Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.

Climate modelling studies predict that the rain forests of the Eastern Amazon basin are likely to experience reductions in rainfall of up to 50% over the next 50-100 years. Efforts to predict the effects of changing climate, especially drought stress, on forest gas exchange are currently limited by uncertainty about the mechanism that controls stomatal closure in response to low soil moisture. At a through-fall exclusion experiment in Eastern Amazonia where water was experimentally excluded from the soil, we tested the hypothesis that plants are isohydric, that is, when water is scarce, the stomata act to prevent leaf water potential from dropping below a critical threshold level. We made diurnal measurements of leaf water potential ( Ψ Ψ Ψ Ψ l ), stomatal conductance ( g s ), sap flow and stem water potential ( Ψ Ψ Ψ Ψ stem ) in the wet and dry seasons. We compared the data with the predictions of the soil-plant-atmosphere (SPA) model, which embeds the isohydric hypothesis within its stomatal conductance algorithm. The model inputs for meteorology, leaf area index (LAI), soil water potential and soil-to-leaf hydraulic resistance ( R ) were altered between seasons in accordance with measured values. No optimization parameters were used to adjust the model. This 'mechanistic' model of stomatal function was able to explain the individual tree-level seasonal changes in water relations ( r 2 = = = = 0.85, 0.90 and 0.58 for Ψ Ψ Ψ Ψ l , sap flow and g s , respectively). The model indicated that the measured increase in R was the dominant cause of restricted water use during the dry season, resulting in a modelled restriction of sap flow four times greater than that caused by reduced soil water potential. Higher resistance during the dry season resulted from an increase in below-ground resistance (including root and soil-to-root resistance) to water flow.

The static chamber method (non-flow-through-non-steady-state chambers) is the most common method to measure fluxes of methane (CH4) from soils. Laboratory comparisons to quantify errors resulting from chamber design, operation and flux calculation methods are rare. We tested fifteen chambers against four flux levels (FL) ranging from 200 to 2300 mu g CH4 M-2 II-1. The measurements were conducted on a calibration tank using three quartz sand types with soil porosities of 53% (dry fine sand, S1), 47% (dry coarse sand, S2), and 33% (wetted fine sand, S3). The chambers tested ranged from 0.06 to 1.8 m in height, and 0.02 to 0.195 m(3) in volume, 7 of them were equipped with a fan, and 1 with a vent-tube. We applied linear and exponential flux calculation methods to the chamber data and compared these chamber fluxes to the reference fluxes from the calibration tank. The chambers underestimated the reference fluxes by on average 33% by the linear flux calculation method (R-Iin), whereas the chamber fluxes calculated by the exponential flux calculation method (R-exp) did not significantly differ from the reference fluxes (p <0.05). The flux under- or overestimations were chamber specific and independent of flux level. Increasing chamber height, area and volume significantly reduced the flux underestimation (p <0.05). Also, the use of non-linear flux calculation method significantly improved the flux estimation; however, simultaneously the uncertainty in the fluxes was increased. We provide correction factors, which can be used to correct the under- or overestimation of the fluxes by the chambers in the experiment. (C) 2012 Elsevier B.V. All rights reserved

Summary• The effects of drought on the Amazon rainforest are potentially large but remain poorly understood. Here, carbon (C) cycling after 5 yr of a large-scale through-fall exclusion (TFE) experiment excluding about 50% of incident rainfall from an eastern Amazon rainforest was compared with a nearby control plot.• Principal C stocks and fluxes were intensively measured in 2005. Additional minor components were either quantified in later site measurements or derived from the available literature.• Total ecosystem respiration (R eco ) and total plant C expenditure (PCE, the sum of net primary productivity (NPP) and autotrophic respiration (R auto )), were elevated on the TFE plot relative to the control. The increase in PCE and R eco was mainly caused by a rise in R auto from foliage and roots. Heterotrophic respiration did not differ substantially between plots. NPP was 2.4 ± 1.4 t C ha )1 yr )1 lower on the TFE than the control. Ecosystem carbon use efficiency, the proportion of PCE invested in NPP, was lower in the TFE plot (0.24 ± 0.04) than in the control (0.32 ± 0.04).• Drought caused by the TFE treatment appeared to drive fundamental shifts in ecosystem C cycling with potentially important consequences for long-term forest C storage.

Determining climate change feedbacks from tropical rainforests requires an understanding of how carbon gain through photosynthesis and loss through respiration will be altered. One of the key changes that tropical rainforests may experience under future climate change scenarios is reduced soil moisture availability. In this study we examine if and how both leaf photosynthesis and leaf dark respiration acclimate following more than 12 years of experimental soil moisture deficit, via a through‐fall exclusion experiment (TFE) in an eastern Amazonian rainforest. We find that experimentally drought‐stressed trees and taxa maintain the same maximum leaf photosynthetic capacity as trees in corresponding control forest, independent of their susceptibility to drought‐induced mortality. We hypothesize that photosynthetic capacity is maintained across all treatments and taxa to take advantage of short‐lived periods of high moisture availability, when stomatal conductance (g s) and photosynthesis can increase rapidly, potentially compensating for reduced assimilate supply at other times. Average leaf dark respiration (R d) was elevated in the TFE‐treated forest trees relative to the control by 28.2 ± 2.8% (mean ± one standard error). This mean R d value was dominated by a 48.5 ± 3.6% increase in the R d of drought‐sensitive taxa, and likely reflects the need for additional metabolic support required for stress‐related repair, and hydraulic or osmotic maintenance processes. Following soil moisture deficit that is maintained for several years, our data suggest that changes in respiration drive greater shifts in the canopy carbon balance, than changes in photosynthetic capacity.

The aim of this study was to identify the sources and depth of water uptake by 15-years old Quercus suber L. trees in southern Portugal under a Mediterranean climate, measuring d 18 O and dD in the soil-plantatmosphere continuum. Evidence for hydraulic lift was substantiated by the daily fluctuations observed in Y s at 0.4 and 1 m depth and supported by similar d 18 O values found in tree xylem sap, soil water in the rhizosphere and groundwater. From 0.25 m down to a depth of 1 m, dD trends differed according to vegetation type, showing a more depleted value in soil water collected under the evergreen trees ()47&) than under dead grasses ()35&). The hypothesis of a fractionation process occurring in the soil due to diffusion of water vapour in the dry soil is proposed to explain the more depleted soil dD signature observed under trees. Hydraulically lifted water was estimated to account for 17-81% of the water used during the following day by tree transpiration at the peak of the drought season, i.e., 0.

Summary1. The Amazon region may experience increasing moisture limitation over this century. Leaf dark respiration (R) is a key component of the Amazon rain forest carbon (C) cycle, but relatively little is known about its sensitivity to drought. 2. Here, we present measurements of R standardized to 25°C and leaf morphology from different canopy heights over 5 years at a rain forest subject to a large-scale through-fall reduction (TFR) experiment, and nearby, unmodified Control forest, at the Caxiuana˜reserve in the eastern Amazon. 3. In all five post-treatment measurement campaigns, mean R at 25°C was elevated in the TFR forest compared to the Control forest experiencing normal rainfall. After 5 years of the TFR treatment, R per unit leaf area and mass had increased by 65% and 42%, respectively, relative to pre-treatment means. In contrast, leaf area index (L) in the TFR forest was consistently lower than the Control, falling by 23% compared to the pre-treatment mean, largely because of a decline in specific leaf area (S). 4. The consistent and significant effects of the TFR treatment on R, L and S suggest that severe drought events in the Amazon, of the kind that may occur more frequently in future, could cause a substantial increase in canopy carbon dioxide emissions from this ecosystem to the atmosphere.

Drought is a major environmental constrain affecting plant performance and survival, particularly in Mediterranean ecosystems. Terpenoids may play a protective role under these conditions, however, observations of drought effects on plant terpenoid emissions are controversial ranging from decreased emissions to unaffected or increased release of terpenoids. In the present study we investigated terpenoid emissions of cork oak (Quercus suber) and gum rockrose (Cistus ladanifer) in response to summer drought stress in 2017. Pre-dawn leaf water potential (ΨPD) decreased from -0.64 to -1.72 MPa in Q. suber and from -1.69 to -4.05 MPa in C. ladanifer, indicating a transition from mild to severe drought along summer. Total terpenoid emissions decreased with drought, but differed significantly between species (p < 0.001) and in response to ΨPD, air temperature and assimilation rates. C. ladanifer emitted a large variety of >75 compounds comprising monoterpenes, sesquiterpenes and even diterpenes, which strongly decreased from 1.37 ± 0.23 μg g-1h-1 to 0.40 ± 0.08 μg g-1h-1 (p < 0.001) in response to drought. Total emission rates were positively correlated to air temperature (p < 0.001). C. ladanifer behavior points toward terpenoid leaf storage depletion and reduced substrate availability for terpenoid synthesis with increasing drought, most likely accelerated by high air temperatures. Q. suber emitted mainly monoterpenes and emissions declined significantly from June (0.50 ± 0.08 μg g-1h-1) to August (0.29 ± 0.02 μg g-1h-1) (p < 0.01). Emission rates were weakly correlated with net assimilation rates (R2 = 0.19, p < 0.001), but did not respond strongly to ΨPD and air temperature. Early onset of drought in 2017 most likely reduced plant metabolism in Q. suber, resulting in diminished, but stable terpenoid fluxes. Calculation of standard emission factors (at 30°C) revealed contrasting emission patterns of decreasing, unaffected, or increasing fluxes of single terpenoid compounds. Unaffected or drought-enhanced emissions of compounds such as α-pinene, camphene or manoyl oxide may point toward a specific role of these terpenoids in abiotic stress adaptation. In conclusion, these results suggest a strong negative, but species- and compound-specific effect of severe drought on terpenoid fluxes in Mediterranean ecosystems.