Quinoa (Chenopodium quinoa) is well-known for high nutritional value and wide adaptability, but it is considered to be heat sensitive. To address this issue, accessions from two tetraploid wild relatives, C. berlandieri and C. hircinum, both native to hot environments, were evaluated alongside lowland and highland ecotypes of cultivated quinoa under field conditions with differing planting dates. Chenopodium berlandieri showed the best yield under the extreme heat experienced during the last planting, followed by lowland quinoa, C. hircinum, and highland quinoa. The yield advantage of C. berlandieri was achieved by maintaining higher grain number/seed set. Pollen viability was positively correlated with seed set under heat stress in cultivated quinoa, indicating its limiting effects. Considerable variation was observed for pollen viability among representative accessions of each species/ecotypes after a 38/33 °C day/night treatment for 5 d, ranging from an 80% reduction observed in highland quinoa to a 30% reduction in C. berlandieri. The most heat-sensitive period for pollen viability was 8-10 d before flowering, corresponding to the early pollen mother cell stage and it was conserved among the different species. In vitro pollen germination tests also demonstrated the heat tolerance of C. berlandieri. Taken together, our results suggest that wild relatives, particularly C. berlandieri, could be crossed with cultivated quinoa to introduce reproductive heat tolerance.

Plant responses to salt stress involve complex processes integrating short- and long-term adaptations, including changes in ion transport, systemic signaling, root architecture, and biomass distribution. A key adaptive mechanism involves the regulation of sodium (Na+) and potassium (K+) ion transport via Class 1 HKT1 transporters, which reduce Na+ accumulation in shoots, thereby enhancing salinity tolerance but at the expense of lateral root development. In this study, we identified differential roles of TMAC2 in modulating ABA accumulation and lateral root development under salt stress in two distinct Arabidopsis genotypes, Col-0 and C24. Overexpression of TMAC2 in the Col-0 background increased ABA accumulation, resulting in reduced lateral root development, suggesting a positive feedback loop involving HKT1, TMAC2, and ABA signaling. In contrast, TMAC2 overexpression in C24 reduced ABA accumulation in lines overexpressing HKT1, indicating genotype-specific differences in the TMAC2-HKT1 interaction. Additionally, we observed that the co-expression of TMAC2 and HKT1 in Col-0 induced ABI4 and ABI5 transcription factors, which are known to mediate salt sensitivity. These findings reveal a regulatory network where TMAC2 and HKT1 modulate salt stress responses through genotype-dependent feedback mechanisms. Our results highlight the complexity of root remodeling under salt stress and the crucial role of genetic background in shaping these adaptive responses. ### Competing Interest Statement The authors have declared no competing interest.

The identification of genes involved in salinity tolerance has primarily focused on model plants and crops. However, plants naturally adapted to highly saline environments offer valuable insights into tolerance to extreme salinity. Salicornia plants grow in coastal salt marshes, stimulated by NaCl. To understand this tolerance, we generated genome sequences of two Salicornia species and analyzed the transcriptomic and proteomic responses of Salicornia bigelovii to NaCl. Subcellular membrane proteomes reveal that SbiSOS1, a homolog of the well-known SALT-OVERLY-SENSITIVE 1 (SOS1) protein, appears to localize to the tonoplast, consistent with subcellular localization assays in tobacco. This neo-localized protein can pump Na+ into the vacuole, preventing toxicity in the cytosol. We further identify 11 proteins of interest, of which SbiSALTY, substantially improves yeast growth on saline media. Structural characterization using NMR identified it as an intrinsically disordered protein, localizing to the endoplasmic reticulum in planta, where it can interact with ribosomes and RNA, stabilizing or protecting them during salt stress.

The rice Zaxinone Synthase (ZAS) gene encodes a carotenoid cleavage dioxygenase (CCD) that forms the apocarotenoid growth regulator zaxinone in vitro. Here, we generated and characterized constitutive ZAS-overexpressing rice lines, to better understand ZAS role in determining zaxinone content and regulating growth and architecture. ZAS overexpression enhanced endogenous zaxinone level, promoted root growth and increased the number of productive tillers, leading to about 30% higher grain yield per plant. Hormone analysis revealed a decrease in strigolactone (SL) content, which we confirmed by rescuing the high-tillering phenotype through application of a SL analogue. Metabolomics analysis revealed that ZAS overexpressing plants accumulate higher amounts of monosaccharide sugars, in line with transcriptome analysis. Moreover, transgenic plants showed higher carbon (C) assimilation rate and elevated root phosphate, nitrate and sulphate level, enhancing the tolerance towards low phosphate (Pi). Our study confirms ZAS as an important determinant of rice growth and architecture and shows that ZAS regulates hormone homoeostasis and a combination of physiological processes to promote growth and grain yield, which makes this gene an excellent candidate for sustainable crop improvement.

Grafting high-yielding tomato varieties onto stress-tolerant rootstocks can mitigate the adverse effects of saline water irrigation on plant tomato productivity in arid regions like Saudi Arabia. This study investigates the efficacy of grafting tomatoes onto both novel and commercial rootstocks to enhance salinity tolerance and its impact on growth, physiological parameters, and yield in a soilless culture system. The experiment involved two water quality levels, 2 (S1) and 4 (S2) dS m−1, two growth media types, volcanic rock (M1) and sand (M2), and six grafting treatments: Tone Guitar F1 non-grafted (G1) (commercial scion), grafted onto itself (G2), Tone Guitar F1* Maxifort F1 (G3) (commercial rootstock), and grafting the scion onto three novel rootstocks, X-218 (G4), X-238 (G5), and Alawamiya365 (G6). Growth, physiology, photosynthetic pigments, and yield improved with lower salinity (2 dS m−1) in volcanic rock and with the grafting treatments (G2–G6) compared to the non-grafted treatment (G1). The best results were achieved with the S1M1G5 treatment, where yield increased by 53% compared to the lowest yield in non-grafted plants grown in sand under higher salinity (S2M2G1). All studied traits were adversely affected under high salinity (S2) in sandy media, with the G1 treatment resulting in the lowest values for these traits.

The rice Zaxinone Synthase (ZAS) gene encodes a carotenoid cleavage dioxygenase (CCD) that forms the apocarotenoid growth regulator zaxinone. Here, we generated and characterized constitutive ZAS -overexpressing rice lines, to better understand ZAS role in determining zaxinone content and regulating growth and architecture. ZAS overexpression enhanced endogenous zaxinone level, promoted root growth and meristem size, and increased the number of productive tillers, leading to an up to 30% higher grain yield per plant. Hormone analysis revealed a decrease in strigolactone (SL) content, which we confirmed by rescuing the high-tillering phenotype through application of a SL analog. Metabolomics analysis revealed that ZAS overexpressing plants accumulate higher amounts of monosaccharide sugars, in line with transcriptome analysis. Moreover, transgenic plants showed higher carbon (C) assimilation rate and elevated root phosphate, nitrate and sulfate level, enhancing the tolerance towards low phosphate (Pi) and indicating a generally better nutrient uptake. Our study shows that ZAS regulates hormone homeostasis and a combination of physiological processes to promote growth and grain yield, which makes this gene an excellent candidate for sustainable crop improvement. Teaser Zaxinone Synthase overexpression modulates rice metabolism and physiology and improves growth and phosphate uptake.

Despite the numerous advances made in our understanding of the physiology and molecular genetics of salinity tolerance, there have been relatively few applications of these to improve the salt tolerance of crops. The most significant advances have historically utilized intraspecific variation, introgression of traits from close crop wild relatives, or, less frequently, introgression from more distant relatives. Advanced lines often fail due to difficulties in the introgression or tracking of traits or due to yield penalties associated with the alleles in nonsaline environments. However, the greatest limitation is that salinity is not a primary trait for breeders. We must close the gap between research and delivery, especially for farmers who have precious few alternatives. These efforts should include a reassessment of old techniques such as grafting current crops with salt-tolerant hybrid rootstocks. Alternatively, future crops can be produced via domestication of salt-tolerant wild species—an approach that is now feasible in our lifetime.

Salinity remains a major inhibitor of crop production in irrigated and marginal lands. The identification of genes involved in salinity tolerance has been predominantly limited to model plants and crop species. However, plants naturally adapted to highly saline environments can provide key insights into mechanisms of salinity tolerance. Plants of the genus Salicornia grow in coastal salt marshes, and their growth is even stimulated by NaCl – much can be learnt from them. We generated genome sequences of two Salicornia species and studied the transcriptomic and proteomic responses of Salicornia bigelovii to NaCl. Through the generation of subcellular membrane proteomes, we found that SbiSOS1, a homolog of the well-known SALT-OVERLY-SENSITIVE 1 (SOS1) protein, appears to localize to the tonoplast, where it could be involved in mediating Na + translocation into the vacuole to prevent toxicity in the cytosol. We identified 11 proteins of interest which, when expressed in yeast, altered salinity tolerance. One of these proteins, SbiSALTY, substantially improves yeast growth on saline media. Structural characterization using NMR showed it to be an intrinsically disordered protein and to localize to the endoplasmic reticulum in planta , where it could interact with ribosomes and RNA, potentially stabilizing or protecting them during salt stress. The study and understanding of the molecular mechanisms providing high salinity tolerance in S. bigelovii is likely to provide significant insights for improving salinity tolerance of crop plants.

EDITORIAL article Front. Plant Sci., 27 July 2022Sec. Plant Abiotic Stress https://doi.org/10.3389/fpls.2022.985057

This article is a Commentary on Zhou et al. (2022), 236: 479–494.

Global use of nitrogen (N) fertilizers has increased sevenfold from 1960 to 1995 but much of the N applied is lost to the environment. Modifying the temporal and spatial distribution of organic N within the plant can lead to improved grain yield and/or grain protein content for the same or reduced N fertilizer inputs. Biotechnological approaches to modify whole plant distribution of amino acids and ureides has proven successful in several crop species. Manipulating selective autophagy pathways in crops has also improved N remobilization efficiency to sink tissues whilst the contribution of ribophagy, RNA and purine catabolism to N recycling in crops is still too early to foretell. Improved recycling and remobilization of N must exploit N-stress responsive transcriptional regulators, N-sensing or phloem-localized promotors and genetic variation for N-responsive traits.

Tartaric acid (TA) is an obscure end point to the catabolism of ascorbic acid (Asc). Here, it is proposed as a “specialized primary metabolite”, originating from carbohydrate metabolism but with restricted distribution within the plant kingdom and lack of known function in primary metabolic pathways. Grapes fall into the list of high TA-accumulators, with biosynthesis occurring in both leaf and berry. Very little is known of the TA biosynthetic pathway enzymes in any plant species, although recently some progress has been made in this space. New technologies in grapevine research such as the development of global co-expression network analysis tools and genome-wide association studies, should enable more rapid progress. There is also a lack of information regarding roles for this organic acid in plant metabolism. Therefore this review aims to briefly summarize current knowledge about the key intermediates and enzymes of TA biosynthesis in grapes and the regulation of its precursor, ascorbate, followed by speculative discussion around the potential roles of TA based on current knowledge of Asc metabolism, TA biosynthetic enzymes and other aspects of fruit metabolism.

The scarcity of freshwater is an increasing concern in flood-irrigated rice, whilst excessive use of nitrogen fertilizers is costly and contributes to environmental pollution. To co-ordinate growth adaptation under prolonged exposure to limited water or excess nitrogen supply, plants employ complex systems for signalling and regulation of metabolic processes. There is limited information on the involvement of one of the most important post-translational modifications (PTMs), protein phosphorylation, in plant adaptation to long-term changes in resource supply. Oryza sativa cv. Nipponbare was grown under two regimes of nitrogen from the time of germination to final harvest. Twenty-five days after germination, water was withheld from half the pots in each nitrogen treatment and low water supply continued for an additional 26 days, while the remaining pots were well watered. Leaves from all four groups of plants were harvested after 51 days in order to test whether phosphorylation of leaf proteins responded to prior abiotic stress events. The dominant impact of these resources is exerted in leaves, where PTMs have been predicted to occur. Proteins were extracted and phosphopeptides were analysed by nanoLC-MS/MS analysis, coupled with label-free quantitation. Water and nitrogen regimes triggered extensive changes in phosphorylation of proteins involved in membrane transport, such as the aquaporin OsPIP2-6, a water channel protein. Our study reveals phosphorylation of several peptides belonging to proteins involved in RNA-processing and carbohydrate metabolism, suggesting that phosphorylation events regulate the signalling cascades that are required to optimize plant response to resource supply.

Quinoa is a crop originating in the Andes but grown more widely and with the genetic potential for significant further expansion. Due to the phenotypic plasticity of quinoa, varieties need to be assessed across years and multiple locations. To improve comparability among field trials across the globe and to facilitate collaborations, components of the trials need to be kept consistent, including the type and methods of data collected. Here, an internationally open-access framework for phenotyping a wide range of quinoa features is proposed to facilitate the systematic agronomic, physiological and genetic characterization of quinoa for crop adaptation and improvement. Mature plant phenotyping is a central aspect of this paper, including detailed descriptions and the provision of phenotyping cards to facilitate consistency in data collection. High-throughput methods for multi-temporal phenotyping based on remote sensing technologies are described. Tools for higher-throughput post-harvest phenotyping of seeds are presented. A guideline for approaching quinoa field trials including the collection of environmental data and designing layouts with statistical robustness is suggested. To move towards developing resources for quinoa in line with major cereal crops, a database was created. The Quinoa Germinate Platform will serve as a central repository of data for quinoa researchers globally.

Growing plants hydroponically is a costing space and changing nutrient solutions on required time is a tedious task. The protocol described here is for nitrogen influx analysis of roots using N label. Preparing a labeled solution is not described here; however, the process for nitrogen influx is constructed well to understand the design of the experiment. After the treatment, the shoots and roots can be seperated and dried in an oven for further analysis.

Water and nitrogen availability limit crop productivity globally more than most other environmental factors. Plant availability of macronutrients such as nitrate is, to a large extent, regulated by the amount of water available in the soil, and, during drought episodes, crops can become simultaneously water and nitrogen limited. In this review, we explore the intricate relationship between water and nitrogen transport in plants, from transpiration-driven mass flow in the soil to uptake by roots via membrane transporters and channels and transport to aerial organs. We discuss the roles of root architecture and of suberized hydrophobic root barriers governing apoplastic water and nitrogen movement into the vascular system. We also highlight the need to identify the signalling cascades regulating water and nitrogen transport, as well as the need for targeted physiological analyses of plant traits influencing water and nitrogen uptake. We further advocate for incorporation of new phenotyping technologies, breeding strategies, and agronomic practices to improve crop yield in water- and nitrogen-limited production systems.

Roots vary their permeability to aid radial transport of solutes towards xylem vessels in response to nutritional cues. Nitrogen (N) depletion was previously shown to induce early suberization of endodermal cell walls and reduce hydraulic conductivity of barley roots suggesting reduced apoplastic transport of ions (Armand et al., 2019). Suberization may also limit transcellular ion movement by blocking access to transporters (Barberon et al., 2016). The aim of this study was to confirm that N depletion induced suberization in the roots of barley and demonstrate that this was a specific effect in response to NO3- depletion. Furthermore, in roots with early and enhanced suberization, we assessed their ability for transporter-mediated NO3- influx. N depletion induced lateral root elongation and early and enhanced endodermal suberization of the seminal root of each genotype. Both root to shoot NO3- translocation and net N uptake was half that of plants supplied with steady-state NO3-. Genes with predicted functions in suberin synthesis (HvHORST) and NO3- transport (HvNRT2.2) were induced under N-deplete conditions. N-deplete roots had a higher capacity for high-affinity NO3- influx in early suberized roots than under optimal NO3-. In conclusion, NO3- depletion induced early and enhanced suberization in the roots of barley, however, suberization did not restrict transcellular NO3- transport.

Degradation of nitrogen-rich purines is tightly and oppositely regulated under drought and low nitrogen supply in bread wheat. Allantoin is a key target metabolite for improving nitrogen homeostasis under stress. The metabolite allantoin is an intermediate of the catabolism of purines (components of nucleotides) and is known for its housekeeping role in nitrogen (N) recycling and also for its function in N transport and storage in nodulated legumes. Allantoin was also shown to differentially accumulate upon abiotic stress in a range of plant species but little is known about its role in cereals. To address this, purine catabolic pathway genes were identified in hexaploid bread wheat and their chromosomal location was experimentally validated. A comparative study of two Australian bread wheat genotypes revealed a highly significant increase of allantoin (up to 29-fold) under drought. In contrast, allantoin significantly decreased (up to 22-fold) in response to N deficiency. The observed changes were accompanied by transcriptional adjustment of key purine catabolic genes, suggesting that the recycling of purine-derived N is tightly regulated under stress. We propose opposite fates of allantoin in plants under stress: the accumulation of allantoin under drought circumvents its degradation to ammonium (NH4+) thereby preventing N losses. On the other hand, under N deficiency, increasing the NH4+ liberated via allantoin catabolism contributes towards the maintenance of N homeostasis.

Intensification of agricultural land and the overuse of inorganic fertilisers has led to soil acidification, depletion of organic matter, and environmental pollution. Approaches to protect soil health, including the enhancement of microbial diversity, are integral to improving crop productivity and food security. Metagenomics has rapidly improved our understanding of soil microbial diversity and function, while genetic techniques have helped to dissect the complex signal exchange between microorganisms and plants. This article presents and evaluates reported beneficial effects of plant growth-promoting bacteria (PGPB), focussing on those capable of mobilising or solubilising nutrients and/or stimulating plant growth when applied to agriculturally important grasses. The agricultural industry has capitalised on these scientific advancements, generating microbial formulations for specific crop responses. However, scientific methodologies must be applied in order to overcome inherent limitations of many PGPBs including their inability to be cultured, their poorly defined or multiple modes of action, a low level of integration with the crop partner, and an unpredictability in translating beneficial plant responses to the field. Novel approaches such as engineered rhizospheres, enhancing endophytic systems, cereal nodule development, and the use of inoculant consortiums will be necessary to sustain growth in the biofertiliser industry.

Turn-over of RNA and catabolism of nucleotides releases one to four ammonia molecules; the released nutrients being reassimilated into primary metabolism. Preliminary evidence indicates that monocots store high levels of free nucleotides and nucleosides but their potential as a source of internal organic nitrogen for use and remobilization is uncharted. Early tillering wheat plants were therefore starved of N over a 5-day time-course with examination of nucleic acid yields in whole shoots, young and old leaves and roots. Nucleic acids constituted ∼4% of the total N pool of N starved wheat plants, which was comparable with the N available from nitrate (NO3-) and greater than that available from the sum of 20 proteinogenic amino acids. Methods were optimized to detect nucleotide (purine and pyrimidine) metabolites, and wheat orthologs of RNA degradation (TaRNS), nucleoside transport (TaENT1, TaENT3) and salvage (TaADK) were identified. It was found that N starved wheat roots actively catabolised RNA and specific purines but accumulated pyrimidines. Reduced levels of RNA corresponded with induction of TaRNS2, TaENT1, TaENT3, and TaADK in the roots. Reduced levels of GMP, guanine, xanthine, allantoin, allantoate and glyoxylate in N starved roots correlated with accumulation of allantoate and glyoxylate in the oldest leaf, suggesting translocation of allantoin. Furthermore, N starved wheat plants exogenously supplied with N in the form of purine catabolites grew and photosynthesized as well as those plants re-supplied with NO3-. These results support the hypothesis that the nitrogen and carbon recovered from purine metabolism can support wheat growth.