全 文 ：Received 7 Jul. 2003 Accepted 16 Dec. 2003
Supported by the State Key Basic Research and Development Plan of China (G1998010200) and the Hi-Tech Research and Development
(863) Program of China (2001AA211031).
* Author for correspondence. E-mail: .
http://www.chineseplantscience.com
Acta Botanica Sinica
植 物 学 报 2004, 46 (7): 867-872
Identification and Microsatellite Markers of a Resistance Gene to Powdery
Mildew in Common Wheat Introgressed from Triticum durum
ZHU Zhen-Dong, KONG Xiu-Ying, ZHOU Rong-Hua, JIA Ji-Zeng*
(Key Laboratory of Crop Germplasm and Biotechnology, Ministry of Agriculture, Institute of Crop Germplasm Resources,
The Chinese Academy of Agricultural Sciences, Beijing 100081, China)
Abstract: A powdery mildew resistance gene in a BC3F2 population, derived from a cross made between
an amphidiploid of Triticum durum Desf.-Aegilops caudata L. and T. aestivum L. cv. Laizhou 953, was
identified. Genetic analysis of resistance to powdery mildew in BC3F2 population and derived BC3F3 families
indicated a single dominant gene controlled the resistance. By bulk segregation analysis, two microsatellite
markers, Xgwm311 and Xgwm382, were identified to be closely linked to the resistance gene with genetic
distance of 5.9 cM and 4.9 cM, respectively. DNA from T. durum accession DR147, Ae. caudata acc. Ae14,
and recurrent parent wheat (Triticum aestivum L.) cv. Laizhou 953 were amplified with primer pairs WMS311
and WMS382, the specific bands related to the resistance gene were only present in T. durum acc. DR147.
Results showed that the resistance gene originated in T. durum acc. DR147. Based on the location of the
linked microsatellite markers, the resistance gene was located on the telomeric region of chromosome
2AL in wheat. Temporarily, the resistance gene was designated as PmDR147. The relation of this gene
and Pm4 was discussed.
Key words: resistance gene; microsatellite; molecular tagging; powdery mildew; durum wheat
Powdery mildew, caused by Erysiphe graminis f. sp.
tritici, is one of the most important diseases of wheat
(Triticum aestivum) and causes serious yield loss in wheat
growing regions with temperate climates. Breeding of re-
sistant cultivars is the economical and environmentally safe
way by reducing the application of fungicides to control
this disease. Up to now, thirty-one gene loci for resistance
to powdery mildew (Pm1– Pm31) have been identified and
assigned to specific chromosomes of wheat (Xie et al., 2001;
Liu et al., 2002; Zeller et al., 2002; McIntosh et al., 2003). A
few of these have been utilized in commercial breeding.
Because E. graminis f. sp. tritici has high genetic variability,
new virulent pathogen mutants can overcome individual
race-specific resistance genes in a relatively short period
(Duan et al., 1998). It is necessary to search for new sources
of resistance to guarantee the progress of resistance
breeding.
Durum wheat (Triticum durum) (2n = 4x = 28; genome
AABB) is a valuable source of genes for diversifying pest
resistance in common wheat. Durum wheat has been re-
ported to carry resistance to leaf rust, stem rust, stripe rust,
loose smut, Fusarium head blight, Septoria nodorum
blotch, Karnal bunt, powdery mildew, and Hessian fly
(McIntosh et al., 1967; Gupta et al., 1991; Cao et al., 2001;
Singh et al., 2001; Mishhra et al., 2001). It also contributes
towards superior quality characteristics.
Molecular markers have been widely used for gene
tagging, gene mapping, and other genetics research. These
DNA markers are not influenced by environmental condi-
tions and are detectable at all plant growth stages. Molecu-
lar markers tightly linked to the trait of interest can be used
in marker-assisted selection (MAS) to improve the efficiency
of conventional plant breeding (Gupta et al., 1999). Among
various molecular markers available, microsatellites, or
simple sequence repeats have been shown to be superior
to other DNA markers with respect to their higher level of
polymorphism and informativeness in hexaploid wheat
(Plaschke et al., 1995; Bryan et al., 1997). Moreover,
microsatellite markers are codominant, PCR-based, and func-
tional across the wild relatives of wheat (Rafalski and
Tingey, 1993; Röder et al., 1998). To date, several
microsatellite maps of wheat have been constructed, with
the microsatellite loci evenly distributed along the chromo-
some lengths to provide excellent coverage of the wheat
genome (Röder et al., 1998; Stephenson et al., 1998;
Pestsova et al., 2000; Gupta et al., 2002). Some microsatellite
markers from hexaploid wheat also have been integrated
into genetic linkage maps of durum wheat (Korzun et al.,
Acta Botanica Sinica 植物学报 Vol.46 No.7 2004868
1999; Nachit et al., 2001). Closely linked microsatellite mark-
ers to many economically important trait genes including a
few powdery mildew resistance genes in wheat have been
identified (Zhu and Jia, 2003).
The present study reports the identification and
microsatellite markers of a powdery mildew resistance gene
in common wheat introgressed from durum wheat.
1 Materials and Methods
1.1 Plant materials
Triticum durum Desf. accession DR147, Aegilops
caudata L. acc. Ae14, recurrent parent wheat (Triticum
aestivum L.) cv. Laizhou 953, cv. Chinese Spring, the nulli-
tetrasomic (NT) stocks N2AT2B, N2BT2A and N2DT2A of
Chinese Spring were from the author’s institute.
A cross was made between the amphidiploid of T. du-
rum acc. DR147/Ae. caudata acc. Ae14 and T. aestivum cv.
Laizhou 953, an agronomically superior wheat cultivar (Kong
et al., 1999). The F1 was backcrossed to Laizhou 953 for
three generations. Then a number of powdery mildew re-
sistant BC3 progenies were selfed. A BC3F2 population and
derived BC3F3 families were used in the study.
1.2 Powdery mildew evaluation
Erysiphe graminis f. sp. tritici (Egt) isolate E09 was
provided by Drs. DUAN Xia-Yu and ZHOU Yi-Lin, Insti-
tute of Plant Protection, The Chinese Academy of Agricul-
tural Sciences. Resistance to powdery mildew of BC3F2
population was identified in the field by inoculating Egt
isolate E09. The method of inoculation, conditions of incu-
bation and disease assessment were according to Sheng
(1991). T. durum acc. DR147, Ae. caudata acc. Ae14, recur-
rent parent wheat cv. Laizhou 953 and BC3F2-derived F3
families were evaluated for resistance to Egt isolate E09 at
the seedling stage in a greenhouse. The method of
inoculation, conditions of incubation and disease assess-
ment were according to Duan et al. (2001). Egt isolate E09 is
a prevalent virulence type in Beijing area with a virulence
formula Pm1, Pm3a, Pm3c, Pm5, Pm7, Pm8, Pm17, Pm19.
Chi-squared tests for goodness of fit were used for devia-
tion of observed data from theoretically expected segrega-
tion ratios. Six infection types of hosts were distinguished:
0=immune, 0; = near immune, 1= high resistant, 2 = moder-
ately resistant, 3 = moderately susceptible, and 4 = high
susceptible (Sheng, 1991; Duan et al., 2001).
1.3 PCR amplification and product analysis
Genomic DNA was extracted from leaf tissue samples as
described by Sharp et al. (1988). Wheat microsatellite mark-
ers employed were described in Röder et al. (1998). Each
PCR reaction contained 80 ng genomic DNA, 0.25 µmol/L
of each primer, 1 U Taq DNA polymerase, 2 µL of 10 ×
PCR buffer containing 15 mmol/L MgCl2, 0.2 mmol/L of
dNTPs in a total volume of 25 µL. PCR was performed in an
MJ Research thermocycler, at 94 ℃ for 5 min, followed by
35 cycles of 94 ℃ for 1 min, 50, 55, or 60 ℃ (based on primer
annealing temperature) for 1 min, and 72 ℃ for 1 min, with
final incubation at 72 ℃ for 5 min before cooling to 4 ℃.
Each PCR product was denatured by adding 8 µL load-
ing buffer ( 98% formamide, 10 mmol/L EDTA, pH 8.0,
0.25% bromo-phenol blue, and 0.25% xylene cyanol) and
denatured for 10 min at 95 ℃ and chilled on ice. Eight µL of
each sample was loaded on 6% polyacyamide (19:1
acrylamide:Bis), 8 mol/L urea and 1× TBE (90 mmol/L Tris-
borate, pH 8.3, and 2 mmol/L EDTA) gels (40 cm length, 20
cm wide, 0.2 cm thick). Samples were then run at 2 500 V, 30
mA, 100 W for approximate 1 h, and the products were
visualized by silver staining (Tixier et al., 1997).
1.4 Microsatellite marker analysis
The identification of microsatellite markers linked to the
resistance gene was accomplished by bulked segregation
analysis (BSA) as described by Michelmore et al. (1991).
The resistant bulk and the susceptible bulk were made by
separately pooling equal amount of DNA from 10 resistant
and 10 susceptible plants from the BC3F2 segregating
population. The markers generating polymorphic
microsatellite fragments between the bulks were further
checked for their linkage to the resistance gene, using 108
plants of the BC3F2 population.
1.5 Linkage analysis
Recombination frequencies (RF) or linkage relationships
between microsatellite markers and the powdery mildew
resistance gene were calculated using Mapmaker 3.0b
(Lander et al., 1987) and converted to cM using the Kosambi
mapping function (Kosambi, 1944). The decimal logarithm
of likelihood ratio (LOD) was used as a test measure of
reliability of linkage. The linkage is reliable if the markers
were placed with an LOD threshold of 3.0.
2 Results
2.1 Powdery mildew resistance
T. durum acc. DR147 and Ae. caudata acc. Ae14 were
resistant to Egt isolate E09, recurrent parent wheat cv.
Laizhou 953 was susceptible to Egt isolate E09. A total of
132 BC3F2 individuals were identified for resistance to pow-
dery mildew in the field. The observed segregation of 100
resistant and 32 susceptible individuals fitted a 3:1 segre-
gation ratio (Table 1). A total of 132 BC3F2-derived F3 fami-
lies were tested with Egt isolate E09 in seedling to deter-
mine genotype of BC3F2 individuals. Thirty families were
ZHU Zhen-Dong et al.: Identification and Microsatellite Markers of a Resistance Gene to Powdery Mildew in Common Wheat
Introgressed from Triticum durum 869
homozygous resistance, 75 families were heterozygous re-
sistant and 27 families were homozygous susceptible. The
observed segregation fitted a 1:2:1 segregation ratio (c2 =
2.59). Both ratios supported a segregation of a single domi-
nant locus. These results showed that the powdery mildew
resistance of the BC3F1 derived from the amphidiploid of T.
durum acc. DR147-Ae. caudata acc. Ae14 is controlled by
single dominant gene.
2.2 Microsatellite markers linked to the resistance gene
Based on the microsatellite map of wheat (Röder et al.,
1998), 136 microsatellite markers were screened to identify
polymorphic microsatellite markers between the resistant
and susceptible DNA bulks. Two primer pairs WMS311
and WMS382 generated polymorphic DNA fragments be-
tween the bulks with size about 170 bp and 130 bp,
respectively. Markers Xgwm311 and Xgwm382 were in the
resistant pool, but absent in susceptible pool. These re-
sults indicated the two microsatellite markers could be linked
to the resistance gene. Further, the two markers were used
to check their linkage to the resistance gene using 108 plants
of segregating BC3F2 population. The data is in Table 1,
illustrated in Figs.1 and 2. Both markers Xgwm311 and
Xgwm382 showed a 3:1segregation ratio, and were closely
linked to the resistance gene with a map distance of 5.9 cM
and 4.9 cM, respectively (Fig.3).
2.3 Origin and chromosomal location of the resistance
gene
Parents of the amphidiploid, T. durum acc. DR147 and
Ae. caudata acc. Ae14, were resistant to powdery mildew.
To determine the origin of the resistance gene in the BC3F2
population, we amplified DNA from T. durum acc. DR147,
Ae. caudata acc. Ae14 and recurrent parent wheat cv.
Laizhou 953 with primer pairs WMS311 and WMS382. The
markers Xgwm311 and Xgwm382 related to resistance gene
were present only in T. durum acc. DR147. The results indi-
cated the resistance gene was from T. durum acc. DR147.
Temporarily, the powdery mildew resistance gene was des-
ignated as PmDR147.
According to microsatellite map of wheat (Röder et al.,
1998), marker Xgwm311 was on wheat chromosome 2AL
and 2DL, while marker Xgwm382 was on wheat chromo-
some 2AL, 2BL and 2DL. To verify the chromosome loca-
tion of markers Xgwm311 and Xgwm382, the DNA from the
NT stocks N2AT2B, N2BT2A and N2DT2A of Chinese
Table 1 Segregation analysis for the resistance gene and microsatellite markers linked to the gene
Gene or marker
Number of Observed number
c23:1 P
BC3F2 D LL
PmDR147 132 100 32 0.04 0.99–0.95
Xgwm311 108 81 27 0.01 >0.99
Xgwm382 108 82 26 0.01 >0.99
D , DR147 or heterozygous; LL, Laizhou 953. c23:1=3.84, P=0.05.
Fig.1. PCR bands amplified from the DNA of selected BC3F2
plants using primer pair WMS311. L, pBR322 DNA/MspⅠ
marker; R, resistance plant; S, susceptible plant; ®, specific
bands linked to powdery mildew resistance gene from DR147.
Fig.2. PCR bands amplified from the DNA of selected BC3F2
plants using primer pair WMS382. L, pBR322 DNA/Msp Ⅰ
marker; R, resistance plant; S, susceptible plant; ®, specific bands
linked to powdery mildew resistance gene from DR147.
Fig.3. Genetic linkage map of powdery mildew resistance gene
PmDR147 and the linked microsatellite markers on wheat chro-
mosome 2A. The genetic distances between gene Pm4 and
Xgwm311 were based on published data by Röder et al. (1998).
Acta Botanica Sinica 植物学报 Vol.46 No.7 2004870
Spring wheat was amplified with primer pairs WMS311 and
WMS382. Both markers were located on wheat chromo-
somes 2A and 2D. Figure 4 shows the location of Xgwm311.
On a molecular linkage map for an intraspecific recombi-
nant inbred population of durum wheat, two nearby
microsatellite Xgwm382 loci are located on chromosome
2AL of durum wheat (Nachit et al., 2001). Since the resis-
tance gene PmDR147 was from T. durum (genome AABB)
and closely linked to the markers Xgwm311 and Xgwm382,
it should be on the long arm of wheat chromosome 2A.
3.2 Relationship between PmDR147 and Pm4
Classical genetic and molecular data show the genes for
disease resistance are not randomly distributed all over the
genome but rather frequently occur in clusters in particular
chromosomes. For example, among the documented 28 pow-
dery mildew resistance gene loci, Pm1, Pm3, Pm4, or Pm5
is a complex locus, and composed of 4, 10, 2, and 5 alleles,
respectively (Hsam et al., 1998; Zeller et al., 1998; Huang et
al., 2003). In our study, we located the gene PmDR147 on
the distal region of long arm of chromosome 2A. Powdery
mildew resistance genes Pm4a and Pm4b are also located
on chromosome 2AL of wheat. Ma et al. (1994) identified
RFLP markers linked to Pm4a using near-isogenic lines.
Two markers Xcdo1231-2A (2) and Xcdo678-2A were found
to be cosegregant with Pm4a. On genetic map of wheat,
RFLP marker Xcdo678 is above microsatellite marker
Xgwm382 on the wheat chromosome 2AL, with a map dis-
tance of 8.3 cM (Röder et al., 1998). The gene PmDR147 is
below marker Xgwm382 with 4.9 cM. An estimative genetic
distance between PmDR147 and Pm4a is 13.2 cM (Fig.3).
Pm4a and Pm4b originated in T. dicoccum and T. carthlicum,
respectively, while PmDR147 derived from T. durum.
Therefore, we assume either the PmDR147 and Pm4a or
Pm4b may be different genes present in a complex region
on chromosome 2AL, or PmDR147 may be an allele of the
complex Pm4 locus. Because of different mapping popula-
tion type and size, the location of the same gene locus on
different genetic maps was probably discrepant. In order to
confirm the genetic relationship between PmDR147 and
Pm4, further allelism tests should be done. However,
whether the gene from DR147 is the same as Pm4 or not,
the markers closely linked to the resistance gene found in
present study can be used for MAS and germplasm
diagnostics.
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