W. Richard Whalley

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.

The effect of soil strength on the yield of wheat

Although it is well-known that high soil strength is a constraint to root and shoot growth, it is not clear to what extent soil strength is the main physical stress that limits crop growth and yield. This is partly because it is difficult to separate the effects of soil drying and high soil strength, which tend to occur together. The aim of this paper is to test the hypothesis that for two different soil types, yield is closely related to soil strength irrespective of difference in soil water status and soil structure. Winter (Triticum aestivum L., cv. Hereward) and spring wheat (cv. Paragon) were grown in the field on two soils, which had very different physical characteristics. One was loamy sand and the other sandy clay loam; compaction and loosening treatments were applied in a fully factorial design to both. Crop growth and yield, carbon isotope discrimination, soil strength, water status, soil structure and hydraulic properties were measured. The results showed that irrespective of differences in soil type, structure and water status, soil strength gave a good prediction of crop yield. Comparison with previous data led to the conclusion that, irrespective of whether it was due to drying or compaction (poor soil management), soil strength appeared to be an important stress that limits crop productivity.

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.

Biological And Physical Processes That Mediate Micro-aggregation Of Clays

The aggregation of clays after the addition of organic materials is described. Clays were incubated with or without added organic matter in the form of grass, straw, or charcoal and needed to be dried to a water potential of −1.5 MPa or less to aggregate. It was the fine fraction of the clay (<0.5 μm) that aggregated after organic additions, and this was apparent even in clays or clay mixtures that had a broad particle size distribution. Grass was more effective than straw at aggregating the clay. When chloroform was added to the samples, there was little aggregation, suggesting that micro-aggregation after the addition of organic material to clay is mainly microbially mediated. The aggregation of clay in the presence of added substrate was temperature dependent, with an optimum between 20 and 30 °C, but in the absence of substrate aggregation increased with temperature without a maximum. This supports the hypothesis that biological mechanisms are playing an important role in aggregation.

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.

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.

Estimating soil strength in the rooting zone of wheat

When roots abstract water thus drying the soil, crop growth may be reduced by increasing strength of soil as well as the lack of water. Strong soil impedes root growth, restricting access to deeper water. As a result, there is a need to estimate soil strength in order to model crop response to dry soil correctly. The strength of soil can be routinely assessed with a penetrometer but measurements are time consuming and hard work to acquire at the frequency required to understand soil-water-plant relations. To make progress, a published relationship that derives penetrometer pressure from both water relations in soil and density was improved to take account of the effects of depth including the friction that results from the increasing hydrostatic pressure. These relationships were then incorporated into an agroecosystem model so that the dynamics of strong soil and its effect on wheat could be simulated. The combined model requires the moisture release curve (but this can be derived from other commonly-measured soil properties), daily rainfall, temperature, and potential evaporation and the agronomy of the crop. Modelled values of penetrometer pressure were simulated well compared with measured values in artificially strengthened (compacted) and weakened (irrigated) soils. Simulations of the strength of soil and the matric potential before anthesis are compared with measured total dry-matter yields of winter wheat in experimental fields. The results lend weight to the hypothesis that wheat yield is limited by the strength of soil in the field and that soil strength, rather than soil matric potential, better explains differences between soils.

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.

Improved Interpretation of Mercury Intrusion and Soil Water Retention Percolation Characteristics by Inverse Modelling and Void Cluster Analysis

This work addresses two continuing fallacies in the interpretation of percolation characteristics of porous solids. The first is that the first derivative (slope) of the intrusion characteristic of the non-wetting fluid or drainage characteristic of the wetting fluid corresponds to the void size distribution, and the second is that the sizes of all voids can be measured. The fallacies are illustrated with the aid of the PoreXpert® inverse modelling package. A new void analysis method is then described, which is an add-on to the inverse modelling package and addresses the second fallacy. It is applied to three widely contrasting and challenging porous media. The first comprises two fine-grain graphites for use in the next-generation nuclear reactors. Their larger void sizes were measured by mercury intrusion, and the smallest by using a grand canonical Monte Carlo interpretation of surface area measurement down to nanometre scale. The second application is to the mercury intrusion of a series of mixtures of ground calcium carbonate with powdered microporous calcium carbonate known as functionalised calcium carbonate (FCC). The third is the water retention/drainage characteristic of a soil sample which undergoes naturally occurring hydrophilic/hydrophobic transitions. The first-derivative approximation is shown to be reasonable in the interpretation of the mercury intrusion porosimetry of the two graphites, which differ only at low mercury intrusion pressures, but false for FCC and the transiently hydrophobic soil. The findings are supported by other experimental characterisations, in particular electron and atomic force microscopy.