Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (10): 2061-2074.doi: 10.3864/j.issn.0578-1752.2026.10.001

• CROP GENETICS & BREEDING·GERMPLASM RESOURCES·MOLECULAR GENETICS • Previous Articles     Next Articles

Analysis of the Breeding Utilization Effects of Loci Related to Wheat Flour Color

ZHAI ShengNan1(), LÜ YingYing1, HE ZhongHu2, XIA XianChun2, MA RuiFeng1, WANG Ying1, LI FaJi1, CAO XinYou1, LI HaoSheng1, HAN Ran1, WANG XiaoLu1, LI JiHu1(), LIU JianJun1()   

  1. 1 Crop Research Institute, Shandong Academy of Agricultural Sciences, Jinan 250100
    2 Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081
  • Received:2025-10-30 Accepted:2025-12-26 Online:2026-05-16 Published:2026-05-20
  • Contact: LI JiHu, LIU JianJun

Abstract:

【Objective】The flour color is a crucial index for evaluating wheat quality. Identifying superior allelic variations at loci associated with flour color and clarifying their breeding utilization effects will provide a foundation for molecular marker-assisted breeding of wheat flour color.【Method】Using functional markers, molecular detection was conducted on 12 flour color-related loci in 166 wheat varieties (lines), including the Psy-A1, Pds-B1, Lcye-A1, Lcye-B1, Lox-B1, Ppo-A1, Ppo-B1, Ppo-D1, Pod-A1, Pod-D1, Pod-2D genes, and the 1B/1R translocation. Combined with the phenotypic measurements of flour color parameters (Yellow pigment content (YPC), L* value, a* value, b* value and whiteness), the influence of different allelic variations on flour color were analyzed to comprehensively and systematically evaluate the breeding utilization effects of each locus.【Result】The flour color of the tested materials exhibited a wide variation range. The average value of YPC was 1.18 μg·g-1, with a range of 0.57-2.96 μg·g-1; The average value of L* was 90.29, ranging from 87.12 to 92.16; the average a* value was -0.86, varying between -1.78 and -0.09; the average b* value was 8.83, with a range of 5.21-14.69; and the average whiteness was 86.78, spanning from 81.35 to 90.30. Environment, genotype, and the interaction between genotype and environment all significantly influenced flour color, with genotype exerting the strongest effect on the phenotypic variations. Psy-A1 and 1B/1R translocation significantly affected YPC, L* value, a* value, b* value and whiteness; Lcye-B1 significantly influenced YPC, a* value, b* value and whiteness; Pds-B1 and Lox-B1 significantly impacted L* value, b* value and whiteness; Pod-2D significantly influenced L* value and whiteness; Lcye-A1 significantly affected L* value; Ppo-A1 and Ppo-D1 significantly influenced a* value. These nine loci had substantial impacts on flour color and exhibited great potential for breeding applications. Wheat varieties (lines) containing Psy-A1b, Pds-B1b, Lcye-A1b, Lcye-B1b, Lox-B1a, Ppo-A1b, Ppo-D1a, Pod-2D-GG, Pod-2D-AG and non-1B/1R translocation exhibited high brightness and whiteness flour color. These genotypes were designated as superior alleles, with distribution frequencies of 34.94%, 20.48%, 97.59%, 66.27%, 26.38%, 50.91%, 57.23%, 48.80%, 15.06% and 51.20%, respectively. As the number of superior alleles increased, the L* value, a* value and whiteness gradually increased, while YPC and b* value gradually decreased. The materials pyramiding 7-8 superior genes exhibited the optimal flour color. There were significant differences in flour color and the allelic variation frequencies of related genes among wheat varieties (lines) from different regions. A total of 22 varieties, such as Zhengyin1, Zimai12 and Wanmai19, carried more than seven superior alleles and could be used as parental materials for breeding wheat varieties with high brightness and whiteness flour.【Conclusion】Psy-A1, Pds-B1, Lcye-A1, Lcye-B1, Lox-B1, Ppo-A1, Ppo-D1, Pod-2D and 1B/1R translocation significantly influence flour color and exhibit strong breeding practicability. Twenty-two wheat varieties, such as Zhengyin1, Zimai12 and Wanmai19, can be used as excellent parents for flour color improvement.

Key words: Triticum aestivum, flour color, molecular marker, superior alleles, breeding utilization

Table 1

Primer information of molecular markers for loci related to flour color"

基因
Gene
标记
Marker
引物序列
Sequence (5′-3′)
扩增片段
Fragment size (bp)
等位基因
Allele
表型
Phenotype
参考文献
Reference
Psy-A1 YP7A F: GGACCTTGCTGATGACCGAG 194 Psy-A1a 高YPC High YPC [6]
R: TGACGGTCTGAAGTGAGAATGA 231 Psy-A1b 低YPC Low YPC
Pds-B1 YP4B-1 F: AAGTTGGTGGGAGTTCCTGTC 652 Pds-B1b [7]
R: TGTCAGCAATGACAAAAGGATGCAT
YP4B-2 F: AAGTTGGTGGGAGTTCCTGTC 382 Pds-B1a [7]
R: GTGAACTCCTGAAACAGAAAGATTG
Lcye-A1 e-LCY3A-3 F: GCAGCAAATAACTAGACCGGGCA 537 Lcye-A1a [8]
R: TGAACTGGGGCACAAACCACG 309 & 230 Lcye-A1b
Lcye-B1 YP3B-1 F: GGATCGATCTCCTGAACAGGATGTC 635 Lcye-B1a [7]
R: CGGCGATGCTCCGATCAGTT No Lcye-B1b
Lox-B1 LOX16 F: CCATGACCTGATCCTTCCCTT 489 Lox-B1a 高LOX活性
High LOX activity
[10]
R: GCGCGGATAGGGGTGGT
LOX18 F: ACGATGTGAGTTGTGACTTGTGA 791 Lox-B1b 低LOX活性
Low LOX activity
[10]
R: GCGCGGATAGGGGTGC
Ppo-A1 PPO18 F: AACTGCTGGCTCTTCTTCCCA 685 Ppo-A1a 高PPO活性
High PPO activity
[13]
R: AAGAAGTTGCCCATGTCCGC 876 Ppo-A1b 低PPO活性
Low PPO activity
Ppo-B1 F-8 F: ACCTTGACTCCAGTGGGACC 400 Ppo-B1a 高PPO活性
High PPO activity
[14]
R: CCGAGTAGAAGTTGCCCAT 400 & 600 Ppo-B1b 低PPO活性
Low PPO activity
Ppo-D1 PPO16 F: TGCTGACCGACCTTGACTCC 713 Ppo-D1a 低PPO活性
Low PPO activity
[15]
R: CTCGTCACCGTCACCCGTAT
PPO29 F: TGAAGCTGCCGGTCATCTAC 490 Ppo-D1b 高PPO活性
High PPO activity
[15]
R: AAGTTGCCCATGTCCTCGCC
Pod-A1 POD-3A1 F: ACGGGAGACGACGAGAAGCAAAG 291 Pod-A1a 低POD活性
Low POD activity
[18]
R: TCGTGGAAGTGTAGGCGAAGA
POD-3A2 F: GTGGCGCAGGGCCTGTCA 766 Pod-A1b 高POD活性
High POD activity
[18]
R: GTTGTCGAACACGTTGGGGGA
Pod-D1 POD-7D1 F: GCTTCGTCCAGGACGCGTT 540 Pod-D1a 低POD活性
Low POD activity
[19]
R: CGAGGAATGGGGGGTTGATG
POD-7D6 F: TGGGCATGGGGCTTCTGCA 640 Pod-D1b 高POD活性
High POD activity
[19]
R: GCGAGGAATGGGGGGTTGATG
Pod-2D Excalibur_c95720
_329-KASP
FAM: GAAGGTGACCAAGTTCATGCTccggacatcatgagggcA Pod-2D-AA 低POD活性
Low POD activity
[20]
HEX: GAAGGTCGGAGTCAACGGATTccggacatcatgagggcG Pod-2D-GG 高POD活性
High POD activity
Common: GCTCCAACTCATCCAGCGTA Pod-2D-AG 中等POD活性
Moderate POD activity
1B/1R H2O F: GTTGGAAGGGAGCTCGAGCTG 1598 1B/1R 高YPC High YPC [22]
R: GTTGGGCAGAAAGGTCGACATC No 非1B/1R
Non-1B/1R
translocation
低YPC
Low YPC

Table 2

Phenotypic variation analysis of flour color-related traits in 166 wheat varieties (lines)"

性状
Trait
环境
Environment
平均值
Mean
标准差
Standard deviation
范围
Range
变异系数
Coefficient of variation (%)
黄色素含量
YPC (μg·g-1)
2020济南2020JN 1.15 0.42 0.53-2.80 36.83
2020德州2020DZ 1.19 0.45 0.51-2.76 37.69
2021济南2021JN 1.09 0.41 0.52-3.06 37.56
2021德州2021DZ 1.28 0.46 0.61-3.23 35.52
平均Mean 1.18 0.43 0.57-2.96 36.36
L*值
L* value
2020济南2020JN 90.06 1.05 87.24-92.25 1.17
2020德州2020DZ 90.02 1.07 86.69-92.06 1.18
2021济南2021JN 90.45 1.05 87.66-92.61 1.16
2021德州2021DZ 90.64 1.03 86.89-92.75 1.14
平均Mean 90.29 0.97 87.12-92.16 1.08
a*
a* value
2020济南2020JN -0.78 0.34 -1.62-0.01 43.57
2020德州2020DZ -0.80 0.37 -1.71-0.14 46.08
2021济南2021JN -0.94 0.38 -2.12-0.07 40.45
2021德州2021DZ -0.94 0.38 -1.91-0.02 40.10
平均Mean -0.86 0.35 -1.78-0.09 40.64
b*
b* value
2020济南2020JN 8.42 1.94 5.02-14.16 23.02
2020德州2020DZ 8.51 1.95 5.17-13.70 22.95
2021济南2021JN 9.14 2.28 5.46-15.76 24.96
2021德州2021DZ 9.26 2.26 5.11-15.16 24.42
平均Mean 8.83 2.04 5.21-14.69 23.15
白度
Whiteness
2020济南2020JN 86.90 1.90 81.84-90.37 2.18
2020德州2020DZ 86.80 1.92 82.00-90.38 2.21
2021济南2021JN 86.69 2.21 80.48-90.65 2.55
2021德州2021DZ 86.75 2.21 81.08-90.73 2.54
平均Mean 86.78 1.98 81.35-90.30 2.28

Table 3

Analyses of variance of flour color-related traits in 166 wheat varieties (lines)"

变异来源
Source of variation
自由度
Df
黄色素含量
YPC (μg·g-1)
L*值
L* value
a*值
a* value
b*值
b* value
白度
Whiteness
平方和
Sum of squares
F
F value
平方和
Sum of squares
F
F value
平方和
Sum of squares
F
F value
平方和
Sum of squares
F
F value
平方和
Sum of squares
F
F value
环境Environment 3 6.40 318.40** 88.18 299.04** 7.22 398.13** 176.15 567.65** 7.05 17.81**
重复Replicate 4 0.55 20.66** 2.14 5.44* 0.13 5.24* 2.25 5.43* 2.55 4.83*
基因型Genotype 165 238.31 215.66** 1230.80 75.89** 158.55 158.92** 5452.96 319.50** 5105.17 234.58**
基因型×环境Genotype×Environment 495 6.91 2.09** 196.77 4.04** 13.97 4.67** 378.18 7.39** 450.64 6.90**
误差Error 636 4.26 62.51 3.85 65.79 83.88

Table 4

The effect of different alleles on flour color"

基因
Gene
等位
基因
Allele
样本数
No. of sample
频率
Frequency
(%)
黄色素含量 YPC (μg·g-1) L*值 L* value a*值 a* value b*值 b* value 白度 Whiteness
平均值±标准差
Mean±SD
范围
Range
平均值±标准差
Mean±SD
范围
Range
平均值±标准差
Mean±SD
范围
Range
平均值±标准差
Mean±SD
范围
Range
平均值±标准差
Mean±SD
范围
Range
Psy-A1 a 108 65.06 1.34±0.39** 0.62-2.96 90.09±0.95** 87.12-92.02 -0.98±0.33** -1.78--0.09 9.57±1.89** 5.88-14.69 86.15±1.86** 81.36-89.94
b 58 34.94 0.87±0.30 0.57-1.92 90.67±0.91 88.46-92.16 -0.65±0.29 -1.51--0.09 7.45±1.56 5.21-12.51 88.01±1.60 83.81-90.30
Pds-B1 a 132 79.52 1.19±0.43 0.57-2.96 90.18±0.95** 87.12-92.16 -0.86±0.34 -1.78--0.09 9.00±2.02* 5.42-14.69 86.62±1.96* 81.36-90.18
b 34 20.48 1.13±0.43 0.57-2.20 90.72±0.97 88.46-92.13 -0.87±0.38 -1.67--0.13 8.12±1.84 5.21-12.51 87.48±1.94 83.16-90.30
Lcye-A1 a 4 2.41 1.04±0.31 0.68-1.42 89.13±0.11* 89.01-89.20 -0.67±0.46 -1.26--0.17 9.38±0.97 8.17-10.36 85.85±0.59 85.02-86.30
b 162 97.59 1.18±0.43 0.57-2.96 90.33±0.94 87.12-92.16 -0.87±0.35 -1.78--0.09 8.82±2.06 5.21-14.69 86.82±2.00 81.36-90.30
Lcye-B1 a 56 33.73 1.30±0.44** 0.58-2.96 90.16±0.89 88.34-91.97 -0.96±0.35** -1.78--0.09 9.39±2.07** 5.41-13.64 86.32±1.96* 82.90-90.30
b 110 66.27 1.12±0.41 0.57-2.47 90.36±1.01 87.12-92.16 -0.82±0.34 -1.75--0.09 8.55±1.98 5.21-14.69 87.04±1.95 81.36-90.30
Lox-B1 a 43 26.38 1.14±0.40 0.61-2.04 90.57±0.95* 88.43-92.13 -0.86±0.30 -1.65--0.30 8.23±1.79* 5.81-11.85 87.25±1.98* 83.10-90.19
b 120 73.62 1.20±0.44 0.57-2.96 90.20±0.96 87.12-92.16 -0.87±0.37 -1.78--0.09 9.02±2.04 5.21-14.69 86.62±1.96 81.36-90.30
Ppo-A1 a 81 49.09 1.23±0.45 0.58-2.96 90.30±0.92 88.63-92.11 -0.93±0.33* -1.78--0.40 8.99±2.05 5.41-14.69 86.70±1.95 81.36-90.30
b 84 50.91 1.13±0.41 0.57-2.20 90.27±1.04 87.12-92.16 -0.81±0.34 -1.65--0.09 8.70±2.05 5.21-13.26 86.87±2.01 83.10-90.30
Ppo-B1 a 133 80.12 1.18±0.44 0.57-2.96 90.25±0.97 87.12-92.16 -0.86±0.36 -1.78--0.09 8.87±2.05 5.21-14.69 86.74±1.96 81.36-90.30
b 33 19.88 1.17±0.36 0.69-2.21 90.44±0.99 88.86-92.11 -0.88±0.32 -1.75--0.30 8.66±2.06 5.85-13.93 87.02±2.05 82.53-90.06
Ppo-D1 a 95 57.23 1.14±0.40 0.57-2.47 90.22±0.95 87.12-91.87 -0.81±0.32* -1.56---0.09 8.76±2.05 5.21-14.69 86.80±1.97 81.36-90.30
b 71 42.77 1.23±0.46 0.58-2.96 90.39±1.01 88.46-92.16 -0.92±0.37 -1.78--0.09 8.93±2.05 5.41-13.64 86.80±2.01 82.90-90.30
Pod-A1 a 72 47.68 1.16±0.40 0.57-1.96 90.38±0.98 88.46-92.16 -0.86±0.35 -1.51--0.09 8.65±2.00 5.21-12.51 86.98±1.94 83.75-90.30
b 79 52.32 1.18±0.41 0.60-2.47 90.18±0.96 87.12-92.02 -0.86±0.34 -1.67--0.09 8.99±1.98 5.41-14.69 86.61±1.93 81.36-90.30
Pod-D1 a 64 43.54 1.19±0.39 0.61-2.47 90.26±0.98 87.12-91.97 -0.86±0.33 -1.56--0.09 8.86±1.95 5.41-14.69 86.76±1.92 81.36-90.30
b 83 56.46 1.15±0.42 0.57-2.20 90.32±0.98 88.34-92.16 -0.85±0.36 -1.67--0.09 8.73±2.03 5.21-13.26 86.88±1.96 83.10-90.30
Pod-2D AA 60 36.14 1.16±0.48 0.57-2.96 90.05±0.83* 88.46-92.16 -0.85±0.40 -1.78--0.09 9.04±1.93 5.21-13.64 86.50±1.76* 82.90-90.30
AG 25 15.06 1.06±0.49 0.58-2.47 90.65±1.02 88.63-92.13 -0.81±0.34 -1.75--0.38 8.09±2.44 5.42-14.69 87.55±2.34 81.36-90.19
GG 81 48.80 1.23±0.37 0.60-2.20 90.37±0.95 87.12-92.02 -0.89±0.31 -1.67--0.09 8.91±1.97 5.53-13.26 86.78±1.97 83.10-89.96
1B/1R N 85 51.20 0.97±0.30** 0.57-2.20 90.47±1.01* 87.12-92.16 -0.72±0.28** -1.67--0.09 8.03±1.75** 5.21-13.26 87.45±1.85** 83.17-90.30
Y 81 48.80 1.39±0.44 0.57-2.96 90.15±0.84 88.34-92.02 -1.02±0.35 -1.78--0.09 9.67±2.01 5.33-14.69 86.11±1.88 81.36-90.08

Fig. 1

Distribution of superior alleles of nine important genes for flour color A: The aggregation of superior alleles in 166 wheat varieties (lines); B: The aggregation of superior alleles in wheat varieties (lines) from different regions. Different letters indicate significant differences of superior gene pyramid number in wheat varieties (lines) from different regions at P<0.05 probability level"

Table 5

Excellent germplasm with more than seven superior alleles"

优异等位基因数目
Number of superior alleles
品种
Varieties (lines)
9 郑引1号、淄麦12 Zhengyin1, Zimai12
8 皖麦19、皖麦50、皖麦52、阿夫、临麦4号、烟农15 Wanmai19, Wanmai50, Wanmai52, Funo, Linmai4, Yannong15
7 矮丰3号、丰产3号、阜936、皖麦53、宿0663、宿农6号、良星99、烟农19、临麦2号、鲁麦23、鲁麦6号、石家庄15、小偃22、小偃81
Aifeng3, Fengchan3, Fu936, Wanmai53, Su0663, Sunong6, Liangxing99, Yannong19, Linmai2, Lumai23, Lumai6, Shijiazhuang15, Xiaoyan22, Xiaoyan81

Fig. 2

Linear regression between the number of superior alleles and flour color A: Correlation analysis of the number of superior alleles with yellow pigment content (YPC); B: Correlation analysis of the number of superior alleles with L* value; C: Correlation analysis of the number of superior alleles with a* value; D: Correlation analysis of the number of superior alleles with b* value; E: Correlation analysis of the number of superior alleles with whiteness"

Table 6

Effects of superior gene pyramid number on flour color of 166 wheat varieties (lines)"

优异等位基因数目
Number of superior alleles
品种数
Number of sample
黄色素含量
YPC (μg·g-1)
L*值
L* value
a*值
a* value
b*值
b* value
白度
Whiteness
1 4 1.95A 89.88C -1.36D 11.65A 84.49D
2 10 1.34B 89.79C -1.00C 10.01B 85.63CD
3 22 1.31B 90.06C -0.96BC 9.53B 86.17C
4 44 1.32B 90.26BC -0.97BC 9.26B 86.47C
5 37 1.11BC 90.22BC -0.83ABC 8.79B 86.77C
6 27 1.03BC 90.28BC -0.73ABC 8.32BC 87.16BC
7 14 0.85C 90.95AB -0.64A 7.03CD 88.50AB
8 6 0.83C 91.23A -0.67AB 6.68D 88.95A
9 2 0.83 90.87 -0.63 6.98 88.50

Table 7

Comparison of flour color of wheat varieties (lines) from different regions"

地区
Region
黄色素含量
YPC (μg·g-1)
L*值
L* value
a*值
a* value
b*值
b* value
白度
Whiteness
国外Foreign 1.33A 89.85BC -0.87AB 9.56A 85.98B
中国安徽Anhui, China 1.01B 91.14A -0.82AB 7.51B 88.32A
中国河北Hebei, China 0.99B 89.80C -0.69A 8.77A 86.49B
中国河南Henan, China 1.29A 90.35BC -0.95B 9.07A 86.68B
中国山东Shandong, China 1.19AB 90.42B -0.94B 8.92A 86.83B
中国陕西Shaanxi, China 1.01B 90.13BC -0.70A 8.44AB 86.94B

Table 8

Distribution frequences of alleles of flour color important genes in wheat varieties (lines) from different regions"

基因
Gene
等位基因
Allele
分布频率Distribution frequences (%)
总计
Total
国外
Foreign
中国安徽
Anhui, China
中国河北
Hebei, China
中国河南
Henan, China
中国山东
Shandong, China
中国陕西
Shaanxi, China
Psy-A1 a 65.06 90.91 26.67 42.86 81.48 72.73 40.00
b 34.94 9.09 73.33 57.14 18.52 27.27 60.00
Pds-B1 a 79.52 90.91 80.00 78.57 83.33 66.67 80.00
b 20.48 9.09 20.00 21.43 16.67 33.33 20.00
Lcye-A1 a 2.41 9.09 0.00 7.14 1.85 0.00 0.00
b 97.59 90.91 100.00 92.86 98.15 100.00 100.00
Lcye-B1 a 33.73 27.27 13.33 21.43 51.85 39.39 16.00
b 66.27 72.73 86.67 78.57 48.15 60.61 84.00
Lox-B1 a 26.38 9.09 53.33 7.14 22.64 51.61 16.00
b 73.62 90.91 46.67 92.86 77.36 48.39 84.00
Ppo-A1 a 49.09 68.18 20.00 57.14 71.70 24.24 32.00
b 50.91 31.82 80.00 42.86 28.30 75.76 68.00
Ppo-D1 a 57.23 54.55 33.33 71.43 74.07 51.52 36.00
b 42.77 45.45 66.67 28.57 25.93 48.48 64.00
Pod-2D AA 36.14 45.45 33.33 42.86 25.93 18.18 64.00
GG 48.80 40.91 20.00 28.57 61.11 72.73 32.00
AG 15.06 13.64 46.67 28.57 12.96 9.09 4.00
1B/1R N 51.20 68.18 80.00 28.57 31.48 63.64 64.00
Y 48.80 31.82 20.00 71.43 68.52 36.36 36.00
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