Summary Dissection of the genetic basis of wheat ionome is crucial for understanding the physiological and biochemical processes underlying mineral accumulation in seeds, as well as for efficient crop breeding. Most of the elements essential for plants are metals stored in seeds as chelate complexes with phytic acid or sulfur-containing compounds. We assume that the involvement of phosphorus and sulfur in metal chelation is the reason for strong phenotypic correlations within ionome. Adjustment of element concentrations for the effect of variation in phosphorus and sulfur seed content resulted in drastic change of phenotypic correlations between the elements. The genetic architecture of wheat grain ionome was characterized by quantitative trait loci (QTL) analysis using a cross between durum and wild emmer wheat. QTL analysis of the adjusted traits and two-trait analysis of the initial traits paired with either P or S considerably improved QTL detection power and accuracy, resulting in the identification of 105 QTLs and 617 QTL effects for 11 elements. Candidate gene search revealed some potential functional associations between QTLs and corresponding genes within their intervals. Thus, we have shown that accounting for variation in P and S is crucial for understanding of the physiological and genetic regulation of mineral composition of wheat grain ionome and can be implemented for other plants.

Tef (Eragrostis tef), a staple crop that originated in the Horn of Africa, has been introduced to multiple countries over the last several decades. Crop cultivation in new geographic regions raises questions regarding the molecular basis for biotic stress responses. In this study, we aimed to classify the insect abundance on tef crop in Israel, and to elucidate its chemical and physical defense mechanisms in response to insect feeding. To discover the main pests of tef in the Mediterranean climate, we conducted an insect field survey on three selected accessions named RTC-144, RTC-405, and RTC-406, and discovered that the most abundant insect order is Hemiptera. We compared the differences in Rhopalosiphum padi (Hemiptera; Aphididae) aphid performance, preference, and feeding behavior between the three accessions. While the number of aphid progeny was lower on RTC-406 than on the other two, the aphid olfactory assay indicated that the aphids tended to be repelled from the RTC-144 accession. To highlight the variation in defense responses, we investigated the physical and chemical mechanisms. As a physical barrier, the density of non-granular trichomes was evaluated, in which a higher number of trichomes on the RTC-406 than on the other accessions was observed. This was negatively correlated with aphid performance. To determine chemical responses, the volatile and central metabolite profiles were measured upon aphid attack for 4 days. The volatile analysis exposed a rich and dynamic metabolic profile, and the central metabolism profile indicated that tef plants adjust their sugars and organic and amino acid levels. Overall, we found that the tef plants possess similar defense responses as other Poaceae family species, while the non-volatile deterrent compounds are yet to be characterized. A transcriptomic time-series analysis of a selected accession RTC-144 infested with aphids revealed a massive alteration of genes related to specialized metabolism that potentially synthesize non-volatile toxic compounds. This is the first report to reveal the variation in the defense mechanisms of tef plants. These findings can facilitate the discovery of insect-resistance genes leading to enhanced yield in tef and other cereal crops.

Pisum fulvum is an annual legume native to Syria, Lebanon, Israel and Jordan. In certain locations, P. fulvum individuals were documented to display a reproductive dimorphism – amphicarpy, with both above and below ground flowers and pods. Herein we aimed to study the possible role of soil texture on amphicarpy in P. fulvum, to investigate the possible bio-climatic associations of P. fulvum amphicarpy and to identify genetic markers associated with this phenotype. A set of 127 germplasm accessions sampled across the Israeli distribution range of the species was phenotyped in two common garden nurseries. Land use and bioclimatic data were used to delineate the eco-geographic clustering of accession's sampling sites. Single nucleotide polymorphism (SNP) markers were employed in genome-wide association study to identify associated loci. Amphicarpy was subject to strong experimental site x genotype interaction with higher phenotypic expression in fine textured soil relative to sandy loam. Amphicarpy was more prevalent among accessions sampled in eastern Judea and Samaria and was weakly associated with early phenology and relatively modest above ground biomass production. Twelve SNP markers were significantly associated with amphicarpy, each explaining between 8 and 12 % of the phenotypic variation. In P. fulvum amphicarpy seems to be a polygenetic trait controlled by an array of genes that is likely to be affected by environmental stimuli. The probable selective advantage of the association between amphicarpy and early flowering is in line with its relative prevalence in drought prone territories subject to heavy grazing.

Worldwide demand for tef (Eragrostis tef) as a functional food for human consumption is increasing, thanks to its nutritional benefits and gluten-free properties. As a result, tef in now grown outside its native environment in Ethiopia and thus information is required regarding plant nutrition demands in these areas, as well as resulting grain health-related composition. In the current work, two tef genotypes were grown in Israel under irrigation in two platforms, plots in the field and pots in a greenhouse, with four and five nitrogen treatments, respectively. Nutritional and health-related quality traits were analyzed, including mineral content, fatty acid composition, hydrophilic and lipophilic antioxidative capacity, total phenolic content and basic polyphenolic profile. Our results show that tef genotypes differ in their nutritional composition, e.g. higher phenolic contents in the brown compared to the white genotype. Additionally, nitrogen availability positively affected grain fatty acid composition and iron levels in both experiments, while negatively affecting total phenolics in the field trials. To conclude, nitrogen fertilization is crucial for crop growth and productivity, however it also implicates nutritional value of the grains as food. These effects should be considered when fertilizing tef with nitrogen, to optimize both crop productivity and nutritional effects.

Background: Characterization and quantification of visual plant traits is often limited to the use of tools and software that were developed to address a specific context, making them unsuitable for other applications. CoverageTool is flexible multi-purpose software capable of area calculation in cm2, as well as coverage area in percentages, suitable for a wide range of applications. Results: Here we present a novel, semi-automated and robust tool for detailed characterization of visual plant traits. We demonstrate and discuss the application of this tool to quantify a broad spectrum of plant phenotypes/traits such as: tissue culture parameters, ground surface covered by annual plant canopy, root and leaf projected surface area, and leaf senescence area ratio. The CoverageTool software provides easy to use functions to analyze images. While use of CoverageTool involves subjective operator color selections, applying them uniformly to full sets of samples makes it possible to provide quantitative comparison between test subjects. Conclusion: The tool is simple and straightforward, yet suitable for the quantification of biological and environmental effects on a wide variety of visual plant traits. This tool has been very useful in quantifying different plant phenotypes in several recently published studies, and may be useful for many applications. © 2019 The Author(s).

Tef (Eragrostis tef (Zucc.) Trotter) is a C4 annual cereal, common in Ethiopia, where it was presumably domesticated. Worldwide interest in tef cultivation and consumption has considerably increased in the last few decades because it is a gluten-free grain with high nutritional value. Here we report on the genetic diversity and heritability in a tef germplasm collection characterized in Israel. A total of 408 accessions of tef held in the Israel Gene Bank were grown in 2015 under common garden (screen-house) conditions for propagation and initial phenotyping. A diversity panel, consisting of 273 accessions representing the entire collection's range of phenotypic diversity, was assembled and evaluated in small field plots in 2016. Further evaluation was conducted in 2017, in single-plant field plots (to eliminate admixtures). A representative plant (plot) was selected from each accession grown in 2017 and its single seed descent progenies where grown in 2018 in single-plant plots. The collection exhibited a wide diversity for each of the measured phenotypic traits, across all four environments. High grain yield was associated in most cases with early flowering time, whereas higher biomass was associated with late flowering. Heritability estimates, calculated based on the 2017, 2018 data, varied between 0.11 for plant biomass and 0.75 for 1000 grain weight. This study shows that tef can successfully grow and produce under irrigated Mediterranean conditions. The wide diversity available in our germplasm collection can provide the foundations for breeding new tef cultivars that are better adapted to these conditions. © Koninklijke Brill NV, Leiden, 2018.

Dissection of the genetic basis of wheat ionome is crucial for understanding the physiological and biochemical processes underlying mineral accumulation in seeds, as well as for efficient crop breeding. Most of the elements essential for plants are metals stored in seeds as chelate complexes with phytic acid or sulfur-containing compounds. We assume that the involvement of phosphorus and sulfur in metal chelation is the reason for strong phenotypic correlations within ionome. Adjustment of element concentrations for the effect of variation in phosphorus and sulfur seed content resulted in drastic change of phenotypic correlations between the elements. The genetic architecture of wheat grain ionome was characterized by quantitative trait loci (QTL) analysis using a cross between durum and wild emmer wheat. QTL analysis of the adjusted traits and two-trait analysis of the initial traits paired with either P or S considerably improved QTL detection power and accuracy, resulting in the identification of 105 QTLs and 617 QTL effects for 11 elements. Candidate gene search revealed some potential functional associations between QTLs and corresponding genes within their intervals. Thus, we have shown that accounting for variation in P and S is crucial for understanding of the physiological and genetic regulation of mineral composition of wheat grain ionome and can be implemented for other plants. © 2019 The Authors The Plant Journal © 2019 John Wiley & Sons Ltd

A near-isogenic line (NIL-7A-B-2), introgressed with a quantitative trait locus (QTL) on chromosome 7AS from wild emmer wheat (Triticum turgidum ssp. dicoccoides) into the background of bread wheat (T. aestivum L.) cv. BarNir, was recently developed and studied in our lab. NIL-7A-B-2 exhibited better productivity and photosynthetic capacity than its recurrent parent across a range of environments. Here we tested the hypothesis that root-system modifications play a major role in NIL-7A-B-2’s agronomical superiority. Root-system architecture (dry matter and projected surface area) and shoot parameters of NIL-7A-B-2 and ‘BarNir’ were evaluated at 40, 62, and 82 days after planting (DAP) in a sand-tube experiment, and root tip number was assessed in a ‘cigar-roll’ seedling experiment, both under well-watered and water-limited (WL) treatments. At 82 DAP, under WL treatment, NIL-7A-B-2 presented greater investment in deep roots (depth 40–100 cm) than ‘BarNir,’ with the most pronounced effect recorded in the 60–80 cm soil depth (60 and 40% increase for root dry matter and surface area, respectively). NIL-7A-B-2 had significantly higher root-tip numbers (∼48%) per plant than ‘BarNir’ under both treatments. These results suggest that the introgression of 7AS QTL from wild emmer wheat induced a deeper root system under progressive water stress, which may enhance abiotic stress resistance and productivity of domesticated wheat.

Aerial thermal remote sensing can provide a means for collecting spatial plant water status data. Many studies have shown their potential in irrigation management but the adaptation of this technology is not straight forward. In this paper, knowledge accumulated in recent years on thermal imagery analysis methodology for water status mapping is summarized aiming at indicating alternatives to calculate the Crop Water Stress Index (CWSI) for commercial scale water status mapping. Based on literature overview, four forms of wet-baselines to calculate CWSI were selected, namely: artificial wet reference surface, two theoretical calculations and a statistical bio-indicator. These baselines were used to calculate CWSI based on multi-temporal aerial thermal images of cotton fields. CWSI based on a statistical bio-indicator and one of the theoretical wet-baselines provided the best correlations. It is argued though that the statistical one is preferable since it includes the plant characteristics and it is farmer-friendly. Based on bio-indicators, leaf water potential maps of three commercial fields were produced on several dates through the season. Water status spatial patterns were not static and the effect of static factors like sandy soil patches also changed through the season. The maps show the importance of in-season variability mapping for rational irrigation management. To improve current variable-rate irrigation, the concept of in-season irrigation management zones (IMZ) based on thermal-images should be considered and integrated with the delineation of static irrigation IMZ.

Productivity of grain crops in semi-arid environments is often affected by drought, which is likely to increase due to predicted climate changes. Wild pea (Pisum fulvum Sibth. & Smith, Pf) accessions sampled across its ecological amplitude in Israel (350–850 mm annual precipitation) were used to assess the genetic diversity for drought responses. We hypothesized that native species evolving under Eastern Mediterranean climate carry adaptive traits to cope with drought stress. Accessions were classified according to single-nucleotide polymorphism variation pattern and habitat ecogeographic parameters. Significant differences were found between the accession groups, but grouping in both systems did not match. Subsequently, 52 Pf accessions and three domesticated pea (P. sativum L.) genotypes were evaluated during 2 yr under well-watered (∼580 mm) and water-limited (∼340 mm) treatments. Total dry matter, grain yield, harvest index, and average grain weight were higher in domesticated pea than wild Pf; however several Pf accessions exhibited lower drought susceptibility indices (i.e., greater stability across environments) than domesticated genotypes. Of special interest are a number of Pf genotypes in which low susceptibility to water stress was coupled with relatively high productivity. The sampling habitats of those low susceptibility–high productivity accessions are characterized by mild (400–530 mm) annual precipitation. Further sampling and evaluation of Pf from such locations may improve our understanding of pea drought adaptation and yield physiology.

Wild emmer wheat (Triticum turgidum ssp. dicoccoides) is considered a promising source for improving stress resistances in domesticated wheat. Here we explored the potential of selected quantitative trait loci (QTLs) from wild emmer wheat, introgressed via marker-assisted selection, to enhance drought resistance in elite durum (T. turgidum ssp. durum) and bread (T. aestivum) wheat cultivars. The resultant near-isogenic lines (BC₃F₃ and BC₃F₄) were genotyped using SNP array to confirm the introgressed genomic regions and evaluated in two consecutive years under well-watered (690–710 mm) and water-limited (290–320 mm) conditions. Three of the introgressed QTLs were successfully validated, two in the background of durum wheat cv. Uzan (on chromosomes 1BL and 2BS), and one in the background of bread wheat cvs. Bar Nir and Zahir (chromosome 7AS). In most cases, the QTL x environment interaction was validated in terms of improved grain yield and biomass—specifically under drought (7AS QTL in cv. Bar Nir background), under both treatments (2BS QTL), and a greater stability across treatments (1BL QTL). The results provide a first demonstration that introgression of wild emmer QTL alleles can enhance productivity and yield stability across environments in domesticated wheat, thereby enriching the modern gene pool with essential diversity for the improvement of drought resistance.

Wild emmer wheat (Triticum turgidum ssp. dicoccoides) is considered a promising source for improving drought resistance in domesticated wheat. Nevertheless, wild germplasm has not been widely used in wheat breeding for abiotic stress resilience. In the current study, a near isogenic line NIL-7A-B-2, introgressed with a drought-related QTL from wild emmer wheat on chromosome 7A, and its recurrent parent, bread wheat cv. BarNir, were investigated under four environments across 2 years—water-limited and well-watered conditions in a rain-protected screen-house (Year 1) and two commercial open field plots under ample precipitation (Year 2). NIL-7A-B-2 exhibited an advantage over BarNir in grain yield and biomass production under most environments. Further physiological analyses suggested that enhanced photosynthetic capacity and photochemistry combined with higher flag leaf area are among the factors underlying the improved productivity of NIL-7A-B-2. These were coupled with improved sink capacity in NIL-7A-B-2, manifested by greater yield components than its parental line. This study provides further support for our previous findings that introgression of wild emmer QTL alleles, using marker assisted selection, can enhance grain yield and biomass production across environments in domesticated wheat, thereby enriching the modern gene pool with essential diversity for the improvement of yield and drought resistance.

BACKGROUND: The metabolite content of a seed and its ability to germinate are determined by genetic makeup and environmental effects during development. The interaction between genetics, environment and seed metabolism and germination was studied in 72 tomato homozygous introgression lines (IL) derived from Solanum pennelli and S. esculentum M82 cultivar. Plants were grown in the field under saline and fresh water irrigation during two consecutive seasons, and collected seeds were subjected to morphological analysis, gas chromatograph-mass spectrometry (GC-MS) metabolic profiling and germination tests. RESULTS: Seed weight was under tight genetic regulation, but it was not related to germination vigor. Salinity significantly reduced seed number but had little influence on seed metabolites, affecting only 1% of the statistical comparisons. The metabolites negatively correlated to germination were simple sugars and most amino acids, while positive correlations were found for several organic acids and the N metabolites urea and dopamine. Germination tests identified putative loci for improved germination as compared to M82 and in response to salinity, which were also characterized by defined metabolic changes in the seed. CONCLUSIONS: An integrative analysis of the metabolite and germination data revealed metabolite levels unambiguously associated with germination percentage and rate, mostly conserved in the different tested seed development environments. Such consistent relations suggest the potential for developing a method of germination vigor prediction by metabolic profiling, as well as add to our understanding of the importance of primary metabolic processes in germination.

Wild emmer wheat (Triticum turgidum ssp. dicoccoides) is considered a promising source for improving stress resistances in domesticated wheat. Here we explored the potential of selected quantitative trait loci (QTLs) from wild emmer wheat, introgressed via marker-assisted selection, to enhance drought resistance in elite durum (T. turgidum ssp. durum) and bread (T. aestivum) wheat cultivars. The resultant near-isogenic lines (BC3F3 and BC3F4) were genotyped using SNP array to confirm the introgressed genomic regions and evaluated in two consecutive years under well-watered (690-710 mm) and water-limited (290-320 mm) conditions. Three of the introgressed QTLs were successfully validated, two in the background of durum wheat cv. Uzan (on chromosomes 1BL and 2BS), and one in the background of bread wheat cvs. Bar Nir and Zahir (chromosome 7AS). In most cases, the QTL x environment interaction was validated in terms of improved grain yield and biomass-specifically under drought (7AS QTL in cv. Bar Nir background), under both treatments (2BS QTL), and a greater stability across treatments (1BL QTL). The results provide a first demonstration that introgression of wild emmer QTL alleles can enhance productivity and yield stability across environments in domesticated wheat, thereby enriching the modern gene pool with essential diversity for the improvement of drought resistance.

Background: Wheat domestication is considered as one of the most important events in the development of human civilization. Wheat spikelets have undergone significant changes during evolution under domestication, resulting in soft glumes and larger kernels that are released easily upon threshing. Our main goal was to explore changes in transcriptome expression in glumes that accompanied wheat evolution under domestication. Methods: A total of six tetraploid wheat accessions were selected for transcriptome profiling based on their rachis brittleness and glumes toughness. RNA pools from glumes of the central spikelet at heading time were used to construct cDNA libraries for sequencing. The trimmed reads from each library were separately aligned to the reference sub-genomes A and B, which were extracted from wheat survey sequence. Differentially expression analysis and functional annotation were performed between wild and domesticated wheat, to identity candidate genes associated with evolution under domestication. Selected candidate genes were validated using real time PCR. Results: Transcriptome profiles of wild emmer wheat, wheat landraces, and wheat cultivars were compared using next generation sequencing (RNA-seq). We have found a total of 194,893 transcripts, of which 73,150 were shared between wild, landraces, and cultivars. From 781 differentially expressed genes (DEGs), 336 were down-regulated and 445 were up-regulated in the domesticated compared to wild wheat genotypes. Gene Ontology (GO) annotation assigned 293 DEGs (37.5 %) to GO term groups, of which 134 (17.1 %) were down-regulated and 159 (20.4 %) up-regulated in the domesticated wheat. Some of the down-regulated DEGs in domesticated wheat are related to the biosynthetic pathways that eventually define the mechanical strength of the glumes, such as cell wall, lignin, pectin and wax biosynthesis. The reduction in gene expression of such genes, may explain the softness of the glumes in the domesticated forms. In addition, we have identified genes involved in nutrient remobilization that may affect grain size and other agronomic traits evolved under domestication. Conclusions: The comparison of RNA-seq profiles between glumes of wheat groups differing in glumes toughness and rachis brittleness revealed a few DEGs that may be involved in glumes toughness and nutrient remobilization. These genes may be involved in processes of wheat improvement under domestication. © 2015 Zou et al.

Wheat domestication and subsequent evolution under domestication occurred since the dawn of agriculture and caused significant genetic changes that affected plant morphology, physiology and phenology. The majority of these traits are quantitative traits controlled by many genes. Correspondingly, the main goal of the current study is genetic dissection of the key domestication trait (brittle rachis) and traits evolved under domestication, based on quantitative phenotyping. Genetic mapping of quantitative trait loci (QTL) affecting brittle rachis, threshability, threshing efficiency, spike harvest index and kernel weight was conducted using a recombinant inbred lines population derived from a cross between Triticum durum and wild emmer wheat. Using a new quantitative phenotyping approach, we discovered novel QTLs underlying rachis fragility, spike threshability and other domestication-related traits and confirmed some of the known putative locations for QTLs affecting these traits. Overall, the number of domestication-related QTLs mapped to the A genome was twofold higher than those found on the B genome, in accordance with the concept of ‘genome asymmetry’, implying that the A genome is dedicated to the control of morphological traits, house-keeping metabolic reactions and yield components. Our results add a new dimension to this important concept and contribute to a better understanding of the initial steps of domestication evolution of cereals. © 2014, Springer Science+Business Media Dordrecht.

'Domestication syndrome' (DS) denotes differences between domesticated plants and their wild progenitors. Crop plants are dynamic entities; hence, not all parameters distinguishing wild progenitors from cultigens resulted from domestication. In this opinion article, we refine the DS concept using agronomic, genetic, and archaeobotanical considerations by distinguishing crucial domestication traits from traits that probably evolved post-domestication in Near Eastern grain crops. We propose that only traits showing a clear domesticated-wild dimorphism represent the pristine domestication episode, whereas traits showing a phenotypic continuum between wild and domesticated gene pools mostly reflect post-domestication diversification. We propose that our approach may apply to other crop types and examine its implications for discussing the timeframe of plant domestication and for modern plant science and breeding. © 2013 Elsevier Ltd.