Vol. 99, No. 10, 2009 1121
Genetics and Resistance
Quantitative Trait Loci Mapping for Adult-Plant Resistance
to Powdery Mildew in Chinese Wheat Cultivar Bainong 64
Caixia Lan, Shanshan Liang, Zhulin Wang, Jun Yan, Yong Zhang, Xianchun Xia, and Zhonghu He
First, second, fifth, sixth, and seventh authors:
Crop Gene Resources and Genetic Improvement, Chinese Academy of Agricultural Sciences (CAAS), 12 Zhongguancun South Street,
Beijing 100081, China; second author: Department of Primary Industries, Victorian AgriBiosciences Center, La Trobe R&D Park, 1 Park
Drive, Bundoora, Vic 3083, Australia; third author: College of Agronomy, Northwest Sci-Tech University of Agriculture and Forestry,
Yangling, Shaanxi 712100, China; fourth author:
Huanghedadao, Anyang, Henan 455000, China; and seventh author:
China Office, c/o CAAS, 12 Zhongguancun South Street, Beijing 100081, China.
Accepted for publication 28 May 2009.
Institute of Crop Science, National Wheat Improvement Center/The National Key Facility for
Cotton Research Institute, Chinese Academy of Agricultural Sciences (CAAS),
International Maize and Wheat Improvement Center (CIMMYT)
Lan, C., Liang, S., Wang, Z., Yan, J., Zhang, Y., Xia, X., and He, Z.
2009. Quantitative trait loci mapping for adult-plant resistance to
powdery mildew in Chinese wheat cultivar Bainong 64. Phytopathology
Adult-plant resistance (APR) is an effective means of controlling
powdery mildew in wheat. In the present study, 406 simple-sequence
repeat markers were used to map quantitative trait loci (QTLs) for APR
to powdery mildew in a doubled-haploid (DH) population of 181 lines
derived from the cross Bainong 64 × Jingshuang 16. The DH lines were
planted in a randomized complete block design with three replicates in
Beijing and Anyang during the 2005–06 and 2007–08 cropping seasons.
Artificial inoculations were carried out in Beijing using the highly
virulent Blumeria graminis f. sp. tritici isolate E20. Disease severities on
penultimate leaves were scored twice in Beijing whereas, at Anyang,
maximum disease severities (MDS) were recorded following natural
infection. Broad-sense heritabilities of MDS and areas under the disease
progress curve were 0.89 and 0.77, respectively, based on the mean
values averaged across environments. Composite interval mapping
detected four QTLs for APR to powdery mildew on chromosomes 1A,
4DL, 6BS, and 7A; these were designated QPm.caas-1A, QPm.caas-4DL,
QPm.caas-6BS, and QPm.caas-7A, respectively, and explained 6.3 to
22.7% of the phenotypic variance. QTLs QPm.caas-4DL and QPm.caas-
6BS were stable across environments with high genetic effects on
powdery mildew response, accounting for 15.2 to 22.7% and 9.0 to
13.2% of the phenotypic variance, respectively. These results should be
useful for the future improvement of powdery mildew resistance in
Additional keywords: Triticum aestivum.
Powdery mildew, caused by Blumeria graminis f. sp. tritici, is
a very destructive foliar disease of wheat (Triticum aestivum L.),
especially in regions with high rainfall and with a maritime or
semicontinental climate (1). Race-specific major resistance genes
(Pm genes) confer complete resistance to powdery mildew result-
ing from hypersensitive responses (HR) to infection (9). How-
ever, this kind of resistance is often not durable, being overcome
by new pathogen races possessing the corresponding virulence
genes (26). Some wheat cultivars exhibit resistance that delays
infection and reduces growth and reproduction of the pathogen on
post-seedling plants. Such resistance has been termed slow mil-
dewing (26), adult-plant resistance (APR) (9), or partial resistance
(11). So far, 58 powdery mildew resistance genes at 39 loci have
been characterized and assigned to specific wheat chromosomes
jsp), and some of them were introduced from wheat-related
species. Most of these genes are race specific and many of them
have been overcome by new pathogen variants.
In contrast, APR genes are likely to be more useful for long-
term disease resistance strategies and, thus, have greater potential
for durable resistance (1). APR genes are expressed quantitatively
and typically have small genetic effects, thereby requiring multi-
ple genes to provide adequate levels of protection against patho-
gens (9,20). Investigations of the chromosomal locations of such
genes and their biological effects are important to ensure their
appropriate deployment in elite germplasm and commercial wheat
APR to powdery mildew is easy to identify in cultivars pos-
sessing no seedling-effective resistance genes or when natural
populations of B. graminis f. sp. tritici overcome any resistance
gene that may be present (1,26). It is time-consuming and
difficult to assess APR phenotypes in breeding populations in the
field (9); therefore, molecular markers will have many advantages
in the selection of genotypes with combinations of APR genes.
Molecular markers have been used to map APR genes for pow-
dery mildew in wheat. For example, quantitative trait loci (QTLs)
for APR were mapped in the Swiss winter wheat Forno (15), the
French winter wheat lines RE714 (5,23) and RE9001 (3), the
North American winter wheat lines Massey (20) and USG3209
(37), the Japanese cv. Fukuho-Komugi (a QTL on 7DS identified
as Pm38) (18), the Korean wheat Suwon 92 (42), and Inter-
national Maize and Wheat Improvement Center (CIMMYT) line
Saar (two QTLs identified as Pm38 and Pm39) (19).
Bainong 64 was a leading wheat cultivar in the Yellow-Huai
wheat region of China from 1993 to 2000, occupying ≈700,000 ha
annually. It showed highly susceptible responses to many isolates
of B. graminis f. sp. tritici at the seedling stage but was highly
resistant at the adult stage (39). This cultivar had good quality,
high yield, and broad adaptability, and was used widely as a
parent in Chinese wheat-breeding programs. The aim of the
Corresponding authors: X. Xia and Z. He;
E-mail addresses: email@example.com and firstname.lastname@example.org
© 2009 The American Phytopathological Society
present study was to identify QTLs for APR to powdery mildew
and associated molecular markers in a doubled-haploid (DH)
population from a cross between Bainong 64 and Jingshuang 16,
and to assess the stability of the effects of the detected QTLs
across different environments. It was expected that such knowl-
edge could lead to a more efficient strategy in breeding for
MATERIALS AND METHODS
Plant materials. The 181 DH lines used in the study were
developed from a cross between Chinese winter wheat cvs.
Bainong 64 and Jingshuang 16, and were generated by the wheat
× maize crossing method (17). Bainong 64 (Bainong 8717/3/Yeda
72-629-52/Shi 82-5594//Bainong 84-4046-1) is susceptible to
Chinese isolates of B. graminis f. sp. tritici at the seedling stage
but highly resistant at the adult-plant stage, whereas Jingshuang
16 is highly susceptible at both the seedling and adult-plant stages
(39). These isolates were collected in Beijing, Henan, Guizhou,
Yunnan, and Sichuan provinces from 1990 through 1998, and
maintained on wheat seedlings in growth chambers with 24 h of
light at 4 to 8°C, in the Institute of Plant Protection, Chinese
Academy of Agricultural Sciences (CAAS).
Field trials. The DH lines and parents were evaluated for APR
to powdery mildew during the 2005–06 and 2007–08 cropping
seasons at the CAAS experimental station of the Institute of Crop
Science in Beijing, and at the CAAS Cotton Research Institute,
Anyang. Field trials were planted in randomized complete blocks
with three replicates. Plots consisted of single 1.5-m rows with 25
cm between rows. In all, ≈75 seeds were sown in each row. The
susceptible parent cv. Jingshuang 16 was planted every 10 rows
as a susceptible check and planted around the test lines to ensure
ample powdery mildew inoculum in spring.
Artificial inoculations in Beijing were carried out using the
highly virulent B. graminis f. sp. tritici isolate E20 prior to the
plants reaching stem elongation. The disease severity on the
penultimate leaf (F-1 leaf) on 10 randomly selected plants from
each line was scored based on the percentage of leaf area covered
by powdery mildew for the first time in 6 weeks after inoculation,
and then after 1 week for the second time when the disease
severity reached a maximum level at ≈20 May. Disease severity
of the 10 selected plants was averaged to obtain mean powdery
mildew severity for each line.
Powdery mildew severities on each line in Anyang were rated
on the percentages of leaf area covered by powdery mildew under
natural infection conditions when the disease severities on Jing-
shuang 16 reached a maximum level at ≈18 May 2006. Field data
from the 2008 trial were not used for analysis because disease
severities failed to reach satisfactory levels; for instance, the
averaged maximum disease severities (MDS) of susceptible check
cv. Jingshuang 16 was <5%.
Statistical analysis. The area under the disease progress curve
(AUDPC) was calculated for each line using the following
formula from Bjarko and Line (2):
)(2 / )
where Xi is the disease severity on assessment date i, Ti is the
number of days after inoculation on assessment date i, and n is the
total times of disease assessments. MDS and AUDPC were used
for the subsequent analysis of variance (ANOVA) and QTL
analysis. SAS software (SAS Institute, Cary, NC) was used to
compute ANOVA. Broad-sense heritability (h2) for powdery
mildew resistance was calculated from the ANOVA, on an
across-environment genotype mean basis, by the formula h2 =
σ2g/(σ2g + σ2ge/e + σ2ε/re), where σ2g, σ2ge, and σ2ε are estimates of
genotypic, genotype–environment interaction, and error vari-
ances, respectively, and e and r are the numbers of environments
and replications per environment, respectively. Each QTL was
represented by a 20-centimorgan (cM) interval with the local
logarithm of odds (LOD) maximum as center. QTLs with over-
lapping 20-cM intervals among different environments were
considered as being in common.
Simple-sequence repeat analysis. To construct the framework
map for the QTL analysis, we chose 406 simple-sequence repeats
(SSRs), including 140 pairs of Beltsville Agriculture Research
Center (BARC) primers (33), 134 pairs of Wheat Microsatellite
Consortium (WMC) primers (8), 81 pairs of Gatersleben Wheat
Microsatellite (GWM) primers (27), 30 pairs of Clermont Ferrand
D-genome (CFD) primers (10), 14 pairs of Gatersleben D-ge-
nome Microsatellite (GDM) primers (25), and 7 pairs of Clermont
Ferrand A-genome (CFA) primers (34).
Bulked segregant analysis. Two DNA bulks were constructed
by mixing equal amounts of DNA from the five most resistant
and the five most susceptible lines, respectively, based on severity
data from all three environments. SSRs showing the same
patterns of polymorphism between the parents and between the
bulks were used to genotype all 181 DH lines. Additional SSRs
on the linkage maps (http://wheat.pw.usd a.gov/GG2/index.shtml,
around the initially indicated loci associated with APR were
chosen to genotype the population for linkage and QTL analyses
as described by Michelmore et al. (22).
Map construction and QTL detection. Linkage groups were
generated with the software Map Manager QTXb20 (21). Genetic
distances between markers were estimated using the Kosambi
(16) mapping function. QTL were detected by composite interval
mapping using the software Cartographer 2.5 (38). An LOD of
2.0 was set to declare QTL as significant. For each QTL, esti-
mates of phenotypic variance (R2) and additive effects at the LOD
peaks were obtained from QTL Cartographer 2.5.
Phenotypic analysis of MDS and AUDPC, and their
correlations and heritabilities. The MDS scores from the three
environments were significantly correlated (P < 0.0001), with
correlation coefficients of 0.61 to 0.76. The frequency distribu-
tions of the powdery mildew severity parameters (MDS and
AUDPC) of the DH lines over the three environments showed
continuous variation, confirming quantitative inheritance (Fig. 1).
The mean MDS of Bainong 64 and Jingshuang 16 were 2.3 and
28.3% in Anyang in 2006, whereas the means were 1.0 and
30.0% and 0.7 and 86.7% in Beijing in 2006 and 2008, respec-
tively. The average of MDS of the DH lines in Beijing over 2
years was 10.4% (range: 0.3 to 86.7%) and 7.3% in Anyang
(range: 0 to 60.0%). In Beijing, the average of AUDPC over 2
years was 51.4 (range: 1.2 to 298.3).
The MDS and AUDPC were significantly correlated for the test
in Beijing over 2 years (r = 0.89, P < 0.0001). The broad-sense
heritabilities of MDS and AUDPC across environments were 0.89
and 0.77, respectively. The ANOVA confirmed a significant
variation among DH lines (Table 1).
QTL for APR to powdery mildew. Four QTLs for APR were
detected in the DH population in three environments (Table 2;
Fig. 2). Using the MDS from the 2005–06 cropping season in
Anyang, three QTLs were detected on chromosomes 1A, 4DL,
and 6BS with additive effects of 4.00, 6.95, and 4.98 explaining
7.4, 20.9, and 13.2% of the phenotypic variance, respectively.
Four MDS QTLs were identified in Beijing in the 2005–06
cropping season, accounting for 9.9, 22.7, 11.5, and 6.3% of the
phenotypic variance, respectively. The additive effects of these
QTLs were 1.68, 2.82, 1.93, and 1.52, respectively. In Beijing in
2007–08, three QTLs were detected on chromosomes 4DL, 6BS,
and 7A, explaining 16.7, 10.3, and 7.1% of the phenotypic vari-
ance, respectively. The additive effects of these QTLs were 6.33,
Vol. 99, No. 10, 2009 1123
6.79, and 5.03, respectively. Using the average values of MDS
from three environments over 2 years, four QTLs were mapped
on chromosomes 1A, 4DL, 6BS, and 7A with additive effects of
3.10, 3.42, 4.36, and 2.90, respectively, explaining 6.7 to 15.2%
of the phenotypic variance. Based on averaged AUDPC values in
Beijing over two cropping seasons, three QTLs were found on the
chromosomes 4DL, 6BS, and 7A explaining 6.7 to 19.0% of the
phenotypic variance, and the additive effects of these QTLs were
16.14, 19.63, and 17.05, respectively. All QTLs identified in this
population were from the resistant parent Bainong 64.
In a previous study, it was estimated that Bainong 64 possessed
three to four genes for APR to powdery mildew, based on
phenotypic data from F2:3 populations of Bainong 64 × Jing-
shuang 16 (40). This was in agreement with the present study,
where four QTLs were identified in Bainong 64 using DH lines
from the same cross.
The QTL in the centromeric region of chromosome 1A was
detected in three environments in the SSR interval Xbarc148–
Xwmc550. This was in a similar position to a QTL found by
Keller et al. (15) but flanked by amplified fragment length poly-
morphism (AFLP) markers Xpsr1201b and Xpsr941M in a
segregating wheat × T. spelta population. Mingeot et al. (23)
detected a QTL in the marker interval Xcdo572b–Xbcd442 in the
French line RE714, and this location is close to QPm.caas-1A
based on the consensus map of Somers et al. (32). Therefore,
these QTLs could be at the same locus or linked closely. In
contrast, Liang et al. (18) identified a QTL in the marker interval
Xgdm33–Xpsp2999 in a DH population from Fukuho-Komugi ×
Oligoculm at a similar location to the Pm3 locus; this position is
clearly different from that of QPm.caas-1A. Two seedling-effec-
tive race-specific resistance genes have been identified on chro-
mosome 1A—namely, Pm25 derived from T. aegilopoides (30)
and Pm17 from rye (12)—however, QPm.caas-1A is unlikely to
be related to those genes. Although QPm.caas-1A had a small
effect (Table 2), it was detected in two environments in 2006 and
the average MDS in three environments and was close to wheat
SSR marker Xwmc550. Its relatively stable contribution to resis-
tance in two environments and its detection in a previous study
indicates a significant role in APR.
Fig. 1. A to E, Frequency distributions of powdery mildew maximum disease severities (MDS) and area under the disease progress curve (AUDPC) values in
doubled-haploid (DH) lines derived from Bainong 64 × Jingshuang 16. A, MDS, Anyang 2006; B, MDS, Beijing 2006; C, MDS, Beijing 2008; D, average MDS
across three environments; and E, average AUDPC, Beijing over 2 years. Mean values for the parents, Bainong 64 and Jingshuang 16, are indicated by arrows.
The QPm.caas-4DL was proximal to the wheat microsatellite
marker Xwmc331 on the long arm of chromosome 4D. Keller et
al. (15) previously detected a QTL at the end of chromosome 4DL
in the marker interval of Xglk302b–Xpsr1101a in a segregating
wheat × T. spelta population. This location was different from that
of QPm.caas-4DL by a map distance of nearly 50 cM (http://wheat.
fore, QPm.caas-4DL is likely to be a new gene conferring APR to
powdery mildew. It had a high LOD score, a large genetic effect
on resistance, and was stably detected across three environments.
The SSR marker Xwmc331, closely linked to QPm.caas-4DL,
could be used for marker-assisted selection in wheat breeding.
QTL QPm.caas-6BS was proximal to the molecular marker
Xbarc79 at a genetic distance of 0.0 to 11.9 cM. Keller et al. (15)
detected a QTL in the AFLP marker interval Xpsr167b–Xpsr964
on chromosome 6BL in a segregating wheat × T. spelta popu-
lation. Based on the consensus map, that gene should be different
from QPm.caas-6BS. Five race-specific seedling-effective resis-
tance genes have been identified on the chromosome 6B: Pm11
(36), Pm12 (14), Pm14 (35), Pm20 (6), and Pm27 (13). Pm11 and
Pm14 are effective only against certain wheat grass B. graminis f.
sp. tritici cultures, and Pm12, Pm20, and Pm27 were derived
from alien species; namely, Aegilops speltoides (14), rye (6), and
T. timopheevii (13), respectively. Thus, QPm.caas-6BS is unlikely
to correspond to a known Pm gene or QTL on chromosome 6B. It
showed a stable genetic effect against powdery mildew across
three environments, explaining 9.0 to 13.2% of the phenotypic
variance. SSR marker Xbarc79, closely linked to QPm.caas-6BS,
could be used for improving wheat powdery mildew resistance in
molecular breeding programs.
The QTL detected at the centromeric region of chromosome 7A
was proximal to the wheat microsatellite marker Xbarc174.
Chantret et al. (4) identified a QTL in the interval Xfba069–
Xgwm344 in an RE714 × Hardi population but that region is quite
distal from the location of QPm.caas-7A based on the consensus
map. Several seedling-effective major gene loci have been located
on chromosome 7A, including Pm1 (29), Pm9 (28), Pm37 (24),
and mlRD30 (31). QPm.caas-7A is unlikely to be any of these
genes or is unlikely to represent the effects of defeated alleles at
any of these loci. It was detected in Beijing in two environments,
and also identified in the mean MDS and AUDPC analyses. This
QTL was proximal to marker Xbarc174 within genetic distance of
the 0.0- to 2.0-cM region. That marker could be used for marker-
In the crosses of the 11 cultivars and advanced lines derived
from Bainong 64, including the released cvs. 04 Zhong 36,
Huaimai 0320, Xu 9908, and Yuanyu 3 and advanced lines
13397, 03 Zhong 35, Bainong 69, Junmai 35, Ruifeng 97-6,
Tianmin 668, and Xumai 4060, all of them have showed moderate
resistance to powdery mildew at adult-plant stage in the field,
TABLE 2. Quantitative trait loci (QTLs) detected for adult-plant resistance (APR) to powdery mildew in the doubled-haploid (DH) population derived from the
cross Bainong 64 × Jingshuang 16
Parametera, year, and location QTLb Interval Positionc LODd Additive effect
R2 (%)e Total R2 (%)
Average MDS in three environments
Average AUDPC in Beijing
a MDS = maximum disease severity and AUDPC = area under the disease progress curve.
b QTL that extend across single one-log support confidence intervals were assigned the same symbol.
c Peak position in centimorgans from the first interval marker.
d Logarithm of odds (LOD) score.
e R2 is the proportion of phenotypic variance explained by the QTL.
TABLE 1. Analysis of variance of maximum disease severities (MDS) values for penultimate leaves and area under the disease progress curve (AUDPC) values
for powdery mildew responses on doubled-haploid (DH) lines derived from the cross Bainong 64 × Jingshuang 16
Parameter, source of variation df Sum of squares Mean of squares
77.11 38.55 1.37
a Asterisks (**) indicate significant at P < 0.0001.
Vol. 99, No. 10, 2009 1125
except for Huaimai 0320 (7,43). In the crosses for the 11 cultivars
and advanced lines, all the other parents were susceptible to
powdery mildew, except for Zhoumai 16, another resistant parent
of Junmai 35 (41,44). This indicates that Bainong 64 is a good
resistant parent in Chinese wheat-breeding programs.
In this study, we detected four QTLs in Bainong 64 which, in
total, explained 34.1 to 50.4% of the phenotypic variance for
powdery mildew response across three environments (Table 2).
QPm.caas-4DL and QPm.caas-6BS had large effects on resis-
tance to powdery mildew, and the molecular markers Xwmc331
and Xbarc79, closely linked to two QTLs, respectively, could be
used in molecular breeding for APR to powdery mildew in wheat.
Caixia Lan and Shanshan Liang contributed equally to this work. This
study was supported by the National Science Foundation of China
(30810214 and 30671294). We thank R. A. McIntosh, Plant Breeding
Institute, University of Sydney, for a critical review of an earlier draft of
Fig. 2. Logarithm of odds (LOD) contours obtained by composite interval mapping for quantitative trait loci on chromosomes 1A, 4DL, 6BS, and 7A affecting
powdery mildew severity in Bainong 64 × Jingshuang 16 doubled-haploid (DH) lines. 06A = maximum disease severities (MDS) in Anyang, 2006 and 06B and
08B = maximum disease severities in Beijing, 2006 and 2008, respectively. Average = average maximum disease severities across three environments; AUDPC =
average of area under the disease progress curve in Beijing over 2 years. Logarithm of odds (LOD) thresholds, 2.0.
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