Abstract
The increasing global population and progressive soil salinization threaten future food security and sustainable agriculture. Halophytes, as salt-tolerant plants adapted to saline environments, represent promising alternative crops and valuable sources of nutrients and bioactive compounds. This review presents a structured synthesis of selected halophytes, with emphasis on wild species of ethnobotanical relevance. The nutritional value of halophytes is discussed with respect to proteins, polysaccharides, lipids, minerals, and vitamins, together with their diverse profiles regarding bioactive compounds, such as polyphenols, terpenes and terpenoids (including carotenoids), alkaloids, saponins and chlorophylls. In addition, the biological activities and available clinical evidence of halophyte-derived compounds are summarized, with Lobularia maritima presented as a representative example. By organizing nutritional and phytochemical data according to compound classes, this review provides a perspective largely absent from previous studies and highlights the potential of halophytes as innovative ingredients for the food and pharmaceutical industries, as well as outlining future research challenges and prospects.
1. Introduction
Plant species differ markedly in their ability to cope with saline environments. While some conventional crops exhibit limited salt tolerance, halophytes—both wild and cultivated—are distinguished by their exceptional capacity to survive and develop under high salinity conditions. This unique adaptation makes halophytes particularly attractive as alternative crops for salt-affected areas, as well as promising reservoirs of nutrients and secondary metabolites with potential applications in the food and pharmaceutical industries. Many halophyte species synthesize a wide array of bioactive compounds of economic and therapeutic relevance; however, they also tend to accumulate elevated levels of sodium and chloride ions. Despite growing interest in these plants, the mechanisms by which salinity regulates the balance between beneficial metabolites and non-nutritional or potentially undesirable compounds remain insufficiently understood, largely due to strong species-specific and environment-dependent variability [1,2].
Global food systems are facing unprecedented pressure as the world population is expected to surpass 9.5 billion by 2050 [3]. Sustainable crop production and food security under these conditions is increasingly challenged by climate change, extreme weather events, and progressive land degradation. Among these factors, soil salinization represents one of the most severe constraints on agricultural productivity, as most staple crops are highly sensitive to elevated salt concentrations and cannot tolerate substantial salinity stress [4]. Consequently, the expansion of saline soils poses a direct threat to global food security.
Salinity adversely affects plant growth by disrupting water uptake, altering ionic homeostasis, and inducing oxidative stress at the cellular level. To persist under such hostile conditions, plants must activate complex defense strategies involving coordinated morphological, physiological, and biochemical adjustments. Halophytes, which are naturally adapted to saline habitats, have evolved highly efficient mechanisms to mitigate salt-induced damage, including ion compartmentalization, osmotic adjustment, and enhanced antioxidant capacity [5]. These integrated adaptations enable halophytes not only to survive but often to thrive in environments that severely impair the growth and productivity of glycophytic crops [6].
Beyond their ecological resilience, halophytes are increasingly recognized for their nutritional and phytochemical richness. They can serve as valuable sources of proteins, polysaccharides, lipids, minerals, vitamins, and a broad spectrum of bioactive compounds, including polyphenols, terpenes and terpenoids (notably carotenoids), alkaloids, saponins, chlorophylls and their derivatives, as well as organic sulfur compounds. Many of these metabolites exhibit antioxidant, anti-inflammatory, and anticancer activities, highlighting the potential of halophytes as functional foods and as sources of bioactive ingredients for pharmaceutical applications. Nevertheless, the relationship between salinity intensity and the accumulation of these beneficial compounds remains complex and incompletely characterized.
This review provides a comprehensive synthesis of current knowledge on the nutritional composition and bioactive potential of selected halophytes, with particular emphasis on wild species of ethnobotanical relevance. Nutritional and phytochemical data are systematically organized according to botanical families and classes of compounds, offering a perspective that is largely absent from previous reviews. In addition, available biological and clinical evidence supporting the health-promoting properties of halophyte-derived compounds is summarized, with Lobularia maritima presented as a representative example. By linking salinity-driven adaptations to metabolite production, this article highlights both the opportunities and challenges associated with the utilization of halophytes as innovative resources for the food and pharmaceutical industries, while outlining key directions for future research in the context of increasing soil salinization and global food insecurity.
Research on halophytes integrates diverse methodological approaches, including phytochemical profiling, molecular biology techniques, controlled cultivation systems, plant breeding strategies, and emerging biotechnological tools. Molecular and genomic analyses have been increasingly applied to elucidate stress-response pathways and metabolite biosynthesis mechanisms, while plant breeding and controlled-environment agriculture aim to enhance productivity and compositional stability. In addition, advanced analytical techniques such as UPLC-MS/MS and metabolomics enable detailed characterization of bioactive compounds. Although a comprehensive methodological review is beyond the scope of this article, these approaches collectively contribute to the expanding knowledge base on halophyte biology and utilization.
The literature analyzed in this review was collected from major scientific databases, including Scopus, Web of Science, PubMed, and Google Scholar. Particular emphasis was placed on publications from the last six years to ensure coverage of the most recent advances in halophyte research and technological developments; however, seminal earlier works authored by recognized experts in the field were also included due to their fundamental importance. The inclusion criteria comprised peer-reviewed original research articles and high-quality review papers, focusing on the production, characterization, biological activity, and application of nutrients and bioactive compounds derived from halophytes.
2. Nutrients from Halophytes
2.1. Proteins
The importance of proteins and amino acids in food, pharmacy and cosmetic industries is unquestionable. These macronutrients are one of the most vital elements of healthy diet, as they provide a source of carbon skeletons and energy, and are important constituents of formulated foods, contributing to features like texture, colour and flavour [7]. In cosmetics, proteins are useful for creating suitable environment for skin and hair, mostly owing to their binding properties [8]. Nowadays, we observe an increasing interest and need for higher availability of properly balanced plant-based proteins, as studies suggest that their increased consumption may contribute to a wide range of positive health outcomes [9].
In this respect, extremophytes, with halophytes among them, may provide a novel source of plant-based proteins characterized by unique structural and functional features. Halophytes represent here a non-conventional protein source that can be produced on saline soils and with the use of brackish water [10,11,12]. Indeed, proteomic studies of halophytes (e.g., Schrenkiella parvula) identified salt-responsive protein folding and stress-related proteins [13]. Additionally, similarly to the case of extremophilic microorganisms, proteins isolated from extremophytic plants represent unique biomolecules that are capable of functioning under severe conditions, which can facilitate their exploitation in a range of industrial technological processes [14].
2.1.1. Protein Content, Profile and Digestibility
Halophytes are generally rich in proteins. In natural conditions, total protein content usually ranges between 10 and 30% of dry biomass, although species such as Salicornica fruticosa and Kosteletzkya virginica accumulate 27 and 32% of protein, respectively [15,16,17]. In halophytes, the metabolism of seed storage proteins (SSP) is affected by various salinity levels. As in the example of Salicornia brachiata, the main fractions of SSP include globulins (over 54%), albumins (34%), glutelins (9%) and prolamins (2%) [18]. Studies of Cakile maritima revealed that although salinity exposure causes a delay in SSP degradation, it constitutes a way of stress priming, boosting vegetative growth after germination [19,20]. Interestingly, when plants grow without being exposed to salinity, the protein level may even reach 45%, as in the case of Halimione portulacoides [21].
The profile of proteins and amino acids may significantly differ between species. It also changes under the influence of environmental conditions, particularly salinity and drought. Stress-exposed plants accumulate higher contents of free amino acids that serve mainly as osmo-protectants [22]. Halophyte proteins are usually low in sulfur-containing amino acids, such as cysteine and methionine, whereas they are rich in leucine, isoleucine, lysine and valine, showing a potential for balancing exogenous amino acid levels in the human diet [12,21]. It has been reported that elevated contents of branched amino acids may also play a role in regulating glucose metabolism in humans [12,23]. The amino acid profile is also reported to be tissue- and organ specific. In the leaves of several halophyte species, increased alanine accumulation was observed compared with the seeds, compensating for the reduced proline content [22].
Considering digestibility of halophyte proteins, this is rather moderate [12,24]. In addition, high salinity conditions reduce protein digestibility in comparison with low or moderate salinity levels [25]. Recent reports suggest that digestibility can be improved by processing, such as blanching, fermentation or enzymatic hydrolysis, eliminating the impact of antinutritive compounds such as oxalates and phenolics [12,26]. Inevitably, the proteomic structure and rearrangements are crucial elements of halophyte adaptations and therefore their unique nature. Most proteins differentially regulated in halophytes are those responsible for basic cellular and growth processes. Comprehensive overview of these functional groups of proteins has been provided by [27], with glycophyte- and halophyte-specific examples of differentially regulated proteins involved in signalling (including phytohormonal pathways), regulation of gene expression, protein biosynthesis, folding and degradation, ion transport, photosynthesis and respiratory pathway, defence responses (including redox metabolism, osmolyte biosynthesis, metal binding), and numerous structural proteins, modulating the structures of cell wall and cell membranes [27]. Therefore, in this review we focus on halophyte proteins with particular characteristics, allowing for their exploitation in food and pharmaceutical production, such as halophytic antioxidant enzymes, stress-protective proteins and stress-associated transcription factors, as well as antimicrobial defensins and thionins. Other important groups of halophyte proteins are briefly characterized in Table 1.
Table 1.
Functional groups of halophyte proteins involved in cellular and growth processes.
2.1.2. Bioactive and Functional Proteins
Apart from nutrition, halophyte proteins may exert functional benefits, as these compounds are often stress-associated and stress-adapted. In particular, halophytic antioxidant enzymes, such as superoxide dismutase (SOD), peroxidases (POD), and catalase (CAT), exhibit elevated basal activity and salt-stable isoforms [33,34]. For example, boosted activity of antioxidant enzymes was reported in halophyte grass Aeluropus littoralis [35] and in sea aster (Aster tripolium) [36]. Salinity-induced isoforms were detected in extreme halophyte Salvadora perisca, revealing the constitutive role of Mn-SOD 1 under various salinity levels [37]. Such salt-tolerant antioxidant enzymes could be utilized as antioxidant agents or biocatalysts for oxidative damage mitigation. Another example is esterase, whose isoform predominating under salinity conditions exhibited elevated activity in three halophyte species from the Amaranthaceae family: Arthrocnemum macrostachyum, Sarcocornia fruticosa and Salicornia europaea [38].
Another group of halophyte proteome constitute stress-protective proteins, such as chaperones and heat shock proteins (HSPs), as well as transcription factors and osmo-protective Late Embryogenesis Abundant (LEA) proteins. Analyses of Koyro et al. [39] revealed that in Aster tripolium a 12-fold increase in abundance of HSP-20 occurred under salinity, whereas HSP-70 was a predominant protein in Suaeda salsa and Porteresia coartata. Chaperonins responsible for proper protein folding were identified also in Cakile maritima [19] and Salicornia brachiata [40]. The latter species was also a source of transgene SbASR-1 encoding LEA protein, enhancing salinity and drought tolerance when overexpressed in groundnut [41]. Dehydrins, a family group II of LEA proteins, characterized by a highly hydrophilic amino acid sequence, were identified in numerous halophytes, including Eutrema salsugenium, Cakile maritima, Chenopodium quinoa, Zostera marina, Phoenix dactylifera, Carex littledalei, Asparagus officinalis, and Lactuca saligna, while in three halophytic species (Puccinillia tenuiflora, Eutrema salsugenium, and Hordeum marinum) their expression was induced by salinity conditions and the response depended on dehydrin architecture [42].
Another group of functional proteins represent Stress-Associated Proteins (SAPs), transcription factors involved in the responses to environmental conditions. Their expression is related to enhanced tolerance towards drought, cold, heavy metals, salinity and biotic stressed [43]. The SAP genes and their encoding proteins have been identified in several halophyte species, including Lobularia maritima, Tamarix hispida, Suaeda glauca, and Cynodon dactylon [44,45,46,47].
As LEA proteins and chaperones are known for stabilizing cellular structures and enzymes under stress, they could be broadly exploited in the food, pharmaceutical and cosmetic industries as protein stabilizers. Isolated SAP proteins from Lobularia maritima have already been tested as a functional preservative from Listeria monocytogenes inoculated in beef meat. In the cases of several strains of human pathogenic bacteria, like Escherichia coli, Staphylococcus epidermis and Pseudomonas aeruginosa, direct application of purified LmSAP protein resulted in more pronounced pathogen inhibition than the use of antibiotics [48].
2.1.3. Antimicrobial Proteins and Peptides
Defensins, thionins, and lipid-transfer proteins (LTPs) belong to the large family of antimicrobial peptides (AMPs) synthesized by a wide variety of organisms, including plants. These compounds are an element of the plant defence system, capable of inactivating bacterial and fungal pathogens due to the interaction with their cellular membranes and alterations in their permeability [49]. Apart from direct antimicrobial properties, AMPs may act as immunomodulators and are involved in regulation of plant ontogeny and stress tolerance [50]. Although studies concerning the role of specific AMPs in halophytes are scarce, their abundance and stress-regulated expression have been confirmed by transcriptome profiling in Suaeda fruticosa [51], Hordeum marinum subsp. marinum [52], and Eutrema salsugineum [53]. There is a prospect that the halophyte-derived AMPs may remain active under high ionic strength, which would be a key advantage over conventional antimicrobial peptides. They could be used as topical antimicrobials and food-grade preservatives that can be processed in more severe environmental regimes.
In summary, it should be highlighted that halophyte-derived proteins combine stress stability with biological activity, making them promising but underexplored candidates for exploitation as a sustainable source of nutritional protein, as well as for development of novel nutraceuticals and pharmaceuticals. The main application constraints to be solved include limited purification and structural characterization of halophyte proteins, scarcity of in vivo pharmacological and toxicological studies, and the need for scalable and reliable recombinant production systems.
2.2. Polysaccharides
Polysaccharides are high-molecular-weight carbohydrate polymers and represent one of the earliest naturally occurring biopolymers. They are ubiquitously present across biological systems, including plants, animals, and microorganisms, and constitute a major class of biologically relevant macromolecules [54,55]. In glycophytes, or salt-sensitive plants, polysaccharides play a limited role in stress-response mechanisms, whereas in halophytes they contribute significantly to adaptation under saline conditions [56]. Due to their unique physicochemical characteristics and broad spectrum of biological functions, including immunomodulatory, antioxidant, and antitumor activities, plant-derived polysaccharides and polysaccharide–protein conjugates have gained increasing attention in recent years [57,58]. Moreover, their relatively low toxicity makes them promising candidates for therapeutic applications, as well as potential alternatives to conventional antioxidants for mitigating oxidative stress [55,59].
In halophytes, bioactive polysaccharides can be classified into several major groups based on their biological activity and industrial relevance (Table 2). These include cell wall-associated polysaccharides, such as pectins and hemicelluloses, storage polysaccharides, and water-soluble mucilages and gums. Pectic polysaccharides, particularly homogalacturonans and rhamnogalacturonans, exhibit antioxidant, immunomodulatory, and metal-chelating properties, which are closely linked to the adaptive strategies of halophytes under saline conditions. Hemicelluloses, including arabinoxylans and mannans, contribute to dietary fiber content and display prebiotic and anti-inflammatory effects. Storage polysaccharides, such as fructans and β-glucans, play a key role in osmotic adjustment and stress tolerance, while also providing well-documented health benefits, including modulation of gut microbiota and immune responses. Additionally, mucilages and plant gums produced by many halophytes exhibit gastroprotective, emulsifying, and stabilizing properties, making them valuable ingredients for pharmaceutical formulations and functional foods. Together, these polysaccharides represent a promising group of natural compounds with multifunctional bioactivity and significant potential for applications in the food and pharmaceutical industries [60,61,62].
Table 2.
Polysaccharides from selected halophytes—types, activities, and potential applications.
Studies on polysaccharides from edible halophytes found that most extracted carbohydrates are pectic polysaccharides, rich in uronic acids (18.9–90.1%), arabinose, galactose and other neutral sugars, with molecular weights from 10 to 2650 kDa. These pectins often show immunomodulatory activity, and structure–activity analyses highlight the roles played by uronic acids, sulfate groups, low methylation, and molecular weight [63]. The recent study on Limonium gmelini separated polysaccharide-rich aqueous fractions from polyphenol-rich ones; the polysaccharide fractions showed minimal antibacterial activity, whereas polyphenol fractions were strongly antibacterial [66]. This suggests that Limonium pectins may be more relevant for other bioactivities (e.g., immunomodulation) than for direct antimicrobial effects, but they remain poorly characterized.
Wang et al. [61] isolated and characterized a bioactive polysaccharide, SabPS-1, from the halophyte Salicornia bigelovii, focusing on its structural features and antitumor activity. SabPS-1 exhibited a molecular weight of 3.24 × 104 Da and was rich in arabinose, galactose, and uronic acids, with a backbone mainly composed of β-D-galacturonic acid, α-L-arabino-furanose, and β-D-galactose residues, along with arabinose-containing side chains. In vitro assays demonstrated that SabPS-1 significantly inhibited HepG2 hepatocellular carcinoma cell proliferation and induced apoptosis, which was associated with the activation of caspase-3 and caspase-8 and the regulation of apoptosis-related proteins Bax and Bcl-2. This study provided one of the first detailed links between the structural characteristics of polysaccharides from S. bigelovii and their antitumor bioactivity, highlighting their potential as natural candidates for anticancer drug development.
The analysis of the monosaccharide composition in Tunisian halophyte Salicornia arabica revealed the presence of arabinose, glucopyranose galactose, glucuronic acid and glucose. The identified polysaccharides displayed an important radical scavenging and reducing power as well as exhibiting effective antimicrobial properties [68].
A new heteropolysaccharide (LmPS) was isolated from Lobularia maritima (a halophyte collected in Tunisia) and showed strong antibacterial activity against foodborne pathogens, including Staphylococcus aureus, Listeria monocytogenes, Salmonella enterica, and Escherichia coli. Additionally, LmPS improved the shelf life of raw minced meat by reducing lipid oxidation and metmyoglobin accumulation, indicating its potential as a natural antioxidant additive in the food industry [69].
Polysaccharides from Carpobrotus edulis (CEP) were successfully optimized and characterized as low-methoxyl pectic polymers rich in galacturonic acid, with a semi-rigid structure and an average molecular weight of ~65 kDa. CEP exhibited strong antioxidant and antiglycation activities, highlighting its potential as functional bioactive compound for applications in food preservation as well as in the pharmaceutical and medical industries [60].
In some coastal halophytes, such as Ruppia maritima, Halodule wrightii and a mangrove Avicennia shaueriana, the presence of sulfated polysaccharides has been reported [56]. These cell wall structural compounds are widespread among various taxa of marine algae, suggesting that they are crucial elements allowing undisturbed growth and development in the marine environment. Comparative analyses conducted by Aquino et al. [56] revealed that in response to high salinity sulphated polysaccharides are also synthesized and incorporated into cell walls in the halophyte species of higher plants, while they are absent in representative glycophyte species. Such data suggest that incorporation of negatively charged polysaccharides into cell walls is an adaptation to high salt environments [56]. In this respect, coastal halophytes and mangroves could serve as alternative or complementary raw materials to algae, thereby reducing dependence on marine harvesting and seasonal variability. As accumulation of sulphated polysaccharides seems to be environmentally regulated, it may also become possible to achieve controlled modulation of their levels in selected species.
Overall, polysaccharides derived from halophytes represent a highly promising class of bioactive compounds for pharmaceutical and food applications. Their distinctive structural characteristics—particularly the abundance of pectic domains, arabinose- and uronic acid–rich fractions, and frequent sulfation—are closely associated with a wide range of beneficial biological effects, including immunomodulatory, antioxidant, and antitumor activities. While the structure–activity relationship is increasingly well established, current evidence is largely limited to preclinical studies, highlighting the need for rigorous clinical validation and comprehensive safety evaluations. Beyond therapeutic potential, halophyte polysaccharides also offer valuable functional properties for the food industry and emerging applications as bio-stimulants to enhance crop tolerance to salinity. Addressing existing knowledge gaps related to mechanisms of action, standardization of extraction protocols, and long-term safety will be essential for their successful translation into commercial pharmaceutical and food products.
2.3. Storage Lipids
Halophytes undergo lipid remodeling to preserve membrane stability and sustain the efficiency of essential metabolic processes. As a physiological consequence of stress exposure, these plants often accumulate storage lipids, predominantly neutral lipids in the form of triacylglycerols (TAGs), in both seeds and highly productive photosynthetic organs. Under saline conditions, halophytes can maintain productivity while increasing TAG content in their leaves, frequently exhibiting elevated levels of saturated fatty acids. At the same time, storage lipids from halophytes are often enriched in polyunsaturated fatty acids (PUFAs), including linoleic and α-linolenic acids, which contribute to membrane fluidity during germination and display well-documented health-promoting properties such as anti-inflammatory and cardioprotective effects. In addition to TAG-bound fatty acids, halophyte storage lipid fractions commonly contain lipid-soluble bioactive compounds, including tocopherols, phytosterols, and minor isoprenoid derivatives, which enhance oxidative stability and nutritional value. Notably, some halophyte species maintain high concentrations of essential omega-3 and omega-6 fatty acids even under seawater irrigation. Collectively, these features position halophytes as promising renewable sources of added-value fatty acids and storage lipids, with significant potential for applications in food production, nutraceuticals, pharmaceuticals, and biotechnology. Furthermore, the unique fatty acid composition and bioactive lipid profiles observed in halophyte seeds make these plants prospective sustainable sources of functional lipids for food, dietary supplements, and pharmaceutical formulations, complementing conventional plant oil crops [70,71].
Seeds of halophytes such as Salicornia bigelovii have been documented to contain high seed oil content (approximately 26–33% total lipids) with a fatty acid profile rich in linoleic acid (C18:2 n-6), oleic acid (C18:1), and other unsaturated fatty acids typical of edible oils, making them comparable to conventional oilseed crops for human nutrition and functional food uses [72]. Similar profiles are reported in other halophytic species, in which TAGs often dominate the seed lipid fraction alongside other minor bioactive components such as phytosterols and tocopherol [73]. Lipidomic analyses have proven to be a valuable tool for assessing the biotechnological and economic potential of halophytes. Custódio et al. [74] demonstrated that the edible leaves of Halimione portulacoides constitute a rich source of functional lipids with nutraceutical and pharmaceutical relevance, while also exhibiting a high degree of lipidome stability under variable nutrient conditions. Using high-resolution mass spectrometry–based lipidomics, the authors identified 175 lipid species, predominantly phospholipids and glycolipids, highlighting the structural and metabolic diversity of halophyte lipids. Importantly, plants cultivated under saline conditions with dissolved inorganic nitrogen and phosphorus concentrations typical of aquaculture effluents maintained a relatively stable lipid profile, whereas nutrient limitation led to a reduction in total phospholipid and glycolipid content and significant alterations in glycolipid composition. These findings underscore the relevance of nutritional regime as a key factor influencing halophyte lipid composition and emphasize the need to define nutrient thresholds to ensure consistent lipid profiles for food, nutraceutical, and biotechnological applications.
Lipidomic profiling conducted by Pan et al. [75] revealed pronounced salt-induced lipidome remodeling in Paspalum vaginatum, particularly in root tissues of salt-tolerant accessions. Under high salinity, enhanced lipid metabolism was observed in roots of salt-tolerant plants, indicating a tissue-specific lipidomic response associated with improved salinity tolerance. These lipidome adaptations suggest an important role of membrane lipid remodeling in maintaining root function under saline stress and highlight lipids as key contributors to salt tolerance mechanisms in halophytic grasses.
Roshandel and Shamsi [76] evaluated the oil yield, fatty acid composition, and physicochemical properties of seeds from several halophytic species, including Atriplex griffithii, Haloxylon ammodendron, Salicornia europaea, and Salsola yazdiana, to assess their suitability as alternative oilseed crops under saline conditions. The studied species exhibited moderate to high total oil contents and a predominance of unsaturated fatty acids, with particularly high proportions of linoleic, α-linolenic, and oleic acids. Among the analyzed halophytes, Salicornia europaea was identified as the most promising source of edible oil due to its high level of unsaturated fatty acids and the absence of erucic acid, resulting in oil quality comparable to that of conventional edible oils. Haloxylon ammodendron also showed strong potential, combining relatively high oil yield with favorable fatty acid composition. Overall, these findings underscore halophytic oilseeds as valuable and sustainable sources of edible oils, capable of cultivation in saline soils or under seawater irrigation, thus providing a practical complement to conventional oil crops. The results reported by Agudelo et al. [15] demonstrated that linoleic, α-linolenic, and arachidic acids were the most abundant fatty acids in three halophytic species, Atriplex halimus, Salicornia fruticosa, and Cakile maritima. These fatty acids belong primarily to the omega-6 and omega-3 series, which are of particular importance in human nutrition. Consistent with findings from other studies, the lipid composition of these halophytes suggests a favorable nutritional profile associated with health-promoting properties. Notably, the elevated lipid content observed in Cakile maritima, even under high salinity conditions, represents an added nutritional value, as its predominant lipid fractions exhibit bioactive properties.
Ben et al. [77] reported a pronounced increase in sulfo-lipid content in the halophyte Crithmum maritimum under saline conditions, indicating a salt-induced remodeling of sulfur-containing membrane lipids. Although the overall fatty acid composition of sulfo-lipids remained largely unchanged, salinity exposure led to specific shifts in polyunsaturated fatty acids, characterized by a decrease in linolenic acid (C18:3) and a concomitant increase in linoleic acid (C18:2). These compositional adjustments suggest a role for sulfo-lipids in modulating membrane properties under salt stress and support the involvement of sulfur-containing lipids as part of the physiological strategies contributing to salt tolerance in halophytes. For clarity and comparison, Table 3 summarizes the functional classification of storage lipids derived from several halophytes.
Table 3.
Functional classification of storage lipids derived from halophytes.
Collectively, the reviewed studies demonstrate that halophytes can synthesize a wide spectrum of storage and membrane lipids with high nutritional and biotechnological value, even under severe salinity and nutrient stress. Lipidomic analyses reveal that halophytic tissues and seeds are rich in triacylglycerols, phospholipids, glycolipids, and sulfur-containing lipids, often characterized by a high proportion of nutritionally relevant omega-3 and omega-6 fatty acids, including linoleic and α-linolenic acids. Several halophyte species produce seed oils with fatty acid profiles and physicochemical properties comparable to conventional edible oils, while others accumulate bioactive polar lipids and lipid-soluble antioxidants with recognized anti-inflammatory, cardioprotective, and nutraceutical potential. Moreover, salt-induced lipid remodeling, particularly in membrane and sulfur-containing lipids, appears to contribute to stress tolerance while simultaneously enhancing the functional value of these lipid fractions. This unique combination of environmental resilience, lipidomic diversity, and health-promoting properties positions halophytes as promising and sustainable sources of functional lipids for food, nutraceutical, and pharmaceutical applications, as well as valuable complements to traditional oil crops in sustainable saline agriculture.
2.4. Minerals
Mineral elements are essential for proper plant growth and development, as well as for the health of animals and humans who consume them. These mineral nutrients are commonly classified into two main groups: macronutrients and micronutrients. Plants absorb mineral elements from the soil in the form of dissolved mineral salts through the root system, and their availability is influenced by numerous factors, including soil physicochemical properties such as pH, salinity, and interactions among individual elements [80]. Previous studies of halophytes of Chenopodiaceae have shown differentiated accumulation of mineral elements in plant organs. Higher concentrations of elements have been reported in leaves compared to stems and roots [81]. The mineral composition of numerous halophytic species has been extensively investigated, as documented in the literature [12,15,16,81,82,83]. Moreover, both wild halophytic plants and cultivated species constitute valuable sources of proteins, dietary fiber, and mineral elements, although it has been shown that halophytes are richer in nutrients [16].
Macronutrients are mineral elements required by plants in relatively large amounts to sustain growth, development, and physiological functions. Among the most important macronutrients, nitrogen (N) is a fundamental component of proteins and chlorophyll, making it essential for plant growth. Phosphorus (P) plays a critical role in energy transfer processes through adenosine triphosphate (ATP), contributing to root development and flowering. It has been found that increased salinity levels (200 mM) enhanced the accumulation of nitrogen, phosphorus, and sodium in Tetragonia decumbens [84]. Potassium (K) is involved in the regulation of plant water balance, the opening and closing of stomata, and the activation of numerous enzymes, whereas calcium (Ca), a structural component of cell walls, contributes to cell stability and regulates various cellular processes. Comparative analyses of halophytic species revealed that Inula crithmoides and Sarcocornia fruticosa are characterized by high concentrations of calcium and potassium, whereas Salicornia ramosissima contains elevated levels of magnesium, copper, and manganese [83]. Studies conducted on Mesembryanthemum crystallinum demonstrated that the highest concentrations of nitrogen (N) and magnesium (Mg) occurred in plants grown without saline treatment, whereas phosphorus (P), potassium (K), calcium (Ca) and sodium (Na) accumulated at higher levels under combined salinity and irrigation interval conditions. A statistically significant effect of this interaction was observed only for iron and calcium, indicating that M. crystallinum can mitigate the adverse effects of salt stress on mineral accumulation [85].
Micronutrients are mineral elements required by plants in much smaller trace amounts, but they are equally essential for proper physiological functioning. Iron (Fe) is essential for chlorophyll synthesis and cellular respiration, with its content increasing under salinity in species such as Atriplex halimus (at 300 mM NaCl) and Salicornia fruticosa (100 mM NaCl) [15]. Significant amounts of essential minerals and trace elements (especially calcium and iron) have been determined in Australian edible halophyte species, Atriplex sp., Suaeda sp., and Sesuvium sp. [12]. Manganese (Mn) activates numerous enzymes and contributes to photosynthesis, while boron (B) supports sugar transport, cell division, and reproductive organ development. Salicornia ramosissima has been identified as an excellent source of manganese (204 μg/g dry weight) [86]. In Salicornia fruticosa, the contents of Mo, B, and Mn increased in response to salinity. Zinc (Zn) functions as a component of enzymes and a plant growth regulator; for example, Cakile maritima exhibits high Zn concentrations that are ten times higher than in Atriplex halimus, although salinity had no effect on the Zn content in other species studied [15]. The use of halophytes as edible plants requires careful consideration, as their ability to survive under extreme conditions also enables the accumulation of potentially harmful heavy metals. Several species within the genus Salicornia are capable of significant heavy metal uptake, including arsenic (As), cadmium (Cd), copper (Cu), lead (Pb), and mercury (Hg) [87]. Similar properties have been reported for Salicornia ramosissima [88] Portulaca oleracea [89,90] and Aster tripolium [91].
Halophytes are plants adapted to saline environments, capable of accumulating various mineral salts as part of their survival strategy. The primary salts stored by halophytes include chlorides, sulfates, and carbonates of sodium, magnesium, and calcium (for example, in Salicornia brachiata leaves Na+ reached 7.6 meq g−1, whereas Cl− was noted to be 4.34 meq g−1) [92]. Sodium chloride (NaCl) is the most common salt, often referred to as rock salt or halite in its mineral form, and represents the main contributor to soil salinity that halophytes must tolerate [93]. For instance, leaves of Atriplex halimus accumulate relatively high amounts of Na+, although the levels remain lower than those found in other culinary halophyte species, making them suitable for human consumption [15]. Similarly, Salicornia fruticosa has been reported to contain 1120 mg Na per 100 g fresh weight. Halophytes generally exhibit higher ash content and elevated NaCl concentrations compared to glycophytes, with Salicornia ramosissima (1.50–2.39 g/100 g FW) and S. fruticosa (2.80–4.84 g/100 g FW) showing the highest levels of NaCl and total chlorides [83]. These properties suggest that halophytes could serve as sources of herbal salts or salt extracts, potentially substituting conventional NaCl in food applications. Halophytes also accumulate sulfate, as well as sodium carbonate, which occurs in smaller amounts and is still absorbed from saline soils. A unique feature of halophytes is their ability to store these salts in specialized vacuoles or bladder cells and, in some species, to excrete excess salts onto the leaf surface in the form of crystals. This unusual mechanism allows halophytes to thrive in environments inhospitable to most salt-intolerant plants (glycophytes) [94,95].
2.5. Vitamins
Across species, halophytes are consistently reported as valuable sources of several vitamins and related bioactive compounds, reflecting their metabolic adaptations to saline environments. These plants frequently accumulate antioxidant compounds, including both fat- and water-soluble vitamins, which contribute to their nutritional value and potential use as functional foods [15,96]. Various methods of extracting vitamins from halophyte biomass have been described [97].
Fat-soluble vitamins reported in halophytes include vitamins A and E, whereas data on vitamins D and K remain scarce or entirely absent (Table 4). No quantitative information on vitamin D (calciferol) content in halophytic species is currently available. This is not unexpected, as plants in general are not considered significant dietary sources of vitamin D, which is predominantly obtained from animal-derived foods and endogenous synthesis in humans following exposure to UVB radiation [98]. In contrast, halophytes can provide appreciable amounts of vitamin A–related compounds. Notable concentrations of retinyl acetate have been reported in Mesembryanthemum nodiflorum and Suaeda maritima, reaching 4.51 mg/100 g dry matter (DM) and 5.39 mg/100 g DM, respectively [16]. These values indicate that certain halophytic species may represent relevant plant-based sources of provitamin A activity. Vitamin E, mainly in the form of tocopherols, has been identified in several halophytic taxa, including Arthrocnemum, Halocnemum, Suaeda, and Sarcocornia species, contributing to their antioxidant potential, although quantitative data vary substantially among species [16,86,99].
Water-soluble vitamins are well-documented in halophytes, primarily consisting of vitamin C and selected B-group vitamins (Table 4). Many edible halophytes exhibit high ascorbic acid content, often at levels comparable to or exceeding those found in conventional leafy vegetables. For example, Sesuvium spp. exhibited exceptionally high vitamin C content, reaching 157 mg/100 g dry weight (DW) [12], while Cakile maritima subsp. maritima exhibited 135.3 mg/100 g fresh weight (FW) [100]. Such levels highlight the nutritional relevance of halophytes as sources of dietary antioxidants.
B-group vitamins are present in halophytes, although their distribution and concentrations vary widely among species and remain insufficiently characterized. Thiamine (B1) and pyridoxine (B6) have been quantified in several edible halophytes, including Inula crithmoides, Mesembryanthemum nodiflorum, Salicornia ramosissima, Sarcocornia fruticosa and multiple Suaeda species [16,101,102,103,104,105]. Riboflavin (B2) data are currently limited to Suaeda arcuata. This has been reported to contain quantified levels of riboflavin, ranging from approximately 0.053 to 0.062% of dry mass, indicating a relatively high B2 content [104,105]. Quantitative information regarding B2 for other halophytes is lacking, which underscores a significant gap in the current understanding of B vitamin profiles in halophytes. Folate (vitamin B9) appears to be one of the better characterized B vitamins in halophytes, especially for various species of the genera Suaeda and Atriplex [101,105]. Folate content was reported in Atriplex spp., reaching 303 µg/100 g DW [12]. In contrast, niacin (B3), pantothenic acid (B5), and biotin (B7) have not been quantitatively assessed in edible halophytes, representing notable analytical gaps. Interestingly, cobalamin (vitamin B12), which is generally absent from higher plants, has been reported in Suaeda aegyptiaca and Suaeda vera, although its origin and bioavailability require further investigation [101].
Table 4.
Vitamins reported in halophytic plant species used as food or investigated for their nutritional value.
3. Extraction, Separation, and Detection Technologies of Nutrients and Bioactive Components in Halophytes
The efficient valorization of halophytes requires appropriate extraction, separation, and analytical strategies to recover and characterize their diverse nutrients and bioactive compounds, including phenolics, flavonoids, glucosinolates, terpenoids, alkaloids, and polysaccharides. Traditional extraction techniques such as maceration and Soxhlet extraction using organic solvents (e.g., methanol, ethanol, hexane) remain widely employed due to their simplicity and reproducibility [106,107]. However, these methods are often time-consuming, require large solvent volumes, and may lead to degradation of thermolabile constituents [106]. Enzymatic hydrolysis has been applied to enhance the release of bound phenolic compounds and other bioactives from plant cell walls under mild conditions, thereby improving extraction yield and selectivity [108].
In recent years, more modern and greener technologies have gained increasing attention. Ultrasound-assisted extraction (UAE) enhances mass transfer through cavitation effects, significantly reducing extraction time and solvent consumption while improving recovery efficiency [109]. Supercritical fluid extraction (SFE), particularly using CO2, represents a selective and environmentally friendly alternative for isolating non-polar and moderately polar compounds, producing high-purity extracts with minimal solvent residues [110]. For compound separation and identification, advanced analytical platforms such as HPLC-DAD, UPLC-MS/MS, and GC-MS are routinely employed for high-resolution profiling, structural elucidation, and sensitive quantification of phytochemicals in complex plant matrices [111]. Compared with conventional analytical approaches, these modern techniques provide superior sensitivity, selectivity, and accuracy, facilitating a deeper understanding of the nutritional and pharmacological potential of halophytes.
4. Bioactive Components in Halophytes
4.1. Polyphenols
Polyphenols represent a broad group of secondary metabolites characterized by the presence of one or more phenolic rings. They are commonly classified into phenolic acids, flavonoids, stilbenes, and lignans, with flavonoids being the most abundant subgroup in higher plants. In plants, polyphenols fulfill crucial physiological and ecological roles, including antioxidant defense, UV protection, regulation of growth, and resistance to biotic stress, particularly pathogens and herbivores [112].
In halophytes, polyphenols are often synthesized at significantly higher levels than in glycophytes. Salinity stress induces excessive production of reactive oxygen species (ROS), and polyphenols act as efficient ROS scavengers, stabilizing cellular membranes and protecting photosynthetic tissues [113,114]. Enhanced accumulation of flavonoids and phenolic acids has been reported in halophytes such as Cakile maritima, Inula crithmoides and Dittrichia viscosa, particularly under saline conditions, indicating that polyphenol biosynthesis is part of a broader stress-adaptation strategy [113,115]. These compounds also participate in signaling pathways involved in stress perception and tolerance [116].
From a food and health perspective, polyphenols are highly valued for their antioxidant, anti-inflammatory, cardioprotective, and anticancer properties [112,117]. In the food industry, they contribute to sensory attributes such as color, bitterness, and astringency, while improving oxidative stability and shelf life [113]. The naturally elevated polyphenol content of halophytes, particularly under stress conditions, positions them as promising alternative sources of functional ingredients that can be produced on saline soils unsuitable for conventional agriculture.
4.2. Terpenes and Terpenoids
Terpenes and terpenoids, collectively referred to as isoprenoids, are synthesized from five-carbon isoprene units and represent the largest and most structurally diverse class of plant secondary metabolites, with terpenoids distinguished by the presence of oxygen-containing functional groups [118]. In plants, these compounds function as phytohormones, membrane components, signalling molecules, and defensive agents, as well as mediators of plant–environment interactions [119].
In halophytes, terpene and terpenoid biosynthesis is frequently enhanced under saline and osmotic stress. Salinity can activate isoprenoid pathways, leading to increased production of volatile terpenes and stress-related terpenoids that protect cellular membranes and photosynthetic structures from oxidative damage [116,120]. Additionally, some halophytes such as Artemisia caerulescens accumulate high levels of essential oils rich in monoterpenes and sesquiterpenes [121], which may serve both as antioxidants and as deterrents against herbivores in harsh saline habitats [118,122].
Terpenes and terpenoids are of major industrial and pharmaceutical importance, with applications spanning flavorings, fragrances, cosmetics, agrochemicals, and medicines [123]. Many exhibit antimicrobial, anti-inflammatory, anticancer, and neuroprotective activities [118]. The ability of halophytes to maintain or even enhance terpene production under salt stress highlights their potential as robust bioresources for terpene-rich extracts, particularly in the context of sustainable production systems.
4.3. Alkaloids
Halophytes are not traditionally recognized as primary sources of classical alkaloids such as morphine or quinine; however, numerous species synthesize characteristic groups of alkaloids with significant physiological functions. These plants have developed a range of physiological and morphological adaptations to survive under extreme environmental conditions, including high salinity, osmotic stress, and intense solar radiation. Among these adaptations, the increased synthesis and accumulation of secondary metabolites, particularly alkaloids, plays a crucial role [2]. Many studies have demonstrated that salt stress induces secondary metabolites, serving as a key mechanism that improves halophyte tolerance to adverse conditions [124,125].
Halophytes frequently contain alkaloids along with phenolics and flavonoids, making them a valuable subject of pharmacognostic and biotechnological research. Selected examples of halophyte-produced alkaloids are presented in Table 5. Numerous studies have focused on the preliminary detection of alkaloids using phytochemical screening assays without further structural elucidation [126,127,128,129]. Many of these studies refer to halophytic species characteristic of mangrove ecosystems, such as Avicennia alba [130], A. marina [126,128,129], A. officinalis [126,127], Rhizophora stylosa [128], Acanthus ebracteatus, Sonneratia alba, Aegiceras corniculatum, Ceriops decandra, Excoecaria agallocha, Lumnitzera racemosa, Rhizophora apiculata, and Rhizophora mucronata [127]. Based on such phytochemical screening tests, several species have been identified as particularly rich in alkaloids, including Acanthus ilicifolius and Heliotropium curassavicum [131], as well as Suaeda fruticosa [132]. These findings have repeatedly confirmed a high alkaloid content within these taxa. Other reports provide only the total number of alkaloids detected in a species, without detailed structural identification. For example, 15 alkaloids in Suaeda salsa and 14 alkaloids in Salicornia europaea were reported [133].
An additional phenomenon of interest is that alkaloids may not always be synthesized directly by the plant itself but rather by its endophytic microorganisms. A notable example is Halocnemum strobilaceum, whose endophyte produces the phenolic alkaloid halociline, exhibiting antioxidant, antimicrobial and antibiofilm activities [134]. In many halophytes, alkaloids exert a wide range of biological effects, including antimicrobial, antidiabetic, antioxidant, anti-inflammatory, and antitumor activities [135]. Their strong free radical scavenging capacity often correlate with high total phenolic content, contributing to cell homeostasis under stress conditions [136,137]. Consequently, halophyte-derived alkaloids are increasingly being investigated as potential bioactive agents for use in pharmaceuticals, nutraceuticals, and functional foods.
Although specific, structurally identified halophyte alkaloids are reported with increasing frequency, most available data are still limited. This indicates that halophyte-derived alkaloids represent promising, yet largely experimental therapeutic leads that require further pharmacological validation and comprehensive clinical investigation.
Table 5.
Selected alkaloids described for halophyte species.
4.4. Saponins
Saponins are amphiphilic glycosides composed of a hydrophobic sapogenin (triterpenoid or steroidal) linked to one or more sugar moieties. In plants, saponins are primarily involved in chemical defense, displaying antifungal, antibacterial, and anti-herbivory properties due to their ability to interact with membrane sterols [148].
Saponins are particularly abundant in representatives of the Amaranthaceae (e.g., Salicornia, Atriplex) and certain Asteraceae species, where they may accumulate in both roots and aerial tissues. Root accumulation is often associated with protection against soil-borne pathogens and high ionic concentrations, whereas in leaves and stems saponins may contribute to defense against herbivores and environmental stress. Salinity stress has been shown to stimulate saponin biosynthesis, likely through activation of the terpenoid pathway and stress-responsive signaling networks. Increased saponin levels under saline conditions may contribute to membrane stabilization, regulation of ion transport, and mitigation of oxidative stress. Their amphiphilic nature allows interaction with lipid bilayers, which may support cellular integrity under osmotic imbalance [116]. Saponins have significant food, pharmaceutical, and industrial relevance. They exhibit cholesterol-lowering, immunomodulatory, anticancer, and anti-inflammatory activities, while also functioning as natural emulsifiers and foaming agents [149]. Their bitterness and potential toxicity require careful processing. Halophytes rich in saponins represent promising non-conventional sources of these bioactive compounds, particularly for cultivation on marginal saline lands.
4.5. Carotenoids
Carotenoids are lipid-soluble pigments, including β-carotene, lycopene, and xanthophylls, the oxygenated derivatives of which comprise lutein, zeaxanthin, violaxanthin, and neoxanthin [150]. In higher plants, carotenoids are indispensable components of chloroplasts, where they are structurally and functionally associated with light-harvesting complexes and reaction centers.
A central role of carotenoids in plants is their contribution to photosynthesis and photoprotection. While they assist chlorophylls in capturing light energy, their most critical function lies in preventing photooxidative damage by dissipating excess excitation energy and quenching reactive oxygen species, particularly singlet oxygen [151]. In addition, carotenoids serve as precursors for apocarotenoids, including the phytohormone abscisic acid (ABA), which plays a pivotal role in regulating plant responses to abiotic stresses such as salinity, drought, and osmotic imbalance [150].
In halophytes, carotenoid metabolism is closely linked to stress adaptation. These plants frequently experience high salinity, intense solar radiation, and oxidative stress, conditions that strongly favor enhanced carotenoid biosynthesis. Numerous studies have shown that halophytes tend to accumulate higher total carotenoid contents and exhibit increased carotenoid-to-chlorophyll ratios compared with glycophytes, particularly under saline conditions [151]. Elevated carotenoid levels contribute to the stabilization of thylakoid membranes, maintenance of PSII efficiency, and reduction of lipid peroxidation caused by salt-induced ROS production [151].
A growing body of research suggests that antioxidant systems, including carotenoids, contribute to salt tolerance in Mediterranean and Iberian halophytes, as salt-tolerant species often show increased levels of non-enzymatic antioxidants under salinity stress, concomitant with improved photosynthetic performance [114,115]. Although carotenoids are often assessed together with other antioxidant compounds such as phenolics and flavonoids, these studies emphasize that photoprotective pigments play a critical role in mitigating oxidative damage in halophytes exposed to sub-optimal environments. This adaptive strategy allows halophytes to sustain growth and reproduction whereas glycophytes suffer severe photosynthetic inhibition.
From a nutritional and industrial perspective, carotenoids are among the most valuable plant-derived bioactive compounds. In human nutrition, several carotenoids—most notably β-carotene—act as provitamin A, essential for vision, immune function, and epithelial integrity. Moreover, carotenoids such as lutein and zeaxanthin are associated with reduced risk of age-related macular degeneration and other chronic diseases, largely due to their antioxidant and anti-inflammatory properties [152,153]. In the food, feed, and cosmetic industries, carotenoids are widely used as natural colorants and functional ingredients, responding to increasing demand for plant-based and clean-label products [154].
The fact that halophytes can maintain or enhance carotenoid biosynthesis under saline and high-light conditions suggests that these plants may serve as alternative and sustainable sources of high-value carotenoids. Cultivation of halophytes on saline or marginal lands could enable carotenoid production without competing with conventional crops for fertile soil or freshwater resources. This highlights halophytes as both model systems for studying stress physiology and promising candidates for future saline agriculture and bio-based industries [152].
4.6. Chlorophylls and Their Derivatives
Chlorophylls are photosynthetically active lipophilic pigments present in the majority of land plants. Apart from numerous studies concerning the changes in total chlorophyll contents under various environmental conditions, the scarcity of information on properties of chlorophylls and chlorophyll-derived compounds could be attributed to the difficulty to obtain chlorophylls and their derivatives in a purified form and to their instability, which triggers a degradation process [155]. Instead, halophytes often maintain high chlorophyll content and enhanced pigment stability, owing to their adaptation to saline and oxidative environments. These extremophytes may therefore constitute valuable sources of chlorophylls and chlorophyll-derived compounds with potential applications in the food, nutraceutical, and pharmaceutical sectors.
The content of chlorophylls in halophytes is usually higher compared to that of conventional leafy vegetables [96]. While moderate salinity conditions promote chlorophyll accumulation, sub-optimally high salinity levels (above 600 mM NaCl) often reduce its accumulation [156,157]. Total content of chlorophyll a + b in aerial tissues of Salicornia ramosissima and Sarcocornia perennis amounts to 28 and 25 mg per 100 g FW, respectively [96]. Pronounced accumulation of total chlorophylls was reported in Mesembryanthemum nodiflorum (280 mg/100 g DW), Sarcocornia fructicosa (290 mg/100 g DW) and Suaeda maritima (280 mg/100 g DW) [16,97]. Such high contents of chlorophylls calculated on a dry matter basis indicate that halophytes can concentrate chlorophylls to levels favorable for bioactive ingredient extraction [97].
Chlorophyll-derived compounds include chlorophyllides, pheophytins, chlorophyllins and pheophorbides. These compounds are usually products of chlorophyll degradation. Pheophytins are formed after Mg2+ ion removal, and pheophorbides after further dechelation [158]. Although chlorophylls are strong antioxidants, all chlorophyll-derivatives exhibit higher antioxidant properties than chlorophylls themselves [155]. Chlorophylls, together with their derivatives, are routinely exploited as food colorants, making the food appearance preferred by the customers. Owing to pronounced antioxidant capacity, chlorophylls and their derivatives are widely used as food additives protecting other ingredients from oxidation [159,160]. They are also potent antioxidants in mammalian cells systems [161], possess antiproliferative and anticancer activity [162,163], and their potential for application as chemo-preventing agents has been well documented [164]. For example, pheophorbide a and pheophytin derivatives function as photosensitizers that generate reactive oxygen species (ROS) when exposed to light, resulting in cancer cell death, which is exploited in photodynamic therapy [165,166].
Apart from a broad spectrum of chlorophyll applications in the food industry and pharmacology, studies concerning halophyte chlorophyll-derivatives are not available, indicating a serious knowledge gap that should be addressed in the immediate future. Recently it has been shown that copper chlorophyllin, a semi-synthetic water-soluble compound, functions as plant bio-stimulant and mediates defense responses in A. thaliana under salinity stress [167], so the prospects for specific features of halophyte chlorophylls seem substantiated. Chlorophyll turnover in halophytes could be significantly modified or enhanced, as the activity of chlorophyll-degrading machinery usually increases under stress [168,169], and greater accumulation of porphyrin is related to stress adaptation [170].
4.7. Glucosinolates in Brassicaceae Halophytes
Glucosinolates (GLSs) are sulfur-containing glycosides (β-thioglucoside-N-hydroxysulphates) predominantly found in members of the Brassicaceae family. GSLs are classified into three main groups based on the amino acid-derived side chain: the aliphatic group is synthesized from methionine, alanine, leucine, isoleucine, and valine, the indolic group with tryptophan, and the aromatic group with phenylalanine (Phe) and tyrosine (Tyr) as precursors [171]. GLSs constitute an element of the defense system against herbivores and are considered antinutrients in human diet [172]. On the other hand, as with many other glucosides, GLSs are highly bioactive and are at least partly responsible for such benefits for human health as anti-cancer, anti-inflammatory, cardioprotective and cholesterol-reducing effects [171,173,174]. Bioactivity of GLSs is primarily attributed to their hydrolysis products, formed after cell damage. Breakdown of GLSs is catalyzed by enzyme myrosinase, leading to formation of several products: isothiocyanates (ITCs), thiocyanates, and nitriles [171].
To date the presence of GLSs has been confirmed in Brassicaceae halophytes, such as Cakile maritima, Cochlearia officinalis, Lepidium latifolium, Crambe maritima and salt-tolerant ecotypes of Brassica oleracea var. palmifolia and Diplotaxis tenuifolia [100,157,175,176,177]. Their synthesis was also predicted in Eutrema salsugineum (syn. Thellungiella salsuginea) by analysis of the genome sequences [178]. These species accumulate both aliphatic (e.g., sinigrin, gluconapin, glucoiberin) and indolic GLS (e.g., glucobrassicin), often at concentrations comparable to or exceeding those of cultivated Brassica vegetables [100]. However, quantitative literature data for GLS content and profile in numerous halophytic species are limited. As GLS are metabolites specific also for other families among Brassicales, research attempts should be directed to evaluate potential for GLS accumulation in the species from Salvadoraceae, Capparaceae or Betaceae families, known for their halophytic nature [10,33].
The most extensively studied glucosinolate metabolites are isothiocyanates (ITCs) classified among the most potent plant-derived anticancer agents. Depending on the precursor GLS, various ITCs may be formed: allyl isothiocyanates, phenethyl isothiocyanates, and sulforaphane (SFN), each of these compounds exhibiting anticarcinogenic activity, from inhibition of cell cycle progression, induction of cell death, epigenetic modulation, to suppression of tumor initiation caused by SFN [179,180]. As GLS levels determined in studies od halophytes are comparable to, or exceed, those of cultivated vegetables, the species may be a promising source of anticancer metabolites.
Halophytes as a source of glucosinolates and their metabolites are still underutilized. Two species containing GLS (Cochlearia officinalis and Brassica oleracea var. palmifolia) have been recently explored as novel leafy vegetables suitable for sustainable indoor farming [156]. Since GLS content varies under various salinity levels [156,175]. controlled saline cultivation may be used to modulate GLS profiles, facilitating targeted production of specific bioactive compounds, i.e., antioxidant supplements or redox-modulating agents. However, understanding of salinity–glucosinolate regulatory networks remains incomplete and requires further studies.
5. Biological Activities and Clinical Studies of the Halophyte Plants: Exemplified by Lobularia maritima
Research on halophytes shows that soil salinity influences not only plant physiology by disrupting metabolism, growth, and development but also the quality of plant material. According to the literature, salt-induced stress weakens the photosynthetic capacity of plants, which may lead to reduced levels of carbohydrates, fatty acids, and proteins [181]. Conversely, under saline conditions, many plant species tend to accumulate higher amounts of nitrogen-containing compounds, including various amino acids (particularly proline, but also alanine, arginine, glycine, serine, leucine, and valine), amides such as glutamine and asparagine, polyamines, and specific stress-related proteins (such as osmotin, dehydrins, and LEA proteins [182].
The concentrations of secondary metabolites often referred to as stress metabolites strongly depend on the species and the environmental conditions in which the plants grow, especially when stress factors are present. Reports on how salinity affects phenolic compounds, chlorophylls, carotenoids, essential oils, and alkaloids remain inconsistent across studies. Nonetheless, several investigations indicate an increase in phenolic substances, including flavonoids (e.g., quercetin and apigenin), anthocyanins, and phenolic acids such as protocatechuic, chlorogenic, caffeic, and trans-cinnamic acids, in plants exposed to salt stress. Regarding essential oils, salinity has been shown to reduce the levels of compounds like anethole, carvacrol, and eugenol, while enhancing the production of chamazulene, α-bisabolol, trans-β-farnesene, γ-terpinene, and linalool. There is also strong evidence that increasing salinity significantly decreases the content of photosynthetic enzymes as well as chlorophyll a, chlorophyll b, and total carotenoids (Table 6).
Liver disorders represent a major health concern due to the liver’s central role in metabolism, detoxification, and the elimination of xenobiotics. Hepatotoxic substances, inappropriate dietary habits, and certain medications can all contribute to liver injury. Because liver impairment can have severe consequences, the increasing incidence of hepatic damage poses a significant challenge for the pharmaceutical industry, regulatory agencies, and healthcare professionals, all of whom are actively seeking new therapeutic strategies. While drugs such as immunosuppressants (thiopurines, methotrexate), corticosteroids (prednisone, prednisolone, budesonide), and TNF-α inhibitors (infliximab, adalimumab, certolizumab pegol, golimumab) are widely used, concerns remain regarding their hepatotoxicity. Consequently, natural plant-derived compounds have gained attention as safer alternatives or complementary therapies for liver disease prevention and treatment [183,184,185].
Epidemiological studies indicate an inverse correlation between antioxidant intake and the prevalence of hepatic diseases, emphasizing the protective potential of bioactive compounds [186]. Plant extracts, especially those rich in phenolic compounds, can mitigate oxidative damage by scavenging free radicals, inhibiting lipid peroxidation, and disrupting redox reactions involving metals such as iron [187]. Iron overload is commonly associated with chronic liver diseases and can exacerbate oxidative stress, accelerating disease progression [188]. In this context, the iron-chelating properties of halophyte Lobularia maritima contribute to its hepatoprotective effects. Bioactive constituents such as tannins, flavonoids, and phenolic acids have been implicated in these protective activities [189]. Experimental studies using the carbon tetrachloride (CCl4) model have confirmed the hepatoprotective properties of L. maritima. CCl4 induces oxidative stress that mimics biochemical and histological features of acute human hepatitis [48,190]. Plant extracts rich in phenolics have been shown to reduce lipid peroxidation and enhance the activity of endogenous antioxidant enzymes under CCl4-induced hepatotoxic conditions [191,192,193]. LC-MS and HPLC analyses have revealed that Lobularia maritima extracts contain a diverse array of bioactive compounds, including phenolic acids and flavonoids, which contribute to its potent antioxidant properties. The aqueous extract has been shown to contain phenolic acids such as protocatechuic, chlorogenic, quinic, p-coumaroylmalic, cinnamic, caffeine, p-coumaric, gallic, salicylic, ellagic, and ferulic acids, as well as flavonoids including catechin, myricetin, naringenin, apigenin 7-O-glucoside, narirutin, rutin, luteolin, kaempferol, and quercetin (Table 6) [48,194]. These compounds exhibit multiple bioactivities, including antioxidant, antibacterial, and hepatoprotective effects. Phenolic acids, for instance, protect against degenerative diseases such as cancer and cardiovascular disorders by scavenging reactive oxygen species, preventing lipid peroxidation, and enhancing LDL resistance to oxidation [195]. Flavonoids like quercetin inhibit lipid peroxidation through several mechanisms, including xanthine oxidase inhibition, iron chelation, direct radical scavenging, enhancement of vitamin C absorption, and protection of structural proteins [196,197,198,199]. Rutin (quercetin rutinoside) has been reported to prevent CCl4-induced elevations in transaminases and modulate pentobarbital-induced sleeping time in rats, demonstrating hepatoprotective activity [200]. Similarly, epicatechin supplementation has been shown to reduce malondialdehyde (MDA) levels and protect against oxidative damage in experimental models of hepatitis [201,202].
Additional phenolic compounds such as gallic, ferulic, and ellagic acids have demonstrated hepatoprotective effects in diverse models of liver injury. Gallic acid protects against paracetamol-induced liver damage, ischemia-reperfusion injury, CCl4-induced acute liver injury, and sodium fluoride-induced oxidative stress [203,204]. In combination with quercetin and rutin, gallic acid shows synergistic protective effects in acetaminophen-intoxicated rats [205]. Ellagic acid provides significant protection against CCl4-induced biochemical changes in the liver and blood and reduces lipid peroxidation in liver microsomes in vitro [206]. Ferulic acid also lowers liver enzyme levels in models of alcohol and polyunsaturated fatty acid-induced hepatic damage, with low doses proving more effective than higher ones [207]. Mechanistically, CCl4 hepatotoxicity involves oxidative stress mediated by the tri-chloromethyl radical (•CCl3) and its peroxyl derivative (CCl3OO•), generated during hepatic metabolism by phase I cytochrome P450 enzymes. These reactive species disrupt lipids and proteins, leading to structural and functional damage of hepatocytes, as confirmed by increases in Thiobarbituric Acid Reactive Substances (TBARS) levels, reductions in glutathione S-transferase (GST) and δ-aminolevulinic acid dehydratase (δ-ALA-D) activities, and histological evidence of fibrosis [208,209,210]. Elevated levels of malondialdehyde (MDA) and extensive parenchymal hepatocyte necrosis observed in CCl4-intoxicated mice are consistent with the oxidative damage induced in hepatocellular membranes [210]. The effectiveness of several hepatoprotective agents in neutralizing free radicals generated during these processes is particularly noteworthy [209]. Within cells, reactive oxygen species (ROS), such as superoxide anion (O2−) and hydrogen peroxide (H2O2), are produced during normal aerobic metabolism. The balance between ROS generation and elimination by antioxidants determines intracellular ROS concentrations. Key antioxidant enzymes—including superoxide dismutase (SOD), catalase (CAT), and glutathione peroxidase (GPx)—detoxify H2O2 and superoxide radicals, maintaining cellular redox homeostasis. Ben Hsouna et al. [210] evaluated the ability of L. maritima extracts to attenuate oxidative stress induced by CCl4. Their findings indicate that administration of L. maritima extract exerts a significant hepatoprotective effect against CCl4-mediated oxidative stress. The extracts markedly reduced CCl4-induced increases in MDA and restored the activity of detoxifying enzymes such as GST and δ-ALA-D. The δ-ALA-D enzyme is a sensitive marker of oxidative stress and impaired metabolic function, particularly in association with detoxifying systems like GST. Prior studies have highlighted the relevance of GST and δ-ALA-D in hepatotoxicity models [186,210,211]. CCl4-induced inhibition of δ-ALA-D underscores the susceptibility of its sulfhydryl groups to oxidation, leading to the accumulation of the prooxidant substrate amino-levulinic acid (ALA), increased intracellular H2O2, and mitochondrial dysfunction [212]. Restoration of GST and δ-ALA-D activities by L. maritima extracts suggests a protective effect against lipid peroxidation of hepatocyte membranes.
Inflammation is another key mechanism implicated in CCl4-induced liver injury and is closely linked to oxidative imbalance [213]. Drug-induced liver damage is often associated with elevated production of proinflammatory cytokines and mediators [214]. Compounds capable of modulating inflammatory responses may therefore have hepatoprotective effects [215,216]. Although Ben Hsouna et al. [217] did not quantify inflammatory mediators, histological analysis revealed severe eosinophilic infiltration in CCl4-treated livers. L. maritima extracts and certain plant-derived triterpenes are known to possess anti-inflammatory activity in experimental models [218]. However, in the same study, L. maritima extract did not significantly reduce eosinophilic infiltration, suggesting that modulation of inflammation may not be a primary mechanism underlying its hepatoprotective effect. More precise assays are needed to clarify this aspect. Overall, ethanol extracts of L. maritima exhibit a strong capacity to counteract oxidative damage induced by CCl4, highlighting their therapeutic potential in preventing and mitigating hepatotoxicity [202]. Flavonoids present in high concentrations in L. maritima extracts, including quercetin and caffeic acid, have shown significant protective effects against liver dysfunction [219]. These findings suggest that the combined action of flavonoids and other phenolic compounds contributes synergistically to reducing oxidative damage induced by the extract, as demonstrated in both in vitro and in vivo studies.
Table 6.
Major classes of bioactive compounds identified in Lobularia maritima, their natural sources, principal biological activities, and the corresponding types of experimental evidence.
6. Future Research Perspectives
Although halophytes exhibit high salinity tolerance, their productivity, distribution, and accumulation of nutritional and bioactive compounds are strongly influenced by climate change–related factors, including temperature fluctuations, altered precipitation patterns, and sea-level rise [6]. Seasonal variations in antioxidant capacity further demonstrate the significant impact of environmental conditions on metabolite biosynthesis and therapeutic potential [223]. A better understanding of these environmental drivers is essential for developing predictive models and improving the quality consistency of halophyte-derived products.
Most halophytes occur as wild plants across diverse habitats, resulting in high variability in yield, nutrient composition, and bioactive compound content. Despite initial efforts using hydroponic and aquaponic systems [102,224] optimized and standardized cultivation protocols remain insufficiently developed, limiting large-scale production and economic feasibility. Future studies should therefore prioritize agronomic standardization, controlled production systems, and breeding strategies aimed at improving productivity and compositional stability. Extraction technique is a crucial step in obtaining bioactive compounds from plants, besides other factors influencing the bioactivity of the extract, such as plant origin, harvesting time, and climate. There is a lack of literature on the extraction of halophytic biomass, particularly for industrial scale uses [225]. Furthermore, extraction solvents affect the extraction yield and may be hazardous to health and harm the environment, like most commonly used organic solvents [226]. Hence, it is desirable to replace them with environmentally friendly solvents. Thus, utilization of the bioactive chemicals found in halophyte plants necessitates further research on extraction techniques and subsequent scale-up for commercial usage.
Food safety remains an important research priority. Some halophytes contain anti-nutritional compounds such as oxalates, which may pose health risks if consumed in excess [26]. Comprehensive long-term studies assessing safe intake levels, potential adverse effects and bioavailability of bioactive compounds are still lacking for many species. [227]. Strengthening clinical validation and establishing clear safety guidelines will be crucial for broader commercial utilization. Advances in plant biotechnology provide promising tools to address these challenges. In vitro culture systems enable controlled studies of salinity responses and metabolic regulation [228,229], while genomic techniques such as CRISPR/Cas9 offer opportunities to enhance nutritional traits and secondary metabolite production relevant to pharmaceutical applications [230]. Nevertheless, the systematic application of these approaches to halophytes remains limited and requires further exploration.
Future research should integrate molecular biology, biotechnology, agronomy, food science, and clinical studies to bridge the gap between experimental findings and industrial implementation. Addressing these priorities will be essential for ensuring the safe, scalable, and sustainable use of halophytes under conditions of increasing soil salinization and global environmental change.
7. Conclusions
Halophytes represent a promising yet still underexploited group of plants with substantial nutritional and bioactive potential. Their adaptation to saline environments is associated with the synthesis of diverse secondary metabolites exhibiting antioxidant, anti-inflammatory, and other health-promoting properties. This review has systematically organized current knowledge on the nutritional composition and phytochemical diversity of selected halophytes, highlighting their relevance for food and pharmaceutical applications.
The evidence gathered demonstrates that halophytes may serve as valuable alternative crops for saline and marginal lands, contributing to sustainable agricultural systems under conditions of increasing soil salinization. At the same time, variability in metabolite composition and productivity, driven by environmental and ecological factors, underscores the complexity of their utilization.
Overall, the findings presented in this review confirm that halophytes hold considerable potential as innovative resources for functional foods and bioactive compounds. However, their broader implementation requires coordinated scientific and technological efforts to translate experimental knowledge into practical applications.
Author Contributions
Conceptualization, A.K.; methodology, A.K. and A.W.; writing—original draft preparation, A.K., A.W., M.A.H., D.K., A.E.-S., R.B.S. and A.B.H.; writing—review and editing, A.K., A.W., M.A.H. and D.K.; supervision, A.K. and A.W.; funding acquisition, A.K. and M.A.H. All authors have read and agreed to the published version of the manuscript.
Funding
This research received no external funding.
Data Availability Statement
The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author.
Acknowledgments
The authors acknowledge that the APC was waived by the journal in connection with their role as Guest Editors of the Special Issue.
Conflicts of Interest
The authors declare no conflicts of interest.
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