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Background and Aims: Because functional-structural plant models (FSPMs) take plant architecture explicitly into consideration, they constitute a promising approach for unravelling plant-plant interactions in complex canopies. However, existing FSPMs mainly address competition for light. The aim of the present work was to develop a comprehensive FSPM accounting for the interactions between plant architecture, environmental factors and the metabolism of carbon (C) and nitrogen (N)., Methods: We developed an original FSPM by coupling models of (1) 3-D wheat architecture, (2) light distribution within canopies and (3) C and N metabolism. Model behaviour was evaluated by simulating the functioning of theoretical canopies consisting of wheat plants of contrasting leaf inclination, arranged in pure and mixed stands and considering four culm densities and three sky conditions., Key Results: As an emergent property of the detailed description of metabolism, the model predicted a linear relationship between absorbed light and C assimilation, and a curvilinear relationship between grain mass and C assimilation, applying to both pure stands and each component of mixtures. Over the whole post-anthesis period, planophile plants tended to absorb more light than erectophile plants, resulting in a slightly higher grain mass. This difference was enhanced at low plant density and in mixtures, where the erectophile behaviour resulted in a loss of competitiveness., Conclusion: The present work demonstrates that FSPMs provide a framework allowing the analysis of complex canopies such as studying the impact of particular plant traits, which would hardly be feasible experimentally. The present FSPM can help in interpreting complex interactions by providing access to critical variables such as resource acquisition and allocation, internal metabolic concentrations, leaf life span and grain filling. Simulations were based on canopies identically initialized at flowering; extending the model to the whole cycle is thus required so that all consequences of a trait can be evaluated., (© The Author(s) 2018. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Background and Aims: Models simulating nitrogen use by plants are potentially efficient tools to optimize the use of fertilizers in agriculture. Most crop models assume that a target nitrogen concentration can be defined for plant tissues and formalize a demand for nitrogen, depending on the difference between the target and actual nitrogen concentrations. However, the teleonomic nature of the approach has been criticized. This paper proposes a mechanistic model of nitrogen economy, NEMA (Nitrogen Economy Model within plant Architecture), which links nitrogen fluxes to nitrogen concentration and physiological processes., Methods: A functional-structural approach is used: plant aerial parts are described in a botanically realistic way and physiological processes are expressed at the scale of each aerial organ or root compartment as a function of local conditions (light and resources)., Key Results: NEMA was developed for winter wheat (Triticum aestivum) after flowering. The model simulates the nitrogen (N) content of each photosynthetic organ as regulated by Rubisco turnover, which depends on intercepted light and a mobile N pool shared by all organs. This pool is enriched by N acquisition from the soil and N release from vegetative organs, and is depleted by grain uptake and protein synthesis in vegetative organs; NEMA accounts for the negative feedback from circulating N on N acquisition from the soil, which is supposed to follow the activities of nitrate transport systems. Organ N content and intercepted light determine dry matter production via photosynthesis, which is distributed between organs according to a demand-driven approach., Conclusions: NEMA integrates the main feedbacks known to regulate plant N economy. Other novel features are the simulation of N for all photosynthetic tissues and the use of an explicit description of the plant that allows how the local environment of tissues regulates their N content to be taken into account. We believe this represents an appropriate frame for modelling nitrogen in functional-structural plant models. A companion paper will present model evaluation and analysis.

Background and Aims: Simulating nitrogen economy in crop plants requires formalizing the interactions between soil nitrogen availability, root nitrogen acquisition, distribution between vegetative organs and remobilization towards grains. This study evaluates and analyses the functional-structural and mechanistic model of nitrogen economy, NEMA (Nitrogen Economy Model within plant Architecture), developed for winter wheat (Triticum aestivum) after flowering., Methods: NEMA was calibrated for field plants under three nitrogen fertilization treatments at flowering. Model behaviour was investigated and sensitivity to parameter values was analysed., Key Results: Nitrogen content of all photosynthetic organs and in particular nitrogen vertical distribution along the stem and remobilization patterns in response to fertilization were simulated accurately by the model, from Rubisco turnover modulated by light intercepted by the organ and a mobile nitrogen pool. This pool proved to be a reliable indicator of plant nitrogen status, allowing efficient regulation of nitrogen acquisition by roots, remobilization from vegetative organs and accumulation in grains in response to nitrogen treatments. In our simulations, root capacity to import carbon, rather than carbon availability, limited nitrogen acquisition and ultimately nitrogen accumulation in grains, while Rubisco turnover intensity mostly affected dry matter accumulation in grains., Conclusions: NEMA enabled interpretation of several key patterns usually observed in field conditions and the identification of plausible processes limiting for grain yield, protein content and root nitrogen acquisition that could be targets for plant breeding; however, further understanding requires more mechanistic formalization of carbon metabolism. Its strong physiological basis and its realistic behaviour support its use to gain insights into nitrogen economy after flowering.

In monocarpic species, during the reproductive stage the growing grains represent a strong sink for nitrogen (N) and trigger N remobilization from the vegetative organs, which decreases canopy photosynthesis and accelerates leaf senescence. The spatiotemporal distribution of N in a reproductive canopy has not been described in detail. Here, we investigated the role of the local light environment on the spatiotemporal distribution of leaf lamina N mass per unit leaf area (SLN) during grain filling of field-grown wheat (Triticum aestivum). In addition, in order to provide some insight into the coordination of N depletion between the different vegetative organs, N dynamics were studied for individual leaf laminae, leaf sheaths, internodes, and chaff of the top fertile culms. At the canopy scale, SLN distribution paralleled the light gradient below the flag leaf collar until almost the end of grain filling. On the contrary, the significant light gradient along the flag leaf lamina was not associated with a SLN gradient. Within the top fertile culms, the time course of total (alive + necrotic tissues) N concentration of the different laminae and sheaths displayed a similar pattern. Another common pattern was observed for internodes and chaff. During the period of no root N uptake, N depletion of individual laminae and sheaths followed a first-order kinetics independent of leaf age, genotype, or N nutrition. The results presented here show that during grain filling, N dynamics are integrated at the culm scale and strongly depend on the local light conditions determined by the canopy structure.

Here, nitrogen management within the plant was compared in an early-senescing maize hybrid and in a late-senescing maize hybrid, both grown under field conditions with a high fertilisation input involving large quantities of fertiliser. We monitored, in representative leaf stages, the changes in metabolite content, enzyme activities and steady-state levels of transcripts for marker genes of N primary assimilation, N recycling and leaf senescence. The hybrids differed in terms of persistence of leaf greenness, the expression of marker genes and the concentration of enzymes used to describe the transition from N assimilation to N recycling. The transcription of leaf-senescence marker genes did not differ. Agronomic studies confirmed the ability of the late-senescing hybrid to absorb and store more N in shoots. Despite the differences in the mode of N management adopted by the two hybrids, we conclude that leaf senescence occurs independently of the source-to-sink transition at the high level of fertilisation used involving large quantities of fertiliser. The possibility of improving N metabolic efficiency in the latest maize hybrids is discussed., (Copyright New Phytologist (2005).)

Shoot architecture is a key component of the interactions between plants and their environment. We present a novel model of grass, which fully integrates shoot morphogenesis and the metabolism of carbon (C) and nitrogen (N) at organ scale, within a three-dimensional representation of plant architecture. Plant morphogenesis is seen as a self-regulated system driven by two main mechanisms. First, the rate of organ extension and the establishment of architectural traits are regulated by concentrations of C and N metabolites in the growth zones and the temperature. Second, the timing of extension is regulated by rules coordinating successive phytomers instead of a thermal time schedule. Local concentrations are calculated from a model of C and N metabolism at organ scale. The three-dimensional representation allows the accurate calculation of light and temperature distribution within the architecture. The model was calibrated for wheat (Triticum aestivum) and evaluated for early vegetative stages. This approach allowed the simulation of realistic patterns of leaf dimensions, extension dynamics, and organ mass and composition. The model simulated, as emergent properties, plant and agronomic traits. Metabolic activities of growing leaves were investigated in relation to whole-plant functioning and environmental conditions. The current model is an important step towards a better understanding of the plasticity of plant phenotype in different environments. [ABSTRACT FROM AUTHOR]

Nitrogen (N) distribution among plant organs plays a major role in crop production and, in general, plant fitness to the environment. In the present study, a process-based model simulating N distribution within a wheat (Triticum aestivumL.) culm during grain filling was developed using a functional–structural approach. A model of turnover of the photosynthetic apparatus was used to describe the fluxes between a common pool of mobile N and each leaf lamina. Grain N accumulation within a time-step was modelled as the minimum between the quantity calculated by a potential function and the N available in the common pool. Nitrogen dynamics in the other organs (i.e. stem, chaff, root N uptake and remobilisation) were accounted for by forced variables. Using a unique set of six parameters, the model was able to simulate the observed N kinetics of each lamina and of the grains under a wide range of crop N supplies and for three cultivars. The time-course of the vertical gradient of lamina N during grain filling was realistically simulated as an emerging property of the local processes defined at the lamina scale. The model described in the present study offers new insight into the interactions between N metabolism, plant architecture and productivity. [ABSTRACT FROM AUTHOR]

To illustrate the development of the source-to-sink transition in maize leaves during the grain-filling period, an integrated physiological-agronomic approach is presented in this study. The evolution of physiological markers such as total leaf nitrogen (N), chlorophyll, soluble protein, amino acid and ammonium contents was monitored from silking to a period close to maturity in different leaf stages of three maize genotypes grown at high and low levels of N fertilization. In addition, the activities of glutamine synthetase (GS) and glutamate dehydrogenase (GDH), two enzymes known to play a direct or an indirect role during leaf N remobilization, were measured. In the three genotypes examined, we found that a general decrease of most metabolic and enzyme markers occurred during leaf ageing and that this decrease was enhanced when plants were N starved. In contrast, such variations were not observed between different sections of a single leaf even at an advanced stage of leaf senescence. We found that there is a strong correlation between total N, chlorophyll, soluble protein and GS activity, which is not dependent upon the N fertilization level, which indicates the N status of the plant, either in a single leaf or during ageing. In contrast, ammonium, amino acids and GDH activity were not subject to such variations, thus suggesting that they are indicators of the metabolic activity of the whole plant in response to the level of N fertilization. The use of these markers to predict the N status of maize as a function of both plant development and N availability is discussed. [ABSTRACT FROM AUTHOR]