• Trait variability in space and time allows plants to adjust to changing environmental conditions. However, we know little about how this variability is distributed and coordinated at different ...organizational levels.
• For six dominant tree species in northeastern Spain (three Fagaceae and three Pinaceae) we quantified the inter- and intraspecific variability of a set of traits along a water availability gradient. We measured leaf mass per area (LMA), leaf nitrogen (N) concentration, carbon isotope composition in leaves (δ13C), stem wood density, the Huber value (Hv, the ratio of cross-sectional sapwood area to leaf area), sapwood-specific and leaf-specific stem hydraulic conductivity, vulnerability to xylem embolism (P
50) and the turgor loss point (P
tlp).
• Differences between families explained the largest amount of variability for most traits, although intraspecific variability was also relevant. Species occupying wetter sites showed higher N, P
50 and P
tlp, and lower LMA, δ13C and Hv. However, when trait relationships with water availability were assessed within species they held only for Hv and P
tlp.
• Overall, our results indicate that intraspecific adjustments along the water availability gradient relied primarily on changes in resource allocation between sapwood and leaf area and in leaf water relations.
Not long ago, textbooks on plant physiology divulged the view that phloem and xylem are separate transport systems with exclusive functions. Phloem was flowing downwards providing roots with ...carbohydrates. Xylem transported water upwards from roots to leaves. This simplified view has changed forever. Today we have a much-refined understanding of the complex transport mechanisms, regulatory functions and surprisingly ingenuous solutions trees have evolved to distribute carbohydrates and water internally to fuel growth and help mediate biotic and abiotic stresses. This review focuses on functional links between tissues of the inner bark region (i.e., more than just phloem) and the xylem, facilitated by radially aligned and interconnected parenchyma cells, called rays. Rays are usually found along the entire vertical axis of tree stems, mediating a number of transport processes. We use a top-down approach to unveil the role of rays in these processes. Due to the central role of rays in facilitating the coupling of inner bark and xylem we dedicate the first section to ray anatomy, pathways and control mechanisms involved in radial transport. In the second section, basic concepts and models for radial movement through rays are introduced and their impacts on water and carbon fluxes at the whole-tree level are discussed. This section is followed by a closer look at the capacitive function of composite tissues in stems where gradual changes in water potential generate a diurnal 'pulse'. We explain how this pulse can be measured and interpreted, and where the limitations of such analyses are. Towards the end of this review, we include a brief description of the role of radial transport during limited availability of water. By elucidating the strong hydraulic link between inner bark and xylem, the traditional view of two separate transport systems dissolves and the idea of one interconnected, yet highly segregated transport network for carbohydrates and water arises.
Plant function requires effective mechanisms to regulate water transport at a variety of scales. Here, we develop a new theoretical framework describing plant responses to drying soil, based on the ...relationship between midday and predawn leaf water potentials. The intercept of the relationship (Λ) characterizes the maximum transpiration rate per unit of hydraulic transport capacity, whereas the slope (σ) measures the relative sensitivity of the transpiration rate and plant hydraulic conductance to declining water availability. This framework was applied to a newly compiled global database of leaf water potentials to estimate the values of Λ and σ for 102 plant species. Our results show that our characterization of drought responses is largely consistent within species, and that the parameters Λ and σ show meaningful associations with climate across species. Parameter σ was ≤1 in most species, indicating a tight coordination between the gas and liquid phases of water transport, in which canopy transpiration tended to decline faster than hydraulic conductance during drought, thus reducing the pressure drop through the plant. The quantitative framework presented here offers a new way of characterizing water transport regulation in plants that can be used to assess their vulnerability to drought under current and future climatic conditions.
Increased tree mortality during and after drought has become a research focus in recent years. This focus has been driven by: the realisation that drought-related tree mortality is more widespread ...than previously thought; the predicted increase in the frequency of climate extremes this century; and the recognition that current vegetation models do not predict drought-related tree mortality and forest dieback well despite the large potential effects of these processes on species composition and biogeochemical cycling. To date, the emphasis has been on understanding the causal mechanisms of drought-related tree mortality, and on mechanistic models of plant function and vegetation dynamics, but a consensus on those mechanisms has yet to emerge. In order to generate new hypotheses and to help advance the modelling of vegetation dynamics in the face of incomplete mechanistic understanding, we suggest that general patterns should be distilled from the diverse and as-yet inconclusive results of existing studies, and more use should be made of optimisation and probabilistic modelling approaches that have been successfully applied elsewhere in plant ecology. The outcome should inform new empirical studies of tree mortality, help improve its prediction and reduce model complexity.
Stomatal regulation presumably evolved to optimize CO2 for H2O exchange in response to changing conditions. If the optimization criterion can be readily measured or calculated, then stomatal ...responses can be efficiently modelled without recourse to empirical models or underlying mechanism. Previous efforts have been challenged by the lack of a transparent index for the cost of losing water. Yet it is accepted that stomata control water loss to avoid excessive loss of hydraulic conductance from cavitation and soil drying. Proximity to hydraulic failure and desiccation can represent the cost of water loss. If at any given instant, the stomatal aperture adjusts to maximize the instantaneous difference between photosynthetic gain and hydraulic cost, then a model can predict the trajectory of stomatal responses to changes in environment across time. Results of this optimization model are consistent with the widely used Ball–Berry–Leuning empirical model (r2 > 0.99) across a wide range of vapour pressure deficits and ambient CO2 concentrations for wet soil. The advantage of the optimization approach is the absence of empirical coefficients, applicability to dry as well as wet soil and prediction of plant hydraulic status along with gas exchange.
Current land surface models struggle to represent the complex and species‐specific manner by which stomata respond to environmental cues, especially soil drought. This paper offers a solution to this problem by assuming that the goal of stomatal regulation is to maximize photosynthetic gain minus hydraulic cost. A trait‐ and process‐based ‘profit‐maximizing’ algorithm predicts realistic stomatal behaviour in response to the gamut of environmental stimuli. This new approach to stomatal optimization theory may prove useful in large‐scale modelling of responses to climate change.
Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a ...conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.
Terrestrial carbon stock mapping is important for the successful implementation of climate change mitigation policies. Its accuracy depends on the availability of reliable allometric models to infer ...oven‐dry aboveground biomass of trees from census data. The degree of uncertainty associated with previously published pantropical aboveground biomass allometries is large. We analyzed a global database of directly harvested trees at 58 sites, spanning a wide range of climatic conditions and vegetation types (4004 trees ≥ 5 cm trunk diameter). When trunk diameter, total tree height, and wood specific gravity were included in the aboveground biomass model as covariates, a single model was found to hold across tropical vegetation types, with no detectable effect of region or environmental factors. The mean percent bias and variance of this model was only slightly higher than that of locally fitted models. Wood specific gravity was an important predictor of aboveground biomass, especially when including a much broader range of vegetation types than previous studies. The generic tree diameter–height relationship depended linearly on a bioclimatic stress variable E, which compounds indices of temperature variability, precipitation variability, and drought intensity. For cases in which total tree height is unavailable for aboveground biomass estimation, a pantropical model incorporating wood density, trunk diameter, and the variable E outperformed previously published models without height. However, to minimize bias, the development of locally derived diameter–height relationships is advised whenever possible. Both new allometric models should contribute to improve the accuracy of biomass assessment protocols in tropical vegetation types, and to advancing our understanding of architectural and evolutionary constraints on woody plant development.
Accurate modelling of drought-induced mortality is challenging. A steady-state model is presented integrating xylem and phloem transport, leaf-level gas exchange and plant carbohydrate consumption ...during drought development.
A Bayesian analysis of parameter uncertainty based on expert knowledge and a literature review is carried out. The model is tested by combining six data compilations covering 170 species using information on sensitivities of xylem conductivity, stomatal conductance and leaf turgor to water potential.
The possible modes of plant failure at steady state are identified (i.e. carbon (C) starvation, hydraulic failure and phloem transport failure). Carbon starvation occurs primarily in the parameter space of isohydric stomatal control, whereas hydraulic failure is prevalent in the space of xylem susceptibility to embolism. Relative to C starvation, phloem transport failure occurs under conditions of low sensitivity of photosynthesis and high sensitivity of growth to plant water status.
These three failure modes are possible extremes along two axes of physiological vulnerabilities, one characterized by the balance of water supply and demand and the other by the balance between carbohydrate sources and sinks. Because the expression of physiological vulnerabilities is coordinated, we argue that different failure modes should occur with roughly equal likelihood, consistent with predictions using optimality theory.
Hydraulic properties control plant responses to climate and are likely to be under strong selective pressure, but their macro‐evolutionary history remains poorly characterised. To fill this gap, we ...compiled a global dataset of hydraulic traits describing xylem conductivity (Ks), xylem resistance to embolism (P50), sapwood allocation relative to leaf area (Hv) and drought exposure (ψmin), and matched it with global seed plant phylogenies. Individually, these traits present medium to high levels of phylogenetic signal, partly related to environmental selective pressures shaping lineage evolution. Most of these traits evolved independently of each other, being co‐selected by the same environmental pressures. However, the evolutionary correlations between P50 and ψmin and between Ks and Hv show signs of deeper evolutionary integration because of functional, developmental or genetic constraints, conforming to evolutionary modules. We do not detect evolutionary integration between conductivity and resistance to embolism, rejecting a hardwired trade‐off for this pair of traits.
We compiled a global dataset of hydraulic traits describing xylem conductivity (Ks), xylem resistance to embolism (P50), sapwood allocation relative to leaf area (Hv) and drought exposure (ψmin), and matched it with global seed plant phylogenies. Individually, these traits present medium to high levels of phylogenetic signal, partly related to environmental selective pressures shaping lineage evolution. Most of these traits evolved independently of each other, being co‐selected by the same environmental pressures. However, the evolutionary correlations between P50 and ψmin and between Ks and Hv show signs of deeper evolutionary integration because of functional, developmental or genetic constraints, conforming to evolutionary modules. We do not detect evolutionary integration between conductivity and resistance to embolism, rejecting a hardwired trade‐off for this pair of traits.
Currently, phloem transport in plants under field conditions is not well understood. This is largely the result of the lack of techniques suitable for the measurement of the physiological properties ...of phloem.
We present a model that interprets the changes in xylem diameter and live bark thickness and separates the components responsible for such changes. We test the predictions from this model on data from three mature Scots pine trees in Finland. The model separates the live bark thickness variations caused by bark water capacitance from a residual signal interpreted to indicate the turgor changes in the bark.
The predictions from the model are consistent with processes related to phloem transport. At the diurnal scale, this signal is related to patterns of photosynthetic activity and phloem loading. At the seasonal scale, bark turgor showed rapid changes during two droughts and after two rainfall events, consistent with physiological predictions. Daily cumulative totals of this turgor term were related to daily cumulative totals of canopy photosynthesis. Finally, the model parameter representing radial hydraulic conductance between phloem and xylem showed a temperature dependence consistent with the temperature-driven changes in water viscosity.
We propose that this model has potential for the continuous field monitoring of tree phloem function.
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