Increased hydraulic constraints in Eucalyptus plantations fertilized with potassium.

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 (As :Al ) 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 Al in K-supplied plots increased the hydraulic constraints on water use. This involved a direct mechanism through halved As :Al 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.

In situ 13CO2 pulse labelling of field-grown eucalypt trees revealed the effects of potassium nutrition and throughfall exclusion on phloem transport of photosynthetic carbon.

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.

Measured and modeled interactive effects of potassium deficiency and water deficit on gross primary productivity and light-use efficiency in Eucalyptus grandis plantations.

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.

Effects of potassium and sodium supply on drought-adaptive mechanisms in Eucalyptus grandis plantations.

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.

Photosynthetic and anatomical responses of Eucalyptus grandis leaves to potassium and sodium supply in a field experiment.

Although vast areas in tropical regions have weathered soils with low potassium (K) levels, little is known about the effects of K supply on the photosynthetic physiology of trees. This study assessed the effects of K and sodium (Na) supply on the diffusional and biochemical limitations to photosynthesis in Eucalyptus grandis leaves. A field experiment comparing treatments receiving K (+K) or Na (+Na) with a control treatment (C) was set up in a K-deficient soil. The net CO2 assimilation rates were twice as high in +K and 1.6 times higher in +Na than in the C as a result of lower stomatal and mesophyll resistance to CO2 diffusion and higher photosynthetic capacity. The starch content was higher and soluble sugar was lower in +K than in C and +Na, suggesting that K starvation disturbed carbon storage and transport. The specific leaf area, leaf thickness, parenchyma thickness, stomatal size and intercellular air spaces increased in +K and +Na compared to C. Nitrogen and chlorophyll concentrations were also higher in +K and +Na than in C. These results suggest a strong relationship between the K and Na supply to E. grandis trees and the functional and structural limitations to CO2 assimilation rates.