TRXB土壤学报Acta Pedologica Sinica0564-3929土壤学报编辑部中国南京trxb-59-6-147910.11766/trxb202012270705S154.2A综述与评论Reviews and Comments微生物残体在土壤中的积累转化过程与稳定机理研究进展Research Progress on Accumulation, Turnover and Stabilization of Microbial Residues in Soil张彬ZHANGBin
Microorganisms are the engine driving the biogeochemical cycling of soil elements and play an important role in the transformation of soil organic matter (SOM). They decompose SOM and release CO2 into the atmosphere through mineralization on the one hand and transform SOM into their cell components through assimilation on the other hand. These cell components can be accumulated in the soil as microbial residues after their death. There is increasing recognition that microbial residues are important precursors of SOM formation and contribute significantly to long-term SOM stabilization. Therefore, this paper calls for scientists to pay more attention and study the role of microbial residues in the accumulation, turnover and stabilization of SOM, and possible underlying mechanisms. To achieve this objective, this paper first explains the processes of how microorganisms continuously produce microbial residues in soils via assimilation and emphasizes the extent to which microbial residues contribute to soil stable carbon (C) pool. Subsequently, this paper introduces the (i) quantification of microbial residues using amino sugar analysis, (ii) conversion of amino sugar data into microbial residue C data to account for the proportion of microbial-derived C in soil organic C and (iii) distinction of original- and newly-formed microbial residues with isotopic labelling techniques to indicate the turnover of microbial residues in soil. Furthermore, this paper summarizes the key external factors influencing the accumulation and turnover of microbial residues. These factors include: (1) nutrition management that can directly influence substrate availability for soil microorganisms and consequently the production and accumulation of microbial residues, even though soil fungi and bacteria may respond differently to substrate addition; (2) tillage practices which generally reduce the accumulation of microbial residues through the destruction of fungal hyphae and breakdown of soil aggregates; (3) land-use change that can permanently impact the contribution of microbial-derived C to soil organic C; and (4) climate change factors which include temperature elevation, elevated CO2 concentration and nitrogen deposition. The fourth section of this paper summarizes the potential stabilization mechanisms of microbial residues in soil, which include chemical protection by attaching to soil mineral surfaces, physical protection by occluding in soil aggregates and delayed decomposition due to the chemical structure of microbial residues. In the last section, some perspectives are provided for the scientific issues that need to be further studied regarding microbial residue contribution to SOM: (a) combine microbial residues with living microbial communities to link with the processes of microbial assimilation from both instantaneous and continuous perspectives; (b) explore the distribution process and stabilization mechanism of microbial residues with soil minerals; (c) investigate the accumulation and turnover of microbial residues in subsurface soils as soil physicochemical properties and microbial community composition change substantially with increase in depth. These discussions will provide a clue to clarify the role of microbial anabolism driving and involving SOM formation and stabilization as well as the underlying relationship between SOM turnover and microbial process in terrestrial ecosystem.
土壤有机质微生物残体周转稳定机理Soil organic matterMicrobial residueTurnoverStabilization国家自然科学基金项目42077085国家自然科学基金项目41977025南京信息工程大学人才启动基金2018r101国家自然科学基金项目(42077085,41977025)和南京信息工程大学人才启动基金(2018r101)资助the National Natural Science Foundation of China42077085the National Natural Science Foundation of China41977025the Startup Foundation for Introducing Talent of NUIST of China2018r101Supported by the National Natural Science Foundation of China(Nos. 42077085 and 41977025)and the Startup Foundation for Introducing Talent of NUIST of China(No. 2018r101)
理论上,跟踪和描述土壤氨基糖的合成转化动态变化,在探究和评价微生物通过同化代谢作用参与有机质转化和持续固碳过程中的作用具有独特的意义和优势[9]。然而,从氨基糖的数量变化上无法准确推知微生物利用外源底物合成氨基糖的动态变化及评价微生物活性的相对变化。从技术手段上,只有实现土壤中新生成的微生物残体和土壤中原有微生物残体的有效区分,才能在化合物分子水平上探讨外源碳素的微生物转化动态与稳定机制。He等[28]建立的气相色谱-质谱联机技术(gas chromatographymass spectrometry,GC-MS)为研究外源标记底物进入氨基糖的富集转化动态提供了强有力的支持。GC-MS技术可将时间因素纳入研究范围,对目标化合物进行动态特征研究,是分析氨基糖中同位素比例变化的有效方法之一[29]。通过加入含13C或15N的标记底物并跟踪13C在氨基糖“碳骨架”的结合或15N在“氨基位”富集随时间的变化,可计算微生物利用标记碳氮合成细胞壁物质的速率和容量,从而反映微生物新陈代谢速率,进而推知微生物对土壤碳氮截获及对SOM循环的贡献[28,30]。通过研究添加活性碳源(葡萄糖)后微生物同化无机氮素(15N标记硫酸铵)合成氨基糖的过程,表明碳源的可利用性是影响微生物同化无机氮素构建细胞组分的主控因素,在有活性碳源存在的条件下,铵态氮较硝态氮(15N标记硫酸铵和15N标记硝酸钾)更易于被微生物优先利用,并且短期内细菌较真菌对活性底物添加的响应更为强烈,表现为细菌残体在底物添加初期快速合成,而真菌残体在后期的合成强度逐渐超过细菌[31]。此外,气相色谱-燃烧-同位素比例质谱联用技术(gas chromatography-combustion-isotopic ratio mass spectrometry,GC-C-IRMS)也可进行自然丰度或人工标记13C底物添加条件下氨基糖δ13C的测定,用于评价微生物参与碳转化及微生物死亡残体介导的有机碳截获的研究中[9]。但应用该技术测定氨基糖的同位素丰度值时,由于衍生过程中会引入碳原子,所以会引起同位素的分馏。另外一种用于氨基糖同位素分析的方法是阴离子交换高效液相色谱法(high-performance anion-exchange liquid chromatography,HPAE-LC-IRMS),由于该方法不需要对样品进行衍生,前期准备简单,同位素矫正更为直接,是一种很有前景的化合物同位素测定方法[32]。但该方法分离氨基糖的精确度较低,尤其是含量较低的甘露糖和胞壁酸,需要进一步优化[11]。
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