全 文 :书!"#$%&
,2016,36(6):1110-1116
犃犮狋犪犅狅狋.犅狅狉犲犪犾.犗犮犮犻犱犲狀狋.犛犻狀.
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456789:;< 犕犢犅=>?@AB
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J K:bJKLcS$Md%ef,EVW;Ig6hij8klmnS$4opq/rstIu(MYB)vw
;I
,
xystgz{|}~PCR^_,j
;Ig|},Zc6。
,4k MYB;I、lmnpq/ MYB;I,1k MYB;I \¡、¢
lmnpq/ MYB;I。stgz{£z;I¤l6¥¦。}~PCR
§
,2k犕犢犅;I(犕狀犕犢犅犑犕狀犕犢犅4)},Zc6¨©ªp,1k犕犢犅;I(犕狀犕犢犅330)}
,Zc6«¬p
,
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¯°¸H¹²pq±pqº[»¼
。
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;
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犐犱犲狀狋犻犳犻犮犪狋犻狅狀犪狀犱犉狌狀犮狋犻狅狀犃狀犪犾狔狊犻狊狅犳犃狀狋犺狅犮狔犪狀犻狀犅犻狅狊狔狀狋犺犲狊犻狊
犚犲犾犪狋犲犱犕犢犅犌犲狀犲狊犻狀犕狌犾犫犲狉狉狔
LIJun1,ZHAOAichun2,LIUChangying2,LRuihua2,LIUXiaoqing1,YUMaode2
(1.GuiyangcolegeofTraditionalChineseMedicine,Guiyang550025,China;2.ColageofBiotechnologyofSouthwestUniversi
ty,Chongqing400715,China)
犃犫狊狋狉犪犮狋:Inourstudy,8anthocyaninsbiosynthesisrelatedMYBcandidategeneswereidentifiedbyusing
bioinformaticstechnologiesandtheexpressionprofilesindifferenttissuesandstagesoffruitsduringthe
developmentanddeterminedbytranscriptomedataandrealtimePCR.Phylogeneticanalysisindicatedthat
4MYBgeneswereclosedtoMYBgenesfrom犞犻狋犻狊狏犻狀犻犳犲狉犪L.,犗狉狔狕犪狊犪狋犻狏犪and犣犲犪犿犪狔狊L.,while
only1MYBgenewasgroupedintooneclusterwiththosefrom犃狉犪犫犻犱狅狆狊犻狊狋犺犪犾犻犪狀犪and犕犪犾狌狊犱狅犿犲狊狋犻犮犪
L..Highleveloftranscriptsofmajorgenesinmaleflowerwasdeterminedbytranscriptomedata.Contin
uousdownregulationexpressionsof犕狀犕犢犅犑and犕狀犕犢犅4duringthedevelopmentofmulberryfruits
wereconfirmed,while犕狀犕犢犅330hadacontinuousupregulationexpression.Therefore,MYBgenes
havepositiveregulationandnegativeregulationduringtheaccumutionofanthocyanins.
犓犲狔狑狅狉犱狊:mulberry;anthocyanins;transcriptionfactors;regulation;expressionanalysis
lmn½¾¿¹À#$6ÁÂà n,ÄÀÅÆ4$6ÇÈ[1]。lmnÉ
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[8]。
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。
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,
noppqlmnS$4o/rJ
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[13]。
6犕犢犅;
I犆1pqlmn4o,½tO#$JKlmn
S$4oIuvÞ
[14]。
OJKLcS$Md%
efEVW;Ig6Rlmn4o/犕犢犅
;Iµ¶ow
,
Lcstgz{|}~
PCR^_JK犕犢犅;Ivx,yz;
IV®lmn¯°/³
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。
1 ef
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ð$Ì\<®¯¥¦4o
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1.2 V W
1.2.1 45 犕犢犅,@XY:ZBCD ENCBI
ªlmn4o/ MYB;I(¢ MdMYB10、
MdMYB110a MdMYBA, VvMYBA, ¡PAP1、PAP2,ZmC1)F;-°±,Lc
BlastP°²|VW;Ig6³´lm°±,
µ¶e10。·hiµ¶ß¸F;-°±N
query°±,²|¢、、tâ;Ig|³
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,
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。
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a,bcdef lmɼ³´SMART
(http://smart.Emblheidelberg.De/smart/set_
mode.cgiNORMA)¬½¾,lm;OÂ
¿y ExPasy (http://web.Expasy.Org/)。y
ClustalW ½¾£°±æ·,>êyGeneDoc·æ·
½¾ÀÁ
。
¿y MEGA4.0ÂÃ6Äof
(NeighborJoining,NJ)ÉųƽÆW。
1.2.3 45678gh 犕犢犅,@XYB>?i
jkHI stz{ÇÀµÈɵ;Ig`
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、
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、
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RPKM¶[Reads/(Fragments)PerKilobaseofex
onmodelperMilionmappedreads]Ò 犕犢犅 ;
I
g6¦
。
1.2.4 678lmBno V®lmnìÚ´
pÓRabinoã[15]ef。·-80℃¹V®
6JÔoÕ
,
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,
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²´530570nmâ_ªã¶,l
mn·Ï=犃530-0.25犃570。
1.2.5 45l 犚犖犃 pq`犮犇犖犃 9:
Ú
0.1gäååÀ³ÚJæ63çJÔo
Õiè
,
péRNAisoêì(TaKaRa)ëNì
§ìÚíRNA,IDNAJîÆêy50μLDEPC
Áï
,
¿y1%ð+ûñïòó²´Óxyôõ
bö´íRNA÷b。DíRNAvø,
DOligo(dT)15ð$,xy MMLV õstJ
11116 = >,ã:VWlmn4o/ MYBstIuRGH
(TaKaRa)4ocDNAù1ú,Ü-80℃¹y。
1.2.6 r\stEuvwxDm犘犆犚no ÌÀ
lmn/犕犢犅;Ilmn¯°÷àû
0/Â
,
=},ZêàüýþÿÆ
,
I
´wÚj
‘
30h’4k~(S0、S3、S4S5)
}ÇJK犕犢犅;Iµ¶ow¦,!w
lmn4o/犕犢犅;I。¬{ 犕犢犅;IË
K-"#°±¿yPrimer5.0$ö8kvw
;I}~PCRð$(1),¿y
2-ΔCt 。} ~ PCR % y SYBR
PremixExTaqTM(TliRNaseHPlus) TaKaRa
¥¦
,
%&|YäPÆ%É()
。
2
2.1 45=yz[\9:;<犕犢犅,@BCD
ªPI&*pqlmnS$4o犕犢犅
;IF;-°±
,
LcBlastP°²| 犕犢犅
;I
。
EVW;Ig6Rç8kÇÈS$4
o/ 犕犢犅 vw;I(2),+À
scaffold¬,2k;I1kZÏu,5k;I¹2
kZÏu
,犕狀犕犢犅犅 à11kZÏu,¸ H¹ß
,-./ex
。
yz;IÀ0261~478kF;
-
,
·u]þE29.2~54.0kD。3k MYB
lm
(MnMYB86、MnMYB11 MnMYB21)ÄÀ
1-Âlm
,MnMYB330ã5klmÄÀ2Âl
m
。
£°±æ· 6 k MYB lm (Mn
MYB308、MnMYB330、MnMYB86、MnMYB4、Mn
MYB11、MnMYB21)F;-°±6Ïà3
R2R3ɼ(41),MnMYBB MnMYBJä
5;I6#ß]
。MYBlm6Ïà5k3Ã
F-
(W)7;,MnMYBJ;Iù3kÃF-89F
-
(Glu,E),MnMYBJ MnMYB4ù4kÃF-
8ÍEF-
(Phe,F):F-(Ser,S)ÚÛ,Mn
MYBB MnMYBJù5kÃF-8;F-
(Tyr,Y)Û
2.2 45 犕犢犅,@XYB{=HI
j?´8kVW MYBlmGH,ÉÅj
MYBlmF;-°±³Æ½ÆW(42)。VW
5k MYBlm¸@Ù2kA,ÇÈ
4o/ MYB;I。MnMYB11、MnMYB330、
MnMYB308、MnMYB86BuDkMyb4(犇犻狅狊狆狔狉狅狊
j1 |}~rTable1 Primersusedinthisstudy
;ICÖGenename ¬Dð$Forwardprimer(5′→3′) ªDð$Reverseprimer(5′→3′)
犕狀犕犢犅308 GGTGGATTTGTGAGCAAAGA GCTGTTGTGGCTGATAAGGA
犕狀犕犢犅330 AAAGCCCACACCAATAAAGG TATCCATCTCAGCCTGCAAC
犕狀犕犢犅86 GCGCGAGCTATTTGTAATGA TGCTGTCAAGGCTTCCATAG
犕狀犕犢犅犅 TCATGTCCCATACAGCAGGT ATCCCTTTATCGCATTCACC
犕狀犕犢犅4 GTTTCGCGAGATCAAAGTCA TCTTAGTGCGACATGCTTCC
犕狀犕犢犅犑 CAAGCATCCATGACATCACA ATTGTCGCAGAGTGATCTGG
犕狀犕犢犅11 AAAGCCACCCGTACTACCAC CATCAACCGCAGGTTTAGAA
犕狀犕犢犅21 AATAATCCCAAGTGCCCAAG GTTTGGCAGAATTGACGATG
j2 45678[\9:;<犕犢犅,@XY:Z
Table2 Identificationofflavonoidsbiosynthesisrelated犕犢犅genesinmulberry
;ICÖ
Genenane
;IEth
GeneaccessionNO.
;IgØ
Genomelocation
ZÏukz
Intronnumber
À0F;-
Encodedaminoacids
u
Molecularweight/kD
}FãòÊ
Isoelectricpoint
犕狀犕犢犅330 XP_010104477.1 scaffold485 2 305 34.1 9.22
犕狀犕犢犅86 XP_010089049.1 scaffold297 2 298 33.8 6.56
犕狀犕犢犅犅 XP_010096371.1 scaffold155 11 478 54.0 8.79
犕狀犕犢犅4 XP_010107438.1 scaffold759 2 304 34.8 8.79
犕狀犕犢犅犑 XP_010109645.1 scaffold751 1 319 36.3 8.81
犕狀犕犢犅11 XP_010107990.1 scaffold959 2 364 41.0 5.85
犕狀犕犢犅21 XP_010099251.1 scaffold20 2 327 37.1 6.06
犕狀犕犢犅308 XP_010090332.1 scaffold137 1 261 29.2 8.97
2111 ! " # $ % & 36G
HIJR2、R3lmɼ;KÃeLJ3ÃF-(W)7;;MnMYBJ.MV(XP_010109645.1);MnMYBB.MV(XP_010096371.1);
MnMYB21.MV(XP_010099251.1);MnMYB86.MV(XP_010089049.1);MnMYB4.MV(XP_010107438.1);MdMYB110a.
¢
(BAJ24837);MdMYB1a.¢(ACQ45201);MnMYB330.MV(XP_010104477.1);MnMYB308.MV(XP_010090332.1);
MnMYB11.MV(XP_010107990.1);DkMyb4.Bu(AB503701);PAP1. \¡(NP_176057);C1.(CAA75509);
P.(AAK81903)
41 VW MYBlm£°±æ·
TheblacklinesindicatedconservedR2andR3domain;conservedTryresidueswasshownbysolidbox;MnMYBJ.犕狅狉狌狊狀狅狋犪犫犻犾犻狊
(XP_010109645.1);MnMYBB.犕狅狉狌狊狀狅狋犪犫犻犾犻狊(XP_010096371.1);MnMYB21.犕狅狉狌狊狀狅狋犪犫犻犾犻狊(XP_010099251.1);MnMYB86.
犕狅狉狌狊狀狅狋犪犫犻犾犻狊(XP_010089049.1);MnMYB4.犕狅狉狌狊狀狅狋犪犫犻犾犻狊(XP_010107438.1);MdMYB110a.犕犪犾狌狊犱狅犿犲狊狋犻犮犪L.(BAJ24837);
MdMYB1a.犕犪犾狌狊犱狅犿犲狊狋犻犮犪L.(ACQ45201);MnMYB330.犕狅狉狌狊狀狅狋犪犫犻犾犻狊(XP_010104477.1);MnMYB308.犕狅狉狌狊狀狅狋犪犫犻犾犻狊
(XP_010090332.1);MnMYB11.犕狅狉狌狊狀狅狋犪犫犻犾犻狊(XP_010107990.1);DkMyb4.犇犻狅狊狆狔狉狅狊犽犪犽犻Thunb.(AB503701);PAP1.
犃狉犪犫犻犱狅狆狊犻狊狋犺犪犾犻犪狀犪(NP_176057);C1.犗狉狔狕犪狊犪狋犻狏犪(CAA75509);P.犣犲犪犿犪狔狊L.(AAK81903)
Fig.1 ProteinmulialigmentofMYBgenesinmulberry
犽犪犽犻Thunb.,AB503701)、 VvMybPA1(犞犻狋犻狊
狏犻狀犻犳犲狉犪,NM_001281231)、C1protein(犗狉狔狕犪
狊犪狋犻狏犪,CAA75509)| P (犣犲犪 犿犪狔狊,
AAK81903)@Ù GroupⅠ。MnMYB21 ¡
(犃狉犪犫犻犱狅狆狊犻狊狋犺犪犾犻犪狀犪)PAP1(NP_176057)
PAP2(NP_176813)、¢(犕犪犾狌狊犱狅犿犲狊狋犻犮犪)Md
MYB10(ACQ45201) MdMYB110a(BAJ24837)@Ù
GroupⅡ。MnMYB4、MnMYBB MnMYBJNà
@Ù2kA,3ÂØÊÃF-7;=ÿàß
¥¹Â
,¸
HpÇÈS$4oDõä5
ÛAO;pq
。
2.3 45犕犢犅,@B>?iHI
Lc·VWstgz{P²||
,
,Q
8k 犕犢犅 ;IRl6à¥(43)。
ä6犕狀犕犢犅86Rl6],ä5g
6N಴YstO
;犕狀犕犢犅犅 Rl|Ð6
ߥ
,
¬
、
Í|ÎÏ6߸
;犕狀
犕犢犅4Rl6H²´Y,ä5g6
߸
;犕狀犕犢犅犑ÍRl6²´Y,
¬
、
ÎÏ|Ð6N಴Y
;犕狀犕犢犅11Rl|Ð
6S÷àߥ
,
¬
、
ÍÎÏ6ß
T
;犕狀犕犢犅21Rl6],Í|Ð6à
31116 = >,ã:VWlmn4o/ MYBstIuRGH
46CÊzU;À1000@`,bVÊ -C¨W;XÛXYYÖ
42 MYB;Iµ¶½Æ/³
Thenodesinthefigureshowthebootstrapvalueonreplications,andthescalerepresentthegeneticdistance
Fig.2 ThephylogenetictreeofMYBgenefamilyinmulberry
43 VWlmn/犕犢犅;I
Fig.3 Transcriptionanalysisofanthocyaninrelated犕犢犅genesinmulberry
T
,
¬|ÎÏN಴Y
;犕狀
犕犢犅330Í|Rl6ߥ;犕狀犕犢犅308
Rl|Ð6¥
,
Í6àT
,
¬|Î
Ï6߸
。
2.4 4:N678
mBoD
V®½uÞl
,
EBÕY}QZo[
\DI]]^35~40d,¬{}"_}õ`
·ak},Z~S0~S5 ã6k~,ä6
S0 bBÕQc,S5 }QZo[,dk~]
efg7d。Lc´~lmnÏ,,QE
S0~S4 5k~6lmnÏ߸,7S4~
S5 S_c6,lmnÏÿÆj·h8þ,ì§
yk~½lmni4o~
(
44)。
2.5 犕犢犅,@XY4NBjkHI
j½U!wVWlmn4o/
犕犢犅;I,Lc}~PCR;Il
mn4oßiük~
(S0、S3、S4 S5)V®
6¦
。
8k;IjQ
vx
(
45),犕狀犕犢犅308},Z4k~S
÷àߥ
,犕狀犕犢犅犑犕狀犕犢犅4kl}
,ZjQªpmn
;犕狀犕犢犅6},Z
6jQo¥p߸
;犕狀犕犢犅犅
jQéªpq¬pmn
;犕狀犕犢犅330
},Zc6jQ¬p
,
=},Z
c6/3jh3þ。
4111 ! " # $ % & 36G
45 VWlmn/犕犢犅;I},Zc6
Fig.5 Expressionanalysisofanthocyaninrelated犕犢犅genesindifferentstagesofmulberryfruits
S0~S5lê0、1、2、3、45};ª。
(
Jr46SzUª)
44 V®,Zc6lmnÏ
S0S5represent0,1,2,3,4,5and6weeksafter
fulbloomoffruits;Thesameasbelow
Fig.4 Thecontentofanthocyaninsduringthe
developmentofmulberryfruits
3 s F
V®,Zc6I]jtl
、
Xu
、
S_
、
v
w
、
x]
、
ÿyãNzÿÆc
,
ÿycB
,
7ÿy~ü«~fZlm
n]4o
[16]。
I´
,
V®6lmn¯°ÚÝÀ
}S_ê/rJ|pq;Ivx|ÿ
。
MYBstIu½#$6>`DstIuµ¶B
,
Ì4RMYB、3RMYB、R2R3MYB| MYBre
lated4k}go[17]。JK§R2R3MYBst
Iupj@SÛApq
、
~Æ|~
q¼
、
S$S$köã£[S}SÆ
c
[18]。
lmnS$4opq6
,WD40、bZ
IP、R2R3MYB、bHLH| HY5ãstIu¸HN
o£[,4
,
ÓÉ;I°±6
xN
yÃ4
,
E7ù¼ÙO/;I
,
p
qlmn#$6¯°
[19]。
OJKEVW;I
gz{P6Rç6klmn4o/ MYB
µ¶vw;I÷àuÞR2R3É。,
Q MnMYB11、MnMYB330 MnMYB308PI
&* l m n 4 o / MYB ; I Dk
Myb4[20]、VvMybPA1[21],¸H½V
Wlmn4opq;IB
。MYB²
pqow
(
AN2、AN4),,Qjz±
pq MYBlm(AtMYBL2)[22]。
tâ FaMYB1 lm C j÷àk;°
pdLNLD/ELXiG/S,¸ Hz²pqIu4
;I
,
E7Ypqlmn4o1
[23]。
OJ
KRçlmn4o/6kR2R3MYBv
w;I},Zc6jQvx
。
犕狀犕犢犅犑犕狀犕犢犅4kl},ZjQªp
mn
,
ªp1/61/8,=
6Nàä5#$6ÇÈ4o/犕犢犅;I
,
ä6I´¸H½VWlmn4o
±pq;I
;犕狀犕犢犅330},Zc6jQ
«¬p
,
=},Zc6/3j3þ,¸
H½VWlmn4o²pq;IB
。
I´VW
犕犢犅;Iµ¶·lmn4o¢、tâå
S¹²pq±pqº[»¼
,
½p犕犢犅
;Iow¹ß]6#
。
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[3] , .köÆ â¡¢ìÚV£lÃK¢¤
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