生态系统演替过程中土壤与微生物碳氮磷化学计量关系的变化

土壤碳(C)、氮(N)、磷(P)化学计量特征会显著影响微生物的生长、群落结构、生物量C:N:P化学计量及其代谢活动。然而生态系统演替过程中土壤-微生物C:N:P化学计量的时间格局及其协调关系还不明确。为此, 该研究收集了2016年5月以前发表的文献中19个生态系统演替序列(包括13个森林、6个草地生态系统)的土壤-微生物生物量C:N:P研究结果, 整合分析了其中土壤-微生物生态化学计量的时间动态, 结果表明: (1)生态系统演替过程中土壤C:N没有一致的时间格局, 而土壤C:P和N:P均随演替进程显著增加, 其中土壤C:N:P与演替时间之间线性关系的斜率与相应演替序列的初始土壤有机C含量呈负相关关系。(2)演替进程中土壤-微生物生物量C:N:P没有一致的时间格局。(3)微生物生物量C占土壤有机C百分比(qMBC)、微生物生物量N占土壤全N百分比、微生物生物量P占土壤全P百分比均随着演替进程而显著增加, 即单位资源所能支持的微生物生物量随着演替进程而增加, 这与宏观生态系统演替理论相符。(4) qMBC随着土壤C:N、C:P和N:P以及C:N、C:P和N:P化学计量不平衡性(即土壤C:N、C:P和N:P分别除以微生物生物量C:N、C:P和N:P)的增加而减小; 其中, C:N、C:P和N:P化学计量不平衡性解释了qMBC变异性的37%-57%, 是演替时间解释率的7-17倍, 表明土壤-微生物生态化学计量关系对qMBC演替动态有重要影响。该研究强调了生态化学计量学理论和生态系统演替理论在土壤微生物时间动态研究中的重要作用, 表明适当地融合生态学宏观理论于土壤微生物研究可以加深对土壤-微生物生态过程的认识。     

土壤微生物对气候变暖和大气N沉降的响应

气候变暖和大气N沉降是近一、二十年来人们非常关注的全球变化现象,它们所带来的一系列生态问题已成为全球变化研究的重要议题。它们不仅影响地上植被生长和群落组成,还直接或间接地影响土壤微生物过程,而土壤微生物对此做出的响应正是生态系统反馈过程中非常重要的环节。该文分别从气候变化对土壤微生物的影响(土壤微生物量、微生物活动和微生物群落结构)和土壤微生物对气候变化的响应(凋落物分解、养分利用与循环以及养分的固持与流失)两个角度,综述近期土壤微生物对气候变暖和大气N沉降响应与适应的研究进展。气候变暖和大气N沉降对土壤微生物的影响更多地反映在微生物群落的结构和功能上,而土壤微生物量、微生物活动和群落结构的变化又会通过改变凋落物分解、养分利用和C、N循环等重要的土壤生态系统功能和过程做出响应,形成正向或负向反馈,加强或削弱气候变化给整个陆地生态系统带来的影响。然而,到目前为止土壤微生物的响应对陆地生态系统产生的最终结果仍是未决的关键性问题。     

氮添加对荒漠草原植物群落组成与微生物生物量生态化学计量特征的影响

大气氮(N)沉降增加加速了土壤N循环, 引起微生物生物量碳(C):N:磷(P)生态化学计量关系失衡、植物种丧失和生态系统服务功能降低等问题。开展N添加下植物群落组成与微生物生物量生态化学计量特征关系的研究, 可为深入了解N沉降增加引起植物多样性降低的机理提供新思路。该文以宁夏荒漠草原为研究对象, 探讨了N添加下植物生物量和群落多样性的变化趋势, 分析了微生物生物量C:N:P生态化学计量特征独立及其与其他土壤因子共同对植物群落组成的影响。结果表明: N添加下猪毛菜(Salsola collina)生物量呈显著增加趋势, 牛枝子(Lespedeza potaninii)生物量呈逐渐降低趋势, 其他植物种生物量亦呈降低趋势但未达到显著水平; 沿N添加梯度, Shannon-Wiener多样性指数、Simpson优势度指数和Patrick丰富度指数均呈先略有增加后逐渐降低的趋势; N添加提高了微生物生物量N含量和N:P, 降低了微生物生物量C:N; 植物群落组成与微生物生物量N含量、微生物生物量C:N、微生物生物量N:P、土壤NO₃ ^--N浓度、土壤NH₄ ⁺-N浓度以及土壤全P含量有较强的相关关系; 微生物生物量C:N:P生态化学计量特征对植物种群生物量和群落多样性变化的独立解释力较弱, 但却与其他土壤因子共同解释了较大变差, 意味着N添加下微生物生物量C:N:P生态化学计量特征对植物群落组成的影响与其他土壤因子高度相关。     

氮添加对荒漠草原植物群落组成与微生物生物量生态化学计量特征的影响

大气氮(N)沉降增加加速了土壤N循环,引起微生物生物量碳(C):N:磷(P)生态化学计量关系失衡、植物种丧失和生态系统服务功能降低等问题。开展N添加下植物群落组成与微生物生物量生态化学计量特征关系的研究,可为深入了解N沉降增加引起植物多样性降低的机理提供新思路。该文以宁夏荒漠草原为研究对象,探讨了N添加下植物生物量和群落多样性的变化趋势,分析了微生物生物量C:N:P生态化学计量特征独立及其与其他土壤因子共同对植物群落组成的影响。结果表明:N添加下猪毛菜(Salsolacollina)生物量呈显著增加趋势,牛枝子(Lespedezapotaninii)生物量呈逐渐降低趋势,其他植物种生物量亦呈降低趋势但未达到显著水平;沿N添加梯度,Shannon-Wiener多样性指数、Simpson优势度指数和Patrick丰富度指数均呈先略有增加后逐渐降低的趋势;N添加提高了微生物生物量N含量和N:P,降低了微生物生物量C:N;植物群落组成与微生物生物量N含量、微生物生物量C:N、微生物生物量N:P、土壤NO3--N浓度、土壤NH4+-N浓度以及土壤全P含量有较强的相关关系;微生物生物量C:N:P生态化学计量特征对植物种群生物量和群落多样性变化的独立解释力较弱,但却与其他土壤因子共同解释了较大变差,意味着N添加下微生物生物量C:N:P生态化学计量特征对植物群落组成的影响与其他土壤因子高度相关。     

冰川前缘土壤微生物原生演替的生态特征——以乌鲁木齐河源1号冰川为例

为了解冰川前缘土壤微生物在原生演替过程中的生态特征及其影响因素,用空间距离代替时间序列,以乌鲁木齐河源1号冰川终碛堤为起点,沿6个不同演替时期(0,4,15,31,43a和对照)的样带采集土样,以冰川附近发育良好的土壤为对照,测定土壤酶活性、微生物氮矿化与脱氨作用以及微生物生物量。结果表明,土壤脲酶、蛋白酶、酸性磷酸酶、芳基硫酸酯酶、蔗糖酶活性、微生物氮矿化及脱氨作用随演替时间而增加,微生物生物量碳和氮变化呈波动状,趋势不明显。相关分析表明,土壤有机质与酶活、微生物生物量存在极显著正相关(P0.01)。1号冰川前缘微生物多样性指数随着演替时间持续增加,但目前仍未达稳定状态。     

土壤微生物生物量碳氮磷与土壤酶化学计量对气候变化的响应机制

微生物和土壤酶是陆地生态系统中生物地球化学循环的重要驱动力,深入理解微生物在生态系统中的调节作用以及气候变化过程中微生物量和土壤酶的响应机制是生态学领域关注的重要科学问题.本研究从气候因素角度出发,基于生态化学计量学理论,综述了微生物和土壤酶在陆地生态系统碳氮磷循环中的作用,以及土壤微生物生物量碳氮磷和土壤酶化学计量对气候变化的响应机制,即: 改变微生物代谢速率和酶活性;调整微生物群落结构;调整微生物生物量碳氮磷与土壤酶化学计量特征;改变碳氮磷养分元素利用效率.最后分析当前研究的不足,并提出了该领域亟待解决的科学问题: 综合阐明土壤微生物和土壤酶对气候变化的响应机制;探究土壤微生物和胞外酶养分耦合机理;深入探究土壤微生物量和土壤酶化学计量特征对气候变化的适应对策.     

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。     

长白山温带森林不同演替阶段土壤化学性质及微生物群落结构的变化

土壤微生物是生态系统物质循环和能量流动的驱动者,其群落结构可以用来表征土壤生态过程及其对地上植被变化的响应机制。本研究采用时空替代法在长白山地区选取了阔叶红松林演替序列的5个不同阶段:杨桦幼龄林、杨桦中龄林、杨桦成熟林、阔叶红松成熟林和阔叶红松过熟林,采用磷脂脂肪酸法(PLFA)测定了土壤微生物群落组成和结构,分析了其随地上植被演替过程的变化,同时比较了不同演替阶段土壤化学性质差异。结果表明:随着演替的正向进行,土壤总有机碳、全碳、全氮、全磷含量显著提高,碳氮比逐渐下降。土壤微生物生物量、群落结构及组成发生明显变化:土壤微生物总PLFAs、细菌PLFAs、革兰氏阳性菌PLFAs、革兰氏阴性菌PLFAs含量显著增加;真菌PLFAs(18:2ω6c)先增加后减少,中期阶段的杨桦成熟林土壤真菌PLFA含量最高,同时细菌/真菌最小;革兰氏阳性菌/革兰氏阴性菌(G+/G-)随着演替的进行逐渐增大。土壤微生物生物量与土壤全碳、总有机碳、全氮、全磷含量呈显著正相关,与碳氮比呈显著负相关;冗余分析(RDA)结果显示,全碳、总有机碳、全氮和碳氮比是影响土壤微生物群落结构的主要因素。本研究表明,随着植被演替的正向进行,土壤质量逐渐提高;土壤微生物群落组成明显改变;土壤微生物群落结构与土壤理化性质显著相关。     

森林次生演替和土壤层次对微生物群落结构的影响

森林次生演替与生态系统结构和功能的动态变化密切相关。大多数研究主要关注植物群落以及土壤有机碳(SOC)的变化,然而土壤微生物群落如何响应森林次生演替还需要进一步探究。本研究以长白山森林次生演替序列(20、80、120、200和≥300年)和两个土壤层次为对象,采用磷脂脂肪酸微生物标志物,探究温带森林次生演替过程中地下微生物群落结构变化。森林次生演替改变了土壤微生物群落结构,主要归因于某些特定微生物类群的变化,演替前期革兰氏阴性菌和腐生真菌占主导,而在演替后期革兰氏阳性菌和丛枝菌根真菌占主导。另外,土壤有机质数量和质量差异是影响微生物群落结构和生物量的主要环境因素。森林演替前期和中期增加的SOC含量促进了微生物生物量,而演替后期增加的难分解芳香族有机组分抑制了微生物生物量合成。土壤层次间理化性质的差异导致微生物群落变化,有机质层高的SOC以及氮含量导致更多微生物生物量的合成。微生物群落在时间和空间尺度的变化及其驱动因素反映了生态系统结构和功能对环境变化的响应。     

森林次生演替和土壤层次对微生物群落结构的影响

森林次生演替与生态系统结构和功能的动态变化密切相关。大多数研究主要关注植物群落以及土壤有机碳(SOC)的变化,然而土壤微生物群落如何响应森林次生演替还需要进一步探究。本研究以长白山森林次生演替序列(20、80、120、200和≥300年)和两个土壤层次为对象,采用磷脂脂肪酸微生物标志物,探究温带森林次生演替过程中地下微生物群落结构变化。森林次生演替改变了土壤微生物群落结构,主要归因于某些特定微生物类群的变化,演替前期革兰氏阴性菌和腐生真菌占主导,而在演替后期革兰氏阳性菌和丛枝菌根真菌占主导。另外,土壤有机质数量和质量差异是影响微生物群落结构和生物量的主要环境因素。森林演替前期和中期增加的SOC含量促进了微生物生物量,而演替后期增加的难分解芳香族有机组分抑制了微生物生物量合成。土壤层次间理化性质的差异导致微生物群落变化,有机质层高的SOC以及氮含量导致更多微生物生物量的合成。微生物群落在时间和空间尺度的变化及其驱动因素反映了生态系统结构和功能对环境变化的响应。     

Changes of the relationships between soil and microbes in carbon,nitrogen and phosphorus stoichiometry during ecosystem succession

AimsThe carbon (C), nitrogen (N) and phosphorus (P) stoichiometry (C:N:P) of soil profoundly influences the growth, community structure, biomass C:N:P stoichiometry, and metabolism in microbes. However, the relationships between soil and microbes in the C:N:P stoichiometry and their temporal dynamics during ecosystem succession are poorly understood. The aim of this study was to determine the temporal patterns of soil and microbial C:N:P stoichiometry and their relationships during ecosystem succession.MethodsAn extensive literature search was conducted and data were compiled for 19 age sequences of successional ecosystems, including 13 forest ecosystems and 6 grassland ecosystems, from 18 studies published up to May 2016. Meta-analyses were performed to examine the sequential changes in 18 variables that were associated with soil and microbial C, N and P contents and the stoichiometry. Important findings (1) There was no consistent temporal pattern in soil C:N along the successional stages, whereas the soil C:P and N:P increased with succession; the slopes of the linear relationships between soil C:N:P stoichiometry and successional age were negatively correlated with the initial content of the soil organic C within given chronosequence. (2) There was no consistent temporal pattern in microbial C:N:P stoichiometry along the successional stages. (3) The fraction of microbial biomass C in soil organic C (qMBC), the fraction of microbial biomass N in soil total N, and the fraction of microbial biomass P in soil total P all increased significantly with succession, in consistency with the theory of succession that ecosystem biomass per unit resource increases with succession. (4) The qMBC decreased with increases in the values of soil C:N, C:P, or N:P, as well as the stoichiometric imbalances in C:N, C:P, and N:P between soil and microbes (i.e., ratios of soil C:N, C:P, and N:P to microbial biomass C:N, C:P, and N:P, respectively). The C:N, C:P, and N:P stoichiometric imbalances explained 37%-57% variations in the qMBC, about 7-17 times more than that explainable by the successional age, illustrating the importance of soil-microbial C:N:P stoichiometry in shaping the successional dynamics in qMBC. In summary, our study highlights the importance of the theories of ecosystem succession and stoichiometry in soil microbial studies, and suggests that appropriately applying macro-ecological theories in microbial studies may improve our understanding on microbial ecological processes.     

Heterotrophic microbial communities use ancient carbon following glacial retreat

When glaciers retreat they expose barren substrates that become colonized by organisms, beginning the process of primary succession. Recent studies reveal that heterotrophic microbial communities occur in newly exposed glacial substrates before autotrophic succession begins. This raises questions about how heterotrophic microbial communities function in the absence of carbon inputs from autotrophs. We measured patterns of soil organic matter development and changes in microbial community composition and carbon use along a 150-year chronosequence of a retreating glacier in the Austrian Alps. We found that soil microbial communities of recently deglaciated terrain differed markedly from those of later successional stages, being of lower biomass and higher abundance of bacteria relative to fungi. Moreover, we found that these initial microbial communities used ancient and recalcitrant carbon as an energy source, along with modern carbon. Only after more than 50 years of organic matter accumulation did the soil microbial community change to one supported primarily by modern carbon, most likely from recent plant production. Our findings suggest the existence of an initial stage of heterotrophic microbial community development that precedes autotrophic community assembly and is sustained, in part, by ancient carbon.     

Do bacterial and fungal communities assemble differently during primary succession?

High‐throughput sequencing technologies are now allowing us to study patterns of community assembly for diverse microbial assemblages across environmental gradients and during succession. Here we discuss potential explanations for similarities and differences in bacterial and fungal community assembly patterns along a soil chronosequence in the foreland of a receding glacier. Although the data are not entirely conclusive, they do indicate that successional trajectories for bacteria and fungi may be quite different. Recent empirical and theoretical studies indicate that smaller microbes (like most bacteria) are less likely to be dispersal limited than are larger microbes – which could result in a more deterministic community assembly pattern for bacteria during primary succession. Many bacteria are also better adapted (than are fungi) to life in barren, early‐successional sediments in that some can fix nitrogen and carbon from the atmosphere – traits not possessed by any fungi. Other differences between bacteria and fungi are discussed, but it is apparent from this and other recent studies of microbial succession that we are a long way from understanding the mechanistic underpinnings of microbial community assembly during ecosystem succession. We especially need a better understanding of global and regional patterns of microbial dispersal and what environmental factors control the development of microbial communities in complex natural systems.     

Microbial dynamics in soils of the Damma glacier forefield show succession in the functional genetic potential

Glacier retreat is a visible consequence of climate change worldwide. Although taxonomic change of the soil microbiomes in glacier forefields have been widely documented, how microbial genetic potential changes along succession is little known. Here, we used shotgun metagenomics to analyse whether the soil microbial genetic potential differed between four stages of soil development (SSD) sampled along three transects in the Damma glacier forefield (Switzerland). The SSDs were characterized by an increasing vegetation cover, from barren soil, to biological soil crust, to sparsely vegetated soil and finally to vegetated soil. Results suggested that SSD significantly influenced microbial genetic potential, with the lowest functional diversity surprisingly occurring in the vegetated soils. Overall, carbohydrate metabolism and secondary metabolite biosynthesis genes overrepresented in vegetated soils, which could be partly attributed to plant–soil feedbacks. For C degradation, glycoside hydrolase genes enriched in vegetated soils, while auxiliary activity and carbohydrate esterases genes overrepresented in barren soils, suggested high labile C degradation potential in vegetated, and high recalcitrant C degradation potential in barren soils. For N-cycling, organic N degradation and synthesis genes dominated along succession, and gene families involved in nitrification were overrepresented in barren soils. Our study provides new insights into how the microbial genetic potential changes during soil formation along the Damma glacier forefield.     

Differential sensitivity of total and active soil microbial communities to drought and forest management

Climate change will affect semiarid ecosystems through severe droughts that increase the competition for resources in plant and microbial communities. In these habitats, adaptations to climate change may consist of thinning—that reduces competition for resources through a decrease in tree density and the promotion of plant survival. We deciphered the functional and phylogenetic responses of the microbial community to 60 years of drought induced by rainfall exclusion and how forest management affects its resistance to drought, in a semiarid forest ecosystem dominated by Pinus halepensis Mill. A multiOMIC approach was applied to reveal novel, community‐based strategies in the face of climate change. The diversity and the composition of the total and active soil microbiome were evaluated by 16S rRNA gene (bacteria) and ITS (fungal) sequencing, and by metaproteomics. The microbial biomass was analyzed by phospholipid fatty acids (PLFAs), and the microbially mediated ecosystem multifunctionality was studied by the integration of soil enzyme activities related to the cycles of C, N, and P. The microbial biomass and ecosystem multifunctionality decreased in drought‐plots, as a consequence of the lower soil moisture and poorer plant development, but this decrease was more notable in unthinned plots. The structure and diversity of the total bacterial community was unaffected by drought at phylum and order level, but did so at genus level, and was influenced by seasonality. However, the total fungal community and the active microbial community were more sensitive to drought and were related to ecosystem multifunctionality. Thinning in plots without drought increased the active diversity while the total diversity was not affected. Thinning promoted the resistance of ecosystem multifunctionality to drought through changes in the active microbial community. The integration of total and active microbiome analyses avoids misinterpretations of the links between the soil microbial community and climate change.     

贡嘎山冰川退缩区植物-土壤反馈与植物间相互作用

植物-土壤反馈是揭示陆地生物群落动态变化的关键环节,为理解植物间相互作用及植被群落变化过程奠定基础。本研究以贡嘎山冰川退缩区原生演替早(5~10年)、中(30~40年)和晚期(80~100年)3个阶段典型土壤以及各阶段优势植物为对象,采用盆栽控制试验,比较优势植物在不同土壤条件下的生物量,并量化植物间相互作用以及植物-土壤反馈的方向与强度,为探究贡嘎山冰川退缩区植被群落演替规律提供依据。结果表明:(1)植物-土壤反馈作用显著影响植物在本土中的生物量,早期沙棘(Hippophae rhamnoides)在本土中生长最差,沙棘的植物-土壤反馈系数为负值;演替中期冬瓜杨(Populus purdomii)的反馈系数趋于零;晚期峨眉冷杉(Abies fabri)在本土中生长最好,峨眉冷杉的反馈系数为正值。(2)混种时,早期沙棘与演替中、晚期植物间相互作用指数为负值;中期冬瓜杨、川滇柳(Salix rehderiana)与演替早、晚期植物的相互作用指数接近于零,晚期植物峨眉冷杉、麦吊云杉(Picea brachytyla)与演替早、中期植物相互作用指数为正值。从植物-土壤反馈的方向来看,贡嘎山植被演替从早期负反馈,中期中性反馈,过渡到晚期正反馈。此外,演替早期沙棘促进演替中晚期植物生长,演替中期冬瓜杨、川滇柳对演替早晚期植物无显著影响,晚期峨眉冷杉、麦吊云杉更利于与演替早中期植物相互竞争。结果显示,植物-土壤反馈与植物间相互作用共同驱动了贡嘎山冰川退缩区植被快速演替,直至顶极群落。