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Halophyte Plants as Potential Sources of Anticancer Agents: A Comprehensive Review

Luísa Custodio
Pedro Garcia-Caparros
Catarina Guerreiro Pereira
1 and
Pedro Castelo-Branco
Centre of Marine Sciences, Faculty of Sciences and Technology, Campus of Gambelas, University of Algarve, 8005-139 Faro, Portugal
Agronomy Department of Superior School Engineering, University of Almeria, 04120 Almería, Spain
Faculty of Medicine and Biomedical Sciences (FMCB), Campus de Gambelas, University of Algarve, 8005-139 Faro, Portugal
Algarve Biomedical Center Research Institute (ABC-RI), 8005-139 Faro, Portugal
Champalimaud Research Program, Champalimaud Center for the Unknown, 1400-038 Lisbon, Portugal
Author to whom correspondence should be addressed.
Pharmaceutics 2022, 14(11), 2406;
Submission received: 13 July 2022 / Revised: 28 October 2022 / Accepted: 4 November 2022 / Published: 8 November 2022
(This article belongs to the Special Issue Advances in Anticancer Agent)


Salt-tolerant plants (halophytes) are widely distributed worldwide in several environments such as coastal salt marshes, sand dunes, and inland deserts. To cope with the harsh conditions that characterize those habitats, which include high salinity and radiation levels, such plants have developed morphological and physiological traits, the latter including the synthesis and accumulation of important secondary metabolites such as alkaloids and polyphenols. While essential in maintaining plant homeostasis, these compounds are highly valued in the medical field for the treatment of several human diseases, including cancer. Cancer is one of the most life-threatening disorders worldwide, which accentuates the need to improve current cancer therapies and minimize potential adverse secondary side-effects. In this context, the pharmacological evaluation of natural compounds has attracted growing interest since nature has already provided some important anti-cancer drugs. This review compiles, for the first time, research regarding the anticancer activity of halophytes from different families, including, whenever possible, the bioactive molecules involved in such therapeutical properties along with possible mechanisms of action. The introduction section provides some pertinent information regarding cancer and a summary of the most important characteristics of halophytes. The next section gives information regarding the in vitro and in vivo cytotoxic properties of several halophyte species, grouped by families, including contents in bioactive metabolites and proposed modes of action, if possible. Lastly, the conclusion presents the most relevant metabolites and/or promising species and extracts that could be further explored in anticancer drug research.

1. Introduction

Cancer is a group of diseases characterized by malignant neoplasms arising from the abnormal and uncontrolled cell proliferation that invades and destroys the surrounding tissue and, if not controlled, can result in death [1,2]. With an established complex link between cancer and aging and the increased risk factors of an unhealthy lifestyle, cancer is undoubtfully a major health problem in developed and developing countries [3,4,5,6,7]. It is estimated that, by 2040, the number of new cancer cases diagnosed per year will be higher than 27.5 million worldwide, with 16.3 million cancer deaths due to population growth and aging [8]. Globally, lung and breast cancers are the most frequently diagnosed and are the leading causes of cancer-related death in men and women, respectively [9]. Significant advances have been made in cancer research in the last 20 years concerning its general biology, prevention, and treatment [10]. Currently, chemotherapy is one of the most common cancer therapeutic approaches, although secondary effects are known and include a negative impact on the immune system, and the effectiveness of this therapeutic approach can be limited by drug resistance [11,12]. Therefore, new anticancer drugs and therapies that concomitantly minimize its harmful side-effects are still in urgent demand.
In the war against cancer, the role of natural compounds has become of crucial interest, and the scientific community has placed great effort in identifying novel sources of antitumoral molecules among medicinal plants and, more recently, marine organisms [10,13]. The pharmacological evaluation of natural compounds has rendered some important anticancer drugs such as paclitaxel and its derivatives from the bark of the Pacific yew tree (Taxus brevifolia Nutt, family: Taxaceae), or vincristine and vinblastine from Madagascar periwinkle (Catharanthus roseus (L.) G. Don, family: Apocynaceae) [14]. However, in addition to glycophyte plants and some marine species, other organisms including halophytes contain metabolites with potential use as anticancer drugs.
Although the definition of halophytes has yet to reach consensus, some authors define them as plants completing their life cycle in environments with a salt concentration of around 200 mM of sodium chloride (NaCl) (20 dS m−1) or more [15,16,17]. They represent nearly 1% of the global flora diversity and inhabit different ecosystems such as beaches, rocky shores, saltmarshes, estuaries, and inland deserts [17,18,19]. Such areas are characterized by stressful abiotic conditions, (e.g., high salinity, radiation levels, and drought), which trigger the production and accumulation of reactive oxygen species (ROS) in the plants, leading to cellular and tissue damage, metabolic disorders, and senescence. Halophytes are equipped with strong antioxidant defense systems to counteract the negative effects of ROS, including antioxidant enzymatic mechanisms and the synthesis of secondary metabolites, such as phenolic compounds, saponins, and alkaloids [18,20]. These molecules have a crucial protective role in the plant and exhibit relevant bioactivities, including antioxidant, anti-inflammatory, and antitumoral, which are linked to beneficial therapeutic properties and could help explain the use of some halophytes in traditional medicine and as food [18,21,22].
There are several studies focused on the anticancer properties of a high number of halophytic species, but this information is scattered in the literature. Therefore, our aim was to compile and summarize existing information regarding the anticancer activity of halophytes, the molecules involved in such properties, and the possible mechanisms of action (Table 1, Table 2, Table 3 and Table 4). The Web of Science database and Google Scholar (as a search engine) were consulted to retrieve the most updated articles on the topic under investigation. The keywords “salt-tolerant plants” and “halophytes” were used alone or in combination with, for example, “anticarcinogenic”, “antitumoral”, or “anticancer”. Only English articles having full text were considered. The contribution was distributed according to the different families and species, which were previously checked in the eHALOPH database [23]. The correspondence between the name of the cell lines referred in the text and the type of cancer can be found in Table 1, together with a list of halophyte species, organs and extracts tested, cell lines/types of cancer assessed, and obtained half-maximal inhibitory concentrations (IC50) values and proposed mechanisms of action, when provided by the authors. However, at the end of the study, a summary is presented referring only to the results obeying the criteria of cytotoxic activity for crude extracts (IC50 < 30 μg/mL; Table 3) and isolated compounds (IC50 ≤ 10 μM; Table 4) considered relevant to proceed for antitumoral applications, as established by the American National Cancer Institute (NCI) [24] and the National Institutes of Health (NIH) [25], respectively.

2. Anticancer Activity of Halophytes

Halophytes are a valuable source of bioactive molecules for the prevention and treatment of human chronic diseases, including cancer [26]. In this review, information is presented by family, namely, Acanthaceae, Aizoaceae, Amaranthaceae, Apiaceae, Asteraceae, Brassicaceae, Convolvulaceae, Cymodoceaceae, Cyperaceae, Fabaceae, Juncaceae, Malvaceae, Myrtaceae, Plantaginaceae, Plumbaginaceae, Poaceae, Portulacaceae, Rhizophoraceae, Solanaceae, and Tamaricaceae. Data regarding the anticancer activity of different halophyte species, including organs and extracts tested, cell lines/types of cancer assessed, and obtained IC50 values, are summarized in Table 1. Table 2 contains detected molecules in the active extracts from halophyte species with anticancer properties, while Table 3 summarizes information on extracts/fractions of selected species that could be further explored in the search for new drug-leads, as established by the American National Cancer Institute (NCI) [24], including cytotoxic activity and chemical composition of the extracts (when possible), cell lines tested, obtained IC50 values, and proposed mechanisms of action. Lastly, data related to the cytotoxic activity of selected isolated compounds, according to the criteria of the National Institutes of Health (NIH) [25], are presented on Table 4.

2.1. Acanthaceae Family

Acanthaceae currently comprises 192 genera and 5504 species growing in Indonesia, Malaysia, Africa, Brazil, and Central America [27,28]. The presence of bioactive metabolites such as alkaloids, phenols, terpenoids, tannins, quinones, cardiac glycosides, saponins, carbohydrates, and flavonoids, highlights the medicinal relevance of this family in the treatment of several diseases, including cancer [29]. This review focuses on the anticancer activity of the halophytic genera Acanthus and Avicennia.
Acanthus ilicifolius L. (holly leaved acanthus) is a mangrove shrub rich in triterpenes, alkaloids, and flavonoids, traditionally used in Chinese and Indian medicine against asthma, headache, and skin diseases [30]. An ethanolic leaf extract from A. ilicifolius significantly reduced the viability of HepG2 cells in a dose-dependent manner (92% of cytotoxicity at 100 μg/mL) via apoptosis induction potentially associated with DNA damage [31] (Table 1). A water root extract from A. ilicifolius reduced the viability of HepG2 cells, as observed in the 3-(4, 5-dimethylthiazol-2-yl)-2, 5-diphenyl tetrazolium bromide (MTT) assay (IC50 value of 39.76 μg/mL after 48 h of incubation) [32], via apoptosis induction related to DNA damage (Table 1). Ethanol extracts from leaves and roots of holly leaved acanthus were also able to reduce the viability of MCF7 cells (breast carcinoma), with IC50 values of 24.22 and 29.20 μg/mL for leaf and root extracts, and of PA1 cells (ovarian carcinoma), with IC50 values of 15.74 μg/mL and 20.00 μg/mL for leaf and root, respectively (Table 1) [33]. The chemical profile of the active extracts was not established. Acanthus ebracteatus Vahl. (holly mangrove), is traditionally used in Thai medicine as an anticancer agent [34]. A protein hydrolysate (<3 kDa) from the aerial parts of A. ebracteatus, reduced the viability of A431 cells, with an IC50 value of 425.9 ng protein/mL after 24 h of incubation (Table 1). Partially purified peptides were obtained from this hydrolysate and resulted in the synergistic effects against cell viability of A431 cells, via apoptosis induction [34].
Avicennia comprises mangroves distributed in the Indo-Western Pacific and Atlantic eastern Pacific areas. Some Avicennia species are effective against cancer due to the presence of different classes of molecules, including alkaloids, phenols, flavonoids, tannins, iridoid glucosides, and terpenoids [35]. A methanol leaf extract from Avicennia alba Blume significantly reduced the viability of MCF7 and HeLa cells, evaluated by the MTT assay, with IC50 values of 57.02 and 44.30 µg/mL, respectively, after 48 h of incubation [36] (Table 1). After treatment with the leaf extract of A. alba both cell lines exhibited morphological modifications, i.e., a reduction in cell size and cell detachment. The extract was analyzed using gas chromatography–mass spectrometry (GC–MS) allowing for the identification of 12 compounds, the most abundant ones being catechol borane (11%), neophytadiene (17%), and hexadecanoic acid (29%) (Table 2), but their cytotoxicity toward the tested cell lines was not established [36]. Moreover, since GC–MS mainly allows for the detection of lipophilic molecules, one cannot exclude the possibility that hydrophilic compounds with cytotoxic properties may be present in the methanol leaf extract but were not detected with the chromatographic system used. A chloroform/methanol extract from leaves of A. alba reduced the viability of WiDr cells (IC50 = 173.78 µg/mL after 24 h of incubation) using doxorubicin as the positive control and induced cell arrest in the G0–G1 phase, resulting in apoptosis induction [37] (Table 1).
Avicennia marina (Forssk.) Vierh, one of the most abundant and common mangrove species, is used in traditional medicine for the treatment of different diseases, including rheumatism and smallpox, and it contains different molecules with therapeutic properties, such as luteolin 7-O-methylether and chrysoeriol 7-O-glucoside [38]. An ethyl acetate extract of leaves and stems of A. marina reduced the viability of MCF7 cells (MTT assay), with values of cell growth inhibition of 65% and 75% after treatment for 48 h with the extract at 100 and 200 µg/mL, respectively [39] (Table 1). When applied at the concentration of 200 µg/mL the extract induced apoptosis in a concentration-dependent manner through ROS production and disruption of the mitochondrial membrane potential (Δψm); however, no PARP-1 cleavage was detected, and a significant decrease in caspase-7 protein levels was observed. A significant level of autophagy was also detected at the same concentration [39]. A methanol extract of the same species reduced the viability of HeLa cells in a concentration-dependent manner (tested concentrations: 50, 100, 250, 500, and 1000 µg/mL) with an IC50 value of 107 µg/mL [40] (Table 1). The chemical profile of the extracts was not established by the authors and, therefore, the bioactive compounds were not identified. In another study targeting the same species [38], water, ethanol, methanol, and ethyl acetate extracts from leaves and seeds were tested toward different human cancer cell lines, namely, breast (AU565, MDA-MB-231, and BT483), liver (HepG2 and Huh7), and NIH3T3 cells. The ethyl acetate extract from leaves had the highest cytotoxicity, especially toward cancer cells, which was attributed by the authors to its highest phenol content. The ethyl acetate extract suppressed xenograft MDA-MB-231 tumor growth in nude mice, and induced apoptosis in cancer cells through apoptosis inhibition and inhibition of cell migration. This extract was further fractionated by column chromatography, and resulting fractions were retested for anticancer properties. The most active factions, F2-5, F3-2-9, and F3-2-10, had IC50 values lower than 20 μg/mL, and they were profiled by 1H-NMR and 13C-NMR, allowing for the identification of the flavonoids luteolin and quercetin standards, and avicennones D and E. The antiproliferative activity of the flavonoids was confirmed [38]. Additional studies about the anticarcinogenic activity of other halophytic species of the Acanthaceae family can be found in Table 1.

2.2. Aizoaceae Family

Aizoaceae contains mostly succulent plants with 1910 species currently recognized, distributed across 125 genera [41,42]. The Mesembryanthemum genus, for example, is used in traditional medicine to treat several diseases such as hepatic conditions and diabetes, and it contains several bioactive compounds, including flavonoids and catechins [43]. An ethanol crude extract from leaves of Mesembryanthemum crystallinum L. (ice plant), and obtained hexane, ethyl acetate, butanol, and water fractions were tested for their effect on the growth of HCT116 cells [44] (Table 1). The application of the ethyl acetate and butanol fractions resulted in a dose-dependent inhibition of cell growth (at 125, 250, and 500 μg/mL, applied for 24 and 48 h), as observed in the MTT assay, a reduction in the levels of intracellular ROS (at 250 and 500 μg/mL, using 2′,7′-dichlorofluorescin-diacetate), and apoptosis induction (at 250 and 500 μg/mL, during 48 h). Treatment with the butanol fraction also resulted in cell-cycle arrest in the G2/M phase [44]. Samples were evaluated for total phenolic content (TPC), using the Folin–Ciocalteau (F–C) assay, and it was observed that the butanol fraction had the highest TPC (5.4 mg gallic acid equivalent/g) [44], suggesting that phenolics may be related to the detected cytotoxicity. However, no attempt was made by the authors to identify the active molecules present in this sample.
The application of ethanol, methanol, acetone, hexane, and diethyl-ether extracts from whole plants of Sesuvium portulacastrum L. (sea purslane) decreased the viability of MDA-MB-231, IMR32, and HCT116 cell lines, as assessed by MTT after 24 h of incubation, using doxorubicin as the standard [45]. The hexane extract had the highest cytotoxic activity with IC50 values of 942.07, 703.40, and 407.87 µg/mL for MDA-MDB-231, IMR32, and HCT116 cell lines (Table 1). Despite the considerably high IC50 values obtained, the possible mechanisms of action of the extracts were evaluated using phase contrast-inverted microscope observation and propidium iodide staining, which revealed the presence of typical features of apoptotic cells, including nuclear condensation, cell shrinkage, and the presence of apoptotic bodies [45] (Table 1). No attempt was made by the authors to unravel the chemical composition of the extracts.

2.3. Amaranthaceae Family

Amaranthaceae currently comprises 2398 species grouped in 182 genera [46], which can be found in a wide range of habitats from arid and semiarid regions to saline environments and subtropical areas; it contains flavonoids, phenolic acids, terpenes, and triterpene saponins, and it displays important pharmacological properties such as anticarcinogenic [47,48].
Arthrocnemum sp. is a small genus of succulent halophytes present in coastal marshes of South Africa, Asia, Mediterranean, Europe, and Northern America, and it contains several compounds such as phenols, flavonoids, and tannins with described therapeutic properties, including antibiotic, hypoglycemic, and antitumoral [49,50]. The application of an 80% methanol extract from shoots of Arthrocnemum indicum (Willd.) Moq. significantly reduced the viability of Caco-2 cells, as observed by the MTT assay [50] (Table 1). Fluorescence microscopy observation through DAPI (4,6-diamidino-2-phenylindole) staining showed that cells exhibited a decline of DNA synthesis, while flow cytometry allowed for the observation of cell-cycle arrest at the G2/M phase after 72 h of exposure to the extract at the concentration of 100 µg/mL [51] (Table 1). The extract was profiled by liquid chromatography/electrospray ionization time-of-flight mass spectrometry (LC/ESI-TOF-MS), and high levels of phenolic compounds were detected such as gallic acid, cyanidin, chrysoeriol, quercetin, catechol, syringic acid, and luteolin (Table 2), although their cytotoxicity toward Caco-2 cells was not established [51]. However, it is known that some of the detected compounds exhibit antitumoral properties. For example, chrysoeriol was identified as a selective inhibitor of the PI3K–AKT–mTOR pathway and, therefore, linked with cell cycle regulatory effects; it also reduced the proliferation of human multiple myeloma cells (RPMI 8226 and KM3, with IC50 values of 26 and 35 µmol/L at 48 h) but not of peripheral blood mononuclear cells (PBMNCs) [52].
Several Atriplex species are halophytic [53]. An ethanol extract from leaves of Atriplex halimus L. (sea orache) significantly reduced the viability of HepG2 cells (IC50 = 54.86 µg/mL after 24 h of incubation) (Table 1) via apoptosis induction linked to the expression of TP53, BCL2, and BAX genes [54] (Table 1). No information was provided by the authors regarding the chemical components of such extract, but it is known that A. halimus is characterized by the presence of different bioactive compounds, including syringetin derivatives and the flavonol glycosides atriplexoside A [3′-O-methylquercetin-4′-O-β-apiofuranoside-3-O-(6′′-O-α-rhamnopyranosyl-β glucospyranoside) and atriplexoside B [3′-O-methylquercetin-4′-O-(5′′-O-β-xylopyranosyl-β-apiofuranoside)-3-O-(6′′-O-α-rhamnopyranosyl-β-glucopyranoside)], which may be related to the detected cytotoxic activity [55,56,57,58,59,60].
Chenopodium contains weedy herbs native to Asia, Europe, and America, rich in phenolics, saponins, and triterpenoids, conferring them therapeutic properties such as laxative, analgesic, and anticarcinogenic [61]. The edible species Chenopodium album L. (lamb’s quarters) is valued in traditional medicine for its anticancer properties [62]. A petroleum ether extract of branches and leaves from C. album reduced the viability of A549 cells in a dose-dependent manner, showing an IC50 of 33.31 μg/mL using gemcitabine as a positive control, which was linked to cell-cycle arrest at the G1 phase [63] (Table 1). Essential oil from whole plants of Chenopodium ambrosioides L. (Mexican tea) decreased the viability of MCF7 cells by inducing DNA fragmentation (IC50 values of 18.75, 9.45, and 10.50 μg/mL at 6, 24, and 48 h, respectively) when compared to the control treatment [64] (Table 1). In another study, an essential oil of the same species also displayed a significant cytotoxic activity toward RAJI cells, with an IC50 value of 1.0 μg/mL, which was probably related to the high levels of ascaridole, detected using GC with flame ionization detection (GC-FID), GC–MS, and proton nuclear magnetic resonance (1H-NMR) [65] (Table 1 and Table 2). Ascaridol is a monoterpene with a strong in vitro capacity to decrease the growth of different tumor cell lines and is, therefore, considered as a strong candidate for the treatment of cancer [65]. Chenopodium quinoa Willd is a pseudocereal originated from the Andes of South America with recognized nutritional and functional properties that rendered the species the status of “functional food” [66]. Polysaccharides were extracted from quinoa seeds with petroleum ether and purified by column chromatography, revealing mainly galacturonic acid and glucose monosaccharides. This polysaccharide fraction was applied toward cancer (SMMC 7721 and MCF7) and “normal” (L02 and MCF 10A) cell lines, for 24 and 48 h, allowing for IC50 values ranging from 53.4 to 121.4 μg/mL, without affecting “normal” cell viability [66] (Table 1).
Salicornia species (glasswort, sea asparagus, or samphire) are edible succulent halophytes highly appreciated in gourmet cuisine due to their organoleptic properties [67]. The ethyl acetate and methanol leaf extracts from Salicornia europaea L. reduced the viability of MCF7 cells with IC50 values of 97.9 and 117.1 μg/mL for the ethyl acetate and methanol extracts, respectively, after 24 h of incubation [68]. A qualitative analysis of the extracts identified tannins and saponins, which may be associated with the in vitro antitumor capacity. Moreover, a GC–MS analysis showed the presence of 32 and 29 compounds in corresponding ethyl acetate and methanol extracts, respectively, which were not assessed individually against the cancer cells [68]. Tannins are phenolic compounds with a high molecular weight, while saponins contain a steroidal or triterpenoid aglycone and one or more sugar chains [69]. Both groups of compounds exhibit antitumoral properties; for example, saponins generally decrease cellular proliferation by binding to cholesterol structures on cell membranes, forming pores and holes by binding to specific receptors followed by the induction of apoptosis, while tannins can hamper cancer cell proliferation through antioxidant processes and apoptosis induction [70,71]. However, the presented results must be analyzed with caution, since the phytochemical analysis reported by the authors only targets lipophilic compounds, without unraveling the possible cytotoxic molecules present in the methanol (polar) extract.
Suaeda comprises around 80 to 100 succulent species distributed in semideserts, deserts, and seashores. These species are usually edible and contain several bioactive components, such as phenolics, flavonoids, and terpenoids, conferring them with bioactive properties, including anticarcinogenic [72,73]. A hexane extract from leaves of Suaeda ruticose (L.) Forssk. (shrubby seablight) reduced the viability of HCT116, HepG2, and MCF7 cell lines, being more effective toward HCT116 cells (IC50 = 17.2 µg/mL) [74] (Table 1 and Table 4). The cytotoxicity of the extract was tested using the sulforodamine B (SRB) assays for 72 h using doxorubicin as a positive control with IC50 values of approximately 0.5 µg/mL. This extract caused cell-cycle arrest at the G0–G1 phase and induced apoptosis, especially in HCT116 cells that exhibited chromatin condensation and membrane blebbing (Table 4). The extract was profiled using liquid chromatography coupled to electrospray ionization quadrupole time-of-flight mass spectrometry (LC–ESI-QTOF-MS/MS) and several molecules were identified, including monoterpenes (dihydrojasmone, jasmolone, terpinene-4-ol), diterpenes (pimaric acid, steviol, and momilactone B), and phenolics (quercinol, zingerone, zingerol, and neovaflan) (Table 4), but its contribution to the detected cytotoxicity was not directly established [74]. However, there are several reports on the antitumoral properties of phenolic compounds [56,57,58,59]. Specifically, zingerone reduced the viability of HCT116 cells through ROS-mediated apoptosis in colon cancer cells [75]. Monoterpenes and diterpenes have reported cytotoxic properties [76,77]. For instance, dihydrojasmone was identified as one of the active compounds toward HeLa and NIH/3T3 cell lines, in a methanol extract from Rumex hastatus D. Don [78], while steviol significantly reduced the viability of MCF7 cells via apoptosis induction [79]. A dichloromethane extract from the shoots of S. fruticosa significantly reduced the viability of A549 (IC50 = 49 µg/mL), DLD1 (IC50 = 10 µg/mL), Caco-2 (IC50 = 140 µg/mL), and HT-29 cells (IC50 = 12 µg/mL), which was assessed using the resazurin reduction test after 48 h of incubation (Table 1 and Table 4). Nevertheless, the bioactive compounds present in the chemical profile of the extracts were not identified [80]. Additional studies on the anticarcinogenic activity of other halophytic species of this family are summarized in Table 1.

2.4. Apiaceae Family

Apiaceae currently comprises 3916 species distributed across 457 genera dispersed worldwide [81], and it is characterized by the presence of polyacetylenes displaying important bioactivities, including anticarcinogenic [82]. In this family, there are two genera of halophytic species, Eryngium sp. and Crithmum sp., with described cytotoxic properties against human cancer cells.
Eryngium includes approximately 250 species present in Eurasia, North Africa, North and South America, and Australia. Eryngium species are frequently used as ornamentals or as a food source and are valued in traditional medicine as a diuretic, to treat diarrhea, headaches, and digestive problems. These properties are potentially associated with the presence of terpenoids, triterpenoids, polyacetylenes, flavonoids, and coumarins [83]. The cytotoxic effects of aqueous extracts from shoots and roots of Eryngium maritimum L. were assessed against human cancer lines (HepG2, HEP-2, and U138MG) and on a “normal” cell line (Vero), using the MTT assay after 24 h of incubation [70] (Table 1). The root extracts were overall more cytotoxic, and the IC50 values ranged from 30.3 to 50 µg/mL. However, a strong reduction in cellular viability was also observed in the “normal” cell line, which suggest no selectivity toward cancer cells [70] (Table 1). Although the phytochemical profile of the extracts was not established by the authors, other reports identified several cytotoxic molecules in E. maritimum, including saponins, which create pores and perforations on the cellular membranes by binding to specific receptors, followed by apoptosis induction by, for example, stimulation of the cytochrome c-caspase 9-caspase 3-pathways [70,71].
Crithmum maritimum L. (sea fennel) is traditionally used as a cooking ingredient and in folk medicine for its stimulating and diuretic effects [84]. An ethyl acetate extract from the whole sea fennel plant reduced the proliferation of Huh7 and HepG2 cell lines, by interfering with the cell cycle, specifically on the shift of phases with increasing number of cells in the G0/G1 phase after 24 h of incubation and in the G2/M phase after 48 h [85]. The extract induced an increase in necrotic and apoptotic cancer cells, assessed by cytofluorimetric analysis although the values of IC50, but the chemical profile of the extract was not assessed [85] (Table 1).

2.5. Asteraceae Family

Asteraceae currently contains 1733 genera and 35,988 species [86], including several used as ornamentals (e.g., Calendula and Chrysanthemum) or for food (e.g., Helianthus annuus) and medicinal (e.g., Artemisia sp. and Echinacea sp.) purposes [87]. Achillea millefolium L. (yarrow) is an aromatic perennial herb with traditional medicinal uses such as wound healing and anti-inflammatory activity, and it contains different classes of bioactive compounds, including flavonoids and terpenoids [88]. A methanol extract from yarrow shoots was applied toward prostate cancer (DU145) and “normal” skin (HFFF2) cells, at different concentrations (20, 100, 500, 1000, and 2000 µg/mL), alone or in combination with bleomycin, an anticancer agent. The yarrow extract was not toxic toward “normal” cells, but had significantly enhanced cytotoxicity induced by bleomycin showing 60% and 49% survival rate at doses of 1000 and 2000 µg/mL, respectively, which may indicate that this extract contains molecules able to improve the effectiveness of bleomycin, while minimizing negative side-effects caused by toxicity toward “normal” cells. [88] (Table 1). The extract was not profiled for chemical components, but there are reports of the presence of several cytotoxic compounds in yarrow, such as achillinin A (guaianolide) and casticin (flavonoid) [88,89]. In another work, petroleum ether, ethyl acetate, methanol, and water extracts were prepared from aerial organs of yarrow and tested for toxicity, using the MTT assay, on human cancer cell lines [90] (Table 1). The strongest cytotoxic effect was observed after application of the ethyl acetate extract on HeLa (IC50 = 0.58 μg/mL) and K562 cells (IC50 = 0.73 μg/mL), followed by the water (MCF-7, IC50 = 0.87 μg/mL) and the petroleum ether extract (K562, IC50 = 0.87 μg/mL) (Table 1 and Table 4) [90]. The cytotoxic activity of these extracts may be associated with its contents in phenolic acids, such as apigenin and chlorogenic, p-coumaric, and rosmarinic acids (Table 4), which showed the capacity to block oncogenic pathways due to the activation of caspases [91]. A methanol extract from aerial parts of Limbarda crithmoides (L.) Dumort (commonly known as Inula crithmoides L.) was able to decrease the viability of acute myeloid leukemia cells (OCI-AML3) when applied at 100 and 200 μg/mL, for 24 h [92] (Table 1). This extract was submitted to a solvent–solvent partitioning, affording n-hexane, dichloromethane, and aqueous methanol-soluble fractions. The hexane and dichloromethane fractions exhibited a strong cytotoxicity toward OCI-AML3 cells at concentrations of 15 or 10 µg/mL, which was ascribed to an increase in apoptotic cells, especially in the G0/G1 phase by the mitochondria-dependent pathway. The hexane extract was then further fractionated, leading to the isolation of two molecules, which were identified by NMR as the thymol derivatives 10-acetoxy-8,9-epoxythymol tiglate and 10-acetoxy-9Z-chloro-8,9-dehydrothymol, with the latter being the most active, causing a decrease in cell viability at 1.25 µg/mL associated with apoptosis induction [92].

2.6. Brassicaceae Family

Brassicaceae is one of the largest dicotyledon family of flowering plants, including model species and commercial crops, with 341 genera and 3921 species recognized at the moment [93,94], but few studies have described the potential anticarcinogenic properties of halophytes belonging to this family. One example is the annual halophyte Cakile maritima Scop. (sea rocket), which is confined to maritime strandlines of sand and has agronomic (oilseed and phytoremediation) and medicinal (diuretic, antiscorbutic, and purgative) properties [95]. Hexane, ethyl acetate, and methanol fractions were obtained from a methanol extract prepared from aerial organs of sea rocket and tested for antiproliferative properties on Caco-2 and HeLa carcinoma cells [96] (Table 1). The hexane fraction significantly reduced the viability of Caco-2 and HeLa cells, with IC50 values of 12 and 126 μg/mL after 24 h of incubation; respectively. Cisplatin was used as the positive control with values of 69 μg/mL (Caco-2) and 85 μg/mL (HeLa). The extract was profiled using GC–MS, and the major molecules identified were, by area, 2-hydroxy-1,8-cineole, decane, and limonene, which may contribute to the detected antiproliferative activity [96] (Table 4). For instance, limonene has been reported with anticarcinogenic activity in HepG2 cells due to apoptosis induction [97].

2.7. Convolvulaceae Family

Convolvulaceae currently contains 50 genera and 1952 species widely distributed in tropical and temperate regions of the world [98,99]. Calystegia soldanella (L.) R. Br. ex Roem. & Schult. (shore bindweed) is a perennial edible herb commonly found in coastal sand dunes and foredunes of South Korea, East Asia, Europe, and the Pacific. Shore bindweed is traditionally used for the treatment of, for example, rheumatic arthritis and scurvy, and it displays relevant biological properties, including anticancer [100]. An 85% aqueous fraction from combined methylene chloride and methanol crude extracts obtained from whole plants of shore bindweed significantly decreased HepG2 cellular viability after 24 h of incubation, in a concentration-dependent manner, via cell-cycle arrest at the G0–G1 and S phases and apoptosis induction [100]. In another study, a methanol crude extract from whole plants of the same species exhibited a potent cytotoxic activity toward A549 cells (human lung cancer) and Col2 cells (human colon cancer), with IC50 values of 8.0 µg/mL and 27.4 µg/mL, respectively [101]. A hydroalcoholic extract from shoots of Cressa cretica L. decreased the viability of HepG2 cells (IC50 value = 2300 µg/mL after 72 h of incubation), by increasing the expression of the proapoptotic protein BAX in detriment to antiapoptotic proteins (BCL2) [102].

2.8. Cymodoceaceae Family

This family presently contains six genera and 17 species of seagrasses [103,104]. A water leaf extract from Cymodocea rotundata Ehrenb Hempr. ex Aschers. was supplemented with silver nitrate (AgNO3, 1 M), to produce silver nanoparticles (AgNPs). AgNPs have several medical applications, including coating of medical devices and wound dressings, and they exhibit cytotoxic activity [105]. The obtained AgNPs exhibited high cytotoxicity toward MG63 cells with an IC50 value of 25.31 µg/mL after 48 h of incubation [105]. AgNPs were also produced by combining a water leaf extract of Cymodocea serrulata (R. Br.) Aschers. & Magnus and AgNO3 (1 M), and they reduced the viability of A549 cells in a direct dose–response manner (IC50 = 100 µg/mL after 24 h of incubation). The higher cytotoxicity of the AgNPs was ascribed to the easy permeability to the cellular barriers and their high affinity to biological macromolecules, as well as their capacity to release ROS [106]. No attempt was made to identify the active molecules present in those samples [105,106]. A hydroethanolic extract from shoots of the same species inhibited the proliferation of HepG2 cells (IC50 value of 82.92 µg/mL after 24 h of incubation with camptothecin as a positive control with an IC50 value of 8 µg/mL) [107]. The extract had a high concentration of tannins, flavonoids, and terpenoids, with described antitumoral properties [108], but no attempt was made by the authors to identify the possible bioactive molecules [107].

2.9. Cyperaceae Family

Cyperaceae currently comprises 92 genera and 5888 species with a cosmopolitan distribution [109,110]. Cyperus rotundus L. (nut grass) is a perennial halophyte species traditionally used in the treatment of several pathologies, such as stomach disorders [111]. Methanol, ethanol, and water extracts from nut grass rhizomes were tested toward MDA-MB-231 cells, for 24 h [112] (Table 1). The highest reduction in cell viability was obtained with the ethanol extract, with an IC50 value of 225 µg/mL, through apoptosis induction via upregulation of the death receptor 4 (DR4), DR5, and proapoptotic BAX, and downregulation of antiapoptotic survivin and BCL2 [112]. The active extract was not chemically profiled, but it is known that the rhizomes of nut grass are rich in several bioactive molecules, including flavonoids, tannins, and sesquiterpenes [113], with reported cytotoxic activity against cancer cell lines.

2.10. Fabaceae Family

This family currently comprises 778 genera and 22,356 species [114] and includes five genera of halophytic species, namely, Alhagi, Glycyrrhiza, Melilotus, Prosopis, and Sesbania, with described anticarcinogenic properties [115].
The genus Alhagi is distributed throughout Asia, Australia, and Europe and used traditionally for the treatment of, for example, gastroenteritis, ulcers, and rheumatoid arthritis [116]. Alhagi maurorum Medik (camelthorn) is used for its anti-inflammatory properties, which are ascribed to the presence of the triterpenoid lupeol [117]. In fact, lupeol was isolated from a methanol extract of camelthorn aerial parts, and it decreased the cellular viability of MCF7 and MDA-MB-231 cell lines, with one-fourth of IC50 values >100 µg/mL (Table 1) [118]. The cytotoxic activity of lupeol was lower than that observed with its epoxide form, and it was related to the increase in mRNA expression levels of apoptosis-related genes (TP53, caspase-3 and BAX) and decrease in BCL2 gene expression [118].
Glycyrrhiza (liquorice) contains legumes endogenous to Asia and southern Europe with reported anti-inflammatory and antiviral properties [119]. There are two Glycyrrhiza halophytic species with anticarcinogenic activity: Glycyrrhiza glabra L. and G. uralensis Fisch. Glycyrrhiza glabra is rich in phenolics, tannins, and especially glycyrrhizin, a triterpenoid saponin [120]. Glycyrrhizin reduced the proliferation of HeLa cells at 320 µM after 24 and 48 h of incubation, via apoptosis, through mitochondrial depolarization. Moreover, nuclear condensation, cell membrane lysis, and disintegration of organelles were observed in treated cells through phase-contrast microscopy [121]. The cytotoxic potential of the roots’ methanol extracts from G. glabra, collected in nine different areas from Italy, Turkey, Syria, Russia, Afghanistan, and Uzbekistan, was assessed against “normal” (HaCaT) and cancer cell lines (A549 and HepG2) using the MTT assay, after 24 h of incubation [122]. Results disclosed variable cytotoxicity levels depending on the samples’ collection location and season, potentially related to the influence of climatic conditions on the chemical composition of the plants. Only one sample from Afghanistan was active toward HepG2 cells (IC50 of 248.5 µg/mL), while four extracts (from Italy, Afghanistan, and Syria) were cytotoxic toward A549 cells (IC50 between 189.1 and 238.9 µg/mL) [122] (Table 1). However, samples were also cytotoxic toward “normal” cells (HaCat) (Table 1). An ethanol root extract of G. glabra significantly reduced the proliferation of HT29 cells, at 200 µg/mL after 24, 48, and 72 h of incubation as detected by the MTT assay [123]. Polymerase chain reaction (PCR) studies revealed a downregulation of heat-shock protein 90 (HS90) gene expression that can be related to the reduction in cellular viability, since the HSP90 prevents tumor cells from undergoing apoptotic death; therefore, its blocking could assist active antitumor effects [124]. A reduction in HeLa cell viability above 80% was reported after application of an aqueous ethanol extract of G. uralensis rhizomes at the concentration of 1.84 mg/mL, but the determination of the IC50 value was not reported [125]. Several cytotoxic compounds were identified in that extract using UPLC–ESI-Q-TOF, including isoquercitrin, 4′demethylpodophyllotoxin glucoside, and podophyllotoxin, all with described cytotoxic properties [125] (Table 2).
Melilotus contains 19 species of annual herbs widely distributed in North Africa and Eurasia [126]. Melilotus indicus L. All. (sweet clover), the only described halophyte within this genus, is used in traditional medicine as an analgesic, and it has reported cytotoxic activity [127,128]. A methanol extract from the aerial parts of sweet clover significantly decreased cell proliferation of HepG2 (IC50 = 16.60 µg/mL) and SNU-182 cells (IC50 = 13.21 µg/mL) after 24 h of incubation, using staurosporine as a positive control (Table 1) [127,128]. The extract was less cytotoxic toward L-02 cells (human “normal” hepatic) (IC50 = 90.9 µg/mL) (Table 1) [127,128]. The application of the extract resulted in an increase in the number of apoptotic cells and loss of mitochondrial membrane potential (Δψm) [128] (Table 1). The chemical composition of the extract was not established by the authors.
Prosopis comprises 44 species, mainly small trees distributed in dry lands of America, Africa, and Asia. Several Prosopis species contain anti-inflammatory, antidiabetic, and anticancer compounds, namely, flavonoids, tannins, phenolics, and alkaloids [129]. Prosopis juliflora Sw. DC. (mesquite) is an invasive species in India with reported ethnomedicinal uses, for the treatment of eye and digestive disorders, to name a few [130]. Mesquite extracts display relevant bioactivities, including anti-inflammatory, ascribed mainly to its content in alkaloids [130]. A leaf methanol extract from Prosopis juliflora Sw. DC. (mesquite) had cytotoxic effects on Molt-4 cells, with IC50 values of 90.5, 42.5, and 20.0 μg/mL after 24 h, 48 h, and 72 h of incubation, respectively, using mitomycin-C (6 µg/mL) as a positive control (Table 1). The extract was less toxic toward “normal” cells (mitogen stimulated T-lymphocyte cultures from peripheral human blood) [130]. A genotoxic assessment using a cytokinesis-block micronucleus assay reported that the number of micronuclei showed an increasing pattern with the application of increasing concentration of the extract [130].
The genus Sesbania comprises 60 to 85 species of herbs, shrubs, and trees distributed mostly in the tropical and subtropical regions of Africa, Asia, Australia, and America. Sesbania species have relevant pharmacological properties such as anti-inflammatory and antidiabetic [131,132,133]. Sesbania grandiflora (Akatti) (sesbania or agathi), is a small perennial tree with high levels of vitamins and minerals associated with anti-inflammatory, analgesic, and antipyretic properties [134,135], and the anticancer properties of its fruit are mentioned in the Ayurvedic literature [136]. Water, ethanol, and acetone leaf extracts of S. grandiflora reduced the viability of IMR32 and HT29 cells with IC50 values of 200 µg/mL after 24 h of incubation using doxorubicin as a control [137]. However, the chemical composition of the extract was not unraveled by the authors. Additional studies regarding the anticarcinogenic activity of other halophytes of this family are depicted in Table 1.

2.11. Juncaceae Family

Juncaceae currently contains eight genera and 522 species [138] of herbs adapted to salty marshes or badly drained soils and accumulate different phytoconstituents such as flavonoids, triterpenes, steroids, and phenolic acid derivatives [139,140]. A phenanthrene, juncunol (1,7-dimethyl-5-vinyl-9,10-dihydrophenanthren-2-ol), was identified in a diethyl ether extract of leaves of Juncus acutus (L.) Torr. ex Retz. (spiny rush) and displayed selective in vitro cytotoxicity toward HepG2, HeLa, and MDA-MB-468 cell lines [141]. Juncunol had an IC50 value of 18 µM in HepG2 cells after 72 h of incubation, determined using the MTT assay, and it induced an increase in the number of apoptotic cells in a concentration-dependent manner (IC50 value ± 25%) accompanied by a decrease in the Δψm [141,142]. Juncunol induced cell-cycle arrest in the G0/G1 phase, while showing no hemolytic properties. In silico studies indicate that that compound seems to bind between GC base pairs and, thus, may act as a DNA intercalator [142].

2.12. Malvaceae Family

This family of flowering plants currently contains 243 genera and 5461 species [143]. A leaf decoction from Thespesia populnea L. Sol. ex Corrêa (portia tree) had a high content of total phenols and flavonoids and showed cytotoxic and antiproliferative properties toward HEP-2 cells, as observed in the MTT (IC50 = 120.02 µg/mL after 24 h of incubation) and SRB (IC50 = 77.06 µg/mL) assays [144]. Treated cells showed apoptotic characteristics including membrane blebbing, cell shrinkage, nuclear and cytoplasmic condensation, and formation of apoptotic bodies [144]. However, no attempt was made to identify the active molecules present in this sample. The chloroform-soluble fraction of a methanol bark extract of portia tree caused a reduction in cellular viability of MDA-MB-231 and MCF7 cells, with IC50 values of 23.97 and 20.62 µg/mL respectively, after 24 h of incubation [145]. Chemical analysis of the extract using GC–MS revealed the presence of steroids such as cis-androsterone and fatty-acid derivatives, which display cytotoxic activity against cancer cell lines [108,145].

2.13. Myrtaceae Family

Myrtaceae presently includes 134 genera and 6614 species [146]. Focus is given to the genus Eucalyptus, especially Eucalyptus camaldulensis Dehnh, because is a salt-tolerant species with reported antitumoral properties. Eucalyptus spp. are mainly cultivated for timber and paper production, but the presence of bioactive compounds, such as triterpenoids, flavonoids, and tannins, confers E. camaldulensis with relevant bioactivities, including antitumoral [147]. Methanol, ethyl acetate, n-butanol, and water extracts from leaves of E. camaldulensis reduced the viability of MCF7 and MDA-MB-231 cell lines, according to the MTT and SRB assays [148]. The ethyl acetate extract had the highest cytotoxicity with IC50 values of 26.7 and 7.9 µg/mL for MTT and SRB in MCF7 cells, and IC50 values of 34.4 and 4.9 µg/mL for MTT and SRB in MDA-MB-231 cells after 24 h of incubation [148]. No attempt was reported by the authors to disclosure the molecules responsible for the detected cytotoxic activity.

2.14. Plantaginaceae Family

This family currently contains 101 genera and 2165 species [149], widely distributed worldwide but with preference for temperate zones [150]. Bacopa monnieri (L.) Wettst (water hyssop) is a wetland macrophyte and contains bacopaside (II) (saponine), with reported anticarcinogenic properties [151,152]. The anticarcinogenic properties of artificially digested (artificial saliva and artificial gastric juice) methanol extracts from B. monnieri reduced the motility (capacity of cell migration in a process of cancer development) of DU145 cells [153]. Nevertheless, the IC50 value was not determined by the authors. The extracts, analyzed by HPLC, were rich in phenolic compounds such as chlorogenic, caffeic, and syringic acids and bacoside A, with reported anticarcinogenic properties [154,155,156,157].
Within the Plantaginaceae family, the Plantago genus is the largest with 275 annual and perennial species widespread around the world, some with traditional uses as antipyretic, anti-inflammatory, and antitumoral agents [158]. Plantago major L. (fleawort) is an important medicinal plant due to its richness in bioactive metabolites, such as alkaloids and flavonoids, especially luteolin-7-O-β-glucoside, with antitumoral properties [159]. Silver particles produced by combining a water extract from P. major seeds and silver nitrate (AgNO3, 0.1 M) significantly decreased the viability of MCF7 cells with an IC50 value of approximately 12 µg/mL after 24 h of incubation [159]. However, no attempt was made by the authors to identify the bioactive compounds present in this extract. Information targeting other Plantaginaceae species is summarized in Table 1.

2.15. Plumbaginaceae Family

Plumbaginaceae encompasses several species adapted to survive under saline conditions and currently includes 1138 species distributed across 26 genera [160]. The genus Plumbago contains 18 species (shrubs or perennial herbs) characterized by the presence of flavonoids, phenols and saponins, conferring several properties including anticarcinogenic [161,162]. The genus Limoniastrum Heist. ex Fabr. comprises the halophytic species Limoniastrum monopetalum (L.) Boiss. (≡ Statice monopetala L.) and Limoniastrum guyonianum Boiss. [163,164], thriving in coastal and saline dry areas of the Mediterranean and northern Saharan Africa [165,166,167,168]. Limoniastrum guyonianum is traditionally used in the treatment of gastric infections [169]. A gall aqueous extract from L. guyonianum reduced the viability of HeLa cells with IC50 values of 170 and 140 μg/mL after 24 and 48 h of incubation, respectively [169]. This extract was rich in flavonoids and induced DNA hypomethylation and apoptosis due to its capacity to arrest cell-cycle progression in G2/M (Table 1). Due to the high levels of polyphenolic compounds detected in the extract, the antiproliferative and proapoptotic effects of such metabolites on HeLa cells were hypothesized by the researchers [169]. In another study, hexane, dichloromethane, ethanol, and methanol extracts from L. densiflorum (Guss.) Kuntze shoots were tested for cytotoxicity on cancer and “normal” cell lines using the resazurin reduction assay, after 48 h of treatment (Table 1) [170]. The dichloromethane extract was the most bioactive sample, with IC50 values of 29 and 85 μg/mL toward A-549 and DLD-1, without significantly reducing the viability of a human skin fibroblast cell line (Table 1). The major compounds present in that extracts were identified using RP-HPLC as trans 3-hydroxycinnamic acid, myricetin and isorhamnetin, which may be related to the observed cytotoxic activity [170].

2.16. Poaceae Family

This grass family currently includes 11,917 species divided across 796 genera [171,172]. In this family, only two genera of halophytes are reported with anticarcinogenic properties, namely, Cynodon and Echinochloa. The most representative Cynodon species is Cynodon dactylon L. Pers (Bermuda grass), a weed with several medicinal properties, such as antidiabetic, diuretic, and purifying, as well as in vitro cytotoxic properties [173]. The in vitro antitumoral properties of petroleum ether, dichloromethane, acetone, methanol/water (3/1), and water extracts from Bermuda grass were appraised on a breast cancer cell line, and cell viability was assessed by evaluating (3H)-hypoxanthine incorporation after 48 h of incubation [174] (Table 1). The highest cytotoxicity values were observed after application of the water, acetone, and petroleum ether extract, with IC50 values of 57.2, 38, and 39 μg/mL (Table 1) [174]. The acetone and ether petroleum extracts were profiled using LC–MS, allowing for the identification of several bioactive anthocyanins, namely, delphinidin, petunidin, malvidin, and cyanidin glucosides (delphinidin-3-O-acetylglucoside, petunidin-3-O-caffeoylglucoside-5-O-glucoside, petunidin-3-O-coumarylglucoside-5-O-glucoside, malvidin-3-O-monoglucoside, delphinidin-3-O-acetylglucoside-pyruvic acid, petunidin-3-O-acetylglucoside-5-O-glucoside, and cyanidin-3,5-O-diglucoside) [174] (Table 2). There is no information related to the specific anticancer properties of the detected anthocyanins; however, several others, such as delphinidin, cyanidin, malvidin, and corresponding glycosides, exhibit antitumoral properties in different cell lines [174]. An ethanol extract (70%) from grains of Echinochloa crus-galli (L.) P. Beauv. (cockspur grass) significantly reduced the viability of HCT-116 and HeLa cell lines, with IC50 values of 11.2 and 12.0 μg/mL, respectively [175] (Table 1). The active ethanolic crude extract was fractionated using n-hexane, chloroform, ethyl acetate, and n-butanol, with the ethyl acetate fraction exhibiting the lowest IC50 value (3.8 μg/mL). Fractionation of this active fraction allowed the isolation and identification of eight phenolic compounds, which exhibited significant cytotoxic activities toward the tested cells [175]. The most active molecules were 5,7-dihydroxy-3′,4′,5′-trimethoxy flavone and 5,7,4′ -trihydroxy-3′,5′-dimethoxy flavone (tricin), the latter with IC50 values of 7.2, 8.6, 10.8, and 19.9 μM against HepG2, HeLa, HCT116, and MCF7 cell lines, respectively, with similar results to the commercial anticancer drug, doxorubicin [175]. Further data on the antitumoral properties of other species of the Poaceae family can be found in Table 1.

2.17. Portulacaceae Family

This family contains one genus with presently 150 species [176] distributed worldwide, including the edible Portulaca oleracea L. (purslane), a halophyte succulent annual plant with important nutraceutical and antioxidant properties [177,178]. The anticarcinogenic activity of this species can be ascribed to the presence of several compounds, such as portulacerebroside A, portulacanones B, and 2,2′-dihydroxy-4′,6′-dimethoxychalcone [179] (Table 2). Portulacerebroside A induced apoptosis in HCCLM3 cells via activation of the p38 MAPK and JNK-triggered mitochondrial death pathway [180]. An ethanol extract from purslane seeds reduced the viability (MTT assay) of HepG2 cells in a concentration-dependent manner with IC50 values of approximately 75 µg/mL after 24 h of incubation via apoptosis induction according to cellular morphology modification, i.e., a reduction in cellular size and adhesion capacity (Table 1) [181]. Oil from seeds of the same species reduced the proliferation of HepG2 and A-549 cancer cell lines, at concentrations ranging from 250 to 1000 µg/mL as observed using the MTT and neutral red uptake assays, after 24 h of incubation in both cancer cell lines [182]. However, the high concentrations of the extract used (up to 1000 µg/mL) must be highlighted; even at 100 μg/mL, no reduction in cell viability was observed (Table 1) When treated with the highest concentration, cells exhibited morphological modifications typical of apoptotic cells, such as loss of cell adhesion capacity, shrinkage, and round shape [182]. However, the chemical profile of the extract was not determined by the authors.

2.18. Rhizophoraceae Family

This family includes mangroves and comprises currently 148 species and 15 genera [183,184]. It has two halophytic species with reported cytotoxic properties against human cancer cells, namely, Bruguiera gymnorhiza (L.) Lam (black mangrove) and Ceriops tagal (Pers.) C. B. Rob. (oriental mangrove) [185,186]. A methanol extract from black mangrove stem bark reduced the proliferation of HeLa, RAJI, and myeloma cell lines, with IC50 values of 133, 504, and 384 µg/mL after 24 h of incubation using doxorubicin as a standard control, via apoptosis induction, since DNA fragmentation was detected by fluorescence with double staining (ethidium bromide–acrydine orange) [187].

2.19. Solanaceae Family

Solanaceae contains 100 genera and 2925 species currently recognized [188], many of them with a high worldwide economic importance as cultivated crops, such as Solanum lycopersicum L. (tomato) [189]. The number of halophytic species with anticarcinogenic activity in this family is scarce, and reports have mainly focused on Lycium barbarum L. (Chinese wolfberry). Chinese wolfberry’s dried fruits are widely used in traditional Chinese medicine for its anticancer properties, potentially associated with the presence of different bioactive compounds, including polysaccharides (proteoglycans), scopoletin (phytoalexin), and 2-O-β-D-glucopyranosyl-L-ascorbic acid (vitamin C analog) [190]. A polysaccharide-rich aqueous extract from Chinese wolfberry fruits reduced the proliferation of SW480 and Caco-2 cell lines, as observed by the crystal violet and MTT assays, with IC50 values of 600 and 650 µg/mL respectively, after 4 days of incubation, with a high capacity to arrest cell-cycle progression in the G0/G1 phase [191]. An ethanol extract from the dried fruit residue of this species was cytotoxic toward A431 cells, with an IC50 value of 873.7 µg/mL after 24 h of incubation, determined by water-soluble tetrazolium-8-[2-(2-methoxy-4-nitrophenyl)-3-(4-nitrophenyl)-5-(2,4-disulfophenyl)-2Htetrazolium] monosodium salt (CCK-8) assay [192]. The same authors conducted in vivo studies in mice submitted to an intraperitoneal injection of 50 mg/kg of the crude extract, reporting a lower volume and weight of the formed tumor and a higher survival rate of tumor-bearing nude mice [192]. Several phenolic amides were isolated from an ethanol extract from the stem of L. barbarum and tested toward glioma cell lines, using the MTS method [180]. 4-O-Methylgrossamide and grossamide exhibited the highest capacity to decrease the viability of GSC-3# and GSC-12# cell lines, especially the latter compound, with IC50 values of 6.40 and 5.85 µg/mL respectively. [193]. The precise mechanism of action was not appraised for the isolated compounds; however, there are reports that phenolic amides can inhibit tumor necrosis factor-alpha (TNF-α) and nuclear factor-kappa B (NF-κB), which may be related to the detected cytotoxic effect [194].

2.20. Tamaricaceae Family

Tamaricaceae contains five genera with currently 110 species [195], widely distributed around the world, and several are halophytic [196,197]. Tamarix plants (salt cedar) contain several bioactive metabolites such as methylferulate and syringic acid with antineoplastic properties [198]. Syringic acid was isolated from a methanol extract of Tamarix aucheriana (Decne.) Baum aerial parts and was cytotoxic toward SW1116 and SW837 cells in a time- (24–72 h of incubation) and dose-dependent manner, with IC50 values of 0.95 and 1.2 mg/mL, respectively [199]. Cytotoxicity was related to an increase in the percentage of apoptotic cells, attributed to the antimitogenic effect of syringic acid [199]. Methylferulate was isolated from the same type of extract and significantly reduced the viability of SW1116 and SW837 cells, with IC50 values of 1.7 and 1.9 mM after 24 h of incubation, respectively, being responsible for cell-cycle arrest in the S and G2/M phases and apoptosis induction, together with a reduction in NF-κB DNA-binding activity [200].
Tamarix gallica L. is traditionally used as an expectorant, antidiarrheal, and laxative, potentially attributed to the presence of ellagic and gallic acids, anthocyanins, tannins, flavonones, isoflavonones, and resveratrol in its leaves [201]. A methanol extract from shoots, leaves, and flowers of T. gallica significantly reduced the viability of Caco-2 cells, related to a decrease in DNA biosynthesis, as observed using fluorescence microscopy after DAPI staining, while a reduction in the percentage of cells in the G0/G1 and S phases was detected using flow cytometry [202].

3. Conclusions and Future Perspectives

As stated in Section 1, we included in this review paper published work, which we considered relevant, related to the cytotoxic effects of halophytes. Such reports included results obtained with crude extracts, fractions, and pure compounds, using in vitro and in vivo assays, and they occasionally presented IC50 values, phytochemical profile of the active samples, and identification of active molecules, along with proposed mechanisms of action. There are some drawbacks of the identified reports. For example, several authors tested considerably high concentrations of the target extracts, up to 1000 mg/mL, in some cases. Moreover, the IC50 values were not reported in a high number of studies, which impairs a critical evaluation of the obtained results. Additionally, several reports failed to establish the chemical composition of the cytotoxic extracts and/or the identification of the molecules responsible for the detected activity. Lastly, information about the effect of the extracts on cell lines from nontumoral origin and, therefore, the selectivity of the extracts toward cancer cells was not always provided. According to the American National Cancer Institute NCI), only crude extracts/fractions displaying IC50 values lower than 30 μg/mL in the preliminary assay are considered promising and should be further explored as sources of antitumoral compounds. Therefore, we consider that extracts from the species and extracts included in Table 3 could be explored in detail targeting the isolation of anticancer leads, namely, C. ambrosioides, S. fruticosa, A. millefolium, C. soldanella, C. rotundata, M. indicus, T. populnea, E. camaldulensis, P. major, P. lanceolata, E. crus-galli, and L. barbarum. Moreover, isolated compounds with IC50 values ≤10 μM are considered promising anticancer leads. Therefore, we highlight the compounds in Table 3, namely, 6′-O-(n-butanol) ilekudinoside B ester (A. marina), 5,7-dihydroxy-3′,4′,5′-trimethoxy flavone and tricin (E. crus-galli), Portulacerebroside A (P. oleraceae), and grossamide (L. barbarum).
Table 1. List of halophyte species, organs and extracts tested, cell lines/types of cancer assessed, and obtained half-maximal inhibitory concentrations (IC50) values.
Table 1. List of halophyte species, organs and extracts tested, cell lines/types of cancer assessed, and obtained half-maximal inhibitory concentrations (IC50) values.
FamilySpeciesOrgans ExtractCell Line IC50Proposed Mechanism of ActionRef.
AcanthaceaeAcanthus ebracteatus VahlShootsProtein hydrolysateA431 cells (skin carcinoma) 425.9 ng/mL [34]
Acanthus ilicifolius L.Leaves and rootsEthyl acetateMCF7 cells (breast carcinoma)
PA1 cells (ovarian carcinoma)
24.22 μg/mL (leaf) 29.20 μg/mL (root); 15.74 μg/mL (leaf) 20.00 μg/mL (root) nd[33]
LeavesEthanolHepG2 cells (hepatocellular carcinoma) 100 μg/mLApoptosis induction (DNA damage)[31]
RootsWaterHepG2 cells (hepatocellular carcinoma) 39.76 μg/mLApoptosis induction (DNA damage)[32]
Avicennia alba BlumeLeavesMethanolMCF7 and HeLa cells (breast and cervical carcinomas)57.02 and 44.30 µg/mLReduction in cell size and cell detachment[36]
LeavesChloroform/methanolWiDr cells (human colon carcinoma)173.78 µg/mLApoptosis induction (cell arrest in the G0–G1 phase)[37]
Avicennia marina (Forssk.) VierhLeavesMethanolHeLa cells (cervical carcinoma)107 µg/mLnd[40]
LeavesMethanol extract and fractionsMDA-MB-231 cells (breast carcinoma)
HEK cells (human embryonic kidney)
MDA-MB-231 cells: Crude extract 250 µg/mL
Active faction (luteonin) 28 µg/mL (97 µM)
Luteonin: apoptosis induction (DNA fragmentation, decreased expression of BCL2, decreased expression of TP53).[203]
FruitsIsolated compounds from ethanol/butanol fractions of ethanol extract GSC-3# and GSC-18# cells (human glioma stem cell lines)6′-O-(n-butanol) ilekudinoside B ester: 12.21 µg/mL (14 µM) for GSC-3# and 5.23 µg/mL (6 µM) for GSC-18#nd[204]
Leaves and stemsEthyl acetateMCF-7 cells (estrogen positive breast cancer) naApoptosis induction (ROS production, disruption of Δψm), decrease in caspase-7 protein levels); autophagy[39]
Leaves and seedsWater, ethanol, methanol, ethyl acetate extracts, and fractions from the latter AU565, MDA-MB-231, and BT483 cells (breast cancer)
HepG2 and Huh7 cells (liver)
NIH3T3 cells (nontumoral)
Fraction 2-5: 0.75 μg/mL on AU565 cells
Fraction 3-2-9: 2.1 μg/mL on AU565 cells
Apoptosis induction (DNA fragmentation, cell nucleus condensation and fragmentation, decreased PARP and caspase-8, increased caspase 3)[38]
LeavesEthyl acetate Xenograft MDA-MB-231 tumor growth in nude micendSuppression of tumor growth[38]
AizoaceaeMesembryanthemum crystallinum L.LeavesEthanol extract, ethyl acetate and butanol fractionsHCT116 cells (colon carcinoma)naEthyl acetate and butanol fractions: reduction of ROS, apoptosis induction). Butanol fraction: cell-cycle arrest at G2/M phase[44]
Sesuvium portulacastrum L.Whole plantMethanol, acetone, hexane, and diethyl-etherMDA-MB-231 cells (breast carcinoma), IMR-32cells (neuroblastoma), HCT116 cells (colon carcinoma) Hexane extract: 942.07, 703.40 and 407.87 µg/mL for MDA-MDB-231, IMR32 and HCT116 cells Apoptosis induction: nuclear condensation, cell shrinkage, of apoptotic bodies[45]
AmaranthaceaeArthrocnemum indicum Willd. Moq.Shoots80% methanolCaco-2 cells (colon carcinoma)naDecline of DNA synthesis, cell-cycle arrest at the G2/M phase.[51]
Atriplex halimus L.LeavesEthanolHepG2 cells (hepatocellular carcinoma)
MCF-7 cells (human breast adenocarcinoma)
A549 cells (lung cancer)
HepG2: 54.86 μg/mL MCF-7: 153.6 μg/mL
A549: 101.9 μg/mL
Apoptosis induction (expression of TP53, BCL2, and BAX genes)[54]
Chenopodium album L.Branches and leavesPetroleum etherA549 cells (lung carcinoma)33.31 µg/mLCell-cycle arrest at the G1 phase[63]
Chenopodium ambrosioides L.Whole plantEssential oilMCF7 cells (breast carcinoma)18.75, 9.45 and 10.50 µg/mL after 6, 24 and 48 h of incubation nd[64]
Chenopodium ambrosioides L.LeavesEssential oil, ethanol extract, dichloromethane fractionRAJI cells (lymphoblast)
K562 cells (lymphoblast)
Essential oil: 1.0 g/mL; dichloromethane fraction: 34.0 g/mL (RAJI cells)
Ethanol extract: 47.0 g/mL (K562 cells)
Chenopodium quinoa Willd.SeedsPolysaccharide fractionSMMC 7721 cells (liver cancer)
MCF7 cells (breast carcinoma)
L02 and MCF 10A cells (“normal” cell line)
121.4 μg/mL (24 h), 53.4 μg/mL (48 h)
83.5 μg/mL (24 h), 64.6 μg/mL (48 h)
Salicornia brachiata RoxbShootsMethanolHepG2 cells (hepatocellular carcinoma)267.84 μg/mLModification of cellular morphology[205]
Salicornia europaea L.LeavesEthyl acetate
and methanol
MCF7 cells (breast carcinoma)97.9 and 117.1 μg/mLnd[68]
Suaeda fruticosa (L.) ForsskLeavesHexaneHCT116 cells (colon carcinoma)
HepG2 cells (hepatocellular carcinoma),
and MCF7 cells (breast carcinoma)
17.2 μg/mL
33 μg/mL
28.1 μg/mL
Apoptosis induction, cell-cycle arrest at the G0–G1 phase, chromatin condensation, membrane blebbing[74]
Suaeda fruticosa (L.) Forssk ShootsHexane, dichloromethane, methanol, and waterA549, DLD-1, Caco-2 and HT-29
cells (lung and colon carcinoma)
Hexane extract: 49, 10, 140 and 12 µg/mLnd[80]
Suaeda monoica ForsskWhole plantEthanol, methanol, acetone, and diethyl etherMDA-MB-231 cells (breast carcinoma)Ethanol: 172.38 µg/mL; methanol: 148.77 µg/mL; acetone: 185.56 µg/mL; diethyl ether: 60.18 µg/mLnd[206]
Suaeda palaestina Eig ZoharyShootsDichloromethaneA549 cells (lung carcinoma)
HepG2 cells (hepatocellular carcinoma)
34.82 µg/mL
30.76 µg/mL
Suaeda salsa L.ShootsAcidic polysaccharide (molecular weight = 53.8 kDa; composition: mannose, rhamnose, glucuronic acid, galacturonic acid, galactose and xylose in a molar ratio of 0.6: 8.0: 1.0: 83.6: 5.0: 7.2).MCF7 and MCF-10A cells (human breast carcinoma)naApoptosis induction (reduction of Δψm, increase in the levels of BAX, cytochrome C, caspase-3 and caspase-9, decrease in the level of Bcl-2[208]
ApiaceaeCrithmum maritimum L.Whole plantHexane, ethyl acetate, methanol, ethanolHuh7 and HepG2 cells (hepatocellular carcinoma)naEthyl acetate extract: cell-cycle arrest at the G0/G1 phase after 24 h of incubation and in the G2/M phase after 48 h; increase in necrotic and apoptotic cells [85]
Eryngium maritimum L.Shoots and rootsWaterHepG2 cells (human hepatocellular carcinoma)
Hep2 cells (human laryngeal epidermoid
U138-MG cells (human glioma)
Vero cells (African green monkey kidney)
Shoots: 32.4 µg/mL; roots: 35.0 µg/mL
Shoots: 50 µg/mL; roots: 30.3 µg/mL
Shoots: 32.9 µg/mL; roots: 16.3 µg/mL
AsteraceaeAchillea millefolium L.ShootsMethanol extract combined with bleomycinDU145 cells (prostate carcinoma) and HFFF2 cells (human non-malignant fibroblasts)ndnd[88]
Aerial partsPetroleum ether, ethyl acetate, methanol, waterK562 cells (human myelogenous leukemia),
HeLa cells (human cervical carcinoma)
MCF7 cells (human breast carcinoma)
A549 cells (human non-small cell lung)
Ethyl acetate: 0.58 µg/mL (HeLa), 0.73 µg/mL (K562)
Water: 0.87 µg/mL (MCF-7)
Petroleum ether: 0.87 µg/mL (K562)
Ethyl acetate extract: pre-G1 apoptosis and cell growth arrest in G2/M (HeLa)
Limbarda crithmoides (L.) DumortAerial partsMethanol extract, n-hexane, dichloromethane, and aqueous methanol-soluble fractions, isolated compoundsOCI-AML3 cells (acute myeloid leukaemia)nd10-acetoxy-9Z-chloro-8,9-dehydrothymol: apoptosis induction [92]
BrassicaceaeCakile maritima Scop.Aerial organsMethanol extract, and n-hexane, ethyl acetate, and methanolCaco2 and HeLa cells (colon and cervical carcinoma)Hexane fraction: 12 and 126 µg/mLnd[96]
ConvolvulaceaeCalystegia soldanella L.Whole plantMethylene chloride and methanol extracts, n-hexane, 85% methanol, n-butanol and water fractionsHepG2 cells (hepatocellular carcinoma)naMethanol fraction: cell-cycle arrest at the G0–G1 and S phases, apoptosis induction[100]
Calystegia soldanella L.Whole plantMethanolA549 cells (human lung cancer)
Col2 cells (human colon cancer)
8.0 µg/mL
27.4 µg/mL
Cressa cretica L.ShootsHydroalcoholicHepG2 cells (hepatocellular carcinoma)2300 µg/mLIncreased BAX, decreased BCL2[102]
CymodoceaceaeCymodocea rotundata Ehrin.LeavesSilver particles produced by combining water extract and silver nitrate (AgNO3, 1 M)MG63 cells (osteosarcoma)25.31 µg/mLnd[105]
Cymodocea serrulata (R. Br.) Aschers. & MagnusLeavesSilver particles produced by combining water extract and silver nitrate (AgNO3, 1 M)A549 cells (lung carcinoma)100 µg/mLnd[106]
ShootsHydroethanolicHepG2 cells (hepatocellular carcinoma)82.92 µg/mLnd[107,108]
CyperaceaeCyperus rotundus L.RhizomesMethanol, ethanol, and water.MDA-MB-231 cells (breast carcinoma)225 µg/mLApoptosis induction via upregulation of the death receptor 4 (DR4), DR5, and pro-apoptotic BAX, and downregulation of antiapoptotic BCL2[112]
FabaceaeAlhagi maurorum MedikAerial organsLupeol (isolated from a methanol extract)MCF7, MDA-MB-231 and MCF 10A cells (breast carcinoma)>100 µg/mLIncreased mRNA expression and level of TP53, caspase-3, and BAX genes, decrease in BCL2 gene expression [118]
Glycyrrhiza glabra L.RootMethanolA549 cells (human lung carcinoma)
HepG2 cells (human hepatocellular carcinoma)
HaCaT cells (immortal human keratinocyte)
189.1–238 µg/mL
248.5 µg/mL
158.8–241.9 µg/mL
RootEthanolHT-29 cells (colon carcinoma)naDownregulation of heat-shock protein 90 (HS90) gene expression[123]
Glycyrrhiza uralensis FischRootEthanol/water (7:3, v/v)HeLa cells (cervical carcinoma)nand[125]
Melilotus indicus L.Aerial partsMethanolHepG2 cells (human hepatocellular carcinoma)
SNU-182 cells (hepatocellular carcinoma)
L-02 cells (human “normal” hepatic)
16.60 µg/mL
13.21 µg/mL
90.9 µg/mL
Increase in the number of apoptotic cells, loss of mitochondrial membrane potential (Δψm)[128]
Prosopis cineraria L. DruceLeavesMethanolMCF7 cells (breast carcinoma) and HBL-100 cells (noncancerous breast) nand[209]
Prosopis juliflora Sw. DC.LeavesMethanolMolt-4 cells (human T-cell leukemia)90.5, 42.5 and 20.0 μg/mL (24 h, 48 h and 72 h of incubation)Increased number of micronuclei [130]
Sesbania grandiflora L.LeavesWater, ethanol,
and acetone
IMR32 and HT-29 cell lines (neuroblastoma and colon carcinoma)200 µg/mL [137]
JuncaceaeJuncus acutus L. Torr. Ex Retz.ShootsDiethyl ether,
chloroform, methanol, and water
HepG2 cells (hepatocellular carcinoma)
S17 cells (murine non-tumoral)
Ether extract: 6.2 and
34 µg/mL
Juncunol: 18 µM
(HepG2 cells)
Juncunol: increased number of apoptotic cells, decrease in the Δψm, cell-cycle arrest in the G0/G1 phase, no hemolytic properties[129,130]
MalvaceaeThespesia populnea Sol. Ex CorrêaLeavesDecoctionHEP-2 cells (epidermoid carcinoma)120.02 µg/mLApoptosis induction (membrane blebbing, cell shrinkage, nuclear and cytoplasmic condensation, apoptotic bodies)[144]
Stem barkMethanolMDA-MB-231 and MCF7 cells (breast carcinoma)23.97 and 20.62 µg/mLnd[145]
MyrtaceaeEucalyptus camaldulensis DehnhLeavesEthyl acetateMDA-MB-231 and MCF7 cells (breast carcinoma)26.7 and 34.4 µg/mLnd[148]
PlantaginaceaeBacopa monnieri L. WettstIn vitro cultures (shoots)Methanol and artificial saliva and gastric juice extractsDU145 cells (prostate cancer) ndnd[153]
Plantago lanceolata L.LeavesEthanolMCF7, AMJ13, MDAMB and CAL51 cells (breast carcinoma)0.674, 0.726,
0.251 and 0.024 mg/mL
Plantago major L.SeedsSilver particles produced by combining water extract and silver nitrate (AgNO3, 0.1 M)MCF7 cells (breast carcinoma) 12 µg/mLnd[159]
PlumbaginaceaeLimoniastrum densiflorum (Guss.) KuntzeShootsHexane, dichloromethane, ethanol, and methanolA-549 cells (human lung adenocarcinoma
DLD-1 cells (human colon carcinoma)
WS-1 cells (human skin fibroblasts)
Dichloromethane extract: 85 µg/mL (DLD-1)-and 29 µg/mL (A-549)nd[170]
Limoniastrum guyonianum BoissGallWaterHeLa cells (cervical carcinoma)170 µg/mLDNA hypomethylation and apoptosis, cell-cycle arrest at G2/M, upregulation of p16INK4A, upregulation of UHRF1 and DNMT1[169]
PoaceaeCynodon dactylon L.Whole plantPetroleum ether, dichloromethane, acetone, methanol/water (3/1) and waterMCF7 cells (breast carcinoma)Water extract: 57.21 μg/mL; acetone extract: 38 μg/mL; petroleum ether extract: 39 μg/mLnd[174]
Whole plantPetroleum etherHEP-2, HeLa, and MCF7 cells (laryngeal, cervical, and breast carcinoma)0.20, 0.62 and 1.02 mg/mLApoptosis induction (DNA fragmentation)[211]
Echinochloa crus-galli L.GrainsEthanol extract, n-hexane, chloroform, ethyl acetate, n-butanol fractions, isolated compoundsMCF7, HCT116, HeLa, and HepG2 cells (breast, colon, cervical, and liver carcinoma)Crude extract: 12.0, 11.2, 18.9, and 14.2 μg/mL
Ethyl acetate fraction: 3.8 μg/mL
PortulacaceaePortulaca oleracea L.SeedsEthanolHepG2 cells (hepatocellular carcinoma)75 µg/mLRound cells with reduced size and adhesion[181]
SeedsOil extractionHepG2 and A549 cells (liver and lung carcinoma)ndLoss of cell adhesion capacity, shrinkage, round shape[182]
RhizoporaceaeBruguiera gymnorhiza L. LamStem barkMethanolHeLa cells (cervical cancer), Raji cells (lymphoma) and myeloma cells (leukemia)133, 504, and 384 µg/mLApoptosis induction (DNA fragmentation)[187]
SolanaceaeLycium barbarum L.FruitsWaterSW480 and Caco-2 cells (colon carcinoma)600 and 650 µg/mLCell-cycle arrest at G0/G1[191]
FruitsEthanolA431 cells (cutaneous squamous cell carcinoma)873.7 µg/mLReduced expression of Ki67 and PCNA proteins, increased expression of caspase-3, reduction of BCL2, downregulation of LC3II, reduced the phosphorylation of ERK1/2, and upregulation of JNK. Reversion of the regulation of Beclin1, LC3II, Bcl-2, and cl-caspase-3[192]
FruitsEthanolBALB/c nude mice transplanted tumor model established by subcutaneous injection of A431 cellsndIncreased survival rate, reduced tumor volume and weight, downregulation of Ki67 and MMP-2
TamaricaceaeTamarix aucheriana (Decne.) BaumShootsSyringic acid isolated from a methanol extractSW1116 and SW837 cells (colorectal carcinoma)0.95 and 1.2 mg/mLIncreased percentage of apoptotic cells[199]
ShootsMethyl ferulate isolated from methanol extractSW1116 and SW837 cells (colorectal carcinoma)1.7 and 1.9 mMCell-cycle arrest in the S and G2/M phases, apoptosis induction, reduction in NF-κB DNA-binding activity[200]
Tamarix gallica L.Shoots, leaves and flowers80% methanolCaco-2 cells (colon carcinoma)naDecreased DNA synthesis, cell-cycle arrest at G2/M phase; modification in the levels of cyclin B1, p38, Erk1/2, Chk1, and Chk2[202]
nd: not determined; Δψm: mitochondrial membrane potential, na: data not available.
Table 2. Identified molecules in active extracts from halophyte species.
Table 2. Identified molecules in active extracts from halophyte species.
FamilySpeciesChemical CompoundsChemical Structure ClassRef.
AcanthaceaeAvicennia alba BlumeCatechol boranePharmaceutics 14 02406 i001Organoboron [36]
NeophytadienePharmaceutics 14 02406 i002Terpenoid (diterpene)[36]
Hexadecanoic acidPharmaceutics 14 02406 i003Fatty acid (saturated)[36]
Avicennia marina (Forssk.) VierhLuteoninPharmaceutics 14 02406 i004Phenolic (flavonoid)[203]
AmaranthaceaeArtrochnemum indicumGallic acidPharmaceutics 14 02406 i005Phenolic (phenolic acid)[51]
CyanidinPharmaceutics 14 02406 i006Phenolic (flavonoid)[51]
ChrysoeriolPharmaceutics 14 02406 i007Phenolic (flavonoid)[51]
QuercetinPharmaceutics 14 02406 i008Phenolic (flavonoid)[51]
CatecholPharmaceutics 14 02406 i009Phenolic (benzenediol)
LuteolinPharmaceutics 14 02406 i010Phenolic (flavonoid)[51]
Chenopodium ambrosioides L. AscaridolPharmaceutics 14 02406 i011Monoterpene[65]
AsteraceaeAchillea millefoliumChlorogenic acidPharmaceutics 14 02406 i012Phenolic (phenolic acid)[90]
p-Coumaric acidPharmaceutics 14 02406 i013Phenolic (phenolic acid)[90]
Rosmarinic acidPharmaceutics 14 02406 i014Phenolic (polyphenol)[90]
ApigeninPharmaceutics 14 02406 i015Phenolic (flavonoid)[90]
Glycyrrhiza sp. GlycyrrhizinPharmaceutics 14 02406 i016Saponin[120,121]
Limbarda crithmoides (L.) Dumort10-acetoxy-8,9-epoxythymol tiglatePharmaceutics 14 02406 i017Thymol derivative[92]
10-acetoxy-9Z-chloro-8,9-dehydrothymolPharmaceutics 14 02406 i018Thymol derivative[92]
BrassicaceaeCakile maritima Scop.2-hydroxy-1,8-cineolePharmaceutics 14 02406 i019Monoterpenoid[96]
DecanePharmaceutics 14 02406 i020Alkane[96]
LimonenePharmaceutics 14 02406 i021Monoterpenoid[96]
FabaceaeAlhagi maurorum MedikLupeolPharmaceutics 14 02406 i022Terpenoid (triterpenoid)[118]
Glycyrrhiza uralensis FischIsoquercitrinPharmaceutics 14 02406 i023(Phenolic) flavonoid[125]
4′-Demethylpodophyllotoxin glucosidePharmaceutics 14 02406 i024Lignan[125]
PodophyllotoxinPharmaceutics 14 02406 i025Lignin[125]
JuncaceaeJuncus acutus L.JuncunolPharmaceutics 14 02406 i026Phenanthrene[141]
PlantaginaceaeBacopa monnieri (L.) WettstBacopaside (II)Pharmaceutics 14 02406 i027Terpenoid (isoprenoid)[153]
Plantago lanceolata L.O-Coumaric acidPharmaceutics 14 02406 i028Phenolic (phenolic acid)[210]
RutinPharmaceutics 14 02406 i029Phenolic (flavonoid)[210]
MyricetinPharmaceutics 14 02406 i030Phenolic (flavonoid)[210]
QuercetinPharmaceutics 14 02406 i031Phenolic (flavonoid)[210]
KaempferolPharmaceutics 14 02406 i032Phenolic (flavonoid)[210]
PlumbaginaceaeLimonium densiflorum (Guss.) Kuntzetrans 3-Hydroxycinnamic acidPharmaceutics 14 02406 i033Phenolic (phenolic acid)[170]
MyricetinPharmaceutics 14 02406 i034Flavonoid (phenolic)[170]
IsorhamnetinPharmaceutics 14 02406 i035Flavonoid (phenolic)[170]
PoaceaeCynodon dactylon L. PersDelphinidin-3-O-acetylglucosidePharmaceutics 14 02406 i036Anthocyanin[174]
Echinochloa crus-galli5,7-Dihydroxy-3′,4′,5′-trimethoxy flavone Pharmaceutics 14 02406 i037Phenolic (flavonoid)[175]
5,7,4′-Trihydroxy-3′,5′-dimethoxy flavone (tricin)Pharmaceutics 14 02406 i038Phenolic (flavonoid)[175]
PortulacaceaePortulaca oleracea L.Portulacerebroside APharmaceutics 14 02406 i039Cerebroside[179,180]
Portulacanones BPharmaceutics 14 02406 i040Phenolic (flavonoid)[179,180]
Portulacanones CPharmaceutics 14 02406 i041Phenolic (flavonoid)[179,180]
Portulacanones DPharmaceutics 14 02406 i042Phenolic (flavonoid)[179,180]
2,2′-Dihydroxy-4′,6′-dimethoxychalconePharmaceutics 14 02406 i043Phenolic (flavonoid)[179,180]
SolanaceaeLycium barbarum L.4-O-Methylgrossamide Pharmaceutics 14 02406 i044Phenolic amide[193]
GrossamidePharmaceutics 14 02406 i045Phenolic amide[193]
Lycium barbarum L.ScopoletinPharmaceutics 14 02406 i046Phenolic (coumarin)[190]
2-O-β-D-Glucopyranosyl-L-ascorbic acidPharmaceutics 14 02406 i047Glucoside[190]
TamaricaceaeTamarix aucheriana (Decne.) BaumSyringic acidPharmaceutics 14 02406 i048Phenolic (phenolic acid)[198,200]
Methyl ferulatePharmaceutics 14 02406 i049Phenolic (phenolic acid)[198,200]
Table 3. Cytotoxic activity of selected halophytes species, including types of extracts, compounds, cell lines tested, obtained IC50 values, and proposed mechanisms of action. The criterion of cytotoxic activity for the crude extracts, as established by the American National Cancer Institute (NCI), was considered as an IC50 < 30 μg/mL upon 48 h or 72 h of incubation [24].
Table 3. Cytotoxic activity of selected halophytes species, including types of extracts, compounds, cell lines tested, obtained IC50 values, and proposed mechanisms of action. The criterion of cytotoxic activity for the crude extracts, as established by the American National Cancer Institute (NCI), was considered as an IC50 < 30 μg/mL upon 48 h or 72 h of incubation [24].
FamilySpeciesOrganExtract/FractionCompound Detected in the ExtractCell linesIC50 ValuesMechanismRef.
ambrosioides L.
Whole plantEssential oilndMCF718.75, 9.45 and 10.50 μg/mL at 6, 24 and 48 hDNA fragmentation[64]
Suaeda fruticosa (L.) Forssk. LeavesHexaneMonoterpenes (dihydrojasmone, jasmolone, terpinene-4-ol), diterpenes (pimaric acid, steviol, and momilactone B) and phenolics (quercinol, zingerone, zingerol, neovaflan)HCT116 17.2 µg/mLCell-cycle arrest at the C0–G1 phase and apoptosis induction[74]
Suaeda fruticosa (L.) Forssk.ShootsDichloromethanendDLD-1
10 µg/mL
12 µg/mL
AsteraceaeAchillea millefolium L. Aerial partsEthyl acetate, water, and petroleum ether extractsEthyl acetate: p-coumaric acid, chlorogenic acid, and apigenin
Water extract: rosmarinic acid
K562 cells
HeLa cells
A549 cells
Ethyl acetate: 0.58 µg/mL (HeLa), 0.73 µg/mL (K562)
Water: 0.87 µg/mL (MCF-7)
Petroleum ether: 0.87 µg/mL (K562)
Ethyl acetate extract: preG1 apoptosis and cell growth arrest in G2/M (HeLa) [90]
BrassicaceaeCakile maritima Scop. Aerial partsHexaneGC-MS, decane, limonene, nonaldehyde, dodecaneCaCo2 12 μg/mLnd[96]
ConvolvulaceaeCalystegia soldanella (L.) R.Br. ex Roem. & Schult. Whole plant85% aqueous
methanol fraction
of crude
methanol extract
ndHepG2 <30 μg/mLCell-cycle arrest at the G0–G1 and S phases, apoptosis induction[100]
Whole plantMethanolndA549 cells (human lung cancer)
Col2 cells (human colon cancer)
8.0 µg/mL
27.4 µg/mL
CymodoceaeCymodocea rotundata EhrenbHempr. ex Aschers.LeavesWaterndMG63 25.31 µg/mLnd105
FabaceaeMelilotus indicus L. All. Aerial partsMethanolndHepG2
16.6 µg/mL
13.21 µg/mL
Increase in the number of apoptotic cells, loss of Δψm.[128]
Prosopis juliflora Sw. DC.LeavesMethanolndMolt-420.0 μg/mL after 72 hReduction in micronuclei or cell proliferation [130]
MalvaceaeThespesia populnea L.
Sol. ex Corrêa
BarkChloroform fraction
of methanol extract
Flavonoids, triterpenes, and tanninsMDA-MB-231
23.97 µg/mL 20.62 µg/mLnd[145]
MyrtaceaeEucalyptus camaldulensisLeavesMethanol, ethyl
acetate, n-butanol,
and water
26.7 µg/mL
7.9 µg/mL
4.9 µg/mL
PlantaginaceaePlantago major L. SeedsWaterndMCF712 µg/mLnd[159]
Plantago lanceolata L. LeavesEthanolFlavonoid glycosides: O-cumaric, rutin, myricetin, quercetin and kaempferolCAL5124 µg/mLApoptosis induction increase in the nuclei condensation [210]
PoaceaeEchinochloa crus-galli (L.) P. Beauv. Grains70% ethanolndMCF7
12.0 μg/mL
11.2 μg/mL
18.9 μg/mL
4.2 μg/mL
Grains 5,7,4-Trihydroxy-3,5-dimethoxy flavone (tricin)MCF7
4.3 μg/mL
4.5 μg/mL
4.5 μg/mL
4.2 μg/mL
SolanaceaeLycium barbarum L. StemsEthanolPhenolic amidesGSC-3#
28 µg/mL
20 µg/mL
nd: not determined; Δψm: mitochondrial membrane potential.
Table 4. Cytotoxic activity of selected isolated compounds from halophytes species, including cell lines tested, obtained IC50 values, and proposed mechanisms of action. The criterion of cytotoxic activity for the isolated compounds was an IC50 value ≤10 μM, according to the National Institutes of Health (NIH) for screening the NCI60 program [25].
Table 4. Cytotoxic activity of selected isolated compounds from halophytes species, including cell lines tested, obtained IC50 values, and proposed mechanisms of action. The criterion of cytotoxic activity for the isolated compounds was an IC50 value ≤10 μM, according to the National Institutes of Health (NIH) for screening the NCI60 program [25].
FamilySpeciesOrganCompound Cell LinesIC50 ValuesMechanismRef.
AcanthaceaeAvicennia marina (Forssk.) VierhFruits6′-O-(n-butanol) ilekudinoside B esterGSC-18# (human glioma stem cell lines)6 µMnd[204]
PoaceaeEchinochloa. crus-galli (L.) P. Beauv.Grains5,7-dihydroxy-3′,4′,5′-trimethoxy flavoneHeLa
3.0 µM
3.0 µM
5,7,4-trihydroxy-3,5-dimethoxy flavone (tricin)HCT-116
10.8 µM
8.6 µM
7.2 µM
PortulacaceaePortulaca oleracea L. Aerial
Portulacerebroside AHCCLM3<3.5 µMApoptosis induction via activation of the p38 MAPK and JNK-triggered mitochondrial death pathway[180]
SolanaceaeLycium barbarum L. StemsGrossamideGSC-3#
10.2 µM
9.3 µM
nd: not determined.

Author Contributions

P.G.-C., conceptualization, data curation, and writing—original draft preparation; P.C.-B., writing—review and editing; C.G.P., writing—review and editing; L.C., conceptualization, supervision, and writing—review and editing. All authors have read and agreed to the published version of the manuscript.


This study received Portuguese national funds from FCT—Foundation for Science and Technology through projects UIDB/04326/2020, UIDP/04326/2020 and LA/P/0101/2020, and the FCT Scientific Employment Stimulus (CEECIND/00425/2017). It also received funding through the project HaloFarMs (PRIMA/0002/2019), which is part of the Partnership on Research and Innovation in the Mediterranean Area (PRIMA) Program supported by the European Union and funded by the national funding bodies of Participating States (FCT in Portugal).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Conflicts of Interest

The authors declare no conflict of interest.


  1. Nurmik, M.; Ullmann, P.; Rodriguez, F.; Haan, S.; Letellier, E. In search of definitions: Cancer-associated fibroblasts and their markers. Int. J. Cancer 2020, 146, 895–905. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  2. ACS 2020. American Cancer Society: Cancer Facts and Figures 2020. Atlanta, Ga: American Cancer Society. 2020. Available online: (accessed on 3 December 2020).
  3. Laconi, E.; Marongiu, F.; De Gregori, J. Cancer as a disease of old age: Changing mutational and microenvironmental landscapes. Br. J. Cancer 2020, 122, 943–952. [Google Scholar] [CrossRef] [PubMed]
  4. GBD 2015 Risk Factors Collaborators. Global, Regional, and National Comparative Risk Assessment of 79 Behavioural, Environmental and Occupational, and Metabolic Risks or Clusters of Risks, 1990–2015: A Systematic Analysis for the Global Burden of Disease Study 2015; Lancet: London, UK, 2016; pp. 388–1659.
  5. Anastasiadi, Z.; Lianos, G.D.; Ignatiadou, E.; Harissis, H.V.; Mitsis, M. Breast cancer in young women: An overview. Updat. Surg. 2017, 69, 313–317. [Google Scholar] [CrossRef] [PubMed]
  6. Wadowska, K.; Bil-Lula, I.; Trembecki, Ł.; Śliwińska-Mossoń, M. Genetic markers in lung cancer diagnosis: A review. Int. J. Mol. Sci. 2020, 21, 4569. [Google Scholar] [CrossRef] [PubMed]
  7. Arbyn, M.; Weiderpass, E.; Bruni, L.; de Sanjosé, S.; Saraiya, M.; Ferlay, J.; Bray, F. Estimates of incidence and mortality of cervical cancer in 2018: A worldwide analysis. Lancet Glob. Health 2020, 8, 191–203. [Google Scholar] [CrossRef] [Green Version]
  8. NCI (National Cancer Institute). Available online: (accessed on 3 January 2021).
  9. Mattiuzzi, C.; Lippi, G. Current cancer epidemiology. J. Epidem. Glob. Health 2019, 9, 217–222. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  10. Nigam, M.; Suleria, H.A.R.; Farzaei, M.H.; Mishra, A.P. Marine anticancer drugs and their relevant targets: A treasure from the ocean. DARU J. Pharm. Sci. 2019, 27, 491–515. [Google Scholar] [CrossRef]
  11. Bracci, L.; Schiavoni, G.; Sistigu, A.; Belardelli, F. Immune-based mechanisms of cytotoxic chemotherapy: Implications for the design of novel and rationale-based combined treatments against cancer. Cell Death Differ. 2014, 21, 15–25. [Google Scholar] [CrossRef] [Green Version]
  12. Wu, Q.; Yang, Z.; Nie, Y.; Shi, Y.; Fan, D. Multi-drug resistance in cancer chemotherapeutics: Mechanisms and lab approaches. Cancer Lett. 2014, 347, 159–166. [Google Scholar] [CrossRef]
  13. Bhanot, A.; Sharma, R.; Noolvi, M.N. Natural sources as potential anti-cancer agents: A review. Int. J. Phyto. 2011, 3, 9–26. [Google Scholar]
  14. Atanasov, A.G.; Waltenberger, B.; Pferschy-Wenzig, E.M.; Linder, T.; Wawrosch, C.; Uhrin, P.; Temml, V.; Wang, L.; Schwaiger, S.; Heiss, E.H.; et al. Discovery and resupply of pharmacologically active plant-derived natural products: A review. Biotech. Adv. 2015, 33, 1582–1614. [Google Scholar] [CrossRef] [PubMed]
  15. Willdenov, C.L. The Principles of Botany, and of Vegetable Physiology; University Press: Edinburgh, UK, 1805. [Google Scholar]
  16. Grigore, M.N. Defining halophytes: A conceptual and historical approach. In Halophytes and Climate Change: Adaptative Mechanisms and Potential Uses; Hasanuzzaman, M., Shabala, S., Fujita, M., Eds.; CABI: Wallingford, UK, 2019; pp. 3–18. [Google Scholar]
  17. Flowers, T.J.; Colmer, T.D. Salinity tolerance in halophytes. New Phytol. 2008, 179, 945–963. [Google Scholar] [CrossRef] [PubMed]
  18. Ksouri, R.; Ksouri, W.M.; Jallali, I.; Debez, A.; Magné, C.; Hiroko, I.; Abdelly, C. Medicinal halophytes: Potent source of health promoting biomolecules with medical, nutraceutical and food applications. Crit. Rev. Biotech. 2012, 32, 289–326. [Google Scholar] [CrossRef] [PubMed]
  19. Hasanuzzaman, M.; Nahar, K.; Alam, M.; Bhowmik, P.C.; Hossain, M.; Rahman, M.M.; Prasad, M.N.V.; Ozturk, M.; Fujita, M. Potential use of halophytes to remediate saline soils. BioMed Res. Int. 2014, 2014, 589341. [Google Scholar] [CrossRef] [PubMed]
  20. Ksouri, R.; Megdiche, W.; Falleh, H.; Trabelsi, N.; Boulaaba, M.; Smaoui, A.; Abdelly, C. Influence of biological, environmental and technical factors on phenolic content and antioxidant activities of Tunisian halophytes. Comptes Rendus Biol. 2008, 331, 865–873. [Google Scholar] [CrossRef]
  21. Shahidi, F.; Ambigaipalan, P. Phenolics and polyphenolics in foods, beverages and spices: Antioxidant activity and health effects–A review. J. Funct. Foods 2015, 18, 820–897. [Google Scholar] [CrossRef]
  22. Arya, S.S.; Devi, S.; Ram, K.; Kumar, S.; Kumar, N.; Mann, A.; Kumar, A.; Chand, G. Halophytes: The plants of therapeutic medicine. In Ecophysiology, Abiotic Stress Responses and Utilization of Halophytes; Springer: Singapore, 2019; pp. 271–287. [Google Scholar]
  23. eHALOPH–Halophytes Database© Version 3.20. The University of Sussex. 2020. Available online: (accessed on 4 November 2020).
  24. Suffness, M.; Pezzuto, J.M. Assays related to cancer drug discovery. In Methods in Plant Biochemistry: Assays for Bioactivity; Hostettmann, K., Ed.; Academic Press: London, UK, 1990; pp. 71–133. [Google Scholar]
  25. Cruz, S.; Gomes, S.E.; Borralho, P.M.; Rodrigues, C.M.P.; Gaudêncio, S.P.; Pereira, F. In Silico HCT116 Human Colon Cancer Cell-Based Models En Route to the Discovery of Lead-Like Anticancer Drugs. Biomolecules 2018, 8, 56. [Google Scholar] [CrossRef] [Green Version]
  26. Renna, M.; Gonnella, M. Ethnobotany, nutritional traits, and healthy properties of some halophytes used as greens in the Mediterranean basin. In Handbook of Halophytes: From Molecules to Ecosystems towards Biosaline Agriculture; Grigore, M.N., Ed.; Springer: Cham, Switzerland, 2020; pp. 1–19. [Google Scholar]
  27. Catalogue of Life. Available online: (accessed on 18 October 2022).
  28. Xu, Z.; Chang, L. Acanthaceae. In Identification and Control of Common Weeds; Springer: Singapore, 2017; Volume 3, pp. 329–338. [Google Scholar]
  29. Khan, I.; Jan, S.A.; Shinwari, Z.K.; Ali, M.; Khan, Y.; Kumar, T. Ethnobotany and medicinal uses of folklore medicinal plants belonging to family Acanthaceae: An updated review. MOJ Biol. Med. 2017, 1, 34–38. [Google Scholar] [CrossRef] [Green Version]
  30. Velmani, S.; Perumal, B.; Santhosh, C.; Vetrivel, C.; Maruthupandian, A. Phytochemical and traditional uses on Acanthus ilicifolius (L). J. Adv. Appl. Scient. Res. 2016, 1, 43–48. [Google Scholar] [CrossRef]
  31. Rajamanickam, E.; Gurudeeban, S.; Satyavani, K.; Ramathan, T. In vitro anticancer effect of Acanthus ilicifolius on hepatocellular carcinoma cells. Asian J. Biochem. 2014, 9, 179–186. [Google Scholar] [CrossRef] [Green Version]
  32. Paul, T.; Ramasubbu, S. The antioxidant, anticancer and anticoagulant activities of Acanthus ilicifolius L. roots and Lumnitzera racemosa Willd. leaves, from southeast coast of India. J. Appl. Pharm. Sci. 2017, 7, 81–87. [Google Scholar] [CrossRef]
  33. Smitha, R.B.; Madhusoodanan, P.V.; Prakashkumar, R. Anticancer activity of Acanthus illicifolius linn. from Chettuva mangroves, Kerala, India. Int. J. Bioassays 2014, 3, 3452–3455. [Google Scholar]
  34. Khamwut, A.; Jevapatarakul, D.; Reamtong, O. In vitro evaluation of anti-epidermoid cancer activity of Acanthus ebracteatus protein hydrolysate and their effects on apoptosis and cellular proteins. Oncol. Lett. 2019, 18, 3128–3136. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  35. Thatoi, H.; Samantaray, D.; Das, S.K. The genus Avicennia, a pioneer group of dominant mangrove plant species with potential medicinal values: A review. Front. Life Sci. 2016, 9, 267–291. [Google Scholar] [CrossRef] [Green Version]
  36. Eswaraiah, G.; Peele, K.A.; Krupanidhi, S.; Kumar, R.B.; Venkateswarulu, T.C. Identification of bioactive compounds in leaf extract of Avicennia alba by GC-MS analysis and evaluation of its in-vitro anticancer potential against MCF7 and HeLa cell lines. J. King Saud. Univ. Sci. 2020, 32, 740–744. [Google Scholar] [CrossRef]
  37. Illian, D.N.; Hasibuan, P.A.Z.; Sumardi, S.; Nuryawan, A.; Wati, R.; Basyuni, M. Anticancer activity of polyisoprenoids from Avicennia alba Blume. in WiDr cells. Iran J. Pharm. Res. 2019, 18, 1477–1487. [Google Scholar] [CrossRef]
  38. Huang, C.; Lu, C.K.; Tu, M.C.; Chang, J.H.; Chen, Y.J.; Tu, Y.H.; Huang, H.C. Polyphenol-rich Avicennia marina leaf extracts induce apoptosis in human breast and liver cancer cells and in a nude mouse xenograft model. Oncotarget 2016, 7, 35874–35893. [Google Scholar] [CrossRef] [Green Version]
  39. Esau, L.; Sagar, S.; Bajic, V.B.; Kaur, M. Autophagy inhibition enhances the mitochondrial-mediated apoptosis induced by mangrove (Avicennia marina) extract in human breast cancer cells. Eur. J. Med. Plants 2015, 5, 304–317. [Google Scholar] [CrossRef] [Green Version]
  40. Devi, A.S.; Joseph, J.; Bhuvaneshwari, V. Antioxidant, anticancerous and cytotoxic effects of Avicennia marina. Int. J. Res. Pharm. Sci. 2018, 9, 996–1001. [Google Scholar] [CrossRef]
  41. Catalogue of Life. Available online: (accessed on 18 October 2022).
  42. Klak, C. Aizoaceae. Bothalia 2011, 41, 292–294. [Google Scholar] [CrossRef]
  43. El-Amier, Y.A.; Haroun, S.A.; El-Shehaby, O.A.; Al-hadithy, O.N. Antioxidant and antimicrobial properties of some wild Aizoaceae species growing in Egyptian desert. J. Environ. Sci. 2016, 45, 1–10. [Google Scholar]
  44. Seo, J.A.; Ju, J. Antioxidant and growth inhibitory activities of Mesembryanthemum crystallinum L. in HCT116 human colon cancer cells. J. Nutr. Health 2019, 52, 157–167. [Google Scholar] [CrossRef] [Green Version]
  45. Chintalapani, S.; Swathi, M.S.; Narasu, M.L. Antiproliferative and apoptosis inducing potential of whole plant extracts of Sesuvium portulacastrum against different cancer cell lines. J. Appl. Pharm. Sci. 2019, 9, 38–43. [Google Scholar] [CrossRef]
  46. Catalogue of Life. Available online: (accessed on 18 October 2022).
  47. Xu, Z.; Deng, M. Amaranthaceae. In Identification and Control of Common Weeds; Springer: Dordrecht, The Netherlands, 2017; pp. 245–302. [Google Scholar]
  48. Mroczek, A. Phytochemistry and bioactivity of triterpene saponins from Amaranthaceae family. Phytochem. Rev. 2015, 14, 577–605. [Google Scholar] [CrossRef]
  49. Nisar, F.; Gul, B.; Khan, M.A.; Hameed, A. Germination and recovery responses of heteromorphic seeds of two co-occurring Arthrocnemum species to salinity, temperature and light. S. Afr. J. Bot. 2019, 121, 143–151. [Google Scholar] [CrossRef]
  50. ElNaker, N.A.; Yousef, A.F.; Yousef, L.F. A review of Arthrocnemum (Arthrocaulon) macrostachyum chemical content and bioactivity. Phytochem. Rev. 2020, 19, 1427–1448. [Google Scholar] [CrossRef]
  51. Boulaaba, M.; Mkadmini, K.; Tsolmon, S.; Han, J.; Smaoui, A.; Kawada, K.; Ksouri, R.; Isoda, H.; Abdelly, C. In vitro antiproliferative effect of Arthrocnemum indicum extracts on Caco-2 cancer cells through cell cycle control and related phenol LC-TOF-MS identification. Evid.-Based Compl. Altern. Med. 2013, 2013, 529375. [Google Scholar] [CrossRef] [Green Version]
  52. Yang, Y.; Zhou, X.; Xiao, M.; Hong, Z.; Gong, Q.; Jiang, L.; Zhou, J. Discovery of chrysoeriol, a PI3K-AKT-mTOR pathway inhibitor with potent antitumor activity against human multiple myeloma cells in vitro. J. Huazhong Univ. Sci. Techol. Med. Sci. 2010, 30, 734–740. [Google Scholar] [CrossRef]
  53. Benzarti, M.; Ben Rejeb, K.; Debez, A.; Abdelly, C. Environmental and economical opportunities for the valorisation of the genus Atriplex: New insights. In Crop Improvement; Springer: Boston, MA, USA, 2013; pp. 441–457. [Google Scholar]
  54. Al-Senosy, N.K.; Abou-Eisha, A.; Ahmad, E.S. In vitro antiproliferation effect of Atriplex halimus L. crude extract on human cell lines by induction of apoptosis and G2/M phase arrest. Egypt Acad. J. Biol. Sci. C Phys. Mol. Biol. 2018, 10, 115–126. [Google Scholar] [CrossRef]
  55. Petropoulos, S.A.; Karkanis, A.; Martins, N.; Ferreira, I.C. Halophytic herbs of the Mediterranean basin: An alternative approach to health. Food Chem. Toxicol. 2018, 114, 155–169. [Google Scholar] [CrossRef] [Green Version]
  56. Al-Hazzani, A.A.; Alshatwi, A.A. Catechin hydrate inhibits proliferation and mediates apoptosis of SiHa human cervical cancer cells. Food Chem. Toxicol. 2011, 49, 3281–3286. [Google Scholar] [CrossRef] [PubMed]
  57. Di Domenico, F.; Foppoli, C.; Coccia, R.; Perluigi, M. Antioxidants in cervical cancer: Chemopreventive and chemotherapeutic effects of polyphenols. Biochim. Biophys. Acta (BBA)—Mol. Basis Dis. 2012, 1822, 737–747. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  58. Roleira, F.M.; Tavares-da-Silva, E.J.; Varela, C.L.; Costa, S.C.; Silva, T.; Garrido, J.; Borges, F. Plant derived and dietary phenolic antioxidants: Anticancer properties. Food Chem. 2015, 183, 235–258. [Google Scholar] [CrossRef] [PubMed]
  59. Boeing, J.S.; Barizão, É.O.; Rotta, E.M.; Volpato, H.; Nakamura, C.V.; Maldaner, L.; Visentainer, J.V. Phenolic compounds from Butia odorata (Barb. Rodr.) Noblick fruit and its antioxidant and antitumor activities. Food Anal Methods 2020, 13, 61–68. [Google Scholar] [CrossRef]
  60. Rha, C.S.; Jeong, H.W.; Park, S.; Lee, S.; Jung, Y.S.; Kim, D.O. Antioxidative, anti-inflammatory, and anticancer effects of purified flavonol glycosides and aglycones in green tea. Antioxidants 2019, 8, 278. [Google Scholar] [CrossRef] [Green Version]
  61. Kokanova-Nedialkova, Z.; Nedialkov, P.; Nikolov, S. The genus Chenopodium: Phytochemistry, ethnopharmacology and pharmacology. Pharmacogn. Rev. 2009, 3, 280. [Google Scholar]
  62. Choudhary, N.; Prabhu, K.S.; Prasad, S.B.; Singh, A.; Agarhari, U.C.; Suttee, A. Phytochemistry and pharmacological exploration of Chenopodium album: Current and future perspectives. Res. J. Pharm. Techol. 2020, 13, 3933–3940. [Google Scholar] [CrossRef]
  63. Zhao, T.; Pan, H.; Feng, Y.; Li, H.; Zhao, Y. Petroleum ether extract of Chenopodium album L. prevents cell growth and induces apoptosis of human lung cancer cells. Exp. Ther. Med. 2016, 12, 3301–3307. [Google Scholar] [CrossRef] [Green Version]
  64. Jia-liang, W.; Dan-wei, M.; Ya-nan, W.; Hong, Z.; Bing, H.; Qun, L.; Zhi-yan, Z.; Jing, F. Cytotoxicity of essential oil of Chenopodium ambrosioides L. against human breast cancer MCF-7 cells. Trop. J. Pharm. Res. 2013, 12, 929–933. [Google Scholar] [CrossRef] [Green Version]
  65. Degenhardt, R.T.; Farias, I.V.; Grassi, L.T.; Franchi Jr, G.C.; Nowill, A.E.; Bittencourt, C.M.D.S.; Wagner, T.M.; Bella Cruz, A.; Malheiros, A. Characterization and evaluation of the cytotoxic potential of the essential oil of Chenopodium ambrosioides. Rev. Bras. Farmacogn. 2016, 26, 56–61. [Google Scholar] [CrossRef] [Green Version]
  66. Hu, Y.; Zhang, J.; Zou, L.; Fu, C.; Li, P.; Zhao, G. Chemical characterization, antioxidant, immune-regulating and anticancer activities of a novel bioactive polysaccharide from Chenopodium quinoa seeds. Inter. J. Biol. Macromol. 2017, 99, 622–629. [Google Scholar] [CrossRef] [PubMed]
  67. Patel, S. Salicornia: Evaluating the halophytic extremophile as a food and a pharmaceutical candidate. 3Biotech 2016, 6, 104. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  68. Samuel, P.; Kumar, V.J.; Dhayalan, D.R.; Amirtharaj, K.; Sudarmani, D.N.P. Bioprospecting of Salicornia europaea L. a marine halophyte and evaluation of its biological potential with special reference to anticancer activity. J. Pharma. Pharma. Sci. 2017, 2, 138. [Google Scholar] [CrossRef]
  69. Kensil, C.R. Saponins as vaccine adjuvants. Crit. Rev. Ther. Drug. Carr. Syst. 1996, 13, 1–55. [Google Scholar]
  70. Yurdakok, B.; Baydan, E. Cytotoxic effects of Eryngium kotschyi and Eryngium maritimum on Hep2, HepG2, Vero and U138 MG cell lines. Pharma. Biol. 2013, 51, 1579–1585. [Google Scholar] [CrossRef]
  71. Yildirim, I.; Kutlu, T. Anticancer agents: Saponin and tannin. Int. J. Biol. Chem. 2015, 9, 332–340. [Google Scholar] [CrossRef] [Green Version]
  72. Munir, U.; Perveen, A.; Qamarunnisa, S. Comparative pharmacognostic evaluation of some species of the genera Suaeda and Salsola leaf (Chenopodiaceae). Pak. J. Pharm. Sci. 2014, 27, 1309–1315. [Google Scholar]
  73. Altay, V.; Ozturk, M. The Genera Salsola and Suaeda (Amaranthaceae) and their value as fodder. In Handbook of Halophytes; From molecules to ecosystems towards biosaline agriculture; Springer: Cham, Switzerland, 2020; pp. 1–12. [Google Scholar]
  74. Saleh, K.A.; Albinhassan, T.H.; Al-Ghazzawi, A.M.; Mohaya, A.; Shati, A.A.; Ayoub, H.J.; Abdallah, Q.M. Anticancer property of hexane extract of Suaeda fruticosa plant leaves against different cancer cell lines. Trop. J. Pharm. Res. 2020, 19, 129–136. [Google Scholar] [CrossRef]
  75. Su, P.; Veeraraghavan, V.P.; Krishna Mohan, S.; Lu, W. A ginger derivative, zingerone-a phenolic compound-induces ROS-mediated apoptosis in colon cancer cells (HCT-116). J. Biochem. Mol. Toxicol. 2019, 33, e22403. [Google Scholar] [CrossRef]
  76. Sobral, M.V.; Xavier, A.L.; Lima, T.C.; de Sousa, D.P. Antitumor activity of monoterpenes found in essential oils. Sci. World. J. 2014, 2014, 953451. [Google Scholar] [CrossRef] [Green Version]
  77. Garcia, C.; Teodósio, C.; Oliveira, C.; Oliveira, C.; Díaz-Lanza, A.; Reis, C.; Duarte, N.; Rijo, P. Naturally occurring Plectranthus-derived diterpenes with antitumoral activities. Curr. Pharm. Des. 2018, 24, 4207–4236. [Google Scholar] [CrossRef] [PubMed]
  78. Ahmad, S.; Ullah, F.; Zeb, A.; Ayaz, M.; Ullah, F.; Sadiq, A. Evaluation of Rumex hastatus D. Don for cytotoxic potential against HeLa and NIH/3T3 cell lines: Chemical characterization of chloroform fraction and identification of bioactive compounds. BMC Compl. Altern. Med. 2016, 16, 308. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  79. Gupta, E.; Kaushik, S.; Purwar, S.; Sharma, R.; Balapure, A.K.; Sundaram, S. Anticancer potential of steviol in MCF-7 human breast cancer cells. Pharmacogn. Mag. 2017, 13, 345–350. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  80. Oueslati, S.; Ksouri, R.; Falleh, H.; Pichette, A.; Abdelly, C.; Legault, J. Phenolic content, antioxidant, anti-inflammatory and anticancer activities of the edible halophyte Suaeda fruticosa Forssk. Food Chem. 2012, 132, 943–947. [Google Scholar] [CrossRef]
  81. Catalogue of Life. Available online: (accessed on 18 October 2022).
  82. Chen, Y.; Peng, S.; Luo, Q.; Zhang, J.; Guo, Q.; Zhang, Y.; Chai, X. Chemical and pharmacological progress on polyacetylenes isolated from the family Apiaceae. Chem. Biodiv. 2015, 12, 474–502. [Google Scholar] [CrossRef]
  83. Wang, P.; Su, Z.; Yuan, W.; Deng, G.; Li, S. Phytochemical constituents and pharmacological activities of Eryngium L. (Apiaceae). Pharm. Crop. 2012, 3, 99–120. [Google Scholar] [CrossRef] [Green Version]
  84. Renna, M.; Gonnella, M.; Caretto, S.; Mita, G.; Serio, F. Sea fennel (Crithmum maritimum L.): From underutilized crop to new dried product for food use. Gen. Res. Crop. Evol. 2017, 64, 205–216. [Google Scholar] [CrossRef]
  85. Gnocchi, D.; Cesari, G.; Calabrese, G.J.; Capone, R.; Sabbà, C.; Mazzocca, A. Inhibition of hepatocellular carcinoma growth by ethyl acetate extracts of apulian Brassica oleracea L. and Crithmum maritimum L. Plant Foods Hum. Nutr. 2020, 75, 33–40. [Google Scholar] [CrossRef]
  86. Catalogue of Life. Available online: (accessed on 18 October 2022).
  87. Gao, T.; Yao, H.; Song, J.; Zhu, Y.; Liu, C.; Chen, S. Evaluating the feasibility of using candidate DNA barcodes in discriminating species of the large Asteraceae family. BMC Evol. Biol. 2010, 10, 324. [Google Scholar] [CrossRef] [Green Version]
  88. Shahani, S.; Hamzekanlu, N.; Zakeri, N.; Hosseinimehr, S.J. Synergistic effect of Achillea millefolium L. combined with bleomycin on prostate cancer cell. Res. Mol. Med. 2015, 3, 12–17. [Google Scholar] [CrossRef]
  89. Li, Y.; Zhang, M.L.; Cong, B.; Wang, S.M.; Dong, M.; Sauriol, F.; Huo, C.-H.; Shi, Q.-W.; Gu, Y.-G.; Kiyota, H. Achillinin A, a cytotoxic guaianolide from the flower of Yarrow, Achillea millefolium. Biosci. Biotechnol. Biochem. 2011, 75, 1554–1556. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  90. Abou Baker, D.H. Achillea millefolium L. ethyl acetate fraction induces apoptosis and cell cycle arrest in human cervical cancer (HeLa) cells. Ann. Agric. Sci. 2020, 65, 42–48. [Google Scholar] [CrossRef]
  91. Anantharaju, P.G.; Gowda, P.C.; Vimalambike, M.G.; Madhunapantula, S.V. An overview on the role of dietary phenolics for the treatment of cancers. Nutr. J. 2016, 15, 99. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  92. Adorisio, S.; Giamperi, L.; Bucchini, A.E.A.; Delfino, D.V.; Marcotullio, M.C. Bioassay-guided isolation of antiproliferative compounds from Limbarda crithmoides (L.) Dumort. Molecules 2020, 25, 1893. [Google Scholar] [CrossRef] [Green Version]
  93. Catalogue of Life. Available online: (accessed on 18 October 2022).
  94. Anjum, N.A.; Gill, S.S.; Ahmad, I.; Pacheco, M.; Duarte, A.C.; Umar, S.; Khan, N.A.; Pereira, M.E. The plant family Brassicaceae: An introduction. In Plant Family Brassicaceae; Springer: Dordrecht, The Netherlands, 2012; pp. 1–33. [Google Scholar]
  95. Arbelet-Bonnin, D.; Ben-Hamed-Louati, I.; Laurenti, P.; Abdelly, C.; Ben-Hamed, K.; Bouteau, F. Cakile maritima, a promising model for halophyte studies and a putative cash crop for saline agriculture. In Advances in Agronomy; Academic Press: Cambridge, MA, USA, 2019; Volume 155, pp. 45–78. [Google Scholar] [CrossRef]
  96. Omer, E.; Elshamy, A.; Taher, R.; El-Kashak, W.; Shalom, J.; White, A.; Cock, I. Cakile maritima Scop. extracts inhibit CaCo2 and HeLa human carcinoma cell growth: GC-MS analysis of an anti-proliferative extract. Pharm. J. 2019, 11, 258–266. [Google Scholar] [CrossRef]
  97. Hafidh, R.R.; Hussein, S.Z.; MalAllah, M.Q.; Abdulamir, A.S.; Abu Bakar, F. A high-throughput quantitative expression analysis of cancer-related genes in human HepG2 cells in response to limonene, a potential anticancer agent. Curr. Cancer Drug Targets 2018, 18, 807–815. [Google Scholar] [CrossRef]
  98. Catalogue of Life. Available online: (accessed on 18 October 2022).
  99. Sultana, R.; Rahman, A.H.M.M. Convolvulaceae: A taxonomically and medicinally important morning glory family. Int. J. Bot. Stud. 2016, 1, 47–52. [Google Scholar]
  100. Lee, J.I.; Kim, I.H.; Nam, T.J. Crude extract and solvent fractions of Calystegia soldanella induce G1 and S phase arrest of the cell cycle in HepG2 cells. Int. J. Oncol. 2017, 50, 414–420. [Google Scholar] [CrossRef] [Green Version]
  101. Min, H.-Y.; Park, H.-J.; Kim, Y.; Lee, E.-J.; Hwang, H.-J.; Park, E.-J.; Lee, S.K. Cytotoxic activities of indigenous plant extracts in cultured cancer cells. Nat. Prod. Sci. 2002, 8, 170–172. [Google Scholar]
  102. Pouraminaei, M.; Mirzaiey, M.R.; Khoshdel, A.; Hajizadeh, M.R.; Mahmoodi, M.; Fahmidehkar, M.A. The effect of Cressa cretica hydroalcoholic extract on apoptosis and the expression of Bcl2, Bax and P53 genes in hepatoma cell line HepG2. Gene Rep. 2020, 20, 100692. [Google Scholar] [CrossRef]
  103. Catalogue of Life. Available online: (accessed on 18 October 2022).
  104. Petersen, G.; Seberg, O.; Short, F.T.; Fortes, M.D. Complete genomic congruence but non-monophyly of Cymodocea (Cymodoceaceae), a small group of seagrasses. Taxon 2014, 63, 3–8. [Google Scholar] [CrossRef]
  105. Abraham, J.; Saraf, S.; Mustafa, V.; Chaudhary, Y.; Sivanangam, S. Synthesis and evaluation of silver nanoparticles using Cymodocea rotundata against clinical pathogens and human osteosarcoma cell line. J. Appl. Pharm. Sci. 2017, 7, 55–61. [Google Scholar] [CrossRef] [Green Version]
  106. Palaniappan, P.; Sathishkumar, G.; Sankar, R. Fabrication of nano-silver particles using Cymodocea serrulata and its cytotoxicity effect against human lung cancer A549 cells line. Spectroch. Acta Part A Mol. Biomol. Spec. 2015, 138, 885–890. [Google Scholar] [CrossRef] [PubMed]
  107. Kalpana, C.S.; Kalaivani, P.; Vanitha, V. Phytochemical screening and effect of Cymodocea serrulata on HepG2-human hepatocellular carcinoma cell line. Inter. J. Res. Pharm. Sci. 2020, 11, 1684–1691. [Google Scholar] [CrossRef]
  108. Sofi, M.S.; Nabi, S.; Mohammed, C.; Sofi, S. The role of phytocompounds in cancer treatment: A current review. J. Med. Plant Stud. 2018, 6, 83–93. [Google Scholar]
  109. Catalogue of Life. Available online: (accessed on 18 October 2022).
  110. Semmouri, I.; Bauters, K.; Léveillé-Bourret, É.; Starr, J.R.; Goetghebeur, P.; Larridon, I. Phylogeny and systematics of Cyperaceae, the evolution and importance of embryo morphology. Bot. Rev. 2019, 85, 1–39. [Google Scholar] [CrossRef]
  111. Al-Snafi, A.E. A review on Cyperus rotundus: A potential medicinal plant. IOSR J. Pharm. 2016, 6, 32–48. [Google Scholar] [CrossRef]
  112. Park, S.E.; Shin, W.T.; Park, C.; Hong, S.H.; Kim, G.Y.; Kim, S.O.; Ryu, C.H.; Hong, S.H.; Choi, Y.H. Induction of apoptosis in MDA-MB-231 human breast carcinoma cells with an ethanol extract of Cyperus rotundus L. by activating caspases. Oncol. Rep. 2014, 32, 2461–2470. [Google Scholar] [CrossRef]
  113. Singh, N.; Pandey, B.R.; Verma, P.; Bhalla, M.; Gilca, M. Phyto-pharmacotherapeutics of Cyperus rotundus Linn. (Motha): An overview. Ind. J. Nat. Prod. Res. 2012, 3, 467–476. [Google Scholar]
  114. Catalogue of Life. Available online: (accessed on 18 October 2022).
  115. Ahmad, F.; Anwar, F.; Hira, S. Review on medicinal importance of Fabaceae family. Pharm. Online 2016, 3, 151–157. [Google Scholar]
  116. Muhammad, G.; Hussain, M.A.; Anwar, F.; Ashraf, M.; Gilani, A.H. Alhagi: A plant genus rich in bioactives for pharmaceuticals. Phyto. Res. 2015, 29, 1–13. [Google Scholar] [CrossRef] [PubMed]
  117. Laghari, A.H.; Memon, S.; Nelofar, A.; Khan, K.M.; Yasmin, A.; Syed, M.N.; Aman, A. A new flavanenol with urease-inhibition activity isolated from roots of manna plant camelthorn (Alhagi maurorum). J. Mol. Struc. 2010, 965, 65–67. [Google Scholar] [CrossRef]
  118. Behbahani, M. Evaluation of in vitro anticancer activity of Ocimum basilicum, Alhagi maurorum, Calendula officinalis and their parasite Cuscuta campestris. PLoS ONE 2014, 9, e116049. [Google Scholar] [CrossRef] [PubMed]
  119. Farag, M.A.; Porzel, A.; Wessjohann, L.A. Unequivocal glycyrrhizin isomer determination and comparative in vitro bioactivities of root extracts in four Glycyrrhiza species. J. Adv. Res. 2015, 6, 99–104. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  120. Al-Snafi, A.E. Glycyrrhiza glabra: A phytochemical and pharmacological review. IOSR J. Pharm. 2018, 8, 1–17. [Google Scholar]
  121. Farooqui, A.; Khan, F.; Khan, I.; Ansari, I.A. Glycyrrhizin induces reactive oxygen species-dependent apoptosis and cell cycle arrest at G0/G1 in HPV18+ human cervical cancer HeLa cell line. Biomed Pharm. 2018, 97, 752–764. [Google Scholar] [CrossRef]
  122. Basar, N.; Oridupa, O.A.; Ritchie, K.J.; Nahar, L.; Osman, N.M.M.; Stafford, A.; Kushiev, H.; Kan, A.; Sarker, S.D. Comparative cytotoxicity of Glycyrrhiza glabra roots from different geographical origins against immortal human keratinocyte (HaCaT), lung adenocarcinoma (A549) and liver carcinoma (HepG2) cells. Phyto. Res. 2015, 29, 944–948. [Google Scholar] [CrossRef] [Green Version]
  123. Nourazarian, S.M.; Nourazarian, A.; Majidinia, M.; Roshaniasl, E. Effect of root extracts of medicinal herb Glycyrrhiza glabra on HSP90 gene expression and apoptosis in the HT-29 colon cancer cell line. Asian Pac. J. Cancer Prev. 2016, 16, 8563–8566. [Google Scholar] [CrossRef] [Green Version]
  124. Gao, C.; Peng, Y.N.; Wang, H.Z.; Fang, S.L.; Zhang, M.; Zhao, Q.; Liu, J. Inhibition of heat shock protein 90 as a novel platform for the treatment of cancer. Curr. Pharm. Des. 2019, 25, 849–855. [Google Scholar] [CrossRef]
  125. Fan, H.; Xuan, J.; Zhang, K.; Jiang, J. Anticancer component identification from the extract of Dysosma versipellis and Glycyrrhiza uralensis based on support vector regression and mean impact value. Anal. Methods 2018, 10, 371–380. [Google Scholar] [CrossRef]
  126. Wu, F.; Ma, J.; Meng, Y.; Zhang, D.; Pascal Muvunyi, B.; Luo, K.; Di, H.; Guo, W.; Wang, Y.; Feng, B.; et al. Potential DNA barcodes for Melilotus species based on five single loci and their combinations. PLoS ONE 2017, 12, e0182693. [Google Scholar] [CrossRef] [PubMed]
  127. Ahmed, S.A.K.; Al-Refai, M. Chemical constituents and cytotoxic activities of the extracts of Melilotus indicus. Eur. J. Chem. 2014, 5, 503–506. [Google Scholar] [CrossRef] [Green Version]
  128. El-Hafeez, A.A.A.; Khalifa, H.O.; Elgawish, R.A.; Shouman, S.A.; El-Twab, M.H.A.; Kawamoto, S. Melilotus indicus extract induces apoptosis in hepatocellular carcinoma cells via a mechanism involving mitochondria-mediated pathways. Cytotechnology 2018, 70, 831–842. [Google Scholar] [CrossRef] [PubMed]
  129. Henciya, S.; Seturaman, P.; James, A.R.; Tsai, Y.H.; Nikam, R.; Wu, Y.C.; Dahms, H.-U.; Chang, F.R. Biopharmaceutical potentials of Prosopis spp. (Mimosaceae, Leguminosa). J. Food Drug Anal. 2017, 25, 187–196. [Google Scholar] [CrossRef]
  130. Sathiya, M.; Muthuchelian, K. Anti-tumor potential of total alkaloid extract of Prosopis juliflora DC. leaves against Molt-4 cells in vitro. Afr. J. Biotech. 2011, 10, 8881–8888. [Google Scholar] [CrossRef] [Green Version]
  131. Bunma, S.; Balslev, H. A review of the economic botany of Sesbania (Leguminosae). Bot. Rev. 2019, 85, 185–251. [Google Scholar] [CrossRef]
  132. Boddawar, G.D.; Dhawale, S.C.; Shaikh, S.S. Assessment of anti-inflammatory potential of Sesbania bispinosa Linn. leaf extracts and fractions by acute and chronic models. Alex. J. Med. 2016, 52, 289–293. [Google Scholar] [CrossRef] [Green Version]
  133. Kapoor, B.B.S.; Purohit, V. Sterol contents from some fabaceous medicinal plants of Rajasthan desert. Indian J. Pharm. Biol. Res. 2013, 1, 13–15. [Google Scholar] [CrossRef]
  134. Tamboli, S.A. Anti-inflammatory activity of Sesbania grandiflora. Ind. Drug. 1996, 33, 504–506. [Google Scholar]
  135. Thamboli, S.A.; Pal, S.C.; Kasture, S.B. Analgesic and antipyretic activity of flowers of Sesbania grandiflora. Ind. Drugs-Bombay 2000, 37, 95–98. [Google Scholar]
  136. Laladhas, K.P.; Cheriyan, V.T.; Puliappadamba, V.T.; Bava, S.V.; Unnithan, R.G.; Vijayammal, P.L.; Anto, R.J. A novel protein fraction from Sesbania grandiflora shows potential anticancer and chemopreventive efficacy, in vitro and in vivo. J. Cell Mol. Med. 2010, 14, 636–646. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  137. Ponnanikajamideen, M.; Nagalingam, M.; Vanaja, M.; Malarkodi, C.; Rajeshkumar, S. Anticancer activity of different solvent extracts of Sesbania grandiflora against neuroblastima (imr-32) and colon (ht-29) cell lines. Eur. J. Biomed. Pharm. Sci. 2015, 2, 509–517. [Google Scholar]
  138. Catalogue of Life. Available online: (accessed on 18 October 2022).
  139. Bús, C.; Tóth, B.; Stefkó, D.; Hohmann, J.; Vasas, A. Family Juncaceae: Promising source of biologically active natural phenanthrenes. Phytochem. Rev. 2018, 17, 833–851. [Google Scholar] [CrossRef] [Green Version]
  140. El-Shamy, A.I.; Abdel-Razek, A.F.; Nassar, M.I. Phytochemical review of Juncus L. genus (Fam. Juncaceae). Arab. J. Chem. 2015, 8, 614–623. [Google Scholar] [CrossRef]
  141. Rodrigues, M.J.; Gangadhar, K.N.; Vizetto-Duarte, C.; Wubshet, S.G.; Nyberg, N.T.; Barreira, L.; Varela, J.; Custódio, L. Maritime halophyte species from Southern Portugal as sources of bioactive molecules. Mar Drugs. 2014, 12, 2228–2244. [Google Scholar] [CrossRef] [Green Version]
  142. Rodrigues, M.J.; Vizetto-Duarte, C.; Gangadhar, K.N.; Zengin, G.; Mollica, A.; Varela, J.; Custódio, L. In vitro and in silico approaches to unveil the mechanisms underlying the cytotoxic effect of juncunol on human hepatocarcinoma cells. Pharm. Rep. 2018, 70, 896–899. [Google Scholar] [CrossRef] [PubMed]
  143. Catalogue of Life. Available online: (accessed on 18 October 2022).
  144. Lindamulage, I.K.; Soysa, P. Evaluation of anticancer properties of a decoction containing Adenanthera pavonina L. and Thespesia populnea L. BMC Compl. Altern. Med. 2016, 16, 70. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  145. Gopalakrishnan, A.; Kariyil, B.J.; John, R.; Usha, P.A. Phytochemical evaluation and cytotoxic potential of chloroform soluble fraction of methanol extract of Thespesia populnea in human breast cancer cell lines. Pharm. Mag. 2019, 15, 150. [Google Scholar] [CrossRef]
  146. Catalogue of Life. Available online: (accessed on 18 October 2022).
  147. Vuong, Q.V.; Chalmers, A.C.; Jyoti Bhuyan, D.; Bowyer, M.C.; Scarlett, C.J. Botanical, phytochemical, and anticancer properties of the Eucalyptus species. Chem. Biodiv. 2015, 12, 907–924. [Google Scholar] [CrossRef]
  148. Hrubik, J.D.; Kaišarević, S.N.; Glišić, B.D.; Jovin, E.Đ.; Mimica-Dukić, N.M.; Kovačević, R.Z. Myrtus communis and Eucalyptus camaldulensis cytotoxicity on breast cancer cells. Zb. Matice Srp. Za Prir. Nauk. 2012, 123, 65–73. [Google Scholar] [CrossRef]
  149. Catalogue of Life. Available online: (accessed on 18 October 2022).
  150. Xu, Z.; Chang, L. Plantaginaceae. In Identification and Control of Common Weeds; Springer: Singapore, 2017; pp. 339–374. [Google Scholar]
  151. Bora, M.S.; Sarma, K.P. Phytoremediation of heavy metals/metalloids by native herbaceous macrophytes of wetlands: Current research and perspectives. In Emerging Issues in the Water Environment during Anthropocene; Springer: Singapore, 2020; pp. 261–284. [Google Scholar]
  152. Smith, E.; Palethorpe, H.M.; Tomita, Y.; Pei, J.V.; Townsend, A.R.; Price, T.J.; Young, J.P.; Yool, A.J.; Hardingham, J.E. The purified extract from the medicinal plant Bacopa monnieri, bacopaside II, inhibits growth of colon cancer cells in vitro by inducing cell cycle arrest and apoptosis. Cells 2018, 7, 81. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  153. Koczurkiewicz, P.; Łojewski, M.; Piska, K.; Michalik, M.; Wójcik-Pszczoła, K.; Szewczyk, A.; Halaszuk, P.; Pekala, E.; Muszyńska, B. Chemopreventive and anticancer activities of Bacopa monnieri extracted from artificial digestive juices. Nat. Prod. Commun. 2017, 12, 337–342. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  154. Yamagata, K.; Izawa, Y.; Onodera, D.; Tagami, M. Chlorogenic acid regulates apoptosis and stem cell marker-related gene expression in A549 human lung cancer cells. Mol. Cell Biochem. 2018, 441, 9–19. [Google Scholar] [CrossRef] [PubMed]
  155. Rezaei-Seresht, H.; Cheshomi, H.; Falanji, F.; Movahedi-Motlagh, F.; Hashemian, M.; Mireskandari, E. Cytotoxic activity of caffeic acid and gallic acid against MCF-7 human breast cancer cells: An in silico and in vitro study. Avicenna J. Phytomed. 2019, 9, 574. [Google Scholar] [CrossRef] [PubMed]
  156. Gheena, S.; Ezhilarasan, D. Syringic acid triggers reactive oxygen species-mediated cytotoxicity in HepG2 cells. Hum. Exper. Toxicol. 2019, 38, 694–702. [Google Scholar] [CrossRef] [PubMed]
  157. Aithal, M.G.; Rajeswari, N. Bacoside A induced sub-G0 arrest and early apoptosis in human glioblastoma cell line U-87 MG through notch signalling pathway. Brain Tumor Res. Treat. 2019, 7, 25. [Google Scholar] [CrossRef] [PubMed]
  158. Goncalves, S.; Romano, A. The medicinal potential of plants from the genus Plantago (Plantaginaceae). Ind. Crop. Prod. 2016, 83, 213–226. [Google Scholar] [CrossRef]
  159. Poor, M.H.S.; Khatami, M.; Azizi, H.; Abazari, Y. Cytotoxic activity of biosynthesized Ag nanoparticles by Plantago major towards a human breast cancer cell line. Rend. Lincei 2017, 28, 693–699. [Google Scholar] [CrossRef]
  160. Catalogue of Life. Available online: (accessed on 18 October 2022).
  161. Caperta, A.D.; Róis, A.S.; Teixeira, G.; Garcia-Caparros, P.; Flowers, T.J. Secretory structures in plants: Lessons from the Plumbaginaceae on their origin, evolution and roles in stress tolerance. Plant Cell Environ. 2020, 43, 2912–2931. [Google Scholar] [CrossRef]
  162. Singh, K.; Naidoo, Y.; Baijnath, H. A comprehensive review on the genus Plumbago with focus on Plumbago auriculata (Plumbaginaceae). Afr. J. Trad. Compl. Altern. Med. 2018, 15, 199–215. [Google Scholar] [CrossRef] [Green Version]
  163. Crespo, M.B.; Lledó, M.D. Two new North African genera related to Limoniastrum (Plumbaginaceae). Bot. J. Linn. Soc. 2000, 132, 165–174. [Google Scholar] [CrossRef]
  164. Lledó, M.D.; Crespo, M.B.; Cox, A.V.; Fay, M.F.; Chase, M.W. Polyphyly of Limoniastrum (Plumbaginaceae): Evidence from DNA sequences of plastid rbcL, trnL intron and trnL-F intergene spacer. Bot. J. Linn. Soc. 2000, 132, 175–191. [Google Scholar] [CrossRef] [Green Version]
  165. Ozenda, P. Flore du Sahara, 2nd ed.; CNRS: Paris, France, 1983. [Google Scholar]
  166. Dijkema, K.S.; Beeftink, W.G.; Doody, J.P.; Géhu, J.M.; Heydemann, B.; Rivas Martinez, S. La Végétation Halophile en Europe (Prés Salés); Council of Europe: Strasbourg, France, 1984. [Google Scholar]
  167. Barone, R.; Scholz, S.; Mesa, R. Limoniastrum monopetalum (L.) Boiss. (Plumbaginaceae), adición a la flora de Fuerteventura (Islas Canarias). Bot. Macarones. 1995, 21, 59–60. [Google Scholar]
  168. Ferrer-Gallego, P.P.; Iamonico, D.; Iberite, M.; Laguna, E.; Crespo, M.B. Lectotypification of two names in Limoniastrum (Plumbaginaceae). Taxon 2014, 63, 1342–1346. [Google Scholar] [CrossRef] [Green Version]
  169. Krifa, M.; Alhosin, M.; Muller, C.D.; Gies, J.P.; Chekir-Ghedira, L.; Ghedira, K.; Mely, I.; Bronner, C.; Mousli, M. Limoniastrum guyonianum aqueous gall extract induces apoptosis in human cervical cancer cells involving p16 INK4A re-expression related to UHRF1 and DNMT1 down-regulation. J. Exp. Clin. Cancer Res. 2013, 32, 30. [Google Scholar] [CrossRef]
  170. Medini, F.; Bourgou, S.; Lalancette, K.; Snoussi, M.; Mkadmini, K.; Coté, I.; Abdelly, C.; Legault, J.; Ksouri, R. Phytochemical analysis, antioxidant, anti-inflammatory, and anticancer activities of the halophyte Limonium densiflorum extracts on human cell lines and murine macrophages. S. Afr. J. Bot. 2015, 99, 158–164. [Google Scholar] [CrossRef]
  171. Catalogue of Life. Available online: (accessed on 18 October 2022).
  172. Hodkinson, T.R. Evolution and taxonomy of the grasses (Poaceae): A model family for the study of species-rich groups. Ann. Plant. Rev. Online 2018, 1, 255–294. [Google Scholar] [CrossRef]
  173. Ashokkumar, K.; Selvaraj, K.; Muthukrishnan, S.D. Cynodon dactylon (L.) Pers.: An updated review of its phytochemistry and pharmacology. J. Med. Plants Res. 2013, 7, 3477–3483. [Google Scholar] [CrossRef]
  174. Khlifi, D.; Hayouni, E.A.; Valentin, A.; Cazaux, S.; Moukarzel, B.; Hamdi, M.; Bouajila, J. LC-MS analysis, anticancer, antioxidant and antimalarial activities of Cynodon dactylon L. extracts. Ind. Crop. Prod. 2013, 45, 40–247. [Google Scholar] [CrossRef]
  175. El Molla, S.G.; Motaal, A.A.; El Hefnawy, H.; El Fishawy, A. Cytotoxic activity of phenolic constituents from Echinochloa crus-galli against four human cancer cell lines. Rev. Bras. Farmacogn. 2016, 26, 62–67. [Google Scholar] [CrossRef] [Green Version]
  176. Catalogue of Life. Available online: (accessed on 18 October 2022).
  177. Xu, Z.; Deng, M. Portulacaceae. In Identification and Control of Common Weeds; Springer: Dordrecht, The Netherlands, 2017; pp. 319–323. [Google Scholar]
  178. Syed, S.; Fatima, N.; Kabeer, G. Portulaca oleracea L.: A mini review on phytochemistry and pharmacology. Int. J. Biol. Biotechnol. 2016, 13, 637–641. [Google Scholar]
  179. Zhou, Y.X.; Xin, H.L.; Rahman, K.; Wang, S.J.; Peng, C.; Zhang, H. Portulaca oleracea L.: A review of phytochemistry and pharmacological effects. BioMed. Res. Int. 2015, 2015, 925631. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  180. Zheng, G.Y.; Qu, L.P.; Yue, X.Q.; Gu, W.; Zhang, H.; Xin, H.L. Portulacerebroside A induces apoptosis via activation of the mitochondrial death pathway in human liver cancer HCCLM3 cells. Phytochem. Lett. 2014, 7, 77–84. [Google Scholar] [CrossRef]
  181. Farshori, N.N.; Al-Sheddi, E.S.; Al-Oqail, M.M.; Musarrat, J.; Al-Khedhairy, A.A.; Siddiqui, M.A. Cytotoxicity assessments of Portulaca oleracea and Petroselinum sativum seed extracts on human hepatocellular carcinoma cells (HepG2). Asian Pac. J. Cancer Prev. 2014, 15, 6633–6638. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  182. Al-Sheddi, E.S.; Farshori, N.N.; Al-Oqail, M.M.; Musarrat, J.; Al-Khedhairy, A.A.; Siddiqui, M.A. Portulaca oleracea seed oil exerts cytotoxic effects on human liver cancer (HepG2) and human lung cancer (A-549) cell lines. Asian Pac. J. Cancer Prev. 2015, 16, 3383–3387. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  183. Catalogue of Life. Available online: (accessed on 18 October 2022).
  184. Nebula, M.; Harisankar, H.S.; Chandramohanakumar, N. Metabolites and bioactivities of Rhizophoraceae mangroves. Nat. Prod. Bioprosp. 2013, 3, 207–232. [Google Scholar] [CrossRef]
  185. Liao, P.C.; Hwang, S.Y.; Huang, S.; Chiang, Y.C.; Wang, J.C. Contrasting demographic patterns of Ceriops tagal (Rhizophoraceae) populations in the South China sea. Austr. J. Bot. 2011, 59, 523–532. [Google Scholar] [CrossRef]
  186. Urashi, C.; Teshima, K.M.; Minobe, S.; Koizumi, O.; Inomata, N. Inferences of evolutionary history of a widely distributed mangrove species, Bruguiera gymnorrhiza, in the Indo-West Pacific region. Ecol. Evol. 2013, 3, 2251–2261. [Google Scholar] [CrossRef]
  187. Warsinah, E.; Sismindari, A.; Susidarti, R.A. Antiproliferative effect and apoptosis induced in human cell lines by Brugiera gymnorhiza barks methanol extract. Indones. J. Cancer Chemoprevent. 2011, 2, 299–303. [Google Scholar] [CrossRef] [Green Version]
  188. Catalogue of Life. Available online: (accessed on 18 October 2022).
  189. Shah, V.V.; Shah, N.D.; Patrekar, P.V. Medicinal plants from Solanaceae family. Res. J. Pharm. Technol. 2013, 6, 143–151. [Google Scholar]
  190. Tang, W.M.; Chan, E.; Kwok, C.Y.; Lee, Y.K.; Wu, J.H.; Wan, C.W.; Chan, R.Y.K.; Yu, P.H.-F.; Chan, S.W. A review of the anticancer and immunomodulatory effects of Lycium barbarum fruit. Inflammopharmacology 2012, 20, 307–314. [Google Scholar] [CrossRef] [PubMed]
  191. Mao, F.; Xiao, B.; Jiang, Z.; Zhao, J.; Huang, X.; Guo, J. Anticancer effect of Lycium barbarum polysaccharides on colon cancer cells involves G0/G1 phase arrest. Med. Oncol. 2011, 28, 121–126. [Google Scholar] [CrossRef] [PubMed]
  192. Zeng, M.; Kong, Q.; Liu, F.; Chen, J.; Sang, H. The anticancer activity of Lycium barbarum polysaccharide by inhibiting autophagy in human skin squamous cell carcinoma cells in vitro and in vivo. Int. J. Polym. Sci. 2019, 2019, 5065920. [Google Scholar] [CrossRef] [Green Version]
  193. Zhu, P.F.; Dai, Z.; Wang, B.; Wei, X.; Yu, H.F.; Yan, Z.R.; Zhao, X.D.; Luo, X.D. The anticancer activities phenolic amides from the stem of Lycium barbarum. Nat. Prod. Bioprospecting 2017, 7, 421–431. [Google Scholar] [CrossRef]
  194. Lee, Y.T.; Hsieh, Y.L.; Yeh, Y.H.; Huang, C.Y. Synthesis of phenolic amides and evaluation of their antioxidant and anti-inflammatory activity in vitro and in vivo. RSC Adv. 2015, 5, 85806–85815. [Google Scholar] [CrossRef]
  195. Catalogue of Life. Available online: (accessed on 18 October 2022).
  196. Sun, L.; Yang, R.; Zhang, B.; Zhang, G.; Wu, X.; Zhang, W.; Zhang, B.; Chen, T.; Liu, G. Phylogenetic relationships among species of Tamarix (Tamaricaceae) in China. Biochem. Syst. Ecol. 2016, 69, 213–221. [Google Scholar] [CrossRef]
  197. Channa, F.N.; Shinwari, Z.K.; Ali, S.I. Phylogeny of Tamaricaceae using PsbA-trnH nucleotide sequences. Pak. J. Bot. 2018, 50, 983–987. [Google Scholar]
  198. Bahramsoltani, R.; Kalkhorani, M.; Zaidi, S.M.A.; Farzaei, M.H.; Rahimi, R. The genus Tamarix: Traditional uses, phytochemistry, and pharmacology. J. Etnophar. 2020, 246, 112245. [Google Scholar] [CrossRef]
  199. Abaza, M.S.; Al-Attiyah, R.A.; Bhardwaj, R.; Abbadi, G.; Koyippally, M.; Afzal, M. Syringic acid from Tamarix aucheriana possesses antimitogenic and chemo-sensitizing activities in human colorectal cancer cells. Pharm. Biol. 2013, 51, 1110–1124. [Google Scholar] [CrossRef] [Green Version]
  200. Abaza, M.S.I.; Afzal, M.; Al-Attiyah, R.A.J.; Guleri, R. Methylferulate from Tamarix aucheriana inhibits growth and enhances chemosensitivity of human colorectal cancer cells: Possible mechanism of action. BMC Comp. Altern. Med. 2016, 16, 384. [Google Scholar] [CrossRef] [Green Version]
  201. KalamUrfi, M.; Mujahid, M.; Badruddeen, J.; Khalid, M.; Khan, M.I.; Usmani, A. Tamarix gallica: For traditional uses, phytochemical and pharmacological potentials. J. Chem. Pharm. Res. 2016, 8, 809–814. [Google Scholar]
  202. Boulabaa, M.; Tsolmon, S.; Ksouri, R. Anticancer effect of Tamarix gallica extracts on human colon cancer cells involves Erk1/2 and p38 action on G2/M cell cycle arrest. Cytotechnology 2013, 65, 927–936. [Google Scholar] [CrossRef] [Green Version]
  203. Momtazi-borojeni, A.; Behbahani, M.; Sadeghi-Aliabadi, H. Antiproliferative activity and apoptosis induction of crude extract and fractions of Avicennia marina. Iran. J. Bas. Med. Sci. 2013, 16, 1203. [Google Scholar]
  204. Yang, X.W.; Dai, Z.; Wang, B.; Liu, Y.P.; Zhao, X.D.; Luo, X. Antitumor triterpenoid saponin from the fruits of Avicennia marina. Nat. Prod. Bioprospecting 2018, 8, 347–353. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  205. Santhanakrishnan, D.; Perumal, R.K.; Kanth, S.V.; Rao, J.R.; Chandrasekaran, B. Antioxidant and cytotoxic effects of methanolic extract of Salicornia brachiata L. in HepG2 cells. Int. J. Res. Pharm. Sci. 2013, 4, 512–517. [Google Scholar]
  206. Sudarshan, S.M.; Chintalapani, S.; Narasu, M.L. Phytochemical analysis and determination of antibacterial and anticancer activities of Suaeda monoica. Asian J. Pharm. Clin. Res. 2019, 12, 91–96. [Google Scholar]
  207. El-Attar, M.M.; Awad, A.A.; Abdel-Tawab, F.M.; Kamel, H.A.; Ahmad, S.A.; Hassan, A.I. Assessment of cytotoxic and anticancer activity of Zygophyllum album and Suaeda palaestina extracts on human liver cancer cell lines. J. Agric. Sci. 2019, 27, 539–544. [Google Scholar] [CrossRef] [Green Version]
  208. Liu, X.; Liu, F.; Zhao, S.; Guo, B.; Ling, P.; Han, G.; Cui, Z. Purification of an acidic polysaccharide from Suaeda salsa plant and its anti-tumor activity by activating mitochondrial pathway in MCF-7 cells. Carbohydr. Polym. 2019, 215, 99–107. [Google Scholar] [CrossRef]
  209. Sumathi, S.; Dharani, B.; Sivaprabha, J.; Raj, S.; Padma, P.R. In vitro growth inhibitory activity of Prosopis cineraria leaves in MCF-7 breast cancer cell line. Int. J. Res. Pharm. Sci. 2013, 4, 40–44. [Google Scholar]
  210. Alsaraf, K.M.; Mohammad, M.H.; Al-Shammari, A.M.; Abbas, I.S. Selective cytotoxic effect of Plantago lanceolata L. against breast cancer cells. J. Egypt Nat. Cancer Instig. 2019, 31, 10. [Google Scholar] [CrossRef]
  211. Venkateswarlu, G.; Rani, T.S.; Vani, M.; Vineela, P.A.J. In-vitro anticancer activity of petroleum ether extract of Cynodon dactylon. J. Pharm. Phytochem. 2015, 4, 164–168. [Google Scholar]
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Custodio, L.; Garcia-Caparros, P.; Pereira, C.G.; Castelo-Branco, P. Halophyte Plants as Potential Sources of Anticancer Agents: A Comprehensive Review. Pharmaceutics 2022, 14, 2406.

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Custodio L, Garcia-Caparros P, Pereira CG, Castelo-Branco P. Halophyte Plants as Potential Sources of Anticancer Agents: A Comprehensive Review. Pharmaceutics. 2022; 14(11):2406.

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Custodio, Luísa, Pedro Garcia-Caparros, Catarina Guerreiro Pereira, and Pedro Castelo-Branco. 2022. "Halophyte Plants as Potential Sources of Anticancer Agents: A Comprehensive Review" Pharmaceutics 14, no. 11: 2406.

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