Simulated nitrogen deposition enhances the performance of an exotic grass relative to native serpentine grassland competitors

Previous research suggests that atmospheric nitrogen (N) deposition may facilitate the invasion and persistence of exotic plant species in serpentine grasslands, but the relative impact of increased N availability on native and exotic competitive dynamics has yet to be clearly elucidated. In this study, we evaluated how increased N deposition affects plant performance and competitive dynamics of five native grasses and forbs (Plantago erecta, Layia gaillardioides, Lasthenia californica, Vulpia microstachys, and Cryptantha flaccida) and the most common invasive grass in Bay Area serpentine grasslands, Lolium multiflorum. Using a growth chamber system, we exposed Lolium in monoculture, and native species grown both in monoculture and in competition with the exotic Lolium, to all four possible combinations of gaseous nitrogen dioxide (NO₂; a dominant atmospheric N pollutant) and soil ammonium nitrate (NH₄NO₃). In monocultures, gaseous NO₂ and soil N addition each increased shoot biomass in Lolium and the natives Layia and Cryptantha. Lolium competitive ability (mean relative yield potential??RYP) increased in response to NO₂ addition plus soil N addition against all native competitors. Lolium and most native species did not show differences in photosynthetic rate and stomatal conductance in response to N addition. Our findings indicate that increasing N deposition and subsequent N accumulation in the soil may confer a competitive advantage to the exotic Lolium over native species by stimulating greater biomass accumulation and N allocation to photosynthetic tissue in the invader.     

荒漠草地土壤微生物生物量和微生物熵对沙漠化的响应

采用空间序列代替时间演替的方法,分析宁夏中北部盐池县荒漠草地不同沙漠化阶段(荒漠草地、固定沙地、半固定沙地和流动沙地)土壤微生物生物量(SMB)和微生物熵(qMB)的变化特征及其影响因子.结果表明:从荒漠草地到流动沙地,土壤微生物生物量碳、氮、磷分别降低46.1%、80.8%和30.0%.随着荒漠草地沙漠化程度的加剧,土壤微生物熵碳(qMBC)、土壤微生物熵氮(qMBN)、土壤微生物熵磷(qMBP)均表现为荒漠草地>固定沙地>半固定沙地>流动沙地,而土壤-微生物化学计量不平衡性(C∶Nimb、C∶Pimb、N∶Pimb)基本呈增加趋势.土壤微生物生物量氮与C∶Nimb呈显著正相关,与N∶Pimb呈显著负相关;土壤微生物生物量磷与C∶Pimb呈显著正相关.冗余分析(RDA)显示,土壤生态化学计量(C∶N、C∶P)对微生物熵碳的负效应最明显.荒漠草地沙漠化显著影响土壤微生物生物量和微生物熵.     

长白山天然次生林土壤微生物量对氮沉降的响应

土壤微生物作为土壤养分的生物驱动因素,氮沉降会改变其活性和生物量,从而打破土壤养分循环动态平衡。氮沉降对热带、亚热带森林以及温带原始林生态系统土壤微生物量影响的研究较多,但对温带天然次生林影响的研究鲜有报道。于2016年5月(春)、7月(夏)和9月(秋)分别对长白山模拟10年氮沉降的控制试验样地——白桦山杨次生林进行了野外调查。控制试验分为3个氮添加处理,对照(CK 0 kg N hm~(-2)a~(-1))、低氮(LN 25 kg N hm~(-2)a~(-1))和高氮(HN 50 kg N hm~(-2)a~(-1)),按照土壤层(0—10 cm和10—20 cm)分别测试了不同处理的土壤微生物量碳(MBC)和氮(MBN)、土壤全碳(TC)、全氮(TN)和全磷(TP)、p H、土壤可溶性有机碳(DOC)和氮(DON)等指标。结果表明:1)土壤p H在氮沉降的作用下显著降低;上层土壤TC、TN在氮沉降下变化较小,下层土壤TC、TN的含量显著增加;氮沉降下春、夏两季土壤TP含量上升,LN处理在秋季对TP有抑制作用;氮沉降对DOC、DON的影响不显著。2)上层土壤MBC春季到秋季呈现递减的趋势,下层土壤呈现先升后降的趋势,HN对MBC有抑制作用,LN对下层土壤MBC有促进作用;土壤MBN由春季到秋季呈现递减的趋势,且上、下层土壤MBN差异显著;氮处理对春、秋两季MBN有促进作用,夏季有抑制作用;氮沉降使春、秋两季MBC/MBN降低,夏季土壤MBC/MBN升高。3)氮处理、季节变化和土层深度对MBC、MBN存在显著影响,其交互影响也显著。总之,长期氮沉降在生长季对土壤微生物量的影响具有季节性差异,且受到土层深度的影响。未来研究在重视年际变化的同时,也要注重时空动态对氮沉降作用表现出的差异性。     

模拟氮沉降对土壤酶活性和微生物组成的影响

氮沉降改变了草地生态系统的氮(N)素循环过程,由此带来的生态学效应已成为当前研究的热点。以乌鲁木齐周边短期围封草地为研究对象,通过模拟氮沉降实验,分析了自由放牧地和围封草地土壤酶活性和微生物组成,结合土壤养分及化学计量特征,探讨了氮沉降对短期围封草地土壤微生物组成及酶活性的影响,为该地区放牧草地的保护、恢复及管理提供理论依据。结果表明:(1)土壤有机碳(SOC)、全氮(TN)、全磷(TP)含量随围封年限的增加总体呈升高趋势,表明围封有利于提高土壤养分含量。与中国草地平均值相比,该草地土壤碳氮比(C/N)相对较高,碳磷比(C/P)、氮磷比(N/P)相对较低,表明该草地土壤有机质分解良好,有利于土壤碳(C)、磷(P)的释放,而土壤N素较为缺乏。(2)就不同围封年限而言,围封3年草地5-20cm层土壤真菌数量高于其它样地;围封3年草地表层土壤蔗糖酶与过氧化氢酶活性最高;围封7年草地放线菌数量最多,说明围封能够促进土壤微生物生长及酶活性的提高。(3)氮素添加对土壤真菌具有抑制作用,N5(4.6gN m^-2 a^-1)、N10(9.8gN m^-2 a^-1)处理显著增加了各样地土壤细菌数量,氮素添加对围封7年草地0-10cm层土壤放线菌无显著影响,而氮沉降显著增加了其它样地5-20cm层土壤放线菌数量,其中N5、N10处理下促进作用最明显;氮素添加对该草地土壤脲酶、蔗糖酶、过氧化氢酶均具有促进作用,N5、N10处理促进作用最明显。综合分析表明,氮沉降可直接或间接影响土壤微生物及酶活性,短期围封作为一种草地管理手段,对退化草地生态系统的修复具有一定作用,并可通过改善土壤理化性质、调节养分含量及其化学计量比来加速退化草地的恢复。     

不同环境条件下土壤微生物对模拟大气氮沉降的响应

研究了林内及林缘两个生境中,在有苔藓覆盖和无苔藓覆盖条件下,人工加氮对土壤理化性质及土壤微生物群落的影响。结果显示加氮使土壤pH下降,有效态氮和有效态磷的含量上升,但不同生境及有无苔藓植物覆盖在一定程度上影响土壤理化性质及其对加氮的反应。苔藓植物覆盖可以缓解加氮引起的土壤酸化及有效氮含量上升压力,促进有效态磷含量上升。不同生境中,土壤微生物对氮沉降的响应亦不同。低氮使林缘生境土壤微生物的胁迫程度减小,中高氮使其胁迫程度上升,而任何加氮均增加林内生境中土壤微生物的胁迫程度。两个生境中,苔藓植物覆盖均可以缓解过量氮沉降对土壤微生物造成的压力,降低过量氮沉降对土壤微生物的伤害,提高土壤微生物的代谢活性。     

藏北高寒草地植被和土壤对不同放牧强度的响应

放牧压力在时间和空间的不均衡分布导致局部土地过度利用和生态退化,然而当前对放牧强度的量化多采用替代性指标或对照试验,缺乏直接监测数据,也缺乏不同草地类型对放牧强度变化的响应差异性研究。以西藏自治区那曲市为研究区,利用佩戴式GPS牛羊定位器构建高精度放牧轨迹数据集模拟放牧强度,构建栅格尺度放牧强度空间分布和划分方法,结合草地群落样方调查,通过Duncan法（Duncan''s multiple comparative analysis）进行多重比较分析,探究自由放牧模式下高寒草甸和高寒草原两类区域植被和土壤对不同放牧强度的响应方式及差异性。本研究可为放牧行为环境效应监测提供新思路,并根据不同草地状况因地制宜提出放牧优化管理策略,助力高寒传统牧区的可持续发展。主要结论有：1）随着放牧强度的增大,高寒草原地上生物量先升高后降低,高强度放牧对高寒草原植被的影响大于高寒草甸。2）高放牧强度下,高寒草原土壤水分显著高于中低强度,高寒草甸土壤容重显著低于低强度。可能原因是放牧压力多集聚于水源附近。高寒草原区土壤的砂粒含量随放牧强度的增大而增加。放牧强度的增大导致草甸上层土壤有机质增加,草甸下层、草原上层、草原下层土壤有机质先增加后减小。草甸上下层土壤全磷含量在低放牧强度下显著低于中高强度。3）植被土壤变化受到自然因子和放牧活动的共同影响。高寒草甸更加耐牧,高强度放牧对高寒草原的负面影响更大,而中度放牧有利于草地尤其是高寒草原的放牧利用。4）放牧生态系统是一个环境-植物-家畜自适应系统。在放牧管理中不能仅通过控制载畜量缓解草地超载,还需要综合考虑生态系统的弹性及各营养级的适应性,合理配置放牧强度,控制季节性超载和局部超载。     

Invasive species cover,soil type,and grazing interact to predict long‐term grassland restoration success

Grasslands are undergoing tremendous degradation as a result of climate change, land use, and invasion by non‐native plants. However, understanding of the factors responsible for driving reestablishment of grassland plant communities is largely derived from short‐term studies. In order to develop an understanding of the factors responsible for longer term restoration outcomes in California annual grasslands, we surveyed 12 fields in Davis, CA, U.S.A., in 2015 that were seeded with native species mixtures starting in 2004. Using field surveys, we investigated how invasive plant richness and cover, native plant richness and cover, aboveground biomass, grazing, soil type, and restoration species identity might provide utility for explaining patterns of restoration success. We found a negative relationship between invasive cover and restoration cover, which was attributed to the slow establishment of seeded species and subsequent dominance by weeds. The relationship between invasive cover and restoration cover was modified by grazing, likely due to a change in the dominance of exotic forbs, which have a more similar growing season to restoration species, and therefore compete more strongly for late season moisture. Finally, we found that soil type was responsible for differences in the identity and abundance of invasive plants, subsequently affecting restoration cover. This work highlights the value of focusing resources on reducing invasive species cover, limiting grazing to periods of adequate moisture, and considering soil type for successful long‐term restoration in California annual grasslands. Moreover, observations of long‐term restoration outcomes can provide insight into the way mechanisms driving restoration outcomes might differ through time.     

Response of fungal, bacterial and ureolytic communities to synthetic sheep urine deposition in a grassland soil

In grazed pastures, soil pH is raised in urine patches, causing dissolution of organic carbon and increased ammonium and nitrate concentrations, with potential effects on the structure and functioning of soil microbial communities. Here we examined the effects of synthetic sheep urine (SU) in a field study on dominant soil bacterial and fungal communities associated with bulk soil and plant roots (rhizoplane), using culture-independent methods and a new approach to investigate the ureolytic community. A differential response of bacteria and fungal communities to SU treatment was observed. The bacterial community showed a clear shift in composition after SU treatment, which was more pronounced in bulk soil than on the rhizoplane. The fungal community did not respond to SU treatment; instead, it was more affected by the time of sampling. Redundancy analysis of data indicated that the variation in the bacterial community was related to change in soil pH, while fungal community was more responsive to dissolution of organic carbon. Like the universal bacterial community, the ureolytic community was influenced by the SU treatment. However, different taxa within the ureolytic bacterial community responded differentially to the treatment. The ureolytic community comprised of members from a range of phylogenetically different taxa and could be used to measure the effect of environmental perturbations on the functional diversity of natural ecosystems.     

湿地土壤微生物功能多样性及碳氮组分对长期氮输入的响应

氮输入对湿地生态系统碳氮循环具有重要影响,研究湿地土壤微生物功能多样性及碳氮组分对氮输入的响应,对于明确湿地土壤碳氮循环微生物驱动机制具有重要意义。依托长期野外氮输入模拟试验,利用Biolog-ECO微平板技术,分析不同浓度氮输入:N1(6 g N m^-2 a^-1)、N2(12 g N m^-2 a^-1)和N3(24 g N m^-2 a^-1)对湿地土壤表层(0-15 cm)和亚表层(15-30 cm)微生物碳源代谢活性、功能多样性和碳氮组分的影响。结果表明:N2处理显著提高了亚表层土壤微生物碳源代谢活性和McIntosh指数,N3处理显著降低了表层土壤微生物Shannon指数和Shannon-evenness指数。随氮输入浓度增加湿地表层土壤微生物对糖类的利用率显著降低,N3处理表层土壤微生物对胺类的利用率以及亚表层土壤微生物对醇类的利用率显著提高。N1处理显著提高了湿地表层土壤全氮和微生物量碳含量;N2、N3处理显著提高了土壤铵态氮、硝态氮含量;N3处理显著降低了土壤pH值。湿地土壤pH、总碳、溶解性有机碳含量是影响微生物碳源代谢活性和功能多样性的重要因素,土壤溶解性有机碳、铵态氮、全氮含量、含水率是影响微生物碳源利用变化的主要因子。     

克氏针茅草原土壤生态化学计量特征对放牧强度的响应

生态化学计量学是研究生态系统中各元素间平衡的科学,对于探究土壤有机碳（C）、全氮（N）、全磷（P）元素循环以及土壤养分限制状况具有重要意义。为了解克氏针茅（Stipa kirschnii）草原土壤生态化学计量特征,以内蒙古克氏针茅草原区为研究对象,在不同放牧强度草场进行采样分析。结果表明：随着放牧强度的增加,C、P含量先降低后升高,N含量先升高后降低;土壤C/N先降低后升高、C/P逐渐升高、N/P先升高后降低。C/N随着放牧强度的增加与有机碳相关性逐渐增大、与全氮的相关性逐渐减小;轻牧区的C/P受有机碳作用有所增强;中牧区N/P受全氮作用增强。C/N主要限制因素是有机碳含量,C/P和N/P主要限制因素是全磷含量。不同放牧强度会影响土壤化学计量比与其他物质的关系,也会改变土壤化学计量比各制约因素的强度。     

土壤跳虫对氮磷养分响应的研究进展

土壤跳虫是土壤分解者中最重要的类群之一,它们通过营养联系将基础食物资源中的物质和能量传递到更高营养级,在地下食物网中具有不可替代的作用。在全球氮磷沉降日趋严重的背景下,土壤跳虫群落如何响应氮(N)、磷(P)沉降的研究还十分有限,严重阻碍了我们对跳虫群落的生态功能、在地下食物网的物质循环和能量流动以及陆地碳(C)循环中作用的科学认识。跳虫功能群的分类标准不一导致相关研究结果可比性差。本文整理了跳虫功能群的3种分类方法,归纳了4种研究跳虫营养关系的方法,总结了N、P养分添加下土壤跳虫的响应机制。总体上,跳虫对N添加的响应多表现为负效应即种群密度降低,而对P添加的响应则多表现为正效应,N、P混施下,跳虫群落的变化更为复杂。未来土壤跳虫对N、P添加响应的研究应聚焦在跳虫功能群的科学划分、跳虫营养结构研究方法的完善和N、P交互对跳虫群落的作用机制上。     

华北盐渍化草地土壤净氮矿化速率对不同水平氮添加的响应

对于养分贫瘠的盐渍化草地生态系统, 大气氮沉降如何影响土壤氮循环过程是一个目前尚未解决的问题。该研究在位于华北地区山西省右玉县境内的盐渍化草地建立了一个模拟氮沉降的试验平台, 设置8个氮添加水平, 分别为0、1、2、4、8、16、24、32 g·m^-2·a^-1 (N0、N1、N2、N4、N8、N16、N24、N32), 生长季5-9月, 每月月初以喷施的方式等量添加NH₄NO₃。从2017年5月到2019年10月, 运用顶盖PVC管法每月一次进行净氮矿化速率的测定同时计算了净氮矿化速率对不同水平氮添加的敏感性。主要结果表明: (1)高水平氮添加(N16、N24、N32)显著增加土壤无机氮库; (2)该盐渍化草地土壤氮矿化以硝化作用为主, 经过3年氮添加以后, 高氮添加(N24、N32)显著促进了土壤净硝化速率, 并且不同氮添加水平在不同的月份和年份中表现出差异性响应; (3)不同氮添加水平对土壤净氮矿化敏感性的影响在不同降水年份差异显著, 短期低水平氮添加提高了土壤净氮矿化的敏感性, 而高水平氮添加降低土壤净氮矿化敏感性; (4)盐渍化草地土壤净氮矿化速率与土壤温度和水分呈正相关关系, 与土壤pH呈负相关关系。因此, 在当前氮沉降增加的背景下, 北方盐渍化草地土壤氮矿化速率对低氮添加的敏感性较高, 结合氮沉降的特点, 未来模型预测应该同时考虑氮沉降对盐渍化草地的可能影响。     

Responses of the functional structure of soil microbial community to livestock grazing in the Tibetan alpine grassland

Microbes play key roles in various biogeochemical processes, including carbon (C) and nitrogen (N) cycling. However, changes of microbial community at the functional gene level by livestock grazing, which is a global land‐use activity, remain unclear. Here we use a functional gene array, GeoChip 4.0, to examine the effects of free livestock grazing on the microbial community at an experimental site of Tibet, a region known to be very sensitive to anthropogenic perturbation and global warming. Our results showed that grazing changed microbial community functional structure, in addition to aboveground vegetation and soil geochemical properties. Further statistical tests showed that microbial community functional structures were closely correlated with environmental variables, and variations in microbial community functional structures were mainly controlled by aboveground vegetation, soil C/N ratio, and NH₄⁺‐N. In‐depth examination of N cycling genes showed that abundances of N mineralization and nitrification genes were increased at grazed sites, but denitrification and N‐reduction genes were decreased, suggesting that functional potentials of relevant bioprocesses were changed. Meanwhile, abundances of genes involved in methane cycling, C fixation, and degradation were decreased, which might be caused by vegetation removal and hence decrease in litter accumulation at grazed sites. In contrast, abundances of virulence, stress, and antibiotics resistance genes were increased because of the presence of livestock. In conclusion, these results indicated that soil microbial community functional structure was very sensitive to the impact of livestock grazing and revealed microbial functional potentials in regulating soil N and C cycling, supporting the necessity to include microbial components in evaluating the consequence of land‐use and/or climate changes.     

Disturbance is more important than seeding or grazing in determining soil microbial communities in a semiarid grassland

A primary goal of ecological restoration is often to return processes and functions to degraded ecosystems. Soil, while often ignored in restoration, supports diverse communities of organisms and is a fundamental actor in providing ecosystem processes and services. We investigated the impact of seeding and livestock grazing on plant communities, soil microorganisms, and soil fertility 3 years after the restoration of a disturbed pipeline corridor in southeastern Arizona. The initial soil disturbance and topsoil treatment, regardless of seeding or grazing, was the most influential factor in determining differences in both plant and microbial communities. Compared with the control, the disturbed and restored sites had greater plant species richness, greater total herbaceous plant cover, greater soil organic matter, higher pH, and differed in soil nutrients. Bacteria and fungi appeared to generally correlate with micro‐environment and soil physiochemical properties rather than specific plant species. The undisturbed control had a smaller proportion of bacterial functional groups associated with the breakdown of plant biomass (polysaccharide decomposition) and a smaller proportion of arbuscular mycorrhizal fungi (AMF) compared with disturbed and restored sites. The ability of the unseeded disturbed site to recover robust vegetation may be due in part to the high presence of AMF. These differences show selection for soil microorganisms that thrive in disturbed and restored sites and may contribute to increased plant productivity. Restoration of specific plant species or ecological processes and services would both benefit from better understanding of the impacts of disturbance on soil microorganisms and soil fertility.     

Response of bacterial communities and plant-mediated soil processes to nitrogen deposition and precipitation in a desert steppe

Plant and Soil - Changes in nitrogen (N) and precipitation levels can substantially alter soil properties and plant growth, thereby altering soil microbial diversity and functionality. We used...     

森林土壤微生物对干旱和氮沉降的响应

干旱和氮沉降深刻影响着人类世森林生态系统的生命活动与物质循环,进而影响全球碳平衡、并反馈作用于气候变化。土壤微生物驱动元素的生物地球化学循环和关键土壤生态过程,在气候变化生物学研究方面具有核心地位和全球重要性。本文综述了干旱和氮沉降对森林土壤细菌和菌根真菌的影响。提出未来应加强全球变化多因子交互作用对土壤微生物多样性、活性与生态功能的研究;建立野外长期定位站,强化亚热带森林生态系统与全球变化研究;注重土壤生物之间互作及网络研究;利用微生物大数据建立相关的机理模型等。从认识微生物多样性和群落组成对全球变化的响应与适应,逐步发展为调控利用微生物群落服务于森林的优化管理、生态资源的合理保护与可持续利用,为充分发挥微生物减缓全球气候变化的作用提供理论基础。     

Summer irrigation,grazing and seed addition differentially influence community composition in an invaded serpentine grassland

Community assembly theory predicts that resource availability, biotic interactions, and dispersal dynamics will determine community composition. Recent work has demonstrated that manipulating these processes or “filters” to exclude exotic species may assist in restoring invaded plant communities. In this study, we began by manipulating an abiotic filter, summer water availability, on the theory that irrigation prior to the growing season could trigger the germination of exotic species during unfavorable environmental conditions. First, we performed a greenhouse experiment to assess the germination traits of 23 native and exotic species at low (16°C, spring) and high (30°C, summer) temperatures. At summer temperatures, we found high emergence of many exotic and native grasses and low emergence of native forbs suggesting that summer irrigation may help deplete the exotic seed bank. In a second experiment, we established field plots to test the efficacy of summer irrigation and simultaneously manipulated a biotic and a dispersal filter, subjecting some plots to grazing and/or native seed addition. Summer irrigation and seed addition had no effect on percent cover or species richness while grazing reduced native cover but increased native species richness and soil nitrogen content. Our data suggest that manipulating grazing (a biotic filter) may be more effective than altering abiotic or dispersal filters when restoring invaded serpentine grassland. However, summer irrigation may also be effective, if applied at lower temperatures or for longer periods.