稻田土壤微生物残体积累对外源秸秆输入的响应研究进展

稻田土壤碳循环是我国陆地生态系统碳循环的重要组成部分。促进稻田生态系统碳的固定及稳定对减缓全球气候变化起着不容忽视的作用。微生物主导的有机碳转化过程是土壤碳循环研究的核心,微生物同化代谢介导的细胞残体迭代积累在土壤有机碳长期截获和稳定过程中发挥重要作用。与旱地土壤相比,关于稻田土壤中微生物残体积累动态对外源有机物质如作物秸秆输入的响应及主要影响因子的认识还相对有限,对微生物通过同化作用参与土壤固碳的过程和机制尚缺乏系统认识。基于此,本文介绍了微生物残体对土壤有机碳库形成和积累的重要性及评价指标,重点探讨了秸秆还田对稻田土壤微生物残体积累动态以及外源秸秆碳形成细胞残体转化过程的影响,分析了影响微生物残体积累转化的主要气候因素和土壤因素,最后提出了未来应借助先进的光谱和高分辨率成像技术并结合同位素示踪对微生物残体的稳定性与机理开展更为深入的研究。     

植物残体向土壤有机质转化过程及其稳定机制的研究进展

土壤有机质的数量和质量不仅是衡量土壤肥力状况的核心要素,其形成、转化及稳定过程还与全球气候变化密切相关。植物残体是土壤有机质的初始来源,但由于其腐解过程的复杂、多变性以及土壤有机质、微生物的高度异质性,植物残体向土壤有机质的转化和稳定机理尚不十分明确。本文介绍并讨论了近年来关于植物残体向土壤有机质转化相关研究的新发现,探讨了微生物源和植物源有机质对土壤有机质的贡献,概述了土壤有机质形成的微生物驱动机制,并综述了植物残体输入后土壤有机质稳定性的相关研究,最后对该研究领域未来的发展进行展望,以期能够为科学地提高土壤的固碳能力提供参考。     

森林次生演替过程中有机质层和矿质层土壤微生物残体的变化

微生物对土壤有机质(SOM)转化和形成具有重要作用,然而微生物残体对SOM贡献的评估仍是目前的热点。以长白山5个森林次生演替序列(20 a、80 a、120 a、200 a和≥300 a)和2个土壤深度(0～5 cm有机质层和5～15 cm矿质层)为对象,利用氨基糖和中红外光谱技术,探究森林次生演替过程中微生物残体变化及其对土壤有机碳(SOC)的贡献。森林次生演替序列80～200 a显著增加了有机质层和矿质层微生物残体含量及其对SOC的贡献,而在演替300 a均显著降低。森林演替80～200a有机质层和矿质层土壤芳香族碳组分/多糖较低,有利于微生物生物量碳(MBC)产生和微生物碳利用(高的MBC/SOC),促进微生物残体积累及其对SOC的贡献;而演替300 a芳香族碳组分/多糖较高,抑制MBC产生和微生物碳利用,导致微生物残体及其对SOC贡献的下降。SOC含量差异导致不同土壤深度微生物残体含量的变化,有机质层高的SOC产生高的MBC,进而刺激微生物残体积累;此外,有机质层难利用SOM组分高于矿质层,导致真菌残体对SOC的贡献比例下降,而细菌残体的贡献增加。     

丛枝菌根真菌介导的土壤有机碳稳定机制研究进展

土壤有机碳(SOC)的稳定是陆地生态系统碳循环的关键过程之一,对维持土壤肥力和减少温室气体排放具有重要意义。以往认为植物残体中难降解性物质的物理保护和腐殖质影响土壤中有机碳库的稳定性。最近的研究结果表明,微生物介导的碳循环过程在土壤有机碳稳定中发挥着重要作用。丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)作为土壤中一类重要的共生微生物,参与植物光合碳向土壤的转运和分配,是陆地生态系统碳循环的重要一环,但其在土壤有机碳稳定中的作用潜力还未得到充分挖掘。基于此,本文估算了植物光合碳在AMF根外菌丝的分配量;总结了AMF介导的土壤有机碳稳定机制,主要包括AMF活体菌丝对碳的截留,分泌物及残体的分子结构抗性和土壤矿物吸附,提高植物源碳的质量和数量,菌丝分泌物及残体的激发效应和稳定土壤团聚体;探讨了影响AMF介导的稳定性有机碳形成的非生物(气候因子、土壤养分和土壤矿物)和生物因子(植物和AMF种类);提出了AMF与土壤有机碳周转互作机理进一步的研究方向,包括探究菌根植物光合碳转化为稳定性SOC的机制,解析不同生态系统中AMF对稳定性SOC的贡献及影响因素,并厘清...     

营养动力学为基础的生物残体分解的数学模型研究

生物残体是由活的生物(包括动物和植物)死亡后形成,如森林砍伐后遗留的树桩,温带森林中每年一度集中产生的枯枝落叶,森林和草原中大型动物死亡遗留的尸体,这些生物残体一般都独自形成一个实体,不与土壤进行均匀的混合,它们的分解,一般主要不是依靠贮积在土壤中的酶或微生物的作用,它们的分解要经过一个在残体内重新滋长微生物的过程,使残体逐渐腐烂变质,进行分解。因此,生物残体的分解,与其说是分解过程,还不如说是一个微生物的培养过程,或者说是这两个过程相互交错,一方面是微生物种群的增长,另一方面是残体中有机物质被微生物分解、转化、利用和复合成新的有机体,对于这样一个复杂的过程,通常适用于土壤有机质分解过程的Stanford-Smith方程^[2]就不适用了,而建立在营养动力学基础上的单种群模型(崔-Lawson方程^[3-5])却可加以转化用来描述这个过程。     

基于固态13C核磁共振波谱研究植物残体分解和转化机制的进展

植物残体在土壤中的分解和转化影响了其养分归还和有机质形成过程。由于缺乏高分辨率的分析方法,对不同气候、植被和土壤类型条件下植物残体在分解过程中化学结构组成的演变特征和机制仍不清楚。核磁共振波谱技术在解析自然有机物化学组成方面具有独特的优势,本文综述了基于固态~(13)C核磁共振波谱(solid-state ~(13)C-NMR spectroscopy)技术评价植物残体的基质质量、解析植物残体的分解速率及其官能团组成的变化特征、揭示土壤腐殖质特性等方面的主要进展。未来针对植物残体分解和有机质形成机制的研究,应该结合稳定性同位素质谱和扫描电镜分析方法,综合分析植物残体中的有机化合物组成和物理结构;从多时空尺度揭示不同类型植物残体中有机碳官能团的降解路径;结合高通量测序和基因芯片分析方法,深入研究土壤微生物群落与植物残体化学结构的协同演变机制,提出不同气候–土壤–植被类型区促进土壤有机质形成的调控措施。     

稳定性同位素技术在土壤重要有机组分循环转化研究中的应用

土壤有机质包含多种不同结构的有机物质,如碳水化合物(中性糖、氨基糖)、蛋白质(氨基酸)、木质素等,这些组分在土壤中的保留时间从几天到几百万年,在有机质循环过程中的积累和转化动态各有特征,其作用和贡献也有所不同.由于有机质各组分本身在土壤中稳定存在,只有利用同位素示踪技术才能定量研究有机质及其组分的循环动态.本文概述了利用稳定同位素示踪技术研究有机质中的一些重要组分的来源、可利用性和转化动态及其生物标识作用的进展情况,从而深入了解土壤有机质循环转化动态及调控机制.     

不同土地利用方式对土壤微生物生物量碳的影响

研究了长期定位试验不同土地利用方式下土壤微生物生物量碳的变化,结果表明:裸地处理的微生物生物量碳最低;不施肥种植作物(CK)或施肥种植作物(NPK)能显著提高土壤微生物量碳含量;NPK配合猪厩肥处理的微生物体碳含量显著高于CK和NPK处理;荒地处理由于返回的植物残体量很大,积累的有机物很多,所以其微生物量碳最大;其次为割草处理和林地处理。有机质含量高的土壤其土壤微生物量也高,土壤微生物碳的周转强度也大,周转时间短,周转速率为:荒地>割草>林地>NPK+C>NPK>CK>裸地。     

土壤孔隙结构与土壤微环境和有机碳周转关系的研究进展

土壤结构是土壤功能的基础,不仅影响土壤养分的供应、水分的保持及渗透、气体的交换等过程,还为土壤微生物提供了物理生境,并调控土壤有机碳的周转这一关键过程。土壤的孔隙特征能够直接、真实地反映土壤结构的好坏;用土壤的孔隙特征作为试验指标能更好地反映土壤结构对这些过程的调节作用。在此基础上,将高度异质性的土壤孔隙结构同土壤微环境的变化和土壤有机碳的周转过程进行定量分析,对深入了解土壤结构在土壤生态系统中的功能至关重要。因此,着重从土壤孔隙结构对土壤微环境的影响及其与有机碳的关系两方面展开,剖析土壤孔隙结构调控作用下的土壤微环境响应过程,阐述土壤孔隙结构对土壤有机碳周转产生的直接、间接影响,强调土壤孔隙结构在调节土壤有机碳周转进程中的重要作用,并对土壤孔隙结构在调节土壤有机碳周转、植物残体分解及其与微生物协调作用机制等方面研究提出展望。     

土壤矿质结合态有机碳形成及稳定机制的研究进展

随着分子生物学的发展，微生物的作用改变了科学界对土壤有机碳(SOC)形成和固持的认知。土壤微生物残体与矿物结合形成矿质结合态有机碳(mineral-associated organic carbon, MAOC)加深了对SOC固存的理解。MAOC是以土壤微生物残体C为主的SOC组分，主要由分子量相对较低且可识别的微生物残体与矿物表面结合而成。由于MAOC对草地和农田生态系统土壤C库的贡献超过50%,且周转时间较长(百年—千年尺度),研究其形成过程和稳定机制已成为碳中和背景下土壤碳汇的焦点。现阶段的研究明确指出，MAOC的形成和稳定不仅与微生物残体C密切相关，还与土壤矿物有着非常紧密的联系。基于此，聚焦土壤微生物“碳泵”调控SOC形成这一前沿科学问题，围绕土壤微生物残体贡献MAOC形成这一科学构架进行概述，旨在揭示不同来源LMW—DC(溶解态低分子量C底物)对MAOC形成的贡献，探讨土壤矿物对LMW—DC选择性吸附机理，探究MAOC贡献稳定C库的影响因素。并对该研究领域未来的发展进行展望，以期能从分子水平出发，探究不同生态系统、土壤类型及土层深度微生物的调控差异，为土壤有机碳固持的研究提...     

无机氮素加入量对玉米秸秆分解过程中棕壤氨基糖含量的影响

作物秸秆还田作为一种调控土壤养分循环、减少氮肥损失、维持和提高土壤有机质水平的有力措施,越来越受到人们的重视[1-2]。还田的秸秆是微生物的碳源和能源,常导致土壤微生物量迅速增加,相应的微生物死亡率和微生物残体积累量也提高[3-4]。根据Appuhn等[5]对微生物细胞壁组分的     

无机氮素加入量对玉米秸秆分解过程中棕壤氨基糖含量的影响

在室内恒温(25℃)培养条件下,通过气相色谱法研究高C/N比玉米秸秆降解过程中微生物来源的氨基糖含量及其占有机质比例的变化及其对无机氮素添加水平(0, 60.3, 167.2, 701.9 mg•N•kg-1土,依次标记为N0, Nlow, Nmed, Nhigh)的响应情况。结果表明：在玉米秸秆分解过程中,土壤中的氨基糖含量及其对有机质贡献的比例随着无机氮素供应水平的增加而增加,即以微生物代谢物形式截获的有机碳/氮相应增多。Nmed和Nhigh处理中氨基糖积累量显著高于Nlow和N0处理。不同微生物来源的氨基糖受外源氮素的影响情况不同,胞壁酸比氨基葡萄糖更易于受到土壤中碳氮供给的影响,具有相对较快的转化速率;而在数量上氨基葡萄糖对土壤有机质的贡献比例显著高于前者;氨基半乳糖在土壤中的积累过程较为缓慢,受外源无机氮素添加水平的影响并不明显。可见,在高C/N比作物残体分解过程中,无机氮素的供应水平是影响土壤中氨基糖积累转化的重要因素之一。但是,过多的无机氮素施入并不能被微生物完全同化利用,因此秸秆还田的土壤中必须要考虑有效氮素的水平问题。     

土壤碳水化合物的转化与累积研究进展

碳水化合物虽然仅占土壤有机质的10%~20%,却是土壤中最活跃的有机碳库,是微生物的主要能源和碳源,并参与土壤团聚体的形成,是土壤有机质和土壤性质研究中的重要指标和对象。本文综述了土壤碳水化合物种类、来源和在团聚体中的分布特征、微生物群落结构对碳水化合物的转化和积聚的影响,同时讨论了耕作、施肥等农业措施对碳水化合物含量和分布的影响。从现有文献资料来看,碳水化合物的含量因土壤粒级不同而异;外源碳水化合物在土壤中的转化和累积与微生物群落结构特征密切相关,细菌在单糖等简单碳水化合物的转化中起主要作用,而复杂碳水化合物则首先在真菌的作用下水解成单糖;真菌和革兰氏阳性菌丰富的土壤更有利于碳水化合物来源碳的累积,但是详细的转化过程和存在形态认识有限,有待深入研究。     

How relevant is recalcitrance for the stabilization of organic matter in soils?

Traditionally, the selective preservation of certain recalcitrant organic compounds and the formation of recalcitrant humic substances have been regarded as an important mechanism for soil organic matter (SOM) stabilization. Based on a critical overview of available methods and on results from a cooperative research program, this paper evaluates how relevant recalcitrance is for the long‐term stabilization of SOM or its fractions. Methodologically, recalcitrance is difficult to assess, since the persistence of certain SOM fractions or specific compounds may also be caused by other stabilization mechanisms, such as physical protection or chemical interactions with mineral surfaces. If only free particulate SOM obtained from density fractionation is considered, it rarely reaches ages exceeding 50 y. Older light particles have often been identified as charred plant residues or as fossil C. The degradability of the readily bioavailable dissolved or water‐extractable OM fraction is often negatively correlated with its content in aromatic compounds, which therefore has been associated with recalcitrance. But in subsoils, dissolved organic matter aromaticity and biodegradability both are very low, indicating that other factors or compounds limit its degradation. Among the investigated specific compounds, lignin, lipids, and their derivatives have mean turnover times faster or similar as that of bulk SOM. Only a small fraction of the lignin inputs seems to persist in soils and is mainly found in the fine textural size fraction (<20 µm), indicating physico‐chemical stabilization. Compound‐specific analysis of ¹³C : ¹²C ratios of SOM pyrolysis products in soils with C3‐C4 crop changes revealed no compounds with mean residence times of > 40–50 y, unless fossil C was present in substantial amounts, as at a site exposed to lignite inputs in the past. Here, turnover of pyrolysis products seemed to be much longer, even for those attributed to carbohydrates or proteins. Apparently, fossil C from lignite coal is also utilized by soil organisms, which is further evidenced by low ¹⁴C concentrations in microbial phospholipid fatty acids from this site. Also, black C from charred plant materials was susceptible to microbial degradation in a short‐term (60 d) and a long‐term (2 y) incubation experiment. This degradation was enhanced, when glucose was supplied as an easily available microbial substrate. Similarly, SOM mineralization in many soils generally increased after addition of carbohydrates, amino acids, or simple organic acids, thus indicating that stability may also be caused by substrate limitations. It is concluded that the presented results do not provide much evidence that the selective preservation of recalcitrant primary biogenic compounds is a major SOM‐stabilization mechanism. Old SOM fractions with slow turnover rates were generally only found in association with soil minerals. The only not mineral‐associated SOM components that may be persistent in soils appear to be black and fossil C.     

Invertebrate control of soil organic matter stability

 The control of soil organic matter (SOM) stability by soil invertebrates is evaluated in terms of their impact on the inherent recalcitrance, accessibility to microorganisms, and interaction with stabilizing substances of organic compounds. Present knowledge on internal (ingestion and associated transformations) and external (defecation, constructions) control mechanisms of soil invertebrates is also reviewed. Soil animals contribute to the stabilization and destabilization of SOM by simultaneously affecting chemical, physical, and microbial processes over several orders of magnitude. A very important aspect of this is that invertebrates at higher trophic levels create feedback mechanisms that modify the spatio-temporal framework in which the micro-food web affects SOM stability. Quantification of non-trophic and indirect effects is thus essential in order to understand the long-term effects of soil biota on SOM turnover. It is hypothesized that the activities of invertebrates which lead to an increase in SOM stability partly evolved as an adaptation to the need for increasing the suitability of their soil habitat. Several gaps in knowledge are identified: food selection and associated changes in C pools, differential effects on SOM turnover, specific associations with microorganisms, effects on dissolution and desorption reactions, humus-forming and humus-degrading processes in gut and faeces, and the modification of invertebrate effects by environmental variables. Future studies must not be confined merely to a mechanistic analysis of invertebrate control of SOM stability, but also pay considerable attention to the functional and evolutionary aspects of animal diversity in soil. This alone will allow an integration of biological expertise in order to develop new strategies of soil management which can be applied under a variety of environmental conditions. Received: 6 April 1999     

Abundance, production and stabilization of microbial biomass under conventional and reduced tillage

Soil tillage practices affect the soil microbial community in various ways, with possible consequences for nitrogen (N) losses, plant growth and soil organic carbon (C) sequestration. As microbes affect soil organic matter (SOM) dynamics largely through their activity, their impact may not be deduced from biomass measurements alone. Moreover, residual microbial tissue is thought to facilitate SOM stabilization, and to provide a long term integrated measure of effects on the microorganisms. In this study, we therefore compared the effect of reduced (RT) and conventional tillage (CT) on the biomass, growth rate and residues of the major microbial decomposer groups fungi and bacteria. Soil samples were collected at two depths (0-5 cm and 5-20 cm) from plots in an Irish winter wheat field that were exposed to either conventional or shallow non-inversion tillage for 7 growing seasons. Total soil fungal and bacterial biomasses were estimated using epifluorescence microscopy. To separate between biomass of saprophytic fungi and arbuscular mycorrhizae, samples were analyzed for ergosterol and phospholipid fatty acid (PLFA) biomarkers. Growth rates of saprophytic fungi were determined by [¹⁴C]acetate-in-ergosterol incorporation, whereas bacterial growth rates were determined by the incorporation of ³H-leucine in bacterial proteins. Finally, soil contents of fungal and bacterial residues were estimated by quantifying microbial derived amino sugars. Reduced tillage increased the total biomass of both bacteria and fungi in the 0-5 cm soil layer to a similar extent. Both ergosterol and PLFA analyses indicated that RT increased biomass of saprophytic fungi in the 0-5 cm soil layer. In contrast, RT increased the biomass of arbuscular mycorrhizae as well as its contribution to the total fungal biomass across the whole plough layer. Growth rates of both saprotrophic fungi and bacteria on the other hand were not affected by soil tillage, possibly indicating a decreased turnover rate of soil microbial biomass under RT. Moreover, RT did not affect the proportion of microbial residues that were derived from fungi. In summary, our results suggest that RT can promote soil C storage without increasing the role of saprophytic fungi in SOM dynamics relative to that of bacteria.     

Plant-N incorporation into microbial amino sugars as affected by inorganic N addition: A microcosm study of N-labeled maize residue decomposition

Carbon (C) and/or nitrogen (N) in plant residues can be assimilated into microbial biomass during the plant residue decomposition before incorporation into SOM in the form of microbial residues. Yet, microbial transformation of plant residue-N into microbial residues and the effects of inorganic N inputs on this process have not been well documented. Here, we undertook a 38-week incubation with a silt loam soil amended with a ¹⁵N-labeled maize (Zea mays L.) residue to determine how the transformation of maize residue-N into soil amino sugars was affected by rates of inorganic N addition. The newly metabolized amino sugars derived from maize residue-N were differentiated and quantified by using an isotope-based gas chromatography-mass spectrometry technique. We found that greater amounts of maize residue-N were transformed into amino sugars with lower inorganic N addition at the early stages of the plant residue degradation. However, the trend was reversed during later stages of decay as greater percentage of maize residue-N (8.6-9.4%) were enriched in amino sugars in the N_med and N_high soils, as compared with N₀ and N_low (7.5-8.2%). This indicated that higher availability of inorganic N could delay the transformation process of plant-N into microbial residues during the mineralization of plant residues. The dynamic transformations of the plant residue-N into individual amino sugars were compound-specific, with very fast incorporation into bacterial MurA_M-new found during the initial weeks, while the dynamics of maize residue-derived GluN exhibited a delayed response to assimilate plant-N into fungal products. The findings indicated differential contributions of maize residue decomposing microorganisms over time. Moreover, we found no preferential utilization of inorganic N over plant residue-N into amino sugars during the incubation course, but inorganic N inputs altered the rate of plant-N accumulation in microbial-derived organic matters. Our results indicated that higher N availability had a positive impact on the accumulation or stabilization of newly-produced microbial residues in the long term.     

Variation of intra‐aggregate organic carbon affects aggregate formation and stability during organic manure fertilization in a fluvo‐aquic soil

The aggregate formation and stability are controlled by the dynamics of soil organic matters (SOM), but how it is related to SOM chemical composition within different‐sized aggregates is largely unknown during manure fertilization. In this study, the variations of intra‐aggregate organic carbon (OC), including intra‐particulate organic matter (iPOM) and mineral‐associated organic matter, were quantitatively and qualitatively analysed, and then, their effects on aggregate formation and stability were assessed under four treatments: control (CK), mineral fertilizer (NPK), reduced manure (30%M) and manure fertilizers (M). Manure application (M) significantly increased macroaggregate proportion, mean weight diameter (MWD), and OC contents within different‐sized aggregates compared to CK, NPK, and 30%M. The OC accumulation of macroaggregate in M was attributed to OC content increase in silt plus clay subfraction rather than iPOM with more labile organic groups; oppositely, in microaggregate it was located in the relatively stable fine iPOM. The macroaggregate formation and stability were controlled by the fine iPOM within macroaggregates, whose abundant polysaccharide‐C and aliphatic‐C after manure fertilization advanced the microbial growth except for Gram‐positive bacteria, which further promoted macroaggregate formation and stability. The free silt plus clay fraction also affected macroaggregate formation and stability, and its polysaccharide‐C derived from microorganisms or decomposing SOM was positively associated with MWD and macroaggregate proportion. Because polysaccharide‐C can be easily associated with mineral particles, further improving micro‐ or macroaggregation. We conclude that continuous manure fertilization could increase labile SOM accumulation within aggregates and then facilitate microbial growth, which collectively are responsible for aggregate formation and stabilization.     

Dynamics of soil amino sugar pools during decomposition processes of corn residues as affected by inorganic N addition

Purpose  

Identifying the impact of inorganic-nitrogen (N) availability on soil amino sugar dynamics during corn (Zea mays L.) residue decomposition may advance our knowledge of microbial carbon (C) and N transformations and the factors controlling these processes in soils. Amino sugars are routinely used as microbial biomarkers to investigate C and N sequestration in microbial residues, and they are also involved in microbial-mediated soil organic matter (SOM) turnover. We conducted a 38-week incubation study using a Mollisol which was amended with corn residues and four levels of inorganic N (i.e., 0, 60.3, 167.2, and 701.9 mg N kg⁻¹ soil). The objective of this study was to examine the effects of inorganic-N availability on fungal and bacterial formation and stabilization of heterogeneous amino sugars during the corn residue decomposition in soil.