A systematic review on the ethnoveterinary uses of mediterranean salt-tolerant plants: Exploring its potential use as fodder, nutraceuticals or phytotherapeutics in ruminant production.

ETHNOPHARMACOLOGICAL RELEVANCE: Salt-tolerant plants are well adapted to the harsh conditions of the Mediterranean region, where have been used traditionally as food and medicines for human and animals. In addition, various species are currently recognized as sources of metabolites with pharmacological, cosmetical and nutraceutical interest. Nevertheless, ethnoveterinary data on salt-tolerant plants are dispersed in the literature and there are few discussions on its veterinary potential. Having in mind the rising interest on organic farming, alternatives to chemical substances in livestock production and concern for animal health and welfare practices, these plants may represent an untapped resource for animal management and veterinary purposes. In this sense, the purpose of this work is to summarize the ethnoveterinary knowledge on salt-tolerant plants described in the Mediterranean region, raising awareness to the potential of this group of plants to be used in veterinary science, targeting especially ruminants. MATERIAL AND METHODS: Literature search (2000-2020) was conducted using Web of Science and Science Direct databases. Ethnoveterinary reports (EVR) concerning salt-tolerant plants were summarized and filtered for ruminants. From the final 29 publications, EVR concerning therapeutic uses were categorized according to its ATCvet code and results analyzed. RESULTS: A total of 221 EVR were identified from 39 plants, belonging to 21 plant families, targetting ruminants. Ten EVR (4.5%) concerned uses of salt-tolerant species as animal feed, while around 75% of therapeutic uses was represented by three categories: alimentary tract and metabolism (QA; n = 75), dermatologicals (QD; n = 53) and genitourinary system and sex hormones (QG; n = 41). Pistacia lentiscus L., Foeniculum vulgare Mill., Dittrichia viscosa (L.) Greuter, Plantago major L. and Hordeum vulgare L. were the most cited species in the latter categories. CONCLUSIONS: The ethnoveterinary knowledge on salt-tolerant species hints some plants of veterinary pharmacological potential, but other species deserve further notice. This information should serve as a basis and, coupled with the currently available scientific data on bioactive properties and chemical composition of salt-tolerant species, inspire additional research on the exploitation of this botanical group, as sources of novel products for ruminant nutrition, health and quality of its products.

Palyno-morphological investigations of halophytic taxa of Amaranthaceae through SEM from Salt range of Northern Punjab, Pakistan.

The pollen morphology of 11 salt tolerant plant species of family Amaranthaceae from the salt range of Northern Punjab, Pakistan has been studied. The palyno-morphological characters were examined using light and scanning electron microscope. The examined all salt tolerant species have a slight difference in size but have similarity in shape, pore ornamentation, and polarity. The observed morphological characters of pollen grains were pollen symmetry, size, shape, pore ornamentation, pore size, number of pores, exine thickness, polar and equatorial diameter and, P/E ratio. Apolar type of pollens has been observed in all species. Shape of pollens was spheroidal. Exine sculpturing of pollen grains was scabrate (six spp), microechinate (four spp), and microechinate-scabrate (one spp). Different pori numbers were observed in different species. The pantoporate aperturate and sunken pore ornamentation have been reported in all species. A pollen taxonomic key was developed using examined morphological characters for the accurate identification of halophytic taxa. The high fertility and low sterility of pollens confirmed that the selected halophytes are well-established in the salt region. The findings highlight the taxonomic significance of pollen morphology in correct identification and differentiation of salt tolerant plant species.

Impact of soil salinity on the microbial structure of halophyte rhizosphere microbiome.

The rhizosphere microbiome plays a significant role in the life of plants in promoting plant survival under adverse conditions. However, limited information is available about microbial diversity in saline environments. In the current study, we compared the composition of the rhizosphere microbiomes of the halophytes Urochloa, Kochia, Salsola, and Atriplex living in moderate and high salinity environments (Khewra salt mines; Pakistan) with that of the non-halophyte Triticum. Soil microbiomes analysis using pyrosequencing of 16S rRNA gene indicated that Actinobacteria were dominant in saline soil samples whereas Proteobacteria predominated in non-saline soil samples. Firmicutes, Acidobacteria, Bacteriodetes and Thaumarchaeota were predominant phyla in saline and non-saline soils, whereas Cyanobacteria, Verrucomicrobia, Gemmatimonadetes and the unclassified WPS-2 were less abundant. Sequences from Euryarchaeota, Ignavibacteriae, and Nanohaloarchaeota were identified only from the rhizosphere of halophytes. Dominant halophilic bacteria and archaea identified in this study included Agrococcus, Armatimonadetes gp4, Halalkalicoccus, Haloferula and Halobacterium. Our analysis showed that increases in soil salinity correlated with significant differences in the alpha and beta diversity of the microbial communities across saline and non-saline soil samples. Having a complete inventory of the soil bacteria from different saline environments in Pakistan will help in the discovery of potential inoculants for crops growing on salt-affected land.

Halophytic herbs of the Mediterranean basin: An alternative approach to health.

Wild native species are usually grown under severe and stressful conditions, while a special category includes halophytic species that are tolerant to high salinity levels. Native halophytes are valuable sources of bioactive molecules whose content is higher in saline than normal conditions, since the adaptation to salinity mechanisms involve apart from changes in physiological functions the biosynthesis of protectant molecules. These compounds include secondary metabolites with several beneficial health effects which have been known since ancient times and used for medicinal purposes. Recent trends in pharmaceutical industry suggest the use of natural compounds as alternative to synthetic ones, with native herbs being strong candidates for this purpose due to their increased and variable content in health promoting compounds. In this review, an introductory section about the importance of native herbs and halophyte species for traditional and modern medicine will be presented. A list of the most important halophytes of the Mediterranean basin will follow, with special focus on their chemical composition and their reported by clinical and ethnopharmacological studies health effects. The review concludes by suggesting future requirements and perspectives for further exploitation of these valuable species within the context of sustainability and climate change.

Nutrient resorption or accumulation of desert plants with contrasting sodium regulation strategies.

Desert plants are thought to rely more heavily on nutrient resorption due to the infertile soil. However, little is known regarding the phylogenetic effects on this traits, specifically for halophytes. Here we determined contents of nitrogen (N), phosphorus (P), potassium (K), sodium (Na), calcium (Ca) and magnesium (Mg) in 36 desert plants in a hyper-arid environment. The patterns of resorption or accumulation of the six elements were compared among plant groups with diverse leaf Na regulation strategies: i.e., euhalophytes (Eu), secretohalophytes (Se), pseudohalophytes (Ps) and glycophytes (Gl). Overall, N, P, K presented strict resorption across all groups, but no more efficient than global estimations. Ca and Mg tended to be resorbed less or accumulated during leaf senescence. Significant phylogenetic signal of both leaf Na content and plant group implies the pivotal role of Na regulation in the adaptation of plants to desert environment. Resorption proficiency, rather than resorption efficiency, is more phylogenetically conservative and more relevant to leaf functional traits.

Mercury uptake by halophytes in response to a long-term contamination in coastal wetland salt marshes (northern Adriatic Sea).

Mercury (Hg) distribution in saltmarsh sediments and in three selected halophytes (Limonium narbonense, Sarcocornia fruticosa and Atriplex portulacoides) of a wetland system (Marano and Grado Lagoon, Italy) following a contamination gradient in sediments was investigated. The Hg uptake was evaluated at the root system level by calculating the enrichment factor (EF) and in the aboveground tissues by means of the translocation factor (TF). The related methylmercury (MeHg) concentrations in the halophytes were also investigated with regard to the location of the sites and their degree of contamination. Hg concentration in halophytes seemed poorly correlated both with the total Hg in rhizo-sediments and with the specific plant considered, supporting the evidence that the chemico-physical parameters of sediments could significantly affect metal availability for plants. Hg concentrations in roots increased with depth and were 20-fold higher than content measured in related rhizo-sediments (high EF). A low content of Hg is translocated in aboveground tissues (very low TF values), thus highlighting a kind of avoidance strategy of these halophytes against Hg toxicity. MeHg values were comparable between the two sites and among species, but the translocation from below- to aboveground plant tissues was more active.

Cloning of PEPC-1 from a C4 halophyte Suaeda aralocaspica without Kranz anatomy and its recombinant enzymatic activity in responses to abiotic stresses.

Phosphoenolpyruvate carboxylase (PEPC) is a key enzyme of C4 photosynthetic pathway and plays an important biochemical role in higher plants and micro organisms. To gain understanding of the role of PEPC in stress adaptation in plant, we cloned PEPC gene from Suaeda aralocaspica, a C4 species without Kranz anatomy, and performed a series of experiments with PEPC gene expressed in Escherichia coli under various abiotic stresses. Results showed that, based on the homology cloning and 5'-RACE technique, the full-length cDNA sequence of PEPC (2901 bp) from S. aralocaspica was obtained, which shares the typical conserved domains to documented PEPCs and was identified as PEPC-1 in accord to the reported partial sequence (ppc-1) in S. aralocaspica. qRT-PCR analysis revealed the expression patterns of PEPC-1 and PEPC-2 (known as ppc-2, another plant type of PEPC) in S. aralocaspica, suggesting that PEPC-1 was up-regulated during seed germination and under NaCl stress, and presented higher level in chlorenchyma than other tissues, which were significantly different with PEPC-2. Afterwards, PEPC-1 was recombinant in E. coli (pET-28a-PEPC) and expressed as an approximate 110 kDa protein. Under various abiotic stresses, the recombinant E. coli strain harboring with PEPC-1 showed significant advantage in growth at 400-800 mmol L(-1) NaCl, 10-20% PEG6000, 25 and 30 °C lower temperature, 50-200 µmol L(-1) methyl viologen, and pH 5.0 and 9.0 condition, compared to control. Further analysis of the enzymatic characteristics of the recombinant PEPC-1 suggests that it was the higher enzyme activity of PEPC-1 which might confer the stress tolerance to E. coli. We speculate that over expression of PEPC-1 is probably related to regulation of oxaloacetate (OAA) in tricarboxylic acid (TCA) cycle in E. coli, which may contribute to further understanding of the physiological function of PEPC in S. aralocaspica.

[Estimation of endophytic bacterial diversity in root of halophytes in Northern Xinjiang by high throughput sequencing].

Objective: We studied the diversity of endophytic bacterial communities in different species of halophytes growing in the same saline habitat, and analyzed the effect of rhizosphere soil physicochemical properties on endophytic bacterial communities. Methods: PCR-based Roche FLX 454 pyrosequencing was applied to reveal the diversity of endophytic bacteria. Results: Endophytic bacterial communities of the 16 species of halophytes mainly included 4 phyla, which were Proteobacteria, Tenericutes, Actinobacteria and Firmicutes. In terms of plant species classification, colonial differences existed among plant species at perspectives of composition of bacterial taxa; in the case of plant genus level, endophytic bacteria of different halophyte plant species but belonging to same plant genus exhibited similarity; as to plant family level, Actinobacteria and Proteobacteria comprised the main abundant phyla of the halophytes belonging to Chenopodiaceae; Proteobacteria comprised the main abundant phyla of the halophytes belonging to Zygophyllaceae; Tenericutes comprised the main abundant phyla of the halophytes belonging to Tamaricaceae; Proteobacteria, Fimicutes and Actinobacteria comprised the main abundant phyla of the halophytes belonging to Plumbaginaceae. The Cl- in rhizosphere soil has significant effect on endophytic bacterial community structure. Moreover, there is a strong correlation between bacterial community and the combination of Cl-, Mg2+ and total nitrogen. Conclusion: Halophytes harbors diverse endophytic bacteria. In the same saline habitat, the distribution of endophytic bacteria showed host plant species-specific, and the Cl- in rhizosphere soil was one of the factors determined the endophytic bacterial community.

Description of a halocin-producing Haloferax larsenii HA1 isolated from Pachpadra salt lake in Rajasthan.

Haloarchaea grow in the extreme environment, such as high salt concentration, and secrete antimicrobial peptides known as halocins. Identification of Haloferax larsenii strain HA1 was carried out using biochemical and molecular methods. Strain HA1 was found as a strict aerobe, catalase positive and Gram negative. It was able to grow optimally at 15 % NaCl (w/v), 42 °C and pH 7.2. Strain HA1 was sensitive to bile acid, was resistant to chloramphenicol and could not utilize arginine. Halocin, produced by strain HA1, was stable up to 100 °C and in a pH range of 5.0-9.0. Antimicrobial activity was not affected by organic solvents, surfactants and detergents, but it was completely lost in the presence of proteinase K, suggesting proteinaceous nature of the compound. It was halocidal against indicator strain Hfx. larsenii HA10. The molecular weight of halocin HA1 was found to be ~14 kDa. These properties of halocin HA1 may be applicable to the preservation of salted foods.

A spatial pattern analysis of the halophytic species distribution in an arid coastal environment.

Obtaining information about the spatial distribution of desert plants is considered as a serious challenge for ecologists and environmental modeling due to the required intensive field work and infrastructures in harsh and remote arid environments. A new method was applied for assessing the spatial distribution of the halophytic species (HS) in an arid coastal environment. This method was based on the object-based image analysis for a high-resolution Google Earth satellite image. The integration of the image processing techniques and field work provided accurate information about the spatial distribution of HS. The extracted objects were based on assumptions that explained the plant-pixel relationship. Three different types of digital image processing techniques were implemented and validated to obtain an accurate HS spatial distribution. A total of 2703 individuals of the HS community were found in the case study, and approximately 82% were located above an elevation of 2 m. The micro-topography exhibited a significant negative relationship with pH and EC (r = -0.79 and -0.81, respectively, p < 0.001). The spatial structure was modeled using stochastic point processes, in particular a hybrid family of Gibbs processes. A new model is proposed that uses a hard-core structure at very short distances, together with a cluster structure in short-to-medium distances and a Poisson structure for larger distances. This model was found to fit the data perfectly well.

Taxonomic complexity in the halophyte Limonium vulgare and related taxa (Plumbaginaceae): insights from analysis of morphological, reproductive and karyological data.

BACKGROUND AND AIMS: Limonium is a well-known example of a group of plants that is taxonomically complex due to certain biological characteristics that hamper species' delineation. The closely related polyploid species Limonium vulgare Mill., L. humile Mill. and L. narbonense Mill. are defined species and can be used for studying patterns of morphological and reproductive variation. The first two taxa are usually found in Atlantic Europe and the third in the Mediterranean region, but a number of intermediate morphological forms may be present alongside typical examples of these species. This study attempts to elucidate morphological, floral and karyological diversity representative of these taxa in the Iberian Peninsula. METHODS: The extent of morphological differentiation was tested through comparison of 197 specimens from both Portugal and Spain using 17 descriptive morphological characters and 19 diagnostic morphometric characters. Analyses of floral morphisms (heterostyly and pollen-stigma dimorphism) and karyological determinations were also conducted. KEY RESULTS AND CONCLUSIONS: Discriminant analysis using morphometric variables reliably assigned individuals in natural populations to their respective groups. In addition, the results provide the first direct evidence that L. narbonense and a new species, LIMONIUM MARITIMUM: Caperta, Cortinhas, Paes, Guara, Espírito-Santo and Erben, SP NOV: , related to L. vulgare are present on Portuguese coasts. Most of these species are found together in mixed populations, especially L. vulgare and L. narbonense. It is hypothesized that taxonomic biodiversity found in sites where distinct species co-occur facilitates the evolutionary processes of hybridization, introgression and apomixis. This study therefore contributes to the elucidation of the taxonomic diversity in L. vulgare-related species and may also help in implementing future conservation programmes to maintain the evolutionary processes generating biodiversity.

Metal phytoremediation by the halophyte Limoniastrum monopetalum (L.) Boiss: two contrasting ecotypes.

The phytoremediation potential of the halophyte Limoniastrum monopetalum for the removal of Cd and Pb from polluted sites is assessed in this work. Two pot experiments were conducted; the first with wild L. monopetalum grown on soil polluted with Cd and Pb irrigated at different salinities, and the second with commonly cultivated ornamental L. monopetalum grown on soil polluted with Cd irrigated also at different salinities. The data revealed that wild L. monopetalum is a Cd and Pb tolerant plant able to accumulate at least 100 ppm of cadmium in its shoots without showing any significant decrease in terms of biomass production, chlorophyll content or water content suggesting that it could be an accumulator of Cd. Pb above-ground accumulation was kept at low levels with the majority of Pb localized in the roots. On the other hand, contrasting results were obtained for ornamental L. monopetalum which although it was found to be also Cd tolerant, Cd accumulation in its tissues was kept at significantly lower levels especially compared to that of the wild ecotype. In addition for ornamental L. monopetalum salinity did not have a positive effect on Cd accumulation and translocation as observed in the wild type and in other halophytes. Analysis of the salt excretion crystals on the leaf surface confirmed that wild and cultivated ornamental L. monopetalum excrete cadmium and lead through their salt glands as a possible metal detoxification mechanism, although the amount excreted by the ornamental L. monopetalum is significantly less.

Repeated evolution of salt-tolerance in grasses.

The amount of salt-affected agricultural land is increasing globally, so new crop varieties are needed that can grow in salt-affected soils. Despite concerted effort to develop salt-tolerant cereal crops, few commercially viable salt-tolerant crops have been released. This is puzzling, given the number of naturally salt-tolerant grass species. To better understand why salt-tolerance occurs naturally but is difficult to breed into crop species, we take a novel, biodiversity-based approach to its study, examining the evolutionary lability of salt-tolerance across the grass family. We analyse the phylogenetic distribution of naturally salt-tolerant species on a phylogeny of 2684 grasses, and find that salt-tolerance has evolved over 70 times, in a wide range of grass lineages. These results are confirmed by repeating the analysis at genus level on a phylogeny of over 800 grass genera. While salt-tolerance evolves surprisingly often, we find that its evolution does not often give rise to a large clade of salt-tolerant species. These results suggest that salt-tolerance is an evolutionarily labile trait in grasses.

Ecological investigation of three geophytes in the Deltaic Mediterranean coast of Egypt.

This study was conducted to investigate the ecological features of three geophytes namely Asparagus stipularis, Cyperus capitatus and Stipagrostis lanata which grow naturally in the Nile Delta coast of Egypt. C. capitatus and S. lanata are growing in non-saline sandy soils and can tolerate drought stress while, A. stipularis is growing in saline and non-saline sandy and calcareous clay soils and can tolerate drought and salt stress. Multivariate analysis of the vegetation of 100 sampled stands supporting growth of the three geophytic species in the study area led to the recognition of four vegetation groups namely, (A) Alhagi graecorum, (B) Cyperus capitatus, (C) Lycium schweinfurthii var. schweinfurthii-Asparagus stipularis and (D) Juncus acutus subsp. acutus. Vegetationally, the vegetation groups associated with the three species can be distinguished into two community types. The first one is psammophytic community comprising vegetation groups A and B that may represent the non-saline sand formations (flats, hummocks and dunes). The second one is halophytic community including vegetation groups C and D that may represent the saline sand flats and salt marsh habitat types, respectively. Sodium adsorption ratio, electrical conductivity, sodium cation, chlorides, silt and sand fractions, pH value, moisture content, bicarbonates and available phosphorus were the most effective soil factors that controlling the abundance and distribution of the plant communities associated with the investigated geophytes. This study showed the ecological features of the selected geophytes in terms of their habitats, associated plant communities and the most edaphic factors controlling their richness and distribution in the study area.

Methyl chloride emissions from halophyte leaf litter: dependence on temperature and chloride content.

Methyl chloride (CH(3)Cl) is the most abundant natural chlorine containing compound in the atmosphere, and responsible for a significant fraction of stratospheric ozone destruction. Understanding the global CH(3)Cl budget is therefore of great importance. However, the strength of the individual sources and sinks is still uncertain. Leaf litter is a potentially important source of methyl chloride, but factors controlling the emissions are unclear. This study investigated CH(3)Cl emissions from leaf litter of twelve halophyte species. The emissions were not due to biological activity, and emission rates varied between halophyte species up to two orders of magnitude. For all species, the CH(3)Cl emission rates increased with temperature following the Arrhenius relation. Activation energies were similar for all investigated plant species, indicating that even though emissions vary largely between plant species, their response to changing temperatures is similar. The chloride and methoxyl group contents of the leaf litter samples were determined, but those parameters were not significantly correlated to the CH(3)Cl emission rate.

Plant species potentially useful in the phytostabilization process for the abandoned CMC mining site in northern Cyprus.

The Cupper Mining Company (CMC)'s site located in Lefke-Gemikonagi, Northern Cyprus has been a continuous source of highly dangerous contamination for the surrounding environment, the Lefke region, and the neighboring ecosystems and settlements. Rehabilitation and reuse possibilities of the CMC site due to its vital importance have kept its place in the agenda of Northern Cyprus. Phytostabilization appears to be a convenient and less expensive method that can immediately be used for reducing the negative impacts of the mining site on the region. The main purpose of this study is to identify potential candidate plant species, adapted to grow on polluted sites, for revegetation in the CMC site. Within this context, the method of the study can be summarized as follows: literature review for examining potential candidate plant species for pyhtostabilization in arid and semiarid regions, especially the ones suitable both for the existing ecological and present conditions of Cyprus; identification of native and/or cultural plant species survived in the heavily polluted mining site, and definition of a number of candidate plant species for the study site. The result of sampling revealed that 23 plant species thrive well in the contaminated site. As a result of the literature review and considering drought, metal, salt tolerant features of semiarid environment in the region, 5 tree, 4 shrub, and 23 herbaceous plant species were proposed for starting revegetation with the purpose of phytostabilization on the CMC mining site.

[Response characteristics of the field-measured spectrum for the four general types of halophyte and species recognition in the northern slope area of Tianshan Mountain in Xinjiang].

Based on the field-measured Vis-NIR reflectance of four common types of halophyte (Achnatherum splendens(Trin.) Nevski, Sophora alopecuroides L., Camphorosma monspeliaca L. subsp. lessingii(L.)Aellen, Alhagi sparsifolia shap) within given spots in the Northern Slope Area of Tianshan Mountain in Xinjiang, the spectral response characteristics and species recognition of these types of halophyte were analyzed. The results showed that (Alhagi sparsifolia shap) had higher chlorophyll and carotenoid by CARI and SIPI index. (Sophora alopecuroides L. was at a vigorously growing state and had a higher NDVI compared with the other three types of halophyte because of its greater canopy density. But its CARI and SIPI values were lower due to the influence of its flowers. (Sophora alopecuroides L.) and (Camphorosma monspeliaca L. subsp. lessingii(L.)) had stable REPs and BEPs, but REPs and BEPs of (Achnatherum splendens(Trin.)Nevski, Aellen, Alhagi sparsifolia shap) whose spectra red shift and spectra blue shift occurred concurrently obviously changed. There was little difference in spectral curves among the four types of halophyte, so the spectrum mixing phenomenon was severe. (Camphorosma monspeliaca L. subsp. lessingii (L.)Aellen) and (Alhagi sparsifolia shap) could not be separated exactly in a usual R/NIR feature space in remote sensing. Using the stepwise discriminant analysis, five indices were selected to establish the discriminant model, and the model accuracy was discussed using the validated sample group. The total accuracy of the discriminant model was above 92% and (Achnatherum splendens(Trin.)Nevski) and (Camphorosma monspeliaca L. subsp. lessingii(L.)Aellen) could be respectively recognized 100% correctly.