干旱胁迫下白皮松苗木电阻抗及生理指标的比较研究
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摘要
白皮松(Pinus bungeana Zucc.)是特产中国的珍贵树种,是我国北方主要的园林绿化树种和山区或干旱地带绿化造林优良树种。白皮松造林适宜采用3～4年生容器苗。本试验对4年生白皮松容器苗进行栽植前干旱(土壤相对含水量分别为B1:75～80%(正常浇水),B2:55～60%(轻度干旱),B3:35～40%(严重干旱))和栽植后干旱(土壤相对含水量A1:75～80%,A2:55～60%,A3:35～40%,A4:15～20%(极严重干旱))处理,通过一系列的生理指标的测定(电解质渗透率、可溶性糖、PV曲线参数等)来了解白皮松在栽植前后能够适应的干旱范围,为白皮松科学造林提供理论依据。通过对这些生理指标的分析,来探讨白皮松耐旱机理。另一方面,栽植前和栽植后干旱处理后,进行白皮松苗针叶和茎的电阻抗分析,并得到电阻抗图谱参数的变化规律,得出这些参数随各个生理参数的响应变化曲线,求证相关性。确定可以表征植物抗旱程度的电阻抗参数。为快速检测和评价植物抗旱性提供新的技术手段。主要结果如下:
1.栽植前干旱B3处理下,针叶含水量无显著降低,电解质渗透率增加,叶绿素含量和最大光合效率(Fv/Fm)降低。表明栽植前干旱B3处理对针叶有一定伤害。栽植后如果正常浇水,这些影响将消除;如果遭遇栽植后干旱,B3处理的受害程度不比B1和B2处理的深,表明白皮松容器苗造林在栽植前遭到严重干旱胁迫(35～40%)时不会影响造林后的表现。然而栽植后严重干旱胁迫A3和A4处理4周后,白皮松针叶的含水量降低,电解质渗透率增加,Fv/Fm降低,表明白皮松受害。因此应避免栽植后初期4周以上严重干旱胁迫。
2.栽植前和栽植后干旱都造成了针叶内可溶性糖和淀粉含量的降低,表明干旱下针叶内碳水化合物在输出。栽植后严重干旱A3和A4处理下,超氧化物岐化酶(SOD)活性随干旱胁迫,表现为“升高-降低-升高”,然后在长期的极严重干旱胁迫(A4)下降低,认为是白皮松针叶内主要的保护酶。过氧化物酶(POD)活性随干旱胁迫降低,只在试验最后,严重干旱处理(A3,A4)的POD活性升高,认为不是白皮松针叶内保护酶系统的关键酶。蛋白质含量在严重干旱胁迫(A3,A4)下,先诱导新的蛋白产生而使总蛋白量增加,然后在干旱后4周受到严重伤害后降低,表明蛋白质是重要的调节物质。超氧阴离子在严重干旱胁迫A4处理下始终保持高含量,表明A4处理下白皮松受到活性氧伤害。
3.栽植前和栽植后干旱对白皮松针叶的含水量和电解质渗透率的影响比对茎更早,更大。白皮松受到干旱后,针叶中产生的可溶性糖和淀粉向茎中转移,使茎中的可溶性糖含量增加,以利于茎的抗旱性。干旱胁迫下茎的淀粉含量降低,可能是转化成为了可溶性糖来增加渗透调节能力。栽植前和栽植后干旱都造成了根系活力的增强。
4.白皮松苗木在初始质壁分离时的相对含水量(RWCtlp)大部分都高于90%,所以认为白皮松苗木的忍耐脱水能力不强。在A3和A4干旱胁迫处理下,主要通过降低初始质壁分离时的渗透势(ψwtlp)和饱和含水时的渗透势(ψssat)来维持最低膨压和保持最大膨压,从而增强抗旱性。在A2和A3处理下,通过增加束缚水含量,和轻微地增加忍耐脱水能力来增强抗旱性。在A1和A4处理下,还可以通过调节细胞壁弹性来增强抗旱性。
5.栽植前干旱处理下,针叶的电阻抗图谱和参数变化不大。栽植后干旱处理引起电阻抗图谱和参数发生改变。针叶的胞内电阻率(ri)和高频电阻(r∞)随干旱时间延长,在A1和A2处理下升高,在A3和A4处理下变化不大。针叶的胞外电阻率(re)和细胞膜时间恒量(τm)在A4处理下大幅度降低。电阻抗参数和针叶的含水量、相对电导率、叶绿素含量、F_v/F_m、可溶性糖含量的变化密切相关。其中,r_e和τ_m变化规律性更强,与F_v/F-m(r=0.54～0.89)、含水量(r=0.74～0.98)、可溶性糖含量(r=0.52～0.96)在3周以上为线性正相关,与相对电导率(r=-0.54～-0.97)、叶绿素含量为线性负相关(r=-0.62～-0.8),因此,认为这两个参数可以作为估测白皮松针叶抗旱性的最佳指标。
6.茎的r_1, r_2, r_e和弛豫时间τ2在严重干旱胁迫A4下降低,r和r_i变化不大;茎的弛豫时间τ_1在A4处理干旱胁迫下,前4周高于A1,在干旱5周后显著降低;在干旱胁迫5周后,茎的弛豫时间的分布系数ψ2在A2,A3和A4处理下比A1处理的高。其中r1和re在A4处理下变化规律更强,与含水量、相对电导率、可溶性糖含量、淀粉含量、根系活力都在3周以上有显著相关性(R2>0.5),与淀粉含量(r=0.51～0.97)、含水量(r=0.71～0.97)在3周以上为线性正相关,和相对电导率为负相关(r=-0.75～-0.99),因此认为这两个参数可以作为估测白皮松茎抗旱性的最佳指标。
Pinus bungeana Zucc. is a Chinese special precious species. It is a kind of main landscaping species in the northern part of China and the excellent tree of afforestation in arid region and mountainous areas. Three or four year-old container seedlings are suitable for the forestation. In this experiment, we treated 4-year-old P. bungeana container seedlings with preplanting (soil relative water content was B1: 75-80% (normal irrigation), B2: 55-60% (light drought), and B3: 35-40% (severe drought)) and postplanting (A1: 75-80%, A2: 55-60%, A3: 35-40%, and A4: 15-20%) drought, measured a series of physiologic indexes (Electrolyte leakage, soluable sugar, PV curve parameters and so on), aiming to find what scope of drought the P. bungeana could tolerate before and after planting, and offer the theoretical guide for scientific forestation. Based on the analysis of the physiologic indexes, it is aimed to explore the mechanism of drought resistance of P. bungeana. In addition, after preplanting and postplanting drought treatments, we measured the electrical impedance parameters of needles and stems of P. bungeana, and followed the change rules of the parameters. The correlation between electrical impedance parameters and various physiological parameters was studied to find the best impedance parameters which can estimate the plant drought-resistance degree, and provide a technical method for fast monitoring and assessing plant drought-resistance. Main results are as follows:
1. Under preplanting B3 treatment (severe drought), water content of needles did not decrease significantly, electrolyte leakage of needles increased, chlorophyll content and maximal PS II efficiency (F_v/F_m ) of needles declined significantly. These changes suggested that needles were damaged to some extent. Such effects would be ignored if normal irrigation was given after planting. However, if postplanting drought happened, injury degree of B3 treatment was not deeper than that of B1 and B2. So for preplanting severe drought of container seedlings would not affect the seedlings performance after planting. After 4 weeks A3 and A4 severe drought treatments, water content of needles decreased, electrolyte leakage of needles increased, and F_v/F_m of needles declined significantly, indicating that needles were injured. We therefore suggest that in the early phase after planting, more than 4 weeks severe drought should be avoided.
2. Preplanting and postplanting drought caused decrease in soluble sugar and starch content of needles, suggesting that the carbohydrates in needles outputed under the drought stress. Under postplanting severe drought A3 and A4 treatments, Superoxide dismutase (SOD) changed with a trend of“rise– decrease– rise”, then decreased in long-term extremely severe drought stress (A4). So SOD is thought as main protective enzyme in needles of P. bungeana seedlings. Peroxidase (POD) activity decreased with drought stress in the early four weeks, and r_ised only after five weeks severe drought treatments (A3, A4). Thus, POD activity is not regarded as key enzyme of protective enzyme system in needles of P. bungeana seedlings. The total protein content increased in severe drought stress (A3, A4) through producing new protein in the early drought phase, and decreased after four weeks ser_ious drought injury, showing that protein was an important regulating substance. Super oxide anion always remained high in severe drought stress (A4), showed that P. bungeana seedlings was damaged by active oxygen under A4 treatment.
3. Effects of preplanting and postplanting drought on water content and electrolyte leakage of needles in P. bungeana seedlings were earlier and larger than those of stems. Soluble sugar and starch produced in needles transfered to stems under drought stress, to make the soluble sugar content of stems increased, and to strengthen drought resistance of the stems. Starch content of stems reduced under drought stress, probably because starch content in stems transformed to soluble sugar to increase the osmotic adjustment ability. Preplanting and postplanting drought caused the enhance of root activity.
4. The relative water content at incipient plasmolysis (RWCtlp) of P. bungeana seedlings was mostly higher than 90%. So we think endurance dehydration capability of P. bungeana seedlings was not strong. Under A3 and A4 drought stress, P. bungeana seedlings mainly decreased osmotic potential at incipient plasmolysis (ψwtlp) and maximum osmotic potential at saturation point (ψssat) to maintain minimum turgor pressure and maximum turgor pressure, thereby increasing drought resistance. Under A2 and A3 drought strss, P. bungeana seedlings increased drought resistance through increasing bound water volume and slightly raising endurance dehydration capability. Under A1 and A4 treatments, P. bungeana seedlings also adjusted cell wall elasticity to increase the drought resistance. 5. After preplanting drought treatments, electr_ical impedance spectra and parameters of needles did not change a lot. Postplanting drought treatments caused the changes in electr_ical impedance spectra and parameters. With prolonged drought, intracellular resistance (r_i) and high frequency resistance (r∞) of needles r_ised under A1 and A2 treatments, and changed little under A3 and A4 treatments. Extracellular resistance (re) and membrane time constant (τ_m) decreased greatly under A4 treatment. Some impedance parameters closely related to water content, relative electr_ical conductivity, chlorophyll content, F_v/F_m , and soluble sugar content in needles. Among these impedance parameters, re andτ_m correlated better, and had linear positive correlations with F_v/F_m (r=0.54～0.89), water content (r=0.74～0.98), and soluble sugar content (r=0.52～0.96) in three weeks, while had negative linear correlations with relative electr_ical conductivity (r=-0.54～-0.97), and chlorophyll content (r=-0.62～-0.8). Therefore, re andτ_m could be regarded as the best parameters to estimate the drought resistance of needles in P. bungeana seedlings.
6. Impedance parameters r_1, r2, re and relaxation timeτ2 of stems decreased, whereas r and r_i did not change a lot under severe drought stress (A4). Relaxation timeτ1 of stems under A4 treatment was higher than that under A1 in the first four weeks, and decreased significantly after five weeks drought. After five weeks drought stress, distr_ibution coefficient (ψ2 ) of relaxation timeτ2 of stems was higher under A2, A3 and A4 treatments than that under A1 treatment. Among the impedance parameters, r_1 and re changed regularly under A4 drought treatment, and had significant correlations with water content, relative electrical conductivity, soluble sugar content, starch content and root activity in three weeks (R~2>0.5), and had positive linear correlations with starch content (r=0.51～0.97) and water content (r=0.71～0.97), negative linear correlations with relative electrical conductivity (r=-0.75～-0.99) in three weeks. So r_1 and re could be regarded as the best parameters to estimate the drought resistance of stems in P. bungeana seedlings.
1.栽植前干旱B3处理下,针叶含水量无显著降低,电解质渗透率增加,叶绿素含量和最大光合效率(Fv/Fm)降低。表明栽植前干旱B3处理对针叶有一定伤害。栽植后如果正常浇水,这些影响将消除;如果遭遇栽植后干旱,B3处理的受害程度不比B1和B2处理的深,表明白皮松容器苗造林在栽植前遭到严重干旱胁迫(35～40%)时不会影响造林后的表现。然而栽植后严重干旱胁迫A3和A4处理4周后,白皮松针叶的含水量降低,电解质渗透率增加,Fv/Fm降低,表明白皮松受害。因此应避免栽植后初期4周以上严重干旱胁迫。
2.栽植前和栽植后干旱都造成了针叶内可溶性糖和淀粉含量的降低,表明干旱下针叶内碳水化合物在输出。栽植后严重干旱A3和A4处理下,超氧化物岐化酶(SOD)活性随干旱胁迫,表现为“升高-降低-升高”,然后在长期的极严重干旱胁迫(A4)下降低,认为是白皮松针叶内主要的保护酶。过氧化物酶(POD)活性随干旱胁迫降低,只在试验最后,严重干旱处理(A3,A4)的POD活性升高,认为不是白皮松针叶内保护酶系统的关键酶。蛋白质含量在严重干旱胁迫(A3,A4)下,先诱导新的蛋白产生而使总蛋白量增加,然后在干旱后4周受到严重伤害后降低,表明蛋白质是重要的调节物质。超氧阴离子在严重干旱胁迫A4处理下始终保持高含量,表明A4处理下白皮松受到活性氧伤害。
3.栽植前和栽植后干旱对白皮松针叶的含水量和电解质渗透率的影响比对茎更早,更大。白皮松受到干旱后,针叶中产生的可溶性糖和淀粉向茎中转移,使茎中的可溶性糖含量增加,以利于茎的抗旱性。干旱胁迫下茎的淀粉含量降低,可能是转化成为了可溶性糖来增加渗透调节能力。栽植前和栽植后干旱都造成了根系活力的增强。
4.白皮松苗木在初始质壁分离时的相对含水量(RWCtlp)大部分都高于90%,所以认为白皮松苗木的忍耐脱水能力不强。在A3和A4干旱胁迫处理下,主要通过降低初始质壁分离时的渗透势(ψwtlp)和饱和含水时的渗透势(ψssat)来维持最低膨压和保持最大膨压,从而增强抗旱性。在A2和A3处理下,通过增加束缚水含量,和轻微地增加忍耐脱水能力来增强抗旱性。在A1和A4处理下,还可以通过调节细胞壁弹性来增强抗旱性。
5.栽植前干旱处理下,针叶的电阻抗图谱和参数变化不大。栽植后干旱处理引起电阻抗图谱和参数发生改变。针叶的胞内电阻率(ri)和高频电阻(r∞)随干旱时间延长,在A1和A2处理下升高,在A3和A4处理下变化不大。针叶的胞外电阻率(re)和细胞膜时间恒量(τm)在A4处理下大幅度降低。电阻抗参数和针叶的含水量、相对电导率、叶绿素含量、F_v/F_m、可溶性糖含量的变化密切相关。其中,r_e和τ_m变化规律性更强,与F_v/F-m(r=0.54～0.89)、含水量(r=0.74～0.98)、可溶性糖含量(r=0.52～0.96)在3周以上为线性正相关,与相对电导率(r=-0.54～-0.97)、叶绿素含量为线性负相关(r=-0.62～-0.8),因此,认为这两个参数可以作为估测白皮松针叶抗旱性的最佳指标。
6.茎的r_1, r_2, r_e和弛豫时间τ2在严重干旱胁迫A4下降低,r和r_i变化不大;茎的弛豫时间τ_1在A4处理干旱胁迫下,前4周高于A1,在干旱5周后显著降低;在干旱胁迫5周后,茎的弛豫时间的分布系数ψ2在A2,A3和A4处理下比A1处理的高。其中r1和re在A4处理下变化规律更强,与含水量、相对电导率、可溶性糖含量、淀粉含量、根系活力都在3周以上有显著相关性(R2>0.5),与淀粉含量(r=0.51～0.97)、含水量(r=0.71～0.97)在3周以上为线性正相关,和相对电导率为负相关(r=-0.75～-0.99),因此认为这两个参数可以作为估测白皮松茎抗旱性的最佳指标。
Pinus bungeana Zucc. is a Chinese special precious species. It is a kind of main landscaping species in the northern part of China and the excellent tree of afforestation in arid region and mountainous areas. Three or four year-old container seedlings are suitable for the forestation. In this experiment, we treated 4-year-old P. bungeana container seedlings with preplanting (soil relative water content was B1: 75-80% (normal irrigation), B2: 55-60% (light drought), and B3: 35-40% (severe drought)) and postplanting (A1: 75-80%, A2: 55-60%, A3: 35-40%, and A4: 15-20%) drought, measured a series of physiologic indexes (Electrolyte leakage, soluable sugar, PV curve parameters and so on), aiming to find what scope of drought the P. bungeana could tolerate before and after planting, and offer the theoretical guide for scientific forestation. Based on the analysis of the physiologic indexes, it is aimed to explore the mechanism of drought resistance of P. bungeana. In addition, after preplanting and postplanting drought treatments, we measured the electrical impedance parameters of needles and stems of P. bungeana, and followed the change rules of the parameters. The correlation between electrical impedance parameters and various physiological parameters was studied to find the best impedance parameters which can estimate the plant drought-resistance degree, and provide a technical method for fast monitoring and assessing plant drought-resistance. Main results are as follows:
1. Under preplanting B3 treatment (severe drought), water content of needles did not decrease significantly, electrolyte leakage of needles increased, chlorophyll content and maximal PS II efficiency (F_v/F_m ) of needles declined significantly. These changes suggested that needles were damaged to some extent. Such effects would be ignored if normal irrigation was given after planting. However, if postplanting drought happened, injury degree of B3 treatment was not deeper than that of B1 and B2. So for preplanting severe drought of container seedlings would not affect the seedlings performance after planting. After 4 weeks A3 and A4 severe drought treatments, water content of needles decreased, electrolyte leakage of needles increased, and F_v/F_m of needles declined significantly, indicating that needles were injured. We therefore suggest that in the early phase after planting, more than 4 weeks severe drought should be avoided.
2. Preplanting and postplanting drought caused decrease in soluble sugar and starch content of needles, suggesting that the carbohydrates in needles outputed under the drought stress. Under postplanting severe drought A3 and A4 treatments, Superoxide dismutase (SOD) changed with a trend of“rise– decrease– rise”, then decreased in long-term extremely severe drought stress (A4). So SOD is thought as main protective enzyme in needles of P. bungeana seedlings. Peroxidase (POD) activity decreased with drought stress in the early four weeks, and r_ised only after five weeks severe drought treatments (A3, A4). Thus, POD activity is not regarded as key enzyme of protective enzyme system in needles of P. bungeana seedlings. The total protein content increased in severe drought stress (A3, A4) through producing new protein in the early drought phase, and decreased after four weeks ser_ious drought injury, showing that protein was an important regulating substance. Super oxide anion always remained high in severe drought stress (A4), showed that P. bungeana seedlings was damaged by active oxygen under A4 treatment.
3. Effects of preplanting and postplanting drought on water content and electrolyte leakage of needles in P. bungeana seedlings were earlier and larger than those of stems. Soluble sugar and starch produced in needles transfered to stems under drought stress, to make the soluble sugar content of stems increased, and to strengthen drought resistance of the stems. Starch content of stems reduced under drought stress, probably because starch content in stems transformed to soluble sugar to increase the osmotic adjustment ability. Preplanting and postplanting drought caused the enhance of root activity.
4. The relative water content at incipient plasmolysis (RWCtlp) of P. bungeana seedlings was mostly higher than 90%. So we think endurance dehydration capability of P. bungeana seedlings was not strong. Under A3 and A4 drought stress, P. bungeana seedlings mainly decreased osmotic potential at incipient plasmolysis (ψwtlp) and maximum osmotic potential at saturation point (ψssat) to maintain minimum turgor pressure and maximum turgor pressure, thereby increasing drought resistance. Under A2 and A3 drought strss, P. bungeana seedlings increased drought resistance through increasing bound water volume and slightly raising endurance dehydration capability. Under A1 and A4 treatments, P. bungeana seedlings also adjusted cell wall elasticity to increase the drought resistance. 5. After preplanting drought treatments, electr_ical impedance spectra and parameters of needles did not change a lot. Postplanting drought treatments caused the changes in electr_ical impedance spectra and parameters. With prolonged drought, intracellular resistance (r_i) and high frequency resistance (r∞) of needles r_ised under A1 and A2 treatments, and changed little under A3 and A4 treatments. Extracellular resistance (re) and membrane time constant (τ_m) decreased greatly under A4 treatment. Some impedance parameters closely related to water content, relative electr_ical conductivity, chlorophyll content, F_v/F_m , and soluble sugar content in needles. Among these impedance parameters, re andτ_m correlated better, and had linear positive correlations with F_v/F_m (r=0.54～0.89), water content (r=0.74～0.98), and soluble sugar content (r=0.52～0.96) in three weeks, while had negative linear correlations with relative electr_ical conductivity (r=-0.54～-0.97), and chlorophyll content (r=-0.62～-0.8). Therefore, re andτ_m could be regarded as the best parameters to estimate the drought resistance of needles in P. bungeana seedlings.
6. Impedance parameters r_1, r2, re and relaxation timeτ2 of stems decreased, whereas r and r_i did not change a lot under severe drought stress (A4). Relaxation timeτ1 of stems under A4 treatment was higher than that under A1 in the first four weeks, and decreased significantly after five weeks drought. After five weeks drought stress, distr_ibution coefficient (ψ2 ) of relaxation timeτ2 of stems was higher under A2, A3 and A4 treatments than that under A1 treatment. Among the impedance parameters, r_1 and re changed regularly under A4 drought treatment, and had significant correlations with water content, relative electrical conductivity, soluble sugar content, starch content and root activity in three weeks (R~2>0.5), and had positive linear correlations with starch content (r=0.51～0.97) and water content (r=0.71～0.97), negative linear correlations with relative electrical conductivity (r=-0.75～-0.99) in three weeks. So r_1 and re could be regarded as the best parameters to estimate the drought resistance of stems in P. bungeana seedlings.
引文
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