干旱胁迫对桢楠幼树渗透调节与活性氧代谢的影响及施氮的缓解效应
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摘要
以二年生桢楠(Phoebe zhennan)幼树为研究对象,采用盆栽控水的方法,探讨了桢楠幼树在干旱胁迫下渗透调节和活性氧代谢的变化,以及施氮对桢楠幼树应对干旱胁迫能力的影响。试验先将土壤含水量调整到4个梯度(田间持水量的80%(80%FC)、50%FC、30%FC和15%FC),1周后测定受胁迫植株的相关生理指标,之后进行3个水平的施氮处理(对照N0,中氮MN,高氮HN,各施氮量分4次(即干旱梯度形成后第7、14、21和28天)分别施入)。在施氮结束后30天(即开始施肥处理后51天)再次测定各项生理指标。结果表明:1)干旱处理7天后,桢楠叶片中游离脯氨酸(Pro)和可溶性糖(SS)含量均随胁迫强度增大而显著增加,重度干旱(15%FC)下的Pro含量增加尤为明显,可溶性蛋白(SP)含量则呈先增加后降低的趋势。施氮后,各种土壤水分状态下的Pro含量进一步增加。水分充足和轻度干旱MN水平下,SS含量也增加,而在中度和重度干旱下的SS含量显著降低,HN水平各干旱状态下SS含量变化均不显著。施氮结束后30天时,80%FC和50%FC下的SP含量表现为施氮组低于对照组,而30%FC和15%FC下则相反。2)施氮前随着干旱胁迫的增强,桢楠幼树叶片中过氧化氢(H2O2)含量、超氧化物歧化酶(SOD)活性、过氧化氢酶(CAT)活性显著上升,而过氧化物酶(POD)活性呈先上升后下降的趋势。施氮后,H2O2含量总体上表现为减少趋势,且MN水平下降幅度最大,HN水平反而不利于降低H2O2的含量。3种酶活性的变化则因干旱程度和施氮水平的不同而呈现出不同的变化趋势。3)施氮前随着干旱胁迫的增强,叶片丙二醛(MDA)含量呈显著上升趋势,相对电导率(REC)先显著下降后显著上升;施氮后,除重度干旱胁迫外,其他各干旱处理植株的MDA含量都表现为在MN水平下有所下降,而在HN水平下有所回升,但在重度干旱时,无论是MN或HN处理,MDA含量均呈上升趋势,表明在重度干旱胁迫下,难以通过施氮的方式缓解干旱胁迫产生的伤害。4)双因素方差分析显示,施氮与干旱胁迫间具有极显著的交互效应。以上结果表明:施一定量的氮肥有利于缓解桢楠幼树受到的干旱胁迫,以年施氮量计,施中氮(N元素质量为1.35 g·株~(–1))对除重度干旱外的干旱胁迫具有一定的缓解作用,但施高氮(N元素质量为2.70 g·株~(–1))时反而会对植株造成不利影响。
Aims Two-year-old seedlings of Phoebe zhennan were used in this study to explore the responses of osmotic adjustment and active oxygen metabolism to drought stress and the mitigation effect of nitrogen application. Methods The soil water content was firstly adjusted to four treatment levels, i.e. 80% of field water holding capacity(80% FC), 50% FC, 30% FC and 15% FC, respectively. The physiological variables of plants were measured after one week, and then three nitrogen application rates, control(N0), medium nitrogen(MN) and high nitrogen(HN) were performed at an interval of 7 days for four times(7 d, 14 d, 21 d and 28 d, respectively). The same physiological variables were determined again one month after the accomplishment of nitrogen application. Important findings 1) The free proline(Pro) and soluble sugar(SS) contents in the leaves increased significantly with the aggravation of drought stress after 7 days of drought, but the content of soluble protein(SP) was firstly increased and then declined. The increase of Pro content was especially obvious under severe drought(15% FC). After nitrogen application, the content of Pro raise further, but the values varied in drought treatment. The SS contents under sufficient water supply(80% FC) and mild drought(50% FC) were decreased by MN, but it did not change significantly when supplied with HN despite the soil water content. After nitrogen application, the SP contents under 80% FC and 50% FC were lower than those of no exogenous N, while they were opposite response under 30% FC and 15% FC. 2) Before nitrogen application, with the aggravation of drought stress, the hydrogen peroxide(H_2O_2) content, superoxide dismutase(SOD) activity, catalase(CAT) activity increased significantly, and the peroxidase(POD) activity showed an up-down trend. After nitrogen application, the content of H_2O_2 was generally deceased at each water condition, with the maximum decrease at MN, while the HN treatment was not conducive to reduce the content of H_2O_2. The activities of three kinds of enzymes responded differently to the severity of drought and the level of nitrogen application. 3) Before nitrogen application, the content of malondialdehyde(MDA) in leaves increased significantly when the soil water content declined to and below 50% FC. The relative electrical conductivity(REC) was decreased at first, and followed by significant increase. Except severe drought(15% FC) stress, the MDA content showed a decreasing trend at MN, but a rebound at HN. As regards severe drought stress, however, the content of MDA increased at both MN and HN, indicating that nitrogen application is not a good choice to alleviate the damage caused by severe drought stress. 4)Two-factor ANOVA revealed an obvious interaction between nitrogen application and drought stress. In conclusion, a proper amount of nitrogen(1.35 g·a~(–1) for each sapling) could somewhat alleviate drought stress no severer than 15% FC on seedlings of Phoebe zhennan, but excessive nitrogen at rate of or more than 2.70 g·a~(–1) per sapling is not recommended.
Aims Two-year-old seedlings of Phoebe zhennan were used in this study to explore the responses of osmotic adjustment and active oxygen metabolism to drought stress and the mitigation effect of nitrogen application. Methods The soil water content was firstly adjusted to four treatment levels, i.e. 80% of field water holding capacity(80% FC), 50% FC, 30% FC and 15% FC, respectively. The physiological variables of plants were measured after one week, and then three nitrogen application rates, control(N0), medium nitrogen(MN) and high nitrogen(HN) were performed at an interval of 7 days for four times(7 d, 14 d, 21 d and 28 d, respectively). The same physiological variables were determined again one month after the accomplishment of nitrogen application. Important findings 1) The free proline(Pro) and soluble sugar(SS) contents in the leaves increased significantly with the aggravation of drought stress after 7 days of drought, but the content of soluble protein(SP) was firstly increased and then declined. The increase of Pro content was especially obvious under severe drought(15% FC). After nitrogen application, the content of Pro raise further, but the values varied in drought treatment. The SS contents under sufficient water supply(80% FC) and mild drought(50% FC) were decreased by MN, but it did not change significantly when supplied with HN despite the soil water content. After nitrogen application, the SP contents under 80% FC and 50% FC were lower than those of no exogenous N, while they were opposite response under 30% FC and 15% FC. 2) Before nitrogen application, with the aggravation of drought stress, the hydrogen peroxide(H_2O_2) content, superoxide dismutase(SOD) activity, catalase(CAT) activity increased significantly, and the peroxidase(POD) activity showed an up-down trend. After nitrogen application, the content of H_2O_2 was generally deceased at each water condition, with the maximum decrease at MN, while the HN treatment was not conducive to reduce the content of H_2O_2. The activities of three kinds of enzymes responded differently to the severity of drought and the level of nitrogen application. 3) Before nitrogen application, the content of malondialdehyde(MDA) in leaves increased significantly when the soil water content declined to and below 50% FC. The relative electrical conductivity(REC) was decreased at first, and followed by significant increase. Except severe drought(15% FC) stress, the MDA content showed a decreasing trend at MN, but a rebound at HN. As regards severe drought stress, however, the content of MDA increased at both MN and HN, indicating that nitrogen application is not a good choice to alleviate the damage caused by severe drought stress. 4)Two-factor ANOVA revealed an obvious interaction between nitrogen application and drought stress. In conclusion, a proper amount of nitrogen(1.35 g·a~(–1) for each sapling) could somewhat alleviate drought stress no severer than 15% FC on seedlings of Phoebe zhennan, but excessive nitrogen at rate of or more than 2.70 g·a~(–1) per sapling is not recommended.
引文
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