Halophytes: nutrients, bioactive compounds, chemical characterization and potential applications

Halófitas: nutrientes, compostos bioativos, caracterização química e aplicações potenciais

Abstract

Halophytes are plants adapted to saline soils found in a diversity of environments with varied climatic conditions and high salinity, developing different strategies to complete their life cycle. Their ecological, morphological, physiological and biochemical adaptations allow resistance to the abiotic stresses suffered and the formation of bioactive compounds that give these plants anti-inflammatory, antiparasitic, antimicrobial, antiviral and antioxidant potential properties. Therefore, this study aimed to compile its different biological activities, its nutritional compounds, potential applications for pharmaceutical and food industries, its biotechnological use. Halophytes have vast potential for pharmaceutical industries, as well as being a sustainable alternative in production of animal feed and a viable possibility for replacing the use of table salt and its various applications in human nutrition. It is necessary to invest in new and diversified research highlighted in this study.

Keywords:
bioindicators; foods; vegetable salt

Resumo

Halófitas são plantas adaptadas a solos salinos encontradas em vários ambientes com condições climáticas variadas e alta salinidade, desenvolvendo diferentes estratégias para completar seu ciclo de vida. Suas adaptações ecológicas, morfológicas, fisiológicas e bioquímicas permitem a resistência aos estresses abióticos sofridos e a formação de compostos bioativos que conferem a essas plantas propriedades anti-inflamatórias, antiparasitárias, antimicrobianas, antivirais e potencial antioxidante. Assim, este estudo teve como objetivo compilar as suas diferentes atividades biológicas, os seus compostos nutricionais, as suas potenciais aplicações para a indústria farmacêutica e alimentar e a sua utilização biotecnológica. As halófitas possuem um vasto potencial para as indústrias farmacêuticas, além de ser uma alternativa sustentável na produção de ração animal e uma possibilidade viável para substituir o uso do sal de cozinha e suas diversas aplicações na alimentação humana. É necessário investir em novas e diversificadas pesquisas destacadas neste estudo.

Palavras-chave:
bioindicadores; alimentos; vegetais salinos

1. Introduction

Term “halophytes” derives from Greek “halos”, which means “salt”, and “phyton”, which means “plant” (Flowers et al., 1986). The term is used to describe a group of plants that are highly capable of growing and developing in environments directly or indirectly influenced by ocean waters, such as mangroves, estuaries, saline deserts, coastal dunes, coastal marshes and lands degraded by inadequate agricultural practices (Hameed and Khan, 2011).

Halophytes are flowering plants adapted to saline soils, wicht are found in a variety of environments with varying salinity and climatic conditions (Duarte and Caçador, 2021), to complete their life cycle in such adverse conditions, halophytes have developed different strategies such as development of juiciness, compartmentalization of toxic ions, synthesis of osmotic, increased activity of antioxidants and synthesis of compatible solutes (Tipirdamaz et al., 2006).

Synthesis of secondary metabolites allows these plants to have differentiated ability to develop in saline habitats (Todorović et al., 2022). Various morphological, physiological, biochemical and ecological adaptations affect halophyte plants so that they can resist at abiotic stresses suffered, such as adaptive roots, osmotic adjustments to maintain water balance, excess saline excretory glands, interactions with local microbiota and with animals themselves. Animals, becoming an important object of study around the world (Grigore, 2008).

High degree of stress suffered by these plants contributes to formation of bioactive compounds that confer anti-infectious, antiparasitic, antimicrobial, anti-inflammatory properties and high antioxidant activity, these compounds being responsible for use of halophyte of various genera and species as medicinal plants (Rodrigues et al., 2023).

Brazil has a diverse flora of halophytes with biotechnological potential for production of biofertilizers, economical for production of food and therapeutic for production of bioactive substances. They are distributed in coastal environments under influence of flooding by salt water, such as in mangroves, occurrence of Rhizophora mangle, R. harrisonii, R. racemosa, Avicennia germinans, A. schaueriana, Conocarpus erecta and Laguncularia racemosa (Costa and Herrera, 2016; Costa and Bonilla, 2017); in salt marsh environments close to mangroves, occurrence of grasses, such as Spartina alterniflora or Sporobolus virginicus (Costa et al., 2009), dicotyledons Batis maritima, Sesuvium portulacastrum, Sarcocornia ambigua and Blutaparon vermiculare, cosmopolitan ferns Acrostichum aureum and Acrostichum danaefolium, grass monocotyledons Spartina alterniflora and Spartina densiflora, Cyperaceae Scirpus maritimus and Scirpus olneyi, reed Juncus kraussii and a shrubby area covered by Myrsine parvifolia and Acrostichum danaefolium (Costa et al., 2003).

Halophyte species (Apium graveolens, Myrsine parvifolia, Paspalum vaginatum and Schinus terebinthifolius) proved to be good natural sources of free phenolic compounds (FPC) when compared to soybeans and rice bran, being first study to report FPC extraction in M. parvifolia and P. vaginatum, and their seringic acid, vanillin and quercetin contents were not affected by salinity (Souza et al., 2018).

Research carried out with Salicornia neei Lag., showed that native species from Brazil has high tolerance to salinity and climate of semiarid northeastern region, highlighting adaptive and promising potential of this species for local cultivation (Alves et al., 2019). Another study tested cultivation of Salicornia neei Lag. with brackish water irrigation or fertilizer, with objective of evaluating growth and biomass gain and found a greater productivity of fresh and dry biomass of plants in the 2^nd harvest, demonstrating the forage potential presented by this halophyte, promising for cultivation in large areas of Brazilian semi-arid region (Alves et al., 2020). Regarding nutritional properties, high levels of minerals are observed, such as K, Mg, Ca, Zn, Fe and Mn (Doncato and Costa, 2018), fatty acids rich in ω-6 and ω-9, in addition to antioxidant and anti-inflammatory phenolic compounds (Souza et al., 2018). A study shows that antioxidant compounds can be indicated for treatment of obesity, including diabetes mellitus (DM), which is a comorbidity closely related to obesity (Khutami et al., 2022). The Figure 1 below shows the different applications of halophytes, based on Grigore and Toma (2017) studies.

Figure 1
Different applications of halophytes.

Presence of bioflavonoids volkensiflavone and fukugetin (Botta et al., 2004) and prenylated xanthones Delle et al., 1984) are some of important compounds found in halophytes. These compounds are associated with biological activities, such as free radical scavenging and antiulcer effect (Yamaguchi et al., 2000), cytotoxicity, inhibition of nitric oxide synthase (Shan et al., 2015), cancer chemoprevention (Ito et al., 2003) and trypanocidal effects (Abe et al., 2004).

Due to different biological activities presented by halophytes, this study aims to describe nutritional and bioactive compounds found, their potential applications aimed at pharmaceutical, nutraceutical, and food industries, as well as their applications in biotechnology.

2. Halophyte Plants Characteristics

Halophyte plants have developed physiological, biochemical and molecular control to deal with salt excess, allowing them to grow and reproduce in extreme conditions (Flowers et al., 2010). One of the characteristics of halophytes is ability to accumulate large amounts of salt in their cells, without suffering damage while completing their life cycle (Flowers and Colmer, 2008), this is possible due to property of compartmentalizing ions in vacuoles of their cells, preventing high loads of salt from substrate from reaching protoplasm, thus achieving regulation of salinity and its tolerance to toxic effects associated with an increase in this concentration (Holanda et al., 2011). In addition, they developed layers of semi-permeable cells for NaCl in epidermis and a serous layer in endodermis that works as a kind of filter for salt and deeper and more extensive roots to fix plant in soil (Glenn et al., 1998, 2013).

Commonly, halophytes carry out some mechanisms of adaptation and tolerance to sodium, such as having special glands that excrete salt excess on surface of leaf, forming a layer of salt crystals (Meng et al., 2018). Some species repress absorption of sodium ions by roots, reducing transport to aerial parts of plants (Sui et al., 2010). Others dilute salt accumulated in leaves (Flowers et al., 2010) or excrete it through structures located in epidermis (Yuan et al., 2015). Furthermore, halophytes developed a specialized root system, composed of adventitious roots capable of tolerating waterlogging (Flowers et al., 2010).

3. Use of Halophytes in Ancient Times

In antiquity, many civilizations in coastal regions or near deserts already had traditional knowledge to treat various infectious diseases using halophyte plants, exploring these plants in search of therapeutic Properties (Grigore and Constantin, 2010). Parts of plant, such as leaves, fruits, seeds, roots and bark, were widely used to prepare teas and topical pastes for therapeutic use (Fan et al., 2019). In some regions of Mediterranean, they were used to treat urinary and respiratory diseases, in addition to consumption in food (Öztürk et al., 2014). Coastal halophyte species along Arabian sea are described for therapeutic purposes such as Amaranthus virdis L. (Amarantaceae) used for treatment of constipation, gallbladder/kidney stones, Album chenopodium L. (Chenopodiaceae) used for treatment of constipation and many other plants described in Qasim et al. (2011).

In ancient Greece, for example, the use of plants from Amaranthaceae genus, such as Salicornia and Salsola, were very common in cooking and medicinal use (Lombardi et al., 2022). In ancient Egypt, some plants were cultivated to produce oils, cosmetics and medicines, in addition to using them in food (Grigore and Constantin, 2010).

It is well known that problem of salinity has affected ancient civilizations, such as those close to saltwater and desert regions. However, practice of cultivating halophyte plants has become promising, which has allowed civilizations to overcome challenges posed by salinity, in addition to providing cultivation alternatives that today have become innovative solutions to some environmental problems (Ventura et al., 2015).

4. Halophytes: Species and Applications

Salicornia is a botanical genus that belongs to Amaranthaceae family, halophytes rich in nutrients, including vitamins, minerals and antioxidants, in addition to high protein contente. In general, Salicornia have a notable importance in human nutrition, especially in coastal regions and in places where food cultivation is hampered due to soil salinity (Mishra et al., 2015). There are several Salicornia species of importance for studies, such as S. bigelovii, a plant native to North America. It is a perennial, succulent plant, with leaves reduced to small scales and its stem is cylindrical and articulated. Distinctive aspect of this species is its bright green color, which contrasts with the marshy and saline áreas (Alfheeaid et al., 2022). Among most studied Salicornia species, S. bigelovii Torr is the most searched species as it is most salt tolerant (Ayala and O’Leary, 1995), which makes them promising for cultivation of this oilseed species in desert areas under irrigation with sea water (Glenn et al., 2013). S. bigelovii has attracted increasing interest due to its potential use in various applications, such as food due to its richness in nutrients such as ascorbic acid and chlorophyll in addition to its characteristic flavor (Lyra et al., 2020; Liu et al., 2020), in production of biofuels due to its oil content (El-Tarabily et al., 2020) and in agricultural and economic potential as biofertilizers, due to this species being associated with a variety of microorganisms that colonize roots of plant and establish a mutualistic relationship (mycorrhiza), which consequently promotes plant growth and also confers benefits to other plants (Pseudomonas sp. (SDT3), Bacillus velezensis (SMT38), Pseudarthrobacter oxydans (SRT15) and Bacillus zhangzhouensis (HPJ40) (Mathew et al., 2020; Valle-Romero et al., 2023). Therapeutic properties of S. bigelovii are attributed to high oil content in its seeds (about 30%), such as linolenic acid (C18:3) n-3 PUFA (77%) essential for cardiovascular functions, as it helps reduce substantially levels of triglycerides and cholesterol (Low-Density Lipoprotein - LDL), in addition to regulating blood pressure, it acts on immune system, reducing risk of chronic diseases (Anwar et al., 2002), acts reducing risk of neurodegenerative diseases, such as Alzheimer's (Alpha-linolenic Acid (ALA) produces Eicosapentaenoic Acid (EPA) (20 carbons) and Docosahexaenoic Acid (DHA) (22 carbons), precursors of a group of eicosanoids (prostaglandins, thromboxanes and leukotrienes) which are anti-inflammatory, antithrombotic, antiarrhythmic and vasodilators). S. europaea, also known as glasswort, is native to coastal regions of Europe, North Africa and Western Asia. It is characterized by having bright and succulent stems, branched and green to reddish in color, with leaves reduced to small scale-shaped structures that surround stems (Zhang et al., 2014). Some compounds are found in this species, including flavonoids, flavanones, chromones, sterols, lignans, aliphatic compounds and triterpenoid saponins (Kim et al., 2021). When analyzing biological and therapeutic properties of S. europaea, antiproliferative activities were observed against neoplastic cells A549 (adenocarcinomic human alveolar basal epithelial cells) with IC50 (concentration that inhibits 50% of cell growth) of 52.35 and 79.39 μm, S. europaea also showed antioxidant, anti-inflammatory, antidiabetic, antimicrobial, antihypertensive and antiviral activities (Ayeleso et al., 2017). S. herbacea is a succulent plant with scale-shaped leaves, it is also considered an edible plant, which has aroused human interest in its use. S. herbacea has bioactive properties such as antioxidants, phenolic compounds, flavonoids, saponins, alkaloids and tannins (Limongelli et al., 2022). The same species of halophyte was found in study of Essaidi et al. (2013) that had eight phenolic acids and eight flavonoids, containing antioxidants that help neutralize free radicals in the body and protect it against cellular damage that is associated with a series of chronic diseases, in hyperglycemia control (Lee et al., 2015), hepatoprotective effect (Yi et al., 2015) and in obesity control (Lee et al., 2023). S. ramosissima, also known as “purple glass”, is a plant native to coastal regions with a temperate climate. It has succulent, fleshy stems that are generally green, but can acquire reddish tones during growth (Limongelli et al., 2022). Its ends provide vitamins, minerals, proteins and amino acids, which make them valuable in cooking and can be consumed as salads, cooked or accompanied with meat or fish dishes (Antunes et al., 2021). This plant can be cultivated in a sustainable hydroponics system, providing a food with a good nutritional profile and safe in terms of toxicological and microbiological studies (Lima et al., 2020) and photoprotective as it contains a variety of phytochemicals as compounds present in ethyl acetate fraction present in the S. ramosissima extract, it was shown to be capable of protecting against ultraviolet (UV) rays (Surget et al., 2015). Furthermore, they are considered potent antioxidants and anti-inflammatory (Isca et al., 2014; Lima, 2023). S. brachiata, native of coastal regions of Asia, as well as other Salicornia are plants that have adaptive morphological structures that are similar to each other, such its ability to adapt to saline environments. In phytochemical prospecting, it was possible to find several metabolites with different bioactivity, such as flavonoids used for nutritional supplementation, saturated and polyunsaturated fatty acids in treatment of inflammatory and antiproliferative diseases, including selenium and sulfur as anticancer agentes (Mishra et al., 2015).

Brazil has a diverse flora of halophytes with biotechnological potential for production of biofertilizers, economical for production of food and therapeutic for production of bioactive substances. They are distributed in coastal environments under influence of flooding by salt water, such as in mangroves, occurrence of Rhizophora mangle, R. harrisonii, R. racemosa, Avicennia germinans, A. schaueriana, Conocarpus erecta and Laguncularia racemosa (Costa and Herrera, 2016; Costa and Bonilla, 2017).

In salt marsh environments close to mangroves, occurrence of grasses, such as Spartina alterniflora or Sporobolus virginicus, dicotyledons Batis maritima, Sesuvium portulacastrum, Sarcocornia ambigua and Blutaparon vermiculare, cosmopolitan ferns Acrostichum aureum and Acrostichum danaefolium, Grass monocotyledons Spartina alterniflora and Spartina densiflora, Cyperaceae Scirpus maritimus and Scirpus olneyi, reed Juncus kraussii and a shrubby area covered by Myrsine parvifolia and Acrostichum danaefolium (Costa et al., 2003). Halophyte species (Apium graveolens, Myrsine parvifolia, Paspalum vaginatum and Schinus terebinthifolius) proved to be good natural sources of free phenolic compounds (FPC) when compared to soybeans and rice bran, being first study to report FPC extraction in M. parvifolia and P. vaginatum and their seringic acid, vanillin and quercetin contents were not affected by salinity (Souza et al., 2018). Research carried out with Salicornia neei Lag., showed that native species from South America has high tolerance to salinity and climate of semiarid northeastern region, highlighting adaptive and promising potential of this species for local cultivation (Alves et al., 2019).

Another study tested cultivation of Salicornia neei Lag. with brackish water irrigation or fertilizer, with objective of evaluating growth and biomass gain and found a greater productivity of fresh and dry biomass of plants in the 2^nd harvest, demonstrating the forage potential presented by this halophyte, promising for cultivation in large areas of Brazilian semi-arid region (Alves et al., 2020). Regarding nutritional properties, high levels of minerals are observed, such as K, Mg, Ca, Zn, Fe and Mn (Doncato and Costa, 2018), fatty acids rich in n-6 and n-9 PUFA (Costa et al., 2014), in addition to antioxidant and anti-inflammatory phenolic compounds (Souza et al., 2018).

More than 100 countries in world are affected by problem of salinization (FAO, 2021). In Asia, halophytes have shown promise for agriculture, biofuels, salinity control and environmental preservation with a diversity of halophyte plants resulting from region's climate variation (Khan and Qaiser, 2006).

Low environmental impact for cultivation of halophyte plants is the great reason for diversified studies around the world focused on production of biomass for food and other products, their ability to tolerate environmental stresses and high levels of salt, their nutrients and bioactive substances make halophyte plants a viable alternative for human and animal nutrition in addition to possibility of rehabilitating degraded areas by removing pollutants from soil and preventing desertification (Khan et al., 2016).

In China, these plants are natural resources of potential economic value, as they provide important grains, fruits and vegetables in human and animal nutrition, as well as raw materials for production of biofuels (Liu and Wang, 2021). Artiplex canescens (Pure Nutri, had its potential for use questioned and applied to nutritionally poor lands, with exploration in viable application in needy areas to extract its potential ecological and economic values (Ma et al., 2022).

Studies in most diverse areas of biotechnological applications, animal feed, green food and gourmet vegetables, are developed in countries such as Iran, Tunisia, Algeria, Israel and Argentina, demonstrating potential for development of most diverse species of halophytes (Farzi et al., 2017; Aouissat et al., 2009; Ventura et al., 2015).

The halophyte Arthrocnemum frutcosum (L) present on Jordanian coast of Dead Sea was studied because it is one of few species in area that for most of year prevails in green color including hot and dry summer months, understanding germination and related ripening with these peculiar types of plants, it seeks to explain relationship between soil solidarity, rain, and germination, environmental and physiological aspects (Saadeddin and Doddema, 1986).

In Portugal, a study on potential of bioactive compounds from aerial parts of halophyte Poliganum maritimum L. evaluated antioxidant and antigenotoxic properties of ethanolic extract, with intention of designing development of products aimed at some health benefit in food and pharmaceutical industrys (Oliveira et al., 2023).

Production of botanical extracts with bioactives from Salicornia ramosissima J. Woods has been explored for application in high-value industries such as nutraceuticals, pharmaceuticals and cosmetics and its residual fraction after extraction can be used for bioenergy or chemicals derived from lignocellulose (production of biogas and bioethanol) (Hulkko et al., 2023).

In central region of Saudi Arabia, plants native to region (Euphorbia chamaesyce, Bassia arabica, Fagonia mollis and Haloxylon salicornicum) had their phytochemical contents studied, with different levels of antioxidant and antimicrobial activities evaluated, and in this study the alcoholic extract with most potent bioactivity was from E. chamaesyce, rich in polyphenols and flavonoid secondary metabolites such as 68.0 mg/g of gallic acid and 39.23 mg/g of quercetin (Rugaie et al., 2023).

In Republic of Serbia, a study reviewed biological activities such as antioxidant, anticancer, antibacterial, antifungal, anti-inflammatory of the Amaranthaceae family with 18 species of halophyte plants involved (Todorović et al., 2022).

Improvement and growth of halophyte plants through accumulation of osmotic agents keeps their biological functions active and their biosynthesis interferes in development stages (Slama et al., 2015).

Lycium humile (Lyciae tribe), a non-endemic shrub from northeastern Chile and northwestern Argentina that inhabits solar (saline) areas in Puna region, was studied as it grows in multi-stressed environments with water deficit, extreme temperatures and high salinity in order to analyze germinal responses of species (Palchetti et al., 2020).

In the United States, a study with a halophyte belonging to Amaranthaceae family of genus Salicornia, demonstrated its potential as food, indicating it as a plant for future, due to its resistance to extreme conditions with high salt content, its culinary relevance and medicinal attributes, suggesting new studies to combat food insecurity and production of biomass through species studied, transformed into pickles, drinks or in natura in salad (Patel, 2016).

5. Nutrients, Biochemical Compounds and Toxicity of Halophytes

Development of a food culture with possibility of less use of fresh water in irrigation, greater nutritional use, energy production from renewable biofuels, development of nutraceutical products and knowledge of some biological activities such as antioxidant, vermifuge, anti-inflammatory and bactericidal, definitely gives halophyte plants exceptional characteristics, directing studies towards more accurate and complete investigations of their nutrients, biochemical compounds, chemical characterization, their possible toxicity, as well as anti-nutritional factors for non-traditional production of food, fuels and chemical products (Souza and Costa, 2021).

5.1. Nutrients

In studies by Wu et al. (2012), Suaeda parsley, popularly consumed as a vegetable in Yellow River Delta, when subjected to water stress presents betaine 10 to 100 times greater than other metabolic substances (ethanol, lactate and alanine), betaine is an amino acid that protects against osmotic inactivation by increasing cell water retention, responsible for reducing percentage of body fat in animal models (fish and chick feed) improving nutritional efficiency and growth (Freitas et al., 2015). Other studies show that for Salicornia europaea L., desalination of powder can be carried out with a view to its use in diet, due to its antihypertensive and antilipogenic properties, helping to control obesity, its extract has a hypoglycemic effect due to inhibition of α-amylase enzyme (Rahman et al., 2021; Li et al., 2020). S. salsa has components such as vitamins, essential amino acids and its seeds contain 40% oil rich in unsaturated fatty acids (Wu et al., 2012), rich in proteins, fiber and carotenoids, indicating its potential as a vegetable and oilseed crop (Coc-Coj et al., 2020; Li and Song, 2019). The percentage of lipids may vary between different species of Salicornia seeds; S. bigelovii presented 29% extraction of total lipids, Salicornia brachiata 29.4%, Salicornia fruticosa 26.4% and Salicornia europaea 27.8% in addition to having a low saponin content in seeds, allowing extracted oil to be used for food purposes, results also demonstrated five components of fatty acids in analyzes of Salicornia bigelovi L. with 72.5% linoleic acid n-6 (C18:2), 13.3% oleic acid (C18:1), 7.40 wt% palmitic acid (C16:0), 2.4 wt% stearic acid (C18:0) and 2.3 wt% n-3 linolenic acid (C18:3) (Al-Rashed et al., 2016).

Biomass of Salicornia ramosissima J. Woods, known as sea asparagus, presented polyunsaturated fatty acids (PUFA 58.2%), saturated fatty acid (SFA 41.0%) and monounsaturated fatty acid (MUFA 1.3%) with a predominance of linoleic acid (34.5%) and palmitic acid (30.9%), in addition to a proportion of 1.5% of n-6 and n-3, suggesting a reduction in risk of cardiovascular diseases. Factors such as flooding, high temperatures and intense ultraviolet radiation trigger a protective effect with production of proanthocyanidins and other flavonoids, as well as bioactive compounds and production of free radicals and aqueous extract of plant matter presented bioactive compounds (30.10 mg GAE/g DM) (Hulkko et al., 2023).

Studies of chemical composition of Salicornia ramosissima and Sacocornia perennis Alpini, demonstrated values of moisture (89.7 and 87.8%), protein (6.61 and 4.28%), lipid (1.32 and 1.52%), ash (40% for both), fiber (11.3 and 15.3%) and carbohydrates (51.3 and 52.3%) respectively. These values were higher when compared to those found in a vegetable commonly used in general diet such as beetroot with moisture (88.39%), protein (1.06%), lipids (0.22%), ash (0. 92%), fiber (2.87%) and carbohydrates (6.54%) (Lopes et al., 2023).

5.2. Bioactive compounds

In research Rugaie et al. (2023), four species of halophyte plants studied (Bassia arabica, Fagonia mollis, Haloxylon salicornicum and Euphorbia chamaesyce, all native to central region of Saudi Arabia), last respectively mentioned, presented higher values of phenolics and flavonoids 68.00 mg/g gallic acid (GAE) and 39.23 mg/g quecertin (QE) respectively, in addition to demonstrating three times (3x) more free radical scavenging activity compared to other species in study. In the same study, sixteen flavonoid compounds were identified among them; rutin, myricetin, luteolin, quercetin, naringenin and kaempferol and natira and phenolics such as gallic and ellagic acid.

Several bioactive compounds are described in research of as flavonol glycosides (Patuletin 3-O- [5'”- O-feruloyl-β-D-apiofuransil (1'→ 2“) -β -Dglucopyranoside]; Patuletin 3-O-β-D-glucopyranoside) and flavonoid glycosides (Spinacetin 3-O-β-D-glucopyranoside; arbutin; 4-hydroxybenzyl-β-D-glucopyranoside) all found in Atriplex littoralis. Flavonoids such as myricetin, quercetin and several glycosylated flavonoids have been described in Artiplex halimus. For Camphorosma monspeliaca, essential oils (α-pinene; citronellyl pentanoate; limonene; pinocarvone; camphene; α-cadinol; octen-3-ol; β-eudesmol) and for Chenopodium ambrosioides, a large list of terpenes (β-myrcene; Cis-β-ocimene; Nerol; Geraniol; Limonene; α-terpinene; α-terpinоlen; β-phellandrene; p-cymene; Trans-pinocarveol; α-terpineol; Isoascaridole; Dihydroascaridole; Cariophyllenepoxide; δ3-carene; δ4-carene; γ-curcumene; α carotene; β-carotene), these compounds being just some of many described relating to species studied in research (Todorović et al., 2022).

6. Animal Feed

As they are rich in nutrients (antioxidants, fatty acids and amino acids), climate change, together with natural and anthropogenic problems in agricultural areas, resulting in increased problems with drought and salinity in several regions of the world (Nikalje et al., 2019), halophytes have important interests applied to various agricultural purposes to maintain ecological balance, in which they are currently being used as alternative plants in the production of feed and forage/fodder for animal feed (Centofanti and Bañuelos, 2019).

Moujahed et al. (2015) report that most African countries suffer from water shortages and salinization of agricultural soils, and this situation worsens every year due to climate change, and for this reason livestock farming suffers from chronic food shortages, essentially during dry periods of year, making it necessary to import a large percentage of animal feed. Halophytes are considered an alternative solution to problems related to food security, some of them, such as Chenopodiaceae, contain undesirable compounds for human consumption and to overcome this disadvantage, they can be offered to animals as a supplement or cultivated in combination with other glycophytes, such as legumes (Scopel, 2019).

Nikalje et al. (2019) highlight that forage crops have high biomass, digestibility and palatability for animals, and that among different halophytes, Desmostachya bipinnata and Panicum turgidum, are potential candidates for feed production, as they contain high levels of proteins, low oxalate, fiber and ash content. In this way, conventional forage crops, such as corn, can be replaced by halophyte plants, but due to relatively high salt content and anti-nutritional properties, there are some restrictions on their use.

Another issue is that salt content of halophytes can be nullified by using a mixture of forage plants for food, such as A. nummularia, when mixed with other herbaceous species and annual grasses (with low salt content) can be a good forage for animals (Attia-Ismail, 2018).

Plants like Aegiceras corniculata, Rhizophora mucronata, Avicennia marina and Ceriops tagal are used for camels and cattle food. In arid and semi-arid regions, Salvadora, Acacia, Prosopis and Ziziphus are incorporated into traditional forages. Salicornia, Chenopodium, Atriplex, Suaeda, Salsola and Kochia are shrubs, while Chloris virgata, C. gayana, Echinochloa turnerana, E. colonum, Aeluropus lagopoides, Sporobolus marginatus, Dactyloctenium sindicum, Puccinellia distans and S. marginatus are popularly used grass species as animal feed and fodder (Shiran et al., 2020).

However, despite promising use of halophytes in production of animal feed and forage, many problems related to cultivation and commercialization still need to be resolved, necessitating creation of market demand for consumption of halophytes (Nikalje et al., 2019).

7. Conclusions

Halophytes are plants adapted to saline environments, capable of growing in adverse conditions. They have an unique characteristics that make them valuable in several áreas.

Extracts and essential oils from these plants can be used in natural and innovative products, meeting growing demand for sustainable products in cosmetic industry.

Halophytes can also be used in production of animal feed, especially in regions affected by drought and salinity. They offer an alternative for production of forage and feed, contributing to sustainability of agriculture in adverse environments.

Furthermore, play a crucial role in remediation of saline soils and waters. However, it is important to highlight that many of these plants have not yet been scientifically evaluated regarding their health benefits and risks with long-term studies, and their use should be carried out with caution. it is essential to invest in research to fully understand their characteristics, properties and possible applications, thus ensuring sustainable and responsible use of these plants for benefit of society and environment.

Acknowledgements

We acknowledge the Graduate Program in Health and Development in the Central-West Region of Brazil, Federal University of Mato Grosso do Sul-UFMS, Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES).

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Publication Dates

• Publication in this collection
  14 Feb 2025
• Date of issue
  2024

History

• Received
  15 June 2024
• Accepted
  22 Nov 2024