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Although widely used in ecology, trait-based approaches are seldom used to study agroecosystems. In particular, there is a need to evaluate how functional trait variability among varieties of a crop species compares to the variability among wild plant species and how variety selection can modify trait syndromes. Here, we quantified 18 above- and below-ground functional traits for 57 varieties of common wheat representative of different modern selection histories. We compared trait variability among varieties and among Pooideae species, and analyzed the effect of selection histories on trait values and trait syndromes. For traits under strong selection, trait variability among varieties was less than 10% of the variability observed among Pooideae species. However, for traits not directly selected, such as root N uptake capacity, the variability was up to 75% of the variability among Pooideae species. Ammonium absorption capacity by roots was counter-selected for conventional varieties compared with organic varieties and landraces. Artificial selection also altered some trait syndromes classically reported for Pooideae. Identifying traits that have high or low variability among varieties and characterizing the hidden effects of selection on trait values and syndromes will benefit the selection of varieties to be used especially for lower N input agroecosystems. [ABSTRACT FROM AUTHOR]

Simulating leaf development from initiation to maturity opens new possibilities to model plant–environment interactions and the plasticity of plant architecture. This study analyses the dynamics of leaf production and extension along a maize (Zea mays) shoot to assess important modelling choices. Maize plants from two cultivars originating from the same inbred line, yet differing in the length of mature leaves were used in this study. We characterized the dynamics of the blade and sheath lengths of all phytomers by dissecting plants every 2–3 days. We analysed how differences in leaf size were built up and we examined the coordination between the emergence of organs and phases of their extension. Leaf extension rates were higher in the cultivar with longer leaves than in the cultivar with shorter leaves; no differences were found in other aspects. We found that (i) first post-embryonic leaves were initiated at a markedly higher rate than upper leaves; (ii) below ear position, sheaths were initiated at a time intermediate between tip emergence and appearance, while above the ear position, sheaths were initiated at a high rate, such that the time interval between the blade and sheath initiations decreased for these leaves; and (iii) ear position also marked a change in the correlation in size between successive phytomers with little correlation of size between upper and lower leaves. Our results identified leaf extension rate as the reason for the difference in size between the two cultivars. The two cultivars shared the same pattern for the timing of initiation events, which was more complex than previously thought. The differences described here may explain some inaccuracies reported in functional–structural plant models. We speculate that genotypic variation in behaviour for leaf and sheath initiation exists, which has been little documented in former studies. [ABSTRACT FROM AUTHOR]

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]

The article evaluates the wheat model WALTer to study competition for light for predicting the tillering dynamics at contrasting sowing densities. Topics discussed include simulation of the three-dimensional (3D) development of the aerial architecture of winter wheat with WALTer, cessation of tillering to sense the spectral quality of light and the impact of the tillering process on crop yield.

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. [ABSTRACT FROM AUTHOR]

Background and Aims Simulating resource allocation in crops requires an integrated view of plant functioning and the formalization of interactions between carbon (C) and nitrogen (N) metabolisms. This study evaluates the functional–structural model CN-Wheat developed for winter wheat after anthesis. Methods In CN-Wheat the acquisition and allocation of resources between photosynthetic organs, roots and grains are emergent properties of sink and source activities and transfers of mobile metabolites. CN-Wheat was calibrated for field plants under three N fertilizations at anthesis. Model parameters were taken from the literature or calibrated on the experimental data. Key Results The model was able to predict the temporal variations and the distribution of resources in the culm. Thus, CN-Wheat accurately predicted the post-anthesis kinetics of dry masses and N content of photosynthetic organs and grains in response to N fertilization. In our simulations, when soil nitrates were non-limiting, N in grains was ultimately determined by availability of C for root activity. Dry matter accumulation in grains was mostly affected by photosynthetic organ lifespan, which was regulated by protein turnover and C-regulated root activity. Conclusions The present study illustrates that the hypotheses implemented in the model were able to predict realistic dynamics and spatial patterns of C and N. CN-Wheat provided insights into the interplay of C and N metabolism and how the depletion of mobile metabolites due to grain filling ultimately results in the cessation of resource capture. This enabled us to identify processes that limit grain mass and protein content and are potential targets for plant breeding. [ABSTRACT FROM AUTHOR]

Background and Aims Improving crops requires better linking of traits and metabolic processes to whole plant performance. In this paper, we present CN-Wheat, a comprehensive and mechanistic model of carbon (C) and nitrogen (N) metabolism within wheat culms after anthesis. Methods The culm is described by modules that represent the roots, photosynthetic organs and grains. Each of them includes structural, storage and mobile materials. Fluxes of C and N among modules occur through a common pool and through transpiration flow. Metabolite variations are represented by differential equations that depend on the physiological processes occurring in each module. A challenging aspect of CN-Wheat lies in the regulation of these processes by metabolite concentrations and the environment perceived by organs. Key Results CN-Wheat simulates the distribution of C and N into wheat culms in relation to photosynthesis, N uptake, metabolite turnover, root exudation and tissue death. Regulation of physiological activities by local concentrations of metabolites appears to be a valuable feature for understanding how the behaviour of the whole plant can emerge from local rules. Conclusions The originality of CN-Wheat is that it proposes an integrated view of plant functioning based on a mechanistic approach. The formalization of each process can be further refined in the future as knowledge progresses. This approach is expected to strengthen our capacity to understand plant responses to their environment and investigate plant traits adapted to changes in agronomical practices or environmental conditions. A companion paper will evaluate the model. [ABSTRACT FROM AUTHOR]

Background and Aims Experimental evidence challenges the approximation, central in crop models, that developmental events follow a fixed thermal time schedule, and indicates that leaf emergence events play a role in the timing of development. The objective of this study was to build a structural development model of maize (Zea mays) based on a set of coordination rules at organ level that regulate duration of elongation, and to show how the distribution of leaf sizes emerges from this. Methods A model of maize development was constructed based on three coordination rules between leaf emergence events and the dynamics of organ extension. The model was parameterized with data from maize grown at a low plant population density and tested using data from maize grown at high population density. Key Results The model gave a good account of the timing and duration of organ extension. By using initial conditions associated with high population density, the model reproduced well the increase in blade elongation duration and the delay in sheath extension in high-density populations compared with low-density populations. Predictions of the sizes of sheaths at high density were accurate, whereas predictions of the dynamics of blade length were accurate up to rank 9; moderate overestimation of blade length occurred at higher ranks. Conclusions A set of simple rules for coordinated growth of organs is sufficient to simulate the development of maize plant structure without taking into account any regulation by assimilates. In this model, whole-plant architecture is shaped through initial conditions that feed a cascade of coordination events. [ABSTRACT FROM AUTHOR]

Background and Aims Sustainable agriculture requires the identification of new, environmentally responsible strategies of crop protection. Modelling of pathosystems can allow a better understanding of the major interactions inside these dynamic systems and may lead to innovative protection strategies. In particular, functional–structural plant models (FSPMs) have been identified as a means to optimize the use of architecture-related traits. A current limitation lies in the inherent complexity of this type of modelling, and thus the purpose of this paper is to provide a framework to both extend and simplify the modelling of pathosystems using FSPMs. Methods Different entities and interactions occurring in pathosystems were formalized in a conceptual model. A framework based on these concepts was then implemented within the open-source OpenAlea modelling platform, using the platform's general strategy of modelling plant–environment interactions and extending it to handle plant interactions with pathogens. New developments include a generic data structure for representing lesions and dispersal units, and a series of generic protocols to communicate with objects representing the canopy and its microenvironment in the OpenAlea platform. Another development is the addition of a library of elementary models involved in pathosystem modelling. Several plant and physical models are already available in OpenAlea and can be combined in models of pathosystems using this framework approach. Key Results Two contrasting pathosystems are implemented using the framework and illustrate its generic utility. Simulations demonstrate the framework's ability to simulate multiscaled interactions within pathosystems, and also show that models are modular components within the framework and can be extended. This is illustrated by testing the impact of canopy architectural traits on fungal dispersal. Conclusions This study provides a framework for modelling a large number of pathosystems using FSPMs. This structure can accommodate both previously developed models for individual aspects of pathosystems and new ones. Complex models are deconstructed into separate ‘knowledge sources’ originating from different specialist areas of expertise and these can be shared and reassembled into multidisciplinary models. The framework thus provides a beneficial tool for a potential diverse and dynamic research community. [ABSTRACT FROM AUTHOR]

Background and Aims The relationship between Septoria tritici, a splash-dispersed disease, and its host is complex because of the interactions between the dynamic plant architecture and the vertical progress of the disease. The aim of this study was to test the capacity of a coupled virtual wheat–Septoria tritici epidemic model (Septo3D) to simulate disease progress on the different leaf layers for contrasted sowing density treatments. Methods A field experiment was performed with winter wheat ‘Soissons’ grown at three contrasted densities. Plant architecture was characterized to parameterize the wheat model, and disease dynamic was monitored to compare with simulations. Three simulation scenarios, differing in the degree of detail with which plant variability of development was represented, were defined. Key Results Despite architectural differences between density treatments, few differences were found in disease progress; only the lower-density treatment resulted in a slightly higher rate of lesion development. Model predictions were consistent with field measurements but did not reproduce the higher rate of lesion progress in the low density. The canopy reconstruction scenario in which inter-plant variability was taken into account yielded the best agreement between measured and simulated epidemics. Simulations performed with the canopy represented by a population of the same average plant deviated strongly from the observations. Conclusions It was possible to compare the predicted and measured epidemics on detailed variables, supporting the hypothesis that the approach is able to provide new insights into the processes and plant traits that contribute to the epidemics. On the other hand, the complex and dynamic responses to sowing density made it difficult to test the model precisely and to disentangle the various aspects involved. This could be overcome by comparing more contrasted and/or simpler canopy architectures such as those resulting from quasi-isogenic lines differing by single architectural traits. [ABSTRACT FROM PUBLISHER]

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. [ABSTRACT FROM PUBLISHER]

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. [ABSTRACT FROM PUBLISHER]

Background and Aims The phenotypes of grasses show differences depending on growth conditions and ontogenetic stage. Understanding these responses and finding suitable mathematical formalizations are an essential part of the development of plant and crop models. Usually, a marked change in architecture between juvenile and adult plants is observed, where dimension and shape of leaves are likely to change. In this paper, the plasticity of leaf shape is analysed according to growth conditions and ontogeny. Methods Leaf shape of Triticum aestivum, Hordeum vulgare and Zea mays cultivars grown under varying conditions was measured using digital image processing. An empirical leaf shape model was fitted to measured shape data of single leaves. Obtained values of model parameters were used to analyse the patterns in leaf shape. Key Results The model was able to delineate leaf shape of all studied species. The model error was small. Differences in leaf shape between juvenile and adult leaves in T. aestivum and H. vulgare were observed. Varying growth conditions impacted leaf dimensions but did not impact leaf shape of the respective species. Conclusions Leaf shape of the studied T. aestivum and H. vulgare cultivars was remarkably stable for a comparable ontogenetic stage (leaf rank), but differed between stages. Along with other aspects of grass architecture, leaf shape changed during the transition from juvenile to adult growth phase. Model-based analysis of leaf shape is a method to investigate these differences. Presented results can be integrated into architectural models of plant development to delineate leaf shape for different species, cultivars and environmental conditions. [ABSTRACT FROM PUBLISHER]

In monocarpic species, during the reproductive stage the growing grams 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 aestivuni). 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 cuims, 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. [ABSTRACT FROM AUTHOR]

• Background and Aims Plants regulate their architecture strongly in response to density, and there is evidence that this involves changes in the duration of leaf extension. This questions the approximation, central in crop models, that development follows a fixed thermal time schedule. The aim of this research is to investigate, using maize as a model, how the kinetics of extension of grass leaves change with density, and to propose directions for inclusion of this regulation in plant models.• Methods Periodic dissection of plants allowed the establishment of the kinetics of lamina and sheath extension for two contrasting sowing densities. The temperature of the growing zone was measured with thermocouples. Two-phase (exponential plus linear) models were fitted to the data, allowing analysis of the timing of the phase changes of extension, and the extension rate of sheaths and blades during both phases.• Key Results The duration of lamina extension dictated the variation in lamina length between treatments. The lower phytomers were longer at high density, with delayed onset of sheath extension allowing more time for the lamina to extend. In the upper phytomers—which were shorter at high density—the laminae had a lower relative extension rate (RER) in the exponential phase and delayed onset of linear extension, and less time available for extension since early sheath extension was not delayed.• Conclusions The relative timing of the onset of fast extension of the lamina with that of sheath development is the main determinant of the response of lamina length to density. Evidence is presented that the contrasting behaviour of lower and upper phytomers is related to differing regulation of sheath ontogeny before and after panicle initiation. A conceptual model is proposed to explain how the observed asynchrony between lamina and sheath development is regulated. [ABSTRACT FROM AUTHOR]

• Background and Aims The production of axillary shoots (tillering) in spring wheat (Triticum aestivum) depends on intraspecific competition. The mechanisms that underlie this competition are complex, but light within the wheat canopy plays a key role. The main objectives of this paper are to analyse the effects of plant population density and shade on tillering dynamics of spring wheat, to assess the canopy conditions quantitatively at the time of tillering cessation, and to analyse the relationship between the tiller bud and the leaf on the same phytomer.• Methods Spring wheat plants were grown at three plant population densities and under two light regimes (25 % and 100 % light). Tiller appearance, fraction of the light intercepted, and red : far-red ratio at soil level were recorded. On six sampling dates the growth status of axillary buds was analysed.• Key Results Tillering ceased earlier at high population densities and ceased earlier in the shade than in full sunlight. At cessation of tillering, both the fraction of light intercepted and the red : far-red ratio at soil level were similar in all treatments. Leaves on the same phytomer of buds that grew out showed more leaf mass per unit area than those on the same phytomer of buds that remained dormant.• Conclusions Tillering ceases at specific light conditions within the wheat canopy, independent of population density, and to a lesser extent independent of light intensity. It is suggested that cessation of tillering is induced when the fraction of PAR intercepted by the canopy exceeds a specific threshold (0·40–0·45) and red : far-red ratio drops below 0·35–0·40. [ABSTRACT FROM AUTHOR]

• Background and Aims Fitting the parameters of models of plant organ growth is a means to investigate how environmental conditions affect plant architecture. The aim of this article is to evaluate some non-linear methods for fitting the parameters of multi-phase models of the kinetics of extension of plant organs such as laminae, sheaths and internodes.• Methods A set of computational procedures was developed allowing parameter-fitting of multi-phase models, using the maximum likelihood criterion, in which phases are identified with reference to ontogenic processes. Two bootstrap methods were compared to assess the precision of the estimates of fitted parameters, and of functions of these parameters such as the final leaf length, and the duration and rate of the rapid extension phase. Methods were applied to an experimental dataset, representing the kinetics of laminae, sheaths and internodes along the maize shoot, for two contrasting densities.• Key Results A set of multi-phase models was proposed to describe the extension of laminae, sheaths and internodes along the shoot. The distinguishable phases differed between laminae, sheaths and internodes. For sheaths and laminae, but not for internodes, the same model could be fitted to all phytomers along the shoot. The variation of parameters along the shoot and between density treatments, as well as derived functions such as the durations of the phases of extension, are presented for laminae. It was the duration of the fast extension period, rather than its rate, which determined the difference in final length between treatments.• Conclusions Such methods permit a large degree of objectivity and facilitate the analysis of such rather complicated but co-ordinated datasets. The work also illustrates some natural limitations of maximum likelihood methods, and viable ways of overcoming them by including a priori knowledge in the model fitting method are discussed. [ABSTRACT FROM AUTHOR]