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Crop Science 43:219-226 (2003)
© 2003 Crop Science Society of America

CROP ECOLOGY, MANAGEMENT & QUALITY

Genotype x Environment Interaction for Grain Color in Hard White Spring Wheat

M. A. Matus-Cádiza, P. Hucl*,a, C. E. Perrona and R. T. Tylerb

a Dep. of Plant Sciences and Crop Development Centre, Univ. of Saskatchewan, 51 Campus Dr., Saskatoon, SK, S7N 5A8 Canada
b Dep. of Applied Microbiology and Food Sci., Univ. of Saskatchewan, 51 Campus Dr., Saskatoon, SK, S7N 5A8 Canada

* Corresponding author (hucl{at}sask.usask.ca)


   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
Improvement of grain color in hard white spring wheat (Triticum aestivum L.) breeding programs depends on understanding the influences of genotype (G), environment (E), and their interaction (G x E). The objectives of this study were to quantify genetic variability for grain color and assess the nature of the G x E interaction in determining grain color in 79 spring wheat genotypes. Twelve check cultivars [seven hard red (HR), four hard white (HW), and one soft white (SW)] and 67 white-seeded Australian (AUS) accessions were grown at two locations across 2 yr. Wheat genotypes differed significantly in agronomic traits, grain protein, and kernel hardness. Grain and meal color were quantified using Hunterlab colorimeter values. Whole grain color values without (L = 40.9–50.4 units; a = 7.0–8.3; b = 13.6–19.1) and with NaOH treatment (L = 22.7–38.1; a = 7.7–9.7; b = 9.2–17.9) varied among genotypes. Using ground meal, color values (L = 80.1–84.9; a = 1.8–2.6; b = 8.9–11.8), yellow pigment content (2.5–4.8 µg g-1), and lutein content (1.8–3.7 µg g-1) varied among genotypes. Genotype x location (L) interactions were not significant for colorimetric and pigmentation variables. The Azallini and Cox test detected one crossover G x year (Y) interaction for grain a-value (without NaOH), one for grain b-value (without NaOH), and 12 for lutein content. Genetic variation exists for grain color among HW genotypes. The noncrossover nature of G x E interactions for grain color indicates that white-seeded genotypes selected as superior in one environment will be superior in other environments.

Abbreviations: a-value, Hunterlab redness value • AUS, Australian • b-value, Hunterlab yellowness value • CPS, Canada Prairie Spring • CWRS, Canada Western Red Spring • CWWS, Canada Western White Spring • E, environment • G, genotype • HR, hard red • HW, hard white • KCRF, Kernen Crop Research Farm • L, location • L-value, Hunterlab lightness value • SF, Seed Farm • SW, soft white • Y, year


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
WITH THE INTRODUCTION of the experimental market class Canada Western White Spring (CWWS) in Canada, kernel color has become an important discriminator between HR (Canada Western Red Spring, CWRS) and HW (CWWS) spring wheat classes. International markets, particularly in Asia where products such as steamed bread and noodle products predominate, often prefer white to red-seeded wheat (Crosbie et al., 1998). Australia is currently the largest world producer of HW wheat (Teetaert, 2000). If HW wheat production increases to significant levels in Canada, then we stand to capture a larger share of the wheat export market. ‘AC Snowbird’ was recently (interim) registered as the first CWWS cultivar in Canada. White-seeded CWWS breeding lines must be readily distinguishable from red-seeded CWRS lines to meet registration standards. Only one Canadian cultivar (i.e., ‘Norquay’) has been associated with visual distinguishability problems regarding kernel color (DePauw and McCaig, 1988). The NaOH color test can overcome the subjectiveness of visually classifying the grain color of troublesome wheat samples (Lamkin and Miller, 1980; DePauw and McCaig, 1988). Selecting for superior grain lightness in CWWS cultivars will depend, in part, on the existence of genetic variation for grain color. To date, little information is available on the presence of genetic variability for grain color in HW spring wheat.

Red kernel color is controlled by up to three loci, located on chromosomes of homologous Group 3, with partial dominance over white kernel color (Metzger and Silbaugh, 1970). In addition to three major loci controlling red seed color, there may be as many as six minor genes influencing grain color (Freed et al., 1976; Reitan 1980). Kernel color is known to be a highly heritable trait (Cooper and Sorrells, 1984); however, color is known to vary in its degree of lightness across environments. Wu et al. (1987) reported that kernel color of white wheat cultivars from northern China was darker when grown in the more humid region of the Lower Yangtze Valley. Wu et al. (1999) reported that some HW winter experimental lines were found to be more sensitive to environmental changes than HR winter check cultivars, but others were found with improved levels of color and color stability across environments when compared with HW winter check cultivars. Peterson et al. (2001) documented variation in grain color in white winter wheat across an array of production environments and cultivars. Only one of 18 HW cultivars grown at five locations in 1996 produced grain that was equal to or lighter than the color standard while only one of the 14 cultivars grown at six locations in 1997 had acceptable grain color. Little information is available to date on the nature of the G x E interaction in determining grain color in HW spring wheat.

Selection for grain color in a HW wheat breeding program requires knowledge of the magnitude of the G x E interaction. If interactions between G x E exist, then genotypes selected as superior in one environment may not be superior in other environments (Baker and Kosmolak, 1977). A significant G x E interaction may be either (I) a noncrossover G x E interaction where the ranking of genotypes remains constant across environments and the interaction is significant because of changes in the magnitude of the response, or (ii) a crossover G x E interaction where a significant change in rank occurs from one environment to another. When selecting genotypes for wide adaptation, plant breeders look for a noncrossover G x E interaction or preferably the absence of a G x E interaction. The objectives of this study were to quantify genetic variability for grain color and assess the nature of the G x E interaction in determining grain color in 79 common spring wheat genotypes.


   MATERIALS AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
RESULTS
 DISCUSSION
 REFERENCES
 
Seventy-nine diverse spring wheat genotypes including 12 check cultivars and 67 white-seeded AUS accessions were used in this study (Table 1). All 79 genotypes possessed a hard endosperm texture, except for the following eight genotypes: ‘AC Reed’, ‘Vectis’, ‘Tatiara’, ‘Sunset’, ‘Katunga’, ‘Eradu’, ‘Reeves’, and ‘Chile Late’. Hard red checks consisted of seven Canadian cultivars from the CWRS class (‘Roblin’, ‘AC Elsa’, ‘AC Barrie’, ‘CDC Teal’, ‘Laura’, ‘AC Domain’, and ‘Columbus’). White-seeded check cultivars consisted of four Canadian cultivars [‘AC Karma’, Canada Prairie Spring (CPS); ‘AC Vista’, CPS; AC Snowbird, CWWS; and AC Reed, SW Spring] and the American cultivar ‘Argent’, a HW Spring wheat. Canadian cultivars were obtained from the Crop Development Centre, University of Saskatchewan, Saskatoon, SK. Argent was obtained from the North Dakota Agricultural Experiment Station, North Dakota State University, Fargo, ND. Australian germplasm lines were obtained from the Australian Winter Cereals Collection, Tamworth, NSW.


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  		Table 1. Means for six agronomic and two quality traits for 79 wheat genotypes grown in Saskatchewan at two locations in 2000 and 2001.

 
The 79 wheat genotypes were grown at two locations in 2000 and 2001 at the University of Saskatchewan. The soil type was a Dark Brown Chernozem (Typic boroll) clay, clay loam at the Kernen Crop Research Farm (KCRF), and a Dark Brown Chernozem clay loam at the Seed Farm (SF). The experimental design was a randomized complete block design with two replications. Individual plots consisted of single rows, 3.6-m long and 0.3-m apart. Plots were sown on 2 May (SF) and 18 May (KCRF) in 2000 and 4 May (SF) and 28 May (KCRF) in 2001 at a rate of 250 seeds m-2 on fallow land. Seeds were treated with the systemic fungicide Vitavax Single Solution (Uniroyal Chemical Ltd., Elmira, ON, Canada; a.i. carbathiin) at the recommended rate. Fertilizer was drilled in with the seed at a rate of 7 kg ha-1 of N and 29 kg ha-1 of P. Weeds were controlled with the herbicides Buctril-M (Aventis CropScience Canada Co., Regina, SK, Canada; a.i. bromoxynil and MCPA) and Puma Super (Aventis CropScience Canada Co.; a.i. fenoxaprop-{rho}-ethyl) at recommended rates. Data were collected for days to heading, days to maturity, and plant height. Plots were combine-harvested at maturity and grain samples were dried using forced air driers. Plots were harvested immediately after maturity of the latest maturing genotype in an attempt to minimize kernel discoloration caused by weathering. Data were collected on grain yield, test weight, and 1000-kernel weight.

Seed samples were cleaned using a C-XT2 Dockage Tester (Simon-Day Ltd, Winnipeg, MB) before quality and grain color analysis. A 3.57-by-19.05-mm rectangular riddle and a 1.98-by-19.05-mm rectangular sieve were used to remove shriveled and cracked seed from grain samples. Kernel hardness was determined using a Brabender automatic micro hardness tester (C.W. Brabender Instruments, Inc., South Hackensack, NJ) as described by Miller et al. (1981). Color measurements were measured with a Hunterlab color difference meter (Color Quest 45/0, Hunter Associates Laboratory, Reston, VA) as L-, a-, and b-values. The L-value designates the lightness of the sample (100 = white and zero = black), a-values designate redness when positive or greenness when negative, and b-values designate yellowness when positive or blueness when negative. Total color difference ({Delta}E) was calculated as follows:

where {Delta} is the difference between AC Snowbird and sample readings. Disposable petri dishes (60-by-15-mm) were filled to half their capacity with {approx}10 g of grain. The dishes were covered with their corresponding lids before grain color determinations using the colorimeter. For grain samples treated with NaOH, the grain within each dish was treated with 15 mL of 1.25 mol L-1 NaOH for 1 h at 24°C based on the method outlined by Lamkin and Miller (1980). Excess NaOH solution was removed from each dish. Dishes were covered with their corresponding lids before grain color determinations. Grain (30 g) was ground to pass through a 1-mm screen on a cyclone sample mill (Udy, Fort Collins, CO) to produce whole meal. Meal samples were placed into resealable plastic bags (75-by-145-mm) for meal color determinations. Moisture content of meal was based on Method 44-15A (American Association of Cereal Chemists, 1995). Protein (N x 5.7) in meal was based on Method 46-30 using an FP-528 Protein/N analyzer (LECO Corp., St. Joseph, MI). Yellow pigment and lutein content in meal were based on Method 14-50 using 85% ethanol instead of water-saturated n-butanol as a solvent (McCaig et al., 1992). Yellow pigment and lutein contents (µg g-1) were calculated using absorbance values taken at 435 and 449 nm, respectively (Johnston et al., 1980).

Statistical analyses were conducted using Minitab Version 13 (Minitab Inc., State College, PA). For balanced ANOVAs, the statistical model included sources of variation due to Y, L, replication within L and Y, G, and interactions involving Y, L, and G. Year, location, replication, and their interactions were regarded as random effects, while G was a fixed effect. Bartlett's test (P = 0.05) was used to test for the homogeneity of variances. The Azallini and Cox test for crossover interactions was used to identify changes in rank order among genotypes at a significance level of P = 0.05 (Baker, 1988). Crossover G x E interactions were calculated for each quadruple combination of 79 genotypes using genotypic means averaged within Y and/or L. Pearson correlation coefficients were calculated among agronomic traits, grain protein, kernel grinding time, color values, yellow pigment, and lutein using means averaged across L and Y. Grand means and average differences between years were tested for significance (P = 0.05) using paired t-tests.


   RESULTS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
DISCUSSION
 REFERENCES
 
Wheat genotypes differed significantly in grain yield, heading, maturity, plant height, test weight, 1000-kernel weight, grain protein, and kernel grinding time (Table 1). Grand means for agronomic traits of HR (n = 7), HW (n = 64), and SW (n = 8) wheat genotypes indicated that the groups were similar in yield, heading, and test weight. Hard red genotypes were earlier maturing, taller, lower in test weight, and higher in grain protein relative to the HW or SW wheat genotypes. Hard white and SW genotypes were similar in maturity, height, kernel weight, and grain protein. Soft white genotypes possessed significantly longer kernel grinding times relative to the HR and HW genotypes.

Wheat genotypes differed significantly in grain and meal color values and yellow pigment and lutein contents (Table 2). For the 64 HW wheat genotypes, grain L-values varied from 42.5 to 49.0, a-values 7.0 to 8.2, b-values 15.5 to 17.6, and {Delta}E 0 to 4.3 units. Argent was significantly darker in grain lightness relative to AC Snowbird. AC Snowbird was significantly darker in grain lightness relative to AC Karma, AC Vista, and 35 HW wheat AUS accessions. Of the 60 HW wheat AUS accessions, ‘Sunelg’ was the darkest in grain color while ‘Kiata’ was the lightest. Yellow pigment (2.5 to 4.8 µg g-1) and lutein (1.8 to 3.7 µg g-1) varied among HW wheat genotypes. ‘Angas’, ‘Schomburgk’, ‘Cascades’, ‘Krichauff’, ‘Beulah’, ‘BT-Schomburgk’, and ‘Tern’ had significantly higher yellow pigment and lutein contents relative to the HW checks. Grand means for color values of HR, HW, and SW genotypes indicated that the groups had significantly different grain and meal color values, but similar in yellow pigment and lutein contents (Table 2). Soft white samples (without and with NaOH) were the lightest and most yellowish followed by HW samples and then HR samples. Hard white and HR samples (without NaOH) were similar in redness, but less reddish than SW samples. Hard red samples were redder than HW samples only after NaOH treatment. Hard white genotypes turned a creamy-yellow color after NaOH treatment while HR genotypes turned a brownish-orange color. Hard red samples had a larger {Delta}E with NaOH treatment than without NaOH treatment. Hard white meal samples were intermediate in lightness and redness, and more yellowish compared with HR and SW samples.


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  		Table 2. Means for grain (without and with NaOH) and meal color values, yellow pigment (YP), and lutein for 79 wheat genotypes.

 
Significant genotypic variation existed for colorimetric and pigmentation variables (Table 3). Nonsignificant G x L interactions were observed for grain color (without and with NaOH), meal color, yellow pigment, and lutein. Significant noncrossover G x Y interactions for grain L-values, grain color (NaOH treated), meal color, and yellow pigment were caused by changes in the magnitude of the response of genotypes across years (data not shown). Significant G x Y interactions were detected for grain redness, grain yellowness, and lutein. The Azallini and Cox test detected one significant crossover G x Y interaction for grain redness (Reeves and Condor), one for grain yellowness (‘Houtman’ and ‘Stiletto’), and 12 for lutein (‘Sunset’ with ‘Dollarbird’ and ‘Hartog’; ‘Carnamah’ with Chile Late, Dollarbird, Hartog, Kiata, ‘Leichhardt’, Sunelg, ‘Tasman’, CDC Teal, AC Reed, and Argent). The relative frequencies of crossover interactions detected for grain redness [(1 crossover quadruple/3081 possible quadruples) x 100 = 0.03%], grain yellowness (0.03%), and lutein (0.39%) were <0.5%. Significant noncrossover G x L x Y interactions for grain L- and b-values were caused by changes in the magnitude of the response of genotypes across Y and L (data not shown). The ANOVA generally remained unchanged for color variables when the seven red checks were removed from the analysis with the exception that the G x L x Y interaction for grain lightness became nonsignificant (data not shown).


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  		Table 3. Analysis of variance for grain (without and with NaOH) and meal color values, yellow pigment (YP), and lutein for 79 wheat genotypes.

 
Mean performance of HR, HW, and SW wheat genotypes grown in 2000 and 2001 were compared to determine if years differed in the study (Table 4). Average monthly precipitation during grain filling and maturation ranged from 82 (July) and 53 mm (August) in 2000 to 48 (July) and 6 mm (August) in 2001. Average precipitation values normally observed are 58 (July) and 37 mm (August), respectively. Thus, seed development and maturation of genotypes in this study occurred under dryer conditions in 2001 relative to 2000. Other monthly weather data were similar for 2000 and 2001 (data not shown). In 2001, lower grain yield, earlier maturities, shorter plant heights, lower kernel weights, and increased grain a-values (no NaOH) were observed for HR, HW, and SW genotypes. Higher test weights and increased grain a-values (without and with NaOH) were observed for HW and SW genotypes. These results suggest that kernels may tend to be smaller with a redder appearance in dry years.


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  		Table 4. Means of six agronomic traits, grain protein, kernel grinding time, grain (without and with NaOH) and meal color values, yellow pigment, and lutein for wheat genotypes.

 
Correlations among grain protein, kernel grinding time, grain and meal color, and yellow pigment and lutein contents were determined for 64 HW genotypes (Table 5). Weak correlations were observed among color variables and grain protein. Weak correlations were observed among color variables and kernel grinding time, except for grain b-values (no NaOH) and kernel grinding time (r = 0.74; P = 0.01). Grain color values (no NaOH) were poor indicators of meal color and yellow pigment and lutein contents. Grain L- and b-values (no NaOH) were highly correlated (r = 0.87; P = 0.01). Yellow pigment and lutein contents were poorly correlated with meal yellowness (r {approx} 0.42; P = 0.01). Yellow pigment and lutein contents were positively correlated (r = 0.99; P = 0.01). No correlations were observed among agronomic and color variables (data not shown).


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  		Table 5. Correlation coefficients among grain protein, kernel grinding time, grain (without and with NaOH treatment) and meal color values, yellow pigment (YP), and lutein for 64 hard white-seeded genotypes.

 

   DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
REFERENCES
 
Significant variation existed among the 79 wheat genotypes for agronomic traits, grain protein, grain hardness, and grain color values (without or with NaOH). Hard red, HW, and SW wheat genotypes were similar in grain yield, heading, and test weight; however, Canadian HR cultivars were earlier maturing, taller, lower in test weight, and higher in grain protein content. Soft white grain samples were the lightest and most yellowish, followed by HW and then HR samples. Caution should be exercised when selecting among HW genotypes because our results suggest that increases in grain b-values are associated with decreases in kernel hardness (r = 0.74; P = 0.01). Similarly, Peterson et al. (2001) cautioned breeders when selecting for increased grain L-values in white winter wheat as increased grain lightness was associated with decreased kernel hardness (r = -0.63; P = 0.01). Hard red genotypes had a larger {Delta}E with NaOH than without NaOH, confirming that NaOH treatment of grain samples is a useful method for distinguishing among HR, HW, and SW wheat genotypes. Our results indicate that genetic variation for grain color exists among HW wheat genotypes. Similarly, Wu et al. (1999) reported that significant variation for grain color existed among genotypes of HW winter wheat. The genetic variation described in our study should be useful in improving the grain lightness of AC Snowbird, a CWWS cultivar with intermediate grain lightness. Future research is needed to determine the number of genes, if any, controlling or influencing white seed coat color in HW wheat.

High yellow pigment content is viewed with disfavor in wheat required for milling and bread making (Moss, 1967). Weak correlations indicated that selecting for lower grain or meal b-values would not select for decreased yellow pigment content among HW wheat genotypes. We recommend the use of yellow pigment extraction, instead of grain or meal b-values, when distinguishing the least from the most yellowish meal entries in a breeding population. Specific attention should be given to populations derived from parents possessing high levels of carotenoids (e.g., Angas, Schomburgk, Cascades, Krichauff, Beulah, BT-Schomburgk, and Tern). Further research into that area is needed because the use of pigment extraction will result in an overall decreased efficiency in breeding programs. Parker and Langridge (2000) proposed using a sequence tagged site marker, linked to a major locus controlling flour color, to select breeding lines with low carotenoid levels.

Stability of grain color across L and Y in HW spring wheat is important to crop improvement programs. Lack of statistically significant G x L interactions for grain and meal color and yellow pigment and lutein contents indicated that the ranking of genotypes remained constant across locations. Significant noncrossover G x Y interactions for grain lightness, grain color (NaOH treated), meal color, and yellow pigment indicated that the ranking of genotypes remained constant across years. The Azallini and Cox test detected only one significant crossover G x Y interaction for grain redness, one for grain yellowness, and 12 for lutein content, indicating that the ranking of few genotypes changed significantly across years. Significant noncrossover G x L x Y interactions for grain L- and b-values were caused by changes in the magnitude of the response of genotypes across Y and L. On the basis of the environments tested within this study, the performance of genotypes was less consistent across years (G x Y) than across locations (G x L). The relative importance of the G x Y interaction suggests a need for testing grain color in more years rather than more locations. Similarly, Wu et al. (1999) reported that the ranking of winter wheat genotypes for grain color generally remained stable across environments. Peterson et al. (2001) suggested that environmentally induced variations in grain protein content, hardness, vitreousness, and kernel size and shape might all contribute to variation in visual grain color. Similarly, our results suggest that HW kernels tend to be smaller with a redder appearance in dry years. Genotypic differences in grain lightness are generally stable across environments. Consequently, improvements made in one environment are likely to be expressed in other environments.


   ACKNOWLEDGMENTS
 
Appreciation is expressed to Ken Jackle, Mike Grieman, Madje Cardenas, and Pam Lynn for their technical assistance. Thanks are extended to the Curator of Seed Gene Bank, Australian Winter Cereals Collection, RMB 944 Calala Lane, Tamworth NSW 2340, for providing small quantities of seed of the AUS accessions used in this study. Funding for this research was provided by a grant from the Saskatchewan Agriculture Development Fund.

Received for publication February 11, 2002.


   REFERENCES
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 




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