443 resultados para FV
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PS II photochemical efficiency (F-v/F-m) of Nostoc flagelliforme was examined after rewetting in order to investigate the light-dependency of its photosynthetic recovery. F-v/F-m was not detected in the dark, but was immediately recognized in the light. Different levels of light irradiation (4, 40 and 400 mu mol photon m(2) s(-1)) displayed different effects on the recovery process of photosynthesis. The intermediate level led to the best recovery of photochemical efficiency; the low light required longer and the high light inhibited the extent of the recovered efficiency. It was concluded that the photosynthetic recovery of N. flagelliforme is both light-dependent and influenced by photon flux density.
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The effects of wind speed on loss of water from N. flagelliforme colonies were investigated indoors in an attempt to assess its ecological significance in field. Wind enhanced the process of waterless; the half-time of desiccation at wind speeds of 2.0 and 3.4 m s(-1) was, respectively, shortened to one-third and one-fifth at 20 degrees C and, to one-sixth and one-eighth at 27 degrees C that of still air. Photosynthetic efficiency was not affected before the wet alga lost about 50% water.
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基于时序逻辑CTL(computation tree logic)的一种扩展CTL-FV对优化编译中的语句交换和变量替换这两种常见变换的保义性条件给出了形式刻画,采用含条件重写规则定义了保义语句交换Texch和保义变量替换Tsub,并基于一种归纳证明框架对它们的保义性进行了证明.此外,基于变换Texch对程序基本块内保依赖语句重排的保义性也给出了一种构造性的证明.
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以小麦品种‘小偃6号’(氮高效品种)和‘长旱58’(氮低效品种)为材料,采用开顶式气室和土培实验研究了大气NH3浓度升高对生长于高、低两种供氮介质下小麦植株不同生育期叶片净光合速率(Pn)、气孔导度(Gs)、叶绿素含量、叶绿素荧光参数(Fv/Fm、Fv/F0)和可溶性糖含量的影响。结果显示:两小麦品种高氨低氮处理植株的Pn、Fv/F0和可溶性糖含量均高于高氨高氮和低氨低氮处理,并在生育后期差异达显著水平(P<0.05),氮低效品种的Gs也符合上述规律且不同处理间差异显著(P<0.05);小麦各生育期高氨高氮处理下植株的Pn均显著低于低氨高氮处理,且两处理间灌浆期的叶绿素含量和灌浆期以前的可溶性糖含量的差异显著(P<0.05),而两处理灌浆期以前的叶绿素荧光参数Fv/Fm在各处理条件下均无显著差异;不同处理间及品种间各项光合特征指标差异缺乏规律性。可见,大气中NH3浓度升高有利于改善低供氮介质条件下小麦植株的氮营养状况,但不同氮效率品种间的响应存在差异。
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大田条件下设置不同的施氮和灌水水平,研究施氮量和灌水量对冬小麦(Triticum aestivum L.)叶片光合作用生理及籽粒形成的影响。结果表明,施氮和灌水提高叶片Fv/Fm和Pn,并显著提高冬小麦籽粒产量、单位面积穗数和每穗粒数。施氮明显降低千粒重,而适量灌水明显提高千粒重。
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大气环境中O3和CO2浓度的升高已经引起了广泛关注,但二者交互作用对城市中树木光合生理的影响机制尚不清楚。蒙古栎(Quercus mongolica)、华山松(Pinus armandi)是沈阳市城市森林的两个重要树种,对大气环境变化的响应具有代表性。本文采用开顶箱法研究了在高浓度O3(80 nmol•mol-1)、CO2(700 μmol•mol-1)及其复合作用下沈阳市蒙古栎、华山松生长、光合、蒸腾的日动态、季节动态变化,揭示了高浓度O3、CO2及其复合作用对两树种光合生理的影响机制,为研究城市森林对全球变化的响应提供重要理论基础。 得出的主要结果如下: 1. 高浓度O3处理后,蒙古栎叶片表现为棕斑型伤害,华山松针叶则为叶尖变黄,两树种光合“午休”程度加重,叶片生长受到抑制。同时,可溶性蛋白、可溶性糖、淀粉含量降低。 两树种叶绿素含量降低,净光合速率下降。蒙古栎光合速率的降低前期主要表现为气孔限制,后期转为非气孔限制为主。华山松光合速率下降则主要受非气孔限制,表明其针叶气孔开闭对光合的影响要小于蒙古栎。造成两树种光合速率下降的非气孔因素是表观量子效率和羧化效率的降低。两树种光呼吸的升高是逆境下的一种保护机制。 在叶绿素荧光方面,两树种Fo、Fm、Fv/Fm、φPSII、ETR、qP降低,NPQ升高,表明高浓度O3作用下, PSII反应中心的开放程度降低、电子传递速率下降,用于光合光化学反应的光能减少,过剩能量增加,热耗散增加。 2. 高浓度CO2处理短期内提高了两树种叶片胞间CO2浓度、羧化效率、表观量子效率和光反应能力(φPSII、ETR和qP),从而提高了蒙古栎、华山松的净光合速率。一段时间后表现出光合下调的现象,羧化效率的降低和光反应能力(φPSII、ETR和qP)的下调可能是发生光合适应的主要原因。 高浓度CO2处理促进两树种叶片的生长,提高了可溶性蛋白、可溶性糖、淀粉含量。日变化曲线趋向单峰曲线,缓解了光合“午休”现象。 3. 复合处理效应,高CO2处理缓解了高O3处理的不利影响,减轻了高浓度O3对叶片的伤害,与单因子O3处理相比较,Pn、AQY、CE、Fv/Fm、qP升高,NPQ降低。另外,在叶片代谢产物、叶绿体色素含量、以及叶生长量的变化上缓解作用均有体现。
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本文系统研究了长白山阔计卜红松林主要树种红松、水曲柳、胡桃揪和锻树在模拟水分胁迫下的生理生态变化。研究表明:(l)土壤水分胁迫对供试树种的生比清况有显:影响。在轻度水分胁迫下(Mw),红松和水曲柳的地下生物量/地上生物量值比对照组(CK)高,而胡桃揪和锻树的地下生物燮/地上生物量比对照组(CK)低。在重度水分胁迫下(LW),供试树种中只有红松的地下生物量/地上生物量比对照组高。(2)土壤水分胁迫对供试树种的生理生态指标有显著影响。在轻度水分胁迫下(Mw),红松和水曲柳的光合速率和水分利用效率比对照组(CK)高,胡桃揪和锻树的光合速率和水分利用效率比对照组(CK)低。在重度水分胁迫下(LW)供试树种的光合速率和水分利用效率都比对照组低。(3)在正常水分条件下(CK),水曲柳光合速率曰变化呈单峰曲线,轻度(Mw)和重度水分胁迫(LW)下均呈"双峰"曲线。正常水分条件下(C)水分利用率的日变化呈双峰曲线,轻度和重度水分胁迫下呈现波动趋势,无明显的波峰和波谷。(4)水曲柳的光合速率和水分利用效率与各因素间多元线性回归的结果表明,三种水分处理下的蒸腾速率、气温、叶温、气孔导度、胞间CO2浓度、光合有效辐射均对光合速率,水分利用效率有一定影响,但影响程度有所差别。(5)土壤水分胁迫对水曲柳、胡桃揪和锻树的初始荧光(Fo)、可变荧光(Fv)、最大荧光(Fm)的影响不大,但却显著影响各树种的PSII原初光能转换效率(Fv/Fm)和PSII潜在活性(Fv/Fo)。(6)土壤水分胁迫影响供试树种几!中的养分含量,但影响程度是不同的,水分胁迫对磷、钾素含量的影响比对氮的大。土壤水分亏缺对各树种叶中的Ca元素无显著影响。红松、胡桃揪和极树叶片中的Mg含量随着土壤水分胁迫的加重也随之逐渐减小,但水曲柳在轻度水分胁迫下叶中的Mg含量最高,关于这一现象还有待以后开展深入的研究。
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数值摄动算法将流体动力学效应耦合进NS方程组和对流扩散(CD)方程离散的数学基本格式(MBS),特别是耦合进最简单的一阶迎风和二阶中心格式之中,由此构建成一系列新的摄动格式(PS).构建PS的主要步骤是将MBS中的通量重构为步长的幂级数,利用空间分裂和导出的高阶流体动力学线性关系式,并引入下游不影响上游的对流运动规律,通过消除重构格式修正微分方程的截断误差诸项求出幂级数的待定系数,由此获得非线性PS.PS的项是MBS中对应项与R△x(及λR△x)之简单多项式的乘积,R△x和λ分别是网格Reynolds数和网格CFL数.PS和MBS使用相同结点,简单性彼此相当,但PS精度高,稳定范围大,例如PS包含了许多绝对稳定高阶迎风和中心有限差分(FD)格式和绝对正型有限体积(FV)格式,这些格式对网格Reynolds数的任意值均为不振荡格式.数值摄动算法因此是构建高精度不振荡CFD格式的新方法.PS用于计算不可压缩流,可压缩流,液滴萃取传质,微通道两相流等,均获得良好数值结果或与已有Benchmark解一致的数值结果.已有文献称数值摄动算法为新型高精度方法和高算法,文中也讨论了一些值得进一步研究的课题
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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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在人类活动导致全球变暖的前提下,由于全球气温的升高,地表水分加速向空中蒸发。从20世纪70年代至今,地球上严重干旱地区的面积几乎扩大了一倍。这一增长的一半可归因于气温升高而不是降雨量下降,因为实际上同期全球平均降水量还略有增长。干旱对陆地植物和农林生态系统产生深远影响,并已成为全球变化研究的一个重要方面。位于青藏高原东部的川西亚高山针叶林是研究气候变暖对陆地生态系统影响的重要森林类型。森林采伐迹地、人工林下和林窗环境作为目前该区人工造林和森林更新的重要生境,其截然不同的光环境对亚高山针叶林更新和森林动态有非常重要的影响。凋落物产生的化感物质可通过影响种子萌发和早期幼苗的定居而影响种群的建立和更新,而人工林和自然林物种以及更新速度的差异性也都受凋落物的影响。 云杉是川西亚高山针叶林群落的重要树种之一,在维持亚高山森林的景观格局和区域生态安全方面具有十分重要的作用,其自然更新能力及其影响机制一直是研究的热点问题。本试验以云杉种子和2年生幼苗为研究对象,从萌发、根尖形态、幼苗生长、光合作用、渗透调节和抗氧化能力等方面研究了不同光环境下水分亏缺和凋落物水浸液对云杉种子和幼苗生长的影响。旨在从更新的角度探讨亚高山针叶林自然更新的过程,其研究成果可在一定程度上为川西亚高山针叶林更新提供科学依据,同时也可为林业生产管理提供科学指导。主要研究结论如下: 水分亏缺在生长形态、光合作用、抗氧化能力、活性氧化对云杉幼苗都有显著影响。总体表现为,水分亏缺导致了云杉幼苗的高度、地径、单株总生物量降低,增加了地下部分的生长;水分亏缺显著降低了云杉叶片中相对含水量、光合色素、叶氮含量,净光合速率和最大量子产量(Fv/Fm),提高了幼苗叶片中膜脂过氧化产物(MDA)的含量;水分亏缺提高了幼苗叶片中过氧化氢(H2O2)含量,超氧荫离子(O2-)生成速率以及脯氨酸和抗氧化系统的活性(ASA, SOD, CAT, POD, APX和GR)。从这些结果可知,植物在遭受水分亏缺导致的伤害时,其自身会形成防御策略,并通过改变形态和生理方面的特性以减轻害。但是,这种自我保护机制依然不能抵抗严重水分亏缺对植物的伤害。 模拟林下低光照条件显著增加单株植物的地上部分生长,尤其是其叶片的比叶面积(叶面积/叶干重),同时其光合色素含量和叶片相对含水量也显著增加,这些改变直接导致植株光合速率和生物量的增加。同时,与高光照水平相比,低光照幼苗的膜脂过氧化产物(MDA)和活性氧物质均较低,显示出低光照比高光照水平对植物的更低的氧化伤害。尽管低光照也导致大部分抗氧化酶活性降低,但这正显示出植物遭受低的氧化伤害,更印证了前面的结论。 凋落物水浸液影响了云杉种子的萌发和根系的生长,更在形态、光合作用、抗氧化能力、活性氧物质以及叶氮水平上显著影响了云杉幼苗,其中,以人工纯林凋落物的影响更有强烈。具体表现在,种子萌发速率和萌发种子幼根的长度表现为对照>自然林处理>人工纯林;凋落物水浸液抑制种子分生区和伸长区的生长,人工林处理更降低了根毛区的生长,使根吸水分和养分困难。对2年生幼苗的影响主要表现在叶绿素含量、光合速率以及叶氮含量的降低;膜脂过氧化产物、活性氧物质和抗氧化酶系统的显著增加。同样的,人工纯林处理对云杉幼苗的影响显著于自然林处理。 在自然生态系统中,由于全球变暖气温升高导致的水分亏缺和森林凋落物都存在森林的砍伐迹地,林窗和林下环境中。我们的研究表明,与迹地或林窗强光照比较,林下的低光照环境由于为植物的生长营造了较为湿润的微环境,因此水分亏缺在林下对云杉幼苗造成的影响微弱。这可以从植物的形态、光合速率以及生物量积累,过氧化伤害和抗氧化酶系统表现出来。另一方面,凋落物水浸液在模拟林下低光照环境对植物的伤害也微弱于强光照环境,这与强光照环境高的水分散失导致环境水分亏缺有关;而人工纯林处理对云杉幼苗的伤害比对照和自然林处理显示出强烈的抑制作用。 Under the pre-condition of global warming resulted from intensive human activities, water in the earth’s surface rapidly evaporates due to the increase of global air temperature. From 1970s up to now, the area of serious drought in the world is almost twice as ever. This increase might be due to the increasing air temperature and not decreasing rainfall because global average rainfall in the corresponding period slightly is incremental. Drought will have profound impacts on terrestrial and agriculture-forest system and has also become the important issue of global change research. The subalpine coniferous forests in the eastern Qinghai-Tibet Plateau provide a natural laboratory for the studying the effects of global warming on terrestrial ecosystems. The light environment significantly differs among cutting blanks, forest gap and understory, which is particularly important for plant regeneration and forest dynamics in the subalpine coniferous forests. Picea asperata is one of the keystone species of subalpine coniferouis forests in western China, and it is very important in preserving landscape structure and regional ecological security of subalpine forests. The natural regeneration capacities and influence mechanism of Picea asperata are always the hot topics. In the present study, the short-term effects of two light levels (100% of full sunlight and 15% of full sunlight), two watering regimes (100% of field capacity and 30% of field capacity), two litter aqueous extracts (primitive forest and plantation aqueous extracts) on the seed germination, early growth and physiological traits of Picea asperata were determined in the laboratory and natural greenhouse. The present study was undertaken so as to give a better understanding of the regeneration progress affected by water deficit, low light and litter aqueous extracts. Our results could provide insights into the effects of climate warming on community composition and regeneration behavior for the subalpine coniferous forest ecosystem processes, and provide scientific direction for the forest production and management. Water deficit had significant effects on growth, morphological, physiological and biochemical traits of Picea asperata seedlings. Water deficit resulted in the decrease in height, basal diameter, total biomass and increase in under-ground development; water deficit significantly reduced the needle relative water content, photosynthetic pigments, needle nitrogen concentration, net photosynthetic rate and the maximum potential quantum yield of photosynthesis (Fv/Fm), and increased the degree of lipid peroxidation (MDA) in Picea asperata seedlings; water deficit also increased the rate of superoxide radical (O2-) production, hydrogen peroxide (H2O2) content, free proline content and the activities of antioxidant systems (ASA, SOD, POD, CAT, APX and GR) in Picea asperata seedlings. These results indicated that some protective mechanism was formed when plants suffered from drought stress, but the protection could not counteract the harm resulting from the serious drought stress on them. Low light in the understory significantly increased seedling above-ground development, especially the species leaf area (SLA), and photosynthetic pigments and relative needle content. These changes resulted in the increase in net photosynthetic rate and total biomass. Moreover, the lower MDA content and active oxygen species (AOS) (H2O2 and O2-) in low light seedlings suggested that low light had weaker oxidative damage as compared to high light. Lower antioxidant enzymes activities in low light seedlings indicated the weaker oxidative damage on Picea asperata seedlings than high light seedlings, which was correlative with the changes in MDA and AOS. Litter aqueous extracts affected seed germination and root system of Picea asperata seedlings. Significant changes in growth, photosynthesis, antioxidant activities, active oxygen species and leaf nitrogen concentration were also found in Picea asperata seedlings, and plantation treatment showed the stronger effects on these traits than those in control and primitive forest treatment. The present results indicated that seed germination and radicle length parameters in control were superior to those in primitive forest treatment, and those of primitive forest treatment were superior to plantation treatment; litter aqueous extracts inhibited the meristematic and elongation zone, and plantation treatment caused a decrease in root hairs so as to be difficult in absorbing water and nutrient in root system. On the other hand, litter aqueous extracts significantly decreased chlorophyll content, net photosynthetic rate and leaf nitrogen concentration of Picea asperata seedlings; MDA, AOS and antioxidant system activities were significantly increased in Picea asperata seedlings. Similarly, plantation treatment had more significant effect on Picea asperata seedlings as compared to primitive forest treatment. In the nature ecosystem, water deficit resulted from elevating air temperature and litter aqueous extract may probably coexist in the cutting blank, forest gap and understory. Our present study showed that water deficit had weaker effects on low light seedlings in the understory as compared to high light seedlings in the cutting blank and forest gap. The fact was confirmed from seedlings growth, gas exchange and biomass accumulation, peroxidation and antioxidant systems. This might be due to that low light-reduced leaf and air temperatures, vapour-pressure deficit, and the oxidative stresses can aggravate the impact of drought under higher light. On the other hand, litter aqueous extracts in the low light had weaker effects on the Picea asperata seedlings than those at high light level, which might be correlative to the water evapotranspiration under high light. Moreover, plantation litter aqueous extracts showed stronger inhibition for seed germination and seedling growth than control and primitive forest treatments.
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由于人类活动所引起的地球大气层中温室气体的富集已导致全球地表平均温度在20世纪升高了0.6 ℃,并预测在本世纪将上升1.4-5.8 ℃。气候变暖对陆地植物和生态系统影响深远,并已成为全球变化研究的重要议题。高海拔、高纬度地带的生态系统对气候变化最敏感。而在高原和高山极端环境影响下所形成的高寒草甸生态系统极其脆弱,对由于温室效应引起的全球气候变化极其敏感,对这些变化的响应更具有超前性。 本研究以川西北高寒草甸植物群落及几种主要物种为研究对象,采用国际山地综合研究中心(ITEX)普遍所采用的增温方法-----开顶式生长室(OTC)模拟气候变暖来研究增温对高寒草甸植物群落结构、物质分配及其主要物种生长和生理的影响,以探讨高寒草甸植物响应与适应气候变暖的生物学和生态学机制。主要研究结论如下: 1、OTC的增温效果 由于地温、地表温度和气温的平均值在OTC内分别高出对照样地0.28℃、0.46℃和1.4℃,这说明本研究所采用的开顶式生长室(OTC)起到了增温的作用;同时,由于温室内与温室外接受的降水量相同,温室内由于热量条件的改善,土壤蒸发和植被的蒸腾作用增强,直接导致了OTC内土壤表层相对湿度的减少。 2、群落结构对增温的响应 由于增温时间较短,增温内外样地的物种组成并未发生改变;但增温后一定程度上改变了植物群落的小气候环境,从而导致物种间的竞争关系被破坏,种间竞争关系的破坏引起群落优势种组成发生相应的改变,在对照样地,鹅绒委陵菜、甘青老鹳草、遏蓝菜和蚤缀是占绝对优势的物种,而在OTC内,小米草、尼泊尔酸模、垂穗披碱草、发草和羊茅的重要性显著增加。 禾草和杂草由于对增温的生物学特性及其资源利用响应的不同,加之增温造成土壤含水量下降等环境因子的改变。与对照样地相比较,OTC内禾草的盖度及生物量都显著增加,而杂草的盖度和生物量则显著下降。 3、植物生长期对增温的响应 OTC内立枯和调落物的生物量在生长季末(10月份)都要小于对照样地的立枯和调落物生物量,而OTC内的地上鲜体生物量在10月份却略高于对照样地。这说明OTC内植物的衰老或死亡得以延缓,而植物的生长期得以延长。 4、群落生物量及分配对增温的响应 OTC内的地上鲜体生物量(10月份除外)和地下0-30cm的根系生物量与对照样地相比较,都出现了不同程度的减少;土壤根系的分配格局也发生了明显的改变,其中,OTC内0-10cm土层的生物量分配比例增加,而20-30cm土层生物量分配比例的减少。 5、群落碳、氮对增温的响应 增温后,OTC内植物群落地上活体和地下活根的碳浓度不同程度的高于对照样地,植物群落的碳库在OTC内也略高于对照样地;而OTC内植物群落地上活体和地下活根的氮浓度不同程度的低于对照样地,其植物群落的氮库与对照样地相比也略有下降。 6、几种主要植物的生长及物质分配对增温的响应 垂穗披碱草在增温后株高、比叶面积和地上生物量均显著地增加;尼泊尔酸模在增温后比叶面积和单株平均生物量积累显著地增加,而各组分中,增温处理使叶的生物量显著增加,而根的生物量却显著下降;鹅绒委陵菜在增温后株高、比叶面积和单株平均生物量积累显著地减少,而各组分中,增温处理使叶和茎的生物量显著减少,根的生物量却显著地增加。 尼泊尔酸模的LMR、RMR、R/S、根部碳含量、碳和氮在叶片与根部的分配比例在增温后显著地增加,而SMR、根部氮含量、碳和氮在茎部的分配比例在增温后却显著地降低;鹅绒委陵菜的RMR、R/S、碳和氮在根部的分配比例在增温后显著地增加,而SMR、LMR、碳在叶片的分配比例在增温后却显著地降低 7、几种主要植物的光合生理过程对增温的响应 增温使垂穗披碱草和尼泊尔酸模叶片中的叶绿素a、叶绿素b、总叶绿素含量显著增加;而鹅绒委陵菜叶片的叶绿素a、叶绿素b、总叶绿素含量在增温后显著减少,类胡萝卜素含量在增温后却显著增加。 增温对3种植物的气体交换产生了显著影响。其中,垂穗披碱草和尼泊尔酸模叶片的光响应曲线在增温后明显高于对照处理,A、E、gs、Pmax、、Rday、AQY和LSP显著增加,而LCP则显著降低;鹅绒委陵菜的光响应曲线在增温后则明显的低于对照处理,A、E、gs、Pmax、、Rday、AQY和LSP显著减少,而LCP则显著增加。 增温后垂穗披碱草和尼泊尔酸模叶片的Fv/Fm、Yield和qP显著增加;而鹅绒委陵菜叶片的Fv/Fm、Yield和qP则显著减少,qN却显著地增加。 8、几种主要植物的抗氧化酶系统对增温的响应 增温使垂穗披碱草和尼泊尔酸模体内抗氧化酶活性和非酶促作用有所提高,植物膜脂过氧化作用降低;鹅绒委陵菜叶片中酶促反应和非酶促反应在增温后也显著提高,但可能由于增温后的土壤干旱超过了鹅绒委陵菜叶的抗氧化保护能力,抗氧化酶活性及非酶促反应(脯氨酸、类胡萝卜素)的提高不足以完全清除干旱诱导形成的过量活性氧,因此叶片的膜脂过氧化程度仍然显著提高。 Enrichment of atmospheric greenhouse gases resulted from human activities such as fossil fuel burning and deforestation has increased global mean temperature by 0.6 ℃ in the 20th century and is predicted to increase in this century by 1.4-5.8 ℃. The global warming will have profound, long-term impacts on terrestrial plants and ecosystems. The ecoologcial consequences arising from global warming have also become the very important issuses of global change research. The terrestrial habitats of high-elevation and high-latitude ecosystems are regarded as the most sensitive to changing climate. The alpine meadow ecosystme, which resulted from the composite effects of mountain extreme climatic factors in Tibetan Plateau, is thus thought to be especially vulnerable and sensitive to global warming. In this paper, the response of plant community and several main species in the alpine meadow of Northewst Sichuan to experimemtal warming was studied by using open-top chambers (OTC). The aim of the this study was to research the warming effects on plant community structure, substance allocation, growth and physiological processes of several mian species, and to explore the biological and ecological mechanism of how the alpine meadow plants acclimate and adapt to future global warming. The results were as follows: 1. Warming effects of OTC The mean soil temperature, soil surface temperature and air temperature in OTC manipulation increased by 0.28℃、0.46℃ and 1.4℃ compared to the control during the growing season. This suggested that the OTC used in our study had increased temperature there. Meanwhile, the OTC manipulation slightly altered thermal conditions, but the same amount of precipitation was supplied to both the OTC manipulation and the control, so higher soil evaporation and plant transpiration in OTC manipulation directly lead to the decrease of soil surface water content. 2. The reponse of community structure to experimental warming The species richness was not changed by the short-term effect of OTC manipulation. However, experimental warming changed the microenvironment of plant community, therefore competitive balances among species were shift, leading to changes in species dominance. In the present study, the dominant plant species in the control plots were some forbs including Potentilla anserine, Geranium pylzowianum, Thlaspi arvense and Arenaria serpyllifolia, however, the importance value of some gramineous grasses including Elymus nutans, Deschampsia caespitosa, Festuca ovina, and some forbs including Euphrasia tatarica and Rumex acetosa significantly increased in OTC. The different biology characteristics and resource utilizations between gramineous grasses and forbs, and enhanced temperature caused change in some environment factors such as soil water content. As a result, the coverage and biomass of gramineous grasses significantly increased in OTC compared to the control, however, the coverage and biomass of forbs singnifciantly decreased in OTC compared to the control. 3. The reponse of plant growing season to experimental warming Both the standing dead and fallen litter biomass in OTC were lower than those in the control in October, and the biomass of aboveground live-vegetation in OTC was higher than that of the control. The results indicated that the senescence of plants was postponed, and the growing season was prolonged in our research. 4. The reponse of community biomass accumulation and its allocation to experimental warming Experimental warming caused the decrease of aboveground live biomass and belowground root biomass except for the aboveground live biomass in October. Experimental warming also had pronounced effects on the pattern of root biomass allocation. In the present study, the root biomass in 0-10cm soil layer increased in OTC manipulation compared to the control, however, the root biomass in the 20-30cm soil layer decreased in OTC manipulation compared to the control. 5. The reponse of community C and N content to experimental warming The C concentration and stock in aboveground live and belowground root both increased in OTC manipulation compared to the control. However, the N concentration and stock in aboveground live and belowground root both decreased in OTC manipulation compared to the control. 6. The reponse of gowth and biomass, C and N alloction of several species to experimental warming Experimental warming significantly increased the height, SLA (specific leaf area) and aboveground biomass of Elymus nutans in OTC manipulation compared to the control. The SLA and total biomass of Rumex acetosa also significantly increased in OTC manipulation compared to control, among the different components of Rumex acetosa, leaf biomass significantly increased, but root biomass significantly decreased in OTC manipulation compared to the control. However, the height, SLA and total biomass of Potentilla anserina significantly decreased in OTC manipulation compared to the control, among the different component of Potentilla anserina, leaf and stem biomass significantly decreased, but root biomass significantly increased in OTC manipulation compared to the control. The LMR (leaf mass ratio), RMR (root mass ratio), R/S (shoot/root biomass ration) and root C concentration of Rumex acetosa significantly increased in OTC manipulation compared to outside control, also, Rumex acetosa allocated relatively more C and N content to leaf and root in response to experimental warming, however, the SMR (stem mass ration) and root N concentration of Rumex acetosa significantly decreased in OTC manipulation compared to outside control, also, Rumex acetosa allocated relatively less C and N content to stem in response to experimental warming. The RMR and R/S of Potentilla anserina significantly increased in OTC manipulation compared to outside control, also, Potentilla anserina allocated relatively more C and N content to root in response to experimental warming, however, the SMR and LMR of Potentilla anserina significantly decreased in OTC manipulation compared to outside control, also, Potentilla anserina allocated relatively less C and N content to leaf in response to experimental warming. 7. The reponse of physiological processes of several species to experimental warming Experimental warming significantly increased chlorophyll a, chlorophyll b and total chlorophyll of Elymus nutans and Rumex acetosa in OTC manipulation compared to outside control. However, chlorophyll a, chlorophyll b, total chlorophyll and carotenoid of Potentilla anserina in OTC manipulation significantly decreased compared to outside control. Experimental warming had pronounced effects on gas exchange of Elymus nutans, Rumex acetosa and Potentilla anserine. In the present study, warming markedly increased the light response curves of Elymus nutans and Rumex acetosa in OTC manipulation compared to outside control, and also singnificantly increased A (net photosynthesis rate), E (transpiration rate), gs (stomatal conductance), Pmax (maximum net photosynthetic rate), Rday (dark respiration rate), AQY (apparent quantum yield) and LSP (light saturation point), but LCP (photosynthetic light compensation) of Elymus nutans and Rumex acetosa in OTC manipulation singnificantly decreased compared to outside control. However, warming markedly decreased the light response curves of Potentilla anserina in OTC manipulation compared to outside control, and also singnificantly decreased A, E, gs, Pmax, Rday, AQY and LSP, but LCP of Potentilla anserina in OTC manipulation singnificantly increased compared to outside control. Experimental warming singnificantly increased the chlorophyll fluorescence kinetics parameters such as Fv/Fm, Yield and qP of Elymus nutans and Rumex acetosa and qN of Potentilla anserina in OTC manipulation, but Fv/Fm, Yield and qP of Potentilla anserina in OTC manipulation singnificantly decreased. 8. The reponse of antioxidative systems of several species to experimental warming Experimental warming tended to increase the activities of antioxidative enzymes and stimulate the role of non-enzymes of Elymus nutans and Rumex acetosa. As a result, MDA content of Elymus nutans and Rumex acetosa decreased. The activities of antioxidative enzymes and non-enzymes of Potentilla anserina also significantly increased in OTC manipulation, but more O2- was produced because of lower soil water content, and the O2- accumulation exceeded the defense ability of antioxidative systems and non-enzymes fuctions. As a result, MDA content of Potentilla anserine still increased in OTC manipulation compared to outside control.
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大气臭氧的损耗导致了地球表面具有生物学效应的紫外线-B(UV-B)辐射增强。同时,大气成分变化中除了UV-B辐射增强外,氮沉降是一个新近出现而又令人担忧的环境问题,其来源和分布正在迅速扩展到全球范围,并不断向陆地和水生生态系统沉降。本试验在四川省境内的中国科学院茂县生态站内进行,以云山、冷杉、色木槭和红椋子幼苗为模式植物,从生长形态、光合作用、抗氧化能力和矿质营养等方面研究了青藏高原东缘4种树苗对全球变化-增强UV-B辐射和氮供应(氮沉降)的响应。该试验为室外盆栽试验,包括四个处理:(1)大气UV-B辐射+无额外的氮供应(C);(2)大气UV-B辐射+额外的氮供应(N);(3)增强UV-B辐射+无额外的氮供应(UV-B);(4)增强UV-B辐射+额外的氮供应(UV-B+N)。其目的:一方面有助于丰富我国对全球变化及区域响应研究的全面认识,进一步完善在全球气候变化条件下臭氧层削减和氮沉降对陆地生态系统影响的内容;另一方面,在一定程度上有助于我们更好的理解在全球变化下森林更新的早期过程。具体结果如下: 增强的UV-B辐射在生长形态、光合、抗氧化能力、活性氧和矿质营养方面对4种幼苗都有显著的影响。UV-B辐射增强对幼苗的影响不仅与物种有关,而且,还与氮营养水平相关。总体表现为,高的UV-B辐射导致了色木槭和红椋子幼苗叶片的皱缩和卷曲,并降低了色木槭幼苗的叶片数和叶重,在额外的氮供应下,云杉、冷杉和红椋子的叶重也显著地降低了;色木槭和红椋子幼苗叶片的解剖结构受到了增强的UV-B辐射的影响,增强的UV-B辐射显著地降低了色木槭叶片的栅栏组织厚度,提高了红椋子叶片的厚度;增强的UV-B辐射显著地降低了4种幼苗的单株总生物量、植物地下部分的生长、总叶绿素含量 [Chl (a + b)]、净光合速率和最大量子产量(Fv/Fm),提高了4种幼苗叶片的膜脂过氧化(MDA含量),改变了植物体不同器官中的矿质元素含量;增强的UV-B辐射提高了冷杉、色木槭和红椋子叶片中的过氧化氢含量(H2O2)、超氧负离子(O2-)生成速率,在额外的氮供应下,云杉叶片中的活性氧含量也显著地提高了;在无额外的氮供应条件下,增强的UV-B辐射显著地提高了4种幼苗叶片中的UV-B吸收物质、脯氨酸含量和抗氧化酶的活性(SOD、POD、CAT、GR和APX)。在额外的氮供应条件下,UV-B辐射的增强却显著地降低了冷杉叶片中脯氨酸含量和红椋子叶片中UV-B吸收物质含量,但是,在4种幼苗叶片中,5种抗氧化酶的活性对UV-B辐射的增强没有明显的规律性,增强的UV-B辐射显著地提高了云杉叶片中的POD、SOD和GR的活性,提高了冷杉叶片中的POD和GR活性,提高了色木槭叶片中的POD、SOD和CAT活性和红椋子幼苗叶片中的POD和SOD活性。从这些结果可知,植物在遭受高的UV-B辐射导致的过氧化胁迫时,植物体内形成了一定的保护机制,但是,这种保护不能抵抗高的UV-B辐射对植物的伤害。 额外的氮供应在生长形态、光合、抗氧化能力、活性氧和矿质营养方面对4种幼苗都有一定的影响,不同幼苗对额外的氮供应响应不同,并且受到UV-B辐射水平的影响。在当地现有的UV-B辐射水平下,额外的氮供应显著地提高了幼苗的单株总生物量、植物地下部分的生长、Chl (a + b)、净光合速率(红椋子除外)、UV-B吸收物质(冷杉除外)、脯氨酸含量(红椋子除外)和部分抗氧化酶的活性,降低了H2O2的含量、O2-的生成速率和MDA含量(红椋子除外),改变了植物体内部分矿质元素含量,显著地提高了云杉和冷杉叶片中的Fv/Fm。这些指标总体表明,在当地现有大气UV-B辐射水平下,额外的氮供应对植物的生长和发育是有利的。在增强的UV-B辐射水平下,4种幼苗的生长形态和光合大部分指标都没有受到额外氮供应的影响,额外的氮供应提高了红椋子幼苗的单株总生物量和Chl (a + b)含量,提高了冷杉和色木槭叶片中的活性氧含量和MDA含量,却降低了红椋子叶片中的活性氧含量;额外的氮供应也提高了云杉、色木槭和红椋子叶片中UV-B吸收物质和脯氨酸含量,降低了冷杉叶片中UV-B吸收物质和脯氨酸含量;在抗氧化酶活性方面,额外的氮供应降低了云杉、冷杉叶片中5种抗氧化酶的活性和红椋子叶片中POD和GR的活性,提高了色木槭叶片中的POD和SOD的活性;4种幼苗植物体内的矿质元素含量对额外的氮供应没有显著的规律性。从这些结果可知,在高的UV-B辐射下,额外的氮供应提高了云杉、冷杉和色木槭幼苗对高的UV-B辐射的敏感性,然而,额外的氮供应却促进了红椋子幼苗的生长,原因可能是,在高的UV-B辐射下,额外的氮供应增加了红椋子叶片的厚度、叶重和叶片数,降低了叶片中活性氧含量的结果。表明在高的UV-B辐射水平下,额外的氮供应降低了红椋子幼苗对高的UV-B辐射的敏感性。 在全球变化的趋势下,UV-B辐射增强和氮沉降可能同时存在,我们的研究表明,与大气UV-B辐射+无额外的氮供应处理相比,增强UV-B辐射+额外的氮供应处理显著地降低了幼苗的单株总生物量(红椋子除外)、Chl (a + b)、净光合速率、Fv/Fm(冷杉除外)和MDA含量(红椋子除外),提高了活性氧含量 (云杉除外)、UV-B紫外吸收物质含量(冷杉除外)、脯氨酸含量和部分抗氧化酶的活性,改变了植物体不同器官中的矿质元素含量。结果表明,在当地现有条件下,全球变化(UV-B辐射增强和氮沉降)对云杉、冷杉和色木槭幼苗的生长是不利,尽管植物体内一些抗氧化性指标提高了,然而,却对红椋子幼苗的单株总生物量的累积没有显著的影响。 The depletion of the ozone led to the increase of ultraviolet-B (UV-B) with biological effects in the earth’s surface. At the same time, except for enhanced UV-B radiation, nitrogen deposition was an anxious environmental problem at present, rapidly expanding to the global scope and continuously depositing to land and aquatic ecosystem. The experiment was conducted in Maoxian Ecological Station of Chinese Academy of Sciences, Sichuan province, China. Picea asperata, Abies faxoniana, Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings were selected as model plants to assess the effects of enhanced UV-B radiation and supplemental nitrogen supply on growth, morphological, photosynthesis, antioxidant and mineral nutrient traits of 4 species seedlings in east Qinghai-Tibetan Plateau. The experiment was potted outdoor, including 4 treatments: (1) ambient UV-B without supplemental nitrogen (control, C); (2) ambient UV-B with supplemental nitrogen (N); (3) enhanced UV-B without supplemental nitrogen (UV-B); (4) enhanced UV-B with supplemental nitrogen (UV-B+N). One hand, it was helpful for enriching our country to comprehensive understanding of the researches in the global change and the region response, further perfecting the effects of the depleted ozone layer and nitrogen deposition on land ecosystem under the global change; the other hand, it was favorable for us to better understanding of the early process of forest renews under the global change. The results were as follows: Enhanced UV-B radiation had significant effects on 4 species seedlings in growth, morphological, photosynthesis, antioxidant and mineral nutrient traits of 4 species seedlings. The effects of enhanced UV-B on plants were not only related with species, but also related with nitrogen nutrient level. Generally, the increase of UV-B radiation led to the shrinkage and curl of leaves in Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, and reduced the number of leaf and leaf weight of Acer mono Maxim seedlings, under supplemental nitrogen supply, leaf weight of Picea asperata, Abies faxoniana and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings significantly also reduced; the anatomical features of leaf in Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings affected by enhanced UV-B radiation, the increase of UV-B radiation markedly reduced the palisade tissue thickness of Acer mono Maxim leaf and enhanced the leaf thickness of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings; the enhanced UV-B radiation significantly reduced total biomass per plant of 4 species seedlings, the growth of the underground parts, Chl (a + b), net photosynthetic rate and maximum potential quantum yield of photosynthesis (Fv/Fm), and increased the degree of lipid peroxidation (MDA content) and changed the content of mineral elements in different parts of plants; the enhanced UV-B radiation also increased the rate of superoxide radical (O2-) production and hydrogen peroxide (H2O2) content in leaves of Abies faxoniana, Acer mono Maxim, Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, under supplemental nitrogen supply, the reactive oxygen species in leaves of Picea asperata seedlings also significantly increased by enhanced UV-B radiation; under without supplemental nitrogen supply, enhanced UV-B radiation evidently induced an increase in UV-B absorbing compounds, proline content and the activities of antioxidant enzymes (SOD, POD, CAT, GR and APX) of leaves in 4 species seedlings. Under supplemental nitrogen supply, enhanced UV-B radiation induced a decrease in proline content of leaves in Abies faxoniana seedlings and UV-B absorbing compounds of leaves in Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, but, there were no obvious rules in the activities of five antioxidant enzymes of 4 species seedling leaves to enhanced UV-B radiation, enhanced UV-B radiation significantly increased the activities of POD, SOD and GR in Picea asperata leaves, the activities of POD and GR in Abies faxoniana leaves and the activities of POD, SOD and CAT in Acer mono Maxim leaves. The results indicated that some protective mechanism was formed when plants were exposed to enhanced UV-B radiation, but the protection could not counteract the harm of high UV-B radiation on plants. Supplemental nitrogen supply had some effects on 4 species seedlings in growth, morphological, photosynthesis, antioxidant and mineral nutrient traits. The response of 4 species seedlings was different to supplemental nitrogen supply, and was affected by UV-B levels. Under local ambient UV-B radiation, supplemental nitrogen supply significantly increased the total biomass per plant, the growth of underground parts, Chl (a + b), net photosynthetic rate (except for Acer mono Maxim seedlings), UV-B absorbing compounds (except for Abies faxoniana seedlings), proline content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings) and the activities of some antioxidant enzymes, and reduced H2O2 content, the rate of O2- production and MDA content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings) and changed the content of mineral elemental in different parts; supplemental nitrogen supply also evidently increased Fv/Fm in Picea asperata and Abies faxoniana seedlings. These results indicated that supplemental nitrogen supply was favorable for the growth of plants under local ambient UV-B radiation. Under enhanced UV-B radiation, mostly parameters in growth and morphology of 4 species seedlings were not affected by supplemental nitrogen supply. Supplemental nitrogen supply increased the total biomass per plant and Chl (a + b) of Swida hemsleyi (Schneid. et Wanger.) Sojak seedling, increased the reactive oxygen species and MDA content in Abies faxoniana and Acer mono Maxim leaves, and reduced the reactive oxygen species in Swida hemsleyi (Schneid. et Wanger.) Sojak leaves; supplemental nitrogen supply also increased UV-B absorbing compounds and proline content in Picea asperata, Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak leaves, decreased UV-B absorbing compounds and proline content in Abies faxoniana leaves; in the activities of antioxidant enzymes, supplemental nitrogen supply significantly reduced the activities of antioxidant enzymes in Picea asperata and Abies faxoniana leaves and the activities of POD and GR in Swida hemsleyi (Schneid. et Wanger.) Sojak leaves, and increased the activities of POD and SOD in Acer mono Maxim leaves; the content of mineral elements in 4 species seedlings was no significantly rule to supplemental nitrogen supply. We knew from the results, under enhanced UV-B radiation, supplemental nitrogen supply made Picea asperata, Acer faxoniana and Acer mono Maxim seedlings more sensitivity to enhanced UV-B radiation, however, accelerated the growth of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings. The reason was probably that supplemental nitrogen supply increased the leaf thickness, leaf weight and leaf number, reduced the reactive oxygen content of leaf in Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings grown under high UV-B radiation. This showed that supplemental nitrogen supply reduced the sensitivity of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings to high UV-B radiation. Under the tendency of the global change, enhanced UV-B radiation and nitrogen deposition may probably coexist. The results showed, compared with the treatment of ambient UV-B radiation without supplemental nitrogen supply, the treatment of enhanced UV-B radiation with supplemental nitrogen supply significantly reduced the total biomass per plants (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings), Chl (a + b), net photosynthetic rate, Fv/Fm and MDA content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings), and increased reactive oxygen content (except for Picea asperata seedlings), UV-B absorbing compounds (except for Abies faxoniana seedlings), proline content and part antioxidant enzymes, and changed the content of mineral elements of different parts. The results indicated that the global change (enhanced UV-B and nitrogen deposition) were not favorable for the growth of plants under local ambient UV-B radiation and nitrogen nutrient level,, though increased some antioxidant indexes, however, the treatment of enhanced UV-B with supplement nitrogen supply did not significantly affect on the biomass accumulation of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings.
Resumo:
本研究通过粗枝云杉不同种群进行的温室半控制试验,采用植物生态学、生理学和生物化学的研究方法,系统地研究了粗枝云杉不同种群抗旱性的生长、形态、生理和生化机理,并结合有关研究进行综合分析,得出主要研究结论如下: 1.粗枝云杉对干旱胁迫的综合反应 粗枝云杉在干旱胁迫下的适应机制为:(1)相对生长速率及植株结构的调整:干旱胁迫下虽然植株相对生长速率显著降低,且有相对较多的生物量向根部分配,但并未发现细根/总根比增加。(2)粗枝云杉对干旱胁迫的光合作用表现为:干旱胁迫显著地降低了控制的理想条件下的气体交换,但干旱胁迫对PSII最大光化学效率(Fv/Fm)没有影响,表明干旱并未影响到光合机构。(3)干旱还影响了很多生理生化过程,包括渗透调解物质(游离脯氨酸)、膜脂过氧化产物、脱落酸(ABA)含量的增加,以及保护酶活性的升高。这些结果证明植物遭受干旱胁迫后发生了一系列的形态、生理和生化响应,这些变化能提高干旱时期植物的存活和生长能力。 2.粗枝云杉不同种群对干旱胁迫反应的种群差异 粗枝云杉三个种群-干旱种群(四川丹巴和甘肃迭部)和湿润种群(四川黑水)对干旱适应不同,这种不同应归因于它们采用的用水策略不同:在水分良好和干旱胁迫条件下,受试种群在相对生长速率和水分利用效率(WUE)方面都表现出显著的种群间差异。与湿润种群相比,干旱种群在两种水分条件下有更高的WUE。粗枝云杉不同种群的碳同位素组分(δ13C)只在干旱胁迫下有显著差异,并且这种差异在水分良好时比干旱胁迫条件下小,说明生理响应和干旱适应性之间的关系受植物内部抗旱机制和外部环境条件(如水分可利用性)或两者互作效应的影响。这些结果说明干旱种群和湿润种群所采用的用水策略不同。干旱种群有更强的抗旱能力,采用的是节水型的用水策略,而湿润种群抗旱能力较弱,采用的是耗水型的用水策略。 3. 遮荫对粗枝云杉不同种群抗旱性影响 干旱胁迫显著降低了全光条件下叶相对含水量(RWC)、相对生长速率、气体交换参数、PSII的有效量子产量(Y),提高了非光化学猝灭效率(qN)、水分利用效率、脯氨酸(PRO)积累、脱落酸(ABA)含量及保护酶活性。然而这种变化在遮荫条件下不明显。我们得出结论适度遮荫降低了干旱对植物的胁迫作用。另一方面,在干旱条件下,与湿润种群相比,干旱种群抗旱性更强,表现在干旱种群净光合速率与单位重量上叶氮含量(Nmass)降低较少。另外,干旱种群表现出更为敏感的气孔导度,更高的热耗散能力(qN)能力、用水效率、ABA积累、保护酶活性,以及更低的总用水量、相对生长速率。这一结果表明这两种群采用不同的生理策略对干旱和遮荫做出反应。许多生长和生理反应差异与这两个种群原产地气候条件相适应。 4. 外源脱落酸(ABA)喷施对粗枝云杉不同种群抗旱性影响 外源ABA喷施在干旱和水分良好条件下均不同程度地提高了根/茎比,表明根和茎对ABA敏感程度不同。实验结果还表明,外源ABA喷施对这两个种群在干旱胁迫期间影响不同。干旱胁迫期间,伴随着ABA喷施,湿润种群净光合速率(A)显著降低,而干旱种群净光合速率变化不明显。另一方面,外源ABA喷施显著提高了干旱条件下干旱种群的单位叶面积重(LMA)、根/茎比、细根/总根(Ft)比、水分利用效率(WUE)、ABA含量, 以及保护酶活性。然而,外源ABA喷施对湿润种群的上述测定指标没有显著影响。这一结果表明干旱种群对外源ABA喷施更为敏感, 反应在更大的气孔导度降低,更高的生物量可塑性,及更高的水分利用效率、ABA含量和保护酶活性。综上所述,我们得出结论,粗枝云杉对外源ABA敏感性因种群的不同而不同。该研究结果可为两个明显不同种群在适应分化方面提供强有力的证据。 Arid or semi-arid land covers more than half of China's land territory. In arid systems, severe shortages of soil water often coincide with periods of high temperatures and high solar radiation, producing multiple stresses on plant performance. Protection from high radiation loads in shaded microenvironments during drought may compensate for a loss of productivity due to reduced irradiance when water is available. Additionally, ABA, a well-known stress-inducible plant hormone, has long been studied as a potential mediator for induction of drought tolerance in plants. Picea asperata Mast., which is one of the most important tree species used for the production of pulp wood and timber, is a prime reforestation species in western China. In this experiment, different population of P. asperata were used as experiment material to study the adaptability to drought stress and population differences in adaptabiliy, and the effects of shade and exogenous abscisic acid (ABA) application on the drought tolerance. Our results cold provide a strong theoretical evidence and scientific direction for the afforestation, and rehabilitation of ecosystem in the arid and semi-arid area, and provide a strong evidence for adaptive differentiation of different populations, and so may be used as criteria for species selection and tree improvement. The results are as follows: 1. A large set of parallel response to drought stress Drought stress caused pronounced inhibition of the growth and increased relatively dry matter allocation into the root; drought stress also caused pronounced inhibition of photosynthesis, while drought showed no effects on the maximal quantum yield of PSII photochemistry (Fv/Fm) in dark-adapted leaves, indicating that drought had no effects on the primary photochemistry of PSII. However, in light-adapted leaves, drought reduced the quantum yield of PSII electron transport (Y) and increased the non-photochemical quenching (qN). Drought also affected many physiological and biochemical processes, including increases in superoxide dismutase (SOD), ascorbate peroxidase (APX) activities, malondialdehyde and ABA content. These results demonstrate that there are a large set of parallel changes in the morphological, physiological and biochemical responses when plants are exposed to drought stress; these changes may enhance the capability of plants to survive and grow during drought periods. 2. Difference in adaptation to drought stress between contrasting populations of Picea asperata There were significant population differences in growth, dry matter allocation and water use efficiency. Compared with the wet climate population (Heishui), the dry climate population (Dan ba and Jiebu) showed higher LMA, fine root/total root ratio and water use efficiency under drought-stressed treatments. The results suggested that there were different water-use strategies between the dry population and the wet population. The dry climate population with higher drought tolerance may employ a conservative water-use strategy, whereas the wet climate population with lower drought tolerance may employ a prodigal water-use strategy. These variations in drought responses may be used as criteria for species selection and tree improvement. 3. The effects of shade on the drought tolerance For both populations tested, drought resulted in lower needle relative water content (RWC), relative growth rate (RGR), gas exchange parameters and effective PSII quantum yield (Y), and higher non-photochemical quenching (qN), water use efficiency (WUE), proline (PRO) and abscisic acid (ABA) accumulation, superoxide dismutase (SOD), ascorbate peroxidase (APX) activities as well as malondialdehyde (MDA) levels and electrolyte leakage in sun plants, whereas these changes were not significant in shade plants. Our study results implied that shade, applied together with drought, ameliorated the detrimental effects of drought. On the other hand, compared with the wet climate population, the dry climate population was more tolerant to drought in the sun treatment, as indicated by less decreases in A and mass-based leaf nitrogen content (Nmass), more responsive stomata, greater capacity for non-radiative dissipation of excitation energy as heat (analysed by qN), and higher WUE,higher level of antioxidant enzyme activities,higher ABA accumulation as well as lower MDA content and electrolyte leakage. Many of the differences in growth and physiological responses reported here are consistent with the climatic differences between the locations of the populations of P. asperata. 4. The effects of exogenous abscisic acid (ABA) application on the drought tolerance For both populations tested, exogenous ABA application increased root/shoot ratio (Rs) under well-watered and drought-stressed conditions, indicating that there was differential sensitivity to ABA in the roots and shoots. However, it appeared that ABA application affected the two P. asperata populations very differently during drought. CO2 assimilation rate (A) was significantly decreased in the wet climate population, but only to a minor extent in the dry climate population following ABA application during soil drying. On the other hand, ABA application significantly decreased stomatal conductance (gs), transpiration rate (E) and malondialdehyde (MDA) content, and significantly increased leaf mass per area (LMA), Rs, fine root/total root ratio (Ft), water use efficiency (WUE), ABA contents, superoxide dismutase (SOD), ascorbate peroxidase (APX) and catalase (CAT) activities under drought condition in the dry climate population, whereas ABA application did not significantly affect these parameters in the wet population plants. The results clearly demonstrated that the dry climate population was more responsive to ABA application than the wet climate population, as indicated by the strong stomata closure and by greater plasticity of LMA and biomass allocation, as well as by higher WUE, ABA content and anti-oxidative capacity to defense against oxidative stress, possibly predominantly by APX. We concluded that sensitivity to exogenous ABA application is population dependent in P. asperata. Our results provide strong evidence for adaptive differentiation between populations of P. asperata.
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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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由于人类活动所引起的地球大气层中温室气体的富集已导致全球地表平均温度在20 世纪升高了0.6 ¡æ,并预测在本世纪将上升1.4-5.8 ¡æ。气候变暖对陆地植物和生态系统产生深远影响,并已成为全球变化研究的重要议题。位于青藏高原东部的川西亚高山针叶林是研究气候变暖对陆地生态系统影响的重要森林类型。森林采伐迹地和人工云杉林下作为目前该区人工造林和森林更新的两种重要生境,二者截然不同的光环境对亚高山针叶林不同物种更新及森林动态有非常重要的影响。 本文以青藏高原东部亚高山针叶林几种主要森林树种为研究对象,采用开顶式增温法(OTCs)模拟气候变暖来研究增温对生长在两种不同光环境下(全光条件和林下低光环境)的几种幼苗早期生长和生理的影响,旨在从更新角度探讨亚高山针叶林生态系统不同树种对气候变暖在形态或生理上的响应差异,其研究结果可在一定程度上为预测气候变暖对亚高山针叶林物种组成和演替动态提供科学依据,同时也可为未来林业生产管理者提供科学指导。 1、与框外对照相比,OTCs 框内微环境发生了一些变化。OTCs 框内与框外对照气温年平均值分别为5.72 ¡æ和5.21 ¡æ,而地表温度年平均值分别为5.34 ¡æ和5.04 ¡æ,OTCs 使气温和地表年平均温度分别提高了0.51 ¡æ和0.34 ¡æ;OTCs框内空气湿度年平均值约高于框外对照,二者分别为90.4 %和85.3 %。 2、增温促进了三种幼苗生长和生物量的积累,但增温效果与幼苗种类及所处的光环境有关。无论在全光或林下低光条件下,增温条件下云杉幼苗株高、地径、分支数、总生物量及组分生物量(根、茎、叶重)都显著地增加;增温仅在全光条件下使红桦幼苗株高、地径、总生物量及组分生物量(根、茎、叶重)等参数显著地增加,而在林下低光条件下增温对幼苗生长和生物量积累的影响效果不明显;冷杉幼苗生长对增温的响应则与红桦幼苗相反,增温仅在林下低光条件下对冷杉幼苗生长和形态的影响才有明显的促进作用。 增温对三种幼苗的生物量分配模式产生了不同的影响,并且这种影响也与幼苗所处的光环境有关。无论在全光或林下低光环境下,增温都促使云杉幼苗将更多的生物量分配到植物地下部分,从而导致幼苗在增温条件下有更高的R/S 比;增温仅在林下低光条件下促使冷杉幼苗将更多的生物量投入到植物叶部,从而使幼苗R/S 比显著地降低;增温在全光条件下对红桦幼苗生物量分配的影响趋势与冷杉幼苗在低光条件下相似,即增温在全光条件下促使红桦幼苗分配更多的生物量到植物同化部分—叶部。 3、增温对亚高山针叶林生态系统中三种幼苗气体交换和生理表现的影响总体表现为正效应(Positive),即增温促进了几种幼苗的生理活动及其表现:(i)无论在全光或林下低光环境下,增温使三种幼苗的光合色素含量都有所增加;(ii)增温在一定程度上提高了三种使幼苗的PSII 光系统效率(Fv/Fm),从而使幼苗具有更强的光合电子传递活性;增温在一定程度使三种幼苗潜在的热耗散能力(NPQ)都有所增强,从而提高幼苗防御光氧化的能力;(iii)从研究结果来看,增温通过增加光合色素含量和表观量子效率等参数而促进幼苗的光合作用过程。总体来说增温对幼苗生理过程的影响效果与幼苗种类及所处的光环境有关,增温仅在全光条件下对红桦幼苗光合过程的影响才有明显的效果,而冷杉幼苗则相反,增温仅在低光条件下才对幼苗的生理过程有显著的影响。 4、增温对三种幼苗的抗氧化酶系统产生了一定的影响。从总体来说,增温使几种幼苗活性氧含量及膜脂过氧化作用降低,从而在一定程度上减轻了该区低温对植物生长的消极影响;增温倾向表明使三种幼苗体内抗氧化酶活性和非酶促作用有所提高,从而有利于维持活性氧代谢平衡。但增温影响效果与幼苗种类所处的光环境及抗氧化酶种类有关,增温对冷杉幼苗抗氧化酶活性的影响仅在林下低光环境下效果明显,而对红桦幼苗抗氧化酶活性的影响仅在全光条件下才有明显的效果。 总之,增温促进了亚高山针叶林生态系统中三种幼苗的生长和生理表现,但幼苗生长和生理对增温的响应随植物种类及所处的光环境不同而变化,这种响应差可能异赋予了不同植物种类在未来气候变暖背景下面对不同环境条件时具有不同的适应力和竞争优势,从而对亚高山针叶林生态系统物种组成和森林动态产生潜在的影响。 Enrichment of atmospheric greenhouse gases resulted from human activities suchas fossil fuel burning and deforestation has increased global mean temperature by 0.6¡æ in the 20th century and is predicted to increase it by 1.4-5.8 ¡æ. The globalwarming will have profound, long-term impacts on terrestrial plants and ecosystems.The ecoologcial consequences arising from global warming have also become thevery important issuses of global change research. The subalpine coniferous forests inthe eastern Qinghai-Tibet Plateau provide a natural laboratory for the studying theeffects of climate warming on terrestrial ecosystems. The light environment differssignificantly between clear-outs and spruce plantations, which is particularlyimportant for plant regeneration and forest dynamics in the subalpine coniferous forests. In this paper, the short-term effects of two levels of air temperature (ambient andwarmed) and light (full light and ca. 10% of full light regimes) on the early growthand physiology of Picea asperata, Abies faxoniana and Betula albo-sinensis seedlingswas determined using open-top chambers (OTCs). The aim of the present study wasto understand the differences between tree species in their responses to experimentalwarming from the perspective of regeneration. Our results could provide insights intothe effects of climate warming on community composition and regeneration behavior for the subalpine coniferous forest ecosystem processes, and provide scientificdirection for the production and management under future climate change. 1. The OTCs manipulation slightly altered thermal conditions during the growingseason compared with the outside chambers. The annual mean air temperature andsoil surface temperature was 5.72 and 5.34 ¡æ (within the chambers), and 5.21 and5.04 ¡æ (outside the chambers), respectively. The OTCs manipulation increased airtemperature and soil surface temperature by 0.51 and 0.34 ¡æ on average, respectively.Air relative humidity was slightly higher inside the OTCs compared with the controlplots, with 90.4 and 85.3 %, respectively. 2. Warming generally stimulated the growth and biomass accumulation of thethree tree species, but the effects of warming on growth and development variedbetween light conditions and species. Irrespective of light regimes, warmingsignificantly increased plant height, root collar diameter, total biomass, componentbiomass (stem, foliar and root biomass) and the number of branches in P. asperataseedlings; For A. faxoniana seedlings, significant effects of warming on all the tested parameters (plant height, root collar diameter, total biomass, and component biomass) were found only under low light conditions; In contrast, the growth responses of B.albo-sinensis seedlings to warming were found only under full light conditions. Warming had pronounced effects on the pattern of carbon allocation. Irrespectiveof light regimes, the P. asperata seedlings allocated relatively more biomass to rootsin responses to warming, which led to a higher R/S. Significant effects of warming onbiomass allocation were only found for the A. faxoniana seedlings grown under lowlight conditions, with significantly increased in leaf mass ratio (LMR) and decreasedin R/S in responses to warming manipulation. The carbon allocation responses of B.albo-sinensis seedling to warming under full light conditions were similar with theresponse of A. faxoniana seedlings grown under low light conditions. Warmingsignificantly decreased root mass ratio (RMR), and increased leaf mass ratio (LMR)and shoot/root biomass ratio (S/R) for the B. albo-sinensis seedlings grown under full light conditions. 3. Warming generally had a beneficial effect on physiological processes of dominant tree species in subalpine coniferous forest ecosystems: (i) Warming markedincreased the concentrations of photosynthetic pigments in both tree species, but theeffects of warming on photosynthetic pigments were greater under low lightconditions than under full light conditions for the two conifers; (ii) Warming tended toenhance the efficiency of PSII in terms of increase in Fv/Fm, which was related tohigher chloroplast electron transport activity; and enhance non-radiative energydissipation in terms of in increase in NPQ, which may reflect an increased capacity inpreventing photooxidation; (iii) Warming may enhance photosynthesis and advancephysiological activity in plants by increasing photosynthetic pigment concentration,the efficiency of PSII and apparent quantum yield (Φ) etc. From the results, theeffects of warming on seedlings’ physiological performance varied between lightenvironment and species. The effects of warming on photosynthesis performance of B.albo-sinesis seedlings were pronounced only under full light conditions, while thephysiological responses of A. faxoniana seedlings to warming were found only underthe 60-year plantation. These results provided further support for the observationsabove on growth responses of seedlings to warming. 4. Warming had marked effects on antioxidative systems of the three seedlings.Warming generally decreased H2O2 accumulation and the rate of O2- production, andalleviated degree of lipid peroxidation in terms of decreased MDA content, whichalleviated to some extent the negative effects of low temperature on the plant growthand development in this region; Warming tended to increase the activities ofantioxidative enzymes and stimulate the role of non-enzymatic AOS scavenging,which helped to create an balance in maintaining AOS metabolites for the threeseedlings. Nevertheless, the effects of warming on antioxidative defense systems werepronounced only under the 60-year plantation for the A. faxoniana seedlings. Incontrast, the marked effects of warming on antioxidative defense systems for the B.albo-sinesis seedlings were found only under the full light conditions. In sum, warming is considered to be generally positive in terms of growth andphysiological process. However, the responses of growth and physiology performanceto warming manipulation varied between species and light regimes. Competitive and adaptive relationships between tree species may be altered as a result of responsedifferences to warming manipulation, which is one mechanism by which globalwarming will alter species composition and forest dynamics of subalpine coniferousforest ecosystems under future climate change.