Genomic insight into the origin, domestication, dispersal, diversification and human selection of Tartary buckwheat.

BACKGROUND: Tartary buckwheat, Fagopyrum tataricum, is a pseudocereal crop with worldwide distribution and high nutritional value. However, the origin and domestication history of this crop remain to be elucidated. RESULTS: Here, by analyzing the population genomics of 567 accessions collected worldwide and reviewing historical documents, we find that Tartary buckwheat originated in the Himalayan region and then spread southwest possibly along with the migration of the Yi people, a minority in Southwestern China that has a long history of planting Tartary buckwheat. Along with the expansion of the Mongol Empire, Tartary buckwheat dispersed to Europe and ultimately to the rest of the world. The different natural growth environments resulted in adaptation, especially significant differences in salt tolerance between northern and southern Chinese Tartary buckwheat populations. By scanning for selective sweeps and using a genome-wide association study, we identify genes responsible for Tartary buckwheat domestication and differentiation, which we then experimentally validate. Comparative genomics and QTL analysis further shed light on the genetic foundation of the easily dehulled trait in a particular variety that was artificially selected by the Wa people, a minority group in Southwestern China known for cultivating Tartary buckwheat specifically for steaming as a staple food to prevent lysine deficiency. CONCLUSIONS: This study provides both comprehensive insights into the origin and domestication of, and a foundation for molecular breeding for, Tartary buckwheat.

Agro-Morphological and Molecular Characterization Reveal Deep Insights in Promising Genetic Diversity and Marker-Trait Associations in Fagopyrum esculentum and Fagopyrum tataricum.

Characterisation of genetic diversity is critical to adequately exploit the potential of germplasm collections and identify important traits for breeding programs and sustainable crop improvement. Here, we characterised the phenotypic and genetic diversity of a global collection of the two cultivated buckwheat species Fagopyrum esculentum and Fagopyrum tataricum (190 and 51 accessions, respectively) using 37 agro-morphological traits and 24 SSR markers. A wide range of variation was observed in both species for most of the traits analysed. The two species differed significantly in most traits, with traits related to seeds and flowering contributing most to differentiation. The accessions of each species were divided into three major phenoclusters with no clear geographic clustering. At the molecular level, the polymorphic SSR markers were highly informative, with an average polymorphic information content (PIC) of over 0.65 in both species. Genetic diversity, as determined by Nei's expected heterozygosity (He), was high (He = 0.77 and He = 0.66, respectively) and differed significantly between species (p = 0.03) but was homogeneously distributed between regions, confirming the lack of genetic structure as determined by clustering approaches. The weak genetic structure revealed by the phenotypic and SSR data and the low fixation indices in both species suggested frequent seed exchange and extensive cultivation and selection. In addition, 93 and 140 significant (p < 0.05) marker-trait associations (MTAs) were identified in both species using a general linear model and a mixed linear model, most of which explained >20% of the phenotypic variation in associated traits. Core collections of 23 and 13 phenotypically and genetically diverse accessions, respectively, were developed for F. esculentum and F. tataricum. Overall, the data analysed provided deep insights into the agro-morphological and genetic diversity and genetic relationships among F. esculentum and F. tataricum accessions and pointed to future directions for genomics-based breeding programs and germplasm management.

Comparative and population genomics of buckwheat species reveal key determinants of flavor and fertility.

Common buckwheat (Fagopyrum esculentum) is an ancient crop with a world-wide distribution. Due to its excellent nutritional quality and high economic and ecological value, common buckwheat is becoming increasingly important throughout the world. The availability of a high-quality reference genome sequence and population genomic data will accelerate the breeding of common buckwheat, but the high heterozygosity due to the outcrossing nature has greatly hindered the genome assembly. Here we report the assembly of a chromosome-scale high-quality reference genome of F. esculentum var. homotropicum, a homozygous self-pollinating variant of common buckwheat. Comparative genomics revealed that two cultivated buckwheat species, common buckwheat (F. esculentum) and Tartary buckwheat (F. tataricum), underwent metabolomic divergence and ecotype differentiation. The expansion of several gene families in common buckwheat, including FhFAR genes, is associated with its wider distribution than Tartary buckwheat. Copy number variation of genes involved in the metabolism of flavonoids is associated with the difference of rutin content between common and Tartary buckwheat. Furthermore, we present a comprehensive atlas of genomic variation based on whole-genome resequencing of 572 accessions of common buckwheat. Population and evolutionary genomics reveal genetic variation associated with environmental adaptability and floral development between Chinese and non-Chinese cultivated groups. Genome-wide association analyses of multi-year agronomic traits with the content of flavonoids revealed that Fh05G014970 is a potential major regulator of flowering period, a key agronomic trait controlling the yield of outcrossing crops, and that Fh06G015130 is a crucial gene underlying flavor-associated flavonoids. Intriguingly, we found that the gene translocation and sequence variation of FhS-ELF3 contribute to the homomorphic self-compatibility of common buckwheat. Collectively, our results elucidate the genetic basis of speciation, ecological adaptation, fertility, and unique flavor of common buckwheat, and provide new resources for future genomics-assisted breeding of this economically important crop.

Nutritional Evaluation of Quinoa Genetic Resources Growing in the Climatic Conditions of Central Europe.

Quinoa displays huge genetic variability and adaptability to distinct climatic conditions. Quinoa seeds are a good source of nutrients; however, the overall nutritional composition and nutrient content is influenced by numerous factors. This study focused on the nutritional and morphologic evaluation of various quinoa genotypes grown in the Czech Republic. Significant differences between years were observed for morphological traits (plant height, inflorescence length, weight of thousand seeds). The weather conditions in the year 2018 were favorable for all the morphological traits. The protein content of quinoa accessions ranged between 13.44 and 20.01% and it was positively correlated to mauritianin. Total phenolic content varied greatly from year to year, while the antioxidant activity remained relatively stable. The most abundant phenolic compounds were the flavonoids miquelianin, rutin, and isoquercetin. Isoquercetin, quercetin, and N-feruoloyl octopamine showed the highest stability under variable weather conditions in the analyzed years. A total of six compounds were detected and quantified in quinoa for the first time. Most varieties performed well under Central European conditions and can be considered a good source of nutrients and bioactive compounds. These data can be used as a source of information for plant breeders aiming to improve the quality traits of quinoa.

Rewiring of the seed metabolome during Tartary buckwheat domestication.

Crop domestication usually leads to the narrowing genetic diversity. However, human selection mainly focuses on visible traits, such as yield and plant morphology, with most metabolic changes being invisible to the naked eye. Buckwheat accumulates abundant bioactive substances, making it a dual-purpose crop with excellent nutritional and medical value. Therefore, examining the wiring of these invisible metabolites during domestication is of major importance. The comprehensive profiling of 200 Tartary buckwheat accessions exhibits 540 metabolites modified as a consequence of human selection. Metabolic genome-wide association study illustrates 384 mGWAS signals for 336 metabolites are under selection. Further analysis showed that an R2R3-MYB transcription factor FtMYB43 positively regulates the synthesis of procyanidin. Glycoside hydrolase gene FtSAGH1 is characterized as responsible for the release of active salicylic acid, the precursor of aspirin and indispensably in plant defence. UDP-glucosyltransferase gene FtUGT74L2 is characterized as involved in the glycosylation of emodin, a major medicinal component specific in Polygonaceae. The lower expression of FtSAGH1 and FtUGT74L2 were associated with the reduction of salicylic acid and soluble EmG owing to domestication. This first large-scale metabolome profiling in Tartary buckwheat will facilitate genetic improvement of medicinal properties and disease resistance in Tartary buckwheat.

Breeding Buckwheat for Nutritional Quality in the Czech Republic.

Buckwheat is a nutritionally valuable crop, an alternative to common cereals also usable in gluten-free diets. The selection of buckwheat genotypes suitable for further breeding requires the characterization and evaluation of genetic resources. The main objective of this work was to evaluate selected phenotypic and morphological traits using international buckwheat descriptors, including total phenolic content and antioxidant activity, on a unique set of 136 common buckwheat accessions grown in 2019-2020 under Czech Republic conditions. In addition, UHPLC-ESI- MS/MS was used to analyze a wide spectrum of 20 phenolic compounds in buckwheat seeds, including four flavanols, three phenolic acids, seven flavonols, four flavones, and two flavanones. Significant differences among years and genotypes were observed for morphological traits (plant height and 1000-seed weight) and antioxidant activity, as well as levels of observed chemical compounds. Antioxidant activity, crude protein content, plant height and rutin content were characterized by higher mean values in 2020 than in 2019 and vice versa for total polyphenol content and 1000-seed weight. Crude protein content was the most stable across years, while total polyphenol content and rutin content varied greatly from year to year. The most abundant phenolic compounds were rutin, hyperoside, epicatechin, catechin, vitexin, isovitexin, orientin and isoorientin. Protein content was negatively correlated with plant height, catechin and epicatechin content. On the other hand, AA and TPC were positively correlated with rutin, hyperoside and chlorogenic acid. Five accessions showed high stability of the evaluated traits under changing conditions within both years of observation. These materials can be used in breeding programmes aimed at improving buckwheat genotypes with emphasis on quality traits.

Effect of irrigation, fertiliser type and variety on grain yield and nutritional quality of spelt wheat (Triticum spelta) grown under semi-arid conditions.

Previous studies reported higher antioxidant and mineral micronutrient concentrations in organic compared to conventional wheat flour, but the reasons are poorly understood. Here we report results from a long-term, factorial field experiment designed to assess effects of variety choice, supplementary irrigation and contrasting fertilization regimes used in organic and conventional production on the nutritional quality and yield of spelt wheat grown in a semi-arid environment. Long-straw (Oberkulmer, Rubiota, ZOR) varieties had 10-40% higher grain Cu, Fe, Mn and Zn concentrations, while the modern, short straw variety Filderstolz had 15-38% higher grain antioxidant activity. Supplementary irrigation and the use of manure instead of mineral NPK as fertilizer had no substantial effect on the nutritional composition of spelt grain, but increased grain yields by ~ 150 and ~ 18% respectively. Overall, this suggests that breeding/variety selection is the most promising approach to improve the nutritional quality of spelt grain in semi-arid production environments.

Resistance of Winter Spelt Wheat [Triticum aestivum subsp. spelta (L.) Thell.] to Fusarium Head Blight.

Resistance to Fusarium head blight (FHB) of spelt wheat was investigated in field trials carried out at three European locations between 2016 and 2018. Resistance was assessed after artificial inoculation by visual scoring of symptoms and the determination of the contamination of grains and glumes with the mycotoxin deoxynivalenol (DON). It was found that typical spelt traits such as tall plant height, lax spikes, and tough glumes play a role as passive resistance factors. Across all test environments, modern spelt varieties with a significantly reduced plant height showed a significantly higher susceptibility to FHB and a higher contamination of the grains with DON compared to old landraces/varieties and plant genetic resources. Similarly, the lowest mycotoxin levels in grains were found only in old landraces and varieties, while the highest DON concentration was observed mainly in modern varieties. The results obtained can be used for the selection of suitable parental material for breeding spelt with improved FHB resistance.

Resequencing of global Tartary buckwheat accessions reveals multiple domestication events and key loci associated with agronomic traits.

BACKGROUND: Tartary buckwheat (Fagopyrum tataricum) is a nutritionally balanced and flavonoid-rich crop plant that has been in cultivation for 4000 years and is now grown globally. Despite its nutraceutical and agricultural value, the characterization of its genetics and its domestication history is limited. RESULTS: Here, we report a comprehensive database of Tartary buckwheat genomic variation based on whole-genome resequencing of 510 germplasms. Our analysis suggests that two independent domestication events occurred in southwestern and northern China, resulting in diverse characteristics of modern Tartary buckwheat varieties. Genome-wide association studies for important agricultural traits identify several candidate genes, including FtUFGT3 and FtAP2YT1 that significantly correlate with flavonoid accumulation and grain weight, respectively. CONCLUSIONS: We describe the domestication history of Tartary buckwheat and provide a detailed resource of genomic variation to allow for genomic-assisted breeding in the improvement of elite cultivars.

Fagopyrum esculentum ssp. ancestrale-A Hybrid Species Between Diploid F. cymosum and F. esculentum.

Fagopyrum cymosum is considered as most probable wild ancestor of cultivated buckwheat. However, the evolutionary route from F. cymosum to F. esculentum remains to be deciphered. We hypothesized that a hybrid species exists in natural habitats between diploid F. cymosum and F. esculentum. The aim of this research was to determine the phylogenetic position of F. esculentum ssp. ancestrale and to provide new thoughts on buckwheat evolution. Different methodologies including evaluation of morphological traits, determination of secondary metabolites, fluorescence in situ hybridization (FISH), comparative chloroplast genomics, and molecular markers were deployed to determine the phylogenetic relationship of F. esculentum ssp. ancestrale with F. cymosum and F. esculentum. The ambiguity observed in morphological pattern of genetic variation in three species revealed that F. esculentum ssp. ancestrale is closely related to F. cymosum and F. esculentum. Flavonoid analysis revealed that F. esculentum ssp. ancestrale is closely related to F. esculentum. Comparative chloroplast genome analysis further supported the close proximity of F. esculentum ssp. ancestrale with F. esculentum. Additionally, molecular marker analysis revealed that F. esculentum ssp. ancestrale exhibits co-dominance with the bands amplified by F. cymosum and F. esculentum. These finding provided supporting evidence in favor of the hypothesis that F. esculentum ssp. ancestrale is a hybrid species between F. cymosum to F. esculentum, which was probably originated by spontaneous hybridization under natural conditions.

Revisiting the versatile buckwheat: reinvigorating genetic gains through integrated breeding and genomics approach.

MAIN CONCLUSION: Emerging insights in buckwheat molecular genetics allow the integration of genomics driven breeding to revive this ancient crop of immense nutraceutical potential from Asia. Out of several thousand known edible plant species, only four crops-rice, wheat, maize and potato provide the largest proportion of daily nutrition to billions of people. While these crops are the primary supplier of carbohydrates, they lack essential amino acids and minerals for a balanced nutrition. The overdependence on only few crops makes the future cropping systems vulnerable to the predicted climate change. Diversifying food resources through incorporation of orphan or minor crops in modern cropping systems is one potential strategy to improve the nutritional security and mitigate the hostile weather patterns. One such crop is buckwheat, which can contribute to the agricultural sustainability as it grows in a wide range of environments, requires relatively low inputs and possess balanced amino acid and micronutrient profiles. Additionally, gluten-free nature of protein and nutraceutical properties of secondary metabolites make the crop a healthy alternative of wheat-based diet in developed countries. Despite enormous potential, efforts for the genetic improvement of buckwheat are considerably lagged behind the conventional cereal crops. With the draft genome sequences in hand, there is a great scope to speed up the progress of genetic improvement of buckwheat. This article outlines the state of the art in buckwheat research and provides concrete perspectives how modern breeding approaches can be implemented to accelerate the genetic gain. Our suggestions are transferable to many minor and underutilized crops to address the issue of limited genetic gain and low productivity.

Jasmonate-responsive MYB factors spatially repress rutin biosynthesis in Fagopyrum tataricum.

Jasmonates are plant hormones that induce the accumulation of many secondary metabolites, such as rutin in buckwheat, via regulation of jasmonate-responsive transcription factors. Here, we report on the identification of a clade of jasmonate-responsive subgroup 4 MYB transcription factors, FtMYB13, FtMYB14, FtMYB15, and FtMYB16, which directly repress rutin biosynthesis in Fagopyrum tataricum. Immunoblot analysis showed that FtMYB13, FtMYB14, and FtMYB15 could be degraded via the 26S proteasome in the COI1-dependent jasmonate signaling pathway, and that this degradation is due to the SID motif in their C-terminus. Yeast two-hybrid and bimolecular fluorescence complementation assays revealed that FtMYB13, FtMYB14, and FtMYB15 interact with the importin protein Sensitive to ABA and Drought 2 (FtSAD2) in stem and inflorescence. Furthermore, the key repressor of jasmonate signaling FtJAZ1 specifically interacts with FtMYB13. Point mutation analysis showed that the conserved Asp residue of the SID domain contributes to mediating protein-protein interaction. Protoplast transient activation assays demonstrated that FtMYB13, FtMYB14, and FtMYB15 directly repress phenylalanine ammonia lyase (FtPAL) gene expression, and FtSAD2 and FtJAZ1 significantly promote the repressing activity of FtMYBs. These findings may ultimately be promising for further engineering of plant secondary metabolism.

Higher antioxidant and lower cadmium concentrations and lower incidence of pesticide residues in organically grown crops: a systematic literature review and meta-analyses.

Demand for organic foods is partially driven by consumers' perceptions that they are more nutritious. However, scientific opinion is divided on whether there are significant nutritional differences between organic and non-organic foods, and two recent reviews have concluded that there are no differences. In the present study, we carried out meta-analyses based on 343 peer-reviewed publications that indicate statistically significant and meaningful differences in composition between organic and non-organic crops/crop-based foods. Most importantly, the concentrations of a range of antioxidants such as polyphenolics were found to be substantially higher in organic crops/crop-based foods, with those of phenolic acids, flavanones, stilbenes, flavones, flavonols and anthocyanins being an estimated 19 (95 % CI 5, 33) %, 69 (95 % CI 13, 125) %, 28 (95 % CI 12, 44) %, 26 (95 % CI 3, 48) %, 50 (95 % CI 28, 72) % and 51 (95 % CI 17, 86) % higher, respectively. Many of these compounds have previously been linked to a reduced risk of chronic diseases, including CVD and neurodegenerative diseases and certain cancers, in dietary intervention and epidemiological studies. Additionally, the frequency of occurrence of pesticide residues was found to be four times higher in conventional crops, which also contained significantly higher concentrations of the toxic metal Cd. Significant differences were also detected for some other (e.g. minerals and vitamins) compounds. There is evidence that higher antioxidant concentrations and lower Cd concentrations are linked to specific agronomic practices (e.g. non-use of mineral N and P fertilisers, respectively) prescribed in organic farming systems. In conclusion, organic crops, on average, have higher concentrations of antioxidants, lower concentrations of Cd and a lower incidence of pesticide residues than the non-organic comparators across regions and production seasons.

Variation of polyphenols and betaines in aerial parts of young, field-grown Amaranthus genotypes.

Amaranthus hybridus and Amaranthus mantegazzianus are commonly cultivated and the entire young fresh plants consumed as vegetables in regions of Africa and Asia. A. hybridus and A. mantegazzianus were cultivated at four sites in three climate regions of the world: Santa Rosa, Argentina; Lleida, Spain; and Prague and Olomouc, both in the Czech Republic. The contents of flavonoids (isoquercitrin, rutin, nicotiflorin), hydroxybenzoic acids (protocatechuic acid, 4-hydroxybenzoic acid, vanillic acid), hydroxycinnamic acids (caffeic acid, p-coumaric acid, ferulic acid), hydroxycinnamyl amides (N-trans-feruloyltyramine, N-trans-feruloyl-4-O-methyldopamine), and betaines (glycinebetaine, trigonelline) were determined. The variation in phytochemical content due to species and cultivation site was analyzed utilizing the multivariate statistical methods of principal component analysis (PCA) and graphical model (GM). The Argentinean samples differed from the three other locations due to higher contents of most compounds. The samples from Spain and the Czech Republic differed from each other in the content of the negatively correlated metabolites trigonelline and the flavonoids. The two amaranth species were separated primarily by a higher content of trigonelline and the two hydroxycinnamyl amides in A. mantegazzianus. The GM showed that the quantities of the different analytes within each compound group were intercorrelated except in the case of the betaines. The betaines carried no information on each other that was not given through correlations with other compounds. The hydroxycinnamic acids were a key group of compounds in this analysis as they separated the other groups from each other (i.e., carried information on all of the other groups). This study showed the contents of polyphenols and betaines in the aerial parts of vegetable amaranth to be very dependent on growth conditions, but also revealed that some of the compounds (trigonelline and the two hydroxycinnamyl amides) may be useful as features of a taxonomic classification.

Chemistry and pharmacology of Rhaponticum carthamoides: a review.

Rhaponticum carthamoides (Willd.) Iljin is a perennial herb, commonly known as a maral root or Russian leuzea, which has been used for centuries in eastern parts of Russia for its marked medicinal properties. This review based on 117 literary sources, with many of them being originally published in non-English languages (mainly in Russian), discusses the current knowledge of traditional uses, chemistry, biological effects and toxicity of this species. Several different classes of compounds were previously isolated from various parts of R. carthamoides of which the main groups are steroids, particularly ecdysteroids, and phenolics (flavonoids and phenolic acids) accompanied with polyacetylenes, sesquiterpene lactones, triterpenoid glycosides and terpenes (essential oil). A comprehensive account of the chemical constituents is given in this review (figures of 120 structures are shown). Various types of preparations, extracts and individual compounds derived from this species have been found to possess a broad spectrum of pharmacological effects on several organs such as the brain, blood, cardiovascular and nervous systems as well as on different biochemical processes and physiological functions including proteosynthesis, work capacity, reproduction, and sexual function. Moreover, the extracts and preparations from the plant, which are hopefully safe, exhibited various additional biological effects e.g. antioxidant, immunomodulatory, anticancerogenic, antimicrobial, antiparasitic and insect antifeedant or repellent activities. The results of data analysis on the chemical, pharmacological and toxicological characteristics of R. carthamoides support the view that this species has beneficial therapeutic properties and indicate its potential as an effective adaptogenic herbal remedy. Finally, some suggestions for further research on chemical and pharmacological properties are given in this review.