ABSTRACT
Soils are complex ecosystems and the pore-scale physical structure regulates key processes that support terrestrial life. These include maintaining an appropriate mixture of air and water in soil, nutrient cycling and carbon sequestration. There is evidence that this structure is not random, although the organizing mechanism is not known. Using X-ray microtomography and controlled microcosms, we provide evidence that organization of pore-scale structure arises spontaneously out of the interaction between microbial activity, particle aggregation and resource flows in soil. A simple computational model shows that these interactions give rise to self-organization involving both physical particles and microbes that gives soil unique material properties. The consequence of self-organization for the functioning of soil is determined using lattice Boltzmann simulation of fluid flow through the observed structures, and predicts that the resultant micro-structural changes can significantly increase hydraulic conductivity. Manipulation of the diversity of the microbial community reveals a link between the measured change in micro-porosity and the ratio of fungal to bacterial biomass. We suggest that this behaviour may play an important role in the way that soil responds to management and climatic change, but that this capacity for self-organization has limits.
Subject(s)
Bacterial Physiological Phenomena , Cell Communication , Ecosystem , Models, Biological , Soil Microbiology , Soil , Computer SimulationABSTRACT
Despite the importance of fungi in soil functioning they have received comparatively little attention, and our understanding of fungal interactions and communities is lacking. This study aims to combine a physiologically based model of fungal growth with digitized images of internal pore volume of samples of undisturbed soil from contrasting management practices to determine the effect of physical structure on fungal growth dynamics. We quantified pore geometries of the undisturbed-soil samples from two contrasting agricultural practices, conventionally plowed (chisel plow) (CT) and no till (NT), and from native-species vegetation land use on land that was taken out of production in 1989 (NS). Then we modeled invasion of a fungal species within the soil samples and evaluated the role of soil structure on the progress of fungal colonization of the soil pore space. The size of the studied pores was > or =110 microm. The dynamics of fungal invasion was quantified through parameters of a mathematical model fitted to the fungal invasion curves. Results indicated that NT had substantially lower porosity and connectivity than CT and NS soils. For example, the largest connected pore volume occupied 79% and 88% of pore space in CT and NS treatments, respectively, while it only occupied 45% in NT. Likewise, the proportion of pore space available to fungal colonization was much greater in NS and CT than in NT treatment, and the dynamics of the fungal invasion differed among the treatments. The relative rate of fungal invasion at the onset of simulation was higher in NT samples, while the invasion followed a more sigmoidal pattern with relatively slow invasion rates at the initial time steps in NS and CT samples. Simulations allowed us to elucidate the contribution of physical structure to the rates and magnitudes of fungal invasion processes. It appeared that fragmented pore space disadvantaged fungal invasion in soils under long-term no-till, while large connected pores in soils under native vegetation or in tilled agriculture promoted the invasion.
Subject(s)
Fungi/growth & development , Models, Biological , Soil Microbiology , Computer Simulation , Fungi/physiology , Mycelium/growth & developmentABSTRACT
Reliable techniques for screening large numbers of plants for root traits are still being developed, but include aeroponic, hydroponic and agar plate systems. Coupled with digital cameras and image analysis software, these systems permit the rapid measurement of root numbers, length and diameter in moderate (typically <1000) numbers of plants. Usually such systems are employed with relatively small seedlings, and information is recorded in 2D. Recent developments in X-ray microtomography have facilitated 3D non-invasive measurement of small root systems grown in solid media, allowing angular distributions to be obtained in addition to numbers and length. However, because of the time taken to scan samples, only a small number can be screened (typically <10 per day, not including analysis time of the large spatial datasets generated) and, depending on sample size, limited resolution may mean that fine roots remain unresolved. Although agar plates allow differences between lines and genotypes to be discerned in young seedlings, the rank order may not be the same when the same materials are grown in solid media. For example, root length of dwarfing wheat (Triticum aestivum L.) lines grown on agar plates was increased by ~40% relative to wild-type and semi-dwarfing lines, but in a sandy loam soil under well watered conditions it was decreased by 24-33%. Such differences in ranking suggest that significant soil environment-genotype interactions are occurring. Developments in instruments and software mean that a combination of high-throughput simple screens and more in-depth examination of root-soil interactions is becoming viable.
ABSTRACT
The slow dynamics of granular flow is studied as an extension of static granular problems, which, as a consequence of shaking or related regimes, can be studied by the methods of statistical mechanics. For packed (i.e. 'jammed'), hard and rough objects, kinetic energy is a minor and ignorable quantity, as is strain. Hence, in the static case, the stress equations need supplementing by 'missing equations' depending solely on configurations. These are in the literature; this paper extends the equilibrium studies to slow dynamics, claiming that the strain rate (which is a consequence of flow, not of elastic strain) takes the place of stress, and as before, the analogue of Stokes's equation has to be supplemented by new 'missing equations' which are derived and which depend only on configurations.
ABSTRACT
In particulate materials, such as emulsions and granular media, a "jammed" system results if particles are packed together so that all particles are touching their neighbours, provided the density is sufficiently high. This paper studies through experiment, theory and simulation, the forces that particles exert upon one another in such a jammed state. Confocal microscopy of a compressed polydisperse emulsion provides a direct 3D measurement of the dispersed phase morphology within the bulk of the sample. This allows the determination of the probability distribution of interdroplet forces, P(f) where f is the magnitude of the force, from local droplet deformations. In parallel, the simplest form of the Boltzmann equation for the probability of force distributions predicts P(f) to be of the form e(-f/p), where p is proportional to the mean force f for large forces. This result is in good agreement with experimental and simulated data.