1、!卷#期$%年&%月!日微生物学报!#$%&()&(*(+&$,&-&$!(#):()*!+,-./01$%!#$%&($()*:!#$%&!#()*+$,&+,-%(&.#$%&%/01$&((%(&2);23%)4)%3,+#%/01$&,5,!#$%&4)%6)75/%)-%(&.#$%&8,5,)+1&.9,:,%;7,($%0?+,A,$&B&$:,)5$#(!0?A3%!3%!#)0%),5;%&.$&6(#1%)4 A,:*3#($3*$%($;?37$:1C1&65%(+4,.(4+&+),)(-)%:$!D&()$%6.,)/)%:$2)+1$%6+)-(0)%
2、:$#D(&,$%!#$%(#)!#$#%&$()*+,-&,.(/&(*+,!0*.*-&)-#./0%0/(1*0#$0(123#4/#.+.(,*/&(5*.2*(&30-)(43(%!#$%(#)!#$#%&$()*+,-&,.(/&(*+,!0*.*-&)-#./0%0/(1*0#$0(123#4/#.+.(,*/&(5*.2*(&30-)(43(%EF!G H&,IF!H$&,JK H(&,0I?!D$3$&6,LIF!G M$%31((!#$%&(&)*+$%,(-,).)/0,12$(3(,4%5,&0)*67,($,8,(/9$)$%(,67,($)5+&-%
3、6&:5+&-%6&:9()$&6#1,5(77,)%/$%,&,;$C%#$+%+(),.7%&6+(#(),.#%&6(,5%,(60()/.)55:5 5,.$4,5 G(,))$&/$51/)7$&N$&6.%,01$&4 A1,.$5,5,.#%&6(,5%,1$O$#,.1,7%)16$&6%/#1,O5/$&,,%=,.P$.&,&.(+,)#$%&%/#1,O%.5()/+,4 FG)73&,6#$:,,)%.51;,.O+#,)$(7(.,5$6,.5#)$&EH%)=5$5%#,./)%7#1,6 O.,)%/.$5,5,./$514 4#1%6,&$+$#555
4、),:,.#1#EH%=5:$)(,%#%&6(,5%,&.C,O)/$51(!$(,)%,))O$);,)$#%&,$&3,+#$%&+1,&6,,=$#1#1,J9#%O,$&6+(#,.5#7#8&%(+/(66%/$51(#7$8&%#+/(6/$51)&.&72 8&%(+/(66%/$51(*72 8&%$+/(6/$51)),5;,+#$:,4 A1,&*)8!F 6,&,5,Q(,&+,%/5#)$&EH%.,7%&5#)#,.1$61 5$7$)$#(229)=$#1;7)&)%6,&,#$+&5$5 51%=,.+,)55%+$#$%&%/5#)$&EH%=$#1;
5、4 9$5.$5(O5;4#,52,2,9$4 F.$#$%&,:$.,&+,%/#1,$.,$/$+#$%&$&+(.,.$&7%);1%6$+,;15$%6$+&.O$%+1,7$+.#4 A1,;#1%6,&=5 5,&5$#$:,#%+$/()%$7,((%!:)&.+,/#)$%&,5%.$(7((%!:)4 4),5,#(.,5+)$O,5;4 9$5.$5(O5;4#,52,2,9$/)%7.$5,5,./$51/%)#1,/$)5#$7,$&01$&47089#-/7089#-/:;7)&);&*)9!F4:4(*.+0-:4:4(*.+0-:N2(2;#6*.0(&
6、6#/0:;#6*.0(&6#/0:F5-&6$0!;:5-&6$0!;:%&3$%2($%)%#3%(3%!?;4=!?!?;4=!?;7)&)$2&%,:9$5.$5(O5;4#,52,2,9$5#1,+(5,%/;5#,(),%5$5,=1$+1=5/$)5#.,5+)$O,.$&=$.;%;(#$%&5%/=1$#,;,)+1(+)%)($%,2$(:5)&.5#)$;,.O55(+)%)(5$=$&,.,5)$&2($&01,5;,P,,B*F&4 F#1,&.%/#1,5$#$,5&.#1,O,6$&$&6%/#1,5,:,$,5%/$%#1+,(),#1,O+#,)$(7
7、/%)#1,/$)5#$7,O,+7,+%&%7$+$7;%)#+(5$&6 5,:,),;)%O,75,&.1$61 7%)#$#$,5=,),),;%)#,.$&D;&,5,Q(+(#(),$4 K&?()%;,,A%)&C%,&$.4(.,5+)$O,.#1,/$)5#%(#O),P%/;5#,(),%5$5$&*;$&$&22%,=1$+1/,+#,.3(:,&$,6$#1,.5,O),7(#$%:5$:%$&$)+(#(),5$,&%)#1=,5#,)&),4*%,),/#,),$#+(5,.,+%&%7$+%55,5$,7)$&,+(#(),%/6$#1,.5
8、,O),7,5,O55&.5%,().$5(/$.(5,5&.).$5).$)$,2,.$#,)&,&+%()$,5%/?()%;,&.%#1,)+%(#)$,5()24 45#,(),%5$5+%$&(,5#%O,5,:,),;)%O,7$&$,&5$:,+(#(),%/.$/,),/$515;,+$,5$,2,.$#,)&,&.D;&4 B$),+,,01$&=5+%&5$.,),.#%O,/),%/$51;5#,(),%5$54A%&6(,5%,$5&,+%&%7$+$7;%)#)$&,/$51 5;,+$,5,&.$5:(,./%)$#5);$
9、.6)%=#1&.6%.#5#,4 R:,)#1,)5,#1,),15 O,&5#,.,7&./%)#%&6(,5%,$&01$&,$#15 O,&.%$&6=,$,$,)$%&7)P,#4 0%&5,Q(,,$,&5$:,+(#(),%/#%&6(,5%,=5.,:,%;,.&.),5(#,.$&,=/$51.$5,5,$&01$&4 A1$5;,),;%)#5$5%#$%&%/;49$5.$5(O5;4#,52,2,9$/)%7.$5,5,.#%&6(,5%,$&01$&/%)#1,/$)5#$7,4#A5=B!5:C 5!5:C 5;AB=D?;C!#BE,0-.0
10、(&%$0F*,.0(&%(/-0%10(&!#BE,0-.0(&%$0F*,.0(&%(/-0%10(&FK$%408 7+1$&,(F.:&+,.&.,B*F);AS86)%5,6,9!F;()$/$+#$%&P$#(AS80%4,J#.,9$&,44 8401$&);4B073A:,+#%)(*1&61$*&6%&);F4K$%?55#,7(,%=,.P$.&,&.(+,)#$%&%:,)#1,!#$,%&!(&)*+#,-.&,)/,&-.)*0)*1#&,2/)*&!3/+,)/%+-+(-&*4-(+#(/+5&!-&(-$)*+&%&(+-,&*#.)6-!)51)6&5+7&.
11、)*&()*6.+*6+#%)(/,655+6()*!8&.45+%+-+(&9+*&+4()6&55$,-.(/+/+4&(4&*6-+&,!5#,1&55!5+-,9)#*+$,45+*&*#!-&)*,&*#)*6:5&(+#).+#)&(+5$*(.&-)*+&1&-#$;(*.)*)+,-(?.)&5)*+:445+.+*(+#%)(/#*-(.)15$6+-5!#$%&()*+,-.%/%*0!#$%&()*+,-.%/%*0AB+-*)1/(6:5(:-+*CD:445+.+*(+#%)(/-(?.)E&25%+-+:+#(4-+4&-+:4+*)*)*+%*-(?.)&
12、5)*+,%/)6/%+-+6:*(+#!$:+,&/&+.6$(.+(+-5)#+(F.4-B+#E+:!&:+-($4+)&(&.&1*),)6&()*,/0!=5,(/+B)&!5+6+55 6*6+*(-&()*%&+(&!5)/+#!$4-+4&-)*1+-)&5#(,5#)5:()*,(/+!&6(+-)&5:4+*)*)*&5)*+,&*#4-+)*1+#.C B5:.+*(#:45)6&(+45&(+,CD:445+.+*(+#%)(/-(?.)E&25%)(/)*6:!&()*&(%&,-0%/!3*1:+5+,-.G:&-&*()*+#(69-+61*)H+#)+&
13、+(,-+#%+-+:+#(&+4&(/1+*)6)($!I-:4,)B+,)/(&B+-&1+%+)1/(1,*1)&,(+-)*&6()B&()*(.&)*(&)*+#&(,&2#&,-*.)*)%+-+)*,+6(+#!$)*(-&4+-)(*+&5()!4!))*J+6()*%)(/!3 456789:5;?-!)(.+(/#!D&6(+-)&5-+()5&()*%&.+*.-)!:*#(*1:+5+,55%)*1(/+.+(/#+6-)!+#)*+&*#)#+*(),)6&()*%&.+&4-+B):5$#+6-)!+#!K+!-&,)/(F;$(+.(D)L-)+:7,M-
14、&*6+)%&+.45$+#,55%)*1(/+.+(/#+6-)!+#)*(/+4-#:6(.&*:&5&*#)*6:!&(+#&(%&,-/!3/+7)#&+-+&6()*%+-+4+-,-.+#+4&-&(+5$,-.(/+=F;1&55+-$!786&-39)*%+.*-6)/*-3%):%*3).&;%&%+.-/*3+.;%/3+&/+(0!786&-39)*%+.*-6)/*-3%):%*3).&;%&%+.-/4/&(+D:,+-+#8&5)*+(D8)&*#6+*(-),:1+#&(E-.)*,-.)*,&*#(/+6+55%+-+-+:4+*#+#)*D8&*#6+*(-)
15、,:1+#&(E-.)*,-.)*!3/+!&6(+-):.%&(/+*,)7+#%)(/!*-15:(&-&5#+/$#+!3/+!&6(+-)&5 5:()*%-)44+#*(&644+-*+(&*#&,+%.)*:(+5&(+-,(&)*+#,-*+(%.)*:(+!$E-4/4/(:*1()6&6)#(/+*B)+%+#(/-:1/&(-&*.)*+5+6(-*.)6-64$!=.*%6%+*%/&-.&%*%9%*0!=.*%6%+*%/&-.&%*%9%*0=*()!)()6+*)()B)($%+(+-.)*+#!$:)*1&*()!)()6#)6).4-+1*&(+#%)
16、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