Nitrous Oxide Emission from Forage Plantain and Perennial Ryegrass Swards Is Affected by Belowground Resource Allocation Dynamics

Soil–plant interactions affecting nitrous oxide (N2O) are not well-understood, and experimental data are scarce. Therefore, a greenhouse experiment was conducted in a 3 × 3 full factorial design, comprising three mineral N fertilizer rates (0, 150 and 300 kg N ha−1) applied to monoculture swards and a binary mixture of Plantago lanceolata and Lolium perenne. The parameters measured included daily N2O emissions, aboveground (AG) and belowground biomass (BG), N and C yields, as well as leucine aminopeptidase (LAP) activity in the soil as an indicator for soil microbial activity. Nitrous oxide emission and LAP were measured using the static chamber method and fluorimetric microplate assays, respectively. Cumulative N2O emissions were about two times higher for P. lanceolata than L. perenne monoculture swards or the mixture (p < 0.05). The binary mixtures also showed the highest N use efficiency and LAP activity, which significantly (p < 0.05) correlated with the C concentration in the belowground biomass. Plantago lanceolata was generally ineffective at reducing N2O emissions, probably due to the young age of the swards. Among the biological factors, N2O emission was significantly associated with biomass productivity, belowground C yield, belowground N use efficiency and soil microbial activity. Thus, the results suggested belowground resource allocation dynamics as a possible means by which swards impacted N2O emission from the soils. However, a high N deposition might reduce the N2O mitigation potential of grasslands

Editorial: Rhizosphere Spatiotemporal Organisation

Formation of the rhizosphere, interface between living plant roots and soil, leads to changes in soil properties, nutrient and water distribution and biogeochemical cycling, and to a selection of unique populations of microorganisms and invertebrates. Dynamic feedback processes between the plant, the soil and the biota govern rhizosphere formation. The Frontiers Research Topic on “Rhizosphere Spatiotemporal Organization” presents contributions which aim to advance our understanding of rhizosphere processes. All of the six articles took the challenge to elaborate on the dynamic interactions and feedback processes in both spatial and temporal contexts

Keeping thinning-derived deadwood logs on forest floor improves soil organic carbon, microbial biomass, and enzyme activity in a temperate spruce forest

Deadwood is a key component of forest ecosystems, but there is limited information on how it influences forest soils. Moreover, studies on the effect of thinning-derived deadwood logs on forest soil properties are lacking. This study aimed to investigate the impact of thinning-derived deadwood logs on the soil chemical and microbial properties of a managed spruce forest on a loamy sand Podzol in Bavaria, Germany, after about 15 years. Deadwood increased the soil organic carbon contents by 59% and 56% at 0–4 cm and 8–12 cm depths, respectively. Under deadwood, the soil dissolved organic carbon and carbon to nitrogen ratio increased by 66% and 15% at 0–4 cm depth and by 55% and 28% at 8–12 cm depth, respectively. Deadwood also induced 71% and 92% higher microbial biomass carbon, 106% and 125% higher microbial biomass nitrogen, and 136% and 44% higher β-glucosidase activity in the soil at 0–4 cm and 8–12 cm depths, respectively. Many of the measured variables significantly correlated with soil organic carbon suggesting that deadwood modified the soil biochemical processes by altering soil carbon storage. Our results indicate the potential of thinned spruce deadwood logs to sequester carbon and improve the fertility of Podzol soils. This could be associated with the slow decay rate of spruce deadwood logs and low biological activity of Podzols that promote the accumulation of soil carbon. We propose that leaving thinning-derived deadwood on the forest floor can support soil and forest sustainability as well as carbon sequestration

Rhizosphere size and shape: Temporal dynamics and spatial stationarity

The soil volume affected by roots – the rhizosphere – is one of the most important microbial hotspots determining the processes, dynamics and cycling of carbon (C), nutrients and water in terrestrial ecosystems. Rhizosphere visualization is necessary to understand, localize and quantify the ongoing processes and functions, but quantitative conclusions are very uncertain because of: 1) the continuum of the parameters between the root surface and root-free soil, i.e., there are no sharp borders, 2) differences in the distributions of various parameters (C, nutrients, pH, enzyme and microbial activities, gases, water etc.) across and along roots, 3) temporal changes of the parameters and processes with root growth as well as with water and C flows. In situ techniques: planar optodes, zymography, sensitive gels, ¹⁴C and neutron imaging as well as destructive approaches (thin layer slicing) have been used to analyze the rhizosphere extent and the gradients of various physico-chemical and biological characteristics: pH, CO2, O2, redox potential, enzyme activities, content of water, nutrients and excess elements, and organic compounds. A literature analysis allows the conclusion that: i) the rhizosphere extent for most of the parameters assessed by non-destructive visualization techniques is 0.5–4 mm but exceeds 4 mm for gases, nitrate, water and redox potential. ii) The rhizosphere extent of nutrients (N, P) is decoupled from the extent of the corresponding enzyme activities. iii) The imbalance between element flows to and uptake by roots may lead to accumulation of excess elements and formation of root carapaces (e.g. CaCO3 rhizoliths, Fe plaque) ranging up to a few cm. iv) All destructive approaches show a much (3–5 times) larger rhizosphere extent compared to visualization techniques. These conclusions are crucial for a mechanistic understanding of rhizosphere properties and functioning, estimation of the nutrient stocks available to roots, and for rhizosphere modelling considering root growth and architecture. Overall, roots function as ecosystem engineers and build their environment, serving as the main factors shaping rhizosphere extent. Sharp gradients are formed within a few days for nutrients and enzymes, but more time is necessary for the establishment of specific microbial communities. Despite the very strong dynamics of most parameters, their stationarity is reached within a few days because the release of C and enzymes or nutrient uptake are very quickly compensated by utilization by surrounding microorganisms or/and sorption and diffusion processes. We conclude that despite the dynamic nature of each property, the rhizosphere gradients, their extent and shape are quasi-stationary because of the opposite directions of their formation processes

Contrasting distribution of enzyme activities in the rhizosphere of European beech and Norway spruce

Recent policies and silvicultural management call for forest regeneration that involve the selection of tree species able to cope with low soil nutrient availability in forest ecosystems. Understanding the impact of different tree species on the rhizosphere processes (e.g., enzyme activities) involved in nutrient mobilisation is critical in selecting suitable species to adapt forests to environmental change. Here, we visualised and investigated the rhizosphere distribution of enzyme activities (cellobiohydrolase, leucine-aminopeptidase, and acid phosphomonoesterase) using zymography. We related the distribution of enzyme activities to the seedling root morphological traits of European beech (Fagus sylvatica) and Norway spruce (Picea abies), the two most cultivated temperate tree species that employ contrasting strategies in soil nutrient acquisition. We found that spruce showed a higher morphological heterogeneity along the roots than beech, resulting in a more robust relationship between rhizoplane-associated enzyme activities and the longitudinal distance from the root apex. The rhizoplane enzyme activities decreased in spruce and increased in beech with the distance from the root apex over a power-law equation. Spruce revealed broader rhizosphere extents of all three enzymes, but only acid phosphomonoesterase activity was higher compared with beech. This latter result was determined by a larger root system found in beech compared with spruce that enhanced cellobiohydrolase and leucine-aminopeptidase activities. The root hair zone and hair lengths were significant variables determining the distribution of enzyme activities in the rhizosphere. Our findings indicate that spruce has a more substantial influence on rhizosphere enzyme production and diffusion than beech, enabling spruce to better mobilise nutrients from organic sources in heterogeneous forest soils

DataSheet_1_Contrasting distribution of enzyme activities in the rhizosphere of European beech and Norway spruce.docx

Recent policies and silvicultural management call for forest regeneration that involve the selection of tree species able to cope with low soil nutrient availability in forest ecosystems. Understanding the impact of different tree species on the rhizosphere processes (e.g., enzyme activities) involved in nutrient mobilisation is critical in selecting suitable species to adapt forests to environmental change. Here, we visualised and investigated the rhizosphere distribution of enzyme activities (cellobiohydrolase, leucine-aminopeptidase, and acid phosphomonoesterase) using zymography. We related the distribution of enzyme activities to the seedling root morphological traits of European beech (Fagus sylvatica) and Norway spruce (Picea abies), the two most cultivated temperate tree species that employ contrasting strategies in soil nutrient acquisition. We found that spruce showed a higher morphological heterogeneity along the roots than beech, resulting in a more robust relationship between rhizoplane-associated enzyme activities and the longitudinal distance from the root apex. The rhizoplane enzyme activities decreased in spruce and increased in beech with the distance from the root apex over a power-law equation. Spruce revealed broader rhizosphere extents of all three enzymes, but only acid phosphomonoesterase activity was higher compared with beech. This latter result was determined by a larger root system found in beech compared with spruce that enhanced cellobiohydrolase and leucine-aminopeptidase activities. The root hair zone and hair lengths were significant variables determining the distribution of enzyme activities in the rhizosphere. Our findings indicate that spruce has a more substantial influence on rhizosphere enzyme production and diffusion than beech, enabling spruce to better mobilise nutrients from organic sources in heterogeneous forest soils.</p

Image_1_Contrasting distribution of enzyme activities in the rhizosphere of European beech and Norway spruce.tif

Recent policies and silvicultural management call for forest regeneration that involve the selection of tree species able to cope with low soil nutrient availability in forest ecosystems. Understanding the impact of different tree species on the rhizosphere processes (e.g., enzyme activities) involved in nutrient mobilisation is critical in selecting suitable species to adapt forests to environmental change. Here, we visualised and investigated the rhizosphere distribution of enzyme activities (cellobiohydrolase, leucine-aminopeptidase, and acid phosphomonoesterase) using zymography. We related the distribution of enzyme activities to the seedling root morphological traits of European beech (Fagus sylvatica) and Norway spruce (Picea abies), the two most cultivated temperate tree species that employ contrasting strategies in soil nutrient acquisition. We found that spruce showed a higher morphological heterogeneity along the roots than beech, resulting in a more robust relationship between rhizoplane-associated enzyme activities and the longitudinal distance from the root apex. The rhizoplane enzyme activities decreased in spruce and increased in beech with the distance from the root apex over a power-law equation. Spruce revealed broader rhizosphere extents of all three enzymes, but only acid phosphomonoesterase activity was higher compared with beech. This latter result was determined by a larger root system found in beech compared with spruce that enhanced cellobiohydrolase and leucine-aminopeptidase activities. The root hair zone and hair lengths were significant variables determining the distribution of enzyme activities in the rhizosphere. Our findings indicate that spruce has a more substantial influence on rhizosphere enzyme production and diffusion than beech, enabling spruce to better mobilise nutrients from organic sources in heterogeneous forest soils.</p

Image_2_Contrasting distribution of enzyme activities in the rhizosphere of European beech and Norway spruce.tif

Recent policies and silvicultural management call for forest regeneration that involve the selection of tree species able to cope with low soil nutrient availability in forest ecosystems. Understanding the impact of different tree species on the rhizosphere processes (e.g., enzyme activities) involved in nutrient mobilisation is critical in selecting suitable species to adapt forests to environmental change. Here, we visualised and investigated the rhizosphere distribution of enzyme activities (cellobiohydrolase, leucine-aminopeptidase, and acid phosphomonoesterase) using zymography. We related the distribution of enzyme activities to the seedling root morphological traits of European beech (Fagus sylvatica) and Norway spruce (Picea abies), the two most cultivated temperate tree species that employ contrasting strategies in soil nutrient acquisition. We found that spruce showed a higher morphological heterogeneity along the roots than beech, resulting in a more robust relationship between rhizoplane-associated enzyme activities and the longitudinal distance from the root apex. The rhizoplane enzyme activities decreased in spruce and increased in beech with the distance from the root apex over a power-law equation. Spruce revealed broader rhizosphere extents of all three enzymes, but only acid phosphomonoesterase activity was higher compared with beech. This latter result was determined by a larger root system found in beech compared with spruce that enhanced cellobiohydrolase and leucine-aminopeptidase activities. The root hair zone and hair lengths were significant variables determining the distribution of enzyme activities in the rhizosphere. Our findings indicate that spruce has a more substantial influence on rhizosphere enzyme production and diffusion than beech, enabling spruce to better mobilise nutrients from organic sources in heterogeneous forest soils.</p