239 resultados para root: shoot ratio
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Hybrid opto-digital joint transform correlator (HODJTC) is effective for image motion measurement, but it is different from the traditional joint transform correlator because it only has one optical transform and the joint power spectrum is directly input into a digital processing unit to compute the image shift. The local cross-correlation image can be directly obtained by adopting a local Fourier transform operator. After the pixel-level location of cross-correlation peak is initially obtained, the up-sampling technique is introduced to relocate the peak in even higher accuracy. With signal-to-noise ratio >= 20 dB, up-sampling factor k >= 10 and the maximum image shift <= 60 pixels, the root-mean-square error of motion measurement accuracy can be controlled below 0.05 pixels.
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The physics-based parameter: load/unload response ratio (LURR) was proposed to measure the proximity of a strong earthquake, which achieved good results in earthquake prediction. As LURR can be used to describe the damage degree of the focal media qualitatively, there must be a relationship between LURR and damage variable (D) which describes damaged materials quantitatively in damage mechanics. Hence, based on damage mechanics and LURR theory, taking Weibull distribution as the probability distribution function, the relationship between LURR and D is set up and analyzed. This relationship directs LURR applied in damage analysis of materials quantitatively from being qualitative earlier, which not only provides the LURR method with a more solid basis in physics, but may also give a new approach to the damage evaluation of big scale structures and prediction of engineering catastrophic failure. Copyright (c) 2009 John Wiley & Sons, Ltd.
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The physics-based parameter: load/unload response ratio (LURR) was proposed to measure the proximity of a strong earthquake, which achieved good results in earthquake prediction. As LURR can be used to describe the damage degree of the focal media qualitatively, there must be a relationship between LURR and damage variable (D) which describes damaged materials quantitatively in damage mechanics. Hence, based on damage mechanics and LURR theory, taking Weibull distribution as the probability distribution function, the relationship between LURR and D is set up and analyzed. This relationship directs LURR applied in damage analysis of materials quantitatively from being qualitative earlier, which not only provides the LURR method with a more solid basis in physics, but may also give a new approach to the damage evaluation of big scale structures and prediction of engineering catastrophic failure. Copyright (c) 2009 John Wiley & Sons, Ltd.
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In present study, the transition of thermocapillary convection from the axisymmetric stationary flow to oscillatory flow in liquid bridges of 5cst silicon oil (aspect ratio 1.0 and 1.6) is investigated in microgravity conditions by the linear instability analysis. The corresponding marginal instability boundary is closely related to the gas/liquid configuration of the liquid bridge noted as volume ratio. With the increasing volume ratio, the marginal instability boundary consists of the increasing branch and the decreasing branch. A gap region exists between the branches where the critical Marangoni number of the corresponding axisymmetric stationary flow increases drastically. Particularly, a unique axisymmetric oscillatory flow (the critical azimuthal wave number is m=0) in the gap region is reported for the liquid bridge of aspect ratio 1.6. Moreover, the energy transfer between the basic state and the disturbance fields of the thermocapillary convection is analyzed at the corresponding critical Marangoni number, which reveals different major sources of the energy transfer for the development of the disturbances in regimes of the increasing branch, the gap region and the decreasing branch, respectively.
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当前大气CO2浓度升高是全球变化的主要趋势之一,CO2浓度升高还会引起全球变暖等其它环境问题,因而CO2浓度浓度升高对植物影响的研究已经成为全球变化领域的焦点。红桦是川西亚高山地区暗针叶林演替初期的先锋树种和演替后期的建群种,在群落演替过程中它对环境因子的响应决定红桦群落的演替进程。本文通过控制CO2浓度的气候室试验,研究了CO2浓度倍增环境下,不同密度水平红桦碳氮固定、分配可能发生的改变,并探讨了升高大气CO2浓度对群体内部竞争的影响。以期通过本研究明确川西亚高山地区代表性物种红桦对未来气候变化的响应,为今后采取措施应对气候变化、妥善进行森林管理提供理论依据和科学指导。主要研究结果如下: 1.升高CO2浓度对红桦幼苗生长的影响以及树皮、树干响应的不同 (1) CO2浓度升高显著促进红桦幼苗的生物量、株高、基茎的生长,同时也改变生物量在体内的分配格局,主要是增加根和主茎、减少叶在总生物量中的比重。(2)树皮和树干对升高CO2浓度的影响有差异,它们对CO2浓度升高的反应程度不同,但反应方向一致。 2.密度的副效应 (1) 增加种植密度对单株生物量、株高和基径的生长具有副效应,也降低升高CO2浓度对红桦生长的正效应。(2) 增加种植密度,显著增加红桦幼苗的群体生物量,从而使红桦群体固定更多的大气CO2气体。可见密度在决定红桦生物量及固碳能力方面具有重要意义。探索适合未来大气CO2浓度升高条件下植物生长的密度,对未来的森林经济生产、生态恢复具有重要意义。 3. 升高CO2浓度对红桦幼苗苗冠结构及冠层内部竞争的影响 (1) 冠幅、冠高、苗冠表面积和苗冠体积等树冠特征均受CO2浓度升高的影响而增加,但是受密度增加的影响而降低。(2) 单位苗冠投影面积叶片数(LDcpa)和单位苗冠体积叶片数(LDcv)均低于相应的现行CO2浓度处理,这主要是由于冠幅和冠高的快速生长所造成的。(3) LDcpa和LDcv的降低表明,红桦在升高CO2浓度的条件下,会作出积极的响应,从而缓解由于群体和个体生长的增加所引起的竞争压力的增加。 4. 升高CO2浓度对红桦幼苗养分元素吸收与分配的影响 (1) CO2浓度升高,植株各器官N、P含量降低,但单株N、P总吸收量均增加。红桦幼苗体内N、P浓度的下降是由于生物量迅速增加引起的稀释效应造成的。(2) CO2浓度升高,N、P向主茎和根的分配增加,向叶片的分配减少,主要是由于前者在总生物量中的比重增加,而后者减少了。(3) CO2浓度升高,氮磷利用效率(NUE和PUE)提高,氮磷累积速率(NAcR和PAcR)显著增加。而NUE和PUE的提高可以有效缓解CO2浓度升高后,亚高山和高山地区森林土壤中养分元素不足对森林生产力的限制。 5. 升高CO2浓度对红桦幼苗群体碳平衡的影响 (1) 升高CO2浓度对植物的光合作用、呼吸速率和生长均具有促进作用。(2) 土壤有机碳含量在实验前期迅速增加,后期积累速率下降。(3) 升高CO2浓度以后,土壤呼吸显著增强;土壤呼吸还具有明显的季节变化。(4) 红桦群体日固碳量受到升高CO2浓度的促进作用。结果(1)-(4)说明所研究群落的碳动态对现行的气候波动是敏感的;所研究群落在作为大气CO2气体的源-汇关系方面至少存在季节间的源汇飘移。(5)种植密度的升高显著增加了群体固碳量。 6. 升高CO2浓度对红桦幼苗生长后期叶片衰老的影响 升高CO2浓度有利于减缓红桦幼苗叶片生长季节末期的衰老。生长季节末期,随着CO2浓度的升高光合速率和可溶性蛋白含量均呈上升趋势,同时MDA(丙二醛)含量下降,保护酶SOD(超氧化物岐化酶)、CAT(过氧化氢酶)活性升高。由此说明,升高CO2浓度有利于减缓生长季节后期叶片的衰老,使叶片维持较高的光合速率,也从生理学的角度支持了本文及前人有关CO2浓度升高促进植物光合和生长的假说及结果。 The increased CO2 concentration is one of the most important problems among global changes. The increase of CO2 will also cause other environmental problems, such as global warming, etc. So the effects of elevated CO2 on plant have drawn sights of many scientists in the research field of global change. Red birch (Betula albosinensis) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of the dark coniferous forests in Western Sichuan, China. It’s response to elevated CO2 may determine the succession process of the community where it lives in. By controlling CO2 at the ambient and twice as the ambient level (ambient + 350 umol mol-1) using enclosed-top chambers (ETC), possible effects of elevated CO2 on carbon fixation and allocation under two plantation densities are investigated. The effects of elevated CO2 on competition within canopy of red birch seedlings are also observed in the present paper. We hope to make sure of the effects of elevated CO2 on the representative species, red birch. And so that, our results could provide a strong theoretical evidence and scientific direction for forest management and afforestation under a future, CO2 elevated world. The results are as fowllows: 1. The effects of elevated CO2 on growth and the different responses of wood and bark of red birch seedlings (1) Elevated CO2 increases the growth of seedling biomass, seedling height and basal diameter of red birch. It also changed the biomass allocation in red birch seedlings. The ratio of root and main stem to all biomass is increased and the ratio of leaf is decreased. (2) Tree bark and wood show different response degree but similar response direction to elevated CO2. 2. Negative effects of planting density (1) The increase of planting density showes negative effects on the individual growth of seedling biomass, seedling height and basal diameter of red birch. It also eliminates the positive effects of elevated CO2 on growth of red birch seedlings. (2) Community biomass is increased by the elevated planting density, which means that the high density red birch community could fix more CO2 than the low density one. These results show that planting density plays an important role in determining biomass and carbon fixation ability of red birch community. Thus, exploring proper planting density becomes economically important for the future, CO2 elevated word. 3. The effects of elevated CO2 on crown architecture and competition within canopy of red birch seedlings (1) Crown width, crown depth, crown surface area and crown volume are all increased under the influence of elevated CO2. (2) Leaf number per unit area of projected crown area (LDcpa) and per unit volume of crown volume (LDcv) are lower under elevated CO2. This is resulted from the stimulated growth of tree crown features. (3) The decrease of LDcpa and LDcv indicate that plants will respond forwardly to reduce the possible increase of competition resulted from stimulated growth of individual plant and collectives in conditions of elevated CO2. 4. The effects of elevated CO2 on nutrition accumulation and allocation of red birch seedlings (1) Contents of N and P decrease due to the prompt increase of biomass of plant organs caused by elevated CO2. However, their accumulations increase under elevated CO2. (2) Elevated CO2 increases the allocation of N, P to main stem but reduced its allocation to leaf for that dry weight of the former increased but the dry weight of the later decreased. (3) Using efficiencies of N, P (NUE and PUE) and their accumulation rates (NAcR and PAcR) are found to increase under elevated CO2. Soil nutrition contents are always the limiting factors for plant growth at subalpine and alpine region. The increased NUE and PUE are helpful to eliminate the nutrition limitation in this area in the future world, when CO2 concentration doubles the ambient. 5. The effects of elevated CO2 on carbon balance of red birch communities (1) Net photosynthetic rates (Pn), dark respiration rates (Rd) and growth are all stimulated by elevated CO2. (2) Content soil organic carbon increases sharply at the primary stage of experiments and then the increasing rates decrease to a low level at later stages. (3) Soil respiration rates increase significantly with the elevation of CO2 concentration. (4) The daily carbon fixations of whole community are heightened by elevated CO2. The results (1)-(4) suggest that, the community being studied are sensitive to current climate change; the studied community, as a sink of atmospheric CO2, is pool-sink alternative between seasons. (5) The carbon fixations are increased along the increase of planting densities. 6. The effects of elevated CO2 on physiological features of leaf senescences of red birch seedlings at the later stage of growing season Elevated CO2 helps to postpone the leaf senescences of red birch at the end of the growth season. CO2 enrichment increases the photosynthetic rates, contents of soluble proteins and photosynthetic pigments. And meanwhile contents of malondialdehyde (MDA) decreases and activities of superoxide dismutase (SOD) and catalase (CAT) are both increased. These results suggest that the senescences of red birch leaves are delayed by elevated CO2, which keep the photosynthetic rates at relatively high levels. Our results lend supports to hypothesis and results on stimulated photosynthetic rates and growth from both other researchers and the present paper.
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土壤是人类赖以生存的自然环境和农业生产的重要资源,世界面临的粮食、资源和环境问题与土壤密切相关,目前危害土壤的主要因素是干旱和重金属污染。杨树具有适应性强、生长快和丰产等特性,本论文以青杨组杨树为模式植物,采用植物生态、生理及生物化学等方法,研究杨树对土壤干旱和锰胁迫的生态生理反应以及种群间差异,研究成果可为我国干旱半干旱地区营造人工林、防止沙漠化提供理论依据,也为恢复与重建重金属污染地区退化生态系统提供科学指导。主要研究结果如下: 1. 青海杨不同种群对干旱胁迫的响应差异 干旱胁迫显著降低了两个青海杨种群(干旱种群和湿润种群)生物量积累,包括株高、基径、干物质积累等,通过植物结构的调整,有更多的生物量向根部分配。干旱胁迫还显著降低了叶绿素和类胡萝卜素含量,增加了游离脯氨酸和总氨基酸含量。另一方面,干旱胁迫诱导了活性氧的积累,作为第二信使,激活了抗氧化系统,包括抗坏血酸(ASA)含量和酶系统如超氧化物歧化酶(SOD),愈创木酚过氧化物酶(GPX),抗坏血酸过氧化物酶(APX)和谷胱甘肽还原酶(GR)。这样,杨树既有避旱机制又有耐旱机制,使其在干旱胁迫下有相当程度的可塑性。与湿润种群相比,干旱种群杨树有更多的生物量分配到根部,积累了更多的游离脯氨酸和总氨基酸来进行渗透调节,并且有更有效的抗氧化系统,包括更高含量的ASA 和更高活性的APX 和GR,这些使得干旱种群杨树比湿润种群杨树对干旱有更好的耐性。 2. 喷施硝普钠(SNP)对青海杨阿坝种群干旱胁迫耐性的影响 干旱胁迫显著的降低了青海杨阿坝种群的生长和生物量积累以及叶片相对含水量,还诱导了脯氨酸的合成以进行渗透调节。干旱胁迫下过氧化氢(H2O2)显著累积从而造成对膜脂和蛋白的伤害,使得丙二醛和蛋白羰基含量升高。干旱胁迫下喷施SNP可以减轻干旱胁迫造成的伤害,包括增加叶片的相对含水量,增加脯氨酸和总氨基酸的积累,并激活抗氧化酶系统如SOD,GPX和APX,从而减少丙二醛(MDA)和蛋白羰基(C=O)的积累,但是在水分良好情况下SNP的效果不显著。 3. 青杨不同种群对锰胁迫的生长与形态响应差异 在同一锰浓度下,干旱种群的耐性指数都要高于湿润种群,这表明青杨对干旱和高锰胁迫具有交叉耐性。两个种群的株高,生物量和叶绿素含量都随锰浓度的升高而逐渐下降。就累积浓度而言,0 和0.1 mM 锰胁迫下,干旱种群积累的锰浓度要高于湿润种群,而在高浓度锰胁迫下(0.5 和1 mM),湿润种群要高于干旱种群。在0,0.1 和0.5 mM下,锰大多积累在根中,叶片次之,茎中最少。而在1 mM,锰更多的积累在叶片中。就累积总量而言,在各个锰浓度胁迫下,根,茎和叶相比,两个种群青杨都是叶片累积的锰总量要高于根和茎。两个种群间比较,对照中没有显著区别,0.1 mM 锰胁迫下,湿润种群根中累积的锰要高于干旱种群,而在地上部中,干旱种群要高于湿润种群。而0.5 和1 mM 锰胁迫下,根、叶、茎+叶、根+茎+叶中,锰累积总量都是湿润种群高于干旱种群。锰胁迫下,青杨叶片数和叶面积包括总叶面积和平均叶面积都显著降低。叶片横切面的光学显微观察结果表明未经锰胁迫的栅栏组织的细胞饱满,海绵组织发达、清晰;胁迫后杨树叶片栅栏组织细胞出现不同程度的皱缩,海绵组织几乎不可见,此外还发现输导组织在胁迫下密度变小和分生组织严重割裂等现象。 4. 青杨不同种群对锰胁迫的生理与生化响应差异 青杨两个种群脱落酸(ABA)含量在锰胁迫下都显著增加,干旱种群的增幅更大。三种多胺含量在锰胁迫下显示了不同的响应趋势:腐胺在两个种群的各个锰处理下都增加,亚精胺只在干旱种群中显著增加,而精胺除了干旱种群在1 mM 下有所增加外,在锰胁迫下变化很小。谷胱甘肽含量随锰浓度升高而增加,在0.5 mM 锰时达到最高值,1mM 时有所下降。植物络合素(PCs)含量与非蛋白巯基(NP-SH)趋势相似,随锰浓度的升高而增加,且干旱种群中含量要高于湿润种群。锰处理还引起氧化胁迫,表现为过氧化氢和丙二醛含量增加。SOD 活性在湿润种群中,在0 到0.5 mM 锰胁迫下活性升高,但在1 mM 锰胁迫时,其活性有所下降。而在干旱种群中,SOD 活性变化较小,并始终维持在一个较高的水平。APX 活性在两个种群中都随锰浓度的升高而增加,干旱种群活性要高于湿润种群。锰胁迫还显著增加了酚类物质的含量,同时GPX 和多酚氧化酶(PPO)活性也随锰浓度的升高而增加。干旱种群的酚类含量和GPX 与PPO 活性都要高于湿润种群。锰胁迫还改变了氨基酸的含量和构成,根据锰胁迫下浓度变化的不同,可以将游离氨基酸分为三组:第一组包括,谷氨酸,丙氨酸和天门冬氨酸,这一组氨基酸含量在锰胁迫下有所下降。第二组包括缬氨酸,亮氨酸和苏氨酸含量在锰胁迫下基本不变化或变化很小。剩下的氨基酸为第三组,这组氨基酸含量在锰胁迫下显著增加,而根据增加的幅度又可以将它们分为两个亚组,丝氨酸,酪氨酸,苯丙氨酸,组氨酸和脯氨酸,在1 mM 下的含量是对照的4 倍以上。异亮氨酸,赖氨酸,精氨酸和甘氨酸含量在1 mM 下是对照含量的2 倍以下。同时,同一锰浓度下,干旱种群比湿润种群积累的氨基酸含量要高。 Soil is the indispensable environment for human survival and important resource foragriculture development. Food and environmental problems facing the world are all closelyrelated to soil and nowadays it is threatened by many factors, among which drought stress andheavy metal pollution are the most serious ones. Poplars (Populus spp.) are importantcomponents of ecosystem and suitable as a source of fuel, fiber and lumber due to their fastgrowth. In this study, different populations of Section Tacamahaca spach were used as modelplants to investigate the adaptability to drought stress and manganese toxicity and differencesbetween populations from dry and wet climate regions. Our results can provide theoreticalevidence for the afforestation and prevention of desertification in the arid and semi-arid areas,and also can supply scientific direction for the reconstruction and rehalibitation of ecosystemscontaminated by heavy metals. The results are as follows: 1. Differences in ecophysiological responses to drought stress in two contrastingpopulations of Populus przewalskii Drought stress not only significantly affected dry mass accumulation and allocation, butalso significantly decreased chlorophyll pigment contents and accumulated free proline andtotal amino acids. On the other hand, drought also significantly increased the levels ofabscisic acid and reactive oxygen species, as secondary messengers, to induce the entire set ofantioxidative systems including the increase of reduced ascorbic acid content and the activities of superoxide dismutase, guaiacol peroxidase, ascorbate peroxidase and glutathioneredutase. Thus the combination of drought avoidance and tolerance mechanisms conferredpoplar a high degree of plasticity in response to drought stress. Compared with the wetclimate population, the dry climate population showed lower dry matter accumulation andallocated more biomass to root systems, and accumulated more free proline and total aminoacids for osmotic adjustment. The dry climate population also showed more efficientantioxidant systems with higher content of ascorbic acid and higher activities of ascorbateperoxidase and glutathione redutase than the wet climate population. All these made the dryclimate population superior in adaptation to drought stress than the wet climate population. 2. Effect of exogenous applied SNP on drought tolerance in Populus przewalskii Drought stress significantly increased hydrogen peroxide content and caused oxidativestress to lipids and proteins assessed by the increase in malondialdehyde and total carbonylcontents, respectively. The cuttings of P. przewalskii accumulated proline and other aminoacids for osmotic adjustment to lower water potential, and activated the antioxidant enzymes such as superoxide dismutase, guaiacol peroxidase and ascorbate peroxidase to maintain thebalance of generation and quenching of reactive oxygen species. Moreover, exogenous SNPapplication significantly heightened the growth performance of P. przewalskii cuttings underdrought treatment by promotion of proline accumulation and activation of antioxidant enzymeactivities, while under well-watered treatment the effect of SNP application was very little. 3. Morphological responses to manganese toxicity in the two contrasting populations ofPopulus cathayana High concentration of manganese caused significant decrease in shoot height andbiomass accumulation. The tolerance index of the dry climate population was significantlyhigher than that of the wet climate population, suggesting the superior Mn tolerance in theformer and the existence of cross-tolerance of drought stress and high Mn toxicity. Injuries tothe leaf anatomical features were also found as the reduced thickness in palisade and spongyparenchyma, the decreased density in the conducting tissue and the collapse and split in themeristematic tissue in the central vein. As for the Mn concentrations in the plant tissues, under0, 0.1 and 0.5 mM, most of the Mn accumulated in the roots, then leaves, and stem the least, while under 1 mM, most of the Mn accumulated in the leaves. As far as the total amounts ofMn extraction are concerned, the leaf extracted more Mn than the root and stem in the twopopulations under various Mn concentrations. There is no difference between the twopopulations under control. Under 0.1 mM, the wet climate population extracted higher Mn inthe root than the dry climate population, while in the shoot, the dry climate populationextracted much more Mn. Under 0.5 and 1 mM, the wet climate population translocated moreMn both in the root and the shoot than the dry climate population. 4. Physiological and biochemical responses to manganese toxicity in the two contrastingpopulations of Populus cathayana Mn treatment resulted in oxidative stress indicated by the oxidation to lipids, proteinsand DNA. A regulated network of defence strategies was employed for the chelation,detoxification and tolerance of Mn including the enhanced synthesis of ABA and polyamines,the accumulation of free amino acids, especially His and Pro, and the activation of theenzymes superoxide dismutase and guaiacol peroxidase. Contents of non-protein thiol,reduced glutathione, phytochelatins and phenolics compounds and activities of superoxide dismutase, guaiacol peroxidase and polyphenol oxidase also increased significantly forantioxidant or chelation functions. The wet climate population not only accumulated lessabscisic acid, free amino acids, phytochelatins and phenolics compounds, but also exhibitedlower activities of superoxide dismutase, guaiacol peroxidase and polyphenol oxidase thusresulting in more serious oxidative damage and more curtained growth.
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干旱胁迫是全球范围内影响植物生存、生长和分布的重要环境因子。岷江上游干旱河谷区,由于生态环境的脆弱性和长期人类活动的干扰和过度利用,导致植被严重退化,水土流失加剧,山地灾害频繁,干旱化和荒漠化趋势明显。这种趋势若不能遏制,将严重阻碍区域社会经济的快速协调发展,并且威胁成都平原地区的发展和长江中下游地区的生态安全。因而开展干旱河谷生态恢复研究成为解决这些问题的关键。水分匮乏是限制干旱河谷生态恢复的关键因子,在全球气候变化的背景下,干旱胁迫在区域尺度上可能会更加严重,并使干旱河谷的生态环境更加恶化。因此,深入研究干旱河谷乡土植物对干旱胁迫的响应和适应机理,具有非常重要的理论和实践意义。 本论文以岷江上游干旱河谷的三种乡土豆科灌木,白刺花(Sophora davidii)、小马鞍羊蹄甲(Bauhinia faberi var. microphylla)和小雀花(Campylotropics polyantha)理论和实践意义。为研究对象,在人工控制条件下设计了4-5个连续性干旱胁迫处理,系统地研究了灌木幼苗的生长、生物量积累和水分利用效率(WUE)、形态结构和生理过程等对干旱胁迫的反应,揭示了幼苗的干旱适应能力及种间差异。主要研究结论如下: 1) 灌木生长和繁殖对干旱胁迫的反应 在干旱胁迫下,幼苗生长速率显著减小,叶片衰老和脱落比率增大,这些变化随着胁迫强度的增加具有累积效应。叶片比茎对干旱胁迫的反应更敏感。在严重干旱胁迫下,幼苗的有性繁殖被限制,但在中等程度干旱胁迫下,幼苗的有性繁殖能力被提高。 2) 灌木生物量积累及其分配和WUE对干旱胁迫的反应 在干旱胁迫下,灌木各器官的生物量都显著减小,但是生物量的分配侧重于地下部分,使得根茎比在干旱条件下增大。幼苗的耗水量(WU)随着干旱胁迫的增加而显著减少。白刺花和小马鞍羊蹄甲WUE在干旱胁迫下降低;小雀花的WUE在中等干旱胁迫下升高。 3) 灌木叶片结构特征对干旱胁迫反应 白刺花叶片具有较为典型的旱生型结构,而小马鞍羊蹄甲和小雀花则为中生型结构。在1至2年的干旱胁迫下,灌木叶片结构组成未发生本质性的改变,主要是细胞大小的变化。在中等和严重干旱胁迫下,叶肉组织厚度明显减小;并且气孔和表皮细胞面积也显著减小,气孔和表皮细胞密度显著增大;叶肉细胞层数、P/S值、表皮厚度等无显著变化。 4) 灌木对干旱胁迫的生理响应 气体交换参数和叶片相对含水量(RWC)在中等干旱胁迫下发生了明显的改变,而叶绿素荧光参数和光合色素含量在严重干旱胁迫下才发生显著变化。在干旱胁迫下,净光合作用速率(Pn)、气孔导度(gs)和RWC呈下降趋势,而叶片温度(Tl)呈增加趋势,蒸腾速率(Tr)的变化不明显。除了日最大Pn减小以外,干旱胁迫对气体交换参数的日变化无显著影响,但是对光合-光响应曲线有显著的影响,使有效光合时间缩短。在严重干旱胁迫下光系统受到损害而代谢减弱,PSⅡ中心的内禀光能转换效率(Fv/Fm)、量子产量(Yield)、光化学淬灭参数(qP)显著降低,而非光化学淬灭参数(NPQ)明显增加。气孔限制和非气孔限制对Pn的影响与干旱胁迫强度有关。在中度胁迫下,气孔限制起主导作用,在严重胁迫下非气孔限制起主导作用,40% FC水分条件可能是灌木由气孔限制向非气孔限制的转折点。 5) 灌木对干旱胁迫的适应能力及其种间差异 三种灌木对干旱胁迫具有较好的适应能力,即使在20% FC,幼苗未因干旱胁迫III而死亡;80% FC适宜于幼苗生长。白刺花生长速率慢,耗水量较少,具有较强的耐旱和耐贫瘠能力,并具有干旱忍受机制,能够在较干旱的环境中定居和生长。小马鞍羊蹄甲和小雀花,生长快,水分消耗量较大,尤其是小雀花,对干旱胁迫的忍受能力较弱,具有干旱回避机制,因而适宜于在较为湿润的生境中生长。综合分析表明,生长速率较慢的物种抗旱能力较强,其更适宜于作为干旱地区植被恢复物种。 Drought is often a key factor limiting plant establishment, growth and distribution inmany regions of the world. The harsh environmental conditions and long-termanthropogenic disturbance had resulted in habitat destruction in the dry valley ofMinjiang river, southwest China. Recently, it tended to be more severe on the vegetationdegradation, soil erosion and water loss, natural disaster, as well as desertification, whichimpact on regional booming economy and harmonious development, and would be verydangerous to the environmental security in the middle and lower reaches of Yangzi River.Therefore, ecological restoration in the dry valley is one of the vital tasks in China. Waterdeficit is known to affect adversely vegetation restoration in this place. Moreover, in thecontext of climate change, an increased frequency of drought stress might occur at aregional scale in the dry valleys of Minjiang River. The selection of appropriate plantingspecies for vegetation restoration in regard to regional conditions is an important issue atpresent and in further. The research on responses of indigenous species to drought stresscould provide insights into the improvement of the vegetation restoration in the dry valleys of Minjiang River. In this paper, the responses of three indigenous leguminous shrubs, Sophora davidii,Bauhinia faberi var. microphylla and Campylotropics polyantha, to various soil watersupplies were studied in order to assess drought tolerance of seedlings, and to compare interspecific differences in seedlings’ responses to drought stress. The results were as follows: 1 Growth and reproduction of shrubs in response to drought stress Seedling growth reduced significantly while leaf senescence accelerated underdrought stress, the cumulative responses to prolonged drought were found. The capacityfor reproduction was limited by severe drought stress, and improved by moderate droughtstress. Leaf responses were more sensitive than shoot to various water supplies. 2 WUE, biomass production and its partitioning of shrubs in response to drought stress Drought stress reduced significantly the total dry mass and their components ofseedlings, and altered more biomass allocation to root system, showing higher R/S ratiounder drought. Water use (WU) and water-use efficiency (WUE) of both S. davidii and B.faberi var. microphylla declined strongly with drought stress. The WU C. polyantha ofalso declined with drought stress, but WUE improved under moderate drought stress. 3 Anatomical characteristics and ultrastructures of leaves in response to drought stress There were xeromorphic for S. davidii leaves and mesomorphic for B. faberi var.microphylla and C. polyantha at the all water supplies. The foundational changes in leafstructures were not found with drought stress. However, mesophyll thickness, the areas ofstomatal and epidermis reduced slightly while the densities of stomatal and epidermisincreased under severe drought stress. Variations in these parameters could mainly be duoto cell size. Other structures did not displayed significant changes with drought stress. 4 Physiological responses of shrubs to drought stress The gas exchange parameters and leaf relative water content (RWC) were affectedby moderate stress, while chlorophyll fluorescence and chlorophyll content were onlyaffected by severe stress. Drought stress decreased net photosynthesis rate (Pn), stomatalconductance, light-use efficiency and RWC, and increased leaf temperature. Therespiration rates (Tr) were kept within a narrower range than Pn, resulting in aprogressively increased instantaneous water use effiecency (WUEi) under drought stress.Moreover, drought stress also affected the response curve of Pn to RAR, there was adepression light saturation point (Lsat) and maximum Pn (Pnmax) for moderate andsevere stressed seedling. However, diurnal changes of gas exchange parameters did notdiffer among water supplies although maximum daily Pn declined under severe stress.VISevere stress reduced Fv/Fm, Yield and qP while increased NPQ and chlorophyll content.Photosynthetic activity decreased during drought stress period due to stomatal andnon-stomatal limitations. The relative contribution of these limitations was associatedwith the severity of stress. The limitation to Pn was caused mainly by stomatal limitationunder moderate drought stress, and by the predominance of non-stomatal limitation undersevere stress. In this case, 40% FC water supply may be a non-stomatal limitation 5 Interspecific differences in drought tolerance of shrubs Three shrubs exhibited good performance throughout the experiment process, evenif at 20% FC treatment there were no any seedlings died, 80% FC water supply wassuitable for their establishment and growth. S. davidii minimized their water loss byreducing total leaf area and growth rate, as well as maintained higher RWC and Pncompared to the other two species under drought stress, thus they might be more tolerantto the drought stress than the other two species. On the contrary, it was found that C.polyantha and B. faberi var. microphylla had higher water loss because of their stomatalconductance and higher leaf area ratios. They reduced water loss with shedding theirleaves and changing leaf orientation under drought stress. Based on their responses, thestudied species could be categorized into two: (1) S. davidii with a tolerance mechanismin response to drought stress; (2) C. polyantha and B. faberi var. microphylla withdrought avoidance mechanism. These results indicated that slow-growing shrub speciesare better adapted to drought stress than intermediate or fast-growing species in present orpredicted drought conditions. Therefore, selecting rapid-growing species might leavethese seedlings relatively at a risk of extreme drought.
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本研究针对川西北高山草甸缺乏科学管理,过度放牧导致草场退化,并由此引发的一系列生态环境问题,选取红原县瓦切乡1996 年草地承包后形成的四个放牧强度草场,即不放牧、轻度(1.2 头牦牛hm-1)、中度(2.0 头牦牛hm-1)和重度放牧(2.9 头牦牛hm-1),作为研究对象,研究了不同放牧强度对草地植物-土壤系统中碳、氮这两个最基本物质的分布格局和循环过程的影响,并探讨了放牧干扰下高山草甸生态系统的管理。 1.放牧对草地植物群落物种组成,尤其是优势种,产生了明显的影响。不放牧、轻度、中度和重度放牧草地群落物种数分别为22,23,26,20 种,群落盖度分别是不放牧96.2%>中度93.6%>轻度89.7%>重度73.6%。随放牧强度的增加, 原植物群落中的优势种垂穗鹅冠草( Roegneria nutans )、发草(Deschampsia caespitosa)和垂穗披碱草(Elymus nutans)等禾草逐渐被莎草科的川嵩草(Kobresia setchwanensis)和高山嵩草(Kobresia pygmaea)所取代成为优势种。同时,随放牧强度的增加,高原毛茛(Ranunculus brotherusii)、狼毒(Stellera chamaejasme)、鹅绒委陵菜(Potentilla anserina)和车前(Plantagodepressa)等杂类草的数量也随之增加。 2.生长季6~9 月份,草地植物地上和地下生物量(0~30cm)都是从6 月份开始增长,8 月份达到最高值,9 月份开始下降。每个月份,通常地上生物量以不放牧为最高,重度放牧总是显著小于不放牧;地下生物量随放牧强度的增加表现为增加的趋势,通常重度和中度放牧显著高于不放牧和轻度放牧草地。不放牧、轻度、中度和重度放牧草地6~9 月份4 个月的植物总生物量平均值分别是1543、1622、2295 和2449 g m-2,但随放牧强度的增加越来越来多的生物量被分配到了地下部分,地下生物量占总生物量比例的大小顺序分别是重度88%>中度82%>轻度76%>不放牧69%。生物量这种变化主要是由于放牧使得群落优势种发生改变而引起的,其分配比例的变化体现了草地植物对放牧干扰的适应策略。 3.植物碳氮贮量的季节变化类似与生物量的变化。每个月份,不同放牧强度间植物地上碳氮的贮量有所不同,一般重度放牧会显著减少植物地上碳氮贮量。植物根系(0~30cm)碳氮贮量随放牧强度的增加表现为增加的趋势,通常重度和中度放牧显著高于不放牧和轻度放牧草地。不放牧、轻度、中度和重度放牧草地6~9 月份4 个月的植物总碳平均值分别是547、586、847 和909 g m-2,根系碳贮量占植物总碳的比例大小顺序分别是重度88%>中度82%>轻度76%>不放牧69%;放牧、轻度、中度和重度放牧草地6~9 月份4 个月的植物总氮平均值分别是17、17、23 和26 g m-2,根系氮贮量占植物总氮的比例大小顺序分别是重度79%>轻度71%>中度70%>不放牧65%。 4. 土壤有机碳贮量(0~30cm)的季节变化表现为7 月份略有下降,8 月开始增加,9 月份达到的最大值。土壤氮贮量的季节变化表现为随季节的推移逐渐增加的趋势。增加的放牧强度不同程度的增加土壤有机碳氮的贮量。不放牧、轻度、中度和重度放牧6~9 月份4 个月的土壤有机碳贮量的平均值分别是9.72、10.36、10.62 和11.74 kg m-2,土壤氮贮量分别为1.45、1.56、1.66 和1.83 kg m-2。土壤中有机碳(氮)的贮量都占到了植物-土壤系统有机碳(氮)的90%以上,但不同放牧强度之间的差异不明显。 5. 土壤氮的总硝化和反硝化,温室气体N2O 和CO2 的释放率的季节变化表现为从6 月份开始增加,7 月份达到最大值,8 月份开始下降,9 月份降为最小值。增加的放牧强度趋向于增加土壤氮的总硝化和反硝化作用,温室气体N2O和CO2 的释放率,通常情况下,中度放牧和重度放牧显著地加强了这些过程。 6.垂穗鹅冠草(Roegneria nutans)和川嵩草(Kobresia setchwanensis)凋落物在不同放牧强度下经过1 年的分解,两种凋落物的失重率及其碳氮的损失率3都随放牧增加表现为增加的趋势。在同一放牧强度下,川嵩草凋落物的失重率和碳氮的损失率都高于垂穗鹅冠草凋落物。 7. 尽管重度放牧显著增加了土壤碳氮的贮量,但同时也显著降低了植被群落盖度,降低了植物地上生物量,因此,久而久之会减少植物向土壤中的碳氮归还率;与不放牧和轻度放牧相比,重度放牧又显著增加了土壤CO2 和NO2 的排放量,这是草地生态系统碳氮损失的重要途径。由此可见,对于这些地处青藏高原的非常脆弱的高山草甸生态系统,长期重度放牧不仅导致植物生产力降低,而且将导致草地生态系统退化,甚至造成土壤中碳氮含量减少。 Long-term overgrazing has resulted in considerable deterioration in alpine meadowof the northwest Sichan Province. In order to explore management strategies for thesustainability of these alpine meadows, we selected four grasslands with differentgrazing intensity (no grazing-NG: 0, light grazing-LG: 1.2, moderate grazing-MG: 2.0,and heavy grazing-HG: 2.9 yaks ha-1) to evaluate carbon, nitrogen pools and cyclingprocesses within the plant-soil system in Waqie Village, Hongyuan County, Sichuan Province. 1. Grazing obviously changed the plant species composition, especially ondominant plant species. Total number of species is 22, 23, 26, and 20 for NG, LG, MGand HG, respectively. Vegetation coverage under different grazing intensity ranked inthe order of 96.2% for HG>93.6% for MG>89.7% for LG>73.6% for NG. Thedominator of HG community shifted from grasses-Roegneria nutans andDeschampsia caespitosa dominated in the NG and LG sites into sedges-Kobresiapygmaea and K. setchwanensis. At the same time, with the increase of grazingintensity, the numbers of forbs, such as Ranunculus brotherusii, Stellera chamaejasme,Potentilla anserine and Plantago depressa, increased with grazing intensity. 2. Over the growing season, aboveground and belowground biomass showed a 5single peak pattern with the highest biomass in August. For each month, abovegroundbiomass usually was the highest in the NG site and lowest in the HG site.Belowground biomass showed a trend of increase as grazing intensity increased and itwas significantly higher in the HG and MG site than in the NG and LG sites. Totalplant biomass averaged over the growing season is 1543, 1622, 2295 and 2449 g m-2for NG, LG, MG and HG, respectively. The proportion of biomass to total plantbiomass for NG, LG, MG and HG is 88%, 82%, 76% and 69%, respectively. Higherallocation ratio for is an adaptive response of plant to grazing. 3. Carbon and nitrogen storage in plant components followed the similar seasonalpatterns as their biomass under different grazing intensities. Generally, heavy grazingsignificantly decreases aboveground biomass carbon and nitrogen compared to nograzing. Carbon and nitrogen storage in root tended to increase as grazing increasedand they are significantly higher in the HG and MG sites compared to the LG and NGsite. Total Carbon storage in plant system averaged over the growing season is 547,586, 847 and 909 g m-2 for NG, LG, MG and HG, respectively, while 17, 17, 23 and 26g m-2 for nitrogen. The proportion of carbon storage in root to total plant carbon forNG, LG, MG and HG is 88%, 82%, 76%, 69%, respectively, while 65%, 71%, 70%and 79% for nitrogen. 4. Carbon storage in soil (0-30cm) decreased slightly in July, then increased inAugust and peaked in September. Nitrogen storage in soil tended to increase withseason and grazing intensity. Total Carbon storage in soil averaged over the growingseason is 9.72, 10.36, 10.62 and11.74 kg m-2 for NG, LG, MG and HG, respectively,while 1.45, 1.56, 1.66 and 1.83 for nitrogen. The proportion of carbon (nitrogen)storage in soil to plant-soil system carbon (nitrogen) storage for NG, LG, MG and HGis more than 90%, which is not markedly different among different grazing intensities. 5. Gross nitrification, denitrification, CO2 and N2O flux rates in soil increasedfrom June to July and then declined until September, all of which tended to increasewith the increase of grazing intensity. Generally, heavy and moderate grazing intensitysignificantly enhanced these process compared to no and light grazing intensity. 6. After decomposing in situ for a year, relative weight, carbon and nitrogen loss in the litter of Roegneria nutans and Kobresia setchwanensis tended to increase asgrazing intensity increased. Under the same grazing intensity, relative weight, carbonand nitrogen loss in the litter of Kobresia setchwanensis were higher than these in thelitter of Roegneria nutans. 7. Although heavy grazing intensity resulted in higher levels of carbon andnitrogen in plant and soil, it decreased vegetation coverage and aboveground biomass,which are undesirable for livestock production and sustainable grassland development.What is more, heavy grazing could also introduce potential carbon and nitrogen lossvia increasing CO2 and N2O emission into the atmosphere. Grazing at moderateintensity resulted in a plant community dominated by forage grasses with highaboveground biomass productivity and N content. The alpine meadow ecosystems inTibetan Plateau are very fragile and evolve under increasing grazing intensity by largeherbivores; therefore, deterioration of the plant-soil system, and possible declines insoil C and N, are potential without proper management in the future.