Search

Limonium bicolor , known horticulturally as sea lavender, is a typical recretohalophyte with salt glands in its leaf epidermis that secrete excess Na+ out of the plant. Although many genes have been proposed to contribute to salt gland initiation and development, a detailed analysis of alternative splicing, alternative polyadenylation patterns, and long non-coding RNAs (lncRNAs) has been lacking. Here, we applied single-molecule long-read mRNA isoform sequencing (Iso-seq) to explore the complexity of the L. bicolor transcriptome in leaves during salt gland initiation (stage A) and salt gland differentiation (stage B) based on the reference genome. We identified alternative splicing events and the use of alternative poly(A) sites unique to stage A or stage B, leading to the hypothesis that they might contribute to the differentiation of salt glands. Based on the Iso-seq data and RNA in situ hybridization of candidate genes, we selected the lncRNA Btranscript_153392 for gene editing and virus-induced gene silencing to dissect its function. In the absence of this transcript, we observed fewer salt glands on the leaf epidermis, leading to diminished salt secretion and salt tolerance. Our data provide transcriptome resources for unraveling the mechanisms behind salt gland development and furthering crop transformation efforts towards enhanced survivability in saline soils. [ABSTRACT FROM AUTHOR]

Salt bladders and salt glands of recretohalophytes are specific salt-secreting structures evolving from trichomes that allow plants to adapt to extreme environmental conditions (such as salinity or drought) by precise regulation of the salt load in metabolically active leaf tissues. The knowledge of the process by which these structures are formed and operate may provide a blueprint for introducing them into cultivated crops thus creating climate-resilient genotypes. This review summarizes the recent progress in understanding mechanisms involved in the development and operation of salt bladders and glands, at physiological, molecular, and genetic levels. We show that these processes differ depending on the anatomical structure of glands (unicellular vs bi-cellular vs multicellular), as well as between salt glands and bladders. We also show that recretohalophytes with different types of salt glands may have different trichome development. We discuss new evidence for the role of vesicular transport in salt secretion and highlight the importance of multi-omics in understanding mechanisms of salt secretion and sequestration in epidermal bladder cells, and their contribution to salinity stress tolerance in plants. [ABSTRACT FROM AUTHOR]

Under salt stress, recretohalophyte Plumbago auriculata tetraploids enhance salt tolerance by increasing selective secretion of Na+ compared with that in diploids, although the mechanism is unclear. Using non-invasive micro-test technology, the effect of salt gland Ca2+ content on Na+ and K+ secretion were investigated in diploid and tetraploid P. auriculata under salt stress. Salt gland Ca2+ content and secretion rates of Na+ and K+ were higher in tetraploids than in diploids under salt stress. Addition of exogenous Ca2+ increased the Ca2+ content of the salt gland in diploids and is accompanied by an increase in the rate of Na+ and K+ secretion. With addition of a Ca2+ channel inhibitor, diploid salt glands retained large amounts of Ca2+, leading to higher Ca2+ content and Na+ secretion rate than those of tetraploids. Inhibiting H2O2 generation and H+-ATPase activity altered Na+ and K+ secretion rates in diploids and tetraploids under salt stress, indicating involvement in regulating Na+ and K+ secretion. Our results indicate that the increased Na+ secretion rate of salt gland in tetraploids under salt stress was associated with elevated Ca2+ content in salt gland. [ABSTRACT FROM AUTHOR]

SUMMARY: Salt glands are the unique epidermal structures present in recretohalophytes, plants that actively excrete excess Na+ by salt secretory structures to avoid salt damage. Here, we describe a transmembrane protein that localizes to the plasma membrane of the recretohalophyte Limonium bicolor. As virus‐induced gene silencing of the corresponding gene LbRSG in L. bicolor decreased the number of salt glands, we named the gene Reduced Salt Gland. We detected LbRSG transcripts in salt glands by in situ hybridization and transient transformation. Overexpression and silencing of LbRSG in L. bicolor pointed to a positive role in salt gland development and salt secretion by interacting with Lb3G16832. Heterologous LbRSG expression in Arabidopsis enhanced salt tolerance during germination and the seedling stage by alleviating NaCl‐induced ion stress and osmotic stress after replacing or deleting the (highly) negatively charged region of extramembranous loop. After screened by immunoprecipitation‐mass spectrometry and verified using yeast two‐hybrid, PGK1 and BGLU18 were proposed to interact with LbRSG to strengthen salt tolerance. Therefore, we identified (highly) negatively charged regions in the extramembrane loop that may play an essential role in salt tolerance, offering hints about LbRSG function and its potential to confer salt resistance. Significance Statement: LbRSG (Reduced Salt Gland) was characterized as a transmembrane protein participating in salt gland development and salt resistance of recretohalophyte Limonium bicolor by interacting with Lb3G16832, and the negatively charged regions in the extramembranous loop were verified to play an essential role in salt tolerance of transgenic Arabidopsis. This study facilitated research on the molecular mechanisms of salt gland development and the design of salt‐tolerant crops for agricultural production in saline soils. [ABSTRACT FROM AUTHOR]

Key message: Exogenous 6-BA can increase endogenous hormone content, improve photosynthesis, decrease Na+ by increasing leaf salt gland density and salt secretion ability, and reduce ROS content so that it can promoteL. bicolor growth. 6-benzyl adenine (6-BA) is an artificial cytokinin and has been widely applied to improving plant adaptation to stress. However, it is rarely reported that 6-BA alleviates salt damage of halophytes. In this paper, we treated Limonium bicolor seedlings, a recretohalophyte with high medicinal and ornamental values, with 300 mM NaCl and different concentrations of 6-BA (0.5, 1.0, and 1.5 mg/L) and measured plant growth, physiological index, the density of salt gland, and the salt secretion ability of leaves. The results showed that exogenous applications 1.0 mg/L 6-BA significantly improved plant growth and photosynthesis, increased cytokinin and auxins contents, K+ and organic soluble matter contents, the activities of SOD, CAT, APX, and POD, and decreased Na+, H2O2, and O2− contents compared to that treated with 300 mM NaCl. Further research showed that exogenous 6-BA significantly increased the density of salt gland and the salt secretion ability of leaves by upregulating the expression of the salt gland developmental genes, therefore, can secrete more excess Na+, and thus reduces the Na+ concentration in leaves, which can alleviate Na+ damage to the species. In all, exogenous 1.0 mg/L 6-BA can increase endogenous hormone, improve photosynthesis, decrease Na+ by increasing secretion ability, and reduce ROS content of L. bicolor so that it can improve the growth. These results above systematically prove the new role of 6-BA in salt tolerance of L. bicolor. [ABSTRACT FROM AUTHOR]

Under salt stress, recretohalophyte Plumbago auriculata tetraploids enhance salt tolerance by increasing selective secretion of Na+ compared with that in diploids, although the mechanism is unclear. Using non-invasive micro-test technology, the effect of salt gland Ca2+ content on Na+ and K+ secretion were investigated in diploid and tetraploid P. auriculata under salt stress. Salt gland Ca2+ content and secretion rates of Na+ and K+ were higher in tetraploids than in diploids under salt stress. Addition of exogenous Ca2+ increased the Ca2+ content of the salt gland in diploids and is accompanied by an increase in the rate of Na+ and K+ secretion. With addition of a Ca2+ channel inhibitor, diploid salt glands retained large amounts of Ca2+, leading to higher Ca2+ content and Na+ secretion rate than those of tetraploids. Inhibiting H2O2 generation and H+-ATPase activity altered Na+ and K+ secretion rates in diploids and tetraploids under salt stress, indicating involvement in regulating Na+ and K+ secretion. Our results indicate that the increased Na+ secretion rate of salt gland in tetraploids under salt stress was associated with elevated Ca2+ content in salt gland.

Hyperplasia and hypertrophy, or their counterparts hypoplasia and hypotrophy, are elements of the adjustment of organ size and function in animals according to their needs under altered environmental conditions. As such processes are costly in terms of energy and biomaterials, it is assumed that they are beneficial for the survival of the individual. The ability of animals to perform such adjustments and the limitations in the scope of the adjustments are considered to be adaptive genetic traits which enable individual animals to survive regularly occurring changes in the environmental conditions in their habitats as long as such changes stay within critical limits. The restructuring of mono-functional glands in ducklings, which serve the animals in getting rid of excess amounts of ingested salt from the body, is presented as an example of complex plastic changes in organ structure. Phenotypic adjustments in these salt glands encompass both reversible processes, when environmental conditions switch back to the original state (‘phenotypic elasticity’), and irreversible ones (‘phenotypic plasticity’ in the narrow sense). As more information on genomes or transcriptomes of non-model animal species becomes available, we will better understand the biological significance of such phenotypic adjustments in animals in their natural environments and the underlying molecular mechanisms.

Salt gland is one of the important epidermal structures of recreto-halophytes in resistance to salt stress, which secrete excessive salt ions out of the body to prevent the plant from being damaged by salt stress. As an important structure to achieve high salt resistance in halophytes, salt glands deserve extensive attention and discussion in the fields of stress physiology, development and evolutionary biology. A large number of studies have been reported in the ultrastructure, physiological function, mechanism of salt secretion and developmental pattern of salt glands. In this paper, we reviewed the research progress of salt gland structure, secretion mechanism and salt gland development, summarized the feasible pathways of salt gland secretion and the regulation mode and key genes of salt gland development, proposed relevant views for future research on salt gland secretion and development, and discussed the role of this unique morphological structure of salt gland on plant salt tolerance. We also proposed the theoretical basis and suggestions for improving plant salt tolerance and breeding salt-tolerant plant varieties. The related studies will be beneficial for in-depth analysis of plant salt tolerance adaptation evolution, cultivation of salt resistant crops, and efficient utilization of saline land. [ABSTRACT FROM AUTHOR]

Land salinization considerably limits crop production. Biological improvement of saline and alkaline land is an important way to achieve efficient land use. It is crucial to study the salt tolerance of halophyte resources in order to explore and improve plant resources through biological improvement. Glaux maritima is a mesophyte halophyte with strong salt tolerance. In this study, we conducted research on the salt tolerance mechanism of G. maritima through phenotypic, physiological, and transcriptomic aspects. The results indicate that leaf crosssections revealed that G. maritima has a salt gland tissue composed of stalk, collecting, and secretory cells, which are trapped in epidermal cells. At the physiological level, the maximum salt tolerance threshold of G. maritima leaves was 600 mM/L. At this concentration, proline content, relative conductivity, and superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT) enzyme activities were maximum. At the transcriptional level, transcriptome data of three experimental groups (N0: 0 mM/L, N3: 600 mM/L, and N4: 800 mM/L) were analyzed, and six essential genes related to proline synthesis and five essential genes related to SOD and CAT enzyme activities were identified. Two genes involved in CAT enzyme activity were also found to play an important role in the MAPK signaling pathway. Trend analysis revealed that the MAPK signaling regulation (37 differentially expressed genes (DEGs)), phytohormone regulation (48 DEGs), glutathione metabolism (8 DEGs), flavonoid and flavonoid biosynthesis (2DEGs), and flavonoid biosynthesis (24 DEGs) pathways played important roles in regulating the salt tolerance of G. maritima. These findings provide valuable information for further studies on the functional characteristics of G. maritima in response to abiotic stress and may contribute to salt resistance breeding of fodder crops for cultivation in saline alkali land. [ABSTRACT FROM AUTHOR]

The levels of polycyclic aromatic hydrocarbons, PAHs, were determined in the liver and salt gland of 19 loggerhead turtles (Caretta caretta Linnaeus, Cheloniidae) stranded along the coasts of the south Tyrrhenian Sea, Italy, from 2019 to 2021. The 16 EPA’s priority PAHs were determined by gas chromatography coupled with mass spectrometry (GC-MS). The average values of ΣPAHs in liver, 139 ± 55.0, were exceptionally high and up to one hundred times those of literature and appeared even more worrying in salt gland, 320 ± 97.6 ng/g, w.w. Naphthalene was the predominant contributor to PAHs richness and accounted for 90.0% and 93.7% of ΣPAHs in the two matrices, highlighting the petrogenic source of exposure. An overall higher bioaccumulation of NAP, more than two-fold, was detected in salt gland and especially in female and adults. Data of PAHs richness highlighted a potential risk of neoplastic disease development and that anthropogenic activities may seriously impair healthy state conditions of C. caretta. populations in south Tyrrhenian Sea.

Hyperplasia and hypertrophy, or their counterparts hypoplasia and hypotrophy, are elements of the adjustment of organ size and function in animals according to their needs under altered environmental conditions. As such processes are costly in terms of energy and biomaterials, it is assumed that they are beneficial for the survival of the individual. The ability of animals to perform such adjustments and the limitations in the scope of the adjustments are considered to be adaptive genetic traits which enable individual animals to survive regularly occurring changes in the environmental conditions in their habitats as long as such changes stay within critical limits. The restructuring of mono-functional glands in ducklings, which serve the animals in getting rid of excess amounts of ingested salt from the body, is presented as an example of complex plastic changes in organ structure. Phenotypic adjustments in these salt glands encompass both reversible processes, when environmental conditions switch back to the original state ('phenotypic elasticity'), and irreversible ones ('phenotypic plasticity' in the narrow sense). As more information on genomes or transcriptomes of non-model animal species becomes available, we will better understand the biological significance of such phenotypic adjustments in animals in their natural environments and the underlying molecular mechanisms. [ABSTRACT FROM AUTHOR]

Halophytes can grow and reproduce in high-salinity environments, making them an important reservoir of genes conferring salt tolerance. With the expansion of saline soils worldwide, exploring the mechanisms of salt tolerance in halophytes and improving the salt tolerance of crops have become increasingly urgent. Limonium bicolor is a halophyte with salt glands that secrete excess Na+ through leaves. Here, we identified an uncharacterized gene Lb1G04794, which showed increased expression after NaCl treatment and was high during salt gland development in L. bicolor. Overexpression of Lb1G04794 in L. bicolor showed promoted salt gland development, indicating that this gene may promote salt gland differentiation. Transgenic Arabidopsis strains overexpressing Lb1G04794 showed increased trichomes and decreased root hairs under normal conditions. Compared with wild type (WT), root growth in the transgenic lines was less inhibited by NaCl treatment. Transgenic seedlings accumulated less fresh/dry weight reductions under long-term salt treatment, accompanied by lower Na+ and malondialdehyde accumulation than WT, indicating that these transgenic lines behave better growth and undergo less cellular damage under NaCl stress. These results were consistent with the low expression levels of salt-tolerance marker genes in the transgenic lines upon salt stress. We conclude that the unknown gene Lb1G04794 positively regulated salt gland development, and promoted salt tolerance of Arabidopsis, offering a new direction for improving salt tolerance of non-halophytes and crops. [ABSTRACT FROM AUTHOR]

The basic region/leucine zipper transcription factors (bZIPs) are one of the largest transcription factor families in eukaryotes, and play a key role in growth, development, and response to abiotic stresses. Limonium bicolor is a typical recretohalophyte, which can excrete excess salts from body to outside through the salt glands located on the stems and leaves to maintain ionic balance in the body. In this study, 58 L. bicolor bZIP members (LbbZIP1 – 58) were identified, which were unevenly distributed on 8 chromosomes and divided into 11 groups. The bioinformatics analysis revealed strong colinearity between LbbZIP s and GmbZIP s of soybean (Glycine max). And promoter cis -acting elements of LbbZIPs were related to stress responses and phytohormone responses. Most LbbZIP members responded to abiotic stresses such as NaCl, PEG, and ABA. Silencing LbbZIP28 in L. bicolor increased salt glands density and salt secretion capacity of leaves, and salt tolerance by means of up-regulating the expression levels of genes involved in salt gland development and ion transport. These results provide new information for LbbZIPs and help to interpret salt gland development and salt secretion in L. bicolor. Our results will provide a reference for a deeper understanding of the molecular foundation of LbbZIPs in L.bicolor. • Limonium bicolor is a pioneer plant for improving and utilizing saline-alkali land. • we identified 58 bZIP genes in L. bicolor genome, and the bioinformatic analysis was carried out. • Silencing LbbZIP28 in L. bicolor increased salt gland density, salt secretion from leaves, and overall salt tolerance. • Our findings provided fundamental information on the structure and evolutionary relationship of LbbZIP gene family in L. bicolor. [ABSTRACT FROM AUTHOR]

Soil salinization is one of the major factors restricting crop growth and agricultural production worldwide. Recretohalophytes have developed unique epidermal structures in their aboveground tissues, such as salt glands or salt bladders, to secrete excess salt out of the plant body as a protective mechanism from ion damage. Three hypotheses were proposed to explain how salt glands secrete salts: the osmotic hypothesis, a hypothesis similar to animal fluid transport, and vesicle-mediated exocytosis. However, there is no direct evidence to show whether the salt gland–secreted liquid contains landmark proteins or peptides which would elucidate the salt secretion mechanism. In this study, we collected the secreted liquid of salt glands from Limonium bicolor, followed by extraction and identification of its constituent proteins and peptides by SDS-PAGE and mass spectrometry. We detected 214 proteins and 440 polypeptides in the salt gland-secreted droplets of plants grown under control conditions. Unexpectedly, the proportion of energy metabolism-related proteins increased significantly though only 16 proteins and 35 polypeptides in the droplets of salt-treated plants were detected. In addition, vesicle transport proteins such as the Golgi marker enzyme glycosyltransferase were present in the secreted sap of salt glands from both control and salt-treated plants. These results suggest that trans-Golgi network-mediated vesicular transport and energy production contributes to salt secretion in salt glands. [ABSTRACT FROM AUTHOR]

Salinity, one of the most detrimental abiotic stresses, reduces growth, development, biomass, and yield of economically important plant species. Dixie grass (Sporobolus virginicus) is a halophytic, perennial C4 grass that quickly adapts to the coastal saline area and is generally used worldwide as fodder for grazing animals. However, studies have hitherto not investigated the physiological and biochemical basis of salt tolerance in dixie grass. The objective of this study was to evaluate the physiological, morphological, and biochemical responses of dixie grass to different levels of salinity (0, 400, 600, and 800 mM NaCl or Na2SO4) under greenhouse conditions. Overall growth traits of dixie grass were significantly reduced, especially in plants exposed to 800 mM Na2SO4. Leaf area of the plants was reduced by 78.49% and 54.52% over the control at 800 mM Na2SO4 and 800 mM NaCl, respectively. Na+ content in the leaf tissues of Na2SO4-treated plants was enriched by 28.88 folds over the control, causing chlorophyll degradation, diminution of photon yield of PSII, and reduction in the net photosynthetic rate. Greater accumulation of free proline and total soluble sugars in plants exposed to Na2SO4 were evident, which might be responsible for the osmotic adjustment. Alternatively, Na+ crystal secretion in dixie grass reduced ion toxicity, especially in the leaf blade. The study demonstrated that dixie grass counteracts the salt-induced ionic imbalance by modulating biochemical, physiological, and morphological attributes. [ABSTRACT FROM AUTHOR]

Limonium bicolor is a dicotyledonous recretohalophyte with several multicellular salt glands on the leaves. The plant can directly secrete excess salt onto the leaf surface through the salt glands to maintain ion homeostasis under salt stress. Therefore, it is of great significance to study the functions of genes related to salt gland development and salt tolerance. In this study, an R1- type MYB transcription factor gene was screened from L. bicolor, named LbMYB48, and its expression was strongly induced by salt stress. Subcellular localization analysis showed that LbMYB48 was localized in the nucleus. LbMYB48 protein has transcriptional activation activity shown by transcriptional activation experiments. The density of salt glands in the leaves and the salt secretion capacity of LbMYB48-silenced lines were decremented, as demonstrated by the leaf disc method to detect sodium ion secretion. Furthermore, salt stress index experiments revealed that the ability of LbMYB48-silenced lines to resist salt stress was significantly reduced. LbMYB48 regulates salt gland development and salt tolerance in L. bicolor mainly by regulating the expression of epidermal cell development related genes such as LbCPC-like and LbDIS3 and salt stress-related genes (LbSOSs, LbRLKs, and LbGSTs) as demonstrated by RNA-seq analysis of LbMYB48- silenced lines. The heterologous over-expression of LbMYB48 in Arabidopsis thaliana improves salt tolerance of plants by stabilizing ion and osmotic balance and is likely to be involved in the abscisic acid signaling pathway. Therefore, LbMYB48, a transcriptional activator regulates the salt gland development of L. bicolor and salt tolerance of L. bicolor and A. thaliana. [ABSTRACT FROM AUTHOR]

Halophytes have evolved specialized strategies to cope with high salinity. The extreme halophyte sea lavender (Limonium bicolor) lacks trichomes but possesses salt glands on its epidermis that can excrete harmful ions, such as sodium, to avoid salt damage. Here, we report a high-quality, 2.92-Gb, chromosome-scale L. bicolor genome assembly based on a combination of Illumina short reads, single-molecule, real-time long reads, chromosome conformation capture (Hi-C) data, and Bionano genome maps, greatly enriching the genomic information on recretohalophytes with multicellular salt glands. Although the L. bicolor genome contains genes that show similarity to trichome fate genes from Arabidopsis thaliana , it lacks homologs of the decision fate genes GLABRA3 , ENHANCER OF GLABRA3 , GLABRA2 , TRANSPARENT TESTA GLABRA2 , and SIAMESE , providing a molecular explanation for the absence of trichomes in this species. We identified key genes (LbHLH and LbTTG1) controlling salt gland development among classical trichome homologous genes and confirmed their roles by showing that their mutations markedly disrupted salt gland initiation, salt secretion, and salt tolerance, thus offering genetic support for the long-standing hypothesis that salt glands and trichomes may share a common origin. In addition, a whole-genome duplication event occurred in the L. bicolor genome after its divergence from Tartary buckwheat and may have contributed to its adaptation to high salinity. The L. bicolor genome resource and genetic evidence reported in this study provide profound insights into plant salt tolerance mechanisms that may facilitate the engineering of salt-tolerant crops. A high-quality, 2.92-Gb, chromosome-scale genome was assembled for the halophyte Limonium bicolor , which has multicellular salt glands. Key genes for the control of salt gland development and salt secretion were identified. Our study suggests that salt glands might share a common origin with trichomes and may have evolved for adaptation to high salinity. [ABSTRACT FROM AUTHOR]

The enzyme soluble adenylyl cyclase (sAC) is directly stimulated by bicarbonate (HCO3-) to produce the signaling molecule cyclic adenosine monophosphate (cAMP). Because sAC and sAC-related enzymes are found throughout phyla from cyanobacteria to mammals and they regulate cell physiology in response to internal and external changes in pH, CO2, and HCO3-, sAC is deemed an evolutionarily conserved acid-base sensor. Previously, sAC has been reported in dogfish shark and round ray gill cells, where they sense and counteract blood alkalosis by regulating the activity of V-type H+- ATPase. Here, we report the presence of sAC protein in gill, rectal gland, cornea, intestine, white muscle, and heart of leopard shark Triakis semifasciata Co-expression of sAC with transmembrane adenylyl cyclases supports the presence of cAMP signaling microdomains. Furthermore, immunohistochemistry on tissue sections, and western blots and cAMP-activity assays on nucleus-enriched fractions demonstrate the presence of sAC protein in and around nuclei. These results suggest that sAC modulates multiple physiological processes in shark cells, including nuclear functions.

With global increases in saline soil, it has become increasingly important to decipher salt-tolerance mechanisms and identify strategies to improve salt tolerance in crops. Halophytes complete their life cycles in environments containing ≥200 mM NaCl; these remarkable plants provide a potential source of genes for improving crop salt tolerance. Recretohalophytes such as Limonium bicolor have salt glands that secrete Na+ on their leaf epidermis. Here, we identified Lb1G04202, an uncharacterized gene with no conserved domains, from L. bicolor, which was highly expressed after NaCl treatment. We confirmed its expression in the salt gland by in situ hybridization, and then heterologously expressed Lb1G04202 in Arabidopsis thaliana. The transgenic lines had a higher germination rate, greater cotyledon growth percentage, and longer roots than the wild type (WT) under NaCl treatments (50, 100 and 150 mM). At the seedling stage, the transgenic lines grew better than the WT and had lower Na+ and malonyldialdehyde accumulation, and higher K+ and proline contents. This corresponded with the high expression of the key proline biosynthesis genes AtP5CS1 and AtP5CS2 under NaCl treatment. Isotonic mannitol treatment showed that Lb1G04202 overexpression significantly relieved osmotic stress. Therefore, this novel gene provides a potential target for improving salt tolerance. [ABSTRACT FROM AUTHOR]

Tamarix nilotica is a perennial halophyte growing naturally at the Mediterranean coastal salt marshes. Vegetative parts were collected in two successive winters and summers to investigate the seasonal effects on the ecomorphological and physiological responses of T. nilotica. The results indicated that either in winter or summer seasons, T. nilotica can reduce the effect of soil salinity by excreting salts outside its body through salt glands. Summer season was characterized by low content of soil moisture (due to rare rainfall), high soil EC, high light intensity and high temperature; therefore, plant induced certain morphoanatomical changes in leaves and stem to face the previously mentioned adverse conditions. The most remarkable changes to reduce transpiration process was found by decreasing leaf area and increasing cuticle thickness and mesophyll tissue thickness. In addition, the most marked physiological changes in summer were the significant increase in total phenols, proline, free amino acids and total soluble sugars. These compounds can work as osmotic regulators and/or antioxidants. These features enhance the defensive mechanism against dehydration and permit T. nilotica to tolerate the stress conditions in salt marsh habitat. [ABSTRACT FROM AUTHOR]

Salt-resistant plants have different mechanisms to limit the deleterious effects of high salt in soil; for example, recretohalophytes secrete salt from unique structures called salt glands. Salt glands are the first differentiated epidermal structure of the recretohalophyte sea lavender (Limonium bicolor), followed by stomata and pavement cells. While salt glands and stomata develop prior to leaf expansion, it is not clear whether these steps are connected. Here, we explored the effects of the five phytohormones salicylic acid, brassinolide, methyl jasmonate, gibberellic acid, and abscisic acid on the development of the first expanded leaf of L. bicolor and its potential connection to salt gland, stomata, and pavement cell differentiation. We calculated the total number of salt glands, stomata, and pavement cells, as well as leaf area and pavement cell area, and assessed the correlations between these parameters. We detected strong and positive correlations between salt gland number and pavement cell area, between stomatal number and pavement cell area, and between salt gland number and stomatal number. We observed evidence of coupling between the development of salt glands, stomata, and pavement cells in L. bicolor , which lays the foundation for further investigation of the mechanism behind salt gland development. [ABSTRACT FROM AUTHOR]

Abstract Background Salt, a common environmental stress factor, inhibits plant growth and reduces yields. Melatonin is a pleiotropic molecule that regulates plant growth and can alleviate environmental stress in plants. All previous research on this topic has focused on the use of melatonin to improve the relatively low salt tolerance of glycophytes by promoting growth and enhancing antioxidant ability. It is unclear whether exogenous melatonin can increase the salt tolerance of halophytes, particularly recretohalophytes, by enhancing salt secretion from the salt glands. Results To examine the mechanisms of melatonin-mediated salt tolerance, we explored the effects of exogenous applications of melatonin on the secretion of salt from the salt glands of Limonium bicolor (a kind of recretohalophyte) seedlings and on the expression of associated genes. A pretreatment with 5 μM melatonin significantly improved the growth of L. bicolor seedlings under 300 mM NaCl. Furthermore, exogenous melatonin significantly increased the dry weight and endogenous melatonin content of L. bicolor. In addition, this treatment reduced the content of Na+ and Cl− in leaves, but increased the K+ content. Both the salt secretion rate of the salt glands and the expression level of genes encoding ion transporters (LbHTK1, LbSOS1, LbPMA, and LbNHX1) and vesicular transport proteins (LbVAMP721, LbVAP27, and LbVAMP12) were significantly increased by exogenous melatonin treatment. These results indicate that melatonin improves the salt tolerance of the recretohalophyte L. bicolor via the upregulation of salt secretion by the salt glands. Conclusions Our results showed that melatonin can upregulate the expression of genes encoding ion transporters and vesicle transport proteins to enhance salt secretion from the salt glands. Combining the results of the current study with previous research, we formulated a novel mechanism by which melatonin increases salt secretion in L. bicolor. Ions in mesophyll cells are transported to the salt glands through ion transporters located at the plasma membrane. After the ions enter the salt glands, they are transported to the collecting chamber adjacent to the secretory pore through vesicle transport and ions transporter and then are secreted from the secretory pore of salt glands, which maintain ionic homeostasis in the cells and alleviate NaCl-induced growth inhibition.

The salt gland is a well-developed osmoregulation organ in marine birds, and its relative size often reflects an individual’s feeding environment and osmoregulation capability. The development and functions of salt glands have been described for the Adélie penguin (Pygoscelis adeliae), but this information has been poorly documented in the other two pygoscelid species: gentoo (P. papua) and chinstrap penguins (P. antarcticus). To describe the growth-related changes in salt gland masses in relation to chick growth, we measured the wet mass of the salt glands collected from dead gentoo and chinstrap chicks during the early breeding period. The mass of the salt glands was linearly proportional to their body measurements, especially to body mass, in both species, and no significant difference was detected between the two species. Penguins are obligate marine dwellers throughout their life cycle, and the development of the salt gland in penguin chicks suggests that their ability to regulate dietary osmotic stress begins at an early stage of development after hatching. Furthermore, the linear relationship between the gland mass and body mass also suggests that the osmoregulation capability may continue to develop as penguin chicks grow. This descriptive note provides novel and quantitative information on the early developmental pattern of salt glands in gentoo and chinstrap penguins.

Salt-resistant plants have different mechanisms to limit the deleterious effects of high salt in soil; for example, recretohalophytes secrete salt from unique structures called salt glands. Salt glands are the first differentiated epidermal structure of the recretohalophyte sea lavender (Limonium bicolor), followed by stomata and pavement cells. While salt glands and stomata develop prior to leaf expansion, it is not clear whether these steps are connected. Here, we explored the effects of the five phytohormones salicylic acid, brassinolide, methyl jasmonate, gibberellic acid, and abscisic acid on the development of the first expanded leaf of L. bicolor and its potential connection to salt gland, stomata, and pavement cell differentiation. We calculated the total number of salt glands, stomata, and pavement cells, as well as leaf area and pavement cell area, and assessed the correlations between these parameters. We detected strong and positive correlations between salt gland number and pavement cell area, between stomatal number and pavement cell area, and between salt gland number and stomatal number. We observed evidence of coupling between the development of salt glands, stomata, and pavement cells in L. bicolor, which lays the foundation for further investigation of the mechanism behind salt gland development.

Limonium Mill. plants are typical recretohalophytes, as they withstand salt stress by secreting excess salt onto the leaf surface through salt glands. However, little is known on the salinity thresholds of these plants and the function of salt glands in salt tolerance.Here, we investigated the salinity thresholds of salt tolerance of the Limonium species L. aureum (Linn.) Hill, L. gmelinii (Willd.) Kuntze, L. otolepis (Schrenk) Kuntze and L. sinuatum (L.) Mill grown with various concentrations of NaCl.The salinity thresholds of L. otolepis, L. aureum, L. sinuatum and L. gmelinii were 300, 350, 400 and 420 mm NaCl, respectively. Correlation analysis indicated that total dry weight, chlorophyll content and intercellular CO2 concentration were highly positively correlated with the total fresh weights of all four Limonium species and could therefore be used as indicators of plant salt tolerance. Furthermore, as the salt gland density on the leaf surface increased, the rate of salt secretion per salt gland also increased, allowing more Na+ to be secreted from the plant. Redundancy discriminant analysis indicated that salt gland density, Na+ content and Na+ secretion rate per salt gland were positively correlated with salt concentration.These observations support the notion that salt glands play important roles in the adaptation of Limonium species to high salinity conditions. [ABSTRACT FROM AUTHOR]

Plant epidermal cells, such as trichomes, root hairs, salt glands, and stomata, play pivotal roles in the growth, development, and environmental adaptation of terrestrial plants. Cell fate determination, differentiation, and the formation of epidermal structures represent basic developmental processes in multicellular organisms. Increasing evidence indicates that C2H2 zinc finger proteins play important roles in regulating the development of epidermal structures in plants and plant adaptation to unfavorable environments. Here, we systematically summarize the molecular mechanism underlying the roles of C2H2 zinc finger proteins in controlling epidermal cell formation in plants, with an emphasis on trichomes, root hairs, and salt glands and their roles in plant adaptation to environmental stress. In addition, we discuss the possible roles of homologous C2H2 zinc finger proteins in trichome development in non-halophytes and salt gland development in halophytes based on bioinformatic analysis. This review provides a foundation for further study of epidermal cell development and abiotic stress responses in plants. [ABSTRACT FROM AUTHOR]

Plant epidermal cells, such as trichomes, root hairs, salt glands, and stomata, play pivotal roles in the growth, development, and environmental adaptation of terrestrial plants. Cell fate determination, differentiation, and the formation of epidermal structures represent basic developmental processes in multicellular organisms. Increasing evidence indicates that C2H2 zinc finger proteins play important roles in regulating the development of epidermal structures in plants and plant adaptation to unfavorable environments. Here, we systematically summarize the molecular mechanism underlying the roles of C2H2 zinc finger proteins in controlling epidermal cell formation in plants, with an emphasis on trichomes, root hairs, and salt glands and their roles in plant adaptation to environmental stress. In addition, we discuss the possible roles of homologous C2H2 zinc finger proteins in trichome development in non-halophytes and salt gland development in halophytes based on bioinformatic analysis. This review provides a foundation for further study of epidermal cell development and abiotic stress responses in plants.

This study aimed to employ plant print identification to screen a wild-type soybean (Glycine soja Seib. and Zucc.) and then hybridize it with a cultivated soybean (Glycine max (L.) Merr.) (Lu, Chin Sci Bull 43:2074–2078, 1998) with a view to produce a salt-resistant variety. Scanning electron microscopy, optical microscopy, and paraffin sectioning were employed to compare two varieties of wild soybeans (#028 and #029) captured from different eco-environments, and the one with a salt gland present –– wild soybean #029 – was chosen as the male parent for hybridization. Considering a soybean variety with low stem as female parent, hybridization was performed and a new variety of salt-resistant soybean resulted, Ji-Yu59, which opened up a new, cost-efficient way of soybean breeding. Under saline-alkali stress, our salt-tolerant wild soybean is anticipated to exhibit antagonism by evolving a series of salt-resistant structures, including a salt-containing vacuolar package and a salt gland. The salt gland would help eliminate salts from the soybean plant; upon maturity, a salt gland is known to be able to break up salts to release their respective ions, effectively reducing salt and alkali stress. Therefore, a wild soybean with salt gland is an excellent choice for the hybridization of salt-resistant soybean varieties. [ABSTRACT FROM AUTHOR]

The rectal gland (RG) is a specialized organ that performs a major role in salt excretion of marine elasmobranch species. We aimed to describe the rectal gland ecomorphology in three stingray species (Hypanus marianae , Hypanus guttatus , and Aetobatus narinari) from benthic and pelagic habitats. The gland is lobulated and has two distinct forms: small compact S-shaped for H. marianae (mean width: RG length ± SD = 1.06 ± 0.23 cm), elongated S-shaped for H. guttatus (2.15 ± 0.06 cm), and pod-shaped for A. narinari (3.14 ± 0.57 cm). Rectal gland lengths were different between species (Kruskal–Wallis; X2 = 39.27; df = 2; P < 0.05). The capsule's dense connective tissue was three times thicker in A. narinari than in the Hypanus species (K–W; df = 2, p = 0.014). The secretory parenchyma of H. marianae was simpler, differing from other species that showed a well-defined lobule arrangement around the central duct. Indeed, the central duct of A. narinari and H. guttatus exhibited branches that invaginated into lobes and originated the lobular ducts, while the central duct lacks branches and opened directly into the secretory tubule in H. marianae. Furthermore, central duct cells in the stratified epithelium of A. narinari showed a higher number (average 7.4 ± 1.51/cells) and cell layer thickness (0.09 mm) than the other species analyzed (ANOVA; F = 6.90, p = 0.01). The morphological differences between the rectal gland of Myliobatiformes analyzed showed that their phylogenetic history could be a better explanation to the changes between basal (H. marianae) and derived (H. guttatus and A. narinari) species. [ABSTRACT FROM AUTHOR]

• Microhairs of Zoysia japonica were observed by electron microscopies. • Adaxial microhairs excreted soluble salts. • Abaxial microhairs excreted insoluble substances. • Excretion differed, but microhair structures were similar between leaf sides. Bicellular trichomes called 'microhairs' are known as 'salt glands' in some Chloridoideae in the Poaceae. The adaxial leaf surface of Zoysia japonica, a salt-tolerant grass, excretes salts, whereas the abaxial leaf surface does not. In this study, we investigated the morphology and excreting-function of the microhairs of Z. japonica , comparing the adaxial and abaxial leaf surfaces. Scanning electron microscopy of the leaf surfaces showed the distribution of microhairs on both sides and revealed significant excretion on the adaxial side but a small excretion on the abaxial side. Energy-dispersive X-ray spectrometry revealed that microhairs on the adaxial side excrete sodium, magnesium, and chlorine, whereas those on the abaxial side excrete or secrete insoluble substances, which sometimes contain silicon. Transmission electron microscopy was used to clarify the ultrastructure of microhairs, as in the first detailed report on Zoysia spp.; however, the microhairs on both leaf sides showed typical structures of bicellular salt glands reported in Chloridoideae. The cap cells of the microhairs had smooth surfaces without epicuticular wax structures. These results indicate that the ultrastructural features of Z. japonica microhairs are important for understanding physiological mechanisms in common with salt-excretion and the excretion or secretion of other substances. [ABSTRACT FROM AUTHOR]

Halophytes can survive and complete their life cycle in the presence of ≥200 mM NaCl. These remarkable plants have developed various strategies to tolerate salinity and thrive in high-salt environments. At the appropriate levels, salt has a beneficial effect on the vegetative growth of halophytes but inhibits the growth of non-halophytes. In recent years, many studies have focused on elucidating the salt-tolerance mechanisms of halophytes at the molecular, physiological, and individual level. In this review, we focus on the mechanisms, from the macroscopic to the molecular, underlying the successful growth of halophytes in saline environments to explain why salt has beneficial effects on halophytes but harmful effects on non-halophytes. These mechanisms include the specialized organs of halophytes (for example, ion compartmentalization in succulent leaves), their unique structures (salt glands and hydrophobic barriers in roots), and their salt-tolerance genes. We hope to shed light on the use of halophytes for engineering salt-tolerant crops, soil conservation, and the protection of freshwater resources in the near future.

Using diffusible iodine-based contrast-enhanced computed tomography (diceCT), we examined the morphology of the oral glands of 12 species of the family Homalopsidae. Snakes of this family exhibit substantial interspecific morphological variation in their oral glands. Particular variables are the venom glands, ranging from large (e.g., Subsessor bocourti) to small (e.g., Erpeton tentaculatum). The supra- and infralabial glands are more uniform in morphology, being the second most developed in almost all the sampled species. Premaxillary glands distinct from the supralabial glands were observed in five species (Myron richardsonii, Bitia hydroides, Cantoria violacea, Fordonia leucobalia, and Gerarda prevostiana), in addition to Cerberus rynchops, the only species in which this condition was previously documented associated with the excretion of salt. In the three species of the saltwater group of homalopsids (C. violacea, F. leucobalia, and G. prevostiana), the premaxillary glands also extend posteriorly, occupying a large area above the supralabial gland, a condition not observed in any other species of snake studied thus far. Character evolution analyses indicate that premaxillary glands differentiated from the supralabial gland and evolved independently three or four times in the family, always in lineages that invaded marine habitats. Our results suggest that the differentiated premaxillary glands are likely salt glands, as is the case in C. rynchops. If corroborated, this increases to six or seven the number of independent evolutionary origins of salt glands in snakes that have undergone an evolutionary transition to marine life., (© 2024 Anatomical Society.)

Under salt stress, recretohalophyte Plumbago auriculata tetraploids enhance salt tolerance by increasing selective secretion of Na + compared with that in diploids, although the mechanism is unclear. Using non-invasive micro-test technology, the effect of salt gland Ca 2+ content on Na + and K + secretion were investigated in diploid and tetraploid P. auriculata under salt stress. Salt gland Ca 2+ content and secretion rates of Na + and K + were higher in tetraploids than in diploids under salt stress. Addition of exogenous Ca 2+ increased the Ca 2+ content of the salt gland in diploids and is accompanied by an increase in the rate of Na + and K + secretion. With addition of a Ca 2+ channel inhibitor, diploid salt glands retained large amounts of Ca 2+ , leading to higher Ca 2+ content and Na + secretion rate than those of tetraploids. Inhibiting H 2 O 2 generation and H + -ATPase activity altered Na + and K + secretion rates in diploids and tetraploids under salt stress, indicating involvement in regulating Na + and K + secretion. Our results indicate that the increased Na + secretion rate of salt gland in tetraploids under salt stress was associated with elevated Ca 2+ content in salt gland., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Duan, Jiang, Lei, Ouyang, Liu, Zhao, Li, Yang, Li, Yi and Gao.)

Background: Salt, a common environmental stress factor, inhibits plant growth and reduces yields. Melatonin is a pleiotropic molecule that regulates plant growth and can alleviate environmental stress in plants. All previous research on this topic has focused on the use of melatonin to improve the relatively low salt tolerance of glycophytes by promoting growth and enhancing antioxidant ability. It is unclear whether exogenous melatonin can increase the salt tolerance of halophytes, particularly recretohalophytes, by enhancing salt secretion from the salt glands. Results: To examine the mechanisms of melatonin-mediated salt tolerance, we explored the effects of exogenous applications of melatonin on the secretion of salt from the salt glands of Limonium bicolor (a kind of recretohalophyte) seedlings and on the expression of associated genes. A pretreatment with 5 μM melatonin significantly improved the growth of L. bicolor seedlings under 300 mM NaCl. Furthermore, exogenous melatonin significantly increased the dry weight and endogenous melatonin content of L. bicolor. In addition, this treatment reduced the content of Na+ and Cl− in leaves, but increased the K+ content. Both the salt secretion rate of the salt glands and the expression level of genes encoding ion transporters (LbHTK1, LbSOS1, LbPMA, and LbNHX1) and vesicular transport proteins (LbVAMP721, LbVAP27, and LbVAMP12) were significantly increased by exogenous melatonin treatment. These results indicate that melatonin improves the salt tolerance of the recretohalophyte L. bicolor via the upregulation of salt secretion by the salt glands. Conclusions: Our results showed that melatonin can upregulate the expression of genes encoding ion transporters and vesicle transport proteins to enhance salt secretion from the salt glands. Combining the results of the current study with previous research, we formulated a novel mechanism by which melatonin increases salt secretion in L. bicolor. Ions in mesophyll cells are transported to the salt glands through ion transporters located at the plasma membrane. After the ions enter the salt glands, they are transported to the collecting chamber adjacent to the secretory pore through vesicle transport and ions transporter and then are secreted from the secretory pore of salt glands, which maintain ionic homeostasis in the cells and alleviate NaCl-induced growth inhibition. [ABSTRACT FROM AUTHOR]

The salt gland is a well-developed osmoregulation organ in marine birds, and its relative size often reflects an individual’s feeding environment and osmoregulation capability. The development and functions of salt glands have been described for the Adélie penguin (Pygoscelis adeliae), but this information has been poorly documented in the other two pygoscelid species: gentoo (P. papua) and chinstrap penguins (P. antarcticus). To describe the growth-related changes in salt gland masses in relation to chick growth, we measured the wet mass of the salt glands collected from dead gentoo and chinstrap chicks during the early breeding period. The mass of the salt glands was linearly proportional to their body measurements, especially to body mass, in both species, and no significant difference was detected between the two species. Penguins are obligate marine dwellers throughout their life cycle, and the development of the salt gland in penguin chicks suggests that their ability to regulate dietary osmotic stress begins at an early stage of development after hatching. Furthermore, the linear relationship between the gland mass and body mass also suggests that the osmoregulation capability may continue to develop as penguin chicks grow. This descriptive note provides novel and quantitative information on the early developmental pattern of salt glands in gentoo and chinstrap penguins. [ABSTRACT FROM AUTHOR]

Copyright of Avian Diseases is the property of American Association of Avian Pathologists, Inc. and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Salt secretion is an important strategy used by the mangrove plant Aegiceras corniculatum to adapt to the coastal intertidal environment. However, the structural, developmental and functional analyses on the leaf salt glands, particularly the salt secretion mechanism, are not well documented. In this study, we investigated the structural, developmental and degenerative characteristics and the salt secretion mechanisms of salt glands to further elucidate the mechanisms of salt tolerance of A. corniculatum. The results showed that the salt gland cells have a large number of mitochondria and vesicles, and plenty of plasmodesmata as well, while chloroplasts were found in the collecting cells. The salt glands developed early and began to differentiate at the leaf primordium stage. We observed and defined three stages of salt gland degradation for the first time in A. corniculatum, where the secretory cells gradually twisted and wrinkled inward and collapsed downward as the salt gland degeneration increased and the intensity of salt gland autofluorescence gradually diminished. In addition, we found that the salt secretion rate of the salt glands increased when the treated concentration of NaCl increased, reaching the maximum at 400 mM NaCl. The salt-secreting capacity of the salt glands of the adaxial epidermis is significantly greater than that of the abaxial epidermis. The real-time quantitative PCR results indicate that SAD2, TTG1, GL2 and RBR1 may be involved in regulating the development of the salt glands of A. corniculatum. Moreover, Na+/H+ antiporter, H+-ATPase, K+ channel and Cl- channel may play important roles in the salt secretion of salt glands. In sum mary, this study strengthens the understanding of the structural, developmental and degenerative patterns of salt glands and salt secretion mechanisms in mangrove recretohalophyte A. corniculatum, providing an important reference for further studies at the molecular level., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Key Message: Exogenous 6-BA can increase endogenous hormone content, improve photosynthesis, decrease Na + by increasing leaf salt gland density and salt secretion ability, and reduce ROS content so that it can promote L. bicolor growth. 6-benzyl adenine (6-BA) is an artificial cytokinin and has been widely applied to improving plant adaptation to stress. However, it is rarely reported that 6-BA alleviates salt damage of halophytes. In this paper, we treated Limonium bicolor seedlings, a recretohalophyte with high medicinal and ornamental values, with 300 mM NaCl and different concentrations of 6-BA (0.5, 1.0, and 1.5 mg/L) and measured plant growth, physiological index, the density of salt gland, and the salt secretion ability of leaves. The results showed that exogenous applications 1.0 mg/L 6-BA significantly improved plant growth and photosynthesis, increased cytokinin and auxins contents, K + and organic soluble matter contents, the activities of SOD, CAT, APX, and POD, and decreased Na + , H 2 O 2 , and O 2 contents compared to that treated with 300 mM NaCl. Further research showed that exogenous 6-BA significantly increased the density of salt gland and the salt secretion ability of leaves by upregulating the expression of the salt gland developmental genes, therefore, can secrete more excess Na - contents compared to that treated with 300 mM NaCl. Further research showed that exogenous 6-BA significantly increased the density of salt gland and the salt secretion ability of leaves by upregulating the expression of the salt gland developmental genes, therefore, can secrete more excess Na + , and thus reduces the Na + concentration in leaves, which can alleviate Na + damage to the species. In all, exogenous 1.0 mg/L 6-BA can increase endogenous hormone, improve photosynthesis, decrease Na + by increasing secretion ability, and reduce ROS content of L. bicolor so that it can improve the growth. These results above systematically prove the new role of 6-BA in salt tolerance of L. bicolor., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)