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CROP BREEDING & GENETICS

Small-Scale Bread-Quality-Test Performance Heritability in Bread Wheat: Influence of High Molecular Weight Glutenin Subunits and the 1BL.1RS Translocation

^a National Agricultural Research Center for Hokkaido Region (Memuro), Shinsei, Memuro, Hokkaido 082-0071, Japan
^b National Agricultural Research Center for Western Region, 6-12-1 Nishi-Fukatsucho, Fukuyama, Hiroshima 721-8514, Japan
^c National Agricultural Research Center for the Hokkaido Region (Sapporo), 1 Hitsujigaoka, Toyohira, Sapporo, Hokkaido 062-8555, Japan

^* Corresponding author (zenta{at}affrc.go.jp).

	ABSTRACT

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Small-scale bread-quality assays (grain protein content, SDS-sedimentation volume [SDSS], and single-kernel characterization system [SKCS] grain hardness) represent important tools in bread wheat (Triticum aestivum L.) breeding. The influence of high-molecular-weight glutenin (HMWG) subunits and the 1BL.1RS translocation on small-scale bread-quality assays and their heritability was investigated for F₃ and F₄ lines of two wheat populations. Lines with HMWG subunits 5+10 at Glu-D1 showed significantly greater SDSS volume and SKCS hardness than those with subunits 2+12 or 4+12. Lines with HMWG subunit 20 at Glu-B1 had a significantly lower SDSS volume than those with HMWG subunits 7+8 or 7+9. F₄ lines bearing the 1BL.1RS translocation had significantly greater protein content. The distribution of SDSS volumes suggested that most bore the undesirable HMWG subunits 2+12, 4+12 or 20, but could be selected against if the SDSS volume <6.0 mL. For SDSS volume, lines with HMWG subunits 5+10 had greater heritability in upward screening, whereas those with HMWG subunits 2+12 or 4+12 had greater heritability in downward screening. The above screening criterion provides an excellent tool for bread quality-based wheat breeding, irrespective of hard x hard or hard x soft wheat crosses.

Abbreviations: FPS, flour particle size • HMWG, high-molecular-weight glutenin • SDS-PAGE, sodium dodecylsulfate polyacrylamide gel electrophoresis • SDSS, sodium dodecylsulfate sedimentation • SKCS, single kernel characterization system

	INTRODUCTION

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IN A WHEAT (Triticum aestivum L.) breeding program targeting bread quality, small-scale quality tests are indispensable in screening early-generation plant materials due to the limitation of sample size. Screening for higher grain protein content is essential in increasing gluten content, which in turn improves loaf volume. To estimate gluten quality, small-scale quality tests that measure swelling power and solubility have been developed: Zeleny test (Zeleny, 1947), sedimentation test (Pinckney et al., 1957), and SDS-sedimentation (SDSS) volume test (Axford et al., 1979). Original or modified SDSS volume tests have been widely employed to predict bread quality in wheat breeding programs (Silvera et al., 1993; Weegels et al., 1996; Takata et al., 1999).

In terms of gluten composition, Payne et al. (1979) showed that the high molecular weight glutenin (HMWG) subunits 5+10 coded by the Glu-D1 locus (Glu-D1d allele) were dominant in high quality bread wheats, whereas subunits 2+12 (Glu-D1a allele) were generally found in lower quality wheats. HMWG subunits coded by the complex Glu-1 loci situated on the long arms of chromosomes 1A, 1B, and 1D of wheat were shown to play important roles in the physical properties of dough (Payne et al., 1984). The HMWG subunits that are associated with improved bread quality were reported to have an additive effect on increasing the SDSS volume and loaf volume (Moonen et al., 1982; Rogers et al., 1990; Takata et al., 2000, 2002, 2003). From the point of view of practical breeding, analysis of the effect of SDSS volume screening on the composition of HMWG subunits will be beneficial because of its high throughput potential.

In wheat cultivars with a 1BL.1RS translocation, the short arm of the 1B wheat chromosome is replaced by the short arm of the 1R rye (Secale cereale L.) chromosome. The 1RS arm of rye has been widely used in breeding programs as a resistance source to several diseases and thus as a source of greater potential productivity (Rajaram et al., 1983; Carver and Rayburn, 1994). On the other hand, serious defects in bread quality such as poor mixing tolerance, dough stickiness, and low bread volume have been reported with the presence of the translocation (Dhaliwal et al., 1987; Dhaliwal and MacRitchie, 1990; Lee et al., 1995). Previous studies suggested that HMWG subunits 5+10 played a compensating role for the loss of bread quality associated with 1BL.1RS translocation (Sreeramulu and Singh, 1994; Pflüger et al., 1998; Martín et al., 2001).

Harder wheat grains produce flour with a larger particle size and greater damaged starch content, characteristics which contribute to greater bread quality (Osborne et al., 1997; Ohm et al., 1998). Grain hardness is largely controlled by puroindolines a and b, proteins encoded by the Pina and Pinb genes, which are tightly linked to the Ha locus on chromosome 5D (Giroux and Morris, 1998). Improvements in the measurement of grain hardness has culminated in the single-kernel characterization system (SKCS), based on the force deformation curve obtained from crushing an individual kernel (Martin et al., 1993).

Grain protein content, SDSS volume, and grain hardness have been shown to display low-moderate, moderate, and high heritability, respectively (Baker et al., 1971; O'Brien and Ronalds, 1984; Fischer et al., 1989; Nishio et al., 2005). It is important to investigate the relationship between screening directions and the heritability of the quantitative traits to optimize selection for breeding (Falconer and Mackay, 1996). However, the influence of screening direction on screening effectiveness and the interactions of HMWG subunits and 1BL.1RS translocation to the heritability of bread quality are poorly documented. From a practical perspective, wheat breeding for bread quality requires an understanding of SDSS volume-based screening criteria, and of the influence of HMWG subunits types or the 1BL.1RS translocation on the heritability of small-scale test-assessed quality. In this study, the heritability of upward and downward screening according to criteria based on grain protein content, SDSS volume, and SKCS grain hardness was investigated using two populations differing in HMWG subunits, 1B type and Pinb allele of their parent lines. The influence of screening direction, HMWG subunits and 1BL.1RS translocation on the heritability of small-scale quality criteria, and the usefulness of SDSS volume and SKCS hardness in the screening of bread-quality wheat were assessed to determine appropriate screening direction (up or down) and strength.

	MATERIALS AND METHODS

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Plant Materials
One hundred seventy-two recombinant inbred lines were derived from the cross ‘Hokushin’/KS 831957//Kitami 72/Satsukei 226, termed hereafter the Hokushin population, and an additional 118 lines were derived from the cross Tohoku 195/Satsukei 226//Kitami 72/KS 831957, termed hereafter the Tohoku population. The F₃ lines were derived from F₂ lines by the bulk method, whereas F₄ lines were derived from F₃ lines through random individual selection. Lines KS 831957, Satsukei 226, and Tohoku 195 are superior quality hard red winter wheats; and Hokushin and Kitami 72 are soft red winter wheats. The two populations and parent lines were sown in the fall of 2001 and 2002, at the National Agricultural Research Center for the Hokkaido Region (Memuro). Each experimental unit consisted of a single 2.0-m-long row, laterally 0.72 m distant from the adjoining row. All seeds were sown at intervals of 9 cm. Nitrogen was applied at a rate of 60 kg ha^–1 at seeding, and an additional 20 kg ha^–1 was applied after the April snow melt.

Small-Scale Bread-Making Quality Tests and Calculation of Heritability
Nitrogen content of wheat grain samples was determined using the Dumas combustion method (Rapid-N, Elementar Americas, Inc., Mt. Laurel, NJ) and converted to percentage of protein by multiplying by 5.7 (American Association of Cereal Chemists, 2000). Assessments of grain hardness by the SKCS method (4100 SKCS, Perten Instruments, Springfield, IL), required samples of 50 or 300 kernels for F₃ or F₄ populations, respectively. Puroindoline genotypes of parent cultivars were determined by the method of Lillemo and Morris (2000). The SDSS volume analysis was performed according to the prolonged micro SDSS volume method of Takata et al. (1999), using a 0.7-g sample ground to pass a 0.3-mm sieve. Grain from F₄ lines was conditioned to 14% moisture and milled in a Quadrumat Junior test mill (Brabender GmbH & Co. KG, Duisburg, Germany). Flour particle size (FPS) was measured by laser diffraction particle-size analyzer (Heros & Rodos, Japan Laser Co., Ltd., Tokyo). The HMWG subunits were confirmed by sodium dodecylsulfate polyacrylamide gel electrophoresis (SDS-PAGE) using 7.5% polyacrylamide gels using flour samples of F₄ lines. Using these samples, the presence of the 40-kDa secalin subunit was checked by SDS-PAGE, to confirm the 1BL.1RS translocation (Koebner and Shepherd, 1986). Consequently, 112 lines from the Hokushin population and 72 lines from the Tohoku population were employed for the sequence analysis to confirm homozygosity of HMWG subunits at Glu-1 and further confirm the 1BL.1RS translocation indicated by SDS-PAGE.

Data were analyzed by multifactorial ANOVA. Quality differences between F₃ and F₄ populations were compared using a paired t test. Tukey–Kramer multiple range tests were employed to differentiate means. To calculate realized heritability, the value of each trait was normalized according to each generation's standard deviation. Realized heritability was estimated from the selection differential and genetic gain between F₃ and F₄ generations that were screened by 20% of F₃ populations both for higher rank (upward screening) or lower rank (downward screening) of each quality index.

	RESULTS

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Characteristics of Parent Cultivars
HMWG subunit phenotype determinations, 1B chromosome types, puroindoline genotypes, grain protein content, SDSS volume, and SKCS hardness of parent cultivars are presented in Table 1. The parents differed by two alleles for Glu-A1, three alleles for Glu-B1, and three alleles for Glu-D1. Only Satsukei 226 bore the 1BL.1RS translocation. All parent cultivars carried the Pina-D1a allele at the Pina locus. Hard wheat parent cultivars (KS 831957, Satsukei 226, and Tohoku 195) bore the Pinb-D1b allele at the Pinb locus, while the soft wheat parent cultivars (Hokushin and Kitami 72) bore the Pinb-D1a allele at the Pinb locus.

Characteristics of F₃ and F₄ Populations
Comparisons of means and standard deviations of grain protein content, SDSS volume SKCS hardness and median FPS diameter between F₃ and F₄ populations are presented in Table 2. For both populations, mean protein content was lower in F₄ than F₃ populations. While the SKCS hardness and median FPS diameter maxima of the two populations were similar, the mean and minimum as well as the range in SDSS of the Hokushin population were less than the Tohoku population. Paired t test comparisons of F₃ and F₄ populations were significantly different for all quality parameters, except for the SDSS volume of the Hokushin population and SKCS hardness of the Tohoku population.

View this table: [in this window] [in a new window]   Table 4. Mean values of grain protein content (PC), sodium dodecylsulfate sedimentation (SDSS) volume, single kernel characterization system (SKCS) hardness, and median flour particle size diameter (FPS) of lines of two wheat (Triticum aestivum L.) populations classified by high-molecular-weight glutenin subunits and 1B chromosomal types.

View larger version (32K): [in this window] [in a new window]   Figure 1. Relationship of SDSS volume in F3 lines to that in F4 lines classified by high-molecular-weight glutenin (HMWG) subunits at Glu-B1 and Glu-D1 across the two wheat populations. Hokushin/KS831957//Kitami 72/Satsukei 226. Tohoku 195/Satsukei 226//Kitami 72/KS 831957.

	DISCUSSION

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In our study, mean comparisons between 1B lines and 1BL.1RS translocation lines showed F₄ translocation-bearing lines had increased protein content (Table 4). This concurs with the observations of Carver and Rayburn (1995) and Amiour et al. (2002), but is contrary to those of Fenn et al. (1994), where 1BL.1RS translocation lines had a lower protein concentration. Despite the same complement of 1BL.1RS translocation in the both populations, a significant negative influence of 1BL.1RS translocation on SDSS volume was detected only in the F₄ Tohoku population (Tables 3 and 4). Contrary to Carver and Rayburn (1995) and Martín et al. (2001), this is likely attributable to differences between the studies in terms of the influence of the genetic background response. Carver and Rayburn (1995) indicated the importance of selecting parents that have strong mixing tolerance and high sedimentation volume for crossing with 1BL.1RS translocation parents. The parent line KS 831957, which had particularly strong mixing tolerance (Maruyama-Funatsuki et al., 2004) and a greater SDSS volume (Table 1), is assumed to have compensated for the negative effect of the 1BL.1RS translocation from Satsukei 226. However, further investigation is necessary to elucidate the reason of different negative effect of 1BL.1RS translocation on SDSS volume in each population. As in the case of Fenn et al. (1994), our study found no significant differences in grain hardness between 1B lines and 1BL.1RS lines, whereas Graybosch et al. (1993), when comparing 1BL.1RS translocation lines to their sisters 1R (1B), found the translocation to have significantly decreased grain hardness. In this study, 1BL.1RS translocation lines had greater heritability of SKCS hardness than 1B lines, but only in the Tohoku population. Carver and Rayburn (1995) noted that the decrease in grain hardness depended on the cross. From a genetic point of view, evidence indicates that quality defects associated with 1BL.1RS are influenced by genetic background, which has a potentially larger effect on quality than 1RS itself (Lee et al., 1995).

Heritability of protein content was found to vary from low to moderate values (O'Brien and Ronalds 1987; Fischer et al., 1989; Nishio et al., 2005). Environmental conditions during the growing season can affect grain protein content (Baker et al., 1971; Fischer et al., 1989) and lower measured heritability. The environmental effects and narrow range of protein contents (9.4– 12.9%) within the lines possessing HMWG subunits 4+12 in the Tohoku population were possible reasons for detecting negative heritability of protein content (Table 5). However, further investigation is required to clarify this question.

A number of studies have reported positive effects of HMWG subunits 5+10 on SDSS volume and loaf volume, but negative effect of HMWG subunits 2+12, 4+12, and 20 (Payne et al., 1987; Rogers et al., 1989; Gupta et al., 1994; Buonocore et al., 1996; Takata et al., 2000, 2002, 2003). The scatter plot of SDSS volume for both populations suggests that most lines bearing HMWG subunits 2+12, 4+12, and 20 could be eliminated by cut-off screening at an SDSS volume of 6.0 mL (Fig. 1). Lines with HMWG subunits 5+10 had greater heritability of SDSS volume in upward screening, whereas lines with HMWG subunits 2+12, 4+12, and 20 were all shown to have greater heritability in downward screening (Table 5). These results confirm the effectiveness of SDSS volume screening in eliminating lines offering poorer bread-making quality, when the population includes undesirable HMWG subunits. Greater heritability of SDSS volume in downward screening in a previous study (Nishio et al., 2005) was postulated to have arisen from the lines possessing HMWG subunits 2+12, 4+12, and 20 more consistently showing lower SDSS volumes (Fig. 1). Nakamura (1999) has shown lines possessing HMWG subunits 2.2+12 and 2+12 are dominant in major wheat cultivars developed in Japan, and the number of lines possessing HMWG subunits 5+10 is very limited. Thus, screening by SDSS volume will be a powerful tool in developing bread-making cultivars in such a genetic background. However, further investigation is needed to verify the effectiveness of SDSS volume in different genetic backgrounds.

In our study, the dominant control of grain hardness by the Pinb gene (Giroux and Morris 1998) in both populations was postulated as a possible reason for the greater heritability of SKCS hardness in crosses involving opposite Pinb alleles (Tables 1 and 5). The mean SKCS hardness was greater in the Tohoku population than in the Hokushin population (Table 2), because of the Pinb alleles in the parent lines (Table 1). However, the mean SKCS hardness was significantly and similarly increased in both populations within the lines possessing HMWG subunits 5+10 (Table 4). This indicates that screening for lines with HMWG subunits 5+10 will be useful in increasing grain hardness, regardless of the combination of Pinb alleles in parent cultivars. Our results concur with the observation that transgenic wheat lines enhanced with the HMWG subunits 5+10 gene showed significantly greater grain hardness than the original wheat line (Rakszegi et al., 2005). Autran (1996) reported HMWG subunits play a major role in endosperm texture. The mechanism of increased grain hardness by HMWG subunits 5+10 is postulated to arise from changes in the glutenin proteins altering the efficiency of deposition and thus overall seed development (Don et al., 2003). However, no direct relationship has been shown between allelic variants of the HMWG subunits and grain hardness. Further investigation is needed to elucidate the mechanism of the effect of HMWG subunits 5+10 on grain hardness.

The effect of HMWG subunits 5+10 in increasing SDSS volume, grain hardness, and FPS provide useful tools in the efficient breeding of quality bread wheat. Lines with HMWG subunits undesirable for bread-making quality were clearly separated on the basis of an SDSS volume criterion, and showed greater heritability in downward screening. Screening using this parameter as a selection criterion will be effective in eliminating lines of inferior bread-making quality both for hard by hard or hard by soft wheat crosses. An improvement in the efficiency of bread quality wheat breeding could thus be achieved by the concurrent screening by SDSS volume and SKCS hardness, along with protein content–based criteria.

	ACKNOWLEDGMENTS

 
We are grateful to S. Takahashi, M. Oizumi, Y. Matsudaira, H. Nagahama, S. Yamabuki, and H. Shibukawa for their technical assistance.

	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 July 17, 2006.

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