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There is a need to generate improved crop varieties adapted to the ongoing changes in the climate. We studied durum wheat canopy and central metabolism of six different photosynthetic organs in two yield-contrasting varieties. The aim was to understand the mechanisms associated with the water stress response and yield performance. Water stress strongly reduced grain yield, plant biomass, and leaf photosynthesis, and down-regulated C/N-metabolism genes and key protein levels, which occurred mainly in leaf blades. By contrast, higher yield was associated with high ear dry weight and lower biomass and ears per area, highlighting the advantage of reduced tillering and the consequent improvement in sink strength, which promoted C/N metabolism at the whole plant level. An improved C metabolism in blades and ear bracts and N assimilation in all photosynthetic organs facilitated C/N remobilization to the grain and promoted yield. Therefore, we propose that further yield gains in Mediterranean conditions could be achieved by considering the source-sink dynamics and the contribution of non-foliar organs, and particularly N assimilation and remobilization during the late growth stages. We highlight the power of linking phenotyping with plant metabolism to identify novel traits at the whole plant level to support breeding programmes., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Contribution of inflorescences to seed filling have attracted great attention given the resilience of this photosynthetic organ to stressful conditions. However, studies have been almost exclusively focused to small grain cereals. In this study, we aimed to explore these responses in quinoa, as a climate resilient seed crop of elevated economic and nutritious potential. We compared the physiological and metabolic performance of panicles and leaves of two quinoa cultivars growing under contrasting salinity levels. Plant growth, photosynthetic and transpiratory gas exchange and chlorophyll fluorescence were monitored in inflorescences and leaves throughout the experiment. At flowering stage, young and mature leaves and panicles were sampled for key metabolic markers related to carbon, nitrogen and secondary metabolisms. When subjected to salt stress, panicles showed attenuated declines on photosynthesis, water use, pigments, amino acids, and protein levels as compared to leaves. In fact, the assimilation rates, together with a high hexose content evidenced an active photosynthetic role of the panicle under optimal and salt stress conditions. Moreover, we also found significant genotypic variability for physiological and metabolic traits of panicles and leaves, which emphasizes the study of genotype-dependent stress responses at the whole plant level. We conclude that quinoa panicles are less affected by salt stress than leaves, which encourages further research and exploitation of this organ for crop improvement and stress resilience considering the high natural diversity., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier GmbH.. All rights reserved.)

High-throughput crop phenotyping, particularly under field conditions, is nowadays perceived as a key factor limiting crop genetic advance. Phenotyping not only facilitates conventional breeding, but it is necessary to fully exploit the capabilities of molecular breeding, and it can be exploited to predict breeding targets for the years ahead at the regional level through more advanced simulation models and decision support systems. In terms of phenotyping, it is necessary to determined which selection traits are relevant in each situation, and which phenotyping tools/methods are available to assess such traits. Remote sensing methodologies are currently the most popular approaches, even when lab-based analyses are still relevant in many circumstances. On top of that, data processing and automation, together with machine learning/deep learning are contributing to the wide range of applications for phenotyping. This review addresses spectral and red-green-blue sensing as the most popular remote sensing approaches, alongside stable isotope composition as an example of a lab-based tool, and root phenotyping, which represents one of the frontiers for field phenotyping. Further, we consider the two most promising forms of aerial platforms (unmanned aerial vehicle and satellites) and some of the emerging data-processing techniques. The review includes three Boxes that examine specific case studies., (© 2021 Institute of Botany, Chinese Academy of Sciences.)

Ideotypic characteristics of durum wheat associated with higher yield under different water and temperature regimes were studied under Mediterranean conditions. The focus of this paper is to provide raw and supplemental data from the research article entitled "Durum wheat ideotypes in Mediterranean environments differing in water and temperature conditions" [1], which aims to define specific durum wheat ideotypes according to their responses to different agronomic conditions. In this context, six modern (i.e. post green revolution) genotypes with contrasting yield performance (i.e. high vs low yield) were grown during two consecutive years under different treatments: (i) winter planting under support-irrigation conditions, (ii) winter planting under rainfed conditions, (iii) late planting under support-irrigation. Trials were conducted at the INIA station of Colmenar de Oreja (Madrid). Different traits were assessed to inform about water status (canopy temperature at anthesis and stable carbon isotope composition (δ 13 C) of the flag leaf and mature grains), root performance (root traits and the oxygen isotope composition (δ 18 O) in the stem base water), phenology (days from sowing to heading), nitrogen status/photosynthetic capacity (nitrogen content and stable isotope composition (δ 15 N) of the flag leaf and mature grain together with the pigment contents and the nitrogen balance index (NBI) of the flag leaf), crop growth (plant height (PH) and the normalized difference vegetation index (NDVI) at anthesis), grain yield and agronomic yield components. For most of the parameters assessed, data analysis demonstrated significant differences among genotypes within each treatment. The level of significance was determined using the Tukey-b test on independent samples, and ideotypes were modelled from the results of principle component analysis. The present data shed light on traits that help to define specific ideotype characteristics that confer genotypic adaptation to a wide range of agronomic conditions produced by variations in planting date, water conditions and season., Competing Interests: The authors declare that they have no known competing financial interests or personal relationships that have, or could be perceived to have, influenced the work reported in this article., (© 2021 Published by Elsevier Inc.)

Durum wheat production is seriously threatened by Fusarium head blight (FHB) attacks in Tunisia, and the seed coating by bio-agents is a great alternative for chemical disease control. This study focuses on evaluating, under field conditions, the effect of seed coating with Trichoderma harzianum , Meyerozyma guilliermondii and their combination on (i) FHB severity, durum wheat grain yield and TKW in three crop seasons, and (ii) on physiological parameters and the carbon and nitrogen content and isotope composition in leaves and grains of durum wheat. The results indicated that the treatments were effective in reducing FHB severity by 30 to 70% and increasing grain yield with an increased rate ranging from 25 to 68%, compared to the inoculated control. The impact of treatments on grain yield improvement was associated with higher NDVI and chlorophyll content and lower canopy temperature. Furthermore, the treatments mitigated the FHB adverse effects on N and C metabolism by resulting in a higher δ 13 Cgrain ( 13 C/ 12 Cgrain) and δ 15 Ngrain ( 15 N/ 14 Ngrain). Overall, the combination outperformed the other seed treatments by producing the highest grain yield and TKW. The high potency of seed coating with the combination suggests that the two microorganisms have synergetic or complementary impacts on wheat.

The euhalophyte species Salicornia europaea is cultivated for oilseed and as a fodder crop in various parts of the world. In saline coastal environments it possesses great potential for the subsistence of the most disadvantaged farmers. We investigated the effect of salinity levels in irrigation water on the germination capacity, shoot biomass and seed productivity as well as diverse quality traits (nitrogen content in shoots and seeds and fatty acids, in seeds) and physiological traits (stable carbon and nitrogen isotopes and ion content) of two accessions collected in the United Arab Emirates (UAE). The three salinity levels tested were irrigation with fresh water (0.3 dS m -1 ), brackish water (25 dS m -1 ) and sea water (40 dS m -1 ). In addition, a hypersaline condition (80 dS m -1 ) was also tested for germination. The best germination rates were achieved with seeds exposed to fresh and brackish water, while imbibition with sea water decreased germination by half and hypersaline water inhibited it almost totally. However, the best irrigation regime in terms of biomass and seed yield involved brackish water. Moreover, rising salinity in the irrigation increased the stable isotope composition of carbon (δ 13 C) and nitrogen (δ 15 N), together with the Na + and K + of shoots and seeds, and the lipid levels of seeds, while the total nitrogen content and the profile of major fatty acids of seeds did not change. Differences between the two ecotypes existed for growth and seed yield with the best ecotype exhibiting lower δ 13 C and higher K + in both shoots and seeds, lower Na + and higher δ 15 N in shoots, and lower N in seeds, together with differences in major fatty acids. Physiological mechanisms behind the response to irrigation salinity and the ecotypic differences are discussed in terms of photosynthetic carbon and nitrogen metabolism., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Coating seeds with bio-control agents is a potentially effective approach to reduce the usage of pesticides and fertilizers applied and protect the natural environment. This study evaluated the effect of seed coating with Meyerozyma guilliermondii , strain INAT (MT731365), on seed germination, plant growth and photosynthesis, and plant resistance against Fusarium culmorum , in durum wheat under controlled conditions. Compared to control plants, seed coating with M. guilliermondii promoted the wheat growth (shoot and roots length and biomass), and photosynthesis and transpiration traits (chlorophyll, ɸPSII, rates of photosynthesis and transpiration, etc.) together with higher nitrogen balance index (NBI) and lower flavonols and anthocyanins. At 21 days post infection with Fusarium, M. guilliermondii was found to reduce the disease incidence and the severity, with reduction rates reaching up to 31.2% and 30.4%, respectively, as well as to alleviate the disease damaging impact on photosynthesis and plant growth. This was associated with lower ABA, flavonols and anthocyanins, compared to infected control. A pivotal function of M. guilliermondii as an antagonist of F. culmorum and a growth promoter is discussed.

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., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Knowledge of the agronomic and physiological traits associated with genetic gains in yield is essential to improve understanding of yield-limiting factors and to inform future breeding strategies. The aim of this paper is to dissect the agronomic and physiological traits related to genetic gain and to propose an ideotype with high yield that is best adapted to Spanish Mediterranean environments. Six semi-dwarf (i.e. modern) durum wheat genotypes were grown in a wide range of growing conditions in Spain during two successive years. Diverse agronomic, physiological and leaf morphological traits were evaluated. Kernels spike -1 was the yield component most affected by the genetic gain. While no interaction between genotype and growing conditions existed for grain yield, the more productive genotypes were characterized by a plant height of around 85 cm, small erect flag leaves, more open stomata, a better balance between N sources and N sinks and a higher capacity to re-fix CO 2 respired by the grain. Moreover, in general the non-laminar parts of the plants play a key role in providing assimilates during grain filling. The high heritability of most of the studied parameters allows their consideration as traits for phenotyping durum wheat better adapted to a wide range of Mediterranean conditions., (Copyright © 2019 Elsevier B.V. All rights reserved.)

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 (δ 13 C) 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 δ 13 C 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 δ 13 C and nitrogen of mature grains. Implications for phenotyping in the context of increasing temperatures are further discussed., (Copyright © 2019 Elsevier B.V. All rights reserved.)

The importance of quinoa has been emphasized considerably in the recent decades, as a highly nutritional crop seed that is tolerant to salinity and amenable to arid agronomical conditions. The focus of this paper is to provide raw and a supplemental data of the research article entitled "Agronomic performance of irrigated quinoa in desert areas: comparing different approaches for early assessment of salinity stress" [1], aiming to compare different approaches for early detection, at the genotypic and crop levels, of the effect of salinity caused by irrigation on the agronomic performance of this crop. A set of 20 genotypes was grown under drip irrigation in sandy soil, amended with manure, at the International Center for Biosaline Agriculture (UAE) for two weeks, after which half of the trial was submitted to irrigation with saline water and this was continued until crop maturity. After eight weeks of applying the two irrigation regimes, pigment contents were evaluated in fully expanded leaves. The same leaves were then harvested, dried and the stable carbon and nitrogen isotope compositions (δ 13 C and δ 15 N) and the total nitrogen and carbon contents of the dry matter analyzed, together with ion concentrations. At maturity yield components were assessed and yield harvested. Data analysis demonstrated significant differences in genotypes response under each treatment, within all assessed parameters. The significant level was provided using the Tukey-b test on independent samples. The present dataset highlights the potential use of different approaches to crop phenotyping and monitoring decision making., Competing Interests: The authors declare that they have no known competing financial interests or personal relationships that have, or could be perceived to have, influenced the work reported in this article., (© 2020 The Author(s).)

Coating seeds with biostimulants is among the promising approaches in crop production to increase crop tolerance to drought stress. In this study, we evaluated the potential of coating durum wheat seeds of the cultivar 'Karim' with thyme essential oil on enhancing seed germination and seedling growth, and on plant growth promotion and induction of drought resistance. Coated seeds were pre-germinated, grown in hydroponics, and grown in pots under controlled well-watered and progressive water/nutrient stress conditions. Seed coating with thyme oil increased germination rate and enhanced seedling growth development in hydroponics. In the pot experiment, thyme oil increased, when well watered, root and shoot development, chlorophyll, nitrogen balance index (NBI), abscisic acid (ABA), anthocyanins and flavonoids in leaves, decreased nitrogen isotope composition (δ 15 N) and increased carbon isotope composition (δ 13 C) of shoots. Increasing water/nutrient stress in control plants induced higher accumulation of ABA and anthocyanins coupled with a transient decrease in chlorophyll and NBI, a decrease in shoot and root development, the Normalized Difference Vegetation Index (NDVI), shoot C content, δ 15 N, and an increase in δ 13 C, revealing the avoidance strategy adopted by the cultivar. Thyme oil had the potential to enhance the avoidance strategy by inducing roots elongation, reducing the loss of shoot and roots dry matter and chlorophyll, maintaining balanced NBI, an decreasing anthocyanins, flavonoids, and δ 13 C via maintaining lower ABA-mediated-stomatal closure. Thyme oil increased shoot N content and δ 15 N indicating preferential uptake of the 15 N enriched NH 4 + . Coating seeds with thyme oil is suggested as a promising alternative approach to improve plant's water and nutrient status and to enhance drought resistance.

The stable carbon (δ 13 C) and oxygen (δ 18 O) isotope compositions in plant matter reflect photosynthetic and transpirative conditions in plants, respectively. However, the nature of hydrogen isotope composition (δ 2 H) 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 δ 2 H in autotrophic (flag leaf), mixotrophic (ears), and heterotrophic (grains and roots) organs. Moreover, δ 2 H was compared with the δ 13 C and δ 18 O 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 δ 13 C, the δ 2 H correlated negatively with stomatal conductance, whereas no correlation was observed for δ 18 O. Moreover, δ 2 H 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 δ 2 H of stem water, plant δ 2 H 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, δ 2 H was associated with postphotosynthetic effects because there are several processes that lead to 2 H-enrichment of carbohydrates. In summary, δ 2 H 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 δ 2 H with grain yield illustrate that this isotope can be used to assess plant performance under different growing conditions., (© 2019 American Society of Plant Biologists. All Rights Reserved.)

Much attention has been paid to understanding the traits associated with crop performance and the associated underlying physiological mechanisms, with less effort done towards combining different plant scales, levels of observation, or including hybrids of autogamous species. We aim to identify mechanisms at canopy, leaf and transcript levels contributing to crop performance under contrasting nitrogen supplies in three barley genotypes, two hybrids and one commercial line. High nitrogen fertilization did not affect photosynthetic capacity on a leaf area basis and lowered nitrogen partial factor productivity past a certain point, but increased leaf area and biomass accumulation, parameters that were closely tracked using various different high throughput remote sensing based phenotyping techniques. These aspects, together with a larger catabolism of leaf nitrogen compounds amenable to sink translocation, contributed to higher crop production. Better crop yield and growth in hybrids compared to the line was linked to a nitrogen-saving strategy in source leaves to the detriment of larger sink size, as indicated by the lower leaf nitrogen content and downregulation of nitrogen metabolism and aquaporin genes. While these changes did not reduce photosynthesis capacity on an area basis, they were related with better nitrogen use in the hybrids compared with the line., (Copyright © 2018. Published by Elsevier B.V.)

Ear density, or the number of ears per square meter (ears/m 2 ), 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 R 2 values (of 0.62-0.75) between the algorithm counts and the manual image-based ear counts for both durum wheat and barley.

Knowledge of the function of the ear as a key organ in the uptake, remobilization and partitioning of nitrogen is essential for understanding its contribution to grain filling and thus guiding future breeding strategies. In this work, four Chinese winter wheat genotypes were grown on a 15 N-enriched nutrient solution. N absorption and further remobilization to the flag leaf, the ear and the mature grains were calculated via the 15 N atom % excess. The results indicated that the high yields of the Chinese wheat genotype were determined by higher grain numbers per ear, with greater plant height and a larger ear size, while the thousand-grain weight did not affect grain yield. In the mature grains, 66.7% of total N was remobilized from the pre-anthesis accumulation in the biomass, while the remaining 33.3% was derived from the N taken up during post-anthesis. From anthesis to 2 weeks after the anthesis stage, the flag leaf remobilized 3.67 mg of N outwards and the ear remobilized 3.87 mg of N inwards from the pre-anthesis accumulation in each plant. The positive correlation between ear N rem and grain N rem indicated that the ear was an important organ for providing N to the grain, whereas the remobilized N stream from the leaves was not correlated with grain N rem , thus indicating that flag leaf N was not translocated directly to the grain. The grain N rem was negatively correlated with the ear N concentration throughout grain filling, which suggested that higher-yielding genotypes had better sink activity in the ear, while Rubisco played a critical role in N deposition. Therefore, to improve yield potential in wheat, the N accumulation in the ear and the subsequent remobilization of that stored N to the grains should be considered. N accumulation and remobilization in the ear may at least be valuable for Chinese breeding programs that aim at optimizing the sink/source balance to improve grain filling.

Sweet pepper is among the most widely cultivated horticultural crops in the Mediterranean basin, being frequently grown hydroponically under cover in combination with CO 2 fertilization and water conditions ranging from optimal to suboptimal. The aim of this study is to develop a simple model, based on the analysis of plant stable isotopes in their natural abundance, gas exchange traits and N concentration, to assess sweet pepper growth. Plants were grown in a growth chamber for near 6 weeks. Two [CO 2 ] (400 and 800 μmol mol -1 ), three water regimes (control and mild and moderate water stress) and four genotypes were assayed. For each combination of genotype, [CO 2 ] and water regime five plants were evaluated. Water stress applied caused significant decreases in water potential, net assimilation, stomatal conductance, intercellular to atmospheric [CO 2 ], and significant increases in water use efficiency, leaf chlorophyll content and carbon isotope composition, while the relative water content, the osmotic potential and the content of anthocyanins did change not under stress compared to control conditions support this statement. Nevertheless, water regime affects plant growth via nitrogen assimilation, which is associated with the transpiration stream, particularly at high [CO 2 ], while the lower N concentration caused by rising [CO 2 ] is not associated with stomatal closure. The stable isotope composition of carbon, oxygen, and nitrogen (δ 13 C, δ 18 O, and δ 15 N) in plant matter are affected not only by water regime but also by rising [CO 2 ]. Thus, δ 18 O increased probably as response to decreases in transpiration, while the increase in δ 15 N may reflect not only a lower stomatal conductance but a higher nitrogen demand in leaves or shifts in nitrogen metabolism associated with decreases in photorespiration. The way that δ 13 C explains differences in plant growth across water regimes within a given [CO 2 ], seems to be mediated through its direct relationship with N accumulation in leaves. The changes in the profile and amount of amino acids caused by water stress and high [CO 2 ] support this conclusion. However, the results do not support the use of δ 18 O as an indicator of the effect of water regime on plant growth.

The flag leaf has been traditionally considered as the main contributor to grain nitrogen. However, during the reproductive stage, other organs besides the flag leaf may supply nitrogen to developing grains. Therefore, the contribution of the ear and other organs to the nitrogen supplied to the growing grains remains unclear. It is important to develop phenotypic tools to assess the relative contribution of different plant parts to the N accumulated in the grains of wheat which may helps to develop genotypes that use N more efficiently. We studied the effect of growing conditions (different levels of water and nitrogen in the field) on the nitrogen contribution of the spike and different vegetative organs of the plant to the grains. The natural abundance of δ 15 N and total N content in the flag blade, peduncle, whole spike, glumes and awns were compared to the δ 15 N and total N in mature grains to trace the origin of nitrogen redistribution to the grains. The δ 15 N and total N content of the different plant parts correlated positively with the δ 15 N and total N content of mature grains suggesting that all organs may contribute a portion of their N content to the grains. The potential contribution of the flag blade to grain N increased (by 46%) as the growing conditions improved, whereas the potential contribution of the glumes plus awns and the peduncle increased (46 and 31%, respectively) as water and nitrogen stress increased. In general, potential contribution of the ear providing N to growing grains was similar (42%) than that of the vegetative parts of the plants (30-40%), regardless of the growing conditions. Thus, the potential ear N content could be a positive trait for plant phenotyping, especially under water and nitrogen limiting conditions. In that sense, genotypic variability existed at least between old (tall) and modern (semidwarf) cultivars, with the ear from modern genotypes exhibiting less relative contribution to the total grain N. The combined use of δ 15 N and N content may be used as an affordable tool to assess the relative contribution of different plant parts to the grain N in wheat.

The role of wheat ears as a source of nitrogen (N) and carbon (C) in the grain filling process has barely been studied. To resolve this question, five wheat genotypes were labeled with 15 N-enriched nutrient solution. N remobilization and absorption were estimated via the nitrogen isotope composition of total organic matter and Rubisco. Gas exchange analyses showed that ear photosynthesis contributed substantially to grain filling in spite of the great loss of C due to respiration. Of the total kernel N, 64.7% was derived from the N acquired between sowing and anthesis, while the remaining 35.3% was derived from the N acquired between anthesis and maturity. In addition, 1.87 times more N was remobilized to the developing kernel from the ear than from the flag leaf. The higher yielding genotypes showed an increased N remobilization to the kernel compared to the lower yielding genotypes. In addition, the higher yielding genotypes remobilized more N from the ears to the kernel than the lower yielding genotypes, while the lower yielding genotypes remobilized more N from the flag leaf to the kernel. Therefore, the ears contribute significantly toward fulfilling C and N demands during grain filling., (© 2016 Institute of Botany, Chinese Academy of Sciences.)

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., (Copyright © 2016 Elsevier Ireland Ltd. All rights reserved.)

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.

Elucidating the relationships between gene expression and the physiological mechanisms remains a bottleneck in breeding for resistance to salinity and drought. This study related the expression of key target genes with the physiological performance of durum wheat under different combinations of salinity and irrigation. The candidate genes assayed included two encoding for the DREB (dehydration responsive element binding) transcription factors TaDREB1A and TaDREB2B, another two for the cytosolic and plastidic glutamine synthetase (TaGS1 and TaGS2), and one for the specific Na(+) /H(+) vacuolar antiporter (TaNHX1). Expression of these genes was related to growth and different trait indicators of nitrogen metabolism (nitrogen content, stable nitrogen isotope composition, and glutamine synthetase and nitrate reductase activities), photosynthetic carbon metabolism (stable carbon isotope composition and different gas exchange traits) and ion accumulation. Significant interaction between genotype and growing conditions occurred for growth, nitrogen content, and the expression of most genes. In general terms, higher expression of TaGS1, TaGS2, TaDREB2B, and to a lesser extent of TaNHX1 were associated with a better genotypic performance in growth, nitrogen, and carbon photosynthetic metabolism under salinity and water stress. However, TaDREB1A was increased in expression under stress compared with control conditions, with tolerant genotypes exhibiting lower expression than susceptible ones., (© 2015 Institute of Botany, Chinese Academy of Sciences.)

Although climate scenarios have predicted an increase in [CO(2)] and temperature conditions, to date few experiments have focused on the interaction of [CO(2)] and temperature effects in wheat development. Recent evidence suggests that photosynthetic acclimation is linked to the photorespiration and N assimilation inhibition of plants exposed to elevated CO(2). The main goal of this study was to analyze the effect of interacting [CO(2)] and temperature on leaf photorespiration, C/N metabolism and N transport in wheat plants exposed to elevated [CO(2)] and temperature conditions. For this purpose, wheat plants were exposed to elevated [CO(2)] (400 vs 700 µmol mol(-1)) and temperature (ambient vs ambient + 4°C) in CO(2) gradient greenhouses during the entire life cycle. Although at the agronomic level, elevated temperature had no effect on plant biomass, physiological analyses revealed that combined elevated [CO(2)] and temperature negatively affected photosynthetic performance. The limited energy levels resulting from the reduced respiratory and photorespiration rates of such plants were apparently inadequate to sustain nitrate reductase activity. Inhibited N assimilation was associated with a strong reduction in amino acid content, conditioned leaf soluble protein content and constrained leaf N status. Therefore, the plant response to elevated [CO(2)] and elevated temperature resulted in photosynthetic acclimation. The reduction in transpiration rates induced limitations in nutrient transport in leaves of plants exposed to elevated [CO(2)] and temperature, led to mineral depletion and therefore contributed to the inhibition of photosynthetic activity., (© 2015 Scandinavian Plant Physiology Society.)

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., (© 2013 Institute of Botany, Chinese Academy of Sciences.)

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., (© 2013 Institute of Botany, Chinese Academy of Sciences.)

Mercury (Hg) is a toxic metal that affects plant growth. Here the effect of Hg exposure on plant growth and leaf gas-exchange together with gene expression in roots is reported for barley. Hg was mainly accumulated in roots and only very small amounts were found in the shoots. Chlorophyll fluorescence and net photosynthesis were not affected by Hg. Nevertheless exposure to Hg reduced shoot and root growth, the shoot to root ratio, stomatal conductance, carbon isotope discrimination and expression of an aquaporin transcript, whereas abscisic acid related transcripts were over-expressed. These results suggested some degree of limitation to water uptake causing a moderate water stress when plants are exposed to Hg. Microarray (MapMan) analysis revealed changes in the transcription of genes involved in nitrogen metabolism, which were accompanied by decreased nitrogen concentrations in the shoots, together with an increase in transcripts associated with secondary metabolism, stress, inhibition of DNA synthesis/chromatin structure and cell organization elements. Moreover, Hg induced the expression of many transcripts known to be involved in the uptake, accumulation, transport and general responses to other heavy metals. It is concluded that barley is able to accumulate high amounts of Hg in roots through several transcriptional, metabolic and physiological adjustments.

This study compared the performance of the stable isotope composition of carbon (δ(13) C), oxygen (δ(18) O) and nitrogen (δ(15) N) by tracking plant response and genotypic variability of durum wheat to different salinity conditions. To that end, δ(13) C, δ(18) O and δ(15) N were analysed in dry matter (dm) and the water-soluble fraction (wsf) of leaves from plants exposed to salinity, either soon after plant emergence or at anthesis. The δ(13) C and δ(18) O of the wsf recorded the recent growing conditions, including changes in evaporative conditions. Regardless of the plant part (dm or wsf), δ(13) C and δ(18) O increased and δ(15) N decreased in response to stress. When the stress conditions were established just after emergence, δ(15) N and δ(13) C correlated positively with genotypic differences in biomass, whereas δ(18) O correlated negatively in the most severe treatment. When the stress conditions were imposed at anthesis, relationships between the three isotope signatures and biomass were only significant and positive within the most severe treatments. The results show that nitrogen metabolism, together with stomatal limitation, is involved in the genotypic response to salinity, with the relative importance of each factor depending on the severity and duration of the stress as well as the phenological stage that the stress occurs., (© 2012 Blackwell Publishing Ltd.)

The need of a better adaptation of crops to drought is an issue of increasing urgency. However, enhancing the tolerance of maize has, therefore, proved to be somewhat elusive in terms of plant breeding. In that context, proper phenotyping remains as one of the main factors limiting breeding advance. Topics covered by this review include the conceptual framework for identifying secondary traits associated with yield response to drought and how to measure these secondary traits in practice.

Plant growth, root characteristics and the stable carbon (δ13C) and oxygen (δ18O) composition were studied in durum wheat. Four recombinant inbred lines with good agronomic adaptation were grown under well watered (WW) and water stress (WS) conditions until mid-grain filling in lysimeters. Gas exchange was measured in the flag leaf just before harvest and then the aerial dry matter (Aerial DM), root weight density (RWD) and root length density (RLD) and the specific root length (SRL) were evaluated and the δ13C and δ18O of the roots, the flag leaf blade and the spike were analysed. Water stress decreased stomatal conductance, plant accumulated transpiration and Aerial DM, whereas δ13C and δ18O increased. Genotypic differences were found for all gas-exchange and root traits and isotope signatures. Aerial DM was positively correlated with RLD, regardless of the water regime, whereas it was negatively correlated with δ13C and δ18O, but only under WW conditions. Moreover, RWD and RLD were negatively related to both δ13C and δ18O under the WW regime, but no clear pattern existed under WS. Our study supports the use of δ13C and δ18O as proxies for selecting root traits associated with higher growth in the absence of water stress.

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., (© 2012 Institute of Botany, Chinese Academy of Sciences.)

• Accurate phenotyping remains a bottleneck in breeding for salinity and drought resistance. Here the combined use of stable isotope compositions of carbon (δ¹³C), oxygen (δ¹⁸O) and nitrogen (δ¹⁵N) in dry matter is aimed at assessing genotypic responses of durum wheat under different combinations of these stresses. • Two tolerant and two susceptible genotypes to salinity were grown under five combinations of salinity and irrigation regimes. Plant biomass, δ¹³C, δ¹⁸O and δ¹⁵N, gas-exchange parameters, ion and N concentrations, and nitrate reductase (NR) and glutamine synthetase (GS) activities were measured. • Stresses significantly affected all traits studied. However, only δ¹³C, δ¹⁸O, δ¹⁵N, GS and NR activities, and N concentration allowed for clear differentiation between tolerant and susceptible genotypes. Further, a conceptual model explaining differences in biomass based on such traits was developed for each growing condition. • Differences in acclimation responses among durum wheat genotypes under different stress treatments were associated with δ¹³C. However, except for the most severe stress, δ¹³C did not have a direct (negative) relationship to biomass, being mediated through factors affecting δ¹⁸O or N metabolism. Based upon these results, the key role of N metabolism in durum wheat adaptation to salinity and water stress is highlighted., (No claim to original US government works. New Phytologist © 2012 New Phytologist Trust.)

The physiological performance of durum wheat and two related amphiploids was studied during the reproductive stage under different combinations of salinity and irrigation. One triticale, one tritordeum, and four durum wheat genotypes were grown in pots in the absence of stress until heading, when six different treatments were imposed progressively. Treatments resulted from the combination of two irrigation regimes (100% and 35% of container water capacity) with three levels of water salinity (1.8, 12, and 17 dS m(-1)), and were maintained for nearly 3 weeks. Gas exchange and chlorophyll fluorescence and content were measured prior to harvest; afterwards shoot biomass and height were recorded, and Delta(13)C, delta(15)N, and the concentration of nitrogen (N), phosphorus, and several ions (K(+), Na(+), Ca(2+), Mg(2+)) were analysed in shoot material. Compared with control conditions (full irrigation with Hoagland normal) all other treatments inhibited photosynthesis through stomatal closure, accelerated senescence, and decreased biomass. Full irrigation with 12 dS m(-1) outperformed other stress treatments in terms of biomass production and physiological performance. Biomass correlated positively with N and delta(15)N, and negatively with Na(+) across genotypes and fully irrigated treatments, while relationships across deficit irrigation conditions were weaker or absent. Delta(13)C did not correlate with biomass across treatments, but it was the best trait correlating with phenotypic differences in biomass within treatments. Tritordeum produced more biomass than durum wheat in all treatments. Its low Delta(13)C and high K(+)/Na(+) ratio, together with a high potential growth, may underlie this finding. Mechanisms relating delta(15)N and Delta(13)C to biomass are discussed.

We compared the performance of different physiological traits that reveal genotypic variations in tolerance to salinity in durum wheat. A set of 114 genotypes was grown in hydroponics for over 3 months. Three conditions: control, moderate (12 dS m -1 ) and severe (17 dS m -1 ) salinity, were maintained for nearly 8 weeks before harvest. The genotype biomass in control conditions correlated with the biomass at the two salinity levels. Subsequently, two subsets of 10 genotypes each were selected on the basis of extreme differences in biomass at the two salinity levels while showing relatively similar biomass in control conditions. Carbon isotope discrimination (Δ 13 C), nitrogen isotope composition (δ 15 N), and the concentration of nitrogen, phosphorus and several ions (K + , Na + , Ca 2+ , Mg 2+ ) were analysed in the two subsets for the three treatments. At 12 dS m -1 , K + concentration, K + /Na + ratio, Δ 13 C and δ 15 N correlated positively and Na + correlated negatively with shoot biomass. Under control conditions and at 17 dS m -1 no correlation was observed. However, the trait that correlated best with genotypic differences in biomass was δ 15 N at 12 dS m -1 . This trait was the first variable chosen at each of the two salinity levels in a stepwise analysis. We consider the possible mechanisms relating δ 15 N to biomass and the use of this isotopic signature as a selection trait.

This article reports the effect of growing conditions on the anatomical and ultrastructural changes associated with the development of photoautotrophy in Gardenia jasminoides Ellis plantlets during shoot multiplication in vitro. Two photosynthetic photon flux densities (PPFD) (50 and 100 µmol m-2 s-1 PPFD: L50 and L100, respectively) and two sucrose concentrations in the culture medium (5 and 30 g L-1: S5 and S30, respectively) were assayed. An increase in PPFD stimulated the development of photosynthetic tissues and led to higher photosynthesis and dark respiration regardless of the sucrose level assayed. However, the effect of sucrose in the medium depended on the PPFD. For the high-PPFD treatment, a low sucrose concentration in the medium stimulated the development of photosynthetic tissues, whereas the opposite effect of sucrose was observed at low PPFD. This study demonstrated that an increase in light intensity to moderate values such as L100 has a beneficial effect on the development of structural changes associated with photoautotrophy. Such an effect is stimulated by low sucrose (S5) levels in the medium. These modifications of usual growing conditions (such as L50 combined with S30) for micropropagation may prove to be useful in mass propagation of gardenia. However, the use of a low sucrose level in addition to conventional growing PPFD may be counterproductive.

Photosynthetic rates and related anatomical characteristics of leaves developed at three levels of irradiance (1200, 300 and 80 umol · m(-2) · s(-1)) were determined in the C4-like species Flaveria brownii A.M. Powell, the C3-C4-intermediate species F. linearis Lag., and the F1 hybrid between them (F. brownii × F. linearis). In the C3-C4 and F1 plants, increases in photosynthetic capacity per unit leaf area were strongly correlated with changes in mesophyll area per unit leaf area. The C4-like plant F. brownii, however, showed a much lower correlation between photosynthetic capacity and mesophyll area per unit leaf area. Plants of F. brownii developed at high irradiance showed photosynthetic rates per unit of mesophyll cell area 50% higher than those plants developed at medium irradiance. These results along with an increase in water-use efficiency are consistent with an increase of C4 photosynthesis in high-irradiance-grown F. brownii plants, whereas in the other two genotypes such plasticity seems to be absent. Photosynthetic discrimination against (13)C in the three genotypes was less at high than at low irradiance, with the greatest change occurring in F. brownii. Discrimination against (13)C expressed as δ (13)C was linearly correlated (r (2) = 0.81; P<0.001) with the ratio of bundle-sheath volume to mesophyll cell area when all samples from the three genotypes were combined. This tissue ratio increased for F. brownii and the F1 hybrid as growth irradiance increased, indicating a greater tendency towards Kranz anatomy. The results indicated that F. brownii had plasticity in its C4-related anatomical and physiological characteristics as a function of growth irradiance, whereas plasticity was less evident in the F1 hybrid and absent in F. linearis.

Several leaf anatomical and ultrastructural characteristics usually related with photosynthetic capacity were examined in two Flaveria species with strong differences in anatomy and their F1 hybrid. Flaveria trinervia (Spreng.) Mohr (C4) was the female parent and F. brownii A.M. Powell (C4-like) was the male parent. Quantitative anatomical analysis was made on transverse sections of leaves at both the light and electron microscope level. Four kinds of photosynthetic tissues were considered: bundle sheath (BS), mesophyll adjacent to the BS, mesophyll not adjacent to the BS, and larger spongy mesophyll cells. Flaveria trinvervia partitioned a larger proportion of its photosynthetic cells to BS and the mesophyll layer adjacent to BS and also possessed larger chloroplasts, especially in BS, than did F. brownii. These results suggest that although F. brownii is very C4-like, its anatomy is not as completely C4 as is the case for F. trinervia. In the F1 hybrid the relative contribution of the different tissues to the total photosynthetic tissue volume and area per unit leaf area was quite similar to that of F. trinervia. On the other hand, the chloroplast density and size of the F1 hybrid were fairly similar to those of F. brownii, especially in BS. Thus, there was no evidence of maternal inheritance in the chloroplast characteristics studied. A negative correlation (P<0.05) between chloroplast size and density was observed among species and relicates within each kind of tissue. This correlation was highest (r=-0.94, P<0.001) for the BS and when values were plotted on a logarithmic scale. Thus, higher chloroplast numbers for F. brownii and the F1 hybrid were offset by larger chloroplasts in F. trinervia. Less complete C4 photosynthesis in F. brownii may be partially due to incomplete development of Kranz anatomy usually associated with C4 photosynthesis.