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PLANT GENETIC RESOURCES

Milling and Bread Baking Traits Associated with Puroindoline Sequence Type in Hard Red Spring Wheat

^a Dep. of Plant Sciences, PO Box 173140, Montana State Univ., Bozeman, MT 59717
^b USDA-ARS Western Wheat Quality Laboratory, Pullman, WA 99164-6394
^c Dep. of Plant Sciences, North Dakota State Univ., Fargo, ND 58105

Corresponding author (jmmartin{at}montana.edu)

	ABSTRACT

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
Recent results have shown that mutations in genes coding for puroindoline a and b (PinA and PinB) are associated with the expression of the hard texture of wheat (Triticum aestivum L.) grain. A majority of hard wheats have a glycine-to-serine mutation in puroindoline b (allele PinB-D1b), or they are devoid of puroindoline a (allele PinA-D1b). Hard wheats with PinA-D1b tend to be harder than those with PinB-D1b. Grain hardness is known to affect milling and baking traits. Our objective was to determine the influence of allelic variation in PinA and PinB on milling and bread quality traits in a recombinant inbred population segregating for PinA-D1b and PinB-D1b. One hundred thirty-nine recombinant inbred lines from the cross `Butte 86' (PinA-D1b allele)/ND2603 (PinB-D1b allele) and parents were grown in a field trial with two replications at two locations. Grain hardness was measured by near-infrared reflectance (NIR) and the single-kernel characterization system (SKCS). Grain was milled and baked for each line. Puroindoline allele type was determined for each line. The PinB-D1b group had significantly softer grain, higher break flour yield, flour yield, milling score, and loaf volume, and lower flour ash and crumb grain score (low score being desirable) than the PinA-D1b group. Significant genetic variability was detected within allelic classes for all traits. The proportion of variation among entry means attributed to puroindoline classes was 34% for break flour yield, 26% for NIR hardness, and 22% for SKCS harness index. Grain hardness was negatively correlated with break flour yield, flour yield, and mixing score and positively correlated with flour ash. Grain hardness was not correlated with loaf volume or crumb grain score. The PinB-D1b allele was more desirable for milling and bread baking, although superior milling and bread quality genotypes could be selected within either class.

Abbreviations: NIR, Near-infrared reflectance • SKCS, Single Kernel Characterization System

	INTRODUCTION

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
WHEAT IS CLASSIFIED into hard and soft classes on the basis of the texture of the grain. These textural classes coincide with differences in milling and end-use properties (reviewed in Pomeranz and Williams, 1990; Morris and Rose, 1996). The distinction between soft and hard classes of wheat is governed by the Hardness (Ha) locus on chromosome 5DS (Mattern et al., 1973; Law et al., 1978) with additional modifying genes contributing to variation within classes (Symes, 1965; Baker, 1977); however, Baker and Sutherland (1991) and Giroux et al. (2000) observed significant genetic variation for grain hardness within crosses of hard wheats.

Greenwell and Schofield (1986) identified friabilin as a marker protein for grain softness which was present in larger amounts on the surface of water-washed starch of soft wheats than from hard wheats (Bettge et al., 1995; Greenblatt et al., 1995; Morris et al., 1994). Friabilin is composed of two major polypeptides termed puroindoline a and puroindoline b. Genes coding for these two proteins, PinA and PinB, are tightly linked to the Ha locus on chromosome 5D (Jolly et al., 1993; Sourdille et al., 1996) and probably function together as the Ha locus. Recent results have shown that mutations in PinA and PinB are associated with the expression of hard texture. Giroux and Morris (1997)(1998) showed that hard texture was completely linked to a glycine-to-serine mutation in puroindoline b (allele PinB-D1b), or the complete absence of the puroindoline a protein (allele PinA-D1b). In a survey of hard wheats, cultivars with the PinA-D1b allele were on average 7 units harder than those with PinB-D1b (Giroux and Morris, 1997; unpublished results). Giroux et al. (2000) further showed that progeny carrying the PinA-D1b allele averaged 4.5 units harder than progeny with PinB-D1b in three hard red spring crosses segregating for PinA-D1b vs PinB-D1b. A more recent survey has found additional mutant alleles of PinA or PinB linked to hard textured grain (Lillemo and Morris, 2000).

Since kernel texture has been shown to be associated with numerous milling and bread quality traits in hard wheats (Slaughter et al., 1992) and Giroux et al. (2000) showed hard wheats with the PinA-D1b allele tend to be harder than those with the PinB-D1b allele, it is possible that allelic variation at the PinA and PinB loci affects milling and bread quality traits. Our objective was to determine the influence of allelic variation in PinA and PinB on milling and bread quality traits in a recombinant inbred population segregating for PinA-D1b and PinB-D1b.

	MATERIALS AND METHODS

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
One hundred thirty-nine hard red spring wheat recombinant inbred lines were derived from the cross `Butte 86'/ND 2603. Lines were derived by single-seed-descent from F<sub>2</sub> to F<sub>6</sub> followed by a generation of seed increase to produce F<sub>6</sub>-derived F<sub>8</sub> lines. Butte 86 carries the PinA-D1b allele while ND 2603 has the PinB-D1b allele. The ND 2603 parent has pedigree `Wheaton'/`Sumai 3'. The 139 recombinant inbred lines and parents were grown at Pullman, WA, and 134 lines and parents were grown at Bozeman, MT, in 1998. Each trial was a randomized complete block design with two replications. Each plot was a single 3-m row with 30 cm between rows. Grain from each plot was threshed for milling and bread quality analyses.

Wheat samples were analyzed for moisture content (Method 44-16) and test weight (Method 55-10) (AACC, 2000). Near-infrared reflectance (NIR) hardness (Method 39-70A) (AACC, 2000) was determined from a single aliquot using a near-infrared reflectance spectrometer (model IA450, Technicon, Hoganas, Sweden) on whole grain meal (0.5-mm screen) (UDY Corp., Fort Collins, CO). The Single Kernel Characterization System 4100 (SKCS) (Perten Instruments North America, Inc., Springfield, IL) was used to estimate grain hardness (SKCS hardness index), kernel weight, and kernel diameter (thickness or outer diameter) using a sample of 300 kernels from each plot.

Wheat was tempered to 145 g kg^-1 (14.5%) moisture (fresh weight basis) and milled on Quadrumat Jr. mills following modifications of Jeffers and Rubenthaler (1977). Mill output was separated using 35- (500 µm) and 100-mesh (150 µm) Tyler test sieves into bran, middlings stock and break flour fractions. Break flour yield was calculated as the proportion of break flour to total products. Middlings were further milled to produce reduction flour and shorts. All tests were conducted on "straight-grade" flour derived from combining the break and reduction flour streams, and expressed as flour yield as a proportion of total products. Milling score was calculated as:

Nitrogen content of wheat and flour was determined on a 0.25-g aliquot of the UDY-ground sample using the Dumas combustion method (model FP-428, Leco Corp., St. Joseph, MI) (AACC, 2000), and converted to percentage protein by multiplying by 5.7. Flour was analyzed for moisture (Method 44-16), protein (Method 46-30), and ash (Method 08-01) (AACC, 2000). All grain and flour parameters are reported on a 120 g kg^-1 (12%) and 140 g kg^-1 (14%) moisture basis, respectively. Mixogram analysis was conducted using the 10-g instrument following Method 54-40A (AACC, 2000).

Bread was baked and scored according to Method 10-10B (AACC, 2000) using an optimum absorption, optimum mixing, 90-min fermentation "straight-dough" comprised of 100 g flour (14% mb), 1.8 g dry active yeast, 1.5 g NaCl, 6 g sucrose, 0.3 g malt extract (60 mg commercial malted barley flour [Amylomalt, Cargill Flour Milling, Ogden, UT] per mL extract), 4 g powdered nonfat dry milk, 3 g partially hydrogenated vegetable shortening with mono- and diglycerides (Crisco, Procter & Gamble, Cincinnati, OH), and 7.5 mg ascorbic acid. Crumb grain was scored on the basis of the consensus score of three experienced bakers using a range of 1 (excellent) to 9 (unsatisfactory).

Puroindoline allele type was determined as previously described (Giroux and Morris, 1998). Briefly, Triton X-114 soluble proteins were extracted from a sample of 10 to 20 seeds and then subjected to sodium dodecylsulfate polyacrylamide gel electrophoresis (SDS-PAGE). Samples were screened for the presence or absence of puroindoline a. Presence of puroindoline a was interpreted as PinB-D1b allele (the serine mutation in PinB) and absence as PinA-D1b allele (devoid of puroindoline a).

Analyses of variance combined across locations were performed for each trait, where locations were considered fixed and replications within locations, entries, and entry x location interaction as random effects. Heritability estimates were computed on a progeny mean basis for each trait. The entry variation was further partitioned by including a fixed effect for puroindoline class and a random effect for entries within class and associated interactions with location. The analyses were carried out by PROC MIXED in SAS (SAS Institute, 1997). Least squares means were obtained for the two puroindoline classes, and the difference between the PinA-D1b and PinB-D1b means was compared with a t-statistic, where the standard error of the difference was computed from the appropriate linear combination of mean squares and degrees of freedom by the Satterthwaite (1946) approximation. Transgressive segregates were defined as those lines that were less than the lowest ranking parent or greater than the highest ranking parent by more than one LSD. The proportion of the variation attributed to the PinA-D1b vs PinB-D1b class difference was determined as the ratio of the class sum of squares to the entry sum of squares. Correlations among traits were computed from entry means. Three lines at the Bozeman location did not provide enough flour for baking. These lines were dropped from analyses for bake absorption, loaf volume, and crumb grain score.

	RESULTS AND DISCUSSION

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
Significant genetic variation was observed for all traits. Entry x location interaction variances were also significant in all instances. However, the location x puroindoline mutation class interaction was not significant for any trait.

The two parents could not be differentiated statistically for any of the traits except flour protein where Butte 86 (PinA-D1b) exceeded ND 2603 (PinB-D1b) (Table 1). There was a trend for Butte 86 to have harder kernels (81.5 vs 72.5 NIR units and 63.8 vs 61.0 SKCS units), higher grain protein (149 vs 139 g kg^-1), higher flour ash (4.07 vs 3.82 g kg^-1), and longer mixing time (3.54 vs 3.05 min), but lower kernel weight (32.3 vs 35.4 mg kernel^-1) than ND 2603.

The NIR and SKCS methods of measuring grain hardness were correlated (r = 0.53; P < 0.01). That is less than the r = 0.87 (P < 0.01) between the two methods Morris et al. (1999) reported using 83 recombinant chromosome 5D substitution lines segregating for soft vs hard grain texture for 5D and r = 0.81 (P < 0.01) for 72 hard wheat samples (Ohm et al., 1998). Both methods were positively correlated (P < 0.01) with flour ash but negatively correlated (P < 0.01) with break flour yield, flour yield, and milling score (Table 3). NIR hardness was positively correlated with mixograph and bake absorption, while SKCS hardness index was not correlated with either. Conversely, SKCS hardness index was negatively correlated with kernel weight (P < 0.01) and kernel diameter (P < 0.05), but NIR hardness was not correlated with either. The greatest disparity between the two methods occurred for wheat and flour protein concentration where NIR hardness was positively correlated (P < 0.05) with both, but SKCS hardness index was negatively correlated (P < 0.01) with both. Wheat protein is often positively related with grain hardness within hard wheats (Giroux et al., 2000; Slaughter et al., 1992). The less than complete association between NIR and SKCS hardness index, and their different associations with some traits may reflect the differing approaches to quantifying grain hardness. NIR estimates hardness through spectral characteristics of particle size distribution. SKCS hardness index results from a force-deformation curve derived from crushing individual kernels that is influenced by moisture, kernel size, and kernel weight (Martin et al., 1993).

View larger version (28K): [in this window] [in a new window]   Fig. 1. Distribution of 45 lines within PinA-D1b class and 89 lines within PinB-D1b class for Near-infrared reflectance (NIR) hardness from a cross between Butte 86 (PinA-D1b allele) and ND 2603 (PinB-D1b allele) determined on the basis of mean of two locations

View larger version (29K): [in this window] [in a new window]   Fig. 2. Distribution of 45 lines within PinA-D1b class and 89 lines within PinB-D1b class for single kernel characterization system (SKCS) hardness index from a cross between Butte 86 (PinA-D1b allele) and ND 2603 (PinB-D1b allele) determined on the basis of mean of two locations

View larger version (29K): [in this window] [in a new window]   Fig. 3. Distribution of 45 lines within PinA-D1b class and 89 lines within PinB-D1b class for percentage break flour yield from a cross between Butte 86 (PinA-D1b allele) and ND 2603 (PinB-D1b allele) determined on the basis of mean of two locations

Hard wheats suffer more starch damage during milling and tend to absorb more water during dough formation than soft wheats (Slaughter et al., 1992). Surprisingly, the harder textured PinA-D1b group did not differ from the PinB-D1b group for mixograph or bake absorption (Table 2). In addition, hardness was only weakly associated with absorption (r < 0.3). This may suggest that our genotypes represented a smaller range in hardness than would be encountered in comparing hard and soft textured wheats.

Giroux and Morris (1997)(1998) observed that hard wheats had either the PinB-D1b allele (the serine mutation in PinB) or the PinA-D1b allele (devoid of puroindoline a). Our results and those from Giroux et al. (2000) have confirmed that PinA-D1b genotypes are harder than PinB-D1b genotypes. Dubreil et al. (1998) observed that puroindoline a concentration was inversely related to grain hardness in a sample of 32 wheats. The softer grain conferred by PinB-D1b may be a function of quantity of puroindoline a, b, or both. Some residual function of PinB may remain in PinB-D1b types versus the more severe mutation in PinA-D1b. Dubreil et al. (1998) also observed that flours that were opposite in bread quality, but lacking puroindoline a, had significantly higher loaf volumes than the same flours reconstituted with puroindolines (80% puroindoline a and 20% puroindoline b). Rheological properties of dough were altered in opposite directions by addition of puroindolines. Dough strength and extensibility were reduced in the poor quality flour but increased when puroindolines were added to the good quality flour. Our results showed lower loaf volume from flours lacking puroindoline a (PinA-D1b allele) compared to those with both puroindoline a and b (PinB-d1b allele). These findings, coupled with those from Dubreil et al. (1998) suggest that quantity of puroindolines and/or the ratio of puroindoline a to b may have a role in dough formation and resultant loaf volume.

If the PinA-D1b or PinB-D1b allele conferred an advantage for improved milling or bread quality traits, breeders could use it as a selectable marker to improve milling or baking quality. Our results showed that the PinB-D1b allele may be more desirable because it conferred a significant advantage over the PinA-Db1 allele through increased break flour and flour yield, milling score, and loaf volume with lower flour ash and crumb grain score (lower crumb grain score being desirable). Significant genetic variability was detected within allelic classes for all traits, indicating that superior milling and bread quality genotypes could be selected within either class.

	ACKNOWLEDGMENTS

 
This research was supported in part by grants from USDA-ARS NRICGP (99-01742) and Montana Wheat and Barley Committee.

	NOTES

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
Contribution No. J-2000-27 Montana Agric. Exp. Stn. Mention of trademark or proprietary products does not constitute a guarantee or warranty by the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products that may also be suitable. This article is in the public domain and not copyrightable. It may be freely reprinted with customary crediting of the source.

Received for publication April 10, 2000.

	REFERENCES

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 

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This Article Abstract Figures Only Full Text (PDF) Free Alert me when this article is cited Alert me if a correction is posted Services Similar articles in this journal Similar articles in ISI Web of Science Alert me to new issues of the journal Download to citation manager Citing Articles Citing Articles via HighWire Citing Articles via ISI Web of Science (50) Citing Articles via Google Scholar Google Scholar Articles by Martin, J.M. Articles by Giroux, M.J. Search for Related Content PubMed Articles by Martin, J.M. Articles by Giroux, M.J. Agricola Articles by Martin, J.M. Articles by Giroux, M.J. Related Collections Crop Physiology & Metabolism Wheat Cell Biology & Molecular Genetics