Drivers of nocturnal stomatal conductance in C3 and C4 plants.

Nocturnal water losses were for long considered negligible, but it is now known that incomplete stomatal closure during the night leads to significant water losses at leaf, plant and ecosystem scales. However, only daytime transpiration is currently accounted for in evapotranspiration studies. Important uncertainties on the drivers of nocturnal water fluxes hinder its incorporation within modelling frameworks because some studies indicate that night-time stomatal drivers may differ from day-time responses. Here, we synthesise the studies on nocturnal stomatal conductance (gn) to determine underlying drivers through a systematic literature review and, whenever possible, meta-analytical techniques. Similar to daytime responses, we found negative effects of vapour pressure deficit, predawn water potential, air temperature, and salinity on gn across the plant species. However, the most apparent trend was an increase of gn from the beginning until the end of the night, indicating significant and widespread endogenous regulation by the circadian clock. We further observed how neither elevated CO2 nor nutrient status affected gn significantly across species. We also did not find any significant associations between gn and elevated ozone or increasing plant age. There was a paucity of studies on climatic extremes such heat waves and also few studies connected gn with anatomical features such as leaf specific area or stomatal density. Further studies are also needed to address the effects of plant sex, abscisic acid concentrations and genotypic variations on gn. Our findings solve the long-term conundrum on whether stomatal responses to daytime drivers are the same as those that during the nighttime.

Radiation and Drought Impact Residual Leaf Conductance in Two Oak Species With Implications for Water Use Models.

Stomatal closure is one of the earliest responses to water stress but residual water losses may continue through the cuticle and incomplete stomatal closure. Residual conductance (g res ) plays a large role in determining time to mortality but we currently do not understand how do drought and shade interact to alter g res because the underlying drivers are largely unknown. Furthermore, g res may play an important role in models of water use, but the exact form in which g res should be incorporated into modeling schemes is currently being discussed. Here we report the results of a study where two different oak species were experimentally subjected to highly contrasting levels of drought (resulting in 0, 50 and 80% losses of hydraulic conductivity) and radiation (photosynthetic photon flux density at 1,500 µmol m-2 s-1 or 35-45 µmol m-2 s-1). We observed that the effects of radiation and drought were interactive and species-specific and g res correlated positively with concentrations of leaf non-structural carbohydrates and negatively with leaf nitrogen. We observed that different forms of measuring g res , based on either nocturnal conductance under high atmospheric water demand or on the water mass loss of detached leaves, exerted only a small influence on a model of stomatal conductance and also on a coupled leaf gas exchange model. Our results indicate that, while understanding the drivers of g res and the effects of different stressors may be important to better understand mortality, small differences in g res across treatments and measurements exert only a minor impact on stomatal models in two closely related species.

Hydraulic and photosynthetic limitations prevail over root non-structural carbohydrate reserves as drivers of resprouting in two Mediterranean oaks.

Resprouting is an ancestral trait in angiosperms that confers resilience after perturbations. As climate change increases stress, resprouting vigor is declining in many forest regions, but the underlying mechanism is poorly understood. Resprouting in woody plants is thought to be primarily limited by the availability of non-structural carbohydrate reserves (NSC), but hydraulic limitations could also be important. We conducted a multifactorial experiment with two levels of light (ambient, 2-3% of ambient) and three levels of water stress (0, 50 and 80 percent losses of hydraulic conductivity, PLC) on two Mediterranean oaks (Quercus ilex and Q. faginea) under a rain-out shelter (n = 360). The proportion of resprouting individuals after canopy clipping declined markedly as PLC increased for both species. NSC concentrations affected the response of Q. ilex, the species with higher leaf construction costs, and its effect depended on the PLC. The growth of resprouting individuals was largely dependent on photosynthetic rates for both species, while stored NSC availability and hydraulic limitations played minor and non-significant roles, respectively. Contrary to conventional wisdom, our results indicate that resprouting in oaks may be primarily driven by complex interactions between hydraulics and carbon sources, whereas stored NSC play a significant but secondary role.