腺と鋤鼻器の役割に関する研究( 要約版(Digest) ) Author(s) 冨安, 洵平 Report No.(Doctoral Degree) 博士(獣医学) 甲第534号 Issue Date 2019-03-13 Type 博士論文 Version none URL http://hdl.handle.net/20.500.12099/77965 ※この資料の著作権は、各資料の著者・学協会・出版社等に帰属します。
Ꮫ ㄽ ᩥ せ ⣙
Ặ ྡ Ᏻ Ὥᖹ 㢟 ┠ ⦾Ṫᮇࡢࣄࢢ࣐Ⴅぬࢥ࣑ࣗࢽࢣ࣮ࢩࣙࣥ࠾ࡅࡿ⫼㒊⓶⭢ 㗟㰯ჾࡢᙺ㛵ࡍࡿ◊✲ ࣄࢢ࣐ࡣ⾜ືᅪෆࡢ≉ᐃࡢᮌయࢆ᧿ࡾࡘࡅ∎ṑᙧࢆࡘࡅࡿ⾜ືࢆ♧ࡍࡇࡀ▱ࡽ ࢀ࡚࠸ࡿࠋࡇࡢ⾜ືࡣձ⦾Ṫᮇቑຍࡋղᡂ⋇㞝࡛ࡣከࡃࡳࡽࢀࡿࡀճளᡂ⋇࡛ࡣໝ࠸ ࢆႥࡂࡃࡿࡶࡢࡢయࢆ᧿ࡾࡘࡅ࡞࠸࠸࠺≉ᚩࡀ࠶ࡾࣄࢢ࣐ࡢໝ࠸ࢥ࣑ࣗࢽࢣ࣮ࢩࣙ ࣥ࠾ࡅࡿ࣐࣮࢟ࣥࢢ⾜ືࡋ࡚⪃࠼ࡽࢀ࡚࠸ࡿࠋࡲࡓඹ㏻ࡋ࡚ㄆࡵࡽࢀࡿ࣐࣮࢟ࣥࢢࡢ ጼໃࡋ࡚❧࡛ࡢ⫼㒊ࢆ᧿ࡿ⾜ື ⫼᧿ࡾ ࡀ▱ࡽࢀ࡚࠸ࡿࠋᮏ◊✲࡛ࡣࣄࢢ࣐ࡢ࣐࣮࢟ ࣥࢢࡢ⾜ື≉ᛶࡽࠕ⦾Ṫᮇࡢ㞝ᡂ⋇࡛ࡣ⫼㒊࡛⓶⭢ࡀⓎ㐩ࡋࡑࡢศἪ≀ࢆᮌ᧿ࡾࡘ ࡅ࡚࠸ࡿࠖ௬ㄝࢆ❧࡚ໝ࠸≀㉁ࢆศἪࡍࡿ⓶⭢࠾ࡼࡧཷᐜࡍࡿឤぬჾࡘ࠸࡚ㄪࡓࠋ ➨ ❶࡛ࡣ◊✲ࡢጞࡵ㞝ᡂ⋇ࣄࢢ࣐⫼㒊࠾ࡅࡿศἪ≀ࡢᏑᅾࡘ࠸࡚ᮇኚࢆ ㄪࡓࠋࡑࡢ⤖ᯝ⦾Ṫᮇ㞝ࣄࢢ࣐ࡢ⫼㒊ṇ୰⥺ୖ࡛ⰾ㤶ࢆᨺࡘἜ⬡ᛶ≀㉁ࡢศἪࡀㄆࡵ ࡽࢀࡓࠋࡋࡓࡀࡗ࡚⦾Ṫᮇ࠾ࡅࡿ㞝ᡂ⋇ࣄࢢ࣐ࡢ⫼᧿ࡾ࡛ࡣ⫼㒊ࡽศἪࡉࢀࡿἜ⬡ ᛶ≀㉁ࢆᮌ᧿ࡾࡘࡅ࡚࠸ࡿࡇࡀ♧၀ࡉࢀࡓࠋḟ⫼㒊ࡢ⓶ࢆ᥇ྲྀࡋ⤌⧊Ꮫⓗ᥈⣴ࢆ ⾜ࡗࡓࠋほᐹࡢ⤖ᯝ⦾Ṫᮇ⫼㒊࡛ࡁࡃⓎ㐩ࡋࡓ⬡⭢ࡀㄆࡵࡽࢀࡓࠋ⫼㒊⬡⭢ࡣἜ⬡ ᛶ≀㉁ࡀㄆࡵࡽࢀ࡞ࡗࡓ㠀⦾Ṫᮇࡣ㏥⦰ࡋ࡚࠸ࡓࠋࡋࡓࡀࡗ࡚⫼㒊࠾ࡅࡿἜ⬡ᛶ≀ ㉁ࡢศἪࡣࡁࡃⓎ㐩ࡋࡓ⬡⭢ࡀ㛵ࡋ࡚࠸ࡿࡇࡀ♧၀ࡉࢀࡓࠋࡉࡽ⬡⭢ࡢࡁࡉ ⾑୰ࢸࢫࢺࢫࢸࣟࣥ⃰ᗘࡢ㛫ࡣ┦㛵ࡀㄆࡵࡽࢀ⬡⭢⣽⬊࡛ࡣࣥࢻࣟࢪ࢙ࣥཷᐜ యࡢⓎ⌧ࡀㄆࡵࡽࢀࡓࠋࡲࡓཤໃಶయ࡛ࡣ⦾Ṫᮇ࠾࠸࡚⫼㒊⬡⭢ࡢⓎ㐩ࡣ☜ㄆࡉࢀ࡞ ࡗࡓࠋࡑࡢࡓࡵࣄࢢ࣐ࡢ⫼㒊⬡⭢Ⓨ㐩ࡣ⦾Ṫᮇୖ᪼ࡍࡿࢸࢫࢺࢫࢸࣟࣥࡼࡾㄪ⠇ࡉ ࢀ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡓࠋ ➨ ❶࡛ࡣ㞝ࣄࢢ࣐ࡢ⫼㒊࣏ࢡࣜࣥ⭢ࡢᮇኚ࠾ࡼࡧ⫼㒊ศἪ≀ࡢ㛵ಀࢆㄪ ࡓࠋ⦾Ṫᮇࡢ↓ฎ⨨ಶయ࡛ࡣ㠀⦾Ṫᮇࡢ↓ฎ⨨ಶయ࠾ࡼࡧ⦾Ṫᮇࡢཤໃಶయẚ࡚ ࣏ࢡࣜࣥศἪࡀஹ㐍ࡋ࣏ࢡࣜࣥ⭢ࡢࢧࢬࡢቑຍࡀㄆࡵࡽࢀࡓࠋ㉸ᚤ⣽ᵓ㐀ほᐹࡢ⤖ᯝ ⦾Ṫᮇ㠀⦾Ṫᮇࡢ㛫࡛⣽⬊ෆᑠჾᐁࡢᙧែᕪ␗ࡀㄆࡵࡽࢀࡓࠋࡲࡓ࡞ศἪᵝᘧࡣ ᮇẖ␗࡞ࡾ⦾Ṫᮇࡣ࣏ࢡࣜࣥศἪࡀஹ㐍ࡋ㠀⦾Ṫᮇࡣ࢚ࢡࢯࢧࢺ࣮ࢩࢫࡢ ஹ㐍ࡀㄆࡵࡽࢀࡓࠋ࢚ࢡࢯࢧࢺ࣮ࢩࢫࡣ⣽⬊㛫┦స⏝⏝ࡉࢀ࣏ࢡࣜࣥศἪࡣእ 㒊⎔ቃᑐࡍࡿሗఏ㐩⏝ࡉࢀࡿ⪃࠼ࡽࢀ࡚࠸ࡿࠋࡋࡓࡀࡗ࡚⦾Ṫᮇࡢ㞝ᡂ⋇ࣄࢢ ࣐࠾࠸࡚⫼㒊࣏ࢡࣜࣥ⭢࡛ࡣ࣏ࢡࣜࣥศἪࡀஹ㐍ࡋໝ࠸ࢩࢢࢼࣝࢆ┒ࢇศἪࡋ ࡚࠸ࡿࡇࡀ♧၀ࡉࢀࡓࠋ௨ୖࡢ⤖ᯝࡼࡾ⬡⭢ྠᵝ࣏ࢡࣜࣥ⭢ࡶ⫼㒊ࡢἜ⬡ᛶ≀㉁ ࡢ⏘⏕㛵ࡋ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡓࠋ ➨ ❶࠾ࡼࡧ➨ ❶ࡢ⤖ᯝࡼࡾ⦾Ṫᮇ࠾࠸࡚㞝ᡂ⋇ࣄࢢ࣐ࡢ⫼㒊࡛ࡣ⬡⭢࠾ࡼࡧ࣏ ࢡࣜࣥ⭢ࡢࡕࡽࡶࡁࡃⓎ㐩ࡋศἪࡀஹ㐍ࡋ࡚࠸ࡿࡇࡀ᫂ࡽ࡞ࡗࡓࠋ✀ࡢ⓶ ⭢ࡢศἪ≀ࡣⓎᛶࡢ㐪࠸ࡽఏ㐩ࡍࡿሗࡀ␗࡞ࡿࡇࡀ᥎ᐹࡉࢀ࡚࠸ࡿࠋ㞝ᡂ⋇ࣄࢢ ࣐ࡢ⫼㒊࡛✀ࡢ⓶⭢ࡢศἪࡢஹ㐍ࡀㄆࡵࡽࢀࡓࡇࡽศἪ≀ࡢΰྜ≀࡛࠶ࡿ⫼㒊 ࡢἜ⬡ᛶ≀㉁ࡣከᵝ࡞ሗࢆఏ㐩ࡋ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡓࠋ ➨ ❶࡛ࡣ⏕Ṫ⾜ື㛵ࢃࡿໝ࠸ࡢཷᐜჾ࡛࠶ࡿ㗟㰯ჾࡘ࠸࡚ᙧែᏛゎᯒࢆ⾜ࡗࡓࠋࡢච⤌⧊ᏛᰁⰍࡢ⤖ᯝࡽࣄࢢ࣐ࡣ *ȘL ඹᙺࡍࡿ ᆺ㗟㰯ཷᐜయࢆ⏝ࡋ✵Ẽ୰ ࡢໝ࠸≀㉁ࢆཷᐜࡋ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡓࠋ௨ୖࡢ⤖ᯝࡼࡾࣄࢢ࣐࠾࠸࡚ࡶ㗟㰯ჾ ࡀᶵ⬟ⓗໝ࠸ࢆឤ▱ࡋ࡚࠸ࡿ⪃࠼ࡽࢀࡓࠋࡲࡓࣄࢢ࣐࡛ࡣ㗟㰯⭢ᮼ⣽⬊ຍ࠼࡚ከ ⣽⬊ᛶୖ⓶ෆ⭢ࡀㄆࡵࡽࢀࡑࢀࡒࢀࡢ⭢⣽⬊ࡢศἪ㢛⢏ࡢᛶ㉁ࡣ⤌⧊Ꮫⓗゎᯒ⤌⧊ Ꮫⓗゎᯒ㉸ᚤ⣽ᵓ㐀ゎᯒࡢ⤖ᯝ␗࡞ࡗ࡚࠸ࡓࠋከ⣽⬊ᛶୖ⓶ෆ⭢ࡣࡇࢀࡲ࡛ሗ࿌ࡀ࡞ ࡃ⭢⣽⬊ࡢⓎ㐩ࡣࣄࢢ࣐ࡢ≉ᚩࡔ⪃࠼ࡽࢀࡓࠋⓎ㐩ࡋࡓ㗟㰯ჾࡢᏑᅾࡣࣄࢢ࣐ࡀ㗟㰯ჾ ࢆ⏝ࡋ࡚ໝ࠸≀㉁ࢆཷᐜࡋ࡚࠸ࡿࡇࡢࡅ࡞ࡾࢡ࣐⛉ື≀࡛ほᐹࡉࢀࡿࣇ࣮ࣞ ࣓ࣥࡣ㗟㰯ჾໝ࠸≀㉁ࢆ㏦ࡾࡇࡴ⾜ືࡔ⪃࠼ࡽࢀࡓࠋ ᮏ◊✲ࡼࡗ࡚⦾Ṫᮇ࠾࠸࡚㞝ࣄࢢ࣐ࡢ⫼㒊࡛⬡⭢࠾ࡼࡧ࣏ࢡࣜࣥ⭢ࡀⓎ㐩ࡋἜ ⬡ᛶ≀㉁ࡢศἪࡀㄆࡵࡽࢀࡓࡇࡽ㞝ᡂ⋇ࡀ⦾Ṫᮇໝ࠸ࢩࢢࢼࣝࢆ⏝ࡋ✚ᴟⓗ ሗⓎಙࢆ⾜࠺ࡇࡀ♧၀ࡉࢀࡓࠋࡉࡽࣄࢢ࣐ࡀᙧែᏛⓗⓎ㐩ࡋࡓ㗟㰯ჾࢆᣢࡗ࡚࠸ ࡓࡇࡽศἪ≀ྵࡲࢀࡿໝ࠸≀㉁ࢆឤཷࡋ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡓࠋ௨ୖࡢ⤖ᯝࡼ ࡾ㞝ᡂ⋇ࣄࢢ࣐ࡢ⫼㒊ࡢศἪ≀ࡣໝ࠸ࢥ࣑ࣗࢽࢣ࣮ࢩࣙࣥ⏝ࡉࢀ࡚࠸ࡿ⪃࠼ࡽࢀ ࡓࠋ
Ꮫ ㄽ ᩥ せ ⣙
Ặ ྡ 720,<$68-XPSHL 㢟 ┠6WXGLHVRQ5ROHVRI%DFN6NLQ*ODQGVDQG9RPHURQDVDO2UJDQV ,QYROYHGLQ2OIDFWRU\&RPPXQLFDWLRQDPRQJ%URZQ%HDUVGXULQJWKH %UHHGLQJ6HDVRQ ⦾Ṫᮇࡢࣄࢢ࣐Ⴅぬࢥ࣑ࣗࢽࢣ࣮ࢩࣙࣥ࠾ࡅࡿ⫼㒊⓶⭢ 㗟㰯ჾࡢᙺ㛵ࡍࡿ◊✲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Ș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
