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Soybean is a major crop in the world and an essential source for minerals, including iron (Fe) and zin (Zn). Deficiency of Fe and Zn in soil and soybean plants result in yield loss and poor seed nutritional qualities. Information on genomic regions and candidate genes controlling seed Fe and Zn accumulation in soybean seed is limited. Therefore, The objective of this research was to identify genetic regions, known as quantitative trait loci (QTL), and candidate genes that control the accumulation of Fe and Zn in soybean mature seeds. A ‘Forrest’ by ‘Williams 82’ (F × W82) recombinant inbred line (RIL) population (n = 306) was used and genotyped using a total of 5405 single nucleotides polymorphism (SNP) markers using Infinium SNP6K BeadChips. A two-year experiment was conducted across two environments: North Carolina in 2018 (NC) and Illinois in 2020 (IL). Only QTL with LOD scores ≥ 2.5, as identified by the composite interval mapping (CIM) method, are reported here. In total, 6 QTL were identified for seed Fe; specifically, 3 QTL (qFe-01-[NC-2018], qFe-02-[NC-2018], and qFe-03-[NC-2018]) were located on chromosomes 1, 2, and 6, respectively, in the NC environment, and 3 QTL (qFe-01-[IL-2020], qFe-02-[IL-2020], and qFe-03-[IL-2020]) were positioned on chromosomes 1, 2, and 12, respectively, in the IL environment. A total of 6 QTL associated with seed Zn were also identified; 4 QTL (qZn-01-[NC-2018]; qZn-02-[NC-2018]; qZn-03-[NC-2018]; and qZn-04-[NC-2018]), respectively on Chr 2, 3, 7, and 19 in NC; and 2 QTL (qZn-01-[IL-2020] and qZn-02-[IL-2020]), respectively, on Chr 5 and 8 in IL. Several functional genes encode Fe- and Zn-proteins, transcription factors, proteins-zinc finger motifs (involved in DNA binding and transcriptional regulation; crosstalk between the regulatory pathways of Zn and Fe transporters) were identified and located within the QTL interval. To our knowledge, and based on the literature available, the QTL identified here on Chr 2 and Chr 6 are novel and were not previously identified. This current research provides a new knowledge of the genetic basis of seed Fe and Zn and the markers associated with QTL. The QTL identified here will contribute to efficient marker assisted selection for higher Fe and Zn content in soybean seeds. The candidate genes and metal-responsive transcription factors may coordinate the expression of both Zn and Fe transporters in response to changes in metal availability, providing new knowledge on minerals uptake and transport mechanisms, allowing for possible genetic engineering application.

Understanding the genetic basis of seed Ni and Mo is essential. Since soybean is a major crop in the world and a major source for nutrients, including Ni and Mo, the objective of the current research was to map genetic regions (quantitative trait loci, QTL) linked to Ni and Mo concentrations in soybean seed. A recombinant inbred line (RIL) population was derived from a cross between ‘Forrest’ and ‘Williams 82’ (F × W82). A total of 306 lines was used for genotyping using 5405 single nucleotides polymorphism (SNP) markers using Infinium SNP6K BeadChips. A two-year experiment was conducted and included the parents and the RIL population. One experiment was conducted in 2018 in North Carolina (NC), and the second experiment was conducted in Illinois in 2020 (IL). Logarithm of the odds (LOD) of ≥2.5 was set as a threshold to report identified QTL using the composite interval mapping (CIM) method. A wide range of Ni and Mo concentrations among RILs was observed. A total of four QTL (qNi-01, qNi-02, and qNi-03 on Chr 2, 8, and 9, respectively, in 2018, and qNi-01 on Chr 20 in 2020) was identified for seed Ni. All these QTL were significantly (LOD threshold > 2.5) associated with seed Ni, with LOD scores ranging between 2.71–3.44, and with phenotypic variance ranging from 4.48–6.97%. A total of three QTL for Mo (qMo-01, qMo-02, and qMo-03 on Chr 1, 3, 17, respectively) was identified in 2018, and four QTL (qMo-01, qMo-02, qMo-03, and qMo-04, on Chr 5, 11, 14, and 16, respectively) were identified in 2020. Some of the current QTL had high LOD and significantly contributed to the phenotypic variance for the trait. For example, in 2018, Mo QTL qMo-01 on Chr 1 had LOD of 7.8, explaining a phenotypic variance of 41.17%, and qMo-03 on Chr 17 had LOD of 5.33, with phenotypic variance explained of 41.49%. In addition, one Mo QTL (qMo-03 on Chr 14) had LOD of 9.77, explaining 51.57% of phenotypic variance related to the trait, and another Mo QTL (qMo-04 on Chr 16) had LOD of 7.62 and explained 49.95% of phenotypic variance. None of the QTL identified here were identified twice across locations/years. Based on a search of the available literature and of SoyBase, the four QTL for Ni, identified on Chr 2, 8, 9, and 20, and the five QTL associated with Mo, identified on Chr 1, 17, 11, 14, and 16, are novel and not previously reported. This research contributes new insights into the genetic mapping of Ni and Mo, and provides valuable QTL and molecular markers that can potentially assist in selecting Ni and Mo levels in soybean seeds.

Soybean seed sugars are among the most abundant beneficial compounds for human and animal consumption in soybean seeds. Higher seed sugars such as sucrose are desirable as they contribute to taste and flavor in soy-based food. Therefore, the objectives of this study were to use the ‘Forrest’ by ‘Williams 82’ (F × W82) recombinant inbred line (RIL) soybean population (n = 309) to identify quantitative trait loci (QTLs) and candidate genes that control seed sugar (sucrose, stachyose, and raffinose) contents in two environments (North Carolina and Illinois) over two years (2018 and 2020). A total of 26 QTLs that control seed sugar contents were identified and mapped on 16 soybean chromosomes (chrs.). Interestingly, five QTL regions were identified in both locations, Illinois and North Carolina, in this study on chrs. 2, 5, 13, 17, and 20. Amongst 57 candidate genes identified in this study, 16 were located within 10 Megabase (MB) of the identified QTLs. Amongst them, a cluster of four genes involved in the sugars’ pathway was collocated within 6 MB of two QTLs that were detected in this study on chr. 17. Further functional validation of the identified genes could be beneficial in breeding programs to produce soybean lines with high beneficial sucrose and low raffinose family oligosaccharides.

Abstract Paraquat is used as a harvest‐aid to desiccate green tissues for increasing harvest efficiency and maintaining seed quality. However, its application can cause significant crop damage and yield loss if applied too early. Limited information is available on determining the optimum time for applying paraquat. Therefore, the objectives of this study were to investigate the effects of the timing (critical stages of seed‐fill) of paraquat application on soybean seed yield, seed quality (germination, viability, hard‐seed, and seed damage), and seed composition. Field experiments were conducted in 2019 and 2020 at Stoneville, MS. Paraquat was applied at a rate of 0.56 kg a.i. ha–1 at growth stages R6 (full seed‐fill), R6.5 (pod cavities completely filled with seeds), or R7 (yellow color/beginning maturity). Cultivars P46A57BX and P48A60X were used. The results showed that the application of paraquat at R6 or R6.5 resulted in significant yield loss for both cultivars in both years, whereas application at R7 resulted in significant yield loss for P46A57BX in both years, but in only 1 yr for P48A60X. Seed germination and viability were significantly increased over the control in 2020 for both cultivars at all three application stages, but with mixed effects in 2019. No seed damage that would result in dockage was observed in any treatment, as seed damage for all treatments was below 2%. Application of paraquat at R6 resulted in significantly higher seed protein, oleic acid, raffinose, and stachyose but lower oil and sucrose. This research demonstrated that the harvest‐aid paraquat significantly reduced seed yield, increased seed protein, oleic acid, raffinose, and stachyose when applied before growth stage R7. Therefore, producers should use caution when applying paraquat for harvest efficiency before R7, as they will also likely reduce seed yield the earlier paraquat is applied.

Charcoal rot, caused by Macrophomina phaseolina, is a major soybean disease resulting in significant yield loss and poor seed quality. Currently, no resistant soybean cultivar is available in the market and resistance mechanisms to charcoal rot are unknown, although the disease is believed to infect plants from infected soil through the roots by unknown toxin-mediated mechanisms. The objective of this research was to investigate the association between seed sugars (sucrose, raffinose, stachyose, glucose, and fructose) and their role as biomarkers in the soybean defense mechanism in the moderately resistant (MR) and susceptible (S) genotypes to charcoal rot. Seven MR and six S genotypes were grown under irrigated (IR) and non-irrigated (NIR) conditions. A two-year field experiment was conducted in 2012 and 2013 at Jackson, TN, USA. The main findings in this research were that MR genotypes generally had the ability to maintain higher seed levels of sucrose, glucose, and fructose than did S genotypes. Conversely, susceptible genotypes showed a higher level of stachyose and lower levels of sucrose, glucose, and fructose. This was observed in 6 out of 7 MR genotypes and in 4 out of 6 S genotypes in 2012; and in 5 out of 7 MR genotypes and in 5 out of 6 S genotypes in 2013. The response of S genotypes with higher levels of stachyose and lower sucrose, glucose, and fructose, compared with those of MR genotypes, may indicate the possible role of these sugars in a defense mechanism against charcoal rot. It also indicates that nutrient pathways in MR genotypes allowed for a higher influx of nutritious sugars (sucrose, glucose, and fructose) than did S genotypes, suggesting these sugars as potential biomarkers for selecting MR soybean plants after harvest. This research provides new knowledge on seed sugars and helps in understanding the impact of charcoal rot on seed sugars in moderately resistant and susceptible genotypes.

Isoflavones are secondary metabolites that are abundant in soybean and other legume seeds providing health and nutrition benefits for both humans and animals. The objectives of this study were to construct a single nucleotide polymorphism (SNP)-based genetic linkage map using the ‘Forrest’ by ‘Williams 82’ (F×W82) recombinant inbred line (RIL) population (n = 306); map quantitative trait loci (QTL) for seed daidzein, genistein, glycitein, and total isoflavone contents in two environments over two years (NC-2018 and IL-2020); identify candidate genes for seed isoflavone. The FXW82 SNP-based map was composed of 2075 SNPs and covered 4029.9 cM. A total of 27 QTL that control various seed isoflavone traits have been identified and mapped on chromosomes (Chrs.) 2, 4, 5, 6, 10, 12, 15, 19, and 20 in both NC-2018 (13 QTL) and IL-2020 (14 QTL). The six QTL regions on Chrs. 2, 4, 5, 12, 15, and 19 are novel regions while the other 21 QTL have been identified by other studies using different biparental mapping populations or genome-wide association studies (GWAS). A total of 130 candidate genes involved in isoflavone biosynthetic pathways have been identified on all 20 Chrs. And among them 16 have been identified and located within or close to the QTL identified in this study. Moreover, transcripts from four genes (Glyma.10G058200, Glyma.06G143000, Glyma.06G137100, and Glyma.06G137300) were highly abundant in Forrest and Williams 82 seeds. The identified QTL and four candidate genes will be useful in breeding programs to develop soybean cultivars with high beneficial isoflavone contents.

Charcoal rot is a major disease of soybean (Glycine max) caused by Macrophomina phaseolina and results in significant loss in yield and seed quality. The effects of charcoal rot on seed composition (seed protein, oil, and fatty acids), a component of seed quality, is not well understood. Therefore, the objective of this research was to investigate the impact of charcoal rot on seed protein, oil, and fatty acids in different soybean genotypes differing in their charcoal rot susceptibility under irrigated and non-irrigated conditions. Two field experiments were conducted in 2012 and 2013 in Jackson, TN, USA. Thirteen genotypes differing in charcoal rot resistance (moderately resistant and susceptible) were evaluated. Under non-irrigated conditions, moderately resistant genotypes showed either no change or increased protein and oleic acid but had lower linolenic acid. Under non-irrigated conditions, most of the susceptible genotypes showed lower protein and linolenic acid but higher oleic acid. Most of the moderately resistant genotypes had higher protein than susceptible genotypes under irrigated and non-irrigated conditions but lower oil than susceptible genotypes. The different responses among genotypes for protein, oil, oleic acid, and linolenic acid observed in each year may be due to both genotype tolerance to drought and environmental conditions, especially heat differences in each year (2012 was warmer than 2013). This research showed that the increases in protein and oleic acid and the decrease in linolenic acid may be a possible physiological mechanism underlying the plant’s responses to the charcoal rot infection. This research further helps scientists understand the impact of irrigated and non-irrigated conditions on seed nutrition changes, using resistant and susceptible genotypes.

Interest in using harvest aids (defoliants or crop desiccants) such as paraquat, carfentrazone-ethyl, glyphosate, and sodium chlorate (NaClO3) have become increasingly important to assure harvest efficiency, producer profit, and to maintain seed quality. However, information on the effects of harvest aids on seed nutrition (composition) (protein, oil, fatty acids, sugars, and amino acids) in soybean is very limited. The objective of this research was to investigate the influence of harvest aids on seed protein, oil, fatty acids, sugars, and amino acids in soybean. Our hypothesis was that harvest aid may influence seed nutrition, especially at R6 as at R6 the seeds may still undergo biochemical changes. Field experiments were conducted in 2012 and 2013 under Midsouth USA environmental conditions in which harvest aids were applied at R6 (seed-fill) and R7 (yellow pods) growth stages. Harvest aids applied included an untreated control, 0.28 kg ai ha−1 of paraquat, 0.28 kg ai ha−1 of paraquat, and 1.015 kg ai ha−1 of carfentrazone-ethyl (AIM); 6.72 kg ai ha−1 sodium chlorate, 1.015 kg ai ha−1 carfentrazone-ethyl; and 2.0 kg ae ha−1 glyphosate. Results showed that the application of harvest aids at either R6 or R7 resulted in the alteration of some seed composition such as protein, oil, oleic acid, fructose, and little effects on amino acids. In addition, harvest aids affected seed composition constituents differently depending on year and growth stage. This research demonstrated the possible alteration of some nutrients by harvest aids. This research helps growers and scientists to advance the understanding and management of harvest aids and investigate possible effects of harvest aids on seed nutrition.

NPK fertilizer briquettes (NPKBriq) and organically enhanced N fertilizer (OENF), as newly developed fertilizer products, are reported to increase maize (Zea mays L.) yield and N use efficiency, but their effects on maize grain composition are unknown. The objective of this study was to determine the effects of NPKBriq and OENF on the protein, oil, fiber, ash, and starch of maize grain. A field study was conducted at Jackson and Grand Junction, TN, during 2012 and 2013, with NPKBriq, OENF, ammonium sulfate ((NH4)2SO4) (+P and K), and urea (+P and K) as the main treatments and 0, 85, 128, and 170 kg N ha−1 as the sub treatments under a randomized complete block split plot design with four replicates. The fiber concentration was more responsive to the fertilizer source than the protein, oil, ash, and starch concentrations. OENF resulted in a higher fiber concentration than NPKBriq at 85 kg N ha−1 in 2013, averaged over the two sites. Both OENF and NPKBriq had nearly no significant effects on the concentrations of the quality attributes compared with ammonium sulfate and urea. In conclusion, the nutrient-balanced NPKBriq exerts the same or greater effects on maize grain quality relative to the commonly used nutrient management practices of urea (+P and K) and ammonium sulfate (+P and K) under normal weather conditions. OENF is an alternate N source to urea and ammonium sulfate for similar to higher maize grain quality.

Information on the effects of agricultural practices such as seeding rate (S), row spacing (RS), herbicide apical treatment (T), and nitrogen application (N) on soybean seed nutrition (protein, oil, fatty acids, sugars, and amino acids) is limited. Although seed composition (nutrition) constituents are genetically controlled, agricultural practices and environmental conditions significantly influence the amount and quality of seed nutrition. Therefore, the objective of this research was to understand the responses of these seed composition constituents to these practices, the environment, and cultivar differences. Two-field experiments were conducted, in 2015 and 2016, in Milan, TN, USA. The experiments were irrigated with four replications and included: two soybean cultivars, two seeding rates, three different row spacings, two N rates, and Cobra herbicide apical treatment. The results showed significant effects of S, RS, N, and T on some seed composition constituents, including protein; oleic, linolenic, and stearic acids; sugars; and some amino acids. The current research demonstrated that single or twin row with a seeding rate of 40,000 seeds ha−1 resulted in higher protein, oleic, some sugars, and some amino acids. However, a high seeding rate of 56,000 seeds ha−1 resulted in lower protein, oleic acid, some sugars, and some amino acids due to plant competition for soil nutrients. Herbicide apical application of Cobra1X resulted in higher linolenic acid and some amino acids. Application of nitrogen resulted in higher protein, linolenic, and some amino acids. This research is beneficial to the scientific communities, including breeders and physiologists through advancing knowledge on the interactions between cultivars and environment for seed nutritional quality selection, and to soybean producers through consideration of best agricultural management to maintain high seed nutritional qualities.

The timing of harvest is a major factor affecting seed quality in soybean, particularly in Midsouthern USA, when rain during harvest period is not uncommon. The objective of this research was to evaluate the effects of time of harvest on soybean seed quality (seed composition, germination, seed coat boron, and lignin) in high germinability (HG) breeding lines (50% exotic) developed under high heat. The hypothesis was that seeds of HG lines possess physiological and genetic traits for a better seed quality at harvest maturity and delayed harvest. A 2-year field experiment was conducted under irrigated conditions. Results showed that, at harvest maturity, the exotic HG lines had higher seed protein, oleic acid, sugars, seed coat boron, and seed coat lignin, but lower seed oil compared with the non-exotic checks (Control), confirming our hypothesis. At 28 days after harvest maturity (delayed harvest), the content of seed protein, oleic acid, sugars, seed coat boron, and seed coat lignin were higher in some of the HG lines compared with the checks, indicating a possible involvement of these seed constituents, especially seed coat boron and seed coat lignin, in maintaining seed coat integrity and protecting seed coat against physical damage. Highly significant positive correlations were found between germination and seed protein, oleic acid, sugars, and seed coat boron and seed coat lignin. Highly significant negative correlation was found between germination and oil, linoleic acid, seed coat wrinkling, shattering, and hard seed. Yields of some HG lines were competitive with checks. This research demonstrated that time of harvesting is an important factor influencing seed protein and oil production. Also, since high oleic acid is desirable for oxidative stability, shelf-life and biodiesel properties, using HG lines could positively influence these important traits. This result should suggest to breeders of some of the advantages of selecting for high seed coat boron and lignin, and inform growers of the importance of timely harvest for maintaining high seed quality.

Charcoal rot (CR) disease caused by Macrophomina phaseolina is responsible for significant yield losses in soybean production. Among the methods available for controlling this disease, breeding for resistance is the most promising. Progress in breeding efforts has been slow due to the insufficient information available on the genetic mechanisms related to resistance. Genome-wide association studies (GWAS) enable unraveling the genetic architecture of resistance and identification of causal genes. The aims of this study were to identify new sources of resistance to CR in a collection of 459 diverse plant introductions from the USDA Soybean Germplasm Core Collection using field and greenhouse screenings, and to conduct GWAS to identify candidate genes and associated molecular markers. New sources for CR resistance were identified from both field and greenhouse screening from maturity groups I, II, and III. Five significant single nucleotide polymorphism (SNP) and putative candidate genes related to abiotic and biotic stress responses are reported from the field screening; while greenhouse screening revealed eight loci associated with eight candidate gene families, all associated with functions controlling plant defense response. No overlap of markers or genes was observed between field and greenhouse screenings suggesting a complex molecular mechanism underlying resistance to CR in soybean with varied response to different environments; but our findings provide useful information for advancing breeding for CR resistance as well as the genetic mechanism of resistance.

Frogeye leaf spot (FLS), caused by Cercospora sojina, causes significant damage to soybean in the U.S. One control strategy is the use of quinone outside inhibitor (QoI) fungicides. QoI resistant isolates were first reported in Tennessee (TN) in 2010. To investigate the disease dynamics of C. sojina, we collected 437 C. sojina isolates in 2015 from Jackson and Milan, TN and used 40 historical isolates collected from 2006-2009 from TN and ten additional states for comparison. A subset of 186 isolates, including historical isolates, were genotyped for 49 single nucleotide polymorphism (SNP) markers and the QoI resistance locus, revealing 35 unique genotypes. The genotypes clustered into three groups with two groups containing only sensitive isolates and the remaining group containing all resistant isolates and a dominant clonal lineage of 130 isolates. All 477 C. sojina isolates were genotyped for the QoI locus revealing 344 resistant and 133 sensitive isolates. All isolates collected prior to 2015 were QoI sensitive. Both mating type alleles (MAT1-1-1 and MAT1-2) were found in Jackson and Milan, TN and recovered from single lesions suggesting sexual recombination may play a role in the epidemiology of field populations. Analysis of C. sojina isolates using SNP markers proved useful to investigate population diversity and to elaborate on diversity as it relates to QoI resistance and mating type.

The new Early Soybean Production System (ESPS), developed in the Midsouth USA, including the Mississippi delta, resulted in higher yield under irrigated and non-irrigated conditions. However, information on the effects of the agricultural practices such as row-type (RT: twin- vs. single-row), row-spacing, (RS), seeding rate (SR), soil-type (ST) on seed nutrition under the ESPS environment in the Mississippi delta is very limited. Our previous research in the Mississippi delta showed these agricultural practices altered seed nutrients in one cultivar only. However, whether these effects on seed nutrients will be exhibited by other soybean cultivars with earlier and later maturities across multiple years are not yet known. Therefore, the objective of this research was to evaluate the effects of agricultural practices and cultivar (Cv) differences on seed nutrition in clay and sandy soils under ESPS environment of high heat and drought. Two field experiments were conducted; one experiment was conducted in 2009 and 2010, and the other in 2008, 2009, and 2010 under irrigated conditions. Soybean were grown on 102 cm single-rows and on 25 cm twin-rows with 102 cm centers at seeding rates of 20, 30, 40, and 50 seeds m(-2). Two soybean cultivars (94M80 with earlier maturity; and GP 533 with later maturity) were used. Results showed that increasing seeding rate resulted in increases of protein, sucrose, glucose, raffinose, B, and P concentrations on both single- and twin-rows. However, this increase became either constant or declined at the higher rates (40 and 50 seeds m(-2)). Protein and linolenic acid concentrations were higher in GP 533 than in 94M80 on both row-types, but oil and oleic acid concentrations were in 94M80 than GP 533. Generally, cultivar GP 533 accumulated more seed constituents in seeds than 94M80. In 2010, there were no clear responses of seed nutrients to SR increase in both cultivars, perhaps due to drier year and high heat in 2010. It is concluded that RT and SR can alter seed nutrition under clay and sandy soils, especially under high heat and drought conditions as in the Mississippi delta.

Therefore, the objective of the current research was to investigate the effects of foliar B nutrition on seed protein, oil, fatty acids, and sugars under water stress conditions. A repeated greenhouse experiment was conducted using different maturity group (MG) cultivars. Plants were well-watered with no foliar B (W − B), well-watered with foliar B (W + B), water-stressed with no foliar B (WS − B), and water-stressed with foliar B (WS + B). Foliar B was applied at rate of 0.45 kg·ha−1 and was applied twice at flowering and at seed-fill stages. The results showed that seed protein, sucrose, fructose, and glucose were higher in W + B treatment than in W − B, WS + B, and WS − B. The increase in protein in W + B resulted in lower seed oil, and the increase of oleic in WS − B or WS + B resulted in lower linolenic acid. Foliar B resulted in higher nitrogen fixation and water stress resulted in seed δ15N and δ13C alteration. Increased stachyose indicated possible physiological and metabolic changes in carbon and nitrogen pathways and their sources under water stress. This research is beneficial to growers for fertilizer management and seed quality and to breeders to use 15N/14N and 13C/12C ratios and stachyose to select for drought tolerance soybean.