微生物对分解底物碳氮磷化学计量的响应和调节机制

微生物分解者的生存和生长策略、群落结构和功能会随着底物化学计量特征而改变, 从而强烈地影响底物的分解速度和元素的周转速率, 进而影响生态系统的功能过程。深入理解微生物生理代谢活动和群落结构与陆地生态系统过程之间的联系及其对全球变化的响应和反馈机理是生态学和全球变化生物学领域值得关注的重大科学命题。该文基于生态化学计量学理论和代谢理论, 首先介绍微生物在陆地生态系统碳氮磷循环中的作用; 然后综述微生物对分解底物化学计量变异性的响应和调节的4种主要机制: (1)调整微生物自身化学计量特征; (2)调整微生物群落结构; (3)产生特定的胞外酶以获取受限制的资源; (4)改变碳氮磷元素利用效率。最后, 通过分析当前研究不足, 提出该领域亟需关注的科学问题有: (1)综合阐明微生物对底物化学计量变化响应的各种机制及其相对重要性; (2)探索全球变化对微生物驱动的碳氮磷循环的影响; (3)探索微生物对底物化学计量变化适应对策的时空变化。     

氮添加土壤微生物群落结构影响微生物碳利用效率

尽管近年来中国氮(N)沉降水平逐渐趋于稳定,但中国东南地区N沉降相比于其他地区仍处于较高水平。N沉降对陆地生态系统碳循环过程的影响不容忽视。微生物碳利用效率(CUE)是指微生物将吸收的碳转化为生物量碳的效率,高微生物CUE意味着高土壤有机碳存储潜力。因此,探究N沉降背景下微生物CUE的变化将有助于进一步认识陆地生态系统土壤碳存储的变化。然而,目前关于N沉降下微生物群落结构的变化如何影响微生物CUE鲜有报道。在福建省泉州市戴云山国家级自然保护区的罗浮栲林通过N添加模拟N沉降。实验共包括三个N添加处理：对照(CT,+0 kg hm^-2 a^-1)、低氮(LN,+40 kg hm^-2 a^-1)和高氮(HN,+80 kg hm^-2 a^-1)。测定不同处理土壤基本理化性质、微生物生物量、酶活性和CUE,并使用高通量测序对微生物群落结构和多样性进行测定。结果表明,N添加显著影响微生物CUE,随着N添加水平的增加,CUE逐渐增加;相反,土壤pH、可提取有机碳(EOC)和微...     

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。     

植物磷获取机制及其对全球变化的响应

磷是植物生长的必需元素,而陆地生态系统普遍存在磷限制,全球变化可能会影响土壤磷循环过程,进一步加剧磷限制,探讨植物磷获取策略对科学预测生态系统生产力如何适应全球变化具有重要意义。该文通过收集和梳理相关文献,从4个方面综述植物的磷获取机制及其对全球变化的响应:1)植物的磷饥饿响应机制;2)植物的磷获取途径和策略;3)土壤微生物对植物磷吸收的影响; 4)植物磷吸收对全球变化(温度升高、氮沉降和降水变化)的响应及其机制。该综述有助于深入理解全球变化背景下植物适应低磷胁迫的机理,也可为养分资源管理实践提供理论依据。     

Stoichiometric homeostasis in response to variable water and nutrient supply in a Robinia pseudoacacia plant–soil system

刺槐植物-土壤系统生态化学计量内稳性对水分和养分变异的响应特征 所有生物体都需要一定比例的元素来维持正常的生理代谢过程,它们的可塑性取决于它们利用外部资源的效率。阐明不同资源供应水平下植物、土壤和土壤微生物生物量生态化学计量特征之间的相互作用非常重要。本研究以一年生刺槐(Robinia pseudoacacia)幼苗为研究对象,测定不同水平水分、氮素和磷素处理下刺槐叶片、细根、土壤和微生物生物量C、N、P含量及其化学计量学指标。结果表明,刺槐叶片、细根、土壤和微生物生物量C、N、P含量及其化学计量特征会对其生存环境水分和养分条件的变化表现出一定程度的可塑性;方差分解分析结果表明,细根计量比解释了微生物生物量计量比方差的很大一部分;结构方程模型进一步揭示了细根计量比和叶片计量比是影响土壤微生物生物量C:N和C:P 的两个直接因素,而细根计量比具有较大的直接作用。此外,内稳性特征分析表明土壤微生物生物量C 和C:P对土壤养分变化较为敏感,其他指标均具有内稳性。这些结果明确了土壤微生物生物量化学计量的重要性,提高我们对不同生境水分和养分供应水平下植物-土壤系统养分循环机理的认识。     

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。     

植被类型变化对土壤微生物碳利用效率的影响研究进展

植被类型变化强烈影响着土壤碳循环。土壤微生物碳利用效率（CUE）是微生物将从环境中获取的碳分配给自身生长的比例，是土壤碳循环的综合指标。研究植被类型变化对CUE的影响有助于从微生物视角理解该过程中的土壤碳动态，可以为评估植被类型变化对土壤质量及生态系统碳循环的影响提供基础，具有重要的理论及实际价值。通过系统查阅相关文献，综述了植被类型变化导致的CUE变化情况，以及该过程中影响CUE的因子与机制。目前，相关研究主要涉及以林地、草地和农业用地为起点或终点的植被变化类型。天然林（原生林、次生林）变化为人工林、林地变化为草地后CUE普遍下降，随终点植被的发展CUE可能恢复至起点水平。植被成熟度越高，发生转变时CUE变化越剧烈。植被类型变化以农业用地为起点或终点时，CUE变化方向的不确定性及幅度的变异性均增加。植被类型变化导致的CUE变化主要受到植被、土壤、微生物因子及其交互作用的驱动，指示CUE的指标、采样季节和土层也会一定程度上影响CUE的变化。今后相关研究应采用直接的CUE测定方法，拓宽研究气候区及植被变化类型，关注植被变化过程中CUE变化的土层差异及动态监测，深入对植被类型变化导致的生态环境因子变化与CUE的关系及作用机制的研究。     

太白山不同海拔森林根际土壤微生物碳利用效率差异性及其影响因素

探索森林根际土壤微生物碳利用效率(CUE)是权衡森林生态系统微生物合成代谢和分解代谢强弱的关键过程。然而不同海拔森林根际土壤微生物CUE的变化规律与影响因子尚不清楚。该研究选取秦岭太白山6个不同海拔的森林根际土壤为研究对象,测定其理化性质、胞外酶活性、微生物群落与植被特征等指标,利用酶化学计量比计算微生物CUE,分析根际土壤微生物CUE沿海拔梯度的变化规律,定量研究其影响因子。结果表明:根际土壤微生物CUE随海拔升高总体呈上升趋势。CUE从最低海拔的0.505至最高海拔的0.527升高了4.36%,但在海拔1 603和2 405m处出现了下降。海拔梯度内根际土壤微生物CUE变化受多种环境因子综合影响,土壤基质的影响(如可溶性有机碳和铵态氮含量)占主导地位,植被因子次之,二者分别解释了CUE变化的17.0%和5.7%,且二者相互作用解释了CUE变化的31.9%。研究结果可为秦岭森林土壤微生物碳同化能力和固碳潜力,以及全球变化背景下森林土壤碳循环提供科学依据。     

全球大气CO2浓度升高对土壤微生物的影响

全球大气CO2浓度升高对土壤微生物生态系统的影响已引起广泛关注。本文从土壤微生物群落结构、微生物区系、土壤呼吸、微生物生物量以及土壤酶活性方面对大气高浓度CO2的响应进行了综述。由于提供高浓度CO2的实验系统、所选植物材料以及土壤特性等的不同,大气CO2浓度升高对土壤微生物群落结构、微生物区系、土壤呼吸、微生物生物量以及土壤酶活性的影响并未得出一致结论。但高浓度CO2对土壤微生物生态系统的影响是存在的。     

罗浮栲林土壤微生物碳利用效率对短期氮添加的响应

作为调节土壤碳矿化过程的重要参数,微生物碳利用效率(CUE)对理解陆地生态系统中的碳循环至关重要。本研究在戴云山罗浮栲林设置对照(0 kg N·hm^-2·a^-1)、低氮(40 kg N·hm^-2·a^-1)和高氮(80 kg N·hm^-2·a^-1) 3个氮添加水平以模拟氮沉降,测定了表层(0～10 cm)土壤基本理化性质、有机碳组分、微生物生物量和酶活性;并利用¹⁸O标记水方法测定土壤微生物CUE,以更好地理解氮沉降加剧对微生物CUE的影响及其影响因素。结果表明: 短期氮添加显著降低了土壤微生物的呼吸速率、碳和氮获取酶活性,但显著增加了土壤微生物CUE。β-N-乙酰氨基酸葡糖苷酶(NAG)/微生物生物量碳(MBC)、微生物呼吸速率、β-葡萄糖苷酶(BG)/MBC、纤维素水解酶(CBH)/MBC和土壤有机碳含量是影响CUE的主要因素,且CUE与NAG/MBC、微生物呼吸速率、BG/MBC和CBH/MBC呈显著负相关,与土壤有机碳呈显著正相关。综上,短期氮添加导致土壤微生物获取碳和氮的成本降低,减少微生物呼吸,从而提高了土壤微生物CUE,这将有助于提高罗浮栲林土壤碳固存潜力。     

Energy use efficiency of soil microorganisms: Driven by carbon recycling and reduction

Carbon use efficiency (CUE) is being intensively applied to quantify carbon (C) cycling processes from microbial cell to global scales. Energy use efficiency (EUE) is at least as important as the CUE because (i) microorganisms use organic C mainly as an energy source and not as elemental C per se, and (ii) microbial growth and maintenance are limited by energy, but not by C as a structural element. We conceptualize and review the importance of EUE by soil microorganisms and focus on (i) the energy content in organic compounds depending on the nominal oxidation state of carbon (NOSC), (ii) approaches to assess EUE, (iii) similarities and differences between CUE and EUE, and (iv) discuss mechanisms responsible for lower EUE compared to CUE. The energy content per C atom (enthalpy of combustion, the total energy stored in a compound) in organic compounds is very closely (R² = 0.98) positively related to NOSC and increases by 108 kJ mol⁻¹ C per one NOSC unit. For the first time we assessed the NOSC of microbial biomass in soil (−0.52) and calculated the corresponding energy content of −510 kJ mol⁻¹ C. We linked CUE and EUE considering the NOSC of microbial biomass and element compositions of substrates utilized by microorganisms. The mean microbial EUE (0.32–0.35) is 18% lower than CUE (0.41) using glucose as a substrate. This definitely indicates that microbial growth is limited by energy relative to C. Based on the comparison of a broad range of processes of C and energy utilization for cell growth and maintenance, as well as database of experimental CUE from various compounds, we clearly explained five mechanisms and main factors why EUE is lower than CUE. The two main mechanisms behind lower EUE versus CUE are: (i) microbial recycling: C can be microbially recycled, whereas energy is always utilized only once, and (ii) chemical reduction of organic and inorganic compounds: Energy is used for reduction, which is ongoing without C utilization.     

Increased microbial growth,biomass, and turnover drive soil organic carbon accumulation at higher plant diversity

Species‐rich plant communities have been shown to be more productive and to exhibit increased long‐term soil organic carbon (SOC) storage. Soil microorganisms are central to the conversion of plant organic matter into SOC, yet the relationship between plant diversity, soil microbial growth, turnover as well as carbon use efficiency (CUE) and SOC accumulation is unknown. As heterotrophic soil microbes are primarily carbon limited, it is important to understand how they respond to increased plant‐derived carbon inputs at higher plant species richness (PSR). We used the long‐term grassland biodiversity experiment in Jena, Germany, to examine how microbial physiology responds to changes in plant diversity and how this affects SOC content. The Jena Experiment considers different numbers of species (1–60), functional groups (1–4) as well as functional identity (small herbs, tall herbs, grasses, and legumes). We found that PSR accelerated microbial growth and turnover and increased microbial biomass and necromass. PSR also accelerated microbial respiration, but this effect was less strong than for microbial growth. In contrast, PSR did not affect microbial CUE or biomass‐specific respiration. Structural equation models revealed that PSR had direct positive effects on root biomass, and thereby on microbial growth and microbial biomass carbon. Finally, PSR increased SOC content via its positive influence on microbial biomass carbon. We suggest that PSR favors faster rates of microbial growth and turnover, likely due to greater plant productivity, resulting in higher amounts of microbial biomass and necromass that translate into the observed increase in SOC. We thus identify the microbial mechanism linking species‐rich plant communities to a carbon cycle process of importance to Earth's climate system.     

Chemical changes in organic matter after fungal colonization in a nitrogen fertilized and unfertilized Norway spruce forest

Background and aims

Mineralization of soil organic matter (SOM) constitutes a major carbon flux to the atmosphere. The carbon use efficiency (CUE) of the saprotrophic microorganisms mineralizing SOM is integral for soil carbon dynamics. Here we investigate how the CUE is affected by temperature, metabolic conditions, and the molecular complexity of the substrate.

Methods

We incubated O-horizon soil samples (with either ¹³C–glucose or ¹³C–cellulose) from a boreal coniferous forest at 4, 9, 14, and 19 °C, and calculated CUEs based on the amount of ¹³C–CO₂ and ¹³C–labelled microbial biomass produced. The effects of substrate, temperature, and metabolic conditions (representing unlimited substrate supply and substrate limitation) on CUE were evaluated.

Results

CUE from metabolizing glucose was higher as compared to cellulose. A slight decrease in CUE with increasing temperature was observed in glucose amended samples (but only in the range 9–19 °C), but not in cellulose amended samples. CUE differed significantly with metabolic conditions, i.e. CUE was higher during unlimited growth conditions as compared to conditions with substrate limitation.

Conclusions

We conclude that it is integral to account for both differences in CUE during different metabolic phases, as well as complexity of substrate, when interpreting temperature dependence on CUE in incubation studies.

     

Evaluating soil microbial carbon use efficiency explicitly as a function of cellular processes: implications for measurements and models

Carbon use efficiency (CUE), the proportion of carbon (C) consumed by microbes that is converted into biomass, is an important parameter for soil C models with explicit microbial controls. While often considered as a single parameter, CUE is an emergent property of multiple microbial processes, including assimilation efficiency, biomass-specific respiration, enzyme production, and respiratory costs of enzyme production. These processes occur over variable time scales and imply different fates for C, and the same emergent CUE value can result when C is allocated in fundamentally different ways (e.g. a high investment in enzyme production vs. a high assimilation cost). We developed a model that represents the individual processes underlying emergent CUE to test how shifts in microbial allocation alter equilibrium soil C pool sizes. We found that an increase in emergent CUE that results from a change in assimilation efficiency, biomass specific respiration, or respiration costs from enzyme production causes soil organic C (SOC) to decline, while the same change in emergent CUE resulting from a change in enzyme production causes SOC to increase. We also used the model to test the sensitivity of CUE from isotopic C tracer estimates to changes in microbial allocation processes. We found that these estimates do not account for the same microbial processes represented by emergent CUE in models. We propose that considering microbial processes explicitly rather than representing CUE as a single parameter can improve data-model integration. In addition, modeling microbial processes explicitly will account for a wider range of possible outcomes from shifts in microbial C allocation, such as when increased SOC results from increasing CUE.     

土壤酶计量揭示了武夷山黄山松林土壤微生物沿海拔梯度的碳磷限制变化

了解土壤胞外酶活性和酶计量的变化对评估山地生态系统土壤养分有效性和微生物的营养限制状况具有重要意义.然而,亚热带山地森林土壤微生物的营养限制状况对海拔梯度变化的响应及其驱动因素尚不清楚.本研究以武夷山不同海拔(1200～2000 m)黄山松林为对象,测定了土壤基本性质、微生物生物量以及与碳(C)、氮(N)、磷(P)循环...     

Does declining carbon‐use efficiency explain thermal acclimation of soil respiration with warming?

Enhanced soil respiration in response to global warming may substantially increase atmospheric CO₂ concentrations above the anthropogenic contribution, depending on the mechanisms underlying the temperature sensitivity of soil respiration. Here, we compared short‐term and seasonal responses of soil respiration to a shifting thermal environment and variable substrate availability via laboratory incubations. To analyze the data from incubations, we implemented a novel process‐based model of soil respiration in a hierarchical Bayesian framework. Our process model combined a Michaelis–Menten‐type equation of substrate availability and microbial biomass with an Arrhenius‐type nonlinear temperature response function. We tested the competing hypotheses that apparent thermal acclimation of soil respiration can be explained by depletion of labile substrates in warmed soils, or that physiological acclimation reduces respiration rates. We demonstrated that short‐term apparent acclimation can be induced by substrate depletion, but that decreasing microbial biomass carbon (MBC) is also important, and lower MBC at warmer temperatures is likely due to decreased carbon‐use efficiency (CUE). Observed seasonal acclimation of soil respiration was associated with higher CUE and lower basal respiration for summer‐ vs. winter‐collected soils. Whether the observed short‐term decrease in CUE or the seasonal acclimation of CUE with increased temperatures dominates the response to long‐term warming will have important consequences for soil organic carbon storage.     

Carbon use efficiency of microbial communities: stoichiometry,methodology and modelling

Carbon use efficiency (CUE) is a fundamental parameter for ecological models based on the physiology of microorganisms. CUE determines energy and material flows to higher trophic levels, conversion of plant‐produced carbon into microbial products and rates of ecosystem carbon storage. Thermodynamic calculations support a maximum CUE value of ~ 0.60 (CUE _max). Kinetic and stoichiometric constraints on microbial growth suggest that CUE in multi‐resource limited natural systems should approach ~ 0.3 (CUE _max/2). However, the mean CUE values reported for aquatic and terrestrial ecosystems differ by twofold (~ 0.26 vs. ~ 0.55) because the methods used to estimate CUE in aquatic and terrestrial systems generally differ and soil estimates are less likely to capture the full maintenance costs of community metabolism given the difficulty of measurements in water‐limited environments. Moreover, many simulation models lack adequate representation of energy spilling pathways and stoichiometric constraints on metabolism, which can also lead to overestimates of CUE. We recommend that broad‐scale models use a CUE value of 0.30, unless there is evidence for lower values as a result of pervasive nutrient limitations. Ecosystem models operating at finer scales should consider resource composition, stoichiometric constraints and biomass composition, as well as environmental drivers, to predict the CUE of microbial communities.     

Optimal metabolic regulation along resource stoichiometry gradients

Most heterotrophic organisms feed on substrates that are poor in nutrients compared to their demand, leading to elemental imbalances that may constrain their growth and function. Flexible carbon (C)‐use efficiency (CUE, C used for growth over C taken up) can represent a strategy to reduce elemental imbalances. Here, we argue that metabolic regulation has evolved to maximise the organism growth rate along gradients of nutrient availability and translated this assumption into an optimality model that links CUE to substrate and organism stoichiometry. The optimal CUE is predicted to decrease with increasing substrate C‐to‐nutrient ratio, and increase with nutrient amendment. These predictions are generally confirmed by empirical evidence from a new database of c. 2200 CUE estimates, lending support to the hypothesis that CUE is optimised across levels of organisation (microorganisms and animals), in aquatic and terrestrial systems, and when considering nitrogen or phosphorus as limiting nutrients.