962 resultados para leaf thickness
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The effects of annealing time and Si cap layer thickness: on the thermal stability of the Si/SiGe/Si heterostructures deposited by disilane and solid-Ge molecule beam epitaxy were investigated. It is found that in the same strain state of the SiGe layers the annealing time decreases with increasing Si cap layer thickness. This effect is analyzed by a force-balance theory and an equation has been obtained to characterize the relation between the annealing time and the Si cap layer thickness. (C) 2001 Elsevier Science B.V. All rights reserved.
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The physical properties of low-temperature-deposited GaN buffer layers with different thicknesses grown by metal-organic vapor-phase epitaxy have been studied. A tentative model for the optimum thickness of buffer layer has been proposed. Heavily Si-doped GaN layers have been grown using silane as the dopant. The electron concentration of Si-doped GaN reached 1.7 x 10(20) cm(-3) with mobility 30 cm(2)/V s at room temperature. (C) 1998 Published by Elsevier Science B.V. All rights reserved.
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青藏高原东缘的亚高山针叶林是长江上游重要的生态屏障,经过近六十年的采伐后,取而代之的是大量人工种植的云杉纯林。目前,这些人工林已经表现出树种单一,结构层次简单等生态问题,其物种多样性及生态效益与同地带天然林相比差距较明显。如何丰富该地区物种多样性,完善人工林生态系统的生态功能是一个十分重要的课题。林下植物是人工林群落的重要组成部分,对维持群落的生物多样性及完善生态系统功能具有明显的作用。因此,研究该地区人工针叶林的林下植被对不同生境的适应性对于理解人工林生态系统物种多样性的形成和维持机制都具有重要的意义。 本文以青藏高原东部亚高山针叶林的主要森林类型----云杉人工林为研究对象,选择林下11种具有不同喜光特性的常见植物,分别设置人工林林冠下及成熟林窗为研究样地,通过对各种植物叶片形态与物质分配特征、叶片解剖学特征、叶片光合生理特性、植物自然分布特征等方面的比较分析,研究林下植物对不同光生境的适应策略及其适应能力,揭示不同物种对人工林生境的适应共性,为西南亚高山地区植被恢复及人工林的经营管理提供科学依据。具体研究结果如下: 在叶片形态和物质分配特征方面:在林窗光生境中,11种林下植物叶片比叶重(LMA)显著高于林下光生境的同种植物。同时,林窗下生长的植物叶片叶片厚度及栅栏细胞长度显著增加,这是影响叶片比叶重变化的直接原因。而多数植物叶重比在两种生境中无明显变化。说明在长期适应自然生境之后,植物可能更多地采取调节叶片组织细胞水平(即叶片功能细胞形态)及叶片器官水平(即单个叶片形态)特征的策略来适应各类生境,而非整株水平上的叶片总比重的增减。 在叶片解剖结构特征方面:多数阔叶物种栅栏组织厚度(PT)、栅栏组织厚度/海绵组织厚度(PT/ST)、栅栏细胞层数及近半数种的气孔密度(SD)在林窗生境中更大或更多,而叶片表皮细胞厚度(UET、LET)气孔长径(SL)及海绵组织厚度(ST)受两种生境影响不大。喜光特性相似的物种在生境适应策略上具有一定的趋同性。 在光合生理特征方面:在林窗生境中多数种植物的最大光合速率(Amax)、暗呼吸速率(Rd)及喜光植物光补偿点(LCP)显著或极显著高于林内生境同种植物。且在同一生境条件下,多数深度耐荫植物比喜光及轻度喜光植物有稍低的Rd和LCP。各植物在林内低光生境中具有更大的内禀光能转化效率,并在中午12:00~14:00之间光强最大的时刻发生了的最深程度的光抑制。多数种能通过调节自身某种光合素含量或色素之间的比例来适应不同的光生境,即通过增加叶绿素含量或降低Chla/b值来适应林内弱光生境,通过提高类胡萝卜素含量或单位叶绿素的类胡萝卜素含量降低强光带来的伤害。绝大多数物种并不采取调节叶片C、N含量的策略来适应不同的光生境。总之,植物部分光合参数(Amax、Rd、LCP)受生境的影响与其自身喜光特性有关,但另一些参数(Fv/Fm日变化、色素含量及比例、叶氮相对含量)受生境影响与其自身喜光特性无明显关联。 在表型可塑性方面:在叶片各表型参数中,器官水平及细胞水平的形态特征参数平均可塑性大于整株水平形态和物质分配特征参数可塑性;叶片光合组织的可塑性大于非光合组织可塑性;反映植物光合能力的参数可塑性大于叶片色素含量参数可塑性。植物叶片形态和物质分配、解剖学特征参数平均可塑性大小与其自身喜光特性基本吻合,即喜光种及轻度耐荫种各参数可塑性最高,深度耐荫种可塑性最小,而这种规律并未在光合生理参数的可塑性大小上体现出来。但是叶片形态和物质分配参数、光合生理参数的平均可塑性水平却大于叶片解剖学参数。 在植物自然分布特征方面:喜光物种云杉幼苗及歪头菜在林内生境中分布密度明显降低,深度耐荫种疏花槭却恰恰相反,更多数物种(7种植物)在两种生境中密度变化趋势不明显。从分布格局来看,7种植物在两种生境中均为聚集分布,但聚集强度为林窗>林内;少数物种桦叶荚迷、直穗小檗、冰川茶藨、黄背勾儿茶在林窗中为聚集型,在林内生境中的分布型发生改变而成为随机型,说明光生境的差异能影响到植物种群的分布特征。但这种影响程度与植物自身的喜光特性无关,同时与各物种叶片表型平均可塑性的大小也无明显关联。 The subalpine coniferous forest area in eastern Qinghai-Tibet Plateau is important ecology-barrier of upriver Yangtze. In past sixty years, those forests had been cut down and replaced with a lot of spruce plantations. At now, there are many ecology problems presenting to us such as singleness species, simple configuration, lower species diversity and ecological benefit than natural forests at the same belt. How to restore the species diversity and enhance the eco-function of the plantations is a very important issue. The understory plants are important part of plantation community, which improved the bio-diversity and eco-function distinctly of forests. So, it is very significance to study the adaptation of understory plants to different environment in plantation, and this study would helping us to understand how plantations to develop and remain their biodiversity. This study was conducted in a 60a spruce plantation in Miyaluo located in western Sichuan, China, and spruce plantation is major types of subalpine coniferous forest in eastern Qinghai-Tibet Plateau. In this paper, the leaf morphological and biomass-distributed characteristics, the anatomical characteristics, the photosynthetic characteristics and the distribution patterns characteristics of eleven different light-requirement understory species grown in two different environments (forest gaps and underneath close canopy) were studied and compared. The purpose of this study was to analyze the adaptation of this forest understory plants, to show up the commonness of these different light-requirement understory species in light acclimation, and to provide some scientific reference to manage and restore the vegetation of subalpine plantation of southwest China. The results were as follows: The leaf morphological and biomass-distributed characteristics: These eleven species in forest gaps had significantly higher dry weight per leaf area (LMA) than those under close canopy. The palisade parenchyma cells of the broad-leaved species in gaps were significantly longer than those grown under the canopy, which been a directed factor for the change of leaf mass per unit area (LMA) in different environment. But the leaf weight ratio (LWR) of most plants species were not evidently changed by the contrasted environments in our study. It was shown the morphological characteristics changing been adopted as a strategy of light acclimation for plants wasn’t on whole plant level (leaf weight ratio) but cellular level (the function cells morphological characteristics) and organic level (the leaf morphological and biomass-distributed characteristics) mostly. The leaf anatomical characteristics: Most broad-leaved plants in gaps increased palisade parenchyma thickness (PT), the palisade parenchyma cell layers and the ratio of palisade to spongy parenchyma (PT/ST). So did as almost about half species in this study in stomatal density (SD). No significant differences in thickness of leaf epidermal cells (UET, LET), stomatal length (SL) and spongy parenchyma (ST) between two environments of most species were observed. The results suggested that species with light-requirement approximately had convergent evolution on adaptation to light condition. The leaf photosynthetic characteristics: The dark respiration rate (Rd) of most plants species, the light compensation point (LCP) of light-demanding plants species in gaps were significantly increased than under close canopy in this study. In a same habitat, most deep-shade-tolerant plants had lower Rd and LCP than those light-demanding plants and slight-shade-tolerant plants. Each species has bigger inherent electron transport rate under close canopy than in gaps, and the greatest photoinhibition happened during 12 to 14 in the daytime. Most species could adapt different light environment by the way of changing their photosynthetic pigments content or the ratio of pigments content. For example, some plants under close canopy increased chlorophyll (Chl) or reduced the values of the ratio Chla/b to adapted the low light condition, some plants in gaps increased carotenoid (Car) or reduced the weight ratio CarChl to avoid been hurt in high light. For most plants, changing the content of C and N in leaf wasn’t a strategy of light acclimation. In conclusion, the variation of some leaf photosynthetic parameters in different light environment such as Fv/Fm, pigments, C and N in leaf related with the light-requirmnet of species, but the others such as Amax, Rd, LCP did not. The leaf plasticity indexes: Among those leaf plasticity indexes, the leaf morphological and biomass-distributed parameters on cellular and organic level were greater than on whole plant level for same species, and the photosynthetic parenchyma parameters were greater than non-photosynthetic parenchyma parameters in same leaf, and photosynthetic capability parameters were greater than photosynthetic pigments content parameters for same species. The average plasticity indexes of leaf morphological and biomass-distributed and anatomical parameters were accordant with plants’ light-requirement approximately: those light-demanding plants and slight-shade-tolerant plants had bigger plasticity indexes than deep-shade-tolerant plants. But this regular wasn’t observed in physiological plasticity indexes for most plants, though the average leaf plasticity indexes of leaf morphological and biomass-distributed, photosynthetic characteristics parameters was greater than the anatomical characteristics parameters. The distribution patterns characteristics: Oppositely to the deep-shade-tolerant specie Acer laxiflorum Pax., the density of light-demanding species Picea asperata Mast. and Vicia unijuga A. Br. in gaps was bigger than under close canopy. Each of the other species has the approximately density in two different environment. The spatial patterns of seven species were aggregated distribution in two environments, but the trend of aggregation of population under close canopy was decrease from in gaps. A few species such as Viburnum betulifoium Batal., Berberis dasystachya Maxim., Ribes glaciale Wall. and Berchemia flavescens Brongn. were aggregated distribution in gaps while random distribution under close canopy. It was shown that the difference between two light environments could affect the distribution pattern of plant population, and the effect didn’t relate with the light-requirement or plasticity indexes of species.
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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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繁殖更新是植物生活史的重要阶段,在退化生态系统中,植物繁殖更新能力往往较差,是植被恢复的限制环节,因而也成为恢复研究重点和核心。本研究选择岷江干旱河谷广泛分布的三种蔷薇:多苞蔷薇(R. multibracteata)、黄蔷薇(R. hugonis)和川滇蔷薇(R. soulieana)为研究对象,通过野外调查,在查明其生长、繁殖更新状况的基础上,采用控制和模拟实验,对种子和幼苗阶段进行了深入研究,综合分析更新潜力,并提出相对应的促进更新和植被恢复措施。主要结论如下: 1)三种蔷薇在岷江干旱河谷广泛分布,生长和繁殖状况良好,结实量大。各生长指标:株高、基径和冠幅,繁殖指标:结实数量、重量和单果重量都具有显著的空间差异性。基径对多苞蔷薇结实量影响最大;而冠幅对黄蔷薇结实量影响最大。海拔和纬度是对蔷薇生长和繁殖影响最大的环境因素,随着海拔和纬度的升高,植株生长更高大,结实量增加;坡度和坡向对其生长和繁殖也有一定影响,随着坡度 和坡向增加,蔷薇生长和结实受到抑制。 2)三种蔷薇在岷江干旱河谷更新现状不佳, 但更新潜力大。活力种子比率低,动物取食以及两年生幼苗的大量死亡是蔷薇更新的主要限制因素。多苞蔷薇和黄蔷薇的结实率低,川滇蔷薇较高。三种蔷薇种子产量大,但种子质量较差,更新具有充足的种源。三种蔷薇都能形成持久种子库,种子库中种子总量大,但有效种子少,黄蔷薇被动物啃食的比例很高,多苞蔷薇和川滇蔷薇也有一部分种子受到动物破坏。三种蔷薇幼苗库组成特征表现为,当年生幼苗所占比例很高,年龄较大幼苗所占比例小。 3)三种蔷薇都具有不同程度休眠,未经处理种子的发芽率极低。黄蔷薇休眠程度最深,为深度生理休眠;多苞蔷薇为中度生理休眠;川滇蔷薇为非深度生理休眠。三种蔷薇种子在形态上发育成熟,种皮具有透水性。蔷薇果果肉和瘦果中含有抑制物质,其浸泡液抑制了油菜种子萌发,果肉抑制作用更强,果肉和瘦果浸泡液的抑制程度分别为:川滇蔷薇>黄蔷薇>多苞蔷薇。切割和硫酸腐蚀提高了川滇蔷薇种子的发芽能力,而对多苞蔷薇和黄蔷薇没有影响。完全去除瘦果果皮和种皮提高了多苞蔷薇种子发芽率,但对黄蔷薇没有影响。赤霉素和烟水对蔷薇种子萌发没有促进作用。三种蔷薇打破休眠所需低温层积时间分别为:黄蔷薇>多苞蔷薇>川滇蔷薇。对于多苞蔷薇和川滇蔷薇,层积前对种子进行硫酸腐蚀或暖温层积能缩短低温层积时间,提高发芽率。对于多苞蔷薇,变温层积中暖温层积和低温层积具有一定的负补性,即延长暖温层积可以缩短种子萌发对低温层积的需要。 4)多苞蔷薇种子形态特征和种子休眠与萌发在不同海拔梯度间存在较大差异。种子采集时间、采集季节和干藏影响多苞蔷薇和川滇蔷薇的种子休眠。多苞蔷薇果实大小、种子大小和千粒重、种皮厚度随海拔升高而增加,而种子饱满率和活力随海拔升高而降低,种子休眠程度也随海拔升高而增加。种皮厚度与种子大小、千粒重成正相关关系,硫酸腐蚀后的种子经过不同时间的低温层积后,种子发芽率与种皮厚度、种子大小、千粒重、海拔成正相关关系。2006 年采集川滇蔷薇和多苞蔷薇种子休眠程度较2005 年低。种子休眠随种子年龄增加而减弱。高温和干旱能减轻多苞蔷薇和川滇蔷薇种子休眠。 5)三种蔷薇的生长和生物量积累在干旱胁迫条件下受到抑制,而生物量分配、叶片形态特征和水分利用特征等都发生了变化。三种蔷薇的根、茎、叶各器官生物量以及总生物量等在干旱胁迫下明显减小,叶片脱落数量增加。在干旱胁迫条件下,较多的生物量分配到地下部分,从而这使R/S 明显增加。比叶面积(SLA)和冠层面积比(LAR)对干旱胁迫的反应不敏感,仅有部分物种在干旱胁迫条件下发生了变化,并且其变化特点在不同年龄幼苗之间有一定差异。干旱胁迫对WUE 的影响在不同物种间存在差异。多苞蔷薇和黄蔷薇的WUE 随着干旱胁迫的增加而增大, 而川滇蔷薇的WUE 则随干旱胁迫增加而减小。在干旱胁迫条件下,多苞蔷薇和黄蔷薇叶片脱落量和生物量减小幅度较川滇蔷薇大,表明其抗旱能力较强。在干旱胁迫条件,三种蔷薇两年生幼苗的生物量减小幅度较当年生幼苗小,表明两年生幼苗的抗旱能力更强。 6)两种植被恢复措施中,幼苗移栽比播种具有更好的植被恢复效果。播种后,蔷薇种子的发芽率较高,但出苗率都很低,即使出苗,幼苗也几乎在一月内全部死亡。 三种微生境条件下(灌木、半灌木和裸地),种子出苗和幼苗成活没有差异。移栽幼苗总体死亡率都比较低,小于20%。特别是两年生幼苗死亡率更低,小于2%。移栽后的幼苗生长状况良好,在整个生长季中,各生长指标不断增加。生境对幼苗的存活率没有显著影响,但对于幼苗的生长和生物量积累有一定影响,裸地更有利于幼苗生长和生物量积累。与当年生幼苗相比,两年生幼苗具有更高的成活率。总之,三种蔷薇在干旱河谷分布广泛、生长繁殖状况良好,结实量大,具有丰富种源,繁殖更新潜力大,但繁殖更新状况不佳;种子散布后动物对种子的取食、种子的深度休眠过程、种子出苗以及当年生幼苗的存活和定居是更新的主要限制环节。水分是影响结实、种子休眠解除和萌发,幼苗存活和定居的最主要的限制因素。在植被恢复中,应在种子成熟季节大量采集种子,在室内打破休眠后进行人工播种,培育两年生幼苗,通过幼苗移栽方式进行植被恢复。川滇蔷薇应栽种在相对湿润的过渡区,而多苞蔷薇和黄蔷薇可以应用于核心区植被恢复。 Regeneration is an important phase in plant life cycle. It has been a key component of ecological restoration in degradation ecosystem in which plants commonly has poor regeneration. In this paper, we investigated the natural growth, propagation and regeneration status of native three rose species, Rosa multibracteata, R. hugonis and R. soulieana, and analysis the limitation in seed germination and seedling establishment stages. Advice on facilitating the use of these plants in restoration based on the results has been proposed. The results were as follows: 1) Three rose plants widely distributed in the dry valley of the Minjiang River, and made a good performance in growth and propagation. There were significant spatial differences in each growth parameter, such as ramet height, basal diameter, crown diameter and propagation parameters including hip number of a clump, hip mass of a clump and a hip mass. Basa diameter was the most important growth parameter influencing fruit number for R. multibracteata and crown diameter was for R. hugoni. Altitude and latitude had the greatest effect on the growth and propagation of rose plants among environmental conditions. Each parameter of growth and propagation increased with the increase of altitude and latitude. In addition, the increase of slope and aspect limited the growth and propagation. 2) Three rose plants had poor natural regeneration, but great regeneration potential. Low seed viability, predation and higher mortality of current year old seedlings were the limitation in regeneration. R. multibracteata and R. hugonis had higher fruiting rates than R. souliean. All three plants produced a great number of seeds, while their viability was poor. Three rose plants had persistent seed banks, with high total seed number but very low viable seed density. Predation was most severe in R. hugonis, and it also existed to some degree in R. multibracteata and R. soulieana. The seedling age-structure was characteristic of current-year seedlings predominating and few older seedlings were observed. 3) Three rose seeds were dormant and untreated seeds germinated with very low germination percentages. The rose seeds had morphological mature embryos, and achenes were permeable. Some inhabit substances existed in hips and achenes for the extracts of hips and achenes inhibited germination of Brassica campestris. The inhibition effect of the extracts of three rose hip and achenes was R. soulieana>R. hugonis>R. multibracteata. Mechanical and H2SO4 scarification increased R. soulieana germination but had no effect on germination of R. hugonis and R. multibracteata seeds. Full removal of pericarp and testa improved the germination of R. multibracteata but did not affect R. hugonis germination. GA3 and smoke water had no positive effect on rose seed germination. The periods of cold stratification required to released seed dormancy was R. hugonis > R. soulieana >R. multibracteata. H2SO4 scarification and warm stratification shortened cold stratification to release dormancy for R. soulieana and R. multibracteata. Warm stratification had complementary effect for cold stratification, i.e. the longer warm stratification seeds received, the shorter cold stratification were required to obtain the same germination percentage. Three rose seeds had different kinds of dormancy; R. hugonis has deep physiological dormancy, R. multibracteata with intermediate physiological dormancy and R. souliean non-deep physiological dormancy. 4)The seeds traits and dormancy of R. multibracteata showed significant difference across altitudes. Year and season of seed collection had significant effect on seed dormancy for both R. souliean and R. multibracteata. Hip size, seed size, seed weight, seed coat thickness and seed dormancy level increased with the increase of the altitude. There were positive relations between seed coat thickness with seed size and seed weight. Germination percentage of seeds treated with H2SO4 scarification following different periods of cold stratification showed positive relation with seed coat thickness, seed size, seed weight and altitude. Seeds of R. souliean and R. multibracteata collected in 2006 had low dormancy level than those collected in 2005. Seed dormancy decreased with increasing seeds age. High temperature and drought were associated with low dormancy level. 5) Seedling growth, the total dry mass and their components of seedlings were reduced, while leaf senescence accelerated under drought stress. More biomass allocation to root system resulted in higher R/S ratio under drought. Water-use efficiency (WUE) of R. multibracteata and R. hugonis increased, while it declined for R. soulieana under drought stress. R. soulieana seedlings had poor drought-resistance capacity it had more senescent leaves, and its reduction of biomass was stronger than two other rose plants under drought. The reduction degree of one year old seedlings under drought stress was slighter than that of current year seedlings. Therefore, one year old seedling was more drought-resistent compared to current year seedlings. 6)Planting seedlings may have better effect in comparison with direct seeding. Most seeds germinated after seeding, but seedling emergence was very low. More than 80 % seedlings from direct seeding died within a months after emergence. Seedling emergence and survival rate did not show difference among microhabitats. Mortality rates of seedlings artificially planted in microhabitats were general lower than 20 %, and the mortality rate of one year old seedlings was lower than 2 %. Each grow parameter including plant height, leaf number and branch number continually increased after planting. Microhabitat type had effect on the growth parameter and biomass production, but it did not influence the seedling survival. Bare land tended to facilitate seedling growth. One year old seedlings had higher survival rate than current year seedlings. In conculsion, the three rose had wide distribution in the dry valley of the Minjiang River. They produced many seeds and had tolerance to drought stress to some degree. But they had poor regeneration in habitats may be caused by predation, seed dormancy,and high mortality in current year seedlings. We recommend that rose plants should be utilized in restoration by planting two-year old seedlings in spring. A large quantity of seeds should be collected artificially in autumn, release seed dormancy in room, and then cultivate two-year old seedlings by seeding in particular container. R. soulieana seedling probably be planted in transition area, and R. multibracteata and R. hugonis can be used in core area of the dry valley of the Minjiang River.