The Hydrogen Isotope Composition δ2H Reflects Plant Performance.

The stable carbon (δ13C) and oxygen (δ18O) isotope compositions in plant matter reflect photosynthetic and transpirative conditions in plants, respectively. However, the nature of hydrogen isotope composition (δ2H) and what it reflects of plant performance is poorly understood. Using durum wheat (Triticum turgidum var durum), this study evaluated the effect of different water and nitrogen growing field conditions on transpiration and how this effect influenced the performance of δ2H in autotrophic (flag leaf), mixotrophic (ears), and heterotrophic (grains and roots) organs. Moreover, δ2H was compared with the δ13C and δ18O in the same organs. Isotope compositions were analyzed in dry matter, the water-soluble fraction, and in water from different tissues of a set of genotypes. Similar to δ13C, the δ2H correlated negatively with stomatal conductance, whereas no correlation was observed for δ18O. Moreover, δ2H was not only affected by changes in transpiration but also by photosynthetic reactions, probably as a consequence of NADPH formation in autotrophic organs. Compared with the δ2H of stem water, plant δ2H was strongly diminished in photosynthetic organs such as the flag leaves, whereas it strongly increased in heterotrophic organs such as grains and roots. In heterotrophic organs, δ2H was associated with postphotosynthetic effects because there are several processes that lead to 2H-enrichment of carbohydrates. In summary, δ2H exhibited specific features that inform about the water conditions of the wheat crop, together with the photosynthetic characteristics of the plant part considered. Moreover, correlations of δ2H with grain yield illustrate that this isotope can be used to assess plant performance under different growing conditions.

Interactive effect of water and nitrogen regimes on plant growth, root traits and water status of old and modern durum wheat genotypes.

MAIN CONCLUSIONS: The selection of the ideal root drought adaptive traits should take into account the production and maintenance of root tissues alongside the capacity to capture soil resources. Ten old and modern Spanish durum wheat (Triticum turgidum L. var durum) genotypes were grown in lysimeters under two contrasting water and nitrogen regimes to study the effect of such growth conditions on: (1) the aerial biomass, (2) the growth and structure of the roots and (3) the relationships of the root structure with aerial biomass, photosynthetic and transpirative characteristics and water use efficiency. Both high water and nitrogen regimes significantly increased aerial biomass. Root dry biomass and root length increased and decreased in response to improved water supply and nitrogen regimes, respectively. No significant correlations were detected between aerial biomass and any root trait under well-watered conditions. Under water stress aerial biomass was negatively correlated with root dry biomass, root length and root weight density and positively correlated with the specific root length, particularly for the subset of old genotypes. The high nitrogen regime significantly enriched the carbon isotope composition of the flag leaf (δ (13)CFL) and hindered the effect of the high water regime on decreasing δ (13)CFL enrichment. Thus, positive correlations of aerial biomass with δ (13)CFL were detected regardless of the water regime. The study revealed: (1) the importance of root traits for higher aerial biomass under the low water regime; (2) that the interaction between nitrogen and the water regime may affect the predictive nature of the δ (13)C in drought breeding programs; and (3) the selection of the ideal root system structure should take into account the metabolic costs of the production and maintenance of root tissues alongside the capacity of capturing resources.

Contribution of the ear and the flag leaf to grain filling in durum wheat inferred from the carbon isotope signature: genotypic and growing conditions effects.

The ear, together with the flag leaf, is believed to play a major role as a source of assimilates during grain filling in C3 cereals. However, the intrusive nature of most of the available methodologies prevents reaching conclusive results in this regard. This study compares the carbon isotope composition (δ(13)C) in its natural abundance in the water-soluble fractions of the flag leaf blade and the ear with the δ(13)C of mature kernels to assess the relative contribution of both organs to grain filling in durum wheat (Triticum turgidum L. var. durum). The relative contribution of the ear was higher in landraces compared to modern cultivars, as well as in response to nitrogen fertilization and water stress. Such genotypic and environmentally driven differences were associated with changes in harvest index (HI), with the relative contribution of the ear being negatively associated with HI. In the case of the genotypic differences, the lower relative contribution of the ear in modern cultivars compared with landraces is probably associated with the appearance in the former of a certain amount of source limitation driven by a higher HI. In fact, the relative contribution of the ear was far more responsive to changes in HI in modern cultivars compared with landraces.

Physiological traits contributed to the recent increase in yield potential of winter wheat from Henan Province, China.

This experiment aims to test the traits responsible for the increase in yield potential of winter wheat released in Henan Province, China. Seven established cultivars released in the last 20 years and three advanced lines were assayed. The results showed that grain yield was positively correlated with harvest index (HI), kernel number per square meter, and aboveground biomass. In addition, the HI and aboveground biomass showed an increasing trend with the year of release. Therefore, we can conclude that bread wheat breeding advances during recent decades in Henan Province, China, have been achieved through an increase in HI, kernel number per square meter, and aboveground biomass. A higher δ(13)C seems also to be involved in these advances, which suggests a progressive improvement in constitutive water use efficiency not associated with a trend towards lower stomatal conductance in the most recent genotypes. However, genetic advance does not appear related to changes in photosynthesis rates on area basis when measured in the flag leaf or the spike, but only to a higher, whole-spike photosynthesis. Results also indirectly support the concept that under potential yield conditions, the spike contributed more than the flag leaf to kernel formation.

High-throughput phenotyping and genomic selection: the frontiers of crop breeding converge.

Genomic selection (GS) and high-throughput phenotyping have recently been captivating the interest of the crop breeding community from both the public and private sectors world-wide. Both approaches promise to revolutionize the prediction of complex traits, including growth, yield and adaptation to stress. Whereas high-throughput phenotyping may help to improve understanding of crop physiology, most powerful techniques for high-throughput field phenotyping are empirical rather than analytical and comparable to genomic selection. Despite the fact that the two methodological approaches represent the extremes of what is understood as the breeding process (phenotype versus genome), they both consider the targeted traits (e.g. grain yield, growth, phenology, plant adaptation to stress) as a black box instead of dissecting them as a set of secondary traits (i.e. physiological) putatively related to the target trait. Both GS and high-throughput phenotyping have in common their empirical approach enabling breeders to use genome profile or phenotype without understanding the underlying biology. This short review discusses the main aspects of both approaches and focuses on the case of genomic selection of maize flowering traits and near-infrared spectroscopy (NIRS) and plant spectral reflectance as high-throughput field phenotyping methods for complex traits such as crop growth and yield.

Remote sensing techniques and stable isotopes as phenotyping tools to assess wheat yield performance: Effects of growing temperature and vernalization.

This study compares distinct phenotypic approaches to assess wheat performance under different growing temperatures and vernalization needs. A set of 38 (winter and facultative) wheat cultivars were planted in Valladolid (Spain) under irrigation and two contrasting planting dates: normal (late autumn), and late (late winter). The late plating trial exhibited a 1.5 °C increase in average crop temperature. Measurements with different remote sensing techniques were performed at heading and grain filling, as well as carbon isotope composition (δ13C) and nitrogen content analysis. Multispectral and RGB vegetation indices and canopy temperature related better to grain yield (GY) across the whole set of genotypes in the normal compared with the late planting, with indices (such as the RGB indices Hue, a* and the spectral indices NDVI, EVI and CCI) measured at grain filling performing the best. Aerially assessed remote sensing indices only performed better than ground-acquired ones at heading. Nitrogen content and δ13C correlated with GY at both planting dates. Correlations within winter and facultative genotypes were much weaker, particularly in the facultative subset. For both planting dates, the best GY prediction models were achieved when combining remote sensing indices with δ13C and nitrogen of mature grains. Implications for phenotyping in the context of increasing temperatures are further discussed.

New avenues for increasing yield and stability in C3 cereals: exploring ear photosynthesis.

Small grain cereals such as wheat, rice and barley are among the most important crops worldwide. Any attempt to increase crop productivity and stability through breeding implies developing new strategies for plant phenotyping, including defining ideotype attributes for selection. Recently, the role of non-foliar photosynthetic organs, particularly the inflorescences, has received increasing attention. For example, ear photosynthesis has been reported to be a major contributor to grain filling in wheat and barley under stress and good agronomic conditions. This review provides an overview of the particular characteristics of the ear that makes this photosynthetic organ better adapted to grain filling than the flag leaf and revises potential metabolic and molecular traits that merit further research as targets for cereal improvement. Currently, the absence of high-throughput phenotyping methods limits the inclusion of ear photosynthesis in the breeding agenda. In this regard, a number of different approaches are presented.

Cereal Crop Ear Counting in Field Conditions Using Zenithal RGB Images.

Ear density, or the number of ears per square meter (ears/m2), is a central focus in many cereal crop breeding programs, such as wheat and barley, representing an important agronomic yield component for estimating grain yield. Therefore, a quick, efficient, and standardized technique for assessing ear density would aid in improving agricultural management, providing improvements in preharvest yield predictions, or could even be used as a tool for crop breeding when it has been defined as a trait of importance. Not only are the current techniques for manual ear density assessments laborious and time-consuming, but they are also without any official standardized protocol, whether by linear meter, area quadrant, or an extrapolation based on plant ear density and plant counts postharvest. An automatic ear counting algorithm is presented in detail for estimating ear density with only sunlight illumination in field conditions based on zenithal (nadir) natural color (red, green, and blue [RGB]) digital images, allowing for high-throughput standardized measurements. Different field trials of durum wheat and barley distributed geographically across Spain during the 2014/2015 and 2015/2016 crop seasons in irrigated and rainfed trials were used to provide representative results. The three-phase protocol includes crop growth stage and field condition planning, image capture guidelines, and a computer algorithm of three steps: (i) a Laplacian frequency filter to remove low- and high-frequency artifacts, (ii) a median filter to reduce high noise, and (iii) segmentation and counting using local maxima peaks for the final count. Minor adjustments to the algorithm code must be made corresponding to the camera resolution, focal length, and distance between the camera and the crop canopy. The results demonstrate a high success rate (higher than 90%) and R2 values (of 0.62-0.75) between the algorithm counts and the manual image-based ear counts for both durum wheat and barley.

Comparative effect of salinity on growth, grain yield, water use efficiency, δ(13)C and δ(15)N of landraces and improved durum wheat varieties.

Supplemental irrigation with low-quality water will be paramount in Mediterranean agriculture in the future, where durum wheat is a major crop. Breeding for salinity tolerance may contribute towards improving resilience to irrigation with brackish water. However, identification of appropriate phenotyping traits remains a bottleneck in breeding. A set of 25 genotypes, including 19 landraces and 6 improved varieties most cultivated in Tunisia, were grown in the field and irrigated with brackish water (6, 13 and 18dSm(-1)). Improved genotypes exhibited higher grain yield (GY) and water use efficiency at the crop level (WUEyield or 'water productivity'), shorter days to flowering (DTF), lower N concentration (N) and carbon isotope composition (δ(13)C) in mature kernels and lower nitrogen isotope composition (δ(15)N) in the flag leaf compared with landraces. GY was negatively correlated with DTF and the δ(13)C and N of mature kernels and was positively correlated with the δ(15)N of the flag leaf. Moreover, δ(13)C of mature kernels was negatively correlated with WUEyield. The results highlight the importance of shorter phenology together with photosynthetic resilience to salt-induced water stress (lower δ(13)C) and nitrogen metabolism (higher N and δ(15)N) for assessing genotypic performance to salinity.