Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (21): 4710-4728.doi: 10.3864/j.issn.0578-1752.2026.21.002

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

Identification of Excellent High Temperature Tolerant Resources in Wheat and Their Association Analysis with Agronomic Traits

YIN XiaoKang1(), FENG JiaYi1(), LI XuKang1, ZHANG ZhaoXu1, MA Chi1, WANG RuoYu1, BAI JiaQian1, LAN LuoBing1, ZHANG LiangFeng1, LIU YiKe2, ZHU ZhanWang2, DING FuGong2, LIU DaTong3, WANG ShuPing1(), FANG ZhengWu1(), ZHANG YingXin1()   

  1. 1 College of Agriculture, Yangtze University/Key Laboratory of Green and Efficient Crop Production in the Middle Reaches of the Yangtze River, Ministry of Agriculture and Rural Affairs/Hubei Key Laboratory of Waterlogging Disaster and Agricultural Use of Wetland, Jingzhou 434025, Hubei
    2 Institute of Food Crops, Hubei Academy of Agricultural Sciences/Hubei Key Laboratory of Food Crop Germplasm and Genetic Improvement, Wuhan 430064
    3 Lixiahe Institute of Agricultural Sciences in Jiangsu Province/Key Laboratory of Wheat Biology and Genetic Improvement for Low & Middle Yangtze Valley, Ministry of Agriculture and Rural Affairs, Yangzhou 225007, Jiangsu
  • Received:2026-04-12 Accepted:2026-06-26 Online:2026-11-01 Published:2026-11-06
  • Contact: WANG ShuPing, FANG ZhengWu, ZHANG YingXin

Abstract:

【Objective】To address the wheat yield loss caused by high temperature stress during grain filling, this study was conducted to clarify the effects of post-anthesis high temperature stress on plant phenotype, agronomic traits, yield and yield components of different wheat genotypes, establish a comprehensive evaluation system for wheat high temperature tolerance suitable for winter wheat regions in the middle and lower reaches of the Yangtze River, screen out high temperature tolerant wheat varieties, and provide identification methods and high quality germplasm resources for wheat high temperature tolerance breeding. 【Method】30 winter wheat varieties were used as experimental materials. On the 7th day after anthesis, two treatments were set for 7 days respectively, including the high temperature treatment (38 ℃ daytime/20 ℃ nighttime) and the control treatment (25 ℃ daytime/20 ℃ nighttime). After treatment, the relative chlorophyll content (SPAD value) of flag leaves, activities of ascorbate peroxidase (APX), peroxidase (POD), superoxide dismutase (SOD) and catalase (CAT), as well as the contents of malondialdehyde (MDA), hydrogen peroxide (H2O2) and superoxide anion radical (O2-·) were measured. After harvest, grain length, grain width, grain weight per spike, thousand-grain weight and yield per plant were determined. Analysis of variance (ANOVA) was firstly adopted to eliminate indicators with no significant differences. All raw data were standardized by the high temperature tolerance coefficient (HTC). Principal component analysis (PCA) combined with the membership function method were performed to calculate the comprehensive D value for evaluating wheat high temperature tolerance. Redundancy analysis (RDA) and stepwise regression were finally used to construct the prediction model. 【Result】High temperature stress during grain filling stage caused varying degrees of leaf yellowing and grain shrinkage in all tested wheat genotypes. Under high temperature treatment, all index varied in the range of -66.82% to 210.07%, with the content of O2-· showing the greatest increase (210.07%) and APX activity showing the greatest decreased (-66.82%). Significant differences in HTC values were observed among different wheat cultivars, with coefficients of variation (CV) ranging from 9.522% to 115.003%. APX activity had the highest CV (115.003%), and grain width had the lowest CV (9.522%). PCA was performed on 7 indicators that met the criteria of MSA > 0.5 and KMO ≥ 0.6. 4 independent comprehensive new components were extracted, which explained 82.814% of the total variance in the original data. Furthermore, cluster analysis based on the comprehensive evaluation value D classified the 30 cultivars into three categories: 5 high temperature tolerant cultivars, 12 moderately high temperature tolerant cultivars, and 13 high temperature sensitive cultivars. The optimal regression model for wheat high temperature tolerance evaluation was established via redundancy analysis and stepwise regression: D=-0.111+0.259X8+0.136X12+0.186X2+0.111X1+0.137X11+0.12X10. The regression values (RV) of each variety were highly correlated with the D values (r=0.996). Flag leaf relative chlorophyll content (X8), yield per plant (X12), POD activity (X2), APX activity (X1), thousand-grain weight (X11) and grain weight per spike (X10) were identified as core indicators for evaluating wheat high temperature tolerance. 【Conclusion】High temperature stress during grain filling stage significantly constrained the growth and development of wheat. Using PCA and cluster analysis, 5 high temperature tolerant wheat cultivars were screened out. A comprehensive evaluation model for wheat high temperature tolerance applicable to the middle and lower reaches of the Yangtze River winter wheat was established, which can be effectively used to evaluate the high temperature tolerance of different wheat varieties at grain filling stage.

Key words: Triticum aestivum L., grain filling stage, high temperature tolerance, comprehensive evaluation, germplasm identification

Table 1

Effects of high temperature treatment during grain filling on various indexes of 30 wheat varieties"

性状
Trait
对照Control 高温处理High temperature treatment 变幅
Range (%)
均值±标准差Mean±SD 均值±标准差Mean±SD
超氧化物歧化酶活性SOD activity (U·g-1·h-1) 761.88±65.36b 896.18±108.88a 17.63
过氧化物酶活性POD activity (U·g-1·min-1) 361.79±76.06a 217.17±87.66b -39.97
过氧化氢酶活性CAT activity (U·g-1·min-1) 593.87±62.71a 539.44±49.96b -9.17
抗坏血酸过氧化物酶活性APX activity (U·g-1) 2.20±1.19a 0.73±0.96b -66.82
丙二醛含量MDA content (μmol·g-1) 29.73±5.89b 42.16±23.80a 41.81
过氧化氢含量H2O2 content (μmol·g-1) 5.53±2.06a 3.80±1.95b -31.28
超氧阴离子自由基含量O2-· content (μg·g-1) 22.74±7.96b 70.51±43.12a 210.07
旗叶叶绿素相对含量Flag leaf SPAD 56.33±4.16a 46.40±12.51b -17.63
穗粒重Grain weight per spike (g) 1.14±0.30a 0.66±0.17b -42.11
粒长Grain length (mm) 5.97±0.45a 6.04±0.39a 1.27
粒宽Grain width (mm) 2.73±0.26a 2.53±0.22b -7.22
千粒重Thousand-grain weight (g) 37.27±7.15a 26.57±7.02b -28.71
单株产量Yield per plant (g) 4.46±1.23a 2.29±0.73b -48.65

Fig. 1

Phenotypic and grain changes in wheat under high temperature stress for 7 days during the grain filling stage CK: Control; HT: High temperature treatment. The same as below. Scale bar for plant height: 25 cm; Scale bar for wheat grain: 5 mm"

Fig. 2

Analysis on dispersion degree of various indicators in wheat under high temperature stress and control during grain filling"

Table 2

Heat tolerance coefficient (HTC) of each single index of different wheat cultivars"

品种Variety X1 X2 X3 X4 X5 X6 X7 X8 X9 X10 X11 X12
楚襄40 Chuxiang 40 0.051 0.374 1.084 0.992 0.790 1.100 1.774 0.410 1.267 0.495 0.674 0.424
楚襄1588 Chuxiang1588 0.404 0.076 0.967 1.172 1.358 0.876 1.159 0.432 0.861 0.431 0.590 0.503
襄麦2305 Xiangmai 2305 0.510 0.182 0.936 0.910 0.557 0.837 0.506 0.687 0.841 0.499 0.449 0.367
华麦216 Huamai 216 0.210 0.632 1.416 0.769 0.506 0.759 2.726 0.652 0.832 0.500 0.606 0.467
垦麦58 Kenmai 58 0.269 0.673 1.299 0.950 0.644 0.617 1.787 0.683 0.858 0.373 0.545 0.303
鄂麦017 Emai 017 0.187 0.696 0.994 0.908 0.346 0.931 3.104 0.751 0.761 0.470 0.487 0.321
鄂麦180 Emai180 0.231 0.768 1.274 0.825 0.413 0.832 1.493 0.922 0.989 0.711 0.962 0.875
长麦2号 Changmai 2 0.088 0.796 1.230 1.107 0.245 1.274 3.690 0.986 0.929 0.670 0.664 0.515
鄂麦229 Emai 229 0.658 0.612 1.051 0.976 0.671 1.109 1.709 0.937 0.903 0.432 0.685 0.378
长辐麦10号 Changfumai 10 0.050 0.670 1.306 0.873 1.162 2.251 3.396 0.655 0.944 0.612 0.754 0.510
H51995 0.055 0.681 1.714 0.826 1.066 1.638 4.077 0.887 0.932 0.478 0.664 0.435
鄂麦026 Emai 026 0.072 0.432 1.074 0.674 0.851 1.147 3.544 0.861 0.968 0.584 0.744 0.292
泰麦1号 Taimai 1 0.977 0.683 1.187 1.030 1.036 0.976 4.176 1.070 1.077 0.820 0.957 0.615
楚科316 Chuke 316 0.165 0.713 1.077 0.924 0.456 0.974 3.338 1.011 0.933 0.479 0.750 0.555
长麦007 Changmai 007 0.254 1.561 1.260 1.009 0.655 1.137 2.422 0.924 0.933 0.813 0.761 1.012
轮选112 Lunxuan112 0.102 0.217 0.935 0.827 0.685 1.078 1.604 0.962 0.973 0.865 0.823 0.906
创红麦2号 Chuanghongmai 2 0.038 0.766 1.199 0.834 0.538 1.731 5.338 0.834 0.853 0.708 0.583 0.826
华麦1913 Huamai 1913 1.159 0.776 1.274 0.977 0.553 1.112 1.697 0.994 0.935 0.460 0.691 0.460
华麦230 Huamai 230 0.093 1.974 1.279 0.843 0.780 1.009 6.323 0.910 0.950 0.470 0.764 0.352
鄂麦008 Emai 008 0.946 0.948 1.121 1.008 0.793 1.022 3.665 0.942 0.980 0.768 0.821 0.566
华麦243 Huamai 243 0.053 0.362 1.110 0.830 0.546 1.120 6.354 1.039 0.973 0.629 0.827 0.399
长辐麦8号 Changfumai 8 0.144 0.627 1.309 0.950 1.134 1.042 2.741 0.931 0.918 0.933 0.708 0.622
鄂810619 E810619 0.227 0.787 1.485 0.814 0.837 0.999 1.616 0.989 0.970 0.375 0.890 0.456
鄂麦235 Emai 235 1.118 0.498 0.958 1.024 1.046 1.315 3.467 0.757 0.886 0.698 0.865 0.698
襄麦2355 Xiangmai 2355 0.022 0.787 1.490 0.872 0.635 3.145 4.131 0.744 0.932 0.575 0.742 0.602
华麦2260 Huamai 2260 1.321 0.743 1.131 0.719 0.630 0.810 0.701 0.948 0.904 0.963 0.701 0.811
金广天麦2号Jinguangtianmai 2 0.065 0.561 1.186 1.110 0.690 2.393 3.079 0.843 0.848 0.241 0.423 0.212
长麦5号Changmai 5 0.036 0.379 1.101 1.139 0.555 4.139 2.742 0.873 0.974 0.610 0.761 0.560
鄂麦30 Emai 30 0.026 0.250 1.203 0.744 0.357 5.380 7.869 0.486 0.879 0.689 0.719 0.435
华麦1912 Huamai 1912 0.804 0.190 0.912 0.892 0.388 2.886 3.792 0.416 0.992 0.774 0.881 0.774
平均值Average 0.344 0.647 1.185 0.918 0.697 1.521 3.134 0.818 0.933 0.604 0.716 0.542
标准差STDEV 0.396 0.381 0.186 0.125 0.270 1.084 1.712 0.190 0.089 0.178 0.136 0.200
变异系数CV(%) 115.003 58.890 15.653 13.640 38.775 71.260 54.630 23.268 9.522 29.394 18.988 36.945

Table 3

KMO and Bartlett's sphericity test of heat tolerance coefficient for each index"

变量筛选阶段
Screening stage
Measure of Sampling Adequacy度量值
Measure of Sampling Adequacy measurement value
Kaiser-Meyer-Olkin度量值
Kaiser-Meyer-Olkin measurement value
Bartlett球形度检验
Bartlett’s
sphericity test
X1 X2 X3 X4 X5 X6 X7 X8 X9 X10 X11 X12
Bef. 0.452 0.411 0.380 0.209 0.199 0.379 0.337 0.350 0.290 0.449 0.434 0.384 0.37 <0.001
Aft. 0.702 0.558 0.583 - - - - 0.633 - 0.643 0.819 0.628 0.65 <0.001

Table 4

Eigenvectors and percentage of accumulated contribution of principal components"

主成分Principle factor CI 1 CI 2 CI 3 CI 4
特征值Eigen value 2.499 1.772 0.929 0.597
贡献率Contribution ratio (%) 35.705 25.316 13.269 8.523
累计贡献率Cumulative contribution ratio (%) 35.705 61.022 74.291 82.814
特征向量Eigen vector X1 0.302 -0.246 0.726 0.245
X2 0.150 0.587 0.173 0.552
X3 -0.062 0.604 -0.274 0.042
X8 0.280 0.438 0.415 -0.462
X10 0.541 -0.158 -0.244 0.128
X11 0.481 0.069 -0.132 -0.549
X12 0.530 -0.088 -0.345 0.320

Table 5

The value of each variety’s comprehensive index (CI), index weight, u(Xj), D value, RV and comprehensive valuation"

品种
Variety
CI 1 CI 2 CI 3 CI 4 u1 u2 u3 u4 D值
D value
RV 综合评价
Comprehensive evaluation
楚襄40 Chuxiang 40 -1.069 -1.037 -1.049 0.385 0.258 0.269 0.121 0.591 0.274 0.280 高温敏感型High temperature-sensitive type
楚襄1588 Chuxiang1588 -1.105 -1.811 -0.256 0.543 0.248 0.095 0.332 0.626 0.254 0.261 高温敏感型High temperature-sensitive type
襄麦2305 Xiangmai 2305 -1.197 -1.425 0.910 0.535 0.224 0.182 0.643 0.624 0.319 0.329 高温敏感型High temperature-sensitive type
华麦216 Huamai 216 -0.844 0.374 -0.601 0.777 0.317 0.586 0.241 0.676 0.424 0.405 高温敏感型High temperature-sensitive type
垦麦58 Kenmai 58 -1.408 0.276 0.316 0.629 0.168 0.564 0.485 0.644 0.389 0.381 高温敏感型High temperature-sensitive type
鄂麦017 Emai 017 -1.226 -0.380 0.682 0.739 0.216 0.417 0.582 0.668 0.383 0.401 高温敏感型High temperature-sensitive type
鄂麦180 Emai180 1.367 0.501 -1.052 -0.659 0.903 0.615 0.120 0.364 0.634 0.632 中等耐高温型
Moderately high temperature-tolerant type
长麦2号Changmai 2 0.027 0.636 -0.099 -0.165 0.548 0.645 0.374 0.472 0.542 0.544 中等耐高温型
Moderately high temperature-tolerant type
鄂麦229 Emai 229 -0.393 -0.152 1.624 -0.564 0.437 0.468 0.833 0.385 0.504 0.516 中等耐高温型
Moderately high temperature-tolerant type
长辐麦10号Changfumai 10 -0.265 0.196 -1.097 0.102 0.470 0.546 0.108 0.530 0.442 0.435 高温敏感型High temperature-sensitive type
H51995 -0.718 1.684 -0.762 -0.296 0.350 0.881 0.198 0.443 0.498 0.459 中等耐高温型
Moderately high temperature-tolerant type
鄂麦026 Emai 026 -0.516 -0.214 0.097 -1.467 0.404 0.454 0.426 0.189 0.401 0.412 高温敏感型High temperature-sensitive type
泰麦1号Taimai 1 1.624 0.111 1.109 -1.124 0.971 0.527 0.696 0.263 0.718 0.715 耐高温型High temperature-tolerant type
楚科316 Chuke 316 -0.014 0.321 0.413 -0.922 0.537 0.574 0.511 0.307 0.521 0.537 中等耐高温型
Moderately high temperature-tolerant type
长麦007 Changmai 007 1.556 1.189 -0.799 2.265 0.953 0.770 0.188 1.000 0.779 0.786 耐高温型High temperature-tolerant type
轮选112 Lunxuan112 1.314 -1.002 -1.082 -1.084 0.889 0.277 0.113 0.272 0.514 0.530 中等耐高温型
Moderately high temperature-tolerant type
创红麦2号
Chuanghongmai 2
0.274 0.125 -1.033 1.313 0.613 0.530 0.126 0.793 0.528 0.529 中等耐高温型
Moderately high temperature-tolerant type
华麦1913 Huamai 1913 0.100 0.405 2.251 0.190 0.567 0.593 1.000 0.549 0.643 0.632 中等耐高温型
Moderately high temperature-tolerant type
华麦230 Huamai 230 -0.194 2.214 0.692 1.259 0.489 1.000 0.585 0.781 0.691 0.711 耐高温型High temperature-tolerant type
鄂麦008 Emai 008 1.083 0.046 1.284 0.292 0.828 0.513 0.742 0.571 0.691 0.696 耐高温型High temperature-tolerant type
华麦243 Huamai 243 0.065 0.077 0.037 -2.335 0.558 0.519 0.410 0.000 0.465 0.474 高温敏感型High temperature-sensitive type
长辐麦8号
Changfumai 8
0.725 0.318 -0.925 -0.002 0.733 0.574 0.154 0.507 0.568 0.556 中等耐高温型
Moderately high temperature-tolerant type
鄂810619 E810619 -0.123 1.494 0.079 -1.581 0.508 0.838 0.422 0.164 0.560 0.546 中等耐高温型
Moderately high temperature-tolerant type
鄂麦235 Emai 235 1.101 -1.252 1.050 0.094 0.833 0.221 0.680 0.528 0.590 0.599 中等耐高温型
Moderately high temperature-tolerant type
襄麦2355 Xiangmai 2355 -0.150 0.940 -1.273 0.287 0.501 0.714 0.062 0.570 0.503 0.483 中等耐高温型
Moderately high temperature-tolerant type
华麦2260 Huamai 2260 1.734 -0.589 1.304 1.509 1.000 0.370 0.748 0.836 0.750 0.741 耐高温型High temperature-tolerant type
金广天麦2号Jinguangtianmai 2 -2.041 0.317 0.884 0.047 0.000 0.573 0.636 0.518 0.331 0.335 高温敏感型High temperature-sensitive type
长麦5号Changmai 5 -0.005 -0.269 -0.545 -1.136 0.539 0.442 0.256 0.261 0.435 0.443 高温敏感型High temperature-sensitive type
鄂麦30 Emai 30 -0.575 -0.863 -1.505 -0.105 0.388 0.308 0.000 0.485 0.312 0.304 高温敏感型High temperature-sensitive type
华麦1912 Huamai 1912 0.874 -2.233 -0.655 0.474 0.772 0.000 0.226 0.611 0.432 0.440 高温敏感型High temperature-sensitive type
权重ω 0.431 0.306 0.160 0.103

Fig. 3

Systematic clustering diagram of 30 wheat varieties"

Fig. 4

Redundancy analysis of 30 wheat cultivars and correlation heatmap between comprehensive evaluation value D and selected indicators A: RDA ordination analysis of heat tolerance coefficient of agronomic traits and physiological indicators in wheat under high temperature stress. The included angle between two explanatory variables represents their correlation: an acute angle indicates a positive correlation, and an obtuse angle indicates a negative correlation. The length of arrows indicates the primary and secondary relationships influencing D value, longer arrows represent stronger explanatory power, and shorter ones correspond to weaker explanatory power. B: Correlation analysis of six indices related to high temperature tolerance coefficient. Red indicates positive correlation, Blue indicates negative correlation. A larger circle diameter represents a greater correlation coefficient. * indicates significance at the P<0.05 level"

Table 6

Coefficient estimation and significance test results of the stepwise regression model"

变量Variable B β t P VIF
-0.111 0.000 -6.572 <0.001 -
X8 0.259 0.348 15.300 <0.001 1.380
X12 0.136 0.193 6.122 <0.001 2.641
X2 0.186 0.499 22.634 <0.001 1.298
X1 0.111 0.309 15.111 <0.001 1.118
X11 0.137 0.131 5.491 <0.001 1.526
X10 0.120 0.150 4.692 <0.001 2.726
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