The hydraulic properties of xylem and phloem differ but the magnitude and functional consequences of the differences are not well understood. Phloem and xylem functional areas, hydraulic conduit ...diameters and conduit frequency along the stems of Picea abies trees were measured and expressed as allometric functions of stem diameter and distance from stem apex. Conductivities of phloem and xylem were estimated from these scaling relations. Compared with xylem, phloem conduits were smaller and occupied a slightly larger fraction of conducting tissue area. Ten times more xylem than phloem was annually produced along the stem. Scaling of the conduit diameters and cross‐sectional areas with stem diameter were very similar in phloem and xylem. Phloem and xylem conduits scaled also similarly with distance from stem apex; widening downwards from the tree top, and reaching a plateau near the base of the living crown. Phloem conductivity was estimated to scale similarly to the conductivity of the outermost xylem ring, with the ratio of phloem to xylem conductivity being c. 2%. However, xylem conductivity was estimated to increase more than phloem conductivity with increasing tree dimensions as a result of accumulation of xylem sapwood. Phloem partly compensated for its smaller conducting area and narrower conduits by having a slightly higher conduit frequency.
Summary
Although transpiration‐driven transport of xylem sap is well known to operate under absolute negative pressure, many terrestrial, vascular plants show positive xylem pressure above ...atmospheric pressure on a seasonal or daily basis, or during early developmental stages. The actual location and mechanisms behind positive xylem pressure remain largely unknown, both in plants that show seasonal xylem pressure before leaf flushing, and those that show a diurnal periodicity of bleeding and guttation. Available evidence shows that positive xylem pressure can be driven based on purely physical forces, osmotic exudation into xylem conduits, or hydraulic pressure in parenchyma cells associated with conduits. The latter two mechanisms may not be mutually exclusive and can be understood based on a similar modelling scenario. Given the renewed interest in positive xylem pressure, this review aims to provide a constructive way forward by discussing similarities and differences of mechanistic models, evaluating available evidence for hydraulic functions, such as rehydration of tissues, refilling of water stores, and embolism repair under positive pressure, and providing recommendations for future research, including methods that avoid or minimise cutting artefacts.
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.
Optimization models of stomatal conductance (g
s) attempt to explain observed stomatal behaviour in terms of cost–benefit tradeoffs. While the benefit of stomatal opening through increased CO2 uptake ...is clear, currently the nature of the associated cost(s) remains unclear. We explored the hypothesis that g
s maximizes leaf photosynthesis, where the cost of stomatal opening arises from nonstomatal reductions in photosynthesis induced by leaf water stress.
We analytically solved two cases, CAP and MES, in which reduced leaf water potential leads to reductions in carboxylation capacity (CAP) and mesophyll conductance (g
m) (MES).
Both CAP and MES predict the same one-parameter relationship between the intercellular: atmospheric CO2 concentration ratio (c
i/c
a) and vapour pressure deficit (VPD, D), viz.
c
i
/
c
a
≈
ξ
/
(
ξ
+
D
)
, as that obtained from previous optimization models, with the novel feature that the parameter ξ is determined unambiguously as a function of a small number of photosynthetic and hydraulic variables. These include soil-to-leaf hydraulic conductance, implying a stomatal closure response to drought. MES also predicts that g
s/g
m is closely related to c
i/c
a and is similarly conservative.
These results are consistent with observations, give rise to new testable predictions, and offer new insights into the covariation of stomatal, mesophyll and hydraulic conductances.
Ecosystem respiration is known to vary following changes in canopy photosynthesis. However, the timing of this coupling is not well understood. Here, we summarize the literature on soil and ecosystem ...respiration where the speed of transfer of photosynthetic sugars from the plant canopy via the phloem to the roots was determined. Estimates of the transfer speed can be grouped according to whether the study employed isotopic or canopy/soil flux-based techniques. These two groups should provide different estimates of transfer times because transport of sucrose molecules, and pressure-concentration waves, in phloem differ. A steady-state and a dynamic photosynthesis/phloem-transport/soil gas diffusion model were employed to interpret our results. Starch storage and partly soil gas diffusion affected transfer times, but phloem path-length strongly controlled molecule transfer times. Successful modelling required substantially different phloem properties (higher specific conductivity and turgor pressure difference) in tall compared with small plants, which is significant for our understanding of tall trees' physiology. Finally, we compared isotopic and flux-based approaches for the determination of the link between canopy photosynthesis and ecosystem respiration. We conclude that isotopic approaches are not well suited to document whether changes in photosynthesis of tall trees can rapidly affect soil respiration.
• Photosynthetic rate is concurrently limited by stomatal limitations and nonstomatal limitations (NSLs). However, the controls on NSLs to photosynthesis and their coordination with stomatal control ...on different timescales remain poorly understood. According to a recent optimization hypothesis, NSLs depend on leaf osmotic or water status and are coordinated with stomatal control so as to maximize leaf photosynthesis.
• Drought and notching experiments were conducted on Pinus sylvestris, Picea abies, Betula Pendula and Populus tremula seedlings in glasshouse conditions to study the dependence of NSLs on leaf osmotic and water status, and their coordination with stomatal control, on timescales of minutes and weeks, to test the assumptions and predictions of the optimization hypothesis.
• Both NSLs and stomatal conductance followed power-law functions of leaf osmotic concentration and leaf water potential. Moreover, stomatal conductance was proportional to the square root of soil-to-leaf hydraulic conductance, as predicted by the optimization hypothesis.
• Though the detailed mechanisms underlying the dependence of NSLs on leaf osmotic or water status lie outside the scope of this study, our results support the hypothesis that NSLs and stomatal control are coordinated to maximize leaf photosynthesis and allow the effect of NSLs to be included in models of tree gas-exchange.
Functional relationships between wood density and measures of xylem hydraulic safety and efficiency are ambiguous, especially in wet tropical forests. In this meta‐analysis, we move beyond wood ...density per se and identify relationships between xylem allocated to fibers, parenchyma, and vessels and measures of hydraulic safety and efficiency. We analyzed published data of xylem traits, hydraulic properties and measures of drought resistance from neotropical tree species retrieved from 346 sources. We found that xylem volume allocation to fiber walls increases embolism resistance, but at the expense of specific conductivity and sapwood capacitance. Xylem volume investment in fiber lumen increases capacitance, while investment in axial parenchyma is associated with higher specific conductivity. Dominant tree taxa from wet forests prioritize xylem allocation to axial parenchyma at the expense of fiber walls, resulting in a low embolism resistance for a given wood density and a high vulnerability to drought‐induced mortality. We conclude that strong trade‐offs between xylem allocation to fiber walls, fiber lumen, and axial parenchyma drive drought resistance in neotropical trees. Moreover, the benefits of xylem allocation to axial parenchyma in wet tropical trees might not outweigh the consequential low embolism resistance under more frequent and severe droughts in a changing climate.
The hydraulic safety‐efficiency trade‐off in neotropical trees is strongly related to trade‐offs in wood volume allocation. Fiber wall volume benefits embolism resistance, fiber lumen contributes to sapwood capacitance and axial parenchyma is associated with higher hydraulic conductivity.
Abstract
On-going climate change is increasing the risk of drought stress across large areas worldwide. Such drought events decrease ecosystem productivity and have been increasingly linked to tree ...mortality. Understanding how trees respond to water shortage is key to predicting the future of ecosystem functions. Phloem is at the core of the tree functions, moving resources such as non-structural carbohydrates, nutrients, and defence and information molecules across the whole plant. Phloem function and ability to transport resources is tightly controlled by the balance of carbon and water fluxes within the tree. As such, drought is expected to impact phloem function by decreasing the amount of available water and new photoassimilates. Yet, the effect of drought on the phloem has received surprisingly little attention in the last decades. Here we review existing knowledge on drought impacts on phloem transport from loading and unloading processes at cellular level to possible effects on long-distance transport and consequences to ecosystems via ecophysiological feedbacks. We also point to new research frontiers that need to be explored to improve our understanding of phloem function under drought. In particular, we show how phloem transport is affected differently by increasing drought intensity, from no response to a slowdown, and explore how severe drought might actually disrupt the phloem transport enough to threaten tree survival. Because transport of resources affects other organisms interacting with the tree, we also review the ecological consequences of phloem response to drought and especially predatory, mutualistic and competitive relations. Finally, as phloem is the main path for carbon from sources to sink, we show how drought can affect biogeochemical cycles through changes in phloem transport. Overall, existing knowledge is consistent with the hypotheses that phloem response to drought matters for understanding tree and ecosystem function. However, future research on a large range of species and ecosystems is urgently needed to gain a comprehensive understanding of the question.
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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