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Plants in dense vegetation perceive their neighbors primarily through changes in light quality. Initially, the ratio between red (R) and far-red (FR) light decreases due to reflection of FR by plant tissue well before shading occurs. Perception of low R:FR by the phytochrome photoreceptors induces the shade avoidance response [1], of which accelerated elongation growth of leaf-bearing organs is an important feature. Low R:FR-induced phytochrome inactivation leads to the accumulation and activation of the transcription factors PHYTOCHROME-INTERACTING FACTORs (PIFs) 4, 5, and 7 and subsequent expression of their growth-mediating targets [2, 3]. When true shading occurs, transmitted light is especially depleted in red and blue (B) wavelengths, due to absorption by chlorophyll [4]. Although the reduction of blue wavelengths alone does not occur in nature, long-term exposure to low B light induces a shade avoidance-like response that is dependent on the cryptochrome photoreceptors and the transcription factors PIF4 and PIF5 [5–7]. We show in Arabidopsis thaliana that low B in combination with low R:FR enhances petiole elongation similar to vegetation shade, providing functional context for a low B response in plant competition. Low B potentiates the low R:FR response through PIF4, PIF5, and PIF7, and it involves increased PIF5 abundance and transcriptional changes. Low B attenuates a low R:FR-induced negative feedback loop through reduced gene expression of negative regulators and reduced HFR1 levels. The enhanced response to combined phytochrome and cryptochrome inactivation shows how multiple light cues can be integrated to fine-tune the plant's response to a changing environment.

Cereals and grasses adapt their structural development to environmental conditions and the resources available. The primary adaptive response is a variable degree of branching, called tillering in cereals. Especially for heterogeneous plant configurations the degree of tillering varies per plant. Functional-structural plant modelling (FSPM) is a modelling approach allowing simulation of the architectural development of individual plants, culminating in the emergent behaviour at the canopy level. This paper introduces the principles of modelling tillering in FSPM, using (I) a probability approach, forcing the dynamics of tillering to correspond to measured probabilities. Such models are particularly suitable to evaluate the effect structural variables on system performance. (II) ‘Dose response curves’, representing a measured or assumed response of tillering to an environmental cue. (III) Mechanistic approaches to tillering including control by carbohydrates, hormones and nutrients. Tiller senescence is equally important for the structural development of cereals as tiller appearance. Little study has been made of tiller senescence, though similar concepts seem to apply as for tiller appearance.

The laminae of leaves optimize photosynthetic rates by serving as a platform for both light capture and gas exchange, while minimizing water losses associated with thermoregulation and transpiration. Many have speculated that plants maximize photosynthetic output and minimize associated costs through leaf size, complexity, and shape, but a unifying theory linking the plethora of observed leaf forms with the environment remains elusive. Additionally, the leaf itself is a plastic structure, responsive to its surroundings, further complicating the relationship. Despite extensive knowledge of the genetic mechanisms underlying angiosperm leaf development, little is known about how phenotypic plasticity and selective pressures converge to create the diversity of leaf shapes and sizes across lineages. Here, we use wild tomato accessions, collected from locales with diverse levels of foliar shade, temperature, and precipitation, as a model to assay the extent of shade avoidance in leaf traits and the degree to which these leaf traits correlate with environmental factors. We find that leaf size is correlated with measures of foliar shade across the wild tomato species sampled and that leaf size and serration correlate in a species-dependent fashion with temperature and precipitation. We use far-red induced changes in leaf length as a proxy measure of the shade avoidance response, and find that shade avoidance in leaves negatively correlates with the level of foliar shade recorded at the point of origin of an accession. The direction and magnitude of these correlations varies across the leaf series, suggesting that heterochronic and/or ontogenic programs are a mechanism by which selective pressures can alter leaf size and form. This study highlights the value of wild tomato accessions for studies of both morphological and light-regulated development of compound leaves, and promises to be useful in the future identification of genes regulating potentially adaptive plastic leaf traits.

Shoot branching plays a pivotal role in the development of the aboveground plant structure. Therefore, to understand branching in relation to the environment, it is not only necessary to integrate the knowledge on mechanisms that regulate branching at multiple levels of biological organisation, but also to include plant structure explicitly. To this end, we propose the application of an established methodology called functional-structural plant modelling.

Intimate relationships exist between form and function of plants, determining many processes governing their growth and development. However, in most crop simulation models that have been created to simulate plant growth and, for example, predict biomass production, plant structure has been neglected. In this study, a detailed simulation model of growth and development of spring wheat (Triticum aestivum) is presented, which integrates degree of tillering and canopy architecture with organ-level light interception, photosynthesis, and dry-matter partitioning. An existing spatially explicit 3D architectural model of wheat development was extended with routines for organ-level microclimate, photosynthesis, assimilate distribution within the plant structure according to organ demands, and organ growth and development. Outgrowth of tiller buds was made dependent on the ratio between assimilate supply and demand of the plants. Organ-level photosynthesis, biomass production, and bud outgrowth were simulated satisfactorily. However, to improve crop simulation results more efforts are needed mechanistically to model other major plant physiological processes such as nitrogen uptake and distribution, tiller death, and leaf senescence. Nevertheless, the work presented here is a significant step forwards towards a mechanistic functional‐ structural plant model, which integrates plant architecture with key plant processes.

Article de revue (Article scientifique dans une revue à comité de lecture); International audience; Plants react to their environment and to management interventions by adjusting physiological functions and structure. Functional-structural plant models (FSPM), combine the representation of three-dimensional (3D) plant structure with selected physiological functions. An FSPM consists of an architectural part (plant structure) and a process part (plant functioning). The first deals with (i) the types of organs that are initiated and the way these are connected (topology), (ii) co-ordination in organ expansion dynamics, and (iii) geometrical variables (e.g. leaf angles, leaf curvature). The process part may include any physiological or physical process that affects plant growth and development (e.g. photosynthesis, carbon allocation). This paper addresses the following questions: (i) how are FSPM constructed, and (ii) for what purposes are they useful? Static, architectural models are distinguished from dynamic models. Static models are useful in order to study the significance of plant structure, such as light distribution in the canopy, gas exchange, remote sensing, pesticide spraying studies, and interactions between plants and biotic agents. Dynamic models serve quantitatively to integrate knowledge on plant functions and morphology as modulated by environment. Applications are in the domain of plant sciences, for example the study of plant plasticity as related to changes in the red:far red ratio of light in the canopy. With increasing availability of genetic information, FSPM will play a role in the assessment of the significance towards plant performance of variation in genetic traits across environments. In many crops, growers actively manipulate plant structure. FSPM is a promising tool to explore divergent management strategies.

Plant responses to the light spectrum under which plants are grown affect their developmental characteristics in a complicated manner. Lamps widely used to provide growth irradiance emit spectra which are very different from natural daylight spectra. Whereas specific responses of plants to a spectrum differing from natural daylight may sometimes be predictable, the overall plant response is generally difficult to predict due to the complicated interaction of the many different responses. So far studies on plant responses to spectra either use no daylight control or, if a natural daylight control is used, it will fluctuate in intensity and spectrum. An artificial solar (AS) spectrum which closely resembles a sunlight spectrum has been engineered, and growth, morphogenesis, and photosynthetic characteristics of cucumber plants grown for 13 d under this spectrum have been compared with their performance under fluorescent tubes (FTs) and a high pressure sodium lamp (HPS). The total dry weight of the AS-grown plants was 2.3 and 1.6 times greater than that of the FT and HPS plants, respectively, and the height of the AS plants was 4-5 times greater. This striking difference appeared to be related to a more efficient light interception by the AS plants, characterized by longer petioles, a greater leaf unfolding rate, and a lower investment in leaf mass relative to leaf area. Photosynthesis per leaf area was not greater for the AS plants. The extreme differences in plant response to the AS spectrum compared with the widely used protected cultivation light sources tested highlights the importance of a more natural spectrum, such as the AS spectrum, if the aim is to produce plants representative of field conditions.

The outgrowth of tiller buds in Poaceae is influenced by the ratio of red to far-red light irradiance (R:FR). At each point in the plant canopy, R:FR is affected by light scattered by surrounding plant tissues. This paper presents a three-dimensional virtual plant modelling approach to simulate local effects of R:FR on tillering in spring wheat (Triticum aestivum). ¿ R:FR dependence of bud outgrowth was implemented in a wheat model, using three hypothetical responses of bud extension to R:FR (unit step, curvilinear and linear response). Bud break occurred when a threshold bud length was reached. Simulations were performed for three plant population densities. ¿ In accordance with experimental observations, fewer tillers per plant were simulated for higher plant population densities. The linear and curvilinear responses caused a delay in the increase in tiller number compared with experimental data. The unit step response approached experimental results best. It is suggested that a model based on relatively simple relations can be used to simulate degree of tillering. ¿ This study has shown that the virtual plant approach is a promising tool with which to address crop morphological and ecological research questions where the determining factors act at the level of the individual plant organ.

International audience; In attempting to increase the reliability of clover contribution in clove/ryegrass systems it is important to understand the roles of (1) specific traits of the clover genotype (2) climate and (3) their interactions in determining clover behaviour in swards. Overwintering and spring growth of white clover (cultivars AberHerald and Huia) grown in binary mixtures with perennial ryegrass were measured at 12 European sites ranging in latitude from Reykjavik, Iceland (64 degrees 30' N) to Pordenone, Italy (46 degrees 30' N). In the overwintering period, tiller density of the grass was assessed and detailed morphological and chemical measurements were made on the clover at each sampling time. During the growing season, the clover contribution to total available biomass was recorded, Detailed climatic data were available at all sites. The annual growth cycle of swards was divided into four functional periods (spring, summer, autumn and winter). Within each functional period community responses were modelled. The models incorporated independent biotic variables characterizing each community within each site at the start of the period and independent variables characterizing the climate at each site during the period. The models were linked dynamically by taking, as response variable(s) for a functional period. the independent biotic variable(s) of the succeeding period. In general, the modelling strategy was successful in producing a series of biologically meaningful linked models. Essential prerequisites for this were (a) the establishment of a well-devised common protocol prior to the experiment and (b) the extensive gradients of climatic and other variables obtained by using numerous sites, AberHerald generally performed as well as. or better than, Hula throughout the annual cycle across the range of climatic conditions encountered, and especially under low temperature conditions in winter and autumn. Clover leaf area index appeared to be a key variable in determining clover performance over winter and through the following growing season. Grass tiller density had a strong negative effect on clover content in spring but only at low temperatures, This emphasizes the importance of a high clover leaf area index in autumn as the main biotic factor related to spring clover content in milder conditions. The importance of climatic variables in the models is their use in explaining the reliability of the contribution of clover in clover,: ryegrass systems. Temperature was the primary climatic determinant of clover response in all periods, having a direct effect on clover content and leaf area index or mediating the effect of the associate species. Radiation strongly influenced Clover dynamics during winter and spring but not in the other periods, possibly because it was confounded with the effect of higher temperature. Precipitation was positively related to clover growth during spring and autumn and was related to tiller density in a complex manner during autumn and winter.