Fast turnover of low molecular weight components of the dissolved organic carbon pool of temperate grassland field soils

Large amounts of low molecular weight (LMW;<250 Da) carbon (C) are lost from roots into the rhizosphere as a consequence of root turnover and exudation. Their rates of turnover after release into the soil remain poorly understood. We extracted soil solution from a temperate grassland Eutric Cambisol, isotopically labeled the glucose and amino acid components, and then re-injected the solution back into the soil. We followed the subsequent evolution of ¹⁴CO₂ and incorporation of the LMW C into the soil microbial biomass or grasses for 48 h. The experiments were performed both on grazed and un-grazed swards in the field, and in the laboratory. In the field, we showed that glucose and amino acids had short half-lives (t_1/2) in soil solution (t_1/2=20-40 min), but that they persisted in soil microbes for much longer. A first-order double exponential model fitted the experimental data well and gave rate constant (k) values of 1.21-2.14 h⁻¹ for k₁ and 0.0025-0.0048 h⁻¹ for k₂. Only small amounts of the added ¹⁴C were recovered in plant biomass (<5% of total added to soil) indicating that plant roots are poor competitors for LMW dissolved organic C (DOC) in comparison to soil microorganisms. The first phase of glucose and amino acid mineralization in the laboratory was slower (t_1/2=40-60 min) than measured in the field reinforcing the importance of making flux measurements in situ. Whilst grazing stimulated below-ground respiration, it exerted only a small influence on the turnover of LMW DOC suggesting that the increase in respiration was due to increased root respiration and not turnover of soil organic matter (SOM). Our results suggest that some components of the LMW DOC pool are turned over extremely rapidly (ca. 4000 times annually).     

Free amino sugar reactions in soil in relation to soil carbon and nitrogen cycling

Amino sugars represent a major constituent of microbial cell walls and hydrolyzed soil organic matter. Despite their potential importance in soil nitrogen cycling, comparatively little is known about their dynamics in soil. The aim of this study was therefore to quantify the behaviour of glucosamine in two contrasting grassland soil profiles. Our results show that both free amino sugars and amino acids represented only a small proportion of dissolved organic N and C pool in soil. Based upon our findings we hypothesize that the low concentrations of free amino sugars found in soils is due to rapid removal from the soil solution rather than slow rates of production. Further, we showed that glucosamine removal from solution was a predominantly biotic process and that its half-life in soil solution ranged from 1 to 3 h. The rates of turnover were similar to those of glucose at low substrate concentrations, however, at higher glucosamine concentrations its microbial use was much less than for glucose. We hypothesized that this was due to the lack of expression of a low affinity transport systems in the microbial community. Glucosamine was only weakly sorbed to the soil's solid phase (K_d=6.4±1.0) and our results suggest that this did not limit its bioavailability in soil. Here we showed that glucosamine addition to soil resulted in rapid N mineralization and subsequent NO₃⁻ production. In contrast to some previous reports, our results suggest that free amino sugars turn over rapidly in soil and provide a suitable substrate for both microbial respiration and new biomass formation.     

Amounts of carbon mineralised and leached as DOC during decomposition of Norway spruce needles and fine roots

Changes in climate or forest management practices leading to increased litter production will most likely cause increased leaching rates of dissolved organic carbon (DOC) from the O horizon. The rhizosphere is often assumed to have a large carbon flux associated with root turnover and exudation. However, little has been done to quantify the amount of DOC originating from root litter. We studied decomposition of fine root and needle litter of Norway spruce (Picea abies) through a combined incubation and leaching experiment in the laboratory using five different litter types: fresh needle litter, aged needles from the litter layer, fresh and dead roots from mineral soil samples, and seven-year-old roots from a previous litterbag study. After respiration measurements, the samples were percolated with artificial throughfall water and DOC and UV absorbance were measured in the leachate. Mineralisation of dissolved organic matter in the leachate and sorption of DOC to ferrihydrite were determined as a measure of DOC ability to be stabilised by iron (hydr)oxide surfaces.The mineralisation rate and DOC production rate of root samples were always lower than that of needle samples. However, root and needle derived dissolved organic matter (DOM) were similar in terms of aromaticity, as indicated by their specific UV absorbance, and ability to be sorbed by ferrihydrite. For seven-year-old roots, a significantly higher fraction of carbon was lost as DOC (30%) than for younger roots (20%). Furthermore, DOM from old roots bound more strongly to ferrihydrite and is mineralised at a lower rate than DOC from younger roots, suggesting that roots at late stages of decomposition, although a small fraction of total litter, significantly contribute to carbon build-up in mineral soils. The slower decomposition rate of roots compared with needles must be taken into account when modelling litter decomposition.     

Carbon-to-nitrogen ratio is a poor predictor of low molecular weight organic nitrogen mineralization in soil

Carbon-to-nitrogen ratio (C:N) has frequently been shown to be a good predictor of the speed of organic residue decomposition and N mineralization in soil. While this relationship appears to work well for complex organic materials (e.g. plant litter), its applicability to smaller organic substrates containing N remains unknown. Here we evaluated whether the intrinsic properties of amino acids and peptides could be used to predict their rate of microbial uptake and subsequent N mineralization. In an agricultural grassland soil we found that C:N, molecular weight, aromaticity and sulphur content provided poor indicators of amino acid bioavailabilityand subsequent NH₄⁺ release into soil. We therefore hypothesize that the position of amino acids along microbial biosynthetic pathways together with internal demand for individual amino acids rather than their C or N content is the primary determinant of N mineralization.     

城市不同功能区绿地土壤微生物量碳氮及溶解性碳氮分布特征及影响因素

为研究城市功能分区对绿地土壤活性碳氮的影响,选择合肥市不同功能区(老商业区、老工业区、居民服务区、近郊森林公园)绿地土壤为研究对象,分析了土壤微生物量碳氮(MBC、MBN)、溶解性碳氮(DOC、DON)、有机碳(SOC)、全氮(TN)以及土壤基本理化性质的差异及其相互关系。结果表明,功能区类型对土壤微生物量碳氮及溶解性碳氮影响显著(P0.05),且0~30 cm土层含量均为近郊森林公园高于城区内各功能区。土层厚度对MBC、DOC、DON含量影响显著,MBC、DON在各功能区中均随土层深度增加而递减,森林公园MBN、DOC均为10~20 cm含量最高。不同功能区MBC/MBN在6~14间波动,以此预测本研究范围内土壤微生物可能以真菌和放线菌为主,细菌较少。本研究未证实DOC与MBC、MBC与DON、DON与MBN间的相关性,MBC+DOC、MBN+DON比MBC、DOC、DON、MBN更能反应土壤活性碳氮库与土壤微生物量。总体来说,相比城区内各功能区,近郊森林公园更利于土壤生物量碳氮及溶解性有机碳氮的积累。     

Protein breakdown represents a major bottleneck in nitrogen cycling in grassland soils

Proteins represent the dominant input of organic N into most ecosystems and they also constitute the largest store of N in soil organic matter. The extracellular protease mediated breakdown of proteins to amino acids therefore represents a key step regulating N cycling in soil. In this study we investigated the influence of a range of environmental factors on the rate of protein mineralization in a grazed grassland and fallow agricultural soil. The protein turnover rates were directly compared to the rates of amino acid mineralization under the same conditions. Uniformly ¹⁴C-labelled soluble protein and amino acids were added to soil and the rate of ¹⁴CO₂ evolution determined over 30 d. Our results indicate that the primary phase of protein mineralization was approximately 20 ± 3 fold slower that the rate of amino acid mineralization. The addition of large amounts of inorganic NO₃⁻ and NH₄⁺ to the soil did not repress the rate of protein mineralization suggesting that available N does not directly affect protease activity in the short term. Whilst protein mineralization was strongly temperature sensitive, the presence of plants and the addition of humic and tannic acids had relatively little influence on the rate of soluble protein degradation in this fertile grassland soil. Our results suggests that the extracellular protease mediated cleavage of proteins to amino acids rather than breakdown of amino acids to NH₄⁺ represents the limiting step in soil N cycling.     

Dissolved organic nitrogen uptake by plants—an important N uptake pathway?

The direct uptake of dissolved organic nitrogen (DON) by plants has the potential to be a primary Factor in ecosystem functioning and vegetation succession particularly in N-limiting environments. Clear experimental evidence to support this view, however, is still lacking. Further, many of the experimental approaches used to assess whether DON is important may be compromised due to the use of inappropriate methods for comparing and quantifying plant available inorganic and organic soil N pools. In addition, experiments aimed at quantifying plant DON capture using dual-labelled (¹⁵N, ¹³C) organic N tracers often do not consider important aspects such as isotope pool dilution, differences in organic and inorganic N pool turnover times, bi-directional DON flows at the soil-root interface, and the differential fate of the ¹⁵N and ¹³C in the tracer compounds. Based upon experimental evidence, we hypothesize that DON uptake from the soil may not contribute largely to N acquisition by plants but may instead be primarily involved in the recapture of DON previously lost during root exudation. We conclude that while root uptake of amino acids in intact form has been shown, evidence demonstrating this as a major plant N acquisition pathway is still lacking.     

Seasonal fluctuation in microbial biomass and activity along a natural nitrogen gradient in a drained peatland

The effects of peat total N on the dissolved N and C concentrations and microbial biomass and activity and their range of seasonal fluctuation were studied in a drained peatland forest in Finland. Seasonal fluctuations in the concentrations of extractable dissolved organic (DON) and inorganic nitrogen (DIN) compounds and extractable dissolved organic carbon (DOC), microbial C and N, ergosterol, net and gross N mineralisation rates were investigated during two growing seasons along a natural peat N gradient in a drained peatland. Significant seasonal fluctuations in NH₄⁺ and DOC concentrations, microbial C and N, but not in ergosterol or microbial C-to-N ratios in the peat, were observed during the 1999 and 2000 growing seasons. The peat total N concentration affected extractable DON and DOC, but not DIN concentrations in the peat. A negative correlation was found between total N concentration in peat and microbial N and C, and a positive correlation between total N and ergosterol, in peat with N concentrations of up to 2%. Gross mineralisation rates did not show any correlation, whereas net mineralisation rates showed a significant positive correlation with the total N concentration of the peat in both 1999 and 2000.     

秸秆还田对稻田土壤溶液中溶解性有机质的影响

研究了麦秆不同方式还田、施用无机氮肥及不同移栽时间对水稻田土壤溶液中溶解性有机碳（DOC）、溶解性有机氮（DON）浓度的影响。结果表明,淹水后土壤溶液中DOC浓度在淹水初期明显增加,而后逐渐下降。添加麦秆这一有机物料,在水稻生长期前2个月内显著提高了DOC,对DON在一段时间内也表现出促进作用,其后差异不显著。施用无机氮肥降低了土壤溶液中DOC、DON浓度。DON的浓度与施肥量呈负相关。水稻推迟移栽,土壤溶液中溶解性有机质（DOM）均显著降低。     

Long-term N addition effects on the C mineralization and DOC production in mor humus under spruce

This study was based on laboratory incubations of mor humus from two N fertilized stands of Norway spruce in Sweden (Skogaby and Stråsan), which had received repeated N additions (100 kg N ha⁻¹ yr⁻¹ as (NH₄)₂SO₄ at Skogaby and 35, 73 and 108 kg ha⁻¹ yr⁻¹ as NH₄NO₃ at Stråsan) during 8 and 24-29 years, respectively. The aim was to investigate long-term N effects on the mineralization of C and production of DOC. Mor humus (Oe and Oa) was incubated in columns at 20 °C for 49 days. Columns were leached once a week with artificial throughfall solution, which was analyzed for DOC, total N, NH₄⁺-N and NO₃⁻-N. Prior to each leaching event, CO₂ evolution from the columns was determined. C-to-N ratios in the N-treated Oe layers at Stråsan (21-24) and Skogaby (24) were significantly lower than those of the controls (Stråsan, 32; Skogaby, 28). The cumulative amount of CO₂-C showed a significant treatment effect in the Oe layer at Skogaby, i.e. 18 and 29 mg C g⁻¹ C in the N treatment and control, respectively. At Stråsan, the cumulative CO₂-C was significantly lower in the N3 treatment compared to the control in both layers (33 compared to 74 mg C g⁻¹ C in the Oe layer and 16 compared to 35 mg C g⁻¹ C in the Oa layer). Neither the DOC nor the DON production showed any significant treatment effects at the two sites. However, DOC was lower in the fertilized Oe layer at Skogaby throughout the incubation. The leaching of DON was highest in the Oe layers at both sites, and DON increased with time at Skogaby while there was a decreasing trend at Stråsan. The DOC-to-DON ratio tended to be lower in the fertilized Oe layers at both sites. The NH₄⁺ leaching at Skogaby decreased in the N-treated Oe and Oa layers. At Stråsan, NH₄⁺ from the Oe layer increased in N2 and control. The NO₃⁻ leaching was low and constant in both Skogaby layers. At Stråsan, NO₃⁻ increased in the Oe layer of N1. Cumulative CO₂ was positively correlated to C-to-N ratio (r2=0.71,p<0.01) and to cumulative DOC (r2=0.63,p<0.05) in the Oe layer at Stråsan. Our conclusion was that long-term N additions caused decreased C-to-N ratios and decreased CO₂ evolution rates. The correlation between CO₂ and C-to-N ratio in the Oe layers at Stråsan may be due to a changed quality of the fertilized forest floor material and presence of more N efficient microorganisms.     

Plant capture of free amino acids is maximized under high soil amino acid concentrations

Free amino acids (AA's) represent a significant source of available N for some plants and soil microorganisms. It can be expected, however, that significant competition will exist between plants and microorganisms for this organic N resource. Our study indicated that microbial capture and utilization of glycine was very rapid at a range of soil solution concentrations (0.1 μM to 10 mM) indicating that significant competition will exist between roots and soil microorganisms. Plant capture of free AA's was maximal at high soil solution concentrations where microbial utilization was slowest. Our results suggest that plant capture of soil dissolved organic N may primarily occur in organic rich patches in soil where concentrations of free AA's are high.     

冻融作用对农田土壤可溶性氮组分的影响

为了解非生长季农田土壤氮素转化过程,采用室内冻融模拟培养试验研究了不同冻融温度和冻融循环次数对东北4种典型农田土壤(棕壤、褐土、草甸土、黑土)可溶性氮组分含量的影响。结果表明:随着冻结温度降低,4种农田土壤可溶性无机氮(DIN,NO_3~–-N+NH_4~+-N)、可溶性有机氮(DON)和可溶性全氮(DTN)含量均显著增加。随着融化温度升高,除NH_4~+-N含量显著升高外,4种农田土壤NO_3~–-N、DON和DTN含量的变化行为受冻结温度和土壤类型的协同影响。随着冻融循环次数增加,棕壤和褐土NO_3~–-N、NH_4~+-N、DON和DTN含量均显著增加;草甸土NO_3~–-N、DON和DTN含量均显著增加,而NH_4~+-N含量显著降低;黑土NO_3~–-N和NH_4~+-N含量均显著降低,而DON和DTN含量则先升高后降低。不同类型土壤受冻融作用影响的响应能力不同,其大小顺序为褐土棕壤、草甸土黑土。可见,冻融作用促进了土壤氮素转化,有利于土壤有效氮的累积,为春季作物生长提供足够的氮素,但同时也增加了土壤氮素流失风险。     

The carbon we do not see—the impact of low molecular weight compounds on carbon dynamics and respiration in forest soils: a review

Dissolved organic matter (DOM), typically quantified as dissolved organic carbon (DOC), has been hypothesized to play many roles in pedogenesis and soil biogeochemical cycles, however, most research to date concerning forest soils has focussed on the high molecular weight (HMW) components of this DOM. This review aims to assess the role of low molecular weight (LMW) DOM compounds in the C dynamics of temperate and boreal forest soils focussing in particular on organic acids, amino acids and sugars. The current knowledge of concentrations, mineralization kinetics and production rates and sources in soil are summarised. We conclude that although these LMW compounds are typically maintained at very low concentrations in the soil solution (<50 μM), the flux through this pool is extremely rapid (mean residence time 1-10 h) due to continued microbial removal. Due to this rapid flux through the soil solution pool and mineralization to CO₂, we calculate that the turnover of these LMW compounds may contribute substantially to the total CO₂ efflux from the soil. Moreover, the production rates of these soluble transitory compounds could exceed HMW DOM production. The possible impact of climate change on the behaviour of LMW compounds in soil is also discussed.     

坡面水文过程与土壤有机碳迁移研究进展

选取中国四种典型区域（东北黑土区、南方红壤丘陵区、北方黄土高原区、西南岩溶区）,概述了不同类型区域坡面水文过程与土壤有机碳迁移的关系,分析了坡面水文过程驱动下土壤有机碳迁移的影响因素。结果表明,不同类型区域的土壤有机碳迁移路径及形式因其水文过程不同而存在差异性,土壤类型、降雨强度、坡度等自然因素会影响土壤有机碳迁移路径及在泥沙中富集;而耕作、施肥、土地利用等人为因素既会改变土壤有机碳含量和外源输入而影响其迁移量,也会影响土壤有机碳迁移路径。在对前人研究成果的总结的基础上,本文认为应加强红壤区及岩溶区不同土壤有机碳迁移路径的同步监测研究,明晰水力侵蚀作用下土壤有机碳迁移转化过程与微生物群落之间的耦合关系,建立考虑多维度、多界面水文过程驱动下土壤有机碳迁移模型。     

pH regulation of carbon and nitrogen dynamics in two agricultural soils

Soil pH is often hypothesized to be a major factor regulating organic matter turnover and inorganic nitrogen production in agricultural soils. The aim of this study was to critically test the relationship between soil pH and rates of C and N cycling, and dissolved organic nitrogen (DON), in two long-term field experiments in which pH had been manipulated (Rothamsted silty clay loam, pH 3.5-6.8; Woburn sandy loam, pH 3.4-6.3). While alteration of pH for 37 years significantly affected crop production, it had no significant effect on total soil C and N or indigenous mineral N levels. This implies that at steady state, increased organic matter inputs to the soil are balanced by increased outputs of CO₂. This is supported by the positive correlation between both plant productivity and intrinsic microbial respiration with soil pH. In addition, soil microbial biomass C and N, and nitrification were also significantly positively correlated with soil pH. Measurements of respiration following addition of urea and amino acids showed a significant decline in CO₂ evolution with increasing soil acidity, whilst glucose mineralization showed no response to pH. In conclusion, it appears that changes in soil pH significantly affect soil microbial activity and the rate of soil C and N cycling. The evidence suggests that this response is partially indirect, being primarily linked to pH induced changes in net primary production and the availability of substrates. In addition, enhanced soil acidity may also act directly on the functioning of the microbial community itself.