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作物学报 ›› 2026, Vol. 52 ›› Issue (6): 1593-1603.doi: 10.3724/SP.J.1006.2026.51106

• 作物遗传育种·种质资源·分子遗传学 •    下一篇

小麦Pod-A1、Pod-D1和Pod-2D位点等位变异对籽粒过氧化物酶活性的遗传效应分析

翟胜男1(), 曹新有1, 李豪圣1, 李吉虎1, 李法计1, 刘金栋2, 夏先春2, 吕莹莹1, 马瑞峰1, 王颖1, 耿洪伟3, 刘建军1,*()   

  1. 1 山东省农业科学院作物研究所 / 小麦育种全国重点实验室, 山东济南 250100
    2 中国农业科学院作物科学研究所 / 国家小麦改良中心, 北京 100081
    3 新疆农业大学农学院 / 生物技术重点实验室, 新疆乌鲁木齐 830052
  • 收稿日期:2025-12-16 接受日期:2026-02-27 出版日期:2026-06-12 网络出版日期:2026-03-05
  • 通讯作者: * 刘建军, E-mail: ljjsaas@163.com
  • 作者简介:翟胜男, E-mail: zsn19870322@163.com
  • 基金资助:
    山东省重点研发计划项目(2023CXPT079-2);山东省自然科学基金项目(ZR2023MC205);山东省自然科学基金项目(ZR2023MC155);泰山学者工程项目(tspd20221108);泰山学者工程项目(tsqn202408304);济南市“新高校20条”资助项目(202228067);财政部和农业农村部国家现代农业产业技术体系建设专项(Wheat,CARS-03-06)

Analysis of the genetic effects of allelic variation at the Pod-A1, Pod-D1, and Pod-2D loci on peroxidase activity in wheat grains

Zhai Sheng-Nan1(), Cao Xin-You1, Li Hao-Sheng1, Li Ji-Hu1, Li Fa-Ji1, Liu Jin-Dong2, Xia Xian-Chun2, Lyu Ying-Ying1, Ma Rui-Feng1, Wang Ying1, Geng Hong-Wei3, Liu Jian-Jun1,*()   

  1. 1 Crop Research Institute, State Key Laboratory of Wheat Improvement, Shandong Academy of Agricultural Sciences, Jinan 250100, Shandong, China
    2 Institute of Crop Sciences, National Wheat Improvement Center, Chinese Academy of Agricultural Sciences, Beijing 100081, China
    3 College of Agronomy, Key Laboratory of Agricultural Biological Technology, Xinjiang Agricultural University, Urumqi 830052, Xinjiang, China
  • Received:2025-12-16 Accepted:2026-02-27 Published:2026-06-12 Published online:2026-03-05
  • Contact: * Liu Jian-Jun, E-mail: ljjsaas@163.com
  • Supported by:
    Key R&D Program of Shandong Province(2023CXPT079-2);Natural Science Foundation of Shandong Province(ZR2023MC205);Natural Science Foundation of Shandong Province(ZR2023MC155);Taishan Scholar Program(tspd20221108);Taishan Scholar Program(tsqn202408304);Funding Program of “20 New Policies for Higher Education Institutions” of Jinan City(202228067);China Agriculture Research System of MOF and MARA(Wheat,CARS-03-06)

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摘要：

小麦籽粒过氧化物酶(peroxidase, POD)活性对加工品质和面制品色泽具有重要影响, 是品质改良的重要目标。本研究以151份国内外小麦品种(系)为材料, 利用分子标记系统检测Pod-A1、Pod-D1和Pod-2D位点的等位变异, 结合多年多点POD活性测定, 解析不同等位变异及其组合对籽粒POD活性的遗传效应。结果显示, 供试材料籽粒POD活性变异丰富, 平均值为674.39 U g-1 min-1, 变幅为431.30~954.81 U g-1 min-1, 变异系数为15.52%; 环境、基因型及其互作均对POD活性产生极显著影响(P < 0.01)。3个位点均显著影响籽粒POD活性, 优异等位变异分别为Pod-A1b、Pod-D1b和Pod-2D-GG。供试材料存在12种等位变异组合, 依据籽粒POD活性将其分为高、中、低POD活性基因组合类型, 三类间POD活性差异均达到显著水平(P < 0.05), 而每类内差异不显著。3个位点对籽粒POD活性的遗传效应依次为Pod-2D > Pod-A1 > Pod-D1。籽粒POD活性随着优异等位基因数目的增加而显著升高(R2 = 0.9681, P < 0.05), 聚合2~3个优异基因的小麦品种(系)籽粒POD活性显著高于聚合0~1个优异基因的材料(P < 0.05)。不同地区小麦品种(系)的籽粒POD活性、等位变异及其组合分布频率差异较大。筛选获得良星66、兰考24和济麦22等9份聚合3个优异等位基因且POD活性大于800 U g-1 min-1的优异种质。本研究为小麦籽粒POD活性分子标记辅助育种提供了理论依据和材料基础。

关键词: 小麦, 加工品质, 面制品颜色, 过氧化物酶, POD基因, 分子标记辅助育种

Abstract:

Peroxidase (POD) activity in wheat grains strongly influences processing quality and the color of wheat-based products, and is therefore an important target for quality improvement. In this study, 151 wheat varieties (lines) from domestic and international sources were used to systematically characterize allelic variation at the Pod-A1, Pod-D1, and Pod-2D loci using molecular markers. Together with multi-year, multi-location measurements of grain POD activity, we evaluated the effects of individual alleles and their combinations on POD activity. Grain POD activity showed substantial genetic variation, with a mean of 674.39 U g-1 min-1, a range of 431.30-954.81 U g-1 min-1, and a coefficient of variation of 15.52%. Environment, genotype, and their interaction all had highly significant effects on POD activity (P < 0.01). All three loci significantly affected grain POD activity, and the favorable alleles were Pod-A1b, Pod-D1b, and Pod-2D-GG, respectively. In total, 12 allelic combinations were identified and classified into high-, medium-, and low-activity combination types based on grain POD activity. POD activity differed significantly among the three types (P < 0.05), whereas no significant differences were detected within each type. The genetic effects of the three loci on grain POD activity followed the order Pod-2D > Pod-A1 > Pod-D1. Grain POD activity increased significantly with the number of favorable alleles (R2 = 0.9681, P < 0.05); varieties (lines) pyramiding 2-3 favorable alleles had significantly higher POD activity than those carrying 0-1 favorable allele (P < 0.05). Significant regional differences were observed in POD activity, allelic variation, and allelic-combination frequencies. Nine varieties, including Liangxing 66, Lankao 24, and Jimai 22, were identified as carrying all three favorable alleles and exhibiting POD activity above 800 U g-1 min-1. These findings provide a theoretical basis and germplasm resources for molecular marker-assisted breeding to optimize wheat grain POD activity.

Key words: Triticum aestivum, processing quality, wheat-based product color, peroxidase, POD genes, molecular marker-assisted breeding

表1

Pod-A1、Pod-D1和Pod-2D位点不同等位变异检测的分子标记信息"

基因
Gene		 分子标记
Molecular marker		 标记类型
Marker type		 引物序列
Primer sequence (5′-3′)		 片段大小
Fragment size (bp)		 等位变异
Allele		 参考文献
Reference
Pod-A1 POD-3A1 STS F: ACGGGAGACGACGAGAAGCAAAG 291 Pod-A1a [24]
R: TCGTGGAAGTGTAGGCGAAGA
POD-3A2 STS F: GTGGCGCAGGGCCTGTCA 766 Pod-A1b [24]
R: GTTGTCGAACACGTTGGGGGA
Pod-D1 POD-7D1 STS F: GCTTCGTCCAGGACGCGTT 540 Pod-D1a [26]
R: CGAGGAATGGGGGGTTGATG
POD-7D6 STS F: TGGGCATGGGGCTTCTGCA 640 Pod-D1b [26]
R: GCGAGGAATGGGGGGTTGATG
Pod-2D Excalibur_c95720_
329-KASP		 KASP FAM: GAAGGTGACCAAGTTCATGCTccggacatcatgagggcA Pod-2D-AA [27]
HEX: GAAGGTCGGAGTCAACGGATTccggacatcatgagggcG Pod-2D-GG
Common: GCTCCAACTCATCCAGCGTA Pod-2D-AG

表2

POD活性的变异分析"

环境 平均值 标准差 范围 变异系数
Environment Mean (U g-1 min-1) Standard deviation Range (U g-1 min-1) Coefficient of variation (%)
2013安阳 2013 AY 514.06 112.27 188.59-858.58 21.84
2013濉溪 2013 SX 507.03 110.64 299.90-852.22 21.82
2014安阳 2014 AY 781.33 129.88 477.48-1145.00 16.62
2014濉溪 2014 SX 894.15 154.29 530.16-1252.00 17.26
平均值 Mean 674.39 104.64 431.30-954.81 15.52

表3

POD活性的方差分析"

变异来源 自由度 平方和 均方 F值
Source of variation df Sum of squares Mean square F value
环境 Environment 3 31,444,477.25 10,481,492.42 1249.20**
重复 Replicate 4 641,996.62 160,499.16 19.13**
基因型 Genotype 150 11,633,333.18 77,555.55 9.24**
基因型×环境 Genotype×environment 449 5,841,814.51 13,010.72 1.55**
误差 Error 528 4,430,213.44 — —

表4

不同等位变异对POD活性的影响"

基因
Gene		 等位变异
Allele		 品种数
Number of varieties		 频率
Frequency
(%)		 POD活性
POD activity
(U g-1 min-1)		 标准差
Standard
deviation		 范围
Range
(U g-1 min-1)		 变异系数
Coefficient of
variation (%)
Pod-A1 Pod-A1a 72 47.68 633.59** 86.70 431.30-865.39 13.68
Pod-A1b 79 52.32 710.99 106.61 519.29-954.81 14.99
Pod-D1 Pod-D1a 64 43.54 654.57* 102.01 461.40-874.21 15.58
Pod-D1b 83 56.46 689.26 104.45 431.30-954.81 15.15
Pod-2D Pod-2D-AA 56 37.09 572.75** 48.21 431.30-651.87 8.42
Pod-2D-AG 22 14.57 652.01 15.01 628.88-676.29 2.30
Pod-2D-GG 73 48.34 758.47 75.24 597.67-954.81 9.92

表5

不同等位变异组合对POD活性的影响"

等位变异组合
Allelic combination		 品种数
Number of
varieties		 频率
Frequency
(%)		 POD活性
POD activity
(U g-1 min-1)		 标准差
Standard deviation		 范围
Range
(U g-1 min-1)		 变异系数
Coefficient of variation (%)
Pod-A1b/Pod-D1a/Pod-2D-GG 16 10.89 781.37 A 65.16 678.74-874.21 8.34
Pod-A1b/Pod-D1b/Pod-2D-GG 35 23.81 759.06 A 86.83 597.67-954.81 11.44
Pod-A1a/Pod-D1b/Pod-2D-GG 13 8.84 740.05 A 60.01 634.82-865.39 8.11
Pod-A1a/Pod-D1a/Pod-2D-GG 7 4.76 734.10 A 52.97 676.75-822.32 7.22
高POD活性基因组合类型 71 48.30 758.14 75.31 597.67-954.81 9.93
High POD activity allelic combination type
Pod-A1a/Pod-D1b/Pod-2D-AG 7 4.76 655.67 B 12.51 637.58-675.77 1.91
Pod-A1b/Pod-D1a/Pod-2D-AG 3 2.04 655.03 B 23.41 628.88-674.02 3.57
Pod-A1a/Pod-D1a/Pod-2D-AG 8 5.45 649.34 B 16.59 629.92-676.29 2.55
Pod-A1b/Pod-D1b/Pod-2D-AG 4 2.72 648.67 B 13.48 632.06-662.73 2.08
中POD活性基因组合类型 22 14.97 652.01 15.01 628.88-676.29 2.30
Medium POD activity allelic combination type
Pod-A1b/Pod-D1b/Pod-2D-AA 7 4.76 583.62 C 45.33 523.44-651.87 7.77
Pod-A1b/Pod-D1a/Pod-2D-AA 12 8.16 576.15 C 40.90 519.29-640.04 7.10
Pod-A1a/Pod-D1b/Pod-2D-AA 17 11.56 573.59 C 59.80 431.30-650.97 10.42
Pod-A1a/Pod-D1a/Pod-2D-AA 18 12.25 565.46 C 45.45 461.40-625.96 8.04
低POD活性基因组合类型 54 36.73 572.75 48.56 431.30-651.87 8.48
Low POD activity allelic combination type

图1

POD基因优异等位变异分布分析 A: 优异等位基因数目与POD活性的回归分析; B: 151份小麦品种(系)优异等位基因聚合情况。"

表6

优异基因数目对小麦品种(系) POD活性的影响"

优异基因数目
Number of
favorable alleles		 品种数
Number of varieties		 POD活性
POD activity
(U g-1 min-1)		 标准差
Standard
deviation		 范围
Range
(U g-1 min-1)		 变异系数
Coefficient of variation (%)
0 18 565.46 C 45.45 461.40-625.96 8.04
1 44 613.60 B 76.29 431.30-822.32 12.43
2 46 712.23 A 88.92 523.44-874.21 12.48
3 39 747.73 A 88.95 597.67-954.81 11.90

图2

不同地区小麦品种(系) POD活性比较分析 由于山西仅3个小麦品种, 因此未进行多重比较分析。不同字母表示不同地区间POD活性差异达显著水平(P < 0.05)。"

表7

不同地区小麦品种(系) POD基因等位变异及其组合分布频率"

等位变异及其组合
Allele or allelic combination		 分布频率 Distribution frequences (%)
总的
All		 国外
Foreign		 安徽
Anhui		 河北
Hebei		 河南
Henan		 山东
Shandong		 陕西
Shaanxi
Pod-A1a 47.68 52.63 50.00 57.14 50.00 16.67 65.22
Pod-A1b 52.32 47.37 50.00 42.86 50.00 83.33 34.78
Pod-D1a 43.54 47.37 57.14 42.86 37.50 40.00 47.83
Pod-D1b 56.46 52.63 42.86 57.14 62.50 53.33 47.83
Pod-2D-AA 37.09 42.11 35.71 42.86 29.17 16.67 65.22
Pod-2D-AG 14.57 15.79 42.86 28.57 12.50 6.67 4.35
Pod-2D-GG 48.34 42.11 21.43 28.57 58.33 76.67 30.43
Pod-A1b/Pod-D1a/Pod-2D-GG 10.89 5.26 0.00 7.14 16.67 21.43 0.00
Pod-A1b/Pod-D1b/Pod-2D-GG 23.81 10.53 7.14 14.29 25.00 46.43 22.72
Pod-A1a/Pod-D1b/Pod-2D-GG 8.84 26.32 0.00 7.14 12.50 0.00 4.55
Pod-A1a/Pod-D1a/Pod-2D-GG 4.76 0.00 14.29 0.00 4.17 7.14 4.55
高POD活性等位基因组合类型 48.30 42.11 21.43 28.57 58.34 75.00 31.82
High POD activity allelic combination type
Pod-A1a/Pod-D1b/Pod-2D-AG 4.76 0.00 7.14 14.29 8.33 0.00 0.00
Pod-A1b/Pod-D1a/Pod-2D-AG 2.04 5.26 14.29 0.00 0.00 0.00 0.00
Pod-A1a/Pod-D1a/Pod-2D-AG 5.45 5.26 14.29 7.14 4.17 3.57 4.55
Pod-A1b/Pod-D1b/Pod-2D-AG 2.72 5.26 7.14 7.14 0.00 3.57 0.00
中POD活性等位基因组合类型 14.97 15.78 42.86 28.57 12.50 7.14 4.55
Medium POD activity allelic combination type
Pod-A1b/Pod-D1b/Pod-2D-AA 4.76 5.26 7.14 0.00 6.25 3.57 4.55
Pod-A1b/Pod-D1a/Pod-2D-AA 8.16 15.79 14.29 14.29 2.08 7.14 9.09
Pod-A1a/Pod-D1b/Pod-2D-AA 11.56 5.26 14.29 14.29 10.42 3.57 18.18
Pod-A1a/Pod-D1a/Pod-2D-AA 12.25 15.79 0.00 14.29 10.42 3.57 31.82
低POD活性等位基因组合类型 36.73 42.10 35.72 42.87 29.17 17.85 63.64
Low POD activity allelic combination type
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