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作物学报 ›› 2022, Vol. 48 ›› Issue (2): 448-462.doi: 10.3724/SP.J.1006.2022.11003

• 耕作栽培·生理生化 • 上一篇    下一篇

基于主成分-聚类分析的不同水分冬小麦晚霜冻害评价

王洋洋1(), 贺利1, 任德超2, 段剑钊1, 胡新2, 刘万代1,*(), 郭天财1, 王永华1, 冯伟1,*()   

  1. 1河南农业大学农学院 / 省部共建小麦玉米作物学国家重点实验室,河南郑州 450046
    2商丘市农林科学研究院小麦研究所,河南商丘476000
  • 收稿日期:2021-01-07 接受日期:2021-04-26 出版日期:2022-02-12 网络出版日期:2021-06-09
  • 通讯作者: 刘万代,冯伟
  • 作者简介:E-mail: wyy65wyy@163.com
  • 基金资助:
    本研究由“十三五”国家重点研发计划项目“粮食丰产增效科技创新”(2017YFD0300204);国家现代农业产业技术体系建设专项资助(CARS-03)

Evaluations of winter wheat late frost damage under different water based on principal component-cluster analysis

WANG Yang-Yang1(), HE Li1, REN De-Chao2, DUAN Jian-Zhao1, HU Xin2, LIU Wan-Dai1,*(), GU Tian-Cai1, WANG Yong-Hua1, FENG Wei1,*()   

  1. 1Agronomy College of Henan Agriculture University / State Key Laboratory of Wheat and Maize Crop Science, Zhengzhou 450002, Henan, China
    2Wheat Research Institute, Shangqiu Academy of Agriculture and Forestry Sciences, Shangqiu 476000, Henan, China
  • Received:2021-01-07 Accepted:2021-04-26 Published:2022-02-12 Published online:2021-06-09
  • Contact: LIU Wan-Dai,FENG Wei
  • Supported by:
    This study was supported by the National Key Research and Development Program of China(2017YFD0300204);the China Agriculture Research System(CARS-03)

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摘要：

探讨不同水分条件下拔节期低温对小麦生理代谢的影响,构建综合生理指数精确评价冻害程度, 对农业减损增效和宏观管理具有重要意义。以弱春性的偃展4110和兰考198以及半冬性的郑麦366和丰德存21为材料,在低温处理前1周进行灌水和不灌水处理,于雌雄蕊分化期将盆栽小麦移至低温模拟室进行低温处理,分别为正常(CK)、-2℃ (T1)、-4℃ (T2)、-6℃ (T3)、-8℃ (T4)和-10℃ (T5)。在处理结束后第2天测定小麦生理指标和荧光参数,在成熟期收获小麦产量。结果表明,不同品种、水分和低温胁迫及其互作对小麦拔节期生理指标及荧光参数均具有显著影响。随着低温胁迫加重,叶片含水量、叶绿素a含量及荧光参数qp、Fv/Fm和Fv/Fo呈持续下降趋势,而可溶性蛋白、脯氨酸和可溶性糖含量及SOD活性表现先升后降特征,但MDA含量则相反。灌水处理在一定程度上缓解了低温胁迫对植株生理代谢的影响,低温对半冬性品种的影响相对较小。利用主成分分析将测定参数转化成4个独立的综合指标,且反映了88.55%的原始信息,并构建出冻害生理综合指数(FIPCI)。聚类分析热图显示的颜色越深,生理指标响应程度越大。依据生理指标聚类热图及FIPCI值将冻害划分为5个级别,且其与产量损失率具有较好的一致性。不同水分处理的产量损失率自T3温度开始差异较大,不灌水和灌水处理分别降低了30.4%~44%和21.0%~29.2%。同一水分条件下,产量损失率在品种间差异为LK198>YZ4110>ZM366>FDC21,半冬性品种低于弱春性品种。综合聚类热图和各处理产量,未受冻(CK)和轻度冻害(T1)的产量损失率低于10%,中度冻害(不灌水的T2、灌水的T2和T3)的产量损失率为10%~30%,重度冻害(不灌水的T3、灌水的T4)的产量损失率为30%~50%,特重度冻害(不灌水的T4和T5、灌水的T5)的产量损失率高于50%。可见,低温前灌水有利于缓解低温胁迫造成的伤害,在一定程度上降低了产量损失。利用主成分-聚类分析构建综合生理冻害指数及模型,可以准确评价小麦晚霜冻害的程度,为灾后产量恢复及决策管理提供科学依据。

关键词: 冬小麦, 晚霜冻害, 生理指标, 荧光参数, 主成分分析, 综合生理指数

Abstract:

To investigate the effect of low temperature at jointing stage on wheat physiological metabolism under different water conditions, a comprehensive physiological index is constructed to accurately evaluate the degree of freezing injury, which is of great significance to agricultural loss reduction, efficiency increase and macro management. Irrigation (W1) and no irrigation (W0) treatments were carried out one week before low temperature treatment using the weak-spring Yanzhan 4110 and Lankao 198, semi-winter Zhengmai 366 and Fengdecun 21 as the experimental materials. At the female and male ear differentiation stages, potted wheat was moved to the low temperature simulation room for low temperature treatment with six levels of normal (CK), -2℃ (T1), -4℃ (T2), -6℃ (T3), -8℃ (T4), and -10℃ (T5), respectively. The physiological indexes and fluorescence parameters were measured on the second day after treatment, and wheat yield was harvested at maturity stage. The results revealed that different varieties, water and low temperature stress, and their interactions had significant effects on wheat physiological indexes and fluorescence parameters at jointing stage. With the aggravation of low temperature stress, leaf water content, chlorophyll a content and fluorescence parameters qp, Fv/Fm, and Fv/Fo showed the continuous downward trends, the contents of soluble protein, proline, soluble sugar, and SOD activity first increased and then decreased, but the content of MDA demonstrated the opposite trend. Irrigation treatment alleviated the influence of low temperature stress on plant physiological metabolism, and the effect of low temperature on semi-winter varieties was relatively lower. Four independent comprehensive indexes were transformed by the principal component analysis, which reflected 88.55% of the original information, and the physiological comprehensive index of freezing injury (FICEI) was constructed. The depth of color in heat map indicated the darker the color, the greater the performance degree of indicators. According to the FIPCI value, the freezing injury was divided into five levels, which was consistent with the yield loss rate. Especially under T3 treatment, the yield loss rate of each variety reduced by 30.4%-44.0% under no irrigation, reduced by 21.0%-29.2% under irrigation treatment. Under the same irrigation condition, yield loss rate of different varieties was LK198>YZ4110>ZM366>FDC21, and the yield loss rate of semi-winter varieties was lower than that of weak spring varieties. According to the results of heat map clustering and the yield of each treatment, the yield loss rate was less than 10% for CK and T1, 10%-30% for W0T2, W1T2 and W1T3, 30%-50% for W0T3 and W1T4, more than 50% for W0T4, W0T5 and W1T5. Irrigating before low temperature was conducive to alleviating the damage caused by low temperature stress and reducing the yield loss. Physiological comprehensive freezing injury index and model constructed by principal component-cluster analysis can accurately evaluate the degree of wheat late frost damage, and provide scientific basis for yield recovery and decision-making management after disaster.

Key words: winter wheat, late frost damage, physiological indicators, fluorescence parameters, principal component analysis, integrated physiological index

表1

冻害生理指标及叶绿素荧光参数方差分析"

处理
Treatment		 含水量
Leaf water content (%)		 叶绿素a
Chl a
(mg g-1 )		 脯氨酸
Proline
(μg g-1)		 丙二醛MDA
(μg g-1)		 光化学
淬灭系数
qp		 PS II最大光学效率
Fv/Fm		 PS II潜在活性
Fv/Fo		 可溶性
蛋白
Soluble protein
(mg g-1)		 可溶性糖
Soluble
sugar
(mg g-1)		 超氧化物歧化酶
SOD
(U g-1)		 F-
value
(0.05)		 F-
value
(0.01)
2018-2019
灌水Irr 36.8** 4.4* 4.0* 2.2 2.0 0.8 3.0 8.2** 1.3 76.0** 3.9 6.9
品种Cul 2.6* 1.9 2.0 1.3 1.8 2.4 2.4 24.8** 6.2* 10.5** 2.5 3.5
温度Tem 21.4** 46.3** 72.8** 44.8** 166.0** 69.1** 257.1** 19.8** 57.9** 40.2** 2.3 3.2
Irr × Cul 7.2** 2.9* 1.9 1.7 1.0 1.7 1.5 16.6** 3.1* 29.6** 2.5 3.5
Irr × Tem 45.1** 25.6** 48.2** 23.4** 85.6** 35.6** 119.7** 19.4** 29.9** 23.8** 2.3 3.2
Cul × Tem 6.8** 23.8** 45.8** 14.1** 123.2** 47.6** 149.8** 20.2** 17.1** 5.1** 1.7 2.0
Irr × Cul × Tem 113.3** 407.6** 319.9** 403.4** 796.0** 786.1** 966.6** 238.2** 223.0** 448.5** 1.7 2.0
2019-2020
灌水Irr 6.2* 17.8** 6.9** 3.2 8.5** 2.3 0.1 2.2 23.1** 15.3** 3.9 6.9
品种Cul 6.1* 0.9 1.0 0.4 1.6 0.3 0.7 3.1* 19.5** 11.3** 2.5 3.5
温度Tem 118.8** 106.7** 158.8** 235.6** 188.8** 435.1** 550.3** 11.4** 39.5** 35.1** 2.3 3.2
Irr × Cul 5.4* 3.2* 1.5 0.7 2.0 0.5 0.4 11.5* 5.3* 8.1* 2.5 3.5
Irr × Tem 106.8** 126.2** 168.6** 232.5** 196.4** 469.6** 248.9** 16.61** 56.7** 36.9** 2.3 3.2
Cul × Tem 42.3** 30.8** 49.9** 63.6** 54.7** 103.1** 189.6** 19.0** 12.9** 19.0** 1.7 2.0
Irr × Cul × Tem 333.2** 408.8** 519.1** 535.0** 209.5** 306.8** 402.0** 206.4** 54.5** 287.5** 1.7 2.0

图1

低温胁迫对冬小麦叶片含水量及叶绿素a含量的影响 W0: 不灌水处理; W1: 灌水处理。"

图2

低温胁迫对冬小麦渗透调节物质含量的影响 W0: 不灌水处理; W1: 灌水处理。"

图3

低温胁迫对冬小麦叶片SOD活性及MDA含量的影响 W0: 不灌水处理; W1: 灌水处理。"

图4

低温胁迫对冬小麦叶片qp、Fv/Fm和Fv/Fo的影响 W0: 不灌水处理; W1: 灌水处理。"

表2

冬小麦生理指标及荧光参数间相关系数"

X1 X2 X3 X4 X5 X6 X7 X8 X9 X10
X1 1
X2 0.762** 1
X3 0.115 0.153* 1
X4 0.687** 0.707** 0.395** 1
X5 0.761** 0.787** 0.201* 0.753** 1
X6 0.771** 0.839** 0.194* 0.736** 0.921** 1
X7 0.785** 0.780** 0.122 0.687** 0.914** 0.896** 1
X8 -0.782** -0.792** -0.259** -0.730** -0.798** -0.801** -0.808** 1
X9 0.271** 0.223** 0.402** 0.486** 0.373** 0.344** 0.332** -0.367** 1
X10 0.464** 0.462** 0.358** 0.584** 0.507** 0.565** 0.519** -0.583** 0.399** 1

表3

冬小麦生理指标及荧光参数的主成分分析"

成分
Component		 初始特征值 Initial eigenvalues 提取载荷平方和 Extraction eigenvalues
总计
Total		 方差百分比
Variance (%)		 累积
Cumulative (%)		 总计
Total		 方差百分比
Variance (%)		 累积
Cumulative (%)
1 6.366 63.658 63.658 6.366 63.658 63.658
2 1.357 13.574 77.232 1.357 13.574 77.232
3 0.603 6.028 83.259 0.603 6.028 83.259
4 0.529 5.290 88.550 0.529 5.290 88.550
5 0.325 3.253 91.803
6 0.265 2.655 94.458
7 0.224 2.241 96.699
8 0.182 1.822 98.521
9 0.087 0.868 99.389
10 0.061 0.611 100

图5

不同水分及低温处理条件下植株生理指标之间的关系 X1、X2、X3、X4、X5、X6、X7、X8、X9和X10分别代表叶绿素a、叶含水量、可溶性糖、脯氨酸、qp、Fv/Fm、Fv/Fo、MDA、SOD和可溶性蛋白。"

表4

10个参数相互矩阵的特征向量分析表"

指标 Index CI1 CI2 CI3 CI4
叶绿素a Chlorophyll a 0.852 -0.231 -0.004 0.052
叶片含水量 Leaf water content 0.868 -0.239 -0.123 0.108
可溶性糖 Soluble sugar 0.319 0.786 -0.414 0.296
脯氨酸 Proline 0.859 0.187 -0.029 0.076
光化学淬灭系数 qp 0.926 -0.141 0.068 0.100
PS II最大光学效率 Fv/Fm 0.933 -0.156 0.006 0.020
PS II潜在活性 Fv/Fo 0.911 -0.216 0.082 0.014
丙二醛 MDA -0.903 0.056 0.057 0.010
超氧化物歧化酶 SOD 0.472 0.625 0.610 0.035
可溶性蛋白 Soluble protein 0.670 0.330 -0.169 -0.640

表5

各主成分参数变量的权重、综合公式Y权重、FIPCI权重"

指标
Index		 CI1 CI2 CI3 CI4 Y权重
Y weight		 FIPCI权重FIPCI weight
叶绿素a Chlorophyll a 0.337681 -0.198300 -0.005150 0.071495 0.216279 0.118316
叶片含水量Leaf water content 0.344022 -0.205170 -0.158400 0.148490 0.213953 0.117044
可溶性糖Soluble sugar 0.126432 0.674734 -0.533140 0.406971 0.182342 0.099751
脯氨酸Proline 0.340455 0.160528 -0.037350 0.104493 0.273059 0.149378
光化学淬灭系数qp 0.367010 -0.121040 0.087569 0.137490 0.259461 0.141939
PS II最大光学效率Fv/Fm 0.369784 -0.133920 0.007727 0.027498 0.247476 0.135383
PS II潜在活性Fv/Fo 0.361065 -0.185420 0.105598 0.019249 0.239482 0.131009
丙二醛MDA -0.357890 0.048073 0.073403 0.013749 -0.244100 -0.133540
超氧化物歧化酶SOD 0.187072 0.536525 0.785545 0.048122 0.273080 0.149390
可溶性蛋白Soluble protein 0.265547 0.283285 -0.217630 -0.879940 0.166942 0.091326

图6

生理生化特性的热图分析 1、2、3和4分别为品种YZ4110、LK198、ZM366和FDC21; X1、X2、X3、X4、X5、X6、X7、X8、X9和X10分别为叶绿素a、叶含水量、可溶性糖、脯氨酸、qp、Fv/Fm、Fv/Fo、MDA、SOD和可溶性蛋白。处理同图1。"

表6

不同品种、水分及低温胁迫条件下FIPCI值、产量及产量损失率"

处理
Treatment		 冻害生理综合指数
FIPCI		 平方产量
Yield (g m-2)		 产量损失率
Rate of loss of yield (%)
YZ4110 LK198 ZM366 FDC21 YZ4110 LK198 ZM366 FDC21 YZ4110 LK198 ZM366 FDC21
W0-CK 0.415 0.394 0.421 0.458 938.0 902.9 941.6 854.4 0 a 0 a 0 a 0 a
W0-T1 0.607 0.622 0.65 0.672 840.0 834.6 895.1 819.2 7.1 b 7.5 b 4.6 b 4.1 b
W0-T2 0.898 0.916 0.994 0.993 698.6 636.3 763.4 705.6 25.6 c 29.6 c 19.0 c 17.3 c
W0-T3 -0.181 -0.251 -0.137 -0.143 526.4 485.6 609.1 593.6 44.0 d 46.2 d 35.3 d 30.4 d
W0-T4 -1.294 -1.334 -1.109 -1.016 352.8 235.7 331.2 272.0 62.4 e 73.9 e 64.8 e 68.1 e
W0-T5 -1.341 -1.449 -1.387 -1.257 147.0 50.2 97.1 124.8 84.3 f 94.4 f 89.7 f 85.4 f
W1-CK 0.384 0.407 0.438 0.418 971.6 966.0 996.8 979.2 0 a 0 a 0 a 0 a
W1-T1 0.717 0.674 0.777 0.818 950.6 936.6 980.0 961.2 2.2 a 3.1 a 1.7 a 1.8 a
W1-T2 1.093 1.090 1.192 1.299 793.8 810.6 870.8 855.6 18.4 b 16.1 b 12.7 b 12.6 b
W1-T3 1.129 1.192 1.299 1.351 718.2 684.6 788.2 756.0 26.1 c 29.2 c 21.0 c 22.8 c
W1-T4 -0.354 -0.416 -0.328 -0.225 520.8 487.2 575.4 603.6 46.4 d 49.5 d 42.3 d 38.4 d
W1-T5 -1.267 -1.470 -1.199 -1.054 233.8 166.6 250.6 285.6 75.9 f 82.7 f 74.8 f 70.8 f

图7

低温胁迫下小麦植株生理指标之间的关系 ROS、SOD、MDA、PRO、SP、SS、LWC、Chl a、Fv/Fm、Fv/Fo和qp分别代表活性氧、SOD酶、MDA、脯氨酸、可溶性蛋白、可溶性糖、叶片含水量、叶绿素a、Fv/Fm、Fv/Fo和qp。"

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