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Drought events or the combination of drought and heat conditions are expected to become more frequent due to global warming, and wheat yields may fall below their long-term average. One way to increase climate-resilience of modern high-yielding varieties is by their genetic improvement with beneficial alleles from crop wild relatives. In the present study, the effect of two beneficial QTLs introgressed from wild emmer wheat and incorporated in the three wheat varieties BarNir, Zahir and Uzan was studied under well-watered conditions and under drought stress using non-destructive High-throughput Phenotyping (HTP) throughout the life cycle in a single pot-experiment. Plants were daily imaged with RGB top and side view cameras and watered automatically. Further, at two time points, the quantum yield of photosystem II was measured with a top view FluorCam. The QTL carrying near isogenic lines (NILs) were compared with their corresponding parents by t-test for all non-invasively obtained traits and for the manually determined agronomic and yield parameters. Data quality of phenotypic traits (repeatability) in the controlled HTP experiment was above 85% throughout the life cycle and at maturity. Drought stress had a strong effect on growth in all wheat genotypes causing biomass reduction from 2% up to 70% at early and late points in the drought period, respectively. At maturity, the drought caused 47–55% decreases in yield-related traits grain weight, straw weight and total biomass and reduced TKW by 10%, while water use efficiency (WUE) increased under drought by 29%. The yield-enhancing effect of the introgressed QTLs under drought conditions that were previously demonstrated under field/screenhouse conditions in Israel, could be mostly confirmed in a greenhouse pot experiment using HTP. Daily precision phenotyping enabled to decipher the mode of action of the QTLs in the different genetic backgrounds throughout the entire wheat life cycle. Daily phenotyping allowed a precise determination of the timing and size of the QTLs effect (s) and further yielded information about which image-derived traits are informative at which developmental stage of wheat during the entire life cycle. Maximum height and estimated biovolume were reached about a week after heading, so experiments that only aim at exploring these traits would not need a longer observation period. To obtain information on different onset and progress of senescence, the CVa curves represented best the ongoing senescence of plants. The QTL on 7A in the BarNir background was found to improve yield under drought by increased biomass growth, a higher photosynthetic performance, a higher WUE and a “stay green effect.”

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 (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.

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 inRhopalosiphum 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.

Background The rising availability of assemblies of large genomes (e.g. bread and durum wheat, barley) and their annotations deliver the basis to graphically present genome organization of parents and progenies on a physical scale. Genetic maps are a very important tool for breeders but often represent distorted models of the actual chromosomes, e.g., in centromeric and telomeric regions. This biased picture might lead to imprecise assumptions and estimations about the size and complexity of genetic regions and the selection of suitable molecular markers for the incorporation of traits in breeding populations or near-isogenic lines (NILs). Some software packages allow the graphical illustration of genotypic data, but to the best of our knowledge, suitable software packages that allow the comparison of genotypic data on the physical and genetic scale are currently unavailable. Results We developed a simple Java-based-software called GenoTypeMapper (GTM) for comparing genotypic data on genetic and physical maps and tested it for effectiveness on data of two NILs that carry QTL-regions for drought stress tolerance from wild emmer on chromosome 2BS and 7AS. Both NILs were more tolerant to drought stress than their recurrent parents but exhibited additional undesirable traits such as delayed heading time. Conclusions In this article, we illustrate that the software easily allows users to display and identify additional chromosomal introgressions in both NILs originating from the wild emmer parent. The ability to detect and diminish linkage drag can be of particular interest for pre-breeding purposes and the developed software is a well-suited tool in this respect. The software is based on a simple allele-matching algorithm between the offspring and parents of a crossing scheme. Despite this simple approach, GTM seems to be the only software that allows us to analyse, illustrate and compare genotypic data of offspring of different crossing schemes with up to four parents in two different maps. So far, up to 500 individuals with a maximum number of 50,000 markers can be examined with the software. The main limitation that hampers the performance of the software is the number of markers that are examined in parallel. Since each individual must be analysed separately, a maximum of ten individuals can currently be displayed in a single run. On a computer with an Intel five processor of the 8th generation, GTM can reliably either analyse a single individual with up to 12,000 markers or ten individuals with up to 3,600 markers in less than five seconds. Future work aims to improve the performance of the software so that more complex crossing schemes with more parents and more markers can be analysed.

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.

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. Electronic supplementary material The online version of this article (10.1186/s13007-019-0472-2) contains supplementary material, which is available to authorized users.

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.

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.

Durum wheat, Triticum turgidum ssp. durum Desf., is an important crop particularly in the Mediterranean basin. Powdery mildew, caused by the pathogen Blumeria graminis f. sp. tritici (Bgt), is a major disease of wheat that results in significant yield losses worldwide. A recombinant inbred line (RIL) population, derived from a cross between durum wheat and wild emmer wheat, T. turgidum ssp. dicoccoides, was used for genomic dissection of quantitative and qualitative resistance loci against wheat powdery mildew based on a genomic map of >600 markers, evenly distributed across the A and B genomes of tetraploid wheat. The genetic analysis of the phenotypic reactions of the RIL population to two Bgt isolates revealed two different resistance mechanisms. The first is monogenic: a wild emmer wheat allele in a single locus conferring complete resistance to Bgt#15, previously designated as PmG16. The second one is polygenic: a set of durum wheat alleles, in five independent QTLs that control partial resistance to Bgt#66 in the RIL population, with a LOD score range of 3.4–19.8. One of them is a major quantitative resistance locus (QRL) that was mapped on chromosome 1A and explains 26.4 % of the variance. In most of the detected QRLs, the durum wheat alleles conferred resistance to powdery mildew. These findings are exceptional in the sense that, so far, only a few Pm alleles originated from a durum wheat background. Therefore, our results emphasize the high potential of exploiting the wide genetic diversity of tetraploid wheat germplasm for wheat breeding using modern wheat genomics tools.

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 (Triticum 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 (WW) 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.

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.

'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.

†BackgroundandAims The harvesting method of wild and cultivated cereals has long been recognized as an important factor in the emergence of domesticated non-shattering ear genotypes. This study aimed to quantify the effects of spike brittleness and threshability on threshing time and efficiency in emmer wheat, and to evaluate the implications of post-harvest processes on domestication of cereals in the Near East. †Methods A diverse collection of tetraploid wheat genotypes, consisting ofTriticumturgidum ssp.dicoccoides ‐ the wild progenitor of domesticated wheat ‐ traditional landraces, modern cultivars (T. turgidum ssp. durum) and 150 recombinant (wild × modern) inbred lines, was used in replicated controlled threshing experiments to quantify the effects of spike brittleness and threshability on threshing time and efficiency. †Key Results The transition from a brittle hulled wild phenotype to non-brittle hulled phenotype (landraces) was associated with an approx. 30 % reduction in threshing time, whereas the transition from the latter to non-brittle free-threshing cultivars was associated with an approx. 85 % reduction in threshing time. Similar trends were obtained with groups of recombinant inbred lines showing extreme phenotypes of brittleness and threshability. †Conclusions In tetraploid wheat, both non-brittle spike and free-threshing are labour-saving traits that increase the efficiency of post-harvest processing, which could have been an incentive for rapid domestication of the Near Eastern cereals, thus refuting the recently proposed hypothesis regarding extra labour associated with the domesticated phenotype (non-brittle spike) and its presumed role in extending the domestication episode time frame.

Modifying plant architecture is considered a promising breeding option to enhance crop productivity. Modern chickpea (Cicer arietinum L.) cultivars with either compound (wild-type) or simple leaf shapes are commercially grown but the relationships between leaf shape and yield are not well understood. In this study, a random sample of ‘Kabuli’ type progeny lines of both leaf types, derived from two crosses between modern American simple leaf cultivars and early-flowering wild-type breeding lines, were planted at different sowing densities. Leaf area development and final grain yield in genotypes of the two leaf types responded differently to changes in sowing densities. Compound leaf lines attained higher leaf area indices and higher grain yields at both low and high sowing densities. Yield responses of the simple leaf lines to increasing sowing density were significantly higher compared to compound leaf genotypes in two of three field experiments. The prospects for utilizing the simple leaf trait as a breeding target for short-season growing areas are discussed.

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.

Revealing the genetic underpinnings of cotton productivity will require understanding both the prehistoric evolution of spinnable fibers, and the results of independent domestication processes in both the Old and New Worlds. Progress toward a reference sequence for the smallest Gossypium genome is a logical stepping-stone toward revealing diversity in the remaining seven genomes (A, B, C, E, F, G, K) that permitted Gossypium species to adapt to a wide range of ecosystems in warmer arid regions of the world, and toward identifying the emergent properties that account for the superior productivity and quality of tetraploid cottons. The greatest challenge facing the cotton community is not genome sequencing per se but the conversion of sequence to knowledge.

Quantitative trait loci (QTLs) for yield and drought related traits were exchanged via marker-assisted selection (MAS) between elite cultivars of two cotton species, Gossypium barbadense (GB) cv. F-177 and Gossypium hirsutum (GH) cv. Siv’on. The resulting near-isogenic lines (NILs) manifested in many cases the expected drought-adaptive traits, but rarely exhibited an advantage in yield relative to the recipient parents. In the current study we characterized the photosynthetic activity of two selected NILs and their recipient parents under dryland and irrigated field conditions. The GB NIL exhibited a stable net rate of CO2 assimilation (A) across a wide range of leaf water potentials with a notable advantage over its recipient parent, F-177, under severe drought, associated with lower stomatal limitation, greater maximum velocity for carboxylation of Rubisco and greater electron transport rate. The GH NIL exhibited greater mesophyll conductance under drought conditions than its recipient parent, Siv’on, but these genotypes did not differ in A. Nevertheless, both NILs did not differ from their recipient parents in yield. This study provides further support to our previous conclusion that MAS is a useful means to enhance drought-adaptive traits of cotton but complimentary conventional breeding is required to achieve high yield potential.

Drought is the major factor limiting wheat productivity worldwide. The gene pool of wild emmer wheat, Triticum turgidum ssp. dicoccoides, harbours a rich allelic repertoire for morpho-physiological traits conferring drought resis- tance. The genetic and physiological bases of drought responses were studied here in a tetraploid wheat popula- tion of 152 recombinant inbreed lines (RILs), derived from a cross between durum wheat (cv. Langdon) and wild emmer (acc# G18-16), under contrasting water availabili- ties. Wide genetic variation was found among RILs for all studied traits. A total of 110 quantitative trait loci (QTLs) were mapped for 11 traits, with LOD score range of 3.0-35.4. Several QTLs showed environmental specificity, accounting for productivity and related traits under water- limited (20 QTLs) or well-watered conditions (15 QTLs), and in terms of drought susceptibility index (22 QTLs). Major genomic regions controlling productivity and related traits were identified on chromosomes 2B, 4A, 5A and 7B. QTLs for productivity were associated with QTLs for drought-adaptive traits, suggesting the involvement of several strategies in wheat adaptation to drought stress. Fifteen pairs of QTLs for the same trait were mapped to seemingly homoeologous positions, reflecting synteny between the A and B genomes. The identified QTLs may facilitate the use of wild alleles for improvement of drought resistance in elite wheat cultivars.

Quantitative trait loci (QTLs) for yield and drought related physiological traits, osmotic potential (OP), carbon isotope ratio (d 13 C, an indica- tor of water use efficiency), and leaf chlorophyll content (Chl), were exchanged via marker-assisted selection (MAS) between elite cultivars of the two cotton species Gossypium barbadense cv. F-177 and G. hirsutum cv. Siv'on. The resulting near isogenic lines (NILs) were examined in two field trials, each with two irrigation regimes, in order to (1) evaluate the potential to improve cotton drought resistance by MAS and (2) test the role of physiological traits in plant productivity. NILs introgressed with QTLs for high yield rarely exhibited an advantage in yield relative to the recipient parent, whereas a considerable number of NILs exhibited the expected phenotype in terms of lower OP (5 out of 9), higher d 13 C (4 out of 6) or high Chl (2 out of 3). Several NILs exhibited considerable modifications in non-targeted traits including leaf morphology, stomatal conductance and specific leaf weight (SLW). In G. barbadense genotypes, yield was correlated negatively with d 13 C and OP and positively with stomatal conductance, SLW and Chl, whereas in G. hirsutum yield was negatively correlated with d 13 C, SLW and Chl. This dissimilarity suggests that each of the respective species has evolved different mechanisms underlying plant productivity. We conclude that the improvement of drought related traits in cotton NILs may lead to improved drought resistance via MAS, but that conventional breeding may be necessary to combine the introduced QTL(s) with high yield potential.

Drought is the main environmental stress limiting crop productivity and yield stability worldwide. Improving drought resistance of crop plants is considered a sustainable and economically viable solution to increasing agricultural productivity under diminishing water availability. The implementation of this solution requires wide explorations of potential genetic resources and in-depth understanding of their adaptive mechanisms and responses to water deficit. In this minireview we summarize the physiological, ecological, and genetic aspects of drought resistance in wild emmer wheat [Triticum turgidum ssp. dicoccoides (Korn.) Thell.] and discuss their implications on wheat improvement. A comprehensive survey of wild emmer wheat populations, collected across the aridity gradient in Israel, revealed wide genetic diversity for drought responses, with a considerable number of wild accessions exhibiting an advantage over durum wheat cultivars. A variety of adaptive traits associated with improved performances u...

Micronutrient malnutrition afflicts over three billion peopleworldwide and the numbers are continuously increasing. Developing genetically micronutrientenriched cereals, which are the predominant source of human dietary, is essential to alleviate malnutrition worldwide. Wheat chromosome 6B derived from wild emmerwheat [Triticum turgidum ssp. dicoccoides (Körn.) Thell] was previously reported to be a source for high Zn concentration in the grain. In the present study, recombinant chromosome substitution lines (RSLs), previously constructed for genetic and physical maps of Gpc-B1 (a 250-kb locus affecting grain protein concentration), were used to identify the effects of the Gpc-B1 locus on grain micronutrient concentrations. RSLs carrying the Gpc-B1 allele of T. dicoccoides accumulated on average 12% higher concentration of Zn, 18% higher concentration of Fe, 29% higher concentration of Mn and 38% higher concentration of protein in the grain as compared with RSLs carrying the allele from cultivated wheat (Triticum durum). Furthermore, the high grain Zn, Fe and Mn concentrations were consistently expressed in five different environments with an absence of genotype by environment interaction. The results obtained in the present study also confirmed the previously reported effect of the wild-type allele of Gpc-B1 on earlier senescence of flag leaves. We suggest that the Gpc-B1 locus is involved in more efficient remobilization of protein, zinc, iron and manganese from leaves to the grains, in addition to its effect on earlier senescence of the green tissues.

Canopy temperature has long been recognized as an indicator of plant water status and as a potential tool for irrigation scheduling. In the present study, the potential of using thermal images for an in-field estimation of the water status of cotton under a range of irrigation regimes was investigated. Thermal images were taken with a radiometric infrared video camera. Specific leaves that appeared in the camera field of view were sampled, their LWP was measured and their temperature was calculated from the images. Regression models were built in order to predict LWP according to the crop canopy temperature and to the empirical formulation of the crop water stress index (CWSI). Statistical analysis revealed that the relationship between CWSI and LWP was more stable and had slightly higher correlation coefficients than that between canopy temperature and LWP. The regression models of LWP against CWSI and against leaf temperatures were used to create LWP maps. The classified LWP maps showed that there was spatial variability in each treatment, some of which may be attributed to the difference between sunlit and shaded leaves. The distribution of LWP in the maps showed that irrigation treatments were better distinguished from each other when the maps were calculated from CWSI than from leaf temperature alone. Furthermore, the inclusion of the spatial pattern in the classification enhanced the differences between the treatments and was better matched to irrigation amounts. Optimal determination of the water status from thermal images should be based on an overall view of the physical status as well as on the analysis of the spatial structure. Future study will involve investigating the robustness of the models and the potential of using water status maps, derived from aerial thermal images, for irrigation scheduling and variable management in commercial fields.

Wild emmer wheat ( Triticum turgidum spp. dicoccoides (Korn.) Thell.), the tetraploid progenitor of cultivated wheat, is a potential source for various agronomical traits, including drought resistance. The objectives of this study were to characterize (1) the genetic diversity for drought resistance in wild emmer wheat, and (2) the relationship between drought responses of the wild emmer germplasm and the ecogeographical parameters of its collection sites. A total of 110 wild emmer accessions consisting of 25 populations and three control durum wheat cultivars were examined under two irrigation regimes, well-watered (’wet’) and water-limited (’dry’). Wide genetic diversity was found both between and within the wild emmer populations in most variables under each treatment. A considerable number of the wild emmer accessions exhibited an advantage in productivity (spike and total dry matter) over their cultivated counterparts. Most wild emmer wheat accessions exhibited a greater carbon isotope ratio ( d d d 13 C, indicating higher water-use efficiency) under the dry treatment and higher plasticity of d d d 13 C relative to the cultivated controls, which may have contributed to the drought adaptations in the former. The most outstanding drought-tolerance capacity (in term of productivity under the dry treatment and susceptibility indices) was detected in wild emmer populations originated from hot dry locations. The results suggest that wild emmer has the potential to improve drought resistance in cultivated wheat.

Testing of the extent to which different complex traits share common genetic control provides a means to distinguish associations that are truly diagnostic of genetic potential for improved adaptation to abiotic stress, from incidental phenotypic correlations. In two generations of progeny from a cross between Gossypium hirsutum and Gossypium barbadense , quantitative trait loci (QTL) mapping was used to evaluate correspondence in genetic control of selected physiological measures and productivity under waterlimited and well-watered environments, respectively. A total of 33 QTLs were detected for five physiological variables [osmotic potential (OP), carbon isotope ratio ( d d d 13 C; indicator of water use efficiency), canopy temperature, chlorophyll a and b ], and 46 QTLs for five measures of crop productivity [dry matter, seed cotton yield (SC), harvest index, boll weight, and boll number]. QTL likelihood intervals for high SC and low OP corresponded in three genomic regions, two of which mapped to homoeologous locations on the two subgenomes of tetraploid cotton. QTLs for d 13 C showed only incidental association with productivity, indicating that high water use efficiency can be associated with either high or low productivity. Different cotton species have evolved different alleles related to physiological responses and productivity under water deficit, which may permit the development of genotypes that are betteradapted to arid conditions.

Silverleaf whitefly (SLW), Bemisia argentifolii Bellows and Perring, is one of the most noxious pests of numerous field and vegetable crops, causing billions of dollars worth of damage throughout the world. SLW is a phloem feeder whose feeding is likely to interfere with phloem transport. The aim of this study was to test the hypothesis that SLW infestation impairs carbohydrate export from source leaves, and consequently increases their carbohydrate content. The youngest fully expanded leaves of cotton (Gossypium hirsutum L., cv. Siv'on), grown under SLW-infested and noninfested conditions, were characterized for their diurnal changes in carbohydrate content and photoassimilate export. SLW infestation induced a considerable reduction in net photosynthetic rate (P n ), coupled with increased sucrose, glucose and fructose and decreased starch concentrations. Export rate was determined after 14 CO 2 pulse-labeling both by in situ monitoring of leaf radioactivity and by analyzing the content and radioactivity of the major carbon metabolites. Radioactive counting indicated a lower rate of 14 C efflux for the infested plants. A similar trend was found for the specific activities of sucrose and the three soluble sugars combined (sucrose, glucose and fructose). A single exponential decay function with asymptote was fitted to the above efflux curves. All the calculated exponential coefficients demonstrated lower export rates after SLW injury. These results indicate that SLW impairs photoassimilate export, suggesting possible down-regulation of P n due to increased foliar soluble sugar contents.

Carbon isotope ratio ( 13 C/ 12 C, expressed with a differential notation as δ 13 C) has been proposed as an indirect selection criterion for plant water-use efficiency (WUE=total dry matter produced or yield harvested/water used). For efficient modification of WUE in breeding programs, it is essential to determine a sampling strategy, which will provide consistent genotypic ranking for δ 13 C and maximum differentiation between genotypes. The effects of growth stages and plant organs on δ 13 C values and their genotypic variations were studied in cotton cultivars grown in the field under two irrigation regimes. Values of δ 13 C in leaf remained stable during peak flowering and boll development stages and significantly increased at boll ripening stage, which could result from the effect of late-season water stress on WUE. δ 13 C varied significantly between plant organs, with lower values obtained in assimilating organs, leaf and bur, and higher values in non-assimilating organs, stem and fiber. This could possibly have resulted from carbon discrimination during secondary metabolism. A non-crossover interaction was found between growth stage and cultivar, whereas plant-organ effect did not interact with cultivars, indicating that a similar ranking of cultivars can be expected with δ 13 C at various growth stages and plant organs. Late season leaf samples and bur of the last boll revealed maximal differentiation among genotypes and water regimes, both among other growth stages as well as among other plant organs. These samples have also revealed the highest correlations between δ 13 C and WUE estimates. It is concluded that leaf sampling during boll ripening stage may be most effective for the assessment of δ 13 C as an indicator of WUE.

Wheat is undoubtedly one of the world's major food sources since the dawn of Near Eastern agriculture and up to the present day. Morphological, physiological, and genetic modifications involved in domestication and subsequent evolution under domestication were investigated in a tetraploid recombinant inbred line population, derived from a cross between durum wheat and its immediate progenitor wild emmer wheat. Experimental data were used to test previous assumptions regarding a protracted domestication process. The brittle rachis (Br) spike, thought to be a primary characteristic of domestication, was mapped to chromosome 2A as a single gene, suggesting, in light of previously reported Br loci (homoeologous group 3), a complex genetic model involved in spike brittleness. Twenty-seven quantitative trait loci (QTLs) conferring threshability and yield components (kernel size and number of kernels per spike) were mapped. The large number of QTLs detected in this and other studies suggests that following domestication, wheat evolutionary processes involved many genomic changes. The Br gene did not show either genetic (co-localization with QTLs) or phenotypic association with threshability or yield components, suggesting independence of the respective loci. It is argued here that changes in spike threshability and agronomic traits (e.g. yield and its components) are the outcome of plant evolution under domestication, rather than the result of a protracted domestication process. Revealing the genomic basis of wheat domestication and evolution under domestication, and clarifying their inter-relationships, will improve our understanding of wheat biology and contribute to further crop improvement.

Quantitative trait loci (QTLs) for yield and drought-related traits were exchanged via marker-assisted selection between elite cultivars of two cotton species, Gossypium barbadense (GB) cv. F-177 and Gossypium hirsutum (GH) cv. Siv'on. Three of the resultant near-isogenic lines (NILs), each introgressed with a different QTL region, expressed an advantage in osmotic adjustment (OA) and other drought-related traits relative to their recipient parents. These NILs and the parental genotypes were field-grown under well-watered and water-limited conditions, and characterized for their metabolic and mineral compositions. Comparisons were then made between (1) GB and GH genotypes, (2) the contrasting water regimes and (3) each NIL and its recipient parent. Hierarchical clustering analysis clearly distinguished between GB and GH genotypes based on either metabolite or mineral composition. Comparisons between well-watered and water-limited conditions in each of the genotypes showed differing trends in the various solutes. The greater concentrations of potassium, magnesium and calcium under water stress, when compared with well-watered conditions, may have enhanced OA or osmoprotection. All NILs exhibited significantly modified solute composition relative to their recipient parents. In particular, increased levels of alanine, aspartic acid, citric acid, malic acid, glycerol, myoinositol, threonic acid, potassium, magnesium and calcium were found under drought conditions in one or more of the NILs relative to their recipient parents. The increased values of these solutes could contribute to the superior capacity of these NILs to cope with drought.

Awns are long, stiff filamentous extensions of glumes in many grasses. In wheat, awns contribute up to 40% of the grain's photosynthetic assimilates, and assist in seed dispersal. Awns accumulate silica in epidermal hairs and papillae, and silica has been positively associated with yield and environmental stress tolerance. Here, the awns of a set of domesticated wheat genotypes and their direct progenitor, Triticum turgidum ssp. dicoccoides were characterized. In addition, the silica concentration in awns was genetically dissected in a tetraploid wheat population of recombinant inbred lines (RILs) derived from a cross between durum wheat (cv. Langdon) and wild emmer (accession G18-16). Scanning electron micrographs revealed a continuous silica layer under the cuticle. Extended silicification was identified in the epidermis cell wall and in sclerenchyma cells near the vascular bundles, but not in the stomata, suggesting that an active process directs the soluble silica away from the water evaporation stream. The number of silicified cells was linearly correlated to silica concentration in dry weight (DW), suggesting cellular control over silicification. Domesticated wheat awns contained up to 19% silica per DW, as compared with 7% in the wild accessions, suggesting selection pressure associated with the domestication process. Six quantitative trait loci (QTLs) for silica were identified in the awns, with a LOD score of 3.7-6.3, three of which overlapped genomic regions that contribute to high grain protein. Localization of silica in the awns and identification of QTLs help illuminate mechanisms associated with silica metabolism in wheat.

Genetic diversity for grain nutrients in wild emmer wheat – A potential for wheat improvement Merav Chatzav1‡, Zvi Peleg1‡, Tzion Fahima2, Ismail Cakmak3, Yehoshua Saranga1* Abstract • Background and Aims - Mineral nutrient malnutrition, and particularly deficiency in zinc (Zn) and iron (Fe), afflicts over three billion people worldwide. In the current study wild emmer wheat (Triticum turgidum ssp. dicoccoides (Korn.) Thell.), the progenitor of domesticated wheats, was tested for (i) genetic diversity in grain nutrient concentrations, (ii) the associations among grain nutrients and their relationships with plant productivity, and (iii) their association with the ecogeographical origin of wild emmer accessions. • Methods - A Total of 154 genotypes, including wild emmer accessions from across the Near Eastern Fertile Crescent and diverse wheat cultivars, were characterized in this two-year field study for grain protein (GPC), micronutrients (Zn, Fe, copper and manganese) and macronutrients (calcium, magnesium, potassium, phosphorus and sulphur) concentrations. • Key Results - A wide genetic diversity was found among the wild emmer accessions for all grain nutrients. The concentrations of grain Zn, Fe and protein in wild accessions were about two-fold greater than in the domesticated genotypes. Concentrations of these compounds were positively correlated with one another, with no clear association with plant productivity suggesting that all three nutrients can be improved concurrently with no yield penalty. A subset of 12 populations revealed a significant genetic variation between and within populations for all minerals. Association between soil characterise of the site of collection and grain nutrient concentrations showed negative associations between soil clay and GPC and between Zn in soil and in grain, the latter suggesting that the greatest potential for grain nutrient mineral is expected in populations from mineral deficient soils. • Conclusions – Wild emmer wheat germplasm offers unique opportunities to exploit favourable alleles for grain nutrient properties that were excluded from the domesticated wheat genepool. Key words: Triticum turgidum ssp. dicoccoides, grain quality, protein, zinc, iron, wheat improvement, micronutrients, macronutrients

Nineteen wild emmer wheat [Triticum turgidum ssp. dicoccoides (Körn.) Thell.] genotypes were evaluated for the grain concentrations of phosphorous (P), potassium (K), sulfur (S), magnesium (Mg), calcium (Ca), zinc (Zn), manganese (Mn), iron (Fe) and cooper (Cu) under five different environments in Turkey and Israel. Each mineral nutrient has been investigated for the (1) genotype by environment (G × E) interactions, (2) genotype stability, (3) correlation among minerals and (4) mineral stability. Among the macronutrients analyzed, grain concentrations of Ca (range 338–2,034 mg kg−1) and S (range 0.18–0.43%) showed the largest variation. In the case of micronutrients, the largest variation was observed in the grain Mn concentration (range 13–87 mg kg−1). Grain concentrations of Fe and Zn also showed important variation (range 27–86 and 39–115 mg kg−1, respectively). Accessions with higher nutrient concentrations (especially Zn and Fe) had also greater grain weight, suggesting that higher grain Zn and Fe concentrations are not necessarily related to small grain size or weight. Analysis of variance showed that environment was the most important source of variation for K, S, Ca, Fe, Mn and Zn, explaining between 44 and 78% of the total variation and G × E explained between 20 and 40% of the total variation in all the minerals, except for S and Zn where its effect accounted for less than 16%. Genotype was the most important source of variation for Cu (explaining 38% of the total variation). However, genotype effect was also important for Mg, Mn, Zn and S. Sulfur and Zn showed the largest heritability values (77 and 72%, respectively). Iron exhibited low heritability and high ratio value between the G × E and genotype variance components , suggesting that specific adaptation for this mineral could be positively exploited. The wild emmer germplasm tested in the current study revealed some outstanding accessions (such as MM 5/4 and 24-39) in terms of grain Zn and Fe concentrations and environmental stability that can be used as potential donors to enhance grain micronutrient concentrations in wheats. Keywords Genotype × environment interaction - Grain quality - Micronutrients - Plant breeding - Mineral stability - Triticum turgidum ssp. dicoccoides

Micronutrient malnutrition, and particularly deficiency in zinc (Zn) and iron (Fe), afflicts over three billion people worldwide, and nearly half of the world’s cereal-growing area is affected by soil Zn deficiency. Wild emmer wheat [Triticum turgidum ssp. dicoccoides (Korn.) Thell.], the progenitor of domesticated durum wheat and bread wheat, offers a valuable source of economically important genetic diversity including grain mineral concentrations. Twenty two wild emmer wheat accessions, representing a wide range of drought resistance capacity, as well as two durum wheat cultivars were examined under two contrasting irrigation regimes (well-watered control and water-limited), for grain yield, total biomass production and grain Zn, Fe and protein concentrations. The wild emmer accessions exhibited high genetic diversity for yield and grain Zn, Fe and protein concentrations under both irrigation regimes, with a considerable potential for improvement of the cultivated wheat. Grain Zn, Fe and protein concentrations were positively correlated with one another. Although irrigation regime significantly affected ranking of genotypes, a few wild emmer accessions were identified for their advantage over durum wheat, having consistently higher grain Zn (e.g., 125 mg kg−1), Fe (85 mg kg−1) and protein (250 g kg−1) concentrations and high yield capacity. Plants grown from seeds originated from both irrigation regimes were also examined for Zn efficiency (Zn deficiency tolerance) on a Zn-deficient calcareous soil. Zinc efficiency, expressed as the ratio of shoot dry matter production under Zn deficiency to Zn fertilization, showed large genetic variation among the genotypes tested. The source of seeds from maternal plants grown under both irrigation regimes had very little effect on Zn efficiency. Several wild emmer accessions revealed combination of high Zn efficiency and drought stress resistance. The results indicate high genetic potential of wild emmer wheat to improve grain Zn, Fe and protein concentrations, Zn deficiency tolerance and drought resistance in cultivated wheat.

The association between allelic diversity and ecogeographi- cal variables was studied in natural populations of wild emmer wheat (Triticum turgidum ssp. dicoccoides (Korn.) Thell.), the tetraploid progenitor of cultivated wheat. Pat- terns of allelic diversity in 54 microsatellite loci were ana- lyzed in a collection of 145 wild emmer wheat accessions representing 25 populations that were sampled across natu- rally occurring aridity gradient in Israel and surrounding regions. The obtained results revealed that 56% of the genetic variation resided among accessions within popula- tions, while only 44% of the variation resided between populations. An unweighted pair-group method analysis (UPGMA) tree constructed based on the microsatellite allelic diversity divided the 25 populations into six major groups. Several groups were comprised of populations that were collected in ecologically similar but geographi- cally remote habitats. Furthermore, genetic differentiation between populations was independent of the geographical distances. An interesting evolutionary phenomenon is high- lighted by the unimodal relationship between allelic diver- sity and annual rainfall (r = 0.74, P < 0.0002), indicating higher allelic diversity in populations originated from habi- tats with intermediate environmental stress (i.e. rainfall 350-550 mm year -1 ). These results show for the first time that the 'intermediate-disturbance hypothesis', explaining biological diversity at the ecosystem level, also dominates the genetic diversity within a single species, the lowest hier- archical element of the biological diversity.

Background AtNHX1, the most abundant vacuolar Na+/H+ antiporter in Arabidopsis thaliana, mediates the transport of Na+ and K+ into the vacuole, influencing plant development and contributing to salt tolerance. In this report, microarray expression profiles of wild type plants, a T-DNA insertion knockout mutant of AtNHX1 (nhx1), and a 'rescued' line (NHX1::nhx1) were exposed to both short (12 h and 48 h) and long (one and two weeks) durations of a non-lethal salt stress to identify key gene transcripts associated with the salt response that are influenced by AtNHX1. Results 147 transcripts showed both salt responsiveness and a significant influence of AtNHX1. Fifty-seven of these genes showed an influence of the antiporter across all salt treatments, while the remaining genes were influenced as a result of a particular duration of salt stress. Most (69%) of the genes were up-regulated in the absence of AtNHX1, with the exception of transcripts encoding proteins involved with metabolic and energy processes that were mostly down-regulated. Conclusion While part of the AtNHX1-influenced transcripts were unclassified, other transcripts with known or putative roles showed the importance of AtNHX1 to key cellular processes that were not necessarily limited to the salt stress response; namely calcium signaling, sulfur metabolism, cell structure and cell growth, as well as vesicular trafficking and protein processing. Only a small number of other salt-responsive membrane transporter transcripts appeared significantly influenced by AtNHX1.

Reduced plant productivity induced by silverleaf whitefly (SLW), Bemisia argentifolii Bellows and Perring, is often associated with reduced net photosynthetic rate (P n ). This study was aimed at characterizing the effects of SLW on photosynthetic activity in cotton and the underlying mechanisms. Cotton (Gossypium hirsutum L. cv. Siv'on) plants were examined under SLW-infested and non-infested conditions and subjected to measurements of gas exchange, chlorophyll content and chlorophyll fluorescence. SLW infestation significantly reduced the P n of cotton by 22-54% throughout the day as well as throughout most of the examined ranges of photosynthetic photon flux density (PPFD) and CO 2 concentrations. Stomatal conductance was also reduced by SLW injury, while intercellular CO 2 concentration (C i ) was not affected, suggesting that stomatal limitation is not the major factor limiting P n . This conclusion was further supported by estimates of stomatal limitation, based on P n /C i curves, which were not affected by SLW infestation. On the other hand, non-stomatal limitation induced by SLW was over 2-fold higher than stomatal limitation. Chlorophyll a and b contents were not modified by SLW infestation. Fluorescence ratio (F v /F m ) and the effective quantum yield of PSII were reduced by less than 10%; thus, impaired photochemical reaction is probably not the only factor limiting P n under SLW infestation. In conclusion, SLW-induced P n reduction in cotton was due to non-stomatal factors, possibly induced by impaired carbon fixation and/or carbohydrate export.

Acacia saligna (Labill.) H.Wendl, a potential crop for forage and wood production, is considered highly drought-resistant. The aim of this study was to characterise some of the physiological traits contributing to drought resistance in A. saligna. Two experiments were conducted: (i) 4-year-old A. saligna were grown in the field under dryland and irrigated treatments and (ii) 6-month-old A. saligna were grown in pots and irrigated to replenish 100% of the transpiration demand (control), or 75% 50% or 25% of the control. Soil-water deficits in the field elicited an increase in osmotic potential in phyllodes. Stomatal conductance was negatively correlated with air vapor pressure deficit under drought conditions in both experiments, whereas under irrigation in the field it was correlated with solar radiation. In the field, dry matter (DM) production under irrigation was only 14% greater (not significant) than under dryland. In the pot experiment, DM production was significantly reduced, and water use efficiency (WUE) and chlorophyll content increased with reduced availability of water. The greater WUE induced by drought could have resulted from stomatal regulation and increased chlorophyll content. Carbon isotope ratios were correlated with the WUE, and may be utilised for selection to further improve the WUE of A. saligna under drought conditions.