Crown architecture affects tree growth through the control of leaf area and its display. Yet the linkages between crown structure, leaf traits, and productivity of elite selections of forest trees and responses to intensive silviculture are not fully understood. It was hypothesized that trees with crown and leaf traits governing efficient light capture and photosynthesis at the canopy scale would be the most productive. To this end, families of loblolly (Pinus taeda) and slash pine (Pinus elliottii) were grown at three experimental sites in the West Gulf Coastal Plain of Texas and Louisiana under two silvicultural treatments, including repeated fertilization with control of competing vegetation (HI), and a control (C) consisting of fertilization at planting. Families and species differed in crown traits and aboveground productivity, and genotype differences increased throughout the first 5 years of stand development. Crown shape was important for light interception and growth initially, but at the onset of canopy closure, crown size, stand leaf area and its distribution within crowns affected canopy light interception and tree growth. Among all families and treatments, aboveground biomass productivity was positively related to absorbed photosynthetically active radiation (APAR) and canopy photosynthesis. Light-use efficiency (ε) varied from 0.41 to 0.56 g MJ-1 among families and was lowest in slash pine. Variability in aboveground biomass growth was related more to stand leaf area and APAR than to differences in light-use efficiency in these young stands. Leaf physiological, chemical and morphological attributes changed within crowns in accordance with developing light availability gradients. Physiological attributes, such as net photosynthesis, were better predictors of family performance when integrated at the canopy level than leaf level in the examined pine species. Crown size, light absorption, and aboveground growth generally ranked higher in the HI treatment than in the control, although the effects of the intensive silvicultural treatments did not differ statistically. Family performance was independent of treatment. Crown and canopy attributes, such as high leaf area index and large crowns with low leaf area density per crown volume, may be useful in the selection of highly productive genotypes of loblolly and slash pine under intensive silviculture.

Crown architecture affects tree growth through the control of leaf area and its display. Yet the linkages between crown structure, leaf traits, and productivity of elite selections of forest trees and responses to intensive silviculture are not fully understood. It was hypothesized that trees with crown and leaf traits governing efficient light capture and photosynthesis at the canopy scale would be the most productive. To this end, families of loblolly (Pinus taeda) and slash pine (Pinus elliottii) were grown at three experimental sites in the West Gulf Coastal Plain of Texas and Louisiana under two silvicultural treatments, including repeated fertilization with control of competing vegetation (HI), and a control (C) consisting of fertilization at planting. Families and species differed in crown traits and aboveground productivity, and genotype differences increased throughout the first 5 years of stand development. Crown shape was important for light interception and growth initially, but at the onset of canopy closure, crown size, stand leaf area and its distribution within crowns affected canopy light interception and tree growth. Among all families and treatments, aboveground biomass productivity was positively related to absorbed photosynthetically active radiation (APAR) and canopy photosynthesis. Light-use efficiency (ε) varied from 0.41 to 0.56 g MJ-1 among families and was lowest in slash pine. Variability in aboveground biomass growth was related more to stand leaf area and APAR than to differences in light-use efficiency in these young stands. Leaf physiological, chemical and morphological attributes changed within crowns in accordance with developing light availability gradients. Physiological attributes, such as net photosynthesis, were better predictors of family performance when integrated at the canopy level than leaf level in the examined pine species. Crown size, light absorption, and aboveground growth generally ranked higher in the HI treatment than in the control, although the effects of the intensive silvicultural treatments did not differ statistically. Family performance was independent of treatment. Crown and canopy attributes, such as high leaf area index and large crowns with low leaf area density per crown volume, may be useful in the selection of highly productive genotypes of loblolly and slash pine under intensive silviculture.

This study investigated the use of optical remote sensing for estimating leaf and canopy scale light use efficiency (LUE) and carbon exchange. In addition, a new leaf level model capable of predicting dynamic changes in apparent reflectance due to chlorophyll fluorescence was developed. A leaf level study was conducted to assess the applicability of passive remote sensing as a tool to measure the reduction, and the subsequent recovery, of photosynthetic efficiency during the weeks following transplantation. Spectral data were collected on newly planted saplings for a period of 8 weeks, as well as gas exchange measurements of LUE and PAM fluorescence measurements. A set of spectral indices, including the Photochemical Reflectance Index (PRI), were calculated from the reflectance measurements. A marked depression in photosynthetic rate occurred in the weeks after outplanting followed by a gradual increase, with recovery occurring in the later stages of the experimental period. As with photosynthetic rate, there was a marked trend in PRI values over the study period but no trend was observed in chlorophyll based indices. The study demonstrated that hyperspectral remote sensing has the potential to be a useful tool in the detection and monitoring of the dynamic effects of transplant shock. Relationships between hyperspectral reflectance indices, airborne carbon exchange measurements and satellite observations of ground cover were then explored across a heterogeneous Arctic landscape. Measurements were collected during August 2008, using the University of Edinburgh’s research aircraft, from an Arctic forest tundra zone in northern Finland as part of the Arctic Biosphere Atmosphere Coupling at Multiple Scales (ABACUS) study. Surface fluxes of CO2 were calculated using the eddy covariance method from airborne data that were collected from the same platform as hyperspectral reflectance measurements. Airborne CO2 fluxes were compared to MODIS vegetation indices. In addition, LUE was estimated from airborne flux data and compared to airborne measurements of PRI. There were no significant relationships between MODIS vegetation indices and airborne flux observations. There were weak to moderate (R2 = 0.4 in both cases) correlations between PRI and LUE and between PRI and incident radiation. A new coupled physiological radiative transfer model that predicts changes in the apparent reflectance of a leaf, due to chlorophyll fluorescence, was developed. The model relates a physically observable quantity, chlorophyll fluorescence, to the sub leaf level processes that cause the emission. An understanding of the dynamics of the processes that control fluorescence emission on multiple timescales should aid in the interpretation of this complex signal. A Markov Chain Monte Carlo (MCMC) algorithm was used to optimise biochemical model parameters by fitting model simulations of transient chlorophyll fluorescence to measured reflectance spectra. The model was then validated against an independent data set. The model was developed as a precursor to a full canopy scheme. To scale to the canopy and to use the model on trans-seasonal time scales, the effects of temperature and photoinhibition on the model biochemistry needs to be taken into account, and a full canopy radiative transfer scheme, such as FluorMOD, must be developed.

L’anticipation des effets des changements climatiques nécessite une bonne compréhension dufonctionnement carboné des écosystèmes continentaux. L’une des principales contraintes liées àl’étude de ces écosystèmes est la forte variabilité à la fois spatiale et temporelle de leurs flux decarbone et de leurs réponses aux contraintes abiotiques. L’usage de méthodes de télédétectionoptiques pourrait permettre de suivre de façon spatialisée le fonctionnement des couverts végétaux.Ce travail vise à évaluer le potentiel de méthodes de télédétection pour décrire la structure et lefonctionnement de couverts végétaux à des échelles spatiales et temporelles variées. Pour ce faire,les relations entre indices optiques et phénomènes biologiques ont été étudiées en suivant unedémarche de transfert d’échelle, des échelles les plus fines aux plus larges. Il a été montré que le PRI(Photochemical Reflectance Index), utilisé en tant qu’indicateur du LUE (Light Use Efficiency), est parnature un signal composite qui reflète principalement la régulation du rendement de laphotosynthèse sur des échelles de temps fines, et la structure et composition biochimique ducouvert à l’échelle de la saison. L’analyse de courbes de réponse du PRI au PAR (PhotosyntheticallyActive Radiation) a permis de déconvoluer ces deux sources de variabilité, via l’introduction duconcept de PRI0 ou PRI d’une feuille idéalement adaptée à l’obscurité. Ce PRI0, capturant la variabilitédu PRI indépendante du LUE, a pu être mesuré à l’échelle de la feuille, et estimé à l’échelle de jeunescouverts végétaux et de la parcelle. Cette variabilité a pu être expliquée à l’échelle de la feuille et dejeunes couverts végétaux par les variations du contenu en pigment des feuilles. A l’échelle depeuplements adultes et de l’année, elle résulte cependant d’effets combinés de la compositionbiochimique et de la structure des couverts qui n’ont pu être séparés. Ces effets sont susceptiblesaux échelles larges de masquer en bonne partie, voire de biaiser la relation entre PRI et LUE. Il a enoutre été montré que la représentativité du PRI est limitée aux strates supérieures des canopées etdépend de la structure du couvert et du climat lumineux, ce qui peut limiter son intérêt en tantqu’estimateur du LUE à l’échelle de l’écosystème. Ces résultats soulignent la nécessité de prendre encompte la structure et la composition biochimique des couverts végétaux dans le cadre d’uneutilisation du PRI en tant que proxy du LUE de l’écosystème. / In order to assess the effect of global warming, a good understanding of carbon functioning ofterrestrial ecosystems is needed. The study of terrestrial ecosystem carbon fluxes and responses toabiotic stress remain challenging due to their high spatial and temporal variability. The use of remotesensing may help us to describe those sources of variability. The aim of this work is to assess thepotential of remote sensing as a way to describe canopy structure and functioning over a broadrange of temporal and spatial scales. The relationships between optical indices and biologicalphenomenon were investigated over a range of increasing scales. The PRI (PhotochemicalReflectance Index), used as a proxy of the LUE (Light Use Efficiency) was shown to be a compositesignal, mainly impacted by the regulation of the LUE at short time scales, and by canopy structureand pigment content at seasonal scale. The analysis of PRI response to PAR (PhotosyntheticallyActive Radiation) allowed us to deconvolve those two sources of variability thanks to theintroduction of the PRI0 defined as the PRI of ideally dark adapted leaves. The PRI0 was shown toefficiently describe the LUE unrelated PRI variability, and could be measured at leaf scale, andestimated at the leaf, canopy and stand scales. This variability could be explained by changes in leafpigment content over the growing season at leaf and canopy scales. At the stand scale and over theyear, this LUE independent PRI variability resulted from combined effects of canopy structure andpigment content, which could not be separated. These effects may result in biased or masked PRIversus LUE relationships at larges scales. Moreover, it was shown that the in-situ PRI measurementsmainly responded to the LUE of sunlit leaves, depending on canopy structure and sky conditions. Thismay considerably hamper the use of the PRI as a proxy of the whole ecosystem LUE. These resultsillustrate the need to take canopy structure and pigment content into account while using the PRI asa proxy of the ecosystem LUE.

Understanding the mechanism of photosynthetic seasonality in Amazonian evergreen forests is critical for its formulation in global climate and carbon cycle models. However, the control of the unexpected photosynthetic seasonality is highly uncertain. Here we use eddy-covariance data across a network of Amazonian research sites and a novel evapotranspiration (E) and two-leaf-photosynthesis-coupled model to investigate links between photosynthetic seasonality and climate factors on monthly scales. It reproduces the GPP seasonality (R-2=0.45-0.69) with a root-mean-square error (RMSE) of 0.67-1.25gCm(-2)d(-1) and a Bias of -0.03-1.04gCm(-2)d(-1) for four evergreen forest sites. We find that the proportion of diffuse and direct sunlight governs the photosynthetic seasonality via their interaction with sunlit and shaded leaves, supported by a proof that canopy light use efficiency (LUE) has a strong linear relationship with the fraction of diffuse sunlight for Amazonian evergreen forests. In the transition from dry season to rainy season, incident total radiation (Q) decreased while LUE and diffuse fraction increased, which produced the large seasonal increase (similar to 34%) in GPP of evergreen forests. We conclude that diffuse radiation is an important environmental driver of the photosynthetic seasonality in tropical Amazon forests yet depending on light utilization by sunlit and shaded leaves. Besides, the GPP model simulates the precipitation-dominated GPP seasonality (R-2=0.40-0.69) at pasture and savanna sites. These findings present an improved physiological method to relate light components with GPP in tropical Amazon. Plain Language Summary Understanding the mechanism of photosynthetic seasonality in Amazonian evergreen forests is critical for its formulation in global climate and carbon cycle models. However, the control of the unexpected photosynthetic seasonality is highly uncertain. Here we use eddy-covariance data across a network of Amazonian research sites and a novel evapotranspiration (E) and two-leaf-photosynthesis-coupled model to investigate links between photosynthetic seasonality and climate factors on monthly scales. It reproduces the GPP seasonality (R2= 0.45-0.69) for four evergreen forest sites. We find that the proportion of diffuse and direct sunlight governs the photosynthetic seasonality via their interaction with sunlit and shaded leaves, supported by a proof that canopy light-use efficiency (LUE) has a strong linear relationship with the fraction of diffuse sunlight for Amazonian evergreen forests. We conclude that diffuse radiation is an important environmental driver of the photosynthetic seasonality in tropical Amazon forests yet depending on light utilization by sunlit and shaded leaves. Besides, the GPP model simulates the precipitation-dominated GPP seasonality (R2= 0.40 similar to 0.69) at pasture and savanna sites. These findings present an improved physiological method to relate light components with GPP in Amazon.

Fotossíntese é o processo biofísico pelo qual energia luminosa é transformada em energia química armazenada em compostos de carbono. A taxa fotossintética instantânea possui um forte padrão assintótico em resposta ao incremento da intensidade luminosa, porém quando integramos a fotossíntese em escalas espaciais e temporais maiores, observa-se um padrão linear de resposta entre radiação interceptada e produção. Esta abordagem permitiu o surgimento de modelos baseados nas taxas de conversão de energia radiante em biomassa seca, ou eficiência do uso da luz (&epsilon;). Valores publicados para o Eucalyptus estão na faixa de 0,5-2,5 g MJ-1, porém se faz necessário um entendimento mais profundo a respeito da sensibilidade destes valores às flutuações do clima e sua sazonalidade. Para isso, as taxas de crescimento, uso e eficiência do uso da luz foram monitoradas quinzenalmente durante 16 meses em parcelas de 18 clones de Eucalyptus, dos 1,3 aos 2,7 anos de idade. Foram testadas as hipóteses de que a produção de madeira aumentaria em função de incrementos no uso e/ou eficiência de uso da luz, assim como estes valores aumentariam respectivamente com incrementos no índice de área foliar e por uma alocação de carbono para o fuste, respectivamente. Os clones apresentaram uma grande amplitude de produtividade (9,9-22,7 Mg ha-1 ano-1) e arquiteturas de copa, capturando entre 65-95% da radiação incidente. Tais valores resultaram em uma eficiência do uso da luz média de 1,5 g MJ-1, variando entre 0,16-3,14 g MJ-1. Apesar de patamares distintos, os valores de eficiência de uso dos clones oscilaram de maneira similar, de modo que a radiação incidente foi a principal variável afetando a eficiência de uso da luz, estando &epsilon; positivamente relacionada a variáveis que expressam períodos de maior disponibilidade hídrica e negativamente relacionado a períodos de menor disponibilidade. Maiores valores de índice de área foliar efetivo (Le) acarretaram em maior interceptação de luz, porém as distintas arquiteturas de copa revelaram diferentes estratégias de captura de luz (0,3 < &kappa; < 0,6). Apesar de uma maior interceptação, não houve correlação significativa com a produtividade, no entanto observou-se uma forte correlação entre eficiência do uso da luz e crescimento em madeira, resultado de uma maior alocação para o fuste. Apesar de evidenciar a relação entre alocação e eficiência, existem outros mecanismos associados às alterações observadas em &epsilon; que apenas uma caracterização completa dos fluxos de carbono pode elucidar. / Photosynthesis is the biophysical process by which light energy is converted into chemical energy stored in carbon compounds. The instantaneous photosynthetic rate has a strong asymptotic pattern in response to increases in light intensity, however when we integrate photosynthesis in larger spatial and temporal scales, there is a linear pattern of response between intercepted radiation and production. This approach has allowed the appearance of models based on radiant energy conversion rates into dry biomass, or light use efficiency (&epsilon;). Published values for Eucalyptus range from 0.5 to 2.5 g MJ-1, but a deeper understanding of the sensitivity of these values to climate fluctuations and seasonality is necessary. For this reason, wood growth rates, light use and efficiency were monitored every two weeks for 16 months at 18 Eucalyptus clones plots, from 1.3 to 2.7 years of age. Our hypothesis was that wood production would be positively related to light use and efficiency, as well these values would increase respectively with increases in leaf area index and carbon allocation to the stem. Clones showed a wide range of productivity (9.9 to 22.7 Mg ha-1 yr-1) and canopy architectures, capturing between 65-95% of incident radiation. Such values resulted in an average light use efficiency of 1.8 g MJ-1, ranging from 0.16 to 3.14 g MJ-1. Although different levels, light use efficiency values for the clones fluctuated similarly. Incident radiation was the main variable affecting the efficiency of dry matter conversion, and &epsilon; values were positively related variables expressing periods of greater water availability and negatively related to periods of lower availability. Larger effective leaf area index (Le) values resulted in higher light interception, but the different canopy architectures revealed different light capture strategies (0.3 < &kappa; < 0.6). Despite a higher interception, there was no significant correlation with productivity; however there was a strong correlation between light use efficiency and wood growth, as a result of increased allocation to the stem. While evidencing the relationship between allocation and efficiency, there are other mechanisms associated with changes in &epsilon; observed that only one full characterization of the carbon fluxes can elucidate.

Os objetivos do estudo foram identificar o melhor modelo de restauração pela manipulação da composição florística e práticas silviculturais, e compreender como este melhor desenvolvimento é explicado pelos fatores que controlam o uso e a eficiência do uso da luz pelo dossel florestal. Com esta fundamentação foi instalado em 2004, na Estação Experimental de Anhembi/ESALQ/USP, o ensaio \"Modelos de restauração de áreas degradadas da Mata Atlântica visando o seqüestro de carbono\", com 20 espécies nativas e oito sistemas de restauração florestal, baseados em um delineamento fatorial 23, com três fatores, e dois níveis por fator: i) Composição florística, com maior (67%) ou menor (50%) percentagem de pioneiras; ii) Espaçamento de plantio (3x1m e 3x2m); e iii) Manejo silvicultural, com manejo usual (maior estresse ambiental) e manejo máximo (minimização do estresse ambiental), referentes ao controle da matocompetição e fertilização. Em todos os tratamentos, até os 42 meses, foram determinados: produtividade primária líquida de madeira (PPLM), com base em inventários semestrais, determinação da densidade básica da madeira e fator de forma de todas as espécies, índice de área foliar (IAF, com calibração de fotos hemisféricas), radiação fotossinteticamente ativa interceptada (RFAI, de postos meteorológicos e o IAF), eficiência do uso da luz (EUL=PPLM/RFAI) e o conteúdo de nitrogênio no dossel. Em relação ao fator Composição, as proporções de 50% ou 67% de pioneiras foram equivalentes, em todas as idades, para PPLM, IAF e EUL. Em relação ao fator Espaçamento, o plantio mais adensado (3x1m) propiciou maior estoque de biomassa aos 42 meses, comparativamente ao espaçamento 3x2m (13,0 Mg.ha-1 versus 10,3 Mg.ha-1), devido às suas maiores produtividades até os 24 meses. Porém, as PPLMs de ambos os espaçamentos não diferiram estatisticamente a partir dos 36 meses (5,5 Mg.ha-1ano-1). A maior produtividade do espaçamento 3x1m foi diretamente relacionada ao seu maior índice de área foliar entre os 36 e 42 meses, comparativamente ao espaçamento 3x2m (0,98 m2m-2 versus 0,71 m2m-2). Em relação ao fator Manejo, o manejo máximo proporcionou aumento de 4 vezes na biomassa de madeira aos 42 meses, comparativamente ao manejo usual (18,6 Mg.ha-1 versus 4,6 Mg.ha-1). Este maior estoque foi explicado por maiores IAFs (1,34 m2m-2 versus 0,34 m2m-2), levando a maiores produtividades primárias de madeira (8,5 Mg.ha-1ano-1 versus 2,4 Mg.ha-1ano-1, aos 42 meses). A interação Espaçamento x Manejo foi significativa para as três variáveis de estudo: IAF, EUL, e PPLM, mostrando que as respostas dos espaçamentos 3x1m e 3x2m à aliviação do estresse ambiental são distintas, e que o espaçamento 3x2m requer uma silvicultura mais intensiva para expressar sua máxima produtividade. Finalmente, a PPLM mostrou-se altamente relacionada com o IAF no verão (36 meses) (r2=0,96), sendo que a incorporação do conteúdo de nitrogênio no dossel elevou ainda mais tal relação (r2=0,99), evidenciando o alto potencial de utilização destas duas variáveis como diagnóstico da qualidade e produtividade de florestas de restauração. Assim, para as condições deste estudo o melhor modelo seria: Composição com 50%:50% de pioneiras:não-pioneiras, espaçamento 3x2m, e necessariamente, manejo máximo, com controle da matocompetição das gramíneas (Brachiaria decumbens) e fertilização. / The purpose of this study was to indentify the most appropriated restoration model by manipulating floristic composition and silvicultural practices, and to understand how the best development was explained by the factors that controls light use and light use efficiency at the canopy. A \"Restoration models for Atlantic Forest disturbed lands aiming carbon sequestration\" experiment was installed in 2004 using 20 Brazilian woody species and 8 forest restoration systems, in a factorial 23 design, with 3 factors and two levels per factor: i) Floristic composition with 67% and 50% of pioneer species; ii) Plant spacing (3x1m e 3x2m); and iii) Silvicultural management with usual (higher environmental stress level); and maximum (lower environmental stress level) silvicultural practices due to weed control and fertilization. Wood net primary production (WNPP), based on inventory surveys and wood density of all 20 species, leaf area index (LAI, with hemispherical pictures), absorbed photossintetically active radiation (APAR, on meteorological stations and LAI), light use efficiency (LUE=WNPP/APAR) and canopy nitrogen content was measured or estimated up to 42 months-old. The floristic composition proportion, with 50% and 67% of pioneers, were statistically equivalent, in all ages, for WNPP, LAI and LUE. The 3x1m spacing provided larger biomass stock at age of 42 months, comparatively to the 3x2m planting (13.0 Mg.ha-1 versus 10.3 Mg. ha-1), due a higher productivity until 24 months. However, WNPP of both spacing were equivalent after 36 months (5.5 Mg.ha-1year-1). The higher productivity of the 3x1m spacing was directly related to its greater leaf area index comparatively to the 3x2m spacing (0,98 m2m-2 versus 0.71 m2m-2). The maximum silvicultural practices provided a 4-fold increase on wood biomass at age 42 months, comparatively to the usual practices (18.6 Mg.ha-1 versus 4.6 Mg.ha-1). This larger stock was explained by a higher LAI (1.34 m2m-2 versus 0.34 m2m-2) leading to higher WNPP (8.5 Mg.ha-1year-1 versus 2.4 Mg. ha-1year-1, at 42 months). The Spacing x Management interaction was significant for LAI, LUE and WNPP, meaning that 3x2m and 3x1m spacing had distinct responses to environmental stress levels, showing that 3x2m depends on intensive silvicultural practices to express its maximum productivity. Finally, the WNPP was strongly related to LAI during summer months (at 36 months) (r2=0,96), and this relationship increased by the inclusion of canopy nitrogen content, showing that these two variables have a potential to be used to evaluate forest restoration quality and productivity. Thus, for the study conditions, the best model was: Composition 50%:50% of pioneer:non-pioneer species, 3x2m spacing, and the maximum silvicultural practices, with total weed control (Brachiaria decumbens) and fertilization.

A produtividade dos plantios de eucalipto no Brasil apresentou ganhos significativos nas últimas décadas devido a avanços em melhoramento genético e silvicultura. Contudo, a produção de madeira representa apenas uma fração da produtividade primária bruta (GPP). Avaliar fluxos e partição de carbono (C) entre os diferentes componentes da floresta, e estudar o uso e a eficiência de uso dos recursos disponíveis é essencial para compreender os mecanismos que controlam a produtividade de plantios intensivamente manejados. O estudo quantificou os fluxos e partição de C e a eficiência de uso da luz para a produção de lenho (LUE) em 12 parcelas em um gradiente natural de produtividade, durante o sétimo ano de um plantio comercial de Eucalyptus grandis. Nessas mesmas parcelas, na escala da árvore, foram avaliadas a dominância do crescimento, produção de lenho e LUE, identificando a representatividade de árvores dominantes e suprimidas na produtividade do povoamento. O estudo do balanço de C e a aplicação da teoria da ecologia da produção em diferentes escalas objetivaram ampliar o conhecimento sobre os processos que governam a produtividade florestal. A heterogeneidade espacial dos atributos do solo e a topografia da área experimental influenciaram fortemente os fluxos componentes da GPP e sua partição, gerando um gradiente de produtividade. A produtividade de lenho variou de 554 gC m-2 ano-1 na parcela com menor GPP a 923 gC m-2 ano-1 na parcela com maior GPP. O fluxo de C para o solo variou de 497 gC m-2 ano-1 a 1235 gC m-2 ano-1 sem relação significativa com GPP. A partição do GPP para produção de lenho aumentou de 0,19 a 0,23, com tendência de aumento com o GPP (R2=0,30, p=0,07). A LUE aumentou em 66% (de 0,25 gC MJ-1 para 0,42 gC MJ-1) com a GPP, como resultado da elevação do fluxo e partição de C para produção de lenho. Ao longo do gradiente de produtividade, parcelas com alta eficiência quântica do dossel também mostraram alta LUE. A dominância do crescimento entre árvores teve forte impacto sobre a produtividade do povoamento. As 20% maiores árvores apresentaram em média 38% da biomassa de lenho e representaram 47% da produção de lenho. Características das folhas sugeriram que a maior produtividade de árvores dominantes, em relação às suprimidas, pode resultar de diferenças no controle estomático e não na capacidade fotossintética. A ecologia da produção na escala da árvore mostrou que os indivíduos dominantes produziram mais madeira por terem absorvido mais radiação e pela maior eficiência do uso da luz, comparativamente às árvores suprimidas. Em média, uma árvore suprimida cresceu 1,2 kg ano-1 de lenho, absorveu 2,9 GJ ano-1 de radiação e teve uma LUE de 0,4 g MJ-1. Já uma dominante cresceu 37 kg ano-1, absorveu 38 GJ ano-1 com mais que o dobro da eficiência (1,01 g MJ-1). Estudos sobre o balanço de carbono e ecologia da produção em diferentes escalas são essenciais para aperfeiçoar o conhecimento sobre os processos que controlam a produtividade de madeira e a fixação de carbono, e aprimorar os modelos ecofisiológicos. / The productivity of the eucalypt plantations in Brazil showed significant increase over the last decades, due to improvement in breeding and silviculture. However, wood production represents only a fraction of the gross primary production (GPP). Assessing carbon (C) fluxes and partitioning among forest components, and evaluate use and use efficiency of the available resources is essential to understand mechanisms driving productivity of intensively managed plantations. The study quantified fluxes and partitioning of C and light use efficiency for stem production (LUE) in 12 plots across a natural gradient of productivity during the seventh year of a commercial Eucalyptus grandis. Within these plots, at tree level, were evaluated growth dominance, stem production and LUE, identifying representativeness of dominant and suppressed trees to stand productivity. The study of C budget and the application of the production ecology theory at different levels aimed increase the knowledge about the processes driving forest productivity. The spatial heterogeneity of soil attributes and topography across the experimental site strongly influenced the component fluxes of GPP and partitioning, generating a gradient of productivity. Stem production ranged from 554 gC m-2 year-1 at the lowest GPP plot to 923 gC m-2 year-1 at the highest GPP plot. Total below ground carbon flux (TBCF) ranged from 497 g C m-2 year-1 to 1235 g C m-2 year-1, with no relationship to ANPP or GPP. Stem NPP:GPP partitioning ratio increased from 0.19 to 0.23 showing a trend of increase with GPP (R2=0.30, p=0.07). LUE increased by 66% (from 0.25 gC MJ-1 to 0.42 gC MJ-1) with GPP, as a result of the increased C partitioned and flux to stem NPP. Across the gradient of productivity, plots with the highest canopy quantum efficiency also showed the highest LUE. Growth dominance between trees showed a strong impact on stand productivity. The 20% larger trees accounted for 38% of stem biomass and represented 47% stem production. Leaf characteristics suggested that dominant trees were more productive, in relation to suppressed, may result in differences on stomatal control and not on photosynthetic capacity. The production ecology at tree level showed that dominant trees produced more wood by absorbing more radiation and due to higher light use efficiency, comparing to suppressed trees. On average, a suppressed tree grew 1,2 kg year-1 of stem, absorbed 2,9 GJ year-1 of radiation with a LUE of 0.4 g MJ-1. Although, a dominant grew 37 kg year-1 of stem, absorbed 38 GJ year-1 of radiation with the double of efficiency (1.01 g MJ-1). Studies regarding carbon balance and production ecology at different levels are essential to improve the knowledge on processes controlling wood production and carbon uptake, and develop ecophysiological models.

Os objetivos do estudo foram identificar o melhor modelo de restauração pela manipulação da composição florística e práticas silviculturais, e compreender como este melhor desenvolvimento é explicado pelos fatores que controlam o uso e a eficiência do uso da luz pelo dossel florestal. Com esta fundamentação foi instalado em 2004, na Estação Experimental de Anhembi/ESALQ/USP, o ensaio \"Modelos de restauração de áreas degradadas da Mata Atlântica visando o seqüestro de carbono\", com 20 espécies nativas e oito sistemas de restauração florestal, baseados em um delineamento fatorial 23, com três fatores, e dois níveis por fator: i) Composição florística, com maior (67%) ou menor (50%) percentagem de pioneiras; ii) Espaçamento de plantio (3x1m e 3x2m); e iii) Manejo silvicultural, com manejo usual (maior estresse ambiental) e manejo máximo (minimização do estresse ambiental), referentes ao controle da matocompetição e fertilização. Em todos os tratamentos, até os 42 meses, foram determinados: produtividade primária líquida de madeira (PPLM), com base em inventários semestrais, determinação da densidade básica da madeira e fator de forma de todas as espécies, índice de área foliar (IAF, com calibração de fotos hemisféricas), radiação fotossinteticamente ativa interceptada (RFAI, de postos meteorológicos e o IAF), eficiência do uso da luz (EUL=PPLM/RFAI) e o conteúdo de nitrogênio no dossel. Em relação ao fator Composição, as proporções de 50% ou 67% de pioneiras foram equivalentes, em todas as idades, para PPLM, IAF e EUL. Em relação ao fator Espaçamento, o plantio mais adensado (3x1m) propiciou maior estoque de biomassa aos 42 meses, comparativamente ao espaçamento 3x2m (13,0 Mg.ha-1 versus 10,3 Mg.ha-1), devido às suas maiores produtividades até os 24 meses. Porém, as PPLMs de ambos os espaçamentos não diferiram estatisticamente a partir dos 36 meses (5,5 Mg.ha-1ano-1). A maior produtividade do espaçamento 3x1m foi diretamente relacionada ao seu maior índice de área foliar entre os 36 e 42 meses, comparativamente ao espaçamento 3x2m (0,98 m2m-2 versus 0,71 m2m-2). Em relação ao fator Manejo, o manejo máximo proporcionou aumento de 4 vezes na biomassa de madeira aos 42 meses, comparativamente ao manejo usual (18,6 Mg.ha-1 versus 4,6 Mg.ha-1). Este maior estoque foi explicado por maiores IAFs (1,34 m2m-2 versus 0,34 m2m-2), levando a maiores produtividades primárias de madeira (8,5 Mg.ha-1ano-1 versus 2,4 Mg.ha-1ano-1, aos 42 meses). A interação Espaçamento x Manejo foi significativa para as três variáveis de estudo: IAF, EUL, e PPLM, mostrando que as respostas dos espaçamentos 3x1m e 3x2m à aliviação do estresse ambiental são distintas, e que o espaçamento 3x2m requer uma silvicultura mais intensiva para expressar sua máxima produtividade. Finalmente, a PPLM mostrou-se altamente relacionada com o IAF no verão (36 meses) (r2=0,96), sendo que a incorporação do conteúdo de nitrogênio no dossel elevou ainda mais tal relação (r2=0,99), evidenciando o alto potencial de utilização destas duas variáveis como diagnóstico da qualidade e produtividade de florestas de restauração. Assim, para as condições deste estudo o melhor modelo seria: Composição com 50%:50% de pioneiras:não-pioneiras, espaçamento 3x2m, e necessariamente, manejo máximo, com controle da matocompetição das gramíneas (Brachiaria decumbens) e fertilização. / The purpose of this study was to indentify the most appropriated restoration model by manipulating floristic composition and silvicultural practices, and to understand how the best development was explained by the factors that controls light use and light use efficiency at the canopy. A \"Restoration models for Atlantic Forest disturbed lands aiming carbon sequestration\" experiment was installed in 2004 using 20 Brazilian woody species and 8 forest restoration systems, in a factorial 23 design, with 3 factors and two levels per factor: i) Floristic composition with 67% and 50% of pioneer species; ii) Plant spacing (3x1m e 3x2m); and iii) Silvicultural management with usual (higher environmental stress level); and maximum (lower environmental stress level) silvicultural practices due to weed control and fertilization. Wood net primary production (WNPP), based on inventory surveys and wood density of all 20 species, leaf area index (LAI, with hemispherical pictures), absorbed photossintetically active radiation (APAR, on meteorological stations and LAI), light use efficiency (LUE=WNPP/APAR) and canopy nitrogen content was measured or estimated up to 42 months-old. The floristic composition proportion, with 50% and 67% of pioneers, were statistically equivalent, in all ages, for WNPP, LAI and LUE. The 3x1m spacing provided larger biomass stock at age of 42 months, comparatively to the 3x2m planting (13.0 Mg.ha-1 versus 10.3 Mg. ha-1), due a higher productivity until 24 months. However, WNPP of both spacing were equivalent after 36 months (5.5 Mg.ha-1year-1). The higher productivity of the 3x1m spacing was directly related to its greater leaf area index comparatively to the 3x2m spacing (0,98 m2m-2 versus 0.71 m2m-2). The maximum silvicultural practices provided a 4-fold increase on wood biomass at age 42 months, comparatively to the usual practices (18.6 Mg.ha-1 versus 4.6 Mg.ha-1). This larger stock was explained by a higher LAI (1.34 m2m-2 versus 0.34 m2m-2) leading to higher WNPP (8.5 Mg.ha-1year-1 versus 2.4 Mg. ha-1year-1, at 42 months). The Spacing x Management interaction was significant for LAI, LUE and WNPP, meaning that 3x2m and 3x1m spacing had distinct responses to environmental stress levels, showing that 3x2m depends on intensive silvicultural practices to express its maximum productivity. Finally, the WNPP was strongly related to LAI during summer months (at 36 months) (r2=0,96), and this relationship increased by the inclusion of canopy nitrogen content, showing that these two variables have a potential to be used to evaluate forest restoration quality and productivity. Thus, for the study conditions, the best model was: Composition 50%:50% of pioneer:non-pioneer species, 3x2m spacing, and the maximum silvicultural practices, with total weed control (Brachiaria decumbens) and fertilization.

Devido ao aprimoramento das práticas silviculturais e do melhoramento genético para seleção de genótipos mais produtivos, a produtividade do Pinus taeda no Brasil é reconhecida entre as maiores do mundo. Mesmo com alta produtividade em determinados sítios, sua variação é consideravelmente grande. Sabe-se que a disponibilidade de água, luz e nutrientes são fatores essenciais para a produção de madeira, onde a eficiência de uso desses recursos influenciam nesta variação do produtividade. Contudo, a ausência de fertilização em plantios de Pinus é majoritária nas áreas comerciais no Brasil, devido ao bom crescimento em solos de baixa fertilidade e por dificilmente apresentarem deficiência visual de nutrientes. Para compreender quais fatores influenciam na produção e partição de biomassa, se faz necessário o estudo da ecologia da produção. Diante disso, o presente trabalho tem como principais objetivos: i) estudar a influência de diferentes classes de solo no crescimento do Pinus taeda; ii) estudar a resposta da produtividade da espécie submetida à fertilização e ao desbaste; iii) avaliar a interceptação de luz; e iv) estimar a eficiência do crescimento (EC), uso da luz (EUL) e do nitrogênio (EUN) em povoamentos de Pinus taeda no sul do Brasil. Utilizou-se para este estudo parcelas quadrigêmeas da rede experimental do programa de Produtividade Potencial do Pinus no Brasil, as quais compõem um gradiente edafoclimático nos estados do Paraná e de Santa Catarina. As parcelas foram submetidas aos seguintes tratamentos: controle (C), fertilizado (F), desbastado (D), desbastado e fertilizado (DF). Foi calculado o índice de sítio, o incremento corrente anual, o incremento médio anual, índice de uniformidade do povoamento, a produtividade primária líquida de madeira e o índice de área foliar (IAF). Realizou-se análises de solo (0-20 cm) e foliar. Os dados das variáveis climáticas calculadas foram obtidos das estações meteorológicas mais próximas aos sítios avaliados. A produtividade mínima e máxima das parcelas controle foi de, respectivamente, 20,3 e 56,9 m³ ha-1 ano-1 com idade média de 13 anos. Houve efeito significativo dos tratamentos F e DF, com ganho máximo no estoque de volume por parcela de 33 e 63%, respectivamente. A resposta da fertilização e da fertilização com desbaste ocorreram em áreas menos férteis e com menor teor de matéria orgânica do solo. A uniformidade dos povoamentos correlacionou-se positivamente com as parcelas de maior área basal em 2014 e com a EC, EUL e EUN calculadas em 2013. Os valores médios da EC, EUL e EUN aos 12,4 anos de idade foram de 2,1 Mg ha-1 IAF-1, 0,55 g MJ-1 e 93 kg lenho kg-1 N ano-1, respectivamente. Houve relação linear positiva da EC, EUL e EUN com a produção de biomassa total acima do solo. Ocorreu maior influência das variáveis edáficas na eficiência do uso dos recursos naturais comparado com as variáveis climáticas. / Due to the improvement of silviculture practices and breeding for the selection of more productive genotypes, the loblolly pine productivity in Brazil is recognized as one of the biggest in the world. Even with the high productivity at certain sites, its variation is large. It is known that the availability of water, light and nutrients are essential for the wood production, and that the use efficiency of these resources influences the productivity variation. However, the absence of fertilization in pine plantations is majority in the commercial areas in Brazil due to good growth in low fertility soils and it hardly presents visual nutrient deficiency. To understand which factors influence the production and biomass partitioning, it is necessary to study the production ecology. Thus, the present work has as main objectives: i) to study the influence of different soil classes in loblolly pine growth; ii) to study the productivity response submitted to fertilization and thinning; iii) to evaluate the light interception; and iv) to estimate the growth efficiency (GE), use of light (LUE) and nitrogen (NUE) in loblolly pine stands in southern Brazil. For this study, it was used quadriplots from the experimental network of Pinus Potential Productivity in Brazil program, which composes an edaphoclimatic gradient in Parana and Santa Catarina states. The plots were submitted to the following treatments: control (C), fertilized (F), thinning (T), thinning and fertilized (TF). It was calculated the site index, the annual increment, the mean annual increment, the stand uniformity index, wood net primary productivity and the leaf area index (LAI). The soil (0-20 cm) and foliar analyzes were held. Data from the calculated climate variables was obtained from the closest weather stations to the evaluated sites. The minimum and maximum productivity of control plots were, respectively, 20.3 and 56.9 m³ ha-1 yr-1 with an average age of 13 years. There was a significant effect of F and TF treatments, with maximum gain in volume stock of 33 and 63%, respectively. The response of fertilization and fertilization with thinning occurred in less fertile areas and with lower soil organic matter content. The stands uniformity correlated positively with the basal area in 2014 and the GE, LUE and NUE calculated in 2013. The mean values of GE, LUE and NUE at the average age of 12,4 years were 2.1 Mg ha-1 LAI-1, 0.55 g MJ-1 and 93 kg wood kg-1 N year-1, respectively. There was a positive linear relationship of GE, LUE and NUE with the production of total biomass above ground. The soil variables influenced more on the use efficiency of natural resources than climatic variables.