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One of the most prominent changes in the rhizospheric soil structure is associated with the formation of a strongly bound soil layer in the surroundings of the root, which is named rhizosheath. In this study, we investigated how root hair elongation, a ubiquitous root morphological trait, affect the stability of rhizosheath aggregates. Using 13CO2 pulse labeling, we tracked the fate of root-derived 13C inputted into the rhizosheath of two Zea mays L. genotypes with contrasting root hair elongation: a mutant with root hair defective elongation (rth3) and a corresponding wild type (WT). In addition, we also investigated the differences between two 13CO2 labeling approaches (single vs. multiple pulse labeling) in the distribution of 13C in the rhizosheath aggregates. We were able to demonstrate that the rhizosheath aggregate stability and the resulting aggregate size distribution follows the same mechanisms irrespective of the root hair elongation. This result reinforces the assumption that other soil properties are more decisive for the soil structure formation in the rhizosheath in comparison to root hair elongation. The majority of recently deposited root-derived C (57%) was found in the macroaggregates. Increasing the number of pulses (multiple pulse labeling approach) resulted in a higher 13C enrichment of the rhizosheath aggregates fractions in comparison to the application of a single pulse. While both labeling approaches resulted in a similar distribution of 13C in the rhizosheath aggregates, the higher enrichment given by multiple pulse labeling allowed the separation of significant differences between the genotypes in plant C allocation in the rhizosheath.

Agroforestry systems are characterized by a high complexity between vegetation components and niche partitioning. In a crop-grass-tree agroforestry system, rape, willow, and grasses were in situ pulse labeled separately with 13CO2 for 6 h, and 13C was traced in shoots, roots, topsoil (0–15 cm) and subsoil (15–30 cm), microbial biomass carbon (C), and dissolved organic C, as well as respiration losses (CO2) up to 28 days after labeling to investigate the effects of vegetation components on C allocation belowground. 13C recovery in roots after 28 days was 7.0% of total assimilated C for grassland, which was 3.5- and 5.2-fold higher than that for rape and willow, respectively. The larger C allocation belowground in grassland was ascribed to its higher root/shoot ratio compared to willow and rape. Grassland facilitated higher accumulation of root-derived C in soil compared to rape (9.2% of recovered 13C) and compared to willow (1.6% of 13C). Willow retained more photosynthetic C aboveground and less was allocated to roots compared to rape. Although the C allocated to the top 15-cm soil was similar between willow and rape, willow facilitated C allocation in deeper soil compared to rape (0.6% vs. 0.2%). This could be explained by the lower microbial activity and subsequent weaker decomposition of rhizodeposits in 15–30-cm depth under willow. The net belowground C inputs in grassland, willow, and rape were 0.53, 0.06, and 0.10 g C m−2 month−1 of vegetation period, including rhizodeposition of 0.24, 0.05, and 0.04 g C m−2 month−1, respectively. Overall, integrating trees and grassland within cropland facilitates higher root-derived C input into soil, thus contributing to the soil C sequestration in agroforestry systems.

Aims Soil priming affects soil N transformation and plant N availability, but few studies have investigated these interactions to date. Methods To address this, we reviewed the literature for studies quantifying soil priming, soil N transformation and plant N uptake. Results Gross N mineralization was strongly controlled by soil priming in studies with plants, while abiotic factors had a minor influence on gross N mineralization. In contrast, soil priming was negatively related to gross N mineralization and had a low explanatory power in incubation studies where substrates are added as surrogates for root exudates. These results indicate that plants support increased N mineralization and that this is not adequately reflected in incubation studies. Additionally, we observed a positive relationship between soil priming and the % of Norg-derived N uptake as well as total N uptake, which demonstrates that priming enhances the availability of N that was previously organically bound and that at least part of the N mineralized during priming was available for plant uptake. Conclusion Our results show that the effect of roots and rhizodeposition leads to a number of processes supporting N mineralization and availability through priming that are not well reflected in incubation studies. To fully capture the interactions between plant roots and their associated microbiota, we recommend focusing research on systems with plants. Additionally, the strong correlation between C and N transformation should be considered in biogeochemical modelling.

Background and aims Plant growth affects soil moisture, mineral N and organic C availability in soil, all of which influence denitrification. With increasing plant growth, root exudation may stimulate denitrification, while N uptake restricts nitrate availability. Methods We conducted a double labeling pot experiment with either maize (Zea mays L.) or cup plant (Silphium perfoliatum L.) of the same age but differing in size of their shoot and root systems. The 15N gas flux method was applied to directly quantify N2O and N2 fluxes in situ. To link denitrification with available C in the rhizosphere, 13CO2 pulse labeling was used to trace C translocation from shoots to roots and its release by roots into the soil. Results Plant water and N uptake were the main factors controlling daily N2O + N2 fluxes, cumulative N emissions, and N2O production pathways. Accordingly, pool-derived N2O + N2 emissions were 30–40 times higher in the treatment with highest soil NO3− content and highest soil moisture. CO2 efflux from soil was positively correlated with root dry matter, but we could not detect any relationship between root-derived C and N2O + N2 emissions. Conclusions Root-derived C may stimulate denitrification under small plants, while N and water uptake become the controlling factors with increasing plant and root growth.

There is a trend of increasing woody biomass in tropical savannas. Here we ask what effect this increase may have on soil carbon pools and fluxes. Using a field experiment we determine the amount of soil carbon directly under grasses, a juvenile tree among grasses and a juvenile tree with no grasses. We also measure CO2 efflux at the soil surface and use gas wells to extract CO2 from several soil depths. Our results show that grasses contribute substantially more than trees to both soil carbon and soil respiration. Grasses also make a disproportionate contribution to the δ13C value of SOC in the tree only treatments. The surface gas efflux data show that soil respiration increased with an increase in volumetric soil moisture and temperature and plots with both grasses and trees had higher respiration rates than plots with trees only or with grasses only. The highest soil respiration is in the top 20 cm of the soil with grasses the primary contributors to both δ13CSOC and δ13CR. Any increase in woody biomass will result in a decline in SOM turnover and nitrogen mineralization rates resulting in higher SOC pools. The associated increases in SOC and above ground biomass will however be associated with negative economic and biodiversity impacts.

As the dynamics and magnitude of rhizodeposition vary considerably among cropping systems, we investigated effects of cropping system on the incorporation of root-derived carbon (C) into Collembola, a dominant taxon of soil microarthropods. In the field, we used 13CO2 to pulse label a crop monoculture (oilseed rape, Brassica napus L.), a mixed-grass community (dominated by Lolium perenne L. mixed with clover Trifolium repens L.), and a tree plantation (willow, Salix schwerinii E.L. Wolf and Salix viminalis L.). During 28 days, the incorporation of 13C was traced in nine species of Collembola including epedaphic (surface-dwelling), hemiedaphic (litter-dwelling), and euedaphic (soil-dwelling) functional groups. Incorporation of 13C into Collembola reached a plateau before day 3 after the labeling in grass and willow, but increased up to day 14 in rape. While euedaphic Collembola incorporated less root-derived C than epedaphic and hemiedaphic Collembola in rape and willow, the incorporation of 13C was similar among functional groups in grass. Differential incorporation of 13C in euedaphic species points to niche differentiation within the same functional group. Our findings highlight that cropping system not only affects the flux of root C into soil mesofauna, being slower in rape than in grass and willow, but also the utilization of root-derived resources by functional groups and species of Collembola. The results indicate that pronounced differences in belowground C inputs between cropping systems affect microbivores as basal species and thereby soil food webs and their functioning and services.

Microbial turnover processes are typically restricted by low substrate availability in the subsoil. We hypothesized that SOM decomposition increases with plant density and decreases with N fertilization: We expected a greater rate of C allocation to the rhizosphere in the topsoil than in the subsoil treatments. In order to simulate different degrees of rhizodeposition, wheat was planted in pots at four different densities. The plants were continuously labeled with 13C-depleted CO2. Soil CO2 efflux was partitioned for root- and SOM-derived CO2. Moreover, we determined the enzyme kinetics by measuring catalytic efficiency and enzyme stoichiometry in both topsoil and subsoil. Shoot biomass and the shoot to root ratio were significantly higher for plants grown in the topsoil compared with the subsoil, which demonstrated higher relative C allocation to root biomass in the subsoil treatment. Despite the similar size of the rhizosphere, root-derived CO2 was always higher in the topsoil compared with the subsoil treatment, indicating enhanced root exudation. Effect sizes of all enzyme activities showed stronger magnitudes for the subsoil treatments. This was in line with a two-times increase of the effect size of SOM decomposition in the subsoil relative to topsoil. Overall, the plants in the subsoil treatments allocated more C to root biomass, less C to shoot biomass, and substantially less C to root exudates. However, the effect sizes of both SOM decomposition and enzyme activities were higher in the subsoil than in the topsoil, reflecting a stronger sensitivity to C inputs.

Kobresia grasslands on the Tibetan Plateau comprise the world’s largest pastoral alpine ecosystem. Overgrazing-driven degradation strongly proceeded on this vulnerable grassland, but the mechanisms behind are still unclear. Plants must balance the costs of releasing C to soil against the benefits of accelerated microbial nutrient mineralization, which increases their availability for root uptake. To achieve the effect of grazing on this C-N exchange mechanism, a 15NH4+ field labeling experiment was implemented at grazed and ungrazed sites, with additional treatments of clipping and shading to reduce belowground C input by manipulating photosynthesis. Grazing reduced gross N mineralization rates by 18.7%, similar to shading and clipping. This indicates that shoot removal by grazing decreased belowground C input, thereby suppressing microbial N mining and overall soil N availability. Nevertheless, NH4+ uptake rate by plants at the grazed site was 1.4 times higher than at the ungrazed site, because plants increased N acquisition to meet the high N demands of shoot regrowth (compensatory growth: grazed > ungrazed). To enable efficient N uptake and regrowth, Kobresia plants have developed specific traits (i.e., efficient above-belowground interactions). These traits reflect important mechanisms of resilience and ecosystem stability under long-term moderate grazing in an N-limited environment. However, excessive (over)grazing might imbalance such C-N exchange and amplify plant N limitation, hampering productivity and pasture recovery over the long term. In this context, a reduction in grazing pressure provides a sustainable way to maintain soil fertility, C sequestration, efficient nutrient recycling, and overall ecosystem stability.

Biopores are hotspots of nutrient mobilisation and shortcuts for carbon (C) into subsoils. C processing relies on microbial community composition, which remains unexplored in subsoil biopores. Phospholipid fatty acids (PLFAs; markers for living microbial groups) and amino sugars (microbial necromass markers) were extracted from two subsoil depths (45–75 cm ; 75–105 cm) and three biopore types: (I) drilosphere of Lumbricus terrestris L., (II) 2-year-old root biopores and (III) 1.5-year-old root biopores plus six 6 months of L. terrestris activities. Biopore C contents were at least 2.5 times higher than in bulk soil, causing 26–35 times higher Σ PLFAs g-1 soil. The highest Σ PLFAs were found in both earthworm biopore types; thus, the highest soil organic matter and nutrient turnover were assumed. Σ PLFAs was 33% lower in root pores than in earthworm pores. The treatment affected the microbial community composition more strongly than soil depth, hinting to similar C quality in biopores: Gram-positives including actinobacteria were more abundant in root pores than in earthworm pores, probably due to lower C bioavailability in the former. Both earthworm pore types featured fresh litter input, promoting growth of Gram-negatives and fungi. Earthworms in root pores shifted the composition of the microbial community heavily and turned root pores into earthworm pores within 6 months. Only recent communities were affected and they reflect a strong heterogeneity of microbial activity and functions in subsoil hotspots, whereas biopore-specific necromass accumulation from different microbial groups was absent.

The applications of biochar (BC) and polyacrylamide (PAM) may have interactive effects on carbon (C) dynamics and sequestration for improving the soil quality and achieving sustainable agriculture. Relative to BC and PAM, rhizodeposits act as C and energy source for microorganisms and may change the mineralization dynamics of soil organic matter (SOM). No attempt has been made to assess the effects of BC, anionic PAM, or their combination on the decomposition of different aged 14C-labeled rhizodeposits. The objective of this study was to investigate the effects of the treatments mentioned above on the decomposition of different aged 14C-labeled maize rhizodeposits. biochar (BC) at 10 Mg ha−1 or anionic PAM at 80 kg ha−1 or their combination (BC + PAM) was applied to soils with/without 2-, 4-, 8-, and 16-day-aged 14C-labeled maize rhizodeposits. After that, the soil was incubated at 22 °C for 46 days. After 2 days of incubation, the total CO2 efflux rates from the soil with rhizodeposits were 1.4–1.8 times higher than those from the soil without rhizodeposits. The cumulative 14CO2 efflux (32 % of the 14C input) was maximal for the soil containing 2-day-aged 14C-labeled rhizodeposits. Consequently, 2-day-aged rhizodeposits were more easily and rapidly decomposed than the older rhizodeposits. However, no differences in the total respired 14CO2 from rhizodeposits were observed at the end of the incubation. Incorporation of 14C into microbial biomass and 66–85 % of the 14C input remained in the soil after 46 days indicated that neither the age of 14C-labeled rhizodeposits nor BC, PAM, or BC + PAM changed microbial utilization of rhizodeposits. Applying BC or BC + PAM to soil exerted only minor effects on the decomposition of rhizodeposits. The contribution of rhizodeposits to CO2 efflux from soil and MBC depends on their age as young rhizodeposits contain more labile C, which is easily available for microbial uptake and utilization.

Biopores are pores or voids in soil produced by roots, by earthworms, or by the occupation of earthworms in root pores, which are considered important microbial hotspots, especially in subsoil. We hypothesized that earthworms (Lumbricus terrestris L.) exert stronger effects on microbial activities than decaying plant roots (of Cichorium intybus L.) in the subsoil because of the addition of pre-digested organic material. We tested this hypothesis by analyzing microbial biomass (Cmic), total organic C (Corg), and activities of eight enzymes (cellobiohydrolase, β-glucosidase, xylanase, acid phosphomonoesterase, leucine aminopeptidase, tyrosine aminopeptidase, chitotriosidase, and n-acetylglucosaminidase) down to 105-cm depth. The Cmic increase was associated with a two- to threefold increase of Corg content in biopores as compared to bulk soil. The highest percentage of Cmic-to-Corg (3.7 to 7.3 %) in the drilosphere demonstrated the enhancement of microbial efficiency for organic matter decomposition by earthworms. The availability of organic matter in biopores increased the activities of C- and N-targeting enzymes by 1.2–11.3 times, but reduced acid phosphomonoesterase activity by 10–40 % in biopores versus bulk soil. Introducing earthworms in root biopores caused 1.5–1.8 times higher microbial biomass and 1.2–1.9 times increased enzyme activities compared to the sole effect of earthworms. Soil depth showed a strong effect on the drilosphere, but only slight effects on the biochemical properties of root biopores and bulk soil. In conclusion, biopores are important microbial hotspots of C, N, and P transformations in subsoil. Earthworms exerted stronger effects on biochemical properties of biopores than decaying roots.

The complexity of ecosystem processes, especially under continuously changing environmental conditions, requires high-resolution insight into ecosystem carbon (C) fluxes. It is essential to gain not only information about relative C balance and fluxes (common for partitioning studies), but also to obtain these in absolute mass units. To evaluate absolute fluxes in belowground C pools, the results of 21-day eddy-covariance and stable isotope labeling experiment in summer 2010, were combined. Eddy-covariance based net ecosystem exchange was measured on extensively managed grassland and separated into underlying assimilation and ecosystem respiration through the use of a C flux partitioning model. Resultant CO2 assimilation served as absolute C input into the ecosystem and was further partitioned by applying the relative C distribution in subsidiary pools, gained by 13C pulse labeling and tracing. The results form eddy-covariance measurements showed that the extensively managed grassland was a significant net C sink of −91 g C m−2 a−1 in 2010. The mean daily assimilation of −7.1 g C m−2 d−1 was partitioned into fluxes of 2.5, 0.8, 0.5, 2.3 and 1.0 g C m−2 d−1 into shoots, roots, soil, shoot respiration and CO2 efflux from soil, respectively. We conclude that the combination of EC measurements with isotope labeling techniques allowed determining the absolute C input into several ecosystem pools. Hence, the study demonstrates an approach to expand atmospheric flux measurements and to gain insight into the importance of individual ecosystem pools for soil C cycling.

Although water conservation in rice production has become increasingly important, the effects of water management on the allocation and dynamics of carbon (C) within the rice-soil system remain unknown. We compared the allocation and dynamics of C assimilated by rice under continuously flooded, non-flooded and alternate water regimes. Rice (Oryza sative L. cv. Luliangyou 996) was labeled with 14CO2 and harvested 7 times within 45 days. More 14C was released from roots into the soil in non-flooded and alternate water regimes treatments. Microbial 14C decreased with time after the labeling and was lowest under flooded condition. Roots and rhizomicrobial respiration followed the order of non-flooded > alternate water regimes > flooded treatment. Water management affected 14C distribution in aggregates with more 14C in macroaggregates in the non-flooded treatment. Estimated amounts of C transferred remaining belowground by rice 45 days after labeling were 1,986, 2,827 and 2,472 kg C ha−1, of which rhizodeposition accounted for about 41 %, 16 % and 30 % of C transferred belowground under non-flooded, flooded and alternate water regimes, respectively. Water management affected the allocation and dynamics of recently assimilated C within the rice-soil system and also changed the relative contribution of rhizodeposition to C transferred belowground. This study suggests the differences in the driving mechanisms of C sequestration under flooded vs. non-flooded and alternate water regimes.