Assessing Level of Development and Successional Stages in Biological Soil Crusts with Biological Indicators

Biological soil crusts (BSCs) perform vital ecosystem services, but the difference in biological components or developmental level still affects the rate and type of these services. In order to differentiate crust successional stages in quantity and analyze the relationship between crust developmental level and successional stages, this work determined several biological indicators in a series of different developmental BSCs in the Shapotou region of China. The results showed that crust developmental level (level of development index) can be well indicated by crust biological indicators. Photosynthetic biomass was the most appropriate to differentiate crust successional stages, although both photosynthetic biomass and respiration intensity increased with the development and succession of BSCs. Based on of the different biological compositions, BSCs were quantificationally categorized into different successional stages including cyanobacterial crusts (lichen and moss coverages <20 %), lichen crusts (lichen coverage >20 % but moss coverage <20 %), semi-moss crusts (moss coverage >20 % but <75 %), and moss crusts (moss coverage >75 %). In addition, it was found that cyanobacterial and microalgal biomass first increased as cyanobacterial crusts formed, then decreased when lots of mosses emerged on the crust surface; however nitrogen-fixing cyanobacteria and heterotrophic microbes increased in the later developmental BSCs. The structural adjustment of biological components in the different developmental BSCs may reflect the requirement of crust survival and material transition.     

Ecological Succession,Hydrology and Carbon Acquisition of Biological Soil Crusts Measured at the Micro-Scale

The hydrological characteristics of biological soil crusts (BSCs) are not well understood. In particular the relationship between runoff and BSC surfaces at relatively large (>1 m²) scales is ambiguous. Further, there is a dearth of information on small scale (mm to cm) hydrological characterization of crust types which severely limits any interpretation of trends at larger scales. Site differences and broad classifications of BSCs as one soil surface type rather than into functional form exacerbate the problem. This study examines, for the first time, some hydrological characteristics and related surface variables of a range of crust types at one site and at a small scale (sub mm to mm). X-ray tomography and fine scale hydrological measurements were made on intact BSCs, followed by C and C isotopic analyses. A ‘hump’ shaped relationship was found between the successional stage/sensitivity to physical disturbance classification of BSCs and their hydrophobicity, and a similar but ‘inverse hump’ relationship exists with hydraulic conductivity. Several bivariate relationships were found between hydrological variables. Hydraulic conductivity and hydrophobicity of BSCs were closely related but this association was confounded by crust type. The surface coverage of crust and the microporosity 0.5 mm below the crust surface were closely associated irrespective of crust type. The δ ¹³C signatures of the BSCs were also related to hydraulic conductivity, suggesting that the hydrological characteristics of BSCs alter the chemical processes of their immediate surroundings via the physiological response (C acquisition) of the crust itself. These small scale results illustrate the wide range of hydrological properties associated with BSCs, and suggest associations between the ecological successional stage/functional form of BSCs and their ecohydrological role that needs further examination.     

黄土丘陵区生物土壤结皮表面糙度特征及影响因素

地表糙度是影响地表径流和侵蚀过程的重要属性.生物结皮在干旱半干旱区广泛分布,是地表糙度的影响因子之一.本文采用链条法测定了黄土丘陵区不同发育阶段生物结皮表面糙度特征,分析了不同发育阶段生物结皮表面糙度对坡向、土壤含水量和冻融作用的响应及其与各理化性质的相关性,初步探索了生物结皮对地表糙度的影响及其相关因素.结果表明: 生物结皮显著改变地表糙度,随着生物结皮从藻结皮向藓结皮演替,其糙度先降低后增加,生物结皮发育形成10年以后,其表面糙度基本趋于稳定;研究区早期形成的藻结皮表面糙度较裸土降低47.0%,深色藻结皮(藓类盖度<20%)较裸土降低20.4%,混生结皮(藓类盖度为20%～60%)和苔藓结皮(藓类盖度>70%)表面糙度与深色藻结皮基本一致;坡向对发育10年以上的生物结皮表面糙度的影响不显著;土壤含水量影响地表糙度特征.研究区浅色藻结皮表面糙度随水分变化较为剧烈;随着生物结皮发育,深色藻结皮、混生结皮和苔藓结皮表面糙度随水分的变化趋于平缓.冻融增加了生物结皮表面糙度.浅色藻结皮经两次冻融后表面糙度增加29.7%;深色藻结皮、混生结皮和藓结皮表面糙度的影响需经过反复冻融才有所体现.生物结皮表面糙度与藓结皮盖度呈显著正相关(P＜0.1).     

荒漠结皮层藓类植物死亡对表层土壤水分蒸发和入渗的影响

水分是荒漠植物生长最主要的限制因子,藓类结皮作为荒漠土壤表层重要覆被物,对土壤水分蒸发入渗具有重要影响。研究表明,在全球气候变化背景下,不确定的降水格局变化导致结皮层藓类植物出现集群死亡现象,但这一过程对荒漠地表土壤水分蒸发与入渗过程的影响及其机理尚不清楚。以古尔班通古特沙漠齿肋赤藓结皮为研究对象,利用便携式渗透计和蒸发仪,研究了结皮层藓类植物死亡对土壤水分蒸发与入渗的影响。结果表明,与裸沙相比,藓类结皮的存在显著抑制了水分入渗,而藓类植物死亡的结皮层抑制作用最大,其初渗速率、稳渗速率和累积入渗量分别是活藓类结皮的39.89%、85.91%及64.48%,仅为裸沙的5.96%、13.13%及20.42%。在水分蒸发初期,裸沙的水分蒸发速率明显高于活藓类结皮和藓类植物死亡的结皮层,但藓类植物死亡的结皮层维持相对稳定的蒸发速率的时间长于裸沙和活藓类结皮,这也导致最终累计蒸发量以藓类植物死亡的结皮层最高、裸沙最低。可见,荒漠生物土壤结皮中藓类植物死亡会明显减少土壤水分入渗、增大水分蒸发,进一步影响荒漠表层土壤水分格局,从而影响生物土壤结皮与维管植物的水分利用关系。     

Development of artificially induced biological soil crusts in fields and their effects on top soil

Aims

Biological soil crusts (BSCs) could improve severe environment ecological conditions by increasing soil moisture, soil nitrogen concentration, and so on. In order to control desertification and recover the destroyed soil fertility utilizing a new means using BSCs, the soil surface was artificially inoculated with Microcoleus vaginatus and Scytonema javanicum. Relationships between the development of the artificially induced biological soil crusts and the distribution and dynamic changes of nitrogen and phosphorus in the soil crusts have been analyzed.

Methods

Crusts of different ages were investigated by measuring soil physical and chemical factors, such as moisture, pH, total and available N content, and total and available P, which were correlated with the depths of the crusts.

Results

This study found that the types of color, shape, and species components of the algal crusts increased with crust development. Soil moisture, total N, available N, and available P increased gradually with crust growth. Soil with crusts was wetter than the controlled naked sandy soil, and a significant correlation was observed between biomass and total nitrogen (r?=?0.946, P?=?0.015). Soil pH was lower than that of control. The scytonemin on the soil surface was exceptionally higher than the other pigments, and all the pigments were mainly distributed at the soil surface level. Though the crusts were mainly distributed on soil surface, the available P was mainly stored below the crust layer.

Conclusions

Pearson correlation tests indicated that artificially inoculated biological crusts could improve soil fertility and micro-environment of the top soil: The development of artificially induced BSCs was very well, and this was favorable to inducing the following crust succession.     

内蒙古黑岱沟露天煤矿植被恢复区生物土壤结皮的固氮潜力

氮素限制在陆地生态系统中普遍存在,在干旱受损生态系统中表现得尤为明显.生物土壤结皮是干旱、半干旱区受损生态系统植被恢复过程中的重要组成部分和氮源贡献者.以黑岱沟露天煤矿植被恢复区广泛分布的两类典型生物土壤结皮(藻类结皮和藓类结皮)为研究对象,通过野外调查采集样品,在实验室条件下测定了两类结皮的固氮活性,分析了其固氮活性对水热因子的响应特征及其与草本、结皮盖度的关系.结果表明：不同演替阶段人工植被及相邻撂荒地和天然植被下生物土壤结皮的固氮活性在9～150 μmol C₂H₄·m^-2·h^-1,藻类结皮(平均为77 μmol C₂H₄·m^-2·h^-1)显著高于藓类结皮(17 μmol C₂H₄·m^-2·h^-1).人工植被区3种常见植被类型下藻类和藓类结皮固氮活性均表现为“灌-草型”显著高于“乔-灌型”和“乔-灌-草型”.藻类和藓类结皮的固氮活性与样品相对含水量(10%～100%)和培养温度(5～45 ℃)均呈显著的二次函数关系,其固氮活性随水分、温度升高均呈先上升后下降的趋势,分别在60%和80%相对含水量时达到最大固氮速率,其最适固氮温度均为25 ℃.藻类结皮固氮活性与草本盖度呈显著的二次函数关系,草本盖度超过20%时固氮活性开始降低,藓类结皮固氮活性与草本植物盖度呈显著负相关.两类结皮固氮活性与其盖度均呈显著的正相关关系,随结皮盖度增加其固氮活性显著升高.露天煤矿植被恢复区两类生物土壤结皮固氮活性差异主要由结皮组成生物体即隐花植物的差异所致,不同植被类型下的水热差异及不同植被演替阶段草本、结皮盖度的差异是影响生物土壤结皮氮固定的关键因子,生物土壤结皮在人工植被区的拓殖发育及其氮输入是系统健康发展的重要标志.     

毛乌素沙地固碳功能菌群落随生物结皮发育阶段的演变特征

生物结皮是干旱区生态系统光合固碳的重要贡献者,固碳功能菌是生物结皮碳固定过程中的关键功能群,而毛乌素沙地生物结皮有关固碳功能菌群落多样性的研究未见详细报道。通过qPCR和高通量测序,研究了毛乌素沙地固碳功能菌基因丰度、群落多样性在生物结皮发育过程中的演变规律及其关键环境因子。结果表明,form IAB、IC和ID的基因丰度均随生物结皮发育呈升高趋势,在地衣结皮最高,在藓结皮又明显降低;form IAB的Chao1和form IC的Shannon指数随生物结皮演替呈显著上升趋势,而 form IC和ID的Chao1则在藻结皮达到峰值。物种组成上,IAB型固碳功能菌以蓝藻门的颤藻目占优势,其相对丰度在藻结皮最高,随生物结皮演替逐渐降低;IC型固碳功能菌在裸沙中以土壤红杆菌目为主, 而在三种生物结皮中以生丝微菌目、亚硝化单胞菌目和红螺菌目占优势; ID型固碳菌则以硅藻门的舟形藻目占优势,其丰度在裸沙中最高。IAB和IC型固碳功能菌群落结构在藻结皮和地衣结皮类似,而与裸沙和藓结皮差异显著,裸沙与藓结皮亦具显著差异。生物结皮演替过程中生物类群和土壤理化特征的改变为固碳功能菌提供了不同的生态位,土壤总有机碳、全氮、铵态氮、全磷、速效磷和pH对固碳功能菌群落进行综合调控,通过对固碳功能菌的筛选作用,最终改变固碳功能菌群落组成和结构。为深入理解荒漠生态系统生物结皮的光合固碳功能的微生物学机制提供了理论支撑。     

古尔班通古特沙漠生物结皮不同发育阶段中藻类的变化

通过在古尔班通古特沙漠南缘相同的地貌部位,选择裸沙、藻结皮、地衣结皮和苔藓结皮4种不同演替阶段中的生物结皮,研究了藻类的种类组成、优势种和生物量的变化.结果表明:(1)在结皮的不同演替阶段,藻类种类组成不同,其常见物种有一定的差异,如裸沙中藻类常见种是脆杆藻2(Fragilaria sp.2)、威利颤藻(Oscillatoria willei)和奥克席藻(Phormidium okenii),藻结皮的常见种是小聚球藻(Synechococcus parvus)、颗粒常丝藻 (Tychonema granulatum)、韧氏席藻(Phormidium retzli);同时在不同发育阶段亦存在一些特有种.(2)在裸沙发育到成熟生物结皮的过程中,藻类的优势物种也发生相应的变化.裸沙、藻结皮、地衣结皮和苔藓结皮的优势种分别是脆杆藻1(Fragilaria sp.1)、具鞘微鞘藻(Microcoleus vaginatus)、具鞘微鞘藻、眼点伪枝藻(Scytonema ocellatum)或集球藻(Palmellococcus miniatus).(3)藻类生物量在生物结皮不同演替阶段差异极显著(P<0.01),在裸沙中藻类生物量最低,随着生物结皮的逐渐发育,藻类生物量明显升高,地衣结皮最高,约是裸沙的8.3倍,当发育至苔藓结皮时,藻类生物量又有所下降.(4)在裸沙中基本为松散的沙粒,随着生物结皮的演替,丝状种类占明显的优势,尤其是具鞘微鞘藻,另外真菌菌丝和苔藓假根分别在地衣结皮和苔藓结皮中起着重要作用.     

Diurnal variations of chlorophyll fluorescence and CO2 exchange of biological soil crusts in different successional stages in the Gurbantunggut Desert of northwestern China

Biological soil crusts (BSCs) formed by different combinations of photosynthetic algae, cyanobacteria, lichens and mosses are well-developed in the Gurbantunggut Desert of northwestern China. To investigate the different responses of BSCs to environmental factors, the diurnal variations of chlorophyll fluorescence and CO₂ exchange of BSCs in different successional stages were measured following artificial rehydration in the field. Results showed that the maximum potential quantum efficiency of PSII (F _v/F _m), the actual PSII efficiency (Φ_PSII) and the relative rate of electron transport as well as net photosynthesis of the different successional BSCs varied similarly and changed markedly with diurnal fluctuations in light and temperature. Further analyses indicated that CO₂ exchange and photosynthetic pigment content of chlorophyll (Chl) a, Chl b and carotenoids increased with the developmental level of BSCs, from cyanobacterial crust to lichen crust to moss crust. The differences in responses of BSCs to environmental factors and photosynthetic pigment content may be partially attributed to differences in species composition and morphological characteristics of the various BSCs. Overall, moss crust is better adapted to a wide range of irradiance and higher temperatures than lichen and cyanobacterial crusts. Therefore, BSCs in a later successional stage are expected to play a more important role in desertification control than those of the earlier stages.     

温带荒漠灌丛和藓类结皮对土壤养分的贡献

灌丛与生物土壤结皮镶嵌分布是温带荒漠常见的地表景观之一,二者的发育均显著影响了地表土壤养分的空间分布特征及循环过程。然而,灌丛和生物土壤结皮对荒漠土壤表层养分的影响存在怎样的差异,二者对养分变化的贡献度如何尚不清楚。因此,选择中国北方典型温带荒漠古尔班通古特沙漠为研究区,以荒漠中的优势灌丛膜果麻黄(Ephedra przewalskii)灌丛和生物土壤结皮发育高级阶段的藓类结皮的结皮层及结皮下层土壤为研究对象,采集不同微生境(裸露地、灌丛下)的裸沙与藓类结皮的土壤样品。为探究不同微生境下不同土层碳、氮、磷和钾养分变化特征,测定了不同土层土壤有机碳(SOC)、全氮(TN)、全磷(TP)、全钾(TK)、速效氮(AN)、速效磷(AP)和速效钾(AK)含量。结果表明:(1)相较于裸露地,灌丛显著提高了藓类结皮不同土层有机碳、全氮和全钾的含量和裸沙全磷的含量,降低了灌丛下藓类结皮土壤全磷的含量。(2)对于速效养分而言,与裸沙相比,裸露地藓类结皮覆盖降低了土壤速效氮含量,增加了速效磷和速效钾含量;而灌丛下藓类结皮覆盖提高了土壤速效氮和速效钾的含量,但降低了速效磷的含量。 (3) 相关性分析显示,在0-2 cm土壤中速效磷与速效氮呈现显著负相关,而在2-6 cm速效磷与速效氮呈现显著正相关(P<0.01)。(4)贡献度分析表明,土壤中灌丛效应对养分的贡献(42.54%)要远大于藓类结皮的贡献(2.43%),但二者交互作用却降低了除速效氮以外的其他土壤养分含量。综上,灌丛、藓类结皮覆盖和土层深度变化对土壤碳、氮、磷和钾养分均具有显著影响(P<0.05),但三者间的交互效应对养分的影响不显著(P>0.05)。相对于裸露地,灌丛与藓类结皮的覆盖均对表层土壤碳、氮、磷和钾养分具有促进作用,且随着土层深度的下降,土壤养分含量呈现显著的下降趋势(P<0.05)。因此,在荒漠生态系统中耐旱灌丛与生物土壤结皮这两个最重要的斑块单元联合调控了微尺度土壤养分的空间异质性变化。     

Crust Composition and Disturbance Drive Infiltration Through Biological Soil Crusts in Semiarid Ecosystems

Soil crusts influence many soil parameters that affect how water moves into and through the soil, and therefore, critically influence water availability, erosion processes, nutrient fluxes, and vegetation distribution patterns in semiarid ecosystems. Soil crusts are quite sensitive to disturbance, and their alteration can lead to modification of the local hydrological regime, thus affecting general functioning of the ecosystem. The aim of this study was to analyze the influence of different types of soil crusts, physical, and biological in different developmental stages, as well as the impact of their disturbance, on infiltration. This was assessed by means of rainfall simulations conducted in two semiarid ecosystems in southeast Spain characterized by different lithologies, topographies, and soil crust distributions. Two consecutive rainfall simulation experiments (50 mm h⁻¹ rainfall intensity), the first on dry soil and the second on wet soil, were carried out in microplots (0.25 m²) containing the most representative soil crust types at each site, each crust type subjected to three disturbance treatments: (a) undisturbed, (b) trampling, and (c) removal. Infiltration in the crusts was higher on coarse- than on fine-textured soils and almost two times greater on dry than on wet soil. Biological soil crusts (BSC) showed higher infiltration rates than physical soil crusts (PSC). Within BSC, infiltration increased as cyanobacterial biomass increased and was the highest in moss crusts. However, late-successional crustose and squamulose lichen crusts showed very low infiltration rates. Trampling reduced infiltration rates, especially when soil was wet, whereas crust removal enhanced infiltration. But this increase in infiltration after removing the crust decreased over time as the soil sealed again due to raindrop impact, making runoff rates in the scraped microplots approach those registered in the respective undisturbed crust types. Our results demonstrate that water redistribution in semiarid ecosystems strongly depends on the type of crusts that occupy the interplant spaces and the characteristics of the soils which they overly, as well as the antecedent moisture conditions of the soil. Disturbance of these crust patches results in increased runoff and erosion, which has important consequences on general ecosystem functioning.     

荒漠化地区生物结皮的研究进展

防沙、治沙和固沙一直以来是荒漠化生态环境建设的热点问题。作者从生物结皮对荒漠化地区生态环境的影响,以及在生态系统演替中起到的重要作用等方面进行了综述,指出了地衣在生物结皮中所占有的重要地位,展望了生物结皮研究与应用的前景。     

古尔班通古特沙漠生物土壤结皮对土壤氮库垂直分布特征的影响

生物土壤结皮在干旱区氮素地球化学循环中具有重要作用,研究不同生物土壤结皮下不同形态氮素含量的变化,解析生物土壤结皮对土壤养分影响过程和范围,有助于进一步理解生物土壤结皮的生态功能。本研究以古尔班通古特沙漠藻-地衣混生结皮和藓类结皮两种生物土壤结皮为研究对象,以裸沙为对照,测定生物土壤结皮层和0—100 cm内8个土层全氮、无机氮、可溶性有机氮、游离态氨基酸氮、微生物生物量氮等氮库含量,和土壤脲酶、硝酸盐还原态酶、亮氨酸氨基肽酶等土壤胞外酶活性。结果表明：1)结皮层各形态氮素含量和各土壤酶活性显著高于其下层土壤,结皮层和结皮下各层土壤氮库整体上表现为藓类结皮>藻-地衣混生结皮>裸沙;土壤氮库各形态氮素含量和土壤酶活性在垂直分布上均呈现先显著下降(0—20 cm)后稳定(20—100 cm)的趋势;在20—30 cm土层,除裸沙的无机氮、铵态氮以及藻-地衣混生结皮的硝态氮外,其余速效氮(无机氮、硝态氮、铵态氮)含量具有增加的特点。2)土壤各氮库含量与全磷、有机碳、电导率、土壤脲酶和亮氨酸氨基肽酶活性呈正相关,与pH、土壤含水率呈负相关。3)利用氮循环相关指标建立土壤氮循环多功能...     

Southern African Biological Soil Crusts are Ubiquitous and Highly Diverse in Drylands,Being Restricted by Rainfall Frequency

Biological soil crusts (BSCs) are found in all dryland regions of the world, including the polar regions. They are also known to occur in the southern African region. Although there were a number of case studies on BSCs from that region, we did not know if they are a normal part of the vegetation cover or just a phenomenon that occasionally occurs here and there. In order to investigate diversity, distribution patterns, and the driving factors of both, we followed a random sampling system of observatories along a transect, stretching from the Namibian-Angolan border down south to the Cape Peninsula, covering seven different major biomes. Biological soil crusts were found to occur in six out of seven biomes. Despite the fact that soil-dwelling algae occurred in the Fynbos biome, crust formation was not observed for hitherto unknown reasons. Seven BSC types were distinguished on the basis of morphology and taxonomic composition: three of them were cyanobacteria-dominated, one with additional chlorolichens, two with bryophytes, one hypolithic type restricted to quartz gravel pavements, and the unique lichen fields of the Namib Desert. Besides 29 green algal species in 21 genera, one heterokont alga, 12 cyanolichens, 14 chlorolichens, two genera of liverworts, and three genera of mosses, these crusts are positioned among the most diverse BSCs worldwide mainly because of the unusual high cyanobacterial species richness comprising 58 species in 21 genera. They contribute considerably to the biodiversity of arid and semi-arid bioregions. Taxonomic diversity of cyanobacteria was significantly higher in the winter rain zone than in the summer rain zone (54 versus 32 species). The soil photosynthetic biomass (chlorophylla/m2), the carbon content of the soil and the number of BSC types were significantly higher in the winter rain zone (U27, 29=215.0, p=0.004 [chla]; U21, 21=135.0, p=0.031 [C]; U27, 29=261.5, p=0.028 [BSC types]; excluding the fog-dominated Namib biome). The winter rain zone is characterized by a lower precipitation amount, but a higher rain frequency with the number of rainy days more evenly distributed over the year. The dry period is significantly shorter per year in the winter rain zone (U8, 9=5.0, p=0.003). We conclude that rain frequency and duration of dry periods rather than the precipitation amount is the main factor for BSC growth and succession. Nitrogen content of the soils along the transect was generally very low and correlated with soil carbon content. There was a weak trend that an increasing proportion of silt and clay (<0.63 mm) in the soil is associated with higher values of BSC chlorophyll content (Pearson correlation coefficient=0.314, p=0.237). Furthermore, we found a significant positive correlation between silt and clay and the number of BSC types (Pearson correlation coefficient=0.519, p=0.039), suggesting that fine grain-size promotes BSC succession and their biomass content. Lichens and bryophytes occurred in BSCs with lower disturbance frequencies (e.g. trampling) only. Crust thickness and chlorophyll content increased significantly from crusts of the early successional type to the late successional crust types. From our results, we conclude that BSCs are a normal and frequent element of the vegetation in arid and semi-arid southwestern Africa, and that rain frequency and duration of dry periods rather than the precipitation amount are the key factors for the development, differentiation and composition of BSCs.     

Biological Soil Crust Rehabilitation in Theory and Practice: An Underexploited Opportunity

Biological soil crusts (BSCs) are ubiquitous lichen–bryophyte microbial communities, which are critical structural and functional components of many ecosystems. However, BSCs are rarely addressed in the restoration literature. The purposes of this review were to examine the ecological roles BSCs play in succession models, the backbone of restoration theory, and to discuss the practical aspects of rehabilitating BSCs to disturbed ecosystems. Most evidence indicates that BSCs facilitate succession to later seres, suggesting that assisted recovery of BSCs could speed up succession. Because BSCs are ecosystem engineers in high abiotic stress systems, loss of BSCs may be synonymous with crossing degradation thresholds. However, assisted recovery of BSCs may allow a transition from a degraded steady state to a more desired alternative steady state. In practice, BSC rehabilitation has three major components: (1) establishment of goals; (2) selection and implementation of rehabilitation techniques; and (3) monitoring. Statistical predictive modeling is a useful method for estimating the potential BSC condition of a rehabilitation site. Various rehabilitation techniques attempt to correct, in decreasing order of difficulty, active soil erosion (e.g., stabilization techniques), resource deficiencies (e.g., moisture and nutrient augmentation), or BSC propagule scarcity (e.g., inoculation). Success will probably be contingent on prior evaluation of site conditions and accurate identification of constraints to BSC reestablishment. Rehabilitation of BSCs is attainable and may be required in the recovery of some ecosystems. The strong influence that BSCs exert on ecosystems is an underexploited opportunity for restorationists to return disturbed ecosystems to a desirable trajectory.     

腾格里沙漠东南缘生物土壤结皮的固氮潜力

以腾格里沙漠东南缘广泛分布的3类典型生物土壤结皮(藻类结皮、地衣结皮和藓类结皮)为研究对象,在野外环境下连续一年(2010年6月至2011年5月)测定了不同类型结皮的固氮潜力、季节变化,及其对环境因子的响应特征.结果表明：整个试验期间,藻类、地衣和藓类结皮的固氮活性分别为14～133、20～101和4～28 mol·m^-2·h^-1,差异显著,这种差异主要是由结皮种类组成的差异所致.3类结皮固氮活性对环境因子的响应特征基本一致.3类结皮固氮活性与降水量关系不显著,但与试验前3天小于3 mm的降水量均呈显著的正相关关系,说明该地区3类结皮在3 mm降水条件下即可达到最大固氮速率.3类结皮固氮活性与试验期温度均呈显著的二次函数关系,随气温升高均呈先上升后下降的趋势,藻类和地衣结皮的固氮活性在超过30 ℃后即迅速下降,而藓类结皮的固氮活性在超过25 ℃后即开始下降,说明不同类型结皮具有不同的固氮适宜温度区间.3类结皮固氮活性的季节变化均表现为秋季>春季>夏季>冬季,夏季高温和冬季低温抑制了结皮固氮酶活性,而春末秋初适宜的水热条件促进了其固氮活性的提高,结皮固氮活性的季节变化主要受水热因子的共同调控.温带荒漠区生物土壤结皮在湿润条件下全年均具有固氮能力,环境因子对其氮固定的控制作用层次分明,水分是影响其固氮速率和持续时间的关键因子,在水分和碳源充足的条件下,温度则是制约其固氮能力的主要因子.     

火电厂周边不同生物结皮细菌群落特征差异及其影响因素

为探究大气降尘重金属污染对矿区周边不同类型生物结皮细菌群落结构的影响,利用高通量测序技术分析位于宁东能源化工基地典型火电厂周边的3类生物结皮(藻结皮ZB、混生结皮HB、苔藓结皮TB)和对照(CK,裸土)的细菌丰度和群落结构,并探讨了影响细菌群落结构的环境因子。结果表明: 不同类型生物结皮的理化性质和重金属含量存在差异,且由于生物结皮对大气降尘重金属的富集作用造成各类结皮均达重度污染级别。在相对丰度排名前10的优势细菌门中,芽单胞菌门、蓝细菌门在不同类型生物结皮之间差异显著。细菌群落α多样性由高到低排序依次为CK>TB>HB>ZB。非度量多维排序(NMDS)结果显示,裸土细菌群落与其他3种生物结皮存在明显差异。相关性分析表明,生物结皮演替对细菌群落组成具有显著影响,细菌多样性和组成与pH、养分、重金属含量等密切相关。放线菌门、绿弯菌门相对丰度与pH值呈显著正相关关系,而与全氮(TN)、全磷(TP)、Pb、Zn、Cd均呈显著负相关关系;冗余分析结果表明,TN、pH、TP、有机碳(SOC)是影响3种生物结皮细菌群落α多样性以及一些优势菌群相对丰度的主要土壤环境因子,而重金属Pb、Zn、Cd是影响细菌群落结构的主要重金属元素,对细菌群落数量和多样性有抑制或刺激作用。说明pH、重金属和养分是影响结皮细菌群落组成的关键因子。总体而言,长期的重金属富集作用会对生物结皮的细菌多样性和群落组成产生影响。     

A culture-independent study of free-living fungi in biological soil crusts of the Colorado Plateau: their diversity and relative contribution to microbial biomass

Molecular methodologies were used to investigate free-living fungal communities associated with biological soil crusts (BSCs), along km-scale transects on the Colorado Plateau (USA). Two cyanobacteria-dominated crust types that did not contain significant lichen cover were examined. Fungal community diversity and composition were assessed with PCR-denaturing gradient gel electrophoresis (DGGE) fingerprinting and sequencing, and fungi-specific quantitative PCR was used to measure fungal population densities as compared with those of bacteria. Our results clearly indicate that free-living fungi, while ubiquitous in BSCs, are less diverse and contribute far less biomass than their bacterial counterparts. Biological soil crust fungal community structure differed from that of uncrusted soils in their surroundings. Phylogenetic analyses placed the majority of BSC fungi within the Ascomycota , confirmed the importance of dematiaceous fungi, and pointed to members of the genera Alternaria and Acremonium as the most common free-living fungi in these crusts. Phylotypes potentially representing novel taxa were recovered, as were several belonging to the Basidiomycota that would not have been readily recognized by culture-dependant means.     

生物土壤结皮对荒漠土壤线虫群落的影响

在干旱的沙漠生态系统中,生物土壤结皮对于沙丘的固定和土壤生物的维持起着相当重要的作用.土壤线虫能敏感的指示土壤的恢复程度,是衡量沙区生态恢复与健康的重要生物学属性,而目前关于生物土壤结皮与土壤线虫的关系研究很少.为探明生物土壤结皮对土壤线虫的影响,以腾格里沙漠东南缘的人工植被固沙区藻结皮和藓类结皮覆盖的沙丘土壤为研究对象,根据固沙年限的不同将样地分为4个不同的区进行采样(1956、1964、1981和1991年固沙区),以流沙区作为对照;同时,在不同季度(4、7、9和12月)分别采集腾格里沙漠东南缘的人工植被固沙区藻结皮和藓类结皮覆盖下不同土层(0-10、10-20和20-30 cm)的沙丘土壤,以沙坡头地区的红卫天然植被区为对照,分析生物土壤结皮下土壤线虫的时空变化.采用改良的Baermann漏斗法进行分离线虫,用光学显微镜鉴定并统计.研究表明:1956、1964、1981和1991年人工植被固沙区的藻结皮和藓类结皮均可显著提高其下土壤线虫多度、属的丰富度、Shannon-Weaver多样性指数、富集指数和结构指数(P＜0.05),这可能是因为生物土壤结皮的存在为土壤线虫提供了重要的食物来源和适宜的生存环境;固沙年限与结皮下土壤线虫多度、属的丰富度、Shannon-Weaver多样性指数、富集指数和结构指数存在显著的正相关关系(P＜0.05),这说明固沙年限越久,越有利于土壤线虫的生存和繁衍;结皮类型显著影响土壤线虫群落,相对于藻结皮而言,藓类结皮下土壤线虫多度与属的丰富度更高(P＜0.05),这说明演替后期的藓类结皮比演替早期的藻结皮更有利于土壤线虫的生存和繁衍.此外,藻结皮和藓类结皮均可显著增加其下0-10、10-20和20-30 cm土层线虫多度和属的丰富度(P＜0.05),但随着土壤深度的增加,这种影响逐渐减弱,表明生物土壤结皮更有利于表层土壤线虫的生存;而且,随着季节的变化,藻结皮和藓类结皮下土壤线虫多度基本表现为秋季＞夏季＞春季＞冬季,这些反映了生物土壤结皮的生物量、盖度和种类组成随着季节变化而变化.因此,腾格里沙漠东南缘的人工植被固沙区生物土壤结皮的存在与演替有利于土壤线虫的生存和繁衍,增加了线虫数量、种类和多样性,这指示了生物土壤结皮有利于该区土壤及其相应生态系统的恢复.     

Soil Loss and Runoff in Semiarid Ecosystems: A Complex Interaction Between Biological Soil Crusts, Micro-topography, and Hydrological Drivers

Biological soil crusts (BSCs) cover non-vegetated areas in most arid and semiarid ecosystems. BSCs play a crucial role in the redistribution of water and sediments and, ultimately, in the maintenance of ecosystem function. The effects of BSCs on water infiltration are complex. BSCs increase porosity and micro-topography, thus enhancing infiltration, but, at the same time, they can increase runoff by the secretion of hydrophobic compounds and clogging of soil pores upon wetting. BSCs confer stability on soil surfaces, reducing soil detachment locally; however, they can also increase runoff, which may increase sediment yield. Although the key role of BSCs in controlling infiltration–runoff and erosion is commonly accepted, conflicting evidence has been reported concerning the influence of BSCs on runoff generation. Very little is known about the relative importance of different BSC features such as cover, composition, roughness, or water repellency, and the interactions of these attributes in runoff and erosion. Because BSC characteristics can affect water flows and erosion both directly and indirectly, we examined the direct and indirect effects of different BSC features on runoff and erosion in a semiarid ecosystem under conditions of natural rainfall. We built structural equation models to determine the relative importance of BSC cover and type and their derived surface attributes controlling runoff and soil erosion. Our results show that the hydrological response of BSCs varies depending on rainfall properties, which, in turn, determine the process governing overland flow generation. During intense rainfalls, runoff is controlled not only by rainfall intensity but also by BSC cover, which exerts a strong direct and indirect influence on infiltration and surface hydrophobicity. Surface hydrophobicity was especially high for lichen BSCs, thus masking the positive effect of lichen crust on infiltration, and explaining the lower infiltration rates recorded on lichen than on cyanobacterial BSCs. Under low intensity, rainfall volume exerts a stronger effect than rainfall intensity, and BSC features play a secondary role in runoff generation, reducing runoff through their effect on surface micro-topography. Under these conditions, lichen BSCs presented higher infiltration rates than cyanobacterial BSCs. Our results highlight the significant protective effect against erosion exerted by BSCs at the plot scale, enhancing surface stability and reducing sediment yield in both high- and low-magnitude rainfall events.