33-3䠊⤖ᯝ
15.4 㼼0.4 15.3㼼0.2
85
䜘䜚ヲ⣽䛻ᙧᡂ䛥䜜䛯⥺⥔䛾ᙧែ䜢ゎᯒ䛩䜛䛯䜑䛻
TEM
䛻䜘䜛ゎᯒ䜢⾜䛳䛯䚹ᅗ3-5
䛾ྛ᮲௳䛻䛚䛡䜛
30
㛫Ⅼ䛾TEM
ほᐹ䛾⤖ᯝ䜢ᅗ3-7
䛻♧䛧䛯䚹GroEL WT䜢0.2
䛾 䝰䝹ẚ䠈GroEL G192N䜢0.05
䛾䝰䝹ẚ䠈GroEL G192I 䜢0.1
䛾䝰䝹ẚ䛷ῧຍ䛧䛯᮲௳䛻䛚䛔䛶㟷ᅄゅ䛷♧䛧䛯㒊ศ䛷䛿
AFM
䛻䛚䛔䛶GroEL WT
䜢0.1
ཪ䛿0.2
䛾䝰䝹ẚ䛷 ῧຍ䛧䛯᮲௳䛷䜏䜙䜜䛯䜘䛖䛺㛗䛔┤⥺≧䛾ᙧែ䛾䜰䝭䝻䜲䝗⥺⥔䛜ほ 䛥䜜䛯䚹䛭䛾୍᪉䛷GroEL G192N䠈GroEL G192I䠈GroEL G192W
䛭䜜䛮䜜䜢0.2
䛾䝰䝹ẚ䛷ῧຍ䛧䛯᮲௳䛻䛚䛔䛶䠈㉥ᅄゅ䛷♧䛧䛯㒊ศ䛻䛿
AFM
䛾GroEL G192W
䜢0.1
䛾䝰䝹ẚ䛷ῧຍ䛧䛯᮲௳䛷䜏䜙䜜䛯䜘䛖䛺⣽䛟▷䛔⥺⥔䛜䛽䛨䜜䛯ᙧែ䜢䛸䛳䛯䜰䝭䝻䜲䝗⥺⥔䛜ほ 䛥䜜䛯䚹GroEL 䛾ῧ ຍ 䛸ῧ ຍ 㔞 䛾㐪 䛔䛻䜘䜚ᙧ ᡂ 䛩䜛䜰䝭䝻䜲䝗⥺ ⥔ 䛾ᙧ ែ 䛜ኚ 䛧䛯䛣䛸䛜⪃ 䛘䜙䜜䜛䚹
TEM
䛾 ᐃ⤖ᯝ䛛䜙⟬ฟ䛧䛯⥺⥔ᖜ䜢⾲3-3
䛻♧䛧䛯䚹GroEL WT㻌 䛿ῧຍ䛩䜛㔞䜢ቑຍ 䛥䛫䛶䜒ᙧᡂ䛥䜜䛯⥺⥔䛾ᖜ䛿䛝䛺ኚ䛜ぢ䜙䜜䛺䛛䛳䛯䚹GroEL G192N 䜒㢮ఝ䛧䛯⤖ᯝ䜢♧䛧䛶䛔䛯䚹䛧䛛䛧䠈GroEL G192N 䛿䠈0.05 䛾䝰䝹ẚ䛻䛚䛔䛶
GroEL G192N
䛾0.2
䛾䝰䝹ẚ䜘䜚䜒⊃䛔⥺⥔ᖜ䛷䛒䛳䛯䚹䛥䜙䛻䠈GroEL G192I䛾ῧຍ㔞䜢ቑຍ䛥䛫䛶䛔䛟䛣 䛸䛷⥺⥔䛾ᖜ䛜⊃䛟䛺䛳䛶䛔䛟䛾䛜☜ㄆ䛥䜜䛯䚹GroEL G192W 䛿0.05
䛾䝰䝹ẚ䛻䛚䛔䛶GroEL G192I
䛾0.2
䛾䝰䝹ẚ䜘䜚䜒⊃䛔ᖜ䛾⥺⥔䛜ほ 䛥䜜䛯䚹0.1 䛸0.2
䛾䝰䝹ẚ䛷GroEL G192W
䜢ῧຍ䛧䛶ᙧᡂ䛧䛯䃐-Synuclein䛾䜰䝭䝻䜲䝗⥺⥔䛾⥺⥔ᖜ䛿ᴟ䜑䛶⣽䛟䠈⥺⥔ᖜ䜢 ᐃ䛩䜛䛣䛸䛜ฟ᮶䛺䛛䛳䛯䚹
86
ᅗ3-7䠊GroEL Ꮡᅾୗ䛻䛚䛡䜛䃐-Synuclein䛾⥺⥔ᙧᡂ䛾TEMほᐹ ᅗ3-5䛻䛚䛡䜛ྛᛂ᮲௳䛾30㛫Ⅼ䛾TEM⏬ീ䜢♧䛧䛶䛔䜛䚹
䝇䜿䞊䝹䝞䞊䛿200 nm䜢⾲䛧䛶䛔䜛䚹
87
⾲3-3䠊ᅗ3-7䛷ほ 䛧䛯䜰䝭䝻䜲䝗⥺⥔䛾⥺⥔ᖜ
*⥺⥔ᖜ䛿ᅗ3-7䜘䜚ᚓ䜙䜜䛯TEM⏬ീ䜘䜚䠈⥺⥔䛾3䞄ᡤ䛻䛚䛡䜛ᖹᆒ್㼼ᶆ‽೫ᕪ 䜢♧䛧䛶䛔䜛䚹―䠖 ᐃྍ
䛥䜙䛻䠈᭱䜒䃐-Synuclein䛾䜰䝭䝻䜲䝗⥺⥔ᙧᡂ䜢ᢚไ䛧䛶䛔䛯
GroEL G192W
䛻ὀ┠䛧 䛶䠈䛾䜰䝭䝻䜲䝗ᙧᡂ䝍䞁䝟䜽㉁䛻ᑐ䛩䜛จ㞟ᢚไ䛾ຠᯝ䜢᳨ウ䛧䛯(ᅗ3-8)䚹䛣䛾⤖ᯝ䛻䛚
䛔䛶䠈0.4 M䛾ሷ㓟䜾䜰䝙䝆䞁Ꮡᅾୗ䛻䛚䛡䜛GroES
䛿6
㛫(360 ศ)䛒䛯䜚䛛䜙⥺⥔᰾䛾ᙧᡂ䛜㛤ጞ䛧䛶䛔䜛䜘䛖䛻䜏䜙䜜䜛䛾䛻ᑐ䛧䠈GroEL G192W䜢ῧຍ䛧䛯᮲௳ୗ䛷䛿
15
㛫(900ศ)䛒䛯䜚䛛䜙䜟䛪䛛䛺⥺⥔᰾䛾ᙧᡂ䛜䜏䜙䜜䛯䚹A䃑1-40䛾ሙྜ䠈GroEL㠀Ꮡᅾୗ䛷䛿ᛂ㛤ጞ䛸䛸䜒䛻⥺⥔᰾䛾ᙧᡂ䛜ጞ䜎䛳䛶䛔䜛䚹䛭䛾୍᪉䛷䠈GroEL WT䛾ῧຍ䛻䛚
䛔䛶䜒
A䃑1-40
䛾⥺⥔᰾ᙧᡂ䛿㐜䛟䛺䛳䛶䛚䜚䠈䛥䜙䛻ᙧᡂ䛥䜜䜛⥺⥔㔞䛿ᖜ䛺ῶᑡ䛜䜏䜙䜜䛯䚹䛣䜜䛻ᑐ䛧䠈GroEL G192W䛾ῧຍ䛿
GroEL WT
䜘䜚䜒䛥䜙䛻⥺⥔᰾䛾ᙧᡂ䛜㐜 ᘏ䛧䠈䛥䜙䛻ᙧᡂ䛥䜜䜛⥺⥔㔞䜒ῶᑡ䛧䛶䛔䜛䛾䛜☜ㄆ䛥䜜䛯䚹䛣䛾⤖ᯝ䜘䜚䠈GroES䛸A䃑1-40
䛾䜰䝭䝻䜲䝗⥺⥔ᙧᡂ䛻ᑐ䛧䛶GroEL G192W 䛿䃐-Synuclein
䛾ሙྜ䛸ྠᵝ䛾ᢚไຠᯝ䜢䜒䛳䛶䛔䜛䛣䛸䛜᫂䜙䛛䛻䛺䛳䛯䚹
Sample
⥺⥔ᖜ*(nm)
GroEL
ῧຍ㔞×0.05
㽢0.1×0.2
䠇GroEL WT
11.6±0.9 11.2±1.8 10.6±3.8 +GroEL G192N 11.1±1.2 10.7±2.8 9.7±2.9
+GroEL G192I 9.1±2.5 8.0±2.0 7.7㼼2.3
+GroEL G192W 7.6±2.2
㻌 㻌 䇷 㻌 㻌 䇷88
ᅗ3-8䠊GroES䠈A䃑1-40䛾䜰䝭䝻䜲䝗⥺⥔䛻䛚䛡䜛GroEL G192W䛾ຠᯝ A䠊GroEL G192W 䛻䜘䜛0.4 M Gdn-HCl Ꮡᅾୗ䛻䛚䛡䜛GroES䛾⥺⥔ᢚไຠᯝ
GroES(䖃)䠈GroES+GroEL G192W(䖃) B䠊GroEL 䛻䜘䜛A䃑1-40䛾⥺⥔ᢚไຠᯝ
A䃑1-40(䖃)䠈A䃑1-40+GroEL WT(䖃)䠈A䃑1-40+GroEL G192W(䖃)
3-3-3䠊␗䛺䜛㛫䛻䛚䛡䜛 GroEL
䜢క䛳䛯䃐-Synuclein䛾⥺⥔ᙧᡂ䃐-Synuclein䛾⥺⥔ᙧᡂ㐣⛬䛻䛚䛡䜛
GroEL
䛾ᙳ㡪䜢䜘䜚ヲ⣽䛻ㄪ䜉䜛䛯䜑⥺⥔ᙧᡂ 㐣⛬䛾␗䛺䜛㛫(0䠈3䠈8䠈24 㛫)䛻GroEL
䛾ῧຍ䜢⾜䛳䛯䚹ᅗ3-9
䛻GroEL WT䠈 GroEL G192N䠈GroEL G192I䠈GroEL G192W
䜢䃐-Synuclein䛻ᑐ䛧䝰䝹ẚ0.2
䛻䛺 䜛䜘䛖䛻0䠈3䠈8䠈24
㛫䛻䛚䛔䛶ῧຍ䛧䛯⥺⥔ᙧᡂ㐣⛬䜢♧䛧䛯䚹GroEL WT 䛿䠈0䠈3 㛫䛾ῧຍ䛻䛚䛔䛶䠈᰾ᙧᡂ䛾㐜ᘏ䛿䜏䜙䜜䜛䛜䠈⥺⥔ఙ㛗㐣⛬䛷䛒䜛8
㛫䛾ῧຍ䛻䛚䛔 䛶䜟䛪䛛䛻ഴ䛔䛶䛔䛯䚹䛧䛛䛧䠈᭱⤊ⓗ䛻ᙧᡂ䛥䜜䛯⥺⥔㔞䛻ᕪ䛿䜏䜙䜜䛺䛛䛳䛯䚹24 㛫 䛻䛚䛡䜛䛶䛾GroEL
䜢ῧຍ䛧䛯᮲௳䛻䛚䛔䛶䠈GroEL䛾ᙳ㡪䛿ぢ䜙䜜䛺䛛䛳䛯䚹GroELG192N䠈GroEL G192I
䛾0䠈3
㛫䛻䛚䛡䜛GroEL
䛾ῧຍ䛿䠈ከᑡ䛾⥺⥔ᙧᡂ䛜䜏䜙䜜 䜛䛜䠈ᙧᡂ䛥䜜䛯㔞䛿ῧຍ䛧䛶䛔䛺䛔ሙྜ䛸ẚ㍑䛧䠈ᖜ䛻ῶᑡ䛧䛶䛔䛯䚹G192W 䛻䛚䛔䛶䠈0䠈3
㛫䛾ῧ ຍ䛷䛿䠈ᙧᡂ 䛥䜜䜛⥺⥔䛿䛺䛛䛳䛯䚹䜎䛯䠈8 㛫䛾 Ⅼ 䛻䛚䛡䜛GroEL
G192X
ྛኚ␗య䜢ῧຍ䛩䜛䛣䛸䛻䜘䜚⥺⥔ఙ㛗䛜㏵୰䛷Ṇ䜎䛳䛶䛔䜛䛾䛜☜ㄆ䛥䜜䛯䚹ᅗ3-8䠊GroES䠈A䃑1-40䛾䜰䝭䝻䜲䝗⥺⥔䛻䛚䛡䜛GroEL G192W䛾ຠᯝ A G EL G192W 䛻䜘䜛0 4 M Gd HCl Ꮡᅾୗ䛻䛚䛡䜛G ES䛾⥺⥔ᢚไຠᯝ
89
ᅗ3-9䠊␗䛺䜛㛫䛻䛚䛡䜛GroELᏑᅾୗ䛷䛾䃐-Synuclein䛾䜰䝭䝻䜲䝗⥺⥔ᙧᡂ (A) GroEL WT䠈(B) GroEL G192N䠈(C)GroEL G192I䠈(D) GroEL G192W
GroEL 0㛫 ῧຍ (ຑ)䠈GroEL 3㛫ῧຍ (ຑ)䠈GroEL 8㛫ῧຍ (ຑ) GroEL 24㛫ῧຍ (ຑ)䠈䃐-Synuclein (ຑ)
90
ᅗ
3-10
䛻ᅗ3-9
䛾ྛ᮲௳䛻䛚䛡䜛30
㛫䛾TEM
⏬ീ䜢♧䛧䛯䚹ᅗ3-9
䛾24
㛫䛻 䛚䛡䜛GroEL
䜢ῧຍ䛧䛯⤖ᯝ䜢ᫎ䛧䠈䛹䛾GroEL
䜢ῧຍ䛧䛯᮲௳䛷䜒ከ䛟䛾⥺⥔䛜䜏䜙 䜜䛯䚹0䠈3㛫䛻䛚䛡䜛GroEL WT
䛾ῧຍ䛷䛿䠈ᅗ3-6
䛷䜏䜙䜜䛯䜘䛖䛻㛗䛟┤⥺≧䛾ᙧែ䜢 䛸 䛳 䛯 ⥺ ⥔ 䛜 䜏 䜙 䜜 䛯 䚹 䛧 䛛 䛧 䠈 䜘 䜚 㐜 䛔 㛫 䛻 䛚 䛡 䜛
GroEL WT
䛾 ῧ ຍ 䛷 䛿 䠈 䃐-Synuclein
䛾⥺⥔ᙧᡂ䛻䜘䜚㢮ఝ䛧䠈䛽䛨䜜䛯ᙧែ䛾⥺⥔䛜䜏䜙䜜䛯䚹䛣䜜䛸䛿ᑐ↷ⓗ䛻䠈GroEL G192W
䛾Ꮡᅾୗ䛻䛚䛡䜛⥺⥔ᙧᡂ䛿䠈䛹䛾㛫䛻䛚䛔䛶䜒୍ᵝ䛻䛽䛨䜜䛯ᙧែ䛾⥺⥔䛜䜏䜙䜜䛯䚹ᅗ
3-10
䛻䛚䛔䛶ほ 䛧䛯䜰䝭䝻䜲䝗⥺⥔䜘䜚ゎᯒ䛧䛯⥺⥔ᖜ䜢⾲3-4
䛻♧䛧 䛯䚹䛣䛾⤖ᯝ䛻䛚䛔䛶䠈GroEL WT 䛿䜰䝭䝻䜲䝗⥺⥔ᙧᡂẁ㝵䛻䛚䛔䛶GroEL
䜢ῧຍ䛩䜛㛫䛜㐜䛟䛺䜛䜋䛹⥺⥔ᖜ䛜⊃䛟䛺䛳䛶䛔䛟䛾䛜☜ㄆ䛥䜜䛯䚹䛣䜜䛿
GroEL G192N䠈GroEL G192I
䛷䜒0䠈3䠈8
㛫䛻GroEL
䜢ῧຍ䛧䛯䛸䛝䛿⥺⥔ᖜ䛜⊃䛟䛺䛳䛶䛔䛟䛾䜢☜ㄆ䛧䛯䚹 䛧䛛䛧䠈Thioflavin-T䛾⺯ගほ 䛻䛚䛔䛶䛻⥺⥔䛜ᙧᡂ䛥䜜䛶䛔䜛㛫䛷䛒䜛24
㛫 䛻GroEL
䜢ῧຍ䛧䛶䜒⥺⥔ᖜ䛿8
㛫䛻GroEL
䜢ῧຍ䛧䛯⥺⥔ᖜ䜘䜚䜒ᗈ䛟䛺䛳䛶䛔䛯䚹 䛣䜜䛿䠈GroEL G192W䛻䛚䛔䛶䜒ྠᵝ䛾⤖ᯝ䛷䛒䛳䛯䚹䛣䜜䜢TEM
䛻䜘䜚ほ 䛥䜜䛯ᙧែᏛⓗ䛺ほⅬ䛸ྜ䜟䛫䜛䛸䠈GroEL WT 䛸
GroEL G192X
ኚ␗య䛷䛿䃐-Synuclein䛾䜰䝭䝻 䜲䝗⥺⥔䛻ཬ䜌䛩ᙳ㡪䛜␗䛺䜛䛣䛸䛜♧၀䛥䜜䜛䚹䛴䜎䜚䠈ᅗ3-6
䛷♧䛧䛯䜘䛖䛻䠈ῧຍ䛩䜛GroEL
䛾✀㢮䛻䜘䜚䠈ᙧᡂ䛥䜜䜛⥺⥔䛾ᙧែ䛜␗䛺䜛䛣䛸䛜ᗘ☜ㄆ䛥䜜䛯䚹91
㻌 㻌 ᅗ3-10䠊␗䛺䜛㛫䛻䛚䛡䜛GroELᏑᅾୗ䛷䛾䃐-Synuclein 䛜ᙧᡂ䛧䛯⥺⥔䛾TEM⏬ീ
㻌 㻌 ྛGroEL 䛿䠈α-Synuclein䛾䜰䝭䝻䜲䝗⥺⥔ᙧᡂ䛻䛚䛔䛶ᕥഃ䛻♧䛧䛯㛫䛻䛚䛔䛶ῧຍ䜢⾜䛳䛯䚹 TEM䛻䜘䜛䜰䝭䝻䜲䝗⥺⥔䛾ほᐹ䛿䠈ᅗ3-7䛾Thioflavin-T assay䛻䛚䛡䜛
30㛫ᙧᡂ䛧䛯⥺⥔䜢⏝䛔䛯ゎᯒ䜢⾜䛳䛯䚹 䝇䜿䞊䝹䝞䞊䛿200 nm䜢⾲䛧䛶䛔䜛䚹
92
⾲3-4䠊ᅗ3-10䛷ほ 䛧䛯䜰䝭䝻䜲䝗⥺⥔䛾⥺⥔ᖜ
*⥺⥔ᖜ䛿ᅗ3-7䜘䜚ᚓ䜙䜜䛯TEM⏬ീ䜘䜚䠈⥺⥔䛾3䞄ᡤ䛻䛚䛡䜛ᖹᆒ್㼼ᶆ‽೫ᕪ 䜢♧䛧䛶䛔䜛䚹―䠖 ᐃྍ
␗䛺䜛㛫䛾
GroEL
䛾ῧຍ䛻䛚䛡䜛Thioflavin-T
⺯ග ᐃ䛸TEM
ほᐹ䛾⤖ᯝ䜘䜚䠈GroEL G192X
ኚ␗య䛿䃐-Synuclein䛾䜰䝭䝻䜲䝗⥺⥔ᙧᡂᵝ䚻䛺≧ែ䛻䛚䛔䛶┦స⏝䛧䛶䛚䜚䠈䛥䜙䛻ᙧᡂ䛥䜜䜛⥺⥔䛾ᙧែ䛜
Gly192
䜢⨨䛧䛯䜰䝭䝜㓟䛻䜘䜚㐪䛔䛜⏕䛨䜛䛾 䜢☜ㄆ䛧䛯䚹䛣䛾ᙧᡂ䛥䜜䜛䜰䝭䝻䜲䝗⥺⥔䛾ᙧែ䛾㐪䛔䛿ᚤᑠ䛺ኚ䛷䛒䜛䚹䛧䛛䛧䠈จ㞟୰㛫య䛾ᙧᡂ䜒䠈⥺⥔ఙ㛗䛻䜒䃐-Synuclein 䛾༢㔞య䛜㛵㐃䛧䛶䛔䜛䚹䛭䛾䛯䜑䠈Gly192 䜢
⨨䛧䛯䜰䝭䝜㓟ṧᇶ䛾䛝䛥䛷䜰䝢䜹䝹䝗䝯䜲䞁䛾ഴ䛝䜢ኚ䛥䛫䛯䛣䛸䛷䃐-Synuclein ༢ 㔞య䛸䛾┦స⏝䛻㐪䛔䛜⏕䛨䠈䛣䜜䛻䜘䜚᭱⤊ⓗ䛻ᙧᡂ䛥䜜䜛⥺⥔䛾ᙧែ䛻䜒㐪䛔䛜㉳䛣 䜛䛸⪃䛘䜙䜜䜛䚹䛣䜜䜢䜘䜚ヲ⣽䛻ㄪ䜉䜛䛯䜑䛻
Seed
Ꮡᅾୗ䛻䛚䛡䜛䃐-Synuclein 䛾⥺⥔䜈䛾
GroEL WT
ཪ䛿GroEL G192W
ኚ␗య䛾ᙳ㡪䜢ゎᯒ䛧䛯䚹䃐-Synuclein䛿⥺⥔᰾䛻༢㔞య䛜⤖ྜ䛧䛶䛔䛟䛣䛸䛷⥺⥔䛾ఙ㛗䛜⏕䛨䜛䚹䛣䜜䛿䠈ண䜑⥺⥔᰾䛜Ꮡᅾ䛩䜛䛣䛸䛷⥺
⥔ఙ㛗䛜ຍ㏿䛩䜛䛣䛸䛜ሗ࿌䛥䜜䛶䛔䜛(106)䚹䛭䛣䛷䠈Seed Ꮡᅾୗ䛷䃐-Synuclein䛾༢㔞 య䛸
GroEL G192W
ኚ␗య䛜┦స⏝䛧䛶䛔䜛䛾䛷䛒䜜䜀GroEL WT
Ꮡᅾୗ䜘䜚䜒⥺⥔ᙧᡂ䛜㐜䛟䛺䜛䛣䛸䛜ண 䛥䜜䜛䚹 ῧຍ㛫 ⥺⥔ᖜ
*(nm)
䝃䞁䝥䝹
EL WT G192N G192I G192W
0h 11±1.8 8.7±1.8 8.3±1.7
䇷3h 8.8±1.1 8.3±0.9 7.8±2.6
䇷8h 8.2±0.8 7.9±0.5 6.9㼼1.2 6.4㼼1.0
24h 7.8±1.1 10㼼1.9 8.4㼼0.4 8.3㼼1.8
93
ᅗ3-11䠊Seed Ꮡᅾୗ䛻䛚䛡䜛䃐-Synuclein䛾⥺⥔ᙧᡂ䛾GroEL䛾ᙳ㡪 A䠖GroEL WT Ꮡᅾୗ
10 % Seed Ꮡᅾୗ䛷䛾䃐-Syn(䖃)䠈
Seed+䃐-Syn+GroEL (䖃)䠈䃐-Syn(䖃)䠈䃐-Syn+GroEL(䖃) B䠖GroEL G192W Ꮡᅾୗ
10 % Seed Ꮡᅾୗ䛷䛾䃐-Syn(䖃)䠈
Seed+䃐-Syn+GroEL(䖃)䠈䃐-Syn(䖃)䠈䃐-Syn+GroEL(䖃)
ᅗ
3-11
䛻♧䛧䛯䜘䛖䛻䠈Seed Ꮡᅾୗ䛷䃐-Synuclein 䛿䠈᰾ᙧᡂ䛜ᛴ㏿䛻⾜䜟䜜䠈䛭䛾 ᚋ䛾ᙧᡂ䛥䜜䜛⥺⥔㔞䜒ቑຍ䛧䛶䛔䜛䜘䛖䛻䜏䜙䜜䛯䚹䛣䜜䛻ᑐ䛧䠈㔝⏕ᆺGroEL
䜢ῧຍ䛧 䛯Seed
Ꮡᅾୗ䛻䛚䛔䛶䛒䜎䜚䛝䛺ኚ䛿䜏䜙䜜䛺䛛䛳䛯䚹䛧䛛䛧䠈GroEL G192W
ኚ␗య 䜢ῧຍ䛩䜛䛸Seed
Ꮡᅾୗ䛻䛚䛡䜛᰾ᙧᡂ䛿ᖜ䛻㐜ᘏ䛧䛶䛔䜛䛣䛸䛜☜ㄆ䛥䜜䛯䚹䛣䛾⤖ᯝ䜘䜚䠈GroEL G192Wኚ␗య䛿䃐-Synuclein༢㔞య䛜