੕ହਯ৶धৰୡदचऽॊে੟भৎ৑भ⎚హ᫖ ੪叞৶৲৾ଢ଼஢ਚ ਯ৶൉ୗউটॢছ঒(iTHEMS)

20 ୡ⣖ᚋ༙࠿ࡽ⌧ᅾࡲ࡛ࠊ⏕≀ࣜࢬ࣒࡟㛵ࡍࡿ㑇ఏᏊࡸࢱࣥࣃࢡ㉁ࡢ◊✲ࡣ㦫␗ⓗ࡟㐍ࢇ࡛ࡁࡓࠋ⌧ᅾࠊ㑇ఏᏊࡸࢱࣥࣃࢡ㉁࡟ࡘ࠸࡚ࡢ▱ぢࢆᩘ⌮ⓗ࢔ࣉ࣮ࣟࢳ࡟ࡼࡗ࡚⤫ྜࡍࢀࡤࠊ㑇ఏᏊࡸࢱࣥࣃࢡ㉁ࡢࡇ࡜
ࡀࢃ࠿ࡿ๓࠿ࡽ▱ࡽࢀ࡚࠸ࡓ⏕≀ࡢ᫬㛫ࡢㅦ࡟ࡏࡲࢀࡿ࠿ࡶࡋࢀ࡞࠸ࠋࡇࡇ࡛ࡣ୺࡟ࠊయෆ᫬ィࡢൾᛶࡢᩘ⌮ⓗ◊✲ࢆ⤂௓ࡋࠊ᳨ドྍ⬟࡞ணᗘ⿵ࢆᑟࡅࡿࡇ࡜ࢆ♧ࡍࠋ≉࡟ࠊ᣺ືࡢ᣺ᖜࡸἼࡢᙧ࡟ࡘ࠸࡚⌮ㄽⓗ⪃ᐹࢆ⾜ࡗ࡚ࠊ ᗘ⿵ൾᛶࡢࡳ࡞ࡽࡎࠊ⏕≀ࣜࢬ࣒ࡢ໬ྜ≀ࢫࢡ࣮ࣜࢽࣥࢢ࡞࡝࡟࠾࠸࡚᣺ືࡢ᫬⣔ิࡢ᝟ሗࢆࡼࡾ⏕࠿ࡍ᪉ἲࡶ㆟ㄽࡍࡿࠋ1.

ࡣࡌࡵ࡟ࡤࣖࣜ࢖࢝ࡢάື㟁఩ࡣࠊ38ᗘ࡟࠾࠸࡚㟁఩ࡢቑ࠼ࡿ

⮬↛⏺࡟ࡣᵝࠎ࡞ࣜࢬ࣒ࡀ࠶ࡿࠋ⿕㣗⪅࡜ᤕ㣗⪅ࡢ᫬㛫ᖏ࡜ῶࡿ᫬㛫ᖏࡢ㛗ࡉࡀ࡯ࡰྠࡌ࡛࠶ࡿࡀࠊ21ᗘ㛵ಀ࡟࠶ࡿ࢝ࣥࢪ࢟࢘ࢧࢠ࡜࣐࢜࢜ࣖࢿࢥࡢಶయᩘ࡟࠾࠸࡚ࡣ㟁఩ࡢῶࡿ᫬㛫ᖏࡢ᪉ࡀࡎࡗ࡜㛗ࡃ㠀ᑐࡣ⣙10ᖺࡢ࿘ᮇ࡛఩┦ࢆࡎࡽࡋ࡚ኚືࡋ1ࠊࣖࣜ࢖࢝

⛠࡞Ἴᙧ࡛࠶ࡿ2ࠋࡇࡢࡼ࠺࡟⮬↛࡟ࡣⰍࠎ࡞Ἴᙧࡀ

ࡢᕧ኱⚄⤒㍈⣴ࡢάື㟁఩ࡣᩘ࣑ࣜ⛊ࡢ࣮࢜ࢲ࣮࡛

࠶ࡿࠋኚືࡍࡿࠋࡇࡢࡼ࠺࡟ࣜࢬ࣒ࡢ࿘ᮇࡣᵝࠎ࡛࠶ࡿࠋࡲ18ୡ⣖࡟Joseph FourierẶࡣࠊ඲࡚ࡢࣜࢬ࣒ࡣ୕ゅࡓ࿘ᮇࡔࡅ࡛࡞ࡃࠊࠕἼࡢᙧࠖࡶⰍࠎ࠶ࡿࠋᙧࡀ୸࠸

㛵ᩘࡢ㊊ࡋ⟬࡛⾲⌧࡛ࡁࡿࡇ࡜ࢆぢฟࡋࡓࣜࢬ࣒ࡶ࠶ࢀࡤࠊ▴ᙧἼ≧ࡢࣜࢬ࣒ࡶ࠶ࡿࠋἼࡢᙧࡀ㸦σ௝ ܾ௝ sin ߱௝ ‫ ݐ‬+ ܿ௝ cos ߱௝ ‫ ݐ‬㸧3ࠋࡈᏑ▱ࡢࡼ࠺࡟ࠊࣇ࣮ᑐ⛠ࡢࣜࢬ࣒ࡶ࠶ࢀࡤࠊ㠀ᑐ⛠ࡢࣜࢬ࣒ࡶ࠶ࡿࠋ౛࠼

࢚ࣜゎᯒࢆ౑࠼ࡤࣜࢬ࣒ࡢ࿘ᮇࢆヲࡋࡃㄪ࡭ࡽࢀࡿࠋ

ᅗ 1 Ἴᙧࡢࡦࡎࡳࢆᐃ㔞໬ࡍࡿ࢖ࣥࢹࢵࢡࢫࠋ᫬⣔ิࡀࢧ࢖ࣥ㛵ᩘ࠿ࡽ㞳ࢀ࡚࠸ࡿ⛬ᗘࢆ⾲ࡍࠋྑୗࡣࣇ࣮࢚ࣜ⣭ᩘࡢ」⣲ᩘ⾲♧ࠋᩥ⊩ 4 ࢆᨵኚࠋ‫ݩ‬g.kurosawa@riken.jp
᫬㛫⏕≀Ꮫ
14

Vol. 26, No. 1 (2020)ࠕⰍࠎ࡞Ἴࡢᙧࡶ࿘ᮇࡢࡼ࠺࡟୍ࡘࡢ㔞࡛⾲ࡏ࡞࠸ࡔࢁ࠺࠿㸽ࠖᐇࡣྍ⬟࡛࠶ࡿࠋ඲࡚ࡢἼࡣ୕ゅ㛵ᩘࡢ㊊ࡋ⟬࡛⾲⌧࡛ࡁࡿࡇ࡜࠿ࡽࠊ୕ゅ㛵ᩘ࡟ࡘ࠸࡚࠸ࡿಀᩘࢆ౑ࡗ࡚ࠊ࿘ᮇࡔࡅ࡛࡞ࡃἼࡢᙧࡶᐃ㔞໬࡛ࡁࡿࠋ౛࠼ࡤࠊ᫬⣔ิࡀࠕࢧ࢖ࣥ㛵ᩘ࠿ࡽ㞳ࢀ࡚࠸ࡿ⛬ᗘ(non-sinusoidal power, NS)ࠖࡣᅗ1ࡢࡼ࠺࡞࢖ࣥࢹࢵࢡࢫ࡛⾲ࡏࡿࡇ࡜ࡀࢃ࠿ࡗ࡚࠸ࡿ4ࠋ᫬⣔ิࡀࢧ࢖ࣥ
࡛࠶ࢀࡤ1ࠊࢧ࢖ࣥ࠿ࡽ㞳ࢀࢀࡤᚲࡎ1ࡼࡾ኱ࡁࡃ࡞ࡿ㔞࡛࠶ࡿࠋ࢖ࣥࢹࢵࢡࢫࡢjࡣ᫬⣔ิ࡟ྵࡲࢀࡿ࿘Ἴ
ᩘࡢᩘࢆ⾲ࡍࠋࡇࡢࡼ࠺࡞࢖ࣥࢹࢵࢡࢫࢆᡭࡀ࠿ࡾ࡟Ἴࡢᙧࡢព࿡ࡀศ࠿ࡿ࠿ࡶࡋࢀ࡞࠸ࠋ࠸ࡁ࡞ࡾᩘᘧࡢㄝ᫂࠿ࡽጞࡵ࡚ࡋࡲࡗࡓࡀࠊ⚾ࡣᩘ⌮ࣔࢹࣝࢆ⏝࠸࡚యෆ᫬ィࡸ෤╀ࡢ◊✲ࢆࡋ࡚࠸ࡿࠋᩘ⌮ࣔࢹࣝࡢୡ⏺࡟ࡣࠊ͆All models are wrong. But
some are useful”࡜࠸࠺⤫ィᏛ⪅George BoxẶࡢゝⴥࡀ࠶ࡿࠋᩘ⌮ࣔࢹࣝࡀ⮬↛⌧㇟ࢆ࡝ࡢࡼ࠺࡟グ㏙ࡋᅗ 2 ᗘ⿵ൾᛶࡢ ᗘ-᣺ᖜ࢝ࢵࣉࣜࣥࢢ௬ㄝࠋᗘࡀୖࡀࡗ࡚཯ᛂࡀ㏿ࡃ࡞ࢀࡤయෆ᫬ィࡢ࿘ᮇࡣ▷ࡃ࡞ࡾࡸࡍ࠸ࠋ ᗘୖ᪼࡜࡜ࡶ࡟᣺ᖜࢆ኱ࡁࡃࡍࡿࡇ࡜࡛ࠊᗘ࡟ᙳ㡪ࡉࢀࡎ࡟᫬ࢆ้ࡵࡿࠋ

ࡓ࡜ࡋ࡚ࡶࠊ࠶ࡃࡲ࡛㏆ఝ࡛࠶ࡿࠋࡑࡢⅬ࡛඲࡚ࡢࣔࢹࣝࡀཝᐦ࡟ࡣ㛫㐪ࡗ࡚࠸ࡿࡢ࡛࠶ࡿࡀࠊᩘ⌮ࣔࢹࣝ

ࣥࣃࢡ㉁ࡢࣜࣥ㓟໬࡟ࡼࡗ࡚ᴫ᪥ࣜࢬ࣒ࡣ⏕ࡌࡿ࡜ࡢ୰࡟ࡣᙺ❧ࡘࡶࡢࡶ࠶ࡿ࡜࠸࠺ព࿡࡛࠶ࡿࠋ⚾ࡣࠊࡍࡿࠋᩘ⌮ࣔࢹࣝࡢ࿘ᮇࢆࢥࣥࣆ࣮ࣗࢱ࣮ୖ࡛ㄪ࡭ࡿ

2011ᖺ࡟Michael MenakerẶࡀ㔠ἑࡢ⛬⫕Ặࡢ◊✲࡜ࠊ ᗘࡀୖࡀࡗ࡚཯ᛂࡀ㏿ࡃ࡞ࡿ࡜࿘ᮇࡣ▷ࡃ࡞ࡾᐊࢆゼၥࡉࢀࡓ࡜ࡁ࡟࠾ヰࡋࡉࡏ࡚ࡶࡽࡗ࡚ࠊࡇࡢゝࡸࡍ࠸ࡇ࡜ࡀࢃ࠿ࡿࠋ౛࠼ࡤࠊ඲࡚ࡢ཯ᛂ㏿ᗘࡀྠ᫬ⴥࢆᩍࢃࡗࡓࠋ᭱ึࡣⱥㄒࡀ⪺ࡁྲྀࢀ࡞ࡃ࡚ࠊࠕࡼࡃ࡟2ಸ࡜࡞ࢀࡤࠊᚲࡎ࿘ᮇࡣ༙ศ࡟࡞ࡿࠋࡋ࠿ࡋࠊ཯ࢃ࠿ࡽ࡞࠸ࡀࠊ࠶࡞ࡓࡢᩘ⌮ࣔࢹࣝࡣwrong࡜ゝࡗ࡚ᛂ࡟ࡼࡗ࡚㏿ᗘࡢ࠸ࡿࡼ࠺ࡔࠋึᑐ㠃࡛ఱࡶㄝ᫂ࡋ࡚࡞࠸ࡢ࡟ࠊ࡞ࡐࡀ㏿ࡃ࡞ࡗࡓ࡜ࡋ࡚ࡶ࿘ᮇࡀ࡯࡜ࢇ࡝ኚ໬ࡋ࡞࠸ሙwrong࡜ゝ࠺ࡢࡔࢁ࠺ࠋࡲࡎ࠸࡞࠵ࠖ࡜ᛮࡗࡓࡢࢆぬྜࡀ࠶ࡿࠋ⚾ࡓࡕࡣලయⓗ࡞ᩘ⌮ࣔࢹࣝࢆ⏝࠸ࠊ࿘ᮇ࠼࡚࠸ࡿࠋᙼࡣゝⴥࡢព࿡ࢆࡺࡗࡃࡾᩍ࠼࡚ࡃࢀࡓࠋࢆᏳᐃ࡟ࡍࡿࡓࡵ࡟ࡣࠊ㧗ᮏ✏࡛ࡣࠊ⡆༢࡞ᩘ⌮ࣔࢹࣝࢆ⏝࠸࡚ලయⓗ࡞ணࡀ࠶ࡿࡇ࡜ࢆ♧ࡋࡓ12ࠋࡀ࡛ࡁࡿࠊ࡜࠸࠺ヰࢆ⤂௓ࡋࡓ࠸ࠋᗘ౫Ꮡᛶࡀ␗࡞ࢀࡤࠊ඲࡚ࡢ཯ᛂ

࡛᣺ᖜࡀ኱ࡁࡃ࡞ࡿᚲせ࡞ࡐ᣺ᖜࡀ㔜せ࡞ࡢ࠿㸽ୖグࡢࡼ࠺࡟㧗࡛཯ᛂࡀ㏿ࡃ࡞ࡿ࡜ࡍࡿ࡜࿘ᮇࡀ▷ࡃ࡞ࡿഴྥࡀ࠶ࡿࠋࡋ࠿
2.

ᩘ⌮ࣔࢹ࡛ࣝࡏࡲࡿ

ᗘ⿵ൾᛶࡢㅦࡋࠊ ᗘ࡜࡜ࡶ࡟ྠ᫬࡟᣺ᖜࡀ኱ࡁࡃ࡞ࡿ࡜ࠊ㑇ఏᏊࡈᏑ▱ࡢࡼ࠺࡟ࠊయෆ᫬ィ࡟ࡘ࠸࡚ࡣ㑇ఏᏊࡸࢱࣥ

࡜ࢱࣥࣃࢡ㉁ࡢࢲ࢖ࢼ࣑ࢡࢫࡀࡵࡄࡿ㌶㐨ࡀ㛗ࡃ࡞ࣃࢡ㉁ࡢ◊✲ࡀ㐍ࡳࠊከࡃࡢࡇ࡜ࡀ᫂ࡽ࠿࡟ࡉࢀ࡚ࡁࡿࡢ࡛࿘ᮇࡢ▷⦰ຠᯝࢆ┦ẅࡋࠊ࿘ᮇࢆᏳᐃ໬ࡍࡿ

ࡓ5ࠋࡑࡢ୰࡛ࠊ

㸦ᅗ2㸧ࠋ௨ୖࡢゎᯒ࠿ࡽࠊ㑇ఏᏊάᛶࣜࢬ࣒ࡀ㧗ᗘ⿵ൾᛶࡣ㔜せ࡞ᮍゎỴၥ㢟࡛࠶ࡿࠋ ᗘ⿵ൾᛶ࡜ࡣࠊ ᗘࡀኚࢃࡗ࡚ࡶయෆ᫬ィࡢ࿘࡛㧗᣺ᖜ࡟࡞ࡿࡇ࡜ࢆண

ᮇࡀ࡯࡜ࢇ࡝ኚ໬ࡋ࡞࠸⌧㇟࡛࠶ࡿ6-11ࠋ⏕యෆࡢ཯ྐẶࡢࢢ࣮ࣝࣉࡀࣛࢵࢺ⏤᮶ࡢᇵ㣴⣽⬊࡛᫬ィ㑇ఏᛂ㏿ᗘࡢከࡃࡣࠊ

Ꮚࡢάᛶࣜࢬ࣒ࢆィ

ᗘࡀ10ᗘୖࡀࡿ࡜2~3ಸ࡟࡞ࡿࠋ

ࢩࢫࢸ࣒ࡢᵓᡂせ⣲ࡀ㧗࡛㏿ࡃ࡞ࡿࡢ࡞ࡽ࿘ᮇࡶ

▷ࡃ࡞ࡾࡑ࠺࡞ࡢ࡟ࠊ࿘ᮇࡀ࡯࡜ࢇ࡝ኚࢃࡽ࡞࠸ࡢࡣ࡞ࡐࡔࢁ࠺ࠋ

ࡋࡓ࡜ࡇࢁࠊணࡢ㑇ఏᏊࡢ᣺ᖜࡣ㧗㏻ࡾ࡯࡜ࢇ࡝࡛ ኱ ࡁ ࡃ ࡞ ࡗ ࡚ ࠸ ࡓ 12 ࠋ

Reverb-ǂ ࡢࡼ࠺࡟᣺ᖜࡀኚ໬ࡋ࡞࠸㑇ఏᏊࡶ࠶ࡗ
ࡓࡀࠊPer2 ࡸCry1 ࡞࡝ከࡃࡢ㑇ఏᏊ࡛άᛶࣜࢬ࣒ᗘ⿵ൾᛶࡢศᏊ࣓࢝ࢽࢬ࣒ࢆ⌮ゎࡍࡿࡓࡵࠊࠕయෆ᫬ィࡢⓎ᣺࡟㛵ࢃࡿ඲࡚ࡢ཯ᛂࡀ㧗ࡋࡓࠋඹྠ◊✲⪅ࡢ㔜ྜྷᗣ࡛㏿ࡃ࡞ࡿ࡜ࡁࠊ࿘ᮇࡀᏳᐃ࡟࡞ࡿ᮲௳ࡣఱ࠿ࠖ࡜࠸࠺ᩘᏛࡢၥࡢ᣺ᖜࡣᗘ࡜࡜ࡶ࡟኱ࡁࡃ࡞ࡗࡓ㸦ᅗ3㸧ࠋ⯆࿡῝࠸ࡇ࡜࡟ࠊPer2-luc ࡢ㑇ఏᏊάᛶࣜࢬ࣒ࡢ᣺ᖜࡣ㧗࡛ᑠࡉࡃ࡞ࡿࠋࡇࢀࡣLuciferaseάᛶࡀ㧗࡛పࡃ㢟࡟⨨ࡁ᥮࠼࡚⪃࠼ࡿࡇ࡜࡟ࡍࡿࠋ඲࡚ࡢ཯ᛂࡀ㧗࡞ࡿࡓࡵ࡛࠶ࡿࡼ࠺ࡔࠋ᭱㏆ࠊఀ⸨ᾈྐẶࡽ࡟ࡼࡗ࡚

࡛㏿ࡃ࡞ࡿ࡜௬ᐃࡍࡿ᰿ᣐࡣ࡞࠸ࡀࠊ⡆༢ࡢࡓࡵࡇࡢࢩ࢔ࣀࣂࢡࢸࣜ࢔࡛ࡶ㧗ࡼ࠺࡟ࡍࡿࠋࡲࡓࠊࢩࣙ࢘ࢪࣙ࢘ࣂ࢚ࡸ࣐࢘ࢫ࡟࠾ࡅ

ࡁࡃ࡞ࡿࡇ࡜ࡀሗ࿌ࡉࢀ࡚࠸ࡿ13ࠋࡶࡋ࠿ࡋࡓࡽࠊ᣺ࡿᐇ㦂⤖ᯝ࡟ࡶ࡜࡙ࡁ5ࠊ᫬ィ㑇ఏᏊࡢ㌿෗ไᚚࡸࢱᖜ࡜᫬㛫⏕≀Ꮫ࡛ᴫ᪥ࣜࢬ࣒ࡢ᣺ᖜࡀ኱ᗘࡢ࢝ࢵࣉࣜࣥࢢࡣከࡃࡢ✀ࡢ࿘ᮇࡢᏳᐃᛶ

Vol. 26, No. 1 (2020)15

ᅗ 3 ࣛࢵࢺᇵ㣴⣽⬊࡟࠾ࡅࡿ᫬ィ㑇ఏᏊ
Ⓨ⌧ࡢ࿘ᮇⓗኚ໬࡜ࡑࡢ ᗘ౫Ꮡᛶࠋᩥ⊩12 ࢆᨵኚࠋ࡟㔜せ࡞ࡢ࠿ࡶࡋࢀ࡞࠸8,12ࠋఱࡀ㔜せ࡞ࡢ࠿ࢃ࠿ࡽ࡞࠸㠃ࡶ࠶ࡿࠋࡑࡇ࡛඲࡚ࡢ㑇ఏᏊࡸࢱࣥࣃࢡ㉁ࢆྵࡴ့ங㢮యෆ᫬ィࢩ࣑࣮ࣗࣞ

3.

యෆ᫬ィࡢᩘ⌮ࣔࢹࣝࡢ௬ᐃࡣṇࡋ࠸࠿㸽ᗘ⿵ൾᛶ࡟ᚲせ࡛࠶⤂௓ࡍࡿయෆ᫬ィࡢ⡆༢ࣔࢹࣝ㸦ᅗ4㸧ࡣࠊ኱ࡁࡃ

ࡉࢀࠊᐇ㝿࡟⣽⬊ࡢᐇ㦂࡛㑇ఏᏊάᛶࣜࢬ࣒

ศࡅ࡚2ࡘࡢ㒊ศ࠿ࡽ࡞ࡿ4ࠋ୍ࡘࡣࢱࣥࣃࢡ㉁࡟ࡼࡿ

ᩘ⌮ࣔࢹ࡛ࣝ᣺ᖜࡢㄪᩚࡀ
ࡿ࡜ணࢱ࣮ࢆ༢⣧໬ࡋࡓࠊ⡆༢࡞ࣔࢹࣝࢆ⏝࠸ࡿࠋࡢ᣺ᖜࡣ኱ࡁࡃ࡞ࡗ࡚࠸ࡓࠋࡇࢀ࡛ᗘ⿵ൾᛶࡢၥ㢟㑇ఏᏊάᛶࡢไᚚࠊࡶ࠺୍ࡘࡣࢱࣥࣃࢡ㉁ࡢከẁ㝵ࣜࡣゎỴ࡜ゝ࠼ࡿࡔࢁ࠺࠿ࠋ᭱ึ࡟㏙࡭ࡓࡼ࠺࡟ࠊᩘ⌮ࣥ㓟໬࡛࠶ࡿࠋ့ங㢮࡛ࡣࠊ⩻ヂᚋࡢPER2ࢱࣥࣃࢡࣔࢹࣝࡣ࠶ࡃࡲ࡛⌧㇟ࡢ㏆ఝ࡛࠶ࡿࠋࡓ࡜࠼ᩘ⌮ࣔࢹ㉁࠿ࡽࡓࡃࡉࢇࡢࣜࣥ㓟໬ࢱ࢖ࣉ࡜」ྜయࡀసࡽࢀ

ࣝࡢண㏻ࡾࡢ⤖ᯝࡀᐇ㦂࡛ᚓࡽࢀࡓ࡜ࡋ࡚ࡶࠊၥ㢟ࡿࡢ࡛ࠊࣔࢹ࡛ࣝࡣከẁ㝵ࣜࣥ㓟໬ࣉࣟࢭࢫ࡜ࡋ࡚⾲ࢆゎỴࡋࡓ࡜ࡣ㝈ࡽ࡞࠸ࠋࡑࡶࡑࡶࣔࢹࣝࡢ௬ᐃ࡜ࡋ⌧ࡍࡿࡇ࡜࡟ࡍࡿࠋ඲㒊࡛4✀㢮ࡢࢱࣥࣃࢡ㉁࠿ࡽ࡞

࡚ࠕ඲࡚ࡢ཯ᛂࡣ㧗࠸ᗘ࡛㏿ࡃ࡞ࡿࠖ࡜ࡋࡓࡀࡇࢀ

ࡿࠋmRNAࡢࢲ࢖ࢼ࣑ࢡࢫࢆ↓どࡋࠊmRNA࠿ࡽࡍࡄࡣṇࡋ࠸ࡢࡔࢁ࠺࠿ࠋ⌧ᐇ࡟ࡣࢩ࢔ࣀࣂࢡࢸࣜ࢔

࡟ࢱࣥࣃࢡ㉁ࡀ࡛ࡁࡿ࡜ࡋࡓࠋࡇࡢ⡆༢࡞ᩘ⌮ࣔࢹࣝ

KaiC ࢱ ࣥ ࣃ ࢡ ㉁ ࡢ ATP ά ᛶ ࡸ 9 ࠊ ࣐ ࢘ ࢫ ࡢ 㓝 ⣲ࡣࠊࡼࡃ◊✲ࡉࢀ࡚࠸ࡿGoodwinࣔࢹࣝ࡜࡯࡜ࢇ࡝ྠ

CK1e/dࡢPER2ࢱࣥࣃࢡ㉁࡟ᑐࡍࡿάᛶ10ࡣࠊ࡜ࡶ࡟

ࡌᙧࢆࡋ࡚࠸ࡿ15ࠋ኱ࡁ࡞㐪࠸ࡣGoodwin࡛ࡣࠊ㌿෗࣭ᗘ࡟౫Ꮡࡋ࡞࠸ࡇ࡜ࡀሗ࿌ࡉࢀ࡚࠸ࡿࠋࡇࢀࡽࡢ஦

⩻ヂไᚚࡢ㡯ࢆᢚไࢱࣥࣃࢡ㉁㸦x4㸧ࡢῶᑡ㛵ᩘ࡜ࡋᐇ࡜ᩘ⌮ࣔࢹࣝࡢ௬ᐃࡣ▩┪ࡍࡿࠋ௬ᐃ࡟㛫㐪࠸ࡀ࠶ࡓࡢ࡟ᑐࡋࠊࡇࡇ࡛ࡣ᣺ືࡍࢀࡤ㛵ᩘࡢᙧࡣఱ࡛ࡶⰋࡗࡓࡢ࡞ࡽࠊ᣺ᖜࡣ

ᗘ⿵ൾᛶ࡟㔜せ࡛ࡣ࡞࠸ࡢࡔࢁ࠸࡜ࡋ࡚࠸ࡿ࡜ࡇࢁ࡛࠶ࡿࠋ㌿෗ࢆ࡝ࡢࡼ࠺࡞㛵ᩘ࡛࠺࠿ࠋࢩ࢔ࣀࣂࢡࢸࣜ࢔ࡸ࣐࢘ࢫ࡛♧ࡉࢀࡓ୍㒊ࡢ཯᭩ࡃ࡭ࡁ࠿ᮏᙜࡢ࡜ࡇࢁࡣࢃ࠿ࡽ࡞࠸ࡢ࡛ࠊ⮬⏤ᗘࢆᛂࡢᗘ㠀౫Ꮡᛶ࡟ᑐᛂࡍࡿࡓࡵࠊḟࡢࡼ࠺࡟ၥ㢟タᣢࡓࡏࡿ᪉ࡀⰋ࠸࡜⪃࠼ࡓ㸦f(x4)㸧ࠋヰࡀࡸࡸ⣽࠿ࡃᐃࢆኚ࠼ࡿࡇ࡜࡟ࡍࡿ㸸ࠕయෆ᫬ィࡢⓎ᣺࡟㛵ࢃࡿ཯࡞ࡗ࡚ࡋࡲࡗࡓࡀࠊᩘ⌮ࣔࢹࣝ࡟ࡘ࠸࡚㐺ษ࡟ࣃ࣓ࣛ

ᛂ࡟ࡣᗘ࡟౫Ꮡࡋ࡞࠸ࡶࡢࡀ࠶ࡿࠋࡑࢀ௨እࡢ཯ᛂ

࣮ࢱࢆ㑅࡭ࡤ᣺ືࡀ㉳ࡁࡿࠋࡣ㧗࠸ᗘ࡛㏿ࡃ࡞ࡿ࡜ࡍࡿ࡜ࠊ࿘ᮇࡀᏳᐃ࡟࡞ࡿ᮲

௳ࡣఱ࠿㸽ࠖᐇࡣᩘ⌮ࣔࢹࣝࡀ⡆༢࡞ࡢ࡛ࠊ࿘ᮇࢆゎࡃࡇ࡜ࡀ࡛
ࡁࡿ4,16ࠋࠕゎࡃࡇ࡜ࡀ࡛ࡁࡿࠖ࡜ࡣࢩ࣑࣮ࣗࣞࢩࣙࣥࢆࡋ࡞ࡃ࡚ࡶ࿘ᮇࡢ᝟ሗࡀᚓࡽࢀࡿ࡜࠸࠺ࡇ࡜࡛࠶

4.

ᗘ⿵ൾᛶ࡟Ἴࡢᙧࡀ㛵ಀ㸽ࡿࠋᘧࡢ⣽࠿࠸ᑟฟࡀᮏ㉁࡛ࡣ࡞࠸ࡢ࡛ヲࡋࡃ㏙࡭࡞ᗘ⿵ൾᛶࡢ᮲௳ࢆ⪃࠼ࡿࡓࡵࠊලయⓗ࡟ᩘ࠸ࡀࠊࡲࡎ୍␒ୗࡢᘧ࡛hx3ࢆᕥ㎶࡟ࠊdx4/dtࢆྑ㎶࡟⌮ࣔࢹࣝࢆ♧ࡋࡘࡘヰࢆ㐍ࡵࡿࠋ࡞࠾ᩘᘧࡢヲ⣽࡟ࡇ࠺ࡘࡍࠋࡍࡿ࡜ࢱࣥࣃࢡ㉁x3ࢆࢱࣥࣃࢡ㉁x4ࡢᘧ࡛ᾘࡔࢃࡿᚲせࡣ࡞ࡃࠊ⚾ࡓࡕࡢᩘ⌮ࣔࢹࣝ࠿ࡽఱࡀࢃ࠿ཤ࡛ࡁࡿࠋࡇࡢ᧯సࢆ⧞ࡾ㏉ࡍ࡜4ኚᩘࡢᩘ⌮ࣔࢹࣝࡿࡢ࠿࠾ఏ࠼࡛ࡁࢀࡤ࡜ᛮ࠺ࠋ20ୡ⣖ᚋ༙࠿ࡽయෆ᫬

ࡀࢱࣥࣃࢡ㉁x4ࡔࡅࡢ1ኚᩘࣔࢹࣝ࡟࡞ࡿࠋḟ࡟᫬ィィ࡟㛵ࢃࡿⓎぢࡀ⵳✚ࡉࢀࡿ࡟ࡘࢀ࡚ࠊヲ⣽࡞ࢩ࣑ࣗ

ࢱࣥࣃࢡ㉁ࡢኚືࡣ୕ゅ㛵ᩘࡢ㊊ࡋ⟬࡛⾲⌧࡛ࡁࡿ࣮ࣞࢱ࣮ࡶᥦ᱌ࡉࢀ࡚ࡁࡓࠋࢩ࣑࣮ࣗࣞࢱ࣮ࡣ⌧ᐇࡢࡣࡎ࡞ࡢ࡛ࠊࢱࣥࣃࢡ㉁x4࡟ࣇ࣮࢚ࣜ⣭ᩘࢆ௦ධࡍࡿࠋ㑇ఏᏊࡢ᣺ࡿ⯙࠸ࢆ෌⌧ࡋࡓࡾࠊኚ␗యࡢ᣺ࡿ⯙࠸ࢆࡇ࠺࠸ࡗࡓ᧯సࢆ࠸ࡃࡘ࠿ᐇ⾜ࡍࡿ࡜ࠊḟࡢ࿘ᮇࡀฟ

ᨵࡵ࡚ண࡛ࡁࡓࡾࣃ࣡ࣇ࡛ࣝ࠶ࡿ14ࠋ୍᪉࡛ࠊ」㞧ࡍࡂ࡚᫬㛫⏕≀Ꮫ
16Vol. 26, No. 1 (2020)࡚ࡃࡿࠋᅗ 4: యෆ᫬ィࢩ࣑࣮ࣗࣞࢱ࣮࡜ࡑࡢ༢⣧໬ࣔࢹࣝࠋ့ங㢮࡛ࡣࠊ⩻ヂᚋࡢ PER2 ࢱࣥࣃࢡ㉁࠿ࡽࡓࡃࡉࢇࡢࣜࣥ㓟໬ࢱ࢖ࣉ࡜」ྜయࡀసࡽࢀࡿࠋලయⓗ࡞㑇ఏᏊࢆྵࡴࢩ࣑࣮ࣗࣞࢱ࣮ࡢゎᯒࡶ᭷⏝࡛࠶ࡿࡀࠊపḟඖࡢࣔࢹࣝࡣゎᯒࡋࡸࡍ࠸ࠋ⡆༢ࣔࢹࣝ࡟࠾࠸࡚ࠊ཯ᛂ࣮ࣞࢺࡢ b ࡣ PER2 ࢱࣥࣃࢡ㉁ࡢ bTRCP ࢧ࢖ࢺࠊp ࡣ FASPS priming ࢧ࢖ࢺࠊh ࡣ FASPS ࢡࣛࢫࢱ࣮ࡢ௚ࡢࢧ࢖ࢺࡢࣜࣥ㓟໬࡟ᑐᛂࡍࡿ࡜௬ᐃࡍࡿࠋ

࿘ᮇ = 2ߨ ൥ଵΤଶἼᙧࡢࡦࡎࡳ(ܰܵ)཯ᛂ࣮ࣞࢺࡢ2ḟᘧ൫ಀᩘࡣ඲࡚ṇ൯൩

compensation㸧6ࡸᗘࢧ࢖ࢡࣝ࡬ࡢᘬࡁ㎸ࡳࢆ෌⌧࡛ࡁ࡞࠸࡜ᛮࢃࢀࡿࠋࡇࡢࡼ࠺࡟ᩘ⌮ࣔࢹࣝࢆ⏝࠸ࡿ࡜ࠊ ᗘ౫Ꮡⓗ࡞Ἴᙧࡢኚ໬ࡀᗘ⿵ൾᛶ࡟㔜せ࡛࠶࿘ᮇࡀࠕἼᙧࡢࡦࡎࡳࠖࡢ࣮ࣝࢺ࡟ẚ౛ࡋ࡚࠸ࡿ࡜࠸ࡿ࡜ࢃ࠿ࡿࠋ࡞࠾ࠊ௒ᅇᩘ⌮ࣔࢹࣝ࠿ࡽᑟ࠸ࡓࡢࡣ࠺ᘧ࡛࠶ࡿࠋศẕࡢࠕ཯ᛂ࣮ࣞࢺࡢ2ḟᘧࠖࡢಀᩘࡣࠊᗘ⿵ൾᛶࡢᚲせ᮲௳࡟ࡍࡂࡎࠊ௚ࡢ࣓࢝ࢽࢬ࣒7-11㸦౛඲࡚ṇࡢᩘ࠿ࡽ࡞ࡿࠋ཯ᛂ㏿ᗘࡀศẕ࡟᮶࡚࠸ࡿࡇ࡜࠼ࡤ཯ᛂࡢࡣࠊ཯ᛂࡀ㏿ࡃ࡞ࡿ࡜࿘ᮇࡀ▷ࡃ࡞ࡾࡸࡍ࠸ࡇ࡜࡟ᑐࡶࡢ࡛ࡣ࡞ࡃࠊඹᏑྍ⬟࡛࠶ࡿࠋ㠃ⓑ࠸ࡇ࡜࡟ࠊࢥࣥᛂࡍࡿࠋ

ࣆ࣮ࣗࢱ࣮ୖ࡛ࡇࡢ࿘ᮇࡢᘧ࠿ࡽ⏕≀Ꮫ࡜ࡋ࡚࡝ࢇ࡞ࡇ࡜ࡀࢃ࠿ᗘ࡟ᑐࡍࡿᏳᐃᛶ㸧ࡢ㔜せᛶࢆྰᐃࡍࡿ
ᗘ⿵ൾᛶࡢ᮲௳ࢆ᥈⣴ࡍࡿ࡜ࠊ ᗘ⿵ൾᛶࢆసࡿ࡜ࡁ࡟ࡣࠊ㧗࡛Ἴᙧࡀࡦࡎࡴ࡜ྠ᫬࡟ࡿࡢ࠿㸽ᐇࡣࠊ ᗘ⿵ൾᛶࡢᚲせ᮲௳ࡀࢃ࠿ࡿ㸦ᅗ5㸧ࠋ᣺ᖜࡶ኱ࡁࡃ࡞ࡗ࡚࠸ࡓ㸦ᅗ5㸧ࠋࡲࡓࠊࡼࡃ▱ࡽࢀ㸦ಀᩘࡀ඲࡚ṇ࡞ࡢ࡛㸧㧗࡚࠸ࡿࣂࣛࣥࢫࢭ࣮࢜ࣜࡣ7ࠊ㸦㏿ࡃ࡞ࡿࡇ࡜࡛㸧࿘ᮇ࡛཯ᛂ࣮ࣞࢺࡀ኱ࡁࡃ࡞ࢀࡤࠊᚲࡎศẕࡣ኱ࡁࡃ࡞ࡿࠋࡑࡋ࡚ᗘ⿵ൾᛶࡢࡓࢆ㛗ࡃࡍࡿ཯ᛂ࡜▷ࡃࡍࡿ཯ᛂࡢࣂࣛࣥࢫࢆㄽࡌࡓࡵ࡟ࡣࠊ཯ᛂ࣮ࣞࢺࡀ࿘ᮇࢆ▷ࡃࡍࡿຠᯝࢆ┦ẅࡍࡿࡶࡢ࡛࠶ࡿࡀࠊࠕ࿘ᮇࢆ㛗ࡃࡍࡿ཯ᛂࠖ࡜ࡣἼᙧࢆࡦ࡯࡝ࠊ㧗ࡎࡲࡏࡿ཯ᛂ࡟࡯࠿࡞ࡽ࡞࠸ࠋ

࡛Ἴᙧࡢࡦࡎࡳࡀ኱ࡁࡃ࡞ࡿᚲせࡀ࠶ࡿࠋᗘ⿵ൾᛶࡢᚲせ᮲௳࡛࠶ࡿࠋࡇࡢ᮲௳ࡣࠊࢩࢫࢸ࣒ࡢ୍㒊ࡀἼᙧࡢࡦࡎࡳ࡜ࡣࠊලయⓗ࡟࡝ࢇ࡞ᙧ࠿࡜࠸࠺࡜ࠊᗘ㠀౫Ꮡ࡛࠶ࡗ࡚ࡶኚࢃࡽ࡞࠸ࠋ࡞ࡐ࡞ࡽ౛࠼ࡤ᫬ィ㑇ఏᏊⓎ⌧ࣜࢬ࣒࡛mRNAࡢቑຍࡍࡿ᫬

ᗘ㠀౫Ꮡ࡞཯ᛂ௨እࡢ཯ᛂ㸦౛࠼ࡤศゎࡸ⬺ࣜࣥ㓟㛫ᖏࡼࡾࡶῶᑡࡍࡿ᫬㛫ᖏࡢ᪉ࡀ㛗࠸Ἴᙧ࡛࠶ࡿࠋ┤໬㸧ࡀ㧗࠸ᗘ࡛㏿ࡃ࡞ࢀࡤࠊྠࡌࡼ࠺࡟ศẕࡀ኱ࡁࡃ࡞ࡿ࠿ࡽࡔࠋࡶࡕࢁࢇ඲࡚ࡢ཯ᛂࡀࢀࡤᗘ㠀౫Ꮡ࡛࠶ᗘ⿵ൾᛶ࡟Ἴᙧࡢࡦࡎࡳࡣᚲせ࡞࠸ࠋࡋ࠿ࡋࡑ

ࡢሙྜࡣࠊ㧗᫬㛫⏕≀Ꮫ࡛࿘ᮇࡀᑡࡋࡔࡅ㛗ࡃ࡞ࡿ⌧㇟㸦overほⓗ࡟ࡣࠊ㌿෗ᅉᏊࡢ⵳✚ࡋ࡚࠸ࡿ᫬㛫ࡀ㛗ࡅࢀࡤࠊ㌿෗ᢚไࡢ᫬㛫ࡶ㛗ࡃ࡞ࡗ࡚࿘ᮇࢆᘏࡤࡍຠᯝࡀ࠶
ࡾࠊ⤖ᯝⓗ࡟ᗘ⿵ൾᛶࢆసࡾࡸࡍࡃࡍࡿࠊ࡜⌮ゎ࡛ࡁࡿࠋࡶࡋ࠿ࡍࡿ࡜ࠊ
ࠕࡇࡢㄝ࡛᫂ࡣࡼࡃศ࠿ࡽ࡞࠸ࠖ

Vol. 26, No. 1 (2020)17ᅗ 5ᗘ⿵ൾᛶ࡟ࡘ࠸࡚
ࡢἼᙧ௬ㄝࠋᗘࡀୖࡀࡗ࡚Ἴᙧࡀࡦࡎࡴ࡯࡝࿘ᮇࢆᏳᐃ࡟ಖࡕࡸࡍ࠸ࠋయෆ᫬ィࡢ ᗘ౫Ꮡࢩ࣑࣮ࣗࣞࢩࣙࣥࢆ⾜࠺࡜ࠊ㧗 ࡛Ἴᙧࡀࡦࡎࡴ࡜ྠ᫬࡟᣺ᖜࡀ኱ࡁࡃ࡞ࡗ࡚
࠸ࡿሙྜ࡟ ᗘ⿵ൾᛶࡣ⏕ࡌࡓࠋ࡜࠸࠺ㄞ⪅ࡶ࠸ࡿ࠿ࡶࡋࢀ࡞࠸ࠋἼᙧࡀ࡝࠺ࡋ࡚࿘ᮇࡦࡎࡳ㸦NS㸧ࡀኚࢃࡽ࡞࠿ࡗࡓ࡜ࡋࡓࡽࠊ⸆ࡣ㸦⏕య࡟ຠࡃࡢ࠿ࠊṇ┤࡞࡜ࡇࢁ⚾ࡓࡕࡶ⌮ゎ࡛ࡁ࡚࠸࡞࠸ෆࡢ࡝ࡇ࠿ࡢ㸧཯ᛂࢆ㐜ࡃࡋ࡚࠸ࡿ࡜ண㒊ศࡀ࠶ࡿࠋ᏶඲࡟ࡣ⌮ゎࡋ࡚࠸࡞࠸ࡢ࡛࠶ࡿࡀࠊᑡ࡟⸆࡟ࡼࡗ࡚Ἴᙧࡀࡦࡎࢇ࡛࠸ࡓࡽࠊ⸆ࡣ⏕యෆࡢ཯ࡋูࡢゅᗘ࠿ࡽㄝ᫂ࢆヨࡳࡓ࠸ࠋ๓ࡢẁⴠ࡛㸦ࢩ࣑ࣗᛂࢆ㏿ࡃࡍࡿ᪉ྥ࡟స⏝ࡋ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿࠋࡇࡢ࣮ࣞࢩࣙࣥࡢ⤖ᯝ࡜ࡋ࡚㸧ࠕᗘ⿵ൾᛶࢆసࡿ࡜ࡁ࡟ࡼ࠺࡟Ἴᙧࡢኚ໬࡟╔┠ࡍࢀࡤࠊ⏕≀ࣜࢬ࣒ࡢ࿘ᮇ࡟࡛Ἴᙧࡀࡦࡎࡴ࡜ྠ᫬࡟᣺ᖜࡀ኱ࡁࡃ࡞ࡗ࡚ຠࡃ໬ྜ≀ࡢࢫࢡ࣮ࣜࢽࣥࢢ࡟ᙺ❧ࡘ࠿ࡶࡋࢀ࡞࠸ࠋ

࠸ࡓࠖ࡜᭩࠸ࡓࠋᐇࡣࠊࡇࢀ࡟ࡣ2ࡘࡢព࿡ࡀ࠶ࡿࠋ⚾ࡓࡕࡣࠊᒸᮧᆒẶࡸJean Michel FustinẶࡽࡀぢࡣࠊ㧗1ࡘࡣࠊ㧗

ࡉࢀࡿࠋ㏫࡛᣺ᖜࢆ኱ࡁࡃࡍࡿࡇ࡜ࡀࠊἼᙧࡢࡦࡎฟࡋࡓRNA࣓ࢳࣝ໬࡟ࡼࡿᴫ᪥ࣜࢬ࣒ࡢไᚚ࡟ࡘ࠸

ࡳࢆ኱ࡁࡃࡍࡿ᪉ἲࡢ1ࡘ࡛࠶ࡿ࡜࠸࠺ࡇ࡜ࠋ2ࡘ┠ࡢ࡚ࠊලయⓗ࡟Ἴᙧࡢᐃ㔞໬ࢆ⾜ࡗࡓ17-19ࠋ⯆࿡ࡢ࠶ࡿព࿡ࡣࠊ㧗࡛᣺ᖜࢆ኱ࡁࡃࡋ࡚ࡶࡦࡎࡳࢆ኱ࡁࡃࡋ᪉ࡣᩥ⊩ࢆཧ↷ࡉࢀࡓ࠸ࠋ࡞࠾ࡼࡃㄗゎࡉࢀࡿࡀࠊୖ࡞ࡅࢀࡤ࿘ᮇࢆᏳᐃ໬ࡋ࡞࠸ࠊ࡜࠸࠺ࡇ࡜࡛࠶ࡿࠋලࡢᘧࡣࠕ࿘ᮇࡀ㛗ࡃ࡞ࡿ᫬࡟ࡣࠊ࠸ࡘࡶἼᙧࡣࡦࡎࡴࠖ

యⓗ࡟ࡣࠊ㑇ఏᏊⓎ⌧ࣜࢬ࣒࡛mRNAࡢቑຍࡍࡿ᫬

࡜࠸࠺ព࿡࡛ࡣ࡞࠸ࠋ౛࠼ࡤࠊࣜࣥ㓟໬㓝⣲C.ǅࡢ㛫ᖏ࡜ῶᑡࡍࡿ᫬㛫ᖏࡢẚࡀࠊ ᗘ࡟ࡼࡽࡎྠࡌࡔ࡜

㜼ᐖ๣PF670462ࡣᴫ᪥ࣜࢬ࣒ࡢ࿘ᮇࢆ㛗ࡃࡍࡿࡀࠊࡍࡿ࡜ࠊࡓ࡜࠼㧗࡛᣺ᖜࢆ኱ࡁࡃࡋࡓ࡜ࡋ࡚ࡶ࿘ᮇ

㜼ᐖ๣ࡢ᭷ࡿ᫬࡜↓࠸᫬࡛ࠕἼᙧࡦࡎࡳ㸦NS㸧ࠖࢆẚࢆᏳᐃ໬ࡍࡿࡢࡣ㞴ࡋ࠸ࠋ࡞ࡐ࡞ࡽࠊ ᗘ࡜࡜ࡶ࡟ศ㍑ࡋ࡚ࡶኚ໬ࡣ࡞࠸ࠋ࿘ᮇࢆ㛗ࡃࡋࡓࡢࡣࠊ㜼ᐖ๣ࡀゎࡀ࡝ࢇ࡝ࢇ㏿ࡃ࡞ࢀࡤࠊ㸦᣺ᖜࢆ኱ࡁࡃࡋࡓ࡜ࡋ࡚

ࣜࣥ㓟໬཯ᛂࢆ㐜ࡃࡋࡓࡓࡵ࡜⌮ゎ࡛ࡁࡿࠋࡇࢀ࡟ᑐ

ࡶ㸧mRNAࡢῶᑡࡍࡿ᫬㛫ᖏࡀ▷ࡃ࡞ࡗ࡚ࠊ⤖ᯝⓗ࡟

ࡋ࡚ࠊ&.ǅࡢ㐣๫Ⓨ⌧⣔ࡶ࿘ᮇࢆ㛗ࡃࡍࡿࡀࠊࡇࡢ࿘ᮇࡣ▷ࡃ࡞ࡿ࠿ࡽࡔࠋࡶࡋࡶ㧗࡛mRNAࡢῶᑡ

᫬ࡣἼᙧࡢࡦࡎࡳࡶቑࡸࡍࠋࣜࣥ㓟໬㓝⣲ࡀቑ࠼࡚཯ࡍࡿ᫬㛫ᖏࡢ๭ྜࡀ኱ࡁࡅࢀࡤࠊྠ᫬࡟᣺ᖜࢆ኱ࡁࡃ

ᛂࡀ㏿ࡃ࡞ࡗ࡚ࡶ࿘ᮇࡀᘏ㛗ࡋࡓࡢࡣࠊྠ᫬࡟Ἴᙧࡀࡍࡿࡇ࡜࡛࿘ᮇࡢ▷⦰ຠᯝࢆ┦ẅࡋࡸࡍࡃ࡞ࡿࠋࡦࡎࢇࡔࡓࡵ࡜⪃࠼ࡽࢀࡿࠋ

5.

6.

Ἴࡢᙧࢆㄪ࡭ࢀࡤ⏕≀ࣜࢬ࣒ࡢࣈࣛࢵࢡ࣎ࢵࢡ࠶࡜ࡀࡁ⚾ࡓࡕࡀィ⟬ࡋࡓ࡜ࡇࢁࠊ㟁Ꮚᅇ㊰ୖࡢ᣺ືࣔࢹࣝࢫࡀࢃ࠿ࡿ㸽Ἴᙧࡢࡦࡎࡳࡣࠊ ᗘ⿵ൾᛶ௨እ࡟ࡘ࠸࡚ࡶᵝࠎ࡞࡛࠶ࡿࣇ࢓ࣥࢹ࢔࣏ࣝ᪉⛬ᘧ࡛ࡶࠊ࿘ᮇ=[ㄪ࿴᣺ືࡢࡇ࡜ࢆᩍ࠼࡚ࡃࢀࡑ࠺࡛࠶ࡿࠋ౛࠼ࡤࠊ⏕యෆࡢ࡝ࡢ

࿘ᮇ][Ἴᙧࡢࡦࡎࡳ]1/2࡜࡞ࡗ࡚࠸ࡓࠋࢩ࢔ࣀࣂࢡࢸࣜ཯ᛂ࡟ാࡃ࠿ࢃ࠿ࡽ࡞࠸⸆๣ࡀ࠶ࡗ࡚ࠊ⏕≀ࣜࢬ࣒ࡢ࢔ࡢKaiCࣜࣥ㓟໬ࣜࢬ࣒ࡢᩘ⌮ࣔࢹ࡛ࣝࡶࠊ࿘ᮇࡣ࿘ᮇࢆ㛗ࡃࡋࡓ࡜ࡍࡿࠋࡇࡢ᫬࡟ࠊ⸆ࢆຍ࠼ࡿ๓࡜ᚋ

Ἴᙧࡢࡦࡎࡳࡢ࣮ࣝࢺ࡟ẚ౛ࡋ࡚࠸ࡓࠋ࡞ࡐࡇࡢ㛵ಀ࡛Ἴᙧࡢࡦࡎࡳࢆẚ࡭ࢀࡤ࣓࢝ࢽࢬ࣒ࢆ▱ࡿᡭࡀ࠿ࡀ⌧ࢀࡿࡢ࠿ࠊ⚾ࡓࡕࡣ⌮ゎ࡛ࡁ࡚࠸࡞࠸ࠋࡶࡋ࠿ࡍࡾࡀᚓࡽࢀࡿྍ⬟ᛶࡀ࠶ࡿࠋୖグࡢ࿘ᮇ࡜Ἴᙧࡦࡎࡳࡿ࡜࠶ࡿ⛬ᗘᬑ㐢ⓗ࡞ࡢ࠿ࡶࡋࢀ࡞࠸ࠋෑ㢌ࡢ⿕㣗⪅ࡢᘧࡣࠊ཯ᛂࡀ㏿ࡃ࡞ࡗ࡚࿘ᮇࡀ㛗ࡃ࡞ࡿ᫬ࡣࠊᚲࡎ-ᤕ㣗⪅⣔ࡢ10ᖺࣜࢬ࣒ࢆ෌⌧ࡍࡿLotka-VolterraࣔἼᙧࡣࡦࡎࡴࡇ࡜ࢆព࿡ࡋ࡚࠸ࡓࠋࡇࡢࡇ࡜࠿ࡽࠊࡶ

ࢹࣝࡸࠊάື㟁఩ࡢHodgkin-Huxleyࣔࢹ࡛ࣝࡶྠࡌ

ࡋࡶ⸆ࢆຍ࠼ࡿ๓࡜ᚋ࡛ほ㛵ಀࡀᡂࡾ❧ࡘ࠿࡝࠺࠿᳨ウࡋ࡚࠸ࡿࠋ

᫬㛫⏕≀Ꮫ
18

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Vol. 26, No. 1 (2020)

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ㅰ㎡ࡇࡇ࡟᭩࠸ࡓෆᐜࡣࠊ㔜ྜྷᗣྐ㸦㏆኱㸧ࠊ⸨ᒸཌᏊ㸦㏆

10. Isojima, Y. et al. CKIepsilon/delta-dependent኱㸧ࠊ㩾἟⪽㸦㏆኱㸧ࠊᮃ᭶ᩔྐ㸦ி኱㸧ࠊ൤ಖఙ࿃

phosphorylation is a temperature-insensitive,㸦⌮◊㸧ࠊᒸᮧᆒ㸦ி኱㸧ࠊJean Michel Fustin㸦ி

period-determining process in the mammalian኱㸧ࡢㅖẶ࡜ࡢඹྠ◊✲࡛࠶ࡿࠋ≉࡟Ἴᙧ࡟ࡘ࠸࡚ࡢ

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11. ఀ⸨㸦୕㍯㸧ஂ⨾Ꮚ. ࢩ࢔ࣀࣂࢡࢸࣜ࢔ࡢᴫ᪥᫬ឤㅰࡋࡓ࠸ࠋࡲࡓヲࡋࡃㄞࢇ࡛࠸ࡓࡔ࠸ࡓᰝㄞ⪅ࡢ᪉ィࢱࣥࣃࢡ㉁KaiCࡀ1᪥ࢆィࡿ௙⤌ࡳ. ᫬㛫⏕≀࡟ࡶᚰ࠿ࡽឤㅰࡋࡓ࠸ࠋ

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Vol. 26, No. 1 (2020)19㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼A. ࢥ࣓ࣥࢺࡢ㏻ࡾࠊ໬Ꮫ཯ᛂࡢ㠀⥺ᙧᛶࡀࡦࡎࡳࢆ኱ࡁ௨ୗࡣࠊᰝㄞ⪅࡜ᇳ➹⪅ࡢࡸࡾ࡜ࡾ࡛ࡍࠋᮏ᮶ࣈࣛ࢖ࣥࢻࡃࡍࡿࡇ࡜ࡣ࠶ࡾࡲࡍࠋࡋ࠿ࡋࠊ⥺ᙧࡢ཯ᛂ㡯ࡢࣃ࣓࣮ࣛ࡟ࡍ࡭ࡁᰝㄞ㐣⛬࡞ࡢ࡛ࡍࡀࡇࡢࡸࡾ࡜ࡾࡀ㠃ⓑࡃࠊࡲࡓࢱࡀࡦࡎࡳࢆ኱ࡁࡃࡍࡿࡇ࡜ࡶ࠶ࡾࡲࡍࠋࡍ࡞ࢃࡕࠊࡦࡎ

ㄞ⪅ࡢ᪉ࠎࡢ⌮ゎࡢᡭຓࡅ࡟࡞ࡿ࡜ឤࡌࡲࡋࡓࠋᰝㄞ⪅࡜ࡳࡢ኱ࡁࡉ࡜㠀⥺ᙧᛶࡣྠ್࡛ࡣ࠶ࡾࡲࡏࢇࠋᴫ᪥ࣜࢬ࣒

ᇳ➹⪅࡛࠶ࡿ㯮⃝ඛ⏕ࡢチྍࢆᚓ࡚ᥖ㍕ࡋࡲࡍࠋࡢไᚚࢿࢵࢺ࣮࣡ࢡࡢࡼ࠺࡟ࡓࡃࡉࢇࡢ໬Ꮫ཯ᛂࡀ㛵ࢃ

Q.ᰝㄞ⪅ࠊA. 㯮⃝ඛ⏕ ࡛ࡍࠋࡿࢩࢫࢸ࣒࡟࠾࠸࡚ࠕࢩࢫࢸ࣒ࡢ㠀⥺ᙧᛶࠖࢆᐃ⩏ࡍࡿࡇ㸦㔜ྜྷ㸧࡜ࡣ㞴ࡋࡃࠊㄞ⪅ࢆΰ஘ࡉࡏ࡚ࡋࡲ࠺ᜍࢀࡀ࠶ࡿࡢ࡛ࠊ㠀⥺ᙧᛶ࡜ࡦࡎࡳࡢ㛵ಀ࡟ࡣゐࢀ࡞࠸ࡼ࠺࡟ࡋࡓ࠸࡜ᛮ࠸㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼 㹼ࡲࡍࠋึᚰ⪅࡟ࡶࢃ࠿ࡾࡸࡍ࠸⥲ㄝ࡛࠶ࡿࠋࡋ࠿ࡋ௨ୗࡢ㒊ศࡣQ. ࡦࡎࡳࡀ኱ࡁࡃ࡞ࡿࡇ࡜ࡀࠊ࡝࠺ࡋ࡚཯ᛂ࣮ࣞࢺࡀ࿘⌮ゎࡀࡸࡸᅔ㞴࡛࠶ࡗࡓࠋᮇࢆ▷ࡃࡍࡿຠᯝࢆ┦ẅ࡛ࡁࡿࡢ࡛ࡋࡻ࠺࠿ࠋྍ⬟࡞ࡽ┤

p.8 "ࡑࡋ࡚ ᗘ⿵ൾᛶࡢࡓࡵ࡟ࡣࠊ཯ᛂ࣮ࣞࢺࡀ࿘ᮇࢆឤⓗ࡞ㄝ᫂ࢆຍ࠼࡚࠸ࡓࡔࡁࡓ࠸ࠋ▷ࡃࡍࡿຠᯝࢆ┦ẅࡍࡿ࡯࡝ࠊ㧗 ࡛Ἴᙧࡢࡦࡎࡳࡀ኱ࡁࡃ࡞ࡿᚲせࡀ࠶ࡿࠋ͇ ࡟㛵㐃ࡍࡿ஦㡯࡟ࡘ࠸࡚㉁ၥࠊせA. ࢥ࣓ࣥࢺ࠶ࡾࡀ࡜࠺ࡈࡊ࠸ࡲࡍࠋṇ┤࡞࡜ࡇࢁࠊ⚾ࡓࡕᮃ࡟࠾ᛂ࠼࠸ࡓࡔࡁࡓ࠸ࠋ⮬㌟⌮ゎ࡛ࡁ࡚࠸࡞࠸㒊ศࡶ࠶ࡿࡢ࡛ࡍࡀࠊ
ࠕ4.

ᗘ⿵ൾᛶ࡟Ἴࡢᙧࡀ㛵ಀ㸽 ࠖࡢ᭱ᚋ࡟ࠊ┤ឤⓗ࡞ㄝ᫂ࢆຍ࠼ࡲQ. ๓༙࡛㧗 ࡛ࡣ᣺ᖜࡀ኱ࡁࡃ࡞ࡿࡇ࡜࡛ࠊ ᗘ⿵ൾᛶࡋࡓࠋ

ࢆసࡗ࡚࠸ࡿ࡜ࡢㄝ᫂ࡀ࠶ࡾࡲࡋࡓࠋୖࡢグ㏙࡜ྜࢃࡏࡿ࡜ࠊ᣺ᖜࡀ኱ࡁࡃ࡞ࡿࡇ࡜ࡣࡦࡎࡳࢆ኱ࡁࡃࡍࡿࡇ࡜ࡢ୍

Q. ࡇࢀࡣࣂࣛࣥࢫࢭ࣮࢜ࣜ࡜⪃࠼࡚ࡼ࠸ࡢ࡛ࡋࡻ࠺࠿ࠋࡘࡢ᪉ἲ࡛࠶ࡿࡇ࡜࡟࡞ࡾࡲࡍࠋࡇࡢゎ㔘ࡣṇࡋ࠸࡛ࡋࡻ཯ᛂࡀ᪩ࡃ࡞ࡿ㐣⛬ࡀ࠶ࢀࡤࠊูࡢ཯ᛂ㐣⛬ࡀ㐜ࡃ࡞ࡿᚲ࠺࠿ࠋせࡀ࠶ࡿࠋࡇࡢูࡢ཯ᛂ㐣⛬࡟࠾ࡅࡿ཯ᛂ࣮ࣞࢺࡀ㛗ࡃ࡞ࡿࡇ࡜ࡣࠊࡦࡎࡳࢆࡘࡃࡿࡇ࡜࡜ྠ್࡜⪃࠼࡚ࡼ࠸ࡢ࡛ࡋ

A. ࡑࡢ㏻ࡾ࡛ࡍࠋࢥ࣓ࣥࢺ࡟ᚑࡗ࡚ࠊ
ࠕ4.

ᗘ⿵ൾᛶ࡟Ἴࡻ࠺࠿ࠋྍ⬟࡞ࡽㄝ᫂ࢆຍ࠼࡚࠸ࡓࡔࡁࡓ࠸ࠋࡢᙧࡀ㛵ಀ㸽 ࠖࡢ᭱ᚋ࡟ᩥ❶ࢆຍ࠼ࡲࡋࡓࠋ
A. 㔜せ࡞ၥ㢟ᥦ㉳࠶ࡾࡀ࡜࠺ࡈࡊ࠸ࡲࡍࠋࡈᣦ᦬ࡢ㏻ࡾࠊ
Q. ࡦࡎࡳࡀ኱ࡁࡃ࡞ࡿ࡜࠸࠺ࡢࡣࠊࢩࢫࢸ࣒ࡢ㠀⥺ᙧᛶἼᙧࡢࡦࡎࡳ࡟㛵ࡍࡿ ᗘ⿵ൾᛶࡢ᮲௳࡜ Ruoff ࡉࢇࡢࣂࡀᙉࡲࡿࡇ࡜࡜ྠ್࡜⪃࠼࡚࠸࠸ࡢ࡛ࡋࡻ࠺࠿ࠋྍ⬟࡞ࡽࣛࣥࢫࢭ࣮࢜ࣜࡣ῝ࡃ㛵ಀࡋࡲࡍࠋ
ࠕ4.

ㄝ᫂ࢆຍ࠼࡚࠸ࡓࡔࡁࡓ࠸ࠋᙧࡀ㛵ಀ㸽 ࠖࡢࢭࢡࢩࣙࣥ࡟ㄝ᫂ࢆຍ࠼ࡲࡋࡓࠋ᫬㛫⏕≀Ꮫ
20Vol. 26, No. 1 (2020)ᗘ⿵ൾᛶ࡟Ἴࡢ