Journal of Cereal Science 48 (2008) 789–793

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Journal of Cereal Science

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Combining ability of barley flour pasting properties

M.X. Zhou a, *, H.B. Li a, Z.H. Chen a, N.J. Mendham b
a
Tasmanian Institute of Agricultural Research, University of Tasmania, P.O. Box 46, Kings Meadows, Tasmania 7249, Australia
b
School of Agricultural Science, University of Tasmania, Private Bag 54, Hobart, Tasmania 7001, Australia

a r t i c l e i n f o a b s t r a c t

Article history: Malt extract is one of the major traits contributing to high malting quality and thus a major objective in
Received 3 December 2007 malting barley breeding programs. Understanding the genetic behaviour of this trait could make
Received in revised form 27 May 2008 selection more efficient. However, the measurement of this trait is very time-consuming and cannot be
Accepted 28 May 2008
done in a single plant since it needs a certain amount of grain. In this paper, 10 different varieties with
different malting quality were selected to make two 6  6 half-diallel crosses for the purpose of studying
Keywords:
the genetic behaviour of flour pasting properties, in particular pasting temperature which has been
Barley
shown to be closely related with malt extract. The pasting properties were measured with a Rapid
Pasting property
Combining ability ViscoAnalyser. Both Australian malting varieties showed significantly higher general combining ability
RVA for lower pasting temperature (higher malt extract). The Japanese malting variety also appeared to be
a good parent for lower pasting temperature. All the other feed varieties which are used in the breeding
programs for improving waterlogging tolerance or salinity tolerance showed significantly higher pasting
temperature. Since specific combining ability was not significant, the selection of parents when breeding
for pasting temperature can be based on the pasting temperature of the parents. Combined with the fact
that only a small amount of grain is needed for the measurement of pasting temperature, the selection
can be made in early segregating generations. Preliminary study showed that a major QTL associated
with pasting temperature was located on chromosome 7H, indicating the possibility of marker assisted
selection for this trait.
Ó 2008 Elsevier Ltd. All rights reserved.

1. Introduction To improve selection, researchers are trying to find indirect se-

Malting quality in barley (Hordeum vulgare L.) is one of the major
objectives in breeding programs. It is a combination of a number of
traits, in which malt extract is a key quality indicator because it
reflects the amount of beer that can be produced from a given
quantity of malt (Bamforth and Barclay, 1993). To effectively select
for quality traits, it is important that these traits have high herita-
bility. Unfortunately most of the quality traits showed relatively
lower heritability because grain and malting quality characters are
generally considered to be influenced by many genetic loci, and
highly dependent on environmental factors (Briggs, 1978; Sparrow,
1971). The heritability of malt extract ranged from 8% (based on
a regression of means of F3 families on F2 plants, Foster et al., 1967),
24% (based on a regression of F4 means on F3 lines, Rasmusson and
Glass, 1965), 46% (based on a DH population data from different
sites and years, Han et al., 1997) to 71% (based on the variance
component method in the F3 generation from a diallel cross, Foster
et al., 1967).
* Corresponding author. Fax: þ61 3 6336 5395.
E-mail address: meixue.zhou@dpiw.tas.gov.au (M.X. Zhou).
lection criteria. Oliveira et al. (1994) showed significant correlation
between starch granule characters and malt extract. They also
reported high heritability of starch granule traits based on geno-
type means in a 14 genotype study but the heritability was low in
the parent–offspring heritability evaluation. However, using starch
granule characters as the selection criteria are not so effective in
a breeding program. QTL analysis provides a better understanding
of the genetic factors influencing the target traits. The long-term
goal of QTL analysis is to improve the traits through molecular
marker assisted selection. Many QTLs for malt extract were repor-
ted (Zale et al., 2000) and some molecular markers have already
been used in breeding programs.
Most of the quality traits, including malt extract, are measured
by micromalting. However, in the very early stages of a barley
breeding program, only a small amount of grain is available for
quality assessment, thus it is difficult to conduct micromalting to
evaluate the malt extract. More efficient and feasible approaches
for identifying genotypes with high malt extract are necessary. In
our previous study, we found that some flour pasting properties
measured by the Rapid ViscoAnalyser (RVA) are closely related to
malt extract. Among the different measurements, pasting temper-
ature showed a very close relationship with malt extract (Zhou and

0733-5210/$ – see front matter Ó 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jcs.2008.05.006
790 M.X. Zhou et al. / Journal of Cereal Science 48 (2008) 789–793
Mendham, 2005). The major advantages of using flour pasting
temperature are (1) cheap and fast, and (2) only 5 g of barley grain
is needed, which makes it possible to test single plants. In this
experiment, two different sets of half-diallel crosses were used to
study the genetic behaviour of flour pasting properties in order to
provide some useful information for plant breeders to select for
high malt quality.
2. Materials and methods
2.1. Diallel cross I
A half-diallel cross was made from six selected cultivars, CM72,
Numar (feed varieties from USA), YU6472, Yan90260 (Chinese feed
varieties), Franklin and Gairdner (Australia malting varieties). These
varieties showed great variation in salt tolerance (Chen et al., 2007)
and the tolerant varieties (CM72 and Numar) are being used in
breeding programs.
2.2. Diallel cross II
Another half-diallel cross was made from TX9425, YYXT, DYSYH
(Chinese feed varieties), Naso Nijo (Japanese malting variety),
Franklin and Gairdner. The three Chinese varieties have shown
much better waterlogging tolerance than the other three malting
varieties (Pang et al., 2004, 2006; Zhou et al., 2007).
2.3. Seed production
For both diallel crosses, the six parents and the 15 F1 populations
were grown at Mt Pleasant Laboratories, Tasmania, Australia. Eight
seeds of each parent/F1 were sown in a 1 m row with 35 cm be-
tween rows. Two replications were applied, with randomized
complete blocks. All the parents and F1s were hand harvested,
threshed and cleaned. Seeds were stored in a cool room (<10  C)
before the measurement of pasting properties.
2.4. DH population for preliminary studies of QTLs for pasting
properties
DH lines (176) developed from the cross between Yerong
(Australian feed variety) and Franklin were grown in a birdcage at
Mt Pleasant Laboratories, Tasmania. Around 50 seeds of each DH
line were sown in a 1.5 m row with 40 cm between rows. Both
parents were used as controls. Two replications were applied, with
randomized complete blocks. All the parents and F1s were hand
harvested, threshed and cleaned. Seeds were stored in a cool room
(<10  C) before the measurement of pasting properties. A linkage
map for this population was constructed using 496 DArT and 28
microsatellite markers (Li et al., 2008) and the positions of the
markers were adjusted according to the consensus map (Wenzl
et al., 2006).
2.5. The measurement of flour pasting properties
Samples for RVA analysis were ground on a Cylotech 1903
Sample Mill fitted with a 1 mm screen. Ground barley meal (4.0 g)
was slurried with distilled water or 0.1 M silver nitrate solution
(25.0 g). The pasting properties of the slurries were determined
with a Rapid ViscoAnalyser (RVA-4D, Newport Scientific, Sydney,
NSW, Australia). The test profile has a starting temperature of 50  C,
which is held for 1 min, raised to 95  C in 3.7 min, held for 2.5 min,
cooled to 50  C in 3.8 min, and held for 2 min. Stirring speed is
960 rpm for 10 s (to ensure dispersion of the grist) and 160 rpm for
the remainder of the test period. The RVA measurements were as
follows. Peak viscosity – highest viscosity during heating; time to
peak viscosity; trough – lowest viscosity after cooling started;
breakdown – peak viscosity minus trough; final viscosity – maximum
viscosity after the temperature had returned to 50  C; setback –
final viscosity minus trough; pasting temperature – temperature
when the increase rate reaches 11.5 RVU in 0.2 min, effectively the
beginning of the rapid increase towards PV (Zhou and Mendham,
2005; Zhou et al., 1998).
2.6. Statistical analysis
Parental lines and F1s were subjected to an analysis of variance.
Combining ability effects were analysed according to Griffing
(1956), method 2 (1/2p(p þ 1)) with fixed model.
The software package MapQTL5.0 (Van Ooijen, 2004) was used
for QTL analysis, as described in Li et al. (2008). Logarithm of the
odd (LOD) threshold values applied to declare the presence of a QTL
were estimated by performing the genome wide permutation tests
using at least 1000 permutations of the original data set for each
trait, resulting in a 95% LOD threshold 2.9. The estimated percent-
age of variance of pasting properties explained by the QTL was
obtained using MQM mapping implemented with MapQTL5.0.
MapChart 2.2 was used to draw the chart of genetic linkage and QTL
data.
3. Results
3.1. Pasting properties of the parents
Table 1 shows the pasting properties of the parents used in both
diallel crosses. Significant differences were found for all the RVA
measurements. The malting varieties (Franklin, Gairdner and Naso
Nijo) showed significantly higher peak viscosity and breakdown.
The pasting temperatures of both Franklin and Gairdner were sig-
nificantly lower than all the feed varieties. The pasting temperature
of Naso Nijo was also significantly or meaningfully lower than the
feed varieties. These results are consistent with results reported
earlier (Zhou and Mendham, 2005). As pasting temperature has the
closest correlation with malt extract, further discussion on pasting
properties will focus mainly on pasting temperature.
3.2. Combining ability of pasting temperature
The pasting temperatures of the parents and F1s from both
diallel crosses are listed in Table 2. ANOVA revealed significant
differences between different parents or F1s. Two common varieties
(Franklin and Gairdner) had similar pasting temperatures in both
diallel crosses. ANOVA of combining ability in flour pasting
temperature from both diallel crosses showed that the variance of
general combining ability (GCA) was highly significant (P < 0.01)
while that of specific combining ability (SCA) was not significant
(Table 3). Not surprisingly, both Australian malting varieties had
negative GCA in both diallel crosses. The Japanese malting variety
Naso Nijo also had negative GCA. None of the feed varieties had
negative GCA (Table 4). Similar results were found for other RVA
measurements.
3.3. Identification of QTLs associated with flour pasting properties
Preliminary studies on QTLs controlling flour pasting properties
were conducted with the DH population between the cross of
Franklin and Yerong. In total, seven significant QTLs were found to
be associated with pasting properties, which included two for
trough (2H and 7H), two for breakdown (1H and 4H), two for time
to peak viscosity (2H and 7H) and one for pasting temperature (7H).
No significant QTLs were found for peak viscosity, final viscosity
and setback. The positions of the detected QTLs are shown in Fig. 1.
 M.X. Zhou et al. / Journal of Cereal Science 48 (2008) 789–793 791

Table 1 Table 3
Pasting properties of the selected parents ANOVA of combining ability in flour pasting temperature

Variety Peak viscosity Trough Breakdown Final Setback Peak Pasting Source DF SS MS F
(RVU) (RVU) (RVU) viscosity (RVU) time temperature Diallel cross I
(RVU) (min) ( C) GCA 5 318.10 63.62 38.42
Diallel cross I SCA 15 33.86 2.26 1.36
CM72 385 238 147 400 162 5.97 84.9 Error 20 / 1.66
Numar 392 262 130 437 175 6.13 87.0
Diallel cross II
YU6472 316 172 144 319 147 5.73 87.9
GCA 5 269.30 53.86 20.43
Yan90260 286 147 139 279 132 5.65 88.1
SCA 15 33.73 2.25 0.85
Gairdner 401 198 203 336 139 5.55 75.5
Error 20 / 2.64
Franklin 423 204 219 345 140 5.67 79.1
LSD0.05 10 15 13 17 6 0.13 4.0 F0.05(5,20) ¼ 2.71; F0.01(5,20) ¼ 4.10; F0.05(15,20) ¼ 2.20; F0.01(15,20) ¼ 3.10.
Diallel cross II
TX9425 365 182 184 329 148 5.60 86.2
Naso Nijo 406 188 218 318 130 5.57 79.5 sprouting in barley seed. The RVA measurements of barley starch
Gairdner 447 219 228 371 153 5.70 76.7
were also correlated with amylose content (Yanagisawa et al.,
Franklin 459 239 221 404 165 5.87 75.9
YYXT 383 199 185 343 144 5.80 85.4 2006). Zhou and Mendham (2005) reported that RVA measure-
DYSYH 368 203 165 341 138 5.80 87.1 ments were closely related to malting extract, with pasting
LSD0.05 46 32 18 74 54 0.39 8.3 temperature showing the closest correlation. When pasting tem-
RVU ¼ Rapid Viscoanalyser Units which are approximately equal to centipoise perature is used as an indirect criterion for malt extract, selection
(cP)  10. for malt extract can be on a single plant since only 5 g of grain is
needed. In the current experiment, the flour pasting temperatures
of malting barley varieties were significantly or meaningfully lower
For pasting temperature, the closest marker is bpb-5348 which than those of feed varieties, confirming the relationships between
explained 14.2% of the variation. The malting barley Franklin alleles malting quality and flour pasting temperature.
decreased the pasting temperature, i.e. provided high malting Hockett et al. (1993) found significant GCA and SCA effects in
extract. More detailed study on the QTLs associated with pasting a diallel cross for malt extract with GCA being the predominant
properties will be conducted using data collected from different factor in explaining variation among cross means. The selection of
sites and different DH populations. parent cultivars should therefore just be based on the parental data
of pasting temperature (Baker, 1978), without considering the
4. Discussion specific combining ability. Two Australian and one Japanese
malting variety showed negative GCA (low pasting temperature, i.e.
The assessment of malt extract in a breeding program is time- higher malt extract). All the other feed varieties which are currently
consuming and resource-intensive. Furthermore, in the early stages used in the breeding programs to improve waterlogging or salinity
of the breeding programs, only a small amount of grain is available tolerance showed positive GCA (lower malt extract). For southern
for the assessment. A more efficient and easier method for Australian breeding programs that target the incorporation of
measuring malt quality will be beneficial to plant breeders. waterlogging or salinity tolerance with high malting quality, both
Glennie-Holmes (1995a) described a rapid method of mashing Franklin and Gairdner (low flour pasting temperature and well
micromalts which is suitable for the selection of crossbreds in the adapted to Australian growth conditions) can be used as base
early generations of a barley breeding program. The method he parents for crossing or backcrossing.
developed allows differentiation of samples of similar quality and The relatively lower heritability of malt extract (Foster et al.,
only uses relatively simple equipment. He then developed another 1967; Han et al., 1997; Rasmusson and Glass, 1965) makes it hard to
simple and feasible method based on the hardness of the micro- select for the trait in the early generations. Therefore, indirect
malt, which allows samples of very poor quality to be confidently selection of a related trait with high heritability would increase
rejected (Glennie-Holmes, 1995b). There are very few reports on efficiency in the early generations. Oliveira et al. (1994) found that
the use of a Rapid ViscoAnalyser, which measures the flour pasting some starch granule characters which had relatively high herita-
properties, to select for malting quality. Bueckert et al. (2007) found bilities were correlated with malt extract. However, it is not so easy
that RVA can be used for the early detection of non-visible to use starch granule traits as selection indices in a breeding
program. The measurement of flour pasting temperature which
was closely related to malt extract (Zhou and Mendham, 2005), can
Table 2 be done with only 5 g of barley grains. This not only makes it
Pasting temperatures ( C) of parents and F1s of both diallel sets possible to select for malt extract based on flour pasting tempera-
Diallel cross I ture effectively in early generations, but also reduces plot size for
CM72 Numar Yan90260 YU6472 Gairdner Franklin each line which is desirable for testing the large early generation
CM72 84.7 86.0 87.6 86.4 85.2 83.8 populations (only a small amount of seeds needed for the
Numar 86.0 87.5 86.8 85.1 81.9
measurement).
Yan90260 88.3 87.6 86.7 83.6
YU6472 87.9 83.6 82.3
Gairdner 79.8 78.7 Table 4
Franklin 77.9 GCA of different varieties for pasting temperature
Diallel Cross II
Diallel cross I
TX9425 Naso Nijo Gairdner Franklin YYXT DYSYH
Parents CM72 Numar Yu6472 Yan90260 Gairdner Franklin
TX9425 86.2 83.0 81.0 84.3 85.5 86.7
GCA 1.33 1.62 2.02 2.26 3.64 3.60
Naso Nijo 79.5 76.7 75.4 82.7 85.5
Gairdner 76.7 75.8 81.9 84.3 Diallel cross II
Franklin 75.9 82.3 82.0 Parents TX9425 Naso Nijo Franklin Gairdner YYXT DYSYH
YYXT 85.4 84.3 GCA 2.37 1.48 2.62 2.83 1.69 2.87
DYSYH 87.1
SE ¼ 1.31; t(20,0.05) ¼ 2.09; t(20,0.01) ¼ 2.85.
792 M.X. Zhou et al. / Journal of Cereal Science 48 (2008) 789–793

1H 2H 4H 7H
7.2 bPb-0405 3.5 bPb-4285 12.7 bPb-9304 0.5 bPb-0108
10.4 Bmac0134

21.1 GMS021 27.0 HVM40

36.5 Ebmac0603
45.7 Bmag0872

BD
61.2 bPb-1762
60.7 Bmag0770

TTPV
63.9 Bmac0090

79.8 HVM68 65.9 bPb-4541

72.5 Bmag0518
77.4 Bmac0093

PT
82.6 bPb-5348
87.4 Bmag0341
T
BD

TTPV

106.2 bPb-4515
108.7 HVHVA1 113.5 Ebmac0679
108.2 bPb-1772

129.2 HVM54

T
133.4 bPb-3484
148.6 bPb-3717
147.3 bPb-3756

161.9 bPb-5015 159.2 bPb-0259

Fig. 1. Quantitative trait loci (QTLs) identified for pasting properties in the doubled haploid population produced from the cross between Franklin and Yerong. Only SSR markers and
important DArT markers (both ends and markers linked with the QTLs) were presented in the map. T: trough; BD: breakdown; TTPV: time to peak viscosity; PT: pasting
temperature.

The climatic conditions at or prior to the time of harvest can
significantly affect quality of barley grains. For example, weather-
damaged barley showed a totally different pattern of pasting
property, which is similar to that of malted barley (Zhou and
Mendham, 2005). The use of 0.1 M silver nitrate can inactivate
several different enzymes during the running of RVA and avoid the
effect of possible weather damage on flour pasting properties (Zhou
and Mendham, 2005). Pasting properties can also be affected by soil
and other environmental conditions. The use of molecular marker
assisted selection can greatly reduce the environmental effect and
increase selection efficiency for malt quality. QTL analysis on malt
extract or related traits not only provides a better understanding of
the genetic factors influencing the trait but also helps to identify
chromosome regions and molecular markers linked to this trait.
Many QTLs associated with malt extract have been reported (as
summarised by Zale et al., 2000), covering nearly all the different
chromosomes. Panozzo et al. (2007) reported QTLs on chromo-
somes 1H and 7H with significant positive effects arising from
Franklin. In our preliminary QTL analysis, seven significant QTLs
were found for different pasting properties. Among them, three
QTLs were found on 7H including one QTL for pasting temperature
with negative contribution (positive effects for malt extract) from
Franklin. This QTL explained 14.2% of the variation in pasting
temperature. It is not clear whether this QTL is at the same position
as that reported by Panozzo et al. (2007) since they did not show
the position in their paper. QTLs identified from just one environ-
ment in this experiment may not be accurate enough since, as
mentioned above, quality traits are easily affected by various
environmental factors. As a follow on study, more detailed QTL
analysis for different flour pasting properties will be done with data
from different locations (years) and different DH populations.
In conclusion, general combining ability for pasting temperature
was highly significant and no significant specific combining ability
was detected. The selection of parents for improving pasting
temperature in a breeding program can be based on the parental
data. Since only a small amount of grain is needed for the
measurement of pasting temperature, the selection can be made in
early segregating generations. Three of seven QTLs for pasting
properties were on 7H, showing the promise of 7H for flour pasting
property related traits. However, as pasting properties are very
complicated traits and can easily be affected by various environ-
mental factors, further QTL studies are needed by using data from
different locations and years.
Acknowledgment
We thank the Australian Grain Research and Development
Corporation (GRDC) for funding the project ‘‘Australia China
Collaboration on Barley Genetic Resources’’ (Project UT8).
References
Baker, R.J., 1978. Issues in diallel analysis. Crop Science 18, 533–536.
Bamforth, C.W., Barclay, A.H.P., 1993. Malting technology and the use of malt. In:
MacGregor, A.W., Bhatty, R.S. (Eds.), Barley Chemistry and Technology.
American Association of Cereal Chemists, St. Paul, MN, pp. 297–354.
Briggs, D.E., 1978. Barley. Chapman and Hall, London.
 M.X. Zhou et al. / Journal of Cereal Science 48 (2008) 789–793
Bueckert, R.A., Lefol, E.B., Harvey, B.L., 2007. Early detection of non-visible sprouting
in barley seed using rapid viscosity analysis. Canadian Journal of Plant Science
87, 3–12.
Chen, Z.H., Zhou, M.X., Newman, I., Mendham, N., Zhang, G.P., Shabala, S., 2007.
Potassium and sodium relations in salinised barley tissues as a basis of differ-
ential salt tolerance. Functional Plant Biology 34, 150–162.
Foster, A.E., Peterson, G.A., Banasik, O.J., 1967. Heritability of factors affecting
malting quality of barley. Crop Science 7, 611–613.
Glennie-Holmes, M., 1995a. Studies on barley and malt with the Rapid ViscoAna-
lyser: [I]. The effect of variations in physical and chemical parameters. Journal of
the Institute of Brewing 101, 11–18.
Glennie-Holmes, M., 1995b. Studies on barley and malt with the Rapid ViscoAna-
lyser: [II]. The effect of modification on viscograms. Journal of the Institute of
Brewing 101, 19–28.
Griffing, B., 1956. Concept of general and specific combining ability in relation to
diallel crossing systems. Australian Journal of Biological Science 9, 463–493.
Han, F., Romagosa, I., Ullrich, S.E., Jones, B.L., Hayes, P.M., Wesenberg, D.M., 1997.
Molecular marker-assisted selection for malting quality traits in barley.
Molecular Breeding 3, 427–437.
Hockett, E.A., Cook, A.F., Khan, M.A., Martin, J.M., Jones, B.L., 1993. Hybrid perfor-
mance and combining ability for yield and malt quality in a diallel cross of
barley. Crop Science 33, 1239–1244.
Li, H.B., Vaillancourt, R., Mendham, N.J., Zhou, M.X., 2008. Comparative mapping of
quantitative trait loci associated with waterlogging tolerance in barley (Hor-
deum vulgare L.). BMC Genomics 9, 401.
Oliveira, A.B., Rasmusson, D.C., Fulcher, R.G., 1994. Genetic aspects of starch granule
traits in barley. Crop Science 34, 1176–1180.
Pang, J.Y., Zhou, M.X., Mendham, N., Shabala, S., 2004. Growth and physiological
responses of six barley genotypes to waterlogging and subsequent recovery.
Australian Journal of Agricultural Research 55, 895–906.
793
Pang, J.Y., Newman, I.A., Mendham, N., Zhou, M.X., Shabala, S., 2006. Microelectrode
ion and O2 fluxes measurements reveal differential sensitivity of barley root
tissues to hypoxia. Plant, Cell and Environment 29, 1107–1121.
Panozzo, J.F., Eckermann, P.J., Mather, D.E., Moody, D.B., Black, C.K., Collins, H.M.,
Barr, A.R., Lim, P., Cullis, B.R., 2007. QTL analysis of malting quality traits in two
barley populations. Australian Journal of Agricultural Research 58, 858–866.
Rasmusson, D.C., Glass, R.L., 1965. Effectiveness of early generation selection for four
quality characters in barley. Crop Science 5, 389–391.
Sparrow, D.H.B., 1971. Some genetical aspects of malting quality. In: Nilan, R.A.
(Ed.), Barley Genetics II. Washington State University Press, Pullman, pp.
559–574.
Van Ooijen, J.W., 2004. MapQTLÒ 5, Software for the Mapping of Quantitative Trait
Loci in Experimental Populations. Kyazma B.V., Wageningen, The Netherlands.
Wenzl, P., Li, H.B., Carling, J., Zhou, M.X., Raman, H., Paul, E., Hearnden, P., Maier, C.,
Xia, L., Caig, V., Ovesná, J., Cakir, M., Poulsen, D., Wang, J.P., Raman, R., Smith, K.,
Muehlbauer, G., Chalmers, K., Kleinhofs, A., Huttner, E., Kilian, A., 2006. A high-
density consensus map of barley linking DArT markers to SSR, RFLP and STS loci
and agricultural traits. BMC Genomics 7, 206.
Yanagisawa, T., Domon, E., Fujita, M., Kiribuchi-Otobe, C., Takayama, T., 2006. Starch
pasting properties and amylose content from 17 waxy barley lines. Cereal
Chemistry 83, 354–357.
Zale, J.M., Clancy, J.A., Ullrich, S.E., Jones, B.L., Hayes, P.M., 2000. Summary of barley
malting quality QTLs mapped in various populations. Barley Genetics News-
letter 30, 44–54.
Zhou, M.X., Mendham, N.J., 2005. Predicting barley malt extract with a Rapid Vis-
coAnalyser. Journal of Cereal Science 41, 31–36.
Zhou, M.X., Robards, K., Glennie-Holmes, M., Helliwell, S., 1998. Structure and
pasting properties of oat starch. Cereal Chemistry 75, 273–281.
Zhou, M.X., Li, H.B., Mendham, N.J., 2007. Combining ability of waterlogging toler-
ance in barley. Crop Science 47, 278–284.