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We examined photosynthetic traits of pre-existing and newly developed flag leaves of four wheat genotypes grown in controlled-environment experiments. In newly developed leaves, acclimation of the maximum rate of net CO2 assimilation (A n) to warm nights (i.e. increased A n) was associated with increased capacity of Rubisco carboxylation and photosynthetic electron transport, with Rubisco activation state probably contributing to increased Rubisco activity. Metabolite profiling linked acclimation of A n to greater accumulation of monosaccharides and saturated fatty acids in leaves; these changes suggest roles for osmotic adjustment of leaf turgor pressure and maintenance of cell membrane integrity. By contrast, where A n decreased under warm nights, the decline was related to lower stomatal conductance and rates of photosynthetic electron transport. Decreases in A n occurred despite higher basal PSII thermal stability in all genotypes exposed to warm nights: T crit of 45–46.5 °C in non-acclimated versus 43.8–45 °C in acclimated leaves. Pre-existing leaves showed no change in A n–temperature response curves, except for an elite heat-tolerant genotype. These findings illustrate the impact of night-time warming on the ability of wheat plants to photosynthesize during the day, thereby contributing to explain the impact of global warming on crop productivity. [ABSTRACT FROM AUTHOR]

Drought stress poses a serious threat to grain formation in wheat. Nitrogen (N) plays crucial roles in plant organ development; however, the physiological mechanisms by which drought stress affects plant N availability and mediates the formation of grains in spikes of winter wheat are still unclear. In this study, we determined that pre-reproductive drought stress significantly reduced the number of fertile florets and the number of grains formed. Transcriptome analysis demonstrated that this was related to N metabolism, and in particular, the metabolism pathways of arginine (the main precursor for synthesis of polyamine) and proline. Continuous drought stress restricted plant N accumulation and reallocation rates, and plants preferentially allocated more N to spike development. As the activities of amino acid biosynthesis enzymes and catabolic enzymes were inhibited, more free amino acids accumulated in young spikes. The expression of polyamine synthase genes was down-regulated under drought stress, whilst expression of genes encoding catabolic enzymes was enhanced, resulting in reductions in endogenous spermidine and putrescine. Treatment with exogenous spermidine optimized N allocation in young spikes and leaves, which greatly alleviated the drought-induced reduction in the number of grains per spike. Overall, our results show that pre-reproductive drought stress affects wheat grain numbers by regulating N redistribution and polyamine metabolism. [ABSTRACT FROM AUTHOR]

Functional–structural plant models are increasingly being used by plant scientists to address a wide variety of questions. However, the calibration of these complex models is often challenging, mainly because of their high computational cost, and, as a result, error propagation is usually ignored. Here we applied an automatic method to the calibration of WALTer: a functional–structural wheat model that simulates the plasticity of tillering in response to competition for light. We used a Bayesian calibration method to jointly estimate the values of five parameters and quantify their uncertainty by fitting the model outputs to tillering dynamics data. We made recourse to Gaussian process metamodels in order to alleviate the computational cost of WALTer. These metamodels are built from an adaptive design that consists of successive runs of WALTer chosen by an efficient global optimization algorithm specifically adapted to this particular calibration task. The method presented here performed well on both synthetic and experimental data. It is an efficient approach for the calibration of WALTer and should be of interest for the calibration of other functional–structural plant models. [ABSTRACT FROM AUTHOR]

Crown roots are the main components of the fibrous root system in cereal crops and play critical roles in plant adaptation; however, the molecular mechanisms underlying their formation in wheat (Triticum aestivum) have not been fully elucidated. In this study, we identified a wheat basic helix-loop-helix (bHLH) protein, TabHLH123, that interacts with the essential regulator of crown root initiation, MORE ROOT in wheat (TaMOR). TabHLH123 is expressed highly in shoot bases and roots. Ectopic expression of TabHLH123 in rice resulted in more roots compared with the wild type. TabHLH123 regulates the expression of genes controlling crown-root development and auxin metabolism, responses, and transport. In addition, we analysed the nucleotide sequence polymorphisms of TabHLH123 s in the wheat genome and identified a superior haplotype, TabHLH123-6B , that is associated with high root dry weight and 1000-grain weight, and short plant height. Our study reveals the role of TabHLH123 in controlling the formation of crown roots and provides beneficial insights for molecular marker-assisted breeding in wheat. [ABSTRACT FROM AUTHOR]

In a previous study we used asymmetric-flow field-flow fractionation to determine the polymer mass (Mw), gyration radius (Rw) and the polydispersity index of glutenin polymers (GPs) in wheat (Triticum aestivum). Here, using the same multi-location trials (4 years, 11 locations, and 192 cultivars), we report the factors that are associated with the conformation (Conf) of the polymers, which is the slope of Log(Rw) versus a function of Log(Mw). We found that Conf varied between 0.285 and 0.740, it had low broad-sense heritability (H 2=16.8), and it was significantly influenced by the temperature occurring over the last month of grain filling. Higher temperatures were found to increase Rw and the compactness and sphericity of GPs. Alleles for both high- and low-molecular-weight glutenin subunits had a significant influence on the Conf value. Assuming a Gaussian distribution for Mw, the number of polymers present in wheat grains was computed for different kernel weights and protein concentrations, and it was found to exceed 1012 GPs per grain. Using atomic force microscopy and cryo-TEM, images of GPs were obtained for the first time. Under higher average temperature, GPs became larger and more spherical and consequently less prone to rapid hydrolysis. We propose some orientations that could be aimed at potentially reducing the impact of numerous GPs on people suffering from non-celiac gluten sensitivity. [ABSTRACT FROM AUTHOR]

Lysine crotonylation (Kcr) is a novel post-translational modification and its function in plant salt-stress responses remains unclear. In this study, we performed the first comprehensive chloroplast crotonylome analysis of wheat seedling leaves to examine the potential functions of Kcr proteins in salt-stress responses. In a total of 471 chloroplast proteins, 1290 Kcr sites were identified as significantly regulated by salt stress, and the Kcr proteins were mainly involved in photosynthesis, protein folding, and ATP synthesis. The identified Kcr sites that responded to salt stress were concentrated within KcrK and KcrF motifs, with the conserved KcrF motif being identified in the Kcr proteins of wheat chloroplasts for the first time. Notably, 10 Kcr sites were identified in fructose-1,6-bisphosphate aldolase (TaFBA6), a key chloroplast metabolic enzyme involved in the Calvin–Benson cycle. Site-directed mutagenesis of TaFBA6 showed that the Kcr at K367 is critical in maintaining its enzymatic activity and in conferring salt tolerance in yeast. Further molecular dynamic simulations and analyses of surface electrostatic potential indicated that the Kcr at K367 could improve the structural stability of TaFBA6 by decreasing the distribution of positive charges on the protein surface to resist alkaline environments, thereby promoting both the activity of TaFBA6 and salt tolerance. [ABSTRACT FROM AUTHOR]

Solar-induced fluorescence (SIF) is a promising proxy for photosynthesis, but it is unclear whether it performs well in tracking the gross primary productivity (GPP) under different environmental conditions. In this study, we investigated the dynamics of the two parameters from October 2020 to June 2021 in field-grown winter wheat (Triticum aestivum) and found that the ability of SIF to track GPP was weakened at low temperatures. Accounting for the coupling of light and temperature at a seasonal scale, we found that SIF yield showed a lower temperature sensitivity and had a lower but broader optimal temperature range compared with light-use efficiency (LUE), although both SIF yield and LUE decreased in low-temperature conditions. The discrepancy between the temperature responses of SIF yield and GPP caused an increase in the ratio of SIF/GPP in winter, which indicated the variation in the relationship between them during this period. The results of our study highlight the impact of low temperature on the relationship between SIF and GPP and show the necessity of reconsidering the dynamics of energy distribution inside plants under changing environments. [ABSTRACT FROM AUTHOR]

Cadmium (Cd) is a highly toxic heavy metal that readily enters cereals, such as wheat, via the roots and is translocated to the shoots and grains, thereby posing high risks to human health. However, the vast and complex genome of allohexaploid wheat makes it challenging to understand Cd resistance and accumulation. In this study, a Cd-resistant cultivar of wheat, 'ZM1860', and a Cd-sensitive cultivar, 'ZM32', selected from a panel of 442 accessions, exhibited significantly different plant resistance and grain accumulation. We performed an integrated comparative analysis of the morpho-physiological traits, ionomic and phytohormone profiles, genomic variations, transcriptomic landscapes, and gene functionality in order to identify the mechanisms underlying these differences. Under Cd toxicity, 'ZM1860' outperformed 'ZM32', which showed more severe leaf chlorosis, poorer root architecture, higher accumulation of reactive oxygen species, and disordered phytohormone homeostasis. Ionomics showed that 'ZM32' had a higher root-to-shoot translocation coefficient of Cd and accumulated more Cd in the grains than 'ZM1860'. Whole-genome re-sequencing (WGS) and transcriptome sequencing identified numerous DNA variants and differentially expressed genes involved in abiotic stress responses and ion transport between the two genotypes. Combined ionomics, transcriptomics, and functional gene analysis identified the plasma membrane-localized heavy metal ATPase TaHMA2b-7A as a crucial Cd exporter regulating long-distance Cd translocation in wheat. WGS- and PCR-based analysis of sequence polymorphisms revealed a 25-bp InDel site in the promoter region of TaHMA2b-7A , and this was probably responsible for the differential expression. Our multiomics approach thus enabled the identification of a core transporter involved in long-distance Cd translocation in wheat, and it may provide an elite genetic resource for improving plant Cd resistance and reducing grain Cd accumulation in wheat and other cereal crops. [ABSTRACT FROM AUTHOR]

Wheat is one of the world's major staple food crops, and breeding for improvement of grain yield is a priority under the scenarios of climate change and population growth. WRKY transcription factors are multifaceted regulators in plant growth, development, and responses to environmental stimuli. In this study, we identify the WRKY gene TaGSNE (Grain Size and Number Enhancer) in common wheat, and find that it has relatively high expression in leaves and roots, and is induced by multiple abiotic stresses. Eleven single-nucleotide polymorphisms were identified in TaGSNE , forming two haplotypes in multiple germplasm collections, named as TaGSNE-Hap-1 and TaGSNE-Hap-2. In a range of different environments, TaGSNE-Hap-2 was significantly associated with increases in thousand-grain weight (TGW; 3.0%) and spikelet number per spike (4.1%), as well as with deeper roots (10.1%) and increased root dry weight (8.3%) at the mid-grain-filling stage, and these were confirmed in backcross introgression populations. Furthermore, transgenic rice lines overexpressing TaGSNE had larger panicles, more grains, increased grain size, and increased grain yield relative to the wild-type control. Analysis of geographic and temporal distributions revealed that TaGSNE-Hap-2 is positively selected in China and Pakistan, and TaGSNE-Hap-1 in Europe. Our findings demonstrate that TaGSNE overcomes the trade-off between TGW/grain size and grain number, leading us to conclude that these elite haplotypes and their functional markers could be utilized in marker-assisted selection for breeding high-yielding varieties. [ABSTRACT FROM AUTHOR]

Saline-alkali soil is a major environmental problem affecting crop productivity. One of the most effective approaches to combat it is to breed stress-tolerant plants through genetic engineering. Shanrong No. 4 (SR4) is an alkaline-tolerant cultivar of bread wheat (Triticum aestivum) derived from asymmetric somatic hybridization between the common wheat cultivar Jinan 177 (JN177) and tall wheatgrass. In this study, we aimed to explore the structure and function of alkalinity stress-responsive long non-coding RNAs (lncRNAs) in wheat. Sequencing was employed to identify the lncRNAs associated with stress tolerance and their corresponding targets. Approximately 19 000 novel lncRNA sequences were detected in SR4 and JN177. Upon exposure to alkaline stress, SR4 differentially expressed 5691 lncRNAs, whilst JN177 differentially expressed 5932. We selected five of them (L0760, L6247, L0208, L2098, and L3065) and generated seedlings of transiently knocked down strains using the virus-induced gene-silencing method. Knockdown of L0760 and L2098 caused the plants to exhibit sensitivity to alkaline stress, whereas knockdown of L6247, L0208, and L3065 increased the ability of plants to tolerate alkaline stress. We constructed lncRNA–miRNA–target-mRNA networks and alkali-response-related lncRNA–target-mRNA association networks to analyse the functions of lncRNAs. Collectively, our results demonstrate that lncRNAs may perform different roles under alkaline stress conditions. [ABSTRACT FROM AUTHOR]

Benzoxazinoids are specialized metabolites that are highly abundant in staple crops, such as maize and wheat. Although their biosynthesis has been studied for several decades, the regulatory mechanisms of the benzoxazinoid pathway remain unknown. Here, we report that the wheat transcription factor MYB31 functions as a regulator of benzoxazinoid biosynthesis genes. A transcriptomic analysis of tetraploid wheat (Triticum turgidum) tissue revealed the up-regulation of two TtMYB31 homoeologous genes upon aphid and caterpillar feeding. TaMYB31 gene silencing in the hexaploid wheat Triticum aestivum significantly reduced benzoxazinoid metabolite levels and led to susceptibility to herbivores. Thus, aphid progeny production, caterpillar body weight gain, and spider mite oviposition significantly increased in TaMYB31 -silenced plants. A comprehensive transcriptomic analysis of hexaploid wheat revealed that the TaMYB31 gene is co-expressed with the target benzoxazinoid - encoded Bx genes under several biotic and environmental conditions. Therefore, we analyzed the effect of abiotic stresses on benzoxazinoid levels and discovered a strong accumulation of these compounds in the leaves. The results of a dual fluorescence assay indicated that TaMYB31 binds to the Bx1 and Bx4 gene promoters, thereby activating the transcription of genes involved in the benzoxazinoid pathway. Our finding is the first report of the transcriptional regulation mechanism of the benzoxazinoid pathway in wheat. [ABSTRACT FROM AUTHOR]

Interannual and local fluctuations in wheat crop yield are mostly explained by abiotic constraints. Heatwaves and drought, which are among the top stressors, commonly co-occur, and their frequency is increasing with global climate change. High-throughput methods were optimized to phenotype wheat plants under controlled water deficit and high temperature, with the aim to identify phenotypic traits conferring adaptative stress responses. Wheat plants of 10 genotypes were grown in a fully automated plant facility under 25/18 °C day/night for 30 d, and then the temperature was increased for 7 d (38/31 °C day/night) while maintaining half of the plants well irrigated and half at 30% field capacity. Thermal and multispectral images and pot weights were registered twice daily. At the end of the experiment, key metabolites and enzyme activities from carbohydrate and antioxidant metabolism were quantified. Regression machine learning models were successfully established to predict plant biomass using image-extracted parameters. Evapotranspiration traits expressed significant genotype–environment interactions (G×E) when acclimatization to stress was continuously monitored. Consequently, transpiration efficiency was essential to maintain the balance between water-saving strategies and biomass production in wheat under water deficit and high temperature. Stress tolerance included changes in carbohydrate metabolism, particularly in the sucrolytic and glycolytic pathways, and in antioxidant metabolism. The observed genetic differences in sensitivity to high temperature and water deficit can be exploited in breeding programmes to improve wheat resilience to climate change. [ABSTRACT FROM AUTHOR]

Mesophyll conductance (g m) is an important factor limiting photosynthesis. However, g m response to long-term growth in variable [CO2] is not well understood, particularly in crop plants. Here, we grew two cultivars of wheat (Halberd and Cranbrook), known to differ in g m under current environmental conditions, in four [CO2] treatments: glacial (206 μmol mol−1), pre-industrial (344 μmol mol−1), current ambient (489 μmol mol−1), and super-elevated (1085 μmol mol−1), and two water treatments (well-watered and moderate water limitation), to develop an evolutionary and future climate perspective on g m control of photosynthesis and water-use efficiency (WUE). In the two wheat genotypes, g m increased with rising [CO2] from glacial to ambient [CO2], but declined at super-elevated [CO2]. The responses of g m to different growth [CO2] also depend on water stress; however, the specific mechanism of g m response to [CO2] remains unclear. Although g m and g m/ g sc (mesophyll conductance/stomatal conductance) were strongly associated with the variability of photosynthetic rates (A) and WUE, we found that plants with higher g m may increase A without increasing g sc, which increased WUE. These results may be useful to inform plant breeding programmes and cultivar selection for Australian wheat under future environmental conditions. [ABSTRACT FROM AUTHOR]

A multilocus genome-wide association study of a panel of 369 diverse wheat (Triticum aestivum) genotypes was carried out in order to examine the genetic basis of variations in nutrient mineral concentrations in the grains. The panel was grown under field conditions for three consecutive years and the concentrations of Ca, K, Mg, Mn, P, and S were determined. Wide ranges of natural variation were detected among the genotypes. Strong positive correlations were found among the minerals except for K, which showed negative correlation trends with the other minerals. Genetic association analysis detected 86 significant marker–trait associations (MTAs) underlying the natural variations in mineral concentrations in grains. The major MTA was detected on the long arm of chromosome 5A and showed a pleiotropic effect on Ca, K, Mg, Mn, and S. Further significant MTAs were distributed among the whole genome except for chromosomes 3D and 6D. We identified putative candidate genes that are potentially involved in metal uptake, transport, and assimilation, including TraesCS5A02G542600 on chromosome 5A, which was annotated as a Major Facilitator Superfamily transporter and acted on all the minerals except K. TraesCS5A02G542600 was highly expressed in seed coat, and to a lesser extent in the peduncle, awns, and lemma. Our results provide important insights into the genetic basis of enhancement of nutrient mineral concentrations that can help to inform future breeding studies in order to improve human nutrition. [ABSTRACT FROM AUTHOR]

Lipoxygenases (Loxs) are dioxygenases that play an important role in plant growth and defense. Loxs affect flour processing quality in common wheat (Triticum aestivum). We conducted a genome-wide association study (GWAS) that identified 306 significant single-nucleotide polymorphisms (SNPs) related to Lox activity in Chinese wheat accessions. Among them, a novel lipoxygenase-encoding (Lpx) gene, TaLpx-B4 , was detected on chromosome 3B in a biparental population. Analysis of mutant wheat lines induced using ethyl methanesulfonate confirmed the role of TaLpx-B4 in modulating Lox activity. A phylogenetic tree of various plant Lpx genes indicated the predominance of the 9-Lpx type in common wheat. Further analysis revealed conserved intron number, exon length, and motif number in the TaLpx gene family. GWAS, linkage mapping, and gene annotation collectively showed that 14 out of 29 annotated TaLpx genes played a critical role in regulating Lox activity in the Chinese wheat accessions. Transgenic wheat grains with knockdown of Lpx family genes by RNAi showed significantly lower Lox activity than the wild type. One TaLpx -RNAi line had significantly reduced starch content and dough stability, and thus possessed relatively superior biscuit quality in soft wheat. Further analysis of the transcriptome, lipid components, and other metabolites revealed that knockdown of TaLpx genes significantly increased biscuit quality via changes in unsaturated fatty acid content as well as in starch, sucrose, and galactose metabolism. Our results provide new insights into the role of the TaLpx gene family that will be beneficial in improving soft wheat flour quality. [ABSTRACT FROM AUTHOR]

Wheat (Triticum aestivum L.) productivity is severely reduced by high temperatures. Breeding of heat-tolerant cultivars can be achieved by identifying genes controlling physiological and agronomical traits when high temperatures occur and using these to select superior genotypes, but no gene underlying genetic variation for heat tolerance has previously been described. We advanced the positional cloning of qYDH.3BL , a quantitative trait locus (QTL) on bread wheat chromosome 3B associated with increased yield in hot and dry climates. The delimited genomic region contained 12 putative genes and a sequence variant in the promoter region of one gene, Seven in absentia , TaSINA. This was associated with the QTL's effects on early vigour, root growth, plant biomass, and yield components in two distinct wheat populations grown under various growth conditions. Near isogenic lines carrying the positive allele at qYDH.3BL underexpressed TaSINA and had increased vigour and water use efficiency early in development, as well as increased biomass, grain number, and grain weight following heat stress. A survey of worldwide distribution indicated that the positive allele became widespread from the 1950s through the CIMMYT wheat breeding programme but, to date, has been selected only in breeding programmes in Mexico and Australia. [ABSTRACT FROM AUTHOR]

Abiotic stress tolerance in plants is pivotal to increase yield stability, but its genetic basis is still poorly understood. To gain insight into the genetic architecture of frost tolerance, this work evaluated a large mapping population of 1739 wheat (Triticum aestivum L.) lines and hybrids adapted to Central Europe in field trials in Germany and fingerprinted the lines with a 9000 single-nucleotide polymorphism array. Additive effects prevailed over dominance effects. A two-dimensional genome scan revealed the presence of epistatic effects. Genome-wide association mapping in combination with a robust cross-validation strategy identified one frost tolerance locus with a major effect located on chromosome 5B. This locus was not in linkage disequilibrium with the known frost loci Fr-B1 and Fr-B2. The use of the detected diagnostic markers on chromosome 5B, however, does not allow prediction of frost tolerance with high accuracy. Application of genome-wide selection approaches that take into account also loci with small effect sizes considerably improved prediction of the genetic variation of frost tolerance in wheat. The developed prediction model is valuable for improving frost tolerance because this trait displays a wide variation in occurrence across years and is therefore a difficult target for conventional phenotypic selection.

A growing number of leaf traits can be estimated from hyperspectral reflectance data. These include structural and compositional traits, such as leaf mass per area (LMA) and nitrogen and chlorophyll content, but also physiological traits such a Rubisco carboxylation activity, electron transport rate, and respiration rate. Since physiological traits vary with leaf temperature, how does this impact on predictions made from reflectance measurements? We investigated this with two wheat varieties, by repeatedly measuring each leaf through a sequence of temperatures imposed by varying the air temperature in a growth room. Leaf temperatures ranging from 20 °C to 35 °C did not alter the estimated Rubisco capacity normalized to 25 °C (V cmax25), or chlorophyll or nitrogen contents per unit leaf area. Models estimating LMA and V cmax25/N were both slightly influenced by leaf temperature: estimated LMA increased by 0.27% °C–1 and V cmax25/N increased by 0.46% °C–1. A model estimating Rubisco activity closely followed variation associated with leaf temperature. Reflectance spectra change with leaf temperature and therefore contain a temperature signal. [ABSTRACT FROM AUTHOR]

The induced dwarf mutant Rht12 was previously shown to have agronomic potential to replace the conventional DELLA mutants Rht-B1b/Rht-D1b in wheat. The Rht12 dwarfing gene is not associated with reduced coleoptile length (unlike the DELLA mutants) and it is dominant, characteristics which are shared with the previously characterized dwarfing genes Rht18 and Rht14. Using the Rht18/Rht14 model, a gibberellin (GA) 2-oxidase gene was identified in the Rht12 region on chromosome 5A. A screen for suppressor mutants in the Rht12 background identified tall overgrowth individuals that were shown to contain loss-of-function mutations in GA2oxidaseA13 , demonstrating the role of this gene in the Rht12 dwarf phenotype. It was concluded that Rht12 , Rht18 , and Rht14 share the same height-reducing mechanism through the increased expression of GA 2-oxidase genes. Some of the overgrowth mutants generated in this study were semi-dwarf and taller than the original Rht12 dwarf, providing breeders with new sources of agronomically useful dwarfism. [ABSTRACT FROM AUTHOR]

Salt and drought-induced RING finger1 (SDIR1) is a RING-type E3 ubiquitin ligase that plays a key role in ABA-mediated responses to salinity and drought stress via the ubiquitination pathway in some plant species. However, its function in wheat (Triticum aestivum) is unknown. Here, we isolated a SDIR1 member in wheat, TaSDIR1-4A , and characterized its E3 ubiquitin ligase activity. DNA polymorphism assays showed the presence of two nucleotide variation sites in the promoter region of TaSDIR1-4A , leading to the detection of the haplotypes Hap-4A-1 and Hap-4A-2 in wheat populations. Association analysis showed that TaSDIR1-4A haplotypes were associated with 1000-grain weight (TGW) across a variety of different environments, including well-watered and heat-stress conditions. Genotypes with Hap-4A-2 had higher TGW than those with Hap-4A-1. Phenotypes in both gene-silenced wheat and transgenic Arabidopsis showed that TaSDIR1-4A was a negative regulator of grain size. Gene expression assays indicated that TaSDIR1-4A was most highly expressed in flag leaves, and expression was higher in Hap-4A-1 accessions than in Hap-4A-2 accessions. The difference might be attributable to the fact that TaERF3 (ethylene response factor) can act as a transcriptional repressor of TaSDIR1-4A in Hap-4A-2 but not in Hap-4A-1. Examination of modern wheat varieties shows that the favorable haplotype has been positively selected in breeding programs in China. The functional marker for TaSDIR1-4A developed in this study should be helpful for future wheat breeding. [ABSTRACT FROM AUTHOR]

Using a field to lab approach, mature deep-rooting traits in wheat were correlated to root phenotypes measured on young plants from controlled conditions. Mature deep-rooting root traits of 20 wheat genotypes at maturity were established via coring in three field trials across 2 years. Field traits were correlated to phenotypes expressed by the 20 genotypes after growth in four commonly used lab screens: (i) soil tubes for root emergence, elongation, length, and branching at four ages to 34 days after sowing (DAS); (ii) paper pouches 7 DAS and (iii) agar chambers for primary root (PR) number and angles at 8 DAS; and (iv) soil baskets for PR and nodal root (NR) number and angle at 42 DAS. Correlations between lab and field root traits (r 2=0.45–0.73) were highly inconsistent, with many traits uncorrelated and no one lab phenotype correlating similarly across three field experiments. Phenotypes most positively associated with deep field roots were: longest PR and NR axiles from the soil tube screen at 20 DAS; and narrow PR angle and wide NR angle from soil baskets at 42 DAS. Paper and agar PR angles were positively and significantly correlated to each other, but only wide outer PRs in the paper screen correlated positively to shallower field root traits. NR phenotypes in soil baskets were not predicted by PR phenotypes in any screen, suggesting independent developmental controls and value in measuring both root types in lab screens. Strong temporal and edaphic effects on mature root traits, and a lack of understanding of root trait changes during plant development, are major challenges in creating controlled-environment root screens for mature root traits in the field. [ABSTRACT FROM AUTHOR]

Molybdenum (Mo), which is an essential microelement for plant growth, plays important roles in multiple metabolic and physiological processes, including responses to drought and cold stress in wheat. Lipids also have crucial roles in plant adaptions to abiotic stresses. The aim of this study was to use glycerolipidomic and transcriptomic analyses to determine the changes in lipids induced by Mo that are associated with Mo-enhanced drought tolerance in wheat. Mo treatments increased the transcript levels of genes involved in fatty acid and glycerolipid biosynthesis and desaturation, but suppressed the expression of genes involved in oxylipin production. Wheat plants supplemented with Mo displayed higher contents of monogalactosyldiacyglycerol (MGDG), digalactosyldoacylglycerol (DGDG), phosphatidylglycerol (PG), phosphatidylethanolamine (PE), and phosphatidylcholine (PC) with increased levels of unsaturation. The levels of MGDG, DGDG, PG, and PC increased under PEG-simulated drought (PSD), and the magnitude of the responses varied in the presence and absence of Mo. Mo increased the accumulation of the most abundant glycerolipid species of C36:6, C34:4, and C34:3 by increasing the expression of genes related to desaturation under PSD, and this contributed to maintaining the fluidity of membranes. In addition, Mo attenuated the decreases in the ratios of DGDG/MGDG and PC/PE that were observed under PSD. These changes in lipids in Mo-treated wheat would contribute to maintaining the integrity of membranes and to protecting the photosynthetic apparatus, thus acting together to enhance drought tolerance. [ABSTRACT FROM AUTHOR]

One way to increase yield potential in wheat is screening for natural variation in photosynthesis. This study uses measured and modelled physiological parameters to explore genotypic diversity in photosynthetic capacity (P c, Rubisco carboxylation capacity per unit leaf area at 25 °C) and efficiency (P eff, P c per unit of leaf nitrogen) in wheat in relation to fertilizer, plant stage, and environment. Four experiments (Aus1, Aus2, Aus3, and Mex1) were carried out with diverse wheat collections to investigate genetic variation for Rubisco capacity (V cmax25), electron transport rate (J), CO2 assimilation rate, stomatal conductance, and complementary plant functional traits: leaf nitrogen, leaf dry mass per unit area, and SPAD. Genotypes for Aus1 and Aus2 were grown in the glasshouse with two fertilizer levels. Genotypes for Aus3 and Mex1 experiments were grown in the field in Australia and Mexico, respectively. Results showed that V cmax25 derived from gas exchange measurements is a robust parameter that does not depend on stomatal conductance and was positively correlated with Rubisco content measured in vitro. There was significant genotypic variation in most of the experiments for P c and P eff. Heritability of P c reached 0.7 and 0.9 for SPAD. Genotypic variation and heritability of traits show that there is scope for these traits to be used in pre-breeding programmes to improve photosynthesis with the ultimate objective of raising yield potential. [ABSTRACT FROM AUTHOR]

Wheat adaptation can be fine-tuned by earliness per se (Eps) genes. Although the effects of Eps genes are often assumed to act independently of the environment, previous studies have shown that they exhibit temperature sensitivity. The number of leaves and phyllochron are considered determinants of flowering time and the numerical components of yield include spikelets per spike and fertile floret number within spikelets. We studied the dynamics of leaf, spikelet, and floret development in near isogenic lines with either late or early alleles of Eps - D1 under seven temperature regimes. Leaf appearance dynamics were modulated by temperature, and Eps alleles had a greater effect on the period from flag leaf to heading than phyllochron. In addition, the effects of the Eps alleles on spikelets per spike were minor, and more related to spikelet plastochron than the duration of the early reproductive phase. However, fertile floret number was affected by the interaction between Eps alleles and temperature. So, at 9 °C, Eps -early alleles had more fertile florets than Eps -late alleles, at intermediate temperatures there was no significant difference, and at 18 °C (the highest temperature) the effect was reversed, with lines carrying the late allele producing more fertile florets. These effects were mediated through changes in floret survival; there were no clear effects on the maximum number of floret primordia. [ABSTRACT FROM AUTHOR]

Iron (Fe) is an essential micronutrient for all organisms. In crop plants, Fe deficiency can decrease crop yield significantly; however, our current understanding of how major crops respond to Fe deficiency remains limited. Herein, the effect of Fe deprivation at both the transcriptomic and metabolic level in hexaploid wheat was investigated. Genome-wide gene expression reprogramming was observed in wheat roots subjected to Fe starvation, with a total of 5854 genes differentially expressed. Homoeologue and subgenome-specific analysis unveiled the induction-biased contribution from the A and B genomes. In general, the predominance of genes coding for nicotianamine synthase, yellow stripe-like transporters, metal transporters, ABC transporters, and zinc-induced facilitator-like protein was noted. Expression of genes related to the Strategy II mode of Fe uptake was also predominant. Our transcriptomic data were in agreement with the GC-MS analysis that showed the enhanced accumulation of various metabolites such as fumarate, malonate, succinate, and xylofuranose, which could be contributing to Fe mobilization. Interestingly, Fe starvation leads to a significant temporal increase of glutathione S -transferase at both the transcriptional level and enzymatic activity level, which indicates the involvement of glutathione in response to Fe stress in wheat roots. Taken together, our result provides new insight into the wheat response to Fe starvation at the molecular level and lays the foundation to design new strategies for the improvement of Fe nutrition in crops. [ABSTRACT FROM AUTHOR]

Since starch is by far the major component of the mature wheat grain, it has been assumed that variation in the capacity for starch synthesis during grain filling can influence final grain weight. We investigated this assumption by studying a total of 54 wheat genotypes including elite varieties and landraces that were grown in two successive years in fields in the east of England. The weight, water content, sugars, starch, and maximum catalytic activities of two enzymes of starch biosynthesis, ADP-glucose pyrophosphorylase and soluble starch synthase, were measured during grain filling. The relationships between these variables and the weights and starch contents of mature grains were analysed. Final grain weight showed few or no significant correlations with enzyme activities, sugar levels, or starch content during grain filling, or with starch content at maturity. We conclude that neither sugar availability nor enzymatic capacity for starch synthesis during grain filling significantly influenced final grain weight in our field conditions. We suggest that final grain weight may be largely determined by developmental processes prior to grain filling. Starch accumulation then fills the grain to a physical limit set by developmental processes. This conclusion is in accord with those from previous studies in which source or sink strength has been artificially manipulated. [ABSTRACT FROM AUTHOR]

Plant peroxisomes are important components of cellular antioxidant networks, dealing with ROS generated by multiple metabolic pathways. Peroxisomes respond to environmental and cellular conditions by changing their size, number, and proteomic content. To investigate the role of peroxisomes in response to drought, dehydration and ABA treatment we took an evolutionary and comparative genomics approach. Colonisation of land required evolution of dehydration tolerance in the absence of subsequent anatomical adaptations. Therefore, the model bryophyte Physcomitrella patens, the model dicot Arabidopsis thaliana and wheat (Tricitcum aestivum), a globally important cereal crop were compared. Three sets of genes namely ‘ PTS1 genes’ (a proxy for genes encoding peroxisome targeted proteins), PEX genes (involved in peroxisome biogenesis) and genes involved in plant antioxidant networks were identified in all 3 species and their expression compared under drought (dehydration) and ABA treatment. Genes encoding enzymes of β-oxidation and gluconeogenesis, antioxidant enzymes including catalase and glutathione reductase and PEX3 and PEX11 isoforms showed conserved up-regulation, and peroxisome proliferation was induced by ABA in moss. Interestingly, expression of some of these genes differed between drought sensitive and resistant genotypes of wheat in line with measured photosynthetic and biochemical differences. These results point to an underappreciated role for peroxisomes in drought response. [ABSTRACT FROM AUTHOR]

Gradients exist in the distribution of storage proteins in the wheat (Triticum aestivum) endosperm and determine the milling properties and protein recovery rate of the grain. A novel image analysis technique was developed to quantify both the gradients in protein concentration, and the size distribution of protein bodies within the endosperm of wheat plants grown under two different (20 or 28 °C) post-anthesis temperatures, and supplied with a nutrient solution with either high or low nitrogen content. Under all treatment combinations, protein concentration was greater in the endosperm cells closest to the aleurone layer and decreased towards the centre of the two lobes of the grain, i.e. a negative gradient. This was accompanied by a decrease in size of protein bodies from the outer to the inner endosperm layers in all but one of the treatments. Elevated post-anthesis temperature had the effect of increasing the magnitude of the negative gradients in both protein concentration and protein body size, whilst limiting nitrogen supply decreased the gradients. [ABSTRACT FROM AUTHOR]

Plant phloem-based defence (PBD) against phloem-feeding insects is characteristic of the sieve occlusion by phloem lectins and β-1,3-glucan callose, both of which are produced under regulation by ethylene and MYB transcription factors. Wheat PBD requires β-1,3-glucan synthase-like proteins GSL2, GSL10, and GSL12, and may also require insect-resistant mannose-binding lectins Hfr-1 and Wci-1, which can accumulate in the phloem upon aphid feeding. This study elucidates whether any of the 73 MYB genes identified previously in the common wheat Triticum aestivum genome plays a role in wheat PBD activation with regard to the GSLs and lectins. Wheat MYB genes TaMYB19, TaMYB29, and TaMYB44 are highly activated in response to infestation of English grain aphid, and their silencing facilitates aphid feeding on wheat phloem and represses wheat PBD responses. Repressed PBD is shown to decrease aphid-induced callose deposition in wheat leaf epidermis and decrease aphid-induced expression of genes GSL2, GSL10, GSL12, Hfr-1, and Wci-1 in wheat leaf tissues. Based on single gene silencing effects, TaMYB19, TaMYB29, and TaMYB44 contribute 55-82% of PBD responses. However, the contributions of TaMYB genes to PBD are eliminated by ethylene signalling inhibitors, while simultaneous silencing of the three TaMYB genes cancels the tested PBD responses. Therefore, TaMYB19, TaMYB29, and TaMYB44 are co-regulators of wheat PBD and execute this function through crosstalk with the ethylene signalling pathway. [ABSTRACT FROM AUTHOR]

The tiller inhibition gene (tin) that reduces tillering in wheat (Triticum aestivum) is also associated with large spikes, increased grain weight, and thick leaves and stems. In this study, comparison of near-isogenic lines (NILs) revealed changes in stem morphology, cell wall composition, and stem strength. Microscopic analysis of stem cross-sections and chemical analysis of stem tissue indicated that cell walls in tin lines were thicker and more lignified than in freetillering NILs. Increased lignification was associated with stronger stems in tin plants. A candidate gene for tin was identified through map-based cloning and was predicted to encode a cellulose synthase-like (Csl) protein with homology to members of the CslA clade. Dinucleotide repeat-length polymorphism in the 5'UTR region of the Csl gene was associated with tiller number in diverse wheat germplasm and linked to expression differences of Csl transcripts between NILs. We propose that regulation of Csl transcript and/or protein levels affects carbon partitioning throughout the plant, which plays a key role in the tin phenotype. [ABSTRACT FROM AUTHOR]

Low phytic acid is a trait desired in cereal crops and can be achieved by manipulating the genes involved either in its biosynthesis or its transport in the vacuoles. Previously, we have demonstrated that the wheat TaABCC13 protein is a functional transporter, primarily involved in heavy metal tolerance, and a probable candidate gene to achieve low phytate wheat. In the current study, RNA silencing was used to knockdown the expression of TaABCC13 in order to evaluate its functional importance in wheat. Transgenic plants with significantly reduced TaABCC13 transcripts in either seeds or roots were selected for further studies. Homozygous RNAi lines K1B4 and K4G7 exhibited 34–22% reduction of the phytic acid content in the mature grains (T4 seeds). These transgenic lines were defective for spike development, as characterized by reduced grain filling and numbers of spikelets. The seeds of transgenic wheat had delayed germination, but the viability of the seedlings was unaffected. Interestingly, early emergence of lateral roots was observed in TaABCC13-silenced lines as compared to non-transgenic lines. In addition, these lines also had defects in metal uptake and development of lateral roots in the presence of cadmium stress. Our results suggest roles of TaABCC13 in lateral root initiation and enhanced sensitivity towards heavy metals. Taken together, these data demonstrate that wheat ABCC13 is functionally important for grain development and plays an important role during detoxification of heavy metals. [ABSTRACT FROM AUTHOR]

The capture of subsoil water by wheat roots can make a valuable contribution to grain yield on deep soils. More extensive root systems can capture more water, but leave the soil in a drier state, potentially limiting water availability to subsequent crops. To evaluate the importance of these legacy effects, a long-term simulation analysis at eight sites in the semi-arid environment of Australia compared the yield of standard wheat cultivars with cultivars that were (i) modified to have root systems which extract more water at depth and/or (ii) sown earlier to increase the duration of the vegetative period and hence rooting depth. We compared simulations with and without annual resetting of soil water to investigate the legacy effects of drier subsoils related to modified root systems. Simulated mean yield benefits from modified root systems declined from 0.1-0.6 t ha-1 when annually reset, to 0-0.2 t ha-1 in the continuous simulation due to a legacy of drier soils (mean 0-32 mm) at subsequent crop sowing. For continuous simulations, predicted yield benefits of >0.2 t ha-1 from more extensive root systems were rare (3-10% of years) at sites with shallow soils (<1.0 m), but occurred in 14-44% of years at sites with deeper soils (1.6-2.5 m). Earlier sowing had a larger impact than modified root systems on water uptake (14-31 vs 2-17 mm) and mean yield increase (up to 0.7 vs 0-0.2 t ha-1) and the benefits occurred on deep and shallow soils and in more years (9-79 vs 3-44%). Increasing the proportion of crops in the sequence which dry the subsoil extensively has implications for the farming system productivity, and the crop sequence must be managed tactically to optimize overall system benefits. [ABSTRACT FROM AUTHOR]

Earliness per se (Eps) genes account for the variation in flowering time when vernalization and photoperiod requirements are satisfied. Genomics and bioinformatics approaches were used to describe allelic variation for 40 Triticum aestivum genes predicted, by synteny with Brachypodium distachyon, to be in the 1DL Eps region. Re-sequencing 1DL genes revealed that varieties carrying early heading alleles at this locus, Spark and Cadenza, carry a subtelomeric deletion including several genes. The equivalent region in Rialto and Avalon is intact. A bimodal distribution in the segregating Spark X Rialto single seed descent (SSD) populations enabled the 1DL QTL to be defined as a discrete Mendelian factor, which we named Eps-D1. Near isogenic lines (NILs) and NIL derived key recombinants between markers flanking Eps-D1 suggest that the 1DL deletion contains the gene(s) underlying Eps-D1. The deletion spans the equivalent of the Triticum monoccocum Eps-Am1 locus, and hence includes MODIFIER OF TRANSCRIPTION 1 (MOT1) and FTSH PROTEASE 4 (FTSH4), the candidates for Eps-Am1. The deletion also contains T. aestivum EARLY FLOWERING 3-D1 (TaELF3-D1) a homologue of the Arabidopsis thaliana circadian clock gene EARLY FLOWERING 3. Eps-D1 is possibly a homologue of Eps-B1 on chromosome 1BL. NILs carrying the Eps-D1 deletion have significantly reduced total TaELF3 expression and altered TaGIGANTEA (TaGI) expression compared with wild type. Altered TaGI expression is consistent with an ELF3 mutant, hence we propose TaELF3-D1 as the more likely candidate for Eps-D1. This is the first direct fine mapping of Eps effect in bread wheat. [ABSTRACT FROM AUTHOR]

Considerable progress has been made in understanding the roles of AGC kinases in mammalian systems. However, very little is known about the roles of AGC kinases in wheat (Triticum aestivum). The necrotrophic fungus Rhizoctonia cerealis is the major pathogen of the destructive disease sharp eyespot of wheat. In this study, the wheat AGC kinase gene TaAGC1, responding to R. cerealis infection, was isolated, and its properties and role in wheat defence were characterized. R. cerealis-resistant wheat lines expressed TaAGC1 at higher levels than susceptible wheat lines. Sequence and phylogenetic analyses showed that the TaAGC1 protein is a serine/threonine kinase belonging to the NDR (nuclear Dbf2-related) subgroup of AGC kinases. Kinase activity assays proved that TaAGC1 is a functional kinase and the Asp-239 residue located in the conserved serine/threonine kinase domain of TaAGC1 is required for the kinase activity. Subcellular localization assays indicated that TaAGC1 localized in the cytoplasm and nucleus. Virus-induced TaAGC1 silencing revealed that the down-regulation of TaAGC1 transcripts significantly impaired wheat resistance to R. cerealis. The molecular characterization and responses of TaAGC1 overexpressing transgenic wheat plants indicated that TaAGC1 overexpression significantly enhanced resistance to sharp eyespot and reduced the accumulation of reactive oxygen species (ROS) in wheat plants challenged with R. cerealis. Furthermore, ROS-scavenging and certain defence-associated genes were up-regulated in resistant plants overexpressing TaAGC1 but down-regulated in susceptible knock-down plants. These results suggested that the kinase TaAGC1 positively contributes to wheat immunity to R. cerealis through regulating expression of ROS-related and defence-associated genes. [ABSTRACT FROM AUTHOR]

The high potential for an acrocentric chromosome originated from a complex reorganization of chromosomes 1HchS and 6HchS from Hordeum chilense in the development of hybrid wheat technology.Cytoplasmic male sterility (CMS) results from incompatibility between nuclear and cytoplasmic genomes, and is characterized by the inability to produce viable pollen. The restoration of male fertility generally involves the introgression of nuclear genes, termed restorers of fertility (Rf). CMS has been widely used for hybrid seed production in many crops but not in wheat, partly owing to the complex genetics of fertility restoration. In this study, an acrocentric chromosome that restores pollen fertility of CMS wheat in Hordeum chilense cytoplasm (msH1 system) is studied. The results show that this chromosome, of H. chilense origin and named Hchac, originated from a complex reorganization of the short arm of chromosomes 1Hch (1HchS) and 6Hch (6HchS). Diversity arrays technology (DArT) markers and cytological analysis indicate that Hchac is a kind of `zebra-like′ chromosome composed of chromosome 1HchS and alternate fragments of interstitial and distal regions of chromosome 6HchS. PCR-based markers together with FISH, GISH, and meiotic pairing analysis support this result. A restorer of fertility gene, named Rf6HchS, has been identified on the short arm of chromosome 6HchS. Moreover, restoration by the addition of chromosome 1HchS has been observed at a very low frequency and under certain environmental conditions. Therefore, the results indicate the presence of two Rf genes on the acrocentric chromosome: Rf6HchS and Rf1HchS, the restoration potential of Rf6HchS being greater. The stable and high restoration of pollen fertility in the msH1 system is therefore the result of the interaction between these two restorer genes. [ABSTRACT FROM AUTHOR]

Iron insufficiency is a worldwide problem in human diets. In cereals like wheat, the bran layer of the grains is an important source of iron. However, the dietary availability of iron in wheat flour is limited due to the loss of the iron-rich bran during milling and processing and the presence of anti-nutrients like phytic acid that keep iron strongly chelated in the grain. The present study investigated the localization of iron and phosphorus in grain tissues of wheat genotypes with contrasting grain iron content using synchrotron-based micro-X-ray fluorescence (micro-XRF) and micro-proton-induced X-ray emission (micro-PIXE). X-ray absorption near-edge spectroscopy (XANES) was employed to determine the proportion of divalent and trivalent forms of Fe in the grains. It revealed the abundance of oxygen, phosphorus, and sulphur in the local chemical environment of Fe in grains, as Fe-O-P-R and Fe-O-S-R coordination. Contrasting differences were noticed in tissue-specific relative localization of Fe, P, and S among the different genotypes, suggesting a possible effect of localization pattern on iron bioavailability. The current study reports the shift in iron distribution from maternal to filial tissues of grains during the evolution of wheat from its wild relatives to the present-day cultivated varieties, and thus suggests the value of detailed physical localization studies in varietal improvement programmes for food crops. [ABSTRACT FROM AUTHOR]

The green area displayed by a crop is a good indicator of its photosynthetic capacity, while chlorophyll retention or ‘stay-green’ is regarded as a key indicator of stress adaptation. Remote-sensing methods were tested to estimate these parameters in diverse wheat genotypes under different growing conditions. Two wheat populations (a diverse set of 294 advanced lines and a recombinant inbred line population of 169 sister lines derived from the cross between Seri and Babax) were grown in Mexico under three environments: drought, heat, and heat combined with drought. In the two populations studied here, a moderate heritable expression of stay-green was found–when the normalized difference vegetation index (NDVI) at physiological maturity was estimated using the regression of NDVI over time from the mid-stages of grain-filling to physiological maturity–and for the rate of senescence during the same period. Under heat and heat combined with drought environments, stay-green calculated as NDVI at physiological maturity and the rate of senescence, showed positive and negative correlations with yield, respectively. Moreover, stay-green calculated as an estimation of NDVI at physiological maturity and the rate of senescence regressed on degree days give an independent measurement of stay-green without the confounding effect of phenology. On average, in both populations under heat and heat combined with drought environments CTgf and stay-green variables accounted for around 30% of yield variability in multiple regression analysis. It is concluded that stay-green traits may provide cumulative effects, together with other traits, to improve adaptation under stress further. [ABSTRACT FROM PUBLISHER]

MYB transcription factors (TFs) play pivotal roles in the abiotic stress response in plants, but their characteristics and functions in wheat (Triticum aestivum L.) have not been fully investigated. A novel wheat MYB TF gene, TaMYB73, is reported here based on the observation that its targeting probe showed the highest salinity-inducibility level among all probes annotated as MYB TFs in the cDNA microarray. TaMYB73 is a R2R3 type MYB protein with transactivation activity, and binds with types I, II, and IIG MYB binding motifs. The gene was induced by NaCl, dehydration, and several phytohormones, as well as some stress-, ABA-, and GA-responsive cis-elements present in its promoter region. Its over-expression in Arabidopsis enhanced the tolerance to NaCl as well as to LiCl and KCl, whereas it had no contribution to mannitol tolerance. The over-expression lines had superior germination ability under NaCl and ABA treatments. The expression of many stress signalling genes such as AtCBF3 and AtABF3, as well as downstream responsive genes such as AtRD29A and AtRD29B, was improved in these over-expression lines, and TaMYB73 can bind with promoter sequences of AtCBF3 and AtABF3. Taken together, it is suggested that TaMYB73, a novel MYB transcription factor gene, participates in salinity tolerance based on improved ionic resistance partly via the regulation of stress-responsive genes. [ABSTRACT FROM PUBLISHER]

WRKY proteins are transcription factors involved in many plant processes including plant responses to pathogens. Here, the cross activity of TaWRKY78 from the monocot wheat and AtWRKY20 from the dicot Arabidopsis on the cognate promoters of the orthologous PR4-type genes wPR4e and AtHEL of wheat and Arabidopsis, respectively, was investigated. In vitro analysis showed the ability of TaWRKY78 to bind a –17/+80 region of the wPR4e promoter, containing one cis-acting W-box. Moreover, transient expression analysis performed on both TaWRKY78 and AtWRKY20 showed their ability to recognize the cognate cis-acting elements present in the wPR4e and AtHEL promoters, respectively. Finally, this paper provides evidence that both transcription factors are able to cross-regulate the orthologous PR4 genes with an efficiency slightly lower than that exerted on the cognate promoters. The observation that orthologous genes are subjected to similar transcriptional control by orthologous transcription factors demonstrates that the terminal stages of signal transduction pathways leading to defence are conserved and suggests a fundamental role of PR4 genes in plant defence. Moreover, these results corroborate the hypothesis that gene orthology imply similar gene function and that diversification between monocot and dicot has most likely occurred after the specialization of WRKY function. [ABSTRACT FROM PUBLISHER]

O2 deficiency during soil waterlogging inhibits respiration in roots, resulting in severe energy deficits. Decreased root-to-shoot ratio and suboptimal functioning of the roots, result in nutrient deficiencies in the shoots. In N2-flushed nutrient solutions, wheat seminal roots cease growth, while newly formed adventitious roots develop aerenchyma, and grow, albeit to a restricted length. When reliant on an internal O2 supply from the shoot, nutrient uptake by adventitious roots was inhibited less than in seminal roots. Epidermal and cortical cells are likely to receive sufficient O2 for oxidative phosphorylation and ion transport. By contrast, stelar hypoxia–anoxia can develop so that H+-ATPases in the xylem parenchyma would be inhibited; the diminished H+ gradients and depolarized membranes inhibit secondary energy-dependent ion transport and channel conductances. Thus, the presence of two transport steps, one in the epidermis and cortex to accumulate ions from the solution and another in the stele to load ions into the xylem, is important for understanding the inhibitory effects of root zone hypoxia on nutrient acquisition and xylem transport, as well as the regulation of delivery to the shoots of unwanted ions, such as Na+. Improvement of waterlogging tolerance in wheat will require an increased capacity for root growth, and more efficient root functioning, when in anaerobic media. [ABSTRACT FROM PUBLISHER]

Intimate relationships exist between form and function of plants, determining many processes governing their growth and development. However, in most crop simulation models that have been created to simulate plant growth and, for example, predict biomass production, plant structure has been neglected. In this study, a detailed simulation model of growth and development of spring wheat (Triticum aestivum) is presented, which integrates degree of tillering and canopy architecture with organ-level light interception, photosynthesis, and dry-matter partitioning. An existing spatially explicit 3D architectural model of wheat development was extended with routines for organ-level microclimate, photosynthesis, assimilate distribution within the plant structure according to organ demands, and organ growth and development. Outgrowth of tiller buds was made dependent on the ratio between assimilate supply and demand of the plants. Organ-level photosynthesis, biomass production, and bud outgrowth were simulated satisfactorily. However, to improve crop simulation results more efforts are needed mechanistically to model other major plant physiological processes such as nitrogen uptake and distribution, tiller death, and leaf senescence. Nevertheless, the work presented here is a significant step forwards towards a mechanistic functional–structural plant model, which integrates plant architecture with key plant processes. [ABSTRACT FROM PUBLISHER]

Grain weight is one of the most important components of cereal yield and quality. A clearer understanding of the physiological and molecular determinants of this complex trait would provide an insight into the potential benefits for plant breeding. In the present study, the dynamics of dry matter accumulation, water uptake, and grain size in parallel with the expression of expansins during grain growth in wheat were analysed. The stabilized water content of grains showed a strong association with final grain weight (r2=0.88, P <0.01). Grain length was found to be the trait that best correlated with final grain weight (r2=0.98, P <0.01) and volume (r2=0.94, P <0.01). The main events that defined final grain weight occurred during the first third of grain-filling when maternal tissues (the pericarp of grains) undergo considerable expansion. Eight expansin coding sequences were isolated from pericarp RNA and the temporal profiles of accumulation of these transcripts were monitored. Sequences showing high homology with TaExpA6 were notably abundant during early grain expansion and declined as maturity was reached. RNA in situ hybridization studies revealed that the transcript for TaExpA6 was principally found in the pericarp during early growth in grain development and, subsequently, in both the endosperm and pericarp. The signal in these images is likely to be the sum of the transcript levels of all three sequences with high similarity to the TaExpA6 gene. The early part of the expression profile of this putative expansin gene correlates well with the critical periods of early grain expansion, suggesting it as a possible factor in the final determination of grain size. [ABSTRACT FROM PUBLISHER]

Invertases cleave sucrose in glucose and fructose, using water as an acceptor. Fructosyltransferases catalyse the transfer of a fructosyl residue between sucrose and/or fructan molecules. Plant fructosyltransferases (FTs) evolved from vacuolar invertases by small mutational changes, leading to differences in substrate specificity. The S-type of enzymes (invertases, sucrose:sucrose 1-fructosyltransferases or 1-SSTs, and sucrose:fructan 6-fructosyltransferases or 6-SFTs) prefer sucrose as the donor substrate while F-type enzymes (fructan:fructan 1-fructosyltransferases or 1-FFTs and fructan:fructan 6G-fructosyltransferases or 6G-FFTs) preferentially use fructan as the donor substrate. Recently, a functional Asp/Arg or Asp/Lys couple in the Hypervariable Loop (HVL) was suggested to be essential to keep Asp in a favourable orientation for binding sucrose as the donor substrate in S-type enzymes. However, the F-type enzyme 1-FFT of Triticum aestivum (Ta1-FFT) also contains the Asp/Arg couple in the HVL, although it prefers fructan as the donor substrate. In this paper, mutagenesis studies on Ta1-FFT are presented. In Ta1-SST, Tyr282 (the Asp281 homologue) seems to be essential in creating a tight H-bond Network (HBN) in which the Arg-residue of the Asp/Arg couple is held in a fixed position. This tight HBN is disrupted in Ta1-FFT, leading to a more flexible Arg-residue and a dysfunctional Asp/Arg couple. A single D281Y mutation in Ta1-FFT restored the tight HBN and introduced typical S-type characteristics. Conclusively, in wheat FTs Asp281 (and its homologues) is involved in donor substrate specificity. [ABSTRACT FROM PUBLISHER]

Most modern wheat cultivars contain major dwarfing genes, but their effects on root growth are unclear. Near-isogenic lines (NILs) containing Rht-B1b, Rht-D1b, Rht-B1c, Rht8c, Rht-D1c, and Rht12 were used to characterize the effects of semi-dwarfing and dwarfing alleles on root growth of ‘Mercia’ and ‘Maris Widgeon’ wheat cultivars. Wheat seedlings were grown in gel chambers, soil-filled columns, and in the field. Roots were extracted and length and dry mass measured. No significant differences in root length were found between semi-dwarfing lines and the control lines in any experiment, nor was there a significant difference between the root lengths of the two cultivars grown in the field. Total root length of the dwarf lines (Rht-B1c, Rht-D1c, and Rht12) was significantly different from that of the control although the effect was dependent on the experimental methodology; in gel chambers root length of dwarfing lines was increased by ∼40% while in both soil media it was decreased (by 24–33%). Root dry mass was 22–30% of the total dry mass in the soil-filled column and field experiments. Root length increased proportionally with grain mass, which varied between NILs, so grain mass was a covariate for the analysis of variance. Although total root length was altered by dwarf lines, root architecture (average root diameter, lateral root:total root ratio) was not affected by reduced height alleles. A direct effect of dwarfing alleles on root growth during seedling establishment, rather than a secondary partitioning effect, was suggested by the present experiments. [ABSTRACT FROM PUBLISHER]

The movement of water into harvest-ripe grains of dormant and non-dormant genotypes of wheat (Triticum aestivum L.) was investigated using Magnetic Resonance Micro-Imaging (MRMI). Images of virtual sections, both longitudinal and transverse, throughout the grain were collected at intervals after the start of imbibition and used to reconstruct a picture of water location within the different grain tissues and changes over time. The observations were supplemented by the weighing measurements of water content and imbibition of grains in water containing I2/KI which stains starch and lipid, thereby acting as a marker for water. In closely related genotypes, with either a dormant or a non-dormant phenotype, neither the rate of increase in water content nor the pattern of water distribution within the grain was significantly different until 18 h, when germination became apparent in the non-dormant genotype. Water entered the embryo and scutellum during the very early stages of imbibition through the micropyle and by 2 h water was clearly evident in the micropyle channel. After 12 h of imbibition, embryo structures such as the coleoptile and radicle were clearly distinguished. Although water accumulated between the inner (seed coat) and outer (pericarp) layers of the coat surrounding the grain, there was no evidence for movement of water directly across the coat and into the underlying starchy endosperm. [ABSTRACT FROM PUBLISHER]

In higher plants there are two forms of ferredoxin NADP+ oxidoreductase (FNR), a photosynthetic pFNR primarily required for the photoreduction of NADP+, and a heterotrophic hFNR which generates reduced ferredoxin by utilizing electrons from NADPH produced during carbohydrate oxidation. The aim of this study was to investigate the presence of multiple forms of FNR in wheat leaves and the capacity of FNR isoforms to respond to changes in reductant demand through varied expression and N-terminal processing. Two forms of pFNR mRNA (pFNRI and pFNRII) were expressed in a similar pattern along the 12 cm developing primary wheat leaf, with the highest levels observed in plants grown continuously in the dark in the presence (pFNRI) or absence (pFNRII) of nitrate respectively. pFNR protein increased from the leaf base to tip. hFNR mRNA and protein was in the basal part of the leaf in plants grown in the presence of nitrate. FNR activity in plants grown in a light/dark cycle without nitrate was mainly due to pFNR, whilst hFNR contributed significantly in nitrate-fed plants. The potential role of distinct forms of FNR in meeting the changing metabolic capacity and reductant demands along the linear gradient of developing cells of the leaf are discussed. Furthermore, evidence for alternative N-terminal cleavage sites of pFNR acting as a means of discriminating between ferredoxins and the implications of this in providing a more effective flow of electrons through a particular pathway in vivo is considered. [ABSTRACT FROM PUBLISHER]

In this study, a pathogen-inducible ERF (ethylene-response factor) gene in wheat, designated TaERF3, was isolated and characterized in detail. The sequence of the TaERF3 protein possesses all of the traits commonly associated with ERFs, but its entire sequence shares low identity with other ERFs of transcription factor families. The results of assays on subcelluar localization, GCC box-binding ability, and transactivation activity indicated that TaERF3 is a nuclear targeting protein and functions as a GCC box-binding transcriptional activator. Following infection with Blumeria graminis, the induction peak of TaERF3 expression occurring at 12 h in the resistant line was about six times higher than that in its susceptible parent. Following infections with Fusarium graminearum or Rhizoctonia cerealis, the TaERF3 maximum inductions in the susceptible line occurring at 12 h were about three or six times higher than those in the resistant lines, whereas after 24 h or 48 h, the transcript inductions in the resistant lines were much higher than that in the susceptible line. Furthermore, the TaERF3 transcript peak induced by salicylic acid (SA) treatment occurred at 4 h, whereas the peaks induced by exogenous ethylene and methyl jasmonate (MeJA) occurred at 24 h, all of which were earlier than those induced by pathogens in the resistant lines. These results suggested that TaERF3 might be mainly involved in the active defence response to B. graminis at an earlier stage through SA signalling, and to F. graminearum and R. cerealis at a later stage through the ethylene/jasmonic acid signalling pathways. [ABSTRACT FROM PUBLISHER]