O aumento para demanda na produção de alimentos associado a possíveis alterações climáticas globais tem promovido um interesse renovado em pesquisas envolvendo o metabolismo fotossintético. Acredita-se que com o melhoramento fotossintético a produtividade das culturas aumente. Para se alcançar este objetivo, além do uso de práticas biotecnológicas, ainda são necessários estudos abordando as limitações intrisecas do processo fotossintético que possam contribuir com a identificação de alvos para a engenharia genética. Neste contexto, esta tese aborda perguntas especificas sobre a regulação da fotossíntese C4 pela nutrição nitrogenada, o deficit hídrico e a interação destes dois fatores. No primeiro capítulo experimental foi desenvolvido um estudo envolvendo os efeitos da nutrição nitrogenada na difusão interna de CO2. Foi observado que o conteúdo de nitrogênio (N) foliar é determinante para a condutância interna de CO2 (? 18O-gm). Um menor conteúdo de N induz reduções iniciais em ? 18O-gm devido menor atividade das enzimas carboxilativas. Conforme o conteúdo de N reduz em folhas mais velhas alterações na antomia passaram a contribuir com a regulação de ? 18O-gm. No segundo capítulo, foi explorada a variação genotipica de cana-de-açúcar quanto a parâmetros relacionados a fotossíntese e o nitrogênio foliar, envolvendo a eficiência de uso de nitrogênio fotossintética (PNUE) e índices de herdabilidade. A variabilidade dos materiais genéticos foi explicada principalmente pelos parâmetros de trocas gasosas que contribuíram com o agrupamento dos materiais em três grupos. A partir deste agrupamento é possível verificar as diferenças entre os materiais em reposta a nutrição nitrogenada. Além disto, parâmetros de trocas gasosas como a taxa de assimilação de CO2 e condutância estomática junto com PNUE apresentaram altos índices de herdabilidade no sentido restrito. Por fim, o terceiro capítulo investigou a regulação do metabolismo de assimilação de CO2 em resposta ao deficit hídrico em folhas de cana-de-açúcar e como o metabolismo de nitrogênio está envolvido neste processo. Diferentes fatores envolvidos na regulação negativa da fotossíntese foram identicados. Interesantemente, alterações nos componentes nitrogenados só contribuíram com a inibição metabólica na fase de estresse severo. Adicionalmente foi observado que a superação da inibição metabólica durante a reidratação é mais rápida quando a cultivar apresenta decréscimos mais lentos no metabolismo de N no período de seca, associado a menores danos na capacidade fotossintética. / Increased demand for food production and the possibility of global climate change has promoted renewed interest in photosynthesis research. It is expected that with the photosynthetic improving the crop productivity increase. For this, besides of the use of biotechnological practices, further studies are needed about intrinsic limitations of the photosynthetic process that can contribute to the identification of targets for genetic engineering. In this context, this thesis discusses specific questions about photosynthesis regulation C4 by nitrogen nutrition, water deficit and the interaction of these two factors. In the chapter I, it was developed a study of the effects of nitrogen nutrition in the internal diffusion of CO2. It was observed that the leaf nitrogen (N) is essential for the internal CO2 conductance (Δ 18O-gm). A lower leaf N induces initial reductions in Δ18O-gm by lower activity of the carboxylation enzymes. With N leaf reduction in older leaves anatomy changes began to contribute to the regulation of Δ18 O-gm. In the chapter II, the genotypic variation of sugarcane to photosynthetic parameters and leaf nitrogen was explored, involving the photosynthetic nitrogen use efficiency (PNUE) and heritability analysis. The variability of the genetic material is explained mainly by gas exchange parameters that contributed to the grouping of materials into three groups. From this, it is possible examine the differences between the materials in response to nitrogen nutrition. Furthermore, gas exchange parameters such as CO2 assimilation rate and stomatal conductance along with PNUE showed high heritability in the narrow sense. Finally, the chapter III investigated the regulation of CO2 assimilation metabolism in response to water deficit sugarcane leaves and how nitrogen metabolism is involved in this process. Different factors involved in the negative regulation of photosynthesis were identified. Interestingly, changes in nitrogen components only contributed to metabolic inhibition in severe stress. Additionally it noted that overcoming metabolic inhibition during rehydration is faster when the cultivar presents slower decreases in N metabolism during water deficit, associated with less damage to the photosynthetic capacity.

Reliable prediction of climate change and its impact on and feedbacks from terrestrial carbon cycles requires realistic representation of physiological and ecological processes in coupled climate-carbon models. This is hampered by various deficiencies in model structures and parameters. The goal of my study is to improve model realism by incorporating latest advances of fundamental eco-physiological processes and further to use such improved models to investigate climate-carbon interactions at regional to global scales. I focus on the CO₂ diffusion within leaves (a key plant physiological process) and large-scale disturbances (a fundamental ecological process) as extremely important but not yet in current models. The CO₂ diffusion within plant leaves is characterized by mesophyll conductance (g[subscript m]), which strongly influences photosynthesis. I developed a g[subscript m] model by synthesizing new advances in plant-physiological studies and incorporated this model into the Community Land Model (CLM), a state-of-art climate-carbon model. I updated associated photosynthetic parameters based on a large dataset of leaf gas exchange measurements. Major findings are: (1) omission of g[subscript m] underestimates the maximum carboxylation rate and distorts its relationships with other parameters, leading to an incomplete understanding of leaf-level photosynthesis machinery; (2) proper representation of g[subscript m] is necessary for climate-carbon models to realistically predict carbon fluxes and their responsiveness to CO₂ fertilization; (3) fine tuning of parameters may compensate for model structural errors in contemporary simulations but introduce large biases in future predictions. Further, I have corrected a numerical deficiency of CLM in its calculation of carbon/water fluxes, which otherwise can bias model simulations. Large-scale disturbances of terrestrial ecosystems strongly affect their carbon sink strength. To provide insights for modeling these processes, I used satellite products to examine the temporal-spatial patterns of greenness after a massive ice storm. I found that the greenness of impacted vegetation recovered rapidly, especially in lightly and severely impacted regions. The slowest rebound occurred over moderately impacted areas. This nonlinear pattern was caused by an integrated effect of natural regrowth and human interventions. My results demonstrate mechanisms by which terrestrial carbon sinks could be significantly affected and help determine how these sinks will behave and so affect future climate. / text

Stomatal conductance and mesophyll conductance for CO2 transport are two key components of diffusive limitations of photosynthesis, since they restrict CO2 flux from the leaf surface to the sub-stomatal cavity and from there to the sites of carboxylation. This thesis summarizes our findings in the field of nature of mesophyll conductance to CO2 transport and its regulation per se and in respect to stomatal conductance.

abstract: Functional traits research has improved our understanding of how plants respond to their environments, identifying key trade-offs among traits. These studies primarily rely on correlative methods to infer trade-offs and often overlook traits that are difficult to measure (e.g., root traits, tissue senescence rates), limiting their predictive ability under novel conditions. I aimed to address these limitations and develop a better understanding of the trait space occupied by trees by integrating data and process models, spanning leaves to whole-trees, via modern statistical and computational methods. My first research chapter (Chapter 2) simultaneously fits a photosynthesis model to measurements of fluorescence and photosynthetic response curves, improving estimates of mesophyll conductance (gm) and other photosynthetic traits. I assessed how gm varies across environmental gradients and relates to other photosynthetic traits for 4 woody species in Arizona. I found that gm was lower at high aridity sites, varied little within a site, and is an important trait for obtaining accurate estimates of photosynthesis and related traits under dry conditions. Chapter 3 evaluates the importance of functional traits for whole-tree performance by fitting an individual-based model of tree growth and mortality to millions of measurements of tree heights and diameters to assess the theoretical trait space (TTS) of “healthy” North American trees. The TTS contained complicated, multi-variate structure indicative of potential trade-offs leading to successful growth. In Chapter 4, I applied an environmental filter (light stress) to the TTS, leading to simulated stand-level mortality rates up to 50%. Tree-level mortality was explained by 6 of the 32 traits explored, with the most important being radiation-use efficiency. The multidimentional space comprising these 6 traits differed in volume and location between trees that survived and died, indicating that selective mortality alters the TTS. / Dissertation/Thesis / Doctoral Dissertation Biology 2017

O aumento para demanda na produção de alimentos associado a possíveis alterações climáticas globais tem promovido um interesse renovado em pesquisas envolvendo o metabolismo fotossintético. Acredita-se que com o melhoramento fotossintético a produtividade das culturas aumente. Para se alcançar este objetivo, além do uso de práticas biotecnológicas, ainda são necessários estudos abordando as limitações intrisecas do processo fotossintético que possam contribuir com a identificação de alvos para a engenharia genética. Neste contexto, esta tese aborda perguntas especificas sobre a regulação da fotossíntese C4 pela nutrição nitrogenada, o deficit hídrico e a interação destes dois fatores. No primeiro capítulo experimental foi desenvolvido um estudo envolvendo os efeitos da nutrição nitrogenada na difusão interna de CO2. Foi observado que o conteúdo de nitrogênio (N) foliar é determinante para a condutância interna de CO2 (? 18O-gm). Um menor conteúdo de N induz reduções iniciais em ? 18O-gm devido menor atividade das enzimas carboxilativas. Conforme o conteúdo de N reduz em folhas mais velhas alterações na antomia passaram a contribuir com a regulação de ? 18O-gm. No segundo capítulo, foi explorada a variação genotipica de cana-de-açúcar quanto a parâmetros relacionados a fotossíntese e o nitrogênio foliar, envolvendo a eficiência de uso de nitrogênio fotossintética (PNUE) e índices de herdabilidade. A variabilidade dos materiais genéticos foi explicada principalmente pelos parâmetros de trocas gasosas que contribuíram com o agrupamento dos materiais em três grupos. A partir deste agrupamento é possível verificar as diferenças entre os materiais em reposta a nutrição nitrogenada. Além disto, parâmetros de trocas gasosas como a taxa de assimilação de CO2 e condutância estomática junto com PNUE apresentaram altos índices de herdabilidade no sentido restrito. Por fim, o terceiro capítulo investigou a regulação do metabolismo de assimilação de CO2 em resposta ao deficit hídrico em folhas de cana-de-açúcar e como o metabolismo de nitrogênio está envolvido neste processo. Diferentes fatores envolvidos na regulação negativa da fotossíntese foram identicados. Interesantemente, alterações nos componentes nitrogenados só contribuíram com a inibição metabólica na fase de estresse severo. Adicionalmente foi observado que a superação da inibição metabólica durante a reidratação é mais rápida quando a cultivar apresenta decréscimos mais lentos no metabolismo de N no período de seca, associado a menores danos na capacidade fotossintética. / Increased demand for food production and the possibility of global climate change has promoted renewed interest in photosynthesis research. It is expected that with the photosynthetic improving the crop productivity increase. For this, besides of the use of biotechnological practices, further studies are needed about intrinsic limitations of the photosynthetic process that can contribute to the identification of targets for genetic engineering. In this context, this thesis discusses specific questions about photosynthesis regulation C4 by nitrogen nutrition, water deficit and the interaction of these two factors. In the chapter I, it was developed a study of the effects of nitrogen nutrition in the internal diffusion of CO2. It was observed that the leaf nitrogen (N) is essential for the internal CO2 conductance (Δ 18O-gm). A lower leaf N induces initial reductions in Δ18O-gm by lower activity of the carboxylation enzymes. With N leaf reduction in older leaves anatomy changes began to contribute to the regulation of Δ18 O-gm. In the chapter II, the genotypic variation of sugarcane to photosynthetic parameters and leaf nitrogen was explored, involving the photosynthetic nitrogen use efficiency (PNUE) and heritability analysis. The variability of the genetic material is explained mainly by gas exchange parameters that contributed to the grouping of materials into three groups. From this, it is possible examine the differences between the materials in response to nitrogen nutrition. Furthermore, gas exchange parameters such as CO2 assimilation rate and stomatal conductance along with PNUE showed high heritability in the narrow sense. Finally, the chapter III investigated the regulation of CO2 assimilation metabolism in response to water deficit sugarcane leaves and how nitrogen metabolism is involved in this process. Different factors involved in the negative regulation of photosynthesis were identified. Interestingly, changes in nitrogen components only contributed to metabolic inhibition in severe stress. Additionally it noted that overcoming metabolic inhibition during rehydration is faster when the cultivar presents slower decreases in N metabolism during water deficit, associated with less damage to the photosynthetic capacity.

京都大学 / 0048 / 新制・課程博士 / 博士(農学) / 甲第21805号 / 農博第2318号 / 新制||農||1065(附属図書館) / 学位論文||H31||N5177(農学部図書室) / 京都大学大学院農学研究科農学専攻 / (主査)教授 白岩 立彦, 教授 奥本 裕, 教授 稲村 達也 / 学位規則第4条第1項該当 / Doctor of Agricultural Science / Kyoto University / DFAM

Les travaux de cette thèse se sont situés autour de la relation entre discrimination isotopique du carbone 13 et échanges gazeux foliaires. Le modèle établi par Farquhar et al. (1982) permet de prédire la discrimination contre le 13C pendant la photosynthèse (delta13C) en tenant compte des processus de diffusion, de carboxylation et décarboxylation engagés pendant la photosynthèse. Cette relation permet d'utiliser delta13C comme indicateur de l'efficience d'utilisation de l'eau (WUE, quantité de carbone fixé en fonction de l'eau consommée), un paramètre particulièrement important dans un contexte de changement climatique, d'agriculture et de sylviculture. Le modèle de delta13C a également été utilisé pour estimer la conductance mésophyllienne au CO2 (gm), un paramètre qui limite fortement la photosynthèse via la disponibilité en carbone dans le chloroplaste. Au cours de nos travaux, nous avons analysé le modèle delta13C pour identifier les paramètres les plus influents dans le modèle, et mis en évidence que l'utilisation du "modèle simple" de delta13C (ignorant gm et les processus de décarboxylation) peut induire un biais important dans l'estimation de WUE. Dans un second temps nous nous sommes concentrés sur les possibles variations à court-terme de gm, un domaine encore sujet a débat. Nous avons confirmé que gm était sensible aux variations de CO2 et d'irradiance sur toutes les espèces d'arbres mesurées dans cette étude. Nous avons aussi montré que ces variations rapides ne peuvent pas être dues a des variations des autres paramètres du modèle, à l'exception possible du paramètre b (discrimination pendant la carboxylation). Nous suggérons que les prochaines études dans ce domaine portent sur (i) la possible variabilité environnementale et génétique du paramètre b et (ii) les mécanismes à l'origine des variations rapides de gm (aquaporines et anhydrases carboniques) / This work was focused on the relationship between isotopic discrimination of 13C during photosynthesis (delta13C) and leaf gas exchange. The model of Farquhar and colleagues (Farquhar et al. 1982) predicts delta13C by accounting for diffusion, carboxylation and decarboxylation processes during the photosynthesis. This relationship is widely used and delta13C is frequently considered as a proxy water use efficiency (WUE, the amount of water required to fix a amount of carbon), an interesting parameter in the context of climate change, crop production and sylviculture. The delta13C model is also used to assess mesophyll conductance to CO2 (gm), that strongly limits photosynthesis via the availability of carbon in the chloroplast. Along this work we analyzed the delta13C model and identified the most important parameters, and highlighted that using the "simple form" of the model (which ignores gm and the decarboxylations) could lead to misestimating WUE. We also focused on the possible rapid variations of gm, a subject still under debate. We confirmed that gm was sensitive to rapid variations of CO2 and irradiance in all species tested in this study. We also showed that apparent rapid variations of gm could not be induced by variations of other parameters in the model, with the exception of parameter b (discrimination during carboxylation). We propose that future studies should focus on (i) the possible environmental and genetic variability of parameter b, and (ii) the physiological processes able to change gm at short time scales (aquaporins and carbonic anhydrase).

Mesophyll conductance is (together with stomatal conductance) a crucial component of diffusion limitations of photosynthesis and it is important to understand the mechanisms of CO2 fluxes through the leaves. Here I tested a new technique for estimation of drawdown in CO2 concentration across hypostomatous leaves based on carbon isotope composition (13C) of leaf cuticle and cuticular waxes isolated from opposite leaf sides.

No description available.

La sequera és una de les majors limitacions per a l’agricultura en general, factor que pot incrementar d’acord amb les prediccions del canvi climàtic. Per això, reduir l’ús de l’aigua en el reg i augmentar l’eficiència en l’ús de l’aigua (EUA) constitueix una de les majors prioritats per aconseguir una agricultura sostenible. L’EUA és un balanç entre guanys de biomassa i les despeses d’aigua. Aquesta Tesi s’ha centrat en l’estudi de 3 dels processos fisiològics que afecten a l’ús de l’aigua i a la producció de la planta en el cas de la vinya, i que per tant poden ser considerats factors potencials per millorar l’EUA: (1) conductància del mesòfil, (2) transpiració nocturna (3) respiració. Els resultats d’aquesta Tesi revelen que la conductància del mesòfil i la respiració són els principals components per millorar l’EUA permetent millorar l’assimilació de carboni o minimitzant les pèrdues del carboni fixat per la fotosíntesis