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。     

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.     

全球森林土壤微生物生物量碳氮磷化学计量的季节动态

土壤微生物生物量在森林生态系统中充当具有生物活性的养分积累和储存库。土壤微生物转化有机质为植物提供可利用养分, 与植物的相互作用维系着陆地生态系统的生态功能。同时, 土壤微生物也与植物争夺营养元素, 在季节交替过程和植物的生长周期中呈现出复杂的互利-竞争关系。综合全球数据对温带、亚热带和热带森林土壤微生物生物量碳(C)、氮(N)、磷(P)含量及其化学计量比值的季节动态进行分析, 发现温带和亚热带森林的土壤微生物生物量C、N、P含量均呈现夏季低、冬季高的格局。热带森林四季的土壤微生物生物量C、N、P含量都低于温带和亚热带森林, 且热带森林土壤微生物生物量C含量、N含量在秋季相对最低, 土壤微生物生物量P含量四季都相对恒定。温带森林的土壤微生物生物量C:N在春季显著高于其他两个森林类型; 热带森林的土壤微生物生物量C:N在秋季显著高于其他2个森林类型。温带森林土壤微生物生物量N:P和C:P在四季都保持相对恒定, 而热带森林土壤微生物生物量N:P和C:P在夏季高于其他3个季节。阔叶树的土壤微生物生物量C含量、N含量、N:P、C:P在四季都显著高于针叶树; 而针叶树的土壤微生物生物量P含量在四季都显著高于阔叶树。在春季和冬季时, 土壤微生物生物量C:N在阔叶树和针叶树之间都没有显著差异; 但是在夏季和秋季, 针叶树的土壤微生物生物量C:N显著高于阔叶树。对于土壤微生物生物量的变化来说, 森林类型是主要的显著影响因子, 季节不是显著影响因子, 暗示土壤微生物生物量的季节波动是随着植物其内在固有的周期变化而变化。植物和土壤微生物密切作用表现出来的对养分的不同步吸收是保留养分和维持生态功能的一种权衡机制。     

喀斯特石漠化地区不同退耕年限下桂牧1号杂交象草植物-土壤-微生物生态化学计量特征

耕地农作物种植与退耕地种草养畜是喀斯特主要的农业生产模式。以喀斯特地区农耕玉米地为对照,研究退耕还草1、5、7a(恢复初期、旺盛期、衰退期)3种年限下桂牧1号杂交象草地植物-土壤-微生物C、N、P生态化学计量特征及内在关联。结果表明:1)牧草地植物地上部分N、P含量均为5a牧草1a牧草7a牧草,C含量则刚好相反;3种退耕年限牧草地植物地上部分C∶N、C∶P、N∶P分别为26.50—33.91、631.70—2254.33、23.89—67.21,且均表现为7a牧草1a牧草5a牧草。2)土壤表层(0—10 cm)C、N、P含量均以玉米地最低,3种退耕年限牧草地中则均为5a牧草地最低;土壤C∶N、C∶P、N∶P在玉米及退耕牧草地之间均无显著差异(P0.05),平均值分别为9.20,27.88,3.38。3)玉米及牧草地土壤MB_C、MB_N、MB_P含量存在显著差异(P0.05);玉米地MB_C/SOC、MB_N/TN、MB_P/TP均高于牧草地,3种退耕年限牧草地中,则均为5a牧草地最高。4)MB_C、MB_P与土壤C、P含量分别呈显著线性正相关(P0.05);植物C、C∶N与土壤C、N含量均呈极显著线性负相关(P0.01)。分析表明,退耕还草地中植物与土壤系统C-N-P化学计量比表现出不一致的时间变化特征,且牧草地植物受P限制严重,尤以恢复旺盛期为甚。     

C:N:P stoichiometry in soil: is there a “Redfield ratio” for the microbial biomass?

Well-constrained carbon:nitrogen:phosphorus (C:N:P) ratios in planktonic biomass, and their importance in advancing our understanding of biological processes and nutrient cycling in marine ecosystems, has motivated ecologists to search for similar patterns in terrestrial ecosystems. Recent analyses indicate the existence of “Redfield-like” ratios in plants, and such data may provide insight into the nature of nutrient limitation in terrestrial ecosystems. We searched for analogous patterns in the soil and the soil microbial biomass by conducting a review of the literature. Although soil is characterized by high biological diversity, structural complexity and spatial heterogeneity, we found remarkably consistent C:N:P ratios in both total soil pools and the soil microbial biomass. Our analysis indicates that, similar to marine phytoplankton, element concentrations of individual phylogenetic groups within the soil microbial community may vary, but on average, atomic C:N:P ratios in both the soil (186:13:1) and the soil microbial biomass (60:7:1) are well-constrained at the global scale. We did see significant variation in soil and microbial element ratios between vegetation types (i.e., forest versus grassland), but in most cases, the similarities in soil and microbial element ratios among sites and across large scales were more apparent than the differences. Consistent microbial biomass element ratios, combined with data linking specific patterns of microbial element stoichiometry with direct evidence of microbial nutrient limitation, suggest that measuring the proportions of C, N and P in the microbial biomass may represent another useful tool for assessing nutrient limitation of ecosystem processes in terrestrial ecosystems.     

Organic matter inputs shift soil enzyme activity and allocation patterns in a wet tropical forest

Soil extracellular enzymes mediate organic matter turnover and nutrient cycling yet remain little studied in one of Earth’s most rapidly changing, productive biomes: tropical forests. Using a long-term leaf litter and throughfall manipulation, we explored relationships between organic matter (OM) inputs, soil chemical properties and enzyme activities in a lowland tropical forest. We assayed six hydrolytic soil enzymes responsible for liberating carbon (C), nitrogen (N) and phosphorus (P), calculated enzyme activities and ratios in control plots versus treatments, and related these to soil biogeochemical variables. While leaf litter addition and removal tended to increase and decrease enzyme activities per gram soil, respectively, shifts in enzyme allocation patterns implied changes in relative nutrient constraints with altered OM inputs. Enzyme activity ratios in control plots suggested strong belowground P constraints; this was exacerbated when litter inputs were curtailed. Conversely, with double litter inputs, increased enzymatic investment in N acquisition indicated elevated N demand. Across all treatments, total soil C correlated more strongly with enzyme activities than soluble C fluxes, and enzyme ratios were sensitive to resource stoichiometry (soil C:N) and N availability (net N mineralization). Despite high annual precipitation in this site (MAP ~5 m), soil moisture positively correlated with five of six enzymes. Our results suggest resource availability regulates tropical soil enzyme activities, soil moisture plays an additional role even in very wet forests, and relative investment in C, N and P degrading enzymes in tropical soils will often be distinct from higher latitude ecosystems yet is sensitive to OM inputs.     

Ecoenzymatic stoichiometry of microbial nutrient acquisition in tropical soils

The relative activities of soil enzymes involved in mineralizing organic carbon (C), nitrogen (N), and phosphorus (P) reveal stoichiometric and energetic constraints on microbial biomass growth. Although tropical forests and grasslands are a major component of the global C cycle, the effects of soil nutrient availability on microbial activity and C dynamics in these ecosystems are poorly understood. To explore potential microbial nutrient limitation in relation to enzyme allocation in low latitude ecosystems, we performed a meta-analysis of acid/alkaline phosphatase (AP), β-1,4-glucosidase (BG), and β-1,4-N-acetyl-glucosaminidase (NAG) activities in tropical soils. We found that BG:AP and NAG:AP ratios in tropical soils are significantly lower than those of temperate ecosystems overall. The lowest BG:AP and NAG:AP ratios were associated with old or acid soils, consistent with greater biological phosphorus demand relative to P availability. Additionally, correlations between enzyme activities and mean annual temperature and precipitation suggest some climatic regulation of microbial enzyme allocation in tropical soils. We used the results of our analysis in conjunction with previously published data on soil and biomass C:N:P stoichiometry to parameterize a biogeochemical equilibrium model that relates microbial growth efficiency to extracellular enzyme activity. The model predicts low microbial growth efficiencies in P-limited soils, indicating that P availability may influence C cycling in the highly weathered soils that underlie many tropical ecosystems. Therefore, we suggest that P availability be included in models that simulate microbial enzyme allocation, biomass growth, and C mineralization.     

Tropical forest and savanna ecosystems show differential impact of N and P additions on soil organic matter and aggregate structure

Landscape transformation and atmospheric nutrient depositions, important global change drivers, are affecting the vegetation and soil properties of natural dry tropical forest and derived savanna ecosystems in India. This study assessed the effect of continuous N and P additions for 6 years on the size distribution and properties of soil aggregates in forest–ecotone–savanna gradient. Addition of N significantly increased the proportion of macroaggregates in forest and ecotone, whereas the same input significantly decreased their proportion in the savanna. Consequently, the stability of soil aggregates increased significantly in forest and ecotone, whereas it decreased significantly in the savanna. The effect of P addition on soil aggregate stability was marginal. N addition also altered the biological and chemical qualities of soil aggregates. It caused increase in microbial biomass C (MBC) associated with macroaggregates in forest and ecotone; however, in savanna, MBC increased in the microaggregates. P addition did not affect the amount of MBC in both types of soil aggregates. Because of rapid accumulation of applied N and P in the microbial biomass, the ratios of MBC to microbial biomass nitrogen (MBN) as well as microbial biomass phosphorous (MBP) were decreased in both aggregates. Overall, the effect of N addition was more marked than that of P addition, suggesting that N is more limiting than P in these dry tropical ecosystems. In the current scenario of N loading, continued soil N loading in forest may lead to increased macroaggregates with associated MBC and MBN and greater aggregate stability. In contrast, the extensively distributed savannas may show the reverse trend leading to a decrease in soil fertility.     

Changes in soil-microbe-exoenzyme C:N:P stoichiometry along an altitudinal gradient in Mt. Datudingzi,Northeast China

海拔变化导致温度、水分、植被等条件的改变会显著影响土壤碳(C_soil)、氮(N_soil)、磷(P_soil)含量及其化学计量特征, 土壤微生物如何通过调整自身生物量和胞外酶化学计量特征进行适应仍不明确。为了研究海拔梯度变化对土壤微生物生物量和胞外酶活性的影响, 探索土壤-微生物-胞外酶C:N:P化学计量特征间的协变性, 该文以黑龙江省雪乡大秃顶子山800、1 100、1 600和1 700 m分布的典型生态系统(针阔混交林、针叶林、岳桦林和草地)为研究对象, 测定其C_soil、N_soil、P_soil含量, 微生物生物量C (C_mic)、N (N_mic)、P (P_mic)含量, 以及微生物获取C (β-1, 4-葡萄糖苷酶, BG), N (几丁质酶, NAG), P (酸性磷酸酶, AP)资源的相关胞外酶活性。结果表明: (1)海拔梯度变化对C_soil和C_mic含量没有显著影响; 不同海拔间土壤和微生物生物量N、P含量存在显著差异。(2) BG和NAG活性随着海拔的升高呈现显著降低趋势, 表明海拔升高导致的温度降低抑制了微生物的活性。(3)海拔对土壤C:N、微生物C:N:P以及胞外酶C:N:P均具有显著影响。胞外酶C:N:P随着微生物与土壤间C:N:P化学计量不平衡性(土壤C:N:P与微生物C:N:P的比值)的增加而逐渐降低。微生物可以通过调整自身生物量以及胞外酶C:N:P适应土壤化学计量特征的变异, 该结果支持了微生物的资源分配理论。     

The elemental stoichiometry of aquatic and terrestrial ecosystems and its relationships with organismic lifestyle and ecosystem structure and function: a review and perspectives

C, N and P are three of the most important elements used to build living beings, and their uptake from the environment is consequently essential for all organisms. We have reviewed the available studies on water, soils and organism elemental content ratios (stoichiometry) with the aim of identifying the general links between stoichiometry and the structure and function of organisms and ecosystems, in both aquatic and terrestrial contexts. Oceans have variable C:N:P ratios in coastal areas and a narrow range approximating the Redfield ratio in deep water and inner oceanic areas. Terrestrial ecosystems have a general trend towards an increase in soil and plant N:P ratios from cool and temperate to tropical ecosystems, but with great variation within each climatic area. The C:N:P content ratio (from now on C:N:P ratio) is more constrained in organisms than in the water and soil environments they inhabit. The capacity to adjust this ratio involves several mechanisms, from leaf re-absorption in plants to the control of excretion in animals. Several differences in C:N:P ratios are observed when comparing different taxa and ecosystems. For freshwater ecosystems, the growth rate hypothesis (GRH), which has consistent experimental support, states that low N:P supply determines trophic web structures by favoring organisms with a high growth rate. For terrestrial organisms, however, evidence not yet conclusive on the relevance of the GRH. Recent studies suggest that the N:P ratio could play a role, even in the evolution of the genomes of organisms. Further research is warranted to study the stoichiometry of different trophic levels under different C:N:P environment ratios in long-term ecosystem-scale studies. Other nutrients such as K or Fe should also be taken into account. Further assessment of the GRH requires more studies on the effects of C:N:P ratios on anabolic (growth), catabolic (respiration), storage and/or defensive allocation. Combining elemental stoichiometry with metabolomics and/or genomics should improve our understanding of the coupling of different levels of biological organization, from elemental composition to the structure and evolution of ecosystems, via cellular metabolism and nutrient cycling.     

Microbial carbon use efficiency in different ecosystems: A meta-analysis based on a biogeochemical equilibrium model

Soil microbial carbon use efficiency (CUE) is a crucial parameter that can be used to evaluate the partitioning of soil carbon (C) between microbial growth and respiration. However, general patterns of microbial CUE among terrestrial ecosystems (e.g., farmland, grassland, and forest) remain controversial. To address this knowledge gap, data from 41 study sites (n = 197 soil samples) including 58 farmlands, 95 forests, and 44 grasslands were collected and analyzed to estimate microbial CUEs using a biogeochemical equilibrium model. We also evaluated the metabolic limitations of microbial growth using an enzyme vector model and the drivers of CUE across different ecosystems. The CUEs obtained from soils of farmland, forest, and grassland ecosystems were significantly different with means of 0.39, 0.33, and 0.42, respectively, illustrating that grassland soils exhibited higher microbial C sequestration potentials (p < .05). Microbial metabolic limitations were also distinct in these ecosystems, and carbon limitation was dominant exhibiting strong negative effects on CUE. Exoenzyme stoichiometry played a greater role in impacting CUE values than soil elemental stoichiometry within each ecosystem. Specifically, soil exoenzymatic ratios of C:phosphorus (P) acquisition activities (EEA_C:P) and the exoenzymatic ratio of C:nitrogen (N) acquisition activities (EEA_C:N) imparted strong negative effects on soil microbial CUE in grassland and forest ecosystems, respectively. But in farmland soils, EEA_C:P exhibited greater positive effects, showing that resource constraints could regulate microbial resource allocation with discriminating patterns across terrestrial ecosystems. Furthermore, mean annual temperature (MAT) rather than mean annual precipitation (MAP) was a critical climate factor affecting CUE, and soil pH as a major factor remained positive to drive the changes in microbial CUE within ecosystems. This research illustrates a conceptual framework of microbial CUEs in terrestrial ecosystems and provides the theoretical evidence to improve soil microbial C sequestration capacity in response to global change.     

Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest

Global changes such as variations in plant net primary production are likely to drive shifts in leaf litterfall inputs to forest soils, but the effects of such changes on soil carbon (C) cycling and storage remain largely unknown, especially in C‐rich tropical forest ecosystems. We initiated a leaf litterfall manipulation experiment in a tropical rain forest in Costa Rica to test the sensitivity of surface soil C pools and fluxes to different litter inputs. After only 2 years of treatment, doubling litterfall inputs increased surface soil C concentrations by 31%, removing litter from the forest floor drove a 26% reduction over the same time period, and these changes in soil C concentrations were associated with variations in dissolved organic matter fluxes, fine root biomass, microbial biomass, soil moisture, and nutrient fluxes. However, the litter manipulations had only small effects on soil organic C (SOC) chemistry, suggesting that changes in C cycling, nutrient cycling, and microbial processes in response to litter manipulation reflect shifts in the quantity rather than quality of SOC. The manipulation also affected soil CO ₂ fluxes; the relative decline in CO ₂ production was greater in the litter removal plots (?22%) than the increase in the litter addition plots (+15%). Our analysis showed that variations in CO ₂ fluxes were strongly correlated with microbial biomass pools, soil C and nitrogen (N) pools, soil inorganic P fluxes, dissolved organic C fluxes, and fine root biomass. Together, our data suggest that shifts in leaf litter inputs in response to localized human disturbances and global environmental change could have rapid and important consequences for belowground C storage and fluxes in tropical rain forests, and highlight differences between tropical and temperate ecosystems, where belowground C cycling responses to changes in litterfall are generally slower and more subtle.     

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.     

N limitation increases along a temperate forest succession: evidences from leaf stoichiometry and nutrient resorption

温带森林演替加剧了氮限制：来自叶片化学计量和养分重吸收的证据 森林生产力和碳汇功能在很大程度上取决于土壤氮和磷的有效性。然而,迄今为止,养分限制随森林演替的时间变化仍存在争议。叶片化学计量和养分重吸收是预测植物生长养分限制的重要指标。基于此,本研究测定了温带森林4个演替阶段所有木本植物叶片和凋落叶中氮和磷的含量,并分析了演替过程中非生物因子和生物因子如何影响叶片化学计量和养分重吸收。研究结果表明,在个体尺度上,叶片氮磷含量在演替末期显著增加,而叶片氮磷比无显著变化;氮的重吸收效率随演替显著增加,然而磷的重吸收效率先增加后减少;氮重吸收效率与磷重吸收效率的比值仅在演替末期显著增加。此外,植物氮素循环对土壤养分的响应比磷素循环更弱。在群落尺度上,叶片氮磷含量随森林演替呈现先降低后升高的趋势,主要受香农-维纳多样性指数和物种丰富度的影响;叶片氮磷比随演替而显著变化,主要由胸径的群落加权平均值决定;氮的重吸收效率增加,主要受物种丰富度和胸径的影响,而磷的重吸收效率相对稳定。因此,氮重吸收效率与磷重吸收效率的比值显著增加,表明随着温带森林演替,氮限制加剧。这些结果可能反映了较高生物多样性群落中物种间对有限资源的激烈竞争,强调了生物因子在驱动森林生态系统养分循环中的重要性,为中国温带和北方森林可持续经营的施肥管理提供了参考。     

黄土高原子午岭林区典型树种叶片N、P再吸收特征

为揭示黄土高原子午岭林区不同演替阶段和植被类型主要树种养分再吸收特征,研究选取4种次生植被树种(白桦、山杨、辽东栎和油松)和2种人工植被树种(刺槐和侧柏),测定其成熟叶、凋落叶和林下土壤碳(C)、氮(N)、磷(P)含量,研究了叶片N、P再吸收率及其与养分指标的关系。结果表明:(1)不同树种叶片养分和林下土壤养分含量存在显著差异,土壤C、N含量和C∶N∶P计量比均表现为演替后期林地(辽东栎和油松)演替前期林地(山杨和白桦)人工林(侧柏和刺槐);(2)不同树种叶片N、P再吸收率分别为17.18%—43.34%和27.13%—58.12%,均表现为演替后期林地人工林演替前期林地,且P的再吸收率总体高于N的再吸收率;(3)不同树种叶片N、P再吸收率与叶片养分指标的关系强于土壤,与养分计量比的相关性大于养分含量的相关性。说明子午岭典型植被会通过叶片N、P再吸收来适应养分限制环境,尤其是演替后期植被再吸收能力更强,研究可为黄土高原植被恢复提供理论依据。     

黄土丘陵区植被与地形特征对土壤和土壤微生物生物量生态化学计量特征的影响

研究黄土丘陵区植被与地形特征对土壤和土壤微生物生物量生态化学计量特征影响有助于深入理解黄土丘陵区不同植被带下土壤和土壤微生物相互作用及养分循环规律.选择黄土丘陵区延河流域3个植被区(森林区、森林草原区、草原区)和5种地形部位(阴/阳沟坡、阴/阳梁峁坡、峁顶)的土壤作为研究对象,利用生态化学计量学理论研究植被和地形对土壤和土壤微生物生物量生态化学计量特征的影响.结果表明: 土壤及土壤微生物生物量碳、氮、磷含量在不同地形之间的差别主要表现在沟坡位置和阴坡高于其他坡位和阳坡.植被类型的变化对两个土层(0～10、10～20 cm)土壤和土壤微生物生物量碳、氮、磷的影响均达到显著水平,坡向对表层(0～10 cm)土壤和土壤微生物生物量碳、氮、磷的影响强于坡位,而在10～20 cm土层,坡位对土壤和土壤微生物生物量碳、氮、磷影响更显著.植被类型显著影响土壤C∶N、C∶P、N∶P和土壤微生物生物量C∶N、C∶P,坡向和坡位仅影响土壤C∶P和N∶P,植被类型的变化是影响土壤C∶N的主要因素.同时,植被类型对土壤养分和微生物生物量碳、氮、磷含量及其生态化学计量特征的影响大于地形因子.标准化主轴分析结果表明,黄土丘陵区不同植被带土壤微生物具有内稳性,特别在草原带,土壤微生物生物量生态化学计量学特征具有更加严格的约束比例.在黄土丘陵区,土壤微生物生物量N∶P或许可以作为判断养分限制的另一个有力工具,若将土壤微生物生物量N∶P与植物叶片N∶P配合使用可能有助于我们更加精确地判断黄土丘陵区的土壤养分限制情况.     

Temporal changes in soil C‐N‐P stoichiometry over the past 60 years across subtropical China

Controlled experiments have shown that global changes decouple the biogeochemical cycles of carbon (C), nitrogen (N), and phosphorus (P), resulting in shifting stoichiometry that lies at the core of ecosystem functioning. However, the response of soil stoichiometry to global changes in natural ecosystems with different soil depths, vegetation types, and climate gradients remains poorly understood. Based on 2,736 observations along soil profiles of 0–150 cm depth from 1955 to 2016, we evaluated the temporal changes in soil C‐N‐P stoichiometry across subtropical China, where soils are P‐impoverished, with diverse vegetation, soil, and parent material types and a wide range of climate gradients. We found a significant overall increase in soil total C concentration and a decrease in soil total P concentration, resulting in increasing soil C:P and N:P ratios during the past 60 years across all soil depths. Although average soil N concentration did not change, soil C:N increased in topsoil while decreasing in deeper soil. The temporal trends in soil C‐N‐P stoichiometry differed among vegetation, soil, parent material types, and spatial climate variations, with significantly increased C:P and N:P ratios for evergreen broadleaf forest and highly weathered Ultisols, and more pronounced temporal changes in soil C:N, N:P, and C:P ratios at low elevations. Our sensitivity analysis suggests that the temporal changes in soil stoichiometry resulted from elevated N deposition, rising atmospheric CO₂ concentration and regional warming. Our findings revealed that the responses of soil C‐N‐P and stoichiometry to long‐term global changes have occurred across the whole soil depth in subtropical China and the magnitudes of the changes in soil stoichiometry are dependent on vegetation types, soil types, and spatial climate variations.     

Land-use change in a tropical mountain rainforest region of southern Ecuador affects soil microorganisms and nutrient cycling

Over the past decades, the tropical mountain rainforest of southern Ecuador has been threatened by conversion to cattle pastures. Frequently, these pastures are invaded by bracken fern and abandoned when bracken becomes dominant. Changes in land-use (forest–pasture–abandoned pasture) can affect soil microorganisms and their physiological responses with respect to soil carbon and nutrient cycling. In situ investigations on litter decomposition and soil respiration as well as biogeochemical characterization of the soil were carried out to identify the driving factors behind. The conversion of forest to pasture induced a pronounced increase in CO₂–C effluxes to 12.2 Mg ha^?1 a^?1 which did not decrease after abandonment. Soil microbial activity and biomass showed a different pattern with lowest values at forest and abandoned pasture sites. With 3445 mg kg^?1 (0–5 cm) microbial biomass carbon (MBC by CFE-method), the active pasture had a more than three times higher value than forest and abandoned pasture, which was among the highest in tropical pasture soils. A shift in the microbial community structure (phospholipid fatty acid, PLFA) was also induced by the establishment of pasture land; the relative abundance of fungi and Gram-negative bacteria increased. PLFA fingerprints of the forest organic layer were more similar to pasture than to forest mineral soil. Chemical properties (pH value, exchangeable cations) were the main factors influencing the respective microbial structure. Bracken-invasion resulted in a decrease in the quantity and quality of above- and belowground biomass. The lower organic substance and nutrient availability induced a significant decline in microbial biomass and activity. After pasture abandonment, these differences in soil microbial function were not accompanied by pronounced shifts in the community structure and in soil pH as was shown for the conversion to pasture. A disconnection between microbial structure and function was identified. Similar soil CO₂–C effluxes between active and abandoned pasture sites might be explained by an underestimation of the effluxes from the active pasture site. All measurements were carried out between grass tussocks where fine-root density was about 2.6 times lower than below tussocks. Thus, lower proportions of root respiration were expected than below tussocks. Overall, soil microorganisms responded differently to changes in land-use from forest to pasture and from pasture to abandoned pasture resulting in pronounced changes of carbon and nutrient cycling and hence of ecosystem functioning.     

Variation of carbon and nitrogen stoichiometry along a chronosequence of natural temperate forest in northeastern China

Aims Carbon (C) and nitrogen (N) stoichiometry contributes to understanding elemental compositions and coupled biogeochemical cycles in ecosystems. However, we know little about the temporal patterns of C:N stoichiometry during forest development. The goal of this study is to explore the temporal patterns of intraspecific and ecosystem components' variations in C:N stoichiometry and the scaling relationships between C and N at different successional stages.Methods Along forest development in a natural temperate forest, northeastern China, four age gradients were categorized into ca. 10-, 30-, 70- and 200-year old, respectively, and three 20 m × 20 m plots were set up for each age class. Leaves, branches, fine roots and fresh litter of seven dominant species as well as mineral soil at depth of 0–10 cm were sampled. A Universal CHN Elemental Analyzer was used to determine the C and N concentrations in all samples.Important findings Intraspecific leaf C, N and C:N ratios remained stable along forest development regardless of tree species; while C, N concentrations and C:N ratios changed significantly either in branches or in fine roots, and they varied with tree species except Populus davidiana (P < 0.05). For ecosystem components, we discovered that leaf C:N ratios remained stable when stand age was below ca. 70 years and dominant tree species were light-demanding pioneers such as Betula platyphylla and Populus davidiana, while increased significantly at the age of ca. 200 years with Pinus koraiensis as the dominant species. C:N ratios in branches and fresh litter did not changed significantly along forest development stages. C concentrations scaled isometrically with respect to N concentrations in mineral soil but not in other ecosystem components. Our results indicate that, leaf has a higher intraspecific C:N stoichiometric stability compared to branch and fine root, whereas for ecosystem components, shifts in species composition mainly affect C:N ratios in leaves rather than other components. This study also demonstrated that C and N remain coupled in mineral soils but not in plant organs or fresh litter during forest development.     

Substrate stoichiometry determines nitrogen fixation throughout succession in southern Chinese forests

The traditional view holds that biological nitrogen (N) fixation often peaks in early‐ or mid‐successional ecosystems and declines throughout succession based on the hypothesis that soil N richness and/or phosphorus (P) depletion become disadvantageous to N fixers. This view, however, fails to support the observation that N fixers can remain active in many old‐growth forests despite the presence of N‐rich and/or P‐limiting soils. Here, we found unexpected increases in N fixation rates in the soil, forest floor, and moss throughout three successional forests and along six age‐gradient forests in southern China. We further found that the variation in N fixation was controlled by substrate carbon(C) : N and C : (N : P) stoichiometry rather than by substrate N or P. Our findings highlight the utility of ecological stoichiometry in illuminating the mechanisms that couple forest succession and N cycling.