Amplified Drought Alters Leaf Litter Metabolome, Slows Down Litter Decomposition, and Modifies Home Field (Dis)Advantage in Three Mediterranean Forests.

In Mediterranean ecosystems, the projected rainfall reduction of up to 30% may alter plant-soil interactions, particularly litter decomposition and Home Field Advantage (HFA). We set up a litter transplant experiment in the three main forests encountered in the northern part of the Medi-terranean Basin (dominated by either Quercus ilex, Quercus pubescens, or Pinus halepensis) equipped with a rain exclusion device, allowing an increase in drought either throughout the year or concentrated in spring and summer. Senescent leaves and needles were collected under two precipitation treatments (natural and amplified drought plots) at their "home" forest and were left to decompose in the forest of origin and in other forests under both drought conditions. MS-based metabolomic analysis of litter extracts combined with multivariate data analysis enabled us to detect modifications in the composition of litter specialized metabolites, following amplified drought treatment. Amplified drought altered litter quality and metabolomes, directly slowed down litter decomposition, and induced a loss of home field (dis)advantage. No indirect effect mediated by a change in litter quality on decomposition was observed. These results may suggest major alterations of plant-soil interactions in Mediterranean forests under amplified drought conditions.

Drought acclimation of Quercus ilex leaves improves tolerance to moderate drought but not resistance to severe water stress.

Increasing temperature and drought can result in leaf dehydration and defoliation even in drought-adapted tree species such as the Mediterranean evergreen Quercus ilex L. The stomatal regulation of leaf water potential plays a central role in avoiding this phenomenon and is constrained by a suite of leaf traits including hydraulic conductance and vulnerability, hydraulic capacitance, minimum conductance to water vapour, osmotic potential and cell wall elasticity. We investigated whether the plasticity in these traits may improve leaf tolerance to drought in two long-term rainfall exclusion experiments in Mediterranean forests. Osmotic adjustment was observed to lower the water potential at turgor loss in the rainfall-exclusion treatments, thus suggesting a stomatal closure at more negative water potentials and a more anisohydric behaviour in drier conditions. Conversely, leaf hydraulic conductance and vulnerability did not exhibit any plasticity between treatments so the hydraulic safety margins were narrower in the rainfall-exclusion treatments. The sequence of leaf responses to seasonal drought and dehydration was conserved among treatments and sites but trees were more likely to suffer losses of turgor and hydraulic functioning in the rainfall-exclusion treatments. We conclude that leaf plasticity might help the trees to tolerate moderate drought but not to resist severe water stress.

Holm oak fecundity does not acclimate to a drier world.

Climate change might impact tree fecundity by altering the relative influences of meteorological and physiological drivers, and by modifying resource investment in reproduction. Using a 13-yr monitoring of Quercus ilex reproduction in a rainfall exclusion experiment, we analysed the interactive effects of long-term increased aridity and other environmental drivers on the inter-annual variation of fecundity (male flower biomass, number of initiated and mature fruits). Summer-autumn water stress was the main driver of fruit abortion during fruit growth. Rainfall exclusion treatment strongly reduced the number of initiated and mature fruits, even in masting years, and did not increase fruit tolerance to severe drought. Conversely, the relative contribution of the meteorological and physiological drivers, and the inter-annual variability of fruit production were not modified by rainfall exclusion. Rather than inducing an acclimation of tree fecundity to water limitation, increased aridity impacted it negatively through both lower fruit initiation due to changes in resource allocation, and more severe water and resource limitations during fruit growth. Long-term increased aridity affected tree reproduction beyond what is expected from the current response to inter-annual drought variations, suggesting that natural regeneration of holm oak forest could be jeopardised in the future.

Substratum influences uptake of radium-226 by plants.

Radium-226, an alpha emitter with half-life 1600 years, is ubiquitous in natural environments. Present in rocks and soils, it is also absorbed by vegetation. The efficiency of 226Ra uptake by plants from the soil is important to assess for the study of heavy metals uptake by plants, monitoring of radioactive pollution, and the biogeochemical cycle of radium in the Critical Zone. Using a thoroughly validated measurement method of effective 226Ra concentration (ECRa) in the laboratory, we compare ECRa values of the plant to that of the closest soil, and we infer the 226Ra soil-to-plant transfer ratio, RSP, for a total of 108 plant samples collected in various locations in France. ECRa values of plants range over five orders of magnitude with mean (min-max) of 1.66 ± 0.03 (0.020-113) Bq kg-1. Inferred RSP values range over four orders of magnitude with mean (min-max) of 0.0188 ± 0.0004 (0.00069-0.37). The mean RSP value of plants in granitic and metamorphic context (0.073 ± 0.002; n = 50) is significantly higher (12 ± 1 times) than that of plants in calcareous and sedimentary context (0.0058 ± 0.0002; n = 58). This difference, which cannot be attributed to a systematic difference in emanation coefficient, is likely due to the competition between calcium and radium. In a given substratum context, the compartments of a given plant species show coherent and decreasing RSP values in the following order (acropetal gradient): roots > bark > branches and stems ≈ leaves. Oak trees (Quercus genus) concentrate 226Ra more than other trees and plants in this set. While this study clearly demonstrates the influence of substratum on the 226Ra uptake by plants in non-contaminated areas, our measurement method appears as a promising practical tool to use for (phyto)remediation and its monitoring in uranium- and radium-contaminated areas.

Rainfall exclusion and thinning can alter the relationships between forest functioning and drought.

Increasing drought caused by the ongoing climate change, and forest management by thinning that aims at mitigating its impact, may modify the current relationships between forest functions and drought intensity and preclude our ability to forecast future ecosystem responses. We used 15 yr of data from an experimental rainfall exclusion (-27% of rainfall) combined with thinning (-30% stand basal area) to investigate differences in the drought-function relationships for each component of above-ground net primary productivity (ANPP) and stand transpiration in a Mediterranean Quercus ilex stand. Rainfall exclusion reduced stand ANPP by 10%, mainly because of lowered leaf and acorn production, whereas wood production remained unaffected. These responses were consistent with the temporal sensitivity to drought among tree organs but revealed an increased allocation to wood. Thinning increased wood and acorn production and reduced the sensitivity of standing wood biomass change to drought. Rainfall exclusion and thinning lowered the intercept of the transpiration-drought relationship as a result of the structural constraints exerted by lower leaf and sapwood area. The results suggest that historical drought-function relationships can be used to infer future drought impacts on stand ANPP but not on water fluxes. Thinning can mitigate drought effects and reduce forest sensitivity to drought.

Stem hydraulic capacitance decreases with drought stress: implications for modelling tree hydraulics in the Mediterranean oak Quercus ilex.

Hydraulic modelling is a primary tool to predict plant performance in future drier scenarios. However, as most tree models are validated under non-stress conditions, they may fail when water becomes limiting. To simulate tree hydraulic functioning under moist and dry conditions, the current version of a water flow and storage mechanistic model was further developed by implementing equations that describe variation in xylem hydraulic resistance (RX ) and stem hydraulic capacitance (CS ) with predawn water potential (ΨPD ). The model was applied in a Mediterranean forest experiencing intense summer drought, where six Quercus ilex trees were instrumented to monitor stem diameter variations and sap flow, concurrently with measurements of predawn and midday leaf water potential. Best model performance was observed when CS was allowed to decrease with decreasing ΨPD . Hydraulic capacitance decreased from 62 to 25 kg m-3 MPa-1 across the growing season. In parallel, tree transpiration decreased to a greater extent than the capacitive water release and the contribution of stored water to transpiration increased from 2.0 to 5.1%. Our results demonstrate the importance of stored water and seasonality in CS for tree hydraulic functioning, and they suggest that CS should be considered to predict the drought response of trees with models.

Recent climate hiatus revealed dual control by temperature and drought on the stem growth of Mediterranean Quercus ilex.

A better understanding of stem growth phenology and its climate drivers would improve projections of the impact of climate change on forest productivity. Under a Mediterranean climate, tree growth is primarily limited by soil water availability during summer, but cold temperatures in winter also prevent tree growth in evergreen forests. In the widespread Mediterranean evergreen tree species Quercus ilex, the duration of stem growth has been shown to predict annual stem increment, and to be limited by winter temperatures on the one hand, and by the summer drought onset on the other hand. We tested how these climatic controls of Q. ilex growth varied with recent climate change by correlating a 40-year tree ring record and a 30-year annual diameter inventory against winter temperature, spring precipitation, and simulated growth duration. Our results showed that growth duration was the best predictor of annual tree growth. We predicted that recent climate changes have resulted in earlier growth onset (-10 days) due to winter warming and earlier growth cessation (-26 days) due to earlier drought onset. These climatic trends partly offset one another, as we observed no significant trend of change in tree growth between 1968 and 2008. A moving-window correlation analysis revealed that in the past, Q. ilex growth was only correlated with water availability, but that since the 2000s, growth suddenly became correlated with winter temperature in addition to spring drought. This change in the climate-growth correlations matches the start of the recent atmospheric warming pause also known as the 'climate hiatus'. The duration of growth of Q. ilex is thus shortened because winter warming has stopped compensating for increasing drought in the last decade. Decoupled trends in precipitation and temperature, a neglected aspect of climate change, might reduce forest productivity through phenological constraints and have more consequences than climate warming alone.

Environmental control of carbon allocation matters for modelling forest growth.

We aimed to evaluate the importance of modulations of within-tree carbon (C) allocation by water and low-temperature stress for the prediction of annual forest growth with a process-based model. A new C allocation scheme was implemented in the CASTANEA model that accounts for lagged and direct environmental controls of C allocation. Different approaches (static vs dynamic) to modelling C allocation were then compared in a model-data fusion procedure, using satellite-derived leaf production estimates and biometric measurements at c. 104 sites. The modelling of the environmental control of C allocation significantly improved the ability of CASTANEA to predict the spatial and year-to-year variability of aboveground forest growth along regional gradients. A significant effect of the previous year's water stress on the C allocation to leaves and wood was reported. Our results also are consistent with a prominent role of the environmental modulation of sink demand in the wood growth of the studied species. Data available at large scales can inform forest models about the processes driving annual and seasonal C allocation. Our results call for a greater consideration of C allocation drivers, especially sink-demand fluctuations, for the simulations of current and future forest productivity with process-based models.

Few multiyear precipitation-reduction experiments find a shift in the productivity-precipitation relationship.

Well-defined productivity-precipitation relationships of ecosystems are needed as benchmarks for the validation of land models used for future projections. The productivity-precipitation relationship may be studied in two ways: the spatial approach relates differences in productivity to those in precipitation among sites along a precipitation gradient (the spatial fit, with a steeper slope); the temporal approach relates interannual productivity changes to variation in precipitation within sites (the temporal fits, with flatter slopes). Precipitation-reduction experiments in natural ecosystems represent a complement to the fits, because they can reduce precipitation below the natural range and are thus well suited to study potential effects of climate drying. Here, we analyse the effects of dry treatments in eleven multiyear precipitation-manipulation experiments, focusing on changes in the temporal fit. We expected that structural changes in the dry treatments would occur in some experiments, thereby reducing the intercept of the temporal fit and displacing the productivity-precipitation relationship downward the spatial fit. The majority of experiments (72%) showed that dry treatments did not alter the temporal fit. This implies that current temporal fits are to be preferred over the spatial fit to benchmark land-model projections of productivity under future climate within the precipitation ranges covered by the experiments. Moreover, in two experiments, the intercept of the temporal fit unexpectedly increased due to mechanisms that reduced either water loss or nutrient loss. The expected decrease of the intercept was observed in only one experiment, and only when distinguishing between the late and the early phases of the experiment. This implies that we currently do not know at which precipitation-reduction level or at which experimental duration structural changes will start to alter ecosystem productivity. Our study highlights the need for experiments with multiple, including more extreme, dry treatments, to identify the precipitation boundaries within which the current temporal fits remain valid.

Explanatory ecological factors for the persistence of desiccation-sensitive seeds in transient soil seed banks: Quercus ilex as a case study.

BACKGROUND AND AIMS: Dominant tree species in northern temperate forests, for example oak and beech, produce desiccation-sensitive seeds. Despite the potentially major influence of this functional trait on the regeneration and distribution of species under climate change, little is currently known about the ecological determinants of the persistence of desiccation-sensitive seeds in transient soil seed banks. Knowing which key climatic and microsite factors favour seed survival will help define the regeneration niche for species whose seeds display extreme sensitivity to environmental stress METHODS: Using the Mediterranean Holm oak (Quercus ilex) forest as a model system, an in situ time-course monitoring of seed water status and viability was performed during the unfavourable winter season in two years with contrasting rainfall, at an instrumented site with detailed climate records. In parallel, the characteristics of the microhabitat and their influence on the post-winter water status and viability of seeds were investigated in a regional survey of 33 woodlands representative of the French distribution of the species. KEY RESULTS: Time-course monitoring of seed water status in natural conditions confirmed that in situ desiccation is the main abiotic cause of mortality in winter. Critical water contents could be reached in a few days during drought spells. Seed dehydration rates were satisfactorily estimated using integrative climate proxies including vapour pressure deficit and potential evapotranspiration. Seed water status was therefore determined by the balance between water uptake after a rainfall event and water loss during dry periods. Structural equation modelling of microhabitat factors highlighted the major influence of canopy openness and resulting incident radiation on the ground. CONCLUSIONS: This study provides part of the knowledge required to implement species distribution models which incorporate their regeneration niche. It is an important step forward in evaluating the ecological consequences of increasing winter drought and environmental filtering due to climate change on the regeneration of the most dominant Mediterranean tree species.

Growth duration is a better predictor of stem increment than carbon supply in a Mediterranean oak forest: implications for assessing forest productivity under climate change.

Understanding whether tree growth is limited by carbon gain (source limitation) or by the direct effect of environmental factors such as water deficit or temperature (sink limitation) is crucial for improving projections of the effects of climate change on forest productivity. We studied the relationships between tree basal area (BA) variations, eddy covariance carbon fluxes, predawn water potential (Ψpd ) and temperature at different timescales using an 8-yr dataset and a rainfall exclusion experiment in a Quercus ilex Mediterranean coppice. At the daily timescale, during periods of low temperature (< 5°C) and high water deficit (< -1.1 MPa), gross primary productivity and net ecosystem productivity remained positive whereas the stem increment was nil. Thus, stem increment appeared limited by drought and temperature rather than by carbon input. Annual growth was accurately predicted by the duration of BA increment during spring (Δtt0-t1 ). The onset of growth (t0 ) was related to winter temperatures and the summer interruption of growth (t1 ) to a threshold Ψpd value of -1.1 MPa. We suggest that using environmental drivers (i.e. drought and temperature) to predict stem growth phenology can contribute to an improvement in vegetation models and may change the current projections of Mediterranean forest productivity under climate change scenarios.

Ecological significance of seed desiccation sensitivity in Quercus ilex.

BACKGROUND AND AIMS: Several widespread tree species of temperate forests, such as species of the genus Quercus, produce recalcitrant (desiccation-sensitive) seeds. However, the ecological significance of seed desiccation sensitivity in temperate regions is largely unknown. Do seeds of such species suffer from drying during the period when they remain on the soil, between shedding in autumn and the return of conditions required for germination in spring? METHODS: To test this hypothesis, the Mediterranean holm oak (Quercus ilex) forest was used as a model system. The relationships between the climate in winter, the characteristics of microhabitats, acorn morphological traits, and the water status and viability of seeds after winter were then investigated in 42 woodlands sampled over the entire French distribution of the species. KEY RESULTS: The percentages of germination and normal seedling development were tightly linked to the water content of seeds after the winter period, revealing that in situ desiccation is a major cause of mortality. The homogeneity of seed response to drying suggests that neither intraspecific genetic variation nor environmental conditions had a significant impact on the level of desiccation sensitivity of seeds. In contrast, the water and viability status of seeds at the time of collection were dramatically influenced by cumulative rainfall and maximum temperatures during winter. A significant effect of shade and of the type of soil cover was also evidenced. CONCLUSIONS: The findings establish that seed desiccation sensitivity is a key functional trait which may influence the success of recruitment in temperate recalcitrant seed species. Considering that most models of climate change predict changes in rainfall and temperature in the Mediterranean basin, the present work could help foresee changes in the distribution of Q. ilex and other oak species, and hence plant community alterations.

Morphological and phenological shoot plasticity in a Mediterranean evergreen oak facing long-term increased drought.

Mediterranean trees must adjust their canopy leaf area to the unpredictable timing and severity of summer drought. The impact of increased drought on the canopy dynamics of the evergreen Quercus ilex was studied by measuring shoot growth, leaf production, litterfall, leafing phenology and leaf demography in a mature forest stand submitted to partial throughfall exclusion for 7 years. The leaf area index rapidly declined in the throughfall-exclusion plot and was 19% lower than in the control plot after 7 years of treatment. Consequently, leaf litterfall was significantly lower in the dry treatment. Such a decline in leaf area occurred through a change in branch allometry with a decreased number of ramifications produced and a reduction of the leaf area supported per unit sapwood area of the shoot (LA/SA). The leafing phenology was slightly delayed and the median leaf life span was slightly longer in the dry treatment. The canopy dynamics in both treatments were driven by water availability with a 1-year lag: leaf shedding and production were reduced following dry years; in contrast, leaf turnover was increased following wet years. The drought-induced decrease in leaf area, resulting from both plasticity in shoot development and slower leaf turnover, appeared to be a hydraulic adjustment to limit canopy transpiration and maintain leaf-specific hydraulic conductivity under drier conditions.

Ground-based optical measurements at European flux sites: a review of methods, instruments and current controversies.

This paper reviews the currently available optical sensors, their limitations and opportunities for deployment at Eddy Covariance (EC) sites in Europe. This review is based on the results obtained from an online survey designed and disseminated by the Co-cooperation in Science and Technology (COST) Action ESO903-"Spectral Sampling Tools for Vegetation Biophysical Parameters and Flux Measurements in Europe" that provided a complete view on spectral sampling activities carried out within the different research teams in European countries. The results have highlighted that a wide variety of optical sensors are in use at flux sites across Europe, and responses further demonstrated that users were not always fully aware of the key issues underpinning repeatability and the reproducibility of their spectral measurements. The key findings of this survey point towards the need for greater awareness of the need for standardisation and development of a common protocol of optical sampling at the European EC sites.

On the differential advantages of evergreenness and deciduousness in mediterranean oak woodlands: a flux perspective.

We assessed the differential advantages of deciduousness and evergreenness by examining 26 site-years of carbon dioxide, water vapor, and energy flux measurements from five comparable oak woodlands in France, Italy, Portugal, and California (USA). On average, the evergreen and deciduous oak woodlands assimilated and respired similar amounts of carbon while using similar amounts of water. These results suggest that evergreen and deciduous woodlands have specific, and similar, ecological costs in mediterranean climates, and that both leaf habits are able to meet these costs. What are the mechanisms behind these findings? Deciduous oaks compensated for having a shorter growing season by attaining a greater capacity to assimilate carbon for a given amount of intercepted solar radiation during the well-watered spring period; at saturating light levels, deciduous oaks gained carbon at six times the rate of evergreen oaks. Otherwise, the two leaf habits experienced similar efficiencies in carbon use (the change in carbon respired per change in carbon assimilated), water use (the change in carbon assimilation per change in water evaporated), and rainfall use (the change in evaporation per change in rainfall). Overall, leaf area index, rather than leaf habit, was the significant factor in determining the absolute magnitude of carbon gained and water lost by each evergreen and deciduous oak woodland over an annual interval; the closed canopies assimilated and respired more carbon and transpired more water than the open canopies. Both deciduous and evergreen mediterranean oaks survive in their seasonally hot/dry, wet/ cool native range by ensuring that actual evaporation is less than the supply of water. This feat is accomplished by adjusting the leaf area index to reduce total water loss at the landscape scale, by down-regulating photosynthesis, respiration, and stomatal conductance with progressive seasonal soil water deficits, and by extending their root systems to tap groundwater.