Grain filling is a critical process for improving crop production under adverse conditions caused by climate change. Here, using a quantitative method, we quantified post-anthesis source–sink relationships of a large dataset to assess the contribution of remobilized pre-anthesis assimilates to grain growth for both biomass and nitrogen. The dataset came from 13 years of semi-controlled field experimentation, in which six bread wheat genotypes were grown at plot scale under contrasting temperature, water, and nitrogen regimes. On average, grain biomass was ~10% higher than post-anthesis above-ground biomass accumulation across regimes and genotypes. Overall, the estimated relative contribution (%) of remobilized assimilates to grain biomass became increasingly significant with increasing stress intensity, ranging from virtually nil to 100%. This percentage was altered more by water and nitrogen regimes than by temperature, indicating the greater impact of water or nitrogen regimes relative to high temperatures under our experimental conditions. Relationships between grain nitrogen demand and post-anthesis nitrogen uptake were generally insensitive to environmental conditions, as there was always significant remobilization of nitrogen from vegetative organs, which helped to stabilize the amount of grain nitrogen. Moreover, variations in the relative contribution of remobilized assimilates with environmental variables were genotype dependent. Our analysis provides an overall picture of post-anthesis source–sink relationships and pre-anthesis assimilate contributions to grain filling across (non-)environmental factors, and highlights that designing wheat adaptation to climate change should account for complex multifactor interactions.

Wheat grain yield is anticipated to suffer from the increased temperatures expected under climate change. In particular, the effects of post-anthesis temperatures on grain growth and development must be better understood to improve crop models. Grain growth and development involve several processes and we hypothesized that some of the most important processes, i.e. grain dry biomass and water accumulation, grain volume expansion and endosperm cell proliferation, will have different thermal sensitivity. To assess this, we established temperature response curves (TRC) of these processes for steady post-anthesis temperatures between 15°C and 36°C. From anthesis to maturity, grain dry mass, water mass, volume and endosperm cell number were monitored, whilst considering grain temperature. Different sensitivities to heat of these various processes were revealed. The rate of grain dry biomass accumulation increased linearly up to 25°C while the reciprocal of its duration linearly increased up to at least 32°C. By contrast, the growth rates of traits contributing to grain expansion, e.g. increase in grain volume and cell numbers, had higher optimum temperatures, while the reciprocal of their durations were significantly lower. These TRC can contribute to improve current crop models, and allow to target specific mechanisms for genetic and genomic studies.

Wheat is one of the most important crops in the world, mainly used for human consumption and animal feed. To overcome the increasing demand in wheat production, it is necessary to better understand the mechanisms involved in the growth of the wheat grain. X-ray computed tomography is an efficient method for the non-destructive investigation of the 3D architecture of biological specimens, which does not require staining, sectioning, or inclusion. In particular, phase-contrast tomography results in images with better contrast and an increased resolution compared to that obtained with laboratory tomography devices. The aim of this study was to investigate the potential of phase-contrast tomography for the study of the anatomy of the wheat grain at early stages of development. We provided 3D images of entire grains at various development stages. The image analysis allowed identifying a large number of tissues, and to visualize individual cells. Using a high-resolution setup, finer details were obtained, making it possible to identify additional tissues. Three-dimensional rendering of the grain also revealed the pattern resulting from the epidermis cells. X-ray phase-contrast tomography appears as a promising imaging method for the study of the 3D anatomy of plant organs and tissues.

Post-anthesis heat shocks, which are expected to increase in frequency under climate change, may affect wheat grain development and lead to significant decreases in grain yield. Grain development occurs in three phases, the lag-phase, the filling-phase, and maturation. The growth of the three main compartments of the grain (outer layers (OLs), endosperm, embryo) is staggered, so that heat shocks affect time- and tissue-specific growth processes differentially depending on their timing. We hypothesized that heat shocks during the lag-phase may reduce final grain size, resulting from a reduction in endosperm cell number and/or a restricted OLs growth. Plants were heated for four consecutive days during the lag-phase or the filling-phase or both phases (lag- and filling-). Heat shocks consisted in four hours a day at 38°C and 21°C for the rest of the day. Controlled plants were maintained at 21/14°C (day/night). For each temperature treatment, kinetics of whole grain and compartment masses and dimensions were measured as well as the endosperm cell number. An early heat shock reduced endosperm cell proliferation. However, the growth patterns neither of endosperm nor of OLs were modified compared to controls, resulting in no differences in final grain size. Furthermore, compared to controls, a single heat shock during the filling-phase reduced both the duration and rate of dry mass accumulation into grains, whereas two consecutive shocks reduced the duration but enhanced the rate of dry mass of accumulation, even when endosperm cell number was reduced. The mean endosperm cell size was shown to be larger after early heat shocks. All together, these results suggest a compensatory mechanism exists to regulate endosperm cell size and number. This process might be a new mechanistic target for molecular studies and would improve our understanding of post-anthesis wheat tolerance to heat-shocks.

Wheat (Triticum aestivum L.) is one of the most important crops as it provides 20% of calories and proteins to the human population. To overcome the increasing demand in wheat grain production, there is a need for a higher grain yield, and this can be achieved in particular through an increase in the grain weight. Moreover, grain shape is an important trait regarding the milling performance. Both the final grain weight and shape would benefit from a comprehensive knowledge of the morphological and anatomical determinism of wheat grain growth. Synchrotron-based phase-contrast X-ray microtomography (X-ray µCT) was used to study the 3D anatomy of the growing wheat grain during the first developmental stages. Coupled with 3D reconstruction, this method revealed changes in the grain shape and new cellular features. The study focused on a particular tissue, the pericarp, which has been hypothesized to be involved in the control of grain development. We showed considerable spatio-temporal diversity in cell shape and orientations, and in tissue porosity associated with stomata detection. These results highlight the growth-related features rarely studied in cereal grains, which may contribute significantly to the final grain weight and shape.