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• Root access to bedrock water storage or groundwater is an important trait allowing plant survival in seasonally dry environments. However, the degree of coordination between water uptake depth, leaf-level water-use efficiency (WUEi) and water potential in drought-prone plant communities is not well understood. • We conducted a 135-d rainfall exclusion experiment in a subtropical karst ecosystem with thin skeletal soils to evaluate the responses of 11 co-occurring woody species of contrasting life forms and leaf habits to a severe drought during the wet growing season. • Marked differences in xylem water isotopic composition during drought revealed distinct ecohydrological niche separation among species. The contrasting behaviour of leaf water potential in coexisting species during drought was largely explained by differences in root access to deeper, temporally stable water sources. Smaller-diameter species with shallower water uptake, more negative water potentials and lower WUEi showed extensive drought-induced canopy defoliation and/or mortality. By contrast, larger-diameter species with deeper water uptake, higher leaf-level WUEi and more isohydric behaviour survived drought with only moderate canopy defoliation. • Severe water limitation imposes strong environmental filtering and/or selective pressures resulting in tight coordination between tree diameter, water uptake depth, iso/anisohydric behaviour, WUEi and drought vulnerability in karst plant communities

• A growing number of field studies report isotopic offsets between stem water and its potential sources that prevent the unambiguous identification of plant water origin using water isotopes. We explored the causes of this isotopic offset by conducting a controlled experiment on the temperate tree species Fagus sylvatica. • We measured δ²H and δ18O of soil and stem water from potted saplings growing on three soil substrates and subjected to two watering regimes. • Regardless of substrate, soil and stem water δ²H were similar only near permanent wilting point. Under moister conditions, stem water δ²H was 11 ± 3‰ more negative than soil water δ²H, coherent with field studies. Under drier conditions, stem water δ²H became progressively more enriched than soil water δ²H. Although stem water δ18O broadly reflected that of soil water, soil–stem δ²H and δ18O differences were correlated (r = 0.76) and increased with transpiration rates indicated by proxies. • Soil–stem isotopic offsets are more likely to be caused by water isotope heterogeneities within the soil pore and stem tissues, which would be masked under drier conditions as a result of evaporative enrichment, than by fractionation under root water uptake. Our results challenge our current understanding of isotopic signals in the soil–plant continuum.

We tested for isotope exchange between bound (immobile) and mobile soil water, and whether there is isotope fractionation during plant water uptake. These are critical assumptions to the formulation of the ‘two water worlds’ hypothesis based on isotope profiles of soil water. In two different soil types, soil-bound water in two sets of 19-l pots, each with a 2-yr-old avocado plant (Persea americana), were identically labeled with tap water. After which, one set received isotopically enriched water whereas the other set received tap water as the mobile phase water. After a dry down period, we analyzed plant stem water as a proxy for soil-bound water as well as total soil water by cryogenic distillation. Seventy-five to 95% of the bound water isotopically exchanged with the mobile water phase. In addition, plants discriminated against 18O and 2H during water uptake, and this discrimination is a function of the soil water loss and soil type. The present experiment shows that the assumptions for the ‘two water worlds’ hypothesis are not supported. We propose a novel explanation for the discrepancy between isotope ratios of the soil water profile and other water compartments in the hydrological cycle.

Version 5 of the Community Land Model (CLM5) introduces the plant hydraulic stress (PHS) configuration of vegetation water use, which is described and compared with the corresponding parameterization from CLM4.5. PHS updates vegetation water stress and root water uptake to better reflect plant hydraulic theory, advancing the physical basis of the model. The new configuration introduces prognostic vegetation water potential, modeled at the root, stem, and leaf levels. Leaf water potential replaces soil potential as the basis for stomatal conductance water stress, and root water potential is used to implement hydraulic root water uptake, replacing a transpiration partitioning function. Point simulations of a tropical forest site (Caxiuanã, Brazil) under ambient conditions and partial precipitation exclusion highlight the differences between PHS and the previous CLM implementation. Model description and simulation results are contextualized with a list of benefits and limitations of the new model formulation, including hypotheses that were not testable in previous versions of the model. Key results include reductions in transpiration and soil moisture biases relative to a control model under both ambient and exclusion conditions, correcting excessive dry season soil moisture stress in the control model. PHS implements hydraulic gradient root water uptake, which allows hydraulic redistribution and compensatory root water uptake and results in PHS utilizing a larger portion of the soil column to buffer shortfalls in precipitation. The new model structure, which bases water stress on leaf water potential, could have significant implications for vegetation‐climate feedbacks, including increased sensitivity of photosynthesis to atmospheric vapor pressure deficit. Key Points An updated soil‐plant‐atmosphere continuum model based on hydraulic theory is implemented in the Community Land Model (version 5) Prognostic leaf water potential replaces soil matric potential as the basis for stomatal conductance water stress Prognostic root water potential is used to implement hydraulic root water uptake, replacing a “soil wilting point” approach

• The significance of shoot surface water uptake (SSWU) has been debated, and it would depend on the range of conditions under which it occurs. We hypothesized that the decline of leaf hydraulic conductance (K leaf) in response to dehydration may be recovered through SSWU, and that the hydraulic conductance to SSWU (K surf) declines with dehydration. • We quantified effects of leaf dehydration on K surf and effects of SSWU on recovery of K leaf in dehydrated leaves of Avicennia marina. • SSWU led to overnight recovery of K leaf, with recovery retracing the same path as loss of K leaf in response to dehydration. SSWU declined with dehydration. By contrast, K surf declined with rehydration time but not with dehydration. • Our results showed a role of SSWU in the recovery of leaf hydraulic conductance and revealed that SSWU is sensitive to leaf hydration status. The prevalence of SSWU in vegetation suggests an important role for atmospheric water sources in maintenance of leaf hydraulic function, with implications for plant responses to changing environments.

We showed that crown roots have a different capacity to transport water compared with seminal roots. Acknowledging such differences between root types is crucial to understand optimal root traits. Abstract The ability of plants to take up water from the soil depends on both the root architecture and the distribution and evolution of the hydraulic conductivities among root types and along the root length. The mature maize (Zea mays L.) root system is composed of primary, seminal, and crown roots together with their respective laterals. Our understanding of root water uptake of maize is largely based on measurements of primary and seminal roots. Crown roots might have a different ability to extract water from the soil, but their hydraulic function remains unknown. The aim of this study was to measure the location of water uptake in mature maize and investigate differences between seminal, crown, and lateral roots. Neutron radiography and injections of deuterated water were used to visualize the root architecture and water transport in 5-week-old maize root systems. Water was mainly taken up by crown roots. Seminal roots and their laterals, which were the main location of water uptake in younger plants, made a minor contribution to water uptake. In contrast to younger seminal roots, crown roots were also able to take up water from their most distal segments. The greater uptake of crown roots compared with seminal roots is explained by their higher axial conductivity in the proximal parts and by the fact that they are connected to the shoot above the seminal roots, which favors the propagation of xylem tension along the crown roots. The deeper water uptake of crown roots is explained by their shorter and fewer laterals, which decreases the dissipation of water potential along the roots.

The ‘two-water-worlds’ hypothesis is based on stable isotope differences in stream, soil and xylem waters in dual isotope space. It postulates no connectivity between bound and mobile soil waters, and preferential plant water uptake of bound soil water sources. We tested the pool-weighted impact of isotopically distinct water pools for hydrological cycling, the influence of species-specific water use and the degree of ecohydrological separation. We combined stable isotope analysis (δ18O and δ²H) of ecosystem water pools of precipitation, groundwater, soil and xylem water of two distinct species (Quercus suber, Cistus ladanifer) with observations of soil water contents and sap flow. Shallow soil water was evaporatively enriched during dry-down periods, but enrichment faded strongly with depth and upon precipitation events. Despite clearly distinct water sources and water-use strategies, both species displayed a highly opportunistic pattern of root water uptake. Here we offer an alternative explanation, showing that the isotopic differences between soil and plant water vs groundwater can be fully explained by spatio-temporal dynamics. Pool weighting the contribution of evaporatively enriched soil water reveals only minor annual impacts of these sources to ecosystem water cycling (c. 11% of bulk soil water and c. 14% of transpired water).

• The mangrove Avicennia marina adjusts internal salt concentrations by foliar salt secretion. Deliquescence of accumulated salt causes leaf wetting that may provide a water source for salt-secreting plants in arid coastal wetlands where high nocturnal humidity can usually support deliquescence whereas rainfall events are rare. We tested the hypotheses that salt deliquescence on leaf surfaces can drive top-down rehydration, and that such absorption of moisture from unsaturated atmospheres makes a functional contribution to dry season shoot water balances. • Sap flow and water relations were monitored to assess the uptake of atmospheric water by branches during shoot wetting events under natural and manipulated microclimatic conditions. • Reverse sap flow rates increased with increasing relative humidity from 70% to 89%, consistent with function of salt deliquescence in harvesting moisture from unsaturated atmospheres. Top-down rehydration elevated branch water potentials above those possible from root water uptake, subsidising transpiration rates and reducing branch vulnerability to hydraulic failure in the subsequent photoperiod. • Absorption of atmospheric moisture harvested through deliquescence of salt on leaf surfaces enhances water balances of Avicennia marina growing in hypersaline wetlands under arid climatic conditions. Top-down rehydration from these frequent, low intensity wetting events contributes to prevention of carbon starvation and hydraulic failure during drought.

Dew is an important water resource for plants in most deserts. The mechanism that allows desert plants to use dew water was studied using an isotopic water tracer approach. Most plants use water directly from the soil; the roots transfer the water to the rest of the plant, where it is required for all metabolic functions. However, many plants can also take up water into their leaves and stems. Examining the dew water uptake pathways in desert plants can lend insight on another all water-use pathways examination. We determined where and how dew water enters plants in the water limited Negev desert. Highly depleted isotopic water was sprayed on three different dominant plant species of the Negev desert—Artemesia sieberi, Salsola inermis and Haloxylon scoparium—and its entry into the plant was followed. Water was sprayed onto the soil only, or on the leaves/stems only (with soil covered to prevent water entry via root uptake). Thereafter, the isotopic composition of water in the roots and stems were measured at various time points. The results show that each plant species used the dew water to a different extent, and we obtained evidence of foliar uptake capacity of dew water that varied depending on the microenvironmental conditions. A. sieberi took up the greatest amount of dew water through both stems and roots, S. inermis took up dew water mainly from the roots, and H. scoparium showed the least dew capture overall.

A common effect of several abiotic stresses is to cause tissue dehydration. Such dehydration is caused by the imbalance between root water uptake and leaf transpiration. Under some specific stress conditions, regulation of root water uptake is more crucial to overcome stress injury than regulation of leaf transpiration. This review first describes present knowledge about how water is taken up by roots and then discusses how specific stress situations such as drought, salinity, low temperature, and flooding modify root water uptake. The rate of root water uptake of a given plant is the result of its root hydraulic characteristics, which are ultimately regulated by aquaporin activity and, to some extent, by suberin deposition. Present knowledge about the effects of different stresses on these features is also summarized. Finally, current findings regarding how molecular signals such as the plant hormones abscisic acid, ethylene, and salicylic acid, and how reactive oxygen species may modulate the final response of root water uptake under stress conditions are discussed.