全 文 ：　遗 传 学 报　Ac ta Gene tica S in ica, Sep tember 2005, 32 (9):969 ～ 978 ISSN 0379 -4172
收稿日期:2004 - 12 -27;修回日期:2005 -04 -12
基金项目:由欧盟项目 (编号:IC18-CT 97 -0172), 浙江省自然科学基金重点项目 (编号:Z303407)和国家自然科学基金(编号:30470985)
资助 [ Supported by the Projec t ofEuropean Commun ity(No. IC18-CT 97-0172) , Natuva l sc ience the Foundat ion Projec t of Zhe-
jiang Prov ince(No. Z303407)and Nat iona l Natural Sc ience Foundation (No. 30470985)]
作者简介:赵坚义(1958 -), 女 , 博士, 研究员 , 研究方向:油菜遗传育种与分子生物学
①　通讯作者。 E-ma il:jy zhao3@yahoo. com;Tel:0571 -86403406
QTL of Three Agronom ically Important Traits and The ir Interactions
w ith Env ironment in a European ×Chinese Rapeseed Population
ZHAO J ian-Y i1, ① , BECKER He iko C. 2 , DING Hou-Dong1 , ZHANG Yao-Feng1 ,
ZHANG Dong-Q ing1 , ECKEWo lfgang2
(1. C rop Research Inst itu te , Zhejiang Academ y o fAgricultural Sc iences, Hangzhou　310021 China;
2. Inst itu te ofAgronom y and P lantB reeding , Georg-Augus t-Univers ity , Goettingen　37075, Germany)
Abstract:A rapeseed popu la tion consisted o f 282 doub led hap lo id (DH) lines de rived from a cross between a
European va lity “ Sa llux” and a C h inese inbred line“Gaoyou” was p lanted in 4 loca tions, 2 in X i’ an and Hang-
zhou, Ch ina, and 2 in Goe ttingen, Germany. F ie ld expermi ents w ere carr ied ou t to ob ta in ag ronom ica lly pheno-
typ ic da ta from above fou r env ironmen ts. A linkage map inc lud ing 125 SSR-marke rs was cons tructed and QTL
ana lyses was pe rfo rmed using m ixed mode l app roach to de tect QTLs show ing add itive(a), ep istasis(aa) as
w e ll as the ir in te ractions w ith environm ents (QE) fo r th ree mi portan t ag ronom ic tra its:p lan t he ight, flow er ing
tmi e and m atu rity. The resu lts dem onstra ted tha t each tra it was con tro lled by severa l QTLsw ith add itive e ffec t
and a number o f QTLsw ith ep ista tic andQE in te raction e ffects. P lan t he igh t was con tro lled bymany QTLs(12
loc iw itha o r comb ined ae , 5 lociw ithae). Add itive e ffects were p redom inan t, to ta lly exp la ined 75% o f the phe-
no typ ic va r ia tion and o ften comb ined w ith d igen ic ep istasis. O f 12 m a in QTLs, 9 showed Gaoyou a lle les decrea-
s ing p lant he igh .t Most o fQTLsw ithQE effects show ed eco log ica lly favou rab le a lle les in d ive rse reg ions. F ive o f
7 ae loc ishow ed Gaoyou a lle les in Hangzhou and a ll theae loc ibu t one had So llux a lle les in two loca tions o f
Germany increas ing p lan t he igh.t The d igen ic ep ista tic ma in e ffec t accounted fo r one th ird o f to ta l add itive m a in
e ffects.
In th is study, w e d iscove red 7 and 8 loc ihav ing s ign ifican t add itive ma in e ffects upon flower ing tmi e and ma-
tur ity, respective ly. O f them, ear ly flowering and m atu rity a lle les w ere respec tive ly 6 and 5 de rived from C h inese
pa ren tGaoyou. A ll these QTLs toge the raccoun ted fo ra round 60% o f the pheno typ ic var ia tion fo r each tra i.t S ig-
n ifican tae inte ractions w ere de tected for flowering tmi e and matur ity and paren ta la lle les showed aml ost even ly
d ispe rsa l a t a llenv ironmen ts. Three o f 8 ma in QTLs fo rma tu r ity were loca ted a t smi ila r o r iden tica lpos itions as
QTLs fo r flow er ing tmi e, w h ich con firmed the close co rre la tion between these two tra its.
Two QTLs fo r p lan t he ight on linkage g roups N14-1 and 19 w ere loca ted a t smi ilar pos itions as QTL fo r
flow er ing tmi e and as a lready known QTLs fo r o il con ten.t Se lec tion fo r reduced p lan t he ight and ea rly flow er ing
m igh t reduce o il con ten .t D igen ic ep is ta tic QTLs both fo r flowering tmi e and m atu rity w ere de tected but much
less mi po rtan t than QTLsw ith add itive e ffects.
Key words:Brassica napus L.;p lan t he igh t; flowering tmi e;matu rity;QTL;QTL and env ironmen t In te rac-
tion
应用中国和欧洲油菜建立的双二倍体群体对 3个
重要农艺性状的 QTL定位以及与环境的互作分析
赵坚义 1, ① , He iko C. Becker2 , 丁厚栋 1 , 张尧锋 1 , 张冬青 1 , Wolfgang Ecke2
(1. 浙江省农业科学院作物与核枝术利用研究所 , 杭州　310021;
2. 德国哥廷根大学作物遗传育种研究所 , 37075)
摘　要:以中国的高油分自交系 “高油”和欧洲高含油量品种 “ So llux”的 F1产生的 282个株系组成的双二倍体
(DH)群体为材料 ,在 125个 SSR标记座位构建的连锁图谱基础上 , 根据在中国和欧洲四个不同环境下的表型鉴
定结果 , 采用混合线性模型基础上的 QTL分析软件 ,对油菜 3个重要农艺性状:株高 , 开花期和成熟期进行了数量
性状基因座位(QTL)的联合定位分析 , 估测了这些 QTL的加性 、上位性以及与环境的互作效应。结果表明各性状
均受多个加性 、加加上位以及与环境互作的 QTL控制。株高受多个 QTL影响(12个位点具有加性或兼有环境互
作效应 , 5个位点具有互作效应), 以加性效应为主 ,加性效应总和可解释定位群体表型变异的 75%左右 , 并多兼有
上位性效应。 12个主效 QTL中 , 9个是 “高油”等位基因相对 “So llux”有降低株高的作用 ,大多数加性 ×环境互作
QTL的有效等位基因具有环境选择特异性。 7个 ae基因座位中 , 5个 “高油”等位基因在杭州种植环境下 , 除一例
外所有在德国环境下的互作基因座中 , “ So llux” 等位基因起着增加株高的作用 ,加加上位性主效总和为加性主效
总和的三分之一。
7个控制花期和 8个控制成熟期的主效 QTL中 ,分别有 6个和 5个是来自 “高油”的等位基因相对 “ So llux”具
有提前开花和成熟的效应 , 这些 QTL的效应总和占到性状表型变异的 60%左右。 5个位于第 2和第 12连锁群中
的 2个大效应 QTL可能和已多次报导的 VFN 1和 VFN 3基因相近或相同。开花期和成熟期两性状均检测到显著
的 ae互作效应 ,双亲等位基因的效应在各环境下呈离散分布。位于 14和 19连锁群上的两个主效株高 QTL同时
也是控制开花期和油分含量的基因位点 ,因而利用这两个位点进行标记辅助筛选时要考虑到对油分含量的影响。
控制成熟期的 8个主效 QTL中有 3个同时也是控制开花期的基因座位 , 证实了开花期和成熟期高度正相关的遗
传基础 , 两个生育性状均表现有较弱的 QTL间加加上位互作 ,但以主效 QTL的作用为主。
关键词:甘兰型油菜;株高;开花期;成熟期;QTL定位;QTL与环境的互作
中图分类号:Q943　　　文献标识码:A　　　文章编号:0379-4172(2005)09-0969-10
　　 P lant he ight is one of the most mi portant
p lant morpho log ica l tra it re la ted to y ie ld potent ia l
inB rassica napus. F lowering tmi e is a major de-
te rm inant o f reg iona l and seasonal adapta t ion of
rapeseed varie ties and c lose ly linked to matu-
rity
[ 1]
. P lant he ight may re la te to lodg ing res is-
tance
[ 2]
and is regarded as unsuitab le formecha-
nized harvestwhen ta lle r than 160 cm. Therefore,
breeders prefer to se lect shorter genotypes. How-
ever, most commerc ia l cu ltivars have shown only
a little reduction in he ight,mostly rang ing from 150
-170 cm , s ince short p lant type norma lly hard to
reach satisfied y ie ld leve l, ind icat ing that the p lant
he ight has a genetic linkage w ith seed y ie ld com-
ponents
[ 1]
. S ign ificant genera l comb in ing ability
(gca)-and spec ific combin ing ab ility (sca)-vari-
ances were found for p lant he ight
[ 3-5] , demonstra-
t ing both additive and dom inant gene effects.
F lowering tmi e is contro lled by a number of
genes in fluenced by day length and verna liza-
t ion
[ 6]
and is often used as a se lect ion index for
maturity ind irec tly. There were many stud ies to an-
alyze the genet ics of flowering tmi e using mo lecu-
la rmarkers inBrassica spec ies. Three QTLswere
detected (VFN1, VFN2, VFN 3) [ 7-8] in Brassica.
napus and two of them (VFN 1 andVFN2) corre-
sponded to two QTLs in B. rapa (VFR 1 and
VFR2) [ 9] . The chromosome reg ions conta in ing
970 遗传学报　A ctaG ene tica S inica　Vo.l 32　No. 9　2005
these two pairs ofQTLs may correspond w ith two
genes (FLC andFR I) that regu la te flower ing tmi e
ofArabidops is tha liana located at the top of chro-
mosome 5 and chromosome 4
[ 8, 10-12 ]
. Butru ille et
a l.
[ 13]
ident ified seven QTLs for flowering tmi e
and two of them may correspond to VFN1 and
VFN 3. Axe lsson eta l.
[ 14 ]
reported that a ll the ma-
jo r flowering QTLs detected in d iffe rent spec ies of
B rass ica could be a resu lt o f dup lica ted copies of
the same ancestra l gene. Camargo and Os-
born
[ 15]
mapped three QTLs for f lowering tmi e in
B. o leracea and detec ted epistasis between two
genom ic reg ions. Mapping stud ies of p lant he ight
and number of prmi ary branches were rare in o il-
seed rape. Butru ille et a l.
[ 13 ]
mapped four QTLs
for p lant he ight and showed ple io tropy of flowering
tmi e at two major loc.i
In the current study, we analyzed the data
from four env ironments, two in Germany and two
in China of a doub led haplo id (DH) popula tion,
which was derived from a cross between a Euno-
pean varie ty (Sollax)and a Chinese inbred lines
(Gaoyou), using the m ixed model approach de-
scribed byW ang et a l.
[ 16]
. The object ives were to
analyse the genet ic basis of p lant he ight, flowering
tmi e and maturity, to d issect the genet ic re la tion-
sh ips among these three ag ronom ica lly mi portant
tra its and to compare w ith prev ious researches.
Informat ion obta ined from th is study may g ive
suggestions for mi provement of agronom ic tra its
by marker-assisted se lec tion to get h igher seed
y ie ld both in China and in Europe.
1　Mater ials andMethods
1. 1　P lantmateria ls and fie ld tr ia ls
Tota l 282 doubled haplo id (DH) lines were
produced from the F1 of a cross between “Sollux”
and “Gaoyou”. So llux was a Germanw inter rape-
seed re leased by ZGW interraps (German Demo-
cratic Republic). Gaoyou is a Chinese inbred line
from varie ty “Gaoyou 605”, developed by Zhe-
jiang Agricu ltura l University. The DH lines, F1 gen-
eration and parents were grown in four environ-
ments in 2000 /2001:two in China (X i’ an and
Hangzhou) and two in Goett ingen, Germany (Re-
inshof andW eende).
A random ized complete b lock des ign w ith two
rep lica t ions was used. The seeds were sown in
double rows for each plo t, w ith rows of 2.5 m
length and d istances of0.33 m between rows and
0.12 m between plantsw ith in rows in Goettingen
and 0.15m in China. The sow ing date was at nor-
mal tmi e in Hangzhou and two locat ions in Ger-
many, while in X i’ an, it was around 10 days
delayed as expected. Three agronom ic tra its-p lant
he ight, f lowering tmi e and maturity were assessed
and recorded. F ive healthy p lants in the m idd le
part o f each plo t were random ly se lec ted and
samp led for measur ing p lant he ight before har-
ves.t F lowering tmi e and maturity were recorded
as the number of days from sow ing until 50%
plants show ing first flowers(TDF) and from so-
w ing unt ilmaturity (siliques be ing ye llow , TDM),
respective ly. The data of p lant he ight was ob-
ta ined from all four locations, while other two tra its
were only investigated at three locations (Weende
was not included).
1. 2　Linkage map and QTL assays
A linkage map was construc ted us ing 125 m i-
crosate llite markers, span ing 1 196 cM of the
rapeseed genome (Kosambi funct ion)w ith an av-
erage in terva l o f9. 6 cM between markers. Deta ils
o f deve loped DH popula tion and SSR map were
described prev iously
[ 17]
.
The M INQUE method
[ 18]
was used to esti-
mate var iance components. The genet ic main
effects(G) and in teract ion effec ts (GE) were
pred icted by the Adjusted Unbiased Pred ict ion
(AUP) method[ 18, 19] . The Jackkn ife method was
app lied for obta in ing estmi ates of pred ictors and
the ir standard errors in a t-test for parameters.
QTLs w ith add it ive and additive × additive
ep istat ic e ffects as we ll as the ir env ironmenta l in-
971ZHAO Jian-Yi et a l. :QTLs o f Th ree Ag ronom ica lly Im po rtan t T raits and Their Interactions…
te ract ions (QE ) were mapped by QTLMapper
vers ion 1. 0
[ 16 ]
. The model, explanat ions of a ll pa-
rameters and deta iled mapp ing procedure were
described prev iously
[ 16, 17]
. QTLsw ith add it ive and
ep ista tic e ffects were filte red us ing a s ign ificance
thresho ld ofP =0. 005. The main effects and the
genotype env ironment interaction effec ts were
tested by a t test w ith the jackkn if ing resampling
procedure. QTLs were presented when genetic
ma in effects (a and aa ) orQE in teract ion effect
(ae and aae)were sign if icantly d iffe rent from ze-
ro (P =0.005).
2　Resu lts
2. 1　Phenotyp ic varia tions
The phenotyp ic frequency d istribut ions of the
282 DH lines for three agronom ic tra its under
three or four env ironments were presented on
F ig. 1. Formost lines, planthe ightwas from 120 to
160 cm in X i’ an, wh ile theywere from 130 to 170
cm in Hangzhou and Goettingen (Re ishof and
W eende) respect ive ly. The distributions of days
from sow ing to flower(DTF) and from sow ing to
maturity(DTM ) were clearly separated in two
groups between locations of China and Germany.
On average, the p lant performance of DH popula-
t ion in Hangzhou showed ta llest p lant he ight, earli-
est flowering tmi e and maturity, while in Reinshof
the flowering tmi e was around two months la ter
and the maturity was two and half months la ter
than in locations of Ch ina (Table 1). The to ta l
grow ing period(DTM) was about ha lf month in
d ifference between parents at a ll locations. W ide
transgressive segregation was observed for p lant
he ight and, to some ex tent, formaturity but not for
f lowering tmi e.
F ig. 1　Frequency d istribu tion o f the So llux /Gaoyou de rived DH lines fo r three ag ronom ic tra its
　　Dom inant effects cou ld be pred icted for a ll
three tra its from the phenotyp ic va lues of F1 and
parents. P lant he ight behaved over dom inant
e ffect, while flowering tmi e and maturity showed
partia l dom inant effects.
The estmi ates of genetic, GE interact ion and
res idua l variances are presented in Table 2. All
th ree agronom ic tra its were main ly contro lled by
genet ic e ffects and the heritab ility caused by addi-
t ive effectwere about50% oreven h igher for each
tra i.t GE interact ion effects were sign if icant and
also s ign ificant residua l e ffects ex isted for a ll tra its.
972 遗传学报　A ctaG ene tica S inica　Vo.l 32　No. 9　2005
Tab le 1　Pheno typ ic va riation o f three ag ronom ic tra its
T raits Loca tions
Parents
Sollux Gaoyou
F1
DH popu lat ion (n=282)
Max M in M ean SD
P lant height(cm) X i’ an 156 100 153 166 78 135 14. 0
Hangzhou 149 147 175 184 106 152 13. 2
Reinshof 161 120 163 180 93 144 14. 4
Weende 155 120 165 177 83 140 15. 2
M ean 155 122 164 177 90 143 14. 2
Days from sow ing to X i’ an 193 176 184 195 175 183 3. 28
f lower(DTF) Hangzhou 189 160 179 190 159 177 6. 60
Reinshof 251 222 245 257 226 245 6. 05
M ean 211 186 203 214 187 202 5. 31
Days from sow ing to X i’ an 246 233 243 249 232 242 3. 58
m aturity (DTM) Hangzhou 239 224 236 243 220 232 4. 00
Reinshof 324 309 317 326 305 316 3. 86
M ean 270 255 265 273 252 263 3. 81
Tab le 2　Estmi a tion o f va riance components for
gene tic e ffec ts and geno type× env ironmen t
in te raction e ffects o f th ree ag ronom ic tra its
Variance com ponent P lant he ight F lowering tmi e Ma turity
VG 186. 0** 19. 4** 10. 3**
VGE 93. 7
* 7. 4** 4. 2**
V
e 86. 6
* 2. 5* 4. 4**
Vp 366. 4
* 29. 4** 18. 9**
Heritability(N)% 50. 8 66. 0 54. 6
Heritability(B)% 76. 3 91. 0 76. 7
　*, **:S ign ificant at the 0. 05 and 0. 01 probab ility leve ls, re-
spect ively;VG , VGE , Ve and Vp are genetic m ain effect, geno-
type × env ironm ent interac tion effec t, res idual and phenotypic
variances, respec t ively.
2. 2　QTL analys is for p lant he ight
The segregation of DH lines for p lant he ight
cou ld be large ly exp la ined by severa l QTLs w ith
additive or ep ista t ic e ffects and the ir complex
in teract ions w ith env ironmen.t In to ta l, 17 QTLs
w ith add it ive (a ) and /or add it ive ×env ironment
in teract ion effec ts (ae ) located on 14 linkage
groups and 7 pairs of loc iw ith ep istat ic (aa) and /
or ep ista tic × env ironment in teract ion effects
(aae)were detec ted (Table 3). O f these QTLs,
12 had sign ificanta effects and alle les from Sollux
at 8 loc i were in the d irec tion of increasing p lant
he ight, wh ile a lle les from Gaoyou on the remain ing
4 loc i were a lso increas ing p lant he igh.t P lant
he ight caused by ind iv idua lmain loc i w ith homo-
zygous genotype could change from 3 to 9 cm in
th is popula tion. A ll 12 QTLs being co llective ly
summed upmay increase (or decrease) 65 cM in
p lant he ight w ith homozygous genotypes and ex-
pla ined 75% of the mean phenotyp ic varia t ion ob-
served in the DH popula t ion as expressed by the
d ifference between lines w ith h ighest and lowest
mean va lues (Table 1). 10 QTLs had sign if icant
ae interaction effects. O f them, five comb ined w ith
a effect and the rest on ly showed ae effects in two
or three env ironments. More ae QTLs were found
in locations of Ch ina than that in Germany. Fur-
thermore, 5 of 7 ae loc i showed Gaoyou alle les in
Hangzhou and all the ae loc i but one had So llux
a lle les in two locat ions of Germany may increase
plant he ight, while in X i’ an, th ree a lle les from Sol-
lux and two a lle les from Gaoyou disp layed positive
effects for p lant he igh.t O f seven pairs of ep ista t ic
loc i for p lant he ight, one showed only aae effec t
and the rema in ing s ix pa irs exh ib ited e itheraa or
aa comb ined aae effects. O f these in teract iona l
QTLs , seven loci inc lud ing two pairs ofQTLs com-
bined a and /or ae effects smi u ltaneously. The
sum of ep ista tic e ffects caused by s ix pa irs of
main loc i accounted for one th ird of tota l add itive
main effects of12 QTLs.
2. 3　QTL analysis for flower ing tmi e and maturity
　　S ince the parents greatly d iffered in flower ing
tmi e and maturity, a strong segregat ion for these
two tra itwere observed in the DH popula tion at a ll
locations (F ig. 1, Tab le 1). Mapping analys is d is-
covered 9 QTLs w ith a and /or ae in teract ion
effects for flowering tmi e (Tab le 4). O f these
QTLs, two showed only ae effects and seven dis-
played a or combined ae effects. The Sollux a l-
973ZHAO Jian-Yi et a l. :QTLs o f Th ree Ag ronom ica lly Im po rtan t T raits and Their Interactions…
le les de layed days to flower on six of seven ma in
QTLs. Collec tive ly the seven ma in QTLs expla ined
about 63% of the phenotyp ic varia t ion in DH pop-
ula t ion (Tab le 1). Add itive × env ironment in ter-
ac tion effects were ident if ied at five genom ic re-
g ions and three of them combined w ith add it ive
main effects. One QTL on N12 combin ing a and
ae effects caused five days earlie r to start flower-
ing in Re inshof when homozygous substitu tion of
Sollux a lle le by a lle les ofGaoyou. A lle les ofae lo-
c iwere d ispersed between parents, show ing QE
in teraction in d iffe rent way at ind ividua lae loc.i
Tab le 3　Estmi a ted add itive (a) and ep istatic(aa) e ffec ts
o fQTL and QTL×env ironmen t interac tions(ae, aae) fo r p lan t he igh t(cm)
N⊕ Marker interval
Pos it ion
(cM) † a
‡ ae in
X i’ an
ae in
Hangzhou
ae in
Reinshof
ae in
W eende
1 HMR292 /HMR327 0. 0 3. 16**
2 HMR364a /HMR300a 6. 0 -2. 09** 1. 18** 1. 24*
3 HMR449a /MR12 4. 0 3. 28** 2. 53**
3 HMR376a /MR197. 1 0. 0 -2. 53** 2. 56**
4 HMR637 /MD2. 2 0. 0 -1. 92** 1. 07* 1. 45**
5 HMR327a /MD21 0. 0 1. 55**
7 MD20 / HMR166 4. 0 1. 67** 1. 70**
9 HMR612b /MR230. 1 2. 0 - 3. 73**
9 HMR359a /HMR370a 0. 0 2. 31** -1. 73**
11-1 MR148 /HMR407a 14. 0 - 1. 42*
12 HMR353b /HMR364b 2. 0 2. 46** -2. 59**
13 MR203. 2 /HMR321 4. 0 1. 68**
14-1 HMR403b /MR229 4. 0 2. 84** -2. 32** 2. 07**
16 HMR348 /MR133 0. 0 4. 64** -1. 59**
16 HMR563 / HMR373 4. 0 - 2. 07**
17 HMR349 /MR127 6. 0 - 2. 02**
19 HMR643a /HMR615a 4. 0 4. 32** 3. 11** -1. 44*
Epistatic e ffec ts
N M arker interval N Marker interva l aa # aae in
X ian
aae in
Hangzhou
aae in
Re inshof
aae in
W eende
1 HMR292 /HMR327c 14-1 HMR 403b /MR229 -1. 64**
2 HMR364a / HMR300a 12 MR132 /HMR399b -1. 77**
3 HMR 449a /MR12 5 MR14 /HMR347 2. 49** -1. 49** 1. 25*
3 HMR364a / HMR300a 9 HMR359a / HMR370a -1. 56** 1. 48**
7 MD20 / HMR166 13 MR64 / HMR376b -1. 33*
9 HMR363a /HMR311b 11-1 HMR314a / HMR335b 2. 78** -1. 98*
12 HMR299c / HMR403a 15 MR97 /MR54 2. 31**
　⊕:L inkage group;*:The s ign if icance level at0. 005 to dec lare the putativeQTL pos it ions and genet ic e ffec ts;**:The s ignif icance lev-
e l at 0. 001 to dec lare the putativeQTL pos it ions and genet ic ef fec ts;†:D istance of the QTL from the f irs tm arker o f the ind icated interval,
g iven in centMi organs (cM) accord ing to the Kosamb im app ing funct ion;‡:TheQTL e ffec t is the pheno typic effect due to the subst itut ion
o f a Gaoyou alle le by an alle le ofSollux;#:A pos it ive s ign of the ep istat ic effect indicates that parental allele com binat ions and a negative
s ign that recomb inant allele comb inat ions increase pheno typic va lues.
　　E ight QTLsw itha or combin ing ae effec ts for
p lant maturity were reso lved (Tab le 4). Except
one show ing only add it ive ma in effect, the rema i-
ning 7 QTLs jo ined w itha andae effects. A ltogeth-
er e ightmain QTLs led to 12 days earlie r o fmatu-
rity upon homozygous genotypes, and collective ly
accounted for about 60% of to ta l varia t ion over
three env ironments. So llux a lle les de layed maturi-
ty on five gene loc i, but caused early maturity on
another three loc.i Furthermore, s ign if icantae in-
te raction w ith re la t ive larger effec ts on severa l
gene loc i were identified and aml ost even ly d is-
pers ion of a lle les between parents was observed
at a ll th ree locations. These m ight be the reasons
for transgrasive segregation of th is tra it observed
in DH popula tion. Three of 8 main QTLs formatu-
rity were the ident ica lQTLs for flowering tmi e.
974 遗传学报　A ctaG ene tica S inica　Vo.l 32　No. 9　2005
Tab le 4　Estmi a ted add itive (a) and add itive×env ironmen t in teraction
(ae) e ffects o f QTL for flowering tmi e (DTF) and matu rity (DTM)
N
Marker
interva l
Pos ition
(cM)
DTF
a ae
†
1 ae2 ae3
DTM
a ae1 ae2 ae3
1 HMR295c /HMR293 2. 0 -1. 22** - 0. 57* - 1. 05** 1. 16**
2 HMR066 /HMR087c 26. 0 1. 54* 1. 68** -1. 17**
3 HMR376a /MR197. 1 0. 0 -0. 79** 0. 85** 0. 77** 0. 58** -0. 89**
3 MR12 /MR3 16. 0 0. 74* - 0. 69** 0. 52*
7 MR133. 2 /MD20 8. 0 0. 85** 0. 56** 1. 19** -1. 96**
11-2 HMR295a /HMR295b 8. 0 1. 15**
12 HMR353b /HMR364b 2. 0 1. 32** - 1. 12** -0. 79** 1. 41** 1. 10**
14-1 HMR403b /MR229 4. 0 0. 71**
15 MR97 /MR54 0. 0 - 0. 56** - 0. 55** - 1. 63** 2. 18**
16 HMR348 /MR133 0. 0 1. 18** 0. 70** 0. 97**
18 HMR335a /HMR588 2. 0 -0. 57** 0. 68**
19 HMR643a /HMR615a 4. 0 1. 07* 0. 87** -0. 75**
19 HMR281 /HMR327b 24. 0 - 0. 67** 0. 93** - 1. 11**
　Note:ae†1 , ae†2 andae†3 represent the QTL w ith add it ive by env ironm enta l interact ion ef fec t in X i’ an, Hangzhou and Re insho f respect ively.
The meaning of other abbrev iat ions and symbols are the same as in Table 3.
　　Ep istas is ofaa effec ts cou ld c learly account
for small part o f genetic bas is of flowering tmi e as
well as for p lantmatur ity (Table 5). Three d igen ic
pa irs of loc iwere detected both for flowering tmi e
and for maturity. The s ign ificant leve l and magn i-
tude of e ffec ts was re la tive ly low. F ive and two
pa irs of loc i w ith additive by environment in terac-
tion effects were identified for flowering tmi e and
maturity respective ly. Most o f them presented as
d ifferentQTL comb inations asmain ep ista tic loc.i
Largeraae effect for flower ing tmi e was identified
at one pair o f loc ion N3 /N14-2, caus ing 10 and 4-
5 days earlie r to flower when two alle les from d if-
fe rent parents homozygous ly recombined in Rein-
shof and the parenta l a lle le comb inations presen-
ted in locations ofCh ina, respective ly. Anotheraae
pa ir o f loc i on N1 /N1 for p lantmatur ity,m ight lead
to 5 and 2 -3 days earlie r to maturity in Germany
and Ch ina locations, respect ive ly, when parenta l
a lle les combinations presented in Germany and
recomb inant a lle le combinations in China. No
ident ica laa and aae pa irs of loc i were detected
between flowering tmi e and maturity.
2. 4　Re lat ionsh ip between three agr icu ltura l tra its
　　Genetic and phenotyp ic corre lat ions among
three agronom ic tra its listed on Table 6 and two
types of corre lat ion were well cons istentw ith each
other. P lant he ight showed sign if icant re la t ion to
f lowering tmi e and maturity, and flowering tmi e a l-
so high ly corre la ted w ith maturity. Mapp ing resu lts
well exp la ined the ir co-re lat ionsh ips. W e found
that in three genom ic reg ions on N14-1, 16 and 19
Tab le 5　Estmi a ted ep ista tic(aa) and ep istas is × environm ent in te raction (aae)
e ffec ts o fQTL fo r flowering tmi e (DTF) and matur ity(DTM)
T rait N Marker interva l N M arker interval aa aae†1 aae†2 aae†3
DTF 1 HMR274b /HMR430 12 HMR364b /GMR132 0. 73** -1. 19**
1 HMR407b /HMR292 13 MR64 /HMR376b -0. 93** -0. 63**
3 HMR376 /MR197. 1 14-2 HMR399a /MR155 - 2. 59** -2. 16** 5. 07**
4 HMR637 /MD2. 2 12 HMR353b /HMR364b -0. 84*
8 HMR577 / HMR613 9 HMR612b /MR230. 1 0. 54** -0. 55*
9 HMR612b /MR230. 1 12 HMR353b /HMR364b -0. 54*
14-1 HMR403b /MR229 17 HMR318 / HMR349 0. 49** -0. 71**
DTM 1 HMR292 /HMR327c 1 HMR293 /HMR274b 1. 22** 1. 47** -2. 77**
7 HMR300c /MR133. 2 13 HMR067 /MR64 -0. 30*
7 MD20 /MR166 13 HMR337 /MR61 0. 40* - 1. 02** -0. 41* 0. 92**
11-1 HMR335b /HMR374 12 HMR299c /HMR403a -0. 46*
　Note:aae†1 , aae†2 andaae†3 represent the QTL w ith ep is tasis by env ironmental interact ion effec t in X ian, Hangzhou and Re inshof, respec-
tively. The meaning of other abbrev iations and sym bols a re the same as in Tab le 3.
975ZHAO Jian-Yi et a l. :QTLs o f Th ree Ag ronom ica lly Im po rtan t T raits and Their Interactions…
Tab le 6　Gene tic(above d iagona l) and
pheno typ ic(be low d iagona l) co rre lations
am ong th ree ag ronom ic tra its
T ra its PH† DTF DTM
PH 1 0. 577** 0. 457**
DTF 0. 554** 1 0. 910**
DTM 0. 423** 0. 956** 1
　†:Abbrev ia tions of tra its are the same as in Tab le 1;*, **:
S ign ificant at the 0. 05, and 0. 01 probab ility leve ls respec tively.
for p lant he ight, QTLswere a lso detected for f low-
ering tmi e and the add it ive effects of these QTLs
were re lat ive ly larger(Tables 3 and 4 );four com-
mon or ad jacent genom ic reg ions w ith add it ive
and one pair o f common loc iw ith ep ista tic e ffects
were found both for p lant he ight and tota l grow th
period (Tab les 3, 4 and 5 );sign if icant corre la tion
between flowering tmi e and maturitym ight be due
to the gene overlapp ing at three additive main loci
on N1, 12 and 16.
3　D iscuss ion
In the present study, we used a QTLmapping
program based on m ixed model to detec t QTLs
w ith addit ive and ep ista tic e ffects, as well as the ir
QE interaction effects for three ag ronom ica lly mi -
portant quant ita tive tra its. The resu lts demonstra-
ted that each tra it was contro lled by severa l add i-
tive main QTLs and a number of QTLs w ith ep i-
stat ic andQE interaction effects. P lant he ight was
contro lled by many QTLs and additive effects
were the predom inant, which are in agreement
w ith prev ious c lass ic genet ic researches
[ 3-5]
. The
varia tion of p lant he ight caused by one maiorQTL
on N16 (HMR348) or N19 (HMR615a) could be
around 9 cm when homozygous substitut ion be-
tween Sollux and Gaoyou alle les ( when SS→
GG , decrease 9 cm and when GG→SS, inc rease
9cm in p lant he ight per locus). The effect o f two
QTLswas genera lly consistent w ith that identified
by Butru ille et a l.
[ 13] , show ing the most s ign ificant
e ffect o f the two QTLs, but they were presented in
d iffe rent genom ic reg ions. In another two linkage
groups N3 and N13, both Butru ille et a l. and our
research found QTLs contro lling p lant he igh.t The
rest o f e ight p lant he ightQTLs in th is study repre-
sented new loci that have not been mapped and
characterized before. QTL informat ion of p lant
he ight is in teresting for getting an optmi a l p lant
structure of rapeseed varie ty by molecu lar des ig-
ning and marker assistant se lec tion. The detected
d igen ic ep istasis loc i and the ir complex comb ina-
t ion w itha orae effects are not reported before in
o ilseed rape to our know ledge. Most o f QTLs
showed Sollux a lle lesm ight increase plant he ight,
while four loc i w ith Gaoyou alle les m ight a lso in-
crease p lant he igh.t The dispersion of a lle le effec t
d irect ions between parents exp la ined the large
transgressive segregation observed in the DH
popula tion. QTLs w ith ae in teract ion effec ts were
sign if icant a t each environmen.t It shou ld be noted
that most o fae QTLs in Hangzhou and a ll but one
in two locations in Goett ingen showed Gaoyou
and Soullx a lle les having the effect o f increas ing
p lant he igh.t The sam ll phenomenon was ob-
served for o il content
[ 17]
as we ll, ind icating genes
for p lant he ight are, to some extent, under the reg-
ula tion of eco log ica l env ironments. The reason
m ight be that the parents se lected for adaptat ion
to the very d iffe rent grow ing conditions. A lle le d is-
tribu tion in expermi ent o f X ian showed that two
Sollux and three Gaoyou alle les increased p lant
he igh.t Th is cou ld be exp la ined that the environ-
ment conditions in X ian, main ly temperature, were
somewhat between Hangzhou and Goettingen
(data not shown).
Butru ille et a l.
[ 13]
detec ted seven loc i for flow-
ering tmi e us ing an inbred backcross popula tion.
The marker loc i, wg6b10a (N2 ) and ec3f12b
(N12), showed the most s ign if icant e ffect, which
were ident if ied c lose ly linked to VFN1 and VFN3,
the major flowering tmi e QTLs reported prev ious-
ly
[ 7 -9]
. In th is study, we discovered seven loc i
hav ing s ign ificant e ffects upon flower ing tmi e and
Gaoyou alle les showed early f lowering on a ll loc i
except one on linkage group 1, resu lt ing in no
c lear transgressive outlie rs observed w ith in popu-
976 遗传学报　A ctaG ene tica S inica　Vo.l 32　No. 9　2005
la tion. Furthermore, the two loc i hav ing the largest
add itive effec t were a lso located on N2 and N12
near marker HMR087c and HMR364b, probably
they are the identica l QTLs reported before
[ 7 -9 , 13]
. A QTL on N12 near marker HMR 364b
was also detected to be combined w ith in three
pa irs of s ign ificant d igen ic ep istas is (aa oraae),
which was reported by Butru ille et a l.
[ 13]
as well
formarkerwg 6b10a. Three QTLs w ith large add i-
tive effec ts for p lant he ight on N14-1, 16 and 19
m ight be p le iotropy for flowering tmi e as reported
prev ious ly
[ 13]
. Th is prov ided the genet ic basis for
s ign ificant genet ic corre la tion between these two
tra its (Table 6). However, QTLs on N14-1 and 19
also pos itive ly were linked w ith QTL for o il con-
tent
[ 17 ]
. Se lection for reduced plant he ight and
early flowering tmi e m ight reduce oil content,
which is mi portant in format ion for breed ing pro-
gram.
P lantmaturity is large ly determ ined by f lower-
ing tmi e. The resu lts from th is study revealed
three of e ight QTLs for maturity overlapp ing w ith
flower ing tmi e. The genetic e ffects of these loci
were re la tive ly larger, which exp la in the h igh ly sig-
nificant corre la tion between these two tra its (Ta-
ble 6). There were three loci w ith which Gaoyou
alle les de layed the maturity, which could be the
exp lanat ion for transgressive segregation ofmatu-
rity observed in the DH popula tion. D igen ic ep i-
stat ic QTLs for flower ing tmi e and maturity were
detected but much less than QTLs w ith add it ive
effects. Th is is genera lly cons istent w ith earlie r
mapping resu lts
[ 13, 15]
.
O f 12 and 7 ma in QTLs for p lant he ight and
flower ing tmi e, both 2 QTLs of each tra it were
identified in the same linkage groups as a lready
reported, but whether they are the identica lQTLs
need to be confirmed after comparat ive mapping
w ith the same f lank ing markers.
C lass ic quantitat ive geneticswere concentra t-
ed mostly on break ing tota l genet ic variat ion of
popula t ions in to d iffe rent components (additive,
dom inance and interaction ) to revea l genera l
properties of the genetic arch itec tures of quant ita-
t ive tra its, while QTLmapp ing d issect each part o f
genetic e ffect in to ind iv idua l genes, thus us ing
marker traced se lect ion to mi prove tra its interest-
ed. In princ ip le, above two methods are comple-
mented each other. The current QTL mapp ing a-
nalys is demonstra te that QTLs w ith additive effec t
are the predom inant part o f genetic bases and
QTLs w ith env ironment in teract ion (QE ) effec t
are a lso substantia l for a ll three agronom ic mi por-
tant tra its. These two factors work ing togethoe
may expla in the s ign ificant genet ic and GE inter-
action variance presented in Table 2.
Doubled hap lo id (DH) lines are complete ly
homozygous genotypes, w ih ich can pror ide unlmi -
ited seed supp lies for repeated expermi ents, while
it’ s major d isadvantage is mi poss ib le to estmi ate
dom inance effects and re la ted types of ep istas is
and in teract ionsw ith env ironmen.t The present re-
search showed that p lant he ight exh ib ited c learly
dom inant e ffect and the other two developmenta l
tra its a lso showed partia l dom inance (Table 1).
For complementing th is def ic iency, Zhang Q et
a l.
[ 20]
proposed to deve lop an F2 popula tion by
mak ing crosses among DH lines random ly, which
named mi mota lized( IF2) populat ion because they
can be reproduced year after year. Using th is IF2
popula tion for mapp ing expermi ent, additive
effects, dom inance effects, epistat ic e ffects of ad-
ditive ×add itive, add it ive ×dom inance, and dom i-
nance×dom inance and the ir in terac tions w ith en-
v ironment of QTLs can be identif ied and ana-
lysed. The IF2 popula tion from So llux /Gaoyou DH
lines were a lready deve loped last year and multi-
ple fie ld expenmi ents are go ing on. More genet ic
in format ion w ill be rece ived and analysed.
The resu lts in th is study revealed a number of
QTLs for three agronom ica lly mi portant tra its and
the ir complex ep istas is and genotype ×environ-
menta l in terac tions. SomeQTLsm ight be ident ica l
w ith these reported prev ious ly wh ile a large num-
ber ofQTLs were new ly ident ified. Mapping resu lts
are in good ag reement w ith the c lass ic genet ic
977ZHAO Jian-Yi et a l. :QTLs o f Th ree Ag ronom ica lly Im po rtan t T raits and Their Interactions…
analys is and demonstrate the re liab ility. The p le i-
otropy or close linkage of QTLs among tra its pro-
v ides the possib ilit ies for jo in t tra its se lection
based onmarkerass istance accord ing to breed ing
purposes and spec ific env ironments.
Acknow ledgments:We are grate fu l to P ro.f D r. Jun
Zhu, Institu te of B iogenes Mathematics, Zhejiang
University, fo r h is va luab le suggest ions on QTL
mapping. Thanks a lso to ZhouW e ijun, Inst itu te of
Agronomy, Zhejiang Univers ity, fo r h is k ind ly pro-
v id ing us h igh o il rapeseed materia ls. W e would
like to express our thanks to Joerg Schonde l-
ma ier, Saaten-Union Resistenzlabor GmbH , Hove-
dissen, Germany, fo r h is contribution to marker
ana lys is.
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