Differences in xylem and leaf hydraulic traits explain differences in drought tolerance among mature Amazon rainforest trees.

Considerable uncertainty surrounds the impacts of anthropogenic climate change on the composition and structure of Amazon forests. Building upon results from two large-scale ecosystem drought experiments in the eastern Brazilian Amazon that observed increases in mortality rates among some tree species but not others, in this study we investigate the physiological traits underpinning these differential demographic responses. Xylem pressure at 50% conductivity (xylem-P50 ), leaf turgor loss point (TLP), cellular osmotic potential (πo ), and cellular bulk modulus of elasticity (ε), all traits mechanistically linked to drought tolerance, were measured on upper canopy branches and leaves of mature trees from selected species growing at the two drought experiment sites. Each species was placed a priori into one of four plant functional type (PFT) categories: drought-tolerant versus drought-intolerant based on observed mortality rates, and subdivided into early- versus late-successional based on wood density. We tested the hypotheses that the measured traits would be significantly different between the four PFTs and that they would be spatially conserved across the two experimental sites. Xylem-P50 , TLP, and πo , but not ε, occurred at significantly higher water potentials for the drought-intolerant PFT compared to the drought-tolerant PFT; however, there were no significant differences between the early- and late-successional PFTs. These results suggest that these three traits are important for determining drought tolerance, and are largely independent of wood density-a trait commonly associated with successional status. Differences in these physiological traits that occurred between the drought-tolerant and drought-intolerant PFTs were conserved between the two research sites, even though they had different soil types and dry-season lengths. This more detailed understanding of how xylem and leaf hydraulic traits vary between co-occuring drought-tolerant and drought-intolerant tropical tree species promises to facilitate a much-needed improvement in the representation of plant hydraulics within terrestrial ecosystem and biosphere models, which will enhance our ability to make robust predictions of how future changes in climate will affect tropical forests.

Plasticity in leaf-level water relations of tropical rainforest trees in response to experimental drought.

The tropics are predicted to become warmer and drier, and understanding the sensitivity of tree species to drought is important for characterizing the risk to forests of climate change. This study makes use of a long-term drought experiment in the Amazon rainforest to evaluate the role of leaf-level water relations, leaf anatomy and their plasticity in response to drought in six tree genera. The variables (osmotic potential at full turgor, turgor loss point, capacitance, elastic modulus, relative water content and saturated water content) were compared between seasons and between plots (control and through-fall exclusion) enabling a comparison between short- and long-term plasticity in traits. Leaf anatomical traits were correlated with water relation parameters to determine whether water relations differed among tissues. The key findings were: osmotic adjustment occurred in response to the long-term drought treatment; species resistant to drought stress showed less osmotic adjustment than drought-sensitive species; and water relation traits were correlated with tissue properties, especially the thickness of the abaxial epidermis and the spongy mesophyll. These findings demonstrate that cell-level water relation traits can acclimate to long-term water stress, and highlight the limitations of extrapolating the results of short-term studies to temporal scales associated with climate change.

Confronting model predictions of carbon fluxes with measurements of Amazon forests subjected to experimental drought.

Considerable uncertainty surrounds the fate of Amazon rainforests in response to climate change. Here, carbon (C) flux predictions of five terrestrial biosphere models (Community Land Model version 3.5 (CLM3.5), Ecosystem Demography model version 2.1 (ED2), Integrated BIosphere Simulator version 2.6.4 (IBIS), Joint UK Land Environment Simulator version 2.1 (JULES) and Simple Biosphere model version 3 (SiB3)) and a hydrodynamic terrestrial ecosystem model (the Soil-Plant-Atmosphere (SPA) model) were evaluated against measurements from two large-scale Amazon drought experiments. Model predictions agreed with the observed C fluxes in the control plots of both experiments, but poorly replicated the responses to the drought treatments. Most notably, with the exception of ED2, the models predicted negligible reductions in aboveground biomass in response to the drought treatments, which was in contrast to an observed c. 20% reduction at both sites. For ED2, the timing of the decline in aboveground biomass was accurate, but the magnitude was too high for one site and too low for the other. Three key findings indicate critical areas for future research and model development. First, the models predicted declines in autotrophic respiration under prolonged drought in contrast to measured increases at one of the sites. Secondly, models lacking a phenological response to drought introduced bias in the sensitivity of canopy productivity and respiration to drought. Thirdly, the phenomenological water-stress functions used by the terrestrial biosphere models to represent the effects of soil moisture on stomatal conductance yielded unrealistic diurnal and seasonal responses to drought.

Effect of 7 yr of experimental drought on vegetation dynamics and biomass storage of an eastern Amazonian rainforest.

*At least one climate model predicts severe reductions of rainfall over Amazonia during this century. Long-term throughfall exclusion (TFE) experiments represent the best available means to investigate the resilience of the Amazon rainforest to such droughts. *Results are presented from a 7 yr TFE study at Caxiuanã National Forest, eastern Amazonia. We focus on the impacts of the drought on tree mortality, wood production and above-ground biomass. *Tree mortality in the TFE plot over the experimental period was 2.5% yr(-1), compared with 1.25% yr(-1) in a nearby control plot experiencing normal rainfall. Differences in stem mortality between plots were greatest in the largest (> 40 cm diameter at breast height (dbh)) size class (4.1% yr(-1) in the TFE and 1.4% yr(-1) in the control). Wood production in the TFE plot was c. 30% lower than in the control plot. Together, these changes resulted in a loss of 37.8 +/- 2.0 Mg carbon (C) ha(-1) in the TFE plot (2002-2008), compared with no change in the control. *These results are remarkably consistent with those from another TFE (at Tapajós National Forest), suggesting that eastern Amazonian forests may respond to prolonged drought in a predictable manner.

Limited acclimation in leaf anatomy to experimental drought in tropical rainforest trees.

Dry periods are predicted to become more frequent and severe in the future in some parts of the tropics, including Amazonia, potentially causing reduced productivity, higher tree mortality and increased emissions of stored carbon. Using a long-term (12 year) through-fall exclusion (TFE) experiment in the tropics, we test the hypothesis that trees produce leaves adapted to cope with higher levels of water stress, by examining the following leaf characteristics: area, thickness, leaf mass per area, vein density, stomatal density, the thickness of palisade mesophyll, spongy mesophyll and both of the epidermal layers, internal cavity volume and the average cell sizes of the palisade and spongy mesophyll. We also test whether differences in leaf anatomy are consistent with observed differential drought-induced mortality responses among taxa, and look for relationships between leaf anatomy, and leaf water relations and gas exchange parameters. Our data show that trees do not produce leaves that are more xeromorphic in response to 12 years of soil moisture deficit. However, the drought treatment did result in increases in the thickness of the adaxial epidermis (TFE: 20.5 ± 1.5 µm, control: 16.7 ± 1.0 µm) and the internal cavity volume (TFE: 2.43 ± 0.50 mm3 cm-2, control: 1.77 ± 0.30 mm3 cm-2). No consistent differences were detected between drought-resistant and drought-sensitive taxa, although interactions occurred between drought-sensitivity status and drought treatment for the palisade mesophyll thickness (P = 0.034) and the cavity volume of the leaves (P = 0.025). The limited response to water deficit probably reflects a tight co-ordination between leaf morphology, water relations and photosynthetic properties. This suggests that there is little plasticity in these aspects of plant anatomy in these taxa, and that phenotypic plasticity in leaf traits may not facilitate the acclimation of Amazonian trees to the predicted future reductions in dry season water availability.