Understanding the underlying mechanisms that account for the impact of potassium (K) fertilization and its replacement by sodium (Na) on tree growth is key to improving the management of forest plantations that are expanding over weathered tropical soils with low amounts of exchangeable bases. A complete randomized block design was planted with Eucalyptus grandis (W. Hill ex Maiden) to quantify growth, carbon uptake and carbon partitioning using a carbon budget approach. A combination of approaches including the establishment of allometric relationships over the whole rotation and measurements of soil CO(2) efflux and aboveground litterfall at the end of the rotation were used to estimate aboveground net production (ANPP), total belowground carbon flux and gross primary production (GPP). The stable carbon isotope (δ(13)C) of stem wood α-cellulose produced every year was used as a proxy for stomatal limitation of photosynthesis. Potassium fertilization increased GPP and decreased the fraction of carbon allocated belowground. Aboveground net production was strongly enhanced, and because leaf lifespan increased, leaf biomass was enhanced without any change in leaf production, and wood production (P(W)) was dramatically increased. Sodium application decreased the fraction of carbon allocated belowground in a similar way, and enhanced GPP, ANPP and P(W), but to a lesser extent compared with K fertilization. Neither K nor Na affected δ(13)C of stem wood α-cellulose, suggesting that water-use efficiency was the same among the treatments and that the inferred increase in leaf photosynthesis was not only related to a higher stomatal conductance. We concluded that the response to K fertilization and Na addition on P(W) resulted from drastic changes in carbon allocation.
Adaptive strategies to improve tree water-use efficiency (WUE) are required to meet the global demand for wood in a future drier climate. A large-scale throughfall exclusion experiment was set up in Brazil to study the interaction between water status and potassium (K) or sodium (Na) availability on the ecophysiology of Eucalyptus grandis trees. This experiment focused primarily on the changes in aboveground net primary production, stand water use, phloem sap and leaf d 13 C, net CO 2 assimilation and stomatal conductance. The correlations between these response variables were determined to gain insight into the factors controlling water-use efficiency in tropical eucalypt plantations. The intrinsic WUE in individual leaves (the ratio of net CO 2 assimilation to stomatal conductance) was estimated at a very short time scale from the leaf gas exchange. Sap flow measurements were carried out to assess the WUE for stemwood production (the ratio of wood biomass increment to stand water use). Averaged over the two water supply regimes, the stemwood biomass 3 years after planting was 173% higher in trees fertilized with K and 79% higher in trees fertilized with Na than in trees with no K and Na addition. Excluding 37% of the throughfall reduced stemwood production only for trees fertilized with K. Total canopy transpiration between 1 and 3 years after planting increased from about 750 to 1300 mm y À1 in response to K fertilization with a low influence of the water supply regime. K fertilization increased WUE for stemwood production by approx. 60% with or without throughfall exclusion. There was a strong positive correlation between phloem sap d 13 C and short-term leaf-level intrinsic WUE. Whatever the water and nutrient supply regime, the gas exchange WUE estimates were not correlated with WUE for stemwood production. Phloem sap d 13 C and leaf d 13 C were therefore not valuable proxies of WUE for stemwood production. The allocation pattern in response to nutrient and water supply appeared to be a major driver of WUE for stemwood production. In areas with very deep tropical soils and annual rainfall <1500 mm, our results suggest that breeding programs selecting the eucalypt clones with the highest growth rates tend to select the genotypes with the highest water-use efficiency for wood production.
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 supply regimes on water resources in tropical regions. This interaction is a major issue because climate change is expected to increase the length of drought periods in many tropical regions and soil water availability in deep soil layers is likely to have a major influence on tree growth during dry periods in tropical planted forests. A process-based model (MAESPA) was parameterized in a throughfall exclusion experiment in Brazil to gain insight into the combined effects of K deficiency and rainfall reduction (37% throughfall exclusion) on the water used by the trees, soil water storage and water table fluctuations over the first 4.5 years after planting Eucalyptus grandis trees. A comparison of canopy transpiration in each plot with the values predicted for the same soil with the water content maintained at field capacity, made it possible to calculate a soil-driven tree water stress index for each treatment. Compared to K-fertilized trees with undisturbed rainfall (+K+W), canopy transpiration was 40% lower for K deficiency (−K+W), 20% lower for W deficit (+K−W) and 36% lower for combined K deficiency and W deficit (−K−W) on average. Water was withdrawn in deeper soil layers for −W than for +W, particularly over dry seasons. Under contrasted K availability, water withdrawal was more superficial for −K than for +K. Mean soil water content down to 18 m below surface (mbs) was 24% higher for −K+W than for +K+W from 2 years after planting (after canopy closure), while it was 24% lower for +K−W and 12% lower for −K−W than for +K+W. The soil-driven tree water stress index was 166% higher over the first 4.5 years after planting for −W than for +W, 76% lower for −K than for +K, and 14% lower for −K−W than for +K+W. Over the study period, deep seepage was higher by 371 mm yr −1 (+122%) for −K than for +K and lower by 200 mm yr −1 (−66%) for −W than for +W. Deep seepage was lower by 44% for −K−W than for +K+W. At the end of the study period, the model predicted a higher water table for −K (10 mbs for −K+W and 16 mbs for −K−W) than for +K (16 mbs for +K+W and 18 mbs for +K−W). Our study suggests that flexible fertilization regimes could contribute to adjusting the local trade-off between wood production and demand for soil water resources in planted forests.
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