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Phosphorus (P) is a major limiting nutrient for plant growth implying an often-intensive competition between microorganisms and plants in the rhizosphere. Increasing the P availability in subsoils may help to mitigate potential future P fertilizer shortages and to overcome P limitations due to droughts, which mainly affect topsoils. Root exudates provide easily available carbon and energy sources for microorganisms to mobilize soil nutrients. Nonetheless, details regarding the distinct processes underlying P mobilization from various P sources (free vs. sorbed PO 4 3−; low molecular vs. complex organic P, e.g. ATP vs. plant litter P) as affected by root exudates are poorly understood, especially in subsoils. This study aimed to identify the controlling factors and microbial processes regulating the availability of organic and inorganic P in top- and subsoils by 33P isotopic labeling. The focus was on the potential key role of root exudates in P mobilization. We found that microbial communities in top- and subsoils used high- and low-available mineral P to a similar extent, but that the subsoil communities were much more efficient in mobilizing and incorporating complex litter-derived organic P. This capability of subsoil communities was even enhanced when root exudates were present. Microbial activity and nutrient-mobilizing mechanisms (e.g., P-related enzymes) clearly increased by root exudate addition, an effect that was generally higher in sub-than in topsoils. We conclude that subsoil communities are well capable of mobilizing and using complex organic P sources, especially if root exudates accelerate overall activity and P cycling. Thus, high root exudation is highly relevant for crops, which depend on subsoil nutrients and litter-derived P. Accordingly, detritusphere P, e.g. in subsoil root channels, is likely to be plant-available because of exudate-induced microbial P (re-)cycling processes. [Display omitted] • Root exudates did not affect growth of the microbial community nor lead to a community shift. • Due to rapid desorption microbes use sorbed P similarly to mineral P added to soil solution. • Low available litter P was mobilized and incorporated more efficient by sub than topsoil communities. • Root exudates increase microbial activity and thus boost P mining process. [ABSTRACT FROM AUTHOR]

Most subsoil carbon (C) turnover occurs in biopore hotspots such as root channels and earthworm burrows. Biopores allocate large C amounts into the subsoil, where a vast capacity for long-term C sequestration is predicted. We hypothesise that organic matter (OM) cycling in biopores depends on their origin. Earthworm and root biopores were induced under field conditions and were sampled from the subsoil (45–75 and 75–105 cm) after two years of biopore formation. The effects of biopore formation on OM decomposition were studied by biomarkers: neutral sugars, cutin and suberin-derived lipids, lignin-derived phenols and free lipids. The degradation stage of OM was biopore type-specific but was only governed by the soil depth in root biopores. Degradation of OM increased from earthworm biopores to root biopores and bulk soil. Hemicelluloses (GM/AX ratio) were more strongly degraded than lignin side-chains (relative change from initial values). Two years of microbial processing during biopore formation increased the GM/AX ratio in earthworm biopores from 0.65 to 1.05 and in root biopores from 0.15 to 1.35 (both relative to source biomasses). Root biopores and bulk soil had the highest GM/AX ratios (1.2–1.3), hinting to rapid processing of plant residues and accumulation of microbial residues. The regular, frequent OM inputs by earthworms stimulated microbial growth and processing of mostly bioavailable OM and, thus, relatively enriched more persistent OM (e.g. lignin). Syringyl subunits of lignin underwent low (ratio changed from 0.35 to 0.55 relative to initial input) and vanillyl subunits underwent almost no processing in earthworm biopores indicating the preferential microbial utilisation of the easily available compounds frequently replenished by earthworm activity. After two years of decomposition of the root detritus, mainly structural plant material was enriched in root biopores. Short periods (6 months) of earthworm activity effectively recharged the highly processed OM in root biopores with fresh OM. In total, deep-rooting catch crops and short-term earthworm activities promote C accumulation in the subsoil followed by biopore-specific microbial processing predominantly governed by the C input frequency. As root biopores are up to 40 times more common than earthworm biopores, they dominate the OM input into subsoils. Such C inputs create several years lasting hotspots for preferential root growth and nutrient mobilisation in the subsoil. We conclude that root- and earthworm-derived biopores are vertical pathways for plant C from the soil surface into the subsoil and for intensive processing of litter C and sequestration of microbial necromass. [ABSTRACT FROM AUTHOR]

Meadows and forests are the main vegetation types in temperate terrestrial ecosystems, and largely contribute to soil carbon (C) stock. Bioavailable C inputs can accelerate microbial decomposition of soil organic matter (SOM), which is known as "priming effect". However, it is still unclear how priming effect, as an important mechanism influencing soil C sequestration, is influenced by spatial transition of vegetation from meadow to forest. To investigate the mechanism of priming effect along a spatial transition gradient of vegetation, a soil incubation experiment with 14C labeled glucose was combined with microbial rDNA sequencing and gene composition prediction. The results showed that with the vegetation transition from meadow to forest soil available phosphorus (P) significantly increased, in contrast to dissolved nitrogen (N) and C which remained unaffected. Moreover, the soil microbial community composition shifted towards a higher relative abundance of K-strategists (Acidobacteria) and a lower abundance of r-strategists (Actinobacteriota) along the vegetation transition from meadow to forest. In the meadow, the microbial community consumed more of the added glucose and increased priming effect. This was accompanied by lower available P but higher soil bacterial gene function encoding P cycling. In contrast, increased soil P availability in forest soils caused a decelerated microbial metabolism of phosphorylated organic compounds within microbial biomass due to decreased microbial demand for P acquisition from SOM, and thus resulted in suppression of the priming effect. Our study showed that P availability and microbial community shifts in spatial transition zones between meadows and forests are important drivers for the priming effect on SOM decomposition. • 14C radioactive isotope labeling was applied in combination with microbial rDNA sequencing. • Soil microbial community in meadow consumed more of the added glucose than that in forest. • Vegetation transition from meadow to forest suppressed the priming effect. • P availability and microbial community shifts are important drivers for the priming effect. [ABSTRACT FROM AUTHOR]

Nitrification acidifies soil, and the produced H+ are neutralized by inorganic carbon (C) in soil leading to irreversible CO 2 emissions. CO 2 released by nitrogen (N) fertilizer-induced acidification is partitioned between solid (CaCO 3 re-precipitation), liquid (dissolved HCO 3 − and CO 3 2−) and gaseous (CO 2) phases. Therefore, quantifying the effects of N fertilization on CO 2 emissions from soil inorganic C is an enormous challenge. 14C-labeled CaCO 3 was used as a model inorganic C to trace the released CO 2 caused by acidification by five fertilizers: chicken manure, urea, KNO 3 , NH 4 NO 3 , and (NH 4) 2 SO 4 added at three N rates. Cropland soil was homogenously mixed with Ca14CO 3 powder and fertilizers, and the emitted CO 2 was trapped in NaOH solution to determine total CO 2 and 14CO 2 efflux originated from inorganic C. Fertilization, particularly ammonium-based fertilizers ((NH 4) 2 SO 4 , NH 4 NO 3), strongly decreased soil pH by 0.35 units over 40 days. All fertilizers except KNO 3 increased total CO 2 emissions by 21%–490% compared to the unfertilized control soil. The fertilization effects on cumulative 14CO 2 emission induced by CaCO 3 neutralization, corresponded to acidification and decreased in the order (NH 4) 2 SO 4 > NH 4 NO 3 > urea > KNO 3 > chicken manure. Ammonium-based fertilizers induced the strongest CO 2 emissions originated from inorganic C, emitting 1.6–4.5 times more 14CO 2 than non-fertilized soils during the first nine days. The total CO 2 emissions from SIC were proportional to the fertilizer dose applied. Therefore, we conclude that both the choice of N fertilizers and their application rates need to be considered to control CO 2 emissions originated from inorganic C. The soil inorganic C losses should be prevented not only because of their irreversible contribution to atmospheric CO 2 , but also to safeguard ecosystem services of CaCO 3 , such as organic matter preservation, soil structure stabilization, and C sequestration. • 14C-labeled CaCO 3 applied to quantify CO 2 emission from soil inorganic carbon (SIC). • CO 2 originated from SIC (CO 2-SIC) increases with nitrification rate of N fertilizers. • Theoretical estimations based on nitrification of N fertilizers overestimates CO 2-SIC emission from soil. • Ignoring the contribution of SIC to total soil CO 2 emission overestimates SOC-originated CO 2. [ABSTRACT FROM AUTHOR]

Alternate wetting-drying (AWD) in rice cultivation controls soil redox conditions and consequently nutrient solubility. Under low redox potential, ferric iron reduction leads to bound phosphate (Fe(III)–P) dissolution, but lack of oxygen retards organic phosphorus (P org) mineralization. Microorganisms accelerate P org mineralization as the redox potential increases during drying, but it is not known which P source will be preferentially taken up by microorganisms and plants. Using double 32/33P labeling, we traced for the first time the P released from inorganic (32P) and organic (33P) sources into microbial biomass P (MBP), phospholipids, and plants under continuous flooding (CF) and AWD. The CF rapidly induced reducing conditions that increased Fe–P dissolution and thus P availability. The AWD had no preference for 32P and 33P incorporation into plants. However, the ratios of 32P in roots to 32P in MBP or to 32P in phospholipids in rooted soil were 4–9 times higher than the respective ratios of 33P. Moreover, the ratios of 33P in roots to 33P in MBP or to 33P in phospholipids were always <0.6. Thus, plants and microorganisms were more competitive for P from Fe–P and from P org , respectively. The AWD increased 32P and 33P incorporation into phospholipids in rooted soil by 53–56% as compared to CF. Also, the ratio of 32P in phospholipids to 32P in MBP was 1.4–2.2 times higher under AWD than under CF. Thus, AWD stimulated microorganisms to allocate more P for cell membrane synthesis as compared to CF. Plants were similarly effective to take up P from inorganic and organic sources. Increased root biomass stimulated arbuscular mycorrhizal symbiosis (represented by 16:1ω5c) and thus 33P recovery in roots under AWD. In turn, the acceleration of microbial P turnover and the microbe-specific preference for P org highlight the importance of arbuscular mycorrhiza for plant P uptake under fluctuating water conditions. [Display omitted] • Continuous flooding increased reductive dissolution of Fe–P and thus P availability. • Microbes allocated more P for cell membrane synthesis under alternate wetting-drying. • Microorganisms in a maintenance state allocated more P for cell compound synthesis. • Rice plants were similarly effective to take up P from inorganic and organic sources. [ABSTRACT FROM AUTHOR]

Plastics accumulating in the environment, especially microplastics (defined as particles <5 mm), can lead to a range of problems and potential loss of ecosystem services. Polyhydroxyalkanoates (PHAs) are biodegradable plastics used in mulch films, and in packaging material to minimize plastic waste and to reduce soil pollution. Little is known, however, about the effect of microbioplastics on soil-plant interactions, especially soil microbial community structure and functioning in agroecosystems. For the first time, we combined zymography (to localize enzyme activity hotspots) with substrate-induced growth respiration to investigate the effect of PHAs addition on soil microbial community structure, growth, and exoenzyme kinetics in the microplastisphere (i.e. interface between soil and microplastic particles) compared to the rhizosphere and bulk soil. We used a common PHAs biopolymer, poly (3-hydroxybutyrate- co -3-hydroxyvalerate) (PHBV) and showed that PHBV was readily used by the microbial community as a source of carbon (C) resulting in an increased specific microbial growth rate and a more active microbial biomass in the microplastisphere in comparison to the bulk soil. Higher β-glucosidase and leucine aminopeptidase activities (0.6–5.0 times higher V max) and lower enzyme affinities (1.5–2.0 times higher K m) were also detected in the microplastisphere relative to the rhizosphere. Furthermore, the PHBV addition changed the soil bacterial community at different taxonomical levels and increased the alpha diversity, as well as the relative abundance of Acidobacteria and Verrucomicrobia phyla, compared to the untreated soils. Overall, PHBV addition created soil hotspots where C and nutrient turnover is greatly enhanced, mainly driven by the accelerated microbial biomass and activity. In conclusion, microbioplastics have the potential to alter soil ecological functioning and biogeochemical cycling (e.g., SOM decomposition). [Display omitted] • Microplastisphere (soil-MPs interface) is localized and visualized by zymography. • MPs stimulates microbial turnover and nutrient efficiency in microplastisphere. • MPs increases soil enzyme activity and shifts bacterial community to K -strategy. • MPs have the potential to alter soil functioning and biogeochemical cycling. [ABSTRACT FROM AUTHOR]

Root morphology reflects plant adaptations to phosphorus (P) deficiency. We hypothesized that changes in rice root morphology reflect P deficiency decrease after ferric iron (Fe(III))-bound phosphate (Fe–P) dissolution in low-redox paddy soils. We developed a novel in-situ 32P phosphor-imaging approach under flooding to estimate P uptake by rice roots released from Fe–P dissolution. 32P-labeled ferrihydrite (31 mg P kg−1) was supplied either (1) in polyamide mesh bags (30 μm mesh size) to prevent roots but not microorganisms from direct Fe–P mobilization, or (2) directly mixed with soil to enable roots and microorganisms unrestricted access to the Fe–P. The establishment of low redox conditions (E h values between −176 and −224 mV) drove the reductive dissolution of Fe–P. Rice root-derived organic acids alone were unable to control Fe–P dissolution, and Fe(III) reduction is predominately a microbially-mediated process. Direct root access to Fe–P raised both the number and mean diameter of crown roots and root tips, and increased P uptake by 149–231%. Crown root elongation rate, 32P activities along roots and root tips were 5–133% higher when roots directly accessed Fe–P compared to Fe–P excluded from roots in mesh bags. Iron accumulation on roots depended on the rice growth stage, but not on their access to Fe–P. Roots' access to Fe–P increased rice crown roots elongation and branching and increased P accessibility under P deficiency. [Display omitted] • Fe–P dissolution increased rice root growth and induced root morphological changes. • Increased root tip numbers raised P uptake by roots from the soil. • Organic acids exuded by rice roots were unable to control Fe–P dissolution. • Iron accumulation on root surface was independent of root's access to Fe–P. [ABSTRACT FROM AUTHOR]

Biogeochemical cycles of phosphorus (P) and iron (Fe) are tightly interlinked, especially in highly weathered acidic subtropical and tropical soils rich in iron (oxyhydr)oxides (Fe(III)). The quantitative contribution of the reductive dissolution of Fe(III)-bound inorganic P (P i) (Fe–P) in low-redox paddy soils may cover the demands of rice plants (Oryza sativa L.) and microorganisms. We hypothesized that microbially-driven Fe(III) reduction and dissolution can cover the P demand of microorganisms but not of the young rice plants when the plants' P demand is high but their root systems are not sufficiently developed. We grew pre-germinated rice plants for 33 days in flooded rhizoboxes filled with a paddy soil of low P availability. 32P-labeled ferrihydrite (30.8 mg kg−1) was supplied either (1) in polyamide mesh bags (30 μm mesh size) to prevent roots from directly mobilizing Fe–P (pellets-in-mesh bag treatment), or (2) in the same pellet form but without a mesh bag to enable roots accessing the Fe–P (pellets-no-mesh bag treatment). Without mesh bags, P i was more available leading to increases in microbial biomass carbon (MBC) by 18–55% and nitrogen (MBN) by 4–108% in rooted soil as compared to P i pellets not directly available to roots. The maximum enzyme activities (V max) of phosphomonoesterase and β-glucosidase followed this pattern. During rice root growth, day 10 to day 33, MBC and microbial biomass phosphorus (MBP) contents in both rooted and bottom bulk (15–18 cm) soil gradually decreased by 28–56% and 47–49%, respectively. In contrast to our hypothesis, the contribution of Fe–P to MBP remarkably decreased from 4.5% to almost zero from 10 to 33 days after rice transplantation, while Fe–P compensated up to 16% of the plant P uptake at 33 days after rice transplantation, thus outcompeting microorganisms. Although both plants and microorganisms obtained P i released by Fe(III) reductive dissolution, this mechanism was not sufficient for the demand of either organism groups. [Display omitted] • 32P application can assess plant-microbial competitive dynamics for P absorbed by Fe. • Contribution of Fe–P to MBP decreased from 4.5% to almost zero with rice growth. • Fe–P compensated up to 16% of rice P uptake during 33 days after rice transplantation. • Rice plants outcompeted microorganisms for P at the tillering stage. [ABSTRACT FROM AUTHOR]

Plants stimulate microbial enzyme production in the rhizosphere, regulating soil organic matter decomposition and nutrient cycling. The availability of labile organic compounds (i.e. exudates) and water is the main prerequisite for such microbial activity and enzyme production, thus shaping the rhizosphere. Root morphology (i.e., root hairs) and exudate composition define the spatial distribution of properties and functions in the rhizosphere. However, the role of root architecture and exudate composition in this spatial self-organization of the rhizosphere remains unknown. We coupled three in situ imaging approaches: 14C imaging to localize exudates, soil zymography to analyze enzyme activity distribution, and neutron radiography for water fluxes to trace the spatial structure of the rhizosphere of three maize genotypes (wild-type, mutant with defective root-hair prolongation rth3 , and mutant with reduced benzoxazinoid content in root exudates bx1). The co-localization of these three soil images revealed the pivotal role of both optimal water content (neutron radiography) and root exudation (14C imaging) for β-glucosidase production by the rhizosphere microbiome and its hydrolytic activity (zymography). Root hairs increased the exudate release and enlarged the spatial extent of increased β - glucosidase activity around the root axis by 35%, leading to a two-fold faster decomposition of 14C exudates compared to the mutant with defective root hairs. In contrast, benzoxazinoids suppressed β - glucosidase activity by 30%, reflecting decreased microbial activity, whereas their absence broadened the rhizosphere. Overall, root hairs in wild-type maize increased microbial activity (i.e. β - glucosidase production), whereas the benzoxazinoids in root exudates suppressed microorganisms. [Display omitted] • Zymography, 14C imaging, and neutron radiography were coupled to localize rhizosphere processes. • Exudates are released at root tip in wild-type maize, but along the whole root in rth3 and bx1 mutants. • Root hairs enlarged the spatial extent of increased β - glucosidase activity around the root by 35%. • Benzoxazinoids (secondary metabolites) suppressed β - glucosidase activity in the rhizosphere by 30%. • Hotspot co-localization revealed that exudates and water increased β - glucosidase activity. [ABSTRACT FROM AUTHOR]

Despite the importance of subsoil carbon (C) deposition by deep-rooted crops in mitigating climate change and maintaining soil health, the quantification of root C input and its microbial utilization and stabilization below 1 m depth remains unexplored. We studied C input by three perennial deep-rooted plants (lucerne, kernza, and rosinweed) grown in a unique 4-m deep RootTower facility. 13C multiple pulse labeling was applied to trace C flows in roots, rhizodeposition, and soil as well as 13C incorporation into microbial groups by phospholipid fatty acids and the long-term stabilization of microbial residues by amino sugars. The ratio of rhizodeposited 13C in the PLFA and amino sugar pools was used to compare the relative microbial stability of rhizodeposited C across depths and plant species. Belowground C allocation between roots, rhizodeposits, and living and dead microorganisms indicated depth dependent plant investment. Rhizodeposition as a fraction of the total belowground C input declined from the topsoil (0–25 cm) to the deepest layer (360 cm), i.e., from 35%, 45%, and 36%–8.0%, 2.5%, and 2.7% for lucerne, kernza, and rosinweed, respectively, where lucerne had greater C input than the other species between 340 and 360 cm. The relative microbial stabilization of rhizodeposits in the subsoil across all species showed a dominance of recently assimilated C in microbial necromass, thus indicating a higher microbial stabilization of rhizodeposited C with depth. In conclusion, we traced photosynthates down to 3.6 m soil depth and showed that even relatively small C amounts allocated to deep soil layers will become microbially stabilized. Thus, deep-rooted crops, in particular lucerne are important for stabilization and storage of C over long time scales in deep soil. Image 1 • Rhizodeposition strongly declined with depth across all plant species. • Microbial stabilization of rhizodeposits increases with depth. • Deep-rooted crops are important for storage and stabilization of rhizodeposition. [ABSTRACT FROM AUTHOR]