Lipid modulation contributes to heat stress adaptation in peanut.

At the cellular level, membrane damage is a fundamental cause of yield loss at high temperatures (HT). We report our investigations on a subset of a peanut (Arachis hypogaea) recombinant inbred line population, demonstrating that the membrane lipid remodeling occurring at HT is consistent with homeoviscous adaptation to maintain membrane fluidity. A major alteration in the leaf lipidome at HT was the reduction in the unsaturation levels, primarily through reductions of 18:3 fatty acid chains, of the plastidic and extra-plastidic diacyl membrane lipids. In contrast, levels of 18:3-containing triacylglycerols (TGs) increased at HT, consistent with a role for TGs in sequestering fatty acids when membrane lipids undergo remodeling during plant stress. Polyunsaturated acyl chains from membrane diacyl lipids were also sequestered as sterol esters (SEs). The removal of 18:3 chains from the membrane lipids decreased the availability of susceptible molecules for oxidation, thereby minimizing oxidative damage in membranes. Our results suggest that transferring 18:3 chains from membrane diacyl lipids to TGs and SEs is a key feature of lipid remodeling for HT adaptation in peanut. Finally, QTL-seq allowed the identification of a genomic region associated with heat-adaptive lipid remodeling, which would be useful for identifying molecular markers for heat tolerance.

Parsimonious root systems and better root distribution can improve biomass production and yield of soybean.

Enhancing the acquisition of belowground resources has been identified as an opportunity for improving soybean productivity worldwide. Root system architecture is gaining interest as a selection criterion in breeding programs for enhancing soil resource acquisition and developing climate-resilient varieties. Here we are presenting two novel characteristics of soybean root system architecture that improve aboveground growth and yield. Eleven selected soybean genotypes were tested under rain-fed conditions in 2019 and 2020 at two locations in South Carolina, in which one of the locations was characterized by compacted soils. The elite SC breeding line SC07-1518RR, exotic pedigree line N09-12854, and slow wilting line N09-13890 were superior genotypes in terms of biomass production, seed yield, and/or water use efficiency. Genotypes N09-12854 and N09-13890 demonstrated reduced root development (based on total root count and length), likely to restrict belowground growth and allocate more resources for shoot growth. This characteristic, which can be referred as a parsimonious root phenotype, might be advantageous for soybean improvement in high-input production systems (characterized by adequate fertilizer application and soil fertility) that exist in many parts of the world. Genotype SC07-1518RR exhibited a similar strategy: while it maintained its root system at an intermediate size through reduced levels of total root count and length, it selectively distributed more roots at deeper depths (53-70 cm). The increased root distribution of SC07-1518RR at deeper depths in compacted soil indicates its root penetrability and suitability for clayey soils with high penetration resistance. The beneficial root phenotypes identified in this study (parsimonious root development and selective root distribution in deeper depths) and the genotypes that possessed those phenotypes (SC07-1518RR, N09-12854, and N09-13890) will be useful for breeding programs in developing varieties for optimal, drought, and compacted-soil conditions.

Alterations in the leaf lipidome of Brassica carinata under high-temperature stress.

BACKGROUND: Brassica carinata (A) Braun has recently gained increased attention across the world as a sustainable biofuel crop. B. carinata is grown as a summer crop in many regions where high temperature is a significant stress during the growing season. However, little research has been conducted to understand the mechanisms through which this crop responds to high temperatures. Understanding traits that improve the high-temperature adaption of this crop is essential for developing heat-tolerant varieties. This study investigated lipid remodeling in B. carinata in response to high-temperature stress. A commercial cultivar, Avanza 641, was grown under sunlit-controlled environmental conditions in Soil-Plant-Atmosphere-Research (SPAR) chambers under optimal temperature (OT; 23/15°C) conditions. At eight days after sowing, plants were exposed to one of the three temperature treatments [OT, high-temperature treatment-1 (HT-1; 33/25°C), and high-temperature treatment-2 (HT-2; 38/30°C)]. The temperature treatment period lasted until the final harvest at 84 days after sowing. Leaf samples were collected at 74 days after sowing to profile lipids using electrospray-ionization triple quadrupole mass spectrometry. RESULTS: Temperature treatment significantly affected the growth and development of Avanza 641. Both high-temperature treatments caused alterations in the leaf lipidome. The alterations were primarily manifested in terms of decreases in unsaturation levels of membrane lipids, which was a cumulative effect of lipid remodeling. The decline in unsaturation index was driven by (a) decreases in lipids that contain the highly unsaturated linolenic (18:3) acid and (b) increases in lipids containing less unsaturated fatty acids such as oleic (18:1) and linoleic (18:2) acids and/or saturated fatty acids such as palmitic (16:0) acid. A third mechanism that likely contributed to lowering unsaturation levels, particularly for chloroplast membrane lipids, is a shift toward lipids made by the eukaryotic pathway and the channeling of eukaryotic pathway-derived glycerolipids that are composed of less unsaturated fatty acids into chloroplasts. CONCLUSIONS: The lipid alterations appear to be acclimation mechanisms to maintain optimal membrane fluidity under high-temperature conditions. The lipid-related mechanisms contributing to heat stress response as identified in this study could be utilized to develop biomarkers for heat tolerance and ultimately heat-tolerant varieties.

Heat stress elicits remodeling in the anther lipidome of peanut.

Understanding the changes in peanut (Arachis hypogaea L.) anther lipidome under heat stress (HT) will aid in understanding the mechanisms of heat tolerance. We profiled the anther lipidome of seven genotypes exposed to ambient temperature (AT) or HT during flowering. Under AT and HT, the lipidome was dominated by phosphatidylcholine (PC), phosphatidylethanolamine (PE), and triacylglycerol (TAG) species (> 50% of total lipids). Of 89 lipid analytes specified by total acyl carbons:total carbon-carbon double bonds, 36:6, 36:5, and 34:3 PC and 34:3 PE (all contain 18:3 fatty acid and decreased under HT) were the most important lipids that differentiated HT from AT. Heat stress caused decreases in unsaturation indices of membrane lipids, primarily due to decreases in highly-unsaturated lipid species that contained 18:3 fatty acids. In parallel, the expression of Fatty Acid Desaturase 3-2 (FAD3-2; converts 18:2 fatty acids to 18:3) decreased under HT for the heat-tolerant genotype SPT 06-07 but not for the susceptible genotype Bailey. Our results suggested that decreasing lipid unsaturation levels by lowering 18:3 fatty-acid amount through reducing FAD3 expression is likely an acclimation mechanism to heat stress in peanut. Thus, genotypes that are more efficient in doing so will be relatively more tolerant to HT.

Comparative Lipidomic Analysis Reveals Heat Stress Responses of Two Soybean Genotypes Differing in Temperature Sensitivity.

Heat-induced changes in lipidome and their influence on stress adaptation are not well-defined in plants. We investigated if lipid metabolic changes contribute to differences in heat stress responses in a heat-tolerant soybean genotype DS25-1 and a heat-susceptible soybean genotype DT97-4290. Both genotypes were grown at optimal temperatures (OT; 30/20 °C) for 15 days. Subsequently, half of the plants were exposed to heat stress (38/28 °C) for 11 days, and the rest were kept at OT. Leaf samples were collected for lipid and RNA extractions on the 9th and 11th days of stress, respectively. We observed a decline in the lipid unsaturation level due to a decrease in the polyunsaturated linolenic acid (18:3) content in DS25-1. When examined under OT conditions, DS25-1 and DT97-4290 showed no significant differences in the expression pattern of the Fatty Acid Desaturase (FAD) 2-1A, FAD2-2B, FAD2-2C, FAD3A genes. Under heat stress conditions, substantial reductions in the expression levels of the FAD3A and FAD3B genes, which convert 18:2 lipids to 18:3, were observed in DS25-1. Our results suggest that decrease in levels of lipids containing 18:3 acyl chains under heat stress in DS25-1 is a likely consequence of reduced FAD3A and FAD3B expression, and the decrease in 18:3 contributes to DS25-1's maintenance of membrane functionality and heat tolerance.

Evaluation of soybean [Glycine max (L.) Merr.] genotypes for yield, water use efficiency, and root traits.

Drought stress has been identified as the major environmental factor limiting soybean [Glycine max (L.) Merr.] yield worldwide. Current breeding efforts in soybean largely focus on identifying genotypes with high seed yield and drought tolerance. Water use efficiency (WUE) that results in greater yield per unit rainfall is an important parameter in determining crop yields in many production systems, and is often related with crop drought tolerance. Even though roots are major plant organs that perceive and respond to drought stress, their utility in improving soybean yield and WUE under different environmental and management conditions are largely unclear. The objectives of this research was to evaluate soybean cultivars and breeding and germplasm lines for yield, WUE, root penetrability of hardpan, and root morphology. Field experiments were conducted at two locations in South Carolina (southeastern United States) during the 2017 cropping season to test the genotypes for yield and root morphology under irrigated and non-irrigated conditions. Two independent controlled-environmental experiments were conducted to test the genotypes for WUE and root penetrability of synthetic hardpans. The slow wilting lines NTCPR94-5157 and N09-13890 had equal or greater yield than the checks- cultivar NC-Raleigh and the elite South Carolina breeding line SC07-1518RR, under irrigated and non-irrigated conditions. The high yielding genotypes NTCPR94-5157, N09-13890, and SC07-1518RR exhibited root parsimony (reduced root development). This supported the recent hypothesis in literature that root parsimony would have adaptational advantage to improve yield under high input field conditions. The high yielding genotypes NTCPR94-5157, N09-13890, NC-Raleigh, and SC07-1518RR and a cultivar Boggs (intermediate in yield) possessed high WUE and had increased root penetrability of hardpans. These genotypes offer useful genetic materials for soybean breeding programs for improving yield, drought tolerance, and/or hardpan penetrability.

Characterization of a soybean (Glycine max L. Merr.) germplasm collection for root traits.

Root systems that improve resource uptake and penetrate compacted soil (hardpan) are important for improving soybean (Glycine max L. Merr.) productivity in optimal and sub-optimal environments. The objectives of this research were to evaluate a soybean germplasm collection of 49 genotypes for root traits, determine whether root traits are related with plant height, shoot dry weight, chlorophyll index, and seed size, and identify genotypes that can penetrate a hardpan. Plants were maintained under optimal growth conditions in a greenhouse. Single plants were grown in mesocosms, constructed of two stacked columns (top and bottom columns had 25 and 46 cm height, respectively, and 15 cm inside diameter) with a 2-cm thick wax layer (synthetic hardpan; penetration resistance, 1.5 MPa at 30°C) in between. Plants were harvested at 42 days after planting. Significant genetic variability was observed for root traits in the soybean germplasm collection, and genotypes that penetrated the synthetic hardpan were identified. Genotypes NTCPR94-5157, NMS4-1-83, and N09-13128 were ranked high and PI 424007 and R01-581F were ranked low for most root traits. Shoot dry weight and chlorophyll index were positively related with total root length, surface area, and volume, and fine root length (Correlation coefficient, r ≥ 0.60 and P-value < 0.0001 for shoot dry weight and r ≥ 0.37 and P-value < 0.01 for chlorophyll index]. Plant height was negatively correlated with total root surface area, total root volume, and average root diameter (|r| ≥ 0.29, P-value < 0.05). Seed size was not correlated with any root traits. The genetic variability identified in this research for root traits and penetration are critical for soybean improvement programs in choosing genotypes with improved root characteristics to increase yield in stressful or optimum environments.

Alterations in wheat pollen lipidome during high day and night temperature stress.

Understanding the adaptive changes in wheat pollen lipidome under high temperature (HT) stress is critical to improving seed set and developing HT tolerant wheat varieties. We measured 89 pollen lipid species under optimum and high day and/or night temperatures using electrospray ionization-tandem mass spectrometry in wheat plants. The pollen lipidome had a distinct composition compared with that of leaves. Unlike in leaves, 34:3 and 36:6 species dominated the composition of extraplastidic phospholipids in pollen under optimum and HT conditions. The most HT-responsive lipids were extraplastidic phospholipids, phosphatidylcholine (PC), phosphatidylethanolamine (PE), phosphatidylinositol, phosphatidic acid, and phosphatidylserine. The unsaturation levels of the extraplastidic phospholipids decreased through the decreases in the levels of 18:3 and increases in the levels of 16:0, 18:0, 18:1, and 18:2 acyl chains. PC and PE were negatively correlated. Higher PC:PE at HT indicated possible PE-to-PC conversion, lower PE formation, or increased PE degradation, relative to PC. Correlation analysis revealed lipids experiencing coordinated metabolism under HT and confirmed the HT responsiveness of extraplastidic phospholipids. Comparison of the present results on wheat pollen with results of our previous research on wheat leaves suggests that similar lipid changes contribute to HT adaptation in both leaves and pollen, though the lipidomes have inherently distinct compositions.

Wheat leaf lipids during heat stress: II. Lipids experiencing coordinated metabolism are detected by analysis of lipid co-occurrence.

Identifying lipids that experience coordinated metabolism during heat stress would provide information regarding lipid dynamics under stress conditions and assist in developing heat-tolerant wheat varieties. We hypothesized that co-occurring lipids, which are up-regulated or down-regulated together through time during heat stress, represent groups that can be explained by coordinated metabolism. Wheat plants (Triticum aestivum L.) were subjected to 12 days of high day and/or night temperature stress, followed by a 4-day recovery period. Leaves were sampled at four time points, and 165 lipids were measured by electrospray ionization-tandem mass spectrometry. Correlation analysis of lipid levels in 160 leaf samples from each of two wheat genotypes revealed 13 groups of lipids. Lipids within each group co-occurred through the high day and night temperature stress treatments. The lipid groups can be broadly classified as groups containing extraplastidic phospholipids, plastidic glycerolipids, oxidized glycerolipids, triacylglycerols, acylated sterol glycosides and sterol glycosides. Current knowledge of lipid metabolism suggests that the lipids in each group co-occur because they are regulated by the same enzyme(s). The results suggest that increases in activities of desaturating, oxidizing, glycosylating and acylating enzymes lead to simultaneous changes in levels of multiple lipid species during high day and night temperature stress in wheat.

Wheat leaf lipids during heat stress: I. High day and night temperatures result in major lipid alterations.

Understanding how wheat (Triticum aestivum L.) plants under high temperature (HT) regulate lipid composition is critical to developing climate-resilient varieties. We measured 165 glycerolipids and sterol derivatives under optimum and high day and night temperatures in wheat leaves using electrospray ionization-tandem mass spectrometry. Levels of polar lipid fatty acyl chain unsaturation were lower in both heat-tolerant genotype Ventnor and susceptible genotype Karl 92 under HT, compared with optimum temperature. The lower unsaturation was predominantly because of lower levels of 18:3 acyl chains and higher levels of 18:1 and 16:0 acyl chains. Levels of 18:3-containing triacylglycerols increased threefold/more under HT, consistent with their possible role in sequestering fatty acids during membrane lipid remodelling. Phospholipids containing odd-numbered or oxidized acyl chains accumulated in leaves under HT. Sterol glycosides (SG) and 16:0-acylated sterol glycosides (ASG) were higher under HT than optimum temperatures. Ventnor had lower amounts of phospholipids with oxidized acyl chains under HT and higher amounts of SG and 16:0-ASG than Karl 92. Taken together, the data demonstrate that wheat leaf lipid composition is altered by HT, in which some lipids are particularly responsive to HT, and that two wheat genotypes, chosen for their differing physiological responses to HT, differ in lipid profile under HT.

Variability of root traits in spring wheat germplasm.

Root traits influence the amount of water and nutrient absorption, and are important for maintaining crop yield under drought conditions. The objectives of this research were to characterize variability of root traits among spring wheat genotypes and determine whether root traits are related to shoot traits (plant height, tiller number per plant, shoot dry weight, and coleoptile length), regions of origin, and market classes. Plants were grown in 150-cm columns for 61 days in a greenhouse under optimal growth conditions. Rooting depth, root dry weight, root: shoot ratio, and shoot traits were determined for 297 genotypes of the germplasm, Cultivated Wheat Collection (CWC). The remaining root traits such as total root length and surface area were measured for a subset of 30 genotypes selected based on rooting depth. Significant genetic variability was observed for root traits among spring wheat genotypes in CWC germplasm or its subset. Genotypes Sonora and Currawa were ranked high, and genotype Vandal was ranked low for most root traits. A positive relationship (R2 ≥ 0.35) was found between root and shoot dry weights within the CWC germplasm and between total root surface area and tiller number; total root surface area and shoot dry weight; and total root length and coleoptile length within the subset. No correlations were found between plant height and most root traits within the CWC germplasm or its subset. Region of origin had significant impact on rooting depth in the CWC germplasm. Wheat genotypes collected from Australia, Mediterranean, and west Asia had greater rooting depth than those from south Asia, Latin America, Mexico, and Canada. Soft wheat had greater rooting depth than hard wheat in the CWC germplasm. The genetic variability identified in this research for root traits can be exploited to improve drought tolerance and/or resource capture in wheat.