重庆石灰岩地区柏木幼苗对水分胁迫的生理生态适应性研究
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摘要
石灰岩地区是三大生态脆弱带之一,该地区岩石裸露率高,土壤瘠薄,保水能力极差,临时性干旱经常发生。能够在石灰岩地区这种干旱环境中生长的植物,它们发展了特殊的适应能力,通过自身器官的保护运动、形态重塑以及代谢过程的调整来适应低水多变的环境,形成特殊的适应机制。利用适生植物自身的抗旱能力来进行石灰岩地区植被的恢复和重建,成为石灰岩生态研究中的重要问题。不同强度或不同类型的水分胁迫很可能导致植物不同程度或方式上的适应性反应。因此,为了全面而深入地了解石灰岩山地适生物种对水分胁迫的适应机制,我们以柏木实生苗为材料,通过盆栽水分受控实验,设定了对照、轻度干旱、中度干旱和重度干旱四个不同的水分胁迫强度,以及胁迫初期、胁迫中期和胁迫末期三个胁迫历时,对其在水分胁迫下的生长特性、光合特性、抗氧化系统、渗透调节等生理特性及复水条件下其修复的可能能力,系统研究一系列适应性反应来揭示柏木的抗旱机制,为石灰岩地区的植被恢复提供理论依据和技术支持。我们得出的主要结果如下:
1.水分胁迫初期,水分胁迫并不能影响柏木幼苗的根长、根系表面积、根系平均直径、根系体积等根系大小参数;但是随着胁迫程度的加剧,包括胁迫历时和胁迫强度的加剧,根系大小参数呈降低趋势。柏木幼苗的苗高和地径都随着胁迫强度的加剧而降低。地上部分的生物量在胁迫初期和胁迫中期并没有受到影响,而胁迫末期干旱处理才与对照有显著差异,但地下部分的生物量随胁迫程度的加重而减低。柏木幼苗的总生物量基本上随着水分胁迫的加剧而降低,但在胁迫初期和中期的轻度干旱下其生物量并没有受到影响。当胁迫强度不够严重时,幼苗的地下部分生物量与地上生物量之比呈降低趋势,但胁迫很严重时,根冠比有增大的趋势。胁迫末期的中度干旱受到水分胁迫影响的根系表面积和根系体积在复水后能够恢复到对照水平,而胁迫中期和胁迫末期的重度干旱复水后,根系所有的形态参数不能够恢复到对照水平。复水后柏木幼苗的苗高没有恢复,而地径有一定程度的恢复。复水促进柏木幼苗根冠和总干物质积累,干旱越重,促进作用越明显,且复水后根冠比升高。
2.柏木幼苗在水分胁迫下净光合速率、蒸腾速率均下降,气孔导度也是随着土壤干旱的发展而降低,但并没有发现胞间CO_2浓度和气孔导度同时降低,相反在胁迫初期和胁迫中期,当气孔导度降低到某一值时,胞间CO_2浓度出现显著的增高,而在胁迫末期维持比较稳定的值。在胁迫初期和胁迫中期的重度干旱以前,随着对照,轻度干旱和中度干旱胁迫程度的加剧,Ls值增大,但在重度干旱下,Ls值降低。而在胁迫末期,与胁迫初期和中期的重度干旱一致,Ls值降低。叶肉导度Pn/Ci在胁迫初期,重度干旱下才开始显著下降,胁迫中期,中度干旱下开始下降,而胁迫末期在轻度干旱下就开始下降。胁迫初期,在重度干旱之前,我们以内在水分利用效率来计算柏木幼苗的水分利用效率。水分利用效率随着水分胁迫梯度而增加。胁迫中期,在重度干旱之前,水分利用效率几乎没有受到水分胁迫的影响,仍然保持着与对照相同水平。而对于胁迫初期和胁迫中期的重度干旱、胁迫末期这几个阶段,我们以瞬时水分利用效率来描述柏木幼苗的水分利用效率,水分利用效率都表现出显著降低的趋势。复水后,光合速率、蒸腾速率、气孔导度和水分利用效率等生理指标都有不同程度的恢复。其中在中度水分胁迫条件前,幼苗各指标的恢复程度较大,出现了补偿效应。而在重度水分胁迫条件下,柏木幼苗的生理功能受损,上述各指标的恢复程度均较小。
3.柏木幼苗的色素含量都在水分胁迫下升高,且升高幅度随着胁迫程度的增加而增加,我们认为与胁迫时间和叶片浓缩有关。复水后叶片扩展生长大大加快,含水量增加,色素含量降低。类胡萝卜素和叶绿素的比率由于水分胁迫而增加。复水后,类胡萝卜素和叶绿素的比率降低。叶绿素a/b并没有随着土壤含水量的变化而发生变化。
4.正常浇水即对照组柏木幼苗的Fv/Fm在0.81-0.82之间,而水分胁迫下柏木幼苗的Fv/Fm显著降低,且胁迫越重,降低幅度越火,同样Fv/F0也表现出相同的趋势。柏木幼苗的光化学猝灭(qP)随着水分胁迫的加剧而呈下降趋势,胁迫越重,影响越严重。NPQ在整个胁迫过程中呈上升趋势,且胁迫越严重,上升的幅度越大。φPSⅡ和ETR也都随着水分胁迫的加剧而呈降低趋势,且胁迫越严重,降幅越大。复水后,柏树幼苗的荧光参数Fv/F0,Fv/Fm,qp,NPQ,φPSⅡ和ETR等在胁迫程度较轻时都能恢复到对照水平,而胁迫严重时,这些指标也能得到恢复,但不能完全恢复到对照水平。
5.柏木幼苗随着水分胁迫的加剧RWC呈下降趋势,且胁迫越严重,下降幅度越大,复水后,胁迫程度较轻时WRC能恢复到对照水平,而重度胁迫下不能完全恢复到对照水平。在土壤干旱胁迫的进程中,柏木幼苗针叶中自由水/束缚水下降,叶片组织的水分由自由水向束缚水转化,束缚水含量升高。而复水后,柏木幼苗立即增加自由水的含量,以此来恢复其正常的生长。随干旱处理时间的延长,柏木幼苗的可溶性蛋白含量呈先增加后降低的趋势。在胁迫处理初期植物体内的不溶性蛋白变为可溶性蛋白以增强渗透调节能力,而在胁迫处理后期干旱胁迫超过植物所能忍耐的阈值,植物体内的合成代谢受阻,蛋白质降解。随着水分胁迫程度的加剧柏木幼苗中MDA含量有升高的趋势,但在胁迫初期的相邻处理间、胁迫中期的轻度干旱和中度干旱之间及其胁迫末期的前三种处理间都没有显著性差异,说明了柏木幼苗对水分胁迫有一个适应过程。而每个处理时期的重度干旱MDA含量都极显著的高于对照,复水后其他处理组的MDA都下降到对照水平,而重度干旱下MDA含量不能下降到对照水平,说明胁迫程度较轻的情况下,植物能够很快的自我修复,而重度干旱下膜结构的受害程度已经超出植物的忍耐限度,不能进行很好的自我修复。在本研究中,随干旱胁迫的加剧,SOD、POD、CAT、ASP几种保护酶均呈现持续上升趋势,复水后,SOD活性均有所下降,但总体上还是高于对照,CAT在重度干旱后复水也未能下降到对照水平,而轻度干旱下CAT含量下降到对照水平。
6.可溶性糖和脯氨酸在干旱逆境下随干旱时间的延长和胁迫程度加大其含量增加,且在复水后基本上都能降低到对照水平,是较为有效的渗透调节物质。K~+含量随着干旱胁迫的增加而显著下降。地上部分在胁迫初期,胁迫中期的轻度干旱和中度干旱Na~+有积累,而胁迫中期的重度干旱及胁迫末期Na~+降低,说明Na~+在干旱较轻时起着渗透调节的作用,而重度干旱后Na~+对其渗透调节不起主要作用。Ca~(2+)、Mg~(2+)含量在柏木幼苗内没有明显的积累现象。脯氨酸、可溶性糖起主要渗透调节作用的物质含量在地下部分即根系中的含量大于地上部分。而Na~+在茎叶中的含量大于根系。
实验表明,柏木幼苗能够通过光合生理、保护酶活性和渗透调节等途径耐受一定程度的水分胁迫,且在水分胁迫较轻时进行补偿生长,能够适应石灰岩山地“临时性缺水”的变水环境。
Karst landscape,one of the three vulnerable ecotones,is very general in the word, especially in the southwest of China.It is often short of water for plants due to low water retention capacity and high exposure of rocks.Therefore,it becomes to be a very important issue to find some drought-resistant plants for vegetation restoration and reconstruction in limestone area.In order to adapt to the drought condition in rocky desert terrain,plants have developed a series of mechanisms,such as organic protective movement,morphology remodeling and metabolic regulation.The plants maybe will have different adaption ways because of different stress strengths or different extents of water stress.In order to understand the suitable trees in limestone area how to resist water stress wholly and completely,we selected Cupressus funebris Endl.as our material,studied the growth characteristics,photosynthetic characteristics,antioxidant system and osmotic adjustment as well as the repair ability after re-watering by water controlled experiment.The experiment was set four water levels,that is,CK,light water stress(LS),middle water stress(MS) and severe water stress(SS) and,three stress stages, that is,early stage,middle stage and end stage.The main results are as following:
1.Water stress had no effects on the length,surface area,average diameter and volume of roots of Cupressusfunebris Endl.seedlings at the early stage.But with the aggravation of stress,they all decreased.The heights and diameters of Cupressus funebris Endl.seedlings dropped with the decline of water content in soil.The above-ground biomasses did not suffer from water stress at the early and middle stage, however,they had significant differences with CK.The under-ground biomasses decreased with the aggravation of stress.For the total biomasses,they also decreased at all three stages,but they had no difference between LS and CK at early and middle stages.The root and shoots ratio of Cupressusfunebris Endl.seedlings had the trend of decrease at the early and middle stages,but no differences at the end stage.When re-watering,the root surface area and root volume that had decreased when suffered MS at the end stage recovered to the level of CK,and they could not come back to the level of CK when suffered SS at the middle and end stages.The height of Cupressus funebris Endl.seedlings did not recover,while the diameter recovered at certain degree. Re-watering promoted the accumulation of above-ground biomass,under-ground biomass and total biomass.This effect was more obvious with lowering of water content in soil.
2.The net photosynthetic rate(Pn),transpiration rate(E) and stomatal conductance (Gs)of Cupressus funebris Endl.seedlings were all decreased under water stress. However,Intercellular CO_2 concentration(Ci) did not decrease with Gs.It increased remarkably at the early and middle stages when Gs decreased to a certain value,and kept steady at the end stage.Stomatal limitation value(Ls) went up with the CK,LS and MS at the early and middle stages,while went down under SS.But at the end stage,it went down from CK to SS.Mesophyll conductance(Pn/Ci) fell remarkably under SS at the early stage.But it fell under MS at the middle stage,and under LS at the end stage. Before SS at the early and middle stage,we calculated water use efficiency by intrinsic WUE,which increased or kept unchangeable with the lessening of water content in soil. While instantaneous water use efficiency was used to SS at early and middle stages and all the treatments at end stage,it behaved decreased.After re-watering,Pn,E,Gs and WUE recovered with different degrees.Compensation effect was visible before serious stress when the photosynthesis indexes recovered greatly,while they recovered little under serious stress due to the hurt of physiological function of Cupressusfunebris Endl. seedlings.
3.Photosynthetic pigment content and the extent increased with the aggravation of water stress,which correlated with the stress time and leaf concentration.After re-watering,the pigment decreased owing to leaf expansion growth and higher water content.Car/Chl went up at water stress while went down after re-water.Chl.a/Chl.b had no change under any stress.
4.Fv/Fm of well-water Cupressusfunebris Endl.seedlings(CK) was 0.81-0.82, nevertheless,it decreased notably under water stress and the extent increased with the aggravation of water stress.Similarly,Fv/F_0 had the same trend,qP decreased,while NPQ increased with the CK,LS,MS and SS.The extent greatened with the aggravation of water stress.φPSⅡand ETR had the same trend with Fv/Fm and Fv/F_0.After re-watering,all the chlorophyll fluorescence indexes came back to the level of CK when stress was not very severe,whereas they could not recovered to the level of CK when stress was severe.
5.Relative water content(RWC) in leaves of Cupressusfunebris Endl.seedlings had the trend of decline and the extent greatened with the aggravation of water stress. After re-watering,RWC could come back to the same as CK when the stress was not very serious but it did not when the stress was too serious.During the period of stress, free water transformed to bound water so that the ratio of free water and bound water decreased.On the contrary,the bound water transformed to free water and so the ratio of free water and bound water increased.Soluble protein of the four treatments increased first and decreased afterwards as treatment was prolonged.MDA of the four treatments consistently increased as treatment was prolonged.But there was no significant difference between the neighboring treatments of the early and end stage, MS and LS of the middle stage.However,MDA of SS at any stage was higher than that of CK,and could not recovere to that of CK when re-watering.As for protective enzyme,SOD,POD,CAT,ASP,they went up continuously with the water content lowering in the soil.Re-watering caused SOD go down,but still higher than that of CK; POD,CAT,ASP come back to that of CK under LS,but not under serious stress.
6.Soluble sugar and proline(Pro) increased consistently with the treatment prolonged and the stress aggravated.After re-watering they could decreased to that of CK.It means that they were good osmotic regulation matter.K~+ declined obviously under water stress.But for Na~+,it increased in the shoots of Cupressusfunebris Endl.at early stage,and also increased under LS and MS at middle stage,nevertheless,it decreased under SS at middle stage and all treatments of end stage.Ca~(2+)、Mg~(2+) did not accumulate not only in the shoots but also in the roots of Cupressusfunebris Endl..The main osmotic regulation matter,soluble sugar and Pro were higher in the roots than that of in the shoots,while Na~+ was opposite.
Experiment showed that,Cupressus funebris Endl.could tolerate a certain extent water stress and adapt the changeable environment of "temporary water deficit" by the ways of adjusting photosynthetic physiology,protective enzyme active,osmotic regulation and its good compensatory growth ability.
1.水分胁迫初期,水分胁迫并不能影响柏木幼苗的根长、根系表面积、根系平均直径、根系体积等根系大小参数;但是随着胁迫程度的加剧,包括胁迫历时和胁迫强度的加剧,根系大小参数呈降低趋势。柏木幼苗的苗高和地径都随着胁迫强度的加剧而降低。地上部分的生物量在胁迫初期和胁迫中期并没有受到影响,而胁迫末期干旱处理才与对照有显著差异,但地下部分的生物量随胁迫程度的加重而减低。柏木幼苗的总生物量基本上随着水分胁迫的加剧而降低,但在胁迫初期和中期的轻度干旱下其生物量并没有受到影响。当胁迫强度不够严重时,幼苗的地下部分生物量与地上生物量之比呈降低趋势,但胁迫很严重时,根冠比有增大的趋势。胁迫末期的中度干旱受到水分胁迫影响的根系表面积和根系体积在复水后能够恢复到对照水平,而胁迫中期和胁迫末期的重度干旱复水后,根系所有的形态参数不能够恢复到对照水平。复水后柏木幼苗的苗高没有恢复,而地径有一定程度的恢复。复水促进柏木幼苗根冠和总干物质积累,干旱越重,促进作用越明显,且复水后根冠比升高。
2.柏木幼苗在水分胁迫下净光合速率、蒸腾速率均下降,气孔导度也是随着土壤干旱的发展而降低,但并没有发现胞间CO_2浓度和气孔导度同时降低,相反在胁迫初期和胁迫中期,当气孔导度降低到某一值时,胞间CO_2浓度出现显著的增高,而在胁迫末期维持比较稳定的值。在胁迫初期和胁迫中期的重度干旱以前,随着对照,轻度干旱和中度干旱胁迫程度的加剧,Ls值增大,但在重度干旱下,Ls值降低。而在胁迫末期,与胁迫初期和中期的重度干旱一致,Ls值降低。叶肉导度Pn/Ci在胁迫初期,重度干旱下才开始显著下降,胁迫中期,中度干旱下开始下降,而胁迫末期在轻度干旱下就开始下降。胁迫初期,在重度干旱之前,我们以内在水分利用效率来计算柏木幼苗的水分利用效率。水分利用效率随着水分胁迫梯度而增加。胁迫中期,在重度干旱之前,水分利用效率几乎没有受到水分胁迫的影响,仍然保持着与对照相同水平。而对于胁迫初期和胁迫中期的重度干旱、胁迫末期这几个阶段,我们以瞬时水分利用效率来描述柏木幼苗的水分利用效率,水分利用效率都表现出显著降低的趋势。复水后,光合速率、蒸腾速率、气孔导度和水分利用效率等生理指标都有不同程度的恢复。其中在中度水分胁迫条件前,幼苗各指标的恢复程度较大,出现了补偿效应。而在重度水分胁迫条件下,柏木幼苗的生理功能受损,上述各指标的恢复程度均较小。
3.柏木幼苗的色素含量都在水分胁迫下升高,且升高幅度随着胁迫程度的增加而增加,我们认为与胁迫时间和叶片浓缩有关。复水后叶片扩展生长大大加快,含水量增加,色素含量降低。类胡萝卜素和叶绿素的比率由于水分胁迫而增加。复水后,类胡萝卜素和叶绿素的比率降低。叶绿素a/b并没有随着土壤含水量的变化而发生变化。
4.正常浇水即对照组柏木幼苗的Fv/Fm在0.81-0.82之间,而水分胁迫下柏木幼苗的Fv/Fm显著降低,且胁迫越重,降低幅度越火,同样Fv/F0也表现出相同的趋势。柏木幼苗的光化学猝灭(qP)随着水分胁迫的加剧而呈下降趋势,胁迫越重,影响越严重。NPQ在整个胁迫过程中呈上升趋势,且胁迫越严重,上升的幅度越大。φPSⅡ和ETR也都随着水分胁迫的加剧而呈降低趋势,且胁迫越严重,降幅越大。复水后,柏树幼苗的荧光参数Fv/F0,Fv/Fm,qp,NPQ,φPSⅡ和ETR等在胁迫程度较轻时都能恢复到对照水平,而胁迫严重时,这些指标也能得到恢复,但不能完全恢复到对照水平。
5.柏木幼苗随着水分胁迫的加剧RWC呈下降趋势,且胁迫越严重,下降幅度越大,复水后,胁迫程度较轻时WRC能恢复到对照水平,而重度胁迫下不能完全恢复到对照水平。在土壤干旱胁迫的进程中,柏木幼苗针叶中自由水/束缚水下降,叶片组织的水分由自由水向束缚水转化,束缚水含量升高。而复水后,柏木幼苗立即增加自由水的含量,以此来恢复其正常的生长。随干旱处理时间的延长,柏木幼苗的可溶性蛋白含量呈先增加后降低的趋势。在胁迫处理初期植物体内的不溶性蛋白变为可溶性蛋白以增强渗透调节能力,而在胁迫处理后期干旱胁迫超过植物所能忍耐的阈值,植物体内的合成代谢受阻,蛋白质降解。随着水分胁迫程度的加剧柏木幼苗中MDA含量有升高的趋势,但在胁迫初期的相邻处理间、胁迫中期的轻度干旱和中度干旱之间及其胁迫末期的前三种处理间都没有显著性差异,说明了柏木幼苗对水分胁迫有一个适应过程。而每个处理时期的重度干旱MDA含量都极显著的高于对照,复水后其他处理组的MDA都下降到对照水平,而重度干旱下MDA含量不能下降到对照水平,说明胁迫程度较轻的情况下,植物能够很快的自我修复,而重度干旱下膜结构的受害程度已经超出植物的忍耐限度,不能进行很好的自我修复。在本研究中,随干旱胁迫的加剧,SOD、POD、CAT、ASP几种保护酶均呈现持续上升趋势,复水后,SOD活性均有所下降,但总体上还是高于对照,CAT在重度干旱后复水也未能下降到对照水平,而轻度干旱下CAT含量下降到对照水平。
6.可溶性糖和脯氨酸在干旱逆境下随干旱时间的延长和胁迫程度加大其含量增加,且在复水后基本上都能降低到对照水平,是较为有效的渗透调节物质。K~+含量随着干旱胁迫的增加而显著下降。地上部分在胁迫初期,胁迫中期的轻度干旱和中度干旱Na~+有积累,而胁迫中期的重度干旱及胁迫末期Na~+降低,说明Na~+在干旱较轻时起着渗透调节的作用,而重度干旱后Na~+对其渗透调节不起主要作用。Ca~(2+)、Mg~(2+)含量在柏木幼苗内没有明显的积累现象。脯氨酸、可溶性糖起主要渗透调节作用的物质含量在地下部分即根系中的含量大于地上部分。而Na~+在茎叶中的含量大于根系。
实验表明,柏木幼苗能够通过光合生理、保护酶活性和渗透调节等途径耐受一定程度的水分胁迫,且在水分胁迫较轻时进行补偿生长,能够适应石灰岩山地“临时性缺水”的变水环境。
Karst landscape,one of the three vulnerable ecotones,is very general in the word, especially in the southwest of China.It is often short of water for plants due to low water retention capacity and high exposure of rocks.Therefore,it becomes to be a very important issue to find some drought-resistant plants for vegetation restoration and reconstruction in limestone area.In order to adapt to the drought condition in rocky desert terrain,plants have developed a series of mechanisms,such as organic protective movement,morphology remodeling and metabolic regulation.The plants maybe will have different adaption ways because of different stress strengths or different extents of water stress.In order to understand the suitable trees in limestone area how to resist water stress wholly and completely,we selected Cupressus funebris Endl.as our material,studied the growth characteristics,photosynthetic characteristics,antioxidant system and osmotic adjustment as well as the repair ability after re-watering by water controlled experiment.The experiment was set four water levels,that is,CK,light water stress(LS),middle water stress(MS) and severe water stress(SS) and,three stress stages, that is,early stage,middle stage and end stage.The main results are as following:
1.Water stress had no effects on the length,surface area,average diameter and volume of roots of Cupressusfunebris Endl.seedlings at the early stage.But with the aggravation of stress,they all decreased.The heights and diameters of Cupressus funebris Endl.seedlings dropped with the decline of water content in soil.The above-ground biomasses did not suffer from water stress at the early and middle stage, however,they had significant differences with CK.The under-ground biomasses decreased with the aggravation of stress.For the total biomasses,they also decreased at all three stages,but they had no difference between LS and CK at early and middle stages.The root and shoots ratio of Cupressusfunebris Endl.seedlings had the trend of decrease at the early and middle stages,but no differences at the end stage.When re-watering,the root surface area and root volume that had decreased when suffered MS at the end stage recovered to the level of CK,and they could not come back to the level of CK when suffered SS at the middle and end stages.The height of Cupressus funebris Endl.seedlings did not recover,while the diameter recovered at certain degree. Re-watering promoted the accumulation of above-ground biomass,under-ground biomass and total biomass.This effect was more obvious with lowering of water content in soil.
2.The net photosynthetic rate(Pn),transpiration rate(E) and stomatal conductance (Gs)of Cupressus funebris Endl.seedlings were all decreased under water stress. However,Intercellular CO_2 concentration(Ci) did not decrease with Gs.It increased remarkably at the early and middle stages when Gs decreased to a certain value,and kept steady at the end stage.Stomatal limitation value(Ls) went up with the CK,LS and MS at the early and middle stages,while went down under SS.But at the end stage,it went down from CK to SS.Mesophyll conductance(Pn/Ci) fell remarkably under SS at the early stage.But it fell under MS at the middle stage,and under LS at the end stage. Before SS at the early and middle stage,we calculated water use efficiency by intrinsic WUE,which increased or kept unchangeable with the lessening of water content in soil. While instantaneous water use efficiency was used to SS at early and middle stages and all the treatments at end stage,it behaved decreased.After re-watering,Pn,E,Gs and WUE recovered with different degrees.Compensation effect was visible before serious stress when the photosynthesis indexes recovered greatly,while they recovered little under serious stress due to the hurt of physiological function of Cupressusfunebris Endl. seedlings.
3.Photosynthetic pigment content and the extent increased with the aggravation of water stress,which correlated with the stress time and leaf concentration.After re-watering,the pigment decreased owing to leaf expansion growth and higher water content.Car/Chl went up at water stress while went down after re-water.Chl.a/Chl.b had no change under any stress.
4.Fv/Fm of well-water Cupressusfunebris Endl.seedlings(CK) was 0.81-0.82, nevertheless,it decreased notably under water stress and the extent increased with the aggravation of water stress.Similarly,Fv/F_0 had the same trend,qP decreased,while NPQ increased with the CK,LS,MS and SS.The extent greatened with the aggravation of water stress.φPSⅡand ETR had the same trend with Fv/Fm and Fv/F_0.After re-watering,all the chlorophyll fluorescence indexes came back to the level of CK when stress was not very severe,whereas they could not recovered to the level of CK when stress was severe.
5.Relative water content(RWC) in leaves of Cupressusfunebris Endl.seedlings had the trend of decline and the extent greatened with the aggravation of water stress. After re-watering,RWC could come back to the same as CK when the stress was not very serious but it did not when the stress was too serious.During the period of stress, free water transformed to bound water so that the ratio of free water and bound water decreased.On the contrary,the bound water transformed to free water and so the ratio of free water and bound water increased.Soluble protein of the four treatments increased first and decreased afterwards as treatment was prolonged.MDA of the four treatments consistently increased as treatment was prolonged.But there was no significant difference between the neighboring treatments of the early and end stage, MS and LS of the middle stage.However,MDA of SS at any stage was higher than that of CK,and could not recovere to that of CK when re-watering.As for protective enzyme,SOD,POD,CAT,ASP,they went up continuously with the water content lowering in the soil.Re-watering caused SOD go down,but still higher than that of CK; POD,CAT,ASP come back to that of CK under LS,but not under serious stress.
6.Soluble sugar and proline(Pro) increased consistently with the treatment prolonged and the stress aggravated.After re-watering they could decreased to that of CK.It means that they were good osmotic regulation matter.K~+ declined obviously under water stress.But for Na~+,it increased in the shoots of Cupressusfunebris Endl.at early stage,and also increased under LS and MS at middle stage,nevertheless,it decreased under SS at middle stage and all treatments of end stage.Ca~(2+)、Mg~(2+) did not accumulate not only in the shoots but also in the roots of Cupressusfunebris Endl..The main osmotic regulation matter,soluble sugar and Pro were higher in the roots than that of in the shoots,while Na~+ was opposite.
Experiment showed that,Cupressus funebris Endl.could tolerate a certain extent water stress and adapt the changeable environment of "temporary water deficit" by the ways of adjusting photosynthetic physiology,protective enzyme active,osmotic regulation and its good compensatory growth ability.
引文
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