Catastrophic Hydraulic Failure and Tipping Points in Plants

Water inside plants forms a continuous chain from water in soils to the water evaporating from leaf surfaces. Failures in this chain result in reduced transpiration and photosynthesis and these failures are caused by soil drying and/or cavitation-induced xylem embolism. Xylem embolism and plant hydraulic failure share a number of analogies to “catastrophe theory” in dynamical systems. These catastrophes are often represented in the physiological and ecological literature as tipping points or alternative stable states when control variables exogenous (e.g. soil water potential) or endogenous (e.g. leaf water potential) to the plant are allowed to slowly vary. Here, plant hydraulics viewed from the perspective of catastrophes at multiple spatial scales is considered with attention to bubble expansion (i.e. cavitation), organ-scale vulnerability to embolism, and whole-plant biomass as a proxy for transpiration and hydraulic function. The hydraulic safety-efficiency tradeoff, hydraulic segmentation and maximum plant transpiration are examined using this framework. Underlying mechanisms for hydraulic failure at very fine scales such as pit membranes, intermediate scales such as xylem network properties and at larger scales such as soil-tree hydraulic pathways are discussed. Lacunarity areas in plant hydraulics are also flagged where progress is urgently needed.

Predicting mangrove forest dynamics across a soil salinity gradient using an individual-based vegetation model linked with plant hydraulics

In mangrove forests, soil salinity is one of the most significant environmental factors determining mangrove forest distribution and productivity as it limits plant water uptake and carbon gain. However, salinity control on mangrove productivity through plant hydraulics has not been investigated by existing mangrove models. Thus, we present a new individual-based model linked with plant hydraulics to incorporate physiological characterization of mangrove growth under salt stress. Plant hydraulics was associated with mangroves nutrient uptake and biomass allocation apart from water flux and carbon gain. The developed model was performed for two-coexisting species of Rhizophora stylosa and Bruguiera gymnorrhiza in a subtropical mangrove forest in Japan. The model predicted that the productivity of both species was affected by soil salinity through downregulation of stomatal conductance, while B. gymnorrhiza trees grow faster and suppress the growth of R. stylosa trees by shading that resulted in a B. gymnorrhiza-dominated forest under low soil salinity conditions (< 28 ‰). Alternatively, the increase in soil salinity significantly reduced the productivity of B. gymnorrhiza compared to R. stylosa, leading to an increase in biomass of R. stylosa despite the enhanced salt stress (> 30 ‰). These predicted patterns in forest structures across soil salinity gradient remarkably agreed with field data, highlighting the control of salinity on productivity and tree competition as factors that shape the mangrove forest structures. The model reproducibility of forest structures was also supported by the predicted self-thinning processes, which likewise agreed with field data. In addition, the mangroves morphological adjustment to increasing soil salinity – by decreasing transpiration and increasing hydraulic conductance – was reasonably predicted. Aside from the soil salinity, seasonal dynamics in atmospheric variables (solar radiation and temperature) was highlighted as factors influencing mangrove productivity in a subtropical region. The physiological principle-based improved model has the potential to be extended to other mangrove forests in various environmental settings, thus contributing to a better understanding of mangrove dynamics under future global climate change.

When Water Availability Is Low, Two Mediterranean Salvia Species Rely on Root Hydraulics

Increase in severity and frequency of drought events is altering plant community composition, exposing biomes to a higher risk of biodiversity losses. This is exacerbated in the most fragile areas as Mediterranean biome. Thus, identifying plant traits for forecasting species with a high risk of drought-driven mortality is particularly urgent. In the present study, we investigated the drought resistance strategy of two Mediterranean native species: Salvia ceratophylloides Ard. (Sc) and Salvia officinalis L. (So) by considering the impact of drought-driven water content decline on plant hydraulics. Well-watered samples of Sc displayed higher leaf and stemsaturated water content and lower shoot biomass than So samples, but similar root biomass. In response to drought, Sc showed a conservative water use strategy, as the prompt stomatal closure and leaves shedding suggested. A drought-tolerant mechanism was confirmed in So samples. Nevertheless, Sc and So showed similar drought-driven plant hydraulic conductance (Kplant) recover ability. Root hydraulic traits played a key role to reach this goal. Relative water content as well as loss of cell rehydration capability and membrane damages, especially of stem and root, were good proxies of drought-driven Kplant decline.

New Representation of Plant Hydraulics Improves the Estimates of Transpiration in Land Surface Model

Transpiration represents more than 30% of the global land–atmosphere water exchange but is highly uncertain. Plant hydraulics was ignored in traditional land surface modeling, but recently plant hydraulics has been found to play an essential role in transpiration simulation. A new physical-based representation of plant hydraulic schemes (PHS) was recently developed and implemented in the Common Land Model (CoLM). However, it is unclear to what extent PHS can reduce these uncertainties. Here, we evaluated the PHS against measurements obtained at 81 FLUXNET sites. The transpiration of each site was estimated using an empirical evapotranspiration partitioning approach. The metric scores defined by the International Land Model Benchmarking Project (ILAMB) were used to evaluate the model performance and compare it with that of the CoLM default scheme (soil moisture stress (SMS)). The bias score of transpiration in PHS was higher than SMS for most sites, and more significant improvements were found in semi-arid and arid sites where transpiration was limited by soil moisture. The hydraulic redistribution in PHS optimized the soil water supply and thus improved the transpiration estimates. In humid sites, no significant improvement in seasonal or interannual variability of transpiration was simulated by PHS, which can be explained by the insensitivity of transpiration demand coupled to the photosynthesis response to precipitation. In arid and semi-arid sites, seasonal or interannual variability of transpiration was better captured by PHS than SMS, which was interpreted by the improved drought sensitivity for transpiration. Arid land is widespread and is expected to expand due to climate change, thus there is an urgent need to couple PHS in land surface models.

Global ecosystem-scale plant hydraulic traits retrieved using model–data fusion

Droughts are expected to become more frequent and severe under climate change, increasing the need for accurate predictions of plant drought response. This response varies substantially, depending on plant properties that regulate water transport and storage within plants, i.e., plant hydraulic traits. It is, therefore, crucial to map plant hydraulic traits at a large scale to better assess drought impacts. Improved understanding of global variations in plant hydraulic traits is also needed for parameterizing the latest generation of land surface models, many of which explicitly simulate plant hydraulic processes for the first time. Here, we use a model–data fusion approach to evaluate the spatial pattern of plant hydraulic traits across the globe. This approach integrates a plant hydraulic model with data sets derived from microwave remote sensing that inform ecosystem-scale plant water regulation. In particular, we use both surface soil moisture and vegetation optical depth (VOD) derived from the X-band Japan Aerospace Exploration Agency (JAXA) Advanced Microwave Scanning Radiometer for Earth Observing System (EOS; collectively AMSR-E). VOD is proportional to vegetation water content and, therefore, closely related to leaf water potential. In addition, evapotranspiration (ET) from the Atmosphere–Land Exchange Inverse (ALEXI) model is also used as a constraint to derive plant hydraulic traits. The derived traits are compared to independent data sources based on ground measurements. Using the K-means clustering method, we build six hydraulic functional types (HFTs) with distinct trait combinations – mathematically tractable alternatives to the common approach of assigning plant hydraulic values based on plant functional types. Using traits averaged by HFTs rather than by plant functional types (PFTs) improves VOD and ET estimation accuracies in the majority of areas across the globe. The use of HFTs and/or plant hydraulic traits derived from model–data fusion in this study will contribute to improved parameterization of plant hydraulics in large-scale models and the prediction of ecosystem drought response.

Evidence for a multicellular symplasmic water pumping mechanism across vascular plant roots

With global warming, climate zones are projected to shift poleward, and the frequency and intensity of droughts to increase, driving threats to crop production and ecosystems. Plant hydraulic traits play major roles in coping with such droughts, and process-based plant hydraulics (water flowing along decreasing pressure or total water potential gradients) has newly been implemented in land surface models. An enigma reported for the past 35 years is the observation of water flowing along increasing water potential gradients across roots. By combining the most advanced modelling tool from the emerging field of plant micro-hydrology with pioneering cell solute mapping data, we found that the current paradigm of water flow across roots of all vascular plants is incomplete: it lacks the impact of solute concentration (and thus negative osmotic potential) gradients across living cells. This gradient acts as a water pump as it reduces water tension without loading solutes in plant vasculature (xylem). Importantly, water tension adjustments in roots may have large impacts in leaves due to the tension-cavitation feedback along stems. Here, we mathematically demonstrate the water pumping mechanism by solving water flow equations analytically on a triple-cell system. Then we show that the simplistic upscaled equations hold in 2- and 3-D maize, grapevine and Arabidopsis complex hydraulic anatomies, and that water may flow uphill of water potential gradients toward xylem as observed experimentally. Besides its contribution to the fundamental understanding of plant water relations, this study lays new foundations for future multidisciplinary research encompassing plant physiology and ecohydrology, and has the ambition to mathematically capture a keystone process for the accurate forecasting of plant water status in crop models and LSMs.