Current knowledge on genus Bassia All.: a comprehensive review on traditional use, phytochemistry, pharmacological activity, and nonmedical applications

Dagmara Wróbel

Bassia All. is a genus from the Amaranthaceae family, which was created by merging selected species belonging to the former Bassia and Kochia genera with those classified to Chenolea, Londesia, Kirilowia and Panderia. The reorganised Bassia genus currently comprises around 20 species, which are annual herbs or perennial subshrubs native to Eurasia and Africa. Bassia plants are well known for their therapeutic applications in folk medicine and traditional medical systems, and they are also used for nonmedical purposes. Some members of this genus, such as Bassia scoparia (syn. Kochia scoparia) is of great medical importance and economic value. The plant is cultivated in some regions of Asia as a crop to collect Kochiae fructus, which is used for both curative and food purposes. Phytochemical studies carried out on Bassia species indicate that these plants synthesize metabolites belonging to different groups of compounds (e.g., triterpene saponins, sterols, flavonoids, fatty acids, lig...

Phytochem Rev https://doi.org/10.1007/s11101-023-09864-1 (0123456789().,-volV) ( 01234567 89().,-volV) Current knowledge on genus Bassia All.: a comprehensive review on traditional use, phytochemistry, pharmacological activity, and nonmedical applications Karolina Grabowska . Weronika Buzdygan . Agnieszka Galanty . Dagmara Wróbel-Biedrawa . Danuta Sobolewska . Irma Podolak Received: 1 July 2022 / Accepted: 7 March 2023 © The Author(s) 2023 Abstract Bassia All. is a genus from the Amaranthaceae family, which was created by merging selected species belonging to the former Bassia and Kochia genera with those classified to Chenolea, Londesia, Kirilowia and Panderia. The reorganised Bassia genus currently comprises around 20 species, which are annual herbs or perennial subshrubs native to Eurasia and Africa. Bassia plants are well known for their therapeutic applications in folk medicine and traditional medical systems, and they are also used for nonmedical purposes. Some members of this genus, such as Bassia scoparia (syn. Kochia scoparia) is of great medical importance and economic value. The plant is cultivated in some regions of Asia as a crop to collect Kochiae fructus, which is used for both curative and food purposes. Phytochemical studies carried out on Bassia species indicate that these plants synthesize metabolites belonging to different groups of compounds (e.g., triterpene saponins, sterols, flavonoids, fatty acids, lignanamides, alkaloids, organic acids). Some of the Supplementary Information The online version contains supplementary material available at https://doi.org/10.1007/s11101-023-09864-1. K. Grabowska · W. Buzdygan · A. Galanty · D. Wróbel-Biedrawa · D. Sobolewska · I. Podolak (&) Department of Pharmacognosy, Medical College, Jagiellonian University, 9 Medyczna Street, Kraków, Poland e-mail: irma.podolak@uj.edu.pl structures are rarely found in the plant kingdom. Biological activity studies carried out on Bassia plants revealed various effects exerted by extracts and isolated compounds, including anti-inflammatory, cytotoxic, antioxidant, antimicrobial, hypoglycemic, anti-obesity, etc. Modern research explained some of the mechanisms of action. This review covers literature from 1935 to 2022, and assembles and discusses data on phytochemistry, biological activity, as well as medical and nonmedical use of the representatives of the genus Bassia. In this review we present the current state of knowledge about the plants of the genus. Keywords Bassia · Kochia · Phytochemistry · Activity · Medical plant · Saponins Abbreviations AAPH 2,2′-Azobis-(2-amidinopropane) ABTS 2,2′-Azino-bis(3-ethylbenzothiazoline-6sulphonic acid) diammonium salt AKT Protein kinase B ALI Acute lung injury ALS Acetolactate synthase ALT Serum alanine transaminase ANP Atrial natriuretic peptide AST Aspartate transaminase BACE-1 β-Site amyloid precursor protein cleaving enzyme 1 Bax Bcl-2-associated X protein BDE Bond dissociation enthalpy 123 Phytochem Rev BHT CG COX-2 CREB DCM DNCB DNFB DPAR DPPH dw ERK FA FoxO FRAP GAE HIF-1α HNE HO i.p. ICAM-1 IL-1β IL-4, IL-5 IL-6 IL-10 INF-γ iNOS i.v. JNK LC50 LDH LPS MAE MAO-B MAPK MCP-1 MIC MPO MMP-9 mTOR NETDNA NF-κB NOS OVA PCA p.o. p38 123 Butylated hydroxytoluene Cathepsin Cyclooxygenase-2 CAMP response element-binding protein Dilated cardiomyopathy 2,4-Dinitrochlorobenzene 1-Fluoro-2,4,-dinitrofluorobenzene Direct passive arthus reaction 2,2′-Diphenyl-1-picrylhydrazyl Dry weight Extracellular-signal-regulated kinase Fatty acid Forkhead family of transcription factors O Ferric reducing antioxidant power Gallic acid equivalents Hypoxia-inducible factor 1-α Human neutrophil elastase Heme oxygenase Intraperitoneally Intercellular adhesion molecule-1 Interleukin-1β Interleukin-4 Interleukin-5 Interleukin-6 Interleukin-10 Interferon-gamma Inducible NO synthase Intravenous C-Jun N-terminal kinases Lethal dose 50% Lactic dehydrogenase Lipopolisaccharide Microwave-assisted extraction Monoamine oxidase-B Mitogen-activated protein kinases, Monocyte chemotactic protein-1 Minimal inhibitory concentration Myeloperoxidase Matrix metalloproteinase-9 Mechanistic target of rapamycin Net-like structures of nuclear DNA Nuclear factor kappa B Nitric oxide synthase Ovalbumin Passive cutaneous anaphylaxis Per os P38-mitogen activated protein kinase PARP PGC-1α PGE2 PI3K PPAR RCA ROS SAR SENP SUMO TAC TFC TLR TPC TBARS TCM TEAC Th TNF-α TrkA VCAM1 VLCFA VEGF VEGFR Poly (ADP) ribose polymerase Peroxisome proliferator-activated receptor-γ coactivator Prostaglandin E Phosphoinositide 3-kinase Peroxisome proliferator-activated receptors Reversed cutaneous anaphylaxis Reactive oxygen species Structure activity-relationship SUMO-specific proteases Small ubiquitin-like modifier protease Total antioxidant capacity Total flavonoid content Toll-like receptor Total phenolic content Thiobarbituric acid reactive substances Traditional Chinese Medicine Trolox equivalent antioxidant capacity Helper T cells Tumor necrosis factor α Tropomyosin receptor kinase A Vascular cell adhesion molecule-1 Very-long-chain fatty acids Vascular endothelial growth factor Vascular endothelial growth factor receptor Introduction The genus Bassia All. is one of the genera found in the Amaranthaceae family. The taxonomy of these plants is problematic, but according to a new reclassification, the genus Bassia comprises circa 20 species (Kaderit and Freitag 2011). Bassia plants are annual herbs or perennial subshrubs native to Eurasia and Africa. As members of the Amaranthaceae family, these species operate C4 photosynthesis, which evolved in dicots to facilitate their growth in arid ecosystems. Therefore, Bassia plants can be found in semideserts or dry steps, and, as halophytes, can exist also in habitats characterized by high salinity. Due to adaptation to different environmental conditions, Bassia species have attracted attention as plants that can be cultivated in regions where other Phytochem Rev Fig. 1 The distribution range of plants in the genus Bassia plant species cannot be planted (Al-Ahmadi et al. 2007; Kaderit and Freitag 2011; Orlovsky et al. 2011; Shelef et al. 2012; Abideen et al. 2015). Today, as can be seen in Fig. 1, Bassia plants are distributed from the Mediterranean region (Europe and North Africa) up to Japan (B. scoparia, B. prostrata, B. hyssopifolia). Several species are characteristic of the arid regions of the Middle East, India, and steps of Central Asia (B. odontoptera, B. indica, B. eriophora, B. stellaris). Some grow in North Africa, in the Sahara region (B. arabica, B. muricata) and South Africa (B. salsoloides, B. dinteri) (Kaderit and Freitag 2011; GBIF 2022). Plants of the Bassia genus, such as B. scoparia, B. prostrata and B. hyssopifolia, have been introduced to North America as high-yielding forage plants with nutritional value or for ornamental purposes. They spread quickly and are currently naturalized in both North and South America (Lauriault et al. 2020; Torbiak et al. 2021; Brignone et al. 2021; Ravet et al. 2021). B. scoparia and B. hyssopifolia have also been introduced to Australia. However, due to the high invasive potential of B. scoparia, a year after introduction, work on the eradication of this plant species began. The elimination process proved successful, and no B. scoparia have been observed in Australia since 2000. In turn, there are some records that B. hyssopifolia still occurs in southeastern Australia, but only in a small area of north-western Victoria state (POWO 2022; Panetta 2015). In Asia, B. scoparia (syn. Kochia scoparia) has great medical importance and economic value. At present, it is cultivated in China and Japan as a crop for both curative and food purposes (Tashiro et al. 2019; Chen et al. 2021; Zou et al. 2021). B. scoparia is a plant recorded in the Chinese Pharmacopeia (Chen et al. 2021), which provides dried fruits (Kochiae fructus, Di-Fu-Zi) which is a TCM product very popular in different parts of Asia. Currently, due to globalization, Kochiae fructus is commercially available worldwide. Given that B. scoparia has welldocumented pro-health effects and is also used as a food, some authors suggest that its fruit should be 123 Phytochem Rev considered for the development of functional foods (Chen et al 2017). Apart from B. scoparia, other Bassia species have also been recommended in traditional medical systems, especially in regions where they grow as wild plants (Hammiche and Maiza 2006; Khare 2007; Altundag and Ozturk 2011; Youssef 2013; Moein et al. 2015; Umair et al. 2017). In addition to the therapeutic value of these plants, they have also been used for nonmedical purposes. For example, some plants of the genus Bassia are being appreciated as human food and livestock feed because of their nutritional value. (Nadelcheva et al. 2007; El Shaer 2010; Khan et al. 2013; Youssef 2013; Akhtar et al. 2018). In recent years, several studies have provided new data on the chemistry and biological activity of Bassia species (Xiae et al. 2018; Houlihan et al. 2019; Abd-ElGawad et al. 2020; Gancheva et al. 2020; Othman et al. 2021a; 2021b; Othman et al. 2022; Said et al. 2021; Petruk et al. 2021). It is noteworthy that not only B. scoparia but also other hitherto unexplored Bassia species attracted scientific attention. Modern research also investigated the potential for new applications of plants from this genus (Abideen et al. 2015; Athinarayanan et al. 2018; Aihemaiti et al. 2018; El-Katori and Al-Mhyawi 2019; Song et al. 2021; Shi et al. 2022; Hozhabralsadat et al. 2022). To the best of our knowledge, there are no papers reviewing available scientific data on the Bassia genus. Although there are some reviews devoted to B. scoparia, they focused exclusively on one part of this plant, namely Kochiae fructus (Al-Sanafi et al. 2018; Zou et al. 2021). Therefore, the main objective of the current review is to provide the first comprehensive overview of the Bassia genus, including the latest information on its phytochemistry, biological activity, as well as its medical and nonmedical use. Methods The search of scientific literature was carried out in various databases including PubMed, Scopus, Google Scholar, Science Direct, and Embase, and covered the period up to 2022 June 17. The key words (as individuals and their different combinations) used to search were: Bassia, Kochia, “phytochemistry”, 123 “phytochemical”, “taxonomic”, “distribution”, “saponins”, “phenolics”, “triterpenes”, “fatty acids”, “flavonoids”, “quantification”, “sterol”, “alkaloid”, “terpenes”, “compound”, “nitrates”, “organic acid”, “oil”, “essential oil”, “monosaccharides”, “carotenoids”, “activity”, “pharmacological”, “extract”, “anti-inflammatory”, “inflammatory”, “oedema”, “anti-allergic”, “psoriasis”, “eczema”, “wound healing”, “traditional medicine”, “food”, “ethnobotany”, “TCM”, “animal model”, “antibacterial”, “antifungal”, “antiparasitic”, “phytoremediation”, “gastric”, “hypoglycemic”, “antioxidant”, “toxicity”, “cytotoxicity”, “cancer cell”, “mechanism”. As key words, the full Latin name of representatives of the genus Bassia (according to the new classification proposed by Kaderit and Freitag (2011), as well as their synonyms and former Latin name (according to the previous taxonomic classification) (as individual or in combination with the above-mentioned keywords) were used. According to the rules of the specific databases, Boolean operators were also used. No filters or time restrictions were applied for publications during the search. Furthermore, the reference lists of all selected reports were searched to identify potentially eligible publications. Only publications that had available full texts were included. Conference communications, posters, and the thesis were excluded. The available information on Bassia species has been divided into several sections, i.e. taxonomy, distribution, phytochemistry, biological activity, medical use, nonmedical use, and toxicology. Publications outside the scope of these thematic sections were rejected. Moreover, publications on the phytochemistry of Bassia species were thoroughly checked, and articles that contained information on preliminary phytochemical analyses of extracts, based only on characteristic reactions of groups of compounds (e. g. foam test), which are not specific, were excluded from the phytochemistry section. ChemDraw 19.0 software was used to draw the chemical structures of compounds. The graphs were performed using Excel 365 software (Microsoft Office) software. The illustrations were made with CorelDraw 2021.5 software. Phytochem Rev Taxonomy Species of the genus Bassia were previously classified to the Chenopodiaceae family. However, the Chenopodiaceae family is currently included in the Amaranthaceae family, which is taxonomically complicated. According to a new classification, proposed by Kadereit and Freitag (2011) based on molecular phylogeny and morphological studies, species belonging to the Bassia and Kochia genera were combined in a subclade renamed Bassia All., which is part of the Bassia/Camphorosma clade. The reorganized genus Bassia All. comprises currently circa 20 species: B. aegyptiaca Turki, El Shayeb & F. Shehata, Bassia angustifolia (Turcz.) Freitag & G. Kadereit, Bassia arabica (Boiss.) Maire & Weiller, Bassia dinteri (Botsch.) A.J. Scott, Bassia eriophora (Schrad.) Asch., Bassia hyssopifolia (Pall.) Kuntze, Bassia indica (Wight) A.J. Scott, Bassia laniflora (S. G. Gmel.) A.J. Scott, Bassia lasiantha Freitag & G. Kadereit, Bassia littorea (Makino) Freitag & G. Kadereit, Bassia muricata (L.) Asch., Bassia odontoptera (Schrenk) Freitag & G. Kadereit, Bassia pilosa (Fisch. & C.A. Mey.) Freitag & G. Kadereit, Bassia prostrata (L.) A.J. Scott, Bassia salsoloides (Fenzl) A.J. Scott, Bassia scoparia (L.) A.J. Scott, Bassia stellaris (Moq.) Bornm., Bassia tianschanica (Pavlov) Freitag & G. Kadereit, Bassia tomentosa (Lowe) Maire & Weiller, Bassia villosissima (Bong. & C.A. Mey.) (Kadereit and Freitag 2011; Hernández-Ledesma et al. 2015). Although some authors indicate ambiguities in the proposed classification, and distinguish more species, for example Bassia eriantha and Kochia monticola, nevertheless these species are denoted in the current classification B. eriophora and B. pilosa, respectively (Akhani and Khoshravesh 2013; Sukhorukov and Kushunina 2020). Apart from Kochia/Bassia species, plants from Chenolea, Londesia, as well as Kirilowia and Panderia have also been transferred into Bassia taxonomic group and renamed Bassia. On the other hand, after taxonomic reorganisation, some formerly known Bassia or Kochia species were excluded from Bassia subclade and grouped in new genera; Eokochia (K. saxicola), Spirobassia (B. hirsuta), Sedobassia (K. sedoides) and Grubovia (B. dasyphylla, K. melanoptera, K. krylovii). Additionally, two perennial species native to the United States, formerly known as Kochia americana S. Watson and Kochia californica S. Watson are now included in the genus Neokochia as Neokochia americana (S. Watson) G.L. Chu & S.C. Sand and N. californica (S. Watson) G.L. Chu & S.C. Sand (Kaderit and Freitag 2011; Kadereit et al. 2014; Hernández-Ledesma et al. 2015). Despite the new classification and nomenclature proposed by Kaderit and Freitag, both new names and their synonyms coexist in the scientific literature. Interestingly, for some species, such as Bassia scoparia, the term Kochia scoparia is used even more frequently. This may be related to the fact that K. scoparia is a plant commonly used in TCM, and according to Flora of China records, Kochia and Bassia were not merged but are still treated as separate genera (Chen et al. 2021). Therefore, in order to avoid any confusion, in the current review, the new terminology according to Kadereit and Freitag (2011) is used for all species of the genus Bassia. The importance of plants of the genus Bassia Numerous ethnobotanical surveys carried out in different regions of Asia and Europe, records found in herbal books referring to traditional medicinal systems, and scientific literature reports, all underline that Bassia plants have been intensively used by humans for therapeutic purposes and as nonmedical products (Hammiche and Maiza 2006; Khare 2007; Nadelcheva et al. 2007; Altundag and Ozturk 2011; Umair et al. 2017; Lin et al. 2021) (see Table 1). It should be mentioned that modern research revealed some new potential applications of Bassia plants, which are currently being investigated. Medical uses of Bassia species Representatives of the genus Bassia have been used in traditional medicine to treat various aliments (see Table 1). One of the most important species from the therapeutic point of view is B. scoparia. The dried fructus of B. scoparia, known as Di-Fu-Zi, has been valued in Traditional Chinese Medicine (TCM). Its application is based on ethnopharmacological data, and modern scientific research (Zou et al. 2021). Today, Kochiae fructus is recommended for urinary system disorders, such as frequent urination or 123 Phytochem Rev Table 1 The ethnobotanical importance of Bassia plants and their use in medical systems, including TCM Plant name Usage Plant part/Form Country (Region) References B. scoparia (syn. K. scoparia) Cardiac tonic Fruits, leaves/ oral China Kirtikar and Basu (1935), Khare (2007), Armenia, Uzbekistan Akopian et al. (2020) Tonic, hypotensive and breath stimulating agent Undef Undef Diuretic Fruits, leaves/oral China Kirtikar and Basu (1935), Khare (2007), Urinary system disorders (frequent urination, urinary incontinence) Gonorrhea Fruit China Zou et al. (2021) Fruit/seed/ decoction China (Gaomi) Lin et al. (2021) Fruit Korea Im et al. (2016), Whang and Hahn (1991) China Skin diseases (itching, eczema, sores), swelling, Fruit urticaria Taiwan Chien et al. (2013), Chen et al. (2015) Dermatitis Fruit Korea Whang and Hahn (1991) Atopic dermatitis Fruit Taiwan Chen et al. (2016) Psoriasis Fruit Taiwan Weng et al. (2016) Male impotence, vaginal discharge, vaginal fungal infections, leucorrhea Fruit China Liu et al. (2012), Zou et al. (2021) Improvement of eyesight, hearing, pain in head, Fruit eyes, ears China Zou et al. (2021) Food/vegetable China (Inner Mongolia) Sachula et al. (2020) Aerial parts/ fried/ steamed with flour Shoots and tender leaves Myanmar Shin et al. (2018) Aerial parts China (Heihe valley) Kang et al. (2012) Food (Tonburi) Fresh fruits Japan Tashiro et al. (2019) Medicinal use household items, Undef South Korea Lee et al. (2017) Chung et al. (2016) Brooms Fodder B. eriophora Aerial parts Bulgaria, Romania, Macedonia, Italy Nedelcheva et al. (2007) Aerial parts Spain Gras et al. (2020) Fresh/ dry aerial parts Pakistan (GilgitBaltistan) Khan et al. (2013) Khan and Qaiser (2006) Ornamental, fibres (mats, baskets) – Pakistan Antirheumatic, snake bite, vermifugal Whole plant, seed oil Saudi Arabia (Onaizah Youssef (2013) province) Food/vegetables and fodder Leaves, stem Pakistan (Karak district) Akhtar et al. (2018) Alzheimer, gingivitis, hair loss Leaf, twig/ infusion Iran (Darab region) Moein et al. (2015) Kidney diseases, antirheumatic, ulcers Seeds, leaves, flowers, seed oil/gargle Saudi Arabia (Onaizah Youssef (2013) province) Analgesic, antiseptic, anti-inflammatory Undef Algiers (Sahara, Oued Righ) Lakhdari et al. (2016) Skin diseases Leaf/cataplasm aerial parts/infusion/oral Morocco (Tata Province) Abouri et al. (2012) Skin diseases: boils, dermatosis, pustules, infected wounds Leaf/poultice Algeria (Tassili N’Ajjer’s) Hammiche and Maiza (2006) Diarrhoea Aerial parts/infusion/ internal Algeria (Tassili N’Ajjer’s) Hammiche and Maiza (2006) Birds appetizer agent Seeds and fruits Saudi Arabia (Riyadh) Sher and Aldosari (2013) B. muricata hypoglycaemic 123 Phytochem Rev Table 1 The ethnobotanical importance of Bassia plants and their use in medical systems, including TCM Plant name B. indica Usage Plant part/Form Country (Region) References Skin diseases: sores Seed oil Saudi Arabia (Tabuk) Al-Harbi (2017) Fodder Seeds, leaves, flowers, seed oil, gargle Saudi Arabia (Onaizah) Youssef (2013) Whole plant Libya (Mediterranean Coast) Louhaichi et al. (2011) Saudi Arabia (Onaizah) Youssef (2013) Sand fixation Whole plant Fuel Seeds, leaves, flowers, seed oil, gargle Heart tonic, cardiac stimulant (weak and irregular heart) Whole plant/ decoction, oil, gargle Saudi Arabia (Onaizah) Youssef (2013) Undef India Khare (2007) Undef India Kirtikar and Basu (1935) Undef India (Rajasthan) Tripathi et al. (1996) Roots, whole plant/ extract Pakistan (Faisalabad) Ahmad et al. (2015) Diuretic Fruits, leaves/ decoction, oil; oral, gargle Pakistan (Hafizabad) Umair et al. (2017) Toothache Fruits, leaves/ decoction, oil; oral, gargle Pakistan (Faisalabad) Ahmad et al. (2015) Blood pressure, constipation, vomiting, internal Whole plant/decoction worms Pakistan (Karak) Khan et al. (2018) Fodder Fresh/ dry aerial parts Pakistan (Karak) Rashid and Marwat (2006) Fresh/ dry aerial parts Pakistan (Tank) Badshah et al. (2012) Fresh/ dry aerial parts Pakistan (GilgitBaltistan) Khan et al. (2013) Fresh/ dry aerial parts Pakistan (Charsadda) Khan and Badshah (2019) Whole plant Libya (Mediterranean Coast) Louhaichi et al. (2011) Fresh/ dry aerial parts fresh Roots, whole plant/ tooth brushes, extract Fresh/ dry aerial parts fresh Pakistan (Muzaffar Garh) Ajaib et al. (2015) Ornamental Fuel Fresh/ dry aerial parts Pakistan (Karak) Rashid and Marwat (2006) Fresh/ dry aerial parts Pakistan (Tank) Badshah et al. (2012) Whole plant Libya (Mediterranean Coast) Louhaichi et al. (2011) Sand fixation 123 Phytochem Rev Table 1 continued7 Plant name Usage Plant part/Form Country (Region) References B. prostrata Urinary system disorders Aerial parts/ decoction/ internal Turkey, (Anatolia) Altundag and Ozturk (2011) Anthelminthic Undef. Armenia, Uzbekistan Akopian et al. (2020) Soap for washing utensils Undef. Pakistan (Karak) Khan et al. (2011) Fodder Aerial parts Kazakhstan Ryabushkina et al. (2008) Whole plant Pakistan (Gilgit) Khan and Khatoon (2008) Aerial parts Bulgaria Nedelcheva, et al. (2007) Branches Pakistan (Baltistan) Abbas et al. (2019) Toothache Brooms B.odontoptera (former name: K. iranica) Roots/tooth brush, whole plant Fuel Branches Fodder, fuel wood Undef. Pakistan (Balochistan) Durrani et al. (2010) Ornamental, fibres (mats, baskets) Undef. Pakistan Khan and Qaiser (2006) undef. The plant part and the form of application were not specified urinary incontinence, and as a remedy in vaginal discharge. Kochiae fructus is also applied in skin, eyes, and ear diseases (Im et al. 2016; Lin et al. 2021; Zou et al. 2021). Recent studies have shown that DiFu-Zi is one of the most prescribed single herbal preparations in traditional Chinese medicine to treat dermatological problems such as eczema, urticaria, psoriasis, and pediatric atopic dermatitis (Chien et al. 2013; Chen et al. 2015, 2016; Weng et al. 2016). The fruit is also used as a key ingredient in herbal formulas to reduce itching, swelling, and sores (Chien et al. 2013; Chen et al. 2015; Weng et al. 2016). In China, the fruits and leaves of B. scoparia have been known as cardiac tonics (Kirtikar and Basu 1935). People in Armenia and Uzbekistan have also used B. scoparia as a breath stimulator and hypotensive agent (Akopian et al. 2020). In addition to B. scoparia, other species of the genus have also been valued as medicinal plants. In the Middle East, as well as in India, the decoction of the entire plant of B. indica has been recommended as a cardiac stimulant (Khare 2007; Youssef 2013; Umair et al. 2017). Ethnobotanical surveys revealed that aerial parts of B. muricata, and B. prostrata have been used by inhabitants of Saudi Arabia and Turkey to treat kidney diseases and as antirheumatic agents 123 (Altundag and Ozturk 2011; Youssef 2013). Furthermore, B. muricata leaves poultices have been applied to the skin as a remedy for boils, ulcers, dermatosis, pustules, and infected wounds in folk medicine of different regions of North Africa and Saudi Arabia (Hammiche and Maiza 2006; Abouri et al. 2012; AlHarbi 2017). B. eriophora has traditionally been used in Iran as an infusion of leaves or twigs in the treatment of gingivitis, hair loss, and the symptoms of Alzheimer’s disease (Moein et al. 2015). Furthermore, recent research has revealed some new applications for Bassia species. B. eriophora was studied as a natural material to obtain cellulose nanofibers. Such structures can be used in various sectors of industry and medicine. Because the produced nanofibers along with their very good mechanical properties were also biocompatible and biodegradable, the possibility of using them in regenerative medicine and tissue engineering should be considered (Athinarayanan et al. 2018). Nonmedical applications of Bassia species In addition to medical applications, Bassia species are also valued as nonmedical products. Fresh fruits of B. scoparia are very popular in Asian cuisine as a Phytochem Rev Fig. 2 Chemical structures of new compounds (at the time of publication) isolated from the genus Bassia 123 Phytochem Rev Fig. 3 Chemical structures of compounds isolated from the genus Bassia: characteristic compound (A), rarely found compounds in the plant kingdom (B) 123 food garnish called Tomburi or ‘plant caviar’ (Tashiro et al. 2019). Some authors also highlight the potential of Kochiae fruits in the development of functional foods (Chen et al. 2017). The shots and tender leaves of B. scoparia are eaten as a vegetable by people from Myanmar and inhabitants of the Heihe valley in China (Kang et al. 2012; Shin et al. 2018). In the inner Mongolian region, aerial parts of the plant are consumed fried or steamed with flour (Sachula et al. 2020). Bassia plants, including B. prostrata, B. scoparia, B. muricata, B. eriophora and B. indica, are also valued as feed for livestock in various districts of Pakistan, Saudi Arabia, and Kazakhstan due to the nutritional values of aerial parts, but equally important to the possibility of their cultivation under unfavourable climatic conditions, even in arid areas (see Table 1) (Khan and Khatoon 2008; Khan et al. 2013; Youssef 2013; Akhtar et al. 2018). In the western United States and the Near East, B. prostrata and B. indica are gaining increasing importance as high-yielding forage plants for grazing (Waldron et al. 2010; El Shaer 2010). Furthermore, ethnobotanical surveys revealed that B. muricata and B. indica have been used by inhabitants of Pakistan and some regions of Saudi Arabia as fuel and firewood (Badshah et al. 2012; Youssef 2013). On the other hand, there are some reports from the USA that another Bassia species, B. prostrata, may be used as a green firebreak, a management tool for wildfires (Harrison et al. 2002). Some reports indicate that aerial parts of B. scoparia have been used in different countries of Europe to produce household items, such as brooms (Nadelcheva et al. 2007; Gras et al. 2020). In turn, the people of Karak district in Pakistan produced a soaplike product from B. prostrata (Khan et al. 2011). B. scoparia, commonly called summer-cypress, is also cultivated as an ornamental plant. Currently, research is being carried out on the possibility of using various species of Bassia (B. indica, B. scoparia) in the phytoremediation of salt- and metal-contaminated soils (Shelef et al. 2012; Moubasher et al. 2015; Aihemaiti et al. 2018; Shi et al. 2022). The possibility of using B. scoparia and its associated rhizosphere microflora is also being considered in the degradation of aromatic hydrocarbons in soils contaminated with crude oil (Moubasher et al. 2015; Song et al. 2021). Furthermore, new potential applications such as the Phytochem Rev Table 2 Saponins isolated from plants of the genus Bassia Aglycone Sugar part Common name of saponin* Species (plant part) References OA C-3: Xyl-(1→3)-GlcA Momordin Ic B. scoparia (FR) Wen et al. (1995), Yoshikawa et al. (1997a, b), Han et al. (2006), Lu et al. (2012b) OA C-3: Xyl-(1→3)-[Glc (1→2)]-β-D-GlcA 2′-O-β-D-glucopyranosyl momordin Ic B. scoparia (FR) OA C-3: Xyl-(1→3)-6-MeGlcA Momordin Ic methyl ester B. scoparia (FR) Wen et al. (1995), Yoshikawa et al. (1997a, b), Han et al. (2006) Wen et al. (1995), Yoshikawa et al. (1997a), Han et al. (2006), Lu et al. (2012b) OA C-3: Xyl-(1→3)-EtGlcA Momordin Ic ethyl ester B. scoparia (FR) Han et al. (2006) OA C-3: Xyl-(1→3)-GlcA C-28: Glc Momordin IIc B. scoparia (FR) Whang and Hahn (1991), Wen et al. (1995), Yoshikawa et al. (1997a), Han et al. (2006), Lu et al. (2012b) OA C-3: Xyl-(1→3)-6-MeGlcA Momordin IIc methyl ester B. scoparia (FR) Han et al. (2006) 2′-O-β-D-glucopyranosyl momordin IIc B. scoparia (FR) Wen et al. (1995), Yoshikawa et al. (1997a), Han et al. (2006), Lu et al. (2012b) Oleanolic acid 3-O-β-Dribopyranosyl-(1→2)-β-Dglucuronopyranoside B. scoparia (FR) Whang and Hahn (1991) C-28: Glc OA C-3: Xyl-(1→3)-[Glc (1→2)]-β-D-GlcA C-28: Glc OA C-3: Rib-(1→2)-GlcA OA C-3: Ara-(1→3)-GlcA Momordin I B. scoparia (FR) Yoshikawa et al. (1997a) OA C-3: Ara (1→3)-GlcA Momordin IIaa B. indica (AP) Mohamed et al. (1998) Oleanolic acid-3-Oglucuronide (Calenduloside E) Oleanolic acid-3-O-βglucopyranoside B. scoparia (FR) Yoshikawa et al. (1997a), Lu et al. (2012b) B. muricata (WP) Kamel et al. (2001) C-28: Glc OA C-3: GlcA OA C-3: Glc OA C-3: GlcA OA C-28: Glc OA C-3: Glc-(1→2)-GlcA Chikusetsusaponin IVa B. muricata (WP) Kamel et al. (2001) B. indica (AP) Mohamed et al. (1998) B. scoparia (FR) Lu et al. (2012b) Chikusetsusaponin V B. indica (AP) Mohamed et al. (1998) Chikusetsusaponin IVa methyl ester B. muricata (AP) Kamel et al. (2001) Oleanolic acid-3,28-βdiglucopyranoside B. muricata (AP) Kamel et al. (2001) C-28: Glc OA C-3: 6‘-Me-GlcA C-28: Glc OA C-3: Glc OA C-3: [3-R1]-6-GlcA C-28: Glc Betavulgaroside IIIa B. indica (AP) Mohamed et al. (1998) OA C-3: [3-R2]-GlcA (2′R,3′S)-3-O-[2′-hydroxy-3′(2-O-glycolyl)-oxopropionic acid-β-Dglucuronopyranosyl]-28-Oβ-D-glucopyranosyl-olean12-en-3β-ol-28-oic acid B. indica (AP) Othman et al. (2021b) C-28: Glc C-28: Glc 123 Phytochem Rev Table 2 continued Aglycone Sugar part Common name of saponin* Species (plant part) References OA C-3: [3-R2]-6-GlcA 3-O-[2′-(2-O-glycolyl)glyoxylyl-β-Dglucuronopyranosyl]-28-Oβ-D-glucopyranosyl-olean12-en-3β-ol-28-oic acida B. indica (AP) Mohamed et al. (1998) Betavulgaroside IVa Betavulgaroside V B. indica (AP) B. indica (AP) Mohamed et al. (1998) Othman et al. (2021b) Betavulgaroside Va B. indica (AP) Mohamed et al. (1998) Oleanolic acid-28-O-β-Dglucopyranosyl ester B. prostrata (WP) Seitimova et al. (2018) C-28: Glc OA OA C-3: [3-R1]-6-GlcA C-3: [Glc(1→2)]- [3R1]-GlcA OA C-3: [Glc(1→2)]- [3R1]-6-GlcA OA C-28: Glc OA C-3: sugar part (Glc) Undef. B. muricata (WP) El-Sayed (1993) OA C-3: sugar part (Glc, Ara) Undef. B. muricata (WP) El-Sayed (1993) OA C-3: sugar part (Glc, Ara, Rha) Undef. B. muricata (WP) El-Sayed (1993) HE C-3: sugar part (nd) Undef. B. muricata (WP) El-Sayed (1993) 22α- hydroxyoleanolic acid Yoshikawa et al. (1997a) C-3: GlcA Kochianoside I B. scoparia (FR) C-3: Xyl(1→3)GlcA Scoparianoside A B. scoparia (FR) Yoshikawa et al. (1997b) Morolic acid C-3: Glc-(1→2)-Xyl (1→3)-GlcA Kochianoside II B. scoparia (FR) C-28: Glc C-28: Glc C-3: Xyl-(1→3)-GlcA 3β,12α- Betulinic acid 3β- dihydroxyolean28,13α-olide Yoshikawa et al. (1997a) C-3: Xyl-(1→3)-GlcA hydroxyolean-13(18)en-28-oic acid Yoshikawa et al. (1997a) Scoparianoside B B. scoparia (FR) Yoshikawa et al. (1997b) C-3: Xyl-(1→3)-GlcA Kochianoside III B. scoparia (FR) Kochianoside IV B. scoparia (FR) Yoshikawa et al. (1997a) C-3: Xyl-(1→3)-GlcA Scoparianoside C B. scoparia (FR) Yoshikawa et al. (1997b) *In the absence of a common name, a chemical term was used a Saponins obtained by Mohamed et al. (1998) as their methyl esters after methylation using ethereal diazomethane; Undef.– undefined—the identity of the saponin cannot be determined due to the lack of a precise compound structure OA Oleanolic acid, HE Hederagenin, FR Fruit, AP Aerial parts, WP Whole plant, Glc β-D-glucoopyranose, GlcA β-Dglucuronopyranose, Ara α-L-arabinopyranose, Xyl β-D-xylopyranose, Rib β-D-ribopyranose, Et Ethyl moiety, Me Methyl moiety, R1 2′-hydroxy-3′-(2-O-glycolyl)-oxo-propionic acid, R2 2′-(2-O-glycolyl)-glyoxylyl use of ethanol extract from fresh parts of the B. muricata as an eco-friendly and effective inhibitor of 123 the corrosion process of the aluminium alloy are also investigated (El-Katori and Al-Mhyawi 2019). Phytochem Rev Phytochemistry The phytochemical studies of representatives of the genus Bassia have been conducted for many years. Notwithstanding, the chemical compositions of only few species of the genus have been investigated so far. Initially, the analyses focused mainly on B. scoparia, but in recent years there has been an increased research interest in phytochemistry of other species of the genus. Until now, more than 300 compounds have been identified in the genus, and some of them were new compounds at the time of publication (Fig. 2). Moreover, several rarely occurring phytochemicals have been detected, including some saponins, flavonoids, steroidal glycosides, or fatty acids. Examples of compounds characteristic of the genus Bassia and at the same time rarely found in the plant kingdom are shown in Fig. 3. Triterpene saponins and triterpenes From a pharmacological point of view, triterpenoid saponins are one of the most important groups of metabolites found in Bassia plants. However, until now, saponins have been isolated solely from three species, B. scoparia, B. indica, and B. muricata. So far, phytochemical studies of Bassia species have led to the isolation of thirty-three different saponins, while twenty-eight of them have been fully characterized by structural studies. These saponins are listed in Table 2. Interestingly, twelve saponins were novel compounds at the time of publication (see Fig. 2). Among saponins, structures with an oleanane skeleton dominated. Within this group, the highest number of saponins represented oleanolic acid glycosides, while compounds with rearranged double bond positions, such as morolic acid and 3β-hydroxyolean-13(18)-en-28-oic acid were less common (Fig. 4). Interestingly, lupane-type saponin was also detected, as well as a fairly rare type of 13,28epoxyoleanane saponins. Sapogenins found in Bassia species are presented in Fig. 4. Almost half of Bassia saponins (42.4%) are bidesmosides, the remaining 57.6% are classified as monodesmosides, usually bearing a sugar chain at C-3 of sapogenin. The sugar part is composed of β-D-glucuronic acid (GlcA), β-D- xylopyranose (Xyl), β-L-arabinofuranose (Ara) and β-D-glucopyranose (Glc). It should be noted that the vast majority (81.8%) of saponins have β-D-glucuronic acid units in the sugar chain attached to C-3 of the aglycone. Among them, 34.4% can be classified as the GOTCAB group (glucuronide oleananetype triterpenoid carboxylic acid 3, 28-O-bidesmosides). It is also worth mentioning that one new saponin (Fig. 2) and five rarely occurring secoglycosidic oleanane saponins, containing acidic constituents at C-3 of glucuronic acid, have been isolated so far exclusively from the aerial parts of B. indica (Mohamed et al. 1998; Othman et al. 2021b). Quantitative research on the content of saponins in Bassia species is limited to Kochiae fructus. Analysis of fruit from various regions of China using the HPLC-ELSD method showed that the saponin level ranges from 3.0 to 5.8% (Xia et al. 2002a; Wang et al. 2014a). In another study, qualitative gas–liquid chromatographic analysis (GLC) of methyl esters of triterpene sapogenins (methyl oleanolate) demonstrated that the sapogenin content ranges from 0.95 to 2.09% in a single seed of B. scoparia, reflecting the content of oleanolic acid-type saponins in plant material (Kernan et al. 1973). Although various saponin compounds have been found in plants of the genus Bassia, the quantitative determination focused on momordin Ic, which is the main saponin of Kochiae fructus (Fig. 3A). Various reports indicate that momordin Ic content in plant material ranges from 0.85 to 3.5% (Xia et al. 2002a; b; Belsevich et al. 2009). According to Hong Kong Chinese Materia Medica Standards, the content of momordin Ic in Kochiae fructus should not be less than 2.3% (HKCMMS 2012). In addition to qualitative studies of the plant material, which reflect the quality of the product, investigations on momordin Ic content were also carried out on various extracts of Kochiae fructus. It can be seen that the amount of momordin Ic in the extract varies greatly depending on the extraction method (Choi et al. 2014; Wang et al. 2014a; Yoo et al. 2017). For example, Wang et al. (2014a), using the HPLC-ELSD method, found that the momordin content in the water extract (under infusion condition for 4 h at room temperature) was 2.88%, while in the 123 Phytochem Rev types of pentacyclic triterpenes, including oleanolic acid, gypsogenin, hederagenin (Setimova et al. 2018; Imran et al. 2017), betulin, betulinic acid and ursolic acid (Imran et al. 2017) were detected in B. prostrata. Oleanolic acid, (Lu et al. 2012b; Zhang et al. 2013) βamyrin and lupeol were also found in B. scoparia, along with some tetracyclic triterpenes such as cycloartenol, 31-norcykloartenol and cycloeucalenol that were detected in Kochiae fructus (Narumi et al. 2001). Sterols, steroidal glucosides and phytoecdysteroids Fig. 4 Types of sapogenins and their share (%) in saponins found in the genus Bassia (graph A), the share (%) of Bassia saponins depending on amount of sugar chains (graph B), the share (%) of Bassia saponins with a specific sugar moiety (graph C) 50% ethanol extract prepared under reflux conditions (4 h) it reached a higher value of 5.75%. In addition to triterpene saponins, free triterpenes have also been reported in Bassia plants. Various 123 Some species of Bassia were analysed for the presence of sterols, and to date, more than 25 sterols have been identified in representatives of the genus (see Table 3). Reports indicate that the most abundant group of sterols were 4-desmethylsterols, but 4αmonomethylsterols were also detected (Narumi et al. 2001). In most of studied species, among 4-desmethylsterols, both Δ5-sterols and Δ7-sterols coexist. The exception was B. indica where exclusively Δ5-sterols were found (Othman et al. 2021b; Javed et al. 2018a, b). Sitosterol and stigmasterol were the main sterol components in all species studied. GC–MS-identification of sterols revealed that fruits of B. scoparia were also abundant in 24-ethyllathosterol (Narumi et al. 2001). In turn, Javed et al. (2018a) underlined that γ-sitosterol was present in considerable amounts in the stem of B. indica. In addition to free sterols, their glycosides have also been reported in some of the Bassia species, including three novel kochiosides A,B,C, belonging to the class of 14α-methylated steroidal glucosides, which are rarely encountered in nature (Fig. 3B) (Imran et al. 2007). It is interesting that some structures containing the 14α-methyl group were found in sea cucumbers (Cordeiro and Djerassi 1990). To date, kochiosides have been isolated solely from B. prostrata (Imran et al. 2007; 2017). Phytoecdysteroids are polyhydroxylated plant sterols that occur in several species from the Amaranthaceae family (Dinan et al. 1998; Das et al. 2021). In the genus Bassia, they were found exclusively in B. scoparia (Dinan et al. 1994). The species was reported to contain 20-hydroxyecdysone and polypodine B (5β,20-dihydroxyecdysone) but the level of phytoecdysteroids varied considerably Phytochem Rev Table 3 Sterols detected in the genus Bassia Species Compound References B. prostrata Sterols: Imran et al. (2007, 2017), Seitimova et al. (2018) β-sitosterol (AP), stigmasterol (AP), spinasterol (AP) Steroidal glucosides: stigmasterol 3-O-β-D-glucopyranoside (AP), β-sitosterol 3-O-β-D-glucopyranoside (AP); kochioside A, kochioside B, kochioside C (WP) B. indica Sterols: Othman et al. (2021b), Javed et al. (2018a, b) β-sitosterol (AP, L, S), γ-sitosterol (S), stigmasterol (S) B. scoparia Sterols: stigmasterol (L,FR), β-sitosterol (L, FR), 24αmethylcholesterol (L), cholesterol (FR), lathosterol (FR), campesterol (FR), 24-methyllathosterol (FR), isofucosterol (FR), 24-ethyllathosterol (FR), avenasterol (FR); 31-norlanosterol (FR), lophenol (FR), obtusifoliol (FR), 4αmethylfecosterol (FR), gramisterol (FR), 4α-methyl-5αcholest-8-en-3β-ol (FR), citrostadienol (FR) Salt and Adler (1985), Narumi et al. (2001), Lu et al. (2012b), Zhang et al. (2013) Steroidal glucosides: daucosterol, (L,FR); Phytoecdysteroids: Dinan et al. (1994) 20-hydroxyecdysone (FR, R, L, S, FL) polypodine B (5β,20-dihydroxyecdysone) (FR, R, F, S, FL) unidentified ecdysteroid (WP) AP Aerial parts, FR Fruit, L Leaves, S Stem, R Root, FL Flos, WP Whole plant between the different morphotic parts of B. scoparia. The radioimmunoassay using DBL-1 antiserum revealed that the highest content of ecdysteroids was found in the roots (143 µg ecdysone equivalents/ g dw). Lower levels were observed in leaves, flowers, stems, and seeds (44–112, 56, 26–37 and 30 µg ecdysone equivalents/g dw, respectively) (Dinan et al. 1994). Phenolics Phenolics, including flavonoids and phenolic acids, are one of the most widely distributed metabolites in the plant kingdom. So far, phytochemical studies of Bassia species have led to the detection of nearly 60 flavonoids. Thirty-one of them were unambiguously identified following structure elucidation of isolated compounds (Lu et al. 2012b; Shaker et al. 2013; Musa et al. 2016; Kamel et al. 2001; Xu et al. 2014; Othman et al. 2021b). The identification of the remaining flavonoids was performed directly in Bassia plant extracts using HPLC with UV detection (Rakhmankulova et al. 2015; Xiao et al. 2018; Petruk et al. 2021) and by combination of chromatographic technique coupled with tandem mass spectrometry. Said et al. (2021) have recently evaluated the flavonoid profile in aerial parts of B. eriophora using LC–ESI–MS/MS in negative-ion mode, which led to tentative detection of 28 compounds (Said et al. 2021). In turn, Xiao et al. (2016) analyzed flavonoids in B. scoparia pollen using atmospheric solid analysis probe mass spectrometry (ASAP-MS) (Xiao et al. 2016). As shown in Table 4 various flavonoids, including flavonols, flavones, and a flavanone, were detected in the Bassia genus. It is interesting to note that free sugar isoflavones have also been found, but so far only in B. scoparia (Zhang et al. 2013). Most of the flavonoids identified in the Bassia genus are glycosidic (75%). Flavonol glycosides, including mainly derivatives of quercetin (35%), kaempferol (15%) and isorhamnetin (15%), are definitely the most abundant. It should be noted that acylated compounds account for 30% of flavonoid glycosides. Their presence was found exclusively in the aerial parts of Bassia species. Kamel et al. (2001) 123 Phytochem Rev Table 4 Flavonoids of the genus Bassia Compound Species (plant part) References Flavonols and dihydroﬂavonols: Quercetin B. scoparia (FR) Lu et al. (2012b) B. eriophora (AP) Xiao et al. (2018) Musa et al. (2016) 3′-methylquercetin B. muricata (AP) Shaker et al. (2013) Isorhamnetin B. scoparia (FR) Lu et al. (2012b) Morin B. prostrata (AP) Rakhmankulova et al. (2015) Fisetin B. prostrata (AP) Rakhmankulova et al. (2015) Dihydroquercitin B. prostrata (AP) Rakhmankulova et al. (2015) B. prostrata (AP) Petruk et al. (2021) Flavones: Luteolin B. eriophora (AP) Musa et al. (2021) 5, 7, 4′-trihydroxy-6,3’-dimethoxyflavone B. scoparia (FR) Lu et al. (2012b) 4′,5, 7-trihydroxy-6-methoxyflavone (Hispidulin) B. scoparia (FR) Lu et al. (2012b) Isoﬂavones Tectorigenin B. scoparia Zhang et al. (2013) Pratensein B. scoparia Zhang et al. (2013) Iriflogenin B. scoparia Zhang et al. (2013) 5,2′-dihydroxy-6,7-methylenedioxyisoflavone (Irisone B) B. scoparia Zhang et al. (2013) 5-hydroxy-6,7-methylenedioxyflavone (Cochliophilin A) B. scoparia Zhang et al. (2013) Quercetin glycosides: Quercetin-3-O-galactoside B. scoparia (FR) Xu et al. (2012) Quercetin 3-O-β-D-glucoside (Isoquercitrin) B. prostrata (AP) Petruk et al. (2021) Quercetin 7-O-β-D-glucopyranoside B. scoparia Xu et al. (2014) B. eriophora (AP) Musa et al. (2016) Quercetin-3,7-O-β-diglucopyranoside Quercetin-3-O-rutinoside (Rutin) B. muricata (AP) B. scoparia (FR) Kamel et al. (2001) Lu et al. (2012b) Wang et. al. (2018) Xiao et al. (2018) B. prostrata (AP) Quercetin-7-O-β-D-sophoroside Rakhmankulova et al. (2015) B. eriophora (AP) Musa et al. (2021) B. scoparia (FR) Xu et al. (2014) Quercetin-3-O-sophoroside B. muricata (AP) Kamel et al. (2001) Quercetin-3-O-(6″-caffeoyl)-sophoroside B. muricata (AP) Kamel et al. (2001) Quercetin-3-O-(6″-feruloyl)-sophoroside B. muricata (AP) Kamel et al. (2001) Said et al. (2021) Quercetin-3-O-(feruloyl)-O-sophoroside B. eriophora (AP) Quercetin-3-O-(sinapoyl)-sophoroside B. eriophora (AP) Said et al. (2021) Quercetin-3-O-β-D-apiofuranosyl-(1→2)-β-Dgalactopyranoside B.scoparia (FR) Xu et al. (2014) Quercetin 3-O-β-D-galactopyranosyl-7-O-β-Dglucopyranoside B. scoparia (FR) Xu et al. (2014) Quercetin 3-O-β-D-apiofuranosyl-(1→2)-β-Dgalactopyranosyl-7-O-β-D-glucopyranoside B. scoparia (FR) Xu et al. (2014) 123 Phytochem Rev Table 4 continued Compound Species (plant part) References Quercetin 3-O-α-L-rhamnopyranosyl-(1→6)-β-Dgalactopyranosyl-7-O-β-D-sophoroside B. scoparia (FR) Xu et al. (2014) 3′-methylquercetin 3-O-α-L-arabinopyranosyl(1→2)-L-α-arabinopyranoside B. muricata (AP) Shaker et al. (2013) Quercetin-3-O-sophoroside-7-O-hexoside B. eriophora (AP) Said et al. (2021) Quercetin-3-O-sophoroside-7-O-dihexoside B. eriophora (AP) Said et al. (2021) Quercetin-3,7-di-O-glycopyranoside B. eriophora (AP) Said et al. (2021) Quercetin-3-O-(feruloyl)-O-trihexosides B. eriophora (AP) Said et al. (2021) Other glycosides: Isorhamnetin-3-O-β-D-glucopyranoside Isorhamnetin-3-O-β-D-rutinoside Isorhamnetin-3-O-β-D-glucopyranosyl-(1→6)-O[α-L-rhamnopyranosyl-(1→2)]-β-Dglucopyranoside B. scoparia (FR) Lu et al. (2012b) B. eriophora (AP) Musa et al. (2016), Said et al. (2021) B. indica (AP) B. prostrata (AP) Othman et al. (2021b) Petruk et al. (2021) B. indica (AP) Othman et al. (2021b) Isorhamnetin-3-O-sophoroside B. eriophora (AP) Said et al. (2021) Isorhamntin-3-O-(feruloyl)-sophoroside B. eriophora (AP) Said et al. (2021) Isorhamnetin-3-O-(feruloyl)-O-hexuronide-Ohexoside B. eriophora (AP) Said et al. (2021) Isorhamntin-3-O-sophoroside-7-O-hexoside B. eriophora (AP) Said et al. (2021) Isorhamnetin-3-O-(feruloyl)-trihexoside B. eriophora (AP) Said et al. (2021) Isorhamnetin-3-O-(feruloyl)-O-tetrahexoside B. eriophora (AP) Said et al. (2021) Kaempferol-3-O-β-D-glucopyranoside (Populin) B. prostrata (AP) Petruk et al. (2021) Kaempferol-3-O-rutinoside B. indica (AP) Othman et al. (2021b) B. eriophora (AP) Musa et al. (2021) Said et al. (2021) Kaempferol-3-O-β-d-glucopyranosyl-(1→6)-O-[βD-galactopyranosyl-(1→3)-2-O-trans-feruloyl-αL-rhamnopyranosyl-(1→2)]-β-D-glucopyranoside B. indica (AP) Othman et al. (2021b) Kaempferol-3-O-dihexoside-7-O-dihexoside B. eriophora (AP) Said et al. (2021) Kaempferol-3-O-(caffeoyl)-O-trihexoside B. eriophora (AP) Said et al. (2021) Kaempferol 3-O-(feruloyl)-O-trihexoside B. eriophora (AP) Said et al. (2021) Kaempferol-3-O-(caffeoyl)-sophoroside B. eriophora (AP) Said et al. (2021) Kaempferol-7-O-hexoside-3-O-dihexosides B. eriophora (AP) Said et al. (2021) Kaempferide-3-O-trihexoside B. eriophora (AP) Said et al. (2021) Rakhmankulova et al. (2015) Naringenin 7-O-neohesperidoside (Naringin) B. prostrata (AP) Acacetin-7-O-β-D-glucoside B. eriophora (AP) Musa et al. (2021) Diosmetin 7-rutinoside B. eriophora (AP) Musa et al. (2021) Apigenin 6,8-di-C-hexosides B. eriophora (AP) Said et al. (2021) Apigenin 6-C-pentoside-8-C-hexoside or B. eriophora (AP) Said et al. (2021) Apigenin 8-C-pentoside-6-C-hexoside FR Fruit; AP Aerial parts 123 Phytochem Rev established the structure of quercetin sophorosides that contained the feruloyl and caffeoyl moiety that were isolated from B. muricata. Furthermore, Said et al. (2021) detected several acylated flavonol glycosides in B. eriophora using the LC–ESI–MS/ MS method. Recently, a novel flavonol tetraglycoside with feruloyl moiety (kaempferol-3-O-β-D-glucopyranosyl-(1→6)-O-[β-D-galactopyranosyl-(1→3)-2O-trans-feruloyl-α-L-rhamnopyranosyl-(1→2)]-β-Dglucopyranoside) (Fig. 2) were isolated from the aqueous methanol extract of the aerial parts of B. indica (Othman et al. 2021b). Although the research on flavonoids of the Bassia genus has been carried out intensively for several years, most quantitative studies focused on determining the total flavonoid content (TFC) or even the total phenolic content (TPC) (see Table S1, Supplementary information). Only a few reports on the quantitative determination of individual compounds have been published (Xiao et al. 2018; Rakhmankulova et al. 2015; Wang et al. 2018; Petruk et al. 2021). Wang et al (2018) measured the content of rutin (0.115 mg/g dw) in B. scoparia fruits by capillary electrophoresis. Rakhmankulova et al. (2015) revealed that naringin (5.03–13.53 mg/g dw) and rutin (1.71–8.13 mg/g dw) were dominant flavonoids in aerial parts of B. prostrata, while free-sugar flavonols, such as isorhamnetin, fisetin, and morin, were found in a lower content (0.11–0.75 mg/g dw, 0.01–0.76 mg/g dw, 0.01–0.03 dw, respectively) (Rakhmankulova et al. 2015). Petruk et al. (2021) also highlighted the high content of rutin (4.96– 8.13 mg/g dw) and the presence of luteolin (0.1– 3.0 mg/g), populin (0.1–6.4 mg/g), and isoquercitrin (0.1–3.4 mg/g) in the aerial parts of B. prostrata (Petruk et al. 2021). The authors suggest that cytotype and habitat can influence the content of flavonoids in plant species (Rakhmankulova et al. 2015; Petruk et al. 2021). Although phenolic acids appear to be widely distributed in plants, data on their occurrence in the genus Bassia are limited. So far, several phenolic acids from both hydroxybenzoic and hydroxycinnamic groups have been detected only in the aerial parts of the species. Chlorogenic acid, caffeic acid, and ferulic acid were determined in shoots of B. scoparia (Lodhi 1979). Seitimova et al. (2016) detected protocatechuic, vanillic, isovanillic, and pcoumaric acids in aerial parts of B. prostrata. The 123 latest report indicates that vanillic acid, o-hydroxybenzoic acid, p-hydroxybenzoic acid, caffeic acid, and methyl caffeate are present in B. indica (Othman et al. 2021b). Some phenolic acids may occur not as a free form, but as part of more complex structures. As mentioned above, flavonoid glycosides bearing feruloyl and caffeoyl moiety have been reported in B. muricata, B. indica and B. eriophora (Kamel et al. 2001; Said et al. 2021; Othman et al. 2021b). Recently, five lignanamides containing the feruloyl moiety were isolated from aerial parts of B. indica (Othman et al. 2021a). Other phenolics reported in the Bassia genus include 6,7-dihydroxycoumarin and tachioside, which were detected in aerial parts of B. indica (Othman et al. 2021b). Fatty acids Some Bassia species were analysed for the presence of oil and fatty acids. The percentage content of oil in the seeds of B. scoparia ranges from 9.7 to 16% (Hegnauer 1964; Weber et al. 2001; Abideen et al. 2015). Due to the fact that seeds are abundant in lipids, most of the studies focused on the identification and quantification of FA in various Bassia seed oils (Kleiman et al. 1972; Escudero et al. 1999; Weber et al. 2001). Data on the analysis of other morphotic parts are scarce and limited to the aerial parts of B. prostrata, B. indica, and B. muricata (Seitimova et al. 2016; Imran et al. 2017; Bibi et al. 2021; Abu Ziada et al. 2015). The reports indicate that Bassia species are the source of saturated and unsaturated fatty acids, especially FA with chain lengths of 16 to 18 carbons. GC–MS analysis revealed that Bassia seed oil contained 54% to 89% linoleic acid (Kleiman et al. 1972; Escudero et al. 1999; Weber et al. 2001; Abideen et al. 2015). Significant amounts of palmitic acid (9%–68%) and oleic acid (4%–17%) were also observed (Kleiman et al. 1972; Weber et al. 2001; Abideen et al. 2015). Similarly, oleic acid (64.9%) and linoleic acid (22.0%) are reported to be the main FA in aerial parts of B. prostrata (Seitimova et al. 2016). It is interesting that very long-chain fatty acids (VLCFA– FA with more than 22C), have been identified in B. scoparia seeds and aerial parts of B. prostrata (Table 6) (Imran et al. 2017; Weber et al. 2001; Abideen et al. 2015). Furthermore, reports revealed Phytochem Rev that members of the Bassia genus accumulate Δ5 unsaturated fatty acids, such as 16:1Δ5, 18:1Δ5, 18:2Δ5,9, 18:3Δ5,9,12, 20:4Δ5,8,11,14 (Kleiman et al. 1972; Escudero et al. 1999; Olagbemiro et al. 1999; Whitney et al. 2004). (Z)-5-hexadecenoic acid (16:1Δ5) (Fig. 3B) is an unusual monounsaturated FA that is considered a potentially useful substance in the control of the mosquito Culex quinquefasciatus, a vector of filariasis in humans. This is since 16:1Δ5 can serve as an intermediate in the production of an insect oviposition pheromone (Olagbemiro et al. 1999; Whitney et al. 2004). (Z)-5-hexadecenoic acid was reported to constitute 12% of the seed oil of B. prostrata and around 5% of the oils of the seeds of B. scoparia as well as B. hyssopifolia (Kleiman et al. 1972). Fatty acids detected in Bassia species are listed in Table 5. Nitrogen-containing compounds Phytochemical analyzes conducted in the 1960s suggested that some Bassia species contain alkaloids. Borkowski and Drost (1965) showed the presence of four compounds in the herb of B. scoparia and B. laniﬂora (formerly classified as B. arenaria) (Borkowski and Drost 1965). B. indica was also reported to contain three alkaloids, which accumulate in seeds (512 mg/100 g dw), pericarp (387 mg/100 g dw), shoot (up to 294 mg/100 g dw) and roots (85 mg/ 100 g dw) (Zahran et al. 1982). The research carried out at that time was mainly based on the reaction of the isolated substances with Dragendorff’s reagent and did not lead to the determination of the structure of alkaloids (Zahran and Wahid 1982). Only DrostKarbowska (1978) on the basis of spectroscopic techniques (UV, MS, NMR) showed that compounds found in aerial parts of B. scoparia are harman and harmine, belonging to the indole alkaloids with βcarboline skeleton (Drost-Karbowska et al. 1978). It is interesting that the most recent studies revealed the presence of a novel alkaloid named bassiamide A (N[(3-(3-methyl-1-oxo-butyl)amino)propyl]-3-(3,4-dihydroxyphenyl)prop-2-enamide in aerial parts of B. indica (Fig. 2) (Othman et al. 2021a). Other constituents reported in the Bassia genus include lignanamides. The presence of N-trans-feruloylmethoxytyramine and N-trans-feruloyltyramine were investigated in B. scoparia (Zhang et al. 2013) and B. indica (Othman et al. 2021a, 2022). N-trans- feruloyltyramine derivative was also tentatively detected by LC–ESI–MS/MS in B. eriophora (Said et al. 2021). Recently, several lignanamides including S-(-)-N-trans-feruloyl nor-metanephrine, S-(-)-Ntrans-feruloyl octopamine, and rarely occurring R-( +)-N-trans-feruloyl octopamine were isolated from aerial parts of B. indica (Fig. 3B) (Othman et al. 2021a, 2022). Species of Bassia genus have been reported to contain proteins and multiple amino acids, and several studies have considered their potential nutritional and feeding value (Davis 1979; Riasi et al. 2008; Waldron et al. 2006, 2020; Seitimova et al. 2016; Houlihan et al. 2019; Nair et al. 2021). For example, Waldron et al. (2006) showed that the crude protein content in B. prostrata ranges from 44.6 to 116.5 g/kg of aerial parts. A similarly high protein content was found in aerial parts of B. scoparia (117 g/kg) (Riasi et al. 2008). El-Adawy et al. (2020) reported a crude protein concentration in aerial parts of B. indica at 12.6%. Some reports underlined that protein content depended on the maturity stage of the plant (Knipfel et al. 1989; Waldron et al. 2006; Nair et al. 2021). Differences in protein level were also observed between various parts of the plant. Seeds, leaves, and stems of B. prostrata were reported to contain 190.6 g, 123.2 g and 55.1 g of protein per kg, respectively (Waldron et al. 2006). Waldron et al. (2020) also observed that protein levels decrease with increasing salinity in the soil (Waldron et al. 2020). Seitimova et al. (2016) revealed that the main amino acids in aerial parts of B. prostrata were glutamic acid (23.46%), alanine (5.82%), proline (3.35%), arginine (3.32), leucine (3.20%) and isoleucine (3.03%) (Seitimova et al. 2016). Houlihan et al. (2019) identified 18 amino acids in water-soluble exudates from B. scoparia seeds with tyrosine and glutamic acid as predominant compounds. It should be noted that some of the Bassia (Kochia) biotypes are resistant to sulfonylurea herbicides (ALS-inhibiting herbicides) (Kumar et al. 2019). Because the acetolactate synthase enzyme (ALS) is the key step in the biosynthesis of branched chain amino acids, some differences in the valine and leucine content were observed between biotypes resistant and susceptible to herbicides. The level of free valine and leucine was elevated in seeds and aerial parts of the resistant Bassia biotype compared to susceptible plants (Dyer et al. 1993; Chodová and Mikulka 2000). 123 Phytochem Rev Table 5 Fatty acids in Bassia sp Fatty acid Abb Species Trivial name B. scoparia B. prostrata B. indica FR/SE SE SE AP AP B. muricata B. hyssopifolia AP SE z 11:0 Undecylic + 12:0 Lauric acid +z 14:0 Myristic acid 15:0 +s,i Pentadecanoic acid 16:0 Palmitic acid 5 16:1Δ 9 (Z)-5-hexadecenoic acid ++ k,o, ++ k, k + 17:0 Margaric acid +w Stearic acid ++k,l,w,an 18:1Δ 9 + k ++ Palmitoleic acid 5 + ± k, ++ k 18:2Δ9,12 Linoleic acid + ++k,e,an + ++k 18:3 Δ α-linolenic acid + + k,e arachidic acid + 20:1Δ11 11-eicosenoic acid +w, 20:4Δ5,8,11,14 Arachidonic acid +e Bohenic acid + 22:1Δ13 13-docosenoic acid +w, Lignoceric acid 15 + k + +k +k +b +a b a +k ++s +* s + + +j +a + z +z s ++k +k + ++k +k + + z +k a an +a +z w, an 22:0 24:0 + k + ++ w,an 20:0 +z +k k ++ ±k 9,12,15 + s +s ++ +k 18:3Δ a k Oleic acid k +* +++ j,b +k +k + k,e s 18:1Δ 18:2Δ5,9 5,9,12 +z + k,w,e,an 16:1Δ 18:0 +z s an + w w, an +i 24:1Δ 15-tetracosenoic acid + 26:0 Cerotic acid +w, 28:0 Montanic acid +i 32:0 Lacceroic acid +i +z an FA Fatty acid;+ + +dominant FA;+ +main (relevant) FA;+present FA;±traces of FA;+* the location of the double bonds have not been determined; a Abeer et al. (2015); z Abu Ziada et al. (2015), an Abideen et al. (2015); b Bibi et al. (2021); e Escudero et al. (1999); i Imran et al. (2017); j Javed et al. (2018a); k Kleiman et al. (1972); l Lu et al. (2012b); o Olagbemiro et al. (1999); s Seitimova et al. (2016); w Weber et al. (2001) Furthermore, the concentration of these amino acids decreased in the susceptible Bassia population after herbicide treatment (Chodová and Mikulka 2000). It is well known that plants of the Amaranthaceae family can accumulate nitrates (Salehi et al. 2019; Liubertas et al.2020; Munekata et al. 2021). Several works have been devoted to quantification of nitrates in Bassia species, especially in B. scoparia (Finley and Sherrod 1971; Steppuhn et al. 1994; Escudero et al. 1999; López-Aguilar et al. 2013). The studies revealed that the content of nitrates in the aerial parts of B. scoparia decreases during the growing season 123 (Finley and Sherrod 1971; López-Aguilar et al. 2013). Steppuhn et al. (1994) also observed that the amount of nitrates can increase from 0.01 to 0.04 g/ kg to 0.5 g/kg after the addition of nitrogen fertilizer. Among other members of the Bassia genus, nitrates were detected also in B. hyssopifolia but only in traces (James et al. 1976). It is interesting that the accumulation of nitrates was not observed in B. prostrata, which is a perennial plant (Waldron et al. 2010; Acar et al. 2016). Phytochem Rev Other compounds Representatives of the genus Bassia can also accumulate organic acids. Among carboxylic acids, oxalic acid was the most common compound. Other compounds such as tartaric acid, malic acids, citric acid, succinic acid, lactic acid, acetic acid, formic acid, and L-ascorbic acid were also detected (Hong et al. 2006; Ma et al. 2011, 2016; López-Aguilar et al. 2013). A low glycolic acid content was found in the root of B. scoparia (Ma et al. 2011). Gluconic acid, citric acid, and oxalic acid were the major organic acids detected in the water-soluble exudates from the seeds of the species. Other acids identified in small amounts in seed exudates included: threonic acid, malic acid, malonic acid, glucaric acid, fumaric acid, lactic acid, ribonic acid, galacturonic acid, 2-ketogluconic acid, glucuronic acid and glucuronic acid 1,5-lactone (Houlihan et al. 2019). It is noteworthy that the level of carboxylic acids varies between the species, its morphotic parts, and maturity stage and depends on environmental factors. B. prostrata is considered a species with a low level of oxalic acid accumulation. The content of oxalates in aerial parts of the species averaged 1.2% (Davis 1979). In turn, studies on organic acid content in aerial parts of B. scoparia provide diverse data, which may result from differences in habitat conditions of the analyzed plants. B. scoparia is an alkali resistant forage plant and is able to survive in extremely unfavourable environments with a pH greater than 10. Several studies indicate that the accumulation of organic acids in the aboveground parts is probably the way to adapt to ecological conditions (Yang et al. 2007; Ma et al. 2011, 2016). The oxalate concentration in B. scoparia shoots (1.4–2.6% of dry weight) can be defined as moderate (Cohen et al. 1989; López-Aguilar et al. 2013). In other studies, the oxalic acid content was much higher and was estimated to be in the range of 12%–17.8% of the dry weight of aerial parts (Yang et al. 2007; Ma et al. 2011, 2016). Hong et al. (2006) revealed that the roots of B. scoparia accumulate higher amounts of oxalic acid (26.76 µmol/g) than the shoots (4.305 µmol/g). A similar relationship was found for other acids, such as malic acid (6.60 µmol/g roots; 1.93 µmol/g shoots), tartaric acid (15.04 µmol/g roots; 1.29 µmol/g shoots) and succinic acid (4.70 µmol/g roots; 2.03 µmol/g shoots) (Hong et al. 2006). It is interesting that, in contrast to control conditions, the concentrations of oxalic acid, succinic acid, malic acid, and tartaric acid were higher in shoots (71.99 µmol/g, 66.73 µmol/g, 45.99 µmol/g, 44.60 µmol/g, respectively) under alkaline stress than in roots of the plant (67.39 µmol/g, 64.0 µmol/g, 26.89 µmol/g, 40.99 µmol/g, respectively) (Hong et al. 2006). Contrary to research reported by Hong et al. (2006), Ma et al. (2011) observed that under control conditions mature leaves contain the highest amount of oxalic acid (8% dry weight), followed by young leaves, old stem, and roots (1% of dry weight), while under salt and alkali stress, the concentration of oxalic acids in mature leaves reached 10% and 12% of their weight, respectively (Ma et al. 2011). Although members of the Bassia genus are not considered aromatic plants, several studies indicate that some of them can produce essential oils (ElShamy et al. 2012; Kianinodeh et al. 2017; AbdElGawad et al. 2020). Recently, thirty-four compounds were identified by the GC–MS method in the essential oil of the aerial parts of B. muricata. The study revealed that the essential oil is rich in terpenes, including sesquiterpenes, oxygenated monoterpenes and diterpenes, which constitute 58.21%, 9.77%, and 1.19% of the oil, respectively. Aromatic compounds (21.92%) and hydrocarbons (8.91%) were also detected. Hexahydrofarnesyl acetone (47.34%) was the main compound of the essential oil of B. muricata, followed by 6-methoxy-1-acetonaphthone (19.92%), n-dotriacontane (3.58%) and endo-borneol (3.24%) and methyl-ionone (3.04%). The other volatile compounds constituted less than 3% of the essential oil (Abd-ElGawad et al. 2020). Thirty-one volatile components were also identified in essential oil from shoots of B. scoparia growing in Egypt. GC– MS analysis revealed that terpenes represented the dominant class of compounds (58.54%), but aliphatic and aromatic hydrocarbons were also present (29.18%). The main compounds were α-thujaplicin (22.91%), phytone (8.66%), dictamnol (7.98%), butylated hydroxytoluene (7.49%), phytol (6.57%) and camphenolone (3.84%) (El-Shamy et al. 2012). Kianinodeh et al. (2017) observed a similar qualitative composition of the essential oil of B. scoparia. However, the essential oil obtained from the fruits of the plant that grows in Iran differed in its quantitative composition because it contained alkanes (n- 123 123 Table 6 Anti-inflammatory activity of extracts and isolated substances of Bassia sp Species; extract Tested doses; Model or assay Administration B. scoparia 50, 200, 500 mg/kg; 70% ethanol extract from fruits Acetic acid- induced vascular permeability in ddY mice positive control: indomethacin (10 mg/kg; p.o.) p.o Carrageenan-induced paw edema in Male Wistar rats positive control: indomethacin (10 mg/kg; p.o.) Compound 40/80 induced edema in ddY mice positive control: diphenhydramine (50 mg/kg; po) Effect References (-) ↑ of vascular permeability induced by acetic Matsuda acid (effect at 200, 500 mg/kg comparable to et al. positive control) (1997a) Prevention ↑ of paw edema (at 200, 500 mg/ kg–significant effect) Prevention ↑ of paw edema (at 500 mg/kg– weak effect) Histamine/serotonin/bradykinin -induced edema in ddY mice Prevention ↑ of paw edema positive control: diphenhydramine (50 mg/kg; po) induced by histamine (significant effect) and serotonin or bradykinin (weak effect) Arachidonic acid-induced ear swelling in ddY mice Prevention ↑ of paw edema positive control: phenidone 20 mg/kg (i.v.) (dose-dependent, significant effect, at 500 mg/ kg comparable to positive control) 10—300 μg/ mL Histamine-induced contraction in isolated guinea pig ileum (ex vivo) (-) of contraction of isolated ileum with IC50 value of 220 μg/mL 10—300 μg/ mL Prekallikrein enzyme activity assay (in vitro) Not active in concentration 100—500 μg/mL B. scoparia ad libidum; aqueous infusion of seeds p.o Carrageenan-induced paw edema in an experimental model of Prevention ↑ of paw edema metabolic syndrome in male Wistar rats (short-lived effect) B. scoparia 150, 250 mg/ kg; p.o methanol extract, fractionated:→ chloroform extract, ethyl acetate extract, butanol extract, Carrageenan-induced paw edema Sprague–Dawley male rats Prevention ↑ of paw edema positive control: indomethacin (100 mg/kg; p.o.) Freud’s complete adjuvant-induced arthritis in Sprague– Dawley male rats Abtulov et al. (2020) Choi et al. (2002) Anti-edema effect (exception: chloroform fraction – not active) positive control: methotrexate (10 mg/kg; i.p.) B. scoparia 1% water extract from fructus Topical application 2,4-dinitrochlorobenzene (DNCB)-induced contact dermatitis in female BALB/mouse model; Histopathological analysis, Western blot analysis B. scoparia 30, 100 or 300 μg/ear Extract ↓ contact dermatitis (via inhibition of the production of inflammatory mediators) 1-fluoro-2,4-dinitrofluorobenzene (DNFB)-induced contact dermatitis mice in BALB/mouse model; Prevents histopathological changes; Histopathological analysis at 300 μg/ear effect similar to dexamethasone positive control: dexamethasone (75 μg/ear) Choi et al. (2014) ↓ ear swelling;↓ production of IL-6, IFN-γ; Jo et al. (2016) Phytochem Rev methanol extracts from fruit ↓ hyperplasia of dermis and epidermis vs. DNCB group; Species; extract Tested doses; Model or assay Administration Effect B. scoparia 100, 200 mg/ kg; (p.o.) ↓ levels of IL-4 and IL-5 in a dose-dependent Lee et al. manner (2011) ethanol extract from fruits B. scoparia methanol extract from fruits OVA induced mouse asthma model; biochemical assay—ELISA 7.5, 13, 30 μg/ LPS-induced raw 264.7 cells in vitro mL References (-) of NO production; ↓ PGE2 and TNF-α release (via blocking NF-kB activation) Shin et al. (2004) Chen et al. (2017) B. scoparia Serine proteases inhibitory assay in vitro; Cathepsin G (CG) (-): IC50 [300 μg/mL; water extract from fruits positive control: elaspol; CG inhibitor CG inhibitor: IC50 =0.29 μM proteinase 3 (-): IC50 [300 μg/mL elaspol: IC50 =0.63 μM Human neutrophil elastase (HNE) inhibition assay positive control: elaspol Selective inhibitory activity: IC50 =79.53 μg/ mL elaspol: IC50 =0.04 μM B. scoparia 70% ethanol extract from fruits 50, 200, 500 mg/kg; p.o Type I allergic model: - forty-eight-h homologus Passive Cutaneous Anaphylaxis (PCA) in Wistar rats at 200, 500 mg/kg: significant ↓ of dye leakage Matsuda caused by PCA et al. (1997b) positive control: DSCG (5 mg/kg, i.v.) one and a half-h heterologous PCA in ddY mice; prednisolone (20 mg/kg; p.o) Type II allergic model: Not active - reversed cutaneous anaphylaxis (RCA) in JW rabbits; positive control: dexamethasone (10 mg/kg) Type III allergic model: At 500 mg/kg: ↓ of paw swelling - direct passive arthrus reaction (DPAR) in JW rabbits positive control: prednisolone (25 mg/kg; p.o.) Type IV allergic model: - sheep red blood cell- induced delayed hypersensivity in ICR mice (SRBC-DTH model) positive control: prednisolone (10 mg/kg; p.o.) - picryl chloride-induced contact dermatitis in ICR mice (PCDT model) positive control: prednisolone (20 mg/kg; p.o.) At 500 mg/kg: ↓ of paw swelling (SRBC-DTH model) and ear swelling (PC-DT model) Phytochem Rev Table 6 continued 123 123 Table 6 continued Species; extract Tested doses; Model or assay Administration B. scoparia 200, 500 mg/ kg; 70% ethanol extract from fruits B. scoparia water extract Compound 48/80-induced pruritogenic model in male ddY mice Effect References (-) of scratching behaviour by 50% vs. control Kubo et al. (1997) group p.o positive control: diphenhydramine (50 mg/kg; p.o.) positive control: inhibition by 74—90% 0.15 g/mL, 0.3 g/mL, 0.6 g/mL; Itching guinea pig model caused by histamine; (-) of scratching behaviour Zou et al. (2021) Carrageenan-induced paw edema in Male Wistar rats Prevention ↑ of paw edema; control: diclophenac 10 mg/mL; i.p significant ↓ of the edema thickness Musa et al. (2016) Itching mice model topical application B. eriophora ethanol extract from aerial parts 250, 500, 750 mg/kg; (i.p.) (1.92±0.067, 1.69±0.112 and 1.58±0.096, respectively) compared to carrageenan injected group with value of 2.404±0.14 Fractions and compounds from Bassia species: Polysaccharide fraction isolated from water extract from Kochiae fructus (molecular weight[300 kDa) 125, 250 mg/ kg; (through gavage) LPS-induced ALI model in male ICR mice (post tests: measurement of MPO activity; neutrophil tissue elastase activity in tissue; protein and pro-inflammatory cytokines in BALF; quantification of NET-DNA) positive control: dexamethasone (10 mg/kg; i.p.) Protective activity against acute lung injury (ALI); Chen et al. (2017) ↓ levels of TNF-α and IL-6 ↓ MPO activity ↓ neutrophil infiltration ↓ HNE activity: IC50 =3. 74 μg/mL ↓ NET-DNA formation Serine proteases inhibitory assay in vitro; Cathepsin G (CG) (-): IC50 [300 μg/mL; positive control: elaspol; CG inhibitor CG inhibitor: IC50 =0.29 μM proteinase 3 (-): IC50 [300 μg/mL elaspol: IC50 =0.63 μM Human neutrophil elastase (HNE) inhibition assay positive control: elaspol Selective inhibitory activity: IC50 =79.53 μg/ mL 100, 200 mg/ kg 2,4-dinitrochlorobenzene- (DNCB)-induced ACD Sprague– Dawley rats; ↓ ear swelling (no significant differences vs. positive control group; significant effect) Topical application (2x/24 h) Histopathological analysis; biochemical assay—ELISA; Western blot; paraffin section immunohistochemical method positive control: prednisolone acetate (2.5 mg/kg) ↓ levels TNF-α and IL-6, IL-18, INF-γ elaspol: IC50 =0.04 μM Flavonoid fraction isolated from ↑ levels of IL-10 ↓ levels of pERK1/2, TLR4, NFκB Xiao et al. (2018) Phytochem Rev 80% ethanol extract of Kochiae fructus Species; extract Tested doses; Model or assay Administration Effect Momordin Ic 6.25, 12.5, 25 μg/mL LPS-induced raw 264.7 cells in vitro Momordin Ic: ↓ production of TNF-α, IL-6, Yoo et al. (2017) PGE2; OA, 20-hydroxyecdysone -not active Momordin Ic 5, 10 mg/kg; oleanolic acid (OA) i.p Freund’s adjuvant-induced arthritis in Sprague–Dawley male Anti-edema effect rats oleanolic acid (OA) 20-hydroxyecdysone References Choi et al. (2002) positive control: aminopyrine (100 mg/kg; p.o.) Momordin Ic 20, 50, 100 mg/kg; p.o Type I allergic model: Significant ↓ of dye leakage caused by PCA - forty-eight-h homologus Passive Cutaneous Anaphylaxis (PCA) in Wistar rats Matsuda et al. (1997b) positive control: DSCG (5 mg/kg, i.v.) - one and a half-h heterologous PCA in ddY mice; prednisolone (20 mg/kg; p.o.) Significant ↓ of dye leakage caused by PCA (at 100 mg/kg effect comparable to positive control) Type IV allergic model: At 50, 100 mg/kg: ↓ of ear swelling - picryl chloride-induced contact dermatitis in ICR mice (PCDT model) Momordin 20,50 100 mg/kg; p.o positive control: prednisolone (20 mg/kg; p.o.) Carrageenan-induced paw edema in Male Wistar rats positive control: indomethacin (10 mg/kg; p.o.) 20, 50,100 mg/ Compound 48/80-induced pruritogenic model in male ddY mice kg; p.o Prevention ↑ of paw edema (significant effect); Matsuda et al. at 50 and 100 mg/kg, after 1 h, the effect (1997a) comparable to positive control At 50 mg/kg (-) of scratching behaviour by 50% vs. control group Kubo et al. (1997) positive control: diphenhydramine (50 mg/kg; p.o.) (–) inhibition; ↑ the increase; ↓ reduction/descrease; ALI Acute lung injury; CG Cathepsin; DNCB 2,4-dinitrochlorobenzene; DNFB 1-fluoro-2,4,-dinitrofluorobenzene; DPAR Direct passive arthus reaction; DSCG sodium cromoglicate; ELISA Enzyme-linked immunosorbent assay; ERK Extracellular-signal-regulated kinase; HNE Human neutrophil elastase; i.v. Intravenous; IL-10 Interleukin-10; IL-4 Interleukin-4; IL-5 Interleukin-5; IL-6 Interleukin-6; INF-γ Interferon-gamma; MPO Myeloperoxidase; NET-DNA Net-like structures of nuclear DNA; NF-κB Nuclear factor kappa B; p.o. Per os; PCA Passive cutaneous anaphylaxis; PCA Passive cutaneous anaphylaxis; PGE2 Prostaglandin; RCA Reversed cutaneous anaphylaxis; TLR Toll-like receptor; TNF-α Tumor necrosis factor α Phytochem Rev Table 6 continued 123 Phytochem Rev tetracosane, n-tricosane, n-docosane, n-henicosane, n-eicosane) as the main constituents (Kianinodeh et al. 2017). Hydrocarbons such as heptacosane, pentacosane, hexacosane, tetracosane, tricosane, and 9-tricosene were also detected in aerial parts of B. indica. In addition, phytol, verbenone, heptanone, p-methoxy acetophenone, phenyl benzene, 2-pentanoyl thiophene and β-ionone were identified as the main volatile compounds (Abou Zeid and El-Khayat 2000). Recently, the polysaccharide rich fraction (KSWP), consisting of carbohydrates (81.82%), uronic acid (6.63%), and protein (8.71%), was isolated from the water extract of the B. scoparia fruit. HPLC analysis revealed that among the monosaccharides in KSWP mannose, rhamnose, glucuronic acid, glucose, galactose, and arabinose were present. Glucose was found to be the predominant sugar in the polysaccharide fraction (Chen et al. 2017). In turn, Houlihan et al. (2019) detected free monosaccharides in water-soluble exudates from seeds of B. scoparia. Among them, fructose, galactose, glucose, and sorbitol were the main compounds. Other sugars, identified in a low amount, included: trehalose, scyllo-inositol, sucrose, mannitol, mannose, rhamnose, melibiose, maltitol, ribitol, xylulose, erythritol, and glucosamine (Houlihan et al. 2019). Other constituents reported in the Bassia genus include macroelements (Na, K, Mg, Ca, P) and microelements (Fe, Cu, Se). These plant species are able to accumulate ions, but the amount depends on various factors, such as soil composition or environmental contamination (Karimi et al. 2005; Yang et al. 2007; Riasi et al. 2008; Ma et al. 2011; Endo et al. 2014; Yamada et al. 2016). Environmental factors, such as salinity and cadmium stress, also affected the content of carotenoids in the Bassia genus. Seasonal variations in the level of carotenoids were also observed (Davis 1979). The published reports indicate that the carotenoids content ranged from 0.13 to1.06 mg/g of fresh aerial parts of the analysed species: B. scoparia, B. prostrata, and B. indica (Karimi et al. 2005; Abeer et al. 2015; Hashem et al. 2019; Ghaffarian et al. 2020). In addition to the phytoconstituents mentioned above, syringaresinol, a compound of the lignan group, was recently isolated from aerial parts of B. indica (Othman et al. 2021a). 123 Biological activity The biological activity studies conducted on the representatives of the genus Bassia focused on several plant species, including B. scoparia, B. eriophora, B. muricata, B. indica, and B. prostrata. It is interesting that most of the studies on B. scoparia were carried out on fruits or seeds of the plant, while the investigations of the other species were mainly focused on the other aerial parts. The various activities of extracts of Bassia species were evaluated, including anti-inflammatory, cytotoxic, hypoglycemic, antioxidant, antimicrobic and tissueprotective potential. However, it should be mentioned that published data on anti-inflammatory hypoglycemic and anti-obesity activity were limited almost completely to Kochiae fructus. It should be noted that not only in vitro tests, but also in vivo studies were conducted. Additionally, in several investigations, an attempt was made to determine which components are responsible for the observed activity of the extract. Anti-inflammatory activity The dried fruit of Bassia scoparia is commonly used in traditional Chinese medicine (TCM) as an antiinflammatory agent, especially in skin disorders (Chien et al. 2013; Chen et al. 2015, 2016; Weng et al. 2016). The anti-inflammatory activity of Kochiae fructus has been well documented in both in vitro studies and in various animal models (see Table 6). In most of the studies, acute inflammation models were performed, including those of edema. Due to the fact that skin inflammation often manifests itself by itching, antipruritic activity was also investigated. The water and ethanol extracts of Kochiae fructus were found to demonstrate the ability to inhibit scratching behavior in animal models (see Table 6). It should be noted that the extracts demonstrated their anti-inflammatory activity in vivo after oral administration and external application. The effect of Kochiae fructus on various types of hypersensitivity was also investigated in animal models (see Table 6). It was found that 70% extract exhibited inhibitory activity in models of I, III and IV-type of allergy (Matsuda et al. 1997b). The activity of extracts in contact allergic dermatitis, Phytochem Rev Fig. 5 The main pathways involved in the anti-inflammatory activity of extracts and compounds from Kochiae fructus. Abbreviations: ERK extracellular signal-regulated kinase; HNE human neutrophil elastase; ICAM-1 intercellular adhesion molecule-1; IL-1β interleukin-1β; IL-6 interleukin-6; IL10 interleukin-10; INF-γ interferon-gamma; JNK c-Jun N-terminal kinases; MAPK mitogen-activated protein kinases; MCP-1 monocyte chemotactic protein-1;MMP-9 matrix metalloproteinase-9; NFκB nuclear factor kappa B;PGE2 prostaglandin E; PPAR peroxisome proliferator-activated receptors; ROS reactive oxygen species; TNF-α tumor necrosis factor α; VCAM-1 vascular cell adhesion molecule-1 which can be classified as type IV hypersensitivity, has been one of the most intensively studied. External application of methanol or water extracts significantly inhibited inflammation in animal contact dermatitis models (Choi et al. 2014; Jo et al. 2016). Choi et al. (2014), revealed that 1% water extract significantly reduced hyperplasia and thickening of the dermis and epidermis (by 35.6% and 39.2%, respectively) of mice in DNCB-induced dermatitis model compared to the animal group without treatment. Furthermore, histological examination by Jo et al. (2016), demonstrated that repeated topical application of methanol extract inhibited epidermal acanthosis, spongiosis and immune cell infiltration (Jo et al. 2016) in mice with DNFB-induced contact dermatitis model. The observed anti-inflammatory effect as inhibition of ear swelling was comparable to dexamethasone (Jo et al. 2016). Matsuda (1997b) observed that, in addition to antiallergic activity in type IV hypersensitivity, Kochiae fructus also influenced direct passive arthus reaction (DPAR) in rats, which is a type III allergic model. In another study, Choi et al (2002) demonstrated the potential of methanol extract in Freund’s complete adjuvant-induced arthritis in Sprague–Dawley rats (Choi et al. 2002). In addition, B. scoparia extracts also showed antiallergic potential in IgE-mediated hypersensitivity (type I). This type of allergy is associated with eczema and asthma (Matsuda et al. 1997b; Lee et al. 2011). Ethanol extracts from fruits reduced the level of interleukins (IL-4, IL-5) and the migration of inflammatory cells into the lungs in the OVA-induced asthma model in BALB/c mice. The effect may be the result of inhibition of matrix metalloproteinase-9 (MMP-9) inhibition and reduction in the expression of adhesive molecules (VCAM-1 and ICAM-1) in lung tissue (Lee et al. 2011). The published research postulates that Kochiae fructus extracts exerted anti-inflammatory effects by influencing various factors involved in the inflammatory response. Kochiae fructus methanol extract 123 Phytochem Rev decreased the level of INF-γ, TNFα, IL-1β and MCP1 in the mouse model with DNCB-induced contact dermatitis (Jo et al. 2016). The methanolic extract was also observed to be a potent inhibitor of LPSinduced TNF-α, NO and PGE2 production in LPSinduced RAW cells 264.7 in vitro. The effect was associated with LPS-induced iNOS and COX-2 gene expression by blocking nuclear factor kappa beta (NFκB) activation (Shin et al. 2004). Furthermore, Choi et al. (2014) point out that the anti-inflammatory activity of B. scoparia can be regulated via NFκB and MAP kinase pathways, such as ERK1/2, p38 and JNK (Choi et al. 2014). Furthermore, recently, Kochiae fructus was found to exert peroxisome proliferator activated receptors (PPAR) α/γ dual agonistic activity, which can alleviate the inflammation process (Jeon et al. 2016). Furthermore, the water extract showed the ability to inhibit human neutrophil elastase (HNE) (IC50 =79.53 μg/mL), the hydrolytic enzyme that can cause damage to elastase-rich tissue, and also promote inflammatory cell migration (Chen et al. 2017). The search for active compounds revealed that various phytochemicals of Kochiae fructus could be associated with an anti-inflammatory effect. Several studies indicated that activity is related to saponins, mainly momordin Ic (Fig. 3B) (Matsuda et al. 1997a, b). This monodesmosidic saponin suppressed the production of pro-inflammatory cytokines, including TNF-α, IL-6 as well as PGE2 in RAW 264.7 cells stimulated with LPS (Yoo et al. 2017). Furthermore, the anti-inflammatory, antiarthritic, and antiallergic potential of the compound was demonstrated in several animal models as well (see Table 6) (Kubo et al. 1997; Matsuda et al. 1997a, b). Furthermore, the antipruritic effect of Kochiae fructus was also attributed to saponins. However, it should be noted that the structure of a particular saponin affected the observed activity. Monodesmosides such as momordin Ic and oleanolic acid 3-O-glucuronide were observed to alleviate scratching behavior, while bidesmosides such as momordin IIc (Fig. 2) were almost ineffective against itching induced using compound 48/80 (Kubo et al. 1997; Matsuda et al. 1998a). However, Xiao et al. (2018) underlined that anti-inflammatory activity is related to the flavonoid fraction which was demonstrated using an animal model of DNCB-induced contact dermatitis. The flavonoid fraction, isolated from Kochiae fructus, 123 containing rutin (24 mg/g) and quercetin (18 mg/g), after topical application, was found to reduce the level of pro-inflammatory cytokines and simultaneously increase the level of anti-inflammatory interleukin IL-10. Additionally, a significant decrease in tissue monocyte infiltration was observed. These effects could be the result of NFκB suppression via the ERK/TLR4 pathway. It should be mentioned that the flavonoid fraction exerted an effect comparable to a positive control (prednisolone acetate; 2.5 mg/kg) (Xiao et al. 2018). In turn, Chen et al. (2017) with the use of bioactivity-guided fractionation revealed that the effect of the water extract of Kochiae fructus should be attributed to the polysaccharide fraction. It significantly reduced the activity of human neutrophil elastase with an IC50 value of 3.74 μg/mL. Furthermore, the polysaccharide rich fraction was also active in the LPS–induced acute lung injury model in mice (Chen et al. 2017). Summarizing the current state of art, the postulated main pathways involved in the antiinflammatory activity of the extracts and compounds from Kochiae fructus are presented in Fig. 5. To our knowledge, in addition to anti-inflammatory investigations on B. scoparia, there is only one report devoted to another species of Bassia. Musa et al. 2016, demonstrated that the ethanol extract of aerial parts of B. eriophora (250, 500 and 750 mg/kg; i.p.) possessed the ability to inhibit swelling in the carrageenan-induced model of paw edema in rats (Musa et al. 2016). Antinociceptive activity The antinociceptive potential of some Bassia species was investigated using an acetic acid-induced writing test and a hot-plate test. The 90% ethanol extract of aerial parts from B. eriophora (250 and 500 mg/kg, p. o.) was found to cause an analgesic effect in both tests. The animals were treated with the extract after 90 min. and showed an increase in reaction time, which was comparable to indomethacin (4 mg/kg) (8.1±0.18; 8.1±0.12; 9.18±0.22, respectively) [control: 5.82±0.13]. In the writhing method, extracts (250 and 500 mg/kg, p.o) also demonstrated significant antinociceptive activity (55.14% and 68.38%) when compared to the control vehicle group, however the effect was slightly worse than Phytochem Rev observed for indomethacin (86%) (Yusufoglu 2015a). Musa et al. (2016) also confirmed the antinociceptive activity of the ethanol extract of the aerial parts of B. eriophora using the hot plate test, but the significant effect was observed only at high doses (500 mg/kg and 750 mg/kg, p.o) (Musa et al. 2016). In turn, Matsuda et al. (1997a, b) revealed that 70% extract of Kochiae fructus (500 mg/kg) demonstrated activity only in the acetic acid-induced writhing model (Matsuda et al. 1997a). Different results were provided by Choi et al. (2002), who investigated the activity of the ethanolic extract of Kochie fructus and fractions (chloroform, ethyl acetate, butanol) derived from the extract. All extracts (100 and 200 mg/kg; p.o.) revealed an analgesic effect in both hot plate and writhing tests in male Sprague–Dawley rats (Choi et al. 2002). Furthermore, the triterpenes isolated from the extract, such as momordin Ic and oleanolic acid (5, 10 mg/kg, i.p.) were also active in both models (Choi et al. 2002). Further studies revealed that momordin Ic (20,50,100 mg/kg, p.o.) alleviated the pain response but only in the second (late) phase of the formalin test, suggesting that its antinociceptive activity is mainly associated with anti-inflammatory activity (Matsuda et al. 1997a). On the other hand, Bassia extracts also showed activity in the hot plate test, which is considered to integrate supraspinal pathways (Deuis et al. 2017). This suggests that apart from anti-inflammatory activity, other mechanisms may also be involved in antinociceptive effect, but this issue requires in-depth research. Cytotoxic activity Most published studies on the cytotoxic activity of the Bassia genus concern B. scoparia species, while two articles describe the activity of B. indica (AbdelHamid et al. 2017; Aboul-Enein et al. 2012) and one B. muricata (Al-Barri et al. 2021). Interestingly, only two of the reviewed papers defined, at least partially, the chemical content of the extracts studied (Cho et al. 2019; Wang et al. 2014a) and one paper provided a full chemical characterization of the essential oil of B. scoparia seeds which was the object of the study (Kianinodeh et al. 2017). Most of the results indicate a rather weak cytotoxic potential of Bassia extracts. The weak cytotoxic activity of the water and ethanol extract prepared from B. indica whole plant against Ehrlich ascites carcinoma cells was observed, with a decrease of 2.88 and 1.6% decrease in cell viability (Abdoul-Enein et al. 2012). A similar weak activity of the ethanolic extract of the whole plant of B. indica was described for HepG2 cells, with IC50 120.5 μg/mL, while for the control cytostatic 5-FU, the IC50 value was 237.56 μg/mL. The extract was combined with 5-FU, and the most effective ratio, namely 5-FU/Bassia 1:2, decreased cell viability up to 48%, with a combination index of 0.996, suggesting a synergistic effect (Abdel-Hamid et al. 2017). In another study, a similar weak effect of Kochiae fructus aqueous and 50% ethanol extracts was investigated on the same HepG2 cells, after 4 and 24 h. Despite the difference in momordin Ic content, both extracts revealed a comparable, weak cytotoxic activity. Extracts at the dose of 1 mg/mL for 4 h or 0.1 mg/mL for 24 h decreased cell viability to approximately 60–70%, while almost total inhibition of cell viability was observed at doses as high as 5 mg/mL (Wang et al. 2014a). The methanolic extract of B. scoparia seeds revealed only weak cytotoxic activity in human lung A549 and colon Col2 cancer cells, with an IC50 exceeding 20 µg/mL (Nam and Lee 2000). Other studies present the cytotoxic impact of ethanolic extract of B. scoparia seeds on human neuroblastoma N-2A cells, with LC50 0.147 mg/mL. Although the authors classified the effect as strong, the value seems to indicate rather weak activity (Mazzio et al. 2009). B. scoparia seed ethanolic (80%) extract was evaluated for the possibility of its use in psoriasis. The in vitro experiment was carried out on HaCaT cells and proliferation was measured by SRB and MTT assays. The extract revealed a rather weak effect in decreasing keratinocyte proliferation, with IC50 185.7 and 125.0 µg/mL for the SRB and MTT assay, respectively (Tse et al. 2006). B. muricata whole plant was extracted with different solvents, namely methanol, ethanol, or the combination of methanol:chloroform:water (12:5:1), and the cytotoxicity of the extracts was examined against human lymphocytes and Chinese hamster ovary CHO cells. Not only was none of the extracts toxic to the tested cells, but an increase in cell viability was also observed during exposure at concentrations of up to 2 mg/mL (Al-Barri et al. 2021). 123 Phytochem Rev Several reports provide data on the mechanism of action of Bassia extracts. The cytotoxic potential of the methanolic extract of B. scoparia seeds was evaluated in a panel of prostate cancer cells, differing in metastatic potential: LNCaP, PC-3, RC-58 T, but also normal prostate epithelial cells RWPE-1, with IC50 89.25, 123.41,141.62 and[250 µg/mL, respectively. The subtoxic dose of the extract (20 µg/mL) significantly suppressed VEGF-induced migration, invasion and formation of capillary-like structures of human umbilical vein endothelial cells (HUVECs) and micro vessels that sprout from rat aortic rings. The extract also downregulated the phosphorylation of VEGFR2 and the level of PI3K/AKT/mTOR, which resulted in decreased angiogenesis in HUVECs (Cho et al. 2019). Similar results on the antiangiogenic effect of Kochiae fructus extract were published by Na et al. (2006). The extract decreased the expression of the antiangiogenic protein HIF-1α (hypoxia-inducible factor 1-α) in HepG2 and HaCaT cells, and also reduced the level of VEGF and iNOS in HaCaT cells. The results indicate that the extract may be an antiangiogenic and anti-inflammatory agent useful in decreasing tumor progression (Na et al. 2006). The methanolic extract of B. scoparia mature fruits was found to influence proliferation rate, cell cycle arrest, the generation of reactive oxygen species (ROS) and the stimulation of apoptosis in human breast MDA-MB-231 cancer cells. The results showed a dose-dependent decrease in cell proliferation, with IC50 36.2 μg/mL. Furthermore, the extract at 25 μg/mL significantly increased the sub-G1 DNA content in cells to 44.7%, when compared to untreated cells, and also increased pro-apoptotic proteins such as cleaved caspase 3, cleaved caspase 8, cleaved caspase 9 and cleaved Poly (ADP-ribose) polymerase (PARP), leading to cell apoptosis (Han et al 2014). In their further studies, the authors examined the effect of Bassia scoparia mature fruit methanolic extract on human (Ca9-22 and HSC-4) and murine (AT-84) oral squamous cell carcinoma, and also on human HaCaT keratinocytes. The extract arrested cancer cells in the sub-G1 phase, and also led the cells to apoptosis, by activating caspase-3 and -9 through the p38 MAPK pathway. Importantly, normal HaCaT cells were not affected (Han et al. 2016). To our knowledge, there is only one report devoted to evaluating the cytotoxic potential and mechanism 123 of action of essential oil from Bassia plants. The essential oil from mature fruits of B. scoparia, obtained by the Clevenger apparatus, which contained alkanes as main constituents (n-tetracosane, ntricosane, n-docosane, n-henicosane, n-eicosane), was tested in human breast cancer MCF-7 and normal HU02 fibroblast cells after 24, 48 and 72 h of incubation. The substance tested inhibited the viability of MCF-7 cells, with IC50 125 μg/mL (72 h), while normal cells were not affected. Furthermore, incubation of MCF-7 cells with essential oil resulted in changes in cellular morphology, manifested as star-shaped cells, vacuolation, cytoplasmic and cellular shrinkage. Pyknotic nuclei were also observed, suggesting stimulation of apoptosis in cells (Kianinodeh et al. 2017). Despite the large amount of in vitro data, so far only one animal study on Bassia extracts has been carried out. The anticancer properties of the methanolic extract of mature B. scoparia fruits were described in vivo, in C3H mice implanted with 3105AT-84 oral squamous cell carcinoma. The animals in the experimental groups received low (1 mg/kg body weight), medium (3 mg/kg body weight) or high (5 mg/kg body weight) doses of the extract, while the control animals were treated with a vehicle of 5% ethanol. The extract was introduced by intratumor injection once every two days (four times in total). The extract significantly reduced tumor volume in a dose-dependent manner, with the highest dose being the most effective. The authors also noted that the extract did not induce systemic toxicity and that no loss of body weight was also observed during treatment (Han et al. 2016). Numerous reports indicate that momordin Ic, (Fig. 3B) a metabolite of saponin group, may be one of the active compounds of Kochiae fructus extracts as it has well-documented cytotoxic activity. Momordin Ic significantly reduced HaCaT cell viability, with IC50 of 168.70 and 76.40 μM/L for 24 and 48 h, respectively. The compound affected cell morphology, resulting in cell shrinkage, but also stimulated cell apoptosis and arrested the cells in S-phase. Furthermore, momordin Ic treatment significantly decreased β-catenin, c-Myc, and VEGF mRNA expression, compared to untreated cells, and modified Wnt/β-catenin pathway activation by affecting β-catenin nuclear distribution. The authors suggest that such activity of momordin Ic may be Phytochem Rev of interest in antipsoriasis therapy in the future (Luo et al. 2021). In another study, the compound promoted the formation of autophagic vacuole and increased the expression of Beclin 1 and LC-3 in a dose- and time-dependent manner in HepG2 cells. The study results also revealed the crosstalk between autophagy and apoptosis stimulation by the compound, which simultaneously induced both processes by suppressing ROS-mediated PI3K/Akt and activating the ROS-related JNK and P38 pathways (Mi et al. 2016). Another study in the same HepG2 cellular model indicated that the compound induced apoptosis in cells, manifested as DNA fragmentation, caspase-3 activation, and PARP cleavage. In addition, momordin Ic stimulated reactive oxygen species (ROS) production and decreased mitochondrial membrane potential, cytochrome c release, down-regulation of Bcl-2 and up-regulation of Bax expression. Activation of p38 and JNK, inactivation of Erk1/2 and Akt, and alterations in the expression of iNOS and HO-1 were also observed after treatment with the compound. These results indicated that momordin Ic induced apoptosis through mitochondrial dysfunction regulated by oxidative stress involving the MAPK and PI3K-mediated iNOS and HO-1 pathways (Wang et al. 2013). In their further studies, the authors investigated the detailed mechanisms of momordin Ic apoptosis stimulation in HepG2 cells. The results demonstrated that momordin Ic activated PPARɣ and inhibited COX-2. PGC-1α and FoxO4 expressions were increased by the PI3K or MAPK pathways, while inhibition of PPARɣ decreased the expression of p-p38 and FoxO4, and restored COX-2 expression. ROS inhibition affected mainly PGC-1α expression, while PPARɣ, COX-2 and FoxO4 expression was almost unaffected (Wang et al. 2014b). In mouse fibroblast NIH 3T3 cells, momordin I suppressed the activator protein-1 (AP-1), an important protein in cellular signaling, responsible for the induction of several genes in response to physiological signals. Furthermore, momordin Ic also decreased the de novo synthesis of the AP-1 protein. The authors also suggest that the inhibitory site of momordin Ic might be in the Jun/Fos dimer and not in DNA, and the basic region of c-Jun is the most probable inhibitory action site (Park et al. 2000). Momordin Ic inhibited the proliferation of KB oral carcinoma cells with IC50 10.4 µg/mL. Furthermore, the compound caused chromosomal DNA fragmentation and at a concentration of 20 µg/mL stimulated apoptosis in cells (approximately 20% apoptotic cells after 72 h) (Seo et al. 2007). The small ubiquitin-like modifier (SUMO) protease, SENP1, plays an important role in cellular inflammation by regulating proteins in SUMOylation. Some studies suggest that momordin Ic might be a novel SENP1 inhibitor. In one of the studies momordin Ic reduced LPS-induced cellular inflammation in the RAW 264.7 macrophages model by depressing SENP1 expression. Furthermore, the effect of SENP1 on the LPS-induced inflammatory response was dependent on the interaction of SENP1Sp3 and the promotion of Sp3 (transcription factor controlling the expression of genes involved in the cell cycle and inflammation response) expression by deSUMOylation of Sp3. Furthermore, momordin Icdepressed Sp3 expression altered the Sp3-nuclear factor (NF)-κB interaction, which decreased cellular inflammation (Zheng et al. 2020). SUMO-specific protease 1 (SENP1), a member of the de-SUMOylation protease family, is elevated in prostate cancer cells and is involved in cancer pathogenesis. Momordin Ιc inhibited SENP1 in vitro and altered the thermal stability of SENP1. Furthermore, the compound increased SUMOylated protein levels, namely hypoxia inducible factor-1α and nucleus accumbens associated protein 1 in PC3 prostate cancer cells, and also reduced SENP1 mRNA levels in cells. The authors suggest that momordin Ic may have therapeutic potential in prostate cancer (Wu et al. 2016). Another study focused on the effect of momordin Ic on a panel of colon cancer cells. The compound decreased the viability of colon cancer CT-116, HCT-8, SW480 and HT-29 cells, with IC50 values ranging from 6.40 to 12.79 µM. As HCT-116 and HCT-8 cells were most vulnerable (IC50 6.40 and 6.83 µM, respectively) they were used in subsequent experiments. Momordin Ic stimulated apoptosis in cells, and arrested cells in the G0/1 phase of the cell cycle. Furthermore, the compound enhanced the SUMOylation of c-Myc, which led to the downregulation of the c-Myc protein. The authors conclude that momordin Ic revealed its cytotoxic effect through the SENP1/c-Myc signaling pathway (Xianjun et al. 2021). In addition to in vitro studies, the antitumor activity of momordin Ic was investigated in an animal model. Momordin Ic was administered to 123 Phytochem Rev Balb/c nude mice implanted with PC3 prostate cancer cells, at a dose of 10 mg/kg daily for 20 days. On day 20, the xenograft tumors treated with the compound were less than those of the control group (p\0.05). The authors observed the accumulation of SUMO1and SUMO2/3-modified proteins in PC3 tumor xenografts treated with momordin Ic. However, a slight decrease in body weight was observed in the momordin Ic-treated group compared to the control group (Wu et al. 2016). Although most of the research on cytotoxic activity has focused on momordin Ic, a single report on other compounds isolated from Bassia species has also been published recently. Three new steroidal glycosides, kochioside 1A1, kochioside 2A1 and kochioside 3A1, isolated from Kochia prostrata, were examined in a brine shrimp lethality assay, and their potential was comparable to the standard drug etoposide, with LD50 8.3201, 8.8205, 8.2310, and 7.4625 μg/mL, respectively (Irfan et al. 2020). Hypoglycemic activity and anti-obesity activity The ethanol extract of Kochiae fructus demonstrated an anti-obesity effect in diet-induced obese mice. Oral administration of extracts together with the high-fat diet (1%, 3%) prevented weight gain. After 9 weeks, a reduction in fat storage was also observed (Han et al. 2006). A similar anti-obesity effect was found for the infusion of B. scoparia seeds prepared from 1.5 g of seeds/100 mL; 3 g of seeds/100 mL; 6 g of seeds/100 mL. Consumption of a high-fat diet with infusions (ad libitum) for 12 weeks by Wistar rats with diet-induced metabolic syndrome resulted in a reduction in retroperitoneal fat weight and the corresponding fat index. Furthermore, infusions dose-dependently inhibited the elevation in cholesterol level induced by diet (Gancheva et al. 2020). The alleviation of the increase in plasma triglyceride levels by Kochiae fructus ethanol extract (250 mg/g) and its total saponins (100 mg/g), after oral administration of the lipid emulsion, was also reported by Han et al. (2006). Further investigation (in vitro) revealed that the antiobesity effect of Kochiae fructus could be associated with saponins, such as momordin Ic, 2′-O-β-D-glucopyranosyl momordin Ic and 2′-Oβ-D-glucopyranosyl momordin IIc which act as pancreatic lipase inhibitors (Han et al 2006). Furthermore, momordin can also induce an antiobesity 123 effect via PPARδ (peroxisome-proliferator activatedreceptor) mediated mechanism (Sasa et al. 2009). PPARs play a role in the regulation of not only lipid but also glucose homeostasis (Gawrieh et al. 2021). The methanol extract (500 mg/kg) of Kochiae fructus and its butanol fraction (200 mg/kg) exhibited a significant hypoglycemic effect in glucose-loaded rats. Oral administration of extracts decreased serum glucose level by 53–58% compared to the control group (Yoshikawa et al. 1997b). Further research based on bioassay-guided fractionation showed that triterpene saponins were responsible for the hypoglycemic activity of the extracts (Yoshikawa et al. 1997b). Dai and Liu, (2002), observed that oral administration of the total saponin fraction of Kochiae fructus also inhibits the increase in serum glucose in alloxan-induced hyperglycemic mice. Structure activity-relationship (SAR) observations on isolated Kochiae fructus saponins revealed that the free carboxyl group at C-17 (28-COOH) in oleanane-type saponins, as well as the presence of the 3-O-glucuronide moiety in the sugar part, is essential for hypoglycemic activity in vivo. Monodesmosides of oleanolic acid that have a glucoside unit in the sugar chain attached to the C-3 of the aglycone were slightly less effective (Yoshikawa et al. 1997b; Matsuda et al. 1998b, 1999a). Furthermore, it was found that the mechanism of hypoglycemic action after oral administration of saponins is related to inhibition of gastric emptying and therefore the transfer of food to the small intestine (Matsuda et al. 1998a, 1999a, 1999b). It depends on serum glucose level and is mediated, inter alia, by capsaicinsensitive sensory nerves (Matsuda et al. 1999b). In addition, an in vitro study revealed that momordin Ic and oleanolic acid 3-O-glucuronide also inhibited glucose uptake in the rat small intestine (Matsuda et al. 1998b). Antioxidant activity in vitro In recent years, phytochemical research has focused on the analysis of phenolic compounds in Bassia species. Phenolics are substances with a well-documented antioxidant effect, therefore, the antioxidant potential of different extracts of Bassia species were also studied. Most articles provide data on the different extracts prepared from aerial parts (leaves, shoots, fruits) of Bassia plants (Wang et al. 2014a; Phytochem Rev Yusufoglu et al. 2015a; Mohammedi et al. 2019; Gancheva et al. 2020) and fractions obtained by partitioning extracts with solvents of increasing polarity (Chemsa et al. 2016; Khalil et al. 2017; Said et al. 2021). Among the studies, the assays based on the DPPH method predominated. The results of studies on the antioxidant potential of Bassia plants are presented in Table S1 (Supplementary information). Khalil et al. (2017) evaluated the antioxidant potential of different plant parts (leaves, steams, roots) of B. eriophora. A significant DPPH radical scavenging effect (IC50 =19.2–46 µg/mL) of the ethyl acetate and butanol fraction of the 70% methanol extract was observed. A slightly lower antiradical potential was determined for the water fraction (IC50 39.9 to 73.2 μg/mL). In turn, both the chloroform and the hexane fraction of the ethanol extract have the lowest ability to eliminate DPPH radicals (IC50 [ 100 μg/mL). It is interesting that no significant differences in activity were found between extracts obtained from different parts of B. eriophora. In another study on B. eriophora, Said et al. (2021), revealed that 40% methanolic fraction obtained by elution of methanol (80%) extract from aerial parts of the plant on the C-18 short column has good antioxidant potential. The fraction had IC50 values of 177.8 µg/mL in the DPPH test and showed concentration-dependent total antioxidant capacity (TAC) and reducing power (RP). The authors underline the positive correlation between TAC and RP with total flavonoid content. It should be noted that the fraction analysed was defined by LC–MS/MS, which led to tentative detection of 28 flavonoids (acetylated and non-acetylated compounds). In the same study, further theoretical calculations on the O– H bond dissociation enthalpy (BDE) allowed the antioxidant activity to be linked to the location of hydroxyl groups in the flavonoid structure. The value of BDE decreased in structures with two adjacent hydroxyl groups. Therefore, such structures should have significant antioxidant potential (Said et al. 2021). Antiradical activity was also investigated in hexane, ethyl acetate, and methanol extracts from aerial parts of B. indica. The results showed that the most potent effect was observed in methanol and water extracts with IC50 values of 1.30 µg/mL and 1.39 µg/ mL as well as TEAC values of 1.44 mM and 3.60 mM in the DPPH and ABTS assays, respectively. The ABTS-free radical elimination effect of the water extract was higher than observed for the reference BHT (TEAC=1.80 mM) and slightly lower than the anti-DPPH potential (IC50 =0.91 µg/mL in DPPH). Compared to those extracts, the scavenging effect of hexane extract was much lower with IC50 values of 2.18 µg/mL in the DPPH test and 0.17 mM Trolox in the ABTS assay. It can be associated with a lower phenolic content in the hexane extract (5 mg PyE/100 g of extract) compared to the methanol and water extract (146 and 72 mg PyE/100 g of extract, respectively) (Bouaziz et al. 2009). Several studies were also conducted to establish the antiradical activity of extracts of B. muricata. Mohammedi et al. (2019) revealed that methanol extracts from the aerial parts had stronger DPPH radical scavenging activity (EC50 =5–5.03 mg/L) than aqueous (EC50 =5.86–5.90 mg/L), ethanol (EC50 = 6.12–6.40 mg/L), acetone (EC50 =6.70–7.14 mg/L) and hexane extracts (EC50 =7.42–8.03 mg/L). However, water extracts were the most potent in the ferric reducing capacity assay (EC50 =1.39–2.60 mg/L) and β-carotene bleaching (EC50 =5.31-6.0 mg/L). The authors noted that the high antioxidant activity of methanol and water extracts is strictly associated with a high polyphenol content (TPC=122.15–144.82 mg of GAE / g of methanol extract and 98–100.12 mg of GAE / g of water extract). Furthermore, in the same study it was observed that the type of extraction technique used (MAE, Soxhlet, maceration) had no significant effect on the measured values of antioxidant activity (Mohammedi et al. 2019). Recently, the essential oil of the B. muricata shoots, extracted by hydrodistillation by Abd-ElGawad et al. (2020), was tested for its DPPH and ABTS radical scavenging activity. To our knowledge, this was the only study on the antioxidant potential of essential oil of any Bassia species. The study revealed that the essential oil eliminated DPPH radicals with an IC50 value of 20.70 µL/mL and acted as anti-ABTS (IC50 =16.32 µL/mL). The authors suggested that considerable antioxidant potential may be associated with a large amount of oxygenated sesquiterpenes (53.18%), detected by the GC–MS method in the oil, including hexahydrofarnesyl acetone (47.34%) (Abd-ElGawad et al. 2020). Unlike other Bassia species, the analysis of the antioxidant potential of B. scoparia focuses on the 123 Phytochem Rev study of dried fruits, which are a valued medicinal agent in traditional Chinese medicine. Wang et al. (2014a) revealed that the water and ethanol (50%) extract of Kochiae fructus possess the ability to reduce the ferricyanide complex (Fe3+) to the ferrous form (Fe2+) in the FRAP assay. Furthermore, extracts not only demonstrated the potential to eliminate DPPH (IC50 =0.24–0.31 mg/mL) and ATBS radicals (IC50 =0.35–0.43 mg/mL) but also showed superoxide anion (IC50 =5.42–6.64 mg/mL) and hydroxyl radical scavenger properties (IC50 =0.23 mg/mL and IC50 =2.25 mg/mL, ethanol extract and water extract, respectively) (Wang et al. 2014a). Reactive hydroxyl radicals, superoxide anion, and their by-products may cause oxidative damage to biomolecules. Wang et al. (2014a) confirmed that water and ethanol (50%) extracts of Kochiae fructus (0.1–7 mg/mL) were able to dose-dependently inhibit the free radical-induced destruction of biomolecules. Extracts (0.1–7 mg/mL) inhibited lipid peroxidation in the range of 3–80% (compared to the control group). The ethanol extract was found to almost completely suppress lipid peroxidation (at 7 mg/mL). Furthermore, at a much lower concentration, 50% ethanol extract (0.1 mg/ mL) significantly suppressed protein oxidative damage by 73%, while water extract exerted similar activity with a dose of 10 mg/mL. It should be noted that both the ethanol and water extracts (at 0.1–7 mg/ mL) effectively protected DNA from oxidative damage by 19–70% and 4–40%, respectively. The authors indicated that the observed effect, especially observed in the ethanol extract, may be related to the presence of momordin Ic. It is interesting that this triterpene saponin was able to protect free radicalinduced protein oxidation in a dose-dependent manner (Wang et al. 2014a). Momordin Ic (at 1 µM= 0.76 µg/mL) showed significant activity in the model of AAPH- as well as Cu2+/H2O2 -mediated oxidative damage ([80% intact proteins) (Wang et al. 2014a). Published reports indicate that some of the extracts of Bassia plants, especially those rich in polyphenols, show significant antioxidant potential. As a result of the different methods of processing the plant material, it is very difficult to compare these results with each other. It should be noted that to date no analyses have been published that compare the antioxidant activity of several species of Bassia under the same conditions. To the best of our knowledge, only one analysis has been performed to date comparing the 123 activity of two species: B. indica and B. muricata (Bouaziz et al. 2009). The study showed significant differences in the content of phenolic compounds and the antioxidant activity of the extracts depending on the type of extractant used (see Table S1, Supplementary information). The water extracts of aerial parts of B. indica showed the strongest antioxidant activity in the ABTS assay, even higher than the reference substance (BHT) (Bouaziz et al. 2009). Although in vitro models may provide preliminary information on the activity of the extracts, they do not always reflect in vivo conditions. Recently, the antioxidant potential of B. scoparia and B. eriophora was investigated using animal models. The studies were related to the hepatoprotective and nephroprotective potential of the extracts (see next section). Hepato-, nephro-, gastro- and neuroprotective activity Kim et al. (2005a), revealed that Kochiae fructus had a hepatoprotective effect against CCl4-induced liver damage in Wistar rats. Pre-treatment with methanol and the butanol fraction derived from the extract (200 mg/kg body weight; p.o.; once a day for 14 days) resulted in a significant reduction in serum levels of aspartate transaminase (AST), alanine transaminase (ALT), lactic dehydrogenase (LDH) and liver concentration of thiobarbituric acid reactive substances (TBARS) in the CCl4 treated rats. Furthermore, both extracts significantly inhibited the reduction in glutathione, glutathione-S-transferase, and glutathione reductase activity produced by CCl4. The study demonstrated that the hepatoprotective effect of B. scoparia was closely related to the antioxidant activity of the extracts and increased the hepatic antioxidant potential (Kim 2005a). Gancheva et al. (2020) came to similar conclusions by examining the protective potential of the infusion of B. scoparia seeds in Wistar rats with diet-induced metabolic syndrome. The animals received a high-calorie diet along with infusions (ad libitum) prepared with the use of 100 mL of boiling water and 1.5 g, 3 g and 6 g of seeds, respectively. After 12 weeks of treatment, all infusions demonstrated antioxidant activity in vivo in a dose-dependent manner. In the group of animals with metabolic syndrome, which were administered infusion made from 6 g of seeds, the most potent decrease in thiobarbituric acid Phytochem Rev reactive substances (TBARS) in serum, comparable to healthy rats on a regular rat chow diet was observed. Furthermore, the histopathological study demonstrated that pre-treatment with infusion prevents the appearance of liver lesions (Gancheva et al. 2020). Although no bioassay-guided fractionations were performed in the investigation, the authors assumed that oleanolic acid and momordin Ic are the hepatoprotective substances in Kochia fructus as they are known to exert the effect (30 mg/g body weight; p.o.; once a day for 14 days) in the CCl4-induced liver damage model in rats (Kim et al. 2005b). It is interesting that in the scientific literature there are also some reports on the gastroprotective potential of momordin Ic, oleanolic acid 3-O-glucuronide against ethanol- and indomethacin-induced gastric mucosal lesions in rats (Matsuda et al. 1998c, 1999c). However, to our knowledge, no reports on such activity of extracts of Bassia species have been published. Among Bassia species, B. eriophora also revealed a preventive potential in vitro, not for liver but kidney damage. To assess plant activity, the CCl4-induced lipid peroxidation model (Albino Wistar rats) and post-test measurement of the level of malondialdehyde in nephritic tissues, as well as histopathological investigation, were used. The study revealed that pretreatment with 90% alcoholic extract from aerial parts of the plant (250 mg/kg body weight, p.o. and 500 mg/kg body weight p.o./six days) influenced kidney functions as a reduction in serum creatinine and urea levels was observed in animals intoxicated with CCl4. Furthermore, the amount of total proteins, which represents the level of damage, was maintained as in normal rats. The extract of B. eriophora also significantly prevented the elevation of malondialdehyde, a marker of oxidative stress, in the kidney tissue of rats intoxicated with CCl4. Finally, histopathological studies confirmed that pre-treatment of animals with B. eriophora extract significantly and dose-dependently reduced injuries such as vacuolization of the cytoplasm of the epithelial lining of renal tubules induced by CCl4 intoxication. The authors emphasize that kidney protective activity results from the antioxidant and free radical scavenging potential and may be related to polyphenol compounds in the extract of B. eriophora (Yusufoglu et al. 2015a). Recently, Othman et al. (2022) demonstrated the neuroprotective effect of methanol extract of aerial parts or B. indica. To our knowledge, this was the only study on such activity of any Bassia species. The study revealed that the extract showed inhibitory activity against MAO-B (IC50 =8.7; selegiline: IC50 = 0.55 μg/mL), BACE-1 (IC50 =28.9 μg/mL; LY2811376: IC50 =0.8 μg/mL) and neurotoxic Aβ142 aggregation (IC50 =40.6 μg/mL; tecrine: IC50 = 1.93 μg/mL) in vitro. Furthermore, B. indica extract reduced the concentration of phosphorylated tauprotein to 6.62 pg/mL (control=10.8 pg/mL; LY2811376: IC50 =2.09 pg/mL). The search for active compounds revealed that phenylpropanoid amides, such as N-trans-feruloyl-3-methoxytyramine and S-(-)-N-trans-feruloyl octopamine, which occur in aerial parts of B. indica, are associated with the neuroprotective effect of the extract. N-trans-feruloyl-3-methoxytyramine (NTFT) exerted potent inhibitory activity against MAO-B, BACE-1 and Aβ1-42 aggregation with IC50 =0.71 μg/mL, IC50 = 5.39 μg/mL, IC50 =0.3 μg/mL, respectively. NTFT also demonstrated a significant effect in reducing the concentration of phosphorylated tau-protein (to 1.62 pg/mL; control=10.8 pg/mL; LY2811376: IC50 =2.09 pg/mL) (Othman et al. 2022). Furthermore, Khan et al. (2021) noted that NTFT exerts a stimulating effect on neurothrophins and the neurogenesis process through the TrkA/ERK /CREB pathway (Khan et al. 2021). Thangnipon et al. (2012) also indicated that the neuroprotective effect of NTFT against AAβ1–42-induced neuronal death may be associated with its antioxidant activity (Thangnipon et al. 2012). These studies on B. indica and isolated phenylpropanoid amides have provided promising results and thus should be the subject of further in-depth analyzes on neuroprotection and prevention of neurodegenerative diseases. Antibacterial, antifungal, and antiparasitic activity Antibacterial activity studies in representatives of the Bassia genus have been conducted mainly on extracts from aerial parts of plants, including fruits of B. scoparia, prepared with the use of different solvents. Most of the research has been carried out using agar diffusion methods, but studies on the determination of minimal inhibitory concentration (MIC) have also been performed. The list of antimicrobial studies is presented in Table S2 (Supplementary information). 123 Phytochem Rev In a preliminary study on the antimicrobial potential of B. eriophora, ethanol extract from aerial parts of the plant was found to inhibit Escherichia coli, Staphylococcus aureus, Pseudomonas aeruginosa, Klebsiella pneumoniae, and Proteus vulgaris growth (Al-Mussawi et al. 2014). Activity toward S. aureus and E. coli was also observed for fractions obtained by partitioning 70% ethanol extract from different parts of B. eriophora plant (leaves, roots, stem) with hexane, ethyl acetate, and water. All extracts, but especially the water fraction from the roots and stems, exhibited high activity towards S. aureus, with inhibition zones between 27 and 40 mm (control: oxacillin 19 mm; erythromycin 22 mm). In turn, moderate activity towards E. coli was observed for the water extract of the roots and stems, as well as the ethyl acetate fraction of the stem (Khalil et al. 2017). In another study, the antibacterial activity of 95% ethanolic extract of aerial parts of B. eriophora against not only methicillin-sensitive S. aureus (MSSA) but also resistant (MRSA) strains were evaluated. The extract was found to inhibit the growth of both MSSA and MRSA strains, with MIC values of 25–75 μg/mL and 75 μg/mL, respectively. Interestingly, the authors also performed analyzes that evaluated the possibility of using B. eriophora extract to create an antibacterial layer on textiles against S. aureus. Furthermore, they indicate that anti-S. aureus fabrics do not lose their antibacterial properties, even after two washing cycles (prepared according to the test method 61(2A)-1996, the American Association of Textile Chemists and Colorists) (Alsaggaf et al. 2018). The potential of B. scoparia for MRSA was also investigated. The nhexane fraction of the Kochiae fructus ethanol extract exhibited significant activity (MIC=7.8–31.25 μg/ mL; ampicilin 31.25–1000 μg/mL; oxacilin 500– 1000 μg/mL), whereas the butanol, ethyl acetate and water fraction did not inhibit the growth of bacteria. Furthermore, synergy in antibacterial activity was observed between the hexane fraction and antibiotics (oxacillin or ampicillin) (Joung et al. 2012). ElShamy et al. (2012), investigated the antibacterial effect of essential oil from B. scoparia shoots. Although the oil significantly inhibited the growth of E. coli, Bacillus subtilis, and S. aureus (MIC= 12.5 μg/mL, 12.5 μg/mL and 25.0 μg/mL, respectively), the effect was lower compared to cefroperazone (MIC=0.8 μg/mL). The observed 123 effect was probably related to α-thujaplicin, which is the major component of essential oil and has a well-documented antimicrobial activity (Yamaguchi et al. 1999; Morita et al. 2001; El-Shamy et al. 2012). Among Bassia species, also alcoholic extracts of B. muricata moderately suppressed the growth of B. subtilis, S. aureus, P. aeruginosa., E. coli, which was determined using the agar well diffusion method (AlBarri et al. 2021). In another study, the ethanol extract from aerial parts of B. muricata (40 extract/ disc) showed antimicrobial activity against E. coli and P. aeruginosa, which was comparable to amoxicilin (25/disc). Additionally, a moderate effect against Vibrio cholerae, Salmonella enterica, and Enterococcus faecalis was also observed (Chemsa et al. 2016). Some reports investigating the antibacterial potential of aerial parts of B. indica can also be found in the literature (Abdel-Hameed et al. 2008; Bouaziz et al. 2009; Bibi et al. 2021; Ahmed et al. 2022). However, they provide contradictory information that can be associated with, among others, different methods of processing plant material. In addition to antibacterial activities, the antifungal potential of various extracts of Bassia plants was also tested. Unfortunately, some of the studies are not well documented, without a corresponding positive control (antifungal drug) and seem not to be too repetitive. Several reports provide preliminary data that alcoholic extracts from aerial parts of B. muricata, B. prostrata, and B. indica have moderate activity towards some pathogenic fungus. B. muricata, B. indica, and B. scoparia inhibited Candida albicans growth (Al-Barri et al. 2021; Ahmed et al. 2022) as well as B. prostrata was active toward Candida glabrata, Microsporum canis, Aspergillus ﬂavus, and Trichophyton longifusus (Imran et al. 2017) (see Table S2). Most reports indicated inactivity of Bassia extracts toward Aspergillus niger or A. fumigatus (Abdel-Hameed et al. 2008; Al-Barri et al. 2021; Ahmed et al. 2022). Additionally, the essential oil obtained from the B. scoparia shoots, with α-thujaplicin as the main compound (22.91%), did not inhibit the growth of A. niger, while it showed moderate activity towards C. albicans (MIC= 12.5 μg/mL) (cefroperazone was used as a positive control, MIC=5.0 μg/mL) (El-Shamy et al. 2012). In another study, Liu et al. (2012) screened the antifungal activity of different extracts of Kochiae fructus (0.2 g fruits/mL) towards Saccharomyces cerevisiae Phytochem Rev and C. albicans. The aqueous extract showed the best fungistatic activity and suppressed growth by 52% and 29%, respectively (control: miconazole (2 mg/ mL) inhibited growth by 89–90%). In turn, the acetone extract exhibited a lower potential for both S. cerevisiae and C. albicans (24% and 15%), while the ethanol extract as well as the hexane extract showed a poor activity (5–10%) (Liu et al. 2012). On the other hand, ethanol extract of B. scoparia fruits (2 mg/mL) strongly inhibited the growth of the plant necrotrophic fungus–Valsa mali (by 96.16% after 96 h). Furthermore, the extract had a negative influence on the functional stability of the mycelium (Xiaoyan et al. 2019). Furthermore, Houlihan et al. (2019) demonstrated that water exudates from B. scoparia seeds suppress the formation of hyphae in Colletotrichum graminicola with a MIC value of 3.125 mg/L. However, it should be mentioned that the exudates were inactive in relation to the other fourteen fungus tested (Houlihan et al. 2019). Significant inhibitory activity against Macrophomina phaseolina was also observed in the n-butanol fraction of the leaf methanolic extract of Kochia indica. At the concentrations tested (1.56–200 mg/ mL), the extract caused a reduction in the biomass of this fungal plant pathogen in the range of 63–92% (Javed et al. 2018b). Unlike antimicrobial studies, data on the antiparasitic activity of Bassia species are scarce. The methanol extract of the seeds of B. scoparia (100 μg/ mL) showed moderate activity (47% inhibition) against the epimastigote stage of Trypanosoma cruzi in axenic cultures (allopurinol was used as a positive control, 67% inhibition) (Lirussi et al. 2004). In a study on the antitoxoplasmal potential of B. eriophora, a methanol extract of aerial parts of the plant (50 μg/mL) displayed very weak activity (5% of inhibition) against Toxoplasma gondii tachyzoites (Al Nasr 2020). In turn, 95% methanol extract of B. indica (100 and 200 μg/mL) was evaluated against Schistosoma mansoni, but no activity was detected (Abdel-Hameed et al. 2008). The reports published so far clearly indicate a low potential of extracts from Bassia species against analysed parasites, which cause disease in humans. However, there are reports that show significant activity of B. scoparia extracts against animal and plant pathogens. Lu et al. (2012a) examined the antiparasitic potential of various extracts of fruits of B. scoparia against the fish pathogen Dactylogyrus intermedius using an in vivo anthelmintic efficacy model in goldfish (Carassius auratus). The methanolic extract of the fruits of the plant demonstrated potent activity with values of EC50 and EC90 of 31.28 mg/L and 52.52 mg/L (in bath), respectively. Furthermore, the acute toxicity assay revealed that the 48 h LC50 values (71.04 mg/L) are higher than therapeutic concentrations. The other extracts tested (petroleum ether, chloroform, ethyl acetate, acetone) exhibited a lower potential in the anthelmintic model and caused fish mortality in the concentration range 90–130 mg/L (Lu et al. 2012a). In turn, Shi et al. (2006) screened the acaricidal activity of extracts from aerial parts of B. scoparia against the spider mites Tetranychus cinnabarinus, Tetranychus urticae, and Tetranychus viennensis, which are parasites on many different plants and crops. All extracts showed systemic and contact toxicity to mites. Chloroform extract caused a strong mite mortality rate (73.3– 89.12%) and the effect was stronger than observed for methanol extract (39.78–44.78%) and petroleum ether extract (41.92–54.21%) (Shi et al. 2006). These studies have provided promising results and thus should be the subject of further in-depth analyses. Other activities In addition to the biological activities mentioned above, some reports also demonstrated other effects observed for some Bassia species. The aqueous extract of Kochiae fructus (200 g/1500 mL) exerted a decrease in stroke volume and pulse pressure in beating rabbit atria. However, this inotropic activity was not mediated by atrial natriuretic peptide (ANP) secretion (Lee et al. 2016). In another study, Kochiae fructus water extract (20 mg/g, p.o.) demonstrated improvement in heart function parameters in the furazolidone-induced DCM model. Furthermore, the extract showed a regulatory effect on Th1/Th2 cell activity, which suppressed myocardial injury in dilated cardiomyopathy (DCM) (Zou et al. 2021). Yusufoglu et al. (2015a) investigated the antipyretic potential of B. eriophora in the rat model of yeast-induced pyrexia. The 90% ethanol extract (250 mg/kg and 500 mg/kg) from the aerial part of the plant significantly reduced the elevated rectal temperature (39.35 °C). 120 min after oral administration, the temperature decreased to 36.68 °C and 123 Phytochem Rev 35.83 °C, while indomethacin (4 mg/kg) reduced the temperature to 35.45 °C. However, the mechanism of action has not been evaluated. The ethanol extract of aerial parts of B. eriophora (250, 500 and 750 mg/kg; i.p.) was found to cause dose-independent skeletal muscle relaxant activity using the rota-rot test in male Swiss albino mice (Musa et al. 2016) as well. Recent studies in animal models with seizures induced by pentylentetrazol and electroshock indicate that essential oil from fresh leaves of B. scoparia (75, 150 and 300 mg/kg; p.o.) had anticonvulsant activity. Unfortunately, this study is only preliminary because no phytochemical investigation was conducted on the composition of essential oil. Therefore, it is not known which phytoconstituents may be responsible for activity (Imade et al. 2020). In turn, Othman et al. (2021b) screened seventeen constituents isolated from hydroalcoholic extract of aerial parts of B. indica as acetylcholinesterase inhibitors in vitro, including a new acylated flavonol tetraglycoside and a rare triglycoside. Among them, the most potent were 6,7-dihydroxycoumarin and quercetin with IC50 values of 3.6 μg/mL and 18 μg/mL, respectively (galantamine was used as a positive control, IC50 = 12.5 μg/mL). Flavonol glycosides: kaempferol-3-Oβ-D-glucopyranosyl-(1→6)-O-[β-D-galactopyranosyl-(1→3)-2-O-trans-feruloyl-α-L-rhamnopyranosyl-(1→2)]-β-D-glucopyranoside (Fig. 2) and isorhamnetin-3-O-β-D-glucopyranosyl-(1→6)-O-[αL-rhamnopyranosyl-(1→2)]-β-D-glucopyranoside (Fig. 3B) demonstrated weak activity (IC50 =250 μg/ mL) (Othman et al. 2021b). Yusufoglu (2015b) investigated the wound healing potency of gel preparations containing 1% and 2% ethanol extract of the aerial part of B. indica. Both formulations showed significant and dose-dependent wound contracting ability in the excision wound model in Albino Wistar rats. The reduction in wound size caused by topical application (once a day for 20 days) of 2% gel was more potent than that observed for 1% gel or Betadine (84.21%, 78.68% and 60.25%, respectively vs. base gel 56.70%). Safety of Bassia species The scientific literature provides only a few data on the toxicity of Bassia plant extracts. It is worth noting, however, that some species have been clinically used in traditional medicinal systems, which 123 also provides information on their safety in humans. One of such plants is B. scoparia. Toxicological studies in mice indicated that Kochiae fructus water extract demonstrated very low oral acute toxicity with an LD50 value of 7.15 g/kg (Zou et al. 2021). Furthermore, in two investigations, it was confirmed that the ethanol extract from aerial parts of B. eriophora also exhibited low toxicity (Yusufoglu 2015b; Musa et al. 2016) and the LD50 after oral administration in mice was estimated at 33.4 g/kg (Musa et al. 2016). However, it should also be noted that the data available in the literature refer to acute toxicity, while there is no information on the chronic use of various Bassia plant extracts. There are only data on the consumption of hay from certain species as feed for livestock. Some species of the genus Bassia can accumulate oxalates and nitrates, which may be associated with signs of toxicosis in animals, but only after chronic consumption of large amounts of hay (Rankins et al. 1991a, b). However, it should be noted that the level of these components in the Bassia plants depended on many factors (maturity stage, environmental factors) (Finley and Sherrod 1971; López-Aguilar et al. 2013; Steppuhn et al. 1994; Yang et al. 2007; Ma et al. 2011, 2016). Furthermore, the observed presence of nitrates and oxalates did not exclude the medical or even food use of other plants from the Amaranthacae family, such as Beta vulgaris, Amaranthus sp., Spinacia oleracea, which also accumulate antinutritional compounds (Jaworska et al. 2005; Gadermaier et al. 2014; Liu et al. 2015; Corleto et al. 2018; Munekata et al. 2021; Avila-Nava et al. 2021). Thus, this indicates that more attention should be paid to the level of potentially toxic compounds before clinical application of Bassia plants. Among the side effects that can appear in contact with plants of the genus Bassia, pollen allergy manifested as allergic rhinitis should also be mentioned. Most allergens found in pollen from the Amaranthaceae family belong to the profilin group (Gadermaier et al. 2014; Elvira-Rendueles et al. 2017). In a study by Zarinhadideh et al. (2015), the first pollen allergen of Bassia scoparia (named Koc s 2) was also characterized. Phytochem Rev Fig. 6 Dependency network of between phytochemical studies and data on the biological activity of extracts and compounds from Bassia sp. (Abbreviations: “x”–not a promising direction of application) Conclusions The representatives of the genus Bassia have been used by humans in different parts of the world for medical as well as nonmedical purposes. This overview gathers and summarizes available scientific data on phytochemistry, biological activity, and significance of plants belonging to the genus, reorganized by Kaderit and Freitag (2011) and denoted Bassia All. Based on the publications covered in the review, for many years the research on Bassia plants focused solely on one species (B. scoparia), and actually only one of its parts, Kochiae fructus, which is a valuable therapeutic agent in TCM. Only in recent years has there been growing interest in the phytochemistry and biological activity of the other species of the genus. Moreover, some investigations were also conducted on other parts of the B. scoparia plant. However, it should be noted that many species of the genus Bassia still remain poorly investigated or unexplored. Phytochemical studies conducted on Bassia species indicate that these plants synthesize metabolites that belong to very different groups of compounds. The latest studies provided information on flavonoids in B. eriophora and B. indica, as well as lignanamides in the latter species. It is interesting that some of the substances found in Bassia plants are unique and rarely found in the plant kingdom. Such structural biodiversity may have a significant impact on the overall biological activity of specific extracts, but this issue requires in-depth research. So far, only some studies on the biological activity of extracts have attempted to answer the question of which compounds are responsible for the observed effect. Most of them focused on bioassay-guided fractionation of active compounds. This approach allows for the selection of a single active compound, but ignores the possibility of a synergistic effect between the phytoconstituents, which often occurs in plant extracts. For example, for many years, the anti-inflammatory activity of Kochiae fructus extracts was attributed 123 Phytochem Rev only to saponins. However, recent studies indicate that flavonoids and, in the case of aqueous extracts, polysaccharides may also be responsible for the effect. Therefore, determining the composition of the extract is extremely important to understand the resultant activity. Equally important are the quantitative analysis of the compounds in the extract and the selection of active markers responsible for the activity. Ensuring the standardization of extracts enables the maintenance of product quality, repeatability of test results, and potential therapy in the future. In the case of research on the activity of extracts from plants of the genus Bassia, although they undoubtedly indicate a multidirectional pharmacological effect of the extracts, including antiinflammatory, cytotoxic, antioxidant, antimicrobial, glucose-lowering and several potential nontherapeutic effects, only some studies provide information on the chemical characterization and quantification of the main components in the extracts. Interestingly, this also applies to in vivo studies. As a result, there is a lack of data that would allow determining the relationship between the phytochemistry of a given species and the activity of its extracts. This is an issue that requires extensive analysis, especially considering some studies that indicate seasonal variability in the content of individual metabolites. However, it should be noted that many of the Bassia species have been used without standardization in traditional healing systems. In such a case, it would be extremely valuable to analyze the qualitative and quantitative profile of the form of the drug used in traditional medicine or at least to identify the main markers. In this review, we try to present the current state of knowledge and research on species of the genus Bassia. After analyzing the published reports, we schematically summarized the network of relationships between phytochemical studies and data on the biological activity of extracts and compounds from individual Bassia species (see Fig. 6). Based on published research on biological activity, one of the most promising directions of research on extracts from plants of the genus Bassia are analyses related to the therapy of inflammatory-related diseases. This aspect is in part in line with the current use of Kochiae fructus in TMC in the treatment of skin diseases. However, it should be mentioned that inflammatory processes are the basis or cause of the 123 progression of many other diseases that are now increasingly common in the developing society, such as cancer, diabetes, and heart disease. Many investigations revealed that extracts (e.g. of B. scoparia, B. eriophora, B. indica) and some isolated compounds (e.g. momordin Ic, flavonoids, N-trans-feruloyl-3methoxytyramine) show not only anti-inflammatory effects, but also significant cytotoxic, glucose lowering, anti-obesity, neuroprotective, and antioxidant effects in vivo. These results indicate a potential for in-depth research on activity against the metabolic syndrome and noncommunicable diseases. The second direction related to potential further research on the biological activity of extracts from plants of the genus Bassia is the antimicrobial and antiparasitic activity. Unfortunately, some of the studies on this issue provide only preliminary information and others were conducted without comparison of results to the corresponding positive control. However, the studies that have been carried out so far indicated that extracts from plants of the genus did not have a broad and nonspecific range of antimicrobial activity, but some of them showed interesting activity against a specific pathogen, such as methicillin resistant S. aureus (MRSA), or against certain fungi or pests causing significant losses in plant crops. These studies should be significantly extended to determine which compounds present in the extracts are responsible for the effect. In recent years, the possibilities of various nonmedical and non-food uses of Bassia species have been explored. Some of them did not produce the expected good results. An example is the use of B. prostrata for the construction of green walls near highways, reducing air pollution (Hozhabralsadat et al. 2022) and the possibility of using B. scoparia oilseeds as a source for the production of biodiesel (Abideen et al. 2015). However, it should be noted that the latter potential application is possible, but not profitable, because of the possibility of using other plants that are much more abundant in the oil. In the literature, there are some single reports on the interesting potential of different nonmedical and medical applications of Bassia plants, such as the production of cellulose nanofibrils from B. eriophora biomass or the use of B. muricata extracts as an ecological corrosion inhibitor. These studies have provided promising results, but require confirmation, and thus should be subject to further in-depth Phytochem Rev research. Another issue is the possible use of some species of Bassia (B. scoparia, B. indica) in soil phytoremediation. Research in this area is very promising from an industrial and ecological point of view. However, these studies indicate the likelihood of the accumulation of toxic substances in Bassia plants, which excludes the possibility of their use for medical purposes. Therefore, this indicates that more attention should be paid to the environmental conditions under which the plant grows or is cultivated in the case of plants of the genus Bassia used for medical purposes. Species of the genus Bassia possess significant adaptation to unfavorable environmental conditions and the possibility of growth and cultivation even in arid areas. Therefore, one of the interesting potentials of Bassia plants may be their cultivation on wasteland and use as a source of bioactive substances. The research conducted indicates the possibility of obtaining (Z)-5-hexadecenoic acid from seeds of various species of Bassia (B. prostrata, B. scoparia, B. hyssopifolia) as an intermediate to produce insect ovioposition pheromone. It is also interesting to obtain momordin Ic, which is considered one of the main compounds of the Bassia scoparia fruit with multidirectional activity. In summary, species of the genus Bassia are characterized by an interesting, complex phytochemical, and pharmacological profile. In addition, some of them, such as B. scoparia, exhibit several activities that justify its use in the traditional medical system. In turn, other species of Bassia, such as B. muricata or B. indica, which also have an established position in folk medicine, have not been subjected to research that justifies their traditional use. However, studies conducted in recent years have provided new data on the phytochemistry and activity of Bassia extracts that go beyond traditional applications. Additionally, considering the possibility of cultivation of some Bassia species, these plants are interesting targets for further research. However, it should be noted that, despite numerous reports, there are still many gaps in the current knowledge on species of the genus Bassia and there is a need to conduct in-depth research on the currently studied species as well as to start the analysis of plants that have not been investigated so far. Funding This research received no external funding. Declarations Conﬂict of interest conflict of interest. The authors declare that they have no Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/. 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