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Crop Science 41:1080-1084 (2001)
© 2001 Crop Science Society of America

CROP BREEDING, GENETICS & CYTOLOGY

Interaction between Allelic Variation at the Glu-D1 locus and a 1BL.1RS Translocation on Flour Quality in Bread Wheat

P. Martín, M. Gómez and J. M. Carrillo*

Unidad de Genética, ETSI Agrónomos, Universidad Politécnica de Madrid, 28040 Madrid, Spain

* Corresponding author (jmcarrillo{at}bit.etsia.upm.es)


   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIAL AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
1BL.1RS wheat (Triticum aestivum L.)-rye (Secale cereale L.) translocation has been widely used in wheat breeding programs all over the world. Serious defects in bread quality have been associated with the presence of the translocation. The aim of this study was to evaluate the cumulative and interaction effects on strength and mixing properties of dough ascribed to the allelic variation for the Glu-D1 locus and to the presence of 1BL.1RS translocation. A group of F4-F5 recombinant lines were developed from the cross between the 1BL.1RS inbred line S-55-87A and ‘Gazul’, a cultivar without the translocation. Lines were cultivated for 2 yr and evaluated by the SDS-sedimentation test, mixograph, and alveograph. A highly significant difference between years was found in grain protein concentration, while 1BL.1RS translocation or Glu-D1 alleles did not influence this parameter. The difference in sodium dodecylsulfate sedimentation (SDSS) volume mean value between lines with high molecular weight (HMW) 5+10 glutenin subunits and lines with HMW 2+12, considering only the set without the 1BL.1RS translocation, was four times lower than this difference in the set with the translocation, reflecting a significant interaction between the effects of Glu-D1 alleles and 1B chromosome type. The detrimental effect on mixing properties caused by 1BL.1RS rye translocation was dependent on the genetic background. No significant differences among means were detected on mixing time and tolerance rate for presence or not of 1BL.1RS chromosomes. The presence of HMW 2+12 glutenin subunits and 1BL.1RS translocation produces a drastic decline in gluten strength, making it difficult to find acceptable quality among genotypes possessing both negative factors.

Abbreviations: AACC, American Association of Cereal Chemists • A-PAGE, acid polyacrylamide gel electrophoresis • HMW, high molecular weight • NIR, near-infrared reflectance • SDS-PAGE, sodium dodecylsulfate polyacrylamide gel electrophoresis • SDSS, sodium dodecylsulfate sedimentation


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIAL AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
THE QUANTITY and quality of the proteins in the wheat endosperm are major determinants of end-use quality (Wall, 1979). High-molecular-weight glutenin subunits, coded by the complex Glu-1 loci on the long arms of 1A, 1B, and 1D chromosomes, are mainly responsible for the viscoelastic properties of dough (Payne et al., 1984). Payne et al. (1979) were the first to observe that the HMW glutenin subunit pair 5+10, coded by Glu-D1 locus (Glu-D1d allele), was found mostly in high quality bread wheats, while the allelic variant 2+12 (Glu-D1a) was generally found in lower quality wheats.

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 chromosome. The 1RS arm of rye has been widely used in breeding programs all over the world as a resistance source to several groups of diseases and as a source of greater potential grain productivity (Rajaram et al., 1983; Moreno-Sevilla et al., 1995). However, serious defects in bread quality such as poor mixing tolerance, superficial dough stickiness and low bread volume have been associated with the presence of the translocation (Zeller et al., 1982; Dhaliwal et al., 1987). For these translocated wheat flours, the changes in the balance between monomeric (gliadins and secalins principally) and polymeric (glutenins) proteins have been related to stickiness and weakness of doughs (Dhaliwal and MacRitchie, 1990; Lee et al., 1995).

The influence of genetic background on the bread quality of 1BL.1RS lines has been shown (Graybosch et al., 1990; Lee et al., 1995; Moreno-Sevilla et al., 1995). Studies on breadmaking quality effects of allelic variation at the Glu-1 loci and 1BL.1RS translocation have been based on the comparison of heterogeneous groups of cultivars or genotypes with different genetic backgrounds (Sreeramulu and Singh, 1994; Pflüger et al., 1998). In these studies, it was suggested that the subunits 5+10 may play a compensanting role for the loss of dough strength associated with the 1BL.1RS translocation. The additive and interaction effects of allelic variation at the Glu-D1 locus and 1BL.1RS translocation have not been reported before this study.

The purpose of this study was to evaluate the cumulative and interaction effects on strength and mixing properties of dough resulting from the allelic variation at the Glu-D1 locus and to the presence of 1BL.1RS translocation using materials with a common genetic background.


   MATERIAL AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
RESULTS
 DISCUSSION
 REFERENCES
 
A cross of S-55-87A, a 1BL.1RS inbred line, and Gazul, a bread wheat cultivar without the 1BL.1RS translocation, was used. Twenty-eight F2 genotypes homozygous for the loci of HMW glutenin subunits were taken from this cross, being homozygous (1B, 1B or 1BL.1RS, 1BL.1RS pair) or heterozygous (1B, 1BL.1RS pair) for the short arm of the 1B chromosome. At the Glu-D1 locus, Gazul possesses HMW 5+10 glutenin subunits (Glu-D1d) and S-55-87 possesses HMW 2+12 glutenin subunits (Glu-D1a). Both parents possess HMW 1 and HMW 7+8 glutenin subunits (Glu-A1a and Glu-B1b alleles, respectively). Glutenins were visualized by sodium dodecylsulfate polyacrylamide gel electrophoresis (SDS-PAGE), whereas the presence or absence of secalins (Gli-B1l allele block described by Metakovsky, 1991) was detected by acid polyacrylamide gel electrophoresis (A-PAGE) in a gel at pH 3.1 with electrophoresis following Lafiandra and Kasarda (1985).

F2 plants, selected for their protein composition, were grown in greenhouse and seeds from each F2 plant were bulked. F4 and F5 lines, were grown in the field in a 2-yr experimental design with two random complete blocks each. Experiments were conducted under field conditions at the Agronomical Engineer University fields at Madrid (Spain) during 1995 and 1996, in plots with three rows 2 m long and spaced 20 cm apart. The soil was a well-drained Typic Xerorthents, with a sandy loam texture. To analyze a population with the same genetic background, the parents were not included in the test.

Grain was tempered to 155 g kg-1 moisture content and 1-mm granule wholemeal flour was obtained with a Udy Cyclone mill (Tecator AB, Sweden). Protein concentration (Pc) was measured with a near-infrared reflectance (NIR) spectroscope (Infra-alyzer 300; Bran-Luebbe, Hamburg, Germany) and SDSS volume was measured according to Mansur et al. (1990). Dough mixing properties were obtained with a 10-g mixograph (National Manufacturing Co., Lincoln, NE). Samples were mixed to optimum water absorption following 54-40A method (AACC, 1995). Optimum mixing time (MT, in min), maximum height (H, in cm), height 3 min after reaching the peak (H3, in cm), and tolerance range or angle formed by the curve peak (TR, in degrees), were measured for both years. Grain from the second year was milled in a Chopin mill and flour was assayed in a Chopin micro-alveograph for 50 g of flour. AACC regulation 54-30 was followed (Faridi and Rasper, 1987), and dough strength (W in 10-4 J) and dough extensibility (L, in cm) were estimated.

The statistical study of the relationship between genetic variability characterized in the lines and bread flour quality variables was conducted through multifactorial variance analysis based on models with fixed effects using the GLM procedure of the SAS system (SAS Institute Inc., 1986). Separation of means was verified with Duncan's test. Principal component analyses (PCA) were conducted with the centered and reduced values of simple parameters measured in the mixograph test (MT, H, H3, and TR) using STAT-ITCF software. Diagrams were obtained in the first principal plane.


   RESULTS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS
DISCUSSION
 REFERENCES
 
Lines were characterized on the basis of their composition for glutenins and secalins: HMW glutenin subunits produced by alleles at Glu-D1 locus (HMW glutenin subunits 2+12 and HMW glutenin subunits 5+10) and 1B chromosome type (1B, heterozygotic, 1BL.1RS) were considered for statistical analysis. The characteristics of the analyzed population were as follow: 1B and HMW glutenin subuints 5+10 (5 lines); 1B and HMW glutenin subunits 2+12 (4 lines); 1BL.1RS and HMW glutenin subunits 5+10 (2 lines), 1BL.1RS and HMW glutenin subunits 2+12 (3 lines); heterozygous type and HMW glutenin subunits 5+10 (7 lines); heterozygous type and HMW glutenin subunits 2+12 (7 lines).

Table 1 shows the mean squares of the multifactorial analysis of variance for a model based on those two sources, their interactions, and the year factor. This model accounts for most of the observed variation in quality parameters estimates (percentage of total mean squares): 83% for Pc, 70% for SDSS, 75% for MT, 62% for TR, or 50% for W. No significant interactions for these quality parameters were detected between the year and the other two sources of variation.


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  		Table 1. Mean squares from the multifactorial analysis of variance for grain protein concentration (Pc), sedimentation volume (SDSS), mixing time (MT), maximum mixogram height (H) and height 3 min after the mixogram peak (H3), tolerance range (TR), dough strength (W) and dough extensibility (L).

 
Grain protein concentration was high as result of low rainfall, especially for the first year (Table 2). The water shortage in the first year made highly significant the difference between years (P < 0.001) for this parameter (Table 1). However, protein concentration was not affected by variation at Glu-D1 locus or by 1B chromosome type.


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  		Table 2. Mean values of flour quality parameters of all genotypes: grain protein concentration (Pc), sedimentation volume (SDSS), mixing time (MT), maximum mixogram height (H) and height 3 min after the mixogram peak (H3), tolerance range (TR), dough strength (W) and dough extensibility (L).

 
Lines possessing HMW glutenin subunits 5+10 had higher SDSS volumes than lines possessing HMW glutenin subunits 2+12 (Table 2). Similar significant differences in SDSS volume were observed between the three classes of 1B chromosome type. Within the non-1BL.1RS set (Table 2), the difference for mean SDSS volumes between lines possessing HMW glutenin subunits 5+10 (93.7 mm) and those with HMW glutenin subunits 2+12 (86.3 mm) was 7.4 mm. However, for the set with the translocation, lines with HMW glutenin subunits 5+10 (85.4 mm) and with HMW glutenin subunits 2+12 (58.2 mm) differed by 27.2 mm. This implies a highly significant interaction (P < 0.001) between the effects of the 1B chromosome type and Glu-D1 alleles (Table 1). This interaction effect was observed for the 2 yr (Table 3), where no significant differences were found on SDSS volume between chromosome types when only lines presenting HMW glutenin subunits 5+10 were considered. However, within lines with HMW glutenin subunits 2+12 the differences were significant for both years.


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  		Table 3. Mean values for grain protein concentration (Pc), sedimentation volume (SDSS), mixing time (MT), maximum mixogram height (H) and height 3 min after the mixogram peak (H3), tolerance range (TR), dough strength (W) and dough extensibility (L), organized by year, Glu-D1 alleles and 1B chromosome type.

 
Regarding mixograph parameters, no significant differences between means were detected in mixing time (MT) and tolerance range (TR) for presence or not of 1BL.1RS translocation. However, the differences for the other mixing parameters (H and H3) were highly significant (Table 1 and 2). Concerning the Glu-D1 allelic variation, the MT and TR values associated with the presence of HMW glutenin subunits 2+12 were 27.3% and 20.5% lower (P < 0.001) than those with HMW glutenin subunits 5+10 respectively. No differences (P < 0.05) were observed for H and H3, respectively. As for protein concentration and SDSS volume, significant differences among years were manifested for mixing time and tolerance range values, but not for H and H3.

The interaction effect detected for SDSS also was detected (P < 0.001) for mixing time values (Table 1). The set of lines possessing identical Glu-D1 alleles were analyzed separately resulting in significant lower MT values for the 1BL.1RS set when compared with 1B type set under presence of HMW glutenin subunits 2+12, but not when HMW glutenin subunits 5+10 were present (Table 2). The interaction effect of the translocation on lines differing in Glu-D1 alleles was even more marked in the second year for the rest of mixing parameters except for TR (Table 3).

Mixograph parameters account for 42.7% of the variance in the principal component analysis (PCA) for the first principal axis and for 37.8% for the second one. Four groups of materials are clearly apparent on illustration of the first factorial plane (Fig. 1), framed by the bisecting line for the two principal axes.

  1. 1B lines carrying HMW glutenin subunits 5+10—white circles—exhibiting the highest values for quality variables.
  2. 1B lines with HMW glutenin subunits 2+12—white triangles, which exhibit high values for H but shorter mixing time.
  3. 1BL.1RS lines with HMW glutenin subunits 5+10—dark circles—with higher MT and TR but lower H.
  4. 1BL.1RS lines with HMW glutenin subunits 2+12—dark triangles—showing the lowest quality properties altogether.



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  		Fig. 1. Principal component analysis of F4 lines from the Gazul x S-66-87A cross with the data from the mixograph parameters.

 
The well-defined division for the observations were based on the Glu-D1 alleles fixed (circles versus triangles) for the mixing time and tolerance range values remained clear even when heterozygous lines for translocation were present on the first factorial plane.

For the alveograph, both the presence of 1BL.1RS translocation and the presence of HMW glutenin subunits 2+12 produced a detrimental effect on dough strength (W) for the material analyzed (Table 2). When they occurred together, their effects were additive and mean value for W dropped significantly to 187.7 x 10-4 J. In contrast, high W values were obtained when non translocated lines possess the HMW glutenin subunits 5+10 (mean value of 405.8 x 10-4 J). When HMW glutenin subunits 5+10 appeared in 1BL.1RS translocated lines, a mean value for W of 298.5 x 10-4 J confirmed the improving role of these glutenins (Table 2).

No significant differences were detected when analyzing all the lines together for dough extensibility (L) values (Table 1). Likewise, no differences in L for 1B chromosome type were detected when only the set of lines having HMW glutenin subunits 2+12 was analyzed. However, in the set of lines possessing HMW glutenin subunits 5+10, the extensibility dropped 32.8% from non-translocated lines (11.3 cm) to 1BL.1RS lines (7.6 cm) (Table 3).


   DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS
 DISCUSSION
REFERENCES
 
A significant influence of year factor on SDSS values and some dough mixing properties was present. This effect could be attributed in a great extent to the different grain protein concentration between years, with the first year 16.6% higher than the second one. Protein concentrations were high as a consequence of water shortage, and those high levels probably increased gluten strength, mixing time and tolerance range. However, grain protein concentration was not affected by Glu-D1 allelic variation or by the presence of the 1BL.1RS translocation, in agreement with Dhaliwal et al. (1987) and Peña et al. (1990), allowing a correct study of these two sources of variation over quality parameters.

It is interesting to note that MT and TR mean values for each year and for each Glu-D1 allele were highly significantly different, but not for each chromosome type. Mixing time is the mixograph parameter the least affected by the translocation (Carver and Rayburn, 1995; Moreno-Sevilla et al., 1995; Martín and Carrillo, 1999). Likewise, no significant differences were found by Dhaliwal et al. (1987) and Martín and Carrillo (1999) in the values related to tolerance to overmixing (TR) between the 1B and 1BL.1RS lines. In contrast, H and H3 were the mixing parameters less affected by the year factor and most independent from Glu-D1 allelic variation, but at the same time, the most affected by the 1B chromosome type.

In 1BL.1RS translocated wheats, the increase of monomeric proteins (secalins coded by genes on the 1RS arm) and the decrease of glutenin concentration (lack of low molecular weight subunits coded by genes on the 1BS arm) are mainly responsible for the negative effects of this translocation on flour quality (Graybosch et al., 1990; Lee et al., 1995). The influence of secalin presence was here clearly detected for dough strength (SDSS and W parameters). The heterozygous lines for 1B chromosome type, which contain only one secalin dose, produced mean values intermediate between those for homozygous classes, confirming the cumulative effect that the replacement of one or two 1BL.1RS chromosomes has on quality. In different genetic backgrounds, Martín and Carrillo (1999) analyzed the dominance effects in the inheritance of quality traits associated to the 1B chromosome type and showed that genes added by the rye chromatin in 1BL.1RS genotypes are quantitatively more important than the genes lost by the 1BS arm in most flour quality parameters. These results confirm those of other authors (Zeller et al., 1982) who have stated that the presence of secalins in flour partly explains the deficiencies in quality of 1BL.1RS lines.

Studying HMW glutenin subunit composition and flour quality of 1B and 1BL.1RS cultivars several authors (Sreeramulu and Singh, 1994; Pflüger et al., 1998) have shown that HMW glutenin subunits 5+10 continue to be an important breeding objective to improve breadmaking quality. Whereas for Bullrich et al. (1998) none of the backgrounds studied could counterbalance the effect of the 1BL.1RS translocation, in the background tested in this experiment, the SDSS and MT parameters for 1B and 1BL.1RS lines have similar mean values when the HMW glutenin subunits 5+10 subunits are present. Thus, the beneficial effect of those subunits on gluten strength seems to be greater in 1BL.1RS than in non translocated wheats. With a lower significance level than for SDSS volume and MT, the additive effect and the interaction between the 1B chromosome type and the Glu-D1 allelic variation were also observed on the other mixing properties and on extensibility, indicating that the detrimental effect caused by the translocation is dependent on the genetic background.

The replacement of the Glu-D1a allele by the Glu-D1d allele would permit selection of lines possessing 1BL.1RS translocation with a substantive improvement on mixing properties. However, it seems that no totally acceptable values for H or H3 parameters will be reached, as indicated in the principal component analysis (Fig. 1), where no translocation line overcomes the second bisecting line. The low range of variation for these two parameters within the 1BL.1RS class indicates that the mixograph variables are more difficult to improve in 1BL.1RS translocated wheats.

Changes on polymeric protein size distribution associated with the allelic variation for Glu-D1 (Gupta and MacRitchie, 1994) seem to affect some quality parameters to a greater extent than changes in monomeric and polymeric protein proportions associated with the presence of 1BL.1RS rye translocation. The simultaneous presence of HMW glutenin subunits 2+12 and 1BL.1RS translocation produced a drastic decline in gluten strength as consequence of a strong interaction between both factors, making the selection for acceptable quality of genotypes possessing both negative factors more difficult.

Detection of additive and interaction effects between the 1B chromosome pair type and the allelic variation for prolamin genes, as identified in this work, could contribute to a more efficient breeding program on these materials through the parental search in recurrent processes.


   ACKNOWLEDGMENTS
 
This work was supported by grant No. AGF 97-937 from the Comisión Interministerial de Ciencia y Tecnología (CICYT) of Spain.

Received for publication June 21, 2000.


   REFERENCES
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 




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