Generalized Münch coupling between sugar and water fluxes for modelling carbon allocation as affected by water status.

A model of within-plant carbon allocation is proposed which makes a generalized use of the Münch mechanism to integrate carbon and water functions and their involvement in growth limitations. The plant is envisioned as a branched network of resistive pathways (phloem and xylem) with nodal organs acting as sources and sinks for sucrose. Four elementary organs (leaf, stem, fruit, root) are described with their particular sink functions and hydraulic attributes. Given the rates of photosynthesis and transpiration and the hydraulic properties of the network as inputs, the model calculates the internal fluxes of water and sucrose. Xylem water potential (Psi), phloem sucrose concentration (C) and turgor pressure (P) are calculated everywhere in the network accounting for osmotic equilibrium between apoplasm and symplasm and coupled functioning of xylem and phloem. The fluxes of phloem and xylem saps are driven by the gradients of P and Psi, respectively. The fruit growth rate is assumed as turgor pressure dependent. To demonstrate its ability to address within-plant competition, the model is run with a simple-branched structure gathering three leaves, eight stem segments, three competing growing fruits and one root. The model was programmed with P-Spice, a software specifically designed for simulating electrical circuits but easily adaptable to physiology. Simulations of internal water fluxes, sucrose concentrations and fruit growth rates are given for different conditions of soil water availability and hydraulic resistances (sensitivity analysis). The discussion focuses on the potential interest of this approach in functional--structural plant models to address water stress-induced effects.

Seasonal variation in xylem pressure of walnut trees: root and stem pressures.

Measurements of air and soil temperatures and xylem pressure were made on 17-year-old orchard trees and on 5-year-old potted trees of walnut (Juglans regia L.). Cooling chambers were used to determine the relationships between temperature and sugar concentration ([glucose] + [fructose] + [sucrose], GFS) and seasonal changes in xylem pressure development. Pressure transducers were attached to twigs of intact plants, root stumps and excised shoots while the potted trees were subjected to various temperature regimes in autumn, winter and spring. Osmolarity and GFS of the xylem sap (apoplast) were measured before and after cooling or warming treatments. In autumn and spring, xylem pressures of up to 160 kPa were closely correlated with soil temperature but were not correlated with GFS in xylem sap. High root pressures were associated with uptake of mineral nutrients from soil, especially nitrate. In autumn and spring, xylem pressures were detected in root stumps as well as in intact plants, but not in excised stems. In contrast, in winter, 83% of the xylem sap osmolarity in both excised stems and intact plants could be accounted for by GFS, and both GFS and osmolarity were inversely proportional to temperature. Plants kept at 1.5 degrees C developed positive xylem pressures up to 35 kPa, xylem sap osmolarities up to 260 mosmol l(-1) and GFS concentrations up to 70 g l(-1). Autumn and spring xylem pressures, which appeared to be of root origin, were about 55% of the theoretical pressures predicted by osmolarity of the xylem sap. In contrast, winter pressures appeared to be of stem origin and were only 7% of the theoretical pressures, perhaps because of a lower stem water content during winter.

Winter stem xylem pressure in walnut trees: effects of carbohydrates, cooling and freezing.

Pressure transducers were attached to twigs of orchard trees and potted trees of walnut (Juglans regia L.) to measure winter stem xylem pressures. Experimental potted trees were partially defoliated in the late summer and early autumn to lower the amount of stored carbohydrates. Potted trees were placed in cooling chambers and subjected to various temperature regimes, including freeze-thaw cycles. Xylem pressures were inversely proportional to the previous 48-h air temperature, but positively correlated with the osmolarity of the xylem sap. Defoliated trees had significantly lower concentrations of stored carbohydrates and significantly lower xylem sap osmolarities than controls. Plants kept at 1.5 degrees C developed xylem pressures up to 40 kPa, just 7% of the theoretical osmotic pressure of the xylem sap. However, exposure to low, nonfreezing temperatures followed by freeze-thaw cycles resulted in pressures over 210 kPa, which was 39% of the theoretical osmotic pressure. A simple osmotic model could account for the modest positive winter pressures at low, nonfreezing temperatures, but not for the synergistic effects of freeze-thaw cycles.

Cryo-scanning electron microscopy observations of vessel content during transpiration in walnut petioles. Facts or artifacts?

The current controversy about the "cohesion-tension" of water ascent in plants arises from the recent cryo-scanning electron microscopy (cryo-SEM) observations of xylem vessels content by Canny and coworkers (1995). On the basis of these observations it has been claimed that vessels were emptying and refilling during active transpiration in direct contradiction to the previous theory. In this study we compared the cryo-SEM data with the standard hydraulic approach on walnut (Juglans regia) petioles. The results of the two techniques were in clear conflict and could not both be right. Cryo-SEM observations of walnut petioles frozen intact on the tree in a bath of liquid nitrogen (LN(2)) suggested that vessel cavitation was occurring and reversing itself on a diurnal basis. Up to 30% of the vessels were embolized at midday. In contrast, the percentage of loss of hydraulic conductance (PLC) of excised petiole segments remained close to 0% throughout the day. To find out which technique was erroneous we first analyzed the possibility that PLC values were rapidly returned to zero when the xylem pressures were released. We used the centrifugal force to measure the xylem conductance of petiole segments exposed to very negative pressures and established the relevance of this technique. We then analyzed the possibility that vessels were becoming partially air-filled when exposed to LN(2). Cryo-SEM observations of petiole segments frozen shortly after their xylem pressure was returned to atmospheric values agreed entirely with the PLC values. We confirmed, with water-filled capillary tubes exposed to a large centrifugal force, that it was not possible to freeze intact their content with LN(2). We concluded that partially air-filled conduits were artifacts of the cryo-SEM technique in our study. We believe that the cryo-SEM observations published recently should probably be reconsidered in the light of our results before they may be used as arguments against the cohesion-tension theory.

Novel Methods of Measuring Hydraulic Conductivity of Tree Root Systems and Interpretation Using AMAIZED (A Maize-Root Dynamic Model for Water and Solute Transport).

Steady-state and dynamic methods were used to measure the conductivity to water flow in large woody root systems. The methods were destructive in that the root must be excised from the shoot but do not require removal of the root from the soil. The methods involve pushing water from the excised base of the root to the apex, causing flow in a direction opposite to that during normal transpiration. Sample data are given for two tropical (Cecropia obtusifolia and Lacistema aggregatum) and two temperate species (Acer saccharum and Juglans regia cv Lara). A hysteresis was observed in the relationship between applied pressure and resulting flow during dynamic measurements. A mathematical model (AMAIZED) was derived for the dynamics of solute and water flow in roots. The model was used to interpret results obtained from steady-state and dynamic measurements. AMAIZED is mathematically identical with the equations that describe Munch pressure flow of solute and water in the phloem of leaves. Results are discussed in terms of the predictions of AMAIZED, and suggestions for the improvement of methods are made.

Use of positive pressures to establish vulnerability curves : further support for the air-seeding hypothesis and implications for pressure-volume analysis.

Loss of hydraulic conductivity occurs in stems when the water in xylem conduits is subjected to sufficiently negative pressure. According to the air-seeding hypothesis, this loss of conductivity occurs when air bubbles are sucked into water-filled conduits through micropores adjacent to air spaces in the stem. Results in this study showed that loss of hydraulic conductivity occurred in stem segments pressurized in a pressure chamber while the xylem water was under positive pressure. Vulnerability curves can be defined as a plot of percentage loss of hydraulic conductivity versus the pressure difference between xylem water and the outside air inducing the loss of conductivity. Vulnerability curves were similar whether loss of conductivity was induced by lowering the xylem water pressure or by raising the external air pressure. These results are consistent with the air-seeding hypothesis of how embolisms are nucleated, but not with the nucleation of embolisms at hydrophobic cracks because the latter requires negative xylem water pressure. The results also call into question some basic underlying assumptions used in the determination of components of tissue water potential using "pressure-volume" analysis.