Mechanisms behind soil N dynamics following cover restoration in degraded land in subtropical China

Purpose

Nitrogen (N) is an important nutrient for re-vegetation during ecosystem restoration, but the effects of cover restoration on soil N transformations are not fully understood. This study was conducted to investigate N transformations in soils with different cover restoration ages in Eastern China.

Materials and methods

Soil samples were collected from four degraded and subsequently restored lands with restoration ages of 7, 17, 23, and 35 years along with an adjacent control of degraded land. A ¹⁵N tracing technique was used to quantify gross N transformation rates.

Results and discussion

Compared with degraded land, soil organic carbon (SOC) and total N (TN) increased by 1.60–3.97 and 2.49–5.36 times in restoration land. Cover restoration increased ammonium and nitrate immobilization, and dissimilatory nitrate reduction to ammonium (DNRA) by 0.56–0.96, 0.34–2.10, and 0.79–3.45 times, respectively, indicating that restoration was beneficial for N retention. There were positive correlations between SOC content and ammonium and nitrate immobilization and DNRA, indicating that the increase in soil N retention capacity may be ascribed to increasing SOC concentrations. The stimulating effect of SOC on ammonium immobilization was greater than its effect on organic N mineralization, so while SOC and TN increased, inorganic N supply did not increase. Autotrophic and heterotrophic nitrification increased with increasing SOC and TN concentrations. Notably, heterotrophic nitrification was an important source of NO₃^??N production, accounting for 47–67% of NO₃^??N production among all restoration ages.

Conclusions

The capacity of N retention was improved by cover restoration, leading to an increase in soil organic carbon and total N over time, but inorganic N supply capacity did not change with cover restoration age.

     

亚热带松林中强酸性土壤自养和异养硝化作用的研究

The occurrence of nitrification in some acidic forest soils is still a subject of debate. Identification of main nitrification pathways in acidic forest soils is still largely unknown. Acidic yellow soil (Oxisol) samples were selected to test whether nitrification can occur or not in acidic subtropical pine forest ecosystems. Relative contributions of autotrophs and heterotrophs to nitrification were studied by adding selective nitrification inhibitor nitrapyrin. Soil NH₄⁺-N concentrations decreased, but NO₃^--N concentrations increased significantly for the no-nitrapyrin control during the first week of incubation, indicating that nitrification did occur in the acidic subtropical soil. The calculated net nitrification rate was 0.49 mg N kg^-1 d^-1 for the no-nitrapyrin control during the first week of incubation. Nitrapyrin amendment resulted in a significant reduction of NO₃^--N concentration. Autotrophic nitrification rate averaged 0.28 mg N kg^-1 d^-1 and the heterotrophic nitrification rate was 0.21 mg N kg^-1 d^-1 in the first week. Ammonia-oxidizing bacteria (AOB) abundance increased slightly during incubation, but nitrapyrin amendment significantly decreased AOB amoA gene copy numbers by about 80%. However, the ammonia-oxidizing archaea (AOA) abundance showed significant increases only in the last 2 weeks of incubation and it was also decreased by nitrapyrin amendment. Our results indicated that nitrification did occur in the present acidic subtropical pine forest soil, and autotrophic nitrification was the main nitrification pathway. Both AOA and AOB were the active biotic agents responsible for autotrophic nitrification in the acidic subtropical pine forest soil.     

Effect of labile and recalcitrant carbon on heterotrophic nitrification in a subtropical forest soil

Carbon (C) is an important factor controlling heterotrophic nitrification in soil, but the effect of individual C components (e.g., labile and recalcitrant C) is largely unclear. We carried out a C amendment experiment in which either labile C (glucose) or a recalcitrant C (cellulose and biochar) was added to a subtropical forest soil. A ¹⁵N-, ¹³C-tracing and MiSeq sequencing study was performed to investigate soil gross heterotrophic nitrification rates, carbon utilization for soil respiration and microbial biomass production and microbial composition, respectively. After 2 days, results showed a significant increase of gross heterotrophic nitrification rate in glucose (GLU) (on average 3.34 mg N kg⁻¹ day⁻¹), cellulose (CEL) (on average 0.21 mg N kg⁻¹ day⁻¹) and biochar (BIO) (on average 0.13 mg N kg⁻¹ day⁻¹) amendment in comparison with the unamended soil (CK) (on average 0.01 mg N kg⁻¹ day⁻¹; p < 0.05). The contribution of heterotrophic nitrification to total soil nitrification was significantly larger in GLU (average 85.86%), CEL (average 98.52%) and BIO (average 81.25%) treatments compared with CK (average 33.33%; p < 0.01). After 2-month amendment, the gross rates remarkably decreased in GLU (average 0.02 mg N kg⁻¹ day⁻¹), and the contribution to total nitrification (average 8.73%) were significantly lower than that in CK (p < 0.05). A decrease in the proportion of heterotrophic nitrification to total nitrification in soil was also observed in CEL (average 38.40%) and BIO (6.74%) treatments. Nevertheless, BIO amendment (compared to CK, GLU and CEL) showed the highest gross heterotrophic nitrification rate, accompanied by a notably higher abundance of specific heterotrophic nitrifiers, i.e. Trichoderma, Aspergillus and Penicillium. These results point to a stimulatory effect of C addition on soil heterotrophic nitrification in the short term, while the stimulatory impact of C amendment diminishes with the decline in easily available C. In addition, a shift of the microbial composition in the long term can possibly be sustained for longer if additional recalcitrant C is available to heterotrophic nitrifiers. The dynamic response of heterotrophic nitrification to labile and recalcitrant C in this study offered an explanation for the positive effect of plantation and plant root exudation on the process.     

Effects of soil moisture on gross N transformations and N2O emission in acid subtropical forest soils

Soil moisture changes, arising from seasonal variation or from global climate changes, could influence soil nitrogen (N) transformation rates and N availability in unfertilized subtropical forests. A ¹⁵?N dilution study was carried out to investigate the effects of soil moisture change (30–90 % water-holding capacity (WHC)) on potential gross N transformation rates and N₂O and NO emissions in two contrasting (broad-leaved vs. coniferous) subtropical forest soils. Gross N mineralization rates were more sensitive to soil moisture change than gross NH₄ ⁺ immobilization rates for both forest soils. Gross nitrification rates gradually increased with increasing soil moisture in both forest soils. Thus, enhanced N availability at higher soil moisture values was attributed to increasing gross N mineralization and nitrification rates over the immobilization rate. The natural N enrichment in humid subtropical forest soils may partially be due to fast N mineralization and nitrification under relatively higher soil moisture. In broad-leaved forest soil, the high N₂O and NO emissions occurred at 30 % WHC, while the reverse was true in coniferous forest soil. Therefore, we propose that there are different mechanisms regulating N₂O and NO emissions between broad-leaved and coniferous forest soils. In coniferous forest soil, nitrification may be the primary process responsible for N₂O and NO emissions, while in broad-leaved forest soil, N₂O and NO emissions may originate from the denitrification process.     

硝化作用驱动下红壤渗漏液的酸化

土壤渗漏液pH对于亚热带酸性土壤的物质迁移和溶液中物质形态具有重要影响。为了研究亚热带酸性土壤硝化作用释放H+与渗漏液pH的关系,以具有不同硝化强度的3个红壤样本为供试材料,分别加入铵态氮0、150和300 mg kg-1,进行112 d的室内土柱模拟淋溶实验。结果表明:酸性土壤的渗漏液并不一定呈酸性。土壤渗漏液pH取决于硝化作用产生H+的速率与土壤酸缓冲能力的综合作用。当硝化作用使渗漏液中NO3-浓度升高至一定程度时,渗漏液pH突然下降,这一临界NO3-浓度与土壤盐基饱和度及加入土壤的铵态氮量呈线性正相关(p0.05)。所以,硝化作用最强的旱地土壤,由于其盐基饱和度达81%,渗漏液始终保持中性;而硝化作用不强、盐基饱和度为21%的灌丛土壤,其渗漏液pH可降至4.0以下。     

Effect of the inhibitors nitrapyrin and sodium chlorate on nitrification and N2O formation in an acid forest soil

An acid forest soil from beech forest gaps, which were either limed or unlimed, and the undisturbed forest was investigated for the type of nitrifying populations and the process of N₂O evolution. To see whether nitrifiers were of heterotrophic or autotrophic origin, the nitrification inhibitors nitrapyrin and sodium chlorate were applied to disturbed soil samples which underwent laboratory incubations. Nitrapyrin inhibits autotrophic nitrification. In different studies, sodium chlorate has been identified as an inhibitor either of autotrophic or of heterotrophic nitrification. In the samples investigated only nitrapyrin inhibited the autotrophic nitrification occurring in the limed soil. Sodium chlorate effectively inhibited heterotrophic nitrification. In the limed forest floor samples, where most autotrophic nitrification occured, sodium chlorate showed no inhibitory effect. In another laboratory incubation experiment, N₂O evolution from undisturbed soil columns, to which the above inhibitors were applied, was investigated. In those samples, in which nitrification had been reduced, neither inhibitor significantly reduced N₂O evolution. Thus it was concluded that the contribution of nitrification to N₂O losses is negligible, and that N₂O evolution arises from the activity of denitrifying organisms. Microbial biomass and respiration measurements showed that the inhibitors did not affect microflora negatively.     

pH 升高对红壤硝化过程产生 N2O 的影响

对红壤添加NaOH培养获得不同pH系列的土壤.通过室内培养试验.研究了3种pH条件下土壤的N_2O排放和无机N的变化情况.结果表明:硝化活性随土壤pH升高而增强:pH升高增加了土壤N_2O的释放;纯化学过程对N_2O散发的贡献随pH的升高而降低;Nitrapyrin在pH 4.8和pH 6.0时表现山硝化抑制作用,在pH 8.5时抑制效果不明显,且提高了培养期间pH8.5土壤N_2O的释放量.     

Nitrogen addition change soil N pools with litter removal or not in subtropical forest

ABSTRACT

There are many nitrogen (N) pools in soil, so their availability and different status can give information about bulk soil response to N deposition. However, the different size of N pools in forest soils and the relationship between them have not been well studied under N deposition when considering the role of litter. Here soil in an N-deposition experiment carried out for 5 years in a broad-leaved forest was used as an object to study the response of N pools to N deposition by stepwise extraction using water or solutions containing 0.5 M K₂SO₄, 2.5 M H₂SO₄ (LPI), or 13 M H₂SO₄ (LPII), and calculation of recalcitrant (RC) N pool. Under N control (CT), soil with the presence of litter had a higher N of 23.8–106.8% in the first four pools, but lower of 80.6% in recalcitrant N pool compared with soil with the absence of litter. In the absence of litter, N addition increased soil N in labile pool but decreased N in the RC pool compared to CT and these impacts were greater at high added N (HN) than low-added N (LN) rates. However, in the presence of litter, LN increased the amount of N in the K₂SO₄- extracted pool and HN reduced that in the water extracted pool. Additionally, LN and HN increased TN in the RC pool and HN increased the total soluble N (TSN) in the LPI and LPII pool. N changes in the water extraction pool were attributed to inorganic N, whereas they were NH₄ ⁺ and soluble organic N (SON) in the K₂SO₄-extracted, LPI, and LPII pools. In the presence of litter, HN increased the SON concentration in the K₂SO₄, LPI, and LPII extractions; thus, SON may be a potentially important N form for N availability. These results suggested that N additions improve the accumulation of N in RC pool with the presence of litter. The different effects of N additions on soil N pool or N form in each pool depend on litter present or not.     

Mechanisms of soil N dynamics following long-term application of organic fertilizers to subtropical rain-fed purple soil in China

In this study, a ¹⁵N tracing incubation experiment and an in situ monitoring study were combined to investigate the effects of different N fertilizer regimes on the mechanisms of soil N dynamics from a long-term repeated N application experiment. The field study was initiated in 2003 under a wheat-maize rotation system in the subtropical rain-fed purple soil region of China. The experiment included six fertilization treatments applied on an equivalent N basis (280 kg N ha⁻¹), except for the residue only treatment which received 112 kg N ha⁻¹: (1) UC, unfertilized control; (2) NPK, mineral fertilizer NPK; (3) OM, pig manure; (4) OM-NPK, pig manure (40% of applied N) with mineral NPK (60% of applied N); (5) RSD, crop straw; (6) RSD-NPK, crop straw (40% of applied N) with mineral NPK (60% of applied N). The results showed that long-term repeated applications of mineral or organic N fertilizer significantly stimulated soil gross N mineralization rates, which was associated with enhanced soil C and N contents following the application of N fertilizer. The crop N offtake and yield were positively correlated with gross mineralization. Gross autotrophic nitrification rates were enhanced by approximately 2.5-fold in the NPK, OM, OM-NPK, and RSD-NPK treatments, and to a lesser extent by RSD application, compared to the UC. A significant positive relationship between gross nitrification rates and cumulative N loss via interflow and runoff indicated that the mechanisms responsible for increasing N loss following long-term applications of N fertilizer were governed by the nitrification dynamics. Organic fertilizers stimulated gross ammonium (NH₄⁺) immobilization rates and caused a strong competition with nitrifiers for NH₄⁺, thus preventing a build-up of nitrate (NO₃⁻). Overall, in this study, we found that partial or complete substitution of NPK fertilizers with organic fertilizers can reduce N losses and maintain high crop production, except for the treatment involving application of RSD alone. Therefore, based on the N transformation dynamics observed in this study, organic fertilizers in combination with mineral fertilizer applications (i.e. OM, OM-NPK, and RSD-NPK treatments) are recommended for crop production in the subtropical rain-fed purple soils in China.     

改变碳输入对南亚热带海岸沙地典型天然次生林土壤呼吸的影响

土壤呼吸是陆地生态系统碳循环的一个重要过程,开展环境因子和改变碳输入对土壤呼吸影响的研究具有重要意义.2015年3月-2016年2月,在南亚热带海岸沙地典型天然次生林中设置去除根系、去除凋落物、加倍凋落物和对照4种处理,采用LI-8100连续观测改变碳输入对土壤呼吸的影响.结果表明:改变碳输入没有显著影响l0cm土壤温度和湿度(P＞0.05);不同处理土壤呼吸速率存在明显的季节变化,表现为夏高冬低,最大值出现在5月或者6月,最小值出现在11月或12月;土壤呼吸速率的年均值为加倍凋落物＞对照＞去除根系＞去除凋落物,不同改变碳输入方式均降低了土壤呼吸的Q10值;矿质土壤呼吸、凋落物呼吸和根系呼吸对土壤总呼吸的贡献分别为41.24％、43.29％和15.45％;不同处理土壤呼吸速率分别与土壤温度和湿度呈显著的指数和线性正相关(P＜0.05),双因子模型能解释土壤呼吸变异的45％ ～ 69％;改变碳输入影响土壤可溶性有机碳和微生物生物量碳,不同处理土壤呼吸速率与可溶性有机碳和微生物生物量碳呈正相关.因此,改变碳输入引起土壤易变碳的变化进而影响土壤呼吸.     

Numbers of autotrophic nitrifiers and nitrification in fertilized forest soil

The most probable number (MPN) method was used to estimate how numbers of autotrophic nitrifiers in Myrtillus-type and Calluna-type pine forest soils in southern Finland were affected by seven different fertilization treatments. No NH⁺₄ oxidizers and only a few hundred NO₂^?1 oxidizers g^?1 of soil were found in unfertilized organic (O) horizons. Ammonium nitrate and nitroform (ureaformaldehyde) had hardly any effect on the nitrifiers. Urea, alone or applied together with apatite + biotite or with apatite + biotite + micronutrients, increased numbers of NH₄⁺ and NO₂^? oxidizers. Wood ash, alone or with apatite, also had a stimulative effect. The effects of the stimulative fertilizers were less in the A₂ horizon than in the O horizon. The MPN counts were considerably affected by the duration of incubation: counts of NH₄⁺ oxidizers kept increasing for at least 8 weeks and counts of NO₂^? oxidizers for at least 15 weeks. These MPN counts were compared with earlier results from incubation experiments on the same soils to find out how they reflect changes in soil nitrification after fertilization.     

Simulated warming and low O2 promote N2O and N2 emissions in subtropical montane forest soil

Journal of Soils and Sediments - The montane subtropical forest soils contain huge nitrogen stocks, and climate warming might drive its volatilization due to the promotion of gaseous losses of...     

Microbial function in adjacent subtropical forest and agricultural soil

Soil microbial communities and their activities are altered by land use change; however impacts and extent of these alterations are often unclear. We investigated the functional responses of soil microbes in agricultural soil under sugarcane and corresponding native soil under Eucalyptus forest to additions of contrasting plant litter derived from soybean, sugarcane and Eucalyptus in a microcosm system, using a suite of complimentary techniques including enzyme assays and community level physiological profiles (CLPP). Initially agricultural soil had 50% less microbial biomass and lower enzyme activities than forest soil, but significantly higher nitrification rates. In response to litter addition, microbial biomass increased up to 11-fold in agricultural soil, but only 1.8-fold in forest soil, suggesting a prevalence of rapidly proliferating ‘r’ and slower growing ‘K’ strategists in the respective soils. Litter-driven change in microbial biomass and activities were short lived, largely returning to pre-litter addition levels by day 150. Decomposition rates of sugarcane and soybean litter as estimated via CO₂ production were lower in agricultural than in forest soil, but decomposition of more recalcitrant Eucalyptus litter was similar in both soils, contradicting the notion that microbial communities specialise in decomposing litter of the dominant local plant species. Enzyme activities and community level physiological profiles (CLPP) were closely correlated to microbial biomass and overall CO₂ production in the agricultural soil but not the forest soil, suggesting contrasting relationships between microbial population dynamics and activity in the two soils. Activities of enzymes that break down complex biopolymers, such as protease, cellulase and phenol oxidase were similar or higher in the agricultural soil, which suggests that the production of extracellular biopolymer-degrading enzymes was not a factor limiting litter decomposition. Enzyme and CLPP analyses produced contrasting profiles of microbial activity in the two soils; however the combination of both analyses offers additional insights into the changes in microbial function and community dynamics that occur after conversion of forest to agricultural land.     

Effects of decreasing water potential on gross ammonification and nitrification in an acid coniferous forest soil

Changes in the soil water regime, predicted as a consequence of global climate change, might influence the N cycle in temperate forest soils. We investigated the effect of decreasing soil water potentials on gross ammonification and nitrification in different soil horizons of a Norway spruce forest and tested the hypotheses that i) gross rates are more sensitive to desiccation in the Oa and EA horizon as compared to the uppermost Oi/Oe horizon and ii) that gross nitrification is more sensitive than gross ammonification. Soil samples were adjusted by air drying to water potentials from about field capacity to around −1.0 MPa, a range that is often observed under field conditions at our site. Gross rates were measured using the ¹⁵N pool dilution technique. To ensure that the addition of solute label to dry soils and the local rewetting does not affect the results by re-mineralization or preferential consumption of ¹⁵N, we compared different extraction and incubation times.T₀ times ranging from 10 to 300 min and incubation times of 48 h and 72 h did not influence the rates of gross ammonification and nitrification. Even small changes of water potential decreased gross ammonification and nitrification in the O horizon. In the EA horizon, gross nitrification was below detection limit and the response of the generally low rates of gross ammonification to decreasing water potentials was minor. In the Oi/Oe horizon gross ammonification and nitrification decreased from 37.5 to 18.3 mg N kg⁻¹ soil d⁻¹ and from 15.4 to 5.6 mg N kg⁻¹ soil d⁻¹ when the water potential decreased from field capacity to −0.8 MPa. In the Oa horizon gross ammonification decreased from 7.4 to 4.0 mg N kg⁻¹ soil d⁻¹ when the water potential reached −0.6 MPa. At such water potential nitrification almost ceased, while in the Oi/Oe horizon nitrification continued at a rather high level. Hence, only in the Oa horizon nitrification was more sensitive to desiccation than ammonification. Extended drought periods that might result from climate change will cause a reduction in gross N turnover rates in forest soils even at moderate levels of soil desiccation.     

Organic nitrogen stimulates the heterotrophic nitrification rate in an acidic forest soil

Many previous studies have demonstrated that heterotrophic nitrification processes play an important role in the production of NO₃⁻ in acidic soils. However, it is not clear whether a low concentration of nitrogenous organic compounds support heterotrophic nitrification processes in natural soils. In this study, we performed an ¹⁵N tracer experiment with a glycine concentration gradient (20, 40, 80, and 160 mg N kg⁻¹) to investigate the effect of the organic nitrogen concentration on the heterotrophic nitrification rate and its relative contribution to the total nitrification of the studied acidic forest soil. This experiment demonstrated that ¹⁵N–NO₃⁻ accumulated over time with all nitrogen treatments in the presence of acetylene, confirming that heterotrophic nitrification occurred even at a low organic nitrogen concentration (20 mg kg⁻¹) in the studied acidic forest soil. In the presence of acetylene, the ¹⁵N–NO₃⁻ concentration in the 20 and 40 mg kg⁻¹ glycine-N treatments was significantly lower than in the 80 and 160 mg kg⁻¹ glycine-N treatments (p < 0.05), indicating that a high organic nitrogen concentration stimulated the heterotrophic nitrification rate. There was no significant difference in the average contribution of heterotrophic nitrification to total nitrification among the different nitrogen treatments, suggesting that the organic nitrogen concentration did not affect the relative contribution of heterotrophic nitrification to total nitrification in the studied acidic soil. Our results confirmed that a low concentration of organic N (20 mg kg⁻¹) supported heterotrophic nitrification in the studied soil. The organic nitrogen concentration stimulates the heterotrophic nitrification rate, but does not affect the relative contribution of heterotrophic nitrification to total nitrification in the studied acidic soil.