-Since about twenty years, hydraulic architecture (h.a.) is, doubtless, the major trend in the domain of plants (and especially trees) water relations. This review encompasses the main concepts and results concerning the hydraulic of architecture of trees. After a short paragraph about the definition of the h.a., the qualitative and quantitative characteristics of the h.a. are presented. This is an occasion to discuss the pipe model from the h.a. point of view. The second part starts with the central concept of embolism and give a review of important experimental results and questions concerning summer and winter embolism. The last part deals with the coupling between hydraulic and stomatal conductances. It discusses the theoretical and experimental relationships between transpiration and leaf water potential during a progressive soil drought, the increase of soil-root resistance and its consequences in term of xylem vulnerability, the factors controlling the daily maximum transpiration and how stomates can prevent "run away embolism". In conclusion different kinds of unsolved questions of h.a., which can be a matter of future investigations, are presented in addition with a classification of trees behaviour under drought conditions. To end, an appendix recalls the notions of water potential, pressure and tension.hydraulic architecture / cohesion-tension theory / summer embolism / winter embolism / drought resistance Résumé -Architecture hydraulique des arbres : concepts principaux et résultats. Sans aucun doute, depuis une vingtaine d'années, l'architecture hydraulique (a.h.) est devenue une approche majeure dans le domaine des relations plantes-eau (et particulièrement pour les arbres). Cette revue présente les principaux concepts et résultats concernant l'a.h. Après un bref paragraphe sur la définition de l'a.h., les caractéristiques qualitatives et quantitatives définissant l'a.h. sont passées en revue. À cette occasion le « pipe model » est discuté du point de vue de l'a.h. La seconde partie commence avec le concept central d'embolie et continue avec une présentation des principaux résultats et questions touchant l'embolie estivale et l'embolie hivernale. La dernière partie analyse le « couplage » entre les conductances hydraulique et stomatique. Il y est discuté des relations théoriques et expérimentales entre la transpiration et le potentiel hydrique foliaire durant la mise en place d'une sécheresse progressive du sol, de l'augmentation de la résistance sol-racines et de ses conséquences en terme de vulnérabilité du xylème, des facteurs contrôlant la transpiration maximale journalière et de quelle manière les stomates peuvent prévenir l'emballement de l'embolie. La conclusion fait état de différentes questions non résolues, qui pourraient faire l'objet de recherches futures et esquisse une classification du comportement des arbres vis-à-vis de la sécheresse. Pour finir, un appendice rappelle les notions de potentiel hydrique, de pression et de tension.architecture hydraulique / théorie de la cohé...

Trees of Juglans regia L. shed leaves when subjected to drought. Before shedding (when leaves are yellow), the petioles have lost 87% of their maximum hydraulic conductivity, but stems have lost only 14% of their conductivity. This is caused by the higher vulnerability of petioles than stems to water‐stress induced cavitation. These data are discussed in the context of the plant segmentation hypothesis.

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.The "cohesion-tension" (CT) theory of sap ascent in plants was proposed more than a century ago by Bö hm (1893) and Dixon and Joly (1894). The theory postulates that (a) the xylem conduits form continuous water columns from the roots to the leaves, (b) the columns are held in place thanks to the capillary pressures that develop in the leaf mesophyll, and (c) leaf transpiration pulls water out of the xylem, which causes water absorption by the roots. A corollary of the theory is that high xylem tensions (negative pressures) must develop inside the xylem conduits. Over the past century a considerable amount of experimental data have been cumulated by plant physiologists, all consistent with the CT theory.However, recent direct measurements of sap pressure with xylem pressure probes (Zimmermann et al., 1994) and direct cryo-scanning electron microscopy (cryo-SEM) observations of xylem vessels content during transpiration (Canny, 1997b(Canny, , 1998b have questioned the validity of the CT theory. New experiments with the xylem pressure probe ...

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.

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.