Core Ideas Plant mucilage and bacterial extracellular polymeric substances (EPS) prevent the breakup of the soil liquid phase. Formation of continuous structures buffers soil hydraulic properties. The release of viscous polymeric substances represents a universal strategy. Plant roots and bacteria are capable of buffering erratic fluctuations of water content in their local soil environment by releasing a diverse, highly polymeric blend of substances (e.g. extracellular polymeric substances [EPS] and mucilage). Although this concept is well accepted, the physical mechanisms by which EPS and mucilage interact with the soil matrix and determine the soil water dynamics remain unclear. High‐resolution X‐ray computed tomography revealed that upon drying in porous media, mucilage (from maize [Zea mays L.] roots) and EPS (from intact biocrusts) form filaments and two‐dimensional interconnected structures spanning across multiple pores. Unlike water, these mucilage and EPS structures connecting soil particles did not break up upon drying, which is explained by the high viscosity and low surface tension of EPS and mucilage. Measurements of water retention and evaporation with soils mixed with seed mucilage show how these one‐ and two‐dimensional pore‐scale structures affect macroscopic hydraulic properties (i.e., they enhance water retention, preserve the continuity of the liquid phase in drying soils, and decrease vapor diffusivity and local drying rates). In conclusion, we propose that the release of viscous polymeric substances and the consequent creation of a network bridging the soil pore space represent a universal strategy of plants and bacteria to engineer their own soil microhydrological niches where stable conditions for life are preserved.

Biohydrogels such as microbial exudates and root‐derived mucilage are soil‐born cross‐linked polymers, able to form porous three‐dimensional networks during water uptake. The gel effect is the variation of soil properties, such as soil hydrology and soil structural stability, resulting from biohydrogel swelling in soil. Conventionally, soil–water–hydrogel interactions are investigated by measuring soil bulk properties such as water retention curves and porosity, without further analyzing the effect of biohydrogel phases in soil on a quantitative basis. Therefore, the evaluation of advanced and novel methods for the characterization of biohydrogel phases in soil and soil–water–hydrogel interactions is necessary. This review evaluates currently available methods for their potential to analyze processes associated to the gel effect. A promising approach to investigate the spatio‐temporal distribution of biohydrogel phases in porous media is based on Nuclear Magnetic Resonance (NMR) such as 1H‐NMR relaxometry, as well as on imaging techniques such as Environmental Scanning Electron Microscopy (ESEM). Especially NMR techniques enable the identification of different water populations based on their differences in the relaxation, and thus the mobility of water molecules in biohydrogels and non‐gel water in soil pores. Rheology measures the flow behavior of biohydrogels, providing information on the structural behavior of the hydrogel network and its gelling mechanism. Soil rheology further quantifies the effect of the biohydrogel phases on the interactions between soil particles, and thus the impact on soil microstructural stability. However, rheology does not elucidate the spatio‐temporal distribution and structural state of biohydrogel phases in soil. All in all, a systematic combination of rheology, NMR and suitable imaging methods seems promising and necessary in order to elucidate the still widely unknown gel effect in soil.

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Polygalacturonic acid (PGA) is considered as a model substance for mucilage to study mucilage–soil interactions, assuming that the gel formation mechanism of mucilage is comparable to the one of PGA. However, some studies question the accepted hypothesis, which states that, like for PGA, this mechanism relies on cross‐links between uronic acid and calcium for mucilage. The aim of this study was therefore to understand the influence of the abundance and degree of esterification of uronic acids and the influence of calcium on the gel formation mechanism in mucilage as compared to model substances. The mucilage used was from chia seeds, as it is easily available in great quantity and has gel properties shared by root mucilage. Results reported here demonstrate that, while the gel formation mechanism of PGA relied on specific cross‐links with calcium and led to heterogeneous gels, low‐methoxy pectin (LMP) formed homogeneous calcium gels also characterized by nonspecific ionic interactions with calcium. On the contrary, despite similar uronic acid content to LMP, chia seed mucilage was mostly governed by weak electrostatic interactions between entangled polymer chains, which conferred the gel poor water retention. Addition of calcium reduced repulsion and molecular expansion, resulting in a reduction of the water content in chia seed mucilage. Finally, the discrepancies between PGA, LMP and chia seed mucilage discredit the use of PGA as model for chia seed mucilage. Comparison with root mucilage is still needed. This study offers the keys for further mechanistic understanding on the influence of mucilage on soil properties.

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PurposeSimultaneously interacting small-scale rhizosphere processes determine emergent plant-scale behaviour, including growth, transpiration, nutrient uptake, soil carbon storage and transformation by microorganisms. Current advances in modelling and experimental methods open the path to unravel and link those processes.MethodsWe present a series of examples of state-of-the art simulations addressing this multi-scale, multi-process problem from a modelling point of view, as well as from the point of view of integrating newly available rhizosphere data and images.ResultsEach example includes a model that links scales and experimental data to set-up simulations that explain and predict spatial and temporal distribution of rhizodeposition as driven by root architecture development, soil structure, presence of root hairs, soil water content and distribution of soil water. Furthermore, two models explicitly simulate the impact of the rhizodeposits on plant nutrient uptake and soil microbial activity, respectively. This exemplifies the currently available state of the art modelling tools in this field: image-based modelling, pore-scale modelling, continuum scale modelling and functional-structural plant modelling. We further show how to link the pore scale to the continuum scale by homogenisation or by deriving effective physical parameters like viscosity from nano-scale chemical properties.ConclusionModelling allows to integrate and make use of new experimental data across different rhizosphere processes (and thus across different disciplines) and scales. Described models are tools to test hypotheses and consequently improve our mechanistic understanding of how rhizosphere processes impact plant-scale behaviour. Linking multiple scales and processes is the logical next step for future research.

Background: Gaseous matter exchanges in soil are determined by the connectivity of the pore system which is easily clogged by fresh root exudates. However, it remains unclear how a hydrogel (e.g., mucilage) affects soil pore tortuosity and gas diffusion properties when drying. Aims: The aim of this viewpoint study is to extend the understanding of gas exchange processes in the rhizosphere by (a) relating it to the patterns formed by drying mucilage within pore space and (b) to give a concept of the effect of drying mucilage on soil gas diffusivity using the combination of experimental evidence and simulations. Methods: To describe the effect of mucilage on soil gas exchanges, we performed gas diffusion experiments on dry soil–mucilage samples and took images of glass beads mixed with mucilage to visualize the formation of mucilage after drying, using Environmental Scanning Electron Microscopy. Finally, we set up simulations to characterize the geometric distribution of mucilage within soil during the drying process. Results: Experiments of gas diffusion show that mucilage decreases gas diffusion coefficient in dry soil without significantly altering bulk density and porosity. Electron microscopy indicates that during drying mucilage forms filaments and interconnected structures throughout the pore space reducing gas phase connectivity. The evolution of these geometric structures is explained via pore scale modelling based on identifying the elastic strength of rhizodeposition during soil drying. Conclusion: Our results suggest that releasing mucilage may be a plant adaption strategy to actively alter gas diffusion in soil.

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