崩岗侵蚀区先锋植物类芦对环境胁迫的生理生态学响应机制
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摘要
类芦是南方崩岗侵蚀区优良的水土保持草种,能快速覆盖裸露山体,成为优势种群。它不仅能在贫瘠和干旱的恶劣环境下生存,而且能形成强大的根网,固土护坡效果极为显著。类芦能在养分贫瘠和干旱的立地上表现出旺盛生命力,是长期适应恶劣生存环境的结果,是长期进化过程中形成的特殊适应性,可能存在着独特的觅寻环境养分和水分机制,有着特殊的生态学意义,但目前国内对类芦适应贫瘠养分和干旱环境的内在机制还了解很少。
有鉴于此,本文根据胁迫植物学的原理和方法,以南方崩岗侵蚀区广泛分布的类芦为研究对象,利用自主设计的环境养分和水分胁迫模拟装置,通过不同的养分和水分胁迫试验,研究养分和水分胁迫条件下类芦的形态学和生理学特征变化,分析类芦在不同环境胁迫条件下形态学和生理学特征的差异,探讨类芦对环境土壤养分和水分的吸收利用策略,挖掘类芦这种特殊生物学特性的生态学适应意义,为揭示类芦适应贫瘠和干旱崩岗侵蚀区土壤的内在机制提供科学依据。主要研究结果如下：
(1)干旱胁迫对类芦地上部分生长有较大影响。干旱胁迫初期不同干旱处理对类芦分蘖和叶片数量影响较大。与充足供水处理相比,中度和轻度干旱胁迫处理类芦分蘖和叶片数量明显较多(p<0.05),但株高生长能力明显减弱。在重度干旱胁迫条件下类芦叶片长度生长明显受阻,但叶片宽度生长较快。随干旱胁迫时间的延长,类芦株高、分蘖能力、叶片数量及最大叶宽生长没有受到明显抑制,但叶片的伸长能力受阻。
(2)干旱胁迫对类芦根系有较大影响,不同处理类芦根系总长度、表面积、平均直径、根系生物量存在明显差异。随干旱胁迫程度的增加,根体积、根平均直径减小；而类芦根系总长度、根表面积、根系生物量、根冠比均比充足供水处理大；在中度干旱胁迫条件下类芦的根系长度、根表面积、根系生物量、地上部生物量、根冠比最大。类芦根系在环境胁迫条件下增强延伸能力、扩大与土壤接触面积可能是其对干旱逆境的形态学响应机制之一。(3)干旱胁迫对类芦光合特性有较大影响。在中度和轻度干旱胁迫条件下类芦叶绿素含量增加,叶绿素a/b比值下降,叶绿素荧光参数F0、Fm、Fv提高,Fv/Fm、Fv/F0下降。类芦叶片的胞间CO2浓度随干旱程度加重而逐渐增加,表现为上午缓慢上升、下午快速上升,16：00达到最大值。净光合速率、气孔导度、蒸腾速率日变化均表现为“单峰型”,为先升后降,12：00达到高峰,日均值随干旱程度加重而逐渐下降。在中度和轻度干旱胁迫下类芦WUE提高,重度干旱处理WUE稍有下降,说明类芦具有较强的耐干旱能力。
(4)不同干旱胁迫处理类芦叶片的生理学指标存在明显差异。干旱胁迫对类芦叶片保护酶活性影响较大,供水充足处理类芦叶片的POD、SOD和CAT活性明显高于轻度胁迫、中度胁迫和重度胁迫处理。从叶片膜系统来看,干旱胁迫越严重,类芦叶片的MDA含量越高,而当水分供应充足时,其MDA含量同样呈升高趋势；叶片相对电导率随胁迫强度的增强逐渐增加。干旱胁迫对类芦叶片可溶性蛋白和芦根系分泌酸性物质的影响表现为随干旱胁迫加剧而有所降低。
(5)不同P素胁迫对类芦生物量分配格局及根系形态指标有明显影响。中度P胁迫处理类芦根系总长度、总表面积和总体积明显大于其它处理,中度P胁迫有利于类芦根长、表面积及体积的增长,但不利于根直径生长。P素胁迫对类芦秆、叶、根和生物积累量的影响均表现出随P胁迫程度的加剧呈上升趋势,然后逐渐下降。
(6)不同N胁迫处理的类芦根系总长度、表面积、根体积和根系生物量存在差异明显。不同N胁迫下根长、根表面、根体积、根系生物量随N胁迫程度增加而逐渐减小。在N素极度匮乏逆境中,类芦生长缓慢,但根系生物量分配比例比地上部分高,这可能是类芦抵抗环境N胁迫的一种形态学机制之一。
(7)不同K胁迫处理类芦生物量分配格局及根系形态指标存在明显差异。不供K处理类芦根系总长度、总表面积和总体积均明显大于其它处理。随供K量的减少,类芦根系平均直径、全株生物量呈递减趋势,根冠比呈增加趋势。
(8)不同P胁迫处理类芦叶片生理学指标存在明显差异。P胁迫一定程度上增强了类芦叶片保护酶活性,与正常供P处理相比,P胁迫处理类芦叶片POD、SOD和CAT活性均明显增强(p<0.05),但随P胁迫程度的加剧,各种酶活性均降低。从叶片膜系统来看,随P胁迫程度加剧,类芦叶片MDA含量和相对电导率逐渐上升,而当P胁迫达到一定程度时,两者呈降低趋势。在中度P胁迫类芦叶片可溶性蛋白和根系分泌酸性物质含量明显高于其它处理。
(9)N胁迫对类芦叶片生理学指标有明显影响。N胁迫对类芦叶片保护酶活性影响明显,正常供N、轻度N胁迫、中度N胁迫处理类芦叶片POD、SOD和CAT活性逐渐增强,但随N胁迫程度的加剧,重度N胁迫和不供N处理保护酶活性依次降低。从叶片膜系统来看,随N胁迫程度的加剧,类芦叶片MDA含量和相对电导率逐渐上升,而当N胁迫达一定程度时,两者均呈降低趋势。重度N胁迫和中度N胁迫处理类芦叶片可溶性蛋白和根系分泌酸性物质含量均明显高于其它处理(p<0.05)。
(10)不同K胁迫处理类芦叶片生理学指标存在明显差异。K胁迫一定程度上可增强类芦叶片保护酶活性,中度K胁迫类芦叶片POD、SOD和CAT活性较高,正常供K处理中各种酶活性最低,明显低于其它K胁迫处理(p<0.05)。从类芦叶片膜系统来看,中度K胁迫处理的MDA含量最高,而叶片相对电导率则随K胁迫程度的加剧呈升高趋势。类芦叶片可溶性蛋白和根系分泌酸性物质含量随K胁迫程度的加剧而表现出先升高后下降趋势。
综合以上研究结果,崩岗侵蚀区先锋植物类芦对环境水分胁迫的响应策略为：当环境土壤田间持水量小于40%形成干旱胁迫时,一方面,芦类的分蘖数量增加,通过增加叶片数量与宽度、减小叶片长度,以减少土壤水分的散失速度。另一方面,干旱胁迫条件下类芦体内叶片胞间CO：浓度、叶绿素、MDA含量和相对电导率显著增加,一定程度上提高了类芦体内的水分利用效率。随干旱胁迫强度的加剧,叶片可溶性蛋白和根系分泌酸性物质含量呈现先增加后降低趋势,但根系生物量呈上升趋势,根冠比不断增加,主要通过根系表面积的扩增以及根系向外延伸,加大根系与土壤的接触面积,增强对环境土壤水分的主动寻觅能力,以维持植株正常生长对水分的需求,这可能是芦类能在极其干旱的环境中正常生长的主要原因。
类芦对P素胁迫、N素胁迫和K素胁迫在一定程度上表现出相似的响应策略,其对环境养分胁迫的响应机制为：当土壤含P量、含N量、含K量分别小于0.50mmol·L-1、3.75mmol·L-1、3.0mmol·L-1形成中度或重度养分胁迫时,类芦叶片POD、SOD和CAT酶活性增强,叶片可溶性蛋白和根系分泌酸性物质含量有所增加,类芦秆、叶、根和全株生物积累量也随之增加,根冠比增大,根系总长度、总表面积和体积明显增加,从而提高了类芦根系在养分胁迫逆境中的主动寻觅养分的能力。但类芦根系的这种主动响应机制只能维持一小段时间,随养分胁迫时间的延长,类芦叶片酶活性及其数量降低,生长变缓,因此类芦对环境贫瘠养分的适应有一定限度。
Neyraudia reynaudiana is a pioneer Plant for water and soil conservation in the southern collapsing hill area. They can recovery rapidly in the bare soil and then become dominance species. Neyraudia reynaudiana can survive and grow in nutrient deficiency and drought soil, which show it may possesses specific mechanisms to find the nutrient and moisture in poor soil. However, the adaption mechanism of Neyraudia reynaudiana to nutrients deficiency and drought condition is still not determined clearly.
For these reasons, the morphological and physiological characteristics of Neyraudia reynaudiana were determined under nutrient and drought stress based on the method of Stress Botany. The differences between the morphological and physiological features were analyzed in different stress treatments. The strategy to find the nutrients and moisture in poor soil was discussed for Neyraudia reynaudiana, which would be benefit for understanding its adaption mechanism to nutrients and drought stress. The main results were as follows:
(1) There was significant effect on the growth of Neyraudia reynaudiana among the different water stress treatments. In the early stage of drought stress, the moisture content in the soil had a greater effect on the number of tillers and leaves of Neyraudia reynaudiana. Compared with adequate water supply treatment, the numbers of tillers and leaf of Neyraudia reynaudiana were significantly more both in moderate and slight water stress treatment, however, the growths of their height were significantly reduced (p<0.05). The growth of leaf length was weaned in the severe drought stress treatment, however, the growth of its leaf width was speeded. AS the drought stress goes, the growth of its height, tiller ability, leaf number and maximum leaf width were inhibited, however, leaf elongation ability weakened significantly.
(2) There was significantly difference in total root length, root surface area, root average diameter, root biomass of Neyraudia reynaudiana among different drought stress treatments, however, not in the root volume. As the drought stress increased, the volume and diameter of roots reduced gradually in different drought stress. Compared with the adequate water treatment, the total length, surface area, biomass of roots, and root-shoot ratio were high significantly under different drought stres. The total length, surface area, biomass of roots, aboveground biomass, root-shoot ratio were the highest under moderate drought stress. Neyraudia reynaudiana had strong drought-tolerant because of large root system with root length and root surface area. The root characters of elongation growth and enlargement root-soil interface of Neyraudia reynaudiana may be one of morphology response mechanisms to drought stress.
(3) The effects of drought stress on photosynthetic characteristics of Neyraudia reynaudiana were significantly different.. Chlorophyll content of Neyraudia reynaudiana increased and chlorophyll a/b was decrease under moderate and slight drought stress. Chlorophyll fluorescence parameters F0, Fm, Fv increased and Fv/Fm, Fv/F0decreased under drought stress. The average intercellular CO2content increased as the drought stress increasing. They increase slowly in the morning and rapidly increase in the afternoon to maximum at16:00. The diurnal variation curves of net photosynthetic rate (Pn), stomatal conductance (Gs), transpiration rate (Tr) were increasing, and then decreasing, a single-peak at12:00. Pn, Gs, and Tr decreased as drought stress increasing. The WUE of Neyraudia reynaudiana increased under moderate and slight drought stress, and decreased slightly under heavy drought stress. Therefore, Neyraudia reynaudiana had strong drought-resistant capacity.
(4) In the different drought stress treatments, there were significantly differences among the physiological process of Neyraudia reynaudiana. The effects of drought stress on protection enzymes of Neyraudia reynaudiana were significant. The activities of POD, SOD and CAT in leaves of Neyraudia reynaudiana under adequate water supply treatment were significantly higher than the slight, moderate and severe stress. From the leaf membrane system, the more serious the drought stress, the higher MDA content of Neyraudia reynaudiana.The MDA content showed the increasing trend under adequate water supply. The relative Conductivity of leaf gradually increased with increasing of drought stress. The effects of drought stress on the leaf soluble proteins and secrete acidic substances in roots of Neyraudia reynaudiana were showed as follows: their contents were decreased with increasing of drought stress.
(5) Among the different low phosphorus stress treatments, there were significant effects on biomass distribution pattern and root morphology indicators of Neyraudia reynaudiana. The total root length, total root surface area and total root volume of Neyraudia reynaudiana were significantly higher than other treatments under moderate stress. The moderate stress was more benefit for growth of root length, root surface area and root volume, however, not conducive to root diameter growth. The effects on stalks, leaves, roots, and biomass accumulation of Neyraudia reynaudiana under the low phosphorus stress showed an upward trend with increase of phosphorus stress and then gradually decline.
(6) Among the different nitrogen stress treatments, there was significantly different in total root length, root surface area, root average diameter, root biomass of Neyraudia reynaudiana, However, not in the average diameter. The root length, root surface area, root volume and root biomass decreased with increase of nitrogen stress. The Neyraudia reynaudiana grew slowly under the extreme shortage of nitrogen in the soil, however, root biomass distribution ratio was higher than the aerial parts which may be an ecological resistance mechanism of Neyraudia reynaudiana under nitrogen stress.
(7) There was significantly difference in biomass allocation pattern and root morphology indicators of Neyraudia reynaudiana under different potassium stress treatments. The total root length, total root surface area and total root volume of Neyraudia reynaudiana without potassium supply treatment were significantly higher than other treatments. The root average diameter of Neyraudia reynaudiana showed a decreasing trend with the decrease of potassium. The whole plant biomass showed an increasing trend and the root-shoot ratio showed a declining trend with the increase of potassium.
(8) There was significantly difference in the leaves physiology indicators of Neyraudia reynaudiana among different phosphorus stress treatments. To some extent, the leaf protection enzymes of Neyraudia reynaudiana were enhanced under low phosphorus stress. Compared with normal phosphorus supply treatment, the activities of POD, SOD and CAT in Neyraudia reynaudiana leaves under low phosphorus stress were significantly increased (p<0.05), however, with the intensification of phosphorus stress, each protection enzymes activities consistently decreased under severe phosphorus stress and not phosphorus supply treatments. From the leaf membrane system, the MDA content and relative conductivity gradually increased with the intensification of phosphorus stress, while the phosphorus stress reaches a certain level, both showed a decreasing trend. The leaf soluble proteins content and secrete acidic substances content in roots of Neyraudia reynaudiana were significantly higher than other treatments under moderate stress.
(9) There was significantly difference in the leaves physiology indicators of Neyraudia reynaudiana among different nitrogen stress treatments. The effects of low nitrogen stress on the leaf protection enzymes of Neyraudia reynaudiana were significantly. The activities of POD, SOD and CAT in Neyraudia reynaudiana under normal nitrogen supply, slight, moderate stress were gradually increased, however, with the intensification of nitrogen stress, each protection enzymes activities decreased in turn among severe nitrogen stress and not nitrogen supply treatments. The MDA content and relative conductivity gradually increased with the intensification of nitrogen stress. When the nitrogen stress reaches a certain level, both showed a decreasing trend. The leaf soluble proteins content and secrete acidic substances content in roots of Neyraudia reynaudiana under severe and moderate stress treatments were significantly higher than other treatments (p<0.05).
(10) There was significantly difference in the leaves physiology indicators of Neyraudia reynaudiana among different potassium stress treatments. To some extent, under low potassium stress the protection enzymes of Neyraudia reynaudiana were enhanced, the activities of POD, SOD and CAT inNeyraudia reynaudiana leaves under moderate potassium stress were higher, and there were the lowest under normal supply potassium treatment, which significantly lower than the low potassium stress treatment (p<0.05). The MDA content was the highest under moderate stress among different treatments from the leaf membrane system of Neyraudia reynaudiana. While the leaf relative conductivity showed a gradually increasing trend with the intensification of low potassium stress. Similarly, the MDA content and the leaf relative conductivity showed a gradually increasing and then decreasing trend with the intensification of potassium stress.
Based on the above results, the response strategies to environmental stress for pioneer plant Neyraudia reynaudiana in the collapsing hill area. were as follows:when field moisture capacity less than40%and moderate or severe drought stress, the tillers number of Neyraudia reynaudiana increased, which improved the coverage of Neyraudia reynaudiana to reduce soil moisture loss by increasing the leaves number and leaf width. At the same time, the average intercellular CO2content, chlorophyll, MDA content and relative conductivity significantly increased, which improved water use efficiency to some extent.The MDA content and the relative conductivity showed a gradually increasing and then decreasing trend with the intensification of drought stress, while the root biomass showed a significantly increasing trend, root-shoot ratio increasing, which mainly through the amplification of the root surface area and root length extending outward to enhance the searching ability for moisture in soi; and meet the demand of its normal growth.
It showed a similar response strategy for Neyraudia reynaudiana to phosphorus nitrogen and potassium stress. When phosphorus content was less than0.50mmol·L-1, the nitrogen content was less than3.75mmol·L-1, and the potassium content was less than3.0mmol·L-1in the soil, or under a moderate or severe nutrient stress, the activities of POD, SOD and CAT in Neyraudia reynaudiana leaves were significantly increased, the leaf soluble proteins content and secrete acidic substances content in roots increased. At the same time, stalks, leaves, roots, and biomass accumulation of Neyraudia reynaudiana also increased, the root-shoot ratio, total root length, total root surface area and total root volume of Neyraudia reynaudiana significantly increased, and then improved root searching ability of Neyraudia reynaudiana under the nutrient stress. However, the active response mechanism to nutrient stress only last for a short time. With nutrient stress for a long time, the enzyme activity and number of the leaves might decrease, and the growth of Neyraudia reynaudiana became slow.
有鉴于此,本文根据胁迫植物学的原理和方法,以南方崩岗侵蚀区广泛分布的类芦为研究对象,利用自主设计的环境养分和水分胁迫模拟装置,通过不同的养分和水分胁迫试验,研究养分和水分胁迫条件下类芦的形态学和生理学特征变化,分析类芦在不同环境胁迫条件下形态学和生理学特征的差异,探讨类芦对环境土壤养分和水分的吸收利用策略,挖掘类芦这种特殊生物学特性的生态学适应意义,为揭示类芦适应贫瘠和干旱崩岗侵蚀区土壤的内在机制提供科学依据。主要研究结果如下：
(1)干旱胁迫对类芦地上部分生长有较大影响。干旱胁迫初期不同干旱处理对类芦分蘖和叶片数量影响较大。与充足供水处理相比,中度和轻度干旱胁迫处理类芦分蘖和叶片数量明显较多(p<0.05),但株高生长能力明显减弱。在重度干旱胁迫条件下类芦叶片长度生长明显受阻,但叶片宽度生长较快。随干旱胁迫时间的延长,类芦株高、分蘖能力、叶片数量及最大叶宽生长没有受到明显抑制,但叶片的伸长能力受阻。
(2)干旱胁迫对类芦根系有较大影响,不同处理类芦根系总长度、表面积、平均直径、根系生物量存在明显差异。随干旱胁迫程度的增加,根体积、根平均直径减小；而类芦根系总长度、根表面积、根系生物量、根冠比均比充足供水处理大；在中度干旱胁迫条件下类芦的根系长度、根表面积、根系生物量、地上部生物量、根冠比最大。类芦根系在环境胁迫条件下增强延伸能力、扩大与土壤接触面积可能是其对干旱逆境的形态学响应机制之一。(3)干旱胁迫对类芦光合特性有较大影响。在中度和轻度干旱胁迫条件下类芦叶绿素含量增加,叶绿素a/b比值下降,叶绿素荧光参数F0、Fm、Fv提高,Fv/Fm、Fv/F0下降。类芦叶片的胞间CO2浓度随干旱程度加重而逐渐增加,表现为上午缓慢上升、下午快速上升,16：00达到最大值。净光合速率、气孔导度、蒸腾速率日变化均表现为“单峰型”,为先升后降,12：00达到高峰,日均值随干旱程度加重而逐渐下降。在中度和轻度干旱胁迫下类芦WUE提高,重度干旱处理WUE稍有下降,说明类芦具有较强的耐干旱能力。
(4)不同干旱胁迫处理类芦叶片的生理学指标存在明显差异。干旱胁迫对类芦叶片保护酶活性影响较大,供水充足处理类芦叶片的POD、SOD和CAT活性明显高于轻度胁迫、中度胁迫和重度胁迫处理。从叶片膜系统来看,干旱胁迫越严重,类芦叶片的MDA含量越高,而当水分供应充足时,其MDA含量同样呈升高趋势；叶片相对电导率随胁迫强度的增强逐渐增加。干旱胁迫对类芦叶片可溶性蛋白和芦根系分泌酸性物质的影响表现为随干旱胁迫加剧而有所降低。
(5)不同P素胁迫对类芦生物量分配格局及根系形态指标有明显影响。中度P胁迫处理类芦根系总长度、总表面积和总体积明显大于其它处理,中度P胁迫有利于类芦根长、表面积及体积的增长,但不利于根直径生长。P素胁迫对类芦秆、叶、根和生物积累量的影响均表现出随P胁迫程度的加剧呈上升趋势,然后逐渐下降。
(6)不同N胁迫处理的类芦根系总长度、表面积、根体积和根系生物量存在差异明显。不同N胁迫下根长、根表面、根体积、根系生物量随N胁迫程度增加而逐渐减小。在N素极度匮乏逆境中,类芦生长缓慢,但根系生物量分配比例比地上部分高,这可能是类芦抵抗环境N胁迫的一种形态学机制之一。
(7)不同K胁迫处理类芦生物量分配格局及根系形态指标存在明显差异。不供K处理类芦根系总长度、总表面积和总体积均明显大于其它处理。随供K量的减少,类芦根系平均直径、全株生物量呈递减趋势,根冠比呈增加趋势。
(8)不同P胁迫处理类芦叶片生理学指标存在明显差异。P胁迫一定程度上增强了类芦叶片保护酶活性,与正常供P处理相比,P胁迫处理类芦叶片POD、SOD和CAT活性均明显增强(p<0.05),但随P胁迫程度的加剧,各种酶活性均降低。从叶片膜系统来看,随P胁迫程度加剧,类芦叶片MDA含量和相对电导率逐渐上升,而当P胁迫达到一定程度时,两者呈降低趋势。在中度P胁迫类芦叶片可溶性蛋白和根系分泌酸性物质含量明显高于其它处理。
(9)N胁迫对类芦叶片生理学指标有明显影响。N胁迫对类芦叶片保护酶活性影响明显,正常供N、轻度N胁迫、中度N胁迫处理类芦叶片POD、SOD和CAT活性逐渐增强,但随N胁迫程度的加剧,重度N胁迫和不供N处理保护酶活性依次降低。从叶片膜系统来看,随N胁迫程度的加剧,类芦叶片MDA含量和相对电导率逐渐上升,而当N胁迫达一定程度时,两者均呈降低趋势。重度N胁迫和中度N胁迫处理类芦叶片可溶性蛋白和根系分泌酸性物质含量均明显高于其它处理(p<0.05)。
(10)不同K胁迫处理类芦叶片生理学指标存在明显差异。K胁迫一定程度上可增强类芦叶片保护酶活性,中度K胁迫类芦叶片POD、SOD和CAT活性较高,正常供K处理中各种酶活性最低,明显低于其它K胁迫处理(p<0.05)。从类芦叶片膜系统来看,中度K胁迫处理的MDA含量最高,而叶片相对电导率则随K胁迫程度的加剧呈升高趋势。类芦叶片可溶性蛋白和根系分泌酸性物质含量随K胁迫程度的加剧而表现出先升高后下降趋势。
综合以上研究结果,崩岗侵蚀区先锋植物类芦对环境水分胁迫的响应策略为：当环境土壤田间持水量小于40%形成干旱胁迫时,一方面,芦类的分蘖数量增加,通过增加叶片数量与宽度、减小叶片长度,以减少土壤水分的散失速度。另一方面,干旱胁迫条件下类芦体内叶片胞间CO：浓度、叶绿素、MDA含量和相对电导率显著增加,一定程度上提高了类芦体内的水分利用效率。随干旱胁迫强度的加剧,叶片可溶性蛋白和根系分泌酸性物质含量呈现先增加后降低趋势,但根系生物量呈上升趋势,根冠比不断增加,主要通过根系表面积的扩增以及根系向外延伸,加大根系与土壤的接触面积,增强对环境土壤水分的主动寻觅能力,以维持植株正常生长对水分的需求,这可能是芦类能在极其干旱的环境中正常生长的主要原因。
类芦对P素胁迫、N素胁迫和K素胁迫在一定程度上表现出相似的响应策略,其对环境养分胁迫的响应机制为：当土壤含P量、含N量、含K量分别小于0.50mmol·L-1、3.75mmol·L-1、3.0mmol·L-1形成中度或重度养分胁迫时,类芦叶片POD、SOD和CAT酶活性增强,叶片可溶性蛋白和根系分泌酸性物质含量有所增加,类芦秆、叶、根和全株生物积累量也随之增加,根冠比增大,根系总长度、总表面积和体积明显增加,从而提高了类芦根系在养分胁迫逆境中的主动寻觅养分的能力。但类芦根系的这种主动响应机制只能维持一小段时间,随养分胁迫时间的延长,类芦叶片酶活性及其数量降低,生长变缓,因此类芦对环境贫瘠养分的适应有一定限度。
Neyraudia reynaudiana is a pioneer Plant for water and soil conservation in the southern collapsing hill area. They can recovery rapidly in the bare soil and then become dominance species. Neyraudia reynaudiana can survive and grow in nutrient deficiency and drought soil, which show it may possesses specific mechanisms to find the nutrient and moisture in poor soil. However, the adaption mechanism of Neyraudia reynaudiana to nutrients deficiency and drought condition is still not determined clearly.
For these reasons, the morphological and physiological characteristics of Neyraudia reynaudiana were determined under nutrient and drought stress based on the method of Stress Botany. The differences between the morphological and physiological features were analyzed in different stress treatments. The strategy to find the nutrients and moisture in poor soil was discussed for Neyraudia reynaudiana, which would be benefit for understanding its adaption mechanism to nutrients and drought stress. The main results were as follows:
(1) There was significant effect on the growth of Neyraudia reynaudiana among the different water stress treatments. In the early stage of drought stress, the moisture content in the soil had a greater effect on the number of tillers and leaves of Neyraudia reynaudiana. Compared with adequate water supply treatment, the numbers of tillers and leaf of Neyraudia reynaudiana were significantly more both in moderate and slight water stress treatment, however, the growths of their height were significantly reduced (p<0.05). The growth of leaf length was weaned in the severe drought stress treatment, however, the growth of its leaf width was speeded. AS the drought stress goes, the growth of its height, tiller ability, leaf number and maximum leaf width were inhibited, however, leaf elongation ability weakened significantly.
(2) There was significantly difference in total root length, root surface area, root average diameter, root biomass of Neyraudia reynaudiana among different drought stress treatments, however, not in the root volume. As the drought stress increased, the volume and diameter of roots reduced gradually in different drought stress. Compared with the adequate water treatment, the total length, surface area, biomass of roots, and root-shoot ratio were high significantly under different drought stres. The total length, surface area, biomass of roots, aboveground biomass, root-shoot ratio were the highest under moderate drought stress. Neyraudia reynaudiana had strong drought-tolerant because of large root system with root length and root surface area. The root characters of elongation growth and enlargement root-soil interface of Neyraudia reynaudiana may be one of morphology response mechanisms to drought stress.
(3) The effects of drought stress on photosynthetic characteristics of Neyraudia reynaudiana were significantly different.. Chlorophyll content of Neyraudia reynaudiana increased and chlorophyll a/b was decrease under moderate and slight drought stress. Chlorophyll fluorescence parameters F0, Fm, Fv increased and Fv/Fm, Fv/F0decreased under drought stress. The average intercellular CO2content increased as the drought stress increasing. They increase slowly in the morning and rapidly increase in the afternoon to maximum at16:00. The diurnal variation curves of net photosynthetic rate (Pn), stomatal conductance (Gs), transpiration rate (Tr) were increasing, and then decreasing, a single-peak at12:00. Pn, Gs, and Tr decreased as drought stress increasing. The WUE of Neyraudia reynaudiana increased under moderate and slight drought stress, and decreased slightly under heavy drought stress. Therefore, Neyraudia reynaudiana had strong drought-resistant capacity.
(4) In the different drought stress treatments, there were significantly differences among the physiological process of Neyraudia reynaudiana. The effects of drought stress on protection enzymes of Neyraudia reynaudiana were significant. The activities of POD, SOD and CAT in leaves of Neyraudia reynaudiana under adequate water supply treatment were significantly higher than the slight, moderate and severe stress. From the leaf membrane system, the more serious the drought stress, the higher MDA content of Neyraudia reynaudiana.The MDA content showed the increasing trend under adequate water supply. The relative Conductivity of leaf gradually increased with increasing of drought stress. The effects of drought stress on the leaf soluble proteins and secrete acidic substances in roots of Neyraudia reynaudiana were showed as follows: their contents were decreased with increasing of drought stress.
(5) Among the different low phosphorus stress treatments, there were significant effects on biomass distribution pattern and root morphology indicators of Neyraudia reynaudiana. The total root length, total root surface area and total root volume of Neyraudia reynaudiana were significantly higher than other treatments under moderate stress. The moderate stress was more benefit for growth of root length, root surface area and root volume, however, not conducive to root diameter growth. The effects on stalks, leaves, roots, and biomass accumulation of Neyraudia reynaudiana under the low phosphorus stress showed an upward trend with increase of phosphorus stress and then gradually decline.
(6) Among the different nitrogen stress treatments, there was significantly different in total root length, root surface area, root average diameter, root biomass of Neyraudia reynaudiana, However, not in the average diameter. The root length, root surface area, root volume and root biomass decreased with increase of nitrogen stress. The Neyraudia reynaudiana grew slowly under the extreme shortage of nitrogen in the soil, however, root biomass distribution ratio was higher than the aerial parts which may be an ecological resistance mechanism of Neyraudia reynaudiana under nitrogen stress.
(7) There was significantly difference in biomass allocation pattern and root morphology indicators of Neyraudia reynaudiana under different potassium stress treatments. The total root length, total root surface area and total root volume of Neyraudia reynaudiana without potassium supply treatment were significantly higher than other treatments. The root average diameter of Neyraudia reynaudiana showed a decreasing trend with the decrease of potassium. The whole plant biomass showed an increasing trend and the root-shoot ratio showed a declining trend with the increase of potassium.
(8) There was significantly difference in the leaves physiology indicators of Neyraudia reynaudiana among different phosphorus stress treatments. To some extent, the leaf protection enzymes of Neyraudia reynaudiana were enhanced under low phosphorus stress. Compared with normal phosphorus supply treatment, the activities of POD, SOD and CAT in Neyraudia reynaudiana leaves under low phosphorus stress were significantly increased (p<0.05), however, with the intensification of phosphorus stress, each protection enzymes activities consistently decreased under severe phosphorus stress and not phosphorus supply treatments. From the leaf membrane system, the MDA content and relative conductivity gradually increased with the intensification of phosphorus stress, while the phosphorus stress reaches a certain level, both showed a decreasing trend. The leaf soluble proteins content and secrete acidic substances content in roots of Neyraudia reynaudiana were significantly higher than other treatments under moderate stress.
(9) There was significantly difference in the leaves physiology indicators of Neyraudia reynaudiana among different nitrogen stress treatments. The effects of low nitrogen stress on the leaf protection enzymes of Neyraudia reynaudiana were significantly. The activities of POD, SOD and CAT in Neyraudia reynaudiana under normal nitrogen supply, slight, moderate stress were gradually increased, however, with the intensification of nitrogen stress, each protection enzymes activities decreased in turn among severe nitrogen stress and not nitrogen supply treatments. The MDA content and relative conductivity gradually increased with the intensification of nitrogen stress. When the nitrogen stress reaches a certain level, both showed a decreasing trend. The leaf soluble proteins content and secrete acidic substances content in roots of Neyraudia reynaudiana under severe and moderate stress treatments were significantly higher than other treatments (p<0.05).
(10) There was significantly difference in the leaves physiology indicators of Neyraudia reynaudiana among different potassium stress treatments. To some extent, under low potassium stress the protection enzymes of Neyraudia reynaudiana were enhanced, the activities of POD, SOD and CAT inNeyraudia reynaudiana leaves under moderate potassium stress were higher, and there were the lowest under normal supply potassium treatment, which significantly lower than the low potassium stress treatment (p<0.05). The MDA content was the highest under moderate stress among different treatments from the leaf membrane system of Neyraudia reynaudiana. While the leaf relative conductivity showed a gradually increasing trend with the intensification of low potassium stress. Similarly, the MDA content and the leaf relative conductivity showed a gradually increasing and then decreasing trend with the intensification of potassium stress.
Based on the above results, the response strategies to environmental stress for pioneer plant Neyraudia reynaudiana in the collapsing hill area. were as follows:when field moisture capacity less than40%and moderate or severe drought stress, the tillers number of Neyraudia reynaudiana increased, which improved the coverage of Neyraudia reynaudiana to reduce soil moisture loss by increasing the leaves number and leaf width. At the same time, the average intercellular CO2content, chlorophyll, MDA content and relative conductivity significantly increased, which improved water use efficiency to some extent.The MDA content and the relative conductivity showed a gradually increasing and then decreasing trend with the intensification of drought stress, while the root biomass showed a significantly increasing trend, root-shoot ratio increasing, which mainly through the amplification of the root surface area and root length extending outward to enhance the searching ability for moisture in soi; and meet the demand of its normal growth.
It showed a similar response strategy for Neyraudia reynaudiana to phosphorus nitrogen and potassium stress. When phosphorus content was less than0.50mmol·L-1, the nitrogen content was less than3.75mmol·L-1, and the potassium content was less than3.0mmol·L-1in the soil, or under a moderate or severe nutrient stress, the activities of POD, SOD and CAT in Neyraudia reynaudiana leaves were significantly increased, the leaf soluble proteins content and secrete acidic substances content in roots increased. At the same time, stalks, leaves, roots, and biomass accumulation of Neyraudia reynaudiana also increased, the root-shoot ratio, total root length, total root surface area and total root volume of Neyraudia reynaudiana significantly increased, and then improved root searching ability of Neyraudia reynaudiana under the nutrient stress. However, the active response mechanism to nutrient stress only last for a short time. With nutrient stress for a long time, the enzyme activity and number of the leaves might decrease, and the growth of Neyraudia reynaudiana became slow.
引文
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