李芳兰,
马晓娟,
景盼盼,
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 银川 750021
基金项目: 国家自然科学基金项目32060425
国家自然科学基金项目31560297
宁夏自然科学基金项目2020AAC03095
详细信息
作者简介:路旭平, 主要从事水稻抗逆分子生物学研究。E-mail: 577861974@qq.com
通讯作者:罗成科, 主要从事植物抗逆分子生物学研究。E-mail: chkluo2002@163.com
中图分类号:S511;S332计量
文章访问数:120
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被引次数:0
出版历程
收稿日期:2020-12-10
录用日期:2021-03-11
刊出日期:2021-07-01
Physiological response strategies of roots of different alkali-tolerant rice varieties to alkali stress
LU Xuping,LI Fanglan,
MA Xiaojuan,
JING Panpan,
LUO Chengke,,
TIAN Lei,
LI Peifu
Agricultural College of Ningxia University/Key Laboratory of Modern Molecular Breeding of Dominant Characteristic Crops in Ningxia, Yinchuan 750021, China
Funds: the National Natural Science Foundation of China32060425
the National Natural Science Foundation of China31560297
the Natural Science Foundation of Ningxia2020AAC03095
More Information
Corresponding author:LUO Chengke, E-mail: chkluo2002@163.com
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摘要
摘要:为明确碱敏感和耐碱水稻品种幼苗根系响应碱胁迫的生理差异和应对策略，本试验以碱敏感水稻品种‘中花11’和耐碱品种‘宁粳52’为材料，将两种碱性盐（NaHCO3和Na2CO3）按照不同摩尔比混合，设3个碱浓度水平（10 mmol·L-1、20 mmol·L-1和30 mmol·L-1）和3个pH水平（8.65、9.55和10.50），模拟出9种碱胁迫环境，研究碱胁迫对供试水稻幼苗根系生长和相关抗逆生理指标的影响；并用逐步回归分析建立最优回归方程，进而筛选出不同胁迫程度下不同水稻品种响应碱胁迫的关键指标。结果表明：1）碱胁迫条件下‘中花11’的根系生长特征（根系总长度、根系总表面积、根系平均直径、根体积）和根系活力降幅大于‘宁粳52’，根系脂氧合酶（LOX）活性、丙二醛（MDA）含量、活性氧（O2·-、H2O2）含量均显著高于‘宁粳52’，而根系渗透调节物质[可溶性糖（SS）、可溶性蛋白（SP）和游离脯氨酸（Pro）]含量、抗氧化酶[超氧化物歧化酶（SOD）、过氧化氢酶（CAT）和过氧化物酶（POD）]活性、还原型抗坏血酸（ASA）和还原型谷胱甘肽（GSH）含量增幅均低于‘宁粳52’。2）在‘中花11’中，20C（碱浓度为20 mmol·L-1，pH 10.50）处理下根系平均直径、根系活力低于30A（碱浓度为30 mmol·L-1，pH 8.65）处理，MDA含量、LOX活性、O2·-产生速率、H2O2含量显著高于30A处理，即高pH低碱浓度对水稻的伤害大于低pH高碱浓度引起的伤害。3）冗余（RDA）分析表明Na+和pH的增加与供试水稻各生长指标的升高呈负相关，与抗逆生理指标的升高呈正相关。4）逐步回归分析表明，ASA、SS和H2O2含量是‘中花11’响应碱胁迫较为敏感的指标；根系平均直径、根系总表面积、根系活力、Pro、SS、SOD、POD和GSH是‘宁粳52’响应碱胁迫较为敏感的指标。综上，碱浓度和pH胁迫均影响了供试水稻根系生长特征和生理特性，在碱化土壤中栽培水稻需要同时考虑碱分组成和pH的影响，不同水稻品种在抵御碱胁迫时会启动不同的防御策略。
关键词:水稻/
碱胁迫/
pH/
根系/
生长/
生理/
响应策略
Abstract:This study aimed to investigate the physiological differences and coping strategies of the roots of rice varieties with different tolerances to alkali stress. In this experiment, alkali-sensitive rice variety 'Zhonghua 11' and alkali-tolerant rice variety 'Ninggeng 52' were subjected to nine alkali stress using two types of alkaline salts (sodium bicarbonate and sodium carbonate) in three alkali concentration levels (10 mmol·L-1, 20 mmol·L-1, and 30 mmol·L-1) and three pH levels (8.65, 9.55, and 10.50). The effects of alkali stress on the root growth and stress resistance of rice seedlings were examined. Through stepwise regression analysis, the optimal regression equation was established, and the key indices of the response of different rice varieties to alkali stress were screened. The results showed that: 1) the decline of root growth characteristics (total root length, total root surface area, average root diameter, and root volume) and root activity of 'Zhonghua 11' were greater than those of 'Ninggeng 52' (P < 0.05). The lysyl oxidase (LOX), malondialdehyde (MDA), superoxide anion (O2·-) producing rate, and hydrogen peroxide (H2O2) content in the roots of 'Zhonghua 11' were significantly higher than those of 'Ninggeng 52' (P < 0.05). The content of osmotic adjustment substances[soluble sugar (SS), soluble protein (SP), free proline (Pro)], the activities of antioxidant enzymes[superoxide dismutase (SOD), peroxidase (POD), catalase (CAT)], reduced ascorbic acid (ASA), and reduced glutathione (GSH) in the roots of 'Zhonghua 11' were lower than those of 'Ninggeng 52'. 2) In 'Zhonghua 11', the average root diameter and root activity of the 20C treatment (20 mmol·L-1, pH 10.50) were lower than those of the 30A treatment (30 mmol·L-1, pH 8.65). The MDA content, LOX activity, O2·- production rate, and H2O2 content of the 20C treatment were significantly higher than those of the 30A treatment, and the damage caused by high pH and a low alkali concentration was greater than that of low pH and a high alkali concentration. 3) Redundancy analysis showed that the increase in sodium ions (Na+) and pH was negatively correlated with the increased growth indexes and positively correlated with increases in the physiological indexes for the tested rice varieties. 4) Stepwise regression analysis showed that the ASA, SS, and H2O2 contents were sensitive indicators of alkali stress in 'Zhonghua 11'. The average root diameter, total root surface area, root activity, Pro content, SS content, SOD activity, POD activity, and GSH content were the sensitive indexes of alkali stress for 'Ninggeng 52'. In summary, alkali and pH stress affected the growth and physiological characteristics of rice roots of rice. The effects of alkali composition and pH must be considered when cultivating rice in alkaline soil, as different rice varieties initiate different defense strategies against different alkali stresses.
Key words:Rice/
Alkali stress/
pH/
Root system/
Growth/
Physiological/
Response strategy
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图1碱胁迫对水稻根系生长特征的影响
ZH11和NG52分别表示水稻品种‘中花11’和‘宁粳52’。CK: 不加碱对照; 10、20和30分别表示碱浓度10 mmol?L–1、20 mmol?L–1和30?mmol·L–1; A、B和C分别表示pH为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); *、**和ns分别表示相同胁迫处理不同品种在P < 0.05、P < 0.01水平差异显著和差异不显著。
Figure1.Effects of alkali stress on root growth characteristics of rice
ZH11 and NG52 respectively indicate rice varieties 'Zhonghua11' and 'Ninggeng52'. CK: no-alkali (control); 10, 20 and 30 respectively indicate alkali contentrations 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 lowercase letters indicate significant difference of the same rice variety among different treatments at P < 0.05 level; * and ** indicate significant difference at P < 0.05 and P < 0.01 levels, and "ns" indicate no significant difference, between two rice varieties under the same treatment, respectively.
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图2碱胁迫对水稻根系活力和渗透调节物质的影响
ZH11和NG52分别表示水稻品种‘中花11’和‘宁粳52’。CK: 不加碱对照; 10、20和30分别表示碱浓度10 mmol?L–1、20 mmol?L–1和30?mmol·L–1; A、B和C分别表示pH为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); *、**和ns分别表示相同胁迫处理不同品种在P < 0.05、P < 0.01水平差异显著和差异不显著。
Figure2.Effects of alkali stress on root activity and osmotic adjustment substances of rice
ZH11 and NG52 respectively indicate rice varieties 'Zhonghua11' and 'Ninggeng52'. CK: no-alkali (control); 10, 20 and 30 respectively indicate alkali contentrations 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 lowercase letters indicate significant difference of the same rice variety among different treatments at P < 0.05 level; * and ** indicate significant difference at P < 0.05 and P < 0.01 levels, and "ns" indicate no significant difference, between two rice varieties under the same treatment, respectively.
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图3碱胁迫对水稻根系脂质过氧化[丙二醛(MDA)含量和脂氧合酶(LOX)活性]和活性氧含量($\text{O}_2^{ \cdot - }$产生速率和H2O2含量)的影响
ZH11和NG52分别表示水稻品种‘中花11’和‘宁粳52’。CK: 不加碱对照; 10、20和30分别表示碱浓度10 mmol?L–1、20 mmol?L–1和30?mmol·L–1; A、B和C分别表示pH为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); *、**和ns分别表示相同胁迫处理不同品种在P < 0.05、P < 0.01水平差异显著和差异不显著。
Figure3.Effects of alkali stress on malonaldehyde (MDA) content, lipoxygenase (LOX) activity, ${\rm{O}}_{\rm{2}}^{ \cdot - }$ producing rate and H2O2 content in rice roots
ZH11 and NG52 respectively indicate rice varieties 'Zhonghua11' and 'Ninggeng52'. CK: no-alkali (control); 10, 20 and 30 respectively indicate alkali contentrations 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 lowercase letters indicate significant difference of the same rice variety among different treatments at P < 0.05 level; * and ** indicate significant difference at P < 0.05 and P < 0.01 levels, and "ns" indicate no significant difference, between two rice varieties under the same treatment, respectively.
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图4碱胁迫水稻根系抗氧化酶[超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)]活性的影响
ZH11和NG52分别表示水稻品种‘中花11’和‘宁粳52’。CK: 不加碱对照; 10、20和30分别表示碱浓度10 mmol?L–1、20 mmol?L–1和30?mmol·L–1; A、B和C分别表示pH为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); *、**和ns分别表示相同胁迫处理不同品种在P < 0.05、P < 0.01水平差异显著和差异不显著。
Figure4.Effects of alkali stress on antioxidant enzymes (superoxide dismutase, SOD; peroxidase, POD; catalase, CAT) activities of rice roots
ZH11 and NG52 respectively indicate rice varieties 'Zhonghua11' and 'Ninggeng52'. CK: no-alkali (control); 10, 20 and 30 respectively indicate alkali contentrations 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 lowercase letters indicate significant difference of the same rice variety among different treatments at P < 0.05 level; * and ** indicate significant difference at P < 0.05 and P < 0.01 levels, and "ns" indicate no significant difference, between two rice varieties under the same treatment, respectively.
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图5碱胁迫对水稻根系还原型抗坏血酸含量(ASA)和还原型谷胱甘肽(GSH)含量的影响
ZH11和NG52分别表示水稻品种‘中花11’和‘宁粳52’。CK: 不加碱对照; 10、20和30分别表示碱浓度10 mmol?L–1、20 mmol?L–1和30?mmol·L–1; A、B和C分别表示pH为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); *、**和ns分别表示相同胁迫处理不同品种在P < 0.05、P < 0.01水平差异显著和差异不显著。
Figure5.Effects of alkali stress on reduced ascorbic acid (ASA) and glutathione (GSH) contents in rice roots
ZH11 and NG52 respectively indicate rice varieties 'Zhonghua11' and 'Ninggeng52'. CK: no-alkali (control); 10, 20 and 30 respectively indicate alkali contentrations 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 lowercase letters indicate significant difference of the same rice variety among different treatments at P < 0.05 level; * and ** indicate significant difference at P < 0.05 and P < 0.01 levels, and "ns" indicate no significant difference, between two rice varieties under the same treatment, respectively.
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图6水稻根系生长及生理指标与胁迫因子间的RDA分析(A为‘中花11’, B为‘宁粳52’)
Pro: 游离脯氨酸含量; SS: 可溶性糖含量; SP: 可溶性蛋白含量; MDA: 丙二醛含量; LOX: 脂氧合酶活性; ${\rm{O}}_{\rm{2}}^{ \cdot - }$: ${\rm{O}}_{\rm{2}}^{ \cdot - }$产生速率; H2O2: H2O2含量; SOD: 超氧化物歧化酶活性; POD: 过氧化物酶活性; CAT: 过氧化氢酶活性; ASA: 还原型抗坏血酸含量; GSH: 还原型谷胱甘肽含量。
Figure6.RDA analysis between root growth and physiological indexes of rice and stress factors (A: Zhonghua11; B: Ninggeng52)
Pro: proline content; SS: soluble sugar content; SP: soluble protein content; MDA: malondialdehyde content; LOX: lipoxygenase activity; ${\rm{O}}_{\rm{2}}^{ \cdot - }$: ${\rm{O}}_{\rm{2}}^{ \cdot - }$ producing rate; H2O2: H2O2 content; SOD: superoxide dismutase activity; POD: peroxidased activity; CAT: catalase activity; ASA: reduced ascorbic acid content; GSH: glutathione content.
下载: 全尺寸图片幻灯片表1各处理碱组成和pH
Table1.Alkali composition and pH of each treatment
| 处理 Treatment | 总碱浓度 Total alkali concentration (mmol?L–1) | pH | 碱浓度 Alkali concentration (mmol?L–1) | |
| NaHCO3 | Na2CO3 | |||
| CK | 0 | 5.50±0.05 | 0 | 0 |
| 10A | 10 | 8.65±0.05 | 9 | 1 |
| 10B | 10 | 9.55±0.05 | 5 | 5 |
| 10C | 10 | 10.50±0.05 | 1 | 9 |
| 20A | 20 | 8.65±0.05 | 18 | 2 |
| 20B | 20 | 9.55±0.05 | 10 | 10 |
| 20C | 20 | 10.50±0.05 | 2 | 18 |
| 30A | 30 | 8.65±0.05 | 27 | 3 |
| 30B | 30 | 9.55±0.05 | 15 | 15 |
| 30C | 30 | 10.50±0.05 | 3 | 27 |
下载: 导出CSV表2不同耐碱性水稻品种生长及生理指标与胁迫程度的逐步回归分析
Table2.Stepwise regression analysis of growth and physiological indexes of different rice varieties with degree of alkali stress
| 胁迫程度 The degree of stress | 品种 Variety | 最优线性回归方程 Best multiple linear regressing equation | |
| 轻度(碱浓度为10 mmol?L–1, pH为8.65、9.55和10.50) Light (10 mmol?L–1 alkaline concentration with pH 8.65, 9.55 and 10.50) | 中花11 ZH11 | Y=?3.613+0.594X16 | (R2=0.963, F=183.909**) |
| 宁粳52 NG52 | Y=85.413+0.260X6?279.923X3?0.021X14 | (R2=0.995, F=516.403**) | |
| 中度(碱浓度为20 mmol?L–1, pH为8.65、9.55; 碱浓度为30 mmol?L–1, pH为8.65) Mild (20 mmol?L–1 alkaline concentration wiht pH 8.65, 9.55; 30 mmol?L–1 alkaline concentration wiht pH 8.65) | 中花11 ZH11 | Y=?37.443+4.270X7 | (R2=0.922, F=82.607**) |
| 宁粳52 NG52 | Y=73.434?257.696X3+0.042X13?2.323X2+0.969X7?71.776X5 | (R2=1.000, F=16 025.683**) | |
| 重度(碱浓度为20 mmol?L–1, pH为10.50; 碱浓度为30 mmol?L–1, pH为9.55和10.50) Sever (20 mmol?L–1 alkaline concentration wiht pH 10.50; 30 mmol?L–1 alkaline concentration wiht pH 9.55 and 10.50) | 中花11 ZH11 | Y=?35.018+13.051X12+0.549X16 | (R2=0.925, F=50.276**) |
| 宁粳52 NG52 | Y=?59.631+0.383X17+8.619X12+0.24X5 | (R2=0.998, F=859.791**) | |
| Y为胁迫程度; X为根系生长及生理参数, 包括根系总长度(X1)、根系总表面积(X2)、根系平均直径(X3)、根体积(X4)、根系活力(X5)、游离脯氨酸含量(X6)、可溶性糖含量(X7)、可溶性蛋白含量(X8)、丙二醛含量(X9)、脂氧合酶活性(X10)、O2·–产生速率(X11)、H2O2含量(X12)、超氧化物歧化酶活性(X13)、过氧化物酶活性(X14)、过氧化氢酶活性(X15)、还原型抗坏血酸含量(X16)、还原型谷胱甘肽含量(X17)。**表示P < 0.01水平显著相关。Y is alkali stress degree. X is growth and physiological parameters of roots, including total root length (X1), total root surface area (X2), average root diameter (X3), root volume (X4), root activity (X5), proline content (X6), soluble sugar content (X7), soluble protein content (X8), malondialdehyde (MDA) content (X9), lipoxygenase (LOX) activity (X10), ${\rm{O}}_{\rm{2}}^{ \cdot - }$ producing rate (X11), H2O2 content (X12), superoxide dismutase (SOD) activity (X13), peroxidased (POD) activity (X14), catalase (CAT) activity (X15), reduced ascorbic acid (ASA) content (X16), glutathione (GSH) content (X17). ** represents significant correlation at P < 0.01. | |||
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