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In subfamily Salsoloideae (family Chenopodiaceae) most species are C4 plants having terete leaves with Salsoloid Kranz anatomy characterized by a continuous dual chlorenchyma layer of Kranz cells (KCs) and mesophyll (M) cells, surrounding water storage and vascular tissue. From section Coccosalsola sensu Botschantzev, leaf structural and photosynthetic features were analysed on selected species of Salsola which are not performing C4 based on leaf carbon isotope composition. The results infer the following progression in distinct functional and structural forms from C3 to intermediate to C4 photosynthesis with increased leaf succulence without changes in vein density: From species performing C3 photosynthesis with Sympegmoid anatomy with two equivalent layers of elongated M cells, with few organelles in a discontinuous layer of bundle sheath (BS) cells (S. genistoides, S. masenderanica, S. webbii) > development of proto-Kranz BS cells having mitochondria in a centripetal position and increased chloroplast number (S. montana) > functional C3–C4 intermediates having intermediate CO2 compensation points with refixation of photorespired CO2, development of Kranz-like anatomy with reduction in the outer M cell layer to hypodermal-like cells, and increased specialization (but not size) of a Kranz-like inner layer of cells with increased cell wall thickness, organelle number, and selective expression of mitochondrial glycine decarboxylase (Kranz-like Sympegmoid, S. arbusculiformis; and Kranz-like Salsoloid, S. divaricata) > selective expression of enzymes between the two cell types for performing C4 with Salsoloid-type anatomy. Phylogenetic analysis of tribe Salsoleae shows the occurrence of C3 and intermediates in several clades, and lineages of interest for studying different forms of anatomy.

Salsola plants are halophytic crops that are distributed worldwide, with more than 100 species figured out in Asia, the Mediterranean region and North Africa. Different Salsola species were reported to exert marked anti-inflammatory activities, whereas the potential anti-inflammatory activities of the three species, S. tetrandra , S. tetragona and S. vermiculata , have not been evaluated. This study provides a comprehensive metabolic study of the shoots and roots of those three species to identify potential anti-inflammatory candidates. An ultra-high performance liquid chromatography mass-mass spectrometry (UHPLC MS/MS) method in conjunction with multivariate analysis principles was utilized in an attempt to decipher their bio-active metabolites and their relevant anti-inflammatory activities. Eighty metabolites were identified in the tested extracts, where nitrogenous compounds and phenolics were highly detected in S. tetragona samples, meanwhile, saponins and phenolic acids were highly dominant in S. tetrendra sample and S. vermiculata samples have a similar chemical profile as S. tetrandra . Concerning the anti-inflammatory activity of the tested extracts, the safety margin of all the tested extracts was higher than that of the standard drug piroxicam. The shoots of the three species demonstrated more potent anti-inflammatory activities compared to the roots. The shoot extract of S. tetrandra was the most biologically active fraction. The obtained results revealed the shoots of the three Salosla species to be promising anti-inflammatory drug candidates of high safety and efficacy that could be used in the pharmaceutical industry.

Some species of Salsoleae (Chenopodiaceae) convert from C₃ photosynthesis during the seedling stage to the C₄ pathway in adult leaves. This unique developmental transition of photosynthetic pathways offers the exceptional opportunity to follow the development of the derived C₄ syndrome from the C₃ condition within individual plants, avoiding phylogenetic noise. Here we investigate Salsola soda, a little-studied species from tribe Salsoleae, using an ontogenetic approach. Anatomical sections, carbon isotope (δ13C) values, transcriptome analysis by means of mRNA sequencing, and protein levels of the key C₄ enzyme phosphoenolpyruvate carboxylase (PEPC) were examined from seed to adult plant stages. Despite a previous report, our results based on δ13C values, anatomy and transcriptomics clearly indicate a C₃ phase during the cotyledon stage. During this stage, the entire transcriptional repertoire of the C₄ NADP-malic enzyme type is detected at low levels compared to a significant increase in true leaves. In contrast, abundance of transcripts encoding most of the major photorespiratory enzymes is not significantly decreased in leaves compared to cotyledons. PEPC polypeptide was detected only in leaves, correlating with increased PEPC transcript abundance from the cotyledon to leaf stage.

Salsola collina Pall has a long history of being used as a traditional medicine to treat hypertension, headache, insomnia, constipation and vertigo. However, only a few biologically active substances have been identified from S. collina. Here, the shoots and roots of S. collina, namely L-Sc and R-Sc, were studied. The primary and secondary metabolites were investigated using ultrahigh-performance liquid chromatography-electrospray ionization-tandem mass spectrometry (UPLC-ESI-MS/MS). A total of 637 putative metabolites were identified and these metabolites were mainly classified into ten different categories. Correlation analysis, hierarchical clustering analysis, principal component analysis and orthogonal partial least squares discriminant analysis of metabolites showed that the L-Sc samples could be clearly separated from the R-Sc samples. Differential accumulated metabolite analysis revealed that most of differential primary metabolites were significantly lower in the L-Sc than in the R-Sc. Conversely, the major differential secondary metabolites had higher levels in the L-Sc than in the R-Sc. Further analysis indicated that the flavonoids were the major putative antioxidant components and most of putative antioxidant components exhibited higher relative concentrations in the L-Sc than the R-Sc. These results improve our understanding of metabolite accumulation and provide a reference for the study of medicinal value in S. collina.

We investigated the possible use of dew as a water source for three desert plant species native to the Negev Desert: an annual Salsola inermis, and two perennials Artemisia sieberi and Haloxylon scoparium, with different rooting depths of 15, 30 and 90 cm, respectively. We quantified dew-water inputs and used stable isotope analyses to determine the proportion of dew as compared to the proportion of soil water each species utilized. Dew was isotopically enriched (δD values ranged from -25 to 5 ‰), relative to rainfall with δD values that ranged from — 40 to — 20 ‰ and relative to soil water with δD values that ranged from — 65 to — 35 ‰. Using a two-source isotope mixing model, we found that S. inermis, A. sieberi and H. scoparium used, on average, 56, 63 and 46 % of their water from dewfall, respectively. Our results suggest that dew-water utilization by Negev Desert plants is highly significant ecologically and thus may be more common than previously thought. In light of future predicted climate change, it may be increasingly important for plants of the Negev Desert to make use of dew as a water resource as it may play an important role in their ability to cope with the associated hydrological constraints predicted for the Negev region.

The genus Salsola L. (Russian thistle, Saltwort) includes halophyte plants and is considered one of the largest genera in the family Amaranthaceae. The genus involves annual semi-dwarf to dwarf shrubs and woody tree. The genus Salsola is frequently overlooked, and few people are aware of its significance. The majority of studies focus on pollen morphology and species identification. Salsola has had little research on its phytochemical makeup or biological effects. Therefore, we present this review to cover all aspects of genus Salsola, including taxonomy, distribution, differences in the chemical constituents and representative examples of isolated compounds produced by various species of genus Salsola and in relation to their several reported biological activities for use in folk medicine worldwide.

Salsola is an important genus in the plant kingdom with diverse traditional, industrial, and environmental applications. Salsola species are widely distributed in temperate regions and represent about 45% of desert plants. They are a rich source of diverse phytochemical classes, such as alkaloids, cardenolides, triterpenoids, coumarins, flavonoids, isoflavonoids, and phenolic acids. Salsola spp. were traditionally used as antihypertensive, anti-inflammatory, and immunostimulants. They attracted great interest from researchers as several pharmacological activities were reported, including analgesic, antipyretic, antioxidant, cytotoxic, hepatoprotective, contraceptive, antidiabetic, neuroprotective, and antimicrobial activities. Genus Salsola is one of the most notorious plant genera from the taxonomical point of view. Our study represents a comprehensive review of the previous phytochemical and biological research on the old world Salsola secies. It is designed to be a guide for future research on different plant species that still belong to this genus or have been transferred to other genera.

Salsola ferganica is a natural desert herbaceous plant in the arid area of western and northwestern China. Because of its salt tolerance and drought resistance, it is of great significance in desert afforestation and sand-fixing capacity. There has been much research on the genes involved in plants under desert stresses in recent years. The application of the best internal reference genes for standardization was a critical procedure in analyzing the gene expression under different types. Even so, the reference gene has not been reported in the application of gene expression normalization of S. ferganica. In this study, nine reference genes (TUA-1726, TUA-1760, TUB, GAPDH, ACT, 50S, HSC70, APT, and U-box) in S. ferganica were adopted and analyzed under six different treatments (ABA, heat, cold, NaCl, methyl viologen (MV), and PEG). The applicability of candidate genes was evaluated by statistical software, including geNorm, NormFinder, BestKeeper, and RefFinder, based on their stability values in all the treatments. These results indicated that the simultaneous selection of two stable reference genes would fully standardize the optimization of the normalization research. To verify the feasibility of the above internal reference genes, the CT values of AP2/ERF transcription factor family genes were standardized using the most (ACT) and least (GAPDH) stable reference genes in S. ferganica seedlings under six abiotic stresses. The research showed that HSC70 and U-box were the most appropriate reference genes in ABA stressed samples, and ACT and U-box genes were the optimal references for heat-stressed samples. TUA-1726 and U-box showed the smallest value in gene expression levels of cold treatment. The internal reference groups of the best applicability for the other samples were U-box and ACT under NaCl treatment, ACT and TUA-1726 under MV stress, HSC70 and TUB under PEG treatment, and ACT in all samples. ACT and U-box showed higher stability than the other genes based on the comprehensive stability ranking of RefFinder, as determined by the geometric mean in this study. These results will contribute to later gene expression studies in other closely related species and provide an important foundation for gene expression analysis in S. ferganica.

Nest-site choice profoundly influences reproductive success and the survival of incubating adult birds. Asian Houbara (Chlamydotis macqueenii) nest in subtly contrasting habitats where the main cause of nest failure is predation. We examined nest-site selection across 3 semiarid shrub habitats that differed in vegetation structure and hypothesized that increased concealment would reduce nest predation. We quantified vegetation structure at 210 nests and 194 random control sites at 2 scales, the “nest area” (50 m radius, considering mean “shrub height” and mean “shrub frequency”) and “nest scrape” (2 m radius, considering a “concealment index”). Variance ratio tests showed that variance in both shrub height and concealment index was lower at nests than at random sites, indicating nonrandom selection. Analysis of the probability of nest occurrence for nest area indicated consistent selection of intermediate shrub heights (shrub height + shrub height2) in the Astragalus, Salsola arbuscula, and S. rigida shrub assemblages (29.5–31.5 cm), although this was not supported statistically in S. rigida because the vegetation available was already similar to the optimal structure. Nest survival analysis, controlling for date, showed that shrub height (but not its quadratic term) in the nest area reduced nest predation rate. Females likely traded off nesting in even taller shrubs that may confer greater nest success against the ability to see approaching danger and thus to reduce the risk of being depredated themselves (head height during vigilance when incubating is ∼30 cm), given that we have no records of females being depredated on the nest. At the nest scrape, females strongly selected better-concealed locations, although the concealment index did not affect nest success. We suggest that concealing the scrape among shrubs may have other roles, such as thermoregulation.

C 4 plants had evolved from C 3 ancestors in at least more than 60 independent lineages of angiosperms, most probably in response to decreasing levels of atmospheric CO 2 and environmental conditions promoting photorespiration. Salsola , Petrosimonia , and Cyathobasis genera of Salsoloideae subfamily contain some species with C 3 cotyledons followed by C 4 leaves. The aim of this study was to determine and compare the biochemical and structural differences between C 3 cotyledons and subsequent C 4 leaves in these genera. The results showed that there were isopalisade C 3 cotyledons in Salsola grandis and isolateral C 3 cotyledons in Cyathobasis fruticulosa , while salsoloid type C 4 Kranz anatomy was present in subsequent leaves. Petrosimonia nigdeensis had dorsiventral C 3 cotyledons and salsoloid-type C 4 leaves. The proteins of C 4 isoform phosphoenolpyruvate carboxylase (PEPC) and pyruvate orthophosphate dikinase (PPDK) enzymes were absent or very sparse in C 3 cotyledons of these species, whereas they were abundant in their C 4 leaves. The proteins of photorespiratory isoform glycolate oxidase (GOX) and glycine decarboxylase-H subunit (GDC-H) were generally higher in cotyledons than leaves. The protein level of photosynthetic isoform ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) enzyme was lower in C 4 leaves compared to C 3 cotyledons. Transcript levels of these enzymes were generally consistent with their protein content except for GOX in S. grandis and S. tragus and glycine decarboxylase complex (GDC) in S. tragus . As a result, we demonstrated that not only the protein and transcript levels of the enzymes required for C 4 pathway increased, but also those levels of C 3 and photorespiratory enzymes decreased during the transition from C 3 cotyledons to C 4 leaves. These results are considered significant in terms of shedding light on the understanding of evolutionary transition from C 3 to C 4 biochemical pathway in a single plant and contributing to C 4 engineering.