୰࡛ࢩ࣮ࢻࣂࣥࢡ࡜ࡋ࡚ẚ㍑ⓗ㛗ᮇ࡟Ⓨⱆࡢᶵ఍ࢆᚅࡘࡇ࡜ࡀྍ⬟࡛࠶ࡿ࡜⪃࠼ࡽࢀࡿࠋࡑࡢࡓࡵ㸪᳜≀✀࡜ࡋ࡚ࡢ༑ศ࡞Ⓨⱆᡂຌ࡜㸪ศᕸᣑ኱ࡢྍ⬟ᛶࡀᢸಖࡉࢀ࡚࠸ࡿ࡜ぢࡽࢀࡿ㸦Bruns & Read 2000㸧
ࠋࡇࡢⓎⱆࡢᡂຌࡣࡑࡢᰤ
㣴※ࡢ☜ಖࡢᡂྰ࡟࠿࠿ࡗ࡚࠸ࡿ࡜⪃࠼ࡽࢀ㸪⳦ᚑᒓᰤ㣴᳜≀࡛࠶ࢀࡤ㐺ษ࡞ඹ⏕⳦ࡢ☜ಖࢆព࿡ࡍ
ࡿࠋ㸲㸬ᚤ⣽✀ᏊࡢⓎⱆ᫬ࡢඹ⏕⳦ ᚤ⣽✀Ꮚࡀぶ࠿ࡽᨺࡓࢀ࡚᭱⤊ⓗ࡟ⴠࡕࡿ᳃ᯘࡸࡑࡢ࿘㎶ࡢᅵተ୰࡟ࡣ㸪ᩘከࡃࡢከᵝ࡞⳦㢮ࡀ⏕ᜥࡋ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿࠋࡇࢀࡽ⳦ᚑᒓᰤ㣴᳜≀ࡢ✀ᏊࡢⓎⱆ࡛ࡣ㸪ࡑࡢ᳜≀✀ࡈ࡜࡟ࡇࡢከᵝ࡞⳦ࡢ୰࠿ࡽ㸪࠶ࡿ≉ᐃࡢ⳦ࢆ✀࣭ᒓ࣭⛉࡞࡝ࡢ࡛ࣞ࣋ࣝ㑅ࢇ࡛ඹ⏕⳦࡜ࡋ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿ㸦Smith& Read 2008㸧ࠋࡍ࡞ࢃࡕ㸪ࡑࡢ✀Ꮚࡀⴠࡕࡿሙᡤ࡟┦ᛶࡢⰋ࠸⳦ࡀᏑᅾࡍࡿࡇ࡜ࡀ㸪Ⓨⱆ࠿ࡽᐃ╔ᡂຌࡢᚲ㡲᮲௳࡟࡞ࡿࠋ ᚤ⣽✀ᏊⓎⱆࡢ㝿ࡢඹ⏕⳦ࢆ᫂ࡽ࠿࡟ࡍࡿࡓࡵࡢⓎⱆᐇ㦂ࡣ㸪୺࡟ࣛࣥ⛉᳜≀ࡸࢩࣕࢡࢪࣙ࢘ࢯ࢘
ࡢ௰㛫ࡢ✀Ꮚࢆᑐ㇟࡟⾜ࢃࢀ࡚࠾ࡾ㸪≉࡟ࣛࣥ⛉᳜≀࡟㛵ࡋ࡚ࡣ㸪᳜≀⑓ཎ⳦࡜ࡋ࡚▱ࡽࢀࡿ୙᏶඲
⳦㢮ࡢࣜࢰࢡࢺࢽ࢔ᒓ⳦ࡀ㸪Ⓨⱆࡢ㝿ࡢඹ⏕⳦࡜ࡋ࡚ẚ㍑ⓗ᪩ࡃ࠿ࡽ▱ࡽࢀ࡚࠸ࡿࠋ
ࡋ࠿ࡋ㏆ᖺ࡛ࡣ㸪ࡑࡢࣜࢰࢡࢺࢽ࢔ᒓ⳦ࡀ」ᩘࡢ⣔⤫࡟ࡲࡓࡀࡗࡓ㞟ྜయ࡛࠶ࡿࡇ࡜ࡸ㸪ࣜࢰࢡࢺࢽ࢔ᒓ⳦࡟ࡣྵࡲࢀ
࡞࠸⳦ࡶࣛࣥࡢඹ⏕⳦࡛࠶ࡿࡇ࡜ࡀ᫂ࡽ࠿࡜࡞ࡾ㸪ࡉࡽ࡟㸪Ⓨⱆ᫬ࡢඹ⏕⳦ࡶከᵝ࡛࠶ࡿࡇ࡜ࡀุ᫂
ࡋ࡚ࡁ࡚࠸ࡿ㸦Smith & Read 2008, Yukawa et al. 2009㸧ࠋࡲࡓࣛࣥࡢሙྜ㸪Ⓨⱆࡢ㝿ࡢඹ⏕⳦ࡣ㸪ぶಶయ࠿ࡽ᳨ฟࡉࢀࡿ⳦࡜୍⮴ࡋ࡚࠸ࡿࡇ࡜ࡀከ࠸ࡼ࠺࡛࠶ࡿ㸦኱࿴࣭㇂ட 2009㸧ࠋࢩࣕࢡࢪࣙ࢘ࢯ࢘ࡢ௰
㛫࡛ࡶ㸪Ⓨⱆ࠿ࡽぶಶయࡢ⳦᰿ࡲ࡛㸪ྠࡌ✀㢮ࡢ⳦ࡀࡘ࠸࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿ㸦Bruns & Read 2000㸪
Leake et al. 2004㸪Yamada et al. 2008㸧ࠋ ࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡢ✀ᏊⓎⱆ࡟㛵ࡍࡿ◊✲ࡣ㸪༙ୡ⣖๓ࡢྂ඾ⓗ࡞◊✲ࡢ౛ࡀ࠶ࡿࡔࡅ࡛㸪ࡈࡃ᭱㏆ࡲ࡛࡯࡜ࢇ࡝ሗ࿌ࡀ࡞ࡉࢀ࡚࠸࡞࠿ࡗࡓࠋࡇࢀࡣ㸪◊✲ࡀ⾜ࢃࢀ࡚࠸࡞࠿ࡗࡓヂ࡛ࡣ࡞ࡃ㸪ࡑࡢⓎⱆࢆぢ࠸ࡔࡍࡇ࡜ࡀ㸪࡞࠿࡞࠿ฟ᮶࡞࠿ࡗࡓࡇ࡜ࡀཎᅉ࡛࠶ࡿ࡜⪃࠼ࡽࢀࡿ㸦Smith & Read 2008㸪
Hynson et al. 2009㸧ࠋࡑࡋ࡚ࡇࡢⓎⱆࡢᕼᑡᛶࡀ㸪࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡢ⏕ែ࡟㸪࡝ࡢࡼ࠺࡟㛵ࢃࡗ࡚࠸ࡿࡢ࠿⯆࿡ࡀᣢࡓࢀ࡚ࡁࡓࠋ≉࡟㸪ࡇࢀࡽ࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡣ㸪⤯ᑐⓗ࡞⳦ᚑᒓᰤ㣴ᛶࡢࢩࣕࢡ
ࢪࣙ࢘ࢯ࢘ࡢ௰㛫࡜㠀ᖖ࡟㏆⦕࡛࠶ࡿࡓࡵ㸪ࡇࢀࡽࡢ᳜≀ࡀࡑࡢ㐍໬ࡢ㐣⛬࡛࡝ࡢࡼ࠺࡟⳦ᚑᒓᰤ㣴ᛶࢆ⋓ᚓࡋ࡚ࡁࡓࡢ࠿ࢆ⪃࠼ࡿୖ࡛㸪ᚲせ࡞᝟ሗ࡛࠶ࡿ࡜⪃࠼ࡽࢀࡿࠋࡉࡽ࡟㸪๓㏙ࡢࡼ࠺࡟࢖ࢳࣖ
ࢡࢯ࢘ࡢ௰㛫ࡢぶಶయࡀసࡿ⳦᰿ࡣ㸪እ⏕⳦᰿⳦࡟ࡼࡿࡶࡢ࡛㸪ࡑࡢ⳦࡟≉␗ᛶࡀ࡞࠸࡜ࡢሗ࿌ࡀ࡞
ࡉࢀ࡚࠸ࡿࠋࡑࡢࡓࡵ㸪ࡇࢀࡽ࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡢⓎⱆࡣ㸪ࣛࣥ⛉ࡸࢩࣕࢡࢪࣙ࢘ࢯ࢘㢮࡞࡝࡜ྠࡌࡼ࠺࡟㸪࠶ࡿ⛬ᗘ≉␗ⓗ࡞⳦ࡀඹ⏕⳦࡜࡞ࡗ࡚࠸ࡿࡢ࠿㸪ࡶࡋࡃࡣぶಶయࡢ⳦᰿࡜ྠࡌࡃ㸪ከᵝ࡞⳦࡜ඹ⏕㛵ಀࢆ⤖ࢇ࡛Ⓨⱆࡍࡿࡇ࡜ࡀྍ⬟࡞ࡢ࠿㸪୍ᒙ⯆࿡ࡀࡶࡓࢀࡿࡼ࠺࡟࡞ࡗࡓࠋࡇࡢࡼ࠺࡞⫼ᬒ࠿ࡽ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࢆᑐ㇟࡟㸪ᙼࡽࡢ✀Ꮚࡀ࡝ࡢࡼ࠺࡞᮲௳࡛Ⓨⱆࡋ㸪࡝ࡢࡼ࠺࡞⳦ࡀⓎⱆࡢ㝿ࡢඹ⏕⳦࡜࡞ࡗ࡚࠸ࡿࡢ࠿ࢆゎ᫂ࡍࡿ࡭ࡃㄪᰝࢆ⾜ࡗࡓ㸦Hashimoto et al. 2012㸧ࠋ

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植物科学最前線 5:124 （2014）㸳㸬࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢⓎⱆㄪᰝ ᚤ⣽✀Ꮚࢆసࡿ᳜≀ࡢⓎⱆࢆᑐ㇟࡜ࡋࡓㄪᰝࡣ㸪㠀ᖖ࡟┠ࡢ⣽࠿࡞ࢼ࢖ࣟࣥࢿࢵࢺ࡞࡝࡟✀Ꮚࢆධࢀ࡚┠ᫍࢆ௜ࡅࡓሙᡤࡢᅵተ୰
࡟ᇙࡵ࡚㸪୍ᐃᮇ㛫ᚋ࡟᥀ࡾฟࡍ᪉ἲࡀ⏝࠸ࡽࢀ࡚࠸ࡿ㸦Rasmussen& Whigham 1998, ㎷⏣࣭㐟ᕝ 2008㸧ࠋ᭱ࡶ୍⯡ⓗ࡞ࡢࡣ㸪ࣇ࢕࣒ࣝ෗┿ࡢࢫࣛ࢖ࢻ⏝࣐࢘ࣥࢺ࡟✀Ꮚධࡾࢿࢵࢺࢆᤄࢇ࡛సࡗࡓ✀Ꮚࣃࢵࢡࢆᅵተ࡟ᇙࡵࡿ᪉ἲ࡛࠶ࡿ㸦ᅗ 5㸧ࠋࡇࢀࡽࡢ᪉ἲࡣ㸪ࣛࣥࡸࢩࣕࢡࢪࣙ࢘ࢯ࢘ࡢ௰㛫ࡢ✀Ꮚࢆᑐ㇟࡟ᡂᯝࢆ࠶ࡆ㸪Ⓨⱆ࡟࠿࠿ࡿ᫬㛫ࡸ㸪ᚲせ࡞᮲௳㸪≉࡟ඹ⏕⳦ࡢ✀ࢆ▱ࡿࡢ࡟᭷ຠ࡛࠶ࡿࠋ ࡇࡢᡭἲࢆ⏝࠸࡚㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢ✀ᏊࡢⓎⱆㄪᰝࢆ㛤ጞࡋࡓࠋࡣࡌࡵࡣ᭱ࡶࡓࡃࡉࢇࡢ✀ᏊࡀᏑᅾࡍࡿ࡜⪃࠼ࡽࢀࡿぶ᳜≀ࡀከࡃぢࡽࢀࡿ᳃ᯘࡢࢥࣟࢽ࣮ෆࡸ㸪ࡑࡢ࿘㎶࡟✀Ꮚࣃࢵࢡࢆᇙ
ࡵ࡚ᵝᏊࢆぢࡿࡇ࡜࡟ࡋࡓࠋࡋ࠿ࡋ㸪᥀ࡾฟࡋࡓࣃࢵࢡ࠿ࡽࡣ඲ࡃ
Ⓨⱆࡣぢࡽࢀ࡞࠿ࡗࡓࠋࡑࢀ௨㝆㸪ᇙࡵࡿሙᡤ㸪῝ࡉ㸪ᮇ㛫㸪ඖࠎᅗ ᚤ⣽✀Ꮚࡢᇙタᐇ㦂࡟⏝࠸ࡓ✀Ꮚࣃࢵࢡ࡜㸦ྑ㸧㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡ
ࢯ࢘✀Ꮚ㸦ᕥ㸧㻌ࡢ✀Ꮚࡢ㉁࡞࡝㸪⪃࠼ࡽࢀࡿ࠸ࢁ࠸ࢁ࡞せᅉࢆ₽ࡍࡼ࠺࡟㸪࠶ࡕࡇࡕ࡟✀Ꮚࣃࢵࢡࢆᇙࡵ࡚᥀ࡾฟࡍ࡜࠸࠺⾜Ⅽࢆ⧞ࡾ㏉ࡋࡓࠋᇙࡵࡓሙᡤࡣ㸪ᶞ✀ࡢ㐪࠺ྛ᳃ᯘෆࡢ࣋
ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢࢥࣟࢽ࣮ෆ㒊ࡸࡑࡢ࿘㎶㸪ᯘ⦕㸪᳃ᯘ࡟㠃ࡋࡓ᧠஘ᆅ㸪∾ⲡᆅ࡞࡝㸪࿘㎶࡟࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡀ⏕⫱ࡋ࡚࠾ࡾ㸪ᙼࡽࡢᑠࡉ࡞✀Ꮚࡀ㣕ࡪྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿሙᡤࢆᵝࠎヨࡋ࡚ࡳࡓ㸦⾲ 1㸪ᅗ 7㸧ࠋ ᭱ึ࡟Ⓨⱆࢆほᐹฟ᮶ࡓࡢࡣ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢ⩌ⴠෆ࡛ࡶ㸪᳃ᯘෆ࡛ࡶ࡞ࡃ㸪኱Ꮫࡢᰯ⯋ࡢᒇୖ࡟ᨺ⨨ࡋࡓ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘࡜ࢩࣛ࢝ࣥࣂࡢⱑᮌࢆ᳜࠼ࡓ௚ࡢᐇ㦂⏝࡟సࡗࡓࣉࣛࣥࢱ
࣮ࡢᅵࡢ୰࡟ணഛⓗ࡟ᇙࡵࡓࣃࢵࢡ࠿ࡽ
࡛࠶ࡗࡓࠋࡇࢀࡀ 2 ᖺ┠ࡢฟ᮶஦࡛㸪௨㝆㸪ࣃࢵࢡࡢᇙタሙᡤࢆ㸪Ⓨⱆࡀほᐹࡉࢀࡓࣉ
ࣛࣥࢱ࣮࡜᮲௳ࡀఝ࡚࠸ࡿ࡜ᛮࢃࢀࡿ㸪ࢩࣛ࢝ࣥࣂᐇ⏕ࡢ⏕࠼ࡓఆ᥇㊧ᆅ㸪㐨㊰⬥ࡢἲ㠃ࡸ㸪࠿ࡃ஘ࢆཷࡅࡓᯘ⦕㒊࡞࡝㸪ᛮ࠸ࡘࡃሙᡤ࠶ࡕࡇࡕ࡟ᇙࡵ࡚ᅇࡗࡓࠋࡑࢀ࡛
ࡶ㸪ࡑࡢᚋ㸰ᖺ㛫඲ࡃⓎⱆࡣぢࡽࢀࡎ㸪᭱ᚋ࡜Ỵࡵ࡚ᮃࢇࡔ 5 ᖺ┠࡟㸪ࡸࡗ࡜ 3 ࢝ᡤࡢⱝ㱋ࡢ࢝ࣂࣀ࢟஧ḟᯘ࡛㸪ࡲ࡜ࡲࡗࡓⓎⱆࢆほᐹࡍࡿࡇ࡜ࡀฟ᮶ࡓࠋ⤖ᒁ㸪Ⓨⱆࡀほᐹࡉࢀࡓࡢࡣ㸪ࡣࡌࡵ࡟Ⓨⱆࢆほᐹࡋࡓࢩࣛ࢝ࣥࣂⱑᮌ࡜࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘
ࢆ࡜ࡶ࡟᳜࠼ࡓࣉࣛࣥࢱ࣮ࡢ 1 ࡘ࡜㸪ᶞ㱋ࡀ⣙ 15 ᖺ⛬ᗘ࡜᥎ ࡉࢀࡿࢩࣛ࢝ࣥࣂࡶᅗ ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢⓎⱆ✀ᏊࠋᕥୖⓎⱆࡢጞࡲࡗࡓ✀Ꮚ࡜ᮍⓎⱆ✀Ꮚ㸦EDU㸸PP⾲㠃࡟ぢ࠼ࡿ㯮Ⰽ⳦⣒ࡣⓎⱆඹ⏕⳦ࡢࡶࡢ࡛ࡣ࡞ࡃᅵተ୰ࡢ⭉⏕ᛶࡢ⳦ࡢ⳦⣒㸧ࠋྑᮍⓎⱆ✀Ꮚ࡜Ⓨⱆẁ㝵ࡢ␗࡞ࡿⓎⱆ✀Ꮚ㸦EDU㸸PP㸧
ࠋᕥୗᰁⰍࡋࡓⓎⱆ✀Ꮚ⣽⬊ෆࡢ⳦⣒㸦EDU㸸wP㸧ࡋࡃࡣࢲࢣ࢝ࣥࣂࡢⱝ㱋ᯘࡢ 3 ࢝ᡤࡔࡅ࡛࠶ࡿࠋࡑࢀ௨እࡢᇙタሙᡤ࠿ࡽࡣ㸪᫂☜࡟Ⓨⱆࡋ࡚࠸ࡿ࡜
ぢ࡞ࡏࡿ✀⓶ࢆ✺ࡁᢤࡅ࡚⏕㛗ࡋ࡚࠸ࡿᐇ⏕ࡣほᐹࡉࢀ࡞࠿ࡗࡓ㸦ᅗ 6㸧ࠋᇙタࡋࡓ✀Ꮚࣃࢵࢡࡣ㸪ண
Y. Hashimoto - 5

植物科学最前線 5:125 （2014）ഛⓗ࡟ᇙࡵࡓࡶࡢࢆ㝖࠸࡚ྜィ 990 ࣃࢵࢡ࡛㸪
ࡑࡢ࠺ࡕⓎⱆࡀぢࡽࢀࡓࡢࡣ 74 ࣃࢵࢡ࡛࠶ࡗࡓ
㸦⾲ 1㸧ࠋ
਀1 ঋॽংॼॖॳখॡ९क़ர৕॑োोञঃॵॡभඇਝৃਚध໧ॉলखञர৕ঃॵॡभ৅࿈૨
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50

0ॺॻঐॶযੵ௔

৺40 ফেॺॻঐॶ

ቬాपে୘‫ق‬॥টॽ‫ش‬஄ਛ‫ك‬

50

0३ছढ़থংযੵ௔

৺15 ফে३ছढ़থং

ऽयैपে୘

150

45 (2.4)

ฝൃ৉

ऩख

ऩख

20

0

௔໿भ࿿એ

ऩख

ऩख

20

0

ᅇ఼඄৉

ऩख

ऩख

20

0

௔෍भ࿿એ

৺3 ফেॲॣढ़থং

ऩख

60

0

ᅇ఼඄৉

৺5 ফেॲॣढ़থং

ऩख

60

0

௔෍भ࿿એ

৺5 ফেॺॻঐॶध॔ढ़ग़॰ঐॶ

ቬాपে୘‫ق‬৵औऩ॥টॽ‫ش‬஄ਛ‫ك‬

20

0

௔෍भ࿿એ

৺5 ফেॺॻঐॶध॔ढ़ग़॰ঐॶ

ቬాपে୘‫ق‬৵औऩ॥টॽ‫ش‬஄ਛ‫ك‬

100௔੊෍भ૬ऌ৉

৺3 ফে३ছढ़থং

ऩख

60

0

௔੊෍भ૬ऌ৉

৺3 ফেॲॣढ़থং

ऩख

60

0

३ছढ़থং੸ઃ௔

৺15 ফে३ছढ़থং

ऽयैपে୘

75

16 (1.0)ॲॣढ़থং੸ઃ௔

৺15 ফেॲॣढ़থংध঑খঐঁথঀय़

ऽयैपে୘

75

10 (2.1)

ৰୡ৷উছথॱ‫ش‬

3 ফেढ़ছঐॶ၈਽

ढ़ছঐॶ௔ऊै఼਄‫ڱعگ‬ট८ॵॺ

50ৰୡ৷উছথॱ‫ش‬

3 ফে३ছढ़থং၈਽

ढ़ছঐॶ௔ऊै఼਄‫ڱعگ‬ট८ॵॺ5

3 (11.9)* ඇਝखञர৕ঃॵॡभअठ৅࿈ऋৄैोञঃॵॡभસ়‫؛‬ถ಍मHashimoto et al. 2012 ૞ස ࡇࡢ⤖ᯝ࠿ࡽ⪃࠼ࡽࢀࡿࡢࡣ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢⓎⱆࡣ㸪ᅵተ୰࡛ᑡ࡞ࡃ࡜ࡶ 5 ࣨ᭶⛬ᗘ࠶ࢀࡤ㉳ࡇࡿྍ⬟ᛶࡀ࠶ࡿࡇ࡜㸪ᅵተ୰࡛෤ᮇࢆ㉺ࡍᚲせࡣ࡞࠸ࡇ࡜㸪ࡲࡓ㸪ᅵተ୰࡛ࡢⓎⱆࡢ㝿࡟㸪࢝ࣥࣂࡢ௰㛫ࡢ≉࡟ⱝ㱋ࡢಶయࡢᏑᅾ
࡜㸪ᡂ⇍ᯘ࡯࡝࡛ࡣ࡞࠸ࡀ࠶ࡿ⛬ᗘ᭷ᶵ≀ࡢሁ✚ࡋࡓᅵተࡢᏑᅾࡀᚲせࡶࡋࡃࡣఱࡽ࠿ࡢᘬࡁ㔠࡟࡞ࡿࡇ࡜࡛࠶ࡗࡓࠋ
≉࡟⯆࿡῝࠸ࡢࡣ㸪ぶಶయࡀ᪲┒࡟⏕⫱ࡋ࡚኱ࡁ࡞ࢥࣟࢽ࣮ࢆసࡾ㸪ከࡃࡢ✀Ꮚࡀᙧᡂࡉࢀ࡚࠸ࡿሙᡤ࡛ࡶ㸪඲ࡃⓎⱆࢆほᐹฟ᮶࡞࠿ࡗࡓࡇ࡜࡛࠶ࡿ㸦ᅗ 7㸧ࠋࣃࢵࢡ࡟ධࢀࡓ✀Ꮚࡣࡇࢀࡽࡢሙᡤ࠿
ࡽ᥇ྲྀࡋ࡚࠸ࡿ࡟ࡶ㛵ࢃࡽࡎ㸪࡛࠶ࡿࠋᅗ ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘✀Ꮚࣃࢵࢡᇙタᆅࠋᕥ✀ᏊⓎⱆࡀぢࡽࢀࡓⱝ㱋ࢩࣛ࢝ࣥࣂ஧ḟᯘࠋྑ✀ᏊⓎⱆࡀぢࡽࢀ࡞࠿ࡗࡓࢺࢻ࣐
ࢶேᕤᯘࡢ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢࢥࣟࢽ࣮㸦ⰼᮇ㸧ࡲࡓ㸪࣐࢝ࣛࢶᯘ㸪ࢺࢻ࣐ࢶᯘࡔࡅ࡛࡞ࡃ㸪ࢩࣛ࢝ࣥࣂᡂ⇍ᯘࡢ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢࢥࣟࢽ࣮ෆ
࡛ࡶ㸪✀Ꮚࣃࢵࢡࢆ 50 ಶᇙࡵࡓࡀ㸪Ⓨⱆࡣࡲࡗࡓࡃほᐹࡉࢀ࡞࠿ࡗࡓࠋࡉࡽ࡟㸪Ⓨⱆࡀぢࡽࢀࡓࢩࣛ
࢝ࣥࣂⱝ㱋ᯘෆ࡛ࡢ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ぶಶయࡢ⏕⫱ሙᡤ࡜㸪Ⓨⱆࡀぢࡽࢀࡓ✀Ꮚࣃࢵࢡࡢᇙタሙᡤࡢ㛫࡟ព࿡ࡢ࠶ࡿ㛵ಀᛶࡣ࡞ࡃ㸪ᯘෆ࡛ࣞ࣋ࣝࡶⓎⱆࡢ㉳ࡇࡿせᅉ࡜ࡋ࡚ࡢぶಶయࡢᏑᅾࡣᚲせ࡞᮲௳࡜ࡣゝ࠼࡞࠸ࡼ࠺ࡔࡗࡓࠋࡘࡲࡾ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢᚤ⣽✀ᏊࡢⓎⱆࡣ㸪ぶࡢᏑᅾ㸪ࡶࡋࡃࡣぶࡢ᰿࡟ࡘ࠸࡚࠸ࡿ⳦ࡢᏑᅾ࡜ࡣ㛵ಀࡀ࡞ࡃ㉳ࡁ࡚࠸ࡿ࡜࠸࠺ࡇ࡜࡟࡞ࡿࠋࡇࢀࡣ㸪ᙼࡽ࡟㏆⦕࡞⤯ᑐⓗ⳦ᚑᒓᰤ㣴᳜≀ࡢࢩࣕࢡࢪࣙ࢘ࢯ࢘ࡢ௰㛫࡛▱ࡽࢀ࡚࠸ࡿⓎⱆᐇ㦂ࡢ⤖ᯝ࡜ࡣ␗࡞ࡿࡶࡢ࡛
࠶ࡗࡓ㸦Leake et al. 2004㸧
ࠋ
Y. Hashimoto - 6

植物科学最前線 5:126 （2014）㸴㸬࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘✀ᏊࡢⓎⱆඹ⏕⳦ ࡑࢀ࡛ࡣ㸪௒ᅇぢࡽࢀࡓⓎⱆ✀Ꮚࡢඹ⏕⳦ࡣ࠸࠿࡞ࡿ✀㢮࡛࠶ࢁ࠺࠿㸦ᅗ 6 ᕥୗ㸧ࠋⓎⱆࡀほᐹࡉࢀ
ࡓࣉࣛࣥࢱ࣮࡜㸪㸱࢝ᡤࡢⱝ㱋࢝ࣂࣀ࢟ᯘ࠿ࡽᚓࡽࢀࡓⓎⱆ✀Ꮚ࠿ࡽ DNA ࢆᢳฟࡋ࡚㸪Ⓨⱆ࡟㛵ࢃࡗ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿ⳦ࡢ rDNA ࡢ ITS 㡿ᇦ࡜ 28S ࡢ୍㒊ࢆቑᖜࡋ࡚㸪ࡑࡢ⳦ࡢ✀ྠᐃࢆヨࡳࡓࠋࡑࡢ⤖ᯝ㸪ࣛࣥࢲ࣒࡟㑅ࢇࡔ 34 ࡢⓎⱆᐇ⏕ࡢࡍ࡭࡚࠿ࡽ㸪ᢸᏊ⳦㢮ࡢࣟ࢘ࢱࢣ⛉㸦Sebacinaceae㸧ࣟ࢘ࢱࢣᒓ㸦Sebacina㸧ࡢ Sebacina vermifera ࡟ࡈࡃ㏆⦕ࡢ⳦ࡔࡅࡀ᳨ฟࡉࢀ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢⓎⱆࡢඹ⏕⳦ࡀ≉␗ⓗ࡛࠶ࡿྍ⬟ᛶࡀ♧ࡉࢀࡓ㸦Hashimoto et al. 2012㸧ࠋ≉࡟㸪㸱࢝ᡤࡢⱝ㱋࢝ࣂࣀ࢟ᯘ
ࡢෆࡢ 1 ࢝ᡤࡣ㸪௚ࡢ 2 ࢝ᡤ࡜⣙ 90km 㞳ࢀࡓ࡜ࡇࢁ࡟఩⨨ࡋ࡚࠾ࡾ㸪ࡲࡓ㸪ᚋ᪥㏣ຍ࡛✀Ꮚࣃࢵࢡࢆᇙࡵ㸪᪂ࡓ࡟᥇ྲྀࡋࡓⓎⱆᐇ⏕ࡢ☜ㄆࢆ⾜ࡗ࡚ࡶ㸪౪ヨࡋࡓࡍ࡭࡚ࡢᐇ⏕࠿ࡽྠࡌ⳦ࡢ DNA ࡀ᳨ฟࡉࢀࡓࡓࡵ㸪ࡇࡢ⳦ࡀ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢⓎⱆ࡟ᙉࡃ㛵ࢃࡗ࡚࠸ࡿ⳦࡛࠶ࡿࡇ࡜ࡣ㛫㐪࠸࡞࠸࡜⪃࠼ࡽࢀࡿࠋࡉ࡚㸪ࡇࡢࣟ࢘ࢱࢣᒓࡢ⳦㸦ࡇࢀࡽࡢ⳦ࡣศ㢮Ꮫⓗぢ┤ࡋࡀ⾜ࢃࢀ࡚࠾ࡾ㸪ᑗ᮶ⓗ࡟✀ྡࡸᒓྡࡢኚ᭦ࡀ⾜ࢃࢀࡿྍ⬟ᛶࡀ㧗࠸㸧ࡀྵࡲࢀࡿࡢࡣ㸪ࣟ࢘ࢱࢣࡢ௰㛫ࡢ୰࡛ࡶ㸪୺࡟ᶞᮌࡢ
᰿࡜ඹ⏕ࡍࡿእ⏕⳦᰿⳦࡜ࡋ࡚ぢࡘ࠿ࡿࢢ࣮ࣝࣉ࡛ࡣ࡞ࡃ㸪⭉⏕⳦㸦ศゎ⳦㸧ࡸ᳜≀ࡢෆ⏕⳦࡜ࡋ࡚ぢࡘ࠿ࡿࢢ࣮ࣝࣉ㸦Sebacinales ࡢ clade B㸧ࡢ⳦࡛࠶ࡗࡓࠋࡍ࡞ࢃࡕ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡣ㸪⏕㛗ࡋࡓᚋ࡟ඹ⏕ࡋ࡚࠸ࡿእ⏕⳦᰿⳦࡜ࡣ␗࡞ࡿ⏕άᵝᘧࡢ⳦ࢆ㸪Ⓨⱆࡢ㝿ࡢඹ⏕⳦࡜ࡋ࡚࠸ࡿࡇ࡜࡟࡞
ࡿࠋࡲࡓ㸪ࡉࡽ࡟⯆࿡῝࠸ࡢࡣ㸪௒ᅇ≉␗ⓗ࡟Ⓨⱆᐇ⏕࠿ࡽ᳨ฟࡉࢀࡓࣟ࢘ࢱࢣᒓࡢ⳦ࡣ㸪୺࡟࣮࢜ࢫࢺࣛࣜ࢔࡛⮬⏕ࡍࡿ 1 ࢢ࣮ࣝࣉࡢࣛࣥ㸦Diuris 㐃㸧ࡢ✀ᏊⓎⱆࡢඹ⏕⳦࡜ࡋ࡚▱ࡽࢀ࡚࠸ࡿ⳦ࡔࡗ
ࡓࡇ࡜࡛࠶ࡿ㸦Warcup 1971㸪Yagame & Yamato 2008㸧ࠋ࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡣ໭༙⌫ࡢࡳ࡟⏕⫱ࡍࡿ᳜≀࡛࠶ࡾ㸪ᆅ⌮ⓗ࡟㠀ᖖ࡟㞳ࢀࡓሙᡤ࡛␗࡞ࡿ⳦ᚑᒓᰤ㣴᳜≀ࡀ㸪ྠࡌ⭉⏕ᛶࡢࣟ࢘ࢱࢣᒓࡢ⳦ࢆ
Ⓨⱆඹ⏕⳦࡟㑅ࡪ㸪཰ᩡ㐍໬ࡀ㉳ࡇࡗࡓྍ⬟ᛶࡶ⪃࠼ࡽࢀࡿࠋ࡞ࡐ≉ᐃࡢ⭉⏕ᛶࣟ࢘ࢱࢣᒓ⳦ࡀ㸪⳦ᚑᒓᰤ㣴ᛶࡢ✀ᏊⓎⱆࡢඹ⏕⳦࡜ࡋ࡚㸪␗࡞ࡿࢢ࣮ࣝࣉࡢ᳜≀࡟᥇⏝ࡉࢀࡓࡢ࠿ࡣ㸪⳦ᚑᒓᰤ㣴᳜≀ࡢ㐍໬ࡸ⏕⌮ᶵᵓࢆ⪃࠼ࡿୖ࡛㸪㘽࡜࡞ࡿせᅉ࠿ࡶࡋࢀ࡞࠸ࠋ

㸵㸬ඹ⏕⳦ࡢ஌ࡾ᥮࠼ ୖグࡢࡼ࠺࡟㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢⓎⱆࡢඹ⏕⳦ࡣ㸪እ⏕⳦᰿ᛶ࡛ࡣ࡞࠸㸦ᶞᮌ᰿࡜ඹ⏕ࡋ࡚࠸࡞࠸㸧ࢱ࢖ࣉࡢࣟ࢘ࢱࢣࡢ௰㛫ࡢ⳦࡛࠶ࡿࡇ࡜ࡀࢃ࠿ࡗࡓࠋࡑࢀ࡛ࡣ㸪Ⓨⱆᚋ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡ
ࢯ࢘ࡢ⏕㛗ࡀ㐍ࢇࡔ㝿࡟㸪Ⓨⱆࡢඹ⏕⳦࡛࠶ࡿࡇࡢ⳦ࡣ㸪࠸ࡘࡲ࡛࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘࡜㛵ࢃࡾ⥆ࡅࡿࡢ࡛࠶ࢁ࠺࠿ࠋࡑࡇ࡛௒ᅇ✀Ꮚࣃࢵࢡࢆᇙࡵ࡚Ⓨⱆࡀぢࡽࢀࡓࢩࣛ࢝ࣥࣂ࡜ࢲࢣ࢝ࣥࣂࡢⱝ㱋ᯘ
2 ࢝ᡤࡢᯘᗋᅵተ୰࠿ࡽ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢࡲࡔࢥࣟࢽ࣮ࢆᙧᡂࡋ࡚࠸࡞࠸ⱝ࠸ಶయࡢ⳦᰿ࢆ
᥇ྲྀࡋ㸪࠶ࢃࡏ࡚࿘ᅖࡢඃ༨ᶞᮌࡢ⳦᰿ࢆ᥇ྲྀࡋ࡚㸪ྠᵝ࡟ DNA ࡢゎᯒࢆ౑ࡗ࡚⳦᰿⳦ࡢ✀ྠᐃࢆ⾜ࡗࡓ㸦⾲ 2㸧ࠋ ࡑࡢ⤖ᯝ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢⱝ࠸ぶಶయࡢ⳦᰿࡛ࡣ㸪ࡑࡢ࡯࡜ࢇ࡝ࢆእ⏕⳦᰿ᛶࡢ⳦ࡀ༨ࡵ
࡚࠾ࡾ㸪Ⓨⱆࡢ㝿ࡢඹ⏕⳦࡛࠶ࡿࣟ࢘ࢱࢣࡢ௰㛫ࡣ㸪ࢩࣛ࢝ࣥࣂᯘ࡛ 108 ⳦᰿∦ࢆㄪ࡭࡚ 5 ⳦᰿ࡢࡳ࠿ࡽ᳨ฟࡉࢀ㸪ࢲࢣ࢝ࣥࣂᯘ࡛ࡣ 53 ⳦᰿ࢆㄪ࡭࡚඲ࡃ᳨ฟࡉࢀ࡞࠿ࡗࡓࠋࡲࡓ㸪ㄪᰝᯘࡢඃ༨ᶞ✀ࡢ࢝ࣂࣀ࢟ࡢእ⏕⳦᰿࠿ࡽࡣ㸪Ⓨⱆࡢ㝿࡟ぢࡽࢀࡓࣟ࢘ࢱࢣࡢ௰㛫ࡢ⳦ࡣ඲ࡃ᳨ฟࡉࢀ࡞࠿ࡗࡓࠋ࣋ࢽ
ࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢ✀Ꮚࡣ㠀ᖖ࡟ᑠࡉࡃ㸪Ⓨⱆࡢᚋ࡟࡝ࡢ⛬ᗘࡢ᫬㛫ࢆ⤒࡚ⴥࢆᙧᡂࡍࡿࡢ࠿᫂ࡽ࠿
࡟࡞ࡗ࡚࠸࡞࠸ࡀ㸪ᑠࡉ࠸࡞ࡀࡽࡶⴥࢆᙧᡂࡋࡓ᫬Ⅼ࡛㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢඹ⏕⳦ࡢ஌ࡾ᥮࠼
Y. Hashimoto - 7

植物科学最前線 5:127 （2014）ࡀጞࡲࡗ࡚࠸ࡿࡼ࠺࡛࠶ࡗࡓࠋ
਀2 ‫ڮ‬ढ़ਚभ଄ೡढ़ংঀय़੸ઃ௔ऊै఼਄खञঋॽংॼॖॳখॡ९क़भ༳உ༳ध३ছढ़থংुखऎमॲॣढ़থংभਗে༳உ༳भRFLP ॱॖউ଄ೡॲॣढ़থং‫؞‬঑খঐঁথঀय़੸ઃ௔

଄ೡ३ছढ़থং੸ઃ௔
ঋॽংॼॖॳখॡ९क़

३ছढ़থংঋॽংॼॖॳখॡ९क़

ॲॣढ़থং୹૥खञ༳உਯ

108

122

53

66

লਠRFLP ॱॖউਯ

34

42

14

26

৅࿈ৎলਠ༳ध৊गRFLP ॱॖউ

5

0

0

0

ુৢखथলਠRFLP ॱॖউਯ4 (5.5%)3 (8.1%)

ถ಍मHashimoto et al. 2012 ૞ස ࡇࡢࡼ࠺࡞ඹ⏕⳦࡜ࡢ㛵ಀᛶࡀ㸪ࡑࡢ௚ࡢ࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫࡟ࡶᙜ࡚ࡣࡲࡿࡢ࠿ࡣ᫂ࡽ࠿࡛ࡣ࡞
࠸ࡀ㸪࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢඹ⏕⳦࡟࠾࠸࡚㸪Ⓨⱆࡢ㝿ࡢ⭉⏕ᛶࡢ⳦࠿ࡽ㸪ගྜᡂ⬟ຊࢆᣢࡗࡓぶಶయ࡛ࡣከᵝ࡞እ⏕⳦᰿⳦࡟஌ࡾ᥮࠼ࡀ࠾ࡁ࡚࠸ࡿࡇ࡜ࡣ⯆࿡῝࠸ࠋࣛࣥࡢ௰㛫ࡸࢩࣕࢡࢪࣙ࢘ࢯ࢘
ࡢ௰㛫࡛ࡣ㸪ከࡃࡢሙྜඹ⏕⳦ࡣ⏕㛗ࡢ㐣⛬࡛ኚ໬ࡋ࡞࠸ࡇ࡜ࡀከࡃ㸪ࡲࡓኚ໬ࡀぢࡽࢀࡿ㝿ࡣ㸪⭉⏕⳦࠿ࡽ⭉⏕⳦㸪ࡶࡋࡃࡣእ⏕⳦᰿⳦࠿ࡽ
እ⏕⳦᰿⳦࡬ࡢኚ໬࡛࠶ࡾ㸪⭉⏕⳦࠿ࡽእ⏕⳦᰿⳦࡬ࡢኚ໬࡛ࡣ࡞࠸
㸦኱࿴࣭㇂ட 2009㸧ࠋ࡛ࡣ㸪ᙼࡽࡀ⏕⫱ࡢ㐣⛬࡛⭉⏕ᛶࡢ⳦࠿ࡽእ⏕⳦᰿ᛶࡢ⳦࡬஌ࡾ᥮࠼ࡿ⌮⏤ࡣఱ࡛࠶ࢁ࠺࠿ࠋ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡣⓎⱆ࠿ࡽⴥࢆᙧᡂࡍࡿࡲ࡛ࡣ㸪᏶඲࡞⳦ᚑᒓᰤ㣴ᛶࡢ⏕ࡁ᪉࡛࠶ࡿࠋ୍᪉㸪ⴥࢆᙧᡂࡋࡓᚋࡢ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡣ㸪๓㏙ࡢ㏻ࡾ㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶ࡛࠶ࡿ࡜ࡢሗ࿌ࡀ࠶ࡾ㸪ࡲࡓ㸪௒ᅇ✀Ꮚࣃࢵࢡࢆᇙタࡋࡓ࣋ࢽࣂࢼ
࢖ࢳࣖࢡࢯ࢘⩌ⴠࡢ࠸ࡃࡘ࠿࡛ⴥࡢᏳᐃྠ఩య 13C ࡢ್ࢆ ᐃࡋࡓ࡜ࡇ
ࢁ㸪ࡸࡣࡾ㸪㒊ศⓗ⳦ᚑᒓᰤ㣴ᛶ࡛࠶ࡿ࡜ぢࡽࢀࡿ್ࢆᚓ࡚࠸ࡿ㸦ᶫᮏ㸪ᮍⓎ⾲ࢹ࣮ࢱ㸧ࠋ୍⯡ⓗ࡟⪃࠼࡚㸪⮬ࡽᅵተ୰ࡢⴠⴥ࡞࡝᳜≀㑇యࢆศゎࡋ࡚࠸ࡿ⭉⏕⳦ࡼࡾࡶ㸪ᶞᮌࢆᐟ୺࡜ࡋ࡚ࡑࡢගྜᡂ⏘≀ࢆࡶࡽࡗ࡚
࠸ࡿእ⏕⳦᰿⳦ࡢ᪉ࡀ㸪ከࡃࡢ⢾ࢆᏳᐃࡋ࡚ᚓࡽࢀࡿ࡜⪃࠼ࡽࢀࡿࠋ࡞ᅗ ࣄࢺࢶࣂ࢖ࢳࣖࢡࢯ࢘
㸦✀Ꮚᙧᡂ᫬ᮇ㸧ࡐ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࡛࢘ࡣ㸪⤯ᑐⓗ࡞⳦ᚑᒓᰤ㣴ᛶࡢ✀ᏊⓎⱆẁ㝵࡛እ⏕⳦᰿ᛶࡢ⳦ࡀඹ⏕⳦࡛࡞࠸ࡢ࠿ࢃ࠿ࡽ࡞࠸ࡀ㸪Ⓨⱆ┤ᚋࡢᑠࡉ࡞ᐇ⏕ࡢẁ㝵࡛ࡣ㸪⭉⏕ᛶࡢ⳦ࢆ㑅ࡪ࣓ࣜࢵࢺࡀ࠶ࡿࡢ࠿ࡶࡋࢀ࡞࠸ࠋ≉࡟࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡢⓎⱆࡀ㸪ᡂ⇍ࡋࡓ᳃ᯘࡼࡾࡶ㸪ࡲ
ࡔᅵተࡢⓎ㐩ࡀ࠶ࡲࡾ㐍ࢇ࡛࠸࡞࠸᮲௳ࡢᯘ࡛ぢࡽࢀ࡚࠸ࡿࡇ࡜࠿ࡽ㸦⾲ 1㸧㸪ඹ⏕⳦ࡢ⏕⫱㐺ᆅࡀ㝈ᐃࡉࢀ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿࠋࡇࡢࡼ࠺࡞せᅉ࡟ࡼࡗ࡚Ⓨⱆሙᡤࡀ㝈ᐃࡉࢀࡿࡇ࡜ࡀ㸪ࡇࡢ᳜≀ࡀ᪂ࡓ࡞ᐃ╔ሙᡤࢆᗈࡆࡿࡢ࡟୍ᙺ㈙ࡗ࡚࠸ࡿࡢ࠿ࡶࡋࢀ࡞࠸ࠋࡲࡓ୍᪉࡛㸪ࡍࡃ࡞ࡃ࡜ࡶぶಶయ࡛⢾ࢆᏳᐃࡋ࡚ᚓࡽࢀࡿ࡜⪃࠼ࡽࢀࡿእ⏕⳦᰿ᛶࡢ⳦࡜ࡢඹ⏕㛵ಀࢆᣢࡘࡇ࡜ࡀฟ᮶ࡿࡼ࠺࡟࡞ࡗࡓࡇ
࡜ࡣ㸪ᙼࡽࡀ኱ࡁ࡞ࢥࣟࢽ࣮ࢆసࡗ࡚ᬯ࠸ᯘᗋ࡛ඃ༨ⓗ࡟⏕⫱ฟ᮶ࡿせᅉ࡛࠶ࡿ࠿ࡶࡋࢀ࡞࠸ࠋࡉࡽ
࡟㸪እ⏕⳦᰿⳦࡜ඹ⏕ࡍࡿࡼ࠺࡟࡞ࡗࡓࡇ࡜ࡣ㸪⳦ᚑᒓᰤ㣴ᛶࡢ⋓ᚓࡢ㐍໬㐣⛬ࡢ๓㐺ᛂ࡜ࡋ࡚ാ࠸
࡚࠸ࡿࡢ࠿ࡶࡋࢀ࡞࠸ࠋྠࡌ࢖ࢳࣖࢡࢯ࢘ࡢ௰㛫ࡢ࢖ࢳࣖࢡࢯ࢘㸦Pyrola japonica㸧࡛ࡣ㸪✀ෆ࡛ࣄࢺࢶࣂ࢖ࢳࣖࢡࢯ࢘࡜࿧ࡤࢀࡿⴥࡢࢧ࢖ࢬࡀᑠࡉࡃ㏥໬ࡋࡓࡶࡢࡀ▱ࡽࢀ࡚࠾ࡾ㸦ᅗ 8㸧㸪⳦ᚑᒓᰤ㣴ᛶࡢ⛬ᗘࡀ㧗ࡲࡗ࡚࠸ࡿᵝᏊࡀぢཷࡅࡽࢀࡿࠋ

Y. Hashimoto - 8

植物科学最前線 5:128 （2014）㸶㸬ࡲ࡜ࡵ 㒊ศⓗ⳦ᚑᒓᰤ㣴᳜≀࡜⪃࠼ࡽࢀࡿ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࢆᑐ㇟࡟㸪ᚤ⣽✀ᏊࡢⓎⱆ࡜ࡑࡢ㝿ࡢඹ⏕⳦࡟ࡘ࠸࡚ࡢ◊✲౛ࢆ⤂௓ࡋࡓࠋ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡣ㸪᏶඲࡞⳦ᚑᒓᰤ㣴ᛶࡢⓎⱆ᫬࡟ࡣ≉␗ⓗ࡞⭉⏕⳦࡜㛵ಀࢆ⤖ࡪࡀ㸪ⴥࡢ⏕㛗࡟కࡗ࡚⳦ࢆ஌ࡾ᥮࠼࡚㸪ᡂ⇍ಶయ࡛ࡣ࿘ᅖࡢᶞᮌࡀ฼⏝ࡍࡿከᵝ࡞እ⏕⳦᰿⳦࡜㛵ಀࢆ⤖ࢇ࡛࠾ࡾ㸪ࡑࡢ⏕ά⎔ࡢ୰࡛ඹ⏕⳦ࢆኚ໬ࡉࡏ࡚࠸ࡿࡼ࠺࡛࠶ࡗࡓࠋࡇ
ࡢࡼ࠺࡞ඹ⏕⳦ࡢ஌ࡾ᥮࠼ࡀ㸪⳦ᚑᒓᰤ㣴ᛶࡢ㐍໬࡬ࡢ๓㐺ᛂ࡜ࡋ࡚ࡢᶵ⬟ࡋ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿ࡜
⪃࠼ࡽࢀࡿࠋࡲࡓ㸪ᬯ࠸᳃ᯘᯘᗋ࡛⏕ࡁࡿ࣋ࢽࣂࢼ࢖ࢳࣖࢡࢯ࢘ࡀ኱ࡁ࡞ࢥࣟࢽ࣮ࢆᙧᡂࡋ࡚ඃ༨ⓗ࡟⏕ࡁࡽࢀࡿ⫼ᬒ࡟ࡣ㸪ࡇࡢࡼ࠺࡞⳦ࢆ฼⏝ࡋࡓᕦࡳ࡞⏕Ꮡᡓ␎ࡀ㛵ࢃࡗ࡚࠸ࡿ࡜⪃࠼ࡽࢀࡿࠋᯘᗋ
࡜࠸࠺ᬯ࠸⎔ቃ࡛ᑠࡉ࡞᳜≀ࡀ⏕ࡁࡿ୰࡛㸪⳦࡟㢗ࡗ࡚⏕ࡁࡿ࡜࠸࠺⏕ࡁ᪉ࡀጞࡲࡗࡓ࡜⪃࠼ࡽࢀࡿࡀ㸪᳃ᯘᅵተ୰࡛␗✀ࡢ᳜≀㛫ࡀ⳦᰿⳦ࡢ⳦⣒࡟ࡼࡗ࡚⧅ࡀࡗ࡚࠾ࡾ㸪㣴ศࡢࡸࡾ࡜ࡾࡀᏑᅾࡍࡿ஦
ᐇࡣ㸪᳃ᯘ⏕ែ⣔ࢆぢࡿୖ࡛᪂ࡓ࡞どⅬࢆ୚࠼࡚ࡃࢀࡿࡔࢁ࠺ࠋㅰ㎡
 ᮏ✏࡛⤂௓ࡋࡓ◊✲ࡢ୍㒊ࡣ㸪⛉Ꮫ◊✲㈝⿵ຓ㔠 14780437 ࡟ࡼࡾࡲࡍࠋࡲࡓ㸪ࡇࢀࡽ◊✲ࡣ㸪ᖏᗈ␆⏘኱Ꮫ࡛ࡢᅧྖ⥤ᏊẶ࣭⚟ᕝᝅẶࡢಟኈㄽᩥ◊✲ࡢ୍㒊࡜ᓥᮏ⧛Ặ࣭Ⲩᮌ㝧ᏊẶࡢ༞ᴗㄽᩥ◊✲ࡢ୍㒊ࡢᡂᯝࢆࡲ࡜ࡵࡓࡶࡢ࡛㸪ᙼࡽࡢປࢆᝰࡋࡲࡠ◊✲࡬ࡢྲྀࡾ⤌ࡳ࡟ឤㅰ࡜ᩗពࢆ⾲ࡋࡲࡍࠋᘬ⏝ᩥ⊩
Bidartondo, M.I. & Read, D.J. 2008. Fungal specificity bottlenecks during orchid germination and development.
Mol. Ecol. 17: 3707-3716.
Bruns, T.D. & Read, D.J. 2000. In vitro germination of non-photosynthetic, myco-heterotrophic plants stimulated
by fungi isolated from the adult plants. New Phytol. 148: 335-342.
Eriksson, O. & Kainulainen, K. 2011. The evolutionary ecology of dust seeds. Perspect. Plant Ecol. Evol. Syst. 13:
73-87.
Hashimoto, Y., Fukukawa S., Kunishi, A., Suga, H., Richard, F., Sauva, M., & Selosse, M.-A. 2012.
Mycoheterotrophic germination of Pyrola asarifolia dust seeds reveals convergences with germination inorchids. New Phytol. 195: 620-630.
Hynson, N.A. & Bruns T.D. 2009. Evidence of a myco-heterotroph in the plant family Ericaceae that lacks
mycorrhizal specificity. Proc. R. Soc. B 276: 4053-4059.
Leake, J.R., McKendrick, S.L., Bidartondo, M., & Read, D.J. 2004. Symbiotic germination and development of
the myco-heterotroph Monotropa hypopitys in nature and its requirement for locally distributed Tricholoma spp.
New Phytol. 163: 405-423.
Massicotte, H.B., Melville, L.H., Tackaberry, L. & Peterson, R.L. 2008. A comparative study of mycorrhizas inseveral genera of Pyroleae (Ericaceae) from western Canada. Botany 86: 610-622.
Rasmussen, H.N. & Whigham, D.F. 1998. Importance of woody debris in seed germination of Tipularia discolor(Orchidaceae). Am. J. Bot. 85: 829-834.
Roy, M., Gonneau, C., Rocheteau, A., Berveiller, D., Thomas, J.-C., Damesin, C., & Selosse M.-A. 2013. Why do
mixotrophic plants stay green? A comparison between green and achlorophyllous orchid individuals in situ.
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Ecol. Monogr. 83: 95-117.
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Tedersoo, L., Pellet, P., Kõljalg, U. & Selosse, M.-A. 2007. Parallel evolutionary paths to mycoheterotrophy inunderstorey Ericaceae and Orchidaceae: ecological evidence for mixotrophy in Pyroleae. Oecologia 151:
206-217.
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Yagame, T., & Yamato, M. 2008. Isolation and identification of mycorrhizal fungi associated with
Stigmatodactylus sikokianus (Maxim. ex Makino) Rauschert (Orchidaceae) Mycoscience 49: 388-391
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Monotropastrum humile var. humile is associated with diverse ectomycorrhizal Russulaceae fungi in Japanese
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኱࿴ᨻ⚽, ㇂ட㧗ᗈ 2009. ࣛࣥ⛉᳜≀࡜⳦㢮ࡢඹ⏕㸬᪥⳦ሗ 50: 21-42.
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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, Saga
University, 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 University
Matsudo, 271-8510, Japan

Tsukuba Botanical Garden, National Museum of Nature and Science
4-1-1, 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࡟ࡼࡗ࡚
ࡏࡗࡏ࡜⮬ศ࡛஧㓟໬Ⅳ⣲࡜Ỉ࠿ࡽ᭷ᶵⅣ⣲࡛࠶ࡿ⢾ࢆྜᡂࡋ࡞ࡀࡽᬽࡽࡋ࡚࠸ࡿࠋ୍᪉㸪⳦ᚑᒓᰤ㣴᳜≀ࡣගྜᡂ࡛ࡣ࡞ࡃ㸪ඹ⏕ࡍࡿ⳦᰿⳦࠿ࡽ᭷ᶵⅣ⣲ࢆࡶࡽࡗ࡚ᬽࡽࡋ࡚࠸ࡿࠋ
ከࡃࡢගྜᡂ᳜≀࡛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໬ࡋ
࡚࠸ࡗࡓࡢ࠿㸪ࡑࡢㅦ࡟㏕ࡗࡓ⚾㐩ࡢ◊✲౛ࢆ⤂௓ࡋ࡚㸪᭱ᚋ࡟ᮏ◊✲ศ㔝࡟࠾ࡅࡿ௒ᚋࡢ
ᒎᮃࢆ⪃࠼ࡿࠋ

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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࠸ࡓ
ࡢ࡛࠶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⣔ࡣ
Davis (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

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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࡟ศࡅࡽࢀࡿࠋ
ࡑࢀࡒࢀࡢඹ⏕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⏕⳦᰿⳦࠿ࡽ
Ⅳ⣲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࠸⣔⤫࡛࠶ࡿࢻ࢘ࢲࣥࢶ
ࢶࢪࡢ௰㛫ࡣ㸪࢔࣮ࣂࢫ࣮࢟ࣗࣛ⳦᰿ᛶ࡛࠶ࡿ࡜ሗ
࿌ࡉࢀ࡚࠸ࡿࡇ࡜࠿ࡽ㸦Abe 2005, Obase & Matsuda
2013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㣴᳜≀࡛࠶ࡿࢠࣥࣜࣙ࢘ࢯ࢘ࡸࢩࣕࢡࢪࣙ
࢘ࢯ࢘ࡢ௰㛫ࡣ㸪ࡇࡢḟ࡟ྂ࠸⣔⤫࡛㸪እ⏕⳦᰿ᛶࡢᢸᏊ⳦㢮࡜ඹ⏕ࡋ࡚࠸ࡿࠋ࠾ࡑࡽࡃࢶ
ࢶࢪ⛉࡛ࡣගྜᡂࢆࡸࡵࡿ㐍໬ࡢ㐣⛬࡛㸪⳦᰿⳦ࡀ AM ⳦࠿ࡽእ⏕⳦᰿⳦࡬ࡍࡾ࠿ࢃࡗ࡚ࡋ
ࡲࡗࡓࡢ࡛ࡣ࡞࠸࠿࡜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⣲ࢆࡶࡽࡗ࡚
ᬽࡽࡋ࡚࠸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⬊Ꮚ
ࡢⓎ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࠾ࡇࡗ
࡚࠸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࠿ࢆ᥎ᐃࡍࡿ
ࡓࡵ࡟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࡜ࢧ
࣑࢞ࣛࣥࡢ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࡜ࡶ
᫂ࡽ࠿࡟࡞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࠿ࡗ
ࡓࠋ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๭ࢆᯝ
ࡓࡋ࡚࠸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⳦ᚑ
ᒓᰤ㣴᳜≀ࡢඹ⏕┦ᡭ࡛࠶ࡿ⳦᰿⳦ࡣ㸪࡝ࡇ࡛࡝ࡢࡼ࠺࡟ᬽ

Y. Ogura-Tsujita - 7

ᅗ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⳦᰿⳦࡬ཷࡅ
Ώࡉࢀࡿࡀ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,
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