86 resultados para Diameter at breast height


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随着社会信息化程度的不断提高,各行各业对远程数字视频监控的应用日益广泛。越来越多的视频数据要求进行实时和无地域性障碍的传输,以求实现更好的资源共享。认证、授权和计费服务在视频监控系统和其他应用系统中都担负了保障系统安全、实体可信赖和费用有效的责任,是网络运营管理中非常重要的部分。 本文以下一代互联网中的视频监控应用需求为背景,介绍了视频监控技术的发展和AAA(Authentication、Authorization、Accounting)服务内容,提出了基于Diameter协议实现AAA服务的方案,设计了支持跨域访问、实时计费、支持IPv4/IPv6双栈的AAA服务系统,并完成了其在视频监控应用中的实现,同时还实现了系统管理和网上营业厅功能的业务支撑系统,测试和验证了AAA服务系统和业务支撑系统的正常工作。 本文工作主要有如下贡献: 1. 参与设计了视频监控系统的整体框架和子系统间接口; 2. 提出了以Diameter协议为基础实现视频监控系统中AAA服务的方案; 3. 设计了视频监控系统中的AAA子系统框架,同时设计了跨域服务方案和实时计费方案; 4. 实现了视频监控系统中的AAA子系统,提供了认证、授权、实时计费、跨域访问等功能,完成了Diameter客户端、代理和服务器实体,测试和验证了AAA服务子系统运行效果; 5. 设计并实现了视频监控系统中的业务支撑子系统,提供了系统管理和网上营业厅功能,展示了业务支撑子系统运行效果。

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Particle velocity distribution in a blowing sand cloud is a reflection of saltation movement of many particles. Numerical analysis is performed for particle velocity distribution with a discrete particle model. The probability distributions of resultant particle velocity in the impact-entrainment process, particle horizontal and vertical velocities at different heights and the vertical velocity of ascending particles are analyzed. The probability distributions of resultant impact and lift-off velocities of saltating particles can be expressed by a log-normal function, and that of impact angle comply with an exponential function. The probability distribution of particle horizontal and vertical velocities at different heights shows a typical single-peak pattern. In the lower part of saltation layer, the particle horizontal velocity distribution is positively skewed. Further analysis shows that the probability density function of the vertical velocity of ascending particles is similar to the right-hand part of a normal distribution function, and a general equation is acquired for the probability density function of non-dimensional vertical velocity of ascending particles which is independent of diameter of saltating particles, wind strength and height. These distributions in the present numerical analysis are consistent with reported experimental results. The present investigation is important for understanding the saltation state in wind-blown sand movement. (C) 2009 Elsevier B.V. All rights reserved.

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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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大气CO2浓度的增加已经成为不可争议的事实。预计本世纪末大气CO2浓度将增加到约700µmol mol-1。森林年光合产量约占陆地生态系统年光合产量的70%。森林树木是一个巨大的生物碳库,约占全球陆地生物碳库的85%。森林树木对CO2的固定潜力是缓解由大气CO2浓度升高引起的未来全球气候变化问题的决定性因子之一。红桦(Betula albosinensis Burk.)是川西亚高山采伐迹地自然或人工恢复的重要树种。本研究以1a红桦幼苗为模式植物,采用人工模拟的方法,研究CO2浓度升高对不同种内竞争强度(种群水平)下红桦幼苗的生理特征、生长、干物质积累及其分配的影响,探讨在种内竞争生长条件下红桦幼苗的“光合适应机理”与生长特征,为西南亚高山森林生产力对未来全球变化的预测提供重要参考。 本研究的主要结果如下: 1)在种内竞争生长条件下红桦幼苗经过CO2浓度升高熏蒸4个月后,叶片出现“光合适应”现象。与对照相比,低种植密度(28株m-2)和高种植密度(84株m-2)条件下的红桦幼苗净光合速率(A)、气孔导度(gs)、蒸腾速率(E)、表观量子产量(AQY)和羧化速率(CE)显著降低,而水分利用效率(WUE)则显著提高。CO2浓度升高处理的红桦幼苗叶片Rubisco活性、单位叶面积N浓度、叶绿素a、叶绿素b和类胡萝卜素浓度都显著降低。但CO2浓度对红桦幼苗的叶绿素a与叶绿素b的比值没有显著影响。CO2浓度升高显著增加红桦幼苗单位叶面积的非结构性碳水化合物(TNC)浓度,结果是红桦幼苗的比叶面积(SLA,cm2 g-1)显著降低。 2)与对照相比,CO2浓度升高处理的红桦幼苗高、基径、单叶面积和侧枝的相对生长速率(R GR)显著提高,尤其在试验处理的早期。CO2浓度升高既增加单株红桦幼苗总叶片数量又增加单叶面积,结果是单株红桦幼苗的总叶面积比对照显著增加。 3)CO2浓度升高处理显著增加红桦幼苗干物质积累(尤其是细根生物量),改变了红桦幼苗生物量的分配格局。与对照相比,CO2浓度升高处理的红桦幼苗叶重比(LWR)、叶面积比(LAR)、叶根重比(Wl/Wr)和源汇重比(leaf weight to non-leaf weight ratio, Wsource/Wsink)显著下降(高种植密度的LWR除外),而根冠比(R/S)则显著增加。在两种种植密度条件下,CO2浓度升高显著增加红桦幼苗根生物量的分配比率,显著降低叶片的生物量分配比率,对主茎、侧枝以及地上生物量的分配比率不变或约有下降。 总之,长期生长在CO2浓度升高条件下的红桦幼苗光合能力下降,并伴随Rubisco活性、叶N浓度、光合色素浓度的显著降低以及TNC浓度的显著增加。支持树木光合速率下降与Rubisco活性、叶N浓度下降以及TNC浓度增加紧密相关的假设。CO2浓度升高处理红桦幼苗的早期相对生长速率大大高于对照,而后期迅速下降,说明红桦幼苗生物量的显著增加主要归功于CO2浓度升高的早期促进作用和叶面积的显著增加。CO2浓度升高显著增加红桦幼苗根系生物量和根冠比,表明红桦幼苗“额外”固定的C向根系转移。 The steady increae of atmospheric CO2 concentration([CO2])has been inevitable fact. Models predict that the atmospheric [CO2] will increase to about 700µmol mol-1 at the end of the twenty-first century. As trees constitute a majoor carbon reservoir–85% of total plant carbon is found in forest, and their ability to sequester carbon is a key determinant of future global change problems caused by increases in atmospheric CO2. In addition to the role of forests in the global carbon cycle, inceased growth could be of economic benefit, for example, offsetting deleterious effects of climatic changes. Betula albosinensis (Burk.) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of mountain forest area, and also is one of important tree species for afforestation in logged area, in southwesten China. In this experinment, Betula albosinensis seedling (one-year-old) was used as the model plant. B. albosinensis seedlings were grown under two all-day [CO2], ambient (about 350 µmol·mol-1) and elevated [CO2] (about 700 µmol·mol-1), and two planting densities of 28 plants per m2 and 84 plants per m2. The objectives were to characterize birch mature leaf photosynthesis, growth, mass accumulation and allocation responses to long-tern elevated growth [CO2] under the influences of neighbouring plants, and to assess whether elevated [CO2] regulated birch mature leaf photosynthetic capacity, in terms of leaf nitrogen concentration (leaf [N]), activity of ribulose bisphosphate carboxygenase (Rubisco), Rubisco photosynthetic efficiency, and total nonstructural carbohydrates (TNC) concentration, and also to provide a strong reference to predict the productivity of subalpine forests under the future global changes. The results are as follows: 1) B.albosinensis seedlings exposed to elevated [CO2] for 120 days, photosynthetic acclimation phenomena occurred. At two planting densities, leaves of birch seedlings grown under elevated [CO2] had lower net photosynthetic rate (A), stomatal conductance (gs), transpiration (E), apparent quantum yield (AQY) and carboxylated efficiency (CE) and higher water use efficiency (WUE), compared to those of B.albosinensis seedlings grown under ambient [CO2]. Based on the leaf area, leaf [N], Rubisco activity and photosynthetic pigments concentrations of B. albosinensis seedlings grown under elevated [CO2] were significantly lower than those grown under ambient [CO2]. The ratio of chlorophyll a to chlorophyll b concentration was not affected by elevated [CO2]. Under elevated [CO2], the TNC concentration per unit leaf area significantly increased, resulting in significant decrease in specific leaf area. Thus leaf photosynthetic capacity of B. albosinensis seedlings would perform worse under rising atmospheric [CO2] and the influences of neighbouring plants. 2) Under elevated [CO2], the relative growth rate (RGR) of B. albosinensis seedlings height, basal diameter, a leaf area and branch length significantly increased, especially at the initial stage of exposure to elevated [CO2], and a leaf area and leaf numbers per B. albosinensis seedling also significantly increased. Thus the total leaf area per B. albosinensis seedling was significantly increased under elevated [CO2]. 3) As the increase of RGR and total leaf area, biomass of B. albosinensis seedling grown elevated [CO2] was higher, compared to that of B.albosinensis seedlings grown at ambient [CO2]. Elevated [CO2] changed the biomass allocation pattern of B. albosinensis seedling. At two planting densities, B. albosinensis seedlings grown elevated [CO2] had lower leaf weight to total weight ratio (LWR), leaf area to total weight ratio (LAR) and leaf weight to non-leaf weight ratio (Wsource/Wsink), but higher root weight to shoot weight ratio (R/S), compared to those of B.albosinensis seedlings grown at ambient [CO2]. Under elevated [CO2], roots biomass to total biomass ratio was signigicantly increased, leaves biomass to total biomass ratio was significantly decreased. The main stem and branch biomass to total biomass ratio were not affected by elevated [CO2]. In conclusion, our results supported the hypothesis that the decline in photosynthetic capacity of C3 plants will appear after long-term exposure to elevated [CO2], accompanying with the significant decrease in Rubisco activity, leaf N concentration, photosynthetic pigments concentration, and significant increase in total non-structural carbohydrates concentration. Our results also have shown that the increase of biomass of B. albosinensis seedlings should be attributed to initial stimulation on RGR and total leaf area resulted from elevated [CO2]. Under elevated [CO2], the extra carbon sequestered by B.albosinensis seedlings transferred into under-ground part because of increase in root biomass and R/S.

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当前大气CO2浓度升高是全球变化的主要趋势之一,CO2浓度升高还会引起全球变暖等其它环境问题,因而CO2浓度浓度升高对植物影响的研究已经成为全球变化领域的焦点。红桦是川西亚高山地区暗针叶林演替初期的先锋树种和演替后期的建群种,在群落演替过程中它对环境因子的响应决定红桦群落的演替进程。本文通过控制CO2浓度的气候室试验,研究了CO2浓度倍增环境下,不同密度水平红桦碳氮固定、分配可能发生的改变,并探讨了升高大气CO2浓度对群体内部竞争的影响。以期通过本研究明确川西亚高山地区代表性物种红桦对未来气候变化的响应,为今后采取措施应对气候变化、妥善进行森林管理提供理论依据和科学指导。主要研究结果如下: 1.升高CO2浓度对红桦幼苗生长的影响以及树皮、树干响应的不同 (1) CO2浓度升高显著促进红桦幼苗的生物量、株高、基茎的生长,同时也改变生物量在体内的分配格局,主要是增加根和主茎、减少叶在总生物量中的比重。(2)树皮和树干对升高CO2浓度的影响有差异,它们对CO2浓度升高的反应程度不同,但反应方向一致。 2.密度的副效应 (1) 增加种植密度对单株生物量、株高和基径的生长具有副效应,也降低升高CO2浓度对红桦生长的正效应。(2) 增加种植密度,显著增加红桦幼苗的群体生物量,从而使红桦群体固定更多的大气CO2气体。可见密度在决定红桦生物量及固碳能力方面具有重要意义。探索适合未来大气CO2浓度升高条件下植物生长的密度,对未来的森林经济生产、生态恢复具有重要意义。 3. 升高CO2浓度对红桦幼苗苗冠结构及冠层内部竞争的影响 (1) 冠幅、冠高、苗冠表面积和苗冠体积等树冠特征均受CO2浓度升高的影响而增加,但是受密度增加的影响而降低。(2) 单位苗冠投影面积叶片数(LDcpa)和单位苗冠体积叶片数(LDcv)均低于相应的现行CO2浓度处理,这主要是由于冠幅和冠高的快速生长所造成的。(3) LDcpa和LDcv的降低表明,红桦在升高CO2浓度的条件下,会作出积极的响应,从而缓解由于群体和个体生长的增加所引起的竞争压力的增加。 4. 升高CO2浓度对红桦幼苗养分元素吸收与分配的影响 (1) CO2浓度升高,植株各器官N、P含量降低,但单株N、P总吸收量均增加。红桦幼苗体内N、P浓度的下降是由于生物量迅速增加引起的稀释效应造成的。(2) CO2浓度升高,N、P向主茎和根的分配增加,向叶片的分配减少,主要是由于前者在总生物量中的比重增加,而后者减少了。(3) CO2浓度升高,氮磷利用效率(NUE和PUE)提高,氮磷累积速率(NAcR和PAcR)显著增加。而NUE和PUE的提高可以有效缓解CO2浓度升高后,亚高山和高山地区森林土壤中养分元素不足对森林生产力的限制。 5. 升高CO2浓度对红桦幼苗群体碳平衡的影响 (1) 升高CO2浓度对植物的光合作用、呼吸速率和生长均具有促进作用。(2) 土壤有机碳含量在实验前期迅速增加,后期积累速率下降。(3) 升高CO2浓度以后,土壤呼吸显著增强;土壤呼吸还具有明显的季节变化。(4) 红桦群体日固碳量受到升高CO2浓度的促进作用。结果(1)-(4)说明所研究群落的碳动态对现行的气候波动是敏感的;所研究群落在作为大气CO2气体的源-汇关系方面至少存在季节间的源汇飘移。(5)种植密度的升高显著增加了群体固碳量。 6. 升高CO2浓度对红桦幼苗生长后期叶片衰老的影响 升高CO2浓度有利于减缓红桦幼苗叶片生长季节末期的衰老。生长季节末期,随着CO2浓度的升高光合速率和可溶性蛋白含量均呈上升趋势,同时MDA(丙二醛)含量下降,保护酶SOD(超氧化物岐化酶)、CAT(过氧化氢酶)活性升高。由此说明,升高CO2浓度有利于减缓生长季节后期叶片的衰老,使叶片维持较高的光合速率,也从生理学的角度支持了本文及前人有关CO2浓度升高促进植物光合和生长的假说及结果。 The increased CO2 concentration is one of the most important problems among global changes. The increase of CO2 will also cause other environmental problems, such as global warming, etc. So the effects of elevated CO2 on plant have drawn sights of many scientists in the research field of global change. Red birch (Betula albosinensis) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of the dark coniferous forests in Western Sichuan, China. It’s response to elevated CO2 may determine the succession process of the community where it lives in. By controlling CO2 at the ambient and twice as the ambient level (ambient + 350 umol mol-1) using enclosed-top chambers (ETC), possible effects of elevated CO2 on carbon fixation and allocation under two plantation densities are investigated. The effects of elevated CO2 on competition within canopy of red birch seedlings are also observed in the present paper. We hope to make sure of the effects of elevated CO2 on the representative species, red birch. And so that, our results could provide a strong theoretical evidence and scientific direction for forest management and afforestation under a future, CO2 elevated world. The results are as fowllows: 1. The effects of elevated CO2 on growth and the different responses of wood and bark of red birch seedlings (1) Elevated CO2 increases the growth of seedling biomass, seedling height and basal diameter of red birch. It also changed the biomass allocation in red birch seedlings. The ratio of root and main stem to all biomass is increased and the ratio of leaf is decreased. (2) Tree bark and wood show different response degree but similar response direction to elevated CO2. 2. Negative effects of planting density (1) The increase of planting density showes negative effects on the individual growth of seedling biomass, seedling height and basal diameter of red birch. It also eliminates the positive effects of elevated CO2 on growth of red birch seedlings. (2) Community biomass is increased by the elevated planting density, which means that the high density red birch community could fix more CO2 than the low density one. These results show that planting density plays an important role in determining biomass and carbon fixation ability of red birch community. Thus, exploring proper planting density becomes economically important for the future, CO2 elevated word. 3. The effects of elevated CO2 on crown architecture and competition within canopy of red birch seedlings (1) Crown width, crown depth, crown surface area and crown volume are all increased under the influence of elevated CO2. (2) Leaf number per unit area of projected crown area (LDcpa) and per unit volume of crown volume (LDcv) are lower under elevated CO2. This is resulted from the stimulated growth of tree crown features. (3) The decrease of LDcpa and LDcv indicate that plants will respond forwardly to reduce the possible increase of competition resulted from stimulated growth of individual plant and collectives in conditions of elevated CO2. 4. The effects of elevated CO2 on nutrition accumulation and allocation of red birch seedlings (1) Contents of N and P decrease due to the prompt increase of biomass of plant organs caused by elevated CO2. However, their accumulations increase under elevated CO2. (2) Elevated CO2 increases the allocation of N, P to main stem but reduced its allocation to leaf for that dry weight of the former increased but the dry weight of the later decreased. (3) Using efficiencies of N, P (NUE and PUE) and their accumulation rates (NAcR and PAcR) are found to increase under elevated CO2. Soil nutrition contents are always the limiting factors for plant growth at subalpine and alpine region. The increased NUE and PUE are helpful to eliminate the nutrition limitation in this area in the future world, when CO2 concentration doubles the ambient. 5. The effects of elevated CO2 on carbon balance of red birch communities (1) Net photosynthetic rates (Pn), dark respiration rates (Rd) and growth are all stimulated by elevated CO2. (2) Content soil organic carbon increases sharply at the primary stage of experiments and then the increasing rates decrease to a low level at later stages. (3) Soil respiration rates increase significantly with the elevation of CO2 concentration. (4) The daily carbon fixations of whole community are heightened by elevated CO2. The results (1)-(4) suggest that, the community being studied are sensitive to current climate change; the studied community, as a sink of atmospheric CO2, is pool-sink alternative between seasons. (5) The carbon fixations are increased along the increase of planting densities. 6. The effects of elevated CO2 on physiological features of leaf senescences of red birch seedlings at the later stage of growing season Elevated CO2 helps to postpone the leaf senescences of red birch at the end of the growth season. CO2 enrichment increases the photosynthetic rates, contents of soluble proteins and photosynthetic pigments. And meanwhile contents of malondialdehyde (MDA) decreases and activities of superoxide dismutase (SOD) and catalase (CAT) are both increased. These results suggest that the senescences of red birch leaves are delayed by elevated CO2, which keep the photosynthetic rates at relatively high levels. Our results lend supports to hypothesis and results on stimulated photosynthetic rates and growth from both other researchers and the present paper.

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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.

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目前,随着全球气候变化的加剧,水分短缺更加明显。在干旱与半干旱地区,水分胁迫是影响植物存活和生长的主要限制因子。同时,随着大气平流层中臭氧浓度的减少,过量的紫外辐射(UV-B)到达地球表面,一些地区的植物不可避免地受到增强UV-B 和水分胁迫的共同作用。文献表明在UV-B 增强的情况下,干旱表现为减弱或增强UV-B 对植物的影响,这与种、品种有一定的相关性。另外,脱落酸(ABA)是近年来研究报道最多的信息调控物质,与植物抗旱性途径有较大的关系,但其对植株抗UV-B 的影响还有待于研究。本论文以滇杨(Populus yunnanensis)为模式植物,从形态和生理方面研究了增强UV-B、干旱和脱落酸对它的影响,并探讨了UV-B 与干旱的互作效应以及喷施脱落酸对植株抗旱性和抗UV-B 能力的影响。主要研究结果如下:1. 增强的UV-B 和干旱胁迫都影响了滇杨的形态生长和生理生化反应。它们都导致了滇杨植株的株高、基茎、整株叶面积、平均叶面积、总生物量和净光合速率的显著降低,使得叶片增厚,过氧化物酶(GPX)活性升高,脯氨酸和花色素苷含量增加,膜脂过氧化程度增大。不同的是干旱显著降低了植株叶片数目,增大了根/冠比(Rs)、细根/总根比(Ft)、提高了内源ABA 含量、碳同位素(δ13C)以及紫外吸收物质含量和超氧化物歧化酶(SOD)的活性,而UV-B 对它们没有影响。干旱与UV-B 的复合作用加剧了任一单独胁迫对植株的抑制,表现为更小的株高、基茎、整株叶面积、平均叶面积、总生物量,更低的光合作用和更高的MDA 含量。而且UV-B 辐射降低了干旱胁迫下生物量分配的可塑性,表现为降低了干旱情况下的Rs 和Ft,ABA 的含量也显著下降,复合胁迫下脯氨酸含量和过氧化氢酶(CAT)的活性比任一单独胁迫时都要低。这些实验结果表明,增强的UV-B 与干旱的复合胁迫加剧了对植株的抑制作用。II2. 干旱情况下同时施加外源ABA 提高了植株的根/冠比、细根/总根比和单位面积叶重,即提高了干旱胁迫下植株对生物量分配的可塑性。而且外源ABA 使干旱胁迫下的长期用水效率、ABA 含量、脯氨酸含量、GPX 活性进一步增加,并有效调节了活性氧代谢的平衡,抑制了受旱植株MDA 的增加。结果表明,外源ABA 的喷施提高了滇杨植株的抗旱性。3. 在增强的UV-B 情况下,外源ABA 加剧了UV-B 对滇杨形态生长的抑制效果,表现为进一步降低了滇杨植株的整株叶面积、平均叶面积、单位面积叶重和总生物量,而且ABA 还降低了UV-B 胁迫下的净光合速率和脯氨酸的含量,增大了MDA 的含量。通过以上的数据我们可以看出,外源ABA 虽然提高了滇杨植株的抗旱性,但却加剧了UV-B 胁迫对植株的抑制作用。Currently, drought is one of the most serious environmental stresses. In arid and semi-aridregions, drought is a major constraint imposed on tree survival and growth. The decrease ofozone layer leads to a significant increase in ultraviolet-B (UV-B, 280-320 nm) radiationreaching the earth surface. In some places, plants suffer both UV-B and water stresssimultaneously. Their combination will increase or decrease the sensitivity of plants to UV-Bstress which lies on the species. On the other hand, abscisic acid (ABA), as a plant homoneand growth regulator, is better for plants resistant to drought stress, but it is uncleared aboutthe relationship between exogenous ABA and supplemental UV-B. In the present study, weemployed Populus yunnanensis Dode as a model species to characterize the growth andecophysiological responses of woody plants to supplemental UV-B, drought and exogenous ABA. The results are as follows:1. Both supplemental UV-B and drought affected the morphological, physiological andbiochemical responses of P. yunnanensis. They decreased the plant height, basal diameter,total leaf area, average leaf area, biomass and photosynthesis, and increased specific leaf mass,the activity of guaiacol peroxidase (GPX), the content of proline, anthocyanins andmalondialdehyde (MDA). However, drought decreased the leaf number and increasedroot/shoot ratio, fine root/total ratio, the activity of superoxide dimutase (SOD) and thecontents of ABA, carbon isotope composition (δ13C), UV-absorbing compounds. Whilesupplemental UV-B had no effects on them. The combination of drought and UV-Baugmented the growth inhibtion acting as further lower plant height and smaller basaldiameter, leaf area, biomass and higher MDA content. And compared with drought stress,root/shoot ratio and fine root/total root ratio decreased under the combination stresses. The photosynthesis, proline content and Catalase (CAT) activity became lower under combinationstresses than that of either stress lonely. According to these results, we suggested that,compared with the effect of stress lonely, the combination of supplemental UV-B and droughtdid not mitigate the harmful effect, but augmented it.2. Under drought conditions, exogenous ABA increased root/shoot ratio, fine root/total rootratio and the specific leaf mass. That was to say exogenous ABA increased plant plasticityunder drought conditions. Also ABA content, proline content, activity of GPX and δ13C wereenhanced further. In addition the enhancement of MDA was restrained. So the resultssuggested that exogenous ABA increased the seedling capacity of resistance to drought.3. Under supplemental UV-B conditions, exogenous ABA augmented the growth restrain ofUV-B to seedlings, which acted as further decreased leaf area, specific leaf mass and biomass.Compared with UV-B stress alone, proline content and photosynthesis were decreased andMDA content was increased under the combination of UV-B and ABA. These resultssuggested that although exogenous ABA increased the seedling capacity of resistance todrought, it augmented the growth restrain of supplemental UV-B to P. yunnanensis.

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杨树具有分布广、适应性强的特征,在生态环境治理和解决木材短缺方面均占有重要位置。青杨(Populus cathayana Rehd.)是青杨派树种的重要成员之一,也是生长较迅速、易繁殖的重要杨树资源。本研究选取了来自不同气候地区的青杨两种群为材料,采用植物生态学、生理学和生物化学的研究方法,系统地研究了青杨对干旱与遮荫、干旱与外源脱落酸(ABA)喷施的生长、形态、生理和生化响应及种群间差异,研究成果可为我国干旱半干旱地区的造林以及生态恢复提供理论依据和科学指导。主要研究结论如下:1.青杨在干旱胁迫下的适应机制为:生长性状及生物量的分配变化:干旱胁迫下虽然植株生长受抑,株高、基茎及各部分生物量都显著减小,但有相对较多的生物量向根部分配,根/冠比以及细/粗根比增加。青杨对干旱胁迫的光合作用表现为:干旱胁迫降低了青杨的净光合速率、蒸腾速率、气孔导度以及光合氮利用效率,提高了瞬时用水效率。干旱还引起了活性氧的产生,使得膜脂过氧化产物丙二醛(MDA)增加,同时也增强了植物抗氧化酶系统(如超氧化物歧化酶(SOD)、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)活性的增加)及非酶系统的能力(如抗坏血酸(AsA)含量的增加)。干旱降低了植物叶片的相对含水量,而促进了渗透调节物质(游离脯氨酸及可溶性糖)的积累,增加了植物的渗调能力。干旱下青杨两种群的内源ABA含量显著增加,碳同位素组分(δ13C)也显著提高。这些结果证明植物遭受干旱胁迫后发生一系列的形态、生理和生化响应,这些变化能提高植物在干旱下的存活和生长能力。2.青杨两种群对干旱胁迫反应的种群差异:与来自湿润地区的汉源种群相比,来自干旱地区的乐都种群在干旱条件下生物量向根系分配的可塑性更强,同时具有更强的抗氧化系统能力,所受到活性氧的伤害也更少,并且累积更多的脯胺酸和ABA,具有更高的δ13C。这些都说明了乐都种群对干旱的适应性比汉源种群更强。两种群对干旱的响应差异应归于它们的用水策略的不同:汉源种群来自湿润地区,采用了耗水型的用水策略,抗旱能力较弱;而乐都种群,来自干旱地区,通常采用节水型的用水策略,有更强的抗旱能力。3.遮荫对青杨两种群抗旱性的影响:遮荫对青杨抗旱性的影响决定于遮荫程度的不同,我们的结果表明中度的遮荫可以有效的提高干旱下植物的生长,对干旱胁迫有明显的缓解作用,具体体现在中度遮荫下受旱植物的叶片相对含水量得到提高,使得植物体内水分状况得到了改善;光合速率并未降低,植物光合氮利用效率增加,说明中度的遮荫并未明显限制植物的碳获得;抗氧化酶活性与膜脂过氧化产物MDA含量的同时降低,说明中度遮荫下所受到的活性氧伤害减少;中度遮荫下的ABA及δ13C的变化也不如在全光下变化明显,这也说明中度遮荫缓解了干旱胁迫。但是重度的遮荫却对干旱胁迫有明显的加剧作用,主要表现在重度遮荫降低了植物的光合速率,严重抑制了植物的生长;同时重度遮荫下脯胺酸含量和抗氧化酶活性的急剧下降,导致了植物渗调能力的下降及膜脂过氧化产物MDA的显著升高;重度遮荫还显著降低了内源ABA的累积和δ13C,降低了植物的抗旱能力。此外,青杨两种群在对干旱和遮荫的响应中,也表现出种群差异。汉源种群,来自湿润且年日照辐射较少的地区,表现出相对更强的耐荫性和需水性。而乐都种群,来自干旱且年日照辐射丰富的地区,表现出相对更强的耐旱性和需光性。这说明了植物对环境胁迫的耐受性是其长期适应原生境的结果,并且来自不同气候地区的两种群在面临环境胁迫时会采取不同的生存策略。4. 外源ABA喷施对青杨两种群抗旱性的影响:外源ABA的喷施可以提高两种群的抗旱性,具体表现为外源ABA喷施促进了青杨根系的生长,显著提高了干旱下植物的根/冠比和细/粗根比,减少了比叶面积;在生理生化方面,外源ABA降低了干旱下植物叶片的气孔导度,降低了蒸腾速率和净光合速率,但提高了瞬时用水效率,提高了叶片的相对含水量,增加了干旱下植物的保水能力。外源ABA进一步增加了干旱下植物内源ABA的积累,促进了植物渗调物质如脯胺酸和可溶性糖的积累,增加了抗氧化酶系统(如SOD、APX、CAT)的活性和非酶系统AsA的含量,降低了活性氧(如超氧阴离子(O2和过氧化氢(H2O2))对植株的伤害。此外,外源ABA还进一步提高了干旱下植物的δ13C,提高了植物的长期用水效率,由此提高了植物的抗旱能力。另一方面,两种群对外源ABA和干旱的响应也有所差别。来自湿润地区的汉源种群,对干旱较为敏感,所受干旱的影响也较大,而外源ABA的喷施对汉源种群抗旱性的提高作用也更为突出。乐都种群,由于其长期适应干旱地区的生长,本身已具有较强的抗旱能力,因此外源ABA喷施对其抗旱性的提高不如对汉源种群的效果明显。由此我们可以得出对于一些抗性弱或干旱敏感的物种或者种群,可以采用外施ABA的方法来提高其抗性。Poplars play an important role in lumber supply, and are important component ofecosystems due to their wide distribution and well adaptation. Populus cathayana Rehd.,which belongs to Populus Sect. Tacamahaca Spach, is one of the most important resources ofpoplars for its fast growth and reproductive. In this study, different populations of P.cathayana were used as experiment material to investigate the adaptability to drought stressand population differences in adaptability, and the effects of shade and exogenous abscisicacid (ABA) application on the drought tolerance. Our results could provide a strongtheoretical evidence and scientific direction for the afforestation, and rehabilitation ofecosystem in the arid and semi-arid area, and provide a strong evidence for adaptivedifferentiation of different populations, and so may be used as criteria for species selectionand tree improvement. The results are as follows:1. A large set of parallel response to drought stress: Drought stress caused pronouncedinhibition of the growth and increased relatively dry matter allocation into the root. For thetwo populations, the shoot height, basal diameter and total biomass were decreased but theroot/shoot ratio and fine root/coarse root ratio were increased under drought conditions;Drought stress caused pronounced inhibition of photosynthesis, decreased the stomatalconductance, transpiration rate, and photosynthetic nitrogen-use efficiency (PNUE) butincreased the instantaneous water use efficiency. Drought significantly improved the levels ofreactive oxygen species and malondialdehyde (MDA) and to induce the entire set ofantioxidative systems including the increase of activities of superoxide dismutase (SOD),ascorbate peroxidase (APX), catalase (CAT) and ascorbate (AsA) content. Drought decreased the leaf relative water content (RWC) but improved the capability of osmotic adjustmentindicated by the higher proline accumulation. Drought also increased the ABA content andcarbon isotope composition (δ13C), which indicating the long period water use efficiency wasimproved under drought. These results demonstrate that there are a large set of parallelchanges in the morphological, physiological and biochemical responses when plants areexposed to drought stress; these changes may enhance the capability of plants to survive andgrow during drought periods.2. Difference in adaptation to drought stress between contrasting populations of P.cathayana: Compared with the Hanyuan population (wet climate), the Ledu population (dryclimate) showed higher root/shoot ratio and water use efficiency, exhibited higherantioxidative systems capability thus resulting in less oxidative damage, accumulated moreABA and free proline content under drought conditions. The results suggested that there weredifferent water-use strategies between the two populations. The Ledu population, whichcomes from dry climate region, with higher drought tolerance, may employ a conservativewater-use strategy, whereas the Hanyuan population, which comes from wet climate, withlower drought tolerance, may employ a prodigal water-use strategy. These variations indrought responses may be used as criteria for species selection and tree improvement.3. The effects of shade on the drought tolerance: The reduction in the availability of lightand water affected the morphological and physiological responses of the two P. cathayanapopulations. In addition, the light environment modified the growth responses of P.cathayana seedlings to varying water environments in different ways depending upon theintensity of the light levels considered. There is an apparent alleviation to drought effects bymoderate shade in P. cathayana seedlings, as indicated by the higher leaf RWC, and unchanged net photosynthesis and PNUE, as well as by the lower antioxditative enzymeactivity, MDA, ABA and δ13C levels, which implied moderate shade did not significantlylimited the carbon acquisition or inhibited the plant growth, but ameliorated the detrimentaleffects of drought. On the other hand, an apparent aggravation to drought effects by severeshade was also observed, as indicated by the pronounced decrease of plant growth and net photosynthesis, the lower total biomass, ABA level, δ13C, free proline content andantioxditative enzyme activity and higher MDA accumulation. By contrast, the twopopulations showed different responses to shade and drought. The Hanyuan population,which comes from a riparian basin having a relatively wet climate and less annual solarradiation, is more sensitive to drought but more tolerant to shade. The Ledu population, whichcomes from a mountainous plateau with less rainfall and with more annual solar radiation, ismore tolerant to drought but more sensitive to shade. The results demonstrated that theendurance of plants to stress is a result of long-term evolution and adaptation to theenvironment, as suggested by the different strategies employed by the P. cathayanapopulations originating from contrasting habitats when they were exposed to drought andshade.4. The effects of exogenous ABA application on the drought tolerance: For bothpopulations under drought conditions tested, exogenous ABA application significantlyimproved the root/shoot ratio, fine root/coarse root ratio, and decreased the specifical leaf area.On the physiological and biochemical traits, exogenous ABA application significantlydecreased stomatal conductance, transpiration rate and net photosythesis but increased theinstance water use efficiency and leaf RWC. On the other hand, exogenous ABA applicationsignificantly increased endogenous ABA, proline, solube sugar and AsA content, as well asSOD, APX and CAT activities, thus reduced the damage of reactive oxygen species. Moreover,the long period water use efficiency as indicated by δ13C was also improved by exogenousABA application. In additionally, there was different responsive between the two populationsto drought and exogenous ABA application. The Hanyuan population, which comes from wetclimate region, is more sensitive to drought, and the effect of exogenous ABA is moreobviously than that in the Ledu population, which comes from dry climate region and is moredrought-responsive. Therefore, we can use exogenous ABA application to improve theresistance of plants, especially for the drought- sensitive species or populations.

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随着全球气候变暖和温室效应加剧,干旱和荒漠化成为威胁人类生存和发展的主要 灾害,许多被子植物对干旱胁迫的生理、生态和生化响应已逐步得以报道,但很少有开 展干旱胁迫对雌雄异株植物的影响方面的研究。由于这类植物在长期进化过程中已经在 生长、性比、生殖格局、空间分布、资源配置和生物量分配等方面形成了明显的性别差 异,因此,干旱胁迫必将对其雌雄植株产生不同的生理生态影响。本研究以青杨为模式 植物,采用植物生态、生理及生物化学等研究方法,系统研究青杨雌雄植株在常温、增 温以及喷施外源脱落酸的条件下对干旱胁迫的响应,揭示其在生长形态、生物量分配、 光合作用、用水效率和生理生化等方面的性别间差异。主要研究结果如下: 1. 青杨雌雄植株对干旱胁迫的综合响应。 与较好水分条件相比,干旱胁迫显著降低了青杨雌雄植株的光合作用和生长发育, 影响了许多生理生化过程,并导致雌雄植株在生长发育、气体交换、用水效率、膜脂抗 氧化和抗氧化系统酶活性方面表现出显著的性别间差异。在较好水分条件下,雌雄植株 之间在株高、基径、生物量、净光合速率、蒸腾速率、用水效率以及丙二醛、脱落酸和 游离脯氨酸等生化物质含量方面均无显著差异。但在干旱胁迫下,雄株在生长发育、气 体交换、水分利用效率、膜脂过氧化保护和抗氧化系统酶活性方面均显著高于雌株,表 现出比雌株更高的株高、基径、叶面积、总叶片数、总生物量、总色素含量、类胡萝卜 素含量、净光合速率、蒸腾速率、羧化效率、光系统II最大光化学效率、内在水分利用 效率、碳同位素组分、过氧化氢酶和过氧化物酶活性等,而在CO2补偿点、比叶面积、 叶绿素a/b、丙二醛、脱落酸和超氧化物歧化酶活性等指标上显著低于雌株。与雌株相比, 雄株表现出更高的干旱胁迫适应能力,而雌株的生长发育和生理生化过程更易遭受干旱 胁迫的影响。 2. 干旱胁迫下的青杨雌雄植株对增温处理的综合响应 与环境温度相比,增温在干旱胁迫前后均显著促进了雌雄植株的生长发育、气体交 换,降低水分利用效率,影响生化物质含量,并促使青杨雌雄植株之间在干旱胁迫下表 现出显著的差异。在较好水分条件下,增温导致雌株的株高、基径、叶面积、总叶片数、 总生物量和超氧化物歧化酶活性显著高于雄株,而用水效率、丙二醛、脱落酸和游离脯 氨酸、抗坏血酸过氧化物酶和过氧化物酶活性低于雄株。在干旱胁迫下,增温将导致雄 株的株高、基径、叶面积、总生物量、净光合速率、蒸腾速率、气孔导度、总色素含量、 相对含水量、过氧化氢酶和抗坏血酸过氧化物酶活性等显著高于雌株,而光系统II 最大 光化学效率、内在水分利用效率、碳同位素组分、丙二醛、脱落酸、游离脯氨酸和超氧 化物歧化酶活性显著低于雌株。与雄株相比,水分较好条件下的增温有利于促进雌株的 生长发育,并在生理生态特征上优于雄株。而干旱胁迫下的增温则加剧了水分胁迫强度, 致使雌株的生长发育遭受比雄株更多的负面影响。 3. 干旱胁迫下的青杨雌雄植株对喷施外源脱落酸处理的综合响应 与对照相比,在干旱胁迫下喷施外源脱落酸可显著增加青杨雌雄植株的生长发育、 气体交换、降低水分利用效率,影响了生化物质含量,并导致青杨雌雄植株之间在干旱 胁迫下表现出显著的生理生态差异。在干旱胁迫下,喷施外源脱落酸致使雌株的株高、 叶面积、叶干重、细根干重、总生物量、净光合速率、蒸腾速率、气孔导度、光系统II 最大光化学效率、非光化学淬灭系数、相对含水量、总光合色素、类胡萝卜素、脱落酸、 超氧化物歧化酶和过氧化物酶活性的增加量显著高于雄株,而根重比、根冠比、细根/ 总根、比叶面积、内在水分利用效率、碳同位素组分、丙二醛、脯氨酸、过氧化氢酶和 抗坏血酸过氧化物酶活性等指标的减少量上显著低于雄株。与对照相比,干旱胁迫下的 喷施外源脱落酸则一定程度能减缓植株遭受胁迫的压力,促进植株生长和气体交换,减 少了植株体内的过剩自由基数量,并促使雌株的生长发育和光合能力显著提高,增强其 抗干旱胁迫能力。 With development of global warming and greenhouse effect, drought and desertification have been became main natural disasteres in resent years. Studies on ecophysiological responses of most angiosperm species to environmental stress have been reported, but little is known about dioecious plant responses to drought stress. Since significant differences on growth, survival, reproductive patterns, spatial distribution, as well as resource allocation between males and females of dioecious plant have been formed during evolutionary process, sexual different ecophysiological responses should be caused by drought stress. In this experiment, Populus cathayana Rehd. was used as model plant to study the sex-related responses to drought by using the ecological, physiological and biochemical methods under normal atmospheric temperature, elevated temperatures and exogenous abscisic acid (ABA) application treatment respectively, and to expose the sexual differences in growth, biomass allocation, photosynthesis, water use efficiency and some biochemical material contents in the males and females of dioecious plant. The results are follows: 1. A large set of parallel responses of males and females of P. cathayana to drought stress Compared with well-watered treatment, drought significantly decreased growth and photosynthesis of P. cathayana individuals, affected some physiological and biochemical processes, and induced males and females to exhibit obvious sexual differences in growth, gas exchange, water use efficiency, lipid peroxidation protection and antioxidant defenses enzyme system. Under well-watered treatment, there were no significant sexual differences in height growth (HG), basal diameter (BD), dry matter accumulation (DMA), net photosynthesis rate (A), transpiration (E), water use efficiency (WUE), and malondialdehyde (MDA), abscisic acid (ABA) and praline (Pro). However, under drought stress, males were found to exhibit higher HG, BD, leaf area (LA), total leaf number (TLA), DMA, total chlorophyll contents (TC), carotenoids content (Caro), A, E, carboxylation efficiency (CE), the maximum efficiency of PSII (Fv/Fm), intrinsic water use efficiency (WUE ), carbon isotope composition (δ13C), catalase (CAT), peroxidase (POD) and lower CO2 compensation point (Γ), specific leaf area (SLA), chlorophyll a/b ratio (Chla/Chlb), MDA, ABA and superoxide dismutase (SOD) than females. The results suggest that males possess greater drought resistance than do females and females suffer more negative effect on growth and development, physiological and biochemical processes than males under drought stress. 2. A large set of parallel responses of drought-stressed males and females of P. cathayana to elevated temperatures Compared with environmental temperature, elevated temperature treatment significant increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents of P. cathayana individuals, and induced males and females to exhibit obvious differences under drought stress. Under good water condition, elevated temperature treatment caused females to show significant higher HG, BD, LA, TLN, DMA, SOD activity, and great lower WUE, MDA, ABA, Pro, ascorbate peroxidase (APX) and POD than do males. On contrary, under drought condition, elevated temperature treatment induced males to exhibit higher HG, BD, LA, DMA, A, E, stomatal conductance (gs), relative water content (RWC), CAT, APX activity but lower Fv/Fm, WUE, δ13C, MDA, ABA, Pro, SOD activity than do females. The results suggest that females will benefit from elevating temperature under good water condition by possessing better ecophysiological processes than that of males, but will suffer from greater negative effects than do males when grown under drought stress with elevated temperature treatment. 3. A large set of parallel responses of drought-stressed males and females of P. cathayana to exogenous ABA application Compared with controls, exogenous ABA application under drought greatly increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents in P. cathayana individuals, and induced males and females to exhibit obvious sexual differences under drought. Under drought stress, exogenous ABA application induced females to exhibit more increases in HG, LA, leaf weight (LW), fine root weight (FRW), DMA, A, E, g, Fv/Fm, non-photochemical quenching coefficient (qN), RWC, TC, Caro, ABA, SOD, POD s activity than males, but to show lower decreases in root/weight ratio (RWR), root mass/foliage area ratio (RF), fine root/total root ratio (FT), SLA, WUE, δ13C, MDA, Pro, CAT, APX than males. The results suggest that exogenous ABA application under drought stress will eliminate negative damages caused by drought stress at a certain extent,promote the growth and gas exchange of plant and decrease the number of superfluous 1O2 in plant cells of males and females of P. cathayana. Furthermore, exogenous ABA application promoted more drought resistance in females than in males by increasing more growth and photosynthetic capacity in females under drought stress.

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臭氧层损耗导致的地球表面UV-B辐射增强以及温室气体增多引起的气候变暖是当今两大全球环境问题。UV-B辐射增强和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。作为世界第三极的青藏高原,UV-B 辐射增强以及气候变暖现象尤为突出。本试验所在林区是青藏高原东缘的主要林区,具有大面积的亚高山人工针叶成熟林,在全球变化背景下该森林的天然更新潜力如何是急待回答的重要问题。基于此,本研究围绕森林树种的种子和幼苗这一更新的重要阶段,开展了气候变暖、UV-B辐射增强和联合胁迫对云杉种子萌发及幼苗定居影响的研究,旨在全球变化背景下,探讨全球变暖、UV-B 辐射增强和联合胁迫是否对西南地区大面积人工亚高山针叶林更新的种子萌发和幼苗定居阶段产生影响。 本文以青藏高原东缘亚高山针叶林主要树种云杉为研究对象,研究云杉种子萌发及幼苗的生长和生理对UV-B辐射增强与气候变暖的响应。采用UV-B荧光灯(UV-lamp)来模拟增强的UV-B 辐射,此外,采用开顶式有机玻璃罩(OTCs)来模拟气候变暖。本试验包括四个处理:(1)大气UV-B 辐射+大气温度(C);(2)大气UV-B 辐射+模拟气候变暖(W);(3)增强的UV-B辐射+大气温度(U);(4)增强的UV-B辐射+模拟气候变暖(U+W)。 根据本试验结果,UV-B辐射增强对云杉种子萌发没有显著影响,它对萌发云杉幼苗的影响主要体现在幼叶展开以后。根据两年的试验结果,增强的UV-B辐射降低了云杉幼苗抗氧化酶活性,降低了抗氧化物质的含量,此外,造成了膜质的过氧化,表现为MDA在针叶中的积累。增强的UV-B照射处理萌发云杉幼苗两年后,幼苗的生长受到显著抑制。我们的结果显示,OTCs分别提高了空气(10 cm)和土壤(5 cm)温度1.74℃和0.94 ℃。增温显著地促进了云杉种子提前萌发,提高了萌发速率和萌发比率,而且,明显地促进了幼苗的生长,表现为株高和生物量累积的显著增长。此外增温还有利于云杉幼苗根的伸长生长以及生物量的累积,这可以使云杉幼苗更好地利用土壤中的水分和营养元素。 根据本试验结果,温度升高显著地促进了增强UV-B辐射下云杉萌发幼苗的生长,这说明,温度升高缓解了UV-B辐射增强对云杉萌发幼苗的负面影响。这种缓解作用可能是温度升高对UV-B辐射增强处理下幼苗的抗氧化系统活性改善的结果。温度升高还缓解了高UV-B辐射对云杉幼苗根生长的抑制作用,这也可能是增温缓解伤害的原因之一。此外,根据我们的试验结果,增温与UV-B辐射增强联合作用(U+W)下云杉萌发幼苗的生长状况好于大气温度与大气UV-B辐射联合(C)处理,表现为株高、地径、根长和生物量积累均高于C处理,因此可以推断,UV-B辐射增强与气候变暖同时存在对萌发幼苗在两年之内的生长没有产生抑制作用,也就是说,气候变暖的缓解作用完全弥补了UV-B辐射增强的有害作用。 同样,增强的UV-B辐射显著影响了云杉幼苗的光合作用,表现为净光合速率(Pn)和表观量子效率(Φ)的提高,此外,根据我们的试验结果,它还造成了PSII的光抑制。增强的UV-B辐射显著抑制了云杉幼苗对营养元素的吸收,表现为大量营养元素、碳、钙、镁和锌含量的降低,但是,它却显著促进了铁在植株体内的积累。增温显著地提高了净光合速率,但是,它对光系统II(PSII)的光化学效率影响不大。温度升高缓解了UV-B增强对云杉幼苗光合作用的伤害,表现为净光合速率、表观量子效率以及PSII光化学效率的提高。此外,温度升高还缓解了UV-B辐射增强对离子吸收的抑制作用。 Enhanced UV-B radiation due to the reduction of O3 layer and global warming induced by increased greenhouse gases in the air have become the two pressing aspects of global climate changes. Moreover, enhanced UV-B radiation and warming have profound and long-term impacts on terrestrial plants and ecosystems, and the studies focusing on the two factors have attracted many attentions. Qinghai-Tibetan Plateau is the third in elevation in the world, and enhanced UV-B radiation and climate warming are especially prominent in this region. Our research located in the main forest belt in the eastern Qinghai-Tibetan Plateau where large areas of subalpine coniferous forests distributed. Based on that, we carried out a research to study the effects of enhanced UV-B radiation and climate warming on seed germination and seedlings growth of seedlings which are the important basic stage in forest regeneration. This research was arranged by a complete factorial design and included two factors (UV-B radiation and temperature) with two levels. The UV-lamps were used to manipulate the supplemental UV-B radiation and open-top chambers (OTCs) were adopted to increase temperature. The four treatments were: (1) C, ambient UV-B without warming; (2) U, enhanced UV-B without warming; (3) W, ambient UV-B with OTCs warming; (4) U+W, enhanced UV-B with OTCs warming. The main results were exhibited as follows: 1. Based on our results in this research, OTCs increased temperature on average 1.74℃ in air (10 cm above ground) and 0.92 ℃ in soil (5 cm beneath ground). Furthermore, OTCs also slightly reduced soil moisture and relative air humidity, however, the differences was not statistically significant. 2. Our results showed that enhanced UV-B had no significant effects on the seeds germination of P. asperata. Enhanced UV-B affected sprouts of P. asperata until the needles unfolded. During two years, enhanced UV-B inhibited the efficiency of the antioxidant defense systems, and as a result, it induced oxidant stress and the accumulation of MDA in needles. After two years of exposure to enhanced UV-B, the growth of P. asperata sprouts was markedly restrained compared with those under ambient UV-B radiation and temperature (C). Warming significantly stimulated the germination speed and increased the germination rate of P. asperata seeds. In the next place, it prominently facilitated the growth of P. asperata sprouts, represented as improvements in stem elongation and biomass accumulation. Furthermore, warming also increased root growth of P. asperata sprouts, which could made sprouts more efficient to use water and nutrient elements in soil. In this research, warming alleviated the deleterious effects of enhanced UV-B on P. asperata sprouts. It markedly stimulated the growth of P. asperata sprouts exposed to enhanced UV-B. The ease effects of warming on the abilities of the antioxidant defense systems might account for its amending effects on growth. After two years of exposure to enhanced UV-B radiation and warming, the growth of P. asperata sprouts was better than those under ambient UV-B radiation without warming (C), which could be seen from the higher plant height, basal diameter, root length and total biomass accumulation compared with C. 3. Enhanced UV-B radiation significantly influenced the photosynthesis processes of two-year old P. asperata seedlings. Our results showed that enhanced UV-B reduced the net photosynthetic rate (Pn) and the apparent quantum efficiency (Φ), and induced photoinhibition of photosynthetic system II (PSII). Enhanced UV-B significantly decreased the concentration of nitrogen (N), phosphorous (P), potassium (K), calcium (Ca), magnesium (Mg) and zinc (Zn), however, it increased the accumulation of iron (Fe) in the whole plant of P. asperata seedlings. Warming significantly stimulated Pn of P. asperata seedlings but it had no prominent impacts on the photochemical efficiency of PSII. In our research, warming also alleviated the harmful effects of enhanced UV-B on photosynthesis and absorption of ions of P. asperata seedlings. It increased Pn, Φ and the photochemical efficiency of PSII in seedlings exposed to enhanced UV-B. Moreover, warming also increased the absorption of ions of the seedlings exposed to enhanced UV-B radiation.

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干旱环境常常由于多变的降水事件和贫瘠土壤的综合作用,表现出较低的生产力和较低的植被覆盖度。全球性的气候变暖和人类干扰必将使得干旱地区缺水现状越来越严竣。贫瘠土壤环境中已经很低的有效养分含量也将会随着干旱的扩大而越来越低。干旱与半干旱系统中不断加剧的水分与养分的缺失将严重限制植物的生长和植被的更新,必然会使得已经恶化的环境恶化速率的加快、恶化范围的加大。如何抑制这种趋势,逐步改善已经恶化的环境是现在和将来干旱系统管理者面临的主要关键问题。了解干旱系统本土植物对未来气候变化的适应机制,不仅是植物生态学研究的重要内容,也对人为调节干旱环境,改善干旱系统植被条件,提高植被覆盖度具有重要的实践意义。 本研究以干旱河谷优势灌木白刺花(Sophora davidii)为研究对象,通过两年大棚水分和施N控制实验和一个生长季野外施N半控制实验,从植物生长-生理-资源利用以及植物生长土壤环境特征入手,系统的研究了白刺花幼苗生长特性对干旱胁迫和施N的响应与适应机制,并试图探讨施N是否可调节干旱系统土壤环境,人工促进干旱条件下幼苗定居,最终贡献于促进植被更新实践。初步研究结论如下: 1)白刺花幼苗生长、生物量积累与分配以及水分利用效率对干旱胁迫和施N处理的适应白刺花幼苗株高、基径、叶片数目、叶面积、根长、生物量生产、相对含水量和水分利用效率随着干旱胁迫程度的增加而明显降低,但地下部分生物量比例和R/S随着干旱胁迫程度的增加而增加。轻度施N处理下幼苗株高、基径、叶片数目、叶片面积和生物量生产有所增加。但重度施N处理下这些生长指标表现出微弱甚至降低的趋势。严重干旱胁迫条件下,幼苗叶面积率、R/S、相对含水量和水分利用效率也以轻度施N处理为最高。 2)白刺花幼苗叶片光合生理特征对干旱胁迫和施N处理的适应叶片光合色素含量和叶片光合效率随着干旱胁迫程度的增加而显著降低,并且PS2系统在干旱胁迫条件下表现出一定程度的光损害。但是比叶面积随着干旱胁迫程度的增加而增加。在相对较好水分条件下幼苗净光合速率的降低可能是因为气孔限制作用,而严重干旱胁迫条件下非气孔限制可能是导致幼苗叶片光合速率下降的主要原因。叶片叶绿素含量、潜在光合能力、羧化效率、光合效率以及RUBP再生能力等在施N处理下得到提高,并因而改善干旱胁迫条件下光合能力和效率。虽然各荧光参数对施N处理并无显著的反应,但是干旱胁迫条件下qN和Fv/Fm在轻度施N处理下维持相对较高的水平,而两年连续处理后在严重干旱胁迫条件下幼苗叶片光合效率受到重度施N处理的抑制,并且Fv/Fm和qN也在重度施N处理下降低。 3)白刺花幼苗C、N和P积累以及N、P利用效率对干旱胁迫和施N处理的适应白刺花幼苗C、N和P的积累,P利用效率以及N和P吸收效率随干旱胁迫程度的增加而显著降低,C、N和P的分配格局也随之改变。在相同水分处理下,C、N和P的积累量、P利用效率以及N和P吸收效率在轻度施N处理下表现为较高的水平。然而,C、N和P的积累量和P利用效率在重度施N处理下不仅没有表现出显著的正效应,而且有降低的趋势。另外,在相同水分条件下白刺花幼苗N利用效率随着施N强度的增加而降低。 4)白刺花幼苗生长土壤化学与微生物特性对干旱胁迫和施N的适应白刺花幼苗生长土壤有机C、有效N和P含量也随干旱胁迫程度的增加而明显降低。干旱胁迫条件下土壤C/N、C/P、转化酶、脲酶和碱性磷酸酶活性的降低可能表明较低的N和P矿化速率。尽管微生物生物量C、N和P对一个生长季干旱胁迫处理无显著反应,但微生物生物量C和N在两年连续干旱胁迫后显著降低。土壤有机C和有效P含量在轻度施N处理下大于重度施N处理,但是有效N含量随着施N强度的增加而增加。微生物生物量C和N、碱性磷酸酶和转化酶活性也在轻度施N处理下有所增加。但是碱性磷酸酶活性在重度施N处理下降低。 5)野外条件下白刺花幼苗生长特征及生长土壤生化特性对施N的适应植物生长、生物生产量、C的固定、N、P等资源的吸收和积累、其它受限资源的利用效率(如P)在轻度施N处理下均有所增加,但N利用效率有所降低。幼苗生物生产量及C、N和P等资源的分配格局在轻度施N处理下也没有明显的改变。白刺花幼苗叶片数目、生物生产量和C、N、P的积累量在重度施N处理下虽然也相对于对照有所增加,但幼苗根系长度显著降低。生物量及资源(生物量、C、N、P)在重度施N处理下较多地分配给地上部分(主要是叶片)。另外,土壤有机C、全N和有效N含量随外源施N的增加而显著增加,土壤pH随之降低,但土壤全P含量并无显著反应。其中有机C含量和有效P含量以轻度施N处理最高。微生物生物量C、N和P在轻度施N处理下也显著增加,而微生物生物量C在重度施N处理下显著降低。同时,转化酶、脲酶、碱性磷酸酶和中性磷酸酶活性在施N处理下也明显的提高,但酸性磷酸酶和过氧化氢酶活性显著降低,其中碱性磷酸酶和中性磷酸酶活性以轻度施N处理最高。 综合分析表明,干旱河谷水分和N严重限制了白刺花幼苗的生长。施N不能完全改变干旱胁迫对白刺花幼苗的抑制的作用,但是由于施N增加土壤N有效性,改善土壤一系列生物与化学过程,幼苗的生长特性也对施N表现出强烈的反应,表现为植物结构与资源分配格局的改善,植物叶片光合能力与效率的提高,植物生长以及利用其他受限资源(如水分和P)的效率的增加,致使植物自身生长及其生长环境在干旱环境下得到改善。但是过度施N不仅不能起到改善干旱胁迫下植物生长环境、促进植物生长的作用,反而在土壤过程以及植物生长过程中加重干旱胁迫对植物的伤害。因此,建议在采用白刺花作为先锋种改善干旱河谷系统环境的实践中,可适当施加N以改善土壤环境,调节植物利用与分配资源的效率,促进植物定居,得到人工促进种群更新的目的。但在实践过程中也要避免过度施N。 Arid regions of the world are generally noted for their low primary productivity which is due to a combination of low, unpredictable water supply and low soil nutrient concentrations. The most serious effects of global climate change and human disturbances may well be those which related to increasing drought since drought stress has already been the principal constraint in plant growth. The decline in total rainfall and/or soil water availability expected for the next decades may turn out to be even more drastic under future warmer conditions. Nevertheless, water deficit is not the only limiting factor in arid and semiarid environments. Soils often suffer from nutrient (especially N and P) deficiencies in these ecosystems, which can also be worsened by climate change. How to improve the poor soil quality and enhance the vegetation coverage is always the problem facing ecosystem managers. The adaptive mechanisms of native plant to future climate change is always the focus in plant ecology, it also plays important roles in improving vegetation coverage by manual controlled programmes. Sophora davidii is a native perennial shrub of arid valleys, which is often predominant on eroded slopes and plays a vital role in retaining ecological stability in this region. It has been found that S. davidii was better adapted to dry environment than other shrubs, prompting its use for re-vegetation of arid lands. A two-years greenhouse experiment and a field experiment were conducted in order to understand the adaptation responses of Sophora davidii seedlings to different water and N conditions, and further explore if additional N supply as a modified role could enhance the adaptation ability of S. davidii seedlings to dry and infertile environment. Two-month old seedlings were subjected to a completely randome design with three water (80%, 40% and 20% water field capacity (FC)) and three N supply (N0: 0, Nl: 92 and Nh: 184 mg N kg-1 soil) regimes. Field experiment was arranged only by three N supplies in the dry valley. 1) The growth, biomass partitioning and water-use efficiency of Sophora davidii seedlings in respond to drought stress and N supply Seedlings height, basal diameter, leaf number, leaf area, root length, biomass production, relative water content (RWC) and WUE were decreased with increase of drought stress. An increase in below-ground biomass was observed indicating a higher root/shoot ratio (R/S) under drought stress conditions. Low N supply increased seedlings height, basal diameter, leaf number, leaf area, and biomass production, but decreased root length. In contrast, these growth characteristics showed little or negative effect to high N supply treatment. Leaf percentages increased with increase of N supply, but fine root percentages decreased. In addition, Low N supply rather than the other two N treatments increased leaf area ratio (LAR), leaf/fine root mass ratio (L/FR), R/S and RWC under severe drought stress (20%FC), even though these parameters could increase with the high N supply treatment under well-watered condition (80%FC). Moreover, Low N supply also increased WUE under three water conditions, but high N supply had little effect on WUE under drought stress conditions (40%FC and 20%FC). 2) Leaf gas exchange and fluorescence parameters of Sophora davidii seedlings in respond to drought stress and N supply Leaf area (LA), photosynthetic pigment contents, and photosynthetic efficiency were decreased with increase of drought stress, but specific leaf area (SLA) increased. Photodamage in photosystem 2 (PS2) was also observed under drought stress condition. The decreased net photosynthetic rate (PN) under relative well-watered water conditions might result from stomatal limitations, but the decreased PN under other hand, photosynthetic capacity by increasing LA, photosynthetic chlorophyll contents, Pnmax, CE, Jmax were increased with increase N supply, and photosynthetic efficiency was improved with N supply treatment under water deficit. Although N supply did a little in alleviating photodamages to PS2 caused by drought stress, low N supply enhanced qN and kept relative high Fv/Fm under drought stress condition. However, high N supply inhibited leaf photosynthetic efficiency, and declined Fv/Fm and qN under severe drought stress condition after two year continues drought stress and N supply. 3) Carbon accumulation, nitrogen and phosphorus use efficiency of Sophora davidii seedlings in respond to drought stress and N supply C, N and P accumulation, NUE , N and P uptake efficiency (NUtE and NUtE ) P N P were decreased with increase of drought stress regardless of N supply. On the other hand, the S. davidii seedlings exhibited strong responses to N supply, but the responses were inconsistent with the various N supply levels. Low N supply rather than the other two N treatments increased C, N and P accumulation, improved NUEP, NUtE and NUtE under corresponding water condition. In contrast, high N supply N P did few even depressed effects on C, N and P accumulation, and NUEP, although NUtEN and NUtEP could increase with high N supply under corresponding water conditions. Even so, a decrease of NUEN was observed with increase of N supply under corresponding water conditions. 4) Soil microbial and chemical characters in respond to drought stress and N supply The content of soil organic C, available N and P were decreased with increase of drought stress. Decreases in C/N and C/P, and invertase, urea and alkaline phosphatase activity were also observed under drought stress conditions, indicating a lower N and P mineralization rate. Although microbial biomass C, N and P showed slight responses to drought stress after one growth period treatment, microbial biomass C and N were also decreased with increase of drought stress after two year continuous treatment. The content of soil organic C and available P showed the stronger positive responses to low N supply than which to high N supply, although than the other two N treatments increased microbial biomass N and invertase activity under severe drought stress condition, even though invertase activity could increase with high N supply treatment under relative well-water conditions. Moreover, low N supply treatment also increased C/P and alkaline phosphatase activity which might result from higher P mineralization, but high N supply did negative effects on alkaline phosphatase activity. 5) The growth characteristics of Sophora davidii seedlings and soil microbial and chemical characters in respond to N supply under field condition Low N supply facilitated seedlings growth by increasing leaf number, basal diameter, root length, biomass production, C, N and P accumulation and absorption, and enhancing the use efficiency of other limited resources as P. Compared to control, however, low N supply did little effect on altering biomass, C, N and P portioning in seedlings components. On the contrary, high N supply treatment also increased leaf number, biomass and C, N and P accumulation relative to control, but significantly decreased root length, and altered more biomass and resources to above-ground, which strongly reduced the ability of absorbing water under drought condition, and thus which might deep the drought stress. In addition, N supply increased soil C, N and available N content, but declined pH and showed little effects on P content. Low N supply showed higher values of soil C and available P content. Low N supply also increased microbial biomass C, N and P, although high N supply decreased microbial biomass C. N supply significantly enhanced soil invertase, urea, alkaline and neutral phosphratase activity, while declined acid phosphratase and catalase activity. Low N supply exhibited higher alkaline and neutral phosphratase activity compared to the others. The results from this study indicated that both drought and N limited the growth of S. davidii seedlings and their biomass production. Regardless of N supply levels, drought stress dramatically reduced the seedlings growth and biomass production. Although plant growth parameters, including basal diameter, height, leaf number, and biomass and their components were observed to be positive responses to low N supply, N supply alone can not alter the diminishing tendency which is caused by drought. available N content increased with increase N supply. In addition, low N supply rather These findings imply that drought played a primary limitation role and N was only the secondary. Even so, appropriate N supply was seemed to enhance the ability that S. davidii seedlings adapted to the xeric and infertile environment by improving soil processes, stimulating plant growth, increasing recourses accumulation, enhancing use efficiency of other limited resources, and balancing biomass and resources partitioning. Appropriate N supply, therefore, would be recommended to improve S. davidii seedling establishment in this region, but excess N supply should be avoided.

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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.

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光是植物赖以生存的重要环境因子,但是植物在获得光的同时不可避免的会受到紫外辐射的伤害。尤其是近年来,人类向大气中排放的大量氮氧化合物和氟氯烃类化合物(CFC’s)引起臭氧分子的分解,导致到达地球表面的紫外辐射增加,特别是UV-B辐射增强。而另一方面,植物对UV-B辐射反应的敏感性在种间和品种间存在差异,主要受植物基因型,生态型和生活型的控制。本项目分别以粗枝云杉和青杨组杨树为模式植物,从形态和生理生化方面分别研究了来自不同水分背景下的粗枝云杉种群和来自不同UV-B背景下的青杨种群在增强UV-B下的反应及其反应差异,并探讨了干旱、喷施外源脱落酸(ABA)对它们抗UV-B能力的影响。研究成果可为生态系统的恢复与重建提供理论依据和科学指导。主要研究结果如下: 1. 粗枝云杉的两个种群,湿润种群(来自四川黑水)和干旱种群(来自甘肃迭部)在水分良好和干旱状况下表现出对增强UV-B的不同响应。同时,干旱对粗枝云杉抗UV-B能力的影响也得到研究:两种胁迫共同作用时,干旱表现出在一定程度上减弱了增强UV-B对粗枝云杉的生理特性的影响。 干旱胁迫显著降低了两个粗枝云杉种群的光合同化速率(A), 气孔导度(gs)和PSII的有效光量子产量(Y), 同时,提高了非光化学猝灭效率(qN)和超氧化物歧化酶(SOD)的活性。与湿润种群相比,干旱种群抗旱性更强,表现为干旱种群拥有更高的SOD和干旱进一步加剧了UV-B的胁迫效应。 本研究中,干旱胁迫单独作用时,显著降低了青杨两个种群的生物量积累和气体交换,具体包括A、gs、蒸腾速率(E)和光合氮利用效率(PNUE),提高了两个种群的瞬时水分利用效率(WUEi)、长期水分利用效率(WUET)、碳同位素组分(δ13C)和氮含量(N)。同时,UV吸收物质和ABA含量也得到积累。另一方面,增强UV-B对青杨两个种群各个指标的影响,同干旱所引起的效应有着相似的趋势。同低海拔种群相比,高海拔种群有着更强的抗旱和抗UV-B能力,具体表现在高海拔种群有着更多的生物量积累,更强的气体交换和水分利用效率及更高水平的ABA和UV吸收物质含量。相比干旱诱导的生物量积累和气体交换的降低,在干旱和增强UV-B两个胁迫同时作用于青杨时,这种降低表现的更为明显。显著的干旱和UV-B的交互作用还表现在WUEi, WUET, δ13C, 可溶性蛋白含量, UV吸收物质含量, ABA, 叶片和茎中的N含量以及C/N比中。 3. 经过一个生长季的试验观察,增强UV-B、外源ABA及两因子共同作用对青杨的生物量积累、气体交换、内源ABA和UV吸收物质含量、抗氧化系统以及碳、氮含量和碳/氮比均产生显著影响。本试验中,青杨的两个种群分别来自中国西南部的不同海拔地区,高海拔种群来自青海大通而低海拔种群来自四川九寨。外源ABA的胁迫为直接喷施ABA到青杨叶片,而增强UV-B胁迫是利用平方波系统分别保证青杨苗暴露于外界UV-B强度和两倍于外界UV-B强度下。 研究结果显示,增强UV-B显著的降低了两个青杨种群的株高、基茎、总叶面积和总生物量等生长指标,同时也导致其A、gs、E和叶片中碳含量的减少。而显著增加了SOD和过氧化物酶(GPx)活性水平,诱导了过氧化氢(H2O2)和MDA的显著增加,促进了UV吸收物质和不同器官中内源ABA含量的显著积累。另一方面,外源ABA引起了青杨光合同化速率的下降,SOD和GPx酶活性的增强,H2O2 和 MDA含量也表现出显著增加,同时,内源ABA含量得到显著累积。同低海拔种群相比,高海拔种群具有更加抗UV-B和外源ABA的特性。显著的UV-B和ABA的交互作用表现在A, E, SOD和GPx活性,以及叶片和根部的内源ABA等一系列指标中。在所有胁迫下,叶片中的碳和氮含量同其在茎和根中的含量显著相关,另外,叶片和茎中的氮含量同茎中的碳含量显著相关。 Sunlight is an indispensable environment factor for plants survival and development. Meanwhile, photosynthetic organisms need sunlight and are thus, inevitably, exposed to UV radiation. Especially for recent years, ultraviolet radiation, especially UV-B reaching the Earth’s surface increased because of depletion of ozone layer resulted from emission of NxO and CFC’s from human activities. On the other hand, the sensitivity of plants to UV-B radiation depends on the species, developmental stage and experimental conditions. In this experiment, two populations of Picea asperata Mast from different water background and two populations of Populus cathayana Rehder from different altitude background were selected as model plants to assess the effects of enhanced UV-B radiation. Morphological and physiological traits induced by enhanced UV-B in each plant species were observed and the different responses were discussed, furthermore the influences of drought and exogenous ABA on responses induced by enhanced UV-B were studied. The study could provide a strong theoretical evidence and scientific direction for the afforestation and rehabilitation of ecosystem. The results are as follows: 1. Different responses of two contrasting Picea asperata Mast. populations to enhanced ultraviolet-B (UV-B) radiation under well-watered and drought conditions were investigated. And the effects of enhanced UV-B on tolerance of drought were also observed in our study that the UV-B exposure may have alleviated some of the damage induced by drought. Two contrasting populations, originating from a wet and dry climate region in China, respectively, were employed in our study. Drought significantly decreased CO2 assimilation rate (A), stomatal conductance (gs) and effective PSII quantum yield (Y), while it significantly increased non-photochemical quenching (qN) and the activity of superoxide dismutase (SOD) in both populations. Compared with the wet climate population, the dry climate population was more acclimated to drought stress and showed much higher activities of SOD and ascorbate peroxidase (APX), and much lower levels of malondialdehyde (MDA) and electrolyte leakage. On the other hand, enhanced UV-B radiation also induced a significant decrease in the chlorophyll (Chl) content in both populations under well-watered conditions, and a significant increase in UV-absorbing compounds in the wet climate population. After one growing season of exposure to different UV-B levels and watering regimes, the increases in MDA and electrolyte leakage, as induced by drought, were less pronounced under the combination of UV-B and drought. In addition, an additive effect of drought and UV-B on A and gs was observed in the wet climate population, and on the activity of APX and qN in the dry climate population. 2. The significant effects of drought, enhanced UV-B radiation and their combination on Populus cathayana Rehd. growth and physiological traits were investigated in two populations, originating from high and low altitudes in south-west China. Our results showed that UV-B acts as an important signal allowing P. cathayana seedlings to respond to drought and that the combination of drought and UV-B may cause synergistically detrimental effects on plant growth in both populations. In both populations, drought significantly decreased biomass accumulation and gas exchange parameters, including A, gs, E and photosynthetic nitrogen use efficiency (PNUE). However, instantaneous water use efficiency (WUEi), transpiration efficiency (WUET), carbon isotope composition (δ13C) and nitrogen (N) content, as well as the accumulation of soluble protein, UV-absorbing compounds and abscisic acid (ABA) were significantly increased by drought. On the other hand, cuttings from both populations, when kept under enhanced UV-B radiation conditions, showed very similar changes in all above-mentioned parameters, as induced by drought. Compared with the low altitude population, the high altitude population was more tolerant to drought and enhanced UV-B, as indicated by the higher level of biomass accumulation, gas exchange, water-use efficiency, ABA concentration and UV-absorbing compounds. After one growing season of exposure to different UV-B levels and watering regimes, the decrease in biomass accumulation and gas exchange, induced by drought, was more pronounced under the combination of UV-B and drought. Significant interactions between drought and UV-B were observed in WUEi, WUET, δ13C, soluble protein, UV-absorbing compounds, ABA and in the leaf and stem N, as well as in the leaf and stem C/N ratio. 3. During one growing season, significant effects induced by enhanced UV-B radiation, exogenous ABA and their combination on biomass accumulation, gas exchange, endogenous ABA and UV-absorbing compounds concentrations, antioxidant system as well as carbon (C) content, nitrogen (N) content and C/N ratio were investigated in two contrasting Populus cathayana populations, originating from high and low altitudes in south-west China. Exogenous ABA was sprayed to the leaves and enhanced UV-B treatment was using a square-wave system to make the seedlings under ambient (1×) or twice ambient (2×) doses of biologically effective UV-B radiation (UV-BBE). Enhanced UV-B radiation significantly decreased height, basal diameter, total leaf area, total biomass, A, gs, E and carbon (C) content in leaves, and significantly increased activities of SOD and guaiacol peroxidase (GPx), hydrogen peroxide (H2O2) and malonaldehyde (MDA) content as well as the accumulation of UV-absorbing compounds and endogenous ABA concentrations among different organs in both populations. In contrast, exogenous ABA showed significant decrease in A and significant increases in activities of SOD and GPx, H2O2, MDA content and the endogenous ABA concentrations. Compared with the low altitude population, the high altitude population was more tolerant to enhanced UV-B and exogenous ABA. Significant interactions between UV-B and ABA were observed in A, E, activities of SOD and GPx, as well as in endogenous ABA in leaves and roots of both populations. Across all treatments, C and N content in leaves was strongly correlated with those were in stems and roots, respectively. Additionally, leaf and stem N content were significant correlated with stem C content.

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黄龙世界自然遗产地岷江冷杉林(Abies faxoniana)生境类型多样,群落结构复杂,群落植物种类组成多样性丰富。揭示不同生境的生物多样性及其差异是认识生物多样性格局、形成及维持机制的前提和进行多样性保育的基础。本文采用样方法对黄龙钙化滩生境、阴坡非钙化生境及半阳坡非钙化生境的岷江冷杉原始林植物群落结构及植物多样性进行了研究。结果表明: 黄龙岷江冷杉林具有明显的复层异龄结构,垂直结构明显,乔木、灌木、草本、苔藓层次分明。共发现高等植物386 种,其中维管植物46 科103 属163 种,苔藓植38 科83 属物223 种。各层片结构及物种组成如下: (1)钙化滩生境、阴坡非钙化生境、半阳坡非钙化生境分别发现乔木18 种、13种、8 种。乔木层均可分为两个亚层,第一亚层优势种均为岷江冷杉,第二亚层主要为岷江冷杉异龄树或其它大高位芽物种。钙化滩生境第一亚层除优势种岷江冷杉外混生有巴山冷杉(Abies fargesii)、粗枝云杉(Picea asperata)以及阔叶树种白桦(Betula platyphylla)等,第二亚层主要为岷江冷杉异龄树;阴坡非钙化生境第一亚层除优势种岷江冷杉外间有巴山冷杉和白桦,第二亚层物种主要为川滇长尾槭(Acer caudatum var. prattii);半阳坡非钙化生境第一亚层除优势种岷江冷杉外混生有巴山冷杉,第二亚层主要为岷江冷杉异龄树。依乔木层优势种的差异,钙化滩生境及半阳坡非钙化生境为岷江冷杉纯林,阴坡非钙化生境为岷江冷杉-川滇长尾槭混交林。不同生境乔木层郁闭度、乔木密度、树高结构、直径结构均存在差异。 (2)钙化滩生境发现灌木41 种,平均盖度为18.49±1.72(%),平均高度为52.12±4.45(cm),优势种为直穗小檗(Berberis dasystachya);阴坡非钙化生境发现灌木30 种,平均盖度为29.33±2.56 (%),平均高度为119.55±8.01 (cm),优势种为箭竹 (Fargesia spathacea) 、唐古特忍冬(Lonicera tangutica) 和袋花忍冬(Lonicera saccata);半阳坡非钙化生境发现灌木29 种,平均盖度为31.35±1.93 (%),平均高度为107.55±4.24 (cm),优势种为箭竹(Fargesia spathacea)。不同生境灌木层结构和物种组成多样性差异显著,钙化滩生境的灌木盖度、高度总体上较非钙化的坡地生境低, 钙化滩生境灌木以小型叶的落叶灌木为主,沟两侧非钙化的坡地生境上则发育了丰富箭竹。 (3)钙化滩生境发现草本46 种,平均盖度为7.18±0.79 (%),平均高度为5.04±0.26(cm),以山酢浆草(Oxalis griffithii)为优势种;阴坡非钙化生境发现草本物种71 种,平均盖度达29.04±2.31(%),平均高度为9.08±0.52(cm),以钝叶楼梯草(Elatostema obtusum)、山酢浆草为优势种;半阳坡非钙化生境草本物种50 种,平均盖度为以8.79±0.82(%),平均高度为7.67±0.43 (cm),以扇叶铁线蕨(Adiantum flabellulatum)、双花堇菜(Viola biflora)、华中蛾眉蕨(Lunathyrium shennongense)、山酢浆草为优势种。阴坡非钙化生境草本层片发育良好,多样性最为丰富,盖度和物种丰富度均显著高于钙化滩生境和半阳坡非钙化生境。 (4)钙化滩生境发现苔藓物种140 种,平均盖度达84.25±1.30 (%),以仰叶星塔藓(Hylocomiastrum umbratum) 等大型藓类为优势种;阴坡非钙化生境发现苔藓物种115 种,平均盖度为79.29±1.64 (%),以刺叶提灯藓(Mnium spinosum)、大羽藓(Thuidium cymbifolium)、毛尖燕尾藓(Bryhnia trichomitra)等个体较小的物种为优势种;半阳坡非钙化生境发现苔藓物种91 种,平均盖度为60.64±1.93 (%),也以刺叶提灯藓为优势种。 (5)钙化滩生境、阴坡非钙化生境、半阳坡非钙化生境的物种数分别为234 种、221 种、175 种。乔木层的Shannon-Wiener 指数分别为0.75 ±0.12、1.87±0.12、1.78±0.07(灌木层,0.44±0.08、1.71± 0.15、2.49±0.06;草本层,0.33±0.13、1.31±0.15 、2.15±0.08; 苔藓层1.30±0.11、2.08±0.04、1.73±0.11,);Pielou 均匀度指数分别为0.45±0.05、0.29±0.06、0.28±0.08(灌木层,0.75±0.03、0.68±0.05、0.52±0.06;草本层,0.68±0.02、0.77±0.02、0.74±0.02;苔藓层,0.40±0.03、0.63±0.02、0.52±0.03);Simpson's 优势度指数分别为0.63±0.06、0.78±0.04、0.83±0.07(灌木层,0.21±0.03、0.28±0.05、0.45±0.06;草本层,0.25±0.02、0.12±0.01、0.17±0.01;苔藓层,0.45±0.04、0.18±0.01、0.31±0.04)。三种生境间乔木层、草本层的Sorenson 群落相似性系数较低, 灌木层、苔藓层的的Sorenson 群落相似性系数较高。 综上所述,黄龙岷江冷杉林的群落结构、植物多样性在三种生境间存在差异性,这将意味着我们在进行黄龙世界自然遗产地的森林经营管理时要较多地关注岷江冷山林群落在不同生境中的差异性。 There were multiplex habitat types, complicated community structure and abundant species composition in the Huanglong World Natural Heritage Site. Uncovering the differences of biodiversity among different habitats was a precondition to understand the distribution, formation and sustaining mechanism of the biodiversity, and the foundation of biodiversity conservation. In the present study, using plenty of quadrants, we investigated the community structure and the biodiversity of the primitive Abies faxoniana forest in different habitats (travertine bottomland, semi-sunny-slope non-calcified habitat and shady-slope non-calcified habitat) in the Huanglong World Natural Heritage Site. The main results are as follows: All the primitive Abies faxoniana forests in the three habitats were uneven-aged with obvious vertical structure including tree layer, shrub layer, herb layer and bryophyte layer. A total of 386 higher plants including 163 vascular plant species (103 generic, 46 families) and 223 bryophyte species (83 generic, 38 families) were investigated. The structure and species composition of each layer are as follows: (1) There were 18, 13 and 8 tree species in travertine bottomland, shady-slope non-calcified habitat and semi-sunny-slope non-calcified habitat, respectively. The tree layers in all habitats can be divided into two clear sub-layers. The upper tree layers were dominated by Abies faxoniana, and the lower tree layers were dominated by uneven-aged Abies faxoniana or other phanerophytes species. There were Abies fargesii , Picea asperata and Betula platyphylla besides the dominated species (Abies faxoniana) in the upper tree layer in travertine bottomland, and the lower tree layers were dominated by uneven-aged Abies faxoniana; There were Abies fargesii and Betula platyphylla besides the dominated species (Abies faxoniana) in the upper tree layer in shady-slope non-calcified habitat, and the lower tree layers were dominated by Acer caudatum var. prattii; There was Abies fargesii besides the dominated species (Abies faxoniana) in the upper tree layer semi-sunny-slope non-calcified habitat, and the lower tree layers were dominated by uneven-aged Abies faxoniana. According to composition percentage of dominate species in tree layer, both the forest in travertine bottomland and in semi-sunny-slope non-calcified habitat could be ranked as pure forest, and the forest in shady-slope non-calcified habitat could be ranked as mingled forest. There were significant differences in crown density, plant density, height structure and diameter structure among the three habitats. (2) A total of 41 shrub species (average coverage 18.49±1.72%; average height 52.12±4.45 ㎝)were found in travertine bottomland, and the dominate species was Berberis dasystachya; A total of 30 shrub species (average coverage 29.33±2.56 %;average height 119.55±8.01 ㎝)were found in shady-slope non-calcified habitat, and the dominate species was Fargesia spathacea, Lonicera tangutica and Lonicera saccata. A total of 29 shrub species (average coverage 31.35±1.93%; average height 107.55±4.24 ㎝) were found in semi-sunny-slope non-calcified habitat, and the dominate species was Fargesia spathacea. There were significant differences in structure and species diversity of the shrub layers among the three habitats. The coverage and height of shrub had lower value in travertine bottomland than in two non-calcified habitats. Moreover, travertine bottomland was dominated by deciduous shrub species with microphyll and non-calcified habitats developed abundant Fargesia spathacea species. (3) A total of 46 herb species (average coverage 7.18±0.79%;average height 5.04±0.26 ㎝)were found in travertine bottomland, and the dominate species was Oxalis griffithii; A total of 71 herb species (average coverage 29.04±2.31%;average height 9.08±0.52 ㎝)were found in shady-slope non-calcified habitat, and the dominate species was Elatostema obtusum and Oxalis griffithii. A total of 50 herb species (average coverage 8.79±0.82%;average height 7.67±0.43 ㎝) were found in semi-sunny-slope non-calcified habitat, and the dominate species was Adiantum flabellulatum, Viola biflora, Lunathyrium shennongense and Oxalis griffithii. Herb layers developed well in shady-slope non-calcified habitat and had the higher species richness and coverage than travertine bottomland and semi-sunny-slope non-calcified habitat. (4) A total of 140 bryophyte species (average coverage 84.25±1.30%)were found in travertine bottomland, and the dominate species was big bryophyte species such as Hylocomiastrum umbratum and so on; A total of 115 bryophyte species (average coverage 79.29±1.64%)were found in shady-slope non-calcified habitat, and the dominate species was small bryophyte species such as Mnium spinosum, Thuidium cymbifolium, Bryhnia trichomitra and so on. A total of 91 bryophyte species (average coverage 60.64±1.93%) were found in semi-sunny-slope non-calcified habitat, and the dominate species was Mnium spinosum. (5) There were 234, 221 and 175 plant species in travertine bottomland, shady-slope non-calcified habitat and semi-sunny-slope non-calcified habitat, respectively. Shannon-Wiener index of the tree layer was 0.75 ±0.12, 1.87±0.12 and 1.78±0.07 (the shrub layer, 0.44±0.08, 1.71± 0.15 and 2.49±0.06; the herb layer, 0.33±0.13, 1.31±0.15 and 2.15±0.08; the bryophyte layer, 1.30±0.11, 2.08±0.04 and 1.73±0.11.) for the three habitats, respectively; Pielou index of the tree layer was 0.45±0.05, 0.29±0.06 and 0.28±0.08 (the shrub layer, 0.75±0.03, 0.68±0.05 and 0.52±0.06; the herb layer, 0.68±0.02, 0.77±0.02 and 0.74±0.02; the bryophyte layer, 0.40±0.03, 0.63±0.02 and 0.52±0.03.) for the three habitats, respectively. Simpson's index of the tree layer was 0.63±0.06, 0.78±0.04 and 0.83±0.07 (the shrub layer, 0.21±0.03、0.28±0.05、0.45±0.06; the herb layer, 0.25±0.02, 0.12±0.01 and 0.17±0.01; the bryophyte layer, 0.45±0.04, 0.18±0.01 and 0.31±0.04.) for the three habitats, respectively. There were low Sorenson index both in the tree layer and in the herb layer among the three habitats, whereas, high Sorenson index occurred both in the shrub layer and in the bryophyte layer. To sum up, there were differences both in community structure and plant diversity among the three different habitats, which means that we should pay more attention to habitats heterogeneities of the primitive Abies faxoniana forest when we take action to manage the forest in the Huanglong World Natural Heritage Site.