丛枝菌根真菌群落沿高寒草原海拔梯度的变化特征

基于丛枝菌根(Arbuscular mycorrhizal,AM)真菌孢子形态学的鉴定,对沿不同海拔(4584、4628、4744、4880、4956 m)梯度采集的高寒草原建群植物根际土壤样品进行了分析。结果表明,高寒草原AM真菌属、种构成均较简单,Acaulospora、Claroideoglomus、Funneliformis、Glomus属见于各海拔梯度,海拔4744 m地带未见Pacispora属,海拔4744、4956 m地带无Scutellospora属分布,Rhizophagus属仅见于海拔4584 m地带。随海拔上升,AM真菌种数、物种丰度均呈显著下降;海拔4584—4880 m范围Shannon-Weiner指数(H)无显著差异,但在最高海拔时显著下降;优势种种数及所占比例与海拔梯度则呈显著正相关(Funneliformis geosporum、Claroideoglomus claroideum为不同海拔梯度优势种);沿海拔梯度,孢子密度基本呈单峰分布格局,峰值出现在海拔4744 m地带;海拔梯度对菌根侵染效应影响显著,菌根侵染率、侵染强度和丛枝丰度随海拔上升均呈显著下降趋势;不同海拔梯度高寒草原AM真菌群落相似度(Sorensen相似性系数0.821—0.969)较高,并在总体上表现出随海拔梯度增大而降低的趋势。土壤pH值、有效磷、有机碳、海拔对AM真菌的群落分布均产生显著影响,尤以海拔的影响最为显著。研究结果对预测高寒草原微生物的作用与影响,以及高寒草原环境对全球变化的响应等提供了理论依据。     

青藏高原高寒草地AM真菌分布及其对近自然恢复的生态作用

超载过牧以及全球气候变化等导致大部分青藏高原高寒草地呈现持续退化态势。青藏高原高寒草地退化致使地上植物群落逐渐发生更替,地下土壤微生物群落多样性和丰富度发生改变。本文旨在探析青藏高原高寒草地丛枝菌根（arbuscular mycorrhizal,AM）真菌的分布特征、对近自然恢复的生理生态效应及其作用机制。青藏高原高寒草地中已报道4目14属61种AM真菌,约占已知AM真菌物种的20%。高寒草地禾本科植物根围AM真菌物种丰度最高,而莎草科植物根围AM真菌孢子密度最高。3种高寒草地植被类型中,高寒草原AM真菌丰度最高（33种）,山地灌丛草原次之（32种）,高寒草甸最低（22种）。高寒草原以光壁无梗囊霉Acaulospora laevis和闪亮和平囊霉Pacispora scintillans为优势种,山地灌丛草原以摩西斗管囊霉Funneliformis mosseae为优势种,高寒草甸以光壁无梗囊霉A. laevis、近明球囊霉Claroideoglomus claroideum和闪亮和平囊霉P. scintillans为优势种。高寒草地土著AM真菌与植物构建的菌根网络可以通过调节营养元素吸收、分配,促进植物建植和生长;但是毒杂草入侵可以改变土著AM真菌物种多样性和菌根网络,限制本地植被的实际生态位扩张。退化高寒草地中,AM真菌群落具有高的环境适应性和恢复力,其不仅调控地上植物群落建植和多样性,同时AM真菌建植也增加了代谢产物-球囊霉素相关土壤蛋白产生,进而协同改善地下土壤微生态系统,为退化高寒草地早期植被恢复塑造土壤生境。因此,AM真菌在退化高寒草地近自然恢复中具有较大的应用潜力。     

西藏高原草地生态系统丛枝菌根真菌的地理分布

西藏高原是地球上极具特色的地理单元,对生物物种的形成与演化具有重要影响。基于孢子形态学鉴定,对从藏东南到藏西北(海拔高差3500 m,年均温、年均降水量差异分别20℃、800 mm)高原热带、亚热带、温带、亚寒带和寒带环境下发育而成的热性草丛、暖性草丛、温性草原、温性荒漠、高寒草甸草原、高寒草原、高寒荒漠和高寒荒漠草原中的AM真菌群落进行研究,结果表明,不同类型草地间AM真菌的群落相似度普遍较低,环境对AM真菌群落具有重要影响。从藏东南到藏西北,不同类型草地间AM真菌的群落相似度呈下降趋势(Jaccard相似性系数从0.52降至0.20),AM真菌群落组成及结构变化渐趋加大,不同草地中的同种植物(包括广谱种、青藏高原特有种)AM真菌群落相似度亦不同。沿藏东南到藏西北环境梯度,随草地寒旱程度的逐步加剧,AM真菌种的丰度,特别是种数、Shannon-Weiner指数在总体上趋于显著(P0.05)下降,孢子密度则在总体上趋于显著提高,优势种比例、Shannon-Weiner指数亦表现出增加的趋势,表明AM真菌物种多样性虽趋于下降,但生存及对环境的适应能力趋于提高。海拔、土壤p H、有效磷和有机碳含量对AM真菌群落组成均具显著影响,但海拔对水热环境的影响决定着土壤环境的变化。因此,海拔和寒旱程度不断上升所导致的p H显著提高、土壤有机碳和有效磷含量显著下降的综合作用影响并决定着AM真菌的群落组成。研究结果对进一步理解西藏高原生物多样性的产生和维持机制等具有重要的参考价值。     

Molecular analysis of arbuscular mycorrhizal fungi colonising a semi-natural grassland along a fertilisation gradient

The community of arbuscular mycorrhizal fungi (AMF) colonizing the roots of Festuca pratensis and Achillea millefolium was characterized in a Swedish pasture at different times, along a gradient of fertilization. The small subunit ribosomal RNA gene was subjected to PCR and denaturing gradient gel electrophoresis (DGGE), sequencing and phylogenetic analysis. The sequences found in this study clustered in 10 discrete sequence groups, seven belonging to Glomus, two to Scutellospora and one to Diversispora. A negative correlation was observed between soil mineral nitrogen and the number of AMF sequence groups in the roots. The frequency of occurrence of AMF in roots decreased dramatically between June and September. No plant-host specificity could be detected.     

Molecular diversity of arbuscular mycorrhizal fungi associated with two co-occurring perennial plant species on a Tibetan altitudinal gradient

Plant communities on Mount Segrila on the Tibetan Plateau show distinct changes at different altitudes, but little information is available on belowground communities of arbuscular mycorrhizal fungi (AMF). Root samples of two co-occurring species, Pennisetum centrasiaticum and Kobresia sp., growing in open grasslands at eight altitudes (3,446–4,556 m) were analyzed for diversity of AMF by PCR, cloning, and sequencing. Dominant plants were well colonized by AMF even at higher altitudes where spore density in rhizospheres decreased dramatically. A total of 29 operational taxonomic units (OTUs) of AMF were detected, and some novel sequence types were found. Acaulosporaceae and Glomeraceae were the dominant families. There was no significant difference in OTU richness along elevational gradients in Kobresia sp., but OTU richness in P. centrasiaticum was higher at intermediate elevations. Elevation, host plant species, and soil variables (pH, soil organic matter, and available P and N) were found to have significant effects on the overall AMF community across all elevations. Fungal community composition differed significantly between the two plant species at each elevation, and the similarity was generally higher at the intermediate elevations. No significant difference in compositional similarity was observed for Kobresia sp. with increasing elevation, but the dissimilarity increased significantly for P. centrasiaticum. These results suggest that host identity is an important determinant for the structure of the AMF communities along the elevational gradients in high altitude environments.     

Arbuscular mycorrhizal fungi associated with sedges on the Tibetan plateau

The arbuscular mycorrhizal (AM) status of nine dominant sedge species and the diversity of AM fungi in Tibetan grassland were surveyed in the autumn of 2003 and 2004. Most of the sedge species and ecotypes examined were mycorrhizal, but Carex moorcroftii and Kobresia pusilla were of doubtful AM status, and Kobresia humilis was facultatively mycorrhizal. This is the first report of the mycorrhizal status of eight of the nine sedge species examined. Intraradical vesicles and aseptate hyphae were the structures most frequently observed. Appressoria, coils, and arbuscules were found in the roots of a few sedge species. A strong negative correlation was found between soil organic matter content and the extent of mycorrhizal colonization. Using trap cultures, 26 species of AM fungi belonging to six genera, Glomus, Acaulospora, Paraglomus, Archaeospora, Pacispora, and Scutellospora, were isolated from the soil samples collected. The frequency of occurrence of different taxa of AM fungi varied greatly. Glomus and Acaulospora were the dominant genera, and Acaulospora scrobiculata was the most frequent and abundant species. The species richness of AM fungi was 2.73 in the study area. Species richness and diversity index differed among the sedge species but were not correlated with soil factors such as pH, available P, or organic matter content.     

西藏高山草原AM真菌生态分布

对西藏高原37种草地植物(建群种或常见种)70个带根土样的AM真菌进行了研究.结果表明:1)在所分离到的5属35种AM真菌中,球囊霉属、无梗囊霉属、盾巨孢囊霉属真菌分别为18、9、6种,内养囊霉属、类球囊霉属各1种.其中,藏南、藏北草原AM真菌分别为4属23种、4属22种,Shannon指数分别为2.31和2.75,但藏北草原AM真菌孢子密度、种的丰度显著高于藏南.2)不同生态区域AM真菌呈共有种较少、特有种较多、优势种差异较大的分布特征.3)高寒草原、山地草甸、高寒草甸草原AM真菌Shannon指数分别为1.91、1.83、1.80,呈严重退化态势的温性草原仅为1.64;海拔4000～4600 m地带,种的丰度最高,海拔4600～5220 m 地带,真菌Shannon指数和物种均匀度最高,分别为2.42和0.79;4)球-囊霉属真菌在不同海拔段均为优势属,但在海拔＜4000 m地带相对多度较高;无梗囊霉属主要分布于海拔＞4000 m地带;盾巨孢囊霉属主要分布于海拔3500～5220 m地带;类球囊霉属主要见于海拔4000～5220 m的藏北高寒草甸草原,少量见于高寒草原环境;内养囊霉属仅见于海拔3500～3700 m的藏南温性草原.     

丛枝菌根真菌对中性紫色土土壤团聚体特征的影响

利用分根装置研究了接种丛枝菌根真菌对中性紫色土土壤团聚体数量、大小和稳定性的影响,分析了土壤团聚体数量、分布和分形维数,并运用通径分析对不同影响因子进行了分析。结果表明:接种G. intraradices和G. mosseae的菌根室土壤有机质和总球囊霉素相关土壤蛋白含量有增加的趋势;湿筛条件下菌根室土壤的平均重量直径(MWD)和几何平均直径(GMD)均显著高于根室土壤,而且降低了土壤分形维数,显示了丛枝菌根真菌提升土壤结构特征的作用。通径分析的结果表明,在影响MWD的众多因子中,菌丝密度对MWD有最大的直接作用。两种菌种对土壤结构均有不同程度的改良作用,反映了丛枝菌根真菌功能的多样性。     

西藏高原不同海拔区域丛枝菌根真菌群落的变化

为了进一步了解丛枝菌根(AM)真菌群落对不同海拔环境的响应,基于孢子形态学鉴定,研究了西藏高原不同海拔区域主要草本植物AM真菌的群落特征、菌根侵染及其变化.结果表明: 藏东南低海拔区(2200~3400 m)、藏中中海拔区(3400~3900 m)和藏北高海拔区(4300~5300 m)的AM真菌分别为11属31种、11属20种和6属14种.随着海拔上升,孢子密度(r=0.978,P<0.01)、物种丰度(r=0.462,P>0.05)均趋提高,优势种、特有种比例大幅增加,Shannon指数(r=-0.945,P<0.01)极显著下降.不同海拔区之间AM真菌群落Sorensen相似性系数(0.526~0.592)较为接近,仅在总体上随海拔差异扩大略趋下降;藏东南低海拔区、藏北高海拔区菌根侵染率无显著差异,但均显著低于藏中中海拔区.各海拔区内,不同海拔梯度对AM真菌群落、根系侵染亦具显著影响,但影响程度、影响趋势因整体海拔环境不同而异.说明西藏高原AM真菌群落趋于生境选择,受控于海拔所主导的水热环境及土壤环境.     

No temperature acclimation of soil extracellular enzymes to experimental warming in an alpine grassland ecosystem on the Tibetan Plateau

Alpine grassland soils store large amounts of soil organic carbon (SOC) and are susceptible to rising air temperature. Soil extracellular enzymes catalyze the rate-limiting step in SOC decomposition and their catalysis, production and degradation rates are regulated by temperature. Therefore, the responses of these enzymes to warming could have a profound impact on carbon cycling in the alpine grassland ecosystems. This study was conducted to measure the responses of soil extracellular enzyme activity and temperature sensitivity (Q₁₀) to experimental warming in samples from an alpine grassland ecosystem on the Tibetan Plateau. A free air-temperature enhancement system was set up in May 2006. We measured soil microbial biomass, nutrient availability and the activity of five extracellular enzymes in 2009 and 2010. The Q₁₀ of each enzyme was calculated using a simple first-order exponential equation. We found that warming had no significant effects on soil microbial biomass C, the labile C or N content, or nutrient availability. Significant differences in the activity of most extracellular enzymes among sampling dates were found, with typically higher enzyme activity during the warm period of the year. The effects of warming on the activity of the five extracellular enzymes at 20 °C were not significant. Enzyme activity in vitro strongly increased with temperature up to 27 °C or over 30 °C (optimum temperature; T_opt). Seasonal variations in the Q₁₀ were found, but the effects of warming on Q₁₀ were not significant. We conclude that soil extracellular enzymes adapted to seasonal temperature variations, but did not acclimate to the field experimental warming.     

Seasonal variation of arbuscular mycorrhizal fungi in temperate grasslands along a wide hydrologic gradient

We studied seasonal variation in population attributes of arbuscular mycorrhizal (AM) fungi over 2 years in four sites of temperate grasslands of the Argentinean Flooding Pampas. The sites represent a wide range of soil conditions, hydrologic gradients, and floristic composition. Lotus glaber, a perennial herbaceous legume naturalised in the Flooding Pampas, was dominant at the four plant community sites. Its roots were highly colonised by AM fungi. Temporal variations in spore density, spore type, AM root colonisation, floristic composition and soil chemical characteristics occurred in each site and were different among sites. The duration of flooding had no effect on spore density but depressed AM root colonisation. Eleven different types of spores were recognized and four were identified. Two species dominated at the four sites: Glomus fasciculatum and Glomus intraradices. Spore density was highest in summer (dry season) and lowest in winter (wet season) with intermediate values in autumn and spring. Colonisation of L. glaber roots was highest in summer or spring and lowest in winter or autumn. The relative density of G. fasciculatum and G. intraradices versus Glomus sp. and Acaulospora sp. had distinctive seasonal peaks. These seasonal peaks occurred at all four sites, suggesting differences among AM fungus species with respect to the seasonality of sporulation. Spore density and AM root colonisation when measured at any one time were poorly related to each other. However, spore density was significantly correlated with root colonisation 3 months before, suggesting that high colonisation in one season precedes high sporulation in the next season.     

不同海拔高度高寒草甸光能利用效率的遥感模拟

利用植被光合模型模拟了藏北高原3个海拔高度(4300,4500 m和4700 m)的高寒草甸生态系统的光能利用效率.海拔4500 m的光能利用效率均值(0.47 g C/MJ)显著高于海拔4300 m(0.38 g C/MJ)和4700 m(0.35 g C/MJ),而海拔4300 m和4700 m两者间差异不显著.相关分析和多重逐步回归分析表明,影响每个海拔光能利用效率季节变化的主要因子为空气温度,相对湿度以及地表水分指数,这3个因子共同解释了99％以上的光能利用效率的季节变化,其中空气温度的贡献最大,相对湿度的贡献次之,地表水分指数的贡献最小,这说明在3个海拔的任何一个海拔高度,温度对光能利用效率季节变化的胁迫作用大于水分对光能利用效率季节变化的胁迫作用.多重逐步线性回归分析表明,生长季节均土壤含水量是决定生长季节均光能利用效率沿海拔高度分布的主导因子.单因子线性回归分析表明,地表水分指数可以定量化高寒嵩草草甸生态系统水分状况,它同时可以反应土壤水分、近地表空气湿度以及生态系统植被含水量状态.因此,在高寒嵩草草甸生态系统,用地表水分指数反应生态系统尺度水分对光能利用效率的胁迫作用是可行的.     

Responses of arbuscular mycorrhizal symbionts to contrasting environments: field evidence along a Tibetan elevation gradient

Plant adaptation to alpine ecosystems is not fully explained by plant physiological and morphological traits. Arbuscular mycorrhizal (AM) associations may be involved in mediating plant performance in response to environmental differences. Little is known, however, as to whether or not a close relationship exists between plant performance and arbuscular mycorrhizal fungus status across environmental gradients. We conducted a field investigation of the performance of six plant species and their associated AM fungi along higher and lower elevation gradients on Mount Segrila in Tibet. In most of our species, we observed higher shoot and inflorescence biomass production and a lower root-to-shoot ratio in the populations at those sites where the species was dominant (intermediate elevation sites) than in populations sampled at the limits of the distribution. The elevation pattern of root colonization differed with plant species on both gradients, and the extraradical hypha development of most species showed a unimodal pattern as did plant growth. The relationship between plant and fungus traits shows that AM fungus development generally matched host plant performance on the lower elevation gradient but not on the higher elevation gradient. This study provides evidence that plant distribution and productivity were significantly related to root and soil colonization by AM fungi, especially under less physically stressful conditions.     

青藏高原气候变化与高寒草地

综述了近五十年来青藏高原气候和高寒草地的变化趋势,阐述了气候变化对高寒草地的可能影响。气候变化主要通过水、热过程及其诱导的环境变化对青藏高原高寒草地产生显著的影响。主要过程包括：气候变化对气候带、植被带、植物、植物群落、农业生产以及生态系统固碳潜力等的影响。从目前的观测和研究结果来看,有关青藏高原气候变化及其对高寒草地的可能影响都还很难得出一致的结论。因此,如何科学评价气候变化及其预测和评价对高寒草地结构和功能的潜在影响,以及如何将已经发生的变化纳入到全球变化模型或评价体系中,以便更加精确地评估气候变化的长期影响,将成为必须要回答的关键科学问题。     

Unimodal pattern of soil hydrophobicity along an altitudinal gradient encompassing Mediterranean,temperate, and alpine ecosystems

Background and Aims

Soil water repellency (SWR, i.e. the reduced affinity for water due to the presence of hydrophobic coatings on soil particles) has relevant hydrological implications on the rate of water infiltration, surface runoff, and overland flow. Here, we test how SWR varies along a 2490 m altitudinal gradient encompassing six ecosystems including Mediterranean, Temperate, and Alpine vegetation types.

Methods

Water repellency, measured by the Molarity of an Ethanol Droplet (MED) test, was quantified in 80 soil samples collected for 16 different elevations. Soil quality was assessed by measuring soil texture, pH, organic carbon, salinity, and nutrient availability.

Results

SWR showed a unimodal pattern along the 2490 m transect, peaking at intermediate elevations. Unexpectedly, SWR was the highest under broad-leaf deciduous forests, and the lowest under evergreen, sclerophyllous Mediterranean vegetation types. The soil organic carbon content, and the pH were the main determinants of water repellency, showing respectively a positive, and a negative correlation with the SWR. In contrast, soil texture and salinity resulted unrelated to the SWR.

Conclusions

With this study we demonstrated a linkage between SWR, vegetation type and soil pH and organic carbon content along the elevation gradient. Further studies are needed to explicitly evaluate the impact SRW on erosion risk at catchment scale in the context of climatic change.

     

亚高山草地土壤原生生物群落结构和多样性海拔分布格局

生物多样性分布格局与维持机制即群落构建机制,是群落生态学研究的热点领域。微生物生态学中的一个关键问题是量化确定性过程和随机过程对微生物群落构建的相对贡献。尽管原生生物是土壤微生物群落中重要的组成部分,在微生物食物链中扮演着关键角色,但与细菌和真菌相比,目前对原生生物群落构建机制知之甚少。运用Illumina Miseq高通量测序技术,分析了五台山亚高山草地生态系统（海拔2000-3061 m范围内）土壤原生生物群落组成和多样性维持机制。结果表明,四个海拔梯度的土壤共获得有效序列520673条,分属于8个超群、24个门、65个纲、125个目、222个科和350个属。门水平上,丝足虫门（Cercozoa）、褐藻门（Ochrophyta）、纤毛门（Ciliophora）和顶复门（Apicomplexa）为主要优势类群。LEfSe分析显示,17个生物标志物对海拔梯度非常敏感,不同海拔梯度富集了不同的原生生物种群。尽管海拔对土壤原生生物群落的α多样性没有显著影响,但非度量多维尺度分析（NMDS）和相似性分析（ANOSIM）结果表明,原生生物群落组成和结构在海拔梯度上存在显著的差异（P<0.05）。冗余分析（RDA）显示海拔、土壤含水量、总氮和植物丰富度指数与原生生物群落结构存在显著相关性（P<0.05）。方差分解分析（VPA）和偏Mantel分析表明环境因子和空间变量对原生生物群落海拔分布格局均有显著影响,但是环境因子的相对作用（7.9%）明显大于空间变量（1.8%）。土壤原生生物群落之间的Bray-Curtis距离与海拔距离呈显著正相关关系（P<0.05）,说明选择过程可能是亚高山草地土壤原生生物群落分布格局的主要驱动因素。零模型分析进一步证实了确定性过程在原生生物群落构建中的相对作用大于随机过程。总之,五台山亚高山草地土壤原生生物群落组成和结构沿海拔梯度具有显著的变化格局,群落构建主要由确定性过程和随机过程驱动,但确定性过程占主导地位。     

Phenological variation in height growth and needle unfolding of Smith fir along an altitudinal gradient on the southeastern Tibetan Plateau

Little is known about variations in tree phenology and their driving forces on the Tibetan Plateau. Herein, we monitored shoot growth and needle unfolding of Smith fir (Abies georgei var. smithii) between 3,800 and 4,360 m a.s.l. in the Sygera Mountains, southeastern Tibetan Plateau. The trees were 0.45–1.48 m high and 12–39 years old. Their phenology was observed every week between May 5 and August 26, 2011. With increasing elevation, shoot growth and needle unfolding started increasingly later, thus indicating a thermal driving force. Although the weekly shoot increment was decreasing with increasing elevation, height growth at various elevations ended in the same week, implying other factors than temperature being responsible for the end of height growth. The accumulated heat sum for the onset of shoot growth appeared to be lower between 4,200 and 4,360 m than between 3,800 and 4,000 m. The anticipated spring warming will likely induce an earlier onset of shoot growth, whereas shoot growth will apparently not benefit from autumn warming. However, the lack of long-term data records precluded a robust statistical test of the underlying cause-and-effect relationships involved in the phenological variations of height growth and needle unfolding.     

Changes in plant biomass and species composition of alpine Kobresia meadows along altitudinal gradient on the Qinghai-Tibetan Plateau

Alpine Kobresia meadows are major vegetation types on the Qinghai-Tibetan Plateau. There is growing concern over their relationships among biodiversity, productivity and environments. Despite the im-portance of species composition, species richness, the type of different growth forms, and plant bio-mass structure for Kobresia meadow ecosystems, few studies have been focused on the relationship between biomass and environmental gradient in the Kobresia meadow plant communities, particularly in relation to soil moisture and edaphic gradients. We measured the plant species composition, her-baceous litter, aboveground and belowground biomass in three Kobresia meadow plant communities in Haibei Alpine Meadow Ecosystem Research Station from 2001 to 2004. Community differences in plant species composition were reflected in biomass distribution. The total biomass showed a de-crease from 13196.96±719.69 g/m2 in the sedge-dominated K. tibetica swamp to 2869.58±147.52 g/m2 in the forb and sedge dominated K. pygmaea meadow, and to 2153.08±141.95 g/m2 in the forbs and grasses dominated K. humilis along with the increase of altitude. The vertical distribution of below-ground biomass is distinct in the three meadow communities, and the belowground biomass at the depth of 0-10 cm in K. tibetica swamp meadow was significantly higher than that in K. humilis and K. pygmaea meadows (P<0.01). The herbaceous litter in K. tibetica swamp was significantly higher than those in K. pygnaeca and K. humilis meadows. The effects of plant litter are enhanced when ground water and soil moisture levels are raised. The relative importance of litter and vegetation may vary with soil water availability. In the K. tibetica swamp, total biomass was negatively correlated to species richness (P<0.05); aboveground biomass was positively correlated to soil organic matter, soil moisture, and plant cover (P<0.05); belowground biomass was positively correlated with soil moisture (P<0.05). However, in the K. pygnaeca and K. humilis meadow communities, aboveground biomass was posi-tively correlated to soil organic matter and soil total nitrogen (P<0.05). This suggests that the distribu-tion of biomass coincided with soil moisture and edaphic gradient in alpine meadows.