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Fertilization is commonly used to increase growth in forest plantations, but it may also affect tree water relations and responses to drought. Here, we measured changes in biomass, transpiration, sapwood-to-leaf area ratio (A s :A l ) and sap flow driving force (ΔΨ) during the 6-year rotation of tropical plantations of Eucalyptus grandis under controlled conditions for throughfall and potassium (K) fertilization. K fertilization increased final tree height by 8 m. Throughfall exclusion scarcely affected tree functioning because of deep soil water uptake. Tree growth increased in K-supplied plots and remained stable in K-depleted plots as tree height increased, while growth per unit leaf area increased in all plots. Stand transpiration and hydraulic conductance standardized per leaf area increased with height in K-depleted plots, but remained stable or decreased in K-supplied plots. Greater A l in K-supplied plots increased the hydraulic constraints on water use. This involved a direct mechanism through halved A s :A l in K-supplied plots relative to K-depleted plots, and an indirect mechanism through deteriorated water status in K-supplied plots, which prevented the increase in ΔΨ with tree height. K fertilization in tropical plantations reduces the hydraulic compensation to growth, which could increase the risk of drought-induced dieback under climate change., (© 2021 John Wiley & Sons Ltd.)

Harvest residue management is a key issue for the sustainability of Eucalyptus plantations established on poor soils. Soil microbial communities contribute to soil fertility by the decomposition of the organic matter (OM), but little is known about the effect of whole-tree harvesting (WTH) in comparison to stem only harvesting (SOH) on soil microbial functional diversity in Eucalyptus plantations. We studied the effects of harvest residue management (branches, leaves, bark) of Eucalyptus grandis trees on soil enzymatic activities and community-level physiological profiles in a Brazilian plantation. We measured soil microbial enzymatic activities involved in OM decomposition and we compared the community level physiological profiles (CLPP) of the soil microbes in WTH and SOH plots. WTH decreased enzyme activities and catabolic potential of the soil microbial community. Furthermore, these negative effects on soil functional diversity were mainly observed below the 0-5 cm layer (5-10 and 10-20 cm), suggesting that WTH can be harmful to the soil health in these plantations.

Wood production in fast-growing Eucalyptus grandis trees is highly dependent on both potassium (K) fertilization and water availability but the molecular processes underlying wood formation in response to the combined effects of these two limiting factors remain unknown. E. grandis trees were submitted to four combinations of K-fertilization and water supply. Weighted gene co-expression network analysis and MixOmics-based co-regulation networks were used to integrate xylem transcriptome, metabolome and complex wood traits. Functional characterization of a candidate gene was performed in transgenic E. grandis hairy roots. This integrated network-based approach enabled us to identify meaningful biological processes and regulators impacted by K-fertilization and/or water limitation. It revealed that modules of co-regulated genes and metabolites strongly correlated to wood complex traits are in the heart of a complex trade-off between biomass production and stress responses. Nested in these modules, potential new cell-wall regulators were identified, as further confirmed by the functional characterization of EgMYB137. These findings provide new insights into the regulatory mechanisms of wood formation under stressful conditions, pointing out both known and new regulators co-opted by K-fertilization and/or water limitation that may potentially promote adaptive wood traits., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

While potassium fertilization increases growth yield in Brazilian eucalyptus plantations, it could also increase water requirements, making trees more vulnerable to drought. Sodium fertilization, which has been shown to promote eucalyptus growth compared to K-deficient trees, could partially mitigate this adverse effect of potassium. However, little is known about the influence of K and Na fertilization on the tree metabolic response to water deficit. The aim of the present study was thus to analyze the transcriptome of leaves sampled from Eucalyptus grandis trees subjected to 37% rainfall reduction, and fertilized with potassium (K), sodium (Na), compared to control trees (C). The multifactorial experiment was set up in a field with a throughfall exclusion system. Transcriptomic analysis was performed on leaves from two-year-old trees, and data analyzed using multifactorial statistical analysis and weighted gene co-expression network analysis (WGCNA). Significant sets of genes were seen to respond to rainfall reduction, in interaction with K or Na fertilization, or to fertilization only (regardless of the water supply regime). The genes were involved in stress signaling, primary and secondary metabolism, secondary cell wall formation and photosynthetic activity. Our focus on key genes related to cation transporters and aquaporins highlighted specific regulation of ion homeostasis, and plant adjustment to water deficit. While water availability significantly affects the transcriptomic response of eucalyptus species, this study points out that the transcriptomic response is highly dependent on the fertilization regime. Our study is based on the first large-scale field trial in a tropical region, specifically designed to study the interaction between water availability and nutrition in eucalyptus. To our knowledge, this is the first global transcriptomic analysis to compare the influence of K and Na fertilization on tree adaptive traits in water deficit conditions., Competing Interests: The authors have declared that no competing interests exist.

Potassium (K) is an important limiting factor of tree growth, but little is known of the effects of K supply on the long-distance transport of photosynthetic carbon (C) in the phloem and of the interaction between K fertilization and drought. We pulse-labelled 2-year-old Eucalyptus grandis L. trees grown in a field trial combining K fertilization (+K and -K) and throughfall exclusion (+W and -W), and we estimated the velocity of C transfer by comparing time lags between the uptake of (13)CO2 and its recovery in trunk CO2 efflux recorded at different heights. We also analysed the dynamics of the labelled photosynthates recovered in the foliage and in the phloem sap (inner bark extract). The mean residence time of labelled C in the foliage was short (21-31 h). The time series of (13)C in excess in the foliage was affected by the level of fertilization, whereas the effect of throughfall exclusion was not significant. The velocity of C transfer in the trunk (0.20-0.82 m h(-1)) was twice as high in +K trees than in -K trees, with no significant effect of throughfall exclusion except for one +K -W tree labelled in the middle of the drought season that was exposed to a more pronounced water stress (midday leaf water potential of -2.2 MPa). Our results suggest that besides reductions in photosynthetic C supply and in C demand by sink organs, the lower velocity under K deficiency is due to a lower cross-sectional area of the sieve tubes, whereas an increase in phloem sap viscosity is more likely limiting phloem transport under drought. In all treatments, 10 times less (13)C was recovered in inner bark extracts at the bottom of the trunk when compared with the base of the crown, suggesting that a large part of the labelled assimilates has been exported out of the phloem and replaced by unlabelled C. This supports the 'leakage-retrieval mechanism' that may play a role in maintaining the pressure gradient between source and sink organs required to sustain high velocity of phloem transport in tall trees., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Global climate change is expected to increase the length of drought periods in many tropical regions. Although large amounts of potassium (K) are applied in tropical crops and planted forests, little is known about the interaction between K nutrition and water deficit on the physiological mechanisms governing plant growth. A process-based model (MAESPA) parameterized in a split-plot experiment in Brazil was used to gain insight into the combined effects of K deficiency and water deficit on absorbed radiation (aPAR), gross primary productivity (GPP), and light-use efficiency for carbon assimilation and stem biomass production (LUEC and LUEs ) in Eucalyptus grandis plantations. The main-plot factor was the water supply (undisturbed rainfall vs. 37% of throughfall excluded) and the subplot factor was the K supply (with or without 0.45 mol K m(-2 ) K addition). Mean GPP was 28% lower without K addition over the first 3 years after planting whether throughfall was partly excluded or not. K deficiency reduced aPAR by 20% and LUEC by 10% over the whole period of growth. With K addition, throughfall exclusion decreased GPP by 25%, resulting from a 21% decrease in LUEC at the end of the study period. The effect of the combination of K deficiency and water deficit was less severe than the sum of the effects of K deficiency and water deficit individually, leading to a reduction in stem biomass production, gross primary productivity and LUE similar to K deficiency on its own. The modeling approach showed that K nutrition and water deficit influenced absorbed radiation essentially through changes in leaf area index and tree height. The changes in gross primary productivity and light-use efficiency were, however, driven by a more complex set of tree parameters, especially those controlling water uptake by roots and leaf photosynthetic capacities., (© 2014 John Wiley & Sons Ltd.)

A basic understanding of nutrition effects on the mechanisms involved in tree response to drought is essential under a future drier climate. A large-scale throughfall exclusion experiment was set up in Brazil to gain an insight into the effects of potassium (K) and sodium (Na) nutrition on tree structural and physiological adjustments to water deficit. Regardless of the water supply, K and Na supply greatly increased growth and leaf area index (LAI) of Eucalyptus grandis trees over the first 3 yr after planting. Excluding 37% of throughfall reduced above-ground biomass accumulation in the third year after planting for K- supplied trees only. E. grandis trees were scarcely sensitive to drought as a result of the utilization of water stored in deep soil layers after clear-cutting the previous plantation. Trees coped with water restriction through stomatal closure (isohydrodynamic behavior), osmotic adjustment and decrease in LAI. Additionally, droughted trees showed higher phloem sap sugar concentrations. K and Na supply increased maximum stomatal conductance, and the high water requirements of fertilized trees increased water stress during dry periods. Fertilization regimes should be revisited in a future drier climate in order to find the right balance between improving tree growth and limiting water shortage., (© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.)

The consequences of diversity on belowground processes are still poorly known in tropical forests. The distributions of very fine roots (diameter <1 mm) and fine roots (diameter <3 mm) were studied in a randomized block design close to the harvest age of fast-growing plantations. A replacement series was set up in Brazil with mono-specific Eucalyptus grandis (100E) and Acacia mangium (100A) stands and a mixture with the same stocking density and 50% of each species (50A:50E). The total fine root (FR) biomass down to a depth of 2 m was about 27% higher in 50A:50E than in 100A and 100E. Fine root over-yielding in 50A:50E resulted from a 72 % rise in E. grandis fine root biomass per tree relative to 100E, whereas A. mangium FR biomass per tree was 17% lower than in 100A. Mixing A. mangium with E. grandis trees led to a drop in A. mangium FR biomass in the upper 50 cm of soil relative to 100A, partially balanced by a rise in deep soil layers. Our results highlight similarities in the effects of directional resources on leaf and FR distributions in the mixture, with A. mangium leaves below the E. grandis canopy and a low density of A. mangium fine roots in the resource-rich soil layers relative to monospecific stands. The vertical segregation of resource-absorbing organs did not lead to niche complementarity expected to increase the total biomass production.