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With current observations and future projections of more intense and frequent droughts in the tropics, understanding the impact that extensive dry periods may have on tree and ecosystem-level transpiration and concurrent carbon uptake has become increasingly important. Here, we investigate paired soil and tree water extraction dynamics in an old-growth upland forest in central Amazonia during the 2018 dry season. Tree water use was assessed via radial patterns of sap flow in eight dominant canopy trees, each a different species with a range in diameter, height, and wood density. Paired multi-sensor soil moisture probes used to quantify volumetric water content dynamics and soil water extraction within the upper 100 cm were installed adjacent to six of those trees. To link depth-specific water extraction patterns to root distribution, fine root biomass was assessed through the soil profile to 235 cm. To scale tree water use to the plot level (stand transpiration), basal area was measured for all trees within a 5 m radius around each soil moisture probe. The sensitivity of tree transpiration to reduced precipitation varied by tree, with some increasing and some decreasing in water use during the dry period. Tree-level water use scaled with sapwood area, from 11 to 190 L per day. Stand level water use, based on multiple plots encompassing sap flow and adjacent trees, varied from ∼1.7 to 3.3 mm per day, increasing linearly with plot basal area. Soil water extraction was dependent on root biomass, which was dense at the surface (i.e., 45% in the upper 5 cm) and declined dramatically with depth. As the dry season progressed and the upper soil dried, soil water extraction shifted to deeper levels and model projections suggest that much of the water used during the month-long dry-down could be extracted from the upper 2–3 m. Results indicate variation in rates of soil water extraction across the research area and, temporally, through the soil profile. These results provide key information on whole-tree contributions to transpiration by canopy trees as water availability changes. In addition, information on simultaneous stand level dynamics of soil water extraction that can inform mechanistic models that project tropical forest response to drought. [ABSTRACT FROM AUTHOR]

With current observations and future projections of more intense and frequent droughts in the tropics, understanding the impact that extensive dry periods may have on tree and ecosystem-level transpiration and concurrent carbon uptake has become increasingly important. Here, we investigate paired soil and tree water extraction dynamics in an old-growth upland forest in central Amazonia during the 2018 dry season. Tree water use was assessed via radial patterns of sap flow in eight dominant canopy trees, each a different species with a range in diameter, height, and wood density. Paired multi-sensor soil moisture probes used to quantify volumetric water content dynamics and soil water extraction within the upper 100 cm were installed adjacent to six of those trees. To link depth-specific water extraction patterns to root distribution, fine root biomass was assessed through the soil profile to 235 cm. To scale tree water use to the plot level (stand transpiration), basal area was measured for all trees within a 5 m radius around each soil moisture probe. The sensitivity of tree transpiration to reduced precipitation varied by tree, with some increasing and some decreasing in water use during the dry period. Tree-level water use scaled with sapwood area, from 11 to 190 L per day. Stand level water use, based on multiple plots encompassing sap flow and adjacent trees, varied from ∼1.7 to 3.3 mm per day, increasing linearly with plot basal area. Soil water extraction was dependent on root biomass, which was dense at the surface (i.e., 45% in the upper 5 cm) and declined dramatically with depth. As the dry season progressed and the upper soil dried, soil water extraction shifted to deeper levels and model projections suggest that much of the water used during the month-long dry-down could be extracted from the upper 2–3 m. Results indicate variation in rates of soil water extraction across the research area and, temporally, through the soil profile. These results provide key information on whole-tree contributions to transpiration by canopy trees as water availability changes. In addition, information on simultaneous stand level dynamics of soil water extraction that can inform mechanistic models that project tropical forest response to drought.

Soil hydraulic conductivity (ksoil) drops significantly in dry soils, resulting in steep soil water potential gradients (ψs) near plant roots during water uptake. Coarse soil grid resolutions in root system scale (RSS) models of root water uptake (RWU) generally do not spatially resolve this gradient in drying soils which can lead to a large overestimation of RWU. To quantify this, we consider a benchmark scenario of RWU from drying soil for which a numerical reference solution is available. We analyze this problem using a finite volume scheme and investigate the impact of grid size on the RSS model results. At dry conditions, the cumulative RWU was overestimated by up to 300% for the coarsest soil grid of 4.0 cm and by 30% for the finest soil grid of 0.2 cm, while the computational demand increased from 19 s to 21 h. As an accurate and computationally efficient alternative to the RSS model, we implemented a continuum multi-scale model where we keep a coarse grid resolution for the bulk soil, but in addition, we solve a 1-dimensional radially symmetric soil model at rhizosphere scale around individual root segments. The models at the two scales are coupled in a mass-conservative way. The multi-scale model compares best to the reference solution (−20%) at much lower computational costs of 4 min. Our results demonstrate the need to shift to improved RWU models when simulating dry soil conditions and highlight that results for dry conditions obtained with RSS models of RWU should be interpreted with caution.

Soil hydraulic conductivity (k soil ) drops significantly in dry soils, resulting in steep soil water potential gradients (ψ s ) near plant roots during water uptake. Coarse soil grid resolutions in root system scale (RSS) models of root water uptake (RWU) generally do not spatially resolve this gradient in drying soils which can lead to a large overestimation of RWU. To quantify this, we consider a benchmark scenario of RWU from drying soil for which a numerical reference solution is available. We analyze this problem using a finite volume scheme and investigate the impact of grid size on the RSS model results. At dry conditions, the cumulative RWU was overestimated by up to 300% for the coarsest soil grid of 4.0 cm and by 30% for the finest soil grid of 0.2 cm, while the computational demand increased from 19 s to 21 h. As an accurate and computationally efficient alternative to the RSS model, we implemented a continuum multi-scale model where we keep a coarse grid resolution for the bulk soil, but in addition, we solve a 1-dimensional radially symmetric soil model at rhizosphere scale around individual root segments. The models at the two scales are coupled in a mass-conservative way. The multi-scale model compares best to the reference solution (−20%) at much lower computational costs of 4 min. Our results demonstrate the need to shift to improved RWU models when simulating dry soil conditions and highlight that results for dry conditions obtained with RSS models of RWU should be interpreted with caution. [ABSTRACT FROM AUTHOR]

Recent studies have identified soil drying as a dominant driver of transpiration reduction at the global scale. Although Arbuscular Mycorrhiza Fungi (AMF) are assumed to play a pivotal role in plant response to soil drying, studies investigating the impact of AMF on plant water status and soil-plant hydraulic conductance are lacking. Thus, the main objective of this study was to investigate the influence of AMF on soil-plant conductance and plant water status of tomato under drought. We hypothesized that AMF limit the drop in matric potential across the rhizosphere, especially in drying soil. The underlying mechanism is that AMF extend the effective root radius and hence reduce the water fluxes at the root-soil interface. The follow-up hypothesis is that AMF enhance soil-plant hydraulic conductance and plant water status during soil drying. To test these hypotheses, we measured the relation between transpiration rate, soil and leaf water potential of tomato with reduced mycorrhiza colonization (RMC) and the corresponding wild type (WT). We inoculated the soil of the WT with Rhizophagus irregularis spores to potentially upsurge symbiosis initiation. During soil drying, leaf water potential of the WT did not drop below −0.8MPa during the first 6days after withholding irrigation, while leaf water potential of RMC dropped below −1MPa already after 4days. Furthermore, AMF enhanced the soil-plant hydraulic conductance of the WT during soil drying. In contrast, soil-plant hydraulic conductance of the RMC declined more abruptly as soil dried. We conclude that AMF maintained the hydraulic continuity between root and soil in drying soils, hereby reducing the drop in matric potential at the root-soil interface and enhancing soil-plant hydraulic conductance of tomato under edaphic stress. Future studies will investigate the role of AMF on soil-plant hydraulic conductance and plant water status among diverse plant species growing in contrasting soil textures.

Recent studies have identified soil drying as a dominant driver of transpiration reduction at the global scale. Although Arbuscular Mycorrhiza Fungi (AMF) are assumed to play a pivotal role in plant response to soil drying, studies investigating the impact of AMF on plant water status and soil-plant hydraulic conductance are lacking. Thus, the main objective of this study was to investigate the influence of AMF on soil-plant conductance and plant water status of tomato under drought. We hypothesized that AMF limit the drop in matric potential across the rhizosphere, especially in drying soil. The underlying mechanism is that AMF extend the effective root radius and hence reduce the water fluxes at the root-soil interface. The follow-up hypothesis is that AMF enhance soil-plant hydraulic conductance and plant water status during soil drying. To test these hypotheses, we measured the relation between transpiration rate, soil and leaf water potential of tomato with reduced mycorrhiza colonization (RMC) and the corresponding wild type (WT). We inoculated the soil of the WT with Rhizophagus irregularis spores to potentially upsurge symbiosis initiation. During soil drying, leaf water potential of the WT did not drop below −0.8MPa during the first 6days after withholding irrigation, while leaf water potential of RMC dropped below −1MPa already after 4days. Furthermore, AMF enhanced the soil-plant hydraulic conductance of the WT during soil drying. In contrast, soil-plant hydraulic conductance of the RMC declined more abruptly as soil dried. We conclude that AMF maintained the hydraulic continuity between root and soil in drying soils, hereby reducing the drop in matric potential at the root-soil interface and enhancing soil-plant hydraulic conductance of tomato under edaphic stress. Future studies will investigate the role of AMF on soil-plant hydraulic conductance and plant water status among diverse plant species growing in contrasting soil textures. [ABSTRACT FROM AUTHOR]

Forest water use has been difficult to quantify. One promising approach is to measure the isotopic composition of plant water, e.g., the transpired water vapor or xylem water. Because different water sources, e.g., groundwater versus shallow soil water, often show different isotopic signatures, isotopes can be used to investigate the depths from which plants take up their water and how this changes over time. Traditionally such measurements have relied on the extraction of wood samples, which provide limited time resolution at great expense, and risk possible artifacts. Utilizing a borehole drilled through a tree's stem, we propose a new method based on the notion that water vapor in a slow-moving airstream approaches isotopic equilibration with the much greater mass of liquid water in the xylem. We present two empirical data sets showing that the method can work in practice. We then present a theoretical model estimating equilibration times and exploring the limits at which the approach will fail. The method provides a simple, cheap, and accurate means of continuously estimating the isotopic composition of the source water for transpiration.

Three-dimensional models of root growth, architecture and function are becoming important tools that aid the design of agricultural management schemes and the selection of beneficial root traits. However, while benchmarking is common in many disciplines that use numerical models, such as natural and engineering sciences, functional-structural root architecture models have never been systematically compared. The following reasons might induce disagreement between the simulation results of different models: different representation of root growth, sink term of root water and solute uptake and representation of the rhizosphere. Presently, the extent of discrepancies is unknown, and a framework for quantitatively comparing functional-structural root architecture models is required. We propose, in a first step, to define benchmarking scenarios that test individual components of complex models: root architecture, water flow in soil and water flow in roots. While the latter two will focus mainly on comparing numerical aspects, the root architectural models have to be compared at a conceptual level as they generally differ in process representation. Therefore, defining common inputs that allow recreating reference root systems in all models will be a key challenge. In a second step, benchmarking scenarios for the coupled problems are defined. We expect that the results of step 1 will enable us to better interpret differences found in step 2. This benchmarking will result in a better understanding of the different models and contribute toward improving them. Improved models will allow us to simulate various scenarios with greater confidence and avoid bugs, numerical errors or conceptual misunderstandings. This work will set a standard for future model development.

Forest water use has been difficult to quantify. One promising approach is to measure the isotopic composition of plant water, e.g., the transpired water vapor or xylem water. Because different water sources, e.g., groundwater versus shallow soil water, often show different isotopic signatures, isotopes can be used to investigate the depths from which plants take up their water and how this changes over time. Traditionally such measurements have relied on the extraction of wood samples, which provide limited time resolution at great expense, and risk possible artifacts. Utilizing a borehole drilled through a tree's stem, we propose a new method based on the notion that water vapor in a slow-moving airstream approaches isotopic equilibration with the much greater mass of liquid water in the xylem. We present two empirical data sets showing that the method can work in practice. We then present a theoretical model estimating equilibration times and exploring the limits at which the approach will fail. The method provides a simple, cheap, and accurate means of continuously estimating the isotopic composition of the source water for transpiration. [ABSTRACT FROM AUTHOR]

Three-dimensional models of root growth, architecture and function are becoming important tools that aid the design of agricultural management schemes and the selection of beneficial root traits. However, while benchmarking is common in many disciplines that use numerical models, such as natural and engineering sciences, functional-structural root architecture models have never been systematically compared. The following reasons might induce disagreement between the simulation results of different models: different representation of root growth, sink term of root water and solute uptake and representation of the rhizosphere. Presently, the extent of discrepancies is unknown, and a framework for quantitatively comparing functional-structural root architecture models is required. We propose, in a first step, to define benchmarking scenarios that test individual components of complex models: root architecture, water flow in soil and water flow in roots. While the latter two will focus mainly on comparing numerical aspects, the root architectural models have to be compared at a conceptual level as they generally differ in process representation. Therefore, defining common inputs that allow recreating reference root systems in all models will be a key challenge. In a second step, benchmarking scenarios for the coupled problems are defined. We expect that the results of step 1 will enable us to better interpret differences found in step 2. This benchmarking will result in a better understanding of the different models and contribute toward improving them. Improved models will allow us to simulate various scenarios with greater confidence and avoid bugs, numerical errors or conceptual misunderstandings. This work will set a standard for future model development. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi (AMF) proliferate in soil pores, on the surface of soil particles and affect soil structure. Although modifications in substrate moisture retention depend on structure and could influence plant water extraction, mycorrhizal impacts on water retention and hydraulic conductivity were rarely quantified. Hence, we asked whether inoculation with AMF affects substrate water retention, water transport properties and at which drought intensity those factors become limiting for plant transpiration. Solanum lycopersicum plants were set up in the glasshouse, inoculated or not with Funneliformis mosseae, and grown for 35 days under ample water supply. After mycorrhizal establishment, we harvested three sets of plants, one before (36 days after inoculation) and the second (day 42) and third (day 47) within a sequential drying episode. Sampling cores were introduced into pots before planting. After harvest, moisture retention and substrate conductivity properties were assessed and water retention and hydraulic conductivity models were fitted. A root water uptake model was adopted in order to identify the critical substrate moisture that induces soil derived transpiration limitation. Neither substrate porosity nor saturated water contents were affected by inoculation, but both declined after substrates dried. Drying also caused a decline in pot water capacity and hydraulic conductivity. Plant available water contents under wet (pF 1.8-4.2) and dry (pF 2.5-4.2) conditions increased in mycorrhizal substrates and were conserved after drying. Substrate hydraulic conductivity was higher in mycorrhizal pots before and during drought exposure. After withholding water from pots, higher substrate drying rates and lower substrate water potentials were found in mycorrhizal substrates. Mycorrhiza neither affected leaf area nor root weight or length. Consistently with higher substrate drying rates, AMF restored the plant hydraulic status, and increased plant transpiration when soil moisture declined. The water potential at the root surface and the resistance to water flow in the rhizosphere were restored in mycorrhizal pots although the bulk substrate dried more. Finally, substrates colonized by AMF can be more desiccated before substrate water flux quantitatively limits transpiration. This is most pronounced under high transpiration demands and complies with a difference of over 1,000 hPa in substrate water potential. [ABSTRACT FROM AUTHOR]

Split root experiments have the potential to disentangle water transport in roots and soil, enabling the investigation of the water uptake pattern of a root system. Interpretation of the experimental data assumes that water flow between the split soil compartments does not occur. Another approach to investigate root water uptake is by numerical simulations combining soil and root water flow depending on the parameterization and description of the root system. Our aim is to demonstrate the synergisms that emerge from combining split root experiments with simulations. We show how growing root architectures derived from temporally repeated X-ray CT scanning can be implemented in numerical soil-plant models. Faba beans were grown with and without split layers and exposed to a single drought period during which plant and soil water status were measured. Root architectures were reconstructed from CT scans and used in the model R-SWMS (root-soil water movement and solute transport) to simulate water potentials in soil and roots in 3D as well as water uptake by growing roots in different depths. CT scans revealed that root development was considerably lower with split layers compared to without. This coincided with a reduction of transpiration, stomatal conductance and shoot growth. Simulated predawn water potentials were lower in the presence of split layers. Simulations showed that this was caused by an increased resistance to vertical water flow in the soil by the split layers. Comparison between measured and simulated soil water potentials proved that the split layers were not perfectly isolating and that redistribution of water from the lower, wetter compartments to the drier upper compartments took place, thus water losses were not equal to the root water uptake from those compartments. Still, the layers increased the resistance to vertical flow which resulted in lower simulated collar water potentials that led to reduced stomatal conductance and growth.

Split root experiments have the potential to disentangle water transport in roots and soil, enabling the investigation of the water uptake pattern of a root system. Interpretation of the experimental data assumes that water flow between the split soil compartments does not occur. Another approach to investigate root water uptake is by numerical simulations combining soil and root water flow depending on the parameterization and description of the root system. Our aim is to demonstrate the synergisms that emerge from combining split root experiments with simulations. We show how growing root architectures derived from temporally repeated X-ray CT scanning can be implemented in numerical soil-plant models. Faba beans were grown with and without split layers and exposed to a single drought period during which plant and soil water status were measured. Root architectures were reconstructed from CT scans and used in the model R-SWMS (root-soil water movement and solute transport) to simulate water potentials in soil and roots in 3D as well as water uptake by growing roots in different depths. CT scans revealed that root development was considerably lower with split layers compared to without. This coincided with a reduction of transpiration, stomatal conductance and shoot growth. Simulated predawn water potentials were lower in the presence of split layers. Simulations showed that this was related to an increased resistance to vertical water flow in the soil by the split layers. Comparison between measured and simulated soil water potentials proved that the split layers were not perfectly isolating and that redistribution of water from the lower, wetter compartments to the drier upper compartments took place, thus water losses were not equal to the root water uptake from those compartments. Still, the layers increased the resistance to vertical flow which resulted in lower simulated collar water potentials that led to reduced stomatal conductance and growth. [ABSTRACT FROM AUTHOR]

As plant roots take up water and the soil dries, water depletion is expected to occur in the vicinity of roots, the so called rhizosphere. However, recent experiments showed that the rhizosphere of lupines was wetter than the bulk soil during the drying period. Surprisingly, the rhizosphere remained temporarily dry after irrigation. Such water dynamics in the rhizosphere can be explained by the drying/wetting dynamics of mucilage exuded by roots. The capacity of mucilage to hold large volumes of water at negative water potential may favor root water uptake. However, mucilage hydrophobicity after drying may temporarily limit the local water uptake after irrigation. The effects of such rhizosphere dynamics are not yet understood. In particular, it is not known how the rhizosphere dynamics vary along roots and as a function of soil water content. My hypothesis was that the rewetting rate of the rhizosphere is primarily function of root age. Neutron radiography was used to monitor how the rhizosphere water dynamics vary along the root systems of lupines during drying/wetting cycles of different duration. The radiographs showed a fast and almost immediate rewetting of the rhizosphere of the distal root segments, in contrast to a slow rewetting of the rhizosphere of the proximal segments. The rewetting rate of the rhizosphere was not function of the water content before irrigation, but it was function of time. It is concluded that rhizosphere hydrophobicity is not uniform along roots, but it covers only the older and proximal root segments, while the young root segments are hydraulically well connected to the soil. I included these rhizosphere dynamics in a microscopic model of root water uptake. In the model, the relation between water content and water potential in the rhizosphere is not unique and it varies over time, and the rewetting rate of the rhizosphere decreases with time. The rhisosphere variability seems an optimal adaptation strategy to increase the water upt

As plant roots take up water and the soil dries, water depletion is expected to occur in the vicinity of roots, the so called rhizosphere. However, recent experiments showed that the rhizosphere of lupines was wetter than the bulk soil during the drying period. Surprisingly, the rhizosphere remained temporarily dry after irrigation. Such water dynamics in the rhizosphere can be explained by the drying/wetting dynamics of mucilage exuded by roots. The capacity of mucilage to hold large volumes of water at negative water potential may favor root water uptake. However, mucilage hydrophobicity after drying may temporarily limit the local water uptake after irrigation. The effects of such rhizosphere dynamics are not yet understood. In particular, it is not known how the rhizosphere dynamics vary along roots and as a function of soil water content. My hypothesis was that the rewetting rate of the rhizosphere is primarily function of root age. Neutron radiography was used to monitor how the rhizosphere water dynamics vary along the root systems of lupines during drying/wetting cycles of different duration. The radiographs showed a fast and almost immediate rewetting of the rhizosphere of the distal root segments, in contrast to a slow rewetting of the rhizosphere of the proximal segments. The rewetting rate of the rhizosphere was not function of the water content before irrigation, but it was function of time. It is concluded that rhizosphere hydrophobicity is not uniform along roots, but it covers only the older and proximal root segments, while the young root segments are hydraulically well-connected to the soil. I included these rhizosphere dynamics in a microscopic model of root water uptake. In the model, the relation between water content and water potential in the rhizosphere is not unique and it varies over time, and the rewetting rate of the rhizosphere decreases with time. The rhisosphere variability seems an optimal adaptation strategy to increase the water uptake of young root segments, which possibly reached new available water, and partly disconnect the old root segments from the already depleted soil [ABSTRACT FROM AUTHOR]