Flowering time is an important trait in wheat breeding as it affects adaptation and yield potential. The aim of this study was to investigate the genetic architecture of flowering time in European winter bread wheat cultivars. To this end a population of 410 winter wheat varieties was evaluated in multi-location field trials and genotyped by a genotyping-by-sequencing approach and candidate gene markers. Our analyses revealed that the photoperiod regulator Ppd-D1 is the major factor affecting flowering time in this germplasm set, explaining 58% of the genotypic variance. Copy number variation at the Ppd-B1 locus was present but explains only 3.2% and thus a comparably small proportion of genotypic variance. By contrast, the plant height loci Rht-B1 and Rht-D1 had no effect on flowering time. The genome-wide scan identified six QTL which each explain only a small proportion of genotypic variance and in addition we identified a number of epistatic QTL, also with small effects. Taken together, our results show that flowering time in European winter bread wheat cultivars is mainly controlled by Ppd-D1 while the fine tuning to local climatic conditions is achieved through Ppd-B1 copy number variation and a larger number of QTL with small effects.
Genomic selection models can be trained using historical data and filtering genotypes based on phenotyping intensity and reliability criterion are able to increase the prediction ability. We implemented genomic selection based on a large commercial population incorporating 2325 European winter wheat lines. Our objectives were (1) to study whether modeling epistasis besides additive genetic effects results in enhancement on prediction ability of genomic selection, (2) to assess prediction ability when training population comprised historical or less-intensively phenotyped lines, and (3) to explore the prediction ability in subpopulations selected based on the reliability criterion. We found a 5 % increase in prediction ability when shifting from additive to additive plus epistatic effects models. In addition, only a marginal loss from 0.65 to 0.50 in accuracy was observed using the data collected from 1 year to predict genotypes of the following year, revealing that stable genomic selection models can be accurately calibrated to predict subsequent breeding stages. Moreover, prediction ability was maximized when the genotypes evaluated in a single location were excluded from the training set but subsequently decreased again when the phenotyping intensity was increased above two locations, suggesting that the update of the training population should be performed considering all the selected genotypes but excluding those evaluated in a single location. The genomic prediction ability was substantially higher in subpopulations selected based on the reliability criterion, indicating that phenotypic selection for highly reliable individuals could be directly replaced by applying genomic selection to them. We empirically conclude that there is a high potential to assist commercial wheat breeding programs employing genomic selection approaches.
Improvement of end-use quality in bread wheat (Triticum aestivum L.) depends on a thorough understanding of the genetic basis of important quality traits. The main goal of our study was to investigate the genetic basis of 1,000-kernel weight, protein content, sedimentation volume, test weight, and starch concentration using an association mapping approach. We fingerprinted 207 diverse European elite soft winter wheat lines with 115 SSR markers and evaluated the genotypes in multi-environment trials. The principal coordinate analysis revealed absence of a clear population but presence of a family structure. Therefore, we used linear mixed models and marker-based kinship matrices to correct for family structure. In genome-wide scans, we detected main effect QTL for all five traits. In contrast, epistatic QTL were only observed for sedimentation volume and test weight explaining a small proportion of the genotypic variation. Consequently, our findings suggested that integrating epistasis in marker-assisted breeding will not lead to substantially increased selection gain for quality traits in soft winter wheat.
Plant height variation in European winter wheat cultivars is mainly controlled by the Rht - D1 and Rht - B1 semi-dwarfing genes, but also by other medium- or small-effect QTL and potentially epistatic QTL enabling fine adjustments of plant height. Plant height is an important goal in wheat (Triticum aestivum L.) breeding as it affects crop performance and thus yield and quality. The aim of this study was to investigate the genetic control of plant height in European winter wheat cultivars. To this end, a panel of 410 winter wheat varieties from across Europe was evaluated for plant height in multi-location field trials and genotyped for the candidate loci Rht-B1, Rht-D1, Rht8, Ppd-B1 copy number variation and Ppd-D1 as well as by a genotyping-by-sequencing approach yielding 23,371 markers with known map position. We found that Rht-D1 and Rht-B1 had the largest effects on plant height in this cultivar collection explaining 40.9 and 15.5 % of the genotypic variance, respectively, while Ppd-D1 and Rht8 accounted for 3.0 and 2.0 % of the variance, respectively. A genome-wide scan for marker–trait associations yielded two additional medium-effect QTL located on chromosomes 6A and 5B explaining 11.0 and 5.7 % of the genotypic variance after the effects of the candidate loci were accounted for. In addition, we identified several small-effect QTL as well as epistatic QTL contributing to the genetic architecture of plant height. Taken together, our results show that the two Rht-1 semi-dwarfing genes are the major sources of variation in European winter wheat cultivars and that other small- or medium-effect QTL and potentially epistatic QTL enable fine adjustments in plant height.
In contrast to high-throughput genotyping which can manage a large number of plants at relatively low cost, phenotyping of many individual genotypes in field trials is still laborious and expensive. Early plant vigour, as an early selection criterion, is a trait that is visually scored due to a lack of suitable phenotyping methods for an accurate detection of this trait in large field trials. A high-throughput phenotyping technique for scoring early plant vigour would enhance the breeding process. This study was conducted to develop a method for scoring phenotypic differences in early plant vigour of 50 winter wheat ( L.) cultivars in a 2-years experiment using a vehicle based multispectral active sensor and two commercially available active sensors, GreenSeeker and CropCircle. Pixel analysis of RGB images revealed to be the most feasible and superior method compared to other possible reference methods. A comparison between the two years 2011 and 2012 confirmed that early plant vigour was affected by genotypic differences. A novel spectral plant vigour index (EPVI) was found to accurately reflect the plant vigour at tillering. Different methods were applied to identify optimal combinations of wavelengths to predict early plant vigour, including multivariate modelling and prediction, contour maps for identifying all possible simple ratios and testing of combined indices. The EPVI and the relative amount of green pixels (RAGP) derived from digital images were significantly related with = 0.98 to each other in both years. A total of 200 plots, 12 m in length, could be measured within 75 min. The EPVI was shown to be an accurate scoring method for the high-throughput screening of large field trials. The rapidity and accuracy of this novel method may contribute to enhanced selection at early growth stages.
The global distribution of croplands is of critical interest to a wide group of end-users. Different crops have their own representative phenological stages during their growing seasons, which differ considerably from other natural vegetation types. During the last decade, the Moderate Resolution Imaging Spectroradiometer (MODIS) has become a key tool for vegetation monitoring because of its high temporal resolution, extensive scope, and rapid availability of various products. However, mixed pixels caused by the moderate spatial resolution produce significant errors in crop area estimation. Here we propose a Crop Proportion Phenology Index (CPPI) to express the quantitative relationship between the MODIS vegetation index (VI) time series and winter wheat crop area. The utility of this index was tested in two experimental areas in China: one around Tongzhou and the other around Shuyang, as representative districts around a metropolis and a rural area, respectively. The CPPI performed well in these two regions, with the root mean square error (RMSE) in fractional crop area predictions ranging roughly from 15% in the individual pixels to 5% above 6.25 km . The training samples containing mixtures of crop types mitigated the challenges of pure end-member selection in a spectral mixture analysis. A small number of training samples are sufficient to generate the CPPI, which is adaptable to other crop types and larger regions. Estimating the CPPI parameters across larger spatial scales helped improve the stability of the model.
► We compared eight crop simulation models for winter wheat at eight sites in Europe. ► Input data was restricted to reflect typical use in large area impact studies. ► Wide ranges of simulated yields at all sites indicate high uncertainties. ► Multi-model mean yield predictions were in good agreement with observations. ► More suited and longer data series are needed for advancing model evaluation studies. We compared the performance of eight widely used, easily accessible and well-documented crop growth simulation models (APES, CROPSYST, DAISY, DSSAT, FASSET, HERMES, STICS and WOFOST) for winter wheat ( L.) during 49 growing seasons at eight sites in northwestern, Central and southeastern Europe. The aim was to examine how different process-based crop models perform at the field scale when provided with a limited set of information for model calibration and simulation, reflecting the typical use of models for large-scale applications, and to present the uncertainties related to this type of model application. Data used in the simulations consisted of daily weather statistics, information on soil properties, information on crop phenology for each cultivar, and basic crop and soil management information. Our results showed that none of the models perfectly reproduced recorded observations at all sites and in all years, and none could unequivocally be labelled robust and accurate in terms of yield prediction across different environments and crop cultivars with only minimum calibration. The best performance regarding yield estimation was for DAISY and DSSAT, for which the RMSE values were lowest (1428 and 1603 kg ha ) and the index of agreement (0.71 and 0.74) highest. CROPSYST systematically underestimated yields (MBE – 1186 kg ha ), whereas HERMES, STICS and WOFOST clearly overestimated them (MBE 1174, 1272 and 1213 kg ha , respectively). APES, DAISY, HERMES, STICS and WOFOST furnished high total above-ground biomass estimates, whereas CROPSYST, DSSAT and FASSET provided low total above-ground estimates. Consequently, DSSAT and FASSET produced very high harvest index values, followed by HERMES and WOFOST. APES and DAISY, on the other hand, returned low harvest index values. In spite of phenological observations being provided, the calibration results for wheat phenology, i.e. estimated dates of anthesis and maturity, were surprisingly variable, with the largest RMSE for anthesis being generated by APES (20.2 days) and for maturity by HERMES (12.6). The wide range of grain yield estimates provided by the models for all sites and years reflects substantial uncertainties in model estimates achieved with only minimum calibration. Mean predictions from the eight models, on the other hand, were in good agreement with measured data. This applies to both results across all sites and seasons as well as to prediction of observed yield variability at single sites – a very important finding that supports the use of multi-model estimates rather than reliance on single models.
The scale mismatch between remote sensing observations and state variables simulated by crop growth models decreases the reliability of crop yield estimates. To overcome this problem, we implemented a two-step data-assimilation approach: first, we generated a time series of 30-m-resolution leaf area index (LAI) by combining Moderate Resolution Imaging Spectroradiometer (MODIS) data and three Landsat TM images with a Kalman filter algorithm (the synthetic KF LAI series); second, the time series were assimilated into the WOFOST crop growth model to generate an ensemble Kalman filter LAI time series (the EnKF-assimilated LAI series). The synthetic EnKF LAI series then drove the WOFOST model to simulate winter wheat yields at 1-km resolution for pixels with wheat fractions of at least 50%. The county-level aggregated yield estimates were compared with official statistical yields. The synthetic KF LAI time series produced a more realistic characterization of LAI phenological dynamics. Assimilation of the synthetic KF LAI series produced more accurate estimates of regional winter wheat yield ( = 0.43; root-mean-square error (RMSE) = 439 kg ha ) than three other approaches: WOFOST without assimilation (determination coefficient = 0.14; RMSE = 647 kg ha ), assimilation of Landsat TM LAI ( = 0.37; RMSE = 472 kg ha ), and assimilation of S-G filtered MODIS LAI ( = 0.49; RMSE = 1355 kg ha ). Thus, assimilating the synthetic KF LAI series into the WOFOST model with the EnKF strategy provides a reliable and promising method for improving regional estimates of winter wheat yield.
Wheat is one of the most important cereals, whose growth and development is strongly limited by drought. This study investigated the physiological and metabolic response of six winter wheat cultivars to drought with the emphasis on the induction of dominant metabolites affected by the treatment and genotypes or both. The plants were exposed to a moderate (non-lethal) drought stress, which was induced by withholding watering for six days under controlled greenhouse conditions. A decline in CO2 assimilation (Pn) and transpiration rate, stomata closure, a decrease in relative water content (RWC) and increase of malondialdehyde content were observed in drought-treated plants of all cultivars. These changes were most pronounced in Ellvis, while Soissons was able to retain the higher RWC and Pn. Among the studied metabolites, sugars (sucrose, glucose, fructose, several disaccharides), organic acids (malic acid, oxalic acids), amino acids (proline, threonine, gamma-aminobutyric acid (GABA), glutamine) and sugar alcohols such as myo-inositol accumulated to higher levels in the plants exposed to drought stress in comparison with the control. The accumulation of several metabolites in response to drought differed between the genotypes. Drought induced the production of sucrose, malic acid and oxalic acid, unknown organic acid 1, unknown disaccharide 1, 2 and 3, GABA, L-threonine, glutamic acid in four (Soissons, Zitarka, Antonija or Toborzo) out of six genotypes. In addition, Soissons, which was the most drought tolerant genotype, accumulated the highest amount of unknown disaccharide 5, galactonic and phosphoric acids. The two most drought sensitive cultivars, Srpanjka and Ellvis, demonstrated different metabolic adjustment in response to the stress treatment. Srpanjka responded to drought by increasing the amount of glucose and fructose originated from hydrolyses of sucrose and accumulating unidentified sugar alcohols 1 and 2. In Ellvis, drought caused inhibition of photosynthetic carbon metabolism, as evidence by the decreased Pn, gs, RWC and accumulation levels of sugar metabolites (sucrose, glucose and fructose). The results revealed the differences in metabolic response to drought among the genotypes, which drew attention on metabolites related with general response and on those metabolites which are part of specific response that may play an important role in drought tolerance.
► Potential yield of winter wheat simulated with EPIC using weather data from 1960 to 2007 was 6.6–9.1 t ha in the North China Plain (NCP), generally increasing from north to south associated with decreasing low temperature stress. ► Based on county-level data from 2004 to 2007, the yield gap in the NCP was 2.7 t ha , 32% of the potential yield. ► Using 80% of the potential yield as an exploitable level, average actual wheat yields in the NCP could be increased by 1.0 t ha or 18%. ► Yield gaps at the county level were inversely correlated with actual yields ( = 0.82). The North China Plain (NCP) is the most important wheat production area in China, producing about two-thirds of China's total wheat output. To meet the associated increase in China's food demand with the expected growth in its already large population of 1.3 billion and diet changes, wheat production in the NCP needs to increase. Because of the farmland reduction due to urbanization, strategies for increasing wheat production in the NCP should be targeted at increasing current yields. To identify options for increasing wheat yields, we analyzed the yield potentials and yield gaps using the EPIC (Environment Policy Integrated Climate) model, Kriging interpolation techniques, GIS and average farm yields at county level. As most (ca. 82%) of the winter wheat in the NCP is irrigated, it is justified to use potential yield as the benchmark of the yield gap assessment. Wheat potential yields simulated with EPIC using daily weather data from 1960 to 2007 at 43 representative sites varied from 6.6 to 9.1 t ha in the NCP, generally increasing from north to south associated with decreasing low temperature stress. Based on the county-level data (2004–2007), the actual wheat yield varied between 2.4 and 7.7 t ha , while the yield gap was between 0.6 and 5.3 t ha (7–69% of the potential yield) across the NCP and decreased with increase of actual yields ( = 0.82). For the entire region, the weighted average actual yield was 5.7 t ha , while the yield gap was 2.7 t ha or 32% of the potential yield. Using 80% of the potential yield as an exploitable level, the average actual wheat yield in the NCP could be increased by 1.0 t ha or 18%. The results provide an indication of the possibilities to increase wheat yields in the NCP.