Individual and interactive effects of warming and nitrogen supply on CO2 fluxes and carbon allocation in subarctic grassland

Climate warming has been suggested to impact high latitude grasslands severely, potentially causing considerable carbon (C) losses from soil. Warming can also stimulate nitrogen (N) turnover, but it is largely unclear whether and how altered N availability impacts belowground C dynamics. Even less is known about the individual and interactive effects of warming and N availability on the fate of recently photosynthesized C in soil. On a 10-year geothermal warming gradient in Iceland, we studied the effects of soil warming and N addition on CO₂ fluxes and the fate of recently photosynthesized C through CO₂ flux measurements and a ¹³CO₂ pulse-labeling experiment. Under warming, ecosystem respiration exceeded maximum gross primary productivity, causing increased net CO₂ emissions. N addition treatments revealed that, surprisingly, the plants in the warmed soil were N limited, which constrained primary productivity and decreased recently assimilated C in shoots and roots. In soil, microbes were increasingly C limited under warming and increased microbial uptake of recent C. Soil respiration was increased by warming and was fueled by increased belowground inputs and turnover of recently photosynthesized C. Our findings suggest that a decade of warming seemed to have induced a N limitation in plants and a C limitation by soil microbes. This caused a decrease in net ecosystem CO₂ uptake and accelerated the respiratory release of photosynthesized C, which decreased the C sequestration potential of the grassland. Our study highlights the importance of belowground C allocation and C-N interactions in the C dynamics of subarctic ecosystems in a warmer world.     

Land Use Alters the Drought Responses of Productivity and CO<Subscript>2</Subscript> Fluxes in Mountain Grassland

Climate extremes and land-use changes can have major impacts on the carbon cycle of ecosystems. Their combined effects have rarely been tested. We studied whether and how the abandonment of traditionally managed mountain grassland changes the resilience of carbon dynamics to drought. In an in situ common garden experiment located in a subalpine meadow in the Austrian Central Alps, we exposed intact ecosystem monoliths from a managed and an abandoned mountain grassland to an experimental early-summer drought and measured the responses of gross primary productivity, ecosystem respiration, phytomass and its components, and of leaf area index during the drought and the subsequent recovery period. Across all these parameters, the managed grassland was more strongly affected by drought and recovered faster than the abandoned grassland. A bivariate representation of resilience confirmed an inverse relationship of resistance and recovery; thus, low resistance was related to high recovery from drought and vice versa. In consequence, the overall perturbation of the carbon cycle caused by drought was larger in the managed than the abandoned grassland. The faster recovery of carbon dynamics from drought in the managed grassland was associated with a significantly higher uptake of nitrogen from soil. Furthermore, in both grasslands leaf nitrogen concentrations were enhanced after drought and likely reflected drought-induced increases in nitrogen availability. Our study shows that ongoing and future land-use changes have the potential to profoundly alter the impacts of climate extremes on grassland carbon dynamics.     

Global change belowground: impacts of elevated CO2, nitrogen,and summer drought on soil food webs and biodiversity

The world's ecosystems are subjected to various anthropogenic global change agents, such as enrichment of atmospheric CO₂ concentrations, nitrogen (N) deposition, and changes in precipitation regimes. Despite the increasing appreciation that the consequences of impending global change can be better understood if varying agents are studied in concert, there is a paucity of multi‐factor long‐term studies, particularly on belowground processes. Herein, we address this gap by examining the responses of soil food webs and biodiversity to enrichment of CO₂, elevated N, and summer drought in a long‐term grassland study at Cedar Creek, Minnesota, USA (BioCON experiment). We use structural equation modeling (SEM), various abiotic and biotic explanatory variables, and data on soil microorganisms, protozoa, nematodes, and soil microarthropods to identify the impacts of multiple global change effects on drivers belowground. We found that long‐term (13‐year) changes in CO₂ and N availability resulted in modest alterations of soil biotic food webs and biodiversity via several mechanisms, encompassing soil water availability, plant productivity, and – most importantly – changes in rhizodeposition. Four years of manipulation of summer drought exerted surprisingly minor effects, only detrimentally affecting belowground herbivores and ciliate protists at elevated N. Elevated CO₂ increased microbial biomass and the density of ciliates, microarthropod detritivores, and gamasid mites, most likely by fueling soil food webs with labile C. Moreover, beneficial bottom‐up effects of elevated CO₂ compensated for detrimental elevated N effects on soil microarthropod taxa richness. In contrast, nematode taxa richness was lowest at elevated CO₂ and elevated N. Thus, enrichment of atmospheric CO₂ concentrations and N deposition may result in taxonomically and functionally altered, potentially simplified, soil communities. Detrimental effects of N deposition on soil biodiversity underscore recent reports on plant community simplification. This is of particular concern, as soils house a considerable fraction of global biodiversity and ecosystem functions.     

Carbon input by roots into the soil: Quantification of rhizodeposition from root to ecosystem scale

Despite its fundamental role for carbon (C) and nutrient cycling, rhizodeposition remains ‘the hidden half of the hidden half’: it is highly dynamic and rhizodeposits are rapidly incorporated into microorganisms, soil organic matter, and decomposed to CO₂. Therefore, rhizodeposition is rarely quantified and remains the most uncertain part of the soil C cycle and of C fluxes in terrestrial ecosystems. This review synthesizes and generalizes the literature on C inputs by rhizodeposition under crops and grasslands (281 data sets). The allocation dynamics of assimilated C (after ¹³C‐CO₂ or ¹⁴C‐CO₂ labeling of plants) were quantified within shoots, shoot respiration, roots, net rhizodeposition (i.e., C remaining in soil for longer periods), root‐derived CO₂, and microorganisms. Partitioning of C pools and fluxes were used to extrapolate belowground C inputs via rhizodeposition to ecosystem level. Allocation from shoots to roots reaches a maximum within the first day after C assimilation. Annual crops retained more C (45% of assimilated ¹³C or ¹⁴C) in shoots than grasses (34%), mainly perennials, and allocated 1.5 times less C belowground. For crops, belowground C allocation was maximal during the first 1–2 months of growth and decreased very fast thereafter. For grasses, it peaked after 2–4 months and remained very high within the second year causing much longer allocation periods. Despite higher belowground C allocation by grasses (33%) than crops (21%), its distribution between various belowground pools remains very similar. Hence, the total C allocated belowground depends on the plant species, but its further fate is species independent. This review demonstrates that C partitioning can be used in various approaches, e.g., root sampling, CO₂ flux measurements, to assess rhizodeposits’ pools and fluxes at pot, plot, field and ecosystem scale and so, to close the most uncertain gap of the terrestrial C cycle.     

Partitioning of ecosystem respiration of CO2 released during land‐use transition from temperate agricultural grassland to Miscanthus × giganteus

Conversion of large areas of agricultural grassland is inevitable if European and UK domestic production of biomass is to play a significant role in meeting demand. Understanding the impact of these land‐use changes on soil carbon cycling and stocks depends on accurate predictions from well‐parameterized models. Key considerations are cultivation disturbance and the effect of autotrophic root input stimulation on soil carbon decomposition under novel biomass crops. This study presents partitioned parameters from the conversion of semi‐improved grassland to Miscanthus bioenergy production and compares the contribution of autotrophic and heterotrophic respiration to overall ecosystem respiration of CO₂ in the first and second years of establishment. Repeated measures of respiration from within and without root exclusion collars were used to produce time‐series model integrations separating live root inputs from decomposition of grass residues ploughed in with cultivation of the new crop. These parameters were then compared to total ecosystem respiration derived from eddy covariance sensors. Average soil surface respiration was 13.4% higher in the second growing season, increasing from 2.9 to 3.29 g CO₂‐C m^?2 day^?1. Total ecosystem respiration followed a similar trend, increasing from 4.07 to 5.4 g CO₂‐C m^?2 day^?1. Heterotrophic respiration from the root exclusion collars was 32.2% lower in the second growing season at 1.20 g CO₂‐C m^?2 day^?1 compared to the previous year at 1.77 g CO₂‐C m^?2 day^?1. Of the total respiration flux over the two‐year time period, aboveground autotrophic respiration plus litter decomposition contributed 38.46% to total ecosystem respiration while belowground autotrophic respiration and stimulation by live root inputs contributed 46.44% to soil surface respiration. This figure is notably higher than mean figures for nonforest soils derived from the literature and demonstrates the importance of crop‐specific parameterization of respiration models.     

脉冲式降水对不同类型草地土壤微生物呼吸碳释放量的影响

降水事件引起土壤短时间内释放大量CO_2的现象常称为降水脉冲效应。降水事件发生后,由于水分和养分可获得性快速提升使土壤微生物呼吸速率快速升高至正常水分状况的数倍,从而导致土壤CO_2大量释放并一定程度上影响着生态系统碳循环过程和土壤碳平衡,尤其在干旱或半干旱地区。利用自主研发的能快速测定土壤微生物呼吸速率的装置,对内蒙古三类典型草原(草甸草原、典型草原和荒漠草原)土壤分别开展土壤复湿实验(60%饱和含水量),并采用高频测定(48 h测定288次)。在土壤复湿后在所有温带草地类型中均发生了明显的脉冲效应,降水脉冲过程中单位有机质(土壤有机碳,SOC)最大呼吸速率(R_(SOC-max))整体表现为荒漠草原(1.59 mg C g~(-1) SOC h~(-1))草甸草原(0.73 mg C g~(-1) SOC h~(-1))典型草原(0.50 mg C g~(-1) SOC h~(-1));而脉冲效应的持续时间(Duration)则表现为典型草原(2.5 h)草甸草原(1.5 h)荒漠草原(0.67 h)。在土壤复湿48 h内,单位土壤微生物呼吸累积量(A_(R_(Soil)))的大小规律与单位土壤微生物呼吸速率R_(Soil)一致,均为典型草原草甸草原荒漠草原;然而,如果用土壤有机质进行标准化,单位有机质呼吸累积量A_(R_(SOC))表现为荒漠草原(9.74 mg C g~(-1) SOC)典型草原(6.54 mg C g~(-1) SOC)草甸草原(3.54 mg C g~(-1) SOC),与当地年降雨频率呈负相关关系,表明降水脉冲效应与土壤长期经历的干旱状况存在密切关系。本研究结果不仅证明在干旱半干旱区域降水脉冲效应的普遍性,同时还启发我们应从国家或区域尺度开展研究,以进一步揭示土壤基质含量、土壤干旱状况等对降水脉冲效应的影响。     

Experimental fire increases soil carbon dioxide efflux in a grassland long‐term multifactor global change experiment

Numerous studies have demonstrated that soil respiration rates increase under experimental warming, although the long‐term, multiyear dynamics of this feedback are not well constrained. Less is known about the effects of single, punctuated events in combination with other longer‐duration anthropogenic influences on the dynamics of soil carbon (C) loss. In 2012 and 2013, we assessed the effects of decadal‐scale anthropogenic global change – warming, increased nitrogen (N) deposition, elevated carbon dioxide (CO₂), and increased precipitation – on soil respiration rates in an annual‐dominated Mediterranean grassland. We also investigated how controlled fire and an artificial wet‐up event, in combination with exposure to the longer‐duration anthropogenic global change factors, influenced the dynamics of C cycling in this system. Decade‐duration surface soil warming (1–2 °C) had no effect on soil respiration rates, while +N addition and elevated CO₂ concentrations increased growing‐season soil CO₂ efflux rates by increasing annual aboveground net primary production (NPP) and belowground fine root production, respectively. Low‐intensity experimental fire significantly elevated soil CO₂ efflux rates in the next growing season. Based on mixed‐effects modeling and structural equation modeling, low‐intensity fire increased growing‐season soil respiration rates through a combination of three mechanisms: large increases in soil temperature (3–5 °C), significant increases in fine root production, and elevated aboveground NPP. Our study shows that in ecosystems where soil respiration has acclimated to moderate warming, further increases in soil temperature can stimulate greater soil CO₂ efflux. We also demonstrate that punctuated short‐duration events such as fire can influence soil C dynamics with implications for both the parameterization of earth system models (ESMs) and the implementation of climate change mitigation policies that involve land‐sector C accounting.     

氮素添加和刈割对内蒙古弃耕草地土壤氮矿化的影响

以内蒙古多伦县恢复生态学试验示范研究站弃耕10余年的草地为研究对象,于2006年起分别设置对照、氮素添加、刈割和氮素添加+刈割4种处理,每种处理6次重复,研究弃耕草地氮素添加和刈割对土壤氮矿化的影响,结合土壤理化性质和植被地上生产力的动态变化,分析弃耕草地土壤氮矿化对植被恢复的响应,为当地草地恢复与重建提供理论依据和数据支持。实验结果表明:1氮素添加显著增加了植物地上净初级生产力(ANPP)和土壤无机氮库,与对照相比分别提高115%和196%,同时显著提高了土壤总硝化速率;但是氮素添加对总氨化速率、土壤微生物生物量碳(MBC)、微生物生物量氮(MBN)、微生物生物量碳氮比(MBC/MBN)、微生物呼吸(MR)以及呼吸熵(q CO2)均无显著影响;2总氨化速率和硝化速率对刈割处理的响应均不显著,但是刈割处理显著降低了土壤MR(P0.05);3氮素添加+刈割处理5—7a后,土壤总氨化和硝化速率均无显著变化;但是氮素添加+刈割处理显著增加了ANPP、土壤无机氮库和q CO2,同时显著降低了MBC和MBC/MBN。这说明在弃耕草地适应性管理中,氮素添加可以显著提高草地生产力,但是长期的氮添加对土壤微生物氮的转化是否有利还值得我们进一步研究。     

Partitioning of belowground C in young sugar maple forest

Background and aims

Trees allocate a high proportion of assimilated carbon belowground, but the partitioning of that C among ecosystem components is poorly understood thereby limiting our ability to predict responses of forest C dynamics to global change drivers.

Methods

We labeled sugar maple saplings in natural forest with a pulse of photosynthetic ¹³C in late summer and traced the pulse over the following 3 years. We quantified the fate of belowground carbon by measuring ¹³C enrichment of roots, rhizosphere soil, soil respiration, soil aggregates and microbial biomass.

Results

The pulse of ¹³C contributed strongly to root and rhizosphere respiration for over a year, and respiration comprised about 75 % of total belowground C allocation (TBCA) in the first year. We estimate that rhizosphere carbon flux (RCF) during the dormant season comprises at least 6 % of TBCA. After 3 years, 3.8 % of the C allocated belowground was recovered in soil organic matter, mostly in water-stable aggregates.

Conclusions

A pulse of carbon allocated belowground in temperate forest supplies root respiration, root growth and RCF throughout the following year and a small proportion becomes stabilized in soil aggregates.     

CO2 flux from soil in pastures and forests in southwestern Amazonia

Stocks of carbon in Amazonian forest biomass and soils have received considerable research attention because of their potential as sources and sinks of atmospheric CO₂. Fluxes of CO₂ from soil to the atmosphere, on the other hand, have not been addressed comprehensively in regard to temporal and spatial variations and to land cover change, and have been measured directly only in a few locations in Amazonia. Considerable variation exists across the Amazon Basin in soil properties, climate, and management practices in forests and cattle pastures that might affect soil CO₂ fluxes. Here we report soil CO₂ fluxes from an area of rapid deforestation in the southwestern Amazonian state of Acre. Specifically we addressed (1) the seasonal variation of soil CO₂ fluxes, soil moisture, and soil temperature; (2) the effects of land cover (pastures, mature, and secondary forests) on these fluxes; (3) annual estimates of soil respiration; and (4) the relative contributions of grass‐derived and forest‐derived C as indicated by δ¹³CO₂. Fluxes were greatest during the wet season and declined during the dry season in all land covers. Soil respiration was significantly correlated with soil water‐filled pore space but not correlated with temperature. Annual fluxes were higher in pastures compared with mature and secondary forests, and some of the pastures also had higher soil C stocks. The δ¹³C of CO₂ respired in pasture soils showed that high respiration rates in pastures were derived almost entirely from grass root respiration and decomposition of grass residues. These results indicate that the pastures are very productive and that the larger flux of C cycling through pasture soils compared with forest soils is probably due to greater allocation of C belowground. Secondary forests had soil respiration rates similar to mature forests, and there was no correlation between soil respiration and either forest age or forest biomass. Hence, belowground allocation of C does not appear to be directly related to the stature of vegetation in this region. Variation in seasonal and annual rates of soil respiration of these forests and pastures is more indicative of flux of C through the soil rather than major net changes in ecosystem C stocks.     

Effect of grazing on carbon stocks and assimilate partitioning in a Tibetan montane pasture revealed by 13CO2 pulse labeling

Since the late 1950s, governmental rangeland policies have changed the grazing management on the Tibetan Plateau (TP). Increasing grazing pressure and, since the 1980s, the privatization and fencing of pastures near villages has led to land degradation, whereas remote pastures have recovered from stronger overgrazing. To clarify the effect of moderate grazing on the carbon (C) cycle of the TP, we investigated differences in below‐ground C stocks and C allocation using in situ ¹³CO₂ pulse labeling of (i) a montane Kobresia winter pasture of yaks, with moderate grazing regime and (ii) a 7‐year‐old grazing exclosure plot, both in 3440 m asl. Twenty‐seven days after the labeling, ¹³C incorporated into shoots did not differ between the grazed (43% of recovered ¹³C) and ungrazed (38%) plots. In the grazed plots, however, less C was lost by shoot respiration (17% vs. 42%), and more was translocated below‐ground (40% vs. 20%). Within the below‐ground pools, <2% of ¹³C was incorporated into living root tissue of both land use types. In the grazed plots about twice the amount of ¹³C remained in soil (18%) and was mineralized to CO₂ (20%) as compared to the ungrazed plots (soil 10%; CO₂ 9%). Despite the higher contribution of root‐derived C to CO₂ efflux, total CO₂ efflux did not differ between the two land use types. C stocks in the soil layers 0–5 and 5–15 cm under grazed grassland were significantly larger than in the ungrazed grassland. However, C stocks below 15 cm were not affected after 7 years without grazing. We conclude that the larger below‐ground C allocation of plants, the larger amount of recently assimilated C remaining in the soil, and less soil organic matter‐derived CO₂ efflux create a positive effect of moderate grazing on soil C input and C sequestration.     

Land-use change in a tropical mountain rainforest region of southern Ecuador affects soil microorganisms and nutrient cycling

Over the past decades, the tropical mountain rainforest of southern Ecuador has been threatened by conversion to cattle pastures. Frequently, these pastures are invaded by bracken fern and abandoned when bracken becomes dominant. Changes in land-use (forest–pasture–abandoned pasture) can affect soil microorganisms and their physiological responses with respect to soil carbon and nutrient cycling. In situ investigations on litter decomposition and soil respiration as well as biogeochemical characterization of the soil were carried out to identify the driving factors behind. The conversion of forest to pasture induced a pronounced increase in CO₂–C effluxes to 12.2 Mg ha^?1 a^?1 which did not decrease after abandonment. Soil microbial activity and biomass showed a different pattern with lowest values at forest and abandoned pasture sites. With 3445 mg kg^?1 (0–5 cm) microbial biomass carbon (MBC by CFE-method), the active pasture had a more than three times higher value than forest and abandoned pasture, which was among the highest in tropical pasture soils. A shift in the microbial community structure (phospholipid fatty acid, PLFA) was also induced by the establishment of pasture land; the relative abundance of fungi and Gram-negative bacteria increased. PLFA fingerprints of the forest organic layer were more similar to pasture than to forest mineral soil. Chemical properties (pH value, exchangeable cations) were the main factors influencing the respective microbial structure. Bracken-invasion resulted in a decrease in the quantity and quality of above- and belowground biomass. The lower organic substance and nutrient availability induced a significant decline in microbial biomass and activity. After pasture abandonment, these differences in soil microbial function were not accompanied by pronounced shifts in the community structure and in soil pH as was shown for the conversion to pasture. A disconnection between microbial structure and function was identified. Similar soil CO₂–C effluxes between active and abandoned pasture sites might be explained by an underestimation of the effluxes from the active pasture site. All measurements were carried out between grass tussocks where fine-root density was about 2.6 times lower than below tussocks. Thus, lower proportions of root respiration were expected than below tussocks. Overall, soil microorganisms responded differently to changes in land-use from forest to pasture and from pasture to abandoned pasture resulting in pronounced changes of carbon and nutrient cycling and hence of ecosystem functioning.     

Long-term carbon exchange in a sparse, seasonally dry tussock grassland

Rainfall and its seasonal distribution can alter carbon dioxide (CO₂) exchange and the sustainability of grassland ecosystems. Using eddy covariance, CO₂ exchange between the atmosphere and a sparse grassland was measured for 2 years at Twizel, New Zealand. The years had contrasting distributions of rain and falls (446 mm followed by 933 mm; long‐term mean=646 mm). The vegetation was sparse with total above‐ground biomass of only 1410 g m^?2. During the dry year, leaf area index peaked in spring (November) at 0.7, but it was <0.2 by early summer. The maximum daily net CO₂ uptake rate was only 1.5 g C m^?2 day^?1, and it occurred before mid‐summer in both years. On an annual basis, for the dry year, 9 g C m^?2 was lost to the atmosphere. During the wet year, 41 g C m^?2 was sequestered from the atmosphere. The net exchange rates were determined mostly by the timing and intensity of spring rainfall. The components of ecosystem respiration were measured using chambers. Combining scaled‐up measurements with the eddy CO₂ effluxes, it was estimated that 85% of ecosystem respiration emanated from the soil surface. Under well‐watered conditions, 26% of the soil surface CO₂ efflux came from soil microbial activity. Rates of soil microbial CO₂ production and net mineral‐N production were low and indicative of substrate limitation. Soil respiration declined by a factor of four as the soil water content declined from field capacity (0.21 m³ m^?3) to the driest value obtained (0.04 m³ m^?3). Rainfall after periods of drought resulted in large, but short‐lived, respiration pulses that were curvilinearly related to the increase in root‐zone water content. Coupled with the low leaf area and high root : shoot ratio, this sparse grassland had a limited capacity to sequester and store carbon. Assuming a proportionality between carbon gain and rainfall during the summer, rainfall distribution statistics suggest that the ecosystem is sustainable in the long term.     

Soil Biological and Chemical Properties in Restored Perennial Grassland in California

Restoration of California native perennial grassland is often initiated with cultivation to reduce the density and cover of non‐native annual grasses before seeding with native perennials. Tillage is known to adversely impact agriculturally cultivated land; thus changes in soil biological functions, as indicated by carbon (C) turnover and C retention, may also be negatively affected by these restoration techniques. We investigated a restored perennial grassland in the fourth year after planting Nassella pulchra, Elymus glaucus, and Hordeum brachyantherum ssp. californicum for total soil C and nitrogen (N), microbial biomass C, microbial respiration, CO₂ concentrations in the soil atmosphere, surface efflux of CO₂, and root distribution (0‐ to 15‐, 15‐ to 30‐, 30‐ to 60‐, and 60‐ to 80‐cm depths). A comparison was made between untreated annual grassland and plots without plant cover still maintained by tillage and herbicide. In the uppermost layer (0‐ to 15‐cm depth), total C, microbial biomass C, and respiration were lower in the tilled, bare soil than in the grassland soils, as was CO₂ efflux from the soil surface. Root length near perennial bunchgrasses was lower at the surface and greater at lower depths than in the annual grass–dominated areas; a similar but less pronounced trend was observed for root biomass. Few differences in soil biological or chemical properties occurred below 15‐cm depth, except that at lower depths, the CO₂ concentration in the soil atmosphere was lower in the plots without vegetation, possibly from reduced production of CO₂ due to the lack of root respiration. Similar microbiological properties in soil layers below 15‐cm depth suggest that deeper microbiota rely on more recalcitrant C sources and are less affected by plant removal than in the surface layer, even after 6 years. Without primary production, restoration procedures with extended periods of tillage and herbicide applications led to net losses of C during the plant‐free periods. However, at 4 years after planting native grasses, soil microbial biomass and activity were nearly the same as the former conditions represented by annual grassland, suggesting high resilience to the temporary disturbance caused by tillage.     

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.     

Interactions between plant growth and soil nutrient cycling under elevated CO2: a meta-analysis

free air carbon dioxide enrichment (FACE) and open top chamber (OTC) studies are valuable tools for evaluating the impact of elevated atmospheric CO₂ on nutrient cycling in terrestrial ecosystems. Using meta‐analytic techniques, we summarized the results of 117 studies on plant biomass production, soil organic matter dynamics and biological N₂ fixation in FACE and OTC experiments. The objective of the analysis was to determine whether elevated CO₂ alters nutrient cycling between plants and soil and if so, what the implications are for soil carbon (C) sequestration. Elevated CO₂ stimulated gross N immobilization by 22%, whereas gross and net N mineralization rates remained unaffected. In addition, the soil C : N ratio and microbial N contents increased under elevated CO₂ by 3.8% and 5.8%, respectively. Microbial C contents and soil respiration increased by 7.1% and 17.7%, respectively. Despite the stimulation of microbial activity, soil C input still caused soil C contents to increase by 1.2% yr^?1. Namely, elevated CO₂ stimulated overall above‐ and belowground plant biomass by 21.5% and 28.3%, respectively, thereby outweighing the increase in CO₂ respiration. In addition, when comparing experiments under both low and high N availability, soil C contents (+2.2% yr^?1) and above‐ and belowground plant growth (+20.1% and+33.7%) only increased under elevated CO₂ in experiments receiving the high N treatments. Under low N availability, above‐ and belowground plant growth increased by only 8.8% and 14.6%, and soil C contents did not increase. Nitrogen fixation was stimulated by elevated CO₂ only when additional nutrients were supplied. These results suggest that the main driver of soil C sequestration is soil C input through plant growth, which is strongly controlled by nutrient availability. In unfertilized ecosystems, microbial N immobilization enhances acclimation of plant growth to elevated CO₂ in the long‐term. Therefore, increased soil C input and soil C sequestration under elevated CO₂ can only be sustained in the long‐term when additional nutrients are supplied.     

Plants Mediate the Sensitivity of Soil Respiration to Rainfall Variability

Soil respiration from grasslands plays a critical role in determining carbon dioxide (CO₂) feedbacks between soils and the atmosphere. In these often mesic systems, soil moisture and temperature tend to co-regulate soil respiration. Increasing variance of rainfall patterns may alter aboveground–belowground interactions and have important implications for the sensitivity of soil respiration to fluctuations in moisture and temperature. We conducted a set of field experiments to evaluate the independent and interactive effects of rainfall variability and plant–soil processes on respiration dynamics. Plant removal had strong effects on grassland soils, which included altered CO₂ flux owing to absence of root respiration; increased soil moisture and temperature; and reduced availability of dissolved organic carbon (DOC) for heterotrophic respiration by microorganisms. These plant-mediated effects interacted with our rainfall variability treatments to determine the sensitivity of soil respiration to both moisture and temperature. Using time-series multiple regression, we found that plants dampened the sensitivity of respiration to moisture under high variability rainfall treatments, which may reflect the relative stability of root contributions to total soil respiration. In contrast, plants increased the sensitivity of respiration to temperature under low variability rainfall treatment suggesting that the environmental controls on soil CO₂ dynamics in mesic habitats may be context dependent. Our results provide insight into the aboveground–belowground mechanisms controlling respiration in grasslands under variable rainfall regimes, which may be important for predicting CO₂ dynamics under current and future climate scenarios.     

Drought decreases incorporation of recent plant photosynthate into soil food webs regardless of their trophic complexity

Theory suggests that more complex food webs promote stability and can buffer the effects of perturbations, such as drought, on soil organisms and ecosystem functions. Here, we tested experimentally how soil food web trophic complexity modulates the response to drought of soil functions related to carbon cycling and the capture and transfer below‐ground of recent photosynthate by plants. We constructed experimental systems comprising soil communities with one, two or three trophic levels (microorganisms, detritivores and predators) and subjected them to drought. We investigated how food web trophic complexity in interaction with drought influenced litter decomposition, soil CO₂ efflux, mycorrhizal colonization, fungal production, microbial communities and soil fauna biomass. Plants were pulse‐labelled after the drought with ¹³C‐CO₂ to quantify the capture of recent photosynthate and its transfer below‐ground. Overall, our results show that drought and soil food web trophic complexity do not interact to affect soil functions and microbial community composition, but act independently, with an overall stronger effect of drought. After drought, the net uptake of ¹³C by plants was reduced and its retention in plant biomass was greater, leading to a strong decrease in carbon transfer below‐ground. Although food web trophic complexity influenced the biomass of Collembola and fungal hyphal length, ¹³C enrichment and the net transfer of carbon from plant shoots to microbes and soil CO₂ efflux were not affected significantly by varying the number of trophic groups. Our results indicate that drought has a strong effect on above‐ground–below‐ground linkages by reducing the flow of recent photosynthate. Our results emphasize the sensitivity of the critical pathway of recent photosynthate transfer from plants to soil organisms to a drought perturbation, and show that these effects may not be mitigated by the trophic complexity of soil communities, at least at the level manipulated in this experiment.     

Precipitation pulses enhance respiration of Mediterranean ecosystems: the balance between organic and inorganic components of increased soil CO2 efflux

In regions characterized by arid seasons, such as the Mediterranean basin, soil moisture is a major driver of ecosystem CO₂ efflux during periods of drought stress. Here, a rain event can induce a disproportional respiratory pulse, releasing an amount of CO₂ to the atmosphere that may significantly contribute to the annual ecosystem carbon balance. The mechanisms behind this pulse are unclear, and it is still unknown whether it is due to the stimulation of autotrophic, heterotrophic and/or inorganic CO₂ fluxes. On the Mediterranean island of Pianosa, eddy flux measurements showed respiratory pulses after rain events following prolonged drought periods, which occurred in the summer of 2003 and 2006. To investigate the mechanisms of this observed enhanced respiration fluxes and partition of the soil CO₂ sources, two water manipulation experiments were performed. The first was designed to estimate the effect of soil rewetting on soil CO₂ efflux, in the different ecosystem types existing on the island (i.e. woodland, ex‐agricultural and Mediterranean shrubland). The second was a soil CO₂ partitioning experiment to investigate the relative contribution of inorganic and organic CO₂ sources to soil respiration, under dry and wet soil conditions. Our results suggest that the pulse in the CO₂ efflux is primarily due to the enhancement of heterotrophic respiration, likely caused by the degradation of easily decomposable substrates, accumulated in soils during the dry period. In fact, the vegetation at the site was senescent and did not play any significant role in CO₂ exchange, as suggested by the absence of diurnal CO₂ uptake in eddy covariance measurements. In addition, soil rewetting did not significantly enhance inorganic CO₂ efflux.     

Patterns of rhizosphere carbon flux in sugar maple (Acer saccharum) and yellow birch (Betula allegheniensis) saplings

Despite its importance in the terrestrial C cycle rhizosphere carbon flux (RCF) has rarely been measured for intact root–soil systems. We measured RCF for 8‐year‐old saplings of sugar maple (Acer saccharum) and yellow birch (Betula allegheniensis) collected from the Hubbard Brook Experimental Forest (HBEF), NH and transplanted into pots with native soil horizons intact. Five saplings of each species were pulse labeled with ¹³CO₂ at ambient CO₂ concentrations for 4–6 h, and the ¹³C label was chased through rhizosphere and bulk soil pools in organic and mineral horizons for 7 days. We hypothesized yellow birch roots would supply more labile C to the rhizosphere than sugar maple roots based on the presumed greater C requirements of ectomycorrhizal roots. We observed appearance of the label in rhizosphere soil of both species within the first 24 h, and a striking difference between species in the timing of ¹³C release to soil. In sugar maple, peak concentration of the label appeared 1 day after labeling and declined over time whereas in birch the label increased in concentration over the 7‐day chase period. The sum of root and rhizomicrobial respiration in the pots was 19% and 26% of total soil respiration in sugar maple and yellow birch, respectively. Our estimate of the total amount of RCF released by roots was 6.9–7.1% of assimilated C in sugar maple and 11.2–13.0% of assimilated C in yellow birch. These fluxes extrapolate to 55–57 and 90–104 g C m^?2 yr^?1 from sugar maple and yellow birch roots, respectively. These results suggest RCF from both arbuscular mycorrhizal and ectomycorrhizal roots represents a substantial flux of C to soil in northern hardwood forests with important implications for soil microbial activity, nutrient availability and C storage.