Long-term grazing alters species composition and biomass of a shrub meadow on the Qinghai-Tibet Plateau

Livestock grazing has long been the most widespread land use on the Qinghai-Tibet Plateau, one of the world's highest ecosystems. However, there has been increasing concern during recent decades because of the rapid increase in livestock numbers. To assess the possible influences of grazing on the vast grassland, a long-term grazing experiment in a shrub meadow on the northern Qinghai-Tibet Plateau was carried out. The experiment included five treatments with different stocking rates and one non-grazing (N) treatment. After 17 years of grazing, treatment differences were clear. The species composition differed markedly between grazing intensities, with a decrease in palatable grass species and an increase in unpalatable forbs at higher grazing intensities. The species richness and species diversity, however, were not significantly different between treatments. Vegetation height decreased significantly at higher grazing intensities. Total above,ground biomass declined considerably and the biomass of forbs increased significantly under the higher grazing intensities. The amount of litter was significantly lower under the higher grazing intensities. The results suggest that long-term grazing alters the species composition, vegetation height and biomass production of the alpine grassland ecosystem without significantly changing species richness.

黄土丘陵区不同坡向及坡位草本群落生物量及多样性研究

以黄土丘陵区纸坊沟流域内的大范家沟阴阳坡和拐沟阴坡坡面主要草本群落为研究对象,分析不同坡向及坡位的草本群落盖度、地上生物量及物种多样性关系。结果表明,阴坡草本群落的物种20多种,较阳坡高30%。阴阳坡面上草本群落生物量和盖度从坡下至坡上逐渐减小。在水分和养分条件好的地段均有灌木入侵。阳坡Margalef和Menhinick丰富度指数均随坡位升高而降低;而阴坡Mafgalef指数随坡位升高而降低,Menhinick指数却随坡位升高而增大;3个均匀度指数和多样性指数在阴阳坡均随坡位的升高而减小。不同坡位引起的植被盖度、Margalef丰富度指数和Alatalo均匀度指数存在显著差异(P<0.05)。坡位与Margalef丰富度指数和Alatalo均匀度指数呈显著相关(P<0.05),与Shannon-wiener多样性指数呈极显著相关(P<0.01)。描述或评价群落多样性、丰富度及均匀度时,应多选几个指标参数,避免单一指标带来的片面性。掌握草本群落自身喜阴阳特性、生境特征、植被组成及多样性特征,充分认识植被演替条件和演替规律,对指导该区植被和生态系统恢复与重建具有重要意义。

云雾山本氏针茅草地群落恢复演替过程中的物种多样性变化动态

采用样方法对云雾山干草原区本氏针茅草地群落进行长期定位监测,对未封育样地和封育5,10,15,20和25年样地群落的平均高度、盖度、多度和地上生物量以及植物多样性进行实地调查分析,以研究围栏封育对草地植物群落特征动态变化的影响。结果表明,围封10年样地群落的高度、多度以及地上生物量最大,而群落的盖度则随封育年限的增加呈显著增加趋势,在封育25年达到最大;围栏封育的不同年限也显著影响了群落的物种多样性,在围封10年样地群落具有最高的多样性指数,围封15年样地具有最高的丰富度指数,而均匀度指数则是在未封育群落中表现为最高值,最低值是在物种丰富度最高时出现。由此可见,围栏封育在一定时间范围内可以显著改善群落的特征,增加群落的生产力,对于本研究地本氏针茅群落的最佳封育期限为10~15年,但围封时间过长在草地退化方面有一定程度的影响。

九寨沟湖泊湿地植物群落与植物地理研究

九寨沟湖泊湿地在维持九寨沟的生态平衡中起着重要的作用，在旅游产业的发展下，湿地生态系统及生物多样性面临着较大的威胁。尽管九寨沟湿地具有重要的生态价值，但目前对其研究尚比较薄弱。湿地植物群落和植物地理研究可以为湿地资源的可持续利用和监测保护提供科学依据。作者从2004年8月到2007年11月对九寨沟湿地的植物物种组成、地理分布、优势植物群落的结构、生长动态、湿地土壤种子库进行了调查研究。主要结果如下： 1. 九寨沟湿地物种组成、地理分布特点及湿地植物群落特点 九寨沟湿地共有苔藓植物8科13属16种，维管植物为48科107属199种。九寨沟湿地植物的地理成份较为丰富，维管植物在科级水平上有7种地理分布型（变型），在属级水平上有13种地理分布型（变型）， 在种级水平上共有29种地理分布型（变型）。九寨沟湿地植物以温带成份和我国特有成份为主，同时兼有热带、亚热带成份和环极—高山成份。九寨沟湿地植物的分布表现出明显的垂直地带性和水平地带性。湿地植物群落可划分21个群落类型，不同植物群落类型的物种多样性及物种组成存在较大的差异。九寨沟湿地植物的物种多样性和群落多样性以及较高的生产力特征，是维持其湿地生态景观多样性和稳定性的基础。 2. 土壤、水环境、海拔等对湿地植物的分布及生物多样性的影响 九寨沟湿地土壤、水等环境因子存在较大的差异。帕米尔苔草和宽叶香蒲等群落的凋落物较多，土壤有机碳、土壤总磷较高，可能是九寨沟湿地的重要土壤碳库。 九寨沟湿地植物沿水环境梯度的分布规律表现为：沉水植物（轮藻—篦齿眼子菜，水苦荬，杉叶藻）——挺水植物（水木贼，芦苇，宽叶香蒲）——湿生草本（苔草、节节草、披散木贼）——湿生灌木（柳灌丛，小檗灌丛）等。海拔也影响湿地植物的物种组成。 水深对物种多样性有影响，水深与物种丰富度负相关。随着水深的增加，水木贼、芦苇、杉叶藻、宽叶香蒲等群落的物种多样性下降；在长期淹水和季节性淹水的地方，水木贼群落物种多样性存在显著差异。土壤总氮与水木贼群落物种丰富度正相关。 3. 土壤营养元素、水环境对植物生长的影响 水深影响湿地植物生物量的分配。芦苇无性系分株在47 cm水深的环境中单株平均生物量最大；在干滩地中（地面水深0 cm），叶生物量百分比最大，而茎生物量百分比最小，茎的生物量百分比和生长速率随水深的增加而增加；在较干的滩地生境中，开花率、花序的生物量百分比明显大于水较深的生境。 水深与水木贼地上生物量负相关，但水木贼地上生物量在长期淹水和季节性淹水的地方没有显著的差异。在水浅的地方，杉叶藻、水木贼、芦苇等植物群落中，其他伴生物种的生物量占样方总生物量的百分比较大。 土壤有机碳、土壤总氮、土壤总磷等对湿地植物生物量的影响比较大：宽叶香蒲地上生物量与土壤总磷正相关；水木贼地上生物量与土壤总氮正相关；杉叶藻地上生物量与土壤有机碳正相关。 水深、土壤营养成分对湿地植物高度、密度等有影响。水木贼的平均高度在季节性淹水的地方比长期淹水的地方低，平均密度在长期淹水的地方比季节性淹水的地方低；除了5月份，其他观察月份水木贼的密度都与水深负相关，同时与土壤有机碳正相关。另外，芦苇密度与土壤有机碳含量正相关，宽叶香蒲密度与水深负相关，帕米尔苔草高度与土壤有机碳负相关。 4. 优势植物群落的动态变化 在优势植物群落中，优势种的高度、密度、盖度、生物量等在群落中占绝对优势。除五花海，水木贼群落的物种组成、高度、生物量在两年间没有显著的变化。芦苇群落的物种丰富度在近两年有所增加。 湿地植物生长表现为明显的季节动态，生长的峰值大多在7月－8月。优势植物群落的物候与水文周期有关。湿地植物群落的物种组成和密度，可以作为对湿地监测和保护的生物指示。 5. 九寨沟湿地土壤种子库特征及其在湿地生物多样性恢复中的作用 水深和现存植被物种丰富度可以解释湿地土壤种子库的变化。水深可以解释表层物种丰富度45%的变化。现存植被物种丰富度可以分别解释10 cm土层、2－5 cm土层及5－10 cm土层土壤种子库45%、48%和25%的变化。 湿地土壤种子库的密度为0－15945粒m-2, 种子库中共发现23个物种。现存植被优势物种和种子库优势物种不同。各层土壤种子库密度和物种丰富度并不存在显著的差异，但第二层土壤种子库密度最大。海拔、现存植被优势种盖度、土壤总磷、土壤总氮、土壤有机碳对湿地土壤种子库的密度和垂直结构没有影响。土壤种子库物种丰富度小于地上植被物种丰富度。湿地土壤种子库与地上植被的相关性不大。在浅水区域，湿地土壤种子库在湿地植被恢复中有一定作用。但在深水区域，保护现存植被更重要。 The lakeshore wetlands are valuable ecological units of the Jiuzhaigou lakes. Pressure for travel industry development pose a continuing and severe threat to the biodiversity-support function of the wetland system. Despite the ecological importance of wetlands in Jiuzhaigou, they are so far poorly studied. Both general plant communties and biogeographical studies are needed in order to attain basis for sustainable use the wetland resources and adequate protection of these areas. The present study was undertaken to examine aquatic plants distribution and the species compositon, structure and growth dynamics of their communities with variations of environmental factors along altitudes, water depth and soil properities gradients in Jiuzhaigou. Analysis of field survey data collected during August 2004 and November 2007 in lakeshore wetlands in Jiuzhaigou National Nature Reserve, Sichuan, China. The results were as following: (i) Species composition and biogeography in wetland vegetation 8 families, 13 genus, 16 species of moss and 48 families, 107 genus and 199 species of vascular plants in Jiuzhaigou wetlands were found. The floristic compositions were abundunt. Ten geographical distribution types at family level, 13 geographical distributions types at generic level and 29 geographical distribution types at specific level in vascular plants were found. Most species in Jiuzhaigou wetlands are temperate elements and Chinese endemic elements, with a few of tropical and subtropical and some circumarctic elements. And the plant distributions show clear vertical and horizontal patterns. There were 21 major wetland plant community types. Species composition and species richness in different plant communities are different. The species diversity and plant community diversity and their high biomass are the basis for the diversity and stability of wetland landscapes in Jiuzhaigou. (ii) Water depth, soil nutrients and altitudes influence on the species diversity and plant distribution. Total phosphorous and organic cabon in soil were higher in C. pamiernensis and T. latifolia communities, where are important cabon reservoirs in Jiuzhaigou wetlands. Along gradients of water depth, among populations of the dominant plant species present: submerged macrophytes (Chara vulgaris, Potagemonton pectinatus, Veronica anagalis-aquatica，Hippuris vulgaris), emergent macrophytes (Equisetum fluviatile, Phragamites australis, Typha latifolia), helophytes (Carex pamirensis )and shrubs (Salix sp., Berberis sp. ). Altitudes influence on the assemblage of plant communities. Water depth negatively correlated with species richness. Specie richness showed differences between permanently flooded sites and seasonally flooded sites in E. fluvatile communities. And total nitrogen in soil was negatively correlated with species richness in E. fluviatile communities. Altitudes show no significant influence on species richness, but in fact, through our analyses, they do have influence on the assemblage of wetland plants. (iii) Water depth, soil nutrients influence on the plant growth Water depth influences the biomass allocation in Phragmities australis. The average aboveground biomass of a single ramet (4.2 g) was the largest in the habitat with water level 47 cm above the soil surface. At the habitat with water level under soil surface 15 cm (－15 cm), the leaf biomass percentage (of the total ramet biomass) was the largest (46.1%), and the height and percentage of ramose ramets ( with branches on stem )(of the total ramets in a plot) were found obviously different. The deeper in water, the larger the biomass percentage and growth rate of stems were. The flowering rate and biomass of panicles were greater in shallow water than those in deep water. Water depth negatively correlated with aboveground biomass of E. fluviatile. However, above-ground biomass of E. fluviatile showed no significant difference between permanently flooded sites and seasonally flooded sites. But in shallow water, more biomasses of accompanying species were found in dominant plant communities such as H. vulgaris communities, E. fluviatile communities and P. australis communities. Water depth, soil nutrients influence on shoot density and shoot length of wetland plants. The shoot density of E. fluviatile was correlated to water depth in all growth months. Annual average density was significantly lower at permanently flooded sites than at seasonally flooded sites. But the annual average shoot length was significantly lower at seasonally flooded sites than at permanently flooded sites. (iv) Growth dynamics of dominant communities in Jiuzhaigou wetland The shoot length and shoot density, coverage and biomass of domiant species were dominated in plant communities. The species composition increased in P. australis communities in recent two years. The species richness in E. fluviatile communities showed no difference between 2005 and 2007. The above-ground biomass and shoot density in Five－flower Lake from July 2005 to July 2007 were significantly different, while in other sites, the differences were not significant. Shoot height, shoot density and above-ground biomass showed significant seasonal changes in all sites. Growth dynamics correlated with the cycle of water levels in lakes. Most plants growth parameters peaked at July or August. The biomass of T. latifolia peaked in August. But the shoot length of T. latifolia in deeper water peaked in July. The shoot length of E. fluviatile increased significantly from May to August except in seasonally flooded sites in Arrow-bamboo Lake. The species composition of communities and shoot density can be used as bioindicators in Jiuzhaigou wetland. (v) Soil seed bank in Jiuzhaigou wetland and its role in vegetation restoration Seed density in all soil layer samples was negatively correlated to water depth. Water depth can explain 45% variance of species richness in surface layer in sediment. Species richness in extant vegetation can explain 45%, 48%, 25% variance of species richness in total 10 cm and in 2－5 cm and 5－10 cm layer sediment respectively. Mean seed densities in wetlands ranged from 0 to 15945 m–2. A total of 23 species germinated in seed bank. The dominant species in seed bank and extant vegetation showed great difference. The total number of species and seedlings that germinated in different layers was not significantly different. But the second layer had the greatest seed density. In shallow water, seed bank can contribute to vegetation restoration, while in deeper water, protection of extant vegetation may be a better strategy.

生姜化感作用机理及其栽培模式研究

近年来，随着对作物重茬（连年种植）障碍原因的深入研究，植物的化感作用越来越受到国内外众多学者的重视。而作为重要调料和药用植物的生姜，其连作障碍也备受关注，系统地研究生姜化感作用将有助于理解和最终解决生姜连作障碍问题。本文通过研究生姜不同部位、不同浓度的水浸液对与其间作的两个物种（大豆和四季葱）种子的萌发及幼苗生长的影响，从而证明生姜化感作用的存在；并通过温室盆栽实验研究了生姜的自毒作用（即研究生姜不同部位、不同浓度的水浸液对其幼苗的形态、生理生化、光合作用、土壤酶、土壤微生物多样性及土壤养分的影响），从而揭示生姜退化和衰老的机制，并为生姜筛选出合适的间作物种提供科学依据，对生姜连作障碍提出科学的解决方法。主要研究结果如下： 1. 与对照相比，生姜所有部位（根茎、茎、叶）、所有浓度（10、20、40、 80 g l-1）的水浸液均抑制了大豆种子和葱籽的萌发率、幼苗生长、水分吸收和脂肪酶活性，并且其抑制程度随着水浸液浓度的增加而增强，其生姜各部位水浸液抑制效应的强弱顺序为茎＞叶＞根茎。这一结果表明生姜根茎、茎、叶含有能够抑制大豆种子和葱籽种子萌发和幼苗生长的水溶性化感物质。根茎是生姜的主要收获部位，而生姜的残株（主要是茎和叶）应该从大田中处理掉以减轻其抑制效应。生姜水浸液中主要化感成分包括：根茎水浸液中主要是丁香酸和伞花内脂；茎水浸液中主要是阿魏酸，且其含量最高为73.4 ug/g；叶水浸液中除了阿魏酸，其他六种物质均检测出来，但含量较高的主要有丁香酸、伞花内脂和香豆酸。 2. 生姜茎和叶不同浓度的水浸液均显著抑制了生姜幼苗的株高、每株叶片数和叶面积，其抑制程度随着水浸液浓度的增加而有所增强，而生姜幼苗每株分枝数差异不显著；同时生姜水浸液也极大程度地影响了生姜幼苗的生物量（包括地下生物量、地上生物量和总生物量，均为鲜重）。在同一浓度下，茎水浸液对生姜幼苗形态指标及生物量指标均显示出最强的抑制作用，叶水浸液次之，根茎水浸液最弱。与对照相比，低浓度的生姜根茎水浸液提高了生姜幼苗叶片内四种抗氧化酶（SOD、POD、CAT、APX）活性，高浓度的根茎水浸液抑制了四种抗氧化酶活性，而茎和叶水浸液均随着浓度的增加而抑制了四种抗氧化酶活性，三种水浸液均随着浓度的增加降低了生姜幼苗叶片内叶绿素的含量，而增加了生姜幼苗叶片的相对电导率和丙二醛含量。同时，三种水浸液均随着浓度的增加降低了生姜幼苗的光合参数（包括胞间CO2浓度、气孔导度、蒸腾速率及净光合速率）。 3. 三种生姜水浸液对所测六种土壤酶活性均产生了不同程度的影响，其中影响最大的是酸性磷酸酶和蔗糖酶，在10 g l-1 时就达到了显著水平，并且所有酶均有随着水浸液浓度增加而增大的趋势；相同部位的水浸液随着浓度的增加，细菌和真菌的数量呈增加趋势，而放线菌的数量呈减少趋势；三种生姜水浸液均随着浓度的增加降低了土壤中有机质的含量，加剧了土壤中硝态氮含量的积累，根茎水浸液对土壤有效磷、速效钾和铵态氮均显示出低浓度提高其含量而高浓度降低其含量的趋势，而茎和叶水浸液则随着浓度的增加均降低了其含量。 4. 与生姜单作相比，所有间作系统均在旺盛生长期和收获期不同程度地提高了土壤酶活性，同时也增加了土壤细菌数量及土壤微生物总数但不显著；所有间作系统在旺盛生长期和收获期均不同程度地影响了土壤真菌及放线菌数量（增加或减少），所有间作系统间的多样性指数差异不显著，除了旺盛生长期四种作物（生姜-大豆-四季葱-大蒜）的间作模式显著降低了多样性指数，其值仅为生姜单作的33.18%；生姜与大豆间作不仅提高了19.6%的生姜产量而且获得了较好的经济效益，并且，所有间作系统均显著抑制了生姜姜瘟病的发生。 5. 不同栽培模式不同程度地影响了收获期生姜的株高、分枝数、根茎产量及内在品质。其中处理2显著地促进了生姜的分枝（10.5%），同时处理2、3和4也促进了生姜的生长（株高分别增加了15.0%、11.4%和14.0%），并且这三个处理提高了生姜的产量；处理2和3能有效提高生姜块茎中维生素C（分别较单作生姜显著提高了3.29%和4.05%）、处理3显著提高了可溶性糖（8.2%）、姜辣素（4.6%）和蛋白质等有益物质的含量，降低硝酸盐有害物质的含量（处理2显著降低了14.0%），改善了姜块的外观和内在品质。并且，生姜与大豆间作具有最高的纯收入和产投比，分别较生姜单作提高了24.80%和8.8%。Recently, allelopathy has been more and more paid attentions by national and foreign scholars with profound research on reasons of crop replanted (continuous planted) obstacle. Ginger rhizome is valuable all over the world either as a spice or herbal medicine and ginger replanted obstacle is also paid attentions. Systematic research on ginger allelopathy will contribute to understanding and ultimate solving problem of ginger replanted obstacle. The effects of ginger aqueous extracts with different parts and concentrations on seed germination and early seedling growth of soybean and chive were studied in this article to testify that ginger existed allelopathy. Furthermore, ginger autotoxicity was also studied by pot experiment in greenhouse (namely research on effects of ginger aqueous extracts with different parts and concentrations on morphological indexes, physiological and biochemical indexes, photosynthesis, soil enzymes, soil microbial diversity and soil nutrients) to reveal mechanism of ginger degeneration and senescence, provide scientific basis for selecting appropriate intercropping species and put forward scientific resolvent for ginger replanted obstacle. The main results were as follows: 1. All aqueous extracts at all concentrations inhibited seed germination, seedling growth, water uptake and lipase activity of soybean and chive compared with the control, and the degree of inhibition increased with the incremental extracts concentration. The degree of toxicity of different ginger plant parts can be classified in order of decreasing inhibition as stem＞leaf＞rhizome. The results of this study suggested that rhizome, stem and leaf of ginger contained water soluble allelochemicals which could inhibit seed germination and seedling growth of soybean and chive. The rhizome is the main harvested part of ginger. The residue (mainly stems and leaves) of the ginger plant should be removed from the field so as to diminish its inhibitory effect. The main allelopathic components of three kind of aqueous extracts were as follows: Rhizome extract chiefly contained syringic acid and vmbelliferone and stem extract mainly contained frulic acid whose content was the highest (73.4 ug/g). The other six substances were detected except of frulic acid, but only contents of syringic acid, vmbelliferone and p-coumaric acid were higher. 2. Stem and leaf aqueous extracts of ginger with different concentrations significantly inhibited plant height, leaf numbers per plant and leaf area, and the degree of inhibition increased with the incremental extracts concentration. However, tiller number per plant of ginger seedling showed no significant difference. At the same time, ginger aqueous extracts also influenced biomass including under-ground biomass, above-ground biomass and total biomass (fresh weight) to a large extent. Under the same concentration, stem aqueous extract showed the mostly inhibitory effect on morphological indexes and biomass indexes of ginger seedling. Rhizome aqueous extract showed the leastly inhibitory effect and leaf aqueous extract was intervenient. Enhanced concentration of ginger aqueous extracts significantly reduced total chlorophyll content, accompanying with increases in memberane permeability (REL) and lipid peroxidation (MDA). Compared with the control, rhizome ginger aqueous extract of lower concentration (10 g l-1) increased the activities of major antioxidant enzymes (superoxide dismutase, SOD; peroxidase, POD; catalase, CAT; ascorbate peroxidase, APX) of ginger leaf tissue and higher concentration inhibited the activities of four antioxidant enzymes. However, stem and leaf aqueous extract inhibited the activities of four antioxidant enzymes with increase in concentration. Meanwhile, enhanced concentration of ginger aqueous extracts significantly reduced photo-parameters of ginger seedling (including CO2 concentration, stoma conductivity, net photosynthesis rate and transpiration rate). 3. Rhizome, stem and leaf ginger aqueous extract showed different effect on six soil enzyme activities, and acid phosphatase and invertase showed significant effect when aqueous extract concentration got 10 g l-1. Furthermore, six soil enzyme activities increased with increase in aqueous extract concentration. Bcterial and fungi number tended to increase while antinomyces tented to decrease with the increase in aqueous extract concentration of identical part. Ginger aqueous extracts reduced soil organic matter content with increased concentration, accompanying with NO3-—N accumulation in soil. Rhizome aqueous extract showed the same tendency for available P, available K and NH4+—N, namely lower concentration increased their contents in soil and higher concentration reduced their contents. While stem and leaf aqueous extracts reduced their contents with the increamental concentration. 4. All intercropping systems increased soil enzyme activities to different extent both at VGS and at HS compared to solo ginger. All intercropping systems increased the colony numbers of soil bacteria and total of soil microbe but not significantly either at VGS or at HS. All intercropping systems increased the colony numbers of soil fungi and actinomytes to a different extent (increase or decrease) both at VGS and at HS. For DI, difference between all cultivation patterns and S-G was not significant either at VGS or at HS except that G-S-C-G whose value was only 33.18% of S-G at VGS significantly decreased. G-S not only increased ginger yield by 19.6% but also obtained better economic benefit. Furthermore, all intercropping systems significantly inhibited occurrence of bacterial wilt of ginger. 5. Different cultivated pattern influenced plant height, tiller numbers, rhizome yields and intrinsic quality of ginger. Treatment 2 significantly facilitated tiller occurring (10.5%). Treatment 2, 3 and 4 promoted ginger growth (plant height respectively increased 15.0%、11.4% and 14.0%) and enhanced rhizome yields. Treatment 2 and 3 effectively increased vitamin C content (significantly increased 3.29% and 4.05% compared to solo ginger). Treatment 3 significantly increased contents of beneficial substances such as soluble sugar (8.2%), gingerols (4.6%) and protein. Treatment 2 significantly decreased contents of deleterious substance namely nitrate (14.0%) and improved appearance and intrinsic quality of ginger rhizome. Furthermore, treatment 2 (ginger/soybean intercropping) could obtain better economic benefit and showed the highest net income and ratio of benefit and cost whose values respectively increased by 24.80% and 8.8% compared to solo ginger.

川西北高寒草甸土壤呼吸研究：季节动态和放牧效应

除植被冠层的光合作用之外，土壤的呼吸作用是陆地生态系统碳收支中最大的通量。土壤呼吸即使发生较小的变化也能显著地减缓或加剧大气中CO2浓度的增加，从而明显影响到全球气候变化。土壤呼吸速率变化与否以及变化的方向可以反映生态系统对环境变化的敏感程度和响应模式。尽管如此，土壤呼吸仍是一个为人们了解不多的生态系统过程。 草地生态系统是陆地生态系统的一个重要组成部分。针对草地土壤呼吸进行野外实验研究和相应方法论的探讨将对区域乃至全球碳源汇性质的准确估算具有重要的科学意义。然而，近几年来关于草地土壤呼吸的主要研究工作都集中在温带草原和部分热带草原，而针对高寒草甸生态系统土壤呼吸的研究报道还很少。 2008年4月至2009年4月期间，我分别在2008年6、8、10、12月和2009年2月和4月分6次对川西北的典型高寒草甸群落的土壤呼吸进行观测，分析了不同类型高寒草甸群落土壤呼吸的季节变化特征以及环境因子和放牧模式对其影响。主要研究结果如下： 1）该地区高寒草甸生态系统在生长季（6月～8月）土壤呼吸速率较大（6.07～9.30μmolCO2¡m-2¡s-1 ） ， 在非生长季（ 12 月～ 2 月） 较小（ 0.16 ～0.49μmolCO2¡m-2¡s-1 ） 。土壤CO2 年累积最大释放量为3963 ～ 5730gCO2¡m-2¡yr-1，其中，生长季土壤CO2的释放量占年总释放量的85%～90%。非生长季占10%～15%。非生长季所占比例略小于冬季积雪覆盖地区的冬季土壤呼吸占年土壤呼吸量的比例（14%～30%）。温度，尤其地温，是影响该地区高寒草甸生态系统土壤呼吸速率的最主要环境因子。土壤呼吸速率与地上生物量和土壤水分之间没有显著相关性，但是土壤含水量过大会导致土壤呼吸速率下降。 2）在观测期内，草丘区的土壤呼吸显著高于对照区的土壤呼吸，其最大土壤呼吸速率为16.77μmolCO2¡m-2¡s-1，土壤CO2 年累积最大释放量为8145gCO2¡m-2¡yr-1，是对照区的近2 倍。由于草丘在高寒草甸中占有较大的面积比例（近30%），因此，它将对高寒草甸生态系统的碳循环起着重要的作用。 3）放牧模式不仅可以影响高寒草甸群落的土壤CO2 排放，而且还可以改变土壤呼吸的温度敏感性（Q10）。本研究表明，在生长季有长期放牧活动干扰时将会增加土壤向大气中释放二氧化碳的速度，促使土壤碳库中碳的流失。禁牧样地的土壤呼吸速率在刚禁牧时先迅速增大，随着禁牧时间的延长土壤呼吸速率将会下降。此外，与其它放牧模式相比，冬季放牧将高寒草甸群落土壤呼吸速率在生长季达到最大值的时间明显向后推迟。不同放牧模式下高寒草甸群落土壤呼吸的Q10 值大小顺序为：禁牧一年群落>冬季放牧群落>禁牧三年群落>夏季放牧群落>自由放牧群落。 4）基于呼吸室技术的观测方法中，测量前的剪草处理可以明显改变该地区高寒草甸群落的土壤温度和土壤呼吸速率。在生长季，剪草处理将使土壤呼吸速率的瞬时响应增加90%左右。由于剪草处理明显增加了剪草样方白天的土壤温度，而土壤温度与土壤呼吸之间存在着极显著的指数相关关系，因而剪草处理导致土壤呼吸速率迅速增加。因此，在高寒地区基于呼吸室技术观测的土壤呼吸应当进行校正。 综上所述，川西北高寒草甸生态系统土壤呼吸速率在生长季较高，而在非生长季较低。土壤温度是影响该地区土壤呼吸的最主要环境因子。在实验观测期，草丘区土壤呼吸速率显著高于对照区的，是对照区土壤呼吸速率的近2倍。由于测量前的剪草处理可以明显改变待测点的土壤呼吸速率，因此，应对在高寒地区基于呼吸室技术观测的土壤呼吸进行校正。 Soil respiration is the second largest component (less than plant phtotosynthesis) of carbon dioxide flux between terrestrial ecosystems and the atmosphere. A minor change in soil respiration rate can significantly slow down or accelerate the increase of atmospheric CO2 concentration that is closely related to global climatic change. In turn, the change in the flux direction and rate of soil respiration may indicate the elasticity and stability of ecosystems to global changes and human disturbance. However, soil respiration is still an ecosystem process that has been poorly understood. Grassland ecosystem is an important component of the terrestrial ecosystem. Accurately estimating the CO2 flux from soil to atmosphere in situ is the key to evaluating the carbon resource and sink regionally or globally. Despite of extensive studies on the temperate and tropic grasslands, the soil respiration of alpine meadows has not substantially been measured. In the current study, soil respiration was measured for an annual cycle from April, 2008 to April, 2009 for the alpine meadow in northwestern Sichuan Province of China to determine the seasonal variation of soil respiration for the typical plant communities. The results are shown as follows: 1) Large seasonal variation of soil respiration was observed in the alpine meadow. The rate of soil respiration was the greatest (6.07～9.30μmolCO2¡m-2¡s-1) in June and the smallest (0.16 ～ 0.49μmolCO2¡m-2¡s-1) occurred from December to February in the non-growing season. The total emission of soil CO2 was estimated as 3963～5730 gCO2¡m-2¡yr-1, 85%~90% of which was released during the growing season, and 10%~15% during the non-growing season which was slightly less than the ratio of winter and annual CO2 flux from soil. Temperature, particularly the soil temperature, was the major environmental factor regulating the soil respiration. Significant and positive relationships were not found between soil respiration and soil moisture and between soil respiration and plant above-ground biomass, but excessive soil water content would decrease in the rate of soil respiration. 2) The rate of soil respiration in grass hummock communities was up to 16.77μmolCO2¡m-2¡s-1, which was about twice as great as in the controls (communities located in low and even sites). Considering the large proportion (about 30% on average) of hummock area in the meadow, it can be concluded that the hummocks played an important role in the carbon cycling of the study ecosystem. 3) Grazing patterns affected the flux of CO2 emission and the temperature sensitivity of soil respiration (Q10) in the alpine meadow. Grazing during growing season increased the rate of soil respiration. The rate of soil respiration increased significantly immediately after the alpine meadow being fenced, but thereafter decreased. In addition, grazing in winter delayed the peak respiration rate relative to the non-grazing mode. The Q10 value was the largest in the non-grazed area for one year, and next came the area with grazing in winter, followed by the non-grazed area for three years, the area with grazing in summer, and the non-limited grazed area. 4) In the chamber-based techniques, clipping manipulation before each measurement increased the transient rate of soil respiration by about 90% in the summer of the alpine meadow. As increase in soil temperature at daytime in the clipped plots by clipping and the exponential relationship between soil respiration and temperature, clipping manipulation led to increase in the rate of soil respiration. This suggested that a correction should be done for the techniques if employed in alpine and cold regions. In summary, the rate of soil respiration in the alpine meadow was the greatest in June and the smallest occurred from ecember to February in the non-growing season. Soil temperature was the major environmental factor regulating the soil respiration. The rate of soil respiration in grass hummock communities was up to 16.77μmolCO2¡m-2¡s-1, which was about twice as great as in the controls. A correction should be done for the techniques if employed in alpine and cold regions, because of the effect of clipping manipulation on soil temperature and respiration.

黄土丘陵沟壑区退耕地植被自然恢复过程——以吴起县双树沟流域为例

采用TWINSPAN分类法对黄土丘陵沟壑区吴起县双树沟流域30个自然恢复草地植被进行分类,并对分类后各植被群落特征和地上生物量进行统计分析。结果表明:在自然恢复条件下,随着退耕年限的不断增加,退耕地植被自然恢复依次经历了猪毛蒿群落—赖草+长芒草群落—赖草+铁杆蒿群落—铁杆蒿群落—铁杆蒿+茭蒿群落5个发展阶段,地带性植被类型铁杆蒿+茭蒿群落在研究区内开始出现,并且已经占有一定优势;随着退耕地植被自然恢复的不断进行,Margalef等丰富度指数以及Shannon-wiener等多样性指数、Pielou等物种均匀度指数和地上生物量都呈现出先减小后增大的发展趋势;在植被自然恢复的稳定阶段,虽然物种丰富度指数和物种多样性指数有一定的增加,但是相对恢复初期来讲还是有所下降,并且有达到与初期相当水平的趋势;物种丰富度指数均在第1恢复阶段最大,而均匀度指数Jsw以及Shannon-wiener指数在第5恢复阶段最高。随着退耕地植被自然恢复的不断进行,植被群落总盖度随着退耕年限的延长而不断增大。

The effect of land management on plant community composition, species diversity, and productivity of alpine Kobersia steppe meadow

Grassland degradation is widespread and severe on the Tibet Plateau. To explore management approaches for sustainable development of degraded and restored ecosystems, we studied the effect of land degradation on species composition, species diversity, and vegetation productivity, and examined the relative influence of various rehabilitation practices (two seeding treatments and a non-seeded natural recovery treatment) on community structure and vegetation productivity in early secondary succession. The results showed: (1) All sedge and grass species of the natural steppe meadow had disappeared from the severely degraded land. The above-ground and root biomass of severely degraded land were only 38 and 14.7%, respectively, of those of the control. So, the original ecosystem has been dramatically altered by land degradation on alpine steppe meadow. (2) Seeding measures may promote above-ground biomass, particularly grass biomass, and ground cover. Except for the grasses seeded, however, other grass and sedge species did not occur after seeding treatments in the sixth year of seeding. Establishment of grasses during natural recovery treatment progressed slowly compared with during seeding treatments. Many annual forbs invaded and established during the 6 years of natural recovery. In addition, there was greater diversity after natural recovery treatment than after seeding treatments. (3) The above-ground biomass after seeding treatment and natural recovery treatment were 114 and 55%, respectively, of that of the control. No significant differences in root biomass occurred among the natural recovery and seeded treatments. Root biomass after rehabilitation treatment was 23-31% that of the control.

Effect of long-term grazing on soil organic carbon content in semiarid steppes in Inner Mongolia

To clarify the response of soil organic carbon (SOC) content to season-long grazing in the semiarid typical steppes of Inner Mongolia, we examined the aboveground biomass and SOC in both grazing (G-site) and no grazing (NG-site) sites in two typical steppes dominated by Leymus chinensis and Stipa grandis, as well as one seriously degraded L. chinensis grassland dominated by Artemisia frigida. The NG-sites had been fenced for 20 years in L. chinensis and S. grandis grasslands and for 10 years in A. frigida grassland. Above-ground biomass at G-sites was 21-35% of that at NG-sites in L. chinensis and S. grandis grasslands. The SOC, however, showed no significant difference between G-site and NG-site in both grasslands. In the NG-sites, aboveground biomass was significantly lower in A. frigida grassland than in the other two grasslands. The SOC in A. frigida grassland was about 70% of that in L. chinensis grassland. In A. frigida grassland, aboveground biomass in the G-site was 68-82% of that in the NG-site, whereas SOC was significantly lower in the G-site than in the NG-site. Grazing elevated the surface soil pH in L. chinensis and A. frigida communities. A spatial heterogeneity in SOC and pH in the topsoil was not detected the G-site within the minimal sampling distance of 10 m. The results suggested that compensatory growth may account for the relative stability of SOC in G-sites in typical steppes. The SOC was sensitive to heavy grazing and difficult to recover after a significant decline caused by overgrazing in semiarid steppes.

西藏高原草原化嵩草草甸生态系统呼吸及碳平衡

The soil respiration and net ecosystem productivity of Kobresia littledalei meadow ecosystem was investigated at Dangxiong grassland station, one grassland field station of Lhasa Plateau Ecosystem Research Station. Soil respiration and soil heterotrophic respiration were measured at the same time by using Li6400-09 chamber in growing season of year 2004. The response of soil respiration and its components, i.e. microbial heterotrophic respiration and root respiration to biotic and abiotic factors were addressed. We studied the daily and seasonal variation on Net Ecosystem carbon Exchange (NEE) measured by eddy covariance equipments and then the regression models between the NEE and the soil temperature. Based on the researches, we analyzed the seasonal variation in grass biomass and estimated NEE combined the Net Ecosystem Productivity with heterogeneous respiration and then assessed the whether the area is carbon source or carbon sink. 1.Above-ground biomass was accumulated since the grass growth started from May; On early September the biomass reached maximum and then decreased. The aboveground net primary production (ANPP) was 150.88 g m~" in 2004. The under-ground biomass reached maximum when the aboveground start to die back. Over 80% of the grass root distributed at the soil depth from 0 to 20cm. The underground NPP was 1235.04 g m"2.. Therefore annual NPP wasl.385X103kg ha"1, i.e.6236.6 kg C ha"1. 2. The daily variation of soil respiration showed single peak curve with maximum mostly at noon and minimum 4:00-6:00 am. Daily variations were greater in June, July and August than those in September and October. Soil respiration had strong correlation with soil temperature at 5cm depth while had weaker correlation with soil moisture, air temperature, surface soil temperature, and so on. But since early September the soil respiration had a obviously correlation with soil moisture at 5cm depth. Biomass had a obviously linearity correlation with soil respiration at 30th June, 20th August, and the daytime of 27th September except at 23lh October and at nighttime of 27th September. We established the soil respiration responding to the soil temperature and to estimate the respiration variation during monsoon season (from June through August) and dry season (May, September and October). The regression between soil respiration and 5cm soil temperature were: monsoon season (June through August), Y=0.592expfl()932\ By estimating , the soil daily respiration in monsoon season is 7.798gCO2m"2 and total soil respiration is 717.44 gCC^m" , and the value of Cho is 2.54; dry season (May, September and October), Y=0.34exp°'085\ the soil daily respiration is 3.355gCO2m~2 and total soil respiration is 308.61 gCC^m", and the value of Cho is 2.34. So the total soil respiration in the grown season (From May to October) is 1026.1 g CO2IT1"2. 3. Soil heterogeneous respiration had a strong correlation with soil temperature especially with soil temperature at 5cm depth. The variation range in soil heterogeneous respiration was widely. The regression between soil heterogeneous respiration and 5cm soil temperature is: monsoon season, Y=0.106exp ' 3x; dry season, Y=0.18exp°"0833x.By estimating total soil heterotrophic respiration in monsoon season is 219.6 gCC^m"2, and the value of Cho is 3.78; While total soil heterogeneous respiration in dry season is 286.2 gCCbm"2, and the value of Cho is 2.3. The total soil heterotrophic respiration of the year is 1379.4kg C ha"1. 4. We estimated the root respiration through the balance between soil respiration and the soil heterotrophic respiration. The contribution of root respiration to total respiration was different during different period: re-greening period 48%; growing period 69%; die-back period 48%. 5. The Ecosystem respiration was relatively strong from May to October, and of which the proportion in total was 97.4%.The total respiration of Ecosystem was 369.6 g CO2 m" .we got the model of grass respiration respond to the soil temperature at 5cm depth and then estimated the daytime grass respiration, plus the nighttime NEE and daytime soil respiration. But when we estimated the grass respiration, we found the result was negative, so the estimating value in this way was not close. 6. The estimating of carbon pool or carbon sink. The NPP minus the soil heterogeneous respiration was the NEE, and it was 4857.3kg C o ha"1, which indicated that the area was the carbon sink.

水分亏缺和凋落物水浸液对云杉幼苗生长和种子萌发的影响

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.

模拟增温与UV-B 辐射增强对云杉种子萌发和幼苗生长的影响

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.

干旱胁迫与施N条件下白刺花（Sophora davidii）幼苗生长及其适应机制

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.

Effect of plateau zokors on vegetation characteristics and productivity of alpine meadow

This research was conducted on alpine meadow site at Menyuan county, Qinghai Province, People's Republic of China to determine the effects of native, subterranean rodent of Qinghai-Tibet grasslands, the plateau zokors (Myospalax baileyi), on seasonal above-and below-ground plant biomass, plant species diversity and productivity. Both total peaks of above-and below-ground biomass were the greatest (413.600 g/m~2 and 2297.502 g/m~2) in the patch no any plateau zokors colonized by plateau zokors over 10 years in August and October, respectively. Both above-and below-ground biomass were significantly increased in the patches where plateau zokors were removed or the burrow systems were abandoned for five years compared to the patches plateau zokors colonized over 10 years. However, both above-and below-ground biomass in abandoned patches were significantly lower than that in uncolonized patches. Monocotyledonous biomass was reduced greatly, but the non-palatable dicots were significantly increased in colonized patches. The palatable biomass of monocots and dicots were increased in abandoned patches. Total plant species diversity was the greatest in uncolonized patchesand least in abandoned patch. The total net primary production in colonized patches was reduced by 68.98% compared with uncolonized patches. Although the patches were without any plateau zokors disturbance for fives years, the total net primary production just reached 58.69% of the uncolonized patches. The above-ground net primary production in abandoned patches increased 28.74% and the below-ground increased 54.91% compared with the colonized patches. We suggest that plateau zokor-induced changes in plant above- and below-ground biomass and species diversity may lead to further alterations of nutrient cycling and trophic dynamics in this alpine meadow ecosystem.