Yann Nouvellon

Influence of nitrogen and potassium fertilization on leaf lifespan and allocation of above-ground growth in Eucalyptus plantations

Eucalyptus grandis (W. Hill ex Maiden) leaf traits and tree growth were studied over 3 years after the establishment of two adjacent complete randomized block designs in southern Brazil. In a nitrogen (N) input experiment, a treatment with the application of 120 kg N ha(-1) was compared to a control treatment without N addition, and in a potassium (K) input experiment a control treatment without K addition was compared to a treatment with the application of 116 kg K ha(-1). Young leaves were tagged 9 months after planting to estimate the effect of N and K fertilizations on leaf lifespan. Leaf mass, specific leaf area and nutrient concentrations were measured on a composite sample per plot every 28 days until the last tagged leaf fell. Successive inventories, destructive sampling of trees and leaf litter fall collection made it possible to assess the effect of N and K fertilization on the dynamics of biomass accumulation in above-ground tree components. Whilst the effects of N fertilization on tree growth only occurred in the first 24 months after planting, K fertilization increased the above-ground net primary production from 4478 to 8737 g m(-2) over the first 36 months after planting. The average lifespan of tagged leaves was not modified by N addition but it increased from 111 to 149 days with K fertilization. The peak of leaf production occurred in the second year after planting (about 800 g m(-2) year(-1)) and was not significantly modified (P < 0.05) by N and K fertilizations. By contrast, K addition significantly increased the maximum leaf standing biomass from 292 to 528 g m(-2), mainly as a consequence of the increase in leaf lifespan. Potassium fertilization increased the stand biomass mainly through the enhancement in leaf area index (LAI) since growth efficiency (defined as the ratio between woody biomass production and LAI) was not significantly modified. A better understanding of the physiological processes governing the leaf lifespan is necessary to improve process-based models currently used in Eucalyptus plantations.

Almost symmetrical vertical growth rates above and below ground in one of the world's most productive forests

Whilst the relationships between growth strategies and leaf traits are well established in functional plant ecology, little attention has been paid to root traits in very deep soil layers. The objective of our study was to compare the vertical velocity of the above- and belowground exploration of the environment for one of the fastest-growing tree species. Fine roots were studied in a chronosequence of intensively-managed Eucalyptus plantations established on highly weathered soils. Here we show that the root front depth was accurately predicted at 85% of mean tree height for stands <20 m in height, in the absence of any physical or chemical barrier. Tree height and root front growth velocities peaked at 0.59 and 0.55 m month−1 respectively 9–10 months after planting, and decreased steadily thereafter. Fast root front displacement might provide a competitive advantage to fast-growing species in forests established on deep soils. Our study may contribute to the debate on the environmental impact of short-rotation plantation forests in the Tropics.

Time Course of Radiation Use Efficiency in a Shortgrass Ecosystem: Consequences for Remotely Sensed Estimation of Primary Production

Abstract A reliable estimation of primary production of terrestrial ecosystems is often a prerequisite for land survey and management, while being important also in ecological and climatological studies. At a regional scale, grassland primary production estimates are increasingly being made with the use of satellite data. In a currently used approach, regional gross, net, and aboveground net primary productivity (GPP, NPP, and ANPP) are derived from the parametric model of Monteith and are calculated as the product of the fraction of incident photosynthetically active radiation absorbed by the canopy (fAPAR) and gross, net, and aboveground net production (radiation-use) efficiencies (ϵg, ϵn, and ϵan); fAPAR being derived from indices calculated from satellite-measured reflectances in the red and near infrared. The accuracy and realism of the primary production values estimated by this approach therefore largely depend on an accurate estimation of ϵg, ϵn, and ϵan. However, data are scarce for production efficiencies of semiarid grasslands, and their time and spatial variations are poorly documented, often leading to large errors for the estimates. In this paper, a modeling approach taking into account relevant ecosystem processes and based on extensive field data was used to estimate time variations of ϵg, ϵn and ϵan of a shortgrass site in Arizona. These variations were explained by variations in plant water stress, temperature, leaf aging, and processes such as respiration and changes in allocation pattern between above- and below-ground compartments. Over the 3 study years, averaged values of ϵg, ϵn, and ϵan were found to be 1.92, 0.74, and 0.29 g DM (MJ IPAR)−1, respectively. ϵg and ϵn exhibited large interannual and seasonal variations mainly due to changes in water limitations during the growing season. Interannual variations of ϵan were much less important. However, for shorter periods, ϵan exhibited very contrasting values from regrowth to senescence. The calculation of ANPP seems less prone to errors due to environmental effects when computed on an annual basis. When estimating GPP and NPP, better results are expected if water limitations are taken into account. This could be possible through the estimation of a water-stress factor by using surface temperature or other indices derived from thermal infrared remote sensing data. The limitations due to temporally varying efficiencies, shown here for shortgrass ecosystems, are also relevant to all drought-exposed ecosystems, particularly those with abundant evergreen or perennial species.

Production and carbon allocation in monocultures and mixed-species plantations of Eucalyptus grandis and Acacia mangium in Brazil

Introducing nitrogen-fixing tree species in fast-growing eucalypt plantations has the potential to improve soil nitrogen availability compared with eucalypt monocultures. Whether or not the changes in soil nutrient status and stand structure will lead to mixtures that out-yield monocultures depends on the balance between positive interactions and the negative effects of interspecific competition, and on their effect on carbon (C) uptake and partitioning. We used a C budget approach to quantify growth, C uptake and C partitioning in monocultures of Eucalyptus grandis (W. Hill ex Maiden) and Acacia mangium (Willd.) (treatments E100 and A100, respectively), and in a mixture at the same stocking density with the two species at a proportion of 1 : 1 (treatment MS). Allometric relationships established over the whole rotation, and measurements of soil CO(2) efflux and aboveground litterfall for ages 4-6 years after planting were used to estimate aboveground net primary production (ANPP), total belowground carbon flux (TBCF) and gross primary production (GPP). We tested the hypotheses that (i) species differences for wood production between E. grandis and A. mangium monocultures were partly explained by different C partitioning strategies, and (ii) the observed lower wood production in the mixture compared with eucalypt monoculture was mostly explained by a lower partitioning aboveground. At the end of the rotation, total aboveground biomass was lowest in A100 (10.5 kg DM m(-2)), intermediate in MS (12.2 kg DM m(-2)) and highest in E100 (13.9 kg DM m(-2)). The results did not support our first hypothesis of contrasting C partitioning strategies between E. grandis and A. mangium monocultures: the 21% lower growth (ΔB(w)) in A100 compared with E100 was almost entirely explained by a 23% lower GPP, with little or no species difference in ratios such as TBCF/GPP, ANPP/TBCF, ΔB(w)/ANPP and ΔB(w)/GPP. In contrast, the 28% lower ΔB(w) in MS than in E100 was explained both by a 15% lower GPP and by a 15% lower fraction of GPP allocated to wood growth, thus partially supporting our second hypothesis: mixing the two species led to shifts in C allocations from above- to belowground, and from growth to litter production, for both species.

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.

Dynamics of soil exploration by fine roots down to a depth of 10 m throughout the entire rotation in Eucalyptus grandis plantations

Although highly weathered soils cover considerable areas in tropical regions, little is known about exploration by roots in deep soil layers. Intensively managed Eucalyptus plantations are simple forest ecosystems that can provide an insight into the belowground growth strategy of fast-growing tropical trees. Fast exploration of deep soil layers by eucalypt fine roots may contribute to achieving a gross primary production that is among the highest in the world for forests. Soil exploration by fine roots down to a depth of 10 m was studied throughout the complete cycle in Eucalyptus grandis plantations managed in short rotation. Intersects of fine roots, less than 1 mm in diameter, and medium-sized roots, 1–3 mm in diameter, were counted on trench walls in a chronosequence of 1-, 2-, 3.5-, and 6-year-old plantations on a sandy soil, as well as in an adjacent 6-year-old stand growing in a clayey soil. Two soil profiles were studied down to a depth of 10 m in each stand (down to 6 m at ages 1 and 2 years) and 4 soil profiles down to 1.5–3.0 m deep. The root intersects were counted on 224 m2 of trench walls in 15 pits. Monitoring the soil water content showed that, after clear-cutting, almost all the available water stored down to a depth of 7 m was taken up by tree roots within 1.1 year of planting. The soil space was explored intensively by fine roots down to a depth of 3 m from 1 year after planting, with an increase in anisotropy in the upper layers throughout the rotation. About 60% of fine root intersects were found at a depth of more than 1 m, irrespective of stand age. The root distribution was isotropic in deep soil layers and kriged maps showed fine root clumping. A considerable volume of soil was explored by fine roots in eucalypt plantations on deep tropical soils, which might prevent water and nutrient losses by deep drainage after canopy closure and contribute to maximizing resource uses.

Do changes in carbon allocation account for the growth response to potassium and sodium applications in tropical Eucalyptus plantations

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.

BAAD: a biomass and allometry database for woody plants

Understanding how plants are constructed—i.e., how key size dimensions and the amount of mass invested in different tissues varies among individuals—is essential for modeling plant growth, carbon stocks, and energy fluxes in the terrestrial biosphere. Allocation patterns can differ through ontogeny, but also among coexisting species and among species adapted to different environments. While a variety of models dealing with biomass allocation exist, we lack a synthetic understanding of the underlying processes. This is partly due to the lack of suitable data sets for validating and parameterizing models. To that end, we present the Biomass And Allometry Database (BAAD) for woody plants. The BAAD contains 259 634 measurements collected in 176 different studies, from 21 084 individuals across 678 species. Most of these data come from existing publications. However, raw data were rarely made public at the time of publication. Thus, the BAAD contains data from different studies, transformed into standard units and variable names. The transformations were achieved using a common workflow for all raw data files. Other features that distinguish the BAAD are: (i) measurements were for individual plants rather than stand averages; (ii) individuals spanning a range of sizes were measured; (iii) plants from 0.01–100 m in height were included; and (iv) biomass was estimated directly, i.e., not indirectly via allometric equations (except in very large trees where biomass was estimated from detailed sub-sampling). We included both wild and artificially grown plants. The data set contains the following size metrics: total leaf area; area of stem cross-section including sapwood, heartwood, and bark; height of plant and crown base, crown area, and surface area; and the dry mass of leaf, stem, branches, sapwood, heartwood, bark, coarse roots, and fine root tissues. We also report other properties of individuals (age, leaf size, leaf mass per area, wood density, nitrogen content of leaves and wood), as well as information about the growing environment (location, light, experimental treatment, vegetation type) where available. It is our hope that making these data available will improve our ability to understand plant growth, ecosystem dynamics, and carbon cycling in the worlds vegetation.

Testing Different Methods of Forest Height and Aboveground Biomass Estimations From ICESat/GLAS Data in Eucalyptus Plantations in Brazil

The Geoscience Laser Altimeter System (GLAS) has provided a useful dataset for estimating forest heights in many areas of the globe. Most of the studies on GLAS waveforms have focused on natural forests and only a few were conducted over forest plantations. This work set out to estimate the stand-scale dominant height and aboveground biomass of intensively managed Eucalyptus plantations in Brazil using the most commonly used models developed for natural forests. These forest plantations are valuable case studies, with large and numerous stands that are very uniform, in which field measurements are precise compared to natural forests. The height of planted Eucalyptus forest stands estimated from waveforms acquired by GLAS were compared with in situ measurements in order to determine the model that produced the best forest height estimates. For our slightly sloping study site , the direct method defined as the difference between the signal begin and the ground peak provided forest height estimates with an accuracy of 2.2 m. The use of statistical models based on waveform metrics and digital elevation models provided slightly better results (1.89 m accuracy) in comparison with the direct method and the most relevant metrics proved to be the trailing edge extent and the waveform extent. Moreover, a power law model was used to fit in situ aboveground biomass to in situ forest height. The results using this model with GLAS-derived heights showed an accuracy for biomass of 16.1 Mg/ha.

Root elongation in tropical Eucalyptus plantations: effect of soil water content

Abstract• Sustainability of Eucalyptus plantations is often questioned in resource-limited environments, especially in areas characterized by soils with poor nutrient and water holding capacities. Yet, field-based observations of fine root dynamics in relation with the seasonality of rainfall are lacking.• This study was undertaken on two Eucalyptus stands planted in the Kouilou Region (south-western Congo), which is characterized by a four-month-long dry season. Fine root (less than 2 mm in diameter) dynamics were studied using rhizotron observations of root elongation in the field.• Fine root elongation rates displayed a seasonal variation in the two stands, with higher elongation rates during the rainy season than during the dry season. Positive and significant correlations were found between fine root elongation rates and soil water content at all depths, but a better correlation was found with soil water content in the deep soil horizon than in the superficial horizons.• These results suggest that the temporal variations in fine root elongation were related to the seasonality of rainfall, and they were probably associated with seasonal changes in tree water status, carbon assimilation and belowground allocation.Résumé• La question de la durabilité des plantations d’Eucalyptus est souvent posée dans les environnements où les ressources sont limitées, en particulier les zones où les sols ont une faible capacité à retenir l’eau et les nutriments et où la saison sèche est longue. Pourtant, les observations in situ de la dynamique racinaire en relation avec la saisonnalité des pluies sont inexistantes.• Cette étude a été réalisée dans deux plantations d’Eucalyptus de la région du Kouilou dans le sud ouest du Congo, qui est caractérisée par quatre mois de saison sèche. La dynamique des racines fines (moins de 2 mm de diamètre) a été étudiée à l’aide de rhizotrons permettant d’observer l’élongation racinaire au champ.• La vitesse d’élongation des racines fines montrait une variation saisonnière dans les deux plantations, avec des vitesses plus élevées en saison des pluies qu’en saison sèche. Des corrélations positives et significatives ont été trouvées entre la vitesse d’élongation des racines fines et la teneur en eau du sol à toutes les profondeurs, mais les meilleures corrélations ont été observées avec la teneur en eau des horizons profonds.• Cela suggère que les variations temporelles de l’élongation des racines fines sont reliées à la saisonnalité des précipitations, et qu’elles sont associées aux changements saisonniers d’état hydrique des arbres, d’assimilation carbonée et d’allocation vers les parties souterraines.