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Rhizosphere wettability decreases upon severe drying leading to periods of prolonged low water content around roots after precipitation or irrigation. These observations were explained by the temporal hydrophobic character of mucilage, while structural alterations of the pore space caused by mucilage, such as pore clogging, remained mostly unexplored. In this study, time-series neutron radiography and a pore network model were used to assess the impact of pore geometry and wettability on water flow following the addition of mucilage in a sand substrate. To do so, we monitored the capillary rise of ethanol and water separately for mucilage contents, ranging from 0.0 to 2.0 mg g−1. A pore network model was developed to analyze the impact of alterations in pore geometry and wettability. Results are compared with analytical solutions of the Lucas-Washburn equation. Rewetting dynamics were explained by a combination of a decrease in effective pore throat size and a global decrease in wettability. The local distribution of wettability, however, appeared of minor importance as dynamics of water imbibition could be matched by a uniform effective contact angle. For 0.1 mg g−1 mucilage content increased wettability was predicted for both approaches: the analytical solution and the pore network model fit. At larger contents, a decrease in wettability occurred which was accompanied by a decrease in derived effective pore and pore throat size. On a minute scale, rewetting appeared to steadily progress at all mucilage contents with accelerated rewetting observed at 0.6 mg g−1 likely related to an increased wetting front length. This study highlights the importance of mucilage on pore geometry in combination with wettability modifications in the rhizosphere. Aside from rhizosphere rewetting, the presented approach provides an opportunity to investigate further wettability-related processes in other soil environments on various spatial scales., Geoderma, 437, ISSN:0016-7061, ISSN:1872-6259

Soils are considered the largest sink of microplastic (MP) in terrestrial ecosystems. However, little is known about the implications of MP on soil physical properties. We hypothesize that low wettability of MP induces soil water repellency, depending on MP content and size of MP and soil particles. We quantified wettability of mixtures of MP and sand. The sessile drop method (SDM) was applied to measure static contact angle (CA) of MP and glass beads at contents ranging from 0 to 100% (w/w). The results are extrapolated to varying combinations of MP and soil particle sizes based on specific surface area. Capillary rise was imaged with neutron radiography quantifying the effect of MP on dynamic CA, water imbibition, and water saturation distribution in sand. At 5% (w/w) MP content, static CA exhibited a steep increase to 80.2 degrees for MP 20-75 mu m and 59.7 degrees for MP 75-125 mu m. Dynamic CAs were approximately 40% lower than static CAs. Capillary rise experiments showed that MP 20-75 mu m reduced water imbibition into sand columns (700-1,200 mu m), with average dynamic CA of 40.3 degrees at 0.35% (w/w) MP content and 51.8 degrees at 1.05%. Decreased water saturation and increased tortuosity of flow paths were observed during imbibition peaking at 3.5% (w/w) local MP content. We conclude, in regions with high MP content. water infiltration and thus MP transport are hindered. Local low wettability induced by MP is expected to limit soil wettability and impede capillary rise., Vadose Zone Journal, 22 (1), ISSN:1539-1663

Soil water repellency is traditionally expressed as contact angle (CA) and measured destructively on exposed surfaces or derived from flow behavior or measured forces. These approaches cannot map local heterogeneities in CA, which typically exist in intact soil. Here, we explore the potential and limitations of in situ measurements of three-dimensional CAs with simplified benchmark tests and in intact sand. The method is based on a protocol for the freely available OpenFOAM software and applied to segmented X-ray computed tomography (X-ray CT) data. This study scrutinizes its suitability under typical vadose zone conditions, as it was originally developed for rock studies in petroleum engineering. The assets of the method are that it allows for accurate measurement of locally varying, equilibrium contact angles and that they can be linked to pore scale features that cause them. This is demonstrated with rehydrated mucilage structures in sand and the associated change in local CAs, where they touch sand grains. In general, the image acquisition time of polychromatic X-ray CT (minutes) precludes the assessments of dynamic CAs that may equilibrate within seconds. Another limitation is that acute CAs are overestimated due to the voxel discretization of interfaces and contact lines in combination with image smoothing. The divergence arises around 60 degrees and is most severe in the limit of vanishing water repellency. In summary, the method enables the mapping of local heterogeneities of equilibrium CAs, though the absolute values should be critically assessed., Vadose Zone Journal, 21 (3), ISSN:1539-1663

Mucilage is a gelatinous high-molecular-weight substance produced by almost all plants, serving numerous functions for plant and soil. To date, research has mainly focused on hydraulic and physical functions of mucilage in the rhizosphere. Studies on the relevance of mucilage as a microbial habitat are scarce. Extracellular polymeric substances (EPS) are similarly gelatinous high-molecular-weight substances produced by microorganisms. EPS support the establishment of microbial assemblages in soils, mainly through providing a moist environment, a protective barrier, and serving as carbon and nutrient sources. We propose that mucilage shares physical and chemical properties with EPS, functioning similarly as a biofilm matrix covering a large extent of the rhizosphere. Our analyses found no evidence of consistent differences in viscosity and surface tension between EPS and mucilage, these being important physical properties. With regard to chemical composition, polysaccharide, protein, neutral monosaccharide, and uronic acid composition also showed no consistent differences between these biogels. Our analyses and literature review suggest that all major functions known for EPS and required for biofilm formation are also provided by mucilage, offering a protected habitat optimized for nutrient mobilization. Mucilage enables high rhizomicrobial abundance and activity by functioning as carbon and nutrient source. We suggest that the role of mucilage as a biofilm matrix has been underestimated, and should be considered in conceptual models of the rhizosphere., Frontiers in Plant Science, 12, ISSN:1664-462X

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. ISSN:1539-1663

The soil offers numerous challenges to life residing in its porous environment. One of these challenges are fluctuations in soil water content which are accompanied by shifts in soil hydraulic properties. In order to avoid undesirable alterations and optimise growth conditions, plants and bacteria engineer their local environment by release of mucilage and EPS (extracellular polymeric substances). So far, modifications of soil properties were mainly attributed to the intrinsic properties of these highly polymeric blends. In this work, we focused on deriving a mechanistic understanding of how mucilage and EPS interact with the soil pore space and how these interactions impact soil hydraulic properties and water dynamics in the rhizosphere and other biological hotspots in soils. Mucilage and EPS are capable of absorbing large volumes of water, increase the viscosity of the soil solution and decrease its surface tension. Upon drying, mucilage turns water repellent. Here, we proposed a conceptual model linking the intrinsic physical properties of mucilage to their impact on soil hydrology. The increase in viscosity is related to the high content of polymers which can form an interconnected network. As the soil dries, mucilage and EPS become increasingly concentrated, the viscosity of the soil solution locally increases and its surface tension decreases. When a critical viscosity is reached and parts of the polymer network are adsorbed to drying surfaces, the retreat of the liquid front is delayed and its break-up due to capillary forces is prevented. This concept is confirmed by microscopy imaging and high resolution X-ray CT, which revealed that mucilage and EPS form filaments and two-dimensional structures in this process. Upon drying in porous media, mucilage at low concentrations (mass of dry gel per mass of dry soil) resulted in the formation of filaments. With increase in initial mucilage concentration, two-dimensional surfaces formed when the water content was relatively high and the liquid phase connected. Complementary measurements of soil hydraulic properties of mucilage amended soils showed how the formation of these continuous two-dimensional structures impacts soil physical properties, such as soil hydraulic conductivity, soil water retention and vapour diffusion. The maintained liquid connectivity in drying soils, which is caused by the high viscosity, low surface tension and interaction of the polymer network with the soil porous matrix, explains why the hydraulic conductivity of a mucilage amended sandy loam was higher at low soil water content when compared to its control, as shown in evaporation experiments. Additionally, the delayed retreat of the liquid phase at a critical mucilage concentration creates an additional matric (capillary) potential and enhances soil water retention. To separate and quantify this matric (capillary) effect from the intrinsic property of the polymer network to absorb water remains an open task. Furthermore, upon severe soil drying, the network of two-dimensional structures reduces vapour diffusion and thus delays soil drying. This effect was illustrated using time series neutron radiography to visualise the drying of mucilage amended sandy loam and a water saturated control. Besides affecting soil hydraulic properties and evaporation rates during soil drying, mucilage impacts the rewetting kinetics. Mucilage amended soils showed water repellency. Precisely, a sharp decrease in wettability was observed near mucilage contents at which one-dimensional structures were replaced by two-dimensional continuous surfaces. Simulation of water drop infiltration experiments in mucilage amended soils showed that the creation of continuous clusters of non-wettable pores induced a substantial decrease in soil wettability, indicated by a transition of water drop penetration time from milliseconds to minutes. Although most experiments presented here were based on simplified systems, such as mucilage amended porous media, we propose that the release of highly polymeric blends into the soil pore space represents a universal strategy of soil organisms. Plants and bacteria engineer the physical properties of their local environment in very similar and astoundingly effective ways. The mechanisms discovered in this thesis lead to hydraulic decoupling of biological hotspots (e.g. the rhizosphere or biocrust) and buffer the erratic fluctuations experienced by soil organisms in these microhydrological niches.

European Journal of Soil Science, 72 (1), ISSN:1351-0754, ISSN:1365-2389

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., Vadose Zone Journal, 18 (1), ISSN:1539-1663

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.