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静岡県における野生動物のE型肝炎ウイルス汚染実態に関する研究

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静岡県における野生動物のE型肝炎ウイルス汚染実態に関する研究
Შ᩺‫ޢ‬ჄƴƓƚǔ᣼ဃѣཋƷᲿ‫׹‬Ꮑ໒ǦǤȫǹ
൲௨ܱ७ƴ᧙Ƣǔᄂᆮ
٤ዊᩉ ⚐ሶ㧔㕒ጟ⋵ⅣႺⴡ↢⑼ቇ⎇ⓥᚲ㧕
ᳰࡩ⼱ᦺ㚅㧔㕒ጟ⋵ⅣႺⴡ↢⑼ቇ⎇ⓥᚲ㧕
ጊ↰ ବඳ㧔㕒ጟ⋵ⅣႺⴡ↢⑼ቇ⎇ⓥᚲ㧕
Ꮉ᫪ ᢥᒾ㧔㕒ጟ⋵ⅣႺⴡ↢⑼ቇ⎇ⓥᚲ㧕
㧝㧚⋡⊛
ㄭᐕ‫ޔ‬㕒ጟ⋵ߢߪࠗࡁࠪࠪ╬ߦࠃࠆㄘᨋ↥‛߳ߩⵍኂ߇ᷓೞൻߒߡ߅ࠅߘߩ㚟㒰ኻ╷߇ᬌ
⸛ߐࠇࠆ৻ᣇߢ‫ࠍࠄࠇߘޔ‬࿾ၞ⾗Ḯߣߒߡ⷗⋥ߒ‫⁚ޔ‬⁸㠽₞⡺㧔ࠫࡆࠛ㧕ߣߒߡ᦭ലᵴ↪ߔ
ࠆขࠅ⚵ߺ߇⋓ࠎߦⴕࠊࠇߡ޿ࠆ‫ޕ‬᳃㑆ਗ߮ߦ⥄ᴦ૕߇ㆇ༡ߔࠆࠫࡆࠛኾ↪ߩ㘩⡺ടᎿᣉ⸳
ߩⒿേߦࠃࠅࠫࡆࠛߩ᥉෸᜛ᄢ߇࿑ࠄࠇ‫ޔ‬₞⡺೑↪⎇ⓥળ߿ࠫࡆࠛᢱℂᢎቶߩ㐿௅╬ߩႎ㆏
ߦࠃࠅࠫࡆࠛߪ⋵᳃ߦᐢߊ⍮ࠄࠇࠆ߽ߩߣߥߞߡ޿ࠆ‫⥸৻ޔߦࠄߐޕ‬ᶖ⾌ࠍⷞ㊁ߦ౉ࠇ‫⋵ޔ‬
ౝߩᄢቇߢߪࠫࡆࠛߩ㕖Ᏹ㘩߳ߩᔕ↪ߦ㑐ߔࠆ⎇ⓥ߿‫ߥࠢ࡯࠾࡙ޔ‬ടᎿ⵾ຠൻߦߟ޿ߡߩᬌ
⸛╬߽ⴕࠊࠇߡ޿ࠆ‫ޕ‬
E ဳ⢄Ἳߪ㧘E ဳ⢄Ἳ࠙ࠗ࡞ࠬ㧔HEV㧕ߦࠃߞߡᒁ߈⿠ߎߐࠇࠆᕆᕈ⢄Ἳߢ‫ޔ‬ᣣᧄࠍߪߓ
߼਎⇇ฦ࿾ߢฦ⒳േ‛ߢߩᗵᨴ߇᣿ࠄ߆ߣߥߞߡ߅ࠅ‫ੱޔ‬₞౒ㅢᗵᨴ∝ߩ৻ߟߣߒߡᵈ⋡ߐ
ࠇߡ޿ࠆ 1)‫ ߪ↢⊒ߩߘޕ‬2000 ᐕએ㒠ో࿖ߢႎ๔ߐࠇ‫ޔ‬2007 ᐕએᓟߪᲤᐕ 50 ଀೨ᓟߩ⊒↢߇
⹺߼ࠄࠇߡ޿ࠆ‫ޕ‬ේ࿃ߪࠫࡆࠛߩ↢㘩‫ߪ޿ࠆ޽ޔ‬ടᾲਇචಽߥ⁁ᘒߢߩ༛㘩╬ߦࠃࠆ߽ߩߣ
⠨߃ࠄࠇߡ߅ࠅ
2,3,4)
‫߽ߡ޿߅ߦ⋵ᧄޔ‬ᐕߦᢙฬߩᖚ⠪ႎ๔߇޽ࠆ‫ޕ‬2007 ᐕߩᖚ⠪ 2 ฬߪ⋵ౝ
ߢ᝝₪ߐࠇߚࠗࡁࠪࠪߩ↢⢄ߩ༛㘩‫ ߮ࠃ߅ޔ‬2009 ᐕߩᖚ⠪ 1 ฬߪࠪࠞߩ↢⡺ߩ༛㘩ߦࠃࠆᗵ
ᨴߢ޽ࠆߣផቯߐࠇߡ޿ࠆ 5)‫ޕ‬ታ㓙ߦ‫ޔ‬៨㘩ኻ⽎േ‛ߩ⡺ߥߤࠫࡆࠛ߆ࠄ HEV ߦᗵᨴߔࠆෂ
㒾ᕈ߇޽ࠆߎߣ߇ฦ⋵ߢߩታᘒ⺞ᩏ
6,7)
ߦࠃࠅ᣿ࠄ߆ߣߥߞߡ߅ࠅ‫ޔ‬㘩ຠⴡ↢਄ߩ໧㗴ߣߒߡ
ෂᗋߐࠇߡ޿ࠆ‫ޕ‬
E ဳ⢄Ἳ࠙ࠗ࡞ࠬᗵᨴࠍ㒐ᱛߔࠆߚ߼‫ޔߪߢ⋵ᧄޔ‬2010 ᐕߦ‫ޟ‬㊁↢േ‛⡺ߩⴡ↢෸߮ຠ⾰
⏕଻ߦ㑐ߔࠆࠟࠗ࠼࡜ࠗࡦ㧔࠾ࡎࡦࠫࠞ࡮ࠗࡁࠪࠪ㧕‫ޠ‬8)ࠍ╷ቯߒ‫ޔ‬㘩⡺ขᛒ⠪╬߳ߩࠫࡆࠛ
ߩⴡ↢▤ℂߦขࠅ⚵ࠎߢ޿ࠆ‫⋵ޕ‬⁸෹ળߢ߽ࠫࡆࠛߩขࠅᛒ޿߿༛㘩ߦߟ޿ߡᵈᗧ༐⿠ࠍⴕ
ߞߡ޿ࠆ‫ޕ‬
ߘߎߢᧄ⎇ⓥߪ‫ߢ⋵ᧄޔ‬㘩⡺೑↪ߐࠇࠆࠗࡁࠪࠪߦ߅ߌࠆ HEV ߩ଻᦭ታᘒࠍ⺞ᩏߒ㘩⡺
╬ߩᳪᨴߩෂ㒾ᕈߦߟ޿ߡᱜ⏕ߥᛠីࠍ⹜ߺࠆߎߣࠍ⋡⊛ߣߒߚ‫ޕ‬
㧞㧚᧚ᢱ߅ࠃ߮ᣇᴺ
㧝㧕᧚ᢱ
2011 ᐕ 5 ᦬߆ࠄ 2012 ᐕ 11 ᦬߹ߢߦ㕒ጟ⋵⁸෹ળຬߦࠃߞߡ᝝₪ߐࠇߚࠗࡁࠪࠪ 118 㗡ߩ
㧙 㧙
ⴊᶧ 112 ᬌ૕‫ޔ‬⢄⤳ 116 ᬌ૕‫♮ޔ‬ଢ 41 ᬌ૕߅ࠃ߮╭⡺ 29 ᬌ૕ࠍ᧚ᢱߣߒߚ‫ⴊޕ‬ᶧᬌ૕ߪណ
ขᓟ಄⬿଻ሽߐࠇߚ߽ߩࠍ࿁෼ߒ‫ࠇߘޔ‬એᄖߩᬌ૕ߪ20 ߢ಄ಓ଻ሽߒߚ‫ޔߚ߹ޕ‬᝝₪ᤨߦ
ࠗࡁࠪࠪߩ૕㊀ࠍផቯߒ‫ޔ‬ᕈ೎ࠍ⏕⹺ߒߚ‫ޕ‬
㧞㧕ELISA ᴺߦࠃࠆ IgG ᛫૕ߩ᷹ቯ
ࠗࡁࠪࠪߩⴊᶧࠍ 3,000rpm ߢ 10 ಽ㑆㆙ᔃಽ㔌ߒߡⴊᷡߣߒߡ↪޿ߚ‫ⴊࠪࠪࡁࠗޕ‬ᷡߩ
HEV IgG ᛫૕ߩ᷹ቯߪ‫ޔ‬Takahashi ࠄ 9)ߩᣇᴺߦḰߓ‫ޔ‬ᮡ⼂᛫૕ࠍ HRP ᛫ࡉ࠲ IgG ᛫૕㧔KPL,
Gaithersburg, MD)ߦᄌᦝߒߡᒰᚲߦߡታᣉߒߚ‫ޕ‬
㧟㧕RT-PCR ᴺߦࠃࠆ HEV ㆮવሶᬌ಴߅ࠃ߮Ⴎၮ㈩೉⸃ᨆ
⢄⤳߅ࠃ߮╭⡺ߪ⚦ಾߒ‫♮ޔ‬ଢߪ Ộߒߡ PBS()ߢ 20%੃೷ߣߒ‫ޔ‬10,000rpm ߢ 10 ಽ㑆㆙
ᔃಽ㔌ᓟ਄ᷡࠍណขߒߚ‫ޕ‬
RNA ᛽಴ߪ‫ⴊޔ‬ᷡ߅ࠃ߮੃೷਄ᷡߦߟ޿ߡᏒ⽼ࠠ࠶࠻㧔QIAamp Viral RNA Mini Kit㧘
QIAGEN㧕ࠍ↪޿ߡⴕߞߚ‫ޕ‬HEV ㆮવሶߩᬌ಴ߪ Li ࠄ 10)ߩᣇᴺߦḰߓߡ‫ޔ‬HEV ㆮવሶߩ ORF2
㗔ၞࠍኻ⽎ߣߔࠆ Nested PCR ࠍታᣉߒߚ‫ޕ‬2%ࠕࠟࡠ࡯ࠬࠥ࡞㔚᳇ᵒേߢ․⇣⊛ࡃࡦ࠼
㧔378bp㧕߇⏕⹺ߐࠇߚ PCR Ⴧ᏷↥‛ߪ‫ޔ‬Ꮢ⽼ࠠ࠶࠻㧔Wizard SV Gel and PCR Clean-Up System㧘
Promega㧕ࠍ↪޿ߡ♖⵾ᓟ࠳ࠗ࡟ࠢ࠻ࠪ࡯ࠢࠛࡦࠬᴺߦࠃࠅႮၮ㈩೉ࠍ᳿ቯߒߚ‫ޕ‬ᓧࠄࠇߚ
Ⴎၮ㈩೉ߪ‫ޔ‬ಽሶ♽⛔᮸⸃ᨆߦࠃࠅ Genbank ߦ⊓㍳ߐࠇߡ޿ࠆ HEV ㆮવሶߩႮၮ㈩೉ߣᲧ
セߒߚ‫ޕ‬
㧟㧚⚿ᨐ
㧝㧕HEV ․⇣⊛ IgG ᛫૕ߩ଻᦭⁁ᴫ
⺞ᩏߒߚ 107 㗡ߩ߁ߜ 53 㗡㧔49.5%㧕ߩࠗࡁࠪࠪ߇ HEV IgG ᛫૕߇㓁ᕈߢ޽ߞߚ㧔⴫ 1㧕‫ޕ‬
⴫ 1 HEV ᛫૕଻᦭⁁ᴫ
േ‛⒳
ᬌ૕ᢙ
IgG 㓁ᕈᢙ(%)
ࠗࡁࠪࠪ
107
53 (49.5%)
㧞㧕HEV ㆮવሶߩᬌ಴⁁ᴫ
HEV ㆮવሶߪ‫ޔ‬118 㗡ߩ߁ߜ 5 㗡㧔4.2%㧕ߩࠗࡁࠪࠪ߆ࠄᬌ಴ߐࠇߚ‫ޕ‬2 㗡ߪⴊᷡߩߺ‫ޔ‬1
㗡ߪⴊᷡ‫ޔ‬⢄⤳߅ࠃ߮♮ଢ߆ࠄ‫ⴊޔ‬ᷡᧂណขߩ 1 㗡ߪ⢄⤳߅ࠃ߮♮ଢ߆ࠄ‫ޔ‬1 㗡ߪⴊᷡ‫ޔ‬⢄
⤳‫♮ޔ‬ଢ߅ࠃ߮╭⡺߆ࠄ HEV ㆮવሶ߇⏕⹺ߐࠇߚ㧔⴫ 2㧕‫ޕ‬
⴫ 2 HEV ㆮવሶߩᬌ಴ (㓁ᕈᢙ / ᬌ૕ᢙ)
േ‛⒳
ⴊᷡ
⢄⤳
♮ଢ
╭⡺
ࠗࡁࠪࠪ
4/112
3/116
3/41
1/29
㧙 㧙
㧟㧕HEV ㆮવሶߩႮၮ㈩೉⸃ᨆ⚿ᨐ
ࠗࡁࠪࠪ߆ࠄᬌ಴ߐࠇߚ HEV ߩㆮવሶဳߪోߡ Genotype4 ဳߢ޽ߞߚ㧔࿑ 1㧕‫ޕ‬ห৻୘૕
ߩ⇣ߥࠆᬌ૕߆ࠄᬌ಴ߐࠇߚ HEV ㆮવሶߩႮၮ㈩೉ߪߘࠇߙࠇ 99%એ਄ߩ⋧หᕈࠍ␜ߒߚ‫ޕ‬
߹ߚ‫ޔ‬5 㗡߆ࠄᬌ಴ߐࠇߚ HEV ߩㆮવሶ㑆ߩ⋧หᕈߪ 98%એ਄ߢ޽ߞߚ‫⛔♽ޕ‬᮸⸃ᨆߦࠃࠅ‫ޔ‬
ߎࠇࠄߪએ೨ߦᗲ⍮⋵ߢᬌ಴ߐࠇߚ HEV ߣㄭ✼ߢ޽ࠆߎߣ߇್᣿ߒߚ‫ޕ‬
࿑ 1 ࠗࡁࠪࠪ߆ࠄᬌ಴ߐࠇߚ HEV ㆮવሶ⸃ᨆ⚿ᨐ
㧠㧚⠨ኤ
ࠗࡁࠪࠪߩ HEV ᗵᨴ੐଀ߪో࿖⊛ߦᄙᢙႎ๔ߐࠇߡ޿ࠆ
1,11,12)
‫⺞ᧄޕ‬ᩏߢߪ⚂ 50%ߩࠗࡁ
ࠪࠪ߇ HEV ᛫૕ࠍ଻᦭ߒߡ޿ߚߎߣ߆ࠄ‫ޔ‬HEV ߪᧄ⋵ߩࠗࡁࠪࠪߦ㜞₸ߦᶐ᷎ߒߡ޿ࠆߎ
ߣ߇್᣿ߒߚ‫ޔߪࠇߎޕ‬2005ޯ2009 ᐕߦታᣉߐࠇߚ⋵ౝࠗࡁࠪࠪ 140 㗡ߦ߅ߌࠆ᛫૕⺞ᩏႎ
๔㧔⋵ౝోၞߢ 34.2%‫᧲ޔ‬ㇱ࿾඙ߢߪ 66.7%㧕ߣห᭽ߩ㜞޿ᶐ᷎⁁ᴫࠍ␜ߔ⚿ᨐߢ޽ߞߚ
11)
‫ޕ‬
౓ᐶ⋵߿ᗲ⍮⋵ߥߤߢߪ‫ޔ‬HEV ߪ㒢ዪߒߚ࿾ၞߢࠗࡁࠪࠪߦᶐ᷎ߒߡ޿ࠆߣႎ๔ߐࠇߡ޿ࠆ
13,14)
‫ޔߚ߹ޕ‬Williams ࠄߪ‫↢⟲ߪࠪࠪࡁࠗޔ‬ᵴࠍߔࠆߚ߼‫✼ⴊߩߘޔ‬㓸࿅ߩਛߢ HEV ᗵᨴ߇
↢ߓߡ޿ࠆߣႎ๔ߒߡ޿ࠆ
15)
‫੹ޕ‬࿁᧚ᢱߣߒߚࠗࡁࠪࠪߪ⋵ౝߩ޿ߊߟ߆ߩ࿾ၞߢߩ᦭ኂ᝝
₪ߦߡᓧࠄࠇߡ߅ࠅ‫ ߇ࠄࠇߘޔ‬HEV ᗵᨴ㓸࿅ߩ৻ㇱߢ޽ߞߚߚ߼ߦ᛫૕଻᦭₸߇㜞₸ߦߥ
ߞߚߣ߽⠨߃ࠆ‫⋵ޔߡߞ߇ߚߒޕ‬ౝߩࠗࡁࠪࠪߦ߅ߌࠆ HEV ߩ࿾ℂ⊛ಽᏓࠍ⹏ଔߔࠆߚ߼
ߦߪ‫ࠅࠃޔ‬ᐢ▸࿐ߩᬌ૕ࠍᓧߡ⹦⚦ߦ⺞ߴࠆᔅⷐ߇޽ࠆߣ⠨߃ࠆ‫ޕ‬
੹࿁ߩ⺞ᩏߢ 4.2%ߩࠗࡁࠪࠪ߆ࠄ HEV ㆮવሶ߇ᬌ಴ߐࠇߚߎߣ߆ࠄ‫߇ࠪࠪࡁࠗߩ⋵ᧄޔ‬
HEV ߦᗵᨴߒߡ޿ࠆታᘒ߇⏕⹺ߐࠇߚ‫ޕ‬น㘩ㇱ૏ߢ޽ࠆ⢄⤳߅ࠃ߮╭⡺߇ HEV ߦᳪᨴߐࠇ
ߡ޿ࠆߎߣ߇್᣿ߒߚߎߣ߆ࠄ‫ߩࠪࠪࡁࠗޔ‬⢄⤳߅ࠃ߮╭⡺ߩ↢㘩޽ࠆ޿ߪടᾲਇචಽߥ⁁
ᘒߢߩ៨㘩ߦࠃߞߡ HEV ߦᗵᨴߔࠆෂ㒾ᕈ߇޽ࠆߎߣࠍ༡ᬺ⠪߅ࠃ߮ᶖ⾌⠪ߦ๟⍮ߔࠆᔅ
ⷐ߇޽ࠆ‫⺞ޔߦࠄߐޕ‬ℂౕེߩ౒↪ࠍㆱߌߡࠫࡆࠛ߆ࠄઁߩ㘩ຠ߳ߩੑᰴᳪᨴࠍ㒐ᱛߒ‫ޔ‬༛
㧙 㧙
㘩ߩ㓙ߦߪචಽߥടᾲ⺞ℂࠍⴕ߁ߎߣࠍᐢႎߔࠆߎߣߪ㊀ⷐߢ޽ࠆߣ⠨߃ࠆ‫♮ޔߚ߹ޕ‬ଢ߆
ࠄ HEV ㆮવሶ߇ᬌ಴ߐࠇߚߎߣ߆ࠄ‫ޔ‬HEV ࠍ଻᦭ߔࠆࠗࡁࠪࠪߩᵴേߦࠃࠅⅣႺਛ߳ߩ
HEV ᳪᨴ߇᜛ᄢߔࠆน⢻ᕈ߇޽ࠆߣ⠨߃ࠄࠇࠆ‫ޕ‬⁸෹ળຬ߅ࠃ߮ࠫࡆࠛಣℂᣉ⸳ߩ૞ᬺຬߥ
ߤߦኻߒߡߪ‫ ࠄ߆ࠪࠪࡁࠗޔ‬HEV ߦᗵᨴߔࠆน⢻ᕈ߇޽ࠆߎߣࠍ๟⍮ߒ‫ߣߩࠪࠪࡁࠗޔ‬Ვ
߅ࠃ߮⸃૕ᤨߩⴊᶧ߅ࠃ߮♮ଢߩขࠅᛒ޿ߦߟ޿ߡᵈᗧ༐⿠ߔࠆᔅⷐ߇޽ࠆߣ⠨߃ࠆ‫ޕ‬
੹࿁ߩ⺞ᩏߢߪ‫ޔ‬5 㗡ߩࠗࡁࠪࠪ߆ࠄᬌ಴ߐࠇߚ HEV ㆮવሶ㑆ߩႮၮ㈩೉ߪ 98%એ਄ߩ⋧
หᕈࠍ␜ߒ‫ޔ‬ㆊ෰ߦㄭ㓞ߩᗲ⍮⋵ߢႎ๔ߐࠇߚ HEV ߣㄭ✼ߢ޽ߞߚ‫⋵ᧄޔࠄ߆ߣߎߩߎޕ‬
ߩࠗࡁࠪࠪߦᶐ᷎ߒߡ޿ࠆ HEV ߪㄭធⅣႺߢ↢ᵴ࿤߇੤Ꮕߔࠆࠗࡁࠪࠪߩ㑆ߢવ᠞ߐࠇ⛮
ઍߐࠇߡ޿ࠆน⢻ᕈ߇޽ࠆߣᕁࠊࠇࠆ‫੹ޔߡߞ߇ߚߒޕ‬ᓟ‫ ࠆߔ᦭଻߇ࠪࠪࡁࠗߩ⋵ᧄޔ‬HEV
ߩ↱᧪⺞ᩏࠍ⛮⛯ߒߡⴕߞߡ޿ߊߎߣߪ‫↢ⴡⴐ౏ޔ‬਄ᭂ߼ߡ㊀ⷐߢ޽ࠆߣ⠨߃ࠆ‫ޕ‬
㧠㧚ෳ⠨ᢥ₂
1)
ᕡశ⵨ઁ㧦േ‛ߢߩ E ဳ⢄Ἳ࠙ࠗ࡞ࠬߩᗵᨴ⁁ᴫ. IASR. 26, 269-70 (2005)
2)
Matsuda, H. et al㧦Severe hepatitis E virus infection after ingestion of uncooked liver from wild
boar. J. Infect. Dis. 188, 944 (2003)
3)
Tei, S. et al.㧦Zoonotic transmission of hepatitis E virus from deer to humn beings. Lancet. 362,
371-3 (2003)
4)
Sonoda, H. et al.㧦Prevalence of hepatitis E virus (HEV) Infection in wild boars and deer and
genetic identification of genotype 3 HEV from a boar in Japan. J. Clin. Microbiol. 42, 5371-4
(2004)
5)
Ꮉ᧛ᰵ਽ઁ㧦㕒ጟ⋵⷏ㇱ࿾ၞߢ⊒↢ߒߚࠪࠞ↢⡺߹ߚߪࠗࡁࠪࠪ↢⢄៨㘩ᓟߩ E ဳᕆᕈ
⢄Ἳ 3 ଀. ⢄⤳. 51, 418-24 (2010)
6)
↰᧛ോઁ㧦ᣂẟ⋵ߩߣ⇓႐಴⩄⽋෸߮ㄘ႐ߦ߅ߌࠆ⽋ߩ E ဳ⢄Ἳ࠙ࠗ࡞ࠬߩ଻᦭⁁ᴫ.
ᣂẟ⋵଻ஜⅣႺ⑼ቇ⎇ⓥᚲᐕႎ. 23, 103-5 (2008)
7)
଻⑼ஜઁ㧦㊁↢േ‛╬ߩ E ဳ⢄Ἳ࠙ࠗ࡞ࠬߩ଻᦭⁁ᴫ. ፉᩮ⋵଻Ⅳᚲႎ. 50㧘 70-3 (2008)
8)
㕒ጟ⋵/ㄘጊ᧛౒↢⺖ࡎ࡯ࡓࡍ࡯ࠫ
http//www.pref.shizuoka.jp/sangyou/sa-325/
9)
Takahashi, M. et al.㧦Swine hepatitis E virus strains in Japan from four phylogenetic clusters
comparable with those of Japanese isolates of human hepatitis E virus. J. Gen. Virol. 84, 851-62
(2003)
10㧕Li, TC. Et al.㧦Detection of hepatitis E virus RNA from the bivale Yamato-Shijimi (Corbicula
Japonica) in Japan. Am. J. Trop. Med. Hyg. 76, 170-2 (2003)
11) ᱞ↰⋥๺㧦㘩ຠᇦ੺࠙ࠗ࡞ࠬᗵᨴ∝. ᣣᧄ㘩ຠᓸ↢‛ቇ㔀⹹. 25, 1-6 (2008)
12) Ꮉᧄᱠઁ㧦ࡅ࠻‫ ࠆߌ߅ߦࠪࠪࡁࠗޔ࠲ࡉޔ‬E ဳ⢄Ἳ࠙ࠗ࡞ࠬߩᗵᨴ⁁ᴫ. 㠽ข⋵ⴡ↢Ⅳ
Ⴚ⎇ⓥᚲႎ. 45, 1-3 (2005)
13) ᭢⾫ม㧦ෘ↢ഭ௛⑼ቇ⎇ⓥ⵬ഥ㊄ 㘩ߩ቟ోᕈ࡮㜞ᐲൻផㅴ⎇ⓥ੐ᬺ‫ࠬ࡞ࠗ࠙ޟ‬ᕈ㘩ਛᲥ
㧙 㧙
ߩ੍㒐ߦ㑐ߔࠆ⎇ⓥ‫ ⃰ޠ‬ᐔᚑ 16 ᐕᐲ✚᜝࡮ಽᜂ⎇ⓥႎ๔ᦠ. 27-32 (2005)
14) ᧻ᶆ෹♿ሶ㧦ᐔᚑ 16-19 ᐕᐲ⑼ቇ⎇ⓥ⾌⵬ഥ㊄‫ޟ‬㊁↢ߩ࠾ࡎࡦࠫࠞߪੱ޽ࠆ޿ߪઁߩേ
‛⒳ߩᗵᨴḮߦߥࠅ߁ࠆߩ߆㧫‫⎇ ⃰ޠ‬ⓥᚑᨐႎ๔ᦠ. 19-24 (2008)
15) Williams TP. et al.㧦Evidence of extrahepatic sites of replication of the hepatitis E virus in a swine
model. J. Clin. Microbiol. 39, 3040-46 (2001)
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