Resolving the influence of lignin on soil organic matter decomposition with mechanistic models and continental-scale data

Confidence in model estimates of soil CO₂ flux depends on assumptions regarding fundamental mechanisms that control the decomposition of litter and soil organic carbon (SOC). Multiple hypotheses have been proposed to explain the role of lignin, an abundant and complex biopolymer that may limit decomposition. We tested competing mechanisms using data-model fusion with modified versions of the CN-SIM model and a 571-day laboratory incubation dataset where decomposition of litter, lignin, and SOC was measured across 80 soil samples from the National Ecological Observatory Network. We found that lignin decomposition consistently decreased over time in 65 samples, whereas in the other 15 samples, lignin decomposition subsequently increased. These “lagged-peak” samples can be predicted by low soil pH, high extractable Mn, and fungal community composition as measured by ITS PC2 (the second principal component of an ordination of fungal ITS amplicon sequences). The highest-performing model incorporated soil biogeochemical factors and daily dynamics of substrate availability (labile bulk litter:lignin) that jointly represented two hypotheses (C substrate limitation and co-metabolism) previously thought to influence lignin decomposition. In contrast, models representing either hypothesis alone were biased and underestimated cumulative decomposition. Our findings reconcile competing hypotheses of lignin decomposition and suggest the need to precisely represent the role of lignin and consider soil metal and fungal characteristics to accurately estimate decomposition in Earth-system models.     

The turnover of carbon pools contributing to soil CO2 and soil respiration in a temperate forest exposed to elevated CO2 concentration

Soil carbon is returned to the atmosphere through the process of soil respiration, which represents one of the largest fluxes in the terrestrial C cycle. The effects of climate change on the components of soil respiration can affect the sink or source capacity of ecosystems for atmospheric carbon, but no current techniques can unambiguously separate soil respiration into its components. Long‐term free air CO₂ enrichment (FACE) experiments provide a unique opportunity to study soil C dynamics because the CO₂ used for fumigation has a distinct isotopic signature and serves as a continuous label at the ecosystem level. We used the ¹³C tracer at the Duke Forest FACE site to follow the disappearance of C fixed before fumigation began in 1996 (pretreatment C) from soil CO₂ and soil‐respired CO₂, as an index of belowground C dynamics during the first 8 years of the experiment. The decay of pretreatment C as detected in the isotopic composition of soil‐respired CO₂ and soil CO₂ at 15, 30, 70, and 200 cm soil depth was best described by a model having one to three exponential pools within the soil system. The majority of soil‐respired CO₂ (71%) originated in soil C pools with a turnover time of about 35 days. About 55%, 50%, and 68% of soil CO₂ at 15, 30, and 70 cm, respectively, originated in soil pools with turnover times of less than 1 year. The rest of soil CO₂ and soil‐respired CO₂ originated in soil pools that turn over at decadal time scales. Our results suggest that a large fraction of the C returned to the atmosphere through soil respiration results from dynamic soil C pools that cannot be easily detected in traditionally defined soil organic matter standing stocks. Fast oxidation of labile C substrates may prevent increases in soil C accumulation in forests exposed to elevated [CO₂] and may consequently result in shorter ecosystem C residence times.     

Estimating soil carbon sequestration under elevated CO2 by combining carbon isotope labelling with soil carbon cycle modelling

Elevated CO₂ concentrations generally stimulate grassland productivity, but herbaceous plants have only a limited capacity to sequester extra carbon (C) in biomass. However, increased primary productivity under elevated CO₂ could result in increased transfer of C into soils where it could be stored for prolonged periods and exercise a negative feedback on the rise in atmospheric CO₂. Measuring soil C sequestration directly is notoriously difficult for a number of methodological reasons. Here, we present a method that combines C isotope labelling with soil C cycle modelling to partition net soil sequestration into changes in new C fixed over the experimental duration (C_new) and pre‐experimental C (C_old). This partitioning is advantageous because the C_new accumulates whereas C_old is lost in the course of time (ΔC_new>0 whereas ΔC_old<0). We applied this method to calcareous grassland exposed to 600 μL CO₂ L^?1 for 6 years. The CO₂ used for atmospheric enrichment was depleted in ¹³C relative to the background atmosphere, and this distinct isotopic signature was used to quantify net soil C_new fluxes under elevated CO₂. Using ¹³C/¹²C mass balance and inverse modelling, the Rothamsted model ‘RothC’ predicted gross soil C_new inputs under elevated CO₂ and the decomposition of C_old. The modelled soil C pools and fluxes were in good agreement with experimental data. C isotope data indicated a net sequestration of ≈90 g C_new m^?2 yr^?1 in elevated CO₂. Accounting for C_old‐losses, this figure was reduced to ≈30 g C m^?2 yr^?1 at elevated CO₂; the elevated CO₂‐effect on net C sequestration was in the range of≈10 g C m^?2 yr^?1. A sensitivity and error analysis suggests that the modelled data are relatively robust. However, elevated CO₂‐specific mechanisms may necessitate a separate parameterization at ambient and elevated CO₂; these include increased soil moisture due to reduced leaf conductance, soil disaggregation as a consequence of increased soil moisture, and priming effects. These effects could accelerate decomposition of C_old in elevated CO₂ so that the CO₂ enrichment effect may be zero or even negative. Overall, our findings suggest that the C sequestration potential of this grassland under elevated CO₂ is rather limited.     

Plant diversity positively affects short‐term soil carbon storage in experimental grasslands

Increasing atmospheric CO₂ concentration and related climate change have stimulated much interest in the potential of soils to sequester carbon. In ‘The Jena Experiment’, a managed grassland experiment on a former agricultural field, we investigated the link between plant diversity and soil carbon storage. The biodiversity gradient ranged from one to 60 species belonging to four functional groups. Stratified soil samples were taken to 30 cm depth from 86 plots in 2002, 2004 and 2006, and organic carbon contents were determined. Soil organic carbon stocks in 0–30 cm decreased from 7.3 kg C m^?2 in 2002 to 6.9 kg C m^?2 in 2004, but had recovered to 7.8 kg C m^?2 by 2006. During the first 2 years, carbon storage was limited to the top 5 cm of soil while below 10 cm depth, carbon was lost probably as short‐term effect of the land use change. After 4 years, carbon stocks significantly increased within the top 20 cm. More importantly, carbon storage significantly increased with sown species richness (log‐transformed) in all depth segments and even carbon losses were significantly smaller with higher species richness. Although increasing species diversity increased root biomass production, statistical analyses revealed that species diversity per se was more important than biomass production for changes in soil carbon. Below 20 cm depth, the presence of one functional group, tall herbs, significantly reduced carbon losses in the beginning of the experiment. Our analysis indicates that plant species richness and certain plant functional traits accelerate the build‐up of new carbon pools within 4 years. Additionally, higher plant diversity mitigated soil carbon losses in deeper horizons. This suggests that higher biodiversity might lead to higher soil carbon sequestration in the long‐term and therefore the conservation of biodiversity might play a role in greenhouse gas mitigation.     

Different types of nitrogen deposition show variable effects on the soil carbon cycle process of temperate forests

Nitrogen (N) deposition significantly affects the soil carbon (C) cycle process of forests. However, the influence of different types of N on it still remained unclear. In this work, ammonium nitrate was selected as an inorganic N (IN) source, while urea and glycine were chosen as organic N (ON) sources. Different ratios of IN to ON (1 : 4, 2 : 3, 3 : 2, 4 : 1, and 5 : 0) were mixed with equal total amounts and then used to fertilize temperate forest soils for 2 years. Results showed that IN deposition inhibited soil C cycle processes, such as soil respiration, soil organic C decomposition, and enzymatic activities, and induced the accumulation of recalcitrant organic C. By contrast, ON deposition promoted these processes. Addition of ON also resulted in accelerated transformation of recalcitrant compounds into labile compounds and increased CO₂ efflux. Meanwhile, greater ON deposition may convert C sequestration in forest soils into C source. These results indicated the importance of the IN to ON ratio in controlling the soil C cycle, which can consequently change the ecological effect of N deposition.     

An oxygen-mediated positive feedback between elevated carbon dioxide and soil organic matter decomposition in a simulated anaerobic wetland

We examined the effects of elevated atmospheric CO₂ on soil carbon decomposition in an experimental anaerobic wetland system. Pots containing either bare C₄‐derived soil or the C₃ sedge Scirpus olneyi planted in C₄‐derived soil were incubated in greenhouse chambers at either ambient or twice‐ambient atmospheric CO₂. We measured CO₂ flux from each pot, quantified soil organic matter (SOM) mineralization using δ¹³C, and determined root and shoot biomass. SOM mineralization increased in response to elevated CO₂ by 83–218% (P<0.0001). In addition, soil redox potential was significantly and positively correlated with root biomass (P= 0.003). Our results (1) show that there is a positive feedback between elevated atmospheric CO₂ concentrations and wetland SOM decomposition and (2) suggest that this process is mediated by the release of oxygen from the roots of wetland plants. Because this feedback may occur in any wetland system, including peatlands, these results suggest a limitation on the size of the carbon sink presented by anaerobic wetland soils in a future elevated‐CO₂ atmosphere.     

蒙古栎叶片及其土壤碳、氮同位素自然丰度对大气CO2浓度升高的响应

大气CO₂浓度升高对土壤氮素转化过程产生重要影响,研究其变化有助于更好地预测陆地生态系统的固碳潜力.氮同位素自然丰度作为生态系统氮素循环过程的综合指标能够有效地指示CO₂浓度升高对土壤氮素转化过程的影响.本研究采用开顶箱CO₂ 熏蒸法研究连续10年的大气CO₂ 浓度升高对我国东北地区蒙古栎及其土壤和微生物生物量碳、氮同位素自然丰度的影响.结果表明: 大气CO₂浓度升高改变了土壤氮循环过程,增加了土壤微生物和植物叶片δ¹⁵N;促进了富¹³C土壤有机碳分解,中和了贫¹³C植物光合碳输入的效果,导致土壤可溶性有机碳和微生物碳δ¹³C在CO₂升高条件下没有发生显著变化.这些结果表明,CO₂浓度升高很可能促进了土壤有机质矿化过程,并加剧了系统氮限制的状态.     

土壤有机碳和氮分解对温度变化的响应趋势与研究方法

总结了土壤中碳和氮贮量与温度的关系、土壤碳和氮分解对温度时空差异和直接加热升温的响应,以及土壤碳和氮分解对低温冻结及冻融循环的响应趋势,讨论了其研究方法的误差和不确定性,并对今后的研究提出了一些建议．气候变暖在短期内将使土壤碳和氮分解加速并引起CO2释放量增加,而长期过程中却并不一定会引起土壤碳和氮分解加速．合理解释不同研究结果的差异,除了需要系统分析土壤碳和氮分解对温度变化响应的机制外,还需要充分认识土壤碳和氮分解对温度变化响应的长期过程和短期过程的差异,以及研究方法、植被、土壤和气候等因素的影响．     

Increased soil carbon storage through plant diversity strengthens with time and extends into the subsoil

Soils are important for ecosystem functioning and service provisioning. Soil communities and their functions, in turn, are strongly promoted by plant diversity, and such positive effects strengthen with time. However, plant diversity effects on soil organic matter have mostly been investigated in the topsoil, and there are only very few long-term studies. Thus, it remains unclear if plant diversity effects strengthen with time and to which depth these effects extend. Here, we repeatedly sampled soil to 1 m depth in a long-term grassland biodiversity experiment. We investigated how plant diversity impacted soil organic carbon and nitrogen concentrations and stocks and their stable isotopes ¹³C and ¹⁵N, as well as how these effects changed after 5, 10, and 14 years. We found that higher plant diversity increased carbon and nitrogen storage in the topsoil since the establishment of the experiment. Stable isotopes revealed that these increases were associated with new plant-derived inputs, resulting in less processed and less decomposed soil organic matter. In subsoils, mainly the presence of specific plant functional groups drove organic matter dynamics. For example, the presence of deep-rooting tall herbs decreased carbon concentrations, most probably through stimulating soil organic matter decomposition. Moreover, plant diversity effects on soil organic matter became stronger in topsoil over time and reached subsoil layers, while the effects of specific plant functional groups in subsoil progressively diminished over time. Our results indicate that after changing the soil system the pathways of organic matter transfer to the subsoil need time to establish. In our grassland system, organic matter storage in subsoils was driven by the redistribution of already stored soil organic matter from the topsoil to deeper soil layers, for example, via bioturbation or dissolved organic matter. Therefore, managing plant diversity may, thus, have significant implications for subsoil carbon storage and other critical ecosystem services.     

Accelerated belowground C cycling in a managed agriforest ecosystem exposed to elevated carbon dioxide concentrations

We investigated the effects of three elevated atmospheric CO₂ levels on a Populus deltoides plantation at Biosphere 2 Laboratory in Oracle Arizona. Stable isotopes of carbon have been used as tracers to separate the carbon present before the CO₂ treatments started (old C), from that fixed after CO₂ treatments began (new C). Tree growth at elevated [CO₂] increased inputs to soil organic matter (SOM) by increasing the production of fine roots and accelerating the rate of root C turnover. However, soil carbon content decreased as [CO₂] in the atmosphere increased and inputs of new C were not found in SOM. Consequently, the rates of soil respiration increased by 141% and 176% in the 800 and 1200 μL L^?1 plantations, respectively, when compared with ambient [CO₂] after 4 years of exposure. However, the increase in decomposition of old SOM (i.e. already present when CO₂ treatments began) accounted for 72% and 69% of the increase in soil respiration seen under elevated [CO₂]. This resulted in a net loss of soil C at a rate that was between 10 and 20 times faster at elevated [CO₂] than at ambient conditions. The inability to retain new and old C in the soil may stem from the lack of stabilization of SOM, allowing for its rapid decomposition by soil heterotrophs.     

Influence of rhizodeposition under elevated CO2 on plant nutrition and soil organic matter

Atmospheric CO₂ concentrations can influence ecosystem carbon storage through net primary production (NPP), soil carbon storage, or both. In assessing the potential for carbon storage in terrestrial ecosystems under elevated CO₂, both NPP and processing of soil organic matter (SOM), as well as the multiple links between them, must be examined. Within this context, both the quantity and quality of carbon flux from roots to soil are important, since roots produce specialized compounds that enhance nutrient acquisition (affecting NPP), and since the flux of organic compounds from roots to soil fuels soil microbial activity (affecting processing of SOM).From the perspective of root physiology, a technique is described which uses genetically engineered bacteria to detect the distribution and amount of flux of particular compounds from single roots to non-sterile soils. Other experiments from several labs are noted which explore effects of elevated CO₂ on root acid phosphatase, phosphomonoesterase, and citrate production, all associated with phosphorus nutrition. From a soil perspective, effects of elevated CO₂ on the processing of SOM developed under a C4 grassland but planted with C3 California grassland species were examined under low (unamended) and high (amended with 20 g m^–2 NPK) nutrients; measurements of soil atmosphere 13C combined with soil respiration rates show that during vegetative growth in February, elevated CO₂ decreased respiration of carbon from C4 SOM in high nutrient soils but not in unamended soils.This emphasis on the impacts of carbon loss from roots on both NPP and SOM processing will be essential to understanding terrestrial ecosystem carbon storage under changing atmospheric CO₂ concentrations.Abbreviations SOM soil organic matter - NPP net primary productivity - NEP net ecosystem productivity - PNPP p-nitrophenyl phosphate     

Integrating plant litter quality,soil organic matter stabilization,and the carbon saturation concept

Labile, ‘high‐quality’, plant litters are hypothesized to promote soil organic matter (SOM) stabilization in mineral soil fractions that are physicochemically protected from rapid mineralization. However, the effect of litter quality on SOM stabilization is inconsistent. High‐quality litters, characterized by high N concentrations, low C/N ratios, and low phenol/lignin concentrations, are not consistently stabilized in SOM with greater efficiency than ‘low‐quality’ litters characterized by low N concentrations, high C/N ratios, and high phenol/lignin concentrations. Here, we attempt to resolve these inconsistent results by developing a new conceptual model that links litter quality to the soil C saturation concept. Our model builds on the Microbial Efficiency‐Matrix Stabilization framework (Cotrufo et al., 2013) by suggesting the effect of litter quality on SOM stabilization is modulated by the extent of soil C saturation such that high‐quality litters are not always stabilized in SOM with greater efficiency than low‐quality litters.     

Elevated CO2 evokes quantitative and qualitative changes in carbon dynamics in a plant/soil system: mechanisms and implications

It is hypothesized that carbon storage in soil will increase under an elevated atmospheric CO₂ concentration due to a combination of an increased net CO₂ uptake, a shift in carbon allocation pattern in the plant/soil system and a decreased decomposition rate of plant residues. An overview of several studies, performed in our laboratory, on the effects of elevated CO₂ on net carbon uptake, allocation to the soil and decomposition of roots is given to test this hypothesis. The studies included wheat, ryegrass and Douglas-fir and comprised both short-term and long-term studies.Total dry weight of the plants increased up to 62%, but depended on nutrient availability. These results were supported by the data on net ¹⁴CO₂ uptake. A shift in ¹⁴C-carbon distribution from shoots to roots was found in perennial species, although this depended on nutrient availability.The decomposition experiments showed that roots cultivated at 700 L L^–1 CO₂ were decomposed more slowly than those cultivated at 350 L L^–1 CO₂. Even after two growing seasons differences up to 13% were observed, although this was found to be dependent on the nitrogen level at which the roots were grown.Both an increased carbon allocation to the soil due to an increased carbon uptake, whether or not combined with a shift in distribution pattern, and a decreased decomposition of root residues will enhance the possibilities of carbon sequestration in soil, thus supporting our hypothesis. However, nutrient availability and the response of the soil microbial biomass (size and activity) play a major role in the processes involved and require attention to clarify plant/soil responses in the long term with regard to sustained stimulation of carbon input into soils and the decomposability of roots and rhizodeposition. Soil texture will also have a strong effect on decomposition rates as a result of differences in the protecting capacity for organic matter. More detailed information on these changes is needed for a proper use of models simulating soil carbon dynamics in the long term.     

Uncertainty in soil carbon accounting due to unrecognized soil erosion

The movement of soil organic carbon (SOC) during erosion and deposition events represents a major perturbation to the terrestrial carbon cycle. Despite the recognized impact soil redistribution can have on the carbon cycle, few major carbon accounting models currently allow for soil mass flux. Here, we modified a commonly used SOC model to include a soil redistribution term and then applied it to scenarios which explore the implications of unrecognized erosion and deposition for SOC accounting. We show that models that assume a static landscape may be calibrated incorrectly as erosion of SOC is hidden within the decay constants. This implicit inclusion of erosion then limits the predictive capacity of these models when applied to sites with different soil redistribution histories. Decay constants were found to be 15–50% slower when an erosion rate of 15 t soil ha^?1 yr^?1 was explicitly included in the SOC model calibration. Static models cannot account for SOC change resulting from agricultural management practices focused on reducing erosion rates. Without accounting for soil redistribution, a soil sampling scheme which uses a fixed depth to support model development can create large errors in actual and relative changes in SOC stocks. When modest levels of erosion were ignored, the combined uncertainty in carbon sequestration rates was 0.3–1.0 t CO₂ ha^?1 yr^?1. This range is similar to expected sequestration rates for many management options aimed at increasing SOC levels. It is evident from these analyses that explicit recognition of soil redistribution is critical to the success of a carbon monitoring or trading scheme which seeks to credit agricultural activities.     

Hierarchical saturation of soil carbon pools near a natural CO2 spring

Soil has been identified as a possible carbon (C) sink to mitigate increasing atmospheric CO₂ concentration. However, several recent studies have suggested that the potential of soil to sequester C is limited and that soil may become saturated with C under increasing CO₂ levels. To test this concept of soil C saturation, we studied a gley and organic soil at a grassland site near a natural CO₂ spring. Total and aggregate‐associated soil organic C (SOC) concentration showed a significant increase with atmospheric CO₂ concentration. An asymptotic function showed a better fit of SOC and aggregation with CO₂ level than a linear model. There was a shift in allocation of total C from smaller size fractions to the largest aggregate fraction with increasing CO₂ concentration. Litter inputs appeared to be positively related to CO₂ concentration. Based on modeled function parameters and the observed shift in the allocation of the soil C from small to large aggregate‐size classes, we postulate that there is a hierarchy in C saturation across different SOC pools. We conclude that the asymptotic response of SOC concentration at higher CO₂ levels indicates saturation of soil C pools, likely because of a limit to physical protection of SOC.     

鼎湖山亚热带常绿针阔叶混交林凋落物及矿质氮输入对土壤有机碳分解的影响

2010年7-12月,选取鼎湖山国家级自然保护区亚热带针阔叶混交林,采用全因子控制试验,研究不同类型的凋落物(针叶和阔叶凋落物)添加及氮处理(加氮模拟氮饱和、减氮模拟根吸收)对表层(0～10 cm)和下层(20～30 cm)土壤有机质分解(呼吸)的影响.结果表明： 2010年7-11月间,两种凋落物的添加使土壤-凋落物系统的呼吸速率显著增加,但这种影响在12月消失.减氮和加氮处理均显著增加了土壤-凋落物系统的呼吸.叶凋落物短期内完全分解,对土壤碳分解和积累的影响十分有限,可能不是该系统中土壤有机质的主要来源.通过减少土壤可利用氮模拟根系对氮的吸收能够明显促进土壤有机质的分解.     

黄土区不同退耕方式下土壤碳氮的差异及其影响因素

研究植被恢复对土壤碳氮动态的影响,对了解陆地生态系统碳氮循环,应对全球温室效应具有重要意义.本研究以黄土丘陵区不同人工恢复植被为对象,以农田为参照,分析了不同人工植被恢复方式对0～100 cm剖面土壤有机碳(SOC)和全氮(TN)含量影响的差异及其影响因素.结果表明: 退耕还林还草显著提高了土壤的SOC和TN含量.退耕后,SOC和TN含量均较农田明显提高.0～100 cm土层SOC平均含量人工乔木林为农田的1.43倍,增幅最大;其次是人工灌木,为1.36倍;最后是人工草地,为1.21倍.0～100 cm土层TN平均含量人工乔木林增幅最大,是农田的1.30倍;其次是人工草地,为1.21倍;而人工灌木增幅最小,为1.13倍.与农田相比,人工恢复植被类型间SOC和TN含量及细根密度的差异在土壤剖面深度上表现出不同,人工乔木和灌木最明显,影响深度＞100 cm;草地最小,仅为60 cm.恢复植被的细根密度、C∶N和凋落物量显著高于农作物,细根密度与SOC、TN呈显著线性相关(P<0.01).细根的质和量以及凋落物量是不同人工恢复植被下SOC和TN含量差异的重要影响因素.