Teresa S. David

Drought-induced photosynthetic inhibition and autumn recovery in two Mediterranean oak species (Quercus ilex and Quercus suber)

Responses of leaf water relations and photosynthesis to summer drought and autumn rewetting were studied in two evergreen Mediterranean oak species, Quercus ilex spp. rotundifolia and Quercus suber. The predawn leaf water potential (Ψ(lPD)), stomatal conductance (gs) and photosynthetic rate (A) at ambient conditions were measured seasonally over a 3-year period. We also measured the photosynthetic response to light and to intercellular CO₂ (A/PPFD and A/C(i) response curves) under water stress (summer) and after recovery due to autumn rainfall. Photosynthetic parameters, Vc(max), J(max) and triose phosphate utilization (TPU) rate, were estimated using the Farquhar model. RuBisCo activity, leaf chlorophyll, leaf nitrogen concentration and leaf carbohydrate concentration were also measured. All measurements were performed in the spring leaves of the current year. In both species, the predawn leaf water potential, stomatal conductance and photosynthetic rate peaked in spring, progressively declined throughout the summer and recovered upon autumn rainfall. During the drought period, Q. ilex maintained a higher predawn leaf water potential and stomatal conductance than Q. suber. During this period, we found that photosynthesis was not only limited by stomatal closure, but was also downregulated as a consequence of a decrease in the maximum carboxylation rate (Vc(max)) and the light-saturated rate of photosynthetic electron transport (J(max)) in both species. The Vc(max) and J(max) increased after the first autumnal rains and this increase was related to RuBisCo activity, leaf nitrogen concentration and chlorophyll concentration. In addition, an increase in the TPU rate and in soluble leaf sugar concentration was observed in this period. The results obtained indicate a high resilience of the photosynthetic apparatus to summer drought as well as good recovery in the following autumn rains of these evergreen oak species.

Disentangling drought-induced variation in ecosystem and soil respiration using stable carbon isotopes

Combining C flux measurements with information on their isotopic composition can yield a process-based understanding of ecosystem C dynamics. We studied the variations in both respiratory fluxes and their stable C isotopic compositions (δ13C) for all major components (trees, understory, roots and soil microorganisms) in a Mediterranean oak savannah during a period with increasing drought. We found large drought-induced and diurnal dynamics in isotopic compositions of soil, root and foliage respiration (δ13Cres). Soil respiration was the largest contributor to ecosystem respiration (Reco), exhibiting a depleted isotopic signature and no marked variations with increasing drought, similar to ecosystem respired δ13CO2, providing evidence for a stable C-source and minor influence of recent photosynthate from plants. Short-term and diurnal variations in δ13Cres of foliage and roots (up to 8 and 4‰, respectively) were in agreement with: (1) recent hypotheses on post-photosynthetic fractionation processes, (2) substrate changes with decreasing assimilation rates in combination with increased respiratory demand, and (3) decreased phosphoenolpyruvate carboxylase activity in drying roots, while altered photosynthetic discrimination was not responsible for the observed changes in δ13Cres. We applied a flux-based and an isotopic flux-based mass balance, yielding good agreement at the soil scale, while the isotopic mass balance at the ecosystem scale was not conserved. This was mainly caused by uncertainties in Keeling plot intercepts at the ecosystem scale due to small CO2 gradients and large differences in δ13Cres of the different component fluxes. Overall, stable isotopes provided valuable new insights into the drought-related variations of ecosystem C dynamics, encouraging future studies but also highlighting the need of improved methodology to disentangle short-term dynamics of isotopic composition of Reco.

Synergy of extreme drought and shrub invasion reduce ecosystem functioning and resilience in water-limited climates

Extreme drought events and plant invasions are major drivers of global change that can critically affect ecosystem functioning and alter ecosystem-atmosphere exchange. Invaders are expanding worldwide and extreme drought events are projected to increase in frequency and intensity. However, very little is known on how these drivers may interact to affect the functioning and resilience of ecosystems to extreme events. Using a manipulative shrub removal experiment and the co-occurrence of an extreme drought event (2011/2012) in a Mediterranean woodland, we show that native shrub invasion and extreme drought synergistically reduced ecosystem transpiration and the resilience of key-stone oak tree species. Ecosystem transpiration was dominated by the water use of the invasive shrub Cistus ladanifer, which further increased after the extreme drought event. Meanwhile, the transpiration of key-stone tree species decreased, indicating a competitive advantage in favour of the invader. Our results suggest that in Mediterranean-type climates the invasion of water spending species and projected recurrent extreme drought events may synergistically cause critical drought tolerance thresholds of key-stone tree species to be surpassed, corroborating observed higher tree mortality in the invaded ecosystems. Ultimately, this may shift seasonally water limited ecosystems into less desirable alternative states dominated by water spending invasive shrubs.

Hydraulic connectivity from roots to branches depicted through sap flow: analysis on a Quercus suber tree

The topology of the xylem network is likely to affect the transport of water, propagation of embolism and plant survival and growth. Few studies have been conducted on the hydraulics of the entire water pathway in trees. We evaluated the hydraulic connections from roots to branches in a mature Quercus suber L. tree, through sap flow responses upon branch severing. Sap flow was recorded in branches, stem and roots by the heat field deformation (HFD) method. Results showed that roots, except for the taproot, were hydraulically connected to all branches, but the rest of the tree (stem, branches and taproot) was highly sectored. In the large roots that showed an integrated response to branch severing, the outer xylem was preferentially connected to the same side branch and the inner xylem to the opposite branch. The hydraulic sectoriality in branches, stem and taproot may be regarded as an adaptive trait to water stress. The integrated hydraulic structure of roots is advantageous under patchy soil conditions, but may allow the spread of root diseases. The HFD sap flow method proved extremely useful to calculate xylem flux connectivity between different organs of a large tree, providing a comprehensive picture of its hydraulic architecture.

The Water Isotopic Version of the Land-Surface Model ORCHIDEE: Implementation, Evaluation, Sensitivity to Hydrological Parameters

Land-Surface Models (LSMs) exhibit large spread and uncertainties in the way they partition precipitation into surface runoff, drainage, transpiration and bare soil evaporation. To explore to what extent water isotope measurements could help evaluate the simulation of the soil water budget in LSMs, water stable isotopes have been implemented in the ORCHIDEE (ORganizing Carbon and Hydrology In Dynamic EcosystEms: the land-surface model) LSM. This article presents this implementation and the evaluation of simulations both in a stand-alone mode and coupled with an atmospheric general circulation model. ORCHIDEE simulates reasonably well the isotopic composition of soil, stem and leaf water compared to local observations at ten measurement sites. When coupled to LMDZ (Laboratoire de Meteorologie Dynamique-Zoom: the atmospheric model), it simulates well the isotopic composition of precipitation and river water compared to global observations. Sensitivity tests to LSM (Land-Surface Model) parameters are performed to identify processes whose representation by LSMs could be better evaluated using water isotopic measurements. We find that measured vertical variations in soil water isotopes could help evaluate the representation of infiltration pathways by multi-layer soil models. Measured water isotopes in rivers could help calibrate the partitioning of total runoff into surface runoff and drainage and the residence time scales in underground reservoirs. Finally, co-located isotope measurements in precipitation, vapor and soil water could help estimate the partitioning of infiltrating precipitation into bare soil evaporation.

Root Function: In Situ Studies Through Sap Flow Research

Sap flow measured by the Heat Field Deformation technique, HFD, is sensitive to flow responses to small changes in water potential gradients within the tree hydraulic systems. When these changes occur abruptly, under experimental treatments (severing, localized irrigation, heavy loading), sap flow movement can be used as a marker to study root functionality, for example root ability to redistribute water and withstand heavy machinery pressure. Experiments also showed that a compensation mechanism may operate in trees, with a temporary increase in the absorbed water due to a preferential use of one part of the root system when another part is damaged or when a water source is lost. Long-term measurements of root sap flow allow distinguishing between water uptake from shallow and deep rooted trees, at different exposures at a forest edge and from healthy and infected trees. Root sap flow can be used as an indicator of tree stress or of the prevailing mechanisms used by trees to survive drought, under irrigation or rain-fed conditions.

Sap flow index as an indicator of water storage use

Abstract Symmetrical temperature difference also known as the sap flow index (SFI) forms the basis of the Heat Field Deformation sap flow measurement and is simultaneously collected whilst measuring the sap flow. SFI can also be measured by any sap flow method applying internal continuous heating through the additional installation of an axial differential thermocouple equidistantly around a heater. In earlier research on apple trees SFI was found to be an informative parameter for tree physiological studies, namely for assessing the contribution of stem water storage to daily transpiration. The studies presented in this work are based on the comparative monitoring of SFI and diameter in stems of different species (Pseudotsuga menziesii, Picea omorika, Pinus sylvestris) and tree sizes. The ability of SFI to follow the patterns of daily stem water storage use was empirically confirmed by our data. Additionally, as the HFD multipointsensors can measure sap flow at several stem sapwood depths, their use allowed to analyze the use of stored water in different xylem layers through SFI records. Radial and circumferential monitoring of SFI on large cork oak trees provided insight into the relative magnitude and timing of the contribution of water stored in different sapwood layers or stem sectors to transpiration.

Root Structure: In Situ Studies Through Sap Flow Research

Sap flow research highlights new perspectives to study in situ root structure of large trees. Several examples demonstrate the ability of the Heat Field Deformation method, HFD, to do this under natural and experimental conditions. Within the latter, localized irrigation, sink- or source-severing trigger sap flow responses that help us to understand the belowground parts of a tree, such as the presence of anastomoses between roots of different trees. The vertical profile of root density, as well as root size around a tree, can be derived from the stem sap flow radial profile. Increase of stem flow due to localized irrigation may be used to distinguish root locations near the corresponding stem sector. Responses of root or stem sap flow when exposing roots using an air-spade or following the severing of roots or branches help us to understand the relationships between different sapwood conducting layers and paths of water between sources and sinks.