不同水稻品种幼苗响应碱胁迫的生理差异及胁迫等级构建

doi:10.16258/j.cnki.1674-5906.2021.08.023

生态环境学报 ›› 2021, Vol. 30 ›› Issue (8): 1757-1768.DOI: 10.16258/j.cnki.1674-5906.2021.08.023

不同水稻品种幼苗响应碱胁迫的生理差异及胁迫等级构建

路旭平(), 李芳兰, 石亚飞, 张娟伟, 杨文伟, 罗成科^*(), 田蕾, 李培富

宁夏大学农学院/宁夏优势特色作物现代分子育种重点实验室,宁夏银川 750021

收稿日期:2021-05-25 出版日期:2021-08-18 发布日期:2021-11-03
通讯作者: * E-mail: chkluo2002@163.com
作者简介:路旭平（1995年生）,男,硕士研究生,主要从事水稻抗逆生理生态研究。E-mail: 577861974@qq.com
基金资助:
国家自然科学基金项目(32060425);宁夏自然科学基金项目(2020AAC03095)

Physiological Differences of Seedlings of Different Rice Varieties in Response to Alkali Stress and Construction of Stress Levels

LU Xuping(), LI Fanglan, SHI Yafei, ZHANG Juanwei, YANG Wenwei, LUO Chengke^*(), TIAN Lei, LI Peifu

Agricultural College of Ningxia University, Key Laboratory of Modern Molecular Breeding of Dominant Characteristic Crops in Ningxia, Yinchuan 75002, China

Received:2021-05-25 Online:2021-08-18 Published:2021-11-03

摘要/Abstract

摘要：

目前关于不同耐碱水稻品种响应碱胁迫的生理差异缺乏系统性的研究,依据生理生化指标计算胁迫指数并确定胁迫等级的方法较少见。为进一步阐明水稻耐碱的生理机制、提高水稻耐碱性和明确水稻抗碱等级,以碱敏感品种“中花11”和耐碱品种“宁粳52”为材料,设3个碱浓度水平（10、20和30 mmol∙L^-1）和3个pH水平（pH 8.65、pH 9.55和pH 10.50）,探究碱胁迫对不同耐碱水稻品种幼苗叶片相对含水量、叶绿素（Chla、Chlb、Chl）含量、叶绿素a/b（Chla/b）、丙二醛（MDA）含量、脂氧合酶（LOX）活性、活性氧（O₂^∙-和H₂O₂）含量、渗透调节物质（Pro、SS、SP）含量、抗氧化酶（SOD、POD和CAT）活性、还原型抗坏血酸（ASA）和还原型谷胱甘肽（GSH）含量的影响,通过主成分分析提取关键生理参数并建立碱胁迫等级。结果表明,（1）在碱胁迫条件下,耐碱品种“宁粳52”能维持更高的Chla、Chlb和Chl含量、Chla/b和叶片相对含水量,保持较低的MDA含量、LOX活性、O₂^∙-和H₂O₂含量,合成更多的SOD、POD和CAT活性、Pro、SS、SP、ASA和GSH含量。（2）通过主成分分析,挑选出Chla、SOD、MDA、H₂O₂、SS、ASA等关键指标,并计算胁迫指数Z,将水稻受害划分为正常（0≤Z<1）、轻度（1≤Z<4）、中度（4≤Z<7）、重度（7≤Z<10）、特重度（10≤Z）5个等级。（3）与20C（碱浓度为20 mmol∙L^-1、pH 10.50）相比,供试材料在30A（30 mmol∙L^-1、pH 8.65）处理下MDA含量、LOX活性、活性氧含量和胁迫指数均下降,说明在碱浓度大于20 mmol∙L^-1时,低碱浓度与高pH对水稻的伤害大于高碱浓度低pH。因此,在碱化土种植水稻时应考虑总盐碱浓度和pH共同对水稻生长的影响。

关键词: 水稻, 碱胁迫, pH, 幼苗, 生理, 胁迫指数

Abstract:

At present, there is a lack of systematic research on the physiological differences of tolerant and susceptible rice varieties in response to alkali stress. The method of calculating the stress index and determining the stress level based on physiological and biochemical indexes is also rarely seen. This study aims at elucidating the physiological mechanism of rice alkali resistance and providing a theoretical basis for improving alkali resistance and classifying alkali resistance levels in rice. The alkali-sensitive cultivar ‘Zhonghua 11’ and alkali-tolerant cultivar ‘Ninggeng 52’ were used as materials, and three alkali concentrations (10 mmol∙L^-1, 20 mmol∙L^-1, and 30 mmol∙L^-1) and three pH levels (pH 8.65, pH 9.55, and pH 10.50) were set to explore the effects of alkali stress on the leaves of rice seedlings. A series of indicators were investigated, including the relative water content, chlorophyll (Chla, Chlb, Chl) content, chlorophyll a/b (Chl a/b), malondialdehyde (MDA) content, lipoxygenase (LOX) activity, reactive oxygen species (O₂^{∙ -} and H₂O₂) content, osmotic adjustment substances (Pro, SS, SP) content, antioxidant enzymes (SOD, POD and CAT) activity, reduced ascorbic acid (ASA) and reduced glutathione (GSH) content. The key physiological parameters were extracted by principal component analysis and the alkali stress levels were established. The results showed that, (1) Under alkali stress, ‘Ninggeng 52’ could maintain higher levels of Chla, Chlb and Chl content, Chl a/b and leaf relative water content, SOD, POD and CAT activity, Pro, SS, SP, ASA and GSH content, and maintain lower levels of MDA content, LOX activity, O₂^{∙ -} and H₂O₂ content. (2) Through principal component analysis, the key indexes such as Chla, SOD, MDA, H₂O₂, SS and ASA were selected for calculation of the stress index Z. The damages in rice were classified into five grades: normal (0≤Z<1), mild (1≤Z<4), moderate (4≤Z<7), severe (7≤Z<10) and extremely severe (10≤Z). (3) The MDA content, LOX activity, active oxygen content and stress indexes of the tested materials decreased under the 30A (alkali concentration 30 mmol∙L^-1 and pH 8.65) treatment compared with that under 20C (alkali concentration 20 mmol∙L^-1 and pH 10.50), indicating that when the alkali concentration was greater than 20 mmol∙L^-1, the damage of low alkali concentration and high pH to rice was greater than that of high alkali concentration and low pH. Therefore, the effects of total saline-alkali concentration and pH on rice growth should be considered in the cultivation of rice in saline-alkali soils.

Key words: rice, alkali stress, pH, seedling, physiology, stress index

中图分类号:

Q945.78
X17

路旭平, 李芳兰, 石亚飞, 张娟伟, 杨文伟, 罗成科, 田蕾, 李培富. 不同水稻品种幼苗响应碱胁迫的生理差异及胁迫等级构建[J]. 生态环境学报, 2021, 30(8): 1757-1768.

LU Xuping, LI Fanglan, SHI Yafei, ZHANG Juanwei, YANG Wenwei, LUO Chengke, TIAN Lei, LI Peifu. Physiological Differences of Seedlings of Different Rice Varieties in Response to Alkali Stress and Construction of Stress Levels[J]. Ecology and Environment, 2021, 30(8): 1757-1768.

图/表 10

表1 各处理碱组成和pH

Table 1 Alkali composition and pH of each treatment

处理 Treatments	c_{(NaHCO3+Na2CO3)}/ (mmol∙L^-1)	pH	n_(NaHCO3)꞉ n_(Na2CO3)	c_(NaHCO3)/ (mmol∙L^-1)	c_(Na2CO3)/ (mmol∙L^-1)
CK	0	5.5±0.05	‒	0	0
10A	10	8.65±0.05	9꞉1	9	1
10B		9.55±0.05	5꞉5	5	5
10C		10.50±0.05	1꞉9	1	9
20A	20	8.65±0.05	9꞉1	18	2
20B		9.55±0.05	5꞉5	10	10
20C		10.50±0.05	1꞉9	2	18
30A	30	8.65±0.05	9꞉1	27	3
30B		9.55±0.05	5꞉5	15	15
30C		10.50±0.05	1꞉9	3	27

图1 碱胁迫对水稻叶片叶绿素含量的影响 n=3;10、20和30分别表示碱水平10、20和30 mmol∙L-1;A、B和C分别表示pH值8.65、9.55和10.50。不同小写字母表示同一品种在不同处理下差异显著（P<0.05）;不同大写字母表示不同品种在相同处理下差异显著（P<0.05）。下同

Fig. 1 Effects of alkali stress on chlorophyll contents of rice leaves n=3; 10, 20 and 30 respectively indicate alkali application levels of 10 mmol∙L-1, 20 mmol∙L-1 and 30 mmol∙L-1; A, B and C respectively indicate pH levels of 8.65, 9.55 and 10.50. Different small letters indicate significant difference of the same rice varieties in the different treatment at 0.05 level; Different capital letters indicate significant difference of different rice varieties in the same l treatment at 0.05 level. The same as below

图2 碱胁迫对水稻叶片相对含水量和渗透调节物质的影响

Fig. 2 Effects of alkali stress on relative water content and osmotic adjustment substances of rice leaves

图3 碱胁迫对水稻叶片脂质过氧化和活性氧含量的影响

Fig. 3 Effects of alkali stress on lipid peroxidation and reactive oxygen species of rice leaves

图4 碱胁迫对水稻叶片抗氧化酶活性的影响

Fig. 4 Effects of alkali stress on antioxidant enzyme activities of rice leaves

图5 碱胁迫对水稻叶片ASA和GSH含量的影响

Fig. 5 Effects of alkali stress on ASA and GSH contents of rice leaves

表2 两种水稻叶片生理参数的主成分分析

Table 2 Principal component analysis of physiological parameter in rice leaves

参数 Parameter	ZH11特征向量 Eigenvector	NG52特征向量 Eigenvector
参数 Parameter	第1主成分 PC1	第1主成分 PC1	第2主成分 PC2
Chl a	-0.996	-0.967	-0.109
Chl b	-0.973	-0.951	-0.244
Chl	-0.994	-0.965	-0.150
Chl a/b	-0.937	-0.099	0.968
RWC	-0.983	-0.848	0.415
SOD	0.834	0.966	0.120
POD	0.878	0.912	-0.092
CAT	0.979	0.948	0.200
MDA	0.957	0.980	-0.030
LOX	0.942	0.975	0.124
H₂O₂	0.983	0.991	-0.091
O₂^·ˉ	0.994	0.989	-0.093
Pro	0.984	0.995	0.065
SS	0.981	0.995	-0.037
SP	0.980	0.989	-0.097
ASA	0.986	0.990	-0.068
GSH	0.985	0.965	-0.031
特征根 Eigenvalue	15.787	14.901	1.321
贡献率 Contribution rate/%	92.867	87.653	7.768
累计贡献率 Cumulative contribution rate/%	92.867	87.653	95.422

表3 Chla、SOD、MDA、H2O2、SS和ASA的权重

Table 3 Weight coefficient of Chla, SOD, MDA, H2O2, SS and ASA

参数 Parameter	ZH11权重系数 ZH11 Weight coefficient/%	NG52权重系数 NG52 Weight coefficient/%
Chl a	16.07	26.02
SOD	9.13	11.11
MDA	13.87	14.24
H₂O₂	17.10	13.43
SS	23.26	15.43
ASA	20.58	19.77

图6 水稻受害等级的划分

Fig. 6 Classification of injury grades in Rice

表4 碱胁迫指数在不同处理下的变化规律

Table 4 Changes of alkali stress index with different treatments

处理 Treatments	ZH11		NG52
处理 Treatments	Z值 Z value	等级 Degree	Z值 Z value	等级 Degree
CK	0	正常 Normal	0	正常 Normal
10A	1.949	轻度 Mild	1.402	轻度 Mild
10B	3.029	轻度 Mild	2.429	轻度 Mild
10C	4.343	中度 Moderate	3.465	轻度 Mild
20A	4.534	中度 Moderate	4.315	中度 Moderate
20B	5.928	中度 Moderate	5.317	中度 Moderate
20C	8.069	重度 Severe	6.824	中度 Moderate
30A	6.733	中度 Moderate	5.936	中度 Moderate
30B	9.411	重度 Severe	8.125	重度 Severe
30C	11.276	特重度 Special severe	9.547	重度 Severe

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不同水稻品种幼苗响应碱胁迫的生理差异及胁迫等级构建

Physiological Differences of Seedlings of Different Rice Varieties in Response to Alkali Stress and Construction of Stress Levels

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