Published online 30 July 2007
Published in Crop Sci 47:1754-1760 (2007)
© 2007 Crop Science Society of America
677 S. Segoe Rd., Madison, WI 53711 USA
PLANT GENETIC RESOURCES
Barley Amylose and ß-Glucan: Their Relationships to Protein, Agronomic Traits, and Environmental Factors
An Hanga,*,
Don Oberta,
Ann Inez N. Gironellab and
Charlotte S. Burtona
a USDA-ARS, Small Grains and Potato Germplasm Research Unit, 1691 S. 2700 W., Aberdeen, ID 83210
b Mathematics Dep., Idaho State Univ., Pocatello, ID 83209. Mention of trade names or commercial products in this article is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the USDA
* Corresponding author (anhang{at}uidaho.edu).
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ABSTRACT
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Amylose and ß-glucan are important components of barley (Hordeum vulgare L.) grain. Factors such as seed yield, test weight, seed plumpness, protein content, field location, and seasonal variation may have an effect on ß-glucan and amylose concentrations, and therefore, require additional study. Twenty-seven barley cultivars and advanced lines with varying levels of ß-glucan (4–9%) and amylose (6–27%) were grown in 2004 and 2005 in replicated plots at three locations in Idaho to evaluate agronomic traits and genotype x environment interactions. Seed yield, test weight, percentage of plump kernels, ß-glucan percentage, amylose/amylopectin ratio, and protein percentage were measured for seed samples from all plots. Estimates of variance components showed that about 49% of the variability in ß-glucan concentration and about 48% in amylose starch levels can be attributed to year, location, year x location, and their interactions with genotype. Amylose was found to be negatively correlated with protein and ß-glucan content but was positively, though weakly, correlated with seed yield and test weight; ß-glucan was positively correlated with protein and percentage plump kernels but showed a weak negative correlation with seed yield.
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INTRODUCTION
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THE QUALITY OF malting, food, and feed barleys (Hordeum vulgare L.) depends on ß-glucan and amylose starch concentrations in the grain. Amylose is a major component of the starch in the barley kernel, usually ranging from 22 to 26%, with 74 to 78% of the kernel composed of amylopectin (Newman and Newman, 1992). High-amylose barley has amylose content >35% of the total starch, while low-amylose or waxy type barley consists of 0 to 10% amylose or 90 to 100% amylopectin (Washington et al., 2000). An extremely low amylose genotype has been found from a mutation in the waxy gene (Patron et al., 2002), and a cultivar with 0% amylose has been reported (Bhatty and Rossnagel, 1992). Conversely, a barley cultivar with amylose exceeding 40% has also been reported (Merritt, 1967). Amylose concentrations in barley are controlled by amylose (amo1) and waxy (wax) genes (Swanston et al., 1995). The amo1 gene is located on chromosome 1HS (5S) (Schondelmaier et al., 1992) and the wax gene is located on chromosome 7HS (1S) (Lundqvist et al., 1997). The interaction between the two genes results in different levels of amylose in grain.
Amylose content may affect the digestibility of feed barley in livestock animals. High-amylose starch is less susceptible to 
-amylase than normal barley starch and forms a stable complex with lipids, which is relatively resistant to enzymatic degradation (Pomeranz et al., 1972; Newman and Newman, 1992). High-amylose barley rations fed to rats resulted in significantly improved bowel health factors (Bird et al., 2004). Foods made from high-amylose grains have also been shown to lower blood glucose, insulin levels, triglycerides, and cholesterol in animals and humans (Behall et al., 1989; Delaney et al., 2003).
Mixed-linkage (1
3) (14)-ß-glucans are major structural components of barley endosperm and aleurone cell walls, comprising 75% of endosperm (Fincher, 1975) and 26% of the aleurone cell walls (Bacic and Stone, 1981). High ß-glucan content is undesirable for malting as it is associated with lowered malt extract values, viscous extracts that lead to filtration difficulties in the brewing process (Ullrich et al., 1986; Fincher, 1992), and the potential production of precipitates in beer (Woodward and Fincher, 1983). In certain animals, ß-glucan is associated with reduced feed efficiency (MacGregor and Fincher, 1993; White, 2000). For food and some feed uses, however, ß-glucan is desirable due to its cholesterol-lowering properties (Oakenfull, 1996). ß-glucan and other soluble fibers are believed to bind serum cholesterol and decrease its absorption, resulting in lower total plasma cholesterol levels in humans and animals, thus reducing the risk of cardiovascular disease (Newman and Newman, 1992; White, 2000).
Genetic studies of ß-glucan in populations derived from doubled haploid (DH) and single seed descent lines from various crosses (Powell et al., 1985) found that ß-glucan in barley is controlled by a simple additive genetic system. Using DH lines derived from the cross ‘Steptoe’/‘Morex’, Han et al. (1995) studied quantitative trait loci (QTL) of ß-glucan content in barley grain and malt. They found that the QTL with the largest effects on barley and malt ß-glucan were located on chromosomes 2H (2), 7H (1), 4H (4), and 5H (7). High ß-glucan is associated with the waxy trait in barley (Newman and Newman, 1992). Genetic and environmental effects on ß-glucan levels in barley grains have been reported (Aastrup, 1979; Powell et al., 1985; Hockett et al., 1987; Fastnaught et al., 1996; Zhang et al., 2001) but limited information is available on the relationship of quality and agronomic traits, environmental factors, and their interactions on amylose and ß-glucan contents. Our objectives were to study the effects of location, year, and their interactions on amylose, ß-glucan, and protein content in various barley genotypes and to determine how they are related to yield, test weight, and percentage of plump kernels.
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MATERIALS AND METHODS
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Plant Material
Twenty-seven hulless and hulled barley genotypes, including cultivars and advanced lines with ß-glucan contents averaging from 4 to >9% and amylose concentrations ranging from 6 to 27%, were obtained from the USDA-ARS National Small Grains Germplasm Research Facility at Aberdeen, ID (Table 1). ‘Baronesse’ is a two-rowed, hulled feed barley cultivar derived from a complex cross. It was developed by the company Nordsaat in Germany and marketed in the USA by WestBred, LLC, Bozeman, MT. It is currently widely grown in Idaho and Montana as feed barley. ‘Azhul’ is a six-rowed, hulless high ß-glucan germplasm released by the USDA-ARS and is the source of high ß-glucan in many current cultivars. ‘CDC Alamo’ is a high ß-glucan, two-rowed, hulless cultivar released by the University of Saskatchewan, and ‘Waxbar’ is a waxy, two-rowed, hulless barley marketed by WestBred, LLC, Bozeman, MT. These cultivars were used as checks.
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Table 1. Barley (Hordeum vulgare L.) genotypes tested for amylose, ß-glucan, protein content, and other agronomic and quality traits in barley.
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Field Trials
Experimental units consisted of seven-row plots, 3.05 by 1.52 m, planted in a randomized complete block design with three replications at each of three locations in Idaho in 2004 and 2005. Plots at Aberdeen and Filer were irrigated, while those at Soda Springs were grown under dryland conditions. These locations vary greatly in elevation and length of growing season. Filer (elevation 1138 m) has the longest growing season while Soda Springs (elevation 1775 m) has the shortest. Aberdeen (elevation 1327 m) is between the two other locations, but is more similar to Filer. Plots were planted at Aberdeen in both 2004 and 2005 at the University of Idaho's Aberdeen Research and Extension Center, and followed a crop of oat (Avena sativa L.) that was harvested for forage. Plots at Filer followed sugarbeet (Beta vulgaris L.) in 2004 and pea (Pisum sativa L.) in 2005. In both 2004 and 2005, the plots at Soda Springs followed malting barley. Plots at all locations were fertilized as recommended for malt barley production. The seeding rate was 105 kg ha–1 at Aberdeen and Filer and 79 kg ha–1 at Soda Springs. Seed yield expressed as kilograms per hectare was adjusted to 110 g kg–1 moisture.
Test Weight and Plump Kernel Analysis
Samples were cleaned and test weight was determined. A 100-g sample was then screened and kernels retained on a sieve with 0.24- by 1.9-cm slotted openings (American Society of Brewing Chemists, 1992) were considered plump.
Chemical Analyses
For amylose and ß-glucan determinations, 5 g of seed was ground in a UDY Cylone sample mill with a 0.5-mm mesh screen and concentrations were determined by using enzyme-specific amylose/amylopectin and mixed-linkage ß-glucan detection assay kits from Megazyme (Megazyme International Ireland Ltd., Wicklow, Ireland). For protein measurement, 40 g of seed was ground with a 1-mm mesh screen and analyzed using a Perten 8611 near-infrared reflectance spectroscopy analyzer (Perten Instruments, Huddinge, Sweden) as described in American Association of Cereal Chemists (1983, Method 39-10).
Statistical Analysis
A combined analysis of variance was done across locations and years using a mixed effects linear model where genotype was fixed and all other effects were considered random. The model is described as:
where Yijkl is the observed response of genotype i (i = 1, 2, ..., 27) in replication j (j = 1, 2, 3) nested in location l (l = 1, 2, 3) of year k (k = 1, 2), µ is the overall mean response, Gi is the effect of the genotype i, Rj(kl) is the effect of replicate j in year k and location l, Yk is the effect of year k, Ll is the effect of location l, (YL)kl, (GY)ik, and (GL)il are the first-order interaction effects, and (GYL)ikl is the second-order interaction effect. The following components of variance: 
R2, Y2, L2, YL2, GY2, GL2, GYL2and Error2 for replication, year, location, year x location, genotype x year, genotype x location, genotype x year x location, and error, respectively, were estimated using the restricted maximum likelihood procedure for the mixed effects model. To correct for nonnormality of residual distributions, all statistical analyses were done on transformed values of amylose, ß-glucan, and plumpness. For amylose, the square root transformation was used; for ß-glucan, the natural logarithm; and for plumpness, the square transformation. The percentage (p) of total variation in an observed response that is due to environmental factors was estimated as follows. Let VTotal = estimated total variation in an observed response = 
R2+Y2+L2+YL2+GY2+GL2+GYL2+2Error and VEnv = estimated variation in an observed response due to environmental factors, that is, year, location, year x location, and their interactions with genotype = Y2+L2+YL2+GY2+GL2+GYL2. Then p = (VEnv/VTotal)100%. Genotype least squares means were estimated and compared using the Tukey–Kramer paired comparisons test. Simple correlation analysis was done to determine the relationships among amylose and ß-glucan, protein, test weight, yield, and percentage of plump kernels. Partial correlation analysis was performed to examine the relationships among these six characteristics when the effects of genotype and all environmental factors were held constant or removed. There were a few missing observations and detectable outliers. Of 486 observations on each response variable, three were missing for amylose, nine for protein, five for seed yield, three for test weight, and two for plumpness. Two observations on test weight were declared outliers based on the scatterplot of the data and on residual analysis. All statistical analyses were performed using the univariate and mixed procedures of SAS (Version 9.1, SAS Institute, Cary, NC).
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RESULTS AND DISCUSSION
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Amylose and ß-glucan means for the 27 genotypes showed significant differences (P < 0.0001), as expected, since they were preselected from cultivars expressing a range of these traits. Mean values for amylose varied from 5.61 to 26.33%, with Genotype 00AH4495 showing the highest mean (Table 2). For ß-glucan, means ranged from 4.20 to 8.94%, with the highest mean recorded for cv. Azhul. Protein ranged from 10.19 to 14.49%, with Genotype 00AH3778 showing the highest mean. Means for other quality traits and agronomic performance are presented in Table 2.
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Table 2. Means of amylose starch, ß-glucan, protein, seed yield, test weight, and seed plumpness of 27 barley genotypes grown at three locations in 2004 and 2005.
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Estimates of the different variance components are shown in Table 3. Of the environmental factors, the components for year and genotype x year x location for amylose were largest, indicating that amylose content of a cultivar or line may vary depending on year and location of planting. An estimated 47.9% of the total variation in amylose content was accounted for by environmental factors and their interactions with genotype. For ß-glucan content, the environmental factor that had the largest component was year x location, indicating that the ß-glucan content of a genotype may vary depending on year and location of planting. An estimated 48.7% of the total variability in ß-glucan content was accounted for by the environmental factors and their interactions with genotype. Estimates of the different variance components for the other quality and agronomic characteristics are presented in Table 3. Simple correlations (Table 4), which included the effects of genotype and environmental factors, showed that amylose was negatively correlated with ß-glucan and protein, but ß-glucan was positively correlated with protein. Amylose was also found to be significantly and positively correlated with seed yield and test weight, and ß-glucan was negatively correlated with seed yield, but the correlations were very low. ß-glucan and percentage of plump kernels were found to be positively correlated. Partial correlation analysis showed that when genotype and environmental factors were held constant or their effects removed, no significant correlation was detected among amylose, ß-glucan, and protein content (Table 5), indicating that the simple correlations found can be attributed to differences in how genotypes respond to varying environmental conditions. Significant partial correlations were also found between protein and test weight, between test weight and yield, between yield and percentage of plump kernels, and between test weight and percentage of plump kernels, but they were too low to be of practical importance.
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Table 3. Estimates of variance components for traits measured in 27 barley genotypes varying for amylose and ß-glucan content grown at three locations in 2004 and 2005.
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Table 4. Pearson's simple correlation coefficients, P values, and sample sizes for traits measured in 27 barley genotypes varying for amylose and ß-glucan content grown at three locations in 2004 and 2005.
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Table 5. Partial correlation coefficients and P values (df = 297) for traits measured in 27 barley genotypes varying for amylose and ß-glucan content grown at three locations in 2004 and 2005.
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The growth of barley for the purpose of extracting ß-glucan has recently become a reality, with premium prices being paid for the production of Azhul (D. Obert, personal communication, 2006). The most desirable barley for human food use would have high protein, high ß-glucan, high amylose, high test weight, high percentage of plump kernels, and high seed yield. Our results, along with those of previous studies, suggest that the effort to develop cultivars with these traits will be difficult due to the high negative correlation between ß-glucan and amylose content. Correlations between ß-glucan and seed yield, and ß-glucan and test weight are both negative. Selection for high ß-glucan will require simultaneous selection for segregants with suitable yield and test weight, but since these correlations are very low, this is not seen as a major problem in the development of high ß-glucan barley cultivars. The positive correlations between ß-glucan and protein and ß-glucan and seed plumpness are both advantageous to breeders as each of these traits are desirable for all end uses involving grain consumption. The development of barley for human consumption may need to be focused on breeding cultivars for specific end uses. For example, due to the high negative correlation between ß-glucan and amylose, barley cultivars developed specifically for ß-glucan extraction will have lower levels of amylose. High-amylose cultivars developed for human consumption will have lower to moderate levels of ß-glucan. Thus a cultivar that meets the needs of all market classes, specifically a high ß-glucan, high amylose type, may not be feasible.
Barley for animal consumption would not require the high levels of ß-glucan desired for human consumption, but all other factors would be similar. The development of barley cultivars with low or moderate levels of ß-glucan would be negatively impacted by the correlation between amylose content and protein, and to a lesser extent by seed plumpness. The positive correlations of amylose content with seed yield and test weight are desirable for achieving breeding progress. Breeding efforts to develop hulless barleys with varying levels of both ß-glucan and amylose are underway.
Seed yields of hulless barley tended to be lower than hulled, due in part to the lack of hull. In this study, Baronesse, the hulled check, had higher yield than all hulless lines. The highest yielding hulless line was 00AH4495, which yielded 85% of Baronesse, while Azhul had the lowest yield, with 58% of Baronesse.
Eslick and Ullrich (1977) compared means of protein and other agronomic traits of the barley cultivar Glacier and an isogenic line with high amylose grown at 21 locations in Montana and Arizona. While protein in the high-amylose isoline was significantly higher than in Glacier, yield, test weight, kernel weight, and percentage of plump kernels values were significantly higher in Glacier than in the high-amylose isoline. Kernel weight was only slightly greater in Glacier than in the high-amylose isoline. In our study, protein was negatively correlated with amylose. The difference in results may be attributed to the lower amylose content of genotypes studied compared with that of the high-amylose (44%) isoline.
Ullrich et al. (1986) compared ß-glucan in low-amylose or waxy genotypes with their isogenic normal genotype. They found that total ß-glucan content was consistently higher in low-amylose or waxy barley than in the normal isotypes. This study also found a negative correlation between amylose and ß-glucan for genotypes across environments. The waxy gene was associated with an increase in barley ß-glucan in another study by Newman and Newman (1992). Hockett et al. (1987) grew barley cultivars in eight Montana environments and found that ß-glucan content was positively correlated to protein content and yield in 1 yr but not significantly correlated in another year. Our data showed that simple correlation between ß-glucan and protein content was significantly positive across environments. Similar results were also found in studies with oat (Peterson et al., 1995). Hockett et al. (1987) indicated that test weight and percentage of plump kernels had positive correlations with ß-glucan in 1983, but were not significant in 1982. Our data showed significant simple correlations between ß-glucan and yield that were negative but positive between ß-glucan and percentage of plump kernels, although these correlations were low.
For the determination of amylose and ß-glucan contents, we used enzyme-specific assay kits from Megazyme, which may explain the differences in amylose from those previously reported. For example, we found that the average amylose content of CDC Alamo was about 6.4%. Washington et al. (2000) evaluated amylose in this cultivar by using both an iodine method and Megazyme assay. Amylose was detected at 4 to 5% for the Megazyme assay and at 6% for the iodine method; Bhatty and Rossnagel (1992) used a modified method from Holm et al. (1986) and reported 0% amylose in CDC Alamo. Accurate determination of quality traits such as amylose/amylopectin, ß-glucan, and protein is difficult and has varied between laboratories. Genotypes with high amylose, however, have been consistently associated with low ß-glucan and vice versa. These results were also observed by Ullrich et al. (1986). The quality traits of starch, protein, and ß-glucan are useful for feed, food, and industry and vary depending on genotype and field environments. Our study showed that genotype x environment interactions affect these characteristics. Additional research on the effects of quality and agronomic traits and their interactions with genotypes would be useful and will possibly improve nutritional values in barley grain.
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ACKNOWLEDGMENTS
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We acknowledge Kathy Satterfield and Chris Evans for technical assistance, and Katherine O'Brien for seed protein measurements.
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NOTES
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All rights reserved. No part of this periodical may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permission for printing and for reprinting the material contained herein has been obtained by the publisher.
Received for publication June 27, 2006.
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REFERENCES
|
|---|
- Aastrup, S. 1979. The effect of rain on ß-glucan content in barley grains. Carlsberg Res. Commun. 44:381–393.[Web of Science]
- American Association of Cereal Chemists. 1983. Approved methods of American Association of Cereal Chemists. 8th ed. AACC, St. Paul, MN.
- American Society of Brewing Chemists. 1992. Methods of analysis. 8th ed. ASBC, St. Paul, MN.
- Bacic, A., and B.A. Stone. 1981. Isolation and ultrastructure of aleurone cell walls from wheat and barley. Aust. J. Plant Physiol. 8:453–495.[Web of Science]
- Behall, K.M., D.J. Scholfield, I. Yuhauiak, and J. Canary. 1989. Diets containing high amylose vs amylopectin starch: Effects on metabolic variables in human subjects. Am. J. Clin. Nutr. 49:337–344.[Abstract/Free Full Text]
- Bhatty, R.S., and B.G. Rossnagel. 1992. Zero amylose lines of hull-less barley. Cereal Chem. 74:190–191.[CrossRef]
- Bird, A.R., C. Flory, D.A. Davies, S. Usher, and D.L. Yopping. 2004. A novel barley cultivar (Himalaya 292) with a specific gene mutation in starch synthase IIa raises large bowel starch and short-chain fatty acids in rats. J. Nutr. 134:831–835.[Abstract/Free Full Text]
- Delaney, B., R.J. Nicolosi, T.A. Wilson, T. Carlson, S. Frazer, G.H. Zheng, R. Hess, K. Ostergren, J. Haworth, and N. Knutson. 2003. ß-glucan fractions from barley and oat are similarly antiatherogenic in hypercholesterolemic Syrian golden hamsters. J. Nutr. 133:468–475.[Abstract/Free Full Text]
- Eslick, R.F., and S.E. Ullrich. 1977. A characterization of three high lysine mutants in relation to their normal isotypes and their environment response. Barley Newsl. 20:39–45.
- Fastnaught, C.E., P.T. Berglund, E.T. Holm, and G.J. Fox. 1996. Genetic and environmental variation in ß-glucan content and quality parameters of barley for food. Crop Sci. 36:941–946.[Abstract/Free Full Text]
- Fincher, G.B. 1975. Morphology and chemical composition of barley endosperm cell walls. J. Inst. Brew. 81:116–122.
- Fincher, G.B. 1992. Cell wall metabolism in barley. p. 413–437. In P.R. Shewry (ed.) Barley: Genetics, biochemistry, molecular biology, and biotechnology. CAB Int., Wallingford, UK.
- Han, F., S.E. Ullrich, S. Chirat, S. Menteur, L. Jestin, A. Sarrafi, P.M. Hayes, B.L. Jones, T.K. Blake, D.M. Wesenberg, A. Kleinhofs, and A. Kilian. 1995. Mapping of ß-glucan content and ß-glucanase activity loci in barley grain and malt. Theor. Appl. Genet. 91:921–927.[Web of Science]
- Hockett, E.A., C.F. McGuire, C.W. Newman, and N. Prentice. 1987. The relationship of barley beta-glucan content to agronomic and quality characteristics. p. 851–860. In S. Yasuda and T. Konishi (ed.) Barley genetics V. Proc. Int. Barley Genet. Symp., 5th, Okayama, Japan. 6–11 Oct. 1986. Sanyo Press, Okayama, Japan.
- Holm, J., F. Bjorek, A. Drews, and N.G. Asp. 1986. A rapid method for the analysis of starch. Starch/Staerke 38:224–226.[CrossRef]
- Lundqvist, U., J.D. Franckowiak, and T. Konishi. 1997. New and revised descriptions of barley genes. Barley Genet. Newsl. 26:22–516.
- MacGregor, A.W., and G.B. Fincher. 1993. Carbohydrates of the barley grain. p. 74–130. In A.W. MacGregor and R.S. Bhatty (ed.) Barley: Chemistry and technology. Am. Assoc. Cereal Chemists, St. Paul, MN.
- Merritt, N.R. 1967. A new strain of barley with starch of high amylose content. J. Inst. Brew. 73:583–585.
- Newman, C.W., and R.K. Newman. 1992. Nutritional aspects of barley seed structure and composition. p. 351–368. In P.R. Shewry (ed.) Barley: Genetics, biochemistry, molecular biology, and biotechnology. CAB Int., Wallingford, UK.
- Oakenfull, D. 1996. Food applications for barley. p. 50–57. In G. Scoles and B. Rossnagel (ed.) Proc. Int. Oat Conf. and Int. Barl. Genet. Symp., Saskatoon, SK. 30 July–6 Aug. 1996. Univ. of Saskatchewan Press, Saskatoon.
- Patron, N.J., A.M. Smith, B.F. Fahy, C.M. Hylton, M.J. Naldrett, B.G. Rossnagel, and K. Denyer. 2002. The altered pattern of amylose accumulation in the endosperm of low-amylose barley cultivars is attributable to a single mutant allele of granule-bound starch synthase I with a deletion in the 5'-non-coding region. Plant Physiol. 130:190–198.[Abstract/Free Full Text]
- Peterson, D.M., D.M. Wesenberg, and D.E. Burrup. 1995. ß-glucan content and its relationship to agronomic characteristics in elite oat germplasm. Crop Sci. 35:965–970.[Abstract/Free Full Text]
- Pomeranz, Y., R.F. Eslick, and G.S. Robbins. 1972. Amino acid composition and malting and brewing performance of high amylose and hiproly barleys. Cereal Chem. 49:629–635.
- Powell, W., P.D.S. Caligari, J.S. Swanston, and J.L. Jinks. 1985. Genetical investigations into ß-glucan content in barley. Theor. Appl. Genet. 71:461–466.[Web of Science]
- Schondelmaier, J., A. Jacobi, G. Fischbeck, and A. Jahoor. 1992. Genetical studies on the mode of inheritance and localization of the amo1 (high amylose) gene in barley. Plant Breed. 109:274–280.[CrossRef]
- Swanston, J.S., R.P. Ellis, and J.R. Stark. 1995. Effects on grain and malting quality of genes altering barley starch composition. J. Cereal Sci. 22:265–273.[CrossRef]
- Ullrich, S.E., J.A. Clancy, R.F. Eslick, and R.C.M. Lance. 1986. ß-glucan content and viscosity of extracts from waxy barley. J. Cereal Sci. 4:279–285.
- Washington, J.M., A. Box, and A.R. Barr. 2000. Developing waxy barley cultivars for food, feed and malt. p. 303–306. In S. Logue (ed.) Barley genetics VIII. Int. Barley Genet. Symp., 8th, Adelaide, Australia. 22–27 Oct. 2000. Univ. of Adelaide, SA, Australia.
- White, P.J. 2000. Dietary fiber. p. 201–213. In C.F. Murphy and D.M. Peterson (ed.) Designing crops for added value. Agron. Monogr. 40. ASA, CSSA, and SSSA, Madison, WI.
- Woodward, J.R., and G.B. Fincher. 1983. Water-soluble barley ß-glucans. Fine structure, solution behaviour and organization in the cell wall. Brew. Dig. 58:28–32.
- Zhang, G., J. Chen, J. Wang, and S. Ding. 2001. Cultivar and environmental effects on (13, 14)-ß-D-glucan and protein contents in malting barley. J. Cereal Sci. 34:295–301.[CrossRef]
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TOP
ABSTRACT
INTRODUCTION
