Acclimation of hydraulic and morphological traits to water deficit delays hydraulic failure during simulated drought in poplar.

The capacity of trees to tolerate and survive increasing drought conditions in situ will depend in part on their ability to acclimate (via phenotypic plasticity) key hydraulic and morphological traits that increase drought tolerance and delay the onset of drought-induced hydraulic failure. However, the effect of water-deficit acclimation in key traits that determine time to hydraulic failure (THF) during extreme drought remains largely untested. We measured key hydraulic and morphological traits in saplings of a hybrid poplar grown under well-watered and water-limited conditions. The time for plants to dry-down to critical levels of water stress (90% loss of stem hydraulic conductance), as well as the relative contribution of drought acclimation in each trait to THF, was simulated using a soil-plant hydraulic model (SurEau). Compared with controls, water-limited plants exhibited significantly lower stem hydraulic vulnerability (P50stem), stomatal conductance and total canopy leaf area (LA). Taken together, adjustments in these and other traits resulted in longer modelled THF in water-limited (~160 h) compared with well-watered plants (~50 h), representing an increase of more than 200%. Sensitivity analysis revealed that adjustment in P50stem and LA contributed the most to longer THF in water-limited plants. We observed a high degree of trait plasticity in poplar saplings in response to water-deficit growth conditions, with decreases in stem hydraulic vulnerability and leaf area playing a key role in delaying the onset of hydraulic failure during a simulated drought event. These findings suggest that understanding the capacity of plants to acclimate to antecedent growth conditions will enable better predictions of plant survivorship during future drought.

Plasticity of the xylem vulnerability to embolism in Populus tremula x alba relies on pit quantity properties rather than on pit structure.

Knowledge on variations of drought resistance traits are needed to predict the potential of trees to acclimate to coming severe drought events. Xylem vulnerability to embolism is a key parameter related to such droughts, and its phenotypic variability relies mainly on environmental plasticity. We investigated the structural determinants controlling the plasticity of vulnerability to embolism, focusing on the key elements involved in the air bubble entry in vessels, especially the intervessel pits. Poplar saplings (Populus tremula x alba (Aiton) Sm., 1804) grown in contrasted water availability or light exposure exhibited differences in the vulnerability to embolism (P50) in a range of 0.76 MPa. We then characterized the structural changes in features related to pit quantity and pit structure, from the pit ultrastructure to the organization of xylem vessels, using different microscopy techniques (transmission electron microscopy, scanning electron microscopy, light microscopy). A multispectral combination of X-ray microtomography and light microscopy analysis allowed measuring the vulnerability of each single vessel and testing some of the relationships between structural traits and vulnerability to embolism inside the xylem. The pit ultrastructure did not change, whereas the vessel dimensions increased with the vulnerability to embolism and the grouping index and fraction of intervessel cell wall both decreased with the vulnerability to embolism. These findings hold when comparing between trees or between the vessels inside the xylem of an individual tree. These results evidenced that plasticity of vulnerability to embolism in hybrid poplar occurs through changes in the pit quantity properties such as pit area and vessel grouping rather than changes on the pit structure.

Assessing inter- and intraspecific variability of xylem vulnerability to embolism in oaks.

The genus Quercus comprises important species in forestry not only for their productive value but also for their ability to withstand drought. Hence an evaluation of inter- and intraspecific variation in drought tolerance is important for selecting the best adapted species and provenances for future afforestation. The presence of long vessels makes it difficult to assess xylem vulnerability to embolism in oak. Thanks to the development of an in situ flow centrifuge equipped with a large rotor, we quantified (i) the between species variability of embolism resistance in four native and two exotic species of oaks in Europe and (ii) the within species variability in Quercus petraea. Embolism resistance varied significantly among species, with the pressure inducing 50% loss of hydraulic conductivity (P50 ) ranging between - 7.0 and -4.2 MPa. Species native to the Mediterranean region were more resistant than pan-European species. In contrast, intraspecific variability in embolism resistance in Q. petraea was low within provenances and null among provenances. A positive correlation between P50 and vessel diameter among the six oak species indicates that the more embolism resistant species had narrower xylem vessels. However, this tradeoff between hydraulic efficiency and safety was not observed between Q. petraea provenances.

PtxtPME1 and homogalacturonans influence xylem hydraulic properties in poplar.

While the xylem hydraulic properties, such as vulnerability to cavitation (VC), are of paramount importance in drought resistance, their genetic determinants remain unexplored. There is evidence that pectins and their methylation pattern are involved, but the detail of their involvement and the corresponding genes need to be clarified. We analyzed the hydraulic properties of the 35S::PME1 transgenic aspen that ectopically under- or over-express a xylem-abundant pectin methyl esterase, PtxtPME1. We also produced and analyzed 4CL1::PGII transgenic poplars expressing a fungal polygalacturonase, AnPGII, under the control of the Ptxa4CL1 promoter that is active in the developing xylem after xylem cell expansion. Both the 35S::PME1 under- and over-expressing aspen lines developed xylem with lower-specific hydraulic conductivity and lower VC, while the 4CL1::PGII plants developed xylem with a higher VC. These xylem hydraulic changes were associated with modifications in xylem structure or in intervessel pit structure that can result in changes in mechanical behavior of the pit membrane. This study shows that homogalacturonans and their methylation pattern influence xylem hydraulic properties, through its effect on xylem cell expansion and on intervessel pit properties and it show a role for PtxtPME1 in the xylem hydraulic properties.