An overlooked mechanism underlying the attenuated temperature response of soil heterotrophic respiration.

Biogeochemical reactions occurring in soil pore space underpin gaseous emissions measured at macroscopic scales but are difficult to quantify due to their complexity and heterogeneity. We develop a volumetric-average method to calculate aerobic respiration rates analytically from soil with microscopic soil structure represented explicitly. Soil water content in the model is the result of the volumetric-average of the microscopic processes, and it is nonlinearly coupled with temperature and other factors. Since many biogeochemical reactions are driven by oxygen (O2) which must overcome various resistances before reaching reactive microsites from the atmosphere, the volumetric-average results in negative feedback between temperature and soil respiration, with the magnitude of the feedback increasing with soil water content and substrate quality. Comparisons with various experiments show the model reproduces the variation of carbon dioxide emission from soils under different water content and temperature gradients, indicating that it captures the key microscopic processes underpinning soil respiration. We show that alongside thermal microbial adaptation, substrate heterogeneity and microbial turnover and carbon use efficiency, O2 dissolution and diffusion in water associated with soil pore space is another key explanation for the attenuated temperature response of soil respiration and should be considered in developing soil organic carbon models.

The role of plant species and soil condition in the structural development of the rhizosphere.

Roots naturally exert axial and radial pressures during growth, which alter the structural arrangement of soil at the root-soil interface. However, empirical models suggest soil densification, which can have negative impacts on water and nutrient uptake, occurs at the immediate root surface with decreasing distance from the root. Here, we spatially map structural gradients in the soil surrounding roots using non-invasive imaging, to ascertain the role of root growth in early stage formation of soil structure. X-ray computed tomography provided a means not only to visualize a root system in situ and in 3-D but also to assess the precise root-induced alterations to soil structure close to, and at selected distances away from the root-soil interface. We spatially quantified the changes in soil structure generated by three common but contrasting plant species (pea, tomato, and wheat) under different soil texture and compaction treatments. Across the three plant types, significant increases in porosity at the immediate root surface were found in both clay loam and loamy sand soils and not soil densification, the currently assumed norm. Densification of the soil was recorded, at some distance away from the root, dependent on soil texture and plant type. There was a significant soil texture × bulk density × plant species interaction for the root convex hull, a measure of the extent to which root systems explore the soil, which suggested pea and wheat grew better in the clay soil when at a high bulk density, compared with tomato, which preferred lower bulk density soils. These results, only revealed by high resolution non-destructive imagery, show that although the root penetration mechanisms can lead to soil densification (which could have a negative impact on growth), the immediate root-soil interface is actually a zone of high porosity, which is very important for several key rhizosphere processes occurring at this scale including water and nutrient uptake and gaseous diffusion.

Isotope fractionation factors controlling isotopocule signatures of soil-emitted N2O produced by denitrification processes of various rates.

RATIONALE: This study aimed (i) to determine the isotopic fractionation factors associated with N2O production and reduction during soil denitrification and (ii) to help specify the factors controlling the magnitude of the isotope effects. For the first time the isotope effects of denitrification were determined in an experiment under oxic atmosphere and using a novel approach where N2O production and reduction occurred simultaneously. METHODS: Soil incubations were performed under a He/O2 atmosphere and the denitrification product ratio [N2O/(N2 + N2O)] was determined by direct measurement of N2 and N2O fluxes. N2O isotopocules were analyzed by mass spectrometry to determine δ(18)O, δ(15)N and (15)N site preference within the linear N2O molecule (SP). An isotopic model was applied for the simultaneous determination of net isotope effects (η) of both N2O production and reduction, taking into account emissions from two distinct soil pools. RESULTS: A clear relationship was observed between (15)N and (18)O isotope effects during N2O production and denitrification rates. For N2O reduction, diverse isotope effects were observed for the two distinct soil pools characterized by different product ratios. For moderate product ratios (from 0.1 to 1.0) the range of isotope effects given by previous studies was confirmed and refined, whereas for very low product ratios (below 0.1) the net isotope effects were much smaller. CONCLUSIONS: The fractionation factors associated with denitrification, determined under oxic incubation, are similar to the factors previously determined under anoxic conditions, hence potentially applicable for field studies. However, it was shown that the η(18)O/η(15)N ratios, previously accepted as typical for N2O reduction processes (i.e., higher than 2), are not valid for all conditions.

A novel grass hybrid to reduce flood generation in temperate regions.

We report on the evaluation of a novel grass hybrid that provides efficient forage production and could help mitigate flooding. Perennial ryegrass (Lolium perenne) is the grass species of choice for most farmers, but lacks resilience against extremes of climate. We hybridised L. perenne onto a closely related and more stress-resistant grass species, meadow fescue Festuca pratensis. We demonstrate that the L. perenne × F. pratensis cultivar can reduce runoff during the events by 51% compared to a leading UK nationally recommended L. perenne cultivar and by 43% compared to F. pratensis over a two year field experiment. We present evidence that the reduced runoff from this Festulolium cultivar was due to intense initial root growth followed by rapid senescence, especially at depth. Hybrid grasses of this type show potential for reducing the likelihood of flooding, whilst providing food production under conditions of changing climate.

Roots, water, and nutrient acquisition: let's get physical.

Improved root water and nutrient acquisition can increase fertiliser use efficiency and is important for securing food production. Root nutrient acquisition includes proliferation, transporter function, exudation, symbioses, and the delivery of dissolved nutrients from the bulk soil to the root surface via mass flow and diffusion. The widespread adoption of simplified experimental systems has restricted consideration of the influence of soil symbiotic organisms and physical properties on root acquisition. The soil physical properties can directly influence root growth and explain some of the disparities obtained from different experimental systems. Turning this to an advantage, comparing results obtained with the same model plant Arabidopsis (Arabidopsis thaliana) in different systems, we can tease apart the specific effects of soil physical properties.

Water supply and not nitrate concentration determines primary root growth in Arabidopsis.

Understanding how root system architecture (RSA) adapts to changing nitrogen and water availability is important for improving acquisition. A sand rhizotron system was developed to study RSA in a porous substrate under tightly regulated nutrient supply. The RSA of Arabidopsis seedlings under differing nitrate (NO3⁻) and water supplies in agar and sand was described. The hydraulic conductivity of the root environment was manipulated by using altered sand particle size and matric potentials. Ion-selective microelectrodes were used to quantify NO3⁻ at the surface of growing primary roots in sands of different particle sizes. Differences in RSA were observed between seedlings grown on agar and sand, and the influence of NO3⁻ (0.1-10.0 mm) and water on RSA was determined. Primary root length (PRL) was a function of water flux and independent of NO3⁻. The percentage of roots with laterals correlated with water flux, whereas NO3⁻ supply was important for basal root (BR) growth. In agar and sand, the NO3⁻ activities at the root surface were higher than those supplied in the nutrient solution. The sand rhizotron system is a useful tool for the study of RSA, providing a porous growth environment that can be used to simulate the effects of hydraulic conductivity on growth.

Soil engineering in vivo: harnessing natural biogeochemical systems for sustainable, multi-functional engineering solutions.

Carbon sequestration, infrastructure rehabilitation, brownfields clean-up, hazardous waste disposal, water resources protection and global warming-these twenty-first century challenges can neither be solved by the high-energy consumptive practices that hallmark industry today, nor by minor tweaking or optimization of these processes. A more radical, holistic approach is required to develop the sustainable solutions society needs. Most of the above challenges occur within, are supported on, are enabled by or grown from soil. Soil, contrary to conventional civil engineering thought, is a living system host to multiple simultaneous processes. It is proposed herein that 'soil engineering in vivo', wherein the natural capacity of soil as a living ecosystem is used to provide multiple solutions simultaneously, may provide new, innovative, sustainable solutions to some of these great challenges of the twenty-first century. This requires a multi-disciplinary perspective that embraces the science of biology, chemistry and physics and applies this knowledge to provide multi-functional civil and environmental engineering designs for the soil environment. For example, can native soil bacterial species moderate the carbonate cycle in soils to simultaneously solidify liquefiable soil, immobilize reactive heavy metals and sequester carbon-effectively providing civil engineering functionality while clarifying the ground water and removing carbon from the atmosphere? Exploration of these ideas has begun in earnest in recent years. This paper explores the potential, challenges and opportunities of this new field, and highlights one biogeochemical function of soil that has shown promise and is developing rapidly as a new technology. The example is used to propose a generalized approach in which the potential of this new field can be fully realized.

Root water potential integrates discrete soil physical properties to influence ABA signalling during partial rootzone drying.

To investigate the influence of different growing substrates (two mineral, two organic) on root xylem ABA concentration ([ABA](root)) and the contribution of the drying root system to total sap flow during partial rootzone drying (PRD), sunflower (Helianthus annuus L.) shoots were grafted onto the root systems of two plants grown in separate pots. Sap flow through each hypocotyl was measured below the graft union when one pot ('wet') was watered and other ('dry') was not. Each substrate gave unique relationships between dry pot matric potential (Psi(soil)), volumetric water content ((v)) or penetrometer resistance (Q) and either the fraction of photoperiod sap flow from roots in drying soil or [ABA](root). However, decreased relative sap flow, and increased [ABA](root), from roots in drying soil varied with root water potential (Psi(root)) more similarly across a range of substrates. The gradient between Psi(soil) and Psi(root) was greater in substrates with high sand or peat proportions, which may have contributed to a more sensitive response of [ABA](root) to Psi(soil) in these substrates. Whole plant transpiration was most closely correlated with the mean Psi(soil) of both pots, and then with detached leaf xylem ABA concentration. Although Psi(root) best predicted decreased relative sap flow, and increased [ABA](root), from roots in drying soil across a range of substrates, the inaccessibility of this variable in field studies requires a better understanding of how measurable soil variables (Psi(soil), (v), Q) affect Psi(root).

Physical effects of soil drying on roots and crop growth.

The nature and effect of the stresses on root growth in crops subject to drying is reviewed. Drought is a complex stress, impacting on plant growth in a number of interacting ways. In response, there are a number of ways in which the growing plant is able to adapt to or alleviate these stresses. It is suggested that the most significant opportunity for progress in overcoming drought stress and increasing crop yields is to understand and exploit the conditions in soil by which plant roots are able to maximize their use of resources. This may not be straightforward, with multiple stresses, sometimes competing functions of roots, and conditions which impact upon roots very differently depending upon what soil, what depth or what stage of growth the root is at. Several processes and the interaction between these processes in soil have been neglected. It is our view that drought is not a single, simple stress and that agronomic practice which seeks to adapt to climate change must take account of the multiple facets of both the stress induced by insufficient water as well as other interacting stresses such as heat, disease, soil strength, low nutrient status, and even hypoxia. The potential for adaptation is probably large, however. The possible changes in stress as a result of the climate change expected under UK conditions are assessed and it appears possible that wet warm winters will impact on root growth as much if not more than dry warm summers.

Effects of soil compaction and irrigation on the concentrations of selenium and arsenic in wheat grains.

Arsenic is a known human carcinogen, whereas Se is essential to human health and human Se intake in some European countries has decreased in recent decades. There have been few reports on the potential effect of soil physical conditions on the uptake of Se and As by food crops. Field experiments were conducted to evaluate the effects of soil compaction and irrigation on Se and As concentrations in wheat grains in two seasons. Grain Se concentration varied from 10 to 115 microg kg(-1), whereas the concentration of As in grain was low (<20 microg kg(-1)). Irrigation significantly decreased grain Se concentration by 30-75%, and the effect was likely due to a dilution as a result of increased grain yield, a competition of sulfate added in the irrigation water on Se uptake and increased leaching losses. Soil compaction significantly decreased grain Se concentration in one season. In contrast, soil compaction increased grain As concentration significantly when the crop was irrigated. The contrasting effects of soil compaction on As and Se uptake are explained by the difference in their mobility in soil and the pathway of ion transport to roots. The observed effect on grain Se is significant for human and animal nutrition, because the concentration varied from sufficient to very low levels in response to the alternation of soil physical conditions in the field. Soil physical conditions have to be taken into account when assessing the bioavailability of the contaminant As or the essential nutrient Se.