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BỘ GIÁO DỤC VÀ ĐÀO TẠO TRƯỜNG ĐẠI HỌC NÔNG NGHIỆ H N I NGUYỄN TÙNG NGHIÊN CỨU M T SỐ ĐẶC TÍNH SINH HỌC CỦA VI RÚT CÚM A/H5N1 CLADE HÂN LẬ Ở VIỆT NAM LUẬN ÁN TIẾN SỸ CHUYÊN NG NH: KÝ SINH TRÙNG V VI SINH VẬT HỌC THÚ Y H N I, NĂM 2013 BỘ GIÁO DỤC VÀ ĐÀO TẠO TRƯỜNG ĐẠI HỌC NÔNG NGHIỆ H N I NGUYỄN TÙNG NGHIÊN CỨU M T SỐ ĐẶC TÍNH SINH HỌC CỦA VI RÚT CÚM A/H5N1 CLADE HÂN LẬ Ở VIỆT NAM CHUYÊN NG NH: KÝ SINH TRÙNG V VI SINH VẬT HỌC THÚ Y MÃ SỐ: 62.64.01.04 NGƯỜI HƯỚNG DẪN KHOA HỌC TS NGUYỄN BÁ HIÊN TS NGUYỄN VĂN CẢM H N I, NĂM 2013 LỜI CAM ĐOAN Tơi xin cam đoan: cơng trình khoa học tôi, số liệu kết nghiên cứu luận án trung thực chưa công bố cơng trình khác Tơi xin cam đoan: giúp đỡ cho việc thực đề tài nghiên cứu hoàn thành luận án cảm ơn Các thơng tin trích dẫn luận án xác nêu rõ nguồn gốc Hà nội, tháng 10 năm 2013 Tác giả Nguyễn Tùng i LỜI CẢM ƠN Trong suốt q trình thực đề tài hồn thành luận án, nhận giúp đỡ nhiều tổ chức cá nhân Nhân dịp xin cảm ơn Ban Giám hiệu, Ban Quản lý Đào tạo, Khoa Thú y, Trường Đại học Nông nghiệp Hà Nội tạo điều kiện cho theo học chương trình đào tạo nghiên cứu sinh trường Tôi xin chân thành cảm ơn Lãnh đạo cán Trung tâm Chẩn đoán Thú y tạo điều kiện thuận lợi giúp đỡ cho tơi hồn thành đề tài nghiên cứu Tơi xin cảm ơn tập thể cán thuộc môn Vi sinh vật – Truyền nhiễm, khoa Thú y, Trường Đại học Nông nghiệp Hà Nội, giúp đỡ q trình học tập nghiên cứu Tơi xin bày tỏ long biết ơn sâu sắc đến Thầy giáo, TS Nguyễn Bá Hiên, Khoa Thú y, Trường Đại học Nông nghiệp Hà Nội TS Nguyễn Văn Cảm – Hội Thú y , người thầy hướng dẫn khoa học, trực tiếp giúp đỡ tơi q trình thực đề tài hồn thành luận án Tơi xin gửi lời cảm ơn đến đồng nghiệp công tác tổ chức quan quốc tế FAO, CDC, USDA hỗ trợ cung cấp tài liệu nguyên liệu cần thiết để thực nghiên cứu Tôi biết ơn gia đình, bạn bè đồng nghiệp ln bên cạnh, động viên giúp đỡ tơi hồn thành nghiên cứu luận án Hà Nội, tháng 10 năm 2013 Tác giả Nguyễn Tùng ii M CL C Lời cảm ơn Lời cam đoan cl c Danh m c ký hiệu, chữ viết tắt Danh m c bảng Danh m c hình MỞ ĐẦU CHƯƠNG TỔNG QUAN TÀI LIỆU 1.1 Lịch sử bệnh cúm gia cầm 1.1.1 Lịch sử bệnh giới 1.1.2 Bệnh cúm gia cầm Việt Nam 1.2 Nguyên nhân bệnh cúm gia cầm 1.2.1 Đặc điểm sinh học phân tử virus cúm gia cầm 1.2.2 Kháng nguyên virus cúm gia cầm 1.2.3 Đặc điểm tiến hóa hình thành genotype virus cúm gia cầm giai đoạn 1996-2008 1.2.4 Tính thích ứng đa vật chủ virus cúm A/H5N1 1.2.5 Cơ chế xâm nhiễm gây bệnh virus cúm A tế bào vật chủ 1.2.6 Độc lực khả gây bệnh virus cúm gia cầm 1.2.7 Triệu chứng lâm sàng gia cầm mắc bệnh cúm 1.2.8 Bệnh tích gia cầm mắc cúm gia cầm 1.3 Chẩn đoán bệnh 1.3.1 Chẩn đoán dựa vào dịch tễ học 1.3.2 Chẩn đoán dựa vào triệu chứng bệnh tích 1.3.3 Chẩn đốn phịng thí nghiệm 1.4 Vacxin phịng bệnh cúm gia cầm 1.4.1 Tình hình sử d ng vắc-xin cúm gia cầm giới 1.4.2 Tình hình sử s ng vắc-xin cúm gia cầm Việt Nam 1.5 Tình hình nghiên cứu cúm gia cầm Việt Nam CHƯƠNG NỘI DUNG – NGUYÊN LIỆU – PHƯƠNG PHÁP NGHIÊN CỨU 2.1 Nội dung 2.1.1 Phân lập giám định virus cúm gia cầm A H N1 từ mẫu dịch ngoáy ổ nhớp 2.1.2 Xác định đặc tính di truyền học virus cúm A/H5N1 clade 2.1.3 Xác định số đặc tính sinh học virus A/H5N1 clade 2.1.4 Xác định đặc tính kháng nguyên (tính tương đồng kháng nguyên) 2.1.5 Xác định hiệu lực vacxin H5N1 Re-1 2.2 Địa điểm nghiên cứu 2.3 Vật liệu phương pháp nghiên cứu 2.3.1 Vật liệu nghiên cứu 2.3.2 Phương pháp nghiên cứu Trang i ii iii v vi viii 4 10 14 20 29 30 32 34 36 36 36 36 37 37 39 41 42 44 46 46 46 46 46 46 46 47 47 47 iii CHƯƠNG KẾT QUẢ NGHIÊN CỨU VÀ THẢO LUẬN 3.1 Phân lập giám định virus cúm gia cầm H5N1 clade 3.2 Xác định đặc tính di truyền virus cúm A H N1 thuộc clade phân lập Việt Nam 3.2.1 Giải trình tự gen HA (H ) phân tích phả hệ sử d ng chuỗi gen H5 3.2.2 Giải trình tự gen NA(N1) phân tích phả hệ dựa chuỗi nucleotide gen N1 3.2.3 Giải trình tự gen phân tích phả hệ dựa chuỗi nucleotide gen M 3.3 Xác định số đặc tính sinh học chủng virus A/H5N1 clade A/chicken/Vietnam/NCVD-016/2008 3.3.1 Tính thích ứng phơi gà (xác định số EID50) 3.3.2 Tính thích ứng tế bào xơ phôi gà (xác định số TCID50) 3.3.3 Kết xác định độc lực virus cúm A/H5N1clade 3.3.4 Kết đánh giá độ thải virus động vật thí nghiệm 3.3.5 Đánh giá khả nhiễm đa phủ tạng virus cúm A/H5N1clade 3.3.6 Xác định đặc tính kháng nguyên chủng virus cúm A H N1 clade phân lập Việt Nam 3.3.7 Đánh giá khả bảo hộ vacxin Re-1 virus cúm A H N1HA clade gà KẾT LUẬN VÀ ĐỀ NGHỊ Tài liệu tham khảo Công trình cơng bố liên quan đến luận án Ph l c 59 59 62 62 71 78 84 84 89 92 103 106 113 116 121 124 135 136 iv DANH M C CÁC KÝ HIỆU, CÁC CH VIẾT T T Viết tắt Tên đầy đủ AI CDC CEF CK cs Ct DEF Dk EID50 ELD50 FAO HA HI HPAI IHC LPAI M MDCK MDEF MDT MEGA NA NCBI NCVD NP NS OIE PA PB1 PB2 RNA RRT-PCR TCID50 WHO Avian Influenza Center of Disease Control and Prevention Chicken Embryo Fibroblast Chicken cộng Cycle Threshold Duck Embryo Fibroblast Duck Embryo Infection Dose 50% Embryo Lethal Dose 50% Food and Agriculture Organisation Hemaglutinin Hemagglutination Inhibition Highly Pathogenic Avian Influenza Immuno Histochemistry Low Pathogenic Avian Influenza Matrix protein Mardine Darby Canine Kidney Muscovy Duck Embryo Fibroblast Mean Death Time Molecular Evolution Genetic Analysis Neuraminidase National Center for Biotechonology Information National Centre for Veterinary Diagnostics Nucleoprotein Non-strutural protein Office International des Epizooties Polymerase acidic Polymerase basic protein Polymerase basic protein Ribonucleic acid Realtime Reverse Transcriptase – Polymerase Chain Reaction Tissue Culture Infection Dose 50% World Health Organisation v DANH M C CÁC BẢNG TT 1.1 1.2 1.3 1.4 2.1 2.2 2.3 2.4 2.5 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 3.9 3.10 3.11 3.12 3.13 3.14 3.15 3.16 3.17 Tên bảng Trang Tổng số trường hợp nhiễm cúm gia cầm A H N1 người báo cáo cho WHO đến 2012 ột số đặc điểm triệu chứng gia cầm mắc cúm A/H5N1 34 Một số loại vacxin phòng cúm gia cầm H N1 sử d ng 40 giới Kết tiêm phòng vacxin cúm gia cầm chương trình quốc gia 41 Bảng tổng hợp số liệu tính tốn theo phương pháp eed-Muench 49 Primer probe để phát virus cúm gia cầm A/H5N1 52 Chuẩn bị mix (hỗn hợp phản ứng cho ealtime T-PCR 52 Chuẩn bị mix (hỗn hợp phản ứng) T-PC giải trình tự gen 55 Trình tự primer để giải trình tự gen(theo quy trình CDC) 56 Kết xét nghiệm virus từ chương trình giám sát biên giới 60 Kết giải trình tự gen phân tích phát sinh loài mẫu 62 virus cúm A H N1 phát Lạng Sơn Danh sách chủng virus cúm A H N1 dùng để so sánh lập 66 phát sinh loài dựa gen H5 So sánh mức độ khác biệt di truyền gen HA chủng 69 virus A H N1 clade với số chủng tham chiếu Danh sách chủng virus A H N1 sử d ng để so sánh lập 74 phát sinh loài dựa N1 So sánh mức độ khác biệt di truyền gen N1 chủng 77 virus cúm A/H5N1 clade với số chủng tham chiếu Danh sách chủng virus cúm A H N1 sử d ng để so sánh lập 80 phát sinh loài dựa gen So sánh mức độ khác biệt di truyền gen chủng virus 83 cúm A/H5N1 clade với số chủng tham chiếu Theo dõi thời gian gây chết phôi 85 Kết theo dõi tỷ lệ sống/chết phôi trứng gây nhiễm virus 86 A/Chicken/Vietnam/NCVD-016/2008 Theo dõi thời gian virus gây nhiễm lên tế bào CEF 89 Kết theo dõi bệnh tích tế bào CEF gây nhiễm virus 90 A/Chicken/Vietnam/NCVD-016/2008 93 Các axit amin vùng “cleavage site” virus cúm H N1 độc lực cao clade số chủng virus tham chiếu Kết đánh giá độc lực virus cúm A/H5N1clade gia 95 cầm Kết theo dõi lâm sàng gà thí nghiệm 96 Đánh giá độ thải virus gây nhiễm b ng virus A/chicken/ 103 Vietnam/NCVD-016/2008 Kết xét nghiệm T-PC số loại phủ tạng gà gây 107 bệnh b ng virus A H N1 clade chuyển đổi sang nồng độ virus vi 3.18 3.19 3.20 Phân bố bệnh tích vi thể nhuộm IHC phát kháng nguyên virus cúm H N1A chicken Vietnam NCVD-016/2008 Kết xác định đặc tính kháng nguyên virus cúm A/H5N1 clade phân lập b ng phản ứng HI Kết theo dõi thí nghiệm cơng cường độc gà tiêm vacxin cúm gia cầm H5N1 Re-1 109 115 118 vii DANH M C CÁC H NH TT 1.1 1.2 1.3 1.4 1.5 1.6 1.7 1.8 1.9 1.10 1.11 1.12 1.13 1.14 1.15 1.16 2.1 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 3.9 3.10 Tên hình Trang Bản đồ phân bố ca cúm A/H5N1trên giới (tính đến 2009) Biểu đồ biểu diễn dịch cúm gia cầm virus cúm A/H5N1 theo thời gian (A) hình thái virus cúm, (B) Hình thái kính hiển vi 10 điện tử hình hệ gen virus cúm A 14 ô cấu trúc kháng nguyên Haemalutinin Neurminidase 15 Sơ đồ minh họa đột biến điểm phân đoạn genvirus cúm A 20 Sơ đồ minh họa tượng trộn kháng nguyên virus cúm 20 A/H5N1và H3N2 Cây phả hệ dựa gen HA virus cúm A H N1 độc lực cao 22 Sự tiến hóa clade virus A/H5N1 theo thời gian 23 Sự phân bố clades virus cúm gia cầm giới từ 200324 2009 Thời gian xuất clade H5N1 Việt Nam từ 2001-2007 25 Hình 1.12 Sự phân bố clade virus A/H5N1 khác theo 27 không gian Các genotype virus cúm gia cầm A H N1 độc lực cao 28 ối quan hệ lây nhiễm thích ứng lồi vật chủ virus cúm A 30 hình chế xâm nhiễm nhân lên virus cúm A tế bào chủ 31 Minh hoạ vùng “Cleavage site” virus cúm độc lực thấp(LPAI) 33 Sơ đồ bố trí primer giải trình tự gen H , N1 virus 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Option VII meeting in Hongkong, 2010 135 PHỤ LỤC 1) T ình tự gen HA củ c c i c A/H5N1HA clade A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 10 20 30 40 50 60 70 | | | | | | | | | | | | | | ATGGAGAAAATAGTGCTTCTTCTTGCAATGATCGGTCTTGTTAAAAGTGATCAGATTTGCATTGGTTACC .A G .A A.C .A A.C .A G A A A C .G A .A .G G A .G G A .A .A .A .A A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 80 90 100 110 120 130 140 | | | | | | | | | | | | | | ATGCAAACAACTCGACAGAGCAGGTTGACACAATAATGGAAAAGAACGTTACTGTTACACATGCTCAAGA .A A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 150 160 170 180 190 200 210 | | | | | | | | | | | | | | CATACTGGAGAAGACACACAACGGGAAGCTCTGCAACCTAGATGGGGTGAAGCCTCTAATTTTGAAAGAC .A .C.A A .C.A A .C.A A A .T .A .C.A A A .T .A .C.A A A .T A A .T .T A A .T .A .C.A G .A .C.A A .C.A A .C.A A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 220 230 240 250 260 270 280 | | | | | | | | | | | | | | TGTAGTGTAGCTGGATGGCTCCTCGGGAACCCAATGTGTGACGAATTTCTCAATGTGTCAGAATGGTCTT A A A A A A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 290 300 310 320 330 340 350 | | | | | | | | | | | | | | ACATAGTGGAGAAGGCCAGTCCAGCCAATGGCCTCTGTTACCCAGGGGATTTCAATGACTATGAAGAACT 136 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ T .A T T T A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 360 370 380 390 400 410 420 | | | | | | | | | | | | | | GAAACACCTATTGAGCAGAATAAACCATCTTAAGAAAATTAAGATCATCCCCAAAAGTTATTGGTCCAAT A.A CA A.A CA A.A CA T G A.A CA T G A.A CA T G T G A.A CA A.A CA A.A CA A.A CA A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 430 440 450 460 470 480 490 | | | | | | | | | | | | | | CATGAAGCCTCATCAGGGGTGAGCGCAGCATGTTCCTATCTGGGAGAGCCCTCCTTTTTCAGAAATGTGG G A G G A T T AGA.T T A A G A T T AGA.T T A G A T T AGA.T T A A T T AGA.T T A G T G G G G A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 500 510 520 530 540 550 560 | | | | | | | | | | | | | | TATGGCTTATCAAAAAGAATAATACATACCCACCAATAAAGGTGAACTACACCAATACCAACCAAGAAGA .T .T .T C .C G A T C G A T C G A .C G A .C T .T .T .T .T A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ 570 580 590 600 610 620 630 | | | | | | | | | | | | | | TCTTTTGGTACTGTGGGGGATTCACCATCCCAATGATGAGAAAGAGCAGATAAGGATCTATCAAAACCCA A GC CA T A GC CA T A GC CA T A .A GC A T T A GC CA T A .A GC T T A GC CA T A .A GC T T A .A GC T T A GC CA T A GC CA T A GC CA T 137 A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ A GC CA T A GC CA T A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 640 650 660 670 680 690 700 | | | | | | | | | | | | | | AACACCTCTATTTCAGTTGGAACATCAACACTAAACCAGAGATTGGTACCAAAAATAGCTACTAGACCCA T CA A T T CA A T T CA A T .A.G C T T CA A T .A C GT T CA.R A T .A C T .A C T .A A T T CA.G A T T CA A T T CA A T T CA A T A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 710 720 730 740 750 760 770 | | | | | | | | | | | | | | AAGTGAACGGGCAAAGTGGACGAATGGAGTTCTTCTGGACAATTTTAAAGCCGAATGATTCTATCAATTT .T A A T A A .C A A A .C A A .A G A .C A G A .C A G A A .A A .A A .A A .A C A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 780 790 800 810 820 830 840 | | | | | | | | | | | | | | CGATAGCAATGGAAATTTCATTGCTCCAGAATATGCATACAAAATTGCCAAGAAAGGGGACTCAGTGATT C - T .G.CA C - T .G.CA C - T .G.CA T T .CA C - T .G.CA T T .CA C - T .G.CA T T .CA T T .CA - C T .CA C - T .G.CA C - T .G.CA C - T .G.CA C - T .G.CA A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 850 860 870 880 890 900 910 | | | | | | | | | | | | | | ATGAAAAGTGAATTGGAATATGGTAACTGCAACACCAAGTGTCAAACTCCAATGGGGGCGATAAATTCTA G .G .G A C A G A C A G A C A A C A A .G .G .G .C .G .C A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 920 930 940 950 960 970 980 | | | | | | | | | | | | | | GTATGCCATTCCACAACATACACCCTCTCACCATCGGGGAATGCCCCAAATATGTGAAATCAAACAGATT G G G 138 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ G G C G G G A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 990 1000 1010 1020 1030 1040 1050 | | | | | | | | | | | | | | AGTCCTCGCGACTGGACTCAGAAATGCCCCCCAAAGAGAG GGAGGAAGAAGGAGAAAAAGAGGA T AAGAGAG G A T AAGAGAG G A T AAGAGAG G A T T - A T AAGAGAG G A T T - A T AAGAGAG G A T T - A T T - A T T A T AAGAGAG G A T AAGAGAG G A T AAGAGAG G A T AAGAGAG G A A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 1060 1070 1080 1090 1100 1110 1120 | | | | | | | | | | | | | | CTATTTGGAGCCATAGCAGGGTTTATAGAGGGAGGATGGCAGGGAATGGTAGATGGTTGGTATGGGTACC C C C C C A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 1130 1140 1150 1160 1170 1180 1190 | | | | | | | | | | | | | | ACCATAGCAATGAGCAGGGGAGTGGATACGCTGCAGACAAAGAATCCACTCAAAAGGCAATAGATGGAAT G T C G .G T C .T A A .G T .A .G T .A .T .A .T .A .G .G .G .G A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 1200 1210 1220 1230 1240 1250 1260 | | | | | | | | | | | | | | CACCAATAAGGTCAACTCGATCATTGACAAAATGAACACTCAGTTTGAGATCGTTGGAAGGGAATTTAAT T A GC T A GC T A GC A GC T A GC A GC T A GC A GCT A GCT A GC T A GC T A GC T A GC T A GC 139 A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 1270 1280 1290 1300 1310 1320 1330 | | | | | | | | | | | | | | AACTTAGAAAGGAGAATAGAAAATTTAAACAAGAAGATGGAGGACGGATTCCTAGATGTCTGGACTTATA G .A G .A G .A A G .A A G .A A A A A A G .A G .A G .A G .A A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 1340 1350 1360 1370 1380 1390 1400 | | | | | | | | | | | | | | ACGCTGAACTTCTGGTTCTCATGGAGAATGAGAGAACTCTAGACTTTCATGACTCAAATGTCAAGAACCT A .A .A .A C T A .A C T .T A .A C T A C T T C T .T A .T A .T A .T A A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 1410 1420 1430 1440 1450 1460 1470 | | | | | | | | | | | | | | TTACGAAAAGGTCCGACTACAGCTTAGGGATAATGCAAAGGAGCTGGGTAACGGTTGTTTTGAGTTCTAC C C C G C C G C C G C G C G C C .W C C C A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 1480 1490 1500 1510 1520 1530 1540 | | | | | | | | | | | | | | CACAAATGTGATAATGAATGTATGGAAAGTGTAAGAAACGGAACGTATGACTACCCGCAGTATTCAGAAG T A G T A G .A A T G .A A T G .G A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ 1550 1560 1570 1580 1590 1600 1610 | | | | | | | | | | | | | | AAGCAAGACTAAGCAGAGAGGAAATAAGTGGAGTAAAAATGGAATCAATGGTAACTTATCAAATACTGTC A T C A T C A T C A .T T C A T C A A .T T C 140 A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ .A C C A .T T C A .T T C A .T T C A T .A T C A T C A T C A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 1620 1630 1640 1650 1660 1670 1680 | | | | | | | | | | | | | | AATTTATTCAACAGTGGCGAGTTCCCTAGCATTGGCAATCATGGTGGCTGGTCTATCTTTATGGATGTGC A A/chicken/Vietnam/NCVD-016/200 A/chicken/Vietnam/NCVD-04/2008 A/chicken/Vietnam/NCVD-03/2008 A/chicken/Vietnam/NCVD-05/2008 A/chicken/Vietnam/NCVD-093/200 A/chicken/Vietnam/NCVD-swab06/ A/chicken/Vietnam/NCVD-swab15/ A/chicken/Vietnam/NCVD-swab16/ A/chicken/Vietnam/NCVD-swab17/ A/chicken/Vietnam/NCVD-swab18/ A/chicken/Vietnam/NCVD-swab19/ A/chicken/Vietnam/NCVD-swab20/ A/chicken/Vietnam/NCVD-swab24/ A/chicken/Vietnam/NCVD-swab25/ A/chicken/Vietnam/NCVD-swab26/ 1690 1700 1710 | | | | | | | TCCAATGGATCGTTACAATGCAGAATTTGCATTTGA 10 2) T ình tự gen NA củ c c i c 20 30 40 50 A/H5N1HA clade A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 10 20 30 40 50 60 70 | | | | | | | | | | | | | | ATGAATCCAAATCAGAAGATAATAACCATCGGATCAATCTGTATGGTAATTGGAATAGTTAGCTTAATGT C C C G A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 80 90 100 110 120 130 140 | | | | | | | | | | | | | | TACAAATTGGGAACATGGTCTCAATATGGGTCAGTCATTCAATTCAGACTAGGAATCAACGCCAAGCTGA .T .T .T .A .AGA .T A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 150 160 170 180 190 200 210 | | | | | | | | | | | | | | ACCGATCAGCAACACTAAATCTCTTACTGAGAAAGCTGTGGCTTCAGTAACATTAGCGGGCAATTCATCT G .G .G .T T .T C G AT A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 220 230 240 250 260 270 280 | | | | | | | | | | | | | | CTTTGCCCCATTAGCGGATGGGCTGTATACAGTAAGGACAACAGCATAAGGATCGGTTCCAAGGGGGATG C 141 A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 290 300 310 320 330 340 350 | | | | | | | | | | | | | | TGTTTGTTATAAGAGAGCCGTTCATCTCATGCTCACACTTGGAATGCAGAACTTTCTTTTTGACTCAGGG C A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 360 370 380 390 400 410 420 | | | | | | | | | | | | | | AGCTTTGCTAAATGACAAGCACTCCAATGGGACAATCAAAGACAGAAGTCCTCACAGAACATTAATGAGT G C G C G C C G .TG C A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 430 440 450 460 470 480 490 | | | | | | | | | | | | | | TGTCCTGTGGGTGAGGCTCCCTCCCCTTATAACTCAAGGTTTGAGTCTGTCGCTTGGTCAGCAAGTGCTT C .T C .T C .T .T A .T A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 500 510 520 530 540 550 560 | | | | | | | | | | | | | | GCCATGATGGCACCAGTTGGTTGACAATTGGAATTTCTGGCCCAGATAATGGGGCTGTGGCTGTATTGAA .C A G C A G C A G C A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 570 580 590 600 610 620 630 | | | | | | | | | | | | | | ATACAATGGCATAATAACAGACACTATCAAGAGTTGGAGAAACAACATACTGAGAACTCAAGAGTCTGAA T A G A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 640 650 660 670 680 690 700 | | | | | | | | | | | | | | TGTGCATGTGTAAATGGCTCTTGCTTTGCTGTAATGACTGATGGACCAAGTAATAGGCAGGCATCATATA T A A T .C A A T .C A A A G A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 710 720 730 740 750 760 770 | | | | | | | | | | | | | | AGATTTTCAAAA TGGAAAAAGGAAAAGTGGTTAAATCAATCGAA-TTGGATGCTCCTAATTATCACTA - - - C T G .G.T - A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 780 790 800 810 820 830 840 | | | | | | | | | | | | | | TGAGGAGTGCTCCTGTTATCCTGATGCTGGCGAAATCACATGTGTGTGCAGAGATTATTGGCATGGCTCA C A C A .A A G A A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 850 860 870 880 890 900 910 | | | | | | | | | | | | | | AATAGGCCATGGTTATCTTTCAATCAAAATTTGGAGTATCAAATAGGGTATATATGCAGCGGAGTTTTTG T C .T G A T C .G G CT A .A T C A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 920 930 940 950 960 970 980 | | | | | | | | | | | | | | GAGACAATCCACGCCCCAATGATGGAACAGGTAGTTGTGGTCCGGTGTCCTCTAACGGGGCATATGGGGT CA A C .G C .C T 990 1000 1010 1020 1030 1040 1050 | | | | | | | | | | | | | | 142 A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 AAAAGGGTTTTCATTTAAATACGGCAATGGTGTTTGGATCGGGAGAACCAAAAGCACTAATTCCAGGAGC A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 1060 1070 1080 1090 1100 1110 1120 | | | | | | | | | | | | | | GGTTTTGAAATGATTTGGGATCCAAATGGGTGGACTGAAACGGACAGCAGCTTTTCGATGAAGCAAGATA .G .G .G C G G G A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 1130 1140 1150 1160 1170 1180 1190 | | | | | | | | | | | | | | TCGTAGCAATAACTGATTGGTCAGGATATAGCGGGAGTTTTGTTCAGCATCCAGAACTGACAGGATTAGA G C G C G C .C A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 1200 1210 1220 1230 1240 1250 1260 | | | | | | | | | | | | | | TTGCATAAGACCTTGTTTCTGGGTTGAGTTGGTCAGAGGGCGGCCCAAAGAGAGCACAATCTGGACTAGT T .G T A A G T A A G A A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 1270 1280 1290 1300 1310 1320 1330 | | | | | | | | | | | | | | GGGAGCAGCATATCTTTTTGTGGTGTAAATAGTGACACTGTGAGTTGGTCTTGGCCAGACGGTGCTGAGT C C C C G G A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/2008 A/CHICKEN/VIETNAM/NCVD-04/2008 A/CHICKEN/VIETNAM/NCVD-05/2008 A/CHICKEN/VIETNAM/NCVD-093/200 1340 1350 | | | | TGCCATTCACCATTGACAAGTA - 3) T ình tự gen M củ c c i c A/H5N1HA clade A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 10 20 30 40 50 60 70 | | | | | | | | | | | | | | ATGAGTCTTCTAACCGAGGTTGAAACGTACGTTCTCTCTATCATCCCGTCAGGCCCCCTCAAAGCCGAGA A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 80 90 100 110 120 130 140 | | | | | | | | | | | | | | TAGCGCAAAAACTTGAAGATGTTTTCGCAGGAAAGAACACTGATCTCGAGGCTCTCATGGAGTGGCTAAA A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 150 160 170 180 190 200 210 | | | | | | | | | | | | | | G G GACAAGACCAATCCTGTCACCTCTGACTAAAGGGATTTTGGGATTTGTATTCACGCTCACCGTGCCCAGT A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 220 230 240 250 260 270 280 | | | | | | | | | | | | | | T GAGCGAGGACTGCAGCGTAGACGCTTTGTCCAGAATGCCCTAAGTGGAAATGGAGATCCAAATAATATGG 143 A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 290 300 310 320 330 340 350 | | | | | | | | | | | | | | G ATAGGGCAGTTAAGCTATATAAGAAGCTGAAAAGAGAAATAACATTCCATGGGGCTAAAGAGGTCGCACT A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 360 370 380 390 400 410 420 | | | | | | | | | | | | | | C A CAGCTACTCAACTGGTGCACTTGCCAGTTGCATGGGTCTCATATACAACAGAATGGGAACGGTAACTACG A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 430 440 450 460 470 480 490 | | | | | | | | | | | | | | G GAAGTAGCTTTTGGTTTAGTGTGTGCCACTTGTGAGCAGATTGCAGATTCACAGCATCGGTCTCACAGAC A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 500 510 520 530 540 550 560 | | | | | | | | | | | | | | G AGATGGCAACTATCACCAACCCACTAATCAGGCATGAGAACAGAATGGTGCTAGCCAGCACTACAGCTAA A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 570 580 590 600 610 620 630 | | | | | | | | | | | | | | C .C .C .C .C GGCTATGGAGCAGATGGCGGGATCAAGTGAGCAGGCAGCGGAAGCAATTGAGATCGCTAATCAGGCTAGG A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 640 650 660 670 680 690 700 | | | | | | | | | | | | | | A CAGATGGTGCAGGCAATGAGGACAATTGGGACTCATCCTAACTCTAGTGCTGGTCTGAGGGACAATCTTC A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 710 720 730 740 750 760 770 | | | | | | | | | | | | | | TTGAAAATTTGCAGGCCTACCAGAAACGAATGGGAGTGCAGATGCAGCGATTCAAGTGATCCTCTTGTTG A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 780 790 800 810 820 830 840 | | | | | | | | | | | | | | TTGCCGCAAATATCATTGGGATCTTGCACTTGATATTGTGGATTCTTGATCATCTTTTCTTCAAATGCAT A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 850 860 870 880 890 900 910 | | | | | | | | | | | | | | T TTATCGTCGCCTTAAATACGGTTTGAAAAGAGGGCCTTCTACGGCAGGGGTACCTGAGTCTATGAGGGAA A/CHICKEN/VIETNAM/NCVD-016/200 A/CHICKEN/VIETNAM/NCVD-03/08 A/CHICKEN/VIETNAM/NCVD-04/08 A/CHICKEN/VIETNAM/NCVD-05/08 A/CHICKEN/VIETNAM/NCVD-093/08 920 930 940 950 960 970 980 | | | | | | | | | | | | | | G G G G GAATACCGGCAGGAACAGCAGAGTGCTGTGGATGTTGACGACGGTCATTTTGTCAACATAGAGTTGGAGT 144 SỐ ĐĂNG KÝ NGÂN HÀNG GEN CAC CHỦNG VI US CÚ CỦA VIỆT NAM (SỬ DỤNG T ONG NGHIÊN CỨU) A H N1 CLADE Số đăng ký gen HA (GenBank) Số đăng ký full Genome (GISAID) A/Chicken/Vietnam/NCVD-016/2008 FJ842476 EPI_ISL_28830 A/chicken/Vietnam/NCVD-03/2008 FJ842481 EPI_ISL_80638 A/chicken/Vietnam/NCVD-04/2008 FJ842482 EPI_ISL_80639 A/chicken/Vietnam/NCVD-05/2008 FJ842483 EPI_ISL_80640 A/chicken/Vietnam/NCVD-093/2008 FJ842480 EPI_ISL_80641 A/chicken/Vietnam/NCVD-swab06/2008 FJ842484 A/chicken/Vietnam/NCVD-swab15/2008 FJ842477 A/chicken/Vietnam/NCVD-swab16/2008 FJ842485 A/chicken/Vietnam/NCVD-swab17/2008 FJ842478 A/chicken/Vietnam/NCVD-swab18/2008 FJ842479 A/chicken/Vietnam/NCVD-swab19/2008 FJ842486 A/chicken/Vietnam/NCVD-swab20/2008 FJ842487 A/chicken/Vietnam/NCVD-swab24/2008 FJ842488 A/chicken/Vietnam/NCVD-swab25/2008 FJ842489 A/chicken/Vietnam/NCVD-swab26/2008 FJ842490 Tên chủng 145 ... Total Quốc gia Cases Death Cases Death Cases Death Cases Death Cases Death Cases Death Cases Death Cases Death Cases Death Cases Death Cases Death Ajerbaijan Bangladesh Cambodia China 1 4 2 1 13... N1 clade 62 Hiện tượng chèn x? ?a axit amin vị trí cleavage site 71 chủng virus A H N1 clade phân lập Việt Nam Cây phả hệ d? ?a gen N1 virus A H N1 clade 73 Cây phả hệ d? ?a gen virus A H N1 clade 79 ... cao 22 Sự tiến h? ?a clade virus A/ H5N1 theo thời gian 23 Sự phân bố clades virus cúm gia cầm giới từ 200324 2009 Thời gian xuất clade H5N1 Việt Nam từ 2001-20 07 25 Hình 1.12 Sự phân bố clade virus