植物科学最前線 7:153 (2016)ࢥࢣ᳜≀࡟࠾ࡅࡿࢽࢵࢳᡓ␎ࡢࡓࡵࡢ⣽⬊ศ໬㖡ⱏ࡟࠾ࡅࡿ↓ᛶⱆࡢศ໬㔝ᮧಇᑦ㥅⃝┒୍㑻ᴬཎᆒ⌮໬Ꮫ◊✲ᡤ⎔ቃ㈨※⛉Ꮫ◊✲ࢭࣥࢱ࣮ࠛ⚄ዉᕝ┴ᶓ὾ᕷ㭯ぢ༊ᮎᗈ⏫ᮾ◊✲Ჷ (ᮾி኱Ꮫ኱Ꮫ㝔᪂㡿ᇦ๰ᡂ⛉Ꮫ◊✲⛉ඛ➃⏕࿨⛉Ꮫᑓᨷࠛ༓ⴥ┴᯽ᕷ᯽ࡢⴥᮾி኱Ꮫ᯽࢟ࣕࣥࣃࢫ⏕࿨Ჷྡྂᒇ኱Ꮫ኱Ꮫ㝔⏕࿨㎰Ꮫ◊✲⛉ࠛྡྂᒇᕷ༓✀༊୙⪁⏫ྡྂᒇ኱Ꮫ኱Ꮫ㝔⏕࿨㎰Ꮫ◊✲⛉⏕≀໬Ꮫ◊✲ᐊ
Toshihisa Nomura1, Seiichiro Hasezawa2, Hitoshi Sakakibara1,3
Cell differentiation for niche strategy in bryophytes-Gemma differentiation in copper mossesKeyword: bryophytes, copper moss, gemma, niche, Scopelophila cataractae
1RIKEN Center for Sustainable Resource Science, 1-7-22 Suehiro-cho, Tsurumi,Yokohama, Kanagawa 230-0045, Japan2

Department of Integrated Biosciences, Graduate School of Frontier Sciences,

The University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa, Chiba 277-8562, Japan3

Department of Biological Mechanisms and Functions, Graduate School ofBioagricultural Sciences, Nagoya University, Furo, Chikusa, Nagoya, 464-8601,
Japan㸯㸬ࡣࡌࡵ࡟⏕≀ࡣ㸪ྛࠎࡢ≉ᛶ࡟㐺ࡋࡓ㒊ศⓗ࡞⎔ቃ㸪⏕ែⓗᆅ఩㸦ࢽࢵࢳ㸧ࢆ᭷ࡋ㸪ࡑࡇ࡛⏕Ꮡࡍࡿࡓࡵ࡟㸪ᰤ㣴※㸪⾜ື㸪⣽⬊ศ໬ࡸᙧែ㸪௦ㅰ⣔㸪⎔ቃࢫࢺࣞࢫ⪏ᛶ࡞࡝ࢆᕦࡳ࡟ኚ໬ࡉࡏྛ⎔ቃ࡟㐺ᛂࡋ࡚࠸ࡿࠋࡇࡢ⏕≀Ꮫ࡛⏝࠸ࡿ⏝ㄒ࡜ࡣᑡࡋ␗࡞ࡿព࿡࡛㸪ᮏ✏ࡢࢱ࢖ࢺࣝ࡟ࡣ㸪ࢽࢵࢳᡓ␎࡜࠸࠺ゝⴥࢆ౑⏝ࡉࡏ࡚㡬࠸ࡓࠋࢽࢵࢳᡓ␎࡜ࡣ㸪⤒Ⴀᡓ␎ࡢ୍ࡘ࡛࠶ࡿࠋࡇࡇ࡛ࡢࢽࢵࢳ࡜࠸࠺ゝⴥࡢព࿡ࡣ㸪౛࠼ࡤ͆ࢽࢵࢳ࡞ᴗ✀͇࡜ゝ࠺ࡼ࠺࡟͆≉Ṧ࡞͇࡜࠸࠺ព࿡ྜ࠸࡛౑ࢃࢀࡿࠋ༶ࡕ㸪ࢽࢵࢳᡓ␎࡜ࡣ㸪୰ᑠ௻ᴗ࡞࡝ࡀ⊂⮬ࡢᢏ⾡ࡸ▱㆑࡞࡝ࢆṊჾ࡟㸪኱௻ᴗࡀᡭࢆฟࡉ࡞࠸ศ㔝㸦࠸ࢃࡺࡿࡍࡁࡲ⏘ᴗ㸧ࡢᴗ✀࡟⊂༨ⓗᒎ㛤ࢆᅗࡿ࡜࠸࠺⤒Ⴀᡓ␎ࡢࡇ࡜࡛࠶ࡿࠋࡇࢀ࡜㢮ఝࡋࡓ⏕ࡁṧࡾᡓ␎ࢆ࡜ࡿ✀ࡣ㸪⏕ែᏛ࡛ࡣࢫ࣌ࢩࣕࣜࢫࢺ࡜࿧ࡤࢀࡿࠋࢫ࣌ࢩࣕࣜࢫࢺࡣ㸪ᵝࠎ࡞⏕≀✀ෆ࡟࠾࠸࡚ࡳࡽࢀࡿࡀ㸪㝣ୖ᳜≀࡟࠾࠸࡚ࢥࢣ᳜≀࡛ࡣ㸪≉Ṧ࡞⏕⫱ᆅࢆ㑅ᢥࡍࡿࢱ࢖ࣉࡢࢫ࣌ࢩࣕࣜࢫࢺࡀ┠❧ࡘ༳㇟ࡀ࠶ࡿࠋ౛࠼ࡤ㸪⏕ࡁࡓⴥࡢୖ࡟╔⏕ࡍࡿ࢝ࣅࢦࢣ㸪㐨㊰⬥࡞࡝࡟⏕⫱ࡍࡿࢠࣥࢦࢣࡸ࣍ࢯ࢘ࣜࢦࢣ㸪
T. Nomura-1BSJ-Review 7:153 (2016)

植物科学最前線 7:154 (2016)㖔ᒣ࡞࡝㖡ࡸ㕲ࡀ㇏ᐩ࡟Ꮡᅾࡍࡿ⎔ቃ࡟⏕⫱ࡍࡿ࣍ࣥࣔࣥࢪࢦࢣࡸ࢖࣐࣡ࢭࣥ࣎ࣥࢦࢣ㸪Ὕ❍࡞࡝ప↷ᗘ⎔ቃࢆዲࡴࣄ࢝ࣜࢦࢣ࡜࠸ࡗࡓ✀ࡀᣲࡆࡽࢀࡿࠋࡇࡢࡼ࠺࡞ࢥࢣ᳜≀ࡓࡕࡣ㸪ఱ࠿ࡋࡽࡢᕤኵࢆจࡽࡋ࡚㸪≉Ṧ⎔ቃࢆぢࡘࡅ㸪ᐃ╔ࡋ㸪㐺ᛂࡋ࡚࠸ࡿ࡜᥎ࡉࢀࡿࠋṧᛕ࡞ࡀࡽከࡃሙྜ㸪ࡑࡢ௙⤌ࡳࡣศ࠿ࡗ࡚࠸࡞࠸ࡀ㸪⣽⬊ศ໬ࡀ⏕ែⓗᆅ఩ࡢ⋓ᚓ࡟㛵୚ࡍࡿ౛ࢆ㸪ࣄ࢝ࣜࢦࢣ Schistostega pennata (Hedw.) F.Weber & D.Mohr ࡟࠾࠸࡚ぢࡿࡇ࡜ࡀ࡛ࡁࡿࠋ
ࣄ࢝ࣜࢦࢣࡣ㸪Ὕ❍ࡸᒾ㛫࡞࡝ప↷ᗘ⎔ቃࢆ⏕ែⓗᆅ఩࡜ࡍࡿཎጞⓗ࡞⹊㢮ࡢ୍✀࡛࠶ࡿࠋࡇࡢ✀ࡣ㸪ప↷ᗘ⎔ቃ࡟㐺ᛂࡍࡿࡓࡵ࡟㸪ࣞࣥࢬ≧⣽⬊࡜࠸࠺≉Ṧ࡞⣽⬊ࢆ㸪⹊㢮ࡢⓎ㐩ึᮇ࡟ࡳࡽࢀࡿ⣒≧ࡢ⤌⧊࡛࠶ࡿཎ⣒య࡟ศ໬ࡉࡏࡿࠋࣞࣥࢬ≧⣽⬊ࡣ㸪ග※࡟ᑐࡋ࡚᪉
ྥᛶࡢ࠶ࡿᙧ≧ࢆࡋࡓ⣽⬊࡛㸪ࡑࡢྡࡢ㏻ࡾࣞࣥࢬࡢᙺ๭࡛ᙅ࠸ගࢆ㞟ගࡋ㸪ⴥ⥳య࡟㞟ࡵࡿാࡁࢆᣢࡘ㸦ᅗ1B㸧ࠋࡇࡢ࡜ࡁ㸪ගྜᡂ࡟฼⏝ࡉࢀ࡞࠸⥳Ⰽࡢගࡣ෌ᖐⓗ࡟཯ᑕࡍࡿࡓࡵ⥳Ⰽ࡟ගࡗ࡚ぢ࠼࡚ࡋࡲ࠺㸦ᅗ1A㸧ࠋᅗ㸯ࣄ࢝ࣜࢦࢣ࡟ࡳࡽࢀࡿప↷ᗘ⎔ቃ࡟㐺ᛂࡍࡿࡓࡵࡢ⣽⬊ศ໬A. ᬯ㜌࡛ගࡗ࡚ࡳ࠼ࡿࣄ࢝ࣜࢦࢣ㸦㯤▮༳, 㛗㔝┴ᒾᮧ⏣ࡢ⮬⏕ᆅ࡟࡚᧜ᙳ㸧
B. ࣄ࢝ࣜࢦࢣࡢࣞࣥࢬ≧⣽⬊,㸦ⓑ▮༳㸸ධᑕග, ⥳▮༳㸸཯ᑕග㸧ࢫࢣ࣮ࣝࣂ࣮ࡣ, 10 ȝm୍᪉㸪ᡃࠎࡀ◊✲ᮦᩱ࡜ࡋ࡚࠸ࡿ࣍ࣥࣔࣥࢪࢦࢣ Scopelophila cataractae (Mitt.) Broth. ࡣ㸪௚ࡢ᳜≀✀ࡢ⏕⫱ࡀᅔ㞴࡞㧗⃰ᗘࡢ㖡ࡀᏑᅾࡍࡿ⎔ቃࡢࡳࢆ⏕ែⓗᆅ఩࡜ࡍࡿࠋᮏ✀ࡀ࡝ࡢࡼ࠺࡟㸪ࡑࡢࡼ࠺࡞ሙᡤࢆぢฟࡋ㸪ᐃ╔ࡍࡿࡢ࠿ࡣ୙࡛᫂࠶ࡗࡓࡀ㸪ᡃࠎࡣ㸪ࡑࡇ࡟ࡶ⣽⬊ศ໬ࡀ㛵୚ࡋ࠺ࡿࡇ࡜ࢆぢฟࡋ࡚࠸ࡿࠋᮏ✏࡛ࡣࡇࢀࡽ࡟ࡘ࠸࡚⤂௓ࡍࡿ࡜ඹ࡟㸪௒ᚋࡢᒎᮃ࡟ࡘ࠸࡚㏙࡭ࡓ࠸ࠋ㸰㸬㖡ࡀ㇏ᐩ࡞⎔ቃࢆ⏕ែⓗᆅ఩࡜ࡍࡿ࣍ࣥࣔࣥࢪࢦࢣ࣍ࣥࣔࣥࢪࢦࢣࡣ㸪⚄♫௖㛶࡟࠾ࡅࡿ㖡ⵌࡁᒇ᰿ࡢୗࡸ㖡㖔ᒣ࿘㎶࡞࡝㸪㧗⃰ᗘࡢ㖡ࢆྵࡴ⎔ቃࢆ⏕⫱ᆅ࡜ࡍࡿ⹊㢮࡟ᒓࡍࡿࢥࢣ᳜≀࡛࠶ࡿ㸦Satake et al. 1988㸧ࠋᮏ✀ࡣࠊୡ⏺୰ࡢ㖡ࡀ㇏ᐩ࡞⎔ቃ࡟ࡳࡽࢀࡿࡀ㸦Show 1989㸧㸪᪥ᮏ࡟࠾࠸࡚ࡣ㸪ᮾிࡢụୖᮏ㛛ᑎ࡛ึࡵ࡚☜ㄆࡉࢀࡓࡓࡵ㸪ࡇࡢ࿴ྡࡀ௜ࡅࡽࢀ࡚࠸ࡿ㸦Sakurai 1934, ᅗ2A㸧ࠋ

T. Nomura-2BSJ-Review 7:154 (2016)

植物科学最前線 7:155 (2016)ᅗ㸰㸬࣍ࣥࣔࣥࢪࢦࢣ࡟ࡘ࠸࡚
A. ụୖᮏ㛛ᑎ஬㔜ሪୗࡢ࣍ࣥࣔࣥࢪࢦࢣ
B. ࣍ࣥࣔࣥࢪࢦࢣࡢཎ⣒యඛ➃㒊࡟ᙧᡂࡉࢀࡓ↓ᛶⱆ, ▮༳㸸↓ᛶⱆ, ▮ᑼ㸸⬺㞳⣽⬊, ࢫࢣ࣮ࣝࣂ࣮ࡣ, 20 ȝmC. ษࡾ㞳ࡉࢀࡓ↓ᛶⱆ, ࢫࢣ࣮ࣝࣂ࣮ࡣ, 20 ȝmD. ↓ᛶⱆࡢⓎⱆ, ▮༳㸸Ⓨⱆࡋࡓ↓ᛶⱆ, ▮ᑼ㸸᪂ࡓ࡟ᙧᡂࡉࢀ࡚࠸ࡿ↓ᛶⱆ, ࢫࢣ࣮ࣝࣂ࣮ࡣ, 50 ȝmࡇࡢࡼ࠺࡞㔜㔠ᒓởᰁ⎔ቃࢆ⏕ែⓗᆅ఩࡜ࡍࡿ᳜≀✀ࡣMetallophytes ࡜࿧ࡤࢀ㸪㔠ᒓࡢ㇏ᐩ࡞⎔ቃ࡟ࡢࡳ⏕⫱ࡍࡿ Obligate metallophytes ࡜㸪㔠ᒓࡀ㇏ᐩ࡛࡞࠸⎔ቃ࡛ࡶ⏕⫱ࡀぢࡽࢀࡿ Facultative metallophytes ࡜࡟༊ูࡉࢀࡿࠋࢥࢣ᳜≀ࡢ୰࡟ࡣ㸪≉࡟㖡ࡢ㇏ᐩ࡞⎔ቃࢆዲࡴ࠸ࡃࡘ࠿ࡢ✀ࡀ࠾ࡾ㸪ࡑࢀࡽࡣ㖡ⱏ㸦Copper mosses㸧࡜⥲⛠ࡉࢀ࡚࠸ࡿ㸦Persson 1956,Shaw 1994㸧ࠋࡇࡢࡼ࠺࡞ࢥࢣ᳜≀✀ࡀ㸪࡝ࡢࡼ࠺࡟ࡋ࡚㖡ࡢ㇏ᐩ࡞⎔ቃࢆ᥈ࡋ㸪ᐃ╔ࡋ࡚࠸ࡿࡢ࠿ࡣ⏕ែ࠾ࡼࡧ⏕⌮Ꮫⓗ࡟⯆࿡῝࠸ࡀ㸪ࡑࡢᐇែࡣ᫂ࡽ࠿࡟࡞ࡗ࡚࠸࡞࠸ ໭ᕝ1987 ࠋࡲࡓ㸪࣍ࣥࣔࣥࢪࢦࢣ࡛ࡣ㸪᭷ᛶ⏕Ṫࡀ㠀ᖖ࡟⛥࡛࠶ࡾ㸦㬼ἑ & బ➉ 2010, Show 1989 24⣽⬊㸦୺࡟ࡣ3⣽⬊㸧࠿ࡽᵓᡂࡉࢀࡿ↓ᛶⱆ㸦ᅗ2B, C, Nomura & Hasezawa 2011, Rumsey &Newton 1989㸧ࢆ⏝࠸࡚↓ᛶⓗ࡟ศᕸࢆᣑ኱ࡉࡏ࡚࠸ࡿ࡜⪃࠼ࡽࢀ࡚࠸ࡿࠋᚑࡗ࡚㸪࣍ࣥࣔࣥࢪࢦࢣࡣ㸪ࢡ࣮ࣟࢼ᳜ࣝ≀ࡢ⏕ែ◊✲ࡢᑐ㇟࡜ࡋ࡚ࡶ⯆࿡῝࠸✀࡛࠶ࡿࠋ㸱㸬࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ㖡࡟ࡼࡿ⣽⬊ศ໬
๓㏙ࡢ࡜࠾ࡾ㸪࣍ࣥࣔࣥࢪࢦࢣࡀ㸪㧗⃰ᗘࡢ㖡ởᰁ⎔ቃ࡟ࡢࡳᐃ╔ࡍࡿ௙⤌ࡳࡣㅦ࡛࠶ࡗࡓࠋࡑࡇ࡛㸪ᡃࠎࡣ㸪࣍ࣥࣔࣥࢪࢦࢣࡢᇵ㣴ᰴࢆసฟࡋ㸪Ⓨ㐩ึᮇ࡟࠾ࡅࡿ㖡ῧຍࡢᙳ㡪ࢆㄪ࡭ࡓࠋࡲࡎ㸪↓ᛶⱆࡢⓎⱆ࡬ࡢ㖡ῧຍࡢᙳ㡪ࢆ᳨ドࡋࡓࡀ㸪㖡ࡢ᭷↓࡟ࡼࡿᙳ㡪ࡣぢࡽࢀ࡞࠿ࡗࡓࠋࡋ࠿ࡋ㸪Ⓨⱆࡋࡓཎ⣒యࢆᇵ㣴ࡋ⥆ࡅࡿ࡜㸪㧗⃰ᗘࡢ㖡ῧຍᇵ㣴᮲௳ୗ࡛ࡣࢡࣟࣟࢿ࣐⣽⬊࠿ࡽ㸪ࡼࡾⓎ㐩ࡋࡓẁ㝵ࡢ࢝࢘ࣟࢿ࣐⣽⬊㸦Nomura et al. 2015, ᅗ3A, C㸧࡬ࡢศ໬ࡀಁ㐍ࡉࢀࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋ୍᪉㸪ప⃰ᗘࡢ㖡Ꮡᅾ᮲௳࡛ࡣ㸪ᮏ✀ࡀ୺࡟ศᕸᣑ኱࡟⏝࠸ࡿ࡜ࡉࢀࡿ↓ᛶⱆࡢᙧᡂࡀㄏᑟࡉࢀࡓ㸦Nomura & Hasezawa 2011, Nomura et al.
T. Nomura-3BSJ-Review 7:155 (2016)

植物科学最前線 7:156 (2016)2015, ᅗ3E, F㸧ࠋࡇࡢ⤖ᯝࡣ㸪࣍ࣥࣔࣥࢪࢦࢣࡀⓎ㐩ࡢึᮇẁ㝵࡟࠾࠸࡚㸪⎔ቃ୰ࡢ㖡⃰ᗘ࡟ᛂࡌࡓ⣽⬊ศ໬ࡢไᚚᶵᵓࢆ᭷ࡍࡿࡇ࡜ࢆ♧၀ࡍࡿࠋᅗ㸱㸬࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ⎔ቃ୰ࡢ㖡⃰ᗘ࡟ᛂࡌࡓ⣽⬊ศ໬⹊㢮ࡢཎ⣒య࡟࠾ࡅࡿ⣽⬊ศ໬ㄪ⠇࡟ࡣ㸪࣮࢜࢟ࢩࣥࡸࢧ࢖ࢺ࢝࢖ࢽࣥ㸪࢔ࣈࢩࢪࣥ㓟࡞࡝ࡢ᳜≀࣍ࣝࣔࣥࡀ㸪㛵୚ࡍࡿࡇ࡜ࡀ▱ࡽࢀ࡚࠸ࡿ㸦Cove et al. 2006㸧ࠋࡑࡇ࡛㸪࣍ࣥࣔࣥࢪࢦࢣ࡛ࡳࡽࢀࡓ㖡౫Ꮡⓗ࡞⣽⬊ศ໬ࡢᶵᵓࢆ᫂ࡽ࠿࡟ࡍࡿࡓࡵ㸪ᮏ✀ࡢཎ⣒యࢆ㖡ῧຍ᮲௳ୗ࡛ᇵ㣴ࡋࡓ㝿ࡢྛ✀᳜≀࣍ࣝࣔࣥෆ⏕㔞ࡢኚ໬ࢆゎᯒࡋࡓࠋࡑࡢ⤖ᯝ㸪㧗⃰ᗘࡢ㖡ῧຍ᮲௳ୗ࡛ࡣ㸪≉࡟࣮࢜࢟ࢩࣥෆ⏕㔞ࡀୖ᪼ࡍࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓ㸦Nomura et al. 2015,ᅗ3B㸧ࠋࡲࡓ㸪ప⃰ᗘࡢ㖡Ꮡᅾ᮲௳࡟࠾࠸࡚ࡶ㸪࣮࢜࢟ࢩࣥࡢῧຍࡣ㸪㖡ῧຍ࡜ྠᵝࡢ⣽⬊ศ໬࡬ࡢຠᯝࢆᘬࡁ㉳ࡇࡋࡓࠋຍ࠼࡚㸪㧗⃰ᗘࡢ㖡࡟ࡼࡾㄏᑟࡉࢀࡿ࢝࢘ࣟࢿ࣐⣽⬊ศ໬ࡢಁ㐍ࡣ㸪࣮࢜࢟ࢩࣥస⏝㜼ᐖ๣㸦PEO-IAA㸧ࡸ㸪࣮࢜࢟ࢩࣥ⏕ྜᡂ㜼ᐖ๣㸦L-࢟ࢾࣞࢽࣥ㸧ࡢฎ⌮࡟ࡼࡾᢚไࡉࢀࡓ㸦Nomura et al. 2015㸧ࠋࡇࢀࡽࡢ⤖ᯝ࠿ࡽ㸪࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ㖡⃰ᗘ࡟ᛂࡌࡓ⣽⬊ศ໬ࡢㄪ⠇࡟ࡣ㸪࣮࢜࢟ࢩࣥࡀ㛵୚ࡍࡿࡇ࡜ࡀ᫂ࡽ࠿࡟࡞ࡗࡓࠋ࣍ࣥࣔࣥࢪࢦࢣࡣ㸪⮬㌟ࡢᣢࡘ㧗࠸㖡⪏ᛶ⬟ࢆ᭱኱㝈࡟ά࠿ࡏࡿ⏕ែⓗᆅ఩࡟ࡢࡳᐃఫ࡛ࡁࡿࡼ࠺࡟㸪ࡇࡢࡼ࠺࡞⎔ቃ࡟ᛂࡌࡓ⣽⬊ศ໬ࡢᶵᵓࢆ⋓ᚓࡋࡓࡢ࡛ࡣ࡞࠸࠿࡜⪃࠼ࡽࢀࡿࠋ௚᪉࡛㸪࣍ࣥࣔࣥࢪࢦࢣ࡜㢮ఝࡋࡓཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆᙧᡂࡢ౛ࡀ㸪ᒾࡢ๭ࢀ┠࡞࡝ⷧᬯ࠸⎔ቃࢆዲࡴᩘ✀ࡢ⹊㢮࡛ࡶሗ࿌ࡉࢀ࡚࠸ࡿ㸦Whitehouse 1980㸧ࠋࡇࢀࡽࡢ✀ࡢሙྜ㸪ᬯ᮲௳ୗ࡟࠾࠸࡚㸪ཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆᙧᡂࡀㄏᑟࡉࢀ㸪ࡇࢀࡣ࠶ࡲࡾ࡟ࡶᬯࡍࡂࡿ⎔ቃ࡟฿㐩ࡋ࡚ࡋࡲࡗࡓሙྜࡢ㏨㑊⟇࡛ࡣ࡞࠸࠿࡜⪃࠼ࡽࢀ࡚࠸ࡿࠋ㖡࡟ᑐࡋ࡚㧗࠸⪏ᛶ⬟ࢆ᭷ࡍ
T. Nomura-4
BSJ-Review 7:156 (2016)

植物科学最前線 7:157 (2016)ࡿ࣍ࣥࣔࣥࢪࢦࢣࡢሙྜ㸪㧗⃰ᗘࡢ㖡ࡀ࡞࠸⎔ቃࡣ㸪➇த┦ᡭࡀᏑᅾࡍࡿࡓࡵ㸪⏕⫱࡟୙฼࡞≧ἣ࡜࡞ࡿࠋㄏᑟࡢせᅉࡣ␗࡞ࡿࡀ㸪ୖグࡢ౛࡜㢮ఝࡋࡓᶵᵓ࡟ࡼࡾ㸪↓ᛶⱆᙧᡂࡀㄏᑟࡉࢀࡿࡶࡢ࡜᥎ࡉࢀࡿࠋࡲࡓ㸪ࣄࣙ࢘ࢱࣥࢦࢣ࡞࡝ࡢཎ⣒యࢆྠ୍ࡢᇵᆅ࡛㛗ᮇ㛫ᇵ㣴ࡍࡿ࡜㸪ཎ⣒య⣽⬊ࡢ㛫࡟⬺㞳⣽⬊㸦Tmema cell㸧࡜࿧ࡤࢀࡿᑠࡉ࡞⣽⬊ࡢᙧᡂࡀࡳࡽࢀࡿࡇ࡜ࡀ࠶ࡿ㸦ᅗ4㸧ࠋࡇࡢ⣽⬊ࡣࡸࡀ࡚Ṛ⁛ࡋࠊཎ⣒యࡀศ᩿໬ࡍࡿࡓࡵࠊࡑࡢ⎔ቃ࠿ࡽ⛣ືࡍࡿᶵ఍ࢆቑࡸࡍ࡜⪃࠼ࡽࢀ࡚࠸ࡿࠋࡇࡢ⬺㞳⣽⬊ࡢศ໬ࡣ㸪࣮࢜࢟ࢩࣥෆ⏕㔞ࡢపୗࡋࡓኚ␗ᰴ࡛ஹ㐍ࡋ㸪࣮࢜࢟ࢩࣥฎ⌮࡟ࡼࡾᢚไࡉࢀࡿࡇ࡜ࡀሗ࿌ࡉࢀ࡚࠾ࡾ㸦Bopp et al. 1991㸧㸪
࣍ࣥࣔࣥࢪࢦࢣ࡜ྠᵝ࡟㸪ཎ⣒య⣽⬊࡟࠾ࡅࡿ࣮࢜࢟ࢩࣥ⃰ᗘࡢపୗࡀ㸪୙㐺࡞⎔ቃ࠿ࡽࡢ㏨㑊ࡢࡓࡵࡢ⣽⬊ศ໬ㄏᑟ࡟㛵୚ࡍࡿྍ⬟ᛶࡀ♧၀ࡉࢀࡿࠋᅗ㸲㸬ࣄࣙ࢘ࢱࣥࢦࢣཎ⣒య࡟ᙧᡂࡉࢀࡓ⬺㞳⣽⬊▮ᑼ㸸⬺㞳⣽⬊, ࢫࢣ࣮ࣝࣂ࣮ࡣ, 20 ȝm㸲㸬࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆᙧᡂⱼⴥయࡸ㸪ࢮࢽࢦࢣ࡞࡝ࡢⴥ≧య࡟ᙧᡂࡉࢀࡿ↓ᛶⱆ࡛ࡣ㸪᳜≀యࡀ❧యⓗ࡛࠶ࡿࡓࡵ࡟⏕⣽⬊࢖࣓࣮ࢪࣥࢢ࡛ࡢᙧᡂ㐣⛬ࡢほᐹ࡟ࡣᢏ⾡ࡸタഛࡀᚲせ࡜࡞ࡿࠋࡑࡢࡓࡵ㸪ࡑࡢᙧᡂ㐣⛬࡟ࡘ࠸࡚ࡢ▱ぢࡣ㸪ษ∦ࢆసᡂࡋ࡚ぢࡽࢀࡓⓎ㐩ẁ㝵ูࡢീ࠿ࡽᚓࡽࢀ࡚࠸ࡿ
㸦Ligrone et al. 1996, Barnes & Land 1908㸧ࠋ୍᪉㸪⹊㢮ࡢཎ⣒య࡟ᙧᡂࡉࢀࡿ↓ᛶⱆࡢሙྜ㸪ཎ⣒యࡀ2ḟඖⓗ࡟ᡂ㛗ࡋ㸪↓ᛶⱆࡣ㏻ᖖ㸪ཎ⣒యࡢඛ➃㒊࡟ᙧᡂࡉࢀࡿࡓࡵ㸪ㄏᑟࡢ᮲௳
ࡀศ࠿ࡗ࡚࠸ࢀࡤほᐹࡀᐜ࡛᫆࠶ࡿࠋ᭱㏆㸪࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆᙧᡂ㐣⛬ࡢࢱ࢖࣒ࣛࣉࢫ᧜ᙳࢆ⾜࠸㸪↓ᛶⱆศ໬᫬ࡢືែࢆᤊ࠼ࡓࡢ࡛㸪ࡇࢀࢆ⤂௓ࡍࡿࠋ࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆࡣ㸪ศ⿣࡜ఙ㛗ᡂ㛗ࢆ⾜࠺ࢡࣟࣟࢿ࣐㡬➃ᖿ⣽⬊㸦▼ᕝ 2015㸧ࡀ㸪ศ໬ࡍࡿࡇ࡜࡛ᙧᡂࡉࢀࡿࠋᙧᡂࡢࢫ࢖ࢵࢳ㸦࣍ࣥࣔࣥࢪࢦࢣࡢሙྜ㸪ప⃰ᗘ㖡ῧຍᇵᆅ࡛ࡢᇵ㣴᮲௳㸧ࡀධࡿ࡜ศ⿣࠾ࡼࡧඛ➃ᡂ㛗ࡋ࡚࠸ࡓ㡬➃⣽⬊ࡀ㸪▷㍈᪉ྥ࡟ᑡࡋ⭾ࡽࡴ࡜ඹ࡟㸪ศ⿣ࡍࡿ㸦ᅗ5A, B, ▮ᑼ㸧ࠋࡑࡢศ⿣ᚋ㸪ඛ➃࠿ࡽぢ࡚ᚋ᪉ࡢ⣽⬊࡛ࡣ᰾ࡀ⣽⬊ࡢᇶ㒊ഃ࡟⛣ືࡋ㸪୙➼ศ⿣࡟ࡼࡾ⬺㞳⣽⬊ࡀᙧᡂࡉࢀࡿ㸦ᅗ5C, ▮ᑼ㸧ࠋ୍᪉㸪1ᅇ┠ࡢศ⿣ᚋ࡟㸪ඛ➃ഃࡢ⣽⬊ࡣఙ㛗ࡋ㸪⬺㞳⣽⬊ࡢᙧᡂ࡟㐜ࢀ࡚෌ࡧศ⿣ࡍࡿࡇ࡜࡛㸦ᅗ5D, ▮ᑼ㸧㸪3⣽⬊࠿ࡽ࡞ࡿ↓ᛶⱆࡀᙧᡂࡉࢀࡿࠋ

T. Nomura-5BSJ-Review 7:157 (2016)

植物科学最前線 7:158 (2016)ࡑࡢᚋ㸪↓ᛶⱆࡢ⣽⬊ෆᵓ㐀࡟ኚ໬ࡀ㉳ࡇࡾ㸪↓ᛶⱆࡀᡂ⇍ࡍࡿ㸦࠾ࡑࡽࡃࡇࡢ᫬࡟⃦⢊ࡸἜ⁲ࡢ⵳✚ࡀ⏕ࡌ࡚࠸ࡿ࡜᥎ࡉࢀࡿࠋ㸧ࠋ᭱ᚋ࡟㸪⬺㞳⣽⬊ࡀ⭾₶໬ࡋࡓᚋ㸪⣽⬊Ṛࢆ㉳ࡇࡍࡇ࡜࡛㸪↓ᛶⱆࡀཎ⣒య࠿ࡽษࡾ㞳ࡉࢀࡿ㸦ᅗ5F▮ᑼ㸧ࠋ↓ᛶⱆࡀษࡾ
㞳ࡉࢀࡓཎ⣒యࡣᖿ⣽⬊࡜࡞ࡾ㸪ඛ➃ఙ㛗ࢆጞࡵࡿ㸦ᅗ5G▮ᑼ㸧ࠋ↓ᛶⱆࢆ᭱㐺࡞᮲௳࡛ᇵ㣴ࡍࡿ࡜㸪ྛࠎࡢ↓ᛶⱆ⣽⬊࠿ࡽࢡࣟࣟࢿ࣐⣽⬊ࡀⓎⱆࡍࡿ㸦ᅗ2D▮༳㸧࣍ࣥࣔࣥࢪࢦࢣཎ⣒యࢆ㸪ᾮయᇵ㣴ࡋ࡚࠸ࡿ࡜㸪↓ᛶⱆ࠿ࡽⓎⱆࡋࡓཎ⣒య࡟෌ࡧ↓ᛶⱆࡀᙧᡂࡉࢀ㸪ୖ㏙ࡢ㐣⛬ࢆ⧞ࡾ㏉ࡍᵝᏊࡀほᐹࡉࢀࡿ㸦ᅗ2D▮ᑼ㸧ࠋࡇࡢࡼ࠺࡟㸪࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆᙧᡂࡢ㐣⛬࡛ࡣ㸪ᰤ㣴⦾Ṫჾᐁ࡛࠶ࡿ↓ᛶⱆ⣽⬊ࡢศ໬㸪୙➼ศ⿣࡟ࡼࡿ⬺㞳⣽⬊ࡢᙧᡂ࡜ࣉࣟࢢ࣒ࣛ⣽⬊Ṛ㸪ษࡾ㞳ࡉࢀࡓ↓ᛶⱆ࠿ࡽࡢཎ⣒య⣽⬊ࡢ෌ศ໬࡞࡝㸪ᵝࠎ࡞⣽⬊ࡢኚᐜࢆ㸪ᩘ⣽⬊࠿ࡽᵓᡂࡉࢀࡿࢩࣥࣉࣝ࡞ᙧែᙧᡂࣉࣟࢭࢫࡢ୰࡛㸪ほᐹࡍࡿࡇ࡜ࡀྍ⬟࡛࠶ࡿ㸦ᅗ6㸧ࠋ㸳㸬௒ᚋࡢᒎᮃᅗ㸳㸬࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆᙧᡂࡢࢱ࢖࣒ࣛࣉࢫ᧜ᙳ࡟ࡼࡿほᐹࢫࢣ࣮ࣝࣂ࣮ࡣ, 25 ȝm๓㏙ࡢࡼ࠺࡟࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆᙧᡂࡢ㐣⛬࡛ࡣ㸪ᵝࠎ࡞⣽⬊ศ໬ࡢ᣺ࡿ⯙࠸ࢆほᐹ࡛ࡁࡿࡓࡵ㸪⣽⬊⏕≀Ꮫ◊✲ࡢᮦᩱ࡜ࡋ࡚⯆࿡῝࠸ᮦᩱ࡛࠶ࡿ࡜⪃࠼ࡽࢀࡿࠋࡲࡓࠊ᳜≀ࡢᰤ㣴⦾Ṫჾᐁᙧᡂ࡟㛵ࡍࡿ▱ぢࢆྲྀᚓࡍࡿࡓࡵࡢᮦᩱ࡜ࡋ࡚ࡶ᭷⏝࡛࠶ࡿࠋ

T. Nomura-6
BSJ-Review 7:158 (2016)

植物科学最前線 7:159 (2016)ᅗ㸴㸬࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆᙧᡂ࡛ぢࡽࢀࡿ⣽⬊ศ໬ࡢኚᐜ࡜ᚠ⎔ⱏ㢮࡟࠾࠸࡚ࡣ㸪᪂⯆ࡢࣔࢹ᳜ࣝ≀࡛࠶ࡿࢮࢽࢦࢣࢆ⏝࠸ࡓ◊✲࠿ࡽ㸪↓ᛶⱆᙧᡂ࡜⿕Ꮚ᳜≀࡟࠾ࡅࡿ⭆ⱆᙧᡂࡢศᏊᶵᵓ㛫ࡢಖᏑᛶࡀ᫂ࡽ࠿࡟ࡉࢀࡘࡘ࠶ࡿ㸦▼ᓮ2014㸧ࠋ୍᪉㸪⹊㢮ࡢከࡃࡢ✀࡟࠾࠸࡚㸪↓ᛶⱆᙧᡂ࡟ࡼࡿ⦾Ṫࡀሗ࿌ࡉࢀ࡚࠸ࡿࡀ㸦Imura 1994㸧㸪ࡑࡢศᏊᶵᵓࡣ㸪Ṥ࡝᫂ࡽ࠿࡟࡞ࡗ࡚࠸࡞࠸ࠋࡲࡓ㸪ࢥࢣ᳜≀ࡢ↓ᛶⱆ࡟ࡣἜ⁲ࡸ⃦⢊⢏ࡢ⵳✚ࡀࡳࡽࢀ㸦Ligrone et al. 1996㸧㸪ࡑࢀࡽࡢྜᡂ࠾ࡼࡧ⵳✚ࡢศᏊᶵᵓ࡟㛵ࡍࡿ᪂▱ぢࡀᚓࡽࢀࡿྍ⬟ᛶࡶ⛎ࡵ࡚࠸ࡿࠋ⌧ᅾ㸪ᡃࠎࡣḟୡ௦ࢩ࣮ࢡ࢚ࣥࢧࢆ⏝࠸ࡓ࣍ࣥࣔࣥࢪࢦࢣࡢࢤࣀ࣒ゎㄞࢆ㐍ࡵ࡚࠾ࡾ㸪᭱㏆㸪ࢤࣀ࣒⦅㞟ᢏ⾡ࢆ⏝࠸ࡓኚ␗ᑟධἲࡢ㛤Ⓨ࡟ࡶᡂຌࡋ࡚࠸ࡿࠋࡉࡽ࡟㸪࣍ࣥࣔࣥࢪࢦࢣࡢᣢࡘ↓ᛶⱆࡢศ໬ㄏᑟࢆேⅭⓗ࡟ࢥࣥࢺ࣮࡛ࣟࣝࡁࡿ࡜࠸࠺≉ᛶ࡟╔┠ࡋ㸪↓ᛶⱆᙧᡂㄏᑟ᮲௳㸦㏻ᖖᇵᆅ㸧࡜㸪ᢚไ᮲௳㸦࣮࢜࢟ࢩࣥ㸪㧗⃰ᗘ㖡ῧຍᇵᆅ㸧㛫࡛ࡢẚ㍑ࢺࣛࣥࢫࢡࣜࣉࢺ࣮࣒ゎᯒࢆᐇ᪋ࡋࡓࠋࡑࡢ⤖ᯝ㸪↓ᛶⱆᙧᡂ᮲௳࡛Ⓨ⌧㔞ࡀஹ㐍ࡋ࡚࠸ࡓ㌿෗⏘≀⩌ࢆぢฟࡍࡇ࡜ࡀ࡛ࡁ㸪୰࡛ࡶ≉࡟㌿෗ㄪ⠇ᅉᏊࡢᶵ⬟࡟ࡘ࠸࡚⌧ᅾゎᯒࢆ㐍ࡵ࡚࠸ࡿࠋࡇࢀࡽࡢゎᯒ࠿ࡽ㸪᳜≀⣽⬊ࡢศ໬ㄪ⠇ࡸ㸪ᮍࡔከࡃࡀ᫂ࡽ࠿࡟࡞ࡗ࡚࠸࡞࠸᳜≀࡟࠾ࡅࡿᰤ㣴⦾ṪჾᐁᙧᡂࡢศᏊᶵᵓ࡟ࡘ࠸࡚㸪᪂ࡋ࠸▱ぢࡀᚓࡽࢀࡿࡇ࡜ࡀᮇᚅࡉࢀࡿࠋࡲࡓ㸪࣍ࣥࣔࣥࢪࢦࢣཎ⣒య࡟࠾ࡅࡿ↓ᛶⱆᙧᡂㄏᑟࡢୖὶᅉᏊࢆ㎺ࡿࡇ࡜࡟ࡼࡾࠊ
ࢫ࣌ࢩࣕࣜࢫࢺࡀ㸪⮬㌟ࡢ⏕ែⓗᆅ఩࡜ࡋ࡚㐺ࡋࡓ⎔ቃ࠿ྰ࠿ࢆឤ▱ࡋ㸪ࢽࢵࢳᡓ␎ࢆᐇ⌧ࡉࡏࡿ௙⤌ࡳࡢ୍➃ࢆゎ᫂ࡋࡓ࠸࡜⪃࠼࡚࠸ࡿࠋ

ㅰ㎡ᮏ✏࡛⤂௓ࡋࡓ◊✲ෆᐜࡢ୍㒊ࡣ㸪᪥ᮏᏛ⾡᣺⯆఍࣭⛉Ꮫ◊✲㈝⿵ຓ㔠㸦11J06080,15K18824㸧
㸪᪥ᮏ⛉Ꮫ༠఍࣭➲ᕝ◊✲ຓᡂ㔠㸦27-420㸧ࡢᨭ᥼ࢆཷࡅ࡚⾜ࢃࢀࡓࠋࡲࡓᮏ◊✲ࡢ㐙⾜࡟࠾࠸࡚㸪ᨭ᥼ࢆ㡬࠸ࡓⓙᵝ᪉࡟ཌࡃᚚ♩ࢆ⏦ࡋୖࡆࡿࠋ

T. Nomura-7
BSJ-Review 7:159 (2016)

植物科学最前線 7:160 (2016)

ᘬ⏝ᩥ⊩
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ࡻ࠺ࡈ⛉Ꮫᢏ⾡༠఍Ꮫ⾡◊✲ຓᡂᡂᯝሗ࿌᭩.
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㬼ἑ⨾✑Ꮚ&బ➉◊୍2010.⛛∗ᒣᆅ࡛෌Ⓨぢࡉࢀࡓ࣍ࣥࣔࣥࢪࢦࢣࡢ⬊Ꮚయཬࡧ⏕Ṫჾᐁࡢᙧᡂ࡟ࡘ࠸࡚⹊ⱏ㢮◊✲10: 10-12.
T. Nomura-8
BSJ-Review 7:160 (2016)