Expression quantitative trait loci analysis of BAN, F3H and TT19 genes in Brassica
napus
Kun Lu, Cunmin Qu, Kai Zhang, Hao Yu, Junxin Lu, Yourong Chai, Jiana Li*
Chongqing Rapeseed Engineering & Technology Research Center; Engineering Research Center of South Upland
Agriculture, Ministry of Education, P. R. China; College of Agronomy and Biotechnology, Southwest University,
Tiansheng Road 2, Beibei, Chongqing 400715, P. R. China
ljn1950@swu.edu.cn drlukun@swu.edu.cn
Abstract—Flavonone 3-hydroxylase (F3H), anthocyanidin
reductase (ANR) encoded by BANYULS (BAN) gene and
glutathione S-transferase encoded by TRANSPARENT TESTA
19 (TT19) gene are three key enzymes in flavonoid pathway,
playing important roles in the synthesis and transport of
anthocyanin and proanthocyanidin. However, little is known
about the expression variations and the regulatory network in
Brassica. In this study, we examined their expression levels in
seeds of 30 days after flowering (DAF) in F
7
recombinant
inbred lines (RILs) derived from a cross between Brassica
napus cultivars Youyan2 (black-seeded) and GH06 (yellow-
seeded), and characterized the locus-level regulatory network
related to B. napus BAN, F3H and TT19 genes. Expression
quantitative trait loci (eQTL) mapping revealed five, seven and
eight eQTLs for BAN, F3H and TT19 genes, respectively.
Marker E4M8 was associated with qBAN-8-2 and qTT19-8-2,
while marker H022L18-2 was linked with qBAN-16-4 and
qF3H-16-6, implying that there might be upstream regulatory
genes in E4M8 and H022L18-2 marker flanking regions. In
further analysis, we identified four trans-eQTLs (qBAN-16-5,
qF3H-16-5, qBAN-16-4 and qF3H-11-2) close to the location of
the major QTL controlling yellow-seeded trait of B. napus. The
200-kb flanking sequences of the four eQTL on B. rapa
chromosome A09 showed well synteny to B. oleracea genome
sequence and partial continuous fragment of chromosome 1 of
Arabidopsis genome, suitable for candidate gene predication of
eQTLs. Based on gene finding results, seven transcription
factors were suggested to be the potential upstream
candidate(s) controlling expression variations of BAN, F3H
and TT19 genes. These results could provide a new approach
for constructing regulatory pathways that contribute to
complex traits, such as yellow-seeded trait.
Keywords-Brassica napus; transparent testa; expression
quantitative trait loci; regulatory network
I. INTRODUCTION
Oilseed rape (Brassica napus, AACC, 2n=4x=38) is one
of main oil crops in the world. It is an important source to the
vegetable oil, meal protein and industry material. Compared
with black-seeded cultivars, yellow-seeded B. napus shows
many good quality traits, such as lower seed coat pigment
content, lower meal fiber content, higher seed oil and meal
protein content, etc. Selecting of yellow-seeded oilseed rape
cultivars with stably inherited yellow seed trait and good
agronomic traits has become the most important breeding
goals. However, it is very slow advance and difficult
improvement to breeding of yellow-seeded B. napus, since
the genetic and molecular mechanisms of yellow-seed trait
were still unclear.
It had been proved that plant seed coat pigments are
mainly composed of polymers of proanthocyanidin (PA),
which is synthesized via the flavonoid-anthocyanin-
proanthocyanidin pathway (simplified as flavonoid pathway
here), a core branch pathway of phenylpropanoid pathway.
F3H, BAN and TT19 are 3 key enzymes in flavonoid
pathway, playing important roles in the synthesis and
transport of anthocyanin and proanthocyanidin. In
Arabidopsis thaliana, the loss of function mutants of the 3
genes all leads to the transformation of the wild-type dark-
brown seed to yellow or pale yellow seed, i.e. transparent
testa (TT) trait [1]. At present, the gene families encoding the
3 key enzymes have been cloned by RACE method in our
lab, which laid the foundation to further study their
molecular and regulatory mechanism.
In order to determine the upstream regulatory network of
the 3 gene families, analysis of transcript abundance was
carried out on RNA from seeds of 30 DAF in RIL
population. Regarding as quantitative traits, the transcript
levels of the 3 gene families were examined by QTL
mapping method for eQTL detection. Using this method, it
was possible to construct regulatory pathways that contribute
to complex traits, demonstrating that the principles of eQTL
mapping could be applied to B. napus.
II. MATERIALS AND METHODS
A. Plant materials and total RNA extraction
The RILs were developed through successive selfing up
to six generations from a cross between yellow-seeded
female parent ‘GH06’ and black-seeded male parent
‘Youyan2’ by single seed descent (SSD). Parental lines and
RILs were sown in field trials at plant breeding station of
Chongqing Rapeseed Technology Research Center
(CRTRC) in 2009 as previously described [2]. Seeds of 30
DAF in 180 F7 recombinant inbred lines were harvested and
used for total RNA isolation. Total RNA was extracted using
the Plant RNA Mini Kit (Watson Biotechnologies, Inc.,
China). To remove contaminated genomic DNA, the total
RNA was treated with RNase-free DNase I (TaKaRa).
Quality and concentration of total RNA samples were
2012 International Conference on Biomedical Engineering and Biotechnology
978-0-7695-4706-0/12 $26.00 © 2012 IEEE
DOI 10.1109/iCBEB.2012.214
272