Boron (B) is an essential trace element required for the physiological functioning of higher plants. B deficiency is considered as a nutritional disorder that adversely affects the metabolism and growth of plants. B is involved in the structural and functional integrity of the cell wall and membranes, ion fluxes (H+, K+, PO43−, Rb+, Ca2+) across the membranes, cell division and elongation, nitrogen and carbohydrate metabolism, sugar transport, cytoskeletal proteins, and plasmalemma-bound enzymes, nucleic acid, indoleacetic acid, polyamines, ascorbic acid, and phenol metabolism and transport. This review critically examines the functions of B in plants, deficiency symptoms, and the mechanism of B uptake and transport under limited B conditions. B deficiency can be mitigated by inorganic fertilizer supplementation, but the deleterious impact of frequent fertilizer application disrupts soil fertility and creates environmental pollution. Considering this, we have summarized the available information regarding alternative approaches, such as root structural modification, grafting, application of biostimulators (mycorrhizal fungi (MF) and rhizobacteria), and nanotechnology, that can be effectively utilized for B acquisition, leading to resource conservation. Additionally, we have discussed several new aspects, such as the combination of grafting or MF with nanotechnology, combined inoculation of arbuscular MF and rhizobacteria, melatonin application, and the use of natural and synthetic chelators, that possibly play a role in B uptake and translocation under B stress conditions.

Root respiratory burst oxidase homologue-dependent H2O2 production serves as an early signal to confer salt tolerance on a grafted cucumber through regulation of Na+ exclusion and stomatal closure.

A fungus Fusarium oxysporum F. sp. niveum (FON) is the causal organism of Fusarium wilt in watermelon. In this study, we evaluated the effect of wheat intercropping on the Fusarium wilt of watermelon. Our results showed that wheat intercropping decreases the incidence of Fusarium wilt of watermelon, likely due to the secretion of coumaric acid from the roots of wheat that dramatically inhibits FON spore germination, sporulation, and growth. The secretion of p-hydroxybenzoic acid, ferulic acid, and cinnamic acid from the roots of watermelon stimulates FON spore germination, sporulation, and growth. The secretion of phenolic acids and organic acids from the roots of watermelon is also promoted by FON infection. However, secretion of phenolic acids and organic acids from the roots of watermelon is substantially reduced under wheat intercropping systems. FON infection increases the accumulation of free and conjugated salicylic acid (SA) in watermelon grown under wheat intercropping systems through isochorismate (ICS) and phenylalanine ammonia-lyase (PAL) pathways. Furthermore, wheat intercropping up-regulates the expression of disease-and defense-responsive genes and improves the activities of corresponding pathogenesis-related (PR) enzymes in the roots of watermelon. In conclusion, the secretion of coumaric acid from the roots of wheat and changes in the composition of phenolic acid and organic acid secretion from the roots of watermelon suppress Fusarium wilt of watermelon under wheat intercropping system. Meanwhile, wheat intercropping also enhanced the resistance of watermelon to FON by up-regulating the expression of disease-and defense-responsive genes in watermelon.

Potassium (K+) is a critical determinant of salinity tolerance, and H2O2 has been recognized as an important signaling molecule that mediates many physiological responses. However, the details of how H2O2 signaling regulates K+ uptake in the root under salt stress remain elusive. In this study, salt-sensitive cucumber and salt-tolerant pumpkin which belong to the same family, Cucurbitaceae, were used to answer the above question. We show that higher salt tolerance in pumpkin was related to its superior ability for K+ uptake and higher H2O2 accumulation in the root apex. Transcriptome analysis showed that salinity induced 5816 (3005 up- and 2811 down-) and 4679 (3965 up- and 714 down-) differentially expressed genes (DEGs) in cucumber and pumpkin, respectively. DEGs encoding NADPH oxidase (respiratory burst oxidase homolog D; RBOHD), 14-3-3 protein (GRF12), plasma membrane H+-ATPase (AHA1), and potassium transporter (HAK5) showed higher expression in pumpkin than in cucumber under salinity stress. Treatment with the NADPH oxidase inhibitor diphenylene iodonium resulted in lower RBOHD, GRF12, AHA1, and HAK5 expression, reduced plasma membrane H+-ATPase activity, and lower K+ uptake, leading to a loss of the salinity tolerance trait in pumpkin. The opposite results were obtained when the plants were pre-treated with exogenous H2O2. Knocking out of RBOHD in pumpkin by CRISPR/Cas9 [clustered regularly interspaced short palindromic repeat (CRISPR)/CRISPR-associated protein 9] editing of coding sequences resulted in lower root apex H2O2 and K+ content and GRF12, AHA1, and HAK5 expression, ultimately resulting in a salt-sensitive phenotype. However, ectopic expression of pumpkin RBOHD in Arabidopsis led to the opposite effect. Taken together, this study shows that RBOHD-dependent H2O2 signaling in the root apex is important for pumpkin salt tolerance and suggests a novel mechanism that confers this trait, namely RBOHD-mediated transcriptional and post-translational activation of plasma membrane H+-ATPase operating upstream of HAK5 K+ uptake transporters.

Tissue tolerance to salinity in Cucurbita is associated with HKT1-mediated Na+ exclusion from the leaf mesophyll, NHX4/6-mediated Na+ sequestration in the leaf vein, K+ retention in the leaf mesophyll, and early ABA-induced stomatal closure.

No abstract

Drought, cold and salinity are the major environmental stresses that limit agricultural productivity. NAC transcription factors regulate the stress response in plants. Pumpkin (Cucurbita moschata) is an important cucurbit vegetable crop and it has strong resistance to abiotic stress; however, the biological functions of stress-related NAC genes in this crop are largely unknown. This study reports the function of CmNAC1, a stress-responsive pumpkin NAC domain protein. The CmNAC1-GFP fusion protein was transiently expressed in tobacco leaves for subcellular localization analysis, and we found that CmNAC1 is localized in the nucleus. Transactivation assay in yeast cells revealed that CmNAC1 functions as a transcription activator, and its transactivation domain is located in the C-terminus. CmNAC1 was ubiquitously expressed in different organs, and its transcript was induced by salinity, cold, dehydration, H2O2, and abscisic acid (ABA) treatment. Furthermore, the ectopic expression (EE) of CmNAC1 in Arabidopsis led to ABA hypersensitivity and enhanced tolerance to salinity, drought and cold stress. In addition, five ABA-responsive elements were enriched in CmNAC1 promoter. The CmNAC1-EE plants exhibited different root architecture, leaf morphology, and significantly high concentration of ABA compared with WT Arabidopsis under normal conditions. Our results indicated that CmNAC1 is a critical factor in ABA signaling pathways and it can be utilized in transgenic breeding to improve the abiotic stress tolerance of crops.

Soil salinity adversely affects the growth and yield of crops, including cucumber, one of the most important vegetables in the world. Grafting with salt-tolerant pumpkin as the rootstock effectively improves the growth of cucumber under different salt conditions by limiting Na+ transport from the pumpkin rootstock to the cucumber scion. High-affinity potassium transporters (HKTs) are crucial for the long distance transport of Na+ in plants, but the function of pumpkin HKTs in this process of grafted cucumber plants remains unclear. In this work, we have characterized CmHKT1;1 as a member of the HKT gene family in Cucurbita moschata and observed an obvious upregulation of CmHKT1;1 in roots under NaCl stress conditions. Heterologous expression analyses in yeast mutants indicated that CmHKT1;1 is a Na+-selective transporter. The transient expression in tobacco epidermal cells and in situ hybridization showed CmHKT1;1 localization at plasma membrane, and preferential expression in root stele. Moreover, ectopic expression of CmHKT1;1 in cucumber decreased the Na+ accumulation in the plants shoots. Finally, the CmHKT1;1 transgenic line as the rootstock decreased the Na+ content in the wild type shoots. These findings suggest that CmHKT1;1 plays a key role in the salt tolerance of grafted cucumber by limiting Na+ transport from the rootstock to the scion and can further be useful for engineering salt tolerance in cucurbit crops.