Nitrogen cycling in desert biological soil crusts across biogeographic regions in the Southwestern United States

Biological soil crusts (BSCs) are thought to be important in the fertility of arid lands as gateways for carbon (C) and nitrogen (N). Studies on the Colorado Plateau have shown that an incomplete internal N cycle operates in BSCs that results in significant exports of dissolved organic N, ammonia and nitrate into the bulk soil through percolating water, thus mechanistically explaining their role as a N gateway. It is not known if this pattern is found in other arid regions. To examine this, we measured rates of major biogeochemical N-transformations in a variety of BSCs collected from the Colorado Plateau and the Mojave, Sonoran and Chihuahuan Deserts. Dinitrogen fixation and aerobic ammonia oxidation were prominent transformations at all sites. We found anaerobic ammonia oxidation (anammox) rates to be below the detection limit in all cases, and at least 50-fold smaller than rates of N₂-fixation, making it an irrelevant process for these BSCs. Heterotrophic denitrification was also of little consequence for the flow of N, with rates at least an order of magnitude smaller than those of N₂-fixation. Thus we could confirm that despite the demonstrable differences in microbial community composition and soil material, BSCs from major biogeographic regions in arid North America displayed a remarkably consistent pattern of internal N cycling. The implications for arid land fertility drawn from previous studies in the Colorado Plateau appear applicable to BSCs across other arid regions of the Southwestern United States.     

Relevance of ammonium oxidation within biological soil crust communities

Thin, vertically structured topsoil communities that become ecologically important in arid regions (biological soil crusts or BSCs) are responsible for much of the nitrogen inputs into pristine arid lands. We studied N(2) fixation and ammonium oxidation (AO) at subcentimetre resolution within BSCs from the Colorado Plateau. Pools of dissolved porewater nitrate/nitrite, ammonium and organic nitrogen in wetted BSCs were high in comparison with those typical of aridosoils. They remained stable during incubations, indicating that input and output processes were of similar magnitude. Areal N(2) fixation rates (6.5-48 micromol C(2)H(2) m(-2) h(-1)) were high, the vertical distribution of N(2) fixation peaking close to the surface if populations of heterocystous cyanobacteria were present, but in the subsurface if they were absent. Areal AO rates (19-46 micromol N m(-2) h(-1)) were commensurate with N(2) fixation inputs. When considering oxygen availability, AO activity invariably peaked 2-3 mm deep and was limited by oxygen (not ammonium) supply. Most probable number (MPN)-enumerated ammonia-oxidizing bacteria (6.7-7.9 x 10(3) cells g(-1) on average) clearly peaked at 2-3 mm depth. Thus, AO (hence nitrification) is a spatially restricted but important process in the nitrogen cycling of BSC, turning much of the biologically fixed nitrogen into oxidized forms, the fate of which remains to be determined.     

Simultaneous measurement of denitrification and nitrogen fixation using isotope pairing with membrane inlet mass spectrometry analysis

A method for estimating denitrification and nitrogen fixation simultaneously in coastal sediments was developed. An isotope-pairing technique was applied to dissolved gas measurements with a membrane inlet mass spectrometer (MIMS). The relative fluxes of three N(2) gas species ((28)N(2), (29)N(2), and (30)N(2)) were monitored during incubation experiments after the addition of (15)NO(3)(-). Formulas were developed to estimate the production (denitrification) and consumption (N(2) fixation) of N(2) gas from the fluxes of the different isotopic forms of N(2). Proportions of the three isotopic forms produced from (15)NO(3)(-) and (14)NO(3)(-) agreed with expectations in a sediment slurry incubation experiment designed to optimize conditions for denitrification. Nitrogen fixation rates from an algal mat measured with intact sediment cores ranged from 32 to 390 microg-atoms of N m(-2) h(-1). They were enhanced by light and organic matter enrichment. In this environment of high nitrogen fixation, low N(2) production rates due to denitrification could be separated from high N(2) consumption rates due to nitrogen fixation. Denitrification and nitrogen fixation rates were estimated in April 2000 on sediments from a Texas sea grass bed (Laguna Madre). Denitrification rates (average, 20 microg-atoms of N m(-2) h(-1)) were lower than nitrogen fixation rates (average, 60 microg-atoms of N m(-2) h(-1)). The developed method benefits from simple and accurate dissolved-gas measurement by the MIMS system. By adding the N(2) isotope capability, it was possible to do isotope-pairing experiments with the MIMS system.     

Nitrogenase activity by biological soil crusts in cold sagebrush steppe ecosystems

In drylands worldwide, biological soil crusts (BSC) form a thin photosynthetic cover across landscapes, and provide vital benefits in terms of stabilizing soil and fixing nitrogen (N) and carbon (C). Numerous studies have examined the effects of climate and disturbance on BSC functions; however, few have characterized these responses in rolling BSCs typical of northern ecosystems in the Intermountain West, US. With temperature increases and shifts in precipitation projected, it is unclear how BSCs in this region will respond to climate change, and how the response could affect their capacity to perform key ecosystem functions, such as providing ‘new’ N through biological N₂ fixation. To address this important knowledge gap, we examined nitrogenase activity (NA) associated with rolling BSCs along a climatic gradient in southwestern Idaho, US, and quantified how acetylene reduction rates changed as a function of climate, grazing (using exclosures), and shrub-canopy association. Results show that warmer, drier climates at lower elevations hosted greater cover of late successional BSC communities (e.g., mosses and lichens), and higher NA compared with colder, wetter climates at higher elevations. Highest NA (0.5–29.3 µmol C₂H₄ m^?2 h^?1) occurred during the early summer/spring, when water was more available than in late summer/autumn. Activity was strongly associated with soil characteristics including pH and ammonium concentrations suggesting these characteristics as potentially strong controls on NA in BSCs. The relationship between grazing and NA varied with elevation. Specifically, lower elevation sites had lower NA at grazed locations, whereas higher elevation sites had higher NA with grazing. At both low and high ends of the elevation gradient, shrub-canopy associated BSCs maintained two to three times higher NA compared to BSCs in the interspace among shrubs. Taken together, our findings indicate that the controls and rates of NA in BSCs vary seasonally and strongly with climate in the Intermountain West, and that drier springs are likely to influence rates of NA more than warmer summers.     

Effect of biological soil crusts on soil elemental concentrations: implications for biogeochemistry and as traceable biosignatures of ancient life on land

Biological soil crusts (BSCs) are topsoil biosedimentary structures built by photosynthetic microbes commonly found today on arid soils. They play a role in soil stabilization and the fertility of arid lands, and are considered modern analogues of ancient terrestrial microbial communities. We determined the concentrations of four biogenic and 21 other elements, mostly metals, in surface soils that hosted BSCs, in the soils underneath those crusts, and in proximate but non-crusted surface soils. The samples were from six sites in the Colorado Plateau highlands and the Sonoran Desert lowlands. In spite of the variability in climate and geologic setting, we found statistically significant overall trends of enrichment in biogenic elements and depletion in non-biogenic elements when BSCs were compared with non-crusted soils. The differences between crusted and non-crusted soils were statistically significant at ∼95% confidence for C, N (enrichments) and for Ca, Cr, Mn, Cu, Zn, As, and Zr (depletions). These trends are best explained by the activity of microbes. As expected, no differences in the concentrations of C, N, P, and S were detected between the soils underneath the crusts and the non-crusted soils, but the former showed depletion of non-biogenic elements, indicating that the leaching effect of crust microbes extends downward in the soil. These patterns speak to the need for a sustained input of allochthonous material, possibly dust, to maintain BSC fertility. These elemental patterns can be considered a biosignature that may be preserved in the rock record and might help identify ancient microbial communities on land.     

沙坡头地区生物土壤结皮的固氮活性及其对水热因子的响应

氮是除水分之外影响干旱区生态系统生物活性的关键因子。生物土壤结皮是干旱半干旱荒漠地表景观的重要组成部分, 也是荒漠生态系统氮素的主要贡献者。通过野外调查采样, 利用开顶式生长室, 模拟不同降水梯度, 采用乙炔还原法连续测定了沙坡头地区典型生物土壤结皮(藻类结皮、地衣结皮和藓类结皮)在其主要固氮活跃期(6-10月, 湿润期)的固氮活性, 及其对水热因子的响应特征。结果表明, 试验期三类生物土壤结皮的固氮活性介于2.5 × 10³-6.2 × 10⁴ nmol C₂H₄·m^-2·h^-1之间, 其中藻类结皮的最高(平均达2.8 × 10⁴ nmol C₂H₄·m^-2·h^-1), 地衣结皮的次之(2.4 × 10⁴ nmol C₂H₄ ·m^-2·h^-1), 藓类结皮的最低(1.4 × 10⁴ nmol C₂H₄·m^-2·h^-1), 差异显著(p < 0.001)。在模拟降水3 mm时, 三类结皮均可达到最大固氮速率, 当发生> 3 mm的降水事件时, 它们的固氮速率无显著增加; 不同结皮的固氮活性与温度均呈显著的负相关关系(r_藻类结皮 = -0.711, r_地衣结皮 = -0.732, r_藓类结皮 = -0.755, p < 0.001), 藻类和藓类结皮的固氮活性的最适温度区间为25-30 ℃, 地衣结皮为20-30 ℃。三类结皮之间的这种固氮差异主要归因于结皮组成生物体即隐花植物的差异, 藻类结皮主要成分为大量的蓝细菌和一些绿藻, 地衣结皮也由大量的固氮藻和真菌共生形成, 而藓类结皮的主要组成部分苔藓植物并不具有固氮作用, 其微弱的固氮量是结皮中混生的少量蓝细菌或地衣所致。     

Co-occurrence of in-stream nitrogen fixation and denitrification across a nitrogen gradient in a western U.S. watershed

It is frequently assumed that nitrogen (N₂) fixation and denitrification do not co-occur in streams because each process should be favored under different concentrations of dissolved inorganic nitrogen (DIN), and therefore these processes are rarely quantified together. We asked if these processes could co-exist by conducting a spatial survey of N₂ fixation using acetylene reduction and denitrification using acetylene block [with and without amendments of carbon (C) as glucose and nitrogen (N) as nitrate]. Rates were measured on rocks and sediment in 8 southeastern Idaho streams encompassing a DIN gradient of 26–615 µg L^?1. Sampling at each site was repeated in summer 2015 and 2016. We found that both denitrification and N₂ fixation occurred across the gradient of DIN concentrations, with N₂ fixation occurring primarily on rocks and denitrification occurring in sediment. N₂ fixation rates on rocks significantly decreased 100× across the DIN gradient in 1 year of the study, and amended (with N and C) denitrification rates increased 10× across the DIN gradient in both years. Multiple linear regression and partial least squares models with environmental characteristics measured at the scale of entire stream reaches showed that C and phosphorus were positive predictors of amended and unamended denitrification rates, but no significant model could explain N₂ fixation rates across all streams and years. This, coupled with the observation that detectable rates of N₂ fixation occurred primarily on rocks and denitrification occurred primarily on sediment, suggests that microhabitat scale factors may better predict the co-occurrence of these processes within stream reaches. Overlooking the potential co-occurrence of N₂ fixation and denitrification in stream ecosystems will impede understanding by oversimplifying the contribution of each process to the N cycle.     

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.     

Denitrification and Nitrogen Fixation in Alaskan Continental Shelf Sediments

Rates of nitrogen fixation and denitrification were measured in Alaskan continental shelf sediments. In some regions, rates of nitrogen fixation and denitrification appeared to be equal; in other areas, rates were significantly different. Potential rates of denitrification were found to be limited primarily by the available nitrate substrate. Major regional differences in rates of denitrification were not statistically significant, but significant differences were found for nitrogen fixation rates in different regions of the Alaskan continental shelf. Estimated net losses of nitrogen from Bering Sea sediments were calculated as 1.8 × 10¹² g of N/yr. Experimental exposure of continental shelf sediments to petroleum hydrocarbons reduced rates of nitrogen fixation and denitrification in some cases but not others. Long-term exposure was necessary before a reduction in nitrogen fixation rates was observed; unamended rates of denitrification but not potential denitrification rates (NO₃⁻ added) were depressed after exposure to hydrocarbons.     

Biological soil crusts increase the resistance of soil nitrogen dynamics to changes in temperatures in a semi-arid ecosystem

Aims

Biological soil crusts (BSCs), composed of mosses, lichens, liverworts and cyanobacteria, are a key component of arid and semi-arid ecosystems worldwide, and play key roles modulating several aspects of the nitrogen (N) cycle, such as N fixation and mineralization. While the performance of its constituent organisms largely depends on moisture and rainfall conditions, the influence of these environmental factors on N transformations under BSC soils has not been evaluated before.

Methods

The study was done using soils collected from areas devoid of vascular plants with and without lichen-dominated BSCs from a semi-arid Stipa tenacissima grassland. Soil samples were incubated under different temperature (T) and soil water content (SWC) conditions, and changes in microbial biomass-N, dissolved organic nitrogen (DON), amino acids, ammonium, nitrate and both inorganic N were monitored. To evaluate how BSCs modulate the resistance of the soil to changes in T and SWC, we estimated the Orwin and Wardle Resistance index.

Results

The different variables studied were more affected by changes in T than by variations in SWC at both BSC-dominated and bare ground soils. However, under BSCs, a change in the dominance of N processes from a net nitrification to a net ammonification was observed at the highest SWC, regardless of T.

Conclusions

Our results suggest that the N cycle is more resistant to changes in T in BSC-dominated than in bare ground areas. They also indicate that BSCs could play a key role in minimizing the likely impacts of climate change on the dynamics of N in semi-arid environments, given the prevalence and cover of these organisms worldwide.     

Diurnal variation of rates of denitrification and nitrogen fixation of periphyton associated with Oryza glumaepatula Steud in an Amazonian Lake

The rates of denitrification and biological nitrogen fixation of the periphyton communities associated with the culms and adventitious root of Oryza glumaepatula in Lake Batata, State of Pará, Brazilian Amazonia, were investigated over the course of 24 h. N₂O was not detected during the period, demonstrating that denitrification was not occurring. The low concentration of nitrate in the water column was indicated as the main factor limiting the denitrification process. The rates of biological nitrogen fixation (BNF) in culms were lowest at night and early morning, while the highest values were measured at 1200 h. BNF rates in periphyton associated with adventitious roots were lowest during the morning, while highest values were measured at 1600 h. The difference in the daily patterns of BNF rates in the periphyton of culms and roots of O. glumaepatula in the euphotic zone was attributed to defferences in the availability of organic carbon which supports this process. In the periphyton of the culms, the BNF process was carried out by heterocystous cyanobacteria, which was photoautotrophs. In the periphyton of the adventitious roots, BNF was carried out by periphytic bacteria, and is heterotrophically mediated. This revised version was published online in July 2006 with corrections to the Cover Date.     

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。     

黄土高原生物结皮覆盖对风沙土和黄绵土溶质运移的影响

干旱和半干旱地区生物结皮的普遍发育显著改变了表层土壤的结构与养分富集特征,但其对土壤养分迁移和淋失的影响目前尚不明确。本研究针对黄土高原风沙土和黄绵土上发育的藓类生物结皮,以Ca²⁺和Cl^-为示踪离子开展溶质穿透试验,对有无生物结皮层及其覆盖下不同深度土壤的溶质运移特征进行了研究。结果表明: 在0～5 cm土层,生物结皮覆盖延缓了风沙土和黄绵土的溶质穿透过程,其Cl^-的穿透时间比无结皮延长了3.83(风沙土)和2.09倍(黄绵土),而Ca²⁺则分别延长了2.50和2.73倍。生物结皮覆盖条件下,表层0～5 cm土壤溶质完全穿透所对应的孔隙体积数比下层5～10 cm土壤更高,且其穿透历时更长;其中,Cl^-的穿透时间分别增加了67.3%(风沙土)和51.8%(黄绵土),Ca²⁺的穿透时间分别增加了8.0%和33.7%。生物结皮覆盖降低了土壤孔隙水流速(37.5%～70.2%);除风沙土的5～10 cm土层外,生物结皮使溶质弥散系数提高了1.73～6.29倍,使溶质弥散度提高了2.77～20.95倍。置换液完全穿透土壤后,风沙土和黄绵土的生物结皮层Ca²⁺含量显著高于无结皮,其分别比无结皮高4.14和2.58倍。研究证实,生物结皮覆盖能够提高表层土壤对养分的吸附与固持能力,从而减少土壤表聚养分的深层渗漏和流失,对干旱和半干旱地区退化土壤的肥力提升与植被恢复具有重要意义。     

Denitrification and nitrogen fixation dynamics in the area surrounding an individual ghost shrimp (Neotrypaea californiensis) burrow system

Bioturbated sediments are thought of as areas of increased denitrification or fixed-nitrogen (N) loss; however, recent studies have suggested that not all N may be lost from these environments, with some N returning to the system via microbial dinitrogen (N(2)) fixation. We investigated denitrification and N(2) fixation in an intertidal lagoon (Catalina Harbor, CA), an environment characterized by bioturbation by thalassinidean shrimp (Neotrypaea californiensis). Field studies were combined with detailed measurements of denitrification and N(2) fixation surrounding a single ghost shrimp burrow system in a narrow aquarium (15 cm by 20 cm by 5 cm). Simultaneous measurements of both activities were performed on samples taken within a 1.5-cm grid for a two-dimensional illustration of their intensity and distribution. These findings were then compared with rate measurements performed on bulk environmental sediment samples collected from the lagoon. Results for the aquarium indicated that both denitrification and N(2) fixation have a patchy distribution surrounding the burrow, with no clear correlation to each other, sediment depth, or distance from the burrow. Field denitrification rates were, on average, lower in a bioturbated region than in a seemingly nonbioturbated region; however, replicates showed very high variability. A comparison of denitrification field results with previously reported N(2) fixation rates from the same lagoon showed that in the nonbioturbated region, depth-integrated (10 cm) denitrification rates were higher than integrated N(2) fixation rates (～9 to 50 times). In contrast, in the bioturbated sediments, depending on the year and bioturbation intensity, some (～6.2%) to all of the N lost via denitrification might be accounted for via N(2) fixation.     

Nitrogen Fixation and Leaching of Biological Soil Crust Communities in Mesic Temperate Soils

Biological soil crust is composed of lichens, cyanobacteria, green algae, mosses, and fungi. Although crusts are a dominant source of nitrogen (N) in arid ecosystems, this study is among the first to demonstrate their contribution to N availability in xeric temperate habitats. The study site is located in Lucas County of Northwest Ohio. Using an acetylene reduction technique, we demonstrated potential N fixation for these crusts covering sandy, acidic, low N soil. Similar fixation rates were observed for crust whether dominated by moss, lichen, or bare soil. N inputs from biological crusts in northwestern Ohio are comparable to those in arid regions, but contribute substantially less N than by atmospheric deposition. Nitrate and ammonium leaching from the crust layer were quantified using ion exchange resin bags inserted within intact soil cores at 4 cm depth. Leaching of ammonium was greater and nitrate less in lichen than moss crusts or bare soil, and was less than that deposited from atmospheric sources. Therefore, biological crusts in these mesic, temperate soils may be immobilizing excess ammonium and nitrate that would otherwise be leached through the sandy soil. Moreover, automated monitoring of microclimate in the surface 7 cm of soil suggests that moisture and temperature fluctuations in soil are moderated under crust compared to bare soil without crust. We conclude that biological crusts in northwestern Ohio contribute potential N fixation, reduce N leaching, and moderate soil microclimate.     

Patterns of diversity for fungal assemblages of biological soil crusts from the southwestern United States

Molecular methodologies were used to investigate fungal assemblages of biological soil crusts (BSCs) from arid lands in the southwestern United States. Fungal diversity of BSCs was assessed in a broad survey that included the Chihuahuan and Sonoran deserts as well as the Colorado Plateau. At selected sites samples were collected along kilometer-scale transects, and fungal community diversity and composition were assessed based on community rRNA gene fingerprinting using PCR-denaturing gradient gel electrophoresis (DGGE). Individual phylotypes were characterized through band sequencing. The results indicate that a considerable diversity of fungi is present within crusted soils, with higher diversity being recovered from more successionally mature BSCs. The overwhelming majority of crust fungi belong to the Ascomycota, with the Pleosporales being widespread and frequently dominant. Beta diversity patterns of phylotypes putatively representing dominant members of BSC fungal communities suggest that these assemblages are specific to their respective geographic regions of origin.     

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.     

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...     

Global change and biological soil crusts: effects of ultraviolet augmentation under altered precipitation regimes and nitrogen additions

Biological soil crusts (BSCs), a consortium of cyanobacteria, lichens, and mosses, are essential in most dryland ecosystems. As these organisms are relatively immobile and occur on the soil surface, they are exposed to high levels of ultraviolet (UV) radiation and atmospheric nitrogen (N) deposition, rising temperatures, and alterations in precipitation patterns. In this study, we applied treatments to three types of BSCs (early, medium, and late successional) over three time periods (spring, summer, and spring–fall). In the first year, we augmented UV and altered precipitation patterns, and in the second year, we augmented UV and N. In the first year, with average air temperatures, we saw little response to our treatments except quantum yield, which was reduced in dark BSCs during one of three sample times and in Collema BSCs two of three sample times. There was more response to UV augmentation the second year when air temperatures were above average. Declines were seen in 21% of the measured variables, including quantum yield, chlorophyll a, UV‐protective pigments, nitrogenase activity, and extracellular polysaccharides. N additions had some negative effects on light and dark BSCs, including the reduction of quantum yield, β‐carotene, nitrogenase activity, scytonemin, and xanthophylls. N addition had no effects on the Collema BSCs. When N was added to samples that had received augmented UV, there were only limited effects relative to samples that received UV without N. These results indicate that the negative effect of UV and altered precipitation on BSCs will be heightened as global temperatures increase, and that as their ability to produce UV‐protective pigments is compromised, physiological functioning will be impaired. N deposition will only ameliorate UV impacts in a limited number of cases. Overall, increases in UV will likely lead to lowered productivity and increased mortality in BSCs through time, which, in turn, will reduce their ability to contribute to the stability and fertility of soils in dryland regions.     

基于CCA的坡面尺度生物结皮空间分布

生物结皮是干旱半干旱地区普遍存在的活地被物,在该区生态系统中具有重要的生态功能。探讨生物结皮的空间分布规律是科学管理该资源的理论基础。选择黄土高原水蚀风蚀交错区六道沟小流域内的典型坡面,通过全面调查并应用GS+和CANOCO统计软件进行分析,探讨了坡面尺度上生物结皮的空间分布特征及其影响因子。结果表明:(1)生物结皮的分布具有明显的空间分异性。沙土区生物结皮以大面积连续分布为主,平均结皮盖度在30%以上;在黄土区则以零星分布为主,结皮盖度大都在20%以下,主要分布在坡的边缘和末端。而生物结皮厚度和抗剪强度的空间变异不大,说明其主要与生物结皮的发育年限有关。(2)典范对应分析(CCA)表明:生物结皮的空间分布与土壤、植被、地表湿度、坡度坡向等有密切关系。其中,土壤类型和生物结皮的空间分布关系最大,可以解释生物结皮空间变异的20%。其次是植被群落类型和地形湿度指数,沙蒿(Artemisia desertorum Spreng.Syst.Veg.)地和小叶杨(Populus simonii Carr.)林地是其最理想的生长环境;其他如坡度、坡向、太阳辐射等也都对生物结皮的分布有一定的影响。总体上,生物结皮具有明显的地形、土壤和植被群落的选择性,偏向于生长在较湿润的沙生植被群落当中。