Salicornia

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Other Names : pickleweed, glasswort, sea beans, sea asparagus, crow’s foot greens, samphire, marsh samphire, "titines de souris" (mouse tits).
Salicornia is a halophyte, belonging to Amaranthaceae family (Singh et al. 2014). In fact, Salicornia name has originated from the Latin word meaning ‘salt’. Studies report that some species, for example Salicornia europaea show tolerance towards salinity as high as 3 % NaCl (Yamamoto et al. 2009). This fleshy plant is found at the edges of wetlands, marshes, sea shores, and mudflats, actually on most alkaline flats (Smillie 2015). It has a geographical distribution spanning 4 continents such as North America, Asia, Africa and Europe. Salicornia in spring and summer is green and fit for consumption, b

Salicornia in autumn is red and purple, with high salt concentration, is not suitable for food purpose</p></div></div></div>

Table 1

Most studied species of Salicornia and their geographical distributions

<thead></thead><tbody></tbody>
No.Botanical nameCommon namesGeographical RangeReferences
1

Salicornia europaea

Common glasswortBritain, France, IrelandZhang et al. (<a href="#CR86" rid="CR86" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674017">2014</a>)
2

Salicornia bigelovii

Dwarf glasswortUSA, MexicoZhang et al. (<a href="#CR87" rid="CR87" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674019">2015</a>)
3

Salicornia brachiata

Umari keeraiIndiaJha et al. (<a href="#CR29" rid="CR29" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673982">2012</a>)
4

Salicornia virginica

American glasswort, pickleweedCanada, USA, MexicoRosso et al. (<a href="#CR65" rid="CR65" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674027">2005</a>)
5

Salicornia maritima

Slender glasswortCanada, USA, Mexico–
6

Salicornia ramosissima

Purple glasswortFrance, IberiaIsca et al. (<a href="#CR25" rid="CR25" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674048">2014</a>)
7

Salicornia herbacea

–KoreaCho et al. (<a href="#CR7" rid="CR7" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674033">2015</a>)
8

Salicornia persica

–IranSingh et al. (<a href="#CR73" rid="CR73" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673986">2014</a>)

Salicornia has been historically used for both non-edible and edible purposes. Usage of the plant as a source of soda (sodium carbonate) for glass making dates back to centuries. Oriental pharmacopeia reports its medicinal uses. The efficacy of Salicornia herbacea against oxidative stress, inflammation, diabetes, asthma, hepatitis, cancer, gastroenteritis has been reported (Essaidi et al. <a href="#CR14" rid="CR14" class=" bibr popnode">2013</a>). Food use was not altogether new, with multiple reports of their consumption as a salt source. However, recent dearth in food availability, quest for sustainable food sources and foraging interest has pushed this genus to prominence. This plant’s aerial parts are consumed in salads or processed into pickles, beverages etc.; an interest that has taken off only in the last few years. This review explores the present status of this genus in the food arena and evaluates its scope ahead.

Food uses

For its saltiness and crunchiness, it is used as a green salad. Even in some cultures, it is considered a delicacy. Only the green, tender parts are recommended for edibility, the reddish being too high in salinity and silica. In some communities, the shoots are processed into beverages like nuruk (a type of fermentation starter), makgeolli (a Korean rice wine), or vinegar (Song et al. <a href="#CR75" rid="CR75" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674029">2013</a>; Kim et al. <a href="#CR36" rid="CR36" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673990">2013</a>). A study found that Salicornia not only stimulates the fermenting microbe propagation but also improves the quality of vinegar (Seo et al. <a href="#CR68" rid="CR68" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674002">2010</a>). Apart from direct consumption, these plants have been found fitting as a source of dietary salt. S. herbacea powder was transformed into spherical granules, which showed potential to be used like NaCl (Shin and Lee <a href="#CR71" rid="CR71" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674035">2013</a>). A study found that 1.5 % of Salicornia salt as a partial substitute for NaCl can be added to frankfurters for texture improvement without any perceivable side effects (Kim et al. <a href="#CR37" rid="CR37" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674023">2014</a>). The positive effect of the fortification was manifested in increased cooking yield and emulsion stability (Kim et al. <a href="#CR37" rid="CR37" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673961">2014</a>). In another interesting study, Salicornia bigelovii salt was discovered to prevent hypertensive effect, normally associated with NaCl. Ameliorative effect on kidney and liver was established which correlated to lower serum creatinine level (Zhang et al. <a href="#CR87" rid="CR87" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673967">2015</a>). Further, superoxide dismutase (SOD) and Na(+)-K(+)-ATPase activity increased while malondialdehyde (MDA) content decreased, suggesting beneficial effect on antioxidant profile of the body (Zhang et al. <a href="#CR87" rid="CR87" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673987">2015</a>). The overall food value of Salicornia has got a boost from the phytochemical profiling studies that have unveiled an array of nutritive components, as discussed below.

Phytochemicals

Salicornia plants have been screened for phytochemical profile and presence of a range of carbohydrates, proteins, oils, phenolic compounds, flavonoids, sterols, saponins, alkaloids, and tannins have been reported. Water and alcohol extraction followed by component profiling has indexed many potentially bioactive compounds. Studies have reported the presence of dietary fibers, bioactive polysaccharides, proteins, lipids, sterols, flavonoids, and minerals (Mg, Ca, Fe, K) in S. herbacea (Essaidi et al. <a href="#CR14" rid="CR14" class=" bibr popnode">2013</a>).

An immunomodulatory polysaccharide was isolated from S. herbacea (Lee et al. <a href="#CR40" rid="CR40" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674015">2006</a>; Im et al. <a href="#CR23" rid="CR23" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674045">2006</a>). Monosaccharide composition analysis of Salicornia brachiata fraction revealed the dominance of rhamnose, arabinose, mannose, galactose, and glucose, with meager presence of ribose and xylose (Sanandiya and Siddhanta <a href="#CR67" rid="CR67" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673964">2014</a>). Proteomic analysis (matrix-assisted laser desorption/ionization-time of flight (MALDI-TOF) mass spectrometry) revealed high protein contents of the seeds. Based on the detection of high disulfide linkages in the globulin proteins, it was deemed suitable for consumption, as sulfur-rich proteins are generally fit for nutrition (Jha et al. <a href="#CR29" rid="CR29" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674037">2012</a>). Gas chromatography mass spectrometry (GC–MS) was used to profile lipidome of S. ramosissima which showed esterified and free fatty acids, fatty alcohol, sterols, alkanes and aromatic acid derivatives. Among the dominant components, palmitic acid, tetracosanol and octacosanol were significant (Isca et al. <a href="#CR25" rid="CR25" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674046">2014</a>). Tetracosanol, the aliphatic alcohol has been identified to possess α-amylase ability, which makes it relevant in diabetes therapy (Jhong et al. <a href="#CR30" rid="CR30" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674003">2009</a>). Octacosanol, the high-molecular-weight aliphatic alcohol has been known to be a component of cholesterol-lowering drugs such as policosanol (Liu et al. <a href="#CR45" rid="CR45" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674012">2015</a>). By metabolomic analysis, S. brachiata was detected to be rich in sulfur amino acids and polyunsaturated fatty acids (PUFAs 55–64 %) (Mishra et al. <a href="#CR48" rid="CR48" class=" bibr popnode">2015</a>). Presence of selenium in S. brachiata was detected (Mishra et al. <a href="#CR48" rid="CR48" class=" bibr popnode">2015</a>). Selenium is an essential micronutrient for growth and robust antioxidant effects, deficiency of which has been documented to impair the immune system (Finley <a href="#CR17" rid="CR17" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674031">2005</a>). In this regard, it appears justified to evaluate dietary extraction of selenium from Salicornia. A study found S. herbacea seed oil to be stable to oxidation and eligible to be used in food processing (Choi et al. <a href="#CR8" rid="CR8" class=" bibr popnode">2014</a>). The oil composed of linoleic acid, oleic acid, arachidic acid, palmitic acid, tocopherol (α, γ, δ type), phenol, chlorophyll, and β carotene, was free of rancidity during a 60 day dark storage period (Choi et al. <a href="#CR8" rid="CR8" class=" bibr popnode">2014</a>). Stigmastanol, 24-ethyl-δ(22)-coprostenol and several other bioactive fatty alcohols were detected (Isca et al. <a href="#CR25" rid="CR25" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674010">2014</a>). In another study, a chlorogenic acid, 3-caffeoyl-4-dicaffeoylquinic acid was detected in S. herbacea extract (Hwang et al. <a href="#CR22" rid="CR22" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674028">2010</a>). Another study identified pentadecyl ferulate, stigmasterol, ergosterol, vanillic aldehyde and scopoletin in S. herbacea (Wang et al. <a href="#CR81" rid="CR81" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674013">2013</a>). Chromatography could detect β sitosterol (1), stigmasterol (2), uracil (3), and isorhamnetin-3-O-β-D-glucopyranoside in S. herbacea (Lee et al. <a href="#CR39" rid="CR39" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673992">2004</a>). Saponins were identified in butanol fraction of S. herbacea extract, some of which exerted antioxidant effect (Kim et al. <a href="#CR35" rid="CR35" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673985">2012</a>). Two new noroleanane-type triterpene saponins, Salbige A and B were isolated from the aerial parts of S. herbacea (Zhao et al. <a href="#CR88" rid="CR88" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674000">2014</a>). A new nortriterpene saponin, bigelovii D with antifungal potential was isolated from the hydroalcoholic extract of S. bigelovii (Shan et al. <a href="#CR69" rid="CR69" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_792533393">2015</a>).These studies contributed towards phytochemical composition of this genus and emphasized the significant role of non-targeted metabolomics in further component analysis.

Antioxidant

Aqueous and methanol extracts of the enzyme-treated S. herbacea possessed potential antioxidant effects as observed in vitro on rat liver microsomal lipid peroxidation (Jang et al. <a href="#CR26" rid="CR26" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674004">2007</a>). The butanol fraction S. herbacea methanol extract exerted scavenging activities attributable to its active principle isorhamnetin 3-O-β-d-glucopyranoside. The glucopyranoside intervened with inflammatory pathways via manipulation of cytokine profile (Kim et al. <a href="#CR34" rid="CR34" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673974">2009</a>). In a rat model, S. herbacea vinegar when orally administered (at 7 g/kg water) showed free radical scavenging and SOD-like activities. Furthermore, the vinegar-administered rats showed higher glycogen accumulation in liver and muscles, higher lactate and ATP metabolism, promoting enzyme activities such as muscle creatine kinase and lactate dehydrogenase, whereas serum fatigue biomarkers such as ammonia, lactate and inorganic acid were markedly decreased (Cho et al. <a href="#CR7" rid="CR7" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673994">2015</a>).

Antiproliferative

Polysaccharides from botanical sources exerting anticancer properties have been well-documented (Chang <a href="#CR6" rid="CR6" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673971">2002</a>; Patel and Goyal <a href="#CR57" rid="CR57" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673977">2012</a>). In this regard, many Salicornia polysaccharides have also shown promise, validated through in vitro and in vivo models. Crude as well as purified polysaccharides from S. herbacea (at 0.5–4 mg/ml) demonstrated anti-proliferation of human colon cancer HT-29 cells when incubated for 24–48 h (Ryu et al. <a href="#CR66" rid="CR66" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674022">2009</a>). The mechanism of the cancer cell death was attributed to cell cycle arrest at G2/M phase, followed by apoptosis. Expression of the p53 tumor suppressor gene and the cyclin-dependent kinase inhibitor 1 (CDK inhibitor p21) were observed (Ryu et al. <a href="#CR66" rid="CR66" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674016">2009</a>). In another study S. herbacea -derived chlorogenic acid, 3-caffeoyl, 4-dicaffeoylquinic acid exerted control on metastasis of human fibrosarcoma HT-1080 cell line (Hwang et al. <a href="#CR22" rid="CR22" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674018">2010</a>). The invasion restraint was mediated through the inhibition of activator protein-1 AP-1 and signaling pathways involving protein kinase C (PKC) delta (repressing phosphorylation of ERK, p38 MAPK, and JNK) and three MAPKs, culminating in reduced activation of matrix metalloproteinase (MMP)-9 (Hwang et al. <a href="#CR22" rid="CR22" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674021">2010</a>). Another study found that pentadecyl ferulate from S. herbacea ethyl acetate extract possesses antioxidant effect and exerts anticancer response towards human hepatocellular liver carcinoma HepG2 and human lung adenocarcinoma epithelial A549 cells, along with phytol and γ-linolenic acid (Wang et al. <a href="#CR81" rid="CR81" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674040">2013</a>). Saponins, Salbige A and B, isolated from S. herbacea exerted antiproliferative activities towards A549 cells, while a pheophorbide (a chlorophyll catabolite) inhibited both A549 and HepG2 cancer cells (Zhao et al. <a href="#CR88" rid="CR88" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673973">2014</a>).

Lipid lowering

Hyperlipidemia is a major cause of morbidity and mortality across the globe. S. herbacea ethanol (50 %) extract exerted lipid lowering in mice model when prescribed for 10 weeks, via suppression of lipogenesis related genes [sterol regulatory element-binding protein 1 (SREBP1a), fatty acid synthase (FAS), and glycerol-3-phosphate acyltransferase (GAPT)] (Park et al. <a href="#CR55" rid="CR55" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673989">2006</a>). In addition, flavonoids from this plant were observed to exert adipogenic inhibition in 3T3-L1 adipocytes (Kong and Seo <a href="#CR38" rid="CR38" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673965">2012</a>). The isorhamnetin compound reduced adipogenic differentiation by down-regulation of peroxisome proliferator-activated receptor-γ (PPARγ), CCAAT/enhancer-binding proteins (C/EBPα), SREBP1, and the adipocyte-specific proteins. Involvement of AMP-activated protein kinase (AMPK) was also observed (Kong and Seo <a href="#CR38" rid="CR38" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674044">2012</a>). In addition, 3-caffeoyl, 4-dihydrocaffeoylquinic acid extracted from this species prevented lipid accumulation by blocking SREBP-1c and FAS through LKB1/SIRT1 and AMPK activation as studied in HepG2 cells (Pil Hwang et al. <a href="#CR60" rid="CR60" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674026">2013</a>). Another mice study on this plant revealed that it can decrease body weight gain by controlling serum leptin and manipulating lipid synthesis genes as SREBP-1c, PPARγ and FAS. Intake of high fat diet along with the plant powder at 50 g/ kg dose conferred significantly better parameters compared to only high fat diet or high fat diet plus NaCl group (Pichiah and Cha <a href="#CR59" rid="CR59" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674054">2015</a>).

Antibacterial

Methanol extract of S. herbacea showed antibacterial activities, mediated by interference with cytochrome P450 CYP1A2, CYP3A4 and CYP2D6 enzymes (Essaidi et al. <a href="#CR14" rid="CR14" class=" bibr popnode">2013</a>). Another study explored the possibility of developing antimicrobial nanoparticles from Salicornia. S. brachiata-based gold nanoparticles were analyzed through a set of standard tools, which revealed its poly-dispersed, crystalline nature and 22–35 nm size (Essaidi et al. <a href="#CR14" rid="CR14" class=" bibr popnode">2013</a>). The particles decimated tested bacteria, manifested in zone of clearance on inoculated plates. Further, the nanoparticles showed synergistic activity with fluoroquinolone antibiotic ofloxacin (Ayaz Ahmed et al. <a href="#CR3" rid="CR3" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674052">2014</a>). Based on these exciting results, follow-up studies ought to be pursued.

Antidiabetic

Diabetes has assumed epidemic proportions in current times, due to pollution, and processed, calorie-rich food consumption, among other causal factors (Olokoba et al. <a href="#CR53" rid="CR53" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674025">2012</a>). Existing panel of antidiabetic drugs provoke side effects (Haque et al. <a href="#CR20" rid="CR20" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673969">2011</a>; Stein et al. <a href="#CR76" rid="CR76" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674043">2013</a>). In this regard, safer options to control hyperglycemia are being searched. Consequently, the ameliorative effect of S. herbacea powder on induced-diabetic rats was studied. When administered alone or recommended with exercise (in the form of swimming), it showed higher expression of liver and muscle glucose transporters GLUT-4 and GLUT-2 (Lee et al. <a href="#CR41" rid="CR41" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673998">2015</a>). Higher glycogen concentration in liver and muscle also corroborated the finding, heralding possible usage of the halophyte as an antidote to control diabetes (Lee et al. <a href="#CR41" rid="CR41" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674008">2015</a>).

Hepatoprotective

Liver is a vital gland for proper functionality of the body (Jarrar et al. <a href="#CR27" rid="CR27" class=" bibr popnode">2001</a>). Most analgesics induce adverse effect on liver tissues and functions (Bessone <a href="#CR4" rid="CR4" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674014">2010</a>). To evaluate hepatoprotective effects of a Korean herbal drink, of which Salicornia was a constituent, this study was conducted. The multi-herb potion ‘taemyeongcheong’ was administered to acetaminophen-stressed mice. At 500 mg/kg dose, the drink conferred protective effects on mice liver. Drop in the level of oxidative enzymes as alanine aminotransferase (ALT), aspartate aminotransferase (AST), alkaline phosphatase (ALP), and lactate dehydrogenase (LDH), and elevation in the level of antioxidative enzymes as catalase, superoxide dismutase, glutathione peroxidase, and glutathione was observed. Decline in expression of hepatic mRNA levels of TNF-α, IL-1β, IL-6, COX-2, and iNOS genes were observed, which can be linked to the lower degree of lesions to liver tissue (Yi et al. <a href="#CR84" rid="CR84" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673981">2015</a>).

Immunomodulatory

Polysaccharides from this plant have shown evidence of eliciting immune response. In this context, some pertinent studies have been discussed below. In an in vitro study, S. herbacea polysaccharides induced nitric oxide (NO) production from mouse peritoneal macrophages and mouse leukaemic monocyte macrophage RAW 264.7, through the activation of nuclear factor-kappaB/Rel (NF-kappaB/Rel) (Lee et al. <a href="#CR40" rid="CR40" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674020">2006</a>). Consequently, the polysaccharide stimulating macrophages that express iNOS gene came forth (Lee et al. <a href="#CR40" rid="CR40" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673996">2006</a>). In another study on S. herbacea, its polysaccharide demonstrated effect on monocyte activation and differentiation into macrophage (Im et al. <a href="#CR23" rid="CR23" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673980">2006</a>). RAW 264.7 cells elaborated cytokines such as tumor necrosis factor (TNF)-alpha and interleukin (IL)-1 beta, and nitric oxide (NO) when incubated with the polysaccharide. Further differentiation into macrophage was determined from higher adherence development in the monocytes (Im et al. <a href="#CR23" rid="CR23" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674036">2006</a>). Enhanced collagen-adherence is known to improve phagocytosis (Newman and Tucci <a href="#CR51" rid="CR51" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674032">1990</a>). Further work by same group of researchers reaffirmed that the purified polysaccharide worked in sync with IFN-γ to induce immune effector molecules as TNF-α, IL-1 β, and NO to differentiate the monocytes into the macrophages (Im et al. <a href="#CR24" rid="CR24" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673995">2007</a>).

Osteoprotective

Bone health is crucial for they make the framework of body and enable proper muscle movement. Nutrient deficiency, genetic or geriatric conditions impose bone malfunctions. Osteoporosis is a key bone disorder, for which one causal agent has been identified as higher bone adipogenesis (differentiation of stem cells into mature adipocytes) (Pino et al. <a href="#CR61" rid="CR61" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674047">2012</a>). In this regard, S. herbacea extract was observed to inhibit adipogenesis via manipulation of PPARγ, CCAAT/enhancer-binding protein (C/EBP)α and SREBP1c. Resultantly, osteogenesis improved, as evidenced in MC3T3-E1 pre-osteoblasts. Osteoblastogenesis markers as alkaline phosphatase (ALP), bone morphogenetic protein (BMP)-2, osteocalcin and collagen type I (collagen-I) lent support to bone formation induction by S. herbacea extract (Karadeniz et al. <a href="#CR32" rid="CR32" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674007">2014</a>).

Antiseptic food additive

Sepsis, caused by the compromised integrity of membrane barrier can be fatal (Li et al. <a href="#CR43" rid="CR43" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673960">2009</a>), so the discovery of the antiseptic effect of Salicornia is interesting, regarding which some relevant findings have been discussed here. High mobility group box 1 protein (HMGB1), a nuclear protein elaborated by activated leucocytes, is released in excess when inflammatory tissue damage renders the membrane porous (Tang et al. <a href="#CR78" rid="CR78" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673999">2010</a>; Lotze and Tracey <a href="#CR46" rid="CR46" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673979">2005</a>; Passali et al. <a href="#CR56" rid="CR56" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674055">2012</a>). In this scenario, inhibitors of HMGB1 are suggested to be a potential treatment for sepsis. S. herbacea-derived caffeoylated quinic acids showed anti-HMGB1 activity which exerted protection towards vasculature (Tuan et al. <a href="#CR79" rid="CR79" class=" bibr popnode">2015a</a>). The purified flavanones and chromone derivatives from the plant suppressed the release of HMGB1 in mice models, barricading the animal intestine from septic shocks (Tuan et al. <a href="#CR80" rid="CR80" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673968">2015b</a>). Role of the phytochemicals in hyperpermeability modulation needs to be investigated further, to better utilize the halophyte in fight against septicemia. While the precise mechanism is yet to be discovered for Salicornia, other studies have attributed anti- HMGB1 activity to be due to activated cholinergic anti-inflammatory pathway (Goldstein et al. <a href="#CR18" rid="CR18" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674024">2007</a>).

Cultivation

Salicornia is mulled to be the right candidate for reclamation of barren lands, salt flats, and sea shores. In short, they can be deemed for seawater agriculture. It is suggested that as global warming threatens to submerge more landmass, and freshwater is depleting, a shift to saline crop might be a viable option (Katschnig et al. <a href="#CR33" rid="CR33" class=" bibr popnode">2013</a>). Few plants can tolerate excess salt and among them few are edible. In this context, Salicornia seems to be a right candidate for cultivation (Singh et al. <a href="#CR73" rid="CR73" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674053">2014</a>). Regarding cultivation, different degrees of success have been observed in different parts of the world. Some of the Salicornia species are being farmed at commercial scale, for biodiesel, animal feed, and salt and oil extraction, e.g. S. bigelovii (Cybulska et al. <a href="#CR9" rid="CR9" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674050">2014</a>). This species produces oleaginous seeds which have been evaluated to be a promising feedstock for biodiesel production (Falasca et al. <a href="#CR15" rid="CR15" class=" bibr popnode">2014</a>). Introduction of Salicornia in arid lands of Saudi Arabia and Africa is being pondered and practiced (Fedoroff et al. <a href="#CR16" rid="CR16" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673972">2010</a>). Heavy metal removal is another possible usage of this plant. Controlled cultivation of various species procured from various habitats showed different results. S. bigelovii was grown in greenhouse conditions, which reflected that crop yield can vary depending on plants sourced from different habitats. Greenhouse milieu reduced biomass and fruit yield, though cultivation was successful (Bresdin et al. <a href="#CR5" rid="CR5" class=" bibr popnode">2016</a>). In addition, species-specific yield was observed, as seen with S. ramosissima, which produced more harvestable biomass than S. dolichostachya (Singh et al. <a href="#CR73" rid="CR73" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673976">2014</a>). Even if subsequent research dismisses Salicornia as unfit for human consumption, they might be purposed for other utilities, like biofuel harvest or livestock feeding.

Associated risks of Salicornia diet

Though plentiful studies have established beneficial effects of Salicornia, it has its share of concerns too. It is important to be aware of possible harmful reactions before consumption. Some alarming facts have been outlined below. Accumulation of heavy metals in the vegetation is a risk to consumer health (Lei et al. <a href="#CR42" rid="CR42" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674039">2015</a>). Wetlands are biodiversity-rich and are critical for ecological balance, but these unique ecosystems across the world are fragile now, in the face of increased anthropogenic activities (industrial effluent release, sewage treatment etc.) (Gutzwiller and Flather <a href="#CR19" rid="CR19" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673970">2011</a>; Anza et al. <a href="#CR2" rid="CR2" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674034">2014</a>). Most wetland plants are afflicted by widespread damage including Salicornia. This succulent is easily affected by metal and oil spill stressors. A study conducted on a marsh of California showed that the species Salicornia virginica suffers stress from heavy metals like chromium and vanadium (Rosso et al. <a href="#CR65" rid="CR65" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673997">2005</a>). Another study reported S. brachiata to be capable of imbibing cadmium, nickel and arsenic salts (Sharma et al. <a href="#CR70" rid="CR70" class=" bibr popnode">2010</a>). Yet another study reports the suitability of different Salicornia species as biomonitors of zinc and copper, emphasizing its relevance in metal remediation from water (Smillie <a href="#CR74" rid="CR74" class=" bibr popnode">2015</a>). The possibility of using Salicornia persica as a biofilter in a constructed wetland for effluent water released from a recirculating mariculture system was studied in Israel (Shpigel et al. <a href="#CR72" rid="CR72" class=" bibr popnode">2013</a>). These aspects might be promising from phyto-remediation perspective, definitely not from consumption standpoint.

In addition, Chenopodiaceae (goosefoot group of plants, a part of Amaranthaceae family) members are known to contain high oxalate content, which might be harmful to consumers (Norman et al. <a href="#CR52" rid="CR52" class=" bibr popnode">2013</a>). A review has described the adverse effects of dietary oxalic acid on consumer health, by reducing calcium bioavailability, causing renal stones, stunting bone growth, preventing blood coagulation etc. (Dolan et al. <a href="#CR12" rid="CR12" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674049">2010</a>).

High salt content in diet is a major risk factor, especially for hypertension. Excess sodium intake is known to hamper with rennin and angiotensin homeostasis, leading to endothelial dysfunction (Drenjančević-Perić et al. <a href="#CR13" rid="CR13" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673963">2011</a>). Halophytes, as their name suggests are known to thrive in saline areas and imbibe salt, storing them in specialized vacuoles (Priyashree et al. <a href="#CR63" rid="CR63" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673993">2010</a>). Ingestion of excess salt can aggravate blood pressure. In addition, Salicornia might contain iatrogenic iodine. A case study resulted that hyperconsumption of S. herbacea can lead to excess iodine in body, causing hypokalemic thyrotoxic paralysis (Yun et al. <a href="#CR85" rid="CR85" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673988">2014</a>). Discontinuation of Salicornia intake restored the potassium level and normalized thyroid imbalance (Yun et al. <a href="#CR85" rid="CR85" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673966">2014</a>).

Another concern is saponin toxicity. Amaranthaceae members have been characterized to contain high quantity of saponins. These glycosides have tissue necrotic (in small intestine, liver, kidney) (Diwan et al. <a href="#CR11" rid="CR11" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674006">2000</a>), gut permeability alteration (Onning et al. <a href="#CR54" rid="CR54" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674011">1996</a>), and adjuvant potential which can provoke immune system (Rajput et al. <a href="#CR64" rid="CR64" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674038">2007</a>).

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Discussion

From historical usage of this halophyte for glass making, the shift towards biofuel harvest occurred (Lieth and Al Masoom <a href="#CR44" rid="CR44" class=" bibr popnode">1993</a>). Salicornia though not primarily or widely consumed; its ingestion as food and medication is, however, not altogether new. Trials and nutritional assessments on it for human edibility are novel. As outlined in above sections, Salicornia both have its pros and cons as a food candidate. Additional research might better illuminate on its relevance for consumption. In this regard, some significant areas pertaining to it have been discussed below.

Chenopodiaceae members are known to contain high amount of crude protein, sulfur and minerals (Norman et al. <a href="#CR52" rid="CR52" class=" bibr popnode">2013</a>), which goes in favor of Salicornia as an edible plant. Mucilages are plant-derived polysaccharides with myriad roles such as food thickeners, binding agents, water holding agents, emulsifier etc. (Nayak et al. <a href="#CR50" rid="CR50" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674030">2010</a>). In this regard, Salicornia as a source of mucilage can be assessed. Amaranthaceae family member plumed cockscomb (Celosia argentea) elaborates an acidic polysaccharide celosian that has been characterized to ameliorate liver injuries (Hase et al. <a href="#CR21" rid="CR21" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673984">1997</a>). The healing effect was linked to immunomodulating effect via tumor necrosis factor-alpha (TNF-α), interleukin-1 beta (IL-1 β) and NO production (Hase et al. <a href="#CR21" rid="CR21" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673991">1997</a>). Consequently, Salicornia can also be evaluated for immune-modulating polysaccharides.

Previously, the adverse effects of saponin as a dietary ingredient have been mentioned. However, medicinally, saponins are crucial with an array of their health benefits reported so far, such as cytotoxic activity (Podolak et al. <a href="#CR62" rid="CR62" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674042">2010</a>). The variable biological effects of saponins stem from their diverse structural configurations. Based on the hydrophobic aglycone moiety, the glycosides can be categorized as triterpenoids, steroids or glycoalkaloidss (Moses et al. <a href="#CR49" rid="CR49" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674056">2014</a>). Amaranthaceae member C. argentea seed contains triterpenoid saponins (celosin) which shows in vitro antitumor and anti-inflammatory properties (Wu et al. <a href="#CR82" rid="CR82" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674001">2011</a>).

Apart from the validated nutrients present in the plant, its food candidature can be assumed from its plant family Amaranthaceae. Many conventional and emerging foods are sourced from this family (Amaranthaceae), which builds trust on the food potential of Salicornia. This family is at the forefront of valuable food sources e.g. beet, spinach, amaranthus and quinoa (Délano-Frier et al. <a href="#CR10" rid="CR10" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673962">2011</a>). Particularly, quinoa (Chenopodium quinoa Willd.), a related halophyte has surged to prominence in recent times, due to its high protein, lipids, fibers, vitamins and mineral contents (Maradini Filho et al. <a href="#CR47" rid="CR47" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673959">2015</a>). Only a few species have been evaluated so far, other species are largely inconspicuous, though there are more than 50 species under this genus. Investigation on these species is expected to reveal myriad other biological benefits. So far, among all Saicornia species, only S. herbacea has been the subject of lipid lowering effect, which provides ample scope to visit the potential of other species. In recent times polysaccharides from many higher plants (Strickland <a href="#CR77" rid="CR77" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674051">2001</a>), mushrooms (Akramiene et al. <a href="#CR1" rid="CR1" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603674041">2007</a>) and seaweeds (Jeong et al. <a href="#CR28" rid="CR28" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673978">2015</a>) have been validated as candidates as immune-therapeutics. These findings build the prospect of recruiting Salicornia as well, in immune modulation.

With due research input, this genus might be popularized for food and medicinal purposes. Pairing with compatible and complementary botanicals might improve efficacy, though cross reaction-caused adverse reactions must be monitored first.

Study on this genus is not a new area and several interesting findings have accumulated over the years. It is beyond the scope of this manuscript to furnish them all, yet it is justified to outline that this genus has been evaluated to shed light on marsh habitat loss, heavy metal accumulation and saline stress tolerance mechanisms of plants etc., Adaptation to salinity is particularly a well-pursued area, for its relevance to acclimatize vulnerable plants. A study reported of improved salt tolerance in transgenic alfalfa grass by over-expression of the S. europaea Na(+)/H(+) antiporter gene SeNHX1 (Zhang et al. <a href="#CR86" rid="CR86" class=" bibr popnode tag_hotlink tag_tooltip" id="__tag_603673975">2014</a>). Another study found that fresh water cultivation of Salicornia yields higher phenolic and flavonoid content over saline water cultivation (Kang et al. <a href="#CR31" rid="CR31" class=" bibr popnode">2015</a>). In addition, the fresh water-grown Salicornia demonstrated higher in vitro cytotoxic effects (Kang et al. <a href="#CR31" rid="CR31" class=" bibr popnode">2015</a>). However, it is the food and medicinal facet that needs to be intensified.

Alien plant invasions are a threat to this marsh plant, which even though at preliminary stage of nutrition research, is a validated critical component of wetland food chain. Dodder (Cuscuta salina), the Convolvulaceae creeper infestation on Salicornia has been observed. C. salina parasitizes S. virginica, eventually killing the latter (Pennings and Callaway <a href="#CR58" rid="CR58" class=" bibr popnode">1996</a>) (Fig. <a href="/pmc/articles/PMC4835422/figure/Fig1/" target="figure" class="fig-table-link figpopup" rid-figpopup="Fig1" rid-ob="ob-Fig1" co-legend-rid="lgnd_Fig1">1</a>b).

Conclusion

Salicornia is touted as a ‘secondary vegetable’, ‘famine food’ and ‘plant for future’. Despite multiple evidences of its health benefits it languishes as a mere marsh plant. As food insecurity looms large, such nutrition sources should not be wasted. Further, saline habitats have low agronomic relevance, so this halophyte can be cultivated to make better use of them. Further investigation in the line of the suggested area is expected to promote its popularity and provide an abundant source of nutrition in the times of ‘food insecurity’.

Compliance with ethical standards

Conflict of interest

The author declares there is no conflict of interest in submission of this manuscript.

References

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Articles from 3 Biotech are provided here courtesy of Springer
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Phytochemicals

Salicornia plants have been screened for phytochemical profile and presence of a range of carbohydrates, proteins, oils, phenolic compounds, flavonoids, sterols, saponins, alkaloids, and tannins have been reported. Water and alcohol extraction followed by component profiling has indexed many potentially bioactive compounds. Studies have reported the presence of dietary fibers, bioactive polysaccharides, proteins, lipids, sterols, flavonoids, and minerals (Mg, Ca, Fe, K) in S. herbacea (Essaidi et al. 2013). An immunomodulatory polysaccharide was isolated from S. herbacea (Lee et al. 2006; Im et al. 2006). Monosaccharide composition analysis of Salicornia brachiata fraction revealed the dominance of rhamnose, arabinose, mannose, galactose, and glucose, with meager presence of ribose and xylose (Sanandiya and Siddhanta 2014). Proteomic analysis (matrix-assisted laser desorption/ionization-time of flight (MALDI-TOF) mass spectrometry) revealed high protein contents of the seeds. Based on the detection of high disulfide linkages in the globulin proteins, it was deemed suitable for consumption, as sulfur-rich proteins are generally fit for nutrition (Jha et al. 2012). Gas chromatography mass spectrometry (GC–MS) was used to profile lipidome of S. ramosissima which showed esterified and free fatty acids, fatty alcohol, sterols, alkanes and aromatic acid derivatives. Among the dominant components, palmitic acid, tetracosanol and octacosanol were significant (Isca et al. 2014). Tetracosanol, the aliphatic alcohol has been identified to possess α-amylase ability, which makes it relevant in diabetes therapy (Jhong et al. 2009). Octacosanol, the high-molecular-weight aliphatic alcohol has been known to be a component of cholesterol-lowering drugs such as policosanol (Liu et al. 2015). By metabolomic analysis, S. brachiata was detected to be rich in sulfur amino acids and polyunsaturated fatty acids (PUFAs 55–64 %) (Mishra et al. 2015). Presence of selenium in S. brachiata was detected (Mishra et al. 2015). Selenium is an essential micronutrient for growth and robust antioxidant effects, deficiency of which has been documented to impair the immune system (Finley 2005). In this regard, it appears justified to evaluate dietary extraction of selenium from Salicornia. A study found S. herbacea seed oil to be stable to oxidation and eligible to be used in food processing (Choi et al. 2014). The oil composed of linoleic acid, oleic acid, arachidic acid, palmitic acid, tocopherol (α, γ, δ type), phenol, chlorophyll, and β carotene, was free of rancidity during a 60 day dark storage period (Choi et al. 2014). Stigmastanol, 24-ethyl-δ(22)-coprostenol and several other bioactive fatty alcohols were detected (Isca et al. 2014). In another study, a chlorogenic acid, 3-caffeoyl-4-dicaffeoylquinic acid was detected in S. herbacea extract (Hwang et al. 2010). Another study identified pentadecyl ferulate, stigmasterol, ergosterol, vanillic aldehyde and scopoletin in S. herbacea (Wang et al. 2013). Chromatography could detect β sitosterol (1), stigmasterol (2), uracil (3), and isorhamnetin-3-O-β-D-glucopyranoside in S. herbacea (Lee et al. 2004). Saponins were identified in butanol fraction of S. herbacea extract, some of which exerted antioxidant effect (Kim et al. 2012). Two new noroleanane-type triterpene saponins, Salbige A and B were isolated from the aerial parts of S. herbacea (Zhao et al. 2014). A new nortriterpene saponin, bigelovii D with antifungal potential was isolated from the hydroalcoholic extract of S. bigelovii (Shan et al. 2015).These studies contributed towards phytochemical composition of this genus and emphasized the significant role of non-targeted metabolomics in further component analysis.

Health Benefits and Uses of Salicornia

  • Skin rejuvenating : An Salicornia herbacea (SH) treatment to B16 melanoma cells decreased the synthesis of melanin and inhibited tyrosinase activity. These results collectively indicate that SH had antioxidative and whitening effects on skin and would be a good candidate for skin rejuvenating agent.
  • Antioxidant : Aqueous and methanol extracts of the enzyme-treated S. herbacea possessed potential antioxidant effects as observed in vitro on rat liver microsomal lipid peroxidation (Jang et al. 2007). The butanol fraction S. herbacea methanol extract exerted scavenging activities attributable to its active principle isorhamnetin 3-O-β-d-glucopyranoside. The glucopyranoside intervened with inflammatory pathways via manipulation of cytokine profile (Kim et al. 2009). In a rat model, S. herbacea vinegar when orally administered (at 7 g/kg water) showed free radical scavenging and SOD-like activities. Furthermore, the vinegar-administered rats showed higher glycogen accumulation in liver and muscles, higher lactate and ATP metabolism, promoting enzyme activities such as muscle creatine kinase and lactate dehydrogenase, whereas serum fatigue biomarkers such as ammonia, lactate and inorganic acid were markedly decreased (Cho et al. 2015).
  • Antiproliferative : Polysaccharides from botanical sources exerting anticancer properties have been well-documented (Chang 2002; Patel and Goyal 2012). In this regard, many Salicornia polysaccharides have also shown promise, validated through in vitro and in vivo models. Crude as well as purified polysaccharides from S. herbacea (at 0.5–4 mg/ml) demonstrated anti-proliferation of human colon cancer HT-29 cells when incubated for 24–48 h (Ryu et al. 2009). The mechanism of the cancer cell death was attributed to cell cycle arrest at G2/M phase, followed by apoptosis. Expression of the p53 tumor suppressor gene and the cyclin-dependent kinase inhibitor 1 (CDK inhibitor p21) were observed (Ryu et al. 2009). In another study S. herbacea -derived chlorogenic acid, 3-caffeoyl, 4-dicaffeoylquinic acid exerted control on metastasis of human fibrosarcoma HT-1080 cell line (Hwang et al. 2010). The invasion restraint was mediated through the inhibition of activator protein-1 AP-1 and signaling pathways involving protein kinase C (PKC) delta (repressing phosphorylation of ERK, p38 MAPK, and JNK) and three MAPKs, culminating in reduced activation of matrix metalloproteinase (MMP)-9 (Hwang et al. 2010). Another study found that pentadecyl ferulate from S. herbacea ethyl acetate extract possesses antioxidant effect and exerts anticancer response towards human hepatocellular liver carcinoma HepG2 and human lung adenocarcinoma epithelial A549 cells, along with phytol and γ-linolenic acid (Wang et al. 2013). Saponins, Salbige A and B, isolated from S. herbacea exerted antiproliferative activities towards A549 cells, while a pheophorbide (a chlorophyll catabolite) inhibited both A549 and HepG2 cancer cells (Zhao et al. 2014).
  • Lipid lowering : Hyperlipidemia is a major cause of morbidity and mortality across the globe. S. herbacea ethanol (50 %) extract exerted lipid lowering in mice model when prescribed for 10 weeks, via suppression of lipogenesis related genes [sterol regulatory element-binding protein 1 (SREBP1a), fatty acid synthase (FAS), and glycerol-3-phosphate acyltransferase (GAPT)] (Park et al. 2006). In addition, flavonoids from this plant were observed to exert adipogenic inhibition in 3T3-L1 adipocytes (Kong and Seo 2012). The isorhamnetin compound reduced adipogenic differentiation by down-regulation of peroxisome proliferator-activated receptor-γ (PPARγ), CCAAT/enhancer-binding proteins (C/EBPα), SREBP1, and the adipocyte-specific proteins. Involvement of AMP-activated protein kinase (AMPK) was also observed (Kong and Seo 2012). In addition, 3-caffeoyl, 4-dihydrocaffeoylquinic acid extracted from this species prevented lipid accumulation by blocking SREBP-1c and FAS through LKB1/SIRT1 and AMPK activation as studied in HepG2 cells (Pil Hwang et al. 2013). Another mice study on this plant revealed that it can decrease body weight gain by controlling serum leptin and manipulating lipid synthesis genes as SREBP-1c, PPARγ and FAS. Intake of high fat diet along with the plant powder at 50 g/ kg dose conferred significantly better parameters compared to only high fat diet or high fat diet plus NaCl group (Pichiah and Cha 2015).
  • Antibacterial : Methanol extract of S. herbacea showed antibacterial activities, mediated by interference with cytochrome P450 CYP1A2, CYP3A4 and CYP2D6 enzymes (Essaidi et al. 2013). Another study explored the possibility of developing antimicrobial nanoparticles from Salicornia. S. brachiata-based gold nanoparticles were analyzed through a set of standard tools, which revealed its poly-dispersed, crystalline nature and 22–35 nm size (Essaidi et al. 2013). The particles decimated tested bacteria, manifested in zone of clearance on inoculated plates. Further, the nanoparticles showed synergistic activity with fluoroquinolone antibiotic ofloxacin (Ayaz Ahmed et al. 2014). Based on these exciting results, follow-up studies ought to be pursued.
  • Antidiabetic : Diabetes has assumed epidemic proportions in current times, due to pollution, and processed, calorie-rich food consumption, among other causal factors (Olokoba et al. 2012). Existing panel of antidiabetic drugs provoke side effects (Haque et al. 2011; Stein et al. 2013). In this regard, safer options to control hyperglycemia are being searched. Consequently, the ameliorative effect of S. herbacea powder on induced-diabetic rats was studied. When administered alone or recommended with exercise (in the form of swimming), it showed higher expression of liver and muscle glucose transporters GLUT-4 and GLUT-2 (Lee et al. 2015). Higher glycogen concentration in liver and muscle also corroborated the finding, heralding possible usage of the halophyte as an antidote to control diabetes (Lee et al. 2015).
  • Hepatoprotective : Liver is a vital gland for proper functionality of the body (Jarrar et al. 2001). Most analgesics induce adverse effect on liver tissues and functions (Bessone 2010). To evaluate hepatoprotective effects of a Korean herbal drink, of which Salicornia was a constituent, this study was conducted. The multi-herb potion ‘taemyeongcheong’ was administered to acetaminophen-stressed mice. At 500 mg/kg dose, the drink conferred protective effects on mice liver. Drop in the level of oxidative enzymes as alanine aminotransferase (ALT), aspartate aminotransferase (AST), alkaline phosphatase (ALP), and lactate dehydrogenase (LDH), and elevation in the level of antioxidative enzymes as catalase, superoxide dismutase, glutathione peroxidase, and glutathione was observed. Decline in expression of hepatic mRNA levels of TNF-α, IL-1β, IL-6, COX-2, and iNOS genes were observed, which can be linked to the lower degree of lesions to liver tissue (Yi et al. 2015).
  • Immunomodulatory : Polysaccharides from this plant have shown evidence of eliciting immune response. In this context, some pertinent studies have been discussed below. In an in vitro study, S. herbacea polysaccharides induced nitric oxide (NO) production from mouse peritoneal macrophages and mouse leukaemic monocyte macrophage RAW 264.7, through the activation of nuclear factor-kappaB/Rel (NF-kappaB/Rel) (Lee et al. 2006). Consequently, the polysaccharide stimulating macrophages that express iNOS gene came forth (Lee et al. 2006). In another study on S. herbacea, its polysaccharide demonstrated effect on monocyte activation and differentiation into macrophage (Im et al. 2006). RAW 264.7 cells elaborated cytokines such as tumor necrosis factor (TNF)-alpha and interleukin (IL)-1 beta, and nitric oxide (NO) when incubated with the polysaccharide. Further differentiation into macrophage was determined from higher adherence development in the monocytes (Im et al. 2006). Enhanced collagen-adherence is known to improve phagocytosis (Newman and Tucci 1990). Further work by same group of researchers reaffirmed that the purified polysaccharide worked in sync with IFN-γ to induce immune effector molecules as TNF-α, IL-1 β, and NO to differentiate the monocytes into the macrophages (Im et al. 2007).
  • Osteoprotective : Bone health is crucial for they make the framework of body and enable proper muscle movement. Nutrient deficiency, genetic or geriatric conditions impose bone malfunctions. Osteoporosis is a key bone disorder, for which one causal agent has been identified as higher bone adipogenesis (differentiation of stem cells into mature adipocytes) (Pino et al. 2012). In this regard, S. herbacea extract was observed to inhibit adipogenesis via manipulation of PPARγ, CCAAT/enhancer-binding protein (C/EBP)α and SREBP1c. Resultantly, osteogenesis improved, as evidenced in MC3T3-E1 pre-osteoblasts. Osteoblastogenesis markers as alkaline phosphatase (ALP), bone morphogenetic protein (BMP)-2, osteocalcin and collagen type I (collagen-I) lent support to bone formation induction by S. herbacea extract (Karadeniz et al. 2014).
  • Antiseptic food additive : Sepsis, caused by the compromised integrity of membrane barrier can be fatal (Li et al. 2009), so the discovery of the antiseptic effect of Salicornia is interesting, regarding which some relevant findings have been discussed here. High mobility group box 1 protein (HMGB1), a nuclear protein elaborated by activated leucocytes, is released in excess when inflammatory tissue damage renders the membrane porous (Tang et al. 2010; Lotze and Tracey 2005; Passali et al. 2012). In this scenario, inhibitors of HMGB1 are suggested to be a potential treatment for sepsis. S. herbacea-derived caffeoylated quinic acids showed anti-HMGB1 activity which exerted protection towards vasculature (Tuan et al. 2015a). The purified flavanones and chromone derivatives from the plant suppressed the release of HMGB1 in mice models, barricading the animal intestine from septic shocks (Tuan et al. 2015b). Role of the phytochemicals in hyperpermeability modulation needs to be investigated further, to better utilize the halophyte in fight against septicemia. While the precise mechanism is yet to be discovered for Salicornia, other studies have attributed anti- HMGB1 activity to be due to activated cholinergic anti-inflammatory pathway (Goldstein et al. 2007).

References

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