Robotized indoor phenotyping allows genomic prediction of adaptive traits in the field.

Breeding for resilience to climate change requires considering adaptive traits such as plant architecture, stomatal conductance and growth, beyond the current selection for yield. Robotized indoor phenotyping allows measuring such traits at high throughput for speed breeding, but is often considered as non-relevant for field conditions. Here, we show that maize adaptive traits can be inferred in different fields, based on genotypic values obtained indoor and on environmental conditions in each considered field. The modelling of environmental effects allows translation from indoor to fields, but also from one field to another field. Furthermore, genotypic values of considered traits match between indoor and field conditions. Genomic prediction results in adequate ranking of genotypes for the tested traits, although with lesser precision for elite varieties presenting reduced phenotypic variability. Hence, it distinguishes genotypes with high or low values for adaptive traits, conferring either spender or conservative strategies for water use under future climates.

Genetic regions determine tolerance to nitrogen deficiency in European elite bread wheats grown under contrasting nitrogen stress scenarios.

KEY MESSAGE: Clustering 24 environments in four contrasting nitrogen stress scenarios enabled the detection of genetic regions determining tolerance to nitrogen deficiency in European elite bread wheats. Increasing the nitrogen use efficiency of wheat varieties is an important goal for breeding. However, most genetic studies of wheat grown at different nitrogen levels in the field report significant interactions with the genotype. The chromosomal regions possibly involved in these interactions are largely unknown. The objective of this study was to quantify the response of elite bread wheat cultivars to different nitrogen field stress scenarios and identify genomic regions involved in this response. For this purpose, 212 elite bread wheat varieties were grown in a multi-environment trial at different nitrogen levels. Genomic regions associated with grain yield, protein concentration and grain protein deviation responses to nitrogen deficiency were identified. Environments were clustered according to adjusted means for grain yield, yield components and grain protein concentration. Four nitrogen availability scenarios were identified: optimal condition, moderate early deficiency, severe late deficiency, and severe continuous deficiency. A large range of tolerance to nitrogen deficiency was observed among varieties, which were ranked differently in different nitrogen deficiency scenarios. The well-known negative correlation between grain yield and grain protein concentration also existed between their respective tolerance indices. Interestingly, the tolerance indices for grain yield and grain protein deviation were either null or weakly positive meaning that breeding for the two traits should be less difficult than expected. Twenty-two QTL regions were identified for the tolerance indices. By selecting associated markers, these regions may be selected separately or combined to improve the tolerance to N deficiency within a breeding programme.

Physiological adaptive traits are a potential allele reservoir for maize genetic progress under challenging conditions.

Combined phenomic and genomic approaches are required to evaluate the margin of progress of breeding strategies. Here, we analyze 65 years of genetic progress in maize yield, which was similar (101 kg ha-1 year-1) across most frequent environmental scenarios in the European growing area. Yield gains were linked to physiologically simple traits (plant phenology and architecture) which indirectly affected reproductive development and light interception in all studied environments, marked by significant genomic signatures of selection. Conversely, studied physiological processes involved in stress adaptation remained phenotypically unchanged (e.g. stomatal conductance and growth sensitivity to drought) and showed no signatures of selection. By selecting for yield, breeders indirectly selected traits with stable effects on yield, but not physiological traits whose effects on yield can be positive or negative depending on environmental conditions. Because yield stability under climate change is desirable, novel breeding strategies may be needed for exploiting alleles governing physiological adaptive traits.

Genetic Analysis of Platform-Phenotyped Root System Architecture of Bread and Durum Wheat in Relation to Agronomic Traits.

Roots are essential for water and nutrient uptake but are rarely the direct target of breeding efforts. To characterize the genetic variability of wheat root architecture, the root and shoot traits of 200 durum and 715 bread wheat varieties were measured at a young stage on a high-throughput phenotyping platform. Heritability of platform traits ranged from 0.40 for root biomass in durum wheat to 0.82 for the number of tillers. Field phenotyping data for yield components and SNP genotyping were already available for all the genotypes. Taking differences in earliness into account, several significant correlations between root traits and field agronomic performances were found, suggesting that plants investing more resources in roots in some stressed environments favored water and nutrient uptake, with improved wheat yield. We identified 100 quantitative trait locus (QTLs) of root traits in the bread wheat panels and 34 in the durum wheat panel. Most colocalized with QTLs of traits measured in field conditions, including yield components and earliness for bread wheat, but only in a few environments. Stress and climatic indicators explained the differential effect of some platform QTLs on yield, which was positive, null, or negative depending on the environmental conditions. Modern breeding has led to deeper rooting but fewer seminal roots in bread wheat. The number of tillers has been increased in bread wheat, but decreased in durum wheat, and while the root-shoot ratio for bread wheat has remained stable, for durum wheat it has been increased. Breeding for root traits or designing ideotypes might help to maintain current yield while adapting to specific drought scenarios.

Grain carbon isotope composition is a marker for allocation and harvest index in wheat.

The natural 13 C abundance (δ13 C) in plant leaves has been used for decades with great success in agronomy to monitor water-use efficiency and select modern cultivars adapted to dry conditions. However, in wheat, it is also important to find genotypes with high carbon allocation to spikes and grains, and thus with a high harvest index (HI) and/or low carbon losses via respiration. Finding isotope-based markers of carbon partitioning to grains would be extremely useful since isotope analyses are inexpensive and can be performed routinely at high throughput. Here, we took the advantage of a set of field trials made of more than 600 plots with several wheat cultivars and measured agronomic parameters as well as δ13 C values in leaves and grains. We find a linear relationship between the apparent isotope discrimination between leaves and grain (denoted as Δδcorr ), and the respiration use efficiency-to-HI ratio. It means that overall, efficient carbon allocation to grains is associated with a small isotopic difference between leaves and grains. This effect is explained by postphotosynthetic isotope fractionations, and we show that this can be modelled by equations describing the carbon isotope composition in grains along the wheat growth cycle. Our results show that 13 C natural abundance in grains could be useful to find genotypes with better carbon allocation properties and assist current wheat breeding technologies.

Management and Characterization of Abiotic Stress via PhénoField®, a High-Throughput Field Phenotyping Platform.

In order to evaluate the impact of water deficit in field conditions, researchers or breeders must set up large experiment networks in very restrictive field environments. Experience shows that half of the field trials are not relevant because of climatic conditions that do not allow the stress scenario to be tested. The PhénoField® platform is the first field based infrastructure in the European Union to ensure protection against rainfall for a large number of plots, coupled with the non-invasive acquisition of crops' phenotype. In this paper, we will highlight the PhénoField® production capability using data from 2017-wheat trial. The innovative approach of the PhénoField® platform consists in the use of automatic irrigating rainout shelters coupled with high throughput field phenotyping to complete conventional phenotyping and micrometeorological densified measurements. Firstly, to test various abiotic stresses, automatic mobile rainout shelters allow fine management of fertilization or irrigation by driving daily the intensity and period of the application of the desired limiting factor on the evaluated crop. This management is based on micro-meteorological measurements coupled with a simulation of a carbon, water and nitrogen crop budget. Furthermore, as high-throughput plant-phenotyping under controlled conditions is well advanced, comparable evaluation in field conditions is enabled through phenotyping gantries equipped with various optical sensors. This approach, giving access to either similar or innovative variables compared manual measurements, is moreover distinguished by its capacity for dynamic analysis. Thus, the interactions between genotypes and the environment can be deciphered and better detailed since this gives access not only to the environmental data but also to plant responses to limiting hydric and nitrogen conditions. Further data analyses provide access to the curve parameters of various indicator kinetics, all the more integrative and relevant of plant behavior under stressful conditions. All these specificities of the PhénoField® platform open the way to the improvement of various categories of crop models, the fine characterization of variety behavior throughout the growth cycle and the evaluation of particular sensors better suited to a specific research question.

Using environmental clustering to identify specific drought tolerance QTLs in bread wheat (T. aestivum L.).

KEY MESSAGE: Environmental clustering helps to identify QTLs associated with grain yield in different water stress scenarios. These QTLs could be useful for breeders to improve grain yields and increase genetic resilience in marginal environments. Drought is one of the main abiotic stresses limiting winter bread wheat growth and productivity around the world. The acquisition of new high-yielding and stress-tolerant varieties is therefore necessary and requires improved understanding of the physiological and genetic bases of drought resistance. A panel of 210 elite European varieties was evaluated in 35 field trials. Grain yield and its components were scored in each trial. A crop model was then run with detailed climatic data and soil water status to assess the dynamics of water stress in each environment. Varieties were registered from 1992 to 2011, allowing us to test timewise genetic progress. Finally, a genome-wide association study (GWAS) was carried out using genotyping data from a 280 K SNP chip. The crop model simulation allowed us to group the environments into four water stress scenarios: an optimal condition with no water stress, a post-anthesis water stress, a moderate-anthesis water stress and a high pre-anthesis water stress. Compared to the optimal water condition, grain yield losses in the stressed conditions were 3.3%, 12.4% and 31.2%, respectively. This environmental clustering improved understanding of the effect of drought on grain yields and explained 20% of the G × E interaction. The greatest genetic progress was obtained in the optimal condition, mostly represented in France. The GWAS identified several QTLs, some of which were specific of the different water stress patterns. Our results make breeding for improved drought resistance to specific environmental scenarios easier and will facilitate genetic progress in future environments, i.e., water stress environments.

Functional mapping of N deficiency-induced response in wheat yield-component traits by implementing high-throughput phenotyping.

As overfertilization leads to environmental concerns and the cost of N fertilizer increases, the issue of how to select crop cultivars that can produce high yields on N-deficient soils has become crucially important. However, little information is known about the genetic mechanisms by which crops respond to environmental changes induced by N signaling. Here, we dissected the genetic architecture of N-induced phenotypic plasticity in bread wheat (Triticum aestivum L.) by integrating functional mapping and semiautomatic high-throughput phenotyping data of yield-related canopy architecture. We identified a set of quantitative trait loci (QTLs) that determined the pattern and magnitude of how wheat cultivars responded to low N stress from normal N supply throughout the wheat life cycle. This analysis highlighted the phenological landscape of genetic effects exerted by individual QTLs, as well as their interactions with N-induced signals and with canopy measurement angles. This information may shed light on our mechanistic understanding of plant adaptation and provide valuable information for the breeding of N-deficiency tolerant wheat varieties.

Applying association mapping and genomic selection to the dissection of key traits in elite European wheat.

KEY MESSAGE: We show the application of association mapping and genomic selection for key breeding targets using a large panel of elite winter wheat varieties and a large volume of agronomic data. The heightening urgency to increase wheat production in line with the needs of a growing population, and in the face of climatic uncertainty, mean new approaches, including association mapping (AM) and genomic selection (GS) need to be validated and applied in wheat breeding. Key adaptive responses are the cornerstone of regional breeding. There is evidence that new ideotypes for long-standing traits such as flowering time may be required. In order to detect targets for future marker-assisted improvement and validate the practical application of GS for wheat breeding we genotyped 376 elite wheat varieties with 3,046 DArT, single nucleotide polymorphism and gene markers and measured seven traits in replicated yield trials over 2 years in France, Germany and the UK. The scale of the phenotyping exceeds the breadth of previous AM and GS studies in these key economic wheat production regions of Northern Europe. Mixed-linear modelling (MLM) detected significant marker-trait associations across and within regions. Genomic prediction using elastic net gave low to high prediction accuracies depending on the trait, and could be experimentally increased by modifying the constituents of the training population (TP). We also tested the use of differentially penalised regression to integrate candidate gene and genome-wide markers to predict traits, demonstrating the validity and simplicity of this approach. Overall, our results suggest that whilst AM offers potential for application in both research and breeding, GS represents an exciting opportunity to select key traits, and that optimisation of the TP is crucial to its successful implementation.

A multi-environmental study of recent breeding progress on nitrogen use efficiency in wheat (Triticum aestivum L.).

KEY MESSAGE: By comparing 195 varieties in eight trials, this study assesses nitrogen use efficiency improvement in high and low nitrogen conditions in European winter wheat over the last 25 years. In a context where European agriculture practices have to deal with environmental concerns and nitrogen (N) fertiliser cost, nitrogen use efficiency (NUE) has to be improved. This study assessed genetic progress in winter wheat (Triticum aestivum L.) NUE. Two hundred and twenty-five European elite varieties were tested in four environments under two levels of N. Global genetic progress was assessed on additive genetic values and on genotype × N interaction, covering 25 years of European breeding. To avoid sampling bias, quality, precocity and plant height were added as covariates in the analyses when needed. Genotype × environment interactions were highly significant for all the traits studied to such an extent that no additive genetic effect was detected on N uptake. Genotype × N interactions were significant for yield, grain protein content (GPC), N concentration in straw, N utilisation, and NUE. Grain yield improvement (+0.45 % year(-1)) was independent of the N treatment. GPC was stable, thus grain nitrogen yield was improved (+0.39 % year(-1)). Genetic progress on N harvest index (+0.12 % year(-1)) and on N concentration in straw (-0.52 % year(-1)) possibly revealed improvement in N remobilisation. There has been an improvement of NUE additive genetic value (+0.33 % year(-1)) linked to better N utilisation (+0.20 % year(-1)). Improved yield stability was detected as a significant improvement of NUE in low compared to high N conditions. The application of these results to breeding programs is discussed.