Structural changes in the peach root system under salt stress. (A) Photo of the peach tree root structure, the figures of roots in the second column are where the data were collected by the root analysis software. The collected data are root length, surface area, root volume, tips, and forks. (B) Fine root distribution heat map. The vast majority of root lengths were distributed in the range of 0 < L ≤ 1, and the root number of NaCl + 200 mg/L PC treatment was the highest in the range of 0 < L ≤ 0.5 under salt stress. Measurements were performed using the intact root system of the peach plant. Measurements of root structural were performed at 10:00 am on 11th day of the salt stress treatment. Bars correspond to 5 cm.

Structural changes in the peach root system under salt stress. (A) Photo of the peach tree root structure, the figures of roots in the second column are where the data were collected by the root analysis software. The collected data are root length, surface area, root volume, tips, and forks. (B) Fine root distribution heat map. The vast majority of root lengths were distributed in the range of 0 < L ≤ 1, and the root number of NaCl + 200 mg/L PC treatment was the highest in the range of 0 < L ≤ 0.5 under salt stress. Measurements were performed using the intact root system of the peach plant. Measurements of root structural were performed at 10:00 am on 11th day of the salt stress treatment. Bars correspond to 5 cm.

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Salt stress is a major adverse abiotic factor seriously affecting fruit tree growth and development. It ultimately lowers fruit quality and reduces yield. Phosphatidylcholine (PC) is an important cell membrane component that is critical for cell structure and membrane stability maintenance. In this study, we found that the addition of external PC s...

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... the root hair growth and volume in the soil also determines the potential of plants for nutrient and water absorption. In this research, we found that salt stress considerably reduced the number of absorbing roots ( Figure 3A). However, the PC supplementation applied along with the salt stress treatment obviously alleviated the decrease in the number of absorbing roots ( Figure 3B). ...
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... this research, we found that salt stress considerably reduced the number of absorbing roots ( Figure 3A). However, the PC supplementation applied along with the salt stress treatment obviously alleviated the decrease in the number of absorbing roots ( Figure 3B). As displayed in Table 2, the salt stress treatment decreased the total length, total surface area, and total volume of the root system. ...
Context 3
... the root hair growth and volume in the soil also determines the potential of plants for nutrient and water absorption. In this research, we found that salt stress considerably reduced the number of absorbing roots ( Figure 3A). However, the PC supplementation applied along with the salt stress treatment obviously alleviated the decrease in the number of absorbing roots ( Figure 3B). ...
Context 4
... this research, we found that salt stress considerably reduced the number of absorbing roots ( Figure 3A). However, the PC supplementation applied along with the salt stress treatment obviously alleviated the decrease in the number of absorbing roots ( Figure 3B). As displayed in Table 2, the salt stress treatment decreased the total length, total surface area, and total volume of the root system. ...
Context 5
... present research evidenced that under salt stress, the exogenous application of PC enhanced the activity of PLD and increased the content of PA in the cells. Notably, the exogenous application of PC increased the root growth of the treated peach seedlings as compared with that in the control, especially the number of fine roots ( Figure 3B). Using transmission electron microscope observations of the root cells, we found that the exogenous administration of PC had a positive effect on maintaining cell integrity and cell-membrane structure. ...

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... Plant membranes, as a key biological barrier, protect cells and organelles from the harmful effects of salt stress 25 , and the stability and function of membrane structural integrity are the basis of normal cell metabolism and overall physiology 26 . Under unfavorable conditions such as salt stress, plants can alter the content and composition of membrane lipids to adapt to the effects of adverse factors 27 . The GO analysis revealed that DEGs were increasingly associated with M. azedarach membranes as the severity of salt stress increased (Fig. 2), which corroborates the above arguments. ...
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Melia azedarach demonstrates strong salt tolerance and thrives in harsh saline soil conditions, but the underlying mechanisms are poorly understood. In this study, we analyzed gene expression under low, medium, and high salinity conditions to gain a deeper understanding of adaptation mechanisms of M. azedarach under salt stress. The GO (gene ontology) analysis unveiled a prominent trend: as salt stress intensified, a greater number of differentially expressed genes (DEGs) became enriched in categories related to metabolic processes, catalytic activities, and membrane components. Through the analysis of the category GO:0009651 (response to salt stress), we identified four key candidate genes (CBL7, SAPK10, EDL3, and AKT1) that play a pivotal role in salt stress responses. Furthermore, the KEGG (Kyoto Encyclopedia of Genes and Genomes) pathway enrichment analysis revealed that DEGs were significantly enriched in the plant hormone signaling pathways and starch and sucrose metabolism under both medium and high salt exposure in comparison to low salt conditions. Notably, genes involved in JAZ and MYC2 in the jasmonic acid (JA) metabolic pathway were markedly upregulated in response to high salt stress. This study offers valuable insights into the molecular mechanisms underlying M. azedarach salt tolerance and identifies potential candidate genes for enhancing salt tolerance in M. azedarach.
... The addition of exogenous PC significantly improves the tolerance of annual peach tree (Prunus persica (L.) Batsch.) to salt stress and mitigates its damage [29]. PC can interact with ACBP, enhancing plants' salt tolerance by increasing PLDδ's activity and further converting PC to PE, PS, and PG to stabilize the cell membrane [30]. ...
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Rice (Oryza sativa L.), a crucial food crop that sustains over half the world’s population, is often hindered by salt stress during various growth stages, ultimately causing a decrease in yield. However, the specific mechanism of rice roots’ response to salt stress remains largely unknown. In this study, transcriptomics and lipidomics were used to analyze the changes in the lipid metabolism and gene expression profiles of rice roots in response to salt stress. The results showed that salt stress significantly inhibited rice roots’ growth and increased the roots’ MDA content. Furthermore, 1286 differentially expressed genes including 526 upregulated and 760 downregulated, were identified as responding to salt stress in rice roots. The lipidomic analysis revealed that the composition and unsaturation of membrane lipids were significantly altered. In total, 249 lipid molecules were differentially accumulated in rice roots as a response to salt stress. And most of the major phospholipids, such as phosphatidic acid (PA), phosphatidylcholine (PC), and phosphatidylserine (PS), as well as major sphingolipids including ceramide (Cer), phytoceramide (CerP), monohexose ceramide (Hex1Cer), and sphingosine (SPH), were significantly increased, while the triglyceride (TG) molecules decreased. These results suggested that rice roots mitigate salt stress by altering the fluidity and integrity of cell membranes. This study enhances our comprehension of salt stress, offering valuable insights into changes in the lipids and adaptive lipid remodeling in rice’s response to salt stress.
... Phosphatidylcholine (PC) is a fundamental type of lipid found in the membranes of most eukaryotic cells. Sun et al. discovered that supplementing peach trees, Prunus persica L. Batsch, with external sources of PC greatly improved their salt tolerance after one year of growth (Sun et al. 2020). Plants responding to low temperatures show an increase in the PC: phosphatidylethanolamine (PE) ratio (Welti and Li 2002;Moellering and Benning 2011). ...
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The membranes of plants are where temperature sensing begins and where freezing injury typically occurs. Barley’s adaptation to and survival after freezing stress is aided by remodelling of its membrane lipid composition. The modifications of individual lipid molecular species in different stress-treated plant species and cultivars can indicate the functions of genes regulating lipid metabolism or signaling. In this study, we employed a membrane lipidomic approach to investigate the response of barley of two cold-tolerant and two cold-sensitive cultivars to freezing temperatures during the barley trefoil stage. A total of 56 predominant lipid compounds changed significantly under freezing stress were identified. Phosphatidic acid (PA), lysophosphatidic acid (LPA) and monogalactosyldiacylglycerol (MGDG) in freezing-tolerant varieties were significantly upregulated under freezing stress, while there was a decrease in freezing-sensitive cultivars. Freezing-tolerant varieties experienced greater changes in lipid composition compared to freezing-sensitive cultivars, which had proportionally smaller changes. In addition, when exposed to short-term cold stress, varieties A and B had lower levels of monoglyceride lipase (MGLL) than varieties C and D. However, under long-term cold stress, the opposite was observed. Additionally, the freezing-tolerant variety A showed a notable increase in the expression of diacylglycerol acyltransferase 1 (DGAT1) after being exposed to 4 °C. Furthermore, SENSITIVE TO FREEZING 2 (SFR2) reached its highest level in all four varieties after being exposed to cold treatment for 48 h. This study indicates that freezing injury in barley leaves is correlated with extensive changes in lipid metabolism and that freezing-tolerant varieties can alleviate freezing injury by membrane lipid remodelling. The study’s outcomes may improve our understanding of barley’s freezing adaptation mechanisms and contribute to breeding for better tolerance.
... 9(10)-EpOME may eventually lead to the accumulation of the phosphatidylcholine (PC), and it has been reported that phosphatidylcholine shows an overall reduction in salt-treated barley roots [34]. Some studies have also revealed that phosphatidylcholine enhances homeostasis in peach seedling cell membrane or in Arabidopsis thaliana in response to salt stress [35,36]. The increase in phosphatidylcholine in DH20 leaves under salt stress might have effects on lipid homeostasis, which could sustain cell expansion and the growth of salt-stressed plants [37]. ...
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Barley is the most salt-tolerant cereal crop. However, little attention has been paid to the salt-tolerant doubled haploids of barley derived from mutagenesis combined with isolated micro-spore culture. In the present study, barley doubled haploid (DH) line 20, which was produced by mutagenesis combined with isolated microspore culture, showed stably and heritably better salt tolerance than the wild type H30 in terms of fresh shoot weight, dry shoot weight, K + /Na + ratio and photosynthetic characteristics. Transcriptome and metabolome analyses were performed to compare the changes in gene expression and metabolites between DH20 and H30. A total of 462 differentially expressed genes (DEGs) and 152 differentially accumulated metabolites (DAMs) were identified in DH20 compared to H30 under salt stress. Among the DAMs, fatty acids were the most accumulated in DH20 under salt stress. The integration of transcriptome and metabolome analyses revealed that nine key biomarkers, including two metabolites and seven genes, could distinguish DH20 and H30 when exposed to high salt. The pathways of linoleic acid metabolism, alpha-linolenic acid metabolism, glycerolipid metabolism, photosynthesis, and alanine, aspartate and glutamate metabolism were significantly enriched in DH20 with DEGs and DAMs in response to salt stress. These results suggest that DH20 may enhance resilience by promoting lipid metabolism, maintaining energy metabolism and decreasing amino acids metabolism. The study provided novel insights for the rapid generation of homozygous mutant plants by mutagenesis combined with microspore culture technology and also identified candidate genes and metabolites that may enable the mutant plants to cope with salt stress.
... The increasing PI and PC content was conducive to maintaining membrane integrity and fluidity [46,47]. PC enhanced homeostasis in peach seedling cell membranes, and increased the plant's salt stress tolerance by means of phosphatidic acid (PA) [48]. In addition, W_vs_L2 had more differential lipids compared to W_vs_L1, implying that the functions of CmnsLTP6.9L ...
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Chestnut (Castanea mollissima Blume) is an important economic tree owing to its tasty fruit and adaptability to environmental stresses, especially drought. Currently, there is limited information about non-specific lipid transfer protein (nsLTP) genes that respond to abiotic stress in chestnuts. Here, a chestnut nsLTP, named CmnsLTP6.9, was identified and analyzed. The results showed that the CmnsLTP6.9 protein localized in the extracellular matrix had two splicing variants (CmnsLTP6.9L and CmnsLTP6.9S). Compared with CmnsLTP6.9L, CmnsLTP6.9S had an 87 bp deletion in the 5′-terminal. Overexpression of CmnsLTP6.9L in Arabidopsis enhanced tolerance to osmotic and drought stress. Upon exposure to osmotic and drought treatment, CmnsLTP6.9L could increase reactive oxygen species (ROS)-scavenging enzyme activity, alleviating ROS damage. However, CmnsLTP6.9S-overexpressing lines showed no significant differences in phenotype, ROS content, and related enzyme activities compared with the wild type (WT) under osmotic and drought treatment. Moreover, lipid metabolism analysis confirmed that, unlike CmnsLTP6.9S, CmnsLTP6.9L mainly altered and upregulated many fatty acyls and glycerophospholipids, which implied that CmnsLTP6.9L and CmnsLTP6.9S played different roles in lipid transference in the chestnut. Taken together, we analyzed the functions of CmnsLTP6.9L and CmnsLTP6.9S, and demonstrated that CmnsLTP6.9L enhanced drought and osmotic stress tolerance through ROS scavenging and lipid metabolism.
... 14 It is a precursor in the phosphatidylcholine, which plays a key role in cell membrane structure and keeps in balance the equilibrium of ROS production and removal. 15,16 Chol-BTH, as an ionic liquid, has better physicochemical properties than BTH, such as thermal stability (with a melting point of 114.3°C due to the hydrogen bonding between ions) and a high dissolution rate (3600 s), which may be appropriate for greenhouse application. 17 In our previous study, the efficiency of BTH and some of its derivatives (including Chol-BTH) in antiviral plant response potential in tobacco has been described. ...
... Choline is a precursor to the synthesis of phosphatidylcholine, 61 a key phospholipid in cell membrane maintenance and signaling molecule, which is also up-regulated in salt and drought stress. 15,16 It also has antioxidant activity, which may support GSH in action; therefore, increased glutathione synthesis might not be not necessary. This may explain the decrease in GSH levels for plants treated with Chol-BTH at 8 dpt. ...
Article
Tomatoes are one of the most important vegetables thanks to their taste attributes and nutritional value. Their cultivation is threatened by various pathogens including viruses. The application of resistance inducers (RI), such as benzo(1,2,3)-thiadiazole-7-carbothioic acid S-methyl ester (BTH) may be used to enhance plant performance against viruses. Here we aimed to compare the impact of BTH and its choline derivative (Chol-BTH) on resistance induction and antioxidant properties of healthy plants and tomato mosaic virus (ToMV)-infected ones. The response of tomato plants to treatment with BTH or Chol-BTH was manifested by increased expression of not only pathogenesis-related (PR) genes but also WRKY and Jasmonate Zim-domain protein (JAZ) genes and increased jasmonic acid (JA) levels. The effect of BTH as a resistance inducer was observed early after application, while with Chol-BTH the plant defense system reacted more strongly after 8 days. The antioxidant properties of RI-treated tomatoes are related to both glutathione content and peroxidase activity. In the case of BTH, an increase in these activities occurred early after application, while in the case of Chol-BTH, the glutathione level was particularly high in the plant early after treatment, and high peroxidase activity was observed 8 days post-treatment. Overall, the collected results indicate that Chol-BTH, due to its physicochemical parameters (e.g., good solubility) and biological activity (increased expression of lignification-related genes, supported by increases in peroxidase activity and total phenolic compounds levels), can also be a very useful agent inducing tomato resistance against viral pathogens.
... 42 Phosphatidylcholine application was also found to enhance homeostasis against salt stress. 43 In contrast, few reports are found in the literature on lysophospholipids except for their role as priming agents of the plant immune system and resistance against pathogens. 44 A detailed comparison between phospholipids and lysophospholipids regarding plants' eects is yet to be reported. ...
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Balanites aegyptiaca (L.) Delile (Zygophyllaceae), also known as the desert date, is an edible fruit-producing tree popular for its nutritional and several health benets. In this study, multi-targeted comparative metabolic proling and ngerprinting approaches were conducted for the assessment of the nutrient primary and secondary metabolite heterogeneity in dierent parts, such as leaves, stems, seeds, unripe, and ripe fruits of B. aegyptiaca using nuclear magnetic resonance (NMR), ultra-performance liquid chromatography (UPLC-MS), and gas chromatography mass-spectrometry (GC-MS) based metabolomics coupled to multivariate analyses and in relation to its cytotoxic activities. NMR-based metabolomic study identied and quantied 15 major primary and secondary metabolites belonging to alkaloids, saponins, avonoids, sugars, and amino and fatty acids. Principal component analysis (PCA) of the NMR dataset revealed a-glucose, sucrose, and isorhamnetin as markers for fruit and stem and unsaturated fatty acids for predominated seeds. Orthogonal projections to latent structure discriminant analysis (OPLS-DA) revealed trigonelline as a major distinctive metabolite in the immature fruit and isorhamnetin as a major distinct marker in the mature fruit. UPLC-MS/MS analysis using feature-based molecular networks revealed diverse chemical classes viz. steroidal saponins, N-containing metabolites, phenolics, fatty acids, and lipids as the constitutive metabolome in Balanites. Gas chromatography-mass spectroscopy (GC-MS) proling of primary metabolites led to the detection of 135 peaks belonging to sugars, fatty acids/esters, amino acids, nitrogenous, and organic acids. Monosaccharides were detected at much higher levels in ripe fruit and disaccharides in predominate unripe fruits, whereas B. aegyptiaca vegetative parts (leaves and stem) were rich in amino acids and fatty acids. The antidiabetic compounds, viz, nicotinic acid, and trigonelline, were detected in all parts especially unripe fruit in addition to the sugar alcohol D-pinitol for the rst time providing novel evidence for B. aegyptiaca use in diabetes. In vitro cytotoxic activity revealed the potential ecacy of immature fruit and seeds as cytotoxic agents against human prostate cancer (PC3) and human colorectal cancer (HCT-116) cell lines. Collectively, such detailed proling of parts provides novel evidence for B. aegyptiaca medicinal uses.
... However, the unsaturated fatty acid contents (linoleic, 13-hydroperoxy-9z,11e-octadecadienoic acid) increased with increasing Cd concentration in S. tonkinensis, consistent with previously reported results [54]. Phosphatidylcholine (PC) is involved in cell membrane synthesis [55]. PC is obtained by the addition of a ...
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Sophora tonkinensis is an important medicinal plant native to China and Vietnam, growing in stony mountains and calcareous or sandstone hill slopes. Cadmium negatively affects the quality of S. tonkinensis , indirectly threatening human health. To elucidate the physiological and metabolic mechanism of S. tonkinensis in response to Cd stress, we conducted a hydroponic greenhouse experiment and measured the growth parameters of S. tonkinensis treated with 0, 20, 40, 60, and 80 µmol L − 1 Cd. Furthermore, metabolite changes in S. tonkinensis roots were investigated using liquid chromatography–tandem mass spectrometry. We identified 380 differential metabolites, and significant differences were detected in the varieties and quantities of amino acids, organic acids, fatty acids, and ketones, which have significant impact on S. tonkinensis stress response to Cd, under different Cd concentrations. S. tonkinensis resisted the stress induced by 40 µmol L − 1 Cd by upregulating lysine, arginine, histidine, glycine, sarcosine, tryptophan, D-proline, DL-serine, cellobiose, linoleic, indole, cytosine, and genistein contents and downregulating betaine, leucine, citrate, oleic, trehalose, and daidzein contents. Linoleic, palmitic, citrate, maackiain, isomaltose, and cinnavalininate contents were upregulated, and L-phenylalanine, sarcosine, indole acetate, and cytosine were downregulated in response to 80 µmol L − 1 Cd stress. KEGG pathway enrichment analysis showed that the isoflavonoid biosynthesis pathway was significantly enriched, indicating its important role in the Cd resistance of S. tonkinensis. This study provides important information for understanding the tolerance and accumulation mechanisms of S. tonkinensis in response to Cd stress, which will contribute to the breeding of medicinal plant.
... Although it is usually maintained at low levels in plants, the synthesis of PA can be activated in minutes when plants are exposed to an environmental stimulus (Testerink and Munnik 2005;Okazaki and Saito 2014). The content of PA can be affected by foliar application of other phospholipids, for example, phosphatidylcholine (PC) (Sun et al. 2022); in turn, the deletion of phospholipase D, which is a key enzyme for PA synthesis, resulted in the increases in other lipids [PC and phosphatidylethanolamine (PE)] in Arabidopsis (Song et al. 2021). ...
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Phosphatidic acid (PA) is a key intermediate for lipid biosynthesis and an important second messenger mediating plant responses to environmental stresses. The objectives were to determine how PA priming could regulate plant tolerance to heat stress and to identify specific lipids and molecular species associated with PA-enhanced heat tolerance in tall fescue (Festuca arundinacea). Plants were exposed to heat stress (38/33 °C) for 14 days with or without foliar application of PA (25 µM). PA-priming increased leaf dry weight, photochemical efficiency, and chlorophyll content and decreased leaf electrolyte leakage. Lipidomic analysis showed that the contents of PA, phosphatidylcholine (PC) and phosphatidylglycerol (PG), and digalactosyl diacylglycerol (DGDG), as well as acyl-chain length of phospholipids decreased under heat stress. The contents of PA-priming regulated major lipid molecular species, including PA (36:6), PG (32:1), PI (34:3), PI (36:5), PI (36:4), PI (36:3), PS (36:2), MGDG (34:4), MGDG (34:3), DGDG (34:3), were positively correlated to physiological traits of heat tolerance, which could serve as potential lipid biomarkers for improving heat tolerance in plants. The enhanced heat tolerance by PA-priming could be mainly attributed to lipid reprogramming with enriched lipids playing roles in stress signaling, and membrane stability and integrity.
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Desiccation tolerance has evolved repeatedly in plants as an adaptation to survive extreme environments. Plants use similar biophysical and cellular mechanisms to survive life without water, but convergence at the molecular, gene and regulatory levels remains to be tested. Here we explore the evolutionary mechanisms underlying the recurrent evolution of desiccation tolerance across grasses. We observed substantial convergence in gene duplication and expression patterns associated with desiccation. Syntenic genes of shared origin are activated across species, indicative of parallel evolution. In other cases, similar metabolic pathways are induced but using different gene sets, pointing towards phenotypic convergence. Species-specific mechanisms supplement these shared core mechanisms, underlining the complexity and diversity of evolutionary adaptations to drought. Our findings provide insight into the evolutionary processes driving desiccation tolerance and highlight the roles of parallel and convergent evolution in response to environmental challenges.