Hydraulic Root Research Articles

Mechanistically derived macroscopic root water uptake functions: The α and ω of root water uptake functions

AbstractWater uptake by plant roots is an important component of the soil water balance. Predicting to what extent potential transpiration from the canopy, that is, transpiration demand, can be met by supply of water from the soil through the root system is crucial to simulate the actual transpiration and assess vegetation water stress. In models that simulate the dynamics of vertical soil water content profiles as a function of water fluxes and soil water potential gradients, the root water uptake (RWU) distribution is represented by macroscopic sink terms. We present RWU functions that calculate sink terms based on a mechanistic model of water flow in the soil–root system. Based on soil–root hydraulics, we define α‐supply functions representing the maximal uptake by the root system from a certain soil depth when the root collar water potential equals the wilting point, ω‐supply factors representing the maximal supply from the entire root system, and a critical ωc factor representing the potential transpiration demand. These functions and factors are subsequently used to calculate RWU distributions directly from potential transpiration or demand and the soil water potentials. Unlike currently used approaches, which define α‐stress functions and ω factors representing ratios of actual uptake to uptake demand, the supply‐based formulations can be derived directly from soil and root hydraulic properties and can represent processes like root hydraulic redistribution and hydraulic lift.

Protein kinase SnRK2.4 is a key regulator of aquaporins and root hydraulics in Arabidopsis.

Soil water uptake by roots is a key component of plant water homeostasis contributing to plant growth and survival under ever-changing environmental conditions. The water transport capacity of roots (root hydraulic conductivity; Lpr ) is mostly contributed by finely regulated Plasma membrane Intrinsic Protein (PIP) aquaporins. In this study, we used natural variation of Arabidopsis for the identification of quantitative trait loci (QTLs) contributing to Lpr . Using recombinant lines from a biparental cross (Cvi-0 x Col-0), we show that the gene encoding class 2 Sucrose-Non-Fermenting Protein kinase 2.4 (SnRK2.4) in Col-0 contributes to >30% of Lpr by enhancing aquaporin-dependent water transport. At variance with the inactive and possibly unstable Cvi-0 SnRK2.4 form, the Col-0 form interacts with and phosphorylates the prototypal PIP2;1 aquaporin at Ser121 and stimulates its water transport activity upon coexpression in Xenopus oocytes and yeast cells. Activation of PIP2;1 by Col-0 SnRK2.4 in yeast also requires its protein kinase activity and can be counteracted by clade A Protein Phosphatases 2C. SnRK2.4 shows all hallmarks to be part of core abscisic acid (ABA) signaling modules. Yet, long-term (>3 h) inhibition of Lpr by ABA possibly involves a SnRK2.4-independent inhibition of PIP2;1. SnRK2.4 also promotes stomatal aperture and ABA-induced inhibition of primary root growth. The study identifies a key component of Lpr and sheds new light on the functional overlap and specificity of SnRK2.4 with respect to other ABA-dependent or independent SnRK2s.

Soil Water Deficit Reduced Root Hydraulic Conductivity of Common Reed (Phragmites australis).

Alterations in root hydraulics in response to varying moisture conditions remain a subject of debate. In our investigation, we subjected common reeds (Phragmites australis) to a 45-day treatment with four distinct soil moisture levels. The findings unveiled that, in response to drought stress, the total root length, surface area, volume, and average diameter exhibited varying degrees of reduction. Anatomically, drought caused a reduction in root diameter (RD), cortex thickness (CT), vessel diameter (VD), and root cross-sectional area (RCA). A decrease in soil moisture significantly reduced both whole- and single-root hydraulic conductivity (Lpwr, Lpsr). The total length, surface area, volume, and average diameter of the reed root system were significantly correlated with Lpwr, while RD, CT, and RCA were significantly correlated with Lpsr. A decrease in soil moisture content significantly influenced root morphological and anatomical characteristics, which, in turn, altered Lpr, and the transcriptome results suggest that this may be associated with the variation in the expression of abscisic acid (ABA) and aquaporins (AQPs) genes. Our initial findings address a gap in our understanding of reed hydraulics, offering fresh theoretical insights into how herbaceous plants respond to external stressors.

Plasma membrane aquaporins regulate root hydraulic conductivity in the model plant Setaria viridis.

The high rate of productivity observed in panicoid crops is in part due to their extensive root system. Recently, green foxtail (Setaria viridis) has emerged as a genetic model system for panicoid grasses. Natural accessions of S. viridis originating from different parts of the world, with differential leaf physiological behavior, have been identified. This work focused on understanding the physiological and molecular mechanisms controlling root hydraulic conductivity and root-to-shoot gas exchange signaling in S. viridis. We identified 2 accessions, SHA and ZHA, with contrasting behavior at the leaf, root, and whole-plant levels. Our results indicated a role for root aquaporin (AQP) plasma membrane (PM) intrinsic proteins in the differential behavior of SHA and ZHA. Moreover, a different root hydraulic response to low levels of abscisic acid between SHA and ZHA was observed, which was associated with root AQPs. Using cell imaging, biochemical, and reverse genetic approaches, we identified PM intrinsic protein 1;6 (PIP1;6) as a possible PIP1 candidate that regulates radial root hydraulics and root-to-shoot signaling of gas exchange in S. viridis. In heterologous systems, PIP1;6 localized in the endoplasmic reticulum, and upon interaction with PIP2s, relocalization to the PM was observed. PIP1;6 was predominantly expressed at the root endodermis. Generation of knockout PIP1;6 plants (KO-PIP1;6) in S. viridis showed altered root hydraulic conductivity, altered gas exchange, and alteration of root transcriptional patterns. Our results indicate that PIPs are essential in regulating whole-plant water homeostasis in S. viridis. We conclude that root hydraulic conductivity and gas exchange are positively associated and are regulated by AQPs.

Natural variation of maize root hydraulic architecture underlies highly diverse water uptake capacities.

Plant water uptake is determined by the root system architecture and its hydraulic capacity, which together define the root hydraulic architecture. The current research aims at understanding the water uptake capacities of maize (Zea mays), a model organism and major crop. We explored the genetic variations within a collection of 224 maize inbred Dent lines and successively defined core genotype subsets to access multiple architectural, anatomical, and hydraulic parameters in the primary root (PR) and seminal roots (SR) of hydroponically grown seedlings. We found 9-, 3.5-, and 12.4-fold genotypic differences for root hydraulics (Lpr), PR size, and lateral root size, respectively, that shaped wide and independent variations of root structure and function. Within genotypes, PR and SR showed similarities in hydraulics and, to a lesser extent, in anatomy. They had comparable aquaporin activity profiles that, however, could not be explained by aquaporin expression levels. Genotypic variations in the size and number of late meta xylem vessels were positively correlated with Lpr. Inverse modeling further revealed dramatic genotypic differences in the xylem conductance profile. Thus, tremendous natural variation of maize root hydraulic architecture underlies a high diversity of water uptake strategies and paves the way to quantitative genetic dissection of its elementary traits.

Dual Inoculation with Rhizophagus irregularis and Bacillus megaterium Improves Maize Tolerance to Combined Drought and High Temperature Stress by Enhancing Root Hydraulics, Photosynthesis and Hormonal Responses

Climate change is leading to combined drought and high temperature stress in many areas, drastically reducing crop production, especially for high-water-consuming crops such as maize. This study aimed to determine how the co-inoculation of an arbuscular mycorrhizal (AM) fungus (Rhizophagus irregularis) and the PGPR Bacillus megaterium (Bm) alters the radial water movement and physiology in maize plants in order to cope with combined drought and high temperature stress. Thus, maize plants were kept uninoculated or inoculated with R. irregularis (AM), with B. megaterium (Bm) or with both microorganisms (AM + Bm) and subjected or not to combined drought and high temperature stress (D + T). We measured plant physiological responses, root hydraulic parameters, aquaporin gene expression and protein abundances and sap hormonal content. The results showed that dual AM + Bm inoculation was more effective against combined D + T stress than single inoculation. This was related to a synergistic enhancement of efficiency of the phytosystem II, stomatal conductance and photosynthetic activity. Moreover, dually inoculated plants maintained higher root hydraulic conductivity, which was related to regulation of the aquaporins ZmPIP1;3, ZmTIP1.1, ZmPIP2;2 and GintAQPF1 and levels of plant sap hormones. This study demonstrates the usefulness of combining beneficial soil microorganisms to improve crop productivity under the current climate-change scenario.

Role of Root Hydraulics in Plant Drought Tolerance

Role of Root Hydraulics in Plant Drought Tolerance

Xylem structure and hydraulic characteristics of deep roots, shallow roots and branches of walnut under seasonal drought

BackgroundDespite the importance of root hydraulics, there is little research on the in situ dynamic responses of embolism formation and embolism repair of roots distributed in different soil depths in response to different water regimes.ResultsThe vessel diameter, hydraulic conductivity, and vulnerability to cavitation were in the order of deep root > shallow root > branch. The midday PLC of shallow root was the highest in the dry season, while the midday PLC of deep root slightly higher than that of branch with no significant difference in the two seasons. The capacity of embolism repair of roots was significantly greater than that of branch both in dry season and wet season. The xylem pressure was in the order of deep roots > shallow root > branch, and it was negative in most of the time for the latter two in the dry season, but positive for both of the roots during the observation period in the wet season. The NSC and starch content in roots were significantly higher than those in branches, especially in the dry season. In contrast, roots had lower content of soluble sugar.ConclusionsThe relatively stable water condition in soil, especially in the deep layers, is favorable for the development of larger-diameter vessels in root xylem, however it cannot prevent the root from forming embolism. The mechanism of embolism repair may be different in different parts of plants. Deep roots mainly depend on root pressure to refill the embolized vessels, while branches mainly depend on starch hydrolysis to soluble sugars to do the work, with shallow roots shifted between the two mechanisms in different moisture regimes. There is theoretically an obvious trade-off between conducting efficiency and safety over deep roots, shallow roots and branches. But in natural conditions, roots do not necessarily suffer more severe embolism than branches, maybe due to their root pressure-driven embolism repair and relatively good water conditions.

Exposure to high nitrogen triggered a genotype-dependent modulation of cell and root hydraulics, which can involve aquaporin regulation.

Root nitrogen acquisition has been proposed to be regulated by mass flow, a process by which water flow brings nutrients to the root surface, depending on a concerted regulation of the root hydraulic properties and stomatal conductance. As aquaporins play an important role in regulating transcellular water flow, we aimed at evaluating the short-term effect of high nitrogen (HN) availability on the dynamics of hydraulic parameters at both the root and cell level and the regulation of aquaporins. The effect of short-term HN (8 mM NO3 - ) treatment was investigated on 12 diverse 15-day-old maize genotypes. Root exposure to HN triggered a rapid (<4 h) increase in the root hydraulic conductivity (Lpr ) in seven genotypes while no Lpr variation was recorded for the others, allowing the separation of the genotypes into two groups (HN-responsive and HN-nonresponsive). A remarkable correlation between Lpr and the cortex cell hydraulic conductivity (Lpc ) was observed. However, while differences in gas exchange parameters were also observed, the variations were genotype-specific and not always correlated with the root hydraulic parameters. We then investigated whether HN-induced Lpr variations were linked to the activity and regulation of plasma membrane PIP aquaporins. While some changes in PIP mRNA levels were detected, this was not correlated with the protein levels. On the other hand, the rapid variation in Lpr observed in the B73 genotype was correlated with the PIP protein abundance in the plasma membrane, highlighting PIP posttranslational mechanisms in the short-term regulation of root hydraulic parameters in response to HN treatment.

Root pressure-volume curve traits capture rootstock drought tolerance.

Living root tissues significantly constrain plant water uptake under drought, but we lack functional traits to feasibly screen diverse plants for variation in the drought responses of these tissues. Water stress causes roots to lose volume and turgor, which are crucial to root structure, hydraulics and growth. Thus, we hypothesized that root pressure-volume (p-v) curve traits, which quantify the effects of water potential on bulk root turgor and volume, would capture differences in rootstock drought tolerance. We used a greenhouse experiment to evaluate relationships between root p-v curve traits and gas exchange, whole-plant hydraulic conductance and biomass under drought for eight grapevine rootstocks that varied widely in drought performance in field trials (101-14, 110R, 420A, 5C, 140-Ru, 1103P, Ramsey and Riparia Gloire), grafted to the same scion variety (Vitis vinifera 'Chardonnay'). The traits varied significantly across rootstocks, and droughted vines significantly reduced root turgor loss point (πtlp), osmotic potential at full hydration (πo) and capacitance (C), indicating that roots became less susceptible to turgor loss and volumetric shrinkage. Rootstocks that retained a greater root volume (i.e. a lower C) also maintained more gas exchange under drought. The rootstocks that previous field trials have classified as drought tolerant exhibited significantly lower πtlp, πo and C values in well-watered conditions, but significantly higher πo and πtlp values under water stress, than the varieties classified as drought sensitive. These findings suggest that acclimation in root p-v curve traits improves gas exchange in persistently dry conditions, potentially through impacts on root hydraulics or root to shoot chemical signalling. However, retaining turgor and volume in previously unstressed roots, as these roots deplete wet soil to moderately negative water potentials, could be more important to drought performance in the deep, highly heterogenous rooting zones which grapevines develop under field conditions.

From hydraulic root architecture models to macroscopic representations of root hydraulics in soil water flow and land surface models

Abstract. Root water uptake is an important process in the terrestrial water cycle. How this process depends on soil water content, root distributions, and root properties is a soil–root hydraulic problem. We compare different approaches to implement root hydraulics in macroscopic soil water flow and land surface models. By upscaling a three-dimensional hydraulic root architecture model, we derived an exact macroscopic root hydraulic model. The macroscopic model uses the following three characteristics: the root system conductance, Krs, the standard uptake fraction, SUF, which represents the uptake from a soil profile with a uniform hydraulic head, and a compensatory matrix that describes the redistribution of water uptake in a non-uniform hydraulic head profile. The two characteristics, Krs and SUF, are sufficient to describe the total uptake as a function of the collar and soil water potential, and water uptake redistribution does not depend on the total uptake or collar water potential. We compared the exact model with two hydraulic root models that make a priori simplifications of the hydraulic root architecture, i.e., the parallel and big root model. The parallel root model uses only two characteristics, Krs and SUF, which can be calculated directly following a bottom-up approach from the 3D hydraulic root architecture. The big root model uses more parameters than the parallel root model, but these parameters cannot be obtained straightforwardly with a bottom-up approach. The big root model was parameterized using a top-down approach, i.e., directly from root segment hydraulic properties, assuming a priori a single big root architecture. This simplification of the hydraulic root architecture led to less accurate descriptions of root water uptake than by the parallel root model. To compute root water uptake in macroscopic soil water flow and land surface models, we recommend the use of the parallel root model with Krs and SUF computed in a bottom-up approach from a known 3D root hydraulic architecture.

Stomatal regulation and water potential variation in European beech: challenging the iso/anisohydry concept.

The iso/anisohydric continuum has been used to classify tree species' drought response strategies. The range over which stomata are regulating leaf water potential (ψl) before turgor loss occurs can be described with metrics such as the dependence of ψl on soil water potential (ψsoil) and the size of 'hydroscape area' (HA), but corresponding field data from adult trees are scarce. We examined the stomatal conductance (gs)-ψl relationship in its temporal (diurnal vs seasonal and interannual) and spatial (within-crown vs between-site) variation in European beech, using extensive ψl and gs measurements in the canopy of four beech stands across a precipitation gradient, and complemented the data set by published ψl and gs measurements in further Central European beech stands (including the extreme 2018 drought) in order to cover the full water potential operation space of the species. Both metrics characterize beech as a strictly anisohydric species with δψl/δψsoil>>1 and HA=4MPa2. However, stomates close sensitively in response to increasing vapor pressure deficit, disproving the widely assumed dependence of large ψl variation on looser stomatal control. Characterizing the water status regulation mechanisms of trees requires separating diurnal from day-to-day variation in ψl and gs. The large diurnal and seasonal ψl variation in beech leaves is partly caused by a low leaf tissue elasticity, suggesting that a whole-plant perspective with consideration of osmotic and elastic tissue properties and stem and root hydraulics is needed for fully understanding ψl regulation and the drought tolerance strategy of trees.

Combining cross-section images and modeling tools to create high-resolution root system hydraulic atlases in Zea mays.

Root hydraulic properties play a central role in the global water cycle, in agricultural systems productivity, and in ecosystem survival as they impact the canopy water supply. However, the existing experimental methods to quantify root hydraulic conductivities, such as the root pressure probing, are particularly challenging, and their applicability to thin roots and small root segments is limited. Therefore, there is a gap in methods enabling easy estimations of root hydraulic conductivities in diverse root types. Here, we present a new pipeline to quickly estimate root hydraulic conductivities across different root types, at high resolution along root axes. Shortly, free-hand root cross-sections were used to extract a selected number of key anatomical traits. We used these traits to parametrize the Generator of Root Anatomy in R (GRANAR) model to simulate root anatomical networks. Finally, we used these generated anatomical networks within the Model of Explicit Cross-section Hydraulic Anatomy (MECHA) to compute an estimation of the root axial and radial hydraulic conductivities (k x and k r , respectively). Using this combination of anatomical data and computational models, we were able to create a root hydraulic conductivity atlas at the root system level, for 14-day-old pot-grown Zea mays (maize) plants of the var. B73. The altas highlights the significant functional variations along and between different root types. For instance, predicted variations of radial conductivity along the root axis were strongly dependent on the maturation stage of hydrophobic barriers. The same was also true for the maturation rates of the metaxylem vessels. Differences in anatomical traits along and across root types generated substantial variations in radial and axial conductivities estimated with our novel approach. Our methodological pipeline combines anatomical data and computational models to turn root cross-section images into a detailed hydraulic atlas. It is an inexpensive, fast, and easily applicable investigation tool for root hydraulics that complements existing complex experimental methods. It opens the way to high-throughput studies on the functional importance of root types in plant hydraulics, especially if combined with novel phenotyping techniques such as laser ablation tomography.

Soil textures rather than root hairs dominate water uptake and soil-plant hydraulics under drought.

Although the role of root hairs (RHs) in nutrient uptake is well documented, their role in water uptake and drought tolerance remains controversial. Maize (Zea mays) wild-type and its hair-defective mutant (Mut; roothairless 3) were grown in two contrasting soil textures (sand and loam). We used a root pressure chamber to measure the relation between transpiration rate (E) and leaf xylem water potential (ψleaf_x) during soil drying. Our hypotheses were: (1) RHs extend root-soil contact and reduce the ψleaf_x decline at high E in dry soils; (2) the impact of RHs is more pronounced in sand; and (3) Muts partly compensate for lacking RHs by producing longer and/or thicker roots. The ψleaf_x(E) relation was linear in wet conditions and became nonlinear as the soils dried. This nonlinearity occurred more abruptly and at less negative matric potentials in sand (ca. -10 kPa) than in loam (ca. -100 kPa). At more negative soil matric potentials, soil hydraulic conductance became smaller than root hydraulic conductance in both soils. Both genotypes exhibited 1.7 times longer roots in loam, but 1.6 times thicker roots in sand. No differences were observed in the ψleaf_x(E) relation and active root length between the two genotypes. In maize, RHs had a minor contribution to soil-plant hydraulics in both soils and their putative role in water uptake was smaller than that reported for barley (Hordeum vulgare). These results suggest that the role of RHs cannot be easily generalized across species and soil textures affect the response of root hydraulics to soil drying.

Herb and conifer roots show similar high sensitivity to water deficit.

Root systems play a major role in supplying the canopy with water, enabling photosynthesis and growth. Yet, much of the dynamic response of root hydraulics and its influence on gas exchange during soil drying and recovery remains uncertain. We examined the decline and recovery of the whole root hydraulic conductance (Kr) and canopy diffusive conductance (gc) during exposure to moderate water stress in two species with contrasting root systems: Tanacetum cinerariifolium (herbaceous Asteraceae) and Callitris rhomboidea (woody conifer). Optical dendrometers were used to record stem water potential at high temporal resolution and enabled non-invasive measurements of Kr calculated from the rapid relaxation kinetics of water potential in hydrating roots. We observed parallel declines in Kr and gc to <20% of unstressed levels during the early stages of water stress in both species. The recovery of Kr after rewatering differed between species. T. cinerariifolium recovered quickly, with 60% of Kr recovered within 2 h, while C. rhomboidea was much slower to return to its original Kr. Recovery of gc followed a similar trend to Kr in both species, with C. rhomboidea slower to recover. Our findings suggest that the pronounced sensitivity of Kr to drought is a common feature among different plant species, but recovery may vary depending on root type and water stress severity. Kr dynamics are proposed to modulate gc response during and following drought.

Root hydraulics adjustment is governed by a dominant cell-to-cell pathway in Beta vulgaris seedlings exposed to salt stress

Soil salinity reduces root hydraulic conductivity (Lpr) of several plant species. However, how cellular signaling and root hydraulic properties are linked in plants that can cope with water restriction remains unclear. In this work, we exposed the halotolerant species red beet (Beta vulgaris) to increasing concentrations of NaCl to determine the components that might be critical to sustaining the capacity to adjust root hydraulics. Our strategy was to use both hydraulic and cellular approaches in hydroponically grown seedlings during the first osmotic phase of salt stress. Interestingly, Lpr presented a bimodal profile response apart from the magnitude of the imposed salt stress. As well as Lpr, the PIP2-aquaporin profile follows an unphosphorylated/phosphorylated pattern when increasing NaCl concentration while PIP1 aquaporins remain constant. Lpr also shows high sensitivity to cycloheximide. In low NaCl concentrations, Lpr was high and 70 % of its capacity could be attributed to the CHX-inhibited cell-to-cell pathway. More interestingly, roots can maintain a constant spontaneous exudated flow that is independent of the applied NaCl concentration. In conclusion, Beta vulgaris root hydraulic adjustment completely lies in a dominant cell-to-cell pathway that contributes to satisfying plant water demands.

Critical advances in understanding ecohydrological processes of terrestrial vegetation: From leaf to watershed scale

<p indent=0mm>The study of ecohydrological processes within vegetated terrestrial ecosystems is one of the new frontiers in ecology, hydrology and global change studies, and one of the most theoretical areas in ecohydrology. In recent years, a great deal of research has been conducted on the interactions between vegetative ecological processes and hydrologic processes, ranging from plant cells scales to whole continent scale. These studies have made great progress in changing the way we think and approach both ecology and hydrology. However, from the perspective of Ecohydrology, a discipline that integrates Ecology and Hydrology, there is an urgent need to systematically summarize the progress of multi-scale theories and methods and to present new ideas on the theoretical frontiers in this emergent field. This review synthesizes and evaluates recent findings and future challenges in the science and modeling of ecohydrological progresses in terrestrial vegetative systems including: (1) Plant water use and regulation; (2) coupled carbon-nitrogen-water cycle processes and modeling; (3) ecological effects of key processes in the hydrologic cycle; (4) influence of vegetation on the generation and alteration of runoff and overland flow; (5) feedback effects from changes in terrestrial vegetation ecological water cycle on localized and regional precipitation pattern. The main progress in these five aspects of ecohydrology at multi-scales and across-scales is systematically summarized, with special attention given to the development of process-based quantitative models. Based on our synthesis of progress in the ecohydrology of terrestrial vegetation systems, we identify the four primary avenues for future research. First, it is necessary to clarify the effectiveness of the “triangular trade-off relationship” among water transmission efficiency and safety, and those mechanisms underlying plant water use strategies given that plant water use strategies and water regulation mechanisms under changing environmental conditions are the foundation of ecohydrological theory. Accordingly, an exploration on the internal associations among this “triangular trade-off relationship” and root hydraulics redistribution, ecohydrological separation, and isohydric characteristics will be needed. Special attention is needed for a better understanding of the multi-scale mechanisms for plant water use and regulation at the individual-community scale. Second, in the light of marginal water consumption during vegetative carbon assimilation, we explore the multi-coupling relationships among marginal water consumption and environmental factors such as light intensity and water availability and variations in the relationship among various vegetation types and growth stages. The ecosystem carbon-water coupling parameters that have clear ecological significance across scales need to be determined and highlighted. It is also important to develop a cross-scale carbon and water flux modeling method based on the canopy optimal conductivity theory. Third, how and to what extent the water availability and its dynamic changes restrict ecosystem carbon and nitrogen process is one of the frontier bottlenecks that need to be resolved to advance global change ecology. It is necessary to systematically clarify the complex interactions and threshold conditions between the water availability, nitrogen supply, and carbon benefits of the ecosystem. Development of mechanism models for the full coupling of ecosystem carbon and nitrogen cycling and hydrologic cycle processes are also crucial. Fourth, the vegetation factors and mechanisms in the terrestrial water cycle remain key issues for continued attention in the future. The core issue is the identification and quantitative description of ecological factors in the formation and dynamic changes in runoff. On a larger spatial scale, it is also vital to accurately analyze the atmospheric feedback and contribution range of changes in precipitation from the vegetation cover changes in the relevant regions based on the understanding of the impact of land surface vegetation structure changes on the land-atmosphere coupling relationship.

Normalization criteria determine the interpretation of nitrogen effects on the root hydraulics of pine seedlings.

Plant hydraulics is key for plant survival and growth because it is linked to gas exchange and drought resistance. Although the environment influences plant hydraulics, there is no clear consensus on the effect of nitrogen (N) supply, which may be, in part, due to different hydraulic conductance normalization criteria and studied species. The objective of this study was to compare the variation of root hydraulic properties using several normalization criteria in four pine species in response to three contrasting N fertilization regimes. We studied four closely related, yet ecologically distinct species: Pinus nigra J.F. Arnold, Pinus pinaster Ait., Pinus pinea L. and Pinus halepensis Mill. Root hydraulic conductance (Kh) was measured with a high-pressure flow meter, and values were normalized by total leaf area (leaf specific conductance, Kl), xylem cross-section area (xylem specific conductance, Ks), total root area (root specific conductance, Kr) and the area of fine roots (fine root specific conductance, Kfr). Controlling for organ size differences allowed comparison of the hydraulic efficiency of roots to supply or absorb water among fertilization treatments and species. The effect of N on the root hydraulic efficiency depended on the normalization criteria. Increasing N availability reduced Kl and Ks, but increased Kh, Kr and especially Kfr. The positive effect of N on Kr and Kfr was positively related to seedling relative growth rate and was also consistent with published results at the interspecific level, whereby plant hydraulics is positively linked to photosynthesis and transpiration rate and fast growth. In contrast, normalization by leaf area and xylem cross-sectional area (Kl and Ks) reflected opposite responses to Kr and Kfr. This indicates that the normalization criteria determine the interpretation of the effect of N on plant hydraulics, which can limit species and treatment comparisons.

Decoupled drought responses of fine-root versus leaf acquisitive traits among six Prunus hybrids

Abstract Aims Predicting drought consequences on forests and fruit crop plantings requires improved understanding of drought responses of both leaf and fine-root resource acquisitive traits (specific leaf area—SLA, specific root surface area—SRA and specific root length—SRL). We hypothesize their responses are coordinated towards integrated plant resource conservation under severe drought. Methods We tested the hypothesis with a greenhouse-based drought experiment on saplings of six Prunus hybrids with a priori known contrasting drought sensitivity. Saplings were subjected to either control (100% field capacity) or severe drought stress treatment (33% evapotranspiration of hybrid-specific control plants). Sample collections were carried out at 30 and at 60 days after the start of treatments, for both control and stressed saplings. Important Findings No hybrid showed concurrent significant decrease of SLA and SRA (or SRL) under severe drought. The fine-root traits of the six hybrids showed two major drought-response scenarios, in particular: (i) increased root tissue density (RTD) and decreased average root diameter without significant change of SRL and (ii) increased RTD and decreased SRL without significant change of average root diameter. Drought responses of leaf gas exchange, SRA, SRL and RTD were closely correlated along a gradient towards resource conservation from control to drought-stressed plants in all hybrids, which was orthogonal to another gradient characterized by a hybrid-dependent decrease of SLA. These findings highlight (i) the multi-dimensionality of root-trait drought responses, (ii) the decoupling between leaf economics and leaf hydraulics and (iii) the covariation of leaf and root hydraulics in terms of trait drought responses. The study contributes to identifying the origin of the multi-dimensionality of root-trait drought response at intraspecific scale, and highlights differential drought–response combinations of leaf and fine-root traits among hybrids to survive under severe soil drought stress.

Declining root water transport drives stomatal closure in olive under moderate water stress.

Efficient water transport from soil to leaves sustains stomatal opening and steady-state photosynthesis. The aboveground portion of this pathway is well-described, yet the roots and their connection with the soil are still poorly understood due to technical limitations. Here we used a novel rehydration technique to investigate changes in the hydraulic pathway between roots and soil and within the plant body as individual olive plants were subjected to a range of water stresses. Whole root hydraulic resistance (including the radial pathway from xylem to the soil-root interface) constituted 81% of the whole-plant resistance in unstressed plants, increasing to >95% under a moderate level of water stress. The decline in this whole root hydraulic conductance occurred in parallel with stomatal closure and contributed significantly to the reduction in canopy conductance according to a hydraulic model. Our results demonstrate that losses in root hydraulic conductance, mainly due to a disconnection from the soil during moderate water stress in olive plants, are profound and sufficient to induce stomatal closure before cavitation occurs. Future studies will determine whether this core regulatory role of root hydraulics exists more generally among diverse plant species.