免费文献传递   相关文献

Research Progress in the Biological Basis of Legume Pod Dehiscence

豆科植物裂荚生物学基础的研究进展



全 文 :!23" !5#
 Vol.23  No.5
! " # $
ACTA AGRESTIA SINICA
   2015$  9%
  Sep.  2015
犱狅犻:10.11733/j.issn.10070435.2015.05.004
ÈÉ*+ÊËb+­KÌ0”•±²
Q R,/8S,@TU

Nñ():AÀ¬­C:=Vº  2*+Nñ()°•+0

78  2730020)
45

Cu5h°89•;<‡/+ú:BŠYSˆRg

wCu*NXY7•ÐJ0*ë)—

PY7:ú\á
UCB9“

GHÎìY7úë

h°89BåCugháG»ÓCDBTY³¬’Š

˜œ¾D¿€+

:
X+=z7:9+3eã¨Kh°89Cu;iés,ÃK³0”•BĒ†ÅÌB˜™gä;iHkâ。
678

h°89

Cu

CuÝ

Cu‡E
9:;<=
:S330.3    >?@AB:A     >CD=:10070435(2015)05092709
犚犲狊犲犪狉犮犺犘狉狅犵狉犲狊狊犻狀狋犺犲犅犻狅犾狅犵犻犮犪犾犅犪狊犻狊狅犳犔犲犵狌犿犲犘狅犱犇犲犺犻狊犮犲狀犮犲
LUODong,WANGYanrong,LIUZhipeng
(StateKeyLaboratoryofGrasslandAgroecosystems,ColegeofPastoralAgricultureScienceandTechnology,
LanzhouUniversity,Lanzhou,GansuProvince730020,China)
犃犫狊狋狉犪犮狋:Poddehiscenceisaveryimportantreproductivetraitduringtheevolutionoflegume.However,
thistraitalsocanleadtoseeddispersalbeforeharvest,whichcausesheavylossesonlegumeseedproduc
tion.Inordertoimprovetheseedyield,lossofpoddehiscencebecomesoneofthekeyagronomicaltraits
targetedbyhumanbreedingselection.Thestudyadvancesofmorphology,physiologyandmolecularbiolo
gyonpoddehiscencetraitsoflegumewereelaboratedinthispaper.Moreover,theexistingproblemsand
developmentaltendencywerealsoprospected.
犓犲狔狑狅狉犱狊:Legume;Poddehiscence;Dehiscencezone;Dehiscencegenes
EFGH
:20140806;IJGH:20141220
KLMN

¬­C:‡Ã˜™@kÁØ
(973ÁØ)“,N、:ËNå;ãùìúB:9+‡Ã”(2014CB138704);¬­®¯°+‡±/‚²
³

-o”(31272492)´ µ
OPQR


(1991),¸ ,>3×»,¼½˜™:,˜™NLGNyÅ9TY8:9•–,Email:luod13@lzu.edu.cn; ÇÈÅÆ
Authorforcorrespondence,Email:lzp@lzu.edu.cn
  h°89Cu5«•á$#mu?–u@B
Á

ABm^CU
,¯
ÌCÿY7BA_
[13]。
K
89®ñˆÎ

Cu589u#;<‡/+€áB
CDSˆRg

KY7Âÿ

SU=ûü9Y‹&g
[ŒCDVÛ

w•():ú+

h°89C*
h
(犌犾狔犮犻狀犲犿犪狓L.)、Dh(犘犻狊狌犿狊犪狋犻狏狌犿 L.)、;

(犔狅狋狌狊犮狅狉狀犻犮狌犾犪狋狌狊L.)、ô©Fqh(犔狌狆犻狀狌狊
犪狀犵狌狊狋犻犳狅犾犻狌狊L.)、Th(犔犪犫犾犪犫狆狌狉狆狌狉犲狌狊L.)!
BCu¦*NXY7•ÐJ0*ë)—

PY7:
ú\áUCB9“
[4]。
GHÊڎ{=:úBá
T

ÎìY7úë

ŒTåCuGYáG»ÓCDB
TY³¬’Š

T˜¾D¿€+

:X+=z7
:9+3eã¨Kh°89Cu:9+RgB‡Ã
˜™;iés

ÎHÒ+”•BĒ

ÃKÅÌU
kh°89BCuƒ@˜™BC:=gä;ik
â

ƒ•G;ŠŒ ›Cu˜™ÎŠkBš>

1 ÊË0,-­”•
©˜™PJ

͋h°89CughBDo
8muu÷s°€†mu¿§+RSƒ.A
[4]。
1.1 mËÍÎ,-
mus°€BDo|â5½ê89Cugh
WXBÞE’Š
。Caviness![5]•*h+°?Å*
hCu58Y7*w”•ƒ@g

ˆY7*wX“
67–muBHªÈ

£Ì
,Luo![6];ŠŒP=
H*hCu5=muBHªÈá—ríƒ@

Mos[7]q@A;E¼Y7á$#,€muUCÈT
€mu&

Cu5Ÿì

ˆ,€muBHªÈÈT€
Bmuw

d–mu€˜™Bßl

+ÆӍH
Ÿ‹ZB@A
。Tsuchiya[8]z{H8*hCugh
ƒ@B‹eÒ~(Igh

@AåCuGYmuB
! " # $ !23"
HªÈ—r*¦DCuBGY

ˆmuu¨

ª¨



JK¨

uLBH¨!Ò~«¬•Ð2yGY
+ÃVJ—Do
。Suzuki![9]Bw‡qPJåC
uGY8DCuGYBmu•u¨

ª¨}†H¨
N/[W”•—rDo



˜™«
[10],
8
9BCuRg]8mu‹p€RSƒ@gW*

ˆ¾D5ëmuœ°l%+WDIjÅBœwn<
]½êB

1.2 mËÏÎlÐ
‚•œ°àHl%DokHs•CuRgB
WX

h°89m=Gum

ë2æuaàá,Ãë
Á

ABm7A

Y7Ë̕+MN‚

8DCu":YƒÈ

åCuYBmuŠ÷
RêB¿§+RS

C•*h+

Ž{Y=":Y
“ŠeCDBœ°l%Do|â5ëABm^I
OüýLBPIQüýH¨WXˆú:B

¿":
*hŎ{*hBR<‡/+

CuݚNBPI
QüýBü:L—r H

ñ*HuazŸ]FB
Úç

E£iñfSHmuUC
[11]。
hs

mu
ÁBm^IOQBu¨=H¨}†uLBH¨8
Cu/¨á—ríƒ@

ҕ*h;<‡/+qT
–Cu/¨ïwBNL@k
[12]。
•Th+

÷å
CuGYBu@P/*;<Š÷WªsBCU

˜™PJ

Ðyl%â^izVÁ

ABm^UC
“]EÅBW–WBБç=›Gdç

E£|
iÊÚThBCu5
[13]。
•"DhÊ
(Vicieae)8
9+

åCuGYmu‚m@BPIæH¨šDC
uB":Y¿

]ÓAï“
[1415]。
•;E¼O
(犔狅
狋狌狊)+,åCuYT–+m@hù;<
。Yang![16]K;E¼OBmu¿§+l%;
iH˜™

@ADCuYmuB+m@hùšåCuYBmuì

\üýNölm—ó


YmuB+m@hù#

ÉPAGåCug

qÍ5E

+m@hù<
/¨8Cu5áðƒ@

hs

˜™@A;E¼m
uB+m@‚PIu6ÕÄNÂ8umBu6NL
f4ã

ˆ8apXYhJKNL]Ó

qÍ5E

+m@‚PIBÕÄN¢;HmuXYUC
[17]。
}‚˜™lmPJ

•h°89B;<‡/+

A
mu‚üýæ•NY‚†pë‚B¿n

m@l%
Bn<

+m@hù
Ð÷2nX
Ñ89Cu/¨ÊÚ½×S|Z“Cu›L

¿ˆ
;<åCuBGY

£s

8CuRgƒ@BR꿧+RSn<
Çn@:•muBCuÝ]Ò­#

ÐPJmuB
CuݕmuUC‡/+@ÒCDÅ¥

CuÝ5
«Á

ABm^2æuaB7AÝ,ëŸæ=ua¡
?*BHLüýæàá

HLüýæhùì

ˆŸæ5ëap¡?@TáB1CõBüý
&

¾Dëmhù
CuA_B
@:5EG”•1eCuÝ,£ÝÜB¤züýz
U

vÌë¦ÚçBÅ¥ÑmuUC
[18]。
[B
5

¿§+@Ah°89umBÁ

ABm^BCu
Ýl%ÃWƒX

‰Doš*

ÊC

•ABm^+
m@æ78We‰M2(eCuÝ

ˆ•ÁBm^
+m@æ IOQl%Bt\zá2¤z。Ð8
rBo°89BuãmÁ

ABm^5ë2eK]
BuaàáBw‡WŠX
[18]。
’]}ú:ÐYl
%Do|â5EGum5ë1e©®;<ˆ…B。
89;<‡/+

©®+E©<@TáABm

]}
®ñàHƒKS(

ˆ©®¡?«@"K@TáÁ
Bm

]}*”•2eua^AB__。3VE8
CuBêΤYGÁBmƒ`@

ë¦Á

ABm
^Bl%Do

ÁBm^UC]FB›GçJ—w
¦ABm^

]}Cu*\ËA•ÁBm^
。Ti
wari![12]•*hmu¿§+˜™+@A,v&½ê
CuBÞE5ÁBm^IOQ2ea“Ÿæüý
BÊ¿

ëWX*hGY

IOQaB“a)
*

mu)DUC

¯ˆ

qw‡«ABm95
CuBêΤY
。Dong![11]•*hCuBvZ˜
™+@A

muUCB@ql%ÃW5Cu݂B
Ÿæüý

ˆ5ABm^CuݚNBPIQüý

PIQBH¨¦ÓA67muUC“]FB›G
ç

Ð|â5ÉüCuUΦABmBçP½

é‚]s

h°89CuRgWø8muu÷
€RS”•ƒ@g

¿muœ°l%z{

h°8
9CuRgø8muBCuݔ•–J—Bƒ@
g

¯ˆ

³0@¦muCuÝB˜™ø•*h+
ƒKá$

bCDh

cdDh

Th!CD,NÅ
9+øew‡



F•Ÿ‹h°89+./U
kmu¿§+˜™

G;ŠŒI™h°89Cuª
«=›Lfk‡Ã

2 ÊË0bk­”•
Cu51eMNB9X=:X:<‡/。mu
•Sá:Xá$Ì

CuÝ@:ŠAă@BÂR
829
!5# = Ê!:h°89Cu:9+‡ÃB˜™;k
À‘

CCuÝüýB,<

m@“}†a“Bl@
Bã<

Ÿæ‚üýBgxçB9“=+“¿æB
Ê¿!

Ð÷À‘ƒXÅ¥½êHmuUC

•*h+
,Christiansen![19]¦*hmuB@
T“#zG3eðà。!1eðà5´Þmu:u
#

Uo#Ì4~5u),Ÿæ\˕Á、ABm^u
aB¡?€á

M2uL

Ãëý“7hi£ƒ7

!2eðà5nÞmuá$#(Uo#Ì6~7u),
CuÝüýKzUÎj“

ý“æk¿

џæüý
ƒJzŸ

A“a

O5Íý“7hlê–7A
Å¥

!3eðà5mÞmub#(Uo#Ì8
~9u),Ÿæüý*ëMC,ý“7h›“,uan
ùzU

˜™PJ

CuÝHLüýKzZ“ÌK
Ÿæüýú:B‚†gÚç5*hCuBÓAÞ
E

ˆ‚†gÚçBú:5ë¦u@‚BHLüý
•RkBNY‚z
‚m@=sm@[ë
HLüýàá

ÕÄN•NL‚ƒ

w[8mu
BK6máoã

EG•‚

sm@æ+

üý6B
NYWX

]}

“KÌÐ2eüýæ•NY‚(C
ML=KL

Bv*Б/¨”•Do

ÐYDÆ
БÑHLàHú:Úç

Å¥•Ÿæ

¿ˆNXC
u
[20]。
3‡/WFDpµsç

ë1e‚Z›Lú
:BÚç]½N

G¾QCu

Dh

cdDh

T
h

qh
(犞犻犮犻犪犳犪犫犪L.)、ô©Fqh、;E¼!h
°89q”•y„BCu›L

C•;E¼+

á
$#

u@¿LüýKz9“B¶5ì¦HLàH
+BPIüý

Ñu@W–WБ=rsÃú:Ú
ç

¿ˆ½êCu
[17]。
¾QCu•:u
(犃狉犪犫犻
犱狅狆狊犻狊狋犺犪犾犻犪狀犪L.)、tA(犅狉犪狊狊犻犮犪犮犪犿狆犲狊狋狉犻狊L.)
!rBo°89+q@A

:uBm=Guã
m

¾DzGa

V•p†Ÿæ3¤z,Ÿæ•a8
V•p“Bz…m^€á

uãmá$Ì

apæ
‚¤HLüýhù<ÉaBsæ¿LüýБÑ
Õm=ú:Úç

¢Ña?V•p¡?BŸæz
U
[21]。
tAm==:uŒƒ„Bl%

[5ë
2eu@Bvb©àáBvbE´Ì@TáGuã
m
[22]。
tABmuë2eaàá,•Ò@T#,
muB2eaë1WÇkBmhù<üý&7
A

îŸæ

uãmS|á$Ì
,2ea•ÚçÅ¥
šL2eNLk[18]。Ð÷lmPJ,rBo°B
Cu›L8h°89BCu›L”•b‹ƒ„g

CuÝ+Ÿæüýi£zŸq8ÔYK¿t

C‹xëwxyst

PIFt

‚ö1,4βzx
x!

½êBüýL=ý“æm{Ê¿@

˜™
PJ

CuÝàHB@Tñ HCughB@:

Ò
+“tBú:=Ÿ@ÒHCDÅ¥
。Agrawal
!
[23]
jkH*hmu+B2YK¿t(PIFt=
‹xëwxyst

BPg

@Aá$#DCuGY
+BPIFt¿mCuÝ2#ÅCuÝ

‰Pg—
rñ›

£A_•åCuGY+|

8£X
“

‹xëwxyst•åCuGYBCuÝ+Â
Ÿ

Pg—rì¦DCuBGY

z{@A

PIF
t}<ŸæBüýLÊ¿

ˆ‹xëwxystk
¿ý“æBm{

¢;u@¿<
。Christiansen
!
[19]
K*hmu;itz{BlmPJ

•´mu
:u#CuÝAH‚ö1,4βzxx=‚ö‹
xëwxyst

ɕnmuá$#ÄÅv*õ

PÄz{@A‚ö‹xëwxyst8Cu0¿L
üýæBeC@

•rBo°89+

C:u

tA!

uãmBCuÔ/¦uãmLBÊ¿†Ÿ
æBzŸ

ˆŸæBzŸÔ/¦üýL~t


pG‚öxëwxyst¢Ñ+“¿æeC
[24]。
t¯*h+B‚öxëwxyst8tA}†:
u+B‚öxëwxystBX†ƒ„gO
57%=43%,‰PÄéÂq”•Do,w5ÒÓ[
8ŸæüýLBÊ¿@

é‚]s

h°89CuB:X›L|8ŠG
2y:!1y5¾QCu,îë¦u@B‚:HLà
Hæüý]ú:BHçWXÑ7AÁ

ABmB¿
LàHÚC

u@UC

3‡/WFpµsç

¾D
ë‚Z¾Q›L½N

!2y5 QCu]tRÖ
Cu

îCu݂BüýL~tKCuÝBŸæ
üý;iC¿

Ÿæüýi£zŸ

u@UC


/Esç67

CtBK¿
),
‚Z¾Q›Lþ@
җrť

Cu@Wú:Úç
)。
h°89CuR
g8¾QCu›L] QCu›Lƒ.A

]«Æ
ƒXÅ¥NXuazŸÑÕY7Cÿ

hs

rB
o°BCu›L8h°89BCu›L”•b‹ƒ
„g

Ð÷ƒ„gG2y89CuBÌ8˜™Î
ŠkBpF

3 ÊË0;Ÿb+­”•
Cu¾DëÁÂE7RÖ

€8Cughƒ
@B‡E5».ŒB³¬

ҊN/|} ß0
ÓKCughBv‚

hŠN/|}¿#£ghÁ
ÌݕBz7Á›L
[25]。
d–‡Eà+=z7
:9+“?BŅ

¬‚s+ƕÐN/ÃH*ë
929
! " # $ !23"


ÃxՊAÄCDám

³0

©./UkC
ughÁÂgz{Bh°89¾D

*h
[2627]、
D
h
[2829]
=ô©Fqh
[3031]。
£s

Ah
(犘犺犪狊犲狅犾狌狊
狏狌犾犵犪狉犻狊L.)[32]、ƒh(犞犻犵狀犪狌狀犵狌犻犮狌犾犪狋犪L.)[33]、
cdDh
(犞犻犮犻犪狊犪狋犻狏犪L.)[34]、wTh(犔犲狀狊犮狌犾犻
狀犪狉犻狊Medic.)[35]=ah(犞犻犵狀犪犪狀犵狌犾犪狉犻狊Ohwiet
Ohashi)[36]+qƒ@w‡。
3.1 ÀÈ
•‡=B20$,d–ÁÂz{NÓBð„,*
hpëgh‡E.
(quantitativetraitlocus,QTL)
kY©áG˜™*hCughB¾èNÓ

P1)。
\1 ÀÈ9Ñ®Ò0ÓÔÊË.30犙犜犔狊/Kž
Table1 SummaryofthereportedQTLs/genesforpoddehiscenceinsoybean
>T
Parents
z{NÓ
Analysismethod
]•7„f
Linkagegroup
v#±
Nearestmarker
¿#no
Variation
explained/%
LODõ
š>˜§
References
Young
(C)
PI1416937
(C)
ANOVA
D1b A725 5.3~7.2 -
E Cr2741 6.0~7.1 -
J B1221 39.1~42.5 -
L A4891 5.0~5.6 -
N A808n 4.1~5.7 -
[26]
Toyomusume
(C)
Hayahikari
(C)
CIM J Sat_093 >50.0 13.8~15.6 [27]
Tokei780
(C)
Hidaka4
(W)
CIM
E Sat_124 9.6 2.7
J Satt215 16.3~21.8 3.3~4.9
[37]
Keunolkong
(C)
Sinpaldalkong
(C)
CIM
A1 Satt385 6.7 8.9
B2 Satt126 6.4 9.7
D1b Satt546 4.9 5.0
J Satt215 45.8 65.4
[38]
Jinbean23
(C)
ZDD2315
(C)
CIM C2 Sat062 49.4 12.15 [6]
  ò:C,Ž{Y;W,":Y
Note:C,Cultivar;W,Wild
  &•1987$,Tsuchiya[8]¥v*„¯ÓK8e*
hNöGA;i¥Þ

@ACugh5ë1~2e‡
EÖLB

ÃÞ.CughBL–ÁÂgG93%。
1997 $,Bailey ![26] K * h G Y ‘Young’=
‘PI416937’B120eNöGABCugh;iQTL
z{

kYÅ1e¾i=‹eœiQTL.Y,zÆY
¦!2、!15、!16=!19…Þ\‚,þ¥KLg
‚öt®àu¨‹g
(restrictionfragmentlength
polymorphism,RFLP)¬±•–%ðH*hBÁÂ7
„׆

ÐG̅B˜™fkH‡Ã

•dÌB15
$<

*hCuƒ@‡EB˜™xÕH"MgB;
k
。2006$,Funatsuki![27]Ñ¥M@ݓÅ×Ó,
QTLz{H1eghzŸB*hCà®öAYf,
\ükYÅ1eÖL*hCughB¾iQTL.
Y

¼«G狇犘犇犎1,õkH3QTL.YY¦!16
™…Þ\‚

Y¦”œCMºÄ
(simplesequencere
peat,SSR)¬±Sat_093=Sat_366’“,hsÞ.Ð
2e¬±’“BÁžŸG2.9cM。2008$,Funat
suki![39]*¥4eŒWXÁÂÁbB*hÅG
‘

|üPJ狇犘犇犎1Y¦ SSR¬±Sat_093=
Sat_366’“,‰@A狇犘犇犎1B!Y‡E•WXBÁ
ÂÁbšŒ‹CY:

2009$,Kang![38]*¥SSR¬±K*hGY
‘Keunolkong’=‘Sinpaldalkong’BCà®ö\A
YfBCugh;i QTLz{,kYÅ1e¾i
=3eœiQTL.Y,3¾iQTL.YY¦!16
™…Þ\‚狇犘犇犎1.YB­#,3eœiQTL.
YzÆY¦!5、!10=!14™…Þ\‚。X“
~ÓvG*hBCugh•WXBÁÂÁbšŒ
ŠkBDog

X$
,Yamada![40]z{HÔe*
hYfBz7±

@AWW>TŒ¥‡ˆGA

N
öF2 ?BGA+,ÖLCughBQTL.Y[k
Y•狇犘犇犎1­#。E£,~ÓvGÖLCuB‡
E•WXBÁÂÁbšt¯*”•ŠkBDo

w
5Cughl¯5ë狇犘犇犎1.YÖLB。2010
$
,Suzuki![41]%ðH狇犘犇犎1B1eìz‰5×
†

¦狇犘犇犎1kY•!16™…Þ\‚B134kb
BÝ܂

ÃvG3ÝÜ|┕10e8Cugh
ƒ@BñŒ‡E

2012$,Luo![6]}*hì?Cà®öAf039
!5# = Ê!:h°89Cu:9+‡ÃB˜™;k
=R
³ÎÉ

G>T

KÒ­A
BCugh;i QTLkY,•!6™…Þ\‚I
jÅ1e¾iQTL.Y,Y¦SSR¬±Satt062=
Satt520’“,¼«G狇犘犇犎61。Ð80/Bw‡
WŠX

67*hCughB¾iQTL.YG狇犘
犇犎1,Y¦!16™…Þ\‚[27,38]。£s,~Óø•
狇犘犇犎61.Y­#@AH1eŒ:uFžé
êPgB‡E犌犾狔犿犪06犵00860.1=1e˜ŠK¿
tB‡E犌犾狔犿犪06犵01500.1,ÐPJ:uF=K¿
t•*hCu‡/+@ҖCDBÅ¥

2013$,Gao![42]K狇犘犇犎1.Y;i´ük
Y

¦ÒkY•!16™…Þ\‚B47kbÝ܂。
X“~ӕ狇犘犇犎1­#@AH1e˜ŠbZIPy
2²E7B‹Q7犌犾狔犿犪16犵25600,3‹Q7ËÌ
1e113bpBM‡Œl]ï“,ƒ@˜™PJ,
bZIPy2²E7BòóPÄ,*NX89üýü:
LB—r H
[43]。


#$…狇犘犇犎1=‹eñŒ‡E].Y•*
h+ QTLkYÅ,ÁB5,Ð÷ñŒ‡E(P=
[W✊ÖLCugh

¬‚+ƎU¿Ò~”Ø
=ÅÖLCughBœŠ‡E

ÃxÕCD"M

2014$,Dong![11]•*hGY+UkH‘à
H<+éêkY

‡EâJä=V

ÁÂ+z{

‡
E;
@ANAC‡E­Ê+
B犛犎犃犜15‡E•*hCu‡/+@ÒH×@C
DBÅ¥

ƒ@˜™PJ

¿":*hŎ{*hR
<‡/+

릛çB».Œ
,犛犎犃犜15‡E‚¹
4kb^B1e’L7 S|#$,ю{*hmuP
IQüý+犛犎犃犜15‡EBPÄòó‚R,›¨È
":*hÎìH15[,NXPIQüýü:Lòó
hù<Ã*ëx§•CuݚN

¿ˆifSCu

×1)。£s,~Ó«犛犎犃犜15=狇犘犇犎1#!
‡EAB犌犾狔犿犪16犵25600âƒ#,|âù†y„
B”Ø…RÖüýü:LB H

E£¿üýKL‚
IV犌犾狔犿犪16犵25600B‡Eâ,¦µ¦S“*
hÁÂRÖø”

:1 ՊÀÈÖÊË.3±Š0×ØÙÚ:
Fig.1 Schematicevolutionmodeloftheresistanceofpoddehiscenceindomesticatedsoybeans
ò
:VB,IO;AL,ŸÝ;FC,PIQüý
Note:VB,vascularbundle;AL,abscissionlayer;FC,fibrecapcels
3.2 ÛÈ
•Dh+©@AC‹8Cughƒ@B‡E

1972$,Blixt[44]\üw‡HDhCugh¾Dë1
e—g‡E犇狆狅1RÖ。1987$,Muehlbauer[45]«
5g‡E犘=犞 â\áDhmu‚m@Bï“,
ˆ‚ m @ B ï “ ⠍ i f S C u
。2002 $,
Weeden![28]•Dh2eCà#öf\+,B¥H1
CughB¾iQTL.Y犇狆狅1,ɵk3.Y
Y¦7„fⅢ‚,X“~ӕ7„fⅦ‚kYÅ1
eœiQTL.Y犇狆狅2。£s,~Ó«7„fV
‚RÖmuÝÞPÄB QTL.Y犌狆qš8RÖ
Cu

ÒPÄ*\áu@‚:HLàHæBñH

d̕2007$,Weeden![29]KDhB5eCà#
öf\;i QTLz{,kYÅ4e8Cughƒ
@BQTL.Y,zÆG:犇狆狅1,犇狆狅2,犌狆}†犖狆,
Ð8’0B˜™lmƒ`@

ˆZkYÅB QTL
.Y犖狆Y¦7„fⅢ‚,RÖu@P/u•(¾z

B:u

3.3 ÜÝÞßÈ
ô©Fqhq”•–UCBCuA_
。1967
139
! " # $ !23"
$
,Gladstones![46]\üw‡H8ô©FqhCu
ghƒ@B2e5g®¯"n‡E:犔犲狀狋狌狊=犜犪狉
犱狌狊。犔犲狀狋狌狊‡ERÖmu‚mLNLB¿n,m
uá$#

â–ãu@E“Kˆ½êBua"KB
çë

ÒPÄEmu‚¤ÞFn
ˆ犜犪狉
犱狌狊‡ERÖu@‚:HLàHBk¿,Ñuaz
Ÿ“EÅBÚçï5

ÒPÄEFŽEF

ì¨=_
¨

=犔犲狀狋狌狊‡EB!Y€Â67。£s,3˜™
«

VW5犔犲狀狋狌狊ø5犜犪狉犱狌狊[OâÊÚCu
5

øÉ8X“”•Ð2e‡E“9âPAGS
|åCug

2005$,Boersma![30]} ô © F q h G Y
‘83A:476’=":Q‘P27255’NöJÕ89eF8?
Cà®öAf\GÅ×f,
*¥—kœÇî®à
u ¨ ‹  g
(microsateliteanchored fragment
lengthpolymorphism,MFLP)•–,K>T=Cà
®öABCugh;iDNA«U׆z{Ã%ð
7„×

d̕2007$,Boersma![47]*¥ MFLP•
–

µkH犔犲狀狋狌狊‡EB2eDNA¬±:LeM1=
LeM2,ÃÞ.Ð2e¬±8犔犲狀狋狌狊‡EBÁž
ŸzÆ2.6=1.3cM。Ҙ™lmøPJÐ2e
¬±8ÉñNöGABPQ˜¹5G95%,w8"
:YBPQ˜¹5O36%。2009$,Boersma
!
[48]
ÝÑ¥ƒXBNӃ™JÕ犜犪狉犱狌狊‡EB3
eDNA¬±:TaM1,TaM2= TaM3,ÃÞ.Ð3
e¬±8犜犪狉犱狌狊‡EBÁžŸzÆG2.1,3.9
=3.9cM。X&,jÕÐ3e¬±8ÉñNöGA
BPQ˜¹5G70%,w8":YBPQ˜¹5O
24%~39%。2010$,Li![49]B¥ã<ÌB
MFLP•–,CZU@H犜犪狉犱狌狊‡EB1eƒ—
gDNA¬±:TaLi,ÃÞ.3¬±8犜犪狉犱狌狊‡E
BÁžŸG1.4cM。80»U@Bz7¬±ƒ
È

3¬±825„š)GY=120„Ÿ5YùPQ
˜¹5–ìÄ94%,E£|./¥¦Fkô©Fq
hYf

3.4 àáÈÉ*+
;E¼Cugh vG51eìÁÂgRS,
Ãë‹e‡ERÖ
[50]。
wThBCugh¾Dë
œŠ‡EÖL

wX“EŋeÒ~‡ERÖ
[51]。
ƒ@˜™PJ

h°89Cugh[5ë1~2e—
g‡E]QTL.YRÖ[42]。
AbdElMoneim![34]•ŒTcdDhåCu
GY˜™+

°?ÅåCu":Y

GùšD

À$

ä

Úú!

8DCuŽ{Y

Gùšì

&$

ӄ

ìú!

BNöF2?+Cugh=åCughB
zŸÈÊG3∶1,Ð8›Gk(1ÁÂ+BzŸk
«S|`@

~Ó½£úecdDhBåCugh
5ë1e5g‡E狀狊ÖL,ˆCugh5ë1e—
g!Y‡E犖犛ÖL。Aliboh![52]LJ}ƒhG
‘BNö=V+

P=HƒhBCugh=åC
ughqë1K!Y‡EÖL。Nelson![31]Kô
©Fqh=;E¼B‡EàºÄ;iHX†gz{
ÌKÈ@A

t¯2e9YB‡Eà•l%‚5ì
¨z
w5ÖLCuB‡E犔犲狀狋狌狊–kY•¤
zûœBXmÝÜ+

•Dh=wTh+

ÖLC
uB‡EX&kYÅHXmÝÜ
[2829]。
ЊAÄB
˜™PJ

h°89“|┕͋ƒ„BCu‡
E]ÁÂR֛L

£s

vZ˜™PJ
[53],
•(犕犲犱犻犮犪犵狅)
+

åCumºÓRB¦€5ë¦犛犎犘 X†
‡E+1e˜Š7B~¿NXB,w5Ҍ\Bz
7Á›Lø¾|ì

é‚]s

h°89+©*ëCuY:=‡
E Fk…

Ð÷Y:=‡E•h°,NZGY
ŒT=z7žµTY+Œ.ÝBž¥0b

¯
ˆ

³0B˜™Ÿ^¦KCu‡EB€ðà


z‡EBâ=Å¥›XŸ¾JH

ÑÕ¿ÁÂR
Öø”㨿#Cu›LBI…EÅHL,

4 ²Ç
Cu5h°,NŽ{=:ú‡/+Xúëƒ@
gvìB(Igh’Š

#$…+ÆÓC:@ò8
9Cu:9+‡ÃB˜™

ˆ•CuB€+=:
X+˜™+

³0ÎHC‹[wøVÓ¿#B
°+Ē

C*h+

Cu”•B¾Q=tRÖ2
y”Ø’“B.AW½H¿

;E¼+

+m@h
ù
"DhÊ8
9+

˜™«C‹R/¨8muPIæBH¨ƒ@gÃW*
[15]。
ˆ•CuBz7:9+˜™+

*¥h°89
©Fk͋8Cughƒ@BY:=‡E

C*
h+ B 狇犘犇犎1,狇犘犇犎61,犛犎犃犜15 = 犌犾狔
犿犪16犵25600,w5狇犘犇犎1=狇犘犇犎61B‡E
âŸFFk

Dh+

©O H犇狆狅1,犇狆狅2,犌狆}
†犖狆 ƒ4e|â8Cughƒ@B‡E,w5F
239
!5# = Ê!:h°89Cu:9+‡ÃB˜™;k
*¥Ÿ‹ŒghDoBGY;iVP

hs

ô
©Fqh+q©Fk犔犲狀狋狌狊=犜犪狉犱狌狊ƒ2e8
Cughƒ@B‡E

wÐ2e‡EBJÅižø
¡¦;ŠŒ˜™



VWh°89CuRgB€+ø5Á
Â+˜™[ƒK—Ì

Œ\PAGWXh°89C
uRgB˜™;kWL¼

Cuƒ@‡E@Bp
ë=ùëøtWâèÍÅ9z7TYBF×

Cu
ƒ@l%8‡E’“BÁÂRÖø”Ÿ¾S(ð„
!

DK‚sĒ

h°89Cu•¾…aöFD
Uk}š3eN/B˜™:


KŒCuRgBh°89‘;i.
/R˜

C‹ãöBh°89¢1QBåCul
%=‡E

ÒÓ5£pBÁ´†

ð¤KÐ÷h°
89;i¥¦=˜™

§ŒåCuâ皛Bh°
89

€8Cughƒ@Bl%=‡E

ÐWø
â$%åCuTY‘BYù´†

ˆ‰âå=˜
™h°89CuBz7ÁÂ+›LB‘



€h°89‘BåCu‡E

LJ
%ðÁÂ7„׆
,QTLkY,Cuƒ@‡EBO
 

ìÇëjº=2‡E!ßà

Kh°89


5h°éÂ89BCugh;i./BÁÂz{

¿ˆJÕ*ëBCuƒ@‡E

£s

pF89’
“”•Bb‹ƒ„g

|ÎìCuƒ@‡EB@
i5

•h°89¨×ô7©89+

͋89B
mu€+l%mnƒ„

E£I…<¾Ð÷89
BCu›L

¦*¦Ò~89+ƒ@Cu‡EB
kY



KåCu‡E;iâVP8z{

d
–z7:9+=‡Eà+B@k

žKh°89+
©ìCuƒ@‡EUk‡EPÄz{

‡E@.g
z{

‡EâJäVP!ŠAʙ

¿‡EàK
LOóCuRgB©üz7ÁÂ+›L

¿ˆAÀ

|/

ßlBS“CuЊCDBÁÂRÖø”

u>?
[1] ª¬!,א«.*h¬ug˜™;k[J].*h°+,2012,30
(6):10301034
[2] YH,;­,¦­:,!..Î"DhCu67EF8Cu+
Q˜™
[J].N)+w,2009,18(3):251254
[3] ;­,¶ÔÆ,Ö®d,!.89‰Š)—z7:9+˜™;k
[J].Å9+w,2009,35(3):381387
[4] =¯©,°3ø.89—±gh˜™;k[J].h°‡Eà+8
ÁÂ+

ø”¡
,2011,2(1):113
[5] CavinessCE.Heritabilityofpoddehiscenceanditsassociation
withsomeagronomiccharactersinsoybeans[J].CropScience,
1969,9(2):207209
[6] LuoR,GongP,ZhaoD,犲狋犪犾.QTLmappingandanalysisof
traitsrelatedtopoddehiscenceinsoybean[J].LegumeGe
nomicsandGenetics,2012,3(3):1420
[7] MosM.Anatomicstructureofbirdsfoottrefoil(犔狅狋狌狊犮狅狉狀犻犮
狌犾犪狋狌狊)pod[J].ZeszytyProblermowePostepov,1987,240:
200213
[8] TsuchiyaT.Physiologicalandgeneticanalysisofpodshatter
inginsoybean[J].JapanAgriculturalResearchQuarterly,
1987,21(3):166175
[9] SuzukiM,FujinoK,FunatsukiH.A majorsoybeanQTL,
狇犘犇犎1,controlspoddehiscencewithoutmarkedmorphologi
calchange[J].PlantProductionScience,2009,12(2):217223
[10]SummersJE,BruceDM,VancanneytG,犲狋犪犾.Podshatterre
sistanceintheresynthesized犅狉犪狊狊犻犮犪狀犪狆狌狊lineDK142[J].
TheJournalofAgriculturalScience,2003,140(1):4352
[11]DongY,YangX,LiuJ,犲狋犪犾.Podshatteringresistanceassoci
atedwithdomesticationismediatedbyaNACgeneinsoybean
[J].NatureCommunications,2014,5:3352
[12]TiwariSP,BhatiaVS.Charactersofpodanatomyassociated
withresistancetopodshatteringinsoybean[J].Annalsof
Botany,1995,76(5):483485
[13]WelshMadduxM M,SpaethSC,MuehlbauerFJ.Morpholo
gyoftheslashedpodtraitinlentil[J].EconomicBotany,
1994,48(1):2934
[14]PateJS.Legumefruitsthestructure:functionequation[C]//
StirtonCH,ZarucchiJLed(s).Advancesinlegumebiology.
SaintLouis:MissouribotanicGardensPress,1989:399415
[15]ButlerA.InvestigationsofpodcharactersintheVivieae[J].
VegetationHistoryandArchaeobotany,2002,11(1/2):127
132
[16]YangJB,SomersDA,WrightRL,犲狋犪犾.Seedpoddehiscence
inbirdsfoottrefoil,犔狅狋狌狊犮狅狀犻犿犫狉犻犮犲狀狊犻狊,andtheirinterspecific
somatichybrid[J].CanadianJournalofPlantScience,1990,70
(1):279284
[17]RepkovaJ,HofbauerJ.Seedpodshatteringinthegenus犔狅狋狌狊
anditsovercoming[J].CzechJournalofGeneticsandPlant
Breeding,2009,45(2):3944
[18]ChildRD,summersJE,BabijJ,犲狋犪犾.Increasedresistanceto
podshatterisassociatedwithchangesinthevascularstructure
inpodsofaresynthesized犅狉犪狊狊犻犮犪狀犪狆狌狊line[J].Journalof
ExperimentalBotany,2003,54(389):19191930
[19]ChristiansenLC,DalDeganF,UlvskovP,犲狋犪犾.Examination
ofthedehiscencezoneinsoybeanpodsandisolationofadehis
cencerelatedendopolygalacturonasegene[J].PlantCeland
Environment,2002,25(4):479490
[20]GrantWF.Seedpodshatteringinthegenus犔狅狋狌狊(Fabace
ae):Asynthesisofdiverseevidence[J].CanadianJournalof
PlantScience,1996,76(3):447456
[21]RajaniS,SundaresanV.The犃狉犪犫犻犱狅狆狊犻狊myc/bHLHgene
犃犔犆犃犜犚犃犣enablescelseparationinfruitdehiscence[J].
339
! " # $ !23"
CurrentBiology,2001,11(24):19141922
[22]c“0,²©~,蘳,!.:uãm@T=UCBR֛
L†•tATY+Bž¥
[J].üý:9+NZ,2008,29(6):
864868
[23]AgrawalAP,BasarkarPW,SalimathPM,犲狋犪犾.Roleofcel
waldegradingenzymesinpodshatteringprocessofsoybean,
犌犾狔犮犻狀犲犿犪狓 (L.)Merril[J].CurrentScience,2002,82(1):
5860
[24]PetersenM,SanderL,ChildR,犲狋犪犾.Isolationandcharacteri
sationofapoddehiscencezonespecificpolygalacturonasefrom
犅狉犪狊狊犻犮犪狀犪狆狌狊[J].PlantMolecularBiology,1996,31(3):517
527
[25]RossIbarraJ,MorrelPL,GautBS.Plantdomestication,au
niqueopportunitytoidentifythegeneticbasisofadaptation
[J].ProceedingsoftheNationalAcademyofSciences,2007,
104(S1):86418648
[26]BaileyMA,MianM,CarterTE,犲狋犪犾.Poddehiscenceofsoy
bean:Identificationofquantitativetraitloci[J].JournalofHe
redity,1997,88(2):152154
[27]FunatsukiH,IshimotoM,TsujiH,犲狋犪犾.Simplesequencere
peatmarkerslinkedtoamajorQTLcontrolingpodshattering
insoybean[J].PlantBreeding,2006,125(2):195197
[28]WeedenNF,BraunerS,PrzyborowskiJA.Geneticanalysisof
poddehiscenceinpea(犘犻狊狌犿狊犪狋犻狏狌犿 L.)[J].Celularand
MolecularBiologyLetters,2002,7(2B):657664
[29]WeedenNF.Geneticchangesaccompanyingthedomestication
of犘犻狊狌犿狊犪狋犻狏狌犿:Isthereacommongeneticbasistothe‘do
mesticationsyndrome’forlegumes?[J].AnnalsofBotany,
2007,100(5):10171025
[30]BoersmaJG,PalottaM,LiC,犲狋犪犾.Constructionofagenetic
linkagemapusingMFLPandidentificationofmolecularmark
erslinkedtodomesticationgenesinnarrowleafedlupin(犔狌
狆犻狀狌狊犪狀犵狌狊狋犻犳狅犾犻狌狊L.)[J].CelularandMolecularBiology
Letters,2005,10(2):331344
[31]NelsonM N,PhanH T,ElwoodSR,犲狋犪犾.Thefirstgene
basedmapof犔狌狆犻狀狌狊犪狀犵狌狊狋犻犳狅犾犻狌狊L.Locationofdomestica
tiongenesandconservedsyntenywith犕犲犱犻犮犪犵狅狋狉狌狀犮犪狋狌犾犪
[J].TheoreticalandAppliedGenetics,2006,113(2):225238
[32]KoinangeEM,SinghSP,GeptsP.Geneticcontrolofthedo
mesticationsyndromeincommonbean[J].CropScience,
1996,36(4):10371045
[33]MohammedMS,RussomZ,AbdulSD.Inheritanceofhairi
nessandpodshattering,heritabilityandcorrelationstudiesin
crossesbetweencultivatedcowpea(犞犻犵狀犪狌狀犵狌犻犮狌犾犪狋犪 (L.)
Walp.)anditswild(var.pubescens)relative[J].Euphytica,
2010,171(3):397407
[34]AbdElMoneim A M.Selectionfornonshatteringcommon
vetch,犞犻犮犻犪狊犪狋犻狏犪L.[J].PlantBreeding,1993,110(2):168
171
[35]LadizinskyG.Seeddispersalinrelationtothedomesticationof
MiddleEastlegumes[J].EconomicBotany,1979,33(3):284
289
[36]IsemuraT,KagaA,KonishiS,犲狋犪犾.Genomedissectionof
traitsrelatedtodomesticationinazukibean(犞犻犵狀犪犪狀犵狌犾犪狉犻狊)
andcomparisonwithotherwarmseasonlegumes[J].Annals
ofBotany,2007,100(5):10531071
[37]LiuB,FujitaT,YanZ,犲狋犪犾.QTLmappingofdomestication
relatedtraitsinsoybean(犌犾狔犮犻狀犲犿犪狓)[J].AnnalsofBota
ny,2007,100(5):10271038
[38]KangS,KwakM,KimH,犲狋犪犾.PopulationspecificQTLsand
theirdifferentepistaticinteractionsforpoddehiscenceinsoy
bean[犌犾狔犮犻狀犲犿犪狓 (L.)Merr.][J].Euphytica,2009,166
(1):1524
[39]FunatsukiH,HajikaM,HagiharaS,犲狋犪犾.Confirmationofthe
locationandtheeffectsofamajorQTLcontrolingpoddehis
cence,狇犘犇犎1,insoybean[J].BreedingScience,2008,58
(1):6369
[40]YamadaT,FunatsukiH,HagiharaS,犲狋犪犾.A majorQTL,
狇犘犇犎1,iscommonlyinvolvedinshatteringresistanceofsoy
beancultivars[J].BreedingScience,2009,59(4):435440
[41]SuzukiM,FujinoK,NakamotoY,犲狋犪犾.Finemappingandde
velopmentofDNA markersforthe狇犘犇犎1locusassociated
withpoddehiscenceinsoybean[J].MolecularBreeding,2010,
25(3):407418
[42]GaoM,ZhuH.Finemappingofamajorquantitativetraitlo
custhatregulatespodshatteringinsoybean[J].Molecular
Breeding,2013,32(2):485491
[43]EhltingJU R,MattheusN,AeschlimanDS,犲狋犪犾.Global
transcriptprofilingofprimarystemsfrom犃狉犪犫犻犱狅狆狊犻狊狋犺犪犾犻
犪狀犪identifiescandidategenesformissinglinksinligninbio
synthesisandtranscriptionalregulatorsoffiberdifferentiation
[J].ThePlantJournal,2005,42(5):618640
[44]BlixtS.Mutationgeneticsin犘犻狊狌犿[J].Agriculturaland
HorticulturalGenetics,1972,30:1293
[45]MuehlbauerFJ.Registrationof‘Alaska81’and‘Umatila’
drypea[J].CropScience,1987,27(5):10891090
[46]GladstonesJS.Selectionforeconomiccharactersin犔狌狆犻狀狌狊
犪狀犵狌狊狋犻犳狅犾犻狌狊and犔.犱犻犵犻狋犪狋狌狊[J].AnimalProductionSci
ence,1967,7(27):360366
[47]BoersmaJG,BuirchelBJ,SivasithamparamK,犲狋犪犾.Devel
opmentoftwosequencespecificPCRmarkerslinkedtothe犾犲
genethatreducespodshatteringinnarrowleafedlupin(犔狌狆犻
狀狌狊犪狀犵狌狊狋犻犳狅犾犻狌狊L.)[J].GeneticsandMolecularBiology,
2007,30(3):623629
[48]BoersmaJG,NelsonMN,SivasithamparamK,犲狋犪犾.Develop
mentofsequencespecificPCRmarkerslinkedtotheTardus
genethatreducespodshatteringinnarrowleafedlupin(犔狌狆犻
狀狌狊犪狀犵狌狊狋犻犳狅犾犻狌狊L.)[J].MolecularBreeding,2009,23(2):
259267
[49]LiX,RenshawD,YangH,犲狋犪犾.Developmentofacodomi
nantDNAmarkertightlylinkedtogene狋犪狉犱狌狊conferringre
ducedpodshatteringinnarrowleafedlupin(犔狌狆犻狀狌狊犪狀犵狌狊狋犻
犳狅犾犻狌sL.)[J].Euphytica,2010,176(1):4958
[50]PhilipsRL,KeimWF.Seedpoddehiscencein犔狅狋狌狊andin
439
!5# = Ê!:h°89Cu:9+‡ÃB˜™;k
terspecifichybridizationinvolving犔.犮狅狉狀犻犮狌犾犪狋狌狊L.[J].Crop
Science,1968,8(1):1821
[51]LadizinskyG.Thegeneticsofseveralmorphologicaltraitsin
thelentil[J].JournalofHeredity,1979,70(2):135137
[52]AlibohVO,KehindeOB,FawoleI.Inheritanceofleafmark,
poddehiscenceanddrypodcolourincrossesbetweenwildand
cultivatedcowpeas[J].AfricanCropScienceJournal,1996,4
(3):283288
[53]FourquinCE,DelCC,VictoriaFC,犲狋犪犾.Achangein
犛犎犃犜犜犈犚犘犚犗犗犉proteinliesattheoriginofafruitmor
phologicalnoveltyandanewstrategyforseeddispersalin
犕犲犱犻犮犪犵狅genus[J].PlantPhysiology,2013,162(2):907917

%&( , -
欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍欍

¦§¨©2016ª《9«âãÉ­》9、ä>å
  《+¬()°+》+、H˜¡5ë()¤¾、+¬()°+08+¬(+*ƒX¾Bé@g+–#


¾D´(,)‡Ã°+=ž¥‡Ã°+˜™W˜

és

”w!

æÅ9ÁÂTY
·
Yù´†
·
z
7ÁÂ+

µÅŽ{
·
:X:<
·
()ȶ•–

89û

ËÌ$‘
·
àK¤·
·
():FŽ
;¸
I

þ
¤
·
ûe
·
 .

O,
·
¹-
·
´†`ü!‘³

•ÆK_G¬‚s()°˜0

]
)、
*Ä0—B°˜

–
+8X»È


+¬()°+

+˜¡Gë%

67E7
、`
 ½Rü78‹$6|¬()°•#v0Ä]0ÄY
ü

G4¥*+×(º1992-2011$786ü¡ŒBŸ5#,Y6《+˜Ÿ5#D³`»》"()é@
yŸ5#P"B\Y。1999-2008、2013-2014$J"¬­®¯°+‡±C:+–#Ä²‡±"´µ;
2015$J+¬°Â´G°•#./²³´µ。1999$J"\¼¬­#",2003、2005$J"!Û、Ù¼
¬­#Î«";2002-2014$ËÌ12ü +¬°+•–ȶ˜™]½¾";Y+¬)+–#"]
™
;2009$J+¬#Â*/+¬¡°+˜™0"Z+¬60$67çB#"]™;2010、2013$ƒJ
"!Û、Ù¼+¬¡¿#Î«",2013$JZÀ¡.³`°";›°•#"]™;2012、2013、
2014$J
+¬()°+

+˜¡*16U,”%1、16+¡,¬‚sªU@i。”#208¬,kô49.50®,|$
kô1188.00®。¬‚ÀŠ78¡9™:CN11-1328/S,¬<¬»78¡9™:ISSN0578-1752,¯ @

:2-138,¬s?™:BM43。《+¬()°+》H˜¡(AgriculturalSciencesinChina,ASA),2002$›



。2012$Ÿ«G《()°++w》(JournalofIntegrativeAgriculture,JIA)。2006$1%ê8¬<
r«¡§ÁElsevier@Å,|˜p½•ScienceDirectLÂ/L_…@i。2009$ SCIв,2014$
JIA67E7G0.833。
JIA*16U,”%20+¡,¬‚sªU@i。”#180¬,¬‚±ô80.00®,|$960.00®。¬
‚ÀŠ78¡9™
:CN10-1039/S,¬<¬»78¡9™:ISSN2095-3119,¯ @?™:2-851,¬s

:1591M。

+¬()°+

+

H˜¡–|LJ|¬1ñ¯°±™

q|L˜[¤ÓA±Ã

¯˜
:100081;ñ³:4¥+@Ä*¸12™《+¬()°+》˜[¤
³¹
:010-82109808,82106281,82105098;Â::010-82106247
ø³
:www.ChinaAgriSci.com;E-mail:zgnykx@caas.cn
.A»

£Fm
539