Export of nitrogenous compounds due to incomplete cycling within biological soil crusts of arid lands

Second only to water among limiting factors, nitrogen controls the fertility of most arid regions. Where dry and wet depositions are weak, as in the western US deserts, N inputs rely heavily on biological N(2) fixation. Topsoil cyanobacterial communities known as biological soil crusts (BSCs) are major N(2) fixation hot spots in arid lands, but the fate of their fixed N remains controversial. Using a combination of microscale and mesoscale process rate determinations, we found that, in spite of theoretically optimal conditions, denitrification rates in BSCs were paradoxically immaterial for nitrogen cycling. Denitrifier populations within BSCs were extremely low. Because of this absence of denitrification, and because of the limitation of respiration and ammonia oxidation by diffusive O(2) supply, we could demonstrate that BSCs function as net exporters of ammonium, nitrate and organic N to the soils they cover, in approximately stoichiometrically equal proportions. Overall export rates during periods of biological activity are in the range of tens to hundreds of mumol-N m(-2) h(-1), commensurate with those of N(2) fixation. These results explain the long-term dependence of BSCs on N(2) fixation, confirm their role in landscape fertility, and provide a robust argument for conservation of these endangered communities.     

High rates of denitrification and nitrous oxide emission in arid biological soil crusts from the Sultanate of Oman

Using a combination of process rate determination, microsensor profiling and molecular techniques, we demonstrated that denitrification, and not anaerobic ammonium oxidation (anammox), is the major nitrogen loss process in biological soil crusts from Oman. Potential denitrification rates were 584±101 and 58±20 μmol N m⁻² h⁻¹ for cyanobacterial and lichen crust, respectively. Complete denitrification to N₂ was further confirmed by an ¹⁵NO₃⁻ tracer experiment with intact crust pieces that proceeded at rates of 103±19 and 27±8 μmol N m⁻² h⁻¹ for cyanobacterial and lichen crust, respectively. Strikingly, N₂O gas was emitted at very high potential rates of 387±143 and 31±6 μmol N m⁻² h⁻¹ from the cyanobacterial and lichen crust, respectively, with N₂O accounting for 53–66% of the total emission of nitrogenous gases. Microsensor measurements revealed that N₂O was produced in the anoxic layer and thus apparently originated from incomplete denitrification. Using quantitative PCR, denitrification genes were detected in both the crusts and were expressed either in comparable (nirS) or slightly higher (narG) numbers in the cyanobacterial crusts. Although 99% of the nirS sequences in the cyanobacterial crust were affiliated to an uncultured denitrifying bacterium, 94% of these sequences were most closely affiliated to Paracoccus denitrificans in the lichen crust. Sequences of nosZ gene formed a distinct cluster that did not branch with known denitrifying bacteria. Our results demonstrate that nitrogen loss via denitrification is a dominant process in crusts from Oman, which leads to N₂O gas emission and potentially reduces desert soil fertility.     

Germination and seed water status of four grasses on moss-dominated biological soil crusts from arid lands

Biological soil crusts dominated by drought-tolerant mosses are commonly found through arid and semiarid steppe communities of the northern Great Basin of North America. We conducted growth chamber experiments to investigate the effects of these crusts on the germination of four grasses: Festuca idahoensis, Festuca ovina, Elymus wawawaiensis and Bromus tectorum. For each of these species, we recorded germination time courses on bare soil and two types of biological soil crusts; one composed predominantly of the tall moss Tortula ruralis and the other dominated by the short moss Bryum argenteum. On the short-moss crust, the final germination percentage was about half of that on bare soil. Also, the mean germination time was 4 days longer on short-mosses than on bare soil. In contrast to the short-moss crust, the tall-moss crust did not reduce the final germination percentage but increased the mean germination time. Similar results were observed in the four grasses studied. To investigate the mechanism by which moss crusts affected germination, we analyzed the water status of seeds on bare soil and moss crusts. Six days after seeding, the water content of seeds on bare soil was approximately twice that of seeds on tall- or short-moss crust. Analysis of the time course of changes in seed weight and water potential in Bromus tectorum revealed that overtime seeds on tall mosses reached higher water content than those on short mosses. The increase in the water content of seeds on tall mosses occurred as the seeds gradually fell through the moss canopy. Taken together, our results indicate that biological soil crusts with distinct structural characteristics can have different effects on seed germination. Furthermore, this study revealed that a biological soil crust dominated by short mosses had a negative effect on seed water status and significantly reduced seed germination.     

Microbial diversity of eolian dust sources from saline lake sediments and biological soil crusts in arid Southern Australia

While microbial communities of aerosols have been examined, little is known about their sources. Nutrient composition and microbial communities of potential dust sources, saline lake sediments (SLS) and adjacent biological soil crusts (BSC), from Southern Australia were determined and compared with a previously analyzed dust sample. Multivariate analyses of fingerprinting profiles indicated that the bacterial communities of SLS and BSC were different, and these differences were mainly explained by salinity. Nutrient concentrations varied among the sites but could not explain the differences in microbial diversity patterns. Comparison of microbial communities with dust samples showed that deflation selects against filamentous cyanobacteria, such as the Nostocales group. This could be attributed to the firm attachment of cyanobacterial filaments to soil particles and/or because deflation occurs mainly in disturbed BSC, where cyanobacterial diversity is often low. Other bacterial groups, such as Actinobacteria and the spore-forming Firmicutes, were found in both dust and its sources. While Firmicutes-related sequences were mostly detected in the SLS bacterial communities (10% of total sequences), the actinobacterial sequences were retrieved from both (11-13%). In conclusion, the potential dust sources examined here show highly diverse bacterial communities and contain nutrients that can be transported with aerosols. The obtained fingerprinting and sequencing data may enable back tracking of dust plumes and their microorganisms.     

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.     

踩踏干扰对生物结皮土壤渗透性的影响

采用人工模拟降雨试验,研究了踩踏干扰对生物结皮土壤渗透性的影响.结果表明:踩踏干扰显著增加了土壤表面粗糙度,增加幅度与干扰强度有关,50%干扰度下表面粗糙度指数较不干扰增加91%.踩踏干扰延长了坡面产流时间,20%～50%干扰度范围内,随着干扰强度的增加,初始产流时间呈线性增加趋势,50%干扰度的初始产流时间较不干扰增加了169.7%.踩踏干扰增加了土壤渗透性,降低径流系数.50%干扰度的土壤累积入渗量较不干扰增加12.6%;去除生物结皮,土壤渗透性降低,累积入渗量较不干扰降低30.2%.50%以下的干扰度未显著增加土壤侵蚀模数.去除生物结皮,土壤侵蚀模数较不干扰增加10倍.生物结皮破碎度低于50%的干扰在不明显增加土壤流失量的前提下,可增加降水入渗,减小径流风险,改善土壤水分状况.     

干旱沙区生物土壤结皮对红砂种子定居的影响及作用机制

为了探清干旱荒漠区生物土壤结皮对易漂移性植物种子定居影响及作用机制,以巴丹吉林沙漠南缘结皮前期以及藻类、地衣、藓类阶段结皮为场所,通过人工补充种子与野外长期监测的方法对红砂种子在生物结皮上流失动态、种子宿存微区特征以及种子宿存量与微区面积间的关系等进行研究。结果表明：(1)生物土壤结皮发育过程中,地表枯落物、微坑、龟裂缝、长藓的面积百分比均发生了显著变化,即：枯落物呈现出增大(藻类阶段)—稳定不变(地衣阶段)—减小(藓类阶段)的趋势,微坑、龟裂缝均先增大后减小(峰值在地衣阶段),长藓持续增大。(2)4个阶段的种子流失速度或定居率两两之间存在显著差异,流失速度大小依次为：结皮前期>藻类阶段>藓类阶段>地衣阶段,定居率大小为：地衣阶段>藓类阶段>藻类阶段>结皮前期。(3)生物土壤结皮上定居的种子仅宿存于枯落物、微坑、龟裂缝、长藓上,这4种微区上定居种子数与其面积之间均存在显著正相关关系,其关系均可用二次项y=ax²+bx+c(a≠0)表示,等面积微区内的定居种子数大小依次为：龟裂缝>枯落物>微坑>长藓。因此认为,生...     

The presence of oxygen in rumen liquor and its effects on methanogenesis

In situ measurement of O₂ in the rumen liquor of cows, sheep and goats using a membrane-covered O₂ electrode revealed the presence of up to 1630 nmol/l O₂; O₂ became undetectable immediately after feeding of animals. The effects of O₂ on H₂ production and methanogenesis in samples of rumen liquor were investigated using a mass spectrometer fitted with a membrane inlet system. Methanogenesis was totally and irreversibly inhibited after short term exposure (about 10 min) to 5 KPa (0·05 atm) O₂; H₂ production was unaffected. Glucose additions produced rapid transient increases in H₂ levels and increased O₂ uptake.     

Progress in the study of algae and mosses in biological soil crusts

Algae and mosses are not only two of the familiar communities in the process of desert vegetational succession, but also have the highest biomass in biological soil crusts. Meanwhile, being the pioneer plants, algae and mosses are involved in the establishment of biological soil crusts, which have great importance in arid environments and play a major role in desert ecosystems, such as being the indicator of the vegetation type, soil-holding, preventing erosion by water and wind, and sand fixation. This paper reviews the advances in the study of algae and mosses in arid and semi-arid areas. It mainly describes the ecological functions of algae and mosses including their influences on water cycle, circulation of substances, and community succession. In addition, the relationships between algae and mosses are discussed. Finally, some suggestions are proposed for the research orientations of algae and mosses in biological soil crusts. Ecologically, algae and mosses have significant ecological importance in arid areas, especially in those areas where environmental problems are becoming increasingly serious.     

干旱沙区生物土壤结皮发育对红砂形态及干物质积累的影响

为了探讨干旱沙区生物土壤结皮发育对红砂形态及干物质积累的影响,以巴丹吉林沙漠南缘已发育不同类型生物土壤结皮并有红砂种群成功定居的区域为研究场所,通过野外监测与室内测定的方法对藻类-地衣、地衣、地衣-藓类结皮上的红砂（当年生、幼株、成株）形态特征及生物量进行了调查研究。结果表明：（1）藻类-地衣结皮演替到地衣-藓类结皮的过程中,当年生红砂形态差异不显著,但5 a以上植株基部分枝长、树冠/侧冠投影面积、主根长均显著减小;同时,地衣-藓类结皮的3-5 a植株基部分枝数明显减少,且5 a以上植株明显矮化。（2）生物土壤结皮发育不仅降低了红砂幼株或成株生物量积累能力,还减小了植株根冠比,且降低/减小程度随结皮演替或株龄的增大逐渐增大。（3）红砂形态、生物量指标与物理或藻类结皮面积百分比呈极显著正相关关系,与藓类结皮面积百分比呈极显著的负相关关系。因此认为,生物土壤结皮的演替导致土壤关键生态因子（如土壤水分）发生变化,进而影响红砂植株生长发育能力,从而使得不同发育阶段结皮上的同龄红砂形态特征及生物量存在差异性。     

Progress in the study of algae and mosses in biological soil crusts

Algae and mosses are not only two of the familiar communities in the process of desert vegetational succession,but also have the highest biomass in biological soil crusts.Meanwhile,being the pioneer plants,algae and mosses are involved in the establishment of biological soil crusts,which have great importance in arid environments and play a major role in desert ecosystems,such as being the indicator of the vegetation type,soil-holding,preventing erosion by water and wind,and sand fixation.This paper reviews the advances in the study of algae and mosses in arid and semi-arid areas.It mainly describes the ecological functions of algae and mosses including their influences on water cycle,circulation of substances,and community succession.In addition,the relationships between algae and mosses are discussed.Finally,some suggestions are proposed for the research orientations of algae and mosses in biological soil crusts.Ecologically,algae and mosses have significant ecological importance in arid areas,especially in those areas where environmental problems are becoming increasingly serious.     

生物土壤结皮的分布影响因子及其监测

生物土壤结皮在荒漠化地区广泛分布。本文详细论述了生物土壤结皮的分布规律及其影响因素,如海拔高度、土壤、维管植物群落水分条件和干扰,以及生物土壤结皮在生态系统和景观变化监测和评价中的作用等。同时对中国西北地区生物土壤结皮的生态学研究和开发应用提供了研究的重点和方向。     

Response of desert biological soil crusts to alterations in precipitation frequency

Biological soil crusts, a community of cyanobacteria, lichens, and mosses that live on the soil surface, occur in deserts throughout the world. They are a critical component of desert ecosystems, as they are important contributors to soil fertility and stability. Future climate scenarios predict alteration of the timing and amount of precipitation in desert environments. Because biological soil crust organisms are only metabolically active when wet, and as soil surfaces dry quickly in deserts during late spring, summer, and early fall, the amount and timing of precipitation is likely to have significant impacts on the physiological functioning of these communities. Using the three dominant soil crust types found in the western United States, we applied three levels of precipitation frequency (50% below-average, average, and 50% above-average) while maintaining average precipitation amount (therefore changing both timing and size of applied events). We measured the impact of these treatments on photosynthetic performance (as indicated by dark-adapted quantum yield and chlorophyll a concentrations), nitrogenase activity, and the ability of these organisms to maintain concentrations of radiation-protective pigments (scytonemin, beta-carotene, echinenone, xanthophylls, and canthaxanthin). Increased precipitation frequency produced little response after 2.5 months exposure during spring (1 April–15 June) or summer (15 June–31 August). In contrast, most of the above variables had a large, negative response after exposure to increased precipitation frequency for 6 months spring–fall (1 April–31 October) treatment. The crusts dominated by the soil lichen Collema, being dark and protruding above the surface, dried the most rapidly, followed by the dark surface cyanobacterial crusts (Nostoc-Scytonema-Microcoleus), and then by the light cyanobacterial crusts (Microcoleus). This order reflected the magnitude of the observed response: crusts dominated by the lichen Collema showed the largest decline in quantum yield, chlorophyll a, and protective pigments; crusts dominated by Nostoc-Scytonema-Microcoleus showed an intermediate decline in these variables; and the crusts dominated by Microcoleus showed the least negative response. Most previous studies of crust response to radiation stress have been short-term laboratory studies, where organisms were watered and kept under moderate temperatures. Such conditions would give crust organisms access to ample carbon to respond to imposed stresses (e.g., production of UV-protective pigments, replacement of degraded chlorophyll). In contrast, our longer-term study showed that under field conditions of high air temperatures and frequent, small precipitation events, crust organisms appear unable to produce protective pigments in response to radiation stress, as they likely dried more quickly than when they received larger, less frequent events. Reduced activity time likely resulted in less carbon available to produce or repair chlorophyll a and/or protective pigments. Our findings may partially explain the global observation that soil lichen cover and richness declines as the frequency of summer rainfall increases.     

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...     

一种从土壤生物结皮中有效提取DNA的方法

在干旱、半干旱地区稀疏的高等植被间, 常有一层生物土壤结皮。       

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.     

高寒沙区生物土壤结皮覆盖土壤碳释放动态

生物土壤结皮广泛分布于荒漠生态系统,能够通过自身的呼吸作用影响土壤碳释放以及区域碳循环过程。本研究在具有典型高寒沙区气候特征的青藏高原东北部（青海共和盆地）,以广泛分布于人工植被恢复区的藻类和藓类结皮为研究对象,裸地为对照,观测了裸地与两种类型生物土壤结皮去除和覆盖土壤碳释放速率的日动态和生长季动态规律,探讨生物土壤结皮对土壤碳释放量的影响。结果表明：生物土壤结皮去除和覆盖土壤碳释放速率日动态和生长季动态特征与裸地一致,均呈“单峰”曲线。生物土壤结皮覆盖土壤的日最大碳释放速率出现于13:00左右,裸地与去除结皮土壤的日峰值均出现于15:00左右,生物土壤结皮的存在使土壤碳释放速率的日峰值出现时间提前2h左右,各观测类型生长季内碳释放速率最大值均出现在8月。在相对干旱年份（2017）,藻类和苔藓结皮覆盖导致土壤碳释放量分别增加了22.07%和85.61%,其中,藻类和苔藓结皮层碳释放量占增加量的67.60%和25.76%;而在相对湿润年份（2018）,藻类和苔藓结皮覆盖导致土壤碳释放量分别增加了139.37%和290.53%,二者结皮层碳释放量分别占增加量的69.09%和45.59%,生物土壤结皮发育促进了土壤的碳释放。温度对土壤碳释放变化的贡献率为48.89%,是高寒沙区土壤碳释放日动态变化的关键驱动因子。因此,在核算荒漠生态系统碳交换过程中,应充分考虑各区域不同类型生物土壤结皮对土壤碳释放产生的影响。     

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.