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作物学报

作物学报 ›› 2024, Vol. 50 ›› Issue (1): 172-186.doi: 10.3724/SP.J.1006.2024.33019

所属专题： 玉米：耕作栽培·生理生化

• 耕作栽培·生理生化 • 上一篇    下一篇

长江中下游地区糯玉米花期耐热性鉴定及评价指标筛选

宋旭东1(), 朱广龙3(), 张舒钰1, 章慧敏1, 周广飞1, 张振良1, 冒宇翔1, 陆虎华1, 陈国清1,2, 石明亮1, 薛林1,2, 周桂生3, 郝德荣1,*()   

  1. 1江苏沿江地区农业科学研究所, 江苏南通226541
    2江苏省现代作物生产协同创新中心, 江苏南京210095
    3扬州大学教育部农业与农产品安全国际合作联合实验室 / 江苏省粮食作物现代产业技术协同创新中心, 江苏扬州 225009
  • 收稿日期:2023-03-27 接受日期:2023-06-29 出版日期:2024-01-12 网络出版日期:2023-07-27
  • 通讯作者: *郝德荣, E-mail: drhao2008@163.com
  • 作者简介:宋旭东, E-mail: xudong.song@foxmail.com; 朱广龙, E-mail: zhuguang2007@163.com

    **同等贡献

  • 基金资助:
    江苏省种业振兴揭榜挂帅项目(JBGS[2021]054);江苏省重点研发项目(BE2021317);南通市基础科学研究项目(JC12022088);江苏省自主创新项目(CX(20)1002);江苏现代农业产业技术体系建设专项资助

Identification of heat tolerance of waxy maizes at flowering stage and screening of evaluation indexes in the middle and lower reaches of Yangtze River region

SONG Xu-Dong1(), ZHU Guang-Long3(), ZHANG Shu-Yu1, ZHANG Hui-Min1, ZHOU Guang-Fei1, ZHANG Zhen-Liang1, MAO Yu-Xiang1, LU Hu-Hua1, CHEN Guo-Qing1,2, SHI Ming-Liang1, XUE Lin1,2, ZHOU Gui-Sheng3, HAO De-Rong1,*()   

  1. 1Yanjiang Institute of Agricultural Sciences, Nantong 226541, Jiangsu, China
    2Jiangsu Collaborative Innovation Center for Modern Crop Production, Nanjing 210095, Jiangsu, China
    3Joint International Research Laboratory of Agriculture and Agri-Product Safety, the Ministry of Education, Yangzhou University / Co-Innovation Center for Modern Production Technology in Grain Crops of Jiangsu Province, Yangzhou 225009, Jiangsu, China
  • Received:2023-03-27 Accepted:2023-06-29 Published:2024-01-12 Published online:2023-07-27
  • Contact: *E-mail: drhao2008@163.com
  • About author:**Contributed equally to this study
  • Supported by:
    Open Competition Project of Seed Industry Revitalization of Jiangsu Province(JBGS[2021]054);Jiangsu Province Key Research and Development Project(BE2021317);Scientific and Technological Project of Nantong City, China(JC12022088);Jiangsu Agriculture Science and Technology Innovation Fund(CX(20)1002);Earmarked Fund for Jiangsu Agricultural Research System

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摘要：

全球气候变暖趋势日趋明显, 高温热害已成为限制糯玉米丰产稳产的主要非生物胁迫之一。鉴定糯玉米的耐热性, 筛选耐高温评价指标, 可为耐热糯玉米品种的选育提供理论支撑。本研究以长江中下游地区主推品种为试验材料, 设置2种增温方式(覆膜增温处理和延迟播种处理)以及自然对照处理(CK), 测定了13个与耐热相关的指标, 采用主成分分析、隶属函数法、聚类分析和逐步回归分析相结合的方法, 对糯玉米花期耐热性进行综合评价。结果显示, 与对照相比, 除叶绿素a和光系统II最大光化学效率外, 各指标在高温处理下的降低幅度均达到显著或极显著水平。利用主成分分析将13个单项指标转换为3个相互独立的综合指标, 其贡献率分别为64.46%、15.06%和7.76%, 代表了糯玉米耐热性87.28%的原始数据信息量; 通过隶属函数计算综合耐热评价值(D值), 将10个参试糯玉米品种划分为3类, 分别为3个耐热型、4个中等耐热型和3个热敏感型。筛选出的耐热型品种为苏玉糯2号、中糯2号和苏玉糯901, 生产中应用这些品种可能在一定程度上减轻高温热害对糯玉米生长发育造成的不利影响。最后, 利用逐步回归分析建立了糯玉米耐热性评价方程预测供试材料的耐热性, 预测值(PV)与D值基本一致, 确定了产量、Fv/Fm、叶面积指数和花粉活力作为糯玉米花期耐热性评价指标。本研究筛选出的耐热品种和鉴定指标可为今后耐热糯玉米品种的选育提供理论依据。

关键词: 糯玉米, 花期耐热性, 综合评价, 鉴定指标

Abstract:

Heat stress has acknowledged as one of the major threats to waxy maize production as a result of global warming gradually seriously. Heat resistance identification and indices screening could provide the theoretical basis in breeding heat-resistant cultivars. In the present study, 13 heat-related parameters of 10 waxy maize cultivars from the middle and lower reaches of Yangtze River area were measured under normal temperature treatment (CK) and two heat stress treatments (artificial warming treatment and delayed sowing treatment). Principal component analysis, membership function method, cluster analysis, and stepwise regression analysis were used to comprehensively evaluate the heat resistance of waxy maize at flowering stage. The results showed that most indices under heat stress decreased significantly compared with CK, except for chlorophyll a and PS II primary light energy conversion efficiency. Three independent comprehensive components were obtained from 13 single traits using a principal component analysis, and their contribution rates respectively were 64.46%, 15.06%, and 7.76%, which represented 87.28% information of the original data. Ten testing cultivars were divided into three heat-tolerance types that were heat tolerance category, medium heat tolerance category, and high temperature sensitive category based on comprehensive heat tolerance values (D) calculating by membership function methods. SYN2, ZN2, and SYN901 were identified as the heat-resistant cultivars, and planting these cultivars could partly alleviate the negative effects of heat stress on waxy maize production. Finally, stepwise regression method was used to establish a predictive equation to evaluate heat tolerance, which indicating that predictive values (PV) calculated by the predictive equation were basically consistent with D value. The results showed that yield, Fv/Fm, leaf area index, and pollen viability could be used to identify heat-resistant cultivars of waxy maize. The heat tolerant cultivars and identification index selected in the study could serve as a basis for subsequent breeding heat tolerant cultivars of waxy maize.

Key words: waxy maize, heat-tolerance at flowering stage, comprehensive assessment, evaluation indexes

图1

不同处理下各糯玉米品种播种时间 A: 对照和覆膜增温处理; B: 延迟播种处理。SYN802: 苏玉糯802; ZN2: 中糯2号; SYN2: 苏玉糯 2号; SYN11: 苏玉糯11; SYN901: 苏玉糯901; SYN639: 苏玉糯639; JDN517: 焦点糯517; SYN1803: 苏玉糯1803; SYN1502: 苏玉糯1502; SYN1704: 苏玉糯1704。"

图2

高温处理期增温棚内(H1)、延迟播种田间高温(H2)与对照处理(CK)日最高温度, 其中虚线代表不同处理的平均温度"

图3

花期不同处理对糯玉米主要农艺性状的影响 LAI: 叶面积指数(cm2 m-2); DB: 地上部干物重(g m-2); ERN: 穗行数; KNR: 行粒数; SR: 结实率(%); Fv/Fm: PSII最大光化学效率; Fv/Fo: PSII潜在活性; Chl a: 叶绿素a (mg g-1); Chl b: 叶绿素b (mg g-1); Chl (a+b): 叶绿素a+b (mg g-1); PV: 花粉活力(%); Yield: 产量(kg hm-2); ETB: 秃尖长度(cm)。*表示P < 0.05的显著水平; **表示P < 0.01的显著水平; ns表示差异不显著。"

附表1

不同处理下10个糯玉米品种13个农艺性状的表现"

品种
Cultivar		 LAI DB ERN KNR SR
CK H1 H2 CK H1 H2 CK H1 H2 CK H1 H2 CK H1 H2
苏玉糯639 SYN639 4.47 3.80 4.04 932.4 843.7 860.5 15.1 12.9 13.3 29.67 23.61 25.08 89.65 72.51 74.28
苏玉糯2号 SYN2 3.56 3.22 3.35 892.5 827.1 860.8 14.0 12.8 12.9 27.33 25.01 25.31 90.48 82.15 84.50
苏玉糯11 SYN11 5.17 4.23 4.61 995.9 885.2 917.7 13.9 11.7 11.9 32.67 26.84 27.24 86.91 68.88 69.90
中糯2号ZN2 4.28 3.91 4.03 878.9 789.7 853.2 13.3 12.5 12.4 27.81 25.74 26.24 92.33 84.37 86.20
苏玉糯1803 SYN1803 4.40 3.89 3.99 949.0 831.4 895.2 14.0 12.8 12.7 34.48 29.26 30.48 90.20 73.20 74.80
焦点糯517 JDN517 5.38 4.43 4.97 1090.4 974.1 1045.5 15.6 13.2 13.2 35.67 28.53 29.87 87.08 69.61 70.99
苏玉糯1502 SYN1502 4.62 4.19 4.21 1154.5 998.1 1035.3 15.6 13.3 13.7 29.33 25.39 26.27 88.08 78.59 79.98
苏玉糯1704 SYN1704 4.54 4.13 4.30 903.5 794.9 821.3 13.8 11.5 11.7 33.33 29.33 30.45 85.60 73.11 76.20
苏玉糯901 SYN901 4.37 4.01 4.11 1096.7 1021.2 1032.7 15.0 12.9 13.1 30.67 28.2 28.74 89.35 79.38 81.50
苏玉糯802 SYN802 5.69 5.09 5.37 1143.7 1016.7 1072.9 14.6 13.2 13.2 30.63 26.67 26.84 87.70 75.25 76.10
品种
Cultivar		 Fv/Fm Fv/Fo Chl a Chl b Chl a+b
CK H1 H2 CK H1 H2 CK H1 H2 CK H1 H2 CK H1 H2
苏玉糯639 SYN639 0.75 0.68 0.68 2.94 2.60 2.64 0.76 0.69 0.71 1.33 1.14 1.13 2.11 1.79 1.81
苏玉糯2号 SYN2 0.76 0.71 0.73 3.20 2.82 2.97 0.76 0.70 0.70 1.35 1.24 1.25 2.18 2.03 2.01
苏玉糯11 SYN11 0.74 0.67 0.70 3.05 2.56 2.8 0.75 0.65 0.66 1.26 1.07 1.09 2.02 1.77 1.74
中糯2号ZN2 0.76 0.70 0.73 3.21 2.86 2.93 0.73 0.69 0.67 1.28 1.15 1.19 2.04 1.81 1.95
苏玉糯1803 SYN1803 0.76 0.70 0.71 3.12 2.70 2.67 0.76 0.70 0.71 1.45 1.32 1.24 2.17 1.89 1.91
焦点糯517 JDN517 0.78 0.71 0.73 3.50 2.98 3.00 0.77 0.70 0.70 1.37 1.16 1.17 2.15 1.86 1.9
苏玉糯1502 SYN1502 0.75 0.69 0.70 3.19 2.73 2.70 0.76 0.71 0.72 1.21 1.10 1.06 2.05 1.88 1.91
苏玉糯1704 SYN1704 0.78 0.71 0.73 3.48 2.99 3.15 0.78 0.72 0.73 1.12 0.95 1.00 1.99 1.79 1.82
苏玉糯901 SYN901 0.76 0.71 0.72 3.25 2.72 2.97 0.72 0.66 0.68 1.43 1.29 1.30 2.14 1.96 2.03
苏玉糯802 SYN802 0.77 0.68 0.69 3.31 2.82 2.94 0.7 0.63 0.64 1.48 1.31 1.31 2.2 1.94 1.99
品种
Cultivar		 PV ETB Yield
CK H1 H2 CK H1 H2 CK H1 H2
苏玉糯639 SYN639 CK H1 H2 CK H1 H2 10030.5 8158.5 8462.9
苏玉糯2号 SYN2 90.3 71.5 74.1 1.54 2.30 2.40 9518.0 8649.5 8779.5
苏玉糯11 SYN11 88.8 76.3 80.5 1.10 1.29 1.21 10627.5 8382.0 8801.5
中糯2号ZN2 87.2 64.7 71.5 1.67 2.90 2.68 9432.0 8527.5 8911.5
苏玉糯1803 SYN1803 91.2 80.1 82.4 0.73 0.81 0.78 10519.5 8896.5 9021.0
焦点糯517 JDN517 90.3 75.5 77.2 1.10 1.53 1.49 10631.0 8469.0 8736.0
苏玉糯1502 SYN1502 91.1 71.9 74.5 1.43 2.72 2.23 9679.5 8163.5 8371.5
苏玉糯1704 SYN1704 89.5 71.6 77.2 1.50 2.45 2.32 10773.0 9064.5 9360.5
苏玉糯901 SYN901 87.1 70.8 73.3 1.33 2.07 2.00 10242.0 9001.0 9191.0
苏玉糯802 SYN802 91.1 76.9 79.7 1.27 2.09 2.14 10123.5 8542.0 8763.0

表1

不同糯玉米品种各单项指标的耐热系数"

品种 Cultivar X1 X2 X3 X4 X5 X6 X7 X8 X9 X10 X11 X12 X13
苏玉糯639 SYN639 0.877 0.914 0.870 0.821 0.819 0.913 0.892 0.908 0.852 0.855 0.806 0.829 1.500
苏玉糯2号 SYN2 0.922 0.946 0.924 0.921 0.921 0.943 0.905 0.928 0.919 0.918 0.880 0.917 1.138
苏玉糯11 SYN11 0.856 0.906 0.855 0.828 0.799 0.927 0.879 0.882 0.868 0.869 0.781 0.808 1.676
中糯2号 ZN2 0.928 0.940 0.931 0.935 0.924 0.936 0.903 0.927 0.915 0.923 0.891 0.925 1.089
苏玉糯1803 SYN1803 0.906 0.920 0.910 0.870 0.821 0.929 0.861 0.926 0.884 0.872 0.846 0.842 1.379
焦点糯517 JDN517 0.874 0.930 0.847 0.819 0.808 0.925 0.853 0.905 0.855 0.877 0.804 0.809 1.728
苏玉糯1502 SYN1502 0.909 0.886 0.867 0.881 0.906 0.925 0.851 0.938 0.895 0.922 0.831 0.850 1.593
苏玉糯1704 SYN1704 0.928 0.895 0.842 0.900 0.873 0.921 0.880 0.923 0.869 0.909 0.827 0.854 1.529
苏玉糯901 SYN901 0.930 0.937 0.870 0.929 0.904 0.941 0.878 0.936 0.903 0.932 0.859 0.881 1.676
苏玉糯802 SYN802 0.918 0.916 0.905 0.887 0.864 0.893 0.871 0.905 0.881 0.884 0.839 0.855 1.627
平均值 Mean 0.905 0.919 0.882 0.879 0.864 0.925 0.877 0.918 0.884 0.896 0.836 0.857 1.494
标准差 SD 0.027 0.020 0.033 0.044 0.049 0.014 0.019 0.017 0.024 0.027 0.034 0.040 0.225
变异系数 CV (%) 2.935 2.125 3.698 5.028 5.680 1.563 2.168 1.900 2.673 3.062 4.112 4.700 15.038

表2

高温处理下各指标耐热系数相关分析"

X1 X2 X3 X4 X5 X6 X7 X8 X9 X10 X11 X12 X13
X1 1.000
X2 0.142 1.000
X3 0.435 0.498 1.000
X4 0.896** 0.065 0.304 1.000
X5 0.862** 0.199 0.579* 0.903** 1.000
X6 0.010 0.448 0.307 0.282 0.275 1.000
X7 0.344 0.631* 0.603* 0.205 0.313 0.341 1.000
X8 0.678* 0.296 0.256 0.859** 0.737* 0.614* 0.484 1.000
X9 0.743* 0.459 0.317 0.841** 0.795** 0.460 0.556 0.942** 1.000
X10 0.717* 0.270 0.373 0.889** 0.834** 0.560* 0.360 0.912** 0.881** 1.000
X11 0.867** 0.344 0.536 0.902** 0.869** 0.485 0.559 0.912** 0.896** 0.884** 1.000
X12 0.861** 0.238 0.530 0.884** 0.914** 0.316 0.581 0.861** 0.895** 0.873** 0.936** 1.000
X13 -0.412 0.002 -0.372 -0.424 -0.501 -0.220 -0.659* -0.568 -0.543 -0.383 -0.573* -0.700* 1.000

表3

各性状主成分的特征向量及贡献率"

主成分 Principle factor CI1 CI2 CI3
特征值 Eigen value 8.38 1.96 1.01
贡献率 Contribution ratio 64.46 15.06 7.76
累计贡献率 Cumulative contribution ratio (%) 64.46 79.52 87.28
特征向量 Eigen vector X1 0.84 0.36 -0.03
X2 0.53 -0.59 0.28
X3 0.71 -0.49 -0.34
X4 0.94 0.21 -0.09
X5 0.91 0.29 -0.07
X6 0.55 -0.05 0.81
X7 0.55 -0.61 0.00
X8 0.75 0.49 0.06
X9 0.94 0.03 0.07
X10 0.82 0.49 0.19
X11 0.97 -0.07 -0.11
X12 0.98 -0.12 -0.06
X13 -0.75 0.49 0.09

表4

各参试材料综合指标值、权重、u(Xj)、D值及综合评价"

品种
Cultivar		 CI(1) CI(2) CI(3) u(X1) u(X2) u(X3) D值
D-value		 排序
Rank		 综合评价
Comprehensive valuation
苏玉糯639 SYN639 2.42 0.46 0.59 0.22 0.61 0.41 0.303 8 高温敏感型 Heat sensitive type
苏玉糯2号 SYN2 2.71 0.36 0.55 0.91 1.00 1.00 0.932 2 耐热型 Heat tolerance type
苏玉糯11 SYN11 2.34 0.53 0.63 0.13 0.43 0.67 0.228 10 高温敏感型 Heat sensitive type
中糯2号 ZN2 2.74 0.35 0.53 0.95 0.98 0.86 0.950 1 耐热型 Heat tolerance type
苏玉糯1803 SYN1803 2.52 0.44 0.57 0.52 0.38 0.72 0.512 6 中等耐热型Medium heat tolerance type
焦点糯517 JDN517 2.34 0.56 0.64 0.20 0.39 0.63 0.269 9 高温敏感型 Heat sensitive type
苏玉糯1502 SYN1502 2.49 0.59 0.59 0.63 0.00 0.64 0.523 5 中等耐热型Medium heat tolerance type
苏玉糯1704 SYN1704 2.50 0.55 0.59 0.57 0.35 0.57 0.533 4 中等耐热型Medium heat tolerance type
苏玉糯901 SYN901 2.54 0.60 0.61 0.79 0.67 0.95 0.787 3 耐热型 Heat tolerance type
苏玉糯802 SYN802 2.47 0.54 0.55 0.43 0.44 0.00 0.393 7 中等耐热型Medium heat tolerance type
权重Weight 0.739 0.173 0.089

图4

基于D值的鲜食糯玉米品种耐热性系统聚类分析图 缩写同图1。"

表5

回归方程的精度分析"

品种
Cultivar		 预测值
Predicted value		 原始值
Primary value		 差值
Difference		 估计精度
Evaluation accuracy (%)
苏玉糯639 SYN639 0.299 0.303 -0.004 98.68
苏玉糯2号 SYN2 0.935 0.932 0.003 99.68
苏玉糯11 SYN11 0.228 0.228 0.000 100.00
中糯2号 ZN2 0.951 0.950 0.001 99.89
苏玉糯1803 SYN1803 0.516 0.512 0.004 99.22
焦点糯517 JDN517 0.272 0.269 0.003 98.88
苏玉糯1502 SYN1502 0.512 0.523 -0.011 97.90
苏玉糯1704 SYN1704 0.542 0.533 0.000 100.00
苏玉糯901 SYN901 0.778 0.787 0.009 98.31
苏玉糯802 SYN802 0.392 0.393 -0.009 98.86

图5

聚类结果中不同耐热型糯玉米品种各性状的表现特征 *和**分别表示在0.05和0.01概率水平差异显著, ns表示在0.05概率水平无显著差异。"

表6

高温处理对不同耐热型糯玉米品种产量的影响"

耐热类型
Tolerance type		 品种
Cultivar		 对照产量
Control yield
(kg hm-2)		 高温产量
High temperature yield (kg hm-2)		 产量下降比例
Relative yield reduction (%)
H1 H2 平均值Mean H1 H2 平均值Mean
耐热型
Heat resistance		 中糯2号 ZN2 628.8 568.5 594.1 581.3 9.59 5.52 7.55
苏玉糯2号 SYN2 634.5 576.6 585.3 581.0 9.12 7.76 8.44
苏玉糯901 SYN901 682.8 600.1 612.7 606.4 12.12 10.26 11.19
高温敏感型
Heat sensitive		 苏玉糯639 SYN639 668.7 543.9 564.2 554.0 18.66 15.63 17.15
焦点糯517 JDN517 708.7 564.6 582.4 573.5 20.34 17.83 19.08
苏玉糯11 SYN11 708.5 558.8 586.8 572.8 21.13 17.18 19.16
中等耐热型
Medium resistance		 苏玉糯1803 SYN1803 701.3 593.1 601.4 597.3 15.43 14.24 14.84
苏玉糯1502 SYN1502 645.3 544.2 558.1 551.2 15.66 13.51 14.59
苏玉糯1704 SYN1704 718.2 604.3 624.0 614.2 15.86 13.11 14.49
苏玉糯802 SYN802 674.9 569.5 584.2 576.8 15.62 13.44 14.53
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