Salinity represents a major constraint on plant development and crop productivity in wheat, which represents one of the most critical sources of dietary calories worldwide. Its detrimental effects are particularly pronounced during the early stages of growth, including seed germination and seedling establishment. Salinity tolerance is a multifaceted trait governed by several interrelated mechanisms, notably ion homeostasis, osmotic adjustment, activation of enzymatic antioxidant systems, and transcriptional regulation of ion transporter genes. In the present study, contrasting wheat genotypes exhibiting differential salinity tolerance were selected from a panel of 172 accessions evaluated under salinity stress (175 mM NaCl) and control conditions (0 mM NaCl). The objectives of the current study are to confirm the underlying physiological and molecular mechanisms conferring salinity tolerance. Key physiological and molecular parameters including Na+, K+, and P homeostasis; activities of major antioxidant enzymes; and expression profiles of the salinity-responsive ion transporter genes TaAVP1 and NHX1 were quantified in six tolerant genotypes and one susceptible genotype. The tolerant genotypes exhibited higher concentrations of Na+ and K+ and elevated activities of all antioxidant enzymes, compared with the susceptible genotype. Furthermore, the tolerant genotypes showed differential expression of TaAVP1 and NHX1: both genes were upregulated in Javelin 48 and Kandahar, whereas they were downregulated in genotype 1018d. Notably, genotype Kule demonstrated the highest Na+ accumulation, accompanied by markedly elevated activities of all major antioxidant enzymes, with ascorbate peroxidase and glutathione reductase increasing by 9.20-fold and 2.32-fold, respectively, under salinity stress. Based on these findings, the tolerant genotypes can be categorized into two functional groups: Javelin 48, Ghati, and 1018d (characterized by high K+ and salinity tolerance) are better suited to soils affected by low Na+ salinity, whereas Kandahar, Kule, and 1049 (characterized by high Na+ and sodicity tolerance) are more adapted to soils with elevated Na+ levels. In conclusion, the tolerant genotypes exhibited distinct, coordinated mechanisms to mitigate salinity stress, underscoring the complexity and plasticity of adaptive responses in wheat.
The peduncle stem plays an essential role in resource allocation and water transport to spike grains under normal conditions. Therefore, exploring peduncle traits and their relationships with spike production under drought stress may provide insights into the mechanisms that mitigate the effects of drought stress on grain yield in wheat. To address this challenge, a panel of 198 highly homozygous and diverse spring wheat varieties was evaluated under normal and drought conditions across two growing seasons. Peduncle traits, namely, length (PL), diameter (PD), and weight (PW), as well as spike traits, namely, spike length (SPL), number of spikelets/spike (NSPS), grain number/spike (GNPS), and grain yield per spike (GYPS), and thousand-kernel weight (TKW), were assessed. We revealed that PW and PD, unlike PL, were significantly and strongly associated with spike traits and grain weight under both conditions. Genome-wide association study (GWAS) revealed that spike and peduncle traits were controlled by different genetic mechanisms, as no stable markers were shared between these two groups. Distinct SNPs found between genotypes with contrasting peduncle traits led to the identification of a key SNP marker in a gene model encoding a protein highly expressed in the peduncle and spike of wheat. Comparing cultivars with low peduncle trait values to cultivars with high peduncle trait values, particularly PW and PD, high peduncle trait cultivars had greater yield-related trait values under both drought and normal conditions. The results of this study shed light on the importance of peduncle traits in enhancing wheat spike productivity under normal and drought stress conditions.
Seed germination performance and storability are fundamental components of seed quality and critical for successful crop establishment. However, information on the impact of different crop production systems on the quality and storability of seed material is still limited. Therefore, the aim of this study was to compare the effects of different crop production systems (ecological, integrated, conventional, and monoculture) on seed germination and predisposition for storage. The research was carried out on four varieties of winter wheat. Seed material was produced within a two-year period, during which different weather conditions occurred. Four germination-related traits were assessed: germination capacity NS (%), total germination (TG%), time to reach 50% germination (t50) and the area under the germination curve (AUC). The results demonstrated that the cultivar, the cultivation system and the year of study had a significant impact on germination characteristics. The ecological system ensured the highest germination rate in fresh seeds. However, in the CD test, the conventional system demonstrated the highest levels of stress resistance and stability, suggesting the best storage potential. The significant system x variety interaction demonstrates the importance of accurate matching of the genotype to the growing conditions to ensure optimal seed quality. Furthermore, the data demonstrated a strong influence of climatic conditions in the year of production, which is crucial for seed vigor.
Water shortage is a major abiotic stress limiting wheat production worldwide, particularly in Iran. This study aimed to identify genetic loci and candidate genes associated with grain yield and related traits in spring bread wheat under water shortage conditions. Genotyping of 111 spring bread wheat genotypes was conducted using the 15K SNP array developed by TraitGenetics (Germany) on the Illumina platform. Phenotypic evaluations of the studied genotypes were conducted from 2020 To 2022 using a lattice square design under both well-watered and water shortage stress environments in Iran. Broad-sense heritability estimates under water shortage stress conditions varied among traits, with the lowest value (12.04
Amylase trypsin inhibitors (ATIs) are a group of proteins in all cereals' seeds, including wheat, barley, rye, and maize. Currently, ATIs are the most studied wheat components since they are linked to celiac disease and baker's asthma. The interest has increased since 2012, when they were shown to trigger the innate immune system and intestinal and extra-intestinal inflammation. Even though ATIs are present in many plant-based diets, the genetic factors underlying these proteins have not yet been investigated in barley. To this end, this study was designed to measure ten ATI proteins in a diverse barley collection grown under field conditions. Ample natural variation among the accessions in ten ATIs has been measured, which showed a substantial role in the accumulation of ATI_total, such as CMa, CMd, CMe, BTI_CMc, and AIBDAI_1. A genome-wide association scan (GWAS) utilizing a large number of molecular markers demonstrated that the accumulation of ATIs was influenced by many small to medium quantitative trait nucleotides (QTNs). Eight QTNs showed the highest association with ATIs, particularly AIBDAI, whereas these QTNs negatively influenced ATI accumulation. Genomic investigations identified Serine/threonine protein phosphatase as a putative candidate gene. Our research provides the initial analysis of the ATI proteins found in barley, which might potentially contribute to enhancing the quality of barley-based food products. As a result, the study demonstrated that cultivars with lower ATI proteins can speed up their integration to improve the quality of barley products and diminish the possibility of some diseases.
The escalating frequency and severity of environmental stressors pose a critical challenge to global crop production. Among the various mechanisms by which plants cope with these conditions, stress memory often referred to as "priming" has emerged as a powerful phenomenon. It enables plants to retain molecular, physiological, and epigenetic information from prior suboptimal conditions, thereby mounting faster and more robust defenses upon re-exposure to similar stresses. This review explores the underpinnings of abiotic stress memory in crop plants, focusing on key signaling pathways, genomic and epigenomic modifications, and the regulatory networks they influence. We further highlight practical avenues for leveraging this knowledge in crop breeding and management, particularly in light of climate change. By developing varieties capable of "remembering" and thus better resisting repeated or simultaneous stress events, agricultural systems can become more resilient while relying less on resource-intensive interventions. Ultimately, integrating stress memory into breeding frameworks offers an innovative approach to enhance crop adaptability and ensure a more secure global food supply.
The generation of stress memory in barley under salt stress conditions is a very important component of plant tolerance and adaptation to adverse environmental conditions. Mechanisms of stress memory allow the plant to "remember" the previous exposure to high salinity and to switch on more quickly pre-established pathways of defense, thus causing less damage and sustaining productivity. Understanding the role of stress memory in barley can thus help inform breeding strategies for salt tolerance cultivar development, which in turn would contribute to stability in crop yield in saline environments using a genome-wide association study (GWAS). To follow the effects of transgenerational, intergenerational, and combined effects of salt stress memory event on the third generation, we compared barley genotypes whose first and second generations were not exposed to salinity stress (C1C2) with the groups that had experienced a single-generation of salt stress either two generations ago (S1C2; first generation salt stress memory effects), or one generation ago (C1S2; second generation salt stress memory effects) and the group that had experienced salinity in both generations (S1S2; combined salt stress memory effects). Our results showed that the historical presence of salt stress, regardless of the number of generations previously exposed, had significant effects on changing various agronomic and physiological traits, including spike length, the number of spikelets and grains per spike, grain weight per spike, and thousand kernel weight. More importantly, the history of stress affected both osmolytes and metabolite levels. These results suggest that pretreatment with salt stress could provoke long-lasting changes in barley growth and stress tolerance, which enhance the capacity of its offspring to tolerate the same kind of stress. Based on GWAS analysis, a total of 100 SNP markers located on all chromosomes showed a highly significant association with several potential candidate genes at p-value ≥ 5 with all the measured morphological and biochemical attributes. Interestingly, the expression of our potential genes under salt stress in cereals plays a central role in the plant’s ability to perceive, transduce, and respond to salinity-induced stress signals. These genes participate in molecular-mediated pathways that activate critical adaptive responses, such as ion homeostasis, ROS detoxification, and hormonal regulation, which are crucial for plant survival and productivity under saline environments. Knowledge of the function of our potential candidate genes is important for the development of genetic and biotechnological strategies for improving stress tolerance in cereals.
Climate change is intensifying soil salinization, posing a major threat to crop establishment and productivity, particularly in arid and semi-arid regions. Barley (Hordeum vulgare L.), one of the most salt-tolerant cereals, offers valuable genetic resources for improving salinity resilience at early growth stages. This study exploited the genetic diversity of the Nested Association Mapping (NAM) population Halle Exotic Barley-25 (HEB-25) to dissect salinity tolerance during germination and seedling developmental stages. First, the HEB-25 parental lines (25 wild barley genotypes and cv. Barke) were evaluated under salinity treatment to identify contrasting responses. Based on this screening, four HEB families (01, 04, 09, and 22) were selected out of 25 HEB families for detailed phenotypic and genomic analysis. Seeds of the selected HEB families were subjected to 40% seawater salinity stress and control treatments to assess germination percentage and seedling traits, including shoot length, root length, fresh weight (FW), dry weight (DW), DW/FW ratio, root-shoot ratio, and salt tolerance index (STI). Substantial variation was observed among families for all measured traits under salinity stress. STI values enabled clear differentiation among families: Family 01 exhibited the most consistent overall tolerance profile, Family 22 showed the strongest sensitivity in biomass traits, and Family 04 displayed a trait-specific response with sensitivity at the family-mean level but exceptional within-family diversity, harboring some of the highest individual TI values across the population. A genome-wide association study was conducted using 32,995 SNP markers. A total of 27 significant SNPs were identified, corresponding to 20 quantitative trait loci (QTLs). Of these, 12 QTLs were detected under control conditions, 16 under seawater treatment, and 21 based on tolerance indices, indicating both constitutive and stress-responsive genetic effects. Gene annotation within these regions revealed approximately 23 candidate genes associated with abiotic stress tolerance, including genes involved in ion transport, osmotic adjustment, kinases and stress signaling pathways. HEB_22_003, HEB_04_087, and HEB_01_013 represent the most promising genotypes for salinity breeding. These findings highlight the effectiveness of combining precise phenotyping with high-resolution genomic analysis in the HEB-25 population to uncover the genetic architecture of salinity tolerance at early developmental stages. We identified 20 salinity-responsive QTLs, including five major-effect loci on chromosomes 2H, 4H, 5H, and 7H that consistently explained the largest share of phenotypic variation. These loci co-localized with candidate genes linked to ion homeostasis, Ca2+-mediated signaling, protein glycosylation, epigenetic regulation, and root system plasticity, revealing key mechanisms underlying early-stage salt adaptation in barley. The strong and contrasting responses of Family 01 and Family 04 provide an excellent genetic framework for functional validation of tolerance alleles. Collectively, these genomic resources establish a robust foundation for QTL pyramiding, marker-assisted breeding, and the development of climate-resilient barley cultivars for saline agroecosystems.
Climate change is intensifying the frequency and severity of abiotic stresses that threaten global food security by reducing crop productivity. Among these, saline stress poses a serious threat to barley (Hordeum vulgare L.) production. These conditions are increasingly prevalent in arid and semiarid regions, as well as in regions with limited access to freshwater resources, making the identification of salt tolerance genes essential for breeding resilient varieties. In this study, we evaluated 400 genotypes from the barley nested association mapping population HEB-25 under control conditions and 40% seawater irrigation to simulate moderate-to-high salinity stress. A genome-wide association study (GWAS) was conducted to identify alleles from wild barley [H. vulgare L. subsp. spontaneum (C. Koch) Thell.] associated with enhanced salt tolerance. Phenotypic evaluation included germination percentage (Ger%), shoot length (SL), root length (RL), root-shoot length ratio, seedling fresh weight, seedling dry weight, and salt tolerance index of the different traits. The HEB-25 families exhibited significant variation in seedling responses to seawater-induced salinity, with contrasting effects on SL, RL, and dry weight. Compared to the elite parental Barke, several genotypes demonstrated high tolerance under seawater stress, maintaining stable Ger% and exhibiting the highest tolerance indices. Moreover, GWAS results identified 60 highly significant single nucleotide polymorphisms associated with seedling growth parameters under both conditions. These findings underscore the value of the HEB-400 panel as a genetic resource for dissecting salinity tolerance mechanisms, identifying stress-adaptive alleles lost during domestication and a source of pre-breeding material for developing genotypes with enhanced salinity tolerance.
Introduction Drought is one of the most damaging abiotic stresses, reducing seed germination, impairing seedling establishment, and ultimately decreasing crop yield. The objectives of the present study were to identify associated SNP markers and genes associated with drought tolerance in wheat at early developmental stages and to explore gene networks that reveal gene-gene interactions under drought stress.Methods A total of 168 genotypes were tested for drought tolerance under 20% PEG (drought stress) and 0% PEG (control). Genome-wide association study, annotation analyses, gene network analysis were performed.Results High genetic variation was observed among genotypes for all traits scored under both conditions. The GWAS identified 130 and 128 significant SNPs under drought and control conditions, respectively. Gene annotation identified 98 genes responsive to drought stress, of which 54 have been previously reported to be associated with drought tolerance. Ten SNPs were common between control and drought treatments, nine of which were located within genes controlling variation in germination percentage. Gene network analysis of these nine gene models showed that they were organized into eight distinct pathways.Discussion These networks were regulated by master genes encoding proteins with diverse biological functions, including rRNA and tRNA methyltransferases, protein kinases, and potassium transmembrane transporters. These findings provide fundamental insights into the genetic basis of drought tolerance in wheat. These results indicate that GWAS is a robust and instrumental approach to identifying the associated genes. The network analysis explored the interactions among several genes, not the effect of each gene individually, which will help us to understand the drought tolerance during early stages in wheat. These findings are fundamental to better understand drought tolerance in wheat.
The article’s aims are to be a comprehensive exploration of factors influencing seed longevity, emphasizing the importance of seed viability for future plant generations, biodiversity conservation, and agricultural productivity. It covers topics related to seed biology, storage conditions, the impact of reactive oxygen species (ROS) and reactive nitrogen species (RNS), the role of seed-associated microbiomes, DNA damage, genetic factors affecting longevity, and the intricate processes involved in seed germination. The discrepancies between artificially aged seeds and those aged naturally in ambient and long-term genebank conditions are discussed, underlining the importance of studying long-term stored seeds to identify genetic markers associated with seed longevity accurately and highlighting the application of genome-wide association studies (GWAS) in uncovering genetic factors.
Hull-less barley (Hordeum vulgare L. var. nudum) is an underutilized form of barley that shows significant potential for broader use in human nutrition due to its exceptional nutritional value and beneficial health effects. This potential was the basis for research within the CROPDIVA project, funded by the Horizon Europe program. Despite its historical importance as a staple food, it currently occupies less than 2% of the total barley production area in Europe. The important agronomic, morphological and grain quality traits were investigated in order to determine their value for agriculture and the food industry. The study included 72 genotypes of winter hull-less barley obtained from the IPK genebank in Gatersleben (Germany), along with two control cultivars sown in 10 replications. The experiments were established during the 2023/24 and 2024/25 at the Rimski šančevi experimental station of the Institute of Field and Vegetable Crops in Novi Sad using an augmented block design. This research focused on assessing phenotypic variation of following characteristics: biomass, thousand grain weight, harvest index, grain color, threshability, test weight and grain protein content. Among the tested genotypes, 67 belong to the six-rowed (6R) type, and 7 genotypes are two-rowed (2R) types. After harvest, the protein content of the seeds was determined by the combustion method according to the DUMAS procedure (AOAC 992.23, 1998). Data analysis employed BLUEs (Best Linear Unbiased Estimators) to correct for spatial variation across the 10 blocks. Descriptive statistics revealed substantial phenotypic variability with the highest coefficients of variation (CV) observed for biomass (27.8%) and thousand-grain weight (27.5%). Conversely, hectoliter mass remained highly stable (CV = 4.2%). Two-rowed genotypes achieved a statistically significantly higher biomass (2,445 g m- ²) and thousand grain weight (49.1 g) compared to six-rowed barley, where the biomass was 1,614 g m- ², and thousand grain weight was 30.0 g. In contrast, the average values of hectoliter mass in six-rowed forms (84.6 kg hL-1) and protein content (12.3%) were higher compared to two-rowed forms (79.2 kg hL-1; 10.3%). Only seven genotypes showed a lower degree of threshability, with some seeds retaining hulls after mechanical dehulling. Aleurone layer analysis identified a predominance of white color, while light blue, dark blue, purple, and black pigments were less frequent. Correlation analysis showed that the most pronounced positive correlation was found between biomass and thousand grain weight (0.56). A modarate correlation was detected between TGW and hectoliter weight (-0.47). Principal component analysis (PCA) confirmed a high degree of phenotypic variability explaining 63.3% of the total phenotypic variability indicating mutual independence of traits related to protein content and yield components. These results confirm that the European collection of hull-less barley represents a valuable genetic resource suitable for use in breeding programs and improvement of cultivars intended for human consumption.
Background Small interfering RNAs (siRNAs), a subclass of small non-coding RNAs (sncRNAs), play crucial roles in regulating seed germination and viability through epigenetic mechanisms like RNA-directed DNA methylation (RdDM). This study presents the first comprehensive investigation of siRNA profiles linked to seed viability and germination in barley (Hordeum vulgare L.), utilizing a unique set of seeds from a single batch subjected to controlled long-term storage. Some seeds lost viability due to moisture exposure from unsealing, creating a natural experimental model to explore vigor effects.Results sRNA sequencing revealed 85,728 differentially expressed siRNAs, with distinct patterns between regenerated, high-viability, and low-viability seeds. Notably, trans-acting siRNAs (ta-siRNAs) showed peak abundance at different imbibition times depending on seed quality, suggesting dynamic regulation. Around 46% of siRNAs were 21 nucleotides, and 54% were 22 nucleotides long. Gene Ontology and degradome analyses confirmed siRNA target genes involved in vital biological processes such as cytochrome complex function, root development, cell maturation, and carbohydrate metabolism. Despite RNA degradation in low-viability seeds, siRNAs remained relatively stable.Conclusions This pioneering research uncovers novel insights into siRNA-mediated control of seed longevity and germination, highlighting the innovative use of stable, well-characterized plant material to disentangle molecular mechanisms underpinning seed vigor and germination success. siRNAs show potential in maintaining seed metabolic activity during dormancy release and germination initiation.
Wheat amylase trypsin inhibitors (ATIs) are prominent allergens in Baker’s asthma and contribute to innate inflammation in non-celiac wheat sensitivity (NCWS), linking them to metabolic and autoimmune diseases. Their tetra-, di-, and monomeric forms, stabilized by disulfide bonds, confer resistance to digestion, baking, and heating. Although proteomic studies reveal minor variation in ATI subtypes among cultivars and major variation among species, the influence of environment and wheat genotype on ATI levels and TLR4-stimulating activity remains unclear. We assessed the effect of the environment on the in vitro inflammatory bioactivity of ATIs extracted from 60 German wheat genotypes focusing on breeding over time between 1891 and 2010, and cultivation across three climatically distinct years. We found considerable genotype-dependent variation in ATI bioactivity that did not correlate with ATI subtype abundance, and observed no consistent difference between old and modern cultivars. ATIs from samples grown in 2019, a warm and dry year, showed reduced TLR4 activity, highlighting the significant impact of environmental conditions on inflammatory ATI bioactivity.
The Rht12 semidwarf mutant can potentially replace the long-used Rht1-B1b/Rht-D1b (DELLA) mutants in wheat breeding. The Rht12 mutant offers advantages over DELLA mutants, particularly in regions with limited rainfall, as it does not reduce coleoptile length, an important trait for such semiarid areas. The dwarfing mechanism in Rht12 involves the overexpression of the GA2-oxidase A13 gene, which reduces the bioactive gibberellin contents, leading to reduced plant height. In this two-year field study, the protein content and composition of the wheat variety 'Maris Huntsman' and its Rht12 mutant line (Triticum aestivum ssp. aestivum) were examined. Changes in the size- and hydrophobicity-based distribution of the total proteins of the samples have been monitored by SE- and RP-HPLC, respectively. Significant differences in protein content and composition have been observed between the Rht12 mutant and wild-type genotype, showing altered protein characteristics (glutenin-to-gliadin ratio, unextractable polymeric protein percentage-UPP%) directly related to techno-functional properties. In addition, the basic quality parameters of the wholemeal samples from both growing seasons were determined with NIR (near-infrared spectroscopy) and revealed that due to the Rht12 mutation, Zeleny sedimentation and gluten content significantly improved.
Background: A genome-wide association scan (GWAS) is a powerful tool for identifying genetic variants and specific loci underlying complex traits. Bread Wheat (Triticum aestivum L.) is one of the primary food resources in the world, and understanding its physiological parameters will help improve agronomic and yield traits. This study investigated single-nucleotide polymorphism (SNP) markers associated with physiological and agronomic traits in bread wheat to inform breeding programs. A diverse panel of 272 bread wheat genotypes was evaluated across two growing seasons (2019–2021) using a 16 × 17 rectangular lattice design with two replications. Key physiological traits, including carbon dioxide exchange and chlorophyll content, and agronomic traits, such as days to heading, days to maturity, flag leaf length, plant height, grain number per spike, grain weight per plant, thousand-grain weight, biological yield, and harvest index, were measured. Genotyping was conducted using a 90K SNP array at Trait-Genetics, Germany, yielding 17,093 high-quality SNPs after filtering for minor allele frequency and missing data (>10%). Results: Population structure analysis grouped the genotypes into five subgroups based on their genetic variation. GWAS was performed using General Linear Model (GLM), Fixed and random model Circulating Probability Unification (FarmCPU), and Mixed Linear Model (MLM), identifying 320, 302, and 27 significant marker-trait associations (MTAs), respectively. Sixteen MTAs were consistently significant across models, including four stable MTAs detected in two cropping seasons. These MTAs harbored 139 high-confidence genes associated with nine traits. Conclusions: These findings provide valuable insights into the genetic architecture of key wheat traits, facilitating targeted breeding strategies to enhance yield.
Amylase/trypsin-inhibitors (ATIs) are known allergens and triggers of non-celiac wheat sensitivity. Until now, ATIs were only quantitated in wheat species. We developed and validated a targeted stable isotope dilution analysis LC-MS/MS method to quantitate ten barley-specific ATIs, including one monomeric and one dimeric amylase-inhibitor, four chloroform/methanol-soluble types, three subtilisin/chymotrypsin-inhibitors and one amylase/subtilisin-inhibitor. After successful validation in terms of precision, recovery and limits of detection and quantitation, the method was applied to 181 barley accessions from the Global EcoSeed panel, comprising 113 two-row and 68 six-row barleys of different genetic backgrounds. The overall ATI content was 1.1-5.2 mg/g, corresponding to 0.7-3.6 % of the total protein content with no clear distinction between two-row and six-row barleys. This study is the first to provide insights on the ATI content and composition of barley, which can be used to make low-ATI foods for special dietary needs.
Spikelet development is an inherited complex trait influenced by several endogenous, genetic, and environmental factors. This study aims to find the genetic factors controlling the maximum yield potential (MYP) in a core set of 250 diverse spring barley accessions. To achieve this, we evaluated a collection of 250 barley spikes at three different developmental stages MYP, green anther (GA), and awn tipping (TIP). We also, measured the sugar (sucrose, glucose, and fructose) contents from each accession of the collection. Six-rowed accessions showed higher phenotypic variation for all studied traits than two-rowed. The spikelet-related traits showed a highly significant positive relationship with the sugar contents. Based on a genome-wide association study (GWAS) outputs, four quantitative trait nucleotide regions (QNRs) containing 37 significant quantitative trait nucleotides (QTNs) were identified, which encompass plausible candidate genes. The highest significant QTNs on 1H were located inside the HORVU.MOREX.r2.1HG0042180 and HORVU.MOREX.r2.1HG0072210 that encodes sugar transporter and auxin response factor. These candidate genes potentially promote sucrose accumulation and auxin during spikelet development by regulating sugar hydrolysis and transport in addition to auxin biosynthesis. Differential expression profiling revealed that these candidate genes are also highly expressed during spike development, confirming their potential role in regulating spikelet survival.