Effects of root dominate over aboveground litter on soil microbial biomass in global forest ecosystems

Background Inputs of above- and belowground litter into forest soils are changing at an unprecedented rate due to continuing human disturbances and climate change. Microorganisms drive the soil carbon (C) cycle, but the roles of above- and belowground litter in regulating the soil microbial community have not been evaluated at a global scale. Methods Here, we conducted a meta-analysis based on 68 aboveground litter removal and root exclusion studies across forest ecosystems to quantify the roles of above- and belowground litter on soil microbial community and compare their relative importance. Results Aboveground litter removal significantly declined soil microbial biomass by 4.9% but root exclusion inhibited it stronger, up to 11.7%. Moreover, the aboveground litter removal significantly raised fungi by 10.1% without altering bacteria, leading to a 46.7% increase in the fungi-to-bacteria (F/B) ratio. Differently, root exclusion significantly decreased the fungi by 26.2% but increased the bacteria by 5.7%, causing a 13.3% decrease in the F/B ratio. Specifically, root exclusion significantly inhibited arbuscular mycorrhizal fungi, ectomycorrhizal fungi, and actinomycetes by 22.9%, 43.8%, and 7.9%, respectively. The negative effects of aboveground litter removal on microbial biomass increased with mean annual temperature and precipitation, whereas that of root exclusion on microbial biomass did not change with climatic factors but amplified with treatment duration. More importantly, greater effects of root exclusion on microbial biomass than aboveground litter removal were consistent across diverse forest biomes (expect boreal forests) and durations. Conclusions These data provide a global evidence that root litter inputs exert a larger control on microbial biomass than aboveground litter inputs in forest ecosystems. Our study also highlights that changes in above- and belowground litter inputs could alter soil C stability differently by shifting the microbial community structure in the opposite direction. These findings are useful for predicting microbe-mediated C processes in response to changes in forest management or climate.

Evidence of considerable C and N transfer from peas to cereals via direct root contact but not via mycorrhiza

Intercropping of legumes and cereals is an important management method for improving yield stability, especially in organic farming systems. However, knowledge is restricted on the relevance of different nutrient transfer pathways. The objective of the study was to quantify nitrogen (N) and carbon (C) transfer from peas to triticale by (1) direct root contact (= R), (2) arbuscular mycorrhizal fungi (AMF; = A), and (3) diffusion (= D). Pea (Pisum sativum cv. Frisson and P2) and triticale (Triticum × Secale cv. Benetto) plants as intercrop were grown for 105 days. Treatment ADR enabled all transfer paths between the two crops. Treatment AD with root exclusion enabled AMF and diffusion transfer between peas and triticale. Treatment A with a diffusion gap barrier only allowed AMF transfer. Pea plants were labelled every 14 days with a 13C glucose and 15N urea solution, using the cotton wick technique. Direct root contact resulted in the highest pea rhizodeposition and thus the largest absolute amounts of N and C transfer to triticale. Root exclusion generally changed composition of rhizodeposits from fine root residues towards root exudates. Pea plant-N consisted of 17% N derived from rhizodeposition (NdfR) in treatment ADR but only 8% in the treatments AD and A, independently of pea variety, whereas pea plant-C consisted of 13% C derived from rhizodeposition (CdfR), without pea variety and transfer path treatment effects. Averaging all transfer path treatments, 6.7% of NdfR and 2.7% of CdfR was transferred from Frisson and P2 to triticale plants. Approximately 90% of this NdfR was transferred by direct root contact from Frisson to triticale and only 10% by AMF, whereas only 55% of CdfR was transferred to triticale by direct root contact, 40% by AMF and 5% by diffusion. Similar percentages were transferred from mutant P2 to triticale. Root exclusion generally changed RD composition from fine root residues towards root exudates.

Hyphal colonization of Rhizophagus irregularis increases unsaturated hydraulic conductivity of a loamy sand distant from roots

<p>Hydraulic properties of mycorrhizal soils have rarely been reported and difficulties in directly assigning potential effects to hyphae of arbuscular mycorrhizal fungi (AMF) arise from other consequences of AMF being present, i.e. their influence on growth and water consumption rates of their host plants that both also influence soil hydraulic properties.</p><p>We assumed that the typical nylon meshes used for root-exclusion experiments in mycorrhizal research can provide a dynamic hydraulic barrier. It is expected that the uniform pore size of the rigid meshes causes a sudden hydraulic decoupling of the enmeshed inner volume from the surrounding soil as soon as the mesh pores become air-filled. Growing plants below the soil moisture threshold for hydraulic decoupling would minimize plant-size effects on root-exclusion compartments and allow for a more direct assignment of hyphal presence to modulations in soil hydraulic properties.</p><p>We carried out water retention and hydraulic conductivity measurements with two tensiometers introduced in two different heights in a cylindrical compartment (250 cm³) containing a loamy sand, either with or without the introduction of a 20 µm nylon mesh equidistantly between the tensiometers. Introduction of a mesh reduced hydraulic conductivity across the soil volumes by two orders of magnitude from 471 to 6 µm d<sup>-1</sup> at 20% volumetric water content.</p><p>We grew maize plants inoculated or not with Rhizophagus irregularis in the same soil in pots that contained root-exclusion compartments while maintaining 20% volumetric water content. When hyphae were present in the compartments, water potential and unsaturated hydraulic conductivity increased for a given water content compared to compartments free of hyphae. These differences increased with progressive soil drying.</p><p>We conclude that water extractability from soils distant to roots can be facilitated under dry conditions when AMF hyphae are present.</p><p> </p>

Effects of litter and root manipulations on soil carbon and nitrogen in a Schrenk’s spruce (Picea schrenkiana) forest

Plant detritus represents the major source of soil carbon (C) and nitrogen (N), and changes in its quantity can influence below-ground biogeochemical processes in forests. However, we lack a mechanistic understanding of how above- and belowground detrital inputs affect soil C and N in mountain forests in an arid land. Here, we explored the effects of litter and root manipulations (control (CK), doubled litter input (DL), removal of litter (NL), root exclusion (NR), and a combination of litter removal and root exclusion (NI)) on soil C and N concentrations, enzyme activity and microbial biomass during a 2-year field experiment. We found that DL had no significant effect on soil total organic carbon (SOC) and total nitrogen (TN) but significantly increased soil dissolved organic carbon (DOC), microbial biomass C, N and inorganic N as well as soil cellulase, phosphatase and peroxidase activities. Conversely, NL and NR reduced soil C and N concentrations and enzyme activities. We also found an increase in the biomass of soil bacteria, fungi and actinomycetes in the DL treatment, while NL reduced the biomass of gram-positive bacteria, gram-negative bacteria and fungi by 5.15%, 17.50% and 14.17%, respectively. The NR decreased the biomass of these three taxonomic groups by 8.97%, 22.11% and 21.36%, respectively. Correlation analysis showed that soil biotic factors (enzyme activity and microbial biomass) and abiotic factors (soil moisture content) significantly controlled the change in soil C and N concentrations (P < 0.01). In brief, we found that the short-term input of plant detritus could markedly affect the concentrations and biological characteristics of the C and N fractions in soil. The removal experiment indicated that the contribution of roots to soil nutrients is greater than that of the litter.

Sodium interception by xylem parenchyma and chloride recirculation in phloem may augment exclusion in the salt tolerant Pistacia genus: context for salinity studies on tree crops

Working in tandem with root exclusion, stems may provide salt-tolerant woody perennials with some additional capacity to restrict sodium (Na) and chloride (Cl) accumulation in leaves. The Pistacia genus, falling at the nexus of salt tolerance and human intervention, provided an ideal set of organisms for studying the influences of both variable root exclusion and potentially variable discontinuities at the bud union on stem processes. In three experiments covering a wide range of salt concentrations (0 to 150 mM NaCl) and tree ages (1, 2 and 10 years) as well as nine rootstock-scion combinations we show that proportional exclusion of both Na and Cl reached up to ~85% efficacy, but efficacy varied by both rootstock and budding treatment. Effective Na exclusion was augmented by significant retrieval of Na from the xylem sap, as evidenced by declines in the Na concentrations of both sap and wood tissue along the transpiration stream. However, while we observed little to no differences between the concentrations of the two ions in leaves, analogous declines in sap concentrations of Cl were not observed. We conclude that some parallel but separate mechanism must be acting on Cl to provide leaf protection from toxicity specific to this ion and suggest that this mechanism is recirculation of Cl in the phloem. The presented findings underline the importance of holistic assessments of salt tolerance in woody perennials. In particular, greater emphasis might be placed on the dynamics of salt sequestration in the significant storage volumes offered by the stems of woody perennials and on the potential for phloem discontinuity introduced with a bud/graft union.

Separation of soil respiration: a site-specific comparison of partition methods

Without accurate data on soil heterotrophic respiration (Rh), assessments of soil carbon (C) sequestration rate and C balance are challenging to produce. Accordingly, it is essential to determine the contribution of the different sources of the total soil CO2 efflux (Rs) in different ecosystems, but to date, there are still many uncertainties and unknowns regarding the soil respiration partitioning procedures currently available. This study compared the suitability and relative accuracy of five different Rs partitioning methods in a subtropical forest: (1) regression between root biomass and CO2 efflux, (2) lab incubations with minimally disturbed soil microcosm cores, (3) root exclusion bags with hand-sorted roots, (4) root exclusion bags with intact soil blocks and (5) soil δ13C–CO2 natural abundance. The relationship between Rh and soil moisture and temperature was also investigated. A qualitative evaluation table of the partition methods with five performance parameters was produced. The Rs was measured weekly from 3 February to 19 April 2017 and found to average 6.1 ± 0.3 MgCha-1yr-1. During this period, the Rh measured with the in situ mesh bags with intact soil blocks and hand-sorted roots was estimated to contribute 49 ± 7 and 79 ± 3 % of Rs, respectively. The Rh percentages estimated with the root biomass regression, microcosm incubation and δ13C–CO2 natural abundance were 54 ± 41, 8–17 and 61 ± 39 %, respectively. Overall, no systematically superior or inferior Rs partition method was found. The paper discusses the strengths and weaknesses of each technique with the conclusion that combining two or more methods optimizes Rh assessment reliability.

Separation of soil respiration; a site-specific comparison of partition methods

Without accurate data on soil heterotrophic respiration (Rh), assessments of soil carbon (C) sequestration rate and C balance are challenging to produce. Accordingly, it is essential to determine the contribution of the different sources of the total soil CO2 efflux (Rs) in different ecosystems, but to date, there are still many uncertainties and unknown regarding the soil respiration partitioning procedures currently available. This study compared the suitability and accuracy of five different Rh/Rs partitioning methods in a subtropical forest: (1) regression between root mass and root derived CO2; (2) root exclusion bags with intact soil blocks; (3) root exclusion bags with hand-sorted roots; (4) lab incubations with minimally disturbed soil microcosm cores; and (5) soil δ13C-CO2 natural abundance. The relationship between Rh and soil moisture and temperature was also investigated. A qualitative evaluation table of the partition methods with five performance parameters was produced. The Rs was measured weekly from February 3rd to April 19th 2017 and found to average 6.1 ± 0.3 Mg C ha−1 y−1. During this period, the Rh measured with the in-situ mesh bags with intact soil blocks and hand-sorted roots were estimated to contribute 49 ± 7 % and 79 ± 3 % of Rs respectively. The Rh percentage estimated with the root mass regression, microcosm incubation and δ13C-CO2 natural abundance were 54 ± 41 %, 8–17 % and 61 ± 39 % respectively. Overall, no systematically superior or inferior Rh/Rs partition method was found. The paper discusses the strengths and weaknesses of each technique with the conclusion that combining two or more methods optimizes Rh assessment reliability.