7
Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tpns20 Download by: [University of California Santa Barbara] Date: 29 February 2016, At: 17:58 Progress in Natural Science ISSN: 1002-0071 (Print) 1745-5391 (Online) Journal homepage: http://www.tandfonline.com/loi/tpns20 Isolation and identification of a novel coronavirus from wild bird Zhang Guihong , Fu Jiadong , Ren Tao , Cao Weisheng , Luo Kaijiang , Xu Chenggang , Xin Chaoan , Jiang Jingwei & Liao Ming To cite this article: Zhang Guihong , Fu Jiadong , Ren Tao , Cao Weisheng , Luo Kaijiang , Xu Chenggang , Xin Chaoan , Jiang Jingwei & Liao Ming (2006) Isolation and identification of a novel coronavirus from wild bird, Progress in Natural Science, 16:12, 1275-1280 To link to this article: http://dx.doi.org/10.1080/10020070612330141 Published online: 25 Feb 2007. Submit your article to this journal Article views: 1 View related articles Citing articles: 1 View citing articles

2006 Isolation and identification of a novel coronavirus from wild bird

  • Upload
    others

  • View
    2

  • Download
    0

Embed Size (px)

Citation preview

Page 1: 2006 Isolation and identification of a novel coronavirus from wild bird

Full Terms & Conditions of access and use can be found athttp://www.tandfonline.com/action/journalInformation?journalCode=tpns20

Download by: [University of California Santa Barbara] Date: 29 February 2016, At: 17:58

Progress in Natural Science

ISSN: 1002-0071 (Print) 1745-5391 (Online) Journal homepage: http://www.tandfonline.com/loi/tpns20

Isolation and identification of a novel coronavirusfrom wild bird

Zhang Guihong , Fu Jiadong , Ren Tao , Cao Weisheng , Luo Kaijiang , XuChenggang , Xin Chaoan , Jiang Jingwei & Liao Ming

To cite this article: Zhang Guihong , Fu Jiadong , Ren Tao , Cao Weisheng , Luo Kaijiang , XuChenggang , Xin Chaoan , Jiang Jingwei & Liao Ming (2006) Isolation and identification of anovel coronavirus from wild bird, Progress in Natural Science, 16:12, 1275-1280

To link to this article: http://dx.doi.org/10.1080/10020070612330141

Published online: 25 Feb 2007.

Submit your article to this journal

Article views: 1

View related articles

Citing articles: 1 View citing articles

Page 2: 2006 Isolation and identification of a novel coronavirus from wild bird

!"#$"%&&’(()*+"),&-’%(-% !"#$%&,’"$%(,!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!)*+*,-*.(//&

"0122".3*4-5’637"86#’631.6#0+7*8+*9"184637"8":;<786(=.683’"$>?@?>(?/)

""A<*B*3C"613<".B+"83.7-13*4*D16##53"3<7BC".E"""A"C<",+"..*B2"84*8+*B<"1#4-*644.*BB*4$FG,67#:,#76"!B+61$*41$+8

!"#$%&’#(%()’)*(&’+’,%&’#(#+%(#-*$,#.#(%-’./"+.#01’$)2’.)"

HIJ’==17<"8K"",9LM764"8K"",NF’A6",;JOP*7B<*8K,QLOR67S768K,

TL;<*8KK68K,TU’;<6"68,MUJ’=M78KC*7684QUJOV78K"""

(!*3*.786.5V*47+78*;"##*K*":0"13<JK.7+1#31.*L87W*.B735,=168KX<"1Y%/&Z(,;<786)

N*+*7W*4V6.+<%>,(//&;.*W7B*4J2.7#Y,(//&

32"&.%,& J8"W*#+"."86W7.1BB3.678C6B7B"#63*4:.",#6.58K"3.6+<*6#BC6-":C7#426.3.74K*6844*B7K863*46B26.3.74K*/=)/

0%Z/(//>(0%Z)$U3BC<"#*K*8",7+B*D1*8+*C6B"-3678*4(=*8[68EJ++*BB7"881,-*.:J\&Z&(@>)3<."1K<NAG];N6,2#7:7+637"8,

+#"878K,81+#*"374*B*D1*8+78K,684686#5B7B-53<*)’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’K*8*7847+63*43<63K."12UB3.678B,7K<3*W"#W*:.",N’J.*+",-78637"8-*3C**8B3.678IY(684=.65;C<7#*K."12UUB3.678B:.",B3.678I%(/684)%Z&&$0%ZB3.678<643<*<7K<*B3’K*8*<","#"K5C73<B3.678_TU[!C<7+<C6B?Y$‘ ,3<1B+#6BB7:7*46B6K."12UUU,*,-*.$03.678B0JU[(/684=)&G?@C<7+<C*.*+#"B*#5.*#63*43"3<*VK*8*":0%ZB3.678-*#"8K*43"K."12U684U!,.*B2*+37W*#5$J2"BB7-#*."#*":26.3.74K*0%ZB3.678,652#65783<*2."+*BB":+"."86W7.1B*W"#137"87B47B+1BB*4$

4*51#.)": 1’$)2’.),,#.#(%-’./","&./,&/.%$6.#&*’(7*(*",-’./"*-#$/&’#(8

;"."86W7.1B*a7B3BC74*#578W6.7"1BE784B":4"G,*B37+684C7#4687,6#B$A<*5":3*8.*B1#378.*B27.6G3".5".783*B3786#7##8*BB78+#1478KW",73,476..<*6684*W*84*63<":3<*78:*+3*4687,6#B[%]$U83<7BB3145,

"8*C7#4-7.4+"."86W7.1BC6B7B"#63*4684+<6.6+3*.G7X*4:.",63"36#":%Y%/#6.58K"3.6+<*6#BC6-B684@>37BB1*B6,2#*B,C<7+<C*.*+"##*+3*4:.", C7#42"1#3.547B3.7-13*478W6.7"1B6.*6B":=168K4"8K]."W78+*,;<786$A<7BB3.678C6B+#"B*#5.*#63*43"6GW76878:*+37"1B-."8+<737BW7.1B-1347B3683#53"0JN0G;"!-52<5#"K*8*37++",26.7B"8C73<W7.1B*B1B78KK*8",7+B*D1*8+*46366W67#6-#*78=*8[68EAV$

9 :%&*.’%$"%()0*&;#)"

%$% V63*.76#B

!7.1B*B:]6.3.74K*+"."86W7.1B(4*B7K863*46B0%Z)C6B7B"#63*468474*837:7*4-53<*R*5Q6-".63"G.5":]"1#3.59**478K684)7B*6B*B;"83."#,V787B3.5":JK.7+1#31.*,;<786$

N*:*.*8+*B3.678B:N*B27.63".5G352*B3.678B78G+#14*4 VZ%(0% V(%@@>,0(:T/Z‘((, V:

J9(@&%@Z,’:V(@Y&&),J.E??(0:Q%/>@Z,’:

V@Y(ZZ),0)%G?‘(J\>(Y‘>(),)F/‘((V:

J9(/(???,’:J9(/>//%),)%Z&&(0:V(%?‘%,V:

J9(/>//Z,’: J9(/>//&), I6’%G?Y (0%:

J\(Y%@%‘, V: J\>(Y‘>Z ), I6’(G?Y(J\>(Y‘(@), =T%G?@(0%: J\>%?>/(,V:

J9>(Y‘(‘,’: J\(‘@%%/), =T(G?@ (0%:

J\(Y%@%&,V:J\>(Y‘>>,’:J\(Y%@%&),I%(/(0:V(%?‘/,V:J\/(@(?Y,’:J\/(@(?&),IY((0:J9>Y(>%Y,V:J9(@&%@Y)$;LGA((0:

L/Z‘>?,V:LZ&/>Y,’:L/Z@/>)C6B6.*B27.63"G.5,13683352*;8*2<."K*87+G352*B3.678B78+#14*4MT%G??(J9(%/‘>Y),0JU[CS(J9>?‘Y(@),I"#3*(0:Q%@?@@;J9>>Z&@Y),=.65(0%:Q%Z/&?,0(:

J9>?Z%@/), R[@Y(> ( V(%Y%Y), T (0%:

J9Z(‘@%?,’:J\/Z>>%Y),[M(0:J\>%?&YY%)$]."W*83.7+1#1BG352* B3.678B 78+#14*4 _%

(J9(@&>/(),M((J9>Y(>%(),A>(J9((‘Z>@),

Dow

nloa

ded

by [

Uni

vers

ity o

f C

alif

orni

a Sa

nta

Bar

bara

] at

17:

58 2

9 Fe

brua

ry 2

016

Page 3: 2006 Isolation and identification of a novel coronavirus from wild bird

!"#$%(&’()*+,*),-"+(.:&/(01()2,3:

&/**02*,),45)2((.:&’*1,*((,3:&’*1,*60)7

.89:;<;:=8>?@AB9C=<D99 (.E’) 9FGDHAC>?9I:@;:J9C9BBK:3;C9=I>H=ALI.E’9FGDHAC>?9I:@;:J9C9BBKM9D9J;CILH8DAN;I9IGHOP89D;F9C?>L.?>?;ACA<%9?9D;C>DHQ9I;:;C9,.AR?@S@;C>&BD;:RL?RD>LTC;=N9DK;?H7

UA?>LV3&OP?D>:?;ACW;?###M>K8RD:@>K9I<DAF.;CA=&F9D;:>C$;A?9:@CALABHSA7&Q%"-,

X3>K;C,*Y=’RLLV&SOSAD9.9?,1Y=’RLLV&SOSAD9.9?,8I(3)Z,>CIOP!"#9C[HF9M9D98DAN;I9IGHU>W>V>$;A?9:@CALABH(X>L;>C)SA7,-?I7

(7, Q9?@AIK

(7,7( %;DRK 8DA8>B>?;AC >CI V3&9P?D>:?;AC .(+K?D>;CN;DRKM>K;CA:RL>?9I?A(6=I>H=ALI.E’9FGDHAC>?9I:@;:J9C9BBK,<ALLAM9IGH;C:RG>?;AC>?*2\<AD+0@ARDK7S@;:J9C9FGDHAK?@>?I;9I;C,+@ARDK8AK?=;CA:RL>?;ACM9D9I;K:>DI9I7

&LL>C?A;:<LR;IKM9D9:ALL9:?9I>K98?;:>LLH,:9C?D;<RB9I>?+666D8F/F;C>?+\,>CI?@9CN;DRKB9CAF;:V3&M>K9P?D>:?9I<DAF?@9KR89DC>?>C?>::ADI;CB?A?@98DA?A:ALKA<?@9UA?>LV3&OP?D>:?;ACW;?###(.;CA=&F9D;:>C$;A?9:@ALABH-?I7)7

(7,7, VU=ESV>F8L;<;:>?;AC,:LAC;CB>CIK9]R9C:=;CBA<N;DRKB9CAF9K9BF9C?K VU=ESV>F8L;<;:>=?;AC>CI:LAC;CBA<N;DRKB9CAF9K9BF9C?KM9D9:>D=D;9IAR?>::ADI;CB?A?@9F9?@AIKA<V9C9?>L7[Z]7U@99P?D>:?9IN;D>LV3&M>KI9KALN9I;C((71!-XOES=?D9>?9IM>?9D>CI?@9CD9N9DK9LH?D>CK:D;G9I7U@9*Y=>CI1Y=’RLLV&SOM9D989D<ADF9I>::ADI;CB?A ?@9 D9:AFF9CI>?;ACK A< ?@9 F>CR<>:?RD9D(U>J>D>)7EAK;?;N9:LAC9KM9D9K9C??A.@>CB@>;$;A&K;>$;ALAB;:>LU9:@CALABHSA7,-?I7<ADCR=:L9A?;I9K9]R9C:;CB7

&?A?>LA<,)8>;DKA<8D;F9DK(U>GL9()M9D9I9=K;BC9I<AD:LAC;CB>CIK9]R9C:;CBA<B9CAF;:K9B=F9C?K>CIKHC?@9K;[9IGH.@>CB@>;$;A&K;>$;ALAB;:>LU9:@CALABHS 7,-?I7

U>GL9(7 ED;F9D8>;DKI9K;BC9I

’ADM>DI V9N9DK9 G8( 1Y=S&&_SSU&&&&S&___&_U&US=*Y 1Y=_UU&&_US&UUUS_S&U_SS=*Y )+6, 1Y=__S&U_S_&&&U_&SUU&&S=* 1Y=&__US_US&S&S&UUUUSUS=*Y )+(* 1Y=_&_&&&&U_U_U_&S_&SSU=*Y 1Y=S&_&&SUSUUU&&&SUS&USS&=*Y (+,(+ 1Y=U__&U_&_UUU&&&_&_UUSU_=*Y 1Y=_&_S__U&UUU&&S&SS&U=*Y )001 1Y=&U__U_UU&&&U&SS_SUS=*Y 1Y=&__UUU&UU&&__U_UUSU_=*Y 066Z 1Y=_S&_&&S&SSUU&&U&&&SSU=*Y 1Y=UU_S_S&U&&U&_&U__S=*Y Z222 1Y=_SS&USU&UU&U_S_S&&=*Y 1Y=UUUSUSU&UU&&&SS&&_U&__&=*Y (6*+0 1Y=USSU&SUU__UUU&&U&_&_&&&=*Y 1Y=&UU&SS&&S&&&SUS&UU_SS=*Y )Z,) 1Y=__S&&U_&_UUU_UU__U&&U=*Y 1Y=US&_UUSS&_U_U_U&&U_S&=*Y ),Z(6 1Y=U_S&UU&S&S&SU__&&SU_&=*Y 1Y=UUS&&S&UU&&_SUU&_US&&&=*Y ((16(( 1Y=UUU_&SU&&BSUU&&U_UU_&&=*Y 1Y=USS&&&S&&SU_UU_&&U&U_=*Y (2*6(, 1Y=S&U&UUS&&S&_UU_UUU__&=*Y 1Y=__USUUU/USS_U&_U&__U&UU=*Y (Z,0(* 1Y=&&U&SSU&SU&S__&V&&&_&SS=*Y 1Y=U&SUSUU_&USSS&_U&SUU&&&_=*Y 2+((+ 1Y=SUUU&&_U&SU___&US&&_&_U&=*Y 1Y=__VU&_SS‘_S&SUSUU&US=*Y (*,+(1 1Y=_&U&&_&_U_S‘__SU&/SS=*Y 1Y=&SU&_S&UU_UVU_UU_‘_&&S&=*Y +1*(Z 1Y=U_UUS‘S&&S&/&S&&U_SU&_U=*Y 1Y=SS&&&&&SV_U&SS&UU&_&U&S=*Y Z)Z(2 1Y=_U&USU&&U__U&S/_UUUUU__=*Y 1Y=UUU&USUU&SS&UU&&U&&&W_&S=*Y (,)0(0 1Y=_USQUUU&UU&&U__U&&_&U&&&=*Y 1Y=_UU_U&_SUSU&W&&SS&S&&_&=*Y +0(() 1Y=_UUSUU_U__UUQU&_&_SU&S&&S=*Y 1Y=UUU__S&_U_U_S_&&UVUU=*Y (*,Z,6 1Y=&&/&UUS_S&S&SU_SS&&&=*Y 1Y=U_USUUUU_UUS&_UUUUS&&=*Y 02(,( 1Y=UU_&&&&SU_&&S&&&&_&S&=*Y 1Y=S&U&&SU&&S&U&&___S&&U=*Y (*2,,, 1Y=S&_UUU_U&_UUUSU__U__=*Y 1Y=US&UUSS&SS&UUU‘_&S&&=*Y 0+6,* 1Y=UU_US‘&&&U__U__&&U_&=*Y 1Y=&UUSS&&SS_UUSUU&__=*Y Z12,+ 1Y=SSU&&_&&S__UU__&&U=*Y 1Y=U&SUSUSU&S&S&S&S&S=*Y Z*,,1 1Y=U_&SS&&V_S__&&&U&&_&=*Y 1Y=&&U_&&VUSSS&&S__&&&U=*Y (,)6,Z 1Y=_&U__U&U&_U_U___UU=*Y 1Y=SUSSUS&UUS&USUU_US=*Y (Z++,2 1Y=_&U_&S&&_&U_&&U_&__&=*Y 1Y=&&_SU&S&U_SSU&USUUSS=*Y (6(6,0 1YV&SO(=’1Y=S&UUU&_&&_&S&USUUU__U

_US=*Y1YV&SO(=V1Y=U_U _&S _U& U&_ &&& &&S&&&_S_US&S=*Y

Z6+

,) *YV&SO=’1Y=__&&_&U&__S&U_U&_SUU=*Y

*YV&SO=V1Y=SU_&USU&_&__U&SS__&USS=*Y

**,

Z2,( MMM7?>CI<7:A7RJ/aARDC>LK EDAB!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

D9KK;C3>?RD>L.:;9C:9 %AL7(Z3A7(, ,66Z

Dow

nloa

ded

by [

Uni

vers

ity o

f C

alif

orni

a Sa

nta

Bar

bara

] at

17:

58 2

9 Fe

brua

ry 2

016

Page 4: 2006 Isolation and identification of a novel coronavirus from wild bird

!"#$%&’&()"*+$,(-’.,&,/*,"+,0#&+#11’,.(*-2%-3,45!6781(91#:#".+(:8#1,.0’-38*2$’&3,.1,$,;#"-&,/*,"+,&#;#’$#2$,’"<,"4#"=7>?!"*:@2,1()1,81,&,"-#-’;,;’1*&,&)1(:.’)),1,"-9,(91#83’@+#$$(+#-’("&0,1,&,$,+-,.#".#"#$%A,.2%BC!6@7!D&()-0#1,8#+=#9,?

! "#$%&’$

E?F D7@GHD#:8$’)’+#-’("()6FI9,"(:,

6FI&-1#’"0#&*&,.)(1DC!8*1’)’+#-’("#".D7@GHD#:8$’)’+#-’("#&.,&+1’2,.#2(;,?!&&3(0"’"J’9?F,0,&*++,&&)*$$%(2-#’",.-3,+BC!)1#9@:,"-&2%9,"(:’+-,1:’"#$DC!D!HK($#",F,

EL),(11(*-’",D7@GHD?

J’9?F? D7@GHD#:8$’)’+#-’("()6FI03($,9,"(:’+DC!?>,

>#1=,1@B5EMMM;$#",F,NOD!HK;$#",&E—EP,+$("’"9&,9:,"-&*&’"9-3,81’:,18#’1&#&&3(0"’"7#2$,F;$#",EL,QOD!HK?

E?E !"#$%&’&()"*+$,(-’.,&,/*,"+,&()6F81(-,’"9,",&#"..,.*+,.#:’"(#+’.&,/*,"+,

6F3%8,1;#1’#2$,1,9’("&()-3,-0(&-1#’"&(>IF#".6FI)0,1,(2-#’",.2%&,/*,"+’"9,03’+3’"+$*.@,.-3,"*+$,(-’.,&)1(:&-#1-+(.,"!7<-(-3,+$,#;@#9,&’-,()681,+*1&(181(-,’"?6F9,",()6FI&-1#’"0#&FREM28$("9,,"+(.’"9NIM#:’"(#+’.&#".+("@-#’"’"9FL8(-,"-’#$C@9$%+(&%$#-’("&’-,&?73,<SH8,1+,"-#9,0#&QN?ILT?U%.1(83’$’+#"#$%&’&().,@.*+,.#:’"(#+’.&,/*,"+,&()6F81(-,’"&3(0,.-3#--3,)’1&-EMC@-,1:’"#$#:’"(#+’.1,&’.*,&()6F81(@-,’"()6FI;’1*&)(1:,.#3%.1(83(2’+1,9’(",03’+30#&81,&*:,.#&-3,&’9"#$&,/*,"+,()681,+*1&(181(-,’"?!:’"(#+’.&,/*,"+,&#--3,+$,#;#9,&’-,()681,+*1&(181(-,’"0,1,DDJDD?

H(:8#1,.0’-31,),1,"+,&-1#’">IF,-3,1,0,1,&,;,1#$’"&,1-’("&#"..,$,-’("&’"6FI&-1#’":#7H7’"&,1-’("#-8(&’-’("RV,!77!#-EFR,!7!!7HH!#-EIN,!#-QNQ,<#-ERR,!#-QNM,!<#-QRM,

7H7#-QRE,!7<<#-IEM#".H7#-IEL?W"-3,(-3,13#".,-3,.,$,-’("&()!77H!7<<7<<7#-

8(&’-’("EEF—EQQ,7#-EQR,!#-ERE,!7<<HH#-IFN—IEF0,1,’.,"-’)’,.?73,1,0,1,&’:’$#1:*-#@-’("&’"5XI#".YXZ4[&-1#’"&,’"&,1-’"97H#".!<#-8(&’-’("QRM,1,&8,+-’;,$%?73,1,0,1,#$&(&(:,"*+$,(-’.,&*2&-’-*-’("&’"6FI&-1#’"?H(:@8#1,.0’-38(&’-’("EN()>IF&-1#’",-3,.,.*+,.#:’"(#+’.&,/*,"+,()6FI3#.#"!&"(C)’"&,1-’("#--3’&8(’"-#".<$%(<)S6,1(6)#-8(&’-’("FFL?

J’9?E&3(0&)(*191(*8&()’"),+-’(*&21("+3’-’&;’1*&,&(Z4[)?<1(*8Z+(*$.2,.’;’.,.’"-(-0(&*291(*8&,’"03’+3<1#%,[’+6,\>]&-1#’"&2,@$("9-(&*291(*8!#".(-3,1&2,$("9-(&*291(*84?<1(*8ZZ+(*$.#$&(2,.’;’.,.’"-(-0(&*291(*8&,

0’-36FI,\XF@LL,#".7\E@LR&-1#’"&+$#&&’)’,.-(&*291(*8H#".(-3,1&-1#’"&-(&*291(*8B?6*2@91(*84&-1#’"&3#.3’939,",-’+’.,"-’-%0’-3&-1#’"&UNE,UFEM#".>IF?Z-0#&&8,+*$#-,.-3#-<1(*8Z;’1*&,&0,1,:*-#-,.)1(:;#++’",;’1*&?!$$<1(*8ZZ;’1*&,&0,1,’&($#-,&)1(:H3’"#’"+$*.’"98*2@$’&3,. ",831(9,"’+@-%8, #". 81(;,"-1’+*$*&@-%8,&-1#’"&?6FI&-1#’"0#&#$&()(*".’"-3’&91(*8?<E@LP’&($#-,)1(:H3’"#()<1(*8ZZZ3#.+$(&,’.,"-’-%0’-3&-1#’"U($-,,#".’-0#&.,.*+,.-3#-’-:’93-2,:*-#-,.)1(: U($-,?<1(*8Z[&-1#’"&0,1,81(;,"-1’+*$*&@-%8,’"),+-’(*&21("+3’-’&;’1*&,&’&(@$#-,.’"H3’"#,-3,6F9,",()03’+30#&.’&-#"-$%1,@$#-,.-(-3(&,()(-3,1;’1*&,&?

E?Q !"#$%&’&()"*+$,(-’.,&,/*,"+,&()6E9,",#"..,.*+,.#:’"(#+’.&,/*,"+,

73,+(.’"91,9’("()6E81(-,’"()-3,-0(&-1#’"&(>IF#".6FI)0#&(2-#’",.2%"*+$,(-’.,&,@/*,"+’"9#"#$%&’&,+(;,1’"9-3,"*+$,(-’.,&)1(:-3,+$,#;#9,&’-,()681,+*1&(181(-,’"-(-3,&-(8+(.("?6E81(-,’"+(.’"91,9’("()6FI&-1#’"0#&FPVP28’"$,"9-3,03’+3,"+(.,&REN#:’"(#+’.&?73,8,1+,"-@#9,()<SH0#&QN?NET?6FI&-1#’"3#.FQC@9$%@+(&%$#-’("&’-,&-31(*93 #"#$%&’&()9$%+(&%$#-’("1,+(9"’-’("&’9"#$&?73,1,0#&#3%.1(83(2’+1,9’("#-8(&’-’("NNM-(NLM’"-3,H@-,1:’"#$,03’+3:’93-2,-3,+("",+-’"91,9’("(),";,$(8,#"."*+$,(+#8@&’.?H(:8#1,.0’-3-3,8*2$’&3,.&,/*,"+,()>IF&-1#’"(<,"4#"=#++,&&’(""*:2,1:>EFPPQ),-3,1,0,1,:#"%"*+$,(-’.,:*-#-’("&’"6E81(-,’"9,",()6FI&-1#’"?

H(1("#;’1*&&-1#’"&()<1(*8Z0,1,+$(&,$%1,$#-@,.-(UNE,>IF#".U($-,?!$$<1(*8ZZ;’1*&,&,’"@+$*.’"98*2$’&3,.",831(9,"’+@-%8,#".81(;,"-1’+*@$*&@-%8,&-1#’"&,0,1,’&($#-,.)1(:H3’"#?6FI&-1#’"

VVEFG1(91,&&’"C#-*1#$6+’,"+, [($?FRC(?FE EMMR 000?-#".)?+(?*=/_!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

(*1"#$&

Dow

nloa

ded

by [

Uni

vers

ity o

f C

alif

orni

a Sa

nta

Bar

bara

] at

17:

58 2

9 Fe

brua

ry 2

016

Page 5: 2006 Isolation and identification of a novel coronavirus from wild bird

!"#"$#%&’()&#*+%,-./"01’(%*1()1+,()1+1!"#2$%#1*1’1(&2&31’(&(451(!11’6+%,-7#(+"&’#"’36+%,-77#(+"&’#,!)&$16+%,-7778&+,#1#!1+1+1-+19

#1’("(&81#:%+;<1+&2"’+1#-&+"(%+49(4-1"’38"22&’1#(+"&’#(=&*.>).

=&*.?. @)4$%*1’1(&2(+11%:AB*1’1.!7CD&#%$"(1#:+%<E)&’".

=&*.>. @)4$%*1’1(&2(+11%:A?-+%(1&’*1’1.!7CD&#%$"(1#&’E)&’".

?.F ;’"$4#&#%:’,2$1%(&31#1G,1’21#%:H-+%(1&’*1’1"’3313,213"<&’%"2&3#1G,1’21

I,2$1%(&31#1G,1’21#"’3313,213"<&’%"2&3#19G,1’21%:H-+%(1&’%:()1#1(!%#(+"&’#!1+1:,+()1+"’"$4J13./)1%-1’+1"3&’*:+"<1%:()1H-+%(1&’%:ABF#(+"&’!"#KLM5-&’$1’*(),1’2%3&’*??N"<&’%"2&3#./)1+1!1+1(!%I9*$42%#4$"(&%’#&(1#"(()1I9(1+<&’"$%:ABF#(+"&’./)1:&+#(BB"<&’%"2&3#%:ABF#(+"&’!1+1)43+%-)&$&2"’3!1+1-+1#,<13"#()1%,(<1<5+"’13%<"&’%:H-+%(1&’,()1&+3%!’#(+1"<?O(%BOO"<&’%"2&3#:%+<13()+11)43+%-)%5&2+19

*&%’#"’3!1+12%’#&31+13"#()+11(+"’#<1<5+"’13%<"&’#.

7’2%<-"+&#%’!&()()1-,5$&#)13#1G,1’21%:PN?96Q(61’C"’0"221##&%’’,<51+:;ROFFBMF),

()1+1!1+1<"’4’,2$1%(&31#,5#(&(,(&%’#5,(’%&’9#1+(&%’%+31$1(&%’&’H-+%(1&’*1’1%:ABF#(+"&’.

C"#13%’H*1’131’3+%*+"<,6+%,-72%,$351:,+()1+3&8&313&’(%#,5*+%,-#;"’3C.S)&$1TUB9VL,6WB9VM,QXOL?,P"I?9VN,"’3PXIF9VF!1+12$%#1$4+1$"(13(%PN?,HFB51$%’*13(%#,5*+%,-;.QBFKK,PB?O,A;7C!Y,AQB9VL,AE?9V> "’3

ML?B !!!.("’3:.2%.,0/Y%,+’"$# @+%*"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""

+1##&’I"(,+"$A2&1’21 D%$.BKI%.B? ?OOK

Dow

nloa

ded

by [

Uni

vers

ity o

f C

alif

orni

a Sa

nta

Bar

bara

] at

17:

58 2

9 Fe

brua

ry 2

016

Page 6: 2006 Isolation and identification of a novel coronavirus from wild bird

!"#$$,%&’(&&")*&+&’,&+-*,+.+*’()+.*’*/%’*&0123,4+#5.,+)*5-64,75689:;1<4+#5.,+)*5

,7568===,".)%"-"#-5(#5-+#/7+#"*+)*5>7568-=".)==46*)’-*".*#/*5>7568=?(@’,:<):

@’,:<: A&/#5,+.+*’(*7++5BC875*+’.,+.+:!=9?’-5#"*+-’.!&’."

2:D E."#/-’-5B.6(#+5*’)+-+F6+.(+-5BG875*+’.,+.+".))+)6(+)"H’.5"(’)-+F6+.(+

G6(#+5*’)+-+F6+.(+".))+)6(+)"H’.5"(’)-+IF6+.(+5BG875*+’.,+.+5B*%5-*7"’.-,C1<".);1<,%+7+54*"’.+)4/-+F6+.(’.,:J&+58+.7+")’.,B7"H+5BG875*+’.5B;1<-*7"’.%"-12K348’.#+.,*&,+.(5)’.,<3$"H’.5"(’)-:J&+7+%"-5.+GI,#/(5-/#"*’5.-’*+"**&+GI*+7H’."#5B;1<-*7"’.,

%’*&".5*&+7’.*&+H’))#+:G 875*+’. %"-&/I)758&’#’(,%&’(&H"/4+4+.+B’(’"#B57*&+’.*+7"(*’5.5BG875*+’.".)LGE:

=.(5H8"7’-5.%’*&*&+864#’-&+)-+F6+.(+5BC<1(>+.9".M"((+--’5..6H4+7:C2NDOO),*&+G875*+’.,+.+5B;1<-*7"’.&").6(#+5*’)+-64-*’*6*’5.-46*.5’.-+7*’5.57)+#+*’5.:

9"-+)5.*&+"."#/-’-5BG875*+’.,+.+,>7568=(56#)4+B67*&+7)’P’)+)’.*5-64,7568-E".)9,5B%&’(&QR1I$S,;E=91<".);E=9%T(#5-+#/7+#"*+)*50D24+#5.,+)*5-64,7568E".);E=923,>U1I$N,

>U2I$N%&’(&&")*&+&’,&+-*’)+.*’*/%’*&>7"/".)E7M$$4+#5.,+)*5-64,75689:>7568==(56#)"#-54+)’P’)+)’.*5!".)V-64,7568-,5B%&’(&;!2I$K,

;V1I$S(#5-+#/7+#"*+)*5V1<OO4+#5.,+)*5-64,7568!".)0"G2I$D,>V1INS%&’(&&")*&+&’,&+-*’)+.I*’*/%’*&01234+#5.,+)*5-64,7568V:;1<-*7"’.&")*&+&’,&+-*’)+.*’*/%’*&WU=9?".)4+#5.,+)*5>7568===(@’,:D):

! "#$%&$$#’(

E-".LGEP’76-,=9?’-875.+*5,+.+*’(H6*"I*’5.".)7+(5H4’."*’5.)67’.,*&+875(+--5B7+8#’("I*’5.,7+-6#*’.,’.*&++H+7,+.(+5B.+% H6*".*-*7"’.-%’*&)’BB+7+.*-+75*/8+-:J&+7+B57+,’*’-)’BB’I(6#**5)+P+#58+BB+(*’P+P"((’.+-","’.-*=9?’.B+(I*’5.:E-"7+-6#*,"P’".’.B+(*’56-475.(&’*’--*’##85-I+--’,.’B’(".*+(5.5H’(*&7+"**5*&+856#*7/’.)6-*7/:=.56787+P’56-%57M,%+54*"’.+)B567(575."P’76-’-5#"*+-B75H%’#)8"7*7’),+,5.+5B%&’(&’-)+-’,."*I+)"-A"7*7’),+(575."P’76-;1<:

A&/#5,+.’*’("."#/-’-5B*&+%&5#+,+.5H+5B;1<&+#8+)6-B’.)56**&+57’,’.5B*&+P’76-:9/"."#/XI’.,*&+-*76(*67"#875*+’.,+.+-(;1,;2,C".)G)

5B;1<,;1<%"-(#6-*+7+)%’*&>7568!P’76-+-4/;1,+.+,*&+P’76-+-’.*&’-,7568"7+"##.+8&75,+.’(I*/8+".)875P+.*7’(6#6-I*/8+=9?-’-5#"*+)B75H!&’I.":;1<&")"&’,&’)+.*’*/%’*&.+8&75,+.’(I*/8+-*7"’.-QU1I$$,JQ2I$O,WU=9?,RU<".)9Q:Y&’#+;2,+.+5B;1<-*7"’.&")"H6(&&’,&+7,+.+*I’(’)+.*’*/ %’*&*&+ !&’.+-+ 875P+.*7’(6#6-I*/8+-*7"’.-WU=9?".)RU<,%&’(&%"-$S:2Z ".)$3:OZ,7+-8+(*’P+#/:J&+/B57H+)"-64,7568==%’*&".5*&+7!&’.+-+=9?’-5#"*+9Q:A&/#5,+.+*’("."#/-’-5BC ,+.+-&5%+)*&"*;1<4+#5.,+)*5>7568===".)&")"&’,&+7,+.+*’(’)+.*’*/%’*&*&+!&’.+-+’-5#"*+-;E=923,>VOI$N".)RU<*&".*&"*

$S21A75,7+--’.G"*67"#;(’+.(+ ?5#:1OG5:12 233O %%%:*".)B:(5:6M/T"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""

567."#-

Dow

nloa

ded

by [

Uni

vers

ity o

f C

alif

orni

a Sa

nta

Bar

bara

] at

17:

58 2

9 Fe

brua

ry 2

016

Page 7: 2006 Isolation and identification of a novel coronavirus from wild bird

!"#$%&’()*#+,-&%./010!"234&435!674’89:;!<5!67***=>1?>1&?.6’4’;>4@>1&404*#+0;5%1&0

()*#+,A):%&’#$,

B13,C, D>/.!34&4;1?;544!"275!;41&34&4,!*#+10!.%;401&@>1&%,

*;=%0?!&01’454’;>%;8E%&’234&40!"*#+=454?!&045F4’,*&!655404%5?>,;>48E%&’234&40!"89:=454?.!04./54.%;4’;!;>4@>1&404*#+10!.%;4006?>%0()*#+,A):%&’#$,0!=%0;>48934&4,G!=4F45,1;=%0&!;;>4?%04"!5H34&4,-&%./010!"H34&40>!=4’;>%;;>4H34&4!"89:=%0?.!04./54.%;4’;!*!=%,8-*#EI,<JKLMN%&’A):O6;F45/’10;%&;./;!#$%&’()*#+,81&?489,8E%&’234&40!"89:=454?.!04./54.%;4’;!;>!04!"@>1&40410!.%;40()*#+%&’#$,=>!0404P64&?40=454%..’10;%&;./54.%;4’;!;>!04!"0;5%1&0"5!Q!;>45?!6&L;5140,()*#+%&’#$Q13>;O4;>4F156.4&;10!.%;401&@>1&%=>45489:=%0!5131&%;4’,-0"%5%0H34&4=%0?!&?45&4’,89:=%0&!;?.!04./54.%;4’;!F%??1&40;5%1&0GCE%&’G9EI=>1?>=454=1’4./604’1&@>1&%,R>454"!54,1;=%0754’1?;4’;>%;;>44F!.6;1!&!"89:=%0%.0!54.%;4’;!;>4Q6;%;1!&!"F%??1&40;5%1&0,8!Q4&4=*#+0;5%1&01&@>1&%%&’!;>45?!6&;5140Q13>;O4;>454?!QO1&%&;0!"F%??1&40;5%1&0%&’"14.’0;5%1&0[K,S],B65;>450;6’/0>!6.’O4?%5514’!6;;!’4;45Q1&4=>4;>4589:=%0;>454?!QO1&%&;!"

*#+10!.%;401&@>1&%%&’1&;5!’6?4’F%??1&40;5%1&0,

!"#"$"%&"’

9 T1&U,%&’A16$,G,-&1Q%.+15!.!3/(E&’4’1;1!&),#41V1&3:8?1L4&;1"1?%&’R4?>&!.!31?%.D5400,9MMN,KS9—KSE,

E A168,W,,W%&3W,,A16T,B,4;%.,H!.4?6.%5?>%5%?;4510;1?0!"071X434&4!"1&"4?;1!60O5!&?>1;10F1560A):10!.%;4’1&@>1&%,@>1&404$!65&%.!"+15!.!3/,EII:,EI(9):KS—SE,

Y W%&3T,J,,W%&3T,A,,B%&<,@,4;%.,-F1%&75!F4&;51?6.60;/741&"4?;1!60O5!&?>1;10(54F14=),@>1&404$!65&%.!"-&1Q%.%&’+4;451&%5/8?14&?40,EII9,YE(9):K:—SE,

: G4@,8,,W41$,U,,W%&3<,$,4;%.,*0!.%;1!&%&’1’4&;1"1?%;1!&!"%F1%&&47>5!34&1?L;/741&"4?;1!60O5!&?>1;10F1560,@>1&404$!65L&%.!"D54F4&;1F4+4;451&%5/H4’1?1&4,EII:,EK(9):YM—:9,

C U>%&3J,T,,U>!6$,T,,@>4&$,<,4;%.,@.!&1&3%&’04P64&?L1&3!"&6?.4!?%701’34&4!"&47>5!7%;>!34&1?%F1%&1&"4?;1!60O5!&L?>1;10F1560()8;5%1&),@>1&404$!65&%.!"+4;451&%5/8?14&?4,

EIIE,EE(K):CEM—CYE,K Z4&R,,A1%!H,,A6![,$,4;%.,8;6’/!&34&4;1?F%51%;1!&0!"C\4&’8975!;41&34&4!"%F1%&1&"4?;1!60O5!&?>1;1010!.%;41&@>1L&%,@>1&404$!65&%.!"+4;451&%5/8?14&?4,EIIE,EE(9):S—M,

S Z4&R,8;6’/!&;>47%;>!34&4010%&’34&4;1?F%51%;1!&01&;>40;56?;65%.75!;41&34&40!"1&"4?;1!60O5!&?>1;10F15604010!.%;4’1&@>1&%,R>4J10045;%;1!&!"8!6;>@>1&%-351?6.;65%.]&1F4501;/,

EII9,:—99:,

INE9 ===,;%&’",?!,6X/V!65&%.0 D5!3"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""

54001&2%;65%.8?14&?4 +!.,9K2!,9E EIIK

Dow

nloa

ded

by [

Uni

vers

ity o

f C

alif

orni

a Sa

nta

Bar

bara

] at

17:

58 2

9 Fe

brua

ry 2

016